After Eden: Behavioral Ecology of Early Food Production in the Near East and North Africa 9780860545057, 9781407346694

Table of contents :
Front Cover
Copyright
Dedication
Table of Contents
List of Tables
List of Figures
Preface
Introduction
CHAPTER 1: THE EVOLUTION AND ADOPTION OF FOOD PRODUCTION STRATEGIES: A THEORETICAL FOCUS
CHAPTER 2: ESTABLISHING COSTS AND BENEFITS FOR EARLY PASTORAL FOOD PRODUCTION: ETHNOGRAPHIC AND HISTORIC LABOR AND PRODUCTIVITY DATA AS RELATED TO THE PREHISTORIC PAST
CHAPTER 3: ESTABLISHING COSTS AND BENEFITS FOR THE EARLY CULTIVATION OF CEREALS: ETHNOGRAPHIC AND HISTORIC LABOR AND PRODUCTIVITY DATA AS RELATED TO THE PREHISTORIC PAST
CHAPTER 4: COMPARATIVE ECOLOGY OF THE RELEVANT ANIMAL AND PLANT SPECIES
CHAPTER 5: RELATIVE ENERGETICS OF PASTORAL STRATEGIES INVOLVING CAMELS, CATTLE, SHEEP AND GOATS
CHAPTER 6: RELATIVE ENERGETICS OF EARLY WHEAT AND BARLEY STRATEGIES
CHAPTER 7: ENERGETIC EXPECTATIONS AND THE ARCHAEOLOGICAL RECORD
APPENDIX A: DROMEDARY CAMEL POPULATION DYNAMICS AND SLAUGHTER OFFTAKE FROM A MODEL HERD OVER A ONE YEAR PERIOD
APPENDIX B: CALORIC PRODUCTIVITY OF THE MODEL CAMEL HERD OVER A ONE YEAR PERIOD
APPENDIX C: AGE SPECIFIC GROWTH RESULTS FOR THE FEMALE COMPONENT OF A MODEL CAMEL HERD OVER A FORTY YEAR PERIOD BASED UPON THE DATA PRESENTED IN APPENDIX A
APPENDIX D: AGE SPECIFIC GROWTH RESULTS FOR THE IMMATURE MALE COMPONENT OF A MODEL CAMEL HERD OVER A FORTY YEAR PERIOD BASED UPON THE DATA PRESENTED IN APPENDIX A
REFERENCES CITED
INDEX

Citation preview

After Eden The Behavioral Ecology of Early Food Production in the Near East and North Africa

Kenneth W. Russell

BAR International Series 3 91 1988

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 S391,1988 'After Eden' © Kenneth W Russell, 1988

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 9780860545057 paperback ISBN 9781407346694 e-book DOI https://doi.org/10.30861/9780860545057 A catalogue record for this book is available from the British Library This book is available at www.barpublishing.com

T o m y m o the r , L o is , a nd my u nc le Wayne I n memo r ium

C u rsed b e t he g round b ecause o fy ou B yt o i l s ha l ly ou e a to fi t A l lt h ed ays o fy ou rl i f e G enes is 3 . 17

CONTENTS

L IST OF T ABLES L IST OF F IGURES

i x

P REFACE

x i

I NTRODUCT ION

1

CHAPTER 1 .

THE E VOLUT ION AND ADOPT ION OF FOOD P RODUCT ION S TRATEG IES : AT HEORET ICAL FOCUS I n t roduc t ion T he P o ten t ia lo fa n E vo lu t iona ry P e rspec t ive B as ic E ne rge t ic Mode ls Why D id H umans B ecome I n vo lved i nC oevo lu t iona ry R e la t ionsh ips w i th P lan ts a nd A n ima ls : A n I s sue W i thou t R ea l Mean ing?

2 .

E STABL ISH ING COSTS AND B ENEF ITS FOR E ARLY P ASTORAL FOOD P RODUCT ION : E THNOGRAPH IC AND H ISTOR IC L ABOR AND PRODUCT IV ITY DATA AS R ELATED TO T HE P REH ISTOR IC P AST I n t roduc t ion P as to ra l L abo rC os ts i nt he E thnog raph ic P resen t a nd t he P reh is to r ic P as t O ve r v iew H e rd ing C os ts E ncampmen tC os ts B u tcher ing C os ts M i lk ing C os ts P as to ra l P roduc t iv i ty i nt he E thnog raph ic P resen t a nd t he P reh is to r ic P as t G ene ra lC ons ide ra t ions B ehav io r Mo rpho logy P hys io logy A du l t H ypo lac tas ia : S ign i f ican t i nt he E vo lu t ion a nd E ne rge t ics o fE a r ly P as to ra lS t ra teg ies? AC ross -Cu l tu ra l Mechan ism f o rt he I n i t ia l C ap tu re a nd C on t ro l o fW i ld H e rds S umma ry

3 .

E STABL ISH ING COSTS AND B ENEF ITS FOR T HE E ARLY CULT IVAT ION OF CEREALS : E THNOGRAPH IC AND H ISTOR IC L ABOR AND P RODUCT IV ITY D ATA AS RELATED TO T HE P REH ISTOR IC P AST I n t roduc t ion C e rea l C u l t iva t ion L abo rC os ts i nt he E thnog raph ic P resen t a nd t h e P reh is to r ic P as t O ve rv iew S o i l P repa ra t ion

7 7 8 10 13

1 5 15 16 16 1 6 17 1 8 18 1 9 19 19 2 2 2 4 2 9 3 3 3 4

3 7 3 7 3 7 3 7 3 8

H a rves t ing T h resh ing S to rage M i l l ing a nd G r ind ing A nnua l L abo ri nE a r ly I r iga t ion S t ra teg ies C e rea l P roduc t iv i ty i nt he E thnog raph ic P resen t a nd t he P reh is to r ic P as t G ene ra lC ons ide ra t ions T he G ram ineae a nd C e rea l S pec ies S ub jec tt oE a r ly H usband ry AF ocus o nA nnua ls a s Opposed t oP e renn ia ls T he E vo lu t ion o fA nnua l S pec ies f r om P e renn ia l P rogen i to rs A nnua ls a nd P redom inan t S e l f -Po l l ina t ion P heno typ ic P las t ic i ty a nd P roduc t ive P o ten t ia l : T he E v idence f o r Whea t T he N a tu re o fP reh is to r ic a nd H is to r ic S eed S e lec t ion a nd t he H is to ry o fE a r ly H yb r id iza t ion E for ts A nc ien ta nd Mode rn C a lo r ic V a lues S umma ry 4 .

COMPARAT IVE E COLOGY OF T HE RELEVANT AN IMAL AND P LANT S PEC IES I n t roduc t ion S ome C ompa ra t ive A t t r ibu tes o fC ame ls , C a t t le , S heep a n d Goa ts I n t roduc t ion Ma in ta in ing T he rma lS tab i l i ty Ma in ta in ing Wa te rB a lance F eed ing E co logy T he U n ique F ea tu res o fC ame l L ocomo t ion C ompa ra t ive H e rd G row th F ac to rs I n t roduc t ion C ompa ra t ive R ep roduc t ion C ompa ra t ive Mo r ta l i ty E co logy , D is t r ibu t ion a nd E a r ly E xp lo i ta t ion o fW i ld Whea ts a nd B a r leys I n t roduc t ion Mode rn E co logy a nd D i s t r ibu t ion P e rspec t ives o n t he E a r ly D is t r ibu t ion a nd E xp lo i ta t ion o f Whea ts a nd B a r leys E co logy o fC u l t iva ted Whea ts a nd B a r leys I n t roduc t ion Wa te ra nd N u t r ien ts S o i ls a nd H yd ro logy o fE a r ly C e rea lC u l t iva t ion V a r ia t ions i nt he A nnua l R e l iab i l i ty o fS pec i f ic Wa te rS ou rces a nd t he Geog raph ic C i rcumsc r ip t ion o fO p t ima lC e rea l L ands

5 .

R ELAT IVE E NERGET ICS OF P ASTORAL S TRATEG IES I NVOLV ING CAMELS , CATTLE , SHEEP AND GOATS I n t roduc t ion C ompa ra t ive H e rd G row th a nd P roduc t iv i ty I n t roduc t ion A nnua l H e rd G row th A nnua l H e rd C a lo r ic P roduc t iv i ty A nnua l L abo rC os ts o fP as to ra l S t ra teg ies I n t roduc t ion B ehav io ra nd H e rd ing L abo rC os ts C ame l H e rd S ubd iv is ions a nd A nnua l H e rd ing L abo r : T he R end i l le D a ta

4 0 4 1 4 2 4 2 4 2 4 3 4 3 4 3 4 6 4 6 4 7 4 8 5 0 5 3 5 3

5 5 5 5 5 5 5 5 5 6 5 7 5 8 5 9 6 0 6 0 6 1 6 3 6 4 6 4 6 4 6 4 6 6 6 6 6 6 6 6 6 8

7 3 7 3 7 3 7 3 7 3 7 3 7 6 7 6 7 6 7 8

C ame l H e rd S ubd iv is ions a nd A nnua l H e rd ing L abo r : T he A l Mu r rah D a ta H e rd S ubd iv is ions a nd A nnua l H e rd ing L abo rf o rC a t t le , S heep a nd G oa t P as to ra l is ts T he H e rd ing D ay v s . D a i ly H e rd ing L abo rH ou rs Max imum H e rd S i ze a nd A nnua l H e rd ing L abo r E ncampmen tT ime Manya t ta a nd H e rd ing C amp L abo rC os ts A nnua l Gab ra E ncampmen tC os ts A nnua l R end i l le E ncampmen tC os ts B lack T en tL abo rC os ts A nnua lA lM u r rah E ncampmen tC os ts T he E f fec ts o fC a rn ivo re P reda t ion o n A nnua l E ncampmen tC os ts M i lk ing T ime B u tche r ing T ime S umma ry o fT o ta lA nnua l P as to ra l L abo rC os ts R e tu rn R a tes f o r P as to ra l S t ra teg ies I n vo lv ing C ame ls , C a t t le , S heep a nd G oa ts R e tum R a tes f o rP as to ra lC ame l H e rds C ompa ra t ive R e tu rn R a tes f o r Mea t -On ly P as to ra l S t ra teg ies C ompa ra t ive R e tu rn R a tes f o r Mea t P l us M i lk P as to ra l S t ra teg ies M i lk P rocess ing a nd P as to ra l R e tu rn R a tes 6 .

RELAT IVE E NERGET ICS OF E ARLY WHEAT AND B ARLEY S TRATEG IES I n t roduc t ion C u l t iva ted Whea ta r id B a r ley Y ie lds I n t roduc t ion Med ieva l Whea t a nd B a r ley Y ie lds i nt he W inches te r B i shop r ic Whea ta nd B a r ley Y ie lds i nt he N ea rE as t F ie ld L abo rC os ts o fC u l t iva t ing Whea t a nd B a r ley U nde rC ond i t ions o fH and T i l lage a nd L i th ic T echno logy I n t roduc t ion H and T i l lage S ow ing L abo r Un de rH and T i l lage R eap ing L abo ri nE a r ly C e rea l S t ra teg ies F ie ld L abo rC os ts o fC u l t iva t ing Whea ta nd B a r ley U nde rC ond i t ions o fA n ima lT rac t ion T i l lage an d E a r ly Me ta lT echno logy I n t roduc t ion R oman F i e ld T ask L abo r U nde rA rd C u l t iva t ion : T he D a ta o fC o lume l la H is tor ica l V a r ia t ions i nt he E f ic iency o fA n ima lT rac t ion T i l lage P rocess ing L abo rC os ts o f Whea ta nd B a r ley I n t roduc t ion I n i t ia l P rocess ing : T h resh ing a nd W innow ing F i na l P rocess ing : G r ind ing a nd P ound ing R e tu rn R a tes f o r Whea t a nd B a r ley E xp lo i ta t ion a nd C u l t iva t ion I n t roduc t ion P os tencoun te r R e tu rn R a tes f o r E xp lo i t ing W i ld Whea tS tands R e tu rn R a tes f o rE a r ly Whea t a nd B a r ley C u l t iva t ion C ompa ra t ive R e tu rn R a tes f o r C u l t iva t ing S o rghum a nd P ea r l M i l le t I n t roduc t ion

7 9 8 2 8 3 8 4 8 5 8 6 8 9 9 0 9 0 9 2 9 3 9 3 9 5 9 7 9 9 9 9 1 00 1 02 1 02

1 09 1 09 1 09 1 09 1 09 11 13 13 13 15 15 18 18 19 1 20 1 24 1 24 1 24 1 26 1 27 1 27 1 27 1 30 1 34 1 34

Mean R e tu rn R a tes f o rC u l t iva t ing P ea r l M i l le ta t G en ie r iV i l lage i n1 949 Mean R e tu rn R a tes f o rC u l t iva t ing S o rghum a t G en ie r iV i l lage i n1 949 7 .

E NERGET IC E XPECTAT IONS AND T HE ARCHAEOLOG ICAL R ECORD I n t roduc t ion P o ten t ia l P rob lems i nt he I n te rp re ta t ion o fF auna l D epos i ts R e levan tt oE a r ly P as to ra l F ood P roduc t ion E ne rge t ics o fA l te rna t ive P as to ra l S t ra teg ies T he R e la t ive R ank ing o fP as to ra l S t ra teg ies a nd t he A ccep ted T empo ra l S equence o fA n ima l D omes t ica t ion A ccoun t ing f o rt he L a te D omes t ica t ion o fC ame ls : T he I mpor tance o fE nv i ronmen ta l C ond i t ions a nd t he P hy logeny o ft he A n ima ls I n vo lved We re C a t t le R ea l ly D omes t ica ted L a te rt han S heep a nd G oa ts? A rchaeo log ica l E xpec ta t ions B ased U pon t he R e la t ive E ne rge t ics o f E a r ly Whea t a nd B a r ley E xp lo i ta t ion a nd C u l t iva t ion I n t roduc t ion When S hou ld Whea t a nd B a r ley H ave E n te red P reh is to r ic D ie ts? R e tu rn R a tes a nd C i rcumsc r ibed E nv i ronmen ts : I mp l ica t ions f o r P os t -P le is tocene H uman D emog raphy , T e r r i to r ia l i ty , a nd S oc ia lC omp lex i ty A rchaeo log ica l E xpec ta t ions B ased U pon E ne rge t ic C ompa r isons B e tween P as to ra la nd C e rea lS ubs is tence Op t ions G ene ra lC ons ide ra t ions E ne rge t ic E xpec ta t ions C ompa r isons w i th t he A rchaeo log ica l R eco rd C onc lus ions

1 34 1 35

1 37 1 37 1 38 1 40 1 40

1 41 1 43 1 47 1 47 1 47

1 49 1 52 1 52 1 53 1 57 1 59

APPEND ICES A .

B .

C .

D .

DROMEDARY CAMEL POPULAT ION DYNAM ICS AND S LAUGHTER OFFTAKE F ROM A MODEL HERD OVER A ONE YEAR P ER IOD

1 61

CALOR IC P RODUCT IV ITY OF T HE MODEL CAMEL H ERD OVER A ONE Y EAR P ER IOD

1 85

AGE S PEC IF IC GRO WTH R ESULTS FOR T HE F EMALE COMPONENT OF A MODEL CAMEL HERD OVER A FORTY YEAR P ER IOD B ASED UPON T HE DATA P RESENTED I NA PPEND IX A

2 05

AGE S PEC IF IC GROWTH RESULTS FOR T HE I MMATURE MALE COMPONENT OF A MODEL CAMEL HERD OVER A FORTY YEAR P ER IOD B ASED UPON THE DATA PRESENTED I N A PPEND IX A

2 09

R EFERENCES C ITED

2 11

I NDEX

2 53

i v

L IST OF TABLES

T ABLE 1 .

P AGE

P hy logene t ic R e la t ionsh ips o fC e rea ls A cco rd ing t ot he Geog raph ic R eg ions o fT he i rI n i t ia l H usband ry

4 5

2 .

H yd ro log ic A ssoc ia t ions o fS i tes E xh ib i t ing E a r ly C e rea l C u l t iva t ion

6 7

3 .

E s t ima ted P roduc t ion o f Whea t a nd A rea C u l t iva ted i nJ o rdan b y R a in fa l l R eg ions

7 1

4 .

C ompa ra t ive P as to ra l H e rd G row th D ynam ics

7 4

5 .

C ompa ra t ive C a lo r ic P roduc t iv i ty f o r P as to ra l H e rds C on ta in ing E xac t ly 1 00 H ead

7 4

C ompa ra t ive C a lo r ic P roduc t iv i ty f o r P as to ra l H e rds C on ta in ing E xac t ly 1 00 H ead When P o ten t ia l P os t -P le is tocene S ize R educ t ions a re A ccoun ted F o r

7 5

7 .

N u mbe ro fH e rde rs S easona l ly R equ i red P e rR end i l le C ame l H e rd

8 0

8 .

C on t r ibu t ion o fE ach Ma le H e rde rA ge C lass t oT o ta lA nnua l R end i l le C ame l H e rd ing L abo r

8 1

S umma ry o fA nnua l C ame l H e rd ing L abo r R equ i remen ts Among t he R end i l le

8 2

6 .

9 .

1 0 . S umma ry o fA nnua l P as to ra l L abo rf o r Mea t -On ly C ame l S t ra teg ies I n vo lv ing H e rds C on ta in ing 1 00 H ead

9 8

1. S umma ry o fA nnua l P as to ra l L abo rf o r Mea t -On ly C a t t le , S heep a nd G oa t S t ra teg ies I n vo lv ing H e rds C on ta in ing 1 00 H ead

9 8

1 2 . S umma r yo fA nnua l P as to ra l L abo rf o r aMea t P l us M i lk C ame l S t ra tegy I n vo lv ing aH e rd C on ta in ing 1 00 H ead a tV a r ious L eve ls o fM i lk P roduc t ion W i th N o rma lO f t take

9 9

1 3 . S umma ry o fA nnua l P as to ra l L abo rf o r Mea t P l us M i lk C a t t le , S heep a nd G oa t S t ra teg ies I n vo lv ing H e rds C on ta in ing 1 00 H ead

9 9

1 4 .

E xpec ted D ec l ine i nt he T o ta l C a lo r ic P roduc t iv i ty o fP as to ra l H e rds When One -Ha l fo ft he L ac tose C on ten t o fT o ta l H e rd M i lk P roduc t ion i sL os t d ue t oF e rmen ta t ion P rocess ing

10 4

1 5 . Whea t a nd B a r ley Y ie ld E xpec ta t ions o fB edou in F a rme rs i nt he N egev

1 2

1 6 . A ve rage Whea t a nd B a r ley Y ie lds ( kg /du ) o nA rab F a rms i nP a les t ine A cco rd ing t o aB r i t ish C omm i t tee R epo r t o n t he C ond i t ion o fA g r icu l tu ra l is ts d a ted J u ly 1 3 , 1 930

112

1 7 .

1 8 .

E xpec ted Whea ta nd B a r ley Y ie lds ( k g /du ) o n Ma r l L ands i nC o rnwa l l a nd D evon i nA .D . 1 668

112

E xpec ted Whea t a nd B a r ley Y ie lds ( k g /du ) U nde rT rad i t iona l A g r icu l tu ra l P rac t ices a tA evan ,T u rkey

112

1 9 .

R e la t ive E f f ic iency o fH and T i l lage U s ing V a r ious I mp lemen ts

1 14

2 0 . T he R e la t ive R eap ing E f ic ienc ies o fP reh is to r ic t h rough Mode rn H and S ick les

1 16

2 1. C ompa ra t ive E f f ic ienc ies ( h rs /du ) o fA l te rna t ive R eap ing I mp lemen ts a nd t he A dd i t iona l L abo rR equ i red f o r Ga the r ing , B ind ing a nd R ak ing

1 17

2 2 . T o ta l L abo rf o r One l uge rum o f Whea t i nR oman A g r icu l tu re A cco rd ing t oC o lume l la ( 1 .12 .1 )

19

2 3 . T o ta l L abo rf o r One l uge rum o fB a r ley i nR oman A g r icu l tu re A cco rd ing t oC o lume l la ( 1 .12 .2 )

19

2 4 .

N o rma l D raugh ta nd Wo rk P o ten t ia lo fV a r ious A n ima ls

1 20

2 5 .

E f ic iency o fR oman S ymme t r ica l A rd T i l lage U s ing O ne Y oke o fO xen ( Two A n ima ls ) A cco rd ing t oC o lume l la ( 1 .4 .8 )

1 22

E f ic iency o fR oman S ymme t r ica l A rd T i l lage U s ing O ne Y oke o fO xen ( Two A n ima ls ) A cco rd ing t oC o lume l la ( 1 .12 .8 )

1 22

R epo r ted T i l lage E f ic ienc ies a nd D raugh ts o fS ymme t r ica l A rds

1 23

2 6 .

2 7 .

2 8 . T i l lage E f ic iency o fJ ames Sma l l 's Sw ing P l ough D rawn b y Two H o rses

1 23

2 9 . T i l lage E f f ic ienc ies a nd D raugh ts o fE a r ly Ame r ican Wooden P loughs

1 23

3 0 .

E f ic iency o fT h resh ing a nd W innow ing Whea t b yT rad i t iona l Me thods i nI r an

1 25

3 1.

E f f ic iency o fT h resh ing a nd W innow ing Whea t U s ing F l a i ls a nd S hee ts i n E a r ly Ame r ican A g r icu l tu re

1 25

E f ic iency o fT h resh ing a nd W innow ing B a r ley U s ing F l a i ls a nd S hee ts i n E a r ly Ame r ican A g r icu l tu re

1 25

3 3 .

M i l l ing L abo rf o r Whea ta s R epo r ted i nV a r ious S ources

1 26

3 4 .

M i l le t P rocess ing L abo r( h rs /kg ) o fS even Women T imed I nK ampamba V i l lage , R hodes ia

1 27

3 2 .

3 5 . S umma ry o fT o ta l C o l lec t ing a nd P rocess ing L abo rf o rO ne K g o fW i ld E i nko rn Whea t

1 28

3 6 .

P os tencoun te r R e tu rn R a tes f o r E xp lo i t ing W i ld E inko rn Whea t

1 28

3 7 .

P os tencoun te r R e tu rn R a tes f o rV a r ious S eed a nd N u t R esou rces i nt he G rea t B as in A s E xpe r imen ta l ly D e r ived b y S imms ( 1984 )

1 29

P os tencoun te r R e tu rn R a tes f o rA lyawa ra A cac ia S eed E xp lo i ta t ion I n A us t ra l ia a sC a lcu la ted b y O 'Conne l la nd H awkes ( 198 1 , 1 984 )

13 0

3 8 .

3 9 . S umma ry o fT o ta l L abo rf o rC u l t iva t ing One D unum o fL and U nde rH and T i l lage a nd E a r ly L i th ic T echno logy w i th Whea t Y ie ld ing 5 0 K g /du o rB a r ley Y ie ld ing 6 0K g /du

1 31

4 0 . S umma ry o fT o ta l L abo rf o rC u l t iva t ing O ne D unum o fL and U nde rH and T i l lage a nd A dvanced L i th ic T echno logy w i th Whea tY ie ld ing 5 0 K g /du o rB a r ley Y ie ld ing 6 0K g /du

1 31

4 1. S umma r yo fT o ta l L abo rf o rC u l t iva t ing One D unum o fL and U nde rA n ima l T rac t ion T i l lage a nd E a r ly Me ta l T echno logy w i th Whea tY ie ld ing 5 0 K g /du o rB a r ley Y ie ld ing 6 0 K g /du

1 31

v i

4 2 .

R epo r ted Ges ta t ion P e r iods i nt he C ame l

1 62

4 3 . C ame lY ean ing R a tes b y D is t r ic t i nK azakhs tan , S ov ie tC en t ra l A s ia , 1 964

1 67

4 4 . T he F ema le Mo r ta l i ty R a tes i nD ah l a nd H jo r t 's S econd Mode l o fC ame l H e rd G row th ( 1 976 :83 )

1 69

4 5 . A ge -Spec i f ic N a tu ra l Mo r ta l i ty a nd R esu l t ing S u rv ivo rsh ip f o r Ma le a nd F ema le C ame lC a lves a tB i kane r , I n d ia

1 70

4 6 . Obse rved Mo r ta l i ty R a tes Among R end i l le C ame l H e rds i n1 976

1 71

4 7 . A ssumed N a tu ra l Mo r ta l i ty a nd R esu l t ing S urv ivo rsh ip R a tes Among D romeda ry C ame ls

1 71

4 8 .

F ema le M o r ta l i ty V a lues

1 74

4 9 .

I mma tu re Ma le Mo r ta l i ty V a lues

1 75

5 0 .

Mean F ema le A ge D i s t r ibu t ions D e r ived f r om t he L as t 1 4Y ea rs o ft he S imu la t ion

1 78

5 1.

I mma tu re Ma le S tab le A ge D is t r ibu t ion

1 78

5 2 .

E xpec ted A ge a nd S ex S t ruc tu re o f aMode l C ame l H e rd C on ta in ing 5 5 R ep roduc t ive ly M a tu re F ema les ( Exc lud ing A du l t Ma les )

1 80

5 3 .

N umbe ro fF ema le C ame ls w h ich c an b eS e rv iced b y aS ing le S tud

1 81

5 4 .

Mode lo ft he R e ten t ion R a te f o r Ma le C ame ls E n te r ing T he i r7 t h Y ea rw hen t he Goa l i s Ma in ta in ing 5t o 6Ma tu re Ma les

1 81

5 5 .

N o rma lO f f take f r om t h e Mode lC ame l H e rd

1 83

5 6 .

Max imum O f f take f r om t he Mode lC ame l H e rd

1 84

5 7 .

S laugh te ra nd P ropo r t iona lY ie ld D a ta o n 1 50 A du l t S oma l i D romeda r ies

1 87

5 8 .

S laugh te ra nd P ropo r t iona lY ie ld D a ta o n 7 4 Ma le T u rkmen ian D romeda r ies Ma in ta ined o n a nA ve rage P l ane o fN u t r i t ion

1 88

5 9 .

S tanda rd A na lys is o f4 0C ame l H umps

1 90

6 0 .

E xpec ted Y ie lds o fE d ib le I t ems P e rC ame l a s aP e rcen tage o fL i vewe igh t

1 90

6 1. A ve rage L i vewe igh ts R epo r ted f o r Ma tu re D romeda r ies

1 91

6 2 .

A ve rage L i vewe igh ts R epo r ted f o r Ma tu re B ac t r ians

1 91

6 3 .

E s t ima ted A ge a nd S ex S pec i f ic L i vewe igh ts f o rC ame ls i nt he Mode l H e rd

1 92

6 4 .

E xpec ted Mea t , O rgan , F a t a nd B lood Y ie lds P e r I n d iv idua l i nt he Mode l H e rd

1 92

6 5 . T o ta l Mea t , O rgan , a nd F a tY ie ld f r om t he Mode lC ame l H e rd w i th N o rma l O f f take

1 93

6 6 . T o ta l Mea t , O rgan , a nd F a tY ie ld f r om t he Mode lC ame l H e rd w i th Max imum M ake

1 93

6 7 .

1 94

S tanda rd A na lyses o fC ame l Mea t P u rchased i nC a i ro

V I

6 8 .

P ro te in a nd F a tC on ten to fT r immed Mea tf r om F e ra l C ame ls i nA us t ra l ia

1 94

6 9 . C a lo r ic V a lue o fC ame l Mea t P u rchased i nC a i ro

1 95

7 0 . C a lo r ic V a lue o f Mea t Ob ta ined f r om F e ra lC ame ls i nA us t ra l ia

1 96

7 1. S tanda rd A na lyses a nd C a lo r ic V a lues o fC e r ta in C a t t le O rgans A cco rd ing t o P yke ( 1970 )

1 96

7 2 . S tanda rd A na lyses a nd C a lo r ic V a lues o fC e r ta in C a t t le O rgans A cco rd ing t o U .S .D .A . ( 1975 )

1 96

7 3 . T o ta lC a lo r ic Y ie ld b y E d ib le I t em f r om t he Mode l C ame l H e rd w i th N o rma l O f f take o ve rO ne Y ea r

1 97

7 4 . T o ta l C a lo r ic Y ie ld b y E d ib le I t em f r om t he Mode l C ame l H e rd w i th Max imum O ff take o ve rO ne Y ea r

1 98

7 5 .

2 00

R epo r ted Y ie lds o fD romeda ry M i lk f o rH uman C onsump t ion

7 6 . A pp rox ima te Y ie lds o fM i lk A cco rd ing t oA n ima l T ype a s Obse rved among K e lT amasheq H e rds i nt he Wes t A f r ican S ahe l

2 01

7 7 .

2 02

L eng th o fL ac ta t ion Among D romeda ry C ame ls

7 8 . S tanda rd A na lyses o fD romeda ry G a rne rs M i lk

2 03

L IST OF F IGURES

F IGURE

P AGE

1 .

D ie tb read th mode l

1

2 .

S ubs is tence s t ra tegy r e tu rn r a tes a nd t he e vo lu t ion o ra dop t ion o ff ood p roduc t ion o p t ions

12

A nnua l d ev ia t ion i nt o ta l p rec ip i ta t ion a tJ e rusa lem f r om t he c umu la t ive mean f o rt h ey ea rs 1 85 1 t h rough 1 960

6 9

A nnua l d ev ia t ion i nt o ta l N i le d i scha rge a tA swan f r om t he c umu la t ive mean f o rt h ey ea rs 1 87 1 t h rough 1 965

6 9

5 .

N umbe ro fh e rde rs r e qu i red s easona l ly p e r R end i l le c ame l h e rd

8 0

6 .

R e la t ive s easona lc on t r ibu t ion t oc ame l h e rd ing l a bo rb y a ge -c lass among R end i l le ma le h e rde rs

8 1

3 .

4 .

7 .

8 .

9 .

1 0 .

1.

1 2 .

R anges o fp o ten t ia l r e tu rn r a tes f o rc ame l h e rds u nde rv a r ious p roduc t ive p oss ib i l i t ies

10 0

R e la t ionsh ips b e tween a nnua l c a lo r ic p roduc t iv i ty a nd r equ i red l a bo rf o r mea t -on ly p as to ra ls t ra teg ies

10 1

R anges o fp o ten t ia l r e tu rn r a tes f o r mea t -on ly p as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts

1 01

R e la t ionsh ips b e tween a nnua l c a lo r ic p roduc t iv i ty a nd r equ i red l a bo rf o r mea tp l us m i lk p as to ra l s t ra teg ies

10 3

R anges o fp o ten t ia l r e tu rn r a tes f o r mea t p l us m i lk p as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts

10 3

R anges o fp o ten t ia l r e tu rn r a tes f o r mea t p l us m i lk p as to ra l s t ra teg ies w hen m i lk p rocess ing i sa ccoun ted f o r

10 6

1 3 . A nnua ld ev ia t ions i n mean w hea t a nd b a r ley y i e lds i nt he W inches te r B i shop r ic f r om t he c umu la t ive m ean f o rt he y ea rs A .D . 1 232 t h rough 1 349 1 4 .

D ev ia t ions i nh oe ing l a bo rp e r h u tc ompound f r om t he c umu la t ive mean f o ra l h u tc ompounds i nG en ie r i V i l lage , Gamb ia , i n1 949

110

114

1 5 . S imp l i f ied h i s to r ic e vo lu t ion o ft he mou ldboa rd p l ough

18

1 6 . V a r ia t ions i nt he mechan ics o ft i lage b e tween s ymme t r ica l a rds a nd p l oughs w i th mou ldboa rds

12 1

1 7 .

1 8 .

R e la t ionsh ips b e tween c a lo r ic p roduc t iv i ty a nd l a bo rf o rw hea t a nd b a r ley c u l t iva t ion a tv a r ious s t ages o ft e chno logy a nd v a r iab le y i e lds

13 2

R anges o fp o ten t ia l r e tu rn r a tes f o rw hea t a nd b a r ley c u l t iva t ion u nde rv a r ious s t ages o ft e chno logy a nd v a r iab le y i e lds

133

1 9 .

2 0 .

2 1.

E ne rge t ic r e la t ionsh ips e xpec ted b e tween e a r ly c e rea l s t ra teg ies a nd mea t -on ly p as to ra ls t ra teg ies

1 54

E ne rge t ic r e la t ionsh ips e xpec ted b e tween e a r ly c e rea l s t ra teg ies a nd m ea tp l us m i lk p as to ra ls t ra teg ies

1 55

E xpec ted v a r ia t ions i nt he e ne rge t ic r e la t ionsh ips b e tween c e rea l s t ra teg ies u nde rc hang ing t echno log ica l c ond i t ions a nd mea t p l us m i lk p as to ra l s t ra teg ies

1 56

2 2 . S imu la ted g row th i nt he t o ta l f ema le a nd i mma tu re ma le c omponen ts o f ap as to ra lc ame l h e rd

1 77

2 3 . S easona lv a r ia t ions i nt he a pp rox ima te m i lk y i e ld f o r h uman c onsump t ion b y a n ima l a so bse rved among K e l T amasheq h e rds i nt he Wes tA f r ican S ahe l

2 01

x

PREFACE

I n f luenced b y t he c u l tu ra l ma te r ia l ism o f Ma rv in H a r r is a nd s t imu la ted b y t he s e lec t ion is t t h ink ing emp loyed i nb ehav io ra l e co logy ( evo lu t iona ry e co logy , s oc iob io logy ) , ag roup o ft h ree U n ive rs i ty o f U tah p ro fesso rs a nd s even g radua te s t uden ts b egan t o i n fo rma l ly mee t o f f c ampus i n1 983 t od iscuss t he i mp l ica t ions o f mode rn e vo lu t iona ry t heory a s a pp l ied t og ene ra l a n th ropo log ica l i s sues . T he t op ics a dd ressed r anged f r om e a r ly h om in id f eed ing s t ra teg ies t h rough f ood s ha r ing among h un te r -ga the re rs , a nd f r om a bo r ig ina l s eed e xp lo i ta t ion i nt he G rea t B as in t h rough c on f l ic ts i n ma le fema le r ep roduc t ive s t ra teg ies . W i th in a s pan o ft h ree y ea rs , membe rs o f wha t c ame t o b e c a l led t he " Hun ter -Ga the re r R esea rch G roup " p roduced s eve ra l s cho la r ly a r t ic les a nd s i x d oc to ra l d i sser ta t ions f ocuss ing u pon t he u t i l i ty o f mode rn e vo lu t iona ry t heory a s a pp l ied t o ab e t te ru nde rs tand ing o fh uman a f fa i rs . T h is b ook i sb ased u pon o ne o ft hose d oc to ra l d i sse r ta t ions . Wr i t ten i n1 986 w i th in t he D epa r tmen t o fA n th ropo logy a tt he U n ive rs i ty o f U tah , t h is wo rk a dd ressed t he a pp l icab i l i ty o f b ehav io ra l e co logy t o a rchaeo log ica l i s sues i n vo lv ing t he e vo lu t ion a nd a dop t ion o fa l te rna t ive f ood p roduc t ion s t ra teg ies i nt he N ea r E as t a nd N o r th A f r ica . T he p resen t b ook ma in ta ins t ha t o r ig ina l manusc r ip t e ssen t ia l ly i n tac t , a l though a dd i t iona l d a ta a nd c o r rec t ions h ave b een i n co rpo ra ted a nd s ome s ec t ions r eo rgan ized . As ub jec t i n dex h as a l so b een i n c luded t o make t he manusc r ip t mo re u se fu l a s ar e fe rence wo rk . R ega rd less o f h ow o ne f ee ls a bou t t he t heo re t ica l a nd me thodo log ica l i s sues r a ised i nt h is w ork , o r t he c onc lus ions d rawn f r om t he a pp roach emp loyed , t he b as ic d a ta o n v a r ious a spec ts o f f ood p roduc t ion wh ich i tc on ta ins ( much o f wh ich i sb rough tt o ge the rf o rt he f i r s tt ime ) s hou ld b e o fu se t ov a r ious s cho la rs . Many i n d iv idua ls c on t r ibu ted t ot he f ocus , c on ten t a nd t echn ica l d eve lopmen t o fb o th t he o r ig ina l d isser ta t ion a nd t h is r ev ised a nd e xpanded manusc r ip t . F o r t he i r c r i t ica l a dv ice , g u idance a nd e d i tor ia l c ommen tary o n v a r ious d ra f ts o f t he o r ig ina l wo rk , Iam i n deb ted t o K r is ten H awkes , L au rence D . L oeb a nd D enn is L .H eske l , whose e f fo r ts a s membe rs o f m y d oc to ra l c omm i t tee a re d eep ly a pp rec ia ted . I n p a r t icu la r , t he i n te l lec tua l s t i mu lus o f K r is ten H awkes , b o th a s amembe ro ft he H un te r -Ga the re r R esea rch G roup a nd n ow a s ac o l league , i s g ra te fu l ly a cknow ledged . F or s cho la r ly c onve rsa t ions a nd a dv ice c once rn ing v a r ious t heo re t ica l a spec ts o f t h is r e sea rch , Iw ish t ot hank R icha rd D .A lexande r , E r ic L .C hamov , Ma rv in H a r r is a nd H i l la rd K ap lan . E ach h as made t he i r own s pec ia l c on t r ibu t ion t ot he r ea l iza t ion o ft h is b ook . F o r u npub l ished manusc r ip ts a nd ' p e rsona l c ommun ica t ions , Imus t e xp ress my a pp rec ia t ion t o A l ison V . G . B e t ts , D ona ld 0 . H en ry , K im R . H i l l , R obe r t K . H i tchcock , P e te rJ ones , M ichae l L i t t le , J ames F . O 'Conne l l a nd W i l l iam I .T o r ry . F o r p ub l ished manusc r ip ts , p e rsona l c ommun ica t ions a nd s ugges t ions n o t i n c luded i nt he o r ig ina ld i sser ta t ion , Iam i n deb ted t oS ebas t ian P ayne , Ga ry 0 . R o l le fson , S teven A . R osen a nd F red Wendo r f . F o rt he i rc r i t ica l c ommen ta ry a nd h e lp fu l s ugges t ions , Iam g ra te fu l t oZ b ign iew T . F i ema , A lan S . L i ch ty , K a ren D . L upo , a nd S teven R . S imms . Z b ign iew T . F i ema k i nd ly t r ans la ted r e fe rences i nR uss ian , w h i le L au rence D . L oeb d i d s of o rw orks i nH eb rew . T o a l o ft h e membe r s o f my f am i ly , a nd e spec ia l ly t o my s i s te rA . K ay R usse l l Sma l lman a nd my c ous in N ico la R . Zwaschka K ledas , Iam t hank fu l f o ry ou r e ncou ragemen t a nd s uppor t o ve r s o many y ea rs . F ina l ly , Iw ish t o t hank t he L . S . B . L eakey F ounda t ion f o r a g ran t t o c onduc t e thnoa rchaeo log ica l r esea rch among b edou in p opu la t ions i ns ou thwes te rn J o rdan d ur ing t he S p r ing o f 1 986 , a low ing f o rt he p e rsona l o bserva t ions i n c luded i nt h is manusc r ip t .

I NTRODUCT ION

R esea rch Goa ls a nd F ocus T h is s tudy p resen ts a n a pp l ica t ion o f b ehav iora l e co logy t o e thnograph ic a nd a rchaeo log ica l d a ta i na n a t temp t t o f u r the r o ur c u r ren t u nde rs tand ing o f t he e vo lu t ion a nd a dop t ion o fa l te rna t ive f ood p roduc t ion s t ra teg ies i nt he N ea r E as t a nd N or th A f r ica d ur ing t he H o locene e poch . A me thodo logy i s d eve loped f o r a pp ly ing e vo lu t ionary t heo ry t o a n u nde rs tand ing o f why p a t te rns o f e a r ly h uman f ood p roduc t ion t ook t he f o rm t hey d id , when a nd where t hey d i d . C os t a nd b ene f i t d a ta a re g ene ra ted f o r a n ima l a nd c e rea l h usband ry s t ra teg ies i n af ash ion c ons is ten t w i th t ha t emp loyed i nr ecen t e vo lu t iona ry s t ud ies o f h u man f o rag ing b ehav io r . E ne rge t ic e xpec ta t ions d e r ived f r om t hese d a ta a re t hen u sed t o e xp lo re t he e x ten t t o w h ich s imp le o p t im iza t ion mode ls m igh t e xp la in t he d ec is ion mak ing a ssoc ia ted w i th t he s pa t ia l a nd t empo ra l a ppea rance o f a l te rna t ive p as to ra l a nd a g r icu l tura l s ubs is tence o p t ions a s r e f lec ted i nt he H o locene e poch a rchaeo log ica l r eco rds o ft he N ea r E as t a nd N or th A f r ica . I n i t ia l ly , r anges o fp oss ib le r e tu rn r a tes ( ene rgy y i e ld / t ime i n ves ted t oo b ta in t ha t y i e ld ) a re e s tab l ished f o r p as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts , a nd a g r icu l tu ra l s t ra teg ies i n vo lv ing t he c e rea ls whea t a nd b a r ley . U s ing t he e ne rge t ic r esu l ts i nc on junc t ion w i th a s imp le o p t im iza t ion mode l , t h ree i s sues r e levan t t o o ur p resen t u nde rs tand ing o f t he e vo lu t ion a nd a dop t ion o fp as to ra l a nd a g r icu l tura l s ubs is tence s t ra teg ies i nt he N ea r E as t a nd N o r th A f r ica a re s pec i f ica l ly a dd ressed . T hese i s sues a re : ( 1 ) I si tr easonab le t o a ssume t ha t p as to ra l s t ra teg ies n ecessa r i ly e vo lved a s a dap ta t ions t oe nv i ronmen ts ma rg ina l t oa g r icu l tu re? , ( 2 ) Wou ld m i lk p roduc t ion h ave b een s i gn i f ican t i nt he e vo lu t ion o f e a r ly p as to ra l s t ra teg ies? , a nd ( 3 ) Why w e re p as to ra l s t ra teg ies i n vo lv ing c ame ls a dop ted l a te r t han o the r p as to ra l a nd c e rea l s ubs is tence o p t ions? P red ic t ions d e r ived f r om t he e co log ica l a nd e ne rge t ic a na lys is o f f ac to rs r e levan tt ot hese i s sues a re t hen t es ted a ga ins t H o locene e poch a rchaeo log ica l d a ta i n o rde r t o e xp lo re t he e x ten t t o wh ich o p t im iza t ion mode ls m igh t e nhance c ur ren t me thods o f a rchaeo log ica l i n fe rence .

O rgan iza t ion a nd S ynops is T he b as ic p rem ise o ft h is s t udy i st ha t s i gn i f ican t a dvances i nu nde rs tand ing t he e vo lu t ion a nd a dop t ion o ff ood p roduc t ion s t ra teg ies a re u n l ike ly u n less man -an ima l -p lan t r e la t ionsh ips a re i n ves t iga ted w i th in t he c on tex t o fg ene ra l t heo r ies wh ich a dd ress t he u nde r ly ing p r inc ip les a f fec t ing s ubs is tence b ehav io r . C hap te r 1w i l l e l abo ra te o n t h is t heme . P rev ious p e rspec t ives o n t he e vo lu t ion a nd a dop t ion o f a l te rna t ive a g r icu l tu ra l a nd p as to ra l s t ra teg ies a re b r ie f ly d i scussed . Wh i le e a r ly r esea rch i n to t he " o r ig ins " o ff ood p roduc t ion h as b een a pp rop r ia te ly r ep laced b y af ocus o n t he c ond i t ions u nde r wh ich a g r icu l ture a nd p as to ra l ism b ecame p ro f i tab le r e la t ive t o c on t inued f o rag ing , p opu la r p red ic t ive mode ls i n vok ing e co log ica l ma rg ina l i ty a nd p roduc t iv i ty p e r u n i t a rea f a i l t o a dequa te ly a dd ress t he u nde r ly ing p r inc ip les a fec t ing s ubs is tence b ehav io r . A n a l te rna t ive a pp roach t o i s sues o fe a r ly f ood p roduc t ion i n vo lves mode ls d e r ived f r om b ehav io ra l e co logy wh ich i n voke d ec is ion mak ing b ased u pon t he r e la t ive r e tu rn r a tes ( ca lo r ic y i e ld / t ime i n ves ted ) o f a l te rna t ive , s ubs is tence o p t ions . T he u nde r ly ing p rem ises o f t h is a pp roach a re t ha t e ne rgy f ue ls r ep roduc t ion , a nd t ha t a n o rgan ism w h ich c ap tu res e ne rgy mo re e f f ic ien t ly t han i t sc ompe t i to rs wou ld e n joy ar e la t ive r ep roduc t ive a dvan tage . I ne f fec t , p a t te rns o f s ubs is tence b ehav io r w h ich e nhance r ep roduc t ive s uccess d ue t o t he e f ic iency o f e ne rgy c ap ture a re e xpec ted t o p e rs is t a nd s p read a t t he e xpense o f t hose wh ich d o s o l e ss e f fec t ive ly . A s aw o rk ing h ypo thes is , i ti sp roposed t ha t f ood p roduc t ion s t ra teg ies s hou ld h ave b een a dop ted w hen mean f o rag ing r e turn r a tes f e l l e qua l t oo r b e low t he mean r e tu rn r a te f o r a l te rna t ive f ood p roduc t ion o p t ions . F ur the rmo re , a va i lab le f ood p roduc t ion o p t ions s hou ld h ave b een a dop ted a cco rd ing t ot he i rr ank ing b y r e la t ive r e tu rn r a tes .

C hap te rs 2 a nd 3 a dd ress v a r ious p rob lems a nd a ssump t ions i na pp ly ing a n e vo lu t iona ry c os t a nd b ene f i t a pp roach t o t he a na lys is o f a nc ien t p as to ra l a nd a g r icu l tura l e ne rge t ics . I n p as to ra l s t ra teg ies i n vo lv ing e x tens ive h usband ry , t he l a bo r c os ts w ou ld p r ima r i ly c ons is t o f h e rd ing , e ncampmen t , b u tche r ing a nd ( po ten t ia l ly ) m i lk ing t a sks , w h i le t he b ene f i ts o b ta ined w ou ld p r ima r i ly c ons is t o f t he c a lo r ies d e r ived f r om a n ima l s l augh te r a nd ( po ten t ia l ly ) m i lk p roduc t ion . I nt he c u l t iva t ion o fc e rea ls s uch a s w hea t a nd b a r ley , l a bo rc os ts wou ld p r ima r i ly c ons is t o fs o i l p repa ra t ion , s ow ing , h a rves t ing , t h resh ing a nd w innow ing , s t o rage a nd g r ind ing t asks , wh i le t he b ene f i ts w ou ld c ons is t o ft he c a lo r ic y i e ld o ft he p rocessed g ra in . S ince t he q uan t i ta t ive c os t a nd b ene f i t d a ta r equ i red f o r t he c ompa ra t ive e ne rge t ic a na lys is o f t hese s t ra teg ies c an o n ly b e o b ta ined f r om s uch i n d i rec t s ou rces a s mode rn e xpe r imen ta l r ep l ica t ion o r f r om h i s tor ic a nd e thnog raph ic d ocumen ta t ion , t he p o ten t ia l e f fec ts o f t echno log ica l i n nova t ions o n l a bo r c os ts a nd h uman s e lec t ion o n p l an t a nd a n ima l p roduc t iv i ty b e tween t he p reh is to r ic p as t a nd t he h i s to r ic a nd e t hnog raph ic p resen ta r e d e ta i led . C hap te r 2a dd resses t h is i s sue i nr e fe rence t op as to ra l s ubs is tence . T he re i sl i t le r eason t o s uspec t ma jor v a r ia t ions t h rough t ime i n p as to ra l l a bo r c os ts , a nd d a ta o n modern " un imp roved " b reeds o f h e rd a n ima ls s hou ld a dequa te ly r e f lec t t he p o ten t ia l r ange o f p roduc t iv i ty e xh ib i ted b y t he i r p reh is to r ic p rogen i to rs . T he p oss ib i l i ty o f p ropor t iona l s i ze r educ t ions among c a t t le , s heep a nd g oa ts u nde re a r ly h usband ry c an b e a ccoun ted f o ro n t he b as is o f a va i lab le d a ta . A n e xam ina t ion o f t he s pec i f ic b ehav io ra l , mo rpho log ica l a nd p hys io log ica l t r a i ts c ommon ly p e rce ived a s s i gn i f ican t i ne i the rt he d omes t ica t ion p rocess , o ri n d i s t ingu ish ing " w i ld " f r om " domes t ica ted " h e rd s pec ies , s ugges ts t ha t t he p o ten t ia l r ange o f p heno typ ic v a r iab i l i ty a nd p l as t ic i ty a l ready e s tab l ished i ns pec i f ic h e rd s pec ies b y t he e nd o f t he P le is tocene h as b een s e r ious ly u nde res t ima ted i nm any p rev ious a na lyses . T he e fec ts o f h uman s e lec t ion p rac t ices o n h e rd a n ima ls p r io r t ot he s even teen th c en tury A .D . h ave b een e qua l ly o ve res t ima ted . I nr e fe rence t ot he p o ten t ia l f o rd a i ry ing i ne a r ly p as to ra l s t ra teg ies , t he a ssumed s i gn i f icance o fa du l t h ypo lac tas ia h as a l so b een o ve r ly s t ressed . F i na l ly , t he e x tens ive u se o f g ame d r ive l i nes a nd c o r ra ls i nh un t ing h e rd s pec ies b y f o rag ing p opu la t ions i nt he p reh is to r ic p as t a nd t he e t hnog raph ic p resen t i sb r ie f ly d i scussed . S uch p rac t ices p rov ide a p robab le mechan ism f o r t he i n i t ia l c ap ture o fw i ld h e rds f o r u se a s p as to ra l h e rds , w h i le t he s ubsequen t c on t ro l o f t hese a n ima ls c ou ld h ave b een a ch ieved t h rough t he e xp lo i ta t ion a nd man ipu la t ion o fp rev ious ly e s tab l ished h e rd a ge -sex c l asses a nd s oc ia lb ehav io rs . C hap te r 3a dd resses t he u se o f h i s to r ic a nd e thnog raph ic d a ta i n d e te rm in ing t he e ne rge t ics o fe a r ly c e rea l c u l t iva t ion . T echno log ica l v a r ia t ions i nl a bo rc os ts c an b e a ccoun ted f o ro nt he b as is o fa va i lab le d a ta , a nd p roduc t iv i ty d a ta o n n onhyb r id l a nd races o fp redom inan t ly s e l f -po l l ina t ing c e rea ls s hou ld a dequa te ly r e f lec t t he p o ten t ia l p roduc t iv i ty o ft he i r p reh is to r ic p rogen i to rs . C lass ica l , med ieva l , a nd mode rn y i e ld d a ta o n n onhyb r id d ry fa rmed w hea ts f a l l w i th in t he u ppe r r ange o fp roduc t iv i ty r epo r ted f o rw i ld w hea t a nd b a r ley s t ands . T h is means t ha t t he p o ten t ia l r ange o f p roduc t ive v a r iab i l i ty a nd p l as t ic i ty a l ready e s tab l ished i nc e rea l p opu la t ions p r io r t o t he e vo lu t ion o f f ood p roduc t ion s t ra teg ies h as b een s er ious ly u nde res t ima ted i n many p rev ious a na lyses . O nt he o the rh and , t he e f fec ts o fh uman s e lec t ion o n p redom inan t ly s e l f -po l l ina t ing c e rea ls p r io r t o t he s even teen th c en tury A .D . h ave b een f r equen t ly o ve res t ima ted . U n l ike p as to ra l s t ra teg ies , h oweve r , t echno log ica l i n nova t ions may b e e xpec ted t oh ave a f fec ted t he e ne rge t ics o fc e rea l c u l t iva t ion t h rough t ime . C hap te r 4a dd resses t he c ompa ra t ive e co logy o ft h e p r inc ip le a n ima l a nd p l an t s pec ies e xam ined i nt h is t e x t , b eg inn ing w i th c ame ls , c a t t le , s heep a nd g oa ts . P hy logene t ic v a r ia t ions b e tween t hese s pec ies w h ich w ou ld a f fec t t he i r r e la t ive p roduc t ive p o ten t ia l a nd h ence d es i rab i l i ty a s p as to ra l h e rd s pec ies i ns pec i f ic e nv i ronmen ts a r ed i scussed . Ma jo rv a r ia t ions a re o bserved i nt he mechan isms b y w h ich e ach s pec ies m a in ta ins t he rma l s t ab i l i ty a nd w a te r b a lance , a nd i nt he w ays i nw h ich e ach o b ta ins f o rage e ne rgy . T he u n ique f e tu res o f c ame l l o como t ion a re a l so d i scussed . I ng ene ra l , b y t he i n he r i ted c ha rac te r is t ics o ft he i r l o como t ion , d romeda ry c ame ls a re l a rge ly e xc luded f r om e xp lo i t ing w e t e nv i ronmen ts whe re c a t t le , s heep a nd g oa ts f a re b e t te r . C onve rse ly , d ue t ot he i r i n he r i ted w a te r r equ i remen ts , mechan isms o f t he rma l r egu la t ion a nd f eed ing e co logy , c a t t le , s heep a nd , t o al e sse re x ten t , g oa ts , a re l a rge ly e xc luded f r om e xp lo i t ing t he h o t /d ry d eser t e nv i ronmen ts w he re d romeda ry c ame ls f a re b e t te r . V a r ia t ions i nt he r e la t ive s uscep t ib i l i ty o ft hese s pec ies t os pec i f ic d i sease f ac to rs w h ich wou ld a f fec t t he i r r ep roduc t ion a nd mo r ta l i ty r a tes a l so a ppea r t ot r ack t hese s ame e nv i ronmen ta l c ond i t ions . 2

T he e co logy o f whea ts a nd b a r leys i st hen e xam ined . Wh i le b a r ley i s mo re s a l t to le ran t a nd x e rophy t ic t han whea t , b o th r equ i re s im i la r e nv i ronmen ta l c ond i t ions f o r o p t imum g row th . V a r ia t ions i n wa te ra nd s o i l n u t r ien ts a s p r ima ry l i m i t ing f a c to rs i nc e rea l g row th a nd p roduc t iv i ty a re s pec i f ica l ly a dd ressed . T he l o ose , mo is t a luv ia l s o i ls a ssoc ia ted w i th n a tura l o r e as i ly o b ta ined i r iga t ion ( e .g . , l a ke -edge , r i ve r ine , a nd s p r ings ide e nv i ronmen ts , a nd h ab i ta ts w i th c oncen t ra ted r uno f f ) o f ten a low f o r e xcep t iona l ly h i gh s eed y i e lds r e la t ive t o mos t d ry fa rm ing c ond i t ions a nd t he d ry p la teau a nd h i l ls lope e nv i ronmen ts i n wh ich a nces t ra l w hea ts a nd b a r leys a ppa ren t ly e vo lved . H oweve r , s uch h i gh ly -p roduc t ive e nv i ronmen ts a re e xpec ted t ob e g eog raph ica l ly c i rcumsc r ibed , a nd wou ld g ene ra l ly c ons t i tu te o n ly asma l l p ropo r t ion o ft he t o ta l a va i lab le a rab le l a nd e ven w hen d ry fa rm ing r eg ions w i th h i ghe r t han a verage r a in fa l l a re i n c luded . R e la t ive ly " ma rg ina l " d ry fa rm ing l a nds a re g ene ra l ly e xpec ted t oc ons t i tu te t he v as t ma jo r i ty o fa l a va i lab le a rab le l a nd . C hap te r 5d e ta i ls t he r e la t ive e ne rge t ics o fa l te rna t ive p as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts . P roduc t ive v a r ia t ions b e tween t hese s pec ies a re a na lyzed u s ing h e rd g row th a nd p roduc t iv i ty mode ls p rev ious ly d eve loped f o rp as to ra l c a t t le , s heep a nd g oa t h e rds , a nd s im i la r mode ls f o r d romeda ry c ame l h erds a s p resen ted i n t he a ppend ices t o t h is manusc r ip t . W i th e ach o ft hese h e rd s pec ies , t he e ne rgy d e r ived f r om m i lk wou ld c on t r ibu te mo re t han h a l f o f t he t o ta l e ne rgy e thnog raph ica l ly e xpec ted f r om p as to ra l p roduc t ion . A s a pp l ied t o t he p reh is to r ic p as t , t h is s ame r e la t ionsh ip i se xpec ted e ven when p o ten t ia l s i ze r educ t ions i np reh is to r ic c a t t le , s heep a nd g oa t p opu la t ions a re a ccoun ted f o r . I fr anked o n t he s o le b as is o f p roduc t iv i ty p e r n umbe ro fa n ima ls i n ah e rd , d romeda ry c ame ls a ppea rt ob e t he mos t p roduc t ive , f o l lowed b y c a t t le , s heep a nd g oa ts r espec t ive ly , wh i le r ank ing i nt e rms o f h e rd g row th r a tes e fec t ive ly r e ve rses t h is o rde r . S ign i f ican t ly , i fr anked a cco rd ing t op roduc t ive p o ten t ia l u nde r x er ic c ond i t ions , c ame ls wou ld a ga in h ead t he l i s t , f o l lowed b y g oa ts , s heep a nd c a t t le , wh i le r ank ing a cco rd ing t o p roduc t ive p o ten t ia l u nder we t /co ld o r we t /wa rm c ond i t ions e f fec t ive ly r eve rses t h is o rde r . T he l abo r c os ts o f p as to ra l s t ra teg ies a re t hen a dd ressed . H erd ing t asks c ons t i tu te t he v as t m a jo r i ty o fa l p as to ra l l a bo rc os ts , a nd w ou ld r ema in r e la t ive ly c ons tan t e xcep t a tt he l owe r a nd u pper r anges o f h e rd s i ze manageab i l i ty u nde r s pec i f ic e nv i ronmen ta l c ond i t ions . I nd iv idua l h erde rs wou ld b e a b le t o manage l a rge r h e rds o f s heep , g oa ts a nd c a t t le t han o f c ame ls , wh ich w ou ld p a r t ia l ly o fse tv a r ia t ions i nt he r e la t ive p roduc t iv i ty o fe qua l -s ized h e rds . R e turn r a tes a re t hen e s tab l ished f o r p as to ra l c ame l , c a t t le , s heep a nd g oa t h e rds o f v a r ious s i zes , s how ing t ha t p as to ra l r e turn r a tes i n c rease a s ap r ima ry f unc t ion o f h e rd s i ze . B ecause o ft he s im i la r i t ies i nc os ts , t he i r r ank ing a cco rd ing t oe ne rgy o b ta ined p e ru n i t o fl a bo r t ime p a ra l le ls t he i r r ank ing a cco rd ing t ot o ta l e ne rgy p roduc t ion a l one . C ame ls wou ld h ead t he l i s t , f o l lowed b y c a t t le , s heep a nd g oa ts r e spec t ive ly . H oweve r , a s w i th g ross p roduc t iv i ty , t he mo re f avorab le r e turn r a tes f o r c ame l h usband ry wou ld o n ly b e e xpec ted u nde r x er ic c ond i t ions , wh i le t hose e s tab l ished f o r c a t t le , s heep , a nd g oa t h e rds c ou ld b e o b ta ined u nde r we t /co ld o r we t /wa rm c ond i t ions , b u t n o t x e r ic c ond i t ions . C hap ter 6 d e ta i ls t he r e la t ive c os ts a nd b ene f i ts o f w hea t a nd b a r ley e xp lo i ta t ion a nd c u l t iva t ion . I n i t ia l ly , y i e ld d a ta o n w i ld a nd c u l t iva ted whea t a nd b a r ley s t ands a re p resen ted , d raw ing f r om n ume rous c l ass ica l , med ieva l a nd modem s ou rces . T he l a bo rc os ts o fe a r ly w hea t a nd b a r ley c u l t iva t ion a re t hen d e ta i led b y d i v id ing a nnua l l a bo rc os ts i n to t he c a tego r ies o fs o i l p repa ra t ion a nd s ow ing , r eap ing , p rocess ing , a nd s t o rage t asks . Wh i le s o i l p repa ra t ion a nd s ow ing l a bo r a re r e la t ive ly c ons tan t p e r u n i t o fl a nd r ega rd less o fa c tua l c e rea l y i e ld , r eap ing , p rocess ing a nd s t o rage l a bo r a re e xpec ted t o i n c rease a s af unc t ion o fy i e ld p e r u n i t o f l a nd . T he e f fec ts t h rough t ime o f v a r ious t echno log ica l i n nova t ions o n t he e f f ic iency o f s o i l t i lage a nd h a rves t ing l a bo ra re a l so d ocumen ted . R e turn r a tes f o r e xp lo i t ing w i ld e i nko rn whea t a nd f o r e a r ly whea t a nd b a r ley c u l t iva t ion u nde r v a r ious y i e ld c ond i t ions a re t hen c a lcu la ted . T hese a re c ompa red w i th r e turn r a tes c a lcu la ted f o rt he h and t i lage o fm i l le t a nd s o rghum i n Wes t A f r ica . C e rea ls wh ich d o n o t e xh ib i t c ompa ra t ive ly h i gh p os tencoun te r r e turn r a tes f o r t he i r e xp lo i ta t ion i nn a tu ra l s tands wou ld h ave b een u n l ike ly c and ida tes f o r e a r ly d omes t ica t ion s i nce t he a dd i t iona l l a bo r c os ts o f c u l t iva t ion wou ld f u r the r d im in ish t he i r r a tes o f e nergy r e tu rn . T he h a rves t ing o f w i ld c e rea l s tands w ou ld g ene ra l ly b e mo re e ne rgy e f ic ien tt han t he i re a r ly d ry fa rm ing c u l t iva t ion , b u t t h is r e la t ionsh ip wou ld b e r eve rsed i ft hey w e re c u l t iva ted i nt hose c i rcumsc r ibed e nv i ronmen ts whe re f l ood r ecess ion o ro the rf o rms o fn a tu ra l i r iga t ion w e re c oup led w i th r i ch s o i ls . Wh i le t o ta l 3

c e rea l y i e lds d o i n c rease a s af unc t ion o ft he amoun t o fl a nd c u l t iva ted , r e turn r a tes r ema in r e la t ive ly c ons tan t a s ar e su l t o fc ompa rab le l a bo r r equ i remen ts p e r u n i to fl a nd , p e rd ens i ty o f c rop h a rves ted , a nd p e r u n i t o f g ra in p rocessed o r s t o red . E xc lud ing t empo ra l v a r ia t ions i n a va i lab le w a te r , i ti st he q ua l i ty o f l a nd c u l t iva ted , n o t t h e q uan t i ty , t ha t d i rec t ly a f fec ts mean c e rea lc u l t iva t ion r e tu rn r a tes . I nC hap te r 7 ,t he e vo lu t iona ry i mp l ica t ions o f r e la t ive e f f ic iency i ne ne rgy c ap ture a re a pp l ied t ot he p as to ra l a nd c e rea l r e turn r a te d a ta g ene ra ted i nC hap te rs 5 a nd 6 i no rde r t o d eve lop a rchaeo log ica l ly t e s tab le h ypo theses c once rn ing t he s pa t ia l a nd t empo ra l e vo lu t ion o f a l te rna t ive p as to ra l a nd c e rea l s ubs is tence o p t ions . P red ic t ions d e r ived f r om t he r e la t ive e ne rge t ics o f p as to ra l a l te rna t ives c on t rad ic t c ur ren t a rchaeo log ica l i n terp re ta t ions wh ich s ugges tt ha t s ma l ls tock w e re d omes t ica ted i nt he N ea rE as ta nd N o r th A f r ica p r io rt oc ame ls a nd c a t t le . T he n a tu re o ft h is s t r ik ing i n cons is tency i ss ubsequen t ly e xp lo red . T he t empo ra l p a radox o f c ame l d omes t ica t ion i sa dd ressed b y a sk ing t he f o l low ing q ues t ion : U nde r w ha t c ond i t ions s hou ld d romeda ry c ame ls b e al a te d omes t ica te r e la t ive t o a l te rna t ive a n ima l o r c e rea l o p t ions? T he c ame l r ep roduc t ion a nd mor ta l i ty d a ta p resen ted i n A ppend ix A a nd c ompa ra t ive d a ta o nt he a t t r ibu tes o fh e rd s pec ies a s p resen ted i nC hap te r 4 a nd C hap te r 5s how t ha t u n l ike t he h usbandry o f c a t t le , s heep , a nd g oa ts , r e tu rn r a tes f o r d romeda ry c ame l h usband ry w ou ld b e s i gn i f ican t ly l owe ri nw e t /co ld o rw e t /wa rm e nv i ronmen ts . S ince t he mean mea t p l us m i lk r e tu rn r a tes e xpec ted f o r d romeda ry c ame l h usbandry i n d eser t e nv i ronmen ts w ou ld g ene ra l ly e qua l o re xceed t he m ean r e turn r a tes f o re i the r whea t o r b a r ley c u l t iva t ion o n ma rg ina l d ry fa rm ing l a nds , o r t hose e xpec ted w i th a l te rna t ive p as to ra l s t ra teg ies b ased u pon c a t t le , s heep o rg oa ts , t he l a te a ppea rance o fc ame l h usband ry r e la t ive t oa l te rna t ive a g r icu l tu ra l a nd p as to ra l o p t ions w ou ld o n ly make s ense i ft he e xpans ion o fx e r ic e nv i ronmen ts was a l so l a te . T h is e xpec ta t ion i s m e t i n b o th t he a rchaeo log ica l a nd p a leoenv i ronmen ta l r e co rds o ft h e N ea rE as t . H oweve r , w h i le t he l a te a ppea rance o f c ame l s t ra teg ies makes s ense o n t he b as is o f t empo ra l a nd s pa t ia l e nv i ronmen ta l t r ends , t he s eem ing ly l a te a ppea rance o fc a t t le s t ra teg ies c anno t b e e xp la ined i nt he s ame way b ecause t he e nv i ronmen ta l r equ i remen ts o fc a t t le a re s im i la rt ot hose o fs heep a nd g oa ts . E xp lana t ions f o rt h is a ppa ren t s equen t ia l d i spa r i ty wh ich a ppea l t ob ehav io ra l a nd s i ze v a r ia t ions b e tween t hese s pec ies a s r e la ted t ot he h uman a b i l i ty t oe f fec t ive ly c on t ro l p as to ra l h e rds c an a l so b ed i sm issed o nt he b as is o ft he d a ta p r esen ted i n C hap te r2 . T he a va i lab le emp i r ica l e v idence i st he re fo re r e exam ined , a nd t he ' a ssump t ions emp loyed i nc u r ren t i n te rp re ta t ions o f t ha t e v idence a re q ues t ioned . T he p r inc ip le e v idence u sed t o s ugges t t he e a r l ie r d omes t ica t ion o f s heep a nd g oa ts i n vo lves t he a ge -sex b i as a ppa ren t i n s pec i f ic f auna l d epos i ts a nd c er ta in mo rpho log ica l c r i te r ia . I np a r t icu la r , i th as b een a ssumed t ha t morpho log ica l ly " w i ld " a so pposed t o" domes t ica ted "t r a i ts ( p r ima r i ly i n vo lv ing r e la t ive s i ze ) a re n ecessa r i ly c o r re la ted w i th p a leoeconom ic s ys tems ( h un t ing v s . h e rd ing ) . W i thou t t hese a ssump t ions , h e re s hown t ob ef l awed , t he c ur ren t d a ta a r e i n conc lus ive o n t he r e la t ive t im ing o fc a t t le , s heep a nd g oa t d omes t ica t ion . S ugges t ions a re m ade f o rf u tu re r esea rch a dd ress ing t h is i s sue . T urn ing t o mean c e rea l r e tu rn r a tes , i ti sn o ted t ha t t h e a ppa ren t p redom inance o fb a r ley o ve rw hea t a st h e p re fe r red c r op o fp r eh is to ry makes s ense o na n e ne rge t ic b as is . S im i la r ly , t he a rchaeo log ica l r eco rd s hows t ha t t he c i rcumsc r ibed m ic roenv i ronmen ts whe re h i gh c e rea l r e tu rn r a tes w ere p oss ib le w e re t he f o cus o fe a r ly c e rea l p roduc t ion . H oweve r , m ean r e tu rn r a tes h i ghe r t han a l f ood p roduc t ion a l te rna t ives a re a l so e xpec ted t o h ave l e d t o i n tense c ompe t i t ion o ve r t hose g eog raph ica l ly c i rcumsc r ibed e nv i ronmen ts . Ap o ten t ia l l i nk b e tween h uman d emog raph ic s h i f ts d ur ing t he H o locene e poch , h ab i ta t s e lec t ion o n t he b as is o f p o ten t ia l r e turn r a tes , a nd e conom ic d e fendab i l i ty i st hen u sed t oe xp lo re a rchaeo log ica l d a ta o nt he t empo ra l a nd s pa t ia l e vo lu t ion o ff ood p roduc t ion a nd i n c reas ing ly c omp lex s oc ie t ies . E ne rge t ic c ompa r isons b e tween c e rea l a nd p as to ra l s ubs is tence o p t ions s how t ha t when c a lo r ies f r om m i lk a re e xc luded f r om p as to ra l p roduc t ion , t h e c u l t iva t ion o f whea t o rb a r ley o n ma rg ina l d ry fa rm ing l a nds g ene ra l ly r e su l ts i n h i ghe r m ean r e turn r a tes t han a l p as tora l s t ra teg ies . H oweve r ,w hen c a lo r ies f r om m i lk a re i n c luded i np as to ra l p roduc t ion , o n ly t he mean r e turn r a tes f o rw hea t o r b a r ley c u l t iva ted i no p t ima l c e rea l p roduc ing e nv i ronmen ts g ene ra l ly 4

e xceed t hose o b ta ined i np as to ra l s t ra teg ies , w i th t he s o le e xcep t ion o f g oa t h e rd ing . T he i n c reas ing r e turn r a tes e xpec ted f o rc e rea l c u l t iva t ion u nde r c hang ing t echno log ica l c ond i t ions f u r the r s ugges t t ha t mea t p l us m i lk p as to ra l s t ra teg ies w ou ld n o t h ave b ecome ma rg ina l t o a ve rage c e rea l s ubs is tence o p t ions u n t i lt he a dven t o fa n ima lt r ac t ion t i lage . Two mu tua l ly e xc lus ive a nd a rchaeo log ica l ly t e s tab le h ypo theses c once rn ing t he s pa t ia l a nd t empo ra l e vo lu t ion o fe a r ly p as to ra l a nd a g r icu l tu ra l f ood p roduc t ion a l terna t ives i nt he N ea r E as t a nd N or th A f r ica a re d e r ived f r om t hese r e su l ts . T hese a re : ( 1 ) I fm i lk p roduc t ion was n o t s i gn i f ican t i nt he e vo lu t ion o fe a r ly p as to ra l s t ra teg ies , a rchaeo log ica l e v idence s hou ld r e f lec t t he a g r icu l tura l e xp lo i ta t ion o f ma rg ina l d ry fa rm ing e nv i ronmen ts p r io r t o t he a ppea rance o f a l te rna t ive p as to ra l s t ra teg ies , a nd ( 2 ) I fm i lk p roduc t ion was s i gn i f ican t i nt he e vo lu t ion o fe a r ly p as to ra l s t ra teg ies , a rchaeo log ica l e v idence s hou ld r e f lec t t he a g r icu l tu ra l e xp lo i ta t ion o f o n ly o p t ima l d ry fa rm ing a nd n a tura l ly i r iga ted c e rea l e nv i ronmen ts p r io r t o t he a ppea rance o f a l te rna t ive p as to ra l s t ra teg ies . T he e xp lo i ta t ion o f ma rg ina l d ry fa rm ing l a nds s hou ld a ppea r a f te r p as tora l s t ra teg ies a re i np lace . On ly t he f i rs t p red ic t ion i sc ons is ten t w i th t he w ide ly f avo red ma rg ina l i ty a nd p roduc t iv i ty p e r u n i t a rea h ypo theses wh ich s ugges t t ha t p as to ra l s t ra teg ies n ecessa r i ly e vo lved a s a dap ta t ions t oe nv i ronmen ts ma rg ina l f o ra g r icu l tu re . C ompa r isons w i th t he H o locene e poch a rchaeo log ica l r eco rds o ft he N ea r E as t a nd N or th A f r ica r evea l t ha t wh i le t he f i r s t p red ic t ion f a i ls t oa ccoun t f o rt he emp i r ica l r eco rd , t he s econd p red ic t ion d oes . T he r e su l ts s uppor t t he p robab le s i gn i f icance o f m i lk p roduc t ion i n t he e vo lu t ion o f e a r ly p as to ra l s t ra teg ies , a nd f u r the r s ugges t t ha t i ti s u n l ike ly t ha t p as to ra l s t ra teg ies n ecessa r i ly e vo lved a s a dap ta t ions t oe nv i ronmen ts ma rg ina lf o ra g r icu l tu re . Ma jo r a spec ts o ft he r e su l ts o b ta ined i nt h is r e sea rch a re t hen s umma r ized . I ti sn o ted t ha t t he t heo re t ica l a pp roach a dop ted h e re g ene ra tes u se fu l b od ies o fq uan t i ta t ive d a ta , b r ings i n to q ues t ion p rev ious ly h e ld i n te rp re t ive a ssump t ions , o f fe rs c oun te r in tu i t ive e xpec ta t ions , s ugges ts n ew d i rec t ions f o r s ubs tan t ive r esea rch , a nd b e t te r a ccoun ts f o r t he a va i lab le emp i r ica l d a ta t han d o a l te rna t ive a pp roaches . F ur the r , t he u t i l i ty o ft he a pp roach u sed h e re f o r a dd ress ing i s sues o f e a r ly f ood p roduc t ion v as t ly e xpands t he r ange o f p o ten t ia l a pp l ica t ions a nd t e s ts o f s imp le e vo lu t ionary c os t -bene f i t mode ls wh ich c an c on t r ibu te t o e xp lana t ions o f s ubs is tence b ehav io r i n aw ide v ar ie ty o fh uman s i tua t ions a nd e nv i ronmen ts .

5

CHAPTER 1

T HE E VOLUT ION AND ADOPT ION OF FOOD P RODUCT ION S TRATEG IES : A THEORET ICAL FOCUS

I n t roduc t ion T he e a r ly e vo lu t ion a nd a dop t ion o f f ood p roduc ing s ubs is tence e conom ies i sw ide ly r ecogn ized a s o ne o ft he m os t s i gn i f ican t e ven ts i nh uman e x is tence . A f te r m i l l ions o fy ea rs a s f o rage rs , i th as t aken l e ss t han 1 5 ,000 y ea rs t o b r ing f o rag ing s ubs is tence s t ra teg ies t o t he b r ink o fe x t inc t ion i n awo r ld n ow d om ina ted b y f ood p roduc ing s oc ie t ies . I r on ica l ly , t he s ea rch f o ra n u nders tand ing o fw hy t h is s h i f t f r om f o rag ing t of ood p roduc t ion t ook p l ace , why i td i d n o t h appen u n t i l s o l a te ap o in t i nh uman e x is tence , a nd w hy i to ccu r red whe re i td i d , h as p roven t o b e a n e lus ive e ndeavo r , a l though s eve ra l g ene ra t ions o f s cho la rs h ave a ccep ted t he c ha l lenge . T he p resen t s t udy a t temp ts t of o rge o ne mo re l i nk i nt h is l o ng c ha in o f i n ves t iga t ion b y e xp lor ing t he p o ten t ia l o f u n i fo rm i ta r ian p r inc ip les d e r ived f r om t he f i e ld o f b ehav iora l e co logy f o r a dd ress ing t he b ehav io ra l c omponen t o ft he f o rage r / food p roduce rt r ans i t ion . A f ter many y ea rs o fe a r ly r esea rch d i rec ted a ti d en t i fy ing t he s pa t ia l a nd t empo ra l o r ig ins o f f ood p roduc t ion , mode rn r e sea rch h as i n c reas ing ly f ocussed u pon t he c omp lex e nv i ronmen ta l a nd b ehav io ra l p rocesses wh ich p resumab ly l e d t o t he f o rage r / food p roducer t r ans i t ion . Gene ra l ly a ccep t ing t ha t h uman b ehav io r h as e vo lved i nr esponse t o c hanges i nt he p hys ica l a nd s oc ia l e nv i ronmen t , s cho la rs h ave i s o la ted e nv i ronmen ta l s e t t ing , c l ima t ic v a r ia t ions , d emog raph ic s h i f ts a nd p opu la t ion g row th a s p r ima ry f a c to rs s i gn i f ican t i nt he p rocess b y wh ich f ood p roduc t ion s t ra teg ies e vo lved . S eve ra l p opu la r mode ls c ur ren t ly e x is t wh ich i n voke o ne o r mo re o ft hese f ac to rs t o o s tens ib ly a ccoun t f o r t he e vo lu t ion o ff ood p roduc t ion ( e .g . , B in fo rd 1 968 ; C h i Ide 1 936 , 1 952 ; C la rk 1 976 ; C ohen 1 977 ; F l anne ry 1 969 , 1 973 ; H a r r is 1 977 ; Moo re 1 985 :48 -49 ; Sm i th a nd Y oung 1 972 ; W r igh t 1 968 , 1 976 , 1 977 ) . U n for tuna te ly , i twou ld s eem t ha t w h i le we h ave i d en t i f ied a nd s t ressed t he c ond i t ions u nde r wh ich t he e vo lu t ion o f f ood p roduc t ion m igh t h ave t aken p lace , we h ave f a i led t o a dequa te ly a dd ress t he u nder ly ing p r inc ip les a f fec t ing s ubs is tence d ec is ions i ng ene ra l ( Moo re 1 985 ) . I na l o f t hese p r io r s t ud ies , i ti st he p resumed p roduc t iv i ty o r r e l iab i l i ty o ff ood p roduc t ion o p t ions wh ich h ave b een a ppea led t oi ns eek ing ac ausa l b ehav io ra l mechan ism t oa ccoun t f o r t he e vo lu t ion o ra dop t ion o fs uch s ubs is tence s t ra teg ies . I ti su sua l ly a ssumed t ha tt he a b i l i ty t o e nsu re o r i n c rease t he p roduc t iv i ty a nd r e l iab i l i ty o f p lan t a nd a n ima l r esources wou ld n ecessa r i ly h ave b een a d es i rab le g oa l u nde r c ond i t ions o f e nv i ronmen ta l o r d emog raph ic s t ress . F ur the r , i th as b een s ugges ted t ha t t he g rea te r p roduc t iv i ty p e r u n i t a rea wh ich i so f ten a ssoc ia ted w i th a g r icu l tura l s t ra teg ies n ecessa r i ly i mp l ies t ha t p as to ra l s ubs is tence mus t h ave e vo lved a s a n a dap ta t ion t o e nv i ronmen ts ma rg ina l f o r a g r icu l tu re ( e .g . , K hazanov 1 984 ; Ga r ra rd 1 984 ; S ah l ins 1 968 ; S poone r 1 97 1 , 1 973 ) . A s a ppea l ing a s s uch p os tu la tes may b e , t hey d o n o t a dd ress t he d ec is ion mak ing w h ich was u ndoub tab ly i n vo lved i n mak ing c ho ices b e tween s eve ra l a l te rna t ive , a nd t he re fo re c ompe t ing , s ubs is tence o p t ions , e ach o fw h ich w ou ld h ave h ad n o t o n ly a ssoc ia ted b ene f i ts , b u t a ssoc ia ted c os ts a s w e l l . On ly a s e lec t n umbe r o fp l an ts a nd a n ima ls b ecame t he f ocus o f h uman f ood p roduc t ion , e ven t hough s eve ra l s pec ies o fe ach w e re o f ten a va i lab le i nt he s ame e nv i ronmen ts . Wha t i sl a ck ing i s ac ommon c oncep tua l f r amewo rk t oa ccoun tf o rt he d i f fe ren t ia l e xp lo i ta t ion a nd man ipu la t ion o f t he p l an t a nd a n ima l r esources a va i lab le i n s pa t ia l ly a nd t empo ra l ly v a r iab le e nv i ronmen ts . I nt he a bsence o f at heore t ica l a pp roach t o u nde rs tand ing t he u nder ly ing p r inc ip les a f fec t ing s ubs is tence d ec is ions , e ach i n s tance o fs ubs is tence c hange mus t n ecessa r i ly a ppea r t o b e a s u n ique a s t he r eg ions o r r esou rces i n vo lved . N o t s urp r is ing ly , mode ls p rev ious ly p roposed f o r t he way i nw h ich f ood p roduc t ion b egan i n v a r ious r eg ions o f t he w o r ld a re

r ecogn ized a s b e ing a s d i s t inc tf r om o ne a no the r a st he a r eas o r r esou rces t o w h ich t hey r e fe r ( F lanne ry 1 973 ; H a r lan 1 977 :359 -360 ) . U n fo r tuna te ly , t h is s i tua t ion h as l e d s eve ra l s cho la rs t o c onc lude t ha t t he e vo lu t ion a nd a dop t ion o f f ood p roduc t ion s t ra teg ies i n v a r ious r eg ions o ccur red i nd i f fe ren t ways f o r d i f fe ren t r easons , a nd t ha t u n iversa l o r u n i fo rm i ta r ian p rocesses we re n o t i n vo lved ( e .g . , B anks 1 984 :245 ; F l anne ry 1 973 :272 ; H a r lan 1 977 :358 -360 ) . l th as b een s ugges ted t ha t a rchaeo log is ts s hou ld n ow e schew s eek ing a t heo re t ica l u nde rs tand ing o fw hy f ood p roduc t ion o ccu r red i nd e fe rence t os imp ly d ocumen t ing t he p rocesses o fc hange w h ich w e re p resumab ly i n vo lved ( Moo re 1 985 :48 -49 ) . T he p resen t s t udy r e jec ts t hese n ega t ive c onc lus ions a s i l founded . T he f a i lure t o d eve lop a c ompe l l ing c oncep tua l f r amewo rk f o r u nde rs tand ing s ubs is tence d ec is ions i sn o t e qu iva len t t o d emons t ra t ing t ha t u n ive rsa l o r u n i fo rm i ta r ian p rocesses a f fec t ing s ubs is tence b ehav ior d o n o t e x is t , n o r i si ts u f f ic ien t t o s ugges t t ha t t he s ea rch f o r s uch p r inc ip les i sn o l o nge r" p ro f i tab le . " T he b as ic p rem ise o ft h is s t udy i st ha t s i gn i f ican t a dvances i nu nde rs tand ing t he e vo lu t ion a nd a dop t ion o ff ood p roduc t ion s t ra teg ies a re u n l ike ly u n less man -an ima l -p lan t r e la t ionsh ips a re i n ves t iga ted w i th in t he c on tex t o f g ene ra l t heo r ies wh ich d o a dd ress t he u nder ly ing p r inc ip les a f fec t ing s ubs is tence b ehav io r .

T he P o ten t ia l o fa n E vo lu t ionary P e rspec t ive T he t heo re t ica l f ocus o f t h is s t udy i sd rawn f r om t h e b as ic p os tu la te o f n eo -Da rw in ian e co logy ( behav io ra l e co logy , e vo lu t ionar y e co logy , s oc iob io logy ) t ha t t he b ehav ior o f a l o rgan isms i ss haped b y n a tura l s e lec t ion . R icha rd D . A lexande r , i np a r t icu la r , h as b een a n o u tspoken p roponen t f o r t he a pp l icab i l i ty o f n a tu ra l s e lec t ion i na dd ress ing t he e vo lu t ion o f h uman b ehav ior ( e .g . , A lexande r 1 978a , 1 978b , 1 979a , 1 979b ) . T he mos t f r equen t ly r a ised o b jec t ion t ot h is p e rspec t ive i st ha t h uman b ehav io r i sf a rt oo v a r iab le , a nd p ar t icu la r ly f a rt oo s ub jec t t oc hange t h rough l e a rn ing , t ob e e xp la ined i ng ene t ic t e rms t h rough n a tu ra l s e lec t ion ( e .g . , S ah l ins 1 976 ) . T he g ene ra l t endency o f s uch o b jec t ions i st o c oncep tua l ize " cu l tu ra l b ehav io rs " a s b e ing d i s t inc t f r om t he r es t o f t he h uman p heno type , a nd t hus n o t s ub jec t t o n a tura l s e lec t ion . T h is i sg ene ra l ly a ccomp l ished b y a n a ppea l t o t he d i f fe rences b e tween i n dependen t modes o ft r a i t t r ansm iss ion , s pec i f ica l ly v i ewed a s g ene t ic v s . l e a rned ( f o r ad iscuss ion o f why n o g ene t ic t r a i t e x is ts o r d eve lops i n dependen t o f i t s p as t o r p resen t e nv i ronmen t , s ee A lexande r 1 978a :155 , 1 978b :251-252 ; F l inn a nd A lexande r 1 982 :388 ) . S uch a rgu men ts ma in ta in t ha t wh i le e a r ly h om in id b ehav io rs w e re ( o r" m igh t h ave b een " ) s ub jec tt oe vo lu t ion b y n a tura l s e lec t ion , o nce t he g ene t ic c apac i ty e vo lved f o r l ea rn ing , behav io ra l t r a i ts f r om n on re la ted i n d iv idua ls , s ubsequen t b ehav io ra l c hanges h ave b een i n c reas ing ly i n dependen t o f c hange i nt he g ene t ic c apac i ty f o r l e a rn ing a nd h ence i n c reas ing ly i n dependen t o f n a tu ra l s e lec t ion . H ence , i ti sa rgued t ha t a ts ome p o in t i nh om in id e vo lu t ion , t he b ehav iora l c omponen t o f t he h uman p heno type ( as d i s t inc t f r om i t s p hys io log ica l a nd morpho log ica l c omponen ts ) b ecame " uncoup led " f r om n a tu ra l s e lec t ion s uch t ha t b ehav io ra l c hange c ou ld o ccur i nt he a bsence o fa ny r e la ted g ene t ic v a r ia t ion ( Ha r r is 1 979 :12 1125 ) . T h is a rgu men t h as b een u sed b y s ome s cho la rs t oo u t r igh t r e jec t a n e vo lu t iona ry a pp roach t o h uman b ehav io r ( e .g . , S ah l ins 1 976 ) , w h i le o the rs h ave u sed i tt op os i t as epa ra t ion b e tween " na tura l " a nd " cu l tu ra l " s e lec t ion ( e .g . , C ava l l i -S fo rza a nd F e ldman 1 981 ; D u rham 1 976 , 1 979 , 1 982 ) . H oweve r , t h is w ou ld s eem t ob e am is lead ing v i ew o ft he p heno type wh ich i g no res b o th t he p r inc ip les o f o rgan ic e vo lu t ion a s we l l a s t he u nder ly ing r easons f o r w hy t he p heno type ( and p ar t icu la r ly i t s b ehav io ra l c omponen t ) s hou ld h ave e vo lved a ta l . T he b ehav io ra l c omponen t o ft he p heno type o fa l o rgan isms e xh ib i t ing f l ex ib le b ehav io rs d i fe rs f r om t he p hys io log ica l a nd mo rpho log ica l c omponen ts i nt he d eg ree t o wh ich i tc an v a ry i nt he a bsence o f g ene t ic c hange . T h is i st he r eason f o r w hy f l ex ib le b ehav io rs w ou ld h ave e vo lved i nt he f i r s t p l ace . B ehav io ra lf l ex ib i l i ty a nd t he f u r the rc apac i ty f o rl e a rn ing a low f o r mo re r ap id r esponses t o v a r ia t ions i nt he p hys ica l a nd s oc ia l e nv i ronmen t t han i sp oss ib le t h rough g ene t ic v a r ia t ions ( A lexande r 1 979a :66 -67 , 1 979b :75 -76 ; F l inn a nd A lexande r 1 982 :388 3 90 ) , w i th d i rec t r epe rcuss ions o n t he r e la t ive r ep roduc t ive s uccess o ft he o rgan isms wh ich b ea rs uch t r a i ts .

8

" Cu l ture " i s at e rm w h ich d es igna tes t he p a r t icu la r a nd e l abo ra te s ys tem o f b ehav io rs a ssoc ia ted w i th h umans , b u t i td oes n o t i mp ly t ha t ad i s t inc t f o rm o f s e lec t ion i sn ecessa r i ly i n vo lved i n t he u se , mod i f ica t ion , o r t r ansm iss ion o f h uman b ehav io ra l t r a i ts . T he g ene t ic c apac i ty f o r " cu l tura l ly t r ansm i t ted b ehav io rs " c ou ld n o t e vo lve a nd b e ma in ta ined u n less , i n g ene ra l , t he r ange o ff l ex ib le b ehav io rs wh ich i ta lows f avo rab ly a fec ts t he r e la t ive r ep roduc t ive s uccess o f t he o rgan ism w h ich b ea rs t h is s pec i f ic g ene t ic c apac i ty . A s n o ted b y F l inn a nd A lexande r ( 1982 :390 ) : P heno types - a nd b ehav io r af o r t ior i- a re ways o fa dop t ing d i f fe ren t s t ra teg ies i nd i f fe ren t e nv i ronmen ts u s ing t he s ame s e ts o f g enes ; a nd p heno types a re p roduc ts o f n a tura l s e lec t ion . C omp lex a l te rna t ive p heno typ ic s t ra teg ies c o r re la ted w i th g ene t ic u n i fo rm i ty a re n o t e v idence a ga ins t t he e f f icacy o f s e lec t ion , b u t i n s tead a re e v idence o f ah i s to ry o fp ower fu l s e lec t ion i n vo lv ing u np red ic tab i l i ty i ne nv i ronmen ta l s h i f ts o ro sc i l la t ions . T he b i o log ica l l i t e ra tu re i s f i led w i th e xamp les . We c anno t o ve remphas ize t he i mpor tance o ft h is p o in t f o r u nde rs tand ing t he i n f luence n a tu ra l s e lec t ion h as h ad o n t he e vo lu t ion o f h uman l ea rn ing , a nd c onsequen t ly o n c u l tura l e vo lu t ion . T he a b i l i ty t o l e a rn , a nd d i rec t iona l t endenc ies i n l e a rn ing o f wha teve r s o r t , a re s ure ly p roduc ts o f n a tura l s e lec t ion . L ea rn ing mus t r e su l t f r om t he a c t ion o f g enes t ha t s pec i fy t ha t t he o rgan ism s hou ld d eve lop i ns uch af ash ion a st ob e a b le t or e spond a dap t ive ly t o a ny s i tua t ion f r om s ome a r ray o fl i ke ly s i tua t ions when t he p a r t icu la ro ne t ha t w i l l e ven tua te c anno t b e i d en t i f ied f a ra head o ft ime . Wha t h as b een f avo red b y n a tura l s e lec t ion i n g enera l i st he a b i l i ty t o r espond a dap t ive ly t o v a ry ing e nv ironmen ta l c ond i t ions . l ti sn o t e xpec ted t ha t t he p ar t icu la r b ehav io rs o f a ny o rgan ism a re n ecessa r i ly c oded i n p a r t icu la r g enes . R a the r , i ti se xpec ted t ha t o rgan isms h ave e vo lved w i th t he g ene t ica l ly d e te rm ined c apac i ty t o a d jus t t he i r b ehav io r t o c hang ing c i rcums tances i n av a r ie ty o f ways ( i nc lud ing l ea rn ing ) i ns uch af ash ion a s t o max im ize t he i r r ep roduc t ive f i t ness . A s r epea ted ly s t ressed b y A lexande r ( 1978a , 1 978b , 1 979b :94 -102 ) , t he e qua t ion wou ld b e , Genes + E nv i ron men t —› Behav ior , a nd t he r o le o f e i the r f ac to r c anno t b e v a l id ly u nderp layed o r o ve remphas ized r e la t ive t ot he o the r . A s a pp l ied s pec i f ica l ly t o t he b ehav io rs o f h umans , a n e xp l ic i t ly e vo lu t iona ry a pp roach a ssumes t ha t h umans h ave b een s e lec ted t o b e e no rmous ly o ppor tun is t ic i nt he i r r e sponses t oc hange i nt he s oc ia l a nd p hys ica l e nv i ronmen t , a nd t ha t s uch b ehav io ra l a d jus tmen ts a re n o t t he p roduc t o f s hor t te rm g ene t ic c hange . l ti se xpec ted t ha t h umans h ave b een s e lec ted t o s e ize o ppor tun i t ies t ha t a re l i ke ly t o e nhance t he i r r e la t ive r ep roduc t ive f i tness . l t wou ld t he re fo re s eem t ha t a n e xp l ic i t ly e vo lu t ionary a pp roach t o i s sues o f h uman f eed ing b ehav io r wou ld p o ten t ia l ly p rov ide ap ower fu l u n i fo rm i ta r ian p e rspec t ive f o r a dd ress ing t he u nder ly ing p r inc ip les a f fec t ing s ubs is tence d ec is ions . R ecen t ly , p red ic t ive b ehav io ra l mode ls d e r ived f r om o p t ima l f o rag ing t heo ry a s d eve loped b y e vo lu t iona ry b i o log is ts a nd e co log is ts ( Cha rnov 1 976a , 1 976b ; C ha rnov a nd O r ians 1 973 ; Em len 1 966 ; K rebs 1 978 ; K rebs a nd D av ies 1 985 :49 -68 ; MacA r thur a nd P ianka 1 966 ; P yke e t a l . 1 977 ; S choener 1 97 1) h ave b een s uccess fu l ly u sed t o i l um ina te v a r ious a spec ts o f f o rag ing b ehav io r among d i ve rse h un te r -ga the re r p opu la t ions ( e .g . , H awkes a nd O 'Conne l l 1 981 ; H awkes e ta l . 1 982 ; H i l l 1 983 ; H i l l e ta l . 1 986 ; O 'Conne l l a nd H awkes 1 98 1 , 1 984 ; S imms 1 984 , 1 987 ; Sm i th 1 983 ; W in te rha lde r 1 981a , 1 981b ) . T he u t i l i ty o f s uch mode ls f o r ab e t ter u nde rs tand ing o f i n sec t ivo r y i nh uman d i e ts h as a l so b een s ugges ted ( Ha r r is 1 985 : 168 -169 ) . T hese p red ic t ive mode ls i n voke d ec is ion mak ing b ased u pon t he r e la t ive e ne rge t ics ( ca lo r ic y i e ld / t ime i n ves ted ) o fe xp lo i t ing a l te rna t ive f ood r esou rces . T he b as ic p rem ise o f t h is a pp roach i s t ha t s i nce e ne rgy d e r ived f r om f ood f ue ls r ep roduc t ion ( Lew in 1 982 ) , o rgan isms wh ich max im ize t he i r f o rag ing e f f ic iency , a s measu red b y t he i r r a te o f e nergy c ap tu re , w ou ld g enera l ly e n joy a r e la t ive ly h i ghe r r ep roduc t ive f i t ness ( MacA r thur a nd P ianka 1 966 ; S choene r 1 97 1 ; C ha rnov a nd O r ians 1 973 ; P yke e ta l . 1 977 ) . l ti s e xpec ted t ha t f o rage rs w ho a re mo re e f f ic ien t wou ld g enera l ly b e a b le t oo b ta in mo re e ne rgy w i th in as pec i f ic p er iod o ft i me , wou ld r educe t he i r p e r iod o f e xposu re t ot he r i sks o ff o rag ing , a nd w ou ld t he re fore b e a b le t od evo te mo re t ime t oo the rf i t ness r e la ted a c t iv i t ies ( Sm i th 1 979 , 1 983 ) . A l l e l se b e ing e qua l , f o rag ing s t ra teg ies t ha t a re mo re e f f ic ien t i nr a tes o fe ne rgy c ap tu re

9

s hou ld b e f avo red b y n a tura l s e lec t ion , a nd w ou ld p e rs is t a nd s p read a tt he e xpense o ft hose t ha t a re l e ss e f ic ien t . Wh i le i th as b een s ugges ted t ha t p red ic t ive mode ls i n vok ing d ec is ion mak ing b ased u pon t he r e la t ive e ne rge t ics o fe xp lo i t ing a l te rna t ive f ood r e sou rces s hou ld h ave a b road a pp l icab i l i ty t oi s sues c once rn ing t he e vo lu t ion a nd a dop t ion o f f ood p roduc t ion s t ra teg ies ( Hawkes e ta l . 1 982 :395 ; H i l l e t a l . 1 986 ; O 'Conne l l a nd H awkes 1 981 : 116 ) , s ubs tan t ive a pp l ica t ions h ave b een f ew i nn umbe r ( e .g . , K eegan 1 986 ; R edd ing 1 98 1 , 1 983 ) . I ns ubsequen t c hap ters o ft h is manusc r ip t , a me thodo logy w i l l b e d eve loped f o r a dd ress ing s pa t ia l a nd t empo ra l i s sues o f e a r ly f ood p roduc t ion i n af ash ion c ons is ten t w i th t ha t emp loyed i ne vo lu t iona ry s t ud ies o f h uman f o rag ing b ehav io r .

B as ic E ne rge t ic Mode ls Two p red ic t ive b ehav io ra l mode ls d eve loped b y b ehav io ra l e co log is ts wh ich i n voke r a tes o fe ne rgy c ap tu re a re p a r t icu la r ly r e levan tt ot he p resen t s t udy . F i rs t , t he d ie t b read th o r o p t ima l d i e t mode l ( Cha rnov 1 976a ; C ha rnov a nd O r ians 1 973 ; Em len 1 966 ; MacA r thur a nd P ianka 1 966 ; S choener 1 971) p red ic ts w h ich r esou rces a f o rage r s hou ld e xp lo i t f r om a n a r ray o f a va i lab le r esou rces e ncoun te red a tr andom i ft he r a te o fe ne rgy c ap ture i sb e ing max im ized . I n i t ia l ly , a va i lab le r e sou rces may b e r anked a cco rd ing t ot h e r a t io o fe nergy r e tu rns e ach i t em p rov ides ( measu red i nc a lo r ies ) t ot he c os t ( o r h and l ing t i me ) o fa cqu i r ing a nd p rocess ing t ha t i t em o nce i th as b een e ncoun te red ( = i t s p os tencoun te r r e tu rn r a te ) . I faf o rage r i s max im iz ing t he r a te o fe ne rgy c ap tu re , ar esou rce i t em r andom ly e ncoun te red s hou ld b e t aken i fa nd o n ly i f i t s r ank ing ( r e tu rn r a te ) i se qua l t oo r g rea te r t han t he r e tu rns t o b e g a ined f o r s ea rch ing f o r , f i nd ing , a nd h and l ing a n i t em o fh i ghe r r ank d ur ing t he s ame p e r iod o ft ime . R esources w h ich f a i l t o mee tt h is c r i te r ion s hou ld n o t b et aken . I n amo re f o rma l s t a temen t ,f o ra l i t ems t ob e i n c luded i nt he d i e t : E i >

E p

h i Whe re

E i =t o ta l e ne rgy ( k ca l ) o fe ach r e sou rce i h i =c o l lec t ing a nd p rocess ing t i me ( g i+p i ) f o re ach r e sou rce i E p =E / (Ts+Tg+Tp ) E =t o ta l e ne rgy i nr esou rces g a the red p e rc o l lec to r T s =t o ta l s ea rch t ime i nt he r e sou rce p a tch T g =t o ta l g a the r ing t ime p e rc o l lec to rf o ra l r esou rces g a the red T p =t o ta l p rocess ing t i me p e rc o l lec to rf o r a l r e sou rces g a the red

A n o p t ima l d i e t s hou ld i n c lude a l p o ten t ia l r e sou rces w i th p os tencoun te r r e tu rn r a tes e qua l t o o r h i ghe r t han t he a ve rage r e tu rns f o r f o rag ing i ng ene ra l , a nd s hou ld e xc lude a l p o ten t ia l r esources w i th p os tencoun te r r e tu rn r a tes l owe r t han a ve rage f o rag ing r e turns . S ign i f ican t ly , w he the r o r n o t ap o ten t ia l r esou rce i si nt he o p t ima l d i e t d oes n o t d epend u pon i t s o wn a bundance , b u t r a the r o n t he a bundance ( o r mo re p rec ise ly t he e ncoun ter r a te ) o f h i ghe r r ank ing r e sources . H igh ranked r e sou rces s hou ld r ema in i nt he d ie t r ega rd less o fh ow r a re t h ey b ecome . H oweve r , i ft he e ncoun te r r a te f o r h i gh ranked r e sources f a l ls , a ve rage f orag ing r e turn r a tes wou ld a l so f a l l a nd l owe r r anked r esources wou ld e n ter t he d ie t . T hese e xpec ta t ions a re s umma r ized i nF i gu re 1 . I n t he p resen t s t udy , t he d ie t b read th o r o p t ima l d i e t mode l i s s pec i f ica l ly u se fu l f o r c oncep tua l iz ing t he c hanges i nh uman d i e t wh ich a ppa ren t ly o ccu r red i nt he p os t -P le is tocene p e r iod . A s n o ted b y H awkes e ta l . ( 1982 :395 ) a nd H i l l e ta l . ( 1986 ) , t he c o inc idence b e tween t he a ppa ren t d ec l ine o r e x t inc t ion o f l a rge mamma l p opu la t ions a nd o ther f auna a to r n ea rt he e nd o f t he P le is tocene a nd t he eme rgence o f " b road s pec trum " s ubs is tence e conom ies ( F lannery 1 969 ) i sc ons is ten t w i th e xpec ta t ions d e r ived f r om t he d ie t b read th o ro p t ima l d i e t mode l .

1 0

K ca l/hr

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A . T r iang les i n d ica te p rey t ypes a r ranged b y r ank , b ased o n r e turns o nce e ncoun te red . C i rc les ma rk r e turns g a ined f r om s ea rch ing f o r a nd h and l ing p rog ress ive ly l owe r r anked p rey .

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B . I f t he e ncoun ter r a te f o r h igh ranked p rey d ec reases , r e tu rns f r om f o rag ing f o r t hose p rey i t ems s hou ld a l so d ec rease . A s a verage r e turns f r om s ea rch ing f o r a nd h and l ing t hese r e sou rces f a l l b e low t hose g a ined f r om h and l ing l owe r ranked p rey , t he l a t te r s hou ld e n ter t he d i e t .

F i gu re 1 . D ie t b read th mode l ( adap ted f r om C ha rnov 1 976a ; O 'Conne l l a nd H awkes 1 984 ) . 1 1

Wh i le t he d ie t b read th o r o p t ima l d ie t mode l s pec i f ica l ly d ea ls w i th t he e xp lo i ta t ion o f r esou rces r andom ly e ncoun te red i n a" f ine g ra ined " e nv i ronmen t , p a tch -cho ice mode ls d ea l w i th t he e xpec ted s pa t ia l a nd t empo ra l e xp lo i ta t ion o f p a tchy e nv i ronmen ts whe re r esou rces a re c l umped , a ga in a ssum ing t ha tt he g oa lo ff o rag ing b ehav io ri st o max im ize t he r a te o fe ne rgy c ap tu re ( Cha rnov 1 976b ; C ha rnov a nd O r ians 1 973 ; MacA r thu r a nd P ianka 1 966 ; P yke e ta l . 1 977 ; S choene r 1 971) . H e re , i ti se xpec ted t ha t af o rage rw ou ld o pe ra te i nt ha t p a tch o r s e t o f p a tches t ha t p roduce t he b es t e ne rgy r e tu rns r e la t ive t ot he t i me c os ts o ft r ave l ing t ot he p a tch , s ea rch ing i t , a nd g a the r ing a nd p rocess ing t he r esou rces e ncoun te red t here . A f u r the r e xpec ta t ion o f s uch mode ls c once rns movemen t b e tween p a tches . A ssum ing t ha t t he f o rag ing p rocess d ep le tes t he r e source l e ve l i na ny p a tch t h rough t ime , i ti se xpec ted t ha t a f o rage rw i l l o n ly r ema in i nt ha t p a tch u n t i l t he r a te o fe ne rgy c ap tu re t ot i me i n ves ted f a l ls e qua l t ot he mean e ne rgy r e tu rn p e ru n i tt ime ( i nc lud ing t r ave lt i me ) o fa l a va i lab le p a tches ( Cha rnov 1 976b ) . A s emp loyed i nt he p resen t s t udy , i ti sp roposed t ha t u nde rd ec l in ing f o rag ing r e tu rn r a tes a t t he e nd o f t he P le is tocene , f ood p roduc t ion s t ra teg ies s hou ld h ave b een a dop ted w hen mean f o rag ing r e turn r a tes f e l l e qua l t o o r b e low t he mean r e turn r a te f o r a va i lab le f ood p roduc t ion o p t ions . T h is e xpec ted r e la t ionsh ip i sd ep ic ted i nF i gu re 2 .F ur the r , a va i lab le f ood p roduc t ion o p t ions s hou ld h ave b een a dop ted a cco rd ing t o t he i r r ank ing b y r e la t ive r e tu rn r a tes . F i na l ly , i ti se xpec ted t ha t v a r ia t ions i nt he s pa t ia l a nd t empo ra l e xp lo i ta t ion o fh ab i ta ts o r h ab i ta t t ypes s u i tab le f o rf ood p roduc t ion s hou ld h ave b een af unc t ion o ft he e ne rgy r e turns f o r f ood p roduc t ion e f for t w h ich c ou ld b e o b ta ined i ns pec i f ic l o ca t ions o r r eg ions . Mo re c omp lex i s sues r e la t ing t om ixed s ubs is tence s t ra teg ies w i th v a r ious c omb ina t ions o f s easona l f o rag ing a nd a g r icu l tu ra l o r p as to ra l o p t ions , o r v a r ious c omb ina t ions o f p lan t a nd a n ima l h usbandry i nf ood p roduc t ion s t ra teg ies , w i l l n o t b e a dd ressed i nt he p resen t s t udy . F o l low ing K eegan ( 1986 ) , i ti ss uspec ted t ha t t he d i e t b read th o r o p t ima l d ie t mode l may w e l l p rove u se fu l i nb e t te r u nde rs tand ing a g r icu l tu ra l s ys tems i n vo lv ing t he d i fe ren t ia l c u l t iva t ion o f v a r ious c rops , a nd b y e x tens ion , t he d i f fe ren t ia l h e rd ing o fv a r ious h e rd a n ima l c omb ina t ions i n p as to ra l s t ra teg ies . S ince t he s p read o ff ood p roduc t ion i n to E u rope a ppea rs t o h ave i n vo lved a" package " o ff i ve a n ima l d omes t ica tes ( ca t t le , s heep , g oa t , p i g a nd d og ) a s w e l l a s w hea t , b a r ley , l e n t i ls a nd s o o n ( Sebas t ian P ayne , p e rsona l c ommun ica t ion 1 987 ) , a n u nde rs tand ing o ft he e ne rge t ics o f m ixed s t ra teg ies i sc e r ta in ly d es i rab le . I n i t ia l ly , h owever , i ts eems mo re p ro f i tab le t ok eep t he i s sues a dd ressed a s s imp le , a nd a s e as i ly t es ted , a s p oss ib le . K ca l/h r P o int i n t ime when f ood p roduc t ion s hou ld e vo lve o r b e a dop ted

WE I

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1— h un t ing-ga the r ing

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p roduc t ion

1 T I ME F i gu re 2 . S ubs is tence s t ra tegy r e tu rn r a tes a nd t he e vo lu t ion o r a dop t ion o ff ood p roduc t ion o p t ions .

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I nt he f o l low ing c hap te rs o ft h is manusc r ip t , e ne rge t ic c os t a nd b ene f i t d a ta f o r s epa ra te p as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts , a nd a g r icu l tura l s t ra teg ies i n vo lv ing whea t a nd b a r ley , w i l l b e g ene ra ted . E ne rge t ic e xpec ta t ions d e r ived f r om t hese d a ta w i l l t hen b e u sed t o e xp lo re t he e x ten t t o w h ich s imp le e ne rgy o p t im iza t ion mode ls m igh t e nhance c u r ren t me thods o f a rchaeo log ica l i n fe rence c once rn ing t he f o rage r / food p roduce r t r ans i t ion i nt he N ea r E as t a nd N or th A f r ica . B e fo re p roceed ing , h oweve r , ab r ie f c ommen ta ry o n a n a l te rna t ive a pp roach t o i s sues o f e a r ly f ood p roduc t ion r ecen t ly p roposed b y D av id R indos ( 1980 , 1 984 ) i si no rde r .

Why D id H umans B ecome I n vo lved i nC oevo lu t iona ry R e la t ionsh ips W i th P lan ts a nd A n ima ls : A n I s sue W i thou t R ea l Mean ing? I n r ecen t works , D av id R indos ( 1980 , 1 984 ) h as a l so a rgued f o r t he u t i l i ty o f a n e vo lu t iona ry a pp roach t o i s sues o f f ood p roduc t ion . F ocuss ing s pec i f ica l ly u pon a g r icu l tura l s t ra teg ies , R indos h as d rawn a t ten t ion t ot he c oevo lu t iona ry i n te rac t ion b e tween h umans a nd p lan ts wh ich o ccur red b o th b e fo re a nd a f te r t he e vo lu t ion o f f ood p roduc t ion s t ra teg ies . I n p ar t icu la r , R indos s ugges ts t ha t t he d omes t ica t ion p rocess was a r esu l t o f c oevo lu t iona ry i n terac t ions o ccur r ing w i th in t he e nv i ronmen t o f ad eve lop ing h uman -p lan t mu tua l ism . Wh i le t h is r esea rch s ha res a n e vo lu t iona ry f ocus w i th t he wo rk o f R indos , t he re a re s i gn i f ican t d i f fe rences . A s w i th s eve ra l e a r l ie r h ypo theses c once rn ing t he e vo lu t ion o f f ood p roduc t ion , R indos a ppea ls t o p resumed i n c reases i n p lan t p roduc t iv i ty d ur ing t he d eve lop ing h uman -p lan t m u tua l ism i ns eek ing ac ausa l b ehav io ra l mechan ism t oa ccoun tf o rt he e vo lu t ion o ra dop t ion o f a gr icu l tura l s ubs is tence s t ra teg ies . Wh i le h e s t resses t he b ene f i ts d e r ived f r om p lan t h usband ry a nd c u l t iva t ion , t he c os ts o f s uch a c t iv i t ies a re g ene ra l ly i g no red . A ppea l ing t o c omp lex " pos i t ive a nd n ega t ive f eedback " mechan isms , h e p os i ts i n c reases i nw i ld p l an t y i e lds d ue t o i n i t ia l h usband ry p rac t ices w h ich a lowed f o r p opu la t ion g row th , wh ich i nt u rn i n c reased t he i n cen t ive f o r s pec ia l ized f ood p roduc t ion . T he s ubsequen t s h i f t t owa rd h i ghe r l e ve ls o f r e sou rce u t i l iza t ion a nd a ssoc ia ted i n c reases i nt he a bundance a nd y i e ld o fd omes t ica tes t hen f avo red t he t r ans i t ion t of u l l a g r icu l tu ra l s ubs is tence s t ra teg ies . Why p l an ts w e re i nt he d i e t , why t he s h i f t t ot he i r i n t ia l h usbandry w ou ld h ave o ccu r red , a nd why s ome p lan ts b u t n o t o the rs we re t he f ocus o ft h is s h i f t , a re n o t c ons ide red s i gn i f ican t i s sues . R indos i n s tead a ssumes t ha t h umans h ave a lways h ad g ene ra l ized d ie ts wh ich i n c luded p lan ts , a nd t ha tt he h uman -p lan t r e la t ionsh ip i ss imp ly " na tu ra l ' : g i ven t ha t amu tua l is t ic r e la t ionsh ip w i th p lan ts w ou ld n ecessa r i ly i n c rease t he c a r ry ing c apac i ty o f ar eg ion . A s h e s ays ( 1984 :14 1) Why h umans b egan t oe s tab l ish c oevo lu t iona ry r e la t ionsh ips w i th p lan ts i s aq ues t ion w i thou t r ea l mean ing . We m igh t a s w e l l a sk why c e r ta in a n ts e s tab l ished c oevo lu t iona ry r e la t ionsh ips w i th f ung i o rc er ta in b i rds w i th s pec i f ic f r u i ts . T he r e la t ionsh ips we re e s tab l ished a s ar esu l t o f t he max im iza t ion o f f i tness i ng i ven s i tua t ions i nt ime a nd s pace ; t hey w e re n e i the r i n ev i tab le n o r d es i rab le , b u t me re ly h appened . T he p resen t s t udy r e jec ts t h is p os i t ion a s c oncep tua l ly i n cons is ten t . I ft he b ehav io r o f a l o rgan isms i ss haped b y n a tu ra l s e lec t ion , ma jo rc hanges i np a t te rns o ff eed ing b ehav io rd o n o t s imp ly h appen , b u t a re t he r e su l t o fd i rec t iona l t endenc ies a s p roduc ts o f n a tura l s e lec t ion . I f , h oweve r , ma jo r c hanges i nf eed ing b ehav io r d o s imp ly h appen w i thou t d i rec t iona l i ty , t hen a n a ppea l t o e vo lu t iona ry t heory t h rough d i fe ren t ia l r ep roduc t ion i su nnecessa ry . T he q ues t ion o f why h umans ( o r a ny o the r o rgan ism f o r t ha t ma t te r ) b ecame i n vo lved i n c oevo lu t iona ry r e la t ionsh ips w i th p ar t icu la r p l an ts o r a n ima ls i se xp l ic i t ly a dd ressed h e re . U l t ima te ly , t he r e la t ive u t i l i ty o f t he p e rspec t ive d eve loped h e re c ompa red t o a l te rna t ives l i ke t ha t o f R indos w i l l d epend u pon t he i n s igh ts w h ich e ach p rov ide .

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CHAPTER 2

E STABL ISH ING COSTS AND B ENEF ITS FOR EARLY P ASTORAL FOOD P RODUCT ION : E THNOGRAPH IC AND H ISTOR IC L ABOR A ND P RODUCT IV ITY DATA AS R ELATED TO THE PREH ISTOR IC P AST

I n t roduc t ion I n o rde r t o a pp ly mode ls d er ived f r om b ehav io ra l e co logy t o i s sues o f e a r ly f ood p roduc t ion , i tw i l l b e n ecessa ry t oe s tab l ish wha tt he r e la t ive e ne rge t ics o fa l te rna t ive p reh is to r ic f ood p roduc t ion s t ra teg ies m igh t h ave b een . H ence , q uan t i ta t ive d a ta r e levan t t o t he c a lo r ic b ene f i ts a nd l a bo rc os ts a ssoc ia ted w i th p reh is to r ic p as to ra l a nd a g r icu l tu ra l s t ra teg ies mus t b e e s tab l ished . Obv ious ly , t he emp i r ica l d a ta r equ i red c an o n ly b e d e r ived f r om s uch i n d i rec t s ources a s modern e xpe r imen ta l r ep l ica t ion , o rf r om h i s to r ic a nd e thnog raph ic d ocumen ta t ion . Y e t , t housands o f y ea rs s epa ra te s uch q uan t i ta t ive d a ta f r om t he p reh is to r ic p as t . G iven t he p o ten t ia l e f fec ts o ft echno log ica l i n nova t ions o n l a bo rc os ts a nd h uman s e lec t ion o n p lan t a nd a n ima l p roduc t iv i ty , t he a pp l icab i l i ty o f h i s to r ic a nd e thnog raph ic c os t a nd b ene f i t d a ta t o a n a na lys is o f t he e ne rge t ics o f p reh is to r ic f ood p roduc t ion may b e q ues t ioned . T he p resen t c hap te rw i l l a dd ress t h is i s sue a s r e la ted t op as to ra l s ubs is tence , wh i le C hap te r 3w i l l s im i la r ly e va lua te d a ta o n c e rea l h usband ry . I n i t ia l ly , s ome b as ic t e rm ino log ica l c l a r i f ica t ions a re r equ i red . T h roughou t t h is manusc r ip t , t he t e rm " husband ry " w i l l b e u sed t o d es igna te t he c a re fu l , f r uga l , a nd t h r i f ty managemen t o f p l an t a nd a n ima l r esou rces . I ti ss pec i f ica l ly u sed t o c onvey a s ense o f e conom ic r esource c onserva t ion . F ur the r , ad i s t inc t ion w i l l b e d rawn b e tween " i n tens ive " a nd " ex tens ive " f o rms o f a n ima l h usband ry . A s a pp l ied t o man -an ima l r e la t ionsh ips , " i n tens ive h usband ry " s hou ld b e u nde rs tood a s a s t ra tegy i n wh ich h umans n o to n ly c on t ro lt he a n ima ls i n vo lved , b u t a l so p rocure o rp roduce t he f o rage r esou rces wh ich t hose a n ima ls c onsume . T he s t a l l , b y re , o rp en ‘ f eed ing o fh e rd a n ima ls w i th h a rves ted n a tura l o r c u l t iva ted f o rage , a nd t he g raz ing o f h e rds o n c u l t iva ted ( o f ten i r iga ted ) p as tu rage s pec i f ica l ly i n tended f o r h e rd c onsump t ion , wou ld c ons t i tu te e xamp les o f i n tens ive h usband ry . T he l abo r c os ts o f f eed ing a n ima ls u nde r c ond i t ions o f i n tens ive h usband ry wou ld t he re fo re r ange f r om t he t ime e xpended i np rocu r ing n a tura l ly a va i lab le f o rage t h rough t ha t o f c u l t iva t ing a p lan t c rop o r c rops f o r t he s o le p urpose o f h erd c onsump t ion . T h is l a t te rp rac t ice n ow t yp i f ies a n ima l h usband ry o n mode rn b reed ing f a rms a nd d a i ry r anches . H oweve r , t he s ta l l feed ing o fc a t t le h e rds i nE u rope u s ing t w igs a nd l e aves f r om p o l la rded e lm , b i rch , o ak , l i me , map le a nd a sh t r ees d a tes t o a t l e as t t he e a r ly f our th m i l lenn ium B .C . ( T roe ls -Sm i th 1 984 ) , wh i le b o th s t a l l feed ing a nd f enced p addocks w e re emp loyed w i th v a r ious h e rd a n ima ls i nE gyp t b y a tl e as tt he t h i rd m i l lenn ium B .C . ( Sm i th 1 969 ) . S ign i f ican t ly , i n tens ive h usband ry s t ra teg ies a re p redom inan t ly , t hough n o t e xc lus ive ly , a ssoc ia ted w i th s eden ta ry a g r icu l tu ra l p opu la t ions . H oweve r , w h i le h umans w ou ld d i rec t ly c on t ro l t he f odde r p rov ided f o r t he i r a n ima ls u nde r c ond i t ions o f i n tens ive h usband ry i n vo lv ing t he p rocuremen t o f s uch n a tura l ly o ccu r r ing f o rage a s l e a ff odde r , f o rage a va i lab i l i ty w ou ld v a ry a s ar e su l t o fb o th n a tu ra l p roduc t ive p rocesses a nd t he i n tens i ty o f h uman e xp lo i ta t ion . H ence , i n i t s s imp les t f o rm , i n tens ive h usband ry s ha res c er ta in a t t r ibu tes w i th w ha t w i l l b et e rmed " ex tens ive h usbandry . " " Ex tens ive h usband ry "w i l l b e s pec i f ica l ly u sed i nr e fe rence t op as to ra l s t ra teg ies i n wh ich h umans c on t ro l t he a n ima ls i n vo lved , b u t r egu la te t he i r f eed ing b y mov ing h e rds t o a va i lab le n a tura l f o rage . T hus , u nde r c ond i t ions o f e x tens ive h usbandry , ah e rd 's p lane o f n u t r i t ion w ou ld p r ima r i ly b e af unc t ion o fs pa t ia l ly a nd t empo ra l ly a va i lab le n a tu ra lf o rage . S ince l o ng te rm p red ic t ions o f t he s pa t ia l a nd t empo ra l a va i lab i l i ty o f s uch r esou rces a re n ecessa r i ly l i m i ted ,

s i nce mass ive s t ock l o sses a re a lways a p oss ib i l i ty , a nd s i nce h e rd a n ima ls a re t he p r ima ry " cur rency " u sed t o p rov is ion o f fsp r ing when t hey e s tab l ish n ew h ouseho lds , i n d iv idua l p as to ra l is ts wou ld d e r ive f ew b ene f i ts f r om e i the r l i m i t ing t he i r h e rd 's g row th , o r f r om c onserv ing p as ture r esources ( Bar th 1 961:102 ; B radburd 1 982 ; B re man a nd d e W i t 1 983 :1346 ; D ah l a nd H jo r t 1 976 :129 ; D yson -Hudson a nd D yson -Hudson 1 969 :79 ; I n go Id 1 980 :89 , 1 34 , 1 78 , 2 01-202 , 2 06 -207 ; W ids t rand 1 975 :149 ; ) . H ence , u nde r c ond i t ions o f e x tens ive h usbandry , i ti s e xpec ted t ha t p as to ra l is ts w ou ld r a re ly e xpend l a bo r i n t he h usbandry o f f o rage r esou rces . S im i la r c ond i t ions wou ld p resumab ly e x is t u nde r i n tens ive h usband ry i n vo lv ing t he p rocuremen t o fn a tu ra l ly o ccu r r ing f o rage . T he t r anshuman t h e rd ing p rac t iced b y e thnog raph ic p as to ra l p opu la t ions i nt he O ld Wo r ld , a nd n i ne teen th c en tury c a t t le r anch ing i nA us t ra l ia a nd t he wes te rn U .S . , wou ld c ons t i tu te e xamp les o f e x tens ive h usband ry . F requen t ly , t he managemen t o f i n d iv idua l o r c ommuna l h e rds f r om a g r icu l tura l v i l lages h as a l so i n vo lved e x tens ive h usbandry , a l though v a r ious c omb ina t ions o f e x tens ive a nd i n tens ive s t ra teg ies a re s ome t imes p rac t iced , o f ten o n a s easona l b as is ( Swee t 1 974 :96 ; Wa tson 1 979 :104 ) . B y c on t ras t , t he e thnograph ic r eco rd o f n omad ic p as to ra l p opu la t ions i nt he O ld Wo r ld wou ld s ugges t t ha t h ere , s ubs is tence h e rd managemen t h as b een p redom inan t ly , i fn o t e xc lus ive ly , b ased u pon e x tens ive h usbandr y . I tw i l l b e a ssumed i nt he p resen t a na lys is t ha t t hese v a r ious managemen t o p t ions a nd c ond i t ions wou ld a l so h ave t yp i f ied a n ima l h usband ry s t ra teg ies i nt he p reh is to r ic p as t . A s w i l l b e s hown , t h is d i s t inc t ion b e tween i n tens ive a nd e x tens ive f o rms o fa n ima l h usband ry i sc r i t ica l f o r e va lua t ing p o ten t ia l t empo ra l v a r ia t ions i n p as to ra l l a bo r a nd p roduc t iv i ty b e tween t he e thnog raph ic p resen t a nd t he p reh is to r ic p as t . W i th t hese c ons ide ra t ions i nm ind , t he l a bo rc os ts o fp as to ra l s t ra teg ies u nde rc ond i t ions o f e x tens ive h usbandr y wou ld p r ima r i ly c ons is t o f h e rd ing , e ncampmen t , b u tche r ing , a nd ( po ten t ia l ly ) m i lk ing t asks , wh i le t he b ene f i ts o b ta ined wou ld c ons is to ft he c a lo r ies d e r ived f r om mea t , f a t , o rgan , b lood , a nd ( po ten t ia l ly ) m i lk p roduc t ion . I n t he f o l low ing t ex t , p oss ib le v a r ia t ions i n t hese c os ts a nd b ene f i ts b e tween t he p reh is to r ic p as t a nd t he h i s to r ic a nd e thnog raph ic p resen t w i l l b e e nume ra ted a nd a na lyzed . A mo re c omp le te a ccoun t ing o ft he c os ts a nd b ene f i ts a ssoc ia ted w i th p as to ra l f ood p roduc t ion w i l l b e p resen ted i nC hap te r 5 . F i xed c os ts , s uch a s t he i n i t ia l c ons t ruc t ion o fp o r tab le d we l l ings o rt he manu fac tur ing o ft o o ls a nd e qu ipmen t , w i l l n o t b e a dd ressed i nt he p resen t s t udy .

P as to ra l L abo rC os ts i nt he E thnog raph ic P resen t a nd t he P reh is to r ic P as t O verv iew Among t he f ou r p as tora l l abo r c os ts u nde r e x tens ive h usbandry men t ioned a bove , t he re s eems l i t le r eason t o s uspec t t ha t ma jo r v a r ia t ions n ecessa r i ly e x is t b e tween t he p reh is to r ic p as t a nd t he e thnog raph ic p resen t wh ich wou ld a f fec t t he e ne rge t ic a na lyses a t temp ted h e re . T h is i sn o tt os ugges tt ha tv a r ia t ions i nl a bo rc os ts d i d n o t o ccu r , b u t r a the rt ha tt hose v a r ia t ions w h ich c an b e r ea l is t ica l ly p os i ted a re e i the rr e la t ive ly m ino r , o rc an b e a ccoun ted f o ro nt he b as is o fa va i lab le d a ta . T hese v a r ia t ions a re e nume ra ted a nd a na lyzed b e low b y r e fe rence t os pec i f ic p as to ra l l a bo rt a sk .

H erd ing C os ts H e rd ing t asks u nde rc ond i t ions o fe x tens ive h usband ry p r ima r i ly c ons is t o ft he t ime s pen t i nd r iv ing a n ima ls t oa nd f r om a va i lab le n a tu ra l p as tu rage , a nd t he t ime s pen t i ng ua rd ing g raz ing h e rds , wa te r ing a n ima ls , a nd s ea rch ing f o rs t rays . A s d ocumen ted l a te r , h e rd ing t asks a ccoun t f o rt he v as t ma jo r i ty o f a l p as to ra l l a bo r c os ts . I ng ene ra l , h e rd ing a nd h e rd managemen t a re t echn ica l ly a ch ieved t h rough t he e xp lo i ta t ion a nd man ipu la t ion o f v ar ious h e rd a n ima l b ehav io ra l t r a i ts . T hese i n c lude g rega r iousness , g ene ra l r esponses t o a ud i tory a nd v i sua l s i gna ls , p a t te rns o f mo the r in fan t i n te rac t ion , a nd t he n a tu ra l s easona l d iv is ion o f h e rds i n to s ubg roups a cco rd ing t o a ge a nd s ex c l asses . S ince t hese b ehav io ra l a t t r ibu tes w ou ld h ave e vo lved d ur ing a dap ta t ion t o o pen e nv i ronmen ts l o ng b e fore t he e nd o f t he P le is tocene

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( Es tes 1 974 ; Ge is t 1 974 ) , t he re s eems l i t le r eason t o s uspec t t ha t p reh is to r ic h e rd ing c os ts w ou ld n ecessa r i ly h ave b een g rea te rt han t hose a ppa ren t i nt he e thnog raph ic p resen t .

E ncampmen t C os ts E ncampmen t t asks u nde rc ond i t ions o f e x tens ive h usband ry p r ima r i ly c ons is t o ft he t ime s pen t i n mov ing r es idence a s ad i rec t r e su l t o ft he r a t iona le o fc oo rd ina t ing h e rds w i th s pa t ia l ly a nd s easona l ly v a r iab le p as tu re a nd wa te r r e sou rces . F ur the r , s i nce t he t ime s pen t i nm ig ra t ion b y n onherde rs i so f ten d evo ted t ov a r ious l a bo rt asks u n re la ted t od i rec t p as to ra l p roduc t iv i ty , s uch a sf o rag ing a nd s oc ia l iz ing , t he l a bo rc os ts o fb o th l o ng a nd s ho r t d i s tance m ig ra t ions w i l l b e r es t r ic ted i nt he p resen t a na lys is t ot he t ime s pen t i nb reak ing e ncampmen ts a nd l o ad ing g ea r , a nd t ha t s pen t i nu n load ing g ea r a nd r ees tab l ish ing e ncampmen ts a t n ew l o ca t ions . A b as ic u nde rs tand ing o f p reh is to r ic a nd e thnog raph ic p as to ra l s he l te rs a nd e ncampmen ts i s t he re fo re r equ i red . Ap rom inen t f ea tu re o ft he ma te r ia l c u l tu re o fc on tempo ra ry p as to ra l is ts i nt he N ea r E as t a nd N or th A f r ica i st he i r u se o f s qua re o r r ec tangu la r " b lack t en ts " o f w oven h a i r ( Dough ty 1 936 :265 -268 ; Mus i l 1 928 :6 1-78 ) . Among t hese p opu la t ions , h e rd a n ima ls a re o ccas iona l ly c o r ra l led i ns t ock e nc losu res , b u t a re mo re f r equen t ly k ep t n ex tt oi n d iv idua lt en ts o rw i th in t en t c ompounds . I n d i s t inc t c on t ras t , many c on tempo ra ry p as to ra l p opu la t ions i n E as t A f r ica c ons t ruc t r ound t en ts o rs he l te rs , a nd i n va r iab ly e nc lose t he i rh e rds i nb rush c o r ra ls . F o r e xamp le , t he R end i l le ( Sa to 1 980 :47 ) , Gab ra ( To r ry 1 973 :173 -178 ) , a nd D assane tch ( G i f fo rd 1 977 :68 -70 , 1 978 :87 ) u t i l ize p or tab le r ound h u ts o r " t en ts " c ons is t ing o fa n ima l s k ins a nd /o rf i be r ma ts d raped o ve r ah em isphe r ica l f r ame o fc u rved w ooden s uppor ts . F ur the r , t he i r h e rd a n ima ls a re c ons is ten t ly q uar tered i nc o r ra ls ( o r " k raa ls " ) c ons truc ted o f t ho rn b ushes , p a r t ia l ly a s ad e fens ive measu re a ga ins t r a id , b u t a l so t o i n h ib i t c a rn ivo re p reda t ion ( G i f fo rd 1 977 :7 1; S a to 1 980 :45 -46 ; S . Sm i th 1 980 :48 1) . I twou ld s eem t ha t a na logous d we l l ings a nd s t ock e nc losures we re a l so emp loyed b y t he e a r l ies t p as to ra l p opu la t ions k nown i nt he N ea r E as t . Wh i le s qua re o r r ec tangu la rt en ts p resumab ly s im i la rt o mode rn b l ack t en ts o f woven h a i r a re a rchaeo log ica l ly i n fe r red i ns ou thwes te rn I r an a s e a r ly a s 6 ,200 -5 ,900 B .C . ( Ho le 1 974 ) , e v idence f r om t he A rab ian P en insu la a nd N or th A rab ia s ugges ts a n e a r l ie r u se o f r ound s he l te rs b y p reh is to r ic p as to ra l is ts . R ecen t s ys tema t ic a rchaeo log ica l s u rveys s ponso red b yt he S a 'ud i A rab ian g ove rnmen t h ave l o ca ted p reh is to r ic p as to ra l e ncampmen ts w i th r ound h u ts o r t en ts t h roughou t a l b u t t he e as te rn , s ou th -cen t ra l , a nd s ou theas te rq r eg ions o f t he c oun t ry ( Adams e t a l . 1 977 :34 -35 ; Ga r rad a nd H a rvey 1 981:142 143 ; G i lmo re e t e l . 1 982 : 13 ,16 ; I n g raham e t a l . 1 98 1:67 -7 1 ; P a r r e t a l . 1 978 :37 -39 ; Z a r ins e t a l . 1 979 :25 -26 , 1 980 :19 -23 , 1 981 :21) . F ur the r , t he r o ck a l ignmen ts wh ich c ha rac te r ize t hese c i rcu la r h u ts , h u t c ompounds , a nd s tock c o r ra ls p hys ica l ly p reda te t he a ppea rance o f t he s qua re a nd r ec tangu la r r ock a l ignmen ts a nd r a in d i tches w h ich c ha rac te r ize a nc ien t a nd modem b l ack t en t e ncampmen ts o n t he A rab ian P en insu la a nd N or th A rab ia ( Be t ts 1 982a , 1 982b :25 ; R ees 1 929 :401 ; Z a r ins e ta l . 1 980 :23 ) . T hus , wh i le s he l te rs s im i la r t o mode rn b l ack t en ts we re a ppa ren t ly emp loyed b y p reh is to r ic p as to ra l is ts i ns ou thwes te rn I r an , e a r ly p as to ra l is ts o nt he A rab ian P en insu la a nd i n N o r th A rab ia u t i l ized r ound h u ts o rt en ts . Wh i le D ough ty ( 1936 :328 ) h ad c ome a c ross s t one c i rc les i nt he l a te n i ne teen th c en tury n ea rt he o as is o fT e ima i nn or thwes te rn S a 'ud i A rab ia wh ich a ppea red t o b e a nc ien t p as to ra l e ncampmen ts , t he b es t -known b u t p rev ious ly u nda ted e xamp les o ft hese e a r ly p as to ra l s i tes w e re t hose f ound i nn o r theas te rn J o rdan , l o ca ted i na nd a round t he A z raq B as in . T hese h ad b een i n i t ia l ly d i scove red , d esc r ibed , a nd p ho tog raphed f r om t he a i r b y RAF p i lo ts f l y ing t he B aghdad t o C a i ro a i r ma i l r ou te a f te r WW I ( Ma i t land 1 927 ; R ees 1 929 ; R i ley 1 982 ) . D i rec t a rchaeo log ica l i n ves t iga t ions h ave s i nce c on f i rmed t ha tt hey a re i n deed a nc ien t h u t c i rc les , h u t c ompounds , a nd s tock c o r ra ls , t he o u t l ines o f wh ich we re " f oss i l ized " i nt he f o rm o f r ock a l ignmen ts r esu l t ing f r om t he c l ea r ing o fc obb le -s t rewn s ur faces d ur ing t he i r c ons t ruc t ion a nd u se ( Be t ts 1 981 , 1 982a , 1 982b , 1 983 , 1 983 -84 , 1 984 ; F i e ld 1 960a ; H e lms 1 981 , 1 982 ) . T hese s i tes w e re s eem ing ly o ccup ied b y p as to ra l is ts i nN or th A rab ia a s e a r ly a st he e i gh th ( Be t ts , i nR o l le r 1 984 :220 ; I n g raham e ta l . 1 98 1:67 ) o r s even th m i l lenn ia B .C . ( Be t ts 1 982b :3 1 ; H e lms 1 981:50 , 1 982 :101) , a s b ased u pon t he c ross -da t ing o f a ssoc ia ted l i t h ics . A s s uch , 1 7

t h ese s i tes c u r ren t ly c ons t i tu te t he e a r l ies t i n d ispu tab le e v idence f o r p as to ra l s oc ie t ies i nt he N ea r E as t . S uch s i tes s eem ing ly c on t inue t oa ppea ri nt he a r chaeo log ica l r eco rd o ft he A rab ian P en insu la a nd N or th A rab ia t h rough t he C ha lco l i th ic p e r iod o ft he f our th m i l lenn ium B .C . , a t w h ich t ime t h ey w e re i n c reas ing ly s upe rseded b ys qua re a nd r e c tangu la rr ock a l ignmen ts . T he s i te p l ans o ft hese a nc ien te ncampmen ts c l ose ly p a ra l le lt hose o f mode rn E as tA f r ican p as to ra l p opu la t ions , s uch a s t he R end i l le , G ab ra , a nd D assane tch . F ur ther , t hey i n c lude b o th l a rge c i rcu la r c ompounds , o f ten c ons is t ing o fa n o u te r r i ng o f h u t c i rc les w i th i n te rna l c o r ra l d i v is ions , a nd sma l le r h e rd ing c amps o f i n d iv idua l s he l te rs o r w indb reaks w i th a ssoc ia ted c o r ra ls . T o a vo id s ubsequen t amb igu i ty i n r e fe r r ing t o t he l a rge c i rcu la r h u t a nd c or ra l e ncampmen ts o f a nc ien t A rab ia , t he Swah i l i t e rm manya t ta w i l l b e s ubs t i tu ted f o r t he i r c ha rac te r iza t ion a s " j e l ly f ish s i tes " b y B ets a nd H o lms . T he t e rm manya t ta i sd e r ived f r om a Maasa iw o rd , a nd d eno tes ac i rcu la r , s em ipe rmanen tp as to ra l h omes tead o rs e t t lemen to fs ome s i ze a nd s ubs tance ( a f te rG ifo rd 1 977 :72 , 1 978 :87 ) . S ign i f ican t ly , t he s pa t ia l p a t te rn ing o ft hese a nc ien t manya t tas a nd h e rd ing c amps i nN or th A rab ia a ppea rs t o r e f lec t p as to ra l d ua l res idence t r anshumance p a t te rns s im i la r t o t hose e t hnog raph ica l ly a t tes ted i nE as tA f r ica . T hey a l so s eem ing ly r e f lec t as easona lf o cus o no rn ea r t he ma rg ins o ft he H o locene l a kes w h ich f o rme r ly e x is ted i nt he b as ins a nd i n te rduna l r eg ions o fe as te rn J o rdan a nd t he A rab ian P en insu la . U n fo r tuna te ly , n o th ing i sa ppa ren t ly k nown o ft he h e rd a n ima ls e xp lo i ted b y t hese a nc ien t p as to ra l is ts . A t p resen t , s heep , g oa ts , c a t t le , a nd c ame ls m us t a l b e c ons ide red a s p oss ib le c and ida tes , a l though f o r r easons d i scussed i n s ubsequen t c hap te rs , c a t t le a nd g oa ts s eem mos t l i ke ly . R ega rd less , s i nce t he p as to ra l s he l te rs a nd e ncampmen ts h i s to r ica l ly a nd e t hnog raph ica l ly d ocumen ted i nb o th t he N ea r E as t a nd A f r ica e xh ib i t l i t le v a r ia t ion i na ppa ren t s t ruc tu re a nd l a you t f r om t hose a rchaeo log ica l ly a t tes ted f o rt he p reh is to r ic p as t , i ts eems r easonab le t oa ssume t ha t e t hnog raph ic a nd h i s to r ic d a ta o n t he l a bo r c os ts a ssoc ia ted w i th t he i r movemen t a nd c ons t ruc t ion s hou ld a l so b e a pp rop r ia te f o ra pp l ica t ion t ot he p reh is to r ic p as t .

B u tcher ing C os ts B u tche r ing l a bo ri n c ludes s uch t a sks a s k i l l ing , e v isce ra t ing , a nd ( s ome t imes ) s k inn ing a n a n ima l , a s w e l l a s t he r educ t ion o f t he r e su l t ing c a rcass i n to c u l ina ry c u ts o f m ea t . H e re , v a r ia t ions i nl a bo r e f f ic iency may b e e xpec ted a s ar e su l t o f s t one v s . me ta l t echno logy . U n fo r tuna te ly , o n ly q uan t i ta t ive d a ta o n b u tche r ing w i th m e ta l t oo ls i sc ur ren t ly a va i lab le . H oweve r , i tw i l l b e s hown t ha t s i nce b u tche r ing l a bo rw ou ld a ccoun t f o r l e ss t han 1% o ft o ta l a nnua l p as to ra l l a bo rc os ts u nde rc ond i t ions o fe x tens ive h usband ry , v a r ia t ions i nb u tche r ing e f f ic iency a s ar e su l to ft e chno log ica l i n nova t ion a re n o t e xpec ted t oh ave s i gn i f ican t ly a f fec ted t he o ve ra l l e ne rge t ics o fp as to ra l s t ra teg ies .

M i lk ing C os ts M i lk ing l a bo r c ons is ts o ft he t ime s pen t i ne x t rac t ing m i lk f r om t he l a c ta t ing f ema les i na h e rd . H oweve r , o n ly i ft he f ema les i np reh is to r ic h e rds w e re m o re r eca lc i t ran tt ob e m i lked t han t he i r modern c oun te rpar ts w ou ld t he l a bor e xpended h ave b een g rea te r t han i n t he e thnog raph ic p resen t . D ur ing t he r ecen t d omes t ica t ion o ft he c ommon e land ( Tauro t ragus o ryx ) , ah eav i ly b u i l t b ov ine a n te lope o fA f r ica s t and ing a pp rox ima te ly 1 . 8 ma tt he s hou lde r , c ond i t ion ing o f y oung f o r f u tu re m i lk ing h as b een a ch ieved b y r ubb ing a nd k nead ing t he u dde rs o ft e the red i n d iv idua ls ( T reus a nd K ravchenko 1 968 :400 ) . A du l t e l and a re o f ten m i lked i ns pec ia l s t a l ls , b u tt h is i sn o t r e qu i red i na lc ases , a nd t e the r ing w ou ld p resumab ly a ch ieve t he s ame e nd . R uss ian e xpe r imen ts w i th d omes t ica t ing e l k a l so i n d ica te t ha tt hese a n ima ls may b e r ead i ly c ond i t ioned t om i lk ing ( W i lk inson 1 972 :108 ) . Among r e indee r , t he mo re t ame a n ima ls k ep t i nd omes t ic h e rds n ea re ncampmen ts a re s a id t or ead i ly s ubm i tt om i lk ing ,w h i le l e ss t ame i n d iv idua ls i np as to ra l h e rds k ep t a way f r om e ncampmen ts mus t b e t e the red t o b e m i lked ( l ngo ld 1 980 :101-103 ) . U n fo r tuna te ly , n o d ocumen ta t ion h as b een f ound o n w he the r v a r ia t ions i nt he r e la t ive r e ca lc i t rance o fd i f fe ren t h e rd s pec ies e x is t i nr e spec t t om i lk ing . H oweve r , t op a r t ia l ly a ccoun t f o r t h is p oss ib i l i ty , t he a dd i t iona l l a bo r o f s omeone c on t ro l l ing i n d iv idua l f ema les d u r ing t he m i lk ing p rocess w i l l b e i n c luded i ns ubsequen t l a bo r c a lcu la t ions . A s d ocumen ted b e low , t he 1 8

f u r the rp oss ib i l i ty t ha tt he p hys io logy o fl a c ta t ion i np reh is to r ic f ema les was s u f f ic ien t ly d i f fe ren t f r om t ha t o ft he i r mode rn c oun te rpa r ts a s t oa f fec t t he l a bo rc os ts o fd a i ry ing s eems u n l ike ly . O bv ious ly , w hen m i lk p roduc t ion i sn o t p a r t o f ap as to ra l s t ra tegy , n e i the r t he l a bo r c os ts o f d a i ry ing n o rt he c a lo r ic b ene f i ts o fm i lk w ou ld a pp ly .

P as to ra l P roduc t iv i ty i nt he E thnog raph ic P resen t a nd t h e P reh is to r ic P as t G enera lC ons ide ra t ions Wh i le s eve ra l e co log ica l f ac to rs a nnua l ly a f fec t t he b ene f i ts d e r ived f r om p as tora l p roduc t iv i ty , d e f ined a st he c a lo r ies o b ta ined f r om mea t , f a t , o rgan , b l ood , a nd ( po ten t ia l ly ) m i lk p roduc t ion , i ts eems r easonab le t o e xpec t t ha t s uch e f fec ts , w h i le s pa t ia l ly a nd t empo ra l ly v a r iab le , wou ld b e c ommon t o b o th t he e thnog raph ic p resen t a nd t he p reh is to r ic p as t . Mo re c r i t ica l i nt he p resen ta na lys is i st he p oss ib i l i ty o ft empo ra lv a r ia t ions i np as to ra l p roduc t iv i ty a nd a ssoc ia ted l a bo rc os ts a s ar e su l to fc umu la t ive v a r ia t ions i nt h ep heno typ ic t r a i ts o fh e rd a n ima ls u nde r c ond i t ions o f h uman h usband ry a nd s e lec t ive man ipu la t ion d ur ing t he t housands o f y ea rs w h ich s epa ra te t he p reh is to r ic p as tf r om t he e t hnog raph ic p resen t . G iven t he e vo lved o ppor tun is t ic a nd man ipu la t ive p roc l iv i t ies o fh umans , i ts eems h i gh ly p robab le t ha t e a r ly p as to ra l is ts w ou ld h ave s e lec ted f o r d es i rab le t r a i ts i nt he i r h e rd a n ima ls t h rough t he s imp le p rocess o fp heno typ ic s e lec t ion a nd p a i r ing . I n deed , i ts eems r easonab le t os uspec tt ha t t he i n i t ia l c ho ice o fw h ich a n ima ls among w i ld p opu la t ions w e re mos t d es i rab le f o ru se a s f ound ing membe rs f o r e a r ly p as to ra l h e rds w ou ld h ave b een ad ec is ion made u pon t he c ompa ra t ive o bse rva t ion o f s uch d es i rab le p heno typ ic t r a i ts ( W i lk inson 1 971) . H oweve r , t he re w ou ld a l so s eem t ob e n a tura l l i m i ts i mposed b y t he a va i lab le p l ane o f n u t r i t ion o n t he c umu la t ive p roduc t ive r e su l ts wh ich c ou ld h ave b een a ch ieved t h rough t he s ubsequen t p heno typ ic p a i r ing o fh e rd a n ima ls u nde rc ond i t ions o fe x tens ive h usband ry . F ur the r , i tw i l l b e s ugges ted t ha tt he p o ten t ia l r ange o fp heno typ ic v a r iab i l i ty a nd p l as t ic i ty a l ready e s tab l ished i n s pec i f ic h e rd s pec ies b y t he e nd o f t he P le is tocene h as b een s e r ious ly u nde res t ima ted i n many p rev ious a na lyses . T hese i s sues w i l l b e e l abo ra ted b e low b y r e fe rence t os pec i f ic b ehav io ra l , mo rpho log ica l , a nd p hys io log ica l t r a i ts c ommon ly p erce ived a s b e ing s i gn i f ican t i ne i the r t he d omes t ica t ion p rocess , o r i n d i s t ingu ish ing " w i ld " f r om " domes t ica ted " h e rd s pec ies . F o l low ing t hese a na lyses ,t he e x is tence among v a r ious h un te r -ga the re rp opu la t ions o fb o th t he p hys ica l means a nd l a bo rc os ts p resumab ly a ssoc ia ted w i th t he i n i t ia lc ap tu re a nd c on t ro l o fw i ld h e rds p r io rt o t he e vo lu t ion o fp as to ra l s t ra teg ies w i l l b e d ocumen ted .

