Prey Movements and Settlement Patterns During the Upper Palaeolithic in Southwestern France 9780860548003, 9781407349244

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Prey Movements and Settlement Patterns During the Upper Palaeolithic in Southwestern France

Ariane M. Burke

BAR International Series 619 1995

Published in 2019 by BAR Publishing, Oxford BAR International Series 619 Prey Movements and Settlement Patterns During the Upper Palaeolithic in Southwestern France © Ariane M. Burke and the Publisher 1995 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 9780860548003 paperback ISBN 9781407349244 e-book DOI https://doi.org/10.30861/9780860548003 A catalogue record for this book is available from the British Library This book is available at www.barpublishing.com BAR Publishing is the trading name of British Archaeological Reports (Oxford) Ltd. British Archaeological Reports was first incorporated in 1974 to publish the BAR Series, International and British. In 1992 Hadrian Books Ltd became part of the BAR group. This volume was originally published by Tempvs Reparatvm in conjunction with British Archaeological Reports (Oxford) Ltd / Hadrian Books Ltd, the Series principal publisher, in 1995. This present volume is published by BAR Publishing, 2019.

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ACKNO WLEDGE MENTS T he s uppor t o f f e l low r e sea r che r s , f a cu l ty a nd s t a f f a t t h e D epar tmen t o f A n thropo logy , N ew Y o rk Un ive r s i ty , t h e L abo ra to ire d 'Ana to tn ie C ompa ree , U n ive r s i te d e P ar i s V I I , t h e I n s t i tu t d u Q ua te rna ire , U n ive r s i te d e B o rdeaux I I , t h e Z ooa r chaeo logy l a bo ra to ry , P e abody Mu seum , a nd t h e C .M .R .A .E . ( M .I .T . & H a rva rd ) i sg r a tef u l ly a cknow ledged . S pe c ia l t h ank s a r e d ue t oD . We lsh w ho d ona ted s eve ra l o fh i s own s amp le s o fh o r se t e e th f o ru se i nt h e c on t ro l g roup . V a luab le c on tr ibu t ion s t ot h i s r e sea r ch w e re o f f e red b y : D r . R anda l l Wh i te , D r . P am C rab t re e , a nd D r . R i ta Wr igh t , N ew Y o rk U n ive r s i ty ; D r , R icha rd Meadow , H ar va rd U n ive r s i ty ; D r . J a cque s C a s tane t , U n ive r s i te d e P ar i s V I I a nd D r . A nne P ike -Tay , V a s sa r . W i thou t t h e ir i n pu t t h is r e sea rch w ou ld n eve r h ave b een c omp le ted . A cce s s t oc ruc ia l a r chaeo log ica l c o l le c t ion s w a s m ade p o s s ib le t h rough t h e a genc ie s o fJ . -P . R igaud , H . L av i l le , F . D e lpe ch , M . L eno ir , J . -P . R ayna l , H . D e lpor te , a nd c o l league s a t t h e I n s t i tu t d u Q ua te rna ire , U n ive r s i te d e B o rdeaux , t h e l a te R . A rambou rou , A . R ou s so t ( Muse& d 'Aqu i ta ine , B o rdeaux ) a nd J . J . C leye t -Me r le ( Muse& N a t iona l d e P reh i s to i re , l e sE yz ie s ) . T hank s a r ed ue t ot h ev ar iou se xcava to r s o ft h e s amp led s i te s , w ho g ene rou s ly a l lowed m e t oa pp ly a n i n he r en t ly d e s t ru c t ive f o rm o fa na ly s is t ot h e ir c o l le c t ion s . L abwo rk f o rt h is r e sea r ch w a s c ar r ied o u t a tt h e D ep t . o fA n th ropo logy A rchaeo log ica l L abo ra to ry , N ew Y o rk U n ive r s i ty , a tt h e I n s t i tu td u Q ua te rna ir e , Un ive r s i te d e B o rdeaux I I , a nd a tt h e L abo ra to ire d 'Ana tom ie C ompa re , Un ive r s i te d e P a r is V I I . A dd i t iona ls uppor tw a s o f f e red b y F . P r a t t , I n s t i tu td u Q ua terna ire , B o rdeaux , B . G o rdon , a nd t h e l a te S . P re s ley , A rchaeo log ica l S urveys o fC anada , a nd A . Mo r a la , Mu se& N a t iona ld eP reh is to ire , l e sE yz ie s . S uppo r to f ad i f f e r en t n a tu re w a s p rov ided t om e b y f r iend s a nd f am i ly t o o n ume rou s t om en t ion i nd iv idua l ly , b u t t ow hom Io we ag rea t d eb t o f t h ank s . S pe c ia l t h ank sa r ed ue t oD r . & Mr s . R igaud , t o Mr s M . S a i l ly , a nd t oJ . -M . G ene s te , f o rt h e ir u nf a i l ing h o sp i ta l i ty . T h i sr e sea rch w a ss uppo r ted b yt h eS oc ia l S c ience s &H uman i t ie sR e sea r ch C ounc i l o fC anada , t h eF rench G ove rnmen t ( Cha teaubr iand S cho la r sh ip ) , a nd b y N ew Y o rk U n ive r s i ty ( J ame s A r thu r S cho la r sh ip ) .

i i

TABLE OF CONTENTS D ed ica t ion A cknow ledgemen t s L i s to fF igu r e s L i s to fA ppend ic e s

i i i x x CHAPTER 1 :I NTRODUCT ION

1 . 1 .0 R eg iona la nd C h rono log ica lF ramewo rk

p . 1 p . 1 p . 1 p . 2

1 . 1 .1 R eg iona lP e r sp ec t ive so nH un te r-Ga th e r e r s 1 . 1 .2 T he S ou thwe s to fF rance : H i s to r ica lO ve r v iew 1 . 1 .3 T he P e r iod o fS t udy : t h eE a r ly U pp e rP a laeo l i th ic

1 . 2G ene ra l Mode l s 1 . 3 S ea sona l i ty a s aMean so fT es t ing S e t t lemen ta nd S ub s is ten ce Mode l s 1 . 4S e t t lemen tS y s tem s : A r chaeo log ica l Mode l sf o rS ou thwe s t er nF ran ce 1 . 5 Me thodo logy f o rO b ta in ing S ea sona lD e t e rm ina t ion sf r om F auna lR ema in s 1 . 6S pec ies o fI n te r e s t 1 . 7S ke le to ch rono log ica lS t ud ie s : C emen tum A na ly s i s 1 . 8T he A r chaeo log ica l S amp le 1 . 9C onc lu s ion

p . 2 p . 2 p . 2 1 1 3 1 1 3 1 1 3 p . 4 p . 4

PART A . SKELETOCHRONOLOGY OF HORSE ( E .c aba l lu s ) TEETH . CHAPTER 2 PART A : H a rd t i s sue s . 2 . 0 2 . 1 2 . 2 2 . 3

I n t roduc t ion B one a nd C emen tum : S t ruc tu re a nd C ompo s i t ion M ine ra l i sa t ion C o l lagen O r ien ta t ion a nd T ype so fB one

P -5 1 1 5 1 1 5 p . 6

PART B :I n c r emen ta lG row th S t r uc tu r e s . p . 6 p . 8 p . 8

2 . 4T e rm ino logy 2 . 5N a tu re o ft h eG row th S t ruc tu re s 2 . 6 .0 T echn ique sf o rO b se r v ing G row th L aye r s 2 . 6 .1 2 . 6 .2 2 . 6 .3 2 . 6 .4 2 . 6 .5 2 . 6 .6 2 . 6 .7

p . 8 p . 8 p . 8

P o la r ized L igh t N a tu ra lL igh t S ta ined H i s to log ica lT h in S ec t ion s T ran sm i t ted E l ec t ron M ic ro scope S cann ing E lec t ron M ic ro scop e H i s tochem ica lA na ly se s M ic ro rad iog raphy

1 1 8 p . 8 1 1 9 1 1 9

2 . 7D i scu s s ion :G row th L aye r s i nC emen t 2 . 8D epo s i t ion o ft h eG row th L aye r s 2 . 9 .0 R a te a nd D eg r ee o fM ine ra l i sa t ion

1 1 9 1 1 9 p . 10

2 . 9 .1 P r ima ry ,S econda ry a nd A cce s so ry G row th L aye r s 2 . 9 .2 A nnua lG row th L aye r s

p . 10 p . 10

i v

2 . 10 .0 C au sa lF ac to r si nt h eC y c l ica lD epo s i t ion o fG row th L aye r s 2 . 10 .1 E nv i ronmen ta lF ac to r s p . 11 2 . 10 .2 E ndog enou sF ac to r s p . 11 2 . 10 .3 G ene ra lC onc lu s ion s p . 12

p . 11

2 . 11 .0 P opu la t ion S t ud ie s

p . 12 CHAPTER 3 : THE CONTROL S A MPLE PART A :T h e Mod em C on t ro lS amp le .

3 . 0 3 . 1 3 . 2 3 . 3

I n t roduc t ion A pp l icab i l i ty o ft h e Mod em C on t ro lt oF o s s i lS pe c ie s E f f ec t so ft h eE nv i ronmen to nt h eE xp r e s s ion o fG row th L aye r s E s tab l i sh ing t h eC on t ro lS amp le

p . 14 p . 14 p . 15 p . 15

P ART B :A na ly s i so fG row th S t ru c tu r e si nt h eT ee th o fE .c aba l lu s . 3 . 4 .0 Me thods a nd Ma te r ia l s 3 . 4 .1 U ndeca lc i f ied T h in S ec t ion s 3 . 4 .2 S cann ing E l ec t ron M ic ro scope 3 . 4 .3 M ic ro rad iog raphy 3 . 4 .4 R ef l ec ted L igh t 3 . 4 .5 D eca lc if ied T h in S ec t ion s 3 . 4 .6 I mag eA na ly s i s

p . 16 p . 16 p . 18 p . 19 p . 19 p . 19 p . 19

3 . 5 .0 R e su l t s

p . 19 3 . 5 .1 S cann ing E l ec t ron M ic ro scope 3 . 5 .2 H i s to log ica lP r epa ra t ion s 3 . 5 .3 M ic ro rad iog raphy 3 . 5 .4 R ef l ec ted L igh t 3 . 5 .5 T ran sm i t ted L igh t

p . 19 p . 19 p . 23 p . 23 p . 23

3 . 6 .0 D i scu s s ion /Con c lu s ion 3 . 6 .1 H i s to log ica ls amp les 3 . 6 .2 T ran sm i t t ed a nd R ef l ec ted L igh t 3 . 6 .3 C on c lu s ion s

p . 23 p . 23 p . 23 p . 24

3 . 7 .0 A pp l ica t ion so fS k e le to ch rono logy 3 . 7 .1 A ge D e te rm ina t ion s :I n t roduc t ion 3 . 7 .2 Mo rphome t r ic A ge D e te rm ina t ion s 3 . 7 .3 S ke le to ch rono log ica l Mean so f D e te rm in ing A ge 3 . 7 .4 C on c lu s ion :A ge D e te rm ina t ion s 3 . 7 .5 D e te rm ina t ion o fS ea son o fD ea th 3 . 8S amp le R e su l t s :S ea sona l i ty

p . 24 p . 24 p . 24 p . 26 p . 27 p . 27 p . 27

PART B . APPL IED SKELETOCHRONOLOGY . CHAPTER 4 : REGIONAL CHRONOLOGICAL , GEOLOG ICAL AND CULTURAL F RA ME WORK OF THE APPL IED RESEARCH . p . 30 p . 30 p . 32 p . 32

4 . 0I n t roduc t ion 4 . 1 G eo log ica lH i s to ry o fS ou thwe s tF ran ce 4 . 2C h rono log ica lB a ckg round 4 . 3 .0 C l ima to log ica lR econ s t ruc t ion s p . 32 p . 35 p . 35

4 . 3 .1 G eo c l ima t ic D a ta 4 . 3 .2 B io c l ima t ic I n d ica to r s 4 . 3 .3 C onc lu s ion 4 . 4 .0 C u l tu ra lF ramewo rk

p . 38 p . 38 p . 38 p . 39

4 . 4 .1 I n t roduc t ion 4 . 4 .2 Magda len ian S y s tema t ic s 4 . 4 .3 C onc lu s ion 4 . 5 .0 I n t roduc t ion :S i te sS amp led

p . 39 p . 39 p . 4 1 p . 4 1 p . 43 p . 43 p . 43 p . 45 p . 45 p . 45 p . 47 p . 47 p . 49

4 . 5 .1 L a Made le ine 4 . 5 .2 R e igna c 4 . 5 .3 C ap B lanc 4 . 5 .4 B adegou le 4 . 5 .5 L auge r ie H au te E s t 4 . 5 .6 S a in t-Ge rma inlar iv ie re 4 . 5 .7 R o cd e Ma r camp s 4 . 5 .8 Mou l in N eu f 4 . 5 .9 D u ru thy 4 . 5 .10 R ond d uB a r ry 4 . 5 .11 E nva l 4 . 5 .12 E sc lauzu r

CHAPTER 5 : APPL ICAT ION OF CE MENTU M ANALYS IS TO THE FOSS IL S A MPLE . 5 . 0I n t roduc t ion 5 . 1 S amp l ing S t ra tegy 5 . 2 Me thod sa nd Ma te r ia ls

p . 50 p . 50 p . 50

5 . 3 .0 R e su l t s

p . 54 p . 54 p . 54 p . 54

5 . 3 .1 C ompa ra t ive S t udy 5 . 3 .2 T ran sm i t ted L igh t 5 . 3 .3 S ea sona lD e te rm ina t ion sb yS i te

5 . 4D i scu s s ion 5 . 5C on c lu s ion : A pp l icab i l i ty o fS ke le to ch rono logy t oF o s s i l Ho r se T ee th

p . 56 p . 56

5 . 6 .0 A r chaeo log ica lI mp l ica t ion so ft h eS t udy

p . 56

5 . 6 .1 I n t roduc t ion p . 56 5 . 6 .2 E tho log ica lC on s ide ra t ion sf o rS ea sona l i ty S t ud ie s p . 56 5 . 6 .3 S ea sona l i ty S t ud ie s : O the rS ea sona l

v i

I n dexe s

p . 59

5 . 7 .0 D i scu s s ion : R eg iona lP a t te r n ing a nd I n d ica to r so fS ea sona l i ty i nt h eF auna l A s semb lage s p . 60 5 . 7 .1 5 . 7 .2 5 . 7 .3 5 . 7 .4

T he T he T he T he

p . 60 p . 6 1 p . 6 1 p . 6 1

D o rdogn e G i ronde Ma s s i fC en t ra l L ande s

5 . 8C on c lu s ion s

p . 62 CHAPTER 6 : HUNTER-GATHERER MODELS .

6 . 0 .0 I n t roduc t ion

1 1 64 p . 64

6 . 0 .1 H u n t e r-Ga the r e rR e sea rch

p . 64

6 . 1 .0 B and t h eo r y :t h eC ompo s i te B and p . 65

6 . 1 .1 C r i t ique o fB and T heo ry

p . 65

6 . 2 .0 G ene ra l S ub s i s tenc e Mode l si nA r chaeo logy 6 . 2 .1 G ene ra l Mode l so fS e t t lemen t O rgan i sa t ion 6 . 2 .2 C r i t ique o fG en e ra lS ub s i s tence Mode l s 6 . 2 .3 C on c lu s ion s :G en e ra lS ub s i s tenc e Mod e l s

1 0. 6 6 p . 66 p .6 6

6 . 3 R eg iona l Mode l so fH un te r-Ga the r e rS e t t lemen ta nd S ub s i s tence

p . 67

6 . 3 .1 I n t roduc t ion : I n teg ra t ing G ene ra lT heo ry a nd A r chaeo log ica lT heo ry p . 67 6 . 4 .0 Mode l so fE a r ly P a laeo l i th ic S e t t lemen ta nd S ub s i s ten ce

p . 68 p . 68 p . 69 p . 70

6 . 4 .1 H e rd F o l low ing Mode l s 6 . 4 .2 S ea sona lS e t t lemen tS h i f t Mode l s 6 . 4 .3 D i scu s s ion : S ea sona lE v iden ce

p . 7 1

6 . 5C onc lu s ion s

CHAPTER 7 : CONCLUS IONS OF THE RESEARCH . p . 72

7 . 0 .0 C emen tum A na ly s i s 7 . 0 .1 G row th S t ru c tu r e si nD en ta lC emen t 7 . 0 .2 O b se r va t ion o fG row th L aye r s 7 . 0 .3 P a t ter no fD epo s i t ion o fC emen t i nE .c aba l lu s

p . 72 p . 72 p . 72

7 . 1 A pp l ica t ion o ft h e Moder nC on t ro lS amp le t oF o s s i lH o r se P opu la t ion s

p . 73

7 . 2 Mo rphome t r ic V e r su s S k e le to ch rono log ica l A na ly se s

p . 73

v i i

7 . 3R eg iona la nd C h rono log ica lF ramewo rk

p . 73

7 . 4 .0 E tho logy o f Ma jo rP rey S pec ie s

p . 74

7 . 4 .1 S ea sona lD a ta f o rR e indee r 7 . 4 .2 S ea sona lD a ta f o rH o r se 7 . 4 .3 E tho logy a s aP red ic to ro fH uman B ehav iou r

p . 74 p . 74 p . 76

7 . 5T he A r chaeo log ica l S amp le a s aR e f lec t ion o fH uman B ehav iou r

p . 76

7 . 6S ea sona lI n d ica to r sa sE v idence o fS ub s i s ten ce a nd S e t t lemen tS y s tem s

p . 76

7 . 7 .0 Mode l so fS ub s i s tenc ea nd S e t t lemen ti nt h eF r en ch E a r ly U ppe r P a laeo l i th ic

p . 77

7 . 7 .1 H e rd F o l low ing Mode l s 7 . 7 .2 T he S ea sona lR ound s Mode l 7 . 7 .3 C onc lu s ion

p . 77 p . 77 p . 77

7 . 8 .0 S ugge s t ion sf o rf u r the rr e sea rch 7 . 8 .1 T heo re t ica lr e sea r ch 7 . 8 .2 C emen tum a na ly s i s

p . 78 p . 78 p . 78

Em ino te s

p . 79

B ib l iog raphy

p . 81

A ppend ix A

p . 97

A pp end ix B

p . 100

A ppend ix C

p . 124

P la te s

p . 127

L is to fF igure s ,T ab le sa nd P ho tograph ic P la te s . T ab le s . 3 . 1 Con t ro lS amp le a nd I n ve s t iga t ive T echn ique s U sed . 3 . 2Con t ro lS amp le .R e su l t s u nde rT ran sm i t ted L igh t . 3 . 2b-Con t ro l S amp le .R e su l ts u nde rR ef lec ted L igh t . 3 . 2c-Con t ro l S amp le . R e su l ts f r om S ta ined S pec imen s . 3 . 3 Re la t ion sh ip B e tween A ge E s t ima tes , N umbe ro fI n c r emen t s a nd C rown H e igh t . 4 . 1 Env ironmen ta l i n d ica to r s . 4 .2 Fauna , L a Made le ine . 4 . 3 Fauna ,L auger ie H au te E s t . 4 .4 Fauna ,R o cd e Ma rcamp s . 4 . 5Fauna ,D u ru thy . 5 . 1 Samp le P roven ience . 5 . 2So r t ing P ro toco l . 5 . 3 Syn the t ic R e su l t so ft h eF o ss i l A na ly s is . F igu r e s . 2 . 1- C ro ss S ec t ion T oo th : R e la t ion sh ip B e tween A nnu l i , LAG s a nd G row th Z one s . 3 . 13 . 23 . 33 . 44 . 14 .24 . 3-

O r ien ta t ion o fT oo th S ec t ion s . R ad ia l S ec t ion : P a t ter no fD epo s i t ion . E s t ima ted A ge v e r sus n o .o fG row th I n c r emen t s . S ea sona lR hy thm o fD epo s i t ion o ft h eG row th L aye r s . T he Aqu i ta ine R eg ion . C o r re la t ing L ines o fE v idence . E nv i ronmen ta lR econ s t ruc t ion f o rt h eA qu i ta ine B a s in .

4 .4- S yn the s is : B iogeog raph ica l R econ s t ruc t ion s f o rt h eA qu i ta ine B a s in . 4 . 55 . 15 . 27 . 1-

L oca t ion o ft h e S amp led S i te s . D en ta l Mea su remen t s . D en ta l Wea rP a t ter n s a nd A n t le rG row th ( R . t a randu s ) . P a leo-Range R econ s truc t ion f o rR e indee r .

P ho tog raph ic P la tes . 3 . 1- Mand ibu la rT oo th R ow , S ag i t ta l a nd B ucca l V iew s . 3 . 2- S .E .M .O b serva t ion o fG row th L aye r s . 3 . 3- H is to log ica lP repa ra t ion . 3 . 4- M ic ro rad iog raph . 3 . 5a-T ran sm i t ted ,O rd ina ry L igh t . 3 . 5b-Po la r ised L igh t . 3 . 6- P r ima ry C emen tum L aye r . 3 . 75 . 15 . 25 . 3-

R e so rp t ion B ays i nE name l . M ic ro rad iog raphy o fF o ss i l T oo th . T ran sm i t ted L igh tO b ser va t ion o fF o ss i l . E f f ec t so fP oo rP rese rva t ion o nO b serva t ion so fG row th L aye r s i nF o ss i lC emen tum .

L is to fA ppend ice s A pp end ix A D en ta l Mea su remen t so fF o ss i l S amp le . A ppend ix B D en s i ty g r aph s ( l um ina nce p rof i les ) f o r c on t ro l s amp le s pec imen s v i ewed u nde r t r an sm i t ted p o la r i sed l i gh t . A pp end ix C R esu l t so ft h e F o s s i l S amp le .

CHAPTER 1 I NTRODUCTION . S ou thwe s tern F rance ( e .g . , B ahn 1 984 ) , p rov id ing e v idence o f s oc ia l i n te rac t ion a tt h e l e ve l o f t h e l a rge r r eg ion , i . e . ,

1 .1 .0 R eg iona l a nd Chrono log ica l F ramework .

t h e Aqu i ta ine B a s in . 1 . 1 .1 Reg iona l P er spec t ive s o n Hun ter -Ga therer s . T he Aqu i ta ine i so f ten s ubd iv ided f o r t h e T he i mpor tance o f ar e g iona l p e r spec t ive

p urpose s o f a r chaeo log ica l i nve s t iga t ion , i n to t r ad i t iona l

f o rt h es t udy o fh un te r -ga the re r si so bv iou si fo ne c on s ide r s

s ubreg ion s ( e .g . , t h e P 6r igord ) wh ich a re a s sumed t o b e

t h a t h uman a c t ion - a s r e f le c ted i n m a te r ia l c u l tu re - i sn o t

s i gn if ican t u n i t s o f a na lys is , w i th o r w i thou t j u s t if ica t ion

t h e r e su l t o f r andom b ehav iou r , b u t i s t h e r e su l t o f a

( Conkey 1 987 ) . Howeve r ,b iogeograph ic r e con s truc t ion sf o r

c u l tura l

s i te s d o n o t e x is t i n

t h e S ou thwe s td iv ide t he r eg ion i n to mean ingfu l s ubreg ion s

i s o la t ion . T hey a r e p a r t o f ar eg iona l p a t te rn o fa dap ta t ion ,

r ough ly c or re spond ing t ot h e se " t rad i t iona l " d iv is ion s ( s ee

s y s tem .

A rchaeo log ica l

i . e . , as e t t lemen ts y s tem . T h is p e r spec t ive a cknow ledge st h e

C hap te r4 ,b e low ) . T he p re sen tr e sea rch e ncompa sse s a l lo f

e x is tence o f " t he l a rge r n a tu ra l a nd c u l tu ra l u n i t s w i th

t h e Aqu i ta ine B a s in , wh i le a cknow ledg ing t h e e x is tence o f

wh ich i nd iv idua l s i te s a r t icu la ted " ( Be t t inge r 1 982 :161 ) .

d i f f e ren t ia lp a t tern ing w i th in t h is r eg ion wh ich may i nd ica te t h a t f u r the r s ubd iv is ion i sp oss ib le .

E thnog raph ic t heor ie so fb and o rgan isa t ion 1 .1 .2 The S ou thwe s t o fF rance : H is tor ica l Overv iew .

( s ee c on tr ibu tor st oL ee & D evore 1 968 ; a nd D ama s 1 969 ) t r ea t t h e r eg iona l b and a s ad ef inab le l e ve l o f h un te r -

T he P a leo l i th ic o f S ou thwe s t F rance h a s

g a the re r s oc ia l o rgan isa t ion , wh ich " may b e i d en t if ied a sa

b een a f o cus o fa r chaeo log ica l i nve s t iga t ion f o r more t h an

r e la t ive ly e ndur ing e n t i ty i nr e la t ion t o ac o mmon r a nge o f e xp lo i ta t ion " ( I ngo ld 1 980 :266 ) . F ur thermo re , t h er e g iona l

1 25 y ea r s . T he f i r s t r i ch f i nds o f a r chaeo log ica l m a ter ia l

b and h a s b een e t hnog raph ica l ly p roven t o h ave c u l tu ra l

f r om

m ean ing f o r i t s membe r s ( i .e . , i ti sd ef ineab le f r om t h e ir

d epar temen t o ft h e Dordogne ) s t imu la ted mode r n s c ien t if ic

own c u l tu ra l p e r spe c t ive ) , a nd i n te rac t ion o n ar e g iona l

i n te re s t i nE uropean P reh is tory .

s c a le

among

d i spe r sed

g roups

o f h un te r -ga the re r s

t h e

s ubreg ion

o f

t h e

Kr igord

( t he

modern

i s T he Aqu i ta ine r eg ion h a s a l ong a nd

d ocumen ted ( e .g . , B rown 1 987 :374 ) . T he r e g ion , t h e ref ore , V ar ia t ion a mong

u n in ter rup ted h i s tory o fh uman o ccupa t ion . Un l ike i t s more

h un te r -ga the re r a dap ta t ion s i s c ommon ly o bserved a t t h e

n or the r ly E u ropean n e ighbou r s , t he Aqu i ta ine r ema ined i c e -

h a s more t h an g eograph ic m ean ing .

r eg iona l l e ve l ( Gamb le 1 986 :40 ) i n d ica t ing t h a t ar e g iona l

f r ee d ur ing t h e l a s t f u l l g l ac ia l p er iod , r e su l t ing i n t h e

s c a le o f a na lys is i s c apab le o f g ene ra t ing m ean ingfu l c ompar ison s . F or t h e se r e a son s , t h e p re sen t r e sea rch i s

a ccumu la t ion o fc u l tura l d epos i ts t ha t s pan m i l lenn ia . T he

c onduc ted a tt h e r eg iona l l e ve l .

b een t h e f o cu s o f much s ed imen to log ica l , p a lyno log ica la nd

l ong s t ra t igraph ic c o lumn s f r om S ou thwe s tern F rance h ave p a laeon to log ica l

r e sea rch

a imed

a t

e s tab l ish ing

a

d epos i t iona l h i s tory o ft h e r e g ion i .e . , ar e la t ive , r eg iona l

A l though r e g iona la na ly s is s t and sa c cu sed

c h ronos tra t igraphy .

o f p lac ing u ndue e mpha s is o n t h e e nv ironmen t , i . e . , o f e s tab l ish ing a n e nv ironmen ta l p r imacy ( Conkey 1 987 :66 ) , t h e i mpor tance o ft h e e nv ironmen ta l c on tex t s hou ld n o t b e

T he c h ronos tra t igraphy o f t h e Aqu i ta ine

m in im ized . A more b a lanced v i ew o f r e g iona l a na lys is

b a s in i sb a sed o n t h e i d en t if ica t ion o f r eg iona l p a t tern s i n

" encompa sse s t h ed e term in ing f e a tu re s o ft h e e nv ironmen t ,

t h e s equence o fd epos i t iona l e ven t s , u sua l ly p re c ip i ta ted b y

t o wh ich g roups mu s t a dap t , b u t a l so t h e c on t inua lp roce s s o f s oc ia l r eproduc t ion [ . .]" ( Gamb le 1 986 :31 ) . A s I ngo ld

c l ima t ic c hange ( e .g . , L av i l le e ta l . 1 980 ) . T he s equence o f

( 1981) a rgue s , t h ee nv ironmen ti mpo se sl im i t ing c on s t ra in t s o n e conom ic p roduc t ion wh ich mu s t b e c ons ide red e ven i f

S ou thwe s t o fF rance h a s t h e refore b een af ocu s o fi n te re s t .

i ti s t h e s oc ia l s ys tem wh ich d e term ine s t h e s t ruc tu re o f

g eograph ica l ly

e conom ic p roduc t ion .

c h rono log ica l s equence w i th s o me d egree o f e xac t itude . I t

g l ac ia l / in te rg lac ia l a nd s t ad ia l/ in te r s tad ia l e ven t s i n t h e C hronos t ra t ig raph ic r e sear ch a l low sa r chaeo log is t s t or e la te i s o la ted

l o ca l

d epos i t s

t o

a r eg iona l

i st h e refore p os s ib le t o s t udy a r chaeo log ica l d epos i t s f r om T he s c a le o f t h e r eg iona l " c a tchmen t " i s

t h e S ou thwe s t o f F rance o n ar eg iona l b as is , w i th s ome

a n i mpo r tan t c ons ide ra t ion ( Brown 1 987 ) . I n o the r w o rd s ,

c on t ro l

t h e r eg ion o f s t udy mu s t b e d ef ined . T he A qu i ta ine B a s in

s y s tema t ics ( Lav i l le e ta L 1 980 :11) .

o ve r

c h rono logy ,

w i thou t

r e ly ing

o n

c u l tura l

e x is t s a s ad i scre te g eograph ica l a nd c l ima t ic u n i t ( Gamb le 1 986 F igu re 3 .9 ; s e e C hap te r 3 , b e low ) wh ich m ay ,

T he c u l tura l c h rono logy o f t he Uppe r

t h erefore , r epre sen t ar eg ion f r om t h e p o in t o fv i ew o ft h e " common r ange o f e xp lo i ta t ion " r e f er red t o i n I ngo ld

P a laeo l i th ic i s at horny i s sue , l a rge ly d ue t o me thodo log ica l p rob lem s s uch a s u se o f d i agnos t ic t oo l t ypes , o rf os s i le s

( 1980 :266 ) . T he movemen t o f r aw a nd e xo t ic m a te r ia ls

d i rec teur s , f o rt empo ra l s e r ia t ion . T h is s e r ia t ion r e s t so nt h e

b e tween

i d en t if ica t ion o f d i scre te c u l tu ra l u n i t s o n t h e b a s is o f

P a laeo l i th ic

s i te s

i s

w e l l

d ocumen ted

i n

1

t ypo log ica le l emen t s( f o s s i le sd i rec teur s ) , wh ich a r ec hosen a rb i t ra r i ly ( Lav i l le e t a l . 1 980 :14 ) . T h is i s n o l onge r u n ive r sa l ly a ccep ted a s av a l id m e thodo log ica l a pp roach

i s p re sen t i n t h e a r chaeo log ica l r e co rd o f S ou thwe s te rn F rance i ss t i l l l a ck ing , h oweve r .

( e .g . , S a cke t t 1 981 :89 ; G amb le 1 986 :24f f ) .

T he i d en t if ica t ion o f a gg rega t ion s i te s i n t h e P a laeo l i th ic l a nd scape ( a s e v idence o f t h e e x i s tence o f ac yc le o f f i s s ion a nd f u s ion ) i sb a sed o n an umb e r o f

1 . 1 .3 T he P er iod o fS tudy : t he Upper P a laeo l i th ic . T he Uppe rP a laeo l i th ic i s ak ey p e r iod f o r

a r chaeo log ica l d a ta , e . g . , s i te s i ze ( Dav id 1 973 ; F re eman 1 973 ; C la rk 1 980 ; Wh i te 1 985 ) a nd v ar iou s m ea su re s o f

t h e s t udy o fh uman s oc ia l d eve lopmen t i nE u rope . Du r ing t h e Uppe rP a laeo l i th ic , ap ro l if e ra t ion o fa r t is t ic ( s ymbo l ic ) a c t iv i ty o ccu r s wh ich a r chaeo log i s t s c on s ide r e v idence o f

d ive r s i ty ( e .g . , C onkey 1 980 ; G amb le 1 983 ; Wh i te 1 985 ) . Howeve r , t h el i nk b e tween t h e se d a ta a nd a gg rega t ion s i te s i s s ome t im e s t e nuou s ( s ee C hap te r 5 ) . F ur the rmor e , t h e

h e igh tened s oc ia l i n te ra c t ion ( s e e F reeman 1 973 ; Wob s t 1 977 ; C onkey 1 980 ; G amb le 1 983 ; J och im 1 983 ; Wh i te 1 985 ) . O the r i nd ica t ion s o fc hange i n t h e h uman a dap t ive s y s tem , s uch a s at r end t owa rd s i n c rea se i n m a te r ia l

r e l iab i l i ty o f t h e se d a ta r e s t s o n s t ra t ig raph ic ( h ence , c h rono log ica l ) c on t ro l , wh ich i sn o te a s i ly a s se s sed (Wh i te 1 987 :264 ) .

c omp lex i ty a nd m ea su re s o ff a una l d ive r s i ty ( e .g . , Me l la r s 1 985 ; J och im 1 987 ; R igaud & S imek 1 990 ) a ppea r d u r ing t h i s t ime , l e ad ing m any r e sea rche r s t o s u spe c t t h a t

1 .2 .0 G enera l mode l s o fs ub s is tence a nd s e t t lemen t .

An umbe ro fa r chaeo log ica l mode ls f o rt h e Uppe r P a laeo l i th ic u se c oncep t s d rawn f r om e ne rge t ic s mode l s s uch a s o p t ima l f o rag ing t h eo ry , e . g . , o p t ima l is a t ion , i nhe ren t i n h e rd f o l low ing mode ls ( s ee S tu rdy 1 975 ; B ahn 1 983 , 1 984 ; G o rdon 1 988 ) . I n t h e se mode ls t h e d en se s t , mo s t a bundan t a nd mo s t p red ic tab le r e sou r ce ( e .g . , r e indee r ) i sa s sumed t ob e t h e m a ins tay o f t h e h uman e conomy .

V a r iou s g ene ra l mode ls a re b or rowed b y a r chaeo log is t s s t udy ing P a laeo l i th ic h un te r -ga the re r a dap ta t ion s . A rchaeo log ica l mode ls f o r t h e e a r ly Uppe r

A l l o f t h e a bove a r chaeo log ica l a s sump t ion sa r e t h e r e su l t o f al a ck o fm idd le r a nge t h eo ry b r idg ing t h eg ap b e twe en g ene ra l mode ls a nd a r chaeo log ica l

P a laeo l i th ic i nS ou thwe s te rn F rance f r equen t ly a s sume t h e v a l id i ty o f c oncep t s d rawn f r om g ene ra l t heo ry , e .g . ,

d a ta ( Be t t inge r 1 982 , 1 991 ) . T he b ody o f g ene ra l t h eo ry ( mode ls d ea l ing w i th t h e c u l tura l s y s tem ) a dd re s sed b y t h e

o p t ima l f o rag ing t h eo ry , e t hnoa r chaeo logy , a nd e t hno logy . T he se g ene ra l mode l sa r ea dap ted f o ru se b ya r chaeo log is t s , b u t h ave n o t b een t ho rough ly t e s ted i n t h e s t udy e nv i ronmen t .

a r chaeo log ica l mode l s i sb a sed o n e t hnog r aph ic d a ta . T he

" d rama t ic c hange s i n t h e c u l tu ra l r epe r to ire " ( R igaud & S imek 1 990 :84 ) o ccu r a t t h e g l a c ia l m ax imum ( s ee a l so Me l la r s 1 985 ; S of f e r 1 987 ; J och im 1 987 ) .

r e su l t ing s i tua t ion i so ne i n wh ich mode l s o f s oc ia l a nd t e r r i tor ia lo rgan isa t ion ,b a sed o nc e r ta in k ey c oncep t sd r awn f r om g ene ra l t h eo ry a r e n o t e a s i ly t e s ted u s ing a r chaeo log ica l d a ta .

Many a r chaeo log is t s ( e .g . , Wh i te 1 985 , 1 987 ; C onkey 1 980 ) , f o r e xamp le , a s sume t h ee x is tence o f af unc t iona ls e t t lemen tp a t te rn b a sed o ne t hnoa rchaeo log ica l o b se rva t ion s a mong t h e Nuna tn iu t ( B inf o rd 1 978 ) , a n I nu i t p eop le . Howeve r , t h is mode l h a s n o t b een f u l ly o pe ra t iona l i sed f o r t h e P a laeo l i th ic ( s ee C hap te r 6 ) .

1 .3 .0 S ea sona l I nd ica tor s a s aMean s o fT e s t ing Mode l s o f S e t t lemen t a nd S ub s is tence . S ea sona l i nd ica to r s a r e o ne o f t h e f ew t e s t s o f t h e p red ic t ion s g ene ra ted b y mode ls o f s e t t lemen t a nd s ub s i s tence . T he d egre e o f m ob i l i ty o f t h e s e t t lemen t s y s tem p red ic ted b yt h eh e rd -fo l low ing mode l , f o re xamp le ,

T he v a l id i ty o fc oncep t s s uch a s mob i l i ty

c an b e t e s ted a ga in s t s e a sona l p a t ter n s o f s i te o ccupa t ion . S ea sona l d i f f e rence s i n r e sou r ce u se a r e at e s t o f t h e d eg ree t ow h ich h uman s ub s is tence s y s tem sa r ec on s t ra ined b y s e a sona l e nv ironmen ta l c hange , i n o the r w o rd s , t h e d eg ree t o wh ich t h e s ub s is tence s y s tem c onform s t o

i nh un te r -ga the re r s y s tem s , a dap ted f r om t h e e t hnog raph ic mode l o f t h e c ompo s i te b and ( s ee d i scu ss ion C hap te r 6 , b e low ) , i so f ten t a ken f o rg ran ted i n mode ls o fP a laeo l i th ic a dap ta t ion s . T he c oncep t o fs oc io te r r i tor ia l mob i l i ty a s a n a spe c t o f h un te r -ga the re r a dap ta t ion s i s i nhe ren t i n a r chaeo log ica l mode l s o f s e a sona l r e s iden t ia l mob i l i ty , o r " s ea sona l r ound s " ( e .g . ,F reeman 1 973 ; S p ie s s 1 979 ; Wh i te 1 987 ) .

e xpec ta t ion s g ene ra ted b y o p t ima l f o rag ing t h eo ry a nd t h e c ompo s i te b and mode l . T he d eg ree o f s ea sona l e conom ic s pe c ia l isa t ion i n as ub s is tence s y s tem c an b e t e s ted u s ing s ea sona l d e te rm ina t ion s a nd e t ho log ica l i n f orma t ion j o in t ly w i th d a ta o n s i te l o ca t ion a nd s pa t ia l s t ruc tu re ( s ee C hap te r 6 ) .

A no the r , r e la ted a spe c t o f t h e c ompos i te b and mode l e ncoun te red u nde r v ar iou s g u ise s i n a r chaeo log ica l mode l s f o r t h e U ppe r P a laeo l i th ic i s t h e n o t ion o f f i s s ion a nd f u s ion . A s ea sona l p a t te rn o f

1 .4 .0 S e t t lemen t s y s tems : a rchaeo log ica l mode l s f or S ou thwe s tern F rance .

a gg rega t ion a nd d i spe r sa l o f t h e h uman p opu la t ion i s a s sumed t o e x is t i n a l l h un te r -ga the re r s y s tem s . T h i s a s sump t ion i mp l ie s al i nk b e tween r e s iden t ia l p a t te rn a nd s oc ia l o rgan i sa t ion ( Gamb le 1 986 ) , b a sed o n e t hnog raph ic

S ea sona l i ty e s t ima te sh ave i mp l ica t ion sf o r c ur ren t a r chaeo log ica l mode ls o fs e t t lemen ta nd s ub s is tence i n S ou thwe s ter n F rance ( s ee C hap te r 6 ) . A l l o f t h e se mode ls s h a re a c oncep t o f t h e s i te a s p a r t o f al a rge r ,

o b serva t ion . C onv inc ing p roof t h a t t h is r e s iden t ia l p a t te rn

2

r e g iona l s e t t lemen t s ys tem . Two b a s ic f o rm s o f e conom ic

t h a t as i ng le k i l l -popu la t ion ( whose p a rame te r s c an t h e ref ore

p a t te rn ing a r ep roposed f o rt h e Uppe rP a laeo l i th ic : r e indee r

b ek nown )i sr ef lec ted i nt h ea r chaeo log ica la s semb lage m ay

h e rd

s ea sona l

b e d i ff icu l t t os u s ta in . T he p re sence o f s e a sona l v a r ia t ion s

s e a sona l

i n p opu la t iona l p a rame te r s ( s uch a s v ar iab le a ge a nd s ex

f o l low ing ( e conom ic

r e s iden t ia l

s h if t s ,

o r

s pec ia l isa t ion ) a nd

r ound s ,

p a t terned

a f te r

s ubs is tence s h if t s . T he d egree o f s eden t ism i nv e r s ion s o f

d i s tr ibu t ion s ,

t h e

p ropose s

s ea sona l d e term ina t ion su s ing morphome tr ic d a ta . F ina l ly ,

g eog raph ica l ly l im i ted s ea sona l movemen t s ) . T he b a s ic

t e chn ique ss uch a st o o th w ea ra na ly s is h ave p roven d if i cu l t

d i f f e rence s b e tween t h e mode ls i nvo lve 1 ) t h e d egree o f

t o u se e ven when s t udy ing l i ve p opu la t ion s ( e .g . , K e lso 1 1

l a t te r mode l v ar ie s

( e .g . ,

J och im

1 983

e . g . ,

K e lso 1 1

1 968 )

f u r the r

c omp l ica te s

e conom ic d ependence o nr e indee ra nd 2 )t h er e la t ive d eg ree

1 968 ; M i l le r 1 974 ; S tu rdy 1 975 ; We lsh 1 975 ) . Un t i l v e ry

o f mob i l i ty i nt h e s y s tem . T he se mode ls g ene ra te d i f f e ren t

r e cen t ly , t h e refo re , a r chaeo log is t s h ave l a cked am ean s o f

e xpe c ta t ion sw i th r ega rd s t os ea sona lr e s iden t ia lp a t te rn ing .

a ccura te ly t e s t ing t h e s e a sona l e xpec ta t ion s o f s e t t lemen t a nd s ubs is tence mode ls f o r S ou thwe s tern F rance .

T he p a t te rn o fr e s idence p red ic ted f o r t h e h e rd f o l low ing mode l i s s ea sona l ly m a rked . r e s iden t ia l

p a t te rn ing

i s

e xpe c ted

t o

S ea sona l

c ovary

S ke le toch rono logy , mo re s pe c if ica l ly t h e

w i th

a na lys is o f g row th l a ye r s

i n d en ta l c emen t ,

p rov ide s

r e con s truc ted s e a sona l movemen t s o f t h e p r imary p rey

a r chaeo log is t s w i th t h e p o s s ib i l i ty o f o b ta in ing a b so lu te

s pec ie s i nt h eh e rd f o l low ing mode l . I n t h e s e a sona l r ound s

s e a sona l d e term ina t ion s f r om i nd iv idua l f o s s i l t e e th . T he

mode l , s e a sona l r e s iden t ia l s h if t s o ccur r ing w i th in a l o ca l

s t udy

r e g ion a r e n o t e xpec ted t o c ovary w i th t h e s e a sona l

i ndependan t o f p oss ib le s e a sona l b ia se s i n t h e s t ruc tu re o f

o f d en ta l

c emen t

h a s

t h e a dvan tage

o f b e ing

d i s tr ibu t ion o f as i ng le s pec ie s , b u t t op a t te rn a ccord ing t o

t h e p rey p opu la t ion . C emen tum a na lys is , t h e refo re , i s a

a more c omp lex r e source s chedu le . D e term ina t ion s o f

p owe rfu l t oo l f o r t h e t e s t ing o f s ub s is tence a nd s e t t lemen t

s e a son o f d ea th f o r a l l p rey s pec ie s t e s t t h ese p red ic t ion s .

mode ls i nt h e P a laeo l i th ic r e cord o fS ou thwes ter nF rance .

R esu l t s o b ta ined f r om t h is r e sea rch a nd f r om o the r s ou rce s a l low u s t ot e s t t h e p red ic t ion s o ft h ese mode ls . 1 .5 .0

Methodo logy

f or

Obta in ing

1 .6 .0 S pec ie s o fI n tere s t . S easona l

R e indee r p l ay a d om inan t r o le i n e a r ly

D e term ina t ion s f rom F auna l R ema in s .

Upper P a laeo l i th ic h uman e conom ie s i n t h e A qu i ta ine r eg ion ( s ee D e lpe ch 1 983 ) . D e ta i led s ke le toch rono log ica l

E a r ly s t ud ie s o fs ea sona l i nd ica tor s r e l ied

a na lys is o f moder n a nd f o ss i l r e indee r h a sb een u nder taken

o n morpho log ica l d i f f e rence s i n f a una f o r d e term in ing t h e

( Re ime r s & No rdby 1 968 ; M i l le r 1 974 ; G o rdon 1 988 ) . T he

s e a sona l i ty o fa r chaeo log ica l a s semb lage s . T h is a pproach i s

s e cond mos ti mpo r tan tp rey s pec ie si nt h eh uman e conom ie s

b a sed

a n t le r

o fS ou thwe s tern F rance d ur ing D rya s 1i sh or se . S i nce o ne

morpho logy , a nd a ge -dependen t v ar ia t ion i n d en ta l w ea r

o n

d e term in ing

s ea sona l

d i f f e rence s

i n

o f t h e d i s t ingu ish ing f e a tu re s o f t h e h uman s ub s is tence

s u rf ace s , c rown h e igh t r educ t ion , e p iphysea l f u s ion a nd

s y s tem d ur ing t h ep er iod o fi n te re s t i s ar e la t ive i n c rea se i n

d en ta le r up t ion p a t te rn s . S ea sona l morpho log ica ld i f f e rence s

f a una l d ive r s i ty ( Me l la r s 1 985 ; J och im 1 987 ; R igaud &

p rov ide d i re c t e v idence o f s ea son o f d ea th . Age d e term ina t ion s b a sed o n morpho log ica l c ha rac ter is t ic s ,

o ther s pec ie s i n to ad i scus s ion o fs e a sona l i nd ica to r s .

