Stable Carbon Isotopes and Prehistoric Diets in the South-Western Cape Province, South Africa 9780860543763, 9781407345604

Table of contents :
Cover
Copyright
Series
TABLE OF CONTENTS
ACKNOWLEGEMENTS
LIST OF FIGURES
LIST OF TABLES
LIST OF MAPS
CHAPTER 1 INTRODUCTION
CHAPTER 2 THE SOUTH-WESTERN CAPE
CHAPTER 3 ARCHAEOLOGICAL RESEARCH INTO THE LATER STONE AGE OF THE SOUTH-WESTERN CAPE
CHAPTER 4 INDIGENOUS FOODS
CHAPTER 5 ISOTOPIC ECOLOGY
CHAPTER 6 FIELD SAMPLING AND LABORATORY ANALYSIS OF THE PREHISTORIC HUMAN FOOD-WEB
CHAPTER 7 ISOTOPIC ANALYSES OF PREHISTORIC HUMAN SKELETONS
CHAPTER 8 IMPLICATIONS OF SKELETAL ANALYSES FOR ARCHAEOLOGICAL RESEARCH IN THE SOUTH-WESTERN CAPE
TABLES
MAPS
REFERENCES

Citation preview

Stable Carbon Isotopes · and Prehistoric Diets in the South-Western Cape Province, South Africa

Judith Sealy

Cambridge Monographs in African Archaeology 15 BAR International Series 29 3 1986

B.A.R.

5, Centremead, Osney Mead, Oxford OX2 ODQ, England.

GENERAL EDITORS A.R. Hands, B.Sc., M.A., D.Phil. D.R. Walker, M.A.

BAR-S29J,1986 :'Stable Carbon Isotopes and Prehistoric Diets in the South­ Western Cape Province'. @

Judith Sealy,1986

The author’s moral rights under the 1988 UK Copyright, Designs and Patents Act are hereby expressly asserted. All rights reserved. No part of this work may be copied, reproduced, stored, sold, distributed, scanned, saved in any form of digital format or transmitted in any form digitally, without the written permission of the Publisher. ISBN 9780860543763 paperback ISBN 9781407345604 e-book DOI https://doi.org/10.30861/9780860543763 A catalogue record for this book is available from the British Library This book is available at www.barpublishing.com

C AMBRIDGE M ONOGRAPHS I N A FRICAN A RCHAEOLOGY G eneral E ditor J ohn A lexander, M .A., P h.D., F .S.A. D ept. A rchaeology, D owning S treet C ambridge C B2 3 DZ, E ngland M anuscripts s hould b e s ubmitted t o D r. A lexander i n t he f irst i nstance. T itles a lready p ublished : 1 .

' The N iger

D elta :

N wanna N zewunwa. 2 .

A spects

o f

i ts

' Prehistoric I nvestigations i n t he R egion o f J enne , M cIntosh a nd R oderick J . M cIntosh.

3 .

P rehistoric E conomy

B AR-S89.

' Off-Site A rchaeology a nd H uman A daptation i n E astern A frica :

' Settlement

B AR-S114.

P atterns

i n

t he

I nterpretation ' b y M artin H all. 6 .

I ron A ge

o f

Z ululand :

A n E cological

B AR-S119.

' The N eolithic P eriod i n t he S udan , M ohammed-Ali.

7 .

A n

S outhern K enya '

' Later P leistocene C ultural A daptations i n S udanese N ubia ' b y Y ousif M ukhtar e l A min.

5 .

b y

Mali ' b y S usan K eech

A nalysis o f R egional A rtefact D ensity i n t he Amboseli , b y R obert F oley. B AR-S97. 4 .

a nd C ulture '

B AR-S75.

c . 6 000 - 2 500 B .C. ' b y Abbas S .

B AR-S139.

' History a nd E thnoarchaeology i n E astern N igeria :

A S tudy o f I gbo-Igala

r elations w ith s pecial r eference t o t he A nambra V alley ' b y P hilip A digwe O guagha a nd A lex I kechukwu O kpoko. 8 .

B AR-S195.

' Meroitic S ettlement i n t he C entral S udan : N ile V alley a nd t he W estern B utana '

A n A nalysis o f

S ites i n t he

b y K hidir A bdelkarim A hmed .

B AR

S 197. 9 .

' Economy

a nd

T echnology

C harles C able. 1 0.

' Frontiers :

i n

t he L ate

S tone

A ge

S outhern A frican A rchaeology T oday '

D .M. A very, M .L . Wilson a nd A .J.B. H umphreys. 1.

o f

S outhern N atal '

b y

B AR-S201. e d . M . H all ,

G . A very ,

B AR-S207.

' Archaeology a nd H istory i n S outhern N igeria :

T he a ncient l inear

e arthworks o f B enin a nd I shan ' b y P .J . D arling . B AR-S215 . 1 2.

' The L atest

S tone A ge

o f

S outhernmost A frica ' b y J anette D eacon .

B AR

S 213. 1 3.

' Fisher-Hunters

a nd N eolithic P astoralists

J ohn W ebster B arthelme.

B AR-S254.

i n E ast T urkana ,

K enya '

b y

1 4.

h e A rchaeology o f C entral D arfur ( Sudan ) b y I brahim M usa M ohammed. B ARS285.

i n t he

i st M illennium A .D . '

T ABLE O F C ONTENTS

A cknowledgements L ist o f f igures L ist of t ables L ist o f m aps

i i i ii v i

C HAPTER 1 I NTRODUCTION

1

C HAPTER 2 T HE S OUTH-WESTERN C APE

4

C HAPTER 3 A RCHAEOLOGICAL R ESEARCH I NTO T HE L ATER S TONE A GE O F T HE S OUTH-WESTERN C APE

1 0

C HAPTER 4 I NDIGENOUS F OODS

2 0

C HAPTER 5 I SOTOPIC E COLOGY

3 4

C HAPTER 6 F IELD S AMPLING A ND L ABORATORY A NALYSIS O F T HE P REHISTORIC H UMAN F OOD-WEB

4 8

C HAPTER 7 I SOTOPIC A NALYSES O F PREHISTORIC H UMAN S KELETONS

7 6

C HAPTER 8 I MPLICATIONS OF S KELETAL A NALYSES F OR A RCHAEOLOGICAL R ESEARCH I N T HE S OUTH-WESTERN C APE

8 3

T ABLES

9 1

M APS

1 24

R EFERENCES

1 32

A CKNOWLEGEMENTS

A l arge n umber o f p eople h ave c ontributed t o t his p roject.

I t w ould

h ave b een i mpossible t o c omplete w ithout t he h elp o f a ll t he f armers w ho k indly a llowed m e a ccess t o t heir f arms ,

p eople w ho d onated s amples ,

b otanists

t o

a nd

z oologists

w ho

h elped

t he r esults of t he a nalyses. M argaret

A very ,

A nne B ean ,

V illiers , W ieland G evers , J enny Jarvis,

m e

i dentify t hem a nd

a nd

i nterpret

Ia m e specially g rateful t o G raham a nd

R ichard B oycott, R obbie G riffiths,

R ichard B rooke , R obert H edges ,

A therton d e

N igel J arman ,

J im J obling, F iona K otze, J ohn L anham, J ulie L ee T horp,

C hristien Malan , W endy Metelerkamp , E ugene Moll, K laas M outon, A lan M orris,

C edric

S taff a nd C ape

P oggenpoel,

s tudents o f

T own

h elped

u npublished

w ith

r esearch

J ohn P arkington ,

t he

w ho

J ohn

V ogel,

A rchaeology

t he

f ield

r esults .

W enner a nd R oyden

D epartment

w ork

T he

a lso h elped

D avid

a nd

p roject

f inance

o f

allowed

w as

t he

U niversity

a ccess

i nitially

o f

t o t heir

s uggested

b y

i t t hrough h is H uman S ciences

R esearch C ouncil - f unded S patial A rchaeology R esearch U nit . f inancial

Y ates .

A dditional

s upport w as p rovided b y t he O ppenheimer I nstitute f or A frican

S tudies a t t he U niversity o f C ape T own a nd b y t he C ouncil a nd

I ndustrial

a nd

gave

R esearch .

u nfailing

N ick

v an

e ncouragement

d er a nd

M erwe

f or S cientific

s upervised t his

s upport.

A g reat

p roject, d eal

of

i nvaluable a ssistance f rom J ohn L anham m ade p ossible t he p reparation o f t his

v olume .

L IST O F F IGURES

F igure 1 G as s eparation l ine i n t he A rchaeometry L aboratory,

U niversity o f C ape T own

5 2

F igure 2 S easonal c hanges

i n t he 61 3 C v alues

o f W atsonia p yramidata

6 5

F igure 3 613 C v alues o f s eaweeds f rom t he s outh-western C ape

6 9

F igure 4 613 C v alues o f s eaweeds f rom p ublications

1 1

6 9

L IST O F T ABLES

T able I M ammal r emains f rom a rchaeological s ites i n t he s outh-western C ape

9 2

T able I I B ird r emains f rom a rchaeological s ites i n t he s outh-western C ape

9 5

T able I II F ish r emains f rom a rchaeological s ites i n t he s outh-western C ape

9 6

T able I V S hellfish r emains f rom a rchaeological s ites i n t he s outh-western C ape

9 7

T able V E dible p lant r emains f rom a rchaeological s ites i n t he s outh-western C ape

9 8

T able V I E dible i ndigenous p lants o f t he s outh-western C ape

9 9

T able V II A natomical d ifferences b etween p lants w ith d ifferent s ub-types o f C4 p hotosynthesis

1 07

T able V III 6 13

C v alues o f c ollagen f rom a nimals i n t he s outh-western C ape

1 08

T able I X 613

C

s pacing b etween d ifferent t issues o f t he s ame a nimal

1 11

T able X ( S 13 C

v alues o f m eat f rom a nimals w ith

b ased d iets

C3 -

i n t he s outh-western C ape

1 12

T able X I C hange i n 613

C

v alues o f a nimal t issues w ith t rophic l evel

1 12

T able X II 613

C

v alues o f t he e dible p arts o f p lants

1 13

T able X III 1 13 5 C

v alues o f m arine a nimals

1 14

T able X IV 413

C

v alues o f m arine a lgae

1 15

T able X V 413

C

v alues o f m arine a lgae f rom d eep a nd

s hallow w ater a t H out B ay ,

C ape P eninsula

1 18

T able X VI P hysiological m easurements o n G reat B arrier R eef s eaweeds

1 19

T able X VII ( S 13 C

v alues o f m arine o rganisms a t d ifferent t rophic l evels

1 20

T able X VIII ( S 13 C

v alues o f m iscellaneous

i ndigenous

f oods f rom t he s outh-western C ape

1 21

T able X IX 613

C

v alues o f a rchaeological a nimals f rom E lands B ay C ave

1 21

T able X X i 5 13 C

v alues o f d ated h uman s keletons f rom

E lands B ay a nd t he O lifants R iver V alley

i v

1 22

T able X XI 1 3

Cv alues o f o ther d ated h uman

s keletons f rom t he s outh-western C ape c oast

1 23

T able X XII M onths i n w hich j uvenile d assies f rom D e H angen w ere k illed

1 23

L IST O F M APS

M ap I Location

of

research

area

1 25

M ap I I G eography o f t he r esearch a rea

1 26

M ap I II V egetation t ypes o f t he s outh-western C ape

1 27

M ap I V L ocation o f a rchaeological s ites m entioned i n t he t ext

1 28

M ap V L ocation o f p lant a nd a nimal s ampling s tations

1 29

M ap V I P roportions o f d ifferent

C3

parts

a nd C4 g rasses i n of

the

s outh-western

C ape

1 30

M ap V II L ocations w here h uman s keletons a nalysed w ere f ound

v i

1 31

C HAPTEH 1

I NTRODUCTION

T his v olume c onsists o f a s table c arbon i sotope a ssessment o f H olocene h uman d iets

i n t he s outh-western C ape ,

S outh A frica ,

a nd t ests

v arious

m odels o f p rehistoric s ubsistence d erived f rom a rchaeological e vidence . S uch a ssessments h ave b een a ttempted i n a n umber o f a reas y ears ,

i n r ecent

i ncluding w oodland N orth A merica ( van d er M erwe a nd V ogel 1 978 ),

D enmark ( Tauber 1 981), V enezuela ( van d er M erwe , R oosevelt a nd V ogel 1 981) a nd t he w est c oast o f C anada ( Chisholm ,

N elson a nd S chwarcz 1 982).

S ome p reliminary w ork h as a lso b een d one

S outh A frica

1 976 ,

1 979;

R awlinson

1 982).

A ll

i n

o f t hese

s tudies

( Silberbauer

h ave ,

h owever,

c onsisted m ainly o f i sotopic a nalyses o f h uman s keletons ,

c onstructing

t heoretical

p rinciples .

d ietary

c arbon

i sotope

r atios

f rom

f irst

S ilberbauer a nd R awlinson r ecognised t hat t his l imited t he a mount o f i nformation t hey c ould g lean f rom s keletal a nalyses , a nd r ecommended t hat f uture r esearch c oncentrate o n o btaining a m ore d etailed p icture o f t he

. 513 C v alues

o f

t he

a ctual

p lants

a nd

a nimals

e aten .

T his,

t herefore, i s t he f irst e xplicitly e cological s tudy o f ap rehistoric h uman f oodweb .

T he f irst

s tep

i n t he

i sotopic

c haracterisation o f t his f oodweb i s t he

i dentification o f t he m ost i mportant a nimal a nd v egetable c omponents o f t he H olocene h uman d iet i n t he s outh-western C ape .

E thnographic s ources

a nd t he f ood-waste r ecovered f rom a rchaeological e xcavations p rovide m uch o f

t his

i nformation .

T he

l ist o f o rganisms o btained

i n f act

i ncludes a f airly r epresentative c ross-section ( in i sotopic t erms) o f t he f auna a nd f lora o f t he a rea . A s election o f t he m ost i mportant i tems w as c ollected f rom e ach o f t he c learly-differentiated e cological z ones o f t he r esearch a rea ,

a nd t he

13

C / 12 C r atios m easured .

T his i nformation

e nables o ne t o e stimate t he i sotopic " signature" o f t he a verage d iet t hat w ould h ave b een c onsumed b y ap erson l iving i n a ny o ne o f t hese e cological

z ones .

b ody t issues, c arbon

i sotope

i t

S ince

t his

" signature"

i s

i ncorporated

i nto p eoples'

i s p reserved i n a rchaeological s keletons .

a nalysis

of

these

1

s keletons,

c ombined

S table

w ith

t he

i nformation o n d iet d escribed a bove ,

t ells u s t he p roportions o f f oods

f rom e ach i sotopically d istinct a rea t hat t he i ndividual c onsumed d uring h is

l ifetime .

T hese

s pent i n e ach z one, about

s easonal

p roportions

o bviously r eflect t he a mount o f t ime

t hus p roviding a n i ndependent t est o f h ypotheses

m ovements

of

p eople

g enerated

f rom

e xcavated

a rchaeological m aterial.

A lmost a ll r ecent a rchaeological r esearch i n t he s outh-western C ape h as b een c onducted w ithin

t he

f ramework o f a s easonal

p roposed b y J ohn P arkington 1 983,

1 984 ) .

( 1972 ,

1 976a ,

1 976b ,

m obility h ypothesis

1 977 ,

1 979 ,

1 980 ,

1 981,

P arkington h as p ostulated s easonal m ovement o f L ate S tone

A ge p eople b etween t he c oast a nd t he i nterior, i n ac ycle d esigned t o m ake m aximum u se o f t emporary p eaks s ources ,

w hile a voiding t roughs .

i n s easonally f luctuating f ood

T here i s s ome a rchaeological e vidence

f or t his , b ut i t i s d ifficult t o p rove o r d isprove c onclusively . t he

g reat

s trengths

o f

t his

h ypothesis

i s

t hat

i t

h as

O ne o f

t estable

c onsequences a nd s table c arbon i sotope a nalysis i s i deally s uited t o s uch t esting .

I f p eople w ere e ngaging i n a s easonal r ound i ncorporating

b oth t he c oast a nd t he r iver v alleys o f t he i nland m ountains, a rchaeological

s keletons

t hen

f rom t hese t wo a reas s hould b e t he r emains o f

t he s ame p opulation , a nd h ave s imilar i sotopic r atios .

Q uite a part f rom t he t esting o f t his e xtremely i nteresting h ypothesis , a b etter u nderstanding o f

t he

di 3 C v alues o f t he

h uman

f oodweb

s keletal r emains w ill p rovide a g reat d eal o f o ther i nformation . m easurements

a re

I ndividuals. r econstructions ,

d irect

T he

r eflections

b iases

i nherent

o f i n

t he m ore

d iets

o f

a nd S uch

p rehistoric

c onventional

d ietary

s uch a s d ifferential p reservation o f d ifferent k inds o f

f ood-waste , d o n ot e nter t he p icture .

M oreover , s mall-scale d ietary

d ifferences w ithin o ne p opulation ( e .g .

d ifferences b etween t he d iets o f

m en

a nd w omen)

i mpossible ,

c an b e

s tudied .

T hese a re o ften d ifficult,

i f n ot

t o d etect a t t he l evel o f s ite-based r econstructions .

T he d13 C v alues o f m odern p lants a nd a nimals t hat w ill b e o btained w ill a lso s erve a s a b aseline f or p alaeoenvironmental r econstructions . m easurements a re c limate-dependent ,

di 3 C

s o t hat o nce w e h ave a g ood i dea o f

w hat t he m odern c arbon i sotope " map" l ooks l ike,

w e w ill b e a ble t o

d etect d eviations f rom t his n orm c orresponding t o p alaeoclimatic s hifts .

2

I t m ay b e,

f or e xample,

t hat

t here w as a w armer p eriod

i n t he m id-

H olocene c aused b y a s light p oleward s hift o f t he z onal c irculation b elts

( van

Z inderen

necessarily

B akker

very

1 978).

general,

p alaeoenvironmental

P redictions

and

knowledge

the

d oes

present

n ot

u sually

r econstruction o f t he c limates o f s mall a reas . r egional

t opographic

f eatures ,

c hanges

o f

i n w ind

t his

k ind

a re

s tate

of

our

a llow d etailed

T hese a re

i nfluenced b y

v elocities o r t he t ype

a nd a mount o f p recipitation , a nd a g reat d eal m ore r esearch i s n eeded b efore p ast c limates i n m ost s mall a reas c an b e a ccurately d escribed ( Parkington 1 984 ) . t o

t he

F uture s table c arbon i sotope r esearch c an c ontribute

s olving o f

s uch p roblems .

I sotopic r atios o f t he b ones o f

t erritorial a nimals f rom w ell-dated a rchaeological

s equences w ill

r eflect p ast c limatic c onditions n ear t he s ite, w hile t hose o f m ore w ide-ranging

a nimals

w ill

p rovide

a n

i ntegrated

m easure

o f

t he

e nvironment o n a l arger s cale .

T his

s tudy w ill

t herefore

f ulfil

s ome

s hort-term r esearch o bjectives ,

a nd c ontribute t o t he s olving o f a n umber o f l onger-term o nes . s quarely

i nto

a m ajor p rogramme

o f

i nvestigation

I t f its

i nto t he L ater S tone

A ge o f t he s outh-western C ape .

T he r est o f t his v olume i s d ivided i nto s even c hapters . d iscussion o f t he g eography ,

g eology ,

w estern

a s

C ape,

a rchaeology a nd

a nd

i s

m eant

C hapter 2 i s a

f auna a nd f lora o f t he s outh-

a b ackground t o

i sotope e cology w hich f ollow .

t he

d iscussions

o f

C hapter 3 c onsists o f a

s hort a ccount o f w hat w e k now a bout t he L ater S tone A ge o f t he a rea . C hapter 4 i s ab road-based a ccount o f i ts e dible p lants a nd a nimals w ith c omments

d rawn

f rom

b oth

e thnographic

a nd a rchaeological

s ources.

C hapter 5 c ontains a d iscussion o f s table i sotope t racer t echniques w ith s pecific

r eference

t o t he

s outh-western

C ape .

T he

v arious

c lasses

o f

p lant a nd a nimal m aterial t o b e a nalysed a re m entioned ,

a nd t he k inds o f

i sotopic v ariation e xpected

C hapter 6 i s a n

i n e ach c ase e xamined .

a ccount o f t he f ieldwork a nd l aboratory p rocedures , o f t he a nalyses. d efined ,

a nd o f t he

r esults

T he i sotopic s ignature o f e ach e cological z one i s

a nd i n C hapter 7 t his

i nformation i s u sed t o i nterpret 61 3 C

r eadings o n a rchaeological h uman s keletons. v olume w ith a d iscussion o f t he

C hapter 8 c oncludes t he

i mplications o f t he r esults f or

a rchaeological r esearch i n t he s outh-western C ape .

3

f uture

C HAPTER 2

T HE S OUTH-WESTERN C APE

T he r esearch a rea

i s t he s outh-western p art o f t he C ape P rovince o f

S outh A frica f rom t he C ape P eninsula i n t he s outh t o L ambert 's B ay i n t he n orth , a nd f rom t he A tlantic c oastline i n t he w est t o t he e dge o f t he

K aroo

i n

d escription

t he

e ast

o f t he a rea ,

( see

Map

I ).

T his

i ts g eography ,

c hapter

c limate,

c onsists

o f

a

f auna a nd f lora,

c oncentrating u pon t hose a spects o f t he e cology w hich w ould h ave b een o f i mportance t o

i ts p rehistoric

i nhabitants .

G EOGRAPHY A ND C LIMATE

S everal d istinct p hysiographic z ones c an b e d istinguished , t hese t end t o r un

f rom

S andveld , t he K aroo .

n orth

t o

s outh

a nd

are

( in

o rder ,

f rom

t he

c oast):

t he

t he C oastal F oreland , t he m ountains o f t he C ape F old B elt a nd T he d iffering g eology a nd r ainfall o f t hese v arious r egions

m ean t hat t hey s upport n oticeably d ifferent v egetations,

a nd h ence

p rovided d ifferent s ets o f r esources t o t he p rehistoric p opulation .

A long t he c oast ,

t he r ainfall v aries f rom a bout 6 00 m m/year a t C ape T own

t o l ess t han 2 00 m m/year a t E lands B ay .

T he F old M ountain B elt t ends t o

b e w etter ,

w ith t he h ighest a reas r eceiving m ore t han 3 500 m m/year ,

t he K aroo,

w hich l ies i n t he r ain-shadow o f t hese m ountains,

d rier ( Weather B ureau 1 965).

b ut

i s m uch

T he w hole o f t he r esearch a rea i s i n t he

w inter r ainfall r egion o f t he s outh-western C ape , s o t hat m ost o f i ts r ain f alls b etween M ay a nd O ctober , a lthough t here c an b e o ccasional s hort s howers i n o ther m onths . i n w inter , b ut i t i s i nfrequent .

S now m ay b e f ound o n t he m ountain t ops F rosts a re m ore c ommon , o ccurring o n

a n average o f a bot 4 0 d ays e ach w inter i n t he m ountains .

N earer t he

c oast t hey a re v ery r are, a nd s easonal v ariations i n t emperature a re l ess p ronounced

( Weather B ureau

1 954 ).

4

G EOLOGY A ND V EGETATION

T he g eology o f t he a rea

i s

v aried .

T he F old M ountain B elt

i tself

c onsists m ainly o f h ard q uartzitic r ocks b elonging t o t he T able M ountain G roup .

T hese w eather s lowly , a nd t he s oils a s t hey a re p roduced t end t o

e rode a way b ecause o f t he s teepness o f t he t errain .

A s ar esult, t he

m ountain s oils a re s hallow , s tony , a cid a nd r ather p oor i n n utrients ( Lambrechts 1 979). A cocks '

T he v egetation t hey s upport

v egetation t ype N o .

f ynbos t ypes.

6 9

( Acocks

1 953),

i s m ountain f ynbos,

o ne o f t he t wo m ajor

F ynbos i s o ne o f t he s ix f loral k ingdoms o f t he w orld .

I t i s c haracterised b y a v ery h igh s pecies d iversity ( more t han 6 000 p lant s pecies h ave b een r ecorded f rom t he f ynbos b iome ), s ingle-species d ominance .

M any p lants h ave v ery r estricted r anges,

s ometimes o nly a f ew h ectares . l ow ,

a nd a l ack o f

F ynbos a lways c ontains e ricoid s hrubs :

e vergreen s hrubs w ith h ard ,

r olled l eaves,

i ncluding

a re a lways p resent :

r eed-like ,

t ufted e vergreen p lants w ith n early l eafless t ubular s tems ;

t he f amily

h eaths ,

E rica s pp .

R estionaceae c omponent,

i s

R estioid h erbs ,

p articularly

p roteoid s hrubs,

n arrow , t oo ,

p rominent

w idespread .

i s p resent i n s ome h abitats .

m edium t o t all s hrubs w ith f airly l arge , ( Taylor 1 972 ).

a nd

T hese a re

l eathery a nd o ften h airy l eaves

F ynbos i s a dapted t o p eriodic f ires ,

w ith u nderground b ulbs,

A t hird

c orms a nd r hizomes

a fter w hich p lants

( chiefly m embers o f t he

I ridaceae a nd L iliaceag ) p roduce s pectacular d isplays o f f lowers ( Kruger 1 979a).

T he p roteoid e lement d oes o ccur

i n m ountain f ynbos,

w here

m embers o f t he P roteaceae a nd o ther f amilies a re s ome o f t he l argest p lants t o b e s een .

T rees a re r are i n t his v egetation t ype,

a lthough

t hey d o o ccur i n v alleys a nd r avines w here t he s oil i s o ften d eeper . T rue m ountain f ynbos i s p oor i n g rasses f or g razing a nimals ,

a nd i n t he

g eophytes w hose c orms w ere s ought a fter b y p rehistoric p eople ( Taylor 1 972 ),

a nd s o w as p robably n ot o ne o f t heir p referred h abitats .

C oastal

f ynbos

( Acocks '

v egetation

m arine s ands a long t he c oast. h ere t han

i t

i s

t ype

N o .

4 7 )

o ccurs o n f ine-grained

T he p roteoid e lement i s l ess i mportant

i n m ountain f ynbos ,

b ut t here a re m ore g rasses ,

m any o f

t hem p alatable a nd n utritious t o g razing a nimals ( Taylor 1 978). o ther i s

T he

v egetation t ype w hich o ccurs o n t hese s ands o n t he c oastal p lain

s trandveld

( Acocks '

v egetation t ype N o .34 ) .

T his

i s

a b road-leaved

s clerophyllous w oody s hrubland c haracteristic o f c oastal s and-dunes , b ut

5

i t m ay b e m ixed w ith c oastal f ynbos . o r s emi-succulent p lants ,

I t i ncludes a n umber o f s ucculent

a nd a nnuals a nd g rasses a re p rominent

i n t he

s pring ( Acocks 1 953) .

A l arge p art o f t he c oastal f oreland h as , g eology a nd v egetation .

b y c ontrast ,

T he d istricts o f M almesbury ,

q uite d ifferent M orreesburg a nd

P iketberg t hat a re t oday i mportant w heat-farming a reas, d erived f rom s hales,

h ave s oils

r ather t han q uartzites a nd s andstones .

T hese

i nclude a g reater p roportion o f f ine s and a nd s ilt t han o ther w estern C ape s oils,

a nd a re

l ess a cid a nd r icher

p otassium ( Lambrechts 1 979).

i n n utrients,

e specially

T oday , t he r are u ncultivated p atches o f

t hese f ertile s oils a re c overed w ith c oastal r enosterveld ( Acocks ' v eld t ype N o .

4 6 ) .

E lytropappus r hinerocerotis

( renosterbos )

i s t he d ominant

s pecies, a nd a lthough t here a re o ther n arrow-leaved s clerophyllous s hrubs ,

e ricoids a nd p roteoids a re r are .

l ower t han i n f ynbos c ommunities , m ore

c ommon

( Acocks

T he

s pecies d iversity

i s f ar

a nd a nnual h erbs a nd g rasses a re m uch

1 953) .

O n t he e astern s ide o f t he F old B elt , b oth g eology a nd v egetation c hange a bruptly a s t he q uartzites a nd s andstones o f w hich t he m ountains a re c omposed g ive w ay t o t he E cca a nd D wyka s hales o f t he K aroo s ystem . v egetation t ype i s A cocks ' s ucculent k aroo ( type N o . 3 1). k ind

o f

k aroo

f ound

i n

h ot,

a rid a reas

r ainfall o f b etween 5 0 a nd 2 00 m m/year .

T his i s t he

w ith w inter o r y ear-round

I t c ontains n umerous s ucculent

s pecies a s w ell a s a g reat m any g eophytes a nd a nnuals ( Acocks 1 953). s pring t here i s p lenty o f g rass ,

T he

I n

a nd e dible g eophytes c an b e c ollected ,

b ut f or t he r est o f t he y ear a nimals h ave t o b rowse o n t he s mall b ushes o r s ucculents .

T HE L ATE P REHISTORIC F LORA A ND F AUNA

T hose b oundaries b etween a nd c haracteristics o f v egetation t ypes t hat a re d etermined b y g eological

s ubstrate a re u nlikely t o h ave c hanged i n

t he l ast f ew t ens o f t housands o f y ears. w estern C ape h as , a ctivity ,

h owever ,

b een c onsiderably d isturbed b y r ecent h uman

p articularly a griculture

k eeping s ince c a . 2 000 B .P .

T he e cology o f t he s outh-

T he m odern

s ince c a .

1 750 A .D .,

v egetation

a nd s tock-

i s w idely c onsidered

t o d iffer i n an umber o f r espects f rom t hat o f t wo t housand y ears a go .

6

I ncreased f ire f requency a nd g razing p ressure h ave p robably r educed t he a mount o f g rass,

a nd s crubby s hrublands

e xpansion ( Levyns 1 927 ; H endey 1 983) . t rue o f t he r enosterveld , r enosterbos)

u ndergone

a c orresponding

T his i s l ikely t o b e p articularly

w here t he p lants m ost

c ommon t oday

( e .g .

a re k nown t o b e w eedy ,

p ioneer s pecies t hat t hrive i n

d isturbed o r r ecently-burned g round .

T he " climax" v egetation o f t he

s hale-derived s oils o f t he c oastal f oreland w as p robably r ather g rassier t han m odern r enosterveld ( Hendey 1 983 ;

I n o ther a reas ,

w ithin

t he

t he c arrying c apacity o f p arts o f t he K aroo t hat

r esearch

a rea

h as

d ecreased

o vergrazing a nd p oor l and m anagement . t his c entury ,

c omm . ).

r ecent c hanges i n t he v egetation a re e ven m ore d ramatic .

W ithin l iving m emory , f all

M oll p ers .

F arms t hat ,

p rovided g ood s pring g razing ,

v egetation ( Nieuwoudt,

p ers .

t remendously

d ue

t o

a t t he b eginning o f

a re t oday a lmost d enuded o f

c omm .).

F actors s uch a s t hese h ave h ad a p rofound e ffect o n t he i ndigenous f auna o f t he r egion .

A lthough a n umber o f s mall a nimals a re s till p lentiful

a ll o ver t he r esearch a rea ,

h erds o f l arge u ngulates a re e xtinct e xcept

i n a f ew s mall g ame r eserves .

I t

i s ,

t he l ikely l ate p rehistoric f aunal

h owever ,

p ossible t o r econstruct

c ommunities f rom a k nowledge o f t he

v egetation, t he r ecords o f e arly t ravellers a nd t he r emains f ound i n a rchaeological e xcavations .

C ommon s mall a nimals i nclude d assies o r r ock h yraxes ( Procavia c apensis) a nd

s mall

b uck

( Raphicerus h are

( Lepus

w idespread ,

s uch

m elanot 4 )

a s

( Raphicerus

a nd g rey d uiker

c apensis) a s

s teenbok

a nd

d une

c ampestris),

( Sylvicapra

m ole

i s t he a ngulate t ortoise

r at

q rimmia ) .

( Bathyergus

g rysbok T he

C ape

s uillus)

( Chersina a nqulata ) .

a re

A ll t hese

a nimals a re f ound i n a ll t he v egetation t ypes d escribed a bove .

T hey a re

c ommon

a nd a re ,

i n t he f ood w aste

e xcavated f rom a rchaeological

s ites ,

i ndeed ,

s till i mportant i n t he d iets o f r ural p eople t oday .

L arger

g razing

a nimals

a re

l ikely

t o

h ave

b een

g rassier a reas o f t he c oastal p lain o r t he K aroo . w ith

i ts

a nimals :

r ich p astures

T he c oastal

t o t he

f oreland

i s k nown t o h ave s upported n umbers o f l arge

e arly t ravellers r ecorded e lephant

r hinoceros

r estricted

( Diceros b icornis),

( Loxodonta a fricana ),

h ippopotamus

7

b lack

( Hippopotamus a mphibius),

z ebra ( Equus b urchelli),

q uagga ( Equus q uagga ),

a nd h artebeest ( Alcelaphus b uselaphus)

e land ( Taurotraqus o ryx )

( Skead 1 980 ) .

T hese s pecies w ere

p robably m ore c ommon i n t he r enosterveld ( or i ts p rehistoric e quivalent) t han i n t he l ess

f ertile c oastal

f ynbos a nd s trandveld ,

a lthough t his

i s ,

a s y et ,

u nproven .

T he

h ighest

p arts o f t he F old B elt m ountains p robably n ever s upported

m any a nimals ,

f or t he r easons a lready m entioned ,

b ut t he f oothills a nd

r iver v alleys w ere r ich h unting g rounds f or i ndigenous p eople a nd e arly s ettlers a like .

S eventeenth- a nd e ighteenth-century r ecords o f a nimals

e ncountered i n t he O lifants R iver v alley m ention h erds o f e lephant w ith s everal h undred a nimals

( Thom 1 958 ).

T he K aroo i s l ikely t o h ave s upported h erds o f l arger a nimals , p ossibly m igratory h erds w hich m oved i nto t he w inter r ainfall a rea t ake a dvantage o f t he n ew g rass .

T here a re h istorical r ecords o f t his

b ehaviour p attern i n s pringbok ( Antidorcas m arsupialis ) E arly f armers

i n s pring t o

i n t he a rea f ollowed a s imilar s chedule ,

( Thunberg

1 795) .

m oving b etween

t he m ountains a nd t he K aroo t o m ake t he m ost o f t emporary p astures ( Borcherds 1 861).

T he v arious v egetation t ypes t hus p robably s upported s ubstantially d ifferent f aunal c ommunities .

V ariations i n b oth t he p lant a nd a nimal

f ood p otential o f t he d ifferent p hysiographic z ones o f t he s outh-western C ape m ean t hat p rehistoric p eople w ere e xploiting a s trongly-patterned e nvironment o ffering a f luctuating f ood s upply .

T HE A TLANTIC C OASTLINE

T he l ast m ajor e cological z one i s t he A tlantic s horeline , ac oast w ellk nown f or i ts h igh p lant a nd a nimal b iomass . s outh-easterly w inds ,

H ere t he c ombination o f

t he B enguela c urrent a nd t he e arth 's s pin d eflect

s urface w aters a way f rom t he c oast s o t hat d eep ,

c old w ater h as t o r ise

u p t o r eplace t hem ( Branch a nd B ranch 1 981) .

T his c old w ater i s r ich i n n utrients w hich s upport l arge p opulations o f p lankton .

P lankton c ontributes, t ogether w ith a lgae, t o t he l owest

t rophic l evel i n af ood-web w hich i ncludes f ilter-feeding s hellfish

8

( such a s m ussels) a nd f ish l ike a nchovies a nd s ardines, l arger c arnivorous f ish . s eals a nd p eople .

M ussels, l impets, f ish , f ish-eating b irds,

c rayfish p rovided a r ich

s torehouse o f f ood f or p rehistoric

S hellfish a re a p redictable r esource t hat p rehistoric

d wellers m ay h ave u sed a s a s taple f ood , t he l arger a nimals m entioned 1 972 ,

t he p rey o f

1 976a ,

T he r elative

1 976b ,

1 980 ;

( H .J .

c oast-

s upplemented o ccasionally b y

D eacon 1 972 ,

1 976 ,

1 983 ,

P arkington

R obertshaw 1 979a ) .

i mportance o f a ll t he d ifferent

w ill b e d iscussed i n d etail i n C hapter 4 .

f ood r esources m entioned

T his i nformation w ill t hen b e

i ntegrated i nto a d iscussion o f w hat i s k nown a bout p rehistoric d iets i n t he s outh-western C ape a s r evealed b y a rchaeological e xcavations .

9

C HAPTER 3

A RCHAEOLOGICAL RESEARCH I NTO T HE L ATER S TONE A GE O F T HE S OUTH-WESTERN C APE

H uman o ccupation o f t he s outh-western C ape d ates b ack a s l east a s f ar a s t he E arly S tone A ge: t here a re a n umber o f A cheulean a nd M iddle S tone A ge

s ites

s cattered

t hroughout

t he

r egion

b ut

t hey

h ave

r eceived

r elatively l ittle a rchaeological a ttention ( Inskeep 1 978 ).

T he v ast

b ulk o f a rchaeological r esearch h as b een d irected t owards t he L ater S tone A ge , t he last t en t o t welve t housand y ears o f p rehistory . w ork h as

r ecently

P arkington

b een

( 1984)

a nd

e xtensively

T hackeray

r eviewed

( 1981).

T his

b y

J .

s hort

D eacon

T his

( 1984),

s ummary

d raws

c onsiderably o n t heir s yntheses .

T he b eginning o f t he L ater S tone A ge m ore o r less c orrelates w ith t he e stablishment o f t he p resent c limatic r egime a s h igher t emperatures a nd t he m elting of t he large Pleistocene i ce s heets r esulted i n ar ise i n s ea l evels a nd a n i ncrease o f s ome f ive d egrees c entigrade i n g lobal t emperatures

c ompared w ith t he

l ast g lacial

D eacon 1 979; J . D eacon 1 984; Vogel 1 983).

m aximum

( Butzer

1 973 ;

H .J .

T hese e vents c onsiderably

c hanged t he e nvironment a nd w ay o f l ife o f p rehistoric p eople a t t he C ape .

D uring t he c old I ce A ge a bout t hirty k ilometres o f p resently

s ubmerged

c oastal

s helf w as a bove s ea l evel,

a nd w as p robably c overed

w ith a g rassy v egetation s upporting large m igratory h erds o f g razing a nimals.

T hese i ncluded a n umber o f s pecies a nd e ven g enera t hat a re

n ow e xtinct,

i ncluding a g iant C ape h orse

( Equus

c apensis),

a g iant

b uffalo ( Pelorovis a ntiquus) a nd a h artebeest ( Megalotraqus p riscu ), a nd s ome s maller a nimals s uch a s a s pecies o f s pringbok ( Antidorcas a ustralis) ( Klein 1 972,

1 974,

1 978, 1 980).

T hese a nimals w ere h unted

a nd/or s cavenged b y b ands o f h unter-gatherers . e thnographic e vidence t hat o f a nimals

i s

t o

b e

t he b est w ay f or h umans t o h unt

o rganised

i n

o perating i n a n e xtensive t erritory . f ollow h erds a round ,

O ne c an d educe f rom

f airly

b ig ,

m obile

l arge h erds

g roups,

e ach

T his s trategy e nables t he g roup t o

a nd m akes t he b est u se o f l arge q uantities o f m eat

w hen t hey a re obtained .

S ites d ating t o t his p eriod a re n ot n early a s

n umerous a s H olocene s ites ,

p erhaps

1 0

b ecause t he p opulation d ensity w as

l ower a nd/or b ecause m ore u se w as m ade o f o pen a reas w here p reservation w as p oor ,

r ather t han c aves

( H .J .

D eacon

1 976 ) .

A s t he c limate b ecame w armer, t he s ea l evel r ose t o m ore o r l ess i ts p resent l evel, s ubmerging t he c ontinental s helf . i ncreased .

P alaeoecological

T he r ainfall p robably

i nformation f rom a n umber o f s ites i n t he

s outhern a nd w estern C ape s eems t o s how t hat t he v egetation b ecame m ore b ushy ( H .J . D eacon 1 972,

1 979; A very 1 982; J . D eacon 1 984; K lein 1 983;

P arkington

w as

1 984 ) .

T here

a r elatively

s udden t echnological

c hange

r eflected i n a rchaeological s equences a ll o ver s outhern A frica .

W ithin

a bout a t housand y ears, a t 1 2 0 00-11 0 00 B .P ., n ew k inds o f s tone t ool a ssemblages a ppeared .

T hese h ad f ew f ormal t ools ,

m ostly m edium t o l arge

s crapers

( mean

l ength

w hat t here w as b eing

> 20 m m ) .

I rregular c ores

w ith m ore t han o ne s triking p latform a nd l arge q uadrilateral f lakes w ere c ommon ,

a s

w ere

m icrolithic

b eautifully-fashioned b one

f laking

t echniques

a nd

t iny

t ools

a nd

b eads .

r etouched

T he

p ieces

s o

c haracteristic o f t he l ater H olocene w ere v ery r are ( J . D eacon 1 984). F ood r emains a t c oastal s ites i nclude m arine f ish , s ome t errestrial a nimals . b ones

o f

b irds ,

s hellfish a nd

B oth c oastal a nd i nland s ites c ontain t he

s ome o f t he P leistocene

l arge g razers,

b ut

m ore u se w as

s tarting t o b e m ade o f t he s mall b ovids t hat w ere s uch a n i mportant f ood s ource l ater o n .

T he f auna i s t herefore t ransitional b etween t hat o f

t he P leistocene a nd t he H olocene l evels o f a n umber o f i mportant s ites ( H .J . D eacon 1 972,

1 976; K lein 1 974,1980).

g reat m any p lant f ood r emains .

T here d o n ot s eem t o b e a

I nterestingly , w hile t his l ifestyle m ust

r epresent a n a daptation t o c hanging c limate ,

i t p ost-dates b y s ome 3 0 00

y ears t he m ajor c limatic s hift a nd v egetation c hanges , s outhern C ape

( J .

D eacon

a t l east

i n t he

1 984 ).

G radually y et a nother k ind o f s tone t ool a ssemblage s pread o ver s outhern A frica .

T he e arliest d ates f or H olocene m icrolithic a ssemblages a re

a round 1 0 0 00 B .P .

i n Z imbabwe ,

N amibia a nd t he n orthern C ape .

B y 7 0 00

B .P . t hey a ppear i n t he e xtreme s outh o f t he C ape P rovince ( J . D eacon 1 984 ) . s till

I n

s ome a reas

( such a s t he s outh-western C ape )

b eing m ade w hen

t he

s eventeenth c entury A .D .

f irst

E uropean

s ettlers

m icroliths w ere a rrived

i n

t he

T hese s mall t ools i ncluded s crapers ( mean

l ength i n t he m id-Holocene < 20 m m ),

a dzes ,

s egments ,

b ladelets, b orers a nd g rooved a nd b ored s tones.