B ehav ior Ac oncep to fh uman c ausa t ion o fn ew b ehav io ra lt r a i ts i nt he i rh e rd a n ima ls i si mp l ic i t i nt he mode rn v erb " domes t ica te , " w h ich w as d e r ived f r om t he p as t p a r t ic ip le d omes t ica tus o ft he Med ieva l L a t in v e rb d omes t icare . Wh i le t h is v e rb o r ig ina l ly c onveyed as ense o f" t oc ause t ob e a t h ome , " i twas e x tended t o man -an ima l r e la t ionsh ips w i th a s ense o f " t o t ame . " T hus , t o " domes t ica te "a na n ima ll i t e ra l ly m eans t oc l a im t h a ta n ima lf r om i t sw i ld s t a te b y" caus ing "i tt ob e mo re e as i ly managed o rc on t ro l led t han i t sw i ld c oun te rpar ts . H ence , i nt he t r ad i t iona l s ense o f t he t e rm , i ti si mp l ic i tt ha tt hese b ehav io ra lt r a i ts m us t n ecessa r i ly h ave b een i n duced b y h uman a gency . T he a l te rna t ive p oss ib i l i ty t ha t , t ov a ry ing d eg rees , h e rd s pec ies m igh t a l ready h ave p ossessed b ehav io ra l t r a i ts w h ich h umans c ou ld h ave e xp lo i ted a nd s ubsequen t ly f a vo red i n t he i r s ubs is tence h e rds , i sn o t a n i n he ren t c oncep t i nt he e t ymo logy o f" domes t ica t ion . " O n a b roader s ca le , i ti ss ugges ted t ha t t h is g ene ra l b i as t owa rds t he h uman c rea t ion o f n ew p heno typ ic t r a i ts h as s t rong ly i n f luenced many p e rcep t ions o f t he e vo lu t ion o f e a r ly f ood p roduc t ion s t ra teg ies . A c tua l ly , w i th v e ry f ew e xcep t ions ( e .g . , t a i l c a r r iage i nd omes t ic d ogs a nd p a t te rns o f a t tack i n mode rn s t ra ins o ff i gh t ing c ocks ) , t h e re s eems t ob e l i t le e v idence t os ugges tt ha t t he b as ic b ehav ior p a t terns o f d omes t ic u ngu la tes , c a rn ivo res , a nd g a l l inaceous b i rds h ave s i gn i f ican t ly d i ve rged f r om t hose o ft he i rw i ld p rogen i to rs ( C lu t ton -B rock 1 981 :55 ; G e is t 1 97 1a , 1 975 ; H a fez a nd S co t t 1 962 :297 ; H a le 1 962a :4 1; S cha l le r 1 977 ) . N o t s urp r is ing ly , t he i n i t ia l 1 9

o bser va t ion o fs pec i f ic b ehav io rp a t te rns i nd omes t ic a n ima ls h as o f ten s t imu la ted r e sea rch i n to s uch p a t te rns i nt he i rw i ld c oun te rpa r ts ( Ha le 1 962b :593 ) . I ng ene ra l , b as ic mo to r p a t te rns a nd s enso ry c apac i t ies h ave s eem ing ly r ema ined q u i te s t ab le u nde rd omes t ica t ion ( Ha le 1 962a :40 4 6 ; S co t t 1 962 :4 ) . Wha t h as a ppa ren t ly c hanged i st he f r equency o r i n tens i ty o f e x tan t b ehav io ra l e xp ress ions a s ar esu l t o fs h i f ts i nr esponse t h resho lds i n duced b y h uman s e lec t ion ( Ha le 1 962a :46 ; R a tne ra nd B o ice 1 975 :9 ) . T ha t n umerous c hanges d o o ccur i nt he b ehav io r o f a n ima ls u nde r t he a l te red e co logy p roduced b y c ond i t ions o f h uman h usbandry i su nden iab le . H owever , t hese s eem t o p r ima r i ly r esu l t f r om s uch f ac to rs a s i mp r in t ing ( o r i mp r in t ing l ike p rocesses ) , e f fec t ive ly i n duc ing t he a n ima l t o b ehave t owa rds h umans a s a membe r o f i t s o wn s pec ies , o r f r om c ond i t ioned r esponses a nd l ea rned a d jus tmen ts a s ar esu l t o f t he f r equency a nd d ura t ion o f t he i r a ssoc ia t ion w i th h umans ( Bask in 1 974 ; C owan 1 974 :929 -930 ; D enenbe rg 1 962 ; G e is t 1 97 1b ; G uh l 1 962 :105 106 ; H a fez 1 968a :213 -2 14 ; H a le 1 962a :36 -37 ; H owa rd 1 965 :463 ; K rebs a nd D av ies 1 985 :258 -260 ; P r ice a nd K ing 1 968 :41-44 ; R a tne r a nd B o ice 1 975 :1112 ; R eed 1 969 :369 ; S co t t 1 962 :4 -6 ; W i lk inson 1 972 :114 ; Woo lpy a nd G insberg 1 967 ) . T h is h as c er ta in ly b een t he c ase d u r ing t he r ecen t d omes t ica t ion o ft he c ommon e l and . I nt he a bsence o f p r io r s e lec t ive b reed ing , t he o f fsp r ing o f c ap t ive e l and h e rds e xh ib i t s uch " domes t ica ted " t r a i ts a s d oc i l i ty , e ase o fh and l ing , a nd r es t ra in t b y f ences , a nd a re s a id t ob e s upe r io r i ns uch t r a i ts r e la t ive t o r ange c a t t le ( Jewe l l 1 969 :104 ; P osse I t 1 963 ; S k inne r 1 967 ) . S im i la r r e su l ts h ave a l so b een n o ted w i th t he r ecen t d omes t ica t ion o f musk -oxen a nd e l k ( W i lk inson 1 97 1 , 1 972 :107 -8 ) . E ven b y s uch s imp le p rocesses a s p rov id ing s t imu l i b eyond t he i ru sua l p e r iod o r i n tens i ty , c er ta in p roduc t ive ly s i gn i f ican t b ehav io rs c an b e p ro longed . F o r e xamp le , e gg l a y ing may b e p ro longed i nn ume rous s pec ies o fb i rds b y t he s imp le e xped ien t o f r emov ing e ggs a s t hey a re l a id ( Ha le 1 962a :36 ) . As im i la r r e su l t may b e o b ta ined i nt he q uan t i ty a nd l e ng th o fl a c ta t ion i n c er ta in s pec ies t h rough h uman m i lk e x t rac t ion ( Cow ie e t a l . 1 980 ; Mepham 1 983 ; S h inde 1 978 ) , ap rocess wh ich , a l ong w i th t he c ond i t ion ing o ft he m i lk e j ec t ion r e f lex , w i l l b e f u r the r d i scussed i nt he s ubsequen t s ec t ion o n p hys io logy . O ve ra l l , many o ft he b ehav io ra l c hanges n o rma l ly a ssumed t o b e a ssoc ia ted w i th e a r ly d omes t ica t ion may b e a t t r ibu ted t o e x tan t p l as t ic i ty i nb ehav io ra l t r a i ts among s pec i f ic a n ima l s pec ies l i v ing w i th in a h uman ly a l te red e co logy . A s n o ted b y B e r ry ( 1969 :2 13 -214 ) a nd H owa rd ( 1965 :463 ) , p heno typ ic f l ex ib i l i ty i sa n e vo lved t r a i t o fa l h i ghe ro rgan isms , a nd t he re i sn o r eason t ob e l ieve t ha t e a r ly d omes t ica t ion n ecessa r i ly i n vo lved c hanges i nt he g eno types o fa n ima ls i no rde rt op roduce n ew p heno typ ic e xp ress ions . T he s ame may b e s a id f o rt he a b i l i ty o f h umans t oc on t ro l h e rd mpvemen ts b y e xp lo i t ing t he i r g ene ra l r esponses t o a cous t ic a nd v i sua l s i gna ls . N o t ing t ha t t he a ud i tory a nd v i sua l s i gna ls u sed t o man ipu la te h e rd movemen ts i nd i f feren t r e indee r h e rd ing s oc ie t ies w e re a l s urp r is ing ly s im i la r , L .S . B ask in ( 1974 , 1 976 ) o ft he R uss ian I n s t i tu te o fE vo lu t ion , Mo rpho logy , a nd A n ima l E co logy f ound t ha t t hese s ame s i gna ls we re a l so e f fec t ive i n c on t ro l l ing t he movemen ts o fb o th f e ra l a nd w i ld c a r ibou h e rds . F ur the r , t he s ame a ud i to ry s i gna ls w e re f o und t o b e e f fec t ive i nc on t ro l l ing t he h e rd movemen ts o f d omes t ica ted s heep a nd y aks . U s ing t hese s ame h e rd ing s i gna ls , B ask in ( 1974 :537 ) a nd h i s c o l leagues w e re s ubsequen t ly a b le t o c on t ro l aw i ld h e rd o fE u ropean b i son , k nown a s " w isen t " ( B i son b onasus ;p oss ib ly c onspec i f ic w i th B ison b i son , s ee C lu t ton -B rock 1 98 1 :66 ) o ve r ap e r iod o fs eve ra l d ays , a nd e ven tua l ly t o d r ive t hem i n to ac o r ra l . S im i la r ly , t hey w e re a b le t od i rec tt he movemen ts o f ah e rd o fm ig ra t ing moose , e ven tua l ly c ap tu r ing t hem i nc o r ra ls a s we l l . N o t s u rp r is ing ly , B ask in c onc luded t ha t p as to ra l h e rd ing t echn iques o r ig ina ted f r om t he k now ledge a nd e xper ience o f a n ima l b ehav io rs a cqu i red b y p reh is to r ic h un t ing p opu la t ions e xp lo i t ing h erds o fg rega r ious a n ima ls . S im i la r ly , t he c on t ro l o ff ema les a nd y oung i nh e rds may b e a ch ieved b y e xp lo i t ing g ene ra l p a t te rns o f mo the r in fan t b ehav io r ., F o r e xamp le , c a r ibou c ows a nd t he i rc a lves d i l igen t ly s eek t o r eun i te a f te r b e ing s epa ra ted i n to d i s tan t g roups ( Len t 1 974 :28 ) , a nd a cco rd ing t o B ask in ( 1974 :539 ) , i n many h e rd s pec ies , d am a nd o f fsp r ing w i l l r e turn t o t he p lace whe re t he l a s t s uck l ing e ven t o ccur red i fb o th a re s ubsequen t ly s epa ra ted . N o t s urp r is ing ly , L uc re t ius ( De R erum Na tu ra I ,3 97 -399 ; 1 968 :70 -71) h ad a l ready n o ted i nt he f i r s t c en tury B .C . t ha t c ows s epa ra ted f r om t he i rc a lves r epea ted ly r e tu rned t ot he i r b y res i ns ea rch o ft he i ro f fsp r ing , w h i le l ambs a nd k i ds wou ld r e turn t o t he " m i lk -swo l len u dde rs " o f t he i r own d ams " a t n a tu re 's b idd ing . " B ask in o bse rved t he e xp lo i ta t ion o f s uch mo the r in fan t t endenc ies b y c ame l p as tora l is ts i n T urkmen is tan . H e rde rs r egu la r ly s epa ra ted f ema les f r om t he i r u nweaned 2 0

y ea r l ings a f te r t he mo rn ing s uck l ing . H e rded i ns epa ra te g roups d u r ing t he d ay , b o th r ead i ly r e turned " home " i nt he e ven ing when t he n ex t s uck l ing p e r iod a pp roached . S eve ra l J apanese e thnog raphe rs h ave a l so n o ted t ha td a i ly c a t t le a nd c ame l h e rd ing a re t echno log ica l ly a ch ieved b yt ak ing i n fan ts a s " hos tages " ( Sa to 1 980 :63 ) . Ih ave p e rsona l ly o bse rved t h is p rac t ice among g oa t h e rd ing p as to ra l is ts i ns ou the rn J o rdan ( pe rsona l o bserva t ion 1 986 ) . A ppa ren t ly , s uch p a t te rns o f ma te rna l in fan t i n te rac t ion a re w ide ly e xp lo i ted i n t he l i ves tock i n dus t ry ( Len t 1 974 :28 ) . I n ma t te rs o f g ene ra l h e rd managemen t , s uch a s r egu la t ing t he b reed ing o f f ema les , m in im iz ing c on f l ic ts b e tween ma les d u r ing ma t ing s easons , a nd a d jus t ing g raz ing t o max im ize p roduc t iv i ty a nd g row th a cco rd ing t o t he c hang ing a t t r ibu tes o f t he a ge a nd s ex c l asses c omp r is ing a h e rd , p as to ra l is ts h e rd ing s heep , g oa ts , c a t t le , a nd c ame ls s easona l ly s ubd iv ide t he ir h erds i n to sma l le r g roups . T hese s ubd iv is ions a re q u i te s im i la r i n t he i r a ge -sex c ompos i t ion t o t he n a tu ra l s easona l s ubd iv is ions o bserved among a l h erds o f g rega r ious b ov ids ( Es tes 1 967 , 1 974 ; H enry 1 975 :380 -381 ; H esse 1 984 :25 1; J a rman a nd J a rman 1 979 ; Mende lssohn 1 974 :737 ; S immons 1 975 -77 ; Wa l ther e t a l . 1 983 :35 -42 ) , g i ra f fes ( Fos te r a nd D agg 1 972 ) , a nd f r ee l iv ing c ame ls ( Bask in 1 974 :538 ; C ompagnon i a nd T os i 1 978 :100 ; G au th ie r -P i l te rs 1 974 :546 ; Gau th ie r -P i l te rs a nd D agg 1 981:78 ) . A ga in , t he e xp lo i ta t ion a nd man ipu la t ion o fe x is t ing p a t te rns o fh e rd b ehav io r i sa ppa ren t . I nl i gh t o ft h is d a ta , t he a ssump t ion t ha t s uch a n ima ls a sp reh is to r ic c a t t le w ou ld n ecessa r i ly h ave b een f o rm idab le b eas ts t o d omes t ica te d ue t o t he i r s i ze a nd f i e rce b ehav io r ( e .g . , B ököny i 1 976 :20 ; C lu t ton -B rock 1 981:66 ; F agan 1 986 :220 ; H o le 1 984 :53 , 5 7 ; I ngo Id 1 980 :139 ) i n v i tes a r eeva lua t ion . T he w i ld p rogen i to r o f d omes t ic c a t t le i sb e l ieved t o h ave b een t he a u rochs ( Bos p r im igen ius ) , t he l a s t k nown s pec imen o f wh ich d i ed i nt he B ia low iaza g ame p rese rve i nP o land i nA .D . 1 627 . Ou rk now ledge o fi t s " w i ld " b ehav io r i sp r ima r i ly d e r ived f r om " r econs t i tu ted " a urochsen . T hese we re i n i t ia l ly p roduced i n t he 1 920s a nd 1 930s i n Mun ich b y c ross ing a nd s e lec t ive ly b ack -b reed ing H unga r ian a nd P odo l ian s teppe c a t t le , S co t t ish H igh land c a t t le , A lp ine b reeds , F r ies ians , a nd Co rs icans f o r t he i r p r im i t ive mo rpho log ica l t r a i ts ( Heck 1 951; Z eune r 1 963 :204 ) . S im i la r e xper imen ts we re c onduc ted i n B e r l in u s ing S pan ish f i gh t ing c a t t le , t he C ama rgue b reed o ft he L owe r R hone , a nd C o rs ican c a t t le ( Heck 1 952 ; Z eune r 1 963 :205 ) . T he r esu l ts h ave b een i n terp re ted a s s ugges t ing t ha t t he a nc ien t a u rochs was f i e rce , t empe ramen ta l , a g i le , a nd e x t reme ly s hy . H owever , i nt he a bsence o f a ge a nd s ex s pec i f ic b ehav io ra l d a ta , i ti sn o t p oss ib le t o a ssess t he c ha rac te r iza t ion o fa l a u rochsen b y s uch t r a i ts . C e r ta in ly , ma tu re ma les i n many h e rd s pec ies e xh ib i t s uch a gg ress ive t r a i ts , p a r t icu la r ly d u r ing ma t ing s easons . H oweve r , f ema les a nd t he i r y oung a re t he p r inc ip le p roduc t ive c omponen t o f p as to ra l s ubs is tence h e rds , a nd v e ry f ew s t ud ma les a re r equ i red . P resumab ly , b ehav io ra l mod i f ica t ions i ny oung ma les wou ld h ave r esu l ted f r om t he i r r ea r ing i na ssoc ia t ion w i th h umans , f ac i l i ta t ing t he i rf u ture c on t ro l a nd e xp lo i ta t ion a s s t uds . F ur the r , i nt he a bsence o f a ny d i rec t k now ledge o ft he p hys io logy a nd b ehav io r o f B os p r im igen ius , i ti sn o t p oss ib le t o d e te rm ine w he ther i n tens ive ly b ack -b red a u rochsen n ecessa r i ly e xh ib i tt he r ange o fb ehav io ra l f l ex ib i l i ty w h ich p resumab ly c ha rac te r ized t he i r p reh is to r ic p rogen i to rs ( I saac 1 962 :200 ) . F ina l ly , t he a va i lab le d a ta a re i n su f f ic ien t t o e va lua te whe the r d es i rab le b ehav io ra l t r a i ts l i ke t hose d ocumen ted e a r l ie r a l so e x is ted wh ich w ou ld h ave f ac i l i ta ted t he h uman man ipu la t ion o fB os p r im igen ius p opu la t ions . E ven i fa f i e rce a nd t empe ramen ta l d i spos i t ion d id c ha rac te r ize s ome B os p r im igen ius p opu la t ions i nE u rope , i tmay b e q ues t ioned whe the rt h is was n ecessa r i ly c ha rac te r is t ic o f a l m embe rs o f as pec ies w i th d i s t inc t p opu la t ions o ccupy ing h ab i ta ts s t re tch ing f r om A s ia t h rough E u rope a nd f r om t he E u ras ian t und ra t ot he I n d ian O cean a nd i n to N o r th A f r ica . S uch t r a i ts , i n a dd i t ion t o r oya l a nd p r iva te c onse rva t ion e f for ts , m igh t p a r t ia l ly a ccoun tf o rw hy t he re we re s t i l l r emnan t p opu la t ions o f B os p r im igen ius i nE u rope t og o e x t inc t a ts uch a l a te d a te . I ne f fec t , t hey wou ld h ave b een l e ss d es i rab le f o r c ap tu re a nd e xp lo i ta t ion a s s ubs is tence a n ima ls r e la t ive t oa l te rna t ive c a t t le p opu la t ions w i th mo re f avo rab le b ehav io ra lt r a i ts . I fB os p r im igen ius p opu la t ions b ea r ing l e ss d es i rab le b ehav io ra l t r a i ts we re t he l a s t c a t t le t o b e i n corpo ra ted i n to h uman s ubs is tence s t ra teg ies , t he i r d escendan ts wou ld p resumab ly e xh ib i t t he mos t p r im i t ive mo rpho log ica l t r a i ts r e la t ive t oo the r c a t t le , a nd h ence h ave b een v i sua l ly t he mos t p rom is ing s ub jec ts f o r mode rn a t temp ts a tb ack -b reed ing . I ne fec t , we may h ave r econs t i tu ted s ome o f t h e l e as td es i rab le b ehav io ra lt r a i ts i np reh is to r ic c a t t le .

2 1

R e tu rn ing t o t he i s sue o f w he the r c hanges i nh e rd a n ima l b ehav iors wou ld n ecessa r i ly a f fec t t he a pp l icab i l i ty o f e thnog raph ic d a ta o n p as to ra l p roduc t iv i ty t o a n a na lys is o f t he p reh is to r ic p as t , t he a nswe r w ou ld s eem t o b e n ega t ive . P resumab ly , p reh is to r ic h un te rs a dop t ing p as to ra l s t ra teg ies wou ld h ave b een a b le t o e xp lo i t a nd man ipu la te b ehav io ra l t endenc ies a l ready p resen t i nc e r ta in h e rd s pec ies . F ur the r , t hose s pec ies e xh ib i t ing s uch t r a i ts wou ld p resumab ly h ave b een p re fe ren t ia l ly c hosen f o r h usband ry o ve r a l te rna t ive s pec ies i nw h ich s uch t r a i ts we re e i the r a bsen t o r l e ss p ronounced . F i na l ly , i ts eems r easonab le t o s uspec tt ha tt hose a n ima ls i n wh ich d es i rab le b ehav io ra lt r a i ts we re mos t s t rong ly ma rked w ou ld h ave b een s e lec t ive ly f avo red a s f ound ing membe rs f o r e a r ly p as to ra l s ubs is tence h e rds , a nd t ha t t hese t r a i ts wou ld h ave b een ma in ta ined a nd r e in fo rced i n s ubsequen t g ene ra t ions t h rough s e lec t ive p heno typ ic p a i r ing a nd p re fe ren t ia l s l augh te rp rac t ices .

Morpho logy C hanges i n a n ima l mo rpho logy u nde r c ond i t ions o f h uman h usband ry h ave b een t he p r ima ry f ocus o f e x tens ive a rchaeo log ica l a nd z oo log ica l a t ten t ion . E a r ly i nt he e vo lu t ion o f a n ima l h usband ry , c er ta in morpho log ica l c hanges s eem ing ly a ppea red i n h e rd a n ima ls . P rom inen t among t hese a re c hanges i nh o rn s i ze a nd s hape among c a t t le , s heep , a nd g oa ts , g ene ra l f ema le h o rn lessness among s heep , c hanges i ns ku l l s i ze a nd s hape among c a t t le , a nd c hanges i nc oa t c o lo ra t ion i ns heep a nd c a t t le ( Bököny i 1 973 ; C lu t ton -B rock 1 981; G r igson 1 969 ; J a rman 1 969 ; P e rk ins 1 969 ; R yde r 1 969 , 1 973 ; Z eune r 1 963 ) . C hanges i nt he f l eece o f c er ta in b reeds o f d omes t ic s heep h ave b een d ocumen ted b y R yde r ( 1969 , 1 973 ) , a l though p rev ious s ugges t ions f o r aN eo l i th ic d a t ing o f s uch c hanges o n t he b as is o f s upposed w oo l a t Q a ta l H üyük a re u n tenab le ( Ryde r 1 965 ) . T he e a r l ies t woo l s amp le e xam ined b y R yde r was f r om a n E gyp t ian t omb o f t he O ld K ingdom ( l a te t h i rd m i l lenn ium B .C . ) , a nd was f ound t ob e ah a i ry med ium woo l . AS cy th ian s heepsk in f r om C en t ra l A s ia d a t ing t o a pp rox ima te ly 4 00 B .C . e xh ib i ted r e la t ive ly f i ne f i b res t r end ing t owa rds t h ose o f mode rn c oa rse -woo led b reeds . Wh i le af ew f i ne -woo ls w e re n o ted b y R yde r among t he D ead S ea S c ro l l p a rchmen ts , y ams f r om R oman p e r iod t ombs i na nc ien t P a les t ine a nd R oman p e r iod E uropean t ex t i les g ene ra l ly e xh ib i ted f l eece i n te rmed ia te b e tween p r im i t ive h a i ry t ypes a nd modern f i ne -woo ls . S im i la r ly , s econd c en tury A .D . t ex t i les f r om t he Cave o ft he L e t te rs i nI s rae l we re a l f ound t o b e o f i n te rmed ia te woo l . T he d eve lopmen t o f woo l t ypes p r io r t o t he f i rs t m i l lenn ium B .C . t hus r ema ins s pecu la t ive . F ur the r , wh i le S ume r ian a nd A ssy r ian t e x ts d o i n d ica te t ha t " woo l " p roduc t ion was b ig b us iness b y t he t h i rd m i l lenn ium B .C . ( Sher ra t 1 981:283 ; S ebas t ian P ayne , p e rsona l c ommun ica t ion 1 987 ) , i ti s n o t e xpec ted t ha t mo rpho log ica l c hanges s uch a st h is wou ld a f fec t t he a pp l icab i l i ty o f modem c a lo r ic b ene f i t d a ta t o a n a na lys is o f p reh is to r ic p as to ra l e ne rge t ics , n o r i s woo l p roduc t ion e xpec ted t o h ave s i gn i f ican t ly i n f luenced p as to ra l s ubs is tence d ec is ion mak ing p r io r t o t he d eve lopmen t o f ma rke t e conom ies . A c tua l ly , t he s i gn i f icance o f ten a t tached t o a ppa ren t m orpho log ica l c hanges p resumed t o b e a ssoc ia ted w i th e a r ly d omes t ica t ion h as f r equen t ly b een q ues t ioned o n t he b as is o f o bse rved v a r iab i l i ty i ns uch t r a i ts among w i ld p opu la t ions ( e .g . , B e r ry 1 969 ; C hap l in 1 969 ; I s aac 1 962 ; J a rman a nd W i lk inson 1 972 ) . F o r e xamp le , t w is ted h o rns h ave b een n o ted among C aucas ian w i ld g oa ts i nt he a ppa ren t a bsence o f i n t rog ress ion w i th d omes t ic g oa ts , a l though t w is ted h orns h ave o f ten b een c ons ide red a mo rpho log ica l t r a i t a ssoc ia ted w i th d omes t ica ted g oa ts ( Bököny i 1 973 :70 ; H esse 1 984 :247 ) . S im i la r ly , " domes t ica ted " r e indeer e xh ib i t mo rpho log ica l t r a i ts wh ich f a l l w i th in t he n a tu ra l r ange o fv a r iab i l i ty a ppa ren t i n" w i ld " r e indee r ( Ban f ie ld 1 961:102 -103 ) . I n s pec i f ica l ly a dd ress ing t he i s sue o f whe ther t he re a re mo rpho log ica l c hanges wh ich i n ev i tab ly a ccompany d omes t ica t ion t ha t c ou ld n o t a r ise b y a ny o ther mechan ism , B e r ry ( 1969 :2 14 ) a nd J a rman a nd W i lk inson ( 1972 :96 ) h ave c onc luded t ha t mos t o ft he mo rpho log ica l c r i te r ia b y wh ich d omes t ica t ion i sc l a imed c ou ld e qua l ly o ccur a s a r esu l t o fp rocesses u n re la ted t od omes t ica t ion . A n e xam ina t ion o ft he a dm i t ted ly l i m i ted e v idence o n t he C ame l idae f u r the r s ugges ts t ha t n e i the r mo rpho log ica l t r a i ts a l one , n o r p heno typ ic t r a i ts c o l lec t ive ly , a re n ecessa r i ly s u f f ic ien t f o r mak ing s uch d i s t inc t ions . N e i the r B ac t r ian n o r d romeda ry c ame ls a ppea r t o h ave b een mod i f ied b y d omes t ica t ion ( Zeune r 1 963 :339 -340 ) , w h ich h as a ppa ren t ly w rough t n o i d en t i f iab le c hanges i nt he i r s ke le ta l mo rpho logy ( C lu t ton -B rock 1 981 :126 ; C ompagnon i a nd T os i 1 978 :100 ; Z a r ins 1 978 :45 ; Z eune r 1 963 :359 ) . T he s ame may b e s a id f o r l amas a nd 2 2

a l pacas i nS ou th Ame r ica ( P i res -Fer re i ra e ta l . 1 976 :489 ) . F ur the r , b o th u ngua rded a nd f e ra l C ame l idae e xh ib i t b ehav io r s c ommon t oc er ta in w i ld h e rd s pec ies wh ich a re w ide ly c ons ide red a s h av ing b een n o rma t ive f o r " w i ld " C ame l idae p opu la t ions ( Bask in 1 974 :538 ; C ompagnon i a nd T os i 1 978 : 100 ; C lu t ton -B rock 1 981:124 125 ; Gau th ie r -P i l te rs 1 974 :546 ; Gau th ie r -P i l ters a nd D agg 1 981 :59 -60 ; P i res -Fer re i ra e ta l . 1 976 :488 -89 ) . E ven t he mode rn r ep roduc t ive t r a i ts o ft he C ame l idae a ppea r t o h ave b een t he p roduc t o ft he p hy logeny o ft he i r w i ld p rogen i to rs a nd t he x e r ic e nv i ronmen ts i n wh ich t hey e vo lved ( Novoa 1 970 ; S ha lash 1 965 : 106 ) . O ve ra l l , i t w ou ld s eem t ha t t he g enera l p heno typ ic a t t r ibu tes o f mode rn C ame l idae we re a l ready e s tab l ished p r io rt ot he e vo lu t ion o fp as to ra l s t ra teg ies . O n t he b as is o fs uch d a ta , i tw i l l b e a ssumed t ha t t he r ange o f mea t , o rgan , f a t a nd b l ood y i e lds f r om u n imp roved s pec ies o fc ame ls i nt he e thnog raph ic p resen t a dequa te ly r e f lec ts t he p o ten t ia l r ange o fs uch y i e lds i nt he p reh is to r ic p as t . T he s i tua t ion w i th c a t t le , s heep a nd g oa ts , h oweve r , s eems l e ss c l ea r , p a r t icu la r ly i nr e la t ion t op o ten t ia l s i ze r educ t ions u nde r c ond i t ions o fh uman h usband ry . E xc lud ing i n s tances o f ma t ing w i th w i ld c oun terpa r ts a nd t he e xchange o f a n ima ls b e tween h e rds , s ubs is tence h e rds w ou ld t end t o b e i n b reed ing p opu la t ions u nde r h usband ry , as omewha t a na logous s i tua t ion t ot he mamma l c o lon iza t ion o fi s lands . I ns t udy ing s uch i s land c o lon iza t ions , F os te r ( 1964 ) f ound t ha t i ng ene ra l , p opu la t ions o f l a rger a n ima ls ( ca rn ivo res , a r t iodac ty ls , a nd l a gomo rphs ) t ended t o d ec rease i ns i ze w hen r ep roduc t ive ly i s o la ted , wh i le p opu la t ions o f sma l le ra n ima ls ( r oden ts ) t ended t oi n c rease i ns i ze . Ad ec rease i ns i ze h as l o ng b een a ccep ted a s av a l id c r i te r ia f o rs epa ra t ing mo rpho log ica l ly " w i ld " f r om " domes t ic " c a t t le , s heep a nd g oa ts i na rchaeo log ica l f auna l a na lyses , a l though t he a ssump t ion o f ad i rec t r e la t ionsh ip b e tween a n ima l s i ze a nd p a leoeconom ic s ys tems ( hun t ing v s . h erd ing ) s eems e x t reme ly d ub ious ( Ja rman 1 969 ; J a rman a nd W i lk inson 1 972 ; P ayne 1 972a ; S axon 1 974 ) . R ega rd less , s uch s i ze r educ t ions m igh t p o ten t ia l ly make c a lcu la t ions o f s l augh te r y i e ld b ased u pon mode rn u n imp roved b reeds o f c a t t le , s heep a nd g oa ts s l igh t ly l ower t han t hose e xpec ted i n t he p reh is tor ic p as t . A t p resen t , t h is p oss ib i l i ty i se x t reme ly d i f f icu l tt oe va lua te . I n t he c ase o f c a t t le , i ti sa ppa ren t t ha t t he s i ze r anges o f r eg iona l B os p r im igen ius p opu la t ions v a r ied w ide ly i nt he p reh is to r ic p as t ( G r igson 1 969 ; J a rman 1 969 ; S axon 1 974 ) . J a rman s umma r ized t he s i tua t ion a sf o l lows ( 1969 :255 -256 ) : T he s t udy o f p reh is tor ic c a t t le o ve r a n e x tended a rea i nb o th t ime a nd s pace s ugges ts t ha t t he e x is tence o f sma l l c a t t le may h ave b een a r ecu r ren t p henomenon wh ich was n o t l i nked i n a l c ases , o r e ven n ecessa r i ly i nt he ma jo r i ty o fc ases , t od omes t ica t ion . Sma l l i n d iv idua ls e x is ted i nt he P le is tocene p opu la t ions o f N or th A f r ica , S ou th - Wes t A s ia , a nd S ou th -Eas t E urope , a ppa ren t ly a c ons is ten t e l emen t , a t l eas t f r om E em o nwa rds , i np opu la t ions o therw ise s im i la r t o t he c on tempo ra ry p opu la t ions o f t he N o r th T empe ra te E u rope b i o tope . F ur the r : T here i sn o o s teo log ica l e v idence t ha t t he sma l l c a t t le o ft he P le is tocene a nd e a r ly P os t -g lac ia ld epos i ts o ft he E as t Med i te r ranean a re d i f fe ren tf r om t hose o f t he s ame s i ze i nt he N eo l i th ic o ft ha t a rea o ro fN or th - Wes t E u rope ; n o r i st he re a ny e v idence t ha t t he s i ze c hanges s een i nc a t t le we re n ecessa r i ly c aused b y f ac to rs o the r t han t hose o pe ra t ing i n many o the r s pec ies a t t he s ame a nd d i f feren t p e r iods . P a r t icu la r ly s i gn i f ican t i nt h is r e spec t i sS axon 's ( 1974 :35 -36 ) c ompa ra t ive a na lys is o ft he r ange a nd mean l eng th o f B os p r im igen ius a s t raga l i f r om t he K eba rah C ave d epos i ts ( Uppe r P a leo l i th ic t h rough N a tu f ian p e r iods ) , P re -po t te ry N eo l i th ic J e r icho , K o rn Ombo , a nd s i tes i n D enma rk a nd H ungary . S axon f ound n o e v idence o f c a t t le s i ze c hange i nt he 1 0 ,000 y ea r K eba rah r eco rd , b u t d id n o te t ha t t he c a t t le p opu la t ions o f p os t -g lac ia l D enma rk a nd l a te N eo l i th ic H unga ry we re s i gn i f ican t ly l a rge rt han a ny o ft he o the r c a t t le p opu la t ions s t ud ied . I t w ou ld s eem t ha t r educed s i ze was a l ready a n e s tab l ished t r a i t i nv a r ious c a t t le p opu la t ions l o ng b e fore t he e vo lu t ion o fp as to ra l s t ra teg ies b ased u pon t he i r h usband ry . 2 3

E ven s o , J a rman 's ( 1969 :255 , F i gu re 9 ) o s teo log ica l a na lyses d o s ugges tt ha t a2 0% s i ze r educ t ion t o t he mode rn mean d id o ccu r i n E as t Med i te r ranean c a t t le p opu la t ions b e tween r ough ly 1 0 ,000 a nd 5 ,000 BC , a s b ased u pon t h i rd mo la r , me taca rpa l , me ta ta rsa l , r ad ius a nd c a lcaneum measu remen ts . A n a pp rox ima te 2 0% r educ t ion i ns i ze b e tween t he L owe r /M idd le P a leo l i th ic a nd N eo l i th ic c a t t le p opu la t ions o f N or th A f r ica i s a l so a ppa ren t i n mean measu remen ts o ft he d i s ta l w id th a nd t h ickness o f me tapod ia ls , l e ng th a nd d i s ta l w id th o ff i rs t p ha langes , a nd t he d i s ta l c ondy la r w id th o f h ume r i ( A . Sm i th 1 980a :494 -496 , F igures 2 0 .2 t h rough 2 0 .5 ) . S im i la r s i ze r educ t ions a re a l so a ppa ren t b e tween P re -po t te ry N eo l i th ic a nd B ronze A ge a n ima l p opu la t ions a t t he J e r icho t e l l ( C lu t ton -B rock 1 979 ) . F rom c ompa r isons b e tween t he mean measu remen ts o f me taca rpa ls , p ha langes , a nd a s t raga l i , as i ze r educ t ion o f a round 1 4 t o 1 8% i nc a t t le s i zes a tJ e r icho b e tween t hese p e r iods i ss ugges ted ( C lu t ton -B rock 1 979 :148 , T ab le 5 ) . A n a pp rox ima te 2 0 t o2 2% r educ t ion i ng oa t s i zes a tJ e r icho i sa l so a ppa ren t f r om mean measuremen ts o f t he d i s ta l w id th o f me taca rpa ls a nd t he l e ng th o f p ha langes , wh i le mean s c im i ta r h o rn c o re measu remen ts s ugges t as i ze r educ t ion o fa round 2 9% ( 1979 :153 , T ab le 6 ) . H oweve r , wh i le C lu t ton -B rock a l so o bse rved s im i la r r educ t ions b e tween t hese t empo ra l p e r iods i nt he measuremen ts o f s heep l i mbs , s he s pec i f ica l ly n o ted t ha t o n ly t he l i mbs o f p reh is to r ic s heep wou ld h ave b een l o nge rt han mode rn s heep , wh i le a c tua l b od i ly p ropor t ions wou ld h ave r ema ined s im i la r t o t he w i ld Ov is o r ien ta l is ( 1979 :153 ) . I ti st he re fo re u nc lea r whe the r t he s i ze r educ t ions a ppa ren t i nl i mb b ones b e tween t he o the r P re -po t tery N eo l i th ic a nd B ronze A ge a n ima l p opu la t ions a tJ e r icho n ecessa r i ly r e f lec t o ve ra l l s i ze r educ t ions u nde r h uman h usbandry , o r p r ima r i ly r educ t ions i nl i mb l e ng ths . T he f ew I r on A ge a nd B yzan t ine p er iod c a t t le a nd g oa t r ema ins f r om J e r icho a re g ene ra l ly e qua lt oo r l a rge rt han e a r l ie rp e r iods ( C lu t ton -B rock 1 979 :148 , T ab le 5 , 1 53 , T ab le 6 ) . C ompa r isons b e tween t he N eo l i th ic a nd mode rn c a t t le r ema ins r ecove red f r om t he T i lems i V a l ley o f Wes t A f r ica s eem ing ly i n d ica te t ha t t he mean d i s ta l c ondy la r w id th o f mode rn h ume r i a re a pp rox ima te ly 1 3% sma l le r t han i nt he N eo l i th ic p opu la t ions , wh i le t he mean o f mode rn p ha langes i s s l igh t ly l a rge r ( A . Sm i th 1 980a :494 , F i gu re 2 0 .3 , 4 96 , F i gu re 2 0 .5 ) . T hese o bserva t ions , c oup led w i th t he l a ck o f c ompa rab le o s teo log ica l s t ud ies o n t he mode rn c a t t le , s heep a nd g oa t b reeds f r om wh ich s l augh te ry i e ld d a ta w i l l b ed e r ived , makes a n a ccura te e va lua t ion o f t he a pp l icab i l i ty o f t h is d a ta t ot he p reh is to r ic p as t e x t reme ly d i f f icu l t . H owever , w i th t he e xcep t ion o f mo rpho log ica l v a r ia t ions d ue t o ap o ten t ia l ly h i gh p l ane o f n u t r i t ion u nder mode rn c ond i t ions o fi n tens ive h usband ry , i ts eems r easonab le t oa ssume t ha t t he r ange o f mea t , o rgan , f a t , a nd b l ood y i e lds o b ta ined f r om u n imp roved c a t t le , s heep a nd g oa t s pec ies u nde r c ond i t ions o f e x tens ive h usband ry i nt he e thnog raph ic p resen t wou ld m in ima l ly r e f lec t t he p o ten t ia l l owe r r ange o f s uch y i e lds i nt he p reh eo r ic p as t . T o p ar t ia l ly a ccoun t f o r p oss ib le r educ t ions i n mean s l augh te r y i e lds b e tween mode rn u n imp roved b reeds a nd t he i rp reh is to r ic p rogen i to rs , c a t t le y i e lds w i l l b e i n c reased 2 0% a nd s heep a nd g oa t y i e lds 3 0% i ns ubsequen t c a lcu la t ions o fp reh is to r ic p as to ra l s t ra teg ies . I nv i ew o ft he a va i lab le d a ta , t hese a d jus tmen ts s eem e x t reme ly g ene rous .

P hys io logy T he p hys io log ica l t r a i ts mos t s i gn i f ican t i n t he p resen t a na lys is o f e thnog raph ic a nd p reh is to r ic p as to ra l p roduc t iv i ty i n vo lve t he c ompa ra t ive p hys io logy o fl a c ta t ion a nd a ssoc ia ted a t t r ibu tes a f fec t ing t he a b i l i ty o f h umans t o p rac t ice d a ir y ing . U n for tuna te ly , t he e phemera l n a ture o f ma te r ia l e v idence a ssoc ia ted w i th p reh is to r ic d a iry ing p rec ludes a d i rec t a rchaeo log ica l a pp roach t o t he a n t iqu i ty o f m i lk e xp lo i ta t ion . A t p resen t , t he e a r l ies t ma te r ia l e v idence f o r d a i ry ing a c t iv i t ies i nt he S aha ra , E gyp t , a nd Mesopo tam ia d a tes r ough ly f r om 4 ,000 t o 2 ,900 B .C . ( S imoons 1 971 , 1 979 ) . T hus , wh i le s ome s cho la rs s ugges t t ha t m i lk p roduc t ion was a n e ssen t ia l a t t r ibu te o ft he d omes t ica t ion p rocess , a tl e as t f o rg oa ts a nd c a t t le ( e .g . , B anks 1 984 :237 ; H a fez 1 968a :203 ; Wendor f a nd S ch i ld 1 980a :278 ; Z eune r 1 963 :152 , 2 4 1) , o the rs s ugges t t ha t w i ld s pec ies i ng ene ra l p roduce o n ly s u f f ic ien t q uan t i t ies o fm i lk t o r ea r t he i r own y oung , a nd t ha t i n c reased m i lk p roduc t ion a nd d a i ry ing we re n ecessa r i ly t he r esu l t o f p os tdomes t ica t ion s e lec t ive b reed ing ( e .g . , B ököny i 1 976 :22 ; F agan 1 986 :2 19 ; D . H a r r is 1 977 :229 -231; H e r re 1 963 :235 ; H o le 1 984 :57 ; S he r ra t 1 981:276 ; Wa tson 1 979 :7 1 , 9 3 ) .

2 4

R ecen t ly , S he r ra t ( 198 1 , 1 983 ) h as a dvanced t he p os i t ion t ha t a l h erd s pec ies w e re f i r s t e xp lo i ted f o r mea t p roduc t ion , a nd o n ly s ome m i l lenn ia l a te r f o r m i lk a nd o the r " seconda ry " p roduc ts . A cco rd ing t o S he r ra t , t h is " seconda ry p roduc ts r evo lu t ion " d id n o t o ccur u n t i l t he f our th m i l lenn ium B .C . , a n i n te rp re t ive p os i t ion t ha t h as b een a dop ted b y many a rchaeo log is ts ( e .g . , D av is 1 976 , 1 984 ; Ga r ra rd 1 984 :117 118 , 1 26 ; L evy 1 983 ; L evy a nd A lon 1 983 ; R osen 1 984 , 1 985 , 1 987 ) . Ad i rec t a t temp t h as a l so b een made t ob u t t ress S he r ra t 's p os i t ion t h rough r ad iog raph ic e v idence o ff ou r th m i l lenn ium o s teopo ros is i nt he me taca rpa ls o fo l de rs heep a nd g oa ts i n t he a bsence o f H a r r is l i nes o r r i cke ts i n j u ven i les , i n te rp re ted a s e v idence o f " con t inued c a lc ium d ra in t h rough p ro longed m i lk ing a nd r epea ted p regnanc ies " ( Sm i th a nd H o rw i tz 1 984 :468 ) . H oweve r , a s n o ted b y C hang a nd K os te r ( 1986 :108 , 10 ) , t h is s t udy c anno t b e a ccep ted a s d e f in i t ive d ue t oo u r p oo ru nde rs tand ing o ft he e t io logy o fo s teopo ros is i nh e rd a n ima ls ( Baker a nd B ro thWe l l 1 980 ; Z ede r 1 978 ) . F ur the r , t he re a re e x tens ive d a ta w h ich s eem ing ly i n d ica te t ha t t he p o ten t ia l o fd a i ry ing s hou ld h ave b een a ma jo rf ac to r i nt he t empo ra l a nd s pa t ia l e vo lu t ion o fp as to ra l s ubs is tence s t ra teg ies . Wh i le t he s ugges t ion t ha t l a c ta t ing f ema les i nw i ld h e rds o n ly p roduce s u f f ic ien t q uan t i t ies o f m i lk f o r t he i r i mmed ia te o f fsp r ing i sg enera l ly c o r rec t , t he f ac to rs wh ich r egu la te t h is r e la t ionsh ip mus t b e u nde rs tood . D ur ing t he c ourse o f l a c ta t ion , t he mamma ry g land c e l ls modu la te b o th t he c ompos i t ion a nd y i e ld o f m i lk t oc omp ly w i th t he c hang ing r equ i remen ts o f t he y oung ( Ho l lmann 1 974 :4 ) , a nd a n i n c rease i n t he n umbe r o f o f fsp r ing c an r esu l t i n i n c reased l e ve ls o f m i lk p roduc t ion ( Tucke r 1 974 :315 ) . F ur the r , i ti sk nown t ha t b o th t he q uan t i ty o fm i lk p roduced a nd t he d u ra t ion o fl a c ta t ion may b e f avo rab ly a l te red i ns pec i f ic h e rd s pec ies t h rough h uman m i lk ing a c t iv i t ies ( Cow ie e t a l . 1 980 :13 ; Mepham 1 983 :6 ; S h inde 1 978 ) . A l though mamma ls d o h ave i n he r i ted l i m i ts o n t he q uan t i t ies o f m i lk wh ich t hey c an r ea l is t ica l ly p roduce i n ag i ven p e r iod o ft ime , t he a c tua ly i e ld a nd ma in tenance o fm i lk s ec re t ion i ss pec i f ica l ly d ependen t u pon n euroendoc r ine s t imu l i a nd me tabo l ic f ac to rs ( Cow ie e t a l . 1 980 ; Mepham 1 983 ; S chm id t 1 97 1; S h inde 1 978 ; T ucke r 1 974 ) . N eu roendoc r ine s t imu l i a re r e la ted t o t he i n tens i ty o f s uck l ing o r h uman m i lk ing a nd t he i r e f fec ts o n h o rmone s ec re t ion r a tes , wh i le me tabo l ic f ac to rs a re r e la ted t od i e ta r y i n take a nd r ou tes o fa ss im i la t ion a nd d i v is ion o f n u t r ien ts b e tween d i f fe ren t o rgans a nd t i ssues o ft he b ody . S ign i f ican t ly , A r is to t le ( H is tor ia A n ima l ium 1 1 1 .2 1) was c l ea r ly a wa re o ft he r e la t ionsh ip b e tween a n a n ima l 's p l ane o fn u t r i t ion a nd i t s m i lk y i e ld ( 1949 :522 ) , a sw e re Wa l te ro fH en ley ( The H usband ry 8 7 , 8 8 ) a nd t he a nonymous a u tho ro fa no the r med ieva l t r ea t ise o n e s ta te managemen t a l so k nown a st he H usband ry ( 33 ) i n t he t h i r teen th c en tury A .D . ( s ee O sch insky 1 97 1 :330 -333 , 4 3 1) . G iven a pp rop r ia te n eu roendoc r ine s t imu l i ( i .e . , s uck l ing a nd /o r ,m i lk ing ) , t he p lane o f n u t r i t ion o ft he l a c ta t ing f ema le h as b een r epea ted ly s hown t oh ave ma rked e f fec ts o n m i lk y i e ld ( A rms t rong a nd P resco t t 1 971 :361; F la t t a nd Moe 1 971:342 ; S chm id t 1 97 1:191-195 ) . A s e ne rgy i n take r i ses , m i lk y i e ld i n c reases t owa rds t he a n ima l 's i n he r i ted p o ten t ia l , a l though a ta d im in ish ing r a te g ene ra l ly a ssoc ia ted w i th a n i n c rease i nl i vewe igh t g a in . Among h e rd s pec ies , t h is r e la t ionsh ip h as b een s pec i f ica l ly n o ted among u n imp roved b reeds o fc ame ls ( B r i t ish N ava l I n te l l igence 1 946 :423 ; K noess 1 977 :4 1-42 ; Mus i l 1 927 :67 , 1 928 :88 -89 ; N ico la isen 1 963 :63 ; R ao e ta l . 1 970 :73 ; T o r ry 1 973 :73 ; Y as in a nd Wah id 1 957 :293 ) , s heep a nd g oa ts ( Ma l tz a nd S hko ln ik 1 984 ; N ico la isen 1 963 :40 , 4 2 ; R edd ing 1 98 1:167 173 ) , a nd c a t t le ( B reman a nd d e W i t 1 983 :1345 ; D ah l a nd H jo r t 1 976 :144 -146 ; D yson -Hudson a nd D yson -Hudson 1 970 :113 ) , a sw e l l a s r e indee r /ca r ibou ( Wh i te a nd L u ick 1 984 ) a nd r ed d ee r( Loudon a nd K ay 1 984 ) . F o r e xamp le , u n imp roved b reeds o fc ame ls i nt he Wes t A f r ican S ahe l w e re o bserved i n 1 972 t o y i e ld 7 t o 9l i t e rs ( app rox ima te ly 7 .2 t o 9 .3 k g ) o f m i lk d a i ly f o r h uman c onsump t ion d ur ing t he we t s eason when p as tu rage was r e la t ive ly l u sh , b u t o n ly 2t o 4l i t e rs ( app rox ima te ly 2 . 1 t o 4 . 1 k g ) d a i ly d ur ing t he h o t s eason u nde r c ond i t ions o f g rea t ly r educed p as tu rage ( S . Sm i th 1 980 :469 ) . S im i la r ly , K a r imo jong c ows i n U ganda w e re o bserved i n 1 957 -58 t o y i e ld 4 .78 p i n ts ( app rox ima te ly 2 . 3 k g ) o fm i lk d a i ly f o r h uman c onsump t ion n ea rt he e nd o ft he r a iny s eason , b u t o n ly 2 . 13 p in ts ( app rox ima te ly 1 k g ) d a i ly b y t he e nd o f t he d ry s eason ( Dyson H udson a nd D yson -Hudson 1 970 :113 ) . Among r e indee r /ca r ibou , p eak m i lk p roduc t ion i ny ea rs o f p oo r n u t r i t ion c an f a l l t o l e ss t han 5 0% o f t ha t a ch ieved i n mo re f avo rab le y ea rs ( Wh i te a nd L u ick 1 984 :227 ) . S im i la r ly , among r ed d ee r r e s t r ic ted t ou n imp roved , h ea the r -dom ina ted h i l l p as tu re , t o ta l m i lk p roduc t ion amoun ted t o a pp rox ima te ly 1 . 6k g d a i ly a tt he b eg inn ing o ft he i r l a c ta t ion , w i th t o ta l p roduc t iv i ty 2 5

s ubsequen t ly d ec l in ing t o a round 0 .55 k g d a i ly b y 9 0 100 d ays a f te r c a lv ing . B y c on t ras t , r ed d ee r r es t r ic ted t oi mp roved p as tu rage o fp e renn ia l r ye g rass u nde rsown w i th c l ove rp roduced a t o ta l o f a round 2 .0 k g o f m i lk d a i ly a tt he b eg inn ing o ft he i r l a c ta t ion , s ubsequen t ly p eaked a t 2 .2 k g d a i ly a t2 0 -30 d ays a f te rc a lv ing , a nd t hen d ec l ined t o 1 . 4 k g d a i ly a t9 0 100 d ays a f te r c a lv ing ( Loudon a nd K ay 1 984 :236 ) . C lea r ly , m i lk y i e ld i sa n a spec t o f l a c ta t iona l s t ra teg ies w h ich e xh ib i ts c ons ide rab le p l as t ic i ty i nr e sponse t oe nv i ronmen t . G iven t he o bserved e fec ts o f s pec i f ic n euroendoc r ine s t imu l i ( i . e . , s uck l ing a nd /o r m i lk ing ) a nd me tabo l ic f ac to rs ( i . e . , p l ane o f n u t r i t ion ) o n m i lk p roduc t ion , t he o bse rva t ion t ha t w i ld h e rd s pec ies o n ly p roduce s u f f ic ien t m i lk t o ma in ta in t he i r own o f fsp r ing i sq u i te m is lead ing when a pp l ied t o t he p o ten t ia l f o r e a r ly h uman d a i ry ing b ehav io r . C er ta in ly , t he b i o log ica l p o ten t ia l f o rt o ta l q uan t i ty o fm i lk p roduced v a r ies among h e rd s pec ies i na cco rdance w i th p r io r s e lec t ive f ac to rs i nt he e nv i ronmen ts i n wh ich t he i r p rogen i to rs e vo lved , b u t t he d eg ree t o w h ich mode rn h e rd s pec ies may a pp roach t he ir i nhe r i ted p o ten t ia l i s l i m i ted b y t he e nv i ronmen ts i n wh ich t hey n ow e x is t . Wh i le h umans h ave s e lec t ive ly b red c er ta in h e rd s pec ies , p ar t icu la r ly c a t t le , i no rde r t o r a ise t he i r i n he r i ted b i o log ica l p o ten t ia lf o r m i lk p roduc t ion , t h is h as b een a p roduc t o fr e la t ive ly r ecen t i n tens ive h usbandry s t ra teg ies w h ich s ta r ted i n s pec i f ic r eg ions o f Wes te rn E urope d u r ing t he s even teen th c en tury A .D . ( Cow ie e t a l . 1 980 ; F usse l l 1 966a ; P resco t t e ta l . 1 930 ; Whea ton -Sm i th 1 963 ) . E ven s o , i tw as n o t u n t i l t he e i gh teen th c en tury t ha t s e r ious a t temp ts w e re f i r s t made t o i mp rove t he e x is t ing r eg iona l b reeds o fE uropean c a t t le t oi n c rease t he i r m i lk p roduc t ion ( Cow ie e ta l . 1 980 :3 ; P resco t t e ta l . 1 930 :1 ; Whea ton -Sm i th 1 963 :57 -58 ) . A nc ien t a ccoun ts o fp l an t a nd a n ima l h usband ry , i n c lud ing C a to 's D e Ag r iC u l tu ra ( 1954 ) , V a r ro 's R erum R us t icarum ( 1954 ) , a nd C o lume l la 's R es R us t ica ( 1948 ) , s uch t ex ts o n n a tu ra l h i s to ry a s A r is to t le 's H is tor ia A n ima l ium ( 1949 ) a nd P l iny 's N a tu ra l is H is tor ia ( 1956 ) , a nd p oe t ic w orks l i ke V i rg i l 's Georg ics ( 1982 ) a nd L uc re t ius ' D e R erum N a tura ( 1968 ) , a re d evo id o fa ny a ccoun ts , s ugges t ions , o r i n t ima t ions o f s e lec t ive b reed ing t o i n c rease t he e x tan t p roduc t ive t r a i ts o f h e rd s pec ies . T he i r s i lence o n t h is s ub jec t i se x t reme ly s i gn i f ican t , f o rt hey a nd t he i r i mmed ia te a nces to rs w e re c l ea r ly t he b ene f ic ia r ies o f t he a n ima l h usband ry e xpe r ience a nd k now ledge a cqu i red a nd emp loyed b yt he p reh is to r ic c a t t le h e rde rs o fE u rope . C a t t le we re d omes t ica ted e a r ly i n G reece , a nd f i gure p rom inen t ly i n t he r eg ion 's s ubsequen t h i s to ry . S im i la r ly , t he I r on A ge s ubs is tence o f n ume rous p opu la t ions i n hab i t ing t he I t a l ian P en insu la i n vo lved c a t t le h e rd ing c oup led w i th c e rea l a g r icu l ture . T he a ccep ted e tymo logy o f " I ta ly " i si t s d e r iva t ion f r om t he a nc ien t G reek t e rm f o r " bu l ls ," i t a l i , a ppa ren t ly b es t6wed u pon t he p en insu la b y t he G reek c o lon is ts o f S ic i ly a nd s ou the rn I t a ly ( Magna G raec ia ) o n a ccoun t o f t he " numbe r a nd b eau ty " o f i t s c a t t le a nd c a lves ( Va r ro 1 . 5 .3 ) . T h is p as to ra l h e r i tage e ven a ppea rs i nt he L a t in wo rd f o r money , p ecun ia , d e r ived f r om p ecus ( " ca t t le " ) , r e f lec t ing ac u l tu ra l p as t i nw h ich af am i ly 's w ea l th w as measu red i nc a t t le ( Ros tov tzef 1 977 :20 ) . N o t s urp r is ing ly , R oman a u tho r i t ies e x to l lt he v i r tues o fh erd ing c a t t le o n wooded l a nd w i th a bundan t p as tu rage a nd f o l iage f o rf odde r ( Va r ro 1 . 5 .11 ; V i rg i l Georg . 1 .142 ) , s t ress t he v a lue o f e lm l e aves i nt he a bsence o ff odde r c rops , wh i le n o t ing o the r a l te rna t ives i na co rns o r t he l e aves o fa sh , p op la r , a nd o ak ( Ca to 3 0 a nd 5 4 ; C o lume l la V 1 .3 .7 a nd X I .2 .99 ) , a nd p romo te t he u se o f a n ima l manure f o r f e r t i l iz ing c u l t iva ted l a nd ( Ca to 5 .8 a nd 3 6 ; C o lume l la 1 . 5 ; P l iny XV I 1 1 .192 ; V a r ro 1 . 38 ) , a l s eem ing ly r e f lec t ions o ft he a n ima l a nd p lan t h usband ry p rac t ices o f E u rope 's e a r ly f ood p roduce rs ( T roe ls -Sm i th 1 984 ) . E ven A r is to t le ( HA V I I I .7 ) n o ted t ha t c a t t le c ou ld b e f a t tened o n e lm l e aves . T hus , i ft echn iques o f s e lec t ive b reed ing t o i n c rease p roduc t ive t r a i ts i nh e rd a n ima ls h ad b een p rac t iced i nt he p reh is to r ic a nd c l ass ica l wo r lds , i t s eems r easonab le t oe xpec tt ha t t hey wou ld m in ima l ly h ave b een men t ioned i ns uch c l ass ica l t ex ts . S ign i f ican t ly , w h i le A r is to t le ( HA 1 1 .20 , 2 1) e xp l ic i t ly n o ted t ha t s ome a n ima ls we re c apab le o f p roduc ing s uper f luous amoun ts o f m i lk f o r h uman c onsump t ion w h i le o the rs w e re n o t , h e a t t r ibu ted s uch v a r ia t ions t o b o th t he d i f fe rences a ppa ren t i n mamma ry s t ruc tures b e tween s pec ies a nd t o d i ve rs i t ies i nt he i r p as turage . F ur the r , when P l iny ( V I I I .69 .176 ) a t t r ibu ted t he p roduc t iv i ty o f E p i ro te c a t t le , t he mos t l a uded b reed o f c l ass ica l a n t iqu i ty , t o t he h usbandry p rac t ices o ff ab led K ing P y r rhus , i tw as b ecause P y r rhus d i d n o t l e t h i s c a t t le b reed b e fore t he a ge o ff ou r , as ound p rac t ice emp loyed e thnog raph ica l ly b y v a r ious p as to ra l p opu la t ions . T he 2 6

p roduc t iv i ty o f E p i ro te c a t t le i sn owhe re a t t r ibu ted t o h uman s e lec t ive b reed ing . F ur the r , i t w ou ld s eem t ha t t he a nc ien t c a t t le b reeds b es t k nown f o r t he i r h i gh m i lk p roduc t ion , s uch a s t h e l a rge c ows o f E p irus i nn o r thwes te rn G reece ( A r is to t le HA 1 .21 ; C o lume l la V I .1 .1-2 ; P l iny V I I I .70 .176 ; V a r ro 1 . 5 .10 ) a nd t he sma l l c ows o fA l t in ium i n Sw i tze r land ( Co lume l la V I .1 .1-2 ; P l iny V I I I .70 .179 ) , w e re g eog raph ica l ly l o ca ted i ne nv i ronmen ts w i th r e la t ive ly a bundan t n a tu ra l p as tu rage ( s ee e spec ia l ly A r is to t le HA 1 1 .21) . A cco rd ing t oA r is to t le ( HA V I I I .7 ) , E p i ro te c a t t le d i d n o t t h r ive e xcep t i nt he r eg ion o fE p i rus , a nd a t temp ts t or ea rt hem e l sewhe re h ad f a i led . B y c on t ras t , i ti sq u i te c l ea r i nt hese t e x ts t ha t t he t h rus t o f a nc ien t b reed ing t echn iques w as t op e rpe tua te e x tan t d es i rab le t r a i ts t h rough p heno typ ic p a i r ing . A s n o ted b y V a r ro ( 1 .5 .9 ) , " t he i r y oung r ep roduce t he c ha rac te r is t ics o ft he p a ren ts " ( 1954 :37 1) . S im i la r ly , L uc re t ius ( De R er . N a t . I ,6 74 ) n o ted t ha t h e rd a n ima ls s uch a s s heep , h o rses , a nd c a t t le " l i ve a cco rd ing t o t he i r o wn k i nd a nd s eve ra l ly k eep t he n a ture o f t he i r p a ren ts a nd c opy t he i r b ehav io r " ( 1 968 :79 ) . Wh i le i nb reed ing ( = " l i ne b reed ing " ) t o ma in ta in p a r t icu la r ly d es i rab le t r a i ts was u n ive rsa l , a t temp ts a tc ross -b reed ing we re e x t reme ly r a re , b e ing l i m i ted t o s uch i n s tances a s S pan ish ewes c rossed w i th l o ca l w i ld r ams , a nd i mpor ted c ocks c rossed w i th I t a l ian h ens ( Co lume l la V I I .2 .4 ,13 ) . Med ieva l t r ea t ises o n h usband ry a re s im i la r ly d evo id o f r e fe rences t o s e lec t ive b reed ing f o r t he i mp rovemen t o f p roduc t ive t r a i ts t h rough c ross -b reed ing , a l though t he p e rpe tua t ion o f e x tan t t r a i ts t h rough p heno typ ic p a i r ing i s a ga in a ppa ren t ( Osch insky 1 97 1) . Y e t , i ti ss pec i f ica l ly t h rough t he c ross -b reed ing o f p rev ious ly i n b red l i nes , r a the r t han t he f i x ing o f e x tan t v a r ian ts a nd t he c onsequen t r educ t ion o f g ene t ic v a r iab i l i ty d ur ing i n b reed ing i t se l f , wh ich p rov ides t he a dd i t ive g ene t ic v a r iab i l i ty f avo rab le f o r a t temp ts a t p roduc t ive i mp rovemen t t h rough s e lec t ive b reed ing ( F ishe r 1 965 :146 -150 ; H owa rd 1 965 :473 -76 ; P r ice a nd K ing 1 968 :36 , 4 3 ) . T he p r ima ry f ac to rs wh ich s eem ing ly l i m i ted a t temp ts a t c ross -b reed ing f o r t he e nhancemen t o f p roduc t ive t r a i ts i n h e rd s pec ies p r io r t o t he s even teen th a nd e i gh teen th c en tu r ies A .D . we re t he a va i lab le p lane o fn u t r i t ion a nd t he c os ts o f al o ng term c omm i tmen tt o p rog rams o f i n tens ive s e lec t ive b reed ing i n l i gh t o f g ene ra l ly s l ow r a tes o f p roduc t ive i mp rovemen t . E ven u nde rt he i n tens ive h usbandry p rac t ices o ft he R oman p e r iod , t he l a ck o f s u f f ic ien t f o rage f o r h e rd a n ima ls i na u tumn a nd w in te r was f r equen t ly a luded t o b y R oman a u tho rs a s as easona l p rob lem ( Ca to 5 4 .2 ; C o lume l la V I .3 .3 , V I .24 .4 ) . E xcep tf o r r a re l o ca l i t ies i nw h ich g ood a rab le l a nd was a va i lab le i nq uan t i ty , t he p roduc t ion o fv a r ious f o rage c rops f o r h e rd c onsump t ion ( e .g . , c l ove r , a l fa l fa , p an ic g rass , b i t te rv e tch , b road b eans , a nd t u rn ips ) h ad t oc ompe te w i th f ood p roduc t ion f o rd i rec t h uman c onsump t ion . G iven t he c ompe t ing c l a ims o f h uman a nd a n ima l f ood r equ i remen ts f o r l i m i ted a rab le l a nd , t he c l a ims o ft he f o rme rg ene ra l ly r e ce ived p re fe rence . A cco rd ing t o T row -Sm i th ( 1957 :258 ) , i t was s pec i f ica l ly t he i n c reased e f for t a nd a ssoc ia ted c os ts p u t i n to t he d eve lopmen t a nd p roduc t ion o f f o rage c rops f o r l i ves tock i n E u rope d u r ing t he s even teen th c en tury A .D . wh ich f i na l ly r a ised l e ve ls o f s t ock n u t r i t ion t o a p o in t whe re t he max imum p roduc t ive p o ten t ia ls o f i n d iv idua l a n ima ls c ou ld b e a dequa te ly a ssessed . T h is a l lowed f o r s ubsequen t a t temp ts a t s e lec t ive c ross -b reed ing a nd p rogeny t e s t ing i ne f for ts t o e nhance t he i r p roduc t ive t r a i ts . P resumab ly , t he c umu la t ive e f fec ts o f a g row ing u rban ma rke t f o r mea t a nd m i lk a nd a dvances i n t he means a nd f ac i l i t ies f o r t r anspo r ta t ion , t he l a t te rc r i t ica l f o r i n i t ia t ing c omme rc ia l d a i ry ing o pe ra t ions , c rea ted as i tua t ion i nw h ich t he p o ten t ia l e conom ic b ene f i ts o f i n c reas ing h e rd p roduc t iv i ty f i na l ly o u twe ighed t he c os ts o f i n c reas ing ly i n tens ive a n ima l h usband ry s t ra teg ies . I ngo Id ( 1980 :14 1-42 ) s im i la r ly n o ted t ha t wh i le t he r ecen t d eve lopmen t o f a n u rban ma rke t f o r r e indeer mea t h as s t imu la ted t h e d eve lopmen t o f g ove rnmen t p rog rams t o b reed r e indeer i no rde r t o e nhance t he ir mea t y i e ld ing q ua l i t ies , c ond i t ions o f e x tens ive h usbandry h ave p osed t he p r ima ry o bs tac le t o t he s uccess o f s uch e f for ts . E x t re me ly e n l igh ten ing i nt h is c on tex t i sI s aac 's c ommen tary ( 1962 ) o n t he i n t roduc t ion o f E u ropean c a t t le b red f o r h i gh mea t a nd m i lk p roduc t ion i n to A f r ica f o r p u rposes o fl a rge -sca le c r oss -b reed ing w i th n a t ive s t ock a f te rt he d i sas t rous r i nde rpes t e p idem ic o f 1 890 t o 1 896 ( s ee S inc la i r 1 979 :3 -6 ) . A cco rd ing t o I s aac ( 1962 :200 ) , t he c on t r ibu t ion o f t he E u ropean s t ock t o t h e c on fo rma t iona l c ha rac te r is t ics a nd p roduc t ive c apac i ty o fn a t ive c a t t le was a l b u t n eg l ig ib le , d ue p r ima r i ly t o n a tu ra l s e lec t ion u nde r c ond i t ions o fe x tens ive h usband ry f avo r ing a n ima ls o f o ve ra l l r uggedness r a the r t han a n ima ls o f i n d i f fe ren t s t am ina t hough o f h i gher p o ten t ia l a s s ou rces o f mea t a nd m i lk . A s n o ted b y Y ouse f e ta l . ( 1968 :244 ) , t he d eve lopmen t o fn ew , mo re p roduc t ive b reeds o f c a t t le t h rough c ross -b reed ing r equ i res l a rge n umbe rs o f a n ima ls 2 7

managed u nde rc a re fu l ly p lanned a nd i n tens ive b reed ing p rog rams , a nd i ss e ldom p rac t ica l f o r c ommerc ia l p roduc t ion i nd eve lop ing c oun t r ies d ue t o t he c os ts a nd manage r ia l p rob lems i n vo lved . A t temp ts t o e nhance t he p roduc t ion o f n a t ive c a t t le b reeds t h rough i n tens ive s e lec t ive b reed ing r equ i re a l o ng te rm c omm i tmen t t o p rocesses w i th s uch s l ow r a tes o f i mp rovemen t t ha t t he e conom ic r e tu rns a re f r equen t ly d ub ious r e la t ive t o t he i n ves tmen t i n vo lved . F o r e xamp le , i n tens ive b reed ing t echn iques a pp l ied t o n a t ive c a t t le i nE gyp t h ave i n c reased t he m i lk p roduc t ion o ft he s ub jec t h e rd 3 0% , b u t r equ i red 8 0 y ea rs t oa ch ieve ( Asker e ta l . 1 955 ) . F ur ther , t he u se o f s uch a n ima ls u nde r e x tens ive h usbandr y w ou ld r esu l t i nd ub ious b ene f i ts , s i nce e ven t he mos t r ecen t b reeds o f m i lch c a t t le w i l l p roduce l e ss m i lk t han u n imp roved b reeds i ft he ir p lane o f n u t r i t ion i s d ep ressed ( Schm id t 1 971:191195 ) . A s s hou ld b e e xpec ted , e thnog raph ic a t temp ts t o i n c rease m i lk p roduc t ion u nde r c ond i t ions o f e x tens ive h usbandry i n vo lve t he p re fe ren t ia l b reed ing o f b e t te r p roduce rs t h rough p heno typ ic p a i r ing , t he movemen t o f h e rds t o t he b es t a va i lab le p as turage , c ons t r ic t ing t he a ccess o fy oung t o t he i r mo thers u dde r a nd /o r f o rc ib le e a r ly wean ing , a nd f r equen t ly t he s l augh te r o f y oung ma les . G iven t he a ppa ren t r ange o fp l as t ic i ty i nl a c ta t ion e xh ib i ted b yt he h e rd s pec ies c hosen f o r e a r ly h usband ry , t he c umu la t ive e fec ts o fs ubsequen t p heno typ ic p a i r ing u nde r c ond i t ions o f e x tens ive h usband ry p r io r t ot he s even teen th a nd e i gh teen th c en tur ies A .D . p r ima r i ly s erved t o r eg iona l ly s tab i l ize mean s ubs is tence h e rd m i lk p roduc t iv i ty w i th in t he u ppe r r ange o f v a r iab i l i ty e xp ressed b yt he i rw i ld p rogen i to rs . M i lk p roduc t iv i ty b y l o ca l ly a dap ted i n b red h erds wou ld h ave r ema ined l i m i ted b y a va i lab le n a tu ra l f o rage i nt he i r r espec t ive e nv i ronmen ts . H ence , e thnog raph ic d a ta o n t he r ange o f m i lk y i e lds a ch ieved i ns ubs is tence h e rds u nde r c ond i t ions o f e x tens ive h usband ry s hou ld a dequa te ly r e f lec t t he p o ten t ia l r ange o f m i lk y i e lds a ch ieved i nt he p reh is to r ic p as t . I ft he c ompa ra t ive p hys io logy o f l a c ta t ion w as a ma jo r f ac to r i nt he s pa t ia l a nd t empo ra l e vo lu t ion o f p as to ra l ism , t hen v a r ia t ions i nl a c ta t ion s t ra teg ies b e tween h e rd s pec ies s hou ld l a rge ly a ccoun t f o r why h umans h ave f ocussed u pon c e r ta in s pec ies t ot he e xc lus ion o fo the rs i n e s tab l ish ing s ubs is tence h e rds . S ign i f ican t ly , s heep , g oa ts , c ame ls , a nd c a t t le h ave i n he r i ted a n e vo lved f unc t iona l a na tomy o f l a c ta t ion w h ich s pec i f ica l ly i n vo lves a m i lk s t o rage s ys tem c omposed o f l a rge c i s te rns , q u i te d i s t inc t f r om t he mo re p reva len t s ys tem o fa l veo la r s to rage f ound among o the r mamma ls , s uch a s p i gs ( C ross 1 977 :194 195 ; G rosveno r a nd Mena 1 974 :236 ) . I n g oa ts , f o r e xamp le , u p t o 8 0% o f t he t o ta l m i lk y i e ld may b e s t o red i n c apac ious u dde rc i s te rns , wh i le i n mos t s pec ies , a l veo la r m i lk p robab ly a ccoun ts f o r r o ugh ly 7 0 9 0% o f t he s t o red v o lume ( C ross 1 977 :194 -195 ) . I ti s s ugges ted, tha t t h is p hys io log ica l d i s t inc t ion i sc r i t ica l f o r u nde rs tand ing b o th h ow h umans w e re a b le t o p rac t ice d a i ry ing i nt he f i r s t p lace , a nd why t hey b ased t he i rs ubs is tence e fo r ts o nt he s pec i f ic h e rd s pec ies t h ey d id . Among s pec ies w i th c apac ious u dde r c i s te rns , t he m i lk e j ec t ion r e f lex s erves t of i l t he c i s te rns , f r om wh ich m i lk e x t rac t ion may b e a ch ieved a ta ny t i me t h rough a p ump ing a c t ion o n t he t ea t , w he the r b y o fsp r ing o r h umans ( C ross 1 977 :196 197 , 1 99 -200 ) . T h is r e f lex , i n t ia ted t h rough t he r e lease o f o xy toc in , i st r igge red b y t he r ecogn i t ion o f o f fsp r ing , u sua l ly b y s i gh t a nd /o r sme l l ( B lax te r 1 971:58 ; C ross 1 977 :199 ; P e te rs 1 978 :25 ; Wa lse r 1 977 :324 -325 ) . H owever , n o t ing t ha t t he m i lk e j ec t ion r e f lex c an a l so b e c ond i t ioned among s pec ies w i th c apac ious u dder c i s terns , C ross ( 1977 :199 -200 ) s ugges ted t ha t t he h i gh r a t io o f c i s te rn t o a l veo la r m i lk h as m in im ized t he i mpo r tance o ft he n eu rohypophys ica l m i lk -e jec t ion m echan ism i nt hese s pec ies . I nt e rms o ff eed ing a nd ma te rna l b ehav io r , t h is s t ra tegy o f l a c ta t ion s eems a dvan tageous among h erd s pec ies wh ich l i ve i no pen t e r ra in a nd b ea r p recoc ious o f fsp r ing ( Wa lse r 1 977 :323 ) . Among t he l a rge b ov ids , o v ids , a nd c ame l ids , f ema les b ea r p recoc ious y oung w i th f o l low ing b ehav io rs s uch t ha t t he y oung a re a b le t of o l low t he i r mo the r s ho r t ly a f te r b i r th , a nd t he mo the r n eed n o tt o ta l ly i n te r rup t h e rf eed ing b ehav io rt os uck le h e ro f fsp r ing . I nd i s t inc t c on t ras t , o n ly a sma l l p ropor t ion o ft o ta l s t o red m i lk may b e w i thd rawn i nt he a bsence o ft he m i lk e j ec t ion r e f lex among s pec ies l a ck ing c apac ious u dde r c i s te rns ( C ross 1 977 :200 -20 1) . F o r e xamp le , m i lk r emova l among p i gs c anno t b e a ch ieved i nt he a bsence o f t he m i lk e j ec t ion r e f lex , wh i le t he e j ec t ion p hase i t se l f l a s ts a round 1 4 s ec ( B raude a nd M i tche l l 1 950 ) . T he m i lk e j ec t ion r e f lex a ppea rs t o r esu l t f r om t he r e lease o f ap u lse o f o xy toc in f o l low ing r e f lex ly t he i n i t ia l massag ing o f t he t ea ts b y t he l i te r ( C ross 1 977 :200 ; Wa lser 1 977 :324 ) . Among r abb i ts , l e ss t han 1 5% o ft o ta l s t o red m i lk may b e w i thd rawn i nt he a bsence 2 8

o ft h e m i lk e j ec t ion r e f lex , w i th n a tura l s uck l ing o ccu r r ing o nce a d ay a nd l a s t ing 2 -5 m inu tes . T he s uck l ing i n duced m i lk e j ec t ion r e f lex i sa l so e ssen t ia lf o rm i lk r emova l among r a ts , whe re i ti s a ppa ren t ly l i m i ted t o b r ie f , i n te rm i t ten t m i lk e j ec t ions o ccur r ing a t i n te rva ls o f 5 -20 m inu tes . O bv ious ly , m i lk ing a n ima ls w i th a n e vo lved t r a i t o f p redom inan t a l veo la r s to rage , e ven i f p oss ib le , wou ld m in ima l ly b e t ed ious , a nd t he b ene f i ts o b ta ined q ues t ionab le r e la t ive t o t he e f fo r t i n vo lved . A lveo la rs t o rage i ss eem ing ly a dvan tageous among s pec ies s uch a s p i gs a nd d eer , w h ich l i v e i nf o res ted a reas o r whe re t he re i ss ome c ove r( Wa lse r 1 977 :323 ) . Wh i le s uch a n ima ls may b ea r p recoc ious y oung , t hese g ene ra l ly e xh ib i t h i d ing b ehav io rs wh i le t he i r mo the rs move a way t o f eed . Mo the rs t he re fo re d i scon t inue t he i r f eed ing b ehav ior t o p e r iod ica l ly r e tu rn t o s uck le t he i ry oung i nh i d ing p l aces o r" nes ts . " S ign i f ican t ly , u n l ike many c erv ids , m ig ra to ry r e indee r /ca r ibou b ea r p recoc ia l y oung w i th f o l low ing b ehav io rs ( Len t 1 974 :24 ) , a nd d o p ossess sma l l u dde rs w i th a l ow r es idua l m i lk v o lume ( Wh i te a nd L u ick 1 984 ) . T he p oss ib le s i gn i f icance o ft h is p hys io log ica l a t t r ibu te i nt he e vo lu t ion o f e a r ly r e indee r /ca r ibou p as to ra l ism h as n o t b een a dd ressed b y p rev ious s cho la rs ( e .g . , I n go Id 1 974 , 1 980 ; K hazanov 1 984 ; L eeds 1 965 ; P a ine 1 957 , 1 964 ; S poone r 1 973 ) , a ppa ren t ly b ecause r e indee ra re n ow r a re ly e xp lo i ted f o rm i lk . T he o bse rva t ion t ha t v a r ious s oma t ic a nd p sycho log ica l s t resso rs c an e as i ly i n h ib i tt he m i lk e j ec t ion r e f lex ( C ross 1 977 :198 ; G rosveno r a nd Mena 1 974 :238 -243 ) i s a l so c r i t ica l i n u nde rs tand ing t he i n i t ia l h uman a b i l i ty t o m i lk u ngu la te s pec ies w i th c apac ious u dde rc i s te rns . Wh i le t he s t ress c rea ted b y h uman a t temp ts t oe x t rac t m i lk f r om s pec ies w i th a l veo la r s t o rage w ou ld p ose a ma jo r p rob lem b y i n h ib i t ing t he r equ i red m i lk e j ec t ion r e f lex , t h is wou ld n o t b e s o w i th s pec ies p ossess ing c apac ious u dde r c i s te rns s o l o ng a s t he r e f lex h ad b een p rev ious ly t r i gge red t h rough t he d am 's r ecogn i t ion o f h e r o f fsp r ing o r b y c ond i t ion ing . T he s ubsequen t r emova l o fm i lk f r om t he c i s te rn c ou ld t hen b e a ch ieved r ega rd less o fa ny c onsequen t s t ress o n t h ea n ima l . l ti ss uspec ted t ha t , d ue t o t he p o ten t ia l c a lo r ic y i e ld f r om d a i ry ing , v a r ia t ions i n t he c ompa ra t ive p hys io logy o f l a c ta t ion ( e .g . , t he d eg ree o f p las t ic i ty i n m i lk y i e ld , a l veo la r v s . c i s te rn s t o rage , a nd v a r ia t ions i nt he mechan isms g overn ing t he m i lk e jec t ion r e f lex ) we re s i gn i f ican t i nt he s pa t ia l a nd t empo ra l d ec is ion mak ing a ssoc ia ted w i th t he e vo lu t ion o fp as to ra l s t ra teg ies b ased u pon s pec i f ic h e rd s pec ies t ot he e xc lus ion o r l a te a dop t ion o fo the rs . S ome s uppor t f o r t h is s ugges t ion e x is ts i ne xpec ta t ions o n t he r e la t ive a nnua l c a lo r ic c on t r ibu t ion o f m i lk a nd mea t t o t o ta l s heep , g oa t , c a t t le , a nd c ame l h e rd p roduc t iv i ty u nde r c ond i t ions o f e x tens ive h usband ry . A ccord ing t o R edd ing 's d a ta ( 1981 :33 ) , m i lk wou ld a ccoun t f o r a pp rox ima te ly 7 0% o ft he a nnua l c a lo r ic y i e ld e xpec ted f r om f l ocks o fs heep , a nd 7 9% o f t he a nnua l c a lo r ic y i e ld f r om h e rds o fg oa ts . D ah l a nd H jo r t 's d a ta o n c a t t le ( 1976 :175 ) s ugges t t ha t m i lk wou ld a ccoun t f o r a pp rox ima te ly 6 2% o f a nnua l c a lo r ic y i e ld , e xc lud ing c a lor ies o b ta ined f r om b leed ing a n ima ls . D a ta d eve loped i n t he p resen t s t udy a s p resen ted i n A ppend ix B s ugges t t ha t m i lk wou ld m in ima l ly a ccoun t f o r a pp rox ima te ly 5 4% , a nd mo re n o rma l ly 7 5% , o ft he a nnua l c a lo r ic y i e ld e xpec ted f r om a h e rd o fc ame ls . F i na l ly , i n ar ecen t s t udy o f e ne rgy e x t rac t ion among t he N g isonyoka s ubsec t ion o f t he p as to ra l T urkana o f n or thwes t K enya , m i lk was f ound t oc on t r ibu te 6 1% o ft o ta l a nd 8 0% o f l i ves tock -de r ived f ood e ne rgy ( Coughenou r e t a l . 1 985 :230 ) . D ue t o t he t r emendous c a lo r ic c on t r ibu t ion made b y m i lk , i tw i l l b e s hown i nC hap te r 5t ha tt he r e tu rn r a tes f o rp as to ra l s t ra teg ies b ased s o le ly u pon m ea t p roduc t ion a re e x t reme ly l ow r e la t ive t o mea t p l us m i lk s t ra teg ies .

A du l t H ypo lac tas ia : S ign i f ican t i nt he E vo lu t ion a nd E ne rge t ics o fE a r ly P as to ra l S t ra teg ies? A no the r f ac tor f r equen t ly c i ted a ga ins t t he p oss ib le s i gn i f icance o f d a i ry ing i n e a r ly p as to ra l s t ra teg ies i sa du l t l a c tose i n to le rance ( Ga r ra rd 1 984 :118 ; S he r ra t 1 981:276 -277 , 1 983 :94 -95 ; S ebas t ian P ayne , p e rsona l c ommun ica t ion 1 987 ) . D ue t o t he s i gn i f icance wh ich many s cho la rs a t tach t ot h is c omp l ica t ing f ac to r , ab r ie fd i scuss ion o ft he i s sues i n vo lved s eems w a r ran ted .

2 9

L ac tose i s ad i saccha r ide wh ich c anno t b e me tabo l ized u n less i ti sh yd ro lyzed i nt h e sma l l i n tes t ine b y t he e nzyme l a c tase i n to i t s c ons t i tuen t monosaccha r ides , g l ucose a nd g a lac tose , w h ich a re p resen t i ne qua l q uan t i t ies . S ome o ft he g l ucose i st hen u t i l ized l o ca l ly , w h i le t he r es t e n te rs t he b loods t ream a l ong w i th g a lac tose , b o th b e ing s ubsequen t ly me tabo l ized b y t he l i ve r . I ft he amoun to fl a c tose i n ges ted e xceeds t he h yd ro ly t ic c apac i ty o ft he a va i lab le l a c tase i n t he sma l l i n tes t ine , mos t o ft he u nhyd ro lyzed l a c tose e n te rs t he l a rge i n tes t ine , d raw ing wa te r o u t o ft he s u r round ing c e l l s t ruc tu res t o a t ta in o smo t ic e qu i l ib r ium , a nd p o ten t ia l ly f e rmen t ing u nde r t he i n f luence o f b ac te r ia i nt he c o lon . O rgan ic a c ids a nd c a rbon d iox ide w i l l t hen b e p roduced , o f ten r esu l t ing i n b loa t ing , f l a tu lence , b e lch ing , n ausea , c ramps a nd a wa tery , e xp los ive d ia r rhea . H umans g ene ra l ly e xh ib i t h i gh l e ve ls o f i n tes t ina l l a c tase i ni n fancy a nd a s ubsequen td ec l ine i nl a c tase a c t iv i ty a f te rt wo t of ou ry ea rs o fa ge ( hypo lac tas ia ) , w i th r e s idua l a c t iv i ty a t a pp rox ima te ly 5 10 % o f i n i t ia l l a c tase l e ve ls . T h is p a t te rn i sc ons ide red n o rma l t h roughou t t he mamma l ian k i ngdom ( F la tz a nd R o t thauwe 1 977 ; F r ied l 1 981; K re tchme r 1 972 ) . A l though ap a t te rn o fa du l t h ypo lac tas ia i sg ene ra l ly o bserved among a ma jor i ty o fh u man p opu la t ions , t hose w i th p r io r h i s to r ies o f d ie ts c on ta in ing m i lk a nd m i lk p roduc ts f r equen t ly e xh ib i tt he p e rs is tence o f i n tes t ina l l a c tase i n to a du l t l i f e ( Cook 1 978 ; C ook a nd A l -To rk i 1 975 ; F r ied l 1 981; J ohnson e t a l . 1 98 1; K re tchme r 1 972 , 1 977 ; McC racken 1 97 1; S imoons 1 979 ) . Among t hose p opu la t ions w i th h i gh f r equenc ies o ft h is t r a i t a re c er ta in A rab p opu la t ions o n t he A rab ian P en insu la , H am i t ic p opu la t ions i n A f r ica , a nd mos t n or the rn E uropeans a nd t he i r d escenden ts i nN o r th Ame r ica a nd A us t ra las ia . T he s cho la r ly c onsensus i st ha tt he p e rs is tence o fi n tes t ina l l a c tase i n to a dul t l i f e i sa n a u tosoma ld om inan tg ene t ic t r a i tw h ich h as b een f avo red among t hose p opu la t ions w i th a h i s to ry o fc onsum ing m i lk a nd d a i ry p roduc ts . A ssum ing t ha t d a i ry ing b egan b e tween 1 0 ,000 a nd 8 ,000 B .C . , as e lec t ive a dvan tage o fo n ly 1 -3% w ou ld b e s u f f ic ien t t o a ccoun t f o r t he p resen t d i s t r ibu t ion o ft h is t r a i t among mode rn p opu la t ions ( F r ied l 1 981 :39 ; S imoons 1 979 :69 ) . H oweve r , i fd a iry ing d i d n o t d eve lop u n t i l t he f our th m i l lenn ium B .C . , a s s ugges ted b y p roponen ts o ft he " seconda ry p roduc ts r evo lu t ion , " t hen a m uch h i ghe r s e lec t ive a dvan tage w i l l h ave t ob e a ccoun ted f o r . Wh i le a ccep t ing t he g ene ra l p os i t ion t ha t t he p e rs is tence o fi n tes t ina l l a c tase i n to a du l t l i f e i s ag ene t ic t r a i t wh ich h as b een f avo red among p as to ra l p opu la t ions , a l though f o r e ne rge t ic , r a the rt han c ons t i tuen t n u t r ien t r easons (con t ra F l a tz a nd R o t thauwe 1 977 ; H a r r is 1 985 ; S he r ra t 1 983 ) , t he a c tua l s i gn i f icance o ft h is t r a i t i nt he e a r ly e vo lu t ion o fp as to ra l s t ra teg ies i su nc lea r . I n i t ia l ly , t he r e la t ionsh ip b e tween l a c tose ma labso rp t ion , i nw h ich i n d iv idua ls a re u nab le t of u l ly me tabo l ize l a c tose d ue t o i n su f f ic ien t l a c tase l e ve ls , a nd l a c tose i n to lerance , i n wh ich i n d iv idua ls e xh ib i t s ymp toms o f d i s t ress f o l low ing c er ta in l e ve ls o f l a c tose i nges t ion , i sn o t a bso lu te ( Johnson e t a l . 1 981; F r ied l 1 981) . A r e la t ive ly h i gh p ropor t ion o f ma labsorbe rs ( app rox ima te ly 2 0 -30 % ) a ppea rt ob e l a c tose t o le ran t ( asymp toma t ic ) , wh i le a sma l l p ropor t ion o f l a c tose a bso rbers ( l ess t han 2 0% ) a ppea r t o b e i n to le ran t . L ac tose ma labso rp t ion a nd l a c tose i n to le rance may t he re fo re b e d i f fe ren t t hough c o r re la ted p heno types . I fc o r rec t , a d i s t inc t ion mus t b e made b e tween ma labso rp t ion , a s measured b y c l in ica l t e s ts , a nd i n to le rance , a s r epor ted i nl i f e h i s to ry r esponses o rd u r ing c l in ica lt es ts . S pec i f ica l ly , t he s tanda rd t es t ing p rocedure f o r l a c tose ma labso rp t ion a nd /o r i n to lerance i n vo lves ap r io r 1 2 h ou rf as tf o l lowed b yt he i n ges t ion o f5 0 100 gm o fl a c tose i nw a te r . T h is i s e qu iva len tt od r ink ing a pp rox ima te ly 1 -2 k go fm i lk o n a n emp ty s t omach . N o t s urp r is ing ly , s uch d osages may p roduce d i s t ress fu l s ymp toms n o t o n ly i nl a c tase d e f ic ien t i n d iv idua ls , b u t i n s ome i n d iv idua ls who h ave f u l l l a c tase a c t iv i ty . F ur the r , ma labso rbe rs a re mo re l i ke ly t od eve lop s ymp toms o n c onsum ing l a c tose i n wa te rt han o n c onsum ing e qua l q uan t i t ies o fl a c tose i nm i lk ( S imoons 1 979 :65 ; Ga l lagher e t a l . 1 974 :419 ; F r ied l 1 981 :44 ) . F ina l ly , wh i le s t ud ies h ave s hown t ha t n ea r ly h a l f o fa l ma labso rbe rs may d eve lop s ymp toms o f i n to le rance a f te rd r ink ing o n ly 8 o z ( r ough ly 0 .25 k g ) o f m i lk o n a n emp ty s t omach ( F r ied l 1 981 :4 1; S imoons 1 979 :66 ) , o the rs t ud ies h ave s hown t ha t mo re t han h a l fo ft he i n d iv idua ls man i fes t ing i n to le rance d eve lop s ymp toms o n ly a f te r d r ink ing b e tween 0 .5 a nd 1 . 0 l i t e rs ( r ough ly 0 .5 -1 k g ) o f m i lk ( F la tz a nd R o t thauwe 1 977 ) . Wh i le t he c on t inued i n ges t ion o f m i lk d oes n o t a ppea r t o i n c rease l a c tose a bsorp t ion t h rough i n c reased o r s us ta ined l a c tase p roduc t ion , i td oes a ppea r t o d im in ish s ymp toms o f i n to lerance t h rough t ime ( Ransome -Ku t i 1 977 :78 ) . L ac tose t o le rance may a l so b e s i gn i f ican t ly i n c reased when i ti s i n ges ted w i th a mea l ( F r ied l 1 981 :4 1) . S ymp toms o f i n to le rance a re a ppa ren t ly a fec ted b yd i e ta ry h ab i ts , a l though ma labso rp t ion i sn o t . T o le ran t ma labso rbe rs may 3 0

t he re fo re g a in s ubs tan t ia l n u t r i t iona l b ene f i ts f r om m i lk e ven i ft he i rt o ta l e ne rgy u t i l iza t ion f r om t he a va i lab le l a c tose i sl e ss t han t ha t o fa bso rbe rs ( F la tz a nd R o t thauwe 1 977 ; J ohnson e ta l . 1 981 :10 ) . U n for tuna te ly , q uan t i ta t ive d a ta o n l a c tose u t i l iza t ion u nde r r educed l a c tase l e ve ls h as n o t b een f ound . On t he b as is o fa va i lab le d a ta ,t he p o ten t ia l s i gn i f icance o fa du l t h ypo lac tas ia i na dd ress ing t he e vo lu t ion a nd e ne rge t ics o f e a r ly p as to ra l s t ra teg ies i sd i f f icu l t t o a dequa te ly e va lua te . I n i t ia l ly , i ti ss imp ly u nknown whe the r t he p e rs is tence o f h i ghe r t han n o rma l i n tes t ina l l a c tase l e ve ls i n to a du l t l i f e was a n a be r ran t h uman t r a i t w i th l ow f r equenc ies p r io rt ot he e vo lu t ion o f e a r ly p as to ra l s t ra teg ies , o r whe the r i tf o r tu i tous ly a ppea red i ns ubsequen t p er iods . P oss ib ly s i gn i f ican t i nt h is c on tex t i st he o bse rva t ion t ha t asma l l p ropor t ion ( gene ra l ly l e ss t han 1 0% ) o f c e r ta in p opu la t ions w i thou t ap r io rh i s tory o fd a i ry ing ( i nc lud ing C h inese a nd f u l l -b looded N a t ive Ame r icans ) a ppea rt ob e a d r if t l a c tose a bso rbe rs ( F r ied ' 1 981 :40 ; K re tchme r 1 977 ; M cC racken 1 97 1; S imoons 1 979 :70 -7 1) . A s men t ioned i nt he f i na l s ec t ion o ft h is c hap te r , d a i ry ing a c t iv i t ies b y f o rager s s easona l ly e xp lo i t ing s pec i f ic w i ld h e rds t h rough t he u se o fd r ive l i nes a nd c or ra ls may h ave b een i n t ia ted p r io rt ot he e vo lu t ion o fp as to ra l s t ra teg ies . C er ta in ly , mode rn f o rage rs i nB o tswana h ave b een o bserved t o ma in ta in i n d iv idua l g oa ts t e the red n ex t t o t he i r h u ts f o r t he e xp l ic i t p urpose o f o b ta in ing m i lk ( Robe r t K . H i tchcock , p e rsona l c ommun ica t ion 1 984 ) . T he p o ten t ia l u t i l iza t ion o f m i lk c on ta ined i nt he s t omaches o fy oung h e rd a n ima ls a nd f r om t he u dde rs o f a du l t f ema les s l augh tered b y f o rage rs mus t a l so b e c ons idered . P o ten t ia l ly , as e lec t ive a dvan tage f o r t he a du l t a b i l i ty t od i ges t l a c tose may h ave e x is ted among c er ta in f o rag ing p opu la t ions l o ng b e fore t he e vo lu t ion o fp as to ra l s t ra teg ies . F i na l ly , i ti sw ide ly r e cogn ized t ha tt he d i lu t ion o ff r esh m i lk w i th wa te r a nd t he r educ t ion o f l a c tose c on ten t t h rough v a r ious me thods o fp rocess ing c an d im in ish o r n ega te t he s i gn i f icance o f a du l t h ypo lac tas ia i n u t i l iz ing m i lk ( F r ied ! 1 981:40 ; Ga l laghe r e t a l . 1 974 ; McC racken 1 97 1:494 ; S imoons 1 979 :68 ) . T he d i lu t ion o f m i lk w i th wa te r was s pec i f ica l ly s ugges ted b y M cC racken ( 197 1 :494 ) a s p o ten t ia l ly s i gn i f ican t i nr educ ing t he i l -e f fec ts o fl a c tose i n to le rance , p resumab ly d ue t o ar esu l t ing d ec rease i nt he p a r t ic le c on ten t o f i n tes t ina l f l u id r e la t ive t o c e l l f l u id a nd a c onsequen t r educ t ion i no smo t ic a c t ion i nt he l a rge i n tes t ine . T he a dd i t ion o f wa te r t ob o th f r esh m i lk a nd m i lk p roduc ts i se thnog raph ica l ly a t tes ted among t he T ua reg ( N ico la isen 1 963 :126 , 2 22 ) . U n fo r tuna te ly , t he a bsence o fq uan t i ta t ive d a ta o n l a c tose a bso rp t ion u nde r r educed l a c tase l e ve ls p rec ludes a n e s t ima t ion o ft he amoun t o f e ne rgy w h ich a n i n d iv idua l m igh t me tabo l ize f r om l a c tose u nde rt hese c ond i t ions . R ega rd less , t he mos t c ommon p rac t ice wh ich wou ld d im in ish o r' n ega te t he i l -efec ts o f a du l t h ypo lac tas ia i st he p roduc t ion o ff e rmen ted m i lk p roduc ts s uch a s s ou red m i lk , y oghu r t , c ul tu red b u t te rm i lk a nd v a r ious c heeses , a l o f wh ich h ave g rea t ly r educed l a c tose c on ten ts . F o r e xamp le , i th as b een s hown t ha t i n d iv idua ls e xpe r ienc ing mode ra te t os eve re s ymp toms o f l a c tose i n to le rance o n d i e ts c on ta in ing 5 0 gm o fl a c tose d a i ly i nt he f o rm o fn on fe rmen ted d a i ry i t ems e xh ib i ted n o s ymp toms o n d i e ts c on ta in ing 5 0 gm o f l a c tose d a i ly i nf e rmen ted d a iry p roduc ts ( Ga l laghe r e t a l . 1 974 ) . S im i la r ly , Mashona t r ibesmen o f S a l isbury , R hodes ia , e xh ib i ted 1 00% i n to le rance t o l a c tose , a l though t hey w e re a b le t oc onsume l a rge q uan t i t ies o f m i lk s ou red b y at r ad i t iona l me thod ( McCracken 1 97 1 :498 ) . A s d i scussed b e low , t echn iques o f f e rmen ta t ion a re a c tua l ly t he d om inan t me thod emp loyed b y p as to ra l is ts t o p reserve t he i r o the rw ise p e r ishab le m i lk r e sou rces . A l though m i lk i sp e r ishab le when f r esh , i t may b e s to red f o r s hor t p e r iods b y v a r ious me thods o f c on t ro l led f e rmen ta t ion ( sour ing ) a nd s ubsequen t p rocess ing , a nd f o r l o nge r p er iods b yv a r ious me thods o fd ehyd ra t ion . U n for tuna te ly , n o s ys tema t ic s t udy h as a s y e t b een c a r r ied o u t o n t he p rocess ing a nd s t o rage o fp as to ra l d a iry p roduc ts , t he amoun ts p rocessed a nd s t o red , o r t he c ompos i t ion a nd n u t r i t iona l q ua l i ty o f t he i t ems p roduced . I ng ene ra l , t he f e rmen ta t ion ( sou r ing ) o ff r esh m i lk a ppea rs t o b e u n ive rsa l ly p rac t iced b y b o th p as to ra l a nd a g r icu l tura l p opu la t ions i nt he N ea r E as t a nd A f r ica . S ou red m i lk f r om c ows , s heep a nd g oa ts i s a l so f r equen t ly c hurned t o p roduce b u t te r a nd b u t te rm i lk , a l though c ame l 's m i lk , f o r r easons d i scussed b e low , i sn o t . D r ied p roduc ts , h oweve r , a re made f r om t he m i lk o f a l f ou r s pec ies ( Bar th 1 960 :78 ; D ah l a nd H jor t 1 976 :159 ; D ickson 1 951:402 ; D yson -Hudson 1 960b :46 ; D yson -Hudson a nd D yson -Hudson 1 969 :80 ; J enn ings -B ram ley 1 907b :133 ; L eese 1 927 :140 ; N ico la isen 1 963 :221; R ao e t a l . 1 970 :76 ; S . Sm i th 1 980 :469 ; Swee t 1 974 :104 ; T o r ry 1 973 :134 ; Wa tson 1 979 :110 ) . 3 1

D u r ing m i lk f e rmen ta t ion , s e lec t m ic roo rgan isms ( l ac t ic s t a r te rs ) h yd ro lyze l a c tose a nd me tabo l ize i t s c ons t i tuen t monosaccha r ides , p roduc ing l a c t ic a c id a nd a c onsequen t p H u n favo rab le f o r t he g row th o f o the r , u ndes i rab le m ic roo rgan isms . L eng th o f f e rmen ta t ion p ropo r t iona l ly r educes t he l a c tose c on ten t o ft he o r ig ina l who le m i lk , w i th t he c a lo r ic c on ten t d ec reas ing a nd l a c t ic a c id c on ten t i n c reas ing a cco rd ing t o t he amoun t o f monosaccha r ides me tabo l ized . B y r educ ing l a c tose l e ve ls , f e rmen ta t ion w ou ld f ac i l i ta te t he c onsump t ion o fd a i r y p roduc ts b y l a c tose ma labso rbe rs , wh i le s imu l taneous ly c r ea t ing c ond i t ions f avo rab le f o r s ho r t t e rm s t o rage b y c rea t ing a n a c id p H . A l l o ft he mos t i mpo r tan t l a c t ic a c id b ac te r ia i nc omme rc ia l u se t oday we re o r ig ina l ly i s o la ted f r om c u l tu res u sed i n mak ing t r ad i t iona l d a i ry p roduc ts , a nd a re n o t k nown t o e x is t o u ts ide t he d a iry e nv i ronmen t ( Ga rv ie 1 984 :48 ) . T he i r g ene ra l p hy logeny r ema ins u nknown . S ign i f ican t ly , wh i le many l a c t ic s t ar te rs c an me tabo l ize b o th g l ucose a nd g a lac tose , t he c ommon ly u sed S t rep tococcus t hermoph i lus , L ac tobac i l lus b u lgar icus , L b . l a c t is , a nd L b . h e lv i t icus o n ly me tabo l ize t he g l ucose h a l f o ft he h yd ro lyzed l a c tose wh i le r e leas ing g a lac tose i n to t he e x te rna l med ium ( Garv ie 1 984 :53 , 5 7 ; Ma rsha l l a nd L aw 1 984 :71 , 7 7 ) . Y oghu r ts a re f e rmen ted u s ing e qua l r a t ios o f S t r . t hermoph i lus a nd L b . b u lgar icus , " Bu lga r ian " b u t te rm i lk u s ing L b . b u lgar icus a l one , a nd l a ban ( t he c u l tu red m i lk p roduc t c ha rac te r is t ic o ft he N ea r E as t ) u s ing a s ta r te r c omposed o f S t r . t he rmoph i lus , L b . b u lga r icus , L euconos toc l a c t is a nd t wo y eas ts ( Ma rsha l l 1 984 : 166 -67 , 1 69 , 1 76 ) . L euc . l a c t is v a r ies f r om t he l a c t ic s t a r te rs men t ioned e a r l ie r i nf o rm ing g as d ur ing t he f e rmen ta t ion o fg l ucose , a l though i n fo rma t ion a bou t i t s a b i l i ty t o h yd ro lyze l a c tose i sn o t a va i lab le ( Ga rv ie 1 984 :55 ) . S t r . l a c t is , u sed t op roduce b u t te rm i lk i n t he U .S . , I r e land a nd S cand inav ia , c an me tabo l ize b o th g l ucose a nd g a lac tose , a l though i t p r ima r i ly me tabo l izes g l ucose when u sed a s ab u t te rm i lk s t a r te r ( Ma rsha l l 1 984 : 163 -66 , 1 74 ) . I n E as t A f r ica , c ow 's u r ine i su sed t o s our m i lk ( Dyson -Hudson a nd D yson -Hudson 1 969 :80 ) . U n fo r tuna te ly , n o i n fo rma t ion o nt he m ic rob io logy o rb i ochem is try o ft h is p ar t icu la rp rocess h as b een f ound . I tw ou ld s eem , t hen , t ha t t he a c tua l s i gn i f icance o fa du l t h ypo lac tas ia i nt he e vo lu t ion a nd a dop t ion o fe a r ly p as to ra l s t ra teg ies h as b een o ve r ly s t ressed a nd m isunde rs tood . P resumab ly , e a r ly p as tora l is ts , l i ke t he i r mode rn d escenden ts , wou ld h ave p rese rved t he i r o therw ise p e r ishab le m i lk r esources t h rough f e rmen ta t ion p rocess ing , a n a c t iv i ty wh ich wou ld h ave g rea t ly d im in ished o r n ega ted t he e f fec ts o f a du l t h ypo lac tas ia among many i n d iv idua ls . T he a c tua l s i gn i f icance o ft h is c omp l ica t ing f ac to r wou ld t he re fo re h ave b een t o i n c rease t he l a bo r c os ts o fd a i ry ing a s ar esu l t o ff e rmen ta t ion p rocess ing , a nd t od im in ish t he c a lo r ic r e tu rn f r om d a i ry ing p ropor t iona l t ot he l a c tose me tabo l ized b y m ic roo rgan isms . B o th o ft hese f ac to rs c an b e a ccoun ted f o r o n t he b as is o f a va i lab le d a ta , a nd w i l l b e c ons ide red i nt he c a lcu la t ions o f p as to ra l r e tu rn r a tes g ene ra ted i nC hap te r5 . I ns ubsequen t a na lys is , i tw i l lb e a ssumed t ha tt he p r ima ry g oa l o fp as to ra l m i lk p rocess ing i st o p roduce i t ems f o r s hor t a nd l o ng te rm s t o rage w i th a m in ima l l o ss o f c a lo r ic v a lue . T he p rocess ing t echn iques w h ich w i l l b e e xam ined i n c lude f e rmen ta t ion ( sou r ing ) , t he c hurn ing o f b u t te r a nd b u t te rm i lk f r om s oured m i lk , a nd t he p roduc t ion o f d r ied m i lk i t ems . H oweve r , t r ad i t iona l c heeses p roduced b y a dd ing r enne t ( ob ta ined f r om t he a bomassum o fs l augh te red k i ds a nd l ambs o r f r om t he i r p ar t ly -d iges ted s t omach c on ten ts ) t o wa rmed m i lk a s ac urd l ing a gen t a nd s ubsequen t ly s hap ing a nd d ry ing t he r esu l t ing c urd ( Bar th 1 96 1:7 ; N ico la isen 1 963 :225 ; Swee t 1 974 :103 -4 ; Wa tson 1 979 :110 ) , w i l l n o t b e c ons ide red . F i rs t , t he p roduc t ion o f t r ad i t iona l c heeses i s n o t u n ive rsa l among p as to ra l is ts , s i nce t hey a re n o t a ppa ren t ly p roduced i n E as t A f r ica ( Dah l a nd H jor t 1 976 :160 ; D yson -Hudson a nd D yson -Hudson 1 969 :80 ; T o r ry 1 973 :134 ) . S econd ly , t r ad i t iona l c heeses a re f r equen t ly e a ten when n ew ly made ( Bar th 1 96 1 :7 ; N ico la isen 1 963 :225 ; Wa tson 1 979 :110 ) , wh i le t hose p roduced f r om g oa t o r s heep m i lk mus t b e f u r the r p rese rved i nb r ine s o lu t ions ( Swee t 1 974 :104 ; Wa tson 1 979 :110 ) . F i na l ly , i nt he p roduc t ion o ft r ad i t iona l c heeses , at yp ica l w hey w i l l c on ta in 0 .3 % f a t , 0 .9% p ro te in , 4 .9% l a c tose a nd t he wa te r s o lub le v i tam ins B 1 , B 2 ( r ibo f lav in ) a nd C ( Ma r t in 1 970 :43 ) , a l o f wh ich wou ld n o rma l ly b e l o s t s i nce o n ly t he c u rd i sr egu la r ly u t i l ized . A s s uch , t r ad i t iona l c heeses a ppea r t o b e s pec ia l ty d i shes u n less p roduced f o r a ma rke t . N o t s u rp r is ing ly , N ico la isen ( 1963 :225 ) s pec i f ica l ly n o ted t ha t t he A hagga rT ua reg o n ly made s uch c heeses when m i lk was p l en t i fu l .

3 2

AC ross -Cu l tu ra l Mechan ism f o rt he I n i t ia l C ap tu re a nd C on t ro l o fW i ld H e rds I ne a r l ie r d i scuss ion , i th as b een a ssumed t ha t p reh is to r ic h un te rs c ou ld h ave c ap tured a nd i n i t ia l ly c on t ro l led w i ld h e rds i ns uch af ash ion a s t o b e a b le t o i s o la te t he p heno typ ica l ly mos t d es i rab le i nd iv idua ls f o r u se a s t he f ound ing membe rs i ns ubs is tence h erds . On t h is s ub jec t , c u r ren t a nd p as tp rac t ices i nr e indee rh un t ing a nd p as to ra l ism a re i n fo rma t ive . T he mechan ism emp loyed f o r c ap tur ing a nd c on tro l l ing b o th w i ld a nd p as tora l r e indee r /ca r ibou h e rds i n vo lves t he u se o f " hun t ing s u r rounds " o r " t rap c o r ra ls " c omposed o f t wo c onve rg ing b a r r ie rs e nd ing i n ac i rcu la r e nc losu re o r c o r ra l . S uch s t ruc tu res a re u sed i n s easona l r oundups f o r s or t ing o u t t he r e indee r o f d i f fe ren t owne rs , s e lec t ing i n d iv idua ls f o r s l augh ter , a nd f o r h erd c on t ro l d ur ing c as t ra t ion a nd e a r ma rk ing ( I ngo Id 1 980 :116 ) . A rchaeo log ica l e v idence f o r t he u se o f s uch h un t ing s ur rounds i n p re -pas to ra l p e r iods i s w idesp read i nN o rway ( Bar th 1 982 :33 -41 , 1 983 ; V o r ren 1 965 ) , a nd t he i r u se b y n ume rous p opu la t ions f o r h un t ing r e indee r /ca r ibou i sh i s to r ica l ly a nd e thnog raph ica l ly r eco rded a c ross A s ia , N or th Ame r ica , a nd G reen land ( B in fo rd 1 978 :235 -242 , 3 91-392 ; H ea rne 1 9 11:120 -121; I ngo Id 1 980 :56 -63 ; Me ldgaa rd 1 983 :267 ; O sgood 1 936 :25 , 1 940 :251-252 ; S turdy 1 972 :164 ) . S uch h un t ing s ur rounds we re a l so u sed i nt he e xp lo i ta t ion o f n ume rous o ther h e rd s pec ies b y v a r ious p opu la t ions wor ld -w ide , i n c lud ing t he N ea r E as t ( con t ra H o le 1 984 :56 ) . I n N o r th Ame r ica , t hey we re emp loyed o n t he G rea t P la ins i nh un t ing b ufa lo ( G i lmo re 1 924 ) , i n Wyom ing , t he G rea t B as in , a nd t he S ou thwes t f o r h un t ing a n te lope ( Egan 1 917 :238 2 4 1; F r ison 1 978 :254 -257 ; H i l l 1 938 :145 -156 ; K luckhohn e t a l . 1 97 1:8 -10 ; S tewa r t 1 938 :82 , 1 75 , 1 941 :218 -221 , 2 72 , 3 28 , 1 943 :266 -267 , 2 93 , 3 59 ) , a nd i nt he G rea t B as in , t he C o lo rado P la teau , a nd t he S ou thwes t f o r h un t ing d eser t b i gho rn s heep ( G ran t 1 980 :9 -10 ; Mu i r 1 901 ; We the r i l l 1 954 ) . I nn o r theas t I n d ia , t hey w e re s pec i f ica l ly u sed t oc ap tu re w i ld e l ephan t h e rds f o r s ubsequen t d omes t ica t ion ( Zeuner 1 963 :284 ) . I n t he N ea r E as t , t hey were e thno g raph ica l ly emp loyed i nt he h un t ing o f g aze l le h e rds b y b edou in p opu la t ions ( Aha ron i 1 946 ; B urkha rd t 1 831 :220 -221 ; Mus i l 1 928 :26 -27 ) . T he ma te r ia ls u sed i n c ons t ruc t ing t hese d r ive l i nes a nd e nc losures v a r ied w ide ly a cco rd ing t ob o th a va i lab le ma te r ia ls a nd t he a n ima ls i n vo lved . T he c onve rg ing b a r r iers wh ich c hanne led t he h e rd i n to t he e nc losu re o r c o r ra l c ou ld v a ry f r om c on t inuous wa l ls o f b rush o r l o ca l r ock , t h rough r ows o f s paced r ock c a i rns o r s t acks o f s od p i nned w i th s t akes . R ows o f s paced u p r igh t p o les o r b rushwood we re a l so u sed . S lende r p o les w i th r ag p ennan ts we re u sed a s t he g u id ing w ings o fg aze l le t r ap c o r ra ls o bserved i nS y r ia i nA .D . 1 604 ( Legge a nd R ow ley -Conwy 1 987 :93 ) . T he e nc losu res o rc o r ra ls , h oweve r , w e re g enera l ly mo re s ubs tan t ia l , b u t a ga in c ou ld b e b u i l t o fe i the r wood o r r ock i nc on t inuous wa l ls o r o pen f ences . A ppa ren t ly , t he I n ca emp loyed s im i la rt ac t ics i na nnua l ly c ap tur ing a nd s hea r ing w i ld g uanaco a nd v i cuña h erds , a l though h ere , l a rge n umbe rs o f h umans were emp loyed i n r ows r a the r t han c ons t ruc t ing w ood o rs t one b a r r ie rs ( Ga rc i lasso d e l aV ega 1 869 :116 -17 ) . Obv ious ly , i fi twe re n o t f o r h i s to r ic a nd e thnog raph ic a ccoun ts , o ur k now ledge o f t he e x ten t t o wh ich t r ap c o r ra ls were emp loyed wou ld b e e x t reme ly l i m i ted . Wh i le h un t ing e nc losures b u i l t o f s t one may b e e xpec ted t o l a s t f o r m i l lenn ia , t hose o fw ood , b rush , o r s od w ou ld n o t . F o r e xamp le , t he t wo h i s to r ic a n te lope d r ive e nc losu res l o ca ted n or thwes t o f t he G rea t S a l t L ake i nt he e as te rn G rea t B as in w e re b u i l t i nt he m id - t o l a te 1 800s o f j un ipe r b ranches c u tw i th s t ee l a xes ( Raymond 1 982 ) . T oday , t hese h i s to r ic s t ruc tu res a re f r ayed a nd w ea the red , w i th mos t o ft he j un ipe r b ranches l y ing f l a t o n t he g round a nd w i th s ec t ions o ft he g u id ing b a r r ie rs n o l o nge r a ppa ren t . H ence , i ts hou ld b e e xpec ted t ha t t he a rchaeo log ica l r e co rd o fp reh is to r ic h un t ing s u r rounds i sn ecessa r i ly f r agmen ta ry , a nd o n ly p a r t ia l ly r e f lec ts t he a c tua l e x ten t o ft he i r p reh is to r ic u se . F or tuna te ly , p reh is to r ic h un te rs i n t he N ea r E as t s ome t imes u sed a va i lab le s t one i n c ons t ruc t ing t r ap c o r ra ls , a nd f o rt h is r eason , e v idence o ft he i ru se h as b een p rese rved i ns uch d i ver se a reas a s t he S ina i ,t he Wad i A raba s ou th o ft he D ead S ea ( Meshe l 1 974 ) , S y r ia ( Legge a nd R ow ley -Conwy 1 987 :9 1) , t he N a fudh D ese r to fn o r the rn S a 'ud iA rab ia ( Adams e ta l . 1 977 ) , a nd s ou thwes te rn S a 'ud i A rab ia ( Ryckmans 1 976 ) . H oweve r , t he mos t e x tens ive e v idence f o r t he i r p reh is to r ic u se i nt he N ea r E as t c omes f r om t he b asa l t r eg ions o f n o r theas te rn J ordan , w he re t housands o f r ock -bu i l t t r ap c o r ra ls h ave b een r eco rded ( Be t ts 1 981 , 1 982a , 1 982b , 1 983 , 1 983 -84 , 1 984 ; H e lms 1 981:39 -47 , 1 982 :99 -101; Ma i t land 1 927 ; R ees 1 929 ; R i ley 3 3

1 982 ) . T hese o f ten o ccur i nc on t inuous c ha ins o f t r ap c o r ra ls f o rmed b y t he l i nkage o f t he i r r ad ia t ing g u ide b a r r ie rs . H e lms ( 1982 :100 ) made ar ough c a lcu la t ion t ha t b e tween e i gh t a nd t en o f t hese c ha ins c ons is ted o f o ve r 1 , 000 i n d iv idua l t r ap c o r ra ls , r ep resen t ing a pp rox ima te ly 4 ,000 km o f s t one wa l ls a nd a round 7 m i l l ion t ons o f b asa l t , c l ose t o t he we igh t o f K u fu 's p y ram id a tG iza . S im i la r c ha ins o f r e indee rt r aps o ccu r i nN o rway ( Ba r th 1 982 , 1 983 ; V o r ren 1 965 ) . U n for tuna te ly , t he d a t ing o ft he t r ap c o r ra ls i nN o r th A rab ia r ema ins p rob lema t ica l s i nce s im i la r s t ruc tu res w e re i nu se t h rough t he e thnog raph ic p resen t , a nd s i nce v e ry f ew d a tab le s ur face a r t i fac ts h ave b een f ound i na ssoc ia t ion w i th t hem . I na nd a round s ome t r ap c o r ra ls , a r t i fac ts o f t he s i x th o r s even th m i l lenn ia B .C . h ave b een r ecove red ( Be t ts 1 981:246 ; 1 982a :186 ; 1 982b :7 , 3 2 , 1 983 :10 , 1 983 -84 :275 , 1 984 :31; H e lms 1 981:46 -47 , 1 982 :100 1 01) . H oweve r , i ts eems p robab le t ha tt he u se o fs uch s t ruc tures e x tends b ack e a r l ie rt han t h is p e r iod . F i rs t , a l o ft he d a tab le ma te r ia l h as b een r ecove red f r om mo re c omp lex f o rms o ft r ap c o r ra ls wh ich s ome t imes o ve r l ie s imp le rf o rms w i th h eav i ly e roded wa l ls ( Be t ts 1 983 -84 :275 ) . F ur the r , manya t ta s i tes h ave b een f ound b u i l t o n to t he g u ide b a r r ie rs o f t r ap c or ra ls ( Be t ts 1 982a :186 ; 1 982b :7 , 3 1) , a nd t hese s ome t imes i mpede a ccess t o t he c o r ra ls t hemse lves ( R i ley 1 982 :353 ) . U n for tuna te ly , a rchaeo log ica l t es ts o f t hese t r ap c or ra ls a nd a ssoc ia ted b u tche r ing s i tes h ave a ppa ren t ly n o t b een u nde r taken . R ega rd less , i ti ss ugges ted t ha t t he i n i t ia l c ap tu re a nd c on t ro l o fw i ld h e rds c ou ld h ave b een a ch ieved t h rough t he u se o f s uch h un t ing s ur rounds o r t r ap c o r ra ls . S ubsequen t h e rd managemen t c ou ld t hen h ave b een a ch ieved t h rough t he e xp lo i ta t ion o f g enera l p a t te rns o f h e rd b ehav io r . E n t ic ing i n t h is r espec t a re p ho tog raphs o f p reh is to r ic r ock e ng rav ings i n e as te rn J o rdan wh ich d ep ic t t r ap c o r ra ls i n wh ich t he ma in e nc losu re i ss ur rounded b y sma l le r s ubs id ia ry c o r ra ls ( Be t ts 1 984 :P la te 1 3 ; B e t ts , i n R o l le r 1 984 :P la te 3 6 , F igure 5 ) , a nd t he e xce l len t a e r ia l p ho tog raph o f as im i la ra nc ien tt r ap c o r ra l a tD umay ri nS y r ia r ecen t ly p ub l ished b y L egge a nd R ow ley -Conwy ( 1987 :89 ) . T hese e xamp les a re v ery s im i la r t o t he p as to ra l r oundup f ences emp loyed b y Swed ish moun ta in L apps f o r s epa ra t ing t he p as to ra l h e rds o f i n d iv idua ls o nce e n t i re h e rds a re c o l lec t ive ly c ap tured ( l ngo ld 1 980 :117 , F igu re 1 8 ) . A lso e n t ic ing a re r epo r ts o fa nc ien t manya t ta s i tes w h ich s eem ing ly p ossess t he g u id ing b a r r ie rs o f t r ap c o r ra ls ( Be t ts 1 982a :86 -88 , 1 983 :9 ; Ma i t land 1 927 :202 ) . O nt he b as is o fa va i lab le d a ta , i ti ss ugges ted t ha t b o th t he p hys ica l means a nd l a bo rc os ts wh ich c an b e a t t r ibu ted t o t he i n i t ia t ion o f p as to ra l s t ra teg ies we re a l ready i np lace among v a r ious h un te r -ga the re r p opu la t ions p r io r t o t he e vo lu t ion o f p as to ra l s t ra teg ies . H ence , a c ons ide ra t ion o f i n i t ia l " buy in " o r " oppo r tun i ty " c os ts f o r t he s h i f t f r om f o rag ing t o p as to ra l s ubs is tence s t ra teg ies i sd eemed u nnecessa ry i nt he p resen t a na lys is . I ti sa l so s ugges ted t ha t t h rough t he u se o ft r ap c o r ra ls , p reh is to r ic f o rage rs c ou ld h ave p rac t iced d a iry ing l o ng b e fo re t he e vo lu t ion o fp as to ra l s t ra teg ies .

S umma ry E thnograph ic a nd h i s to r ic d a ta o n p as to ra l l a bo r a nd p roduc t iv i ty d o n o t a ppea r t o b e i n app rop r ia te f o r u se i ne s tab l ish ing t he r ange o fp o ten t ia l r e tu rn r a tes f o r p reh is to r ic p as to ra l s t ra teg ies . S ince v a r ia t ions i ns l augh te ry i e lds d ue t op o ten t ia l s i ze r educ t ions i nc a t t le , s heep a nd g oa ts c an b e a ccoun ted f o ro n t he b as is o fa va i lab le d a ta , t he mos tc r i t ica lo b jec t ion wh ich m igh t b e r a ised i n vo lves t he i s sue o f whe the rd a i ry ing was o r was n o ts i gn i f ican t i ne a r ly p as to ra l s t ra teg ies . Wh i le t h is i s sue c anno t b e a dd ressed t h rough d i rec t a rchaeo log ica l e v idence a l one , i ti ss pec i f ica l ly i ns uch c ases t ha t t he t heo re t ica l a pp roach a dop ted h e re c an b e o f v a lue i n a rchaeo log ica l i n te rp re ta t ions . S ince t he c a lo r ies d e r ived f r om m i lk c ons t i tu te a s i gn i f ican t p ropo r t ion o ft he t o ta l e ne rgy y i e ld e xpec ted f r om p as to ra l c ame l , c a t t le , s heep a nd g oa t h e rds , t he r e turn r a tes f o r mea t -on ly a nd mea t p l us m i lk p as to ra l s t ra teg ies a re e xpec ted t ov a ry w ide ly r e la t ive t oo ne a no the r , a nd r e la t ive t o a l te rna t ive o p t ions i n vo lv ing c e rea l h usband ry . H ence , a l terna t ive b ehav io ra l p red ic t ions d e r ived f r om t he r e la t ive e ne rge t ics o f t hese f ood p roduc t ion o p t ions may b e t es ted a ga ins t e x tan t a rchaeo log ica l d a ta i ns uch af ash ion a s t o a dd ress t he i s sue o f whe the r d a i ry ing w ou ld o r wou ld n o t h ave b een s i gn i f ican t i nt he e vo lu t ion o fe a r ly p as to ra l s t ra teg ies .

3 4

I n s ubsequen t c hap te rs , i tw i l l b e s hown t ha t s i nce mea t -on ly p as to ra l s t ra teg ies c ons is ten t ly e xh ib i t l owe rr e tu rn r a tes t han t h ose o b ta ined i nc e rea l h usbandry o n ma rg ina ld ry f a rm ing l a nds , t he a dop t ion o f mea t -on ly p as to ra l s t ra teg ies w ou ld o n ly make s ense i nt he a bsence o f a l te rna t ive c e rea l h usbandry o p t ions o n a g r icu l tura l ly ma rg ina l l a nds . F rom t h is o bserva t ion , i ti s e xpec ted t ha t t he a rchaeo log ica l r eco rd s hou ld r e f lec t t he s eden ta ry o ccupa t ion a nd e xp lo i ta t ion o f ma rg ina l d ry fa rm ing l a nds p r io rt oe v idence f o rt he a dop t ion o f p as to ra l a l te rna t ives i fm ea t w as t he o n ly p roduc t e x t rac ted . T h is p red ic t ion w ou ld a l so b e c ons is ten tw i th " ma rg ina l i ty h ypo theses "f o rt he e vo lu t ion o fp as to ra l s t ra teg ies . B y c on t ras t , i tw i l l b e s hown t ha t mea t p l us m i lk p as to ra l s t ra teg ies o f ten e xh ib i t h i ghe r r e turn r a tes t han a l b u t t h e mos t f a vo rab le o f a g r icu l tu ra l e nv i ronmen ts . I fe a r ly p as to ra l is ts e x t rac ted m i lk a sw e l la s m ea t ,t he a r chaeo log ica l r e co rd s hou ld r e f lec tt h e e vo lu t ion o fp as to ra l s t ra teg ies p r io r t o t he s eden ta ry o ccupa t ion a nd e xp lo i ta t ion o f ma rg ina l d ry fa rm ing l a nds . H ence , i fd a i ry ing w as s i gn i f ican t i nt he e vo lu t ion o f e a r ly p as to ra l s t ra teg ies , e v idence o f p as to ra l s oc ie t ies s hou ld a ppea r i nt he a rchaeo log ica l r e co rd c on tempo raneous w i th , o r j u s t s ubsequen t t o , t he s eden tary e xp lo i ta t ion o f t hose g eog raph ica l ly c i rcumsc r ibed h ab i ta ts w he re t he h i ghes t r e tu rn r a tes f o rc e rea l h usbandry w ou ld h ave b een o b ta ined .

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CHAPTER 3

ESTABL ISH ING COSTS AND B ENEF ITS FOR THE E ARLY CULT IVAT ION OF CEREALS : E THNOGRAPH IC AND H ISTOR IC L ABOR AND P RODUCT IV ITY DATA AS RELATED TO THE P REH ISTOR IC P AST

I n t roduc t ion Wh i le a d i s t inc t ion b e tween i n tens ive a nd e x tens ive f o rms o f a n ima l h usband ry was n ecessa r i ly emp loyed i nt h e p rev ious c hap te r , as im i la rd i s t inc t ion b e tween f o rms o fa gr icu l tu ra l p rac t ices i sd eemed u nnecessa ry f o r e va lua t ing p o ten t ia l t empo ra l v a r ia t ions i nt he l a bo r a nd p roduc t iv i ty a ssoc ia ted w i th c e rea l h usband ry . Wh i le " ex tens ive " a nd " i n tens ive " d es igna t ions a re o f ten a pp l ied i n a n th ropo logy t o d i s t ingu ish h uman f r om a n ima l mo t ive p owe r , o r t o d i s t ingu ish l ow y i e ld " subs is tence a g r icu l ture " f r om h i gh y i e ld " ma rke t o r ien ted a g r icu l ture , " i t w i l l b e s hown t ha t ac ommon r ange o fy i e lds p e r u n i t a rea a re p oss ib le r ega rd less o f mo t ive p ower , o r s ubs is tence o r ma rke t mo t iva t ion . F ur the r , i ti ss ugges ted t ha t s uch p rac t ices a s a pp ly ing n a tura l f er t i l ize r t o s o i ls o r i r iga t ing c rop l a nd , o f ten c i ted a s a t t r ibu tes wh ich d i s t ingu ish " i n tens ive " s ho r t fa l low f r om " ex tens ive " l o ng fa l low s ys tems ( Boserup 1 965 ) , p r ima r i ly s erved t o r ep l ica te e x tan t n a tu ra l c ond i t ions a t f avo rab le b u t g eograph ica l ly c i rcumsc r ibed l o ca l i t ies w he re l e ss l a bo r was r equ i red f o rs im i la rp roduc t ive r e su l ts . I nt he s ubsequen tt e x t , t he t e rm " cu l t iva t ion "w i l l b e s pec i f ica l ly u sed i nr e fe rence t of o rms o f c e rea l h usbandry i nw h ich s o i ls h ave b een p u rpose ly t i led i np repa ra t ion f o r s ow ing o r t r ansp lan t ing . T h is u sage i sc ons is ten t w i th t he p r inc ip le mean ing o ft he Med ieva l L a t in v e rb c u l t ivare ( " to t i l " ) , a s d e r ived f r om t he v e rb c o lere ( " to t i l " ) t h rough t he p a r t ic ip le c u i tus . B y e x tens ion , t he p lan t ing o fc e rea ls t h rough t he u se o f ad ibb le s hou ld a l so b e c ons ide red a s " cu l t iva t ion ." S econda ry mean ings f r equen t ly a t tached t o t he c u l t iva t ion o f c e rea ls , s uch a s " t end ing " o r " p romo t ing , " s hou ld b e u nde rs tood w i th in t he c on tex t o f s o i l d i s turbance . I nt he a bsence o f p u rpose fu l s o i l d i s turbance , t he t end ing o r p romo t ion o f c e rea ls t h rough s uch a c t ions a s s easona l ly b u rn ing p as tu re o rg rass s t ands , weed ing n a tu ra l s t ands , o rs ow ing s eeds o n u n t i l led l a nd , s hou ld b e u nde rs tood w i th in t he g ene ra l c on tex t o f" husband ry " a s p rev ious ly d e f ined . I nt he c u l t iva t ion o fc e rea ls s uch a s whea t a nd b a r ley , l a bo rc os ts w ou ld p r ima r i ly c ons is t o f s o i l p repa ra t ion , s ow ing , h a rves t ing , t h resh ing , w innow ing , s t o rage , a nd g r ind ing t a sks , wh i le t he b ene f i ts wou ld c ons is t o f t he c a lo r ic y i e ld o f t he p rocessed g ra in . I nt he f o l low ing t ex t , p o ten t ia l ly s i gn i f ican t v a r ia t ions i nt hese c os ts a nd b ene f i ts b e tween t he p reh is tor ic p as t a nd t he e thnog raph ic p resen t w i l l b e e numera ted a nd a na lyzed . F i xed c os ts , s uch a s t he i n i t ia l c ons truc t ion o f s ta t iona ry d we l l ings , t he manu fac tur ing o f t oo ls a nd e qu ipmen t , a nd t he ma in tenance c os ts o ft r ac t ion a n ima ls , w i l l n o tb e a dd ressed i nt he p resen t manusc r ip t .

C e rea l C u l t iva t ion L abo rC os ts i nt he E thnog raph ic P resen t a nd t he P reh is to r ic P as t O ve rv iew Wh i le t echno log ica l i n nova t ions s i nce t he E p ipa leo l i th ic h ave d emons t rab ly a f fec ted t he l a bo r c os ts o fs pec i f ic t a sks i nc e rea l c u l t iva t ion , s ome t asks a nd t he i r a ssoc ia ted l a bo r c os ts h ave e xh ib i ted r emarkab le c on t inu i ty t h rough t ime . F o r e xamp le , t he h and s ow ing o f c e rea ls w as s t i l l c ommon i nt he U .S . w e l l i n to t he n i ne teen th c en tury , a s was t he t h resh ing o fw hea t a nd o the r sma l l g ra ins t h rough f l a i l ing b y h and o r t r ead ing w i th a n ima ls ( Co t t re l l 1 955 : 120 ; R og in 1 931:157 -160 , 2 04 -207 ; U .S . D epa r tmen t o f L abo r 1 899 :432 -433 , 4 46 -447 , 4 70 -473 ) . I nt he f o l low ing e va lua t ion o f t he l a bo r c os ts o f c e rea l h usband ry i nt he e thnog raph ic a nd h i s to r ic

p resen t a s r e la ted t ot he p reh is to r ic p as t , a t ten t ion w i l l f o cus u pon t hose v a r ia t ions i ns pec i f ic t asks a ssoc ia ted w i th h uman v s . a n ima l mo t ive p owe r , a nd v a r ia t ions d ue t os uch t e chno log ica l i n nova t ions a s s t one v s . me ta l i mp lemen ts .

S o i lP repa ra t ion E a r ly c e rea l c u l t iva t ion p resumab ly i n vo lved h uman mo t ive p owe ri ns o i l p repa ra t ion u s ing s uch t oo ls a s d igg ing s t icks , h oes , s pades , o rf oo t -p loughs . S ince t he u se o ft hese i mp lemen ts f o rt i lage h as c on t inued i n to t he e t hnog raph ic p resen t , s ome q uan t i ta t ive l a bo rd a ta i sa va i lab le f o r u se i nt he p resen t a na lys is . U n fo r tuna te ly , w h i le l a bo rd a ta o n t he u se o f wooden s pades ( Fo rde 1 931) a nd f oo t -p loughs ( Cu rwen 1 927 ) a re a va i lab le , l abo r d a ta o n s eedbed t i lage u s ing d igg ing s t icks h as n o t b een l o ca ted , wh i le d a ta o n h oe t i lage i s l i m i ted t o t he l o ng h and led d raw h oes w i th f l a t me ta l b l ades c u r ren t ly u sed i nA f r ica ( Haswe l l 1 953 ) . E ven s o , i ti ss ugges ted t ha t t he l a bo r v a r ia t ions a ppa ren t b e tween me thods o f h and t i lage a nd t hose emp loy ing a n ima l t r ac t ion c oup led w i th c u l t iva t ion i mp lemen ts a re f a r mo re s i gn i f ican t f o r t he p resen t a na lys is . B y a t l e as t t he f our th m i l lenn ium B .C . , a n ima l t r ac t ion p l ough ing i mp lemen ts w i th s ymme t r ica l s ha res w e re b e ing u sed f o rt i lage i nt he N ea r E as t a nd E u rope ( E l l ison 1 982 :180 ; F usse l l 1 966b :177 ; H op fen 1 969 :44 -47 ; S he r ra t 1 981 :266 -272 , 1 983 :9 1-92 ; S teensbe rg 1 977 :5 ) . B u i l t o f wood a nd c apab le o f s c ra tch ing a r e la t ive ly s ha l low g roove a cco rd ing t o t he i r s hape , t hese i mp lemen ts d epos i ted l o osened s o i l e qua l ly o n b o th s i des o f t he f u r row c rea ted . C o l loqu ia l ly t e rmed " sc ra tch p loughs " o r " na i l p loughs , " t hese s ymme t r ica l i mp lemen ts a nd t he i r s ubsequen t a symme t r ica l c oun terpar ts p r io r t o t he d eve lopmen t i nt he s even teen th c en tur y A .D . o f mode rn a symme t r ica l p loughs w i th c u rved mou ldboa rds c apab le o f t r u ly i n ver t ing f u r row -s l ices , a re f o rma l ly k nown a s " a rds " o r " a rd p l oughs , " ad e r iva t ion o ft he L a t in a ra t rum . A l though e t hnog raph ic a nd h i s to r ic e f f ic iency d a ta o n a rd t i lage a re a va i lab le , i tmay b e q ues t ioned whe the rv a r ia t ions i na rds t h rough t i me wou ld make s uch d a ta i n app l icab le t ot he p reh is to r ic p as t . N umerous s cho la rs h ave p resen ted a ccoun ts o f t he r eg iona l d eve lopmen t a nd mechan ica l v a r ia t ions o fa rds t h rough t ime ( Abe rg 1 956 ; B i shop 1 936 ; C u rwen 1 938 :143 -147 ; G lob 1 945 ; Gow 1 9 14 ; H a r r ison 1 915 -16 ; H un t ing fo rd 1 932 :237 -233 ; Mann ing 1 964 ; P ayne 1 947 , 1 957 ; S teensbe rg 1 936 ; Z ag reb 1 952 ) , a nd i ti sa ppa ren t t ha t p r ior t ot he d eve lopmen t o f mode rn p l oughs , t he o ve ra l l e f ic iency o fa rd t i lage w ou ld n o t h ave g rea t ly v a r ied e xcep t b y s pec i f ic d ra f t r equ i remen ts a nd s o i l c ha rac te r is t ics . H owever , t he f r equen t a sser t ion t ha t mode rn a symme t r ica l p l oughs w i th c urved mou ldboa rds h ad r ep laced a rds i nE u rope p r io rt o t he s even teen th c en tury A .D . , t hus a f fec t ing t i lage e f f ic iency , s eems u nwa r ran ted . A ppa ren t ly i n f luenced b y n a t iona l is t ic , p o l i t ica l , a nd /o r e thn ic mo t iva t ions ( e .g . , J acob -F r iesen 1 934 ; R y tz 1 935 ; Z ag reb 1 952 ) , o rb y c on fus ion o ve rv a r ia t ions i nr equ i red d ra f t , t he p resence o ra bsence o fc ou l te rs , t he o ccas iona l u se o f whee led f o reca r r iages , a nd f unc t iona l d i f fe rences b e tween f l a t , t i l ted o rb en t s l ide -boa rds o n a rds a s o pposed t o mode rn c urved mou ldboa rds o n p l oughs , t he s i gn i f icance o f v a r ia t ions i n E u ropean a rds p r io r t o t he s even teen th c en tury h as b een b lown o u t o f p ropor t ion t o t he i r b as ic u nchang ing n a tu re f r om t he p reh is to r ic p as t i n to t he h i s tor ic a nd e thnog raph ic p resen t . Wh i le i ti s w ide ly r ecogn ized t ha t t he wooden a rds e thnog raph ica l ly emp loyed i nt he N ea r E as t a nd t h roughou t t he a r id a nd s em ia r id r eg ions o f t he w o r ld h ave c hanged l i t le f r om t he i r p reh is to r ic c oun te rpar ts ( Av i tsur 1 965 ; H op fen 1 969 :47 -55 ; S impson 1 930 :66 ; V a r isco 1 982 ; Wa tson 1 979 :74 -75 ) , t he h i s tory o f a rds i n E u rope h as b een s ub jec t t o n ume rous m isconcep t ions . T he my th ica l s i gn i f icance o f ag rea t S axon o r m ed ieva l h eavy p lough w i th w hee led f o reca r r iage a nd a c urved mou ldboa rd r equ i r ing t he d ra f t o f e i gh t o xen e xemp l i f ies s uch p rob lems . B ased u pon t he f i rs t c en tury A .D . r e fe rence o f P l iny ( XV I 1 1 .47 -48 ) t o b o th t he o ccas iona l u se o f e igh t o xen i n ,I t a ly f o r d ra f t a nd t he " r ecen t " i n ven t ion o f a whee led f o reca r r iage f o rp laumo ra t i ( =" p loum R ae t i "? ) i nR ae t ia G a l l iae ( =t he f oo th i l ls o ft he I t a l ian A lps a nd e as te rn Sw i tze r land ) , c oup led w i th t he med ieva l u se o fa n e igh t o xen p lough t eam a s t he b as is f o rl a nd d i v is ion a nd f i sca l measu remen t i ne s ta te a ccoun t ing , s eve ra l p rom inen t s cho la rs h ave s ugges ted t he g radua l d eve lopmen t a nd d i f fus ion i n E u rope o f h eavy p l oughs w i th mou ldboa rds a nd whee led f o reca r r iages d u r ing t he D a rk a nd M idd le A ges ( e .g . , v an B a th 1 963 :62 -63 ; B loch 1 93 1:49 -65 ; C u rwen 1 927 :280 ; Wh i te 1 940 :151152 , 1 55 , 1 962 :4 1-46 ) .

3 8

S een a s a" r evo lu t ion " i na g r icu l tu ra l t echno logy , t h is " heavy p l ough " h as b een c red i ted w i th c aus ing c hanges i nf i e ld s hapes f r om s qua re t or e c tangu la rs t r ips , s av ing p easan t l a bo rb u t f o rc ing t he p oo l ing o f t he i r o xen i n c oope ra t ive a g r icu l tura l c ommun i t ies l e ad ing t o t he e vo lu t ion o f t he mano r ia l s ys tem , e l im ina t ing t he n eed f o r c rossp lough ing , i mp rov ing f i e ld d ra inage , a l low ing f o rt he e xp lo i ta t ion o fh eav ie r s o i ls , a nd s o i mp rov ing a g r icu l tu ra l p roduc t iv i ty a s t o make p oss ib le t he a ccumu la t ion o f s urp lus f ood s upp l ies l e ad ing t o p opu la t ion g row th , t he s pec ia l iza t ion o f l abo r , i n c reased u rban iza t ion , a nd e ven t he g row th o f " l e isure ." U n for tuna te ly , s uch g rand a sser t ions s eem i l founded . T he u se o f l a rge t eams o f o xen w i th h eav i ly b u i l t a rds o r p l oughs u sed s pec i f ica l ly f o r b reak ing l o ng fa l low l a nd , a nd e spec ia l ly t hose u sed f o r i n t ia l ly b reak ing v i rg in l a nd w i th d ense s o i ls a nd o f ten t ang led r oo t masses , s hou ld n o t b e s u rp r is ing a ta ny p o in t i nt ime o r s pace . T he u se o f b o th h eav i ly -bu i l t w ooden a nd l a te r s t ee l b reak ing p l oughs w i th whee led f o reca r r iages a nd c ou l ters ( t he l a t te r b e ing a n e dged b l ade s e t v er t ica l ly i nf r on t o ft he p l oughsha re t o c u t s od ) o n t he v i rg in p ra i r ie o ft he U .S . d u r ing t he n i ne teen th c en tu ry c ommon ly r equ i red d ra f ts o f s i x t o t we lve o xen y oked i np a i rs , w i th s i x teen a ppa ren t ly b e ing n o c ur ios i ty a nd a s many a s t wen ty four b e ing r epo r ted i no ne c on tempo ra ry a ccoun t ( Rog in 1 93 1:45 -49 ) . F requen t ly , t he c os t o fh i r ing p ro fess iona l o pe ra to rs t od o t h is i n t ia l b reak ing o fv i rg in l a nd e xceeded t ha t o ft he l a nd i t se l f ( Rog in 1 93 1:51) . I nt he l a te e i gh teen th a nd e a r ly n i ne teen th c en tu ry i nt he U .S . , " s t rong " p l oughs r equ i r ing t he d ra f t o f f ou r a nd s ome t imes s i x o xen w e re f r equen t ly u sed f o r b reak ing l a nd wh ich h ad p rev ious ly l a in f a l low f o r s eve ra l y ea rs ( Ed inbu rgh E ncyc lopaed ia 1 832 :336 ; R og in 1 931:5 -7 ) . S ubsequen t p l ough ing , h oweve r , was a ch ieved u s ing sma l le r a nd l i gh te r p l oughs r equ i r ing t he d ra f t o fa t mos t , t wo o xen . N o t s u rp r is ing ly , h und reds o fB ronze A ge p e t rog lyphs i nt he I t a l ian Ma r i t ime A lps d ep ic t ing a rds a nd d ra f t t eams f r equen t ly p or t ray d raf ts o ft wo o xen , a l though a t l e as t s i x e ng rav ings d ep ic t f our o xen y oked i np a i rs , a nd o ne d ep ic ts s i x o xen y oked i np a i rs ( B ickne l l 1 9 13 :40 ,42 ) . Wh i le a n e i gh t o xen p l ough t eam was d emons t rab ly u sed i n med ieva l E u rope a s au n i t o f f i sca l measu re ( F inbe rg 1 95 1; L enna rd 1 945 ; R icha rdson 1 942 ) , t eams o ft wo t ot we lve o xen a re a l so r eco rded , a nd i ti sa ppa ren t t ha t a n e i gh t o xen f i sca l measu remen t d oes n o t p rov ide a p i c tu re o f l i f e i nt he f i e lds . F u r the r , t he n ume rous med ieva l s cenes o fp l ough ing p resen ted i n S teensberg 's h i s tory o f E u ropean p l ough t ypes ( 1936 :263 -265 ) f r equen t ly d ep ic t t eams o f t wo o xen o r h o rses , b u t s ome t imes d ep ic t t eams o ff ou r . A s n o ted b y P ayne ( 1947 :102 106 ) , t he re i sn o e v idence i ni l um ina ted med ieva l manusc r ip ts w h ich c an b e u sed t o s uppor t t he s ugges t ion t ha t ah eavy S axon o r med ieva l p l ough r equ i r ing e no rmous d ra f t h ad l e d t o a r evo lu t ion i n E uropean a g r icu l tu re . Wha t a re d ep ic ted a re a rds w i th s l ide -boa rds ( = " f ins , " " mou ld -s troke rs , " " sh i f t ing -ea rs , " a nd " g round -w res ts " ) , s ome o f w h ich p ossessed whee led f o reca r r iages a nd r equ i red h eav ie rd ra f t . Whee led f o reca r r iages w e re s pec i f ica l ly u se fu l o n t he h eavy p l oughs u sed f o r b reak ing v i rg in o r l o ng fa l low l a nd , a nd w e re c er ta in ly emp loyed w i th a rds i nc e r ta in r eg ions o f E urope d u r ing t he f i r s t c en tu ry A .D . H oweve r , a s s t a ted b y L oudon ( 1883 :97 ) : . . the w hee l i st o b e c ons ide red a s o f n o i mpor tance i ns e t t ing a p l ough f o r wo rk ; b u t p ass ing l i gh t ly o ve rt he s ur face , i tw i l l b eo f ma te r ia l a i d i nb reak ing u p o ld l e ys , o rg round whe re f l i n ts , r ocks , o r r oo ts o ft r ees o ccu r , a nd i nc or rec t ing t he d ep ress ion o ft he s ha re f r om a ny s udden o bs t ruc t ion , a sw e l l a s i nb r ing ing i tq u ick ly i n to wo rk a ga in , when t h rown o u tt owa rds t he s ur face . P resumab ly , t he d eve lopmen t a nd u se o f whee led f o reca r r iages d u r ing t he f i r s t c en tu ry A .D . i n R ae t ia G a l l iae r e f lec ts e fo r ts a tt ha tt ime t ob r ing i n c reas ing ly mo re d i f icu l t a nd p rev ious ly l e ss d es i rab le l a nd u nde r c u l t iva t ion . T he c on t inued u se o f f o reca r r iages i n E urope w ou ld h ave b een p a r t icu la r ly s i gn i f ican t d u r ing t he b reak ing o f v i rg in l a nd r ec la imed d ur ing s ubsequen t p e r iods o f d e fo res ta t ion . D esp i te t he n ume rous m isconcep t ions o f many s cho la rs c once rn ing t he d eve lopmen t o f t he modern p l ough i n E u rope , s ome a u tho rs w e re we l l a wa re t ha t p r io r t o t he s even teen th c en tury A .D . , a n ima l t r ac t ion t i lage i mp lemen ts i n E urope h ad c hanged l i t le f r om t hose emp loyed b y t he a nc ien t G reeks a nd R omans ( e .g . , Ed inburgh E ncyc lopaed ia 1 832 :238 ; H op fen 1 969 :59 ; L oudon 1 883 :389 ; P ayne 1 947 :102 -109 , 1 957 :79 -80 ) . C oncern ing t he 3 9

e vo lu t ion o f mode rn p l oughs c apab le o f i n ve r t ing f u r row -s l ices , i twou ld s eem t ha t Wes te rn E urope a cqu i red t he mode l f o r c ur ved mou ldboa rds f r om t he F a r E as t i nt he s even teen th o r e a r ly e i gh teen th c en tur ies . F l a t , t i l ted , a nd b en t s l ide -boa rds we re f r equen t ly a t tached t o a rds i nE u rope p r io r t o t he s even teen th c en tu ry , b u t wh i le t hese s erved t oc l ea rt he c u t f u r row f r om t he u n t i l led s o i l , t hey d id n o t h ave t he c apab i l i ty o f i n ver t ing f u r row -s l ices . T he mode l f o r t he mode rn c urved mou ldboa rd c ame f r om C h ina i nt he s even teen th o r e a r ly e i gh teen th c en tury , a nd i n i t ia l ly b ecame k nown i n wes te rn E urope i n t he r eg ion o f B raban t , af o rme r d uchy n ow d iv ided b e tween t he N e the r lands a nd B e lg ium ( Hop fen 1 969 :59 ) . D u tch a nd F l em ish p l oughw r igh ts a ppa ren t ly made t he f i r s t a t temp ts a ta t tach ing c urved mou ldboa rds t o a rds a tt ha t t i me ( A l len 1 847 :229 ; Ed inbu rgh E ncyc lopaed ia 1 832 :238 ; L oudon 1 883 :389 ; P ayne 1 957 :83 ) . T hese i n nova t ions b ecame k nown t o p l oughw r igh ts i n n or the rn E ng land i nt he e a r ly e i gh teen th c en tury , a nd i n1 730 , t he f i rs t c omme rc ia l ly p roduced mode rn p l ough w i th c urved mou ldboa rd a nd t r iangu la r f r ame , k nown a s t he " Ro the rham " p l ough b u t o f ten t e rmed t he " Du tch P lough , " was p a ten ted ( A l len 1 847 :229 ; Ed inburgh E ncyc lopaed ia 1 832 :238 -239 ; L oudon 1 883 :391; Ma rsha l l 1 982 ) . I n 1 767 , J ames Sma l l p a ten ted a l i gh t s w ing p lough , i n co rpora t ing t he c urved mou ldboa rd a nd t r iangu la r b ody o f t he R o the rham p l ough w i th f ea tures o ft he o ld S co tch p l ough , t he l a t te r d e r ived f r om t he s ymme t r ica l a rd b u t p ossess ing s l ide -boa rds ( Fen ton 1 976 :40 -42 ; L oudon 1 883 :390 ) . T he s ubsequen t p roduc t ion i nt he l a te e i gh teen th c en tury o f c as t i ron p loughs w i th d es igns d e r ived f r om Sma l l 's s w ing p lough c omp le tes t h is b r ie f h i s tory o f t he e a r ly d eve lopmen t o f mode rn p loughs w i th c urved mou ldboa rds . F ina l ly , t he n o t ion t ha t mode rn p loughs w i th mou ldboa rds e l im ina ted t he n eed f o r c rossp lough ing i sa l so e r roneous . C er ta in ly , s eedbed p repa ra t ion f o r c e rea ls s uch a s whea t a nd b a r ley w hen a rds a re u sed r equ i res a n i n i t ia l b reak ing o ff a l low a nd a tl e as tt wo s ubsequen t c rossp lough ings . T h is p rac t ice was r ecommended i n t he f i rs t c en tury A .D . b y C o lu me l la ( 1 .4 .4 ,8 ,11) , i nt he n i n th c en tury A .D . b yt he I s lam ic a g ra r ian l q im B ab i l ( E I -Sama r ra ie 1 972 :59 ) , i nt he t h i r teen th c en tu ry b y Wa l te r o f H en ley ( Osch insky 1 97 1:325 ) , a nd i nt he f our teen th c en tu ry b y a l -Ma l ik a l -A fda l a l 'Abbas i b n ' A l i i nh i s t r ea t ise o n Y emen i te a g r icu l tu re ( Se r jean t 1 974 :4 1) . T h ree t o f ou r p l ough ings we re a l so c ommon i ns eedbed p repa ra t ion f o r whea t i n E ng l ish a g r icu l tu re u nde rC ha r les I( Lenna rd 1 932 :3 1) . N o t s u rp r is ing ly , a t l e as t t h ree p l ough ings w i th a n a rd a re e thnog raph ica l ly r epor ted a s m in ima l f o rc e rea l a g r icu l tu re i nY emen ( Va r isco 1 982 : 168 ) a nd E th iop ia ( Gams t 1 969 :79 ) , a nd a re a ppa ren t i nt he t ime c os ts r epor ted f o r a rd t i lage i n E gyp t ( Fakbour i 1 972 :35 ; Amma r 1 954 :27 ) . H oweve r , a t l e as t o ne c rossp lough ing was m in ima l ly r equ i red i nc e rea l c u l t iva t ion w i th e a r ly mou ldboa rd p l oughs i no rde r t o c u t a nd p u lve r ize t he i n t ia l f u r row -s l ices c rea ted d ur ing t he b reak ing o ff a l low , a nd at h i rd p l ough ing was r ecommended i ft he s econd p l ough ing d i d n o t p roduce t he d es i red p u lve r iza t ion o ft he s o i l ( S tephens 1 866 :181) . A cco rd ing t o R og in ( 193 1:21) , t h ree p l ough ings w e re f r equen t ly emp loyed i nt he U .S . p r io r t ot he 1 840s f o r s eedbed p repa ra t ion f o r whea t , a nd f our w e re c ons ide red d es i rab le i n c er ta in c i rcums tances . A cco rd ing t ot he Ed inbu rgh E ncyc lopaed ia ( 1832 :335 ) , h eavy c lays s e ldom r ece ived mo re t han t wo p l ough ings f o r s eedbed p repa ra t ion , a l though l o amy o r s andy s o i ls n o rma l ly r ece ived t h ree p l ough ings ( a lso R og in 1 931 :233 ) . H owever , i twas n o t u n t i l t he 1 840s t ha t t he r e la t ive p er fec t ion o f s t ee l p loughs , t he a dven t o f f i e ld c u l t iva to rs , a nd i mp rovemen ts i nt he c ons t ruc t ion o f s p ike too thed h a r rows made mo re t han o ne p l ough ing s upe r f luous i nt he U .S . ( Rog in 1 931 :233 ) . I ti st he re fo re s ugges ted t ha t t he g ene ra l e f ic iency o fa rd t i lage h as r ema ined r e la t ive ly c ons tan t t h rough t he p resen t , v a ry ing o n ly b y r equ i red d ra f t a s ar esu l t o f a rd s i ze a nd s o i l c ha rac te r is t ics . A s s uch , e thnog raph ic a nd h i s to r ic l abo r d a ta o n a rd t i lage s hou ld b e q u i te a pp l icab le t ot he p reh is to r ic p as t .