S imek 1 990 ) , t h e p re sen t r e sea rch i si n tended t o i n c lude

c omb ined w i th p opu la t ion p arame te r s f o rt h es pec ie ss t ud ied ( e .g . , s e a son o fb i r th ) , a l so p rov ide e v idence o f s ea son o f

1 .7 .0 S ke le tochrono log ica lS tud ie s : C emen tu rn Ana ly s is .

d ea th . I n o rde r t oa pp ly c emen tum a na ly s is t oa S ea sona l

d a ta

f o r

r e inde e r

o n

t a rge t s pe c ie s , ad e ta i led h i s to log ica l s t udy a nd c a refu l

morpho logy a r e d i scus sed f o r t h e Uppe r P a laeo l i th ic i nt h e

a na lys is o f t h e n a ture o f t h e g row th s t ruc ture s o b served ,

A qu i ta ine r eg ion ( e .g . , C heyn ie r e t a l .

b a sed

1 952 ; B ouchud

i n c lud ing c ompar ison w i th g row th s t ruc ture s o bserved i n

1 966 ; L acor re 1 956 ) . T he a u thor s u se t h e se s e a sona l d e term ina t ions t op red ic th uman r e s idence p a t terns . I ns ome

o ther s pe c ie s , mus t b e u nder taken ( s ee C hap te r 2 ) . T h is

c a se s , h oweve r , c on f l ic t ing mode ls a r e s uppor ted u s ing t h e

d e term ina t ions b a sed o n i n cor rec t i d en t if ica t ion o fp r ima ry

s ame f a una l d a ta ( e .g . , f o rB adegou le , c on t ra s t C heyn ie re t

g row th

a L 1 952 a nd L acor re 1 956 ) . T h is c onfu s ion i sl a rge ly d ue

c ompar ison s w i th o the r s pec ie s .

t o

m e thodo log ica l

p rob lem s

i nhe ren t

i n

t h e

u se

m in im ise s t h e p o ss ib i l i ty o f p roduc ing e r roneou s s ea sona l l a ye r s

o r

f a lse

a s sump t ions

b a sed

o n

h a s ty

o f

morpho logy ,w h ich h ave h ampe red b o th t h ee l abora t ion a nd

B efore u nder tak ing s uch a p rog ram o f

t e s t ing o fs e t t lemen t mode ls .

r e sea rch , i ti sn ece s sa ry f o rt h e a r chaeozoo log is tt ob ecome a cqua in ted w i th t h e l a rge b ody o f l i t e ra tu re c once rn ing

T aphonom ic

h uman

m ine ra l ized t i ssue s . Much c onfus ion h a s a r isen c oncer n ing

p roce s s ing a c t iv i t ie s m ay d i f f e ren t ia l ly a f f e c t t h e v ar iou s

t h e u se o fc emen tum a na ly s is i na r chaeo logy a s ar e su l t o f

c l a sse s

o f

morpho log ica l

Morpho log ica l d a ta

c r ea te

p roce s se s d a ta

( s ee

o ve r lapp ing

a nd C h .4 , a ge

b e low ) .

t h eu se o fd i f f e ren to bserva t ion t e chn ique sa nd n on s tanda rd

c a tegor ie s

t e rm ino logy .

A n

a t temp t

i s m ade

t o s t anda rd ize

t h e

( Sp ie s s 1 979 ) r e su l t ing i nr e la t ive r a the r t h an a bso lu te a ge

l a nguage u sed a nd e s tab l ish a s t anda rd p rocedu re f o r t h e

d e term ina t ions . T h is r equ ire s t h a t ap opu la t ion b e t h eo b jec t

o bserva t ion o fg row th l a ye r s i nc emen t . V ar iou s t e chn ique s

o f s t udy , r a the r t h an i s o la ted r ema in s . T he d e term ina t ion

o f p o ten t ia l i n te re s t t o a r chaeo log is t s s t udy ing c emen tum

3

I nhe ren t p rob lem s w i th t h e m e thodo logy a r e d e l inea ted . A

g row th l a ye r s a re c ompa red ( s ee C hap te r s 2& 3 ) .

p rac t ica l a pp l ica t ion o f t h i s a na ly s i s o n f o s s i l h o rse p opu la t ion si se xp lo red , e xpand ing t h ed a tabase o fs e asona l

B e cau se t h e p a t te rn o f d epo s i t ion o f g row th l a ye r si nc emen ti se s sen t ia l ly e ndogen ic ( g ene t ica l ly d e te rm ined ) , t hough t r igge red b y e xogenou s ( e x te rna l )

i n f orma t ion f o rt h el a te Uppe rP a laeo l i th ic i nt h eS ou thwe s t o fF rance .

f a c tor s s uch a s s e a sona l e nv ironmen ta l c hange ( s ee d i scu s s ion , C hap te r 2 ) , ac on t ro l s amp le f o r t h e t a rge t s pe c ie s mu s t b e a s semb led . T he p u rpo se o f t h e c on t ro l s amp le i st od e term ine t h e t im ing o f t h e d epo s i t ion o f t h e d i f f e ren t c emen tum g row th l a ye r s . T he c on t ro l g roup c on s is t s o fa n ima ls o fk nown s e a son o fd ea th . F o ro bv iou s r ea son s , t h e refore , t h e c on t ro l s amp le c on s is t s o f mode rn s pe c imen s o ft h e t a rge t s pe c ie s . T he a pp l ica t ion o f am ode rn c on t ro lg roup t o af o s s i lp opu la t ion r e s t s o n t h ea s sump t ion o f ad egre eo f g ene t ic u n ifo rm i ty ( s ee C hap te r 2 ) , i mp ly ing t h a t n o c hange s i nt h e e ndogenou sp a t te rn o fc emen tum d epo s i t ion h ave o ccu r red . C on t ro l a n ima ls a r e o b ta ined f r om e nv ironmen t s w i th r ough ly s im i la r p a t te rn s o fs e a sona l i ty , t hu se l im ina t ing ap os s ib le e xogenou ss ou rce o fd if e ren t ia l e xpre s s ion o f t h e g row th l a ye r s . F o r t h e p u rpo se s o f t h is r e sea r ch , l a t i tud ina lv a r ia t ion i sc on s ide red t h ed e te rm in ing f a c to r ( i .e . , at empe ra te z one c on t ro l i s e s tab l ished f o r e xam in ing f o s s i l p opu la t ion s f r om t empe ra te z one s ) . 1 .8 .0 The A rchaeo log ica l S amp le . T he a r chaeo log ica l s amp le f o rt h ep re sen t r e sea rch o r ig ina te s f r om t we lve s i te s i n S ou thwe s te rn F rance c on ta in ing B adegou l ian a nd L owe r Magda len ian o ccupa t ion l e ve ls . E x is t ing c o l le c t ion s , f i rm ly a t t r ibu tab le t oD rya s 1( be tween 1 8 a nd 1 4 K yr s B .P .) o n t h e b a s i s o f c h rono s t ra t igraph ic i n f orma t ion , w e re s amp led . T he s amp led s i te s c ome f r om a v a r ie ty o f l o ca l e nv i ronmen t s w i th in t h e s t udy r eg ion . T he p re sence o f h o r se i n a n a s semb lage i s ap re requ is i te f o r i n c lu s ion i n to t h e s amp le p opu la t ion , i n o rde r t o t e s t mode ls o f e conom ic s pec ia l isa t ion b a sed o nr e indee r . T he s amp le a l so r e f le c t sa n a t temp t t o i n c lude s i te s w i th mo re t h an o ne s ea sona l i nd ica to r , i n o rde r t o c on s t ruc t a n a rgumen t b ea r ing o n s e t t lemen ta nd s ub s is tence s ys tem s . A l l o ft h e s i te s i nt h e s amp le a r e c ave o r r ock she l te r l o ca t ion s . T h is i mpo se s r e cogn ised l im i ta t ion s o n t h e p re sen t r e sea rch , s i nce r eg iona l s t ud ie s h ave s hown t h a tn ume rou s o the r s i te f o rm s e x is t i nt h e A qu i ta ine B a s in ( s ee Wh i te 1 987 ; R igaud & S imek 1 987 :56 ) . Howeve r , t h i s l o ca t iona l b ia s r ef le c t s t h e r e a l i t ie s o ff a una l p re serva t ion . R e su l t so ft h ep re sen ts t udy a r e c omb ined w i th o t he ri nd ice s o fs e a sona l i ty i nt h ed i scu s s ion o fp a t te rn ing i nt h es e a sona l d a ta ( Chap te r 4 ) . 1 .9 .0 C onc lus ion s . T he p u rpo se o ft h is r e sea r ch , t h e refo re , i s t wofo ld . 1 )T he i mp l ica t ion so fs ea sona l d e term ina t ion s f o r h o r se a nd r e indee r , o b ta ined f r om c emen tum a na ly se s , f o r e x is t ing mode l s o f s ub s is tence a nd s e t t lemen t i n t h e S ou thwe s t o f F rance a r e d rawn . 2 ) T he a pp l ica t ion o f c emen tum a na lys is t o an ew s pe c ie s , E . c aba l lu s , i st e s ted . 4

CHAPTER 2 . THE SKELETOCHRONOLOGY OF HORSE TEETH 2 .0 I n troduc t ion .

d e scr ibed a s at wo -pha se p roce ss : p roduc t ion o ft h eo rgan ic m a t r ix

T h is C hap te r s e rve s a s a n i n t roduc t ion t o

p recede s

m ine ra l iza t ion

( e .g . ,

S imk iss

1 975 ;

t h es t udy o fm ine ra l iza t ion a nd g row th l a ye r s i nm ine ra l ized

G l imche r 1 976 ; C a s tane t 1 982 ) . A d escr ip t ion o f t h e p roce s s o f f o rma t ion o f t h e o rgan ic m a tr ix , wh ich i s

t i ssues .T he r e levan tb io log ica ll i te ra tu re c oncern ing g row th

s e c re ted b y o s teob la s t so n t h e g row ing s urf ace o ft h e b one ,

s t ruc tu re s i nb one a nd c emen t i sr ev iewed a s an ece ssary

i sf ound i nJ one s e ta l . ( 1985 ) . Minera l iza t ion i st h e r e su l t

b ackground f o rt h e a r chaeozoo log is t i n te re s ted i nt h e s t udy

o f c omp lex h i s tochem ica l r e ac t ion s , i nvo lv ing h ormona l ,

o fd en ta l c emen t . T he r ange o ft e chn ique s u sed t oo b serve

b lood -ca lc ium a nd e nzyme l e ve ls ( S imk is s 1 975 ) l i nked t o

g row th l a ye r s i nb one a nd c emen t i sa l so r e v iewed .

e ndocr ine s e c re t ion s (War ren 1 963 ) . T he o rgan ic ma tr ix

PART A : HARD T ISSUES .

c r y s ta l f o rma t ion ( s o l id p ha se g row th ) i s t hough t t o b e

f o rm s ag row th s u rf ace f o r t h e s o l id p ha se . T he r a te o f r egu la ted b y t h ep a ra thyro id t h rough a d ju s temen t si nb lood 2 .1 B one a nd C emen tum : S truc ture a nd

c a lc ium l e ve ls ( S imk iss 1 975 ) . B lood i ss uper sa tu ra ted w i th

c ompo s i t ion .

c a lc ium a nd o the rs o l id p ha se c omponen t s wh ich p rec ip i ta te t o f o rm b one c rys ta ls . E nzyme l e ve ls i n t he b lood a r e C emen tum i s am ine ra l iz ed t i ssue c l ose ly

r e la ted t o b one ( Franc i l lon -V ie i l lo t e t a l .

t h ough t t o a c t a s i nh ib i to r s t o c rys ta l f o rma t ion a nd t hu s

1 991 :474 ; d e

c on t ro l t h e r a te o fb one g row th ( S imk is s 1 975 ) .

R icq ls e ta l . 1 992 :8 ) . B one a nd c emen tum a r e c omposed o f a pprox ima te ly 6 5% i n organ ic c omponen t s ( c a lc ium p ho spha te ) a nd a bou t 3 5% o rgan ic c omponen t s ,

Wh i le n ume rou s h ypo these s h ave b een

w i th

p ropo sed t o e xp la in t h e p roce ss o f b one m ine ra l iza t ion

r e la t ive ly f ew c e l ls p e r v o lume /ma s s o f m a tu re t i ssue

( e .g . , G l imche r 1 976 ; S imk iss 1 975 ) , t h e se p roce sse s a r e

( S imk iss 1 975 ; G l imche r 1 976 ; C a s tane t 1 982 ) . C e l ls i n c emen tob la s t s

s t i l l i mprope r ly u nde r s tood t oday ( Boyde 1 980 ) . T he i mpor tan t f e a ture o f m ine ra l iza t ion i st h a t t he s o l id -pha se

embedded i n t h e t i ssue ( S chm id t & K e i l 1 972 :202 ) . T he

t r an sforma t ion i s n o t a ccomp l ished i n as i ng le s t ep . A n

o rgan ic ma tr ix wh ich f o rm s t h eb a s ic s t ruc tu ra l c omponen t o fc a lc if ied t i s sue s i sc ompo sed o fc o l lagen f i b re s ( S imk is s

i n i t ia l s o l id p ha se i s d epo s i ted a s p oor ly c ry ta l l ine h yd roxyapa t i te m ine ra ls . As e r ie so ff u r the rt r ansforma t ion s

1 975 ; G l imche r 1 976 ) . S ome 9 5% o f t h e e x t race l lu la r

r e su l t s i nt h e m a tu re c rys ta l l ine p hase ( G l imche r 1 976 ) . I n

o rgan ic ma tr ix i sc o l lagenou sp ro te in ( G l im che r 1 976 ) . T he n on -co l lagenou s c omponen t s a r e h igh ly v a r iab le i nq ua l i ty

o t he r word s , t h e d egree o fm ine ra l iza t ion i nb one t i s sue i s n e i ther a na b so lu te n or ac on s tan tr e la t ionsh ip . O lder t i s sue s

a nd q uan t i ty a st h ey a r ea l te red o rl o s td ur ing m ine ra l iza t ion

w i l l t e nd t o b e r e la t ive ly more m inera l ized t han y ounge r

( G l imche r 1 976 ) . T he s o l id p ha se o fb one a nd c emen t ( i .e . ,

t i s sue s , r ega rd le s s o f t h e o rgan isa t ion o f t h e b one ma t r ix

c emen t ,

t h e

c emen tocy te s ,

m inera l

p ha se )

a r e

i s

s e cre ted

b y

p r inc ipa l ly

c omposed

o f

( Ca s tane t 1 982 ; R icq ls e ta l . 1 992 :18 ) . T h is p henomenon

h ydroxyapa t i te c rys ta ls , af o rm o fc a lc ium p hospha te , i . e . ,

w i l la f fe c tt h es t udy o fi n cremen ta l s t ruc ture s i nc emen t a nd

C a , o( PO 4) , (OH ) , ( H i l lson 1 986 ) .

i sc r uc ia l t o wha t f o l low s .

T he re e x is t b a s ic f e a tu re s c o mmon t oa l l

B one a nd c emen tum s hou ld n o tb es een a s

c o l lagen s , b u t t h e re a r e s i gn if ican t d i f f e rence s b e twe en

i n er to r " dead " s t ruc ture s . T he se t i ssue sa r e , i nf a c t , p ar to f

m inera l ized ( bone ) a nd n on -m ine ra l ised c o l lagen . B one

l i v ing o rgan s w i th a c on t inued d ynam ism , e ven when

c o l lagen p os se sse s au n ique s t ruc tu re a nd h i s tochem is try ( G l imche r 1 976 :84 )a nd p l ays a ni mpor tan tr o le a s ag row th

m a tu re . B one i st hough t t oa c t a s a" r eservo ir " o fe s sen t ia l i o n s ( G l imche r 1 976 ) . B one r emode l ing , wh ich c on t inue s t h roughou ta no rgan ism 's l i f e t ime , p lays ar o le i nr egu la t ing

s u rf ace f o r c r ys ta ls ( t h rough ap roce s s k nown a s e p i taxy , s e e S imk iss 1 975 ; a nd a s an uc lea t ion c a ta lys t f o r c ry s ta ls , s e e G l imche r 1 976 :60 ) . A t t h e mo le cu la r l e ve l , b one

b lood c a lc ium l e ve ls a l though b one i t se lf i sn o t i n d i r ec t c on tac t w i th t h e c i rcu la tory s y s tem b u t i s med ia ted b y t h e

c o l lagen i s s t ruc tura l ly o rgan ised i n to f i br i ls . F ibr i ls a r e

b one m embrane ( S imk is s 1 975 ) . T he e xac t n a ture o f t h is

o rgan ised

m ed ia t ion i sn o t y e t p e rfe c t ly u nder s tood .

i n

a h igh ly

s pe c if i c

h exagona l

p ack ing

a r rangemen t i n to l a rge r s t ruc tu ra l u n i t sk nown a sf i bre sa nd i n to e ven l a rge r u n i t s k nown a s f i b re b und le s ( G l imche r

Dur ing t h e g row th o f t h e o rgan ism b one

1 976 ) . B e cau se o ft h e ir p ack ing a r rangemen t , b one c o l lagen f i br i ls c on t ro l t h e g row th r a te o f t h e s o l id p ha se ( S imk is s

i sr e s t ruc tu red , t a k ing o n i t sm a tu re f o rm . T h is g row th m ay b e a symme tr ica l ( a x ia l ly a nd l ong i tud ina l ly ) . O nce t h e

1 975 ;

s ke le ton h a s a ch ieved i t s m a tu re f o rm , h oweve r , g row th

G l imche r

1 976 ) a nd ,

u l t ima te ly , t h e d eg ree o f

m ine ra l iza t ion ( G l imche r 1 976 ) .

d oe s n o t c ea se b u t i sa c t iva ted b y f unc t iona l c on s t ra in t s

2 .2 M inera l iza t ion

m ine ra l i o n l e ve ls i n t h e b lood i s a ccomp l ished t h rough

( G l imche r

1 976 ) .

F ur thermo re ,

i t s r o le

c on t inued b one " t ur nove r " o r r emode l ing . B one d epos i t ion c an

b e mos t

s imp ly 5

i n r egu la t ing

2 .3 C o l lagen o rgan i sa t ion a nd t ype s o fb one .

d epo s i ted l a ye r s h ave a l owe r c e l l space v o lume f r a c t ion

T h re e m a in t ype s o fb one m a t r ix c an b e d i s t ingu ished o nt h eb a s is o ft h eo rgan isa t ion o ft h ec o l lagen

t h an l a ye r s d epo s i ted a t af a s te r r a te , w i th ar e su l tan tl owe r d en s i ty ( Ca s tane t 1 982 ; B oyde 1 987 ) . T h i s d i s t inc t ion w i l l b ei mpor tan t when c on s ide r ing t h en a tu r e o fm ine ra l iza t ion p a t te rn s i nc emen t ( b e low ) .

f i b re s c ompo s ing t h em : w oven b one ; l ame l la r b one ; a nd p a ra l le l -f ibe red b one ( Pr i tcha rd 1 956 ; E n low & B rown 1 956 :7 -8 ; C a s tane t 1 982 :31 -37 ; F r anc i l lon -V ie i l lo t 1 991 :509 -510 ;d eR iq le se ta l . 1 991 ) . Woven b one h a sp oo r

I nt h e c a se o fh o r se c emen tum , e xam ined d u r ing t h e c ou r se o f t h is r e sea rch , c em en tum l a ye r s o bserved c on ta in ed l a cunae ( e v idence o f c emen tocy te s )

s pa t ia lo rgan isa t ion a nd c on ta in sn ume rou so s teocy te s wh ich a r e v o lum inou s , g lobu la r a nd h ave d eve loped c y top la sm ic e x ten s ion s . Woven b one c an b e mo re d en se ly m ine ra l iz ed t h an l ame l la rb one . L ame l la r b one i s mo re h igh ly s pa t ia l ly o rgan ised , c ompo sed o f s ucce s s ive l a ye r s o f c o l lagen f i b r i ls , w i th amo re o rde r ly a r rangemen t o fo s teocy te s t h an w oven b one . P a ra l le lf i be red b one ( f ound a tt h el e ve l o ft h e

r ega rd le s s o f t h e i r l o ca t ion o n t h e t o o th . T he d i s t inc t ion b e tween a ce l lu la ra nd c e l lu la rc emen t m ay t h e re fo re n o tb e r e levan t f o ra l l s pec ie s . A s ar e su l t , t h ep r inc ipa lf unc t iona l a s se t o fc emen t wh ich c once rn s u s h e re i st h e o rgan isa t ion o f t h e c o l lagen f i b re m a t r ix t he s ou rce o fo b se rved m a jo r s t ruc tu ra l v a r ia t ion s ( O sbo rn 1 981 :198 ) .

c or t ica l d i aphy se s ) r ep re sen t s a n i n te rmed ia te l e ve l o f o rgan isa t ion .

PART B I ncremen ta l G row th S truc ture s 2 .4 T erm ino logy

S eve ra l d i f f e ren tt ype so fc emen tum h ave a l so b een d i s t ingu i shed o n t h e b a s i s o f c o l lagen f i b re o rgan isa t ion a nd o r ien ta t ion . S chm id t a nd K e i l ( 1972 :298 ) d i s t ingu ish b e twe en " l a t t ice c emen tum " ( e qu iva len t t o w oven b one ) a nd " o s teocemen tum " ( e qu iva len t t op a ra l le l f i be red b one ) . O sbo rn ( 1981 ) d i s t ingu i she s f ou r s epa r a te t y pe s o fc emen tum : a f ib r i l la r c emen t ( r a re ) ; e x t r in s ic f i b re c emen t ; i n t r in s ic f i b re c emen t a nd m ixed f i b re c emen t . T he se t e rm s r e f e r t o t h e o r ig in o f t h e c o l lagen f i b re s m ak ing u p t h e m a t r ix o f t h e c emen t . E x t r in s ic f i b re s ( s ome t ime s r e f e red t o a s S ha rpey 's f i b re s ) a r e i n c luded

T he l i te ra tu re c once rn ing g row th s t ruc tu re s i nc emen t c an b e c onf u s ing ( e spe c ia l ly t h e n on s pe c ia l is t l i t e ra tu re ) d ue t o t h e l a ck o f s t anda rd ised d ef in i t ion s f o r t h e o b served s t ruc tu re s ( no tab ly LAG s a nd a nnu l i )a nd t h ef a c tt h a ta u tho r sd on o ta lway sm ake e xp l ic i t wh ich c l a s s if ica tory s y s tem t h ey a r e u s ing t od i f f e ren t ia te t ype s o fc emen t . I nt h ef o l low ing d i s cu s s ion ,g row th l a ye r s w i l l r e f e r t o a ny l a ye r ing i n c em en t o r p r ima ry c or t ica l b one wh ich i st h e r e su l t o fa ppo s i t iona l ( s ome t ime s t e rmed i n c remen ta l ) g row th ( Monk s 1 981 ) . T he t e rm g row th z one s w i l l d e s igna te t h e r e la t ive ly t h ick , o paque l a ye r s o f

f r om t h ep e r iodon ta ll i gamen t( Boyde & J one s 1 972 ; B oyde 1 980 ; O sborn 1 981 ) . I n t r in s ic f i b re s a r e p roduced b y c emen tob la s t s( O sbo rn 1 981 ; B oyde 1 987 ) .I n t r in s ic c emen t i sf ound i nt h e a p ica l r eg ion s o fo lde r t e e th , p l ay s n o p a r t i n t h e s uppor t ing s y s tem o f t h e t oo th a nd m ay b e o f a t r an s i to ry n a tu re ( O sbo rn 1 981 :203 ) . E x tr in s ic c emen t i s c on s ide red " p r ima ry " , i . e . , f o rmed p r io r t o f unc t iona l o cc lu s ion , s l ow ly a nd e ven ly f o rmed a nd w e l l m ine ra l iz ed ( O sborn 1 981 :198 ) . M ixed f i b re c emen t ( c on ta in ing b o th e x t r in s ic a nd i n t r in s ic f i b re s , O sbo rn 1 981 :199 ) c an b e c on s ide red t o f o rm t h e b u lk o f t h e c emen tum d epo s i t . R e la t ive p ropor t ion so fi n t r in s ic a nd e x t r in s ic f i bre s wou ld f o rm t h e m a jo r d i s t inc t ion b e twe en g row th z one s a nd a nnu l i /LAG ' , a cco rd ing t oB oyde ( 1987 ) .

" s umme r " o r a c t ive b one a nd c emen t g row th ( Peabody 1 958 , 1 961 ; C a s tane t 1 981 , 1 982 ; B ag l in ie re e ta l . 1 990 ) wh ich a r e i s o t rop ic ( Ca s tane t 1 981 , 1 982 ) . A nnu l ir e f e r s t o t h e n a r rowe r , a n iso t rop ic ( Ca s tane t 1 981 , 1 982 ) b and s o f r e la t ive ly t r an s lu scen t c emen t ( Bag l in ie re e t a l . 1 990 ) d e scr ibed b y P eabody ( 1958 , 1 961 ) , Wa r ren ( 1963 ) a nd C a s tane t ( 1981 , 1 982 ) . L ine s o f A r re s ted G row th ( LAGs ) d e s igna te t h in , c h romoph i le d i scon t inu i t ie s ,o r " r e s t " l i ne s , i n c emen t , s ome t ime s s i tua ted w i th in a nnu l i ( when t h es e l a t te ra r ep re sen t ) , s ome t ime sa l te rna t ing w i th g row th z one s when a nnu l i a r e n o t m a rked o r a r e a b sen t ( Ca s tane t 1 979 , 1 981 , 1 982 ; B ag l in ie re e t a l . 1 990 ) . LAG s r ep re sen t " i nunda ted s u rf ace r eg ion s " (Wa r ren 1 963 :25 ) a nd c o r re spond t o t h e " adhe s ion l i ne s " o f K leveza l a nd

S ome c onfus ion a r ises i n t he n omenc la tu re , s i nce a v a r ie ty o f c r i te r ia a r e u sed t o d i s t ingu ish d i f f e ren t t y pe s o fc emen t ( s e e d e R icq le s e ta l . 1 992 :9 ; F ranc i l lon -V ie i l lo t 1 991 :499 ) . F o r e xamp le , c e l lu la r c on ten t i su sed a s ac r i ter ion f o r d i s t ingu ish ing

K le inenbe rg ( 1967 ) , t h e " i ncremen ta lg row th l i ne s " o fG rue & J en sen ( 1979 ) , a nd t h e " r e s t l i ne s " o f K leveza l & Sm ir ina ( 1969 ; a s q uo ted i n C a s tane t 1 982 ) . ( See F igu re 2 .1 ) .

b e tween a ce l lu la r a nd c e l lu la r c emen t ( S chm id t & K e i l 1 972 ; B oyde & J one s 1 972 ; B oyde 1 987 ) . I n s ome s t ud ie s , a ce l lu la r a nd c e l lu la r c emen t a r e d e sc r ibed a s a l te rna t ing s t ruc tu re s ( S chm id t &

A n o paque b ody ( e .g . , g row th z one s i n c emen t ) w i l l a ppea r b r igh t ( wh i te ) u nde r r ef le c ted n a tu ra l l i gh t a nd d a rk ( b lack ) u nde r t r an sm i t ted n a tu ra l l i gh t . A t r ans lucen t b ody ( e .g . , a nnu l i i nc emen t ) w i l l h ave i n ve r se p roper t ie s , a ppea r ing d a rk u nde r r e f le c ted n a tu r a l l i gh ta nd

K e i l 1 972 ) . O sbo rn ( 1981 ) , h oweve r , l im i t st h ed i s tr ibu t ion o fp r ima ry ( i .e . , a ce l lu la r ) c emen t t o at h in l a ye rd epo s i ted p r io rt om a tu ra t ion o ft h et oo th . O b se rva t ion so fa l te rna t ing l a ye r s o f wha t a r e d e sc r ibed a s " c e l lu la r " a nd " a ce l lu la r " c emen t m ay b e ar e su l t o f t h e f a c t t h a t c emen tum i s

b r igh t u nde r t r an sm i t ted l i gh t . LAGs a ppea r b r igh t u nde r o rd ina ry t r an sm i t ted l i gh t a nd h igh ly b i re f r ingen t u nde r p o lar ised l i gh t( Bag l in ie re e ta l . 1 990 ; F ranc i l lon -V ie i l lo te t

d epo s i ted i ns ucce s s ive l a ye r s a t av a r iab le r a te ( d e R icq le s e t a l . 1 992 ; F ranc i l lon -V ie i l lo t 1 991 ) . Mo re s l ow ly

a l . 1 991) . 6

F igure 2 . 1 S pa t ia lR e la t ionsh ip B e tween Annu l i , LAG s a nd Grow th Z one s i n Equ id Den ta l C emen tum .

T ransver se s ec t ion o f mand ibu lar mo lar o fh or se : ANN . =a nnu lus . LAG = l i gn o fa rre s ted g row th . PER = p er iodon ta ll i gamen t . N o te t he p re sence o fp er iodon ta l ( Sharpey 's ) f i bre s p ene tra t ing t he t oo th 's s urface . M AND . = - - mand ibu lar b one . j u . =c e men tu m j unc t ion . L a . =l a cuna ( c emen tocy te ) .

7

2 .5 Na ture o ft he g row th s t ruc ture s .

c oncen t ra t ion s o f m inu te p o re s pace s ( r e la ted t o c o l lagen p ack ing ) a nd /o r b ounda r ie s b e twe en m ed ia w i th d if e ren t r e f rac t ion i ndexe s . Unde r t r an sm i t ted o rd ina ry l i gh t , d i f f e rence s i nt h e o r ien ta t ion o fc o l lagen f i br i l s w i l la ppea r a s a l te rna t ing b r igh t a nd d a rk l a ye r s ( H i l lson 1 986 :173 ) .

S eve ra l d if e ren t i n ve s t iga t ive m e thod s h ave b een u sed t o o bse rve g row th s t ruc tu re s i nb one a nd c emen t . A s u mma ry o f e ach o f t h e se me thod s i sg iven b e low , f o l lowed b y a d i scu s s ion o f t he c onc lu s ion s r e ga rd ing t h e n a tu re o f g row th l i ne s w h ich c an b e d rawn

G row th z one s , w h ich a r e o paque b od ie s , a ppea r a s r e la t ive ly d a rk l a ye r s wh i le a nnu l i a ppea r r e la t ive ly b r igh t ( Franc i l lon -V ie i l lo te ta l . 1 991 :508 ) .

f r om t h em . 2 .6 .0 T e chn ique s f or E xam in ing G row th S truc ture s .

P o l ished c emen tum s u rf ace sv i ewed u nde r o rd inar yr e f le c ted l i gh tr evea l ap a t te rn o fa l ter na t ing b r igh t b and s c o r re spond ing t o g row th z one s a nd d a rk b and s ,

2 .6 .1 P o lar ised l i gh t :

c or re spond ing t o a nnu l i /LAGs ( Franc i l lon -V ie i l lo t e t a l . 1 991 ) .

E xam ina t ion o fu nde ca lc if ied ,g round t h in s e c t ion s u nde rp o la r ised l i gh ti sa n e xce l len t t e chn ique f o r I nve s t iga t ing t he s t ruc tu ra l a nd h i s to log ica l d if e rence s b e tween g row th l a ye r s i nc emen t a nd b one , wh ich a ppea r a s c on t ra s t ing p a t te rn s o f d a rk a nd l i gh t a cco rd ing t o t h e

2 .6 .3 S ta ined , H i s to log ica l Th in S ec t ion s . D em ine ra l ised , s t a ined m icro tomed ( f roz en )t h in s e c t ion s , moun ted i na na queou sm ed ium ,h ave a l so b een u sed t oi n ve s t iga te b and ing i nb one a nd c emen t . When s t a ined w i th E h r l ich 's h ema toxy l in , f o r e xamp le ,

o r ien ta t ion o ft he f i br i l la ra nd m ine ra lc omponen t( Peabody 1 961 ; S chm id t & K e i l 1 972 ; Myr ick 1 980 ; O sbo rn 1 981 ) . S ince f o s s i l ma te r ia l c anno t b e d eca lc if ied , i ti sa l so o ne o f t h e f ew me thod s o fo b se rv ing g row th l a ye r s i nf o s s i l t e e th . E a r ly o b serva t ion so fg row th s t ruc tu re si nb one a nd c emen t

c emen t a nd p r ima ry c or t ica l b one s how a l te rna te b and s o f w ide , p oo r ly s t a ined t i s sue a nd t h in , d a rk ly s t a ined t i s sue

i n c lude d e scr ip t ion sb y C ape & K i tch in ( 1930 , a sq uo ted i n S chm id t&K e i l 1 972 , a nd c . f . K e i l 1 954 , I b id .) w ho s t a te

( Re ime r s & N o rdby 1 968 ; K leveza l 1 970 , 1 980 ; G rue & J ensen 1 979 ) . T he t h in , c h romoph i le b and s c or re spond t o a nnu l i o r , mo re o f ten , t o LAGs ( Boyde 1 980 ; C a s tane t 1 981 , 1 982 ) .I ns t a ined ,d eca lc if ied s e c t ion s ,t h ei mpo r tance o f t h e o r ien ta t ion o f t h e f i b re ma t r ix i n d e term in ing t h e c ha ra c te r i s t ic so ft h ei n c remen ti sa ppa ren t( Ca s tane t 1 982 ) . A n umbe r o f d i f f e ren t s t a in ing a gen t s , e .g . , Maye r 's

t h a t c emen t c on s is t s o fa l te rna te l a ye r s o fb i ref r ingen t a nd i s o t rop ic t i s sue u nde rp o la r ised l i gh t . I mage sa r ep roduced u nde rp o la r i sed l i gh t t h rough d oub le r e f rac t ion ( i .e . , b i re f r ingence ) . L igh t i s r e f rac ted i n to t wo e me rgen t b eam s wh ich , i n t h e c a se o f a pa t i te c rys ta ls a nd c o l lagen f i b re s , a r eo r ien ta ted 1 )p a ra l le l t ot h el ong a x is o ft h ec o l lagen f i b re ( S chm id t&K e i l 1 972 ; H i l lson 1 986 ) a nd 2 ) p e rpend icu la r t o t ha t a x i s ( H i l lson 1 986 ) . T he d i f f e rence i nr e f rac t ive i n dexe s b e twe en t h e se t wo b eam s i sk nown a s b i ref r ingence a nd h a s ap o s i t ive v a lue f o r c o l lagen f i b r i ls ( H i l lson 1 986 ) a nd t h e o ppos i te s i gn f o r a pa t i te ( Schm id t&K e i l 1 972 ) .

h ema lun ( Grue & J en sen 1 979 ; A r to i s&S a lmon 1 981 )a nd s i lve r n i t ra te ( d 'Hua r t 1 980 ) , h ave b een u sed t o p roduce d i ch rom i sm i n c emen t a nd b one , h igh l igh t ing t h e h i s tochem ica l d i f f e rence s b e tween a nnu l i a nd g row th z one s i nt h e se t i s sue s ( S chm id t&K e i l 1 972 ; C a s tane t 1 981 ) . 2 .6 .4 T ran sm i t ted E lec tron Micro scope ( TE M ) . T he re h ave n o tb e en m any s pe c if ic s t ud ie s

Unde ca lc if ied p o la r i sed t h in s e c t ion s o f c emen t s how al a ye red s t ruc tu re o fd i f f e ren tp a th d en s i t ie s ,

o fg row th l a ye r s i nc emen tc a r r ied o u tu s ing t h et r an sm i t ted e l e c t ron m ic ros cope . U nde r TE M , s em i th in p repa ra t ion so f p r ima ry c or t ica l b one s how a l te rna t ing e l e c t ron -opaque a nd n on -opaque b and ing c or re spond ing t o LAGs a nd g row th

i . e . , b r igh tne s s ( Schm id t & K e i l 1 972 ) . T he se c hange s c or re spond t o c hange s i n t he o r ien ta t ion o f c on s t i tuen t c o l lagen f i b re s , b e t ray ing a n a d ju s tmen ti nt he d i re c t ion o f

z one s , r e spe c t ive ly ( Ca s tane t 1 981 ) . E x is t ing s t ud ie su s ing TE M s how t h a t g row th z one s , a nnu l i a nd LAGs i nb one c an b e d i s t ingu i shed o n t h e b a s is o ft h e o rgan isa t ion o ft h e

g row th ( I b id .) . Ne ighbou r ing l a ye r sw i th d i f f e ren tt a ngen t ia l f i bre o r ien ta t ion a re r e ad i ly d i s t ingu i shed b y t h e ir o ppo s ing s i gn s , a cco rd ing t o S chm id t & K e i l ( 1972 :302 ) . R ecen t o bserva t ion s n o te t h a t a nnu l i a re a n iso t rop ic a nd a l te rna te w i th g row th z one s , wh ich a re i s o t rop ic ( Ca s tane t 1 981 , 1 982 ) . I n t h ea nnu l i , c o l lagen f i br i ls a r e o r ien ted i n amore r egu la r f a sh ion t han i ng row th z one s , r e semb l ing t h ose i n l ame l la r o r p a ra l le l f i be red -bone ( Ca s tane t 1 981 , 1 982 ) .

c o l lagen f i br i ls c ompos ing t h em a nd r e la ted d i f f e rence s i n m ine ra l d en s i ty , f i br i l la r d en s i ty a nd g round s ub s tance ' ( e .g . , C a s tane t 1 981 ) . T he se o b serva t ions p robab ly h o ld t r ue f o r c emen t , w h ich i s at i s sue c l o se ly r e la ted t ob one , a s a l ready s t a ted .

LAGs a re more b i ref r ingen t t han e i the r a nnu l i o r g row th z one s ( I b id . ; F ranc i l lon -V ie i l lo te ta l . 1 991 ) .

2 .6 .5 S cann ing E lec tron M icro scope . S cann ing e l ec t ron m ic ro scopy c an b eu sed t o e xam ine g round , e t ched s u rf ace s o f c emen t a nd b one . E tch ing w i th a c id ( e .g . o r thopo sphor ic a c id , H i l l son 1 986 :170 ) r e ve a ls t h e s t ruc tu re o ft h e t i s sue , r e su l t ing i na p a t te rn o f r i dge s a nd t r ough s wh ich c o r re spond s t o t h e p a t te rn o fb r igh ta nd d a rk i n c remen t s s e en u nde r p o la r ised l i gh t a nd t h e p a t te rn o f mo re a nd l e s s c h romoph i le b and s

2 .6 .2 O rd inary L igh t I mage s o f i n tac t d en ta l t h in se c t ion s u nde r t r an sm i t ted o rd ina ry l i gh t a l so s how al a yer ing i nc emen t . T h is l a ye r ing may r e su l t f r om b i ref r ingence ( compa rab le t o p o la r ised i mage s ) , l i gh t s ca t te r ing p a t te rn s d ue t o 8

s e en i n s t a ined h i s to log ica l s e c t ion s ( Boyde 1 980 ; Hohn 1 980 ) . T he " t rough " r eg ion s ( s how ing l ow r e l ie f )

m ine ra l iza t ion i s l e ss

c or re spond t o r e la t ive ly more m inera l iz ed a r eas ( Boyde

o r ien ta t ion o f t h e c o l lagen f i bre s when d e term in ing t h e

1 980 :68 ) . T h is i s c onf i rmed b y C a s tane t 's s t udy u s ing g rea ter

o p t ica l e xpre ss ion o ft h e g row th l a yer s . A s C a s tane t s t a te s : " ce s f a ib le s v ar ia t ion s d e f o r te s t e neur s m ine ra le s

m ine ra l iza t ion ( p e rv o lume ) wa s d i scove red t ob e g ene ra l ly

n ' in ter v iennen t p resque p a s d an s l ' expre s s ion o p t ique d e s

c ha rac ter is t ic o f LAGs ( Ca s tane t 1 979 ) .

marque s d e c ro issance s que l le t ique s" ( Ca s tane t 1 982 :60 ) . A ccord ing t o C as tane t , a nnu l i a r e a n iso t rop ic b ecau se t h e

2 .6 .6 H is tochem ica l Ana lys is .

c omponen t c o l lagen f i br i ls a re r egu la r ly o r ien ted ( a nd

q uan t i ta t ive

m icrorad iography ,

where

a

( K leveza l p er s . c omm . 1 991) . V ar ia t ion i n t h e d egree o f s i gn if ican t t han c hange s

i n t h e

c or re spond g enera l ly t o al ame l la r b one s t ruc ture ) ; g row th H is tochen a ica la na lyse sh ave r evea led t h a t

z one s

a re

i s o t rop ic

b ecause

o f

t h e ir

v ar iab le

f i b re

t h e re a r e b a s ic d i f f e rence s i nt h e c hem ica l c o mpos i t ion o f

o r ien ta t ion s , c or re spond ing s t ruc tu ra l ly t o woven b one a nd

a nnu l i a nd g row th z one s ( G l imche r 1 972 ; C a s tane t 1 982 ) .

o cca s iona l ly t op a ra l le l -f ibe red b one ( Ca s tane t 1 982 ) .

Annu l i a r e b a soph i l ic , a s r evea led i np a r t b y t h e ir r e ac t ion t ov ar iou s c hem ica ls ( K leveza l &K le inenbe rg 1 967 ; B oyde

C o l lagen a nd m ine ra l a l ignmen t a r e b o th

1 980 ; C a s tane t 1 981 ) . T he re i s , f u r thermo re , ad i f f e rence

a ppa ren t c ause s o f t h e p o lar isa t ion p a t tern s o bserved i n

i n t h e c oncen t ra t ion o fg lycosam inog lycane s i n a nnu l i a nd

i n tac tg round s e c t ion s ( Schm id t&K e i l 1 972 ; Myr ick 1 980 ;

g row th z one s( Ca s tane t 1 982 :56 ) . C ompo s i t iona la na lys is o f c emen tum h as a l so r evea led aq uan t i ta t ive d i f f e rence i nt h e

C a s tane t 1 982 ) . Howeve r , s i nce c rys ta ls i nb one o r ien tt h e ir l ong a x is p a ra l le l t o t h e l ong a x is o ft h e f i bre b und le s t h a t

r e la t ive p e rcen tage o fs u lfu rp re sen ti nt h ed i f f e ren tt ype so f

s e rve a st h e ir ma tr ix ( Boyde & J one s 1 972 ; S chm id t&K e i l

g row th l a ye r ( K leveza l 1 988 ; p e r s . c omm . 1 991) .

1 972 ) , t h ec rys ta l lograph ic a r rangemen ti nb one a nd c emen t i sp r imar i ly d ependen t o n t h e a r rangemen t a nd o r ien ta t ion

2 .6 .7 M icrorad iography .

o f t h e c o l lagen f i bre s ( Schm id t & K e i l 1 972 ; C a s tane t 1 982 ) .

M icrorad iog raphy a l so r e vea ls ab anded o r i n cremen ta l

s t ruc tu re

i n c emen t

a nd

b one ,

g ene ra l ly

a ppear ing a s a l te rna t ive ly r ad iopaque a nd r ad io lu scen t

A l li nves t iga t ions t od a te , t h e refo re , s how t h a t t h e o p t ica l a nd p hys ica l e xpre ss ion o fg row th l a ye r s i n

l a ye r s

c emen t a re ma in ly t h e r e su l t o fc o l lagen f i bre o rgan isa t ion

( Hohn

1 980 ;

d 'Huar t

1 980 ;

C a s tane t

1 982 ) .

A ccord ing t o C a s tane t t h is d i f f e rence i n r ad iopac i ty i s

a nd v ar ia t ions i nc e l ld en s i ty ( Ca s tane t 1 982 ; H i l lson 1 986 ) .

i nf luenced b y t h e d i f f e rence s o fp oros i ty ( c e l lu la r d en s i ty ) b e tween a nnu l i a nd g row th z one s b u t a l so b y r e la t ive d i f f erence s i n m ine ra l d en s i ty ' b e tween LAGs a nd g row th

T he f i r s t o bserve r s o fg row th s t ruc tu re s i n b one a nd c emen t w e re c oncer ned w i th d e scr ib ing a nd

z one s ( Ca s tane t 1 981 , 1 982 ) a s r evea led b y q uan t i ta t ive

e xp la in ing t he a ppa ren t p hys ica l c ha rac ter is t ic s r e spon s ib le

m icrorad iograph ic s t ud ie s ( Ca s tane t 1 979 ) .

f o r t h e o p t ica l q ua l i t ie s o ft h e s t ruc ture s ( d e R icq ls 1 975 ,

2 .7 D i scu s s ion : G row th l ayer s i n c emen t .

empha s iz ing t he f unc t iona l s i gn if icance o f c emen t ( e .g . ,

1 979 ) . O sborn T he l a yer ing o b ser ved b y t h e v a r iou s

Wi th

t h e

1 981) ,

a dop t ion o f a c l a s s if ica tory h oweve r ,

f unc t iona l

s y s tem

e xp lana t ion s

o f

c emen tum l a ye r ing a re s ough t . R a te o f d epos i t ion i st h e

me thod s d i scu s sed a bove c or re spond s t o t h e p resence o f

mo s t l i ke ly c au se o fs t ruc tura l v ar ia t ion i nc emen t .

g row th l a ye r s , c omposed o f g row th z one s , a nnu l i a nd LAGs . An umbe r o f p os s ib le e xp lana t ion s f o r t h e o p t ica l

2 .8 D epo s i t ion o f t he g row th l ayer s .

a nd p hy s ica l e xpre ss ion o f g row th l a ye r s i n b one a nd c emen t h ave b een o f f e red : c hange s i n m ine ra l d en s i ty ;

G row th l a ye r sa r ed epos i ted a ppos i t iona l ly

c e l lu la r d en s i ty ; h i s tochem ica l d i f f e rence s ; a nd c o l lagen

i nb one , c emen t a nd d en t in ( Peabody 1 961 ; War ren 1 963 ;

f i bre o r ien ta t ion .