1

b acked p oints a nd

M any o f t he f ormal

t ools a nd s ome u nretouched f lakes w ere h afted ;

a f ew m ounted a rtefacts

a nd a g reat m any b earing t races o f m astic h ave b een f ound .

T here w ere

a lso b one t ools , t ortoiseshell b owls , o strich e gg-shell b eads a nd w ater c ontainers, d igging s ticks, w ooden p egs, b ows a nd a rrows a nd l eather b ags ,

s andals a nd c lothes

( J .

D eacon 1 984 ).

W e c an s ee f rom s keletal r emains t hat t he p eople w ho m ade t hese t ools w ere t he a ncestors o f t he m odern B ushmen . n ot

u ncommon,

b oth

a ssociated b eads ,

F ood r emains t hat

w e

i n

c aves

o rnaments ,

t ools ,

i nclude a nimal b ones :

r ecognise

f rom

h istoric

a ntelope s uch a s s teenbok , t he m ost

a nd

i n

t he

L ate S tone A ge b urials a re o pen,

a nd

f requently h ave

o chre a nd s ometimes p ainted s tones .

a fter 8 0 00 t imes a re

d uiker a nd r hebok ,

f requently-eaten a nimals.

M ost

B .P .

o nly t hose

f ound .

S mall

s pecies

b rowsing

d assies a nd t ortoises w ere s ites

c ontain p lant

f ood

r esidues, p articularly t he t ough , f ibrous c orm c asings t hat s urround t he e dible

c orms o f a number o f s pecies o f I ridaceae .

c ommon .

P eople a long t he c oast a te f ish ,

a nd s tranded w hales

( see T ables I -IV).

S eeds,

t oo ,

s hellfish , m arine b irds ,

a re

s eals

W e k now f rom t he r ecords o f

e arly E uropean t ravellers i n S outh A frica t hat h oney , c aterpillars, l ocusts ,

t ermites ,

( Sparrman

1 785;

b irds '

e ggs ,

T hunberg

a lthough m any o f t hese

l izards a nd s nakes w ere a lso

1 795;

i tems

B arrow

1 801-1804;

i mportant

W aterhouse

1 932),

l eave n o a rchaeologically-identifiable

r esidues.

T here a re a f ew f eatures p articular t o s ites o f t his t ime p eriod i n t he s outh-western C ape t hat r equire m ention .

E lands B ay C ave

J ohn

C ave

P arkington )

a nd t he n earby T ortoise

( excavated b y

( excavated b y T im R obey )

w ere u noccupied f rom 8 0 00 t o 40 00 B .P . ( Parkington 1 976a , 1 981, R obey

1 984).

T his

h as

b een

t entatively

i nterpreted

1 980,1984 ) a s t he c onsequence o f a t emporary i ncrease

1 984;

( Parkington

i n a ridity ,

m aking

a n a lready m arginal a rea u ninhabitable

( as p roposed f or t he i nterior o f

t he s ubcontinent b y J . D eacon ( 1974)).

A lternatively , i t m ight b e t he

r esult o f a s mall r ise i n s ea l evel .

A" mid-Holocene t ransgression" h as

b een w ell d ocumented b y F lemming ( 1977 ),

a nd i f t he s hellfish-rich r ocks

n ear t hese s ites w ere p ermanently s ubmerged ,

t here m ight h ave b een

l ittle r eason f or p eople t o g o t o E lands B ay ( Parkington 1 984). a bout

F rom

4 0 00 t o 2 0 00 B .P . t hese s ites h ad a c lassic m id-Holocene b acked

1 2

b lade a nd s egment i ndustry ( Parkington 1 976a , o f t he

l arge

s hell

m iddens

a ccumulated a t t his t ime ,

t his

s tretch

o f

Ag reat m any

c oast

p robably

a nd t he s mall s andy d eflation h ollows i n t he

S andveld w ere a lso o ccupied . t o d eal w ith ,

a long

R obey 1 984 ) .

T hese r ather e nigmatic s ites a re d ifficult

s ince t hey u sually c ontain o nly s urface s tone t ools w ith

l ittle o r n o s tratified d eposit o r d ateable o rganic m aterial . s hifting s ands c ontinuously c over a nd r e-expose s ites,

M oreover ,

a nd m ay m ix

t ogether a rtefacts t hat w ould b e c learly s eparated i n u ndisturbed s ites w ith g ood s tratigraphy .

N evertheless ,

M anhire ( 1984 ) f eels t hat m ost o f

t hese w ere o ccupied b etween 4 0 00 a nd 2 0 00 B .P .

a nd n ot e arlier o r

l ater.

T HE S EASONAL M OBILITY H YPOTHESIS

T he f ramework f or m ost o f t he r ecent a rchaeological r esearch i n t he w estern C ape h as b een J ohn P arkington 's s easonal m obility h ypothesis ( Parkington 1 972, 1 976a , 1 980, 1 981, 1 983, 1 984).

M odels o f t his k ind

h ave b een a pplied t o p rehistoric h unter-gatherers

i n

t he w orld f or s ome t ime

v arious

( see f or e xample H iggs e t a l.

p arts

1 967).

o f

T hey

g enerally i nvolve t he p ostulation o f ap attern o f s easonal t ranshumance d esigned t o m ake t he b est u se o f t he g reatest p ossible r ange o f f ood a nd n on-food r esources w ithout over-exploiting a ny o ne p articular i tem . T here a re b oth m aterial a nd n on-material b enefits t o b e g ained f rom s uch al ifestyle : r egular c ontacts w ith o ther b ands a re o f s ocial i mportance , a llowing r enewal o f k inship t ies a nd e xchange o f i nformation a bout d istant f ood r esources . p articular a rea

T his m ay b e e ssential t o s urvival

i f a ny

i s s truck b y a n e cological d isaster s uch a s a d rought .

N ineteenth c entury e thnographic r ecords f rom t he s outhern C ape d escribe s easonal m ovements o f t he l ast b ands o f S an b etween t he c oast a nd t he m ountains ( H .J . D eacon 1 969 , 1 970, 1 976).

T he w estern C ape , w ith i ts

s trong n orth-south e nvironmental z onation a nd t he d iffering r esource p otential o f t he v arious z ones , r eadily l ends i tself t o t he f ormulation o f a s easonal m obility h ypothesis .

T he e vidence f or s uch s easonal m obility i s b ased o n e xcavations a t t wo c ave s ites :

D e H angen i n t he F old M ountain B elt a nd E lands B ay C ave o n

t he c oast ( see M ap I I).

T he f ood r emains f rom D e H angen i ncluded a

n umber o f i ndicators o f s ummer o ccupation ,

1 3

w hile m any o f t hose f rom

E lands B ay s uggested w inter o ccupation .

T he d eposits a t D e H angen c ontained l arge n umbers o f d assie b ones, o f w hich m any w ere t hose o f i mmature a nimals .

D assies h ave a r estricted

b reeding s eason , t he y oung b eing b orn i n l ate O ctober o r e arly N ovember . A s t hey g row, t heir d eciduous t eeth e rupt a nd a re i n t urn r eplaced b y p ermanent t eeth a t v ery s pecific , w ell-defined a ges . a t D e H angen ,

O f t he d assie j aws

4 5% w ere t hose o f y oung a nimals w ith d entition s uch t hat

t hey w ere m ost l ikely t o h ave b een k illed s ome t ime b etween O ctober a nd M ay ( 54% w ere a dult o r a diagnostic i ndividuals).

T here w ere a lso a

l arge n umber o f t ortoises i n t his s ite ( more t ortoises t han a ll t he o ther a nimals p ut t ogether),

a nd t ortoises a re m uch m ore a ctive a nd

h ence e asier t o c ollect i n s ummer t han i n w inter .

A n umber o f t he w ads

o f g rass u sed a s b edding c ontained p lants w ith i nflorescences , g rasses

i n q uestion f lower i n s ummer .

s eeds o f s ummer f ruits ( Nylandtia ,

P lant f ood r emains

E ucle@ ,

O lea

l arge q uantities o f c orm c asings f rom I ridaceae .

i ncluded t he

a nd R hus), T he

a nd t he

a s w ell a s

s tarchy c orms o f

t hese p lants a re a t t heir largest - a nd m ost p alatable - d uring t he s ummer ,

w hen t he p lants a re d ormant .

e aten a s a s taple f ood,

T hey a re t hought t o h ave b een

s ince t hey a re a p redictable r esource t hat

p eople c ould h ave r elied o n f inding e very d ay , p eriods .

H ilary D eacon

( 1976 )

h as

o r c ould h ave s tored f or

s uggested t hat p rehistoric s easonal

m ovements

i n t he s outhern C ape w ere g eared t o t he a nnual

I ridaceae ,

e specially W atsonia ,

c ycle o f

a nd t hat p eople m ay h ave m oved t o t he

c oast d uring t he t imes w hen t he c orms w ere u npalatable ( see C hapter 4 f or a d iscussion o f t he g rowth c ycle o f t he I ridaceae). t oo , i n

h as p roposed ( 1972 , t he

F old

Mountain

P arkington,

1 976a ) t hat t emporary s hortages o f p lant f oods B elt

i n w inter w ould h ave

i ncreased t he

a ttractiveness o f t he c oast .

E lands B ay C ave

i s

s ituated o n t he A tlantic

c oastline a bout s ixty

k ilometres f rom D e H angen .

L ike m ost c oastal

p lant f ood r emains .

i s

i mportant t o n ote t hat p lants a re p reserved

i n t he H olocene d eposits,

a nd a n umber o f w ads o f b edding g rass a nd

I t

s ites ,

t his

c ave h as

f ew

f ragments o f s eaweeds h ave b een r ecovered ( Parkington 1 980), b ut t he I ridaceae

c orm

c asings

f ragmentary a t E lands B ay .

s o

prominent

a t

D e

H angen

are

r are

a nd

A l ikely s ubstitute s taple f ood w ould h ave

b een s hellfish , p articularly t he l impets a nd m ussels c ommon o n t he r ocky

1 4

o utcrops b elow t he c ave . c ave .

V ast q uantities o f t he s hells a re f ound i n t he

T hese a nimals, t oo , a re a d ependable r esource t hat c ould h ave

p rovided t he b asis o f t he p rehistoric c oast-dwellers ' m enu ( H .J . 1 972,

1 976 ,

1 979a ).

1 983;

P arkington

1 972,

1 976a ,

1 976b ,

1 980;

D eacon

R obertshaw

S hellfish w ould h ave b een s upplemented b y t he m eat o f c oastal

b irds a nd s eals,

f ish ,

c rayfish a nd s tranded w hales .

e xtraordinarily r estricted p upping s eason ,

S eals h ave a n

t he f emales

i n a g iven

r ookery g iving b irth d uring t he s ame w eek e ach y ear a nd c onceiving a gain t he f ollowing w eek . t o s outh ,

b ut

T he a ctual c alendar w eek v aries s omewhat f rom n orth

i s g enerally i n N ovember .

M easurements o f t he m andibles

o f j uvenile s eals c an r eveal t heir a ges w ith c onsiderable p recision , a nd h ence t he t ime o f y ear a t w hich t hey d ied .

T he a nimals i n t he E lands

B ay C ave d eposits a re n early a ll y earlings k illed b etween J uly a nd O ctober .

T his s hort t ime p eriod s eems t o b e c onfirmed b y t he a nalysis

o f s easonal g rowth r ings o n a s mall t he s ame s ite

( Fletemeyer 1 977 ).

s ample o f a dult m ale c anines f rom

T here i s s ome d oubt a s t o e xactly h ow

p rehistoric p eople o btained s eals ,

s ince t hey t end t o b reed o n o ffshore

i slands a nd r arely c ome o n t o t he m ainland u nless s ick o r d ying . a nimals a re w ashed a shore, p articularly d uring w inter s torms .

D ead I t i s

p ossible t hat t here u sed t o b e ar ookery n ear E lands B ay a nd t hat t he y oung s eals w ere a ctively h unted .

T hey a re e asier t o c atch t han a dults ,

a nd y earlings a re i n p eak c ondition s ince t hey a re s uckled u ntil t he n ew b lack p ups a re b orn .

I t w ould t herefore m ake s ense f or h unters t o

c oncentrate o n y earling s eals ,

b ut t he p attern i s

i f t he a nimals w ere s cavenged .

T here a re ,

t he s easons a nd a ges a t w hich s eals d ie , n oted t hat :

" some

l ess e asy t o e xplain

h owever , p eaks a nd t roughs i n a nd P arkington

( 1976a :113 )

h as

r ather u ncontrolled p ersonal o bservations a long t he

A tlantic c oast s uggest t hat s eals o f a ll s izes m ay b e w ashed a shore ,

b ut

t hat b lack p up c arcasses w ill b e c lose t o o ff-shore r ookeries , a nd t hose o f n ewly-weaned y earlings

i n r ough w inter

p roportion o f w inter w ashups" .

s eas

m ay

f orm a s ignificant

T he i ncidence o f b lack p ups

i n t he c ave

s ample i s v ery l ow , a nd t here i s n o r eason w hy t hey s hould n ot h ave b een e aten i f t hey w ere a vailable ( as a t N elson B ay C ave o n t he s outh c oast). H ence t he i mplication h ere i s t hat t here w as n o n earby r ookery , a nd t hat t he y earling s eals w ere p robably s cavenged o ff t he b eaches .

1 5

A l ess r eliable i ndicator o f w inter o ccupation m ay b e t he l arge n umber o f b lack m ussels

i n b oth c ave a nd o pen s ites a t E lands B ay .

T hese c an

b e h arvested f rom t he r ocks a ll y ear r ound , b ut P arkington s uggests t hat t hey w ould h ave b een

s omewhat

p henomenon o f " red t ide" .

s afer t o e at

i n w inter ,

S udden " blooms" o r p opulation e xplosions o f

v arious r ed p lankton s pecies o ccur i n t hese w aters . f requent i n s ummer ,

d ue t o t he

T hey a re m ore

s ince t his i s t he s eason d uring w hich t he s outh-

e asterly w inds p revail ,

h elping t o d eflect s urface w aters a way f rom t he

c oast s o t hat d eep , n utrient-rich w ater w ells u p t o r eplace t hem ( Branch a nd B ranch 1 981) . t o m an , w hile

O ne o f t hese o rganisms ( Gonyaulax c atenella ) i s t oxic

a nd i s c oncentrated b y f ilter-feeding s hellfish s uch a s m ussels , g razing

m olluscs

( e .g .

l impets)

a re

u naffected .

G onyaulax

c atenella d oes n ot k ill t he m ussel , b ut e ven o ne c ontaminated m ussel c an k ill a p erson .

S hellfish m ay r emain t oxic f or u p t o f our m onths a fter a

r ed t ide ( Grindley a nd S apeika 1 969),

s o i t i s b etter t o a void u sing

t hem a s f ood f or t he e ntire s eason d uring w hich r ed t ides a re l ikely .

S everal o ther f actors m ay h ave i ncreased t he r ange o f f ood r esources a vailable t o h unter-gatherers a t E lands B ay d uring t he w inter .

T his i s

t he s eason d uring w hich f emale w hales c ome c lose i nshore t o c alve , a nd i t s eems p robable t hat t he w hale m ortality r ate , c arcasses w ashed a shore ,

w ould h ave

a nd t he n umber o f

i ncreased a t t his t ime .

T he b ones

o f s everal s pecies o f b irds t hat a re w inter v isitors t o t he n earby. V erlore V iel

( e .g .

f lamingoes) h ave b een f ound t hroughout t he H olocene

l evels o f t he c ave d eposits .

W inter i s a lso t he t ime w hen t he v iel,

s wollen b y w inter r ains, u sed t o b reak t hrough i nto t he s ea , c arrying w ith i t o rganic m aterial t hat w ould a ttract f ish , s teenbras ( Lithognathus l ithognathus).

e specially t he w hite

T his i s o ne o f t he c ommonest

f ish i n t he c ave d eposits .

T here i s t hus a c onsiderable a mount o f a rchaeological e vidence f or s ummer o ccupation a t D e H angen a nd w inter o ccupation a t E lands B ay C ave . T his i s s upported b y an umber o f a ccounts o f s ummer m eetings b etween e arly E uropean t ravellers

a nd

" Soaqua " o r h unter-gatherers

t he O lifants R iver V alley ( tabulated i n P arkington 1 976a ,

i n o r n ear 1 977 ) .

S ome

o f t hese p eople a dvised V an M eerhoff i n F ebruary 1 661 t o t ravel u p t he v alley o f t he O lifants R iver , r outes i mpractical

s aying t hat l ack o f w ater m ade o ther

( in P arkington 1 977 ) .

6

T his,

t hen ,

w as

t he o riginal

h ypothesis w as p roposed .

f orm

i n w hich t he

P arkington s aw

s easonal

i nland s ites

i n

m obility t he

F old

M ountain B elt, p articularly t hose i n t he O lifants R iver Valley , a nd c oastal s ites a t E lands B ay a s d ifferent s egments o f o ne i ntegrated s ubsistence e conomy .

P eople m oved b etween t he v arious r esource z ones i n

s uch a w ay a s t o e xploit p eaks r esources ,

i n a nimal,

w hile a voiding t roughs .

a nd p articularly p lant,

" People w ill b e a ttracted t o a reas

b ecause b y c omparison w ith o thers t hey o ffer a m ore s ecure s ubsistence b ase a t p articular t imes i n t he s easonal c ycle . a reas m ay n ot f luctuate a t a ll t heir

a ttraction

f luctuate a nd ,

m ay

s easonally i n t erms o f w hat t hey o ffer ,

f luctuate

t herefore ,

M oreover , a lthough s ome

s imply

b ecause

r elatively s o d o t hey "

t heir

c ompetitors

( Parkington

1 977 :156 ) .

T he O lifants R iver V alley , p articularly , w as s een a s a f ocus o f h unterg atherer s ettlement , s ummer

m onths

p roviding a l ife-giving s ource o f w ater i n t he d ry

( when ,

c ongregate t here) .

p resumably,

I n w etter m onths ,

l arge

n umbers

o f

p eople

w ould

p eople c ould h ave m oved o ut

i nto

t he S andveld o r t he K aroo t o t ake a dvantage o f t emporary a bundances o f p lant f oods, a nd p erhaps a nimals a ttracted b y n ew g rass .

T entative

s upport f or t his h ypothesis c omes f rom t he r ock a rt o f t he O lifants R iver V alley ,

w hich s eems t o d epict u nusually l arge n umbers o f s cenes

i nvolving g roup c o-operation ( Manhire e t a l.

I n

r ecent

p apers,

P arkington

m obility h ypothesis m ore

( 1983,

s ecurely

1 984)

1 983) .

h as

f itted

i nto a c hronological

n ow e xpects t hat i t w ill b e f ound t o a pply ,

h is

s easonal

f ramework .

H e

i n t he f orm o utlined a bove ,

o nly t o t he p eriod b etween 4 0 00 B .P . a nd 2 0 00 B .P .

P rior t o t his,

t here w as t he h iatus i n h uman o ccupation o f E lands B ay ,

a nd a fter 2 0 00

B .P .

t he s ystem w as p robably s eriously d isrupted b y t he a rrival o f

d omesticated a nimals a nd p ottery y et

k nown w hether t hese

o r " Hottentots",

s outh-western C ape .

I t

i s n ot

i tems w ere a cquired f rom e lsewhere b y t he

a lready e xisting p opulation , g roup o f p eople .

i n t he

o r w hether t hey w ere b rought b y a s eparate

T he f irst E uropean s ettlers d istinguished t he " Khoi" w ho w ere p ractising p astoralism ,

f rom t he " San" o r

" Bushmen" w ho w ere h unter-gatherers, b ut t he t wo w ays o f l ife w ere t o s ome e xtent i nterchangeable: a p astoralist w ould h unt a nd g ather t o s upplement h is d iet , a nd m ight b e d riven t o d o s o f ull-time i f h is s tock d ied o r w ere s tolen .

H unter-gatherers h unted o ther p eoples ' d omestic

1 7

, t • A s imals , a nimal

w hen t hey c ould ,

( Marks 1 972) .

a s t hough t hey w ere j ust a nother k ind o f g ame

N evertheless ,

t he a dvent o f t his n ew w ay o f l ife

m ust h ave h ad a t remendous e ffect o n t he a ctivities o f h unter-gatherers : s eventeenth c entury t ravellers t ell o f h erds o f t housands o f s heep a nd c attle o n t he f ertile

l ow-lying g rasslands o f • t he s outh-western C ape .

T hese h erds m ust h ave d riven m any o f t he w ild a nimals o ut o f t he b est g razing

l ands .

H unter-gatherers p robably t ook r efuge i n t he m ore

m ountainous a reas, a s s uggested f or t he s outh c oast b y J anette D eacon ( 1972 ), o r a long t he c oastline w here t he e nvironment w as l ess d isrupted .

T his p attern i s c onfirmed b y m any s mall a rchaeological s ites i n t he l ittle r ocky h ills t hat d ot t he S andveld .

M ost o f t hese a re s mall r ock

s helters t hat w ere n ot o ccupied p rior t o 20 00 B .P .

T he d eposits t hey

c ontain h ave w ads o f g rass u sed a s b edding , u sually a round t he w alls o f t he s helter .

I n t he m iddle a re t he a shy r emains o f f ires ,

s hells, p ottery , g rindstones a nd w ood-working a dzes. p lant f oods s uddenly b ecome m uch m ore a bundant . p aintings .

a f ew m arine

T he r emains o f

M any s helters h ave r ock

T his i s t he k ind o f s ite f ound i n t he F old M ountain B elt

b etween 4 0 00 a nd 2 0 00 B .P . , b ut a fter 2 0 00 B .P . t he p attern s preads t o t hese S andveld h illocks ( Manhire 1 984).

A lthough t here m ust h ave

b een s ome o ccupation o f t he d eflation h ollows a fter 2 0 00 B .P . , s ince s ome o f t hem h erders ),

c ontain p ottery ( which m ight h ave b een l eft t here b y

P arkington a nd M anhire s ee t he m ain f ocus o f h unter-gatherer

s ettlement m oving o ut o f t he S andveld a t t his t ime .

T here a lso s eems t o

h ave b een a d ifferent p attern o f e xploitation o f s hellfish a t E lands B ay i n t he p ost 2 0 00 B .P . p eriod .

O f 6 7 c harted m iddens a t E lands B ay ,

o nly f ive a re w ithout ( or h ave l ower l evels w ithout) p ottery . p ottery m iddens t end t o b e v ery l arge i ndeed , s hell

s ituated

o utcrops .

i n o ptimal

m assive a ccumulations o f

l ocations n ext t o s hellfish-rich

r ocky

P ost-pottery m iddens a re s maller , m uch m ore n umerous a nd

f ound s pread o ut over g reater a reas , p articularly o bvious s hellfish b eds . r eliance o n s hellfish ,

n ot n ecessarily r ight n ext t o T he p attern s uggests i ncreased

p robably t o t he p oint w here i naccessible o r

o therwise l ess d esirable s ources w ere h aving t o b e d rawn u pon . p atterns

P re-

o f

p roposals,

s ite

d istribution

s eem

t o

f it

i n

w ith

T hus

P arkington 's

a lthough t he d ifficulty o f d istinguishing h unter -gatherer

s ites f rom h erder s ites h as n ot b een o vercome ,

1 8

a nd i t

i s p ossible t hat

s ome o f t he l ate s hell m iddens c ould h ave b een l eft b y p astoralists r ather t han b y h unter-gatherers

( Hausman

1 98 ( )) .

T his w as t he s ituation a t t he C ape w hen t he f irst E uropean s ettlers a rrived i n t he s eventeenth c entury A .D . b y s hips

T here h ad b een p eriodic v isits

e n r oute t o t he E ast b efore t his ,

a nd t raded w ith s ome o f t he l ocal s eriously d isrupted t heir l ives .

a nd s ailors h ad e ncountered

i nhabitants,

b ut h ad p robably n ot

V ery s oon a fter t he e stablishment o f

t he C olony o f t he C ape o f G ood H ope, t rading a nd h unting p arties w ere s ent o ff i nto t he i nterior , a nd t he t erritory t hey o pened u p w as r apidly a ppropriated b y f armers . p revious

i nhabitants ,

T his l ed t o a c onflict o f

i nterests w ith t he

m any o f w hom w ere k illed a nd t he r emainder d riven

n orthwards s o t hat b y t he e nd o f t he e ighteenth c entury n one w ere i n t he r esearch a rea .

1 9

l eft

C HAPTER 4

I NDIGENOUS F OODS

T his c hapter

i s m eant t o p rovide a b road o verview o f t he k inds o f

i ndigenous p lant a nd a nimal f oods a vailable i n t he v arious r esource z ones o f t he s outh-western C ape . c an b e

o btained

e xcavations, s election

f rom

a nalysis

T he p icture w ill b e a w ider o ne t han o f

f ood-waste

f rom

a rchaeological

b ecause s uch f ood-waste i s t he p roduct o f t wo s eparate

p rocesses .

T he

f irst

i s

t he

c hoice

o f

f oods

m ade

b y

p rehistoric p eople, a nd t he s econd i s t he b ias c aused b y d ifferential p reservation o f t he

v arious k inds o f r emains

i n a rchaeological

s ites .

A lthough t he e cological ( what w as a vailable) a nd t he a rchaeological ( what w as a ctually u tilised ) p ictures w ill u ltimately b e i ntegrated , i s

i mportant t o

c onsider e ach a spect

i t

s eparately b efore e valuating t he

s ignificance o f t he f ood-waste p atterning i n a rchaeological s ites .

H unter-gatherers ,

e ven w hen l iving i n s uch m arginal e nvironments a s t he

K alahari D esert, d o n ot u sually e at a ll t he d ifferent k inds o f f ood a vailable t o t hem . r anking o f f oods,

E thnographic e vidence i ndicates t hat t here i s a w ith a f ew p referred s pecies

a nimals ) p roviding t he b ulk o f t he d iet ,

( of b oth p lants a nd

i .e . a cting a s s taples .

O ther

s pecies m ay b e e aten o ccasionally f or v ariety , o r m ay b e i ntensively e xploited w hen t he p referred f oods a re d ifficult t o g et , d rought .

a s i n t imes o f

P eople t hus c arry a round i n t heir h eads l ong c hecklists o f

p otential f oods , b ut m ay a ctually e ver c onsume o nly a s mall f raction o f t hem ( Tanaka 1 976; L ee 1 978 , r icher ,

1 979).

T he s outh-western C ape i s am uch

m ore v aried e nvironment t han t he K alahari,

a nd s o i t s eems

l ikely t hat p rehistoric h unter-gatherers i n t his a rea h ad a g reat d eal o f c hoice a s t o w hat t hey w ere g oing t o e at .

L ists o f f ood-waste f rom

a rchaeological s ites c ertainly r eflect c onsiderable b iases t owards s ome p lants a nd a nimals a nd a way f rom o thers . a c ertain e xtent, t here

i s

l ikely

A lthough t his d oes r elate ,

t o

t o s ome i tems b eing e asier t han o thers t o o btain , a lso

t o

b e

a n

e lement

o verlooked i n a rchaeological s tudies .

2 0

o f

c onscious

c hoice o ften

P LANT F OODS

H .J . D eacon ( 1976) s uggested t hat,

i n t he s outhern C ape, p eople m oved

a cross t he l andscape i n a s easonal c ycle g eared t o t he a vailability o f I ridaceae c orms , ( 1972 , w as a n

p articularly t hose o f t he g enus W atsonia .

1 976a ) h as p roposed t hat , i mportant

i n t he w estern C ape t oo ,

i nfluence o n p rehistoric b ehaviour ,

P arkington

t his r esource

l eading p eople t o

s pend t he s ummer m onths i n t he r iver v alleys o f t he F old M ountain B elt g athering c orms a s o ne o f t heir s taple f oods , t he c oast .

a nd t he w inter m onths o n

H ere s hellfish w ould p rovide a s e asily-collected s ubstitute

f or p lant f oods a t at ime w hen t he c orms a re u npalatable b ecause t hey a re p utting u p n ew s hoots .

A t t his s tage t he c orm qa re p inkish ,

a v ery b itter r esin a nd u npleasant t o e at .

f ull o f

J ust b efore t he p lant

f lowers, a n ew c orm s tarts t o b e f ormed : t his i s w hite a nd g ood t o e at . I t g oes o n g rowing d uring t he s pringtime w hen t he p lant i s f lowering, a nd i s b est g athered w hen a t i ts m aximum s ize j ust a fter t he f lowers h ave d ied o ff .

T he p lant t hen u ndergoes a p eriod o f d ormancy d uring t he

s ummer , a dnd t he c orms c an a lso b e c ollected a t t his s tage, u ntil n ew g rowth b egins

i n a utumn ( Silberbauer 1 974;

t here a re u ndoubted s easonal

H .J .

p eaks a nd t roughs

D eacon 1 976).

i n t he

T hus

a vailability o f

I ridaceae c orms, a nd t here a re t wo s trands o f e vidence s upporting t he i dea t hat t hey w ere s taple f oods .

T hese a re,

e vidence

a nd ,

f rom

a rchaeological

e arly s ites .

t ravellers,

f irstly ,

s econdly ,

e thnographic

t he

r emains

f rom

A n umber o f e arly t ravellers r ecorded I ridaceae

c orms a s a rticles o f d iet a mongst t he i ndigenous p eoples o f t he C ape, i ncluding K olbe w ho i n 1 685 w rote : c all O nions ( ajuine ),

" The S isyrinchia ,

T ho ' t hey a re n othing l ike O nions ,

o r t aste . . .They h ad b etter b een c all 'd P otatoes , c ome s omewhat n ear .

t he C ape-Europeans e ither i n s hape

f or t hey a re R oots t hat

W hen b oi rd t hey e at m uch l ike c hestnuts .

T hey a re

v ery p lentiful a t t he C ape i n S eptember a nd t he T wo o r T hree f ollowing m onths —their R oots d iffer i n c olour a s w ell a s s ize ; o thers a d ark r ed , b lack

c olour"

d escription a s r espectively . " uientje"

t o

( in

S mith

b eing o f

( of w hich i s t he S ort f rom t he H ills ) o f a

1 966:24-25).

s pecies

S mith

o f S paraxis ,

( 1966)

i nterprets t his

W atsonia a nd

H exaglottis

T he o ld w ord " alum " g ave r ise , t hrough " ajuintje" a nd " uintjie",

I ridaceae s till ( 1795-96 ),

a nd o thers

s ome b eing w hite ,

B arrow

a c ommon

n ame

f or

i n w idespread u se t oday . ( 1801-4 )

a nd L ichtenstein

a n umber

o f

s pecies

S parrman ( 1785), ( 1812-15 )

a ll

o f

T hunberg

m entioned t he

i mportance o f t he " iris" t o l ocal p eople , a nd t he f irst t hree a uthors a lso d escribed i ts u se a s as easonal i ndicator ,

t he t ime o f t he y ear

b eing e xpressed i n r elation t o t he f lowering t ime o f t he " uintjies" .

A lmost w ithout e xception ,

a rchaeological

s ites

i n w hich p lant

r emains

a re p reserved c ontain c orm t unics a nd c orm b ases o f I ridaceae .

C orm

t unics c an o ften b e i dentified t o g enus l evel b y t he c haracteristic p atterning o f t heir i nterlocking r ibs; t he m ost i mportant g enera s eem t o b e W atsonia a nd B abiana i n t he m ountains, a nd M oraea a nd G ladiolus i n t he S andveld ( see T able V ).

T his p robably r elates t o t he d iffering

a vailability o f t hese p lants i n t he m ountains a nd t he S andveld .

A ll

f our g enera a re f ound i n b oth e nvironments, b ut W atsonia a nd B abiana p refer r ocky g round ,

w hereas M oraea a nd G ladiolus d o w ell i n d eep s andy

s oil.

T he p resence o f l arge q uantities o f t hese p lants i n e xcavated s amples i s d ue i n p art t o s heer g ood l uck :

t heir f ibrous c orm c asings a nd c orm

b ases t end t o b e w ell-preserved b ecause t hey a re s o t ough a nd d ry .

I t

i s q uite l ikely t hat t here w ere o ther i mportant p lant f oods t hat w e d o n ot k now a bout b ecause t heir r emains h ave a ll d ecayed a way . j ust o ne e xample ,

G rielum h umifusum ,

s andy a reas o f t he w est c oast ,

a s mall

c reeping p lant c ommon

T hese m ay b e e aten r aw ,

i nto a f lour a nd u sed t o m ake a p orridge t hat

a pparently t astes s imilar t o o atmeal .

T his i s s till t oday a s taple f ood

o f p oor r ural p eople i n p arts o f t he N orthern C ape , b e f ound a ll y ear r ound r emarkably n utritious, p otatoes ( Archer 1 982 ) .

i n

h as t hick s limy r oots t hat c an e asily b e

d ug u p ( far m ore e asily t han m ost I ridaceae). o r d ried a nd g round

T o t ake

s ince t he r oots c an

( Metelerkamp a nd S ealy 1 983).

I t

i s a lso

w ith a k ilojoule c ontent s imilar t o t hat o f G rielum i s ,

n ot h ave a ny h ard o uter c asing ,

h owever , a s oft p lant ,

t he r oots d o

a nd i t w ould b e u nlikely t o l eave

q uantities o f i dentifiable r emains i n a rchaeological s ites .

T his

i s m erely m eant t o

i llustrate t hat

i t

i s q uite p ossible t hat

a rchaeologists h ave o ver-emphasised t he r ole o f I ridaceae c orms i n p rehistoric m enus ,

a t l east i n t he s outh-western C ape .

u ndoubtedly i mportant ,

W hile t hey w ere

a nd p robably d id a ttract f oraging b ands t o a reas

w here t hey w ere a bundant,

i t m ay w ell b e o ver-stating t he c ase t o

s uggest t hat t heir a bsence i n a ny p articular a rea o r a t a ny s eason

i mposed s ignificant r estraints o n p rehistoric o ccupation .

T able V I p resents a l ist o f t he p lants t hat a re k nown t o b e e dible o ccurring

i n t he r esearch a rea .

M uch o f t he

o riginally f rom S mith ( 1966),

w ho ,

i nformation w as e xtracted

i n t he p rocess o f d ocumenting t he

c ommon n ames o f S outh A frican p lants, c ollated m uch o f t he e xisting l iterature o n

e dible

s pecies .

c onfirmed b y o ther a uthors

A g reat

( Mason

1 972 ;

d eal

o f

i t

h as

s ince

b een

L e R oux a nd S chelpe 1 981;

K idd

1 983; F ox a nd N orwood Y oung 1 983), i n c onversation w ith b otanists a nd f armers

o r

by

c omprehensive, a ltogether ,

p ersonal t he

l ist

c ompared

a rchaeological s ites . h uge

d iscrepancy:

e xperiment. i s

w ith

Although

s urprisingly f ewer

t han

l ong

3 0

u nlikely

- over

s pecies

2 00

to

b e

s pecies

i dentified

f rom

A c ombination o f f actors i s r esponsible f or t his

s election

o f

f avourite

s pecies

b y

p rehistoric

c ollectors , p oor p reservation o f s oft p lants i n s ites , a nd t he r elative p aucity o f e xcavated s ites c ontaining a ny p lant m aterial a t a ll - m ost o f t he e xcavated s ites i n t he r esearch a rea a re c oastal s ites , n otoriously p oor i n p lant r emains. k inds

O ther f actors, t oo , i nfluence t he

( and q uantities ) o f p lant f ood r efuse

p articularly d ifficult p roblems.

w hich a re

i n s ites ,

s eeds p osing s ome

A lmost a ll t he f ruits a nd b erries

e aten b y m an a re a lso f avourite f oods o f b aboons a nd b irds, w hich m ay b ring t hem i nto c aves .

I t i s q uite l ikely t hat t his s ort o f f ood , a nd

o thers w hich d o n ot r equire p rocessing o r c ooking ,

w ere o ften e aten b y

h umans a way f rom a reas w e r ecognise a s a rchaeological

s ites .

T hey m ay

h ave b een c onsumed d uring g athering e xpeditions, o r i n t he c ourse o f o ther d ay-to-day a ctivities a nd o ne c an s eldom b e c ertain t hat t he e xcavated r emains e ven o f p lants k nown t o b e e dible a re a r eliable g uide t o t heir s tatus a s p rehistoric h uman f oods. p rudent

t o

c onsider

a w ider

r ange

o f

I t w as t herefore d eemed

p lants

t han

t hose

a ctually

i dentified f rom a n a rchaeological c ontext i n a ssessing t he i sotopic s ignature o f t his c omponent o f p rehistoric d iet .

An umber o f e dible p lants h ave c ommon n ames t hat i nclude t he p refix H ottentots- o r B ushmans/Boesmans. ( Maurocenia f ranqularia

T hese i nclude H ottentots ' c herry

w hich h as a s mall

r ed f ruit),

H ottentotsbrood

( Dioscorea e lephantipes , w hich h as a l arge f leshy t uber ), H ottentotskool ( several s pecies o f T rachyandra

o f w hich t he u nopened g reen f lower b uds

c an b e e aten ),

( Brabejum s tellatifolium ,

H ottentotsamandel

2 3

w ith a lmond -

s haped n uts w hich a re e dible o nce t hey h ave b een l eached i n r unning w ater t o g et r id o f t he c yanide t hey c ontain) a nd m any o thers . a re

a lso

a v ariety

" kukumakranka" ,

o f

n ames

d erived

f rom

K hoisan

w ords

T here s uch

ag eneral t erm f or a ll t he s pecies o f G ethyllis ,

w ith v ery f ragrant e dible f ruits;

" bietou" ,

a s

l ilies

u sed f or s everal m embers o f

t he f amily A steraceae, b ut p articularly C hrysanthemoides m onilifera w hich h as j uicy e dible b erries ;

" kannipr ,

p lant w ith a l arge f leshy e dible f ruit ; k inds

o f A sclepiadaceae w ith b ig

P achypodium ,

a nd " kambro" ,

f rom

c ommon F ockea

t he

K hoisan

u sed f or s everal

( Smith

1 966).

i ncluding

T he s pecific L atin

i n t he r esearch a rea ,

n ame

a p arasitic

f leshy e dible t ubers,

B rachystelma a nd , p articularly , F ockea .

n ame o f t he m ost d erived

H ydnora a fricana ,

F .

T hese

c omaru ,

i s

t antalising

a ssociations o f K hoisan p eople w ith p articular p lants p robably o ften r eflect

s ome

r eal,

i f

e xtinct

p reoccupation

w ith

the

s pecies,

p articularly i n t he c ases w here t he K hoisan w ord h as s urvived .

T he p lants

l isted

i n

T able

V I

w ould

n ot

a ll

h ave

i mportance i n t he d iets o f p rehistoric p eople .

b een

a ttempt a ny s uch a ssessment . w idespread ,

w hile

o thers

t ubers

a nd

s eeds

c ontribution

B roadly s peaking , a re

h owever ,

a bundant a nd

m ore p articular a bout t heir h abitat;

f ormer p robably m ade a g reater t otal t han d id t he l atter .

i mpossible t o

a nd t his s tudy d oes n ot

C ertain s pecies a re ,

a re

e qual

U nfortunately , e ven

r ough e stimates o f t heir r elative s ignificance a re a lmost m ake w ith t he k nowledge w e h ave a t p resent,

o f

" better"

t o t he

t he

K hoisan d iet

s torage o rgans s uch a s c orms a nd

f oods

( in

t hat

t hey

p rovide

m ore

k ilojoules) t han l eaves, f lowers o r f ruits, a lthough t he l atter m ay c ontain i mportant v itamins o r b e c onsumed a s as ource o f w ater r ather t han o f e nergy .

T he c omments t hat f ollow a re b ased l argely o n p ersonal

o bservations.

O f t he p lants l isted i n T able V I a s h aving e dible f lowers o r n ectar , o nly t hree c ases a re t he a ctual

f lowers e aten .

C rassula a lpestris ,

i n a

d rylands s pecies f ound i n t he K aroo , h as a p eppery f lower-stalk e aten a s ad elicacy b y c hildren . b een

i mportant

f oods

T he t wo p lants i n t his c ategory t hat m ight h ave i n

prehistory

( waterblommetjies ) a nd T rachyandra s pp .

a re

Aponogeton

d istyachos

T he y oung f lowers o f b oth t hese

p lants a re u sed t oday a s g reen v egetables, o ften a s a n i ngredient i n m eat s tews .

T hey a re w idespread ,

b ut a vailable f or o nly a s hort t ime

2 4

i n

s pring o r e arly s ummer . t hey a re i n s eason .

C onsiderable q uantities c an b e c ollected w hen

S ome n utritional d ata o n T rachyandra i s a vailable ,

i t h as a s urprisingly h igh p rotein c ontent a nd k ilojoule v alue ( Archer 1 982 ) .

T he o ther p lants

l isted m erely c ontain s mall a mounts

o f

s weet

n ectar w hich c an b e s ucked o ut o f t he f lowers .

I n t he g roup w ith e dible l eaves o r s tems, a ll t he A sclepiadaceae a nd A lbuca

a re

i mportant

m ainly

a s

s ources

o f w ater .

A sclepiadaceae m entioned a re f ound m ostly a vailable a ll y ear r ound , b ecome

i n t he

s pecies

K aroo .

o f

T hey a re

b ut s ome ( e .g . H oodia a nd C aralluma)

v ery b itter a fter p rolonged d ry p eriods

1 983).

T he

( Metelerkamp a nd

c an

S ealy

A lbuca i s a vailable o nly i n t he s pring i n t he f oothills o f t he

m ountains a nd o n t he c oastal p lain . b ases o f t he s tems o f P hragmites,

F lour c an b e m ade f rom t he p owdered P rionium a nd T ypha ( Smith 1 966 ),

a ll

o f w hich a re c ommon a long r ivers a nd s treams e verywhere i n t he s outhw estern C ape ,

a nd c an b e f ound a ll y ear r ound .

T he l eaves o f H ypertelis

s alsoloides a nd O xalis s pp . a re c hewed r aw o r , c ooked i n s tews .

i n t he c ase o f O xalis,

O ther s pecies m entioned h ave g reen l eaves t hat c an b e

u sed a s v egetables .