H arves t ing W i th t he e xcep t ion o fh and s t r ipp ing , t he u se o fs i ck les f o r r eap ing c e rea ls r ep resen ts t he o ldes t a nd mos t l o ng l ived me thod o f h a rves t ing sma l l g ra ins . P r io r t o t he d eve lopmen t o f 4 0

h o rse -d rawn mach inery i nt he 1 830s , t he o n ly s e r ious c ompe t i to rt ot he s i ck le was t he s cy the w h ich , a l though a va i lab le much e a r l ie r , was n o t p opu la rf o r h a rves t ing c e rea ls i nt he U .S . a nd E u rope p r io rt ot he e nd o ft he e i gh teen th c en tu ry , a ppa ren t ly d ue t oi t s d es t ruc t ion o ft he s t raw u sed f o r t ha tch ( Rog in 1 931 :69 -72 ; T ay lo r 1 844 ) . T h roughou t A s ia , t he N ea r E as t , A f r ica , E uropean R uss ia , a nd mos t o f s ou theas te rn E u rope , t he s i ck le h as b een a lmos t e xc lus ive ly u sed f o r h arves t ing c e rea ls f r om t he p reh is to r ic p as t t o t he p resen t d ay ( Hop fen 1 969 :10 11 03 ; R og in 1 931:69 ) . B o th e thnog raph ic d a ta o n c e rea l h arves t ing e f f ic iency u s ing me ta l s i ck les ( Av i tsur 1 966 ; H aswe l l 1 953 ; L e rche 1 968 ) a nd e xpe r imen ta l d a ta o n r eap ing w i th v a r ious f o rms o f a nc ien t s i ck les w i th s t one i n ser ts ( Ko robkova 1 98 1; S teensbe rg 1 943 ) a re a va i lab le f o r u se i nt he p resen t a na lys is . H ence , v a r ia t ions i nh arves t ing e f f ic iency b e tween v a r ious s t one a nd me ta l s i ck les w i l l b e a ccoun ted f o r i nt he p resen t a na lys is .

T hresh ing A s n o ted e a r l ie r , t h resh ing b yf l a i ls o rt he t r ead ing o fa n ima ls was s t i l l c ommon i nt he U .S . u n t i l t he e a r ly n i ne teen th c en tu ry . H oweve r , t he t echn iques o f e a r ly c e rea l t h resh ing r ema in u nknown . A s s ugges ted b y H a r lan ( 1967 :200 ) , t he s epa ra t ion o f emme r a nd e inko rn whea t g ra ins f r om t he i rg l umes may h ave b een a ch ieved b y p ound ing i n wooden mo r ta rs , r e su l t ing i n b o th t h resh ing a nd c oa rse g r ind ing f o r p o r r idge o r g roa ts i n as i ng le o pe ra t ion . U n for tuna te ly , t i med d a ta o nt h is f o rm o fp rocess ing i su nava i lab le . A l te rna t ive ly , e a r ly g ra ins m igh t h ave b een t h reshed b y t he g raddan me thod u s ing f i re a s o bserved b y Ma r t in Ma r t in i nt he Wes te rn I s les o fS co t land a round 1 695 . A woman s i t t ing d own t akes ah and fu l o fc o rn , h o ld ing i tb yt he s t a lks i nh e rl e f t h and , a nd t hen s e ts f i re t ot he e a rs , w h ich a re p resen t ly i n af l ame . S he h as a s t ick i nh e r r i gh t h and , wh ich s he manages v e ry d ex te rous ly , b ea t ing o f f t he g ra in a tt he v e ry i n s tan t when t he h usk i sq u i te b u rn t ;f o ri fs he m iss o ft ha t s he mus t u se t he k i ln , b u t e xpe r ience h as t augh t t hem t h is a r t t o p er fec t ion . T he c o rn may b e s o d ressed , w innowed , g round , a nd b aked w i th in a n h ou r a f ter r eap ing f r om t he g round [ Ma r t in 1 934 :244 ] . As im i la ra ccoun t o ff i re th resh ing w hea t i sc on ta ined i nt he f ou r teen th c en tu ry A .D . t r ea t ise o n Y emen i te a g r icu l tu re b y a l -Ma l ik a l -A fda l a l 'Abbas i b n ' A l i ( Se r jean t 1 974 :42 ) : When o ne wan ts t o make p a rched g ra in o fi t[ whea t ] , l e t h im c u t f r om h a l f -way u pt he s t ock when t he c rop h as j u s t b egun t ot u rn y e l low a nd t he e a r h as f i led . I ti sb ound i n to s heaves , e ach s hea f wha t t he h and c an g rasp , a nd o ne l i gh ts a f i re w i th af l ame b u t n o smoke , a nd b u rns o f ft he h a i rs [ awn ] o ft he e a ru n t i l t hey a re b u rn t u p a nd t h e g ra in i ni ti sc ooked . I ti sr ubbed w i th t he h and o n ap a lm l e a f b aske t -wo rk t r ay , o rb ea ten w i th as t ick i n ag oa t -woo l b ag , t hen w innowed . A s i mp l ied b y C la rk ( 1968 : 112 ) a nd H i l lman ( 1978 :169 ) , s uch p rac t ices m igh t a ccoun t f o r t he f r equen t d i scovery o fc a rbon ized g ra ins i np reh is to r ic d epos i ts . U n for tuna te ly , c onc ise l abo r d a ta o n t h is me thod o f t h resh ing i s u nava i lab le . I ti s s uspec ted , b u t c anno t b e d emons t ra ted , t ha t f i re th resh ing c an b e a ch ieved i nl e ss t ime t han a l te rna t ive me thods , b u t t ha t t h is r e la t ive e f f ic iency w ou ld b e r educed a s ar e su l t o fc ha r r ing a p ropo r t ion o ft he g ra in s o p rocessed . I nt he p resen t a na lys is , t h resh ing l a bo r c os ts w i l l b e b ased u pon Wa tson 's q uan t i ta t ive d a ta o n t h resh ing b y a n ima l t r ead ing i nI r an ( 1979 :82 ) a nd U .S . D epa r tmen t o f L abo r d a ta o n t h resh ing w i th f l a i ls i nt he U .S . i n1 829 -30 ( 1899 :433 , 4 7 1 , 4 73 ) . H oweve r , e x t ra t ime w i l l b e a lowed i no rde rt op a r t ia l ly a ccoun tf o rp reh is to r ic g ra in w h ich was n o tf r ee th resh ing .

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S torage C e rea lc u l t iva t ion g ene ra l ly p roduces l a rge q uan t i t ies o fg ra in w i th in ar es t r ic ted p e r iod o fa y ea r , a nd t he re fo re r equ i res s ome means o ff ood s t o rage . Two b as ic t ypes o f g ra in s to rage s ys tems a re c ommon ly r ecogn ized a cco rd ing t o whe the rt hey a re v en t i la ted o r n on -ven t i la ted ( Food a nd A g r icu l tu re O rgan iza t ion 1 983 :65 ) . V en t i la ted s ys tems a re g ene ra l ly a ssoc ia ted w i th e nv i ronmen ts w i th r e la t ive ly h i gh a tmosphe r ic h um id i ty , a nd a low t he s t o red g ra in t oc on t inua l ly d ry b y e xposu re t o a i r movemen t . S uch s ys tems r educe h ea t ing d ue t o i n c ip ien t m o ld g row th , b u t o f ten e xpose t he g ra in t o f r esh i n sec t a t tack . B y c on t ras t , n onven t i la ted s ys tems a re n o rma l ly a ssoc ia ted w i th r e la t ive ly d ry e nv i ronmen ts , a nd r equ i re t he g ra in t ob e a to rb e low i t s s pec i f ic mo is ture c on ten t p r io r t o s t o rage . S uch s ys tems may a low f o r s l igh t v en t i la t ion , b u t u n l ike p urpose fu l ly v en t i la ted s ys tems , d o n o t a low t he g ra in t o b e e xposed t o f r esh i n sec t a t tack . U nde rg round s t o rage i ns ea led c av i t ies o r p i ts r ep resen ts a c ommon , w idesp read , a nd v ery a nc ien t me thod o f n onven t i la ted s t o rage i nr e la t ive ly d ry e nv i ronmen ts . When f i led w i th g ra in , t hese p a r t ia l ly a i r t igh t c on ta ine rs a pp roach t he c oncep t o f h e rme t ic s t o rage , i n wh ich i n sec ts p resen t i no ro n t he g ra in a re a sphyx ia ted b yt he i ro wn d ep le t ion o f l ow o ve ra l l o xygen c on ten t , w i th r e in fes ta t ion v i r tua l ly i mposs ib le when we l l -sea led ( Food a nd A g r icu l ture O rgan iza t ion 1 983 :65 , 8 8 ) . V a ry ing i nd es ign f r om s qua re o r r ound p i ts l i ned w i th c ha f f a nd ma ts a nd s ea led w i th c haf a nd c ompac ted e a r th , t h rough n a r row -necked c on ica l p i ts p l as te red w i th mud a nd s ea led w i th s t one s l abs a nd c l ay , s uch u nde rg round s t o rage s ys tems a re s t i l l u sed f r om A f r ica t h rough t he N ea r E as t a nd i n to I n d ia . E thnog raph ica l ly , t he B aha rvand o f s ou thwes te rn I r an ( Ho le 1 978 :153 ) , t he Rwa la o fN o r th A rab ia ( Mus h l1 927 :122 ) , t he T ua reg o f t he n or the rn S aha ra ( N ico la isen 1 963 :206 -207 ) , a nd t he K e l T amasheq o ft he S ahe l ( S . Sm i th 1 980 :47 1-472 ) a re r epor ted t os t o re w i ld a nd /o rc ul t iva ted g ra ins i nt h is f ash ion . T hese i n s ta l la t ions a re s a id t o b e e asy t o c ons t ruc t , r equ i re l i t le i nt he way o fb u i ld ing ma te r ia ls , e l im ina te t he r i sk o ff i re , make p i l fe rage d i f f icu l t , a nd c oncea l t he amoun t o fg ra in i n s t o re ( Food a nd A g r icu l ture O rgan iza t ion 1 983 :88 ) . P resumab ly , t hese o r s im i la r s to rage i n s ta l la t ions w e re u sed i nt he p reh is to r ic p as t . U n fo r tuna te ly , n o q uan t i ta t ive l abo rd a ta o n t he i r c ons t ruc t ion h as b een l o ca ted . H oweve r , e ven i nt he i rs imp les tf o rms , t he c ons truc t ion o fs uch i n s ta l la t ions r ep resen ts f i xed c os ts , t he l abo r o f wh ich i ss p read o u t o ver t he n umbe r o fy ea rs f o r w h ich t hey may b e u sed . H ence , ad i rec t a ccoun t ing o f s to rage l abo r i sp rec luded i nt he p resen t a na lys is . H oweve r , a n a rb i t rar y 5% r educ t ion o ft o ta l g ra in y i e ld w i l l b e i n c luded i n s ubsequen t e nerge t ic c a lcu la t ions i no rde r t op a r t ia l ly a ccoun t f o r b o th s t o rage l a bo r a nd t he p a r t ia l l o ss o fg ra in e xpec ted d u r ing s t o rage .

M i l l ing a nd G r ind ing E thnog raph ic a nd e xpe r imen ta l d a ta a re a va i lab le o n t he g r ind ing o f whea t a nd b a r ley f o r f l ou r ( e .g . , C urwen 1 94 1; J asny 1 944 :158 ; Mo r i tz a nd J ones 1 950 :594 ) . U n for tuna te ly , mos t o ft h is d a ta r e la tes t ot he m i l l ing o ff i ne f l ou rf o rt he b ak ing o fb read u s ing r o tary h and -que rns . On ly o ne r e fe rence t ot he m i l l ing o f whea t f l ou r u s ing e a r l ie r h and -que rns h as b een l o ca ted ( Forbes 1 965 :93 ) . H oweve r , e a r ly g ra ins we re p resumab ly c onsumed a s g roa ts o r p or r idge , a nd w ou ld h ave r equ i red l e ss g r ind ing o r p ound ing t han t ha t a ppa ren t i nt he p roduc t ion o f f l ou r . I no rde rt o e va lua te t h is p oss ib i l i ty , R icha rds ' e f ic iency d a ta ( 1951 :104 ) o n t he c oa rse r educ t ion o f m i l le t f o r p o r r idge i n R hodes ia u s ing wooden mo r ta rs w i l l u sed f o r c ompa ra t ive p urposes . A nnua lL abor i nE a r ly I r iga t ion S t ra teg ies A s d ocumen ted i nt he s ubsequen t s ec t ion o n c e rea l p roduc t iv i ty , s upp ly ing wa te r t o c e rea l c rops t h rough i r iga t ion g ene ra l ly r esul ts i ns t ab i l iz ing mean y i e lds w i th in t he r ange o f y i e lds e xpec ted i ne i the rd ry fa rm ing o nf e r t i le s o i ls w i th f a vo rab le r a in fa l l o rr uno f f c ond i t ions , o r t hose a ppa ren t ly o b ta ined i nd öc rue o r " f lood r ecess ion " a g r icu l ture ( Ha r lan a nd P asque reau 1 969 ) . S ince e a r ly c e rea l c ul t iva t ion was s eem ing ly f ocussed o n g eog raph ica l ly c i rcumsc r ibed h ab i ta ts whe re f e r t i le s o i ls we re a ssoc ia ted w i th f avo rab le r a in fa l l , r uno f f , f l ood r e cess ion , o r p e renn ia l wa te r r esou rces r equ i r ing l i t le o r n o i r iga t ion l abor ( Do lukhanov 1 98 1:375 -377 ; H a r lan 1 977 :37 1; H a r lan e t a l . 1 976 :16 ; M i l le r 1 980 :33 1-333 ; Moo re 1 979 , 1 982 ; S he r ra t

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1 980 ) , a n a ccoun t ing o ft h e e f fec ts o fl a te r l a rge -sca le i r iga t ion s t ra teg ies o n t he e ne rge t ics o f c e rea l c u l t iva t ion w i l l n o tb e u nde r taken i nt he p resen t s t udy .

C e rea l P roduc t iv i ty i nt he E thnog raph ic P resen t a nd t he P reh is to r ic P as t Gene ra lC ons ide ra t ions A s p rev ious ly n o ted f o ra n ima l p roduc t iv i ty , i ti sf r equen t ly a ssumed o r a sser ted t ha t c e rea l p roduc t iv i ty h as d rama t ica l ly i n c reased s i nce t he p reh is tor ic p as t a s ar e su l t o fh uman s e lec t ion f o r c rop y i e ld c ha rac te r is t ics a nd o the r d es i rab le t r a i ts . C e r ta in ly , r a tes o f i n c rease i nc e rea l p roduc t iv i ty s i nce t he e a r ly n i ne teen th c en tury A .D . h ave b een d rama t ic , b u t ab ackwa rd e x t rapo la t ion o ft hese mode rn t r ends y i e lds a bsurd r esu l ts . F o r e xamp le , al e as t -squa res t r end o f B r i t ish whea t y i e lds f r om 1 932 t h rough 1 852 i mp l ies a z ero y i e ld a round A .D . 1 575 , a nd a s im i la r e x tens ion o f F rench whea t y i e lds f r om 1 932 t h rough 1 8 15 i mp l ies a z e ro y i e ld a round A .D . 1 625 ( Benne t t 1 935 :12 ) . Obv ious ly , mode rn t r ends i n t he i n c reased p roduc t iv i ty o f c e rea ls c anno t b e p ro jec ted i n to t he p reh is to r ic p as t . l ti sr easonab le t oe xpec t t ha t e a r ly a g r icu l tu ra l is ts i nb o th t he O ld a nd N ew Wo r lds wou ld h ave s e lec ted f o r d es i rab le t r a i ts i nt he i r c e rea l c rops , p a r t icu la r ly i nt e rms o ft he i r p roduc t ive c ha rac te r is t ics . H oweve r , i ti sa ga in s ugges ted t ha t t he f ocus o f s uch a c t iv i t ies i n vo lved t he r ange o fn a tura l v a r iab i l i ty i nd es i rab le t r a i ts a l ready p resen t among c e r ta in c e rea l s pec ies , a nd t ha t p r io r t o t he e igh teen th c en tury A .D . , s e lec t ive p lan t b reed ing f o r t he e nhancemen t o f d es i rab le t r a i ts w as u n l ike ly i na l b u t af ew c e rea l s pec ies . F u r the r , i ti sa ppa ren tt ha t many s h i f ts i nt he f r equenc ies o f s pec i f ic morpho log ica l t r a i ts , s uch a s u p r igh t c u lms a nd n onsha t te r ing s p ikes , we re p robab ly t he r esu l t o f n a tura l o r u n in ten t iona l s e lec t ion a s ar esu l t o f g row th i n d ense c u l t iva ted s tands , h a rves t ing me thods a nd t h resh ing p rac t ices , r a the rt han t he r esu l ts o f i n ten t iona l b reed ing ( Boh rer 1 972 ; E vans 1 976 :110 ; F e ldman 1 976 :126 ; H a r lan e t a l . 1 973 :313 -321, 1 976 :6 -8 ; H i l lman 1 978 :167 ; K is lev 1 984 :63 ; S immonds 1 976 ; Z ohary 1 969 :60 ; Z oha ry 1 973 :622 ) . E ven h e re , t he p resumed s i gn i f icance o f n onsha t te r ing s p ikes among d omes t ica ted w hea ts s eems o ve r ly s t ressed , s i nce H a r lan f ound t ha t w i ld w hea ts when a lmos t r i pe " can b e h a rves ted w i th a s i ck le w i th o n ly m ino r l o sses " ( 1967 :198 ) . F ur the r , d omes t ica ted e inko rn ( T r i t icum monococcum ) a nd emme r ( T r i t icum d i coccum ) a re s a id t oh ave ar ach is w h ich i so n ly s l igh t ly l e ss b r i t t le t han t he i rw i ld c oun te rpa r ts ( H . J a rman 1 972 :17 18 ; P e rc iva l 1 92 1 :17 1 , 1 89 , 3 64 ) . A s l a te a s t he f i r s t c en tury A .D . , t he R oman a g ronom is t C o lume l la wa rned t ha t b a r ley s hou ld b e r ap id ly h a rves ted d ue t o i t s t endency t oq u ick ly s ha t te r w hen r i pe ( 1 .9 .15 ) . E ven i n t he m id -n ine teen th c en tury , S tephens ( 1866 :329 ) s t ressed t he n ecess i ty o f n o t a l low ing whea t , b a r ley o r o a ts t o b ecome t oo r i pe b e fo re h arves t ing l e s t e ach " cas t i t s s eed w i th t he s l igh tes t w ind . " H e t he re fo re s ugges ted t ha t " I n e ve ry c ase , i ti s much b e t te r t o r eap t he c rop b e fo re i ti s r i pe , t han t o a l low i tt o s tand u n t i l t oo r i pe " ( S tephens 1 866 :329 ) . Wh i le n onsha t te r ing s p ikes m igh t a low f o r al o nge rp e r iod d u r ing w h ich c e rea ls c ou ld b e h a rves ted , i t i sn o t e xpec ted t ha tt h is t r a i tw ou ld a f fec tt he e ne rge t ics o fc e rea l e xp lo i ta t ion o rc u l t iva t ion .

T he G ram ineae a nd C e rea lS pec ies S ub jec t t oE ar ly Husband ry A l l t r ue c erea ls b e long t o t he g rass f am i ly ( Gram ineae ) w i th in t he s ubc lass Monoco ty ledones ( wh ich i n c ludes p a lms , o rch ids , l ies , a nd g rasses ) o ft he a dvanced c l ass o f p l an ts k nown a s A ng iospe rmae , p l an ts w h ich a re p o l l ina ted t h rough f l o ra l s t ruc tures a nd b ea r s eeds e nc losed i na n o va ry o r p od . D ur ing t he c ou rse o f t he i r p hy logeny , g rasses e vo lved i n s evera l d is t inc t h ab i ta ts wo r ld -w ide , r esu l t ing i n d i f fe ren t t r a i ts a nd d i s t r ibu t ion p a t te rns b e tween g rass s ub fam i l ies a nd t he s pec ies t hey c on ta in ( Gou ld a nd S haw 1 983 :105 ) . T he d i s t r ibu t ion p a t terns o ft hese s ub fam i l ies a nd t he g ene ra l t r a i ts a ssoc ia ted w i th t he i rc ons t i tuen t s pec ies a re m in ima l ly r ecogn ized a s p l ay ing s i gn i f ican t r o les i nt he s pa t ia l e vo lu t ion o fe a r ly f ood p roduc t ion ( e .g . , H a r lan 1 977 ; H a r lan a nd Z oha ry 1 966 ; S tem le r 1 980 ; Z oha ry 1 969 ) . Ab as ic u nde rs tand ing o ft he g ene ra l p hy logeny a nd e co logy o fc e rea l g rasses i st he re fo re wa r ran ted b e fo re t he n a tu re o fe a r ly man /p lan t r e la t ionsh ips i sa dd ressed .

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I n c ur ren t p hy logene t ic g rass t axonomy , t he G ram ineae f am i ly i s d iv ided i n to s i x s ub fam i l ies a nd 2 5 t r ibes c on ta in ing a pp rox ima te ly 6 00 g ene ra a nd a round 7 ,500 s pec ies ( Ba rna rd 1 964 ; Gou ld a nd S haw 1 983 :2 ) . O f t h is t o ta l n umbe r o f s pec ies , o n ly 2 9 ( o r r o ugh ly 0 .4% ) r ep resen t k nown o r s uspec ted c e rea ls s ub jec t t o e a r ly h uman h usband ry f o r f ood p roduc t ion . T ab le 1 s u mma r izes t he p hy logene t ic r e la t ionsh ips b e tween t hese c e rea ls a cco rd ing t ot he g eog raph ic r eg ions o ft he i r a ppa ren t e a r ly h usband ry . S pec ies i n t he s ub fam i ly F es tuco ideae ( = P oo ideae ) e vo lved i n t empe ra te l a t i tudes e xh ib i t ing c oo l o rc o ld s easons . H ence , s ho r t d ays a nd l ow t empe ra tures f avo r r oo t g row th a nd a bundan t t i te r ing i nt he f es tuco id g rasses , w h i le s ubsequen t l o ng d ays a nd mode ra te ly h i gh t empe ra tures f avo r l e a f a nd s t em g row th o n e s tab l ished t i le rs ( Ba rna rd 1 964 :116 ; G ou ld a nd S haw 1 983 : 105 , 1 09 , 14 ) . F l owe r p roduc t ion among t he f es tuco ids i ss pec i f ica l ly p romo ted b y i n c reas ing d ay l e ng th ( Ba rna rd 1 964 :136 ; J ones 1 983 :179 -180 ; V ince -P rue 1 975 : 13 , 1 4 , 2 62 -263 ) . A s e xamp les o f c o ld t o le rance among t he F es tuceae , b a r ley i s c apab le o f g e rm ina t ing a tt empe ra tu res a s l ow a s3 ° C( Robe r ts 1 969 : 172 ) , a nd w hea t a t3 .0 -4 .5° C ( Lev i t t 1 969 :397 ) . A s t he h ab i ta ts o f many f e s tuco id g rasses b ecomes i n c reas ing ly h o t a nd d ry , t he g ra in ma tu res a nd t he i r r ep roduc t ive c yc le i sc omp le ted . A l though t he F es tuceae e vo lved i n h ab i ta ts l y ing n or th a nd s ou th o f t he t r op ics , s ome s pec ies h ave b ecome a dap ted t o h i gh a l t i tude h ab i ta ts i nt he moun ta inous r eg ions o ft r op ica l l a t i tudes ( Gou ld a nd S haw 1 983 : 114 ) . S pec ies i nt he s ub fam i ly P an ico ideae p r ima r i ly e vo lved i nt r op ica l a nd s ub t rop ica l l a t i tudes . H ence , p an ico id g rasses , u n l ike t he F es tuco ideae , g ene ra l ly r equ i re h i gh t empera tu res f o r v ege ta t ive g row th ( Ba rna rd 1 964 :118 ; S tem le r 1 980 :508 ) , w i th f l owe r f o rma t ion p romo ted b y d ec reas ing d ay l e ng th ( Ba rna rd 1 964 :136 ; H a r lan e t a l . 1 973 :316 ; J ones 1 983 :179 180 ; S tem le r 1 980 :508 ; V ince -Prue 1 975 :25 -26 ) . A s e xamp les o f c o ld i n to le rance among t he P an ico ideae , s o rghum a nd p ea r l m i l le t r equ i re af r os t f r ee p e r iod f o r g row th a nd t empe ra tures a bove 1 0 ° Cf o rg e rm ina t ion a nd eme rgence ( Bagna l l 1 979 :67 ; N o rman e ta l . 1 984 :34 ) . Membe rs o f t he P an ico ideae c ons t i tu te t he ma jo r i ty o ft he g rasses f ound i nt he t r op ics a nd s ub t rop ics , a l though s eve ra l p an ico id s pec ies a re a l so f ound i nt he E as te rn U .S . ( Gou ld a nd S haw 1 983 :117 -118 ) . H oweve r , many p an ico id s pec ies a re u nab le t o r ep roduce a tt hese h i ghe r l a t i tudes , s i nce t he s ho r te r d ays t ha t i n t ia te t he i r f l owe r ing a re g enera l ly f o l lowed i n t empe ra te z ones b y k i l l ing f r os t ( S tem le r 1 980 :508 ) . T h is i sp ar t icu la r ly t r ue f o r t he p an ico id c u l t iva rs ( Ha r lan e t a l . 1 973 :316 ) , a nd h as r ecen t ly b een s ugges ted a s s i gn i f ican t i n u nde rs tand ing d emog raph ic a nd s ubs is tence v a r iab i l i ty among c o rn a g r icu l tura l is ts i nt he p reh is tor ic S ou thwes t o ft he U .S . ( Pe te rsen 1 985 ) . T he w a rm s eason E u ropean m i l le ts ( Se ta r ia i t a l ica a nd P an icum m i l iaceum ) a re n o tab le e xcep t ions t ot hese g ene ra l g eog raph ic t r ends i n t he O ld Wo r ld , a s t hey h ave b ecome a dap ted t ot he b e l t o fs umme r r a in fa l l a c ross t empe ra te E u ras ia ( Ha r lan 1 977 :379 -380 ) . L i ke t he P an ico ideae , s pec ies i nt he s ub fam i ly O ryzo ideae p r ima r i ly e vo lved i nt r op ica l a nd s ub t rop ica l e nv i ronmen ts , a nd a re a l so c o ld s ens i t ive d u r ing t he i r g erm ina t ion , g row th , a nd s eed p roduc t ion p e r iods ( Jones 1 983 :199 ; N o rman e t a l . 1 984 :87 , 9 0 ; Z so ldos a nd K a rva ly 1 979 : 124 ) . A s ian r i ce ( O ryza s a t iva ) , f o re xamp le , w i l l n o tg e rm ina te a tt empera tu res b e low 1 6 ° C( No rman e ta l . 1 984 :87 ; R ober ts 1 969 :167 ) . A s among t he P an ico ideae , f l owe rf o rma t ion i n t he O ryzo ideae i s a l so p romo ted b y d ec reas ing d ay l e ng th ( Ba rna rd 1 964 : 136 ; J ones 1 983 :179 -180 ; V ince -P rue 1 975 :12 , 4 1) . G rasses o f t h is s ub fam i ly a re c ha rac te r is t ica l ly a dap ted t o mo is t , ma rshy , o r a qua t ic h ab i ta ts , a nd a re w ide ly d i s t r ibu ted i nt he t r op ica l a nd s ub t rop ica l r eg ions o ft he wo r ld ( Gou ld a nd S haw 1 983 :105 , 1 27 -128 ) . S ome o ryzo id s pec ies , h oweve r , h ave b ecome a dap ted t o h ab i ta ts i nt empera te N or th Ame r ica , i n c lud ing A nnua l W i ld r ice ( Z izan ia a qua t ica ) a nd N o r the rn W i ld r ice ( Z izan ia p a lus t r is ) , t he l a t te r r epor ted f r om N ew E ng land t h rough t he G rea t L ake s t a tes a nd n or th i n to c en t ra l C anada ( Gou ld a nd S haw 1 983 :128 , 3 25 ) . H oweve r , w h i le m embe rs o ft he g enus Z izan ia h ave l o ng b een e xp lo i ted a s af ood r esource b y N a t ive Ame r icans , r i ce a g r icu l ture s pec i f ica l ly d eve loped f r om t he e xp lo i ta t ion o fs pec ies i nt he g enus O ryza , a l membe rs o f wh ich e vo lved i n t he mo is t t r op ica l a nd s ub t rop ica l h ab i ta ts o f t he O ld a nd N ew Wo r lds ( Gou ld a nd S haw 1 983 :321) . F u r the r , o ft he 1 5 t o 2 0 s pec ies w i th in t he g enus O ryza , o n ly A f r ican r i ce ( O ryza g labe r r ima ) a nd A s ian R ice ( O ryza s a t iva ) f o rmed t he b as is f o rc e rea l a g r icu t tu res , a nd o n ly t he l a t te rb ecame t he d om inan t c e rea lo f ama jo ra g r icu l tu ra lc omp lex .

4 4

T ab le 1

P hy logene t ic R e la t ionsh ips o fC e rea ls A cco rd ing t ot he Geog raph ic R eg ions o fT he i r I n i t ia l H usband ry

TR IBES I NS UBFAM ILY

GRASS S UBFAM ILY

#

CULT IVAR S PEC IES I N TR IBE

1 t ha t h ave c u l t iva rs

b y R eg ion o fI n i t ia l H usband ry

A rund ino ideae

3

N one

0

N one

B ambuso ideae

2

N one

0

N one

C h lor ido ideae

8

E rag ros teae

2

A FR ICA :l e f t ( E rag ros t i s t e f f ) , f i nge r m i l le t

O ryzo ideae

1

O ryzeae

2

A FR ICA : A f r ican r i ce ( O ryza g labbe r ima )

A nd ropo g oneae

P an ico ideae

A veneae F es tuco ideae ( = P oo ideae )

TOTAL

2 5

6

AS IA :J ob 's t ea rs ( Co ix l a chrymajob t ) MESOAMER ICA : ma ize ( Zea mays )

1 0

A FR ICA : p ea r l m i l le t ( Penn ise tum ame r icanum ) , G u inea m i l le t ( B rach ia r ia d e f lexa ) , c ommon f on io ( D ig i ta r ia e x i l is ) , b l ack f on io ( D ig i tar ia i b u rua ) AS IA : s l ende rm i l le t ( Pan icum m i l iare ) , J apanese m i l le t ( Ecenoch loa f r umen tacea ) , k odo m i l le t ( Paspa lum s c rob icu la tum ) E URAS IA :f o x ta i l m i l le t ( Se ta r ia i t a l ica ) , p an ic m i l le t ( Pan icum m i l iaceum ) MESOAMER ICA : p an ic g rass ( Pan icum s onorum )

6

NEAR E AST : s and o a t ( Avena s t r igosa ) , c e rea l o a ts ( A . s a t iva , A . b yzan t ina ) , E th iop ian o a ts ( A . a byss in ica , A . v av i lov iana ) E ASTERN U .S . : mayg rass o rc ana ryg rass ( Pha lar i sc a ro l in iana )

6

NEAR E AST : e i nko rn w hea t ( T r i t icum monococcum ) , emme rw hea t ( T . d i coccum ) , t imopheev i ( T . t imophee d , b read whea t ( T . Xa es t ivum ) , b a r ley ( hordeum v u lga re ) , r y e ( Seca le c e rea le )

9

T r i t iceae

AS IA :A s ian r i ce ( O ryza s a t iva ) A FR ICA : s o rghum ( Sorghum b ico lo r )

3

2 P an iceae

( E leus ine c oracana )

2 9

4 5

S pec ies i nt he s ub fam i ly C h lo r ideae s eem ing ly e vo lved i ns pa t ia l ly d i ve rse , w a rm , s em ia r id e nv i ronmen ts w i th h i gh t empe ra tu res , h i gh l i gh t i n tens i t ies , a nd l ow s o i l mo is ture , a nd a re c ons ide red t he mos t s uccess fu l o f t he g rass s ub fam i l ies i ns uch h ab i ta ts ( Gou ld a nd S haw 1 983 :105 ) . I n s ome r eg ions o f t he wa rm , s em ia r id S ou thwes t o f t he U .S . , membe rs o f t h is s ub fam i ly c omp r ise mo re t han 5 0% o f t he g rass b i omass ( Gou ld a nd S haw 1 983 :120 ) . U n for tuna te ly , d a ta o n t empe ra tu re r esponse a nd f l owe r ing among c h lo r ido id g rasses h ave n o t b een l o ca ted .

AF ocus On A nnua ls a s Opposed t oP e renn ia ls Wh i le t he s ub fam i l ies C h lo r ido ideae , O ryzo ideae , P an ico ideae , a nd F es tuco ideae c on ta in b o th p e renn ia l a nd a nnua l s pec ies , a l o ft he k nown o rs uspec ted c e rea l c u l t iva rs e xh ib i t a nnua l l i f e h i s to ry h ab i ts . F ur the r , o n ly A s ian r i ce ( O ryza s a t iva ) a nd r y e ( Seca le c erea le ) m igh t h ave b een s e lec ted f r om i n i t ia l ly p e renn ia l p opu la t ions , a l though t he i r d er iva t ion f r om a l ready e s tab l ished a nnua l s pec ies s eems mo re p robab le ( Ha r lan e ta l . 1 973 :3 17 ) . P e renn ia ls a re e ssen t ia l ly p l an ts w i th l i f e c yc les t ha t s pan mo re t han a s i ng le g row ing s eason . Wh i le s ome p e renn ia ls d i e b ack t ot he g round d ur ing c o ld s easons a nd s ubsequen t ly r ev ive f r om t he i r r oo ts , mos t s imp ly a dd n ew g row th e ach y ea r w i th l i t le l o ss o rd es t ruc t ion o f a bove -g round p a r ts . I nd i s t inc t c on t ras t , t he l i f e c yc les o fa nnua l p l an ts a re c omp le ted w i th in a s i ng le g row ing s eason . P e renn ia l i ty a nd a nnua l i ty t hus r ep resen t d i s t inc t l i f e h i s to ry v a r ia t ions i np lan t r ep roduc t ive s t ra teg ies . T ra i ts a ssoc ia ted w i th p e renn ia l a nd a nnua l s t ra teg ies i n c lude t he d iageo t rop ism o f n ew t i le rs i np e renn ia l g rasses , w h ich c an r oo t a nd b ecome i n dependen t mo re r ead i ly t han t he u p r igh t t i le rs o f many a nnua ls ( Ba rna rd 1 964 :145 ) , a s w e l l a s v a r ia t ions i ns eed s e t a nd s i ze . W i ld p e renn ia l g rasses a re n o to r ious ly p oo ro re r ra t ic s eed s e t te rs , a nd af u l l s e t o fs eed i ss a id t o b e u ncommon ( Ha r lan e ta l . 1 973 :317 ) . I nc on t ras t , a nnua l g rasses d epend e n t i re ly u pon r ep roduc t ion t h rough s eeds ( Bake r 1 959 :181) , a nd t hus s e t s eed much b e t ter t han p e renn ia ls , g ene ra l ly p roduc ing l a rge rq uan t i t ies o fs eed w i th in a mo re r es t r ic ted p e r iod o ft ime ( Cohen 1 97 1 ; H a r lan e ta l . 1 973 :317 ; L l oyd 1 980 :85 -86 ; P a l t r idge a nd D enho lm 1 974 ; Why te 1 977 :214 ) . A lso , t he s eeds o fa nnua l g rasses a re o f ten l a rge rt han t hose o fp e renn ia ls (Why te 1 977 :2 14 ) , a l though s eed s i ze v a r ia t ions a re o f ten p ronounced b e tween r aces o fw i ld s pec ies , a s n o ted among p opu la t ions o fw i ld b a r ley a nd emme r ( Ha r lan a nd Z ohary 1 966 ; N evo e t a l . 1 979 ; Z ohary 1 969 ) . I nt e rms o fh arves t ing l a bo r a nd s eed y i e ld , H a r lan h as f ound t ha t d ue t o t he i nde te rm ina te g row th o f p e renn ia ls , a t temp ts t o o b ta in a n o p t imum s eed y i e ld d u r ing h a rves t ing e xpe r imen ts o f ten i n vo lved a low ing t he e a r l ies t s eed t o s ha t te r b e fo re s t a r t ing t o h a rves t , a nd f u r the r was ted t he l a te r ma tu r ing s eed ( Ha r lan e ta l . 1 973 :314 ) . I n ap rox ima te s ense , t he e a r ly h uman f ocus o n a nnua l g rass s pec ies f o r e xp lo i ta t ion a s c e rea l c rops was p robab ly r e la ted t ot he i r mo re f avo rab le s eed s i ze , s e t t ing , a nd p roduc t ion t r a i ts r e la t ive t o p e renn ia l g rasses . Mo re s pec i f ica l ly , i ts eems r easonab le t o e xpec t t ha t a g rea te r y i e ld o f s eed p e r u n i t t ime s pen t h a rves t ing c ou ld b e o b ta ined f r om c e r ta in a nnua l s pec ies a s o pposed t op e renn ia ls . D epend ing u pon t he c a lo r ic v a lue o ft he s eed y i e ld a nd t he p rocess ing t ime i tr equ i res , i ti sg ene ra l ly e xpec ted t ha t t he e xp lo i ta t ion o f s pec i f ic a nnua l s pec ies wou ld o f ten b e e ne rge t ica l ly mo re e f f ic ien t t han t he e xp lo i ta t ion o f p erenn ia l a l te rna t ives . A s d ocumen ted i nC hap ter 6 ,s ome s uppor t f o rt h is g ene ra l e xpec ta t ion i sf ound i n ac ompa r ison o ft he e xpe r imen ta l h a rves t ing d a ta o fH a r lan ( 1967 ) f o rw i ld e i nko rn whea t i n T urkey a nd S imms ' ( 1984 , 1 987 ) e xper imen ta l h a rves t ing d a ta f o r v a r ious p e renn ia l s eed r esou rces i nt he G rea t B as in .

T he E vo lu t ion o fA nnua lS pec ies f r om P erenn ia l P rogen i tors T he ma jo r i ty o fa l monoco ty ledons a re p e renn ia ls , a nd p erenn ia l i ty i sw ide ly a ccep ted a s t he p r im i t ive c ond i t ion i n g rasses ( e .g . , B a rna rd 1 964 :144 ; S tebb ins 1 957 :340 ; Why te 1 977 :213 ) . A nnua l s pec ies s eem ing ly a rose , a nd c on t inue t oa r ise , f r om p e renn ia l p rogen i to rs . S ign i f ican t ly , t he b reed ing s ys tem o f mos t p e renn ia l g rasses i n vo lves c ross -po l l ina t ion , wh i le many o f t he a nnua l g rasses a re p redom inan t ly s e l f -po l l ina to rs ( Bake r 1 959 :181; B a rna rd 1 964 :141-142 ; F ryxe l l 1 957 ; S tebb ins 1 957 ) . Wh i le s e l f incompa t ib le a nnua ls ( even d ioec ius 4 6

o nes ) a re k nown , t hey a re p ropor t iona te ly r a re ( Bake r 1 959 : 18 1) . S tebb ins ( 1957 ) i np ar t icu la r h as d rawn a t ten t ion t o a ppa ren t t r ends f r om s e l f incompa t ib le p erenn ia ls t o s e l f -po l l ina ted a nnua ls i n s eve ra l g rass g ene ra , s ugges t ing t ha t t he s e lec t ive f ac to rs r espons ib le f o r t he e vo lu t ion o f a nnua l s pec ies f r om p e renn ia l p rogen i to rs o f ten f avo red t he c oevo lu t ion o f as e l f f e r t i l iz ing t r a i t . Wh i le c omp lex p hys io log ica l , b i ochem ica l , o r c y to log ica l f ac to rs may b e i n vo lved i nt he e vo lu t ion o f a nnua l s pec ies , i ti sg ene ra l ly p roposed t ha t n a tu ra l s e lec t ion wou ld f avo r t he e vo lu t ion o f a nnua l s pec ies f r om p e renn ia l p rogen i to rs when p l an t g row th i sl i m i ted b y t ime o r b y l i m i t ing r esources ( e .g . , C ohen 1 97 1; P a l t r idge a nd D enho lm 1 974 ; Why te 1 977 ) . I n p a r t icu la r , i th as b een n o ted t ha t a nnua ls mos tf r equen t ly o ccu ri nt empo ra r i ly a va i lab le h ab i ta ts ( Baker 1 959 :181) . N o t ing t ha t h ighe r f r equenc ies o f a nnua ls t o p e renn ia ls a re s eem ing ly c o r re la ted w i th e nv ironmen ts whe re s easona l a r id i ty i s c omb ined w i th r e la t ive ly h i gh t empera tures ( app rox ima te ly 2 5 -30 0 ) , Why te ( 1977 :214 -215 ) h as s ugges ted t ha t t he c hange f r om a p e renn ia l t o a n a nnua l h ab i t i n vo lves s ome c omb ina t ion o f g rea t o r i n c reased i n tens i ty o f d es icca t ion a nd a w ide d i u rna l a nd s easona l r ange b e tween h i gh a nd l ow t empe ra tu res . I n e f fec t , a nnua l i ty i ss een a s a mechan ism f o r e scap ing s easona l ly u n favo rab le e nv i ronmen ts , w i th a nnua ls b e ing a dap ted t ot he a ch ievemen t o f mass r ep resen ta t ion a nd s eed p roduc t ion w h i ls tf avo rab le c ond i t ions i nt he i rh ab i ta t l a s t .

A nnua ls a nd P redom inan tS e l f -Po l l ina t ion Wh i le t empo ra l ly a va i lab le h ab i ta ts f avo r s hor te r g ene ra t ion t imes o f ten l ead ing t o a nnua l i ty , p i onee r a nnua ls i n i t ia l ly c o lon iz ing s uch h ab i ta ts w ou ld b e i n ap os i t ion whe re t he o ppor tun i t ies f o r c ross fe r t i l iza t ion we re r a re . U nde r t hese c ond i t ions , i ti sg ene ra l ly a ccep ted t ha t ah igh s e lec t ive a dvan tage wou ld a cc rue t o c o lon is ts c apab le o f s e l f -po l l ina t ion ( A l la rd 1 975 :115 ; B ake r 1 959 :181; C ha rnov 1 982 :234 -235 ; F ranke l a nd Ga lun 1 977 :51; F ryxe l l 1 957 :156 ; J a in 1 976 :485 ; L Loyd 1 979 :76 ; b u t s ee J a in 1 976 :473 -475 ; L l oyd 1 980 :80 , 8 4 8 7 ) , e ven a tt he p o ten t ia l e xpense o f r educed p opu la t ion v a r iab i l i ty d ue t o i n b reed ing ( Baker 1 959 :187 -188 ) . O f t he 2 9 k nown o r s uspec ted c e rea l c u l t iva rs men t ioned i nt h is t ex t , o n ly ma ize ( Zea mays ) , p ea r l m i l le t ( Penn ise tum amer icanu m ) , a nd J ob 's t ea rs ( Co ix l achryma job i ) a re p redom inan t c ross -po l l ina to rs , wh i le s o rghum ( Sorghum b ico lo r ) a nd r y e ( Seca le c erea le ) a re p a r t ia l ly s e l f - a nd p a r t ia l ly c ross fer t i l ized . A l l o ft he r es t a re p redom inan t s e l f -po l l ina to rs ( see F ryxe l l 1 957 :177 -182 ) . F o r e xamp le , t he r a te o f n a tu ra l o u tc ross ing i nc u l t iva ted b a r ley i s a pp rox ima te ly 1 -2 % ( A l la rd e ta l . 1 968 :117 ; J a in 1 976 :479 ) , i nf i nge r m i l le t a round 1% ( F ryxe l l 1 957 : 179 ; P u rseg love 1 976 ) , i nr i ce a pp rox ima te ly 0 .4 1 .07 % ( Rober ts e ta t . 1 96 1 :1085 ) , a nd i s max ima l ly 6 a nd 1 0% r e spec t ive ly i n whea ts a nd o a ts ( F ranke l a nd Ga lun 1 977 :18 ) . B y c on t ras t , p ea r l m i l le t i sh i gh ly s e l f incompa t ib le , w h i le ma ize e xh ib i ts 9 5% n a tu ra l o u tc ross ing , s o rghum 1 8 -76 % o u tc ross ing , a nd r y e a pp rox ima te ly 5 0% o u tc ross ing ( F ryxe l l 1 957 :18 1-182 ) . A c tua l ly , p redom inan t a s o pposed t o o b l iga tory s e l f fer t i l iza t ion i n p lan ts d oes n o t n ecessa r i ly r esu l t i ng ene t ica l ly u n i fo rm p opu la t ions ( A l la rd 1 965 ; A l la rd e ta l . 1 968 ; N evo e ta t . 1 979 ) . W i th in p opu la t ions o f p redom inan t ly s e l f -po l l ina t ing g rasses , i n d iv idua l p lan ts o f ten d i f fer f r om o ne a no the r w i th r espec t t o s i ng le - a nd mu l t ip le -un i t p o lymorph isms a nd a l so f o r c on t inuous ly v a ry ing c ha rac te rs . H oweve r , d ue t op redom inan t s e l f -po l l ina t ion , s uch v a r ia t ions a re p r ima r i ly c a r r ied b y d i f fe ren t h omozygo tes w i th in t ha t p opu la t ion ( A l la rd 1 975 :124 ; B rown e t a l . 1 978 :54 -57 ; N evo e ta l . 1 979 :832 ) . E ven w i th l ow f r equenc ies o f o u tc ross ing b e tween v a r ious c oex is t ing g eno types , n ew v a r iab i l i ty a r ises i ne ach g enera t ion t h rough r ecomb ina t ion a nd s eg rega t ion . T he c on t inued e vo lu t ion o f t he s ys tem s eem ing ly i n vo lves t he s l ow s o r t ing o u t a nd i n co rpo ra t ion o f t he o ccas iona l s upe r io r g eno type p roduced b y c on t inued s egrega t ion a nd r ecomb ina t ion t h rough b o th s tab i l iz ing a nd d i rec t iona l s e lec t ion ( A l la rd 1 965 ; A l la rd e ta l . 1 968 ) . N o t s urp r is ing ly , g ene t ic d i f fe ren t ia t ion b e tween p opu la t ions o ft he s ame i n b reed ing p l an t s pec ies i s ma rked , f r equen t ly e xh ib i t ing c l ina l v a r ia t ion i na ssoc ia t ion w i th p rog ress ive c hanges i nr a in fa l l , t empera ture , e l eva t ion , s o i lt ypes , a nd o the re co log ica l f a c to rs ( A l la rd e ta l . 1 968 :96 4 7

9 7 , 1 21) . F u r the r , s uper imposed o n s uch b road g eog raph ica l v a r ia t ions i s a" mosa ic " p a t te rn o f v a r ia t ion r e f lec t ing a dap ta t ions t ol o ca l e nv i ronmen ts . F o re xamp le , Z oha ry ( 1969 :55 ) n o ted t he p resence o f morpho log ica l ly d i s t inc t e cogeog raph ic r aces o f w i ld b a r ley ( Hordeu m s pon taneum ) i n I s rae l , wh i le N evo e t a l . ( 1979 ) d emons t ra ted t ha t many p a t te rns o f s uch v a r ia t ion i nt he g ene p oo l o fw i ld b a r ley a re p red ic tab le e co log ica l ly , e spec ia l ly b yc omb ina t ions o ft empe ra ture , r a in fa l l , h um id i ty , a nd s o i l t y pes . S im i la r e cogeog raph ic v a r ia t ions h ave b een n o ted among w i ld o a ts ( Avena b arba ta ) i nb o th I s rae l ( Nevo e t a l . 1 979 :831) a nd C a l i fo rn ia ( A l la rd 1 975 :117 122 ) , a nd among w i ld e i nko rn whea t ( T r i t icum b oeo t icum ) a nd t he g oa tgrass A eg i lops s qua rosa ( Ha r lan a nd Z oha ry 1 966 ;Z oha ry 1 969 ; Z oha ry e ta l . 1 969 ) . O ve ra l l , g ene t ic v a r ia t ions i n ap redom inan t ly s e l f -po l l ina t ing g rass s pec ies i n c lude c l ina l , r eg iona l , a nd l o ca l p a t te rns wh ich s ome t imes d i sp lay s ha rp g eog raph ic d i f feren t ia t ion o ve r s hor t d is tances . T hus , p redom inan t ( bu t n o t o b l iga tory ) s e l f -po l l ina t ion a l lows f o r t he p e rpe tua t ion o ft he p resen t ly b es t a dap ted g eno types f o r v a r ious s pec i f ic m ic roenv i ronmen ts w i th o ccas iona l o u tc ross ing p rov id ing t he f l ex ib i l i ty r equ i red f o r t he d eve lopmen t o f ad i ve rs i ty o fg eno types o f i n c reas ing f i t ness i ns pec i f ic n i ches ( A l la rd 1 965 :72 -74 ; A l la rd e ta l . 1 968 : 12 1 , 1 24 -125 ; S tebb ins 1 957 :344 -346 ) . H oweve r , r ega rd less o f t he i r b reed ing s t ra tegy , p heno typ ic v a r iab i l i ty a nd p l as t ic i ty a re q u i te ma rked among a l a nnua lg rass s pec ies , p a r t icu la r ly i nr e spec tt ot he i rp roduc t ive t r a i ts .

P heno typ ic P las t ic i ty a nd P roduc t ive P o ten t ia l : T he E v idence f o r Whea t A s n o ted e a r l ie r , a nnua l g rasses h ave e vo lved t oa ch ieve mass r ep resen ta t ion a nd s eed p roduc t ion i nt empo ra l ly f avo rab le e nv i ronmen ts ( Cohen 1 971 ; P a t t r idge a nd D enho lm 1 974 ) . As i gn i f ican t a spec t o ft h is a dap t ive s t ra tegy i se no rmous p heno typ ic p las t ic i ty i nt he i rp o ten t ia l f o rs eed p roduc t ion ( A l la rd 1 965 :76 ) . F o r e xamp le , i na n u n favo rab le s eason , w i ld o a ts w i l l g row a s s ma l l p l an ts p roduc ing a sf ew a so ne s eed e ach , b u t i n af a vo rab le s eason , w i l lg row a s much l a rge r p lan ts p roduc ing h und reds o f s eeds e ach ( A l la rd 1 965 :76 ) . S im i la r ly , w i ld whea ts o ccupy ing p oo r , t h in , a nd r ocky s o i ls i nt he i r n a tu ra l h ab i ta ts r espond w i th i n c reased s eed p roduc t ion when i n t roduced t o r i che r , mo re f avo rab le h ab i ta ts ( Fe ldman 1 976 :124 ) , wh i le t i le r ing i n whea t , b a r ley , r ye , a nd o a ts i ss t rong ly i n f luenced b y s o i l f e r t i l i ty , a va i lab le mo is tu re , t empe ra tu re , a nd d ay l e ng th ( Ha r lan e t a l . 1 973 :3 16 ) . E ven t he y i e ld f r om t he s ame s eed o n d i fe ren t p a r ts o ft he s ame f i e ld f r equen t ly v a r ies g rea t ly , w i th s uch v a r ia t ions b e ing e ven mo re p ronounced b e tween r eg ions ( Co t t re l l 1 955 :29 ) . T he e f fec ts o fs o i l n u t r ien ts o nt he p heno typ ic p l as t ic i ty a ssoc ia ted w i th c e rea l p roduc t iv i ty i sr ead i ly a ppa ren t i nr e la t ion t ow hea t . When g rown o n s o i l d e f ic ien t o f n i t rogen , w hea ts y i e ld p oo r ly i nb o th q uan t i ty a nd q ua l i ty o fg ra in ( S tephens 1 866 :455 ) . H owever , a s e xemp l i f ied b y e xpe r imen ts c onduc ted o ve r a1 0 y ea r p e r iod a t R o thams ted , E ng land d u r ing t he m id n i ne teen th c en tu ry , t he mean y i e ld o f whea t p e r a c re c ou ld b e d oub led t h rough t he a pp l ica t ion o f n i t rogen r ich P eruv ian g uano i nt he t o ta l a bsence o f s e lec t ive b reed ing ( Ca i rd 1 853 :460 4 62 ) . G iven t he e normous p heno typ ic p l as t ic i ty e xh ib i ted b y a nnua l g rasses , i ti sn o t s urp r is ing t of i nd t ha t n a tu ra l s t ands o fw i ld w hea t c an o ccu r i nd ens i t ies a pp rox ima t ing t hose o f mode rn , c u l t iva ted , n onhyb r id whea ts ( Ha r lan a nd Z oha ry 1 966 :1078 ) . F o r e xamp le , e s t ima tes o f t he p roduc t iv i ty o fw i ld s t ands o fm ixed whea t a nd b a r ley i nt he e as te rn Ga l i lee r ange f r om 5 0 t o8 0 k g p e rd unum o fl a nd ( = 1 , 000 m2 ,0 . 1 h ec ta re , o r0 .247 a c res ) i nr a iny y ea rs ( Zoha r y 1 969 :56 ) . T h is r ange o fw i ld y i e lds i sq u i te c ompa rab le t ot he e xpec ted r ange o f whea t y i e lds r epo r ted f o r s pa t ia l ly a nd t empo ra l ly d i ve rse d ry fa rm ing c ond i t ions . F o r e xamp le , u s ing C o lume l la 's d a ta o n t he s ow ing r a te f o r w hea t o n l and o f a verage q ua l i ty ( 1 .9 .1) a nd h i s d a ta o n e xpec ted whea t y i e ld t h roughou t mos t o f I t a ly ( 1 1 .3 .4 ) , aR oman f a rme ro ft he f i r s t c en tu ry A .D . s ow ing 5 mod i io fw hea t ( 1 mod ius = 6 .74 k g ) p e r i uge rum o fl a nd ( = 2 .518 d unums ) c ou ld e xpec t ay i e ld o f 2 0 mod i ip e r i u gerum , o r a pp rox ima te ly 5 3 .5 k g p e r d unum ( see D uncan -Jones 1 982 :370 -7 1 f o r u n i t c onve rs ions ) . S im i la r ly , p apy r i o ft he s even th c en tu ry A .D . r ecove red f r om t he B yzan t ine t own o f N essana i nt he N egev d ese r t o fs ou the rn I s rae l i n d ica te a n a ve rage l o ca l whea t y i e ld o f a round 5 0 k g p er d unum ( Maye rson 1 955 :55 , 1 960 :18 ) . 4 8

F o r med ieva l E urope i nt he f our teen th a nd f i teen th c en tu r ies , L enna rd ( 1922 :20 -25 ) c a lcu la ted ar ange o fa ve rage whea ty i e lds o fa pp rox ima te ly 5 0t o5 7k g p e rd unum o nt he b as is o fv a r ious mano r ia l a ccoun ts . B enne t t ( 1935 :2 1) f u r the rd e te rm ined t ha t a ve rage B r i t ish whea t y i e lds f r om A .D . 1 200 t h rough 1 450 amoun ted t oa pp rox ima te ly 5 4 t o6 1 k g p e rd unum , b ased u pon h i s a na lys is o ft he w hea ty i e ld d a ta p rev ious ly s umma r ized b y L enna rd ( 1922 ) , B eve r idge ( 1927 ) , a nd o the rs . H oweve r , t he mos t c onc ise a ccoun t ing o f med ieva l whea t y i e lds i sT i tow 's a na lys is ( 1972 ) o f n ume rous mano r ia l a ccoun ts i nt he W inches te r b i shop r ic b e tween L ondon a nd B r is to l d u r ing t he t h ir teen th a nd f our teen th c en tur ies . D u r ing t he 7 1 y ea rs f r om 1 232 t h rough 1 349 f o r wh ich T i tow was a b le t oe s tab l ish w hea t y i e lds o n a pp rox ima te ly 4 1 mano rs ( 1972 :83 -91) , t he mean a nnua l y i e ld was 5 2 .8 k g p e r d unum , w i th a r ange o f a nnua l means f r om a pp rox ima te ly 2 9 t h rough 7 2 k g p e r d unum . F i na l ly , e nqu i r ies o ft he R oya l S oc ie ty i n to a gr icu l tu re i nC o rnwa l l a nd D evon u nde rC ha r les Ie s tab l ished t ha tt he mean y i e ld o f whea t o n ma r l l a nds i nA .D . 1 668 amoun ted t o5 4 k g p e rd unum ( Lenna rd 1 932 :40 ) . R e tu rn ing t ot he N ea r E as t , s im i la r whea t y i e lds we re r epo r ted d u r ing t he B r i t ish manda te i nP a les t ine . I n ar epo r t c omp i led d u r ing t he l a te 1 920s b yt he Z ion is t A g r icu l tu ra l E xpe r imen ta l S ta t ion a tT e l A v iv , t he a ve rage w hea t p roduc t iv i ty o fA rab f a rms i nP a les t ine was g i ven a s 5 0 k g p er d unum ( i n S impson 1 930 :177 ) . I n ar epor t b y aB r i t ish c omm i t tee o n t he c ond i t ion o f a g r icu l tura l is ts i n P a les t ine d a ted J u ly 1 3 , 1 930 , t he mean d ec la red whea t y i e ld i n 1 04 " r ep resen ta t ive " A rab v i l lages was 4 8 k g p e rd unum , a l though " se lec ted e v idence " i n d ica ted a mean y i e ld o f5 7 k g p e rd unum a nd a n " of ic ia l e s t ima te " p l aced whea t p roduc t iv i ty a t6 7 k g p e r d unum ( S impson 1 930 :185 ) . F ina l ly , d a ta c omp i led d u r ing a s urvey o f 2 7 J ew ish a g r icu l tura l c o lon ies i nP a les t ine i n1 926 i n d ica tes amean whea t y i e ld t ha ty ea ro fa round 7 2 k g p e rd unum ( Adams 1 928 : T ab le 1 3 ) . N o t s u rp r is ing ly , b edou in f a rme rs i nt he c en t ra l N egev i nt he l a te 1 950s c ons ide red 5 0 t o 7 0 k g o f whea t p e r d unum a s b e ing t he r ange f o r " good " h a rves ts , w i th 3 0 t o 5 0 k g b e ing " sa t is fac to ry " ( Maye rson 1 960 :18 ) . T he mean y i e ld o f whea t i na l o fJ o rdan d u r ing t he t en y ea r p er iod f r om 1 957 t h rough 1 966 was a pp rox ima te ly 5 2 k g p e r d unum ( S te i t ieh a nd Smad i 1 974 :10 , T ab le 1 ) , wh i le t he mean y i e ld o nt he E as t B ank a l one d u r ing t he s i x y ea rp er iod f r om 1 967 t h rough 1 972 amoun ted t o a pp rox ima te ly 6 9 k g p e r d unum ( S te i t ieh a nd Smad i 1 974 :13 , T ab le 2 ) . I ft r ea ted a s as i ng le r eco rd , mean J o rdan ian whea t p roduc t iv i ty d u r ing t h is 1 6 y ea r p er iod was a pp rox ima te ly 5 8 k g p e rd unum , w i th ar ange o fa nnua l mean y i e lds f r om 1 7 t o 1 00 k g p e rd unum . Wh i le t hese p roduc t ion f i gures p r ima r i ly r e la te t o mean p roduc t iv i ty o n a ve rage l a nds u nde r a ve rage e nv i ronmen ta l c ond i t ions , much h i ghe rd ry fa rm ing whea t y i e lds we re e xpec ted o n r i che r s o i ls u nde r mo re f avo rab le c ond i t ions . T hus , C ice ro 's s ow ing a nd r e turn r a tes f o r whea t i nt he r i ch L eon t in i d i s t r ic t o f S ic i ly i nt he f i r s t c en tury A .D . ( I n V et . , 1 . 3 .112 ) i n d ica te y i e lds r ang ing f r om 1 07 t o 1 60 k g p e r d unum , w i th 1 28 k g p e r d unum b e ing a ve rage . V a r ro ( 1 .44 ) r epor ted t ha t whea t y i e lds i ns ome s ec t ions o fE t ru r ia w e re a s h i gh a s 2 00 k g p e rd unum . S im i la r ly , i n d iv idua l d r y fa rm ing whea ty i e lds a s h i gh a s 1 84 k g p e rd unum w e re r epo r ted i nt he W inches ter mano r ia l a ccoun ts o f t he t h i r teen th a nd f ou r teen th c en tur ies ( T i tow 1 972 :88 ) . I n t he e nqu i r ies o ft he R oya l S oc ie ty u n de rC ha r les I ,w hea t y i e lds i nD evon i n1 668 we re a s h i gh a s 1 34 t o 1 68 k g p e r d unum ( Lenna rd 1 932 :40 ) . I nt he 1 926 s urvey o f2 7 J ew ish a g r icu l tu ra l c o lon ies i nP a les t ine , whea t y i e lds a s h i gh a s 1 35 k g p er d unum w e re r eco rded ( Adams 1 928 : T ab le 1 3 ) , wh i le Z oha ry ( 1969 :56 ) n o ted a r ange o f whea t a nd b a r ley y i e lds f r om 5 0 t o 1 50 k g p er d unum f o r a rd a g r icu l tu re i nt he e as tern Ga l i lee . U nde r " excep t iona l " c ond i t ions , b edou in f a rme rs i nt he N egev e xpec ted awhea t y i e ld o fa round 1 00 k g p e rd unum ( Maye rson 1 960 :18 ) , wh i le d ry fa rm ing whea t y i e lds o f 1 46 k g p e rd unum a re e xpec ted i ng ood y ea rs i nt he u p land r eg ions o f mode rn J o rdan ( S te i t ieh a nd Smad i 1 974 :19 , T ab le 4 ) . T he h igher d ry fa rm ing whea t y i e lds r epor ted f o r r i che r s o i ls u nder f avorab le e nv i ronmen ta l c ond i t ions a pp rox ima te t hose a ch ieved u nde r i r iga t ion a g r icu l ture . F o r e xamp le , i n t he v i l lage o f A svan , T u rkey , H i l lman ( 1973 :226 -227 ) f ound t ha t p r ior t o t he i n t roduc t ion o fc hem ica lf er t i l ize rs a nd mode rn f a rm mach ine ry , l o ca l d ry fa rm ing whea t y i e lds a ve raged 6 3 k g p e r d unum , wh i le whea t y i e lds u nde r l ow in tens i ty i r iga t ion a ve raged 10 k g p e r d unum . I nt he J o rdan V a l ley , y i e lds o f n onhyb r id whea t u nde r i r iga t ion a re e xpec ted t o r ange f r om 12 t o 1 91 k g p e r d unum ( S te i t ieh a nd Smad i 1 974 :19 , T ab le 4 ) . Wh i le p r ima r i ly r e levan t t o b a r ley y i e lds ( wh ich a re g ene ra l ly s l igh t ly h i ghe rt han whea t y i e lds ) , J acobsen a nd A dams ( 1958 : 1252 ) f ound i nt he i r a na lys is o f a nc ien t t ex tua l d a ta f r om t he s i te o f G i rsu i n Mesopo tam ia t ha t l o ca l g ra in y i e lds o f2 53 .7 l i t e rs p e r d unum a round 2 ,400 B .C . h ad d ec l ined 4 9

t o 1 46 l i t e rs p e rd unum b y 2 , 100 B .C . , p r ima r i ly r e f lec t ing t he d e le te r ious e f fec ts o fs a l in iza t ion u nde r i r iga t ion c ond i t ions . I ft hese y i e lds a re a pp l ied t o whea t , ad ec l ine f r om 1 70 k g p er d unum t o9 8 k g p e rd unum o ve r a4 00 y ea r p e r iod i si n d ica ted , a ssum ing ac onve rs ion a t0 .67 k g p e r l i t e ro fw hea t ( K rame r 1 982 :39 ) . T he mos t p roduc t ive l a nd u nde r i r iga t ion i nt he N ea r E as t a ppea rs t o b e t ha t o ft he N i le , whe re Amma r 's ( 1954 :253 ) d a ta o n t he v i l lage o f S i lwa i n d ica te a ve rage whea t y i e lds o f a pp rox ima te ly 2 26 k g p e r d unum o n l a nd f e r t i l ized w i th a c omb ina t ion o f n i t rous s o i l b rough t f r om t he n e ighbo r ing c l i f fs , a n ima l d ung , a nd o ven a shes . A cco rd ing t o Wendo r f a nd S ch i ld ( 1980a :271) , t he whea t y i e lds o b ta ined b y r ecen t ly s eden te r ized b edou in o n t he N i le f l oodp la in n ea r l s na r ange f r om 5 t o 6a rdabs ( 1a rdab = 1 44 k g ) p e r f eddan o f l a nd ( = 1 .038 a c res o r4 .2 d unums ) , s ugges t ing y i e lds o f 1 7 1 t o2 06 k g p e rd unum . S urp r is ing ly , o ne o ft he i r i n fo rman ts n o ted t ha t i nt he n e ighbo r ing Wad i R amd i in , ar eg ion c on ta in ing a pp rox ima te ly 1 7 d unums o fa rab le l a nd p rev ious ly c u l t iva ted o nce e ve ry t h ree t of i ve y ea rs a f te r i n f requen t r a ins , w hea t y i e lds a s h i gh a s 8a rdabs p e rf eddan , o ra round 2 74 k g p e rd unum , were o f ten o b ta ined ( Wendo r f a nd S ch i ld 1 980a :27 1) . A ppa ren t ly , r i ch s o i ls w a te red b y n a tu ra l r uno f f c an p roduce w hea t y i e lds h i ghe rt han t hose n o rma l ly o b ta ined a l ong t he N i le 's f l oodp la in . F o r c ompa r ison , o f f ic ia l s t a t is t ics ( Un i ted A rab R epub l ic 1 965 :86 -89 ) o n whea t p roduc t iv i ty t h roughou t a l l o f E gyp t f r om 1 950 t h rough 1 964 i n d ica te a mean whea t y i e ld o f 2 58 k g p e r d unum , w i th a n a nnua l r ange o fy i e lds f r om 2 0 1 t o3 00 k g p e rd unum . I n c on t ras t t o t he whea t y i e ld d a ta p resen ted a bove , mode rn h yb r id whea ts c er ta in ly e xh ib i t g rea te r p roduc t iv i ty . F o r e xamp le , h yb r id whea ts p roduced 4 47 k g p e r d unum i n D enma rk a nd 4 45 k g p e r d unum i nt he N e the r lands d ur ing 1 965 ( Ayk royd a nd D ough ty 1 970 :43 ) , wh i le mode rn h yb r id whea ts i nt he U n i ted K ingdom p roduced 4 04 k g p e rd unum i n 1 970 a nd 5 21 k g p e r d unum i n 1 980 ( Spedd ing 1 983 :471) . H oweve r , t o p ro jec t s uch i n c reases i n ac l ina l f ash ion i n to t he p as t wou ld c er ta in ly b e e r roneous . F o r i n s tance , t he a ve rage y i e ld o f n onhyb r id d ry fa rmed whea t o n p easan t f a rms i nt he C en t ra l A g r icu l tu ra l R eg ion o fR uss ia f r om 1 883 t h rough 1 900 o n ly amoun ted t oa pp rox ima te ly 6 90 l b s p e ra c re , o r r ough ly 7 7 k g p e r d unum ( Benne t t 1 935 :2 1) , h a rd ly s pec tacu la r w hen c ompa red t o an a tura l p roduc t iv i ty o f5 0t o8 0 k g p e rd unum a s n o ted e a r l ie rf o rw i ld s t ands o fm ixed whea t a nd b a r ley u nde rf avo rab le c ond i t ions i nt he e as te rn Ga l i lee . Wh i le h umans h ave s e lec t ive ly b red whea ts t o i n c rease t he i r i nher i ted p roduc t ive p o ten t ia l , s uch p rac t ices d i d n o t b eg in u n t i l t he l a te e i gh teen th a nd e a r ly n i ne teen th c en tu r ies A .D . i nr eg ions o fw es te rn E u rope . T he re a re n o v a r ia t ions i n whea t y i e lds p r io r t ot h is p er iod wh ich c anno t b e a t t r ibu ted t o t he n a tura l p heno typ ic p las t ic i ty o f whea ts g iven r eg iona l v a r ia t ions i ns o i lf e r t i l i ty ,t he u se o ff e r t i l ize r , c l ima te , a nd r a in fa l l o ri r iga t ion .

T he N a ture o fP reh is tor ic a nd H is tor ic S eed S e lec t ion a nd t he H is to ry o fE a r ly H yb r id iza t ion E f fo r ts I nl i gh to ft he t empo ra l a nd s pa t ia ld a ta o n whea ty i e lds p resen ted a bove , r eade rs m ay n ow b e s urp r ised t ol e a rn t ha t t he c a re fu l s e lec t ion o fd es i rab le s eed f o rt he n ex t y ears s ow ing h as p robab ly b een p rac t iced o n ay ea r ly b as is s i nce t he i n i t ia t ion o fc e rea l h usbandry . P resumab ly r e f lec t ing p reh is to r ic p rac t ices , V a r ro ( 1 .52 .1) a dv ised s e t t ing a s ide t he l a rges t a nd b es t e a rs o f g ra in f o r u se a s n ex t y ea r 's s eed s t ock . T h is s ugges t ion w as r epea ted b y C o lume l la ( 1 .9 .11) , w ho a dded t ha t i ft he y i e ld t ha t y ea r was b e t te r t han a ve rage , s eed s t ock s hou ld b e s e lec ted f r om t he t h reshed g ra in o n t he b as is o fl a rges t s i ze a nd h eav ies t we igh t . T h is l a t te r s ugges t ion was r epea ted b y P l iny ( XV 1 1 .195 ) . T he c on t inu i ty o fs uch p rac t ices i n to t he p resen t i sr e f lec ted i nt he 1 903 e d i t ion o ft he E ncyc lopaed ia B r i tann ica , whe re f a rme rs w e re a dv ised t os e lec tt he i r whea t s eed s t ock b y s epa ra t ing " t he h eav ies t a nd p l umpes t g ra ins " ( 1903 :533 ) . S im i la r ly , a g r icu l tu ra l is ts i nA f r ica s e lec t t he b es t h eads o fg ra in f o r n ex t y ea r 's s eed s t ock ( Ha r lan e ta l . 1 973 :321; H aswe l l 1 953 :31) . F ur the r , t he u n ive rsa l n a tu re o f s uch p rac t ices i s s eem ing ly r e f lec ted i nt he i n c reased mean s i ze o f s eeds f r equen t ly o bserved among a ppa ren t n once rea l c u l t iva rs i nt he N ew Wo r ld ( Asch a nd A sch 1 978 , 1 980 ; S immonds 1 976 ; S t rueve r a nd V ickery 1 973 :1205 -9 ; W i lson 1 98 1; Y a rne l l 1 965 , 1 977 , 1 978 ) , i n c lud ing s umpweed ( I va a nnua ) , s un f lowe r ( He l ian thus a nnuus ) , a nd g oose foo t ( Chenopod ium s p . ) . W i th y ea r ly s e lec t ion f o r s eed s i ze a nd w e igh t o ve r s eve ra l m i l lenn ia , why h aven ' t mean whea t y i e lds g radua l ly i n c reased t h rough t ime? F ur the r , w hy s hou ld i tb et ha t s ome r a ces o fw i ld 5 0

emmer , b a r ley , a nd o a ts p roduce s eeds a s l a rge o r l a rge rt han s ome r aces o ft he i r c u l t iva ted c oun terpar ts ( Zoha ry 1 969 :58 ; H a r lan e t a l . 1 973 :319 )? T he c ommon a nswer t o t hese q ues t ions l i es i nt he k nown e f fec ts o fd i rec t iona l s e lec t ion f o rs i ng le t r a i ts i np redom inan t ly s e l f p o l l ina t ing c e rea ls . C er ta in ly , sma l ler s eeds g ene ra l ly p roduce sma l l -seeded p rogeny , w h i le l a rge s eeds g ene ra l ly p roduce l a rge -seeded p rogeny . H oweve r , i nt he a bsence o ff o rced c ross -b reed ing a nd a ssoc ia ted a dd i t ive g ene t ic v a r iab i l i ty , e xpe r imen ts i nt he d i rec t iona l s e lec t ion o fs uch t r a i ts a s s eed s i ze a nd we igh t i np redom inan t ly s e l f -po l l ina t ing c e rea ls i n d ica te t ha t t he n e t e f fec t i s g ene ra l ly t or a ise t he mean o ft hese t r a i ts t ow i th in t he u ppe rr ange o fv a r iab i l i ty i ns eed s i ze a nd w e igh t e xh ib i ted b y t he i n i t ia l b ase p opu la t ion ( s ee A l la rd e t a l . 1 968 :99 103 , 11-113 ) . F u r the r , w h i le d i rec t iona l s e lec t ion f o ra ny o ne c ha rac te rc hanges t he m ean o ft ha tc ha rac te r i n t he d i rec t ion o ft he s e lec t ion , c hanges a l so o ccur i no the r , s eem ing ly u n re la ted t r a i ts , w i th i n tens ive s e lec t ion o f ten r educ ing t he p er fo rmance o f p rogeny a s measu red b y s eed y i e ld ( A l la rd e ta l . 1 968 :113 ) . I ti sf o r t hese r easons t ha t t he t empo ra l ly a nd s pa t ia l ly d i ve rse w hea t y i e ld d a ta u nder a ve rage d ry fa rm ing c ond i t ions p resen ted e a r l ie r f a l l w i th in t he u ppe r r ange o f p roduc t iv i ty r epor ted f o r w i ld s t ands u nde r f avo rab le e nv i ronmen ta l c ond i t ions , a nd f u r the r e xp la in why s ome w i ld c erea l r aces p roduce s eeds a s l a rge o r l a rge r t han r aces o f t he i r c u l t iva ted c oun terpa r ts . S eed s e lec t ion b y s i ze a nd w e igh t o n ay ea r ly b as is f o rt housands o fy ea rs h as s pec i f ica l ly s e rved t os t ab i l ize t he mean y i e ld o fp redom inan t ly s e l f -po l l ina t ing c e rea ls t h rough b road ly b ased mass s e lec t ion , r e su l t ing i nt h e c rea t ion o fl a nd races o fc e rea ls h i gh ly a dap ted t o r e g iona l e nv i ronmen ta l c ond i t ions a nd w h ich p ossess p roduc t ive c ha rac te r is t ics f a l l ing w i th in t h e u ppe r r ange o fv a r iab i l i ty e xh ib i ted b yt he i rw i ld p rogen i to rs . T ha tt he p e rpe tua t ion , r a the rt han i mp rovemen t , o fh i gh mean y i e ld w as t he e xp l ic i tg oa lo f s uch p rac t ices i nt he p as ti sr e f lec ted i nV i rg i l 's G eo rg ics ( 1 . 197 ) : Ih ave s een h ow s eeds t ha ta re s o r ted w i th t he u tmos tc a re W i l ld egene ra te , u n less e ve ry y ea rt he t a sk O f p i ck ing o u tt he l a rges t , o ne b yo ne , i sr epea ted [ 1 982 :34 -5 ] . T h is f o rm o f s e lec t ion s hou ld n o t b e c on fused w i th s e lec t ive b reed ing , n o r s hou ld i tb e e qua ted w i th g radua l ly i n c reas ing y i e lds . Among c u l t iva ted , p redom inan t ly s e l f -po l l ina t ing c erea l p opu la t ions , o ccas iona l n a tu ra l h yb r id iza t ion w i th w eed r a ces o n t he ma rg ins o ff i e lds d oes o ccu r , b r t s e ldom , i fe ve r , o n a mass ive s ca le ( Ha r lan 1 965 : 174 ; H a r lan a nd Z ohar y 1 966 :1076 ;Z oha ry e ta l . 1 969 ) . T he t y p ica l p a t tern i so ne i n vo lv ing a l o ca l ized e rup t ion o f ah yb r id s wa rm . S uch c ases a re i n te rm i t ten t i n t i me a nd s pace , e rup t ing l o ca l ly f o r ag ene ra t ion o rt wo , a nd t hen q u ick ly s ubs id ing . T h rough t h is p rocess , t h e l o ca l ly a dap t ive t r a i ts o fw eed r a ces a re i n t roduced i n to c u l t iva ted c r ops . I th as b een s ugges ted t ha t w i ld emme r e vo lved a s ar e su l t o f s uch n a tura l h yb r id iza t ion b e tween s t ands o fw i ld e i nko rn a nd t he g oa tg rass A eg i lops s pe l to ides ( Ha r lan a nd Z oha ry 1 966 :1078 ) , a n e ven t wh ich h ad p resumab ly o ccu r red b yt he e a r ly K eba ran p e r iod j u dg ing f r om t he emme r f ound a t t he s i te o f N aha l O ren i n I s rae l ( Noy e t a l . 1 973 ) . F ur the r , r ecu r ren t n a tura l i n t rog ress ion b e tween c u l t iva ted emme r a nd t he g oa tg rass A eg i lops s quar rosa h as b een s ugges ted a s l e ad ing t o t he e vo lu t ion o f t he b read w hea ts t e rmed T r i t icum X a es t ivum ( Fe ldman 1 976 :123 ; Z oha ry 1 969 :60 -63 ; Z oha ry e ta l . 1 969 :63 -64 ) . H oweve r , t he i n ten t iona l c r oss -b reed ing o fp redom inan t ly s e l f -po l l ina t ing c e rea ls t oe nhance t he i rp roduc t ive t r a i ts i sn o t a t tes ted t ou n t i lt he e nd o ft he e i gh teen th c en tury A .D . Wh i le t he n a tu re o f c ross -po l l ina t ion w as a nc ien t ly r ecogn ized i nr espec t t o ma le a nd f ema le d a te p a lms , t he p rac t ice o fa r t i f ic ia l p o l l ina t ion i np a lms was s o le ly d i rec ted t owa rds t he p roduc t ion o f f r u i t r a the r t han f o r p urposes o f b reed ing ( Dzhapa r idze 1 967 :2 ; R ober ts 1 965 :11-12 ) . U nder n a tura l c ond i t ions , d a te p a lms p roduce r ough ly e qua l n umbe rs o f ma le a nd f ema le t r ees , b u t o n ly f ema les p roduce d a tes . U nde rc u l t iva t ion , g row ing l a rge n umbe rs o f n on f ru i t ing ma les i s aw as te fu l u se o f l a nd , s i nce a l a rge n umbe r o ff ema les c an b e h and p o l l ina ted f r om as ma l l n umbe ro f ma les . I twas n o t u n t i l t h e i n ves t iga t ions o fR udo lph J akob C ame re ro ft he U n ive rs i ty o fT üb ingen i nt he l a te s even teen th a nd e a r ly e i gh teen th c en tur ies A .D . t ha t t he p oss ib i l i t ies o f s e lec t ive 5 1

h yb r id iza t ion i np redom inan t c ross -po l l ina t ing p l an ts s uch a s s p inach , h emp , a nd ma ize w e re f i rs t s ugges ted ( Dzhapa r idze 1 967 :20 -2 1; R ober ts 1 965 :12 -15 ) . F o l low ing h i s l e ad , e a r ly b o tan is ts i nt he e i gh teen th c en tu ry p roduced t he f i r s t i n ten t iona l h yb r ids o f c a rna t ions , a pp le t r ees , p r im roses , c ucumbe rs , ma ize , a nd o the r p redom inan t c ross -po l l ina t ing p l an ts . I n 1 760 , J oseph G o t t l ieb K o l reu te r b egan as e r ies o fp rec ise h yb r id iza t ion e xpe r imen ts w h ich s erved t o b o th e s tab l ish b as ic p l an t h yb r id iza t ion t echn iques a nd t o r e i fy t he p roduc t ive p o ten t ia ls o f s uch p rac t ices ( Dzhapa r idze 1 967 :29 ; R obe r ts 1 965 :34 -6 1) . E ven s o , i twas n o t u n t i l 1 795 t ha t T homas A nd rew K n igh t , w ho f ounded t he L ondon H or t icu l tu ra l S oc ie ty i n 1 804 t o p romo te t he s c ien t i f ic s e lec t ion o f h yb r ids , r ecogn ized t ha t p redom inan t s e l f -po l l ina t ing c e rea ls c ou ld b e h yb r id ized b y i n te r -sow ing d i f fe ren t v a r ie t ies i n t he s ame f i e ld , a nd t h rough t h is p rocess p roduced t he f i rs t k nown h yb r id whea t r aces ( Dzhapa r idze 1 967 :33 ; P e rc iva l 1 92 1:36 1 ; R ober ts 1 965 :114 ) . S ubsequen t ly , s eve ra l c e rea l h yb r ids w e re d eve loped b y v a r ious h o r t icu t tura l is ts d u r ing t he n i ne teen th c en tu ry , a l though i t w as n o t u n t i l 1 848 t o 1 85 1 t ha t h yb r id whea ts w e re g rown o n al a rge s ca le ( Pe rc iva l 1 921 :36 1) . S ign i f ican t ly , D a rw in ( 1893 ) was we l l -awa re o f t he d i f f icu l t ies e ncoun te red b y h i s c on tempo ra r ies i n t he i r a t temp ts t o e nhance t he p roduc t ive t r a i ts o f whea ts t h rough t he c rea t ion o f h yb r id s t ra ins , l ead ing h im t o d oub t t ha t s uch p rac t ices h ad c ha rac te r ized t he p reh is to r ic p as t . B u t whe the r i na nc ien t t imes s e lec t ion was me thod ica l ly p ursued we may w e l l d oub t , when w e h ea r h ow l a bo r ious t he w ork h as b een f ound b y L e C ou teur a nd H a l le t t . A l though t he p r inc ip le o f s e lec t ion i ss o i mpor tan t , y e t t he l i t le wh ich man h as e f fec ted , b y i n cessan t e f fo r ts d u r ing t housands o f y ea rs , i n r ende r ing t he p l an ts mo re p roduc t ive o r t he g ra ins mo re n u t r i t ious t han t hey w e re i nt he t ime o ft he o l d E gyp t ians , wou ld s peak s t rong ly a ga ins t i t s e f f icacy . B u t w e mus t n o t f o rge t t ha t a te ach s uccess ive p e r iod t he s t a te o f a g r icu l tu re a nd t he q uan t i ty o f manure s upp l ied t o t he l a nd w i l l h ave d e te rm ined t he max imum d eg ree o f p roduc t iveness ; f o r i twou ld b e i mposs ib le t o c u l t iva te a h i gh ly p roduc t ive v a r ie ty , u n less t he l a nd c on ta ined a s u f f ic ien t s upp ly o ft he n ecessa ry c hem ica l e l emen ts [ Da rw in 1 893 :336 ] . H e re , D a rw in h ad r ecogn ized t wo i mpo r tan t r easons f o r w hy t he e a r l ier s e lec t ive b reed ing o f s e l f -po l l ina t ing c e rea ls w ou ld h ave b een u ncommon . S ince a nnua l g rasses h ad e vo lved w i th e no rmous p heno typ ic p las t ic i ty i n s eed p roduc t ion , t he f ocus o fe a r ly h uman a t temp ts t o i n c rease c e rea l p rodu c t iv i ty l o g ica l ly i n vo lved b o th s e lec t ion b y s eed s i ze a nd we igh t a nd s o i l f e r t i l iza t ion , wh i le o p t imum p roduc t iv i ty was r ecogn ized a s b e ing a ch ieved o n r i ch s o i ls w i th f avo rab le r a in fa l l a nd /o rr unof p a t te rns . N a tu ra l o r a r t i f ic ia l means o f i r iga t ing r i ch s o i ls we re a l so r ecogn ized a s f avorab ly a fec t ing c e rea l p roduc t iv i ty . H oweve r , t he e f for t r equ i red t o p roduce h yb r id l and races o f p redom inan t s e l f p o l l ina t ing c e rea ls wou ld h ave h ad l i t le p ayof i nt he a bsence o f i n c reas ing ly i n tens ive s o i l f e r t i l iza t ion , a nd f u r the r r equ i red c omm i tmen t a nd p a t ience w i th n o g ua ran ty o f c ons is ten t ly i n c reased p roduc t ive r esu l ts . F o re xamp le , T homas K n igh td esc r ibed h i s i n t ia l h yb r id w hea ts t ra ins a sf o l lows : S ome o ft hose Io b ta ined we re e xce l len t , o the rs v ery b ad , a nd n one o ft hem p e rmanen t . B y s epa ra t ing t he f i rs t v a r ie t ies , a mos t a bundan t c rop was p roduced , b u t i nq ua l i ty was n o t e qua l t o t he q uan t i ty ; a nd a l t he d i sca rded v a r ie t ies a ga in a nd a ga in made t he ir a ppea rance [ as q uo ted i n R ober ts 1 965 :114 ] . I na dd i t ion t oe a r l ie rs uccesses i np roduc ing p l an t h yb r ids a nd t he g row th o ft he f i r s ts eed s t ock t r ade b e tween Ge rmany , F rance a nd B r i ta in i nt he l a te s even teen th a nd e a r ly e i gh teen th c en tur ies ( Food a nd A g r icu l tu re O rgan iza t ion 1 975 :v i i ) , K n igh t l i ved i na n e nv i ronmen t wh ich p rov ided as t rong e conom ic i n cen t ive f o r h i st ed ious e f fo r ts a tp roduc ing h yb r id w hea ts . B r i ta in h ad c eased t o b e s e l f -su f f ic ien t i ng ra in p roduc t ion d u r ing t he s econd h a l f o ft he e i gh teen th c en tury , a nd h ad s ubsequen t ly b ecome d ependen t u pon i mpor ts t o s upp lemen t h ome p roduc t ion ( M inch in ton 1 953 ) . D u r ing t he l a s t t we lve y ea rs o ft he e igh teen th c en tury , 5 2

t h is p rob lem i nB r i t ish a g r icu l tu re b ecame a cu te . F rom 1 789 t h rough 1 795 , a l b u t t wo B r i t ish h a rves ts we re e i the r s can t o r p oo r , a nd d u r ing t he f ou r y ea r p er iod f r om 1 792 t h rough 1 795 , whea t p r ices r o se 1 51 % , l e ad ing t o w idesp read c i v i l d i scon ten t a nd f ood r i o ts . I n Oc tobe r o f 1 795 , t he c a r r iage o f t he k i ng was a t tacked b y r i o te rs o n h i s way t o o pen p a r l iamen t . I nt h is e nv i ronmen t , K n igh t s u re ly r ecogn ized t he p o ten t ia l e conom ic a nd s oc ia l r ewa rds wh ich wou ld a cc rue t o a nyone who c ou ld s uccess fu l ly i n c rease B r i t ish whea t p roduc t iv i ty t h rough s e lec t ive b reed ing . T he s e lec t ive b reed ing o f p redom inan t s e l f -po l l ina t ing c e rea ls i nt he p reh is to r ic p as t i s t he re fore u n l ike ly . I ft he s e lec t ive b reed ing o f c e rea ls was p rac t iced a t a l , i t mo re p robab ly o ccur red w i th t he p redom inan t c ross -po l l ina t ing c e rea ls ma ize , p ea r l m i l le t , a nd J ob 's t ea rs , o r t he p a r t ia l s e l f , p ar t ia l c r oss -po l l ina t ing s o rghums a nd r yes . I n te res t ing ly , t he mos t d rama t ic c hanges i nt he mo rpho log ica l t r a i ts o f d omes t ic c e rea ls a re a ssoc ia ted w i th t hese s pec i f ic s pec ies , i n c lud ing i n c reased s i ze o f i n f lo rescences , " condensa t ion " t h rough t he s ho r ten ing o f i n te rnodes a nd b ranches , a nd " mu l t ip l ica t ion "t h rough i n c reases i nr ank o r r ow n umbe ro r i nt he n umbe r o f b ranches o ft he i n f lo rescence ( Ha r lan e ta l . 1 973 :3 15 -3 18 ) . H oweve r , s i nce n a tu ra l h yb r id iza t ion b e tween i n d iv idua ls wou ld a nnua l ly o ccur i nf i e lds o ft hese p ar t icu la r c e rea ls , t he s imp le s e lec t ion o f n ex t y ea r 's s eed s t ock f r om p heno typ ica l ly p rom is ing i n d iv idua ls c ou ld r esu l t i nt he s ame mo rpho log ica l s h i f ts i n dependen t o fi n ten t iona l b reed ing . P resumab ly a wa re o fs uch p rocesses , H op i a nd N ava jo c o rn a g r icu l tu ra l is ts i nt he S ou thwes t l a v ish e x t reme c a re i n t he s e lec t ion o f s eed s t ock ( H i l l 1 938 :27 -28 ) , b u t a l so a ss ign s epa ra te p l o ts o f l a nd f o r t he i r d i f fe ren t v a r ie t ies o f c o rn i na n a c t ive a t temp t t o p erpe tua te t he s pec i f ic t r a i ts o f e ach v a r ie ty ( B rown e t a l . 1 952 :599 ) . R ega rd less , s e lec t ion f o r s eed s i ze a nd w e igh t , b u t n o t s e lec t ive b reed ing , c ha rac te r ized t he h usband ry o f p redom inan t s e l f -po l l ina t ing c e rea ls p r io r t o t he e nd o f t he e igh teen th c en tury A .D . T he e f fec t o fs uch p rac t ices was t o s t ab i l ize mean p roduc t iv i ty w i th in t he u ppe r r ange o f p roduc t ive v a r iab i l i ty e xh ib i ted b y t he i r w i ld p rogen i to rs . H ence , e thnog raph ic a nd h i s to r ic d a ta o n n on -hyb r id , p redom inan t ly s e l f -po l l ina t ing c e rea ls d oes n o t a ppea r t o b e i n app rop r ia te f o ru se i nc a lcu la t ing t he r ange o fp o ten t ia l c e rea l y i e lds i nt he p reh is to r ic p as t .

A nc ien ta nd Modern C a lo r i c V a lues T he c a lo r ic v a lue o fw hea ts , a nd p resumab ly o the rc e rea ls a sw e l l , d oes n o ta ppea rt oh ave s i gn i f ican t ly c hanged a s ar e su l to fd omes t ica t ion . I ns t anda rd a na lyses , c u l t iva ted whea ts h ave b een f ound t o c on ta in 6 0 -80% c arbohyd ra tes , 8 15 % p ro te ins , a nd 1 . 5 -2 .0 % f a ts ( Fe ldman 1 976 :121) . T h is r ep resen ts ar ange o fc a lo r ic v a r iab i l i ty f r om 2 ,855 t o3 ,980 k ca l /kg . H oweve r , t he mean c a lo r ic c on ten t o f modem h a rd w hea ts i sc ommon ly s e t a t3 ,320 k ca l /kg , a nd mode rn s o f tw hea ts a t3 ,330 k ca l /kg ( Ayk royd a nd D ough ty 1 970 :18 ) . W i ld e inko rn c o l lec ted b y H a r lan i nT u rkey ( 1967 :198 ) was f ound t oc on ta in 2 2 .82% c rude p ro te in , 6 0 .04% n i t rogen f ree e x t rac t ( mos t ly c a rbohyd ra tes o the r t han f i b re ) , a nd 2 .64 % f a t . F rom t hese d a ta , t he a pp rox ima te e ne rgy v a lue o ft he w i ld e i nko rn h a rves ted b y H a r lan wou ld b e 3 ,567 k ca l /kg . Mode rn h i gh q ua l i ty h a rd r ed w in te r whea t a na lyzed a tt he s ame t ime was f ound t o c r in ia in 1 4 .5 % c r ude p ro te in , 6 8 .7% n i t rogen f ree e x t rac t a nd 1 .62 % f a t , s ugges t ing a n e ne rgy c on ten t o f 3 ,474 k ca l /kg . T hese d a ta s ugges t t ha t mode rn whea ts a c tua l ly c on ta in mo re c a rbohyd ra tes b u t s l igh t ly l e ss p ro te in a nd f a t t han w i ld e i nko rn , a nd may e xh ib i t c a lo r ic v a lues wh ich a re a round 2 . 7% l owe r t han t he i rw i ld c oun te rpa r ts .

S umma ry A s p rev ious ly o bserved f o r ' h i s tor ic a nd e thnograph ic d a ta o n p as to ra l l abo r a nd p roduc t iv i ty , s im i la r d a ta o n c e rea l c u l t iva t ion d oes n o t a ppea r t o b e i n app rop r ia te f o r u se i n e s tab l ish ing t he r ange o fp o ten t ia l r e tu rn r a tes f o r p reh is to r ic c e rea l s t ra teg ies . H oweve r , wh i le i ti ss uspec ted t ha t t he e ne rge t ics o f p as to ra l s t ra teg ies i n vo lv ing e x tens ive h usband ry h ave v a r ied l i t le s i nce t he p reh is to r ic p as t , t echno log ica l i n nova t ions h ave c e r ta in ly a f fec ted t he e ne rge t ics o f c e rea l c u l t iva t ion . A s w i l l b e d ocumen ted i nC hap te r 6 , s uch i n nova t ions h ave a ppa ren t ly i n c reased t he r e tu rn r a tes f o r c e rea l c u l t iva t ion t h rough t ime . I n C hap ter 7 , t he e f fec ts o f t hese s h i f ts i n a l te r ing t he e ne rge t ic r e la t ionsh ips b e tween p as to ra l a nd c e rea l s ubs is tence s t ra teg ies w i l l b e e xam ined . 5 3

CHAPTER 4

COMPARAT IVE E COLOGY OF THE RELEVANT AN IMAL AND P LANT S PEC IES

I n t roduc t ion I nt h is c hap te r , d a ta r e levan tt oa n u nde rs tand ing o fv a r ia t ions i nt he c ompa ra t ive e co logy o f p as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts , a nd c u l t iva t ion s t ra teg ies i n vo lv ing w hea t a nd b a r ley , a re p resen ted . Wh i le c a lcu la t ions o fp as to ra l a nd c e rea l r e tu rn r a tes o n ly r equ i re d a ta o nt he c a lo r ic p roduc t iv i ty a nd l a bo rc os ts a ssoc ia ted w i th e ach s t ra tegy g i ven t he s pec i f ic a n ima ls o rp l an ts i n vo lved , d a ta o n t he g ene ra l e co logy o fe ach s pec ies i sr equ i red f o r c ompa ra t ive e ne rge t ic a na lyses o f a l te rna t ive f ood p roduc t ion o p t ions u nder s pec i f ic e nv i ronmen ta l c ond i t ions . I n i t ia l ly , v a r ia t ions i nc er ta in e vo lved c ha rac te r is t ics o f c ame ls , c a t t le , s heep a nd g oa ts w h ich wou ld a fec tt he i rr e la t ive d es i rab i l i ty a s p as to ra l h e rd a n ima ls i ns pec i f ic e nv i ronmen ts a re p resen ted . S ubsequen t ly , ma thema t ica l mode ls o fp as to ra l c ame l , c a t t le , s heep a nd g oa t h erd g row th d ynam ics u nde r a ve rage e nv ironmen ta l c ond i t ions a re e xam ined , n o t ing v a r ious d i sease f ac to rs t ha t c ou ld p o ten t ia l ly a l te r g row th a nd c onsequen t p roduc t iv i ty e xpec ta t ions b ased u pon t hem . S ubsequen t ly , t he e co logy , n a tu ra l d i s t r ibu t ion a nd e a r ly e xp lo i ta t ion o fw i ld w hea ts a nd b a r leys a re d i scussed , a l ong w i th s im i la rd a ta o n t he i r e a r ly c u l t iva t ion . N o t ing t ha t h yd ro log ic a nd s o i l n u t r ien t c ond i t ions c ons t i tu te p r ima ry l i m i t ing f ac to rs i nc e rea l g row th a nd p roduc t iv i ty , a t ten t ion i s f ocussed u pon t he g eograph ic c i rcumsc r ip t ion o f " op t ima l " m ic roenv i ronmen ts f o re a r ly c e rea l f ood p roduc t ion s t ra teg ies .

S ome C ompa ra t ive A t t r ibu tes o fC ame ls , C a t t le , S heep a nd G oa ts I n t roduc t ion T axonom ica l ly , c ame ls b e long t o t he f am i ly C ame l idae o ft he i n f rao rde r T y lopoda o ft he o rde r A r t iodac ty la . C a t t le , s heep a nd g oa ts b e long t o t he f am i ly B ov idae o f t he i n f rao rde r P eco ra o ft he s ubo rde r R um inan t ia o ft he o rde r A r t iodac ty la . T he r ad ia t ion o ft he B ov idae a nd C ame l idae c o inc ides w i th t he d eve lopmen t o f w idesp read g rass lands i nt he M iocene a nd P l iocene , a nd s eve ra l member s o fb o th f am i l ies e vo lved w i th s im i la r ma jo rf o regu t f e rmen ta t ion c hambe rs o r c ompa r tmen ts . T hese p recede t he t r ue s t omach whe re d iges t ive e nzymes a nd a c id / a re s ec re ted . I n t he a n te r io r c hambe rs , m ic robes u t i l ize t he c e l lu lose a nd r e la ted p o ly accha r ides o f i n ges ted f o rage , p roduc ing v o la t i le f a t ty a c ids w h ich t hese h erb ivores t hen d iges t . Wh i le r um ina t ion , i n wh ich f o res tomach d iges ta i s r egurg i ta ted , r echewed a nd r e swa l lowed i n af r equen t , r egu la r p a t te rn , o ccu rs among a l t he P eco ra a nd T y lopoda , c ame ls a re u sua l ly n o t a ss igned t ot he s ubo rde r R um inan t ia b ecause t he i r f o regu t c ons is ts o ft h ree c hambe rs r a the rt han t he f ou r c hambe rs u sua l ly f ound among " t rue " r um inan ts ( C lu t ton -B rock 1 981:123 ; Gau th ie r -P i l te rs a nd D agg 1 981:3 -5 ; H ume a nd Wa rner 1 980 :673 -677 ; J an is 1 976 :769 -770 ; Mac fa r lane 1 964 :5 13 ; R edd ing 1 98 1:54 -58 ; S chm id t -N ie lsen 1 979 :34 -38 ) . D ur ing t he c ourse o ft he i r p hy logeny , c ame ls , c a t t le , s heep a nd g oa ts a l so e vo lved w i th s eve ra l mo re d i s t inc t a t t r ibu tes a ssoc ia ted w i th s pec i f ic e nv i ronmen ta l c ond i t ions . I np a r t icu la r , t he c ame l 's c e leb ra ted a dap ta t ion t o x e r ic e nv i ronmen ts r ead i ly i s o la tes i tf r om a l o f t he B ov idae , w i th t he p ar t ia l e xcep t ion o f t he g oa t . I nt he f o l low ing s ec t ions , s ome o f t he mo re d i s t inc t ive a t t r ibu tes o f t hese f ou r h e rd s pec ies a re d i scussed w h ich w ou ld s pa t ia l ly a nd t empo ra l ly a fec tt he r e la t ive r e turn r a tes o fp as to ra l s t ra teg ies b ased u pon t hem .

Ma in ta in ing T he rma lS tab i l i ty Mamma ls p roduce me tabo l ic h ea t when t r ans fo rm ing f ood i n to work , a nd a l so a bso rb e nv i ronmen ta l h ea t f r om d i rec t o r i n d i rec t s o la r r ad ia t ion . T h is c umu la t ive h ea t l o ad mus t b e b a lanced w i th h ea t l o ss i no rde rf o r mamma ls t o ma in ta in t he rma l s t ab i l i ty . H ea t l o ss i sp r ima r i ly a ch ieved b y r ad ia t ion , c onvec t ion a nd c onduc t ion f r om t he b ody t ot he e nv i ronmen t , o r b y t he e vapo ra t ion o f wa te rf r om t he s k in a nd r e sp i ra to ry p assages ( B ianca 1 968 :101103 ) . I na dd i t ion t ov a r ious b ehav io ra l s t ra teg ies , d e fense a ga ins t e xcess ive d i rec t o r i n d i rec t a bso rp t ion o fs o la r r ad ia t ion may b e a ch ieved b y r educ ing h ea t r e ta in ing b ody i n su la t ion o r b y i n c reas ing t he r e f lec tance o f t he h a i r . C onve rse ly , e xcess ive h ea t l o ss t o t he e nv i ronmen t mus t a l so b e c oun tered , g ene ra l ly t h rough t he c onserva t ion o f h ea t t h rough i n su la t ion , t he i n te rna l p roduc t ion o f mo re h ea t , o r b y v a r ious b ehav io ra l s t ra teg ies . I ng ene ra l , e ne rgy e xpend i tu re i n d e fense a ga ins t c o ld i s mo re e conom ica l ly a ch ieved b y i n c reas ing b ody i n su la t ion t han b y i n c reas ing h ea t p roduc t ion ( B ianca 1 968 : 104 ) . C a t t le ma in ta in r e la t ive ly c ons tan t b ody t empe ra tu res w h ich d i u rna l ly v a ry a round 2 ° C , a nd p r ima r i ly d i ss ipa te e xcess b ody h ea t b y s wea t ing ( Mac fa r lane 1 968a : 172 , 1 986b :267 ; S chm id t N ie lsen 1 979 :72 -77 ; Y ouse f e ta l . 1 968 :273 , F i gu re 1 7 -4 ) . P an t ing p l ays a s econda ry r o le i n t he i r h ea t r egu la t ion . F o r e xamp le , t he Z ebu o r B rahman c a t t le ( Bos i nd icus ) o f mo re s ou the r ly l a t i tudes s ec re te a pp rox ima te ly 6 t imes mo re s wea t t han t hey e vapo ra te a s wa te r f r om t he i r r esp i ra tory t r ac t , ap rocess i n vo lv ing g rea te rv o lume a nd mo re e f fec t ive c oo l ing t han t ha tf ound among t he E uropean c a t t le ( Bos t aurus ) o f mo re n or the r ly l a t i tudes ( B ianca 1 968 :110 ; Mac fa r lane 1 968a :172 ; S chm id t -N ie lsen 1 979 :77 ) . I ng ene ra l , t he mo re h ea t t o le ran t Z ebu o r B rahman b reeds o f c a t t le h ave s hor t , smoo th a nd h i gh ly r e f lec t ive c oa ts ( Mac fa r lane 1 968a :168 , 1 968b :266 ; S chm id t -N ie lsen 1 979 :78 ) . B yc on t ras t , s heep a nd g oa ts p r ima r i ly r e ly o n p an t ing t od i ss ipa te e xcess b ody h ea t , wh i le s wea t ing p lays a s econda ry r o le , p r ima r i ly among s heep ( Mac fa r lane 1 968a :17 1-72 ; R edd ing 1 981 :194 -199 ; S chm id t -N ie lsen 1 979 :98 100 ; T e r r i l l 1 968 :248 , 2 50 ) . A l though s heep e xh ib i t o n ly mode ra te d i urna l t empe ra tu re v a r ia t ions o f a round 1 . 5° t o 2 ° C( Mac fa r lane 1 968b :265 ; S chm id t -N ie lsen 1 979 :95 ) , s ome e v idence e x is ts t ha t g oa ts may b e a b le t o l e t t he i r b ody t empe ra tu re r i se mo re t han s heep ( Redd ing 1 98 1 :194 ) . T rop ica l a nd d eser t s heep g ene ra l ly h ave h a i r ie r t y pes o f f l eece t han mo re n or the rn b reeds , w i th f i be rs l y ing n ea r ly p a ra l le l w i th t he s k in p roduc ing a c oa t wh ich i s smoo th a nd r e f lec t ing i n af ash ion s im i la rt ot ha t o fg oa ts ( Mac fa r lane 1 968a : 168 , 1 968b :266 ) . S uch c oa ts a low f o r c onvec t ive c oo l ing a nd t he s ho r t -wave r e f lec t ion o f i n com ing r ad ia t ion w h i le n e i the r wa te r n o r h ea t i st r apped t o a ny e x ten t . L oose , l o ng a nd t ang led woo l o r h a i r -woo l f l eeces p rov ide i n su la t ion f r om b o th h i gh a nd l ow t empe ra tures , b u t wh i le c onvec t ion r emoves h ea t f r om t he o u te r l a ye rs , az one o f h ea ted a i r d eve lops i nt he s t i l l i n te rs t ices o ft he w oo l t ow h ich t he s un p ene t ra tes . A s ar e su l t , t he re i sl e ss e f fec t ive c onvec t ive a nd l o ng -wave r ad ian t em iss ion o f h ea t f r om t h is s ubsur face e n t rapmen t z one t han t he re i sf r om t he s u r face . B reeds w i th t hese t ypes o ff l eece may b ecome o ve rhea ted i nt he s un ( Mac fa r lane 1 968a :168 ) . F i ne f l eece , s uch a st ha tf ound i nt he me r ino s heep o fS pa in , A f r ica , R uss ia a nd A us t ra l ia p reven ts a f r ee a i r movemen t . I tt he re fo re r es is ts t he i nwa rd c onvec t ion o f h ea t a nd p rov ides f o r a n e xponen t ia l f a l l o f t empe ra ture a c ross t he i n su la t ion ( Mac fa r lane 1 964 :511; 1 968a :166 ; S chm id t -N ie lsen 1 979 :96 -98 ) . C ame ls d o n o t p an t , a nd l i ke c a t t le , p r ima r i ly d i ss ipa te e xcess b ody h ea t b y s wea t ing ( Mac fa r lane 1 968b :267 ) . H oweve r , u n l ike c a t t le , t he d romeda ry c ame ls ( Came /us d romeda r ius ) o f mo re s ou the r ly , h o t /d ry d ese r ts h ave a n e vo lved c apac i ty f o r af l uc tua t ing h ea t l o ad , w i th d a i ly b ody t empe ra tu re v a r ia t ions o f u p t o 6 ° Co r mo re b e ing a ccep tab le ( Gau th ier -P i l te rs 1 974 :545 ; S chm id t -N ie lsen 1 979 :44 -45 ) . T hus , r a the r t han d i ss ipa t ing a l l a cqu i red h ea t b y e vapo ra t ion , t he d romeda ry c ame l i sa b le t oi n te rna l ly s t o re h ea td u r ing t he d ay t ob e d i ss ipa ted l a te rb yc onduc t ion a nd r ad ia t ion i nt he c oo l o fn i gh ts . T h is c apac i ty r educes h ea tf l ow f r om t he e nv i ronmen t d u r ing t he d ay a s i t s b ody t empera tu re r i ses , a nd f u r the r p os tpones u n t i l l a te r i n t he d ay t he momen t when e vapo ra t ive c oo l ing mus t b e u sed t o p reven t af u r the r r i se i nt he i r h ea t l o ad . Whe the rt h is c apac i ty e x is ts t ot he s ame e x ten t among t he B ac t r ian c ame ls ( Came /us b ac t r ianus ) o f mo re n or the r ly c o ld /dry d ese r ts h as n o t b een e xpe r imen ta l ly d emons t ra ted ( Bu l l ie t 1 975 :31) . H oweve r , L eese ( 1927 :11) c asua l ly o bse rved t ha t B ac t r ians c ou ld n o t w i ths tand p ro longed p e r iods o f e x t reme h ea t r e la t ive t o t he c apac i t ies o f d romeda r ies . I n g ene ra l , B ac t r ians a re n o tc ons ide red w e l l a dap ted t oh o tc ond i t ions ( Mac fa r lane 1 964 :510 ) . 5 6

D romeda ry c ame l c oa ts e xh ib i t an or th -sou th c l ina l v a r ia t ion , w i th s ho r te ra nd l i gh te rw in te r c oa ts p reva len t i ns ou the r ly l a t i tudes , a nd t h icke r , d a rke r w in te r c oa ts p reva len t i nn o r the r ly l a t i tudes ( Leese 1 927 :10 ; Mac fa r lane 1 964 :510 -511, 1 968b :266 ) . F ur the r , t he w in te r c oa ts o f B ac t r ian c ame ls a re l o nge r , h eav ie r , t h icke r a nd d a rke r t han t hose o f d romeda r ies , a nd t he amoun t o f h a i r r e ta ined a f te r s hedd ing i sg ene ra l ly g rea te r ( Gau th ie r -P i l te rs a nd D agg 1 98 1 :3 ; L eese 1 927 :11 ; L eona rd 1 894 :96 -97 ; Mac fa r lane 1 964 :510 ) . A s n o ted b y S chm id t -N ie lsen ( 1979 :47 ) , wh i le t he c oa t o ft he d romeda ry s erves t o p rov ide i n su la t ion w h ich d im in ishes h ea t f l ow f r om t he e nv i ronmen t , ab a lance mus t b e s t ruck b e tween t h is d es i rab le t r a i t a nd t he n eed t oa low f o rt he d i ss ipa t ion o fi n te rna l ly s t o red h ea tt h rough c onduc t ion a nd r ad ia t ion . I fr anked a cco rd ing t ot he i r o ve ra l l a b i l i ty t o ma in ta in t he rma l s t ab i l i ty u nde r h ea t s t ress , d romeda ry c ame ls wou ld b e a tt he t op o ft he l i s t , f o l lowed b y g oa ts , s heep a nd f i na l ly c a t t le . H owever , d romeda ry c ame ls f a l l t o t he b o t tom o f t he l i s t when c o ld e nv i ronmen ts a re c ons ide red . E uropean c a t t le ( Bos t aurus ) a nd B ac t r ian c ame ls ( Came /us b ac t r ianus ) p ossess s eve ra l t r a i ts wh ich make t hem b e t te r a dap ted t oc o ld e nv i ronmen ts t han t he i r mo re s ou the r ly c oun te rpa r ts ( Gau th ie r -P i l te rs a nd D agg 1 98 1:3 ; H ense l 1 968 :183 ; L eese 1 927 :11; L eona rd 1 894 :98 ; Y ouse f e t a l . 1 968 :233 ) . F ur the r , b o th s heep a nd g oa ts a re w e l l -adap ted f o r c o ld e nv i ronmen ts , a l though g oa ts a re g ene ra l ly mo re s ens i t ive t o c o ld t han a re s heep ( Scha l ler 1 977 :69 -75 ; T e r r i l l 1 968 :249 -250 ) . D romeda ry c ame ls , h oweve r , a re q u i te p oo r ly a dap ted t o w e t /co ld c ond i t ions , b e ing e x t reme ly s uscep t ib le t of a ta l r e sp i ra tory d i seases w i th h i gh d ea th r a tes o ccu r r ing among c a lves ( Bha rgava e t a l . 1 963 : 187 188 ; C ross 1 917 :110 -114 ; D ickson 1 951:418 ; J enn ings -B ram ley 1 906 :205 ; L ancas te r 1 981:10 , 1 01; L eese 1 927 :282 -291; L eona rd 1 894 :221-225 ; T o r ry 1 973 :72 , 1 27 ) .

Ma in ta in ing Wa te rB a lance Mamma ls l o ose b ody wa te r t h rough e vapo ra t ion , u r ina t ion a nd d e feca t ion . Wh i le s ome d ep le t ion o f b ody wa te r may b e t o le ra ted f o r s hor t p e r iods , a l wa te r l o ss mus t e ven tua l ly b e o fse t b y a n e qua l i n take . I ng ene ra l , mamma ls o b ta in wa te r d i rec t ly b y d r ink ing , o r i n d i rec t ly f r om t he wa te r c on ta ined i nt he i r f ood o r f r om t he o x ida t ion o f h yd rogen i nt he i r f ood ( Ha fez 1 968b :13 ) . C a t t le u nder e x tens ive h usbandry , a nd p a r t icu la r ly c a lves , a re g ene ra l ly u nab le t o mee t t he i r wa te r r equ i remen ts s imp ly b y e a t ing f r esh p l an ts ( Behnke 1 980 :26 ) . F ur the r , t he i r wa te r c onsump t ion i sh i gh r e la t ive t o c ame ls , s heep a nd g oa ts ( Schm id t -N ie lsen 1 979 :79 -80 ) . A s t he i r wa te r i n take i n c reases u nde r h ea t s t ress , t he i ru r ine o u tpu t a nd f r equency o fu r ina t ion a l so i n c rease , wh i le u r ine c oncen t ra t ions d ec rease . T hus , t he i r wa te r i n take a ppea rs t o i n c rease mo re t han i sr equ i red t oc ove r i n c reased e vapo ra t ion , a nd i ti sl i ke ly t ha t c a t t le d r ink i n c reas ing q uan t i t ies o f wa te r u nde r h ea t s t ress i no rde rt oc oo l t he i r b od ies d i rec t ly r a the r t han t o a vo id d ehyd ra t ion ( Schm id t -N ie lsen 1 979 :80 ) . I ng ene ra l , Z ebu o r B rahman c a t t le ( Bos i nd icus ) h ave ad a i ly wa ter t u rnove r o f a round 2 5% l e ss t han E u ropean c a t t le ( Bos t aurus ) u nde r s im i la r amb ien t t empe ra tures a nd c onsum ing t he s ame f eed ( Ha fez 1 968a :205 , T ab le 1 5 -2 ; Mac fa r lane 1 964 :524 , 1 968a :173 , T ab le 1 2 -4 ) . E ven s o , a l c a t t le l o se wa ter a tg rea te r r a tes t han d o o the r r um inan ts u nde r d eser t c ond i t ions , a nd t he i r a b i l i ty t o r educe s uch o u t f low i s s ma l l ( Mac fa r lane 1 964 :536 ; , 1 968b :27 1 , 2 73 ) . S heep a nd g oa ts a re b o th a b le t o mee t much o f t he i r wa te r r equ i remen ts f r om f o rage d ur ing p e r iods o fl u sh g raz ing a nd b rows ing , a nd g ene ra l ly r equ i re l e ss f r equen t wa te r ing t han d o c a t t le d u r ing d ry s easons ( Coppock 1 985 ; D yson -Hudson a nd McCabe 1 985 ; D ah l a nd H jo r t 1 976 :239 , 2 49 -250 ; R edd ing 1 981:200 ; T o r ry 1 973 :126 ) . F ur the r , b o th s heep a nd g oa ts a re a b le t or educe t he i r wa te r l o ss v i au r ine e xc re t ion u nde r h ea t s t ress , w i th g oa ts b e ing mo re t han t w ice a s e f fec t ive i nt h is p rocess t han s heep ( Mac fa r lane 1 968a :175 -176 ; R edd ing 1 981 :200 ; S chm id t -N ie lsen 1 979 : 101) . Ove ra l l , g oa ts u se l e ss wa te r t han s heep , a nd s heep o n ly a bou t h a l f a s much a s c a t t le ( Mac fa r lane 1 968a :176 , 1 968b :270 -27 1) . L i ke s heep a nd g oa ts , c ame ls a re a b le t o mee t much o f t he i r wa te r r equ i remen ts f r om f o rage d ur ing p er iods o f l u sh g raz ing a nd b rows ing ( Coppock 1 985 ; D yson -Hudson a nd McCabe 1 985 ; Gau th ie r -P i l te rs 1 974 :544 -545 ; Gau th ie r -P i l ters a nd D agg 1 981:50 ; T o r ry 1 973 :119 ) , b u t e xceed a l o ther s t ock i n t he i r e f f ic iency o f wa te r u se a nd t o le ra t ion o f d ehyd ra t ion u nde r h ea t s t ress . D romedary c ame ls c an t o le ra te awa te r l o ss e qu iva len t t o2 7%

5 7

o ft o ta l b ody we igh t , a nd c an t hus w i ths tand wa te rd ep r iva t ion f o r l o nge rp e r iods t han a ny o the r mamma l e xposed t os im i la rc ond i t ions ( Schm id t -N ie lsen 1 979 :62 -63 , 7 0 ) . Wa te r l o ss t h rough f eces i sa l so g rea t ly r educed i nc ame ls . A cco rd ing t oS chm id t -N ie lsen ( 1979 :60 ) , f r ee -g raz ing c a t t le w i l l l o se o ve r 5 00 gm o f wa te r p e r 1 00 gm o f f eces , wh i le d romedar y c ame ls f ed o n d ry d a tes a nd h ay w e re o bse rved t o l o se o n ly 1 09 gm o f wa te r p e r 1 00 gm o ff eces . Gau th ie r -P i t te rs a nd D agg ( 198 1 :62 -63 ) s im i la r ly r epo r ted t ha t wh i le c a t t le may l o se 2 0 -40 l i t e rs o f wa te r p e r d ay t h rough f eces , c ame ls o n ly l o se 6 -7 l i t e rs o r l e ss p er d ay . U nde r w a te r s t ress , c a t t le a re a b le t o r educe t he i r n o rma l f eca l wa te r l o ss b y 4 0% a f te r t h ree d ays w i thou t wa ter , s heep 5 5% a f te rs i x d ays w i thou t wa te r , a nd c ame ls 6 0 -62% a f te rf i ve d ays w i thou tw a te r ( Mac fa r lane 1 968b :273 ) . F u r the r , wh i le d romeda ry c ame ls c an e xc re te l a rge q uan t i t ies o fd i lu te u r ine w hen wa te ri s p l en t i fu l , t he i r p ower fu l k i dneys c an a l so p roduce e x t reme ly c oncen t ra ted u r ine u nde r wa te r d ep r iva t ion ( Gau th ier -P i t te rs a nd D agg 1 981 :66 ; S chm id t -N ie lsen 1 979 :54 -57 ) . T he c ame l 's a b i l i ty t o p roduce u r ine w i th a h i gh c oncen t ra t ion o f s a l ts n o t o n ly a l lows f o r a r educed e xpend i tu re o f wa te rf o r s ec re t ion , b u t a l so a lows t hem t od r ink h i gh ly s a l ine wa te r a nd t o e a t p lan ts w i th h i gh s a l ts a nd o xa l ic a c id c on ten ts ( Schm id t -N ie lsen 1 979 :55 -56 ) . D romeda ry c ame ls a re a b le t ot o le ra te wa te rc on ta in ing 5 .5% N aC I ,w h i le s heep c an o n ly t o le ra te wa te rw i th 1 . 3% N aC I a nd c a t t le 1 . 0% N aC I ( Gau th ie r -P i t te rs a nd D agg 1 98 1 :66 -67 ) . D ue t ot he i r l ow r a tes o f wa te rt u rnove r , c ame ls r equ i re l e ss f r equen t wa te r ing t han o the r l i ves tock . R end i l le i n fo rman ts ma in ta in t ha t d u r ing d ry s easons , t he i r c ame ls c ou ld s urv ive f o r u pt os even w eeks w i thou t wa te r , w h i le t he i rs heep a nd g oa ts c ou ld n o t s urv ive f o r mo re t han a w eek a nd t he i r c a t t le f o r o n ly f our d ays ( Sa to 1 980 :11) . T h is a cco rds we l l w i th Mac fa r lane 's o bserva t ion ( 1968b :272 ) t ha t u nde r wa te r d ep r iva t ion , c ame ls c ou ld s urv ive f o r mo re t han 1 5 d ays , s heep a pp rox ima te ly s i x t o t en d ays , a nd c a t t le o n ly t h ree t o f i ve d ays . C ame ls i nt he A lge r ian h amada d u r ing s umme r t empe ra tu res o f 4 3 -46 ° Cw i th l e ss t han 1 0% h um id i ty o n ly r equ i red wa te r ing e very t h ree t o f ou r d ays ( Gau th ie r -P i l ters a nd D agg 1 981:52 ) . Among N g isonyoka h e rds i nn o r thwes te rn K enya , c ame ls g ene ra l ly d r ink e ve ry 4t o 6d ays d u r ing t he d ry s eason , a du l t sma l ls tock e ve ry o the r d ay , a nd c a t t le d a i ly ( Coppock 1 985 ; C oughenou r e t a l . 1 985 :625 , n .5 0 ; D yson -Hudson a nd McCabe 1 985 ) . S im i la r ly , A l Mu r rah c ame ls i nt he h ea t o fS umme r i nt he R ub ' a l -Kha l i d ese r t o fS a 'ud i A rab ia o n ly r equ i re wa te r e ve ry f ou rd ays ( Co le 1 975 :52 ) . I fo n ly f o rt he i r i n f requen t wa te r ing r equ i remen ts , c ame ls c ou ld u t i l ize f eed ing a reas t ha t wou ld b e i naccess ib le t o o the r d omes t ic s tock ( Gau th ie r -P i l te rs a nd D agg 1 981:53 ; S pence r 1 973 :17 ; T o r ry 1 973 :81) . I fr anked a cco rd ing t o m in ima l wa te r r equ i remen ts a nd e f f ic iency o f wa te r u se , c ame ls w ou ld a ga in b e a t t he t op o f t he l i s t , f o l lowed b y g oa ts , s heep a nd f i na l ly c a t t le . R e la t ive s uscep t ib i l i ty t o d i sas t rous h e rd l o sses d ue t o d rough t s hou ld p resumab ly f o l low t he s ame r ank ing . I ns ou the rn E th iop ia i n1 973 , 8 8% o ft he t o ta l c a t t le a s o pposed t o 6 9% o ft he t o ta l c ame ls p resen t i n1 972 a re s a id t oh ave d ied a s ar esu l t o fd rough t ( Dah l a nd H jo r t 1 976 :115 ) . S im i la r ly , d ur ing t he 1 980 -1981 d rough t i nn or thwes te rn K enya , N g isonyoka p as to ra l is ts l o s t 3 7% o f t he i r a du l t c ame ls a nd g oa ts , 4 3% o ft he i r a du l t s heep a nd 5 4% o f t he i r a du l t c a t t le ( McCabe 1 984 ) .

F eed ing E co logy U nder e x tens ive h usband ry , c a t t le a nd s heep p redom inan t ly d er ive t he i r f o rage e ne rgy f r om g raz ing o n h e rbaceous s pec ies , wh i le c ame ls a nd g oa ts a re l e ss s e lec t ive f eede rs , f r equen t ly d e r iv ing a l a rge p ropo r t ion o f t he i r e ne rgy f r om a dd i t iona l ly b rows ing o n woody s pec ies ( Asad 1 964 :52 ; B ehnke 1 980 :26 -29 ; C oppock 1 985 ; C oughenour e t a l . 1 985 :620 6 21 ; D ah l a nd H jor t 1 976 :244 , 2 53 ; D yson -Hudson a nd D yson -Hudson 1 970 :98 -99 , 1 09 ; H a fez 1 968b :203 -204 ; J ohnson 1 969 :9 ; Mac fa r lane 1 964 :535 ; R edd ing 1 981:72 -78 ; S a to 1 980 :15 , 1 8 -20 , 6 1; T o r ry 1 973 :119 ) . T hese f eed ing p a t terns a re e x t reme ly s i gn i f ican t i n r e la t ion t o t he r e la t ive p roduc t ive p o ten t ia l o f t hese h e rd s pec ies i n s pec i f ic e nv i ronmen ts , s i nce ma rked s easona l v a r ia t ions o ccur i nt he a va i lab i l i ty o ff o rage e ne rgy f r om h erbaceous ( he rb , g rass , v i ne ) a s o pposed t ow oody ( s h rub , t r ee ) s pec ies i nd ese r t , s teppe a nd s avanna e nv i ronmen ts .

5 8

F or e xamp le , t he s t eppe r ange lands o ft he S ahe l , l i ke o the r s t eppe , d ese r t a nd s avanna r ange lands , ma in ly c ons is t o ft ho rn -sh rubs d om ina ted b y a nnua l g rasses whose a c t ive p er iod o fg row th i sr e s t r ic ted t ot h et wo o rt wo a nd ah a l f mon ths o ft he s ho r t r a iny s eason ( B reman a nd d e W i t 1 983 : 134 1) . T h is a c t ive g row ing p e r iod i so n ly s l igh t ly l o nge r f o r t he i n d igenous p e renn ia l g rasses . I n d ry s easons a nd d rough t y ea rs , h erbaceous p roduc t ion i sm in ima l o r n onex is ten t . F ur ther , h erbaceous f o rage q ua l i ty , a s measu red b y i t s p ro te in c on ten t , r ap id ly d ec l ines a f te r t he r a ins c ease ( B reman a nd d e W i t 1 983 :1343 , 1 346 ; C oughenour e t a l . 1 985 :622 -623 ) , wh i le w oody f o rage ma in ta ins g reen b i omass much l o nge r i n to d ry s easons ( Coughenou r e ta l . 1 985 :622 ) . S ign i f ican t ly , a n i n ve rse r e la t ionsh ip b e tween t he q ua l i ty o f h e rbaceous b i omass a s measured b y i t s p ro te in c on ten t , a nd t o ta l h e rbaceous y i e ld u nde r r a in fa l l g rad ien ts , h as b een o bse rved i nt he S ahe l , S ou th A f r ica , a nd t he U n i ted S ta tes ( B reman a nd d e W i t 1 983 :1343 , 1 346 ) . I ng ene ra l , l ow wa te ra va i lab i l i ty p roduces asma l l amoun t o fh e rbaceous b i omass o fh i gh p ro te in c on ten t , wh i le h i gher wa te r a va i lab i l i ty r esu l ts i n ag rea te r h erbaceous b i omass o f i n c reas ing ly i n fe r ior q ua l i ty . D ue t o t he i r wa te r r equ i remen ts , t he r ange o f g raz ing i s mo re r es t r ic ted f o r c a t t le t han m ixed s heep a nd g oa t h e rds , w i th a l t h ree s pec ies b e ing much mo re c on f ined t han t he r ange o fc ame ls ( Mac fa r lane 1 968a :176 ) . I nh o tw ea the r , c a t t le h e rds a re g ene ra l ly l i m i ted t of o rag ing w i th in ao ne d ay 's wa lk o fw a te rs ou rces a nd s heep a nd g oa ts at h ree d ay 's wa lk , wh i le c ame l 's c an make ac i rcu i t o fs i x d ays o u t a nd as im i la rt ime b ack ( Mac fa r lane 1 968b :273 ) . T hus , i nd ry s easons , c a t t le a re g ene ra l ly r es t r ic ted t og raz ing w i th in a4 t o 1 3 km r ad ius o f wa te r s ources ( Dah l a nd H jor t 1 976 :239 -240 ; S pence r 1 973 :15 ; T ony 1 973 :81) . F o r e xamp le , R end i l le c ame ls may b e h e rded 5 0 -70 km f r om wa te rho les d u r ing d ry s easons , wh i le m ixed s heep a nd g oa t h e rds a re r e s t r ic ted t ow i th in a 2 0 -30 km r ad ius o ft he n ea res t wa terho les ( Sa to 1 980 :11 , 6 6 ) . S im i la r ly , Gab ra c ame l h e rds we re o bserved t of eed o n p as tures u p t o 4 8 km f r om we l ls d u r ing d ry s easons , wh i le G ab ra c a t t le h e rds w e re l i m i ted t og raz ing w i th in a 1 3 km r ad ius o f w e l ls ( Tor ry 1 973 :119 ) . H oweve r , w i th g ood p as tu rage c ond i t ions , Gab ra c ame ls c ou ld e xp lo i t p as tures u p t o 8 1 km f r om we l ls , t he i r sma l ls tock u p t o 4 8 km , a nd t he i r c a t t le 4 0 km ( Tor ry 1 973 : 8 1) . S ince a s i gn i f ican t p ropor t ion o f t he s hor t te rm a bundance o f h i gh q ua l i ty h erbaceous f o rage o ccu rs b eyond t he i r g raz ing r ad ius f r om wa te r s ou rces , much o ft he a nnua l b i omass i s u nava i lab le t o c a t t le ( Coughenou r e t a l . 1 985 :623 ) . F requen t ly , c a t t le mus t b e p rema tu re ly w i thd rawn f r om r eg ions c on ta in ing h erbaceous p l an ts w i th h i gh p ro te in c on ten t , n o t f o r l a ck o f f o rage , b u t f o r l a ck o f a va i lab le wa te r ( B reman a nd d e W i t 1 983 :1344 ) . T o al e sse r d eg ree , t hese s ame c ons t ra in ts o n t he a b i l i ty t ou t i l ize s easona l ly a va i lab le h e rbaceous f o rage wou ld a pp ly t o m ixed s heep a nd g oa t h e rds . Came ls , h oweve r , a re a b le t o e xp lo i t t he a nnua l a bundance o fp r ime h e rbaceous f o rage i nx e r ic r eg ions b e t te rt han a l o the rl i ves tock .

T he U n ique F ea tu res o fC ame lL ocomo t ion J us t a s c ame ls a re n o tt r ue r um inan ts , t hey a re a l so , i n as ense , n o tt r ue u ngu la tes . U n l ike c a t t le , s heep a nd g oa ts , t he c ame l 's f ee t a re n o t h ooved , b u t r a the rc ons is t o fl a rge c u taneous p ads w i th t wo a n ter io r t oena i ls ( C lu t ton -Brock 1 981:121-122 ; Gau th ier -P i l ters a nd D agg 1 98 1 :102 ; Mac fa r lane 1 964 :5 12 ) . T he i rf ee t a re c ompa ra t ive ly l a rge , w i th c ush ions a nd e l as t ic s t ruc tu res b e tween t he d i ve rg ing p ha langes a nd t he i rt h ick s o les , p rov id ing f o r b o th s t ab i l i ty i n s o f t s and a nd i n su la t ion a ga ins t h i gh s ur face t empe ra tu res . B ac t r ian c ame ls , h oweve r , d i f fer f r om d romeda r ies i nh av ing h arde r s o les a nd t oena i ls w h ich e x tend b eyond t he p ads o ft he i r f ee t ( Leese 1 927 :11; L eona rd 1 894 :97 -98 ) . D u r ing a s t r ide , t he c ame l 's s o le i sp laced d own f l a t o n t he g round a s i ts h i f ts i t s we igh t o n to t he f oo t . A s we igh t i sp u t o n t he p ad , i t s a rea i n c reases b y a bou t o ne th i rd . T he c ame l s ubsequen t ly p ushes o f f r om e ach f oo t w i th i t s t oes . Wh i le c ame ls c an s t ep w i thou t much d i scom for t o n sma l l p ebb les a nd e ven s t ony g round , t hey p re fe r t o f o l low w ind ing t r a i ls whe re t he re a re n o l o ose s tones ( Gau th ie r -P a te rs a nd D agg 1 981:102 ; L eona rd 1 894 :208 , 3 26 ) . A lso d i s t inc tf r om c a t t le , s heep a nd g oa ts i st ha t t he c ame l 's ma in f o rce o fl o como t ion , l i ke t ha t o f g i ra f fes , i si n i t s f o requa r te rs r a the r t han i t s h i ndqua r te rs ( Gau th ie r -P i l te rs a nd D agg 1 98 1 :105 -106 ; L eona rd 1 894 :206 -207 ) . A s ar e su l t , w hen a c ame l c omes t o ah i l l , i t" pu l ls " i t s b ody u p s low ly , a nd i s f u r the r u nab le t o j ump o ve r o bs tac les . C ame ls f a re p oo r ly i n 5 9

moun ta inous r eg ions , a l though B ac t r ians g ene ra l ly d o b e t te r t han d romeda r ies ( Leona rd 1 894 :208 ) . T he c ame l i sa l so u n ique i nb e ing t he o n ly s pec ies t ha t g ene ra l ly p aces r a the rt han t r o ts when mov ing a t a mode ra te s peed , r a is ing b o th l e gs o n t h e s ame s i de a t a pp rox ima te ly t he s ame t ime ( Dagg 1 974 ; Gau th ie r -P i t te rs a nd D agg 1 981:10 1) . T h is s i gn i f ican t u se o fu n i la te ra l s uppo r t ing l e gs , wh ich i sa l so a ppa ren t t o al e sse r e x ten t among g i ra f fes , s eems p a r t icu la r ly s u i ted f o r l o ng legged a n ima ls i np reven t ing t he l e gs o n o ne s i de f r om h i t t ing e ach o the r a s t hey s w ing f o rwa rd i nl o ng s t r ides , a nd a l so a lows t he musc les o n o ne s i de o ft he b ody t o wo rk a lmos t i nu n ison t o p u l l t he l e gs f o rwa rd . A ppa ren t ly , i ni t s a dap ta t ion t od ese r t e nv i ronmen ts w i th f l a t t e r ra in a nd h i gh v i s ib i l i ty , t he c ame l 's e vo lu t ion h as s ac r i f iced maneuve rab i l i ty a nd g a i t s t ab i l i ty f o rf r eedom o fs w ing f o rt he l e gs a nd h ence l e ng th o fs t r ide ( Gau th ie r -P i l ters a nd D agg 1 981:108 ) . Wh i le t hese a t t r ibu tes o fl o como t ion p a r t icu la r ly s u i t t he c ame l t od ry d eser t e nv i ronmen ts , t he c omb ina t ion o f l o ng l e gs , u n i la te ra l s uppor t , a nd p added f ee t make d romeda ry c ame ls v i r tua l ly h e lp less o n we t , s l ippe ry s ur faces ( Gau th ier -P i l te rs a nd D agg 1 981: 102 , 11; Mac fa r lane 1 964 :512 ; T o r ry 1 974 :67 ) . T he d e le ter ious n a ture o f t hese a t t r ibu tes was u n ive rsa l ly a pp rec ia ted b y t he v e te r ina r ians a nd e xpe r ienced o f ice rs o fB r i t ish c ame l c o rps . A s d esc r ibed b y Ma jo r A . G . L eona rd o f t he S econd E as t L ancash i re R eg imen t , av e teran o f n ume rous c ampa igns emp loy ing c ame ls i nI n d ia , A fghan is tan a nd t he S udan ( 1894 :208 ) : I n we t a nd s l ippe ry , s o f t a nd b oggy g round h e [ t he d romeda ry c ame l ] i s mo re o r l e ss u se less , b ecomes n ervous a nd a nx ious , a nd s eems t o l o se a l c on f idence i nh imse l f . H is f ee t a ppea rt ol o se a l h o ld o ft he f o rme r , a nd h e s l ips a nd s l ides a bou t ; wh i le i nt he l a t te r h e s i nks d own a cco rd ing t ot he y i e ld ing n a tu re o ft he s o i l , l o ses h i s h ead , f l ounde rs , f a l ls , a nd p robab ly r up tures , o r i fo ve rwe igh ted d i s jo in ts h i s h i ps , o rs p l i ts h imse l f u p . I np a r t icu la r , B r i t ish v e te r inar y s u rgeons n o ted t he t endency o ft he d romeda ry 's f o re o r h i nd l e gs t o s l ide o u twa rds o n muddy o r s l ippery g round , r esu l t ing i nf r ac tures o f t he h umerus , d i s loca t ions o f h i p j o in ts , a nd r up tures o f musc les ( Cross 1 917 :14 ; L eese 1 927 : 119 ) . N o t s u rp r is ing ly , o f f ice rs i nc ha rge o fc ame l c orps we re s t rong ly a dv ised t od i scon t inue ma rches i f r a ins made t he p assage s l ippery ( Leese 1 927 :119 ; L eona rd 1 894 :209 ) . H oweve r , B ac t r ian c ame ls a ppa ren t ly f a re b e t te r o n we t a nd s l ippe ry s o i ls t han d o d romeda r ies d ue t o b o th t he v a r ia t ions i nt he i r f oo t s t ruc ture n o ted e a r l ie r , a nd t ot he i r s ho r te r l e gs ( Leona rd 1 894 :97 -98 , 3 26 ) . T hese p ecu l ia r i t ies o f d romeda ry c ame l l o como t ion s eem ing ly p ose f ew p rob lems f o r c ame l p as to ra l is ts i nt he d eser ts o f t he N ea r E as t a nd N or th A f r ica , f o r e thnog raph ies f r om t hese r eg ions a re d evo id o f r e fe rences t o t he d i f f icu l t ies e xper ienced b y c ame ls o n we t , s l ippery s o i ls . H owever , t h is i sn o t s o f o r c ame l p as tora l is ts i n E as t A f r ica . T o r ry ( 1973 :83 , 1 974 :67 ) e xp l ic i t ly n o ted t ha t t he Gab ra wou ld move t he i r c ame l c amps f r om muddy a nd s l ippe ry l o ca t ions d u r ing r a iny s easons b ecause o ft he d i f f icu l t ies e xpe r ienced b y t he i rc ame ls , a nd b ecause t hey b ecame n ervous , a nx ious , a nd a t temp ted t oe scape i fc on f ined i nk r aa ls w i th we t , s l ippe ry s ur faces . S im i la r ly , S a to ( 1980 :22 ) s t a ted t ha t t he R end i l le wou ld r emove t he i r c ame ls f r om a reas wh ich b ecame muddy o r s wampy i nr a iny s easons d ue t ot he i rt endency t o s u f fe r s p ra ined l i mbs , n o t ing t ha t s p ra ins a nd b roken b ones we re o ne o ft he p r ima ry f ac to rs r e spons ib le f o r mor ta l i ty among a do lescen t a nd a du l t c ame ls ( 1980 :22 ) .

C ompa ra t ive H e rd G row th F ac to rs I n t roduc t ion E xc lud ing t he e fec ts o ft empo ra l a nd s pa t ia l e nv i ronmen ta l v a r iab i l i ty , a nnua l p as to ra l h e rd p roduc t iv i ty i sp r ima r i ly a f unc t ion o f t o ta l h e rd s i ze a nd g row th , h uman c ropp ing s t ra teg ies g i ven s pec i f ic h usband ry g oa ls , a nd t he p roduc t ive a t t r ibu tes o ft he h e rd 's c ons t i tuen t a ge a nd s ex c l asses . S ince b o th h erd s i ze a nd t he n umbe r o f i n d iv idua ls i ni t s a ge a nd s ex c l asses c hange d ur ing t he t empo ra l s pan o f ay ea r , a nd s i nce l o ng te rm s t ud ies o f i n d iv idua l p as to ra l h erds h ave n o t b een c onduc ted , ma thema t ica l mode ls o fp as to ra l h e rd g row th d ynam ics w i l l b e emp loyed i n Chap te r 5t o c a lcu la te t he e xpec ted p roduc t iv i ty o f h e rds u nder s pec i f ic 6 0

managemen t s t ra teg ies . T he f o l low ing s ec t ions d e ta i l s pec i f ic e co log ica l f ac to rs wh ich w ou ld a fec t p as to ra l h e rd g row th d ynam ics a nd a ssoc ia ted h e rd p roduc t iv i ty among c ame ls , c a t t le , s heep a nd g oa ts a s d ocumen ted i n t hese mode ls a nd i n e thnog raph ic a ccoun ts , a n ima l h usband ry manua ls a nd v e te r ina ry s ou rces . P opu la t ion g row th i n ap as to ra l h e rd e xpe r ienc ing n o em ig ra t ion o r i mm ig ra t ion i st he d i f ference t h rough t ime b e tween t he h e rd 's b i r th r a te a nd i t s d ea th r a te . T he b i r th r a te o f a p as to ra l h e rd i sp r ima r i ly a r esu l t o ft he s pec ies ' g es ta t ion l e ng th , b reed ing f r equency , a ge a t f i r s t p a r tu r i t ion ( b ir th ing ) , y ean ing r a te ( t he n umbe r o fa nnua l b i r ths p e r r ep roduc t ive ly ma ture f ema le ) , a nd h uman i n te r fe rence i nt he i r n a tu ra l b reed ing p rocess . T he d ea th r a te o f ap as to ra l h e rd r esu l ts f r om b o th t he n a tu ra l mor ta l i ty f ac to rs t ow h ich t ha t s pec ies i ss ub jec t a nd f r om h uman c ropp ing s t ra teg ies . I nd eve lop ing ma thema t ica l mode ls o fp as to ra l h e rd g row th d ynam ics , t he v a lues u sed f o r e s tab l ish ing a s pec ies ' b i r th a nd d ea th r a tes a re g ene ra l ly d e r ived f r om t he a va i lab le d a ta i n s uch a f ash ion a s t o a pp rox ima te t he a ve rage r esu l ts e xpec ted u nde r n o rma l c ond i t ions o f e x tens ive h usbandry , r ea l iz ing t ha t s uch v a lues may f l uc tua te w i ld ly f r om y ea r t oy ea r e ven i n t he s ame r eg ion . T h is b i r th a nd d ea th r a te d a ta i st hen u sed t o mode l a verage e xpec ted p as to ra l h e rd g row th d ynam ics a nd t o d e te rm ine t he a ge a nd s ex s t ruc tu re o f a n e s tab l ished h e rd . W i th t h is i n fo rma t ion , a nnua l p roduc t iv i ty f r om h e rds o fv a r ious s i zes i sc a lcu la ted o n t he b as is o f s pec i f ic h uman c ropp ing s t ra teg ies a nd d a ta o n t he a ve rage e xpec ted p roduc t iv i ty o f i n d iv idua l h e rd a n ima ls b y t he i r a ge a nd s ex c l asses , a ga in r ea l iz ing t ha t a c tua l p roduc t iv i ty w ou ld v a ry a cco rd ing t oe nv i ronmen ta l c ond i t ions a nd t he p roduc t ive c ha rac te r is t ics o fs pec i f ic r eg iona l b reeds . Mode ls o f p as tora l h erd g row th d ynam ics a nd p roduc t iv i ty h ave b een p rev ious ly d eve loped f o rd romeda ry c ame ls , c a t t le , s heep a nd g oa ts ( Dah l a nd H jo r t 1 976 ; R edd ing 1 98 1 ; S a to 1 980 ) . T he p resen t a na lys is w i l l s pec i f ica l ly f ocus u pon t he mode ls o f D ah l a nd H jor t ( 1976 :28 -76 , 1 40 181) f o r p as to ra l c a t t le h e rds i nA f r ica o f Z ebu ( Bos i nd icus ) a nd Z ebu d e r ived b reeds , a nd t hose o f R edd ing ( 1981) f o r N ea r E as te rn b reeds o f s heep a nd g oa ts . H oweve r , p rev ious mode ls o fc ame l h e rd g row th a nd p roduc t iv i ty a s d eve loped b y D ah l a nd H jo r t ( 1976 :78 -86 , 1 84 198 ) a nd S a to ( 1980 :28 -3 1) wou ld s eem mo re a pp rop r ia te f o r e x t reme r a the r t han a ve rage e xpec ted c ond i t ions u nde r e x tens ive h usband ry , d ue p r ima r i ly t o t he c ame l r ep roduc t ion , mo r ta l i ty , a nd p roduc t iv i ty d a ta wh ich t hey emp loyed . F o rt hese r easons , r ev ised mode ls o fd romeda ry c ame l h e rd g row th a nd o f f take h ave b een g ene ra ted i nA ppend ix A , w i th r ev ised c a lcu la t ions o f h e rd p roduc t iv i ty p resen ted i nA ppend ix B . T hese d a ta w i l l b e u sed i nt he p resen t a na lys is .

C ompara t ive R ep roduc t ion Age a tF i rs t P ar tu r i t ion . Wh i le c ame ls may b eg in t ob reed a s e a r ly a s t h ree y ea rs o fa ge , t hey a re n o t g ene ra l ly b red u nder e x tens ive h usbandry u n t i l t he i r f our th t o s i x th y ea r ( see A ppend ix A ) . S im i la r ly , c a t t le may b eg in t o b reed a s e a r ly a s t he i r s econd y ea r , b u t a re n o t g ene ra l ly b red u nde r e x tens ive h usbandry u n t i l t he i r t h i rd t of i th y ea r ( Behnke 1 980 :3 1; D ah l a nd H jor t 1 976 :33 -34 ; D yson -Hudson a nd D yson -Hudson 1 969 :88 , 1 970 :111) . B y c on t ras t , s heep a nd g oa ts a re g ene ra l ly a lowed t ob ea rt he i rf i r s to f fsp r ing a tb e tween o ne a nd t wo y ea rs o f a ge ( Behnke 1 980 :31 ; R edd ing 1 98 1:94 100 ) . Ges ta t ion P e r iod . L eng ths f o rc ame l g es ta t ion g ene ra l ly v a ry f r om 1 t o 1 3 mon ths , w i th a 1 2 t o 1 3 mon th g es ta t ion p e r iod b e ing w ide ly s ugges ted ( s ee A ppend ix A ) . Among p as to ra l c a t t le , t he g es ta t ion p e r iod i sg ene ra l ly g i ven a s 9t o 9 .5 mon ths ( Behnke 1 980 :31 ; T o r ry 1 973 :111; W i l l iamson a nd P ayne 1 965 :175 ) . F o r s heep a nd g oa ts , s t a ted l e ng ths o fg es ta t ion v ary f r om t h ree t o s i x mon ths ( Asad 1 964 :51, 1 970 :15 ; B ehnke 1 980 :29 ; S a to 1 980 :27 ; S pence r 1 973 :13 ; W i l l iamson a nd P ayne 1 959 :27 1 , 2 90 ) . Obv ious ly , t hese v a r ia t ions i n g es ta t ion l e ng th a f fec tt he f r equency a t wh ich membe rs o fe ach s pec ies c an b reed . B reed ing F requency . F ema le c ame ls w i l l n o rma l ly g i ve b i r th t oo ne c a l f e ve ry t wo t ot h ree y ea rs , w i th e nv i ronmen ta l f a c to rs f r equen t ly l i m i t ing c a lv ing t oo ne p eak s eason e ach y ea r ( s ee A ppend ix A ) . S im i la r ly , c a t t le u nde r e x tens ive h usband ry c an b reed o nce e very 1 4 t o 2 4 mon ths , w i th e nv i ronmen ta l f ac to rs f r equen t ly r es t r ic t ing c a lv ing t oo ne p eak s eason p e r y ea r ( Dah l a nd H jor t 1 976 :36 ; D yson -Hudson a nd D yson -Hudson 1 969 :88 , 1 970 :111) . T he 6 1

b reed ing f r equency o fs heep a nd g oa ts s eems s t rong ly i n f luenced b y d ay l e ng th a cco rd ing t o l a t i tude , w i th b i r ths e xpec ted y ea r round s ou th o f2 7 ° l a t i tude , b u tw i th s easona l c hanges i nd ay l e ng th g ene ra l ly r es t r ic t ing t he b reed ing s eason n o r th o f 2 7 ° l a t i tude ( Redd ing 1 981 :126 ) . R edd ing ( 198 1:8 1-94 ) c onc luded t ha t s heep a nd g oa ts i nt he N ea r E as t wou ld g ene ra l ly b e r es t r ic ted t ob reed ing o nce ay ea r , a l though h a l f o ft he e wes c ou ld l amb t w ice i n1 8 mon ths a nd a l d ams c ou ld b reed t w ice i n 1 2 mon ths i ft he i r p as tu rage was e spec ia l ly g ood . I n mo re s ou the r ly l a t i tudes , e wes may l amb u pt ot h ree t imes i nt wo y ea rs ( Dah l a nd H jor t 1 976 :90 -9 1 , 9 7 ; R edd ing 1 98 1:125 -126 ) , wh i le d ams may b reed f r om t h ree t o f our t imes e very t wo y ea rs ( Dah l a nd H jo r t 1 976 :92 ) . Y ean ing Ra tes . T he y ean ing r a te i s t he n umbe r o f b i r ths e xpec ted a nnua l ly p e r r ep roduc t ive ly ma ture f ema le i n ah e rd , a nd mus t t ake i n to a ccoun t t he f r equency o f b o th m u l t ip le b i r ths a nd r ep roduc t ive f a i lu res . Y ean ing r a tes o fa round 0 .4 o ccu r among c ame ls , a nd a re q u i te l ow b y c ompa r ison w i th t hose o fo the r d omes t ic s t ock . T hese l ow r a tes a re p r ima r i ly d ue t o t he c ame l 's b i enn ia l b reed ing p a t te rn , ar a r i ty o f mu l t ip le b i r ths , a nd r e la t ive ly h i gh a bor t ion r a tes ( see A ppend ix A ) . Y ean ing r a tes f o r p as to ra l c a t t le g ene ra l ly r ange f r om 0 .4 t o 0 . 8 ( Dah l a nd H jo r t 1 976 :35 ; T o rn y 1 973 :112 ) . D ah l a nd H jo r t a ssumed a r e la t ive ly l ow r a te o f 0 .58 i nt he i r mode l o f" no rma l " h e rd g row th ( 1976 :64 ) . F o ru n imp roved b reeds o f N ea r E as te rn s heep a nd g oa ts u nde r e x tens ive h usband ry , R edd ing ( 198 1 :110 ) e s tab l ished a y ean ing r a te o f0 .8 f o rs heep a nd 1 . 2 f o rg oa ts , t he l a t te rd ue t oh i gher r a tes o f mu l t ip le b i r ths among g oa ts . I n mo re s ou the r ly l a t i tudes , y ean ing r a tes may r i se t o 1 . 5 f o rs heep a nd 2 .0 f o rg oa ts a s ar e su l t o fn onseasona l b reed ing ( Dah l a nd H jo r t 1 976 :91 , 9 3 ; R edd ing 1 98 1:125 , 1 27 ) . D i seases a nd Abor t ion . R ep roduc t ive f a i lu res f r equen t ly o ccu r among l i ves tock a s ar e su l t o fs pec i f ic d i seases t ha t d i rec t ly a f fec t t he i r r ep roduc t ive o rgans , a nd n onspec i f ic d i seases t ha t r educe t he i r r ep roduc t ive e f f ic iency a s aw ho le . V a r ia t ions i nt he r eg iona l d i s t r ibu t ion o f s uch d i seases a nd t he r e la t ive s uscep t ib i l i ty o f h e rd s pec ies t ot hem wou ld a cco rd ing ly a l ter t he i r e xpec ted r ep roduc t ive r a tes i ns pec i f ic e nv i ronmen ts . A l though r ep roduc t ive f a i lu res c ommon ly o ccu r a s ac omp l ica t ion o f n ume rous d i seases , a bor t ions a re p ar t icu la r ly c ommon w i th b ruce l los is i n fec t ions . I n g ene ra l , c a t t le a re mos t s uscep t ib le t o t h is d i sease , f o l lowed b y g oa ts a nd s heep ( J ubb a nd K ennedy 1 963 :1:422 4 27 ) , w i th c ame ls a ppa ren t ly b e ing a s s uscep t ib le a s c a t t le ( K lös a nd L ang 1 982 :228 ; R u t te r 1 963 :119 -120 ) . P as teure l los is ( haemo r rhag ic s ep t icaem ia ) a l so a ppea rs t o b e f r equen t ly a ssoc ia ted w i th a bo r t ions i na l f our s pec ies ( Jubb a nd K ennedy 1 963 :1:172 -175 ; K lös a nd L ang 1 982 :227 -129 ; R u t te r 1 963 :121-122 ) . H oweve r , wh i le c a t t le , s heep a nd g oa ts f r equen t ly a bor t a s ac omp l ica t ion o f c on tag ious p l eu ropneumon ias ( Jubb a nd K ennedy 1 963 :1:170 -172 ) , c ame ls a re g ene ra l ly r es is tan t t o t hese s pec i f ic p neumon ias ( C ross 1 917 :111 ; R u t te r 1 963 :123 ) . E ven s o , c ame ls a re e spec ia l ly p rone t o a bo r t ion a s ac omp l ica t ion o f t r ypanosoma l i n fec t ions c aused b yf l age l la ted p ro tozoa s p read p r ima r i ly b y t aban id ( b i t ing ) f l i es a nd p oss ib ly b y t i cks ( C ross 1 917 :91 ; D ah l a nd H jor t 1 976 :245 ; L eese 1 927 :223 , 3 22 ; R u t ter 1 967 :6 13 ; S a lyaev 1 956 :483 ) . Wh i le c a t t le a re a l so s uscep t ib le t ot r ypanosomes , s ome p opu la t ions s u f fe r f ewe r c omp l ica t ions t han c ame ls , a nd l i ke s heep a nd g oa ts , p r ima r i ly a c t a s r ese rvo i rs o ft he i n fec t ion t o o the r a n ima ls ( Dah l a nd H jor t 1 976 :245 ; L eese 1 927 :223 ) . U n l ike e p idem ic b ac te r ia l d i seases wh ich r ap id ly s p read i nr eg ions a nd s ubsequen t ly s ubs ide , t r ypanosom iases p ose ac ons tan t , t hough f r equen t ly s easona l t h rea t . P andem ic f r om I n d ia a nd S ov ie t C en t ra l A s ia t h rough Mo rocco a nd Wes t A f r ica ( Leese 1 927 :223 -254 ; R u t te r 1 967 :6 12 ; S a lyaev 1 956 :485 ) , t he p r imary v ec to r o f t r ypanosom iases a re t aban id ( b i t ing ) f l i es wh ich b reed i n wa rm s easons n ea r o pen wa te r s ources a nd wooded a nd l ow ma rshes ( Cross 1 9 17 :94 -95 ; L eese 1 927 :219 -223 ; R u t te r 1 967 :6 11-6 13 ; S a lyaev 1 956 :483 -485 ) . We t /wa rm e nv ironmen ts i nhab i ted b y t aban id f l i es wou ld t here fore b e p a r t icu la r ly d e t r imen ta l t o c ame l h e rd g row th a nd p roduc t iv i ty . L eese a c tua l ly s ugges ted t ha t t he f a i lu re o f c ame ls t o t h r ive i n mo re s ou ther ly l a t i tudes i nA f r ica was a d i rec t r esu l t o ft he i n c reas ing p reva lence o f t aban id f l i es s uch a s t se tse c l ose r t o t he e qua to r ( 1927 :2 19 ) , a n o p in ion e choed b y B u l l ie t ( 1975 :40 ) a nd J ohnson ( 1969 :8 ) . I ng ene ra l , c ame l p as to ra l is ts i n t he d ese r ts o f mo re n or the r ly l a t i tudes s easona l ly a vo id k nown t r ypanosome z ones ( D ickson 1 95 1:418 ; L eese 1 927 :250 -25 1) , a l though i n E as t A f r ica , t h is i sn o t a lways p oss ib le ( Sa to 1 980 :20 , 6 7 ) .