K leveza l & K le inenbe rg 1 981) . E a r ly a t temp t s t o e xp la in t h e d i f f e rence s

T he

c on s ide rab le

1 967 ; K leveza l

1 970 ; Monk s

t im ing o f t h is a ppos i t iona l g row th i s o f i n te re s t

t o r e sea rche r s

i n t h e

b io log ica l

i no p t ica l e xpre s s ion o ft h eg row th z one s a nd a nnu l i t e nded

s c ience sb ecau se i th a sb een a s soc ia ted w i th c yc l ica l g row th

t o c oncen t ra te o n t h e e f f ec t s o f m ine ra l d en s i ty (War ren

o ft h e a n ima l o rgan ism .

1 963 ; K leveza l & K le inenbe rg 1 967 ) . I n t heory , s i nce h ydroxyapa t i te c r ys ta ls w e re k nown t ob e o fo ppos i te s i gn t o c o l lagen ( S chm id t & K e i l

E a r ly t heor ie s l i nk ing c hange s

i n t h e

1 972 ) wh ich i s p o s i t ive

s t ruc tu re o fb ony t i ssue s t or a te o fd epos i t ion i n c lude t h ose

( H i l lson 1 986 ) a nd s i nce r emova l o f HA c r ys ta ls t h rough

o f Ampr ino ( 1947 , a nd 1 967 , a s q uo ted i nK leveza l 1 980

d em inera l isa t ion

i n c reased

a nd i nC a s tane t 1 982 ) a nd E g l i ( 1946 ; a sq uo ted i nS chm id t

b i ref r ingence ( Ke i l 1 954 ) , t h e a n iso t ropy o ft h e a nnu l i w a s

& K e i l 1 972 ) . E g l i a s soc ia ted a r re s ted d epos i t ion , i . e . , t h e

wa s

s hown

t o

r e su l t

i n

c ons ide red t ob e ar e su l t o ft h e ir i n c rea sed m ine ra l d en s i ty

f o rma t ion o fa n a r re s ted g row th l i ne o r LAG , w i th ap e r iod

( K leveza l & K le inenbe rg 1 967 ) .

o ft o o th e rup t ion ( Schm id t&K e i l 1 972 ) . Ampr ino 's ( 1 947 ) h ypo the s is , s ugge s t ing r e la t ive d i f f e rence s i n t h e s peed o f

S t r ic t ly s peak ing , h oweve r , t he r e la t ive

d epos i t ion o f t h e d i f f e ren t t ype s o f b one t i ssue ( p a ra l le l -

d eg re e o fm ine ra l iza t ion b e tween a nnu l i a nd g row th z one s

f i be red , l ame l la r a nd woven b one , s e e a bove , F ranc i l lon -

d oe sn o td i f f e rg rea t ly i nb one ( Ca s tane t 1 979 )o ri nc emen t 9

V ie i l lo t e t a l .1991 ) h a s b e en v e r if ied t h rough s t ud ie s o f f l uo re scen t m a rk ing i n b one ( de B uf f ren i l 1 980 ; C a s tane t

A cce s so ry g row th l a ye r s h ave b een o b se rved b y m any i nve s t iga to r s ( K leveza l & K le inenbe rg 1 967 ; R e ime r s & No rdby 1 968 ; S axon & H igham 1 968 ;

1 980 , 1 982 ) . G row th z one s i nb one , g ene ra l ly f o rmed o f w oven -f ibe red o rl ame l la rb one , c or re spond t o mo re r ap id ly

M i tche l l 1 969 ; M i l le r 1 974 ; G rue & J en sen 1 979 ; i n te r a l i i ) . T he se l a ye r s a r e t yp ica l ly e xp re s sed a st h inne r , mo re e pheme ra l b and s , s ome t ime s d i scon t inuou s a nd g ene ra l ly l a ck ing i na ny r egu la r p e r iod ic i ty ( K leveza l &K le inenbe rg 1 967 ; R e ime r s & No rdby 1 968 ; K levez a l 1 970 ; G rue & J en sen 1 979 ; Monk s 1 981 ) . A g rea te r w id th o f c em en t

d epo s i ted f o rma t ion s t h an a nnu l i , wh ich a r e g ene ra l ly f o rmed o f p a ra l le l f i be red o r l ame l la r b one ( I b id . , F ranc i l lon -V ie i l lo te ta l . 1 991 ) . LAG s o n t h e o the r h and , r e p re sen t " i nunda ted s ur f ace r eg ion s " (Wa r ren 1 963 :28 ) , i . e . , p e r iod so f" a r re s ted a c t iv i ty" ( Grue & J en sen 1 979 :3 ) , c or re spond ing t o p hys iochem ica l t r an sforma t ion s o n t h e

d epo s i t ion ( e .g . , aw ide r z one ) i s c apab le o f r eg is te r ing

t i s sue s u rf ace d u r ing a momen ta ry a r re s t o f g row th ( Ca s tane t 1 981 :15 ) , a nd a s s uch a re n o t t h e r e su l t o ft i s sue

mo re n ume rou s s e conda ry e ven t s i n t h e f o rm o f s ma l l , a cce s so ry b and s ( Ca s tane t 1 982 ) . T he e xp re s s ion o f t h e se

a ppos i t ion .

a cce s sory i n c remen t sm ay i n te r f e re w i th i d en t if ica t ion o ft h e p r ima ry ( a nnua l ) i n c remen t s , e . g . , t h is i sa ppa ren t ly t h e

2 .9 .0 Ra te a nd D egree o fM inera l isa t ion .

c a se w i th d ome s t ic d og s ( Can is f am i l ia r is ) ( s e e G rue 1 976 ; G rue & J en sen 1 979 ) . B one r emode l l ing a nd o the r mo rphogene t ic p roce s se s m ay a l so b e r e spon s ib le f o r s ome o ft h e o b se rved a c ce s sory l a ye r ing ( Ca s tane t 1 981 ) .

K leveza l & K le inenbe rg a s soc ia te i n c remen ta l g row th w i th c hange s i n t h e r a te o f m ine ra l iza t ion ( K leveza l & K le inenbe rg 1 967 :17 ) w i thou t m ak ing e xp l ic i t t h e m echan ism wh ich wou ld r egu la te t h is p roce s s . J one s &B oyde ( 1972 )a s soc ia te t h is m ine ra l iza t ion p a t te rn w i th t h er e la t ive p ropor t ion so fe x tr in s ic ( Sha rpey 's f i b re s ) a nd i n t r in s ic f i b re s , i nt u rn r e la ted t os peed o ft i s sue d epo s i t ion . O the r a u tho r s l i nk s peed o f d epo s i t ion w i th d eg ree o f m ine ra l iza t ion , a s a c on sequence o f t h e p ropo r t ion o fe x t r in s ic f i bre sa nd c e l ls p e rv o lume o ft i s sue ( e .g . , O sbo rn 1 981 ) . A cco rd ing t o t h is h ypo the s is , r a p id c emen togene s is , wh ich r e su l t s i n t h e i n c lu s ion o f g rea te r n umbe r so fe x t r in s ic c o l lagen f i b re s i n to t h eo rgan ic m a t r ix a nd t h e e mbedd ing o f c emen tocy te s , r e su l t s i n al e s se r m ine ra l d en s i ty p e rv o lume o ft i ssue ( Boyde 1 980 ; O sborn

T he f o rma t ion o fs e conda ry g row th l a ye r s h a s b een l i nked w i th " ba s ic e ven t s i n a n a n ima l 's l i f e " ( K leveza l 1 970 :266 ) . S ome o f t h e " e ven t s " p ropo sed i n c lude t h e r u t ( Low & C owan 1 963 , a s q uo ted i n Monk s 1 981 ; R e ime r s & No rdby 1 968 ) , n u r s ing ( K leveza l 1 970 ) a nd t h e a t ta inmen t o f s exua l m a tu r i ty ( K leveza l & K le inenbe rg 1 967 ; M i tche l l 1 969 ; K leveza l 1 970 )b u tt h e se f a c to r s h ave n o tb een c onc lu s ive ly p roven i na ny b u t af ew p opu la t ion s ( M i l le r 1 974 ) . 2 .9 .2 Annua l G row th L ayer s .

1 981 ) . 2 .9 .1 P r imary , L ayer s .

S econdary

a nd

A cce s sory

B io log is t s inve s t iga t ing inc remen ta lg row th l i ne si n aw ide v a r ie ty o fv er teb ra te s pe c ie sf r om d ive r s if ied e nv ironmen t s h ave emp ir ica l ly i d en t if ied ay ea r ly c yc le o f

G row th

c emen tum f o rma t ion , c on s is t ing g ene ra l ly o f as i ng le p a i red g row th z one + a nnu lu s /LAG ( Peabody 1 961 ; L ow & C owan 1 963 ; McEwan 1 963 ; K leveza l & K le inenbe rg 1 967 ; R e ime r s & No rdby 1 968 ; M i t che l l 1 969 ; G rue & J en sen 1 979 ) . T he s ame p a t te rn h a s b een e s tab l ished b y h e rpe to log is t s f o r v a r iou s p o ik i lo the rm ic s pe c ie s (Wa r ren 1 963 ; C a s tane t e t a l . 1 979 ; C a s tane t 1 980 ; d e B uf f ren i l 1 980 ) a nd b y r e s ea r che r s s t udy ing m a r ine m amma l s ( s ee r ev iew i n S chef f e r & Myr ick 1 980 ) . T he se emp ir ica l

O nce t h e a s soc ia t ion b e twe en l a ye r ing i n b ony t i s sue sa nd r a te o fg row th h ad b een e s tab l i shed , m any s c ien t is t s b egan s e a r ch ing f o r c or re la t ion s b e tween t h e o b served p a t te rn s a nd k nown p hys io log ica l e ven t s . I tw a s s oon d i scove red t h a t g row th l a ye r s c ou ld b e d iv ided i n to t h ree c a tegor ie s : p r ima ry i n c remen t s , p os se s s ing a p red ic tab le p e r iod ic i ty , c o r re spond ing t o as pe c ie s -spe c if ic a nnua l g row th c yc le ; s e conda ry i n cremen t s p o sse s s ing

o bserva t ion s , b a sed o n s t ud ie so fc on t ro l g roup so fa n im a l s o f k nown a ge a nd s ea son a t d ea th , a r e s uppor ted b y e xper imen ta l s t ud ie s i nvo lv ing t h e u se o ff l uo roma rke r s a s

p red ic tab le s ub -annua lp e r iod ic i ty ;a nd a cce s so ry i n cremen t s w hose e xac t t im ing i s f a r l e s s p red ic tab le a nd i n d iv idua l ised .

" benchma rk s " t or e co rd t h e p o s i t ion o f c emen t g row th a t p rec ise i n te rva l s ( K leveza l 1 980 ; C a s tane t 1 980 ; d e

E a r ly o b se rva t ion o fa n a ppa ren t ly l i nea r r e la t ion sh ip b e tween t h e n umbe r o f g row th l a ye r s a nd c emen t w id th wou ld i nd ica te t h e e x is tence o f ac on s tan t g row th r a te o f c emen t ( M i tche l l 1 969 ) . T h is s t a temen t i s q ua l if ied t oday b y s t ud ie si nd ica t ing t h a t ac on s tan tn umbe r o f p r ima ry l a yer s i sd epo s i ted a nnua l ly , b u t t h a t t h e t o ta l t h ickne s s o f t h e l a ye r s v ar ie s r e la t ive t o t h e a ge o f t h e i n d iv idua l ( Wa r ren 1 963 ; K leveza l 1 970 , 1 988 ; C a s t ane t 1 982 ; H i l lson 1 986 ) . F unc t iona l s t re s se s , t r igge r ing c emen tum d epo s i t ion , w i l l h ypo the t ica l ly a l so a f f ec t t h e w id th o ft h e a nnua l l a ye r s b u tn o t t h e ir n umbe r s ( K leveza l

I n t empe ra te c l ima te s , t h e n orma l ly a ccep ted t ime f o r t h e f o rma t ion o f t h e a nnu l i /LAG i s w in te r , h ence t h e o f t en u sed d e s igna t ion " w in te r l i ne " ( Re ime r s &N o rdby 1 968 ; S axon & H igham 1 968 ; M i tche l l 1 969 ; M i l le r 1 974 ) . C au t ion mu s t b e e xe r c i sed , h oweve r , s i nce G rue & J en sen , d ocumen t ing i n c remen ta l g row th i n 5 2 s pec ie s , h ave d i scove red t h a t " w in te r " l i ne s may b e

1 970 ; p e r s . c omm . t oa u tho r , 1 991 ) .

a nnu l i /LAG s f o r a ny

B uf f ren i l 1 980 ; G o rev ich & S t ewa rd 1 980 ) .

f o rmed a to the rt ime sd ur ing t h ey ea r( Grue & J en sen 1 979 ; s e e a l so C a s tane t 1 980 ) . T he t ime o f f o rma t ion o f o ne

s pe c ie s

mu s t t h e refo re b e

e mp ir ica l ly e s tab l ished . N eve r the le s s , t h ea u tho r s s t a te t h a t 1 0

a nnu l i /LAGs t y p ica l ly f o rm d ur ing a t h ree -mon th p e r iod a nd t h a t m amma ls i n A rc t ic a nd T empera te r eg ion s f o rm

R educed f ood i n take , ap laus ib le c au se o f p hys io log ica l g row th s l owdown a nd h ence a nnu l i a nd /or

o ne l i ne p e r y ea r r ega rd le ss o f a ge o r s exua l d i f f e rence s

LAG f o rma t ion , i sn o t o n ly d ue t oe x terna l i n f luence s s uch a s t h e s ever i ty o f w in te r i n t empe ra te a nd a r c t ic r e g ion s ,

( Grue & J en sen 1 979 ) .

h oweve r . G rue n o ted t h a t m ink , r a ised o n f a rm s a nd f e d a s t eady d i e t , r educe t h e ir f ood i n take d ur ing w in te r mon th s

T he a nnua l i n cremen ta l g row th c yc le i s i d en t if ied w i th a n a nnua l c yc le o fp hys io log ica l g row th a t

( Grue 1 976 ) . T h is i mp l ie s ac hange i n i n te rna l m e tabo l ic

t h e l e ve l o f t h e o rgan ism ( K leveza l & K le inenbe rg 1 967 ;

r a te s ,

K leveza l 1 970 ; M i l le r 1 974 ; C a s tane t e ta l . 1 979 ; C a s tane t

H igham 1 968 ; P ike -Tay 1 991 ) .

p erhaps t r iggered b y p ho toper iod ism ( Saxon &

1 981 ; i n te r a l i i ) . T he f a c t t h a t s eve ra l d i f f e ren t t ype s o f b ony t i ssue f o rm i n c remen ta l l i ne s i n s ynch rony , i . e . , d en t in ,

c emen t

a nd

p er ios tea l

b one ,

s uppor t s

t h is

2 .10 .2 Endogenou s f ac tor s .

i d en t if i ca t ion ( K leveza l 1 970 ; 1 980 ) . A ge a nd s e a sona l d a ta o b ta ined t h rough a na lys is o f i n cremen ta l g row th c o inc ide w i th

s im i la r

d a ta

o b ta ined

t h rough

v ar iou s

o the r

T he p re sence o f a n e ndogenou s g row th r egu la t ing c emen tum d epos i t ion r a te a nd i t s

r hy thm ,

morphome tr ic m e thod s i n l i v ing s pe c imens a nd i n s ome

s t ruc tu ra l e xpre ss ion , h a s b e en p roposed (War ren 1 963 ;

c a se sp rov ide b e t te re s t ima te s ( Nova row sk i 1 965 ; G i lber t&

K leveza l 1 970 , 1 980 ; K leveza l & M ina 1 973 ; G rue &

S to l t 1 970 ; M i l le r 1 974 ; d 'Huar t 1 980 ; P asca l & D e la t t re

J ensen 1 979 ; C a s tane t 1 980 , 1 981 , 1 982 ) . E xper imen ta l

1 981 ; S t a l l ib ra s s 1 982 ; P asca l & K ovaks 1 983 ; Que ré &

s t ud ie s

P a sca l 1 983 , 1 984 ) , f u r the r s uppor t ing t h e i d en t if ica t ion o f

h ypo the s is o fa ne ndogenou sr hy thm . H ibe rna tor sw i l lf o rm

o f

h ibe rna tor s

a nnua l

a nd

i n cremen t

d ome s t ica te s b e ing

p reven ted

t h e

a n a nnua l p e r iod ic i ty i nt h e f orma t ion o fp r imary l a ye r s i n

a n

c emen t . F luoroma rke r s t ud ie s a l so c onf irm t h is h ypo the s is

h iberna t ing ,i . e . , i nt h ea b sence o fe nv ironmen ta li nf luence s

f r om

f o r t e r re s tr ia l m amma ls ( K leveza l 1 980 ; 1 988 ) , m ar ine

( K leveza l

m amma ls ( Go rev ich & S t ewar t 1 980 ) , amph ib ian s , a nd

i n f luence s o n ly r e su l t s i n aw eake r e xp re s s ion o f t h e

r e p t i le s

p r imary i n cremen t s , n o t t h e ir e r ad ica t ion , s uppo r t ing t h e

( F ranc i l lon

1 979 ;

C a s tane t

1 982 ;

C ae tano

&

C a s tane t 1 987 ) .

1 980 ,

d e sp i te

s uppor t

1 988 ) . T he a b sence o f e nv ironmen ta l

p re sence o fo n togene t ic c on t ro l o ft h e ir f o rma t ion ( Grue & J en sen 1 979 ; C a s tane t 1 982 ) . T he p re sence o f g row th

2 .10 .0 C au sa l F ac tor s i n t he C yc l ica l D epos i t ion o f

mark s i n b o th e ndo therm s a nd e c to therm s i s a l so b e s t e xp la ined b y t h e h ypo the s is o f a n e ndogenou s r hy thm

B ony T i s sue s .

s e condar i ly 2 .10 .1 Env ironmen ta l f ac tor s .

r e info rced

a nd

e nv ironmen t ( Ca s tane t 1 982 ) .

s ynch ron ised

e ndocr ine a c t iv i ty i n t r op ica l v er tebra te s , S evera l

c ausa l

f a c tor s

f o r t h e a nnua l

w i th

t h e

T he p re sence o f s e a sona l r e su l t ing i n

c yc l ica l i n cremen ta l g row th , c anno t b e a s cr ibed t om a rked

g row th c yc le o fc emen t h ave b een p ropo sed o ve rt h e y ea r s .

s e a sona l f l uc tua t ions a nd s uppor t s t h e h ypo the s is o f a n

S i nce

t h e

e ndogenou s r hy thm (War ren 1 963 ) . A more e x t reme v i ew

s y s tema t ic a ff i l ia t ion o fs pe c ie s a nd t h ep a t tern o ff orma t ion

i st h a t o f Ma r sh ( 1980 ) , who s ugge s t s t h a t l a ye r d epo s i t ion

t h e re

i s n o

a ppa ren t

c or re la t ion

b e tween

o ft h e a nnua l l a ye r s ( Grue & J en sen 1 979 ) , e nv ironmen ta l

i sa n e ndogenou s r hy thm b eg inn ing a tb i r th ( i .e . , t r igge red

i nf luence s h ave l o ng b een c ons ide red r e spon s ib le f o r t h e ir

b y

e xp re s s ion . M i tche l l c on s ide rs t h e c or re la t ion w i th b ody

a synch rony i n s pe c ie s w i th a n e x tended b i r th ing s e ason ,

g row th t ob e ar e su l to fg ene ra l p hys io log ica lc ond i t ion ( t he

e .g . , t h e d ugong ( Dugong d ugong ( M i l le r ) ) .

b i r th

a l one ) ,

t hu s

e xp la in ing

a ppa ren t

s ea sona l

" p rox ima l f a c tor ") i n t u rn u l t ima te ly c au sed b y d i e tary r e s tr ic t ion s( t he " u l t ima te f a c tor ") ( M i tche l l 1 969 :283-284 ) .

A l though t h e p r inc ipa l c au sa l f a c tor o f

K leveza l & K le inenbe rg c on s ide r p er iod ic c hange s i n

d i f f e ren t ia l i n cremen ta l g row th a ppea r s t ob e e ndogenou s ,

i n tens i ty o f f e ed ing o r , a l terna t ive ly , h ormona l c hange s t o

e x te rna l f a c tor s a r e s t i l l o pe ra t ive i n mos t c a se s . I ts e em s

b e r e spon s ib le ( 1967 ) .

i mprobab le t h a t o bserved

I n tens i ty o f f e ed ing h a s b een

g row th

( 1963 ) , c i ted b y K leveza l & K le inenbe rg ( 1967 ) , a s t h e

c o inc iden ta l . T he d eg ree o fc ana l isa t ion ( e .g . , t h ed eg ree o f

p r inc ipa l e nv ironmen ta l f a c tor r e spon s ib le f o r s t ruc tu ra l

c or re la t ion

c hange s i n c emen t .

S axon & H igham ( 1968 ) c on s ide r

f l uc tua t ion a nd t h e r e la t ive w id th o f c emen tum l a ye r s ) i s

r educed f o od i n take t ob eo n ly i nd ire c t ly c au sa l . A ccord ing t ot h e se a u tho r s , r educed f ood i n take r e su l t s i n ar educ t ion

M i tche l l o bse rve s n o c or re la t ion b e tween t h e w id th o f

o f

c emen tum d epos i t ion i nc erv ids a nd t h e s ever i ty o fw in te r s

t h e

f unc t iona l ,

b iome chan ica l

s t re sse s

s t i l l

s t imu la t ing

a nd

c hange s

c or re la t ions b e tween a nnua l

p ropo sed b y b o th McEwan ( 1963 ) a nd L ow & C owan

b e tween

i mprope r ly

i n t h e t h e

e nv ironmen t

amp l i tude

u nder s tood ,

o f

h oweve r

a r e

m e re ly

e nv ironmen ta l ( Boyde

1 987 ) .

c emen tum g row th a nd i ti st h is s t re s s r e duc t ion wh ich i st h e

( M i tche l l 1 969 ) b u t C a s tane t n o te s t h a t t h e e xpre s s ion o f

d i re c t c au se o f t h e f o rma t ion o f a nnua l " w in te r r e s t l i ne s "

i n cremen t s i n r ep t i le s m ay b e a t tenua ted b y t h e l a ck o f

( S axon & H igham 1 968 ) . L ieberman ( 1993 ) h a s s hown t h a t

c l ima t ic c on t ra s t s ( s ee a l so K leveza l 1 980 ) . G rue n o ted t h a t

b o th r e duced n u tr i t ion a nd b iomechan ica l s t re sse s c an b e

t h e e xpre ss ion o f p r ima ry i n c remen ta l l i ne s w a s o n ly

r e spon s ib le f o r t h e f o rma t ion o f a nnu l i i n g oa ts . R educed

a f f ec ted i n t hose d ome s t ica te s t h a t w e re e n t ire ly r emoved

n u tr i t ion w a ss hown t or e su l ti ns l owe rt i ssue d epos i t ion a nd

f r om s ea sona l e nv ironmen ta l f l uc tua t ion s ( Grue 1 976 ) .

c onsequen t h yperm ine ra l isa t ion ; c hange s i nb iomechan ica l

G rue 's c onc lu s ion i sc ha l lenged , h oweve r , b y e v idence o f

s t re ss w e re s hown t o r e su l t i n v ar ia t ions i n c o l lagen f i bre

a nnua l i n cremen ta l b and ing i n h uman s who a r e l a rge ly

o r ien ta t ion , p e rce ived a s b and ing ( L ieberman 1 993 ) .

b uf f e red f r om t h e ir e nv ironmen t s ( S chm id t & K e i l 1 972 ; 1

O sbo rn 1 981 ; S to t t , S i s &L evy 1 982 ) .

i na ncho r ing t e e th i nt h ea l veo la rc av i ty o fm amma ls ( Saxon & H igham 1 968 ) , c emen t i st h e p re f e r r ed t i s sue f o r t h e c o l lec t ion o fv i ta l d a ta ( Boyde 1 980 ; Monk s 1 981 ) .

T he h ypo the s is o f s e conda ry s e a sona l s ynch ron isa t ion , c oup led w i th t h e p re sence o fa n i nhe r i ted , s pec ie s -spec if ic e ndogenou sr hy thm ,e xp la in sw hy f o rma t ion o fi n c remen t s i n membe r s o ft h e s ame e co log ica l s y s tem i s n o t n ece s sa r i ly s ynch ron ised ( Grue & J en sen 1 979 ) . S ea sona l i nf luence s won ' t b e e xpre s sed t h e s ame w ay b e tween s pec ie s ( K leveza l 1 980 ; C a s tane t 1 980 ) .

Wi ld l if e

b io log is t s

s t udy ing

l i v ing

p opu la t ion s a r e c once rned w i th e s tab l ish ing t h e a ge o ft h e i nd iv idua l sp re sen t i nt h e t a rge t g roup . T h is i n f orma t ion i s u sed t oe s tab l i sh a n a ge p rof i le wh ich i si nt u rn u sed i nt h e m anagemen t o f t h e p opu la t ion ( s e t t ing h un t ing l im i t s ,

E ndogenou sp hy s io log ica lr hy thm sa r es t i l l

d i agno s ing h ea l th p rob lem s a nd a na ly s ing p opu la t ion g row th ) . A n umbe r o f r e sea r che r s h ave s ucce s sfu l ly p ionee red t h eu se o fs ke le toch rono logy ( t he u se o fg row th l a ye r s i n c emen t o r o the r b ony t i s sue s ) f o r t h e s t udy o f

i mp rope r ly u nde r s tood , l a rge ly b e cau se o f t h e n umbe r o f v a r iab le sw h ich i n f luence t h e ir e xpre s s ion . T he re i sn oc l ea r

l i v ing p opu la t ion sf o r aw ide r a nge o fs pe c ie s ( Novakow sk i 1 965 ; A dam s & Wa tk in s 1 967 ; R e ime r s & No rdby 1 968 ;

e vo lu t iona ry s i gn if icance f o rs ynch ron i sa t ion o fe ndogenou s a nd s e a sona l r hy thm s , n o r i s t h e re a ny c l ea r e co log ica l

M i tche l l 1 969 ; G i lbe r t & S to l t 1 970 ; M i l le r 1 974 ; H a l l -

2 .10 .3 G enera l c onc lu s ions .

Ma r t in 1 976 ; d 'Hua r t 1 980 ; Q u6 re & P a sca l 1 983 ; C ae tano e ta l . 1 985 ; McCu l lough & B e ie r 1 985 ; C a s tane t 1 986 -87 ; C a s tane t , N ewman & S a in t -G iron s 1 988 ; Wa t t s & G a sk in 1 989 ; i n te r a l i i ) .

c on sequence o f t h is s ynch rony ( En r igh t 1 970 ) . T he o n ly e xp lana t ion f o rt h is mechan ism m igh tb et h a t : " Endogenou s r hy thm ic i ty s imp ly p rov ide s t h e o rgan ism w i th at empo ra l i n e r t ia , a t ime -dependen t r e ad ine s s t o r e spond t o e nv i ronmen ta l s t imu l i i n a n a pp ropr ia te m anne r " ( En r igh t 1 970 :236 ) . I nt h e c a se o fe ndogenou sg row th r hy thm s , t h e s en s i t iv i ty o f v a r iou s t i s sue s i n r e co rd ing c hange s i n a n o rgan ism s ' g row th r a te w i l l v a ry , d epend ing l a rge ly o n t h e ir own g row th r a te , a nd m ay m a sk t h e p r ima ry c yc le ( K leveza l 1 980 ) .

F or mo s t s pe c ie s , ac oun t o ft h e n umbe r o fp r ima ry i n c remen t s p re sen t i nt h e c emen tum o f ag iven t oo th , c omb ined w i th ac a l ib ra t ion f a c to r , y i e ld s t h e a ge i n y ea r s o ft h e a n ima l ( McEwan 1 963 ; M i tche l l 1 967 ; M i l le r 1 974 ) . T he c a l l ib ra t ion f a c tor i so b ta ined b ye s tab l ish ing t h e a ge o f f o rma t ion o f t h e f i r s t a nnu lu s , f o r e a ch d en ta l a na tom ica l p o s i t ion , e mp ir ica l ly v e r if ied f o r t h e t a rge t

An umbe ro fp hy s io log ica lg row th c yc le s o f a n a ppa ren t ly e ndogenou s n a tu re w i th d i f f e ren t p e r iod ic i t ie sf i nd t h e ir e xpre s s ion i nh a rd t i s sue s . A l though t h e se w i l l n o t b e o f much u se t ou s i nt h e c on tex t o f t h e p re sen ts t udy , t h ey b ea rm en t ion b ecau se t h ey r e inf o r ce t h e h ypo the s is o fa n a nnua l p hy s io log ica l c yc le . Ame logene s i s p roduce s c ro s s s t r ia t ion sw i th ad a i ly ( 24 h ou r ) p e r iod ic i ty ( Myr ick 1 980 ; O sbo rn 1 981 ; H i l lson 1 986 ; B oyde 1 987 ) . T he re may a l so b e ac o r re la t ion b e tween t h e B rown s t r iae o f R e tz iu s , o bserved i n e name l , a nd a s even d ay c yc le ( H i l lson 1 986 ; B oyde 1 987 ) . I n d en t ine , d a i ly s t r iae c a l led v on E bne r 's l i ne s h ave b e en o b se rved ( Boyde 1 980 , 1 987 ; H i l lson 1 986 ;O sbo rn 1 981 )c or re spond ing h ypo the t ica l ly t o t i da l r hy thm s when o ccur r ing i n ma r ine m amma ls ( Ma r sh

s pec ie s ( M i l le r 1 974 ; G rue & J ensen 1 979 ) . S ome a u thor s c on s ide rt h ea ge a te rup t ion ( f o rt h ea na tom ica l p o s i t ion )t o b e t h e c a l ib r a t ion f a c to r ( Sp inage 1 976 ) . A l though c emen tum g row th b eg in s p r io r t o e rup t ion o f t h e t oo th ( Low & C owan 1 963 ; S axon & H igham 1 968 ) , t h e t ime o f f o rma t ion o ft h ef i r s ta nnu lu s /LAG i sn o ta lways c o inc iden t w i th t o o th e r up t ion ( McEwan 1 963 ; G i lbe r t 1 966 ; K leveza l &K le inenbe rg 1 969 ; M i l le r 1 979 ; G rue & J ensen 1 979 ) . S ome a u tho r s s ugge s tt h a tt h ep re sence o f a n a nnu lu s a l low s u s t o s i tua te t h e d ea th o f a n i nd iv idua l w i th in af a i r ly s ho r t , t h ree mon th t ime -f rame ( e .g . , G rue & J en sen 1 979 ; b u t s e e K leveza l 1 987 ) . G row th z one s , h oweve r , a r e g ene ra l ly f o rmed o ve r al o nge r t ime p e r iod ( I b id .) . S ome a u tho r s h ave s ugge s ted t h a t i ti sp o s s ib le t o e s tab l ish t h e s e a son o fd ea th w hen t h e f i na l i n c remen t i sa z one b yc a lcu la t ing t h e amoun to fc emen tg row th wh ich w a s u nderway w hen t h ea n ima l d i ed ( Bou rque , Mor r is & S p ie s s

1 980 ) . A lso i n ma r ine m amma ls , a nnua l g roup ing s o f 1 3 s t r iae h ave b e en o bserved wh ich a ppea r t oc o r re la te w i th a l una r r hy thm ( Myr ick 1 980 ) . 2 .11 .0 P opu la t ion s tud ie s .

1 978 ; G o rdon 1 988 ; P ike -Tay 1 991 ) . T he c a lcu la t ion i s e xp re s sed a s ap ropor t ion o fe xpe c ted g row th ( ba s ed o nt h e t h ick ne s s o ft h ep rev iou s z one ) b ea r ing i nm ind c hange s i n mo rpho log ica l r a te o fg row th d ue t os uch f a c to r s a s a ge o f t h e i nd iv idua l ( I b id . ) . H oweve r , i t h a s n o t b een s a t i sf ac tor i ly p roven t h a td epo s i t ion o ft h e z one i s al i nea r , a nd q uan t if iab le , e ven t( K leveza l 1 987 , 1 988 ; S p ie s s 1 990 ) .

D en t ine a nd e name l a r e n o t o f much u se i n p opu la t ion s t ud ie s o f t e r re s t r ia l m amma l s , s i nce t h e i r g row th i s i mpeded b y an umbe r o f f a c to r s i n c lud ing i nf i l l ing o ft h ep u lpa rc av i ty ( d en t ine )a nd m a tu ra t ion o ft h e c rown ( e name l ) ( K leveza l & K le inenbe rg 1 967 ; G rue & J en sen 1 979 ) a l though t h ey m ay b e u sefu l i n e s t ima t ing s P ason o f d ea th . T he l ong b one s o f t e r re s t r ia l v er teb ra te s a r e l im i ted b y r e so rp t ion a nd r emode l l ing o f t h e t i s sue s , wh ich i n te r f e re sw i th r e ten t ion o fi n c remen ta lg row th l a ye r s

Results

obtained

t hrough

s ke le toch rono logy c ompa re f a vou rab ly w i th r e su l t s o f mo re t r ad i t iona l c ynegene t ic s t ud ie s , s uch a sc r own h e igh t , t oo th w ea r , t o o th e r up t ion a nd r e p lacemen t , l i ve w e igh t a nd e ye

d ur ing t h e c ou r se o f a n o rgan i sm s ' l i f e span ( K leveza l & K le inenbe rg 1 967 ) . Due t ot h el a ck o fm a jo rt i s sue t u rnove r ( a s i n b one ) a nd c on t inued a c t ive g row th t h roughou t a n o rgan ism s ' l i f e , r e su l t ing f r om t h e f unc t iona l r o le i tp l ay s

l e n s m a s s , f o r e xamp le ( s e e c ompa r ison s o f f e r ed i n Nova row sk i 1 965 ; G i lbe r t&S t o l t 1 970 ; H a l l - Ma r t in 1 976 ; H aa s 1 977 ; Q ué rd & P a sca l 1 983 ; Wa t t s & G a sk in 1 989 ) . 1 2

T hey h ave t h e a dvan tage o fb e ing a bso lu te , whe rea s o the r me thod s ( s uch a s t oo th w ea r a na lys is ) o f f e r o n ly r e la t ive i n forma t ion a nd i n s ome c a se s a r e i n adequa te f o r t h e a s se s smen t o fa ge i no lde ra n ima ls ( e .g . , t o o th e rup t ion a nd r ep lacemen t s equence s ) a nd t he refore a r e more u sefu l f o r i n te r -popu la t iona l a na lyse s ( S ta l l ibra s s 1 982 ) . A c or re spondence b e tween t h e g row th s t ruc tu re s o bserved i n r e cen t b ony t i s sue s a nd a na logou s s t ruc tu re s o bse rved i nf o s s i l s pe c imens h a sb een s ugge s ted : " f os s i l a nnu la t ion s a r e s im i la r i n e ve ry r e spec t t ot ho se i n r e cen t v er tebra te s" ( Peabody 1 961 :63 ) ( s ee a l so K leveza l 1 970 ; d e R icq ls 1 975 ) . P rudence i sa dv isab le , h oweve r , s i nce t h e b io logy o f e x inc t p opu la t ion s c an

o n ly b e

i mpe rf ec t ly u nde r s tood ( Ca s tane t , Meun ie r & d e R icq ls 1 976 ) . F os s i ls u nde rgo d i agene t ic p roce s se sr e su l t ing i nl o s s o f c o l lagen , b u t t h e m ine ra l c omponen t o f t h e t i ssue s a nd t h e ir

c ry s ta l lograph ic

a l ignmen t

m ay

r ema in

i n tac t ,

p re serv ing i n cremen ta l s t ruc ture s( Remy 1 974 ) . T h is mean s t h a t wh i le c ynegene t ic a ge d e term ina t ion s s uch a s ma ss o f t h e e ye l e n s , b ody w e igh t , e t c , c anno t b e a pp l ied t o f o ss i l p opu la t ions , s ke le toch rono logy i s s t i l l a pp l icab le . A f ew b io log is t sh ave u sed s ke le toch rono logy t oa na lyse p opu la t ion s t ruc tu re i nf o ss i lp opu la t ion s( de R icq ls 1 975 ; C a s tane t & B aez 1 988 ; K leveza l p e r s . c omm . 1 991 ) b u t o n t h e who le i ti s a r cheo log is t s who h ave b een i n te re s ted i n a pp ly ing s ke le toch rono logy t o f o s s i l ma ter ia l ( S axon & H igham 1 968 ; S p ie ss 1 976 , 1 990 ; C oy e ta l . 1 982 ; S t a l l ibra s s 1 982 ; S ave l le & B ea t t ie 1 983 ; Morey 1 983 ; G ordon 1 988 ; P ike T ay 1 991 ; L ieberm an e ta l . 1 990 ) .

1 3

CHAPTER 3

3 .0 I n troduc t ion .

THE CONTROL SA MPLE . r e su l t o f t h e l o ca l d ome s t ica t ion o f d if e ren t w i ld p opu la t ion s ( e .g . , G rove s 1 974 ; 1 986 ) .E bha rd t 's ( Ebha rd t

T h is C hap te re xam ine st h ea pp l icab i l i ty o f ac on t ro l s amp le o f mode rn , d ome s t ic h o r se f o r t h e s t udy o fd en ta lc emen t i nP a laeo l i th ic h o r se p opu la t ion s( Pa r tA ) . T he r equ i red a s sump t ion s a r e e xam ined a nd j u s t if ied . T he

1 954 , 1 957 , a s q uo ted i n G rove s 1 974 ) a na lys is f u r the r c omp l ica te s t h e s t udy o f P le is tocene r ad ia t ion s b y i n t roduc ing t h e n o t ion o f f ou r " t ype s " o fe qu id , i d en t if ied t h rough a na ly s is o f b ehav iou ra l t r a i t s i n e x is t ing e qu id p opu la t ion s . T he re e x is t s n o i n con t rove r tab le p roof o f a mu l t ip le o r ig in o f t h e c aba l l ine s t ra in , h oweve r (Wa r ing

a na ly s is o ft h e c on t ro l s amp le r e su l t s i n as chedu le f o r t h e f o rma t ion o fg row th l a ye r s i ne qu id d en ta l c emen tum ( Pa r t B ) .

1 983 :9 ) .F u r the rmore ,L undho lm 's mode l( Lundho lm 1 949 , a s q uo ted b y G rove s 1 986 ) o f as epa ra t ion o f t h e " t rue h o r se " i n to a n E a s te rn a nd We s te rn g roup ( l iv ing i n t h e

PART A : THE MODERN CONTROL SA MPLE . 3 .1 App l icab i l i ty o f t he modern c on tro l t o f o s s i l

E u ra s ian s t eppe a nd s epa ra ted b y t h e A lp s ) i sa ccep ted b y s ome mode rn a u thor s .

s pec ie s . T he p u rpo se o ft h e mode rn c on t ro ls amp le i st oe s tab l ish t h e t ime o ff o rma t ion o fi n cremen ta l g row th s t ruc tu re s i n t h e c emen t o f h o r se t e e th i n o rde r t o a pp ly t h e se d a ta t ot h e s t udy o ff o s s i l p opu la t ion s o fh or se . T he u se o f amode rn s pe c ie s f o r t h e s t udy o f af o s s i l o ne r e s t s o n t h e a s sump t ion o f u n if orm i ty . T h is a s sump t ion i s s uppor ted b y t h e f a c t t h a t t h e g ene t ic d i s tance b e tween

B io log is t ss t udy ing e qu id p opu la t ion sp r io r t od ome s t ica t ion i no u r r eg ion o f i n te re s t h ave i nt h e p a s t i d en t if ied t h re e " w i ld " p opu la t ion s : t h et a rpan ( Equu sf e ru s gme l in i ) t h eP rz ,ewa lsk i h o r se ( Equu s f e ru sp rzewa lsk i )a nd t h e f o re s t p ony ( Equu s f en i s s y lva t icu s ) ( An ton inu s 1 913 , a s q uo ted i n G rove s 1 986 a nd i n C lu t ton -Brock 1 987 ; Z eune r 1 963 , a s q uo ted i n War ing 1 983 ) , l a rge ly o n t h e

E quu s f e ru s , t h e w i ld a nce s to r o f t h e d ome s t ic h or se , a nd E quu s c aba l lu s , t h e d om e s t ic h o r se , d oe s n o t a ppea r t ob e g rea t ( we a r e s t i l l d ea l ing w i th e s sen t ia l ly t h e s ame h o r se n ow a nd d u r ing t h e P len ig lac ia l , a bou t 1 4 ,000 t o 1 8 ,000 B .P . a go )( s ee b e low ) . I n f a c t , t h e re e x is t s ah igh d eg re e o f mo rpho log ica la nd b ehav iou ra ls im i la r i ty a mong e x is t ing e qu id p opu la t ion s ,s ugge s t ing ad eg re eo fg ene t ic i nva r iance a mong a l lm embe r s o ft h eg enu sE quu s( L ev ine 1 982 :224 ) .

b a s is o fh i s tor ica lr e co rd sa nd e x tan tb re ed s . T he i d en t i ty o f t h e se p opu la t ion s i sp rob lema t ic , h oweve r , s i nce G rove s p o in t s o u t t h a t E . f .s y lva t icu s a nd E . f .p r ezwa lsk i a r e mo re o r l e s s t h e s ame t h ing ( Grove s 1 986 :19 ) a nd e l sewhe re , t h a t E . f .p rezwa lsk i i si nd i s t ingu ishab le f r om E quus c aba l lu s ( Grove s 1 974 :38 ; s ee a l so C lu t ton -Brock 1 987 :83 ) . T he i d en t i ty o fT a rpan a s at r ue w i ld d e scendan t o f P le is tocene h o r se i sn ow a l so i n d oub t ( C lu t ton -B rock 1 987 :82 ) .

Membe r s o f t h e g enu s E quu s , i n c lud ing t h e mode rn d ome s t ic h o r se , E quu s c aba l lu s , e vo lved f r om as i ng le c ommon a nce s to r i nt h e L a te P l iocene : P l ioh ippu s (Wa r ing 1 983 ) . Du r ing t h e P le is tocene , r ad ia t ion a nd d i f f e ren t ia t ion o ccu r red , r e su l t ing i nt h ec r ea t ion o ft h e s i x c ur ren t ly r e cogn ised e qu id s pec ie s ( S impson 1 951 ; Wa r ing 1 983 ; N ob is 1 971 a nd Z eune r 1 963 , a s q uo ted i n Wa r ing 1 983 ; G rove s 1 986 ( bu ts e e G rove s 1 974 ) . Du r ing t h eL a te P le i s tocene , e qu id s , a " s ing le , v a r iab le , i n te rb re ed ing g roup " r ad ia ted i n to A f r ica , A s ia a nd E u rope , w he re p opu la t iona l i s o la t ion r e su l ted i n s pec ia t ion ( S imp son 1 951 :26 ) . F o l low ing t h e l a s t G lac ia l p e r iod , e qu id s d i sappea red f r om t h eN ew Wo r ld . T he " t rue h or se " ( Equu s f er n s ) a ppea red i n t h e E u r a s ian l ow land s d u r ing t h is t ime ( Nob i s 1 971 a s q uo ted i n Wa r ing 1 983 ) a nd d i f f e ren t ia ted

P a laeon to log is ts s tudy ing Upper P le is tocene f o s s i l p opu la t ion s i n S ou thwe s te rn E u rope i d en t ify d i f f e ren t e vo lu t iona ry l i ne s f o r e qu id so n t h eb a s is o f g ro s s mo rpho log ica l t r a i t s ( Pra t t 1 976 ; D e lpe ch 1 983 ) . Dur ing t h e Wurm , as i ng le l i ne ( Equu sa f f . s t e inhe imen s is ) r ad ia te s t h roughou t E u ra s ia a nd g ive s r i se t oE quu s f e ru s g erman icu s ( s ucceeded b y EQ UUS f e ru sg a l l icu s ) a nd EQ UU S s t eh l in i ( g iv ing r i se t o E quu s h yd run t inu s ) ( Pr a t t 1 976 ; G rove s 1 986 ) . T he e xac t r e la t ion sh ip b e twe en t h e t wo L a te P le is tocene e qu id s p re sen t i n E u rope , E quu s f e ru s a nd E quus h ydnm t inu s , i s n o t c l ea r e xcep t t h a t t h ey a r e morpho log ica l ly d i s t inc t ( P ra t t 1 976 ; G rove s 1 986 :47 ; C lu t ton -Brock 1 987 ) . T he s pec ie s E quu s f e ru s , s ub sp ec ie s g a l l icu s( P ra t t 1 976 )i st ob et h eo b jec to ft h ep res en t s t udy .

i n to a n umbe r o f d i f f e ren t e co type s ( Zeune r 1 963 ) o r g eog raph ica l r a ce s ( Wa r ing 1 983 ) .