S pecies

w ith

e dible

r oots,

c orms

o r

t ubers

d istribution , a vailablity a nd p alatability .

vary

w idely

i n

t heir

I ridaceae a re p lentiful a nd

w idespread b ut w ith s trong s easonal f luctuations i n a vailability a nd p alatability . w idespread ,

O xalis s pp . a nd G rielum h umifusum a re a lso c ommon a nd

a nd

s eem

l ess

s ubject

t o

s easonal

c ycles.

h yacinthoides i s a bundant a ll o ver t he r esearch a rea t oday , n oticeable w hen

i t

f lowers

i n

s pring,

b ut

s ince

i t

I t h as l arge c orms a nd

a nd i s v ery

f lourishes

d isturbed g round s uch a s a bandoned f ields ( Archer 1 982), b een l ess c ommon i n t he p ast.

C yanella

o n

i t m ay h ave

i s t oday s till

e aten a s as taple i n p arts o f N amaqualand ( Archer 1 982).

A nnesorhiza

a nd C yphia a re m ainly K aroo a nd c oastal p lain g enera , p lants o ften o ccur i n g roups b ut a re v ery d ifficult t o r ecognise e xcept w hen t hey p ut u p l eaves a nd s tems i n t he s pring .

B rachystelma i s t oday v ery r are i ndeed ,

b ut F ockea c omaru i s s till q uite c ommon

i n t he K aroo a nd w ell-known a s

a n e dible p lant.

y ear r ound ,

e lephantipes .

I t c an b e f ound a ll

T he t ubers o f t his p lant ,

b een r ecorded ( Archibald 1 967), p rehistoric p eople ,

a s c an D ioscorea

o f w hich 7 00 l b .

s pecimens h ave

m ust h ave p rovided o ccasional f easts f or

b ut t hey a re s low-growing a nd r equire a f airly

s pecific h abitat ( Bayer p ers. c omm .), s o w ere p robably n ever a s taple f ood .

A llium ,

A sparagus a nd P elargonium a re a ll

w idespread a nd c onsistently a vailable ,

s till r elatively

a lthough A llium i s m ore v isible

d uring i ts w inter/spring g rowing s eason .

F ruits

o f t he p lants

T able

V I p robably m ade w elcome

o ccasional a dditions t o t he K hoisan d iet.

M ost a re available o nly

s easonally ,

m entioned

i n

u sually i n l ate s ummer o r a utumn ,

a lthough o dd b erries c an

b e f ound o n s pecies s uch a s C olpoon c ompressum d uring m uch o f t he y ear ( Mason

1 972 a nd o wn o bservations).

N ylandtia s pinosa

S andveld s pecies v alued f or i ts s weet R hus

j uicy b erries r ich

i s a c ommon i n

V itamin C .

E uclea a nd C arpabrotus a re v ery c ommon a ll o ver t he c oastal

p lain a nd F old M ountain B elt .

T he p lant f ood c omponent o f t he K hoisan d iet p robably p rovided t he b ulk o f t he f ood e aten , w ith o ccasional m eat f rom s uccessful h unts o r , f rom

d omestic

s tock .

T his

p attern

i s

c ommon

g athering s ocieties i n m iddle l atitudes ( Lee 1 968 ),

t o

m ost

l ater ,

h unting a nd

a nd i s c onfirmed f or

S outh A frica ( at l east f or p astoralist/hunter-gatherers i n h istoric t imes ) b y e thnographic a ccounts s uch a s t hat o f T en R hyne :

" These

( bulbs

a nd r oots) a re d ug u p b y t he w omen, f or i t i s t he w omen w ho p rovide t he f ood f or t he m en ,

t he m en l ook a fter t he h uts a nd t he h erds o r e lse a re

o ccupied i n w ar" ( Schapera 1 933:129).

K olbe a lso d escribed h ow w omen

c ollected p lants: " Every m orning , e xcepting w hen h er h usband g oes a h unting o r c ollect

f ishing,

c ertain

w hich h appens n ot

r oots .-for

t he

v ery o ften,

s ustenance

o f

s he g oes o ut t o

t he

f amily"

( Kolbe

1 731:160).

S mall a nimals s uch a s t ortoises a nd d assies ( rock-rabbits) w ere a lso g athered r ather t han h unted ,

a nd t hese a re t he m ost

i dentified f rom w estern C ape s ites ( see T able I ) .

c ommon a nimals

L ike p lants ,

t hey a re

r elatively d ependable r esources t hat p eople c ould r ely o n t o k eep t hem g oing

f rom d ay t o d ay ,

P arkington

( 1976a )

s upplemented b y o ccasional

t abulates

n ine

l arger a nimals.

r eferences t o K hoisan f oods

t ravellers a t t he C ape b etween t he 1 650 's a nd 1 820 's .

b y

e arly

O f t hese, e ight

m ention r oots o r p lant f oods, f ive r efer t o i nsects, c aterpillars o r a nts '

e ggs

a nd

f ive

t o

b ig

g ame,

9E ,

u sually

a dding

t hat

i t

i s

o nly

o ccasionally t aken .

I n a ddition ,

t ortoises,

d assies a nd h oney a re

s pecifically m entioned .

A NIMAL F OODS - T ERRESTRIAL A NIMALS

T ortoises a re s till v ery c ommon i n t he w estern C ape , v isible

a nd a re e specially

i n s pring a nd s ummer w hen t hey a re m ost a ctive .

T he

vast

m ajority a re C hersina a nqulata , t he a ngulate t ortoise w hich o ccurs a long t he S outh A frican c oast f rom E ast L ondon t o t he m outh o f t he O range R iver .

I ts p referred h abitat

i s c oastal s andveld ,

b ut

i t

i s f ound a ll

o ver t he r esearch a rea e xcept i n v ery h igh m ountainous r egions ( Greig a nd B urdett

1 976 ).

A verage

l ength

i s a bout

1 5-20

c entimetres a nd e ach

a nimal w eighs u p t o f ive p ounds ( Loveridge a nd W illiams 1 957) s lightly s maller t han f emales).

( males a re

A lthough t hey a re n ow a p rotected

s pecies , m any p eople s till c onsider t he m eat a d elicacy .

S mall n umbers o f o ther t ortoises a lso o ccur i n t he w estern C ape: t he n orthern p art o f t he r esearch a rea i s t he s outhern l imit o f t he r ange o f H omopus

s ignatup _ ,

t he p adlopertjie .

w ould p rovide m uch u ncommon .

l ess

P sammobates

m eat

T his

t han

g eometricus ,

C .

i s a s mall f lat t ortoise , a nqulata a nd

i s

t he g eometric t ortoise ,

t o t he s outh-western C ape ( Greig a nd B urdett 1 976).

i n

a ny i s

i t

c ase

e ndemic

I t i s n ow a v ery

r are , e ndangered s pecies a lthough i t m ay f ormerly h ave b een m ore c ommon . N o e xamples o f t hese s pecies h ave b een p ositively

i dentified f rom

a rchaeological s ites , a nd i t s eems l ikely t hat t he b ulk o f t he t ortoises e aten b y p rehistoric p eople w ere C .

a nqulata .

i dentified f rom a lmost e very a rchaeological

s ite

T ortoises h ave b een i n t he r esearch a rea .

I n s ome c ases ( e .g . a t D e H angen) t here a re m ore t han t hree t imes a s m any t ortoises a s a ll t he o ther a nimals p ut t ogether .

A ngulate t ortoises c an e asily b e c ollected i n q uite l arge n umbers , e ven t oday , i n s pring a nd s ummer .

I n w inter t hey a re m uch l ess a ctive a nd

m ore d ifficult t o s ee , b ut s till a vailable .

A dditional e vidence o f t he

r eliance S tone A ge p eople p laced o n t hem a s af ood s ource c omes f rom t he w ork o f K lein a nd C ruz-Uribe ( 1983),

w hose m easurements o f t ortoise

b ones f rom t he s outhern C ape s ites o f D ie K elders a nd B yeneskranskop i ndicates t hat t he p opulation m ay h ave b een " farmed d own" s ince L ate P leistocene/Early H olocene .

t he

T ortoises d ating f rom t his p eriod a re

2 7

l arger ,

o n a verage ,

t han t hose f rom t he L ate H olocene ,

s uggesting t hat

o ver-exploitation l ed t o ag eneral r eduction i n t ortoise s ize a fter a bout 6 0 00 B .P .

D assies ( rock-rabbits o r h yraxes: P rocavia c apensis) a re a lso c ommon . T hey a re s ocial

a nimals t hat l ive i n c olonies o n m ountains o r r ocky

o utcrops ( Bigalke 1 978 ) a ll o ver S outh A frica . a nd a nimals c an b e h unted a ll

y ear r ound ,

T he m eat i s g ood e ating , p eople d o t his t oday b y

c utting t he a nimals o ff f rom t he r ocks a nd r unning t hem d own, c hasing t hem

i nto

c revices

o r h unting

t hem w ith d ogs .

A n umber

o f

e arly

t ravellers ' r eports m ention m eeting " Soaquas" o r " Sonquas" o r " Somquas a lias B ushmen" w ho c arried r oasted r ock-rabbits w ith t hem ( Waterhouse

1 932 :117-118 ,

T hom

1 958 :315 ,

3 49 ,

3 81,

3 82).

i n s kin b ags T hey w ere o ne

o f t he i tems u sed t o b arter f or t obacco .

O f t he

l arger a nimals

i dentified f rom a rchaeological

a ntelope o f t he g enus R aphicerus a re b y f ar t he m ost I ).

c ommon

s mall

( see T able

T hese a re t erritorial a ntelope t hat l ive i n p airs, a nd a re s till

f airly w idespread i n t he w estern C ape t oday . a n

s ites,

e ndemic s pecies

i n t he f ynbos

m elanotis

( Bigalke 1 978),

( steenbok ) o ccurs a ll o ver s outhern A frica , t he g rey d uiker .

R .

b ut R .

( grysbok )

c ampestris

a s d oes S ylvicapra g rimmia ,

G rysbok a nd d uiker a re b rowsing a ntelope ,

b ut s teenbok

a re m ore o pportunistic f eeders a nd e at g rass a s w ell a s b ushes , a vailable

( Dorst a nd D andelot 1 970,

t hat t he a rchaeological

i s

A dams 1 980) .

i f i t i s

K lein ( 1981) b elieves

s pecimens o f t hese t hree s pecies w ere p robably

c aught b y s etting s nares a long t heir h abitual r uns t hrough t he b ush . T heir a ge p rofile r esembles t hat o f a l ive p opulation ,

w hich m eans t hat

t he h unting t echnique u sed d id n ot d ifferentiate a mong a nimals a ccording t o t heir a ges, a nd s naring i s t he e asiest w ay t o c atch n on-gregarious s edentary a nimals .

R aphicerus

a nd

S ylvicapra a re

H olocene t han i n L ate P leistocene d eposits . s how

a ny

d ecrease

i n

s ize

t hrough

U ribe 1 983) a nd s hellfish ( Buchanan e t a l .

O ther a nimals

I nterestingly ,

t ime t hat

e xcessive e xploitation o r " farming d own" ,

m uch m ore c ommon

t hey d o n ot

m ay b e a ttributed t o

a s t ortoises

( Klein a nd C ruz-

1 978 ) d o .

i dentified i n r elatively l arge n umbers f rom w estern

s ites i nclude h ares a nd d une m ole r ats. h are i n t he r esearch a rea ,

C ape

T here a re s everal s pecies o f

o f w hich t he C ape h are ( Lepus c apensis)

2 8

i n

i s

t he m ost c ommon .

I t i s f ound a ll o ver A frica i n s avannah a reas, b eing a

g razing r ather t han a b rowsing a nimal . k ilograms ( Smithers 1 966 ).

L arge s pecimens w eigh u p t o t wo

H ares a re s olitary o r l ive i n p airs , a nd a re

p robably a ll t erritorial ( Bigalke 1 978).

D une m ole r ats ( Bathyergus

s uillus) a re b urrowing r odents a bout t he s ize o f a l arge g uinea p ig a nd a re v ery c ommon i n s andy s oils . w estern C ape

( Bigalke

l eaves a nd g rasses . t hem

o ut

o f

t heir

1 978 ) .

i s e ndemic t o t he

T hey e at I ridaceae c orms ,

s outh-

t uberous r oots ,

W ith a l ittle p ractice, o ne c an t rap t hem o r d ig b urrows,

a nd d uring w et

s ometimes w ashed o ut a nd c an b e c omm .) .

T his s pecies

C ape w inters,

c ollected o n t he

s urface

t hey a re

( Jarvis p ers .

T hey a re s till e aten i n c ountry a reas t oday .

T he o ther a nimals l isted i n T able I o ccur i n s uch s mall n umbers t hat t hey p robably f eatured o nly o ccasionally i n t he K hoisan d iet .

T he s mall

c arnivores

j ust

( mongooses a nd

j ackals )

a nd t he b aboons

m ay w ell

h ave

d ied i n t he s ites ( most o f w hich a re c aves) r ather t han b een t aken t here b y p rehistoric p eople .

P orcupines a re p resent i n s mall n umbers ,

q uite a l arge p roportion o f s ites . a nimals,

T here a re r emarkably f ew r eally b ig

o f w hich e land a re t he m ost c ommon .

t hat K lein ( 1978,

b ut a t

T his

i s a d ocile

s pecies

1 981) p roposes c ould h ave b een d riven i nto t raps o r

o ver t he e dges o f c liffs .

O ne c ould a rgue t hat t he s mall n umbers o f

l arge a nimals m ight b e t he r esult o f b utchering p ractices ( for i nstance , c utting t he m eat o ff t he b ones a t t he k ill s ite s o a s n ot t o h ave t o c arry h eavy b ones b ack t o t he c ave),

p erhaps,

o f t he d eliberate

s mashing o f l arge b ones f or t he m arrow t hey c ontain .

W hile t hese m ight

b e

c ontributing

f actors,

t hey

o r,

s hould

a lso

h ave

a pplied

i n

t he

P leistocene a nd t he d iminishing r atio o f l arge t o s mall a nimals f rom P leistocene t o H olocene f aunal a ssemblages d oes s eem s trong e vidence o f a r eal t he

s hift

f auna

o f

i n d ietary e mphasis . c oastal

s ites,

S eals a re a c onsistent

a nd

s tranded

w hales,

c omponent o f

a lthough

o nly

i ntermittently a vailable , w ould h ave p rovided h uge a mounts o f f ood .

T able I t hus

i ndicates h ighly s elective h unting o f a nimal f oods

L ater S tone A ge . f rom

i n t he

D assies c onstitute 1 9% o f t he i dentifiable m ammals

a rchaeological

s ites

i n

t he

r esearch

a rea ,

R aphicerus

S ylvicapra 2 2% ( 30% i f t he u nidentified s mall b ovids a re i ncluded ), m ole r ats 1 2% a nd h ares 6 % .

a nd d une

A ny o ther s pecies c ontributes l ess t han 5 % .

A lthough a ll t hese a nimals a re s mall,

2 9

a nd a g reat m any d assies w ould b e

r equired t o p rovide a s m uch m eat a s as ingle e land , t he s mall a nimals a re c onsistently p resent i n a ll t he s ites , w hile t he l arger a nimals a re n ot .

T his c an p robably b e t aken t o d emonstrate t heir g reater i mportance

a s s taple f oods .

B IRDS

S trong

p atterning

a rchaeological

i s

a lso

o bvious

s ites ( Table I I) .

i mportant a rticle o f d iet:

c aves

S ome b irds ,

t he

b ird

s pecies

f ound

i n

L and b irds d o n ot s eem t o h ave b een a n

t he o nly s pecies t hat o ccurs r egularly

( although i n v ery s mall n umbers) c apensis .

i n

i s t he C ape f rancolin F rancolinus

p articularly s mall p asserines a nd c rows ,

n est i n

( McLachlan a nd L iversidge 1 978) a nd t hose f ound i n s ites a re

p robably n ot p rehistoric f ood r emains .

O n t he w hole ,

s ay t hat m ost l and b irds w ere n ot h unted o n a ny s cale ,

i t s eems s afe t o a lthough t hey m ay

o ccasionally h ave b een s cavenged , o r a s ick o r i njured b ird k illed .

T he

p icture i s v ery d ifferent w hen i t c omes t o c oastal b irds: c ormorants, p enguins

a nd

g annets

( in

t hat o rder ) p rovided a s ignificant p roportion

o f t he d iet o f L ate S tone A ge c oastal p eople .

T he

C ape

c ormorant

( Phalacrocorax

s outh-western C ape c oast .

A very

w hite-breasted c ormorant

i s

a bundant

a ll

I t e ats p elagic s urface f ish ,

o ther t hree s pecies o f c ormorant , a nd L iversidge 1 978 ) .

c apensis)

( P .

w hich a re

u nlike t he

i nshore f eeders

( 1981) p oints o ut t hat P .

a long t he

( McLachlan

c apensis a nd t he

c arbo) a re t he o nly t wo m embers o f t he

g roup t hat r egularly s pend t ime o n t he s hore ;

t he o ther b irds

l ive a nd

b reed o n o ffshore i slands a nd d o n ot u sually c ome o n t o t he m ainland u nless s ick o r d ying .

T his m eans t hat,

a lthough P .

c arbo m ight h ave b een h unted , t he o ther c ormorants ,

c apensis a nd P .

p enguins a nd g annets

a re m ore l ikely t o h ave b een s cavenged f rom t he b eaches . n umbers

o f d ead o r d ying b irds a re w ashed a shore:

C onsiderable

8 0% o f a ll

m ortality a t E lands B ay o ccurring b etween N ovember a nd A pril. s easonal

p eak

i s m ore p ronounced f or s ome s pecies

( Morus

b ird T his

c apensis) t han

f or o thers ( Spheniscus d emersus), d epending o n h ow m arkedly s easonal t heir

b reeding

l uvenile b irds , d eaths . p resent

p attern

i s.

a nd b reeding

O ther m arine b irds i n s mall n umbers:

M uch s tress

o f

t he

m ortality

i s a c ontributing f actor

s uch a s a lbatrosses ,

t hese s pecies ,

' 2C 1

o ccurs

t oo ,

a mongst i n

a dult

p etrels a nd p rions a re

c an b e s cavenged o ff w est

c oast b eaches .

A f ew r epresentatives o f t he a vian f auna o f e stuaries,

l agoons a nd r ivers a re f ound a t n earby s ites . p elicans

T he d ucks , f lamingoes a nd

f rom E lands B ay p robably c ome f rom t he V erlore V iel ,

a nd t he

p elicans a t S tofbergsfontein f rom L angebaan L agoon .

F ISH

M arine f ish a re w ell r epresented i n a ll c oastal s ites .

T he d ominant

s pecies o n t he w est c oast a re h ottentot ( Pachymetopon b lochii), w hite s teenbras

( Lithognathus l ithognathus),

g lobiceps)

a nd

h aarders

s tumpnose a re m arine

( Mugil

s pecies ,

a nd

w hite s tumpnose ( Rhabdosargus

L iza

s pecies).

b ut t heir y oung s wim

S teenbras

i nto e stuaries

a nd s oon

a fter h atching , a nd r emain t here u ntil t hey a re s exually m ature, w hen t hey r eturn t o t he s ea . s alinities,

f rom a lmost f resh t o s ea-water , a nd a re c ommon ( and m ost

e asily c aught) t he s ea: r ocks,

H aarders c an t olerate a w ide r ange o f w ater

i n l agoons a nd e stuaries .

T he h ottentot i s o nly f ound i n

i t i s a n i nshore s pecies u sually c aught w ith a l ine o ff t he

o r i n k elp b eds

( Poggenpoel p ers .

c omm . ) .

J udging f rom t he f igures i n T able I II, e mphasis

t here s eems t o b e a s hift

f rom s teenbras a nd s tumpnose i n t he n orthernmost s ites

B ay a nd T ortoise C ave ) t o h ottentot i mmediately

t o

t he

e ast

o f

i t

i n

( Elands

i n s ites o n t he C ape P eninsula a nd

( Hout

S mitswinkel B ay C ave a nd R ooiels C ave ) .

B ay C ave ,

S imonstown

M idden,

T his m ay r eflect t he f act t hat

t here i s a l arge e stuary n ear E lands B ay C ave a nd T ortoise C ave , b ut n ot n ear t he P eninsula s ites ( Poggenpoel p ers .

c omm . ) .

M ost o f t he h aarders

c ome f rom S tofbergsfontein , a nd w ere u ndoubtedly c aught i n L angebaan L agoon.

S mall n umbers o f o ther f ish a re p resent : g elb en ( Coracinus c apensis) u sually c aught w ith a l ine f rom r ocks ,

o r o n s andy b eaches .

i s

T he o ther

s pecies l isted m ight a ll o ccasionally b e c aught i n t his w ay , t here b eing v ery

f ew

d eep-water

f ish

i ndeed .

T he

s noek

( Thyrsites

a tun)

a nd

k ingklip ( Genypterus c apensis) f rom E lands B ay C ave a re d eep-water f ish , b ut t here i s a n a rea o f v ery d eep w ater j ust a round t he c orner f rom t his s ite w here a m ountain r ises s traight u p o ut o f t he s ea .

3 1

D eep-water f ish

c ould

c onceivably

c ome q uite c lose

i nshore h ere

( Poggenpoel

p ers.

c omm.).

A t

R ooiels

s ome w armer-water f ish a re e vident.

( Petrus r upestris),

c arpenter

T he r ed s teenbras

( Arqyrozona a rqyrozona)

a nd p oenskop

( Cymatoceps n asutus) a re I ndian O cean s pecies f ound i n a rchaeological s ites a long t he s outh c oast .

F reshwater f ish a re r are K lipfonteinrand , B oontjies r iver , 1 977).

i n s ites .

T here a re t hree L abeo s eeberi

s and- o r m udfish p resumably p resent

f rom

i n t he n earby

a nd c ertainly i n t he s lightly m ore d istant D oom n( Hall

A ndriesgrond , a s ite n ext t o t he O lifants R iver , y ielded t he

b ones o f af ew C lanwilliam y ellowfish ( Barbus c apensis) ( Poggenpoel p ers . c omm .). s mall) f aunal

O ne u nidentified f ish i s r epresented i n t he ( admittedly s ample f rom A spoort,

( Smith a nd R ipp 1 978).

o n t he b anks o f t he D oom nR iver

O n t he w hole ,

f reshwater f ish d o n ot s eem t o

h ave b een a n i mportant f ood r esource i n t he s outh w estern C ape .

S HELLFISH

S hells a re b y f ar t he m ost c ommon f ood-waste a t a ny c oastal s ite i n t he w estern C ape .

T here a re v ast m iddens a long t he g reater p art o f t he

r ocky s hores ( sandy s hores s upport f ew s hellfish) c onsisting a lmost e ntirely o f l impet a nd m ussel m eridionalis)

a nd

v arious

s hells .

s pecies

T he b lack m ussel

o f

l impet

e specially P . g ranatina a nd P . g ranular's , p rehistoric p eople a t t he c oast.

( the

( Choromytilus

g enus

P atella),

w ere a m ajor f ood s ource f or

T able I V d emonstrates t he h eavy

r eliance o f p rehistoric p opulations o n o nly t hree s pecies o f s hellfish . T he p attern i s n ot a s m arked a t R ooiels a nd S mitswinkel B ay , w here a w ider v ariety o f s hellfish i s available i n t he w armer w ater o f t he I ndian O cean .

H ere t he b rown m ussel ( Perna p erna) a nd a w ide r ange o f

P atella a nd o ther p rehistoric d iet,

s pecies

m ade

a s ignificant

c ontribution

t o

b ut a s s oon a s o ne r eaches t he c old w aters o f t he

A tlantic O cean a t H aut B ay C ave, t he d ominance o f C . m eridionalis, P . g ranatina a nd P . q ranularis

i s a pparent.

c ommon w est c oast i ntertidal s pecies o f

T hese w ere t he t hree m ost s hellfish , a nd c ould a ll b e

o btained a t l ow t ide e very d ay , e xcept d uring s torms. T oday , e dulis, t he E uropean m ussel,

M ytilus

i s t he m ost c ommon m ussel i n w est c oast

3 2

k elp b eds ,

b ut

i t h as n ot b een i dentified f rom a rchaeological m aterial

a s y et, a nd s o i s t hought t o b e a r ecent i mportation ( Grant a nd C herry 1 985).

P atella b arbara , P . a rgenvillei a nd P . c ochlear a re u sually

f ound i n d eeper w ater a nd a re c ollectable o nly a t s pring l ow t ides .

P .

m iniata i s v irtually a n i nfra-tidal s pecies ( Branch a nd B ranch 1 981).

T he p ercentage o f C horomytilus i n w est c oast s ites r anges f rom 1 0% t o m ore t han 6 0% o f t he s hell p resent, w hile P atella r anges f rom 2 0% t o n early 9 0% . s hells,

S ome m iddens a re c omposed a lmost e ntirely o f C horomytilus

o thers

a re

n early

a ll

P atella ,

a nd

c ompletely s atisfactory e xplanation f or t his h as

s ome

a re

m ixed .

N o

s o f ar b een a dvanced .

I t h as b een s uggested ( Robertshaw 1 977) t hat P atella m iddens m ight b e t he r esult o f s ummer o ccupations , b y r ed t ides i n s ummer . f eeders , w hen

s ince C horomytilus

i s o ften p oisoned

A s P atella a re g razers r ather t han f ilter-

t hey a re u naffected b y r ed t ides a nd c an s afely b e e aten e ven

the m ussels

a re toxic.

I n

s ome

c ases,

variations

i n the

C horomytilus : P atella r atio m ay s imply r eflect d ifferences i n t he l ocal a vailabilities o f t he s pecies, s heltered r ocks , t he w aves

s ince P atella u sually p refer f airly

w hereas C horomytilus d oes b est o n o utcrops

e xposed t o

( Branch a nd B ranch 1 981).

I t i s o bvious t hat p rehistoric f ood p rocurement i n t he s outh-western C ape w as a nything b ut a r andom p rocess . f ew

i tems a re f ound i n l arge n umbers

s ites ,

I n e ach c ategory o f f oods, a

i n t he m ajority o f a rchaeological

a r ange o f o ther i tems i s f ound i n s maller q uantities ,

a nd t here

a re a g reat m any f oods t hat m ight h ave b een c onsumed , b ut o f w hich w e h ave

n o

r ecord .

T hese

c lear p atterns

h ave b een

s imulated

i n t he

c ollection o f s amples o f e dible p lants a nd a nimals f or c arbon i sotopic a nalysis.

3 3

C HAPTER 5

I SOTOPIC E COLOGY

I n o rder t o u nderstand t he m ajor p art o f t his s tudy , w hich c onsists o f a n

i sotopic a ssessment o f t he d iets o f t he H olocene

s outh-western C ape , o f t he a rea .

i nhabitants o f t he

o ne n eeds s ome a ppreciation o f t he

i sotopic e cology

T his r equires a s lightly d ifferent v iew o f t he e nvironment

f rom t hat u sually h eld b y plant a nd a nimal e cologists. broad

s ubdivisions

of

the

area

i nto

c ategories

I t i nvolves

based

on

p hotosynthetic p athways o f t he p lants a t t he b ase o f t he f oodweb . i n t urn d etermine t he

i sotopic

the T hese

c omposition o f t he b odies o f h erbivores

a nd c arnivores a nd , u ltimately , p eople i n t he a rea .

B ACKGROUND T O T HE C ARBON C YCLE

T he b asis o f t he i sotopic evaluation o f t he f oodweb r ests o n t he f act t hat c arbon e xists

i n n ature

w hich a re a ssimilated a t

i n t hree m ain f orms,

1 00:1:10

1 0

( or r ather ,

1 4

C t o t he a tmosphere)

r adioactive a nd s o d isintegrates w ith t ime , T heir p roportions c an c hange ,

h owever ,

b ut

1 3

c arbon r eservoirs ' 3

s ources o f f ractionation

C

12 C:1 3 C:1 4

C /'

i n t he g lobal

2C

( change

( Libby 1 955). 1 2

C a nd

I 3

i n t he

I 3

i n

1 3

C /1 2 C r atio )

c arbon d ioxide t o a d ifferent

C4

a nd C AN.

C3

p hotosynthesis ,

o r t he C alvin c ycle ,

c arbon a ssimilation .

I t

s tarts

i nto a t hree-carbon m olecule, c ells o f t he l eaf .

14

i s

C

C a re s table . 1 2

C ,

C c ontent o f

r atios r evealed t hat o ne o f t he m ajor

T here a re t hree p hotosynthetic p athways,

a tmospheric

i s a bout

c arbon c ycle ( Bolin 1 974 ).

i s t he p hotosynthetic

f ixation o f a tmospheric c arbon d ioxide b y g reen p lants 1 957).

C ,

C , r eacts a l ittle f aster .

T his p henomenon p roduces s ystematic d ifferences

E arly m easurements o f

14

i n c hemical r eactions s ince

b eing s lightly s maller a nd l ighter than

v arious

C a nd

i t was b efore t he t esting o f n uclear w eapons

a dded l arge a mounts o f

t he

13

s lightly d ifferent r ates b y l iving o rganisms .

Averaged o ut over t he w hole globe, t he r atio o f -

'2 C,

w ith

e xtent .

( Craig 1 953,

e ach f ractionating T hey a re c alled

C3,

i s t he m ost w idespread s ystem o f

t he

c onversion

o f

p hosphoglyceric a cid ,

c arbon

d ioxide

i n t he m esophyll

T his r eaction r equires t he e nzyme r ibulose-1,5-

3 4

d iphosphate . p hosphates

N ext ,

p hosphoglyceric

i n t he p resence

n icotinamide r eactions .

t he

a denine

o f a denosine

d inucleotide

a cid

i s

r educed t o t riose

t riphosphate p hosphate

T hirdly , r ibulose-1,5-diphosphate

( ATP )

a nd

( NADPH)

r educed

f rom

l ight

i s r egenerated a s f ive

t riose p hosphates a re c onverted i nto t hree p entose p hosphates a nd t he c ycle i s r epeated .

T he o verall e quation f or t he r eaction i s :

3 CO2 + 9 ATP + 6 NADPH + 6 H+ + 5 H 20 › T riose-P + 9 ADP + 8 P1 + 6 NADP

w here Pi = i norganic p hosphate A DP = a denosine d iphosphate

( Zelitch 1 979 )

P lants u sing t his p hotosynthetic c ycle i nclude n early all t rees a nd s hrubs a nd t hose g rasses f ound i n t emperate e nvironments, e .g . E urope a nd N orth A merica . w inter r ainfall

A lmost a ll p lants g rowing

a reas

i n s haded f orests a nd

i n

( including w inter r ainfall d eserts s uch a s t hose

o f t he w estern C ape a nd P eru)

a re

C3 .

T he

c ontrolling f actor

i s

a

c ombination o f r adiation a nd l ow t emperatures d uring t he g rowing s eason .

C4

photosynthesis:

d icotyledonous

p lants ,

a nd A maranthaceae , t he

C4

f ixed

A great

many

tropical

c hiefly

m embers

o f t he

t he

m esophyll

P hosphoenolpyruvate p lus a nd t he r eaction o xaloacetate c hloroplasts,

f amilies

u se a nother p hotosynthetic p athway .

o r H atch-Slack p athway, i n

grasses

i nto

n ext

T his i s k nown a s

t he

4 -carbon

c ompound

p lants . )

o xaloacetate .

r educed

t o

m alate

a nd/or

c arboxylase . a spartate

T he

i n

t he

a nd t hen t ransported t o t he b undle s heath c ells w here ( Bundle s heath c ells a re t hick-walled c ells

r ich i n c hloroplasts w hich s urround t he v ascular b undles C4

C henopodiaceae

c arbon d ioxide i s c onverted i nto o xaloacetate ,

d ecarboxylation o ccurs.

o f

a f ew

i n w hich c arbon d ioxide i s i nitially

i s c atalysed b y p hosphoenolpyruvate

i s

and

T he C O2 t hus p roduced g oes

t he r emaining 3 -carbon c ompound

i s

i n t he l eaves

i nto t he C alvin c ycle ,

r eturned t o t he

w hile

l eaf b lade c ells t o

r e-enter t he H atch-Slack c ycle b y r e-carboxylation ( Zelitch 1 979 ) .

T he

C4

p athway c an b e d ivided i nto t hree s ub-types o n t he b asis o f t he

e nzyme m ost a ctive i n d ecarboxylating t he 4 -C o rganic a cid i n t he b undle s heath c ells .

T here a re also c orresponding a natomical d ifferences

( tabulated i n T able V II) .

I n m alate f ormers , t he d ecarboxylation o ccurs

3 5

i n t he

p resence o f

N ADP-malic

e nzyme

( NADP-me ).

I n a spartate f ormers ,

t he e nzyme i s e ither P EP-carboxykinase ( PEP-ck ) o r N AD-malic e nzyme ( NAD-me )

T he

( Ellis e t a l .

p athway t hus

C4

1 980).

i ncorporates t he

i nitial c arboxylation s tep . p lants,

a nd

c ounterparts ,

C3

b ut d iffers

o f

t his

f actor

a nd

t he

CI ,

i n t he

t han i n

lack

o f

C3

l ight

c rops t wice o r t hree t imes a s p roductive a s t heir

C4

i .e .

C O2 t o p lant b iomass

c ycle,

T his i s m ore e fficient i n

a c ombination

r espiration m akes

C3

t hey a re m ore e fficient c onverters o f a tmospheric ( Zelitch 1 979 ).

C AM p hotosynthesis ( crassulacean a cid m etabolism) i s f ound m ostly i n s ucculents .

I t i s n ot r eally a s eparate p athway ,

t he a bility t o s witch b etween e nvironmental c onditions

T he

i mportance o f

t he

f act

t hat

C3

t he

t wo

d ifferent e xtents . 0.

a nd

C3

( Osmond

C4

a nd

C4

c onsisting r ather o f

p hotosynthesis a ccording t o

1 978 ).

p hotosynthesis t o t he a rchaeologist l ies i n

m echanisms

f ix

a tmospheric

I 2

C

a nd

13

C

t o

A tmospheric c arbon d ioxide h as a 6I 3 C v alue o f a bout

T his m eans t hat

i t

i s d epleted i n

C

13

b y s even p arts p er

t housand ( or p arts p er m ille) r elative t o a u niversally r ecognised s tandard ,

t he m arine

l imestone f rom t he P eedee

C arolina ( Craig 1 953,

1 957).

613

C

C

C

s ample

C/

12

i t i s

Cs ample

- 1 )x 1 000° C l

/0

0

12 ,

, standard

D uring p hotosynthesis, b oth C,

L ike a ll m arine l imestones,

0.

s o t hat m ost n atural s ubstances h ave n egative

=(

1 3

13

S outh

v alues , c alculated a s:

13

13

I 3

i n

T he P DB S tandard , a s i t i s c alled , h as

b een a ssigned a v alue o f 0 °/ 0 r elatively r ich i n

f ormation

C3

a nd

C4

p lants f ix

I 2

C

i n p reference t o

b ut t he s election i s c onsiderably m ore s evere i n t he

C3

T his o bviously c ontributes t o t he r elatively l ow y ield o f p lants .

p rocess. C3

c rop

I n t he C alvin c ycle t he t otal f ractionation i nvolved i s a bout

19 ,5 ° /0 0 : t he f oliage o f

C3

p lants h as a n a verage v alue o f 26 ,5°Vo o

( range 20 t o 35 °/ o o)c ompared w ith 12 ,5 °/ 0 0( range 9 t o 16° / 0 0)f or C4

p lants ( Vogel e t a l. 1 978,

v an d er M erwe 1 982).

3 6

T he r ange f or

C3

p lants

i s m ore t han t wice a s g reat a s t hat f or

l ow a s 35°/ 0 0 a re u nusual . a lgae a nd

p lants ,

C4

T hey h ave b een f ound s o f ar o nly i n m arine

i n p lants f rom d ense f orests ,

w here t he t ree c anopy p revents

m ixing o f t he f orest a ir w ith t he f ree a tmosphere . c anopy e ffect)

r esults

n egative o n t he

f orest

b ut v alues a s

i n a g radient o f f loor

( where

T he l atter c ase

i sotopic

r otting

l eaf

( the

v alues f rom v ery l itter

r eleases

i sotopically l ight C O2 )t o v alues w hich a re l ess n egative a t t he l evel o f t he t ree-tops .

V ogel

( 1978a ) h as r ecorded a 61 3 C v alue o f 27°/0 0

f or a ir o n t he f loor o f a f orest i n B avaria .

S imilar v alues h ave b een

r eported f rom A mazonian r ain f orests b y M edina a nd M inchin

( 1980).

O ther f actors b esides l ocal d ifferences i n t he c arbon d ioxide c ycle a lso i nfluence t he 613 p athway .

C

v alues o f p lants,

e ven w ithin o ne p hotosynthetic

T hese i nclude t emperature , m oisture a vailability , s alinity ,

l ight i ntensity a nd t he n utrient s tatus o f t he s oil ( Bender a nd B erge 1 979, H attersley 1 982). o f h abitats t han

T he f act t hat

C3

p lants g row i n aw ider r ange

p lants a nd a re t herefore e xposed t o m ore

C4

v ariation

i n a ll t hese f actors p robably c ontributes t o t heir g reater 6 ' 3C r ange .

C ARBON I SOTOPES I N A NIMALS

T he l ack o f o verlap i n t he i sotopic r atios o f p assing o n o f t hese t racing . 613

C

S ome

c haracteristic

A h erbivore e ating

C3

r atios

C3

t o

a nd

C4

c onsumers

p lants a nd t he a llows

a nd/or C4based f ood w ill

d ietary

h ave t issue

v alues r eflecting t he p roportions o f e ach g roup o f p lants c onsumed . a dditional

f ractionation

t akes

p lace

i n

t he

c ourse

o f

t he

c onversion o f p lant m aterial t o a nimal s kin , h air , m uscle , b one o r o ther t issues ,

a nd t here a re d ifferent f ractionation f actors f or e ach o f t hese

p rocesses.

L iving o rganisms m aintain a c arbon i sotope b alance: t he

t otal n umber o f

I 2

Ca nd

I 3

Ca toms e ntering a n o rganism i n f ood , d rink

a nd i nspired C O2 e quals t he n umber s tored i n b ody t issues , r espired a s C O2 ( DeNiro a nd E pstein a s a c arbon i sotope s eparator ,

1 978 ).

H owever ,

r ecombining

I 2

C a nd

e xcreted a nd

t he o rganism a cts I 3

C i n d ifferent

p roportions d uring a ll t he p hysiological p rocesses r equired t o k eep i t a live .

T he d ifferent f ractionation i n d ifferent t issues p robably h as

t wo c omponents: t he f irst a rising o ut o f t he o rdinary d iscrimination b etween

i sotopes o ccurring

i n e very c hemical

r eaction .

T he

s econd m ay

d epend o n t he d irect " inheritance" o f t he i sotopic s ignatures o f c ertain

3 7

k inds o f f oods u sed p referentially t issues .

i n t he s ynthesis o f p articular

F or e xample , d ietary f ats t hat a re n ot i mmediately r equired a s

a s ource o f e nergy a re p robably c onverted d irectly i nto a nimal f at . E xcess

p rotein

i n

t he

d iet,

h owever ,

m akes

c ontribution t o f atty t issue ( DeNiro 1 977;

a l ess

s ignificant

v an d er M erwe 1 982) .

T hese m etabolic p athways a re n ot a t a ll w ell u nderstood , a s y et , i mplications f or d ietary t racing a re c lear . e qually

t o

a ll

m easurements

t he

o f

p roportions o f

b ody

t hese

t issues

t issues

o f

a nd C4 based f oods

C3

I f n ot a ll f oods c ontribute

c onsumers,

c annot

b ut t he

t hen

s imple

b e t ranslated d irectly c onsumed .

T his

61 3 C i nto

i s o ne o f t he

p roblems t hat w ill h ave t o b e c onfronted l ater i n t his s tudy .

T his u ncertainty a lso c omplicates t he d iet-to-tissue f ractionation f actors t hat a n umber o f a uthors h ave a ttempted t o m easure ( DeNiro a nd E pstein 1 978 ;

L and ,

L undelius a nd V alastro 1 980 ;

V ogel

1 978b ;

M erwe a nd V ogel 1 978; C hisholm , N elson a nd S chwarcz 1 982). g iven s pecies ,

a g iven d iet

i sotopic c omposition .

v an d er F or a ny

s hould p roduce t issues w ith a p redictable

T his i s t rue , b ut w ithin b road l imits:

l aboratory

a nimals f ed o n t he s ame d iet s how i ndividual v ariation o f u p t o 2 °/ c, ( DeNiro

a nd

E pstein

p roportions o f S outh A frica .

C3

1 978).

Vogel

e t

al.

( 1978)

e stimated t he

a nd C4 g rasses i n t he d ifferent v egetation t ypes o f

T hey

v aried

U ngulate t issue s amples

f rom a lmost

( hair ,

h ide ,

b one ,

1 00%

C3

t o a lmost

f lesh a nd f at )

1 00% C4 .

f rom a n umber

o f b iomes y ielded a verage d iet-tissue f ractionation f actors o f + 5 ,3 ° /0 0 f or b one c ollagen ,

+ 3

0/ 0

0 f or f lesh a nd 3 ° /0 0 f or f at ( Vogel 1 978b ) .

T he v alues a re s omewhat d ifferent f rom t hose o btained b y D eNiro f or l aboratory m ice . d ifferences

a nd t his

b etween

c an p robably b e a ttributed t o m etabolic

s hort-lived m ice a nd l arge ,

D ifferences i n d iet p robably a lso p lay a p art : f ractionation f actors f or a nimals w ith t han t hose f or a nimals w ith C4 d iets . m ay

b e

o ne

p aragraphs ,

a spect s ince

a nd C4 g rasses .

o f

t here T he

t he s eem

C3

l ong-lived u ngulates .

t here a re h ints t hat t he

d iets m ay b e s lightly s maller

( Sullivan a nd K rueger 1 981) .

p roblem

d iscussed

t o b e n utritional

i n

c oncentrated i n t hese c ells ,

p revious

d ifferences

l atter a re m ore f ibrous a nd

o f t heir t ough b undle s heath c ells ),

t he

T his

b etween

i ndigestible

t wo C3

( because

a nd s ince m ost o f t he n utrients a re

a re g enerally p oorer f oods f or h erbivores

3 8

t han a l .

g rasses.

C3

1 973;

T heir p rotein c ontent m ay a lso b e l ower ( Caswell e t

C aswell a nd R eed 1 976 ).

M ARINE C ARBON C YCLE

T he c arbon c ycle i n t he o ceans

i s s omewhat d ifferent f rom t hat o n l and .