6 2

C ompa ra t ive Mo r ta l i ty Mo r ta l i ty R a tes i nH e rd G row th Mode ls . A s n o ted a bove , t he d ea th r a te o f ap as to ra l h e rd r esu l ts f r om b o th t he n a tu ra l mo r ta l i ty f ac to rs t o wh ich t he s pec ies i ss ub jec t , a nd f r om h uman c ropp ing s t ra teg ies . I nt he mode ls o f c ame l h e rd g row th d eve loped i nA ppend ix A , c ame ls h ave amax imum l i f espan o f2 2t o2 3 y ea rs , w i th 3 8% o fa l n ew ly -bo rns d y ing b e fo re t he i rf ou r th y ea r , a nd t he t o ta l a du l t p opu la t ion h av ing a mo r ta l i ty r a te o f 3% p e r y ea r . I nD ah l a nd H jor t 's mode l o f" no rma l " c a t t le h e rd g row th ( 1976 :45 -8 , 6 4 -65 ) , c a t t le h ave a max imum l i f espan o f 1 3 y ea rs , w i th 3 6% o f a l n ew ly -bo rns d y ing b e fo re t he i rf i f th y ea r , a nd t he t o ta l a du l t p opu la t ion h av ing a mor ta l i ty r a te o f 5% p e r y ea r . I nR edd ing 's mode ls o fs heep a nd g oa t h e rd g row th i n t he N ea r E as t ( 198 1:80 136 ) , b o th s pec ies h ave a max imum l i f espan o fn i ne y ea rs , w i th 3 2% o f a l l ambs a nd 4 5% o fa l k i ds d y ing d ur ing t he i rf i r s t y ea r , a nd b o th a du l t e wes a nd d oes s ub jec t t o at o ta l mor ta l i ty r a te o f1 8% p e ry ea r . D iseases a nd Mo r ta l i ty R a tes . I na dd i t ion t ot he g ene ra l e nv i ronmen ta l c ond i t ions d e ta i led e a r l ie r , d i sease f ac to rs w ou ld a l so a l te rt he e xpec ted mo r ta l i ty r a tes emp loyed i nt hese mode ls o fa ve rage p as to ra l h e rd g row th d ynam ics . I ng ene ra l , wh i le o ne h e rd s pec ies w i l l s uccumb t o i t s o wn p a r t icu la rs e t o fd i seases , i ti so f ten a b le t or es is t t hose wh ich a re f a ta l f o r a no the r ( van d e r Wa l t a nd J ansen 1 968 :2 17 ) . V a r ia t ions i nt he d i s t r ibu t ion o ft hese d i seases a nd t he r e la t ive s uscep t ib i l i ty o f e ach h e rd s pec ies t o t hem wou ld a cco rd ing ly a l te r t he i r mo r ta l i ty r a tes a nd p o ten t ia l p roduc t iv i ty i ns pec i f ic e nv i ronmen ts . Wh i le a l h e rd a n ima ls a re s ub jec tt o av a r ie ty o f d i seases , o n ly t he mos t p rom inen tf a ta l d i seases o fl i ves tock w i l l b e n o ted h e re . C ame ls , c a t t le , s heep a nd g oa ts a re a l s uscep t ib le t oa n th rax ( C ross 1 917 : 102 104 ; J ubb a nd K ennedy 1 963 :1:295 -299 ; L eese 1 927 :268 -270 ; R u t te r 1 963 :119 ) , b u t o n ly c ame ls , c a t t le a nd s heep a re s uscep t ib le t o b lack q ua r te r ( C ross 1 917 :104 107 ; J ubb a nd K ennedy 1 963 :2 :402 -403 ; R u t te r 1 963 :120 ) . O f t he v a r ious p ox d i seases o fl i ves tock , o n ly s heeppox i s e spec ia l ly v i ru len t ( 2 -50% mo r ta l i ty ) , wh i le c ow , g oa ta nd c ame l p ox a re u sua l ly b en ign w i th v ery l ow mor ta l i ty r a tes ( C ross 1 917 :79 ; J ubb a nd K ennedy 1 963 :2 :525 -229 ; K lös a nd L ang 1 982 :230 ; L eese 1 927 :260 -264 ) . H oweve r , L eese ( 1927 :26 1) a l so n o ted t ha t wh i le c ame l p ox was s e ldom f a ta l among y oung c ame ls i no the rs easons , i twas k nown t od es t roy u pt o4 0% o ft he c a lves i n ah e rd d u r ing r a iny s easons , a ppa ren t ly i n c reas ing i nv i ru lence u nde r wa rm a nd w e t c ond i t ions . F oo t a nd mou th d i sease i sp a r t icu la r ly v i ru len t among y oung u ngu la tes , a f fec t ing c a t t le , s heep a nd g oa ts ( Jubb a nd K ennedy 1 963 :2 :6 -7 ) . C ame ls , h owever , a ppea r t o h ave a c ons iderab le d eg ree o f n a tura l i mmun i ty t o t h is d i sease ( C ross 1 917 :109 ; K lös a nd L ang 1 982 :229 ; L eese 1 927 :273 ) . R inde rpes t ( " ca t t le p lague " ) i sh i gh ly c on tag ious a nd f a ta l among c a t t le , a nd g enera l ly mo re v i ru len t among g oa ts t han s heep ( J ubb a nd K ennedy 1 963 :2 :2 12 2 ) . A ga in , c ame ls s eem t o h ave a c ons ide rab le d eg ree o f i mmun i ty t o t h is d i sease ( Cross 1 9 17 :85 ; L eese 1 927 :272 ) . D u r ing t he d i sas t rous A f r ican r i nde rpes t e p idem ic o f 1 890 1896 ( S inc la i r 1 979 :3 -6 ) , c ame ls w e re a c tua l ly i mpor ted f r om S oma l ia t oS ou th A f r ica i nt he h ope t ha t t hey c ou ld b e u sed t o r ep lace o xen ( Ru t te r 1 967 :6 11-6 13 ) . U n fo r tuna te ly , t he c ame ls i n s tead s ufe red f r om t r ypanosom iases . Men t ioned e a r l ier i nr e la t ion t o t he r e la t ive ly h i gh i n c idence o f a bo r t ions among c ame ls , t r ypanosom ias is i st he mos t s e r ious a nd d ead ly p ro tozoa l d i sease t o wh ich c ame ls a re p a r t icu la r ly s uscep t ib le ( C ross 1 917 :87 ; K lös a nd L ang 1 982 :225 ; L eese 1 927 :217 -254 , 3 22 ; R u t te r 1 967 :6 11) . I na l c ame l c oun t r ies , ah eavy morb id i ty o fu p t o3 0% a nd amor ta l i ty r a te o f a round 3% c an b e a t t r ibu ted t o t r ypanosoma l i n fec t ions ( K lös a nd L ang 1 982 :225 ; R u t te r 1 967 :611) . T he d eb i l i ta t ing e f fec ts o f t r ypanosoma l i n fec t ions a l so r educe t he c ame l 's r es is tance t o o the r d i seases , s uch a s p as teu re l los is a nd v a r ious o the r p neumon ias ( Cross 1 917 :91 , 11; L eese 1 927 :270 -27 1 , 2 82 -283 ) . D romeda ry c ame ls a re a l so p a r t icu la r ly s uscep t ib le t o e nzoo t ic e ncepha lomye l i t is ( 100 % mo r ta l i ty ) , ad i sease a ppa ren t ly s p read b yt i cks b u t wh ich i sn o tt r ansm iss ib le t oo the rd omes t ic l i ves tock ( K lös a nd L ang 1 982 :229 -230 ; O r jechow a nd P jedkow 1 952 ) . P resumab ly , t h is i st he f a ta l t i ck -bo rne c ame l d i sease f ea red b y t he R end i l le a nd Gab ra o f K enya ( J ones 1 984 :126 ; T o r ry 1 973 :82 -83 , 1 974 :67 ) . C a t t le a re a l so s uscep t ib le t ot he i r o wn t i ck -bo rne d i sease i nE as t A f r ica , k nown a s " Eas t C oas t f eve r , " b u t w h i le mor ta l i ty i sh i gh among t he c a t t le who c on t rac t t he d i sease , t he n umber o f a c tua l d ea ths s eems t o b e l ow ( Dyson -Hudson 1 960a , D yson H udson a nd D yson -Hudson 1 969 :8 1 , 1 970 :122 ) . I ng ene ra l , t he Z ebu o r B rahman c a t t le ( Bos 6 3

i nd icus ) o f mo re s ou the r ly l a t i tudes e xh ib i t a ma rked r e s is tance t ot i ck -bo rne d i seases r e la t ive t o b o th E u ropean c a t t le ( Bos t aurus ) a nd d romeda ry c ame ls ( Dah l a nd H jo r t 1 976 :245 ; H a fez 1 968b :11; J ohnson 1 969 :9 ; Mac fa r lane 1 964 :536 ; S pence r 1 973 :12 ; S u ther land 1 968 : 158 ; T o r ry 1 973 :407 ; v an d e r Wa f t a nd J ansen 1 968 :226 ) . S ign i f ican t ly , t he i n c idence o f d i sease c a r ry ing t i cks i n E as t A f r ica a ppea rs t o i n c rease i nr eg ions w i th g rea te r mo is ture a nd h i ghe r h um id i ty ( Dyson -Hudson 1 960a :254 ; T o r ry 1 973 :82 , 4 11) . I ng ene ra l , c ame ls , c a t t le , s heep a nd g oa ts s eem e qua l ly s uscep t ib le t ov a r ious e p idem ic b ac ter ia l d i seases a f fec t ing b o th r ep roduc t ion a nd mo r ta l i ty r a tes , w i th s pec ies -spec i f ic r e s is tance t oo ne ma jo rd i sease f r equen t ly o f fse t b y ag rea te rs uscep t ib i l i ty t oo ther s . H oweve r , i ti sa l so a ppa ren tt ha t ma jo r e nv i ronmen ta l d i f fe rences e x is t b e tween t he c ond i t ions f a vo rab le f o r c ame l h usband ry a nd t hose f avo rab le f o r c a t t le , s heep a nd g oa t h usbandry . Wh i le t he r emarkab le a dap ta t ion o fd romeda ry c ame ls t o h o t /dry e nv i ronmen ts i sw ide ly r ecogn ized , t he h i ghe r mo r ta l i ty a nd l owe r r ep roduc t ion r a tes e xpec ted among d romeda r ies i n we t /wa rm a nd w e t /co ld e nv i ronmen ts r e la t ive t oc a t t le , s heep a nd g oa ts s eems e qua l ly s t r ik ing .

E co logy , D is t r ibu t ion a nd E a r ly E xp lo i ta t ion o fW i ld Whea ts a nd B a r leys I n t roduc t ion C y togene t ica l ly , d omes t ica ted whea ts a nd b a r leys b e long t o t he s ame s pec ies a s t he i r w i ld c oun terpa r ts , t he p r imary d i s t inc t ion b e tween t hese r a ces b e ing a h i gh f r equency o fs eed s ha t te r ing t r a i ts i nw i ld p opu la t ions a nd ah i gh f r equency o fn onsha t te r ing t r a i ts i nd omes t ica ted p opu la t ions . "W i ld " w hea ts a nd b a r leys f r ee ly i n terb reed w i th " domes t ica ted " whea ts a nd b a r leys , e xh ib i t p er fec t c h romosome p a i r ing i nt he F 1 g ene ra t ion , a nd p roduce f u l ly f e r t i le h yb r ids . I n t he f o l low ing d i scuss ion , c ommon ly u sed e p i the ts a nd g ener ic n ames w i l l b e emp loyed , e ven t hough t hey i mp ly g rea te r d i f fe rences b e tween t axa t han t he b i o log ica l e v idence c an s uppo r t ( Ha r lan 1 977 :358 -358 ; H a r lan a nd Z oha ry 1 966 :1075 ; H a r lan e ta l . 1 972 ; K i s lev 1 984 :63 ; Z ohary 1 969 ; Z oha ry e ta l . 1 969 ) .

Mode rn E co logy a nd D is t r ibu t ion I n t he i r n a tura l h ab i ta ts , w i ld e inko rn whea ts ( T r i t icum b oeo t icum ) a re e daph ica l ly a ssoc ia ted w i th b asa l t ic s o i ls , ma r ls a nd l i mes tones , a nd a re s a id t o b e mo re mesophy t ic a nd t o le ran t o fc o ld t han w i ld b a r leys . T hey a re p resen t ly f ound i ns t ands r ang ing f r om 0 -2 ,000 m i n e l eva t ion . W i ld emme rs ( T . d icocco ides ) a re s l igh t ly mo re r es t r ic ted i nt he i r d i s t r ibu t ion a nd e co logy t han e inko rn , f o r wh i le t hey t oo s how c l ose a f f in i t ies w i th b asa l t ic a nd h a rd l i mes tone s l opes , t hey a re r a re o rc omp le te ly a bsen to n ma r l s o i ls . L i ke e i nko rn , emme ri sd i s t r ibu ted o ve r aw ide a t t i tud ina l amp l i tude ,w i th s t ands e x is t ing f r om 1 00 m b e low s ea l e ve l i nt he J o rdan V a l ley t h rough 1 ,500 -1 ,600 m a bove s ea l e ve l . W i ld b a r leys ( Hordeu m s pon taneum ) a re mo re x e rophy t ic b u t l e ss c o ld to le ran t t han w i ld whea ts , a nd a re r a re ly f ound a bove 1 ,500 m i n e l eva t ion . Wh i le t hey o ccu r i nd ense m ixed s t ands c on ta in ing e i nko rn a nd /o r emme r o n t he h eavy s o i ls a ssoc ia ted w i th h a rd l i mes tone a nd b asa l t ic b ed rocks , w i ld b a r leys a re mo re w ide ly d i s t r ibu ted t han w i ld whea ts , w i th s tands e x tend ing i n to wa rm s teppe a nd d eser t r eg ions , f r equen t ly o ccu r r ing a l ong d ry wa te r c ou rses . I nt he i r p resen t d i s t r ibu t ion , mass ive s t ands o f r obus t a nd l a rge -seeded r aces o f w i ld e inko rn o ccur o n t he moun ta inous s l opes o f s ou theas te rn T urkey , n or the rn I r aq a nd w es te rn I r an . S im i la r ly r obus t a nd l a rge -seeded r a ces o f w i ld emme r a nd b a r ley o ccur i n mass ive s tands i nt he u ppe r J o rdan V a l ley o n t he b asa l t ic p l a teaus a nd l i mes tone s l opes o f E as tern Ga l i lee , G i lead a nd t he Go lan . T hese d ense s t ands a re g ene ra l ly r es t r ic ted t o r eg ions r ece iv ing 4 00 1 ,000 mm o f a nnua l r a in fa l l ( Ha r lan 1 967 ; H a r lan a nd Z oha ry 1 966 ; Z oha ry 1 969 ) .

P erspec t ives o n t he E ar ly D is t r ibu t ion a nd E xp lo i ta t ion o f Whea ts a nd B ar leys Wh i le t he p reced ing d a ta r e la te t o t he k nown a t t r ibu tes a nd d i s t r ibu t ions o fw i ld w hea ts a nd b a r leys i n t he p resen t , t he i r P le is tocene p as t r ema ins s pecu la t ive . A rgu ing f r om a bundance a nd s eem ing ly i mbued w i th t he b e l ie f t ha t c e rea l c u l t iva t ion n ecessa r i ly r ep resen ts 6 4

at r emendous a dvance ( r ead " p rog ress " ) i nh uman e x is tence , s eve ra l s cho la rs h ave a t temp ted t o a ccoun t f o r t he a ppa ren t f a i lure o f h umans t o e xp lo i t w i ld c e rea l s t ands o r t o a dop t c e rea l c u l t iva t ion s t ra teg ies i nt he N ea r E as t p r io r t o t he E p ipa leo l i th ic a nd N eo l i th ic p e r iods b y p os tu la t ing v a r ious s cena r ios c once rn ing t he P a leo l i th ic n a tu re a nd d i s t r ibu t ion o ft hese p l an ts . F o r e xamp le , i th as b een s ugges ted t ha tw i ld whea ts a nd b a r leys s imp ly d i d n o t e x is t i na nnua l f o rms p r ior t o a pp rox ima te ly 1 ,000 B .P . ( Why te 1 977 ) , t ha t t hey e vo lved i nN or th A f r ica a nd d id n o t s p read i n to t he N ea r E as t u n t i l t he e nd o f t he P le is tocene ( Ho le 1 984 :50 ; Wr igh t 1 977 :296 ) , o r t ha t t hey o n ly e x is ted i nt he N ea r E as t i n sma l l , u np red ic tab le ( r ead " l ess d es i rab le " ) r e fug ia s t ands a tl owe r e l eva t ions a nd d i d n o t b ecome a bundan t a nd p red ic tab le ( r ead " mo re -des i rab le " ) u n t i l t he i r e xpans ion i n to h i ghe r e l eva t ions a t t he c l ose o f t he P le is tocene ( Hen ry 1 983 : 150 ; H en ry e ta l . 1 98 1 :54 ; Wr igh t 1 968 ) . Wh i le e a r ly d a tes f o r t he e xp lo i ta t ion o f b a r ley i n E gyp t c an n o l o nge r b e ma in ta ined ( Wendo r f a nd S ch i ld 1 980b ; Wendo r f e ta l . 1 984 ) , t he d i scove ry o f emmer i nK eba ran l e ve ls a t N aha l O ren ( Noy e ta l . 1 973 ) a nd b a r ley i nK eba ran l e ve ls a tE in Gev I( Ba r -Yose f 1 970 ; H en ry 1 983 ) s ugges ts amuch e a r l ie rd a te f o rt he e vo lu t ion o fa nnua l c e rea ls i nt he N ea r E as t ( p r ior t o 1 5 ,000 B .P . ) t han a lowed f o r i n Why te 's s cena r io . F ur the r , w h i le a n e l eva t iona l s h i f t i nc erea l p opu la t ions c ou ld a ccoun tf o rv a r ia t ions i nt he r eg iona l d i s t r ibu t ion o fc e rea l s t ands , i ti su nc lea r w hy s uch s h i f ts wou ld make s tands mo re o r l e ss p red ic tab le . F ur the r , b o th t heo re t ica l c ons idera t ions a nd emp i r ica l d a ta s ugges t t ha t t he re i sn o n ecessa ry c o r re la t ion b e tween t he a bundance o f a n i n d iv idua l f ood i t em a nd i t s d es i rab i l i ty a s af ood r esou rce ( Cha rnov 1 976b ; C ha rnov a nd O r ians 1 973 ; Em len 1 966 ; H awkes a nd O 'Conne l l 1 98 1 ; H awkes e ta l . 1 982 ; H i l l 1 983 ; H i l l e ta l . 1 986 ; K rebs 1 978 ; MacA r thu r a nd P ianka 1 966 ; O 'Conne l l a nd H awkes 1 981 , 1 984 ; P yke e ta l . 1 977 ; S imms 1 984 , 1 987 ) . F i na l ly , i ts eems f a r mo re p robab le t ha t whea ts a nd b a r leys e vo lved i nt he N ea r E as t , r ega rd less o fw he the r s im i la r s t ands e x is ted i nP le is tocene N o r th A f r ica . H a r lan a nd Z oha ry ( 1966 :1079 ) h ad p rev ious ly s ugges ted t ha t t he d ense s t ands o f emme r a nd b a r ley n ow f ound i nt he u ppe r J o rdan V a l ley o nce e x tended s ou th a nd e as t o n to t he l i mes tone a nd b asa l t r eg ions o ft he E s -Sa l t P la teau o fn o r the rn J o rdan a nd t he J ebe l D ruze o f s ou the rn S y r ia . H oweve r , t hese l i mes tone a nd b asa l t e nv i ronmen ts i n wh ich whea ts a nd b ar leys a ppa ren t ly e vo lved c on t inue a c ross s ou the rn S y r ia a nd n or the rn J ordan i n to n or thwes te rn S a 'ud i A rab ia . I tt he re fo re s eems p robab le t ha t a nces t ra l w hea ts a nd b a r leys w ou ld h ave b een p resen t among t he a nnua l g rass c omponen t o ft he s avanna g rass lands wh ich f o rme r ly s t re tched a c ross t he A rab ian P en insu la , a nd w e re e i the r r educed t o , o r a ch ieved t he i r p resen t , mo re r e s t r ic ted d i s t r ibu t ion a tt he c l ose o ft he P le is tocene . F ur the r , t he i n te rp re ta t ion o f al a te e xp lo i ta t ion o fc e rea ls a s b ased u pon t he a bsence o f s t one m i l l ing e qu ipmen t o n s eve ra l P a leo l i th ic a nd E p ipa leo l i th ic s i tes may a c tua l ly b e i l f ounded . I n i t ia l ly , l i mes tone a nd b asa l t " g r inds tones , " " pounde rs , " a nd mo r ta rs h ave b een f ound d u r ing r ecen t s urveys o n b o th U ppe r P a leo l i th ic a nd E p ipa leo l i th ic s i tes i n wha t a re n ow t he d eser ts o f n o r theas te rn J o rdan ( Ga r ra rd e ta l . 1 975 -77 , 1 985 ) . T hese i n c lude s i tes i nt he Wad i e l -J i la t ( Dhoba i ) whe re e a r l ie ra rchaeo log ica l s ur veys a nd e xcava t ions f a i led t or epo r ts uch i t e ms ( Waech te r a nd S e ton -W i l l iams 1 938 ) . H eav i ly c ura ted a ndes i te a nd me tamo rphosed s ands tone g r ind ing s l abs may a l so b e s een t h roughou tt he l e ve ls o ft he K eba ran r ockshe l te r i n Wad i Madamagh ( Denn is L .H eske l , p e rsona l c ommun ica t ion 1 985 ; A lan S . L i ch ty , p e rsona l c ommun ica t ion 1 978 ; p e rsona l o bse rva t ion 1 986 ) , a l though t hese we re u n repor ted b y t he e xcava to ro ft he s i te ( K i rkb r ide 1 958 ) . T he e x tens ive e thnog raph ic a nd h i s to r ic u se o f wooden mor ta rs a nd p es t les f o r t h resh ing a nd p ound ing c e rea ls i n to g roa ts ( Ha r lan 1 967 :199 -200 ; H e lbaek 1 969 :401-402 ; Mus i l 1 928 :91; N ico la isen 1 963 :235 ; R icha rds 1 951 :104 ; S e r jean t 1 974 :44 ; S . Sm i th 1 980 :472 ) a nd ab i as t od epos i t ion o n l a te r s i tes c rea ted b yt he c ura t ion a nd r euse o fe a r l ie rg r ind ing s t ones ( S imms 1 985 ) f u r the rd im in ish t he a bsence o fg round s t one i n a rchaeo log ica l c on tex ts a s as i gn i f ican t i n d ica t ion t ha t c e rea ls w e re e i the r n o t b e ing e xp lo i ted , o rw e ren ' t a va i lab le f o re xp lo i ta t ion: I nt he p resen t a na lys is , i tw i l l b e a ssumed t ha t s t ands o fa nnua l whea ts a nd b a r leys w ere a va i lab le f o r e xp lo i ta t ion b y t he e a r ly E p ipa leo l i th ic ( app rox ima te ly 1 8 -10 ,000 B .C . ) , i fn o t e a r l ie r . A s s uch , i tw i l l a l so b e a ssumed t ha t c e rea l c u l t iva t ion was a s ubs is tence o p t ion b y a t l e as t t h is p er iod o ft ime . H ence , f ac to rs r e spons ib le f o r t he i n c lus ion o r e xc lus ion o fc e rea ls i n t he d i e ts o fp reh is to r ic f o rage rs , a nd f o rt he e vo lu t ion a nd a dop t ion o fc e rea l c u l t iva t ion , w i l l b e s ough t i nt he r e la t ive e ne rge t ics o fa l te rna t ive d i e ta ry a nd s ubs is tence o p t ions u nde rc hang ing e nv i ronmen ta l c ond i t ions d u r ing t he t e rm ina l P le is tocene a nd t he H o locene e poch . 6 5

E co logy o fC u l t iva ted Whea ts a nd B a r leys I n t roduc t ion I nt he N ea r E as t a nd N or th A f r ica , d ry fa rm ing , a nd f r equen t ly i r iga t ion s t ra teg ies , h ave p redom inan t ly i n vo lved a w in te r c e rea l monocu l ture b ased u pon a f a l low /whea t o r b a r ley r o ta t ion ( de B r ichambau t a nd Wa l len 1 963 :6 ; S impson 1 981:27 ) . A s n o ted i n C hap te r 3 , w hea ts a nd b a r leys h ave i n he r i ted a n a nnua l r ep roduc t ive s t ra tegy o r ien ted t owa rds a ch iev ing mass r ep resen ta t ion a nd s eed p roduc t ion w h i ls t f avo rab le c ond i t ions i n t he i r h ab i ta t l a s t . A ssoc ia ted w i th t h is r ep roduc t ive s t ra tegy i sp heno typ ic f l ex ib i l i ty a low ing f o r aw ide r ange o f p roduc t ive p o ten t ia l r e la t ive t os pec i f ic l i m i t ing f ac to rs i nt h e i rt empo ra l h ab i ta ts .

Wa ter a nd N u t r ien ts F o r whea ts a nd b a r leys c u l t iva ted i nf avo rab le e nv i ronmen ts , a s w i th w i ld g rasses a nd r ange lands i ng ene ra l , wa te r a nd s o i l n u t r ien ts r ep resen t t he p r ima ry l i m i t ing f ac to rs t o p l an t g row th a nd p roduc t iv i ty ( B reman a nd d e W i t 1 983 ; G u ise 1 969 ; H a l l e t a l . 1 979 ; H i l lman 1 973 :230 ; S impson 1 981; Web ley 1 972 :173 ) . U nde r c ond i t ions o f s easona l ly r es t r ic ted r a in fa l l ,w a te rs t ress i sp a r t icu la r ly s i gn i f ican t i nr educ ing c r op y i e lds a nd r ange land p roduc t iv i ty . Wa te r s t ress i st he p r inc ip le g row th inh ib i t ing f ac to r among g rasses i nr eg ions r e ce iv ing l e ss t han a pp rox ima te ly 2 00 -300 mm o f a nnua l r a in fa l l ( B reman a nd d e W i t 1 983 :1342 ) , w i th r egu la r d ry fa rm ing i ns uch a reas f r equen t ly c ons ide red i mposs ib le o r e x t reme ly h aza rdous w i thou t i r iga t ion ( de B r ichambau t a nd Wa l len 1 963 :4 , 9 10 , 2 9 -33 ; Maye rson 1 960 :19 ; S impson 1 981 :27 ) . A s n o ted e a r l ie r , d ense s t ands o f w i ld w hea ts a nd b a r leys a re g ene ra l ly r es t r ic ted t o b asa l t a nd l i mes tone h ab i ta ts r ece iv ing 4 00 1 ,000 mm o fa nnua l r a in fa l l ( Zoha ry 1 969 :55 ) . A bove t he 2 00 -300 mm t h resho ld , n u t r ien ts ( pa r t icu la r ly n i t rogen a nd p hospho rus ) i n c reas ing ly b ecome t he p r inc ip le g row th inh ib i t ing f ac to rs ( B reman a nd d e W i t 1 983 :1342 ) . I ng ene ra l , b a r leys a re mo re s a l t to le ran t t han w hea ts ( G ibson 1 974 :10 ; J acobsen a nd A dams 1 958 : 1252 ; S impson 1 981 :27 ) i na dd i t ion t o b e ing mo re x e rophy t ic , a nd t he re fo re d o b e t te r a nd a re mo re f r equen t ly c u l t iva ted t han w hea ts i n a reas o f l owes t r a in fa l l a nd o n s em isa l ine s o i ls ( de B r ichambau t a nd Wa l len 1 963 :40 ; Maye rson 1 960 :17 ; E I -Sama r ra ie 1 972 :85 ; S impson 1 981 :27 ) . A s w i l l b e d i scussed i nC hap te r 6 ,b a r leys h ave a c a lo r ic c on ten t e qua l t o whea ts , b u t o f ten o u tp roduce t hem i ns eed y i e ld p e ru n i t a rea w i th l e ss l a bo r r equ i red , f u r the ra dd ing t ot he i rd es i rab i l i ty a s ac e rea l c rop r e la t ive t ow hea ts .

S o i ls a nd H yd ro logy o fE a r ly C erea lC u l t iva t ion Wh i le b o th whea t a nd b a r ley a re h i gh ly p roduc t ive o n h eav ie r c l ay s o i ls w i th g ood wa te r h o ld ing c apac i t ies , t he p r ima ry f ocus o fe a r ly c e rea l c u l t iva t ion i nt he N ea r E as t , N or th A f r ica , C en t ra l A s ia a nd E u rope was o n l i gh t s o i ls w i th l owe rw a te r re ta in ing c apac i t ies ( C la rk 1 968 :97 ; S her ra t 1 980 :315 , 1 983 :98 ; S impson 1 981:27 ; Web ley 1 972 :171, 1 73 ) . T h is i n i t ia l a ssoc ia t ion may b e p a r t ia l ly u nde rs tood b y r e fe rence t oe a r ly t i lage t echno logy a nd t he l a bo r r equ i red f o r s eedbed p repa ra t ion , s i nce l i gh t s o i ls wou ld b e i n he ren t ly e as ie r t o c u l t iva te w i th h and t oo ls t han t he d ense rc l ay s o i ls ( She r ra t 1 980 :321 , 1 98 1:293 ; S impson 1 981 :27 ; Web ley 1 972 :17 1 , 1 73 ) . F ur the r , t he f r equen t a ssoc ia t ion o f c l ay s o i ls w i th f o res t g row th wou ld p resumab ly h ave made t he e xp lo i ta t ion o fs uch l a nds l e ss a t t rac t ive t han t hose w i th l i gh te r s o i ls u nde r c ond i t ions o fl i t h ic t echno logy d ue t ot he h i ghe r l a bo rc os ts e xpec ted f o r i n t ia l ly c l ea r ing ( I ve rsen 1 956 ; K ap lan 1 985 : 12 ) a nd s ubsequen t ly b reak ing d ense v i rg in c l ay s o i ls T he l o ca t ions w he re l i gh t s a i ls w e re e xp lo i ted f o r e a r ly c e rea l c u l t iva t ion we re a lmos t u n ive rsa l ly a ssoc ia ted w i th l o ca l ized s u r face w a te ro rh i gh g roundwa te r , r esu l t ing f r om p e renn ia l s p r ing f l ows , s pa te r uno f f o n a l luv ia l f ans , c oncen t ra ted r uno f f i n c a tchmen ts , a nd f l ood r ecess ion e nv i ronmen ts a round p onds , ma rshes , l a kes a nd s t reams ( Moo re 1 985 :15 16 ; S he r ra t 1 980 ) , e f fec t ive ly p rov id ing i r iga t ion c ond i t ions w i th l i t le o r n o a ssoc ia ted l abo r i n ves tmen t . T ab le 2s umma r izes s ome o ft he g ene ra l h yd ro log ica l a ssoc ia t ions o fv a r ious s i tes w he re e a r ly c erea l c u l t iva t ion i sa t tes ted o r s t rong ly s uspec ted . T he p robab le s i gn i f icance o fa c o r re la t ion b e tween e a r ly c e rea l c u l t iva t ion a nd r eg iona l ly c i rcumsc r ibed h yd ro log ica l e nv i ron -

6 6

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P e renn ia l s p r ing o n a luv ia lf an B o th o n E uph ra tes f l oodp la in a nd s easona lw ad is N ex tt o ma r sh / lake

9 -8 th

D o r re l l 1 978

9 -8 th 9 th 8 th

C oncen t ra ted r uno f f i n wad i M ic roembaymen t o n p e renn ia l s t ream C oncen t ra ted r unof i n wad i

7 -6 th 7 -6 th

S he r ra t 1 980 :315 Moo re 1 982 :229 V an Z e is t a nd B akke r H ee res 1 979 :168 R a ikes 1 966 R o l le fson 1 984 :3 , 1 2

6 th?

R o l le fson a nd K a fa f i 1 985 :63 -64

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B ra idwood a nd H owe 1 960 :26

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A l lt h ree n ex tt o i n te rmon tane l a ke b as ins O n a luv ia lf an o fS e le rek i R ive r A tw iden ing o fr i ve rv a l ley f l oodp la in On a luv ia lf an o fC a rsamba R ive r

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P ayne 1 972b :194

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A l l n ex tt o s easona l ly f l ooded n a tu ra l d ep ress ions

7 -6 th 6 -5 th 6 -4 th

H o le e ta l . 1 969 :21-22 "

E as tern S aha ra S i tes

E -75 -6 E -75 -8 E -77 -5 /5A

N ab ta P l aya N ab ta P l aya E lK o r te in P laya

7 -6 th 6 -5 th 6 -5 th

Wendo r f a nd S ch i ld 1 980a :84 -9

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A l l o n e dges o fa luv ia lf ans

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men ts i nt he e vo lu t ion o fa g r icu l tu ra l s t ra teg ies a nd i n c reas ing ly c omp lex s oc ia l s t ruc tu res w i l l b e d i scussed i ns ubsequen t s ec t ions .

V a r ia t ions i nt he A nnua lR e l iab i l i ty o fS pec i f ic Wa ter S ou rces a nd t he Geog raph ic C i rcumsc r ip t ion o f Op t ima lC e rea lP roduc ing L ands F o ra reas i n wh ich e a r ly c e rea l c u l t iva t ion was d ependen t u pon d i rec t r a in fa l l , s pa te r u no f f , a nd c oncen t ra ted r unof i n sma l l c a tchmen ts , v a r ia t ions i na nnua l l o ca l r a in fa l l may b e e xpec ted t o h ave d i rec t ly a fec ted a nnua l c e rea l y i e lds . E ven w hen l ong te rm t r ends i n r eg iona l p rec ip i ta t ion a re e xc luded , l o ca l v a r ia t ions i na nnua l r a in fa l l a re q u i te p ronounced . F o re xamp le , o ne o f t he l o nges t , c on t inuous ly r eco rded r a in fa l l r eco rds a va i lab le i nt he wo r ld i st ha t o f J e rusa lem , whe re measu remen ts h ave b een made s i nce 1 851 ( U . S . D epa r tmen to fC omme rce 1 966 :193 196 ) . F i gure 3 d iag rams t he d ev ia t ion i nt o ta l a nnua l r a in fa l l a tJ e rusa lem f r om t he mean f o rt he 1 08 y ea rp e r iod f r om 1 851 t h rough 1 960 ( t wo y ea rs m iss ing ) . A s c an b e s een , t he amoun t o fv a r ia t ion i nt o ta l p rec ip i ta t ion f r om y ea rt oy ea rc an b e q u i te e x t reme , w i th r e la t ive ly b r ie f c l us ters o f h i gh a nd l ow r a in fa l l y ea rs b e ing r ead i ly a ppa ren t . H owever , t he p e r iod f r om 1 922 t h rough 1 935 h ad c ons is ten t ly b e low -ave rage r a in fa l l , r e f lec t ing t he m ino r w o r ld -w ide c l i ma t ic f l uc tua t ion wh ich i n c luded t he M idwes t d rough t a nd " Dus t -bow l " o f t he U .S . T he s uccess o r f a i lu re o f y ea r t o y ea r c e rea l c u l t iva t ion u nde r t hese c ond i t ions wou ld f r equen t ly d epend u pon v a r ia t ions i nb o th t he t im ing a nd q uan t i ty o fa nnua l p rec ip i ta t ion o n ar e la t ive ly l o ca l s ca le . F o r a reas i n wh ich e a r ly c e rea lc u l t iva t ion was d ependen t u pon n a tu ra l i r iga t ion f r om f l ood r ecess ion , d i scha rge v a r ia t ions i nt he d ra inage s ys tems s upp ly ing wa te r may b e e xpec ted t o h ave a fec ted t he amoun t o fa rea i n unda ted , a nd t hus t he t o ta l a rea a va i lab le f o rc u l t iva t ion . F o r e xamp le , o ne o ft he l o nges t , c on t inuous ly r eco rded d i scha rge r eco rds f o ra ny r i ve r i nt he w o r ld i st ha t o ft he N i le , whe re measu remen ts a tA swan h ave b een made s i nce 1 871 ( C lawson e ta l . 1 97 1 :209 -210 ) . F i gu re 4 d iag rams a nnua l d ev ia t ions i nt he t o ta l d i scha rge o ft he N i le f r om t he mean d i scha rge f o r t he 9 5 y ea r p e r iod f r om 1 871 t h rough 1 960 . A s w i th t he r a in fa l l r eco rd a t J e rusa lem , v a r ia t ions i nt he t o ta l d i scha rge o ft he N i le h ave o f ten b een q u i te e x t reme f r om y ea r t o y ea r , a l though c l us ters o f h i gh a nd l ow d i scha rges s eem mo re p ronounced t han s im i la r c l us te rs i na nnua l r a in fa l l . H oweve r , i n af l ood recess ion e nv i ronmen t , t he p r inc ip le e f fec t o f s uch v a r ia t ions wou ld b e t oa l te rt he t o ta l a rea a va i lab le f o rc u l t iva t ion a s af unc t ion o ft he t o ta l a rea i n unda ted . F i e lds a tl owe r l e ve ls n ea r t he e dge o ft he N i le wou ld n o rma l ly b e i n unda ted e ven w i th l ow f l oods , a nd o n ly t hose f i e lds a t h i ghe r l e ve ls f u r the r f r om t he N i le w ou ld b e a dve rse ly a fec ted ( Hu rs t a nd P h i l l ips 1 93 1 :2 1) . T ha t a ma jo r s h i f t o ccu r red i nt he c l ima t ic r eg ime o ft he N i le j u s t b e fo re t he t u rn o ft he c en tury h as l o ng b een r ecogn ized ( Hurs t 1 936 : 1038 ; R ieh l e ta l . 1 979 :1548 ) , a n e ven t w h ich t empo ra l ly c o r re la tes w i th s ecu la r r a in fa l l c hanges t h roughou t mos t o f t he t r op ics . A s s umma r ized b y K raus ( 1955 : 198 ) : T he a ve rage a nnua l v o lume o fp rec ip i ta t ion d ec reased r a the r s udden ly a tt he t u rn o ft he c en tury . T h is c hange , wh ich o ccu r red af ew y ea rs l a te rt han a th i ghe r l a t i tude , b rough t d rough t -o ra tl e as t d ry c ond i t ions -t ot r op ica l A us t ra l ia , p a r ts o ft he P ac i f ic , C o lomb ia , Mex ico , NE . B raz i l , t he C a r ibbean , A f r ica , SW . A s ia a nd C ey lon . As im i la r s h i f t o ccur red i nt he C o lo rado R ive r s ys tem a round 1 930 ( R ieh l e t a l . 1 979 :1549 ) . H owever , t he p r inc ip le e f fec t o fs uch s h i f ts i nr a in fa l l r eg imes wou ld a ga in b e t o a l te rt he t o ta l a rea i n unda ted b y n a tura lf l ood ing . F o r a reas i n wh ich e a r ly c e rea l c u l t iva t ion was d ependen t u pon p e renn ia l s p r ing f l ows , a nnua l a nd s easona l f l uc tua t ions i nf l ow c ond i t ions may b e e xpec ted t o h ave a f fec ted t he amoun t o fa rea r ece iv ing wa te r a nd t hus t he t o ta l a rea a va i lab le f o r c u l t iva t ion . U n for tuna te ly , l o ng te rm f l ow r eco rds f o r p e renn ia l s p r ings a re n o t a va i lab le . F u r the r , i nt he a rchaeo log ica l r eco rds o fe a r ly c e rea l c u l t iva t ion i nt he N ea r E as t a nd N o r th A f r ica , t he s i te o fJ e r icho w i th i t s s p r ing o f ' A in e s -Su l tan a ppa ren t ly r ep resen ts t he o n ly k nown l o ca t ion whe re e a r ly c e rea l c u l t iva t ion w ou ld h ave b een s pec i f ica l ly t i ed t o ap e renn ia l s p r ing f l ow . Wh i le i ts eems u n l ike ly

6 8

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Y EAR F i gu re 4 . A nnua l d ev ia t ion i nt o ta l N i le d i scha rge a tA swan f r om t he c umu la t ive mean f o rt he y ea rs 1 87 1 t h rough 1 965 . T h is r eco rd c ove rs ap e r iod o f9 5 y ea rs . 6 9

t ha t J e r icho was n ecessa r i ly u n ique i nt h is a ssoc ia t ion , t he re a re s eve ra l e co log ica l f ac to rs w h ich w ou ld make s uch a g r icu l tu ra l ly f avo rab le m ic roenv i ronmen ts r a re i na ny l a ndscape . I n i t ia l ly , w h i le n ume rous p e renn ia l s p r ings e x is t a l ong t he r i f t o f t he J o rdan V a l ley , t he i r s easona l f l ow c ha rac te r is t ics a re l a rge ly d e te rm ined b y t he n a tu re o ft he a qu i fe rs w h ich f eed t hem ( l on ides 1 939 :249 -25 1) . I n p ar t icu la r , i th as b een n o ted t ha t s p r ings i n t he u ppe r o r n or the rn J o rdan V a l ley a re f ed b y a qu i fe rs i nS enon ian s t ra ta , a nd g ene ra l ly e xh ib i t ac ons tan t f l ow l a rge ly u na f fec ted b y r a in fa l l . I nt he c en t ra l J o rdan V a l ley , s p r ings a re o f ten f ed b y a qu i fe rs i nC enoman ian s t ra ta , a nd e xh ib i tw ide v a r ia t ions i ns easona lf l ow s t rong ly i n f luenced b y r a in fa l l ( e .g . , ' A in A u ja ; B lake a nd Go ldschm id t 1 947 :405 -406 ) . S ou th o ft h is r eg ion t h rough t he Wad i A raba , s p r ings a r ise i na l o ft he e xposed s t ra ta , w i th s ome e xh ib i t ing c ons tan tf l ows a nd o the rs h i gh ly v a r iab le f l ows . ' A in e s -Su l tan , l i ke t he n ea rby ' A in O ut , i sa ppa ren t ly f ed b y ar eservo i r a qu i fe r r u nn ing t h rough d o lom i te a nd l i mes tone T uron ian -Cenoman ian s t ra ta . T h is r ese rvo i r i sc ha rged b y r a in fa l l i nt he u ppe r wa te rshed o ft he J udaean H i l ls , a nd t he s p r ings a ssoc ia ted w i th i teme rge u nde r p ressu re ( B lake a nd Go ldschm id t 1 947 :405 ; D o r re l l 1 978 :11) . T he c omb ined d i scha rge o f' A in e s -Su l tan a nd t he O ut s p r ings i nt he h i s to r ic p resen t a ve rages 0 .7 m3/ sec i n aw e t y ea r a nd 0 .4 m3/ sec i n ad ry y ea r , w i th a n a nnua l a ve rage o f0 . 5 m3/ sec ( Gove rnmen t o fP a les t ine 1 947 :10 ) . A l though t he f l ow a t' A in e s -Su l tan i n c reases w i th r a in fa l l i nt he J udaean H i l ls w i th in o n ly o ne o r t wo d ays t o a max imum o f 0 .37 m3 / sec a f te r h eavy r a ins , i t s f l ow i ss t i l l c .0 .26 m3/ sec a tt he e nd o ft he d ry s eason , s ugges t ing t ha t max imum f l ow c ond i t ions r esu l t f r om t he o ver topp ing o f i t s r eservo i r a qu i fe r f r om wh ich a r e la t ive ly c ons tan t b ase f low i sn o rma l ly ma in ta ined ( Dor re l l 1 978 :11) . A cco rd ing t o D o r re l l , b o th t he s i ze o ft he a qu i fe r i n vo lved a nd t he s ou rce o fi t sc ha rge f r om r a in fa l l i nt he u ppe re l eva t ions o ft h e J udaean H i l ls d im in ish t h e e fec ts o fs ma l ls ca le c l ima t ic v a r ia t ions o n s p r ing f l ow . A no the r f ac to r p resumab ly s i gn i f ican t i nt he e a r ly a g r icu l tu ra l u t i l i ty o f p e renn ia l s p r ings wou ld b e t he way i n wh ich t hey eme rged a nd d i spe rsed . Mos t s p r ings a l ong t he J o rdan R i f t , s uch a s ' A in O l t , eme rge i n wa te r c ourses o ccas iona l ly o r p e r iod ica l ly s ub jec t t o h i gh r uno f f f r om v i o len t d ownpours i nt he s ur round ing h i l ls , a nd t hus f l ow i nd eep ly i n c ised b eds when t hey eme rge o n to f oo th i l l f ans ( Do r re l l 1 978 : 11) . T he f l ow o f' A in e s -Su l tan , h oweve r , d oes n o t eme rge i n s uch a d ra inage a nd i sn o t r e in fo rced b y h i gh r uno f f . Emerg ing o n to a n e ven ly s l op ing f an w i th f e r t i le , n onsa l ine s o i ls w i th al o ca l g rad ien to f5 0 mo ve r 1 . 5 km ( =0 . 03 m /m , o r 1 :30 ) , i ti sp robab le t ha t t he r esu l t ing p reh is to r ic s t ream w as b ra ided , w i th r e la t ive ly s ha l low c hanne ls s p l i t t ing a nd r e jo in ing d owns lope ( Do r re l l 1 978 : 12 ) . S ince w in te rc e rea ls i nt he J o rdan V a l ley h ave a n e s tab l ished wa te rd u ty o fa pp rox ima te ly 4 m3 / du /day ( l on ides 1 939 :xx i i -xx i i i ) , t he mode rn f l ow f r om ' A in e s -Su l tan c ou ld m in ima l ly i r iga te 1 , 170 d u o f l a nd d u r ing a t en h our i r iga t ion d ay a ssum ing a m in imum f l ow o f 0 .26 m 3/ sec w i th h a l f o f t ha t f l ow u sed f o r c u l ina ry p u rposes o r l o s t t h rough e vapo ra t ion ( 0 .13 m 3/ sec x 6 0 s ec = 7 .8 m3 / h r x1 0 h rs = 4 ,680 m3 I 4 m3 p e r d u =1 , 170 d u ) . U n less t he p reh is to r ic f l ow o f ' A in e s -Su l tan was l e ss t han t ha t i nt he h i s to r ic p resen t ( wh ich s eems u n l ike ly ) , a much l a rge r a rea t han t h is c a lcu la ted m in imum c ou ld h ave b een i r iga ted i nt he p reh is to r ic p as t w i th l i t le r equ i red l a bo r . Wh i le t he m ic roenv i ronmen t a tJ er icho may n o t h ave b een t o ta l ly u n ique , i ts eems d oub t fu l t ha t s im i la r c ond i t ions wou ld h ave b een o b ta ined a t many o the rs i tes . I ne va lua t ing e xpec ted v a r ia t ions i ny ea rt oy ea r c e rea l p roduc t iv i ty a s ar esu l t o fa nnua l f l uc tua t ions i ns pec i f ic wa te r s ou rces , i ti sa ppa ren t t ha t t he mos t e x t reme v a r ia t ions wou ld o ccur i nl o ca l i t ies d ependen t u pon d i rec t r a in fa l l o r r uno f f i nr eg iona l ly sma l l c a tchmen ts . I n t hese e nv i ronmen ts , b o th l o ca l v a r ia t ions i nr a in fa l l a nd r eg iona l f l uc tua t ions d ue t o s h i f ts i n c yc lon ic s ys tems wou ld r e su l t i ne x t reme v a r ia t ions i ne xpec ted c e rea l y i e lds . B yc on t ras t , a reas d ependen t u pon r uno f f i ne x tens ive c a tchmen ts , a nd p a r t icu la r ly f l ood r ecess ion e nv i ronmen ts i n vo lv ing ma jo r d ra inage s ys tems , a re e xpec ted t o e xh ib i t l e ss e x t reme v a r ia t ions i n c e rea l p roduc t iv i ty f r om y ea r t o y ea r . I n t hese e nv i ronmen ts , l o ca l v a r ia t ions i nr a in fa l l wou ld o f ten b e o f fse t b y t he r eg iona l e x ten t o ft he t o ta l c a tchmen t , w i th ma jo rf l uc tua t ions p r ima r i ly r es t r ic ted t o s h i f ts i nt o ta l c yc lon ic s ys tems a nd c hanges i nr a in fa l l r eg imes . F ur the r , u nder f l ood -recess ion c ond i t ions , l ow ly ing l a nds wou ld g ene ra l ly b e i n unda ted e ven u nde r c ond i t ions o fl ow f l ood . U n l ike r eg ions d ependen t u pon d i rec t r a in fa l l o r 7 0

r uno f f i n sma l l c a tchmen ts , t he i nab i l i ty o f t he h yd ro log ic s ys tem t o mee t a gr icu l tura l r equ i remen ts w ou ld b e p r ima r i ly r es t r ic ted t o c ompa ra t ive ly i n f requen t s i tua t ions i n vo lv ing a p rec ip i ta t ion f a i lure a c ross e x tens ive r eg ions , a s w i th t he f a i lure o f t he N i le f l ood i n 1 913 ( MacDona ld 1 921 :5 ) . Obv ious ly , t empo ra l v a r ia t ions i nt he s easona l i ty o f f l ood ing b e tween d ra inage s ys tems wou ld h ave made s ome r eg ions mo re a t t rac t ive f o r e a r ly c u l t iva t ion t han o thers . F i na l ly , a reas wa te red b y p e renn ia l s p r ing f l ows f r om r e se rvo i ra qu i fe rs may b e e xpec ted t o e xh ib i t t he l e as t f l uc tua t ion o f a l h yd ro log ic e nv i ronmen ts . P resumab ly , o n ly t he d ep le t ion o f t h e r ese rvo i ra qu i fe rt h rough al o ng te rm r echa rge f a i lu re , o rt ec ton ic a c t iv i ty a l te r ing t he f l ow o f t he s p r ing , w ou ld c ause d rama t ic f l uc tua t ions . S ign i f ican t ly , when t he a rea l e x ten t o f e ach o f t hese h yd ro log ic e nv i ronmen ts i s c ons ide red , a reas d ependen t u pon d i rec t r a in fa l l a nd r uno f f i n sma l l c a tchmen ts a re e xpec ted t or ep resen t t he ma jo r i ty o fa l a va i lab le a rab le l a nd , w h i le r eg ions d ependen t u pon e x tens ive c a tchmen ts a nd f l ood r ecess ion wou ld r ep resen t f a r mo re c i rcumsc r ibed r eg ions . H oweve r , t he m os t c i rcumsc r ibed , l e as t v a r iab le , a nd r a res to fa l a g r icu l tu ra l ly f avo rab le e nv i ronmen ts wou ld b e t hose w he re p e renn ia l s p r ings f l ow f r om r ese rvo i r a qu i fe rs o n to f e r t i le l a nds . S ign i f ican t ly , t he mo re f avo rab le e co log ica l c ond i t ions a ssoc ia ted w i th c i rcumsc r ibed e nv ironmen ts p ossess ing f e r t i le s o i ls a nd n a tura l o re as i ly o b ta ined i r iga t ion w ou ld g ene ra l ly r esu l t i ng rea te r c e rea ly i e lds t han a l b u tt he mos tf avo rab le o fd i rec t r a in fa l l a nd s ma l lc a tchmen t e nv i ronmen ts . Wh i le n o t d i rec t ly a pp l icab le t o t he h yd ro log ica l e nv i ronmen ts d i scussed a bove , t he a na lys is o f S te i t ieh a nd Smad i ( 1974 :19 , T ab le 4 ) o f whea t p roduc t ion b y r eg ion i n mode rn J o rdan e xemp l i f ies t he p ropor t iona l r e la t ionsh ips e xpec ted b e tween a g r icu l tura l z ones w i th v a ry ing h yd ro log ica l q ua l i t ies . T ab le 3 s umma r izes t he i r p ro jec t ions f o r modern whea t p roduc t ion i nJ o rdan a cco rd ing t o r a in fa l l r eg ions , n o t ing t he t o ta l p e rcen tage o fa l a rab le l a nd w h ich e ach r eg ion r ep resen ts . R eg ions e xpec ted t o p roduce 4 0 -60 k g /du o n a pp rox ima te ly 1 50 -300 mm o f r a in fa l l a nnua l ly , wh ich wou ld t echn ica l ly make t hem " a r id z ones " ( de B r ichambau t a nd Wa l len 1 963 :4 ) , r ep resen t a pp rox ima te ly 5 6 .9 % o fa l a va i lab le a rab le l a nd . R eg ions e xpec ted t o p roduce 8 1 k g /du o n 3 00 -400 mm o fa nnua l r a in fa l l , wh ich makes t hem " s em ia r id z ones " ( de B r ichambau t a nd Wa l len 1 963 :4 ) , r ep resen t a no the r3 1 .3 % o fa l a va i lab le a rab le l a nd . B y c on t ras t , e ven when t he a g r icu l tu ra l ly mo re favo rab le u p land r eg ions e xpec ted t o p roduce 1 05 k g /du o n r a in fa l l g rea te r t han 4 00 mm a re i n c luded w i th i r iga ted l ands p roduc ing 1 47 k g /du , t hese " op t ima l " c e rea l p roduc ing z ones wou ld o n ly r ep resen t 1 .8 % o f a lo ft he a rab le l a nd i nJ o rdan . S ince t he t o ta l a rea o fi r iga ted l a nd i nt hese d a ta r e f lec ts mode rn c ond i t ions w i th l a rge -sca le h yd rau l ic p ro jec ts a l ready i np l ace ( e .g . , t he Gho re c ana l s ys tem ) , t h e p reh is to r ic r a t io b e tween " op t ima l " a nd l e ss favo rab le l a nds w ou ld h ave b een e ven mo re p ronounced .

T ab le 3 E s t ima ted P roduc t ion o f Whea t a nd A rea C u l t iva ted i nJ o rdan b y R a in fa l l R eg ions

E NV IRONMENT R a in fa l l R eg ion ( mm ) D ese r t E as te rn P la in Wes te rn P la in U p land G ho re

1 50 -250 2 50 -300 3 00 -400 4 00+ i r iga ted

SO W ING RATES AND Y IELDS ( k g /du ) S ow ing A ve rage R ange o fY ie lds Y ie ld R a te " Bad " -" Good " 4 -5 , 5 -7 8 -15 1 5 1 0 -15

4 0 6 4 8 1 1 05 1 47

a dap ted f r om S te i t ieh a nd Smad i ( 1974 : 19 , T ab le 4 )

7 1

1 0 -70 3 5 -90 5 0 -117 7 5 -146 12 -191

ARABLE AREA I N AN " AVERAGE " YEAR %o fT o ta l D unums A rab le 4 16 ,000 6 93 ,000 6 10 ,000 1 80 ,000 5 0 ,000

2 1.3 3 5 .6 3 1.3 9 .2 2 .6

A l though t he r a in fa l l , c e rea l y i e lds a nd a rea l e x ten t o fs pec i f ic a g r icu l tura l z ones i no the r c oun t r ies wou ld c er ta in ly v a ry f r om t hose o f J o rdan , i ti se xpec ted t ha t s im i la r p ropor t iona l r e la t ionsh ips wou ld s t i l l o b ta in . I ng ene ra l , h i gh ly f avo rab le a g r icu l tura l a reas p ossess ing r i ch s o i ls w i th h i gh r a in fa l l o r n a tura l o r e as i ly o b ta ined i r iga t ion c ond i t ions a re e xpec ted t o r ep resen t ar e la t ive ly sma l l p ropor t ion o f t he t o ta l a va i lab le a rab le l a nd . F o r p urposes o f g ene ra t ing a rchaeo log ica l e xpec ta t ions f r om c ompa r isons b e tween t h e r e la t ive e ne rge t ics o f p as to ra l a nd c e rea l s ubs is tence o p t ions , v a r ia t ions i nw hea t a nd b a r ley y i e lds o n " ave rage " a nd " ma rg ina l " l a nds w i l l b e e s tab l ished i nC hap te r6 .

S umma ry S eve ra l p hy logene t ic v a r ia t ions e x is t b e tween c ame ls , c a t t le , s heep a nd g oa ts wh ich wou ld a f fec t t he i r r e la t ive p roduc t ive p o ten t ia l a nd h ence d es i rab i l i ty a s s ubs is tence h erd a n ima ls i ns pec i f ic e nv i ronmen ts . Ma jo rv a r ia t ions a re o bserved i nt he mechan isms b y wh ich e ach s pec ies ma in ta ins t he rma l s t ab i l i ty a nd wa te r b a lance , a nd i nt he ways i nw h ich e ach o b ta ins f o rage e ne rgy . T he u n ique f ea tu res o fc ame l l o como t ion m us t a l so b e c ons ide red . I n g ene ra l , b y t he i n he r i ted c ha rac te r is t ics o f t he i r l o como t ion , d romedary c ame ls a r e l a rge ly e xc luded f r om e xp lo i t ing w e t e nv i ronmen ts w he re h oo fed c a t t le , s heep a nd g oa ts f a re b e t ter ( Mac fa r lane 1 964 :5 12 ) . C onve rse ly , d ue t ot he i r i n he r i ted w a te r r equ i remen ts , mechan isms o f t he rma l r egu la t ion , a nd f eed ing e co logy , c a t t le , s heep a nd ,t o al e sse re x ten t , g oa ts , a r e l a rge ly e xc luded f r om e xp lo i t ing t he h o t /d ry d ese r t e nv i ronmen ts w he re d romeda ry c ame ls f a re b e t te r . V a r ia t ions i nt he r e la t ive s uscep t ib i l i ty o ft hese s pec ies t o s pec i f ic d i sease f a c to rs wh ich w ou ld a f fec t t he i r r ep roduc t ion a nd mor ta l i ty r a tes a l so a ppea r t o t r ack t hese s ame g enera l e nv i ronmen ta l c ond i t ions . I n s ubsequen t e nerge t ic c ompar isons b e tween t he r e la t ive d es i rab i l i ty o f t hese f ou r h e rd s pec ies f o r e a r ly p as to ra l s ubs is tence , t empo ra l a nd s pa t ia l v a r ia t ions i ng ene ra le nv i ronmen ta lc ond i t ions w i l lb et a ken i n to a ccoun t . When t he e co logy o fw hea ts a nd b a r leys a re e xam ined , i ti sa ppa ren tt ha t a l though b a r ley i s mo re s a l t to le ran t a nd x e rophy t ic t han whea t , b o th r e qu i re s im i la r e nv i ronmen ta l c ond i t ions f o r o p t imum g row th a nd p roduc t iv i ty . I np a r t icu la r , w a te r a nd s o i l n u t r ien ts e x is t a s p r imary l i m i t ing f a c to rs i nt he i rg row th a nd p roduc t iv i ty ,w i th t he l o ose , m o is t a luv ia l s o i ls a ssoc ia ted w i th n a tura l o r e as i ly o b ta ined i r iga t ion ( e .g . , l a ke -edge , r i ve r ine a nd s p r ings ide e nv i ronmen ts , a nd h ab i ta ts w i th c oncen t ra ted r uno f f ) p rov id ing w ha t a ppea r t o b e o p t imum c ond i t ions f o r e a r ly c erea l p roduc t ion s t ra teg ies . S uch e nv i ronmen ts , h oweve r , a re a l so e xpec ted t o b e g eog raph ica l ly c i rcumsc r ibed , a nd w ou ld g ene ra l ly c ons t i tu te o n ly as ma l l p ropo r t ion o ft h e t o ta l a va i lab le a rab le l a nd e ven when r eg ions w i th h i ghe r t han a verage r a in fa l l a re i n c luded . P resumab ly , t h is r e la t ionsh ip w ou ld h ave h ad d emog raph ic a nd s oc ia l c onsequences w h ich , i n t u rn , wou ld h ave a f fec ted t he s pa t ia l a nd t empo ra l e vo lu t ion o r a dop t ion o f c e rea l f ood p roduc t ion s t ra teg ies . T hese i mp l ica t ions w i l l b ef u r the re xp lo red i nC hap te r7 .

7 2

CHAPTER 5

RELAT IVE E NERGET ICS OF P ASTORAL S TRATEG IES I NVOLV ING CAMELS ,CATTLE , SHEEP AND GOATS

I n t roduc t ion I nt h is c hap te r , q uan t i ta t ive d a ta o n t he c a lo r ic b ene f i ts a nd a ssoc ia ted l abor c os ts o f p as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts a re d e ta i led , a nd t he r e la t ive e ne rge t ics o f t hese s t ra teg ies e s tab l ished . I n i t ia l ly , ma thema t ica l mode ls o f p as to ra l c ame l , c a t t le , s heep a nd g oa t h e rd g row th a nd p roduc t iv i ty a re u sed t o e s tab l ish e xpec ted a nnua l c a lo r ic y i e lds f r om h e rds o fs pec i f ic s i zes u nde r mea t -on ly a nd mea t p l us m i lk managemen t s t ra teg ies . T he a nnua l l a bo r c os ts a ssoc ia ted w i th o b ta in ing t hese c a lo r ic b ene f i ts a re s ubsequen t ly e xam ined , f ocuss ing u pon t he b e t te r d ocumen ted t ime a loca t ion d a ta a va i lab le f o rc ame l s t ra teg ies . B ased u pon t hese c os t a nd b ene f i t d a ta , r e la t ive r e turn r a tes f o r mea t -on ly a nd mea t p lus m i lk c ame l , c a t t le , s heep a nd g oa t p as to ra l s t ra teg ies a re c a lcu la ted f o r s ubsequen t c ompa ra t ive a nd p red ic t ive p u rposes .

C ompa ra t ive H e rd G row th a nd P roduc t iv i ty I n t roduc t ion A s d i scussed i nC hap te r4 ,t he c a lcu la t ions o fp as to ra l h e rd g row th a nd a ssoc ia ted a nnua l c a lo r ic p roduc t iv i ty p resen ted i n t h is manusc r ip t a re b ased u pon ma thema t ica l mode ls d eve loped f o r p as to ra l c a t t le h e rds i nA f r ica o fZ ebu ( Bos i nd icus ) a nd Z ebu -de r ived b reeds b y D ah l a nd H jor t ( 1976 :28 -76 , 1 40 181) , h e rds c on ta in ing N ea r E as te rn b reeds o f s heep a nd g oa ts a s g ene ra ted b y R edd ing ( 1981) a nd d romeda ry c ame l h erds a s d eve loped i nt he a ppend ices o ft h is b ook . A s n o ted a tt ha t t ime , t hese mode ls a pp rox ima te t he a ve rage r e su l ts e xpec ted u nde r n o rma l c ond i t ions o f e x tens ive h usband ry , r ea l iz ing t ha t a c tua l h e rd g row th a nd p roduc t iv i ty wou ld v a ry a ccord ing t o e nv i ronmen ta l c ond i t ions a nd t he p roduc t ive c ha rac te r is t ics o fs pec i f ic r e g iona l b reeds .

A nnua lH erd G row th T ab le 4 s umma r izes c ompa ra t ive p as to ra l h e rd g row th d a ta f o rd romeda ry c ame ls , c a t t le , s heep a nd g oa ts a s d e r ived f r om t he mode ls p rev ious ly d i scussed . Wh i le t he e xac t r a tes o f g row th wou ld c e r ta in ly f l uc tua te a cco rd ing t o e nv i ronmen ta l c ond i t ions , p as to ra l c ame l a nd c a t t le h e rds a re g ene ra l ly e xpec ted t o g row a t much s l owe r r a tes t han s heep o r g oa t h e rds , p r ima r i ly d ue t o t he v a r ia t ions i nt he i r r ep roduc t ive t r a i ts d ocumen ted i nC hap ter 4 . I n mo re s ou ther ly l a t i tudes u nde r e x t reme ly f avo rab le e nv i ronmen ta l c ond i t ions a nd mo re f r equen t b reed ing , f l ocks o f s heep c ou ld p o ten t ia l ly g row a t a n a nnua l r a te a s h i gh a s 2 6% , a nd g oa t h e rds a t ar a te a s h i gh a s 4 1% ( Dah l a nd H jo r t 1 976 :97 106 ) . S im i la r ly d rama t ic i n c reases i nt he r a tes o fp as to ra lc ame l a nd c a t t le h e rd g row th u nde r mo re f avo rab le c ond i t ions a re u n l ike ly , w i th o p t im is t ic c a lcu la t ions s ugges t ing a max imum a nnua l c a t t le h e rd g row th o fa round 1 % ( Dah l a nd H jo r t 1 976 :6 1) a nd a max imum a nnua l c ame l h e rd g row th o f a round 8% ( Dah l a nd H jor t 1 976 :83 ) .

A nnua lH e rd C a lo r ic P roduc t iv i ty T ab le 5 s umma r izes c ompa ra t ive p as to ra l p roduc t iv i ty f o r h erds c ons is t ing o f 1 00 t o ta l h ead , a s d e r ived f r om t he s ame mode ls . T he d a ta f o r c ame l h erds r e f lec t t he i r e xpec ted p roduc t iv i ty u nde rc ond i t ions o f n o rma lo f f take , i nw h ich e xcess a do lescen t ma les a nd e l de r ly

T ab le 4 C ompa ra t ive P as to ra l H e rd G row th D ynam ics

DROMEDARY CAMEL 1

CATTLE 2

SHEEP 3

GOAT 3

% o f R ep roduc t ive ly Ma tu re F ema les i n aH e rd

3 9

3 2

5 9

5 0

Y ean ing R a te

0 .39

0 .58

0 .80

A nnua l % I n c rease

4 .5

3 .4

6 .3

1.5

F i n i te R a te o f N a tu ra l I n c rease

1 .045

1 .034

1 .063

1 . 115

I n t r ins ic R a te o f I n c rease

0 .044

0 .033

0 .061

0 . 109

1C a lcu la ted 2F rom

—

1 .20

f r om d a ta p resen ted i nA ppend ix A

D ah l a nd H jor t ( 1976 : 6 4 -5 , 1 57 , 2 83 )

3C a lcu la ted

f r om R edd ing ( 198 1 : 10 , 1 22 , A ppend ix I ,A ppend ix I ) , a ssum ing t ha t af l ock o fs heep w i th 1 00 r ep roduc t ive ly m a tu re e wes w ou ld c on ta in 1 69 t o ta l h ead , a nd t ha t ah e rd o fg oa ts w i th 1 00 r ep roduc t ive ly ma tu re d oes wou ld c on ta in 1 99 t o ta l h ead

T ab le 5 C ompa ra t ive C a lo r ic P roduc t iv i ty f o r P as to ra l H e rds C on ta in ing E xac t ly 1 00 H ead

SOURCE OF CALOR IES

DROMEDARY CAMELS 1 k caVy r %

CATTLE 2

SHEEP 3

k ca l /y r

%

k ca l /y r

GOATS 3 %

k ca l /y r

%

S laugh te r M i lk

3 ,786 ,154 1,273 ,060

2 5 .1 7 4 .9

3 ,420 ,480 5 ,497 ,065

3 8 .4 6 1.6

7 45 ,377 1 ,780 ,189

2 9 .5 3 99 ,679 2 0 .8 7 0 .5 1 ,524 ,987 7 9 .2

TOTAL

1 5 ,059 ,214

1 00

8 ,917 ,545

1 00

2 ,525 ,566

1 00

1 ,924 ,666

1 00

1C a lcu la ted

f r om d a ta p resen ted i nA ppend ix B ,a ssum ing c ond i t ions o fn o rma l m i lk y i e ld , n o rma l o f f take a nd a5 0% u se ö fn a tu ra ld ea ths

2C a lcu la ted

f r om D ah l a nd H jo r t ( 1976 : 1 75 , T ab le 7 -6 ) , e xc lud ing b l eed ing , i n c lud ing a5 0% u se o fn a tu ra l d ea ths

3C a lcu la ted

f r om R edd ing ( 198 1 : 2 33 , T ab le I X -2 ) , a t a7 5% u se o fn a tu ra l d ea ths a nd a ssum ing t ha t af l ock o fs heep w i th 1 00 r ep roduc t ive ly ma tu re e wes wou ld c on ta in 1 69 t o ta l h ead , a nd a h e rd o fg oa ts w i th 1 00 r ep roduc t ive ly ma ture d oes wou ld c on ta in 1 99 t o ta l h ead

7 4

ma les a nd f ema les a re r egu la r ly c u l led , a nd w h ich i n c ludes a5 0% u se o ft he a n ima ls s chedu led t o d ie o f n a tura l c auses ( s ee A ppend ix B ) . T he d a ta f o r s heep a nd g oa t h e rds r ep resen ts e xpec ted p roduc t iv i ty when o n ly o ne b reed ing s eason o ccurs a nnua l ly . G iven t he h i ghe r g row th r a tes men t ioned e a r l ie r , s heep f l ock p roduc t iv i ty may i n c rease b y a s much a s 1 9 .7% a nd g oa t h e rd p roduc t iv i ty b y a s much a s 2 9 .5 % o ve r t he l e ve ls p resen ted i n T ab le 5 u nde r e x t reme ly f avorab le e nv i ronmen ta l c ond i t ions a nd mo re f r equen t b reed ing . H oweve r , a s d ocumen ted i ns ubsequen t c a lcu la t ions , e ven i ft hese i n c reases i ng row th r a tes we re t o r esu l t i ne qu iva len t i n c reases i ne xpec ted p roduc t iv i ty , s uch i n c reases wou ld b e l a rge ly o f fse t b y a t tendan t i n c reases i nt o ta l p as to ra l l a bo rc os ts . I na cco rdance w i th t he d i scuss ion i nC hap te r 2c once rn ing p o ten t ia l p os t -P le is tocene s i ze r educ t ions among c a t t le , s heep a nd g oa ts , T ab le 6 s umma r izes t he p roduc t ive v a r ia t ions e xpec ted a ssum ing a2 0% i n c rease i nt he s l augh te rp roduc t iv i ty o fc a t t le a nd a3 0% i n c rease i n t he s l augh ter p roduc t iv i ty o f s heep a nd g oa ts among e a r ly p as to ra l h e rds , h o ld ing m i lk p roduc t ion c ons tan t . P a r t icu la r ly s t r ik ing i nt hese d a ta i st he o bser va t ion t ha t m i lk p roduc t ion wou ld a ccoun tf o r mo re t han h a l f o ft he a nnua l c a lo r ic p roduc t iv i ty e xpec ted p e r h e rd r ega rd less o ft he s pec ies i n vo lved , e ven when p o ten t ia l p os t -P le is tocene s i ze r educ t ions a re a ccoun ted f o r . A s p rev ious ly n o ted , ar ecen t s t udy o fe ne rgy e x t rac t ion among t he N g isonyoka s ubsec t ion o ft he p as to ra l T urkana o fn or thwes t K enya , w ho h e rd a l f ou r o ft hese s pec ies , h as s hown t ha t m i lk c on t r ibu tes 6 1% o f t o ta l a nd 8 0% o f l i ves tock -de r ived f ood e nergy ( Coughenour e t a l . 1 985 :230 ) . Wh i le s uch d a ta s uppo r t t he s i gn i f icance o fm i lk -de r ived e ne rgy i n mode rn p as to ra l s oc ie t ies , i tmay s t i l l b e q ues t ioned w he the rd a i ry ing was s im i la r ly s i gn i f ican t i nt he e vo lu t ion o f e a r ly p as to ra l s t ra teg ies . I no rde rt o e ven tua l ly a dd ress t h is i s sue , s ubsequen t s ec t ions o ft h is c hap te r w i l l d e ta i l t he d rama t ic e ne rge t ic v a r ia t ions w h ich e x is t b e tween mea t -on ly a nd mea t p l us m i lk p as to ra l s t ra teg ies . O n t he c ompa ra t ive b as is o ft he c a lo r ic p roduc t iv i ty e xpec ted f r om h e rds o fe qua l s i ze , t he r e su l ts p resen ted i nT ab le 5w ou ld s ugges tt ha t u nde r a ve rage e xpec ted p as to ra l c ond i t ions i n t he e thnog raph ic p resen t , d romeda ry c ame ls wou ld b e a pp rox ima te ly 1 . 7 t imes a s p roduc t ive a s Z ebu a nd Z ebu -der ived A f r ican c a t t le , 6t imes a s p roduc t ive a s a nnua l ly b reed ing N ea r E as tern s heep , a nd 7 .8 t imes a s p roduc t ive a s a nnua l ly b reed ing N ea r E as tern g oa ts . T he r e su l ts p resen ted i nT ab le 6w ou ld s im i la r ly s ugges tt ha t i ne a r ly p as to ra l s t ra teg ies , d romeda ry c ame ls w ou ld h ave b een a pp rox ima te ly 1 . 6 t imes a s p roduc t ive a s A f r ican c a t t le , 5 .5 t imes a s p roduc t ive a s a nnua l ly b reed ing N ea r E as te rn s heep , a nd 7 .4 t imes a s p roduc t ive a s a nnua l ly b reed ing N ea r E as te rn g oa ts . H oweve r , a s d i scussed i ns ubsequen t s ec t ions o ft h is c hap te r , t he a b i l i ty t o e f fec t ive ly manage l a rge r h e rds o fc a t t le a nd sma l ls tock r e la t ive t o c ame ls wou ld p a r t ia l ly o f fse t t hese p roduc t ive v ar ia t ions .

T ab le 6 C ompa ra t ive C a lo r ic P roduc t iv i ty f o r P as to ra l H e rds C on ta in ing E xac t ly 1 00 H ead When P o ten t ia l P os t -P le is tocene S ize R educ t ions a re A ccoun ted F o r

SOURCE OF CALOR IES

DROMEDARY CAMELS k ca l /y r %

CATTLE

SHEEP

k ca l /y r

` ) / 0

k ca l /y r

GOATS %

k ca l /y r

` ) / 0

S laugh te r M i lk

3 ,786 ,154 1,273 ,060

2 5 .1 7 4 .9

4 , 104 ,576 5 ,497 ,065

4 2 .7 5 7 .3

9 68 ,990 1 ,780 ,189

3 5 .2 5 19 ,583 6 4 .8 1 ,524 ,987

2 5 .4 7 4 .6

TOTAL

1 5 ,059 ,214

1 00

9 ,601,641

1 00

2 ,749 ,179

1 00

1 00

7 5

2 ,044 ,570

A nnua l L abo r C os ts o fP as to ra l S t ra teg ies I n t roduc t ion F ou r p r inc ip le p as to ra l l a bo r c os ts w i l l b e e xam ined i nt he f o l low ing s ec t ions . T he f i r s t o f t hese i st he l a bo r r equ i red i na c tua l h e rd ing , wh ich p r ima r i ly c ons is ts o ft he t ime s pen t i nd r iv ing a n ima ls t o a nd f r om p as tu rage , g ua rd ing g raz ing h e rds , wa te r ing a n ima ls , a nd s ea rch ing f o r s t rays . A s m igh t b e s u rm ised , mos t o ft he a nnua l l abo rc os ts o f p as to ra l s t ra teg ies c ons is t o f t he t ime s pen t i nt hese a c t iv i t ies . T he s econd s e t o f l a bo r c os ts t o b e c ons ide red c ons is ts o f t he t ime s pen t i n movemen t o f e ncampmen ts . F ina l ly , t he t ime s pen t i n m i lk ing a nd s l augh te r ing a n ima ls w i l l b e e xam ined . Wh i le n ume rous s cho la rs h ave d ocumen ted a nd a na lyzed t he t r anshumance p a t te rns o f v a r ious p as to ra l s oc ie t ies , f ew h ave q uan t i f ied t he l a bo r c os ts o f p as to ra l f ood p roduc t ion . I n g ene ra l , t he b es tq uan t i ta t ive l a bo rd a ta r e la te t oc ame l p as to ra l is ts , a nd t hese d a ta w i l l b e u sed t o p a r t ia l ly r econs t ruc t s uch c os ts f o r s t ra teg ies i n vo lv ing c a t t le , s heep a nd g oa ts . I n i t ia l ly , t he p o ten t ia l e f fec ts o f a n ima l f eed ing a nd s oc ia l b ehav io rs o n a nnua l h e rd ing l a bo r r equ i remen ts w i l l b e e xam ined .

B ehav ior a nd H e rd ing L abo rC os ts Came ls t end t o move wh i le f eed ing , a nd may c ove r a s much a s 5 km i n 2 .5 h ours , d epend ing u pon t he a va i lab le p as tu rage ( Gau th ie r -P i l te rs a nd D agg 1 98 1 :33 ) . I n as i ng le d ay , a c ame l h e rd may c ove r mo re t han 2 0 km w h i le g raz ing ( Gau th ier -P i l te rs 1 974 :543 ) . F ur the r , i n d iv idua l membe rs o f ac ame l h e rd g ene ra l ly t end t o s p read a par t w h i le f eed ing a nd a re l e ss s oc ia lt han c a t t le o r s ma l ls tock ( Coppock 1 985 ; M cCabe 1 984 ) . B y c on t ras t , c a t t le a re s a id t o move mo re s l ow ly w h i le f eed ing , t end t o s t ay i n mo re c ompac t g roups , a nd d o n o tt end t os p read a pa r to r wande r( Dah l a nd H jor t 1 976 :247 ; J ohnson 1 969 :9 ; T o r ry 1 973 :117 18 ) . When h e rded i nm ixed f l ocks , g oa ts a re s a id t o s p read o u t more t han s heep , w i th t he l a t te r s tay ing t i gh t ly b unched w h i le f eed ing ( Johnson 1 969 :8 ; S a to 1 980 :43 ) . I ng ene ra l , g oa ts e xh ib i t al o ose r h e rd s t ruc tu re t han f l ocks o f s heep ( Ha fez a nd S co t t 1 962 :321 ; S cha l le r 1 977 :192 -94 , 1 98 -200 ) . Wh i le g oa ts may t r ave l a t af as ter p ace a nd c ove r g rea te r d i s tances t han s heep ( Behnke 1 980 :27 -8 ; N ico la isen 1 963 :46 ) , t hey a re s a id t o move a t as omewha t s l owe r p ace t han s heep w h i le f eed ing ( Bask in 1 974 :536 , 5 39 ) . When d is turbed , g oa ts h ave a g rea te r t endency t o s ca t te r t han d o s heep ( Ha fez a nd S co t t 1 962 :321) . E ven s o , A sad ( 1964 :51-2 ) , N ico la isen ( 1963 :46 ) , a nd S pencer ( 1973 :13 ) n o ted t ha t p or t ions o f s heep f l ocks w e re mo re l i ke ly t han g oa ts t o s t ray wh i le f eed ing o r when u na t tended d u r ing t he n i gh t . I fa lowed t o , c ame ls w i l l f eed 8t o 1 2 h ou rs ad ay , r ega rd less o ft he q ua l i ty o ft he p as tu rage ( Gau th ier -P i l te rs 1 974 :543 ; Gau th ie r -P i t te rs a nd D agg 1 98 1:36 ) . I ns t udy ing f ood i n take b y c ame l h e rds i nS ou the rn A lge r ia a nd Maur i tan ia , a ve rage d a i ly f eed ing t imes v a r ied f r om 8h ou rs i n May t h rough A ugus t , 1 0 t o 1 h ou rs i n Ma rch , a nd 1 t o 1 2 h ours i nN ovembe r ( Gau th ie r P i l te rs a nd D agg 1 981 :37 , T ab le 5 ) . A cco rd ing t oT o r ry 's o bse rva t ions o n t he h e rd ing d ay o ft he Gab ra ( 1973 :140 ) , m i lk ing a nd g raz ing w ou ld o ccupy a bou t t we lve h ou rs p e r d ay , af i gu re w h ich a l so i n c luded t r ave l t i me t o a nd f r om p as turage . Mos t s pec i f ica l ly , S a to ( 1980 :38 ) n o ted t ha t a t t he h e igh t o f t he d ry s eason , R end i l le s e t t lemen t -based h e rd ing i n vo lved 1 3 h ou rs d a i ly , w h ich i n c luded a n a ve rage o f 7 .6 h ou rs o f t r ave l t ime t o a nd f r om p as turage . B y c on t ras t , s imu l taneous R end i l le c amp b ased h e rd ing i n vo lved a pp rox ima te ly 1 .5 h ou rs d a i ly , wh ich i n c luded a n a ve rage o f4 . 1 h ou rs o f t r ave l t ime t o a nd f r om p as turage . T hus , d ur ing t he h e igh t o f t he d ry s eason , R end i l le s e t t lemen t -based h e rd ing a lowed c ame ls t of eed o n ly 5 .4 h ou rs d a i ly , w h i le t he i r c amp -based h e rd ing a lowed c ame ls t of eed a pp rox ima te ly 7 .4 h ou rs d a i ly . P resumab ly , d ur ing t he h e igh t o f t he w e t s eason , t r ave l t ime b e tween e ncampmen t a nd p as tu rage w ou ld b e r educed , w h i le f eed ing t ime w ou ld b e i n c reased . I n ad e ta i led s t udy o f t he c a t t le h e rd ing F u lan i o f Wes t A f r ica , v an R aay ( 1975 : 106 -29 ) n o ted t ha tt he i rc a t t le w e re a lowed t og raze a pp rox ima te ly 1 0 h ou rs p e rd ay , w i th a m in imum o f 7 .5 h ours w hen f odde r was a bundan t a nd amax imum o f1 2 .5 h ours w hen i twas s ca rce . S im i la r

7 6

d a ta o n s heep a nd g oa ts h as n o t b een l o ca ted , a l though S a to ( 1980 :43 ) n o ted t ha t t he R end i l le h e rd t he i r sma l ls tock " f rom s un r ise t os unse t e ve ry d ay . " D ur ing t he s umme r mon ths , c ame ls t end t od i scon t inue f eed ing d u r ing t he e x t reme h ea t o f m idday a nd i n s tead l i e c l us te red t oge the r i n as i ng le l o ca t ion wh i le r um ina t ing ( Gau th ie r P u t te rs 1 974 :547 ; Gau th ie r -P i l te rs a nd D agg 1 981 :35 ) . A cco rd ing t o Mus i l ( 1928 :337 ) , c ame ls i nN or th A rab ia wou ld d o s o b e tween n oon a nd t wo o rt h ree i nt he a f te rnoon . S a to ( 1980 :35 , 4 3 ) s im i la r ly n o ted t ha t b o th R end i l le c ame ls a nd sma l ls tock d i scon t inue f eed ing b e tween 1 2 :30 a nd 3 :30 , r e t i r ing i n s tead t o t he s hade o ft r ees t o r um ina te wh i le t he i r h e rde rs r es ted . P resumab ly , p as to ra l c a t t le e xh ib i tt he s ame t endency . I ti sa l so d u r ing t he f ou rt o f i ve h o t tes t mon ths o ft he y ea rt ha t c ame ls e xh ib i t e xcep t iona l f i de l i ty t o f am i l ia r s ources o f wa ter , r e turn ing t o t hem o n t he i r own f r om d i s tan t p as tures a t i n te rva ls o f 3t o 1 0 d ays ( Co le 1 975 :39 , 5 2 ; Gau th ie r -P i t te rs 1 974 :547 ; Gau th ie r -P i l te rs a nd D agg 1 981:50 -53 , 7 8 ; J ohnson 1 969 :126 ) . T hus , u nde r c ur ren t c ond i t ions o f d im in ished p reda t ion ( bo th w i ld a nd h uman ) , p as to ra l is ts i nS a 'ud i A rab ia , s ou the rn A lge r ia , T r ipo l i tan ia , a nd T ibes t i a re a b le t o l e ave t he i r c ame l h e rds u ngua rded d u r ing t he s umme r mon ths ( Co le 1 975 :52 ; Gau th ie r -P i l te rs a nd D agg 1 981:78 , 8 3 ; J ohnson 1 969 :126 ) , c on f iden t t ha t t he h e rds w i l l f eed o n wha teve r p as tu rage t hey m igh t f i nd a nd k now ing t ha t t hey w i l l p e r iod ica l ly r e tu rn t o s pec i f ic wa te r s ou rces . P resumab ly , t he c ame l 's s easona l f i de l i ty t o s pec i f ic wa ter s ou rces wou ld h ave b een as i gn i f ican t f ac to r i nt he h uman a b i l i ty t o i n t ia l ly c ap tu re a nd c on t ro l w i ld h e rds . T he T u rkmen is o fC en t ra l A s ia s t i l l e xp lo i tt h is t r a i tt oc ap tu re f e ra l c ame ls n ea rd raw w e l ls ( Bask in 1 974 :538 ) . D ur ing r a iny s easons , h oweve r , t h is s i tua t ion i sr eve rsed . T he r a iny s eason r esu l ts i n i n c reased p as tu rage o f h i ghe r mo is tu re c on ten t , l owe rt empe ra tures , h i gher r e la t ive h um id i ty , a nd t he f o rma t ion o fs ca t te red d eser to rs avannah r a in p oo ls , p onds , a nd p l ayas . A st hey a re sn o l o nge r d ependen t u pon p e renn ia l s ources o f wa te r , c ame ls t end t o s t ray mo re f r equen t ly d u r ing we t s easons a nd t hus r equ i re mo re i n tens ive h e rd ing a nd t he p e r iod ic t r ack ing a nd r ecove ry o ft he a n ima ls w h ich d o s t ray ( Co le 1 975 :48 ; Gau th ie r -P i t te rs a nd D agg 1 98 1 :83 ) . U nder d ry s eason c ond i t ions a nd w i th r educed p reda t ion , c ame l h e rd ing l abo r may b e r educed t o t he t ime s pen t d raw ing wa te r when a h e rd r e turns t o a wa te r s ource , as i tua t ion s eem ing ly b a lanced b y i n c reased managemen t r equ i remen ts a nd d ec reased wa ter ing l abo r d u r ing w e t s easons . I ng ene ra l , b o th c a t t le ( Tony 1 973 :85 , 18 ) a nd g oa ts ( Asad 1 964 :51) a re s a id t o r ead i ly r e turn t o e ncampmen ts f r om g raz ing w i thou t h av ing t o b e d r iven . O ve ra l l , t he l e ng th o f ah e rd ing d ay f o r c ame l , c a t t le , s heep a nd g oa t p as to ra l is ts s eems g enera l ly e qu i tab le , e ven t hough t he e xac t a l loca t ion o f l abo r b y h e rd ing t ask a nd a ge o f h e rde r i s e xpec ted t o v a ry a cco rd ing t ot he s pec ies a nd e co log ica l c ond i t ions I n vo lved ( see Coppock 1 985 ; McCabe 1 984 ) . T he mos t s t r ik ing v a r ia t ion i nh e rd ing l abo r b e tween s pec ies a ppea rs t o i n vo lve v a r ia t ions i nt he a dd i t iona l h e rd ing l a bo r r equ i red a s ar esu l t o f s easona l h e rd s ubd iv is ions . E f fec t ive p as to ra l h e rd managemen t d ur ing , a nd j u s t s ubsequen t t o , t he d e l ivery a nd ma t ing s eason (s ) r equ i res t he d i v is ion o fh e rds i n to sma l le rg roup ings a cco rd ing t oa ge a nd s ex c l asses . T h rough t he c on t ro l a nd man ipu la t ion o f t hese h e rd s ubd iv is ions , p as to ra l is ts a re a b le t o r egu la te t he b reed ing o fi n d iv idua l f ema les , m in im ize c on f l ic ts r esul t ing f r om ma t ing b ehav io rs among ma les , r egu la te t he d i s tances wh ich t he r e la t ive ly w eake ry oung m us tt r ave lt of eed , a nd a d jus t g raz ing t o max im ize m i lk a nd mea t p roduc t ion a cco rd ing t ot he c hang ing p hys io log ica l , mo rpho log ica l , a nd b ehav io ra l a t t r ibu tes o ft he a ge a nd s ex c l asses c omp r is ing ah e rd . H oweve r , t he f o rma t ion o f h e rd s ubd iv is ions a l so i n c reases t he n umbe r o fh e rde rs , a nd h ence h erd ing l abo r , r equ i red a nnua l ly p e r h e rd . A l though t he u se o f h e rd s ubd iv is ions i sb e t te r -documen ted among c ame l p as to ra l is ts , i ti sr ead i ly a ppa ren tt ha t t hey a re s im i la r ly emp loyed i nh e rd ing c a t t le , s heep a nd g oa ts . T he K a r imo jong , f o r e xamp le , a re s a id t o h e rd t he i rc a lves s epa ra te ly f r om t he i r mo the rs , wh i le e n t i re h e rds a re s easona l ly s ubd iv ided i n to t wo o r mo re h e rds f o r f eed ing a nd b reed ing p urposes ( Dyson -Hudson 1 960b :45 ; D yson -Hudson a nd D yson -Hudson 1 969 :88 , 1 970 :107 , 1 09 ; M ichae l L i t t le , p e rsona l c ommun ica t ion 1 985 ) , ap rac t ice s im i la r ly o bse rved among N g isonyoka p as to ra l is ts ( Dyson -Hudson a nd McCabe 1 985 ; McCabe 1 984 ) . T he K e l T amasheq a re a l so s a id t o h e rd c a lves s epa ra te ly f r om t he i r mo the rs ( S . Sm i th 1 980 :477 ) . S im i la r ly , B a r th n o ted t ha t t he B asse r i h e rded t he i r l ambs a nd k i ds s epa ra te ly f r om t he a du l t 7 7

membe rs o ft he i r h e rds ( 196 1:7 ) , a nd H esse ( 1984 :250 ) n o ted t ha t ma le s heep a nd g oa ts a r e o f ten h erded s epa ra te ly i no rde rt o r egu la te b reed ing a nd t o a ss is t i nh e rd movemen t . Wh i le n o t ing t ha t sma l ls tock i n s ou the r ly l a t i tudes d o n o t e xh ib i t t he f i xed s easona l b reed ing o bserved i n mo re n or ther ly l a t i tudes , S a to ( 1980 :54 ) o bse rved t ha t e ach R end i l le s heep a nd g oa t h e rd was n o rma l ly managed i ni n fan t , j u ven i le , a nd a du l t -ado lescen t s ubd iv is ions y ea r r o und . D yson -Hudson a nd D yson -Hudson ( 1970 : 107 ) s im i la r ly n o ted t ha t t he sma l ls tock h e rds o f t he K a r imo jong " may b e d i v ided i n to s eve ra l h e rds . " T he f o l low ing s ec t ions d e ta i l t h e r a t iona le a nd r equ i red l a bo ro ft hese s easona l h e rd s ubd iv is ions u s ing t he b e t te r -documen ted d a ta a va i lab le o nc ame l s t ra teg ies .

C ame lH e rd S ubd iv is ions a nd A nnua lH erd ing L abo r : T he R end i l le D a ta Wh i le T o r ry ( 1973 :89 -90 ) b r ie f ly n o ted t ha t Gab ra c ame l h e rds w e re s ubd iv ided a cco rd ing t o a ge -sex c l asses f o r e f fec t ive managemen t d u r ing t he i r b reed ing s eason , t he mos t e xp l ic i t e t hnog raph ic a ccoun t o ft he r a t iona le a nd s t ruc tu re o fs uch s ubd iv is ions i st ha t p resen ted b y S a to a sb ased u pon h i s o bse rva t ions among t he R end i l le o fN o r the rn K enya . E ve ry c ame l h e rd i sd i v ided i n to 2 -5 h e rd ing g roups a cco rd ing t o t h e g row th s t age o ft he c ame ls a nd t he a va i lab i l i ty o f manpowe r , i no rde rt op l an f o r e fec t ive r ep roduc t ion a nd m i lk s upp ly f o r b o th s uck l ing c ame ls a nd h umans . B as ica l ly , i ti sd i v ided i n to t wo g roups a tt i mes o the rt han t he d e l ivery a nd t he ma t ing s easons : ag roup o fa du l t a nd a do lescen t c ame ls ( gaa l ) a nd ag roup o f j u ven i le c ame ls w h ich h ave b een w eaned b u t h ave n o t y e t r eached ma tu r i ty ( n io loho o r u rbo l ) . A s t he c a lves a re b o rn , a n a dd i t iona l t h i rd g roup ( i l iban ) c omposed o fs uck l ing i n fan ts a nd t he i r mo the rs , i sf o rmed . S oon t he rea f te r , when c ame ls b eg in t ob ecome s exua l ly e xc i ted , ag roup o f a du l t a nd a do lescen t c ame ls i sf i r s t d i v ided i n to t wo g roups t o f ac i l i ta te r ep roduc t ive e f f ic iency , a nd t o p reven t e xc i ted s t ud c ame ls f r om f i gh t ing among t h emse lves . E ach g roup h as o ne a c t ive s t ud . O ne g roup ( gaa l imoo lo ) i s f o rmed t op rov ide f o r h uman s ubs is tence a nd i sc omposed o fo ne s t ud c ame l , s eve ra lp ack c ame ls a nd many m i lk ing c ame ls . T he o t he r( gaa l i fo r r ) i sf o rmed f o r t he p urpose o fr ep roduc t ion . T h is l a t te rg roup i sc omposed o fo ne s t ud c ame l a nd a du l t -ado lescen tc ame ls o fb o th s exes . I nt he l a t te r h a l f o ft he ma t ing s eason , at h i rd g roup ( gaa l ihoopu ) o fa du l t a nd a do lescen t c ame ls i sf o rmed a nd i sc omposed o f o ne s t ud c ame l a nd t hose f ema les s t i l l n o t p regnan t i nt he a bove -men t ioned t wo g roups . T h is t h i rd g roup i s r a re ly o bserved , e spec ia l ly i n c amp -based h erd ing ( v ide i n f ra ) , b ecause i tc anno t b e f o rmed u n less t he re a re e nough s t ud c ame ls ( a t l e as t t h ree ) a nd h e rde rs . F i na l ly , a s s oon a s t hey b ecome p regnan t , f ema le c ame ls a re g a the red t oge the r i n to a g aa l i for r g roup . A f te r t he e nd o f t he ma t ing s eason , t he c ame ls o fg aa l ihoopu a re r e turned t ot he i rp rev ious g roup a nd t he g aa l imoo lo s t ud c ame l i ss h i f ted t o t he g aa l i for r . T hus t wo g roups o f a do lescen t a nd a du l tc ame ls a re f o rmed , g aa l imoo lo a nd g aa l i for r . A s i n fan t c ame ls a re g radua l ly w eaned t hey a re s h i f ted t o g aa l imoo lo , t oge the rw i th t he i r mo the rs , a nd a f ter t hey h ave b een c omp le te ly weaned t he mo ther in fan t g roup i sd i v ided i n to a g aa l imoo lo a nd a j u ven i le g roup [ Sa to 1 980 :26 -27 ] . T hus , t he g ene ra l p a t te rn o fh e rd s ubd iv is ions p rac t iced b y t h e R end i l le i n vo lves t he d i v is ion o f ah erd i n to a du l t -ado lescen t a nd j u ven i le g roups f o r t he ma jo r i ty o ft he y ea r ( app rox ima te ly e i gh t mon ths ) . H oweve r , d u r ing t he d e l ive ry a nd ma t ing s eason , h e rds a re d i v ided i n to a n a du l t -ado lescen t ma t ing h e rd , a n ,a du l t -ado lescen t s ubs is tence h erd , a nd a mo the r in fan t g roup ( app rox ima te ly t wo m on ths ) . S ubsequen t t ot he d e l ive ry a nd ma t ing s eason , h erds a r e r e s t ruc tu red i n to a du l t -ado lescen t a nd mo the r in fan t g roups ( app rox ima te ly t wo mon ths ) . A f te r t h is , h e rds a re a ga in r e s truc tured i n to a du l t -ado lescen t a nd j u ven i le g roups , a nd t h e s easona l c yc le b eg ins a ga in ( Sa to 1 980 :55 -56 ) . Why as t ud c ame l i si n c luded w i th t he s ubs is tence h e rd ( gaa l imoo lo ) i su ns ta ted . S a to ( 1980 :53 -54 ) a l so r e co rded s easona l c hanges i nt he h e rde r p opu la t ion s t ruc tu re o f R end i l le c ame l c amps a nd t he ir r e la t ionsh ip t o t he c hang ing s t ruc ture o f c ame l h erd 7 8

s ubd iv is ions . T ab le 7s umma r izes h i sd a ta . P lease n o te t ha t a tR end i l le c ame lc amps , h e rd ing i s a n e xc lus ive ly ma le o ccupa t ion , a l though y oung f ema les o f ten h e rd t hose c ame ls r e ta ined a t m a in r es iden t ia l s i tes ( Sa to 1 980 :378 ) . F rom S a to 's d a ta , i twou ld s eem t ha t w hen a R end i l le c ame l h e rd i sd i v ided i n to a du l t -ado lescen t a nd j u ven i le g roups , o n ly 1 . 51 h e rde rs a re r equ i red d a i ly . H oweve r , when h e rds a re d i v ided i n to a n a du l t -ado lescen t ma t ing h erd , a n a du l t a do lescen t s ubs is tence h e rd , a nd a mo the r in fan t g roup , 2 .55 h e rde rs a re r equ i red d a i ly . F i na l ly , when a h e rd i sd i v ided i n to a du l t -ado lescen t a nd mo the r in fan t g roups , o n ly 1 . 82 h e rde rs a re r equ i red d a i ly ( Sa to 1 980 :53 -54 ) . F i gu re 5d iag rams t hese s easona l c ame l h e rd ing l a bo r r equ i remen ts among t he R end i l le f o r ao ne y ea rp e r iod . P ar t icu la r ly s t r ik ing i st he p e rcen tage c on t r ibu t ion o f e ach h erde r a ge -c lass t o t o ta l R end i l le c ame l h e rd ing r equ i remen ts . T ab le 8 s umma r izes t h is p e rcen tage c on t r ibu t ion b y a ge c l ass , wh i le F igu re 6 d iag rams t he i r r e la t ive h e rd ing l a bo r c on t r ibu t ions b y s eason o ve r ao ne y ea r p e r iod . I ng ene ra l , i tw ou ld s eem t ha t y oung ma les 7t o 2 1 y ea rs o fa ge a re h e rde rs y ea r r o und a nd p rov ide a pp rox ima te ly 7 2% o ft o ta l a nnua l c ame l h erd ing l a bo r . U nma r r ied b u t a du l t m a les ( app rox ima te ly 2 2 -3 1 y ea rs o l d ) c on t r ibu te 2 3% o fa nnua l ly r equ i red c ame l h erd ing l a bo r . B yc on t ras t , a du l t ma r r ied m a les ( 3 1+ y ea rs o l d ) a re o n ly i n vo lved w i th c ame l h e rd ing d u r ing t he p r inc ip le b reed ing a nd c a lv ing p e r iod u nde rc ond i t ions o f max imum h e rd s ubd iv is ions . O ve ra l l , a du l t ma r r ied ma les p rov ide o n ly 5% o ft o ta l a nnua l c ame l h e rd ing l abo r . T ab le 9 s umma r izes t h e d a i ly c ame l h e rd ing l a bo r r equ i remen ts o ft he R end i l le o ve r ao ne y ea r p e r iod . T he r esu l ts s ugges t t ha t e ach R end i l le c ame l h e rd r equ i res a pp rox ima te ly 6 32 d ays o f h e rd ing l a bo r p er y ea r . S a to 's d a ta o n R end i l le c ame l h e rd s ubd iv is ions a nd a nnua l h e rd ing l a bo r s hou ld a l so a pp ly t o Gab ra c ame l h e rds , a s i ss ugges ted b y T o r ry 's b r ie f o bserva t ions ( 1973 :89 -90 ) . F u r the r , i ti se xpec ted t ha tt hese a nnua l l a bo r r equ i remen ts wou ld a l so a pp ly t oc ame l h e rds i n N o r th A f r ica a nd t he N ea r E as t , s i nce s easona l h e rd s ubd iv is ions a re p a t te rned b y t he a nnua l r ep roduc t ive c yc le e xh ib i ted b y a l c ame ls . S ome s uppor t f o r t h is s ugges t ion e x is ts i nC o le 's o bse rva t ions o nA l Mur rah c ame l h e rd s ubd iv is ions i nS a 'ud i A rab ia .

C ame lH erd S ubd iv is ions a nd A nnua lH erd ing L abor : T he A l Mu r rah D a ta Wh i le l e ss c onc ise t han S a to 's o bse rva t ions among t he R end i l le , C o le ( 1975 :36 -38 ) h ad e a r l ie r r eco rded t he c ame l h e rd s ubd iv is ions u t i l ized b yt he A l Mu r rah b edou in o fS a 'ud i A rab ia . C o le n o ted t ha t t he A l Mu r rah d i v ided t he i r c ame l h e rds i n to f our d i s t inc t g roups . T he f i rs t o f t hese was c omposed o ff ema les w i th n ew ly -bo rn c a lves . A cco rd ing t oC o le , t h is g roup r equ i red t h e mos t h e rd ing a t ten t ion , s i nce y oung c a lves c anno t move r ap id ly a tf i r s t a nd mo the rc ame ls m us t d r ink a t l e as t o nce aw eek e ven d u r ing t he r a iny s eason u n less g rasses a re p a r t icu la r ly p l en t i fu l . F ur the r , t he m i lk o ft hese c ame ls i sr i ch a nd p l en t i fu l , a nd d eemed e spec ia l ly " swee t " b y t he A l Mu r rah . A s ar esu l t , t h is h e rd s ubd iv is ion i sc on t inuous ly g ua rded , a l though i ti so n ly a lowed t o g raze c l ose t o t he e ncampmen t d o t o i t s r es t r ic ted mob i l i ty . T he h e rd ing o f t h is s ubd iv is ion i su sua l ly t he r e spons ib i l i ty o ft he e l des tt een -aged s on o ft he h e rd o wner . T he s econd h e rd s ubd iv is ion n o ted b y C o le c ons is ted o f l a c ta t ing f ema les w i th u nweaned b u t o l de r y oung . T h is g roup i ss a id t op rov ide " l ess -swee t m i lk , " b u t i sa b le t o move f as te r a nd g raze f a r ther f r om t he e ncampmen t t han t he o the rf ema le in fan t d i v is ion s i nce t he i r y oung a re m o re mob i le . T h is s ubd iv is ion was a l so g ua rded c on t inuous ly . U sua l ly , t he h e rd ing o f t h is g roup was e n t rus ted t o at een -aged s on , ay oung man , o r ay oung u nma r r ied d augh ter . T he t h i rd h e rd d i v is ion c ons is ted o fp regnan tc ame ls . T h is s ubd iv is ion was a lowed t og raze o nt he i r o wn n ea r c amp d ur ing t he d ay , a nd o n ly r equ i red s omeone t o r ound t hem u p a nd d r ive t hem b ack t oc amp b e fo re s unse t . T he f our th s ubd iv is ion n o ted b y C o le c ons is ted o ft he ma le c ame ls u sed a s c a r r ie rs , t he s t ud c ame l o rc ame ls , a nd t h e r i d ing c ame ls , mos t o f wh ich we re f ema le . When n o t i nu se , b o th t he c a r r ie rs a nd t he r i d ing c ame ls w e re h obb led a nd l e f t t og raze o n t he i r own n ea rc amp . L i ke t he s ubd iv is ion c omposed o f p regnan t c ame ls , t h is g roup r equ i red m in ima l h e rd ing l a bo r . C o l lec t ive ly , t he h e rd d i v is ion c ons is t ing o f p regnan t f ema les a nd t ha t c ons is t ing o f c a r r ie rs , s t uds , a nd r i d ing c ame ls r equ i red o ne a dd i t iona l p a r t t ime h erde r . A s s uch , C o le s t a ted t ha t e ach A l Mur rah c ame l h e rd r e qu i red t he d a i ly l abo r o ft wo f u l l t ime h e rde rs a nd o ne p a r t t ime h e rde r ( 1975 :38 , 1 6 1) . 7 9

T ab le 7 N umbe ro fH e rde rs S easona l ly R equ i red P e rR end i l le C ame l H e rd

HERDER CENSUS BY AGE C LASS ( ALL MALES ) S easona l U nma r r ied U nc i rcums ized H e rd D a te o f Ma r r ied 1 6 -21 7 15 >3 2 2 2 -32 T o ta l S ubd iv is ion C ensus A A A( Mean ) B C

1 0 /5 /75 3 /31/76 5 /30 /76 7/ 22 /76

1 3 1 1 1 2 2 0 1 7

6 9 7 .5 1 2 1 3

0 0 0 8 0

1 6 1 3 1 4 .5 1 6 2 1

3 5 3 3 3 4 5 6 5 1

#o f H e rds P resen t 2 1 2 4 2 2 .5 2 2 2 8

#o f H e rde rs P e rH e rd 1 .67 1 .38 1 .51 2 .55 1 .82

K ey t o H e rd S ubd iv is ions A =A du l t -ado lescen t a nd j u ven i le g roups ( Sep tembe rt h rough A p r i l ) B =A du l t -ado lescen t ma t ing h erd , a du l t -ado lescen t s ubs is tence h e rd , a nd mo the r in fan t g roups ( May t h rough J une ) C =A du l t -Ado lescen t a nd mo the r in fan t g roups ( J u ly t h rough A ugus t )

Nu mber o f Herders

3

2

J an.

March

D ry Season ,

May

July

Main Rainy Season

Sept.

D ry Season

Nov. Autu mn Rains

F i gu re 5 . N umbe ro fh e rde rs r equ i red s easona l ly p e r R end i l le c ame l h e rd .

8 0

T ab le 8 C on t r ibu t ion o fE ach Ma le H e rde rA ge C lass t oT o ta l A nnua l R end i l le C ame l H e rd ing L abo r

HERD SUBD IV IS ION

% OF ANNUAL HERD ING L ABOR CONTR IBUTED Ma r r ied U nma r r ied U nc i rcums ized > 32 2 2 -32 1 6 -21 7 15

A( Mean ) B C

0 1 4 .3 0

2 2 .1 2 1.4 2 5 .5

3 5 .3 3 5 .7 3 3 .3

4 2 .7 2 8 .6 4 1.2

Mean

4 .8

2 3 .0

3 4 .8

3 7 .5

K ey t o H erd S ubd iv is ions A =A du l t -ado lescen t a nd j u ven i le g roups ( Sep tembe rt h rough A p r i l ) B =A du l t -ado lescen t ma t ing h e rd , a du l t -ado lescen t s ubs is tence h e rd , a nd mo the r in fan t g roups ( May t h rough J une ) C =A du l t -ado lescen t a nd mo the r in fan t g roups ( J u ly t h rough A ugus t )

60 7-1 5

o f Total Herders

40 16 -21

30

L .

22 -31

20

10 32 +

o J an.

March

I I 1 Dry Season i

May 1

J uly

Main SRainy

Sept.

I , Dry Season i

Nov. I Autu mn Rains

F igu re 6 . R e la t ive s easona lc on t r ibu t ion t oc ame l h e rd ing l a bo rb y a ge c l ass among R end i l le ma le h e rde rs .

8 1

i 1 I i

T ab le 9 S umma ry o fA nnua l C ame l H e rd ing L abo r R equ i remen ts Among t he R end i l le

MONTH

SEASON

J anua ry F eb rua ry Ma rch

D RY

- -

#o f d ays

H erd D iv is ions

#o f H e rde rs p e rD ay

#o f H e rd ing D ays /Mon th

3 1 2 8 3 1

A A A

1 .51 1 .51 1 .51

4 6 .81 4 2 .28 4 6 .81

A

1 .51 2 .55 2 .55

4 5 .30 7 9 .05 7 5 .00 5 6 .42 5 6 .42 4 5 .30 4 6 .81 4 5 .30 4 6 .81

-

MA IN R A INY S EASON

A p r i l May J une

3 0 3 1 3 0

D RY

J u ly A ugus t S ep tember

3 1 3 1 3 0

A

1 .82 1 .82 1 .51

Oc tobe r N ovembe r D ecembe r

3 1 3 0 3 1

A A A

1 .51 1 .51 1 .51

AUTUMN RA INS

TOTALS

3 65

/ / / / / / / / / / / / / / / / / / / / / / / / / / / / / / / / / / / / / / /

- -

6 32 .31

K ey t oH e rd D iv is ions A =A du l t -ado lescen t a nd j u ven i le g roups B =A du l t -ado lescen t ma t ing h e rd , a du l t -ado lescen t s ubs is tence h e rd , a nd mo the r in fan t g roup C =A du l t -ado lescen t a nd mo the r in fan t g roups

T he A l Mu r rah h e rd s ubd iv is ions r eco rded b y C o le c l ose ly a pp rox ima te t hose r e co rded b y S a to f o r R end i l le c ame l h e rds . H oweve r , C o le 's a ccoun t l a cks t he s easona l d ynam ics a ppa ren t i nS a to 's d a ta . P resumab ly , t he A l Mu r rah , l i ke t he R end i l le , mus t s easona l ly r es t ruc tu re t hese s ubd iv is ions i no rde r t oa ch ieve e f fec t ive h e rd managemen t w h i le m in im iz ing r equ i red h e rd ing l abo r . I n p ar t icu la r , C o le 's a sser t ion t ha t t he A l Mur rah w ou ld ma in ta in a h e rd s ubd iv is ion c omposed o ff ema les w i th n ew ly -bo rn y oung f o rt he c ou rse o fa n e n t i re y ea r makes l i t le s ense . Wha t C o le h as a ppa ren t ly p resen ted a re t he max imum s easona l h e rd s ubd iv is ions wh ich t he A l Mu r rah emp loy . T hus , h i s o bserva t ion t ha t e ach A l Mu r rah h e rd r equ i red t he d a i ly a t ten t ion o f 2 .5 h e rde rs ( Co le 1 975 :38 , 1 6 1) w ou ld s eem ing ly r e la te t o max imum s easona l A l M ur rah h erde r r equ i remen ts d ur ing t he ma t ing a nd b i r th ing s eason o ft he i rc ame ls . S ign i f ican t ly , t h is i s t he s ame max imum d a i ly h e rde r l a bo r r equ i remen t n o ted b y S a to f o rt he R end i l le d u r ing t he i r c ame ls ' ma t ing s eason . I ns ubsequen t l a bo rc a lcu la t ions , i tw i l lt he re fo re b e a ssumed t ha t c ame l h e rds i nN o r th A f r ica a nd t he N ea r E as t wou ld a l so r equ i re 6 32 d ays o fa nnua l h e rd ing l a bo r . H erd S ubd iv is ions a nd A nnua lH erd ing L abo rf o r C a t t le , S heep a nd G oa tP as to ra l is ts Wh i le t he h e rd ing c os ts o fa l p as to ra l h e rds wou ld m in ima l ly c ons is t o fo ne h e rde rf o r3 65 d ays a y ea r , t he a dd i t iona l c os ts o f s easona l h e rd s ubd iv is ions a re n o t we l l -documen ted f o r c a t t le , s heep a nd g oa t h e rds . A cco rd ing t o D yson -Hudson a nd D yson -Hudson ( 1969 :88 , 1 970 :109 ) , t he K a r imo jong n o rma l ly d i v ide t he i rc a t t le h e rds i n to t wo o r mo re s ubd iv is ions , w i th o ne s ubd iv is ion k ep t a th e rd ing c amps y ea r round , a nd t h e o the r k ep t a tt he ma in s e t t lemen t f o r a s much o ft he y ea r a s p oss ib le . When g raz ing b ecomes s pa rse n ea rt he ma in s e t t lemen t , t h is l a t te rs ubd iv is ion may j o in t he c amp h e rd , o ri s moved a way a s as epa ra te u n i t . P resumab ly , s ubd iv is ions f o r b reed ing p u rposes a re a l so made . I nt he a bsence o f mo re c onc ise d a ta , i tw i l l b e a ssumed t ha t t he t o ta l a nnua l l abo r c os ts o f c a t t le h e rd ing a pp rox ima te t he 6 32 d ays r equ i red f o rc ame l h e rds .

8 2

A cco rd ing t oT ony ( 1973 :105 -6 , 1 974 :97 -8 ) , e ach Gab ra s ma l ls tock h e rd r equ i red o ne t o t h ree h erders , r e f lec t ing v a r ia t ions i ns easona l l a bo r r equ i remen ts a s ar esu l t o f a ge -sex s ubd iv is ions . S a to ( 1980 :27 ) was a ga in mo re s pec i f ic i n d ocumen t ing t he r a t iona le a nd s t ruc ture o f R end i l le sma l ls tock s ubd iv is ions . D ur ing t he i rf i r s t mon th , i n fan t s heep a nd g oa ts a re k ep t i np ens a te ncampmen ts , w i th w ean ing a nd t he a d jus tmen t t od r ink ing a nd f eed ing o n t h e i r o wn a ch ieved d u r ing t h e i rs econd a nd t h i rd mon ths . D u r ing t he i rf o ur th a nd f i f th mon ths , t he n ow j u ven i le s heep a nd g oa ts a re f o rmed i n to a m ixed g roup f o rr egu la rh erd ing a way f r om e ncampmen ts . F rom t he i rs i x th mon th o n , t he n ow a do lescen t s heep a nd g oa ts a re m ixed w i th a du l ts f o rr egu la rh e rd ing . T he s ame p a t te rn a ppa ren t ly a pp l ies t o Gab ra ( Tony 1 974 :97 -8 ) a nd T ua reg ( N ico la isen 1 963 :36 -9 , 4 6 ) sma l ls tock h e rds . F rom t hese d a ta , as ma l ls tock h e rd w i th o n ly o ne b reed ing s eason s hou ld m in ima l ly r equ i re o ne f u l l t ime h e rde rf o rt he ma in h e rd o fa du l ts f o r3 65 d ays ay ea r ,o ne p a r t t ime h e rde r f o rt wo mon ths ( app rox ima te ly 3 1 h e rd ing d ays ) t o manage i n fan t s ubd iv is ions f eed ing a round e ncampmen ts , a nd o ne f u l l t ime h e rde r f o r t wo mon ths ( app rox ima te ly 6 1 h e rd ing d ays ) t o manage t he j u ven i le s ubd iv is ion . A nnua l h e rd ing r equ i remen ts f o r asma l ls tock h e rd w i th o n ly o ne b reed ing s eason s hou ld t he re fo re amoun tt oa pp rox ima te ly 4 57 h e rd ing d ays ( 365 +3 1 + 6 1 =4 57 ) . I ft wo d i s t inc t b reed ing s easons o ccu r red , a nnua l h e rd ing c os ts w ou ld p resumab ly r i se 2 0% , t oa pp rox ima te ly 5 49 d ays ( 365 +6 2 +1 22 =5 49 ) . N onseasona l , y ea r round b reed ing among sma l ls tock w ou ld p resumab ly r equ i re t wo f u l l t i me h e rders f o ra du l t -ado lescen t a nd j u ven i le s ubd iv is ions , a n a dd i t iona lf u l l t ime h e rde rf o r a s epa ra te b reed ing h e rd , a nd o ne p ar t t ime h e rde r y ea r round f o r t he i n fan ts f eed ing n ea r e ncampmen ts . T h is w ou ld amoun t t o 3 .5 h e rde rs r equ i red d a i ly f o r 3 65 d ays a y ea r , o r a n a s tound ing 1 , 278 d ays o f h e rd ing l a bo r p e r y ea r . A cco rd ing t o S a to ( 1980 :54 -5 ) , R end i l le s ma l ls tock h e rds d o r equ i re a pp rox ima te ly 3 .2 h e rde rs d a i ly a s ar esu l t o fn on -seasona l , y ea r r o und b reed ing . P resumab ly , s uch t r emendous l a bo rc os ts a re a tl e as t p a r t ia l ly o f fse tb yg rea t ly i n c reased h e rd p roduc t iv i ty . U n for tuna te ly , n o q uan t i ta t ive d a ta e x is t o n t he o vera l l p roduc t iv i ty o f sma l ls tock h e rds ma in ta ined u nde rs uch c ond i t ions . I ns ubsequen t e ne rge t ic c a lcu la t ions , R edd ing 's p roduc t iv i ty d a ta o n s easona l ly b reed ing N ea r E as tern s heep a nd g oa ts w i l l b e u t i l ized , a ssum ing a h e rd ing l a bo r r equ i remen t o f4 57 h e rd ing d ays a nnua l ly . H ypo the t ica l c a lcu la t ions w i l l a l so b e d e r ived f r om h i s d a ta i no rder t o e va lua te t he p o ten t ia l e f fec ts o n t he e ne rge t ics o fs heep a nd g oa t s t ra teg ies w hen s ma l ls tock b reed t w ice ay ea r , a ssum ing ah e rd ing l a bo rr equ i remen to f5 49 h e rd ing d ays a nnua l ly .

T he H e rd ing D ay v s . D a i ly H e rd ing L abo rH ou rs I ti sa ppa ren tt ha to nce ap as to ra l h e rd h as b een d r iven t op as ture a nd b eg ins t of eed , t he s ubsequen t g ua rd ing o f t ha t h e rd d oes n o t i n vo lve c on t inuous h e rd ing l a bo r . F o r e xamp le , T ony c ha rac te r ized c ame l h e rd ing l a bo r among t he Gab ra a sf o l lows ( 1973 : 140 ) : Ih ave j o ined c ame l h e rdboys o n an umbe r o f o ccas ions , a nd wha t mos t i mp ressed me a bou t t h is a c t iv i ty w as t he e no rmous amoun t o f h e rd ing t i me s pen t i ns hee r l o a f ing . E xcep tf o rt he e fo r t o ft ak ing t he b eas ts t op as tu re a nd g u id ing t hem b ack h ome , r e la t ive ly l i t le t ime o r e ne rgy i ss pen t i nd i rec t ing t he ir movemen ts . T hey mus t b e w a tched , h oweve r , a nd p e rhaps o nce e very f ew h ours t he mo re c onsc ien t ious h e rde rs w i l l c heck t os ee i fa l a n ima ls a re p resen t , e spec ia l ly w hen c ame ls h ave n o t d runk f o rs eve ra l d ays a nd a re t hus i n c l ined t ow ande ro f fi nt he d i rec t ion o ft he w a te rh o les . F ur the r , S a to c ha rac te r ized R end i l le c ame l h e rd ing l a bo ri n as im i la rf a sh ion ( 1980 :35 ) : When t he c ame ls a re n o tt oo d i spe rsed , t he h e rde r may r e s tc om for tab ly i nt he s hade o ft r ees , p ay ing o n ly i n te rm i t ten t a t ten t ion t o make s ure t ha tt he a n ima ls d o n o tw ande rt oo f a ra nd t ha tt hey a re f eed ing p roper ly . J enn ings -B ram ley ( 1907a :23 -24 , 3 1) a l so c ommen ted o nt he l ow l abor r equ i remen ts o fc ame l h e rd ing i nt he S ina i , s ugges t ing t ha t m uch o f ah e rders t i me w as a c tua l ly s pen to no the rt a sks .

8 3

H is l e isu re h ou rs - a nd t hey a re many - a re s pen t i np l ay ing t he f l u te , k n i t t ing t he c ame l 's woo l , w h ich t he w omen h ave s pun , i n to c aps o rt w is t ing i ti n to t he a ka l ( r ope ) w i th w h ich t he c ame ls a re f a s tened a tn i gh t , o r mer ica -co rd f o r k eep ing o nt he k ou f ia o r h ead h andke rch ie f [ J enn ings -B ram ley 1 907a :24 ] . J enn ings -B ram ley a dd i t iona l ly n o ted t ha t w hen a do lescen t f ema les w e re e ngaged i nh e rd ing , t he i rt i me w as o f ten s pen t i ns p inn ing w oo lo rh a i r( 1907a :3 1) . A cco rd ing t oC o le 's o bserva t ions among t he A l Mu r rah ( 1975 :37 ) , h erde rs i nc ha rge o fl a c ta t ing f ema les w i th i n fan t y oung c ou ld e xpec t t o s pend a t l e as t p a r t o f t he i r h e rd ing d ay s oc ia l iz ing w i th v i s i tors ( Co le 1 975 :37 ) . S im i la r ly , y oung B agga ra h e rde rs w e re o bserved t o p oo l t he i r r espec t ive c a t t le h e rds w hen g rasses w e re p l en t i fu l s o t ha t t hey c ou ld c ha t a nd p l ay t oge ther a s t he a n ima ls g razed ( Cunn ison 1 966 :69 ) . Mos t i mpo r tan t ly , M us i l ( 1928 :337 ) n o ted t ha t o nce aRwa la h e rde rh ad d r iven h i s c ame ls t o p as ture , " t he h erdsman s i ts d own i ns ome h i ghe r p l ace t he b e t ter t o o ve r look h i s h e rd , s i ngs , c a rves s ome th ing , o r s eeks e d ib le p l an ts a nd b u lbs . " D i ckson ( 195 1:60 ) a l so n o ted t ha t M u ta i rc ame l h e rde rs w ou ld l o ok f o rt r uf les o r h un t sma l l a n ima ls " t ow h i le a way t he l o ng d ay . " S im i la r ly , S a to ( 1980 :9 ) n o ted t ha tt he R end i l le w ou ld c o l lec t e d ib le p l an ts a nd h oney w h i le o n r ou te t o p as tu res . S . Sm i th ( 1980 :480 ) n o ted t ha t K e l T amasheq c ame l h e rde rs i nt he S ahe l s easona l ly c o l lec ted w i ld g ra ins e ncoun tered w h i le h e rd ing . D ona ld 0 . H enry ( pe rsona l c ommun ica t ion 1 984 ) a l so o bse rved i ns ou the rn J o rdan t ha t c ame l h e rde rs o f ten s pend p a r t o f t he i r h erd ing d ay f o rag ing f o r p l an t r e sou rces . P e te r J ones ( pe rsona l c ommun ica t ion 1 985 ) n o ted t ha t Maasa i c a t t le h e rde rs i nE as t A f r ica p rocu red a nd c a rved s t ick t oo ls w h i le h e rd ing , a nd w as t o ld t ha t Maasa i c h i ld ren w ou ld f o rage f o r r oo ts w h i le h e rd ing . F i na l ly , D yson -Hudson a nd D yson -Hudson ( 1969 :89 , 1 970 :117 ) n o ted t ha t K a r imo jong h e rde rs wou ld s easona l ly g a the r w i ld f r u i ts a nd b e r r ies wh i le h e rd ing t he i r c a t t le i ns pec i f ic r eg ions . D yson -Hudson ( 1960b :46 ) f u r the r n o ted t ha t t hey w ou ld d i g e d ib le r oo ts a nd g a the r h oney w h i le h erd ing . I t w ou ld t he re fo re s eem t ha t a tl e as t p a r t o ft he t i me s pen t i n mov ing a h e rd t o p as tu rage , a nd p a r t icu la r ly i ng ua rd ing a h e rd o nce i tb eg ins f eed ing , i sa c tua l ly d evo ted t o n on -he rd ing a c t iv i t ies , i n c lud ing s oc ia l iz ing , h and t a sks , a nd f o rag ing f o rf ood i t ems u n re la ted t ot he d i rec t p roduc t iv i ty o ft he h e rd . I n s ubsequen t c a lcu la t ions , a n e i gh t h our mean h e rd ing d ay w i l l b e a ssumed a s a pp rop r ia te f o rt he t ime a c tua l ly s pen td a i ly i nd i rec t h e rd ing l a bo r . I n a1 2 h our h e rd ing d ay ,t h is a ssump t ion a lows f o ur h ou rs f o rf o rag ing , s oc ia l iz ing , r e s t ing a nd h and t a sks , w i th e i gh t h ou rs o f mean d a i ly l a bo rd i rec t ly d evo ted t ot he p as to ra l h e rd . I ti ss t rong ly s uspec ted , h oweve r ,t ha t t h is s t i l l i n c ludes t ime a c tua l ly a loca ted t on onhe rd ing a c t iv i t ies . H oweve r , a tt h is r a te , t he 6 32 d ays o f h e rd ing l a bo r p e r c ame l o r c a t t le h e rd a s e s tab l ished a bove w ou ld amoun t t o 5 , 056 h ours o f l abo r a nnua l ly , w h i le t he 4 57 h e rd ing d ays r equ i red f o r as ma l ls tock h e rd b reed ing o nce ay ea rw ou ld amoun tt o3 ,656 h ours o fl a bo r a nnua l ly , o r4 ,392 h ours w i th t wo b reed ing s easons . T hese h e rd ing l a bo r c os ts w i l l b e emp loyed i ns ubsequen t c a lcu la t ions o f p as to ra l e ne rge t ics .

Max imum H e rd S ize a nd A nnua lH e rd ing L abo r P resumab ly ,t he re i sa n u ppe rl i m i tt ot he a c tua l s i ze o f ap as to ra l h e rd o r h e rd s ubd iv is ion w h ich c an b e r e a l is t ica l ly managed b yo ne h e rde rg i ven s uch f a c to rs a st he b ehav io ra l a t t r ibu tes o f t he s pec ies i n vo lved , t he d i f f icu l ty o ft he t e r ra in , t he p resence o f p reda to rs , t he a ge a nd a b i l i t ies o f t he h erde r , a nd p rox im i ty t o e nemy t e r r i tor ies . U n for tuna te ly , a va i lab le e t hnog raph ies r a re ly d ocumen t s easona l h e rd s ubd iv is ions a nd t he i r r e qu ired l a bo r , l e t a l one t he n umbe r o f a n ima ls e ach c on ta ined . P resumab ly , t he a dd i t iona l l a bor r equ i red f o r t he h usbandry o f h e rds a nd h e rd s ubd iv is ions e xceed ing e f fec t ive h erd ing l i m i ts w ou ld p ar t ia l ly o f fse tt he i n c reases i np roduc t iv i ty e xpec ted w i th l a rge rh e rds . Among t he G ab ra o fE as tA f r ica , t he l a rges t manageab le c ame l h e rd s ubd iv is ion i ss a id t o r a re ly e xceed 7 0 a n ima ls ( lo rr y 1 973 :85 , 1 974 :67 ) , a l though i nJ o rdan , o ne man ( o f ten al i t le b oy ) i ss a id t o e as i ly c on t ro l ah e rd o f5 0 c ame ls ( Köh le r 1 984 :203 ) . H oweve r , a ccord ing t o Gau th ie r -P i l te rs a nd D agg ( 1981 :14 1) , " excep tf o rd raw ing w a te rf o rt he a n ima ls , i ti sa lmos t a s e asy t oh e rd 2 00 c ame ls a s5 0 . " F rom t he a va i lab le d a ta , a n i n tu i t ive a pp rox ima t ion o ft h e u ppe r l i m i t o ft o ta l c ame l h e rd s i ze w h ich c ou ld b e managed w i th amax imum s easona l l a bo rf o rce o f 2 .5 h erde rs w ou ld s eem ing ly l i e s omewhe re b e tween 1 50 a nd 2 00 h ead , a nd i tw i l l b e 8 4

a ssumed i ns ubsequen tc a lcu la t ions t ha tt he a c tua l l i m i tl i es a round 1 50 h ead o fa l a ge a nd s ex c l asses . H oweve r , c a t t le , s heep , a nd g oa ts c an a ppa ren t ly b e managed i nl a rge rh e rds t han c ame ls w i thou t i n c reas ing r equ i red a nnua l h e rd ing l a bo r . A cco rd ing t od a ta c o l lec ted b yt he K a ramo ja D i s t r ic t V e ter ina ry O f ice ,t he m ean s i ze o f1 K a r imo jong c a t t le h e rds w as 1 7 1 h ead , e xc lud ing a n u n reco rded n umbe r o f c a lves k ep t a t c a t t le h e rd ing c amps ( Dyson -Hudson a nd D yson H udson 1 970 :112 , T ab le 3 ) . F ur the r , w h i le t he o p t imum s i ze o fK a r imo jong c a t t le h e rds i ss a id t o b e f r om 1 00 t o 2 00 h ead , i ti so n ly " w i th h und reds o f c a t t le " t ha t as i ng le h e rd mus t n ecessa r i ly b e s p l i t i n to s epa ra te u n i ts f o re f fec t ive h e rd ing ( Dyson -Hudson 1 960b :52 ) . Among t h e N g isonyoka o fn o r thwes te rn K enya , c a t t le a re o f ten k ep t i nh e rds o f 1 00 t o 1 50 a n ima ls ( Dyson -Hudson a nd M cCabe 1 985 ; M cCabe 1 984 ) . T o r ry ( 1973 :85 ) o bser ved d i sc re te g raz ing h e rds o f1 50 t o2 00 c a t t le among t he G ab ra . A l though Wa tson ( 1979 :93 ) s ugges ted t ha t o n ly 1 50 t o 2 00 s heep a nd g oa ts c ou ld b e m anaged b yo ne v i l lage h e rde ri nI r an , s im i la r ly m ixed s ma l ls tock h e rds o fu pt o3 00 a n ima ls a re n o rma l ly managed b y o ne a do lescen t h e rde r a ged 8t o 1 8 among t he N g isonyoka o f K enya ( Dyson -Hudson a nd McCabe 1 985 ; M cCabe 1 984 ) . B a r th ( 1960 :345 ; 1 961 :6 , 2 2 ) n o ted t ha t 3 00 t o 4 00 s heep a nd g oa ts c ou ld b e h e rded b y o ne s hephe rd w i thou t d ogs , e ven i nq u i te d i f f icu l t t e r ra in . Sw id le r ( 1 972 :74 ) f u r the r n o ted t ha t u p t o 5 00 s heep c ou ld b e s uperv ised b y o ne B rahu i h e rde r w i th a d og . T ony ( 1973 :85 ) o bse rved d i sc re te g raz ing h e rds o f 3 00 -400 h ead o fs heep a nd g oa ts among t he G ab ra , a l though s ma l ls tock h e rds w e re u sua l ly s ubd iv ided f o r h erd ing p u rposes when t hey e xceeded 3 00 t o 3 50 h ead , a ppa ren t ly t o d im in ish t he d i f f icu l t ies e xpe r ienced b y t h e i r y oung h e rde rs ( 1973 :105 ) . I nt he a bsence o f f u r the r d a ta , h e rd ing l a bo rc os ts w i l l b e h e ld c ons tan tf o r h e rds c on ta in ing u pt o1 50 c ame ls , 2 00 c a t t le , a nd 4 00 s heep a nd g oa ts . I tw i l l t he re fo re b e a ssumed t ha t h e rds sma l le r t han t hese p os tu la ted s i ze l i m i ts s hou ld g ene ra l ly r equ i re t he s ame amoun t o fa nnua l h e rd ing l a bo ra s l a rge r h e rds . I ng ene ra l , ah e rd s ubd iv is ion c on ta in ing o n ly 2 0a n ima ls w ou ld s t i l l r equ i re t he s ame amoun to fd a i ly f eed ing t i me a s o ne c on ta in ing 1 00 a n ima ls , r ega rd less o f a va i lab le p as turage . F ur the r , e f fec t ive m anagemen t f o r ah e rd c on ta in ing o n ly 2 0 a do lescen t a nd a du l tf ema les w ou ld s t i l l i n vo lve t he s ame s easona l h erd s ubd iv is ions a s t hose emp loyed w i th l a rge r h e rds . O n ly i nc ases o f c oopera t ive h e rd ing b e tween h ouseho lds w ou ld s ma l le r h e rds n ecessa r i ly r equ i re l e ss a nnua l h e rd ing l a bo r . H owever , i nt h e p resen t a bsence o fs u f f ic ien t d a ta t o m ode l s uch r e la t ionsh ips , t h e a nnua l h e rd ing l a bo rc os ts e s tab l ished e a r l ier w i l l b e h e ld c ons tan t f o ra l h e rds c on ta in ing f ewe ra n ima ls t han t he s i ze l i m i ts p rev ious ly n o ted . Wha t wou ld p resumab ly c hange w i th i n c reas ing ly l a rge r i n d iv idua l h e rds w ou ld b e t he f r equency o fs t ray a n ima ls a nd t he n umbe ro fl a bo rh ou rs s pen td raw ing w a te rf o r l a rge rh e rds d u r ing d ry s easons . Wh i le i ti sr easonab le t o e xpec t t ha t v a r ia t ions i nt he t i me a nd e ne rgy r e qu i remen ts o f s uch h e rd ing t asks i ns pec i f ic e nv i ronmen ts w ou ld a fec t r e la t ive p as to ra l r e tu rn r a tes a nd h ence d ec is ion mak ing , t he a ssump t ion o f a n e i gh t h our mean h erd ing d ay m us t n ecessa r i ly s uf ice f o r t he p resen t c a lcu la t ions . A s a pp l ied h e re , i td oes n o t ma t te r w he the r ah e rde rs pends e i gh th ou rs ad ay g ua rd ing , mov ing , o rw a te r ing ap as to ra l h e rd . P resumab ly , t he c umu la t ive n umbe r o f a n ima ls f eed ing i n ag i ven p a tch o f p as turage w ou ld e ven tua l ly a f fec tt he f e ed ing e f f ic iency o ft he a n ima ls h e rded i nt ha t p a tch . A s a va i lab le f o rage d ec l ined , ap o in t w ou ld b e r eached a tw h ich i n d iv idua l h e rde rs w ou ld h ave t oc hoose b e tween t he c os ts a nd b ene f i ts o fc on t inu ing t o h e rd t he i r a n ima ls i nt ha t p a tch r e la t ive t ot he c os ts a nd b ene f i ts o f mov ing t h e i r h e rds t oa l te rna t ive p as tu res . Wh i le d a i ly h e rd ing l a bo rc os ts a s c a lcu la ted h e re wou ld r ema in c ons tan t i ne i the r c ase , movemen t t o n ew p as tu res wou ld r e su l t i nt he a dd i t iona l l a bo rc os ts a ssoc ia ted w i th movemen ts o fe ncampmen t .

E ncampmen t T ime T he b as ic t r anshumance p a t te rns o fp as to ra l is ts i nt h e N ea rE as t a nd N o r th a nd E as tA f r ica m ay b e b road ly c ha rac te r ized a s i n vo lv ing h o r izon ta l a nd /o r v er t ica l p a t te rns o f s easona l r e source e xp lo i ta t ion ( J ohnson 1 969 :165 -176 ) . I n p ar t icu la r , t he l a bo r c os ts o f mov ing e ncampmen ts p r ima r i ly r e su l tf r om t he p roduc t ive r a t iona le o f mov ing h e rds t on ew l o ca t ions i n a cco rdance w i th t he c hang ing s easona l d i s t r ibu t ion a nd d ep le t ion o f p as ture a nd w a te r 8 5

r esources , a nd s easona l c l i ma t ic v a r ia t ions . C oo rd ina t ing h e rds w i th s pa t ia l ly a nd t empo ra l ly v a r iab le p as tu re a nd wa te r r esou rces may b e a ch ieved b y s easona l ly mov ing b o th h e rds a nd h ouseho lds t o n ew l o ca t ions , a nd /o r b y d ua l res idence p a t te rns i n vo lv ing t he s easona l e s tab l ishmen t o f sma l le r h e rd ing c amps i nl o ca l i t ies away f r om ma in r e s iden t ia l e ncampmen ts . S uch movemen ts may r ange f r om s ho r t moves i n vo lv ing h und reds o f me te rs t h rough s easona l m ig ra t ions o f at housand o r mo re k i lome te rs . I ti n i t ia l ly s eems r easonab le t o e xpec t t ha t b o th l e ng th o f m ig ra t ions a nd f r equency o f e ncampmen t movemen t wou ld a fec t t he r e la t ive l a bo rc os ts o f a l te rna t ive p as to ra l s t ra teg ies u nde r r eg iona l ly s pec i f ic e nv i ronmen ta l c ond i t ions . H oweve r , wh i le t he i n c lus ion o fl a bo r c os ts a ssoc ia ted w i th t he f r equency o f r equ i red movemen ts o f e ncampmen t s eems j u s t i f ied , i ti s u nc lea r t o wha t e x ten t t he t ime a nd e ne rgy e xpended b y n onhe rde rs d ur ing m ig ra t ion i s n ecessa r i ly ad i rec tc os t o fp as to ra lf eed ing s t ra teg ies . Among p as to ra l is ts , ma jo r h ouseho ld i t e ms s ee m t ob e u n ive rsa l ly t r anspor ted o n p ack a n ima ls , a nd t he o p t ion t or i de a n ima ls d u r ing l o ng m ig ra t ions i sf r equen t ly e xp lo i ted . H oweve r , i nt he c ompa ra t ive ly s hor t m ig ra t ions emp loyed b y p as to ra l is ts i nE as t A f r ica , mos t i n d iv idua ls c hoose n o tt o r i de . E ven h e re , h oweve r , p regnan t w omen , t he e l de r ly , t he s i ck , a nd t h e y oung d o r i de d u r ing m ig ra t ions ( Sa to 1 980 :61; T o r ry 1 974 :60 , 6 6 ) . Wh i le t he K a r imo jong k eep d onkeys f o r t r anspor t , t hey a ppa ren t ly d o n o t r i de t hem d u r ing m ig ra t ions ( Dyson -Hudson a nd D yson -Hudson 1 970 :100 ) . I n I r an , d onkeys , mu les a nd h o rses a re f r equen t ly u sed f o r b o th p ack ing a nd r i d ing ( Ba r th 1 96 1:7 , 1 3 , 1 5 ; H o le 1 978 : 140 ) . P resumab ly , e a r ly p as to ra l is ts c ou ld h ave c ached s easona l ly u t i l ized h ouseho ld i t ems i n t he g ene ra l r eg ions where t hey we re r equ i red , b u i l t t empo ra ry s he l te rs f r om l o ca l ly a va i lab le ma te r ia ls , a nd g ene ra l ly t r ave l led w i th f ew h ouseho ld i t ems ( Ho le 1 978 :149 -50 ) . S ign i f ican t ly , t he l a bo r a c t iv i t ies o ft hose i n d iv idua ls n o t d i rec t ly i n vo lved i nh e rd ing d u r ing m ig ra t ions i sq u i te d i ve rse . I ti n c ludes h and tasks ( e .g . , s p inn ing ) , s oc ia l iz ing , a nd mos t i mpo r tan t ly , f o rag ing i nr eg ions a l ong o r n ea r m ig ra t ion r o u tes ( Ca r ru the rs 1 9 10 :230 ; D ough ty 1 936 :37 1) . U n fo r tuna te ly , t he d eg ree t o wh ich p as to ra l d ec is ion mak ing m igh t b e i n f luenced b y t he r e la t ive c os ts a nd b ene f i ts o f c oord ina t ing h e rd a nd e ncampmen t movemen t w i th s easona l f o rag ing o ppor tun i t ies c anno t b e a dd ressed o n t h e b as is o fa va i lab le d a ta . E ven s o , t he t ime a loca ted t of o rag ing a c t iv i t ies b y n onhe rde rs d u r ing m ig ra t ions i sn o t n ecessa r i ly ac os t o fp as to ra l ism . I nt he e thnog raph ic p resen t , e conom ic a nd s oc ia l i n te rac t ion w i th s eden ta ry o r n omad ic p opu la t ions o n o r n ea r m ig ra t ion r ou tes a l so o ccu r red a t t hese t imes . A ga in , i ti s u nc lea rw he the rt he t ime a loca ted t os uch a c t iv i t ies i sn ecessa r i ly ac os to fb e ing ap as to ra l is t . F o r t hese r easons , t he l abo r c os ts o f b o th s hor t a nd l o ng -d is tance m ig ra t ions w i l l b e r es t r ic ted i nt he p resen t a na lys is t ot he t ime s pen t i nb reak ing e ncampmen ts a nd l o ad ing g ea r , a nd t ha t s pen t i nu n load ing g ea ra nd r ees tab l ish ing e ncampmen ts a tn ew l o ca t ions . I no rde rt o c a lcu la te t he t o ta l l a bo r h ou rs s pen t a nnua l ly i ne ncampmen tt a sks , i ti sn ecessa ry t oe s tab l ish t he a c tua l l abo r h ou rs r equ i red p e r move o f s pec i f ic t ype o f e ncampmen t . I n t he f o l low ing s ec t ions o n t he l a bo r c os ts o f b reak ing a nd r ees tab l ish ing e ncampmen ts , c ompa ra t ive t ime d a ta o n manya t tas a nd h e rd ing c amps i nE as t A f r ica , a nd b l ack t en t e ncampmen ts i nt h e N ea r E as t a nd N o r th A f r ica w i l l b e e xam ined .