T he d i s t inc t ion b e twe en " b reed s " o fh o r se a nd t r ue s pe c ie s i s n o t a lways m ade c l ea r a nd l e ad s t o

E a r ly t h eo r ie s o f a mu l t ip le o r ig in f o r d ome s t ic h o r se ( e xp la in ing t h e e x i s tence o f t h e d i f f e ren t a nc ien t a nd mode rn b reed s ) h ave , o n t h e w ho le , b een a bandoned i nf a vou ro f am ode lo fd i f f e ren tt y pe sb e long ing t o as i ng le s pe c ie s ( S imp son 1 951 ; N ob is 1 971 , a s q uo ted i n Wa r ing 1 983 ; C lu t ton -B rock 1 987 ) . Howeve r , s ome

c onfu s ion i nt h ed i scu s s ion o ft h e o r ig in o fd ome s t ic h o r se . T he i d en t if ica t ion o fs pec ie s t h rough a ne xam ina t ion o ft h e i r g ros s mo rpho logy i s n o tor iou s ly d i ff icu l t . F ive e x tan t s pe c ie s o f w i ld e qu id a r e c ur ren t ly r e cogn ised : E quu s h em ionu s ; E quu sa s inu s ; E quu sq uagga ; E quu sg revy i ; a nd E quu s z eb ra ( S impson 1 951 ) . I n a dd i t ion t h e re a r e t wo d ome s t ic s pec ie s : E quus c aba l lus a nd E quu s a s inu s .

a u tho r s b e l ieve t h a t t h e d i ve r s i ty o f moder n b reed s i sa

1 4

C lad is t ic a na ly s is o ft h e e qu id s r e su l t s i nt h e i d en t if ica t ion o f t wo d i s t inc t g ene t ic c omp lexe s :

I n t h e c a se o f t h e h or se , c on t ro l led b reed ing d oe s n o t a ppea r t oh ave a l te red t h e g ene t ic s t ock

E quu s a nd A s inu s

( Bene t t 1 980 , a s q uo ted i nG rove s 1 986 ) .

t o a ny g rea t e x ten t ( Lev ine 1 982 ; War ing 1 983 ; B e rge r 1 986 ; C lu t ton -Brock 1 987 ) . O s teo log ica l c hange s wh ich a ccompany d ome s t ica t ion a nd wh ich a r ep robab ly i nd ica tor s

T he s oc ia l s t ruc tu re o fe qu id s l e nd s i t se lf t o t h e c r ea t ion o f d i scre te p opu la t ion s , r e su l t ing i n t h e

o fg ene t ic s e lec t ion a r e n o t d ocumen ted i ne qu id s ( Koh le r R o l lef son 1 989 :142 ,145 ; An thony 1 991 :252 ) . We t h e refore

a ppea rance o f morpho log ica l t r a i t s c ha rac ter is t ic o f l o ca l p opu la t ion s ( e .g . , c oa t c o lor ) .

T h is c ou ld e xp la in t h e

f e e lj us t if ied i no u ru se o fd ome s t ic a n ima ls t oe s tab l ish t h e

a ppa ren t d ive r s i ty o f p hy s ica l

t ype s o f h or se i n t h e

c on tro l s amp le .

p reh is tor ic p a laeon to log ica l a nd a r chaeo log ica l r e cord s , a c cord ing t o War ing ( 1983 :10 ) . E v idence o fp hy s ica l t ype s o f e qu id c oex is t ing i n t h e p a laeon to log ica l a nd

C on t ro ls amp le s pec imen so b ta ined f o rt h is s t udy c ome f r om aw ide v ar ie ty o f s ou rce s , r ang ing f r om

a r chaeo log ica l r e co rd i sn o t , a s G rove s s t a te s , e v idence o f

Nor th Amer ica t o F rance a nd P o land . A l l w e re s t ockya rd

s ympa t ry a nd t hu s o f t h e e x is tence o f s epa ra te s pec ie s

h or se s , h oweve r , a nd a ss uch w e re r a ised o u tdoor s . T h is i s

( Grove s 1 986 :23 ) , s i nce o u r c h rono log ica l c on t ro l i sn o t

r e levan t

f i ne -gra ined e nough t op erm i t t h is a s sump t ion .

d ome s t ica te s h a s o n ly b een p roven t ob e a f fe c ted i nc a se s

s i nce

t h e e xpre s s ion

o f i n c remen ta l

l i ne s i n

o f e x t reme b uf f er ing f r om t h e e nv ironmen t , i . e . , c a se s I ti st r ue t h a tp reh is tor ic a r t is t ic d ep ic t ion s

whe re t h ea n ima ls w e re r a ised i ndoo r so ns t eady d i e t s( Grue

o fe qu id s i n S ou thwe s tern F rance i n d ica te t h e p re sence o f

1 976 ; G rue & J en sen 1 979 ) .

d i f f e ren t morpho log ica lc ha rac te r s ( Grove s 1 974 :180 -181) . N ever the le s s , i ti s f r u i t le s s , i n t h e c on tex t o f t h e p re sen t

D if f e rences i nt h ee nv i ronmen ta lc on tex t s

s t udy , t o a t temp t t o t r ace t h e p o ss ib le d e r iva t ion s o f t h e

o f

g eograph ica lb reed sa t te s ted b o th i n modern p opu la t ions a nd

p opu la t ion s i n S ou thwe s te rn F rance u ndoub ted ly e x is t

t h e

modern

c on tro l

p opu la t ion s

a nd

P len ig lac ia l

i nt h e a r chaeo log ica l r e co rd . A s B e rge r s t a te s : " A l though

( De lpech

t h ey d i f f e r mo s t f r om o ne a no the r i n t h e ir c h romosome s a nd e x te rna l a ppea rance [ . .] a l l s pec ie s a re r ema rkab ly

c on s ide red when i n te rp re t ing t h ec on t ro ls amp le . Howeve r ,

s im i la r i nr e produc t ion , p hys io logy , f e ed ing p a t te rn s , a n t i -

i n cremen ta l g row th r e su l t ing f r om e xp lo i ta t ion o fd i f f e ren t

p reda tory b ehav io r , f i gh t ing s t y le s , s t omach a na tomy a nd

r ange s w i th in t h e s ame c l ima t ic z one h ave n o tp roven t ob e

1 983 ;

i n tra spec if ic

C .O .H .M . A . P. 1 988 ) . T he se mu s t b e

d i f f e rence s

i n

t h e

p a t tern

o f

c yc l ica l

g ene ra l b ehav ior " ( Be rge r 1 986 :14 -15 ) . I n o the r word s ,

s i gn if ican t , i . e . , t h e t im ing o f t h e p r imary i n c remen t s

t h e re e x is t s ab road a dap t ive t ype f o r e qu id s ( S impson

a ppea r s t o b e u naf fe c ted ( Re ime r s & Nordby 1 968 ) . A s

1 951 :217 ; War ing 1 983 ; L ev ine 1 982 ) l i t t le a f f ec ted b y

S p ie ss s t a te s i nh i s d i scu ss ion o fc l ima t ic i n f luence s o n t h e

s e le c t ive p re s sure s d ur ing t h e p roce ss o f d ome s t ica t ion

t im ing o fi n c remen ta l g row th i np opu la t ion so f wh i te t a i led

( C lu t ton -Brock 1 987 :80 ;K oh le r -Ro l lef son 1 989 :145 ) wh ich

d ee r : " The t im ing o ff a l l p hys io log ica l c hange s [ . .] wou ld

w i l l b e t h e o b jec t o fo u r s t udy .

n o t b e e xpec ted t o v a ry s ubs tan t ia l ly w i th p a s t c l ima t ic z ebra

c hange f o r ag iven l a t i tude " ( Sp ie s s 1 990 :31 ) . T he wo rk ing h ypo the s is a dop ted h e re , t h e refo re , i st h a ta n ima ls r a ised i n

b urche l l i a nd o ne o fE quu s f er n s p rzewa lsk i w e re d ona ted

t empera te r eg ion sw i th in t h es ame b road l a t i tude a nd c an b e

b y t h e S an D iego z oo a nd t h e t e e th w ere s e c t ioned f o r

e xpe c ted t op rov ide amode l o f i n cremen ta l g row th u sefu l

c ompa ra t ive p urpose s . T he se s pe c imen s h ave t h ea dvan tage

i nt h e i nve s t iga t ion o ff o ss i l e qu id t e e th f r om S ou thwe s tern

o fw e l l -documen ted l i f e -h is tor ie sa nd d e ta i led p o s t -mor tem s .

F rance .

O ne

h em i -mand ib le

o f

E quu s

C ompar ison o f t h e se e qu id s pe c imen s w i th t h e d ome s t ic h or se c on t ro l s ugge s t s ag r ea t s im i lar i ty i n t h e p a t te rn o f

C on s is ten t

r e su l t s

o b ta ined

f r om

o u r

c emen tum d epos i t ion a mong e qu id s pec ie s ( s e e r e su l t s ,

c on tro l a nd We lsh 's ( 1972 ) s amp le o f t we lve i nd iv idua ls

b e low ) .

f r om t he s ame p opu la t ion s uppor t t h e wo rk ing h ypo the s is

3 .2 E f fe c t s o f t he e nv ironmen t o n t he e xpre s s ion o f

g eograph ica l o r ig in s o ft h e i nd iv idua l s pe c imen s .

t h a t t h e c on t ro l s amp le i s u naf fe c ted b y t h e d i f f e ren t g row th s t ruc ture s . 3 .3 E s tab l ish ing t he c on tro l s amp le . T he a pp l icab i l i ty o f ad ome s t ic c on tro l s amp le t o aw i ld p opu la t ion c an b ed ef ended o nt h eg round s

A s w e h ave a l ready s e en ( Chap te r 2 ,

t h a t t h e e f f ec t s o f d ome s t ica t ion o n t h e f o rma t ion o f

a bove ) i n cremen ta l g row th o f h a rd t i s sue s i n b one a nd

i n c remen ta l g row th l i ne s a ppea r s t ob e m in ima l ( Saxon &

c emen t a ppea r s t o b e r egu la ted o n togene t ica l ly ( K leveza l

H igham 1 969 ; G rue 1 976 ; G rue & J ensen 1 979 ) . S ince t h e u nde r ly ing c ause o fc yc l ica l i n c remen ta l g row th a ppea r s t o

o fd epos i t ion o ft h e g row th i n cremen t s mu s t b e e mp ir ica l ly

b e

g ene t ic ,

t h e

e nv ironmen t

t h e

e s tab l ished f o r t h e t a rge t s pec ie s . T he refo re , ac on t ro l

p er iod ic i ty a nd t h e e xp re s s ion o f t h e i n cremen t s i . e . , t h e ir

s amp le mu s t b e e s tab l ished . T he s t ruc tu re a nd p os i t ion o f

c on t ra s tedne s s

t h e f i na l g row th i n cremen t ( t he o ne f o rm ing a tt h e t ime o f

( s ee

d i scus s ion

m e re ly

i nf luence s

1 970 ; C a s tane t 1 982 ; i n te ra l i i ) . T h is m eans t h a tt h ep a t tern

i n C hap te r

2 ) .

Whe re

d ome s t ica t ion h a s r e su l ted i ng ene t ic s e lec t ion , h oweve r , i t

d ea th )

i sh ypo the t ica l ly p o ss ib le t h a t e ndogenou s r hy thm ic i ty h a s

s pe c imens o f k nown d a te o f d ea th . I d ea l ly , af u l l , y ea r ly

b e en a f f e c ted a nd w i l l d i f f e r f r om t h a t e ncoun te red i nw i ld

c yc le o fg row th s hou ld b er ep re sen ted i nt h ec on t ro l s amp le

p opu la t ion s o ft h e s ame s pe c ie s .

a nd i nd iv idua ls f r om d i f f e ren t a ge c l a s se s s hou ld a l so b e 1 5

i s o bserved a t d i f f e ren t t ime s o f t h e y ea r i n

p re sen t . T h is a l low s f o r mo rpho log ica l d i f f e rence s i n c emen tum d epo s i t ion b e tween t h e a ge c l a s se s t o b e

i n c i so r sa r eo fd i f f e ren tt h ickne s sa nd y i e ld d i f f e ren tr e su l t s . B ecau se o ft h e t h inne s s o fc emen tum d epo s i ts o n i n c isor s , m o la r sa nd p remo la r sa r ep ref e r red i nt h is a na lys i s( s ee a l so

d e te c ted , s pec if ica l ly d i f f e rence s i n t h e a moun t o f t i s sue d epo s i ted ( Ca s tane t 1 982 :50 ; S t a l l ib ra s s 1 982 :119 ) t h e t im ing o fd epo s i t ion ( K levez a l p e r s . c omm . 1 991 ) a nd t h e

M i tche l l 1 963 ) . D e c iduou s t e e th e xam ined p re sen t al a rge n umbe r o fa nnu l i 'i nc em en t , b ea r ing n o r e la t ion sh ip t ot h e a ge o f t h e a n ima l , m i lk t e e th a r e t h e ref ore a l so om i t ted f r om t h e s t udy .

n a tu re o ft h e t i s sue s d epo s i ted . I n t h e c a se o f modem E quu s. c aba l lu s , h oweve r , d i ff icu l t ie s a r ise i n t h e a cqu is i t ion o fs pe c imen s .

Mand ibu la r mo la r r ow s ( r igh t s ided ; s e e P la te 3 . 1 ) w e re s l ow ly d ef le shed i n ah ea ted s o lu t ion o f

Nor th Ame r ican s ou rce sw e re t a pped f i r s t ,t h e mo s tr e l iab le s ou rce b e ing t h e N ew B o l ton C en te r , a p a r t o f t h e

w a te r a nd c hem ica l e nzyme s ( c omm e rc ia l ly a va i lab le i n s ome d e te rgen t p owde r s ) . T he s o lu t ion w a s n o ta l lowed t o c ome t o ab o i l , i n o rde r t op reserve t h e c emen tum f r om t h e rma l s hock s . N ex t , m and ib le sw e re d ipped i ns ucce s s ive

v e te r ina ry s choo l o ft h e Un ive r s i ty o fP enn sy lvan ia . I tw a s o r ig ina l ly i n tended t h a t 3h o r se s d ecea sed t owa rd s t h e 1 5 th o fe ach mon th b ea cqu ired o ve rt h ep e r iod o fo ne f u l ly ea r . Unf or tuna te ly , d i ff icu l t ie s a rose m ak ing i ti mpos s ib le t o c o l lec tt h e e xpe c ted n umbe r o fa n ima l s - d u r ing t h e mon th s

b a th s o f e t hy l a l coho l ( i nc rea s ing c oncen t ra t ion b y 1 0% i n c remen t s u n t i l 1 00 % s o lu t ion r e ached ) t od ry a nd f i x t i s sue s . T e e th c hosen f o r s t udy w e re r emoved f r om t h e

o fJ u ly ,D ecembe r , J anua ry , Ma rch a nd A pr i ln os pec imen s w e re c o l le c ted .

m and ib le s w i th a d jacen t b one ( a lveo la r w a l l s ) i n ta c t u s ing ah and saw . E rup t ion /wea r s t age i n f o rma t ion a nd c rown

Ho r se s d e s t royed a t t h e N ew B o l ton C en te r o r s en t f o r p o s t -mor tem c ome f r om aw ide v a r ie ty

h e igh t sw e re r e co rded i no rde r t oe s t ima te i nd iv idua la ge a t d ea th ( f f . L ev ine 1 982 ) . An umbe r o ft e e th w e re p repa red a nd v i ewed p e r m and ib le u s ing av a r ie ty o ft e chn ique s . S ee T ab le 3 . 1 .

o f s ou r ce s . A l though t h is r ep re sen t s a l o s s o f s t anda rd i sa t ion , t he v a r iab i l i ty t hu si nduced i sp robab ly n o t v e ry s i gn if ican t( s ee S ec t ion 3 .2 -E f f e c to ft h eE nv ironmen t o nt h eE xp re s s ion o fG row th S t ruc tu re s , a bove ) . T he a ge o f

3 .4 .1 Unde ca lc if ied T h in S ec t ion s .

t h e h o r se s o b ta ined f r om t h e N ew B o l ton C en te r w a s n o t k nown ; a n e rup t ion a nd u se -wea r t a b le f o r mo la r s a nd p remo la r sw a su sed t oe s t ima te a ge a td ea th ( L ev ine 1 982 ) .

P e t rog r aph ic t h in se c t ion sw e re p roduced u s ing amod if ied v e r s ion o fs t anda rd m e thodo logy ( s ee G o rdon 1 988 ; CMRAE L abo ra tory

I n o rde r t oc omp le te t h e c on t ro l s amp le , a n a ba t to ir i nB o rdeaux , F r ance , w a s c on tac ted . Mon th ly t r ip s t oc o l le c t f r e sh ly s l augh te red a n im a ls w e re a r r anged . O ne t o t wo s pe c imen s p e r mon th w e re c o l le c ted i n

P rocedu re 1 988 ) . O cc lu sa ls u rf ace s ( we l la bove t h eg i ng iva ) a nd r oo t t i p sw e re s e c t ioned of u s ing a h i gh speed d i amond s aw i no rde r t od ebu lk t h e s pe c imen s . T he t e e th w e re p l aced i nr i g id p la s t ic t r ay s a nd l a be l led . N ex t , t h ey w e re i mpregna ted i n at wo -pa r t r oom cu r ing e poxy l i qu id r e s in c omme r c ia l ly a va i lab le a s S co tchca s t B rand E lec t r ica l R e s in 5 (5235 ) . T he s amp le s w e re p l ac ed i n av a cuum c hambe r f o r a pp rox ima te ly 1 5 m inu te s ( a t 0 Mba r s ) w h i le t h e e poxy w a s s t i l l p o lyme r i s ing i no rde r t o i n su re p rope r p ene t ra t ion o ft h e r e s in . T he r e su l tan t b lock o f r e s in w a s d ebu lked w i th a h igh speed s aw t o f a c i l i ta te h and l ing .

Novembe r , D ecembe r , J anua ry a nd A p r i l . T he a pprox ima te a ge o f t h e a n ima ls w a s e s t ima ted u s ing L ev ine 's e rup t ion /wea rt a b le sa nd c rown h e igh tt a b le s( Lev ine 1 982 ) . A n ima ls d e s t royed a tt h e a ba t to ir i nB o rdeaux a r e s h ipped l i ve f r om aw ide v a r ie ty o f s ou rce s , i n c lud ing P o land a nd t h e Un i ted S t a te s . T h re es amp le sw e re c o l le c ted w i th t h ea i d o fS .P re s ley , A rchaeo log ica l S urvey so fC anada . A t temp t s t o o b ta in s amp le s f r om b reede r s i n t h e P y r6n6e s , whe re s t ock i sn orma l ly k ep ti ns em i -w i ld c ond i t ion s , h ave b e en

I n t h e f i r s t s t age s o f t h e s t udy , t e e th e mbedded i n r e s in b lock s w e re c a ref u l ly o r ien ted a nd

u n succe s sfu l . I n s um , o ve r t h e c ou r se o f t h ree y ea r s e i gh teen a n ima ls o fk nown d a te o fd ea th w e re c o l lec ted ( 1 6 E quu s c aba l lu s , 1E quu s f e ru s p rzewa lsk i a nd 1 E quu s z ebra ) , r ep re sen t ing a9 -mon th c o l le c t ion p e r iod . I na dd i t ion

s e c t ioned l o ng i tud ina l ly ( bucco l ingua l ly )u s ing al ow speed d i amond s aw ( Bueh le r I s ome t ) . S ee F igu re 3 . 1 . T he c u t m u s t b e p e rpend icu la r t o t h e c emen tum s u r f ace s i nce t h e t h ickne s s o fi n c remen t sw i l lv a ry i fv i ewed a ta n a ng le a nd b e cau se a nnu l i m ay a ppea r d oub le d ue t oa n o p t ica l e f ec t i fv i ewed o n as l an t( Pa s ca l 8 4Q ur6 1 983 ) . F o rt h i sr e a son i n fund ibu lae mu s t b e a vo ided s i nce a r e curv ing s u rf ace i s t o o d i ff icu l tt oa l ign p e rpend icu la r i ly .

t o t h e se a n ima ls , t oo th s amp le s f r om t we lve a n ima ls o f k nown s e a son o fd ea th , p rev iou s ly s t ud ied u nde r r e f le c ted l i gh tb yD . We lsh ( 1975 ) , a r e i n c luded i nt h ef i na la na ly s is . PART B . ANALYSIS OF GRO WTH S TRUCTURES I N MODERN EQU ID TEETH .

2 . T he e xpo sed f a ce o f t h e m e s ia l h a lf o f t h e t o o th w a s p o l ished w i th s ucce s s ive g r i t s i z e s t o r emove g r ind ing m a rk s .

3 .4 .0 Me thod s a nd Ma ter ia l s . Ho r se s a r e h e te rodon t ; t h e ir mo la r s a nd i n c iso r s a r e a na tom ica l ly d i s t inc t . I nc i so r s h ave v e ry t h in c emen tum d epo s i t s wh ich a r e d i ff icu l tt oi n te rpre t ( s e e a l so L eade r -W i l l iam s 1 979 ) . L ab ia l a nd l i ngua l c emen t p ad s i n

3 .T he p o l ished f a ce w a s g lued t o as l ide w i th a r a ld i te o r , a l te rna t ive ly , w i th a1 : 5 p r epa ra t ion o ft h e S co tchca s t r e s in ( f o r b e t te r a dhe s ion o ft h e s pec imen d ur ing h and l ing ) . 1 6

Modern

Sample

#

Control

TABLE 3 .1 Sample: Preparation

Transmitted Light

Methods

Used.

Stained Histological

S . E. M.

NB

1 .1

Ml, M2

NB

1 .2

Ml, M2

P2

NB

1 .3

P3, P4

P4, M3

NB

2 .1

M1

NB

3 .1

M1

NB

4 .1

M1

NB

4 .2

M1

NB

4 .3

Radiography

*unreadable

ASC

1

Il

ASC

2

Il

ASC

5

I l

CON

20

Ml, P2, P3

M2

CON

21

Ml, P2, P3

M2

CON

22

Ml, P2, P3

M2

CON

2 3

Ml, P2, P3

M2

CON

2 4

Ml, P2, P3

M2

CON

25

Ml, P2, P3

1 7

M1

F igur e 3 .1 Or ien ta t ion o ft he t oo th s e c t ion s .

4 . T he g lued s e c t ion w a s moun ted i n as l ide h o lde r a nd t h inned u s ing al ow speed s aw t o at h ickne ss o f 01 mm .

5 .v i ewed w i thou t s l ide c ove r , u nde r at h in f i lm o fw a te r . 6 .A sa bove , m ic ropho tog raph sw e r et a ken o fe a ch s amp le .

5 .T he s l ide w a st h en g round t o at h ickne s so f 040 m ic ron s o n s ucce s s ive ly f i ne r g r i t a b ra s ive s u s ing a M in ime t p o l ishe r .

T ee th f r om s i x i n d iv idua ls w e re p r epa red i nt h is w ay . 3 .4 .2 S cann ing E lec tron M icro scopy .

6 .T he s l ide w a sc he cked u nde r t r ansm i t ted , p o lar ised l i gh t b efo re a pp l ica t ion o f as l ide c ove ru s ing F i she r 's P ermoun t .

Two s amp le s w e r e p repa red f o r s c ann ing e l ec t ron m ic ro scopy . T he mo s t r e l iab le p repa ra t ion o f a s e c t ioned s u rf ace w a s a s f o l low s :

M icropho tograph sw e re p roduced f o re ach s pec imen ( u s ing K odak P lu s -X P an 1 25 A SA 3 5nun f i lm ) .

1 .E ach t o o th w a ss e c t ioned r a d ia l ly ( b ucco l ingua l ly )u s ing al ow s peed d i amond s aw ( Bueh le r I s ome t ) .

As e cond p repa ra t ion m e thod e vo lved a s t h e s t udy p rog re ssed . T ee th w e re e mbedded a s a bove , f o l low ing wh ich as e r ie s o fp a ra l le l r a d ia l c u t s w e re m ade

2 . T he s pe c imen w a s t h en g round b y h and o n as t r ip

i ne ach t oo th , 2mm a pa r t , u s ing ah igh speed d i amond s aw w i th a 5 " b l ade , f i t ted w i th a s pe c imen -ho lde r a nd

g r inde r ( Hand ime t 1 ,B ueh le r ) o n p rogre s s ive ly f i ne r g r i t a b ra s ive s w i th a r e c t i l inea r m o t ion . T h is t e chn ique p red ic tab ly p a t terned p r epa ra t ion a r t if ac t s i n t h e l i ne o f m o t ion .

c a l l ib ra ted g u ide . 2 .T he s e c t ion s w e re p o l ished o n o ne s i de o n p rog re s s ive ly f i ne r g r i t a b ra s ive s u s ing al a p -whee l .

3 . T he f i na l s u r f a ce w a s p o l ished f o r am in imum o f t wo m inu te s w i th a c l o th l a pwhe e l u s ing a0 .06 m ic ron s i l ica s l u r ry .

3 .P o l ished f a ce sw e re g l ued t o7 .5nun x4 .0mm s l ide sw i th E po tek 3 01 , at wo -pa r t r e s in p roduced b y E po tecny .

4 . T he s pe c imen w a s t h en r i n sed w i th d i s t i l led w a te r a nd 4 . T he e xposed f a ce w a s g round b y h and o n al a p -whe e l u n t i l t h e t h in -se c t ion r e a ched 040 m ic rons , p o l ished w i th af i ne a b ra s ive ( s i l ica s l ur ry ) a nd ,

c he cked u nde r a n i n ve r ted o p t ica l m icro scope ( Re iche r t , A u s tr ia ) a t l ow m agn if ica t ion ( 10x ) f o r t h e p re sence o f p o l ish ing a r t if a c t s . I f s c ra tche s w e r e s t i l l v i s ib le t h e p o l ish ing p rocea lu re w a sr e pea ted u n t i lt h es u r f a ce w a s f r e e 1 8

o f s c ra tche s .

t n ic ro tome ( Le i tz K ryoma t 1 700 ) . A s e r ie s o f p a ra l le l s e c t ions , 1 8 m icrons t h ick , w e re p roduced p e r s pec imen .

5 . T o p roduce t opograph ica l d e ta i l t h e s u rf ace o f t h e s pec imen w a s e t ched w i th a 0 .7252 g / 100m 1 s o lu t ion o f

3 .T he s e c t ion s w e re c o loured i nE h r l ich 's h ema toxy l in f o r

p hosphor ic a c id a s s ugge s ted b y G an t t ( 1979 ) f o r t wo

1 5 m inu te s a nd r i nsed i nt a p w a te r .

m inu te s . T he s pec imen w a s t h en r i nsed i n d i s t i l led w a te r a nd t h en a b so lu te e t hano l b efore b e ing a i r -dr ied .

4 .

S l ide s

w e re

p repa red

u s ing

A quamoun t

moun t ing

m ed ium . 6 . T he s pe c imen w a s t h en moun ted o n a l um inum S EM moun t ing s t ud s w i th l ow -re s is tance c on tac t c emen t ( Erne s t

5 . O b serva t ion s w e re m ade u nde r o rd ina ry t r ansm i t ted

F u l lum I n c .) a nd s pu t te r c oa ted w i th g o ld ( PS -2 C oa t ing

l i gh t .

Un i t , I n terna t iona l S c ien t if ic I n t s trumen ts ) t oe nhance S EM v i ew ing .

3 .4 .6 I mage Ana lys is .

3 .4 .3 M icrorad iography .

I n cremen ta l g row th i s m ea su red f r om i mage s p roduced f r om s c anned m icropho tog raphs ( s ee

Undeca lc if ied

t h in

s e c t ions

f r om

a ppend ix A ) . T he a moun t o f g row th i s e xp re ssed a s a

o ne

modern a nd t h ree f o ss i ls pe c imen sw e re m ic rorad iographed .

p e rcen tage o f t h e t o ta l g row th e xpec ted , b a sed o n t h e o b served w id th o f t h e p rev iou s ( p enu l t ima te ) i n cremen t

1 . F ree t h in s e c t ions w e re p roduced u s ing a h igh -speed

( Gordon 1 989 ; S p ie ss 1 990 ; P ike -Tay 1 991 ; i n te r a l i i ) .

d i amond s aw a nd t empo rar i ly f i xed t os l ide s u s ing C anada

A rea m ea su remen t s o ft h e f i na l a nd p enu l t ima te i n cremen t s

B a lsam a nd p o l ished t o at h ickne ss o f 0200 m icrons .

w e re c a lcu la ted u s ing i mage p ro ce s s ing s of twa re ( Image 1 . 23) . I n c remen t s w e re c oun ted a nd m ea su red o n

2 . S pec imen s w e re d e tached f r om t h e s l ide s i n at o luene b a th ( 3 h our s ) .

h i s togram s ( op t ica l d ens i ty p rof i le s ) b a sed o n a ve rage p ixe l l um inance v a lue s f o r 8 -p ixe l w ide c ros s s e c t ions o f t h e

3 . F ree t h in s e c t ions w e re p o s i t ioned o n K odak SO -343

a L

p ho tog raph ic f i lm , 4 0cm f r om t h e r ad ia t ion s ource a nd

h i s togram s p e r i mage .

s c anned i mage s n orma l t ot h e t o o th s u rf ace ( L ieberman 1 990 ) .

R e su l ts w e re a ve raged

e

f r om t wo t o t h ree

e xposed t o 4 0 K vo l ts a t 1 5 mA f o r 3 0 m inu te s . T he f i lm w a s d eve loped .

3 .5 .0 R e su l t s .

4 . M ic ropho tographs o ft h e e xposed f i lm w e re t a ken u nde r

R e su l t s

t r an sm i t ted o rd ina ry l i gh t a tl ow m agn if ica t ion .

o f

t h e

d i f f e ren t

m e thods a r e p re sen ted i n T ab le s 3 .2 ,

p repara t ion

3 .2b a nd 3 .2c

( b e low ) . 5 . O b se rva t ion s w e re made o f t h e m icropho tograph s a nd c ompa r ison s m ade w i th a ) u nde ca lc if ied t h in s e c t ion s

3 .5 .1 S cann ing E lec tron M icroscope .

p roduced f r om t h e s ame t oo th b )d e ca lc if ied , s t a ined f r ozen t h in -se c t ion s ( s ee b e low ) f r om t h e s ame i n d iv idua l .

I n cremen ta l s t ruc ture si ne qu id c emen ta r e v i s ib le u nde r S EM a f te r a pp ropr ia te s amp le p repa ra t ion . S e e P l a te 3 .2 . A c id e t ch ing p roduced t opog raph ica l d e ta i l

3 .4 .4 R ef le c ted L igh t .

i nt h e c emen tum o fh or se t e e th b y more d e ep ly e t ch ing t h e F or ty -four

t e e th ,

r ep resen t ing

t we lve

a nnu l i ,

i . e . , t h ose i n c remen t s w i th a h ighe r i n organ ic

a n ima ls , w e re o b ta ined f r om D . We lsh f r om h i s ( 1975 )

( m ine ra l ised ) c on ten t r e la t ive t o t h e o rgan ic c omponen t

c on tro l s amp le f o r v i ew ing u nde r r ef le c ted l i gh t . T he t e e th

( c o l lagen f i b re s a nd c e l l c on ten t ) .5

w e re p rev ious ly e mbedded i nb lock so fr e s in , s e c t ioned a nd R e su l ts u s ing S EM a re d i re c t ly c ompa red

p o l ished u s ing al a pwhee l (We lsh 1 975 ) .

w i th t h in -se c t ions f r om t h e s ame s pec imens v i ewed u nde r 1 . A l i gh t s ource i s p os i t ioned a t a n a ng le t o t h e t oo th

t r ansm i t ted , p o la r ised l i gh t . T he c emen to -ename l j unc t ion i s c l ea r ly v i s ib le a s as e r ie s o f r e sorp t ion b ays o n t h e

s u rf ace .

s u rf ace o f t h e e name l , a ccomoda t ing a p r ima ry l a ye r o f c emen t ( s e e P la te 3 . 3 )6.

2 . O bserva t ions a r e m ade u nde r b inocu la r m agn ify ing g l a sse s a tl ow m agn if ica t ion ( 10x ) .

3 .5 .2 H is to log ica l P repara t ion s . 3 .4 .5 D e ca lc if ied , S ta ined Th in S ec t ion s . T he v o lume a nd f o rm o fc emen tocy te s i n 1 .M , s f r om 5 s pe c imens w e re d e ca lc if ied u s ing a 5% s o lu t ion o f HNO3 f o r a bou t 3d ays , a nd t h en i mme r sed i n

t h ep r ima ry i n cremen t ( t he z one i mmed ia te ly a d jacen tt ot h e e name l /cemen t b oundary ) a r e d i s t inc t a nd i n d ica te a

a7 0% s o lu t ion o fa l coho l f o r 1d ay a nd r i nsed i nd i s t i l led

p robab le r ap id d epo s i t ion . T he LAGs s t a in d a rk ly , i . e . , a r e

w a te r .

h ema toxy l inoph i l ic , a nd c an b e f o l lowed a l ong t h e who le l e ng th o ft h e t o o th . C emen tocy te s a nd r e sorp t ion b ays a l so

2 . F rozen t h in -sec t ions w e re p roduced u s ing a c ryos ta t

s t a in mo re d a rk ly a nd a r e r e la t ive ly n umerou s w i th in t h e 1 9

Table 3 .2 Control Sa m ple: Results under Transmitted L ight ( Provenience/Sample number/Month of death/Estimated age**/teeth prepared a s thin s ections/# increments/Identity , f inal inc./% growth o f l ast growth z one, % of previous z one) Provenience Sample

Mo.

Age

Pos.

Bordeaux

CON 2 3

1

6

M1 P 2

9 9

l ag zone*

6 7.1 2 0.1

Bordeaux

CON 2 4

1

9 .5

M1

9

lag*

3 1

P enn.

NB 4 .1

2

5

M1

zone

3 3.1

P enn.

NB 4 .2

2

1 2+

M1

1 0-11

lag*

1 6

A .S.C.

ASC 1

5

n /a

I l

6

z one

4 0.6

A .S.C.

ASC 2

5

n /a

I i

2

z one

6 2.8

Bordeaux

CON 2 5

5

4

M1

6

l ag

1 15

Penn.

NB 1 .1

6

2

M1 M2

6 -8 8

z one z one

7 3.2 8 0

Penn.

NB 1 .2

6

2

M1 M2

5 4 -5

z one z one

1 00 1 5

Penn.

NB 1 .3

6

1 3

P 3 P 4

6 -8 8

z one z one

5 0+ 4 3.5

San D iego

P .horse

7

2 .5

I l 1 2

6 5 -6

z one z one

n/a 4 9.9

A .S.C.

ASC 5

8

n /a

I i

5

z one

4 3.8

P enn.

NB 2 .1

1 0

2 .5

M1 P 4

6 8 -9

z one z one

5 0.6 1 00

P enn.

NB 3 .1

1 1

3 .5

M1 M2

6 4

l ag z one*

1 00 1 00

Bordeaux

CON 2 0

1 1

9

M1 P 3 P 2

6 5 -6 8 -9

l ag l ag l ag

7 4 1 00 6 2

Bordeaux

CON 2 1

1 2

7

M1 P 3

1 0 7 -8

l ag l ag

1 03.5 9 5

Bordeaux

CON 2 2

1 2

1 0

P 2

1 0

l ag

1 01

* Unclear.

* * E stimated age,

2 0

# Inc.

1 1

I .D.

Percent

eruption/crown height.

Table 3 .2b Results under r eflected l ight Samples provided by D . Welsh.

( Sample provenience/Specimen number/Sample number/Month o f death/Identity of f inal increment) Provenience Sable S I

I sland S I

I

S e

I

e l

Specimen # E -2-4 e l

Sample # 3 93 3 94

W-16-3

3 95 3 96

M-16-2

3 97 3 98

M-13-4a

3 99 4 00

M-9-11

4 01 4 02

M-15-3

Month 3

_5 1

4 03 4 04

M-9-20

4 05 4 06

M-9-18

4 07 4 08

M-12-2

4 09 4 10

W-18-7

4 11 4 12

M-9-12

1 -12**

M-11-2

1 3-24**

ID Final LAG*

S I

5

S I S I

* LAG f inal with intermittent ( M2. -M2 more r ectangular cement i slet between metastyle and entoconid s horter, wider i n Ml. sharper curvature to M 2 r oot.

-M2 more r ectangular -M2 more s harply curved r oot a nd o cclusal s urface.

-PM vestibular f old more marked -PM l ingual f old wider, more developed ptychostylid. protocone f latter i n M . -M root angled towards f ront o f mouth. -PM parastyles and metastyles thicker than i n M .

0 Z

the angle o f the o cclusal s urfaces viewed l ingually ' s o pposite i n P 3 & P 4. P 3 " squarer" than P 4.

PREMOLARS

c n

t i t 0 . 4 4 ) i n t 0 U )

Figure Dental

5 .1

Measurements

1 1 1

5 3

Recorded.

d 'Aqu i ta ine ) w e re s amp led . R e su l t s f r om a s i ng le l e g ib le s pec imen o u t o f at o ta l o f f i ve t e e th i nd ica te s s u mme r ( May -Oc t ) e xp lo i ta t ion o fh o r se .

m ic roscope , u s ing t h e d en s i ty p rof i le s a s a n i n te rp re t ive g u ide . 5 .3 .0 R e su l t s .

C ap B lanc : 5 .3 .1 C ompara t ive s tudy . L eve ls

a t t r ibu tab le

t o

t h e

L owe r

Magda len ian ( Rou sso t 1 972 ; p e r s .conun . 1 990 ) f r om t h e P eyr i l le ( a ka . L a lanne ) a nd R ou s so t c o l le c t ion s w e re s amp led . B o th w arm ( may -O c t ) a nd c o ld s ea son p re sence i s

Unde r r e f le c ted l i gh t , o p t ica l c on t ra s t i n t h e f o s s i l t e e th w a sv e ry w eak a nd e ven n on -ex is tan t i nt h e c a se o f s t a ined t e e th . C on sequen t ly , u se o f r ef le c t ed l i gh t w a s n o t d eve loped f u r the r . R e su l t s f r om m ic ro rad iog raphy

i n d ica ted . C o ld s e a son o ccupa t ion p robab ly o ccur red d ur ing D e c Jan o n ly .

w e re c ompa rab le ; g row th l a ye r sw e re v i s ib le ( t hough al i t t le d i f fuse ) i nt h e r a d iog raph s b u t v i s ib i l i ty i ns t a ined s amp le s w a s r e la t ive ly p oo r ( s e e P la te 5 . 1) .

B adegou le :

5 .3 .2 T ran sm i t ted L igh t .

L eve ls a t t r ibu tab le t o t h e L owe r Magda len ian f r om t h e c o l le c t ions h ou sed a t t h e Muse e N a t iona l d e s E yz ie s w e re s amp led . F ou r t e e th ( a l l l e g ib le ) i n d ica te b o th w arm ( May -Oc t ) s e a son a nd c o ld ( Dec -Ap r ) s e a son p re sen ce a tt h e s i te .

G row th l a ye r s w e re c l ea r ly v i s ib le u nde r t r an sm i t ted p o la r ised l i gh t i n t h e f o s s i l p repa ra t ion s ( s ee P la te 5 .2 ) . r ep roduced C hap te r 3 , t wo b road ,

R e su l t s o f t h e i n d iv idua l f o s s i l s amp le s a r e i n A ppend ix C . A s a l r eady d i scu s sed ( i n a bove ) s ea sona l d e te rm ina t ion s a r e l im i ted t o 6 -mon th c a tegor ie s r ough ly e qu iva len t t oc o ld

L auger ie H au te E s t : T he P ey rony c o l lec t ion , ( Mu s6e N a t iona l

s ea son /warm s e a son , i . e . , N ovembe r -Apr i l a nd May O c tobe r , r e spec t ive ly . T he c o ld s e a son c an b e f u r the r

d e P rA is to ire d e s E yz ie s ) , a nd t h e B o rde s c o l lec t ion ( I n s t i tu t d u Q ua ter n a i re , B o rdeaux ) w e re s amp led . T e e th f r om t h e P ey rony s e r ie s a t t r ibu ted t o Magda len ian I I ( S .

s ubd iv ided i n to : N ovembe r ( " au tumn " ) ; D e cembe r Janua ry ( "w in te r " ) a nd F ebrua ry -Apr i l ( " la te w in te r " "spr ing " ) . I t

Made le ine , p e r s .c omm . 1 990 )i nd ica te p redom inan t ly w a rm

i su nde r s tood t h a tt h ese c a tegor ie s d on o tn ece s sar i ly r ef le c t a c tua l " s ea son s " e x i s t ing d ur ing t h e P len ig lac ia l i nt h e s t udy r e g ion .

s e a son o ccupa t ion , w i th a s i ng le t oo th i n d ica t ing D e c Jan . r e s idence . R e su l t s f o rc omb ined L owe r Magda len ian l e ve ls ( Mag .0 I I I ) f r om t h e B o rde s s e r ie s i n d ica te b o th D ec -Ap r . a nd May -Oc t s e a son r e s idence . T he c omb ined r e su l t s o f

T he r a te o fa t t r i t ion r e su l t ing f r om p oo r ly

b o th c o l le c t ion s i n d ica te s t h a t L auger ie H au te E s t w a s p o ten t ia l ly o ccup ied a ta l l t ime s o ft h e y ea r .

p rese rved s pe c imen s i sr e la t ive ly l ow o ve ra l l : 2 5% o f t h e t e e th d id n o t p roduce l e g ib le s l ide s . C emen tum g row th l a ye r s w e re o b se rvab le i nt h eo the r 7 5%o ft h e s pe c imen s .

S a in t -Germa in la -r iv ire : 5 .3 .3 S ea sona l d e term ina t ion s f or e ach s i te b a sed o n d en ta l i ncremen ta l a na ly s is o f h or se r ema in s .

S amp le s w e re o b ta ined f r om l e ve ls c . 1 , c . 3 a nd c . 4 o ft h eG . T re co l le e xcava t ion s , s e r ie s h oused a t t h e I n s t i tu t d u Q ua ter na ire , B o rdeaux . T he t e e th a t te s t t o w arm s e a son ( May -Oc t ) e xp lo i ta t ion o fh o r se w i th as i ng le

T h i s s e c t ion p rov ide s as i te b y s i te s umma ry i f s e a sona l d e term ina t ion s r e su l t ing f r om c emen tum a na ly s is o ff o s s i l h o r se t e e th , c a rr ied o u t a bove ( Tab le 5 .3 , a nd s e e A ppend ix C ) . T he a r chaeo log ica l i mp l ica t ion s o f t h e se r e su l ts w i l lb ed i scu ssed i nc omb ina t ion w i th o the rs e a sona l d a ta f r om e a ch o ft h e s amp led s i te s ( b e low ) .

f a l l ( Nov . ) k i l l . R oc d e Ma r camps : B o th l o c i w e re s amp led ( Ro c d e Ma r camp s 1a nd 2 )f r om t h e M . L eno ir e xcava t ions , s e r ie s

L a Made le ine :

h oused a t t h e I n s t i tu t d u Q ua terna ire , B o rdeaux . R e su l t s i n d ica te y ea r r ound , i . e . , May -Oc t . , Novembe r a nd D e c -

T he s amp le o fh o r se t e e th o b ta ined f o rt h e p re sen t s t udy o r ig ina te s f r om B ouv ie r 's L owe r Z one , h or izon 2 7 ( a t tr ibu ted t o t h e P len ig lac ia l , D e lpe ch p e r s .conun . 1 990 ) . P redom inan t ly w arm s e a son e xp lo i ta t ion o fh o r se ( May -Oc t ) i si nd ica ted , w i th D e c Jan . e xp lo i ta t ion

A p r . , e xp lo i ta t ion o fh o r se . Mou l in N euf :

a l so i nd ica ted .

E v idence f r om l e ve ls 2 a nd i n f . i n d ica te s p o ten t ia l y e a r r ound e xp lo i ta t ion o f h o r se w i th o ne N ov . ,

R e ignac :

o ne D ec Jan . , o ne F eb -Ap r . a nd t wo May -Oc t . d ea ths . .

L eve ls a t tr ibu ted t o t h e L owe r Magda len ian ( Rou sso t p e r s .comm . 1 990 ) f r om b o th t h e Hu ina rd a nd R ou s so t c o l le c t ion s ( hou sed a t t h e Mu s&

D uru thy : T he A rambou rou c o l lec t ion sh oused a tt h e

5 4

T ab le 5 . 3 S yn the t ic R e su l t s f or t he F o s s i l S amp le . S ea sona l D e term ina t ion s

S i te N ame

N

N.

L auger ie a

1 3

5

L auger ie b

" Summer " May -Oc t

N ov

N=2

0

2

1

3

D ec -Jan

F eb -Apr

"W in ter " N ov -Apr

7

6

0

1

0

1

2 2

1 2

8

0

3

1

4

B adegou le

4

4

1

0

2

1

3

Du ru thy 4

1 3

1 0

7

0

1

2

3

Duru thy 5

5

4

4

0

0

0

0

Made le ine

9

7

6

0

1

0

1

1 4

1 4

7 ( 8 )

1

4

1

6

E nva l

4

3

2

0

1

0

1

E sc lauzu r

4

3

3

0

0

0

0

1 0

5

3 (4 )

1

0

0

1

Mou l in N eu f

5

5

2

1

1

1

3

Marca n ip s 1

8

6

3

1

2

0

3

Ma r camp s 2

3

2

0

0

1

1

2

Mar camp s t

1

8

3

1

3

1

5

R e ignac

5

1

1

0

0

0

0

C ap B lanc

3

2

1

0

1

0

1

L auger ie t

R ond d u B arry

S t G e rma in

N = Numbe r o fs amp le s . N * =N umbe r o fl e g ib le s amp le s . L auger ie a = L auger ie Hau te E s t , B orde s c o l le c t ion . L auge r ie b = L auger ie H au t e E s t , P eyrony c o l le c t ion . Mar camp s 1& 2 = R oc d e Ma r camp s l o cu s 1& l o cu s 2 . ( For i nd iv idua l s amp le r e su l t s s e e A ppend ix C )

5

R e serve A rcheo log ique d 'Ar thous , w e re s amp led . L eve l 4 ( Mag .IV / p os t -P len ig lac ia l )y i e lded e v idence o f May -Oc t . ,

5 .5 Conc lu s ion : App l icab i l i ty o f S ke le tochrono logy t o F os s i l Hor se T ee th .