P rimary

s ources o f c arbon h ere

i nclude

n ot

d ioxide ,

b ut a lso b icarbonates a nd c arbonates .

o nly d issolved C arbonates

c arbon

( e .g . t he P DB

m arine l imestone s tandard ) a re i sotopically v ery h eavy ( enriched i n 1 3

s o o rganisms u sing t hese c ompounds a s t heir s ources o f c arbon t end

C ),

t o b e a t t he p ositive e nd o f t he s cale . v alues : o nes .

w arm w ater o rganisms a re

T emperature i nfluences 61 3 C

i sotopically h eavier t han c old w ater

613 C v alues f or m arine p lankton r ange f rom 25 ,7+2 ,7°/o

0

( mean +

o ne s tandard d eviation) f or s amples f rom w ater w ith a t emperature o f 0 -15°C , t o 20 ,8+1,6°/ 1 968;

D euser 1 970).

0 0

a t 1 5-30°C ( Sackett e t a l . 1 965 ; D egens e t a l .

D ifferences over t he s ame t emperature r ange a re

s lightly g reater f or m arine a lgae

( Parker

i s n ot n early a s w ell-understood a s d iphosphate ( the e nzyme f ound i n c arboxylase ( as

i n

C4

C3

1 964 ) .

A lgal

i t m ight b e,

C

b ut b oth r ibulose

p lants) a nd p hosphoenolpyruvate

p lants) h ave b een i dentified i n s eaweeds

a nd J oshi 1 973; S teemann N ielsen 1 975; T ing 1 976). 613

p hotosynthesis

( Karekar

S ome s pecies h ave

v alues v ery s imilar t o t hose o f t errestrial C3 p lants , w hile o thers

m ore c losely r esemble CI, t ypes E pstein 1 961;

P arker 1 964;

B lack a nd B ender 1 976 ;

( Wickman 1 952;

S ackett e t a l.

1 965 ;

D eNiro a nd E pstein 1 981;

C raig 1 953;

S mith a nd E pstein 1 971; S chell 1 983).

Am ixture o f b oth c ategories c ontributes t o t he d etritus w hich , w ith p lankton ,

P ark a nd

f orms t he b ase o f t he m arine f ood-web .

t ogether

T he a verage 61 3

C

v alue o f t his b ase o bviously v aries f rom p lace t o p lace w ith s pecies a vailability a nd t emperature . o f S outh A frica , 1 976 ), r anging

F or t he A tlantic O cean o ff t he w est c oast

w here s ea t emperatures r ange f rom 8 °C t o 1 6°C ( Simons

t he f irst t ier o f c onsumers a re f rom

19 ,4

t o

15 ,9°/

0

0

( see

f ilter-feeders w ith T able

X III).

6I 3 C v alues

G razing m olluscs ,

c arnivorous f ish a nd f ish-eating a nimals s uch a s s eals a nd c oastal b irds b ecome

p rogressively m ore

p ositive d ue

o ne t rophic l evel t o a nother . m ore d etail

t o

s tepwise

f ractionation

f rom

T he m arine f ood-web w ill b e d iscussed i n

i n C hapter 6 .

3 9

T here i s t hus a n e xtremely w ide r ange o f p ossible 81 3 C v alues i n a ll t he p lants a nd a nimals t hat c an b e e aten b y h umans . g razing o nly o n

C4

g rasses m ight b e a s p ositive a s 6

f at o f a b rowsing a nimal n egative a s

38°/0 0 .

m uch n arrower .

T he m eat o f a n u ngulate °/ a

0 , w hereas t he

i n a t ropical r ain f orest c ould w ell b e a s

I n a ny o ne e nvironment,

h owever ,

t his r ange i s

W e w ill n ow c onsider e ach o f t he f our e nvironmental

z ones i n t he r esearch a rea w ith a v iew t o e stimating , f rom t he d ata o n c limate a nd f lora p resented i n C hapter 2 , t he k inds of p hotosynthetic p athways a nd t he v ariations

i n

i sotopic r atios o ne w ould e xpect i n e ach

z one.

P HOTOSYNTHETIC P ATHWAYS I N T HE S OUTH-WESTERN C APE

V ogel e t a l . v egetation

( 1978 ) e stimated t he p roportions o f t ype

i n

S outh

A frica .

C3

A ccording

s trandveld a nd m ountain f ynbos e ach h ave 0 -5%

C4

a nd t o

C4

g rass i n e ach

t heir

g rass ,

b reakdown,

c oastal

f ynbos

a nd c oastal r enosterveld h ave 5 -25% a nd s ucculent K aroo 9 5-100% ( see M ap V I).

T hese f igures,

n umber o f g rass

h owever ,

r efer t o t he p ercentage o f t he t otal

s pecies p resent

t hat

a mount o f g round c overed b y e ach t ype . b otanical p oint o f v iew , t racing . s pecies

a re

C3

o r

C4 ,

r ather t han t he

W hile t his i s s ensible f rom a

g round c over i s m ore

i mportant

T he o nly c omment o n t his i s i n a later p aper : i n p redominantly

C4

i n d ietary

" the

C3

g rass

a reas a re f ound t o b e m ostly r are a nd

l ocalised a nd v ice v ersa" ( Vogel 1 978b :298).

I n t he s econd p aper , h e

a ctually s implifies t he p revious s cheme i nto t hree c ategories:

> 90%

m ixed a nd > 90%

s tarting

C.

T hese t wo c lassifications p rovide a u seful

C3,

p oint f or a c loser e xamination o f t he a rea .

I SOTOPIC R ATIOS O F A NIMALS I N T HE R ESEARCH A REA

I n t heory , g rass ,

a n u nselective p ure g razer ( i .e . a n a nimal t hat e ats o nly

n ot b ushes ,

a nd c onsumes

e verything

i n

i ts p ath w ithout p icking

o ut s ome p alatable s pecies a nd l eaving o thers b ehind ) w ould r eflect t he r elative a mounts o f g round c overed b y n o s uch a nimal e xists .

C3

a nd

C4

g rasses .

U nfortunately

M any s pecies p reviously t hought t o h ave d efinite

s trong p references f or g razing o r b rowsing a re b eing f ound t o b e m uch m ore o pportunistic f eeders u npublished d ata ) .

( Lee T horp 1 983; L ee T horp a nd v an d er M erwe ,

M oreover ,

t he d egradation o f t he e nvironment

4 0

i n t he

r esearch a rea , a nd t he f act t hat m uch o f i t i s n ow u nder c ultivation, m eans t hat g razers t hat l ived t here i n p rehistoric t imes l onger d o s o .

F inally ,

s pecies, p alatable

a nd a ttribute

C3

( in

g rassland b y a

C4

i t t o t he s elective overgrazing o f t he m ore

g rass b y c attle ,

C3

E vans a nd T isdale

1 973) d escribe t he t ake-over o f a

A .

z ebra ) n o

i t i s q uite p robable t hat m any g razers a re i n

f act h ighly s elective a bout t he g rasses t hey e at : C aswell e t

( e .g .

s heep a nd m ule d eer .

A lthough a ll t hese f actors c omplicate t he I nterpretation o f t he 613 C v alues o f a nimals c ollected i n t he r esearch a rea , o ne w ould s till e xpect a r ange o f v alues - f or a nimals t hat e at s ome g rass - b ecoming m ore p ositive f rom t he 0 -5% t he 9 5-100%

C4

N orth A merica

z one . ( Land ,

m oisture a ffects 813

C4

a reas t hrough t he

T his p attern h as b een d emonstrated f or d eer i n L undelius a nd V alastro

C

v alues

( Tieszen 1 979 ),

i f t here w ere a g radient w ithin t he 0 -5% P eninsula ,

i ntermediate c ategories t o

C4

1 980).

i f s oil

i t w ould n ot b e s urprising a rea f rom p arts o f t he C ape

w ith u p t o 6 00 m m o f r ain p er y ear ,

r eceives l ess t han 2 00 m m ( see C hapter 2 ).

M oreover ,

t o E lands B ay w hich

A lthough m ore o r l ess t he

s ame k inds o f p lants g row i n b oth p laces, c limatic d ifferences c ould w ell r esult i n s mall d ifferences i n t heir i sotope r atios ( see d iscussion e arlier i n t his c hapter a bout t he g reater v ariability i n t he el 3 c v alues o f

C3

p lants

c ompared w ith

C4

p lants).

C limatic a nd o ther d ifferences

( frequency o f f ires, g razing p ressure) m ight a lso r esult i n d ifferent a reas w ithin o ne

v egetation t ype h aving d ifferent a mounts o f g rassland

c ompared w ith b ushy s crub . w ould t hen i ngest m ore b ushy p atches .

C4

A m ixed f eeder

( grazer a s w ell

a s b rowser )

f ood i n g rassier a reas, a nd m ore

C3

f ood i n

H endey ( 1983 ) g ives a c oncise a ccount o f t he p roblem o f

t he r ole p layed b y g rasses i n f ynbos v egetation .

H e s uggests t hat a

m oderate i ncrease i n f ire f requency f avours t he s pread o f g rasslands. I f f ires a re t oo f requent ,

h owever ,

E lytropappus r hinocerotis,

t he r enosterbos) m ay s uccessfully c ompete

w ith g rasses,

i n t he s hrubland c ommunities f ound

c oastal

r esulting

c ertain p ioneer s hrubby b ushes

l owlands t oday ( Heydorn a nd T inley 1 980).

m ammals t hat

u sed t o b rowse o n t hese s hrubs

e land) a re n o l onger f ound i n t he w estern C ape . r educe t he s hrub c over ,

A lso ,

( elephant,

( e .g .

i n t he

t he l arge

r hinoceros,

T hey p robably h elped

t hus p romoting t he g rowth o f s tands o f g rass

( Hendey 1 983).

4 1

Y et a nother c omplication m ay e xist i n

g rasslands .

C4

I t h as b een s hown ,

i n as tudy d one i n N amibia , t hat t he t hree s ubtypes o f d ifferent d istributions .

W hile

C4

g rasses o n t he w hole a re a dapted t o

c limates w here t he t emperature d uring t he g rowing s eason a re f ound i n q uite m oist a s w ell a s a rid a reas. c ommonest i n w etter r egions ,

i s h igh ,

t hey

M alate f ormers a re

w hereas a spartate f ormers f avour d ry a reas .

T he l atter c an b e s ubdivided i nto N AD-me a nd P EP-ck, p redominating

g rasses h ave

C4

i n a reas o f v ery l ow o r u npredictable

w ith t he f irst

r ainfall .

P EP-ck

g rasses d o b est w here t he r ainfall i s a bout 3 50 m m/year ( Ellis e t a l. 1 980 ) .

C ontrolled

l aboratory

e xperiments

s lightly h eavier 61 3 C v alues ,

s how m alate

o n a verage ,

f ormers

t han a spartate f ormers ,

w ithin t he a spartate g roup P EP-ck a re s omewhat e nriched i n w ith N AD-me g rasses t hese

( Hattersley 1 982).

i sotope r atio d ifferences a re

t o h ave

I t

a nd

C c ompared

1 3

i s n ot y et k nown w hether

s ignificant

i n n atural

v egetation .

H attersley d oubts i t , a nd c orrelation o f p ublished 61 3 C v alues f or S outh A frican

C4

g rasses

( Vogel

e t

a l.

1 978;

E llis

e t

a l.

1 980)

w ith

a natomical e vidence f or t heir d ivision i nto t he v arious s ubtypes ( Ellis 1 977) d oes n ot r eveal s tandard

d eviation

s ignificantly large d ifferences

f or

e ach

t ype

i s

a s

f ollows:

( mean + o ne

M alate

f ormers:

12 ,4+1,0 ° /0 0 , n =31; P EP-ck a spartate f ormers : 13 ,1+1,1 ° /0 0 , n =17 ; a nd N AD-me a spartate f ormers: 13 ,5+1,0°/

0 0

,n =14 ).

H owever , t hese s amples

w ere v ery r andomly c ollected f rom w idely s cattered l ocalities , w ith

v ery

d ifferent

m icroclimates.

N o-one h as

s ystematically a long a t ransect f rom w et t o d ry o therwise

u niform

p ossibility o f

C4

e nvironment.

U ntil

subtype a ffecting t he

t his

c arbon

C is

h as

s o

p robably

f ar

s ampled

g rassland i n a n b een d one,

i sotope

v alues

t he

o f t he

f ood-web i n d ifferent a reas m ust b e b orne i n m ind .

T here a re t hus a n umber o f f actors t hat c ould c omplicate t he . 513 o f a nimals

c ollected

i n t he r esearch a rea .

C

v alues

T he s ampling p rogramme w as

s et u p w ith a ll t hese p roblems i n m ind , a nd i n t he h ope t hat t he r esults w ould h elp d etermine w hich f actors,

i f a ny ,

a ctually i nfluence t he

i sotopic s tructure o f t he f ood-web .

I SOTOPIC R ATIOS O F E DIBLE P LANTS I N T HE R ESEARCH A REA

S o f ar , t he d iscussion h as r eferred m ainly t o a nimal s amples .

T here a re

g rey a reas i n o ur k nowledge o f t he i sotope r atios o f p lants a s w ell,

4 2

a lthough t he s ituation h ere i s l ess i nvolved b ecause p lants a re p rimary p roducers r ather t han c onsumers, s o o ne w hole l evel o f f ractionation t hat c omplicates a nimal

v alues d oes n ot e xist f or p lants.

( 5 13 C

a ttempt w ill b e m ade t o i nvestigate t hree m ain p roblems . t he d egree o f

v ariation o f c arbon

a n i ndividual.

T his

T he f irst

i s

i n d ifferent p arts o f

T he v ast m ajority o f t he p ublished

h ave b een o n f oliage . p athway ,

i sotope r atios

A n

m easurements

d I3 C

i s u seful a s a n i ndicator o f p hotosynthetic

b ut v ery o ften t he f oliage i s n ot t he p art o f t he p lant e aten

b y h umans .

F ruits ,

b erries ,

c orms o r t ubers a re i nformation

s eeds ,

f lowers o r s torage t issues

f requently c onsumed ,

available

o n

t heir

i sotope

a nd t here r atios.

i s

l ittle

i s

k nown t hat

d ifferent t issues i n a p lant d o h ave d ifferent 6I 3 C v alues :

L eavitt a nd

L ong ( 1982 ) r eport a c onstant d ifference o f 2 -4°/ a nd w ood o f j uniper t rees .

I t

v ery

s uch a s

0 0

b etween t he l eaves

O ther a uthors h ave d one s imilar m easurements

o f w oody a nd n on-woody p lants ( Craig 1 953;

M ooney e t a l. 1 977).

A ll

f ound d ifferences o f af ew p er m il b etween d ifferent t issues f rom t he s ame i ndividual.

T his i s n ot s urprising w hen o ne c onsiders t hat t he

d ifferent p arts o f p lants c ontain d ifferent p roportions o f v arious c hemical c ompounds .

T hese v ary i n t heir i sotopic c omposition : S mith a nd

J acobson ( 1976) a nalysed a p otato t uber a nd f ound t hat, s lices o f p otato h ad a 6I 3 C r eading o f 25,8 °/ 27 ,1 ° /

0

0 ,

s ugars

23 ,8°/

0

0 ,

0

s tarch 25 ,5°/o

. P ark a nd E pstein ( 1960 ,

w hile w hole

,a mino a cids w ere a nd a C C1 4 e xtract

1 961) a nd D egens e t a l .

( 1968) a lso

d iscovered t hat l ipid f ractions o f p lants w ere m arkedly i sotopically l ighter t han o ther f ractions . o f

C3

V ogel

( 1982) h as m easured t he 6 ' 3C v alues

p lant p roteins a s c ompared w ith l eaves a nd s eeds .

T o c haracterise t he p lant m enu i sotopically , r equire c onsideration .

F irstly ,

i t

a n umber o f v ariables

i s n ecessary t o a nalyse s ome o f t he

f ood p lants o f t he w estern C ape t o f ind t he 6I 3 C v alues o f t heir e dible p arts , T able

r ather t han t heir f oliage . V I) .

A f ew m ay b e C AM ,

e .g .

T he v ast m ajority a re

( members o f t he f amily A maranthaceag). a re c ertainly

C3 ,

p lants

( see

s ome o f t he s ucculent A sclepiadaceae

i mportant a s w ater s ources i n t he K aroo .

I ridaceae ,

C3

O ne o r t wo m ay p erhaps b e

S taple f oods ,

C4

i ncluding a ll t he

b ut i t w ould b e u seful t o h ave s ome i dea o f

t he r ange o f i sotope v alues t he e dible t issues d isplay .

4 3

T he

s econd p oint w orth

i nvestigating

p lants i n d ifferent s easons .

i s t he v ariation i n 61 3 C i n t hese

L owdon a nd D yck ( 1974) f ound a 6 ° 0 0

d ifference i n m aple t ree l eaves i n s pring c ompared w ith l ate a utumn , a nd L eavitt a nd L ong ( 1982 ) r eport a s easonal v ariation o f 30 /0 0 i n j uniper l eaves.

T hese d ifferences c an b e a ttributed t o s easonal t emperature v ariations, a nd h ence t o i ncreased a ctivity i n t he p lant i n s ome s easons w hile i n o thers

i t

i s d ormant .

c oefficient

f or

L eavitt a nd L ong c alculated t hat t he t emperature

t heir

j unipers

w as

a bout

0,27 °/ 0 0

0

C1 .

T his

f luctuation w ould b e p articularly i nteresting i n p lants w ith c orms, b ecause o f t he c onsiderable a lterations i n t he c omposition o f c orms t hrough t he y ear .

S easonal d ifferences i n

613 C w ould t hen b e t he r esult

o f t emperature c hanges a nd o f v ariations i n t he c hemical t he p lants .

T he c ycle b egins a s n ew g rowth i s

c omposition o f

i nitiated i n t he a utumn ,

a nd s tarchy p hotosynthetic p roducts a ccumulate t o f orm a c orm , t hen d ormant d uring t he f ollowing d ry s eason . f avourable e nvironmental c onditions f ormation o f n ew l eaves a nd s tems ,

w hich i s

W ith t he o nset o f m ore

i n t he a utumn i t

i s u sed u p i n t he

w hich i n t urn m ake a nother n ew c orm .

I t h as b een s hown t hat t he l evels o f t he v arious c omplex p olysaccharides t hat

c onstitute t he

s tored n utrients

c hange s easonally ( Waher 1 974 ) . c onsiderable

i sotopic

c hanges

i n t he

c orms o f W atsonia s pp .

T his d ynamic c ycle o ught t o r esult i n

t he

c orms

i n

d ifferent

a lthough o nly a s mall p art o f t he 61 3 C r ange w ould b e

i n

s easons,

i mportant

i n t he

h uman f ood-web i f t hese p lants w ere e aten o nly a t c ertain t imes o f y ear .

A n umber o f p lant f oods a re o f c ourse n ot a menable t o t his k ind o f s tudy , a s t hey a re a vailable f or o nly a s hort s eason .

E dible f lowers

s uch a s t hose o f A ponogeton a nd T rachyandra a nd m any f ruits a nd b erries f all

i nto

t his

c ategory ,

b ut

t here

a re

s pecies

f or

w hich

u seful

i nformation m ight b e f orthcoming f rom a s easonal s tudy .

T he f inal s ource o f v ariation i n t he i sotope r atios o f p lant f oods i s t hat b etween d ifferent e nvironment a nd d ifferent

i ndividuals o f t he s ame

( 2) d ifferent e nvironments.

i ndividuals o f t he s ame s pecies

s pecies

i n

( 1 )

t he

s ame

A ny v ariation b etween

i n t he s ame

e nvironment m ust

b e d ue m erely t o t he s light v ariations i n n ormal m etabolic p rocesses t hat i nevitably o ccur i n l iving o rganis m.

4 4

T his v ariability m ust b e

k nown

s o

t hat

d ifferences

i t

c an b e

b etween

e nvironments .

u sed a s

i ndividuals

a b aseline o f

t he

i n t he m easurement

s ame

s pecies

l ocal d ifferences

t emperature ,

d ifferent

T hese d ifferences h ave b een d emonstrated i n l aboratories ,

b ut h ave n ever b een a ssessed i n n atural e nvironments . t o

i n

o f

i n t he c arbon d ioxide c ycle ,

m oisture a vailability ,

s alinity ,

T hey m ay b e d ue

o r t o d ifferences

l ight

i ntensity o r

i n

s oil

n utrients ( Degens e t a l. 1 968; B ender a nd B erge 1 979; F arquhar 1 980; H attersley 1 982).

I SOTOPIC R ATIOS O F M ARINE O RGANISMS

T he

f inal

T here

g roup o f p rehistoric f oods

c onsists o f t he m arine o rganisms .

i s a c onsiderable a mount o f l iterature o n m arine p lants a nd

a nimals, w ith a l arge n umber o f p ublished 61 3 C v alues ( Wickman 1 952; C raig 1 953, 1 965 ;

1 957; P ark a nd E pstein 1 961; P arker 1 964;

D egens e t a l.

1 968 ;

S mith a nd E pstein 1 971;

S ackett e t a l.

B lack a nd B ender 1 976 ;

T hayer e t a l. 1 978; D eNiro a nd E pstein 1 981; R au e t a l. 1 981; S chell 1 983).

N one o f t hese r efer s pecifically t o t he S outh A frican s ituation ,

h owever , a nd a lthough m any m arine s pecies a re v ery w idespread , t heir i sotopic r atios v ary w ith w ater t emperature ( as d escribed a bove), p H, e tc .

( Deuser a nd D egens

1 967 ).

A t t he t ime w hen t he r esearch f or t his p roject w as p lanned , o nly a p ilot s tudy o f

6 '

c ompleted . M erwe ,

3C

i n t he m arine f ood-web o f t he w estern

C ape

h ad b een

T his c omprised a s eries o f r esults o btained b y v an d er

V ogel a nd R awlinson o n m aterials c ollected i n t he O udekraal k elp

b ed ( Cape P eninsula ) o r r aised i n t he l aboratory b y F ield a nd G riffiths ( Rawlinson 1 982 ) .

T he r esults i nclude p lants a nd a nimals f rom d ifferent

t rophic l evels i n t he k elp b ed f ood-web ,

a nd s howed t hat t here i s q uite

c onsiderable f ractionation f rom t he p hytoplankton

( -17 ,9°/

0 0

)t hrough

f ilter-feeders s uch a s m ussels t o s cavengers r ight a t t he t op o f t he c hain

( crayfish a t 13 ,1 °/ 0 0 ) .

O ne o f t he m ost

i nteresting a spects o f

t his s et o f r esults w as t he v alues f or t wo d ifferent k inds o f k elp ; o ne , E cklonia m axima , h ad a C4like 6 ' 3C v alue ( -12 ,0°/ „) w hereas t he o ther , L aminaria p allida , l ike

w as C3like ( -20 ,9°/

( Giqartina a t 22,0 ° /

0 0

0 0

) .

, P achymenia a t

O ther a lgae w ere a lso 21,9°/

0 0

) ,

C3 -

r aising t he

q uestion o f w hether t he v alue f or E cklonia w as a nomalous o r w hether

4 5

t here i s i ndeed a g roup o f a lgae w ith r elatively l ow

C /1 2 C r atios a nd

1 3

a nother w ith m uch h igher r atios .

T he f irst p roblem i n t he m arine c ycle i s t herefore t o a scertain t he d egree o f 613 C v ariability a mong s eaweeds .

T his i s n ecessary b efore

p roceeding w ith t he a nalysis o f o rganisms h igher i n t he f ood-web , s eaweeds s ea .

( together w ith p hytoplankton ) a re t he p rimary p roducers

I t

s ince i n t he

i s n ot k nown w hether s eaweeds w ere e ver e aten d irectly b y

p rehistoric p eople ,

b ut a n umber o f a nimals f eed o ff t hem ,

a nd t hey

c ontribute a l arge p art o f t he d etritus t hat s ustains f ilter-feeders . T heir c arbon i sotope r atios t hus d irectly d etermine t hose o f a ll t he h igher t rophic l evels i n t he i nshore m arine f ood-web .

T he s econd a spect o f t he m arine c arbon c ycle t o b e f ractionation a t d ifferent t rophic l evels . s ample d id y ield s ome i nformation o n t his, a nimals a t t he t op o f t he f ood-chain . c ormorants ,

g annets ,

i nvestigated

A lthough t he O udekraal i t d id n ot

A lso ,

i nclude m any

T hese , e specially f ish , s eals,

p enguins a nd c rayfish w ere e aten

i n p rehistoric t imes .

i s t he

i n l arge n umbers

a nalysis o f f urther s amples w ill h elp

d etermine t he d egree o f i sotopic v ariation b etween

i ndividual a nimals ,

a nd b etween a nimals c ollected f rom d ifferent l ocalities .

L astly ,

i t i s n ecessary t o d etermine a ny s easonal v ariations i n t he el 3 C

c ontent o f s mall

f ilter-feeding a nimals .

O ne w ould e xpect t o s ee

s easonal v ariations o nly i n s mall o rganisms w hich r eplace m ost o f t heir c arbon

i n

l ess

r epresent a n

t han

a y ear .

i ntegration o f

( The

s everal

61 3 C v alues y ears '

d iet . )

o f

l arger

a nimals

T he

c arbon t urnover

t ime i n b lack m ussels ( or a t l east i n p arts o f t hem ) i s f airly s hort . T his s pecies s pawns t wice a y ear , i ts b ody w eight i n e ach s pawning .

a nd a n i ndividual m ay l ose u p t o h alf C arbon i n t he r eproductive s ystem i s

t herefore r eplaced r egularly , a lthough c arbon i n o ther p arts o f t he b ody m ay

r emain

M ussels,

f ixed

t his

m uch

l onger

b eing f ilter-feeders,

p eriodically d escribed

f or

a fter

t he

i n C hapter 2 .

( Griffiths

p ers .

c omm .).

c onsume t he p lankton t hat " blooms"

u pwellings

o f

c old ,

n utrient-ri d '

T hese u pwellings a re m ore f requent

b eing t he t ime w hen t he

B ranch 1 981) .

p eriods

s outh-easterly w inds p revail

i n

w aters s ummer ,

( Branch a nd

I t i s t herefore p ossible t hat m ussels c ollected i n s ummer

h ave s lightly d ifferent 61 3 C v alues f rom m ussels c ollected i n w inter .

4 6

T his

c hapter h as

s ummarised t he r easons

d ifferent c lasses o f l iving o rganisms .

i s

o f

i nterest

t o

1 3

C /

1 2

C v ariations

i n

T he f iner d etails o f t he w ays i n

w hich p lants a nd a nimals r espond t o c hanges f actors

f or

z oologists

i n a ll t hese e nvironmental

a nd

b otanists,

r ather

t han

a rchaeologists, a nd n o a ttempt w ill b e m ade t o p erform s ophisticated e xperiments h ere .

T he a pproach w ill b e s pecifically a rchaeological,

a imed m erely a t a ssessing t he r ange o f 61 3 C v alues o f t he i ndigenous f oods o f t he r esearch a rea .

T his l isting o f t he p ossible s ources o f

i sotopic v ariation i s m eant a s a f ramework w ithin w hich t o d esign a s ampling s trategy , r ather t han a s as eries o f s ubjects f or e xhaustive r esearch .

C hapter 5 h as e xamined t he f actors l ikely t o p roduce 61 3 C

v ariations i n t hree m ajor c ategories o f h uman f oods .

C hapter 6 w ill g o

o n t o d evelop a s ampling s cheme d esigned t o a scertain t he r ange o f c arbon i sotope v alues i n t he e dible p lants a nd a nimals o f e ach o f f our r esource z ones

i n t he w estern C ape .

4 7

C HAPTER 6

F IELD S AMPLING A ND L ABORATORY A NALYSIS O F T HE P REHISTORIC H UMAN F OOD-WEB

I n o rder t o c haracterise t he f ood-web o f t he i sotopic

t erms,

u ndertaken ,

a large-scale

s ample

s outh-western

c ollection

C ape

i n

p rogramme w as

i ncluding b oth t errestrial a nd m arine f lora a nd f auna .

T he

s pecimens w ere t hen a nalysed f or t heir s table c arbon i sotope r atios. T he

c ollecting p rogramme

s tretched o ver s ome t wo y ears ,

w hich

m ade

p ossible t o p rovide a s et o f v alues f or a f ull a nnual c ycle, a ppropriate . f rom

A lthough t he s pecimens a re a ll m odern ,

s ampling

s tations

i n

r elatively

l aboratory r esults b ecame available,

i t

w here

t hey w ere c ollected

u ndisturbed

a reas.

A s

t he

t he c ollecting p rogramme w as

a ltered t o f ill i n t he o verall p icture o r t o e xplicate p uzzling r esults . I n t his , p art,

t he p roximity o f l aboratory a nd f ieldwork p layed a n

j oining

c hapter ,

t he

f ield

a nd

t wo

c omponents

l aboratory

i nto a c oherent

p rocedures

a re

i mportant

w hole .

t herefore

I n

t his

d escribed

t ogether w ith t he i sotopic r esults .

1 . T HE F IELD S AMPLING P ROGRAMME

1 . 1

P lants

A f airly

i ntensive

p lant

b eginning o f t his p roject , e ventually a nalysed .

s ampling

p rogramme

w as

i nitiated

a lthough n ot a ll t he s amples

a t

t he

c ollected w ere

P lants w ere o btained :

( 1 ) o n f ield t rips t o s elected f ield s tations , ( 2 ) f rom a c onvenient a nd w ell-established c ultivated c olony o f W atsonia ( for t he s easonal

s tudy ),

( 3 ) o n e xcursions w ith e thnographic i nformants , ( 4 ) b y d onation f rom o ther f ieldworkers ,

1 . 1 .1

F ield

o r

f armers ,

e tc .

S tations

R egular p lant c ollections w ere m ade a t E lands B ay ( in t he s trandveld v egetation t ype ), a t D e H angen i n t he N ardouwsberg a nd a t B oontjieskloof ( in a rid f ynbos) .

S ince m any p lant f oods a re o nly s easonally a vailable ,

4 F 1

r egular c ollecting t rips a re i mperative i f o ne i s t o g ain a c omplete p icture o f t he r esources w ere m ade

o ffered b y

i n N ovember 1 981

I n a ddition ,

a ny p articular a rea .

a nd F ebruary ,

M ay ,

t o D oornbosch i n t he K aroo ( May ,

A ndriesgrond

S everal t rips w ere a lso m ade

J uly a nd S eptember 1 982 ) a nd o ne o r t wo

e ach t o P iketberg m ountain

K aroo ) ,

J uly a nd S eptember 1 982 .

t wo-monthly m onitoring o f a n a rea o f m ountainside a t K alk

B ay o n t he C ape P eninsula w as c arried o ut .

t rips

C ollections

( mountain

( in t he O lifants

f ynbos ),

R iver

t he c oastal f ynbos/renosterveld b oundary )

A spoort

V alley )

( succulent

a nd H opefield

( on

( see M aps I II a nd V ).

T hese s ampling s tations w ere c hosen b ecause t hey w ere r elatively l arge t racts o f u ncultivated l and .

a ll

s ituated

i n

T hey w ere a lso p icked t o

y ield a r easonably r epresentative c ross-section o f t he v egetation t ypes o f

the

s outh-western

A ndriesgrond,

C ape.

S everal

B oontjieskloof ,

A spoort)

( Elands are

Bay,

also

D e

n ear

Hangen,

e xcavated

a rchaeological s ites .

1 . 1 .2 C ultivated W atsonia c olony

O riginally ,

s easonal

s tudies o f t he

ó' 3 C v alues w ithin a s ingle p lant

s pecies w ere p lanned a t s ome o f t he f ield s tations m entioned a bove . T his

r apidly b ecame

i mpractical,

s ince m ost o f t he f ield s tations w ere

s ome d istance f rom t he l aboratory .

M any m ore s amples,

c ollected a t

s hort i ntervals, w ere n eeded f or a s easonal s tudy t han f or a ny o f t he o ther r esearch o bjectives,

s o a l arge c olony o f c ultivated W atsonia

p yramidata o n t he P eninsula w as c hosen f or t his p urpose . h ad

t he

a dvantage

o f

b eing

a ccessible

a nd

a lso

o f

T his c olony c ontaining

a

s ufficiently l arge n umber o f p lants t o a llow a t otal o f m ore t han 6 0 i ndividuals ( five e ach m onth f or a y ear ) t o b e r emoved .

W ild W atsonias

u sually g row i n s mall c lumps, s o t hat i t i s v ery d ifficult t o f ind 6 0 w ild p lants g rowing i n e xactly t he c olony h ad b een u ndisturbed ,

s ame m icroclimatic

e xcept b y w eeding ,

1 . 1 .3 E xcursions w ith E thnographic

I nformants

M ost

i n

o f

t he

e thnographic

p lants

c ollected

i nformants .

t he

c onditions .

T he

f or a t l east f ive y ears .

K aroo

w ere

p ointed

o ut

b y

T he e dible p lants o f t he K aroo h ave n ot b een

a s w ell d ocumented a s t hose o f t he l ess a rid a reas,

4 9

a nd s ome o f t hem

( e .g .

F ockea

c omaru )

a re

v ery d ifficult f or a n

i nexperienced p erson t o

f ind b ecause m ost o f t he p lant i s b elow t he g round . t rips a t D oornbosch w ith a n e thnographic

i nformant y ielded a n umber o f

p lants t hat w ould n ot o therwise h ave b een o btained , r ecords o f e dible

s pecies ,

T wo c ollecting

a s w ell a s t wo n ew

a nd t wo n ew m edicinal u ses

( Metelerkamp a nd

S ealy 1 983).

1 .1.4 D onations

S everal p lants w ere d onated b y i nterested a rchaeologists a nd b y f armers . F iona A rcher ,

w ho i s d oing e thnobotanical w ork i n N amaqualand ,

p rovided

an umber o f s amples i ncluding t he H ydnora a fricana a nalysed .

1 .2 A nimals

M ost o f t he a nimals i n T ables V III,

X a nd X I w ere e ither f ound d ead

d uring p lant c ollecting t rips , o r d onated b y f armers . t ortoises, k ills .

b at-eared f oxes a nd e ven a s teenbok w ere t aken f rom r oad-

M any o f t he s pecimens f or w hich o nly c ollagen v alues a re g iven

d ied n atural d eaths , b ush .

S amples o f h ares ,

a nd t he b ones w ere f ound w hile w alking t hrough t he

A n umber o f f armers d onated p arts o f a nimals t hey h ad t rapped o r

s hot, a nd a f ew s amples c ame f rom n ature r eserves a t C hurchaven o r a t C ape P oint

1 .3

( the l atter m ainly v ia t he S outh A frican M useum ).

M arine S pecimens

M arine s pecimens ,

t oo ,

w ere o btained i n a v ariety o f w ays:

s ome a lgae w ere c ollected c ollected b y s cuba d ivers .

i n

i ntertidal

r ock p ools ,

s hellfish a nd

w hile o thers w ere

A n i nitial s et o f s amples w as d onated b y t he

Z oology D epartment a t t he U niversity o f C ape T own .

M any m arine b irds ,

o ne o f t he s eals a nd t he w hale b one w ere f ound w ashed u p o n t he b each . T he a lbatross a nd p enguin w ere d onations f rom t he P ercy F itzpatrick I nstitute

f or

r espectively , F isheries .

A frican

O rnithology

a nd

t he

S outh

A frican

M useum

w hile t he o ther s eal w as d onated b y t he D epartment o f S ea

F ish w ere p urchased f rom f ishermen .

5 0

2 . L ABORATORY P ROCEDURE

A ll a nalyses w ere d one i n t he A rchaeometry L aboratory a t t he U niversity o f C ape T own .

T hree s teps a re i nvolved: s ample p reparation ,

d ioxide p roduction a nd p urification , t he m ass

a nd c arbon

c arbon

i sotope m easurement o n

s pectrometer .

2 . 1 S ample P reparation

S ample p reparation t echniques b e a nalysed .

M eat, p lants, m arine a lgae a nd s hellfish r equire v ery

l ittle p re-treatment . c ontaminants , m ore

c areful

v ary a ccording t o t he t ype o f m aterial t o

T hese s amples w ere m erely c leaned o f a ny s urface

f reeze-dried a nd g round t o a p owder . p reparation :

o n b one c ollagen , d ecalcifying

B one s amples r equire

a ll t he m easurements

i n t his s tudy w ere d one

t he o rganic f raction o f b one .

T his w as e xtracted b y

w hole

r equired f or c omplete

b one

i n

1 % t o

5 % h ydrochloric

d ecalcification

v aries

a cid .

d epending o n

T he

t ime

s ample

s ize

a nd o n w hether t he s ample c onsists m ainly o f c ompact o r o f c ancellous b one .

I t r anges f rom a f ew d ays t o m ore t han a m onth .

T hereafter , t he

c ollagen w as s oaked i n d istilled w ater w hich w as c hanged r egularly u ntil a ll t races o f a cid w ere r emoved . a nalysis.

B one c ollagen,

I t w as t hen f reeze-dried a nd r eady f or

r ather t han w hole b one,

i s t he p referred

m aterial f or 613 C r eadings o n a rchaeological s amples s ince t he s tronglyb ound p rotein c hains t hat m ake u p c ollagen m aintain t heir

i sotopic

i ntegrity w hen t he b one i s b uried ( Tamers a nd P earson 1 965 ; S ellstedt e t a l .

1 966 ;

T aylor a nd S lota 1 979 ;

I n a ddition ,

T aylor 1 980).

t he s low t urnover t ime o f c ollagen c arbon

a dult a nimals

( more t han t en y ears

i n h umans

i n t he b ones o f

( Stenhouse a nd B axter

1 979 )) m eans t hat t he r eadings o btained r epresent l ong-term a veraging o f t he d iet.

2 . 2 G as P roduction a nd P urification

S ample c ombustion w as c arried o ut i n s ealed g lass t ubes a s d escribed b y S ofer ( 1980).

Af ive-milligram s ample, e xcess c opper o xide ( at l east

h alf a g ram) a nd a t wist o f s ilver w ire w ere l oaded i nto g lass t ubes w hich w ere t hen e vacuated t o l ess t han 1 0 -2

5 1

T orr ,

s ealed o ff w ith a

g lassblower 's t orch a nd c ombusted

i n af urnace o vernight .

I nitially ,

P yrex t ubes a nd a c ombustion t emperature o f 5 00°C w ere u sed, b ut t his w as

f ound

p lants ),

t o

g ive

c ombustion o n

s ome

s amples

( particularly

s o q uartz t ubes a nd a t emperature o f 8 00°C w ere s ubstituted .

A fter c ombustion, d ioxide, m etal.

i ncomplete

t he c arbon

i n t he s ample i s i n t he f orm o f c arbon

w hile s ome o f t he c opper o xide h as

b een

r educed t o

c opper

A ny s ulphur o r h alides p resent s hould h ave r eacted w ith t he

s ilver w ire , w hich a lso a cts a s ac atalyst .

S ome s ulphur d ioxide m ay

a lso c ombine w ith c opper o xide t o f orm c opper s ulphate ( Sofer 1 980). T he t ube a lso c ontains w ater d own s lowly ,

v apour a nd

i f t he f urnace h as b een

c ooled

a ll t he n itrogen s hould b e p resent a s n itrogen g as,

N2 .

S low c ooling a llows r ecombination o f t he m ixture o f o xides o f n itrogen t hat

m ay h ave

f ormed

i nitially .

m etal w hile t he n itrogen p roduces

T he o xygen r e-oxidises N2

r educed c opper

( Sofer 1 980).

T he t ube i s t hen r emoved f rom t he f urnace , t he e nd s cored t o e nsure e asy b reakage ,

a nd

i nserted

D esmarais a nd H ayes

i nto

( 1976 ).

a " cracker"

o f

t he

t ype

d escribed

T he c racker i s a ttached t o a s tainless

F igure I : Gas s eparat ion l ine i n t he A rchaeornetry L aboratory ,

U .C .T .

5 2

b y

s teel

g as-separation

l ine

a nd

t he w hole

l ine

e vacuated t o

< 10 4

T orr .

D uring t his p rocedure t he l ower e nd o f t he s ample t ube i s i mmersed i n l iquid n itrogen , i t

c ontains.

w hich f reezes a ll t he c arbon d ioxide a nd w ater v apour

N itrogen

r emains

g aseous

s o

t hat

w hen

t he

l ine

i s

e ventually r eady f or u se, t he t ube i s c racked a nd t he n itrogen p umped a way ( after m easurement o n a m anometer ), w ater

l eaving t he c arbon d ioxide a nd

v apour b ehind .

T hese t wo s ubstances a re

s eparated b y p assing t he m ixture t hrough c old

t raps i mmersed i n a n a lcohol/dry i ce s lurry , t hus f reezing o ut w ater v apour b ut n ot c arbon d ioxide .

W hen s ufficiently f ree o f w ater ,

c arbon d ioxide y ield i s m easured o n a m anometer a nd t he g as f rozen a c lean t ube w ith

l iquid n itrogen .

T he t ube

i s

t he i nto

s ealed o ff w ith a

g lassblower 's t orch a nd i s t hen r eady f or i nsertion i nto a nother c racker o n t he m ass s pectrometer .

2 .3 M easurement o n t he M ass S pectrometer

T he M icromass 6 02E m ass s pectrometer u sed f or d13 C m easurements i s a 9 0° s ector d ouble-collector i nstrument w ith a d ual i nlet s ystem .

A ll

m easurements

w ere d one a gainst a l aboratory r eference g as t hat h as

b een r elated t o t he C hicago P DB m arine l imestone s tandard b y c alibration a gainst

s ix

N BS 1 6 ,

1 7 ,

N ational 1 8 ,

1 9 ,

T ables

B ureau o f S tandards

2 0 a nd 2 1 .

i sotopic

r eference m aterials ,

R esults o f t he a nalyses a re t abulated

i n

T he r eproducibility o f t he m ethod o n h omogenised

s amples i s b etter t han 0 ,1 ° /

0 o

.

3 . R ESULTS

3 . 1 T errestrial A nimals

T here a re f our t ables o f t he r esults o f a nalyses o f t errestrial a nimals . T able V III p resents

( 0

a nimals

i n d ifferent

c ollected

3C

r eadings o n c ollagen f rom t he s ame r esource z ones.

T able

m easurements o n m eat a nd c ollagen f rom t he s ame i ndividuals . a l ist o f 613 C v alues f or m eat

s pecies o f

I X

c ontains

T able X i s

f rom a nimals e ating a C3based d iet a nd

5 3

T able X I

i s a c omparison o f t he m eat-collagen s pacings f rom h erbivores

a nd c arnivores c ollected f rom o ne a rea .

T he s pecies o f a nimals

i nitially s elected f or T able V III h ad t o f ulfil

t hree r equirements: they h ad t o b e i mportant i n t he d iets o f p rehistoric p eople i n t he a rea they h ad t o o ccur i n m ore t han o ne o f t he r esource z ones , they h ad t o b e w idespread t oday s o t hat

s amples

a nd

w ould b e

r eadily

a vailable.