Manya t ta a nd H erd ing C amp L abor C os ts A s p rev ious ly n o ted , p as to ra l is ts i nt he s avannas a nd d eser ts o f E as t A f r ica u t i l ize t wo b as ic t ypes o fe ncampmen ts ; manya t ta s e t t lemen ts a nd h e rd ing c amps . P hys ica l ly , manya t ta s e t t lemen ts c ons is t o f ac l us te ro fi n d iv idua l h u ts a nd s tock c o r ra ls , a nd r ep resen t t he c ommon e ncampmen t o f s eve ra l f am i l ies . F o r e xamp le , S a to ( 1980 :47 ) n o ted t ha t i nh i s s urvey o f 2 2 R end i l le manya t tas , a n a ve rage s e t t lemen t c ons is ted o f2 2 .2 h u ts , w i th a r ange b e tween 9 a nd 5 0 h u ts . Among t he Gab ra , manya t tas c ons is ted o f a n a ve rage o f 1 7 h u ts ( Tor ry 1 974 :60 ) , a l though s e t t lemen ts c on ta in ing a s many a s 4 2 h u ts we re n o ted ( To r ry 1 973 :196 ) . T he p hys ica l l a you t o f manya t tas i n E as t A f r ica v a ry w ide ly , p resumab ly a s ap r ima ry f unc t ion o f t he h e rd a n ima ls i n vo lved , t he i n tens i ty o f c a rn ivore p reda t ion , d e fense r equ i remen ts f r om p o ten t ia l r a ids , a nd l o ca l t opog raph ic r e l ie f . D assane tch manya t tas e s tab l ished i na reas s ub jec t t o r a ids b y n e ighbo r ing p opu la t ions c ons is t o f ac en t ra l c l us ter o f h u ts s u r rounded b y b rush s t ock c or ra ls ( G i f fo rd 1 977 :71-3 , 1 978 :84 ) . Among B agga ra c a t t le 8 6

n omads , manya t tas c ons is t o f a n o u te r r i ng o f h u ts s ur round ing i n terna l b rush c or ra ls ( Cunn ison 1 966 :65 , 6 9 -70 ) . Among t he Gab ra , manya t tas c ons is to f ac i rc le o fh u ts w i th c ame l c o r ra ls c ons t ruc ted a round t he i r o u te r c i rcu i t , a nd sma l ls tock c o r ra ls c ons t ruc ted a l ong t he i r i n ne rc i rcu i t ( Tony 1 973 :196 ) . E ven mo re e l abo ra te manya t tas a re c ons t ruc ted b y t he R end i l le ( Sa to 1 980 :45 -7 ) . T hese c ons is t o f ac en t ra l c i rc le f o r s oc ia l iz ing wh ich i ss ur rounded b y a b rush f ence . A round t h is c en t ra l c i rc le , b rush s t ock c o r ra ls a re c ons t ruc ted . H u ts a re t hen e rec ted i n ar i ng a round t he s t ock c o r ra ls , w i th t he e n t i re s e t t lemen t s ubsequen t ly e nc losed b y a n o u te rt ho rn f ence . B y c on t ras t t o manya t tas , h e rd ing c amps a re l e ss s ubs tan t ia l , g ene ra l ly c ons is t ing o f i n d iv idua l s he l te rs , s imp le w indb reaks , o r i n fo rma l " s leep ing a reas , " a nd a ssoc ia ted s t ock e nc losures ( G i f fo rd 1 977 :76 , 1 978 :88 ; S a to 1 980 :53 ; T o r ry 1 973 :178 ) . S ince t hese a re p r ima r i ly o ccup ied b y af ew h e rde rs , mos t h ouseho ld i t ems a re l e f t a tt he manya t tas . S he l ter L abo r . L abo rd a ta o n t he e rec t ion o rc ons t ruc t ion o ft he v a r ious s he l te rs u sed b y p as to ra l is ts i nE as t A f r ica i sr a re ly r epo r ted i nt he a va i lab le e t hnog raph ies . A cco rd ing t oW i l l iam I . T o r ry ( pe rsona l c ommun ica t ion 1 985 ) , e ach ma t a nd s k in " t en t " o f t he Gab ra r equ i red a pp rox ima te ly t wo h ou rs t o e rec t a f te r b e ing u n loaded , w i th t wo t ot h ree p eop le c on t r ibu t ing l a bo r . S im i la r ly , t hese " t en ts " c ou ld b e d i sassemb led i n a bou t o ne h our , a nd p acked a nd l o aded i na pp rox ima te ly 4 5 m inu tes , a ga in w i th t wo t o t h ree p eop le c on t r ibu t ing l a bo r . T o r ry n o ted , h owever , t ha t t he l a bo r o f t he women i n vo lved was o f ten i n te r rup ted b y o the r t a sks . P resumab ly , s im i la r l abo rc os ts w ou ld a pp ly t ot he ma t a nd s k in t en ts u sed b yt he D assane tch a nd R end i l le . I n te res t ing ly , t he t ime c os ts s ugges ted b y T o r ry f o r t he e rec t ion o f Gab ra " t en ts " a re s eem ing ly c l ose t ot hose r epo r ted f o r t he e rec t ion o f N ue r h u ts b y E vans -P r i tcha rd ( 1940 :65 6 ) , b u t g rea te r t han t hose r epo r ted f o r T ua reg h u ts b y N ico la isen ( 1963 :353 , 3 59 ) . U n l ike t he G ab ra , R end i l le , o r D assane tch , t he N ue ro fS udan c ons t ruc t sma l l , r ound , g rass o r r eed h u ts p l as te red w i th d ung . L oca l ly p rocu red g rasses a nd r eeds a re t i gh t ly p acked i n a n u p r igh t p os i t ion i n an a r row , c i rcu la rt r ench t of o rm wa l ls . T hese a re t hen b ound t oge the r a tt he i rt ops a nd e x te rna l ly p las te red w i th d ung t o f o rm a h u t . A cco rd ing t o E vans -P r i tcha rd ( 1940 :66 ) , " t hese l i gh t dwe l l ings c an b e e rec ted i n af ew h ou rs . " U n fo r tuna te ly , t he n umbe r o f l a bo re rs i n vo lved was n o t s t a ted . Wh i le s eve ra l T ua reg g roups a l so c ons t ruc t r ound , d ome -shaped ma tt en ts s im i la rt ot hose o f t he Gab ra , R end i l le , a nd D assane tch , mos t ma t a nd s k in t en ts o f t he A y r T ua reg h ave a r ec tangu la r g round p lan a nd a d ome -shaped o r b a r re l v au l ted s upe rs t ruc ture ( N ico la isen 1 963 :351 , 3 56 -58 ) . H owever , w h i le t he i r s hape i sd i f fe ren t , t hese a re § t i l l c omposed o f ma ts a nd /o r s k ins d raped o ve r af r ame o f c urved wooden s uppor ts wh ich a re l a shed t oge the r . N ico la isen s umma r ized t he l a bo r i n vo lved i n e rec t ing a n A y r T ua reg ma t t en t a s f o l lows ( 1963 :353 ) : T he wo rk s eems t o b e v e ry e asy t o t he women who measu re n o d i s tances . T hey s eem t oc a r ry o u tt he i r wo rk a u toma t ica l ly . Ih ave n o t b een a b le t oc on t ro l t he amoun t o f t ime n eeded f o r o ne o r t wo w omen t o e rec t a ma t t en t o ft he t ype men t ioned , f o r t he women a re f r equen t ly i n te r rup ted i n t he i r wo rk b y many o the r h ouseho ld d u t ies , b y t he i r c ry ing b ab ies , o r b y v i s i to rs f r om o the r c amps , b u t Ib e l ieve t ha t o ne w oman c an e rec t t h is e l abo ra te t en t s t ruc tu re i n a bou t h a l f a n h ou r . E ven w i th a dd i t iona l t ime f o r u n load ing , s t ow ing , a nd s e t t ing u p h ouseho ld i t ems , T ua reg e ncampmen t c os ts s eem s omewha t l e ss t han t hose g i ven f o rt he Gab ra . F ur the r , T ua reg ma t t en ts a ppa ren t ly r equ i re mo re l a bo r t o e rec t t han t he i r s k in t en ts , a s was f u r the r s t a ted b y N ico la isen i nh i sc ompa r ison o ft he r e la t ive me r i ts o fT ua reg ma t a nd s k in t en ts ( 1963 :359 -60 ) : l ti s more d i f f icu l t t o e rec t t he ma t t en t t han t he s k in t en t , b u t s k in t en ts v ery o f ten c ons t i tu te a h eav ie rb u rden . T he s hee t o fs k in , o r mo re c o r rec t ly l e a the r , i sh eav ie r t han t he c ove r ing o f ma ts , a nd t hough i ti st r ue o f many s k in t en ts t ha t t hey h ave a l e ss e l abo ra te s t ruc ture t han t he ma t t en ts , t he f o rme r n eed a s ys tem o f p egs a nd p o les t o f as ten t he s hee t , a nd t hese h ave t o b e t r anspor ted e xcep t whe re s u i tab le n a tu ra l b oughs a re a bundan t . T he re a re a dvan tages a nd d i sadvan tages i n ma tt en ts a s w e l l a s i ns k in t en ts . D u r ing t he 8 7

v e ry h o t s eason t he ma t t en t i s b y f a r t he b es t t ype o f dwe l l ing ; i ti s c ompa ra t ive ly h i gh a nd t he ma t c ove r ing i sn o t h ea ted t o a ny e x ten t b y t he s un . D u r ing t he r a iny s eason t he ma t t en t i sl e ss p rac t ica l t han t he s k in t en t . R a in may f a l l s o h eav i ly a nd a bundan t ly t ha t i tp ene t ra tes t he ma tc ove r ing , a nd e ven i fi td oes n o t i tw i l l b e n ecessa ry t or emove a l ma ts a f te r ah eavy s howe r i n o rde rt od ry t hem i nt he s un . T he s k in t en t i sn o rma l ly e rec ted q u icke rt han t he ma t t en t , i tp ro tec ts e f f ic ien t ly a ga ins t r a in , b u t i s , o n t he o the r h and , mos t u np leasan t d u r ing t he h o t s eason . S tock E nc losure L abor . E ven l e ss c onc ise i st he a va i lab le d a ta o n t he l a bo r r equ i red t o c ons t ruc t a nd ma in ta in t ho rnbush s t ock e nc losures . P resumab ly , t he s i ze , c ons t ruc t ion , a nd i n te rna l d i v is ions o f e ach s t ock e nc losu re wou ld v a ry a cco rd ing t o s uch f ac to rs a s a va i lab le ma te r ia ls , t he a n ima ls i n vo lved , s easona l h e rd s ubd iv is ions , i n tens i ty o f c a rn ivo re p reda t ion , l e ng th o fo ccupancy , a nd s i ze o f h e rd . A s o n ly o ne e xamp le , a t l e as t t h ree d i s t inc t e nc losu re p a t te rns a re s easona l ly emp loyed b y t he R end i l le a tt he i r c ame l h e rd ing c amps a s ar e su l t o f c hanges i nh e rd s ubd iv is ions ( Sa to 1 980 :51-2 ) . U n for tuna te ly , g i ven t he a bsence o fa va i lab le t ime a loca t ion d a ta , ac onc ise mode l l ing o f s uch s easona l v a r ia t ions a s t hey w ou ld a f fec t e ncampmen t l a bo rc os ts i sn o t c u r ren t ly p oss ib le . T ony ( 1973 :137 ) d i d n o te t ha tt he Gab ra t ook " no mo re t han h a l f ad ay "t oe rec tt ho rnbush e nc losu res a t a manya t ta , w i th 3 0 m inu tes t oa n h ou r s ubsequen t ly s pen t d a i ly i n mend ing o r r e in fo rc ing t hose e nc losu res o ccup ied b y c ame ls ( To r ry 1 973 :86 , 1 37 ) . U n for tuna te ly , t he n umbe ro fw o rke rs i n vo lved i nt hese a c t iv i t ies was n o t r eco rded . I nt e rms o fa c tua l l a bo rc os ts , P e te r J ones ( pe rsona l c ommun ica t ion 1 985 ) h as o bserved t ha t among t he Maasa i , t h ree t o f ou r ma les c ou ld c ons t ruc t as ubs tan t ia l e nc losu re f r om s c ra tch i nt h ree t of ou r h ou rs , a l though s ubsequen t r e in fo rcemen t w ou ld o ccur o ve rt he s pan o f af ew d ays . I nt he a bsence o ff u r the r o bserva t iona l d a ta , t hese c ons t ruc t ion a nd ma in tenance c os ts w i l l b e emp loyed i ns ubsequen t l a bo rc a lcu la t ions f o rt he s t ock e nc losu res s pec i f ica l ly a ssoc ia ted w i th manya t tas . P resumab ly , l e ss l a bo rw ou ld b e e xpended i nt he c ons t ruc t ion o f l e ss -subs tan t ia l , s ho r t t e rm e nc losu res , a s when R end i l le h e rde rs e rec t o ve rn igh t c o r ra ls w hen t hey a re u nab le t o r e tu rn t ot he i r ma in e ncampmen ts ( Sa to 1 980 :39 ) . F u r ther , i ti se xpec ted t ha t t he f r equency o f c a rn ivo re p reda t ion wou ld v ary s easona l ly , t hus a f fec t ing t he n a ture o f r equ i red s t ock e nc losures a t h e rd ing c amps . F o r e xamp le , c a rn ivo res i nR end i l le t e r r i tor y , i n c lud ing j a cka ls , h yenas , l i ons , a nd l y caons , s easona l ly t r ack h e rds o f w i ld h e rb ivo res i n to h i ghe r e l eva t ions d ur ing t he d ry s easons , t hus a bandon ing t he a r id l ow lands whe re t he R end i l le h erd t he i r c ame ls ( Sa to 1 980 :2 1-2 ) . A s ar esu l t , w hen t he R end i l le a re mos tf r equen t ly mov ing t h e i rc ame l h e rd ing e ncampmen ts , c a rn ivo re p reda t ion i sl ow a nd s ubs tan t ia l e nc losures wou ld b e l e ss c r i t ica l . H oweve r , d u r ing t he r a iny s easons when c ame l e ncampmen ts a re moved l e ss o f ten , c a rn ivo re p reda t ion wou ld b e mo re f r equen t , a nd mo re s ubs tan t ia l e nc losures w ou ld b e r equ i red . T hese s easona l d ynam ics w i l l n eed t o b e a ccoun ted f o r i nc a lcu la t ing t he l a bo r e xpended i nt he c ons t ruc t ion o f v a r ious s easona l h e rd ing e ncampmen ts . H oweve r , f o r t he mo re p e rmanen t manya t tas , e ncampmen t l a bo rc os ts s hou ld r ema in r e la t ive ly c ons tan t . Manya t ta L abor C os ts . S ince manya t tas r ep resen t ac l us te red s e t t lemen t o f i n d iv idua l h u ts a nd c o r ra ls , t he p ropor t iona l c on t r ibu t ion o f e ach f am i ly 's l abo r t o e s tab l ish ing s uch s e t t lemen ts wou ld g ene ra l ly c ons is t o ft he t ime t hey s pend i ns e t t ing -up t he i r o wn h u t , a nd t he t ime s pen t i nc ons t ruc t ing a nd s ubsequen t ly ma in ta in ing s ubs tan t ia l t ho rnbush e nc losu res f o r t he i r s t ock . A ssum ing t ha t e ach b reak ing a nd r ees tab l ishmen t o f a ma t a nd s k in h ouseho ld r equ i res t he l abo r o f t h ree p eop le f o r f our h ou rs , a pp rox ima te ly 1 2 h ours o f t o ta l h u t l abo r w ou ld b e r equ ired f o r e ach c omp le te movemen t o f ah ouseho ld . F ur the r , s ubs tan t ia l t ho rnbush s t ock e nc losu res c an a ppa ren t ly b e b u il tw i th a n i n ves tmen t o f 9t o 1 6 h ou rs o ft o ta l l a bo r , w i th a p os tu la ted mean o f 1 2 .5 h ours . T hus , e ach manya t ta movemen t w ou ld r equ i re a pp rox ima te ly 2 4 .5 h ou rs o ft o ta l e ncampmen t l abo r p e r h ouseho ld . F o l low ing T ony 's d a ta o n c ame l e nc losure ma in tenance , o ne a dd i t iona l h our p e r d ay o f e ncampmen t wou ld s ubsequen t ly b e r equ i red . H e rd ing C amp L abo rC os ts . B yc on t ras tt o manya t tas , t he l a bo rc os ts o fe s tab l ish ing l e ss p e rmanen t h e rd ing c amps s hou ld p resumab ly b e much l e ss . I nt he a bsence o fo bser ved l a bo r d a ta , i tw i l l b e a ssumed t ha t r ough ly t wo th i rds o f t he t ime i n ves ted i n s ubs tan t ia l s t ock e nc losu res a t manya t tas w ou ld b e r equ i red f o r we t s eason h erd ing c amp e nc losures , o r a pp rox ima te ly e i gh t h ou rs o ft o ta l l abo r . A n a dd i t iona l f our h ou rs o fl a bo rf o rt he c ons t ruc t ion o f 8 8

as imp le s he l te rf o rt he h e rde r sw i l l a l so b e a ssumed ,t h us r a is ing t he l a bo rc os ts o fe s tab l ish ing aw e ts eason h e rd ing e ncampmen tt o1 2 h ours o ft o ta l l a bo r . I na dd i t ion , i tw i l l b e a ssumed t ha t f o r e ach s ubsequen t d ay s pen t a t a we t s eason e ncampmen t , 4 5 m inu tes o f c o r ra l m a in tenance w ou ld b e r equ i red d a i ly . F i na l ly , f o r d ry s eason e ncampmen ts o ccup ied f o ro n ly o ne o rt wo d ays u nde rc ond i t ions o fr e duced p reda t ion , af our h ou r e nc losu re l a bo r c os t w i l l b e a ssumed , w i th o ne a dd i t iona l h ou r a lowed f o rt he c ons t ruc t ion o f as imp le s l eep ing a rea . N o e nc losure ma in tenance c os ts s hou ld e x is t , s i nce t hese w ou ld b e a bandoned a f te ro ne o rt wo d ays . T hese c a lcu la ted e ncampmen t c os ts w i l l b e s ubsequen t ly a pp l ied t ot he s easona l h e rd a nd r e s idence m ovemen ts o ft he Gab ra a nd R end i l le i no rde rt oe s tab l ish a pp rox ima te a nnua l e ncampmen t c os ts among E as t A f r ican p as to ra l is ts . H owever , i ti sf i r s t n ecessary t o b r ie f ly d i scuss c e r ta in o bse rved v a r ia t ions b e tween t he manya t ta movemen ts a nd a ssoc ia ted h e rd ing p a t te rns o ft he G ab ra a nd R end i l le . F requency o f Manya t ta Movemen ts . Wh i le S a to ( 1980 :50 ) s t a ted t ha t e ach R end i l le manya t ta r ema ins i no ne l o ca t ion f o ra n a ve rage o ft wo t ot h ree mon ths , t he G ab ra a re s a id t o r e loca te t he i r manya t tas e ve ry t h ree t os i xw eeks , mov ing 1 2 t o 1 5 t i mes ay ea r ( 1 3 .5 mean ) w i th a r ange b e tween 9 a nd 1 7 moves ( To r ry 1 973 :218 -19 ) . T hese v a r ia t ions i n manya t ta m ovemen t s eem ing ly r e f lec t v a r ia t ions i nt he p re fe r red h e rd ing p a t te rns o f t he Gab ra a nd R end i l le . Wh i le t h e Gab ra p r ima r i ly h e rd t he i rc ame ls f r om manya t tas a nd t he i rs ma l ls tock f r om h e rd ing c amps ( To r ry 1 973 :66 , 7 9 ) , t he R end i l le p r ima r i ly h e rd mos t o ft he i r l i ves tock f r om h e rd ing c amps f o r much o ft h ey ea r( Sa to 1 980 :56 ) . I na dd i t ion t ot he d ep le t ion r a te o fa va i lab le p as tu rage a s ar e su l t o fh usband ing s eve ra l h e rds i n as i ng le l o ca l i ty , af u r ther f ac to r i n f luenc ing t he f r equency o f Gab ra manya t ta m ovemen ts i s s a id t o b e t he i n fes ta t ion o f c o r ra ls a nd l i ves tock e xc remen t w i th t i cks ( Rh ip icepha lus p u lche l lus ) w h ich b ea r ad i sease o r d i seases l e tha l t o c ame ls . T hese t i cks c anno t b e c on t ro l led b y p e r iod ica l ly c l ean ing t he c o r ra ls , s i nce t hey l i ve p a r t o ft he i r l i f e c yc le b enea th t he s o i l ( To r ry 1 973 :82 -3 , 1 974 :67 ) . T he R end i l le a l so move t he i r manya t tas t o e scape o u tb reaks o ft i cks ( Sa to 1 980 :50 ) w h ich a re s im i la r ly s a id t o c ause l e tha l i n fec t ions among t he ir c ame ls ( J ones 1 984 :126 ) . T hus , f o r r easons o f b o th p as ture d ep le t ion a nd t i ck i n fes ta t ion , i ti se xpec ted t ha t R end i l le manya t tas w ou ld b e moved a t l e as t o nce e ver y t wo m on ths w hen t he i rc ame l h e rds a re p resen t .

A nnua lG ab ra E ncampmen tC os ts A s t he Gab ra p re fe rt op r ima r i ly h e rd t he i rc ame ls f r om manya t tas , a nd s i nce t hese a re m oved a pp rox ima te ly 1 4 t i mes a y ea r a t 2 4 .5 h ours o f l a bo r p er movemen t , e ach Gab ra h ouseho ld s pends a pp rox ima te ly 3 43 h ou rs ay ea r i n manya t ta e ncampmen t l a bo r . S ince o ne a dd i t iona l h our p e rd ay o fs ubsequen t e ncampmen t w ou ld b e r equ i red f o r c o r ra l ma in tenance ( 3 65 d ays -1 4 d ays o f movemen t =3 5 1 s t a t iona ry d ays ) , a n a dd i t iona l3 5 1 h ou rs o fl a bo rw ou ld b e s pen t a tt hese manya t tas . F o r a pp rox ima te ly f ou r mon ths o u t o f e very y ea r ( r ough ly 1 23 d ays ) , c ame l h e rd s ubd iv is ions r equ i re we t s eason h e rd ing a t s epa ra te e ncampmen ts ( To r ry 1 973 :89 -90 ) . A ssum ing t ha t G ab ra we t s eason h e rd ing c amps w ou ld b e moved o nce e very f i ve d ays a s among t he R end i l le ( Sa to 1 980 :39 ) , 2 5 moves a t 1 2 h ours o f l a bo r p e r m ove w ou ld c os t a G ab ra h erde r o r h erde rs a pp rox ima te ly 3 00 h ou rs o f f u r the r e ncampmen t c os ts . S ince a n a dd i t iona l 4 5 m inu tes p er d ay o fs ubsequen t w e t s eason e ncampmen t w ou ld b e r equ i red f o r e nc losu re ma in tenance ( 123 d ays -2 5 d ays o f movemen t =9 8 s t a t iona ry d ays ) , a pp rox ima te ly 7 4 h ou rs o fa dd i t iona l l a bo rw ou ld b er equ i red . O ve r ao ne y ea r p e r iod , t he t o ta l Gab ra e ncampmen t l a bo r w h ich may b e a t t r ibu ted t o p as to ra l r equ i remen ts wou ld amoun tt o1 , 068 h ou rs s pen t i n at o ta l o fo n ly 3 9 a nnua l m oves .

8 9

A nnua lR end i l le E ncampmen tC os ts F a r mo re c omp lex i st he h e rd ing p a t te rn r eco rded f o rt h e R end i l le o ve r a1 5 m on th p e r iod o fo bse rva t ion ( Sa to 1 980 :55 -6 ) . D ur ing t h is p e r iod , c ame ls w e re h e rded f r om manya t tas f o r o n ly 5 .5 mon ths , r ough ly 3 7% o ft he p e r iod o fo bse rva t ion . H oweve r , f o r a pp rox ima te ly h a l f o f t h is t ime , o n ly b aggage c ame ls a nd l a c ta t ing f ema les w i th i n fan t y oung , r ep resen t ing 8 11 % o f a n a ve rage R end i l le c ame l h e rd , w e re a c tua l ly h e rded f r om manya t tas ,wh i le t he r e s t w e re h usbanded a t c ame l h e rd ing c amps ( Sa to 1 980 :36 -7 ) . F or t he o ther 9 .5 mon ths o f o bse rva t ion , a l c ame ls w e re k ep t a th e rd ing c amps ( Sa to 1 980 :5 -6 ) , a l though b aggage c ame ls w e re p resumab ly s t i l l r e ta ined a t t he manya t tas . T he p ropor t ion o f R end i l le manya t ta e ncampmen t c os ts w h ich may b e d i rec t ly a t t r ibu ted t ot he i r h usbandry o fc ame ls w i l lt h e re fo re b e r e s t r ic ted i ns ubsequen t c a lcu la t ions t ot he 4mon ths i nw h ich c ame l h e rds a nd s ubd iv is ions a r e a c tua l ly h e rded f r om manya t tas . A ssum ing t ha t R end i l le manya t tas w ou ld b e moved a tl e as to nce e ve ry t wo m on ths w hen c ame l h e rds a re p resen t , t wo movemen ts o f amanya t ta m ay b e a t t r ibu ted t oc ame l h usband ry c os ts . A t 2 4 .5 h ours p e r move , 4 9 h ou rs o f e ncampmen t l a bo r w ou ld b e s pen t p e r R end i l le h ouseho ld . S ince o ne h ou ro fl a bo r p e rd ay o fs ubsequen t e ncampmen tw ou ld b e r e qu i red f o r e nc losu re ma in tenance ( 135 d ays -2movemen td ays = 1 33 s t a t iona ry d ays ) , a n a dd i t iona l 1 33 h ou rs o fl a bo rw ou ld b e e xpended a tt hese manya t tas . When S a to 's ( 1980 : 10 ) mon th ly c l ima t ic s umma ry f o rt h e r eg ion o ccup ied b yt h e R end i l le i sc omb ined w i th h i sd a ta o nt he i rs easona l h usband ry o fc ame ls a th e rd ing c amps ( 1 980 :55 -6 ) , i ti s a ppa ren t t ha t c ame l h e rds a nd s ubd iv is ions a re h e rded a t d ry s eason c amps f or a pp rox ima te ly 1 66 d ays p e r y ea r ( r ough ly 5 .5 mon ths ) , a nd a t w e t s eason c amps f o r a pp rox ima te ly 1 37 d ays p e ry ea r ( r ough ly 4 .5 mon ths ) . S ince w e t s eason c ame l h e rd ing c amps a re moved o n a ve rage o nce e ve ry f i ve d ays ( Sa to 1 980 :39 ) , a pp rox ima te ly 2 7 moves a t 1 2 l a bo r h ou rs p e r move amoun ts t o 3 24 h ou rs o f w e t s eason e ncampmen t c os ts . S ince a n a dd i t iona l 4 5 m inu tes p e r d ay o fs ubsequen t w e t s eason e ncampmen t w ou ld b e r e qu i red f o r e nc losure ma in tenance ( 137 d ays -2 7 movemen t d ays = 10 s t a t iona ry d ays ) , a pp rox ima te ly 8 3 h ou rs o fa dd i t iona l l a bo rw ou ld b e s pen t . D u r ing t he 1 66 d ays o fd ry s eason e ncampmen t , a pp rox ima te ly 1 00 moves a t f i ve h ours p e r move a dds af u r the r 5 00 h ours t o t he i r a nnua l e ncampmen t c os ts . O ve r ao ne y ea r p e r iod , t he t o ta l R end i l le e ncampmen t l a bo rw h ich may b e a t t r ibu ted t o c ame l h usband ry amoun ts t o1 , 089 h ou rs s pen t i n at o ta lo f1 29 a nnua l moves . B yc ompa r ison , t he a nnua l e ncampmen t l a bo r c os t c a lcu la ted a bove f o r t h e Gab ra ( 1 ,068 h ou rs ) i so n ly 2 1 h ou rs l e ss t han t ha t c a lcu la ted f o rt he R end i l le . D ue t ot he a nnua l h e rd ing a nd e ncampmen t v a r ia t ions n o ted e a r l ie r b e tween t he Gab ra a nd R end i l le , t he c ompa rab i l i ty o f t he i r a nnua l e ncampmen t c os ts i sq u i te s t a r t l ing . Am ean a nnua l e ncampmen t l a bo rc os t o f 1 , 079 h ou rs a s d e r ived f r om t he Gab ra a nd R end i l le c a lcu la t ions w i l l t he re fo re b e u sed i n s ubsequen t r e fe rence t o p as to ra l h e rd ing e ncampmen t c os ts w hen b rush s t ock e nc losures a re r e qu i red . Wh i le p r ima r i ly b ased u pon c ame l s t ra teg ies , i tw i l l b e a ssumed t ha t s im i la re ncampmen tc os ts o ccu ri nc a t t le , s heep a nd g oa t s t ra teg ies .

B lack T en tL abo rC os ts Movemen ts o fb l ack t en t e ncampmen ts i nN or th A f r ica a nd t he N ea r E as t d o n o t a lways i n vo lve t he t r anspor ta t ion o fa l h ouseho ld b e long ings , n o rd o t hey a lways i n vo lve t h e u se o f l a rge t en ts . L a rge t en ts a re p r ima r i ly u sed d ur ing w in te r mon ths , a nd a re o n ly r equ i red i no the r s easons f o r p u rposes o fs oc ia l iz ing a nd g ues t a ccommoda t ion . F o r m uch o ft he y ea r , s ma l le r t en ts o rp a r ts o ft he l a rge rw in te rt en ts a re u sed i n s tead , a nd e ven t hese a re u nnecessa ry i ft he w ea the r i st empera te ( Co le 1 975 :41 , 4 4 ; D ough ty 1 936 :257 , 2 65 -266 ; J enn ings -B ram ley 1 906 :258 , 1 907a :26 ; M us i l 1 928 :72 -73 ) . F or e xamp le , o n l o ng m ig ra t ions d ur ing t he s umme r mon ths , t he Rwa la b edou in o n ly p i tch s ma l l t en ts , a nd t hen o n ly i ft hese a re n eeded b y c h i ld ren ( Mus i l 1 928 :72 ) . T he K abab ish b edou in o ft he S udan w i l l o n ly p i tch at en t r o o f d u r ing m ig ra t ions ( Asad 1 970 :38 ) , w h i le t he A l Mu r rah o n ly p i tch a s i ng le t en t w a l l a s aw ind b reak d u r ing t he t empera te w ea the r o f t he f a l l s eason a nd d u r ing m ig ra t ions ( Co le 1 975 :41 , 4 4 ) . S im i la r ly , t he s heep a nd g oa t h e rd ing B asse r io fI r an e rec t s ma l le rb l ack t en ts d ur ing m ig ra t ions , o rn one a ta l i ng ood w ea the r( Ba r th 1 96 1 : 1112 ) .

9 0

When l a rge r t en ts a nd o the r h ouseho ld i t ems a re n o t r equ ired , t hey a re s t o red o r " c ached " a t s pec i f ic l o ca t ions . Among t he A l Mur rah , t en ts a nd h eav ier h ouseho ld i t ems a re c ached a tt he i r s ummer we l ls t h roughou t t he f a l l s eason ( Co le 1 975 :4 1) . Among t he b edou in o f S ina i , w in te r t en ts we re o f ten s to red i n r ockshe l te rs f o r a l b u t t he f our mon ths o f t he i r r equ ired u se ( Jenn ings -Bram ley 1 906 :258 ; 1 907a :26 ) . D ough ty ( 1936 :322 -323 ) a l so r e co rded t ha t Moah ib b edou in u sed a" sec re t " c ave i nt he H a r ra b asa l t d ese r t o fs ou the rn S y r ia f o r s easona l ly s to r ing t he i r e xcess h ouseho ld i t ems a nd f ood , a nd f u r ther n o ted t ha t o ther b edou in g roups s easona l ly c ached t he i r w in te r t en ts a nd e xcess h ouseho ld i t ems u nde r l ow m ounds o fd ry d eser t s and . B edou in a re a l so k nown t ob u i ld sma l l , s i ng le room s t o rage s t ruc tu res o f u nhewn s t ones o r m udb r icks f o r s easona l ly c ach ing h ouseho ld i t ems . D ough ty ( 1936 :323 ) o bse rved a nc ien t e xamp les o f s uch s t ruc tures i n S ina i , wh i le A l ison B e t ts ( pe rsona l c ommun ica t ion 1 984 ) r e cen t ly o bse rved s uch s t ruc tu res c u r ren t ly i nu se i nt he b asa l t d eser ts o fn o r theas te rn J o rdan . S ou theas t o f t he o as is o f H a ' i l i nn o r the rn S a 'ud i A rab ia , W inne t t a nd R eed ( 1973 :56 ) f ound s eve ra l a nc ien t e xamp les o f s uch s t o rage s t ruc tures . I n t he r ubb le o f o ne , aT hamud ic i n sc r ip t ion r ough ly d a t ing t o t he t h i rd c en tury A .D . a nnounced " The s t o res o f S auka t b . M as ika t " (W inne t t a nd R eed 1 973 :56 ) . P oss ib le a nc ien t e xamp les h ave a l so b een n o ted i n L u r is tan ( G i lber t 1 983 :113 ) . T he a n t iqu i ty o f s uch p as to ra l p rac t ices r ema ins u ndocumen ted , a l though t he s easona l s t o rage o r c ach ing o f e xcess h ouseho ld i t e ms a nd g ra in s upp l ies a t s pec i f ic l o ca t ions s hou ld b e e xpec ted i ne a r l ie r p e r iods a s w e l l . A s ar esu l t o fs uch s t o rage , t he l a bo r r equ i red i n mov ing b l ack t en t e ncampmen ts s hou ld v a ry s easona l ly . U n for tuna te ly , o n ly af ew s cho la rs h ave d ocumen ted t he t ime r equ i red t o e rec t , o r b reak d own a nd l o ad , ac omp le te b l ack t en t e ncampmen t . J enn ings -B ram ley ( 1907a :30 ) s imp ly n o ted t ha t among t he b edou in o f S ina i , " s t r ik ing a nd p i tch ing t he t en ts c an b e d one i nt wo h ou rs ." B a r th ( 196 1:15 ) s im i la r ly n o ted t ha t among t he B asse r i o f s ou the rn I r an , " t he t o ta l p r ocess o fb reak ing c amp may t ake a bou t 11 / 2 h ours . " S ince e ach d i sman t l ing o fc amp i mp l ies al a te r e rec t ion o n an ew s i te , t he a va i lab le d a ta w ou ld s ugges t t ha t a pp rox ima te ly f ou r h ou rs p e r move o fc omp le te h ouseho ld wou ld b e r equ i red . A cco rd ing t o D ough ty ( 1936 :351) , sma l le r s umme r t en ts c ou ld b e p i tched i no n ly t h ree m inu tes . A l low ing a n a dd i t iona l 2 7 m inu tes f o r p ack ing a nd l o ad ing , o r u n load ing a nd r ees tab l ish ing a r educed q uan t i ty o f h ouseho ld i t ems , e ach i n s tance o f b reak ing o r r e es tab l ish ing sma l le r e ncampmen ts w ou ld r equ i re a pp rox ima te ly 3 0 m inu tes . T hus , e ach m ove a nd r ees tab l ishmen t o f a n e ncampmen t c ons is t ing o f r educed h ouseho ld i t ems a nd i n vo lv ing sma l le rt en ts o rt en t s ec t ions wou ld s eem ing ly r equ i re a pp rox ima te ly o ne h ou r . At h i rd p e rmu ta t ion o f e ncampmen t l a bo r r equ i remen ts w ou ld r esu l t d u r ing p e r iods o f l o ng -d is tance m ig ra t ion . A t s uch t imes , an ew e ncampmen t wou ld h ave t o b e e s tab l ished n i gh t ly . H oweve r , wh i le a l l h ouseho ld b e long ings w ou ld h ave t o b e u n loaded n i gh t ly , o n ly r e qu i red h ouseho ld i t ems w ou ld n eed t o b e u npacked , a nd sma l le r t en ts o r t en t s ec t ions w ou ld g ene ra l ly s u f f ice f o rs l eep ing f ac i l i t ies . I nt he a bsence o fd i rec t o bser va t iona l d a ta , i tw i l l b e a ssumed t ha t e ach d i sman t l ing a nd r ees tab l ishmen t o f ab l ack t en t e ncampmen t d ur ing m ig ra t ions wou ld o n ly r equ i re h a l f o f t he t ime e s tab l ished a bove f o r d isman t l ing a nd r e es tab l ish ing c omp le te h ouseho lds , o ra pp rox ima te ly t wo h ou rs o fl abo rp e rd ay o fm ig ra t ion . T hese f i gures , h oweve r , d o n o t r e f lec t t he t o ta l h ours o f l abor r equ i red p e r move o f e ncampmen t , s i nce mo re t han o ne p e rson wou ld p resumab ly b e e ngaged i ns uch a c t iv i t ies . Wh i le a l l membe rs o f ah ouseho ld may c on t r ibu te l abo r t o b reak ing a nd r ees tab l ish ing e ncampmen ts , t he t ask p r ima r i ly f a l ls u pon t he w omen o f ah ouseho ld . O lde r ma les p r ima r i ly c on t r ibu te l abo r o n ly i n mov ing , l o ad ing , a nd u n load ing t he h eav ies t o b jec ts ( Asad 1 970 :45 ; B a r th 1 96 1 :16 ; C o le 1 975 :64 ; D ickson 1 951:73 ; D ough ty 1 936 :257 -258 ; Mus i l 1 928 :62 ) . U n fo r tuna te ly , t he a c tua l n umbe ro fi nd iv idua ls i n vo lved i ns uch e ncampmen t a c t iv i t ies h as n o t b een q uan t i f ied i nt he a va i lab le e thnog raph ies . I nt he a bsence o f d i rec t o bserva t iona l d a ta , i tw i l l b e a ssumed t ha t t he f u l l t ime l a bor o f t h ree p eop le , p r ima r i ly women a nd c h i ld ren , i sr equ i red i nd i sman t l ing o r s e t t ing -up b lack t en t e ncampmen ts . A t t h is r a te , a pp rox ima te ly 1 2 h ours o f t o ta l l abor wou ld b e r equ i red p e r d i sman t l ing a nd r ees tab l ishmen t o f ac omp le te b l ack t en t h ouseho ld , wh i le o n ly t h ree h ou rs o f t o ta l l abor wou ld b e r equ i red p e r move o f ar educed h ouseho ld o r h e rd ing e ncampmen t . F i na l ly , a pp rox ima te ly s i x h ours o f t o ta l e ncampmen t l a bo r wou ld b e r equ i red d a i ly d ur ing 9 1

m ig ra t ions when a l h ouseho ld i t e ms a re b e ing t en t e ncampmen t l abo rc os ts w i l l b e a pp l ied t o movemen ts o ft he A l Mu r rah b edou in o fS a 'ud i e ncampmen tc os ts among p as to ra l is ts who d o

moved , b u t n o t n ecessa r i ly u t i l ized . T hese b lack C o le 's d a ta o nt he s easona l h e rd a nd r e s idence A rab ia i no rde rt oe s tab l ish a pp rox ima te a nnua l n o t b u i ld b rush s t ock e nc losu res .

A nnua lA l Mu r rah E ncampmen tC os ts C o le s umma r ized t he s easona l movemen ts o ft he A l Mu r rah a sf o l lows ( 1975 :39 ) : D u r ing t he c ou rse o f ay ea r ly c yc le , t he A l Mur rah move o ve r av as t e xpanse o ft e r r i to ry . T he A l ' A zab , o ne o ft he mos tw ide rang ing g roups [ o f t h e A l Mu r rah ] , m ig ra te f r om t he i r s umme rc amps t of a l l p as tures t ha t l i e s ome 2 00 m i les t ot he s ou thwes t . W in te ra nd s p r ing p as tu res l i e a s f a r a s 6 00 m i les f r om t he f a l l p as tures . T hus , u nde r n o rma l c i rcums tances , t he A l ' Azab t r ave l a m in imum o f 1 , 200 m i les p er y ea r , f r om h ome b ase t o f a l l , w in te r , a nd s p r ing p as tu res a nd b ack a ga in t oh ome b ase a tt he i rw e l ls d u r ing t he s umme r . I nc on t ras tt ot he Gab ra a nd R end i l le o fE as t A f r ica , w ho e xp lo i t s easona l v a r ia t ions i nw a te ra nd p as tu re r esources o ccur r ing w i th in a s i ng le c i rcumsc r ibed r eg ion , t he A l Mur rah s easona l ly e xp lo i t t wo c i rcumsc r ibed r eg ions o fs easona l ly a va i lab le r e sources wh ich a re s epa ra ted b yv as t e xpanses o fl e ss d es i rab le t e r r i tory t h rough w h ich t hey a nnua l ly m ig ra te . T h is s easona l e xp lo i ta t ion o fg eog raph ica l ly d i s tan t a nd r eg iona l ly c i rcumsc r ibed w a te ra nd p as tu re r esou rces a l so c ha rac ter izes t he s easona l movemen ts o fo the r p as to ra l is ts i nt h e N ea r E as t a nd N o r th A f r ica , i n c lud ing t he B asse r i ( Ba r th 1 96 1 : 1) , Rwa la ( Johnson 1 969 :38 -46 ; Mus i l 1 928 :7 -17 ; R aswan 1 930 ) , Mu ta i r ( D ickson 1 95 1:47 -51 , 2 50 -25 1 , 5 63 ; J ohnson 1 969 :46 -53 ) , S hamma r , ' Ama ra t , H owe ta t , a nd S ha ra ra t ( Raswan 1 930 :489 -501) o f N o r th A rab ia , a nd t he c ame l n omads o f C y rena ica ( Behnke 1 980 :68 -9 ; J ohnson 1 969 :95 -6 ) , S ou the rn A lger ia ( Gau th ie r -P i l te rs a nd D agg 1 981 :140 -42 ) , a nd t he n or thwes te rn S aha ra ( Johnson 1 969 :114 1 9 ) . On ly i n r eg ions whe re p ronounced t opog raph ic r e l ie f r esu l ts i n s easona l ly a va i lab le p as tu re a nd wa te r r esou rces i nc l ose rp rox im i ty t oo ne a no the ri s al o ng -d is tance c ame l h e rd ing p a t tern b roken i n mo re n o r the r ly l a t i tudes . A s o ne e xamp le , t he r es t r ic ted a nnua l r ange o ft he Z uwe ida H owe ta t o f S ou the rn J o rdan i n vo lves t he s easona l e xp lo i ta t ion o f h i gh land a nd l ow land wa te r a nd p as tu re r esou rces s epa ra ted b y a n e l eva t iona l s h i f t o fo ve r 1 ,000 m i nl e ss t han 1 0 km ( Henry 1 984 :4 -15 ) . When C o le 's a nnua l c l ima t ic s umma ry f o rt he r eg ions e xp lo i ted b yt he A l Mu r rah ( 1 975 :39 4 0 ) i sc omb ined w i th h i s d a ta o n t he s easona l r hy thm o fA l Mu r rah h e rd ing a nd t en t m ovemen t ( 1975 :41-53 ) , t he f o l low ing s yn thes is emerges . D u r ing t he s umme r s eason ( r ough ly 9 2d ays ) , h ouseho lds r ema in s t a t iona ry a t t r ad i t iona l w e l ls . D ur ing mos t o ft he f a l l s eason ( r ough ly 9 2 d ays ) , w in te r t en ts a nd h eav ie r h ouseho ld i t e ms a re c ached n ea r t he i r we l ls , a nd h ouseho lds e s tab l ish h e rd ing c amps w i th sma l le r t en ts o r t en t s ec t ions , mov ing e ncampmen ts e ve ry t wo d ays . A tt he e nd o f F a l l , e n t i re h ouseho lds u nde r take a l o ng -d is tance m ig ra t ion o f 1 5 d ays w i th a l h ouseho ld b e long ings , mov ing n o r th t ow in te r p as tures . D u r ing t he w in te r s eason ( r ough ly 7 4 d ays ) , e n t i re h ouseho lds make s ho r t moves e very t h ree t of ou r d ays ( mean o f3 . 5 d ays ) , w i th i n d iv idua l h ouseho lds o f ten c amp ing i nc l ose p rox im i ty t oo ne a no the r . D u r ing m os to ft he s ubsequen t s p r ing s eason ( r ough ly 7 2 d ays ) , t he w in te r e ncampmen t p a t te rn i sa ppa ren t ly c on t inued , a l though i n d iv idua l h ouseho lds c amp f u r the r a par t . F ina l ly , i n l a te s p r ing , e ach h ouseho ld u nder takes a mo re l e isure ly m ig ra t ion t o t he s ou th o ve r a pp rox ima te ly 2 0 d ays , e ven tua l ly r e tu rn ing a ga in t ot he i rs umme rw e l ls . E ach y ea r , t he A l Mur rah s eem ing ly e s tab l ish a pp rox ima te ly 4 3 e ncampmen ts i n vo lv ing l a rge w in te rt en ts a nd a l h ouseho ld i t ems ( 146 t o ta l d ays +3 . 5 =4 2 moves , + 1e ncampmen t a t s ummer w e l ls ) , 4 6 e ncampmen ts i n vo lv ing sma l ler t en ts ( o r t en t s ec t ions ) a nd a r e duced q uan t i ty o fh ouseho ld i t ems ( 92 t o ta ld ays ÷ 2=4 6 moves ) , a nd f u r the rs pend a pp rox ima te ly 3 5 d ays i nl o ng -d is tance m ig ra t ion w i th a l h ouseho ld i t ems . U s ing b l ack t en t e ncampmen tc os ts o f 1 2 h ou rs p e r move o fc omp le te h ouseho ld , t h ree h ou rs p e r move o f r educed h ouseho ld , a nd s i x h ou rs p e r d ay o f m ig ra t ion , a s e s tab l ished e a r l ie r , t he a nnua l l a bo r c os ts o f e ncampmen t t ime among t he A l Mur rah wou ld amoun t t o a pp rox ima te ly 8 64 h ou rs s pen t i n at o ta l o f 1 24 moves . I ti ss uspec ted t ha t i fs u f f ic ien t d a ta we re c o l lec ted , c ompa rab le a nnua l c ame l re la ted e ncampmen t c os ts wou ld b e f ound among t he Rwa la a nd M u ta i r b edou in o f N o r th A rab ia d ue 9 2

t os im i la r i t ies b e tween t hem a nd t he A l Mu r rah i nt he i rg ene ra lc l ima t ic r eg ime , s easona l p a t te rn o fp as tu re a nd w a te r r esou rce e xp lo i ta t ion , a nd e ncampmen t movemen ts . T he n eed t o p e r iod ica l ly move h e rds t o f r esh p as turage i sa l so e xpec ted t o r esu l t i n c ompa rab le a nnua l e ncampmen t c os ts f o r o ther h e rd s pec ies . F o r e xamp le , t he s heep a nd g oa t h e rd ing B asse r i o f I r an r e loca te e ncampmen ts a pp rox ima te ly 1 20 t imes a y ea r ( Bar th 1 96 1:15 ) , q u i te c ompa rab le t ot he 1 24 a nnua l e ncampmen t moves among A l Mur rah c ame l p as to ra l is ts , a nd t he 1 29 a nnua l moves o fe ncampmen t a ssoc ia ted w i th R end i l le c ame l h e rds .

T he E fec ts o fC arn ivo re P reda t ion o nA nnua l E ncampmen t C os ts A s c a lcu la ted a bove , a nnua l A l Mur rah e ncampmen t c os ts ( 864 h ou rs ) a re a pp rox ima te ly 2 15 h ou rs l e ss t han t he mean o fa nnua l Gab ra a nd R end i l le e ncampmen t c os ts ( 1 ,079 h ou rs ) . S ince b l ack t en ts s eem ing ly r equ i re t he s ame amoun t o f l a bo rt o b reak o r r ees tab l ish a s E as t A f r ican " t en ts , " t he p r inc ip le s ource o f t hese o bse rved v a r ia t ions l i es i nt he a dd i t iona l l abo r e xpended i n c ons t ruc t ing a nd ma in ta in ing b rush s tock e nc losures i n t he mo re s ou ther ly l a t i tudes o fE as t A f r ica , whe re c a rn ivo re p reda t ion may a ccoun tf o r a nnua l l o sses o f 5t o 1 0% o f t o ta l l i ves tock ( J ones 1 984 :100 ) . B y c ompa r ison , t he f r equency o f e ncampmen t movemen ts s eems f a r l e ss s i gn i f ican t i na ccoun t ing f o rs uch v a r ia t ions i nl a bo rc os ts , s i nce A l Mu r rah c ame l h e rde rs move t he i r e ncampmen ts a pp rox ima te ly 1 24 t imes a y ea r , t he R end i l le a pp rox ima te ly 1 29 t imes a y ea r , a nd t he Gab ra a pp rox ima te ly 3 9 t imes a y ea r . H ence , a s ar esu l t o f c ons t ruc t ing a nd ma in ta in ing b rush s t ock e nc losu res i ne nv i ronmen ts w i th h i ghe r d ens i t ies o f c a rn ivo res , p as to ra l e ncampmen t c os ts s eem ing ly i n c rease b y r ough ly 2 5% . I ts eems h i gh ly p robab le t ha t b rush s t ock e nc losures wou ld h ave b een s im i la r ly emp loyed b y e a r ly p as to ra l is ts i nN o r th A f r ica a nd t he N ea rE as t . F o re xamp le , t he c a rn ivo res i n d igenous t ot he f o rme r s avannah g rass lands o f P le is tocene A rab ia i n c luded d ogs , j a cka ls , wo lves , f oxes , h yaenas , l i ons , p an the rs , c hee tahs , a nd l y nx ( Ca r ru the rs 1 910 :235 ; H a rpe r 1 945 :274 -276 , 2 92 -297 ; H a r r ison 1 968 :195 -220 , 2 73 -313 ; Moun t fo r t 1 965 :59 ; N e lson 1 973 :16 1-169 ) . I n t he e thnog raph ic p resen t , wo lves h ave b een n o ted a s a ma jo r p reda to r o f p as to ra l l i ves tock ( Dough ty 1 936 :259 , 4 52 -453 , 4 73 ; Ho le 1 978 :144 -146 ; J enn ings -B ram ley 1 905 :126 -128 , 1 906 :205 , 1 907a :31) , a nd a re p rom inen t i n t h is r o le i nA rab ic f o lk lo re ( Mus i l 1 928 :2 1-22 , 3 48 ) . P an the rs , l i ons a nd c hee tahs w ere p resumab ly h e rd p reda to rs i nt he N ea r E as tt h rough a t l e as t t he m id -n ine teen th c en tu ry A .D . ( Dough ty 1 936 :373 ; J enn ings -B ram ley 1 905 :128 129 ; M us i l 1 928 :20 ; R aswan 1 947 :53 -54 ) , a l though t he i n t roduc t ion o f mode rn weapons a nd v eh ic les h as l e d t ot he n ea r e x t inc t ion o ft he p an the r , t he p robab le e x t inc t ion o f t he c hee tah , a nd t he t o ta l e x t inc t ion o ft he l i on ( C la rk 1 981 ; Moun t fo r t 1 965 :59 ; N e lson 1 973 :16 1-162 ) . T he p re Is lam ic r ock a r t o fS a fa i t ic b edou in p opu la t ions i nN o r th A rab ia o f ten d ep ic ted l i ons a nd l i on h un ts ( L i t tmann 1 904 :107 ) , wh i le p re Is lam ic S a fa i t ic a nd T hamud ic i n sc r ip t ions f r equen t ly men t ion l i ons i n n umerous l o ca t ions i n N or th A rab ia ( W inne t t 1 957 :7 -8 ; W inne t t a nd R eed 1 970 :13 , 1 973 :77 ) . I tt he re fo re s eems r easonab le t o a ssume t ha t p reh is to r ic p as to ra l h e rds i nt he N ea r E as t w ou ld h ave b een s ub jec t t o l e ve ls o f c a rn ivo re p reda t ion g rea ter t han t hose a ppa ren t i nt he e thnog raph ic p resen t , a s t hey p resumab ly we re d ur ing t he H o locene e poch i nN or th A f r ica . Wh i le t he p o ten t ia l o f c a rn ivo re p reda t ion d u r ing d ay l igh t h ours c an b e c oun te red b y h e rders r ema in ing w i th o r n ea r t he i r h e rds t h roughou t t he t ime t ha t t hey a re p as tured a way f r om e ncampmen ts , t he p o ten t ia l o fn i gh t t ime p reda t ion c ou ld b e r educed t h rough t he c ons t ruc t ion o f b rush s tock c o r ra ls a nd e nc losures s im i la r t o t hose emp loyed i n E as t A f r ica b y modern p as to ra l is ts . T he a dd i t iona l l a bo rc os ts o fc ons t ruc t ing a nd ma in ta in ing b rush s t ock e nc losures w i l l t he re fo re b e i n c luded i ns ubsequen t c a lcu la t ions o f p o ten t ia l r e turn r a tes f o r a l l e a r ly p as to ra l s t ra teg ies .

M i lk ing T ime A s men t ioned i n Chap te r2 ,t he mos t c r i t ica l f ac to r i nt he h uman a b i l i ty t oe x t rac t m i lk f r om c ame ls , c a t t le , s heep a nd g oa ts i n vo lves t he p r io r s t imu la t ion o ft he m i lk e j ec t ion r e f lex . Once 9 3

t he m i lk h as b een " l e t -down " i n to t he s t o rage c i s te rns a s ar e su l t o ft h is r e f lex , t he s ubsequen t e x t rac t ion o ft he m i lk p resen t i nt he c i s te rn may b e a ch ieved b y man ipu la t ing t he d am 's t ea ts . Among t hese s pec ies , t he m i lk e j ec t ion r e f lex i si n t ia ted b y ar e lease o f o xy toc in t r i ggered p r ima r i ly t h rough t he mo thers r ecogn i t ion o f h e r o f fsp r ing b y s i gh t a nd sme l l . P as to ra l is ts u n ive rsa l ly e xp lo i t t h is t r a i t i no rde rt oa ch ieve m i lk e x t rac t ion , u sua l ly b y s imp ly p resen t ing t he d am w i th h e ro f fsp r ing p r io rt om i lk ing . T h is c ross -cu l tura l e xp lo i ta t ion o f a n o f fsp r ing 's s i gh t a nd sme l l i nt r igger ing t he m i lk e j ec t ion r e f lex i nt he i r mo the rs i sa l so a ppa ren t i nt he mo re e xo t ic t echn iques emp loyed b y p as to ra l is ts when a t temp t ing t o ma in ta in l a c ta t ion a nd s ecu re m i lk ing i nf ema les wh ich h ave l o s t t he i r y oung t h rough n a tu ra l d ea th o r s l augh te r . F o r e xamp le , b o th t he R end i l le ( Sa to 1 980 :31) a nd t he T ua reg ( N ico la isen 1 963 :57 ) w i l l f l ay t he h i de f r om ad ead i n fan t a nd u se t h is a s a n " e f f igy " o r " dummy " c a l f wh ich t hey t hen p resen t t ot he mo the r . S im i la r ly , when t he Rwa la c u l l t he w eak o f fsp r ing o f ag ood m i lke r , t hey o f ten p resen t h e rw i th af os te rc a l fw rapped i nt he s k in o f h e r r ea l o f fsp r ing ( Mus i l 1 928 :22 ) . A l te rna t ive ly , t he Rwa la wou ld r emove a nd k i l l an ew ly b o rn c a l f b e fo re t he mo the r c ou ld s ee o r sme l l i t ,s ubsequen t ly i n t roduc ing a s ubs t i tu te c a l f smea red w i th a l i t le b lood f r om t he c a l f wh ich was k i l led ( Musa 1 928 :87 ) . S ubs t i tu te o f fsp r ing a re a l so emp loyed b y t he R end i l le ( Sa to 1 980 :31) a nd T ua reg ( N ico la isen 1 963 :57 ) . F i na l ly , b o th t he T ua reg ( N ico la isen 1 963 :59 ) a nd t he Rwa la ( Mus i l 1 928 :87 ) f u r the r d ea l t w i th t he r e jec t ion o f f os te r o f fsp r ing o n t he b as is o f sme l l b y t y ing t he mo the r 's n os t r i ls c l osed . T echn iques s im i la rt ot hese we re a l so emp loyed w i th c a t t le i nb o th E u rope a nd A f r ica i no rde rt o s ecure m i lk ing ( Amo roso a nd J ewe l l 1 963 :130 -131 ; N ico la isen 1 963 :51) . H oweve r , i n t he f o l low ing s ec t ion , o n ly t he l a bo rc os ts o fe x t rac t ing m i lk a l ready p resen t i n ac ame l 's u dde rw i l l b e c ons ide red , s i nce t he u se o f t hese e xo t ic a nd mo re l a bo r ious d a i ry ing t echn iques i s s eem ing ly r es t r ic ted t or e la t ive ly f ew a n ima ls . F requency o fD a i ly M i lk ing . S ince t he l abo rc os ts o fm i lk e x t rac t ion f r om p as to ra l h e rds w i l l b e c a lcu la ted o n t he b as is o ft ime s pen t p e r k i log ram o f m i lk y i e ld , n e i the rt he n umbe ro fd a i ly m i lk ings n o rt he n umbe ro fm i lke rs i n vo lved w i l l a f fec tt he r e su l ts . E ven s o , b o th o ft hese f ac to rs a re w o r thy o fb r ie f d i scuss ion . I ng ene ra l , i ti ss ta ted t ha t c ame ls a re u sua l ly m i lked t w ice d a i ly ( Rao e ta t . 1 970 :73 ; Y as in a nd Wah id 1 957 :293 ) . I ndeed , t he R end i l le ( Sa to 1 980 :3 1) , Gab ra ( To r ry 1 973 :73 ) , a nd A y r T ua reg ( N ico la isen 1 963 :59 ) a re s a id t o n o rma l ly m i lk t he i r c ame ls t w ice d a i ly , o nce i nt he mo rn ing a nd o nce i nt he e ven ing . I nd i s t inc t c on t ras t , t he Rwa la a re s a id t o o n ly m i lk t he i r c ame ls o nce i nt he mo rn ing ( Mus i l 1 928 :88 , 3 37 ) , wh i le t he A l Mu r rah a re a l so s a id t oo n ly m i lk t he i r c ame ls o nce , b u t i n t he e ven ing ( Co le 1 975 :38 ) . F ur the r , D ough ty ( 1936 :304 , 3 69 ) s t a ted t ha t t he b edou in o f n or thwes t A rab ia o n ly m i lked c ame ls w i th i n fan ts i nt he e ven ing , w h i le c ame ls wh ich h ad l o s t t he i r y oung were m i lked t w ice d a i ly , mo rn ing a nd e ven ing . Mo re f r equen t m i lk ings a re a l so r eco rded . T he G ur rah , S oma l i ( To r ry 1 973 :74 ) , a nd K e l T amasheq ( S . Sm i th 1 980 :469 ) a re s a id t o m i lk t he i rc ame ls t h ree t imes d a i ly , wh i le t he T u rkana s ome t imes m i lk t he i r c ame ls f ou rt o f i ve t imes d a i ly ( To r ry 1 973 :74 ) . P resumab ly , s ome o f t he v a r ia t ion a ppa ren t i nt he f r equency o f r epor ted d a i ly m i lk ings r e la tes t os easona l v a r ia t ions i nm i lk p roduc t ion , a s s ugges ted b y T o r ry ( 1973 :74 ) . Af u r the rf ac tor was n o ted b y S a to ( 1980 :31) , w ho o bserved t ha t t he R end i l le wou ld m i lk t he i r c ame ls u p t o f our o r f i ve t imes d a i ly when t he d am was p roduc ing c o los t rum a f te r p ar tur i t ion . P resumab ly , t he f r equency o f d a i ly m i lk ings among a l l p as to ra l is ts w ou ld v a ry a cco rd ing t o s uch f ac to rs . S ign i f ican t ly , b o th K noess ( 1977 :41) a nd Y as in a nd Wah id ( 1957 :293 ) n o ted t ha t d a i ly m i lk p roduc t iv i ty i nc ame ls may b e i n c reased 1 0 -12 % b y mo re f r equen t m i lk ings , a nd s im i la r r esul ts i np roduc t iv i ty h ave a l so b een n o ted w i th t he f r equency o f m i lk ing among d a i ry c a t t le ( Sh inde 1 978 :352 -358 ) . U n for tuna te ly , t he a va i lab le q uan t i ta t ive d a ta i si n suf ic ien tf o r ad e ta i led mode l l ing o fs uch r e la t ionsh ips . N umbe ro fM i lker s . F ur the rv a r ia t ions s eem ing ly e x is tf o rt he n umbe ro fm i lke rs p e ra n ima l . Among t he R end i l le ( Jones 1 984 :126 ) , Gab ra ( To r ry 1 973 :73 ) , a nd A y r T ua reg ( N ico la isen 1 963 :59 ) , t wo ma les a re s a id t o s imu l taneous ly m i lk e ach c ame l . H owever , Mus i l ( 1928 :88 ) s eem ing ly i mp l ies t ha t o n ly o ne p e rson n orma l ly m i lked e ach Rwa la c ame l . R ega rd less , i ti s s uspec ted t ha t v a r ia t ions i nt he n umbe r o f m i lke rs p e r a n ima l o r h e rd wou ld o n ly a fec t t he l e ng th o f t ime i tt akes t o m i lk t ha t h e rd b u t n o t t o ta l m i lk ing l a bo r . W i th t wo p eop le m i lk ing s imu l taneous ly , t he t ask wou ld p resumab ly t ake h a l f a s m uch t ime wh i le t he t o ta l l a bor c os t w ou ld r ema in c ons tan t . 9 4

T he T ime C os ts o fM i lk ing . M i lk e x t rac t ion f r om s pec ies w i th c apac ious u dde rc i s te rns i s a ch ieved b y h o ld ing at ea tw i th t h ree o rf o ur f i nge rs w h i le p ress ing t he o ppos i te s i de o ft he t ea t w i th t he t humb ,w i thou t p u l l ing . T he r e su l t ing p ressu re makes t he m i lk s pu r t i n as t rong j e t i n to a r e cep tac le . A ssum ing t ha tp as to ra l h e rds a re m i lked a tt he e ncampmen t i nt he p resence o ft he i r y oung , t he l a bo rc os ts o fd a i ry ing w ou ld p r ima r i ly c ons is t o ft he t ime i tt akes t od o t he a c tua l m i lk ing . U n fo r tuna te ly , n oo bse rva t iona l d a ta h ave b een f o und o nt he t i me c os ts o fm i lk ing p as tora l h e rd a n ima ls . H oweve r , a cco rd ing t ot he G u iness B ook o f Wo r ld R eco rds ( 1985 :252 ) , t he h and m i lk ing r eco rd f o rd a iry c a t t le w as s e t i n1 937 i nO k lahoma ,w hen 1 20 g a l lons w e re m i lked i n1 2 h ours . A t 1 0 g a l lons p e r h ou r , a nd w i th o ne g a l lon o f m i lk b e ing e qua l t o 3 .904 k g ( USDA H andbook N o . 8 1 , 1 976 ) , a pp rox ima te ly 1 . 54 m inu tes o f e x t rac t ion t ime w as r equ i red p e r k i log ram o fm i lk ( 60 m inu tes 1 -3 9 .04 k g =1 . 54 m in /kg ) . G iven t he s im i la rp hys io logy o fc ame l a nd b ov id l a c ta t ion , i ti ss uspec ted t ha t t he e f f ic iency o f m i lk e x t rac t ion v i a t he t ea t s ph inc te r s hou ld a l so b e c ompa rab le . A ssum ing t ha t ap as to ra l is tw ou ld b e a pp rox ima te ly h a l f a s e f ic ien t a s a n i n d iv idua l a t temp t ing t os e t aw o r ld m i lk ing r eco rd , a pp rox ima te ly t h ree m inu tes o fm i lk ing t i me w ou ld b e r equ i red p e rk go fm i lk e x t rac ted r ega rd less o ft he s pec ies i n vo lved . I na dd i t ion t ot he d i rec t t i me c os ts o fm i lk ing , a lowance w i l l a l so b e made f o rc on t ro l l ing r e ca lc i t ran t f ema les d ur ing t h e m i lk ing p rocess . Among t he A y rT ua reg , f ema le c ame ls w h ich a re r e s t less a nd t here fo re d i f icu l tt o m i lk a re s imp ly h obb led ( N ico la isen 1 963 :6 1) , ap rac t ice o f ten men t ioned i nc onnec t ion w i th m i lk ing c a t t le i nE u rope a nd A f r ica , a nd o ne t ha t o f ten a ppea rs i n a nc ien t E gyp t ian d ep ic t ions o f d a i ry ing ( Amo roso a nd J ewe l l 1 963 :133 -135 ) . P resumab ly , ah obb led a n ima lc ou ld b e m i lked b yo ne p e rson , a l though h av ing t oh obb le t hem w ou ld i n c rease t o ta l m i lk ing t i me . I nt he a bsence o fo bse rva t iona l d a ta , i tw i l l b e a ssumed t ha t o ne a dd i t iona l p e rson w ou ld b e r equ i red d u r ing t he m i lk ing p rocess i no rde r t o c on t ro l t he f ema le a nd h e ra t tendan t o fsp r ing . A s s uch , t he l a bo rc os ts o fm i lk ing a s p resen ted a bove w i l l b e d oub led i ns ubsequen t c a lcu la t ions ( = 6m inu tes /kg m i lk ) . T he E nerge t ics o f D a i ry ing . D ah l a nd H jo r t ( 1976 :154 ) c a lcu la ted t he e ne rgy v a lue o f c ow 's m i lk a t8 08 k caVkg , a l though t he s t anda rd a na lys is wh ich t hey u sed ( 1976 :153 ) w ou ld s ugges t a n e ne rgy c on ten t o f 8 39 k ca l /kg . B o th v a lues a re s l igh t ly h i ghe r t han t he 6 69 .5 k ca l /kg e xpec ted f o r c ame l 's m i lk ( Append ix B ) . T he c a lor ic c on ten t o f g oa t 's m i lk i s a pp rox ima te ly 6 68 k ca l /kg , w h i le t ha t o f s heep i sa pp rox ima te ly 1 , 096 k ca l /kg ( Redd ing 1 98 1: 177 , T ab le V I -2 ) . A t a n e x t rac t ion r a te o f 1 0 k g /h ro ft o ta l l a bo r , 1 0 ,960 k ca l /h r c ou ld b e o b ta ined f r om m i lk ing c ap t ive s heep , 8 ,080 k ca l /h rf o r m i lk ing c ap t ive c a t t le , 6 ,695 k caVh rf o r m i lk ing c ap t ive c ame ls , a nd 6 , 680 k ca l /h rf o rm i lk ing c ap t ive g oa ts . T hese p os tencoun te r r e tu rn r a tes a re q u i te h i gh r e la t ive t o t hose o f o ther f ood i t ems e xp lo i ted b y mode rn f o rage rs i nS ou th Ame r ica ( Hawkes e ta l . 1 982 :389 , T ab le 3 ; H i l l e ta l . 1 986 :Tab le 2 )a nd A us t ra l ia ( O 'Conne l l a nd H awkes 1 984 :5 13 , T ab le 1 ) , a nd v a r ious r e sou rces p reh is to r ica l ly a nd e t hnog raph ica l ly e xp lo i ted i nt he G rea t B as in ( S imms 1 984 :93 -94 , T ab le 5 , 1 987 ) . S ince f o rage rs w e re a ppa ren t ly e xpend ing l a bo ri nt he c ap tu re a nd c on t ro lo fw i ld h e rds t h rough t he u se o fg ame d r ive l i nes a nd c o r ra ls p r io rt ot he e vo lu t ion o fp as tora l s t ra teg ies ( a s n o ted i n C hap te r 2 ) , as t rong e ne rge t ic i n cen t ive wou ld h ave p resumab ly e x is ted f o r a dd i t iona l ly e x t rac t ing m i lk f r om t he l e ss reca lc i t ran t f ema les among h e rd s pec ies p ossess ing c apac ious u dde rc i s te rns . P o ten t ia l ly , i n i t ia ld a iry ing a c t iv i t ies may h ave p receded t he e vo lu t ion o fp as to ra l s ubs is tence s t ra teg ies .

B u tche r ing T ime T o S laugh ter a C ame l . T he mos t e xp l ic i t a ccoun t o f c ame l s l augh te r a nd b u tche r ing p rac t ices b yp as to ra l is ts i st ha tg i ven b y M us i la sb ased u pon h i so bse rva t ions among t he Rwa la b edou in . I ft hey w ish t os l augh te r ac ame l , t hey make i tk nee ld own ,j enawwehunaha , t i e a l i t s l e gs , p ress i t s h ead t ot he g round , a nd t he b u tche rs hou ts : " Lay i t s h ead o n o ne s i de ! r udd r a sha . H et hen t h rus ts as ha rp d agge ri n to i t st h roa t , manha r , a nd c u ts t h rough t ot h e r i gh t a nd l e f t . T he c ame lt r emb les a nd makes ag u rg l ing n o ise , t i r ta ied , a nd t he b l ood f l ows e i the ri n to av esse l o ru pon t he g round . T he b edou ins c u t o f t he t a i l o ft he s l augh te red c ame l , c u t t h rough t he s k in o n i t s 9 5

b ack f r om t he r oo ts o ft he t a i l a c ross t he h ump a s f a r a s t he n ape , r a reb , a nd t hen f l ay i t ,j as lehunaha . When t he s k in h as b een r emoved , t hey c u t o f f t he f r on t l e gs , t hen t he n eck a nd t he h i nd l e gs , a nd o pen t he b e l ly . T he o n ly t h ings wh ich a re t h rown a way a re t he c on ten ts o f t he s t omach a nd t he i n tes t ines , f a r t , a l t he r es t b e ing c l eansed a nd made u se o f[ Mus i l 1 928 :96 ] . T he T ua reg me thod o fc ame l s l augh te ri sq u i te s im i la r : C o rds a re a t tached v er yf i rm ly a round t he l e gs o ft h e a n ima lw h ich i sk ep tt ot he g round b y s eve ra l men who s i t o n i t s b ack a nd d raw i t s h ead b ackwa rds . T he a r tery i st he reupon o pened a tt he l owe rp a r to ft he n eck c l ose t ot he c hes t , a nd t he k n i fe i st h rus t i n to t he a n ima l a tt h is p l ace , b u t n o t s o d eep ly a st ot ouch t he h ea r t o rt he e n t ra i ls [ N ico la isen 1 963 :65 ] . H owever , s i nce t he T ua reg make l i t le u se o f ac ame l 's h i de , t he i r b u tche r ing p rac t ices a re s omewha t d i fe ren tf r om t hose r epor ted f o rt he Rwa la . Wh i le o the r a n ima ls a re s k inned i ns uch away t ha t t he h i de c an b e r emoved e n t i re a s p oss ib le , t he c a rcass o f ac ame l i sc u t u p b e fo re s k inn ing . T he f o ur l e gs w i th t he i rt h ighs a re c u t l o ose c l ose t ot he c a rcass , wh ich i st he reupon c u t i n to s eve ra l p ar ts b y means o f al a rge c hopp ing -kn i fe o r a n a xe . T he h i de i s t hus r emoved f r om t he c a rcass a f te rp a r t i t ion ing ; b u t i ti sd i f f icu l t t or emove t he h i de a s i ts t icks f i rm ly t ot he mea t , a nd o n ly p a r t o fi ti st aken o f f f o r l a te r u se [ N ico la isen 1 963 :65 ] . U n for tuna te ly , t he t ime a c tua l ly s pen t i n b u tche r ing c ame ls h as n o t b een r epor ted i nt he a va i lab le l i t e ra tu re . T h is i sa l so t he c ase w i th o the r p as to ra l s pec ies , a l though af ew s t a temen ts o n b u tche r ing t imes f o ro the ra n ima ls d o e x is t . A va i lab le B u tche r ing T imes . A l l o ft he a va i lab le d a ta o n b u tche r ing t imes a re b ased u pon p opu la t ions c u r ren t ly u s ing me ta l t oo ls , i n c lud ing a xes a nd k n ives . U n fo r tuna te ly , s cho la rs r ep l ica t ing a nc ien t b u tche r ing p rac t ices w i th s t one t oo ls ( e .g . , F r ison 1 978 ) h ave c ons is ten t ly f a i led t o s t a te a ny t imed r esu l ts . H ence , v a r ia t ions i n b u tche r ing t imes a s af unc t ion o f t echno log ica l v a r ia t ions c anno t b e a dd ressed a tp resen t . T he b es t a va i lab le d a ta o n b u tche r ing c os ts a re t hose p resen ted b y H awkes e t a l . ( 1982 :389 ; a s emended b y H i l l e ta l . 1 986 ) , w h ich a re b ased u pon o bserva t ions among A che f o rage rs i nE as te rn P a raguay . T he A che a re a b le t o b u tche r a3 0 k g wh i te l ipped p ecca ry i n a pp rox ima te ly 0 .25 h ou rs . H oweve r , t he A che s imp ly e v isce ra te a nd q ua r te r ak i l l i np repa ra t ion f o rc ook ing , w i th s ubsequen t c u ts o f mea t b e ing made j u s t p r io rt oc onsump t ion . F ur the r , h i des o r s k in a re n o t r emoved d ur ing t he b u tche r ing p rocess ( K im H i l l , p e rsona l c ommun ica t ion 1 985 ) . S im i la r ly , a n i n d iv idua lA lyawa ra f o rage ri nA us t ra l ia i sa b le t oe v isce ra te a nd q ua r te r a2 5k g k anga roo i na pp rox ima te ly 0 .25 h ou rs , a ga in w i thou t s k inn ing . I no ne o bserved i n s tance , ac ow w e igh ing r ough ly 3 00 -350 k g was e v isce ra ted a nd q ua r te red w i thou t s k inn ing b yt wo A lyawa ra ma les i na pp rox ima te ly 0 .5 h ours , wh ich wou ld amoun t t o a pp rox ima te ly o ne h ou r o f t o ta l b u tche r ing t ime , e xc lud ing i t s s ubsequen t r educ t ion i n to c u l ina ry c u ts ( James F . O 'Conne l l , p e rsona l c ommun ica t ion 1 985 ) . C ompa rab le b u tche r ing t imes a re i mp l ied b y B in fo rd 's d a ta o n t he N unam iu t ( 1978 ) . AD a I I S heep , w h ich w e igh a bou t 5 4 k g , was s ho t b y o ne o f B in fo rd 's i n forman ts ( 1978 :142 -144 ) . A t t he k i l l s i te , t he a bdom ina l c on ten ts a nd a l me tapod ia ls a nd p ha langes we re r emoved . U pon r e turn ing t o c amp , t he s ubsequen t b u tche r ing o f t he c a rcass , f r om s k inn ing t h rough i t s r educ t ion t o s t o rab le c u ts o f mea t , r equ i red o n ly 1 6 .8 m inu tes . A l low ing a n a dd i t iona l 1 3 m inu tes f o rt he i n i t ia l b u tche r ing a tt he k i l l s i te , a pp rox ima te ly 0 .5 h ou rs o ft o ta l b u tche r ing t i me we re r equ i red . F u r the r , B in fo rd ( 1978 :52 ) n o ted t ha t b e tween l a te a f te rnoon a nd n i gh t fa l l , a s i ng le N unam iu t h un te r was a b le t os k in , e v isce ra te , a nd h a lve s even b u l l c a r ibou , wh ich w e igh a bou t 1 00 k g e ach ( 1978 :136 ) . P resumab ly , s omewha t l e ss t han o ne h ou r o f b u tche r ing t ime was r equ i red p er c a r ibou , a l though t h is w ou ld n o t i n c lude t he t ime r equ i red t o r educe t he h a lved c a rcasses i n to s pec i f ic c u ts .

9 6

S im i la rb u tche r ing c os ts w e re r epor ted f o rt he B lack foo t o ft he N or the rn G rea t P la ins , who w ou ld s k in , e v isce ra te , a nd q uar te rb ufa lo a f te r ak i l l . W iss le r 's a ccoun t o ft he i rf i e ld b u tche r ing w as a sf o l lows ( 19 10 :42 ) : T he o n ly t oo l u sed i nb u tche r ing was ah eavy me ta l k n i fe . T he b one o ft he f o re l e g was o f ten u sed a s ac l ub t o b reak sma l le r b ones a nd j o in ts . T he t ime r equ i red f o r c u t t ing u p ac a rcass was s hor t , a s men n ow l i v ing c l a im t o h ave b een a b le t ob u tche rf r om f i ve t ot we lve b ufa lo i n ad ay . W iss le r 's a ccoun t o ft he t i me r equ i red t of i e ld p rocess b u f fa lo c l ose ly p a ra l le ls t he n i ne teen th c en tury o bse rva t ions o f F a the r Geo rges A n to ine B e !cour t . I n al e t ter d a ted S ep tembe r 2 5 , 1 845 , B e lcour t p rov ided t he f o l low ing a ccoun t o fB lack foo t b u f fa lo b u tche r ing ( 1944 :16 ) : T o d ress a nd b u tche r o ne o ft hese a n ima ls i sq u i te a n a rduous t ask , b u t o ur f o lk g o t o i tw i th aw i l l a nd s k i l l t r u ly a s ton ish ing . S ome o ft hem h ave b een k nown t o k i l l a nd d ress t en b u f fa lo , w i thou t a ny a ss is tance , i n l e ss t han t en h ours . I tt he re fo re a ppea rs t ha t ab u f fa lo , w i th a n o rma l a du l t w e igh t o fa round 1 , 000 k g ( Wa l lach a nd B oeve r 1 983 :3 19 ) , c ou ld b e s k inned , e v isce ra ted , a nd q ua r te red i na bou t o ne h ou r . F rom t he a va i lab le d a ta , i twou ld s eem t ha t t he l i vewe igh t o fl a rge r a n ima ls a c tua l ly h as a m in ima l e fec to n t he t o ta l b u tche r ing t ime r equ i red t os k in , e v isce ra te , a nd q ua r te rt he c a rcass . H oweve r , t he s ubsequen t c os ts o f r educ ing a h a lved o rq ua r te red c a rcass i n to s pec i f ic c u ts o f mea t w ou ld p resumab ly v a ry a cco rd ing t o b o th c a rcass s i ze a nd t he c u l ina ry c u ts d es i red . F ur the r , t he p rocess ing o fs econda ry p roduc ts , s uch a s s i new , w ou ld a l so i n c rease b u tche r ing c os ts . P resumab ly , s uch f a c to rs w e re r espons ib le f o r Y e l len 's o bserva t ion ( 1977 :280 ) t ha t t he ! Kung s pen t a pp rox ima te ly t wo h ours i n s k inn ing a nd b u tcher ing a 1 25 k g w i ldebees te . H oweve r , g i ven t he c ompa ra t ive t ime d a ta p resen ted a bove , L ee 's o bse rva t ion ( 1979 :278 ) t ha t ma le ! Kung b u tche r ing a nd c ook ing t ime wou ld v a r yf r om o ne h our f o r ag ame b i rd t of ou rh ou rs f o r awa r thog a nd 1 5 h ou rs f o r al a rge a n te lope , i su n fa thomab le . I nt he a bsence o ff u r the rq uan t i ta t ive d a ta , i tw i l l b e a ssumed t ha t a max imum t o ta l o f2 .5 h ours o f b u tche r ing t ime w ou ld b e r equ i red f o r e ach s l augh te red c ame l o r c ow . T h is a lows amp le t i me f o rt he r educ t ion o f al a rge c a rcass i n to f a i r ly sma l l c u l ina ry c u ts . S ince i ti sp robab le t ha t t he b u tche r ing c os ts p e ra n ima l f o r s heep a nd g oa ts wou ld b e l e ss t han t hose a ssoc ia ted w i th c ame ls o rc a t t le d ue t os ma l le rb ody s i ze a nd c onsequen t ly t he n umbe ro fc u ts r equ i red f o r c u l ina ry r educ t ion , o n ly 0 .5 h ou rs o fb u tche r ing t ime p e ra n ima lw i l l b e a ssumed . Wh i le t hese a re o bv ious ly r ough a pp rox ima t ions , i n accuracy h ere i sn o t e xpec ted t o s e r ious ly a fec t c a lcu la t ions o ft o ta l a nnua l p as to ra l l a bo rc os ts . A sw i l l b e s hown , e ven i n mea t o n ly p as tora l s t ra teg ies , b u tche r ing c os ts wou ld c ons t i tu te l e ss t han 1% o ft o ta l a nnua l l abo r c os ts . F ur the r , t he p rocess ing t ime f o r g ame a n ima ls h as b een s hown t o b e an eg l ig ib le c omponen t o ft o ta l h un t ing c os ts when c ompa red t ot he t ime e xpended b y f o rage rs i np u rsu i t o fg ame i t e ms ( H i l l e ta l . 1 986 ) .

S ummary o fT o ta lA nnua lP as to ra lL abo rC os ts Mea t -On ly S t ra teg ies . I ft he g oa l o fe a r ly p as to ra l s t ra teg ies was s imp ly t he p roduc t ion o f c a lo r ies f r om mea t , o rgan , a nd f a t y i e lds , a nnua l l a bo r c os ts w ou ld s o le ly c ons is t o f h e rd ing , e ncampmen t , a nd b u tche r ing t a sks . T ab le 1 0 s umma r izes t he e xpec ted a nnua l l abo rc os ts f o r mea t -on ly c ame l s t ra teg ies i n vo lv ing a h e rd c on ta in ing e xac t ly 1 00 h ead u nder c ond i t ions o f n o rma l a nd max imum o f f take ( s ee A ppend ix A ) . T ab le 1 s im i la r ly s umma r izes t he e xpec ted a nnua l l a bo rc os ts f o r mea t -on ly c a t t le , s heep a nd g oa t s t ra teg ies i n vo lv ing h e rds c on ta in ing e xac t ly 1 00 h ead . F rom ac a t t le h e rd o ft h is s i ze , a n o f f take o f a pp rox ima te ly 1 2 a n ima ls w ou ld b e p oss ib le d u r ing ay ea r i f5 0% o ft he a n ima ls d y ing o f n a tura l c auses we re c onsumed i na dd i t ion t ot he c u l l ing o fe xcess a n ima ls i ns uch a f ash ion a st oa low t he h e rd t oc on t inue t og row a t3 . 5% a nnua l ly . F rom ah e rd o f1 00 s heep , a n o f f take o f a pp rox ima te ly 3 0 a n ima ls wou ld b e e xpec ted i f7 5% o f a n ima ls d y ing o f n a tu ra l c auses w e re c onsumed i na dd i t ion t ot he c u l l ing o fe xcess a n ima ls i ns uch af a sh ion a st oa low 9 7

T ab le 1 0 S umma ry o fA nnua l P as to ra l L abo rf o r Mea t -On ly C ame l S t ra teg ies I n vo lv ing H e rds C on ta in ing 1 00 H ead

ACT IV ITY

NORMAL OFFTAKE L abo r %o f T o ta l H rs ( h rs )

MAX IMUM OFFTAKE L abo r % o f ( h r s ) T o ta l H rs

H erd ing E ncampmen t B u tche r ing

5 ,056 1 ,079 2 0

8 2 .2 1 7 .5 0 .3

5 ,056 1 ,079 3 3

8 2 .0 1 7 .5 0 .5

T o ta l

6 , 155

1 00 .0

6 , 168

1 00 .0

T ab le 1 S umma ry o fA nnua l P as to ra l L abo rf o r Mea t -On ly C a t t le , S heep a nd G oa t S t ra teg ies I n vo lv ing H e rds C on ta in ing 1 00 H ead

ACT IV ITY

CATTLE L abo r % o f T o ta l H rs ( h rs )

SHEEP %o f L abo r ( h rs ) T o ta l H rs

GOATS L abo r %o f ( h rs ) T o ta l H rs

H e rd ing E ncampmen t B u tcher ing

5 ,056 1 ,079 3 0

8 2 .0 1 7 .5 0 .5

3 ,656 1 ,079 1 5

7 7 .0 2 2 .7 0 .3

3 ,656 1 ,079 1 7

7 7 .0 2 2 .7 0 .3

T o ta l

6 , 165

1 00 .0

4 ,750

1 00 .0

4 ,752

1 00 .0

t he f l ock t oc on t inue t og row a ta n a nnua l r a te o f6 .3% . F rom ah e rd o f1 00 g oa ts , a pp rox ima te ly 3 4 a n ima ls w ou ld b e a va i lab le f o rs l augh te r i f7 5% o fa l a n ima ls d y ing o fn a tura l c auses we re u t i l ized i na dd i t ion t o t he c u l l ing o f e xcess a n ima ls i ns uch a f ash ion a s t o a low t he h erd t o c on t inue t og row a ta n a nnua l r a te o f1 .5% . I n b o th t ab les , e ncampmen t c os ts i n c lude t he c ons t ruc t ion a nd ma in tenance o f b rush s t ock e nc losures . I ft hese we re n o t r equ i red , t o ta l a nnua l mea t -on ly l a bo r c os ts wou ld d rop a pp rox ima te ly 3 .5 % f o rc ame l a nd c a t t le s t ra teg ies , a nd 4 .5% f o rs heep a nd g oa t s t ra teg ies . Mea tP lus M i lk S t ra teg ies . I ft he g oa lo fe a r ly p as to ra l s t ra teg ies i n c luded t he p roduc t ion o f c a lo r ies f r om m i lk , t he c os ts o fs uch s t ra teg ies w ou ld i n c lude m i lk ing t ime i na dd i t ion t oh e rd ing , e ncampmen t , a nd b u tche r ing l a bo r . U s ing t he p roduc t iv i ty d a ta g ene ra ted i n A ppend ix B , T ab le 1 2 s umma r izes t he e xpec ted a nnua l l a bo r c os ts f o r mea t p l us m i lk c ame l s t ra teg ies i n vo lv ing n o rma l o f take a nd v a ry ing l e ve ls o f m i lk p roduc t ion f o r ah e rd c on ta in ing 1 00 h ead . S uch a h e rd i se xpec ted t o p roduce a pp rox ima te ly 1 0 ,352 k g o f m i lk u nde r c ond i t ions o f l ow p roduc t iv i ty , 1 6 ,958 k g u nde r n o rma l p roduc t iv i ty , a nd 2 2 ,6 10 k g u nde r h i gh p roduc t iv i ty . T ab le 1 3 s im i la r ly s umma r izes e xpec ted a nnua l l a bo rc os ts f o r mea t p l us m i lk c a t t le , s heep a nd g oa t s t ra teg ies i n vo lv ing h e rds c on ta in ing e xac t ly 1 00 h ead . U nde r a ve rage c ond i t ions , a p as to ra l c a t t le h e rd o ft h is s i ze i se xpec ted t o p roduce a pp rox ima te ly 6 ,803 k g o f m i lk , af l ock c on ta in ing 1 00 s heep i se xpec ted t o p roduce 1 , 624 k g o f m i lk , a nd a h erd c on ta in ing 1 00 g oa ts i se xpec ted t op roduce 2 ,283 k g o fm i lk .

9 8

T ab le 1 2 S umma ry o fA nnua l P as to ra l L abo rf o r aMea t P l us M i lk C ame lS t ra tegy I n vo lv ing aH e rd C on ta in ing 1 00 H ead a tV a r ious L eve ls o fM i lk P roduc t ion W i th N o rma l O f f take

ACT IV ITY

L OW M ILK Y IELD L abo r % o f ( h rs ) T o ta l H r s

ILK Y IELD NORMAL M %o f L abo r T o ta l H rs ( h rs )

H IGH M ILK Y IELD % o f L abo r ( h rs ) T o ta l H rs

H e rd ing E ncampmen t M i lk ing B u tcher ing

5 ,056 1 ,079 1 ,035 2 0

7 0 .3 1 5 .0 1 4 .4 0 .3

5 ,056 1 ,079 1 ,696 2 0

6 4 .4 1 3 .7 2 1.6 0 .3

5 ,056 1 ,079 2 ,261 2 0

6 0 .1 1 2 .8 2 6 .9 0 .2

T o ta l

7 , 190

1 00 .0

7 ,851

1 00 .0

8 ,416

1 00 .0

T ab le 1 3 S umma ry o fA nnua l P as to ra l L abo rf o r Mea t P lus M i lk C a t t le , S heep a nd G oa tS t ra teg ies I n vo lv ing H e rds C on ta in ing 1 00 H ead

ACT IV ITY

CATTLE L abo r %o f ( h rs ) T o ta l H rs

SHEEP L abo r ( h rs )

GOATS

% o f T o ta l H rs

L abo r ( h rs )

% o f T o ta l F i rs

H e rd ing E ncampmen t M i lk ing B u tcher ing

5 ,056 1 ,079 6 80 3 0

7 3 .9 1 5 .8 9 .9 0 .4

3 ,656 1 ,079 1 62 1 5

7 4 .4 2 2 .0 3 .3 0 .3

3 ,656 1 ,079 2 28 1 7

7 3 .4 2 1.7 4 .6 0 .3

T o ta l

6 ,845

1 00 .0

4 ,912

1 00 .0

4 ,980

1 00 .0

A ga in , t he e ncampmen t c os ts emp loyed i nt hese t ab les i n c lude t he c ons t ruc t ion a nd ma in tenance o fb rush s tock e nc losu res . I ft hese w e re u nnecessa ry , a nnua l l a bo r c os ts w ou ld d rop a pp rox ima te ly 3% f o rc ame l a nd c a t t le s t ra teg ies , a nd 4 .4% f o rs heep a nd g oa t s t ra teg ies .

R e turn R a tes f o r P as to ra l S t ra teg ies I n vo lv ing C ame ls , C a t t le , S heep a nd Goa ts R e tu rn R a tes f o rP as to ra lC ame lH e rds U s ing t he c os t a nd b ene f i t d a ta p resen ted a bove , r e turn r a tes f o r p as to ra l c ame l h e rds u nde r v a r ious c ond i t ions o f p roduc t iv i ty a nd h e rd s i ze h ave b een c a lcu la ted . T he r esu l ts a re p resen ted i nF igu re 7 .S eve ra ld rama t ic v a r ia t ions i nr e tu rn r a tes a re o bserved b e tween d i s t inc t c ond i t ions o fp roduc t iv i ty . F o r e xamp le , max imum o f f take , w h ich a lows f o r n o h e rd g row th a nd i n c ludes a 1 00 % u se o f a l n a tura l d ea ths , r esu l ts i n a5 7% i n c rease o ve r t he r e tu rn r a tes e xpec ted u nde r n o rma l o f f take , w h ich a l lows f o r c on t inued h e rd g row th a nd i n c ludes a 5 0% u se o fa l n a tu ra l d ea ths . A ssum ing c ond i t ions o fn o rma l o f f take , r e turn r a tes f o r amea t p l us m i lk s t ra tegy u nde rc ond i t ions o fh i gh m i lk y i e ld i n ah e rd c on ta in ing 1 50 h ead a re 4 7% g rea te rt han t he r e tu rn r a tes e xpec ted u nde r c ond i t ions o f l ow m i lk y i e ld f o r ah e rd o f e qua l s i ze . Mos t s i gn i f ican t ly , t he r e tu rn r a tes f o r amea t p l us m i lk s t ra tegy u nde rc ond i t ions o fl ow m i lk y i e ld i na h e rd c on ta in ing 1 50 h ead a re 1 33% g rea te r t han t hose e xpec ted f o r amea t o n ly s t ra tegy i na 9 9

3000 -

2500 -

1 0 3

t c

I —

2000 -

1500 — t it." ° e t >

1000 —

. . -

nortne1 o llte M b

c 500 —

_

-

0

10

30

50

70

90

1 10

130

1 1 50

NU MBER OF CA MELS I N HERD F i gure 7 . R anges o f p o ten t ia l r e turn r a tes f o r c ame l h e rds u nder v a r ious p roduc t ive p oss ib i l i t ies .

h e rd o fe qua l s i ze u nde rc ond i t ions o fn o rma lo f f take , w h i le u nde rc ond i t ions o fh i gh m i lk y i e ld , r e turn r a tes w ou ld b e 2 42% g rea te rf o r am ea t p l us m i lk s t ra tegy . I ns ubsequen t c ompa ra t ive c a lcu la t ions , c ame lc os ts a nd b ene f i ts w i l lb eb ased u pon n o rma lo f t take a nd n o rma l m i lk y i e ld .

C ompa ra t ive R e turn R a tes F o r Mea t -On ly P as tora lS t ra teg ies F i gu re 8 d i ag rams t he r e la t ionsh ips e xpec ted b e tween a nnua l c a lor ic p roduc t iv i ty ( s o l id l i nes , r ead o ft he v e r t ica l s ca le o nt he l e f t ) a nd a nnua l l a bo rr e qu i remen ts ( b roken l i nes , r e ad o f t he v e r t ica l s ca le o nt he r i gh t ) f o r mea t -on ly c ame l , c a t t le , s heep a nd g oa t s t ra teg ies w i th h e rds o fv a r ious s i zes . T he d i agona l ly s haded a reas r e f lec t t he r a nge o fv a r ia t ion e xpec ted b e tween t he c a lo r ic p roduc t iv i ty o fe t hnog raph ic h e rds a nd p o ten t ia l p roduc t iv i ty when s i ze r educ t ions u pt o2 0% f o rc a t t le a nd 3 0% f o rs heep a nd g oa ts a re a ccoun ted f o r . L abo r c os ts a s c a lcu la ted h e re e xh ib i t l i t le s l ope w i th i n c reas ing h e rd s i ze , s i nce o n ly s l augh te rl a bo ri se xpec ted t oi n c rease . F o re xamp le ,t h e a nnua l l a bo rc os ts o f mea t -on ly c ame l s t ra teg ies i n c rease b y o n ly 2 6 h ou rs f r om h e rds c on ta in ing 1 0 a n ima ls ( 6 ,137 h r s ) t o h e rds c on ta in ing 1 50 a n ima ls ( 6 ,163 h r s ) . S im i la r ly , t he a nnua l l a bo r c os ts o fc a t t le s t ra teg ies o n ly i n c rease b y 5 7 h ou rs f r om h e rds c on ta in ing 1 0 a n ima ls ( 6 ,138 h r s ) t o h e rds c on ta in ing 2 00 h ead ( 6 ,195 h r s ) . F o rs heep , a n i n c rease o f5 1 h ours i se xpec ted b e tween f l ocks c on ta in ing 1 0 a n ima ls ( 4 ,744 h r s ) a nd f l ocks c on ta in ing 4 00 h ead ( 4 ,793 h r s ) . F o rg oa t h e rds , a n i n c rease o f 5 9 h ours i se xpec ted b e tween h e rds c on ta in ing 1 0 a n ima ls ( 4 ,744 h r s ) a nd h e rds c on ta in ing 4 00 h ead ( 4 ,803 h r s ) .

1 00

—1 0

20

C 13 1 : 1 1( 103E I E IO8V -1Tif 1NNV

(m illions o f c a lor ies )

ANNUAL CALOR IC Y IELD

25 -

-

c att le 1 5 —

c a me ls g oats

10 —

T- 1 - 1 - 1 -5 - 1 - r,

50

1 00

MT- 1- 1 - 1

150

I

200

r

250

n

j

300

-r

r

GOATS r I

350

, ,

U,

400

NU MBER OF AN I MALS I N HERD

F i gu re 8 . R e la t ionsh ips b e tween a nnua l c a lo r ic p roduc t iv i ty a nd r equ i red l abor f o r mea t -on ly p as to ra l s t ra teg ies . L abo r c os ts ( b roken l i nes ) r ead o f f t he s ca le o n t he r i gh t . Ca lo r ic b ene f i ts ( so l id l i nes ) r ead o f f t he s ca le o n t he l e f t . P o ten t ia l i n c reases i nc a lor ic b ene f i ts when p oss ib le p os t -P le is tocene s i ze r educ t ions a re a ccoun ted f o ra re i n d ica ted b yt he d iagona l s had ing .

0

50

100 150 200 250 300 NU MBER OF AN I MALS I N HERD

350

400

F i gu re 9 . R anges o fp o ten t ia l r e turn r a tes f o r mea t -on ly p as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts . I n c reases i nr e tu rn r a tes d ue t o g rea te r c a lo r ic y i e ld when p o ten t ia l p os t -P le is tocene s i ze r educ t ions a re a ccoun ted f o ra re i n d ica ted b yt he d iagona l s had ing . 1 01

F igu re 9 d iag rams e xpec ted r anges o f p o ten t ia l mea t -on ly r e tu rn r a tes f o r c ame l , c a t t le , s heep a nd g oa t h erds o f v a r ious s i zes , b ased u pon t he c os t a nd b ene f i t d a ta p resen ted i n F i gu re 8 . T he r e tu rn r a tes f o r c ame ls a re t hose e xpec ted u nder c ond i t ions o f n o rma l o f f take . T he d iagona l ly s haded a reas r ep resen tt he e xpec ted r ange o fr e tu rn r a tes w hen a n i n c rease i n mea t p roduc t iv i ty u pt o2 0% f o rc a t t le a nd 3 0% f o rs heep a nd g oa ts i sa ccoun ted f o r .

C ompara t ive R e tu rn R a tes f o r Mea tP lus M i lk P as to ra lS t ra teg ies F igure 1 0 d iag rams t he r e la t ionsh ips e xpec ted b e tween a nnua l c a lo r ic p roduc t iv i ty ( so l id l i nes ) a nd l a bo r r equ i remen ts ( b roken l i nes ) f o r mea t p l us m i lk p as to ra l s t ra teg ies i n vo lv ing h e rds o f v a r ious s i zes . T he l a bo r a nd p roduc t iv i ty d a ta f o r c ame ls r e f lec t e xpec ta t ions u nde r c ond i t ions o f n o rma l m i lk y i e ld a nd n o rma l o f f take . A ga in , t he d i agona l ly s haded a reas r e f lec t e xpec ted c a lo r ic i n c reases when p o ten t ia l s i ze r educ t ions a re a ccoun ted f o r . I n mea t p l us m i lk s t ra teg ies , l a bo r c os ts e xh ib i t ag rea te r s l ope b y h e rd s i ze t han i n mea t -on ly s t ra teg ies d ue t o t he a dd i t iona l l a bo rr equ i red w i th i n c reas ing m i lk y i e lds . B ased u pon t hese c os t a nd b ene f i t d a ta , F i gu re 1 d i ag rams c a lcu la ted r anges o fp oss ib le mea t p l us m i lk r e turn r a tes f o rc ame l , c a t t le , s heep a nd g oa t h e rds o fv a r ious s i zes . F o rc a t t le , t he d i agona l ly s haded a rea r e f lec ts t he p o ten t ia l 7 .7% i n c rease i nr e tu rn r a tes e xpec ted when a 2 0% s i ze r educ t ion i sa ccoun ted f o ri nc a lcu la t ions o fs l augh te ro f f take . S im i la r ly , t he d i agona l ly s haded a reas a ssoc ia ted w i th s heep a nd g oa t r e tu rn r a tes r e f lec tt he p o ten t ia l 8 .9% i n c rease i n s heep r e tu rn r a tes a nd 6 .2% i n c rease i ng oa t r e turn r a tes when s i ze r educ t ions o f 3 0% a re a ccoun ted f o r . A s n o ted e a r l ie r i n t h is c hap ter , w i th more f r equen t b reed ing u nde r f avorab le e nv i ronmen ta l c ond i t ions , s heep f l ock g row th r a tes may i n c rease b y a s much a s 1 9 .7% a nd g oa t h erd g row th r a tes b y a s much a s 2 9 .5% o ve r t he r a tes e xpec ted u nde r c ond i t ions o fo ne a nnua l b reed ing . H owever , e ven i ft hese i n c reases i n g row th r a tes p roduced e qu iva len t i n c reases i np roduc t iv i ty , e xpec ted r e turn r a tes w ou ld o n ly s l igh t ly i n c rease , s i nce t o ta l p as to ra l l a bo r wou ld a l so i n c rease . F o r e xamp le , h e rd ing c os ts w ou ld i n c rease a pp rox ima te ly 2 0% d ue t ot he a dd i t iona l h e rd s ubd iv is ions r equ i red b y as econd b reed ing s eason ( 3 ,656 h r s /y rv s 4 ,392 h rs /y r ) , w h i le m i lk ing a nd b u tche r ing t asks a re p ropo r t iona l t ot o ta l m i lk a nd s l augh te rp roduc t ion . On ly e ncampmen t c os ts a s c a lcu la ted h e re w ou ld r ema in c ons tan t . W i th af l ock o f 1 00 s heep , t o ta l a nnua l l abo r c os ts wou ld i n c rease a pp rox ima te ly 1 6 .4% , a nd a round 1 6 .7% f o r af l ock o f 4 00 h ead . W i th a h e rd o f 1 00 g oa ts , t o ta l a nnua l l a bo r wou ld i n c rease a pp rox ima te ly 1 6 .9 % , a nd a round 1 8 .3% f o r ah e rd o f4 00 h ead . A s ar esu l t , a1 9 .7% i n c rease i nt he t o ta l p roduc t iv i ty o f af l ock o fs heep d ue t o as econd b reed ing s eason w ou ld o n ly i n c rease r e tu rn r a tes b y a pp rox ima te ly 3% , wh i le a2 9 .7 % i n c rease i n t o ta l g oa t h e rd p roduc t iv i ty wou ld o n ly i n c rease r e tu rn r a tes b y a pp rox ima te ly 112 % . T he r anges o fp o ten t ia l mea t p l us m i lk r e tu rn r a tes p resen ted i nF i gu re 1 a ssume t ha tt he t o ta lc a lo r ic c on ten to ff r esh m i lk , i n c lud ing c a lo r ies d e r ived f r om l a c tose ( m i lk s uga r ) , a re u t i l ized b y t he p as to ra l p opu la t ion . F ur the r , t hey d o n o t a ccoun t f o re i the rt he l a bo r c os ts o fc onver t ing f r esh m i lk i n to s to rab le d a i ry p roduc ts , n o r a ny r educ t ion i n c a lo r ic c on ten t d u r ing m i lk p rocess ing . C ons is ten tw i th t he d i scuss ion p resen ted e a r l ie ri nC hap te r 2o n a du l t h ypo lac tas ia a nd m i lk f e rmen ta t ion p rocess ing , t he p o ten t ia l e f fec ts o ft hese f ac to rs o n p as to ra l r e turn r a tes w i l l b e a ccoun ted f o r .

M i lk P rocess ing a nd P as to ra lR e tu rn R a tes B ased u pon t he d a ta p rev ious ly p resen ted i nC hap te r 2o n t he f e rmen ta t ion p rocess ing o f m i lk , T ab le 1 4s umma r izes t he p e rcen tage r educ t ion e xpec ted i nt he t o ta l c a lo r ic p roduc t iv i ty o f c ame l , c a t t le , s heep a nd g oa t h e rds w hen a l m i lk i ss ou red p r io rt oc onsump t ion , a ssum ing t ha t a l l l a c tose i s h yd ro lyzed b u t o n ly t he g lucose h a l f o f t he l a c tose i s me tabo l ized b y m ic roo rgan isms . Wh i le s ou r ing m i lk may i n vo lve i n i t ia l ly h ea t ing f r esh m i lk t o " b lood " o r " body t empe ra ture " ( Bar th 1 960 :7 ; D ickson 1 951:402 ; Swee t 1 974 :104 ; Wa tson 1 979 :110 ) , p resumab ly t of o rce a mo re r ap id f e rmen ta t ion , i td oes n o t a ppea rt ob e n ecessa ry ( J enn ings 1 02

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300

350

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NUMBER OF AN I MALS I N HERD

F i gure 1 0 . R e la t ionsh ips b e tween a nnua l c a lo r ic p roduc t iv i ty a nd r equ i red l a bo rf o r mea t p l us m i lk p as to ra l s t ra teg ies . L abo r c os ts ( b roken l i nes ) r ead o f f t he s ca le o n t he r i gh t . C a lo r ic b ene f i ts ( so l id l i nes ) r ead o f t he s ca le o n t he l e f t . I n c reases i nc a lo r ic b ene f i ts w hen p oss ib le p os t -P le is tocene s i ze r educ t ions a re a ccoun ted f o ra re i n d ica ted b yt he d i agona l s had ing .

3000

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T o 2000 SHEEP 1500 c GOATS f _ 1 000

500

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50

100

1 50

200

250

300

350

400

NUMBER OF AN I MALS I N HERD F i gure 1. R anges o f p o ten t ia l r e turn r a tes f o r mea t p l us m i lk p as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts . E xpec ted i n c reases i n r e tu rn r a tes w hen p o ten t ia l p os t P le is tocene s i ze r educ t ions a re a ccoun ted f o r a re i n d ica ted b yt he d i agona l s had ing . 1 03

T ab le 1 4 E xpec ted D ec l ine i nt he T o ta l C a lo r ic P roduc t iv i ty o fP as to ra l H e rds When One -Ha l fo ft he L ac tose C on ten to fT o ta l H e rd M i lk P roduc t ion i sL os td ue t oF e rmen ta t ion P rocess ing

AN IMAL

%o fT o ta l M i lk C a lo r ies D e r ived F rom L ac tose C on ten t

%o fT o ta lA nnua l P as to ra l C a lo r ic P roduc t iv i ty F rom M i lk

E xpec ted ` ) / 0D ec l ine I nT o ta l P as to ra l C a lo r ic P roduc t iv i ty When 1 / 2 o f L ac tose i nM i lk i sL os t

C ame ls C a t t le S heep G oa ts

2 9 .3 2 3 .4 2 0 .9 2 6 .3

7 4 .9 6 1.6 7 0 .5 7 9 .2

1.0 7 .2 7 .4 1 0 .4

B ram ley 1 907b :133 ; Mus i l 1 928 :89 ; N ico la isen 1 963 :22 1-22 ; S . Sm i th 1 980 :469 ) . N ico la isen s pec i f ica l ly n o ted t ha t f r esh m i lk was s imp ly p u t i n to t he s k in b ags u sed f o r s our ing , w i th t he b ags p laced i nt he s un o r n ea r af i re d ur ing t he w in te r , o r p l aced i nt he s hade o ft en ts i nt he s umme r ( 1963 :222 ) . W i th l i t le l a bo r r equ i red , t he t o ta l e ne rge t ic e f fec to fs our ing a l m i lk wou ld c ons is t o f ap e rcen tage r educ t ion e qu iva len tt ot he c a lo r ic r educ t ions p resen ted i nT ab le 1 4 . A s n o ted e a r l ie r , p o ten t ia l i n c reases i ns l augh te rp roduc t iv i ty d ue t ol a rge r a n ima ls i ne a r ly p as to ra l s t ra teg ies w ou ld h ave i n c reased mea t p l us m i lk c a t t le h e rd r e tu rn r a tes b y 7 .7 % o ve r t hose e xpec ted e thnog raph ica l ly , s heep f l ock r e tu rn r a tes b y 8 .9 % a nd g oa t h e rd r e turn r a tes b y 6 . 2% , w h i le c ame l h e rd r e tu rn r a tes wou ld n o t h ave c hanged . I ft he r educ t ions i nr e tu rn r a tes p resen ted i nT ab le 1 4 s imu l taneous ly c ha rac te r ized e a r ly p as to ra l s t ra teg ies d ue t o ar equ i red s our ing o f a l m i lk i na ssoc ia t ion w i th l a c tose ma labso rbe rs , e xpec ted r e turn r a tes f o r e a r ly p as to ra l s t ra teg ies r e la t ive t ot hose c a lcu la ted e a r l ie rf o rt he e t hnog raph ic p resen t w ou ld h ave b een 0 . 5% g rea te rf o re a r ly c a t t le h e rds , 1 . 5% g rea te rf o re a r ly s heep f l ocks , 4 .2 % l e ss f o re a r ly g oa t h e rds , a nd 1 % l e ss f o re a r ly c ame l h e rds . S ou red m i lk i st he p re l im ina ry med ium w h ich i sc hu rned f o r t he p roduc t ion o f c u l tured b u t te rm i lk a nd b u t te r . P laced i n to s k in b ags o f v a r ious s i zes , t he s ou ged m i lk i sc hurned b y r o l l ing , s hak ing o r r ock ing t he b ag b ack a nd f o r th . S ome t imes , t he c hurn ing b ag i ss uspended f r om at r ipod o rs t ick b race ( Ba r th 1 96 1 :8 ; D ickson 1 95 1:402 ; Wa tson 1 979 :104 ) , p a r t icu la r ly i f i ti sl a rge ( N ico la isen 1 963 :222 ) . Sma l l b a l ls o r b i ts o f b u t te r a re f o rmed wh ich f l oa t t o t he s u r face , wh i le t he r e s t o ft he s ou red m i lk b ecomes b u t te rm i lk . T he b u t te rg lobu les may b e u sed d i rec t ly , o r me l ted a nd a lowed t o c oo l . A cco rd ing t o Swee t ( 1974 :104 ) , c hu rn ing b y r o l l ing a s k in b ag b ack a nd f o r th t a kes " an h ou ro r mo re . " A cco rd ing t o Wa tson ( 1979 : 110 ) , c hu rn ing i na s k in b ag s uspended f r om at r ipod r equ i res " one -ha l f h our o r s o . " F i na l ly , D ickson ( 1951:402 ) s t a ted t ha t c hu rn ing w i th a s uspended s k in b ag r equ i res " an h ou ro r mo re . " P resumab ly , s uch f ac to rs a s f a t c on ten t , d eg ree o ff e rmen ta t ion , a nd t he q uan t i ty o fs oured m i lk b e ing p rocessed w ou ld a l te rt he r equ i red c hu rn ing t ime . C ame l 's m i lk , h oweve r , i s r a re ly i fe ve r c hurned f o r b u t ter a nd b u t term i lk ( D ickson 1 95 1 :4 16 ; L eese 1 927 : 140 ; Mus i l 1 928 :89 ; R ao e ta l . 1 970 :74 ) . I nc ame l 's m i lk , t he f a t i sf i rm ly b onded t ot he p ro te ins ( Rao e t a l . 1 970 :74 ) . I nt he e xpe r imen ta l c hu rn ing o f s oured c ame l 's m i lk , o n ly 2 o z ( 56 .7 gm ) o fb u t te r w e re o b ta ined f r om 3 l b( r ough ly 1 . 4 k g ) o f m i lk a f ter f ou r h ou rs o fc hu rn ing ( Leese 1 927 : 140 ) . A s s uch , t he p rocess ing o f s ou r m i lk b y c hu rn ing i so n ly a pp rop r ia te f o rc a t t le , s heep a nd g oa t s t ra teg ies . I no rde rt oa ccoun tf o rt he a dd i t iona l l a bo ro fs uch p rocess ing , i tw i l l b e a ssumed t ha t 2 5% o fa l m i lk p roduc t ion f r om c a t t le , s heep a nd g oa t h e rds i sc hurned i n to b u t te ra nd b u t te rm i lk a ta r a te o f 1 . 5 h ou rs o fl a bo rf o re ve ry 4k go fs ou red m i lk ( r ough ly 4l i t ers , t he a pp rox ima te amoun t o fm i lk w h ich asma l l c hu rn ing b ag w ou ld h o ld ) . T he e xpec ted i n c rease i nt o ta l a nnua l p as to ra l l a bo r amoun ts t o a pp rox ima te ly 1 , 276 h ou rs f o r c a t t le h e rds c on ta in ing 2 00 h ead ( 13 ,606 k g m i lk x0 .25 = 3 ,40 1 .5 k g p rocessed 4 k g p e r b a tch = 8 50 .4 b a tches x 1 . 5 h rs = 1 , 275 .6 h r s ) . 1 04

F o rs heep f l ocks c on ta in ing 4 00 h ead , 6 09 a dd i t iona l h ou rs wou ld b e r equ i red ( 6 ,497 k g m i lk x 0 .25 = 1 ,624 .3 k g p rocessed 4 k g p e r b a tch =4 06 .1 b a tches x 1 . 5 h rs = 6 09 .2 h rs ) . F o rg oa t h e rds c on ta in ing 4 00 h ead , 8 56 a dd i t iona l h ours wou ld b e r equ i red ( 9 ,132 k g m i lk x 0 .25 = 2 ,283 k g p rocessed +4k gp e rb a tch =5 70 .8 b a tches x1 . 5 h rs =8 56 .2 h rs ) . D r ied m i lk i t e ms f o r l o ng te rm s t o rage a re a l so p roduced b y many p as to ra l a nd s eden ta ry p opu la t ions . B u t te rm i lk " cheeses " o r " m i lk c akes " a re p roduced b y h ea t ing c u l tured b u t te rm i lk u n t i l t he s o l ids c u rd le ( N ico la isen 1 963 :224 ; Wa tson 1 979 :110 ) o r u n t i l t he l i qu id i sr educed l e av ing a t h ick p as te ( Swee t 1 974 :105 ) . T he r es idues a re t hen s haped a nd d r ied i nt he s un . A cco rd ing t o D ickson ( 195 1 :402 ) , as im i la rd r ied p roduc t i sp roduced b y s imp ly b o i l ing s ou red m i lk " f o r a t l e as t 2h ours u n t i l i tb ecomes t h ick a nd g ranu la ted , " a f te r wh ich t he r es idue i s s haped a nd d r ied . T h is p rocess was a ppa ren t ly u sed b y t he Rwa la t o make " cheeses " f r om s oured c ame l 's m i lk when t he a va i lab le m i lk s upp ly e xceeded t he i r a b i l i ty t o u t i l ize i t( Mus i l 1 928 :89 ) . B ar th r eco rded t ha t s im i la r p roduc ts we re p roduced f r om s our m i lk " by s imp le p ress ing i n ag auze l ike b ag " wh ich s e rved t os epa ra te t he " cu rds "f r om " sou r whey " ( 196 1 :89 ) . B o th c ou ld t hen b e d r ied f o r s t o rage , a l though t he " sour whey " was f r equen t ly d i sca rded b y t he t r iba l s ec t ion wh ich h e s t ud ied . S uch d r ied p roduc ts c an a ppa ren t ly b e s t o red f o r a tl e as t a y ea r ( Bar th 1 961 :8 ; Wa tson 1 979 :110 ) , a nd a re c rumb led a nd r econs t i tu ted w i th wa te r f o r c onsump t ion ( D ickson 1 95 1 :402 ; Swee t 1 974 :105 ; Wa tson 1 979 :110 ) . T he d r ied " cheeses " m en t ioned b y Mus i l ( 1928 :89 ) a nd n o ted b y J enn ings -B ram ley f r om h ea rsay ( 1907b :135 ) a re t he o n ly r e fe rences n o ted f o r a ny l o ng te rm s t o rage i t em p roduced f r om c ame l 's m i lk . U n l ike c a t t le , s heep a nd g oa ts , c ame ls w i l l s upp ly m i lk v i r tua l ly y ea r round , a nd t he s torage o f m i lk p roduc ts a ga ins t l e an s easons w i thou t m i lk w ou ld p resumab ly b e l e ss c r i t ica l . I tw i l l b e a ssumed t ha tu nde rc ond i t ions o fn o rma l m i lk p roduc t iv i ty , 5% o ft o ta l p as to ra l m i lk p roduc t ion i sp rocessed i n to d r ied m i lk p roduc ts , r ega rd less o ft he h e rd s pec ies i n vo lved . A t a r a te o f2 .5 h ou rs f o re ve ry 4k g o fs ou red m i lk , t o ta l a nnua l l a bo rc os tf o r ac ame l h e rd c on ta in ing 1 50 h ead wou ld i n c rease a pp rox ima te ly 7 89 h ours ( 25 ,257 k g m i lk x 0 .05 = 1 , 262 .9 k g p rocessed + 4 k g p e r b a tch = 3 15 .7 b a tches x 2 .5 h rs = 7 89 .3 h rs ) . F o r ah e rd o f c a t t le c on ta in ing 2 00 h ead , a n a dd i t iona l 4 25 h ours o f l a bo r w ou ld b e r equ i red ( 13 ,606 k g x0 .05 = 6 80 .3 k g +4k g p e r b a tch = 1 70 .1 b a tches x 2 .5 h rs = 4 25 .3 h rs ) . F o r af l ock o f s heep c on ta in ing 4 00 h ead , a n a dd i t iona l 2 03 h ou rs wou ld b e r equ i red ( 6 ,497 k g m i lk x0 .05 = 3 24 .9 k g p rocessed + 4 k g p e r b a tch = 8 1 .2 b a tches x 2 .5 h rs = 2 03 h rs ) . F o r ah e rd o f g oa ts c on ta in ing 4 00 h ead , a n a dd i t iona l 2 86 h ou rs o fl a bo r wou ld b e r equ i red ( 9 ,132 k g m i lk x0 .05 = 4 56 .6 k g +4 k g p e rb a tch = 14 .2 b a tches x2 .5 h rs = 2 85 .5 h rs ) . W i th t he p rocess ing d a ta g ene ra ted a bove , t he e ne rge t ic e f fec ts o f m i lk p rocess ing o n p as to ra l r e tu rn r a tes c an b e p a r t ia l ly a ccoun ted f o r . I n i t ia l ly , t he s ou r ing o fa l m i lk wou ld r educe t o ta l a nnua l c a lo r ic p roduc t iv i ty b y 1%f o rc ame l h e rds , 7 . 2% f o rc a t t le h e rds , 7 .4% f o rf l ocks o f s heep a nd 1 0 .4% f o r h e rds o fg oa ts . I na dd i t ion , t he p as to ra l l a bo r c os ts a ssoc ia ted w i th a c ame l h erd c on ta in ing 1 50 h ead wou ld i n c rease a pp rox ima te ly 9 . 1% d ue t ot he a dd i t iona l l a bo r r equ i red i np rocess ing 5% o ft he t o ta l m i lk y i e ld i n to d r ied m i lk p roduc ts ( 789 a dd i t iona l h is 8 ,689 h is w i thou t s uch p rocess ing = 0 .0908 ) . T he l a bo r c os ts o f ac a t t le h e rd c on ta in ing 2 00 h ead wou ld i n c rease a pp rox ima te ly 2 2 .6 % d ue t ot he a dd i t iona l l abo r o f 1 , 276 h ou rs r equ i red i n p rocess ing 2 5% o f t he t o ta l m i lk y i e ld i n to b u t te r a nd b u t te rm i lk , a nd t he 4 25 h ours e xpended i np rocess ing 5% o ft he t o ta l m i lk y i e ld i n to d r ied m i lk p roduc ts ( 1 ,276 + 4 25 = 1 , 701 h is + 7 ,526 h rs w i thou t s uch p rocess ing = 0 .226 ) . T he l a bo r c os ts o f as heep f l ock c on ta in ing 4 00 h ead wou ld i n c rease a pp rox ima te ly 1 4 .9 % d ue t o t he a dd i t iona l 6 09 h ours r equ i red i n s im i la r ly p roduc ing b u t te r a nd b u t te rm i lk , a nd t he 2 03 h ou rs r equ i red i n mak ing d r ied m i lk p roduc ts ( 609 + 2 03 = 8 12 h r s +5 ,443 h rs w i thou t s uch p rocess ing = 0 . 149 ) . F i na l ly , t he l a bo r c os ts a ssoc ia ted w i th a h e rd o fg oa ts c on ta in ing 4 00 h ead wou ld i n c rease a pp rox ima te ly 2 0% d ue t o t he a dd i t iona l 8 56 h ou rs r equ i red i n p rocess ing b u t te r a nd b u t term i lk , a nd t he 2 86 h ou rs r equ i red i n mak ing d r ied m i lk p roduc ts ( 856 + 2 86 = 1 , 142 h rs + 5 ,7 16 h rs o f l abo r w i thou t s uch p rocess ing = 0 .2 ) . F o r ac ame l h e rd c on ta in ing 1 50 h ead u nde r c ond i t ions o f n o rma l m i lk p roduc t ion , t he e xpec ted r e turn r a te o f2 ,600 k ca l /h r ( 22 ,588 ,82 1 k ca l +8 ,689 h rs ) w ou ld d rop t o2 , 12 1 k caVh r ( 20 ,104 ,051 k ca l 4 9 ,478 h r s ) w i th m i lk p rocess ing , an e td ec l ine i nr e tu rn r a te o fa pp rox ima te ly 1 8 .4 % ( 2 ,600 -2 ,121 = 4 79 + 2 ,600 = 0 . 184 ) . F o r ac a t t le h e rd c on ta in ing 2 00 h ead , t he e xpec ted r e turn r a te o f2 ,370 k ca l /h r ( 17 ,835 ,090 k ca l 7 ,526 h rs ) w ou ld d rop t o 1 ,794 k ca l /h r -

4 -

1 05

( 16 ,550 ,964 k ca l 9 ,227 h rs ) w i th m i lk p rocess ing , an e t d ec l ine i nr e tu rn r a te o fa pp rox ima te ly 2 4 .3 % ( 2 ,370 1 ,794 = 5 76 2 ,370 = 0 .243 ) . F o r af l ock o f s heep c on ta in ing 4 00 h ead , t he e xpec ted r e turn r a te o f1 , 856 k ca l /h r ( 10 ,102 ,265 k ca l F5 ,443 h r s ) wou ld d rop t o 1 , 496 k ca l /h r ( 9 ,354 ,698 k ca l I -6 ,225 h rs ) w i th m i lk p rocess ing , an e t d ec l ine i nr e tu rn r a te o f a pp rox ima te ly 1 9 .4 % ( 1 ,856 -1 ,496 = 3 60 f -1 , 856 = 0 . 194 ) . F i na l ly , f o r ah e rd o f 4 00 g oa ts , t he e xpec ted r e tu rn r a te o f 1 , 347 k ca l /h r ( 7 ,698 ,665 k ca l 5 ,7 16 h r s ) wou ld d rop t o 1 , 006 k ca l /h r ( 6 ,898 ,004 k ca l 6 ,858 h rs ) w i th m i lk p rocess ing , an e t d ec l ine i n r e tu rn r a te o f a pp rox ima te ly 2 5 .3 % ( 1 ,347 1 ,006 = 3 4 1 i -1 , 347 = 0 .253 ) . A ssum ing t ha tt he r anges o fp as to ra l r e tu rn r a tes p rev ious ly p resen ted i nF i gu re 1 w ou ld r ep resen t t he u ppe r r ange o f p o ten t ia l p reh is to r ic r e tu rn r a tes , t he d a ta g ene ra ted o n m i lk p rocess ing c an b e u sed t o e s tab l ish a l owe r r ange f o r p o ten t ia l p reh is to r ic r e tu rn r a tes . T he e xpec ted c a lo r ic l o ss d ue t o f e rmen t ing m i lk a nd t he a dd i t iona l l a bo r h ours r equ i red f o r p rocess ing m i lk h ave b een u sed t o r eca lcu la te t he b ase l ine r e tu rn r a tes p rev ious ly p resen ted i n F igu re 1 , e xc lud ing t he a dd i t iona l e ne rgy r e tu rns e xpec ted a s ar esu l t o f p o ten t ia l s i ze r educ t ions among c a t t le , s heep a nd g oa ts . F i gu re 1 2 d iag rams t he r esu l ts o b ta ined . F o rc ame l s t ra teg ies , t he s o l id l i ne a tt he u ppe r e nd o fp o ten t ia l r e turn r a tes r e f lec ts t hose e xpec ted w hen a l c a lo r ies f r om f r esh m i lk a re u t i l ized , a s p rev ious ly p resen ted i nF i gu re 1. T he s o l id l i ne a tt he l owe r e nd o fp o ten t ia l r e turn r a tes r e f lec ts t hose e xpec ted when 5 0% o fa l c a lo r ies a va i lab le i nt he l a c tose c on ten t o ff r esh m i lk a re l o s tt h rough t he f e rmen ta t ion o fa l m i lk p roduced , a nd w hen 5% o f t o ta l m i lk p roduc t ion i sp rocessed i n to d r ied m i lk p roduc ts . T he v e r t ica l ly s haded a rea t hus r ep resen ts t he r ange o f p o ten t ia l r e turn r a tes b e tween t hese t wo e x t remes . F o rc a t t le , s heep a nd g oa t s t ra teg ies , t he r anges o fp o ten t ia l r e tu rn r a tes r ep resen ted b y t he d iagona l ly s haded a reas a re t hose p rev ious ly p resen ted i nF i gu re 1 , a nd r e f lec t e xpec ted r e turn r a tes w hen a l c a lo r ies f r om f r esh m i lk a re u t i l ized a nd s i ze r educ t ions u pt o2 0% f o rc a t t le 3000 — _

RETURN RATE ( kca l/hr )

2500-

CAMELS CATTLE

2000-

1500— _

1000GOATS 500:

0 0

50

100

150

200

250

300

350

400

NUMBER OF AN I MALS I N HERD F i gu re 1 2 . R anges o f p o ten t ia l r e turn r a tes f o r mea t p l us m i lk p as to ra l s t ra teg ies when m i lk p rocess ing i sa ccoun ted f o r . T he u ppe rs o l id l i ne o ft he r ange o fr e tu rn r a tes f o rc ame ls a nd t he d iagona l ly s haded a reas f o rc a t t le , s heep a nd g oa t r e tu rn r a tes a re t hose p rev ious ly p resen ted i nF i gu re 1 . T he v e r t ica l ly s haded a reas r e f lec t l owe rr e turn r a tes d ue t ot he l o ss o fc a lo r ies a nd a dd i t iona l l a bo rc os ts a ssoc ia ted w i th m i lk p rocess ing . 1 06

a nd 3 0% f o rs heep a nd g oa ts a re a ccoun ted f o r . T he v er t ica l ly s haded a reas r e f lec t t he r ange o fe xpec ted r e tu rn r a tes w hen c a lo r ies a re l o s td ue t om i lk f e rmen ta t ion a nd a dd i t iona l l a bo ri s e xpended i nm i lk p rocess ing . T he l owe r l i nes o ft he v er t ica l ly s haded a reas r e f lec t e xpec ted r e turn r a tes w hen 5 0% o f a l c a lo r ies f r om t he l a c tose o f f r esh m i lk a re l o s t d ue t o t he f e rmen ta t ion o fa l m i lk o b ta ined f r om ah e rd , a nd w hen 2 5% o fa l m i lk i sc hu rned i n to b u t te ra nd b u t te rm i lk a nd 5% o fa l m i lk i sp rocessed i n to d r ied m i lk p roduc ts . E xpec ted i n c reases i nt o ta l c a lo r ic p roduc t iv i ty d ue t o p o ten t ia l s i ze r educ t ions h ave b een e xc luded f r om t hese l a t ter c a lcu la t ions . T he r anges o fp o ten t ia l p as to ra l r e tu rn r a tes p resen ted i nF i gure 1 2 w i l l b e s ubsequen t ly u sed i n C hap ter 7 t o g ene ra te e xpec ta t ions c once rn ing t he e vo lu t ion o f e a r ly p as to ra l s t ra teg ies . I n t he f o l low ing c hap te r , c ompa rab le r e turn r a tes f o r c e rea l c u l t iva t ion w i l l b e e s tab l ished .

1 07

CHAPTER 6 RELAT IVE E NERGET ICS OF E ARLY WHEAT AND BARLEY S TRATEG IES

I n t roduc t ion I nt h is c hap te r , t he r e la t ive e ne rge t ics o f whea t a nd b a r ley e xp lo i ta t ion a nd c u l t iva t ion w i l l b e e xam ined i no rde r t o e s tab l ish r anges o f p o ten t ia l r e tu rn r a tes f o r e a r ly c e rea l s t ra teg ies c ompa rab le t o t hose e s tab l ished i n C hap te r 5f o r e a r ly p as to ra l s t ra teg ies . A f te r a n i n i t ia l d i scuss ion o fw hea t a nd b a r ley y i e ld d a ta f r om s pa t ia l ly a nd t empo ra l ly d i ve rse l i t e ra ry s ou rces , p roduc t iv i ty d i v is ions b e tween b road ly d e f ined c a tegor ies o f " op t ima l " a nd " ma rg ina l " a g r icu l tu ra l l a nds a re e s tab l ished f o r u se i ns ubsequen t e ne rge t ic c ompa r isons b e tween e a r ly p as to ra l a nd c e rea l s ubs is tence o p t ions . T he l abo rc os ts o fw hea t a nd b a r ley c u l t iva t ion u nde r c ond i t ions o f h and t i lage w i th l i t h ic t e chno log ies , a nd l a te r a rd t i lage w i th e a r ly me ta l t echno logy , a re t hen d ocumen ted . C a lo r ic r e tu rn r a tes a re s ubsequen t ly c a lcu la ted f o r t he e xp lo i ta t ion o f s t ands o fw i ld e i nko rn whea t , a nd f o r t he e a r ly c u l t iva t ion o f whea t a nd b a r ley g i ven t empo ra l v a r ia t ions i n a g r icu l tura l t e chno logy . C ompa ra t ive d a ta o n t he e ne rge t ics o fp ea r l m i l le t a nd s o rghum c u l t iva t ion i n Wes t A f r ica u nder c ond i t ions o ft r ad i t iona l h and c u l t iva t ion a re a l so p resen ted .

C u l t iva ted Whea ta nd B a r ley Y ie lds I n t roduc t ion I n C hap te r 3 , y i e lds r epo r ted f o r c u l t iva ted whea ts i nt empo ra l ly a nd s pa t ia l ly d i sc re te l i t e ra ry s ources w e re e xam ined r e la t ive t oe s t ima tes o fm ixed whea t a nd b a r ley y i e lds i nn a tura l s t ands i no rder t o e va lua te t he p o ten t ia l o f i n c reased c e rea l p roduc t iv i ty d u r ing t he m i l lenn ia w h ich h ave e l apsed s i nce e a r ly c u l t iva t ion s t ra teg ies i n t ia l ly e vo lved . I twas d e te rm ined a tt ha t t i me t ha t whea ts , l i ke o the r a nnua l c e rea ls , e xh ib i t t r emendous p heno typ ic p l as t ic i ty i nt he i r p roduc t ive p o ten t ia l r e la t ive t ot he c ond i t ions u nde rw h ich t hey a re g rown , a nd t ha tt he re wou ld a ppea r t o b e n o v a r ia t ions i nw hea t y i e lds p r io r t o t he l a te e i gh teen th a nd e a r ly n i ne teen th c en tur ies A .D . wh ich c ou ld n o t b e a t t r ibu ted t o r eg iona l d i f fe rences i ns o i l f e r t i l i ty , t he u se o f f e r t i l ize r , c l ima te , a nd r a in fa l lo ri r iga t ion . I n t he f o l low ing s ec t ions , p roduc t iv i ty d a ta o n c u l t iva ted w hea ts a nd b a r leys w i l l b e e xam ined i ng rea te rd ep th i no rde rt oe s tab l ish r anges o fp o ten t ia l y i e lds f o ru se i ns ubsequen t e ne rge t ic c a lcu la t ions f o r e a r ly c e rea l s t ra teg ies . I n p ar t icu la r , a t ten t ion w i l l f ocus u pon e s tab l ish ing a n a pp rop r ia te mean p roduc t iv i ty d i v is ion b e tween l a nds o f ma rg ina l a nd o p t ima l q ua l i ty . I n i t ia l ly , a nnua l v a r ia t ions a nd l o ng te rm means i nc u l t iva ted w hea t a nd b a r ley y i e lds w i l l b e e xam ined b y r e fe rence t ot he Med ieva l r e co rds c omp i led b y T i tow ( 1972 ) .

Med ieva l Whea ta nd B a r ley Y i e lds i nt h e W inches te rB ishop r ic A s d iscussed i n C hap ter 4 , T i tow ( 1972 ) c omp i led a g r icu l tura l y i e ld r ecords f o r a pp rox ima te ly 4 1 mano rs i nt he W inches te r B i shop r ic b e tween L ondon a nd B r is to l d u r ing t he t h ir teen th a nd f our teen th c en tur ies A .D . F i gure 1 3 d iag rams v a r ia t ions i nt he mean a nnua l y i e lds o f whea t a nd b a r ley i nt h is r eg ion d u r ing t he 7 1 y ea rs b e tween 1 232 a nd 1 349 f o rw h ich T i tow p rov ided d a ta ( 1972 : A ppend ix G , A ppend ix H ) . Med ieva l y i e lds /ac re h ave b een c onver ted t o k g /du a ssum ing a 3 95 l b( 179 .17 k g ) Med ieva l q ua r te r ( T i tow 1 972 :148 ) a nd a s t a tu te a c re e qua lt o4 .05 d u .

BARLEY Mean S td. dev . Med ian M in . Max . Range

= 80.5 = 12.1 = 82.4 = 52.7 = 109.8 = 57.1

Y IELD ( kg)/ DU NU M

0 10 20

h

0

10

20

W HEAT Mean S td . dev . Median M in.

= 52.8 = 9 .0 = 53.5 = 29.2

Max . = 71.7 Range = 42.5 0 10 11177 m iss ing data

1240

1260

20 1280

1300

1320

0

10

20

1340

YEAR

F i gu re 1 3 . A nnua l d ev ia t ions i n mean whea t a nd b a r ley y i e lds i nt he W inches ter B i shop r ic f r om t he c umu la t ive mean f o rt he y ea rs A .D . 1 232 t h rough 1 349 , a s d e r ived f r om d a ta c omp i led b y T i tow ( 1972 : A ppend ix G , A ppend ix H ) . A l though t h is f i gu re s pans a 17 y ea r p e r iod , y i e ld d a ta f o r4 6o ft hese a re m iss ing . H ence , o n ly y i e lds f o r7 1 y ea rs a re a c tua l ly r ep resen ted .

I n i t ia l ly , i ti sa ppa ren tt ha t y ea rt oy ea rv a r ia t ions i nd ry fa rm ing y i e lds c an b e q u i te d rama t ic , w i th r e la t ive ly b r ie f c l us te rs o f a bove o r b e low mean y i e lds f r equen t ly o ccu r r ing . F ur the r , i ti s a ppa ren tt ha t a l though a bove o rb e low mean w hea t a nd b a r ley y i e lds o f ten t r ack o ne a no the r i n t he s ame y ea r , t hey d o n o t a lways d o s o e qua l ly . A t o the rt i mes , a bove mean y i e lds i no ne c rop a re m i r ro red b y b e low mean y i e lds i nt he o the r . P a r t icu la r ly s t r ik ing i nt h is r espec t a re t he v a r ia t ions i n whea t a nd b a r ley y i e lds wh ich o ccu r red b e tween 1 335 a nd 1 348 , t he y ea rs i mmed ia te ly p reced ing t he a r r iva l o f B ubon ic P lague i n E ng land i n 1 349 . G iven t he e r ra t ic n a ture o fy ea rt oy ea r p roduc t iv i ty , i tw ou ld s eem p robab le t ha t i ne va lua t ing t he r e la t ive c os ts a nd b ene f i ts o f c e rea l s ubs is tence o p t ions , h umans w ou ld g ene ra l ly f ocus u pon l o ng te rm means o r e xpec ted r anges o fv a r iab i l i ty o ve r ap e r iod o fs eve ra l y ea rs , r a the rt han b as ing t he i r s ubs is tence d ec is ions o n s hor t term ( c rop t oc rop ) e va lua t ions . 10

Whea ta nd B ar ley Y i e lds i nt h eN ear E as t A s n o ted i nC hap te r3 ,t h e mean w hea t y i e ld o f5 2 .8 k g /du d e r ived f r om T i tow 's Med ieva l B r i t ish d a ta c l ose ly a pp rox ima tes t he 5 3 .5 k g /du mean w hea ty i e ld d e r ived f r om C o lume l la 's f i r s t c en tu ry A .D . d a ta o n l a nd o fa ve rage q ua l i ty i nR oman I t a ly ( 1 .9 .1 ,1 1 1 .3 .4 ) . T hese f i gu res a l so f a l l c l ose t ot he 5 0 k g /du a ve rage w hea t y i e ld c a lcu la ted b y Maye rson ( 1955 :55 , 1 960 :18 ) f r om s even th c en tury A .D . p apy r i r ecove red f r om t he B yzan t ine t own o f N essana i nt he N egev d ese r to fs ou the rn I s rae l . T he p oss ib i l i ty t ha t aw hea t y i e ld o fa round 5 0 -54 k g /du may r e f lec t a r ea l is t ic p roduc t iv i ty d i v is ion b e tween t he y i e lds e xpec ted o n " ma rg ina l " a nd " op t ima l " l a nds f o r whea t c u l t iva t ion w ou ld a l so g a in s uppor t f r om Maye rson 's i n fo rman t d a ta o n t he y i e lds e xpec ted b y b edou in c u l t iva to rs i nt he c en t ra l N egev i nt he l a te 1 950s . T ab le 1 5 s umma r izes t h e e xpec ted w hea t a nd b a r ley y i e ld d a ta c o l lec ted b y Maye rson f r om h i s b edou in i n fo rman ts ( 1 960 :18 ) . A s c an b e s een , a mean w hea t y i e ld o fa round 5 0 -54 k g /du a ccords w e l l w i th t he e xpec ta t ions o f b edou in c u l t iva to rs c once rn ing t he d i v is ion b e tween w ha t t hey c ons ide red " sa t is fac tory " a nd " good " y ea rs . A s f u r the rd ocumen ted b e low , a5 0 k g /du mean y i e ld f o rw hea t d oes s eem q u i te a pp rop r ia te f o r e s tab l ish ing a p roduc t iv i ty d i v is ion b e tween b road ly c once ived c a tego r ies o f" op t ima l " a nd " ma rg ina l " a g r icu l tura l l a nds . H owever , i nr e fe rence t ob a r ley y i e lds , b edou in e xpec ta t ions f a l l s ho r t o ft he 8 0 .5 k g /du mean y i e ld d e r ived f r om T i tow 's M ed ieva ld a ta . F rom a ne xam ina t ion o ff u r the ry i e ld d a ta f r om t he N ea r E as t , i twou ld a ppea rt ha t , i ng ene ra l , mean Med ieva l b a r ley y i e lds i nt he W inches te r B i shop r ic w e re ,f o rw ha teve rr easons , s l i gh t ly h i ghe rt han t hose n o rma l ly e xpec ted o na ve rage l a nd i nt he N ea rE as t . F o re xamp le , T ab le 1 6s umma r izes t he a ve rage w hea t a nd b a r ley y i e lds e xpec ted o n A rab f a rms i n Manda te P a les t ine a cco rd ing t o a1 930 g ove rnmen t r epor t ( i n S impson 1 930 :185 , A ppend ix 2 4 ) . S im i la r y i e ld f i gures f o r A rab f a rms i nt he l a te 1 920s we re g i ven i n ar epo r t c omp i led b y t he Z ion is t A g r icu l tu ra l E xpe r imen ta l S ta t ion a tT e l A v iv , w he re a ve rage w hea t p roduc t iv i ty was p l aced a t 5 0 k g /du , a nd a ve rage b a r ley y i e lds a t 6 0 k g /du ( i n S i mpson 1 930 : 177 ) . S uch v a lues a l so a cco rd w e l l w i th t he a ve rage y i e lds o f4 8 k g /du f o rw hea t a nd 6 3 k g /du f o rb a r ley d ec la red i n1 04 A rab v i l lages a sp resen ted i nT ab le 1 6 . I nf u r the rr e fe rences t o b a r ley c u l t iva t ion b y b edou in i nI s rae l a nd J o rdan , Web ley ( 1972 :173 , 1 75 ) n o ted a ve rage y i e lds o f4 4 a nd 7 5 k g /du n ea rT e l l G eze r d epend ing u pon t he s pec i f ic s o i ls b e ing c u l t iva ted , wh i le H e lms ( 1981:187 ) n o ted a ve rage y i e lds o f 4 0 -60 k g /du o n m ud f la ts i n n or theas te rn J o rdan . T hese whea t a nd b a r ley y i e lds c l ose ly a pp rox ima te t hose w h ich w ere n o rma l ly e xpec ted o n ma r l l an ds i nC o rnwa l l a nd D evon i nA .D . 1 668 a s e s tab l ished b y e nqu i r ies o ft he R oya l S oc ie ty ( Lenna rd 1 932 :39 , T ab le I , 4 0 , T ab le I V ) . T hese l a t te r e xpec ta t ions a re s umma r ized i nT ab le 1 7 , a ssum ing a 6 0 l b( 27 .2 k g ) b ushe l f o r whea t a nd a 5 6 l b( 25 .4 k g ) b ushe l f o r b a r ley ( S tephens 1 866 :435 , 4 46 ) . I ns ubsequen tc a lcu la t ions o fp o ten t ia l p reh is to r ic r e tu rn r a tes , mean y i e lds o f5 0 k g /du f o r w hea t a nd 6 0 k g /du f o r b a r ley w i l l b e a ssumed a s a pp rop r ia te p roduc t iv i ty d i v is ions b e tween " op t ima l " a nd " ma rg ina l " l a nds u nde r w hea t a nd b a r ley c u l t iva t ion . I n a cco rdance w i th t he d i scuss ion o f s o i ls a nd h yd ro logy p resen ted i nC hap te r 4 , i ti se xpec ted t ha t t hese o r l owe r mean y i e lds w ou ld c ha rac te r ize t he ma jo r i ty o fa la va i lab le a rab le l a nds . F o rt he mo re c i rcumsc r ibed " op t ima l " e nv i ronmen ts w i th r i che rs o i ls a nd n a tu ra l o re as i ly o b ta ined s ou rces o fi r iga t ion , h i gher m ean y i e lds a re c er ta in ly e xpec ted . F o r e xamp le , T ab le 1 8 s umma r izes H i l lman 's ( 1 973 :226 -227 , a nd A ppend ix 1 )o bse rva t ions o n a ve rage whea t a nd b a r ley y i e lds u nde rt r ad i t iona ld r y fa rm ing a nd i r iga t ion p rac t ices i nt he v i l lage o f Aran , T u rkey , s how ing t he i n c reases a ch ieved w i th i r iga t ion . F o rc ompa r ison , l o n ides ( 1939 :xx i l i ) s ugges ted t ha tt h e a verage y i e ld o fw hea t o n i r iga ted l a nd i nt he J o rdan V a l ley was 1 00 k g /du , w h i le a b edou in i n fo rman t o f Wendo r f a nd S ch i ld ( 1980a :27 1) s ugges ted w hea t y i e lds a s h i gh a s 2 74 k g /du u nde r r unof c u l t iva t ion d ur ing g ood y ea rs i nt he Wad i R amd i in i nE gyp t . I tw i l l t he re fo re b e a ssumed t ha t mean w hea t a nd b a r ley y i e lds i nt he mo re c i rcumsc r ibed e nv ironmen ts p ossess ing r i ch s o i ls w i th h i gh r a in fa l l o rn a tu ra l o re as i ly o b ta ined i r iga t ion c ou ld h ave r anged a tl e as ta s h i gh a s2 00 k g /du i nt he p reh is to r ic p as t .

11

T ab le 1 5 Whea ta nd B a r ley Y ie ld E xpec ta t ions o fB edou in F a rme rs i nt he N egev

I NFORMANT Y IELD E VALUAT ION

B ARLEY ( kg /du ) Y ie ld S ow ing r a te

WHEAT ( kg /du ) Y ie ld S ow ing R a te

E xcep t iona l G ood S a t is fac to ry P oo r

1 00 5 0 -70 3 0 -50 0 -20

A l l 5 -7

2 00 7 0 -80 4 0 -60 0 -20

A l l 6 -7

a dap ted f r om Maye rson ( 1960 : 18 )

T ab le 1 6 A ve rage Whea t a nd B a r ley Y ie lds ( k g /du )o nA rab F a rms i nP a les t ine A cco rd ing t o aB r i t ish C omm i t tee R epo r td a ted J u ly 1 3 , 1 930

CROP

A S D ECLARED I N 1 04 V ILLAGES

" SELECTED E V IDENCE "

" OFF IC IAL E ST IMATE "

Whea t B a r ley

4 8 6 3

5 7 5 4

6 7 7 4

a dap ted f r om S impson ( 1930 : 185 , A ppend ix 2 4 )

T ab le 1 7 E xpec ted Whea t a nd B a r ley Y ie lds ( kg /du ) o n Ma r lL an ds i nC o rnwa l l a nd D evon i nA .D . 1 668

CHARACTER OF S EASON

WHEA 1 S ow ing R a te

Good Mean B ad

Y ie ld

B ARLEY S ow ing R a te Y ie ld

8 1 5 4 3 4

6

9

9 4 5 6 4 4

c a lcu la ted f r om L enna rd ( 1 932 :39 , T ab le I ,4 0 ,T ab le I V )

T ab le 1 8 E xpec ted Whea t a nd B a r ley Y ie lds ( k g /du ) U nde rT rad i t iona l A g r icu l tu ra l P rac t ices a te a r l , T u rkey

GRO W ING COND IT IONS D ry -cu l t iva t ion I r iga t ion

WHEAT S ow ing R a te 9 .2 1 2 .0

Y ie ld

B ARLEY S ow ing R a te

6 3 10

s ou rce o fd a ta : H i l lman ( 1973 :226 -227 , a nd A ppend ix I ) 12

5 1 3

Y ie ld 4 1 15

F i e ld L abo rC os ts o fC u l t iva t ing Whea ta nd B a r ley U nde rC ond i t ions o fH and T i l lage a nd L i th ic T echno logy I n t roduc t ion A s p rev ious ly n o ted i nC hap te r 3 ,t he p r inc ip le l a bo rc os ts o fc u l t iva t ing c erea ls s uch a s w hea t a nd b a r ley i n c lude t h e f i e ld t a sks o fs o i l p repa ra t ion , s ow ing , a nd h a rves t ing , t he i n i t ia l p rocess ing t asks o ft h resh ing a nd w innow ing t he g ra in , a nd f i na l p rocess ing t asks i n vo lv ing m i l l ing b y g r ind ing o r p ound ing . F o r p urposes o f e va lua t ing t empo ra l s h i f ts i nt he r e la t ive e ne rge t ics o fc e rea l c u l t iva t ion d ue t ot echno log ica l c hanges , l a bo rc os ts w i l l b e g ene ra ted f o r s t ra teg ies i n vo lv ing h and t i lage u nde r e a r ly a nd a dvanced l i t h ic t echno log ies , a nd a rd t i lage w i th a n ima lt r ac t ion a nd e a r ly me ta lt e chno logy . I nt he f o l low ing s ec t ions , q uan t i ta t ive l a bo rd a ta o nf i e ld t asks i n vo lv ing h and t i lage a nd l i t h ic t e chno logy w i l l b ee xam ined .

H and T i l lage E a r ly c e rea l c u l t iva t ion p resumab ly i n vo lved t he u se o f h and t i lage i mp lemen ts s uch a s d i gg ing s t icks , h oes , s pades o r f oo t -p loughs . U n for tuna te ly , w h i le s eve ra l s cho la rs h ave w i tnessed a nd p ho tog raphed t i lage u s ing d i gg ing s t icks , n o q uan t i ta t ive d a ta o n s eedbed p r epa ra t ion u s ing t hese i mp lemen ts h ave b een f ound . T ab le 1 9 , h oweve r , s umma r izes t he r e la t ive e f f ic ienc ies o fd i f fe ren t h and t i lage t echn iques a sd e r ived f r om v a r ious s ou rces . T he b es t q uan t i ta t ive d a ta o n h and t i lage a ppea r t o b e t hose p resen ted b y H aswe l l ( 1953 ) , who r eco rded t he l a bo r e xpended p e r h u t c ompound i nG en ie r i V i l lage i nt he Gamb ia u s ing mode rn l i gh t , l o ng -hand led , f l a t -b laded d raw h oes t oc u l t iva te p l o ts f o rt r ansp lan t ing r i ce a l ong r i ve rf l a ts a f te rf l ood r e cess ion . F i gu re 1 4d i ag rams t he v a r ia t ions i nm ean t i lage l a bo rp e r u n i t a rea wh ich s he r eco rded f o r 1 9 h u t c ompounds ( Haswe l l 1 953 :Append ix 2 ,A ppend ix 6 ) . A s s hou ld b e e xpec ted , d i fe ren t h u t c ompoun ds e xpended w ide ly d i ve rgen t amoun ts o fl a bo r i nt i lage . T he g rea tes t v a r ia t ion , h oweve r , i so bse rved f o r C ompound R , c omposed o f e xs l aves who h ad r e cen t ly s e t t led i nt he v i l lage . U n fo r tuna te ly , i ti sn o t p oss ib le t o d e te rm ine w he the rt he a ppa ren t ly e xcess ive l a bo re xpen ded b yt hese p eop le was a r e su l t o fs uch f a c to rs a s b reak ing n ew l a nd f o r c u l t iva t ion , i n expe r ience w i th t i lage t echn iques , o r t he e f fec ts o f d eb i l i ta t ing d i sease (s ) o n l a bo ro u tpu t . I ng ene ra l , am ed ian l a bo re xpend i ture o fa pp rox ima te ly 2 0 h r s /du s eems a pp rop r ia te f o rh oe t i lage u nde rt hese c ond i t ions , a nd i ti st h is v a lue w h ich i s p r esen ted i nT ab le 1 9 . P resumab ly , t i lage f o rs eedbed p repa ra t ion i ng row ing w hea t o rb a r ley w ou ld r equ i re mo re l a bo rt han t ha t e xpended i np repa ra t ion f o rt r ansp lan t ing r i ce . I nr e fe rence t oh and t i lage u s ing modern me ta l s pades , H op fen ( 1969 :43 ) n o ted t ha to ne p e rson c ou ld t i l a pp rox ima te ly 2 0 m2 o f med ium a rab le s o i l p e r h ou rt o ad ep th o fa round 2 2 cm . A ssum ing t ha t t i lage t o ad ep th o f 1 cm wou ld r equ i re h a l f a s much t ime , ar ange o f e f ic iency f r om 2 5 -50 h r s /du h as b een c a lcu la ted . T he e f f ic iency g i ven f o rH op it i lage u s ing aw ooden s hove l i sd e r ived f r om d a ta p resen ted b y F o rde ( 1931:389 -390 ) . A cco rd ing t o h i s o bserva t ions , t he H op i w ou ld t i l t he i r c o rn f i e lds w i th b l aded t oo ls o fw ood r e semb l ing s pades p r io rt op l an t ing c o rn i no rde rt od im in ish wa ter e vapo ra t ion f r om t he s o i l . F ou ro rf i ve d ays w e re n o rma l ly e xpended f o rw eed ing a nd c l ea r ing a f i e ld o f a round o ne a c re i ns i ze , a nd t w ice t ha t t ime f o r " hoe ing " w i th w ooden s pades . A ssum ing t ha t 8t o 1 0 d ays w e re t he re fo re s pen t i nt i l ing o ne a c re a nd t ha t a n 8t o 1 0 h our w o rk d ay was i n vo lved , a pp rox ima te ly 1 6 t o 2 5 h r s /du w e re e xpended . H oweve r , s i nce H op i t i lage was i n tended t o b reak t he s o i l i no rde r t o i n h ib i t e vapo ra t ion r a the r t han t o a c tua l ly p repa re as eedbed , a dd i t iona l l a bor w ou ld p resumab ly b e n eeded t op roduce t he f i l th r equ i red f o rw hea to rb a r ley c u l t iva t ion u s ing s uch i mp lemen ts . T he e f f ic iency g i ven f o rt h ef oo t -p lough ( casch rom ) i sd e r ived f r om C u rwen 's o bse rva t ions o n t he u se o f t h is i n s trumen t o n t he i s land o f S kye . T h rough t he u se a c aschrom , " a man r e ckons t o' p lough 'a bou t at e n th o f aS co ts a c re i n ad ay -a bou tt w ice a s much a s h ew ou ld b e a b le t o d o w i th a n o rd ina ry s pade o r f o rk " ( Curwen 1 927 :269 ) . A ssum ing a n 8t o 1 0 h our w o rk ing d ay , a pp rox ima te ly 1 7 -21 h rs /du wou ld b e e xpended i nt i lage w i th a f oo t -p lough ( e xc lud ing t ime s pen t i nb reak ing c l ods ) , a nd 3 4 -42 h r s /du w i th s pades o rf o rks .

13

T ab le 1 9 R e la t ive E f f ic iency o fH and T i l lage U s ing V a r ious I mp lemen ts

CULT IVAT ION TOOL

HRS /DU

L ong -hand led me ta l h oe Me ta l s pade H op i wooden s hove l F oo t -p lough ( casch rom ) S pade o rf o rk

2 0 2 5 -50 1 6 -25 1 7 -21 3 4 -42

SOURCE H aswe l l 1 953 H op fen 1 969 :43 F o rde 1 931:389 -390 C u rwen 1 927 :269

r ecent settlers

: 50-: 1 :

Mean S td. dev. Med ian M in. Max . Range .0

24.7 14.3 19.9 13.5 79.0 65.5

r eligious l eader 10

v illage smiths

30 -

= = = = = =

o

5

10

: v illage head t raders

A I BIC

[D IEIFIGINI

II KILI MINI OIPIOIRISIT

1

HUT CO MPOUND F i gu re 1 4 . D ev ia t ions i nh oe ing l a bo r p e r h u t c ompound f r om t he c umu la t ive mean f o r a l h u t c ompounds i n Gen ie r i V i l lage , Gamb ia , i n1 949 . C a lcu la t ions f o rt h is f i gure we re b ased u pon d a ta p resen ted b y H aswe l l ( 1953 :Append ix 2 ,A ppend ix 6 ) .

14

I ns ubsequen t e ne rge t ic c a lcu la t ions f o re a r ly w hea t a nd b a r ley c u l t iva t ion , al a bo rc os t o f 3 5 h rs /du w i l l b e a ssumed a s a n o p t im is t ic a pp rox ima t ion f o rh and t i lage u s ing d i gg ing s t icks o r p r im i t ive h oes w i th s t one b l ades . I tw i l l a l so b e a ssumed t ha t i nl a te r p e r iods , r e f inemen ts i n e a r ly t i lage i mp lemen ts c ou ld h ave r educed h and t i lage l abo r t o a pp rox ima te ly 3 0 h rs /du . P o ten t ia l v a r ia t ions i nt he d eg ree o ff i l th r equ i red f o r whea t a s o pposed t ob a r ley c u l t iva t ion w i l l n o t b e a ccoun ted f o ru nde rc ond i t ions o fh and t i lage .