D ec Jan . a nd F eb -Ap r . p re sence . L eve l 5 ( Mag .I I I / P len ig lac ia l ) y i e lded e v idence o f w arm (May -Oc t ) s e ason

I n c onc lu s ion i ti sp os s ib le t o s t a te t h a t s ke le tochrono logy i s a v a l id t e chn ique f o r o b ta in ing

p re sence o n ly .

d e term ina t ion s o f s e a son o f d ea th f r om f o s s i l e qu id t e e th . T h in s ec t ion s v iewed u nde r t r ansm i t ted , p o lar ised l i gh t

R ond d u B ar ry :

p roduced t h e b e s t r e su l ts . T he a r chaeo log ica l s i gn if icance Ho r se t e e th f r om R ond d u B a r ry , f ou i l le s

o f t h e s ea sona l d e term ina t ions o b ta ined f r om t h e f o s s i l

R . d e B ay le d e H ermen s , l e ve ls F 2 a nd E 3 , a t te s t t os even

s pec imens w i l l b e d i scu ssed i nt h e f o l low ing s e c t ions .

w a rm s ea son ( May -Oc t )d ea th s , o ne f a l lk i l l( Nov ) , a nd f i ve c o ld s ea son d ea ths ( one F eb -Ap r . a nd f ou r De c Jan ) .

5 .6 .0 A rchaeo log ica l Imp l ica t ion s o ft he S tudy .

E nva l :

5 .6 .1 I n troduc t ion . B ou rde l le

B o th r e inde e r a nd h or se r ema in s a r e

e xcava t ion s ( b ack o ft h er ock she l ter ) , ac o l lec t ion h oused a t

Ho r se

t e e th

f r om

t he

t h e I n s t i tu t d u Qua te rna ire , B ordeaux , i nd ica te o ccupa t ion

e x tens ive ly u sed i nt h is s t udy a s i nd ica to r so fs ea sona l i ty i n S ou thwes te rn F rance . B efore p roceed ing a ny f u r the r w i th

d ur ing t h e p e r iod f r om May t o O c t a nd d ur ing t h e c o ld

a n a na lys is o ft h e s ea sona l i n d ica tor s f o r t h es amp led s i te s ,

s e a son , p robab ly De c Jan .

t h e refore , t h ee t ho logy o ft h er e inde e r a nd h o r se , a sw e l la s

E sc lauzur :

p rey s pec ie s i n t h e S ou thwe s t , w i l l b e d i scussed . T he

t h e k nown b ehav iou r o f S a iga a n te lope , a no the r i mpor tan t empha s is i nt h ef o l low ing d i scu s s ion i so ns oc ia l g roups a nd S ea sona l i n forma t ion f o rh or se ( c o l le c t ion

s e a sona l p a t te rns o f movemen t .

T aur isson ) s how s ap a t tern o f May -Oc t . h o r se p reda t ion o n ly ( 3 /3 l e g ib le s pec imen s ) .

D iscu ss ion o ft h e e t ho logy o fm a jo r p rey s pec ie s i sf o l lowed b y d i scu s s ion o ft h ed i f f e ren tt e chn ique s u sed i n t h e p a s t t o o b ta in s e a sona l d e term ina t ions f o r t h e

5 .4 D iscus s ion : App l ica t ion o f S ke le tochrono logy t o

s amp led s i te s . Ac ompar ison b e tween r e su l ts o b ta ined u s ing

F o s s i l Hor se T ee th .

t h e se t e chn ique s a nd s ke le toch rono log ica l d e term ina t ions i s o f f ered .

T h in s e c t ion s w e re n o tc ons ide red l e g ib le

T he

w hen o bscu red b y w ha t a ppea red t o b e m ine ra l s t a in ing

f o rego ing

d i scuss ions

p rov ide

e s sen t ia l i n forma t ion f o r i n te rp re t ing t h e o bserved p a t ter n s

a nd /or when t h ey f a i led t o d emon s t ra te s uff ic ien t o p t ica l

i n s ea sona l d e term ina t ions d e scr ibed b e low ( s ec t ion s 5 .4

c on t ras t ( s ee P la te 5 .3 ) . T he se a l tera t ion s i nt h e f o ss i l t e e th

a nd f f .) .

a r e p robab ly d ue t o d i agene t ic c hange s i n t h e c emen tum s t ruc ture . T he e xac t c ause s o f p os t -depos i t iona l s t ruc tu ra l

5 .6 .2 E tho log ica l c ons idera t ion s f or s ea sona l ity s tud ie s . Rang ifer t arandus :

c hange s i n c emen tum a nd b one a r e a s y e t u nknown . M inera l s t a in ing m ay b e t h e r e su l t o f m e ta l o x ides a nd s e d imen t i nf i l t ra t ing t h e c emen tum , o bscur ing t h e g row th

T he e t ho logy o f t h e r e indee r , R ang if e r

l a ye r s . I n e x treme c ase s t h e c e l lu lar s t ruc ture o ft h ec emen t

t a randus , h a s b een a bundan t ly a nd t ho rough ly r e sea rched

i sd e s troyed . I no t her c ase s , f ung i a t tack t h ec emen t ( wh ich

( e .g . , B anf ie ld 1 954 ,

i s 2 4 -26

% o rgan ic ) mak ing o bserva t ions d i ff icu l t o r

S turdy 1 972 ; M i l le r 1 974 ; i n te r a l i i ) . A cco rd ing t o t h e se

i mpos s ib le . R emy ( 1974 ) h a s s ugges ted , a f te r m icroscop ic e xam ina t ion o f f o s s i l t e e th f r om t he Que rcy , F rance , t h a t

a n ima ls a r e c ha rac te r is t ic o f t h is c o ld -adap ted s pec ie s

s peed o f d epos i t ion i s af a c to r i np re serva t ion , s i nce r ap id

( I b id .) . T he se movemen t s a r e d i c ta ted b y d i f f e ren t f e ed ing

b ur ia l wou ld c r ea te a dve r se c ond i t ions f o rb ac ter ia l g row th .

s t ra teg ie s d ur ing t h e c o ld a nd w arm p er iod s o f t h e y ea r .

1 961 ; K e lsa l l 1 957 , 1 960 , 1 968 ;

s t ud ie s , b iannua l m igra t ion s i n vo lv ing l a rge n umber s o f

Movemen t s a r e v ar ious ly r epor ted a s h i gh ly p red ic tab le P oo r ly p re served t e e th w ere n o t a lways

( B inford 1 978 ; Go rdon 1 988 :11) o r e x treme ly v ar iab le

i d en t if iab le m ac roscop ica l ly . C lu s ter s o f p oor ly p re served

( Ke lsa l l

s amp le s

g iven

r e qu iremen ts a r e l i gh t , s of t s now c ove r a nd a bundan t ,

e xcava t ion u n i t s . F o r e xamp le , e i gh t s amp le s f r om a t o ta l

c on t inuous l i chen c over ( Ke lsa l l 1 960 :13 ; S t urdy 1 975 :68 ;

w e re

i d en t if ied

wh ich

c o inc ided

w i th

1 957 :22 ) .

I n

w in te r ,

t h e

a n ima l 's

p r inc ipa l

o ft h ir teen , t a ken f r om P eyrony 's c o l le c t ion f r om L auge r ie

C hernov 1 985 :123 ,125 ) . T he s umme r d i e t c ons is t s ma in ly

H au te E s t , w e re i leg ib le . S im i la r i ly , f our o u t o f f i ve s amp les f r om R e ignac w e re i leg ib le . T e s t t r ia ls s hou ld b e

o fs edge s a nd g ra s se s ( Sp ie s s 1 979 :31) a nd s umme r r ange s a r e f r equen t ly f ound o n f l oodp la in t e r race s ( Go rdon 1 988 :5 ) . Due t ot h e l ow p roduc t iv i ty o f l i chen s ( Chernov

c onduc ted b efore e x tens ive s amp l ing , i no rde r t od e term ine t h e s u i tab i l i ty o ft h e p roposed a s semb lage s .

1 985 :124 )

m igra t ion

i s d e s ireab le

i n o rde r

t o a vo id

o ve rgraz ing ( S turdy 1 975 :68 ; S p ie s s 1 979 :64 ) . S umme r f o rage a rea s , t h e refore , r a re ly c o inc ide w i th w in te r r a nge s

5 6

Figure

5 .2

S ea sona l i ty o f Mand ibu la r T oo th E rup t ion E ven t s E ven t

P o s s ib le A gea

M , e rup t ing DM , h i gh r e l ie f w ea r M , i no cc lu s ion , M2a b sen t

M 2

3 -5 1 -7 3 13 1 015 1 015 1 2-22

P r emo la r s e rup t ing

1 5-29 2 2-29

M , e rup t ing DM , mode ra te r e l ie f i n , M3 a b sen t M , e rup t ing

P o s s ib le S easona l i ty° m id -Augu s t t om id -Oc tobe r m id June t om id -Decembe r m id -Augu s t t om id June m id -Ma rch t om id -Augus t m id -Ma rch t om id -Augus t m id June t om id -Ma rch a ny t i me o fy ea r m id -Ma rch t om id -Sep tembe r

a Age i sg i ven i n mon th s a f te r c a lv ing .

° B a sed o n am id -May c a lv ing p e r iod .

V ar ia t ion i na n t le r g row th a nd s hedd ing c yc le i nc ar ibou o fv a r iou s a ge sa nd s exe s . T he v er t ica l a x is r ep re sen t s r e la t ive s i ze o fa n t le r ; t he h or izon ta l a x i s i s ac a lendr ic s ca le . A : 6 -yea r -o ld a nd o lder ma le s . B :4 - a nd 5 yea r -o ld ma le s . C : N onp regnan t 4 -yea r -o ld a nd o lde r f ema les . D : A l l p regnan t f ema le s . E l : 3 yea r -o ld m a le s . E 2 : 2 -yea r -o ld ma le s . F : 2 - a nd 3 yea r -o ld n onp regnan t f ema le s . G : F awns o f the -yea r . top:

Dental Eruption Patterns ( Rangifer tarandus). bottom: Antler Growth ( R. tarandus). From

Spiess

@ Academic

( 1979: Press,

table

3 .7;

Figure

with permission.

5 7

3 .7).

( Banf ie ld 1 954 ;K e lsa l l 1 960 , 1 968 ; C hernov 1 985 ) . W in te r r a nge s a r e u sua l ly v ery l a rge ; K e lsa l l ( 1957 :61) s t a te s t h a t

We lsh 1 975 :181 ; L ev ine 1 979 :86 ; A n thony 1 991 :252 ) .

w in te rr anges i nt h e C anad ian B ar ren land s o n ly s uppor t 1 5

r a nge s wh ich m ay o ve r lap (We lsh 1 975 :187 ; L ev ine 1 979 :86 ; C lu t ton -Brock 1 987 :85 ) . T he re i s ar epor ted ly h igh d eg ree o f r ange f i de l i ty (We lsh 1 975 :193 ; B e rge r

H e rds a r e g enera l ly n on te rr i tor ia l ' ' ,

a n ima ls p e r s qua re m i le . C a lv ing g round s a re t yp ica l ly e l eva ted , r ugged a nd p oor ly v ege ta ted ( Ke lsa l l 1 968 :177 ) . T he d i s tance s c ove red d ur ing m igra t ion r e la te t o t h e

1 986 :53 ;

d i s tance s

( Sp ie s s

u npred ic tab i l i ty o f movemen t s ) . H arem , o r f am i ly , b and s

1 979 :64 ) . S ea sona l , a l t i tud ina l s h if t s m ay f u lf i l l m igra t ion

a r e more p red ic tab le i n t h e ir u se o f t h e h ome r a nge s

r equ iremen t s , a nd i n mosa ic e cozone s l ong movemen ts m ay

( An thony 1 991 :253 ) .

b e tween

a d jacen t

e co log ica l

z one s

b u t

s e e

War ing

p o sses s ing h ome

1 983 :159 ,

o n

o cca s iona l

n o t b e n ece ssa ry ( Banf ie ld 1 954 ; S turdy 1 975 :69 ) . T he h ome r ange s a re s ea sona l ly a d jus ted P regnan t

f ema le s

i n i t ia te

s u mme r

t h rough movemen t s o f v ar iab le d egree (We lsh 1 975 :144 ;

movemen t s ( Ke lsa l l 1 957 , 1 960 , 1 968 ; M i l le r 1 974 :63 ) . I n

War ing 1 983 :159 ;B e rger 1 986 :60 ) . S uch m igra t ions e nsu re

c on tras tt op urposefu l s pr ing movemen t s , m ovemen ts i nt h e

a cce ss

f a l l a r e d e scr ibed a s " confu sed " , e spe c ia l ly a round t h e

t o s e a sona l ly

a va i lab le

r e sou rce s

a nd

p reven t

l a te s u mme r ( po s t

o ve rpa s tur ing ( Be rge r 1 986 :57 ) .T he h o r se 's movemen t sa r e b locked b y s t eep t e r ra in ( Be rger 1 986 :33 ) . I nw in te r , h e rd s

c a lv ing ) a ggrega t ions a r e r epor ted , h ypo the t ica l ly d i c ta ted

s e ek c ove r i n a r ea s o f h igh r e l ief where t h ey s ome t ime s

b y t h e s oc ia l n eed s o f t h e a n ima ls , r e g roup ing f o r f a l l

c oncen t ra te (We lsh 1 975 :144 ) . A l terna t ive ly f o re s ted a r ea s

m igra t ion ( M i l le r 1 974 :62 ) .

a r e s ough to u t ( Be rge r 1 986 :60 ) . A l t i tud ina lm igra t ions a r e

t r ee l ine ( Ke lsa l l

1 968 :174 ) . L a rge ,

a l so r epor ted ( Berge r 1 986 ) . F a l l movemen t s a r e t r igge red b y s nowf a l l . B ir ths t a ke p lace i n May / June ( Grove s

B ouchud 's ( 1966 ) mode l o f y ea r r ound

1 974 :63 ; We lsh 1 975 :27 ) .

r e s idence o f r e indee r h e rd s i n t h e Do rdogne i mp l ie s a c hange i nt h ee t ho logy o fR ang if e rt a randu s . A n a l terna t ive s ugge s t ion i n vo lve s t h e p resence o f t wo s ub spec ie s o f r e indee r i n t h e Do rdogne : R ang if e r t a randus a r c t icus , t h e

Dur ing t h e l a te P a laeo l i th ic i n t h e A qu i ta ine B a s in , h o r se p robab ly f o l lowed t h es hor t -d is tance

t undra v ar ie ty , a nd R ang if e r t a randu s s y lva t icu s , af o re s t

m igra t ion p a t ter n o bserved

i n f e ra l

dwe l l ing s ub spec ie so fr e indee r whose s e a sona l movemen t s

(War ing

1 986 :50 ) . N ea r t h e Ma s s if

1 983 :159 ; B e rge r

p opu la t ion s t oday

a r e g rea t ly r e duced ( Cheyn ie re ta l1 952 ) . T he e x is tence o f

C en t ra l a nd P yrenean f oo th i l ls s e a sona l movemen t s wou ld

t wo s ympa tr ic s ub -spec ie s o fr e indee r i nt h e Do rdogne , h a s

p robab ly h ave c ompr ised s hor t a l t i tud ina l s h if t s i n r ange

b e en d ef in i t ive ly r e jec ted b y D e lpech ( 1 986 :68 ; s e e a l so

( t ranshumance ) . I na l ll i ke l ihood ,l a te P a leo l i th ic m igra t ion s

1 983 :154 ) .

w ou ld n o tg ene ra l ly h ave i nvo lved l a rge n umber so fa n ima ls

L aco r re

( 1953 ,

1 956 )

h a s

c r i t ic ised

t h e

( I b id .) 1 9.

s ugge s t ion o f a n e t ho log ica l e vo lu t ion f o r r e indee r o n t h e g round s o f g ene t ic i mprobab i l i ty o ve r s u ch a s hor t t ime s pan , g iven e nv ironmen ta l c ond i t ions b e l ieved t o e x is t d ur ing t h e P len ig lac ia l ( a lso , s e e S turdy 1 975 :68 ) . I ts e em s

S a iga t a tar ica :

l i ke ly , t h e refo re , b a sed o n modern e t ho log ica l d a ta , t h a t r e indee r w e re m igra tory d ur ing t h e P len ig lac ia l . D e lpe ch

T he s a iga a n te lope ( Sa iga t a tar ica ) i sa n

( 1987 :13 )s ugge s t ss hor t , e a s t -we s t movemen t sb e tween t h e

i mpor tan ts pec ie s i ns ome h uman e conom ie s i nt h e G ironde

Ma ss if C en tra l a nd t h e G ironde , w i th h e rd s s pend ing mo s t

d ur ing D rya s I( s ee D e lpech 1 983 , 1 989 ) . T he s a iga i sa

o f t h e t ime i n t h e m idd le a l t i tude " co re " a r ea o f t h e

c o ld -adap ted u ngu la te n ow f ound i na r id , s t epp ic o r s em i -

Kr igo rd , w i th h ypo the t ica ln or th -sou th m igra t ion a l ong t h e

d e se r t h ab i ta ts ( Ha r r ing ton 1 981 :196 ) . I ti sa n e x treme ly

n or ther n f l ank s o ft h e P yr6n6e s .

mob i le , m igra tory a n ima l ( Har r ing ton 1 981 :193 ) . L a rge h e rd s o f s a iga h ave b een r epor ted d ur ing s pr ing a nd f a l l

E quus f er n s :

m igra t ions i n t h e U .S .S .R . ( Bobr insk i e t a l 1 944 :247 ; q uo ted

i n Gordon

1 988 :42 ) .

Movemen t s a r e p os s ib ly

O bserva t ion s o f moder n f e ra l g ra s s land

d i c ta ted b y s now c ove r a nd o ther f a c tor s s uch a s w a te r

a nd /or d ese r t h o r se s , s upp lemen ted w i th h i s tor ic a ccoun ts

( Har r ing ton 1 981 :196 ) . B ir ths o ccur i n May / June a nd t h e

o f t r ue w i ld h orse s ,

r u t t a ke s p l ace i nl a te Novembe r ( Ha rr ing ton 1 981 :194 ) .

b ehav iour

o f

c an

P le is tocene

b e u sed t o r e cons truc t h or se

p opu la t ions

t h e

( Be rge r

1 986 :228 ; A n thony 1 991 :252 ) . T he h o r se i s an on se le c t ive

T he s a iga i s n o t n orma l ly t o leran t o f

f e ede r a nd c an b e f ound i na r ea s o fl ow q ua l i ty v ege ta t ion ,

b roken

e . g . , s andy , s a l ine s t eppe s a nd " ha rd t e r ra in" ( Grove s 1 974 :58 ; We lsh 1 975 ; War ing 1 983 :9 ; B e rge r 1 986 :16 ; b u t

1 981 :196 ; D e lpech 1 983 :206 ) , w h ich e xp la ins i t s d i s tr ibu t ion i n t h e G ironde a nd r e la t ive a bsence i n t h e

s e e F reeman 1 973 :9 ) . T he b a s ic s oc ia l u n i t o ft h e h or se i s

Dordogne , w he re i ti so n ly f ound i nr i ve r v a l leys ( De lpech

t h e " f am i ly" , i . e . , f ema le

+ o f f spr ing (We l ls & v on

t e r ra in

a nd

a r ea s

o f h igh

r e l ief ( Har r ing ton

1 983 :F igu re 4 0 ) . T he L ande s r eg ion d ur ing t h e f u l lg lac ia l -

G o ld schm id t -Ro thsch i ld 1 979 ) .

w i th i t s c on t inen ta l d une s - w i l l h ave b een i nhosp i tab le t o t h is a nd most o ther s pec ies ( Ib id .:210) . S ke le tochrono log ica l a na lyse s o f s a iga i n S ou thwe s ter n

T he h o r se l i ve s i ns ma l l s o c ia l g roups o r h e rd s , c ons is t ing o fe i the ra )o ne s t a l l ion , a bou t 1 6 m a re s ,

F rance h ave n o t y e t b een u nder taken .

t h e ir f o a ls , y ea r l ings a nd t wo -yea ro ld o f f spr ing o rb ) l o ose g roups o fj uven i le a nd b ache lo r m a le s ( Grove s 1 974 :58 -59 ; 5 8

1 987 :74 ) .

5 .6 .3 S ea sona l i ty s t ud ie s : o ther s ea sona l i nd ice s . Much c onfu s ion a r ise s d ue t ot h ed i f f e ren t

E rup t ion t a b le s

c an

b e

u sed

t o a ge

m e thodo log ie su sed t oo b ta in s e a sona l d a ta i nS ou thwe s tern F rance . Two m a in c l a sse s o f s e a sona l d a ta c an b e

r e indee r u p t o 1 7 mon ths o f a ge , a ccord ing t o Gu i l l ien

d i s t ingu ished o n t h e b a s is o f w he the r o r n o t p opu la t iona l

( 1974 :77 ) a nd K e lsa l l ( 1968 :28 ) . B e cause o f t h e w ide c onf idence i n terva ls i nvo lved t h is me thod i s l e ss p rec ise

( 1977 :137 ) o r u p

a na lyse s ( t he r e cons t ruc t ion o f p opu la t ion p a rame te r s ) a r e u sed . S ea sona l d e term ina t ion s a r e o f ten b a sed o n t h e

t o 2 1

mon th s ,

t h an o nce b e l ieved ( B info rd D e lpech

i d en t if ica t ion o f a ge c ohor t s w i th in a r e cons truc ted p opu la t ion . G iven a k nown d a te o f b i r th f o r at a rge t s pec ie s, t h e a ge o ft h e y ounge s t c ohor t c an b e i nd ica t ive

a ccord ing t o M i l le r

1 973 :23 ; S p ie s s

1 983 :166 )2 1. F ur thermo re ,

v ar ia t ion

1 979 :71 ; b e tween

i nd iv idua ls o ft h e s ame p opu la t ion ( Ke lsa l l 1 968 :25 ; S p ie s s 1 979 :71 ,77 ) a nd b e twe en d i f f e ren t p opu la t ions ( M i l le r

o ft h es ea son . D en ta le rup t ion /wea rs e r ie s ( f o rr e indee r , s e e

1 974 :15 ) c an b e c ons ide rab le ,

B ouchud 1 966 ; M i l le r 1 974 ; G u i l l ien 1 977 ; f o r h or se , s e e

v ar ia t ion i sn o ta ppa ren ti nB ouchud 's ( 1966 )a ge ing c r i ter ia

s e e

F igure 5 .2a .

T h is

We lsh 1 975 ; L ev ine 1 979 ) a nd m e tr ica l i n forma t ion , a r e

( Sp ie ss 1 979 :71) . T he g ene ra l u t i l i ty o f d en ta l e rup t ion

o f t en u sed t os e g rega te a n ima ls a ccord ing t oa ge .

t a b le s f o r a r chaeo log is t s i s t h e refore q ue s t ionab le o n t h e b a s is o fb o th t h e r a r i ty o f c omp le te d en ta l s e r ie s ( Lacor re

Af ew b a s ic a s sump t ion s a r e m ade w hen

1 953 :551 ; B info rd 1 973 :239 )a nd i n suff ic ien tk now ledge o f

u s ing p opu la t iona l a na lys is o n a r chaeo log ica l f a una . T he m a in

a s sump t ion

i s

t h a t

a k i l l -popu la t ion

c an

t h e d ynam ics o fd eve lopmen t i nt h e s amp led p opu la t ion .

b e

r e con s t ruc ted f r om a r chaeo log ica la s semb lage s , a nd t h a tt h is p opu la t ion r ef le c t s 1 956 :306 ) .

a n a tura l

T aphonom ic

o ne

( L acor re

p roce s se s

c an

S ea sona l d a ta wh ich d o n o t r e ly d i rec t ly

1 953 :551 ; a f fe c t

o n p opu la t ion a na lys is i n c lude : morpho log ica l a na lys is , e . g . , a n t le r g row th s t age s ( Lar te t &C hr is ty 1 873 ; B anf ie ld

t h e

r e pre sen ta t ion o f c l a sse s o f d a ta , h oweve r , e . g . , s o f t ,

1 954 ; B ouchud 1 966 ; S tu rdy 1 975 ) a nd c emen tum a na lys is ,

s ummer a n t le r o f r e indee r m ay n o t s urv ive ( Lacor re

i . e . , t h e a na lys is o f g row th s t ruc ture s i n c emen t ( f o r

1 965 :303 ; S tu rdy 1 975 :64 ) . H uman p roce s s ing a c t iv i t ie s

r e inde e r , s e e McEwan 1 963 ; M i l le r 1 974 ; S p ie ss 1 979 ;

m ay

G o rdon 1 988 ; f o r h or se : We lsh 1 975 ; O lsen 1 988 ; t h is s t udy ) .

a l so

a f f ec t

c l a sse s

o f b one .

F o r

e xamp le ,

t h e

a s t raga lu s , u sed b y B ouchud ( 1966 )t os e x a nd a ge r e inde e r , i sa n e l emen t w i th o n ly l ow t o mode ra te u t i l i ty ( B info rd 1 978 :74 )a nd m ay b ed i scr im ina ted a ga in s ti na r chaeo log ica l a s semb lage s ( Spe th 1 983 :101-102 , 10 , s t udy ing B i son ) . F ina l ly , v ar iou s d eg ree s o f s e x a nd a ge s eg rega t ion e x is t

R e indee rs hed t h e ir a n t ler sa tk nown t ime s o f t h e y ea r a nd a n t le r morpho logy i sa f f ec ted b y s ea sona l

s e a sona l ly i nn a tu ra l p opu la t ions o f g iven s pe c ie s a nd w i l l

c hange s ( Banf ie ld 1 954 , 1 961 ; K e lsa l l 1 968 ) . B a sed o n t h is

b e r ef le c ted i nt h e k i l l -popu la t ion .

i n forma t ion , an umbe r o f a r chaeo log is ts h ave a t temp ted t o r e con s truc t s e asona l mor ta l i ty p a t te r n s i n f a una l a s semb lage s ( e .g . , L ar te t & C hr is ty 1 873 ; B ouchud 1 966 ;

Ap rob lem w i th p opu la t ion r e cons truc t ion s f o r r e inde e r b a sed o n a r chaeo log ica l a s semb lage s , i st h e

S turdy 1 975 ) . T he d eve lopmen t a nd s hedd ing o f a n t le r s

e x is tence o f s e a sona l ly v a r iab le a ge a nd s e x s eg rega t ion

v ar ie sa ccord ing t oa ge a nd s ex , h oweve r , ( s ee F igu re 5 .2b )

w i th in

m ak ing t h e s egrega t ion o f s ex /age c l a sse s o f a n t le r n e ce s sa ry ( Ke lsa l l 1 968 :151 ; S turdy 1 975 :57 ; S p ie ss

t h e h e rd

( Ke lsa l l

1 960 :22 ,

1 968 :156f f ;

M i l le r

1 974 :77 , ) .

1 979 :99 ; D e lpe ch 1 983 :152 ) . T he u se o f c a s t a n t le r s f r om " Sma l l h e rds i n s ma l l a r ea s u sua l ly s howed a

a r chaeo log ica l a s semb lage s i s d oub ly p rob lema t ic , f o r r e a son s s t a ted a bove a nd b e cause t h e a s sump t ion mu s t b e

s t rong b i a s i n f a vour o f o ne o r more a ge o r s e x

m ade t h a t t h e a n t le r w a s n o t b e ing c o l le c ted s epa ra te ly ( Sp ie s s 1 979 :97 ; D e lpech 1 983 :166 ) .

g roup s " ( Ke lsa l l 1 968 :150 ) . A s ar e su l t , i ti se ven d i ff icu l t t o s t udy t h e p opu la t iona l

B ouchud ( 1966 ) a t temp ted t o r ef ine t h e

d ynam ic s o fl i ve h e rds , e xcep t d ur ing t h e ir d i spe r sed p hase

morpho log ica l a na lys is o fa n t le r b y d e term in ing t h e d eg ree

i n A ugu s t a nd

( Ke lsa l l

o f c ompac tne s s o f t h e p ed ic le u s ing X -rays . B ouchud 's

1 978 :155 ) . A ge /sex s egrega t ion a l so e x is ts i nh o r se ( f am i ly

p r ior

t o a nd

d ur ing

t h e

r u t

( 1966 ) f i nd ings h ave b een h eav i ly c r i t ic ised , h oweve r( e .g . ,

v s . b ache lo r h e rd s ) . T h is f a c tn eed s t ob ec ons ide red b efo re

L aco r re 1 953 ; 1 956 ; B info rd 1 973 ; S turdy 1 975 ; Gu i l l ien

t h e a s sump t ion t h a t a n a r chaeo log ica l a s semb lage c an b e

1 977 ; S p ie s s

u sed t o r e cons truc t p opu la t ion p arame te r s f o r h or se ( e .g . ,

s hedd ing v a ry a ccord ing t o i n d iv idua l a ge a nd s ex , i . e . ,

1 979 ) . R a te s o f c ompac t ion a nd t ime o f

L ev ine 1 979 ; O lsen 1 988 ; A n thony 1 991) c an b e m ade ,

c onf idence i n terva ls f o r s e a sona l d e term ina t ions b a sed o n

e ven a s sum ing ac a ta s troph ic p rof i le , .

a n t le r morpho logy a r e w ide r t h an B ouchud 's ( 1966 ) s t udy s how s ( S turdy 1 975 :61 ; S p ie ss 1 979 :99 ) . B ouchud w a s u nab le t os eg rega te h i sd a ta a l ong t h ese l i ne s , h oweve r , t hus

T he u se o f t o o th w ea r t a b le s r e qu ire s a n

i n va l ida t ing h i ss t udy ( Laco r re 1 953 :548 ; S turdy 1 975 :57 ) .

a s sump t ion o f c ons tan t w ea r t h roughou t t h e y ea r a nd t h roughou t t h e l i f e c yc le . T h is a s sump t ion d oe s n o t h o ld i n m odern p opu la t ions o f e i the r r e inde e r o r h or se ( Ke lsa l l

s ke le ta l

1 968 :28 ; M i l le r 1 974 :76 ; S t u rdy 1 975 :55 ; We lsh 1 975 ; G u i l l ien 1 977 :138 ; o r r e d d e e r , R ow ley -Conwy e t a l .

p ar t s

V ar ia t ion i nt h er e pre sen ta t ion o fd i f f e ren t i n a n a r chaeo log ica l a s semb lage i s a n

i n d ica tor o f h uman b ody p ar t p ref e rence s , t h e t e chno logy a va i lab le f o rp roce ss ing a nd s t or ing , a nd l o g is t ica ld ec is ion s 5 9

r e la ted t od i s tance f r om c amp a nd h un t ing s t r a tegy ( B info rd 1 978 ; S p ie s s 1 979 :24 ) . G iven t h a t b ody p ar t p ref e rence i s b a sed o n k nown n u tr i t iona l v a lue s ( B inf o rd 1 978 )2 a nd i s

5 .7 .1 S easona l i nd ica tor s i n t he D ordogne .

f a ir ly c on s tan t a mong c a r ibou d ependen t p eop le s ( Sp ie s s 1 979 ) , a nd i f Magda len ian t e chno logy i st r ea ted a s ag iven , e l emen t p a r t r e p re sen ta t ion c an t h eo re t ica l ly b e u sed t o r e con s truc t t h e se l o g is t ica l d e c i s ion s ( e .g . , B oy le 1 990 ) .

Wh i te ( 1985 )n o te st h a tl o ca t iona ld a ta f o r L a Made le ine , R e ignac a nd L auger ie H au te E s t , a l l o f wh ich a r e s i tua ted o n f o rd s o r c onf luence s , a r e s ugge s t ive o f i n te rcep t ion , o r m ig ra t ion h un t ing . T h is i mp l ie s t h e u se

A s sum ing s e a sona l c hange s i n h uman mob i l i ty p a t te rn s , r e la ted t oc hange s i n r e sou r ce a va i lab i l i ty , t h e se l og is t ica l

o f r i ve r v a l leys , e i the r a s s e t t ing s f o r n a tu r a l t r ap s , o r f o r t h e ir c hanne l ing e f f e c t , w h ich w i l l h ave a f f e c ted l a rge , s ea sona l a gg rega t ion s ( Bu r ch 1 972 :347 ) . A cco rd ing t o B info rd ( 1978 :392 ) , a mong t h e Nunam iu t w a te r t r ap s a r e c ommon f a l l i n te rcep t s t ra teg ie s , b u t a r e n o t u sed i n t h e s pr ing w hen p ound sa r ep ref e r red . T he s ke le toch rono log ica l d a ta d o n o t s uppo r t t h e e x ten s ion o fB info rd 's o b ser va t ion t oo u r s i te s , h oweve r ( s ee d i scus s ion , b e low ) .

d e c is ion s c an b e c on s ide red r e la ted t o s e a sona l i ty ( e .g . , B oy le 1 990 ) . U se o fs ke le ta lp a r tr e p re sen ta t ion i nt h is w ay , h oweve r , r equ i re s s eve ra l a s sump t ion s . " As sum ing t h a te conom ic c on s ide ra t ion o ft h e s or t c on s ide red a nd q uan t if ied b y B info rd ( 1978 ) w e re k ey t ot h e d e c is ion m ak ing p roce s s a nd a s sum ing

L oca t iona l d a ta , i . e . , h uman u se o fr i ve r v a l ley l o ca t ion s , e spe c ia l ly a tf o rd s o r c onf luence s (Wh i te 1 985 ) , i s al i ke ly i n d ica to r o f h un t ing s t ra tegy b u t i s s ea sona l ly a mb iguou s . S pr ing a nd f a l l m ig ra t ion s a r e

t h a t t h e b one p re serva t ion i n ag i ven a s semb lage r ef lec t s d ec is ion b a sed o n t h e se c on s ide ra t ion s " ( Gray son 1 988 :65 ) .

c apab le o f b e ing c hane l led b y r i ve r v a l ley s ( Bu r ch 1 972 :347 ) a l though K e lsa l l ( 1957 :59 ) n o te s t h a t t h i s i s e spec ia l ly t h e c a se i nt h e s pr ing . F u r the rmo re , i ns umme r , r e indee r t e nd t o " bunch " n ea r w a te r cou r se s ( Ke lsa l l 1 969 :16 ) m ak ing t h e se t h e o n ly p red ic tab le l o ca t ion s f o r e ncoun ter ing t h e s e a sona l ly d i spe r sed h e rd s .

C au t ion s hou ld t h e ref o re b e e xe rc ised w hen i n te rp re t ing e conom ic a c t iv i t ie s a nd s e a sona l i ty f r om p a t terns o f a na tom ica l e l emen t r e p re sen ta t ion . C emen tum a na lys is o f f e r s t h e h ope o f o b ta in ing a b so lu te s e a sona l d e te rm ina t ion s , u s ing s i ng le t e e th a nd i g no r ing s e x /age s egrega t ion i n t h e t a rge t p opu la t ion . A l though t h e a s sump t ion o f ac ons tan t r a te o f a ppo s i t ion o fc emen td ur ing t h ey ea r ly c yc le i sp robab ly n o t

A t l a Made le ine , t h e d e ep s t ra t igr aph ic s equence ( e v idence f o r f r equen t s i te r e -u se ) c oup led w i th r e la t ive ly s t ab le N ISP v a lue s ( Boy le 1 990 :220 )i sc on s is ten t w i th B inf o rd 's ( 1978 ) d a ta d ocumen t ing t h e r e -occupa t ion o fs t ra teg ica l ly p l a ced r e s iden t ia l l o ca t ion s i nt h ec on tex to f i n te r cep t /m ig ra t ion h un t ing . I nd ica to r s o f f a l l /w in te r o ccupa t ion , s uch a s l o ca t iona l d a ta ( s e e B o rde s 1 972 :442 ; Wh i te 1 985 :131 ) a nd r e inde e r d en t i t ion ( De lpe ch p e r s .comm . t o Wh i te 1 985 :156 ) s e em ing ly c on t rad ic t s e a sona l d e term ina t ion s o b ta ined b y s ke le toch rono logy . Gordon 's ( 1988 :198 ; p er s .comm .1991) d a ta f o r Magda len ian I V o ccupa t ion s s how l i t t le o r n o e v idence o f f a l l r e indee rk i l ls , b u tr a the r , e v idence o fw in te r( Dec -Ma r ) a nd p redom inan t ly s pr ing ( Ap r -May ) k i l ls . D a ta f o r h o r se mor ta l i ty ( t h is s t udy ) d ur ing t h e L owe r Magda len ian s equence s how s p redom inan t ly w a rm s e a son , ( May -Oc t ) , d ea th s . 23

v a l id , b road s e a sona l a t tr ibu t ion sa r e p o s s ib le u s ing g row th s t ruc tu re s i n c emen t ( f o r h o r se s e e We lsh 1 975 ; O lsen 1 988 ; C hap te r 3 , a bove ; f o r r e inde e r s e e McEwan 1 963 ; M i l le r 1 974 ; S p ie s s 1 979 ; G o rdon 1 988 ) . T h is m e thod i s l e s s r e s tr ic ted i ni t s a pp l ica t ion t h an o the r s , w hen d ea l ing w i th r e la t ive ly s ma l l a r chaeo log ica l p opu la t ion s ( Sp ie s s 1 979 :69 ) . 5 .7 .0 D iscu s s ion : r eg iona l p a t tern ing a nd i nd ica tor s o f s ea sona l i ty i nt he f auna l a s semb lage s . I n t h e f o l low ing d i s cus s ion , p a t tern s o f s e a sona l i ty w i l l b e e xam ined i n f ou r s ub reg ions o f S ou thwe s te rn F rance : t h e Do rdogne ; t h e G ironde ; t h e Ma s s if C en t ra l , i n c lud ing t h e H au te L o ire a nd t h e A l l ie r ; a nd t h e L ande s . T he se d i v is ion sa r en o t " t aken f o rg ran ted " ( Conkey 1 987 :69 ) , b u t r ef le c t t o pog raph ica l a nd b iogeog raph ica l r e a l i t ie s d i scu s sed i n C hap te r 4 , p ar t A

T he l a ck o f c onco rdance b e twe en G o rdon 's r e su l t s a nd o t he r s e a sona l i n d ica to r s m ay b e d ue t o av ar ie ty o fc i r cum s tance s : 1 )f a l lr e inde e rd ea th sm ay b e m is iden t if ied i n G o rdon 's s t udy' o r 2 ) h uman s a t L a Made le ine m ay h ave h un ted r e indee r p r ima r i ly d u r ing t h e s pr ing m ig ra t ion a nd a l so i nt h e ir w in te rr a nge , k i l l ing o n ly af ew n on -m ig ra tory a n ima ls ( e .g . , b u l ls , n on -f e r t i le o r i n jured c ow s' ) i n t h e f a l l a nd h un t ing h o r se a s a n a l terna t ive r e sou rce w hen r e inde e rw e re a b sen to rd i spe r s ed ( e .g . , i nt h e i r s umme r r a nge ) . I n t h e l a t te r c a se , t h e e r ro r l i e s w i th t h e i n terpr e ta t ion o f t h e d en ta l e r up t ion s e r ie s f o r

( s up ra ) . T he d i scu s s ion w i l l s how t h a t s i te s f r om t h eD ordogne ( L a Made le ine , L auger ie H au te E s t , R e ignac , C ap B lanc , B adegou le a nd E sc lauzu r ) s ha re s im i la r i t ie s i n f a una l p a t te rn ing , a s d o s i te s f r om t h e G ironde ( Sa in t G erma in la -r iv ire , R oc d e Ma rcamp s a nd Mou l in N eu f ) . S i te s f r om t h e Ma s s if C en t ra l ( Enva l a nd R ond d u B ar ry ) d o n o t s ha re t h e s ame a ff in i ty . D uru thy , b e ing t h e o n ly r ep re sen ta t ive o f t h e d i scu s sed s epe ra te ly .

L ande s /Py ra i6e s

A t lan t ique s ,

r e indee r ( s e e d i scu s s ion , b e low ) . T he m o r ta l i ty p a t te rn f o r h o r se s ugge s t s ap robab le " e x ten s ion" o fh uman o ccupa t ion b eyond t h e p er iod o fs e a sona l a va i lab i l i ty o fr e inde e r .

i s

A tR e ignac , B oy le ( 1990 :226 )s ugge s t st h e 6 0

p a t tern o f a na tom ica l p ar t r e pre sen ta t ion f o r r e indee r i s

r e indee r d en ta l s e r ie s , B oy le ( 1990 :263 ) i n terp re t s t h e

c ons is ten tw i th B inford 's d a ta f o r f a l l /w in te ra nd l a te s pr ing

p a t te r n o f e l emen t r epre sen ta t ion a s i n d ica t ing f a l l a nd s pr ing o ccupancy a t R oc d e Ma rcamp s : r e su l t s f o r s a iga ,

h un t ing

c amps .

T he

b u lk

c urve

f o r

h or se

s ugge s t s

h or se a nd r e indee r s ugge s t s pr ing h un t s a nd r e su l t s f o r

e ncoun te r -ba sed h un t ing o ccur ing d ur ing t h e f a l l ( I b id .) . S ke le to ch rono log ica l d a ta f o r h or se i s m eag re , b u t t h e s i ng le l e g ib le s pe c imen d ocumen t s aw arm s e a son ( May -

b ov ine s s ugge s t a u tumn k i l ls ( I b id .) . Go rdon 's ( 1 988 :200 ) d a ta s uppo r t s at h e s is o fw in te r /spr ing ( Dec - May ) r e indee r

O c t ) k i l l . Wh i le Wh i te ( 1985 :131) s ugge s t s t h a t r i ve r

e xp lo i ta t ion .S ke le tochrono log ica ld a ta f o rh or se ( t h is s t udy )

c ros s ing l o ca t ions a r ec ons is ten tw i th f a l lm igra t ion h un t ing ,

i n d ica te s b o th w arm a nd c o ld s e a son mor ta l i ty .

G o rdon 's ( 1 988 :212 ) d a ta f o r r e indee r i n t h e Uppe r Magda len ian l e ve ls l a ck e v idence f o r f a l l k i l ls , s how ing

A t Mou l in N euf , w in te r ( De c -Ma r ) k i l ls

p redom inan t ly s pr ing a nd w in te r mor ta l i t ie s .

o f r e indee r ( Gordon 1 988 :205 ) a nd b o th w a rm s e a son a nd c o ld s ea son d ea ths f o r h or se , ( t h is s t udy ) a r e d ocumen ted .

A t

C ap

B lanc ,

G o rdon

( 1988 :159 ) -

d ocumen t s w in te r /spr ing ( De c - May ) mor ta l i ty p a t terns f o r

5 .7 .3 S easona l i nd ica tor s i n t he Ma s s if C en tra l .

r e indee r . H or se mor ta l i ty o ccu r s d ur ing b o th t h e c o ld a nd w arm s e a son s , i . e . , t h e s i te w a s u sed d ur ing p o ten t ia l ly a l l

A t R ond d u B a r ry , s ke le tochrono log ica l

s e a son s , a c co rd ing t ot h e p resen t s t udy .

d a ta f o r h o r se ( t h is s t udy ) , t h e d om inan t p rey s pe c ie s ( Aa jane 1 986 ) , i nd ica te s b o th w arm ( May -Oc t ) a nd c o ld

A t B adegou le , B ouchud a nd c o l league s

( Nov -Apr i l ) s e a son mor ta l i ty . T he e v idence f o r e x tens ive

h ave p roposed y ea r r ound h uman r e s idence d ur ing t h e

p roce ss ing o f h or se r ema ins s ugge s t s t h a t t h e p a t ter n o f

S o lu t rean a nd L owe r Magda len ian ( Cheyn ie r e ta l . 1 952 ;

e xp lo i ta t ion i sd i s s im i la r t o t h a t o f r e inde e r i n r e indee r -

B ouchud

d om ina ted a s semb lage s . I n o the r w o rds , t h e e t ho logy o f

1 966 ) .

T h is p roposa l h a s b een c on te s ted o n i t se lf i s

h or se i s c ond i t ion ing t h e h uman e xp lo i ta t ion p a t ter n s .

e s sen t ia l ly r e ta ined ) b y L acor re ( 1953 ; 1 956 ) . Gordon 's d a ta f o r L owe r Magda len ian ( Mag .0 l e ve ls , p e r s .comm .)

G o rdon 's ( 1988 :219 ) d a ta f o r r e la t ive ly s c a rce r e indee r a t

l e ve ls d o n o t e n t ire ly c on trad ic t t h e h ypo the s is o f y ea r

T h is r a ise s t h e p os s ib i l i ty t h a t t h e a r ea a round R ond d u

r ound s i te -use , e xcep t f o r a n a bsence o f e a r ly /m id -w in te r r e inde e r d ea ths ( Go rdon 1 988 :154 ) . T h is i st h e o n ly s i te i n

B ar ry m a rk s t h e e x treme o f aw in te r r a nge f o r r e indee r . S i nce b u l ls u sua l ly v en tu re much f a r the r t h an c ow s i n to

t h e D o rdogne f o r wh ich Gordon d ocumen ts f a l l ( Sep t -Nov ) k i l ls . H o r se mor ta l i ty a c cord ing t oo ur r e su l t s , o nce a ga in ,

w in te r r ange s ( M i l le r 1 974 :63 ) , i t wou ld b e i n te re s t ing t o s e x t h e r e inde e r r ema in s a tt h is s i te . T he l a ck o f s umme r

i n c lude s b o th s e a son s d ur ing t h e L owe r Magda len ian .

k i l ls a nd t h e o ve ra l l n umer ica l d ef ic i t o f r e indee r a tR ond

m e thodo log ica l

g rounds

( bu t

t h e

h ypo the s is

R ond d u B ar ry i n d ica te s o n ly w in te r ( Dec -Ma r ) m or ta l i ty .

d u B ar ry s ugge s t s t h a t s u mme r r a nge s w e re n o t n ea rby . T h is

A t L auger ie H au te E s t , B ouchud 's ( 1966 )

m ay

e xp la in

t h e a ppa ren t

d i ve r s i ty o f e co tone s

e xp lo i ted a tR ond d u B a r ry d ur ing t h e L owe r Magda len ian .

s ugge s t ion o fp e rmanen t h uman r e s idence d ur ing t h eL owe r Magda len ian ( b a sed o n r e indee r a n t le r a nd t o o th d a ta ) i s c ons is ten tw i th t h e s ke le toch rono log ica l d a ta f o rh or se ( t h is

A t E nva l , w he re r e indee r i st h e d om inan t

s t udy ) . L oca t iona l d a ta p o in t s t om igra t ion h un t ing (Wh i te

p rey s pe c ie s , t h e mor ta l i ty p a t ter n f o r h o r se ( t h is s t udy )

1 985 :131) . C emen tum a na lys is o fr e indee r t e e th d ocumen ts

i n d ica tes b o th w arm a nd c o ld s e a son e xp lo i ta t ion . R e indee r

p redom inan t ly s pr ing a nd s ome w in te r( i .e . , D e c - May ) k i l ls

t e e th , s amp led f r om U ppe r /F ina l Magda len ian l e ve ls s how

( Gordon 1 988 :192 ) .

p redom inan t ly s pr ing mo r ta l i t ie s ( Apr -May ) , i mp ly ing a

5 .7 .2 S easona l i nd ica tor s i n t he G ironde .

d ea ths ( Go rdon 1 988 :176 ) . T he d a ta a va i lab le s ugge s t s t h a t h uman e conom ie s a t E nva l a nd R ond d u B a r ry a r e

m igra t ion / in te rcep th un t ,w i th o cca s iona ls umme r( J un -Aug )

d i ss im i la r . A l though r e inde e r r ema ins a n i mpor tan t 5 .7 .4 S easona l i nd ica tor s i n t he L ande s .

p rey s p e c ie s i n t h e G ironde , o t he r s pe c ie s s uch a s s a iga , h o r se a nd b ov ine s f r equen t ly d om ina te o r c o -dom ina te i n d iv idua l f a una l a s semb lage s ( Boy le 1 990 :136 ; a nd s e e

L owe r Magda len ian a nd Magda len ian I V

D e lpe ch 1 983 ) . S a iga d om ina te st h e f a una a tS a in tG erma in

l e ve ls a tD uru thy w e re s amp led . I nt h e L owe r Magda len ian

a nd a tR oc d e Ma rcamp s , f o r e xamp le , c o -occur ing w i th

l e ve l h or se i st h e p redom inan t p rey s pe c ie s , f o l lowed b y

h or se , r e indee r a nd b ov ine s ( De lpe ch 1 983 ; L eno ir 1 989 ) .

r e indeer ( De lpe ch 1 983 ) . I n t h e Magda len ian I V l e ve l , r e indeer d om ina te s a nd h o r se i so n ly s e condary ( I b id .) .