T here a re f ive

c olumns

i n T able V III,

r esource z ones ( the c oastal p lain , h ave b een m entioned u p t o n ow .

a lthough o nly t hree t errestrial

t he F old M ountain B elt a nd t he K aroo ) T he f irst,

r espectively c orrespond t o t hese a reas .

s econd a nd f ourth c olumns T he " Karoo m argin"

m erely r efers t o t he e xtreme e dge o f t he K aroo v egetation ,

c olumn

a lmost o n t he

K aroo/Fold M ountain B elt b oundary .

T he r eason f or t he s eparation o f

t his

i s

a rea

f rom

t he

" Karoo

p roper"

o bvious

b e

l ooks

a t

d iscussed

t he

c ollected h ere .

t he s ection o n h ares.

T he " Bushmanland" c olumn i s m eant t o p rovide

o ther c olumns,

w ill

o ne

m easurement o n t he h are

a nimals f rom a n u nequivocally

T his

i f

u nder

e nvironment f or c omparison w ith t he

C4

a lthough t he n orthern

C ape

i s

a ctually o utside t he

r esearch a rea .

T he c riteria a bove l ed t o t he c hoosing o f s teenbok , h ares , t ortoises a nd d assies.

A ll o ccur

a rchaeological

s ites

i n c onsiderable n umbers ( see T able

s outh-western C ape t oday .

I ) ,

a nd a ll

i n t he f ood-waste f rom

a re

S teenbok a re k nown t o

i nclude

s ome g rass

i n

A dams 1 980),

s o

( Smithers

t heir 613

v alues w ere e xpected t o c hange a long a t ransect f rom t he

c oastal p lain t o t he K aroo . d uiker o r g rysbok , ( Smithers 1 966 ;

D orst a nd D andelot 1 970;

i n t he

t heir d iet C

1 966;

s till a bundant

S teenbok w ere c hosen i n p reference t o g rey

s ince t he l atter t wo a re v irtually p ure b rowsers

D orst a nd D andelot

1 970;

A dams 1 980) .

w ould n ot b e e xpected t o c hange f rom t he c oastal p lain t o t he

C3

b ushes/

C4

C3

T heir 61 3 C v alues

b ushes/C 3 g rass

g rass o f t he K aroo .

g rysbok a re r estricted t o h abitats w ithin t he f ynbos a nd

s o

a re

n ot

s teenbok d o 17,4°/ 0 0

f ound

i n t he K aroo

( Bigalke

i n t he K aroo .

I n a ddition,

v egetation t ype ,

1 978).

i ndeed c hange f rom 19,4 t o 21,4 °/ 0

o f t he

T he 0

v alues

f or

a t t he c oast t o

T his p resumably r eflects a n i ncrease o f a t

5 4

l east 2° /

o 0

i n t he 613 C v alue o f t he f ood b eing c onsumed .

A dams ( 1980)

o bserved t wo s teenbok f eeding i n t he S trandveld , a nd f ound t hat t he f requency o f g rass i n t heir d iets w as 9 ,5 a nd 4 ,5% r espectively; t he r est o f t he t ime t hey w ere b rowsing .

T he K aroo s teenbok w as p robably

e ating m ore t han 1 0% g rass, s ince a c hange o f 61 3 C v alue f rom 26,5 t o 12,5°/00

i n o nly 1 0% o f i ts d iet w ould p robably b e i nsufficient t o

i ncrease t he a verage 613 t he s pring ,

C

v alue o f t he w hole d iet b y 2° /

0 0

. A t l east i n

m ore g rass i s a vailable i n t he K aroo t han i n t he S trandveld

a nd s o am ixed f eeder w ould p robably e at m ore o f i t . G illian P uttick ( 1980), S trandveld c ommunity ,

J ohn G lyphis a nd

i n a s tudy o f R aphicerus f eeding h abits f ound t hat

i n a

t hese a nimals a ltered t heir d iet

s easonally s o a s t o m ake t he b est u se o f f resh n ew plant g rowth a nd t o a void t he s econdary c hemicals t hat a ccumulate i n m any p lants a t c ertain t imes o f y ear. u tilised , s hrubs,

I n s ummer ,

all

i n a utumn t here

i s m ore o r

l ess

i n t he g round l ayer o f t he v egetation a re

p robably b ecause o f t he g rowth o f w inter g rass .

f rom B ushmanland i s s lightly l ess e nriched i n K aroo i n t he r esearch a rea :

e qually

i s a s trong p reference f or b road-leaved

a nd i n w inter p lants

p referred ,

v egetation

1 3

T he s teenbok

Ct han t he o ne f rom t he

t his m ay b e d ue t o i ndividual v ariation ( see

b elow ) o r p erhaps i t w as b rowsing m ore a nd g razing l ess .

S everal s pecies o f h are o ccur i n t he s outh-western C ape , t he m ost c ommon b eing L epus c apensis ,

t he C ape h are .

L epus s axatilis ,

t he b ush o r s crub

h are , B unolagus m onticularig , t he B ushman h are a nd P ronolagus r upestrig , t he r ed r ock h are, a re also f ound . ( Smithers

1 966 ,

D orst a nd D andelot

f rom a v ariety o f

l ocalities .

f rom 18,0 a nd 20,0 °/

A ll f eed almost e ntirely o n g rass 1 970) .

T he

L epus c apensis w as c ollected

61 3 C v alues

f or

i ts

c ollagen

r ange

. o n t he c oastal p lain t o 15,1° /a . i n t he

n orthern C ape ( see T able V III).

T he c oastal

r eadings f all

m ore o r l ess

w ithin t he r ange o f v alues f or o ther a nimals i n t his z one , p erhaps l ying s lightly t o t he p ositive s ide .

A larger s ample w ould b e r equired t o

s how w hether t his i s m erely i ndividual v ariation , o r r eflects s ome r eal, s light d ifference . i nteresting : v egetation t he

T he s pecimen f rom t he K aroo m argin i s e xtremely

a lthough ( Acocks '

K aroo/fynbos

p redominantly

c3.

i t w as

c ollected

s ucculent K aroo ), b oundary,

w here

i n a n a rea o f d istinctly K aroo

i t w as o nly a f ew k ilometres f rom t he

g rass

s pecies

m ust

s till

b e

M eat ( but n ot c ollagen ) f rom t wo i ndividuals f rom t he

F old M ountain B elt w as a lso a nalysed .

T he r esults a re , a s e xpected,

t ypically

( Table

C3

X ).

I t

i s

i nteresting t hat t he h are

f rom t he

n orthern C ape, a n a rea t hat s upposedly h as a lmost 1 00% C4 g rasses, v ery i sotopically l ight c onsidering t hat g razer .

i s

i t i s s upposed t o b e l argely a

A nimals t hat e at n othing b ut C4 g rasses h ave 61 3 C v alues o f

a round 7°/, : , 0 .

C learly , t here a re e ither c onsiderable q uantities o f

g rasses i n t he n orthern C ape,

C3

o r e lse t his h are o btained s ignificant

a mounts o f i ts f ood b y b rowsing .

A t t he b eginning o f t he s ampling p rogramme , t oo ,

w ould y ield

d iet .

s ome e nvironmental

i t w as h oped t hat t ortoises ,

i nformation d ue t o g rass

T ortoises a re e asy t o c ollect ,

i n t heir

a nd t he c arapaces o f d ead a nimals

a re c ommon a nd r eadily i dentifiable ,

s o a l arge c ollection o f s pecimens

w as a massed .

p lain ,

m argin

a re

T hose f rom t he c oastal C hersina

a nqulata ,

P sammobates t entorius

a nd

v erroxii.

t he m ountains a nd t he K aroo

t hose

f rom

t he

n orthern

C ape

Very l ittle i nformation o n t ortoise

d iet i s available: i n f act t here i s n o r eliable p ublished s tudy .

D r

B ill B ranch o f t he P ort E lizabeth M useum i s m onitoring t he a ctivities o f a g roup

o f

C hersina

a nqulata

i n

t he

e astern

C ape,

a nd

h e

h as

a

c ollection o f a bout f ifty a s y et u nidentified p lants t hat h e h as w atched t ortoises

c onsume .

a re n o g rasses , p erhaps a s s mall e astern C ape

M ost a re s mall

a nnuals o r w eeds .

T here

a nd p erennial b ushes s eem t o b e b rowsed r arely ,

e xcept

s eedlings

h erbaceous

( Branch p ers .

c omm . ).

T he v egetation

i n t he

i s o f c ourse v ery d ifferent f rom t hat o f t he w estern C ape ,

b ut t his k ind o f p icture w ould c ertainly f it i n w ith t he T able V III, w hich s how p ure

C3

d iets.

8I 3 C v alues

i n

R emarks m ade b y h erpetologists

w orking i n t he r esearch a rea c onfirm t hat t ortoises a re n ot u sually s een e ating g rass

i n t he w ild ,

a nd a re r eluctant t o d o s o i n c aptivity ( John

G reig a nd A therton d e V illiers , t hat t ortoises S trandveld ,

p ers .

i n v egetation r ich

h ave

b een

s een

c omm . ) .

J ohn G reig a lso m entioned

i n s ucculent p lants,

e ating

t hese

a nd

t hat

a ttracted t o a p lant b y b rightly c oloured f lowers, c onsume . t he

t hey

o ften

w hich t hey t hen

w hich h ave

i nconspicuous f lowers .

v alues o f t ortoises i n T able V III a re r emarkably c onsistent , s how a n e ntirely C3 d iet . b e u sed a s e nvironmental

C3 -

a re

T his d oes n ot s eem t o b e a b ehaviour p attern l ikely t o l ead t o

c onsumption o f g rasses ,

i mportant

s uch a s t he

c omponent

T he 61 3 C

a nd c learly

A lthough t his m eans t hat t hese a nimals c annot i ndicators ,

i t

i s u seful

o f p rehistoric h uman d iet

w ould h ave c ontributed a

d erived i sotopic s ignature i n a ny e nvironment .

5 6

t o k now t hat s uch a n

D assies,

t oo,

w ere s uch a r egular i tem o f d iet f or H olocene h unter-

g atherers t hat i t w as d eemed e ssential t o m easure t heir 81 3 C v alues. T here i s s ome d oubt a s t o t he d etails o f t heir d iet: M ilton ( 1978) h as s uggested t hat P rocavia c apensis b rowsing a nd g razing a ctivity m aintains t he a reas b are o f v egetation o ften s een s urrounding r ocky o utcrops i n t he w estern C ape . a nnual

T his m eans t hat t hey w ould e at a m ixture o f b ushes,

w eeds a nd g rass .

S hortridge

( 1934) a nd S mithers

( 1966) b oth

c laim t hat P . c apensis e ats l eaves, g rass, t wigs, f ruit a nd b erries . D orst a nd D andelot

s ay t hat " they f eed m ostly o n g rass b ut a lso o n

b erries, s mall f ruits, b ark , l ichens a nd l eaves" ( Dorst a nd D andelot 1 970:153). V III

H owever , t he i sotope values f or d assie c ollagen i n T able

r eflect

a d iet

m ade

u p

m ore

o r

l ess

e ntirely

o f

b ushes

a nd

h erbaceous p lants w ith l ittle o r n o g rass .

T he s pecimen f rom t he K aroo y ields a value r ight i n t he m iddle o f t he C3derived r ange ,

w hereas a ll t he o ther a nimals f rom t his a rea r eflect a

C4 g rass c omponent i n t heir d iets. l ike t ortoises,

a re p ure b rowsers a nd w ill h ave C3 derived 61 3 C v alues

i n a ny e nvironment. f rom e lsewhere

I t t herefore s eems t hat d assies,

I t w ould b e i nteresting t o a nalyse s ome d assies

i n S outh A frica t o s ee w hether t his p attern i s c onfirmed .

A s t he a nimal s ampling p rogramme p rogressed , t he s hort l ist a bove . p lain ,

o ther s pecies w ere a dded t o

S pringbok w ere r eadily a vailable f rom t he c oastal

a nd f rom t he K aroo .

T hey a re r eally a n a rid e nvironment

n ot f ound o n t he c oastal p lain i n h istoric t imes,

a lthough t hey a re

r ecorded f rom t he O lifants R iver V alley ( Skead 1 980) . m ixed f eeders a nd

t heir

( Shortridge 1 934 ;

i sotope

r atios

S mithers 1 966 ;

a re

u ngulates t hat u sed t o o ccur

p robably

H owever ,

t hey a re

D orst a nd D andelot 1 970),

n ot

i n t he a rea,

s pecies

u nlike

t hose

o f

o ther

b ut n o l onger d o s o .

T he

s pecimen f rom t he K aroo i s 1 ,9 t o 2 ,6° / 0 0 m ore p ositive t han t hose f rom t he c oast . s till

A n a nimal

( -18 ,4

c ompared

r ange o f i ndividual c hange

t heir

f rom t he n orthern w ith

19 ,0 °/a

v ariation ,

p roportions

o f

b ut,

0

C ape

) .

T his

c omments

b rowsing

a nd

i n t he d iscussion o f v alues

r eports a v alue o f 16,5 °/, N amibia

( Table

s omewhat i s

m ore p ositive

c ertainly w ithin t he

l ike m any m ixed f eeders ,

r elative a vailability o f b ush a nd g rass ( see

i s

s pringbok

g razing a ccording t o t he

i n t heir

i mmediate e nvironment

f or s teenbok ).

V ogel

( 1978b )

f or a s pringbok f rom K eetmanshoop i n

I X ).

5 7

D une m ole r ats , b ut

e at

t oo ,

r oots,

a re m ixed f eeders .

s tems,

l eaves,

s tructure o f t heir g ut ( Jarvis,

p ers.

c oastal p lain

m argin ( -19 ,2°/0

0

b ulbs,

t ubers

a nd

g rasses

s eems a dapted t o a h igh-bulk ,

c omm .) . ( -22 ,1

T hey a re e ntirely h erbivorous ,

T he

0/ 0

813

0 , 20 ,3

v alues

C

°/ 0

o f

t he

0 a nd 17 ,8

)a re m ainly C3 derived .

a nd

t he

l ow-nutrient d iet

s pecimens

f rom

t he

/ 0 0 ) a nd f rom t he K aroo

0

T he l atter v alue c onfirms t he

s uggestion m ade i n t he d iscussion o f t he h are f rom t he K aroo m argin: n amely ,

t hat t here i s a g radual c hange-over f rom

M ountain B elt t o c hange

t han

v egetation

g rasses i n t he K aroo .

C4

t hat

b etween

t ype

v egetation

c oinciding

s andstone a nd s hale

w ith

g rasses i n t he F old

I t s eems a m uch m ore g radual t ypes,

t he

C3

a d istinct

g eological

c hange

b oundary

i n

b etween

i s v isible t o t he w est o f t he l ocality w here t hese

s amples w ere c ollected .

U nfortunately n o d une m ole

r ats

c ould b e

o btained f rom t he " Karoo p roper" a nd i t m ay b e t hat t hey d o n ot o ccur f urther i nto t he K aroo , p ers.

T he

( Jarvis

c omm .).

i sotope r atios

h ares ,

p referring t he s andy s oils o f t he c oast

o f h erbivores

s hown t o b e p artly g razers

( steenbok ,

s pringbok a nd d une m ole r ats ) a nd t hose s hown t o b e p ure b rowsers

( tortoises

a nd

d assies)

a re

almost

c ollected o n t he c oastal p lain .

i ndistinguishable

i n s pecimens

O ne c an i nfer f rom t his t hat e ven

g razing a nimals o n t he c oastal p lain e ither d o n ot e at

C4

a ll,

i sotopically

o r c onsume

i nsignificant. e nriched

i n

13

C

s uch

s mall

q uantities

T he o nly e xception ( -18

°

/

0

0

)

p erhaps e ven s ome s eaweed .

i s

t hat

t hey

a re

o ne h are t hat

i s

a nd m ay h ave b een e ating s ome C4

C4

g rasses a t

r elatively g rasses o r

g rasses d o o ccur i n t he c oastal plain,

a nd T hemeda t riandra a nd C ymbopogon s pp . a re r egarded a s t ypical o f c ertain p lant c ommunities

( Acocks 1 953 ),

a lthough t hey a re n ever c ommon .

Q uite p ossibly ,

h erbivores a void t hem w hen o ther s pecies a re a vailable

( see d iscussion

i n p revious

T he

l ast

t wo a nimals

c hapter ).

l isted

i n T able

s ources f or p rehistoric p eople .

B aboons

V III w ere n ot w ere

i ncluded

i mportant f ood f or a n

e ntirely

d ifferent r eason : t hey e at m any o f s ame i tems , p articularly p lants , t hat H olocene

h unter-gatherers

d id .

T heir

c onsisting o f m any s pecies o f r oots ,

d iet

c orms ,

i s

v egetable,

b erries a nd a lso t he s eeds ,

y oung s hoots a nd j uicy l ower s tems o f g rasses. c onsumed ,

l argely

S ome a nimal

f ood i s

e specially l ocusts, c aterpillars a nd t ermites ( Dorst a nd

5 8

D andelot

1 970;

D avidge

a lready d iscussed ,

1 976).

J ust a s f or t he h erbivorous a nimals

t he b oundary b etween a n e ntirely C3 based a nd a

p artly C4based d iet f or b aboons o ccurs a t t he F old M ountain B elt/Karoo I nterface.

B at-eared f oxes a re p erhaps

i n t he

r ecent

e xtreme

i mmigrants t o t he r esearch a rea

e ast).

T his

s pecies

i s

e xtending

( except

i ts

r ange

s outhwards a nd w estwards f rom t he K aroo i nto t he s outh-western C ape, w hich m ay b e a n i ndication o f t he i ncreasing a ridification o f t he f ynbos b iome ( Hendey 1 974,

1 983).

I t

i s p rincipally i nsectivorous,

e ating

t ermites a nd o ther l arge i nsects ,

b ut w ill a lso t ake e ggs a nd t he y oung

o f g round-nesting b irds ,

b erries a nd t uberous r oots

1 934 ;

S mithers

1 966 ;

l izards ,

D orst a nd D andelot

( Shortridge

1 970 ).

T able V III s hows t hat b at-eared f ox c ollagen 613 C r atios a re C3based o n t he c oastal c omponent

p lain a nd

i n

t he

i n t he F old

K aroo

a nd

t he

M ountain B elt,

n orthern

C ape .

b ut I n

i nclude a

s ome

c ases

C4

t he

r eadings a re c omparable t o t hose o f h erbivores f rom t he s ame z one ( at t he c oast a nd

i n t he m ountains),

b ut t he s pecimen f rom t he K aroo i s

i sotopically m uch h eavier t han t he h erbivorous a nimals

i n t hat

c olumn .

T his m ay r elate t o t he p roportion o f t he d iet t hat i s m ade u p o f a nimal f ood.

I t

i s

v ery c lear f rom T able V III t hat

i ndigenous a nimal f oods f rom t he

w estern C ape a re e ntirely C3 based i n t he c oastal p lain a nd t he F old M ountain

B elt.

T hese

t wo

z ones

c annot

b e

d istinguished

f rom

o ne

a nother , a nd t heir b rowsing a nd g razing a nimals h ave v irtually i dentical s table c arbon i sotope r atios .

T he

C3

z one e xtends a l ittle w ay i nto t he

s ucculent K aroo v egetation t ype,

b ut g razing a nimals s oon s tart t o

r eflect

t he

a

C4

g rass

c omponent

i n

l atter

e nvironment .

A nimals

c ollected n ear A spoort h ave 61 3 C v alues c omparable t o s pecimens f rom t he n orthern C ape , g rass

a n u nequivocally

( tortoises ,

d assies)

c4

g rass z one .

A nimals t hat d o n ot e at

s till h ave C3derived 61 3 C v alues ,

a nd t hese

a re w ithin t he r ange o f v ariation f or t he s ame s pecies f rom t he c oastal p lain a nd t he F old M ountain B elt. g rass ( steenbok, 2 - 3 °/ 0 0 .

H erbivorous a nimals t hat e at s ome

s pringbok a nd b aboons) s how a 61 3 C i ncrease o f a bout

T he m agnitude o f t his s hift i s a lmost l ow e nough t o b e

a ttributed t o i ndividual

variation ,

5 9

b ut b ecause

i t

i s a c onsistent

i ncrease

i n f our s eparate

s pecies w ith w idely d iffering d iets ,

o ne m ay

c onfidently a ccept t hat i t r epresents t he c rossing o f a r eal e cological b oundary.

T he n umbers a nd d istributions o f s amples i n T able V III a re o bviously f ar f rom

i deal .

q uestions

I n o rder t o p rovide a bsolutely w atertight a nswers

p osed

( probably

a t

e cological b ase .

i n t he p revious

l east

z one .

t hirty

c hapter ,

o ne w ould n eed

i ndividuals)

o f

e ach

t o t he

l arge n umbers

a nimal

f rom

e ach

T his w ould p rovide a s tatistically s ignificant d ata

U nfortunately ,

s uch a s ample w ould t ake m any y ears t o c ollect ,

o r

r equire t he w holesale s laughter o f t he f ew r emaining i ndigenous a nimals i n t he s outh-western C ape .

T he p icture t hat e merges f rom t he s amples w e

a lready h ave s eems s ufficiently c lear a nd c onsistent t o a llow u s t o d raw a t l east s ome c onclusions .

T wo o ther p ieces o f f irst

i nformation

i s t he a mount o f

c an b e e xtracted f rom T able V III .

813 C v ariation o ne c an e xpect a mongst d ifferent

i ndividuals o f t he s ame s pecies i n t he s ame e nvironment. f rom B oontjieskloof ,

T wo b aboons

t hree s pringbok f rom C hurchaven a nd t hree d assies

a nd t wo d une m ole r ats f rom E lands B ay w ere m easured .

T he r ange o f 61 3 C

v alues i n e ach s et o f r esults v aried f rom 00 /0 0 t o 2 ,5 b aboons w ith i dentical d rowned

T he

c arbon i sotope r atios w ere b oth

i n a f arm d am .

T hey w ere t he s ame a ge ,

°/ 0

0 .

T he t wo

j uveniles

f ound

h ad p robably b een

m embers o f t he s ame t roop , a nd a s s uch w ould b e q uite l ikely t o h ave h ad a lmost

i dentical d iets .

T he t hree d assies f rom E lands B ay a re a b etter

s ample : a ll w ere a dult b ut t hey w ere o f d ifferent a ges , d ifferent t imes. d iffered b y 2 ,2 ° / f or

a nimals

a nd c ollected a t

T he m ost p ositive a nd m ost n egative 61 3 C v alues 0 0

r aised

,w hich i s i n g ood a greement w ith t he f igure o btained i n

a l aboratory

o n

a c ontrolled

d iet.

T heir

i ndividual v ariation w as 2° /, 0 ( DeNiro a nd E pstein 1 978 ).

T he o ther q uestion t hat c an n ow b e a nswered r elates t o p ossible t rends i n

t he

i sotopic

c omposition

o f a nimals

w here

g radient ( within o ne v egetation t ype) ( e .g .

t here

i s

a c limatic

f rom t he r elatively h igh

r ainfall o f t he C ape P eninsula t o t he l ow r ainfall o f E lands B ay ). f igures i n T able V III d o r eveal s uch a t rend: w hich t he E lands

s ame

B ay ,

s pecies o f a nimal

t he

v alues

f rom

T he

i n t he t hree c ases

i n

c ould b e o btained f rom C ape P oint a nd

E lands

6 0

B aY a re

a lways

h eavier

( more

p ositive ) t han t hose f rom C ape P oint . f or t ortoises ,

T he d ifference i s 2 ,2 t o 2 ,8°/0 0

2° /0 0 f or s teenbok ,

a nd

1 ,8 t o 4 ,3 °/ 0 0 f or d une m ole

r ats.

T able I X p resents t he r esults o f a nalyses o f d ifferent t issues f rom t he s ame i ndividual f or t hree k inds o f a nimals: t ortoises, s pringbok a nd b at-eared f oxes .

T here a re e ight t ortoises , s ix o f w hich a re C hersina a ngulatg , w hile t wo a re P sammobates t entorius 1 3

C

v erroxii .

M eat a nd c ollagen s how

C3derived

v alues , a nd i n C hersina a ngulatg , t he c ollagen i s 2 ,2+0 ,6°/0

+ o ne s tandard d eviation ) m ore p ositive t han t he m eat .

( mean

T he m ean

v alue

f or t he c ollagen i s 22 ,4+1,4 ° / 00 . R eferring b ack t o T able V III,

t his

s eems s omewhat l ess n egative t han t he c ollagen r eadings f or m ost a nimals w ith a

C3

d iet.

i n t ortoises ,

P erhaps t his i s d ue t o s ome s light d ietary p reference

o r m aybe t he d iet-collagen s pacing i s d ifferent f or c old-

b looded t ortoises a nd w arm-blooded m ammals.

( There i s, o f c ourse, a

c hance t hat t he d ifference - i f i t i s s ignificant - i s c aused b y t he m ammals

c onsuming

a v ery

s mall

p roportion o f

g rass .)

C4

T he

t wo

P sammobates t entorius v erroxii y ielded e xtremely i nteresting r esults: t he v alues f or U CT 7 81 a re n o d ifferent f rom t hose f or C . a nqulata , O CT 7 82 h as m eat m ore p ositive t han i ts c ollagen . t ime ,

t o t he a uthor 's k nowledge ,

b ut

T his i s t he f irst

t hat t his p henomenon h as b een r eported .

T he a nalyses w ere r epeated t o c heck t heir a ccuracy , y ielding e ssentially t he s ame r esults ( the s mall d ifferences a re d ue m erely t o t he r epeat a nalyses h aving b een d one o n d ifferent s amples t he a nimal). e xpected

13

T here C/

12

C

f rom d ifferent

i s n o o bvious r eason f or t his

r atios,

a lmost

n othing

i s

m etabolism a nd p rocesses o f c ollagen s ynthesis ,

i nversion o f t he

k nown b ut

p arts o f

a bout

t ortoise

t here m ay b e

s ome

b iochemical d ifference b etween r eptiles a nd m ammals t hat w ould e xplain i t .

I t i s c ertainly a t opic f or f uture i nvestigation .

F ive s pringbok w ere a nalysed f rom

w ork r eported b y V ogel

s pacing i s 2 ,6+0,9°/

0 0

i n t his

s tudy ,

( 1978b).

a nd t hree

v alues o btained

T heir a verage m eat-collagen

,a nd t hat f or f ive b at-eared f oxes 2 ,2+0 ,7 ° /

0 0

.

W hen o ne c ompares t hese w ith t he a verage f or C . a ngulata ( 2 ,21 -0,6°/ 0 0 ), a ll t hree f igures a re r emarkably s imilar ,

c onsidering t he d iversity o f

t he a nimals ( a r eptile , a n u ngulate a nd a c arnivore).

6 1

T he v alues a re

t oo

s cattered

s pacings

i n

a dditional

t o

e nable

C3 based

a s

a nalyses

p hysiologists,

t he

d etection o f d ifferent

o pposed

m ight

t o

r eveal

C4based

f ood-webs .

i nformation

o f

t hese d ata s eem a dequate f or t he

a rchaeological d iets.

m eat-collagen

u se

A lthough t o

a nimal

r econstruction o f

T hey e nable e stimation o f t he c arbon i sotope

r atios o f t he m eat t hat w as a ctually e aten f rom a nalyses o f t he b ones r ecovered f rom a rchaeological s ites .

T able X I

c ontains t he 613

C

v alues o f m eat a nd c ollagen f rom t hree

h erbivores a nd t wo c arnivores c ollected f rom o ne s mall a rea .

T he m eat-

c ollagen s pacings a re n ot s ignificantly d ifferent i n t he t wo g roups ( 2 ,6 , 2 ,5 a nd 2 ,8°/ a re e ating

C3

v s 2 ,6 a nd 1 ,9°/

0 0

0 0

) .

A ssuming t hat t he h erbivores

p lants w hose a verage 61 3 C v alue i s 26 ,5°/o 0 , t heir d iet-

c ollagen s pacing i s + 5,9 °/ 0 0 . h erbivores h ave a d iet-collagen

C arnivores e ating t he m eat o f t hese s pacing o f + 4 ,7°/

0

0 .

T here h as b een

s ome s peculation ( van d er M erwe 1 982) t hat s pacings b etween d ifferent t issues o f a n a nimal,

a nd q uite p ossibly d iet-tissue s pacings a s w ell,

m ay b e s maller w hen t he d iet c ontains a l arge p roportion o f a nimal p roducts .

T his m ay r elate t o t he p ossible i ncorporation o f " blocks" o f

r eady-made p roteins ,

c arbohydrates a nd f ats ,

b odies o f c arnivores .

o r p arts t hereof ,

i nto t he

H erbivores, p erhaps, h ave t o b reak d own t hese

c ompounds f rom p lants i nto t heir c onstituent b uilding-blocks b efore r ecombining t hem i nto f orms m ore a ppropriate f or a nimals .

T his w ould b e

l ikely t o i nvolve a g reater d egree o f f ractionation t han o ccurs c arnivores.

I f t his i s t rue t hen o ne w ould e xpect t he m eat-collagen

s pacing i n c arnivores t o b e s maller t han i n h erbivores , v ariable t hat

i n

b ut i t s eems s o

l arge n umbers o f s amples w ould b e r equired t o t est t his

h ypothesis.

T he d iet-collagen s pacing o f + 5 ,9°/ I t

i s

s imilar t o V ogel 's

( 1978b )

0 0

f or h erbivores s eems r easonable .

v alue o f + 6 ,1 ° /

0 0

f or S outh A frican

u ngulates , a lthough r e-calculation o f t his f igure f rom t he d ata a ctually p resented i n h is p aper y ields a v alue o f 5 ,3 ° /

0 0

. T his i s c loser t o t he

f igure f or h uman p opulations i n w oodland N orth A merica g iven b y v an d er M erwe

a nd

V ogel

( 1978),

i .e .

+ 5,1 °/

0

.

C hisholm

c alculated t hat t his v alue s hould b e + 6 ,14 -0 ,4°/ h uman p opulation o f t he O ttawa v alley . t he d iet,

0 0

a l.

( 1982)

f or t he p rehistoric

D ifferences i n t he c omponents o f

t he k ind o f a nimal b eing s tudied o r e rrors

6 2

e t

i n t he e stimation

o f

t he

813

v ariation . s mall

v alue

C

o f

t he

d iet

p robably

a ccount

f or

s ome

o f

t his

W e k now , f or i nstance, t hat t he d iet-collagen s pacing i n

a nimals

w ith a h igh m etabolic

d ifferent f rom

t hat

f or

r ate

l arger a nimals

o btained v alues o f + 2,8 a nd + 3,9 °/

a nd a s hort

l ifespan

( DeNiro a nd E pstein

, f or m ice).

I t i s,

i s

( 1978)

t herefore,

p robably u nrealistic t o g roup s mall h erbivores ( e .g . h ares) t ogether w ith

l arge a ntelope .

T able X l ists t he 813 C3

v egetation z ones

K aroo m argin ) .

C

v alues o f f ifteen m eat s amples f rom a nimals f rom

( the c oastal p lain ,

t he F old M ountain B elt a nd t he

T hey r ange f rom 20 ,9°/, 0 f or a f rancolin ( a s eed-eater )

t o 25 ,6°/0 0 f or a t ortoise ,

w ith a m ean o f 23 ,6+1,3 ° /, 0 .

p robably a r easonable e stimate f or t he s table t he t errestrial a nimal c omponent s outh-western u ngulates

C ape .

( It

i s

c arbon

i sotope

T his

i s

r atio

o f

i n t he d iet o f H olocene p eople

t he

s ame

a s

t he

i n E ast a nd S outh A frica ( Lee T horp ,

a verage

f or

i n t he

b rowsing

u npublished d ata ,

V ogel

1 978b).)

T ables V III-XI t hus p rovide a nswers t o m ost o f t he q uestions a bout 61 3 C v alues o f t errestrial a nimals t hat w ere p osed i n C hapter 5 . t hat a nimals o n t he c oastal p lain ,

W e n ow k now

i n t he F old M ountain B elt a nd e ven o n

t he e dge o f t he K aroo h ave C3 based d iets, a nd t hat t he average 81 3 C v alue o f t heir m eat

i s

l ikely t o b e a bout 23,6 °/

0

.

I ndividual

v ariation b etween a nimals o f t he s ame s pecies r aised o n t he s ame d iet c an b e u p t o 2 ,5°/,

0

a nd c limatic g radients w ithin t he w estern h alf o f

t he r esearch a rea m ay a ffect t he i sotope r atios b y a s imilar a mount . T here d oes n ot,

h owever ,

s eem t o b e a ny d ifference b etween s pecimens

f rom V ogel's o riginal 5 -25% a nd s pecimens

C4

g rass a rea ( e .g . t hose f rom H opefield)

f rom t he 0 -5% a rea .

A p rehistoric h unter-gatherer w ould

h ave h ad t o g o s ome d istance i nto t he K aroo t o f ind a nimals t hat w ere i sotopically h eavier f rom e ating b e l ikely t o b e o nly 2 -3 ° /, S ome

i mportant

i tems

0

C4

g rasses ,

t hey w ould

m ore p ositive t han t hose f rom t he c oast .

i n t he p rehistoric m enu

n ot s how a ny s hift a t a ll,

a nd e ven t hen ,

( tortoises ,

d assies )

w ould

s ince t heir d iets d o n ot i nclude g rasses.

T he m eat-collagen s pacings o f t hree k inds o f a nimal m easured ( tortoises , s pringbok a nd b at-eared f oxes) a veraged o ut f airly c onsistently a t j ust o ver 2° / s ee

0 0

,a lthough i ndividual v ariation w as t oo g reat t o e nable o ne t o

i f t he v alues w ere d ifferent

6 3

i n

C3

a nd

C4

e nvironments,

o r

f or

c arnivores

c ompared w ith h erbivores .

D iet-collagen

s pacings

s eem

s lighly l ower f or c arnivores t han f or h erbivores .

3 .2 T errestrial P lants

613

v alues

C

f or e dible p arts o f p lants

p resented i n T able X II . t wo

s pecies

f rom t he r esearch a rea a re

T he m ajority a re c learly

o f s ucculent p lants

f rom t he K aroo

t he e xceptions a re

C3 ;

( Caralluma m ammillaris

a nd H oodia s p . ) a nd H ydnora a fricana f rom N amaqualand . p arasite ,

u sually o n t he r oots o f E uphorbia s pp .

T he l atter

T he s pecimen a nalysed

w as p arasitic o n t he r oots o f E uphorbia g ummifera i n ah ot, E uphorbia

i s

l argely a C AN g enus

i s a

( Mooney e t al.

d ry a rea .

1 977) a nd s o c ould

e asily s upply i sotopically h eavy p hotosynthetic p roducts t o a p arasite i n t his k ind o f e nvironment. w ere e ver

i mportant

a s

I t i s u nlikely t hat a ny o f t hese p lants

f ood ,

H oodia a nd

C aralluma b eing

f or t heir w ater c ontent t han t heir k ilojoule v alue 1 982).

N o

i nformation

i s

c onsumed

( Smith 1 966 ;

available o n t he n utritional

m ore

A rcher

q uality o f

H ydnora.

T he o ther p lants

l isted a re a ll

C3

s pecies .

T he s table c arbon

i sotope

r atios o f t heir e dible p arts a re s lighly d ifferent f rom t he s tandard f or C3 f oliage ( -26 ,5 °/ p ositive

t han

25 ,4 1 -1,8°/

0 0

0

) , b ut t he v ariation i s r andom .

26,5 °/

p H ,

, s ome

more n egative,

. E ven A ponogeton d istyacho

a n e ntirely u nremarkable k nown t o b e

0 0

C3 ,

i

v alue o f 26 ,9°/

S ome a re m ore

w ith

a m ean

of

af resh-water p lant , y ields

0 0

.

F resh-water p lants a re

o n t he w hole, b ut t hey a re a ffected b y f actors s uch a s

w ater s tate a nd c arbon d ioxide s ource ( Broecker a nd W alton 1 959;

O ana a nd D eevey 1 960).

T hey c an a lso b e v ery l ight d ue t o t he " lake

e ffect" w hich i s s imilar t o t he " canopy e ffect" .

V ogel h as r eported

s ome

G erman

v alues

a pproaching

50

0

/ 0

f or p lants

f rom

l akes

( Vogel

1 980).

3 .2 .1 V ariability i n t he 61 3 C V alues o f T errestrial P lants

F igure 2 c ontains m ore s pecific i nformation o n ( 1) b etween d ifferent t issues i n t he s ame p lant ,

v ariation i n 61 3 C

( 2 ) b etween t he s ame t issue

i n d ifferent i ndividuals i n ag roup o f p lants c ollected a t t he s ame t ime a nd p lace,

a nd ( 3) b etween s amples o f t he s ame s pecies c ollected a t

6 4

- 30 -

-29

-28

°

61 3c poB

Ne w

O ld c or m

c or m

27

O ld 26

L eaf

•

c or m

l ib

•

• Ne w c or m 25

• °

F lo wer

-24

•

-23

2 2

S EP

OCT

NOV

O ld c or ms o f

D EC

JAN

F EB

MAR

APR

M AY

JU N

JUL

AUG

S EP

OCT

1 982

•

Ne w c or ms o f 1 982

•

Ne w c or ms o f 1 983

b eca me o ld cor ms o f 1 983

Leaf F lo wer

F igure 2 : S easona l c hanges i n 61 3C v a lues o f Watson ia p yra m idata

d ifferent t imes o f y ear . p yramidata ,

A ll

t hese m easurements

a l arge s pecies o f W atsonia w ith p ink f lowers .

i ndividuals w ere d ug u p f rom t he s ame f or a y ear .

w ere d one o n W atsonia f ive

l arge p atch o f p lants e ach m onth

I n a h omogenised s ample o f f ive

N ovember 1 982 ,

A bout

i ndividuals

c ollected

t he n ew c orms h ad a 61 3 C v alue o f 27 ,9°/. . ,

i n

w hile t he

f lowers w ere 24 ,5°/0 0 ,a d ifference o f 3 ,4 °/. 0 b etween d ifferent t issues o f t he s ame p lants . p lant t o p lant:

T hese s pacings c an d iffer c onsiderably f rom

t he d ifferences b etween t he n ew c orms a nd t he o ld c orms

o f s ix s pecimens c ollected i n J uly 1 983 v aried f rom 0 ,1 t o 1 ,4°/,

o

.

T he l atter s ample a nd t hat c ollected i n M ay 1 983 g ive a n i dea o f t he i ndividual v ariation t o b e e xpected i n t he s ame t issues i n d ifferent p lants .

T he

dI 3 C v alues o f n ew c orms o f f ive p lants

d iffered f rom o ne a nother b y u p t o 1 °/

a

. .

f or s ix n ew c orms c ollected i n J uly 1 983 , c orresponding o ld c orms .

6 5

c ollected

i n

M ay

T his f igure r ose t o 1 ,8°/a 0 w hile i t w as 1 ,3 ° /.

0

f or t he

F igure 2 a lso s hows t he c yclical s easonal

v ariation t hat o ccurs

613 C v alues o f W atsonia p yramidata c orms.

i n t he

T he m ost p ositive a nd m ost

n egative v alues f or h omogenised s amples o f f ive n ew c orms w ere 23 ,2 a nd 29 ,2°/0

0

:a d ifference o f 6 °/0 0 .

w ave w ith t he m ost p ositive F ebruary ),

T he c urve a pproximates a s inusoidal

v alues o ccurring

i n

s ummer

( December-

t he p eriod d uring w hich t he p lant i s d ormant a fter f lowering .

613 C v alues f or t he n ew c orms r emain m ore o r l ess c onstant d uring t hese m onths ,

b ecoming i ncreasingly n egative a s t he p lant i nitiates n ew g rowth

i n a utumn a nd t he s tarch s tored i n t he c orm i s u sed t o m ake n ew l eaves a nd f lowers .

T he

i sotope r atios a re a t t heir l ightest w hen t he p lants

a re f lowering ( in S eptember-October ). a s t he n ew c orm g rows , t he

c ycle

i s

T hey t hen r apidly b ecome h eavier ,

u ntil t he p eriod o f s ummer d ormancy ,

r epeated .

E ach

y ear 's

" new

c orms"

w ill

a fter w hich b ecome

t he

f ollowing y ear 's " old c orms" .

T he

c ycle

u ndergone

b y

p hotosynthetic p roducts

W atsonia i s a c omplex o ne; d irect

i mportance

i n

p lants

s uch a s

f ortunately o nly a s mall p art o f i t i s o f

t o a rchaeologists .

C orms

s ummer - a t t his s tage t hey a re w hite ,

a re

m ost p alatable

i n t he

f loury a nd o f m aximum s ize .

F rom

a utumn o nwards a stringent w aste-products a re d eposited i n t hem a nd t hey b ecome p inkish ,

r esinous a nd u npleasant t o e at .

T he n ew c orms t hat

s tart t o b e f ormed a re w hite, b ut t hey a re v ery s mall u ntil a fter t he f lowering s eason .

T hus t he b est t ime t o e at t hem i s i n s ummer ,

613 C v alues a re a t t heir m ost p ositive .

w hen t he

I t i s l ikely t hat o nly t he m ost

p ositive p art o f t he r ange o f i sotope v alues d escribed a bove w ill b e i mportant

i n

t he

human

f ood-web;

f or

t his

s ample,

p erhaps,

23to-25°/0 0 .

T he g raph i n F igure 2 i s o f t he s ame s hape a s t hose d escribed b y L owdon a nd

D yck

h owever ,

( 1974)

a nd

L eavitt

a nd

L ong

( 1982).

I t

i s

n ot

a s

n eat,

s ince t hese a uthors w ere m easuring s easonal d ifferences i n

t rees,

a nd s o c ould t ake m onthly s amples f rom t he s ame p lant .

f ound

d ifferences

o f

6 a nd

3 °/

0 i n

m aple

a nd

j uniper

T hey

l eaves

r espectively a t d ifferent t imes o f y ear .

F igure d 13

2 t hus

p rovides

a n

e stimate

o f t he

i ntra- a nd

i nter-individual

Cv ariation a nd a lso t he s easonal v ariation i n a t l east o ne i mportant

f ood p lant

i n t he s outh-western C ape .

6 6

N ot a ll o f t his

i s l ikely t o b e

r eflected i n t he h uman f ood-web , o nly a t c ertain t imes o f y ear .

s ince W atsonias w ere p robably e aten

SI 3 C v ariation d ue e ither t o i ndividual

d ifferences o r t o t he s eason o f s ampling w ould b e u nlikely t o e xceed 2 °/ 0

i n W atsonia p yramidata .

T his

w ell,

b ecause W atsonia h as m ore s trongly-differentiated s easonal g rowth

p hases t han m ost o ther p lants .