S ow ing L abo r U nde rH and T i l lage N o d i rec t r e fe rences t ot he amoun t o f l a bo r e xpended i ns ow ing a nd h a r row ing s eed b y h and i ne a r ly a g r icu l tu ra l s t ra teg ies h as b een f ound . H oweve r , t he s ow ing o f c e rea l s eed b y h and r ema ined q u i te c ommon i n t he t he U .S . u n t i l we l l i n to t he n i ne teen th c en tury A .D . A cco rd ing t o d a ta p resen ted i nt he T h i r teen th A nnua l R epor t o f t he C omm iss ioner o f L abor ( U .S . D epa r tmen t o f L abo r 1 899 :433 , U n i t 3 ,4 7 1 U n i t 2 6 , 4 73 , U n i t 2 7 ) , t he s ow ing o f whea t a nd b a r ley s eed b y h and r equ i red 1 . 25 1 .42 h rs /ac re i n1 829 -30 , o r a pp rox ima te ly 0 .31-0 .35 h r s /du . T he re s eems l i t le r eason t o s uspec tt ha tt he re h ave b een ma jo rt empo ra l v a r ia t ions i n t h e e f f ic iency o fs ow ing w hea t a nd b a r ley s eed b y h and , a nd t hese l abo rc os ts w i l l b e a ccep ted a s a pp rop r ia te f o re a r ly c e rea l s t ra teg ies . H oweve r , t he s ubsequen t h a r row ing o fh and -b roadcas t s eed i ne a r ly Ame r ican A g r icu l ture w as a ch ieved u s ing b rush h a r rows d rawn b y h o rses , a nd r equ i red o n ly 2 .5 -2 .8 h rs /ac re , o r a pp rox ima te ly 0 .62 -0 .7 h rs /du . U n fo r tuna te ly , t he me thods b yw h ich whea t o rb a r ley s eed we re h a r rowed u nde r c ond i t ions o f h and c u l t iva t ion h ave n o t b een p rev ious ly a dd ressed . I nt he i r w i ld s t a te , t he s p ike le t i nw hea ts a nd t he t r ip le t i nb a r ley s e rve a s s pec ia l ized a r row -shaped s eed d i ssem ina t ion a nd p l an t ing u n i ts ( Zoha ry 1 969 :58 ) . T he s hape , s ha rpness o ft he i r b ase ( t he r ach is s egmen t ) , a nd t he h a i r iness a nd a wns o fs p ike le ts a nd t r ip le ts c umu la t ive ly r esu l t i n e f fec t ive a r row l ike d ev ices f o r t he n a tu ra l i n ser t ion o f s eed i n to t he s o i l , w i th o n ly t he a wns p ro t rud ing f r om t he g round a f te rf r u i t b u r ia l . T h reshed s eed w ou ld o bv ious ly l a ck s uch d ev ices . F u r the r , i nt empe ra te w e t c l ima tes , t he o p t ima l d ep th f o r s eed ing i ss a id t o b e a round 5 cm b e low t he s ur face , a nd a pp rox ima te ly 1 0 cm i n mo re a r id z ones ( S teensbe rg 1 976 :272 ) . S ome m eans o fh a r row ing s own s eed mus t h ave e x is ted i ne a r ly c e rea lc u l t iva t ion . P resu mab ly , h uman t r ac t ion c ou ld h ave b een u sed i nd raw ing b rush o r b oa rd h a r rows a c ross s own l a nd . A l te rna t ive ly , s own f i e lds c ou ld h ave b een h a r rowed u s ing wooden r akes . I n R oman a gr icu l tu re , Med ic c l ove r ( l uce rne , a l fa l fa ) was h and s own a nd h a r rowed w i th sma l l w ooden r akes ( Co lume l la 1 . 10 .27 ) , at a sk r equ i r ing t wo l a bo re rs t he l e ng th o f awo rk ing d ay p e r i u ge rum o fl and ( 1 .12 .6 ) . A ssum ing a 1 0 h our a g r icu l tura l w o rk d ay , 2 0h rs / iugerum o r r ough ly 7 . 9 h r s /du was r equ i red . I nt he a bsence o ff u r the rq uan t i ta t ive d a ta , at o ta l l a bo rc os to f 8h r s /du f o rs ow ing a nd h a r row ing w i l l b e a ssumed i ns ubsequen t c a lcu la t ions o ft he e ne rge t ics o fe a r ly w hea t a nd b a r ley c u l t iva t ion u nde rc ond i t ions o fh and t i lage . T he r e l iab i l i ty o fC o lume l la 's l a bo r d a ta w i l l b e a dd ressed b e low i nr e fe rence t oa rd t i lage . S ince i tw i l l b e a ssumed t ha t s ubsequen t t asks s uch a s weed ing o r o the r f o rms o ff i e ld t end ing w e re u ncommon i ne a r ly whea t a nd b a r ley c u l t iva t ion , t he f i na l f i e ld l abo rc os t o fe a r ly c e rea l s t ra teg ies w ou ld c ons is to ft he t ime s pen t i nh a rves t ing t he c rop .

R eap ing L abor i nE ar ly C e rea lS t ra teg ies L abo r d a ta o n t he r eap ing e f ic iency o fv a r ious s t one s i ck les a re g i ven i nt he r ep l ica t ion s t ud ies o f S teensbe rg ( 1943 ) f o r h a rves t ing p l o ts s own w i th t wo rowed b a r ley a nd a sma l l p e rcen tage o f o a ts , a nd t he s t ud ies o f K o robkova ( 1981) f o r h a rves t ing p l o ts o f whea t . T hese s t ud ies p r ima r i ly i n vo lved r eap ing e xpe r imen ts w i th r ep l ica te s i ck les , u s ing b o th a nc ien t a nd mode rn s t one b lade i n se r ts . T ab le 2 0 s umma r izes t he i re f f ic iency d a ta o nv a r ious f o rms o fe a r ly s t one s i ck les , a nd i n c ludes a dd i t iona l e f f ic iency d a ta f o r r ep l icas o f e a r ly me ta l s i ck les a nd e thnog raph ic a nd h i s to r ic d a ta o n t he u se o f mode rn i r on s i ck les . F o r t he e a r l ies t c e rea l s t ra teg ies , ar eap ing l a bo ro fa pp rox ima te ly 2 8 -33 h r s /du s eems a pp rop r ia te . B y l a ter p e r iods , r e f inemen ts i ns t one s i ck les h ad s eem ing ly r educed h a rves t ing l a bo r t o a pp rox ima te ly 1 6 -19 h rs /du . A ppa ren t ly , e a r ly f o rms o f me ta l s i ck les w e re n o t n ecessa r i ly mo re e f f ic ien t t han d eve loped f o rms o fs i ck les w i th s t one i n se r ts .

15

T ab le 2 0 T he R e la t ive R eap ing E f ic ienc ies o fP reh is to r ic t h rough Mode rn H and S ick les

GENERAL FORM

U nha f ted F l akes

DESCR IPT ION OF S ICKLE

HRS /DU

SOURCE

4 1 .6 -33 .3

K o robkova 1 98 1 :340 S teensbe rg 1 943 :23

S t ra igh tf l i n tw /s econda r y f l ak ing

3 1.6 2 4 .6

E a r ly D je i tun s i ck le w / 2f l i n ts E a r ly D je i tun s i ck le w / 3f l i n ts S ick le w / 3o bs id ian i n se r ts S i ck le w / 2f l i n ts w /2 nd ry f l ak ing S ick le w / 5o bs id ian i n ser ts

4 1.6 -33 .3 3 3 .3 3 3 .3 2 7 .8 2 7 .8

Obs id ian p i ece w /o s econda ry f l ak ing C u rved f l i n tw /s econda ry f l ak ing

11

S ten i ld s i ck le w / a nc ien tf l ake S ten i ld s i ck le w / mode rn f l ake F i rs tY a langach -depe /S ten i ld s i ck le S econd Y a langach -depe /S ten i ld s i ck le

2

2 6 .7 3 6 .5 2 3 .8 2 3 .8

C rescen t -shaped , 1f l i n t s i ck le C rescen t -shaped , 1f l i n t s i ck le A s a bove , w / s t ra igh t t oo thed e dge

2 0 .0 2 5 .6 2 7 .0

L a te D je i tun , 2i n se r ts , d en t icu la te L a te D je i tun , 3i n ser ts , d en t icu la te

1 8 .5 1 8 .5

E a r ly T r ipo lye w / 4f l i n t i n ser ts E a r ly T r ipo lye w / 5f l i n t i n ser ts S homu tepe w / o bs id ian i n se r ts

K o robkova 1 98 1 :340

S teensbe rg 1 943 :23 I t

K o robkova 1 98 1:340

S teensbe rg 1 943 :23

K o robkova 1 98 1:340

1 8 .5 1 8 .5 1 6 .7 -18 .5

K o robkova 1 98 1:340

C urved h o rn s i ck le w /f l i n t i n se r ts A s a bove b u tw i th l o nge rb l ade L a te T r ipo lye w /s aw too th f l ak ing

1 6 .7 -18 .5 1 6 .7 -18 .5 1 5 .2

K o robkova 1 981 :340

C rescen t r ic b ronze , s e r ra ted e dge B ronze k nob s i ck le , smoo th e dge H a l f -moon s i ck le , c o ld fo rged c oppe r

1 9. 3 -22 .8 2 0 .0 -23 .0 1 8 .5

S teensbe rg 1 943 :23

1 0 .7 8 .7 8 .8 8 .0 -16 .0 8 .0 2 .7 -4 .0 4 . 1-4 .9

S teensbe rg 1 943 :23

)

I r on , I r on , I r on , I r on , I r on , I r on , I r on ,

smoo th -edge0 S lovak ian s i ck le s e r ra ted Ga l ic ian s i ck le s e r ra ted e dge smoo th -edged , sma l l s i ze smoo th -edged , med ium s i ze smoo th -edged , l a rge s i ze 1 829 1830 s i ck les , b a r ley -whea t

I r on , smoo th -edged , t h in w hea t s t and

16

1 .6

K o robkova 1 98 1:340

K o robkova 1 98 1:340 A v i tsu r 1 966 : iv A v i tsu r 1 966 :v U .S . D ep t . o f L abo r 1 899 L erche 1 968 :37

H oweve r , t h ree p rob lems e x is t i nt he u se o ft h is c omp i led d a ta f o r e ne rge t ic c a lcu la t ions . I n i t ia l ly ,t he d a ta p resen ted i nT ab le 2 0 p r ima r i ly r e la te t or eap ing l a bo ro n ly , w h i le a dd i t iona l t i me w ou ld b e r equ i red t o g a the r a nd b i nd t he r eaped g ra in . T o p a r t ia l ly a ccoun t f o rt he a dd i t iona l l a bo r r equ i red i nt hese t asks , q uan t i ta t ive d a ta p resen ted b y T ay lor ( 1844 ) w i l l b e emp loyed . T ay lo r s pec i f ica l ly c onduc ted r eap ing e xpe r imen ts w i th v a r ious h a rves t ing i mp lemen ts a nd t eams o fh a rves te rs . I nr eap ing w i th s cy thes , t wo mowe rs r eaped a nd f i ve a t tendan ts g a the red a nd b ound t he c u t g ra in . I nr eap ing w i th s i ck les , s i x r eape rs w e re emp loyed w i th o ne a t tendan t g a the r ing a nd b i nd ing t he g ra in . H is d a ta , p resen ted i nS co tch u n i ts o f a rea d ur ing a n e xp l ic i t 1 0 h ou r wo rk d ay , a re s umma r ized i nT ab le 2 1 . When t he t o ta lt ime s pen t i ns i ck le r eap ing a l one i ss ub t rac ted f r om t he t o ta l t i me s pen t i nr eap ing , g a the r ing a nd b i nd ing , i ti sa ppa ren t t ha t t he l a t te rt a sks r equ i red a pp rox ima te ly o ne a dd i t iona l h ou rp e rd unum o fc rop . T ay lo r 's d a ta a l so i n d ica te t ha t w hea tg ene ra l ly r equ i res mo re r eap ing l a bo rt han b a r ley , a r e la t ionsh ip a l so a ppa ren t i nt he r eap ing d a ta g i ven b yt he U .S . D epar tmen t o fL abo r ( 1899 ) a s p rev ious ly p resen ted i nT ab le 2 0 . A s w i l l b e n o ted b e low , t h is s ame r e la t ionsh ip a ppea rs i n C o lume l la 's d a ta o n R oman h a rves t ing . T o p a r t ia l ly a ccoun tf o rt h is d i f fe rence ,t he l a bo rc os ts o f r eap ing i ne a r ly c e rea l c u l t iva t ion w i l l b e s e t a t3 3 h r s /du f o r whea t a nd 2 8 h r s /du f o rb a r ley . F o r l a te rc e rea l s t ra teg ies u nde ra dvanced l i t h ic t echno logy , r eap ing l a bo rw i l l b e s e t a t1 9 h rs /du f o r w hea t a nd 1 6 h rs /du f o r b a r ley . A dd ing a n e x t ra h our t o a ccoun t f o r g a the r ing a nd b i nd ing a c t iv i t ies r a ises t hese e s t ima tes t o3 4 h rs /du f o r whea t a nd 2 9 h r s /du f o rb a r ley u nde re a r ly l i t h ic t e chno logy , a nd 2 0 h rs /du f o r whea t a nd 1 7 h rs /du f o r b a r ley u nde r l a te r l i t h ic t echno logy . T hese l a t te rf i gu res s hou ld a l so a pp ly t oe a r ly me ta l s i ck les . F i na l ly , t he d ens i ty o ft he c e rea l c rop b e ing r eaped i se xpec ted t o a f fec t t he amoun t o f l a bo r r equ i red t or eap i t .T h is f ac to r e xp l ic i t ly a ccoun ts f o rt he l ow l a bo rc os t o f1 .6 h r s /du g i ven b y L e rche ( 1968 :37 ) f o r r eap ing whea t i n I r an , s i nce a v e ry t h in s tand was b e ing h a rves ted ( e .g . , L e rche 1 968 :33 -34 , F i gu re 1 ) . D ense r s t ands u ndoub tab ly r equ i re mo re l a bo rt or eap , a s r e f lec ted i nt he o the r v a lues g i ven f o r h a rves t ing c e rea ls w i th i r on s i ck les . U n for tuna te ly , t he e f fec ts o fd ens i ty o fc rop o n r eap ing l a bo r h ave n o t b een q uan t i f ied i np rev ious s t ud ies . I ns ubsequen t e ne rge t ic c a lcu la t ions f o r e a r ly whea t a nd b a r ley c u l t iva t ion , r eap ing l abo r w i l l n ecessa r i ly b e c a lcu la ted a tt he r a tes e s tab l ished a bove , r ega rd less o fd ens i ty o fc rop . T he p r inc ip le e f fec t o ft h is a ssump t ion w i l l b e t he i n f la t ion o f r eap ing l a bo rc os ts f o r t h in c rops a nd t he d ep ress ion o f r eap ing l a bo r f o r d ense c rops . Wh i le t h is w i l l t end t o f l a t ten t he c urve f o r r e tu rn r a tes i n e a r ly c e rea l s t ra teg ies , i tw ou ld n o t a f fec t r e la t ive e ne rge t ic c ompa r isons b e tween whea t a nd b a r ley s ubs is tence o p t ions , n o r w ou ld i ta f fec t c ompa r isons w i th l a te r c e rea l s t ra teg ies , s i nce t he s ame a ssump t ion o n r eap ing l a bo rw i l l n ecessa r i ly b e emp loyed . I t a l so s eems u n l ike ly t ha t t h is a ssump t ion wou ld a fec t r e la t ive e ne rge t ic.compa r isons b e tween c e rea l a nd p as to ra l r e tu rn r a tes e xcep t a t t he e x t reme l owe r a nd u ppe r e nds o f c erea l p roduc t iv i ty . I ns umma r y , f i e ld l a bo rc os ts f o rw hea t a nd b a r ley c u l t iva t ion u nde rc ond i t ions o fe a r ly l i t h ic t e chno logy w i l l b e c a lcu la ted o nt he b as is o f3 5 h rs /du f o rt i lage , 8h rs /du f o rs ow ing , a nd at o ta l h a r ves t ing l a bo ro f3 4 h r s /du f o r whea t a nd 2 9 h rs /du f o rb a r ley . C umu la t ive ly , t h is r ep resen ts a t o ta l f i e ld l a bo ro f7 7 h rs /du f o r whea t a nd 7 2 h r s /du f o r b a r ley . A s l i t h ic t oo ls a nd a g r icu l tu ra l

T ab le 2 1 C ompa ra t ive E f f ic ienc ies ( h rs /du ) o fA l te rna t ive R eap ing I mp lemen ts a nd t he A dd i t iona l L abo r R equ i red f o r Ga the r ing , B ind ing a nd R ak ing

I MPLEMENT S cy the Smoo th s i ck le S e r ra ted s i ck le

REAP ING ONLY Whea t B a r ley 1 .5 9 .1 1.8

I I I

1 .0 4 .8 6 .0

REAP ING + Whea t B a r ley 5 .3 1 0 .6 1 3 .7

c a lcu la ted f r om T ay lo r( 1844 :263 ) 17

I I I

3 .5 5 .6 7 .0

D IFFERENCE Whea t B a r ley 3 .8 1 .5 1 .9

I I I

2 .5 0 .8 1 .0

t echn iques we re r e f ined a nd p er fec ted , i tw i l l b e a ssumed t ha t t i lage l abo r d ropped t o 3 0 h rs /du , wh i le t o ta l r eap ing l a bo r d ropped t o 2 0 h rs /du f o r w hea t a nd 1 7 h rs /du f o r b a r ley . S ow ing l abo rs hou ld h ave r ema ined c ons tan t . C umu la t ive ly ,t h is r ep resen ts at o ta l f i e ld l a bo ro f 5 8 h rs /du f o r whea t a nd 5 5 h rs /du f o r b a r ley u nde r c ond i t ions o f h and t i lage a nd a dvanced l i t h ic t echno logy .

F i e ld L abo rC os ts o fC u l t iva t ing Whea t a nd B a r ley U nde rC ond i t ions o fA n ima lT rac t ion T i l lage a nd E a r ly Me ta lT echno logy I n t roduc t ion A s d i scussed i n Chap te r 3 ,b y a tl e as t t he f our th m i l lenn ium B .C . , a n ima l t r ac t ion t i lage i mp lemen ts k nown a s a rds w i th s ymme t r ica l s ha res w e re b e ing u sed f o rc u l t iva t ion i nt h e N ea r E as t a nd E u rope . Work ing a td ep ths o fa pp rox ima te ly 1 0 -30 cm ( Av i tsu r 1 965 :v -v i ; S teensberg 1 964 :128 , 1 976 :27 1; V a r isco 1 982 :166 167 ) , s ymme t r ica l a rds l o osen s o i l e qua l ly o n b o th s i des o f t he f u r row c rea ted . I n a reas whe re d ra inage w as a p rob lem a nd s eedbed r i dges d es i rab le , sma l l f l a t , b en t o rt i l ted s l ide -boa rds w e re a t tached t ob o th s i des o fa n a rd 's s ha re a t t he f i na l p l ough ing t ow iden t he f u r row a nd c rea te " r a f te rs " o r r i dges , ap rac t ice a t tes ted b y a t l e as t t he f i rs t c en tury B .C . ( e .g . , V a r ro 1 . 29 .2 ; V i rg i l Georg . 1 .72 ; s ee A i tken 1 956 :10 1102 ; Mann ing 1 964 :56 ; S teensbe rg 1 976 ) . A st he d ense r , mo is tu re re ta in ing c l ay s o i ls o fE u rope c ame u nde rc u l t iva t ion , l a rge rs l ide b oa rds we re a t tached t oo ne s i de o ft he a rd , p roduc ing a n a symme t r ica l a rd c apab le o fc u t t ing a nd p ar t ia l ly p u lve r iz ing f u r row -s l ices ( bu t n o t f u l ly i n ver t ing t hem ) , ad eve lopmen t i n t ia l ly a t tes ted i nt he f i r s t c en tury A .D . i nt he f oo th i l ls o ft he I t a l ian A lps a nd e as te rn Sw i tze r land ( P l iny X V I I I .48 ) . N ume rous v a r ia t ions o nt he a symme t r ica l a rd w e re emp loyed i nE u rope u n t i lt h e l a te s even teen th o r e a r ly e i gh teen th c en tury , when c urved mou ldboa rds c apab le o ff u l ly i n ver t ing f u r row -s l ices we re d eve loped f r om C h inese p ro to types , r e su l t ing i nt he f i rs t t r ue p l oughs . F i gure 1 5 d iag rams t hese t h ree b as ic s tages i n t he h i s to r ic e vo lu t ion o f mode rn w es te rn p loughs . S ince t he d a ta p rev ious ly s umma r ized i nT ab le 2 0 i n d ica te t ha t e a r ly me ta l s i ck les w e re n o t mo re e f f ic ien tt han a dvanced s i ck les w i th s t one i n se r ts ( a l though t hey p resumab ly h ad al o nge r w o rk ing l i f e ) , t he f o l low ing s ec t ions w i l lf ocus u pon e s tab l ish ing s ymme t r ica l a rd t i lage l a bo rf o r whea t a nd b a r ley c u l t iva t ion i ne a r ly a n ima lt r ac t ion s t ra teg ies .

a . S ymme t r ica la rd

b . A symme t r ica l a rd w i th s l ide -boa rd

c . P lough w i th mou ldboa rd

F igure 1 5 . S imp l i f ied h i s to r ic e vo lu t ion o f t he mou ldboa rd p lough ( a f ter H op fen 1 969 ) . S ymme t r ica l a rds ( a ) we re b e ing u sed i nt he N ea r E as t a nd E urope b y a t l e as t t he f o ur th m i l lenn ium B .C . A symme t r ica l a rds w i th l a rge s l ide -boa rds ( b ) a re i n i t ia l ly a t tes ted i nE u rope i n t he f i r s t c en tu ry A .D . P loughs w i th mo ldboa rds ( c ) w e re s ubsequen t ly d eve loped i nE urope i n t he l a te s even teen th o r e a r ly e i gh teen th c en tury A .D . f r om C h inese p ro to types . 18

R oman F ie ld T ask L abo rU nde rA rd C u l t iva t ion : T he D a ta o fC o lume l la T he R oman a g ronom is t L uc ius J un ius Mode ra tus C o lume l la i nh i s R es R us t ica p rov ided t wo d e ta i led a ccoun ts o ft h e t o ta l d ays o ff i e ld l a bo r e xpended b y f a rme rs i nt he f i r s t c en tury A .D . f o r whea t a nd b a r ley c u l t iva t ion i nI t a ly . B y r e fe rence t ot he n umber o f mod i i( 1 mod ius = 6 .74 k g ) o f w hea t o r b a r ley c ommon ly s own p e r i ugerum ( =2 .518 d u ) o f l and , C o lume l la s umma r ized t o ta lf i e ld l a bo ra sf o l lows ( 1948 :187 ) : A nd n ow t o r e ckon u p t he n umbe ro fd ays ' l a bou r r equ i red t ob r ing t ot he t h resh ing f loor w ha t we h ave c omm i t ted t o t he e ar th , f ou r o r f i ve mod i i o f c ommon w hea t t a ke u pf our d ays ' wo rk o ft he p l oughman , o ne o ft he h a r rowe r , t wo o ft he h oe rf o rt he f i r s t h oe ing a nd o ne f o rt he s econd , o ne o ft he weede r , a nd o ne a nd a h a l f o ft he r eape r -a t o ta l o f t en a nd o ne -ha l f d ays o f l a bour [ 1 .12 .1] . F ur the r : F i ve mod i io f b a r ley r equ i re t h ree d ays ' l a bour o ft he p l oughman , o ne d ay o f h a r row ing , o ne a nd ah a l fo fh oe ing , a nd o ne o fr eap ing -s i x a nd ah a l fd ays i n a l [ 1 .12 .2 ] . A ssum ing a 1 0 h ou ra g r icu l tu ra l wo rk d ay , T ab le 2 2 s umma r izes C o lume l la 's l a bo rd a ta f o r whea t c u l t iva t ion , wh i le T ab le 2 3d oes s of o rb a r ley .

T ab le 2 2 T o ta l L abo rf o r One l uge rum o f Whea t i nR oman A g r icu l tu re A cco rd ing t oC o lume l la ( 1 .12 .1)

TASK P lough ing H a r row ing i s t h oe ing 2 nd h oe ing Weed ing R eap ing TOTAL

HRS / IUGERUM ( =2 .5 18 DU )

HRS /DU

4 0 1 0 2 0 1 0 1 0 1 5

1 5 .9 4 .0 7 .9 4 .0 4 .0 6 .0

1 05

4 1.8

T ab le 2 3 T o ta l L abo rf o r One l uge rum o fB a r ley i nR oman A g r icu l tu re A cco rd ing t oC o lume l la ( 1 .12 .2 )

TASK

HRS / IUGERUM ( =2 .5 18 DU )

HRS /DU

P lough ing H a r row ing H oe ing R eap ing

3 0 1 0 1 5 1 0

1.9 4 .0 6 .0 4 .0

TOTAL

6 5

2 5 .9

19

O n ac ompa ra t ive b as is , l e ss t i lage , h oe ing a nd r eap ing l a bo rw e re a ppa ren t ly e xpended i nc u l t iva t ing b a r ley t han whea t i nR oman a g r icu l ture . F u r the r , b a r ley f i e lds were s eem ing ly n o t w eeded . I n i t ia l ly , v a r ia t ions i nt he r eap ing l a bo r f o r whea t a nd b a r ley h ave a l ready b een d i scussed . F u r the r , t he weed ing o f whea t a nd b a r ley f i e lds i sn o t u n ive rsa l ly p rac t iced i nt he e thnog raph ic p resen t , a nd C o lume l la h imse l f n o ted t ha t weed ing was u ncommon i nc u l t iva t ing c e rea ls i nE gyp t a nd N o r th A f r ica ( 1 .11 .3 ) . F i na l ly , wh i le whea t was a ppa ren t ly s own a f te rt h ree a nd p e rhaps f ou rt i l ings i nR oman a g r icu l tu re , b a r ley was s own a f te ro n ly t wo t i l ings ( Co lume l la 1 . 9 .15 ) . T h is l a t te rp rac t ice r uns c on t ra ry t oa ccep ted o p in ion i nt he n i ne teen th c en tury A .D . , f o r a s n o ted b y S tephens ( 1866 :607 ) : " . . w hea t t h r ives b es t w hen t he s o i l i ss omewha t f i rm , a nd n o tt oo much p u lve r ized -whe reas t he l a nd c anno t b e i nt oo p u lve r ized as t a te f o rb a r ley . " E ven s o , i tw i l l b e a ssumed t ha t C o lume l la 's d a ta a dequa te ly r e f lec ts c ommon t i lage p rac t ices f o r w hea t a nd b a r ley c u l t iva t ion i nR oman I t a ly , a nd p resumab ly t h e p reh is to r ic p as t i ng ene ra l . S ince n o o the r h i s to r ic o re thnog raph ic s ources h ave b een f ound w h ich a re a s d e ta i led i n e nume ra t ing t he t o ta l f i e ld l a bo rs o fa rd c u l t iva t ion , C o lume l la 's d a ta s eem p rom is ing f o ru se i n s ubsequen t c a lcu la t ions o ft he e ne rge t ics o fw hea t a nd b a r ley c u l t iva t ion u nde r c ond i t ions o f e a r ly a n ima lt r ac t ion t i lage . T he f o l low ing s ec t ion d e ta i ls s uppo r t f o rt he a ccu racy o fh i s d a ta o n t i lage l abo r u s ing s ymme t r ica l a rds , a nd b y e x tens ion , t he p robab le a ccu racy o f h i s d a ta o n h a r row ing w i th wooden r akes .

H is to r ica l V ar ia t ions i nt he E f f ic iency o fA n ima l T rac t ion T i l lage S eve ra l f ac to rs may b e e xpec ted t oa fec tt he e f f ic iency o fa l f o rms o fa nc ien t a nd modem a n ima l t r ac t ion t i lage . I n i t ia l ly , t he d ens i t ies o fv a r ious s o i ls , t he d ep th o ft i lage , a nd t he s i ze o f t he i mp lemen t emp loyed may a l l b e e xpec ted t o a f fec t t he d raugh t r equ i red f o r t i lage . H oweve r , o nce t he r equ i red d raugh t i sa ch ieved , t he s peed a tw h ich t i lage p roceeds w ou ld b e p r ima r i ly d e te rm ined b y t he wo rk ing p ace o f t he a n ima l o r a n ima ls emp loyed . U n less e x t ra a n ima ls we re a va i lab le , t he amoun t o f l a nd wh ich c ou ld b e t i led i n ad ay wou ld d epend u pon t he n umber o fh ou rs f o r wh ich a n a n ima l c ou ld b e wo rked i nd raugh t . T ab le 2 4 s umma r izes t he n o rma l d raugh t a nd wo rk p o ten t ia l o f a n ima ls c ommon ly u sed i nd raugh t a cco rd ing t o d a ta p resen ted b y H op fen ( 1969 :10 11 , T ab le 2 ) . When emp loyed i nt eams , t o ta l d raugh t c apac i ty w ou ld i n c rease , a l though t he a ve rage s peed o f wo rk wou ld n o t . F ur the r , t he u se o ft eams w i th ac umu la t ive d raugh t g rea te rt han t ha t r equ i red f o rt i lage w ou ld d im in ish t he e ne rgy e xpended b y e ach a n ima l , t hereby i n c reas ing t he l e ng th o f t ime d u r ing wh ich t hey c ou ld b e w o rked . F ina l ly , b o th t he w id th o f f u r rows a nd t he n umbe r o f r equ i red t u rn ings o f t he i mp lemen t a nd t eam w ou ld a f fec t t he amoun t o f t ime a c tua l ly s pen t o n t i l ing a u n i t o f l and , w i th l e ss t ime r equ i red w i th b roade r a nd l o nge rf u r rows ( S tephens 1 866 :166 167 ) . I n r e fe rence t o t he mechan ics o ft i lage b y s ymme t r ica l a rds a nd t he t i lage o f p l oughs , e ach b reaks t he s o i l i nq u i te d i s t inc t f a sh ions , a s d i ag rammed i n F i gu re 1 6 . A s ymme t r ica l a rd

T ab le 2 4 N o rma l D raugh ta nd Wo rk P o ten t ia lo fV a r ious A n ima ls

AN IMAL

A ve rage We igh t ( k g )

L i gh t h o rse O x B u f fa lo C ow Mu le D onkey

4 00 -700 5 00 -900 4 00 -900 4 00 -600 3 50 -500 2 00 -300

A pp rox ima te D raugh t ( kg )

A ve rage S peed o f Wo rk ( km /h r )

H o rsepowe r D eve loped

D a i ly Wo rk ( h rs )

6 0 -80 6 0 -80 5 0 -80 5 0 -60 5 0 -60 3 0 -40

3 .6 2 .2 -3 .1 2 .9 -3 .2 2 .5 3 .2 -3 .6 2 .5

1 .00 0 .75 0 .75 0 .45 0 .70 0 .35

8 + 6 -8 — 2 -3 8 + —

a dap ted f r om H op fen ( 1969 :10 11 , T ab le 2 )

1 20

s camna

a

B . P lough w i th mou ldboa rd t i lage ( a f te r S tephens 1 866 ) . C urved mou ldboa rds c u ta nd f u l ly i n ve r tf u r row -s l ices , l e av ing n o b a lks b e tween f u r rows u n less i mp rope r ly u sed .

A . S ymme t r ica l a rd t i lage ( adap ted f r om S teensberg 1 957 -58 ) . B a lks ( scamna ) b e tween f u r rows ( a ) m us t b eo b l i te ra ted b y c rossp lough ing o r ( b ) b y g u id ing t he t he a rd i na l terna te ly u p r igh t a nd s l op ing p os i t ions .

F i gu re 1 6 . V a r ia t ions i nt he mechan ics o ft i lage b e tween s ymme t r ica l a rds a nd p l oughs w i th mou ldboa rds .

b reaks s o i l i n af u r row s l igh t ly l a rger t han t he w id th o f i t s s o le . I fc onsecu t ive f u r rows d o n o t s u f f ic ien t ly o ve r lap , u n t i l led r i dges o r b a lks ( t e rmed s ca mna i n L a t in ) a re l e f t . Where a s ubsequen t c rossp lough ing i s p oss ib le , s ca mna c an b e o b l i te ra ted . H oweve r , where c rossp lough ing i sd i f icu l t ( e .g . , n a r row f i e lds o n s l opes o r b e tween r ows o ft r ees i no rcha rds ) , s camna mus t b e e l im ina ted b y c u t t ing c onsecu t ive f u r rows w i th t he a rd g u ided i na l te rna te ly u p r igh t a nd s l op ing p os i t ions ( Co lu me l la 1 .2 .25 ; S teensberg 1 957 -58 ) . B o th o f t hese t echn iques w ou ld e f fec t ive ly d oub le t he n umbe r o f p asses wh ich mus t b e made w i th a s ymme t r ica l a rd i no rde rt oo b ta in c omp le te t i lage when i n t ia l ly b reak ing p rev ious ly f a l low l a nd . B y c on t ras t , a symme t r ica l a rds a nd p l oughs c u t f u r row -s l ices , a nd w ou ld o n ly l e ave s camna i f i mp rope r ly u sed . T he a dd i t iona l l a bo r r equ i red f o rt i l ing f a l low l a nd u s ing s ymme t r ica l a rds i s a ppa ren t i nC o lume l la 's d e ta i led a na lys is o ft he l a bo rs pen t i np l ough ing w i th t hese i mp lemen ts . C o lume l la p rov ided t wo d e ta i led a ccoun ts o fa rd t i lage l a bo rb y s pec i f ic t ask . A f te ra dv is ing t ha t l a nd b e p l oughed a f te r ar a in h ad made c u l t iva t ion l a bo r e as ier , C o lume l la n o ted t he f o l low ing ( 1948 :133 ) : B u t ai ugerum o f s uch l a nd i sp repa red w i th f our d ays ' l a bou r ; f o r i ti sb roken e as i ly i nt wo d ays , g one o ve r as econd t ime i no ne , at h i rd t ime i nt h ree fou r ths o f ad ay , a nd i sf o rmed i n to r i dges a nd s own i no ne fou r th o f ad ay [ 1 .4 .8 ] . A ssum ing a 1 0 h ou r wo rk ing d ay , T ab le 2 5 s umma r izes C o lume l la 's d a ta . I ti st h is a ccoun t ing o f t i lage l a bo r w h ich C o lume l la a ppa ren t ly u sed i nh i s s umma ry o ft o ta l f i e ld l a bo rf o r c u l t iva t ing w hea t a s p rev ious ly p resen ted i nT ab le 2 2 . I na no the rp assage d e ta i l ing t he amoun t o fl a nd w h ich c ou ld b e wo rked i no ne y ea rw i th a y oke o fo xen ( t wo a n ima ls ) , C o lume l la p rov ided as econd a ccoun t o ft he t i lage l a bo r e xpended f o rc e rea l c rops s uch a s w hea t ( 1948 : 191) : S eeds t ha t a re s own a t t he f our th p l ough ing r equ i re , f o r t wen ty f ive i ugera , o ne h und red a nd f i f teen d ays ' l abour o f t he p l oughmen ; f o r s uch a p lo t o f g round , h oweve r h a rd , i s b roken i n f i ty d ays , r e -p loughed i n t wen ty f ive , p l oughed at h i rd t i me a nd t hen s own i nf o r ty d ays [ 1 .12 .8 ] . A ga in a ssum ing a 1 0 h our w ork ing d ay , T ab le 2 6 s umma r izes t h is d a ta . I nt h is s econd a ccoun t , a n a dd i t iona l 6h rs / iuge rum ( 2 .4 h r s /du ) h as b een i n c luded i nt he l a bo rc os ts o ff i na l t i l ing a nd s ow ing . S ince at h i rd p l ough ing wou ld h ave r equ i red 3 h rs /du a nd r i dg ing a nd s ow ing a no the r 1h r /du , af ou r th p l ough ing a t 2 .4 h rs /du p r io r t o r i dg ing a nd s ow ing i sa ppa ren t ly i n d ica ted . 1 21

T ab le 2 5 E f f ic iency o fR oman S ymme t r ica l A rd T i l lage U s ing O ne Y oke o fO xen ( Two A n ima ls ) A cco rd ing t oC o lume l la ( 1 .4 .8 )

TASK

HRS / IUGERUM ( =2 .518 DU )

HRS /DU

B reak ing f a l low 2 nd p l ough ing 3 rd p l ough ing R idg ing a nd s ow ing

2 0 .0 1 0 .0 7 .5 2 .5

7 .9 4 .0 3 .0 1 .0

TOTAL

4 0 .0

1 5 .9

T ab le 2 6 E f f ic iency o fR oman S ymme t r ica l A rd T i l lage U s ing O ne Y oke o fO xen ( Two A n ima ls ) A cco rd ing t oC o lume l la ( 1 .12 .8 )

TASK

HRS /25 I UGERA ( =62 .95 DU )

HRS /DU

5 00 2 50

7 .9 4 .0

4 00

6 .4

1 , 150

1 8 .3

B reak ing f a l low 2 nd p l ough ing 3 rd p l ough ing t h rough s ow ing TOTAL

I ft h is i sc o r rec t , t he l abo r a ssoc ia ted w i th a f our th p l ough ing was 'n o t i n co rpora ted i n to C o lume l la 's s umma ry o fn o rma l f i e ld l a bo rf o rc u l t iva t ing whea t a s p rev ious ly p resen ted i nT ab le 2 2 . Wh i le t h ree t i l ings w i th as ymme t r ica l a rd p r io rt os ow ing w hea tw as c ommon p rac t ice i nt he a nc ien t w o r ld , n i n th a nd t en th c en tu ry A .D . s ou rces f r om t he N ea r E as t a l so n o te a f our th p l ough ing p r io rt o s ow ing w hea t d epend ing u pon t he q ua l i ty o ft he s o i l ( E I -Sama r ra ie 1 972 :59 ; S e r jean t 1 974 :4 1) . F o r c ompa ra t ive p urposes , T ab le 2 7 s umma r izes a dd i t iona l a ccoun ts o f t he l abo r e xpended i na rd t i lage a s g i ven i nv a r ious h i s to r ica l a nd e thnog raph ic s ources . A ga in , a1 0 h ou r w o rk ing d ay h as b een a ssumed f o r a ccoun ts s t a t ing t he amoun t o fl a nd t i led i n a" day . " T he i n i t ia l f i gures p rov ided b y A v i tsu r ( 1965 : iv ) a re s a id t o r e f lec t t he t o ta l r ange o ft he a rd 's " po ten t ia l c apac i ty , " w h i le t he s econd s e t r e f lec t t he a rd 's " usua l " p er fo rmance . I nl i gh t o ft h is a dd i t iona l d a ta , i twou ld s eem t ha t t he l a bo r r eco rded b y C o lume l la f o r i n t ia l ly b reak ing f a l low , amoun t ing t o 7 .9 h r s /du , r e f lec ts t he a dd i t iona l t ime r equ i red t ot i l au n i t o f l a nd w hen c u t t ing c onsecu t ive f u r rows w i th t he a rd i na l te rna te ly u p r igh t a nd s l op ing p os i t ions , a s p rev ious ly d iscussed . H owever , s ubsequen t t i l lage w i th a s ymme t r ica l a rd g ene ra l ly r equ ired a pp rox ima te ly 3 -4 h rs /du . O ve ra l l , t he l a bo r d a ta p resen ted b y C o lume l la f o r a rd t i lage s eems e x t reme ly a ccu ra te f o re ach o ft he t i lage t asks wh ich h e e nume ra ted . Wh i le e ne rge t ic c a lcu la t ions o f c e rea l c u l t iva t ion u s ing t he l a te r a symme t r ica l a rds a nd p l oughs w i l l n o t b e made i nt he p resen t s t udy , ab r ie f e va lua t ion o ft he a ppa ren t i n c reases i n t i lage e f f ic iency a ssoc ia ted w i th t he i r u se i s wa r ran ted f o r c ompa ra t ive p urposes . A l though r e l iab le t i lage e f f ic iency d a ta f o r a symme t r ica l a rds h ave n o t b een l o ca ted , e x tens ive d a ta d o e x is t o n t he t i lage e f f ic iency o fe a r ly p l oughs . F o re xamp le , t he 1 832 e d i t ion o ft he E d inbu rgh 1 22

T ab le 2 7 R epor ted T i l lage E f f ic ienc ies a nd D raugh ts o fS ymme t r ica l A rds

SOURCE

L OCAT ION

DRAUGHT

HRS /UN IT AREA

HRS /DU

P a les t ine P a les t ine P a les t ine

2o xen 2o xen 2o xen

1 0 /3 -3 .75 d u 1 0 / 1-4 d u 1 0 /2 .5 -3 d u

2 .7 -3 .3 2 .5 -10 3 .3 -4 .0

Y emen

1d onkey

0 .5 / 100 m3

5 .0

V a r isco 1 982 :167

I r an

2o xen

1 0 /3 ,700 m3

2 .7

A lber ts 1 963 :350

M ishna Oha lo t1 7A A v i tsu r 1 965 : iv I f

E ncyc lopaed ia c on ta ins t he f o l low ing c ommen ta ry o n t he e f f ic iency o f J ames Sma l l 's e a r ly s w ing p l ough , a n i mp lemen tw i th awooden b ody a nd ac as t i r on mou ldboa rd i n t ia l ly d eve loped i n1 767 ( 1832 :239 -240 ) : T h is i mp lemen t i s. .t he b es t c ons t ruc ted s w ing p l ough i nt he i s land ; a nd b y means o fi taman a nd t wo h o rses w i l l , w i th a l e ase , p l ough a n E ng l ish a c re p e r d ay , e xcep t i np a r t icu la rs easons , when t he s o i lf r om d rough t h as b ecome h a rd a nd o bs t ina te . We h ave o f ten s een aS co tch a c re ( one f i th l a rge rt han E ng l ish measu re ) p l oughed i nt wo j ou rneys , o r n i ne h ours ' wo rk ; b u t t h is was e i the r w hen t he l a nd was s eed f u r rowed , o rw i th as o i lo f al o ose a nd f r i ab le n a tu re . T ab le 2 8 s umma r izes t hese d a ta . F o r c ompa ra t ive p urposes , T ab le 2 9 p resen ts s im i la r e f f ic iency d a ta o n e a r ly wooden p l oughs i nt he U .S . a sc omp i led f r om v a r ious s ou rces b y R og in ( 1931:16 17 ) . T hese i mp lemen ts w e re o rd inary s w ing p l oughs w i th a wooden mou ldboa rd c a rved f r om a w ind ing t r ee a nd s hod w i th i r on p l a te ( Rog in 1 93 1 :5 ) . I nl i gh to ft hese d a ta , i ti s

T ab le 2 8 T i l lage E f f ic iency o fJ ames Sma l l 's Sw ing P lough D rawn b y Two H o rses

P LOUGH ING COND IT IONS

HRS /UN IT AREA

HRS /DU

A ve rage s o i l S eed f u r row ing , l o ose s o i l

1 0 /s ta tu te a c re ( 4 .05 d u ) 9 /Sco tch a c re ( 4 .86 d u )

2 .5 1 .9

s ou rce o fd a ta : E d inburgh E ncyc lopaed ia ( 1832 :239 -240 )

T ab le 2 9 T i l lage E f f ic ienc ies a nd D raugh ts o fE a r ly Ame r ican Wooden P loughs

LOCAT ION

DATE

DRAUGHT

HRS /ACRE ( 4 .05 DU )

HRS /DU

Massachuse t ts N ew E ng land R hode I s land N ew E ng land V i rg in ia

1 815 1 819 1 820 -21 1 793 -99

4o xen 4o xen 2o xen 2h o rses 2h o rses

1 0 .0 1 0 .0 1 0 .0 6 .7 1 0 .0 -13 .3

2 .5 2 .5 2 .5 1 .7 2 .5 -3 .3

c a lcu la ted f r om R og in ( 1931 :16 17 ) 1 23

n o t s urp r is ing t ha t t he s t anda rd o f p l ough ing i nt he m id -n ine teen th c en tury i nt he U .S . was t aken a t 1 0 h rs /ac re ( S tephens 1 866 :167 ) , o r a pp rox ima te ly 2 .5 h rs /du . A t a n a ve rage e f f ic iency o f2 .5 h rs /du , e a r ly p l oughs a ppa ren t ly c u t o n ly 0 . 5 1 .5 h ou rs o f t he amoun t o ft ime i tt ook t ot i l ad unum o fp rev ious ly b roken l a nd w i th as ymme t r ica l a rd . I tw ou ld s eem t ha tw h i le i n nova t ions i na n ima l t r ac t ion t i lage i mp lemen ts d i d r esu l t i ni n c reased e f f ic iency a s measu red b y l abo r h ours p e r u n i t a rea , t he a dvances a ch ieved d o n o t a pp roach t he magn i tude o f e f f ic iency i n c rease a ppa ren t b e tween h and t i lage a nd e a r ly a n ima l t r ac t ion t i lage u s ing s ymme t r ica l a ids . I n s ubsequen t c a lcu la t ions o f t he e ne rge t ics o f e a r ly c e rea l c u l t iva t ion i n vo lv ing s ymme t r ica l a rds a nd a n ima lt r ac t ion , C o lume l la 's p l ough ing a nd h a r row ing d a ta a s s umma r ized i nT ab les 2 1 a nd 2 2 w i l l b e emp loyed . H is d a ta o n h oe ing a nd w eed ing l a bo r , h oweve r , w i l l b e e xc luded . I tw i l l t he re fo re b e a ssumed t ha t whea t r equ i red 1 6 h rs /du a nd b a r ley 1 2 h r s /du i n t i lage a nd s ow ing l a bo r , w i th b o th r equ i r ing a n a dd i t iona l 4h r s /du i nh a r row ing . W i th ar eap ing l a bo ro f2 0 h rs /du f o r whea t a nd 1 7 h r s /du f o rb a r ley u s ing e a r ly me ta l s i ck les , t o ta lf i e ld l a bo ri n e a r ly a n ima lt r ac t ion c ul t iva t ion wou ld b e 4 0 h r s /du f o r whea t a nd 3 3 h rs /du f o rb a r ley .

P rocess ing L abo rC os ts o f Whea t a nd B a r ley I n t roduc t ion I n i t ia l whea t a nd b a r ley p rocess ing l abo r i n vo lves t h resh ing a nd w innow ing t he r eaped g ra in , w h i le f i na l p rocess ing i n vo lves v a r ious f o rms o fg ra in m i l l ing . A s p rev ious ly d i scussed i n C hap te r 3 , t he t echn iques b y wh ich e a r ly c e rea ls we re p rocessed r ema ins u nknown . F i re t h resh ing may h ave b een u sed , a l though n o c onc ise d a ta e x is t o n t he e f f ic iency o f s uch t echn iques . T he b es t t ha t c an b e a ch ieved h e re i st o u se a va i lab le q uan t i ta t ive d a ta o n t he t h resh ing a nd w innow ing o f u n imp roved l a nd races o fw hea t a nd b a r ley b y e a r ly t echn iques , a low ing f o rt he g rea te r l a bo r wh ich was p resumab ly e xpended i nt h resh ing c e rea ls wh ich w e re n o t f r ee th resh ing . T he f o l low ing s ec t ions d e ta i l t he l a bo r c os ts o f i n t ia l p rocess ing t a sks , f o l lowed b y d a ta o n t he f i na l p rocess ing o f g ra ins b y g r ind ing o r p ound ing c e rea ls u s ing p r im i t ive i mp lemen ts .

I n i t ia lP rocess ing : T hresh ing a nd W innow ing Two s e ts o f q uan t i ta t ive d a ta o n t he i n i t ia l p rocess ing o f whea t a nd b a r ley h ave b een l o ca ted . T he f i r s t o ft hese i sd er ived f r om o bser va t ions o n t h resh ing a nd w innow ing w hea t b y t r ad i t iona l me thods i nt he v i l lage o f H asanab lad , I r an , a s o bserved a nd r eco rded b y Wa tson ( 1979 :80 -82 ) . H e re , r eaped g ra in was s p read o n ap repa red e a r then t h resh ing f l oo r , t r ead b y f i ve o xen , w innowed i nt he w ind w i th w ooden f o rks t o s epa ra te t he g ra in f r om t he c ha f f , a nd f i na l ly w innowed u s ing s i eve -baske ts . A cco rd ing t oh e ro bser va t ions ( 1979 :82 ) : T o t h resh o u t 1 00 mann ( one x arva r ) o f whea t , t he f a rme r mus t d r ive t he o xen a round o ve r t he g ra in f o r t we lve t o f i teen h ours , w innow i tw i th aw innow ing f o rk f o rt h ree t of ou rh ours , a nd f i na l ly w innow i tw i th as i eve l ike b aske tf o ro ne a nd ah a l ft ot wo h our s . H e rp rocess ing d a ta f o r 1 00 mann ( c . 3 00 k g ) o fw hea t a re s umma r ized i nT ab le 3 0 , a ssum ing t ha t o n ly o ne l a bo re r was i n vo lved . F ur the rq uan t i ta t ive d a ta o n t he i n i t ia l p rocess ing o fg ra ins a re c on ta ined i nt he T h i r teen th A nnua lR epo r to ft he C omm iss ioner o fL abor ( U .S . D epar tmen t o fL abo r 1 899 : 4 33 , 4 7 1 , 4 73 ) , whe re s epa ra te d a ta we re p resen ted o n whea t a nd b a r ley p rocess ing i n 1 829 -30 . H e re , t h resh ing was a ch ieved u s ing wooden f l a i ls , w h i le w innow ing c ons is ted o ft wo l abo re rs c r ea t ing ab reeze b yf l app ing as hee tt ob l ow a way c haf w h i le at h i rd l a bo re rp ou red t he whea t a nd c ha f f f r om a h e igh t , t he g ra in t hen p ass ing t h rough a c oa rse s i eve o r r i dd le n ea r t he f l oo r ( Rog in 1 931 :154 ) . T ab le 3 1 s umma r izes t he p rocess ing l abo r d a ta f o r a l wo rke rs a s p resen ted f o r w hea t , w h i le T ab le 3 2 d oes s of o rb a r ley , a ssum ing a6 0 l b( 27 .22 k g ) b u f o r whea t a nd a5 6 l b ( 25 .4 k g ) b u f o r b a r ley ( S tephens 1 866 :435 , 4 46 ) .

1 24

T ab le 3 0 E f ic iency o fT h resh ing a nd W innow ing Whea t b y T rad i t iona l Me thods i nI r an

TASK

HRS /300 KG

HRS /KG

T h resh ing ( 5o xen ) F ork W innow ing S ieve W innow ing

1 2 .0 -15 .0 3 .0 -4 .0 1 .5 -2 .0

0 .040 -0 .050 0 .010 -0 .013 0 .005 -0 .007

TOTAL

1 6 .5 -21 .0

0 .055 -0 .070

c a lcu la ted f r om Wa tson ( 1979 :82 )

T ab le 3 1 E f f ic iency o fT h resh ing a nd W innow ing Whea t U s ing F l a i ls a nd S hee ts i nE a r ly Ame r ican A g r icu l tu re

TASK H au l ing S heaves t ob a rn T h resh ing a nd s t ack ing s t raw W innow ing w i th s hee ts Ga the r ing a nd s ack ing --------- TOTAL

HRS /20 BU ( C . 5 44 .32 KG )

HRS /KG

4 .00 1 3 .33 1 0 .00 -12 .0 3 .33 -4 .0

0 .007 0 .025 0 .018 -0 .022 0 .006 -0 .007

3 0 .66 -33 .33

0 .056 -0 .061

- - -

c a lcu la ted f r om U .S . D epa r tmen t o fL abo r( 1899 )

T ab le 3 2 E f f ic iency o fT h resh ing a nd W innow ing B a r ley U s ing F l a i ls a nd S hee ts i nE a r ly Ame r ican A g r icu l tu re

TASK

HRS /30 BU ( C . 7 62 .03 KG )

HRS /KG

H au l ing S heaves t ob a rn T h resh ing a nd s t ack ing s t raw W innow ing w i th s hee ts Ga the r ing a nd s ack ing

4 .00 1 5 .00 1 2 .75 4 .25

0 .005 0 .020 0 .017 0 .006

TOTAL

3 6 .00

0 .048

c a lcu la ted f r om U .S . D epa r tmen t o fL abo r( 1899 )

1 25

O n ac ompa ra t ive b as is , t he i n i t ia l p rocess ing c os ts a s p resen ted i nt hese t empo ra l ly a nd s pa t ia l ly d i ve rse s ources a re q u i te c l ose , as urp r is ing r e su l t c ons ide r ing t he d i f fe rences i n p rocess ing t echn iques a nd t he n umber o f l abo re rs i n vo lved . H oweve r , wh i le a n i n i t ia l p rocess ing c os t o fa pp rox ima te ly 0 .06 h rs /kg a s d e r ived f r om t hese d a ta s eems a pp rop r ia te f o r u se i nc a lcu la t ing t he l a bo r c os ts a ssoc ia ted w i th f r ee th resh ing g ra ins , a lowance m us t b e made f o r e a r ly g ra ins wh ich we re n o t f r ee th resh ing . T o t h is e nd , i tw i l l b e a ssumed t ha t e a r ly c e rea ls r equ i red a t l e as t t w ice t he l abor t o t h resh a s f r ee th resh ing g ra ins . A s a pp l ied t o Wa tson 's d a ta , i n i t ia l p rocess ing c os ts wou ld r i se t o 0 .095 -0 .12 h rs /kg , a nd a s a pp l ied t ot he D epa r tmen t o f L abo r 's d a ta , l a bo r c os ts wou ld r i se t o a pp rox ima te ly 0 .068 -0 .086 h r s /kg . A n i n i t ia l p rocess ing c os t o f0 . 1 h rs /kg w i l l t he re fo re b e a ssumed a s a pp rop r ia te f o r e a r ly c e rea ls w h ich w e re n o t f r ee th resh ing .

F ina lP rocess ing : G r ind ing a nd P ound ing A s p rev ious ly n o ted i nC hap te r3 , mos t o ft he a va i lab le l a bo rd a ta o nt he f i na l p rocess ing o f c e rea ls r e la te t ot he p roduc t ion o ff i ne f l ou r g round f r om w hea t f o rt he b ak ing o fb reads . T ab le 3 3 s umma r izes t hese d a ta a sc omp i led a nd p resen ted i nv a r ious s ou rces . F o rbes ' d a ta r e la te t o t he p roduc t iv i ty o f t wo R oman s l aves a s d er ived f r om a n u nspec i f ied a nc ien t s ou rce , wh i le J asny 's d a ta we re c omp i led f r om v a r ious mode rn s ou rces w o r ld -w ide . C u rwen 's d a ta p resen t t he r esu l ts o f r ep l ica t ion e xpe r imen ts , a s d oes t he i n t ia l e n t ry b y Mo r i tz a nd J ones . T he i r s econd e n t ry was d e r ived f r om a n i n dependen t s ou rce d a t ing t ot he t u rn o ft he c en tu r y . P resumab ly , much o ft he v a r ia t ion a ppa ren t b e tween t hese f i gu res d er ives f r om v a r ia t ions i nt he f i neness o f t he f l our b e ing p roduced . T h is i sp ar t icu la r ly a ppa ren t i nt he e xpe r imen ts c onduc ted b y C u rwen ( 194 1) a nd Mo r i tz a nd J ones ( 1950 ) , w he re g ra in was r epea ted ly p assed t h rough r o ta ry h and q ue rns o fs t one i no rde rt op roduce af i ne f l ou ra s c l ose t o mode rn f l ou r a s p oss ib le . I ts eems u n l ike ly t ha t s uch p rac t ices n ecessa r i ly c ha rac te r ized t he p reh is to r ic p as t . E a r ly g ra ins we re p robab ly c onsumed a s c oa rse g roa ts o r p o r r idge , a nd w ou ld h ave r equ i red l e ss p rocess ing t i me t han t ha t a ppa ren t i nt he p roduc t ion o ff i ne f l our . I nt h is r e spec t , R icha rds ' e f f ic iency d a ta ( 1951:104 ) o n t he i n t ia l p rocess ing a nd f i na l r educ t ion o f m i l le t f o r p o r r idge i nR hodes ia p rov ide b o th a c heck o n p rev ious e s t ima tes f o r i n t ia l whea t a nd b a r ley p rocess ing , a sw e l l a s au se fu l l abo rc os tf o rt he p ound ing o fm i l le t i nw ooden mo r ta rs . T ab le 3 4 s umma r izes h e r d a ta o n s even women t imed d u r ing m i l le t p rocess ing . U s ing t he mean f i gures d e r ived f r om h er d a ta , a pp rox ima te ly 0 . 117 h rs /kg were s pen t i n t he i n i t ia l p rocess ing ( t h resh ing a nd w innow ing ) o f m i l le t , c l ose t ot he 0 . 1 h r s /kg a dop ted f o r t he i n i t ia l p rocess ing l a bo r e xpec ted f o r whea t a nd b a r ley wh ich we re n o t f r ee th resh ing . F ur the r , a0 .209 h rs /kg m i l l ing c os t f a l ls j u s t b e low t he 0 .29 h rs /kg r epor ted b y F o rbes f o r h and q ue rns a nd J asny f o r r o ta ry q ue rns u sed i nm i l l ing whea t ( r e fe rt oT ab le 3 3 ) .

T ab le 3 3 M i l l ing L abo rf o r Whea t a s R epo r ted i nV a r ious S ou rces

I MPLEMENT

#OF PEOPLE WORK ING

Y IELD /UN IT T IME

HRS /KG

SOURCE F orbes 1 965 :93 J asny 1 944 : 158

H and q ue rns R o ta ry q ue rn R o ta ry q ue rn R o ta ry q ue rn R o ta ry q ue rn R o ta ry q ue rn R o ta ry q ue rn

2 1 2 1 1 1 1

7k g /h r 3 3 k g /4 h r s 1b u ( 60 k g ) /4 h r s 2s ee rs ( 1 .8 k g ) /h r 1l b( 0 .454 k g ) /0 .5 -0 .75 h r s 1 2 .5 o z ( 0 .35 k g ) /0 .25 h rs Il b( 0 .454 k g ) /h r

0 .29 0 . 12 0 .29 0 .56 1 . 1-1 .65 0 .71 2 .20

S co t t ish q ue rn

1

1 5 l b( 6 .8 k g ) /h r

0 . 15

1 26

I f

C u rwen 1 94 1:29 C urwen 1 94 1 :31 Mo r i tz a nd J ones 1 950 :594

T ab le 3 4 M i l le t P rocess ing L abo r( h rs /kg ) o fS even Women T imed i nK ampamba V i l lage , R hodes ia

SUBJECT

T HRESH ING

W INNO W ING

MORTAR POUND ING

TOTAL

0 .270 0 .472 0 . 108 1 0 .093 0 . 152 0 .067 0 .268 2 0 .050 0 .275 0 .048 0 . 180 0 .047 3 0 . 167 0 .267 4 0 .060 0 .040 0 .200 0 .037 0 . 138 5 0 .025 0 .292 0 .048 0 .213 0 .030 6 0 .343 0 .508 0 .067 0 .098 7 ------- ------- ----- -- -------- - -0 .209 0 .326 MEAN 0 .050 0 .067 a dap ted f r om R icha rds ( 195 1: 104 )

I nt he a bsence o ff u r the r d a ta , a0 .3 h rs /kg m i l l ing c os t w i l l b e a ssumed i ns ubsequen t c a lcu la t ions a s a pp rop r ia te f o r t he f i na l p rocess ing o f w hea t a nd b a r ley , r ega rd less o f t he a r chaeo log ica l p e r iod i n vo lved .

R e tu rn R a tes f o r Whea t a nd B a r ley E xp lo i ta t ion a nd C u l t iva t ion I n t roduc t ion U s ing t he c os t a nd b ene f i t d a ta p resen ted a bove , i ti sn ow p oss ib le t oc a lcu la te p o ten t ia l r e tu rn r a tes f o rv a r ious c e rea l s t ra teg ies b ased u pon w hea t a nd b a r ley . I n i t ia l ly , p os tencoun te r r e tu rn r a tes f o r e xp lo i t ing w i ld whea t s t ands i nt he N ea r E as t w i l l b e c a lcu la ted f o r e ne rge t ic c ompa r isons w i th t he r e tu rn r a tes p rev ious ly e s tab l ished f o r v a r ious s eed r esources i nt he G rea t B as in a nd A us t ra l ia . R e tu rn r a tes w i l l t hen b e g ene ra ted f o r whea t a nd b a r ley c u l t iva t ion u nde r v ary ing y i e ld c ond i t ions a nd s pec i f ic l e ve ls o f a g r icu l tura l t echno logy . F o r c ompa ra t ive p urposes , t he a ve rage r e tu rn r a tes e xpec ted f o r p ea r l m i l le t a nd s o rghum c u l t iva t ion i n Gen ie r i V i l lage , Wes t A f r ica , u nde rt r ad i t iona l h and t i lage t e chn iques w i l l a l so b e e s tab l ished .

P os tencoun te rR e tu rn R a tes f o rE xp lo i t ing W i ld Whea tS tands P ub l ished d a ta o n t he c os ts a nd b ene f i ts o f e xp lo i t ing w i ld whea t s tands a re c u r ren t ly l i m i ted t ot he e xper imen ta l h a rves t ing o fw i ld e i nko rn ( T r i t icum b oeo t icum ) b y H a r lan ( 1967 ) o n t he s l opes o f av o lcan ic moun ta in i nt he P rov ince o f D iya rbak i r i ns ou theas te rn T urkey . A s n o ted e a r l ier , t h is r eg ion o fT urkey c on ta ins mass ive s t ands o fr obus t a nd l a rge -seeded r aces o fw i ld e i nko rn , a nd H a r lan 's d a ta may o n ly b e t aken a s a pp rop r ia te f o rt he e xp lo i ta t ion o fs uch p a tches . H a r lan ( 1967 :197 -98 ) was a b le t o h a rves t 2 .05 k g /h r o f t o ta l p l an t ma te r ia l b y h and s t r ipp ing , a nd 2 .45 k g /h r u s ing a r econs t ruc ted f l i n t s i ck le . S ince t he a c tua l g ra in c on ten t r ep resen ted 4 6% b y we igh t o ft he h a rves ted ma ter ia l , a pp rox ima te ly 0 .94 k g /h r we re h a rves ted b y h and s t r ipp ing ( r ough ly 1 . 1 h r s /kg ) , a nd 1 . 13 k g /h rw i th t he s i ck le ( r ough ly 0 .9 h rs /kg ) . U s ing t he 0 . 1 h rs /kg t h resh ing a nd w innow ing c os t a nd t he 0 . 3 h r s /kg m i l l ing c os t e s tab l ished a bove , T ab le 3 5 s umma r izes t he l a bo r r equ i red p e rk i log ram o fw i ld e i nko rn c o l lec ted . U s ing a c a lo r ic c on ten t o f 3 ,567 k ca l /kg a s d e r ived f r om t he s tanda rd a na lys is o f w i ld e i nko rn s amp les p resen ted b y H a r lan ( 1967 :198 ) , T ab le 3 6 p resen ts t he r esu l t ing p os t . e ncoun te rr e turn r a tes . I ft he h a rves ted g ra in w e re s t o red a nd a 5% l o ss o ccu r red a s ar e su l t , 1 27

T ab le 3 5 S umma ry o fT o ta l C o l lec t ing a nd P rocess ing L abo rf o rO ne K go fW i ld E inko rn Whea t

HAND S TR IPP ING %o f L abo r T o ta l H r s ( h rs )

ACT IV ITY C o l lec t ing T h resh ing a nd w innow ing M i l l ing TOTAL

F L INT S ICKLE % o f L abo r ( h r s ) T o ta l H rs

1 . 1 0 . 1 0 .3

7 3 .3 6 .7 2 0 .0

0 .9 0 . 1 0 .3

6 9 .2 7 .7 2 3 .1

1 .5

1 00 .0

1 .3

1 00 .0

T ab le 3 6 P os tencoun te r R e tu rn R a tes f o r E xp lo i t ing W i ld E inko rn Whea t

METHOD OF HARVEST ING

COLLECT ING AND P ROCESS ING T IME ( HRS /KG )

KCAL /KG

R ETURN RATE KCAL /HR

H and s t r ipp ing F l in t s i ck le

1 .5 1 .3

3 ,567 3 ,567

2 ,378 2 ,744

r e tu rn r a tes wou ld d rop f r om 2 ,378 t o2 ,259 k ca l /h rf o r h and s t r ipp ing , a nd f r om 2 ,744 t o2 ,607 k ca l /h r f o r s i ck le r eap ing . S uch r e tu rn r a tes a re much h i ghe r t han t hose p rev ious ly e s t ima ted f r om H a r lan 's d a ta ( 1 ,500 k caVh r ) b y H awkes a nd O 'Conne l l ( 1981:624 ) d ue t o t he h i gher p rocess ing l abo r ( 1h r /kg ) a nd l owe rc a lor ic c on ten t ( 3 ,000 k ca l /kg ) wh ich t hey a ssumed . Wh i le t he r e turn r a tes c a lcu la ted h e re a re q u i te h i gh , t hey s hou ld b e u nde rs tood a s o n ly r ep re s en ta t ive o f t he u ppe r r ange o f e xpec ted v a r iab i l i ty i n e xp lo i t ing a l te rna t ive s t ands o fw i ld e inko rn . F o rc ompa ra t ive p urposes , T ab le 3 7 s umma r izes p os tencoun te r r e tu rn r a tes f o re xp lo i t ing v a r ious s eed , n u t a nd a co rn r e sources i nt he G rea t B as in o ft he wes te rn U .S . a s e s tab l ished b y S imms ( 1984 , 1 987 ) t h rough e xper imen ta l c o l lec t ing a nd p rocess ing . I nt hese e xpe r imen ts , v a r ious g a the r ing t echn iques we re emp loyed w i th w i ld c e rea ls , i n c lud ing h and s t r ipp ing , s hak ing , a nd t he o ccas iona l u se o f as t ick s eed b ea te r . C e rea l p rocess ing t echn iques i n c luded s un d ry ing , h and r ubb ing o fc haf , p a rch ing o fs eeds /b rac ts u s ing c oa ls i n woven b aske ts , a nd b reeze w innow ing , d epend ing u pon t he s pec i f ic r esource b e ing p rocessed . L abo r c os ts f o r m i l l ing c e rea ls , h oweve r , we re n o t i n c luded i nS imms ' s t udy . F o r t h is r eason , t he r e tu rn r a tes p resen ted i nT ab le 3 7 wou ld b e s l igh t ly l e ss i fm i l l ing w e re a ccoun ted f o r . F ur the r , t he c o l lec t ing a nd p rocess ing c os ts g i ven f o r a co rns d o n o t i n c lude l e ach ing , s i nce a co rns o ft he Gambe l Oak h ave a l owe rt ann in c on ten t t han t he b e t te r k nown C a l i fo rn ia s pec ies , a nd c an b e e a ten w i thou t l e ach ing i ft hey a re n o t i n ges ted i nl a rge q uan t i t ies ( S imms 1 984 :243 , 1 987 :125 ) . F o r p i nyon n u ts , c o l lec t ion t echn iques r anged f r om p ick ing w ho le c ones a nd k nock ing t he s eeds o u t w i th a s t ick t h rough t he c o l lec t ion o f i n d iv idua l n u ts wh ich h ad n a tura l ly d ropped o rw e re s haken f r om t r ees . A cco rd ing t oS imms ' c a lcu la t ions ( 1 984 :238 , 1 987 :123 ) , t he u se o f ma ts p laced u nde r t he t r ees p r io r t o s hak ing c ou ld h ave p o ten t ia l ly i n c reased t he c o l lec t ion r a te e s tab l ished f o r p i ck ing u p i n d iv idua l n u ts b y 3 .5 t imes , i n c reas ing p o ten t ia l r e tu rn r a tes t o3 ,283 -4 ,263 k ca l /h r .

1 28

T ab le 3 7 P os tencoun te r R e tu rn R a tes f o rV a r ious S eed a nd N u t R esources i nt he G rea t B as in A s E xpe r imen ta l ly D e r ived b y S imms ( 1984 , 1 987 )

T YPE

S eeds

RESOURCE

KCAL /KG

RETURN RATE KCAL /HR

2 .8 2 .2 -3 .0 2 .5 2 .7 1 .7 -10 .1

3 ,660 2 ,750 3 ,000 2 ,790 3 ,050

1 ,307 9 17 -1,250 1 ,200 1 ,033 3 02 -1,794

5 .0 5 .9 7 .2 -7 .8 6 .8 -8 .0 5 .9 -10 .5

3 ,510 3 , 160 3 ,650 3 ,340 2 ,800

7 02 5 36 4 68 -507 4 18 -491 2 67 -475

6 .9 -9 .1

2 ,740

3 01-397

8 . 1-10 .0

2 ,610

2 61-322

8 .3 -14 .9 1.2 -22 .2 1 2 .8 1 5 .8 -17 .4 1 6 .2 -27 .8 3 1.0

2 ,420 3 ,070 2 ,590 2 ,540 2 ,430 2 ,820

1 62 -292 1 38 -274 2 02 1 46 -161 8 7 -150 9 1

COLLECT ING AND PROCESS ING T IME ( HRS /KG )

T ansymus ta rd ( Descu ra in ia p inna ta ) S a l ina W i ld R ye ( E lym i ss a l inas ) N u t ta l S hadsca le ( A tnp lex n u t ta l l i ) S hadsca le ( A t r ip lex c on fe r t i f lo ra ) B u l rush ( Sc i rpus s p . ) B arnya rd g rass ( Ech inocho loa c rusga l l i ) P eppe rg rass ( Lep id ium f r emon t i i ) S un f lowe r ( He l ian thus a nnus ) B lueg rass ( Poa s p . ) G r . B as in W i ld R ye ( E lymus c i ne reus ) I n d ian R ice G rass ( O ryzops i s h ymeno ides ) R eed C ana r yG rass ( Pha lar i s a rund inacea ) S c ra tchg rass ( Muh lenbe rg ia a sper i fo l ia o r S po robo l is a spe r i fo l ius ) F ox ta i l B a r ley ( Hordeumjuba tum ) S edge ( Ca rex ) S a l tg rass ( D is t ich l is s t r ic ta ) P ick leweed ( A l len ro l fea o cc iden ta l is ) S qu i r re t ta i l G rass ( S i tan ion h ys t r ix )

A co rns

Gambe l Oak ( Que rcus g ambe l l i )

3 .4

5 ,060

1 ,488

N u ts

P inyon P ine ( P inus monophy l ia )

4 .5 -5 .8

4 ,880 6 ,336

8 411 ,408+

On t he b as is o fr e la t ive p os tencoun te r r e tu rn r a tes , t he e xp lo i ta t ion o fw i ld e inko rn whea t i nt he N ea r E as t a ppea rs t o b e e ne rge t ica l ly mo re e f f ic ien t t han t he e xp lo i ta t ion o f a ny o ft he w i ld ( and p redom inan t ly p e renn ia l ) c e rea l r e sou rces i nt he G rea t B as in s t ud ied b y S imms . I n g ene ra l , t he s ame may b e s a id f o r t he t r ad i t iona l e xp lo i ta t ion o f a cac ia s eeds i nA us t ra l ia b y A lyawa ra f o rage rs . T ab le 3 8 s umma r izes p os tencoun te r r e tu rn r a te d a ta f o r A lyawa ra a cac ia e xp lo i ta t ion a s p resen ted b y O 'Conne l l a nd H awkes ( 198 1 , 1 984 ) . T he o n ly a cac ia r esou rces w i th a h ighe r r e tu rn r a te t han e inko rn whea t a re u n r ipe A cac ia c or iacea s eeds wh ich a re c onsumed w i thou t e x tens ive p repa ra t ion . On a c ompa ra t ive b as is , w i ld s tands o f r obus t a nd l a rge -seeded r aces o f e i nko rn ( and p resumab ly emme r a nd b a r ley a sw e l l ) wou ld s eem ing ly r ep resen t e ne rge t ica l ly a t t rac t ive f ood r esources r e la t ive t o many a l te rna t ive c e rea ls a nd o the r s eed r esources . Wh i le p resen t e ne rge t ic d a ta s e ts f o rw i ld c e rea l r esou rces a re e x t reme ly l i m i ted , i ti ss uspec ted t ha t s im i la r c ompa ra t ive e ne rge t ic r e la t ionsh ips w i l l o b ta in b e tween t he w i ld p rogen i to rs o f t hose c e rea ls w h ich l a te r b ecame t he f ocus o f f ood p roduc t ion s t ra teg ies , a nd a l te rna t ive c e rea l r esou rces a va i lab le i nt he s ame e nv i ronmen ts wh ich d i d n o t . P o ten t ia l ly , r e la t ive p os tencoun ter r e tu rn r a tes f o r e xp lo i t ing s t ands o fw i ld c e rea ls m igh t a ccoun t f o r why o n ly 2 9 o ft he a va i lab le 7 ,500 s pec ies o f c e rea ls wo r ld -w ide ( r ough ly 0 .4 % ) b ecame t he f ocus o fh uman f ood p roduc t ion s t ra teg ies . C e rea ls wh ich d o n o t e xh ib i t r e la t ive ly 1 29

T ab le 3 8 P os tencoun te r R e turn R a tes f o rA lyawa ra A cac ia S eed E xp lo i ta t ion I nA us t ra l ia a s C a lcu la ted b yO 'Conne l l a nd H awkes ( 1 98 1 , 1 984 )

ACAC IA E XPLO ITED A cac ia c or iacea ( un r ipe ) A . c o r iacea ( r i pe ) A . a neu ra A . c ow leana o the ra cac ias

COLLECT ING AND PROCESS ING T IME ( HRS /KG )

KCAL /KG

RETURN RATE KCAL /HR

0 .60 > 5 .25 6 .50 6 .50 6 .50

2 ,600 3 ,551 3 ,778 3 ,589 3 ,500

4 ,333 < 676 5 80 5 52 5 38

h i gh p os tencoun te r r e tu rn r a tes f o r t he i r e xp lo i ta t ion i n n a tura l s tands wou ld b e u n l ike ly c and ida tes f o rf ood p roduc t ion s t ra teg ies , s i nce t he a dd i t iona l l abo r c os ts o fc u l t iva t ion w ou ld f u r the r d im in ish t he i r r a tes o f e ne rgy r e turn t o l a bo r e xpended . A s w i l l b e d ocumen ted b e low , c u l t iva ted whea t , b a r ley , p ea r l m i l le t a nd s o rghum c on t inue t oe xh ib i t h i ghe r r e tu rn r a tes u nde r c ond i t ions o fh and t i lage t han a ny o ft he w i ld c e rea l r esou rces e xam ined b y S imms i nt he G rea t B as in .

R e turn R a tes f o rE a r ly Whea ta nd B a r ley C u l t iva t ion U n l ike t he e xp lo i ta t ion o fw i ld s t ands , c u l t iva t ion s t ra teg ies r equ i re t he r e ten t ion o fs eed f r om t he t o ta l g ra in y i e ld f o ru se a s s eeds tock i ns ow ing ac r op d u r ing t he s ubsequen t y ea r . A s p rev ious ly p resen ted i nT ab les 3 ,1 5 , 1 7 , a nd 1 8 , s ow ing r a tes f o r whea t a nd b a r ley v a ry w ide ly i nb o th t ime a nd s pace . R ega rd less , s ow ing e xper imen ts i nt he m id -n ine teen th c en tury A .D . i n d ica ted t ha t am idd le c ourse o fs ow ing amode ra te q uan t i ty o fs eed was t ob e p re fe r red o ve r t h ick o r t h in s ow ing ( S tephens 1 866 :119 -12 1) , wh i le mo re r ecen t s t ud ies i nt he N ea r E as t i n d ica te t ha tv a ry ing t he s ow ing r a te o f whea tf r om 6t o 1 4 k g /du d oes n o t s i gn i f ican t ly i n f luence e i ther t he r esu l t ing y i e ld o r t he p ro te in c on ten t o f whea t c rops (Wo rze l la 1 969 :512 ) . I n s ubsequen t e ne rge t ic c a lcu la t ions , s ow ing r a tes w i l l b e a ssumed a t 6,k g whea t /du a nd 7 k g b a r ley /du f o r a l l l a nds . Wh i le t h icke r s ow ings o n r i che r l a nds u ndoub tab ly o ccur red , i ti sn o t e xpec ted t ha t s uch v a r ia t ions wou ld s i gn i f ican t ly a fec t r e la t ive e ne rge t ic c ompar isons b e tween f ood p roduc t ion s t ra teg ies a s c a lcu la ted h e re . C ons is ten t w i th t he p r io r d i scuss ion o f s to rage p resen ted i n C hap ter 3 , i tw i l l a l so b e a ssumed t ha t a l g ra in i ss t o red , d u r ing wh ich a5% l o ss o ccu rs . T hus , i nc a lcu la t ing t he c os ts a nd b ene f i ts p e r u n i t a rea f o r c u l t iva t ing whea t p roduc ing 5 0 k g /du , t h resh ing a nd w innow ing c os ts wou ld i n vo lve t he l abo r r equ i red f o r i n t ia l ly p rocess ing 5 0 k g o f g ra in . H oweve r , m i l l ing c os ts a nd c a lo r ic b ene f i ts wou ld r e la te t ot h is i n t ia l5 0 k go fg ra in m inus 6k gf o rs eeds tock ( =4 4 k g ) a nd a 5% l o ss ( 2 .2 k g ) i ns t o rage , o r 4 1 .8 k g o f f i na l g ra in y i e ld . S im i la r ly , t he c os ts a nd b ene f i ts p e r u n i t a rea f o r b a r ley p roduc ing 6 0 k g /du wou ld i n c lude t he l a bo r o ft h resh ing a nd w innow ing 6 0 k g , wh i le m i l l ing c os ts a nd c a lor ic b ene f i ts wou ld r e la te t ot h is i n i t ia l y i e ld m inus 7 k gf o rs eed ( =5 3 k g ) a nd a5% l o ss ( 2 .7 k g ) d u r ing s t o rage , o r5 0 .3 k go ff i na lg ra in y i e ld . S ince f r ee th resh ing g ra ins d o n o t s eem ing ly a ppea ru n t i lt he s i x th m i l lenn ium B .C . ( Ha r lan 1 967 :201 , 1 977 :364 ; H e lbaek 1 966 ) , t h resh ing a nd w innow ing c os ts f o r h and t i lage s t ra teg ies u nder e a r ly l i t h ic t echno logy w i l l b e c a lcu la ted a t 0 . 1 h rs /kg o f t o ta l g ra in y i e ld . H oweve r , f o r h and t i lage u nde r a dvanced l i t h ic t echno logy a nd f o r e a r ly a n ima l t r ac t ion s t ra teg ies , t h resh ing a nd w innow ing w i l l b ec a lcu la ted a t0 .06 h r s /kg o ft o ta l g ra in y i e ld . T ab le 3 9 s umma r izes t he t o ta l l a bo r e xpec ted f o r c u l t iva t ing a d unum o f whea t o r b a r ley p roduc ing a ve rage y i e lds u nde r c ond i t ions o f h and t i lage a nd e a r ly l i t h ic t echno logy , w h i le T ab le 4 0 s im i la r ly s umma r izes t he e xpec ted l a bo r p e r d unum w i th a ve rage whea t a nd b a r ley y i e lds u nde r c ond i t ions o f h and t i lage a nd a dvanced l i t h ic t echno logy . F i na l ly , T ab le 4 1 1 30

T ab le 3 9 S umma ry o fT o ta l L abo rf o rC u l t iva t ing One D unum o fL and U nde rH and T i l lage a nd E a r ly L i th ic T echno logy w i th Whea t Y ie ld ing 5 0 K g /Du o rB a r ley Y ie ld ing 6 0 K g /Du

ACT IV ITY T i l l ing S ow ing a nd H a r row ing R eap ing T h resh ing a nd w innow ing M i l l ing TOTAL

WHEAT %o f T o ta l H rs

L abo r ( h rs )

BARLEY L abo r % o f T o ta l H rs ( h rs )

3 5 .0 8 .0 3 4 .0 5 .0 1 2 .5

3 7 .0 8 .5 3 6 .0 5 .3 1 3 .2

3 5 .0 8 .0 2 9 .0 6 .0 1 5 .1

3 7 .6 8 .6 3 1.2 6 .4 1 6 .2

9 4 .5

1 00 .0

9 3 .1

1 00 .0

T ab le 4 0 S umma ry o fT o ta l L abo rf o rC u l t iva t ing One D unum o fL and U nde r H and T i l lage a nd A dvanced L i th ic T echno logy w i th Whea t Y ie ld ing 5 0 K g /Du o r B a r ley Y ie ld ing 6 0 K g /Du

ACT IV ITY T i l l ing S ow ing a nd H a r row ing R eap ing T h resh ing a nd w innow ing M i l l ing TOTAL

WHEAT %o f T o ta l H rs

L abo r ( h r s ) 3 0 .0 8 .0 2 0 .0 3 .0 1 2 .5

4 0 .8 1 0 .9 2 7 .2 4 .1 1 7 .0

7 3 .5

1 00 .0

BARLEY L abo r ` ) / 0o f ( h rs ) T o ta l H rs 3 0 .0 8 .0 1 7 .0 3 .6 1 5 .1 , 7 3 .7

4 0 .7 1 0 .9 2 3 .0 4 .9 2 0 .5 1 00 .0

T ab le 4 1 S umma ry o fT o ta l L abo rf o rC u l t iva t ing One D unum o fL and U nde rA n ima lT rac t ion T i l lage a nd E a r ly M e ta lT echno logy w i th Whea tY ie ld ing 5 0 K g /Du o r B a r ley Y ie ld ing 6 0 K g /Du

ACT IV ITY T i l l ing a nd s ow ing H a r row ing R eap ing T h resh ing a nd w innow ing M i l l ing TOTAL

L abo r ( h rs )

WHEAT %o f T o ta l H rs

BARLEY L abo r % o f ( h rs ) T o ta l H rs

1 6 .0 4 .0 2 0 .0 3 .0 1 2 .5

2 8 .8 7 .2 3 6 .1 5 .4 2 2 .5

1 2 .0 4 .0 1 7 .0 3 .6 1 5 .1

2 3 .2 7 .7 3 2 .9 7 .0 2 9 .2

5 5 .5

1 00 .0

5 1.7

1 00 .0

1 31

s umma r izes t he t o ta l l a bo r e xpec ted p e r d unum w i th a verage w hea t a nd b a r ley y i e lds u nde r c ond i t ions o f a n ima l t r ac t ion t i lage a nd e a r ly me ta l t echno logy . O n ac ompa ra t ive b as is , a n a pp rox ima te 2 2% i n c rease i nt he l a bo re f ic iency f o rw hea tc u l t iva t ion a nd a2 1% i n c rease i nt he l a bo r e f ic iency o f b a r ley c u l t iva t ion w ou ld e x is t b e tween e a r ly a nd a dvanced f o rms o fh and t i lage s t ra teg ies a s c a lcu la ted h e re . F ur the r , a n a pp rox ima te 4 1% i n c rease i nt he e f ic iency o f w hea t c u l t iva t ion a nd a4 5% i n c rease i nt he e f f ic iency o fb a r ley c u l t iva t ion w ou ld e x is tb e tween e a r ly h and t i lage a nd i n i t ia l a n ima lt r ac t ion s t ra teg ies . F i gure 1 7 d iag rams t he r e la t ionsh ips e xpec ted b e tween t he l a bo r c os ts o f w hea t a nd b a r ley c u l t iva t ion u nde r e ach o f t hese t echno log ica l s cena r ios ( b roken l i nes , r ead o f f t he v e r t ica l s ca le o nt he r i gh t ) a nd t h ef i na lc a lo r ic b ene f i ts o fw hea ta nd b a r ley a tv a r ious y i e lds a f te r t he r e ten t ion o f s eeds tock a nd s t o rage l o ss i sa ccoun ted f o r ( s o l id l i nes , r ead o f f t he v er t ica l s ca le o n t he l e f t ) . I nt h is f i gure , t he l a bo r c os ts a ssoc ia ted w i th h and t i lage a nd e a r ly l i t h ic t e chno logy , h and t i lage a nd a dvanced l i t h ic t echno logy , a nd a n ima l t r ac t ion t i lage w i th e a r ly me ta l t echno logy , a re l a be led " A , " " B , " a nd " C " r e spec t ive ly . O n t he b as is o ft hese d a ta , F i gure 1 8 d i ag rams t he c hanges e xpec ted i nr e tu rn r a tes f o r w hea t a nd b a r ley c u l t iva t ion u nde rt hese t h ree t e chno log ica lc ond i t ions ( aga in l a be led " A , " " B , " a nd " C " ) , g i ven v a r iab le g ra in y i e lds p e r d unum . S ince t he e f f ic iency o f c u l t iva t ing c e rea ls i n c reases t h rough t i me d ue t ot e chno log ica l i n nova t ions , c u l t iva t ion r e tu rn r a tes a l so i n c rease . A sd i scussed e a r l ie r ,t h e p roduc t ive d i v is ion b e tween " ma rg ina l " a nd " op t ima l " a g r icul tu ra l l a nds i sa ssumed t o l i e b e tween 5 0 k g /du f o r whea t a nd 6 0 k g /du f o r b a r ley . D ue t o i n c reased e f ic iency i nc u l t iva t ing c e rea ls a s ar e su l t o ft echno log ica l i n nova t ions , t he e ne rge t ic d i v is ion b e tween " ma rg ina l "a nd " op t ima l " l a nd a s measu red b y r e tu rn r a tes w ou ld s h i f t u pwa rd t h rough t ime .

1 50

500:

. • • • • •

• •

1 00 °

400= • •• • • •

-

• •

-

41 1 1

300

-75

• • • • • •

a t e l v iheat-

1 4 1 4

N O W

200i

1 25

• •• • • -

a

-50

C

b

HOBV1 dO

F INAL CALOR IC Y IELD/ DU

600=

e lN ei

100z

-25

0 0

25

50

75

100

1 25

1 50

1 75

200

SEED Y IELD (Kg/ Du)

F i gu re 1 7 . R e la t ionsh ips b e tween c a lo r ic p roduc t iv i ty a nd l a bo rf o rw hea t a nd b a r ley c ul t iva t ion a tv a r ious s t ages o ft e chno logy a nd v a r iab le y i e lds . C a lo r ic b ene f i ts ( s o l id l i nes ) r ead o f f t he s ca le o nt he l e f t . L abo rc os ts ( b roken l i nes ) r ead o ft he s ca le o nt he r i gh t . " A " d es igna tes l a bo r c os ts f o r w hea t a nd b a r ley c u l t iva t ion u nder c ond i t ions o f h and t i lage a nd e a r ly l i t h ic t e chno logy . " B " s im i la r ly d es igna tes l a bo rc os ts u nde rc ond i t ions o fh and t i lage a nd a dvanced l i t h ic t echno logy . " C "d es igna tes l a bo rc os ts u nde rc ond i t ions o fa n ima lt r ac t ion t i lage a nd e a r ly me ta l t echno logy . 1 32

6000 L t

h .

= 5000 bee

V

. 2 4000 0

t

-

B

L u < 3000 r c

c r

z I • - 2000 _ ,

1000

O

7 _

Margina l" 1 111 -

-L Tr1171

0

25

50

1 -1 1

"Opti mal" 1 1

1

1

75

1-

1 1

100

1 1

1

1 1

125

1

1

1

1 1,

150

1

1 ,111

175

200

Y IELD ( kg/du) F i gu re 1 8 . R anges o f p o ten t ia l r e tu rn r a tes f o r whea t a nd b a r ley c u l t iva t ion u nde r v a r ious s t ages o f t echno logy a nd v a r iab le y i e lds . " A " d es igna tes c ond i t ions u nder h and t i lage a nd e a r ly l i t h ic t echno logy , " B " d es igna tes h and t i lage a nd a dvanced l i t h ic t echno logy , a nd " C " d es igna tes a n ima l t r ac t ion t i lage w i th e a r ly me ta l t echno logy . S ince t he p roduc t ive d i v is ion b e tween " op t ima l " a nd " ma rg ina l " l a nd o ccu rs a t ay i e ld o fa round 5 0 k g /du f o r whea t ( s o l id l i ne ) a nd 6 0 k g /du f o r b a r ley ( dashed l i ne ) , t he e ne rge t ic d i v is ion b e tween t hese c a tego r ies o f l a nd a s m easu red i nr e turn r a tes s h i f ts u pwa rd a s af unc t ion o f i n c reas ing t echno log ica l e f f ic iency t h rough t ime . I na l c ases , t he u ppe r r ange o f p o ten t ia l r e turn r a tes u nde r " op t ima l " c ond i t ions w ou ld b e r es t r ic ted t o ar e la t ive ly sma l l p ropo r t ion o ft he t o ta l a va i lab le a rab le l a nd . T he l owe r r a nge o fr e turn r a tes u nde ra ve rage a nd " ma rg ina l " c ond i t ions s hou ld c ha rac te r ize t he ma jo r i ty o fa la va i lab le a rab le l a nd .

T he c hang ing n a tu re o ft h is e ne rge t ic d i v is ion i sd ep ic ted i nt he l owe r l e f t -hand c o rne r o f F i gu re 1 8 . As o l id v er t ica l l i ne d rawn u pwa rd f r om whea t y i e ld ing 5 0 k g /du i n tersec ts w i th e ach o f t he d i f feren t r e turn r a te c urves e xpec ted u nder s pec i f ic t echno log ica l c ond i t ions a t p rog ress ive ly h i ghe r l e ve ls . Ad ashed v er t ica l l i ne d rawn u pwa rd f r om b a r ley y i e ld ing 6 0 k g /du s im i la r ly d ep ic ts t he s h i f ts e xpec ted i nb a r ley r e tu rn r a tes g i ven t he s ame c hanges i na g r icu l tu ra l t e chno logy . U nde rc ond i t ions o fh and t i lage w i th e a r ly l i t h ic t echno logy , t he e ne rge t ic d i v is ion b e tween " ma rg ina l " a nd " op t ima l " a g r icu l tura l l a nds a s measured i nr e tu rn r a tes wou ld o ccu r b e tween 1 ,469 k ca l /h r f o r whea t y i e ld ing 5 0 k g /du a nd 1 ,794 k ca l /h r f o r b a r ley y i e ld ing 6 0 k g /du . U nde rc ond i t ions o fh and t i lage a nd a dvanced l i t h ic t echno logy , t h is d i v is ion wou ld s h i f t u pwa rd t o b e tween 1 ,888 k ca l /h r f o r whea t y i e ld ing 5 0 k g /du a nd 2 ,266 k ca l /h r f o r b a r ley y i e ld ing 6 0 k g /du . W i th t he a dven t o f a n ima l t r ac t ion t i lage a nd e a r ly me ta l t echno logy , t h is e ne rge t ic d i v is ion wou ld r i se t ob e tween 2 ,50 1 k ca l /h r f o r whea t y i e ld ing 5 0 k g /du a nd 3 ,230 k ca l /hr f o r b ar ley y i e ld ing 6 0 k g /du . I n g ene ra l , t he r ange o f r e turn r a tes b e low t hese p roduc t iv i ty d iv is ions s hou ld r e f lec t t hose e xpec ted o n a ma jo r i ty o f a l a va i lab le a rab le l a nd , w h i le t he r ange o fr e turn r a tes a bove t hese p roduc t iv i ty d i v is ions s hou ld r e f lec t t hose e xpec ted o n am uch sma l le rp ropo r t ion o fa l a va i lab le a rab le l a nd .

1 33

I nt he e ne rge t ic c ompa r isons b e tween c e rea l a nd p as to ra l s ubs is tence s t ra teg ies w h ich w i l l b e p resen ted i n Chap te r 7 , t hese g ross e ne rge t ic d i v is ions b e tween " ma rg ina l " a nd " op t ima l " a g r icu l tura l l a nds u nde r c hang ing t echno log ica l c ond i t ions w i l l b e u sed t o g ene ra te a rchaeo log ica l ly t es tab le h ypo theses c once rn ing t he r e la t ive s pa t ia l a nd t empo ra l e vo lu t ion a nd a dop t ion o f a l te rna t ive s ubs is tence o p t ions d ur ing t he H o locene e poch o ft he N ea r E as t a nd N or th A f r ica . B e fo re p roceed ing , h oweve r , s ome e ne rge t ic c ompa r isons b e tween t he w hea ta nd b a r ley r e tu rn r a tes c a lcu la ted a bove a nd r e turn r a tes f o rp ea r l m i l le t a nd s o rghum ,t he mos t i mpo r tan tc e rea l d omes t ica tes o fA f r ica , w i l l b e i n s t ruc t ive .

C ompa ra t ive R e tu rn R a tes f o rC u l t iva t ing S o rghum a nd P ea r l M i l le t I n t roduc t ion F o r c ompa ra t ive p urposes , r e tu rn r a tes f o r c u l t iva t ing p ea r l m i l le t ( Penn isetu rn amer icanum ) a nd s o rghum ( Sorghum b i co lo r ) u nde r t r ad i t iona l me thods o f h and t i lage w i l l b e g ene ra ted f r om t he q uan t i ta t ive d a ta p resen ted b y H aswe l l ( 1953 ) a s d ocumen ted i nG en ie r i V i l lage i n Gamb ia , Wes t A f r ica . T hese d a ta s pec i f ica l ly i n vo lve l abo rc os ts u s ing mode rn m e ta l i mp lemen ts a nd mean y i e lds f o rt he e n t i re v i l lage a s r eco rded i n1 949 . N o a t temp tw i l l b e made t oa ccoun tf o r amo re p r im i t ive c u l t iva t ion t echno logy , n o rc an i tb e a ssumed t ha t mean y i e lds i n 1 949 a t Gen ie r i V i l lage n ecessa r i ly r e f lec t l o ng te rm mean y i e lds i no the r s avanna r eg ions o f A f r ica . F ur the r , c u l t iva ted r aces o f p ea r l m i l le t a nd s o rghum e xh ib i t s t r ik ing mo rpho log ica l d i fe rences f r om t he i rw i ld p rogen i to rs . H ence , t he r e tu rn r a tes wh ich w i l l b e d e r ived f r om t hese d a ta may o n ly b e t aken a s a dequa te ly r e f lec t ing mode rn r e tu rn r a te p oss ib i l i t ies , b u t m ay n o t n ecessa r i ly r e f lec t p o ten t ia l p reh is to r ic r e tu rn r a tes . S o rghum a nd p ea r l m i l le t a re n o t o n ly t he p r ima ry s ubs is tence c rops o f Gen ier i V i l lage , b u t a l so r ep resen t t he d om inan t c e rea l d omes t ica tes o f wha t h as b een t e rmed t he " A f r ican S avanna C omp lex , " much a s whea t a nd b a r ley r ep resen t t he d om inan t c e rea l d omes t ica tes o f t he " Nea r E as te rn C omp lex " ( Ha r lan 1 977 ) . Wh i le b o th p ea r l m i l le t a nd s o rghum a ppea rt oh ave b een d e r ived f r om w i ld p rogen i to rs w h ich e vo lved i nt he s avanna z ones o fA f r ica s ou th o ft he S aha ra a nd n o r th o ft he e qua to r , p ea r l m i l le t i sd ec ided ly mo re d rough t res is tan t a nd x e rophy t ic t han s o rghum ( Ha r lan 1 977 :374 -376 ) . A s p rev ious ly d i scussed i nC hap te r 3 , p ea r l m i l le t , l i ke ma ize ( Zea mays ) a nd J ob 's t ea rs ( Co ix l a ch rymajob t ) , i s ap redom inan t c ross -po l l ina to r , w h i le s o rghum , l i ke r y e ( Seca le c e rea le ) , i sp a r t ia l ly s e l f - a nd p a r t ia l ly c ross -po l l ina t ing ( F ryxe l l 1 957 ) . A lso l i ke ma ize , c u l t iva ted v a r ie t ies o f s o rghum a nd p ea r l m i l le t e xh ib i t i n c reases i nb o th i n f lo rescence s i ze a nd n umbe r r e la t ive t ot he i rw i ld c oun te rpa r ts ( Hada r %e ta l . 1 973 :3 16 , 3 18 ) . H ence , u n l ike t he y i e lds o fa nc ien t a nd mode rn l a nd races o ft he p redom inan t ly s e l f -po l l ina t ing w hea ts a nd b a r leys , t he y i e lds o f mode rn s o rghums a nd p ea r l m i l le ts may b e e xpec ted t ob e g ene ra l ly h i ghe rt han t hose o b ta ined f r om t he i rp reh is to r ic p rogen i to rs .

Mean R e turn R a tes f o rC u l t iva t ing P ea r lM i l le t a tG en ie r i V i l lage i n1 949 B y r eco rd ing t he t o ta l l a bo r e xpended i nf i e ld p repa ra t ion , weed ing , b i rd s ca r ing ( wh ich i s c r i t ica l i nm i l le tc u l t iva t ion ) a nd h arves t ing p e ra c re among 1 9 h u t c ompounds i n1 949 , H aswe l l ( 1953 :32 ) f ound t ha t t he mean l a bo r e xpended b y v i l lage rs i nc u l t iva t ing e a r ly ma tu r ing m i l le t was 2 67 h rs /ac re , o r a pp rox ima te ly 6 5 .9 h rs /du . F ur ther , t he mean y i e ld o f e a r ly m i l le t was e s tab l ished a t 2 85 l b s ( r ough ly 1 29 .3 k g ) /ac re ( 1953 :39 ) , o r a pp rox ima te ly 3 1 .9 k g /du . T h is r ep resen ts a r e tu rn o f 1 . 07 l b s ( r ough ly 0 .485 k g ) /h r ( 1953 :57 ) . U s ing t he 0 . 117 h rs /kg mean t h resh ing a nd w innow ing l a bor f o r m i l le t r eco rded b y R icha rds ( 1951:104 ) a s p rev ious ly p resen ted i nT ab le 3 4 , a n a dd i t iona l 3 .7 h rs wou ld h ave b een e xpended t o o b ta in t he i n i t ia l 3 1 .9 k g /du y i e ld . S ince m i l le t s ow ing r a tes a t Gen ier i r anged f r om 5 13 l b s ( r ough ly 2 . 3 -5 .8 k g ) /ac re ( Haswe l l 1 953 :52 ) , a pp rox ima te ly 0 .57 -1 .4 k g /du s eeds tock wou ld h ave b een r e ta ined f o r p lan t ing a c rop d u r ing t he n ex t y ea r . A ssum ing a 1k g /du s ow ing r a te , 3 0 .9 k g /du w ou ld b e l e f t a s t he s t o rab le y i e ld . W i th a 5% l o ss d ur ing s t o rage ( app rox ima te ly 1 . 6 k g ) , t he f i na l y i e ld wou ld d rop t oa pp rox ima te ly 2 9 .3 k g /du . A t a mean 0 .2 1 h rs /kg m i l l ing c os t ( R icha rds 1 939 :104 ) , a n a dd i t iona l 6 . 2 h r s o fl a bo rw ou ld h ave b een r equ i red , r a is ing t he t o ta l l a bo rc os ts o fo ne d unum o f m i l le t p roduc ing 3 1 .9 k g /du t o7 5 .9 h ou rs ( 66 + 3 .7 + 6 .2 = 7 5 .9 ) . A t ac a lo r ic c on ten t o f3 ,650 k ca l /kg ( Ayk royd a nd D ough ty 1 970 :18 , T ab le 2 ) , t he f i na l y i e ld o f 2 9 .3 k g /du 1 34

w ou ld r ep resen t 1 06 ,359 k ca l . A t 7 5 .9 h ou rs o f l a bo r , t he mean r e turn r a te f o r c u l t iva t ing a m ode rn r ace o fp ea r l m i l le t a tG en ie r i V i l lage i n1 949 u nde rh and t i lage w i th mode rn i mp lemen ts a moun ted t oa pp rox ima te ly 1 , 40 1 k ca l /h r . B y c ompa r ison w i th t he r e tu rn r a tes e s tab l ished e a r l ie r f o r whea t a nd b a r ley c u l t iva t ion , m i l le t y i e ld ing 3 1 .9 k g /du s eem ing ly e xh ib i ts a r e tu rn r a te wh ich f a l ls s ho r t o f a l whea t a nd b a r ley s t ra teg ies y i e ld ing 5 0 a nd 6 0 k g /du r espec t ive ly . I ft he r e tu rn r a te f o rc u l t iva t ing m i l le t a t G en ie r i V i l lage i n1 949 a dequa te ly r e f lec ts t he mean r e tu rn r a te f o rm i l le t u nde r h and c u l t iva t ion i ng ene ra l , whea t a nd b a r ley c u l t iva t ion u nde r a ve rage c ond i t ions s hou ld r ep resen t f a r mo re a trac t ive c e rea l s ubs is tence o p t ions t han m i l le tc u l t iva t ion .

Mean R e turn R a te f o rC u l t iva t ing S o rghum a tG en ie r i V i l lage i n1 949 A s w i th p ea r l m i l le t , H aswe l l t abu la ted t he t o ta l l abo r e xpended p e r a c re i nt he f i e ld p r epa ra t ion , p l an t ing , weed ing a nd h a rves t ing o fs o rghum among 1 9 h u tc ompounds a t Gen ie r i V i l lage i n 1 949 . T he mean l a bo r e xpended i n s uch t asks was 6 7 h rs /ac re ( 1953 :32 ) , o r a pp rox ima te ly 1 6 .5 h rs /du . S o rghum a ppa ren t ly r equ i res o n ly 2 5% o ft he t o ta l l a bo r p e rd unum r e qu i red i nm i l le t c u l t iva t ion . T he mean y i e ld o f s o rghum t ha t y ea r was 1 58 l b s ( r ough ly 7 1 .7 k g ) /ac re ( 1953 :39 ) , o r a pp rox ima te ly 1 7 .7 k g /du . T h is r ep resen ts a r e tu rn o f 2 .36 l b s ( r ough ly 1 . 07 k g ) /h r ( 1953 :57 ) a s c ompa red t o t he m i l le t r e tu rn o fa round 0 .485 k g /h r . A ssum ing ( w i th r e se rva t ions ) t ha t t he t h resh ing a nd w innow ing o f s o rghum wou ld r equ i re t he s ame 0 . 117 h r s /kg a s m i l le t , a n a dd i t iona l 2 . 1 h rs wou ld h ave b een e xpended t oo b ta in t he i n i t ia l 1 7 .7 k g /du y i e ld . Wh i le H aswe l l d id n o t p rov ide t he s ow ing r a te u sed f o r s o rghum a t Gen ie r i , i ti ss a id t ha t s o rghum i sn o rma l ly s own a t ar a te o f0 .3 -0 .9 k g /du u nde rd ry fa rm ing c ond i t ions ( Why te e ta l . 1 959 :372 ) . A ssum ing a 1k g /du s ow ing r a te , t he s t o rab le y i e ld a t Gen ie r i wou ld amoun tt o 1 6 .7 k g /du . W i th a 5% l o ss d u r ing s t o rage ( app rox ima te ly 0 .8 k g ) , t he f i na l y i e ld wou ld d rop t o a pp rox ima te ly 1 5 .9 k g /du . A ga in a ssum ing ( w i th r eserva t ions ) t ha t t he m i l l ing o f s o rghum r equ i res t he s ame 0 .21 h r s /kg l a bo r a s m i l le t , a n a dd i t iona l 3 .3 h rs o f l a bo r wou ld h ave b een r equ i red , r a is ing t he t o ta l l a bo r c os t o f o ne d unum o f s o rghum y i e ld ing 1 7 .7 k g /du t o 2 1 .9 h ou rs ( 16 .5 + 2 . 1 +3 .3 = 2 1 .9 ) . A t ac a lo r ic c on ten t o f3 ,550 k ca l /kg ( Ayk royd a nd D ough ty 1 970 :18 , T ab le 2 ) , t he f i na l y i e ld o f 1 5 .9 k g /du wou ld r ep resen t 5 6 ,445 k ca l . A t 2 1 .9 h ou rs o f l a bo r , t he mean r e tu rn r a te f o rc u l t iva t ing a mode rn r ace o fs o rghum a t Gen ie r i V i l lage i n1 949 u nde rh and t i lage w i th mode rn i mp lemen ts amoun ted t oa pp rox ima te ly 2 ,577 k ca l /h r . B yc ompa r ison w i th t he r e tu rn r a tes e s tab l ished e a r l ie rf o r whea t a nd b a r ley c u l t iva t ion , t he r e tu rn r a te f o rs o rghum y i e ld ing o n ly 1 7 .7 k g /du a t Gen ie r i V i l lage f a l ls c l ose t ot he 2 ,50 1 k caVh r e xpec ted f o r whea t y i e ld ing 5 0 k g /du u nde rc ond i t ions o fa n ima l t r ac t ion t i lage a nd e a r ly me ta l t e chno logy , b u t l e ss t han t he 3 ,230 k ca l /h re xpec ted f o r b a r ley y i e ld ing 6 0 k g /du u nde r s im i la r c u l t iva t ion c ond i t ions . F ur the r , t he 2 ,577 k ca l /h r e s t ima ted f o r s o rghum c u l t iva t ion i s much h i gher t han t he 1 ,401 k ca l /h r e xpec ted f o r m i l le t y i e ld ing 3 1 .9 k g /du u nde r e qu iva len t t e chno log ica l c ond i t ions . I nl i gh t o ft hese c ompa ra t ive r e tu rn r a tes , i ti sn o t s u rp r is ing t ha t wh i le s o rghum i sl e ss d rough t res is tan t a nd h ence e co log ica l ly mo re d emand ing t han p ea r l m i l le t , i t h as b ecome t he mos t i mpor tan t A f r ican c e rea l d omes t ica te wo r ld -w ide « ( Ha r lan a nd S tem le r 1 976 :465 ) . Wh i le t hese c ompa ra t ive d a ta o n t he e ne rge t ics o f A f r ican c e rea l d omes t ica tes a re o bv ious ly l i m i ted , t hey d o p rov ide i n te res t ing a venues f o r f u ture s ubs tan t ive r esea rch . A s d i scussed i nC hap te r3 ,p ea r l m i l le t i s ap redom inan t c ross -po l l ina to ra nd s o rghum ap a r t ia l s e l f , p a r t ia l c ross -po l l ina to r . L i ke c o rn a nd J ob 's T ea rs , t hey e xh ib i t t he mos t d rama t ic c hanges i n m o rpho log ica l t r a i ts among a l d omes t ica ted c e rea ls . Y e t , e ven w i th s uch mo rpho log ica l ( and c onsequen t ly p roduc t ive ) s h i f ts i np l ace , t he r e turn r a tes a ssoc ia ted w i th t he i rc u l t iva t ion s eem h a rd ly s pec tacu la r when c ompa red w i th t hose o b ta ined i nt he c u l t iva t ion o f t he f es tuco id c e rea ls whea t a nd b a r ley i n mo re n or the r ly l a t i tudes . P o ten t ia l ly , t he l a te ra ppea rance o fc e rea l c u l t iva t ion s t ra teg ies i n s ub -Saha ran A f r ica was a r esu l t o f l owe r e ne rgy r e turn r a tes f o r c u l t iva t ing t he a va i lab le p an ico id c e rea l s pec ies w h ich h ad e vo lved i n t h is r eg ion , a nd a s omewha t l a te rs h i f tt o mode rn e nv i ronmen ta l c ond i t ions , a s d ocumen ted i nC hap te r7 .

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CHAPTER 7

E NERGET IC E XPECTAT IONS AND T HE ARCHAEOLOG ICAL RECORD

I n t roduc t ion I nt h is c hap te r , t he e ne rge t ic d a ta d eve loped i nC hap te rs 5 a nd 6w i l l b e u sed t oe xp lo re t h e e x ten tt ow h ich s imp le o p t im iza t ion mode ls d e r ived f r om b ehav io ra l e co logy m igh t e nhance c u r ren t m e thods o fa rchaeo log ica l i n fe rence c oncern ing t he s pa t ia l a nd t empo ra l e vo lu t ion a nd a dop t ion o f a l te rna t ive p as to ra l a nd c e rea l s ubs is tence o p t ions i nt he N ea r E as t a nd N or th A f r ica . Wh i le t he re a re s eve ra l a rchaeo log ica l a nd e thnog raph ic i s sues w h ich c ou ld b e a dd ressed u s ing t he d a ta g enera ted i nt h is s t udy , i ti sn o t t he i n ten t o ft he p resen t a na lys is t o a dd ress mo re i s sues t han a r e n ecessa ry t ot e s tt he g ene ra l u t i l i ty o ft he e vo lu t iona ry a pp roach a dop ted h e re . H ence , t he s ub jec ts a dd ressed i nd ep th i nt h is c hap te r a re r e s t r ic ted t os pec i f ic i s sues o fg ene ra l a r chaeo log ica lc once rn . A f ter a n i n t ia l d i scuss ion o fp o ten t ia l p rob lems i nt he i n te rp re ta t ion o fz ooa rchaeo log ica l d epos i ts r e levan tt oe a r ly p as tora l ism , t he r e la t ive e ne rge t ics o fa l te rna t ive p as to ra l s t ra teg ies a r e e xam ined i no rder t o f o rmu la te a rchaeo log ica l ly t es tab le e xpec ta t ions c oncern ing t he s pa t ia l a nd t empo ra l e vo lu t ion a nd a dop t ion o f s t ra teg ies i n vo lv ing s pec i f ic h e rd s pec ies . T empo ra l e xpec ta t ions b ased u pon t he r e la t ive e ne rge t ics o fp as to ra l s t ra teg ies a s c a lcu la ted i nt h e p resen t s t udy wou ld i n ver t t he c ommon ly a ccep ted s equence o f g oa ts , s heep , c a t t le a nd c ame ls a st he o rde r i nw h ich t hese s pec ies w e re d omes t ica ted . T he n a tu re o ft h is s t r ik ing i n cons is tency i se xp lo red . T he r e la t ive e ne rge t ics o f e a r ly c e rea l s t ra teg ies a re t hen e xam ined , b r ie f ly n o t ing t ha t s pec i f ic e ne rge t ic e xpec ta t ions a cco rd w e l l w i th c e r ta in c ommon ly a ccep ted i n te rp re ta t ions o f t h e a r chaeo log ica l r eco rd . T es ts o f mo re s ubs tan t ive i mp l ica t ions o ft he r e su l ts p resen ted h e re a re c u r ren t ly p rec luded b y t he l a ck o fa pp rop r ia te d a ta s e ts . A t ten t ion i st he re fo re f ocussed u pon t he i n c lus ion o fc erea ls i np reh is to r ic d i e ts a nd t he s pa t ia l a nd t empo ra l a ppea rance o f c e rea l c u l t iva t ion s t ra teg ies i nt he N ea r E as t a nd N or th A f r ica i no rder i to p rov ide i n terp re t ive i n s igh ts a nd d i rec t ions f o rf u tu re r e sea rch . C ompa r isons a re t hen made b e tween t he r e la t ive e ne rge t ics o f p as to ra l a nd c erea l s ubs is tence o p t ions i no rde r t o g ene ra te a rchaeo log ica l ly t es tab le e xpec ta t ions c once rn ing t h e s pa t ia l a nd t empora l a ppea rance o f a l te rna t ive p as to ra l a nd c erea l s ubs is tence o p t ions . I n i t ia l ly , p o ten t ia l v a r ia t ions i nt he e xchange v a lue o fc e rea l a nd a n ima l c a lo r ies a re d i scussed . T he s ub jec t o f m ixed s ubs is tence s t ra teg ies i n vo lv ing v a r ious c omb ina t ions o f f o rag ing , p as to ra l a nd c e rea l s ubs is tence o p t ions i st hen c ons ide red . Wh i le t he p resen t s t udy p rov ides s eve ra l i n s igh ts i n to t he d i rec t ions w h ich f u ture r e sea rch o n t h is s ub jec t m igh t t ake , t he a va i lab le e ne rge t ic d a ta a re b es t a pp l ied t ob roade ri s sues c oncern ing t he s pa t ia l a nd t empo ra l a ppea rance o fd i s t inc tp as to ra l a nd c e rea ls ubs is tence s t ra teg ies . M u tua l ly e xc lus ive e ne rge t ic e xpec ta t ions c once rn ing t he s pa t ia l a nd t empo ra l e vo lu t ion a nd a dop t ion o f c erea l a nd p as tora l s ubs is tence s t ra teg ies a re g enera ted t h rough a c ompa r ison o ft he r e turn r a tes f o r e a r ly c e rea l s t ra teg ies , a nd mea t -on ly a nd mea t p l us m i lk p as tora l s t ra teg ies . T hese e xpec ta t ions a re t hen t es ted a ga ins t H o locene e poch a r chaeo log ica ld a ta f r om t he N ea r E as t a nd N or th A f r ica w h ich r e f lec t t he s pa t ia l a nd t empo ra l a ppea rance o fp as to ra l a nd c e rea l s ubs is tence s t ra teg ies .

P o ten t ia l P rob lems i nt he I n te rp re ta t ion o fF auna l D epos i ts R e levan tt oE a r ly P as to ra l F ood P roduc t ion I nC hap te r 2 ,t he u se o fg ame d r ive l i nes a nd t r ap c o r ra ls f o rt he i n i t ia l c ap tu re a nd c on t ro l o f h e rd a n ima ls was p os i ted a s o ne mechan ism b y wh ich h un te r -ga the re r p opu la t ions m igh t h ave e s tab l ished e a r ly s ubs is tence h erds . Once p as to ra l s t ra teg ies we re a dop ted , i ts eems r easonab le t o e xpec t t ha t h e rd h usbandry t h rough v a r ious p a t te rns o fs e lec t ive a ge -sex c u l l ing wou ld h ave b een p rac t iced , a nd t ha t a rchaeo log ica l f auna l d epos i ts s hou ld g ene ra l ly r e f lec t s uch p a t te rns o f e conom ic d ec is ion mak ing ( Dav is 1 983 ; H esse 1 982 , 1 984 ; K le in a nd C ruz U r ibe 1 984 ; P ayne 1 972a , 1 973 ; P erk ins 1 964 , 1 973 ; R edd ing 1 98 1 , 1 983 ) . H oweve r , t he u se o f a ge a nd s ex p ro f i les a s r econs t ruc ted f r om f auna l d epos i ts t o i n terp re t p a t te rns o f h uman c onsump t ion a nd h e rd managemen t s t ra teg ies i s a ma t te r o f s ome d eba te ( Bake r a nd B ro thwe l l 1 980 ; C hang a nd K os te r 1 986 ; C o l l ie r a nd Wh i te 1 976 ; H enry 1 975 ; H esse 1 982 ; J a rman a nd W i lk inson 1 972 ; P ayne 1 972a ; S immons 1 975 -76 ; W i lk inson 1 976 ) . I n i t ia l ly , t he a b i l i ty t o e f fec t ive ly i n terp re t f auna l ma te r ia ls i sc omp l ica ted b y v a ry ing c ond i t ions o f b one d epos i t ion , s urv iva l , r ecovery a nd i d en t i f ica t ion , a nd i s made p a r t icu la r ly d i f f icu l t b y o ur l i m i ted u nde rs tand ing o f t he w ide r ange o f n a tura l a nd c u l tu ra l p rocesses a fec t ing t he c rea t ion o f a c tua l f auna l a ssemb lages ( B in fo rd 1 978 , 1 98 1; G i f fo rd 1 977 , 1 978 , 1 981; O 'Conne l l 1 987 ; P ayne 1 972a ) . I nt he a bsence o f s t ud ies wh ich s pec i f ica l ly a dd ress d i f ferences a nd s im i la r i t ies i nt he n a tura l a nd c u l tura l p rocesses a f fec t ing f auna l d epos i ts c rea ted b y h un te r -ga the re r a nd p as to ra l p opu la t ions , t he i n te rp re t ive u t i l i ty o f a ge a nd s ex p ro f i les t od ife ren t ia te s uch s t ra teg ies r ema ins s pecu la t ive . I n g enera l , i ti sf r equen t ly a ssumed t ha t u nde r h erd managemen t s t ra teg ies f o r f ood p roduc t ion , p as to ra l f auna l d epos i ts s hou ld b e d om ina ted b y r e la t ive ly y ounge r ma les a nd o lde r f ema les wh ich a re p re fe ren t ia l ly s l augh te red o ve r h ea l thy a nd p roduc t ive f ema les o f a l a ges ( Chang a nd K os te r 1 986 : 108 -109 ; D av is 1 983 :55 , 6 0 ; K le in a nd C ruz -Ur ibe 1 984 :86 -87 ; P ayne 1 972a :71 , 7 6 -77 , 1 973 :281-282 ; H esse 1 982 :412 ; L egge 1 972 :122 123 ) . T he l o g ic o ft h is a ssump t ion i st ha t r ega rd less o f whe the rp as to ra l managemen t i n vo lves mea t a nd /o rm i lk p roduc t ion , f ema les i ne i ther c ase c ons t i tu te t he p r ima ry p roduc t ive u n i t i n ah e rd , a nd y oung ma les s hou ld b e k i l led e i the ra s s oon a s t he r a t io o ff eed c onsumed t ow e igh t g a ined b eg ins t o r i se , o r a s s oon a s t he y i e ld o f m i lk f r om t he i r mo the rs i sn o t e ndangered . On ly af ew ma les wou ld n eed t o b e r e ta ined i n to o lde r a ges f o r b reed ing o r r i sk a ve rs ion p urposes . A l though s ome d i f fe rences i n t he a ge a t wh ich ma les a re p re fe ren t ia l ly s l augh te red a re t he re fo re e xpec ted b e tween mea t -on ly a nd mea t p l us m i lk s t ra teg ies ( Payne 1 973 :281-282 ) , i ti s s uspec ted t ha t t he a ge a nd s ex p ro f i les r esu l t ing f r om t hese d i f fe rences wou ld o n ly b e d i s t inc t ive when t he i mpor tance o f m i lk c omp le te ly o u twe ighs t ha t o f mea t ( Sebas t ian P ayne , p e rsona l c ommun ica t ion 1 987 ) . Wh i le t he e xac t n umbe r o f s e lec t ma les r e ta ined i n to o l de r a ges i se xpec ted t ov a ry b y s pec i f ic c i rcums tances , t h is g ene ra l p a t te rn o fs e lec t ive a ge -sex c u l l ing wou ld max im ize t o ta l h e rd g row th a nd p roduc t iv i ty . T h is a ssumed c u l l ing p a t te rn t he re fo re s eems q u i te r easonab le g i ven o bse rved p as tora l c ond i t ions u nde r e x tens ive h usband ry , whe re l o ng te rm p r ed ic t ions o ft he s pa t ia l a nd t empo ra l a va i lab i l i ty o fp as tu rage a re n ecessa r i ly l i m i ted , mass ive s t ock l o sses a re a lways p oss ib le , a nd h e rd a n ima ls c ons t i tu te t he p r ima ry means t op rov is ion o fsp r ing when t hey e s tab l ish n ew h ouseho lds ( Bar th 1 961:102 ; B radburd 1 982 ; B remen a nd d e W i t 1 983 :1346 ; D ah l a nd H jo r t 1 976 :129 ; D yson -Hudson a nd D yson -Hudson 1 969 :79 ; I n go Id 1 980 :89 , 1 34 , 1 78 , 2 01-202 , 2 06 -207 ; W ids t rand 1 975 :149 ) . I n g ene ra l a pp l ica t ion , h oweve r , t h is a ssumed p a t te rn o f s e lec t ive a ge -sex p as to ra l managemen t i s f r equen t ly r educed t o t he mo re e as i ly e s tab l ished p ropor t iona l s l augh ter p ro f i les o f i mma tu re /young v s . ma ture a n ima ls , w i th a ppa ren t " young k i l l " s l augh te r p rac t ices u sed t o i n fer h e rd managemen t s t ra teg ies ( Bököny i 1 969 ; D av is 1 983 :55 -56 ; K le in a nd C ruz U r ibe 1 984 :86 ; L egge 1 972 :121 , 1 22 ; P e rk ins 1 964 , 1 966 , 1 973 ; R odden 1 962 :272 ) , e spec ia l ly when a s h i f t t o s uch p a t te rns c an b e d emons t ra ted o ve r aw ide a rea d u r ing a r e la t ive ly s hor t p e r iod o ft ime ( Hesse 1 982 :40 1-4 10 ) . S ome d i rec t s uppo r t f o rt he c o r re la t ion o f a" young k i l l " s l augh ter p ro f i le w i th a c tua l p as to ra l h e rd managemen t s t ra teg ies a ppa ren t ly e x is ts i n ac o l lec t ion o f g oa t mand ib les f r om a round p as to ra l e ncampmen ts i nt he s ou the rn S ina i , whe re mos t we re o bserved t o b e f r om i n d iv idua ls b e tween 6 a nd 1 2 mon ths o f a ge ( Dav is 1 983 :56 ) . S im i la r ly , among f r agmen ts o fd omes t ic g oa t b ones c o l lec ted f r om H o t ten to t

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v i l lages , B ra in ( 1969 :16 17 ) f ound t ha t n ea r ly o ne -ha l f o ft he 6 4 a n ima ls m in ima l ly r ep resen ted w e re i mma ture . U n for tuna te ly , i nt he g ene ra l a bsence o fe xp l ic i t d a ta o n t he a ge a nd s ex p ro f i les o ff auna l d epos i ts c rea ted b y h un te r -ga the re r p opu la t ions , t he u se o f s uch p ro f i les t o i n fe r h uman managemen t s t ra teg ies r ema ins s pecu la t ive . E ven i fa c omp le te ( o r " adequa te " ) r ecovery o f z ooa rchaeo log ica l mor ta l i ty d a ta i sa ssumed a nd t he n ume rous p rob lems o f s amp le b i as a nd t aphonom ic p rocesses i g no red , t he i mp l ic i t a ssump t ion t ha t h un ter -ga the re r p opu la t ions wou ld n ecessa r i ly p roduce f auna l d epos i ts w i th mo re r andom a ge -sex p ro f i les t han t hose e xpec ted f r om p as to ra l is ts may b e s e r ious ly q ues t ioned ( Co l l ie r a nd Wh i te 1 976 ; H enry 1 975 ; J a rman a nd W i lk inson 1 972 ; P ayne 1 972a :77 ; S immons 1 975 -76 ; W i lk inson 1 976 ) . A s o n ly o ne e xamp le , L egge a nd R ow ley -Conwy ( 1987 :91) n o te t ha t when s ta lk ing ( r a the r t han d r iv ing ) g aze l le h erds , h un te rs p redom inan t ly s e lec t y ounge r a n ima ls i np r ime c ond i t ion , o f ten ma in ly ma les . H un te r -ga the re r p reda t ion o n s easona l a ge -sex h e rd s ubd iv is ions a nd o the r v a r ia t ions i nh un t ing me thods may t he re fo re p roduce f auna l d epos i ts w i th a ge a nd s ex p ro f i les ma tch ing t h ose e xpec ted u nde r h e rd managemen t s t ra teg ies . F ur the r , i th as f r equen t ly b een s ugges ted t ha t h e rd h usbandry t h rough s e lec t ive a ge -sex c u l l ing m igh t h ave b een p rac t iced p r io r t o t he e vo lu t ion o r a dop t ion o f p as to ra l s ubs is tence s t ra teg ies ( Banks 1 984 :2 10 ; J a rman 1 969 :256 -257 ; Moo re 1 985 :22 -23 ) . S uch p rac t ices m igh t a ccoun t f o r t he h i gh r a t io o f ma les t o f ema les ( up t o 1 0 :1) f ound b y S turdy ( 1975 ) i n h i s r eana lys is o ft he f auna ld epos i ts a tt he U ppe r P a leo l i th ic r e indee rk i l l s i te a tS te l lmoo r , Ge rmany , w h ich d a tes t oa round 1 3 ,000 B .C . H e rd managemen tt h rough s e lec t ive s l augh te r p rac t ices b y p reh is to r ic h un te rs h as a l so b een p os i ted f o r h e rds o f r ed d ee r ( M . J a rman 1 972 ) , g aze l le ( Legge 1 972 ; S axon 1 974 ; Moo re 1 985 :22 -23 ) , i b ex ( Tche rnov a nd B a r -Yose f 1 982 ) a nd g oa ts ( Bököny i 1 969 :225 , 1 973 :7 1) . Wh i le s eve ra l s cho la rs ( Co l l ie r a nd Wh i te 1 976 ; H en ry 1 975 ; H esse 1 982 , 1 984 ; S immons 1 975 -76 ) h ave s ugges ted t ha t n a tura l v a r ia t ions i nt he a ge -sex c ompos i t ion o fs easona l ly h un ted h e rds m igh t a l so s kew t he a ge -sex r a t ios a ppa ren t i n f auna l d epos i ts i n dependen t o f h e rd managemen t s t ra teg ies , t he i n d isc r im ina te s l augh ter o f h e rds d r iven d ur ing a nnua l m ig ra t ions s hou ld i n c lude a l a ge -sex c l asses p resen t d u r ing t he s eason o ft he d r ive ( Legge a nd R ow ley -Conwy 1 987 :9 1) . U n fo r tuna te ly , t he p reh is to r ic a nd e thnog raph ic e x ten t o f h e rd h usband ry a c t iv i t ies b y h un ter -ga the re rs ( i fs uch a c tua l ly o ccu r red ) , c anno t b e a dequa te ly a dd ressed o n t he b as is o f a va i lab le d a ta . A cco rd ing t o a N unam iu t i n forman t o f B in fo rd ( 1978 :235 -236 ) , f ema le r e indee r /ca r ibou w i th y oung w e re r e leased f r om t r ap c o r ra ls d ur ing d r ives . Ab edou in i n fo rman t i ns ou thwes te rn J o rdan s im i la r ly s t a ted t ha t d ur ing t he i n d iv idua l h un t ing o fg aze l le , t he k i l l ing o ff ema les w i th y oung was c ons ide red h aram , a n A rab ic t e rm g ene ra l ly u sed i nr e fe rence t o r e l ig ious a bom ina t ions ( pe rsona l o bserva t ion 1 986 ) . T he I n ca a re a l so s a id t o h ave p rac t iced s e lec t ive a ge -sex c u l l ing d u r ing t he i rd r ives o fw i ld g uanaco a nd v i cur ia h e rds ( Ga rc i lasso d e l a V ega 1 869 :116 -117 ) . S im i la r ly , n i ne teen th c en tury N ava jo a l so a ppea r t o h ave o ccas iona l ly p rac t iced s e lec t ive s l augh te r d ur ing a n te lope d r ives , a l though i n d isc r im ina te s l augh te r was s eem ing ly mo re c ommon ( K luckhohn e ta l . 1 97 1 :9 -10 ) . E ven s o , i twou ld s eem t ha t i n d isc r im ina te s l augh te r was mos t c ommon d ur ing g ame d r ives , w i th d r ives t empo ra l ly s tagge red i n s pec i f ic r eg ions i n o rde r t o a l low h erds t o d emog raph ica l ly r ecove r f r om s uch i n tense p reda t ion . I ti sa l so d i f f icu l t t o c oncep tua l ize h ow h un te rs wou ld b ene f i t f r om c onser va t ion p rac t ices d u r ing i n d iv idua l s ta lk ing o r g ame d r ives . P resumab ly , t he c onserva t ion o f h e rds t h rough s e lec t ive a ge -sex c u l l ing wou ld n o t b ene f i t a h un te r -ga the re r i ft he managed h e rd was s ub jec t t o p reda t ion b y o the r h un te r -ga the re rs . I ti s t he re fo re d i f f icu l t t o s ee h ow i n d iv idua ls w i th in a s oc ie ty wou ld b ene f i t f r om s uch f o rms o f c onserva t ion u n less o the r membe rs o f t he i r own a nd s u r round ing s oc ie t ies we re s im i la r ly c ons t ra ined . P o ten t ia l ly , i tmay b e a mo re e nergy e f ic ien t s t ra tegy t oe xc lude l a c ta t ing f ema les w i th y oung f r om s l augh te r r e la t ive t ot he c u l l ing o f n on lac ta t ing f ema les o r ma les i ng enera l . S easona l v a r ia t ions i nt he r e la t ive d es i rab i l i ty o fs pec i f ic a ge -sex c l asses m igh t a l so p l ay ar o le ( Spe th a nd S p ie lman 1 983 ) . N o f i na l r eso lu t ion o ft hese i s sues c an b e o f fe red a tt h is t ime , a nd f u r the r r esea rch i sc er ta in ly wa r ran ted . I nt he p resen t a na lys is , a l te rna t ive i n te rp re ta t ions o f r e levan t f auna l d epos i ts w i l l n ecessa r i ly b e e va lua ted w i th in t he b roade rt heo re t ica l c on tex t o f r e la t ive p as to ra l e ne rge t ics u nde r s pec i f ic e nv i ronmen ta l c ond i t ions . I ts hou ld a l so b e r emembe red t ha t i fc on t inued h e rd g row th a nd p roduc t iv i ty was a s d es i rab le a g oa l i ne a r ly p as to ra l s t ra teg ies a s i td emons t rab ly i si nt he e thnog raph ic p resen t , 1 39

e a r ly p as to ra l is ts s hou ld b e e xpec ted t o h ave t aken e ve ry r easonab le o ppor tun i ty t o o b ta in mea t f r om w i ld a n ima ls r a ther t han f r om t he i r s ubs is tence h e rds . Ac l ea r i mp l ica t ion o f t he mass ive g aze l le d r ives u nder taken b y e thnoh is to r ic b edou in p opu la t ions i n t he N ea r E as t ( Aha ron i 1 946 ; B u rkha r t 1 83 1:220 -22 1; Mus i l 1 928 :26 -27 ) i st ha t l a rge n umbe rs o fw i ld a n ima l b ones w ou ld p e r iod ica l ly e n te rt he a rchaeo log ica l r e co rd l e f t b y p as to ra l is ts . I ti ss uspec ted t ha t u nde r s uch c ond i t ions , f auna l d epos i ts c rea ted b y p as to ra l p opu la t ions w ou ld l o ok v e r y s im i la r t ot he f auna l d epos i ts c rea ted b y h un te r -ga the re r g roups . A s n o ted b y G i f fo rd e ta l . ( 1 98 1) , t he p o ten t ia l o f p as tora l p reda t ion o n w i ld a n ima ls wou ld s eem ing ly b e e ven g rea te r i n t he p reh is to r ic p as t . H ence , t he f r equenc ies a t wh ich p as to ra l h e rd s pec ies a re r ep resen ted a te a r ly p as to ra l s i tes may a c tua l ly b e i nverse ly r e la ted t o t he i r t r ue e conom ic s i gn i f icance . B y e x tens ion , e a r ly p as to ra l f auna l d epos i ts w i th l ow f r equenc ies o fo ne p o ten t ia l ly d omes t ica ted s pec ies r e la t ive t oo the rs may a c tua l ly r e f lec t t he e conom ic i mpor tance o ft he f o rme r r e la t ive t o o the r l e ss v a lued a nd mo re e xpendab le d omes t ica tes . H e re a ga in , a n e vo lu t iona ry a pp roach s eem ing ly h o lds g rea te r s ubs tan t ive i n te rp re t ive p o ten t ia l t han c on t inued s pecu la t ion d evo id o ft heo re t ica l d i rec t ion .

A rchaeo log ica l E xpec ta t ions B ased U pon t he R e la t ive E nerge t ics o fA l te rna t ive P as to ra l S t ra teg ies

T he R e la t ive R ank ing o fP as to ra lS t ra teg ies a nd t he A ccep ted T empora lS equence o fA n ima l D omes t ica t ion A s p rev ious ly d i scussed i nC hap te r 1 , af undamen ta l mode l d e r ived f r om o p t ima l f o rag ing t heory s ugges ts t ha t i n d iv idua l r esou rces c an b e r anked r e la t ive t o o ne a no the r a cco rd ing t o t he r e turn r a tes ( k ca l /h r ) f o r e xp lo i t ing t hem . R esources a re e xpec ted t o b e i n c luded o r e xc luded f r om t he d i e t a s af unc t ion o ft he i r r ank r e la t ive t oo the r a va i lab le r esou rces a nd t he e ncoun te r r a te ( dens i ty , a bundance ) o fh i ghe r r anked r e sou rces . A ssum ing t ha t c ame ls , c a t t le , s heep a nd g oa ts w e re e ach a va i lab le i ns u f f ic ien t n umbe rs t oe s tab l ish p as to ra l h e rds , t he e ar ly e vo lu t ion a nd a dop t ion o fp as to ra l s t ra teg ies s hou ld h ave i n vo lved a n i n i t ia l e conom ic d ec is ion c once rn ing wh ich o f t hese h e rd s pec ies w as mos t d es i rab le f o r p as to ra l e xp lo i ta t ion . I ft h is d ec is ion was b ased u pon r e la t ive e ne rge t ics , t he r ank ing o fp as to ra l s t ra teg ies b y t he i r r e spec t ive r e tu rn r a tes s hou ld p red ic t t he t empo ra l o rde r i nw h ich p as to ra l s t ra teg ies b ased u pon e ach s pec ies a ppea ri nt he a r chaeo log ica l r e co rd . I nC hap te r5 ,c ompa r isons b e tween t he e xpec ted c a lo r ic p roduc t iv i ty o f i n d iv idua l a n ima ls i n d ica ted t ha t c ame ls wou ld g enera l ly b e mo re p roduc t ive t han c a t t le , c a t t le mo re p roduc t ive t han s heep , a nd s heep mo re p roduc t ive t han g oa ts . I tw as s ubsequen t ly d emons t ra ted t ha t t h is s ame o rde r w ou ld h o ld f o r t he r e la t ive r ank ing o f p as to ra l s t ra teg ies , a s s hown b y t he r anges o fp o ten t ia l r e tu rn r a tes f o r mea t -on ly a nd mea t p l us m i lk s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts ( r e fe rt o F i gu res 9 ,1 a nd 1 2 ) . D epend ing u pon e xac t h e rd s i ze , i twas e s tab l ished t ha t u nder c ond i t ions o f e i the r mea t -on ly o r mea t p l us m i lk s t ra teg ies , p as to ra l s ubs is tence i n vo lv ing c ame ls o rc a t t le w ou ld g ene ra l ly e xh ib i t h i ghe r p o ten t ia l r e tu rn r a tes t han p as to ra l s ubs is tence i n vo lv ing s heep o r g oa ts . I fr anked o n t he b as is o f t he i r r e la t ive r e tu rn r a tes , c ame ls s hou ld g ene ra l ly b e p re fe r red f o r p as to ra l e xp lo i ta t ion o ve r c a t t le , c a t t le o ve r s heep , a nd s heep o ve rg oa ts . T h is r e la t ive r ank ing e f fec t ive ly i n ve r ts t he c ommon ly a ccep ted t empo ra l s equence f o r t he d omes t ica t ion o f t hese s pec ies . A cco rd ing t o c ur ren t o p in ion , s heep a nd g oa ts we re d omes t ica ted p r ior t o c a t t le , a nd a l t h ree s pec ies p r io r t oc ame ls ( Berge r a nd P ro tsch 1 973 : B ököny i 1 973 , 1 976 ; C lu t ton -Brock 1 981; C ompagnon i a nd T os i 1 978 ; F agan 1 986 :220 ; Gau t ier 1 980 ; P e rk ins 1 973 ; P ro tsch a nd B e rger 1 973 ; R eed 1 969 ; Z a r ins 1 978 ; Z eune r 1 963 ) . T h is s t r ik ing i n cons is tency i n v i tes b o th ar eexam ina t ion o ft he a ssump t ions emp loyed i n e s tab l ish ing t he r e la t ive e ne rge t ic r ank ings d e r ived f r om t h e d a ta p resen ted i nC hap te r5 ,a nd a c r i t ica l e va lua t ion o f t he a ssump t ions p rev ious ly emp loyed i na ccep ted i n te rp re ta t ions o ft he emp i r ica l d a ta r e levan tt oe a r ly a n ima l d omes t ica t ion .

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A ccoun t ing f o rt h eL a te D omes t ica t ion o fC ame ls : T he I mpo r tance o fE nv i ronmen ta lC ond i t ions a nd t h e P hy logeny o ft h eA n ima ls I n vo lved A s d i scussed i n C hap ter 4 , t he re a re ma jo r e nv i ronmen ta l d i f fe rences b e tween t he c ond i t ions f avo rab le f o r c ame l h usband ry a nd t hose f avo rab le f o r t he h usbandry o f c a t t le , s heep a nd g oa ts . I np a r t icu la r , i twas n o ted t ha t d ue t ot he i r r e la t ive p hy logeny , d romedary c ame ls w ou ld g ene ra l ly b e h i gh ly p roduc t ive i nt he r e la t ive ly h o t /d ry e nv i ronmen ts w he re c a t t le , s heep a nd g oa ts w ou ld f a i r p oo r ly , w h i le c a t t le , s heep a nd g oa ts w ou ld g ene ra l ly b e h i gh ly p roduc t ive i nt he r e la t ive ly w e t /co ld o rw e t /wa rm e nv i ronmen ts w he re c ame ls w ou ld f a i rp oo r ly . I ti st he re fo re e xpec ted t ha t s pa t ia l a nd t empo ra l v a r ia t ions i nt hese e nv i ronmen ta l c ond i t ions w ou ld d rama t ica l ly a l te r p red ic t ions b ased u pon c ompa r isons o f t he r e la t ive r e turn r a tes c a lcu la ted f o r p as to ra l s t ra teg ies i n vo lv ing d romeda ry c ame ls u nde r h o t /d ry c ond i t ions , a nd t hose i n vo lv ing c a t t le , s heep a nd g oa ts u nde r w e t /wa rm o r w e t /co ld c ond i t ions . H ence , i ti s s ugges ted t ha t t he l a te a ppea rance o fd romeda ry c ame l s t ra teg ies w ou ld o n ly make s ense i n t he s pa t ia l a nd t empo ra lc on tex to fe xpand ing d eser te nv i ronmen ts r e la t ive t op r io rw e t /wa rm o r w e t /co ld c ond i t ions . S ince c ame ls a ppa ren t ly w en t e x t inc t i nN o r th A f r ica n ea rt he e nd o ft he U ppe r P l e is tocene ( Bu l l ie t 1 975 ; G au th ie r -P i t te rs a nd D agg 1 98 1 ; G r igson 1 983 :312 ) , d a ta f o r t e s t ing t h is p ropos i t ion i si n i t ia l ly s ough to nt he A rab ian P en insu la . Wh i le t he e x is t ing p a leoc l ima t ic a nd a r chaeo log ica ld a ta r e levan tt ot e s t ing t h is p ropos i t ion c u r ren t ly a l low f o r o n ly t he b roades t o f g ene ra l iza t ions , t he eme rg ing r eco rd s eem ing ly s uppor ts t he e xpec ted s pa t ia l a nd t empo ra l c o r re la t ion b e tween d romeda ry c ame l d omes t ica t ion a nd e xpand ing d ese r t e nv i ronmen ts r e la t ive t o p r io r w e t /wa rm o r w e t /co ld c ond i t ions . U n l ike t he L a te P l e is tocene a nd H o locene p a leoenv i ronmen ta l r e co rd e s tab l ished f o rt he Z agros M oun ta ins o fw es tern I r an ( W r igh t 1 968 , 1 977 :282 -284 ; v an Z e is t 1 967 , 1 969 ) , r e sea rch o nt he A rab ian P en insu la ( Ga r ra rd a nd H a rvey 1 98 1 ; H ö tz l a nd Z ö t l 1 978 ; K a ise re ta l . 1 973 ; M cC lu re 1 97 1 , 1 976 , 1 978 ; S akaguch i 1 978 ) i n d ica tes L a te P le is tocene a nd H o locene e nv i ronmen ta l f l uc tua t ions m o re c l ose ly a pp rox ima t ing r o ugh ly c on tempo raneous v a r ia t ions i n t he p a leoenv i ronmen ta l r e co rds e s tab l ished f o r E as t a nd N o r th A f r ica ( Banks 1 984 :19 -59 ; B u tze r e ta l . 1 972 ; Gasse e ta l . 1 980 ; H am i l ton 1 982 ; M c In tosh a nd M c In tosh 1 981:602 -605 ; S t ree t a nd G rove 1 976 ; T a lbo t 1 980 ; Wendor f a nd H assan 1 980 ; Wendor f a nd S ch i ld 1 980a :234 -241) . D ur ing t he L a te P le is tocene , s ha l low w a rm wa te r l a kes w i th t empo ra l ly v a r iab le l e ve ls e x is ted i nt he R ub ' A l K ha l i o fs ou the rn S a 'ud i A rab ia ( McC lu re 1 976 ) , t he J ubba B as in o ft he G rea t N e fud s and s ea o fn o r the rn S a 'ud i A rab ia ( Ga r ra rd a nd H a rvey 1 981) , a nd t he D amascus a nd P a lmy ra B as ins o f mode rn S y r ia ( Ka ise re ta l . 1 973 ; S akaguch i 1 978 ) . D u r ing t h is p e r iod , much o ft he A rab ian P en insu la w as a ppa ren t ly c ove red b y s avanna g rass lands s uppo r t ing a f auna l b i omass s im i la rt ot ha to f mode rn E as t A f r ica . T h is s eem ing ly i n c luded c a t t le a nd b u f fa lo p opu la t ions ( Bos p r im igen ius a nd B uba lus ) a td r ie re nv i ronmen ta l e x t remes a nd H ippopo tamus a tc ompa ra t ive ly mo is ter e x t remes ( Ho lm 1 960 ; McC lure 1 97 1 , 1 978 ) . F rom a pp rox ima te ly 2 3 ,000 t o7 ,500 B .C . , i n c reas ing ly a r id c ond i t ions p reva i led ( Hö tz l a nd Z ö t l 1 978 :304 ) . I nt he R ub ' A l K ha l i , h ype ra r id i ty a ssoc ia ted w i th t he d i sappea rance o f l a kes a nd t he e x tens ive d epos i t ion o f r ed e o l ian s ands o ccur red f r om r o ugh ly 1 5 ,000 t o7 ,000 B .C . , a ppa ren t ly w i th a b r ie f ame l io ra t ion a round 1 3 ,000 t o9 ,000 B .C . a s r e f lec ted i nm in ima l a nd l ow ly ing p l aya l a ke b ed d epos i ts ( McC lu re 1 976 , 1 978 ) . F rom a pp rox ima te ly 7 , 500 t o 2 ,500 B .C . , T he A rab ian P en insu la e xpe r ienced a H o locene " s ubp luv ia l " w e t p hase marked b y t wo t o f ou r r e la t ive ly h um id p e r iods w i th i n c reased p rec ip i ta t ion a nd s t ronge r s ur face r uno f f i nw ad is ( 1 lö tz l a nd Z ö t l 1 978 :30 1-303 ; M cC lu re 1 978 ) . I nt he R ub ' A lK ha l i , t h is p hase i s ma rked b y as e r ies o fv a r iab le p l aya l a ke s t ands w i th a ssoc ia ted b ov id b ones a nd d a tes f r om r ough ly 7 , 000 t o 4 ,000 B .C . , a f te r w h ich t he p l ayas d i sappea red a nd mode rn h ype ra r id c ond i t ions s e t i n( McC lure 1 976 , 1 978 ) . I n ves t iga t ions i n to t he p a leoenv i ronmen t o f t he J ubba B as in i nn or thern S a 'ud i A rab ia ( Ga r ra rd a nd H a rvey 1 98 1) r e vea led ap a ra l le l e nv i ronmen ta l s equence w i th av i r tua l ly i d en t ica l r ad ioca rbon d a t ing t o McC lu re 's f i nd ings i nt h e R ub 'A lK ha l i ( 1 976 , 1 978 ) . A s n o ted b y Ga r ra rd a nd H a rvey ( 1981 :14 1) , t he p oo r ly d a ted l a ke s t and s equence i nt he a l -Ja f r B as in o fs ou thern J o rdan ( Huck r iede a nd W iesemann 1 968 ) w ou ld a l so s eem t of o l low t he s ame p a t te rn a s t ha t e s tab l ished i nt he J ubba B as in a nd t he R ub ' A l K ha l i . Wha t i sl a ck ing i s as equence o f r ad ioca rbon d a tes m in ima l ly e qu iva len t t o t he 2 8 d a tes o b ta ined b y M cC lure ( 1976 ) f o r l a ke s t ands i nt he R ub ' A lK ha l i . 1 41

l ti sa ga ins t t h is eme rg ing p a leoenv i ronmen ta l r eco rd f o r t he A rab ian P en insu la a nd i t s o ve ra l l c ong ruence w i th e x tens ive p a leoenv i ronmen ta l d a ta f r om E as t a nd N or th A f r ica t ha t t he maze o f c on f l ic t ing p a leoenv i ronmen ta l r econs t ruc t ions f o r t he c en t ra l a nd s ou the rn L evan t r ecen t ly s umma r ized b y H en ry ( 1983 :106 -116 ) mus t b e e va lua ted . T he g ene ra l e nv i ronmen ta l r econs t ruc t ion s ugges ted b y Moo re ( 1985 :8 -12 ) f o r t he l a te P le is tocene a nd H o locene o ft he L evan t , A na to l ia a nd c en t ra l Z ag ros mo re c l ose ly p a ra l le ls t ha t p resen ted h e re f o r r eg ions f u r the r s ou th . R ega rd less o fh ow i n cong ruen t c l ima t ic e ven ts may h ave b een b e tween d i f fe ren t r eg ions o f t he " Fe r t i le -Crescen t , " i ti ss t rong ly s uspec ted t ha t t he e vo lu t ion a nd a dop t ion o f f ood p roduc t ion s t ra teg ies i nt he N ea r E as t c anno t b e u nde rs tood i n dependen t o f t he a ppa ren t p os t -P le is tocene v a r iab i l i ty a nd e ven tua l d e te r io ra t ion o f e x tens ive s avanna e nv i ronmen ts o n t he A rab ian P en insu la . A s n o ted b y McC lu re ( 197 1 :47 ) : I ts hou ld b e k ep t i nm ind t ha t t he s o -ca l led " Fer t i le -C rescen t " p robab ly d oes n o t t ake o n a ny r ea l e co log ica l o r c u l tura l s i gn i f icance f r om t he d emog raph ic s tandpo in t u n t i l d es icca t ion s e ts i n a nd d r ives p opu la t ions t o i t . I tt hen b ecomes c on t ras t ing ly " f er t i le " c ompa red t ot he a reas i tb o rde rs . I ts eems p laus ib le t ha t t he i n i t ia l d e te r io ra t ion o f s avanna g rass land e nv i ronmen ts o n t he A rab ian P en insu la b e tween r ough ly 2 3 ,000 t o 7 ,500 B .C . wou ld h ave b een l i nked w i th d rama t ic s h i f ts i nh uman , p l an t a nd a n ima l d emog raphy , r e su l t ing i nd ec l in ing f o rag ing r e turn r a tes u l t ima te ly l e ad ing t o t he i n i t ia l e vo lu t ion a nd a dop t ion o f f ood p roduc t ion s t ra teg ies i n f avo rab le l o ca l i t ies o ft he r emnan t " Fer t i le -C rescen t " d u r ing t he n i n th a nd e i gh th m i l lenn ia B .C . Wh i le t h is s cena r io w i l l b ef u r the rd eve loped i nt he s ubsequen t a na lys is o fc e rea l r e tu rn r a tes a s at en ta t ive e ne rge t ic mode l f o rt he f o rage r / food p roduce rt r ans i t ion i nt he N ea r E as t a nd N or th A f r ica , i ti st he p o ten t ia l c o r re la t ion b e tween c hang ing e nv i ronmen ta l c ond i t ions a nd t he l a te d omes t ica t ion o fc ame ls w h ich i so fi mmed ia te c once rn h e re . I fd romeda ry c ame ls h ad b een af o cus o fa n ima l h usband ry e fo r ts a s e a r ly a st he n i n th a nd e igh th m i l lenn ia B .C . ( no e v idence o f wh ich i sc ur ren t ly k nown ) , t hey s hou ld h ave b een d ropped i nf avo r o f c a t t le , s heep a nd g oa ts d u r ing t he s ubsequen t w e t /co ld o r w e t /wa rm H o locene " subp luv ia l " p hase b e tween a pp rox ima te ly 7 ,500 a nd 2 ,500 B .C . I n N or th A f r ica , h ump less c a t t le , a nd e ven tua l ly s heep a nd g oa ts , w e re c e r ta in ly t he e conom ic f ocus o fp as to ra l p opu la t ions d u r ing t he H o locene we t p hase , a s r e f lec ted i nb o th a rchaeo log ica l r ema ins a nd c on temporaneous r ock a r t ( Banks 1 984 ; C la rk 1 980 ; E ps te in 1 97 1:213 -221 ; Mc In tosh a nd Mc In tosh 1 981 ; A . Sm i th 1 980a , 1 980b ; Wendor f a nd H assan 1 980 ; Wendo r f a nd S ch i ld 1 980a ) . I n S a 'ud i A rab ia , r ock a r t a l so b e l ieved t o d a te t o t he Ho locene we t p hase p redom inan t ly d ep ic ts h ump less c a t t le a nd g oa ts , w i th c ame ls a nd o r yx c onsp icuous ly a bsen t o r r a re ( Ga r ra rd a nd H a rvey 1 981 :142 ; Z a r ins 1 978 :44 , 1 982 :25 ) . Wha t a re b e l ieved t o b e p as tora l s i tes l o ca ted a l ong t he f o rme r s ho re l ines o fH o locene l a kes o n t he A rab ian P en insu la f r equen t ly c on ta in c a t t le b ones , a nd Z a r ins ( 1982 :25 , 2 7 ) h as s ugges ted t ha t b y a tl e as t 5 ,000 B .C . , t he h un t ing o fc a t t le h ad b een r ep laced b y c a t t le h e rd ing . H oweve r , a n e ven e a r l ie rd a te f o rt he e vo lu t ion o fc a t t le p as to ra l s t ra teg ies o nt he A rab ian P en insu la s eems p robab le . P resumab ly , c a t t le wou ld h ave b een t he p r inc ip le h e rd a n ima l o ft he manya t ta b u i ld ing p as to ra l is ts o f N or th A rab ia i nt he e i gh th o r s even th m i l lenn ia B .C . ( Be t ts 1 982a , 1 982b :31 , 1 984 ; H e lms 1 98 1:50 ; I n g raham e ta l . 1 98 1 :67 ; B e t ts , i nR o l le r 1 984 :220 ) , ap ropos i t ion wh ich c an e ven tua l ly b e t es ted o nce t he a rchaeo logy o ft hese s i tes moves b eyond t he me t icu lous d raw ing o f s ur face l i t h ic ma te r ia ls . J udg ing f r om t he f auna l r ema ins r ecove red a tt he N eo l i th ic s i te o f ' A in Ghaza l i nn o r th -cen t ra l J o rdan ( Ro l le fson 1 983 , 1 984 , 1 985 , 1 986 ; R o l le fson a nd S immons 1 986 ) , t he C ha lco l i th ic s i te o fJ awa i nn o r theas te rn J o rdan ( He lms 1 981 , 1 982 ) a nd C ha lco l i th ic s i tes i nt he J uday id B as in o f s ou the rn J o rdan ( Henry 1 982 ) , c a t t le , s heep a nd g oa ts w e re p robab ly t he p r inc ip le d omes t ica tes h e rded i nN o r th A rab ia t h rough a t l e as t t he f our th m i l lenn ium B .C . I ti st he re fo re s uspec ted t ha t f u tu re f auna l r esea rch a tp as to ra l s i tes o f t he H o locene w e t p hase o n t he A rab ian P en insu la w i l l f u r the r c on f i rm p a ra l le l s ubs is tence p a t te rns i nt h is r eg ion t ot hose a l ready e s tab l ished f o r c on tempo raneous p as to ra l p opu la t ions i nN o r th A f r ica . N o t u n t i lt he t h i rd m i l lenn ium B .C . d oc ame l r ema ins a nd r e l ie f r ep resen ta t ions a tt he s i te o f Umm a n -Na ro nt he c oas to fA bu D hab i ( F r i fe l t 1 979 :574 ; H och 1 977 ) a nd c ame l r ema ins a tt he s i te o fH i l i o n t he Oman P en insu la ( Tos i 1 974 :162 ) s ugges t t he p robab le d omes t ica t ion o ft he 1 42

d romeda ry c ame l o nt he A rab ian P en insu la . F u r the ra f ie ld , l a te f ou r th a nd t h i rd m i l lenn ia B .C . s i tes i n E gyp t h ave p roduced t he e a r l ies t k nown f i gu r ines d ep ic t ing d romedary c ame ls ( McC lure 1 97 1 :49 , 6 4 ; R ip insky 1 974 , 1 975 :297 ; Z eune r 1 963 :350 ) , w h i le r ope made f r om c ame l h a i rw as f o und i na n e a r ly t h i rd m i l lenn ium g ypsum q ua r ry i nt he n o r the rn F a iyum s ca rp ( Ca ton -Thompson 1 934 ) . Wh i le p resumab ly r e la t ing t ot he c o ld /d ry a dap ted B ac t r ian c ame l , t he e a r l ies t f i nds o fc ame l b ones , a r t is t ic r ep resen ta t ions o fc ame ls , a nd c ame l h a i r a nd d ung i n e as tern I r an a nd s ou the rn T urkmen is tan o ccur i n m id four th t h rough t h i rd m i l lenn ia B .C . c on tex ts a t s eve ra l s i tes , s ugges t ing a n i n c reas ing a ssoc ia t ion w i th h umans i n terp re ted a s r e f lec t ing a no ngo ing p rocess o fc ame l d omes t ica t ion d u r ing t h is p e r iod ( Compagnon i a nd T os i 1 978 ) . A l though Z a r ins ( 1978 ) h as c o r rec t ly c au t ioned t ha t t hese s ca t te red d a ta d o n o t p rov ide " f i rm e v idence "f o rt he d omes t ica t ion o ft he c ame l a s e a r ly a st he f our th t ot h i rd m i l lenn ia B .C . , t he s imu l taneous a ppea rance d u r ing t h is p e r iod o fa n i n c reas ing a ssoc ia t ion b e tween h umans i ns e t t led c ommun i t ies a nd c ame ls s eems i n dependen t ly c on f i rmed i nn or the rn E gyp t , t he A rab ian P en insu la , a nd e as te rn I r an a nd s ou the rn T urkmen is tan . I n t he a bsence o f a r chaeo log ica ld a ta f r om p as to ra l s i tes o ft h is p e r iod o nt he A rab ian P en insu la a nd t he l a ck o fa s u i tab le e xp lana t ion f o rw hy s e t t led c ommun i t ies w ou ld b ecome i n c reas ing ly i n vo lved w i th w i ld c ame l p opu la t ions a t t h is t i me , t he a u tho r f o l lows C ompagnon i a nd T os i ( 1978 ) , R ip insky ( 1975 :297 ) a nd Z eune r ( 1963 :364 ) i ns ugges t ing t ha t c ame l d omes t ica t ion w as i n i t ia ted b y a t l e as t t he f ou r th a nd t h i rd m i l lenn ia B .C . Wh i le f u r the r d a ta a re c er ta in ly r equ i red , ac l ose t empo ra l a ssoc ia t ion b e tween t he d omes t ica t ion o f t he c ame l a nd t he e xpans ion o f d eser t e nv i ronmen ts s ubsequen t t o p r io r w e t /wa rm o r w e t /co ld c ond i t ions d ur ing t he H o locene w e t p hase o n t he A rab ian P en insu la s eems h i gh ly p robab le .