G ordon 's ( 1988 :223 ) d a ta f o r r e inde er a t S a in t -Germa in la -r iv ire s ugge s t ap a t tern o fp redom inan t ly

A rambou rou a nd D e lpech i n d ica te f a l la nd

f a l l / la te -w in ter ( S ep t - Ma r ) r e inde e r mor ta l i ty a nd s i ng le

w in te r o ccupancy a t D uru thy i n h or izon c . 4 ( Magd .IV ) b a sed o n o b served f a una l a s soc ia t ions ( Arambourou

s umme r

( J un -Aug )

a nd

f a l l

( Sep t -Nov )

k i l ls .

S ke le toch rono log ica ld a ta f o rh o r se ( t h is s t udy )i sc ons is ten t

1 976 :24 ; D e lpe ch 1 983 :113 ) . G o rdon 's ( 1988 :172 )d a ta f o r

w i th G o rdon 's d a ta , s how ing b o th w arm a nd c o ld s e a son

h or izon 4c on t rad ic t s t h e h ypo the s is o f f a l l /w in te rp re sence

m or ta l i ty .

o n ly , w i th e v idence f o r as i ng le s pr ing o r e a r ly s umme r ( Apr -May / Jun -Aug ) k i l l . R e su l t s o f t h e p resen t s t udy i n d ica te h or se w a s h un ted d ur ing b o th c o ld a nd w arm

S uppor ted b y L acor re 's ( 1951) s t udy o f 6 1

p robab ly h un ted o n a n e ncoun te r b a s is i n t h e Do rdogne

s e a sons i n Magd .IV l e ve ls a t Du ru thy' .

( Boy le 1 990 ; a f te r B inf o rd 1 978 :265 ) . T h is i sc on s is ten t w i th t h e k nown b ehav iou r o f t h e s pe c ie s , wh ich e x is t s i n

Dur ing t h eL owe r Magda len ian a tD uru thy ( hor izon c . 5 ) , s e a sona l d a ta f o r h o r se ( t h is s t udy ) i n d ica te s

s ma l l s o c ia l g roup ing s , w e l l -spaced i nt h e l a ndscape ( s ee d i s cu s s ion a bove ) . Ho r se a ta l l s i te s e xcep t R e ignac ( whe re t h e s amp le i s r e duced t o as i ng le l e g ib le s l ide ) s how s e v idence o fh av ing b een e xp lo i ted d ur ing b o th c o ld ( Dec .

w arm s ea son ( May -Oc t ) mor ta l i ty o n ly . T h is i n f orma t ion i nd ica te s t h a t t h e s ugge s ted w in te r e xp lo i ta t ion o fr e indee r ( A rambou rou 1 976 ; D e lpech 1 983 ) d oe s n o t r e p re sen t t h e s ea sona l l im i t o f h uman r e s idence a t t h e s i te . S pe c ie s r epre sen ta t ion f o r b o th Magda len ian I I a nd I V l e ve ls

t oA p r . ) a nd w arm ( May t oO c t .) s e a son s , e xc lud ing " f a l l" ( Nov .) .

i nd ica te s t h a t b ov ine s w e re e conom ica l ly i mpo r tan t ( De lp ech 1 983 :113 ) . I n o rde r f o r t h is d i scu s s ion t o b e c omp le te , t h e ref ore , s e a sona l d e te rm ina t ion s f o r b ov ine s

Two i n te rp re ta t ion s o f t h e c omb ined s e a sona l d a ta a r e p o s s ib le 1 ) t h e s i te s s urveyed w e re o ccup ied y ea r r ound a nd t h e r e sou r ce b a se s h if t ed

mu s t a l so b e m ade .

s e a sona l ly o r2 ) s i te s w e re p r imar i ly o ccup ied i nt h e w in te r a nd s pr ing ( Dec -May ) when r e indee r w e re a va i lab le , a nd h o r se w a s u sed t o s upp lemen t d w ind l ing n umbe r s o f

5 .8 C onc lu s ion s . F auna l a s semb lage s f r om t h e D o rdogne

r e indee r d ur ing t h e l a te s p r ing b efo re a s h if t t o s umme r r e s idence s . I n a ny c a se , h o r se i s a r e sour ce w i th a c omp lemen ta ry s e a sona l i ty , i . e . , e xp lo i ted o u t s ide t h e p er iod o f r e inde e r m igra t ion , p robab ly o n a n e ncoun te r

a r e c ha rac ter ised b y t h e n umer ica l d om inance o f r e indee r a nd t h e s e condary i mpor tance o fh o r se ; o the r t a xa s uch a s r ed d ee r , b ov ine sa nd i b ex f o rm am inu scu le c omponen to f t h e i d en t if ied s pe c ie s ( s ee D e lpe ch 1 983 ) . T he p redom inance o f r e inde e r , wh i le p a r t ly a r ef le c t ion o f

b a s is , p o s s ib ly d ur ing t h e mon i tor ing o fr e indee r m ig ra t ion ( B info rd 1 978 ) .

r eg iona l b iogeography , i sa l so u ndoub ted ly ar ef le c t ion o f h uman h un t ing p a t te rn s a ccomoda t ing t o t h e e t ho logy o f t h e ir p rey .

I nt h eG i ronde , s ea sona l i n forma t ion f r om t h e t h re e s i te s s urveyed c an b e i n te rp re ted s im i lar i ly . I na l l t h re e c a se s e v idence f o r w in te r e xp lo i ta t ion o f r e inde e r s ugge s t s a n a cce s s ib le w in te r r a nge e xp lo i ted o n a n e ncoun te r b a s is . F ur thermo re , a tS a in t -Ge rma in la -r iv ire , R oc d e Ma rcamp s a nd Mou l in N euf , s pr ing ( Ap r -May ) r e inde e rm o r ta l i t ie ss ugge s ti n te rcep t ion o fm ig ra to ry h e rd s . T he i n c idence o f s umme r a nd f a l l r e indee r k i l ls a t S a in t G erma in i sl ow e nough t o s ugge s t i n d iv idua l ( e ncoun te r )

T he p a t tern o f a na tom ica l p ar t r ep re sen ta t ion f o r r e indee r i n t h e D o rdogne i sc on s is ten t w i th B info rd 's ( 1978 ) g ourme t c urve a cco rd ing t o B oy le ( 1990 ) . T h is p a t te rn o fe l emen t r ep re sen ta t ion i si n d ica t ive o f t h e i n te rcep t ion o f m ig ra to ry h e rd s a l ong p red ic ted m igra t ion r ou te s a nd b u lk p roce s s ing o f l a rge n umbe r s o f c a r ca sse s ( f . B info rd 1 978 ) . E ncoun te r -ba sed h un t ing o f

e xp lo i ta t ion o f n on -m ig ra tory a n ima ls . T he s e a sona l i n f orma t ion f o r h o r se i nt h e G ironde i n d ica te s ap a t ter no f w arm a nd c o ld s e a son e xp lo i ta t ion i n c lud ing f a l l ( Nov . ) .

r e inde e r i n t h e Dordogne p robab ly a l so o ccur red d ur ing t ime s o f h e rd d i spe r sa l , i mp ly ing t h a t 1 ) w in te r a nd /o r s umme r r ange s w e re a cce s s ib le t oh un te r s , o r2 )a n ima ls a t t h e e x t reme l im i t so ft h e se r a nge sw e r ea c ce s s ib le . A n im a ls r ema in ing i nw in te rr ange s a f te r s pr ing m ig ra t ion o rs imp ly l a gg ing b eh ind t h e m a in m ig ra t ion m ovemen t c ou ld a l so h ave b een e xp lo i ted o n a n e ncoun te r b a s is . T he n umbe r s i nvo lved w i l ln e ce ssa r i ly h ave b een s ma l l , h oweve r , a nd t h e f r agmen ta t ion p a t ter n s g ene ra ted i n t h e a s semb lage s a r e

E v idence f r om t h e G ironde c an t h e refo re b e i n te rp re ted a si n d ica t ive o ff a l l ( Nov ) , w in te r( De c /Ma r ) a nd s p r ing ( Ap r / May )o ccupancy o ft h es i te ss urveyed , p lu s e i the r 1 )e a r ly w a rm s e a son ( May ) a nd /o rl a te w a rm s ea son ( S ep t -Oc t ) r e s idence o r 2 ) w arm s e a son r e s idence ( May O c t ) . I n o rde r t o g a in a f u l l a pp re c ia t ion o f s ea sona l e conom ic a c t iv i ty i n t h e G ironde , h oweve r , i t wou ld b e n e ce s sa ry t o e x tend t h e s ke le to chrono log ica l s t udy t o i n c lude s a iga a nd p e rhap s b ov ine s , w h ich i sb eyond t h e s cope o ft h e p re sen t r e sea rch .

s ubme rged b y t h e p a t ter n g ene ra ted b y m ig ra t ion / in te r cep t h un t ing . A cco rd ing t o s ke le to ch rono log ica l d a ta , r e indee ri nt h eD o rdogne s i te ss urveyed w e re p redom inan t ly

T he s e a sona l d a ta f o r t h e Ma s s if C en t ra l ,

h un ted i n t h e w in te r a nd s pr ing ( De c -May ) . H uman s , t h e refo re , w e re e xp lo i t ing t h e s pec ie s d ur ing a tl e a s t o ne o f

a tR ond d u B ar ry a nd E nva l , s ugge s t t wo e conom ie s w i th v e ry d i f f e ren t s e a sona l s t ruc tu re s . A t E nva l , s e a sona l d a ta s ugge s t s a s im i la r p a t ter n t o t h a t d e scr ibed f o r t h e D o rdogne ( a bove ) , i . e . , s pr ing m igra t ion / in te r cep t h un t o f r e indee r , w i th o cca s iona l e ncoun te r h un t ing d ur ing t h e c ou r se o f t h e s umme r a nd e ncoun te r h un t ing o f h o r se d ur ing b o th w a rm a nd c o ld s e a son s . N o te t h a t o cca s iona l

t h eb iannua l m ig ra t ion s , i . e . , i ns pr ing , a st h eh e rd sm oved o u t o f w in te r r a nge s . R e indee r a l so a ppea r t o h ave b e en h un ted i nw in te rr a nge s , d u r ing t h e t ime o fm ax imum h e rd d i spe r sa l ( Bu r ch 1 972 :345 ; M i l le r 1 974 :63 ) . T h is i mp l ie s t h e u se o f t wo d i s t inc t a nd s e a sona l ly d i c ta ted h un t ing s t ra teg ie s f o r t h e s ame s pe c ie s , n ame ly i n te r cep t a nd e ncoun te rh un t ing ,a sh a sb een e thnogr aph ica l ly d ocumen ted ( e .g . , B info rd 1 978 )2 7.

e v idence o f r e indee r mor ta l i t ie s d u r ing t h e s umme r d o n o t c on t r ad ic t t h e h ypo the s is o fb iannua l m igra t ion s .

o f

A tR ond d u B ar ry , i nc on t ra s t , e ncoun te r -

a na tom ica l p ar t s f o r h o r se i n d ica te t h a t t h is s pe c ie s w a s

b a sed h un t ing ( on ly ) o f r e indee r a t t h e e x treme s o f i t s

B u lk

c urve s

i n

t h e

d i s tr ibu t ion

6 2

w in te r r ange i ss ugge s ted . T h is m ay e xp la in t h e a ppa ren t d ive r s i ty o f s pec ie s e xp lo i ted h e re ( s e e D e lpech 1 983 ) . Hor se , t h e d om inan t p rey s pe c ie s a t R ond d u B ar ry , i s p o ten t ia l ly e xp lo i ted y ea r r ound . T he h i gh d eg ree o fb one f r agmen ta t ion s ugge s t s h o r se w a s i n ten s ive ly u t i l ised . I n o rde r t ou nde r s tand t h e s e a sona l e conomy a tR ond d u B a r ry mo re f u l ly , i ti s s ugge s ted t h a t t h e s e a sona l p a t tern o f mor ta l i ty o f i b ex ( t he s e cond mo s t i mpor tan t s pec ie s h e re ) s hou ld b e s t ud ied . I n s u mma ry , i na l l t h e s ub reg ion s o ft h e Aqu i ta ine b a s in

e xam ined

e xcep t

t h e Ma s s if C en t ra l ,

s e a sona l d a ta s ugge s t s t h a t h o r se w a s e xp lo i ted a s a c omp lemen tary r e source , h un ted e i the r d ur ing mon i tor ing o fr e indee r m igra t ions ( a nd p e rhaps s a iga m igra t ion s i nt h e G ironde ) , o rd ur ing p er iods w hen r e inde e r w e re n o t l o ca l ly a va i lab le ( o r a tl e a s t n o t e xp lo i tab le o n a n i n te rcep t b a s is , i . e . ,

when

t h ey

a r e

l o ca l ly

d i spe r sed ) .

F ur thermo re ,

f r agmen ta t ion a nd e l emen t r e p re sen ta t ion p a t tern s i n d ica te t h a t i n a l l o f t h e r e g ions ( i nc lud ing t h e Mas s if C en t ra l ) h or se w as i n ten s ive ly p roce s sed . We m ay c onc lude t h a t h or se w as e xp lo i ted o n a n e ncoun te r b a s is , i n am anne r c on s is ten t w i th i t s e t ho logy . T he e v idence , t h e refo re , i n d ica te s t h a t a t l e a s t s ome h um ans w e re r e s iden t i n t h e A qu i ta ine B a s in d ur ing t imes o ft h e y ea r when r e inde e r w e re n o t a cce ss ib le o r n o t e xp lo i ted . T he i mp l ica t ion f o r p roposed m ode ls o f h e rd -fo l low ing , s uch a s a dop ted b y G o rdon ( 1988 ) i s o bv iou s a nd w i l l b e d i scu s sed mo re f u l ly i n t h e n ex t C hap te r .

6 3

CHAPTER 6 HUNTER -GAT HERER M ODELS . o bv iou s why a r chaeo logy h a s r e sponded f a vou rab ly t o e co log ica l t r ends i na n thropo log ica l t heory .

6 .0 .0 I n troduc t ion . T he s t udy o fp reh is tor ic h un te r -ga the re r s

A s ar e su l t o f empha s is o n e nv ironmen ta l

o f ten r e l ie s o n e t hnograph ic r e sea rch t op rov ide mode ls o f

r e la t ion s , t he s t udy o f s oc ia l r e la t ions a nd t h e c u l tura l

h uman s oc io ter r i tor ia l o rgan isa t ion . T he t h eore t ica l b a s is

s upe r s truc tu re ( s en su I ngo ld 1 980 :8 )h a s b een l ef tt os oc ia l

f o r

f o r

a n thropo logy ( bu t s e e C onkey 1 980 ) . C u l tu re e co log is t s

i n terpre t ing t h e p a s t i so f t en l ef t u n s ta ted , h oweve r . T h is

( and a l a rge n umbe r o f a r chaeo log is t s i n g ene ra l ) b y a nd

C hap ter w i l l p re sen t ag ene ra l mode l o f h un te r -ga the re r

l a rge h ave s imp ly a dop ted ag ene ra l , d escr ip t ive mode l o f p reh is tor ic h un te r -ga the re rs oc ia l o rgan isa t ion a sd eve loped

t h e

u se

o f e t hnog raph ic

a na logy

a s

a t oo l

b and o rgan isa t ion a nd e xam ine t h ea r chaeo log ica lc on tex to f i t s ' u se . G ene ra l s ub s is tence mode ls c ur ren t ly u sed b y a r chaeo log is t s s t udy ing t h e P le is tocene w i l l a l so b e

i n

L ee

&

DeVo re

( 1968 ) .

T h is

mode l

empha s ize s

p re sen ted , f o l lowed b y ad i scus s ion o f v ar iou s mode ls o f

r a the r t han l a nd tenu re ( s ee b e low ) .

h un ter -ga the re r s e t t lemen ta nd s ubs is tence i nS ou thwes te rn

d eve lopmen t o f m idd le r ange

F rance a pp l icab le t ot h e L owe r Magda len ian .

f o rma t ion p rocesse s , a nd g enera l t h eor ie s o fs e t t lemen ta nd

f l ex ib i l i ty , mob i l i ty a nd a cce ss t o r e source s , i . e . , l a nd -use W i th t h e f u r the r

t h eo ry ,

i n te re s t i n s i te

s ubs is tence s ys tem s , a r chaeo log is ts h ave b egun m app ing t h is mode l o f b and o rgan isa t ion o n to t h e a r chaeo log ica l

6 .0 .1 Hun ter -Ga therer R e search .

l a ndscape ( e .g . , D av id 1 973 ; B ahn 1 977 , 1 983 ; S p ie s s T he r e search

p r io r

s t a ted

t o t h e

i n te re s t

1 960 s ,

1 979 ; C onkey 1 980 ; B inford 1 980 , 1 982 , 1 983 ; G amb le

o f h un te r -ga the re r

a ccord ing

1 983 ;J och im 1 983 ; S t raus 1 983 ; Wh i te 1 985 , 1 987 , 1 990 ) .

t o B e t t inge r

( 1991 :47 ) , w a s n o t h un te r -ga the re r a dap ta t ions p er s e , b u t r a the r c u l tura l e vo lu t ion . F o r a r chaeo logy , t h e i mp l ica t ion

6 .1 .0 B and t heory : t he c ompos i te b and .

o f a n e vo lu t ionary p e r spec t ive i s t h a t mode rn h un te r g a the re r s ,

a s

r e pre sen ta t ive s

o f

a s t age

o f

S ince t h eg round -b reak ing Man t h e Hun te r s ympos ium ( Lee & D eVo re 1 968 ) , ag ene ra l mode l o fb and

c u l tu ra l

d eve lopmen t , c an b e c on s ide red v a l id mode ls f o r p a s t h un t ing a nd g a ther ing a dap ta t ion s . I n o the r words

o rgan isa t ion' , w i th d emograph ic , s pa t ia l a nd e conom ic

e vo lu t iona ry a n thropo logy e ncouraged a r chaeo log is ts t o

i mp l ica t ions , h as g a ined c ommon c ur rency i na n th ropo logy

p roceed b y d i rec t e t hnog raph ic a na logy .

a nd n o tab ly i na r chaeo logy . A ccord ing t ot h is mode l , b as ic h un t ing

s oc ie ty

i s c omposed

o f l o ca l

b ands ,

whose

e t hnograph ic

c ompos i t ion a nd t e r r i tory v ar ie s s e a sona l ly a nd wh ich f o rm

a na logy , u n suppor ted b yt h eo re t ica l mode ls , t os t udy Uppe r

p ar t o f al a rge r , e ndogamous , l i ngu is t ic c ommun i ty ( e .g . ,

T he

f e a s ib i l i ty

o f u s ing

P a laeo l i th ic h un t ing s oc ie t ie si nE u rope i sq ue s t ionab le . T he

L ee & D eVore 1 968 :8 ) . More s pe c if ica l ly , t h e re e x is ts a n

u se o fs pe c if ic a r c t ic a nd s uba rc t ic e t hnog raph ic mode ls f o r

o rgan isa t iona l

h i e ra rchy

t h es t udy o ff u l lg l ac ia l a dap ta t ions i nS ou thwe s tern F rance ,

" macroband")

2 )

f o r

( " rn ic roband s" ; " hun t ing g roup s") , a nd 3 ) s hor t l ived t a sk g roup s ( Lee & D eVo re 1 968 ; H e lm 1 969 ; R oge r s 1 969 ;

e xamp le ,

h a s

b een

c r i t ic ised

o n

t h e

b a s is

o f

i n compa t ib i l i ty b e tween t h e t wo s t udy e nv ironmen t s ( De lpech 1 983 :172 ) . S pe c ie sd ive r s i ty e ncoun te red i n Uppe r

o f

a n umbe r

1 ) t h e o f

r eg iona l b and

s ma l ler

l o ca l

( o r

b and s

D ama s 1 969 ; McKennan 1 969 ; L eacock 1 969 ; S lobod in 1 969 ; A sch n d ; S harp 1 977 ; I ngo Id 1 980 ; R iche s 1 981) .

P a laeo l i th ic f auna l a s semb lage s i n t h e S ou thwe s t i su n l ike a ny e ncoun te red t oday a mong a r c t ic o r s uba rc t ic p eop le s ( Sp ie ss 1 979 :134 ) .

T he r e g iona l b and i sa n e ndogamous u n i t ( Roger s 1 969 ; L eacock 1 969 ; McKennan 1 969 ; A sch n d ; h as

Myer s 1 986 ) c omposed o f an umbe r o f l o ca l b ands l i nked

a nd

b y t i e s o f k insh ip o r b y r e c iproca l r e la t ionsh ip s b e tween

h un te r -ga the re r s

b and m embe r s ( Maus s 1 904 -5 ; D ama s 1 969 ; B a l ick i 1 970 ;

( Be t t inge r 1 991 :49 ) . T he a dop t ion o f o p t ima l f o rag ing

A sch n d ; L ee 1 979 ; R iches 1 981) . B and o rgan isa t ion ,

More i n corpora ted

e l emen ts

n eofunc t iona l ism

i n to

r e cen t ly , o f t h e

a n thropo logy

e co log ica l s t udy

o f

t h eo ry

t h eory ( e .g . , F r ison 1 978 ; c on tr ibu to r s t o Win terha lde r &

t h e refore ,

Sm i th 1 981 ; B u tze r 1 982 ) f r om t h e b io log ica l s c ience s

d egre e o fa r t icu la t ion b e tween r eg iona l a nd l o ca l b and s i s

( MacAr thu r & P ianka 1 966 ; C harnov 1 976 ) i sa n e xamp le

v ar iab le , e .g . , t h e I nu i t r e g iona l b and d escr ibed b y Maus s

o fw ha t h a sb een c a l led c u l tura l e co logy , o rh uman e co logy

( 1904 -5 ) i sr e la t ive ly more " amorphou s" t h an t h e P in tup i r e g iona l b and d escr ibed i n Mye r s ( 1986 ) .

( Fr ison 1 978 ; L ee 1 979 :2 ; Win te rha lde r 1 981 :15 ; B u tze r

i s s uppor ted b y k insh ip a nd e xchange . T he

1 982 :22 ; B e t t inge r 1 982 :191 ; K eene 1 983 :139 ) . A m a jor i mp l ica t ion o f t h is " new e co logy" ( Be t t inge r 1 991 :49 ) f o r

L oca l b ands a r e o fv ar iab le s i ze , b u t a r e

a r chaeo logy i s empha s is o n t h e r e la t ions b e tween t h e

u sua l ly c omposed o f as ma l l n umbe r o f f am i l ie s o r p eop le

e nv ironmen t

t h e

( 2 -3 f am i l ie si n Maus s 1 904 -5 :87 ; 4 -8 i nS lobod in 1 969 :59 ;

t e chnoenv ironmen t af o cu s o f s t udy ( Be t t inger 1 982 :191) .

a nd

t h e

s oc ia l

s y s tem ,

m ak ing

2 0 -75 p eop le i n McKennan 1 969 :102 ; D ama s 1 969 :122 ;

S i nce t h e t e chnoenv ironmen t i sm a ter ia l ly r e cove rab le , i ti s

R oge r s 1 969 :26 ; 1 0 -30 p eop le i nL ee 1 979 :56 ) . T he s pa t ia l

6 4

d i sper sa l a nd s ma l l s i ze o fl o ca l b and s i sf r equen t ly s t re s sed

d rawn f r om b and t heory , n o tab ly s oc ia l a nd t e r r i tor ia l

a s a n a dap ta t ion t o s eve re e nv ironmen t s , o r o ne s i nw h ich

mob i l i ty , f l ex ib i l i ty a nd e nv ironmen ta l s ens i t iv i ty , h ave

r e source s a r e p a tchy , u neven ly d i s tr ibu ted , o ru np red ic tab le

i ndeed p roven u sefu l i n s t ud ie s o f Uppe r P a laeo l i th ic a dap ta t ions ( s ee b e low ) .

( e .g . ,

Mau s s

1 904 -4 :58 ,65 ;

R oge r s

1 969 :41 ;

L e e

&

D eVo re 1 968 ; S harp 1 977 ; H eff ly 1 981) . 6 .2 .0 G enera l S ub s is tence Mode ls i n A rchaeo logy . Hun te r -ga the re rb and s g o t h rough ac yc le o f a ggrega t ion a nd d i spers a l , w i th r eg iona l b and s f o rm ing

A s a l ready

s t a ted ,

c u l tura l e co logy i s

c ore s iden t ia l g roup s i n t he n uc lea ted o r a gg rega ted p ha se

p r imar i ly c oncerned w i th r e la t ions b e tween t h ee nv ironmen t

( Mau ss 1 904 -5 ; L ee & D eVore 1 968 ; H e lm 1 969 ; L eacock

a nd t h e s oc ia l s y s tem. T he e nv ironmen t i sp e rce ived a s

1 969 ; R oge r s 1 969 ; Dama s 1 969 ; L ee 1 979 ) . T h is c yc le i s

ad e term inan t o f s oc ia l a nd e conom ic p o ten t ia l ( J och im

t h e r e su l t o f a n i n terna l s oc ia l d ynam ism o f r epu ls ion a nd

1 976 :45 ; F r ison 1 978 :19 ; L e e 1 979 :117 ; B inford 1 982 :12 ;

a t trac t ion , o r " d ia lec t ic" , c ha rac ter is t ic o f h un te r -ga the re r

B u tze r 1 982 :7 ; B e t t inge r 1 982 :191 ; K e l ly 1 983 :277 , 3 02 ;

s oc ie t ie s ( Leacock 1 969 :3 ; L e e 1 979 :4 ; K eene 1 983 :143 )

G amb le 1 986 :35 ) .T he refo re , t h ec u l tura ls upe r s truc tu re , a n

wh ich may f unc t ion s oc ia l ly a s a m ean s o f r e so lv ing

a r chaeo log ica l ly

c onf l ic t ( Lee & D eVore 1 968 :9 ) .

i n d ire c t ly , t h rough i t s e f f e c t o n t h e s oc ia l s ys tem .

I nhe ren t t ot h e c yc le o ff i s s ion /fus ion i sa

u nre cove rab le

e n t i ty ,

i s s t ud ied

o n ly

C u l tura le co log is t sa re p r imar i ly c oncerned

f l u id i ty o r f l ex ib i l i ty i n s oc ia l a nd t e r r i tor ia l o rgan isa t ion

w i th i nve s t iga t ing t h ree m a in a rea s whe re t h ee cosys tem a nd

( e .g . , Mau s s

t h e s oc ia l s y s tem i n te rac t : 1 ) s e t t lemen t s ys tem s ( e .g . , J och im 1 976 ; H ef f ly 1 981 ; B u tze r 1 982 ) ; 2 ) d emograph ic

1 904 -5 :50 ; L ee & D eVo re 1 968 ; R oge r s

1 969 ; L eacock 1 969 ; B a l ick i 1 970 ; L e e 1 979 ; H ef f ly 1 981) . T h is i nhe ren t f l ex ib i l i ty , wh ich f a c i l i ta te s mob i l i ty a nd

a r rays ( e .g . , Sm i th 1 981) ; a nd 3 ) s ubs is tence s ys tem s ( d ie t

i n forma t ion e xchange , c an b e v i ewed a s a n a dap t ive , r i sk

c ompo s i t ion )( e .g . ,J och im 1 976 :11 ; W in te rha lde r 1 981 :22 ;

r educ ing s t ra tegy ( Hef f ly 1 981 ; R iche s 1 981 ; K e ene 1 981 ;

K eene

G amb le 1 986 ; i n te r a l i i ) .

p r inc ipa l a r ea s , t h e s ub s is tence s ys tem i so f ten v iewed a s h av ing

1 983 :139 ; c ausa l

B e t t inge r

p r ior i ty

1 982 :103 ) .

( e .g . ,

J och im

O f t h e se t h ree 1 976 :13 ;

K e l ly

Aggrega t ion a nd d i spe r sa l ( o r f i s s ion a nd

1 983 :277 ) . O ne o f t he mo re p owe rfu l t oo ls a r chaeo log is t s

f u s ion ) i s k eyed t o s ea sona l c hange s i n t h e e nv ironmen t

h ave a dap ted f o r s t udy ing s ubs is tence i so p t ima l f o rag ing

( Maus s 1 904 -5 ;D ama s 1 969 ; L eacock 1 969 ; R odge r s 1 969 ;

t heory .

McKennan 1 969 ; B a l ick i 1 970 ; H e lm 1 969 ; S harp 1 977 ) . " nuc lea t ing

T he re e x is t av ar ie ty o f g ene ra l mode ls

c en ter s" ( McKennan 1 969 :102 ) , a re s e en a s d e te rm inan ts

S ea sona l

c oncen tra t ion s

o f

r e source s ,

o r

w h ich c ome u nde r t h e umb re l la o fo p t ima l f o rag ing t heory .

f o r t he t im ing a nd s i ze o f t h e a ggrega t ion s ( e .g . , D ama s

A l l t h e mode ls h ave a n e nv ironmen ta l f o cus . T hey a l so

1 969 :122 ) . I n o the r words t h e p roduc t iv i ty o f t h e l o ca l e nv ironmen t p l ays a r o le i nd e term in ing t h e p a rame te r s o f

a s sume t h a t af undamen ta l l og ic , b a sed o n o p t im isa t ion , i s o pe ra t ive i n h un te r -ga the re r a dap ta t ion s ( J och im 1 976 :10 ;

s oc io ter r i tor ia l o rgan isa t ion ( e .g . , Mau s s 1 904 -5 ; S t ewa rd

B info rd

1 938 :236 , 1 977 :48 -49 ) . J us t h ow g rea t ar o le i s am a t te r

D urham 1 981 :229 -230 ; Sm i th 1 981 :53 ; H ef f ly 1 981 :162 ;

f o r d eba te ; t h eg ene ra l s ubs is tence mode ls r e v iewed b e low ,

B u tze r

f o r e xamp le , p lace a g rea t d ea l o f i mpor tance o n t h e

B e t t inge r 1 991 :106 ) . F ina l ly ,t h e mode ls u se t h e " un ive r sa l"

e nv ironmen t a s ad e term inan t o fs oc ia l o rgan isa t ion .

c ur renc ie s o fe ne rgy a nd t ime , b ased p r imar i ly o n t h e work

1 978 :81' ; 1 982 :258 ;

E a r le Ma r t in

o f MacAr thu r &

P ianka

1 980 :25 ;

R iche s

1 983 :612 ;

( 1966 ) ,

1 981 :227 ;

K eene

( J och im

1 983 :144 ;

1 976 ;

L e e

1 979 :43 , 4 43 ; H ef f ly 1 981 :143 ; W in te rha lde r 1 981 :20 ;

6 . 1 .1 C r i t ique s o f B and Theory .

Sm i th 1 981 :38 ; B u tze r 1 982 :20 ; K eene 1 983 :143 3' ) . C on tac t

b e tween

h un t ing -ga the r ing

e cono m ie s a nd o the r f o rm s o f e conom ic o rgan isa t ion i sa

T he s imp le s t o p t ima l f o rag ing mode l , t h e

much more p erva s ive p henomenon among h i s tor ic a nd

D ie t B read th mode l , w a s f o rmu la ted b y MacAr thur &

modern h un te r -ga the re r g roups t h an e a r ly e t hnographe r s

P ianka ( 1966 ) a s ac o s t re turn i n equa t ion . I n t h is mode l ,

s u spec ted . R e cons truc t ions o fh un te r -ga the re re conom ic a nd

i t em s a r e c ons ide red f o r i n c lus ion i n to a f o rage r 's d i e t o n

s oc ia l

t h e b a s is o ft h e ir r a te o fe ne rgy r e turn ( mea sured i ne ne rgy

o rgan isa t ion a re e s sen t ia l ly b a sed o n s t ud ie s o f

a ccu l tura ted p eop le ( Leacock n d :43 ; L e e&D eVo re 1 968 :7 )

r e tu r ned a s af unc t ion o f p repa ra t ion t ime ) r e la t ive t o t h e

n o

o ve ra l l r a te o f r e turn f o r ad i e t i n c lud ing a l l h ighe r r anked

l onge r

i nhab i t ing

" t he

o p t imum

p o r t ion

o f

t h e ir

t r ad i t iona l r ange " ( Lee & D eVo re 1 968 :5 ) . T he u se o f

t ype s o f f ood i t em ( mea su red i n e ne rgy r e turned a s a

modern e thnograph ie s f o r t h e s t udy o f p re -con tac t p er iod s

f unc t ion o fb o th p repa ra t ion a nd s e a rch t ime ) . I ft h e r a te o f

mus tt he refore p roceed w i th c au t ion ( I b id . ;L eacock 1 969 :1 ;

r e tu r n f o r t h e f ood i t em c ons ide red i s g rea te r t han t h e

McKennan 1 969 :94 ) .

o vera l l r a te o f r e tu r n , t h e i t em w i l l b e i n c luded .

A n

i mpor tan t

t h is

N ever the le s s ,

c r i t ica l

e va lua t ion

o f

( a nd

c oun te r in tu i t ive )

i mp l ica t ion

o f

i nequa t ion i st h a t af ood i t em i sc ons ide red i ndependen t ly o f

a ccu l tura t ion a nd f o cus o n t h e g ene ra l c ha rac ter is t ic s o ft h e

i t s a bundance ( Be t t inge r 1 991 :86 ) .

e t hnograph ie s c o l le c ted h a s r e su l ted i n ag ene ra l mode l u sefu l f o r u nde r s tand ing p a s t f orm s o f h un te r -ga the re r s oc ia l o rgan isa t ion ( L ee & D eVore

T he s ame b a s ic i nequa t ion i su sed i n t h e

1 968 :6 ) . C oncep t s

P a tch C ho ice mode l b u t i nt h is f o rmu la t ion , p a tch t ype s a r e 6 5

s ub s t i tu ted f o r i nd iv idua l f ood i t em s . T he i mp l ica t ions o f i n crea s ing a nd d ec rea s ing a bundance i n t h is mode l a r e s omewha t d i f f eren t f r om t h e D ie t B read th mode l s i nce

J och im ( 1976 ) i n t roduce s a " g r av i ty" mode l t oe xp la in s e t t lemen tp a t terns i nt e rm s o fs ub s is tence

r e sou rce a bundance w i l l a f fe c t t r ave l t ime b e tween p a tche s a nd t h e refore , p a tch c ho ice ( Be t t inge r 1 991 :90 ) . 3'

( ba sed o n a" goa l s a t isf ier " mode l ) . Wobs t ( 1977 ) u se s a n i d ea l iz ed h exagona l l a t t ice p a t tern o fs e t t lemen t p l acemen t ( wh ich max im ize s s oc ia l c on tac t ) a nd m ap s i to n to t h e

H is tor ic

a nd

p reh is tor ic

s ys tem s h ave b een s ucce ssfu l ly

s ubs is tence

l a nd scape ,

a l low ing

a n

e ne rge t ics

mode l

t o p red ic t

s t ud ied u s ing g ene ra l

" d is tor t ion s " o ft h e l a t t ice . Op t ima l f o rag ing mode ls c an b e

s ub s is tence mode ls s uch a s t h e D ie t B read th a nd P a tch C ho ice mode ls ( e .g . , F r ison 1 978 ; Y e sne r 1 981 ;

a pp l ied t o t h e d e scr ip t ion a nd a na lys is o f s e t t lemen t s y s tem s . S ee , f o r e xamp le , d e scr ip t ion s o fh un te r -ga the re r

Win te rha lder

1 981b ) .

Mo re

c omp lex

f o rmu la t ions

o f

s e t t lemen t s u s ing o p t ima l f o rag ing t h eory a s a n o rgan is ing p r inc ip le ( Fr ison 1 978 ; L ee 1 979 ; S i lbe rbaue r 1 981) .

o p t ima l f o rag ing t h eo ry ( i .e . , n on -con t ingency mode ls ) i n c lude t h e l i nea r p rog ramm ing mode l ( e .g . , K eene 1 981 ; Win te rha lde r ( b ind ing

1 986 ) .

I n t h is mode l ,

c ons tra in t s )

c an

b e

a dded d imens ions

c on s idered .

T he

6 .2 .2 C r i t ique s o f G enera l S ubs is tence Mode ls .

l i nea r

p rog ramm ing mode l a nd mode ls l i ke i tr ep re sen t a t temp t s t o a dap t a n e co log ica l mode l f o r a n thropo log ica l u se b y

T he re a re an umbe ro fd i ff icu l t ie si n he ren t i na dap t ing g ene ra l mode ls s uch a s o p t ima l f o rag ing t h eory

i n c lud ing

t ot h es t udy o fh uman s oc ie t ie s ( e .g . , Du rham 1 981 ; K eene

c oncep t s

s u ch

a s

r i sk

a nd

s ha r ing

( e .g . ,

W in te rha lde r 1 986 ) . T h is k ind o fa dap ta t ion i s an ece s sary

1 983 ; B e t t inge r 1 991) . T he s t anda rd o b jec t ion t ot h eu se o f

f e a ture

e co log ica l mode ls i n a n thropo logy i s t h a t b io log is t s a r e

o f

s c ien t if ic

c r oss -f er t i l isa t ion

( Keene

1 983 ;

B e t t inge r 1 991) .

d ea l ing w i th ag ene t ica l ly t r an sm i t ted s ys tem o fb ehav iour , w he rea s s oc ia l r ep roduc t ion d oe sn o tu nde rgo t h es ame k ind h un ter -ga therer

o f s e lec t ion 3 ( Keene 1 983 :142 ) . I n o the r word s , c u l tu ra l

s ub s is tence , wh ich r e semb le o p t ima lf o rag ing mode ls i nt h a t

O ther

t h ey c a lcu la te c os t -re tu rn s a nd s e ek o p t ima l s o lu t ion s t o

e co logy i s more c oncerned w i th e co logy t han w i th c u l tu re . T he a ppropr ia tene ss o fu s ing e ne rgy a s amea su re o ff i tne s s

s ub s is tence , i n c lude t hose p re sen ted b y J och im ( 1976 ) a nd

i s n o t u n ive r sa l ly a ccep ted , e ven w i th in t h e b i o log ica l

B info rd

p r inc ip le sd er ived f r om g ame s t h eo ry ,c ons ide r ss ub s is tence

s c ience s ( Keene 1 983 :143 ) . Howeve r , a s ah eur is t ic d ev ice , o p t ima l f o rag ing t heory r ema ins u sefu l f o r c ompa r a t ive

c ho ice s

a na lyse s o fh uman e conom ic b ehav iou r .

( 1978 ) . i n

mode ls

J och im 's

t e rm s

o f

o f

( 1976 )

mode l ,

c u l tu ra l ly

wh ich

d e term ined

u se s

c a lor ic

" s a t isf ac t ion l e ve ls " . B info rd 's ( 1978 ) a na lys is o fb u tchery p a t terns mode ls s ub s is tence d ec is ion s o n as im i la r s c a le .

I ti so bv ious t h a tv ar iab le su naccoun ted f o r mode ls p resen ted h e re ( e .g . , c u l tu ra l

T he Mod if ied G ene ra l U t i l i ty I ndex , o r M .G .U .I . , i s

i n t h e

e s sen t ia l ly a n e ne rge t ic s mode l ; h uman d e c is ions r ega rd ing

v a r iab le s ) e nsure t h a t t h e a s sump t ion o f o p t im isa t ion i s r a re ly m e t ( Du rham 1 981 :220 ; Sm i th 1 981 :63 ; Mar t in

b u tche ry a r e made r a t iona l ly o n ac os t re turn b a s is ( s ee B e t t inge r 1 991 :108 , F igu re 5 .8 ) .

s imp le

1 983 :613 ) . A s B e t t inge r ( 1991 :113 ) s t a te s , h oweve r : " 1 . .. 1 c ons truc t ive c r i t ic s o f o p t ima l f o rag ing t h eory s e e s u ch m isma tche s ( be tween e xpe c ted a nd o bserved b ehav iou r s )a s

6 .2 .1 G enera l Mode ls o f S e t t lemen t O rgan isa t ion .

o ppor tu in i t ie s t od eve lop b e t te r mode ls ." G ene ra l s ub s is tence mode ls c an b eu sed a s ab a s is f o r t h e mode l l ing o f t e r r i tor ia l o rgan isa t ion . V i ta -

T he

r ema in ing

c r i t ic ism

o f

o p t ima l

F inz i & H iggs ( 1970 ) i n t roduced t h e c oncep t s o f a nnua l

f o rag ing t heory i sp ragma t ic , n ame ly , t h ey a r e d i ff icu l t t o

t e rr i to r ie s a nd c a tchmen t a rea s ,

i . e . , e conom ic z one s

a pp ly t o g iven p reh is tor ic s i tua t ion s . T he f u l l r a nge o f

a round h uman s e t t lemen t s , i na na t temp tt oq uan t ify p a t ter n s

v a r iab le s ( i .e . , t h e s ubs is tence c ho ice s a va i lab le ) a r e r a re ly

o f s pa t ia l o rgan isa t ion i n t e rm s o f a n e ne rge t ic s mode l

k nown i na r chaeo log ica l s i tua t ion s , m ak ing t h e mode ls j u s t

( c os t -re tu rn

r e v iewed d i ff icu l tt oo pera t iona l iz e (W in te rha lde r 1 981 :19 ;

o f

e xp lo i ta t ion

o f

r e sou rces

w i th in

t h e

c a tchmen t a r ea ) . B inford ( 1982 :7 ) u se s p r inc ip le s d er ived

V i l la

f r om m idd le r ange r e sea rch , i n h i s c a se e s sen t ia l ly s i te

1 991 :109 ) .

1 983 :5 ;

R igaud

&

S imek

1 987 :60 ;

B e t t inge r

f o rma t ion p roce s se s , t o mode l s pa t ia l o rgan isa t ion i nt e rm s o fe conom ic z ona t ion : " c amp r ange s " , o r c a tchmen t a r ea s , a nd " annua l r ange s" , o ra nnua l t e rr i tor ie s . B inford 's ( I b id .)

6 .2 .3 C onc lus ion s : g enera l s ubs is tence mode l s .

mode l i sb ased o n a n a nnua l c yc le o f e conom ic a c t iv i ty w i th in a na nnua lr a nge c on ta in ing f unc t iona l ly d i f f e ren t ia ted s i te s , e . g . , r e s iden t ia l c amps a nd h un t ing c amps ( B inford

I ft h eg ene ra l t h eory a nd mode ls p re sen ted a bove a r e n o t w i thou t f l aws , c oncep t s eme rg ing f r om t h em

1 982 :11 ; b u t s e e K oe t je 1 987 :167 ) . A r e g iona l b and s '

h ave p roven u sefu l f o r a r chaeo log is t s s t udy ing t h e U ppe r

t e r r i to ry c on ta ins an umbe ro fa nnua l r ange s e xp lo i ted a t6 1 0 y ea r i n terva ls ( B info rd 1 982 :11) . L oca t iona l s h if t s

P a laeo l i th ic . Many a r chaeo log ica ls e t t lemen ta nd s ub s is tence

b e tween a nnua l r a nge s d o n o t c ons t i tu te ar eo rgan isa t ion o f

s t ruc tura l f l ex ib i l i ty i nh un te r -ga the re r s ys tem s , a so u t l ined

t h e s y s tem ( I b id .) . De c is ions c once rn ing s pa t ia l d i spe r sa l a nd s i te l o ca t ion a r e e s sen t ia l ly o p t im iz ing a nd m ay

i nt h e mode l o f b and o rgan isa t ion p re sen ted a bove . I n t h e

c or re la te w i th s e a sona l i n d ica to r s ( I b id .) .

s e t t lemen ta nd s ubs is tence w i l lb ep re sen ted wh ich u se t h ese a nd o the r c oncep t s d er ived f r om b and t h eory - w he the r

mode ls , f o r e xamp le , a s sume a n i nhe ren t mob i l i ty a nd

f o l low ing s e c t ion , mode ls o f Upper P a laeo l i th ic r e g iona l

6 6

i mp l ic i t ly o r e xp l ic i t ly . T he E ne rge t ics

mode ls ,

w h ich

u se

t h e

" un iver sa l" p r inc ip le s o f e ne rgy a nd t ime , h ave p red ic t ive v a lue

a s h eur is t ic d ev ice s ( Durham

1 981 :229 ;

a s soc ia t ion

o f i n te rcep t /m igra t ion

h un t ing w i th a c ommuna l h un t ing s t ra tegy i s b a sed o n e t hnog raph ic a na logy . Howeve r , c ommuna l h un t ing o f a

B u tze r

m igra tory a n ima l d oes n o t a lways i mp ly t h e s ame l e ve l o f

1 982 :229 ; K eene 1 983 :149 ; B e t t inge r 1 991 :105 ) i n t h e

s oc ia l i n tegra t ion , a s ar ev iew o f e t hnograph ic d a ta s how s

s t udy o f p a s t h un te r -ga the re r s ub s is tence a nd s e t t lemen t

( s ee c on tr ibu tor st oL ee & D eVore 1 968 ; S p ie s s 1 979 ) . T he

s y s tem s - p rov ided e nv ironmen ta l v ar iab le s a r e c on tro l led . F ur thermo re , t h e c oncep t s o f r a t iona l i ty a nd r e la t ive c os t ,

i n f e rence t h a t b a se c amp s f r om wh ich m ig ra tory g ame i s

d eve loped b y t h ese mode ls , a r e f r equen t ly i mp l ic i t i n

a dd i t iona l s uppor t ing e v idence .

i n te rcep ted a r e a ggrega t ion c amps c anno t b ed rawn w i thou t

m ode ls o f U ppe r P a laeo l i th ic h un te r -ga the re r o rgan isa t ion ( s ee b e low ) .