A s

i s p robably t rue f or o ther p lants a s

i n t he a nimal s ample ,

d ifferences o f

t his m agnitude w ould b e u nlikely t o h ave m uch e ffect o n t he average v alues

f or

p lant

C3

f oods

i n

T able X II.

T hey w ould p robably n ot

c onstitute a m ajor s ource o f e rror i n c alculations o f t he

i sotopic

s ignature o f t he h uman f ood-web i n t he s outh-western C ape .

3 .3 T he

T he

M arine F ood-web

f inal

s et

o f

s amples

o f

i ndigenous

f oods

c onsists

o f

m arine

o rganisms a vailable a t t he c oast .

3 .3 .1

P rimary P roducers

- A lgae

Ar ange o f a lgae w as c ollected , n ot b ecause t hey w ere e aten b y h umans ( although t hey m ight d etermining t he T heir

613

b ut

b ecause

t hey

a re

r ange

f rom 34 ,5 t o

9°/a 0 .

T he m ost

n egative s pecies t end t o b e d eep-water r ed w eeds, p ositive a re

i mportant

i n

i sotope r atios o f o rganisms h igher i n t he f ood-web .

v alues

C

h ave b een)

i ntertidal g reens

( see T able X IV).

i sotopically

w hereas

t he m ost

W hen t hese r esults a re

d rawn o ut i n ah istogram ( Figure 3 ), o ne c an s ee t hat t he r ed s eaweeds e ncompass a lmost t he w hole

r ange o f

t end t o c luster a t t he p ositive e nd . b etween 9 a nd 25 25to-30°/

o 0

0 /0 0

g reens a nd b rowns

i s a g reat d eal o f o verlap

.

v alue s eems a pparent .

C

T here

w hereas

,b ut o nly r ed w eeds e xhibit v alues a s n egative a s

O n f urther e xamination o f t he d ata , 613

v alues ,

s ome

c orrelation b etween d epth a nd

T here i s a c ertain a mount o f z onation o f

g reen, b rown a nd r ed w eeds o n t he s ub-littoral s hore: r ed l ight, w ith i ts l ong w avelength , w ater ,

i s

f iltered o ut b y o nly a f ew c entimetres o f s ea-

s o t hat s eaweeds w ith c omplementary p igmentation ( green ) c an o nly

s urvive i n t he i ntertidal o r s hallow e dge o f t he s ubtidal z one . g reen l ight p enetrates m uch f urther i nto t he w ater , c an

l ive

a t

f ar

g reater

d epths .

6 7

B rown

w eeds

Y ellow-

s o t hat r ed s eaweeds a re

m ost

c ommon

a t

i ntermediate d epths .

O n t he S outh A frican c oast, p hotosynthetically

s ignificant l ight i s l imited t o t he t op t hirty m etres o f s ea-water ,

a nd

t his d etermines t he b oundaries o f p ossible s eaweed c olonisation ( Simons 1 976).

O f t he s pecies

i n T able X IV ,

t he m ost

i sotopically n egative a re t hose

c ollected f rom t he d eepest w ater ( although t his w as n ot v ery d eep , 6 -8

m etres).

a gardhii,

T hese

T hamnophyllis

B otryocarpa p rolifera ,

b inderianum . w ater

i nclude

h ave

A ll a re r eds ,

d isciaera ,

o nly

K allymenia

H ymenena v enosa a nd N euroglossum

b ut s ome o ther r eds c ollected f rom s hallow

c omparatively

p ositive

c arbon

i sotope

r atios

o rbitosg. , G elidium Kistoides a nd P orphyra c apensi§) .

( Aeodes

T o t est w hether

d eep-water s eaweeds d o i ndeed h ave m ore n egative 6I 3 C v alues t han t hose f rom s hallow w ater ,

s imultaneous

c ollections o f t he s ame s pecies f rom

d eep ( 25-29 m etres) a nd s hallow ( a f ew m etres ) w ater w ere m ade a t V ulcan R ock o ff H out B ay .

T he r esults a re s hown i n T able XV .

d oes c ontribute t o v ery n egative c arbon i sotope r atios : d eeper

s amples

a re

m ore

n egative

t han

t he

D epth c learly i n a ll c ases t he

s hallow

o nes,

d ifference v aries f rom 1 ,5° / 0 0 ( Hymenena v enosa ) t o 1 1,3 °/0 c ornea ). t hese

0

b ut

t he

( Pachymenia

S everal o f t he s pecies w ith v ery l ight i sotopic r atios d isplay

v alues

i n

s hallow

a s

w ell

a s

i n

d eep

w ater :

N euroglossum

b inderianum , H ymenena v enosa a nd B otryocarpa p rolifera a ll h ave 6I 3 C v alues b elow 30 ° /

0 0

i n b oth s amples .

T he r esults f or E cklonia m axima

a nd L aminaria p allida a re i nteresting : p revious m easurements h ad y ielded C3 like values f or L aminaria ( -20,9°/

0 0

a t O udekraal, 19 ,0°/

P aternoster ) a nd C4like v alues f or E cklonia ( -12 ,0 14 ,1%o a t P aternoster ) .

0/ 00

0 0

a t

a t O udekraal a nd

T he n ew s amples g ive v alues o f 16 t o 17 ° /0

0

f or b oth s pecies i n s hallow w ater , a nd 22 t o 23 °/ 0 0 i n d eep w ater . E arlier s amples o f L aminaria w ere p robably f rom m uch d eeper w ater t han t hose o f E cklonia .

A lthough d epth i s a c ontributing f actor t o t he f inal 6I 3 C v alue o f a m arine a lga , o f

v alues

t here m ust a lso b e s ome o ther r easons f or t he l arge

b etween d ifferent

s pecies .

t hat s ome a lgae c ontain e nzymes o f d ehydrogenase ( Ting 1 976). p hotosynthesis

P hysiological C4

w ork h as

r ange

r evealed

a cid m etabolism s uch a s m alate

K arekar a nd J oshi ( 1973) a lso o bserved

i n c ertain m arine a lgae ,

s eems t o b e p roduced .

6 8

b ut

n oted t hat

C4

o nly a spartate

R ange o f v ar iat ion

w ith in

4

0

3

< > )

2

A

2

A

A

o ne s pec ies

A A

A

GREE NS




A

No . o f s pec ies

6

A

5

A

REDS

4

A

3

A A A

A

A

A

2

-25

20

15

-5

A A A A A A A

. A

A

1

A A

61 3 cpDB

23 5

F igure 3 : 61 3C v a lues o f s ea weeds f rom t he S outh -Wes tern Cape

( 1

> R ange

o f v ar iat ion w ith in o ne s pec ies

0 0 0 0 0 0 0, 0 > 0

3

0

GREENS

0

A A A A

A

A ( A A

BRO WNS

> < .A

A

No . o f s pec ies

A

4

2

1 . -5

RE DS

A

3

, AA 10

A A A A AA A A : 1 5 2 0

A

A 2

2 5

A

3 o

-2 3 5 6 13

F igure 4 : 6. 1 3 C

CPDB

v a lues o f s ea weeds f ro m p ub l icat ions

6 9

O ther a lgae c ontain r ibulose d iphosphate ( Steemann N ielsen 1 975), o r e xhibit

f eatures

c haracteristic o f C3 p hotosynthesis ,

d ioxide c ompensation p oint .

T his

s uch a s a c arbon

i s t he p oint a t w hich a n e quilibrium

i s r eached b etween t he r ate o f p hotosynthetic c arbon d ioxide f ixation a nd t he r ate o f c arbon d ioxide l oss f rom t he p lant i n l ight a nd d ark r espiration .

L aboratory v alues a re t ypically 3 0-80 p pm C O2 f or C3

p lants a nd l ow v alues n ear 0 p pm f or C4 p lants.

T olbert a nd G arey

( 1976) m easured t his f eature i n an umber o f m arine a lgae, s ome o f t hem t he s ame s pecies a s T ing u sed i n h is s tudy o f N ADH m alate d ehydrogenase a ctivity .

T he r esults , t ogether w ith t he 6I 3 C v alues g iven b y B lack a nd

B ender ( 1976) a re t abulated i n T able X VI.

S ome o f t he c arbon d ioxide

c ompensation p oints a pproach t hose o f C4 p lants E nteromorpha) p lants .

w hile o thers

( Avrainvillea e recta)

t he v alue f or

a re m ore

l ike C3

U nfortunately t here d oes n ot s eem t o b e a ny c lear c orrelation

b etween C O2 c ompensation p oint , b e

( e .g .

t hat

a ll

t he

e nzyme a ctivity a nd 6I 3 C v alue .

m easurements

w ould

h ave

t o

b e

d one

o n

I t m ay

t he

i ndividual p lants f or t hese r elationships t o b e p erceived .

s ame

F actors

u nimportant i n t errestrial p hotosynthesis h ave t o b e t aken i nto a ccount i n i nterpreting t his k ind o f d ata i n m arine a lgae :

b oth T ing ( 1976) a nd

H ough ( 1976) h ave s uggested t hat t here m ay b e e xtensive r ef i xation o f r espired c arbon d ioxide i n t he s ea b ecause t he d iffusional r esistance o f s eawater t o c arbon d ioxide i s 1 05 t i• mes t hat o f a ir .

I n

s pite o f t his

v ery u nsatisfactory s tate o f t heoretical

k nowlege o f

m arine a lgal p hotosynthesis , an umber o f 6I 3 C v alues h ave b een p ublished ( Wickman 1 952; C raig 1 953; P ark a nd E pstein 1 961; P arker 1 964; S ackett e t a l. 1 965; S mith a nd E pstein 1 971; B lack a nd B ender 1 976; D eNiro a nd E pstein 1 981; S chell 1 983).

T hese a re d istributed , o n t he w hole, i n a

v ery s imilar m anner t o t he s amples f rom t he w estern C ape ( see F igure 4 ) , e xcept t hat t he n umber o f v ery n egative v alues r eported i s f ar f ewer . M any m easurements s how c onsiderable

I 3

Ce nrichment,

l eading t o s ome

s peculation a s t o w hether b icarbonate a s w ell a s d issolved c arbon d ioxide c ould b e a cting a s a s ource o f c arbon ( 1980)

h as

p ointed

o ut

t hat

i t

i s

( Craig

u nnecessary

t o

1 954).

a ssume

V ogel t his

i n

e xplaining p ositive 61 3 C v alues : t he i ncreased d iffusional r esistance t o C O2

i n

w ater

B icarbonate

c ompared

m ay a ct

a s

w ith

a ir

a c arbon

7 0

c an

a ccount

r eservoir ,

f or b ut

a ny t he

d ifferences . f ractionation

a ssociated w ith i ts e quilibrium c onversion t o c arbon d ioxide e nsures t hat t he g as i s i sotopically l ight ( Deuser a nd D egens

1 967) .

T he a lgae r eported i n t he l iterature a re m ore o r l ess e venly d ivided b etween g reen ,

b rown a nd r ed s pecies ,

a nd a re p robably l argely s hallow

w ater p lants .

O nly o ne i s i sotopically l ighter t han 30

s ample o f H alymenia d urvillaei f rom C ape F lattery , b y B lack a nd B ender ( 1976). t hat R UDP c arboxylase i s

0 /0 0

:t his i s a

A ustralia ,

m easured

T hey r emarked t hat " although w e c onclude

i nvolved i n C O2 f ixation i n H alymenia i t a lso

a ppears t hat H alymenia h as

s ome f eatures o f C O2 a ssimilation w hich a re

n ot c lear , a nd w hich s hould b e i nvestigated f urther , s ince t his i s a n u nusual d egree o f

3 .3 .2

I 2

C e nrichment"

( Black a nd B ender 1 976 :28 ).

M arine F auna

A g reat d eal m ore w ork w ill b e n eeded b efore t he p atterning i n t he 6' 3 C v alues o f m arine a lgae i s w ell u nderstood . s tudy , h owever ,

F or t he p urposes o f t his

i t i s e nough t o k now t he c arbon i sotope r atios o f t he

d etritus f ormed b y m echanically a nd b iologically d egraded a lgae . d epends

n ot o nly o n t he o riginal

T his

6 ' 3C v alues o f t he d ifferent s pecies,

b ut a lso o n t heir r elative a bundances .

I n S outh A frican w aters m uch o f

t he b iomass i s c oncentrated i n k elp b eds:

l arge s tands o f L aminaria

p allida a nd/or E cklonia m axima a nd a ssociated f auna a nd f lora .

E cklonia

i s t he d ominant k elp i n c ommunities o ccurring c lose i nshore ,

w hile

L aminaria

1 981).

p redominates

i n

d eeper

w ater

( Branch

a nd

B ranch

D etritus f rom t hese s eaweeds , t ogether w ith p hytoplankton , p rovides f ood f or f ilter-feeders s uch a s m ussels , A s

o ne

p roceeds

u p t he

m arine

s ponges ,

s ea-cucumbers a nd r ed-bait .

f ood-chain,

t he

6 '

3C

o rganisms b ecome i ncreasingly p ositive ( see T able X VII) . a r ange o f t rophic l evels ,

f rom f ilter-feeders

( crayfish) w ere e aten b y p rehistoric p eople . b ased d iets

v alues

o f t he

O rganisms f rom

( mussels) t o s cavengers T his m eans t hat m arine-

i ncluded i tems w ith a r ange o f 6' 3 C v alues .

P robably ,

t hey

w ere u sually m ade u p o f am ixture o f t he a nimals c atalogued i n T able X III , w hose m ean v alue i s 15 ,64 -1,6°/

o

( excluding t he w hale ) .

C ollagen f rom a n u nidentified s pecies o f w hale h ad a c arbon i sotope r eading o f 20 ,2 ° /0 0 , m uch l ike t he v alues f or t errestrial a nimals w ith C3 based d iets i n T able V III. O ther m easurements o n w hales, h owever ,

7 1

h ave

b een

r ather

l ess

n egative

t herefore c loser t o t he a verage

( Schoeninger

a nd D eNiro

1 984)

a nd

i sotopic c omposition o f m arine f oods .

T his s eemed p uzzling a t f irst,

b ut t he c ollagen w as s ubseqently d e-

f atted a nd t hen f ound t o h ave a ó13 C v alue o f 12 ,5°/0 0 .

T he d ifference

b etween t hese t wo v alues m ust b e d ue t o t he p resence o f f ats ,

b ut t hese

w ould h ave h ad t o h ave b een v ery i sotopically d epleted i ndeed t o a ccount f or s uch a l arge s hift . c ollagen ,

I f t he f at w ere a bout 8 °/ 0

a s i s t he c ase f or u ngulates ( Vogel 1 978b ) ,

0

l ighter t han

t hen t he o riginal

s ample w ould h ave h ad t o b e a lmost e ntirely f at f or t here t o b e s uch a l arge d ifference .

T his w as n ot t he c ase .

s pacing b etween t he

A g reatly i ncreased i sotopic

v arious t issues o f w hales w ould b e c onsistent w ith

t he i dea t hat l arger f ractionation f actors f or b igger a nimals, u ngulates

e .g .

c ompared w ith m ice ( see d iscussion o n p age 3 8 ) a re a f unction

o f s lower m etabolic r ate a nd g reater l ongevity . e aten b y p rehistoric p eople i ncluded f at ,

I f t he p arts o f w hales

w hich s eems l ikely s ince t his

i s a n i tem i n s hort s upply i n m ost h unter-gatherer d iets, t hen w hales m ay h ave m ade t he a verage i sotopic s ignature o f m arine-based h uman d iets s omewhat m ore n egative .

D irect m easurements o f t he

i sotopic r atios o f

w hale m eat a nd b lubber a re o bviously n eeded t o r esolve t his q uestion . I t i s d ifficult t o e stimate t he c ontribution w hales m ade t o p rehistoric d iets: w hale s trandings a re r egular , i f i nfrequent o ccurrences a long t hese c oasts t oday .

T hey w ere p robably m uch m ore c ommon i n t he d ays

b efore c ommercial h unting d rastically r educed w hale p opulations.

A

s ingle a nimal w ould h ave p rovided a v ast a mount o f f ood, w hich c ould p erhaps h ave b een s tored b y d rying, o r b y b urying i n t he s and , w hich, w hen w et ,

i s a s aline a naerobic e nvironment .

O n a verage, t he 81 3 C v alue o f am ixed m arine-based d iet i n t he s outhw estern C ape w as p robably a bout 16,5 °/ f igure o f 17 ,84 -1,4 ° /,

0 .

T his i s s imilar t o t he

o btained b y C hisholm e t a l. ( 1982) f or t he w est

c oast o f C anada .

T able X III a lso l ists t he

13

C/

12

C

r atios o f b lack m ussels c ollected f rom

E lands B ay a t d ifferent t imes o f t he y ear . c lustered , n ot ,

A ll t he v alues a re t ightly

w ith n o d iscernable s easonal v ariation . S easonal f actors d o _ 13 t herefore , c ontribute t o t he s pread o f e C v alues o f p rehistoric

h uman d iets .

7 2

4 . E XTRAPOLATION B ACK I NTO T HE P AST

T he a nalyses a lready d iscussed p rovide a r easonably c omplete p icture o f t he

m odern

i sotopic

e cology

o f

t he

s outh-western

C ape .

T hese

s amples

w ere c arefully c hosen t o m inimise p ossible c ontamination f rom e xotic v egetation ( including c ultivated p lants) .

N evertheless , t he p ossibility

t hat r ecent e cological d isturbances h ave a ffected t he p icture g ained m ust

b e

c onsidered .

( overgrazing,

T he

k inds

o f

d isturbances

b ut p robably m uch l ess e ffect o n

t he i sotopic s tructure o f t he v egetation, d etermined b y c limate a nd ,

j ustified ,

H olocene

t here ( see

s ince t his i s p rincipally

t o a l esser e xtent,

m odern c ollections o f e dible

M oreover ,

d iscussed

i ncreased f ire f requency) h ave a t remendous i mpact o n

p lant e cology a nd c ommunity s tructure ,

s eemed

a lready

g eology .

i ndigenous p lants a nd a nimals t herefore

b ut

t his

a ssumption

o bviously

h ave

b een

s ome

s mall

c limatic

that

c ould

well

C hapter

3 )

T he u se of

d istribution o f t he d ifferent v egetation t ypes .

r equires

t esting .

c hanges d uring t he

have

i nfluenced

the

I n o rder t o e nsure t hat

t he r eadings o btained o n t he m odern e nvironment c an i ndeed b e u sed i n t he

i nterpretation o f t he 6I 3 C v alues o f p rehistoric h uman s keletons ,

a

s hort s eries o f a nimal b ones f rom a n a rchaeological e xcavation w ere a lso a nalysed.

T he s ite c hosen w as E lands B ay C ave ,

s ince t his h as t he l ongest s equence

o f d eposits o f a ny e xcavated s ite i n t he r esearch a rea .

O nly a nimals o f

w hich m odern s pecimens h ad a lready b een a nalysed w ere u sed , r esults u sed :

c ould

b e

c ompared .

h ares a nd s teenbok .

s imilar 613

C

T horp 1 983 ),

T wo

s pecies

o f g razers/mixed

s o t hat

t he

f eeders

w ere

B rowsing a nimals h ave b een s hown t o h ave

v alues i n v ery d ifferent e nvironments ( Table V III, L ee

a nd s o c annot b e u sed a s e nvironmental

i ndicators .

M arine

a nimals c an o nly b e e xpected t o s how c hanges i n t heir 6I 3 C v alues i f t he s ea t emperature c hanges .

A n i ncrease i n t he p roportion o f C4 g rasses o n

t he c oastal p lain i n p rehistoric t imes

i s t he e nvironmental

c hange m ost

l ikely t o h ave o ccurred , a nd t his s hould b e r eflected i n t he 6I 3 C v alues o f p rehistoric g razers o r m ixed f eeders . t he

o ne

m ost

l ikely t o

i nvalidate

c onclusions a bout p rehistoric d iets

b ased o n t he m odern d ichotomy b etween t he a nd c oastal t errestrial f oods .

7 3

S uch a c hange w ould a lso b e

i sotopic s ignatures o f m arine

A s eries o f p rehistoric h are a nd s teenbok b ones w ere c hosen f rom t he s pecimens i dentified b y R ichard K lein .

T he s ample o f d iagnostic f aunal

m aterial i s n ot l arge, s o o nly s even b ones w ere c hosen . d ifficult t o t ell,

e specially f rom p ost-cranial

( It i s o ften

m aterial,

w hether t he

v arious b ones f rom a ny e xcavation u nit a re f rom t he s ame a nimal o r n ot . T his

p roblem ,

t ogether w ith a r eluctance t o d estroy

v aluable

c ranial

b ones, m eant t hat o nly a f ew s amples w ere t aken, a lthough m ore m ay b e a nalysed i n t he f uture .)

T he r esults o f t he a nalyses a re t abulated

i n

T able X IX .

T he 613

C

v alues a re s omewhat s urprising a nd n ot e ntirely c onsistent .

T hose f or s teenbok a re s lightly m ore p ositive t han t he m odern r eadings ( -18,5°/ 00 18,6°/00

i n L ayer 6 ,

17,3 a nd 18,4 °/ 0 0 i n L ayer 9 a nd 19,4 a nd

i n L ayer 1 2 c ompared w ith t wo v alues o f 19 ,4°/0 0 f or m odern

s teenbok a t E lands B ay ). i n

a n

e nvironment

T hese a re t he k inds o f v alues o ne w ould e xpect

w ith

s omewhat

m ore

C4 g rass

t han

t oday .

c onclusion i s n ot b orne o ut b y t he r eadings o n h ares ( -22 ,6°/ 6 a nd 20 ,7 ° /

i n L ayer

0 0

h ares

1 2

o n t he

i n L ayer

c ompared w ith m odern v alues o f 20 ,0 a nd

18,0 '7 0

f or

a nything ,

m ore n egative t han t he m odern o nes .

g rass t han s teenbok d o ,

0 0

T his

c oastal

p lain).

T hese

values

a re,

i f

S ince h ares e at m ore

o ne w ould e xpect a ny i ncrease i n

C4

g rass i n

p rehistoric t imes t o s how u p m ore s trongly i n h ares t han i n s teenbok ( as i n t he " Bushmanland" c olumn

i n T able V III).

O ne c an o nly c onclude t hat

t he r elatively p ositive v alues f or s teenbok a re n ot c aused b y e ating g rass;

C4

t hey m ust h ave b een e ating s ome o ther p lants n ot f avoured b y

h ares, p erhaps s ucculents, o r m aybe s ome s alt-marsh g rasses f rom t he e dge o f t he V erlorevlei e stuary ( then o pen t o t he s ea ,

n ow c ut o ff f rom

i t).

T his

i s o bviously a t opic f or f urther i nvestigation ;

E lands B ay C ave

i tself

a re a lso r epresented t o b e a nalysed . a bove .

t he

( grazing a nimals s uch a s b uffalo a nd h artebeest

i n t he d eposits) a nd o ther n earby s ites w ill h ave s hould h elp e xplain t he

r ather p uzzling r esults

I t i s s afe, h owever , t o c onclude t hat t here h as b een n o m ajor

i ncrease p eriods

T his

m ore s amples f rom

i n t he

C4

g rass c omponent o f t he c oastal p lain d uring t he

t o w hich t hese a nimals d ate .

a nalysis

o f

m odern

a nimals

7 4

c an

T he

i sotopic p icture g ained f rom

t herefore

b e

a pplied

t o

t he

i nterpretation o f 6I 3 C r esults o n H olocene h uman s keletons e xcavated i n t he s outh-western C ape .

5 . S UMMARY

T he d ata p resented i n t his c hapter a llow t he e stimation o f t he 6I 3 C v alues o f p rehistoric h uman d iets i n e ach o f t he f our r esource z ones a vailable t o H olocene h unter-gatherers d efined b y P arkington

( 1972 ,

1 976a ).

i n t he s outh-western C ape , a s

A verage f igures a re

a bout 16 ,5° / 0 0 f or m arine f oods a nd 24° /

l ikely t o b e

0 f or t errestrial f oods o n

t he c oastal p lain a nd i n t he F old M ountain B elt .

I n t he K aroo, p lant

f oods a nd t he m eat o f b rowsing a nimals h ave t he s ame i sotopic r atios a s t hey w ould o n t he c oast o r i n t he m ountains, a nimals

i s a bout 2°/ .

0

m ore p ositive .

b ut t he m eat o f g razing

T here i s a s light i sotopic

g radient f rom t he h igh-rainfall C ape P eninsula t o t he m ore a rid a rea a round E lands B ay .

I n a ddition , t he d iet-collagen a nd m eat-collagen

s pacings h ave b een m easured f or s everal g roups o f a nimals,

a nd t he

i ndividual v ariation b etween a nimals o f t he s ame s pecies i n t he s ame a rea a ssessed . m arine

T he e dible p arts o f an umber o f s pecies o f p lants a nd

o rganisms

h ave b een a nalysed,

v ariations e stimated .

a nd

s easonal

a nd

i ndividual

T he o verall i mpression i s t hat p eople i n t he

s outh-western C ape h ad t wo c learly d istinguishable s ources o f d iet a vailable t o t hem s eafood a t 16 ,5° /

( short o f t rekking d eep

i nto t he K aroo),

. a nd t errestrial f ood a t 24 °/0 . .

n amely

Af ew p reliminary

a rchaeological m easurements i ndicate t hat t his p attern i s l ikely t o h old t rue

t hroughout

t he H olocene .

U sing t hese

t wo

v alues,

i t

i s

n ow

p ossible t o e valuate d ietary i nformation a nd t heories a bout s ubsistence s trategies a rrived a t f rom a rchaeological r emains .

7 5

C HAPTE l e 7

I SOTOPIC A NALYSES O F P REHISTORIC H UMAN S KELETONS

I n a ttempting t o a ssess h uman d iets i n a rchaeology ,

t he m ost d irect

e vidence c omes f rom t he h uman s keletons t hemselves.

T he c hemical

c onstituents o f b ones r eflect p rehistoric d iet i n a m uch m ore d irect w ay t han d oes t he f ood-waste r ecovered f rom e xcavations , a n umber

o f b iases

d iscussed

m easurements r epresent

i n

l ong-term

C hapter

4 .

i ntegrations

w hich i s s ubject t o I n addition,

o f a n

i ndividual 's

s uch d iet ,

p robably m ore t han 1 0 y ears i n t he c ase o f 61 3 C ( Stenhouse a nd B axter 1 979).

T HE S KELETAL S AMPLE

A c onsiderable

n umber

o f

p rehistoric

r ecovered f rom t he s outh-western C ape . i ndividuals

( listed

i n

R awlinson

h umans

s keletons

h ave

b een

S eventy-four r elatively c omplete 1 982)

a re h oused

i n t he

A natomy

D epartment o f t he U niversity o f C ape T own M edical S chool o r i n t he S outh A frican M useum a nd a re available f or s tudy .

U nfortunately ,

a l arge

p roportion w ere c ollected l ong a go , o r w ere e xposed b y e roding l andscape f eatures o r b y c onstruction p rojects a nd s o h ave l ittle o r n o a ssociated a rtefactual

m aterial

r adiocarbon d ated

( all

o r o ther

c ontextual

i nformation .

A f ew

t hese a re l isted i n T ables X X a nd X XI),

a re

b ut t he

v ast m ajority a re n ot .

I n v iew o f c urrent a rchaeological r esearch o bjectives i n t he s outhw estern C ape , p articularly t he h ypotheses a dvanced a bout c hanges i n s ubsistence p atterns d uring t he H olocene,

o nly d ated s keletons a re

i ncluded

h ave

t he b one

i n t his i tself .

s tudy . A n

N ineteen

s keletons

a dditional

h alf d ozen

o r

r adiocarbon s o

d ates

o n

h ave a ssociated

a rtefactual m aterial t hat e nables r ough d ating - u sually p ottery w hich , w henever f ound i n w estern C ape s ites, h as s o f ar d ated t o t he last t wo m illennia .

Ab asic a ssumption h ere i s t hat t he s keleton i s d irectly

a ssociated w ith t he p ottery .

7 6

S KELETONS I N T HE A RCHAEOLOGICAL C ONTEXT

T he t otal n umber o f s keletons d ated b y b oth t hese m ethods

i s s mall - a

m ere t wenty-four i ndividuals .

M oreover , o nly f ourteen c ome f rom t he

a rea

m obility

h ypothesis

i n

X X).

f or

w hich

f ormulated

t he

( these

s easonal

a re

l isted

T able

o bviously n ot a menable t o s tatistical a nalysis ,

w as

o riginally

A s ample t his

s ize

i s

e specially a fter h aving

b een d ivided i nto s ix c ategories, a nd s o t he c omments b elow s hould b e r egarded a s q ualitative .

T he s keletons f rom E lands B ay a nd t he O lifants R iver V alley ( Table X X ) w ere d ivided i nto t hree c hronological c ategories : 4 0 00

- 2 0 00

B .P .

a nd y onger t han

2 0 00

B .P .

o lder t han 8 0 00 B .P . , T hose

i n t he

c ategory p re-date t he h iatus i n h uman a ctivity a t E lands B ay .

f irst

T hey a re

t he r emains o f p eople w ho w ere l iving a t t he t ime w hen t he s ea-level h ad r ecently r eached m ore o r l ess i ts p resent p osition, a nd t he c limate s tabilised i nto a H olocene p attern . T hey w ere p robably s till h unting s ome o f t he l arge g razing a nimals t hat a re n ow e xtinct

( see C hapter 3 ) .

T heir l ifestyle i s l ikely t o h ave b een d ifferent f rom t hat o f l ater H olocene p eople i n an umber o f w ays ,

i ncluding s ocial o rganisation , t ool

t echnology a nd s ubsistence b ehaviour .

T here a re n o c oastal s keletons d ating t o b etween 8 0 00 a nd 4 0 00 B .P . S equences a t E lands B ay c ertainly s how a r eal a bsence o f p rehistoric s ettlement t here a t t his t ime , s ections o f t he c oast ,

b ut t here i s l ittle d ata a s y et f or o ther

s o w e h ave n o w ay o f k nowing h ow l ocalised t his

p henomenon w as .

T hree c oastal a nd f our i nland s keletons f rom t his a rea f all i nto t he 40 00 - 2 0 00 B .P . c ategory . c lassic

T his i s t he t ime f or w hich P arkington 's

v ersion o f t he s easonal m obility h ypothesis

T here a re p lenty o f s ites ,

i s m ost p lausible .

b oth i nland a nd a t t he c oast ,

w as n ot y et a d isruptive f orce . a nimals) a nd b y g athering ,

a nd p astoralism

P eople l ived b y h unting ( mostly s mall

a lthough p lant f oods w ere p erhaps n ot a s

i mportant a s t hey w ere l ater t o b ecome .

T here a re t hree i nland s keletons f rom t he p ost-2 0 00 B .P . t ime p eriod , w hich m ust h ave b een o ne o f r apidly c hanging l ifestyles a s p astoralism

7 7

s pread t hrough t he r esearch a rea .

I ndigenous h erds o f w ild a nimals w ere

p robably p artly r eplaced b y d omestic s tock .

P arkington ( 1983)

h as

h ypothesised t hat t he f ocus o f h unter-gatherer s ettlement s hifted t o a reas l ess a ttractive t o p astoralists s eeking g razing f or t heir a nimals , p erhaps t he s ea s hore a nd r ocky ,

m ountainous a reas,

i nstead o f t he

c oastal f oreland .

S TABLE C ARBON I SOTOPE R ATIOS O F S KELETONS

V iewed a gainst t his a rchaeological b ackground , s keletons

l isted

i n T able X X a re s tartling .

t he

81 3 C v alues o f t he

T he i sotope r atios o f t he

c oastal s keletons a re m arkedly d ifferent f rom t hose o f t he i nland o nes , a t

l east

i n t he 4 0 00 - 2 0 00 B .P .

p eriod .

c oastal g roup v ary f rom 12 ,3 t o 17 ,4°/ 17 ,3 t o 19°/0 0 .

1 3

0 0

C/

12

Cm easurements f or t he

,w hile i nland o nes v ary f rom

I f o ne l ooks a t t he c oastal s keletons a s o ne g roup ,

i ncluding t hose i n T able X XI a nd a lso t hose l isted i n R awlinson t he m ean + o ne s tandard d eviation i s 14 ,1+1,9°/

0 0

( 1982 ),

.

T his i s n ot a t a ll w hat o ne w ould h ave p redicted f rom t he a rchaeology o f t he a rea .

As easonally t ranshumant p opulation w hich i ntermittently

e xploited t he r esources o f t he s ea-shore s hould h ave b one c ollagen w ith 613 C v alues

i ntermediate b etween t he

v alues o ne w ould e xpect f or p ure

C3 based d iets a nd p urely m arine d iets i mportantly ,

( see n ext p aragraph).

o ne w ould e xpect t he s keletons f ound i nland a nd t hose f ound

a t t he c oast t o h ave s imilar i sotope r atios . c ase . t hat

T his i s c learly n ot t he

T he i sotope r atios o f s ome o f t he c oastal i ndividuals i ndicate a v ery

p erhaps, 1 00% .

M ost

l arge

p roportion

o f

t heir d iet

w as

i n s ome c ases ( e .g . U CT 1 12 a t 11,2 °/ 0

0

o f

m arine

o rigin,

,T able X XI) a lmost

T he i nland s keletons a re t he r emains o f p eople w ho m ay w ell h ave

v isited t he c oast - t he h ighest v alue f or a 4 0 00 - 2 0 00 B .P . i nland s keleton ( -17 ,3 ° /

0 0

f or S AM-AP 1 449 ) o verlaps w ith t he l owest v alue f or

ac oastal s keleton ( -17 ,4 ° / E lands B ay ). h owever ,

a nd

0 0

f or a n a s y et u naccessioned s keleton f rom

T hey d id n ot s pend a l arge p roportion o f t heir t ime t here , i t

i s

c lear

t hat

i f t hey d id p lan

t heir

s ubsistence

a ctivities i n s uch a w ay a s t o m ake t he m ost o f s easonal f ood r esources , t heir p rogramme w as n ot t he s ame a s t hat o f t he c oast d wellers . s omewhat s urprising,

T his i s

e specially i n v iew o f t he c omparatively s hort

d istance b etween t he O lifants R iver V alley a nd t he c oast .

7 8

I t w ould o bviously b e e xtremely u seful t o b e a ble t o t ranslate t hese 613

v alues i nto p ercentages o f m arine a nd t errestrial

f ood c onsumed .

U sing t he e stimates o f t he i sotopic r atios o f a ll t he d ifferent k inds o f f ood a rrived a t i n t he p revious c hapter, s omeone e ating a n e ntirely m arine d iet ( average 6I 3 C v alue 16 ,5° / 0 0 ), a nd h aving a d iet-collagen f ractionation f actor o f + 5,1 °/, , s hould h ave a b one c ollagen 6I 3 C r eading o f 11,4°/o 0 .

( Interestingly , a lthough t he s keletons o f c oastal

p eople e xhibit a r ange o f i sotope v alues d epending o n t he a mounts o f s eafood 11

0

/ ,

1 981,

i n t heir d iets, 0

t here

s eems t o b e a c ut-off p oint o f a bout

a t t he p ositive e nd o f t he s cale a ll o ver t he w orld ( Tauber

C hisholm ,

N elson a nd S chwarcz 1 982).) A t t he o ther e nd o f t he

s cale , a n e ntirely t errestrial d iet ( average 61 3 C v alue 24°/0 0 ) s hould p roduce b one c ollagen o f 18,9 °/

0 •

F ifty p ercent m arine a nd f ifty

p ercent t errestrial f ood s hould g ive b one c ollagen o f 15 ,2° /0

0

.

C alculations s uch a s t hese a re p robably m ore o r l ess c orrect, b ut t oo m uch r eliance s hould n ot b e p laced o n t he e xact p roportions o f t he d ifferent f oods c onsumed s ince a n a s y et u nsolved m etabolic p roblem m ust b e c onsidered h ere .

T his

i s t he q uestion o f t he p ossible d ifferential

c ontribution o f t he v arious c omponents o f f oods t o a ny p articular t issue i n t he c onsumer , p rotein

i n t his c ase b one c ollagen .

s ynthesis

i s

a c omplicated

d ifferent s tarting m aterials i n a ny p rotein , a cids .

c ollagen

C ollagen i s a p rotein , a nd

p rocess

w hich

i s m ade u p o f e ssential

a nd n on-essential

L ike a mino

E ssential a mino a cids h ave t o b e d erived f rom p rotein i n t he

a cids i n b one c ollagen a re e ssential. e ssential

a mino

a cids

( mainly

O ne-fifth o f t he a mino

T he o ther f our-fifths a re n on-

g lycine

m anufactured f rom d ietary c arbohydrate .

a nd

T his

p raline) i s

t he

t hat

p referred

i ndividual w hose d iet c ontains p lenty o f c arbohydrate ,

d iet i s r ich i n p rotein b ut l ow i n c arbohydrate , a cids

u se o f

i ndividuals w ith d ifferent d iets .

d iet, t hey c annot b e s ynthesised b y t he b ody .

i n a n

c an m ake

c an

b e

p athway

b ut i f t he

t he n on-essential a mino

i n b one c ollagen m ay a lso b e d erived f rom d ietary p rotein .

I n a

" normal" m odern W estern d iet c onsisting m ainly o f c arbohydrate w ith a p rotein s upplement , a s m uch a s f our-fifths o f b one c ollagen c ould b e c arbohydrate-derived

( Gevers ,

p ers .

7 9

c omm . ).

M ost p rehistoric h unter-gatherer a nd p astoralist d iets w ere p robably n ot a s r ich i n c arbohydrate a s t hose o f m odern W esterners . m any f oods ( Iridaceae c orms ,

s hellfish )

N evertheless,

c ontain c onsiderable a mounts o f

c arbohydrate t hat m ust b e r eflected i n h uman s keletal c ollagen .

I n

o rder f or a ll c ollagen 6I 3 C r eadings t o b e d irectly c omparable , t he c ollagen s hould c ome f rom p eople w hose d ietary p rotein/carbohydrate r atio i s s tandardised . p robably n ot t he c ase .

I n t he s ample i n T ables X X a nd X XI,

t his i s

I f c oastal p eople g athered s hellfish a s a s taple

( 15% p rotein - u npublished l ists c ompiled b y B ill B uchanan) w hereas i nland p eople c ollected I ridaceae c orms

( 3-7% p rotein a ccording t o

A rcher 1 982 ) t here i s l ikely t o b e s ome b uilt-in b ias i n t he 6 ' 3C v alues o f t he t wo g roups.

M oreover ,

i f u nderground p lant f ood c ollection

b ecame m ore i mportant i n t he l ast 2 0 00 y ears , b oosting t he c arbohydrate c omponent o f t he d iet a t t he e xpense o f p rotein ,

t his c an b e e xpected t o

h ave a n e ffect o n c ollagen 6I 3 C v alues .

F ortunately , t he s ituation i n t he s outh-western C ape i s l ess c omplicated t han

i t

c ould h ave

s ignatures

b een .

T he t errestrial

f ood

f or b oth t heir p rotein a nd c arbohydrate

r esources

h ave

c omponents ,

C3

w hereas

m arine o rganisms l ikely t o h ave b een e aten b y p rehistoric p eople a re i sotopically m uch h eavier i n b oth p rotein a nd c arbohydrate . t hat t he o bserved d ifference

i n 613

C

T his m eans

v alues b etween i nland a nd c oastal

p rehistoric h uman s keletons i s l ikely t o b e m eaningful .

T here i s a c onsiderable a mount o f v ariation i n t he i sotope r atios o f t he c oastal s keletons :

t he 613

f rom 11,2 t o 1 .7,4 °/ 0

0

C

v alues o f t hose i n T ables X X a nd X XI r ange

. ( The i nland s ample i s t oo s mall t o r eveal

i nformation o f t his k ind .)

T he o ldest

s keleton

i ndividual f rom E lands B ay a lmost 1 1 0 00 y ears o ld , 0

i n

t he

s ample,

a n

h as a 6I 3 C v alue o f

. T his p erson a te l ittle e lse b esides m arine f oods f or a t

l east t he l ast 1 0 y ears o f h is l ife , a nd s o m ust h ave s pent m uch , a ll o f h is t ime a t t he c oast .

i f n ot

T his i s p articularly i nteresting i n v iew

o f t he f ood-waste d ating t o t his t ime p eriod r ecovered f rom E lands B ay C ave .

T hese r emains c ontain a h igh p roportion o f t errestrial a nimal

b ones ,

m uch h igher t han l ater H olocene l evels,

a nd o ne m ight t herefore

h ave e xpected e arly H olocene c oastal s keletons t o h ave h ad a m ore t errestrially-orientated d iet t han t heir l ater c ounterparts .

T his d oes

n ot s eem t o b e t rue: t hree o ut o f t he f our E lands B ay s keletons d ating

8 0

t o 80 00 - 1 0 0 00 B .P . h ad s trongly m arine-based d iets. i llustration o f t he d ifference b etween s tudies,

a nd

o f

t he

I ndividuals a lmost

f act

t hat

t wo

c annot

i ndividual-based

always

b e

e quated .

c ertainly m oved b etween a n umber o f a rchaeological

s ites d uring t heir l ifetimes . d epends o n l ocal

t he

s ite-based a nd

T his i s ag ood

T he f ood-waste i n a ny p articular s ite

e nvironmental

c onditions

w hich m ay o r m ay n ot

b e

r eflected i n t he i sotope r atios o f t he b ones o f t he p eople w ho u sed t hat s ite ,

d epending o n h ow m uch t ime t hey s pent t here .

T he p ost-2 0 00 B .P .

s keletons i n T able X X r equire s ome c omment . A ll c ome

f rom t he O lifants R iver V alley , w e h ave n ot s o f ar b een a ble t o f ind a ny a t E lands B ay . 4 660) .

O f t he t hree ,

T he o ther t wo ,

r espectively ,

o nly o ne i s c ertainly p rehistoric ( SAM-AP

w ith r adiocarbon d ates o f 2 50+90 a nd 2 70+100

f all

i nto

t he

n on-linear

p art

o f

t he

B .P .

r adiocarbon

c alibration c urve a nd s o m ay i n f act d ate a nywhere f rom t he s eventeenth t o t he n ineteenth c enturies

A .D .

T hey c ould ,

o f f arm l abourers f rom h istoric t imes, b orne

t herefore b e t he

r emains

a nd t his p ossibility m ust b e

i n m ind t hroughout t he d iscussion t hat f ollows.

T here

i s n o

i nformation o n b urial p attern o r a ssociated a rtefacts t hat m ight h elp t o s olve t his p roblem ,

s ince b oth i ndividuals a re r epresented o nly b y

s kulls p resented t o t he S outh A frican

M useum e arly t his

c entury .