We re C a t t le R ea l ly D omes t ica ted L a te rt h an S heep a nd G oa ts? A l though e nv ironmen ta l s h i f ts wou ld s eem ing ly a ccoun t f o r w hy t he c ame l was d omes t ica ted l a te rt han c a t t le , s heep o rg oa ts , as im i la re xp lana t ion f o rt he c ommon ly a ccep ted l a te a ppea rance o fd omes t ica ted c a t t le r e la t ive t os ma l ls tock s eems u n l ike ly . A s d i scussed i n C hap te r 4 , wh i le c a t t le a re i n deed e co log ica l ly mo re d emand ing t han s heep a nd g oa ts d ue p r ima r i ly t o t he i r g rea te r w a te r r equ i remen ts , t h is d i s t inc t ion i sh a rd ly a s d rama t ic a s t he n ume rous d i f fe rences o bse rved b e tween t he e co log ica l r equ i remen ts o fc ame ls a nd t hose o f t he B ov idae i ng ene ra l . F u r the r , a s p rev ious ly n o ted i nC hap te r2 ,p reh is to r ic c a t t le p opu la t ions s eem ing ly o ccup ied n ume rous h ab i ta ts f r om A s ia t h rough E urope , a nd f r om t he E u ras ian t und ra t ot he I n d ian S ubcon t inen t a nd N o r th A f r ica . T he i r w idesp read d i s t r ibu t ion i nt he N ea r E as tf r om t he U ppe rP a leo i i th ic i sw e l l -a t tes ted a ta rchaeo log ica l s i tes . Wh i le c ame l r ema ins a r e s t r ik ing ly a bsen t i n mos tc ases , c a t t le b ones o ccur i nt he f auna l d epos i ts o ft he K eba ran s i te o f Wad i Madamagh a nd P re -Po t te ry N eo l i th ic B B e idha i ns ou the rn J o rdan ( Pe rk ins 1 966 ) , A u r ignac ian t h rough N a tu f ian l e ve ls a tK eba rah C ave ( Saxon 1 974 ) , t he N a tu f ian l e ve l a tH ayon im T e r race ( Hen ry e ta l . 1 981) , Meso l i th ic t h rough P re -Po t tery N eo l i th ic l e ve ls a tJ e r icho ( C lu t ton -B rock 1 979 :147 ) , Meso l i th ic a nd N eo l i th ic T e l l A bu H urey ra ( Ducos 1 978 ; L egge a nd R ow ley -Conwy 1 987 ; Moo re 1 985 ) a nd N aha l O ren ( Noy e ta l . 1 973 ) , t he P re -Po t tery N eo l i th ic B s i tes o f ' A in G haza l i nJ o rdan ( Ro l le fson 1 983 , 1 984 , 1 985 , 1 986 ; R o l le fson a nd S immons 1 986 ) , N aha ! D ivshon i nt he N egev ( Se rve l lo 1 976 ; T chernov 1 976 ) a nd Wad i T be ik i nt he s ou the rn S ina i ( Ba r Y ose f 1 984 ; T che rnov a nd B a r -Yose f 1 982 ) , A ce ram ic N eo l i th ic S ube rde ( Pe rk ins 1 969 ; P e rk ins a nd D a ly 1 968 ) , C an H asan I l , H ac i la ra nd A e ik l i H üyük ( Payne 1 972 , 1 985 ) , A ce ram ic N eo l i th ic J a rmo ( P ro tsch a nd B e rge r 1 973 ; R eed 1 969 ) a nd T epe Gan j D a reh ( Hesse 1 984 ) , a nd v a r ious s i tes i nt he D eh L u ran P la in p r io r t o 5 ,500 B .C . ( Ho le e ta l . 1 969 ) . I tw ou ld s eem t ha t c a t t le p opu la t ions w e re b o th p resen t a nd w ide ly e xp lo i ted t h roughou t t he N ea r E as t d ur ing t he E p ipa leo l i th ic a nd N eo l i th ic p e r iods . Wh i le t hese c a t t le p opu la t ions w e re p resumab ly " t e the red " t o r eg ions w i th r e la t ive ly mo re r e l iab le w a te rs ou rces ,t hey w e re c e r ta in ly a va i lab le f o rp as to ra le xp lo i ta t ion i n many o ft h e s ame e nv i ronmen ta l l o ca l i t ies w he re e a r ly s heep a nd g oa t d omes t ica t ion a re p os i ted . M in ima l ly , i f c a t t le w ere n o t d omes t ica ted i nt he N ea r E as t p r io r t o s heep a nd g oa ts , t hey s hou ld a t l e as t h ave b een d omes t ica ted c on tempo raneous ly w i th t hem .

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I fe nv i ronmen ta l f ac to rs c anno t a ccoun tf o r al a te d omes t ica t ion o fc a t t le r e la t ive t os heep a nd g oa ts , a nd i f ,a s p rev ious ly d i scussed i nC hap te r 2 ,t he a ppea l t o b ehav io ra l a nd s i ze v a r ia t ions a s r e la ted t o t he h uman a b i l i ty t o e f fec t ive ly c on t ro l h erd a n ima ls wou ld a l so b e i n su f f ic ien t t o a ccoun t f o r t h is s i tua t ion , t hen t he c r i te r ia b y wh ich t he l a te d omes t ica t ion o f c a t t le r e la t ive t o sma l ls tock h as b een p os i ted mus t b e q ues t ioned . I n i t ia l ly , i th as b een s ugges ted t ha t t he e a r l ies t e v idence f o r t he d omes t ica t ion o f s heep a nd g oa ts d a tes r espec t ive ly t o t he m id -n in th a nd e a r ly e i gh th m i l lenn ia B .C . ( Be rge r a nd P ro tsch 1 973 :221; B ököny i 1 969 :222 , 1 973 :69 , 7 3 -74 , 1 976 , 1 977 ; P e rk ins 1 964 , 1 973 ; P ro tsch a nd B e rger 1 973 :238 -239 ) , w h i le t he e a r l ies t e v idence f o r c a t t le d omes t ica t ion h as b een p rev ious ly d a ted t o t he e a r ly t o m id -seven th m i l lenn ium B .C . ( Be rge r a nd P ro tsch 1 973 :74 ; B ököny i 1 976 ; Gau t ie r 1 980 :332 , 3 36 ; M i lo jc ic e ta l . 1 962 ; P erk ins 1 969 ; P ro tsch a nd B e rge r 1 973 :238 ) . T he p r ima ry c r i te r ia b y w h ich t he d omes t ica ted s t a tus o ft hese s pec ies h as b een d e te rm ined i n vo lve c hanges i nt he morpho logy o fh o rn c o res a nd a ve rage s i ze a nd s i ze r anges , a nd e v idence f o r s e lec t ive a ge o r a ge -sex c u l l ing a s p resumab ly r e f lec t ing h usband ry p rac t ices . A s p rev ious ly d i scussed a bove a nd i nC hap te r2 ,t he a c tua l u t i l i ty o fs uch c r i te r ia f o r i d en t i fy ing p a leoeconom ic a c t iv i t ies ( hun t ing v s . h e rd ing ) i so f ten o pen t oq ues t ion . F o r e xamp le , t he i d en t i f ica t ion o fd omes t ic s heep a tZ aw i C hem i S han ida r , r ep resen t ing t he o n ly e v idence wh ich c an b e u sed t o s uppo r t t he h e rd ing a nd d omes t ica t ion o f s heep a s e a r ly a s t he m id -n in th m i l lenn ium B .C . , p r ima r i ly r es ts u pon a n i n c rease i nt he p ropor t iona l c on t r ibu t ion o fs heep t ot he f auna l d epos i t t h rough t ime a nd t he h i gh p e rcen tage o f i mma tu re s heep i nt he s amp le ( Pe rk ins 1 964 , 1 973 ) . Mo rpho log ica l ly , t he s heep we re s t i l l " w i ld . " S eem ing ly i n r esponse t o c r i t ic ism o f t h is i n terp re ta t ion i n l i gh t o f e qua l ly p laus ib le a l te rna t ives wh ich d o n o t i mp ly a h erd ing p a leoeconomy , f u r the r s uppor t f o rt he d omes t ica ted s t a tus o f t he s heep a tZ aw i C hem i was s ough t i no bse rved v a r ia t ions b e tween t he c hem ica l c ompos i t ion a nd m ic ros t ruc ture o f t he b ones o f a rchaeo log ica l ly d e f ined " w i ld " a nd " domes t ica ted " s heep a nd g oa ts ( D rew e ta l . 1 97 1 ; P erk ins 1 973 ) . H oweve r , a t temp ts t ov e r i fy s uch s t ruc tura l d i f fe rences u s ing b ones f r om mode rn w i ld a nd d omes t ic s heep p opu la t ions i n d ica te t ha t n o s i gn i f ican t d i f fe rences e x is t b e tween t hese g roups i n t he i r c hem ica l c ompos i t ion o r m ic ros t ruc tu re ( Zede r 1 978 ) . T hus , wh i le t he Z aw i C hem i f auna l d epos i ts may r e f lec t e i the r s e lec t ive s l augh te r p rac t ices o n t he b as is o fa ge o r p reda t ion o n s easona l h e rd s ubd iv is ions ( nurse ry h e rds ) , t hey d o n o t n ecessa r i ly r e f lec t t he d omes t ica t ion o fs heep o r t he e x is tence o f ap as to ra l e conomy . E xc lud ing t he d ub ious s ugges t ion t ha t t he l i m i ted s heep b one s amp le a t T epe A s iab i n wes te rn I r an , c on ta in ing 8 0% a du l ts ( a l l ma le ) , r e f lec ts " con t ro l led h un t ing wh ich m igh t b e c onnec ted w i th d omes t ica t ion " i n t he e a r ly e i gh th m i l lenn ium B .C . ( Bököny i 1 973 :74 ; 1 976 :2 1) , t he a va i lab le d a ta s eem ing ly s ugges t t ha t am id - t ol a te e i gh th m i l lenn ium B .C . d a te may b e mo re a pp rop r ia te f o rt he i n i t ia l a ppea rance o fs heep h e rd ing s t ra teg ies . On t he b as is o f as i ng le h o rn less f ema le d i scove red i nt he l owe r l e ve ls a tA l i K osh , i th as b een s ugges ted t ha t d omes t ic s heep we re p resen t i nt he D eh L uran P la in b yt he m id - t ol a te e i gh th m i l lenn ium B .C . ( Ho le 1 969 :278 -281) , wh i le d omes t ic s heep h ave a l so b een r epo r ted a t Qayönü T epes i i n s ou thwes te rn A na to l ia a round 7 ,000 B .C . ( Reed 1 969 :37 1) . A m id -e igh th m i l lenn ium d a te w ou ld a l so s eem a pp rop r ia te f o rt he i n i t ia l a ppea rance o fg oa t h e rd ing s t ra teg ies . T he e a r ly e i gh th m i l lenn ium B .C . d a te f o rt he d omes t ica t ion o fg oa ts r e s ts s o le ly u pon t he f auna l r ema ins e xcava ted a tT epe A s iab i nt he Mah idash t V a l ley o f wes te rn I r an ( Bököny i 1 973 , 1 976 , 1 977 ) . I nt he a bsence o fo the r mo rpho log ica l c r i te r ia , t he d omes t ica ted s t a tus o ft hese g oa ts was i n fe r red f r om t he d i scovery o ft h ree h o rn c o res e xh ib i t ing a s l igh t ly t w is ted s hape w i th f l a t c ross -sec t ions . Wh i le t hese t h ree s pec imens o n ly r ep resen ted a sma l l p ropor t ion o f t he t o ta l h orn c o re i n ven to ry o ft he s i te ( app rox ima te ly 7 -8% ) , t he i n t ia l s t ages o fd omes t ica t ion we re i n fe r red ( Bököny i 1 973 :7 1) . H oweve r , B ököny i h imse l f n o ted t ha t t w is ted h o rns a re k nown t o a ppea r i nC aucas ian w i ld g oa t p opu la t ions a t l ow f r equenc ies ( 1973 :70 ) , a nd a s s ugges ted b y H esse i nr e fe rence t og oa t h o rn c o res ( 1984 :247 ) , " we d o n o t y e t k now e nough a bou t t he v a r ia t ion i nw i ld g oa t mo rpho logy t ob e c er ta in t ha t t he s l igh t d eg ree o f d i ve rgence f r om t he e xpec ted p a t te rn i ss i gn i f ican t . " F ur the r s uppor t f o rt he d omes t ica ted s t a tus o fs ome o ft he g oa ts a tA s iab was s ough t i n t he a ge -sex r a t io a ppa ren t i nt he d epos i t . S ince a pp rox ima te ly 8 0% o ft he g oa t r ema ins w e re f r om a du l t a nd ma ture a n ima ls ( 95% ma les ) , i tw as s ugges ted t ha t y oung i n d iv idua ls w e re b e ing 1 44

k ep t a nd t amed , wh i le t he w i lde r a du l ts ( p r ima r i ly ma les ) w ere b e ing s l augh te red ( Bököny i 1 973 :7 1) . A s n o ted b y H esse ( 1982 :4 13 , 1 984 :259 ) , a mo re p robab le e xp lana t ion f o r t he A s iab g oa t b one d epos i t w ou ld b e t he e xp lo i ta t ion o f s easona l ma le g oa t b ands b y h un te rs . T h rough amo re c r i t ica l e va lua t ion o ft he f auna ld a ta f r om s eve ra l s i tes i nt he Mah idash t V a l ley , H esse ( 1984 ) h as a l te rna t ive ly s ugges ted t ha t t he e vo lu t ion o f ag oa t h e rd ing s ubs is tence i s n o ta ppa ren t i nt he a rchaeo log ica l r e co rd o ft he r eg ion u r n i lt he m idd le o ft he e i gh th m i l lenn ium B .C . T he mo re c onv inc ing e v idence u sed t os uppor t t h is i n te rp re ta t ion i n c ludes r eg iona l m id e i gh th m i l lenn ium s h i f ts i nt he a ge -sex r a t io o f g oa t f auna l d epos i ts f r om v a r ious s i tes a s p o ten t ia l ly r e f lec t ing t he a dop t ion o f s e lec t ive a ge -sex c u l l ing s t ra teg ies c ons is ten t w i th e xpec ted p as to ra l p rac t ices , a nd c on tempo raneous c ap r ine f oo t p r in ts o n s ome o ft he mud b r icks a tT epe G an j D a reh ,t h e l a t te rr easonab ly a ssumed a s r e f lec t ing d omes t ic a n ima ls o nt he s i te wh i le b r icks w e re b e ing d r ied ( Hesse 1 984 ) . A ga in , w i th t he e xcep t ion o f af ew med ia l ly f l a t tened g oa t h o rn c o res , at r a i to fq ues t ionab le s i gn i f icance i nl ow f r equenc ies , t hese a n ima ls w e re s t i l l mo rpho log ica l ly " w i ld . " S im i la rt ot h e a ppa ren t i n f la t ion o fe a r ly d a tes f o rt he d omes t ica t ion o fs heep a nd g oa ts i s t he p robab le d ep ress ion o fd a tes f o r t he d omes t ica t ion o fc a t t le . Wh i le s ome s cho la rs ( e .g . , J a rman 1 969 :240 ) h ave l o ng b een c r i t ica l o ft he p resumed i d en t i f ica t ion o f c a t t le h un t ing o r h e rd ing e conom ies t h rough t he u se o f mean s i ze a nd s i ze r anges t od i v ide c a t t le i n to " w i ld " a nd " domes t ic " c a tego r ies , s i ze p a rame te rs r ema in t he p r ima ry c r i te r ia b y w h ich e a r ly c a t t le p a leoeconom ics a re i n fe r red . A s e xp l ic i t ly d i scussed b y B anks ( 1984 :203 -204 , 2 31) , s uch o s teome t r ic a na lyses , a tb es t , w ou ld o n ly s e rve t oi d en t i fy t he mo rpho log ica l d imens ion o ft he d omes t ica t ion p rocess , w h i le t he b ehav io ra l d imens ion o fd omes t ica t ion wou ld h ave a l ready o ccur red o ve r ap r io r i n de te rm ina te p e r iod t h rough h usbandry a nd h e rd ing ( a lso P ayne 1 985 :114 ) . A s n o ted i nC hap te r2 ,i fmo rpho log ica lc r i te r ia m us t n ecessa r i ly b ed emons t ra ted i n o rde rt od ocumen tt ha td omes t ica t ion h as o ccu r red o ri so ccu r r ing , t he re w ou ld b e n o e v idence t ha t c ame ls w e re e ve rd omes t ica ted . I twas a l so n o ted a tt ha tt ime t ha t i nt he i n te rp re ta t ion o f e a r ly c a t t le p a leoeconom ies o nt he b as is o fs i ze r educ t ions , a ppa ren t r eg iona l v a r ia t ions i nt he s i ze r anges o fp reh is to r ic c a t t le p opu la t ions d im in ish t he r e l iab i l i ty o fs uch o s teome t r ic c r i te r ia ( J a rman 1 969 ;J a rman a nd W i lk inson 1 972 :88 -90 ; P ayne 1 972a ) . H oweve r , i fi ti sa ssumed t ha t o s teome tr ic e v idence f o r d ec reased s i ze d oes r e f lec t t he e nd s tage o f i n i t ia l c a t t le d omes t ica t ion , t hen t he i re a r ly h usband ry a nd h e rd ing m us t n ecessa r i ly p reda te t ha t e v idence . H oweve r , u n l ike t he s i tua t ion w i th s heep a nd g oa t r ema ins , a ge -sex s l augh te r s chedu les f o r t h e c a t t le c omponen t o fr e levan t f auna l d epos i ts h ave r a re ly b een e xam ined , t hus p rec lud ing a n e qu iva len t e va lua t ion o ft h ec a t t le d omes t ica t ion p rocess . I n E urope , t he e a r l ies t r ad ioca rbon d a tes a ssoc ia ted w i th mo rpho log ica l ly d e f ined d omes t ic c a t t le c ome f r om t h e s i tes o f A rg issa -Magu la a nd N ea N ikomede ia i nG reece , a nd c l us te ra round 7 , 000 t o6 , 900 B .C . ( Be rge ra nd P ro tsch 1 973 ; M i lo je i6 e ta l . 1 962 ; P ro tsch a nd B e rge r 1 973 ; R odden 1 962 ) . T he e a r l ies t d omes t ica ted c a t t le i nt he N ea r E as t h ave b een a ssumed t ob et h ose o fQ a ta l H üyük i nA na to l ia ( Pe rk ins 1 969 ) . H e re , c ompa r isons o ft he d i s ta l w id th o f1 2 c a t t le h ume r if r om l e ve ls d a t ing t oa round 6 ,400 B .C . h ad amean s i ze o f1 02 .0 mm a nd ar ange f r om 8 5 .3 t o1 08 .0 mm ,w h i le 3 3 c a t t le h ume r if r om al e ve ld a t ing t oa pp rox ima te ly 5 , 800 B .C . h ad amean s i ze o f8 6 .3 mm a nd ar ange f r om 6 3 .0 t o1 05 .0 mm . G iven t he a ppa ren t s i ze r educ t ion b e tween t hese d epos i ts , t he c a t t le o f5 ,800 B .C . w e re i d en t i f ied a s " cer ta in ly " d omes t ic . H oweve r , t hose o f 6 ,400 B .C . w e re i d en t i f ied a s " p robab ly " d omes t ic , p r ima r i ly b ecause t he h i gh p e rcen tage o fl i mb b one f r agmen ts i nt hese e a r l ie rd epos i ts " s t rong ly i mp l ies t ha t t hey we re s l augh te red c l ose b y " ( Perk ins 1 969 ) . A cco rd ing t o P e rk ins ( 1973 :280 ) , t he s ma l l s i ze o ft he c a t t le s pec imens r e cove red f r om A ce ram ic H ac i la r may r e f lec t mo rpho log ica l ly d omes t ica ted c a t t le a tt h is s i te a se a r ly a s7 , 000 B .C . , a l though t h is i su ncer ta in d ue t ot he sma l l s amp le i n vo lved . R ega rd less , i ft he s ame b ehav io ra l c r i te r ia emp loyed a t Q a ta l H üyük t o i n fe r " p robab ly " d omes t ica ted c a t t le a t6 , 400 B .C . h ad b een a pp l ied t ot he c a t t le r ema ins f r om t he D eh L u ran P la in i ns ou thwes te rn I r an ( Ho le e ta l . 1 969 :298 -308 ) , t he h i gh f r equency o f a r t icu la ted l i mb b ones i nl e ve ls p reda t ing t h e 5 ,500 t o5 ,000 B .C . a ppea rance o fr e la t ive ly sma l le r , a nd h ence " domes t ica ted " c a t t le , w ou ld h ave s ugges ted a l o ng p reced ing p e r iod o f " i nc ip ien t c a t t le d omes t ica t ion " s t re tch ing b ack t o t he m id -e igh th m i l lenn ium B .C . H oweve r , t he D eh L u ran c a t t le r ema ins p reda t ing 5 ,500 B .C . w e re a l i d en t i f ied a s " w i ld , " b ased u pon s i ze c ompa r isons w i th p resumab ly w i ld c a t t le r ema ins f r om E ng l ish a nd D an ish Meso l i th ic s i tes . I nl i gh t o fk nown r e g iona l v a r ia t ions i nt he s i ze o fp os t -P le is tocene c a t t le p opu la t ions , J a rman ( 1969 :239 ) n o ted

1 45

t ha t " As i ze r e la t ionsh ip b e tween E ng l ish a nd D an ish M eso l i th ic c a t t le a nd N eo l i th ic c a t t le f r om I r an n eed n o tb e r e levan tt o ad i scuss ion o fs i ze c hanges i ne i the ra rea . " A ppa ren t ly a wa re o ft he d ub ious n a tu re o ft he o s teome t r ic c r i te r ia c ommon ly u sed i nt he p a leoeconom ic i n te rp re ta t ion o f c a t t le r ema ins , C lu t ton -B rock ( 1979 :155 ) c au t ious ly s ugges ted t ha t e ven t hough t he c a t t le o f P re -Po t te ry N eo l i th ic J e r icho a ppea r t o b e mo rpho log ica l ly " w i ld "d ue t ot he i rs i ze ,t hey m igh tw e l l h ave b een a tt he e a r ly s t ages o ft am ing a nd c on t ro l i nt he s even th m i l lenn ium B .C . H e re , t he p oss ib i l i ty o f c on temporaneous c a t t le , s heep a nd g oa t d omes t ica t ion h as a t l e as t b een p os i ted . S im i la r ly , D ucos ( 1978 ) h as s ugges ted t ha tw h i le t he c a t t le r ema ins f r om M u reyb i ta re among t he l a rges t k nown i nt h e N ea r E as t , a ge d i s t r ibu t ion d a ta a nd a n i n c rease i nt he i r r e la t ive a bundance may r e f lec t t h e e a r ly s t ages o ft he i r d omes t ica t ion b y t he m id -seven th m i l lenn ium , r ough ly c on tempo raneous w i th t he p roposed d omes t ica t ion o f s heep a nd g oa ts a tt he s i te ( a lso L egge a nd R ow ley -Conwy 1 987 :95 ; Moo re 1 985 :23 ) . A t P re -Po t te ry N eo l i th ic B ' A in G haza l ( c . 7 ,250 -6 ,200 B .C . ) , a r e la t ive ly h i gh d eg ree o fp ha lang ia l p a tho logy , s l igh t i n d ica t ions o fr e la t ive s i ze r educ t ion , a nd c l ay f i gu r ines o fc a t t le w i th i mp ress ions o ft w is ted f i b res t h a te nc i rc led t he n eck a nd h ead , h ave b een c au t ious ly c i ted a s p oss ib le i n d ica t ions t ha tc a t t le d omes t ica t ion m ay h ave b een o ccu r r ing ( Ro l le tson 1 985 :53 , 1 986 :50 ) . F i na l ly , t he e a r l ies t c a t t le r ema ins i n A f r ica t o b e i d en t i f ied a s d omes t ica ted o n t he o s teo log ica l c r i te r ia o fr educed s i ze a re t hose r e cove red f r om t h ree s i tes i nt he N ab ta B as in o f t he e as te rn S aha ra . T hese r ema ins o ccu r i na ssoc ia t ion w i th r ad ioca rbon d a tes p l ac ing t h ese s i tes i nt he e a r ly , m idd le a nd l a te s even th m i l lenn ium B .C . r e spec t ive ly ( Banks 1 984 :78 -79 ; Gau t ie r 1 980 :332 , 3 36 ; Wendor f a nd S ch i ld 1 980a :103 , 1 08 , 1 32 ) . S l igh t ly e a r l ier e v idence f o r d omes t ica ted c a t t le h as a ppa ren t ly b een f ound i nt he B i rK i se iba r eg ion o ft he e as te rn S aha ra , a pp rox ima te ly 1 00 km w es t o f N ab ta P laya ( Wendor f e t a l . 1 984 ) . I th as t he re fo re b een s ugges ted t ha t " ca t t le -keep ing " i nt he e as te rn S aha ra may a c tua l ly d a te a s e a r ly a s 7 , 500 B .C . ( Banks 1 984 :2 ; Wendor f e ta l . 1 984 ) . T he c umu la t ive e v idence w ou ld s eem ing ly i n d ica te t h a t d omes t ica ted c a t t le , a sd e f ined b y o s teome t r ic e v idence o f r educed s i ze , w e re a l ready p resen t b y t he e a r ly s even th m i l lenn ium B .C . i nG reece a nd N o r th A f r ica , a nd w e re p o ten t ia l ly p resen t a tt he s ame t ime i nA na to l ia a nd J o rdan . S ince t he i n i t ia l h usband ry a nd h e rd ing o fc a t t le w ou ld p resumab ly p reda te t h is p e r iod , i n i t ia l c a t t le d omes t ica t ion may w e l l f a l l w i th in t he e i gh th m i l lenn ium B .C . , a s i sc u r ren t ly s ugges ted f o r N or th A f r ica . D omes t ica ted g oa ts , a s d e f ined b y b ehav io ra l c r i ter ia ( s e lec t ive a ge -sex c u l l ing , h oo f p r in ts i nv i l lage b r icks ) , a nd d omes t ica ted s heep , a s d e f ined b y t he c ur ren t ly u nques t ioned c r i te r ia o ff ema le h o rn lessness , s eem ing ly a ppea ra s e a r ly a s t h e m id e i gh th m i l lenn ium BC . H oweve r , w h i le n o t c ommon ly a dd ressed i n mos t f auna l s t ud ies , mo rpho log ica l e v idence o fr educed s i ze i ng oa ts , a nd p oss ib ly s heep , d oes n o t o ccu r u n t i l t he s even th m i l lenn ium B .C . i nt he f auna l s amp le f r om J e r icho ( C lu t ton -B rock 1 981:60 ; C lu t ton B rock a nd U erpmann 1 974 :273 ) , w h i le a s t a t is t ica l ly s i gn i f ican t r educ t ion i nt he l e ng th o fg oa t me taca rpa ls i sn o t a ppa ren t u n t i l a f te rt he P re -Po t te ry N eo l i th ic B p e r iod i ns amp les f r om o t he r s i tes i nt he r eg ion ( Sm i th a nd H orw i tz 1 984 :47 1) . I tt he re fo re s eems p robab le t ha t t he p os tu la ted l a te a ppea rance o fd omes t ica ted c a t t le r e la t ive t os heep a nd g oa ts i nt he N ea rE as t p a r t ia l ly r e f lec ts t he d i f fe ren t ia l a pp l ica t ion o ft he v a r ious c r i te r ia b y w h ich t he d omes t ica t ion p rocess i s t r ad i t iona l ly i n fer red , r a ther t han a n a ccura te i n terp re ta t ion o f t empora l p a leoeconom ic e ven ts . H av ing i n tu i t ive ly d e te rm ined t ha t s heep a nd g oa ts mus th ave b een d omes t ica ted p r io r t o c a t t le , t he a rchaeo log ica l r eco rd h as s eem ing ly b een d i f fe ren t ia l ly i n te rp re ted t o s uppor t t h is l o ng -s tand ing b e l ie f . A f u r ther c omp l ica t ion i n a dd ress ing t he e a r ly d omes t ica t ion o f c a t t le i st he l ow f r equenc ies a t wh ich c a t t le r ema ins g ene ra l ly o ccu r a tE p ipa leo l i th ic a nd N eo l i th ic s i tes i nt he N ea r E as t r e la t ive t o s heep a nd g oa t r ema ins . G ene ra l ly i n terp re ted a t N eo l i th ic s i tes a s r e f lec t ing t he m ino re conom ic s i gn i f icance o f" w i ld "c a t t le r e la t ive t od omes t ic s heep a nd g oa ts , s uch p a t terns may a l te rna t ive ly r e f lec tt h e g rea te re conom ic i mpor tance o fp as to ra lc a t t le h e rds r e la t ive t o s heep a nd g oa ts . F rom t he p e rspec t ive o f r e la t ive e ne rge t ics , i ti se xpec ted t ha t s heep a nd g oa ts s hou ld b e p re fe ren t ia l ly s l augh te red o ve r c a t t le ( and c ame ls ) , a nd w ou ld t he re fo re e n te rf auna l d epos i ts a th i ghe rf r equenc ies t han c a t t le ( o rc ame ls ) w heneve r m ixed h e rd ing s t ra teg ies w e re p rac t iced . T he h i gh r e turn r a tes e xpec ted f o r p as to ra l c a t t le h e rds r e la t ive t os heep a nd g oa ts u nde rb o th mea t -on ly a nd mea t p l us m i lk s t ra teg ies ( F igu res 9a nd 1 2 ) may a c tua l ly p rov ide ama te r ia lb as is f o rt h e h i gh r e l ig ious e s teem i nw h ich N eo l i th ic c a t t le a t Q a ta l H üyük ( Me l laar t 1 975 : 108 110 ) a nd ' A in G haza l ( Ro l le fson 1 983 , 1 986 ) w e re a ppa ren t ly 1 46

h e ld , u l t ima te ly l e ad ing t ot h e p opu la r n o t ion t ha t c a t t le we re i n t ia l ly d omes t ica ted p r ima r i ly f o r r e l ig ious p urposes ( e .g . , H e r re 1 963 ; I s aac 1 962 ) . A l though t he d omes t ica t ion o fc a t t le p r io rt o , o r c on tempo raneous w i th , t he d omes t ica t ion o fs heep a nd g oa ts i nt he N ea r E as tc anno t b e d emons t ra ted , t he a va i lab le e v idence i se qua l ly i n su f f ic ien t f o r d emons t ra t ing t ha t s heep a nd g oa ts we re n ecessa r i ly d omes t ica ted p r io r t o c a t t le . P o ten t ia l ly , a ge -sex s l augh te r s chedu les f o r t he c a t t le c omponen t o f f auna l d epos i ts f r om p rev ious ly e xcava ted s i tes i nt he N ea r E as t m igh t b e u sed t oa dd ress t h is i s sue , a l though w e w i l l n eed t ok now f a r mo re a bou t t he s e lec t ive s l augh te r p rac t ices o f e thnog raph ic h un te r g a the re ra nd p as to ra l p opu la t ions t han w e d o a tp resen t i fs uch a na lyses a re t ob ec onv inc ing . I t i sa l so s t rong ly s uspec ted t ha t c r i t ica l d a ta f o r a dd ress ing t he t empo ra l a ppea rance o f c a t t le d omes t ica t ion i nt he N ea r E as t r ema in u nexcava ted i nt he e a r ly p as to ra l s i tes o f N or th A rab ia a nd t he A rab ian P en insu la .

A rchaeo log ica l E xpec ta t ions B ased U pon t he R e la t ive E ne rge t ics o fE a r ly Whea t a nd B a r ley E xp lo i ta t ion a nd C u l t iva t ion I n t roduc t ion O f t he mo re t r iv ia l e xpec ta t ions d e r ived f r om a n e nerge t ic a na lys is o f e a r ly whea t a nd b a r ley c u l t iva t ion , t he r e la t ive ly h i ghe r r e tu rn r a tes e xpec ted f o r b a r ley c u l t iva t ion s uppor t t he s ugges t ion t ha t b a r ley was t he mos t i mpor tan t o ft hese e a r ly c e rea ls u n t i l h i s to r ic t imes ( Ha r lan 1 967 :201 , 1 977 :360 ; H a r lan a nd Z oha ry 1 966 : 1075 ) . H oweve r , wh i le t he re a re s eve ra l ma jo r i s sues a nd a ssump t ions c once rn ing t he e a r ly e vo lu t ion a nd a dop t ion o fs ubs is tence s t ra teg ies i n vo lv ing c erea ls w h ich c ou ld b e a dd ressed t h rough e ne rge t ic a na lyses , s ubs tan t ive t es ts a re c u r ren t ly h ampe red b yt he l a ck o fa pp rop r ia te d a ta s e ts . F o re xamp le , i twas s ugges ted i nC hap te r 6t ha t c e rea ls wh ich d o n o t e xh ib i t r e la t ive ly h i gh p os tencoun te r r e tu rn r a tes f o r t he i r e xp lo i ta t ion i n n a tura l s tands wou ld h ave b een u n l ike ly c and ida tes f o r e a r ly f ood p roduc t ion s t ra teg ies , s i nce t he a dd i t iona l l abo r r equ i red f o r t he i r c u l t iva t ion wou ld f u r the r d im in ish t he i r r a tes o f e ne rgy r e turn t o l a bo r e xpended . O s tens ib ly , t h is s imp le e ne rge t ic r e la t ionsh ip b e tween c e rea l s pec ies wou ld l a rge ly a ccoun t f o r why s o f ew c e rea ls b ecame t he f ocus o f h uman f ood p roduc t ion r e la t ive t o t he v as t n umbe r o f c erea l s pec ies wh ich w e re , a nd s t i l l a re , a va i lab le f o re xp lo i ta t ion . H oweve r , i no rde rt of u r the rd eve lop a nd t es t t h is p ropos i t ion i nN or th A f r ica a nd t he N ea r E as t , e ne rge t ic c os t a nd b ene f i t d a ta wou ld m in ima l ly b e r equ i red o nt he w i ld p rogen i to rs o ft he r e levan t c u l t iva ted s pec ies , t he n ume rous w i ld c e rea ls a nd s eed r esou rces wh ich v a r ious f ood p roduc ing p opu la t ions c on t inue t o e xp lo i t , b u t d o n o t c u l t iva te ( C la rk 1 976 :77 -78 , 1 980 :576 ; C u r ru thers 1 910 :343 ; I r v ine 1 952 ; Mus i l 1 926 :15 -16 , 1 927 :122 124 , 1 928 :15 16 ; N ico la isen 1 963 :179 -181 , 2 06 -207 ; S . Sm i th 1 980 :470 -472 ; W i l l iams a nd F a r ias 1 972 ) , a nd a ny w i ld c e rea ls a nd s eed r esou rces w h ich a re a l so a va i lab le t ot hese p opu la t ions , b u t a re n o t e xp lo i ted . U n for tuna te ly , wh i le H a r lan 's ( 1967 ) w e l l -known e xpe r imen ta l d a ta o n w i ld e i nko rn h a rves t ing a re w ide ly c i ted i nt he a rchaeo log ica l l i t e ra tu re , c ompa rab le r esea rch h as n o t b een c onduc ted o n t he w i ld p rogen i to rs o fo the rc e rea l d omes t ica tes i nt he N ea r E as t a nd N o r th A f r ica , l e t a l one o n o the rw i ld c e rea l r esou rces . I nt he f o l low ing s ec t ions , t he e ne rge t ic i mp l ica t ions o ft he d a ta g enera ted f o r whea t a nd b a r ley e xp lo i ta t ion a nd c u l t iva t ion w i l l t he re fo re b e u sed t o p rov ide i n s igh ts i n to t he p rocesses w h ich m igh t h ave b een o pe ra t ing i nt he i n i t ia l i n c lus ion o fc e rea ls i n to f o rag ing d i e ts , a nd t he s pa t ia l a nd t empo ra l a ppea rance o f c e rea l c u l t iva t ion s t ra teg ies i nt he N ea r E as t a nd N or th A f r ica .

When S hou ld W i ld Whea ta nd B a r ley H ave E n te red P reh is tor ic D ie ts? A s d i scussed i nC hap te r 6 ,p os tencoun te r r e tu rn r a tes f o r e xp lo i t ing s tands o f r obus t a nd l a rge -seeded r aces o f e i nko rn a ppea r t o b e q u i te h i gh r e la t ive t o t hose e s tab l ished f o r o ther c e rea l a nd s eed r esources w h ich a re , o r p rev ious ly we re , e xp lo i ted b y v a r ious f o rag ing 1 47

p opu la t ions . I fp os tencoun te rr e turn r a tes f o rt h e e xp lo i ta t ion o fr obus t a nd l a rge -seeded r a ces o f emme ra nd b a r ley a re a s h i gh o r h i ghe rt han t hose e s tab l ished f o rw i ld e i nko rn , a l o ft h ese r esources w ou ld e xh ib i t h i ghe r p os tencoun te r r e turn r a tes t han t hose e s tab l ished f o r m os t o f t he r oo t a nd s eed r e sou rces k nown t o h ave b een p reh is to r ica l ly e xp lo i ted i nt he G rea t B as in ( S imms 1 984 , 1 987 ) , a nd s eve ra l p l an t a nd a n ima l r e sou rces c ur ren t ly i n c luded i nt he d i e t o f A che f o rage rs i nE as te rn P a raguay ( Hawkes e ta l . 1 982 ;a s emended i nH i l l e ta l . 1 986 ) . H oweve r , i ts hou ld n o t b e c onc luded f r om s uch c ompa ra t ive o bserva t ions t ha t i nt h e p reh is to r ic N ea rE as t , r obus ta nd l a rge -seeded r a ces o fw hea ts a nd b a r ley s hou ld h ave e n te red t he d i e t o ff o rage rs a s s oon a s t hey w e re a va i lab le . A ga in , o n t he e xpec ta t ion t ha t f o rage rs w ou ld g ene ra l ly max im ize t he i r e ne rgy r e turn r a tes , r e sou rce i t ems s hou ld b e i n c luded i nt h e d i e t i nt he o rde ro ft he i rr ank ing b y r e tu rn r a tes o n ly s o l o ng a st h e r e tu rn r a te f o re ach r e sou rce e xceeds t he r e turns g a ined f o r s ea rch ing , f i nd ing a nd h and l ing a l te rna t ive r esou rces o fh i ghe r r ank . I nt he G rea t B as in ,t he s pa t ia l a nd t empo ra l i n c lus ion o re xc lus ion o fs eve ra l l owe rr a nk ing i t ems i nt he d i e to ff o rage rs w as s eem ing ly ar e su l to fv a r iab i l i ty i nt he e ncoun te r r a tes ( d ens i ty , a bundance ) o f h i ghe r r ank ing i t e ms , p a r t icu la r ly l a rge g ame ( S imms 1 984 :167 -169 , 1 987 ) . S im i la r ly , l ow r ank ing s eed r e sources a ppea rt oh ave b een t empo ra l ly a nd s pa t ia l ly i n c luded o r e xc luded f r om r e cen t A lyawa ra d i e ts i na cco rdance w i th v a r ia t ions i nt h e o ve ra l l r e tu rn r a tes f o r h i gher r ank ing r esources u nder f l uc tua t ing r a in fa l l c ond i t ions ( O 'Conne l l a nd H awkes 1 981:110 , 1 984 :516 ) . F u r the r , o fa pp rox ima te ly 3 0 ma jo rr e sou rces i n c luded i nt he d i e t o f mode rn A che f o rage rs i ne as te rn P a raguay f o r w h ich r e turn r a tes h ave b een e s tab l ished ( Hawkes e ta l . 1 982 :389 , T ab le 3 ; a s emended i nH i l l e ta l . 1 986 :Tab le 2 ) , o n ly f i ve o r s i x o ft he l owes t r ank ing i t ems e xh ib i t r a tes l e ss t han t hose c a lcu la ted i nC hap te r 6f o rw i ld e i nko rn o b ta ined t h rough h and s t r ipp ing ,w h i le o n ly n i ne o rt en i t ems e xh ib i t r a tes w h ich a r ee qua lt o , o rl e ss t han , w i ld e i nko rn r eaped w i th a p r im i t ive s i ck le . S ign i f ican t ly , t he l a rges t g ame a n ima l i nt he j ung les o fe as te rn P a raguay i st he wh i te l ipped p ecca ry , w i th a n a pp rox ima te a du l t w e igh t o f 3 0 k g ( K im H i l l , p e rsona l c ommun ica t ion 1 985 ) . I tw ou ld t he re fore s eem p laus ib le t ha t i n h i gh b i omass e nv i ronmen ts , s uch a s t h ose c on ta in ing l a rge h erb ivo res a nd h e rd a n ima ls s ub jec tt og ame d r ives , e ven t he h i ghes t r a nk ing w i ld c e rea ls c ou ld h ave b een e xc luded f r om t he d i e t , o r r e duced t o am ino rc omponen t o ft h e d i e t , a s af unc t ion o ft he e ncoun te r r a tes ( dens i ty , a bundance ) o f h i ghe r r ank ing r e sou rces . W i ld c e rea ls , i fa va i lab le , may t he re fo re h ave b een i n c luded a nd d ropped f r om f o rag ing d i e ts a t v a r ious t i mes p r io rt ot he E p ipa leo l i th ic p er iod . When v i ewed f r om t h is p erspec t ive , t he a va i lab le e v idence i s ,s imp ly i n su f f ic ien t t o a scer ta in w he the r t he a ppa ren t i n c lus ion o f emme r a nd b a r ley i nK eba ran p e r iod d i e ts b y a round 1 3 ,000 B .C . ( Ba r Y ose f 1 970 ; H en ry 1 983 ; N oy e ta l . 1 983 ) m igh t r e f lec t t he r e cen t e vo lu t ion o f r obus t a nd l a rge -seeded a nnua ls f r om p e renn ia l p rogen i to rs u nde r c ond i t ions o f i n c reased a nnua l c l ima t ic v a r iab i l i ty i nt he p os t -P le is tocene p er iod ( Why te 1 977 ) , d ec l in ing e ncoun te r r a tes ( dens i ty , a bundance ) f o r h i gher r ank ing r esources u nder t hese s ame e nv i ronmen ta ls h i f ts ,o r ac omb ina t ion o fs uch f ac to rs . P resumab ly , t h is i s sue c ou ld b e a dd ressed t h rough d i rec t p a leobo tan ica l r e sea rch t o e s tab l ish whe the r r obus t a nd l a rge -seeded c e rea ls w e re p resen t i nt he N ea r E as t p r io r t o 2 0 ,000 B .C . H owever , i ts eems p robab le t ha t s uch d a ta o n t he G ram ineae i ng ene ra l w i l l b e e x t reme ly s ca rce a nd d i f f icu l t t oo b ta in . A n a l te rna t ive a nd c omp lemen ta ry a pp roach w ou ld b e t o e s tab l ish whe the r o ve ra l l f o rag ing r e turn r a tes i ne nv i ronmen ts w h ich s t i l l c on ta in h e rds o f l a rge mamma ls w ou ld b e s u f f ic ien t ly h i gh t o r educe e ven h i gh r ank ing c e rea ls t o am ino r c omponen to ft he d i e t , o rt oe xc lude t hem e n t i re ly . I nt h is r e spec t , i ti si mpe ra t ive t ha t e ne rge t ic d a ta b e r eco rded among f o rag ing s oc ie t ies i n s uch e nv i ronmen ts b e fore b o th d i sappea r f o reve r .

1 48

R e tu rn R a tes a nd C i rcumsc r ibed E nv i ronmen ts : I mp l ica t ions f o rP os t -P le is tocene H uman D emog raphy , T e r r i to r ia l i ty , a nd S oc ia l C omp lex i ty A s d i scussed i nC hap te rs 3 , 4a nd 6 , s o l o ng a s c e rea ls w ere n o t r emoved f r om t he g enera l l a t i tudes i nw h ich t h ey e vo lved , s o i l a nd w a te r f ac to rs w ou ld h ave c ons t i tu ted t he p r ima ry l i m i t ing f ac to rs i nc e rea l c u l t iva t ion a nd p roduc t iv i ty . U nde r c ond i t ions o fh and t i lage , l i gh t s o i ls w i th n a tura lo re as i ly o b ta ined i r iga t ion a re e xpec ted t op roduce r e la t ive ly h i gh y i e lds w i th a m in imum o fr equ i red l a bo r , a nd t hus p rov ide h i ghe r r e turn r a tes r e la t ive t od ense r s o i ls r equ i r ing g rea te rp repa ra t ion l a bo r ,o rs o i ls w i th u n re l iab le o rh i gh ly f l uc tua t ing w a te rs upp l ies . I n g ene ra l , i ti sa l so e xpec ted t ha t o p t ima l a g r icu l tu ra l h ab i ta ts w ou ld b e c ompa ra t ive ly s ca rce r e la t ive t o a l p o ten t ia l ly a rab le l a nds . I n i t ia l ly , t he e ne rge t ic e xpec ta t ion t ha t s uch l o ca l i t ies s hou ld h ave b een t he f ocus o fe a r ly c e rea l c u l t iva t ion s t ra teg ies s eems s t rong ly s uppor ted b y t he k nown s pa t ia l l o ca t ions o f ma jo rs i tes w he re e a r ly c e rea l p roduc t ion i sa t tes ted o rs uspec ted ( s ee T ab le 2 ) . H oweve r , t h e i mp l ica t ions o f t h is e nerge t ic o bserva t ion g o f a r b eyond t he s imp l is t ic e xpec ta t ion o f as pa t ia l c o r re la t ion b e tween e a r ly a g r icu l tu ra l s i tes a nd f avo rab le b u t c i rcumsc r ibed h ab i ta ts . T he d i f fe ren t ia l o ccupa t ion a nd e xp lo i ta t ion o fh ab i ta ts o r h ab i ta t t y pes i na cco rdance w i th t he e ne rgy r e tu rn r a tes w h ich c ou ld b e o b ta ined i nt hem w ou ld s eem ing ly p rov ide a u se fu l i n te rp re t ive a pp roach f o r u nde rs tand ing b o th w hy f o rage rs w ou ld a dop t f ood p roduc t ion s t ra teg ies i nt he f i r s t p l ace , a nd w hy i n c reas ing t e r r i to r ia l i ty a nd s oc ia l c omp lex i ty w ou ld s ome t imes a ccompany s uch s ubs is tence s h if ts . D raw ing f r om t heo re t ica l c ons idera t ions o f t he e vo lu t iona ry f ac to rs w h ich wou ld a f fec t h ab i ta t s e lec t ion ( F re twe l l a nd L ucas 1 970 :18 -26 ) a nd b ehav io ra l e xpec ta t ions d e r ived f r om o p t ima l f o rag ing mode ls , O 'Conne l l a nd H awkes ( 1981:114 -116 ) a dd ressed t he p rocesses w h ich m igh t h ave b een o pe ra t ing i nt he a ppa ren t p a t tern a nd r a te o f t he c o lon iza t ion o f A us t ra l ia b y p reh is to r ic f o rage rs . T hey p os i ted t ha t , i ng ene ra l , t he o rde r i nw h ich h ab i ta ts o r h ab i ta tt ypes w e re o ccup ied s hou ld h ave v a r ied d i rec t ly w i th t he n e t e ne rgy r e turn r a tes g a ined f r om e xp lo i t ing t hem . H ab i ta ts i nw h ich e ne rgy r e tu rns w e re c ompa ra t ive ly l ow s hou ld h ave r ema ined u noccup ied u n t i l t h e r e tu rns i nc ompa ra t ive ly " be t ter ' h ab i ta ts f e l l t ot he s ame l e ve l . Movemen t i n to n ew h ab i ta ts s hou ld t he re fo re h ave b een a f unc t ion o f p opu la t ion i n c rease ( e i ther t h rough i n terna l g row th o r i mm ig ra t ion ) w h ich d ep le ted o r d im in ished a va i lab le r e sources i no ccup ied h ab i ta ts t o ap o in t w he re movemen t t o a l te rna t ive h ab i ta ts was mo re e f f ic ien t , o r e nv i ronmen ta l c hanges wh ich r educed r e turn r a tes i n o ccup ied h ab i ta ts o r i n c reased t hose i nu noccup ied h ab i ta ts . Wh i le t h is mode l p or tends t o i n teg ra te w ha t w ere p rev ious ly s een a s c on f l ic t ing i n terp re ta t ions o f t he h uman c o lon iza t ion p rocess i n to a s yn the t ic e vo lu t iona ry e xp lana t ion c ons is ten t w i th t he a rchaeo log ica l , p a leoenv i ronmen ta l a nd e t hnog raph ic r e co rds o fA us t ra l ia , t he s ame mode l w ou ld s eem ing ly h ave b road a pp l icab i l i ty t o t he f o rage r / food p roduc t ion t r ans i t ion i nt he N ea rE as t a nd N o r th A f r ica . P resumab ly , t e rm ina l P a leo l i th ic p opu la t ions o n t he A rab ian P en insu la a nd i nN o r th A f r ica w ou ld h ave t r acked t he c hanges i ne nv i ronmen t w h ich a ppa ren t ly o ccur red a tt he e nd o ft he P le is tocene , l a rge ly a bandon ing i n c reas ing ly d es icca ted h ab i ta ts i np re fe rence t ot he r emnan t a nd c ompa ra t ive ly c i rcumsc r ibed h ab i ta ts w he re f o rag ing r e turn r a tes r ema ined r e la t ive ly h i gh . W i th i n c reas ing ly d ense h uman o ccupa t ion i nr e la t ive ly c i rcumsc r ibed r eg ions u nde rc ond i t ions o fc on t inued e nv i ronmen ta l d ec l ine , af u r ther d rop i nf o rag ing r e tu rn r a tes t h rough t he d ec l ine o r d ep le t ion o f h i gh r ank ing r e sources s hou ld h ave l e d t o t he i n c lus ion o f l owe r r ank ing r e sources i n to t he d i e t , a nd p o ten t ia l ly t he h usbandry o f r ema in ing h i gh r ank ing r e sou rces . I f a ve rage f o rag ing r a tes c on t inued t o d ec l ine , t hey w ou ld h ave e ven tua l ly f a l len e qua l t o t he l owe r mean r e turn r a tes e xpec ted f o r d i rec t f ood p roduc t ion b ased u pon s pec i f ic p l an t a nd a n ima l s pec ies . E ven i nt he s imp l is t ic f o rm p resen ted h ere , t h is i n te rp re t ive mode l s eem ing ly r e ce ives b road s uppo r t f r om t he a rchaeo log ica l a nd p a leoenv i ronmen ta l r e cords o ft he N ea r E as t a nd N or th A f r ica . A f te r p r ior o ccupa t ion , t he v i r tua l a bandonmen t o f t ha t p or t ion o f t he A rab ian P en insu la r ep resen ted b y m odern S a 'ud i A rab ia d ur ing t he p er iod o f p os t -P le is tocene d es icca t ion p reced ing t he H o locene w e t p hase i sw ide ly a t tes ted i nr e cen t s ys tema t ic r eg iona l s u rveys ( Adams e ta l . 1 977 ;G a r ra rd a nd H a rvey 1 98 1 ;G i lmo re e ta l . 1 982 ; I n g raham e ta l . 1 98 1 ; P a r r e ta l . 1 978 ; Z a r ins e ta l . 1 979 , 1 980 ) . D ur ing t h is s ame p e r iod f u r the r n or th i nw ha t was b ecom ing t he " Fer t i le C rescen t ," i n c reas ing ly l a rge E p ipa leo l i th ic s i tes c l us te red i na ppa ren t ly f avo rab le h ab i ta ts a nd s eem ing ly r e f lec t ing i n c reas ing h uman p opu la t ion d ens i ty a nd 1 49

c o r respond ing r educ t ions i nl o ca l g roup f o rag ing r ange h ave l o ng b een r ecogn ized ( Ba r -Yose f 1 975 :372 -373 , 1 986 :158 159 ; H enr y 1 983 :10 1, 1 5 1; H en ry e t a l . 1 98 1:55 ; Moo re 1 979 :63 6 5 , 1 982 :227 -229 , 1 985 :12 -13 , 4 6 -48 ) . T he i n c reased i mpo r tance a nd i n tens i f ica t ion o f g aze l le h e rd e xp lo i ta t ion a ppa ren t among N a tu f ian a nd o the r E p ipa leo l i th ic p opu la t ions d u r ing t h is p e r iod ( Dav is 1 982 , 1 983 ; Ga r ra rd 1 984 ; H en ry 1 975 ; L egge 1 972 ; L egge a nd R ow ley C onwy 1 987 ; Moo re 1 982 ; S immons 1 975 -77 ) a l so a ccords we l l w i th s uch e ne rge t ic e xpec ta t ions .Thus , b y t he n in th m i l lenn ium B .C . , l a rge s eden tary o r s em iseden ta ry c ommun i t ies o f i n tens ive f o rage rs e x is ted a t s e lec t e nv i ronmen ta l l o ca t ions , w i th t he h usbandry o fc er ta in p l an t a nd a n ima l r esou rces f r equen t ly s ugges ted ( Hen ry 1 983 ; H en ry e t a l . 1 981 ; Moo re 1 979 , 1 982 , 1 985 ) . B y t he e nd o f t he l a te r E p ipa leo l i th ic p e r iod ( r ough ly 1 0 ,000 -8 ,500 B .C . ) a nd d ur ing t he e a r l ies t p e r iod o f t he A ce ram ic N eo l i th ic ( r ough ly 8 ,500 7 ,600 B .C . ) , f ood p roduc t ion s t ra teg ies w e re u nde rway i nf a vo rab le m ic roenv i ronmen ts ( Moo re 1 982 :229 , 1 985 :22 -28 ) . As eem ing ly p a r ,a l le l s equence o f e ven ts a ppea rs i nt he a rchaeo log ica l r eco rd o f N or th A f r ica . A f te r p r io r o ccupa t ion , t he S aha ra was v i r tua l ly a bandoned d u r ing t he h ype ra r id p os t P le is tocene p e r iod p reced ing t he H o locene we t p hase ( C la rk 1 980 :562 ; Mc In tosh a nd M c In tosh 1 981 :604 ; Wendor f a nd S ch i ld 1 980a :257 ) . D u r ing t he s ame p e r iod , s i tes a l ong t he N i le i n b o th s ou the rn E gyp t a nd t he S udan i n c reased i nb o th n umbe r a nd s i ze , r e f lec t ing i n c reas ing h uman p opu la t ion d ens i t ies ( C la rk 1 980 :552 -558 ; H assan 1 980 :437 -438 , 4 46 ) . T he h i gh f r equenc ies o f f i sh , wa ter fow l a nd s he l l f ish r ema ins , a nd t he n ea r ly u n ive rsa l a ppea rance o fq uan t i t ies o fg r ind ing s t ones , s eem ing ly r e f lec ts a s imu l taneous b roaden ing o f t he d ie t t o i n c lude l owe r r ank ing f ood r e sources i n ap a t te rn s im i la rt ot ha t c on tempo raneous ly o bse rved i nt he N ea r E as t ( F lannery 1 965 , 1 969 , 1 973 ) . C o r respond ing d ec reases i nl o ca l g roup f o rag ing r ange , t he s eden ta ry o r s em iseden ta ry o ccupa t ion o f s pec i f ic s i tes i nf avo red l o ca t ions , a nd t he h usband ry o fc er ta in p lan t a nd a n ima l r e sources a re a l so p robab le ( C la rk 1 980 :555 -558 ) . U n fo r tuna te ly , s ubsequen t a rchaeo log ica l d a ta w h ich m igh t r e f lec t t he i n i t ia l a dop t ion o f f ood p roduc t ion s t ra teg ies a l ong t he N i le a re m iss ing . E x t reme N i le f l ow v a r ia t ions d u r ing t h is c r i t ica l p e r iod may h ave i n f luenced t he a rchaeo log ica l v i s ib i l i ty o fr e levan t s i tes ( C la rk 1 980 :556 ; H asan 1 980 :446 ; Wendo r f a nd S ch i ld 1 980a :274 , 2 76 , 1 980b :262 ) . I ti sp resen t ly u nknown i f i n tens ive f o rag ing c on t inued a l ong t he N i le w i th p opu la t ions a dop t ing f ood p roduc t ion a s t hey em ig ra ted i n to t he e as te rn S aha ra d u r ing t he H o locene w e t p hase , o rw he the rf ood p roduc t ion e vo lved a ts pec i f ic l o ca t ions a l ong t he N i le w i th t he s ubsequen t e xpans ion o fs uch p opu la t ions i n to t he e as te rn S aha ra a s c l ima t ic c ond i t ions ame l io ra ted . R ega rd less , t he g ene ra l c ong ru i ty o f a rchaeo log ica l a nd p a leoenu i ronmen ta l d a ta i nt he N ea r E as t a nd N o r th A f r ica w i th e xpec ta t ions d e r ived f r om a s imp le i n te rp re t ive mode l wh ich a ccoun ts f o rt he s pa t ia l a nd t empo ra l e vo lu t ion o ff ood p roduc t ion s t ra teg ies i nt e rms o fr e la t ive e ne rge t ics m in ima l ly s uppor ts t he e f f icacy o f s uch a n a pp roach . I ndeed , a n e vo lu t ionary a pp roach t o b ehav io ra l i s sues b ased u pon t he r e la t ive e ne rge t ics o f a l te rna t ive s ubs is tence o p t ions wou ld s eem ing ly h o ld f u r the r i mp l ica t ions f o r o ur u nde rs tand ing t he s pa t ia l a nd t empo ra l a ppea rance o fi n c reas ing ly t e r r i to r ia l a nd c omp lex s oc ie t ies i nt he N ea r E as t a nd N o r th A f r ica . C ons is ten t w i th e vo lu t iona ry mode ls o f t e r r i to r ia l i ty i n vok ing e conom ic d e fendab i l i ty ( Dyson -Hudson a nd Sm i th 1 978 ; K rebs a nd D av ies 1 985 :86 -88 ) , i ts eems r easonab le t o e xpec tt ha tt he h i gh r e tu rn r a tes wh ich c ou ld b e o b ta ined b yc u l t iva t ing c e rea ls i nc i rcumsc r ibed e nv i ronmen ts w i th r i ch s o i ls a nd n a tura l o r e as i ly o b ta ined i r iga t ion wou ld h ave p rov ided a s i tua t ion i n wh ich t he b ene f i ts d e r ived f r om p e rmanen t ly o ccupy ing a nd e xp lo i t ing t hem o u twe ighed t he s oc ia l c os ts o ft he i r e xc lus ive u se a nd d e fense . F o r e xamp le , t he c ons t ruc t ion •o f d e fense w orks e nc i rc l ing t he s i te o f J e r icho d ur ing t he P re -Po t tery N eo l i th ic A p e r iod ( r ough ly 8 ,500 -7 ,600 B .C . ) , a ppa ren t ly i n c lud ing aw ide r ock c u t d i tch a nd amass ive s t one c i rcu i t wa l lw i th a tl e as to ne t owe r , t empo ra l ly c o inc ides w i th b o th t he i n i t ia t ion o f c e rea l c u l t iva t ion o n t he s i te ( Hop f 1 969 ; K enyon 1 970 :331) a nd t he f i na l m i l lenn ium o ft he p os t -P le is tocene h ype ra r id p e r iod o n t he A rab ian P en insu la p reced ing t he H o locene w e t p hase . G iven t he c ompa ra t ive ly u n ique g eomorpho log ica l a nd h yd ro log ica l s e t t ing o fJ e r icho a s d i scussed i nC hap te r4 ,i ts eems p l aus ib le t ha t t he h i gh mean r e tu rn r a tes f o r c erea l c u l t iva t ion wh ich c ou ld h ave b een o b ta ined a t J e r icho p rov ide a n e ne rge t ic e xp lana t ion f o r w hy t h is p a r t icu la r s i te w ou ld e xh ib i t t he e a r l ies t a ppa ren t e v idence f o r t he 1 50

p ronounced t e r r i to r ia l i ty a nd s oc ia l c omp lex i ty n o rma l ly a ssumed a s c ha rac te r is t ic f o r m uch l a te r " c iv i l iza t ions . " F u r the r , t he c ompa ra t ive ly u n ique s e t t ing o f J e r icho a nd a c ons ide ra t ion o f a l te rna t ive e ne rge t ic o p t ions u nde r c hang ing e nv i ronmen ta l a nd d emog raph ic c i rcums tances s eem ing ly p rov ide a nswe rs f o r why at e r race wa l l a l one wou ld h ave b een s u f f ic ien t f o r t i f ica t ion d u r ing t he s ubsequen t P re -Po t tery N eo l i th ic B p e r iod ( r ough ly 7 ,600 -6 ,000 B .C . ) , a nd why t he re i ss o l i t le e v idence o fo t he rf o r t i f ied s i tes i nt he N ea r E as t u n t i l l a te rp e r iods . T he r e la t ive ly b r ie f e f f lo rescence o f c omp lex i ty a tJ e r icho o ccu r red a tt he e nd o f al o ng p e r iod o fd ec l in ing s ubs is tence o p t ions i nt he N ea r E as t , wh i le t he s ubsequen t H o locene we t p hase w ou ld h ave o pened v as t t r ac ts o f l a nd f o r b o th a g r icu l tura l a nd p as to ra l e xp lo i ta t ion d u r ing t he P re -Po t tery N eo l i th ic B p e r iod . P resumab ly , i n c reased f o rag ing o ppo r tun i t ies wou ld h ave r e tu rned t ot hose r eg ions whe re p l an t a nd a n ima l c ommun i t ies b ecame r ees tab l ished i n s u f f ic ien t d ens i ty . A f ter 6 ,000 B .C . , w i th t he e nv i ronmen t s h i f t ing t owa rds mode rn c ond i t ions a nd s ubs is tence o p t ions a ga in d ec l in ing , t he t e r r i to r ia l i ty r e f lec ted i nJ e r icho 's d e fens ive wa l ls a ga in a ppea red a ts uch s i tes a s S ama r ran T e l l e s -Sawwan ( Abu a l -Soo f 1 968 ) a nd C hoga Mam i ( Oa tes a nd Oa tes 1 976 :104 ) i nt he s i x th m i l lenn ium ,H ac i la r i nt he f i th m i l lenn ium ( Me l laa r t 1 975 :116 ) , a nd J awa i nt he f our th m i l lenn ium ( He lms 1 981) . T hus , t he r i se o f i n c reas ing ly c omp lex s oc ie t ies was s eem ing ly p os tponed u n t i l f u r the r e nv i ronmen ta l a nd d emog raph ic s h i f ts o ccu r red a tt he e nd o ft he H o locene we t p hase . S uch c ons ide ra t ions g rea t ly d im in ish t he f o rce o fo b jec t ions r ecen t ly r a ised b y B a r -Yose f ( 1986 :158 ) t ov i ew ing t he wa l ls o fJ e r icho a s d e fens ive . C oncur ren t w i th t he H o locene we t p hase o n t he A rab ian P en insu la , p as tora l s oc ie t ies a ppa ren t ly s p read a c ross t he r e tu rn ing g rass lands , wh i le h un te r -ga the rer p opu la t ions s eem ing ly r eoccup ied t he g ene ra l r eg ion o ft he r e juvena ted l a kes i nt he R ub ' A l K ha l i ( Edens 1 982 ; F i e ld 1 955 , 1 958 , 1 960a , 1 960b ; Sm i th a nd Ma ran j ian 1 962 ; Z a r ins e ta l . 1 979 ; Z eune r 1 954 ) . S im i la r ly , t h is p e r iod o f c l i ma t ic ame l io ra t ion i sv i ewed a s b e ing r espons ib le f o r a n i n c rease i nt he n umbe r o fk nown s i tes i nt he N egev a nd S ina i d u r ing t he P re -Po t tery N eo l i th ic Bp e r iod , c ur ren t ly i n terp re ted a s r e f lec t ing h un te r -ga the re r e conom ies ( Ba r Y ose f 1 984 , 1 986 :159 ; N oy e t a l . 1 981:88 ; T che rnov a nd B a r -Yose f 1 982 ) . D u r ing t h is p e r iod i nN or th A f r ica , p as tora l s oc ie t ies a ppea red i n t he e as tern S aha ra , wh i le c on temporaneous r eoccupa t ion i nt he c en t ra l , s ou the rn a nd wes te rn S aha ra r e f lec ts h un te r -ga the rer p opu la t ions w i th e conom ies b ased u pon h un t ing , f i sh ing a nd w i ld g ra in e xp lo i ta t ion ( C la rk 1 980 :562 -568 ; M c In tosh a nd Mc In tosh 1 981 :605 -608 ; A . Sm i th 1 980a :452 -453 , 4 63 ) . P as tora l s ubs is tence s ubsequen t ly s p read a c ross t he S aha ra a t t he e xpense o f p r io r f o rag ing s t ra teg ies i n c on junc t ion w i th d ec l in ing e nv i ronmen ta l c ond i t ions i nl a te rp er iods o ft he H o locene we t p hase ( C la rk 1 980 :565 -568 ; Mc In tosh a nd Mc In tosh 1 981 :606 -608 ; A . Sm i th 1 980a , 1 980b ; Munson 1 976 ) . W i th t he d ec l ine o f t he H o locene we t p hase a f te r 6 ,000 B .C . a nd t he e ven tua l o nse t o f mode rn a r id c ond i t ions o n t he A rab ian P en insu la a nd N or th A f r ica , ap e r iod o f d ec l in ing s ubs is tence o p t ions s im i la rt ot ha t wh ich p receded t he H o locene we t p hase wou ld h ave a ga in o ccu r red . P resumab ly , f o rag ing a nd f ood p roduc ing p opu la t ions wou ld o nce mo re h ave t r acked t hese e nv ironmen ta l s h i f ts , w i th o ccupa t iona l d ens i ty s h i f t ing f r om i n c reas ing ly d es icca ted h ab i ta ts i np re fe rence t ot hose r e la t ive ly c i rcumsc r ibed r eg ions w i th mo re f avo rab le c ond i t ions . H oweve r , i nt he " Fer t i le C rescen t , " p r io r p opu la t ions o f f ood p roduce rs h ad n o t d i sappea red d u r ing t he H o locene w e t p hase , a nd t he mos tf avo rab le a g r icu l tu ra l l o ca l i t ies we re a l ready o ccup ied . T h is f i na l i n f lux o fa dd i t iona l p opu la t ions i n to wha t was b ecom ing t he " Fer t i le C rescen t " a tt he e nd o ft he H o locene w e t p hase s hou ld h ave r esu l ted i ni n c reased c ompe t i t ion o ve re ne rge t ica l ly o p t ima l l o ca l i t ies , a nd amo re i n tens ive e xp lo i ta t ion o fr e la t ive ly l e ss favo rab le r eg ions . I ts eems r easonab le t o s uspec t t ha t u nde r t hese c ond i t ions , i n d iv idua ls wou ld h ave b een i n c reas ing ly w i l l ing t oi n ves t l abo r i n mo re i n tens ive wa te rc on t ro l a nd d e l ivery s ys tems i n t hose r eg ions whe re h i gh c e rea l r e turns wou ld h ave o f fse t i n c reased l a bo rc os ts . T h is p rocess i sp a r t icu la r ly a ppa ren t i nc en t ra l Mesopo tam ia , whe re S ama r ran s e t t lemen t a nd s ubs is tence i nt he s i x th m i l lenn ium was s eem ing ly b ased u pon t he r egu la ru se o fi r iga t ion f o r c rops ( Moo re 1 985 :59 ; O a tes 1 969 :122 -127 , 1 972 :302 -303 ) . A s s umma r ized b y Moo re ( 1985 :59 ) : T he r egu la r u se o f i r iga t ion , l i ke t he e v idence o f s e t t lemen t p l ann ing a nd d e fens ive s t ruc tu res , i n d ica tes t ha t t hese c ommun i t ies w e re o rgan iz ing t he i r a f fa i rs i n a mo re t ho roughgo ing manne r . T hey h ad t hus a dvanced s i gn i f ican t ly 1 51

b eyond af o rma l N eo l i th ic e x is tence a nd w ere o n t h e t h resho ld o fd eve lop ing a nu rban w ay o fl i f e [ emphas is a dded ] . T hese a r t i f ic ia l ly c rea ted o p t ima l e nv i ronmen ts s hou ld h ave , i nt u rn , b ecome t he f o cus o f i n c reas ing c ompe t i t ion a nd t e r r i to r ia l i ty ( w i tness T e l l e s -Sawwan a nd C hoga Mam i ) . B yt he l a te s i x th m i l lenn ium B .C . , U ba id s e t t lemen ts o nt he a luv ia l p l a in o fs ou theas te rn I r aq a l so e xh ib i ted a n u rban c ha rac ter w i th a h e rd ing a nd a g r icu l tu ra l e conomy s uppo r ted b y i r iga t ion ( Moo re 1 985 :60 ) . B yt he l a te f o ur th m i l lenn ium B .C . , ma tu re u rban c i v i l iza t ion a nd t he f i r s t r e cogn izab le s t a tes h ad eme rged . C ons is ten t w i th e ne rge t ic e xpec ta t ions a nd p os tu la ted d emog raph ic s h i f ts , a ppa ren t p opu la t ion i n c reases d u r ing t he f i f th t om id th i rd m i l lenn ia B .C . , c o inc iden tw i th t he f i na l s h i f t t o mode rn e nv i ronmen ta l c ond i t ions , h ave p rev ious ly b een v i ewed a s a n i mpo r tan tf ac to ri nt he r i se o fs uch i n c reas ing ly c omp lex a nd t e r r i to r ia ls oc ie t ies i nM esopo tam ia ( e .g . , A dams a nd N issen 1 972 ; C a rne i ro 1 970 ; G ibson 1 973 ; Y oung 1 972 ) . I ts hou ld b e n o ted t ha t t h is e n t i re p rocess i sq u i te c ons is ten tw i th A lexande r 's " Ba lance o f P ower H ypo thes is " ( 1979 :222 -233 ) t o a ccoun t f o r t he e vo lu t ion o f i n c reas ing ly c omp lex h uman s oc ie t ies . H oweve r , b y f o cuss ing a t ten t ion u pon e xpec ted c o r re la t ions b e tween p os t P le is tocene e nv i ronmen ta l c hanges , a t tendan t s h i f ts i nh uman s ubs is tence a nd d emog raphy , a nd t he c i rcumsc r ip t ion o f o p t ima l f ood p roduc t ion l o ca l i t ies , t he p resen t s t udy e xpands A lexande r 's o r ig ina l h ypo thes is b y a ccoun t ing f o rw hy i n c reas ing ly c omp lex s oc ie t ies a ppea red whe re a nd w hen t hey d id . H e re a ga in , t he c onco rdance b e tween a rchaeo log ica l a nd p a leoenv i ronmen ta l d a ta , a nd s imp le b ehav io ra l e xpec ta t ions b ased u pon r e la t ive e ne rge t ics , w ou ld s eem ing ly s uppo r tt he e f f icacy o fa n e vo lu t iona ry a pp roach t oa r chaeo log ica l i n fe rence .

A rchaeo log ica l E xpec ta t ions B ased U pon E ne rge t ic C ompa r isons B e tween P as to ra l a nd C e rea l S ubs is tence Op t ions

G ene ra lC ons ide ra t ions I n t he f o l low ing s ec t ions , e ne rge t ic e xpec ta t ions c once rn ing t he s pa t ia l a nd t empo ra l e vo lu t ion a nd a dop t ion o fa l te rna t ive p as to ra l a nd c e rea l s ubs is tence o p t ions a re g ene ra ted b y c ompa r isons b e tween t he r e tu rn r a tes o f s uch s t ra teg ies . T hese e xpec ta t ions a re t hen c hecked a ga ins t H o locene e poch a rchaeo log ica l d a ta r e levan t t o t he s pa t ia l a nd t empo ra l a ppea rance o f p as to ra l a nd c e rea l s ubs is tence s t ra teg ies . B e fo re p roceed ing , h owever , t he p o ten t ia l m ix ing o fs ubs is tence o p t ions a nd v a r ia t ions i nt he r e la t ive e xchange v a lue o fp as to ra l a nd c e rea lc a lo r ies m us tb e a dd ressed . I n i t ia l ly , i ts eems p robab le t ha t e a r ly f ood p roduc t ion s t ra teg ies o f ten i n vo lved a m ix o f s ubs is tence o p t ions ( F lannery 1 965 :1247 -1256 , 1 969 :73 -100 ; H o le e t a l . 1 969 ; Moo re 1 985 :27 ) . C e r ta in ly t h is h as b een t he c ase i nt he e t hnog raph ic a nd e thnoh is to r ic p resen t . P as to ra l is ts o f ten f o rage f o rw i ld p l an t r e sou rces a s o ppo r tun i t ies p e rm i t , h un tw i ld g ame i fi ti s a va i lab le , a nd s easona l ly p l an t c e rea ls i fs u i tab le h ab i ta ts a re a va i lab le a nd s chedu l ing o r h ouseho ld d ua l res idence p a t te rns p e rm i t i t .S im i la r ly , a g r icu l tura l p opu la t ions f r equen t ly k eep l i ves tock ( o f ten i nc ommuna l h e rds ) , c o l lec t w i ld p l an t r e sou rces a nd h un t i fs uch o ppo r tun i t ies e x is t . " Pure " o r " spec ia l ized " f ood p roduc t ion s t ra teg ies i n t he s ense o f a n e xc lus ive d ependence o n e i the r p as to ra l o r a g r icu l tura l p roduc ts s eems u n l ike ly e xcep t i n a marke t e conomy . Many , i fn o t a l , o ft he e a r l ies t f ood p roduc ing s oc ie t ies i nt he N ea r E as t a nd N or th A f r ica a ppea rt oh ave p rac t iced s uch m ixed e conom ies . P resumab ly ,t he d ec is ion mak ing w h ich u nde r l ies s uch m ixes r e s ts u pon t he r e la t ive c os ts a nd b ene f i ts o f a l terna t ive s ubs is tence o p t ions a nd a t temp ts t o m in im ize t he r i sk o f s ubs is tence f a i lu re w hen o n ly o ne o r af ew s ubs is tence o p t ions a re p u rsued . H oweve r , g i ven t he g ross r e so lu t ion o fe ne rge t ic d a ta o n p as to ra l a nd c e rea l s ubs is tence o p t ions a sg enera ted i nt he p resen t s t udy a nd v a r ious p rob lems i na dequa te ly i d en t i fy ing m ixed s t ra teg ies i nt he a rchaeo log ica l r e co rd , i ti ss ugges ted t ha t e ne rge t ic a na lyses o fm ixed s t ra teg ies s hou ld i n i t ia l ly b e c onduc ted among e t hnog raph ic p opu la t ions w i th t he r e su l ts s ubsequen t ly a pp l ied t o a rchaeo log ica l d a ta . F o r t e s t ing p urposes i nt he p resen t s t udy , t he t empora l a nd s pa t ia l a ppea rance o fa g r icu l tu ra l s t ra teg ies w i l l b e measu red b yt h e e x is tence o fs eem ing ly s eden ta ry c ommun i t ies w i th e v idence o f c e rea l f ood p roduc t ion , w h i le t he s pa t ia l a nd t empora l 1 52

a ppea rance o fp as to ra l s t ra teg ies w i l l b e measured b y t he e x is tence o fs eem ing ly t empo ra ry e ncampmen ts w i th e v idence o f p as to ra l f ood p roduc t ion a t s ome d i s tance f r om k nown s eden tary c ommun i t ies . A more d i f f icu l t p rob lem t o e va lua te i ng ene ra t ing a nd t es t ing p red ic t ions d e r ived f r om c ompa r isons b e tween t he r e tu rn r a tes o fp as to ra l a nd c e rea l s ubs is tence s t ra teg ies i n vo lves t he p robab i l i ty t ha t p l an t a nd a n ima l c a lo r ies s hou ld n o t b e c ons ide red e qu iva len t . A s d i scussed a tl e ng th b y H i l l e ta l . ( 1986 ) , s eve ra l s t ud ies s ugges tt ha tt he p ro te in a nd f a t ( l i p ids ) o fa n ima l p roduc ts w ou ld b e o fg rea te rb i o log ica l v a lue t han p l an t c omponen ts ( e .g . , C raw fo rd 1 975 ; C raw fo rd e ta l . 1 98 1) . A s ar e su l t , i ti se xpec ted t ha t p l an t -de r ived c a lo r ies s hou ld b e v a lued s omewha t l e ss t han a n ima l -de r ived c a lo r ies . P resumab ly , t he e xchange v a lues o fa n ima l a nd p l an tc a lo r ies w ou ld s h i f t r e la t ive t oo ne a no the ra s af unc t ion o ft he amoun to fe ach w h ich i sa l ready a va i lab le . Wh i le t h ese c ons ide ra t ions w ou ld b e p a r t icu la r ly c r i t ica l i nt he c ase o fh i gh c a rbohyd ra te r oo t c rops , t he n a tu re o f s uch r e la t ionsh ips b e tween p as to ra l p roduc ts a nd c e rea ls h as n o t b een e xp lo red . F o r s ubsequen t p red ic t ive p urposes , a n e qua l e xchange r a te b e tween c e rea l a nd a n ima l c a lo r ies w i l l b e a ssumed , r ea l iz ing t ha t t h is p o ten t ia l ly i n f la tes t he a c tua lv a lue o fc e rea lc a lo r ies . I ti ss uspec ted , h oweve r , t ha t as l igh t ly h i ghe re xchange r a te f o r a n ima l o ver c e rea l c a lor ies w ou ld a c tua l ly r e in fo rce t he s pa t ia l a nd t empora l e xpec ta t ions g ene ra ted b e low .

E ne rge t ic E xpec ta t ions F i gure 1 9 d i ag rams t he e ne rge t ic r e la t ionsh ips e xpec ted b e tween e a r ly c e rea l s t ra teg ies a nd mea t -on ly p as to ra l s t ra teg ies . T he p os tencoun te r r e tu rn r a tes g i ven f o r w i ld e i nko rn a re t hose p rev ious ly p resen ted i nT ab le 3 6 . T he r ange o f r e turn r a tes d es igna ted a s " A " r e f lec ts t he r ange o f r e tu rns e xpec ted b e tween w hea t a nd b a r ley c u l t iva t ion o n a ve rage l a nds u nde r c ond i t ions o fh and t i lage a nd e a r ly l i t h ic t e chno logy a s p rev ious ly p resen ted i nF i gure 1 8 . T he r anges o fp o ten t ia l r e tu rn r a tes f o r p as to ra l h e rds o fv a r ious s i zes u nde r mea t -on ly s t ra teg ies a re t hose wh ich w e re p rev ious ly p resen ted i n F i gu re 9 . A s c an b e s een , t he r e tu rn r a tes e xpec ted f o rc u l t iva t ing c e rea ls o n l a nds o fa ve rage q ua l i ty u nde rc ond i t ions o fh and t i lage a nd e a r ly l i t h ic t echno logy a re c ons is ten t ly g rea te rt han t he r anges o fr e turn r a tes e xpec ted f o r a l m ea t -on ly p as to ra ls t ra teg ies . F i gure 2 0 d i ag rams t he e ne rge t ic r e la t ionsh ips e xpec ted b e tween e a r ly c e rea l s t ra teg ies a nd m ea t p l us m i lk p as to ra ls t ra teg ies . I na dd i t ion t ot he r e tu rn r a te d a ta o nw i ld e i nko rn a nd t he r a nge o fr e turn r a tes f o rw hea t a nd b a r ley c u l t iva t ion a s p resen ted i nF i gu re 1 9 ,t he r e tu rn r a tes e xpec ted f o r w hea t a nd b a r ley y i e ld ing 1 00 k g /du u nde r c ond i t ions o f, h and t i lage a nd e a r ly l i t h ic t e chno logy h ave a l so b een i n c luded . T hese l a t te rf i gures w ou ld r o ugh ly r e f lec t t he mean r e turn r a tes e xpec ted f o rc e rea lc u l t iva t ion o no p t ima l l a nds a sp rev ious ly d i ag rammed i nF i gu re 1 8 . T he r anges o fp o ten t ia l r e tu rn r a tes f o rp as to ra l h e rds o fv ar ious s i zes a re t h ose w h ich w e re p rev ious ly p resen ted i nF i gu re 1 2w hen m i lk p rocess ing i sa ccoun ted f o r . A sc an b e s een , o n ly t he r e turn r a tes f o r h a rves t ing w i ld s t ands o r f o r c u l t iva t ing c e rea ls i no p t ima l e nv i ronmen ts w ou ld c ons is ten t ly f a l l h i ghe r t han t he r e turn r a tes e xpec ted f o r l a rge c ame l o r c a t t le h e rds . L a rge h e rds o f s heep y i e ld r e turn r a tes e qua l t o , o r s l igh t ly a bove , t hose e xpec ted f o rw hea t a nd b a r ley c u l t iva t ion o n a ve rage l a nds . T he l e as t e f f ic ien t s t ra tegy i nt e rms o fe nergy r e tu rn r a tes i n vo lves h e rds o fg oa ts . Wh i le t he i s sue c anno t b e a dequa te ly a dd ressed w i th c ur ren t ly a va i lab le d a ta , i ti ss uspec ted t ha t g oa t h e rd ing m ay h ave i n i t ia l ly e vo lved a s ac omp l imen ta ry s ubs is tence s t ra tegy t oc on t inued h un t ing a nd g a the r ing , o ra s a n a d junc t o fo the rp as to ra l o r c e rea l s ubs is tence o p t ions . F or c ompa ra t ive p urposes , F i gure 2 1 d iag rams t he s h i f t ing e ne rge t ic r e la t ionsh ip e xpec ted b e tween mea t p l us m i lk p as to ra l s t ra teg ies a nd c e rea l s t ra teg ies u nder c hang ing t e chno log ica l c ond i t ions . T he r ange o f r e tu rn r a tes d es igna ted " A " a ga in r e f lec ts t he r ange o f r e tu rns e xpec ted b e tween w hea t a nd b a r ley c u l t iva t ion o n a ve rage l a nds u nde r c ond i t ions o f h and t i lage a nd e a r ly l i t h ic t e chno logy . U s ing t he o the rd a ta p rev ious ly p resen ted i nF i gu re 1 8 , t h e r ange o f r e turn r a tes d es igna ted a s " B " s im i la r ly r e f lec ts t he r e tu rns e xpec ted o n a ve rage l a nds u nde rc ond i t ions o fh and t i lage a nd a dvanced l i t h ic t echno logy , w h i le t he r ange o fr e turn r a tes d es igna ted a s " C " r e f lec ts t hose e xpec ted o n a ve rage l a nds u nde r c ond i t ions o fa n ima l t r ac t ion t i lage a nd e a r ly me ta lt e chno logy .

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c 1 000

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F i gure 1 9 . E ne rge t ic r e la t ionsh ips e xpec ted b e tween e a r ly c e rea l s t ra teg ies a nd m ea t -on ly p as to ra l s t ra teg ies . T he r e tu rn r a te c ur ves f o rm ea t -on ly p as to ra l s t ra teg ies a re t hose p rev ious ly p resen ted i nF i gu re 9 .T he r ange o f r e tu rn r a tes d es igna ted a s " A " r ep resen ts t he e ne rge t ic d i v is ion b e tween " op t ima l " a nd " ma rg ina l " a g r icu l tu ra l l a nd u nde rc ond i t ions o fh and t i lage a nd e a r ly l i t h ic t echno logy a s p rev ious ly p resen ted i nF i gu re 1 8 . F o r c ompa ra t ive p urposes , t he p os tencoun te r r e tu rn r a tes f o r e xp lo i t ing w i ld e i nko rn s t ands a s p rev ious ly p resen ted i nT ab le 3 6a re a l so p l o t ted .

1 54

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1 50

200

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300

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NUMBER OF AN I MALS I N HERD F i gu re 2 0 . E ne rge t ic r e la t ionsh ips e xpec ted b e tween e a r ly c e rea l s t ra teg ies a nd mea t p l us m i lk p as to ra l s t ra teg ies . T he r e tu rn r a te c urves f o r mea t p l us m i lk p as to ra l s t ra teg ies a re t hose p rev ious ly p resen ted i nF i gu re 1 2 w hen m i lk p rocess ing i sa ccoun ted f o r . T he r ange o fr e turn r a tes d es igna ted a s " A " r ep resen ts t he e nerge t ic d i v is ion b e tween " op t ima l " a nd " ma rg ina l " a g r icu l tura l l a nd u nde r c ond i t ions o f h and t i lage a nd e a r ly l i t h ic t echno logy a s p rev ious ly p resen ted i n F i gure 1 8 . F o r c ompa ra t ive p u rposes , t he p os tencoun ter r e turn r a tes f o r e xp lo i t ing w i ld e i nko rn s t ands a s p rev ious ly p resen ted i n T ab le 3 6 a re a l so p l o t ted . T he e xpec ted r e turn r a tes f o r w hea t a nd b a r ley y i e ld ing 1 00 k g /du o n " op t ima l " l a nds u nde r c ond i t ions o fh and t i lage a nd e a r ly l i t h ic t e chno logy a re s im i la r ly p resen ted . T hese l a t te rr e turn r a tes r e f lec tt hose w h ich a re e xpec ted f o re a r ly a g r icu l tu re a ts uch f a vo rab le b u t g eog raph ica l ly c i rcumsc r ibed s i tes a sJ e r icho , M u reyb i t , A bu H u rey ra a nd T e l l A swad .

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NUMBER OF AN I MALS I N HERD F i gu re 2 1 . E xpec ted v a r ia t ions i nt he e ne rge t ic r e la t ionsh ips b e tween c erea l s t ra teg ies u nde r c hang ing t echno log ica l c ond i t ions a nd mea t p l us m i lk p as to ra l s t ra teg ies . T he r anges o fr e turn r a tes d es igna ted a s " A , " " B " a nd " C " r e f lec t t he s h i f t ing e ne rge t ic d i v is ion b e tween " op t ima l " a nd " ma rg ina l " a g r icu l tura l l a nd a s t e chno log ica l e f f ic iency i n c reases t h rough t ime ( r e fer t o F i gure 1 8 ) . " A " d es igna tes t h is d i v is ion u nder c ond i t ions o f h and t i lage a nd e a r ly l i t h ic t echno logy . " B " d es igna tes t h is d i v is ion u nde r c ond i t ions o f h and t i lage a nd a dvanced l i t h ic t echno logy . " C " d es igna tes t h is d i v is ion u nde r c ond i t ions o fa n ima l t r ac t ion t i lage a nd e a r ly me ta lt e chno logy . T he r e turn r a te c urves f o r mea tp l us m i lk p as to ra l s t ra teg ies a re t hose w h ich w e re p rev ious ly p resen ted i nF i gu re 1 2w hen m i lk p rocess ing i sa ccoun ted f o r .

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Two mu tua l ly e xc lus ive a nd a rchaeo log ica l ly t es tab le e xpec ta t ions c once rn ing t he s pa t ia l a nd t empora l e vo lu t ion a nd a dop t ion o f p as to ra l a nd c e rea l s ubs is tence o p t ions eme rge f r om t hese e ne rge t ic c ompa r isons . I n i t ia l ly , t he d a ta d iag rammed i nF i gu re 1 9 l e ad t ot he e xpec ta t ion t ha t mea t -on ly p as to ra l s t ra teg ies s hou ld o n ly a ppea ri nt he a rchaeo log ica l r eco rd a f te ra l l a nds o f o p t ima l a nd a ve rage q ua l i ty a re b e ing a g r icu l tura l ly e xp lo i ted , a nd s hou ld a ppea r c on tempo raneous ly w i th , o r s ubsequen t t o , t he e xp lo i ta t ion o f a g r icu l tu ra l ly ma rg ina l l a nds . I f d a i ry ing was n o t s i gn i f ican t i nt he e a r ly e vo lu t ion a nd a dop t ion o fp as to ra l s t ra teg ies , a s s eve ra l s cho la rs h ave ma in ta ined ( e .g . , B ököny i 1 976 ; F agan 1 986 :2 19 ; Ga r ra rd 1 984 ; D . H a r r is 1 977 ; H e r re 1 963 ; L evy 1 983 ; R osen 1 984 , 1 985 , 1 987 ; S he r ra t 1 981 , 1 983 ; Wa tson 1 979 ) , t hen t h is s pa t ia l a nd t empo ra l p red ic t ion s hou ld b e s uppor ted b y t he a rchaeo log ica l r eco rd . T h is e xpec ta t ion i sa l so q u i te c ons is ten t w i th a l te rna t ive p red ic t ive mode ls i n vok ing e co log ica l ma rg ina l i ty a nd p roduc t iv i ty p e ru n i t a rea t os ugges t t ha t p as to ra l s t ra teg ies mus t h ave e vo lved a s a dap ta t ions t o e nv i ronmen ts ma rg ina l f o r a g r icu l ture ( e .g . , Ga r ra rd 1 984 ; K hazanov 1 984 ; S ah l ins 1 968 ; S poone r 1 97 1 , 1 973 ) . B y c on t ras t , t he d a ta d i ag rammed i nF i gu re 2 0 l e ad t ot he e xpec ta t ion t ha t i fd a i ry ing was s i gn i f ican t i nt he e vo lu t ion o fe a r ly p as to ra l s t ra teg ies , c e rea l s t ra teg ies s hou ld s t i l l a ppea r i nt he a rchaeo log ica l r eco rd i n o p t ima l a g r icu l tura l h ab i ta ts p r ior t o t he a ppea rance o f p as to ra l s t ra teg ies , b u t p as to ra l s t ra teg ies s hou ld a ppea rp r io rt ot he a g r icu l tura l e xp lo i ta t ion o fa ve rage a nd ma rg ina l l a nds . F ur the r , t he d a ta d i ag rammed i nF i gu re 2 1 l e ad t ot he e xpec ta t ion t ha t mea t p l us m i lk p as to ra l s t ra teg ies w ou ld o n ly h ave b ecome a dap ta t ions t oe nv i ronmen ts ma rg ina l t o a g r icu l tu re o n a ve rage l a nds w i th t he a dven t o fi n c reas ing ly e f f ic ien t a g r icu l tu ra lt echno log ies . I f d a i ry ing was s i gn i f ican t i nt he e a r ly e vo lu t ion a nd a dop t ion o f p as to ra l s t ra teg ies , a s was s uspec ted o n t he b as is o f t he d a ta o n l a c ta t ion p resen ted i n C hap te r 2a nd a s p rev ious ly s ugges ted b y s eve ra l s cho la rs ( e .g . , B anks 1 984 ; H a fez 1 968a ; Wendor f a nd S ch i ld 1 980a ; Z euner 1 963 ) , t hen t h is a l te rna t ive s pa t ia l a nd t empo ra l e xpec ta t ion s hou ld b e s uppor ted b y t he a rchaeo log ica l r eco rd .

C ompa r isons w i th t he A rchaeo log ica lR ecord Much o f t he a rchaeo log ica l r eco rd r e levan t t ot es t ing t he e xpec ta t ions g ene ra ted a bove h as a l ready b een p resen ted i np rev ious s ec t ions o ft h is manusc r ip t , a nd w i l l n o t b e r epea ted h e re . H oweve r , a s p rev ious ly n o ted , t he e a r l ies t p os tu la ted a ppea rance o f c e rea l c u l t iva t ion o ccu rs i nt he n i n th t oe i gh th m i l lenn ia B .C . a ts uch s i tes a s Mu reyb i t , A bu H u rey ra , T e l l A swad a nd J e r icho u nde r c ond i t ions wh ich c an b e c ons ide red a s a g r icu l tu ra l ly o p t ima l ( Do r re l l 1 978 ; H op f 1 969 ; K enyon 1 970 ; Moo re 1 979 , 1 982 , 1 985 ; S he r ra t 1 980 ; V an Z e is t a nd B akke rH ee res 1 979 ) . P as to ra l s i tes s ubsequen t ly a ppea r i nN o r th A rab ia a nd N o r th A f r ica i nt he e i gh th o rs even th m i l lenn ia B .C . ( Banks 1 984 ; B e t ts 1 982b ; H e lms 1 981 ; I n g raham e ta l . 1 98 1; B e t ts , i nR o l le r 1 984 ; Wendo r f a nd S ch i ld 1 980a ; Wendor f e ta l . 1 984 ) . B yc on t ras t , t he e xp lo i ta t ion o fi n c reas ing ly ma rg ina l a g r icu l tura l a reas t empo ra l ly f o l lowed t he a ppea rance o f p as to ra l s ubs is tence s t ra teg ies . F o r e xamp le , l a te N eo l i th ic a nd H assuna f a rm ing c ommun i t ies d o n o t i n i t ia l ly a ppea r o n t he r o l l ing A ssy r ian p l a ins o f n o r thern I r aq u n t i l t he s i x th m i l lenn ium B .C . ( Moo re 1 985 :56 -58 ) . A t H assuna i t se l f , p e rmanen t v i l lage l i f e may h ave b een p receded b y s easona l p as to ra l o ccupa t ion ( Oa tes 1 972 :30 1-302 ) . B y t he m id -s ix th m i l lenn ium B .C . , H a la f ian s e t t lemen ts a ppea ra c ross t he n or the rn p l a ins o fI r aq s u i tab le f o rd ry f a rm ing ( Moo re 1 985 :55 , 5 8 ) . D u r ing t he s i x th a nd f i th m i l lenn ia B .C . , t he d i s t r ibu t ion o f l a te N eo l i th ic c ommun i t ies i n m uch o f t he L evan t r e f lec ts a n e xpans ion i n to f o res t z ones , w i th a t tendan t a xes a nd a dzes p resumab ly r e f lec t ing t he l a bo r e xpended i n wood land c l ea rance ( Moo re 1 985 :50 -54 ) . Wh i le d eve lopmen ts i n c e rea l c u l t iva t ion a long t he N i le a re p oo r ly u nders tood p r io rt ot he a ppea rance o ft he f u l ly d eve loped N eo l i th ic o f Me r imde a nd F ayum A s i tes n ea rt he e nd o ft he f i f th o ri nt he e a r ly f our th m i l lenn ium B .C . , c e rea l c u l t iva t ion i nt he r es t o f N or th a nd Wes t A f r ica i n i t ia l ly a ppea rs a s a n a d junc t o f e a r ly p as to ra l s t ra teg ies , o r i s s eem ing ly a dop ted b y p rev ious ly p as to ra l p opu la t ions ( Banks 1 984 ; Mc In tosh a nd Mc In tosh 1 98 1 ; Munson 1 976 ; Wendo r f a nd S ch i ld 1 980a ; Wendo r f e ta l . 1 984 ) . S equen t ia l ly , i ts eems t ha t o p t ima l a g r icu l tu ra l a reas w e re i n deed t he i n t ia l f o cus o fh uman f ood p roduc ing e f fo r ts , w h i le p as to ra l s t ra teg ies e vo lved s hor t ly t he rea f te r . S ubsequen t ly , t he e xp lo i ta t ion o fi n c reas ing ly ma rg ina l a g r icu l tu ra l a reas o ccu r red . E ven c u r ren t i n terp re ta t ions o f t he e vo lu t ion o r a dop t ion o ff ood p roduc t ion s t ra teg ies i nt he N egev a nd S ina i , f r equen t ly c i ted i ns uppor t o f al a te d a te f o rd a i ry ing d ue t ot he l a te d a te a t wh ich p as to ra l s t ra teg ies s eem ing ly 1 57

a ppea r ( e .g . , L evy 1 983 ; L evy a nd A lon 1 983 ; R osen 1 984 , 1 985 , 1 987 ) , f i t e xpec ta t ions d e r ived f r om ac ons ide ra t ion o ft he e ne rge t ics o f mea t p l us m i lk p as to ra l s t ra teg ies . I nt he N egev a nd S ina i , h un te r -ga there r e conom ies a re s a id t o h ave c on t inued i n to t he l a te r p e r iods o f t he N eo l i th ic ( e .g . , B a r -Yose f 1 98 1 , 1 984 ; R osen 1 985 , 1 987 ; T che rnov a nd B a r -Yose f 1 982 ) . T he e a r l ies t f ood p roduc t ion i nt hese a reas i sb e l ieved t o h ave i n vo lved a g r icu l tu re a tl a te N eo l i th ic s i tes c en te red u pon s p r ings a l ong t he n o r the rn N egev c oas ta l p la in i n a ssoc ia t ion w i th wa te r r e ten t ive s o i ls ( Levy 1 983 ; L evy a nd A lon 1 983 :107 ) . T he s ubsequen t e as twa rd e xpans ion o f C ha lco l i th ic s i tes i nt he f i th a nd f ou r th m i l lenn ia B .C . i s b e l ieved t o h ave i n vo lved t he d eve lopmen t o f p as to ra l s ubs is tence c oncom i tan t w i th t he a g r icu l tura l e xp lo i ta t ion o ft r ough -shaped v a l leys , u sua l ly a l ong v a l ley ma rg ins i nc l ose p rox im i ty t of l oodp la in d epos i ts i nt he v a l ley b o t toms , a nd i n vo lv ing f l oodwa te r a nd r uno f f i r iga t ion ( Levy 1 983 :24 -25 ; L evy a nd A lon 1 983 :107 ) ) . P resumab ly , a g r icu l tu ra l r e turn r a tes i ns uch r e la t ive ly f avo rab le l o ca t ions wou ld h ave m in ima l ly e qua l led o r e xceeded t hose e xpec ted o n l a nds o f " ave rage " q ua l i ty i nt he N egev g i ven t he h i s tor ic a nd e thnog raph ic y i e ld d a ta d ocumen ted f o r t h is r eg ion ( Maye rson 1 960 :18 , 1 955 :55 ) a s p rev ious ly d i scussed i nC hap te rs 3 a nd 6 . I nt he c en t ra l N egev a nd S ina i , t he p auc i ty o fk nown l a te N eo l i th ic a nd C ha lco l i th ic s i tes r e la t ive t o t he n or the rn N egev i st hough t t o r e f lec t ad emog raph ic d ec l ine i nt hese s ou the rn r eg ions ( Rosen 1 985 , 1 987 ) . H oweve r , o ft he k nown t e rm ina l N eo l i th ic /ea r ly C ha lco l i th ic s i tes i nt he c en t ra l N egev ( Rosen 1 984 , 1 985 , 1 987 ) a nd s i tes o ft he f i th a nd f our th m i l lenn ia i n S ina i ( Ba r -Yose f 1 984 :155 -156 ) , s im i la r a rch i tec tura l p a t te rns o f wha t a ppea r t o h ave b een c en t ra l a n ima l e nc losures w i th a t tached r ooms o rt umu l i a re i n terp re ted a s r e f lec t ing p as to ra l s ubs is tence . D u r ing t he s ubsequen t E a r ly B ronze A ge o ft he m id four th t oe a r ly t h i rd m i l lenn ia , t he n umbe ro fk nown s i tes d rama t ica l ly i n c reases i nt he c en t ra l N egev , w i th a n e conomy b ased u pon p as to ra l ism s upp lemen ted b y v a ry ing d eg rees o fa g r icu l tu ra l a c t iv i t ies s uspec ted ( Rosen 1 985 , 1 987 ) . H ence , a l though s eem ing ly l a gg ing b eh ind d eve lopmen ts i n s ur round ing r eg ions , a n e xpec ted p a t tern f o r t he e vo lu t ion o r a dop t ion o f a l te rna t ive f ood p roduc t ion o p t ions a s d e r ived f r om t he e ne rge t ics o f mea t p l us m i lk p as to ra l s t ra teg ies i sa l so s eem ing ly a t tes ted i nt he N egev a nd S ina i . A c tua l ly ,t h is a rchaeo log ica l r e co rd f r om t he N egev a nd S ina i may mo re c l ose ly ma tch s uch e ne rge t ic e xpec ta t ions t han i tc u r ren t ly d oes u nde r p resen t i n terp re ta t ions . T he s ugges t ion t ha t P re -Po t te ry N eo l i th ic B s ubs is tence i nt hese a reas w as b ased s o le ly u pon h un t ing a nd g a the r ing r es ts h eav i ly u pon t he a ssumed a bsence o f d omes t ica te s pec ies i n t he f auna l d epos i ts r ecove red f r om s i tes o ft h is p er iod , whe re i b ex i ss a id t o b e w e l l -a t tes ted . H oweve r , g i ven t he k nown d i f f icu l t ies i no s teo log ica l ly d i s t ingu ish ing i b ex ( Cap ra i b ex ) f r om g oa t ( Capra a egagrus ) e xcep t t h rough ma le h o rn c ores ( Ba r -Yose f 1 98 1:225 , 2 26 , P ayne 1 972a :79 -80 ; Sm i th e ta l . 1 984 :472 ; T che rnov a nd B a r -Yose f 1 982 :23 ) , t he i d en t i f ica t ion o f " i bex " a tN aha l D ivshon i n t he N egev ( Serve l lo 1 976 :351; T che rnov 1 976 :69 ) a nd t he p reponde rance o f " i bex " a s i d en t i f ied i nt he f auna l d epos i ts a t Wad i T be ik a nd o the rc on tempo raneous s i tes i nt he s ou the rn S ina i ( Ba r -Yose f 1 981 :225 -226 , 1 984 :153 ; T che rnov a nd B a r -Yose f 1 982 :23 -24 , 3 3 3 4 ) d o n o t a low f o rt he b road a sser t ion t ha t d omes t ica te s pec ies we re d e f in i te ly n o t h e rded i n t hese r eg ions d u r ing t he P re -Po t te ry N eo l i th ic B p e r iod . C oup led w i th t he f r equen t o ccu r rence o f m i l l ing s t ones a nd g ra in s t o rage f ac i l i t ies ( Ba r -Yose f 1 98 1 :224 -226 , 1 984 :148 -155 ; R osen 1 985 , 1 987 ; T chernov a nd B a r -Yose f 1 982 :19 ) , am ixed s ubs is tence i n vo lv ing g oa t h e rd ing s upp lemen ted b y h un t ing a nd w i ld c e rea l e xp lo i ta t ion o rc e rea l h usbandry c an b e a l te rna t ive ly p os i ted . A s s uch , t he e vo lu t ion o f f ood p roduc t ion s t ra teg ies i nt he N egev a nd S ina i wou ld mo re c l ose ly t r ack r ough ly c on tempo raneous s ubs is tence s h i f ts e l sewhe re i nt he N ea r E as t a nd N o r th A f r ica . I t wou ld t here fo re a ppea r t ha t e nerge t ic e xpec ta t ions i n vo lv ing mea t -on ly p as tora l s t ra teg ies f a i l t o a dequa te ly a ccoun t f o rt he emp i r ica l d a ta , a s wou ld p red ic t ive mode ls wh ich s ugges t t ha t p as to ra l ism n ecessa re e vo lved a s a n a dap ta t ion t o e nv i ronmen ts ma rg ina l f o r a g r icu l tu re . B y c on t ras t , e ne rge t ic e xpec ta t ions i n vo lv ing mea t p l us m i lk p as to ra l s t ra teg ies wou ld s eem ing ly a ccoun t f o r b o th t he s pa t ia l a nd t empo ra l a ppea rance o fp as tora l a nd c erea l f ood p roduc t ion s t ra teg ies i nt he a rchaeo log ica l r eco rds o ft he N ea r E as t a nd N or th A f r ica . H ence , t he p robab le s i gn i f icance o f d a i ry ing i n t he e vo lu t ion o f e a r ly p as tora l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts s eems s us ta ined , w h i le t he c ommon p e rcep t ion o f p as to ra l s t ra teg ies a s a dap ta t ions t oe nv i ronmen ts ma rg ina l f o r a g r icu l tu re s eems u n l ike ly . A t b es t , e a r ly p as to ra l s t ra teg ies w e re o n ly ma rg ina lt oa g r icu l tu re p rac t iced i ne x t reme ly f a vo rab le

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l o ca l i t ies , a nd w ou ld n o t h ave b ecome ma rg ina l t ol e ss f a vo rab le a g r icu l tu ra l o p t ions u n t i l t he a dven to fi n c reas ing ly e f f ic ien t a g r icu l tu ra l t echn iques .

C onc lus ions I t was t he e xp l ic i t p u rpose o f t h is manusc r ip t t o p resen t a n a pp l ica t ion o f b ehav io ra l e co logy t o e thnog raph ic a nd a rchaeo log ica l d a ta i n a n a t temp t t o f u r ther o ur c ur ren t u nde rs tand ing o ft he e vo lu t ion a nd a dop t ion o f a l te rna t ive f ood p roduc t ion s t ra teg ies i nt he N ea r E as t a nd N or th A f r ica d ur ing t he H o locene e poch . T o t h is e nd , a me thodo logy was d eve loped f o r a pp ly ing e vo lu t ionar y t heory t o a n u nders tand ing o f w hy p a t te rns o f e a r ly h uman f ood p roduc t ion t ook t h ef o rm t hey d i d , w hen a nd w he re t hey d i d . C os t a nd b ene f i t d a ta w e re g ene ra ted f o r a n ima l a nd c e rea l h usband ry s t ra teg ies i n af a sh ion c ons is ten t w i th t ha t emp loyed i nr e cen t e vo lu t iona ry s t ud ies o f h uman f o rag ing b ehav io r . E ne rge t ic e xpec ta t ions d e r ived f r om t hese d a ta w e re t hen u sed t o e xp lo re t he e x ten t t o w h ich s imp le o p t im iza t ion mode ls m igh t e xp la in t he d ec is ion mak ing a ssoc ia ted w i th t h e s pa t ia l a nd t empo ra l a ppea rance o fa l te rna t ive p as to ra l a nd a g r icu l tu ra l s ubs is tence o p t ions a s r e f lec ted i nt he H o locene e poch a rchaeo log ica l r e co rds o ft h eN ea rE as ta nd N o r th A f r ica . U s ing r anges o fp oss ib le r e tu rn r a tes f o rp as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts , a nd a g r icu l tu ra l s t ra teg ies i n vo lv ing w hea ta nd b a r ley ,t h ree p a r t icu la ri s sues r e levan t t o o ur p resen t u nders tand ing o f t he e vo lu t ion a nd a dop t ion o f p as to ra l a nd a g r icu l tura l s ubs is tence s t ra teg ies i nt h e N ea r E as t a nd N or th A f r ica w e re s pec i f ica l ly a dd ressed . T hese i s sues w e re : ( 1 ) I si tr easonab le t o a ssume t ha t p as to ra l s t ra teg ies n ecessa r i ly e vo lved a s a dap ta t ions t o e nv i ronmen ts ma rg ina l t o a g r icu l ture? , ( 2 ) Wou ld m i lk p roduc t ion h ave b een s i gn i f ican t i nt he e vo lu t ion o fe a r ly p as to ra l s t ra teg ies? , a nd ( 3 ) Why w e re p as to ra l s t ra teg ies i n vo lv ing c ame ls a dop ted l a te rt han o t he rp as to ra l a nd c e rea ls ubs is tence o p t ions? T he r e su l ts i n i t ia l ly s ugges tt ha t i ti su n l ike ly t ha tp as to ra l s t ra teg ies e vo lved a s a dap ta t ions t o e nv i ronmen ts ma rg ina l f o r a g r icu l tu re . B o th t heo re t ica l c ons ide ra t ions a nd emp i r ica l d a ta i n d ica te t ha t p as to ra l s t ra teg ies e vo lved p r ior t ot he a g r icu l tu ra l e xp lo i ta t ion o f ma rg ina l d ry f a rm ing l a nds . A t b es t , p as to ra l s t ra teg ies w e re i n i t ia l ly ma rg ina l t o c u l t iva t ion s t ra teg ies i n o p t ima l a g r icu l tura l a reas , b u t c er ta in ly n o t a g r icu l tura l s t ra teg ies i ng ene ra l . On ly w i th t he a dven t o fi n c reas ing ly e f f ic ien t a g r icu l tu ra l t echno log ies i n vo lv ing a n ima l t r ac t ion t i lage w ou ld p as to ra l s t ra teg ies h ave b ecome ma rg ina l t o mos t a g r icu l tura l o p t ions . On t h is i s sue , t he e vo lu t iona ry a pp roach a dop ted h e re h as l e d t o t he s ubs tan t ive r eeva lua t ion o f ac ommon ly a ccep ted i n terp re ta t ion o ft he s equence b y wh ich f ood p roduc t ion s t ra teg ies e vo lved . T he r esu l ts o ft h is i n ves t iga t ion f u r the r s ugges t t ha t t he re i sg ood r eason t o b e l ieve t ha t d a i ry ing w ou ld h ave b een s i gn i f ican t i nt he e vo lu t ion a nd a dop t ion o f p as to ra l s t ra teg ies i n vo lv ing c ame ls , c a t t le , s heep a nd g oa ts . T he e ne rgy r e turns f o r mea t -on ly p as to ra l s t ra teg ies a re s o l ow a st op rec lude t he e vo lu t ion o fp as to ra l s ubs is tence b ased u pon t hese s pec ies u n t i l a l a gr icu l tu ra l o p t ions w e re f i led , as i tua t ion c on t ra ry t ot he emp i r ica l d a ta . On t h is i s sue , t he e vo lu t iona ry a pp roach a dop ted h ere h as a lowed f o r t he s ubs tan t ive i n ves t iga t ion o f a n o the rw ise a rchaeo log ica l ly i n tang ib le q ues t ion . F i na l ly , t he l a te a dop t ion o fp as to ra l s t ra teg ies i n vo lv ing c ame ls w ou ld s eem ing ly make s ense i n l i gh t o f c on tempo raneous e nv i ronmen ta l c hanges . Wh i le p rev ious ly p e rce ived a s p rov id ing a n e xce l len t p a rad igm f o r p as to ra l ism a s a n a dap ta t ion t o ma rg ina l e nv i ronmen ts , i t w as a c tua l ly t he d eve lopmen to f" ma rg ina l " e nv i ronmen ts w h ich made t he c ompa ra t ive ly m o re p ro f i tab le c ame l s t ra teg ies p oss ib le . O n t h is i s sue , t he e vo lu t iona ry p e rspec t ive a dop ted h e re h as a lowed f o rc oun ter in tu i t ive e xpec ta t ions w h ich b e t te ra ccoun tf o rt he emp i r ica l d a ta . I n a dd i t ion t o t hese i n s igh ts , , s eve ra l a dd i t iona l f ac to rs a ppa ren t ly s i gn i f ican t i nt he e vo lu t ion o fe a r ly f ood p roduc t ion s t ra teg ies h ave a l so b een i d en t i f ied d u r ing t he c ourse o ft h is w ork . F o r p as to ra l s t ra teg ies , t h e h uman a b i l i ty t oe xp lo i t a nd man ipu la te e x tan t p hys io log ica l a nd b ehav io ra lt r a i ts among s pec i f ic h e rd s pec ies w as s eem ing ly c r i t ica l i nt h e i n i t ia lc ap tu re a nd s ubsequen t c on t ro l o fw i ld p opu la t ions . F ur the r , t hese t r a i ts p resumab ly a f fec ted t he c ho ice o f b as ing p as tora l s t ra teg ies o n s pec i f ic h e rd s pec ies t ot he e xc lus ion o fo the rs i n wh ich s uch t r a i ts w ere e i the ra bsen t o r l e ss p ronounced . T he p robab le u se o fg ame d r ive l i nes a nd c o r ra ls i nc ap tu r ing a nd s e lec t ing f ound ing p opu la t ions f o r p as to ra l h e rds h as a l so b een p roposed a s a n a l te rna t ive t o t he p opu la r n o t ion t ha t p as to ra l h e rds n ecessa r i ly d eve loped f r om " pe ts " 1 59

ma in ta ined b y a g r icu l tu ra l is ts . T he a ssump t ion t ha t c a t t le d omes t ica t ion n ecessa r i ly f o l lowed s ma l ls tock d omes t ica t ion h as s im i la r ly b een q ues t ioned . I nl i gh t o ft he h i gh r e tu rn r a tes e xpec ted f o rc u l t iva t ion s t ra teg ies i nc e r ta in c i rcumsc r ibed e nv i ronmen ts , i th as b een s ugges ted t ha t t e r r i to r ia l i ty a nd i n c reas ing s oc ia l c omp lex i ty among a g r icu l tura l p opu la t ions e xp lo i t ing s uch l o ca t ions s hou ld b e e xpec ted . W i th p o ten t ia l ly h i gh r e turn r a tes r e la t ive t oa l a l te rna t ive f ood p roduc t ion o p t ions a nd t he c oncom i tan t a t t r ibu tes o f g eog raph ic c i rcumsc r ip t ion a nd t empo ra l ly p red ic tab le s t o res o ff ood , t he b ene f i ts d e r ived f r om p e rmanen t ly o ccupy ing a nd e xp lo i t ing s uch e nv i ronmen ts a re e xpec ted t o h ave o u twe ighed t he s oc ia l c os ts o ft he i r e xc lus ive u se a nd d e fense . When v i ewed f r om t h is p e rspec t ive , t he p rev ious a noma ly o fJ e r icho 's e a r ly w a l ls f i na l ly makes s ense . F i na l ly , i th as b een s ugges ted t ha t mean f o rag ing r e tu rn r a tes i nt he N ea r E as t a nd N o r th A f r ica p r io r t o t he e vo lu t ion o ff ood p roduc t ion s t ra teg ies m us t h ave b een q u i te h i gh . I nt h is r espec t , i th as b een s ugges ted t ha t t he p os t -P le is tocene v a r iab i l i ty a nd e ven tua l d e ter io ra t ion o f h i gh b i omass s avanna e nv i ronmen ts i nt he A rab ian P en insu la a nd N or th A f r ica w e re s i gn i f ican t e co log ica l f a c to rs i nt he s pa t ia l a nd t empo ra l e vo lu t ion a nd a dop t ion o f f ood p roduc t ion s t ra teg ies i nt hese r eg ions . I np a r t icu la r , i ts eems l i ke ly t ha t ab e t te r u nde rs tand ing o fh uman , a n ima l a nd p l an t r e sponses t op os t -P le is tocene e nv i ronmen ta l s h i f ts o n t he A rab ian P en insu la a nd N o r th A f r ica i sc r i t ica l f o r ab e t te ru nde rs tand ing o ft he E p ipa leo l i th ic , N eo l i th ic , C ha lco l i th ic a nd E a r ly B ronze A ge s equences i nt he e n t i re " Fer t i le C rescen t . " T he a ppa ren t a ssoc ia t ion b e tween i n c reased p opu la t ion d ens i t ies a nd t h e e vo lu t ion o fi n c reas ing ly c omp lex s oc ie t ies i n t he " Fer t i le C rescen t " c o inc iden t w i th t he o nse t o f modern e nv i ronmen ta l c ond i t ions i nt h e N ea r E as t a nd N o r th A f r ica a l so w a r ran ts f u r the ri n ves t iga t ion . I n ab roade r c on tex t , t he i n i t ia l p rem ise t ha t s i gn i f ican t a dvances i nu nde rs tand ing t he e vo lu t ion a nd a dop t ion o f f ood p roduc t ion s t ra teg ies a re u n l ike ly u n less man -an ima l -p lan t r e la t ionsh ips a re i n ves t iga ted w i th in t he c on tex t o f g ene ra l t heo r ies wh ich a ddress t he u nder ly ing p r inc ip les a f fec t ing s ubs is tence b ehav io r a l so s eems u phe ld . I ft he u t i l i ty o f a t heo re t ica l a pp roach t ob ehav io ra l i s sues i st ob ej u dged b y i t sa b i l i ty t og enera te u se fu l b od ies o f q uan t i ta t ive d a ta , t o b r ing i n to q ues t ion p rev ious ly h e ld i n terp re t ive a ssump t ions , t o o f fe r c oun te r in tu i t ive e xpec ta t ions , t os ugges t n ew d i rec t ions f o rs ubs tan t ive r e sea rch , a nd t ob e t te r a ccoun t f o r t he a va i lab le emp i r ica l d a ta t han a l te rna t ive a pp roaches , t hen t he e vo lu t ionar y p e rspec t ive a dop ted h e re mus t b e v i ewed a s e x t reme ly u se fu l . T h is a pp roach a l so e xpands t he r ange o fp o ten t ia l a pp l ica t ions a nd t e s ts o fs imp le e vo lu t iona ry c os t -bene f i t mode ls w h ich c an c on t r ibu te t o e xp lana t ions o fs ubs is tence b ehav io r i n aw ide v a r ie ty o f h uman s i tua t ions a nd e nv i ronmen ts . T he p resen t s t udy t he re fo re p rov ides o ne mo re l i nk i nt he d eve lop ing s yn thes is o f mode rn e vo lu t iona ry b i o logy a nd c u l ture t heo ry .

1 60

APPEND IX A

DROMEDARY CAMEL POPULAT ION D YNAM ICS AND S LAUGHTER OFFTAKE F ROM A MODEL HERD OVER A ONE YEAR P ER IOD

I n t roduc t ion T he a ppea rance o f c ame l p as to ra l ism i nt he N ea r E as t a nd N o r th A f r ica s ubsequen t t o a l te rna t ive p as to ra l a nd c e rea l f ood p roduc t ion o p t ions makes a n a ccu ra te u nde rs tand ing o ft he c os ts a nd b ene f i ts o fc ame l h e rd ing i mpe ra t ive f o r e ne rge t ic a na lyses . C u r ren t mode ls o fc ame l h e rd g row th d ynam ics a nd p roduc t iv i ty a s d eve loped b y D ah l a nd H jo r t ( 1976 :78 -86 , 1 84 -98 ) a nd S a to ( 1980 :28 -3 1) a re d eemed i n adequa te f o ru se i ne s tab l ish ing r ea l is t ic b ene f i ts d a ta f o r c ame l h usbandry i n g ene ra l d ue t o p rob lems i n t he c ame l r ep roduc t ion , mor ta l i ty , a nd p roduc t iv i ty d a ta w h ich t hey emp loyed . I nt h is a ppend ix , r ev ised mode ls o fd romeda ry c ame l h e rd g row th d ynam ics a re g ene ra ted t h rough a n a na lys is o f a va i lab le d a ta o n c ame l r ep roduc t ion a nd mor ta l i ty . B ased u pon t he r esu l ts o fp opu la t ion g row th s imu la t ions f o rt he ma le a nd f ema le c omponen ts o f ad romeda ry h e rd u s ing t h is r ep roduc t ion a nd mo r ta l i ty d a ta , a mode l c ame l h e rd i sg ene ra ted f r om wh ich s l augh te r p roduc t iv i ty i sc a lcu la ted o ve r ao ne y ea r p e r iod . I no rde r t o f ac i l i ta te t he e ne rge t ic a na lys is o f mea t -on ly a nd m ea t p l us m i lk p as to ra l s t ra teg ies , s epa ra te c a lcu la t ions f o r" no rma l " a nd " max imum " o f f take ( s laugh te r y i e ld ) o ve r ao ne y ea r p e r iod a re p resen ted . T hese o f f take d a ta a re t hen u sed i nA ppend ix B ,w here t he t o ta l c a lo r ic p roduc t iv i ty o fb o th s l augh te r a nd m i lk p roduc t ion f r om t he mode l h e rd i sd e te rm ined .

R ep roduc t ion i nD romeda ry C ame ls Gene ra lC ons ide ra t ions P opu la t ion g row th i n ap as to ra l h e rd e xpe r ienc ing n o em ig ra t ion,o r i mm ig ra t ion i st he d i f fe rence t h rough t i me b e tween t he b i r th r a te a nd t he d ea th r a te . T he b i r th r a te o f ap as to ra l h e rd i sp r ima r i ly af unc t ion o ft he s pec ies ' g es ta t ion l e ng th , b reed ing f r equency , a ge a tf i rs t p ar tu r i t ion ( b ir th ing ) , y ean ing r a te ( t he n umbe r o f a nnua l b i r ths p e r r ep roduc t ive ly ma ture f ema le ) , a nd h uman i n ter fe rence i nt he i r n a tu ra l b reed ing p rocess . T he f o l low ing s ec t ions p resen t d a ta r e levan t t o q uan t i fy ing s uch v a r iab les f o r t he r ep roduc t ion o f d romeda ry c ame l h e rds ma in ta ined u nde r c ond i t ions o f e x tens ive h usbandry . T he r ep roduc t ive v a lues u sed i n t he p resen t mode ls o fd romedary h e rd g row th p r ima r i ly v ary f r om t hose o fp r io r mode ls i nt he y ean ing r a tes emp loyed .

Ges ta t ion P er iod A s n o ted b y N ovoa ( 1970 :12 ) , c ame l g es ta t ion l e ng ths a re d i f f icu l t t o p rec ise ly e s tab l ish s i nce t he n umbe r o f c opu la t ions d ur ing t he ma t ing s eason o f ten o bscure t he e xac t t ime o f f e r t i l iza t ion . F igures f o rt he g es ta t ion p e r iod i nc ame ls g ene ra l ly v ar y f r om 1 t o 1 3 mon ths . A l i s t o fp ub l ished g es ta t ion l e ng ths i sp resen ted i nT ab le 4 2 . I n aq uan t i ta t ive s t udy o f3 3 B ikane r i c ame ls a tt he Gove rnmen t C ame l B reed ing F a rm , B ikane r , I n d ia , t he mean l e ng th o fg es ta t ion was f ound t o b e 3 89 .87 + 2 . 1 d ays ( Meh ta e ta l . 1 962 ) . E ach f ema le i nt h is s t udy was s e rv iced o n ly o nce , a nd n o s i gn i f ican t v a r ia t ions i nt he g es ta t ion p e r iod w e re n o ted a s ar e su l t o ft he s ex o ft he c a l f , t he s equence o fc a lv ing , o rt he mon th o fp ar tur i t ion .

T ab le 4 2 R epor ted G es ta t ion P e r iods i nt h e C ame l

S PEC IES

C . d romedar ius

C .b ac t r ianus

GESTAT ION P ER IOD

SOURCE

1 2 mon tns 1 2 13 mon ths 1 2 mon ths 1 3 mon ths 1 mon ths 1 2 mon ths 1 2 mon ths 1 2 mon ths 3 15 -360 d ays 1 3 mon ths 1 3 mon ths 3 89 d ays 1 2 13 mon ths 1 2 mon ths 113 mon ths 1 2 mon ths 1 3 mon ths 1 2 13 mon ths 1 2 13 mon ths 1 3 mon ths 1 2 mon ths 1 mon ths 1 2 -13 mon ths 3 15 -360 d ays 3 70 d ays 1 2 13 mon ths

A sad 1 964 :5 1 , 1 970 :15 B ehnke 1 980 :31 B odenhe ime r 1 935 :126 , 1 954 :162 B r i t ish N ava l I n te l l igence 1 946 :507 C lu t ton -B rock 1 981:124 C o le 1 975 :37 D ickson 1 951 :4 13 D ough ty 1 936 :26 1 F ow le r 1 978 :791 G au th ie r -P i l te rs a nd D agg 1 981 :92 J enn ings -B ram ley 1 906 :202 K lös a nd L ang 1 982 :224 K öh le r 1 984 :202 L eese 1 927 :95 L eona rd 1 894 :119 M us i l 1 928 :333 P h i lby 1 922 :130 R ao e ta l . 1 970 :72 S a to 1 980 :25 S pence r 1 973 :11 S . Sm i th 1 980 :469 Swee t 1 970 :159 T o ny 1 973 :68 , 1 974 :67 Wa l lach a nd B oeve r 1 983 :3 18 W i l l iamson a nd P ayne 1 959 :322 Y as in a nd Wah id 1 957 :293

4 06 -408 d ays 1 3 mon ths 3 89 -406 d ays 4 06 d ays 3 64 -392 d ays 3 89 -406 d ays

B a rm incev , i nN ovoa 1 970 :12 C lu t ton -B rock 1 981:124 F ow le r 1 978 :7Q1 K lös a nd L ang 1 982 :224 P odbe rezk in , i nN ovoa 1 970 :13 Wa l lach a nd B oeve r 1 983 :3 18

T he B i kane r i c ame ls , h oweve r , w e re k ep t u nde r h yg ien ic c ond i t ions a nd we re s t a l l f ed w i th ac oncen t ra te a nd r oughage r a t ion i na dd i t ion t ob e ing a lowed t og raze s i x t o e i gh t h ou rs e ach d ay . A s n o ted b y L eese ( 1927 :95 ) , c ame ls may c a r ry t he i rc a lves s l igh t ly l o nge rw hen g raz ing i s p l en t i fu l . A s w i th p r io r mode ls , a1 2 mon th g es ta t ion p e r iod w i l l t he re fore b e a ssumed f o r d romeda ry c ame ls ma in ta ined u nde rc ond i t ions o fe x tens ive h usband ry .

B reed ing F requency I ti sw ide ly a g reed t ha t f ema le c ame ls w i l l n o rma l ly g i ve b i r th t oo ne c a l f e ve ry t wo t ot h ree y ea rs ( Bha rgava e t a l . 1 963 :187 ; B odenhe ime r 1 954 :162 ; B u l l ie t 1 975 :265 ; D ah l a nd H jor t 1 976 :80 ; G au th ier -P i l te rs a nd D agg 1 98 1:94 ; K öh le r 1 984 :202 ; L eese 1 927 :95 -96 ; L eona rd 1 894 :119 121 ; N ovoa 1 970 :5 ; W i l l iamson a nd P ayne 1 959 :322 ; Y as in a nd Wah id 1 957 :293 ) . H oweve r , a ccord ing t o s t ud ies b y Y uz l ikaev a nd A khmed iev ( 1965 :62 ) , 7 5% o f f ema le d romeda r ies a nd 5 4 .5 % o fB ac t r ian f ema les w i l l c ome i n to h ea t s hor t ly a f te rc a lv ing . A cco rd ing t os eve ra l o the r s t ud ies s u mma r ized b y S ha lash ( 1965 : 107 ) , t h is u sua l ly o ccu rs w i th in 1 4t o2 5 d ays a f te rp ar tur i t ion . I ti st he re fo re p oss ib le t o man ipu la te f ema le c ame ls t o c a l f t w ice w i th in a t wo a nd o ne h a l f t o t h ree y ea r p e r iod ( Gau th ie r -P i l te rs a nd D agg 1 976 :94 ; K noess 1 977 :44 ; W i l l iamson a nd P ayne 1 959 :322 ; Y as in a nd Wah id 1 957 :293 ) . 1 62

H oweve r , t he n ecessa ry c ond i t ions f o r a ch iev ing s uch f r equen t b reed ing w ou ld r a re ly b e o b ta ined u nde rt r ad i t iona l s ys tems o fe x tens ive h usband ry . L eese s umma r ized t he s i tua t ion a s f o l lows ( 1927 :96 ) : T ha t ac ow s hou ld , n o rma l ly , g o emp ty e ach a l te rna te y ea r s eems t o b e a p rov is ion o f n a ture t o p ro tec t h e r f r om a s t ra in s he c ou ld n o t e ndure u nde r o rd ina ry d eser t c ond i t ions ; ac ow w h ich h as c a lved i nt wo c onsecu t ive y ea rs i s g ene ra l ly p re t ty p oo r a tt he e nd o fi t . E ven u nde r c ond i t ions o f a r t i f ic ia l f eed ing c omb ined w i th g ood g raz ing , L eese ( 1927 :96 ) n o ted t ha t af o rced i n c rease i nt he b reed ing f r equency l e f t c onsecu t ive c a lves w i th l e ss m i lk t han t hey wou ld h ave o b ta ined u nde r n o rma l c ond i t ions , a nd f u r ther s ugges ted t ha t s uch " t r icks " s hou ld o n ly b e p rac t iced o nce o rt w ice i nt he l i f e t ime o f ar ep roduc t ive f ema le . F o rt hese r easons , i ti se xpec ted t ha t t he n a tu ra l a b i l i ty o ff ema le c ame ls t o b reed mo re f r equen t ly t han o nce e very t wo y ea rs w ou ld n o rma l ly b e p roh ib i ted u nde rc ond i t ions o fe x tens ive h usband ry .

S easona l i ty o fB reed ing B o th ma le a nd f ema le c ame ls e xh ib i t ap ronounced b reed ing s eason wh ich , a l though v ary ing b y e nv i ronmen ta l l o ca l i ty , i sg ene ra l ly r e s t r ic ted t ot he mon ths o f S ep tembe r t h rough J une . S tud ies o n t he r ep roduc t ive e ndoc r ine c yc le o f ma le c ame ls i nt he N egev D eser t , I s rae l , i n d ica te a p e r iod o f i n c reas ing g e rm c e l l a c t iv i ty b e tween O c tobe r a nd J anua ry , w i th ap eak i n a c t iv i ty b e tween F eb rua ry a nd Ma rch ( Bodenhe ime r 1 954 :162 -163 ) . As im i la r s t udy o fo va r ian f unc t ions i nf ema le d romeda r ies a tC a i ro b y S ha lash ( 1965 :106 ) s ugges ts t ha t t he ma jo r i ty o f o va r ian a c t iv i ty o ccu rs f r om D ecembe rt h rough May , a l though o p t imum a c t iv i ty o ccu r red d u r ing Ma rch w i th ad rop i na c t iv i ty d u r ing F eb rua ry . T h is ma rked s easona l i ty o f t he c ame l 's b reed ing c yc le c o inc ides w i th t he s eason o f o p t imum v ege ta t ion g row th o n d ese r t r anges . C oup led w i th a 1 2 mon th g es ta t ion p e r iod , t he re h as a ppa ren t ly b een s t rong s e lec t ion among c ame ls f o r t he c oo rd ina t ion o f t he i r b reed ing p e r iod a nd p ar tur i t ion w i th max imum s easona l r esou rce a va i lab i l i ty . A s d i scussed b y N ovoa ( 1970 :3 ) : T he p a t tern o f t he r ep roduc t ive c yc le a ppea rs t o r e la te t o t he h a rsh e nv i ronmen t i nw h ich t he c ame l p rogen i to rs e vo lved . On ly c a lves b o rn i na r es t r ic ted p e r iod o ft he y ea r h ad a ny c hance o f s urv iva l . S e lec t ion h as , t hus , b een t owa rds at y pe o fa n ima lw i th ab reed ing s eason c o inc id ing w i th t he mos t f avo rab le t ime o ft h e y ea r . I na reas w i th b e t te re nv i ronmen ta lc ond i t ions , s exua l a c t iv i ty d oes n o tf o l low s uch ar e s t r ic ted p a t te rn . H ence , t he ma t ing s eason o fd romeda r ies i nN or th A f r ica g ene ra l ly o ccu rs b e tween D ecembe r a nd M ay ; i nI s rae l f r om J anua ry t o Ma rch ; i nP ak is tan f r om D ecembe rt h rough Ma rch ; i nI n d ia f r om N ovembe r t o F eb rua ry ; a nd i nS oma l i land f r om A ugus t t o May ( Bodenhe ime r 1 954 :162 ; Gau th ie r -P i l ters a nd D agg 1 981 :92 -93 ; N ovoa 1 970 :6 ; Y as in a nd Wah id 1 957 :293 ) . A cco rd ing t o L eese ( 1927 :89 ) , t he b reed ing s eason i nt he S aha ra e x tends f r om S ep tember t h rough J anua ry , wh i le i nS oma l i land i tg ene ra l ly o ccu rs e i the r i nJ une o r f r om S ep tembe r t h rough N ovember , t he t wo r a iny s easons o f E as t A f r ica . H oweve r , a s n o ted b y S a to ( 1980 :26 ) , t he ma rked s easona l i ty o fc ame l b reed ing i ss t i l l a ppa ren t i n mo re s ou the r ly l a t i tudes , s i nce o n ly a f ew c a lves ( " none o ro ne p e rh e rd " ) a re a c tua l ly b o rn i nK enya d u r ing t he s hor te ra u tumn r a iny s eason . S im i la r ly , t he b reed ing s eason o fw i ld B ac t r ian c ame ls i nt he Gob i D eser t was o bserved t o o ccu r i nJ anua ry a nd F eb rua ry , wh i le aw in te r o r s p r ing s eason h as b een n o ted f o r B ac t r ian c ame ls i nb o th Mongo l ia a nd i nz oos i nt he N or the rn H em isphe re ( Bu l l ie t 1 975 :285 ; Gau th ie r P i l te rs a nd D agg 1 981:93 ; N ovoa 1 970 :6 -7 ) . A ccord ing t o Y uz l ikaev a nd A khmed iev ( 1965 :62 ) , t he n o rma l b reed ing s eason o f c ame ls i nC en t ra l A s ia i sf r om J anua ry t h rough Ma rch . H oweve r , t he p rox ima te mechan isms i n f luenc ing t he s easona l i ty a nd d ura t ion o f s exua l a c t iv i ty i nt he C ame l idae r ema in p oo r ly u nde rs tood . W i l l iamson a nd P ayne ( 1959 :322 ) h ave s ugges ted t ha t t he r es t r ic ted r u t t ing s eason o f ma le c ame ls i sa ssoc ia ted w i th l e ng then ing 1 63

d ays ; b e tween J anua ry a nd Ma rch i nt he N o r the rn H em isphe re , a nd J une t h rough S ep tembe r i nt he S ou the rn H em isphe re . S easona l t empe ra tu re , h um id i ty , a nd r a in fa l l v a r ia t ions h ave a l so b een s ugges ted a s s i gn i f ican t p rox ima te f ac to rs i nt he c ame l 's b reed ing s eason ( Bha rgava e t a l . 1 963 :187 ; B u l l ie t 1 975 :4 1-42 , 2 85 ; D ah l a nd H jo r t 1 976 :80 ; L eese 1 927 :99 ; T o r ry 1 973 :68 ) .

H uman I n ter fe rence i nt he B reed ing C yc le T he p r inc ip le mo t iva t ion f o r p as to ra l is ts t o mod i fy o rr e s t r ic tt he n a tu ra l b reed ing s eason o f t he i r h e rd a n ima ls wou ld s eem t ob e t he c oo rd ina t ion o fp a r tu r i t ion w i th a va i lab le p as tu rage i n s uch a f ash ion a s t o max im ize m i lk p roduc t ion a s w e l l a s t he v i ab i l i ty a nd g row th o f y oung a n ima ls . F ur the r , i tw ou ld s eem g ene ra l ly a dvan tageous t os chedu le t he b reed ing s eason s o a s t oa vo id i n ter fe rence d ur ing p e r iods o fm ig ra t ion . A cco rd ing t o C o le ( 1975 :37 , 4 9 ) , t he A l -Mu r rah b edou in o f S aud i A rab ia b reed t he i r c ame ls i nt he l a te F a l l a nd e a r ly W in te r , w h i le b o th t he Rwa la b edou in o fN or th A rab ia ( Lancas te r 1 981:411; Mus i l 1 927 :453 , 1 928 :332 , 6 00 ) a nd t he M u ta i r b edou in o f K uwa i t ( D ickson 1 951 :4 11) b reed t he i r c ame ls i nD ecembe r a nd J anua ry . I nt hese c ases , t he c oo rd ina t ion o f p ar tur i t ion w i th o p t imum g raz ing p o ten t ia l b e fo re o r a f te r m ig ra t ion t o t r ad i t iona l w in te r r ange lands i sa ppa ren t . A s n o ted e a r l ie r , t he a b i l i ty o ff ema le c ame ls t ob reed m o re f r equen t ly t han o nce e very t wo y ea rs w ou ld g ene ra l ly b e d e le te r ious t o t he i r h ea l th u nde r n orma l c ond i t ions o f e x tens ive h usband ry , r esu l t ing i n d ec reased m i lk p roduc t ion a nd l e ss -v iab le o f fsp r ing . P as tora l is ts a ppa ren t ly c on t ro l t he r ep roduc t ion o f i n d iv idua l f ema les a nd c oo rd ina te t he i r p a r tu r i t ion w i th a va i lab le p as tu rage b y s easona l ly s ubd iv id ing t he i r h e rds , e f fec t ive ly i s o la t ing f ema les w i th n ew ly -bo rn y oung f r om a ccess b y t he s t ud c ame l o r c ame ls ( Co le 1 975 :36 -37 ; D ickson 1 951:4 11 , 4 19 ; Gau th ie r -P i l te rs a nd D agg 1 981:80 ) . H oweve r , t hese s easona l s ubd iv is ions i n c rease t he t o ta l n umbe ro fh e rde rs r equ i red p e rh e rd , a nd t he re fo re i n c rease t he a nnua l l a bo r c os ts o fc ame l p as to ra l ism .

P ube r ty a nd F e r t i le L i fespan P uber ty i nc ame ls , d e f ined a s t ha t p o in t i nt he i r l i f e c yc le a t wh ich r ep roduc t ion f i r s t b ecomes p oss ib le , i sg ene ra l ly a ch ieved a tt h ree t of ou ry ea r s o fa ge . F ema les , a nd a ppa ren t ly ma les , w i l l g ene ra l ly r ema in f e r t i le t h rough a tl e as tt he i rt wen t ie th y ea r . Wh i le i n d iv idua l c ame ls may l i ve t ob e 4 0 t o5 0 y ea rs o fa ge , t he i r n o rma l b reed ing l i f e g ene ra l ly e nds b e tween 2 0 a nd 3 0 y ea rs o f a ge . ( B r i t ish N ava l I n te l l igence 1 946 :507 ; G au th ie r -P i l te rs a nd D agg 1 981:93 ; L eese 1 927 :96 , 1 37 ; L eona rd 1 894 :119 -120 ; N ovoa 1 970 :5 ; R ao e t a l . 1 970 :72 ; W i l l iamson a nd P ayne 1 959 :322 ; Y as in a nd Wah id 1 957 :294 ) . H oweve r , c ame l p as tora l is ts f r equen t ly i n te r fe re w i th t he n a tura l b reed ing t endenc ies o f t he i r h e rd a n ima ls . F ac to rs a ppa ren t ly s i gn i f ican t i nt he i r i n ter fe rence a re d i scussed i nt he f o l low ing s ec t ions .

H uman I n te r fe rence i nt he C ame l 's A ge a tF i r s t P ar tur i t ion A s n o ted b y L eese ( 1927 :96 ) , t he b reed ing o f af ema le c ame l i nh e r t h i rd y ea r may h ave d e le te r ious e f fec ts o n b o th t he g row th o f t he y oung d am a nd t he v i ab i l i ty o f h e r o f fsp r ing u n less t he mos tf a vo rab le o fc ond i t ions a re ma in ta ined . S im i la r ly , ma le c ame ls d o n o t a ppa ren t ly a ch ieve f u l l r ep roduc t ive s t am ina u n t i l s i x o rs even y ea rs o fa ge ( Novoa 1 970 :5 ; S a to 1 980 :25 ; W i l l iamson a nd P ayne 1 959 :322 ) . H uman i n ter fe rence i nt h e a ge a t wh ich c ame ls a re f i r s t b red a ppea rs t or e la te t os uch f a c to rs . A l low ing c ame ls t o b reed a tt h ree t of ou r y ea rs o fa ge w ou ld s eem a s t ra tegy b es t s u i ted f o rt he c ond i t ions n o rma l ly o b ta ined u nde r i n tens ive h usband ry , a s i sf r equen t ly t he c ase i nt he S ov ie t U n ion ( Te ren t jev , i n D ah l a nd H jor t 1 976 :79 a nd N ovoa 1 970 :5 ) . E ven s o , i n I n d ia ( Leese 1 927 :96 ; R ao e ta l .1970 :72 ) , P ak is tan ( Yas in a nd Wah id 1 957 :294 ) , S oma l ia ( Ma res , i n 1 64

D ah l a nd H jo r t 1 976 :79 ) , K enya ( Sa to 1 980 :25 ) , a nd t he t r op ics i ng ene ra l ( W i l l iamson a nd P ayne 1 959 :322 ) , f ema le c ame ls a re f i r s t b red i n t he i r f our th o r f i th y ea r . A cco rd ing t o N ico la isen ( 1963 :56 ) , f ou r i sa l so t he a ge a t wh ich t he T ua reg b edou in o f t he c en t ra l a nd s ou the rn S aha ra n o rma l ly b reed t he i rc ame ls . H oweve r , S pence r ( 1973 :11) s t a ted t ha t f ema le c ame ls w e re f i r s t b red i nK enya i nt he i r s i x th y ea r , wh i le B ehnke ( 1 980 :3 1) n o ted t ha t f ema les i nL i bya b o re t he i r f i r s t y oung a tf i ve t o s i x y ea rs o f a ge . A cco rd ing t o Mus i l ( 1928 :333 ) , t he Rwa la b edou in o f N o r th A rab ia f i r s t b reed f ema les i nt he i r s i x th y ea r . Among t he M u ta i r b edou in o f K uwa i t , D ickson s t a ted t ha t f ema les we re f i r s t b red i nt h e i re i gh th y ea r ( 195 1 :4 13 ) . Swee t h as s t a ted t ha tf ema les a re g ene ra l ly b red i n N or th A rab ia i nt he i r f i f th ( 1970 :276 ) o r s i x th y ea rs ( 1969 :159 ) , wh i le K öh le r ( 1984 :202 ) n o tes t ha t f ou r t os i x y ea rs o fa ge i sn o rma l i nJ o rdan . I th as a l so b een s ugges ted t ha t f ema le c ame ls i nt he M idd le E as t a re n o t g ene ra l ly e xpec ted t o b eg in b ea r ing u n t i l t he ir s i x th y ea r ( Bu l l ie t 1 975 :265 ) . T hese v a r ia t ions i nt he p re fe r red a ge a tw h ich f ema les a re f i r s t a lowed t ob reed a ppea rs t o r e f lec t a n a d jus tmen t o f b reed ing a ge a cco rd ing t o t he p hys io log ica l d eve lopmen t o f y oung d ams a nd t he i r c onsequen t a b i l i ty t o e f fec t ive ly f eed o n a va i lab le p as turage . When f eed ing , c ame ls f i r s t g rasp t he v ege ta t ion b e tween t he i r p rehens i le l i ps a nd t hen e i the r c l ip o r s t r ip t he v ege ta t ive ma t te r b e tween t he i r l owe r i n c iso rs a nd t he i r u ppe r g um ( Gau th ie r -P i l te rs a nd D agg 1 98 1 :34 ) . A s ap r ima ry s t ruc tura l f ea tu re o ft he c ame l 's f eed ing mechan ism , t he c ond i t ion o fa f ema le 's i n c iso rs w ou ld d i rec t ly a f fec t i t s f eed ing e f f ic iency . A cco rd ing t o S ha lash ( 1965 : 107 ) , ma lnu t r i t ion i st he p r ima ry c ause o f s ub fuc t ion ing o va r ies , o ne o f t he t h ree mos t c ommon ly o bse rved s exua l d i sorde rs i n f ema le c ame ls . F ur ther , L eese ( 1927 :99 ) n o ted t ha t a n i nadequa te f ood s upp ly was o ne o f t he t wo p r inc ip le c auses o f h i gh a bor t ion r a tes among c ame ls . Wh i le c ame ls d eve lop s i x t empo ra ry i n c iso rs i nt he i r f i r s t y ea r ( age c l ass 0 1) , t hese a re n o rma l ly w o rn t os t umps b y t he a ge c l ass 4 -5 ( C ross 1 9 17 :4 ; L eese 1 927 :134 ; W i l l iamson a nd P ayne 1 959 :324 ) . On ly i nt he i r s i x th y ea r ( age c l ass 5 -6 ) d o t hey d eve lop t he i r f i r s t t wo p e rmanen t i n c iso rs , a nd t h is n umbe r i n c reases t of ou r i na ge c l ass 6 -7 a nd f i na l ly t os i x i n c iso rs i n a ge c l ass 7 -8 ( C ross 1 9 17 :5 ; Gau th ier -P i l te rs a nd D agg 1 981:34 ; L eese 1 927 :134 135 ; Mus i l 1 928 :338 ; W i l l iamson a nd P ayne 1 959 :324 ) . S ince p erenn ia l s h rubs a re d om inan t i nt he c ame l 's d i e t i nN or th A f r ica , P a les t ine , S ina i , a nd N or th A rab ia ( Behnke 1 980 :29 ; B odenhe ime r 1 935 :126 ; B r i t ish N ava l I n te l l igence 1 946 :5 10 ; Gau th ie r -P i l te rs a nd D agg 1 98 1:39 -43 ; J enn ings -B ram ley 1 906 :256 -257 ; J ohnson 1 969 :48 ; Mus i l 1 928 :337 -338 ) , i ti ss uspec ted t ha t p as to ra l is ts i nt hese r i e g ions g ene ra l ly d e lay t he a ge a t wh ich f ema les a re f i r s t b red a s af unc t ion o ft he i r d en ta l d eve lopmen t . U nde rt hese c ond i t ions , ad e layed b reed ing w ou ld e nhance a f ema le 's a b i l i ty t o i n ges t s uf ic ien t q uan t i t ies a nd v a r ie t ies o f f ood t o e nsu re f e r t i l i ty , t he d eve lopmen t o f v i ab le o f fsp r ing , a nd a c ons is ten t f l ow o fm i lk d ur ing l a c ta t ion . I nt he p resen t mode l o fc ame l h e rd g row th , t h is p re fe rence f o r a l a te rb r eed ing o ff ema les w i l l b e a ssumed .