T he u se o f t h e a r ea l e x ten s ion o f s i te s a s a n i nd ica t ion o f s ea sona l a ggrega t ion f o r t h e p urpose o f h un t ing m igra tory g ame i sc r i t ic ised b y D e lpe ch , w ho n o te s

6 . 3 .0 R eg iona l Mode ls o f P a laeo l i th ic Hun ter -

t h a t t opography a nd a rea l s i te e x tens ion a r e n o tn e ce ssar i ly

G a therer S e t t lemen t a nd S ub s is tence .

a s soc ia ted w i th m igra t ion h un t ing ( 1983 :172 ) . T he u se o f d i ver s i ty a s a n i ndex o f s oc ia l i n te rac t ion ( a s soc ia ted w i th

6 . 3 .1 I n troduc t ion :

a n a gg rega ted b and ) i s c r i t ic ised b y C la rk ( 1980 :621) .

I n tegra t ing G enera l Theory a nd

A rchaeo log ica l Da ta .

S t y l is t ic d ive r s i ty may n o t b e au sefu l m ea su re o fr e g iona l i n terac t ion d ue t o t h e r a r i ty o f s t y l is t ica l ly m a rked o b je c t s

I n t h e f o l low ing s e c t ion w e w i l l e xam ine a n umbe r

o f a t temp t s

t o r e cons truc t

r e cove red

h un te r -ga the re r

f r om

L owe r

1 980 :621 ; S t raus

Magda len ian

1 980 :625 ; Hahn

c on tex t s

( C la rk

1 980 :623 ; G uen the r

s ubs is tence a nd s e t t lemen t o rgan isa t ion d ur ing t h e e a r ly

1 980 :622 ) . I t i s a l so a n a mb iguou s m ea su re o f s oc ia l

U ppe r P a laeo l i th ic i nt h e A qu i ta ine B a s in . I n mos t c a se s , a

i n te rac t ion , s i nce c omp lex i n te rac t ion s m ay h ave b e en

mode l o f c ompos i te ( r eg iona l / loca l ) b and o rgan isa t ion i s

s uppor ted b y s ymbo l ism i n p er ishab le m ed ia ( Gamb le 1 983 :206 ) o r i n med ia wh ich h ave b een i g no red ( s e e Wh i te

a s sumed ,

w i th i t s i mp l ied p a t te rn o f a gg rega t ion a nd

d i spe rsa l , c ond i t ioned a tl e a s t i np ar t b y e co log ica l f a c to r s

1 990b ) . A r chaeo logy , i n o the r word s , h a s f a i led t o p rove

( e .g . , s e asona l i ty ) . T he b a s is f o r a s sum ing t h a t t h is mode l

c onc lu s ive ly t h a t t h e e thnograph ic mode l o ft h e c ompos i te

o f b and o rgan isa t ion i sa pp l icab le t o S ou thwe s tern F rance

b and i sa pp l icab le t oS ou thwe s ter nF rance d ur ing t h e Uppe r P a laeo l i th ic .

d ur ing t h e P l en ig lac ia l r e s t s i np ar t o n t h e i d en t if ica t ion o f ap a t te rn o fa ggrega t ion a nd d i spe r sa l .

Aggrega t ion s i te s a s d ef ined b y b and T he

i den t if ica t ion

o f

P a laeo l i th ic

t h eory a r e s ome t ime sc onfused w i th b a se c amp s , a sd ef ined

a ggrega t ion s i te s i sb a sed o n t h e p re sence o f o ne o r mo re

i n B info rd 's ( 1982 ) f unc t iona l mode l o f h un te r -ga the re r

o f t h e f o l low ing a r chaeo log ica l f e a ture s . 1 ) E v idence f o r

s e t t lemen t s ( e .g . , C la rk 1 980 :621) . B info rd 's ( 1982 ) mode l

c oope ra t ive i n te rcep t , o r m igra t ion , h un t ing o f s e a sona l ly a va i lab le g ame ( Sp ie s s 1 979 :11 ; Wh i te 1 985 ; 1 987 :268 ) . 2 )

o f s e t t lemen t o rgan isa t ion , b ased o n ar eg iona l n e twork o f f unc t iona l ly d i f f eren t ia ted s i te s , d oe s n o t a ddre s s t h e

R e la t ive ly h i gh a s semb lage d i ve r s i ty ( Dav id

1 973 :288 ;

q ue s t ion o fa ggrega t ions i nt e rm s o fb and t h eory , h oweve r .

C onkey 1 980 :612 ) . 3 ) S i te e x ten s ion ( Dav id 1 973 :288 ;

I ti sn o t c l ea r h ow s ea sona l ly o ccur r ing r e s iden t ia l c amps , f o r e xamp le , m igh t r e la te t oa ggrega t ion s i te s i nt h e s ense

F re eman 1 973 :40 ; H ayden 1 980 ; C la rk 1 980 :621 ; Wh i te 1 987 :268 -9 ) . 4 ) E v idence o fh e igh tened s oc ia l i n te rac t ion ,

i n tended b y b and t heory . F ur thermore , t h e mode l i sb ased

i nt h e f o rm o fs t y l is t ic d ive r s i ty ( Fre eman 1 973 :40 ; Wob s t

o n p os t -con tac t e t hnography ( of t h e Nunam iu t , B info rd

1 977 :327 ; C onkey 1 980 :612 ; G amb le 1 983 :203 ; Wh i te

1 978 )

1 985 :152 ) .

a r chaeo log ica l

a nd

a l though t h e re a nd

i s a n

h i s tor ica l

a t temp t t o i n tegra te d a ta

( I b id .) ,

t h e

a ppropr ia tene ss o fu s ing t h is mode l f o r s t udy ing p reh is tor ic N one o f t h e se f e a tu re s o f f e r u nequ ivoca l

h un te r -ga the re r s i s u ncer ta in , e spe c ia l ly i n t h e l i gh t o f

p roof o ft h ee x is tence o f as oc ia l ly i n tegra ted r e g iona l b and ,

v ar iou s e t hnographe r 's c ommen t s , d i scu ssed a bove ( s e e

h oweve r . F i r s t o f a l l , " Sea sona l movemen t a nd o r w i ld

L eacock n d :43 ; L e e & D eVore 1 968 :5 ) .

h arve s t ing o f r e sou rce s a r e n o t t h e s ame a s a n o rgan ized a ggrega t ion a nd d i spe r sa l p a t tern" ( Conkey 1 980 :613 ) . I n

A n umbe r o f a r chaeo log is t s h ave u sed

t h e c a se o fw i ld h arve s t ing o f ad ense , s e a sona l ly a va i lab le

B inford 's s t udy

m igra tory s pec ie s s uch a s r e inde e r , " i n te rcep t" h un t ing i s

P a laeo l i th ic s e t t lemen t , e .g . , s ugge s t ing t h a t o pen a i r s i te s

o f ten

S t rau s

i n S ou thwe s tern F rance r ep re sen t s umme r h un t ing c amp s

t h a t s i te s whe re

( Gaussen 1 979 ; S p ie ss 1 979 ; Wh i te 1 985 , 1 987 ) . K oe t je ( 1987 :167 ) a nd C la rk ( 1980 :621) f i nd , h oweve r , t h a t t h e

i n f er red

1 983 :221) . r e indee r

( B inford

1 978 ;

S p ies s

T he a s sump t ion i s m ade

r ema in s

d om ina te t h e

f a una l

1 979 :11 ;

a s semb lage a r e

t empora ry , s e a sona l k i l l s i te s ( Bu rch 1 972 :363 ) . T h is p a t tern o f b ehav iou r i s mo re p rope r ly i n den t if ied t h rough

( 1982 ,

1 983 ) a s a mode l

f o r U ppe r

e conom ic f unc t ion o f p ar t icu la r s i te t ype s ( e .g . , o pen -a ir s i te s ) c anno t b e i n f er red u s ing s pa t ia l a na ly s is o r m ea su re s o f d ive r s i ty' . A ccord ing t o t h e se f i nd ings , t h e refo re ,

r e cons truc t ions o f t h e s t ruc tu re o f t h e p rey p opu la t ion a nd c ompa r ison w i th t h e " na tu ra l" p opu la t ion ( e .g . , S p ie s s

B inford 's mode l i sd i ff icu l tt oo pera t iona l ise . I nt h is c a se a t

1 979 ; P ike -Tay 1 991) a nd e l emen t r e pre sen ta t ion p a t ter n s

l e as t , a s K oe t je ( 1987 :167 ) s t a te s : t h e " pa t ter n s g ene ra ted

i nt h e f a una l r e co rd ( B info rd 1 978 ; B oy le 1 990 ) .

b y e t hnoa rchaeo logy a r e n o t a dequa te t o i n terp re t t h e 6 7

( a r chaeo log ica l ) r e cord ."

o f h e rd f o l low ing f o r t h e S ou thwe s t o f F rance . I n B ahn 's mode l ( I b id .) , h uman p opu la t ions f o l low r e indee r

T he f o l low ing s e c t ion w i l l d i scu ss a n umbe r o f mode ls o f e a r ly Uppe r P a laeo l i th ic s e t t lemen t

movemen t s 1 ) f r om t h e P er igo rd t o t h e P yrenee s , a nd 2 )

s y s tem s , l a rge ly b a sed o n r econs truc t ions o f s ubs is tence s t ra teg ie s . Many o f t he se r econs truc t ions a re b a sed o n

P yrenee s . e conom ic

u nproven a s sump t ions c once r n ing s oc ia l o rgan isa t ion a nd

( 1977 :245 ,

s i te f unc t ion ( above ) .

1 983 :185 ) ; 3 ) ac yc le o f p opu la t ion a gg lome ra t ion a nd d i spe r sa l ( 1977 :246 , 1 983 :185 ) ; 4 ) r a t iona l r e sou rce

f r om E a s t t o We s t , a l ong t h e n or thern f l ank s o f t h e

None the le ss , many o f t h em a r e

t e s tab le u s ing f a una l ma ter ia l a s am ean s o f e s tab l ish ing

T he a s sump t ions o f t h is mode l i n c lude : 1 ) s pec ia l isa t ion , i . e . , d ependence o n r e indee r 1 983 :179 ) ;

2 ) h uman

mob i l i ty

( 1977 :245 ,

s e a sona l p a t te rns a nd i d en t ify ing e conom ic s t ra teg ie s , s uch

m anagemen t ( 1977 :246 ) ; a nd f i na l ly , 5 ) l o ng -d is tance h erd

a s e conom ic s pec ia l isa t ion .

movemen t s f o r r e indee r ( 1977 :246 )3 6.

6 .4 .0 Mode ls o fP a laeo l i th ic s e t t lemen t a nd s ubs is tence :

B ahn

p red ic t s a c l u s te r o f r e indeer -

d om ina ted s i te s i nt h e P er igord a nd i nt h e l owe r P yrenee s , c or re spond ing t o s ea sona l a gg lome ra t ion s ( 1977 :246 ,

6 .4 .1 Herd f o l low ing mode ls .

1 983 :185 ) . Howeve r , t h is p red ic t ion c an a l so b e made i n H e rd -fo l low ing a s ap a t te rn o fe conom ic

t h e c a se o f " c a sua l m ig ra t ion h un te r s " ( S tu rdy 1 975 :73 ) ,

o rgan isa t ion d ur ing t h e Uppe r P a laeo l i th ic i ss ugge s ted b y

w ho a r ej u s ta s l i ke ly t os i tua te t h em se lve s a long m igra t ion

D av id ( 1973) , S turdy ( 1972 , 1 975 ) , B ahn ( 1977 , 1 983 ) a nd

c or r ido r s .

Go rdon ( 1988 ) , t o n ame b u t af ew . B u rch 's ( 1972 :346 )

h ypo the s is , t h e refore .

L oca t iona l

e v idence d oe s n o t v er ify

e i the r

o bse rva t ion , t h a t " peop le c anno t f o l low t h e h erd s " , i so f ten q uo ted i n c r i t ic ism o f t he h erd f o l low ing mode l ( e .g . ,

A rchaeo log ica l e xo t ic

c onne c t ions ma ter ia ls

s uch

a r e

a s

S p ie s s 1 979 :20 ; Wh i te 1 990a :14 ) . Howeve r , B urch h im se lf

s t y l is t ic s im i lar i t ie s a nd

s e en a s

n ow c i tes e v idence c on trad ic t ing h i s e a r l ie r s t a temen t

s uppor t ing a mode l

( Bu rch 1 991 :443 ) . I n f a c t , e t hnog raph ic e v idence s uppor t s h e rd f o l low ing a s av i ab le e conom ic s t ra tegy ( e .g G ordon

1 977 :250 ; s e e a l so Go rdon 1 988 ) . Howeve r , t h e c onnec t ions c i ted a r e amb iguou s ,s i nce e xo t ic ma ter ia ls m ay

1 981 ; B urch 1 991 P . I n a n a t temp t t oc l ar if y h i s p os i t ion ,

b e t h e p roduc t o fe xchange s n o t e mbedded i np rocu remen t

B u rch ( 1991 :440 ) n ow w r i te s :

a c t iv i t ie s (Wh i te 1 990a :26 -7 ) . A r t is t ic l i nks , c i ted a s p roof

o f Nor th -Sou th movemen t

( Bahn

o f mob i l i ty i nB ahn ( 1983 ) , a r eu sed i nJ och im 's ( 1983 :217 ) " Peop le who " f o l low t h e h erds " d o n o t a t temp t t o

s t udy t o s uppo r t a mode l o f d ec reased mob i l i ty . F ina l ly ,

k eep u pw i th as pec if ic s e t o fa n ima ls b u t move i n

c u l tu ra ll i nk sa nd s im i lar i t ie sa r ep red ic ted b yt he c ompo s i te

at ime ly manne r f r om t h e w in te r r ange o f t h e

b and mode l ( s ee a bove ) , w i thou t p os tu la t ing l ong -d is tance

b reed ing p opu la t ion i n t h e t a iga t o i t s s umme r

p opu la t ion

r ange o n t h e t undra ."

c ons ide red a no pen s ys tem , c r ea t ing a" cu l tura l c on t inuum "

movemen t s ,

s i nce

t h e

r e g iona l

b and

i s

( McKennan 1 969 :98 ) . T he

b a s ic

a s sump t ion

o f

t h e

h e rd

f o l low ing mode l i s , o fc our se , t h a t r e inde e r a r e m igra tory .

Ar e cen ta pp l ica t ion o ft h eh e rd -fo l low ing

T h is a s sump t ion i sn o t i n q ues t ion h e re ( s ee d i scus s ion i n

mode l t ot h e U ppe r P a laeo l i th ic i nt he S ou thwe s to fF rance

C hap te r 5 , a bove ) .

s eden tary a s o pposed t on omad ic t a randu s p opu la t ions , b u t

h a sb een a t temp ted b y Gordon ( 1988 ) . A sa bove , t h is mode l p red ic t s h uman m igra t ion f r om Nor thern w in te r r ange s t o

s imp ly aq ue s t ion o fd egree o f mob i l i ty ( Bu rch 1 972 :345 ) .

S ou ther ns u mme rr a nge s i nt h e A qu i ta ine B a s in d ur ing b o th

" The re i s n eve r a ny

q ue s t ion o f

t h e a P a laeo l i th ic

L owe r

a nd

Uppe r

Magda len ian .

G ordon

u se s

S tu rdy ( 1972 , 1 975 ) p ropose s amode l o f

s ke le tochrono logy t oo b ta in a b so lu te s easona ld e term ina t ion s

h e rd -fo l low ing

We s ter n

o f r e indee r mo r ta l i ty a s at e s t o f t h is mode l . O nce a ga in ,

G ermany , b a sed o n t he s ea sona l i ty a nd d i s tr ibu t ion o f

e conomy

i n

a s sump t ions i n c lude : 1 )e conom ic d ependence o n r e indee r ;

a r chaeo log ica l s i te sa t tr ibu ted t ot h e Magda len ian . T he s i te s

2 ) r e s iden t ia l mob i l i ty ; a nd 3 ) l ong d i s tance r e indee r

s urveyed b y t h e a u tho r ( I b id .) a re c on s ide red p a r t o f a

m igra t ions o u t o f t h e Dordogne t o P yrenean

s i ng le s e t t lemen t s ys tem o n t h e b a s is o f t h e ir c u l tu ra l

g round s ( Gordon 1 988 ) .

c on tempo rane i ty . T he i mp l ica t ion - t h a t h erd

f o l low ing

p roduce s

s e t t lemen t

a n

e x treme ly

s t ab le ,

e ven

s t a t ic ,

c a lv ing

Go rdon 's s amp le i n c lude s f a una l ma te r ia l

p a t te rn - i sd i ff icu l t t oa ccep t g iven t h e d ynam ic n a tu re o f

f r om

b o th h un ter -ga the re r a dap ta t ions a nd r e indee r m igra t ions .

c on temporaneou s o n t h e b a s is o f c u l tura l a t tr ibu t ions . T he u se o f c u l ture s t ra t igraph ic d e s igna t ions a s u n i t s o f s t udy

T he

a s sump t ions

o f

S turdy 's

mode l

a w ide

v ar ie ty

o f

e xcava ted

s i te s

c ons ide red

h a sb e en c r i t ic iz ed e l sewhe re a nd n eed n o tb er epea ted h e re

i n c lude : 1 )y ea r r ound h uman d ependence o n r e indee r a nd

( e .g . ,

2 ) ah igh d egree o f h uman mob i l i ty . T he se a s sump t ion s ,

1 991 :10 -11) .

C lark

1 980 :621 ;

D e lpech

1 988 :281 ;

P ike -Tay

s ha red b y B ahn ( 1977 , 1 983 ) a nd Gordon ( 1988 ) , m ay b e v a l id

f o r

We s te rn

G ermany

b u t

a r e

n o t

n e ces sa r i ly

G o rdon 's h ypo the s is tha t h uman r e indee r

a pp l icab le t oS ou thwe s tern F rance ( s e e d i scus s ion , b e low ) .

f o l lowe r s f o rm d i sc re te s oc ia l g roup s t h a t move w i th e qua l ly d i scre te h e rd s -i sb a sed o n h i s s t udy o f C anad ian

B ahn ( 1977 , 1 983 ) h a s s ugge s ted amode l 6 8

B ar ren land c ar ibou h un ter s( Gordon 1 975 , 1 981) . Howeve r ,

more t h o rough ly t e s ted r e g iona l ly t han h a s b een t h e c a se .

t h e d egree o f s oc ia l c l osu re i mp l ied b y G ordon 's mode l

Al a rge r v ar ie ty o f s i te t y pe s s hou ld b e i n c luded f o r s t udy

s e em s u n rea l is t ic g iven e t hnograph ic d a ta ( e .g . , L ee &

i no rde r t oa vo id g ene ra l is ing f r om wha tm ay b e ar e s tr ic ted

D eVore 1 968 :11 ; McKennon 1 969 :98 ) . T he i mpor tance o f

p or t ion o f al a rge r s y s tem ( s ee Wh i te 1 990a :17 ) .

ma in ta in ing i n forma t ion n e tworks i n s e a sona l ly v ar iab le e nv ironmen t s h a s b een e mpha s ized b y an umbe r o fa u tho r s ( Sah l in s

1 972 :221 ;

Wobs t

1 977 ;

Sm i th

1 978 ;

S easona l e v idence : t h e c a se a ga ins t h e rd -fo l low ing :

S p ie s s

1 979 :115 ; Moore 1 981 ; Sm i th 1 981 :43 ;G amb le 1 983 :205 ) . r e v iewed

F ina l ly , s ea sona l d a ta f o r t h e L owe r Magda len ian r e su l t ing f r om t h e p re sen t r e sea rch d o n o t s uppor t t h e h ypo the s is o f h e rd f o l low ing . R a the r , t h e

a bove ( S tu rdy 1 972 ; 1 975 ; B ahn 1 977 , 1 983 ; G o rdon 1 988 )

e v idence s ugge s t s t h a t h or se w a s e xp lo i ted b y h uman s

T he

h e rd -fo l low ing

mode ls

s ha re a n a s sump t ion o f h uman e conom ic d ependence o n

r e s iden t i n t h e Dordogne a nd t h e G ironde d ur ing t h e

r e indee r ( s ee a l so S t rau s 1 987 ) . T he p e rcep t ion o fr e indee r

h ypo the t ica l p e r iod o f r e indee r o u t -m igra t ion , i . e . , May -

a s e conom ica l ly d om inan t d ur ing t h e Uppe r P a laeo l i th ic

O c t . ( De lpech 1 983 :174 ) . I nt h e Ma ss if C en tra l a nd L ande s

m ay r e su l t f r om a n h i s to r ica l s amp l ing b i a s t owa rds c ave

s i te s s urveyed h or se w a s a ppa ren t ly a l so h un ted i n t h e

s i te s a nd r ock s he l ter s i n ar e s t r ic ted p or t ion o f t h e s t udy r eg ion , h oweve r ( R igaud & S imek 1 987 :55 ) .

a b sence o fs e a sona l c oncen tra t ions o fr e indee r . T he a l ter na t ive h ypo the s is i so f a more

D esp i te a ppea rance s , t h e a s sump t ion o f

g eog raph ica l ly r e s tr ic ted p a t ter n o f r e s iden t ia l mob i l i ty ,

r e indee r d om inance i s n o t a lways m e t i n S ou thwe s tern

a ccompan ied b y s e a sona l s h if t si nr e source u se ( e .g . , D av id 1 973 ; S p ie s s 1 979 ; J och im 1 983 ; Wh i te 1 985 , 1 987 , 1 990a ;

F rance . S ee f o r e xamp le S p ie ss ' ( 1979 ) d i scu ss ion o f A br i P a taud a nd D e lpe ch 's ( 1987 ) s yn the s is o f f a una l

S t raus 1 983 , 1 987 ; P ike -Tay 1 991) . V ar iou s mode ls o f

r epresen ta t ions . T he a va i lab i l i ty o fa l terna t ive r e source s i n

s e a sona l s e t t lemen t s h if t w i l l b e r ev iewed i nt h e f o l low ing

t h e P er igo rd - amosa ic e nv ironmen t ( De lpe ch 1 983 , 1 988 ;

s e c t ion .

D e lpe ch e a L 1 983 ) - w i l l n o t h ave m ade r e s iden t ia l s h if t s d ur ing p er iod s o f r e indee r d i spe r sa l an ece ss i ty ( S turdy

6 .4 .2 S ea sona l s e t t lemen t s h if t mode ls .

1 975 :74 ; S p ie s s

1 979 :138 ; Wh i te 1 990a :19 ; P ike T ay

T he P er igord .

1 990 :12 ) . T he a s sump t ion o f y ea r r ound d ependence o n r e indee r t h e refo re , i sn o t p roven f o r t h e P er igord .

J och im ( 1983 , 1 987 ) p re sen t s amode l o f r e s tr ic ted mob i l i ty f o rt h eU ppe rP a laeo l i th ic . I nt h is mode l ,

T he a s sump t ion t h a t r e inde er m igra ted

e co log ica l p re s sure d ur ing t h e f u l l g l ac ia l p er iod r e su l t s i n

l o ng d i s tance s f r om Nor th t o S ou th i nt h e A qu i ta ine B a s in d ur ing t h e U ppe r P a laeo l i th ic ( Bahn 1 977 , 1 983 ; G o rdon

e conom ic

s pe c ia l isa t ion

o n

s a lmon

r e source s

i n

1 988 ) i se qua l ly u nproven .

C ompre ssed z ona t ion i n t h e

o fb ig g ame h un t ing ( J och im 1 987 :327 ) . T he a s sump t ion i s

P yrenee s ( Freeman 1 973 ; S t rau s 1 983 ; B ahn 1 983 ) a nd s im i la r c ond i t ions i n t h e P er igord ( Sp ie s s 1 979 ; D e lpe ch

t o h uman r a nge r e s tr ic t ion , a s a r e su l t o f e xpand ing

S ou thwe s ter nF rance ( J och im 1 983 :217 )a nd i n tens if ica t ion m ade t h a t p opu la t ion p re s su re o ccur s i nt h e S ou thwe s t d ue

1 983 , 1 988 ; D e lpech e ta l . 1 983 ; Wh i te 1 985 , 1 990a ) , d o n o t s uppor t t h e h ypo the s is o f l o ng -d is tance h e rd

n or thern i c e s hee t s ( J och im 1 983 :217 ,

1 987 :321) . T he

p red ic t ions

a r e :

movemen t s , s i nce t h e r e indee r 's s e a sona l n u tr i t iona l n e eds

mob i l i ty , i n c rea sed t e rr i tor ia l i ty , a nd t h e a ppea rance o f

c ou ld b e m e t b y m in ima l movemen ts ( S tu rdy 1 975 :74 ) .

s e asona l p opu la t ion a grega t ion s ( I b id .) .

Gordon ( 1988 ) r e cons truc t s h e rd r a nge s b ased

o n

t h e

h ypo the s is

t h a t

w in te r

a nd

r e su l t ing

f r om

t h is

mode l

d ecrea sed

F auna l e v idence i n s uppor t o f i n ten s ive

s u mme r

s a lmon f i sh ing i ss c an ty ( Bahn 1 983 :184 ; Me l la r s 1 985 :274 -

s e a sona l i t ie s f r om t h e s t udy r eg ion m a rk t h e e x treme s o fa r a nge . Howeve r , h i sr e cons truc t ion o f No r th -Sou th r e inde e r

2 81 ; J och im 1 987 :326 ; P ike -Tay 1 991 :15 ) . E v idence o f

m igra t ion i su nsuppor ted b y D e lpe ch 's m e tr ica l a na lys is o f

d r iv ing f o r ce f o r t h e mode l , i s amb iguou s . T he a ppa ren t

r e indee r s ugge s t ing s epa ra te P er igo rd ian a nd P yrenean p opu la t ions ( De lpe ch 1 983 :163 ; Wh i te 1 990b :70 ) . F oe ta l

p opu la t ion i n crea se ( J och im 1 987 :323 ) , m ay a l so r e pre sen t

b one a nd a du l t ma le b o is d e mas sac re s ugge s t s umme r

as h if t i n s e t t lemen t p a t ter n ( e .g . , Wh i te 1 985 ) o r s ome

r a nge s n ea rer t h e Do rdogne ( De lpech 1 983 :165 ) . A s a n

o the r h i s tor ica l p roce s s ( J och im 1 987 :323 ) .

i n crea sed d emograph ic p re ssure i n t h e S ou thwe s t , t h e mu l t ip l ica t ion o f s i te s , w h ich g i ve s r i se t o t h e n o t ion o fa

a l terna t ive h ypo the s is D e lpech ( 1983 :165 ,174 ) s ugge s t s A rchaeo log ica ls uppor tf o rJ och im ' smode l

s hor t m ig ra t ions w i th in t h e c ore a r ea o ft h e Dordogne ( t he G ironde i sd e s igna ted ap er iphe ra l a rea o f t h e r a nge ) w i th

i n c lude s i n tens if ica t ion o fa r t is t ic a c t iv i ty , a nd e v idence o f

p o ss ib le w in te r r ange s i n t h e Ma s s if C en t ra l o r m idd le

i n cre a sed s o c ia l i n tegra t ion a s soc ia ted w i th t h ed eve lopmen t

v a l ley o f t h e Do rdogne .

o f s ma l le r o r more c l osed g roup s a nd a s soc ia ted t e r r i tor ie s ( J och im 1 983 :217 , 1 987 :328 ) . A s a l ready n o ted a bove ,

T he t h ird a s sump t ion , s ha red b y S turdy

h oweve r , i n tens if ica t ion o f a r t is t ic a c t iv i ty m ay s imp ly b e

( 1972 , 1 975 ) , B ahn ( 1977 , 1 983 ) , a nd G ordon ( 1988 ) , i so f

u nre cogn ised p r ior t o t h e L ower Magda len ian ( Gamb le

ah igh d egre e o fh uman mob i l i ty .T h is a s sump t ion c onform s

1 983 :206 ) . F ur thermo re , t h ea ppearance o fc ave a r t u sed a s

t o e xpe c ta t ions g ene ra ted b y b and t h eory , b u t s hou ld b e

s uppo r t ing e v idence f o r i n crea sed s eden t ism b y J och im ( 1983 :217 , 1 987 :328 ) , i su sed t o s uppor t ah i gh ly mob i le 6 9

mode l o f p opu la t ion movemen t i n B ahn ( 1977 , 1 983 ; s e e

a nd r ock s he l te r s .

b e low ) .

e conom ic o rgan isa t ion o n s ea sona l s i te u se a re e xp lored

T he e f f ec t s o f t wo b a s ic f o rm s o f

S p ie s s( 1979 )p re sen t s amode lo fs e a sona l r e s iden t ia l s h if t s f o r t h e Aur ignac ian /ea r ly P er igo rd ian i n

as e a sona l r ound s mode l , p a t te r n s 1t o 4 ) . A r e levan t

t h e P er igo rd . S p ie s s ( 1979 :234 ) s ugge s t s as h if t f r om c ave s i te sa nd s he l ter s i nt h en ar row v a l leys o ft h eP er igord , l i ke

s e a sona l r ound s mode l d oe s n o t n e ce ssar i ly p roduce a

( e .g . , h erd f o l low ing , " nomad ic" p a t ter n s 5a nd 6 , v e r su s f e a tu re o f t h e ir d i scus s ion i st h e ir d emons t ra t ion t h a t t h e s harp ly d i f f e ren t ia ted p a t tern o fs easona l s i te -u se .

t h e V ezk re , t o o pen -a ir s i te s a nd u npro tec ted v a l leys , o r F reeman ( 1973 :40 ) p red ic t s ap a t tern o f s ea sona l a l t i tud ina l s e t t lemen t s h if t s i nt h e P yrenee s . H e i s

a l terna t ive ly t os i te s l i ke L a Qu ina , i nt h e C ha ren te ( Sp ie ss 1 979 :233 ) .

u nab le t oi d en t ify t he p red ic ted p a t tern o fs ea sona l h un t ing f o l low ing

c amp s a nd l ow land b ase c amps e xpe c ted u s ing af unc t iona l

a s sump t ions : 1 )e a r ly Uppe rP a laeo l i th ic h un ter sw e re b and

mode l o f s e t t lemen t o rgan isa t ion , h oweve r ( I b id .) . S t rau s

o rgan ised ( 1979 :184 ) ;2 )h un te r sm ade r a t iona l ,m ax im iz ing

( 1983 )a l so p ropo se s amode l o fl im i ted s ea sona l s e t t lemen t

S p ie s s

( I b id .)

m ake s

t h e

c ho ice s ( 1979 :134 ) ; 3 ) t h e S ou thwe s t o f F rance w a s a

s h if ts

p er ig lac ia l , mosa ic e nv ironmen tw i th h igh s pe c ie s d i ve r s i ty

a va i lab i l i ty o f r e indee r a nd o the r g ame . I n S t raus ' mode l

i n

t h e

( 1979 :134 ) ; 4 ) i n mo sa ic e nv i ronmen ts u ngu la te s d o n o t r e ach m ax ima l c oncen tra t ions b u t a r e r e la t ive ly d i spe r sed

Duf aure a nd Duru thy )a r ep red ic ted f a l l /w in te ro ccupa t ions ,

( 1979 :134 ) ; a nd 5 )h uman h un te r sa r e t h e refo re e xpe c ted t o

a nd s umme r r e s idence s a re p red ic ted i n t h e u p land s o r

b e more e c le c t ic i n mosa ic e nv ironmen t s ( 1979 :247 ) , a nd

c oa s ta l a rea s .

( 1983 :221 ,

P yrenee s ,

c or re la ted

w i th

t he

s e a sona l

1 987 :163 ) l ow a l t i tude s i te s ( s uch a s A br i

s pa t ia l ly d i spe r sed ( 1979 :248 ) . C lo t tes

( 1987)

b e l ieves

e v idence

s uppor t ing b o th e conom ic s t ra teg ie s ( he rd f o l low ing

S p ie s s ( 1979 ) p red ic t s t h e e x is tence o f

a nd

s e asona l r ounds , i . e . , b a se c amps w i th s pec ia l ized h un t ing

s ma l l b and s o f 2 0 -30 p eop le , as e a sona l p a t te rn o f s i te o ccupa t ion , a nd a l a ck o f s pe c ia l ised h un t ing t e chn ique s .

c amp s" ) e x is t s i n t h e P yrenee s . F ur the rmore , h e s t a te s

A ccord ing t o t h e a u thor ( I b id .) , t h e se p red ic t ions a r e

t h a t t h e t wo s t ra teg ie s a re n o t e xc lus ive ( 1987 :305 ) , b u t

a c tua l ized a t Abr i P a taud . A l though r e indee r a r e i mpor tan t

r a the r , w ere b o th u sed i nt he P yrenee s - i nk eep ing w i th t h e

a s ap ropor t ion o fu ngu la te MN I a tA br i P a taud" , s pec ie s

c omp lex i ty a nd d ive r s i ty o f t h e r eg iona l e nv ironmen t

d ive r s i ty i sh igh ( Sp ie ss 1 979 :134 ) i nd ica t ing a n e c lec t ic

( 1987 :305 , 3 25 ) .

h un t ing s t ra tegy . T he a ge a nd s e x d i s tr ibu t ions o fr e indee r r ema ins o f f e r n o e v idence o f d r ive s o r o the r s pe c ia l ised

" Ce s d ive r se s s t ra teg ie s n e s on t d 'a i l leu r s p a s e xc lu s ive s

h un t ing t a c t ics a tt h e Abr i P a taud ( Sp ie ss 1 979 :185 ) .

l e s

u nes

d e s

a u t re s ,

s ur tou t

d an s

l ' env ironnemen tp yreneen ,c omp lexe e td iver s if ie ." ( C lo t te s 1 987 :305 ) .

T he p a t ter n o fs e a sona l i ty a tA b r i P a taud d oe s n o t c or re la te w i th e i the r c hange s i n c u l tu ra l p ha se s ,

6 .4 .3 D iscu s s ion : S ea sona l E v idence .

s i te e x ten s ion o r v ar ia t ion i n t h e r e la t ive p ropor t ion o f r e indee r i n t h e t o ta l u ngu la te a s semb lage s ( Sp ie s s

I n t h e a rchaeo log ica l mode ls r ev iewed

1 979 :186 ) . S p ie s s ' d a ta s ugge s t , t h e refo re , t h a t ap a t tern o f s ea sona l r e s iden t ia l s h if t s o ccur red d ur ing t h e e a r ly Uppe r P a laeo l i th ic a sp ar to f al ong term r eg iona l a dap ta t ion i nt h e

a bove e v idence o f s ea sona l ly ma rked s i te o ccupa t ions i s

P e r igo rd . I n as t udy o f s ea sona l mor ta l i ty p a t ter n s f o r r ed d ee r f r om Uppe rP er igo rd ian c on tex ts , P ike -Tay ( 1991 :12 )

S turdy 1 975 ; S p ie ss 1 979 ; Wh i te 1 985 , 1 987 , 1 990 ; S t rau s

s uppor t s t h is mode l .

t ob e v er if ied : as ke le tochrono log ica l s t udy o fa l l u ngu la te s

p red ic ted ( Bo rde s e t a L 1 972 , p a t ter n 2 ; F reeman 1 973 ; 1 983 ; Gordon 1 988 ) . A t Abr i P a taud t h is p red ic t ion s e em s s ugge s t s

m a rked

f a l l

a nd

w in te r

o ccupa t ion

( Sp ie s s

1 979 :192 , T ab le 6 .2 ) . Howeve r , a par t f r om s i x r e indee r ,

Wh i te ( 1985 , 1 987 , 1 990a ) p ropose s a mode l o f s e a sona l s e t t lemen t f o r t h e B adegou l ian i n t h e

o n ly t wo h o r se s ,

P er igo rd ,

s ucce s sfu l ly s e c t ioned i n S p ie s s ' s t udy.°( I b id .) . T he s ma l l

b a sed

o n

a n

o bserved

c onnec t ion

b e tween

t wo r ed d ee r a nd o ne b ov ine w e re

l o ca t iona l d a ta a nd a r ea l s i te e x ten s ion a s a n i n d ica tor o f

s amp le o f u ngu la te s o ther t han r e indeer ,

a ggrega t ion .

p o ten t ia l ly r e spons ib le f o r t h e l im i ted s e a sona l d i s tr ibu t ion

L ike S p ie s s ( 1979 ) , Wh i te ( 1990a :21-22 )

t h e refo re ,

i s

o fp rey mor ta l i t ie s .

p ropose s as h if t f r om w in te r c amp s s i tua ted i n s he l te red v a l leys s uch a s t h e V ez : e re , t o s umme r c amp s s i tua ted i n

S ea sona l e v idence f r om s ke le to ch rono logy ,

i n te r f luv ia l a r ea s , o r p e rhap s w e s t o ft h e Do rdogne . L a rge s i te s s i tua ted a t c ro ss ing p o in ts o n t h e V ef e re r i ve r a r e

p resen ted h e re , d oe s n o t s uppor t a mode l o f s ea sona l ly

i n f er red f a l la nd /o rs pr ing a gg lome ra t ion s , o r ien ted t owa rds

r e s tr ic ted s i te o ccupa t ion s . T he re i se v idence f o rc on t inu i ty

t h e i n te rcep t ion o fm igra t ing r e indee r h e rds ( I b id .) . As h if t

o f h uman

i n s ubs is tence b a se ,

r e s idence s h if t s - i ft h e l a t te r a re c or re la ted w i th r e indee r

a ccompany ing s e a sona l r e s iden t ia l

o ccupa t ion

b eyond

t h e

p red ic ted

s e asona l

movemen t s4 1 -a t mo s t o f t he s i te s s amp led . T h is p a t tern

move s , i sp red ic ted (Wh i te 1 990a :23 ) .

r e semb le s B o rde s e ta L ( 1972 :28 ) p roposed p a t te rn 2 . T he s i x

f a c t t h a t s o m any a na lyse s h ave c oncen tra ted o n t h e

p a t terns o f s e t t lemen t u se f o r t h e P er igo rd , b a sed o n

s e a sona l mor ta l i ty p a t tern s o f r e indee r ( e .g . , a t Duru thy , A rambourou 1 979 ) a nd s i tes c on ta in ing h igh p ropor t ion so f

B o rde s

e t a l .

( 1972 :28 )

p ropo se

e v idence f o rc on t inuou so rp er iod ica l r e -occupa t ion o fc ave 7 0

r e indee rr ema ins ( e .g . , B ahn 1 977 :246 )m ay a c coun tf o rt h e

Howeve r , s i nce m igra t ion p a th s c an b e v a r iab le , o ne o ft h e

a ppa ren t l im i ted s e a sona l d i s tr ibu t ions o b served i nt h ep a s t .

r equ iremen t s o f i n te rcep t h un t ing i sa ccura te i n forma t ion

6 .5 .0 C onc lu s ion s .

b and s

d ur ing t h e h e rd mon i tor ing s t age s . T h is s ugge s ts t h a t l o ca l i n S ou thwe s te r n F rance w e re

i n te rac t ing o n

a

r eg iona l b a s is . T he re e x is t s a s y e t n o i n con trover tab le p roof t h a t t h e c ompo s i te b and mode l i s a pp l icab le t o

T he mos t o bv ious b ar r ie r t o i n terp re t ing

S ou thwe s tern F rance d ur ing t h e l a s t f u l l g l ac ia l p e r iod .

t h e c ur ren t s amp le i s t h e t ime span i n vo lved ( s ee Wh i te

N eve r the le s s , an umbe ro fp r inc ip le so fs o c ia la nd t e r r i tor ia l

1 987 :264 ) . We a re u nab le t o e s tab l ish w he the r t h e s i te s

o rgan isa t ion d eve loped b y

s amp led a re t r u ly c on tempora ry , i . e . , p ar t o f t h e s ame

b and

t h eory

f o rm

a b a s ic

c omponen t o f a r chaeo log ica l mode ls . C oncep t s s uch a s

s y s tem , b eyond e s tab l ish ing t h a t t h ey a r e p ar t o f as im i la r

s oc ia l f l ex ib i l i ty a nd mob i l i ty , f o r e xamp le , a r e k ey t o

c u ltura l p a t tern ( t he L owe r Magda len ian ) i n t h e s ame

mode ls o fh e rd -fo l low ing a nd s e a sona l r e s idence s h if ts ( s ee

c l ima tochrono log ica l p er iod ( t he p l en ig lac ia l ) . Dur ing t h is

a bove ) .

p er iod , n ume rous s e t t lemen t s y s tem s h if t s a r e p red ic tab le e . g . , a t ar a te o fe very 6 -10 y ea r s i nB info rd 's ( 1982 , 1 983 ) T he

c oncep t

o f a s e t t lemen t

s y s tem

mode l . T he c omp lex i ty o f t h e s i tua t ion i s o bv iou s' . Mo reove r , t h e s i te s s amp led h ere a r e a l l r o ck s he l te r s a nd

i n vo lv ing f unc t iona l ly d i f f e ren t ia ted s i te s ( e .g . , F reeman 1 973 ; B info rd 1 982 , 1 983 ; S t rau s 1 983 , 1 987 ; Wh i te 1 985 ,

m ay o n ly r epresen t ap a r t o f h ypo the t ica l ly l a rge r s y s tem ( Conkey 1 980 :611 ; Wh i te 1 990a :19 ) .

1 987 , 1 990 ) s t em s f r om t h e se p r inc ip le s o f f l ex ib i l i ty a nd mob i l i ty , a nd a n a s sumed p a t tern o f g roup f i ss ion a nd f u s ion , i n t u rn d ependen t o n r e sou rce a va i lab i l i ty . A s m en t ioned a bove , h oweve r , t h e re a r ed i ff icu l t ie si nhe ren ti n

T he l im i ta t ions o f t h e d a ta c ur ren t ly a va i lab le f o r s t udy a nd t h e p rob lem s i nhe ren t i n s amp l ing

t e s t ing s i te f unc t ion i nt h e a r chaeo log ica l r e co rd . Howeve r ,

f a una l m a ter ia l f r om b o th o pen a i r a nd s he l te red c on tex t s

i fa r chaeo logy h a s f a i led t od ev ise t e s t s f o r t h e h ypo the se s

m ake s r e conc i l ing t h ese d i ff icu l t ie s an ea r i mpo s s ib i l i ty .

t h a te me rge f r om t h eg ene ra l t h eor ie sa nd mode ls p ropo sed ,

T h is f a i lure c an b e s e en a s p ar t o f al a rge r f a i lu re i n

s ome o ft h e a s sump t ions i n vo lved a r e t e s tab le . D irec t ly o r

a r chaeo logy t om a tch g enera l t h eory t ot h e l im i ted d a ta se t s

i n d ire c t ly , d e term in ing t h e s e a son o f s i te o ccupa t ion i sa

a va i lab le ( Be t t inge r 1 991 :82 ) .

p a r t ia l t e s t o f a l l o f t h e a bove a s sump t ion s . A p a t ter n o f s e a sona l ly r e s tr ic ted s i te o ccupa t ion s , f o r e xamp le , i s a

I tm ay , t h e refore , b e d i ff icu l t t os e e h ow a r chaeo log ica l d a ta c an b e u sed r e l iab ly t o m ea sure p a s t

p l aus ib le t e s t o ft h e d egree o f mob i l i ty o ft h e s ys tem .

e conom ic a nd s oc ia l b ehav iour ( R igaud & S imek 1 987 :48 ) . S ea sona l d i f f e rence s i nr e sou rce u se a r e a

N eve r the le ss , a s i s a ppa ren t h e re , i ti sp os s ib le t o d raw

t e s to ft h ed eg ree t ow h ich t h e h uman s ub s is tence s ys tem i s

m ean ingfu l c onc lus ions f r om t h e s t udy o f f a una l m a ter ia l

d e term ined b y v a r ia t ions i nt h e a va i lab i l i ty o f g iven p rey .

w i th r ega rds t o mode ls o f s e t t lemen t a nd s ub s is tence

S ome o ft h e a s sump t ions g ene ra ted b y g ene ra l s ubs is tence

s y s tem s a nd p a t terns o f s e asona l mob i l i ty . T he se p a t ter n s

mode ls s uch a s o p t ima l f o rag ing , e . g . , t h e o p t im isa t ion o f c os t s a nd b enef i ts , c an b e a s se ssed w i th t h is s e a sona l d a ta .

a r e , o fc our se , g rounded i nh uman b ehav iour .