T hey

a re d escribed i n t he c atalogue a s " Bushman" s kulls, a nd c ertainly t he o verall s uch

c ranial m orphology a nd t ooth-wear p atterns a re

a d escription.

u nremarkable

One

s table c arbon

of

them

i sotope

f irst

c omponent

t ime

i n

i n t he d iet .

m arine/terrestrial

i nland

c onsistent w ith a s easonal

m ust

0

has 0 0

e ntirely

, b ut t he o ther

,a s d oes S AM-AP 4 660 .

s keletons,

an

S uch v alues,

s how a s ubstantial

m arine

T hese i ndividuals p robably d id e at f airly m ixed

d iets .

t he 4 0 00-2 0 00 B .P .

2 6)

r atio o f 18 ,1 °/

( SAM-AP 2 7) h as a r eading o f 16° /0 f or t he

( SAM-AP

c onsistent w ith

I sotope

r eadings

s uch a s

m obility h ypothesis

s keletons ,

t hese a re

f ar m ore

t han t hose o btained f or

b ut a ny d efinite c onclusions a bout

a wait t he a nalysis o f a l arger p rehistoric

s ample o f b oth

t his

i nland

a nd c oastal s keletons .

T here i s , p ost-2

h owever ,

0 00

B .P .

n o s upport ,

c oastal

e ither i n t his d ata o r i n t he r esults f or

s keletons

f rom f urther t o t he

s outh

( see T able

X XI) f or t he t heory t hat a p opulation t hat o ccasionally v isited t he c oast

i n t he e arly a nd m id-Holocene w as d riven b y c ompetition w ith

8 1

p astoralists t o s pend m ore t ime t here i n t he l ast 2 0 00 y ears.

I t i s

q uite p ossible t hat t he p opulation a t t he c oast d id i ncrease i n t he l ast 20 00 y ears, t hus a ccounting f or t he p roliferation o f a rchaeological s ites , a t l east a t E lands B ay ,

d ating f rom t his r ecent p eriod .

H owever ,

t his d id n ot r esult f rom t he b reakdown o f a p reviously c yclical s ystem a s p ostulated b y P arkington

( 1983 ).

I sotopic m easurements t herefore c onfirm t hat p eople w ere e xploiting c oastal r esources t hroughout t he H olocene,

a nd a ll a long t he c oast.

W hat i s u nexpected i s t he i ntensity o f e xploitation b y s ome i ndividuals, a nd a lso t he a mount o f v ariation b etween

i ndividuals .

P opulation

d ensities m ay h ave c hanged , a nd c ompetition f or r esources i ncreased , t hus l eading t o a n i ncrease i n t he n umber o f p eople a t t he c oast, b ut a daptations

t o

t hese

p ressures

d id

n ot

i nvolve

p reviously i ntermittently e xploited r esource .

8 2

g reater

u se

o f

a

C HAPTER 8

I MPLICATIONS O F S KELETAL A NALYSES F OR A RCHAEOLOGICAL R ESEARCH I N T HE S OUTH-WESTERN C APE

S ince t he

r esults o f t he

p resented

i n

C hapter

i nterpretation o f t he

i sotopic a nalyses

7 c learly

d o

n ot

o f t he h uman

f it

i nto

t he

s keletons p revailing

a rchaeology o f t he s outh-western C ape,

i t

i s

a ppropriate t o e xamine t his i nterpretation m ore c losely i n a n a ttempt t o r econcile v iew .

t he

m ore

c onventional

a rchaeological

a nd

i sotopic p oints

T he b est p lace t o b egin i s w ith t he e vidence t hat

f ormulation o f t he s easonal m obility h ypothesis . s ummarised

i n

C hapter

i nconsistencies

l ed t o t he

T his h as a lready b een

3 s o d iscussion h ere w ill

p ointing o ut o f p roblems a nd

o f

b e

l imited t o t he

i n t his a pproach ,

a nd t he

m ention o f n ew d ata t hat h as e merged f rom m ore r ecent r esearch .

M ost o f t he s easonal i ndicators u sed t o d emonstrate s ummer o ccupation o f t he F old M ountain B elt a nd w inter o ccupation a t t he c oast a re p ositive i ndicators .

T he

c ase f or b oth p henomena

i s f airly c onvincing,

b ut

i nsufficient a ttention h as b een p aid t o t he p roblem o f s howing t hat w inter o ccupation o f t he m ountains a nd s ummer o ccupation a t t he c oast d id n ot o ccur .

I t i s d ifficult t o f ind e vidence t hat p eople w ere n ot

s omewhere a t a ny p articular s eason ,

b ut t his m ust b e d one ,

o r s ome o ther

w ay f ound t o s how t hat t he p eople w ho w ere a t t he c oast i n o ne s eason w ere t he s ame a s t hose w ho w ere i nland a t a nother .

T he t echnique o f m easuring j uvenile d assie a nd s eal b ones h as a b uilt-in b ias .

I t h as b een s hown q uite c onclusively t hat m any o f t he d assies a t

D e H angen w ere k illed i n t he s ummer , a nd t hat t he s ite m ust t herefore h ave b een o ccupied i n s ummer . s ite a re j uvenile , i n T able X XII 1 971).

v ery

a nd c an b e a ged ,

f ew o f

t he r esults o f t his a geing a re s hown

( a t able f irst p ublished i n P arkington a nd P oggenpoel

T hese f igures h ave b een

t hese d assies

4 6% o f t he d assie m andibles f rom t his

i nterpreted a s f ollows :

" Almost

a ll

o f

c ould h ave b een k illed b etween N ovember a nd F ebruary a nd t hem

c ould

( Parkington a nd P oggenpoel

o nly

h ave

1 971:30).

b een

k illed

T echnically ,

d uring t his

t he

i s t rue ,

w inter" b ut t he

s easonality i s n ot a s c learly d emonstrated a s t hese a uthors w ould h ave u s b elieve .

T heir c onclusion m eans t hat w e h ave t o a ssume t hat t he 2 7

8 3

d assies

w hose

d entition

i ndicates

S eptember t o N ovember w ere , p eriod ,

w hile t he

2 6

t hat

i n f act ,

i ndividuals

t hey

w ere k illed a ny t ime f rom

c lustered t owards t he e nd o f t his

w ho

c ould h ave d ied d uring F ebruary ,

M arch o r A pril h appen t o h ave s uccumbed a t t he b eginning o f t heir r ange . A s imilar s kewing o f t he o ther 3 1 a nimals i s n ecessary .

T hus ,

a lthough

n early a ll t he d assies c ould h ave b een k illed b etween N ovember a nd M arch,

i t

i s

m ore

l ikely

t hat

t hey

w ere

a ctually

k illed

b etween

S eptember a nd A pril - a t ime r ange e xtending over t wo-thirds o f t he y ear .

F ive a nimals w ere k illed i n t he o ther t hird o f t he y ear .

a ddition,

t he

5 4%

o f

t he

j aws

i n

t he

s ample

t hat

a re

a dult

I n o r

a diagnostic a re a ssumed t o b e t hose o f a nimals k illed a t t he s ame t ime o f y ear a s t he d iagnostic

i ndividuals .

W hen o ne t akes

i nto a ccount t he

g reater e ase w ith w hich j uvenile d assies c an b e k illed ,

i t s eems q uite

p ossible t hat p eople c oncentrated o n d assies a t t imes a t w hich t hey w ere e asy t o o btain,

a nd , p erhaps, o n o ther f ood r esources a t o ther t imes.

W hen u sing t his k ind o f e vidence t o s how t hat a s ite w as o ccupied i n o ne s eason o nly ,

i t i s v ital

t o b e a ble t o s how t hat i t w as n ot

o ccupied a t

o ther s easons.

T his a lso a pplies t o o ther s easonal i ndicators, s uch a s t he s talks o f b edding g rass f rom D e H angen w hich h ave i nflorescences o n t hem . f lower i n s ummer , i s

a lso

t he

a vailable .

s o t hese m ust h ave b een c ollected i n s ummer ,

o nly

s eason

w hen

d ry

g rass

I n w inter t he g rass i s y oung ,

s uitable

f or

o f

t ortoise

b ones

a t

e vidence o f s ummer o ccupation . a ctive , t hem

t oo,

h as

b een

T he h igh t aken

a s

i s t rue t hat t ortoises a re m ore

i n s ummer ,

i n t he m iddle o f w inter .

i s

g reen a nd d amp a nd p eople m ay

s ite,

W hile i t

a nd h ence m ore v isible ,

e ven

t his

b ut t his

b edding

w ell h ave p referred t o u se s ome o ther m aterial f or b edding . i ncidence

G rasses

i t

i s q uite p ossible t o f ind

A r ecently

i nitiated

t ortoise

m onitoring p rogramme a t E lands B ay c onfirms t his .

S easonal e vidence f rom p lant f ood r emains, t oo , T he s eeds o f E uclea s pp"

0 18q ,

X "

a re

i n P arkington

1 976a)

i s a t b est a mbiguous .

R hus a nd N ylandtia ( referred t o a s " Seed

c ertainly f rom

f ruits

t hat

w ould

h ave

r ipened i n s ummer , b ut t he k inds o f p lant f oods t hat a re a vailable i n w inter a re g enerally s oft a nd f leshy a nd w ould n ot l eave a ny t race i n a n a rchaeological

s ite

( e .g

T rachyandra ,

A lbuca ,

C yphia a nd m any o thers).

M oreover , a ll t he a bove f ruits a re e aten b y b irds a nd b aboons a nd a re

8 4

k nown t o b e c arried i nto c aves b y n on-human a gencies . a c ave

s ite

i s

n ot,

t herefore,

a n

o ccupation o f t he s ite b y h umans .

i nfallible

T heir p resence i n

i ndicator

C orms o f M oraea

o f

s ummer

B abiana a nd W atsonia

a re m ost p alatable a nd e asily f ound i n s ummer , b ut a re a lso a vailable a t o ther s easons.

T he b aboons i n t he C ape P oint N ature R eserve i nclude

c orms i n t her d iets e very m onth e xcept J une a nd J uly ( Hall

1 962 ).

T he f inal s trand o f e vidence w hich s uggests s ummer o ccupation o f t he m ountains

c omes

f rom

r eports

b y

e arly t ravellers .

P arkington

r ecords

t hat " almost a ll m eetings w ith S oaqua i n t he O lifants R iver V alley t ook p lace

d uring

t he

d ry

s eason"

( Parkington

1 977:152),

p robably a lso t he b est t ime t o u ndertake t rips

b ut

i nto t he

t his

i nterior .

w as T he

e vidence i s n ot c onfirmed b y e arly t ravellers w ho d id n ot m eet S oaqua i n t he O lifants R iver V alley d uring t he w et s eason .

Q uite o bviously , ar iver v alley w ould h ave b een a g ood p lace t o b e w hen t he a nnual w atercourses w ere d ry d uring t he s ummer,

a nd s ome o f t he

r eports o f e arly t ravellers c onfirm t hat t he K hoisan d id a dopt t his p olicy .

W hether o r n ot t hey a lso s pent t he w et s eason " away f rom t he

r iver i n o rder t o e xploit t he m ore t emporary p otentials t o t he e ast a nd w est"

( Parkington

1 977:156 )

r emains a s ubject

p lace w here t hey d id n ot s pend i t ,

f or f uture

r esearch .

a t l east o n a ny r egular b asis ,

O ne

w as a t

t he c oast .

S easonal i ndicators f rom E lands B ay C ave a re j ust a s i nconclusive a s t hose f rom D e H angen .

T he p ost-4 0 00 B .P . l ayers c ontain t he b ones o f

b irds t hat a re w inter v isitors t o t he n earby V erlore V lei e stuary , a s f lamingoes.

T he s eal m andibles a re

s uch

l argely t hose o f y earlings,

q uite p ossibly c ollected w ashed-up o n t he b each r ather t han h unted ( see d iscussion

i n

C hapter

3 ) .

S eals

h ave a r estricted b reeding

s eason a nd

p ups b ecome y earlings i n w inter , w hich i s a lso t he t ime w hen w ash-ups f rom w inter s torms a re m ost

c ommon .

P erhaps

t he

c ombination o f

w inter w eather a nd s uddenly h aving t o f ind t heir o wn f ood , b eing a ble t o d rink t heir m others ' m ilk ,

s tormy

i nstead o f

i s af atal o ne f or s ome p ups.

T he i dea t hat b lack m ussels a re b est e aten i n w inter b ecause t hey a re l ess l ikely t o b e t oxic f rom r ed t ide o rganisms t han t hey a re i n s ummer , w as o riginally a dvanced w ith c aution i t h as b een r epeated a s a t ruism

( Parkington

1 976a ).

N evertheless ,

i n s ubsequent w ork o n w est

8 5

c oast

s ites

( Robertshaw 1 977,

1 979a ,

1 979b , B uchanan e t a l. 1 978).

S ince m ussels

c an r emain t oxic f or u p t o f our m onths a fter a r ed t ide h as d isappeared ( Grindley a nd S apeika 1 969), o ne c an a rgue t hat i t w ould b e e xtremely u nsafe t o e at m ussels a t a ny s eason o n a s hort E ating m ussels a t a ll c oastal

r equires a n

v isit t o t he

s hore .

i ntimate a nd f ull-time k nowledge o f

w aters .

S ome p ositive e vidence f or s ummer o ccupation o f E lands B ay C ave i s s tarting t o e merge . m ortality

B each s urveys r eveal t hat 8 0% o f t he s horebird

i n t his a rea o ccurs b etween N ovember a nd A pril,

a nd t hat 8 0%

o f t he C ape c ormorants r ecovered f rom t he c ave w ere c ollected d uring t hese m onths.

M oreover , t he p resence o f j uvenile C ape c ormorants o n

o pen m iddens n earby a lso p oints t o s ummer o ccupation A very 's

w ords:

" It

( Avery

1 981) .

I n

i s b ecoming a pparent t hat p eak a vailability o f

d ifferent r esources d oes n ot n ecessarily c oincide a nd t hat a ny s trategy i nvolving e xploitation o f t he

v arious p eaks m ust b e m ore c omplex t han

t he s traight f orward s easonal t ranshumance s o f ar p roposed "

T hus ,

( 1981:86 ).

s everal l ines o f e vidence p oint t o t he n ecessity f or t he s easonal

m obility h ypothesis t o b e r eviewed . d o n ot

c onfirm t hat

S table c arbon i sotopic m easurements

t he c oast a nd t he m ountain b elt s upplied d ifferent

f acets o f a s ingle s ubsistence e conomy f or p rehistoric p eople .

M uch o f

t he e vidence o n w hich t he t heory w as o riginally b ased r equires r ee xamination ,

w ith p articular a ttention p aid t o d emonstrating t hat s ites

w ere n ot o ccupied i n c ertain s easons .

T he a pproach e mployed u p t o n ow ,

i . e . " Ruling o ut o ccupation o f a rchaeological s ites i s g enerally t ackled b y r uling

i n o ccupation n earby a nd s uggesting a m ovement b etween s ites"

( Parkington

1 976a:77),

i s n o

l onger a dequate .

M oreover ,

s ince t he

r ecognition o f t he

l ikely e ffects o f p astoralism o n s easonal

( Parkington

1 984),

t ypical

1 983,

D e H angen

c an

n o

l onger b e

r egarded a s

s ummer o ccupation s ite i n t he m ountains ( Parkington 1 971,

T he w hole o f t he s equence i n t his c ave p ost-dates 2 0 00 B .P . , f alls

i n

t he

p eriod

f or

w hich

P arkington

f ragmentation o f t he h unter-gatherer p opulation b ased

i s

a ccess

t o

a v ariety

o f

r esource

n ow

z ones

a

1 977).

s o t hat

i t

h ypothesising

i nto s mall

i n t he m ountains a nd o thers o n t he c oast.

s cheme ,

m obility

b ands ,

s ome

A ccording t o t his w ould

b e

g reatly

r estricted a nd t he b ands i n t he m ountains w ould h ave t o r emain t here f or m uch o f t he y ear .

T his i s ap lausible h ypothesis, b ut m eans t hat m uch

8 6

o f t he e vidence t hat w as p reviously t aken t o i ndicate t hat s easonal m obility o ccurred w ill h ave t o b e t urned a round t o s how t hat i t d id n ot o ccur , o r a t l east n ot i n t he f orm o riginally p roposed .

T he

i dea t hat

r esources ,

s ome g roups

o f K hoisan p eople

c oastal

s pending m ost o f t heir t ime c ollecting s hellfish a nd c ombing

t he b eaches f or s tranded w hales , J an

r elied h eavily o n

v an R iebeeck m entions

" strandlopers"

a rrival a t t he C ape i n 1 652 : t hree t ribes o f p eople , S trandlopers o r

s eals a nd c oastal b irds i n h is

j ournal

i s a n o ld o ne . s oon a fter h is

" This T able V alley w as v isited a nnually b y

s imilar i n d ress a nd c ustoms .

O ne i s c alled t he

W atermen , b ecause t hey l ive o n m ussels w hich t hey

f ind o n t he r ocks a nd

s ome r oots f rom t he e arth _ .

T hey .

h ave n o

c attle . - t he W atermen l ive p ermanently i n t his T able Valley" ( Thom 1 952 :81) .

O n s everal o ccasions D utch s ettlers h oping t o t rade f or s heep

a nd c attle a t S aldanha B ay w ere d isappointed t o e ncounter " no H ottentots o ther t han d estitute S trandlopers" o n t he c oast, h aving m oved i nland

( Thom

1 958 :152 ,

b ecause o f t heir p overty : t ortoises,

1 93 ,

2 43).

t hose w ith c attle

T he D utch d espised t hem

" these p oor p eople h ad n othing t o b arter b ut

o strich e gg s hells a nd s imilar t rash"

( Thom 1 952:93)

c attle-owning K hoi s eem a lso t o h ave v ictimised t hem : t heir

f ollowing

h ereabouts ,

a re

r ebels

b eing f ar a way ,

a nd

o utlaws

w eak a nd p oor ,

s ome o f t he o ther t ribes c ome t his w ay , h iding-places"

A t

l east

.

a nd

" All o f t hem w ith

t hese

p eople

a re l eft a lone ,

l iving

e xcept w hen

w hen t hey a re d riven

i nto t heir

( Thom 1 954 :172).

o ne a rchaeologist

h as

s upported t he

i dea t hat t here w ere s ome

L ater S tone A ge p eople w ho l ived o n t he s hore n early a ll t he t ime . G arth S ampson ( 1974 ) h as d ivided S outhern A frican s tone t ool a ssemblages w ith " Cape C oastal" p ottery ( thick q uartz-tempered p ottery , o ften w ith r ounded o r p ointed b ases, t hree t ypes ,

v essels

a nd h aving lugs o r s pouts)

o ne o f w hich h e c alls " Strandloper" .

i nto

T hese a ssemblages a re

a ssociated w ith s hell m iddens, a nd h ave n umbers o f large u nretouched f lakes a nd f laked c obbles ,

b ut f ew f ormal t ools .

T his v iew h as n ot b een p opular i n r ecent y ears, m ost a rchaeologists p referring t o s ee c oastal s ites a s s easonally-used s egments o f a n a nnual r ound 1 976b ,

( Deacon, 1 977,

H .J .

1 981).

1 969,

1 970,

P arkington

1 972,

1 976,

P arkington

( 1976a) a nd R obertshaw

8 7

1 972,

1 976a,

( 1979a)

h ave

s uggested t hat h istorical r ecords o f p eople w ho s pent a l arge p roportion o f t heir t ime o n t he c oast m ight

i ndeed

b e

a ccurate,

H olocene h unter-gatherers . ( trading w ith s ailors)

( as q uoted a bove f rom T hom 1 952 , b ut

t hat

I nstead ,

t hese

p eople

w ere

1 954 , n ot

1 958 )

t ypical

t hey h ad a dapted t o a n ew r esource

w hich m ade y ear-round c oastal o ccupation a n

a ttractive a nd v iable p roposition .

T his b ehaviour p attern w ould t hus b e

e xpected t o h ave o ccured o nly a fter E uropean e xploration o f t he c oast .

T he d ata p resented h ere r efute b oth t hese h ypotheses, a t l east i n t he w estern

C ape .

S ome

( though

b y

n o

m eans

a ll)

o f t he p eople

w hose

s keletons h ave b een f ound a t t he c oast d o s eem t o h ave l ived a lmost e ntirely o ff m arine f oods, ( 1965).

a s e xpected b y S ampson ( 1974) a nd b y G ee

L ee b elieved t hat h unter-gatherers w ith a ccess t o

c oastal

r esources w ould h ave u sed t hem a s a s taple j ust a s t he ! Kung S an r ely o n t he m ongongo n ut .

M oreover ,

i t i s a b ehaviour p attern w ith a t ime-depth

o f a t l east 1 0 0 00 y ears, a s t he c onsistently p ositive c arbon i sotope r atios f or s ome c oastal s keletons t hroughout t he H olocene s how .

A lice H ausman,

i n a p hysical

p opulations

i n S outh A frica

m easurements

o f t he

c oastal

a nthropological ( Hausman

m embers

t hose o f s keletons f rom e lsewhere

1 980)

o f

s tudy o f p rehistoric

f ound t hat

t he

c ranial

h er s ample w ere d ifferent f rom

i n t he c ountry .

T his m ust m ean t hat

t here w as s ome g enetic, a s w ell a s e conomic d ifferentiation b etween i nland a nd c oastal p opulations .

M ore w ork n eeds t o b e d one o n t his,

s ince h er s amples w ere s mall a nd d istributed t hrough a l arge t ime r ange , b ut a s t he n umber o f d ated s keletons i mprove . p rep .) o f

M any a uthors

h ave

c oastal

i ncreases,

( Parkington 1 976b , a bout

t he s ituation w ill

R obertshaw 1 979a ,

e xpressed

r eservations

w hether

r esources

( particularly s hellfish ,

t he

B uchanan i n

c arrying

i f t hese w ere a s taple )

w ould b e s ufficient t o s upport a h uman p opulation o f a ny s ize t han a f ew m onths o f t he y ear .

c apacity

f or m ore

P arkington ( 1976b) c alculated t hat a

s ingle k ilometre o f r ocky s horeline a t E lands B ay c ould s upport t en p eople f or s ix m onths o f t he y ear i f t hey h arvested 4 0% o f t he s hellfish p opulation , m ussels .

a nd

T his

i f a t hird o f t heir d iet

i s a v ery s hort

c onsisted o f l impets

l ength o f c oastline ,

a nd e ven

i f p eople

h ad t o g o s ome d istance o ver s andy b eaches t o r each t he n ext o utcrop ,

a s w ould h ave b een t he c ase a t E lands B ay ,

8 8

a nd

r ocky

t he t otal a rea

r equired t o n ecessary

s upport a b and w ould p robably b e n o g reater t han t hat

i nland .

O ther s ites h ave a ccess t o g reater l engths o f r ocky s horeline .

A t

P aternoster t en k ilometres o f r ocky c oast a re a vailable w ithin a f ive k ilometre r adius o f t he m idden e xcavated b y R obertshaw .

T wenty p eople

c ull i ng t hirty p ercent o f t he s hellfish p opulation a nd e ating a d iet w hich w as o ne-third s hellfish c ould h ave l ived t here a ll y ear r ound ( Robertshaw

1 979a).

A ll

t he

t errestrial

p lant

a nd

a nimal

f oods

a vailable i n t he c oastal p lain a nd t he F old M ountain B elt a re a lso t o b e f ound n ear t he c oast .

C ertain p lants m ay b e s omewhat l ess a bundant ,

b ut

i t w ould c ertainly n ot h ave b een n ecessary t o g o v ery f ar t o a chieve a v aried a nd b alanced d iet .

C ONCLUSIONS

613 C

m easurements o f a n umber o f h uman s keletons f rom t he s outh-western

C ape o f S outh A frica ,

c ombined w ith a k nowledge o f t he

o f t he f oods a vailable

i n d ifferent

r esource z ones ,

t wo s ubsistence p atterns a mongst H olocene p eople . 40 00 - 2 0 00 B .P . time p eriod ,

i sotopic

i ndicate a t

r atios l east

I n t he c ritical

i nland-dwellers c onsumed v ery l ittle

m arine-based f ood , w hereas c oastal d wellers a te c onsiderable a mounts. T hese r esults c ontradict s ubsistence s trategy m odels w hich i nvolve s easonal m ovements b etween c oast a nd m ountains .

A b etter m odel

w ould b e o ne

r esources

i s

c ontained ,

w ay o f l ife .

i n w hich t he

r egarded a s a s eparate,

a nd p ossibly s ometimes

a nnual

b ut

r ound ,

s elf-

T he r emains o f i nland p eople a re t oo s canty t o

a llow a ny c onclusions t o b e d rawn a bout b ehaviour ,

e xploitation o f c oastal

t heir p atterns

o f

s ubsistence

i t i s q uite p ossible t hat t hey d id h ave s ome s ort o f a nd m ay o ccasionally

s easonal c ycle w as,

h owever ,

h ave

v isited

t he

c oast .

T his

v ery d ifferent f rom t hat o f p eople w ho

l ived o n t he s ea-shore .

T he

i sotopic d ata a lso d emonstrate t hat

i ntensive c oastal e xploitation

i s n ot a n ew p henomenon i n t he w estern C ape, f ar a s t he b eginning o f t h2 H olocene . s ome

i ndividuals

l ived

i most

T hroughout t he l ast 1 0 G OO y ears ,

e ntirely

8 9

i t d ates b ack a t l east a s

o ff

m arine

f oods

O ther

s keletons f ound a t t he c oast,

h owever, a re t he r emains o f p eople w ho

c onsumed m uch m ore t errestrial

f ood .

a s

n ot

y et,

u nclear

c hronologically i nvestigation .

a nd

w e

d o

p atterned .

I t

T he m eaning o f t his v ariation i s , k now

i s

w hether

c ertainly

i t

i s

s patially

a t opic

f or

o r

f uture

W hile c oastal p opulation d ensity m ay w ell h ave i ncreased

d uring t he l ast 2 0 00 y ears i n r esponse t o t he a rrival o f p astoralism a t t he C ape ,

a daptations t o t his n ew w ay o f l ife d id n ot

i nclude m ore

i ntensive u se o f p reviously i ntermittently-exploited m arine f oods .

T hese c onclusions h ave c onsiderable i mplications f or a rchaeological r esearch i n t he s outh w estern C ape .

F urther w ork o n s easonal p opulation

m ovements m ust s how t hat p eople w ere n ot i n p articular a reas i n s ome s easons , a s w ell a s t hat t hey w ere t here i n o thers .

S tudents o f c oastal

s ites w ill h ave t o r ecognise t hat t hese a re l ikely t o r epresent a l arger s egment o f t he s ubsistence e conomy o f c ertain p rehistoric g roups t han p reviouly s upposed . v arious

E stimates o f g roup s ize ,

f oods a nd t he r elative

k ilojoule i ntake f rom

i mportance o f d ifferent

n eed t o b e a djusted a ccordingly .

r esources

w ill

T he p ossible r elationships b etween ,

a nd s ocial o rganisation w ithin t wo e conomically d iscrete g roups

i n o ne

s mall g eographical a rea c onstitute a f ascinating n ew f ield o f s tudy .

9 0

TABLES

0 c r , i A

, i e r

• ,: r r n , i

A

A

I

( 8L61

id dT b d

I

A

r n

A

. 4 , I

c r

e . C V O N Ic v , i 0 V

Co

40 / 1

I

9 14 4 1-w S ) g loodsv

il oqbuT(/Pd ) p uoabsaTipuv

( IL6I

' Teoduabbod uo lbuTxxed ) u aburg a u

9

( P9L6I

A

x

C o

4 / 1 / D

r i rl

Co

11 0 4 buTX/Pd )

g oo p fda u l I• • ••

uoqbupgrd ) ( P9L6I i A rg s ,purIg

C D

C o C O e r r 1 r 1 e 1

r 1 r 1

r ,)

C o C o r 1

IA

4 . 1

( *dead u r GAPJ

( P6L ' LL6I

' Aaqou ) a sToqio1

U IA

C O

C o

im P t 4PqaPc [oH )

4 : 1

J aqsouialrd

( P6L61 im eqsqlegad ) u Taquogsbae o lgogg ( q 1P 6L6I

gl e P l isql 8q0U ) p urTTH a a 3 {Tn I G

( LL6I

( L961

r 1

r • • V C V LA Ir l C

A rg q nog

' PeadS 9 s bbrw ) L ud .q .aquog ( 1961 ' MqTwS ) s TaTood

X X X

MAMMAL

REMAINS

FROM ARCHAEOLOGICAL SITES

IN THE

SOUTH-WESTERN CAPE

( 6L61

I n r n r I L A c o X ( N C V

0 5 C )

4 )

5 §

4 111 : r 1 1 11. 5 :

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c s 4

L n

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A

( 9L6I ' c ldTH

A

* . e r r I

•

c v X r 4 N N

I f ) C O

C 4

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( 6L61 ' uogbuT) (led ) p uoibsaTipuv

-

r I

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t 1 U

C O

I s . -

I n

C )

r I

-

U

C1 41 C / '1 41 /

r I

( e9L6I ' uolbuTX Ted )

3 0 0P r I a Ta I n

( P9L6T ' uoqbu p lied ) A pg sj oupT3 ( ' c iaad u T ' Aa c toa ) a nuo a sTozioI ( P6L

r I

" I

r i

f

' LL6I ' mu l le=gagod ) i agsouiaqvd

C 41 / 1

( e6L6T im egsgiegot t ) u Taquogsbialgoqs im P t IslaegoU ) p ueIT3 i exTre g

( LL6I

( L961

1 4

ju eue l l 3n E )

' peadS

r I

A r 's g noH

9sbbPW) b iaqpquog

( 1861 '11 3111 S ) s TaTood

a )

( sheep & g oats )

( q 'e6L6I

rI

t y ,

9 3

r f

c o e n

0 r -

e n

c r ,

( 8L6I

9 M1TWS ) 4200dSV

( 6L6I ' uo lbu P (Xed ) p uoabsaTzpuv ( IL6I ' IaOduabbod 9 U045U p (JPd )

▪

r n

r • -

*

u abupH a g ( P9L6I

45 uP lard ) A pe sgp upTa

N J

( -dead u T ' Aagg i b i ) am° a sTcqacq ,

1 4 N J

( r9L6T

( P6L

'1. 1 0 45 uP (1 r d ) p oo p fdaTo

' LL6I

1 10

N

*

I -

( N N C O i n

1m0 . 1s4iagog )

x aqscuaagrd ( P6L6I ' mu t islaaga en u Taquogsbiacgoqs ( q 'r6L6I

r i

0 1 c v t N

i t *

r n

c o

r 1 e n

Im Pgsglegod ) p urTT3 a axTno l

( 1161 ' uvue .qpnE ) A vg q noH ( L96I ' Peeds 9 s f ibpw ) E t zac muog ( 1861 '14 4 -r wS ) s TaTood

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I TES I N T H E S O UTH-WE STERN C A PE B I RD R E MAINS F R OM A R CHAEOLOG ICAL S

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=

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0 . 3 4 4 . .. . . .

0) 6 4

c u

0.

0

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1 4 0 1 1 1

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6 4 3. I :7 e .-•

•P. . -44 4 0 Or C .) 0 = 44 CO 1 . . . Co 4. eV C CV . e . .. 0. 0 IV C D C . ). .-. . . . . . . .

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F I SH R EM AINS F ROM A RCHAEOLOG ICAL S ITES I N T HE S OUT H-WESTERN C APE

in

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T A BL E I V S H E L L F ISH R E M A IN S F R O M A R C H AE OL OG ICA L S I T E S I NT H E S O UT H -W E STE R N C A PE

R o o ie l s S m it sw in k e l B a y

P e r na p e r n a a s %t o ta l s h ell C h o ro t y t ilu s m e r id io n a l is a s % t o ta l s h e l l P a te l l a s p p . a s %t o ta l s h el l O t h e r s p p . a s %t o ta l s h e ll

P a t e l l a mp u la r i s a s %t o ta l P a te l la P a t e ll a g r a n a tina a s %t o ta l P a te l la P a t e l l a b a rb a ra a s %t o tal P a te lla P a te l l a a r ge nv ill e i a s %t o ta l P a te l la P a t e l l a c o c h le a r a s %t o ta l P a te ll a P a t e l l a o c u lu s a s %t o ta l P a tel la P a te l l a m i n ia ta a s %t o ta l P a te lla P a t e ll a l o n g ico s ta a s %t o ta l P a te l la O t h e r s p p . a s %t o ta l P a tella

H o u t B a y

D u iker E i la n d

S t ofb er g s E l a nd s f o n te in B a y

3 4 1 2 4

5 1 2 3 2 6

1 0 8 6 4

3 7 5 2 1

5 9 2 7 1 4

8 1 7 x x 2 5 2 4 x x 2 6

1 9 4 3 4 4 1 5 2 2 I

3 7 5 1 x x x x

1 8 5 2 2 S 3 6 -

2 3 8 2 4 -

1 2

b

3 4

D i ep k l o of

3 2 ) 5 ) 2 6 6 9

x 3 x 5 5 7 1 . , 3

6 2 3 1 7

? ? ? ? ? ? ? ? ?

x p r e sen t ? i n f or m a t io n n o t a v a ilab le

T h e n u m b er s i nT a b le s I-Vr e f er t oL a te : l o lo c ele f o od w a s te . M o ! ' ; o f t h e s i te s l i sted d o n o z , i na n y c a se , d a t e b a ck f u r th er t h a n t h e m—J Ho l cle n e . E l a rd s B a y C a ve , T o r to ise C a ve a n d A sw .r t a l l h a v e e a r ly H o loce ne d e p o sit s ( a l tho u gt t h e l o w er l a y er s a t A s p o or t a r e u n da ted ) ; b u t t h e se f a u n a l a s se mb la g e s a r e n o t i n c lu de d i nT ae s I-V .

9 7

T A BLE V E D IBLE P L ANT R E MA INS F R OM A R CHAE OLOG ICAL S I TE S I N T H E S O UTH -WE STERN C A PE

1

2

3

R e s tio s e tiger ( s eed e d ib le b u t m a le i n f lore scence s f o un d ) W i lldenow ia s t ria ta ( s eed s a n d s e ed -ca se s ) C a nnamo is s p . ( s eed s ) F i cin ia s p . ( i nflore scences , ? b edd ing ) L e ucadendro n s p . ( s eed s a n d s e ed -ca se s )

+ +

D i o scorea e l ep han tip e s ( b ar k o f t u b er ) E u clea t o m ento sa ( s eed s a n d l e av es )

+ + +

+ +

E u clea l i n ear is ( s eed s a n d l e ave s )

+ +

E u c lea s p . O l ea a f r icana ( s eed s ) R h u s u n du la ta ( l eave s )

+ +

R h u s s p . ( s eeds ) S e ca le c e r ea le ( s tem s a n d e a rs ) ( ? )

+ + + +

P e largo n iu m s p p . ( t uber c a sing s ) B r ab eium s t ella tifo lium ( s eed c a s ing s )

+ +

+

C a rp abro tu s e d ulis ( s tem s ) H o raea s p . ( c o re s a n d c o rm c a sing s ) B a b iana s p . ( c orm s a n d c o rm c a sings )

+ e + + +

+ + + +

+

W a tsonia s p . ( c orm s a n d c o rm c a s ing s ) ? L yc ium s p . ( t horn a n d t w ig ) ? A sparagu s s p . ( t w ig ) G l ad io lu s s p .

+ +

N y land tia s p inosa ( s eed s ) D i o spyros s p , H e xaglo ttis s p .

+ y

+ + + ? +

C o ty ledon p a n icula ta

K e y

+

+ +

+4

A n tholyza p l ica ta O x a l is s p p .

1.D e H a n gen ( P ark ington a n d P o ggenpo e 2 A n dr iesgrond ( R ob ey e t a l . 1 9 78 ) 3 D i epk loof ( P ark ing to n 1 9 76a )

+

.1 7 1 ;

t r a ce

+ .p r esen t + + a b undan t

9 8

+ ,

T ABLE V I

E DIBLE I NDIGENOUS P LANTS O F T HE S OUTH-WESTERN C APE

1 . P LANTS W ITH E DIBLE R OOTS/CORMS/TUBERS.

F amily : A piaceae A nnesorhiza s pp.

( anyswortel)

C hamarea c apensis E . & Z . G lia q ummifera S ond.

F amily :

( gammare ,

( gliwortel,

m oerwortel, d ronkwortel)

A ponocynaceae

P achypodium b ispinosum A .DC.

F amily :

v inkelbol)

( bobbejaankos, k afferkambro)

A raliaceae

C ussonia t hyrsiflora T hunb .

( kiepersol,

k iepersolboom ,

c abbage t ree)

F amily: A sclepiadaceae B rachystelma s pp.

( hottentotsbrood ,

k ambro)

C eropegia s pp. F ockea s pp.

F amily :

e sp. F .

c omaru N .E. B r .

( kambro)

C ampanulaceae

C yphia s pp .

( baroe ) .

P hillips

( 1917) l ists e ight s pecies o f C yphia f rom

t he s outh-western C ape, n amely C . p hyteuma , C . i ncisa , s chechteri ,

F amily:

C . d igitatg . l ,

C .

v olubilis a nd

C . c renata .

( boesmanuintjie)

D ioscoreaceae

D ioscorea e lephantipes

F amily:

z eyheriana ,

C .

C yperaceae

C yperus u sitatus B urch.

F amily:

C .

C . b ulbosa ,

( L 17 . 4e rit . )

E ngl .

G eraniaceae

P elargonium s pp.

9 9

( hottentotsbrood ,

o lifantsvoet)

F amily :

I ridaceae

A ntholyza p licata L .F . A ntholyza r inqens L .

( hanekom)

( rotstert)

A ll B abiana s pp. G ladiolus s pp . H exaglottis s pp. I xia s pp. L apeirousia s pp. M oraea

s pp .

e sp .

M .

f ugax

( de

e dible ,

a f ew a re p oisonous .)

la R oche)

J acq .

( Not all

R omulea s pp. S paraxis s pp. T ritonia s pp. W atsonia s pp. a nd p robably o ther g enera.

F amily:

L iliaceae

A llium d reqeanum K unth. A sparagus s pp.

( ajuin,

( katdoring ,

w ild o nion, w ildeui)

h aakdoring ,

w ild a sparagus)

E riospermum s pp. M assonia d epressa H outt. T ulbaqhia a lliacea L .F.

( suikerkannetjie,

b otterkannetjie)

( wild g arlic)

F amily: N ymphaceae N ymphaea c apensis T hunb.

F amily :

( blouwaterblom ,

b lue w ater l ily)

O xalidaceae

A ll O xalis s pp .

( suring)

F amily : P ortulacaceae A nacampseros s pp.

F amily: R osaceae G rielum h umifusum T hunb.

)p ietsnot,

G rielum q randiflorum ( L .) D ruce

)d uikerwortel

1 00

M oraeas a re

F amily:

T ecophilaceae

g yanella h yacinthoides L .

2 .

( raap,

r aaptol,

l ady's h and)

P LANTS W ITH E DIBLE L EAVES A ND/OR S TEMS.

F amily:

A izoaceae

H ypertelis s alsoloides A dams

( haassuring,

T etra wnia f ruticosa L .

( slaaibos)

S celetium s pp .

s trictum L .

k ougoed ).

F amily:

e sp .

L eaves

B ol.

( kanna,

( pigweed ,

m isbredie).

S ome e xotic s pp.

A raceae

c onflicting r eports a bout t his p lant ,

H oodia s pp.

e sp.

( ghaap ,

T richocaulon s pp.

C .

i t m ay o r m ay n ot b e e dible .)

m ammillaris N .E.

B r.

( aroena)

n gaap)

( ghaap ,

n gaap)

( L.F.)

A it.

( doringslaaibos)

T archochnanthus c amphoratus L .

( kanferhout)

L eaves c hewed a s a n arcotic.

C unoniaceae

C unonia c apensis L . y oung b racts

F amily:

( rooiels ,

b otterboom ) O ily s ubstance a t t he b ases o f

e dible .

G eraniaceae

P elargonium s pp.

F amily:

( There are

A steraceae

D idelta s pinosa

F amily:

varkblom)

A sclepiadaceae

C aralluma s pp.

F amily:

H ottentots-

c hewed a s a n arcotic .

Z antedeschia a ethiopica ( L .) S preng ( arum l ily ,

F amily:

k ougoed ,

A maranthaceae

A maranthus s pp.

F amily:

S .

b raksuring)

G raminae

P hraqmites c ommunis T rin.

( fluitjiesriet,

1 01

S onquasriet)

F amily :

J uncaceae

P rionium s erratum D r@ge

F amily:

L iliaceae

A lbuca

s pp .

L eighton

e sp .

A .

( palmiet)

a ltissima

( Jacq .)

D ryand a nd A .

c anadensis

( L .)

( slymstok )

A llium d regeanum K unth. A sparagus s pp.

( ajuin,

( katdoring ,

w ild o nion, w ildeui)

h aakdoring ,

w ild a sparagus)

B ulbine s pp. T rachyandra s pp.

( hottentotskool)

T ulbaghia a lliacea L .

F amily:

( wild g arlic,

w ildeknoffel)

M esembryanthemaceae

C arpanthea p omeridiana ( L .)

N .E .

C onophytum t runcatellum N .E. B r. D orotheanthus s pp.

B r.

( vetkousie)

( clock p lant)

( Bokbaai v ygies)

M esembryanthemum c rystallinum L . ( brakslaai ,

F amily :

( suring)

P olyqonaceae

E mex a ustralis S teinh. R umex

i ce p lant )

O xalidaceae

A ll O xalis s pp.

F amily:

s outslaai ,

s pp .

( duiweltjie,

( tongblaar ,

C ape s pinach)

r ooisuring ,

v eldsuring)

( European

p lants

i ntroduced b y t he f irst W hite s ettlers .)

F amily :

P olypodiaceae

P teridium a quilinum K uhn .

( bracken f ern )

Y oung u nopened l eaves e dible ,

b ut l arge l eaves p oisonous .

F amily :

P ortulaceae

A nacampseros s pp.

F amily:

T yphaceae

T ypha l atifolia L .

s sp.

c apensis R ohrb.

1 02

( bulrush,

p apkuil)

3 .

P LANTS W ITH E DIBLE F LOWERS O R N ECTAR.

F amily:

A ponogetonaceae

A ponogeton s pp.

F amily:

e sp. A .

d istyachos L .f.

A sclepiadaceae

M icroloma t enuifolium

( L.)

M icroloma s agittatum ( L.)

F amily:

B r.

( kannetjies) ( bokhorinkies)

C rassulaceae ( boesnap)

G raminae

A gropyron d istichum P .

F amily:

S chum. R .

C rassula a lpestris T hunb.

F amily:

( waterblommetjie)

B eauv.