Human I n ter ference i nt he C ame l 's F e r t i le L i fespan B o th L eese ( 1927 :96 ) a nd W i l l iamson a nd P ayne ( 1959 :322 ) n o ted t ha t f ema le c ame ls a re n o rma l ly b red o n ly u n t i l , o r s hor t ly a f te r , t he i r t wen t ie th y ea r . T h is i sc e r ta in ly t he c ase i n S oma l ia ( Ma res , i nD ah l a nd H jo r t 1 976 :80 ) a nd K enya ( Sa to 1 980 :25 ; T o r ry 1 974 :67 ) , w he re c ame ls o l der t han 2 0 y ea rs a re n o rma l ly s l augh tered f o r mea t . T ua reg i n fo rman ts c ons ide red t he a ve rage n a tura l e nd o f af ema le 's b reed ing l i f e a s o ccur r ing a t 2 5 y ea rs o f a ge , b u t r ecommended t ha t a ny c ows o lder t han 2 0 s hou ld b e s o ld o r s l augh te red ( N ico la isen 1 963 :63 ) . Among t he Rwa la ( Mus i l , 1 928 :334 , 5 47 ) a nd Mu ta i r b edou in ( D ickson 1 951 :413 ) , ma le a nd f ema le c ame ls b eyond 2 0 y ea rs o fa ge a re c ons ide red o l d , a nd f r om t h is y ea r u n t i l t he i rd ea th a re r e fe r red t ob yt e rms d i fe ren tf r om t hose a pp l ied t o" m idd le -aged " c ame ls . Whe ther d ue t od ec l in ing f e r t i l i ty , m i lk p roduc t ion , o r p robab i l i ty o f s urv ivo rsh ip , f ema les o lde r t han 2 0 y ea rs o f a ge a re n o rma l ly c u l led f r om p as to ra l h e rds . T he e f fec ts o f h uman i n ter fe rence i nt he a ge o ff i r s t p a r tu r i t ion a nd t he f e r t i le l i f espan o fc ame ls w i l l b e a ccoun ted f o r i nt he p resen t mode ls o fh e rd g row th .

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C ame l Y ean ing R a tes Y ean ing r a tes a re t he mos t c r i t ica l r ep roduc t ive v a r iab le i n h e rd g row th mode ls . T he y ean ing r a te i st he n umbe r o f b i r ths e xpec ted a nnua l ly p e r r ep roduc t ive ly ma tu re f ema le i na h e rd , a nd mus t t ake i n to a ccoun t t he f r equency o f b o th mu l t ip le b i r ths a nd r ep roduc t ive f a i lures . S ince y ean ing r a tes d i rec t ly a f fec t a nnua l h e rd g row th a nd t hus p o ten t ia l h e rd p roduc t iv i ty t h rough t he c u l l ing o fe xcess a n ima ls , i n accu racy i ny ean ing r a tes w i l l s i gn i f ican t ly a f fec t c a lcu la t ion o f p as to ra l b ene f i ts . T he mos t s i gn i f ican t d ev ia t ion f r om p r io r mode ls i nt he r ep roduc t ive v a lues emp loyed i nt he p resen t a na lys is i n vo lves y ean ing r a tes . W i thou t ac ons ide ra t ion o f mu l t ip le b i r ths a nd r ep roduc t ive f a i lu res , t he b i enn ia l c a lv ing o f c ame ls w ou ld i mp ly a0 . 5 y ean ing r a te . T h is e s t ima ted v a lue w as emp loyed b y D ah l a nd H jo r t i n t he i r mode ls o f c ame l h e rd g row th ( 1976 :81-82 ) . H oweve r , wh i le t he i n c idence o f t w in a nd t r ip le p regnanc ies i nf ema le c ame ls i st oo l ow t or a ise t h is e s t ima ted 0 . 5 y ean ing r a te ( Bha rgava e t a l . 1 963 ; L eese 1 927 :95 ; S ha lash 1 965 :106 ) , d a ta p resen ted b e low s ugges t t ha t r ep roduc t ive f a i lures d ue t oi n fe r t i l i ty a nd a bor t ions wou ld s i gn i f ican t ly l owe r i t . B y c on t ras t , S a to ( 1980 :28 -3 1) emp loyed a l owe ry ean ing r a te o fa pp rox ima te ly 0 .2 i nh i s mode l o fc ame l h e rd g row th among t he R end i l le o f n o r the rn K enya . H oweve r , w h i le t h is l owe r v a lue may h ave b een a pp rop r ia te f o r R end i l le h e rds a tt he t ime o f S a to 's f i e ld r e sea rch , i ti s s eem ing ly t oo l ow t oc ha rac te r ize g ene ra l c ame l h e rd g row th , a nd i sp robab ly m o re a pp rop r ia te f o r h e rds ma in ta ined u nde r r ep roduc t ive ly d e le te r ious c ond i t ions . I n fe r t i l i ty . S ha lash ( 1965 :107 ) n o ted t ha t i na dd i t ion t of a u l ty managemen t , d i sease f ac to rs w e re t he p r imary c ause o f i n fer t i l i ty i nc ame ls . T h is i n c ludes b o th s pec i f ic d i seases d i rec t ly a f fec t ing t he r ep roduc t ive o rgans , a s we l l a s n onspec i f ic d i seases wh ich r educe t he o ve ra l l r ep roduc t ive e f f ic iency o ft he a n ima l . T he re a re a tl e as t 3 1 s pec i f ic s exua l d i so rde rs i nf ema le c ame ls wh ich a f fec t t he i r f e r t i l i ty , i n c lud ing i n fan t i le u te rus , p yome t ra a s ap robab le r esu l t o f i n fec t ions , a nd s ub func t ion ing o va r ies a s ap robab le r e su l t o f ma lnu t r i t ion ( Sha lash 1 965 :105 1 07 ) . Wh i le t he o ve ra l l f r equency o f s t e r i l i ty i nf ema le c ame ls i ss a id t o b e v er y l ow ( Leese 1 927 :95 , 3 22 ) , i n fer t i l i ty i ng ene ra l w ou ld t end t o l owe r t he 0 .5 y ean ing r a te emp loyed i nt he mode ls o fD ah l a nd H jo r t . A bo r t ions . P r ima ry d a ta o n a bor t ions i nc ame ls i sp rov ided b y t he r esea rch o fB ha rgava e t a l . ( 1963 ) a t t he B ikane r b reed ing f a rm i n I n d ia . I n t h is s t udy , 2 27 o bse rved p regnanc ies r e su l ted i n2 05 l i ve b i r ths a nd 2 2 a bor t ions . T h is w ou ld s ugges t a n a bor t ion r a te o f9 .7 % o ft o ta l p regnanc ies . I na l c ases , t r ypanosom ias is w as s uspec ted a s t he p r inc ip le c ause o f a bo r t ion ( Bha rgava e t a l . 1 963 :188 ) . B o th L eese ( 1927 :99 , 3 22 -323 ) a nd C ross ( 1917 :91) h ad p rev ious ly o bserved t ha t a bo r t ions among c ame ls w e re g ene ra l ly t he r esu l t o f e i the r t r ypanosom ias is o r as ca rc i ty o f f ood . S ince l ower y ean ing r a tes among c ame ls s ub jec t t o t r ypanosom iases wou ld s i gn i f ican t ly a f fec t t o ta l h e rd p roduc t iv i ty b y r educ ing h e rd g row th , a b r ie f d i scuss ion o ft he n a tu re a nd d i s t r ibu t ion o ft hese d i seases i sw a r ran ted . T rypanosom ias is . T rypanosom iases a re d i seases c aused b y f l age l la ted p r o tozoa s p read b y b l oodsuck ing i n sec ts , p r ima r i ly t aban id f l i es wh ich s easona l ly b reed i nr eg ions n ea r o pen wa te r s ources a nd wooded a nd l ow ma rshes ( Leese 1 927 :217 -254 ; R u t te r 1 967 :611-6 13 ; S a lyaev 1 956 :483 -485 ) . Wh i le r eg iona l ly v a ry ing i nf r equency a cco rd ing t o t h e a va i lab i l i ty o f t he i r v ec to r 's b reed ing g rounds , t hese d i seases a f fec t c ame ls f r om K azakhs tan i n S ov ie t C en t ra l A s ia ( Sa lyaev 1 956 :485 ) t h rough Mo rocco a nd Wes t A f r ica ( Leese 1 927 :222 -254 ; R u t te r 1 967 :6 12 ) . I th as b een s ugges ted t ha t i nt he i r n a t ive h ab i ta ts , a pp rox ima te ly 3 0% o ft he d i seases f ound among O ld Wo r ld c ame ls a re c aused b y t r ypanosomes ( K lös a nd L ang 1 982 :225 ; R u t te r 1 967 :6 11) . R eg ions a l ong ma jo r r i ve rs ( e .g . , t he T ig r is , E uph ra tes , J o rdan , a nd N i le ) , c e r ta in d eser t o ases ( e .g . , A z raq i n e as te rn J o rdan ) , a nd v a r ious d ese r t wad is w ere n o ted b y L eese a s s umme rt r ypanosome z ones ( 1927 :250 ) . J enn ings -B ram ley s pec i f ica l ly n o ted t ha t t he Wad i e l A in i nS ina i a nd t he F ayum d ep ress ion i nE gyp t w e re k nown t r ypanosome z ones ( 1905 :131) , a s was t he r eg ion a round A in e l - Waz i r i nt he Wad i e l -Hes i , s ou thwes t o f A qaba ( 1913 :83 ) . D ickson n o ted t ha t i nI r aq , t r ypanosom ias is w as s p read among c ame ls b y b i t ing f l i es wh ich o n ly f l our ished d u r ing t he s umme r a l ong t he b anks o f t he E uph ra tes R ive r a nd t he S ou th H ama r L ake ( 1951 :56 ) .

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A cco rd ing t o R u t te r ( 1967 :6 12 ) , t r ypanosom ias is i s" w idesp read " i n S aud i A rab ia , o f " mode ra te i n c idence " i nJ o rdan , a nd o f" l ow s po rad ic i n c idence " i nS y r ia . H oweve r , i na n F AO r epo r tt ot he g ove rnmen to fS aud iA rab ia ( 1963a : 14 ) ,t he a va i lab le d a ta o nt r ypanosom ias is was c ons ide red t oo u n re l iab le t oa ccess i t s i n c idence i nS aud i A rab ia . S ince t r ypanosom iases c an a l so b e s p read b y t aban id f l i es wh ich b reed i nf r esh d ung ( Leese 1 927 :249 ; R u t te r 1 967 :6 15 ) , h eavy r a ins i nt he d eser t c an p o ten t ia l ly l e ad t o s po rad ic o u tb reaks o ft hese d i seases ( Ru t te r 1 967 :615 ) . A cco rd ing t o D ickson ( 1951 :4 18 ) , t he Mu ta i r b edou in o f K uwa i t u nde rs tood t he r eg iona l a nd s easona l a spec ts o ft hese d i seases a nd t he i r c o r re la t ion w i th t he b reed ing a reas o fb i t ing f l i es . T hus , n one o f t he Mu ta i r wou ld w i l l ing ly move t he i r c ame ls i n to t he v i c in i ty o f t he E uph ra tes d ur ing t he s umme r . L eese h ad a l so n o ted t ha t i n I r aq , " t he n a t ives k now b y e xpe r ience t he wo rs t a reas , a nd a vo id t hem i nt he h o t wea the r " ( 1927 :250 ) . T h is w as a l so t he c ase among c ame l p as to ra l is ts i nA lge r ia ( Leese 1 927 :251 ) . P r imary t r ypanosome z ones c an t he re fo re b e s easona l ly a vo ided , a l though s po rad ic o u tb reaks o f t r ypanosome i n fec t ion c an p o ten t ia l ly o ccu r e ven i nr e g ions d evo id o fp e rmanen t wa te r s ou rces . T he a bor t ion r a te o bserved among p regnan t c ame ls a tt he B ikane r b reed ing f a rm wou ld t he re fore s eem a pp rop r ia te f o r c ame ls r a ised i n o the r l o ca t ions a s we l l . C umu la t ive ly , r ep roduc t ive f a i lu res a s ar e su l t o fs t e r i l i ty a nd a bo r t ions c an b e e xpec ted t os i gn i f ican t ly l owe r t he y ean ing r a te o fc ame ls . Observed Y ean ing R a tes . A s n o ted b y N ovoa ( 1970 :17 ) a nd Gau th ie r -P i l te rs a nd D agg ( 1981:93 ) , q uan t i ta t ive d a ta o n o bserved y ean ing r a tes among c ame l h e rds i s e x t reme ly s ca rce . H oweve r , t he a va i lab le d a ta d o s ugges t r a tes s ubs tan t ia l ly l owe r t han t he 0 .5 y ean ing r a te emp loyed i nt he mode ls o f D ah l a nd H jor t ( 1976 ) . I n aS ov ie t s t udy o fc ame l r ep roduc t ion b y Y uz l ikaev a nd A khmed iev ( 1965 :62 ) , y ean ing r a tes b y d i s t r ic t d u r ing 1 964 we re r epo r ted f o r t he p rov ince o f K azakhs tan , S ov ie t C en t ra l A s ia , a s o b ta ined u nde r " t rad i t iona l " me thods o f c ame l b reed ing . T he i r r e su l ts a re p resen ted i nT ab le 4 3 . A cco rd ing t o N ovoa ( 1970 :17 ) , t hese y ean ing r a tes a re s im i la rt ot hose f ound among c ame l h e rds i nv a r ious o the r c oun t r ies . I n deed , t he 0 .39 mean d e r ived f r om t hese d a ta i sv ery c l ose t o t he 0 .4 y ean ing r a te e s tab l ished f o r G ab ra c ame l h e rds i nn or the rn K enya i n1 969 ( Tony 1 973 :7 1 , 7 4 ) . I ti st he re fo re s u rp r is ing t of i nd t ha t amuch l owe ry ean ing r a te was s ugges ted b y t he d a ta g a the red b y S a to ( 1980 ) o n R end i l le c ame l h e rds i nn o r the rn K enya . I n 1 976 , t he 6 04 a du l t a nd a do lescen t f ema les i n2 0 s epa ra te R end i l le h e rds g ave b i r th t o o n ly 11 c a lves ( Sa to 1 980 :26 , 2 8 ) . S a to t he re fo re c onc luded t ha t a pp rox ima te ly o ne c a l f w ou ld b e b o rn a nnua l ly f o r e ve ry f i ve f ema les , s ugges t ing a 0 . 184 t o 0 .2 y ean ing r a te among R end i l le c ame ls . T hese l owe ry ean ing r a tes w e re u sed b y S a to i nh i s mode l o fc ame l h e rd g row th ( 1980 :30 ) . S ign i f ican t ly , S a to ( 1980 :20 , 2 7 ) n o ted t ha t i nR end i l le t e r r i to ry , o u tb reaks o ft aban id f l i es b ea r ing t r ypanosomes f l uc tua te y ea r ly i nb o th t empo ra l a nd s pa t ia l d imens ions , a nd a s ar e su l t , t he R end i l le d i d n o tw i thd raw t he i rc ame ls f r om f l y in fes ted r eg ions u n less t he i ra n ima ls a c tua l ly

T ab le 4 3 C ame lY ean ing R a tes b y D i s t r ic t i nK azakhs tan , S ov ie t C en t ra l A s ia , 1 964

D ISTR ICT

Y EAN ING RATE

K zy l -O rd insky G u ryevsky U ra lsky C h imken tsky

0 .36 0 .37 0 .38 0 .43

Mean

0 .385

a dap ted f r om Y uz l ikaev a nd A khmed iev ( 1965 :62 )

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c on t rac ted t he d i sease . F ur the r , S a to ( 1980 :10 , 2 2 ) a l so n o ted t ha t t he re h ad b een a lmos t n o p rec ip i ta t ion b e tween 1 968 a nd 1 975 i nt he l ow lands whe re t he R end i l le h e rd t he i rc ame ls . I ti s t he re fo re p oss ib le t ha t t he R end i l le c ame ls g i v ing b i r th i n1 976 h ad b een d eb i l i ta ted b y s eve ra l y ea rs o f r e la t ive ly p oo r p as turage . A s s uch , ay ean ing r a te o f a round 0 .2 s eems mo re a pp rop r ia te f o rc ame l h e rds ma in ta ined u nde r d e le te r ious b reed ing c ond i t ions . I nt he p resen t mode ls o f c ame l h e rd g row th , a0 .39 y ean ing r a te w i l l b e a ssumed f o r a l b reed ing f ema les . T h is r a te i ss ubs tan t ia l ly l e ss t han t he 0 . 5 y ean ing r a te emp loyed i nt he mode ls o f D ah l a nd H jo r t ( 1976 :8 1-82 ) , y e t h i ghe rt han t he 0 . 184 t o0 . 2 y ean ing r a te emp loyed b y S a to ( 1980 :28 -3 1) i n mode l l ing R end i l le h e rd g row th i n1 976 .

S ex R a t io T he s ex r a t io r e f lec ts t he r e la t ive p e rcen tage o f ma les a nd f ema les e xpec ted p e r 1 00 l i ve b i r ths . H av ing e s tab l ished a n a pp rop r ia te y ean ing r a te , t he s ex r a t io c an b e u sed t o c a lcu la te t he a nnua l i n c rease e xpec ted among t he ma le a nd f ema le c omponen ts o f ah e rd . I n as t udy o f5 16 p regnan t c ame l u te r i e xam ined f o rt h e d e tec t ion o ft he s ex r a t io , S ha lash ( 1965 :106 ) f ound t ha t 4 7 .48% we re ma le a nd 5 2 .52% w e re f ema le . H oweve r , t h is w as a n a u topsy s t udy , a nd t he r esu l ts d o n o t t he re fo re r ep resen t as ex r a t io b ased u pon o bse rved b i r ths . O f t he 2 05 l i ve b i r ths o bse rved a tt he B i kane r b reed ing f a rm i nI n d ia , 1 0 1 we re ma le a nd 1 04 w e re f ema le , t hus s ugges t ing a s ex r a t io o f4 9 .3% ma les a nd 5 0 .7% f ema les ( Bha rgava , S ha rma , a nd S ingh 1 963 ) . I nt he p resen t s t udy , as ex r a t io o f4 9 % ma les a nd 5 1% f ema les w i l l b e emp loyed . I nt he p r io r mode ls o fD ah l a nd H jo r t ( 1976 ) a nd S a to ( 1980 ) , a5 0 :50 s ex r a t io w as a ssumed .

Mo r ta l i ty Among D romeda ry C ame ls G ene ra lC ons idera t ions E thnograph ic a nd managemen t l i t e ra tu re p r ima r i ly c on ta in q ua l i ta t ive r a the r t han q uan t i ta t ive mo r ta l i ty d a ta , a l though i ti sw ide ly ma in ta ined t h a t y oung c ame ls a re s ub jec tt ot he h i ghes t mor ta l i ty r a tes ( Leese 1 927 :97 ; L eona rd 1 894 :123 ; W i l l iamson a nd P ayne 1 959 :323 ) . S a to ( 1980 ) u t i l ized o bse rved mor ta l i ty r a tes among R end i l le h e rds i nh i s mode l o fc ame l h erd g row th . H is d a ta w i l l b e e xam ined b e low . H oweve r , l a ck ing q uan t i ta t ive mo r ta l i ty d a ta , D ah l a nd H jor t ( 1976 :8 1-83 ) s imp ly g ene ra ted t wo a l te rna t ive mode ls o f c ame l h e rd g row th i n vo lv ing d i s t inc t s u rv ivo rsh ip c u rves , b o th o fw h ich i mp ly h i gh mo r ta l i ty r a tes among y oung c ame ls . I nt he i rf i r s t mode l , t hey a ssumed a3 0 % mor ta l i ty r a te among f ema le c ame ls d ur ing t he i r f i r s ty ea r , a f te rw h ich a l s u rv ivo rs l i ved t i ld ea th a ta ge 2 1 . T h is w ou ld a l so i mp ly a3 0% d ea th ra te among a l n ew ly -bo rn c ame ls t h rough t h ree y ea rs o f a ge i fma le a nd f ema le d ea th ra tes w e re e qua l . I nt he i r s econd mode l , t h is 3 0% mor ta l i ty r a te w as r e ta ined , w h i le f u r ther i n tu i t ive r a tes w e re a dded t o o b ta in " mo re r ea l is t ic mo r ta l i ty f i gures . " T hese a dd i t iona l f i gures i mp ly a 5 3% t o ta l d ea th r a te f o ra l n ew ly b o rn c ame ls t h rough t he i rt h i rd y ea r . T he mor ta l i ty r a tes f o rf ema le c ame ls w h ich we re a ssumed i nt he i r s econd mode l a re p resen ted i nT ab le 4 4 . I n l i gh t o f r e co rded c ame l mor ta l i ty d a ta u nava i lab le t o D ah l a nd H jor t , i tw ou ld s eem t ha t t he ir i n t ia l 3 0% mo r ta l i ty a ssump t ion w as mo re a ccu ra te w h i le t he i rs econd s e to fv a lues w e re t oo h i gh . I nt he f i r s t p lace , L eona rd ( 1894 :123 ) n o ted t ha t t he " g rea t mo r ta l i ty among n ew ly -bo rn c ame ls u pt of ou ry ea rs o l d " was " gene ra l ly p u t d own a t5 0 p e rc en t , " ar a te wh ich h e o bv ious ly c ons ide red e xcess ive g i ven t he c on tex t o f h i s c ommen t ( t he i n t roduc t ion o f O ld Wo r ld c ame ls i n to t he N ew Wor ld ) . D ah l a nd H jo r t 's s econd mode l p resumes as l igh t ly h i ghe rd ea th ra te u nde r t he a ssump t ion t ha t n o s i gn i f ican t d i f fe rences b e tween ma le a nd f ema le d ea th ra tes e x is t . H oweve r , q uan t i ta t ive d a ta s ugges t as i gn i f ican t ly h i ghe rd ea th ra te among y oung ma le c ame ls , amoun t ing t o1 . 86 ma le d ea ths f o re very f ema le d ea th ( Bha rgava e ta l . 1 963 :187 ) .

16 8

T ab le 4 4 T he F ema le Mo r ta l i ty R a tes i nD ah l a nd H jo r t 's S econd Mode lo fC ame l H e rd G row th ( 1976 :83 )

A ge C lass

Mo r ta l i ty a s a° A o ft he a ge c l ass

S u rv ivo rsh ip a s a% o ft he a ge c l ass

0 1 1 -2 2 -3 3 -4 4 -14 1 4 -15 1 5 -16 1 6 -17 1 7 -18 1 8 -19 1 9 -20 2 0 -21

3 0 2 2 1 4 6 2 7 1 1 1 6 2 1 2 5 3 0 4 2

7 0 7 8 8 6 9 4 9 8 9 3 8 9 8 4 7 9 7 5 7 0 5 8

A tt h is r a t io , D ah l a nd H jor t 's s econd s e t o f mo r ta l i ty v a lues w ou ld i mp ly t ha t a round 8 3% o f a l n ew ly -bo rn c ame ls d i e b e fo re t hey r each f our y ea rs o f a ge , av a lue c l ea r ly t oo h i gh . Mo re r easonab le i st he i r f i r s t mode l , wh ich i mp l ies t ha t a round 4 3% o f a l n ew ly -bo rn c ame ls d ie b e fo re t hey r each f ou ry ea rs o fa ge , a ga in u s ing a 1 . 86 : 1 ma le t of ema le d ea th r a t io . Wh i le D ah l a nd H jor t a ssumed i nt he i r s econd mode l t ha t f ema le mor ta l i ty r a tes wou ld b eg in t o i n c rease i nt he i r f our teen th t o f i f teen th y ea r , t he v a lues t hey u sed wou ld i mp ly t ha t a round 8 7% o f a l f ema les s urv iv ing t he i r t h i r teen th y ea r wou ld b e d ead b y t he e nd o f t he i r t wen ty f i rs t y ea r . A s p rev ious ly n o ted , t he n o rma l l i f e e xpec tancy o fc ame ls r anges f r om 2 0 t o 3 0 y ea rs o fa ge . T hus , t he h i gh mor ta l i ty v a lues u sed b y D ah l a nd H jor tf o rf ema le c ame ls a f te r a ge c l asses 4 14 w ou ld s eem t ob e d ep ressed b y a tl e as tf ou ro r mo re y ea rs . Wh i le o n ly t wo q uan t i ta t ive s t ud ies o n n a tura l mor ta l i ty among d romeda ry c ame l h e rds h ave b een f ound , t hese d o a low f o r t he e s tab l ishmen t o f mo re r ea l is t ic c ame l mor ta l i ty r a tes t han t hose u sed i nt he h e rd g row th mode ls o fD ah l a nd H jor t ( 1976 ) .

Mor ta l i ty R a tes U nde rI n tens ive H usband ry a t B ikane r , I nd ia F rom at h ree y ea r s t udy o f n a tu ra l mo r ta l i ty among h e rds a tt he B ikane r C ame l B reed ing F a rm i nI n d ia , B ha rgava e ta l . r eached t he f o l low ing c onc lus ions ( 1 963 :187 ) : T he mo r ta l i ty r a te f r om b i r th t ot h ree y ea rs o fa ge i nc ame l -ca lves was c a lcu la ted t o b e 1 9 .5 1 p e r c en t , 4 0 d ea ths o ccu r r ing o u t o f2 05 n o rma l b i r ths . O u t o ft o ta l 4 0 mor ta l i t ies , 2 6w e re ma le c a lves a nd 1 4w e re f ema le . T he r eason a t t r ibu ted t ot h is h eavy mo r ta l i ty i n ma le c a lves was d ue t op re fe ren t ia l t r ea tmen t g i ven t o f ema le c a lves i nt he h e rd . T he f ema le c a lves r ema in i nt he h e rd a l t he i r l i f e a nd s uck le t he m i lk o ft he d am f o r ag rea te rp e r iod . T h is w ou ld s ugges t t ha t 6 5% o fa l d ea ths t ot h ree y ea rs o fa ge o ccu r among ma le c ame ls , a nd i n d ica tes ad ea th r a t io o f1 . 86 ma le d ea ths t oe ver yf ema le d ea th . B ha rgava e t a l . f u r the r s u mma r ized n a tura l l o sses among t he 2 05 n o rma l b i r ths i nt he B i kane r h e rd a sf o l lows ( 1963 : 187 ) : O f t he 4 0 c a lves t ha t d i ed , 3 3 d i ed w i th in f i r s t s i x mon ths o fb i r th , o ne b e tween 6t o 1 2 mon ths , o ne b e tween 1 1 / 2 t o 2y ea rs , a nd f i ve b e tween t wo a nd t h ree 1 69

y ea rs o fa ge . T h is s hows t ha t n ea r ly 8 2 .5 p e rc en t o fm o r ta l i ty t ook p l ace b e fo re t he c a lves r eached s i x mon ths o fa ge . T hese d a ta h ave b een c onver ted i n to a n a ge a nd s ex s pec i f ic s umma ry i nT ab le 4 5 o n t he a ssump t ion t ha t t he o bserved ma le t o f ema le d ea th r a t io r ema ined c ons tan t t h rough a l a ge c l asses . Ap e rusa l o fp os tmor tem r epor ts " gave ag ene ra l i mp ress ion t ha t mos t o ft he d ea ths w ere d ue t o r esp i ra tory a nd d i ges t ive t r oub les " ( Bha rgava e ta l . 1 963 : 188 ) . T hey f u r the r s ugges ted t ha t i ng ene ra l , " young c ame ls a re mo re s uscep t ib le t o i n fec t ious d i seases a nd c hanges i n w ea the r c ond i t ions t han o l de r o nes " ( 1963 : 187 ) . E thnog raph ic d a ta wou ld s ugges t t ha t t hese s ame f ac to rs a re p r ima r i ly r e spons ib le f o r h i gh mo r ta l i ty r a tes among y oung c ame ls i np as to ra l h e rds u nde r c ond i t ions o f e x tens ive h usbandry ( e .g . , D ickson 1 95 1:4 18 ; J enn ings -B ram ley 1 906 :205 ; L ancas te r 1 98 1:10 , 1 0 1; Mus i l 1 928 :368 ; T o r ry 1 973 :72 , 1 27 ) . H oweve r , s i nce t he c ame ls a t t he B ikane r B reed ing F a rm we re ma in ta ined u nder c ond i t ions o f i n tens ive h usband ry , i ts eems r easonab le t os uspec t t ha t t he mo r ta l i ty r a tes i nT ab le 4 5 wou ld b e l owe r t han t hose w h ich n o rma l ly o ccu r u nde rc ond i t ions o fe x tens ive h usband ry .

Mo r ta l i ty R a tes U nder E x tens ive H usband ry i n N or thern K enya S a to ( 1980 :29 ) r e co rded n a tura l l o sses o ve r ao ne y ea r p e r iod among 2 0 c ame l h e rds o f t he R end i l le i nN or the rn K enya . H is d a ta a re s umma r ized i nT ab le 4 6 a cco rd ing t ot he a ge s ex c l asses w h ich h e emp loyed . When c ompa red w i th t he B ikane r mor ta l i ty d a ta i nT ab le 4 5 , t he o bse rved mor ta l i ty f o r " i n fan ts " among R end i l le h e rds f a l ls j u s t u nde rt he mor ta l i ty r a tes i na ge c l ass 0 1 a s r eco rded a tB ikane r . I f" j uven i le " i sd e f ined a s c ame ls i na ge c l asses 1 2 a nd 2 -3 , t he B ikane r d a ta wou ld s ugges t a3 .5 % mor ta l i ty r a te f o r j u ven i les , c ompa red t o t he 1 . 1% mo r ta l i ty r a te r eco rded f o r R end i l le h e rds . O ve ra l l , t he R end i l le d a ta w ou ld s eem ing ly s ugges t a l owe r mor ta l i ty r a te f o r y oung c ame ls u nde r e x tens ive h usbandry i nn or the rn K enya t han t ha t o b ta ined u nde r i n tens ive h usband ry a tB ikane r , I n d ia . T hese r e su l ts a re q u i te s urp r is ing s i nce t he Gab ra , l i v ing j u s t n o r th o ft he R end i l le , ma in ta in t ha t a pp rox ima te ly 4 0t o5 0% o ft he i ry oung c ame ls d i e b e fo re r each ing ma tu r i ty ( To r ry 1 973 :7 1) . I nt he a bsence o ff u r the r d a ta , i tw i l l b e a ssumed t ha t t he p rox im i ty o f t he B ikaner a nd R end i l le mor ta l i ty r a tes s imp ly s ugges ts t ha t w i th f avo rab le c ond i t ions , c ame l mor ta l i ty r a tes u nde r e x tens ive h usband ry wou ld b e c l ose t o t he r a tes e xpec ted u nde r n o rma l c ond i t ions o f i n tens ive h usband ry .

N a tu ra lC ame l Mo r ta l i ty R a tes A ssumed i nt he P resen t S tudy Wh i le t he B ikane r ma le t of ema le d ea th r a t io s eems a pp rop r ia te f o ru se i ne s tab l ish ing t he n a tu ra l mor ta l i ty r a tes t o b e u sed i n mode l l ing c ame l h e rd g row th , t he B ikane r mor ta l i ty r a tes s eem t oo l ow f o rt he c ond i t ions n o rma l ly e xpec ted u nde r e x tens ive h usband ry . T h is s hou ld b e T ab le 4 5 A ge -Spec i f ic N a tu ra l Mo r ta l i ty a nd R esu l t ing S u rv ivo rsh ip f o r Ma le a nd F ema le C ame l C a lves a t B i kane r , I n d ia

A ge C lass 0 -1 1 -2 2 -3

N umbe r i n A ge C lass Ma le F ema le 1 01 7 9 7 8

1 04 9 2 9 2

n D ea ths i A ge C lass Ma le F ema le

,

2 2 1 3

1 2 0 2

c a lcu la ted f r om d a ta p resen ted b y B ha rgava e ta l . ( 1963 ) 1 70

M o r ta l i ty a s a% l ass o fa ge c Ma le F ema le 2 2 1 4

1 2 0 2

S ur v ivo rsh ip a s a% o fa ge c l ass Ma le F ema le 7 8 9 9 9 6

8 8 1 00 9 8

T ab le 4 6 O bserved M o r ta l i ty R a tes Among R end i l le C ame l H e rds i n1 976

A ge C lass I n fan t J uven i le A do lescen t a nd A du l t

N umbe ri n A ge C lass Ma le F ema le

n D ea ths i A ge C lass Ma le F ema le 6 ( bo th ) 1

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9

1 74

6

1

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Mo r ta l i ty a s a% o ft he A ge C lass F ema le Ma le 9 .1 2 0 .9 . 1 ( bo th ) 1 3 .5

1 .8

S u rv ivo rsh ip a s a% o ft he A ge C lass Ma le F ema le 7 9 .1 9 0 .9 ( bo th ) 9 8 .9 9 6 .5

9 8 .2

a dap ted f r om S a to ( 1980 )

e spec ia l ly t r ue f o r t he p reh is to r ic p as t . S ince t he B ikane r d a ta s ugges t t ha t a round 2 0% o f a l n ew ly -bo rn c ame ls w i l l d i e b e fo re t hey a ch ieve t h ree y ea rs o fa ge u nde rc ond i t ions o fi n tens ive h usbandry , a pp rop r ia te mor ta l i ty r a tes u nde r a nc ien t c ond i t ions o fe x tens ive h usband ry s hou ld p resumab ly f a l l b e tween t he B i kane r r a te a nd t he 5 0% d ea th ra te e x t reme p rev ious ly n o ted b y L eona rd ( 1894 : 123 ) f o r n ew ly -bo rn c ame ls t h rough t he i rf our th y ea r . U s ing t he 1 . 86 :1 d ea th r a t io o b ta ined f r om t he B ikane r s t udy a nd R u t te r 's ( 1967 :6 11) 3% mo r ta l i ty r a te f o r a du l t c ame ls i nt r ypanosome r eg ions , t he n a tu ra l mor ta l i ty a nd c onsequen t s ur v ivo rsh ip r a tes p resen ted i nT ab le 4 7 h ave b een f o rmu la ted . T hese r a tes a ssume t ha t 4 7% o fa l n ew ly -bo rn ma le c ame ls a nd 2 8% o fa l n ew ly -bo rn f ema le c ame ls w ou ld d i e b e fo re t he i r f our th y ea r u nde r c ond i t ions o f e x tens ive h usbandry o n d eser t r anges . T he o ve ra l l d ea th ra te f o r t h is s ame p e r iod o ft ime t he re fo re amoun ts t o3 8% o f a l n ew ly -bo rn h e rd a n ima ls , ar a te a lmos t t w ice t ha t o bserved u nde r c ond i t ions o f i n tens ive h usband ry a t B i kane r , y e t l e ss t han b o th t he e x t reme 5 0% r a te p r ev ious ly n o ted a nd t he 4 0 t o5 0 p e rcen t d ea th ra te s ugges ted b y G ab ra i n fo rman ts ( To r ry 1 973 :7 1) . T he i n c reas ing mo r ta l i ty r a tes a ssumed f o ra ge c l asses 2 1-22 a nd 2 2 -23 w ere i n c luded u nde rt he e xpec ta t ion t ha t mos t c ame ls wou ld n a tura l ly d i e b e tween t he i rt wen t ie th a nd t h i r t ie th y ea r . T he v a lues a ss igned t hese a ge g roups w e re a ssumed t o b e t he i n ve rse o ft he f ema le mo r ta l i ty r a tes i na ge c l asses 1 -2 a nd 2 -3 . l ti sf u r the r a ssumed t ha t n a tu ra l mor ta l i ty r a tes f o r ma les s u rv iv ing t he i r t h i rd y ea r wou ld s ubsequen t ly p a ra l le l f ema le d ea th ra tes f o rt he s ame a ge c l asses .

T ab le 4 7 A ssumed N a tu ra l Mo r ta l i ty a nd R esu l t ing S urv ivo rsh ip R a tes Among D romeda ry C ame ls

A ge C lass 0 1 1 -2 2 -3 3 -21 2 1-22 2 2 -23

Mo r ta l i ty a s a` ) / 0 o ft he A ge C lass Ma le I F ema le 3 4 1 7 3 4 6

1 8 6 4 3 4 6

1 71

S u rv ivo rsh ip a s a% o ft he A ge C lass Ma le F ema le 6 6 8 9 9 3 9 7 9 6 9 4

8 2 9 4 9 6 9 7 9 6 9 4

A ccoun t ing f o rt he E fec ts o fH uman Managemen t S t ra teg ies I n t roduc t ion S ince ma le c ame ls a t ta in t he ma jo r i ty o ft he i ra du l ts i ze b y4 -6 y ea rs o fa ge ( W i l l iamson a nd P ayne 1 959 :320 -32 1) , t he r e ten t ion o f ma les s urv iv ing t he i rs i x th y ea r i ne xcess o ft h e n u mbe r r equ i red f o r s t ud a nd b aggage s e rv ice wou ld p r ima r i ly s er ve t o i n c rease g raz ing c ompe t i t ion a nd p o ten t ia l ly i n c rease t he f r equency o fc ommun icab le d i seases a nd p a ras i te l o ad w i th in t he h e rd . B y c on t ras t , a l r ep roduc t ive ly ma ture f ema les c ou ld p o ten t ia l ly c on t r ibu te d i rec t ly t ob o th h e rd g row th a nd m i lk p roduc t ion . C ame l p as to ra l is ts wou ld p resumab ly t ake t hese f a c to rs i n to c ons idera t ion w hen d ec id ing h ow b es t t o manage t he i r h e rds g i ven s pec i f ic p roduc t iv i ty g oa ls . F ur the r , i ti sa ppa ren tt ha t p as to ra l is ts i n te r fe re w i th b o th t he a ge a tw h ich f ema les a re f i r s t b red a nd t he i rf e r t i le l i f espan , a s d i scussed e a r l ie r . I ti sn ecessa ry t oa ccoun t f o r t he e f fec ts o fs uch managemen t s t ra teg ies w hen mode l l ing p as to ra l h e rd g row th .

I n te r fe rence w i th t he A ge o fF i rs tP a r tu r i t ion C ons is ten t w i th t he n a tura l b i enn ia l b reed ing p a t te rn o f c ame ls a nd a managemen t p re fe rence f o r t he l a te r b reed ing o fy oung f ema les a s d i scussed e a r l ie r , i tw i l l b e a ssumed i n t he p resen t mode l t ha t h a l f o ft he r ep roduc t ive ly ma tu re a do lescen t f ema les wou ld b e b red f o r t he f i r s tt ime i na ge c l ass 5 -6 , a nd t ha tt he o the rh a l f wou ld b e b red f o rt he f i r s tt ime a tt he e nd o f t he s ubsequen t y ea r . F i rs t p a r tu r i t ion f o r t he f o rme r g roup w ou ld t he re fo re o ccur a tt h e e nd o f a ge -c lass 6 -7 , a nd f o rt he l a t te rg roup a tt he e nd o fa ge c l ass 7 -8 . T h is s chedu l ing a ccommoda tes a d e fer red a ge o f f i r s t p a r tur i t ion wh i le a low ing f o r c on t inuous a nnua l h e rd g row th r a the rt han d i scon t inuous g row th r es t r ic ted t o a l te rna te y ea rs . I t a l so a lows t he e a r l ies t b red f ema les t he p o ten t ia l o fh av ing f o u rp e rmanen t i n c iso rs d u r ing t he i r g es ta t ion p er iod , a nd af u l l s e t o fs i x i n c iso rs a f ter p a r tu r i t ion a nd d u r ing l a c ta t ion . T he s econd g roup o ff ema les w ou ld p o ten t ia l ly h ave a l s i x i n c iso rs a tt he t i me t hey a re f i r s t b red .

I n te r fe rence i nt he F er t i le L i fespan o fF ema les C ons is ten t w i th t he p r io r d i scuss ion o f h uman i n te r fe rence i nt he f e r t i le l i f espan o ff ema le c ame ls , i tw i l l b e a ssumed t ha t h a l f o fa l e l de r ly f ema les a re l a s t b red i nt he i rt wen ty f i rs t y ea r , w h i le t he o the r h a t f a re l a s t b red a tt he e nd o f t he i r t wen ty -second y ea r . T he f o rme r g roup w ou ld t he re fo re d rop t he i r l a s t c a l f a tt he e nd o ft he i rt wen ty f i rs t y ea ra nd t hus b e a va i lab le f o r s l augh te r a tt he e nd o ft he i rt wen ty -second y ea r , wh i le t he o t he r h a l fw ou ld d rop t he i rl a s tc a l f a t t he e nd o f t he i r t wen ty -second y ea r a nd t hus b e a va i lab le f o r s l augh te r a t t he e nd o f t he i r t wen ty th i rd y ea r . Wh i le t he n a tu ra l f ema le s urv ivo rsh ip r a tes f o r a ge c l asses 2 1-22 a nd 2 2 -23 w e re s e t a t9 6% a nd 9 4% r espec t ive ly i nT ab le 4 7 , t he p reced ing a ssump t ions r educe t hese r a tes t o4 8% f o ra ge c l ass 2 1-22 , w h i le n of ema les w ou ld s u rv ive b eyond a ge c l ass 2 2 -23 . T hese f u r the r mode l l ing a ssump t ions a low e ach f ema le c ame l t he p o ten t ia l o fp roduc ing e i gh t c a lves i nh e r l i f e t ime , af i gu re w h ich a g rees w e l l w i th S pence r 's s t a temen t ( 1973 :11) t ha t R end i l le c ame ls i nK enya p roduce s even t ot en c a lves i n al i f e t ime , a nd t he e i gh t c a l f a ve rage n o ted f o rc ame ls i nS oma l ia ( Ma res , i nD ah l a nd H jo r t 1 976 :80 ) a nd K enya ( To r ry 1 974 :67 ) . T h is i sa l so c ons is ten t w i th t he s i x t o n i ne c a l f a ve rage n o ted f o r c ame ls i nt he S ina i b y J enn ings B ram ley ( 1906 :203 ) .

H uman C ropp ing S t ra teg ies T he a nnua l s l augh te ro fh e rd a n ima ls t h rough h uman c r opp ing s t ra teg ies c ou ld p o ten t ia l ly a fec t h erd g row th , d epend ing u pon t he e x ten t o f s l augh te r p rac t iced . I fa h e rd managemen t s t ra tegy i n vo lved max im iz ing t o ta l p roduc t iv i ty w h i le a low ing f o r c on t inued h e rd g row th , o n ly i n f i rm o r i n fe r t i le f ema les s hou ld n o rma l ly b e c u l led f r om t he h e rd wh i le a l f e r t i le f ema les wou ld b e r e ta ined . H oweve r , ma les s u rv iv ing t h rough a do lescence , e nd ing i na ge c l ass 5 -6 , s hou ld o n ly b e r e ta ined a s r equ i red f o rs t ud a nd b aggage s erv ice . T h is s t ra tegy w i l l b e t e rmed " no rma l c u l l ing ." I ft he managemen t s t ra tegy i n s tead i n vo lved max im iz ing t o ta l p roduc t iv i ty t h rough z e ro 1 72

h e rd g row th , a do lescen t f ema les i ne xcess o ft he n umbe r r equ i red t o ma in ta in ac ons tan t h e rd s i ze w ou ld b e a dd i t iona l ly s ub jec tt oc ropp ing . T h is s t ra tegy w i l l b et e rmed " max imum c u l l ing . " C ons is ten t w i th c on t inued h e rd g row th u nde r n o rma l c u l l ing s t ra teg ies , g row th i nt he f ema le p opu la t ion o f ah e rd w i l l b e s imu la ted f o r a l a ge c l asses , a ssum ing t ha t a l s u rv iv ing f ema les a re l a s t b red a tt he e nd o f t he i r t wen ty -second y ea r . H oweve r , g row th i nt he ma le p opu la t ion o f ah e rd w i l l o n ly b e s imu la ted t h rough a ge c l ass 5 -6 . T he s ubsequen t c u l l ing o r r e ten t ion o f ma les s u rv iv ing a ge c l ass 5 -6 w i l l b e a dd ressed l a te r i ne s t ima t ing t o ta l o f take i n e xcess a do lescen t ma les f r om ac ame l h e rd o fk nown s i ze a nd a ge -sex c ompos i t ion . B e fore p roceed ing t o t he s imu la t ions , i ts hou ld b e n o ted t ha t w h i le t he l a te r e ten t ion o f ma les u nde r n orma l a nd max imum c u l l ing s t ra teg ies a s d e f ined a bove w ou ld s e rve t o max im ize t o ta l s l augh te r p roduc t iv i ty , i t wou ld p o ten t ia l ly d o s o a t t he e xpense o f t o ta l h e rd m i lk p roduc t ion . F o re xamp le , c ame l p as to ra l is ts w i l l o f ten k i l l an ew ly -bo rn ma le c a l f a nd s ubs t i tu te a f o s te r -ca l f i ni t s p l ace i no rde rt oa ch ieve ac on t inued f l ow o fm i lk f r om s eve ra lf ema les w h i le o n ly o ne c a l f i sa c tua l ly n u rs ing ( Co le 1 975 :27 , 4 9 ; Swee t 1 970 :276 ) . I nt h is f ash ion , t he p o ten t ia l y i e ld o f m i lk f o r h uman c onsump t ion c an a ppa ren t ly b e i n c reased . Mus i l made s eve ra l r e fe rences t ot he u se o ft h is s t ra tegy among t he Rwa la ( 1928 :22 , 8 7 -88 , 2 09 ) , w h i le I b n S a 'ud , f ounde r o f t he mode rn K ingdom o f S a 'ud i A rab ia , p e rsona l ly e xp la ined t he p rac t ice t o P h i lby ( 1973 :54 ) . A ppa ren t ly , a managemen t s t ra tegy o r ien ted t owa rds max im iz ing m i lk p roduc t iv i ty a tt he e xpense o f s l augh te r p roduc t iv i ty i sq u i te c ommon among c er ta in c ame l h e rd ing p opu la t ions . T h is a l te rna t ive managemen t s t ra tegy w i l l n o t b e s imu la ted i nt he p resen t a na lys is .

F ema le P opu la t ion G row th F rom a F ound ing H e rd C on ta in ing T en R ep roduc t ive ly Ma tu re F ema les G row th S imu la t ion V a lues T he mor ta l i ty v a lues u sed i nt he p resen t mode l o f g row th i nt he f ema le p opu la t ion o f a c ame l h e rd a re p resen ted i nT ab le 4 8 . T he f i r s t c o lumn o ft he t ab le , l a be led p (x ) , p resen ts t he p ropo r t iona l s urv ivo rsh ip o ff ema les f r om a ge xt ox +1 y ea rs a s p rev ious ly p resen ted i nT ab le 4 7 . H oweve r , t he s u rv ivo rsh ip v a lues f o r a ge c l asses 2 1-22 a nd 2 2 -23 h ave b een c o r rec ted t o a ccoun t f o r h uman c ropp ing , a s p rev ious ly d i scussed . T he s econd c o lumn , l a be led 1 ( x ) , p resen ts t he p ropo r t ion o ff ema les l i v ing f r om b i r th t oa ge x ,a nd was c a lcu la ted f r om t he v a lues g i ven i nc o lumn p ( x ) . S ince a 0 .39 y ean ing r a te i sa ssumed f o r a l b reed ing f ema les i nt he a bsence o f a ge s pec i f ic r a tes , t he 4 9 :51 ma le t of ema le s ex r a t io i mp l ies t ha t 0 . 199 f ema le c a lves s hou ld b e p roduced a nnua l ly b y e ach f ema le o fb reed ing a ge .

F ema le P opu la t ion G row th S imu la t ion T he r ep roduc t ion a nd m or ta l i ty v a lues g i ven a bove h ave b een u sed t o s imu la te e xpec ted g row th o ve r a4 0 y ea r p e r iod , s t a r t ing w i th af ound ing p opu la t ion o f 1 0 r ep roduc t ive ly ma ture f ema les r ang ing i na ge f r om 7t o 1 6 y ea rs ( Append ix C ) . T he y ounges t o ft hese i si na ge c l ass 7 -8 , w h i le t he o ldes t i si na ge c l ass 1 6 17 . S ince t he r ep roduc t ive ly ma ture f ema les i nt h is i n t ia l f ound ing p opu la t ion r ep resen t s urv ivo rs f r om e a r l ie r n ew ly -bo rn f ema le g roups o f u nknown s i ze , t he i r s ubsequen t s ur v ivorsh ip i sb ased u pon t he v a lues i nc o lumn p (x ) o f T ab le 4 8 a s r e ca lcu la ted t o r ep resen t i n d iv idua l s urv ivo rsh ip i n to s ubsequen t y ea rs . F ema les s u rv iv ing i n to a ge c l ass 2 2 -23 w ou ld n o t c on t r ibu te f u r the ro fsp r ing . T he g row th s imu la t ion b eg ins a tt he e nd o fy ea r 0 .S ince a l b i r ths i nt he h e rd a re e xpec ted t oo ccu r a tt he e nd o fe ach y ea r , t he f ound ing p opu la t ion o f 1 0 f ema le c ame ls wou ld g i ve b i r th t of ema le o f fsp r ing a tt he e nd o fy ea r0 .W i th ay ean ing r a te o f0 .39 a nd as ex r a t io o f4 9 ma les t o5 1 f ema les , e ach o ft he 1 0 r ep roduc t ive f ema les i nt he f ound ing h e rd w ou ld p roduce 0 . 199 f ema le o fsp r ing ( 0 .39 x 0 . 51 = 0 . 199 ) . T hus , t he 1 0 r ep roduc t ive f ema les wou ld h ave 1 . 99 n ew ly -bo rn f ema le o f fsp r ing a tt he e nd o fy ea r0 ,f o r at o ta l p opu la t ion o f1 .99 f ema les .

1 73

T ab le 4 8 F ema le Mo r ta l i ty V a lues

A ge C lass

p ( x )

0 1 1 -2 2 -3 3 -4 4 -5 5 -6 6 -7 7 -8 8 -9 9 -10 1 0 -11 1-12 1 2 -13 1 3 -14 1 4 -15 1 5 -16 1 6 -17 1 7 -18 1 8 -19 1 9 -20 2 0 -21 2 1-22 2 2 -23

. 82 . 94 . 96 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 97 . 48 . 00

1 . 000 . 820 . 771 . 740 . 7 18 . 696 . 675 . 655 . 635 . 6 16 . 598 . 580 . 563 . 546 . 529 . 5 13 . 498 . 483 . 469 . 455 . 441 . 428 . 2 10

A tt he b eg inn ing o fy ea r 1i nt he s imu la t ion , e ach o ft he o r ig ina l f ound ing f ema les d ec l ines 3% f r om n a tu ra l mo r ta l i ty b e fo re e n te r ing t he s ubsequen t a ge c l ass . T hus , t he 1f ema le i na ge c l ass 7 -8 i ny ea r 0d ec l ines b y 3% , l e av ing 0 .97 f ema les t oe n te r a ge c l ass 8 -9 o fy ea r 1 .T he s ame o ccurs w i th t he o the r 9 r ep roduc t ive ly ma ture f ema les . T hus , t he o r ig ina l 1 0 r ep roduc t ive ly ma tu re f ema les i ny ea r 0d ec l ine f r om n a tu ra l mo r ta l i ty , l e av ing 9 .7 r ep roduc t ive ly ma tu re f ema les a tt he s t a r t o fy ea r 1 .S im i la r ly , t he 1 . 99 n ew ly -bo rn f ema les i na ge c l ass 0 1 o f y ea r 0d ec l ine f r om n a tu ra l mo r ta l i ty l e av ing 1 . 63 f ema les t oe n te r a ge c l ass 1 -2 i ny ea r 1 .H e re , t he e qua t ion i s1 . 99 i n d iv idua ls i na ge -c lass 0 1 t imes t he p ropo r t iona l s urv ivorsh ip i n to a ge c l ass 1 2 a s g i ven i nc o lumn 1 ( x ) o fT ab le 4 8 , o r 1 . 99 x0 .82 = 1 . 63 . A tt he e nd o fy ea r 1 ,t h e 9 .7 r ep roduc t ive ly ma tu re f ema les a ga in g i ve b i r th , p roduc ing 1 . 93 f ema le o f fsp r ing ( 9 .7 x0 . 199 = 1 . 93 ) . T hus , a t t he e nd o f y ea r 1 ,t he t o ta l f ema le p opu la t ion o f t he h erd c ons is ts o f 1 3 .26 i n d iv idua ls . T h is p rocess i st hen r epea ted f o r e ach s ubsequen t y ea r . A t t he e nd o f y ea r 1 ,t h e 9 .7 r ep roduc t ive ly ma ture f ema les i na ge c l asses 8 -9 t h rough 1 7 -18 d ec l ine b y n a tu ra l m o r ta l i ty ( 3% ) , l e av ing 9 .4 r ep roduc t ive ly ma tu re f ema les t o e n te r a ge c l asses 9 10 t h rough 1 8 19 i n y ea r 2 .T he 1 . 93 f ema les i na ge c l ass 0 1 o fy ea r 1d ec l ine b y n a tu ra l mor ta l i ty ( 18% ) , l e av ing 1 . 58 f ema les t oe n te ra ge c l ass 1 2 o fy ea r2 .S im i la r ly ,t he 1 . 63 f ema les i na ge c l ass 1 -2 o fy ea r 1d ec l ine b y n a tu ra l mo r ta l i ty ( 6% ) , l e av ing 1 . 53 f ema les t oe n te r a ge c l ass 2 -3 o fy ea r2 .A tt he e nd o fy ea r 2 ,t he 9 .4 r ep roduc t ive ly ma ture f ema les a ga in g i ve b i r th , a dd ing 1 . 87 n ew ly -born f ema les t oa ge c l ass 0 1 o fy ea r2 .T hus , a tt he e nd o fy ea r2 ,t he t o ta l f ema le p opu la t ion o ft he h e rd c ons is ts o f 1 4 .38 i n d iv idua ls .

S imu la t ion R esu l ts T he a ge s pec i f ic g row th r esu l ts o ve r t he 4 0 y ea r p e r iod o ft he s imu la t ion a re p resen ted i n A ppend ix C . D u r ing t he 4 0 y ea rs o ft he s imu la t ion , t he t o ta l f ema le p opu la t ion i n c reased 5 .66 t imes , w i th a mean a nnua l g row th r a te o f4 .5% . T he t o ta l f ema le p opu la t ion f i r s t d oub led i nt he 1 74

e i gh teen th y ea r a nd d oub led a ga in i n t he t h ir ty four th y ea r . H oweve r , t he n umber o f r ep roduc t ive ly ma ture a nd p o ten t ia l ly m i lk p roduc ing f ema les d i d n o t d oub le u n t i l t he t wen ty s even th y ea r . I nt he s ubsequen t y ea r , t he n umbe r o ff ema les i na ge c l ass 2 2 -23 f i r s t b egan t o a nnua l ly i n c rease , b e ing t he l a s t a ge c l ass t o d o s o . F rom t h is y ea r t h rough t he e nd o f t he s imu la t ion ( 13 t o ta l y ea rs ) , m ean a nnua l g row th r ema ined c ons tan t a t4 .5% , t he o bserved mean f o r t he e n t i re 4 0 y ea r p e r iod . T he p ropor t iona l r ep resen ta t ion o f r ep roduc t ive ly i mma ture ( age c l asses 0 -6 ) a nd ma tu re f ema les ( age c l asses 6 -23 ) s t ab i l ized a t4 6% a nd 5 4% r e spec t ive ly i n t h et wen ty th i rd y ea r , a nd d i d n o t d ev ia te f r om t hese v a lues t h rough t he e nd o ft he s imu la t ion .

C ompa r isons W i th P rev ious Mode ls A s p rev ious ly d i scussed , D ah l a nd H jor t ( 1976 :83 -84 ) g enera ted t wo s imu la t ions o f e xpec ted g row th i nt he f ema le p opu la t ion o f a mode l c ame l h erd b ased u pon t wo d i s t inc t m o r ta l i ty c urves a nd a0 .5 y ean ing r a te . I nt he i rf i r s t s imu la t ion , a nnua l h e rd g row th was 7 .5 % , w i th t he t o ta l f ema le p opu la t ion d oub l ing i n1 y ea rs a nd t he n umbe r o fr ep roduc t ive ly ma ture f ema les d oub l ing i n 1 5 y ea rs . I nt he i r s econd mode l , wh ich a ssumed much h i ghe r mor ta l i ty r a tes , a nnua l h e rd g row th w as o n ly 1 . 5% , w i th t he t o ta l f ema le p opu la t ion d oub l ing i n5 1 y ea rs a nd t he n u mbe r o f r ep roduc t ive ly ma ture f ema les d oub l ing " i n we l l o ve r 5 0 y ea rs . " I nt e rms o f t he c ond i t ions e xpec ted u nde r e x tens ive h usband ry , i ti ss ugges ted t ha t t he i r f i rs t mode l w ou ld a pp rox ima te e xpec ta t ions u nde r e xcep t iona l ly f avo rab le c ond i t ions , wh i le t he r e su l ts o f t he i r s econd mode l wou ld b e more a pp rop r ia te f o r e xpec ted g row th u nder e x t re me e nv i ronmen ta l c ond i t ions . I nS a to 's m ode lo fc ame l h e rd g row th ( 1 980 :28 -3 1) , i twas a ssumed t ha to n ly 0 . 109 f ema le c a lves w ou ld b e e xpec ted a nnua l ly p e r r ep roduc t ive f ema le i nt he h e rd . H oweve r , t h is l ow y ean ing r a te was p a r t ia l ly o f fse t b y S a to 's u se o f much l owe r mor ta l i ty r a tes t han t hose a ssumed i nt he p resen t mode l . A s ar e su l t , S a to c a lcu la ted t ha t a nnua l g row th i nt he f ema le c omponen t o f ac ame l h e rd w ou ld b e a round 3 .4% , af i gure o n ly s l igh t ly l owe r t han t ha t d e r ived i nt he p resen t s imu la t ion . S pec i f ica l ly , S a to 's mode l s eems mos t a pp rop r ia te f o r c ame l h e rds e xpe r ienc ing l ow f e r t i l i ty r a tes , b u t c u r ren t ly e x is t ing u nde r c ond i t ions i nw h ich i n fan t mor ta l i ty h as b een r educed t ot he l e ve ls e xpec ted u nde r c ond i t ions o fi n tens ive h usband ry .

I mma tu re Ma le P opu la t ion G row th F rom aF ound ing H e rd C on ta in ing T en R ep roduc t ive ly Ma tu re F ema les G row th S imu la t ion V a lues T he mor ta l i ty v a lues u sed i nt he p resen t mode l o fg row th i nt he ma le p opu la t ion o f ac ame l h e rd t h rough a ge c l ass 5 -6 a re p resen ted i nT ab le 4 9 . T hese v a lues w ere d e r ived i nt he s ame f a sh ion a s t he f ema le mo r ta l i ty v a lues p rev ious ly p resen ted i nT ab le 4 8 . T he 0 .39 y ean ing r a te a ssumed f o r a l b reed ing f ema les a nd t he 4 9 :5 1 ma le t of ema le s ex r a t io i mp l ies t ha t 0 . 191 ma le c a lves s hou ld b e p roduced a nnua l ly b y e ach f ema le o fb reed ing a ge .

T ab le 4 9 I mma tu re Ma le Mo r ta l i ty V a lues

A ge C lass

p ( x )

0 1 1 -2 2 -3 3 -4 4 -5 5 -6

. 66 . 89 . 93 . 97 . 97 . 97

1 75

1 .000 . 660 . 587 . 546 . 530 . 5 14

Ma le P opu la t ion G row th S imu la t ion T he b i r th a nd mor ta l i ty v a lues g i ven a bove h ave b een u sed t o s imu la te e xpec ted g row th o ve r a4 0 y ea r p e r iod ( Append ix D ) , a ga in a ssum ing t he s ame f ound ing p opu la t ion o f 1 0 r ep roduc t ive ly ma ture f ema les a s b e fo re . G row th i n t he ma le c omponen t o f t he h e rd i s c a lcu la ted i nt he s ame f a sh ion a s g row th i nt he f ema le c omponen t o ft he h erd . T he s imu la t ion a ga in b eg ins a t t he e nd o f y ea r 0 ,w hen t he 1 0 r ep roduc t ive ly ma tu re f ema les g i ve b i r th t o ma le o f fsp r ing . W i th ay ean ing r a te o f0 . 39 a nd as ex r a t io o f4 9 ma les t o5 1 f ema les , e ach o f t he 1 0 r ep roduc t ive f ema les i nt he f ound ing h e rd i se xpec ted t o p roduce 0 . 191 ma le o f fsp r ing ( 0 .39 x 0 .49 = 0 . 191) . T he i mma tu re ma le p opu la t ion o f t he h e rd t he re fo re c ons is ts o f1 . 9 1 ma les i na ge c l ass 0 1 a tt he e nd o fy ea r0 . A tt he b eg inn ing o fy ea r 1i nt he s imu la t ion , e ach o ft he 1 . 91 ma les i na ge c l ass 0 1 o fy ea r 0d ec l ine f r om n a tu ra l mo r ta l i ty , l e av ing 1 . 26 i n d iv idua ls t oe n te r a ge c l ass 1 -2 o fy ea r 1 .H e re , t he e qua t ion i s1 . 91 i n d iv idua ls i na ge c l ass 0 1 t imes t he p ropor t iona l s urv ivo rsh ip i n to a ge c l ass 1 -2 a s g i ven i nc o lumn 1 ( x ) o fT ab le 4 9 ( 0 .660 ) , o r 1 . 9 1 x0 .660 = 1 . 26 . T he o r ig ina l 1 0 r ep roduc t ive f ema les a tt he e nd o fy ea r 0o ft he s imu la t ion d ec l ine f r om n a tu ra l mo r ta l i ty d ur ing y ea r 1t o9 .7 f ema les . A t t he e nd o fy ea r 1 ,t hese 9 .7 s urv iv ing f ema les a ga in g i ve b i r th t o ma le o f fsp r ing , y i e ld ing 1 . 85 ma le o f fsp r ing i na ge c l ass 0 1 o fy ea r o ne . T hus , a tt he e nd o fy ea r 1 , t he i mma tu re ma le p opu la t ion c ons is ts o f 2 .76 i n d iv idua ls . T h is s ame s equence i st hen r epea ted f o r e ach s ubsequen t y ea r o ft he s imu la t ion .

S imu la t ion R esu l ts T he a ge s pec i f ic g row th r esu l ts o ve r t he 4 0 y ea rs o f t he s imu la t ion a re p resen ted i n A ppend ix D ,w h i le a n i l us t ra t ion o fs imu la ted g row th i nb o th t he i mma tu re ma le a nd t o ta lf ema le p opu la t ions o ft he h e rd i sp rov ided i nF i gu re 2 2 . D u r ing t he 4 0 y ea rs o ft he s imu la t ion , t h e t o ta l i mma ture ma le p opu la t ion i n c reased 1 2 .7 t imes w i th a mean a nnua l g row th o f 6 .7% . T he t o ta l i mma ture ma le p opu la t ion f i r s t d oub led i nt he s econd y ea r , a nd s ubsequen t ly d oub led e very 1 5 y ea rs . D u r ing t he l a s t 1 3 y ea rs o ft he s imu la t ion , mean a nnua l g row th was a t4 .6% . N e i the r D ah l a nd H jo r t ( 1976 ) n o r S a to ( 1980 ) mode l led g row th f o rt he i mma tu re ma le c omponen t o f a h e rd .

T he A ge a nd S ex S t ruc tu re o fa n E s tab l ished C ame l H e rd G ene ra lC ons idera t ions T he a ge d i s t r ibu t ion o f ah e rd i st ha t p ropo r t ion o ft he t o ta l h e rd f o rmed b yt he m embers o f e ach a ge c l ass . S o l o ng a sf e r t i l i ty a nd mor ta l i ty r a tes r ema in c ons tan t , a l c on t inuous ly b reed ing a n ima ls t end a symp to t ica l ly o ve r t ime t o ar ecur r ing o r s t ab le a ge d i s tr ibu t ion . S easona l b reede rs may a t ta in e i the r ap a t te rn o fr epea ted c yc l ing o fa ge d i s t r ibu t ion , o r , a s w i th h e rded s heep a nd g oa ts , as t ab le a ge d i s t r ibu t ion ( Redd ing 1 981:129 -130 ) . S im i la r ly , t he s ex d i s t r ibu t ion o f ah e rd i st ha t p ropo r t ion o ft he t o ta l h e rd f o rmed b y t he members o f e ach s ex . H e re , t he h uman a d jus tmen t o f s ex r a t io i n ah e rd t h rough s e lec t ive c u l l ing e f fec t ive ly d e te rm ines t he r esu l t ing p a t te rn . U s ing s uch d a ta , t he a ge a nd s ex s t ruc tu re o f ah e rd o f k nown s i ze c an b e c a lcu la ted . F ur the r , s i nce t he p roduc t iv i ty o f i n d iv idua l a n ima ls v a r ies a cco rd ing t o t he i r a ge a nd s ex s pec i f ic a t t r ibu tes , t hese d a ta a low f o r amo re a ccu ra te a ccoun t ing o ft he p roduc t iv i ty e xpec ted f r om a mode l h e rd t h rough b o th c u l l ing s t ra teg ies a nd t he u t i l iza t ion o fa n ima ls wh ich d i e f r om n a tu ra lc auses .

F ema le A ge D i s t r ibu t ion i n aC ame lH e rd A s p rev ious ly n o ted , t he p ropo r t iona l r ep resen ta t ion o f r ep roduc t ive ly i mma ture a nd ma tu re f ema les i nt he g row th s imu la t ion s t ab i l ized a t4 6% a nd 5 4% r espec t ive ly i nt h e t wen ty t h i rd y ea r , a nd r ema ined a t t hese l e ve ls t h rough t he e nd o ft he s imu la t ion . T he a ge -spec i f ic 1 76

70 —

NU MBER OF I ND IV IDUALS

60 —

50 —

t ota l f e ma les

40 —

30 —

20 —

1 0 —

0 0

I m mature ma les

I 5

I

1 0

i

I

1 5

20

I

25

I

30

I 35

1 40

NUMBER OF YEARS F i gu re 2 2 . S imu la ted g row th i nt he t o ta l f ema le a nd i mma tu re ma le c omponen ts o f ap as to ra l c ame l h e rd .

d i s t r ibu t ion , h owever , d id n o t a pp roach s t ab i l i ty u n t i l t he t wen ty -seven th y ea r . T ab le 5 0 p resen ts t he mean a ge d i s t r ibu t ion a ch ieved f r om t he t wen ty -seven th y ea r o f t he s imu la t ion t h rough t he f o r t ie th y ea r ( 14 y ea rs t o ta l ) . Wh i le t he r e la t ive p ropor t ions o f t he p r ima ry a ge c l asses 0 -6 a nd 6 -23 r ema ined s tab le t h roughou t t h is p e r iod , a ge -spec i f ic p ropo r t iona l d ev ia t ions o f 1% i n a ge c l asses 7 -8 t h rough 1 7 -18 d id o ccur , p resumab ly d ue t o r ound ing e r ro r .

I mma tu re Ma le A ge D is t r ibu t ion i n aC ame lH erd T he a ge -spec i f ic p ropor t iona l r ep resen ta t ion o f i mma ture ma les i n t he g row th s imu la t ion a ch ieved s tab i l i ty i nt he t h ir ty -second y ea r . T ab le 5 1 p resen ts t h is s t ab le a ge d i s t r ibu t ion . F rom t he t h ir ty -second y ea r t h rough t he e nd o f t he s imu la t ion , t he p ropor t iona l r ep resen ta t ion o f t hese a ge c l asses v a r ied f r om y ea rt oy ea r b y a round 0 . 1% , a ga in p resumab ly d ue t o r ound ing e r ro r .

A Mode l C ame l H e rd I no rder t o c a lcu la te a nnua l o f f take ( s laugh te r y i e ld ) , i tw i l l b e n ecessary t o e s tab l ish a mode l h e rd o f s pec i f ic s i ze a nd a ge -sex c ompos i t ion . P re fe rab ly , t h is mode l h e rd s hou ld b e o f " ave rage " s i ze a cco rd ing t o e thnog raph ic a ccoun ts . H oweve r , a s s hou ld b e e xpec ted , b o th e thnog raph ic a nd h i s to r ica l a ccoun ts o f t he s i ze o f c ame l h e rds ma in ta ined b y n omad ic p as to ra l is ts v a ry w ide ly i nb o th t i me a nd s pace .

1 77

T ab le 5 0 Mean F ema le A ge D is t r ibu t ions D e r ived f r om t he L as t 1 4Y ea rs o ft he S imu la t ion

A ge C lass

%o fT o ta l P opu la t ion

0 1 1 -2 2 -3 3 -4 4 -5 5 -6

1 8 8 7 6 6

6 -7 7 -8 8 -9 9 -10 1 0 -11 1-12 1 2 -13 1 3 -14 1 4 -15 1 5 -16 1 6 -17 1 7 -18 1 8 -19 1 9 -20 2 0 -21 2 1-22 2 2 -23

6 5 5 4 4 4 4 3 3 3 3 2 2 2 2 2 1

5 4

T o ta l

1 01

1 00

4 6

T ab le 5 1 I mma tu re Ma le S tab le A ge D i s t r ibu t ion

A ge C lass

%o fT o ta l P opu la t ion

0 1 1 -2 2 -3 3 -4 4 -5 5 -6

2 9 1 8 1 5 1 4 1 3 1 2

T o ta l

1 01

I n many e thnog raph ic a ccoun ts , a n a t temp t was made t oe s tab l ish t he m in ima l n umber o f h e rd a n ima ls r equ i red t o s uppor t as i ng le b edou in f am i ly . T hus , Swee t ( 1969 :160 16 1 ) n o ted t ha t h e r b edou in i n fo rman t i nK uwa i t r ega rded h imse l f a s w e l l -o f f w i th a h e rd c on ta in ing 1 8 ma ture f ema les . Swee t a l so n o ted t ha t e a r ly t wen t ie th c en tury B r i t ish e s t ima tes o ft r i ba l h erd s t reng th a l lowed 2 0 c ame ls a s a n a ve rage f i gu re p er n omad ic t en t . S im i la r ly , L ancas te r ( 1981:99 , 1 39 -140 ) n o ted t ha t 1 5 t o 2 0 c ame ls ( sex n o t s t a ted ) w ere c ons ide red a s t he 1 78

m in imum n umbe r r equ i red t o s uppor t as i x membe r Rwa la f am i ly . A cco rd ing t o J ohnson ( 1969 :10 ) , c ame l h e rd ing b edou in i nt he S aha ra m in ima l ly r equ i re 1 0 t o 2 5 h ead ( sex n o t s pec i f ied ) t o s uppor t o ne f am i ly . Ah e rd c on ta in ing 2 0 ma le a nd f ema le c ame ls was a l so s ugges ted b y G au th ie r -Pee rs a nd D agg ( 198 1 :14 1) a st he m in imum s i ze r equ i red t os uppor t a f am i ly . A cco rd ing t oA sad ( 1 970 :52 ) , t he K abab ish b edou in o ft he S udan m in ima l ly r equ i red 2 0 t o2 5 h ead ( bo th s exes i mp l ied ) t os us ta in a h ouseho ld . Wh i le h e rds c on ta in ing 1 0 t o 2 5 r ep roduc t ive ly ma ture f ema les wou ld s eem t o b e t he m in imum r equ i red f o r s us ta in ing a s i ng le f am i ly , a ve rage h e rds a ppea r t o b e much l a rger . Among t he R end i l le o fN or the rn K enya , c ame l h e rd s i zes w e re o bse rved t or ange f r om 2 4t o7 3 h ead , w i th a n a ve rage h e rd s i ze o f4 9 h ead ( Sa to 1 980 :28 ) . H oweve r , t he R end i l le a l so k eep h e rds o f s heep a nd g oa ts . Among t he A hagga r T ua reg , a" sma l l h e rd " c on ta ined 2 0 t o 4 0 h ead , wh i le a " very l a rge h e rd " m in ima l ly c on ta ined a round 1 00 h ead , r ega rd less o f s ex ( N ico la isen 1 963 :101) . Among t he K abab ish b edou in o ft he S udan , t he max imum n umbe r o f c ame ls o bserved i n ah erd w as 1 50 h ead ( Asad 1 964 :52 ; 1 970 : 16 ) . Among t he Mu ta i r b edou in o fK uwa i t , D ickson ( 195 1:4 13 ) n o ted t ha t e ach h e rd c on ta ined a round 7 0 c ame ls ( s ex n o t s pec i f ied ) . F o rt he Rwa la b edou in o fN o r th A rab ia , Mus i l ( 1928 :336 ) n o ted t ha t l a rge h e rds c on ta ined 7 0 t o 8 0 h ead ( s ex n o t s pec i f ied ) . U n for tuna te ly , i ti su nc lea r w he the r t hese f i gu res r e fe r t o e n t i re h e rds , o r o n ly h e rd s ubd iv is ions . S ince T o r ry ( 1974 :67 ) n o ted t ha t t he l a rges t manageab le c ame l h e rd d i v is ions among t he Gab ra o f E as t A f r ica r a re ly e xceeded 7 0 a n ima ls , i ti ss uspec ted t ha t D ickson a nd Mus i l a l so r eco rded t he s i ze o f m anageab le h e rd s ubd iv is ions , n o t e n t i re h e rds . S ome s uppor t f o r t h is s ugges t ion i sf ound i n t h e d a ta p resen ted b y C o le ( 1 975 :36 ) o nt he s i ze o fh e rds among t he A l' A zab s ec t ion o ft he A l M u r rah b edou in o fS a 'ud i A rab ia . H e re , c ame l h e rds w e re o bse rved t oc on ta in b e tween 4 0 t o 7 5 ma tu re f ema les , w i th 5 0 m a tu re f ema les b e ing a ve rage . M uch l a rge rc ame l h e rds h ave a l so b een r epo r ted . A cco rd ing t o Gau th ie r -P i l te rs a nd D agg ( 1 98 1 :14 1) , s ome R egu iba t b edou in i nt he n or thwes te rn S aha ra i n1 97 1 h ad h e rds o f 1 , 000 o r m o re c ame ls . F o r t he p resen t s t udy , h oweve r , a h erd c on ta in ing a pp rox ima te ly 5 0 r ep roduc t ive ly ma tu re f ema les s eems mo re a pp rop r ia te f o r i n t ia l mode l l ing . S ince t he 5 5 r ep roduc t ive ly ma ture f ema les e xpec ted i n ah e rd c on ta in ing 1 01 t o ta l f ema les a s p rev ious ly p resen ted i nT ab le 5 0 r ough ly a pp rox ima tes t he a ve rage n umbe ro fa du l t f ema les i nt he h e rds o ft r ad i t iona l c ame l b edou in i nS a 'ud i A rab ia , i n t ia l c a lcu la t ions o fa nnua l o f f take w i l l b ased u pon t hese f i gu res . C oup led w i th t he d a ta o n i mma ture ma les p resen ted i n T ab le 5 1 , t he a ge a nd s ex s t ruc tu re o ft he mode l c ame l h e rd wou ld b e t ha t p resen ted i nT ab le 5 2 . T he mode l h erd c ons is ts o f a pp rox ima te ly 4 0% r ep roduc t ive ly ma ture f ema les , 3 4% i mma ture f ema les , a nd 2 6% i mma ture ma les , e xc lud ing a ny a du l t ma les k ep t f o r s t ud a nd b aggage p urposes . I ft hese v a lues a re a pp l icab le t oA l Mu r rah h e rds c on ta in ing 4 0t o7 5 ma tu re f ema les , t he t o ta l s i ze o f t he i r h e rds s hou ld r ange f r om a pp rox ima te ly 9 9 t o 1 86 h ead , w i th a ve rage h e rds c on ta in ing a pp rox ima te ly 1 24 h ead , a ga in e xc lud ing a du l t ma les . U s ing t he 0 .39 y ean ing r a te e s tab l ished e a r l ier , t he mode l h erd wou ld c on ta in a pp rox ima te ly 2 2 l a c ta t ing f ema les a tt he b eg inn ing o ft he y ea r ( 55 ma ture f ema les x 0 .39 = 2 1 .45 ) , a nd t he r ep roduc t ive ly ma ture f ema les s urv iv ing a tt he e nd o f t he y ea r wou ld t hen p roduce a pp rox ima te ly 1 f ema le a nd 1 0 ma le c a lves . F ur the r , t he mode l h e rd wou ld c on ta in f ou r ma les wh ich w ou ld b e e n te r ing a ge c l ass 6 -7 a tt he e nd o ft he y ea r , a nd t hus b e s ub jec tt o c u l l ing . H oweve r , i no rde rt oc a lcu la te t he t o ta l e xpec ted o f f take f r om t h is h erd o ve r ao ne y ea r p e r iod , i ti sf i rs t n ecessa ry t oe s tab l ish t he n umbe ro f ma les e n te r ing a ge c l ass 6 -7 wh ich w ou ld h ave t ob e r e ta ined f o rs ubsequen t s t ud a nd b aggage p u rposes . F ur the r , i tw i l l b e n ecessa ry t o a ccoun t f o r a dd i t iona l o f f take f r om t hose a n ima ls s chedu led t od i e o fn a tu ra l c auses d ur ing t he y ea r . T hese s ub jec ts a re a dd ressed i nt he f o l low ing s ec t ions .

T he R e ten t ion o fA do lescen t Ma les i nt he Mode l C ame l H e rd I no rde r t oc a lcu la te t he a c tua l n umbe r o fa do lescen t ma les e n te r ing a ge c l ass 6 -7 w h ich w ou ld b e a va i lab le f o r s l augh te r , i ti sf i r s t n ecessary t o e s tab l ish h ow many wou ld h ave t o b e r e ta ined f o r s t ud a nd b aggage p urposes .

1 79

T ab le 5 2 E xpec ted A ge a nd S ex S t ruc tu re o fa Mode l C ame l H e rd C on ta in ing 5 5 R ep roduc t ive ly Ma tu re F ema les ( Exc lud ing A du l t Ma les )

A ge C lass

F ema le

Ma le

0 1 1 -2 2 -3 3 -4 4 -5 5 -6 6 -7 7 -8 8 -9 9 10 1 0 -11 1-12 1 2 -13 1 3 -14 1 4 -15 1 5 -16 1 6 -17 1 7 -18 1 8 -19 1 9 -20 2 0 -21 2 1-22 2 2 -23

1 8 8 7 6 6 6 5 5 4 4 4 4 3 3 3 3 2 2 2 2 2 1

1 0 6 5 5 5 4

T o ta l

1 01

3 5

T he R e ten t ion o f Ma les a s S tuds Wh i le t he a va i lab le d a ta o n t he n umbe r o fs t ud c ame ls k ep t i nh e rds v a r ies , i ti sr ead i ly a ppa ren t t ha t v e ry f ew a re a c tua l ly r equ i red . T ab le 5 3 s umma r izes s ome o ft he p ub l ished d a ta o n t h is s ub jec t . I tw i l l b e a ssumed h e re t ha t a tl e as t o ne s t ud ma le i sm in ima l ly r equ i red p e r3 0 r ep roduc t ive ly ma tu re f ema les , a nd t hus t he mode l h e rd w ou ld r equ i re a pp rox ima te ly t wo s t ud c ame ls .

T he R e ten t ion o f Ma les f o r T ranspo r t A cco rd ing t o C o le 's d a ta ( 1975 :37 -38 ) , t he n i ne h ouseho lds o ft he A l K hurb i s ubd iv is ion o f t he A l ' Azab s ec t ion o f t he A l Mur rah b edou in k ep t b e tween 2 0 a nd 3 5 ma le b aggage c a r r ie rs . T hus , t wo t of ou r b aggage c a r r ie rs w e re r equ i red p e rh ouseho ld . T h is a g rees w e l lw i th t he o bserva t ions o f Gau th ie r -P i l te rs a nd D agg ( 1981:137 ) i n N or th A f r ica , w ho n o ted t ha t mov ing a h ouseho ld c amp " i nvo lves t wo t o f our c ame ls c a r ry ing t he t en t , s add les , s ome c a rpe ts , o ne o r t wo w ooden o r me ta l c hes ts , w oo len o r l e a the r b ags c on ta in ing c l o thes a nd p rov is ions , g uerbas , wa te rd rums , a nd t he c ook ing p o t . " l tw i l l t he re fo re b e a ssumed t ha t i nt he mode l c ame l h e rd t he re w ou ld b e t h ree t o f ou r ma tu re ma les k ep t a s c a r r iers . A ny e xcess b aggage r equ i remen ts c ou ld b e c ove red b y u s ing e i the r y ounge r ma les o r i n d iv idua ls d rawn f r om t he f ema le p opu la t ion o ft he h e rd . I nt he mode l h e rd , at o ta l o ff i ve t o s i x ma tu re ma les wou ld t he re fo re n eed t o b e ma in ta ined . E ach y ea r , n a tu ra l mor ta l i ty a nd t he c u l l ing o fo lde r ma les wou ld r educe t he n umber o f ma ture ma les , t hus r equ i r ing t he r e ten t ion o f ac e r ta in n umbe ro f ma les e n te r ing a ge c l ass 6 -7 . 1 80

T ab le 5 3 N umbe ro fF ema le C ame ls w h ich c an b e S e rv iced b y aS ing le S tud

N o . o fF ema les

L oca t ion

3 0 t o5 0 6 0 t o8 0 1 00 7 0 3 0 t o7 0 1 5 t o5 0 2 5 t o5 0 3 0 t o5 0 5 0 5 0 t o8 0

S udan T un is ia K uwa i t g ene ra l g ene ra l g ene ra l K enya K enya g ene ra l P ak is tan

S ource A sad 1 970 :17 B u rgeme is ter , i nK noess 1 977 :40 D ickson 1 951:4 11 Gau th ie r -P i l te rs a nd D agg 1 98 1:92 L eese 1 927 :98 -99 L eona rd 1 894 :121 S a to 1 980 :26 T o r ry 1 973 :69 W i l l iamson a nd P ayne 1 959 :322 -323 Y as in a nd Wah id 1 957 :294

A Mode lo f Ado lescen t Ma le R e ten t ion R a tes T he mode l p resen ted i n T ab le 5 4 was g ene ra ted t o e s tab l ish t he n umbe r o f ma les e n te r ing a ge c l ass 6 -7 w h ich wou ld h ave t o b e r e ta ined a nnua l ly i no rde r t o ma in ta in t he r equ i red f i ve t os i x ma ture ma les . C ons is ten t w i th t he s ugges t ions o f L eese ( 1927 : 134 137 ) o n p urchas ing c ame ls f o r p ack w ork , i th as b een a ssumed t ha t b aggage c a r r ie rs a s we l l a s s t uds w ou ld o u t l ive t he i ru t i l i ty b yt h e e nd o ft he i rt wen t ie th y ea r a nd wou ld b e s ubsequen t ly c u l led . A nnua l s urv ivo rsh ip f o r ma les b e tween t he i r s even th a nd t wen t ie th y ea r a s p rev ious ly p resen ted i nT ab le 4 7 was 9 7% , t he e f fec ts o fw h ich a re mode l led i nt he p resen t s imu la t ion b y a ssum ing t ha t w ho le a n ima ls e n te r t he ma ture ma le p opu la t ion o ft he h e rd a nd s ubsequen t ly d ec l ine b y 3% e ach y ea r . S ince t he mode l h e rd w ou ld a l ready c on ta in t he r equ i red f i ve t o s i x a du l t ma les , t he p resen t s imu la t ion b eg ins w i th ma le s urv ivo rsh ip v a lues i n a l te rna te a ge c l asses wh ich c umu la t ive ly t o ta l 5 .7 a du l t ma les . l ti sa ssumed t ha te ach s ubsequen ty ea ri n

T ab le 5 4 Mode l o ft he R e ten t ion R a te f o r Ma le C ame ls E n te r ing T he i r7 th Y ea rw hen t he G oa l i s Ma in ta in ing 5t o 6Ma tu re Ma les

A ge C lass 6 -7 7 -8 8 -9 9 -10 1 0 -11 1-12 1 2 -13 1 3 -14 1 4 -15 1 5 -16 1 6 -17 1 7 -18 1 8 -19 1 9 -20 T o ta l

0

1

Y ea r 3

2

1 .00 . 97

1 .00 . 97

. 94

. 9 1

. 94 . 9 1 . 86

. 8 1

. 76

. 76

5 .70

. 69 . 67

5 .87

. 76 . 74

. 72

. 69 . 67

. 8 1 . 78

. 74

. 72

. 72

. 86 . 83

. 78

. 74

. 72 . 69

. 67

5 .70

5 .87

1 81

. 97 . 9 1

. 86

. 8 1

. 76

. 94 . 89

. 83

. 78 ,

6

1 .00 . 97

. 89

. 83 . 8 1

5

. 9 1

. 89 . 86

4

5 .70

. 67 5 .87

5 .70

w h ich t he c umu la t ive ma le p opu la t ion d rops b e low t h is 5 .7 t o ta l a s ar e su l t o f n a tura l m o r ta l i ty a nd c u l l ing , o ne ma le wou ld h ave t o b e r e ta ined f r om t he t o ta l a do lescen t ma le p opu la t ion e n te r ing a ge c l ass 6 -7 . T he 5 .7 a du l t ma les a t t he b eg inn ing o f y ea r 0o f t he s imu la t ion d ec l ine f r om n a tu ra l mor ta l i ty a nd c u l l ing t o 4 .87 a du l t ma les b y t he b eg inn ing o f y ea r 0 + 1, t hus r equ i r ing t he r e ten t ion o f 1a do lescen t ma le e n te r ing a ge c l ass 6 -7 . T h is r a ises t he a du l t ma le p opu la t ion t o 5 .87 a n ima ls a tt he b eg inn ing o fy ea r 1 .D u r ing t he c our se o fy ea r 1 ,t hese 5 .87 a du l t ma les d ec l ine f r om n a tura l mor ta l i ty , l e av ing 5 .7 a du l t ma les a tt he b eg inn ing o f y ea r 2 . S ince t h is n umbe r i st he m in imum r equ i red t o mee t b aggage a nd s t ud p u rposes , n o a do lescen t ma les e n te r ing a ge c l ass 6 -7 a tt he b eg inn ing o fy ea r 2n eed b e r e ta ined . H owever , d u r ing t he c ou rse o fy ea r 2 ,t hese 5 .7 a du l t ma les a ga in d ec l ine f r om n a tu ra l mor ta l i ty a nd c u l l ing , l e av ing 4 .87 a du l t ma les b yt he b eg inn ing o fy ea r3 .T hus , o ne a do lescen t ma le e n te r ing a ge c l ass 6 -7 a tt he b eg inn ing o f y ea r 3n eeds t o b e r e ta ined , a ga in r a is ing t he a du l t ma le p opu la t ion t o 5 .87 a n ima ls . I ti sa ppa ren t t ha t t he r e ten t ion o fo ne a do lescen t ma le e very o the r y ea r wou ld m a in ta in t he a du l t ma le p opu la t ion a tt he r equ i red f i ve t os i x p opu la t ion l e ve l . I na l te rna te y ea rs , t h e 0 . 67 ma les i na ge c l ass 1 9 -20 d ec l ine f r om n a tu ra l mo r ta l i ty t o 0 .65 ma les , p rov id ing a n a nnua l o f f take o f0 .325 e l de r ly ma les e n te r ing a ge c l ass 2 0 -21 . S i nce a n a dd i t iona l 0 . 16 t o0 . 17 a du l t ma les i no the r a ge c l asses a re s chedu led t o d i e o f n a tu ra l c auses i na l terna te y ea rs , t o ta l mor ta l i ty b y n a tu ra l c auses a nd h uman c u l l ing i nt he a du l t ma le h e rd a l terna tes b e tween 0 .49 a nd 0 .5 ma les a nnua l ly . T hus , t he a nnua l e xpec ted o f f take f r om a do lescen t ma les s u rv iv ing i n to t he i r s even th y ea r wou ld c ons is t o ft he t o ta l n umbe ro f ma les e n ter ing a ge c l ass 6 -7 m inus t he 0 . 5 i n d iv idua ls r equ i red a nnua l ly t o ma in ta in t he a du l t m a le p opu la t ion .

O f f take F rom T he Mode l C ame l H e rd O ve r aO ne Y ea r P e r iod N a tu ra l Mor ta l i ty a nd E xpec ted O f f take I na dd i t ion t ot he n umbe ro fa n ima ls s l augh te red d u r ing t he y ea ru nde r n orma l o r max imum c u l l ing s t ra teg ies , i ti sr easonab le t o e xpec t t ha t ap ropo r t ion o f t he t o ta l n umber o f a n ima ls s chedu led t o d i e o f n a tu ra l c auses e ach y ea r w ou ld a l so b e c onsumed . F o r e xamp le , among t he s heep a nd g oa t h e rd ing B akh t ia r i o fs ou thwes te rn I r an , R edd ing ( 1981:133 ) o bse rved t ha t s i ck a n ima ls a nd t hose b i t ten b y s nakes o r a t tacked b y w o lves w e re o f ten s l augh te red a nd c onsumed when i mpend ing d ea th was a ppa ren t . K a r imo jong c a t t le h e rde rs i nU ganda s im i la r ly s l augh te r a nd c onsume s i ck o r d y ing c a t t le ( Dyson -Hudson 1 960 :51 ; D yson -Hudson a nd D yson -Hudson 1 970 :110 ) . D ough ty ( 1936 :231) s pec i f ica l ly r eco rded t he c onsump t ion o f af ema le c ame l w h ich h ad d i ed i nc a lv ing , a nd n o ted t ha t i ng ene ra l , b edou in wou ld k i l l a nd c onsume a n ima ls l i ke ly t od i e ( 1936 :499 ) . P h i lby ( 1973 : 125 ) a l so r e co rded t he s l augh te r a nd c onsump t ion o f a n e xhaus ted c ame l . C o le ( 1975 :27 ) s im i la r ly n o ted t ha t t he A l Mu r rah w ou ld s l augh ter a nd c onsume s i ck c ame ls when d ea th was i mm inen t , a nd s uch was a l so t he c ase among b o th t he T ua reg ( N ico la isen 1 963 :63 ) a nd Gab ra ( To r ry 1 973 :115 -16 ) . A s n o ted b y L eese ( 1927 : 140 ) , wh i le I s lam ic L aw p roh ib i ts t he c onsump t ion o fa n ima ls wh ich h ave n o t b een r i t ua l ly s l augh te red , t h is p roh ib i t ion i sn o t g ene ra l ly c ons ide red r e levan t s o l o ng a st h e d y ing a n ima l 's t h roa t i sc u t b e fo re t he h ea r t s t ops b ea t ing . H oweve r , a cco rd ing t o Gab ra i n fo rman ts ( To r ry 1 973 :115 ) , l i t le mo re t han h a l f o f t he c ame ls w h ich d i e f r om n a tu ra l c auses a re u sua l ly c onsumed u n less t hey h ave d i ed i nm asses f r om e p idem ics . A5 0% u t i l iza t ion o fa l c ame ls s chedu led t od i e f r om n a tura l c auses t h e re fo re s eems a pp rop r ia te f o r n o rma l c ond i t ions . I ns ubsequen t c a lcu la t ions , i tw i l l b e a ssumed t ha t 5 0% o fa l a n ima ls s chedu led t od i ef r om n a tu ra l c auses wou ld a l so b e c onsumed i na dd i t ion t ot he a n ima ls s l augh te red u nde r n o rma l c u l l ing s t ra teg ies . T h is s l augh te r y i e ld w i l l b e t e rmed " no rma l o f f take . " H oweve r , i nc a lcu la t ing t he max imum s l augh te r p o ten t ia l o ft he mode l c ame l h erd , i tw i l l b e a ssu med t ha t 1 00% o fa l a n ima ls s chedu led t o d ie o f n a tura l c auses wou ld b e c onsumed i n a dd i t ion t o t he a n ima ls s l augh te red u nder a max imum c u l l ing s t ra tegy . T h is s l augh te r y i e ld w i l l b e t e rmed " max imum 1 82

o f f take . " Wh i le t h is l a t te r y i e ld i su n rea l is t ica l ly h i gh , i th as b een a dop ted h e re i no rder t o max im ize t he t o ta l p o ten t ia l mea t , o rgan , f a t a nd b l ood p roduc t iv i ty e xpec ted f r om t he mode l h e rd t o f ac i l i ta te t he l a te r c ompa ra t ive e ne rge t ic a na lys is o f mea t -on ly a nd mea t p l us m i lk p as to ra l s t ra teg ies .

N o rma lO f f take f r om t he Mode lC ame lH e rd N o rma l o f f take f r om t he mode l h e rd o ve r ao ne y ea rp e r iod wou ld c ons is t o f a5 0% u se o f a l a n ima ls s chedu led t od i e o fn a tu ra l c auses a nd t he c u l l ing o fe xcess a do lescen t ma les a nd e l de r ly ma les a nd f ema les . D ur ing t he c ou rse o ft he y ea r , t he f ou r ma les i na ge c l ass 5 -6 o ft he m ode l h e rd d ec l ine a s ar esu l t o f n a tura l mor ta l i ty , l e av ing 3 .88 ma les e n te r ing a ge c l ass 6 -7 . S ince 0 .5 ma les mus t b e r e ta ined a nnua l ly , 3 .38 ma les i na ge c l ass 6 -7 a re c u l led . A s p rev ious ly n o ted , a pp rox ima te ly 0 .33 ma les e n te r ing a ge c l ass 2 0 -2 1 wou ld a l so b e c u l led a nnua l ly . T hus , t he t o ta l n umbe r o f a do lescen t a nd e l de r ly ma les c u l led f r om t he mode l h erd a tt he e nd o f a y ea r wou ld amoun tt o3 .7 1 a n ima ls . T he t wo f ema les i na ge c l ass 2 1-22 o ft he mode l h e rd a l so d ec l ine f r om n a tura l mor ta l i ty d u r ing t he y ea r , l e av ing 1 . 92 f ema les e n te r ing a ge c l ass 2 2 -23 . H a l f o ft hese a re t hen c u l led , y i e ld ing 0 .96 f ema les . F ur the r , t he o ne f ema le i na ge c l ass 2 2 -23 a l so d ec l ines f r om n a tura l m o r ta l i ty , l eav ing 0 .94 t o b e c u l led a tt he e nd o f t he y ea r . T hus , t he t o ta l n umbe r o f e l de r ly f ema les c u l led f r om t he h e rd a tt he e nd o f ay ea r amoun ts t o1 . 9 a n ima ls . W i th a 5 0% u se o ft he t o ta l n umbe r o f a n ima ls s chedu led t o d ie f r om n a tura l c auses ( = 5 .04 a dd i t iona l a n ima ls ) , n o rma l o f f take f r om t he mode l h e rd wou ld amoun t t o a pp rox ima te ly 1 0 .65 a n ima ls o ve rt he c ou rse o fo ne y ea r . T h is o ff take i ss umma r ized i nT ab le 5 5 . T hese f i gures s ugges t t ha t n o rma l o f f take a t t he e nd o f o ne y ea r wou ld amoun t t o a pp rox ima te ly 7 .5 % o f t he 1 42 i n d iv idua ls i n t he mode l h e rd a t y ea r 0 + 1 d ay . T h is i s s ubs tan t ia l ly h i ghe r t han t he 3% a nnua l o f take f r om a c ame l h e rd a s c a lcu la ted b y D ah l a nd H jo r t ( 1976 :196 ) . H oweve r , w h i le t hese a u tho rs a ssumed t ha t a l ma les i na ge c l ass 5 -6 wou ld b e c u l led a nnua l ly , t hey d i d n o t i n c lude o f f take f r om c u l l ing o l de r a n ima ls n o rt he u se o f a ny a n ima ls s chedu led t od ie o fn a tu ra lc auses . F ur the r , n a tu ra l mor ta l i ty among ma les i na ge c l ass 5 -6 a nd t he n eed t o r e ta in a p o r t ion o f t he s u rv ivo rs i n o rde r t o ma in ta in t he a du l t ma le c omponen t o ft he h e rd we re a l so n o t c ons ide red . I fo f f take i nt he p resen t s t udy we re s im i la r ly c a lcu la ted , i twou ld amoun tt oa pp rox ima te ly 2 . 8% o ft he t o ta l h e rd , v e ry c l ose t ot he 3% o ff take c a lcu la ted b y D ah l a nd H jor t . S a to 's mode l o f c ame l h erd g row th ( 1980 ) d id n o t i n c lude c a lcu la t ions o fa nnua lo ff take .

T ab le 5 5 N o rma l O f f take f r om t he Mode l C ame l H e rd

AGE C LASSES 0 1 1 -4 4 -7 7 -23 TOTAL

NATURAL DEATHS #o fH e rd A n ima ls a t #o fD ea ths b y Y ea r0 + 1 d ay Y ea r0 +365 d ays Ma le F ema le Ma le F ema le 1 0 1 6 9 6 4 1

NORMAL OFFTAKE 5 0% U se o fa l D ead A n ima ls b y N o rma l C u l l ing Y ea r0 +365 d ays a tY ea r 1 + 1 d ay Ma le F ema le Ma le F ema le

1 2 3 1 8 4 9

3 .40 •1 . 16 0 .27 0 . 18

1 .98 1 .01 0 .54 1 .52

1 .70 0 .58 0 . 14 0 .09

0 .99 0 .51 0 .27 0 .76

3 .38 0 .33

1 .90

1 01

5 .01

5 .05

2 .51

2 .53

3 .71

1 .90

1 83

-

Max imum O f f take f r om t he Mode lH e rd Max imum o f f take f r om t he mode l h e rd o ve r ao ne y ea rp e r iod wou ld c ons is t o f a1 00% u se o f a l a n ima ls s chedu led t o d i e o f n a tura l c auses , t he c u l l ing o fe xcess a do lescen t ma les a nd e l de r ly ma les a nd f ema les , a nd t he c u l l ing o fa l a do lescen tf ema les i ne xcess o ft hose r equ i red t o ma in ta in t he h e rd a t ac ons tan t p opu la t ion s i ze . D ur ing t he c ourse o f t he y ea r , t he a du l t f ema le p opu la t ion i na ge c lasses 7 -23 o f t he mode l h e rd w ou ld d ec l ine b y 1 .52 t o ta l i n d iv idua ls a s ar e su l t o f n a tu ra l mor ta l i ty . I na dd i t ion , f ema les i na ge c l ass 6 -7 wou ld d ec l ine b y0 . 18 a s ar esu l to fn a tu ra l mo r ta l i ty . F i na l ly , 1 . 9 e l de r ly f ema les w ou ld b ec u l led a tt he e nd o ft he y ea r . T hus , t he f ema le p opu la t ion i na ge c l asses 6 -23 w ou ld d ec l ine b y 3 .6 i n d iv idua ls o ve rt he c ou rse o ft he y ea r . S ince 5 .82 f ema les i na ge c l ass 5 6w ou ld b e a va i lab le f o r s l augh te r a t t he e nd o f t he y ea r , t he r e ten t ion o f 3 .6 o f t hese t o ma in ta in t he a du l t f ema le p opu la t ion o ft he h e rd w ou ld a low f o rt he c u l l ing o f 2 .22 a do lescen t f ema les e n te r ing a ge c l ass 6 -7 . W i th t he a dd i t iona l u se o f 1 00% o f a l a n ima ls d y ing o fn a tu ra l c auses ( = 1 0 .6 a dd i t iona l a n ima ls ) a nd t he c u l l ing o fe xcess a do lescen t ma les a nd e l de r ly ma les a nd f ema les , max imum o f f take f r om t he mode l h e rd wou ld amoun tt oa pp rox ima te ly 1 7 .89 a n ima ls , o r a round 1 2 .7% o f t he o r ig ina l 1 42 a n ima ls i nt he mode l h e rd a ty ea r0 + 1 d ay . T ab le 5 6 s u mma r izes t he s l augh te r y i e ld u nde r max imum o f f take . W i th t he d a ta p resen ted i nt h is a ppend ix , i tw i l l b e p oss ib le t oc a lcu la te t he e xpec ted c a lo r ic r e tu rns f r om t he mode lc ame l h e rd o ve r ao ne y ea rp e r iod u nde rn o rma l a nd max imum o f f take s t ra teg ies . S uch c a lcu la t ions a re made i nA ppend ix B .

T ab le 5 6 Max imum O f f take f r om t he Mode l C ame l H e rd

1 00% U se o fa l D ead A n ima ls b y Y ea r0 +365 d ays Ma le F ema le

Max imum C u l l ing a tY ea r 1 + 1 d ay Ma le F ema le

0 1 1 -4 4 -7 7 -23

3 .40 1 . 16 0 .27 0 . 18

1 .98 1 .01 0 .54 1 .52

3 .38 0 .33

2 .22 1 .90

T o ta l

5 .01

5 .05

3 .71

4 .12

A ge C lasses

1 84

APPEND IX B

CALOR IC P RODUCT IV ITY OF T HE MODEL CAMEL HERD OVER A ONE Y EAR P ER IOD

I n t roduc t ion T he t o ta l e ne rgy p roduc t ion o ft he mode l c ame l h e rd g ene ra ted i nA ppend ix A o ve r ao ne y ea r p e r iod wou ld c ons is t o ft he c a lo r ies d e r ived f r om b o th o f f take a nd m i lk p roduc t ion . I nt h is a ppend ix , t he c a lo r ic y i e ld d e r ived f r om n o rma l a nd max imum o f f take i sd e te rm ined ,f o l lowed b y s im i la rc a lcu la t ions f o rl ow , n o rma l a nd h i gh l e ve ls o fm i lk p roduc t iv i ty . I no rde rt oc a lcu la te t h e t o ta l m ea t , o rgan , f a t a nd b l ood y i e ld f r om t he mode l c ame l h erd u nde rn o rma l a nd max imum o f f take a sp resen ted i nA ppend ix A , i ti sn ecessa ry t oe s tab l ish t he q uan t i t ies o fe ach p roduc t e xpec ted p e r i n d iv idua l c ame l i ne ach a ge a nd s ex c l ass . S ince t he q uan t i t ies o fs uch i t ems o b ta ined p e ri n d iv idua lw ou ld p resumab ly v a ry a cco rd ing t oi t s r eg iona l b r eed , p r io r p l ane o fn u t r i t ion , a ge , s ex , a nd s eason o fd ea th o rs l augh te r , i tw ou ld h ave b een p re fe rab le t o e s tab l ish mean y i e lds a cco rd ing t oa ge a nd s ex s pec i f ic d a ta d er ived f r om t he o bserva t ion o ft empo ra l ly a nd g eog raph ica l ly d i s t inc t c ame l h e rds ma in ta ined u nde rc ond i t ions o f e x tens ive h usband ry . H oweve r , t he a va i lab le d a ta a re s o s pa rse a s t o p rec lude s uch p r ec is ion . I n i t ia l ly , r easonab le e s t ima tes o fa ge a nd s ex s pec i f ic mea t , o rgan ,f a ta nd b l ood y i e lds p e r a n ima la re d e r ived f r om t he a va i lab le q uan t i ta t ive d a ta .A ne s t ima t ion o ft he t o ta l m ea t , o rgan ,f a t a nd b l ood y i e ld e xpec ted f r om t he mode lc ame l h e rd i st hen c a lcu la ted f o rn o rma la nd max imum o f f take s t ra teg ies . T he c a lo r ic v a lues o fi n d iv idua l e d ib le i t ems a re s ubsequen t ly e s tab l ished , a nd t he t o ta lc a lo r ic y i e ld f o rn o rma la nd max imum o f f take s t ra teg ies i sc a lcu la ted . I no rde rt oe s tab l ish t h ey i e ld o fm i lk e xpec ted f r om t he mode l c ame l h e rd o ve r ao ne y ea r p e r iod , i ti sn ecessa ry t o i n i t ia l ly d e te rm ine t he q uan t i ty o fm i lk e xpec ted p e r l a c ta t ing f ema le . S i nce t he q uan t i ty a nd c ompos i t ion o f m i lk c ou ld p o ten t ia l ly v a ry b y r t e g iona l b reed , g raz ing c ond i t ions , a nd l e ng th o ft i me b e tween p a r tu r i t ion a nd s amp l ing , t he c a lcu la t ion o ft o ta l m i lk y i e ld f r om t he mode l h e rd s hou ld b e b ased u pon r esea rch s pec i f ica l ly a dd ress ing s uch v a r ia t ions w i th in t empo ra l ly a nd g eog raph ica l ly d i sc re te c ame l h e rds ma in ta ined u nder c ond i t ions o f e x tens ive h usband ry . H oweve r , t he a va i lab le d a ta a re a ga in s o s pa rse a s t o p r ec lude s uch p rec is ion , a l though r easonab le e s t ima tes o fe xpec ted y i e lds p e r i n d iv idua l c an b e e s tab l ished . U s ing s uch e s t ima tes , c a lcu la t ions a re made f o rv a r ious m i lk y i e lds f r om t he mode l h e rd o ve r ao ne y ea r p e r iod i no rde rt of a c i l i ta te p roduc t iv i ty c ompa r isons b e tween mea t -on ly a nd m ea t p l us m i lk p as to ra l s t ra teg ies . A f te r e s tab l ish ing t he c a lo r ic c on ten t o f c ame l 's m i lk , t he c a lo r ic y i e lds f r om l ow , n o rma la nd h i gh l e ve ls o fm i lk p roduc t iv i ty a re c a lcu la ted f o rc ompa r isons w i th t h ec a lo r ic y i e lds o b ta ined u nde rn o rma l a nd max imum o f f take .

T o E a t aC ame l T he t o ta l amoun t o f mea t , o rgans , f a t a nd b l ood o b ta ined f r om ac ame l a re n o t e qu iva len t t ot h e a n ima l 's l i vewe igh t , s i nce s uch i t ems a s b one , s k in , s t omach c on ten ts , a nd i n tes t ines a re n o t n o rma l ly c onsumed ( Mus i l 1 928 :96 ) . P ar t icu la r ly i n s t ruc t ive o n t he p repa ra t ion a nd c onsump t ion o f s l augh te red a n ima ls i st he a ccoun t o fJ enn ings -B ram ley a s b ased u pon h i s o bse rva t ions among t he b edou in o fS ina i . A l l k i nds o f mea t a re g ene ra l ly b o i led , t he w ho le o ft he f a t o ft he a n ima l b e ing p u t i n to t he p o t a s w e l l . T he e n t ra i ls , t he l i ve r a nd t he h ea r t a re u sua l ly

s en t o n b e fo re , t ot he menz i l , t ob e e a ten w h i le t he r e s t i ss t i l l c ook ing , t hese o dds a nd e nds r equ i r ing l e ss t ime , a nd b e ing p repa red s imp ly e nough , v i z . , b y b e ing t h rown o nt ot he h o tc i nde rs a nd a lowed t ob urn f o r af ew m inu tes . T he c h i ld ren w ho c o l lec t i nn umbe rs r ound t he menz i l ,w hen af eas t i sg o ing o n , a r e t h rown mos t o ft hese b i ts w h ich a re n o t c ons ide red a sf o rm ing p a r t o ft he r e a l mea l , b u tt a ke t he p l ace o fh ors d ' oeuv res [ J enn ings -B ram ley 1 907 :13 11321 T he f a t ty h ump o ft he c ame l i sa l so c onsumed . C hunks o fh ump t i ssue a re s ome t imes r oas ted o rf r ied , a l though i ng ene ra l , t he e n t i re h ump i sb o i led d own f o rg rease t ob e u sed a s b u t te ro r s hor ten ing ( D ickson 1 951 :190 , 4 16 ; D ough ty 1 936 :231; J enn ings -B ram ley 1 907b : 13 1 ; Mus i l 1 928 :96 ) . I ti su nc lea r , h oweve r , w he the r b one ma r row i sa l so c onsumed b y c ame l n omads . M us i l ( 1928 :70 ) men t ioned t he u se o f ma r row i nt he p repa ra t ion o f h i des , b u t n o r e ferences h ave b een n o ted w h ich d ocumen t i t s a c tua l c onsump t ion . T he c onsump t ion o fb l ood f r om c ame ls i s a l so p oo r ly d ocumen ted . A cco rd ing t o Mus i l ( 1928 :96 ) , t h e b l ood o fs l augh te red c ame ls was e i the rc o l lec ted b yt he Rwa la i n av esse l o rs imp ly a lowed t od ra in u pon t he g round , a l though h e a l so s t a ted t ha t Rwa la w ou ld c onsume t he b l ood o fs l augh te red a n ima ls ( Mus i l 1 928 :97 ) . P resumab ly , t he t o ta l c on t r ibu t ion o f b l ood t o t he Rwa la d i e t i sm in ima l . Wh i le t h e T ua reg o ccas iona l ly b l eed c ame ls t od r ink , t hey d o n o t c ons ider b l ood t ob e ap rope rd i sh ( N ico la isen 1 963 :122 -23 , 2 33 -35 ) . Whe the r t he T ua reg c onsume t he b l ood o f s l augh te red c ame ls i s u ndocumen ted . I nE as tA f r ica , t he Gab ra ( Tony 1 973 :77 -78 , 1 974 :65 ) , T urkana ( Dah l a nd H jo r t 1 976 :193 -94 ; C oughenou r e t a l . 1 985 :620 ) , a nd R end i l le ( Sa to 1 980 :8 -9 ) b l eed c ame ls t o d r ink , a nd wou ld p resumab ly c onsume t he b l ood o f s l augh te red c ame ls . P reh is to r ic b l ood c onsump t ion f r om s l augh te red c ame ls w i l lt he re fo re b ea ssumed . I nc a lcu la t ing t he t o ta lc a lo r ic y i e ld f r om s l augh te red c ame ls , i tw i l lb e a ssumed t h a tc a rcass mea t , mea t s c raps , b l ood , t ongue , l i ve r , h ear t , a nd k i dneys wou ld b e c onsumed , a l ong w i th h ump , r ena l , a nd r ende red f a t . C a lo r ic y i e lds f r om p e r iod ica l ly b l eed ing l i ve c ame ls o rf r om s uch i t ems a s ma r row , b ra ins a nd l u ngs w i l l n o t b e i n c luded . T he a dd i t iona l c a lo r ic c on t r ibu t ion o f s uch i t ems i se xpec ted t ob e r e la t ive ly i n s ign i f ican t b yc ompa r ison w i th t hose i t ems w h ich c an b e a ccoun ted f o r .

M ea t , O rgan , F a t a nd B lood Y ie ld F rom S laugh te red C ame ls Me thodo logy Ac onven ien t me thod f o rc a lcu la t ing t he e xpec ted y i e ld o fc a rcass mea t , o rgans , a nd f a t f r om i n d iv idua la n ima ls i st oe s tab l ish mean p ropo r t iona lv a lues f o re ach i t em a s ap e rcen tage o f mean l i vewe igh t . Wh i le t h is a ssumes i s ome t r ic a s o pposed t o a lome t r ic s ca l ing ( Mar t in 1 984 :87 -88 ) , i ti st he b es tt ha tc an b e a ch ieved g i ven t he a va i lab le d a ta o nc ame l s l augh te r .

S laugh te ra nd P ropo r t iona l Y i e ld D a ta Two q uan t i ta t ive s t ud ies o n t he s l augh te r a nd p ropo r t iona l y i e lds o f e d ib le i t ems f r om d romeda ry c ame ls h ave b een l o ca ted . O ne i sa n I t a l ian s t udy c onduc ted a ta na ba t to i ri nc en t ra l S oma l ia w h ich i n vo lved t he s l augh te ro f1 50 a du l tc ame ls o ft h e l a rge B enad i ra nd B asso G iuba b reeds ( Cong iu 1 953 ) . T he r e su l ts o ft h is s t udy a re p resen ted i nT ab le 5 7 . Af a r m o re d e ta i led s t udy w as c onduc ted o ve r at h ree y ea r p e r iod i n S ov ie t T u rkmen is tan , a nd i n vo lved t he s l augh te r o f 1 28 ma le T urkmen ian d romeda r ies o ft he A rwan b reed ( Kuzne tzov e ta l . 1 970 ) . T he i rr e su l ts f r om t he s l augh te ro f7 4 ma le c ame ls ma in ta ined o n " ave rage " p l anes o fn u t r i t ion s eem mos t r e levan tt ot he p resen ts t udy , a nd t hese a re p r esen ted i nT ab le 5 8 . O n ac ompa ra t ive b as is , i ti sa ppa ren tt ha tt he mean l i vewe igh to fa du l t ma le B enad i ra nd B asso G iuba b reeds ( 554 k g ) i sh i ghe r t han t ha t o ft h e A rwan b reed ( 468 k g ) . F u r the r , t he s l augh te ro fB enad i ra nd B asso G iuba b reeds r e su l ted i nm o re f avo rab le d ress ing r a tes ( c a rcass w e igh t / l ivewe igh t ) t han among t he A rwan b reed ( 56% v s 5 0 .5 % r e spec t ive ly ) . I nt h e p resen t s t udy , amean d ress ing r a te o f5 3% o fl i vewe igh tw i l lb ea ssumed a sa pp rop r ia te f o ra lc ame ls .

1 86

T ab le 5 7 S laugh te ra nd P ropo r t iona lY ie ld D a ta o n 1 50 A du l tS oma l i D romeda r ies

S EX

MALE

F EMALE

SAMPLE S IZE MEAN L IVE WE IGHT

5 0 5 54 k g

1 00 5 14 k g

I TEM c a rcass e n t i re h ump l i ve r h ea r t k i dneys e n t i re h ead l u ngs h ide

k g

%o f l i vewe igh t

k g

% o f l i vewe igh t

3 09 .00 1 5 .25 1 2 .00 2 .34 2 .56

5 5 .78 2 .75 2 . 16 0 .42 0 .46

2 86 .00 2 0 .21 1.80 2 . 17 2 .88

5 5 .64 3 .93 2 .30 0 .42 0 .56

1 5 .00 4 .45 5 8 .25

2 .71 0 .80 1 0 .51

1 2 .94 4 .42 4 8 .28

2 .52 0 .86 9 .39

c a lcu la ted f r om d a ta g i ven i nC ong iu ( 1 953 ) H oweve r , b e fo re e s tab l ish ing t he p ropor t iona l y i e ld o ft o ta l mea t e xpec ted f r om i n d iv idua ls i nt h e mode l h e rd o n t he b as is o ft hese s t ud ies , i ti sn ecessa ry t od e te rm ine w ha t t he r a t io o f m ea t t o b one m igh t b e f o r c ame l c a rcasses . F ur the r , i ti sn ecessa ry t o b r ie f ly d i scuss t he p o ten t ia lf o rs easona la nd s ex v a r ia t ions i nc ame l h ump f a ty i e lds .

B on ing R a te C a rcass w e igh ts a re n o t e qu iva len t t o mea t y i e lds , s i nce t he c a rcass c on ta ins b one . U n for tuna te ly , n o d a ta o n b on ing r a tes f o r c ame l c a rcasses h ave b een f ound . H owever , a cco rd ing t o K uzne tzov e t a l . ( 1970 :125 ) , t he r a t io o f mea t t o b one i nc ame l c a rcasses i s g ene ra l ly mo re f a vo rab le t han i nc a t t le c a rcasses . S ince p ub l ished p e rcen tages o fb one i nt he c a rcasses o f u n imp roved b reeds o fc a t t le i nE as t A f r ica a re g i ven a t 1 0% , 1 2% , a nd 1 4 .8 % ( Dah l a nd H jo r t 1 976 :169 ) , a1 0% b one c on ten t f o rc ame l c a rcasses s eems g ene rous , a nd w i l l emp loyed i ns ubsequen t c a lcu la t ions . G iven t he 5 3% mean d ress ing r a te e s tab l ished e a r l ie r , a1 0% b one c on ten t f o r t he r e su l t ing c a rcass w ou ld i mp ly t ha t mea t y i e ld f r om ac ame l c a rcass s hou ld r ep resen t4 7 .7% o f l i vewe igh t . F ur the r , t he d a ta o fK uzne tzov e ta l . ( 1970 ) o n s upp lemen ta ry mea t o b ta ined f r om b u tche r ing s c raps a nd s c rap ing t he h i de i n d ica te a dd i t iona l mea t y i e lds o f a round 0 .7% o f l i vewe igh t . T hus , t he t o ta l y i e ld o f mea t e xpec ted f r om t he s l augh te ro f ac ame l s hou ld amoun t t or o ugh ly 4 8 .4% o fl i vewe igh t . O ve ra l l , t he a ssump t ion t ha t 4 8 .4% o fl i vewe igh t w ou ld r ep resen t t he u se fu l y i e ld o f mea t f r om c ame ls s eems q u i te l i be ra l . F o r c ompa r ison , N ewman ( 1975 :97 ) s ugges ted t ha t a pp rox ima te ly o ne th i rd o ft h e l i v ewe igh to ff e ra lc ame ls i nA us t ra l ia c ou ld b e r ega rded a su se fu l m ea tf o rt he d omes t ic t ab le a nd manu fac tur ing mea t ma rke ts . E ven w h i le a cknow ledg ing t ha t p as to ra l p opu la t ions w ou ld p r esumab ly b e l e ss d i sc r im ina t ing i nt he i r s e lec t ion o f mea ts t han c on tempo ra ry A us t ra l ians , am ea t y i e ld o f 4 8 .4% o f l i vewe igh t s eems e x t reme ly g ene rous i n c ompa r ison w i th amea ty i e ld o fo n ly 3 3% . V a r ia t ions i n Hump F a tY i e ld Wh i le t he s easona l s t o rage o f f a t among t r ue u ngu la tes i sp ar t ia l ly a ch ieved t h rough s ubcu taneous f a t d epos i t ion o ve r a n e x tended a rea o f t he b ody , s uch s t o rage i nc ame ls i s 1 87

T ab le 5 8 S laugh te ra nd P ropo r t iona l Y ie ld D a ta o n 7 4 Ma le T u rkmen ian D romeda r ies Ma in ta ined o n a n A ve rage P lane o fN u t r i t ion

AGE GROUPS

2-3YEARS

4YEARS AND O LDER

SAMPLE S IZE MEAN L IVE WE IGHT

5 2 3 69 k g

2 2 4 68 k g

-

-

UN ITS OF MEASURE

Mea t

E d ib le I t ems

F a t

c a rcass f r om s c raps f r om h i de

- -

0 / 0o f l i vewe igh t

k g

k g

0 / 0o f l i vewe igh t

1 85 .00 1 .40 1 .40

5 0 .10 0 .38 0 .38

2 38 .00 1 .50 1 .40

5 0 .90 0 .32 0 .30

t ongue l i ve r h ea r t k i dneys

0 .52 5 .80 1 .81 1 .45

0 . 15 1 .57 0 .50 0 .40

0 .73 6 .20 2 .20 1 .86

0 . 15 1 . 32 0 .47 0 .40

h ump r ena l r endered

3 .30 1 .35 1 .90

0 .90 0 .38 0 .51

5 .70 1 .90 2 .80

1 . 22 0 .41 0 .60

1.42 6 7 .20 1 7 .90 2 .80 5 .90 5 .40 2 7 .60 7 .77 1 3 .00 6 .03

3 . 10 1 8 .20 4 .85 0 .80 1 .60 1 .48 7 .50 2 . 11 3 .50 1 .64

1 4 .20 7 8 .60 2 4 .20 3 .76 7 .40 6 . 15 3 6 .00 9 .79 1 6 .40 9 .51

3 .03 1 6 .80 5 . 17 0 .80 1 . 56 1 .30 7 .70 2 .09 3 .50 2 .02

3 69 .00

1 00 .10

4 68 .30

1 00 .10

E n t i re h ead S tomach w i th c on ten ts I n tes t ines w i th c on ten ts L ungs F ee t L owe r l e gs H ide S p leen , t r achea , e t c . B lood ( es t ima ted ) Was te TOTALS 1

1N o t e xac t n umbe rs d ue t o r ound ing a dap ted f r om K uzne tzov e ta l . ( 1970 )

l o ca l ized i nt he i r we l l -known h umps ( Gau th ie r -P i l te rs a nd D agg 1 98 1:7 1; L eese 1 927 :13 -14 ; P ond 1 984 :20 -22 ) . T h is p ecu l ia rt r a i t i nc ame ls a ppa ren t ly r ep resen ts o ne a spec to ft h ec ame l 's e labo ra te a dap ta t ion t o p rob lems o f h ea t r egu la t ion a nd wa ter c onserva t ion i n d eser t e nv i ronmen ts ( Gau th ie r -P i l te rs a nd D agg 1 981 :71 ; S chm id t -N ie lsen 1 959 :15 1, 1 979 :48 -49 ) . A s d i scussed i nC hap te r5 ,t he c ame l h as a n a ccep tab le b ody t empe ra tu re r ange o fa round 3 5 t o 4 1° C , a nd c an t hus i n te rna l ly s t o re h ea t g a ined f r om t he e nv i ronmen t d ur ing t he d ay t o b e d i ss ipa ted i n t he c oo l o f n i gh ts t h rough c onduc t ion a nd r ad ia t ion w i thou t n ecessa r i ly e xpend ing wa te r i nt he p rocess . B y s t o r ing f a t i nt he h ump r a the rt han s ubcu taneous ly o ve r a n e x tens ive a rea o f t he b ody , h ea t s t o red i n te rna l ly d u r ing t he d ay c an f l ow u n impeded t o t he s k in 's s ur face t ob e d i ss ipa ted a tn i üh t ( Schm id t -N ie lsen 1 979 :42 -45 ) . S ince t he amoun t o f h ump fa t s t o red b y c ame ls wou ld v a ry a cco rd ing t o t he q uan t i ty a nd q ua l i ty o f p as tu rage a va i lab le t o t hem , t he y i e lds o ff a t f o r h uman c onsu mp t ion s hou ld v ary a cco rd ing t ot he s eason o f s l augh te r . E a r ly e thnog raphe r s i nN o r th A rab ia , f o r e xamp le , o f ten n o ted t ha t c ame ls g rew " f a t a nd s t rong " a nd e xh ib i ted " en la rged h umps " d u r ing w in te r a nd e a r ly s p r ing when v ege ta t ion was p l en t i fu l , b u t " l angu ished " a nd " became l ean " d u r ing s umme r a nd f a l l when v ege ta t ion d ec reased ( D ickson 1 951:257 , 4 14 ; D ough ty 1 936 :259 -260 , 3 23 ;

1 88

J enn ings -B ram ley 1 906 :256 -257 ; Mus i l 1 928 :155 , 3 13 , 6 00 ) . U n for tuna te ly , q uan t i ta t ive d a ta o n s easona l v a r ia t ions i nc ame l h ump fa t s t o rage h ave n o t b een f ound . A cco rd ing t o D ough ty ( 1936 :23 1) , t he h ump o f af a t f ema le c ame l p u rchased d ur ing t he w in ter a t K ha iba r , S a 'ud i A rab ia , we ighed " a lmos t a n h und red w e igh t , " wh ich wou ld b e s omewha t l e ss t han 4 6 t o 5 1 k g . I n C ong iu 's s t udy i nS oma l ia ( 1953 :189 -190 ) , t he mean w e igh t o f e n t i re h umps f r om a du l t B enad i r a nd B asso G iuba d romeda r ies was 1 5 .25 k g f o r ma les a nd 2 0 .2 1 k g f o rf ema les , r ep resen t ing 2 .75 a nd 3 . 93 % o f l i vewe igh t r espec t ive ly . Mann ( 1 954 :493 -494 ) s t a ted t ha tt he " y ie ld o ff a t " p e rc ame l s l augh tered a ta n a ba t to i r i nS oma l ia w as a pp rox ima te ly 1 6 t o 2 7 k g , a l though t h is a ppa ren t ly r ep resen ts e n t i re h ump we igh ts i nl i gh t o f C ong iu 's d a ta . H oweve r , i twou ld s eem t ha t c ame l h umps w ou ld g ene ra l ly we igh s omewha t l e ss t han s uch f i gures s ugges t , e ven u nde r c ond i t ions o f o p t ima l p as turage . F o r e xamp le , G au th ie r -P i l te rs a nd D agg ( 198 1:7 1) s ugges ted t ha t w i th g ood f o rage , ad romeda ry 's h ump w ou ld n o rma l ly we igh b e tween 1 0 a nd 1 5 k g . F ur the r , H i l le l ( 1982 :59 ) s ugges ted t ha t aw e l l f ed a nd we l l -wa te red c ame lw ou ld h ave ah ump w e igh ing a bou t 1 0 k g . I nt he a bsence o f e xp l ic i t b reed , a ge , s ex , a nd s easona l i ty d a ta o n t he we igh ts o fc ame l h umps , C ong iu 's r esu l ts f r om S oma l ia w i l l b e a dop ted a s a n o p t im is t ic a pp rox ima t ion o f e xpec ted h ump y i e lds . H oweve r , a s w i th c a rcass we igh ts , h ump we igh ts a re n o t e qu iva len t t o f a t y i e lds , s i nce t he h ump i sb ound t oge the r b y f i b rous t i ssue a nd w a te r . S ince t he h ump i s g enera l ly r educed t o f a t , o n ly a p ropor t ion o ft he t o ta l h ump w e igh t i se qu iva len t t o a c tua l f a t y i e ld . I na n a na lys is o fc ame l h umps i nE gyp t , N as r e ta l . ( 1965 :255 ) f ound t ha t a pp rox ima te ly 4 2 t o 5 2% o ft o ta l h ump w e igh t c ons is ted o ff a t . T he r esu l ts o ft he i r a na lys is a re p resen ted i n T ab le 5 9 . A s a pp l ied t o C ong iu 's d a ta o n t he mean we igh ts o f ma le a nd f ema le c ame l h umps , a nd a ssum ing o n ly m in ima l v a r ia t ions i nt he h ump we igh t t o l i vewe igh t r a t io a s ar esu l t o fa ge ( see K uzne tzov e ta l . 1 970 :124 ) , i tw i l l b e e xpec ted t ha t h ump fa t y i e lds i ny ounge r f ema les wou ld amoun t t o 1 .63 % o f l i vewe igh t , wh i le t he h ump fa t y i e lds f o r o lde r f ema les wou ld amoun t t o 2 . 03 % o f l i vewe igh t . S im i la r ly , h ump fa t y i e lds i ny ounge r ma les wou ld amoun t t o 1 . 2% o f l i vewe igh t , wh i le t he h ump fa t y i e ld f o r o lde r ma les s hou ld amoun t t o 1 . 43 % o f l i vewe igh t . F o r c ompa ra t ive p u rposes , K uzne tzov e t a l . ( 1970 ) f ound t ha t t he h ump fa t y i e ld f o r a du l t ma le c ame ls amoun ted t o 1 . 22% o fl i vewe igh t , wh i le f o ry ounge r ma les t he y i e ld was 0 .9 % . A ga in , i t i ss uspec ted t ha t t he p ropo r t iona l r ep resen ta t ions f o r h ump fa t y i e lds a dop ted h e re a re q u i te g ene rous . I na dd i t ion t of a t r educed f r om t he h ump , sma l l q uan t i t ies o ff a t c ou ld a l so b e d e r ived f r om n eph r ic d epos i ts a nd f r om r ende r ing . U s ing mean v a lues f r om t he s l augh te r r esu l ts o f K uzne tzov e t a l . ( 1970 ) , a dd i t iona l f a t y i e lds amoun t ing t o 0 .95% o f l i vewe igh t a re e xpec ted p e ra n ima l .

P ropor t iona l Y i e lds o fE d ib le I t ems P e rC ame l T he e xpec ted y i e lds o f e d ib le i t ems f r om s l augh te red c ame ls a s d e r ived f r om t he d a ta p resen ted a bove a re s umma r ized i n T ab le 6 0 , whe re y i e lds p e r i t em a re e xp ressed a s a p e rcen tage o f l i vewe igh t . T he v a lues f o r mea t a nd f a t y i e lds a re t hose d e r ived d u r ing t he p reced ing a na lys is . F o r t ongue a nd b l ood y i e lds , t he o n ly a va i lab le v a lues a re t hose o f K uzne tzov e t a l . ( 1970 ) . F ur ther , t he h i ghe r h ea r t y i e ld v a lues o f K uzne tzov e ta l . ( 1970 ) h ave a l so b een a dop ted . H oweve r , C ong iu 's ( 1953 ) h i ghe rv a lues f o r l i ve r a nd k i dney y i e lds among a du l t d romeda r ies h ave b een p re fe ren t ia l ly a dop ted o ve rt he l owe r v a lues g i ven b y K uzne tzov e ta l . ( 1970 ) , a nd a re e qua l ly a pp l ied t oy ounge ra n ima ls o nt he a ssump t ion o fm in ima lv a r ia t ions i np ropo r t iona l y i e ld a s ar e su l t o fa ge . On t he b as is o ft h is d a ta , e d ib le i t ems a s d e f ined i nt h is s t udy wou ld a ccoun t f o r r ough ly 5 8% o ft he l i vewe igh t o fd romeda ry c ame ls .

C ame l L i vewe igh ts A ge a nd s ex s pec i f ic l i vewe igh t d a ta f o r c ame ls a re e x t reme ly s pa rse . K noess ( 1977 :42 ) n o ted t ha t i np l ow ing e xpe r imen ts i nE th iop ia , t he w e igh t o f as even y ea r o ld A fa r d romedary w as 3 7 1 k g , wh i le an i ne y ea ro l d o ft he s ame b reed we ighed 4 30 k g . H oweve r , A fa rc ame ls a re g ene ra l ly c ons ide red sma l l b y c ompa r ison w i th c ame ls f r om o the r r e g ions ( Dah l a nd H jo r t 1 89

T ab le 5 9 S tanda rd A na lys is o f4 0 C ame l H umps

AGE OF CAMEL

SEX

% P ROTE IN

% WATER

% F AT

` ) / 0 ASH

5y ea rs o ro l de r

ma le f ema le

9 .40 8 .50

4 7 .12 4 9 .06

4 3 .40 4 1.47

0 .43 0 .965

l e ss t han 5y ea rs

ma le f ema le

9 .67 9 .62

3 8 .11 3 8 .36

5 1.85 5 1.64

0 .33 0 .43

a dap ted f r om N as re ta l . ( 1965 )

T ab le 6 0 E xpec ted Y ie lds o fE d ib le I t ems P e rC ame l a s aP e rcen tage o f L i vewe igh t

I TEMS

0 -4 YEARS Ma le F ema le

4OR MORE YEARS Ma le F ema le

Mea t T ongue L i ve r H ea r t K idneys F a t B lood

4 8 .40 0 . 15 2 . 16 0 .50 0 .46 2 .38 3 .50

4 8 .40 0 . 15 2 .30 0 .50 0 .56 2 .98 3 .50

4 8 .40 0 . 15 2 . 16 0 .50 0 .46 2 . 15 3 .50

4 8 .40 0 . 15 2 .30 0 .50 0 .56 2 .58 3 .50

TOTAL

5 7 .55

5 8 .39

5 7 .32

5 7 .99

1 976 :190 ) . F o r e xamp le , N ewman ( 1975 :97 ) n o ted t ha t t he a ve rage l i vewe igh t o ft wo t of our y ea r o ld f e ra l d romeda r ies i nA us t ra l ia r anged f r om 4 50 t o6 50 k g . Mo re r e l iab le d a ta o n t he l i vewe igh ts o f ma le c ame ls a re c on ta ined i nt he s t udy b y K uzne tzov e ta l . ( 1970 ) , a s p r ev ious ly p resen ted . O f 7 4 ma le T u rkmen ian c ame ls ma in ta ined o n a n " ave rage " p lane o f n u t r i t ion , t he mean l i vewe igh t o ft wo t ot h ree y ea r o l ds was 3 69 k g , wh i le t he mean o ft hose f ou ry ea rs a nd o lde r was 4 68 k g . I nC ong iu 's s t udy ( 1953 ) o f t he r e la t ive ly l a rge B enad ir a nd B asso G iuba b reeds i nS oma l ia ,t he mean l i vewe igh to f1 50 a du l t ma le a nd f ema le c ame ls w i th am ean a ge o f 1 0 t o 1 2 y ea rs w e re f ound t ob e 5 54 k g a nd 5 14 k g r espec t ive ly . T he o n ly o the r a ge s pec i f ic l i vewe igh t d a ta f o r c ame ls c ons is t o f r eco rded b i r thwe igh ts . F o r e xamp le , t he mean b i r thwe igh t o f ma le c ame ls a tB ikane r , I n d ia , was 3 8 k g , wh i le R uss ian s t ud ies s ugges t 3 5 t o 4 0 k g a s t he mean b i r thwe igh t f o r a l n ew ly -bo rn c ame ls ( s ee D ah l a nd H jor t 1 976 :191) . F i na l ly , Wa l lach a nd B oeve r ( 1983 :326 ) s ugges t t ha t 3 0 t o4 5 k g i st h e n orma l r ange o fb i r th w e igh ts f o rd romeda r ies . H oweve r , wh i le a ge a nd s ex s pec i f ic l i vewe igh t d a ta f o r a du l t c ame ls a re q u i te s pa rse , s eve ra l a u tho rs h ave r eco rded a ve rage w e igh ts o r we igh t r anges . T h is a dd i t iona l l i vewe igh t d a ta f o r d romeda r ies a re s umma r ized i nT ab le 6 1 , wh i le t ha t f o r B ac t r ians a re s umma r ized i n T ab le 6 2 f o rc ompa ra t ive p u rposes . I ng ene ra l , i tw ou ld s eem t ha t B ac t r ian c ame ls w e igh more t han d romeda r ies , a nd t ha t ma les i nb o th s pec ies a re l a rge r t han f ema les . H oweve r , i nt he a bsence o f mo re s pec i f ic d a ta , p a r t icu la r ly o n t he f eed ing c ond i t ions a nd b reeds o ft h e c ame ls i n vo lved , i ti sg enera l ly n o t p oss ib le t o e va lua te t he a pp l icab i l i ty o f t hese d a ta t o c ame ls ma in ta ined u nde rc ond i t ions o fe x tens ive h usband ry .

1 90

T ab le 6 1 A ve rage L i vewe igh ts R epor ted f o r Ma tu re D romeda r ies

KG

S EX

4 00 -700 3 00 -500 5 00 -550 4 54 -558 5 32 -558 4 55 -523 3 50 -400 4 50 -650 4 50 -700

-

5 50 5 00 5 00 4 50 5 00

M F M -

LOCAT ION

BREED

N egev S aha ra S oma l ia I n d ia

B enad i r f e ra l -

N . S oma l ia A us t ra l ia S oma l ia S oma l ia S udan -

SOURCE B odenhe ime r 1 935 :125 Gau th ie r -P i l te rs a nd D agg 1 981 :38 Gau th ie r -P i l te rs a nd D agg 1 981:7 L eese 1 927 :14 L eese 1 927 :14 L eona rd 1 894 :96 Mason a nd Mau le , i nD ah l a nd H jor t 1 976 :190 N ewman 1 975 :97 Wa l lach a nd B oeve r 1 983 :198 C ozz i , i nD ah l a nd H jor t 1 976 :190 C ozz i , i nD ah l a nd H jo r t 1 976 :190 F ow le r 1 978 :791 Mason a nd Mau le , i nD ah l a nd H jor t 1 976 : 190 Wa l lach a nd B oeve r 1 983 :3 18

K abab ish -

T ab le 6 2 A ve rage L i vewe igh ts R epo r ted f o r Ma tu re B ac t r ians

KG 6 51-970 4 55 -850 7 00 7 00

S EX

LOCAT ION

BREED

SOURCE

M F

R uss ia R uss ia

-

-

K a lmyk K a lmyk -

T e ren t jev , i nD ah l a nd H jor t 1 976 :190 T e ren t jev , i nD ah l a nd H jo r t 1 976 :190 F ow le r 1 978 :791 Wa l lach a nd B oeve r 1 983 :318

I tw ou ld s eem t ha t l o ca l u n imp roved b reeds o fc ame ls v a ry w ide ly i nt he i r s i ze r anges , a s w as e a r l ie r o bse rved f o r P le is tocene a nd p os t -P le is tocene c a t t le p opu la t ions . T he b es t t ha t c an b e a ch ieved i nt he p resen t a na lys is i st oe s tab l ish r ea l is t ic a ge a nd s ex s pec i f ic l i vewe igh ts f r om t he a va i lab le d a ta , r ea l iz ing t ha t r eg iona l v a r ia t ions e x is t a cco rd ing t o t he u n imp roved b r eed o r b reeds f ound i ns pec i f ic l o ca l i t ies . Wh i le l i vewe igh ts a s h i gh a s 7 00 k g a re n o ted f o r d romeda r ies , t hese s eem t o b e e xcep t iona l a nd p resumab ly r ep resen t w e igh t r e su l ts o b ta ined t h rough t he ma in tenance o f a h i gh p l ane o f n u t r i t ion . S ince t he B enad i r b reed o fd romeda r ies f ound i nK enya a nd s ou the rn S oma l ia a re g ene ra l ly c ons ide red t ob e l a rge b y c ompa r ison w i th o the rc ame l b reeds ( Dah l a nd H jo r t 1 976 :190 ; Gau th ie r -P i l te rs a nd D agg 1 98 1:7 ) , t he mean l i vewe igh ts emp loyed i nt he p resen t s t udy s hou ld b e s omewha t l e ss t han t he 5 54 a nd 5 14 k g mean l i vewe igh ts o bserved f o r ma les a nd f ema les r espec t ive ly among t he B enad i r a nd B asso G iuba b reeds b y C ong iu ( 1 953 :189 ) . T hus , 4 50 t o 5 70 k g may b e s ugges ted a s ag ene rous u ppe r we igh t r ange f o r m a tu re d romeda r ies r a ised u nde rc ond i t ions o fe x tens ive h usband ry . F o rc ame ls j u s t e n te r ing a du l thood ( age c l asses 4 -5 t h rough 6 -7 ) , al i vewe igh t r ange f r om 4 00 a nd 5 00 k g s eems g ene rous , wh i le a r ange f r om 2 00 t o4 50 k g s hou ld r easonab ly a pp ly t o c ame ls i na ge c l asses 1 -2 t h rough 3 -4 . D ur ing t he i r f i r s t y ea r , t he r ange o f l i vewe igh ts s hou ld p resumab ly f a l l b e tween 3 5 a nd 2 50 k g . A ssum ing t ha t d romedary ma les wou ld g ene ra l ly o u twe igh f ema les b y a pp rox ima te ly 4 0 k g i na du l thood , t he e s t ima tes o f mean l i vewe igh ts g i ven

1 91

T ab le 6 3 E s t ima ted A ge a nd S ex S pec i f ic L i vewe igh ts f o rC ame ls i nt he Mode l H e rd

AGE C LASS 0-1 1-4 4-7 7-2 3

L IVE WE IGHT ( KG ) Ma les F ema les 1 48 3 35 4 65 5 30

1 38 3 15 4 35 4 90

i nT ab le 6 3 w i l l b e a ssumed a s a pp rop r ia te f o ra l b u t t he l a rges t b reeds o fd romeda r ies . T hese mean l i vewe igh ts a low e ach a n ima l i n a ge c l asses 4 -7 a pp rox ima te ly 8 8 t o 8 9% o f t he i r e xpec ted mean a du l t w e igh t .

Mea t , O rgan , F a t a nd B lood Y ie ld F rom t he Mode l C ame l H e rd U s ing t he v a lues f o re xpec ted p roduc t iv i ty p e r i n d iv idua l c ame l a s p rev ious ly p resen ted i n T ab le 6 0 a nd t he l i vewe igh t d a ta p resen ted i nT ab le 6 3 , t he e xpec ted mea t , o rgan , f a t a nd b l ood y i e lds p e r c ame l i nt he mode l h e rd h ave b een c a lcu la ted . T he r esu l ts a re p resen ted i n T ab le 6 4 . U s ing t hese v a lues a nd t he n o rma l o f take d a ta p rev ious ly p resen ted i nT ab le 5 5 , t he t o ta l s l augh te ry i e ld e xpec ted f r om t he mode l c ame l h e rd h as b een c a lcu la ted . T he r e su l ts a re p resen ted i nT ab le 6 5 . T ab le 6 6 p resen ts t he r e su l ts s im i la r ly o b ta ined u nde r max imum o f take a s p rev ious ly s u mma r ized i nT ab le 5 6 . F o rc ompa r ison , D ah l a nd H jo r t ( 1976 :192 ) o n ly c a lcu la ted o f f take f r om t he s l augh te ro fs i x y ea ro l d a do lescen t ma les , a ssum ing t ha t e ach c a rcass w ou ld y i e ld 2 25 k g o f mea t a nd 2 0 k g o f h ump f a t . S a to ( 1980 ) d i d n o t make a ny c a lcu la t ions o fc a lo r ic y i e ld f r om c ame l h e rd o f take . H av ing e s tab l ished t he e xpec ted w e igh t y i e ld p e r e d ib le i t em f o r t he mode l c ame l h e rd u nde r n o rma l a nd max imum o f f take , i tw i l l n ow b e p oss ib le t o c onve r t t hese v a lues t ot o ta l e ne rgy y i e ld o nce t he c a lo r ic v a lue p e rk i log ram o fe ach e d ib le i t em h as b een e s tab l ished .

T ab le 6 4 E xpec ted Mea t , O rgan , F a t a nd B lood Y ie lds P e r I n d iv idua l i nt he Mode l H e rd

S EX

F ema le

Ma le

A ge C lasses

Mean L i vewe igh t

Mea t

F a t

0 1 1 -4 4 -7 7 -23

1 38 3 15 4 35 4 90

6 6 .8 1 52 .5 2 10 .5 2 37 .2

4 . 1 9 .4 1.2 1 2 .6

0 .2 0 .5 0 .7 0 .7

3 .2 7 .3 1 0 .0 1.3

0 1 1 -4 4 -7 7 -22

1 48 3 35 4 65 5 30

7 1.6 1 62 .1 2 25 .1 2 56 .5

3 .5 8 .0 1 0 .0 1.4

0 .2 0 .5 0 .7 0 .8

3 .2 7 .2 1 0 .0 1.5

1 92

Y IELD P ER I TEM ( KG ) T ongue L i ve r H ea r t

K idneys

B lood

0 .7 1 .6 2 .2 2 .5

0 .8 1 .8 2 .4 2 .7

4 .8 1.0 1 5 .2 1 7 .2

0 .7 1 .7 2 .3 2 .7

0 .7 1 .5 2 . 1 2 .4

5 .2 1.7 1 6 .3 1 8 .6

T ab le 6 5 T o ta l M ea t , O rgan , a nd F a tY ie ld f r om t he Mode l C ame l H e rd w i th N o rma l O f f take

S EX

F ema le

M a le

TEM ( KG ) Y IELD P ER I L i ve r H ea r t T ongue

A ge C lasses

N umbe r C onsumed

Mea t

F a t

0 1 1 -4 4 -7 7 -23

0 .99 0 .51 0 .27 2 .66

6 6 .1 7 7 .8 5 6 .8 4 50 .7

4 . 1 4 .8 3 .0 2 3 .9

0 .2 0 .3 0 .2 1 .3

3 .2 3 .7 2 .7 2 1.5

0 1 1 -4 4 -7 7 -22

1 .70 0 .58 3 .52 0 .42

1 21.7 9 4 .0 7 92 .4 1 07 .7

6 .0 4 .6 3 5 .2 4 .8

0 .3 0 .3 2 .5 0 .3

5 .4 4 .2 3 5 .2 4 .8

1 0 .65

1 ,767 .2

8 6 .4

5 .4

8 0 .7

TOTAL

K idneys

B lood

4 .8 0 .8 0 .7 5 .6 0 .8 0 .9 4 .1 0 .6 0 .7 4 5 .8 4 .8 5 . 1 ------ -1 .2 1 .2 8 .8 1 .0 0 .8 6 .8 5 7 .4 8 .1 7 .4 1 . 1 1 .0 7 .8 1 8 .3

1 7 .9

1 41.1

T ab le 6 6 T o ta l Mea t , O rgan , a nd F a tY ie ld f r om t he Mode l C ame l H e rd w i th Max imum O f f take

S EX

F ema le

M a le

Y IELD P ER I TEM ( KG ) T ongue L i ve r H ea r t

A ge C lasses

N umbe r C onsumed

Mea t

F a t

0 -1 1 -4 4 -7 7 -23

1 .98 1 .01 2 .76 3 .42

1 32 .3 1 54 .0 5 81.0 8 11.2

8 .1 9 .5 3 0 .9 4 3 .0

0 .4 0 .5 1 .9 2 .4

6 .3 7 .4 2 7 .6 3 8 .7

0 -1 1 -4 4 -7 7 -22

3 .40 1 . 16 3 .65 0 .51

2 43 .4 1 88 .0 8 21.6 1 30 .8

1 2 .0 9 .3 3 6 .5 5 .8

0 .7 0 .6 2 .6 0 .4

1 7 .89

3 ,062 .3

1 55 .1

9 .5

TOTAL

K idneys

B lood

1 .4 1 .6 6 . 1 8 .6

1 .6 1 .8 6 .6 9 .2

1 0 .9 8 .4 3 6 .5 5 .9

2 .4 2 .0 , 8 .4 1 .4

2 .4 1 .7 7 .7 1 .2

9 .5 1.1 4 2 .0 5 8 .8 -1 7 .7 1 3 .6 5 9 .5 9 .5

1 41.7

3 1.9

3 2 .2

2 21.7

C a lo r ic Y ie ld o f Mea t , O rgans , F a t a nd B lood f r om t he Mode l C ame l H e rd S tandard A na lyses o fC ame l Mea ta nd S easona l V a r ia t ions i nF a tC on ten t Two d i f fe r ing c ompos i t iona l a na lyses o f c ame l mea t h ave b een l o ca ted i nt he a va i lab le l i t e ra tu re . T he f i r s t o ft hese w as a s t udy o n t he e f fec ts o f a ge a nd s ex o n t he c ompos i t ion o f c ame l mea t s o ld i nt he mea t ma rke ts o fC a i ro ( Nas r e ta l . 1 965 ) . I nt h is s t udy , 2 0 ma le a nd 2 0 f ema le c a rcasses w e re c hosen f o r a na lys is . F o re ach s ex , 1 0 c a rcasses w e re o fi n d iv idua ls l e ss t h an 5y ea rs o fa ge , a nd 1 0 w e re o fi n d iv idua ls o l de rt han 5y ea rs o fa ge . F rom e ach c a rcass , 1 k g e ach o f r ump , c huck , a nd r i b mea t was s ub jec ted t o s t anda rd a na lys is . T he c ompos i t iona l r e su l ts o ft h is s t udy a re p resen ted i nT ab le 6 7 . Wh i le m in ima lv a r ia t ions w e re f ound a s ar e su l to f s ex , h i ghe r p ro te in a nd c r ude f a t r a tes w e re o bserved among o l de rc ame ls t han y ounge ro nes . P ar t icu la r ly s t r ik ing i st he l ow f a t c on ten t o f a l s amp les ( mean 0 .97% ) , s ugges t ing l e an m ea t . I ft h is l ow f a t l e ve lw as i n deed r ep resen ta t ive o fc ame l mea t i ng ene ra l , i t s c a lo r ic c on ten t 1 93

T ab le 6 7 S tanda rd A na lyses o fC ame l Mea t P u rchased i nC a i ro

° / . 3F AT

° A ASH

7 6 .10 7 6 .14 7 6 .30

0 .86 0 .95 0 .96

0 .72 0 .79 0 .86

2 1.77 2 1.65 2 2 .03

7 6 .68 7 6 .41 7 5 .82

0 .69 1 .00 1 .60

1 .01 1 .05 0 .77

r ump c huck r i b

2 0 .33 1 9 .95 1 9 .88

7 8 .04 7 8 .72 7 8 .09

0 .96 0 .78 1 .00

0 .78 0 .63 0 .98

r ump c huck r i b

1 9 .98 2 0 .02 2 0 .26

7 8 .54 7 8 .41 7 7 .85

0 .83 0 .98 0 .98

0 .69 0 .62 0 .88

MEAN OF A LL CUTS , 5+YEARS

2 2 .02

7 6 .24

1 .01

0 .86

MEAN OF A LL CUTS ,