( suikerriet,

L iliaceae

M assonia d epressa H outt.

( suikerkannetjie,

T rachyandra s pp.

f alcata

F amily:

e sp.

( heuningblom,

e sp.

P .

r epens

( L.)

( hottentotskool)

k ruidjie-roer-my-nie)

L .

( soetpop ,

( suikerbossie)

r ooipoppies,

k annie)

T hymelaeaceae

S truthiola c iliata ( L.)

3 .

K unth.

S crophulariaceae

H yobanche s anquinea L .

F amily:

( L.F.)

P roteaceae

P rotea s pp.

F amily:

T .

b otterkannetjie)

M elianthaceae

M elianthus m ajor L .

F amily:

s trandkoring)

L am.

( heuningbossie,

s troopbossie)

P LANTS W ITH E DIBLE S EEDS.

F amily:

A sclepiadaceae

M icroloma s aqittatum ( L.) M icroloma t enuifolium

( L.)

R .

B r.

S chum.

( bokhorinkies) ( kannetjies)

1 03

F amily:

C rassulaceae

C otyledon p aniculata L .f .

F amily:

A ll

b otterbos)

F abaceae

A cacia k arroo H ayne

F amily:

( botterboom,

( acacia,

k areedoring, m imosa).

G um a lso e aten.

M esembryanthemaceae

C arpabrotus

s pp .

e sp .

C .

e dulis

( L . ) N .E .

B r .

( hottentotsvy ,

p erdevy) a nd C . q uadrifidus L . B ol. ( elandsvy).

s uurvy ,

A lso C . a cinaciforme

a nd C . s auerae .

F amily :

P roteaceae

S eeds o f a ll s pp.

p robably e dible, b ut e specially:

B rabejum s tellatifolium L .

( wild almond ,

h ottentotsamandel,

b itter

a lmond) L eucadendron s pp. L eucospermum s pp . b randhoutboom ,

F amily :

L .

c onocarpodendron

( L .) B uek .

( bobbejaanbos,

g oudsbos)

R estionaceae

L arge-seeded C annamois

5 .

e sp .

s pp .

e .g .

v irgata S teud .

W illdenowia s triata T hunb . ( besemsriet ,

( Sonquasriet) a nd

o lifantsriet)

P LANTS W ITH E DIBLE F RUITS.

F amily :

A maryllidaceae

A ll G ethyllis s pp.

F amily :

( koekoemakranka ,

k ukumakranka)

A nacardiaceae

A ll R hus s pp.

F amily :

A steraceae

C hrysanthemoides

m onilifera

T .

N orl.

b ietou)

1 04

( boetabessie,

b rother

b erry ,

F amily :

C elastraceae

C assine p arvifolia S onder ( mountain s affronwood) C assine p eragua L .

( lepelhout,

M aurocenia f rangularia M iller

k ububessie, b astard s affronwood) ( Hottentots c herry)

F amily: E benaceae D iospyros a ustro-africana D e W inter ( kritikom) D iospyros q labra ( L .) D e W inter D iospyros

r amulosa

( E .

( bloubessie, b lue b erry)

M eyer e x A .

D C)

D e W inter ( Namaqua kritikom,

a ppelkoosbessie) D iospyros w hyteana ( Hiern.) F . W hite E uclea l inearis H iern. E uclea r acemosa M urr.

( smalblaar ) ( seeghwarrie)

E uclea t omentosa E . M ey.

F amily :

( L.) V erdc.

j akkalsbos,

c argosang)

( wilde-ertjie)

H ydnoraceae

H ydnora a fricana T hunb.

F amily:

( heuningghwarrie,

F abaceae

D ipamon l ignosus

F amily:

( bladder-nut)

( jakkalskos,

b obbejaankos,

k annip)

I ridaceae

A ll R omulea s pp .

( froetang)

F amily: L auraceae C assytha c iliolata N ees.

F amily: A ll

v rouehaar ,

M esembryanthemaceae

C arpabrotus s pp .

e sp .

C . e dulis

p erdevy ) a nd C . q uadrifidw i L . B ol . C arpanthea p omeridiana ( L .,

F amily :

; L .) N .E . B r .

( elandsvy )

N .E. B r.

( vetkousie)

M oraceae

F icus c ordata T hunb. F icus

d evil's t resses)

( Nama c :u3 f ig)

i licina S onder e x M ic : ,

( laurel f ig)

F amily: O leaceae O lea c apensis L .

( ironwood,

/ sterhout)

1 05

( hotnotsvy,

s uurvy,

O lea e uropaea L .

s ubsp .

a fricana ( Miller ) P .S . G reen ( wild o live ,

o lien-

h out) O lea e xasperata J acq.

F amily :

( coast o live,

b asterolienhout)

P odocarpaceae

P odocarpus

l atifolius

( Thunb .) R . B r .

e x M irb .

( yellowwood ,

o pregte

g eelhout)

F amily:

P olyqalaceae

N ylandtia s pinosa ( L.)

F amily:

F amily:

f .) K urz.

i nerme

( L .f .)

K unth .

b okdrolletjie ,

t urkey-berry )

( sumach ,

p ruimbas)

( melkhout,

m ilkwood)

S crophulariaceae

H alleria l ucida L .

( kinderbossie,

n utseng)

S olanaceae

L ycium f errocissimum M iers.

F amily:

( Cape d ate ,

S apotaceae

S ideroxylon i nerme L .

F amily :

( katdoring)

S antalaceae

C olpoon c ompressum B ergius

F amily:

b okbessie)

R ubiaceae

C anthium

F amily:

( skilpadbessie,

R hamnaceae

S cutia m yrtina ( Burm.

F amily:

D umort.

( bokdoring)

T iliaceae

G rewia

o ccidentalis

F amily :

V itaceae

R hoicissus

t omentosa

L .

( kruisbessie,

( Lam .)

W ild

a nd

b uttonwood)

D rummond

( wild

v ine,

b obbejaantou)

( From P hillips 1 917; a nd S chelpe 1 981;

S mith 1 966 ; L evyns 1 966 ;

P algrave 1 981; K idd 1 983.)

1 06

M ason 1 972; L e R oux

T A B LE V I I A N AT OM ICA L D I F F ERE NCE S B E T WEE N P L ANT S W I T H D I F FE R E NT S U B -T YPE S O F C4 P H OTO S YNT HE S IS

A s p a r ta te f o r mer s

M a la te f o r m er s S in g le K r a n z s h ea th w i th c e n tr ifug al c h lo rop la s t s a r ou nd t h e l a rg er v a scu lar b u nd le s

D o ub l e b u nd le s h ea th w i th v a r iab le c h lo rop la s t p o s it io n ( s e e b e low )

N o c e l l s a r e p r e se n t b e tw ee n t h e m e ta x y lem v e s se l e l e m e n ts a n d t h e K r anz s h ea th c e l l s o f t h e f i r s t o r de r v a sc u l a r b u nd l e s

M e s to n e , o r i n ne r b u nd le s h ea th , p r e s e n t b e tw ee n m e ta xy l em e l em e n t s a n d K r a n z s h ea th c e ll s

W e ll d e v e lo p e d g r a na l a ck in g i nt h e c h lo rop la s t s

W e ll d e ve lop ed g r a na p r e sen t i nt h e c h lo rop la sts

L o w a i to ch o nd r ia l f r eq u en cy

H ig h m i to cho nd r ia l f r eq u en cy

N o p o stil lum ina tio n c a r b on d i o x ide b u r s t

P o st illu m ina t ion c a rb on d i o x id e b u r st o b se rv ed

M A D -m e s u b ty p e C h lo rop la s t s r o u nd i n ne r w a l ls o f K r a nz s h ea th c e l l s M e sop hy ll c o n s ist s o f as i n g le l a y e r o f d i st inc t , r a d ia lly a r ra n ged n a rro w t a b u lar c e l ls

P E P ck s u b typ e C h lo ro p la st s r o u nd o u ter w a l l s o f K r an z s h ea th c e ll s R ad ia l a r r an gem en t i r re gu la r a n d c h lor en ch yma c e l ls l a r g er a n d l e s s e l o n ga ted t h an i n H A D me s u b typ e

( f r om E l l is e t a l . 1 9 8 0 )

1 07

T A BLE V I II 813

CV ALUE S

O F C O LL AGE N F R OM A N IMAL S I NT H E S O UT H -WE STE RN C A PE

( i n °/0

K a ro o p r op er

0)

C o a s tal P l a in

F o ld M o un ta ins

K a roo m a rg in

B u sh ma nl a nd

T o r to ises ( C her sina a n gu la ta ,

U C T 7 8 0

U C T 7 7

U C T 7 2 4

O C T 7 8 2

e x cep t f o r B u shma nland w h ere b o th s p ec imen s a r e P s ammob a te s t e n tor iu s v e r rox ii )

2 3 ,0 ( C P )

2 3 ,9 ( D eH )

2 3 ,3 ( D b )

2 4 ,9

U C T 7 6 2 3 ,1

U C T 7 9 2 2 ,2

( K )

( B jkf )

U C T 7 8 1 2 2 ,8

U C T 7 8 2 2 ,5 ( P a t ) U C T 1 0 75 2 0 ,2 ( E B ) U C T 1 0 76 2 0 ,8 ( E B )

H a re s ( L ep u s c a p ensis )

U C T 7 2 8

U C T 7 3 4

O C T 7 3 8

1 8 ,0 ( D w )

2 1 ,0 ( O b )

1 5 ,1

O C T 7 3 3 2 0 ,0 ( H f )

S t eenbo k ( R ap h iceru s c a mp e str is )

U C T 7 3 5

O C T 7 2 2

O C T 7 2 1

2 1 ,4

1 7 ,4

1 8 ,5

( C P )

( A p )

O C T 0 6 3 2 3 ,5 ( B ok ) O C T 7 3 2 2 1 ,2 ( H f ) O C T 8 7 6 1 9 ,4 ( E B ) O C T 1 0 74 1 9 ,4 ( E B )

1 08

C o a sta l P l a in S p r ingbo k ( A n t idorca s m a r su 2 . 1 2 1ü )

F o ld M o un ta in s

K a ro o m a rg in

K a roo p r op e r

B u shma nl a nd

U C T 8 1 9

U C T 7 2 0

U C T 8 2 2

2 0 ,9 ( C h )

1 9 ,0 ( A p )

1 8 ,4 ( K im b erl ey )

U C T 8 2 0 2 1 ,2 ( C h ) U C T 8 2 1 2 1 ,6 ( C h )

D a s s ie s ( P r o ca v ia c a p en s i s )

U C T 8 7 8

U C T 7 8 3

U C T 7 2 3

2 2 ,1 ( E B )

2 1 ,1 ( A g )

2 1 ,4 ( A p )

U C T 8 7 9 2 0 ,4 ( E B ) U C T 8 3 1 9 ,9 ( E B )

D u n e m o le r a ts ( B a th ye r gu s s u illu s )

U C T 8 7 7 2 2 ,1 ( P e n )

U C T 7 1 7 1 9 ,2 ( D b )

U C T 8 0 2 0 ,3 ( E B ) U C T 1 0 77 1 7 ,8 ( E B )

B a b o o n s ( P ap io u r sinu s )

O C T 8 1 2 0 ,3

O C T 7 1 8 1 8 ,1

( B jkf )

( A p )

U C T 8 2 2 0 ,3 ( B jk f )

•B a tea red f o xe s ( O to cyo n m e g a lo t is )

U C T 8 4 2

O C T 8 4 1

U C T 8 7 5

U C T 8 7 2

1 9 ,6 ( P ik )

1 9 ,2 ( D el )

1 5 ,2 ( A P )

1 5 ,7

O C T 8 7 3 1 4 ,2

1 n9

K e y t o T a b l e V I II E a c h d13 C v a lu e r e f er s t o o n e a n imal : w h er e t h er e a r e t h r e e n u m b er s f o r , s a y , d a ss ie s f r om E l a n d s B a y , e a ch r e f er s t o as e p ara te i n d ividual . L o calit ie s w h e re t h e a n imals w e re c o lle c ted a r e s h ow n o n M a p V ;t h e a b b r evia t io n s u s e d i n T a ble V I I I a r e a s f o llow s : C P P e n P a t C h H f P i k E B D e H B j k f A g A p B o k D b D w

C a p e P o in t N a tur e R e serv e C a p e P e n in su la K o eb er g P a ter no s ter C h urch av en H o p ef ie ld P i k e tb erg E l a nd s B a y D e H a n gen B o on tjie sk loof A n d r ie sgrond A s p oo r t B o k baa i D o o rnb o s D w a r sk e r sb o s

T h e U C T n u mb er g i v en i s t h e l a b o ra to ry a n a ly sis n u mb er f o r t h a t a n im al .

T A BLE I X 813

CS P A C IN G

B E TWEE N D I F F E RE NT T I SSU E S O FT H ES A KE A N IMAL

U C T n o .

( i n

0/••)

B o ne c o lla gen

H e a t

D i ff er en ce

T O R T O ISE S 7 7 7 6 8 3 6 8 3 7 8 3 8 8 3 9

D eH a n gen ( C h e r s in a a n qu la ta ) K o eb e r g ( C . a n g u l a ta ) J o n k e r sh oek n o . 1( C . a n gu la ta ) J o n k e r sho ek n o . 2( C . a n qu la ta ) J o n k e r sho ek n o . 3( C . a n qu la ta ) J o n k e r sho ek n o . 4( C . a n qu la ta )

2 3 ,9 2 3 ,1 2 2 ,9 2 0 ,1 2 1 ,4 2 2 ,7

2 5 ,8 2 5 ,3 2 5 ,1 2 3 ,2 2 3 ,8 2 4 ,0

1 , 9 2 , 2 2 , 2 3 , 1 2 , 4 1 , 3 H e a n = 2 ,2 + 0 ,6 °/ • 0

7 8 1 7 8 2

D eT u in ( P saa n ob a te s t e n to r iu s v e r ro x i i ) C a lv in ia ( P . t e n t o r iu s v e r ro x i i )

2 2 ,8 2 4 ,6 2 5 ,2

2 4 ,4 2 4 ,0 2 4 ,4

1 , 6 0 ,6 0 ,8

2 0 ,9 2 1 ,2 2 1 ,6 1 8 ,4 2 0 ,0 2 0 ,8 1 8 ,9 1 6 ,5

2 4 ,0 2 4 ,3 2 5 ,0 2 1 ,8 2 0 ,8 2 2 ,6 2 1 ,9 1 8 ,5

3 , 1 3 , 1 3 , 4 3 , 4 0 , 8 1 , 8 3 , 0 2 , 0 H e a n= 2 ,6

S P R IN GBOK 8 1 9 6 2 0 8 2 1 8 2 2 8 2 3 • • •

C h u r ch a ven C h u r ch av en Y C h u r ch av en Z K i a b e r ley H u t ch in so n P h il l ip stow n D eA a r K e e ta a n sho op

+ 0 ,9 °/ 1 0

B A T -E A RED F O XE S 8 4 2 8 4 1 8 4 0 8 7 3 8 7 2

P i k e tb er g D eH a ng en N i e uw o ud tv il le B r a n d v le i K e n h a r d t

1 9 ,6 1 9 ,2 2 0 ,8 1 4 ,2 1 5 ,7

2 2 ,0 2 1 ,6 2 2 ,6 1 5 ,4 1 8 ,9

2 , 4 2 , 4 1 , 8 1 , 2 3 , 2 H e an = 2 ,2 + 0 ,7 0 / 0 0

*F r o m V o g el 1 9 78 b .

1 11

T A BLE X 813 CV A L UE S O F M E AT F R OM A N IM AL S W I T H

U C T n o . 7 3 4 7 1 7 2 7 2 8 7 3 3 7 3 2 7 3 5 7 3 1 7 3 7 8 1 9 8 2 0 8 2 1 7 8 7 7 6 7 7

C3 —

B A SED D I ET S I NT H E S O UT HN E ST ERN C A PE

A n ima l

L o cal ity

L e p u s c a p e n s is ( h a re ) L e p u s c a p en s is ( h a r e ) L e p u s c a p e n s is ( h a r e ) L e p u s c a p en s is ( h a re ) L e pu s c a p e n s is ( h a r e ) R a p h icer u s c a mp e s tr is ( s te enb ok ) R a p h iceru s c a mp e str is ( s te enb o k ) S y lv icapra g r im m ia ( d u ike r ) T a uro tragu s o r y x ( e land ) A n tido r ca s m a r sup ia l is X ( s p r ingb o k ) A n t ido r ca s n a r sup ia l is Y ( s p r ingb o k ) A n tido r ca s n a r sup ia l is Z ( s p r ingb o k ) F r anco l inu s c a pe n s is ( f r a n co l in ) C h er s ina a n g u la ta ( a ng u la te t o r to ise ) C h er sina a n gu la ta ( a ng u la te t o r to i se )

413

D o o rnb o s C e d arb er g C e da rb er g D w a r sk er sb o s H o p ef ie ld H o p ef ie l d C a pe P o in t H o p ef ie l d C h ur cha v e n C h ur ch av e n C h ur chav e n C h u rchav e n Y z erfo n te in K o eb erg D e H a ng en

C (Veo)

2 3 ,0 2 2 ,2 2 2 ,0 2 3 ,6 2 2 ,6 2 3 ,7 2 4 ,3 2 3 ,5 2 3 ,7 2 4 ,0 2 4 ,3 2 4 .9 2 0 ,9 2 5 ,3 2 5 ,8 M e an 2 3 ,6 +1 , 3 °/ ..

T A BL E X I C H AN GE I N 813 CV A L UE S O FA N IMAL T I S SUE S W I T H T R O P H IC L EVEL

U C T n o .

A n im a l

B o n e c o l la g en

( i n °/ 0 0 )

H e a t

D i f fe r e n ce

7 3 3 7 3 2 7 3 1

H e rb ivo res H a r e ( L epu s c a p en s is ) S t e e nbo k ( R ap h icer u s c a mp e s tr i s ) D u ik er ( S y lv icapr a R r imm ia )

2 0 ,0 2 1 ,2 2 0 ,7

2 2 ,6 2 3 ,7 2 3 ,5

2 , 6 2 , 5 2 , 8

7 3 0 7 2 9

C a r n ivor es L y nx ( F e l is c a ra ca l ) J a ck a l ( C an is m e so mela s )

1 9 ,2 1 7 ,9

2 1 ,8 1 9 ,8

2 , 6 1 , 9

A l l a n im a l s c o lle c ted a t H o p ef ie ld .

1 12

T A BL E X I I 413

U C Tn o 7 5 6 V a r io u s

1 0 3 8 4 7 5 7 7 5 8 7 5 9 7 6 1 7 6 0 • • 8 5 8 6 8 7 8 9 8 9 0 1 0 4 7 1 0 4 6 1 0 45

C

V A L UE S O F T H E E D IBLE P A R T S O F P L A NT S

L o cality

P l a n t M o r a ea f u ga x c o r m s ( n = 3 ) W a t son ia p y r a m id a ta c o rm s ( N ew c o r m s c o l le c ted e v e r y m o n th o v er ap e r iod o f o n e y e a r . ) N y la n d tia s p in o sa b e r r ie s C a rp ab r o tu s s p . s e ed s a n d s a p G r ie ln h u n if u su s r o o ts ( n = 5 ) P r ion iu m s e r r a tu m s t em D i o scor ea e l ep h a n tip es t u b e r F o c k ea c o m a ru t u b er ( n = 1 ) C a ra l lu ma m a n s il lar i s s t em ( n = 1 ) C a r a lln a m a s t il lar is H y p er tel is s a l so l o id e s l e av e s C y a n e lla h y a c in t h o id e s c o rm s ( n = 3 ) A l l iu m d r e gea n u m b u lb s ( n = 12 ) A p o n o ge to n d i sty a cho s f l owe r s H y d no r a a f r ica n a f r u it ( n = 1 ) H o o d ia s p . s t em ( n = 1 ) C y p h ia d i g ita ta s u b sp .d ig ita ta t u b er ( n = 1 ) C o lp oo n c o mpr e s su m b e rr ie s O x a l is s p p . c o r m s

61 3 CO h 0 )

E l and s B a y P e n in sula

2 6 ,7 2 3 ,2 t o 2 9 ,2

E l an d s B a y E l a n d s B a y E l an d s B a y B o on tjie sk lo of D o or nb o s D o o r nb o s D o o r nb o s

E l a nd s B a y D o or nb o s E l and s B a y N a maq ua la nd D o or nb o s A n d r ie sgro nd

2 2 ,3 2 3 ,2 2 6 ,3 2 7 ,0 2 8 ,2 2 4 ,4 1 0 ,9 1 4 ,4 2 3 ,7 2 6 ,3 2 4 ,7 2 6 ,9 1 1 ,4 1 1 ,9 2 3 ,0

P e n in su la E l a nd s B a y

2 6 ,2 2 6 ,4 M e a n f o r C3 v a lu e s o n ly : 2 5 ,4+ 1 .8 0/ „

*F r o m R o b er t so n K a r o o ( M o o ne y e t a l . 1 9 77 )

1 13

T A BL E X I I I 613 CV A LUE S O F M A R INE A N IMA L S ( a l l t e a t )

U C Tn o . 0 9 5 0 9 7 1 0 13 1 0 14 1 0 15 1 0 16 1 0 17 1 0 18 5 1 9 5 8 5 0 9 6 1 0 2 0 1 0 2 1 1 0 2 2 1 0 2 3 1 0 19 7 4 2 7 6 6 7 6 9 5 8 6 7 4 0 7 6 7 7 6 8 7 4 3 7 0 7 4 1 8 1 8 7 8 6 5 1 7

A n imal

L o calil n dd a te ya

C h o rom y t ilu s n e r id io na l i s ( n = 10 ) C .m e r id iona l is ( n = 3 ) C . s e r id io na l is C .m e r id iona l is C . m e r id io na l is ( n = 7 ) C .m e r id io na l is ( n = 2 ) C . a e r id iona l is ( n =8 ) C .a e r id iona l is ( n =9 ) A u la co m aa t er B u r nup e na s p . D o na x s e rr a ( n = 16 ) P a te l la g r a na t ina P a te lla g r anu lar is P a te lla a r g env ille i P a tella c o ch lear H a l io t is m i d a e ( n = 1 ) L i za r a mad a ( n = 1 ) L i tho g na thu s l i tho g na th u s ( n = 1 ) P a chy m e top o n b l o ch ii ( n = 1 ) P a chy s e top o n b l och ii ( n = 1 ) S p h en iscu s d e mu r su s ( n = 1 ) P h ala cro co ra x c a p e n s is ( n = 1 ) H o ru s c a p en s is ( n = 1 ) D i oned ea m e la nop hr is ( n = 1 ) A r c to c ep ha lu s p u s il lu s ( n = 1 ) A r c toc eph a lu s p u s illu s ( n = 1 ) J a su s l a la nd ii ( n = 1 ) J a su s l a la nd ii J a su s l a la nd i i ( n = 1 )

E l a nd s B a y 1 / 2/ 76 E l a nd s B a y 2 4 / 4/ 76 E l a nd s B a y 1 / 1 1/ 8 1 E l a nd s B a y 1 / 3/ 82 E l a nd s B a y 2 4 / 5/ 82 E l a nd s B a y 1 0 / 7/8 2 E l a nd s B a y 1 5 / 7/8 2 E l a nd s B a y 1 2 / 9/8 2 O u d ek r a a l O u d ek ra a l E l a nd s B a y 2 7 /4/ 76 E l a nd s B a y 1 / 11/8 1 E l a nd s B a y 1 / 11/ 8 1 E l a nd s B a y 1 / 11/ 8 1 E l a nd s B a y 1 / 1 1/8 1 P a ter n o s te r E l a nd s B a y K a lk B a y E l a nd s B a y O u d ekr a a l S a lda n h a B a y E l a nd s B a y Y s terf o n te in H o u t B a y P a tern o s te r R o u t B a y P a tern o s ter E l a nd s B a y O u d ek ra a l

413

C ('/ • 0 )

1 9 ,4 1 8 ,1 1 7 ,4 1 6 ,2 1 5 ,9 1 6 ,4 1 6 ,4 1 6 ,2 1 6 ,7 1 4 ,8 1 7 ,2 1 5 ,2 1 3 ,6 1 2 ,3 1 2 ,8 1 6 ,8 1 5 ,8 1 4 ,9 1 6 ,5 1 5 ,9 1 5 ,1 1 6 ,3 1 5 ,0 1 6 ,4 1 4 ,3 1 4 ,6 1 4 ,0 1 4 ,6 1 3 ,1 M e an = 1 5 ,6 +1 , 6

7 8 8 7 8 8

W h a l e ( s p e c ie s u n k no w n ) c o llag en S a me w h a le c o llagen a f ter d e -fa t t ing

1 14

2 0 ,2 1 2 ,5

0/ ,0

T A BL E X I V 413 CV A L UE S O F M A R IN E A L GAE

L o ca l ity

U C T n o .

413

C (°/ oo)

D I V IS IO N : R H OD O PH YT A ( r ed w e e d s )

7 9 7 7 9 6

C L A SS : F l o r id eoph y cea e O R D E R : C r yp to n em ia le s F A M IL Y :K a llym en ia c ea e T h a m no p h y llis d i sc ig era K a l ly m en ia a g a rdh i i

P a ter no s ter P a ter no ster

3 4 ,5 3 3 ,0

8 1 5 8 0 7 1 0 9 0 5 2 0

F A M IL Y : C r yp to n es ia cea e A e o d e s o r b ito sa P a ch y t e n ia c o r nea P a chy s e n ia c a r no sa P a chy m e n ia s p .

P a terno ster P a terno ster O u d ekraa l O u dek ra a l

1 4 ,6 2 3 ,7 1 7 ,8 2 1 ,9

7 9 4 8 0 0 8 0 2

O R D E R : C e ra m ia le s F A M IL Y :D e l es ser iac ea e B o tryo ca r p a p r o l if er a H y t e n e na v e no sa N e u ro g lo s sum b i nd e r ia num

P a ter no ste r P a ter no ster P a ter no ste r

3 2 ,8 3 2 ,4 3 2 ,6

8 2 9 8 2 6 8 3 1

F A M IL Y : C e ram ia c Pae C a rp o b lep har i s f l a c c id a C e ra m iu m p l an um C e n tr o c er a s c l av ula tus

P a terno ster P a ter no st er E l an d s B e y

1 9 ,9 2 1 ,2 1 7 .8

7 9 8 8 1 0

O R D E R : R h o dyn en i -l e s F A M IL Y : R h o dy s e r a c ea e E p y t en ia o b tu sa R h o dy n en ia s p .

P a ter no ste r P a terno ste r

3 1 ,3 2 0 ,1

7 9 5 7 9 5 8 2 5 1 0 88 8 0 9 5 1 8

O R DE R : G i ga rt in i e s F A M IL Y : G i gar t ir z ea e G i gar t in a r a d u l i G i g a r t in a r a d u la G i gar t ina b r a c te n ta G i gar t in a b r a c teea G i ga r t ina s p . G i ga r t in a s p .

P a te rno ster P a ter no s ter P a terno s ter O u dek raa l P a ter no ste r O u d ek ra a l

2 6 ,1 2 4 ,8 2 2 ,3 1 6 ,5 2 6 ,1 2 2 ,0

8 2 4 8 1 3 8 1 3

F A M IL Y : P l o cam ia eea e P l o ca m ium r i g idu e P l o ca m iu m c o r a ll t: n iza P l o ca n iu m c o r a l l eT h iz a

P a ter no s ter P a ter no st1 7 : P a ter no st e r :

3 2 ,6 3 3 ,3 3 3 ,9

8 2 7

F A M IL Y : S p haeroco 'za ceae H e r ing ia s p .

P a ter no ste

2 6 ,4

1 15

U C T n o .

L o c a l ity

413

C

( ° / 4 s )

8 1 7 8 0 6 8 0 6

F A M IL Y : S a r co co d ia c eae T r ema to carp u s a f f in is T r em a to carpu s f r ag il is T r em a tocarp u s f r a g il is

P a te r n o ster P a te r no s ter P a te r no s ter

1 9 ,4 2 1 ,3 1 8 ,0

8 0 1

F A M IL Y : N e aa s to ma c ea e N e ma stona l a n c eo la ta

P a te r no ster

3 0 ,4

8 1 2 8 3 2 0 9 3

O R D E R : N e ta lia les F A M IL Y : G e l id ia c ea e G e l id iun v e r sico lo r G e l id iva p r i sto id e s G e l id iu m p r i sto id e s

P a ter no s ter F r a n sk r a a l P e n i n su la

2 1 ,0 1 7 ,5 1 4 ,5

8 1 6 8 3 5 0 9 2

C L A SS : B a ng iop h yc ea e O R DE R : B a ng ia le s F A M IL Y : B a ng ia c ea e P o rp hy ra c a p e n si s P o rp h yra c a p en s is P o rp h y ra c a p e n s is

P a te rn o s te r E l a n d s B a y P e n in su la

1 4 ,2 1 8 ,2 1 4 ,9

7 4 7 4 5 2 1

O R D E R : L a m ina r ia le s F A M IL Y :La m ina r ia c ea e L a m ina r ia p a l l ida L a m inar ia p a ll ida L a a inar ia p a l l id a

P a te r no s ter P a te r no s te r O u d e k ra a l

2 0 ,3 ( s t ip e ) 1 9 ,0 ( f ro nd s ) 2 0 ,9

7 5 7 5 5 2 2

F A M ILY : A l ar ia c ea e E c k lo n ia a x im a E c k lo n ia m a x i l la E c k lon ia a x ima

P a te r no ster P a te r no s ter O u d e k ra a l

1 5 ,2 ( s t ip e ) 1 4 ,1 ( f r on d s ) 1 2 ,0

1 0 8 9

O R DE R : F u ca l e s F A M IL Y : F u ca ceae A x illar ie lla c o n s tr ic ta

O u d ek ra a l

1 7 ,8

8 3 3 8 3 3

F A M IL Y : C y sto se ir a ce a e B i fur ca r iop sis c a p e n s is B i fu r ca r iop s is c a p e n s is

F r an skr a a l F r an s kr a a l

1 4 ,5 1 4 ,9

8 2 8

F A M IL Y : S a r ga s sac ea e S a r ga ssua l o n g ifo l iu m

P a te r no s ter

1 7 ,7

8 0 8

O R DE R : D e saa re s t ia le s F A M IL Y : D e sm a r e st ia c ea e D e sa ar est ia f i r aa

P a te r no s ter

2 3 ,0

0 4

O R D E R : E c to carp a l e s F A M IL Y : S p erm a to ch n a c ea e C h ord a r iop s is c a p e n si s

P a te r n o s te r

1 0 ,6

D I V IS ION : P H A E OP H YT A ( b r ow n w e ed s )

1 16

O C T n o .

8 1 1

L o ca l ity F A M IL Y : S p la chn id ia cea e S p la ch n id iu n r u go su m

813

C (Voo)

P a ter no ster

1 0 ,8

O R D E R : U l o tr icha le s F A M IL Y : U l va c ea e E n te rot orp h a s p . U l va c a p en s is U l va s p . U l va s p .

P a tern o s ter O u d ek raa l E l and s B a y P e n in su la

1 1 ,2 1 6 ,0 1 0 ,2 1 3 ,8

O R D E R : C l ad op h o ra le s F A M IL Y : C l ado p hora cea e C l ad op h ora s p . C l a d op h o ra s p .

E l a nd s B a y E l a nd s B a y

1 1 ,7 1 4 ,0

O R D E R : S i p ho na le s F A M IL Y : C o d ia ceae C o d ius f r a g il e C o d iu n f r a g ile C o d ium p r o stra tue C o d ium p r o str a tun

P a ter no ster O u d ek raa l P a ter no ster O u d ek raa l

1 8 ,6 1 6 ,0 1 7 ,3 1 0 ,3

D I V IS IO N : C H L O ROP H YTA ( g re en w e ed s )

7 9 9 1 0 85 8 3 0 0 9 4

8 3 4 8 3 4

8 0 3 1 0 8 7 8 1 4 1 0 86

1 17

T A BL E X V 413 CV A L UE S O F M A R INE A L GAE F R O M D E E P A N D S H A LL O W W A T ER A T B O UT B A Y

S p ec ie s

C A PE P E N IN SUL A

1 3

Cv a lu e i nd e ep w a ter ( c a . 2 5 . )

1 3

6C v a lu e i n s h a l low w a ter ( 1 -3 . )

E c k lo n ia m a x ima ( f ro nd s )

2 3 ,5 ( U CT 9 7 7 )

1 6 ,3 ( U CT 9 7 6 )

L a n ina r ia p a l lid a ( f ro nd s )

2 2 ,7 ( U CT 9 8 9 )

1 7 ,0 ( U CT 9 8 8 )

P a c hym en ia c a rno sa

1 8 ,0 ( U CT 9 7 9 )

1 4 ,0 ( U CT 9 7 8 )

P a chy me n ia c o rnea

3 4 ,2 ( U CT 9 8 1 )

2 3 ,0 ( U C T 9 8 0 )

N e uro g lo s su m b i nd er ia nua

3 5 ,0 ( U CT 9 8 3 )

3 1 ,5 ( U CT 9 8 2 )

H y m e nen a v e no sa

3 4 ,7 ( U C T 9 8 5 )

3 3 ,2 ( U CT 9 8 4 )

B o try o ca rp a p r o l if e ra

3 4 ,2 ( U CT 9 8 7 )

3 2 ,0 ( U CT 9 8 6 )

1 18

T A BLE X V I P H YS IO L O G IC AL M E A SURE ME NT S O N G R E AT B A R R IER R E E F S E AVE ED S

S p e c ies

C h lo r op h y ta E n terom o rp h a f l exu o sa H a l im ed a m a cro lob a H a l in ed a o p un tia H a l is ed a c y lind ra c ea C h lor o d e sm is f a s tig ia ta A v r a inv illea e r e c ta B o o d lea c o mp o s ita P h a eoph y ta S a r ga ssu m s p . P a d ina t e nu is T u rb ina r ia o r na ta R h o dop h y ta L a ur en c ia s p . H a l m e n ia d u r v illa ei

T o ta l C

1 O r ga n ic C

2 1 ,2 1 4 ,8 1 4 ,4 9 ,8 1 9 ,5 -

2 1 ,3 1 5 ,8 2 1 ,3 1 5 ,3 2 5 ,8 1 8 ,2 -

9 6 4 6 3 7 1 5 4x 1 0 8 6 9 2 8

1 4 ,1 1 1 ,4 9 ,2

1 1 ,3

1 8 5 3

1 1 ,0 3 2 ,0 3 2 ,4

1 5 ,4

2

5 8 5

6

4 2 3

x P E P c a rb o xy la se p r e sen t 1

1 63C i n 6/0. ( f r o m B l a c k a n d B e nd e r 1 9 76 ) . 2 N A D PH m a la te d e h y d ro ge na se i nm i cr omo l e p e r m i n . p e r a g . c h l or op hy l l ( f rom T i n g 1 9 76 ). 3 c a rb on d i o x id e e q u il ib r iu m p o in t i nm g . p e r l i tr e ( f rom T o lb er t a n d G a r ey 1 9 76 ).

1 19

T A BL E X V I I 613

C

V A L U E S O F M A R INE O R GA N ISM S A TD I F FE RE NT T R OP H IC L E VE L S

S c av e ng er s U C T 5 1 7

C r ay f ish

( J a su s ) : 1 3 ,1 ° /o

U C T 5 8 6 F i sh ( P a chy m e top o n ) : 1 5 ,9 o /o 0 U C T 5 1 5 S t a r f ish ( H en r ic ia ) : 1 3 ,3 0/0 U C T 5 9 0 S t a rf ish ( H a r tha ster ia s ) : 1 3 ,5 Yo e

F i lter -Fe ed e r s U C T 5 8 3 H y s id s : 1 6 ,5 0/0 U C T 5 1 9 M u s se ls ( A u la com ya ) : 1 6 ,7 0/0 4 U C T 5 1 6 S e a cu cu mb er ( P en tra c ta ) : 1 4 ,3 040 U C T 5 8 9 R e d -b a it ( P y u r a ) : 1 5 ,6 ° /0

P r ima ry P r od u c er s A l g ae U C T 5 2 0 P a ch ya e n ia : 2 1 ,9 O /0 U C T 5 1 8 G i ga r t in a : 2 2 ,0 0/0 U C T 5 2 1 L a m inar ia : 2 0 ,9 0/ „ U C T 5 2 2 E c k lo n ia : 1 2 ,0 0/0 P h y top la nk to n a n d z o op la nk ton U C T 5 8 1 : 1 9 ,8 U C T 5 8 2 : 1 7 ,9 O /0

A l l o r gan ism s f r om t h e O u d ek raa l k e lp b e d .

1 20

T A BLE X V III i 51 3

CV A LUE S O F M I SCELLANE OUS I N D IGENOU S F O OD S F R O M T H E S O UTHN E STE RN C A PE

U C T n o .

L o ca l ity

7 1 6

R o ck p i geo n e g g

8 8 7 2 6

H o ney C l anw ill ia l y e llowf ish ( B arb us c a p ensis ) m e a t ( n = 1 )

7 2 5

B a rbu s a n dr ew i m e at ( n = 1 )

413

C (0 4 0

B o on tlie sk lo of

2 2 ,8

H o p ef ield D o om nR i ver

2 3 ,7 1 9 ,0

H e x R i ver

2 2 ,6

T A BLE X I X 413 C V A LU ES O F A R CHAEOLOG ICAL A N IM ALS F R OM E L AND S B A Y C A VE

81 3 C (Voo)

U C T n o .

1 0 /8 1 0 79

L a yer 6 G a K 4335 1 20+85 B P ( t op o f l a yer ) G a K 4337 1 5 20+80 B P ( b o t tom o f l a yer ) S t eenb ok H a re

1 0 8 0 1 0 8 1

L a yer 9 G a K 4339 2 9 5 0+ 115 B P ( b o ttom o f l a yer ) S t eenbok S t eenb ok

1 7 ,3 1 8 ,4

1 0 82

L a yer 1 2 P t a-686 9 6 00+9 0 B P S t eenb ok

1 9 ,4

1 0 8 3

S t eenbok

1 8 ,6

1 0 84

H a re

2 0 ,7

1 21

1 8 ,5 2 2 ,6

T A BLE X 413

C

V A LU E S O F D A T ED H U M AN S K EL E T ON S F R OM E L AND S B A Y A N D T H E O L IF ANT S R I VE R V A L L E Y .

S K E LE T AL R E G IST ER N U MBER

R A D IO CAR BO N D A T E B . P .

413

C

( Vo )

E L AN D S B A Y U C T 2 4

O x A 45 5 P t a 17 5 4 O x A 4 77

2 4 0 0+ 10 0 3 8 357 5 0 4 0 50+ 10 0

1 3 ,9 1 4 ,0 1 5 ,9

P t a 18 29 P t a 3086 O x A 4 56 O x A 4 78

8 0 00+9 5 9 7 5 0+ 100 9 8 007 16 0 1 0 8 6 0+ 18 0

1 3 ,5 1 2 ,3 1 7 ,4 1 2 ,5

2 5 0+9 0 2 7 07 i00 4 0 +8 0

1 6 ,1 1 8 ,1 1 6 ,0

O L IF AN T S R I VE R V A LL EY S A M AP 2 7 S A M -AP 2 6 S A M AP 4 6 6 0

O x A 45 2 O x A 458 O x A 4 5 4

U C T 3 1 S A M -AP 1 4 49

P t a 38 69 O x A 45 3

2 1 00+ 7 0 2 30+ 100

1 8 ,4 1 7 ,3

( W ye g a ng )

U C T 3 4

P t a 164 2 O x A 45 7

3 5 40+6 0 3 8 5 0+ 8 0

1 9 ,0 1 9 ,0

( K l ip f on te inra n d ) ( A n dr ie sg ro nd )

N . B . P t a ( P r e to r ia ) d a te s a r e c o rr ec ted f o r 41 3C , O x A ( O xf or d r a d io c ar bo n a c c e le ra to r ) d a te s a r e n o t . S k e le ta l r e g is ter n u mb er s a r e t h e a c ce s s io n n u mb er s o f s k e le ton s i nt h e p h y s ica l a n th rop o lo g ica l c o l le c tio n s o f t h e U n iv er s i ty o f C a p e T o w n M e d ica l S c hoo l ( U CT n u m b e r s ) o r t h e S o u th A f r ica n M u seu m ( S AM AP n u mb er s ). U n nu mb e red s k e le to n s a r e a w a i ti n g a c c e ss io n i nt h e f o r m er i n s ti tu t io n . U C T 3 1 h a s p r ev io u s ly b e e n r e p o r ted ( S e a ly a n d v a n d e r M e rw e 1 9 8 5 ) a s b e in g s o m e 7 0 0 0 y e a r s o l d , b u t h a s j u st b e en r e -d a ted a f ter h a v ing b e en f o u nd t o b e c o n tam ina ted w i th af o ss il c a rb o n -co n ta in ing m a ter ia l .

1 22

T A BLE X I 413 CV A L UE S O F O T H E R D A T ED H U MA N S K E L ET O N S F R O M T H E S O UTH -WE STE R N C A P E C O A ST .

S K E LE TAL R E G ISTER N U MBE R

L O CA L IT Y

U C T 6 0 S A M -AP 5 0 8 3 S A M AP 1 4 4 3 U C T 1 6 2 U C T 1 2

R A D IO CAR B ON D A T E B . P .

S a lda n ha Y s terf on te in G o r do n 's B a y Y s terf o n te in D a r l in g

P t a 20 05 P t a -9 26 P t a 2 309 P t a -9 29 P t a 2 003

613 C ( °/ 0 0 )

9 5 5 +5 0 1 4 9 0+ 55 2 0 5 0+ 5 0 2 8 8 0+ 50 4 4 5; 5 0

1 4 ,6 1 4 ,5 1 1 ,8 1 1 ,5 1 1 ,2

N o te : A d d it iona l 413 Cv a lu e s a r e a v a ilab le f o r 1 7 a r ch a eo lo g ica l c o a sta l s k e le t o n s f r om t h is a r ea ( R aw l in so n 1 N 2 ). T h e se a r e u n d a t ed f i nd s w i tho u t a s so c ia ted c u l tu ra l m a ter ia l . T h e ir m e a n 4 Cv a lu e +o n e s t a nd ar d d e v ia t io n i s 1 4 ,5 + 1 ,7 0/0 ( r a n g e f r om 1 1 t o 1 7 ,7 ' V" ) .

T A BL E X II M O N T H S I N W H ICH J U VE N IL E D A SS IE S F R OM D E H A N GE N W E RE K I L LED

S ept.

Oct.

Nov.

D ec.

Jan.

F eb.

M ar.

Apr.

May

June

July

Aug.

2 7