T he d egree o fe conom ic s pe c ia l isa t ion a nd d ependence o n ag i ven r e sou rce c an a l so b e e xam ined . T h is

r e sea r ch

h a s

p re sen ted

e v idence

s ugge s t ing e x tended s e asona l s i te -use f o r an umbe r o fs i te s i n t h e A qu i ta ine r e g ion' . T h is r e su l t i sn o t p red ic ted b y b and t h eory o r b y t h e v a r ious a r chaeo log ica l

mode ls

d i scus sed ( a bove ) , g iven e nv ironmen ta l c ond i t ions b e l ieved t o h ave e x is ted d ur ing t h e f u l l g l ac ia l p er iod

i n t h e

A qu i ta ine B a s in . I n a n e nv ironmen t w i th s t rong ly m a rked s e a sona l i ty , t h e mode ls o f b and o rgan isa t ion , s ubs is tence a nd s e t t lemen t , r e v iewed a bove , p red ic t s e a sona l r e s idence s h if t s . I n s tead , t h e e v idence p re sen ted h e re s ugges t s t h a t s e a sona l s h if t s i n t h e h un t ing s t ra teg ie s e mp loyed , r a the r t h an r e s iden t ia l s h if t s , o ccur red w hen t h e r e source b a se s h if ted ( e .g . , a f te r t h e p er iod o fr e indee r m ig ra t ion ) r a the r t h an a s ar e su l t o f r e s iden t ia l s h if t s ( s e e d i scus s ion i n C hap te r 5 , a bove ) . T he s oc ia l c o r re la te s o f t h e se s e a sona l s h if t s i n h un t ing s t ra tegy , f r om e ncoun te r t o i n te rcep t h un t ing ( s e e C hap te r 5 , a bove ) , s t ra teg ie s

a r e

n o t ,

o f t h em se lve s ,

a r e s ub t le .

I n te rcep t

e v idence

o f s oc ia l

i n te rac t ion o n t h e s c a le o f t h e c ompo s i te , r e g iona l b and' . 7 1

CHAPTER 7 CONCLUSION . 7 .0 C emen tum A na lys is

o f p r ima ry g row th l a ye r s i sa n o n togene t ic ( e ndogenou s ) p roce s s , t r igge r ed b y e xogenou s e ven t s ( e .g . , s e a sona l

7 .0 .1 G row th S t ruc tu re s i nD en ta l C emen t : E .

e nv ironmen ta l i n f luence s ) . O b se rva t ion s o f c emen tum f o rma t ion s i nh o r se t e e th a r e c on s is ten t w i th t h ed epo s i t ion o f as i ng le , a nnua l p r ima ry c emen tum l a ye r , c on s is t ing o f

c aba l lu s . T h ree t ype s o f g row th s t ruc tu re a r e o b servab le i nt h e d en ta l c emen t o f E . c aba l lu s : LAGs ( o r L ine so fA r re s t ed G row th ) , a nnu l ia nd g row th z one s . T he se

o ne g row th z one p lu so ne LAG ( a s soc ia ted w i th a na nnu lu s , w hen p re sen t ) . P r ima ry l a ye r s i nc emen t a r e u sed f o r t h e d e term ina t ion o fs e a son o fd ea th o fh or se .

d i f f e ren t s t ruc tu re s a r e d i s t ingu ished o n t h e b a s is o f t h e ir i n t r in s ic o p t ica l q ua l i t ie s : LAGs a r e t r an s lucen t , g row th z one s a r e o paque a nd i s o t rop ic , a nd a nnu l i a r e l e s s o paque t h an z one s a nd f a in t ly a n iso t rop ic . S ince g row th z one s i n c emen t r e p re sen t ap er iod o f a c t ive t i s sue f o rm a t ion , t h ey a r e g ene ra l ly w ide r t h an a nnu l i o r LAGs , wh ich r e p re sen t e p isode s o f s l owe r g row th a nd g row th c e s sa t ion , r e sp ec t ive ly .

7 .0 .2 O b serva t ion o fG row th L aye r s . V a r iou s t e chn ique s f o r o b ser v ing g row th s t ruc tu re s i nc emen t e x is t : 1 )u nde ca lc if ied t h in s e c t ion sa r e v i ewed u nde r t r an sm i t ted l i gh t ; 2 ) u nde ca lc if ied t h in s e c t ion s a r e m ic ro rad iog raphed ; 3 ) p o l ished t h ick s e c t ion s a r e v i ewed u nde r r ef le c ted l i gh t ; 4 ) p o l ished a nd e t ched t h ick s e c t ion s a r e v i ewed u nde r S cann ing E lec t ron M ic ros cope ; 5 )d e ca lc if ied , s t a ined t h in s e c t ion sa r ev i ewed m ic roscop ica l ly . A c o r re spondence e x is t s b e tween t h e

Sma l l d i f f e rence s i n t h e r e la t ive p ropor t ion s o f m ine ra l a nd o rgan ic c omponen t s o f t h e v a r iou s t ype so fc emen th ave b een o b served i n an umbe ro f t e r re s tr ia l s pec ie s . T he se d i f f e rence s a r e b e l ieved t ob e l e s s i mpor tan t t h an t h e d i f f e rence s i n t h e b a s ic p a t te rn o f o rgan isa t ion o ft h ec o l lagen m a tr ix c ompo s ing e a ch t ype o f g row th l a ye r . G row th z one s c on ta in r e la t ive ly m o re c e l ls a nd t h e i rc o l lagen f i b re m a tr ix i sg ene ra l ly s im i la rt ot h a to f " woven " b one . I no the r word s , t h e f i b re s i ng row th z one s a r e a r ranged w i thou t a ny s pa t ia l o rde r ( o r " woven" ) , c au s ing i s o t ropy . T owa rd s t h e e nd o f ag row th z one , t h e

g row th l a ye r s o b served i n e qu id c emen tum u s ing e ach m e thod , i . e . , t h e se t e chn ique s a r e e qu iva len t f o r t h e p urpo se s o f v i ew ing p r ima ry g row th s t ru c tu re s i n h o r se c emen t . F o r t h e p u rpo se o f v i ew ing p r ima ry g row th l a ye r s , s oph is t ica ted t e chn ique s s u ch a s S EM a nd m ic rorad iog raphy p roduce u nnece ssa ry d e ta i l . T he u se o f t e chn ique s mo re g ene ra l ly a cce s s ib le t oa r chaeo log is t s , s uch a s t r an sm i t ted a nd r ef le c ted l i gh t , i ss ugge s ted .

f i br i l la r a r rangemen t o f t h e c o l lagen m a tr ix b e come s p rog re s s ive ly mo re p a ra l le l . C emen tum a nnu l ia r eg ene ra l ly o rgan ised l i ke l ame l la r b one , i . e . , t h ey c on ta in f ewe r c e l ls t h an g row th z one s a nd t h e c o l lagen m a t r ix i s h igh ly o rgan ised ,c on ta in ing c l ose ly p acked p a ra l le l f i b re s , c au s ing i s o t ropy . LAGs a r ei nunda ted s u rf ace r eg ions r e su l t ing f r om p e r iod s o fg row th c e s sa t ion a nd p hys iochem ica l c hange o f t h e c emen tum s u rf ace , r a the r t h an t i s sue d epo s i t ion .

I n f o s s i l ised h o r se t e e th , t h e o rgan ic c omponen t o f c emen t h a s b e en r emoved . C on tra s t s i nt h e r e la t ive p e r cen tage o f o rgan ic a nd i n o rgan ic c on s t i tuen t s , w h ich a r e e nhanced b y d e ca lc if ica t ion a nd e t ch ing , a r e t h e refo re a l so l o s t . C onsequen t ly , S EM a nd h i s to log ica l p repa ra t ion s a r e n ega t ive ly a f f e c ted . I n t h e c a se o f h eavy m ine ra ls t a in ing d ue t od i agene t ic p roce s se s , o p t ica lc on t ra s t

C emen tum s t ruc tu re s i n h o r se t e e th c or re spond t o g row th s t ruc tu re s i n b one a nd c emen t d e scr ibed e l sewhe re . O b serva t ion s o f h o r se t e e th s uppor t t h ee x is tence o fd i f f e rence si nt i s sue c ompo s i t ion a nd m a tr ix o rgan isa t ion i n t h e d i f f e ren t t ype s o f c emen tum g row th l a ye r . G row th z one s i n h o r se c emen t c on ta in r e la t ive ly

b e twe en g row th l a ye r s i n f o s s i l t e e th m ay b e g rea t ly d im ished o r l o s t . T he d i f f e ren t g row th l a ye r s i n h e av i ly s t a ined t e e th a r e f r equen t ly n o t v i s ib le i n r ef le c ted l i gh t . T he r e la t ive u sef u lne s s o f t r an sm i t ted l i gh t a na ly s is f o r e xam in ing f o s s i l t e e th h i nge s o n t h e p o s s ib i l i ty o f u s ing c ompu te r -a ided d i g i ta l e nhancemen t t o i n c rea se o p t ica l c on t ra s t s .

m o r ec e l ls t h an a nnu l i , a nd a r eg ene ra l ly t h icke rt h an a nnu l i o r LAGs . When v i ewed u nde r t r an sm i t ted l i gh t g row th z one s a r e i s o t rop ic . LAGs ( a nd a nnu l i , when p re sen t ) a r e t h inne r , r e la t ive ly t r an sm i t ted l i gh t .

c e l l -poo r

a nd

a n iso t rop ic

7 .0 .3 P a t te rn o fD epo s i t ion o fC emen t i nE . c aba l lus .

u nde r T he p a t te rn o f d epo s i t ion o f e qu id c emen t i s d e s cr ibed , b a sed o n ac on t ro l o f 1 6 a n ima ls o fk nown d a te o f d ea th , v i ewed u nde r t r ansm i t ted l i gh t , a nd 1 2 a n ima l s ( de ad i n Ma rch ) c o l lec ted b y D . We lsh ( 1975 ) a nd v i ewed

I n t e r re s tr ia l s pec ie s , t h e re e x is t s a d i s t inc t ion b e tween p r ima ry a nd s e conda ry l a ye r ing i n c emen t . P r ima ry l a ye r s i n b one a nd c emen t a r e g ene ra l ly

u nde r r ef le c ted l i gh t . T he o paque g row th z one i n e qu id c emen t f o rm s f r om May t oO c tobe r . C emen tum d epo s i t ion s l ow s i n N ovembe r , w hen a n i n term i t ten t a nnu lu s m ay f o rm , a nd s t op s d ur ing t h ep er iod f r om D e cembe r t oA pr i l ,

d epos i ted i n ap red ic tab le , a nnua l c yc le . S e conda ry l a ye r s a r e l e s s p red ic tab le a nd r ef lec t av ar ie ty o f s ub -annua l e ven t s a f f ec t ing a no rgan ism 's d eve lopmen t . T he f o rma t ion

7 2

when a nnu l i a nd /or LAGs a r e g ene ra l ly o bservab le o n t h e

r e qu ire s t h e a s sump t ion t h a t as i ng le p opu la t ion i sb e ing

c emen tum s u rf ace . I n term i t ten t z one f o rma t ion m ay b eg in

s t ud ied . T h is a s sump t ion i sn o ta lways v a l id i na r chaeo logy .

a ga in i n Ma rch /Apr i l , n o t e xceed ing

1 0% o f e xpec ted

I n c ompar ison , c emen tum a na lys is d oes n o t r equ ire a ny

g row th ( b a sed o n t h ew id th o ft h ep receed ing g row th z one ) .

a s sump t ions r ega rd ing t h e s t udy p opu la t ion , a nd t h e refore i s b e t te r

s u i ted

t o t h e

a na lys is

a r chaeo log ica l a s semb lage s .

o f f o ss i l

t e e th

f r om

N ega t ive o r w eak p os i t ive

T he r a te o fd epo s i t ion o ft h eg row th z one s

c o r re la t ions b e tween c rown h e igh t , e rup t ion /wear s t age a nd

i nh o r se c emen t i sn o tl i nea r , b u tv ar ie sb e tween i n d iv idua ls

t h en umbe ro fp r imary g row th i n cremen ts w ere o b ta ined f o r

a nd d ur ing a n i nd iv idua l 's l i f e span . A n umbe r o f c au sa l

t h e

f a c tor s a r e p roposed ; f o r e xamp le , a ge' .

c or respondance b e tween t h e se t h ree m eans o f e s t ima t ing

T he r a te o f

p re sen t

c on t ro l

s amp le ,

i n d ica t ing

a

p oo r

a ge .

c emen t =g row th a l so v ar ie sw i th in t h es ame d en t i t ion a nd , t o al e sse r e x ten t , b e twe en s ucce s s ive p a ra l le l s e c t ion s o f t h e s ame t oo th , p robab ly a s ar e su l t o f b iome chan ica l

T he d i ff icu l t ie s i n he ren t i n i n terpre t ing

s t resse s . T he p resen t s amp le i st o o s ma l l t ot e s t p ropo sed

s e a sona l i n forma t ion i n as pe c if ic a r chaeo log ica l c on tex t

f a c tor s a f f ec t ing r a te o ft i ssue d epos i t ion . Al a rge r c on t ro l

u s ing t h e s ome t ime s c onf l ic t ing morphome tr ic d a ta a r e

s amp le

a ppa ren t i n t h e d i scu ss ion o f s ea sona l e v idence f r om a

s hou ld b e p repa red

r e la t ionsh ip b e tween a ge ,

i n o rde r

t o e s tab l ish t h e

s e lec t ion o f B adegou l ian a nd L ower Magda len ian s i te s i n

h ea l th , c l ima t ic f l uc tua t ion s ,

b iomechan ica l s t re sse s o n d en t i t ion a nd r a te o f c emen tum

S ou thwe s ter n F rance ( s ee C hap te r 6 ) . T he c on tr ibu t ion o f

d epos i t ion .

d en ta l c emen tum a na lys is t o ad i scuss ion o f s ea sona l i ty i s t h e refore a ppre c iab le .

7 .1 App l ica t ion o f a Modern C on tro l t o F os s i l Hor se 7 .3 R eg iona l a nd Chrono log ica l F ramework .

P opu la t ion s . P os s ib le d i f f e rence s b e tween modern a nd f o ss i l

p a t terns

o f

c emen tum

d epo s i t ion

c aused

T he g l ac ia l m ax imum , o r D rya s I ,( a bou t

b y

1 8 ,000 -14 ,000 B .P . ) is h igh ly r e cogn isab le s t ra t igraph ica l ly

e nv ironmen ta l ( i .e . , e xogenous ) i nf luence s , a r e a t tenua ted

i n S ou thwe s tern F rance a nd i s a n a i d t o e s tab l ish ing t h e

b y c o l lec t ing a l l s pec imen s f r om t h e s ame b road l a t i tud ina l

r e la t ive c on temporane i ty o fa r chaeo log ica l s i te s . A dd i t iona l

b and . A s im i la r o n togene t ic p a t tern o f p r imary c emen t

c h rono log ica l c on tro l i sp rov ided b y c u l tura l s ys tema t ics :

g row th c an b e a s sumed i nt h e c a se o fL a te P le is tocene a nd

c u l tura l

mode rn h o r se s .

Magda len ian a r e s t ud ied i nt h e p re sen t r e search .

T h is a s sump t ion i s s uppor ted b y

t h e

l e ve ls

d e s igna ted

a s

B adegou l ian

o r

L owe r

i d en t if ica t ion o f ab road a dap t ive t y pe f o r h o r se , E . f e ru s , T he S ou thwe s t o f F rance , o r A qu i ta ine

wh ich h a s n o t s pec ia ted s i nce t h e L a te P le is tocene . T he c ompa ra t ive ly r e cen t r a d ia t ion o f t r ue c aba l l ine s i n to

B a s in , i s ag eograph ica l ly d ef inab le r eg ion wh ich h a s b een

E ura s ia h a s r e su l ted i n t h e c rea t ion o f c on tempo ra ry ,

t h e s ub jec to fe x tens ive a r chaeo log ica l r e search . T he r e su l t

g eograph ica l ly

s ha re

i s aw ea l th o fa r chaeo log ica l d a ta , i n c lud ing e nv ironmen ta l

d eve lopmen t ,

a nd b iogeograph ica lr e con s truc t ions .R eg iona lc l ima to log ica l

n u tr i t ion a nd s oc ia l b ehav iour . T h is i mp l ie s ah igh d eg ree

r e cons truc t ions f o r t h e g l ac ia l m ax imum ( s ee C hap te r 4 )

s im i lar i t ie s

s epa ra ted

p hys ica l

i n p hys io logy ,

t y pe s ,

r eproduc t ion ,

wh ich

o f g ene t ic u n iform i ty a mong m embe r s o f t h e c aba l l ine

r e vea l

s pec ie s .

w in te r s a nd r a p id s p r ing t h aw s . T he re i sn o e v idence o f

m arked

s e a sona l i ty , w i th w arm s u mme r s ,

c o ld

e x ten s ive p ermaf ro s t z one s i nt h e r eg ion , e xcep tn ea ra l p ine 7 .2 Morpho log ica l V er su s S ke le tochrono log ica l

i c e s hee ts . S t r ic t ly s peak ing , t h e refore , t h e Aqu i ta ine i sn o t

Ana lyse s .

ap er ig lac ia l e nv ironmen t d ur ing D rya s I . R a the r , t h e r e g iona ll a nd scape i sc ha rac ter ised b yp redom inan t ly s t epp ic T he re e x is tv ar iou sm orpho log ica lm e thods

f e a tu re s , w i th av a r ie ty o fm icroc l i ma te s i nf luenced b y l o ca l

f o r e s t ima t ing a ge a td ea th a nd s e a son o fd ea th . T hey h ave

t o pog raphy a nd p rox im i ty t op er ig lac ia l e nv ironmen ts i nt h e

t h e a dvan tage o f b e ing n on -de s truc t ive . T he a na lys is o f

n or th , a nd A lp ine c ond i t ion s t ot h e e as t a nd s ou th .

d en ta l

w ea r /erup t ion

p a t terns ,

c rown

h e igh t ,

a n t le r T he A qu i ta ine b a s in d ur ing t h e g l ac ia l

morpho logy a nd e p iphysea l f u s ion p a t ter n s c an p rov ide e v idence o f i n d iv idua l a ge . T h is e v idence c an b e u sed , i n

m ax imum c an b e d i v ided i n to t h ree m a jo r b iogeog raph ic

c on junc t ion w i th k nown s e a sona l p opu la t ion s t ruc ture s f o r

z one s r ough ly c o inc id ing w i th 1 ) t h e d epar temen t o f t h e

a g iven s pe c ie s ,

i n dex o f s e a sona l i ty .

D ordogne , 2 ) t h e d epar temen ts o ft h e G ironde a nd L ande s ,

D iff icu l t ie s i n heren t i n t h e u se o f t h e se morphome tr ic

3 ) t h e r e g ions b o rder ing t h e P yr6ne ;e s a nd t h e Ma s s if

t o p rov ide a n

t e chn ique sf o rs t udy ing h or se i n c lude : a lome tr ic d i f f e rence s

C en tra l . T he f i r s t z one , i . e . , t h e D ordogne , i s a mosa ic

i nt oo th h e igh t b e tween a nd w i th in h o r se p opu la t ions ; i n ter

e nv ironmen t w i th n ume rous t r ans i t ions f r om i n terf luv ia l t o

a nd i n tra -popu la t iona l d i f f e rence s i n t oo th w ea r p a t ter n s ,

r i ve r v a l ley e co tone s , c on ta in ing a c o ld s t eppe -adap ted

a nd t h e f a c t t h a t a l l o cc lusa l s u rf ace s a r e r epor ted t ow ea r

f a una d om ina ted b y r e indee r ( R . t a randus ) . T he s e cond ,

f l a t b y 6y ea r s o f a ge . T he a ge a t w h ich e r up t ion i s

c oa s ta l z one i s g ene ra l ly more o pen a nd f l a t te r t e r ra in ,

c omp le te l im i ts a pp l ica t ion o f e rup t ion t a b le s t o4 -6 y ea r s

c ha rac ter ised

f o r h o rse . F ur thermore , a l though t o o th h e igh t a nd w ea r p a t tern s c an b e u sed t oe s tab l ish r e la t ive a ge c l a s se s , t h is

d om ina ted b y S a iga ( S . t a ta r ica ) i nt h e n or th a nd i n c lud ing b i son ( B . p r iscu s ) , h or se ( E . f e rus ) a nd r e inde er . I n t h e

b y

a c o ld ,

d ry

s t eppe -adap ted

f a una ,

t h ird z one - t h e more e l eva ted s ou thern a nd e a s tern m a rg ins 7 3

o f t h e A qu i ta ine - mon tane e l emen t s , e . g . , c hamo is r up icap ra ) a nd i b ex ( L.i b ex ) , a r e c ha rac ter is t ic o ft h e l o ca l

l ong n or th / sou th p a t ter n o f m ig ra t ion , b e twe en P y renean s umme r r a nge s a nd t h e D o rdogne r e g ion , i sc on s ide red b y G o rdon ( 1988 ; a nd s e e D e lpe ch 1 987 ) .

f a una . 7 .4 .0 E tho logy o f Ma jor _ Pm i S pec ie s .

T he mode l p ropo sed i n F igu r e 7 . 1 ( a dap ted f r om B u rch 1 972 ) i sc ons is ten tw i th s e a sona l d a ta s ugge s t ing t h a t w in te r r a nge s w e re a c ce s s ib le t oh uman s i n t h e G ironde ( hypo the t ica l ly a tt h e p e r iphery o f t h e w in te r r ange ) a s w e l l a s t h e D ordogne ( t he c en te r o f t h e w in te r r ange ) . I ti si n te re s t ing t on o te t h a t o t he rw ise r a re s umme r k i l ls o fr e indee r a r e r epor ted f o r s i te s i nt h e G ironde s u ch a sS t -Germa in la r iv ire ,F ongaban , V idon a nd Mongaud ie r

B iogeograph ica l a nd c l i mat ic r e con s t ruc t ion sg u ide t h e a pp l ica t ion o f mode rn e t ho log ica l p rof i le s t o t h e s t udy o f f u l l g l ac ia l f auna i nS ou thwe s te rn F rance . R e indee r ( 1. t a randu s ) a nd h o r se ( L . f e ru s ) , c on s t i tu te t wo o ft h e m o s t i mpo r tan t r eg iona l g l ac ia l f a una f o r S ou thwe s ter n F r ance . T he i mpor tance o f o the r p rey s pe c ie s ( e .g . , S . t a ta r ica i nt h e G ironde ) i sa cknow ledged , b u t t h e i r s t udy e xceed s t h e s cope o ft h is r e sea r ch .

( i n G o rdon 1 988 ) -w he re s u mme r a nd w in te r r a nge s p o ten t ia l ly o ve r lap ( s ee F igu re 7 . 1) . T he a bundance o fo the r g regar iou s p rey s pe c ie s i n t h e G i ronde d ur ing s umme r

I n t h e c a se o fr e inde e r , s pr ing m ig ra t ion

mon th s , n o tab ly s a iga , m ay e xp la in w hy a r chaeo log ica l a s semb lage s f r om z one 2 a r e n o t h eav i ly d om ina ted b y r e indee r .

i s t r igge red b y t h e r e qu iremen t s o f g r av id f ema le s f o r i s o la ted , b a r ren t e r ra in t o c a lve i n ; s now c ove r a nd a va i lab i l i ty o f s u i tab le f odde r g ene ra l ly t r igge r f a l l m igr a t ion . S ea sona l r e indee r movemen t s d ur ing t h e f u l l g l a c ia l a r e p red ic ted . T he se m ig ra t ion s w i l l h ave b een

A s a l ready s t a ted , G o rdon 's ( 1988 ) c emen tum s t udy o f r e indee r t e e th i n d ica te s ap a t ter n o f

r e la t ive ly s ho r t i nt h e D o rdogne , d ue t o t h e p rox im i ty o f t opograph ica l ly a nd f l or is t ica l ly v a r ied e nv ironmen t s ( i .e . , p o ten t ia lc a lv ing g round s ,w in te ra nd s umme rf o rage a r ea s ) . H e rd s i n t h e D o rdogne m ay h ave b een s ma l le r t ho se

w in te r / spr ing r e indee r e xp lo i ta t ion b y h uman s i n t h e D ordogne . H is d a ta s ugge s t a s im i la r p a t te rn i n t h e G ironde . A l though Go rdon u se s t h e se r e su l t s t os uppo r ta n h ypo the s is o f l o ng , n or th / sou th m ig ra t ion s , t h ey a r e n o t i n con s is ten t ap a t te rn o f e a s t -we s t m ig ra t ion e i the r , i fo ne c on s ide r s am ode l r e indee r h ab i ta t s uch a s p ropo sed i n F igu re 7 . 1 .

o b se rved i nm o re o pen t u ndra o rb ar ren land e nv ironmen t s , w he re s pe c ie s d i ve r s i ty i s l ow , a nd t h e ir m igra tory movemen t s m ay t o pog raph ica l ly .

h ave

b een

l e ss

" channe led"

Unfo r tuna te ly , t h e d a ta a va i lab le a r e i n suff ic ien t f o r t h ep u rpo se o ft e s t ing r a nge r e con s t ruc t ion s

M idd le a l t i tude r eg ion s i nt h e Do rdogne p robab ly c on s t i tu ted t h e " co re " a r ea , o r k ey h ab i ta t , f o r r e inde e rd ur ing t h eg l a c ia l p er iod' . N e ighbour ing r eg ion s , s u ch a st h e f oo th i l ls o ft h e Ma s s if C en t ra l a nd t h e G ironde ,

f o r t h e Ma s s if C en t ra l a nd P y renean r e g ion s . Howeve r , s e a sona l e v idence f r om R ond d u B a r ry d oe s n o ts ugge s tt h e e x is tence o fs umme rr ange s f o rr e inde e rn ea rby . T he f a una l

w ou ld h ave b e en p e r iphe ra l t o t h e c o re r a nge . R e indee r h e rd movemen t si nz one 2m ay h ave b e en l o nge rt h an i nt h e D o rdogne , d ue t o t h e l o nge r d i s tance s e x is t ing b e tween s e a sona l ly e xp lo i ted e cozone s . I n f a c t , r e la t ive ly l ong s e a sona l movemen t s c an b e p red ic ted f o r a l l o ft h e mob i le , l a rge -bod ied h e rb ivo re s i n hab i t ing t h e w e s tern p or t ion o f t h e A qu i ta ine d ur ing D rya s I .

e v idence f r om D uru thy , o n t h e o t he rh and , s ugge s t sw in te r p re sence o f r e indee r c on s is ten t w i th s hor t , a l t i tud ina l m ig ra t ion s r a the r t h an a l o ng n or th / sou th p a t ter n o f movemen t s u ch a s s ugge s ted b y G o rdon ( 1988 ) . S ince a l l o f t h e a bove r ange r e con s truc t ion s c l a im t o b e s uppo r ted b y e x is t ing t opog raph ica l a nd s ea sona l e v idence , i ti so bv iou s t h a t w e d o n o t h ave e nough d a ta f o r t h e p u rpose o f t e s t ing

T he r e la t ive ly s hor t s e a sona l movemen t s o f h o r se a r e p r ima r i ly t r igge red b y s now c ove r a nd t h e a va i lab i l i ty o f f odde r . I n w in te r , h o r se s f r equen t ly s e ek s he l te r i nv a l ley b o t tom s . D ur ing t h e g l a c ia l p er iod i n t h e D o rdogne s m a l l h e rd s ( a bou t 4 10 h o r se s ) p robab ly f o l lowed s ho r t , a l t i tud ina ls e a sona l movemen t s . Ho r se m ay

h ypo the t ica l m ig ra t ion p a t te rn s y e t . T h is i s p robab ly a r ef le c t ion o f a more g ene ra l p rob lem i n he ren t i n t h e c on s truc t ion o fp a laeon to log ica l mode l sf r om a r chaeo log ica l a s semb lage s .

h ave f o l lowed r e la t ive ly l o nge rm igra t ion s i nt h e o pen , l e s s d i ve r s if ied e co sy s tem o ft h e G ironde .

O ne c onc lu s ion c an b e d rawn f r om t h e d a ta a t h and , h oweve r . R ega rd le s s o f t h e l o ca t ion o f s umme r r a nge s a nd c a lv ing g round s , t h e D o rdogne i sa l i ke ly w in te rh ab i ta t f o r b o th r e inde e r a nd h o r se , p rov id ing

7 .4 .1 S ea sona l D a ta f o r R e indee r : R ange R e cons t ruc t ion s .

v a l ley s he l te r a s w e l l a s e s cape f r om u nf avou r ab le s now c ond i t ions . T he a va i lab le s e a sona l d a ta s uppor t t h i s c onc lu s ion .

A ne a s t /we s tp a t tern o fr e inde e rm ig ra t ion d ur ing D rya s I , f r om t h e G ironde t o t h e D o rdogne , i s p roposed b y an umb e r o f a u tho r s ( e .g . , L a cor re 1 953 ; B ouchud 1 966 ; D av id 1 973 ; S tu rdy 1 975 ; D e lpe ch 1 983 , 1 987 ) . S im i lardy , ap a t te rn o f movemen t f r om h igh t ol ow a l t i tude r a nge s n e a r t h e Ma s s if C en t ra l a nd P yr& i6e s i s

7 .4 .2 S ea sona l D a ta f o r H o r se . When c emen tum a na lys i s i s d one o n a r chaeo log ica l s amp le so fh o r se t e e th f r om t h ir te en r ough ly c on tempo ra ry B adegou l ian a nd L owe r Magda len ian s i te s i n

p ropo sed ( e .g . , F reem an 1 974 ; D e lpech 1 987 ) . Ar e la t ive ly 7 4

F igure 7 . 1 P ropo sed R econ s truc t ion o fR e indeer R ange s i nS ou thwe s tern F rance Dur ing t he P len ig lac ia l .

T he e x tremes o ft he w in ter a nd s u mmer r ange s a re u nknown . Over lap b e tween w inter a nd s u m mer r anges c orre sponds t o t he C ore A rea o ft he r e indeer h ab itat .

7 5

h uman h un t ing s t ra teg ie s d ur ing t h e U ppe r P a laeo l i th ic . T he se s h if t sp robab ly a ccompany s ub s is tence s h if t sb a sed o n

S ou thwe s te rn F rance , t h e r e su l t s g ene ra l ly s how ap a t te rn o f h o r se e xp lo i ta t ion t h a t i sn o t s ea sona l ly m a rked , a nd o ccu r s p o ten t ia l ly t h roughou t t h e e n t ire y ea r . T he se r e su l t s

t h ea va i lab i l i ty o fm ig ra to ry ( i .e . ,c l umped )g ame . I n te r cep t h un t ing o fr e indee r a l te rna te s w i th e ncoun te r -ba s is h un t ing o f d i spe r sed p rey ( s uch a s h o r se a nd s e a sona l ly d i spe r sed r e indee r ) . T h i s p a t te rn i sv a l id i na l l o f t h e s ub reg ion s o f t h e A qu i ta ine B a s in s t ud ied .

a r e c on s is ten t w i th t h e e xpec ta t ion o f s hor t , p o s s ib ly a l t i tud ina lm ig ra t ion f o rh o r se , g ene ra ted b y t h ee t ho log ica l d a ta p re sen ted a bove . 7 .4 .3 E tho logy a s aP red ic tor o f Human B ehav iou r .

I n o rde r t og a in ad eepe r a pp rec ia t ion o f h ow h uman s e t t lemen ta nd s ub s is tence s y s tem sa r er e f le c ted i nt h ef a una l s amp le , h oweve r , i ti sn e ce s sa ry t oe xpand t h e s ke le toch rono log ica l s t udy o f s e a sona l i ty t o i n c lude o the r p rey s pe c ie s s u ch a ss a iga a nd b ov ine si nt h e G ironde ( z one

I t i s p o s s ib le t o g ene ra te p red ic t ion s c once rn ing h uman s e t t lemen t p a t te rn s f r om t h e e t ho log ica l i n f orm a t ion d i scu ssed a bove . 1 )T he o p t ima l p l acemen t o f s e a sona l s i te so rh un t ing s t and s i nt r an s i t iona le cozone s a nd a l ong l i ke ly m ig ra t ion p a th s , e . g . , w a te r cou r se s . 2 ) Win te r

2 ) , a nd c hamo is a nd i b ex i nt h e f oo th i l ls o f t h e P y r6n6e s a nd Ma s s if C en t ra l ( z one 3 ) . A s a l re ady s t a ted , t h i s i s b eyond t h e s c ope o ft h e p re sen t r e sea r ch .

o ccupa t ion o fv a l ley l o ca t ions i nt he Do rdogne . Ac o ro l la ry p red ic t ion i sf o rw in te ro ccupa t ion o fl ow land l o ca t ions n ea r t h e Ma s s if C en t ra l a nd P y ren & s . Wh i te ( 1985 ) p o in t s o u t t h e n umbe r o f s i te s c on sp icuou s ly l o ca ted n ea r p o ten t ia l r e indee r h e rd c ro s s ing s a l ong t h e We e re d ur ing t h e e a r ly Uppe r P a laeo l i th ic . B oy le 's ( 1990 ) a na ly s is o f r e inde e r b one f r agmen ta t ion i nt h es amp le r eg ion i sa l so c on s is ten tw i th a n

7 .6 .0 S ea sona l I nd ica tor s a s E v idence o f S ub s is tence a nd S e t t lemen t S y s tem s I n t e rm s o f e a r ly U ppe r P a laeo l i th ic s e t t lemen t s y s tem s , t wo i n te rp re ta t ion s o f t h e c omb ined s e a sona l d a ta f o r h o r se a nd r e inde e r a r e p o s s ib le . 1 )S i te s i n t h e s amp le r e g ion a r e u sed y ea r r ound a nd t h e

h ypo the s is o fh uman i n te rcep t ion o f m igra to ry r e indee r i n v a l ley b o t tom l o ca t ion si nt h e Nr igo rd . T he se d a ta s uppo r t

s ub s is tence b a se s h if t s s e a sona l ly . 2 ) S i te s s amp led a r e

t h e p red ic ted o p t ima l p l a cemen t o f s i te s a l ong m ig ra t ion r ou te s , w he the rb e tween i n te rf luv ia l /f luv ia le cozone sw i th in

p r imar i ly o ccup ied w hen r e inde e r a r e l o ca l ly a va i lab le i n l a rge c oncen t ra t ion s . H o r se s upp lemen t sd w ind l ing r e indee r n umbe r s j u s t b ef o re a nd p e rhap s j u s t a f t e r s e a sona l m ig ra t ion o ft h eh e rd s , p r io r t o ar e s iden t ia l s h if t . I ne i the r

t h e Do rdogne , o r b e tween p o ten t ia l w in te r a nd s umme r r ange s i n t h e G ironde a nd t h e Do rdogn e . T he s e cond p red ic t ion ( w in te r o ccupa t ion o ft h e Do rdogne ) i sv er if ied b y f a una l e v idence p re sen ted a bove . T he c o ro l la ry , w in te r

c a se , a t a l l t we lve o f t h e s i te s s amp led h e re , h or se i sa " comp lemen ta ry " r e sou r ce -d im in ish ing t h es e a sona l i mpac t o fr e inde e ra va i lab i l i ty o n l o ca l h uman e conom ie s . A mode l o fy e a r r ound h uman e conom ic d ependence o n r e inde e r i s t h e refo re n o t s uppor ted b y t h e se r e su l t s . T he i mp l ica t ion s o ft h i s c onc lu s ion f o r h e rd f o l low ing m ode l s a r e d i s cu s sed b e low .

o ccupa t ion o f l ow land s i te s , i s v er if ied a t Du ru thy ( Py renee s ) a nd R ond d u B a r ry ( Ma s s if C en t ra l ) . 7 .5 T he A rchaeo log ica l S amp le a s aR e f lec t ion o f Human B ehav iour . F rom a n a r chaeo log ica l p e r spec t ive , t h e a bsence o fs umme r r e indee r k i l ls i nt h e D o rdogne ( z one 1 )

7 .6 .1 G enera l T heory a nd A rchaeo log ica l Mode ls o f S e t t lemen t a nd S ub s is tence .

c ou ld b e ar e su l t o f t wo t h ings : 1 ) as h if t i n t h e h uman s ub s is tence s y s tem d ur ing t h e s u mme r s e a son mak ing t h e l a t te r a r chaeo log ica l ly i n v is ib le i fo ne r e l ie s o n as i ng le s pe c ie s a sa n i n dex o fs e a sona l i ty , 2 ) as amp le b i a s a ga in s t

Mo s t s e t t lemen t a nd s ub s is tence m ode ls p ropo sed f o r t h e S ou thwe s t o f F rance d ur ing t h e l a s t f u l l

ap or t ion o f al a rge r , s e a sona l ly a nd p e rhap s g eograph ica l ly

g l ac ia l m ake b a s ic a s sump t ion sa bou th un te r -ga the re rs o c io -

d i f f e ren t ia ted s e t t lemen t s y s tem . T he p re sen t r e sea rch a t temp t s t oa dd re s s t h e se p os s ib i l i t ie sb y t e s t ing t h e s ea son

t e r r i tor ia l o rgan isa t ion a nd s ub s is tence . T he se a s sump t ion s a r e d rawn f r om g ene ra l m ode ls n o tp roven t ob ea dap ted t o

o fs i te o ccupa t ions r eg iona l ly , u s ing mo re t h an o ne s pe c ie s a s a n i n dex o fs e a sona l i ty .

t h e s t udy o fP a laeo l i th ic s y s tem s , e . g . , t h ec ompo s i te b and . I n t h is m ode l , an umbe r o f s m a l l , s p a t ia l ly d i spe r sed , e xogamou s l o ca l b and s t a ke a dvan tage o f s e a sona l ly a gg rega ted r e sou rce s t o f o rm e ndogamou s , c o re s iden t ia l g roup ing s ( i .e . , r e g iona l b and s ) s uppor ted b y k insh ip a nd e xchange . T he c ompo s i te b and m ode l i mp l ie s as e a sona l p a t te rn o f f i s s ion a nd f u s ion w h ich h a s n o t b e en

T he d a ta c o l lec ted h e re a r gue f o r t h e e x ten s ion o fh uman s i te -u se i nt h e A qu i ta ine B a s in b eyond s i ng le sea son o ccupa t ion s . T he s ma l l n umbe r o f a n ima ls ( ba sed o n MN I ) a va i lab le a t e ach s i te s ugge s t s t h e p o s s ib i l i ty o fs po rad ic , s hor t te rm e p isode s o fs i te -u se o n a

c onv inc ing ly i d en t if ied i nS ou thwe s te rn F rance ( s ee C hap te r 6 ) . N eve r the le s s , " un ive r sa l" c oncep t s d e r ived f r om b and

y e a r r ound b a s is . F ur thermo re , t h e e v idence p o in t s t o t h e e x is tence o f s e a sona l s ub s is tence s h if t s . T he se s h if t s a r e

t h eo ry , s u ch a st h e i n he ren t s oc ia l a nd t e rr i to r ia l f l ex ib i l i ty

l i ke ly t ob e l i nked t os h if t s i nh un t ing s t ra tegy .

o f h un te r -ga the re r s y s tem s a nd t h e ir p o ten t ia l m ob i l i ty , f o rm b a s ic c omponen t s o fm any a r chaeo log ica l mode l s f o r t h e F rench U ppe r P a laeo l i th ic ,

S ke le toch rono log ica le v idence f r om h o r se a nd r e indee r t e e th , c omb ined w i th B oy le 's ( 1990 ) a na ly s is o f b one f r agmen ta t ion p a t tern s , s ugge s t s e a sona l s h if t s i n 7 6

O p t ima l F o rag ing T heory i sa no the r a r ea

mob i l i ty t h an t h a t p roposed b y t h e h e rd f o l low ing mode l .

o f g ene ra l t h eory f r om wh ich k ey c oncep t s f o r t h e i n ve s t iga t ion o f P a laeo l i th ic s e t t lemen t s y s tem s h ave b een b e

I n s umma ry , t h e refore , s ea sona l d a ta f o r t h eB adegou l ian a nd L owe r Magda len ian r e su l t ing f r om t h e

o pe ra t iona l ise c l , g i ven t h e c ur ren t s t a te o fk now ledge a bou t

p re sen t r e sea rch d o n o t s uppor t amode l o fh e rd f o l low ing

t h eP a laeo l i th ic e nv ironmen t . Howeve r , t h ey h ave p rov ided

f o r t h e S ou thwe s t o fF rance d ur ing D rya s I -n o r d o f a una l

d r awn .

T he

mode ls

d er ived

f r om

i t

c anno t

a r chaeo log is t s w i th am eans o fe va lua t ing t h e r e la t ive c os t s

d a ta f r om t h e G ironde a nd Ma s s if C en tra l .

o f g i ven r e sou rce s' , a nd t h e refore a m ean s o f c ompara t ive ly e va lua t ing h um an d ec is ions c oncern ing

h ypo the s is

s ubs is tence . T he n o t ion o f e conom ic r a t iona l i ty , i . e . , o p t im isa t ion , a s i n t roduced b y O p t ima l F orag ing a nd o the r ,

r e s iden t ia l mob i l i ty , a ccompan ied b y s ea sona l s h if t s i n

s im i la r

e ne rge t ic s

mode ls ,

i s f r equen t ly

i mp l ic i t

t o r e indee r

g eog raph ica l ly

a nd

h e rd

A n a l ter na t ive

f o l low ing i s o f a more

s e a sona l ly

r e s tr ic ted

p a t ter n

o f

r e source u se .

i n

a r chaeo log ica l mode ls .

7 .7 .2 T he S ea sona l R ounds Mode l .

7 .7 .0 Mode ls o f S ub s is tence a nd S e t t lemen t i n t he

p ropo sed f o r t h e e a r ly Uppe r P a laeo l i th ic i nt h e S ou thwe s t

V ar iou s

" s ea sona l r ound s" mode ls a r e

o fF rance . A l l o f t h e mode ls d i scu ssed ( s ee C hap te r 6 ) a r e

F rench Upper P a laeo l ith ic .

b a sed o n a n a s sump t ion o f s e a sona l r e s idence s h if t s . I n o t he r w ords , t h e mode ls p red ic t m a rked s e a sona l c hange s

7 . 7 .1 H erd F o l low ing Mode ls .

i n s i te u se p a t terns . Howeve r , s e a sona l e v idence f r om S ome e t hnograph ic e v idence s uppo r t sh e rd f o l low ing a s a v i ab le e conom ic s t ra tegy f o r h uman s .

s ke le toch rono logy , p re sen ted h e re , d oe s n o t s uppor t t h e p red ic t ion o f s e a sona l ly m a rked s i te -u se . O n t h e c on t ra ry ,

H oweve r , t h ea pp l ica t ion o ft h eh e rd f o l low ing mode l t ot h e

t h e re i se v idence o fc on t inued s i te -use t h rough t wo o r more s e a sona l c hange s .

F rench e a r ly Uppe r P a laeo l i th ic r e qu ire s an umbe r o f a s sump t ion s .

T he f i r s t ma jor a s sump t ion i s o f h uman A no the r p a t ter n c an b e p roposed , more

e conom ic r e l iance o n r e indee r . T he s e cond i so fm igra tory b ehav iour o n

t h e

p ar t

o f r e indee r

h e rds .

T he

g eograph ica l ly r e s tr ic ted t h an t h eh e rd -fo l low ing mode l a nd

t h ird

mo re c omp lex t h an a s imp le s e a sona l r ound mode l . F o r

a s sump t ion i so fh uman r e s iden t ia l mob i l i ty .

e xamp le , C lo t te s ( 1987 :305 , 3 25 ) s t a te s t h a t e v idence o f e conom ic

s pec ia l ised m igra t ion h un t ing ( imp ly ing h e rd f o l low ing ) amd

r e l iance o n r e indee r i sb a sed o n a" c la ss ic" s i te t y pe f o r

e v idence o f" s ea sona l r ound s " b o th e x is ti nt h eP yrenee s . I n

S ou thwes ter nF rance , i . e . , t he Nr igo rd c ave o r s he l te r s i te

o the r words t h e t wo s t ra teg ie s a r e n o t mu tua l ly e xc lu s ive

w i th

b u t r a the r , r epresen t c omp lemen tary s e a sona l s t ra teg ie s . I

T he

a s sump t ion

a r e indee r -dom ina ted

o f

f a una l

h uman

a s semb lage .

T h is

w ou ld a rgue t h a t t h is i sa l so t h e s i tua t ion i nt h e A qu i ta ine B a s in d ue ing t h ee a r ly Uppe rP a laeo l i th ic .T he e v idence j us t

a s sump t ion m ay o n ly b e s ea sona l ly v a l id , h oweve r . S i te s i n o t he ra r ea s o ft h eA qu i ta ine a r ed om ina ted b y o the rs pec ie s , e . g . , R ond d u B a r ry ( i n t he f oo th i l ls o ft h e Ma s s if C en tra l ) a nd R oc d e Ma rcamps ( i n t h e G ironde ) . T he i n e scapab le

p re sen ted s ugge s t s ap a t ter n o fs e a sona l ly m a rked s h if t s i n

i mp l ica t ion i s t h a t e cono m ic s pe c ia l isa t ion ( i .e . , h e rd f o l low ing ) i sn o t ar e g iona l ly v a l id mode l f o r t h e A qu i ta ine

t h an s easona l ly marked r e s iden t ia l s h if t s .

B a s in .

7 . 7 .3 C onc lu s ions .

h un t ing s t ra tegy , f r om e ncoun te rt oi n te rcep th un t ing ,r a ther

S ea sona l d a ta o b ta ined f r om c emen tum

S ea sona l d a ta f o r B adegou l ian a nd L owe r

a na lys is a r e u sed h e re a s at e s t o f t h e s e cond a nd t h ird a s sump t ion s r e ga rd ing h uman mob i l i ty . T he l i ke l ihood o f

Magda len ian s i te sr e su l t ing f r om t h ep re sen tr e sea rch d on o t

m ig ra tory b ehav iou r o n t h e p ar t o f r e indee r d ur ing D rya s

s uppor t as imp le , s e a sona l r ounds mode l . I n s tead , t h is

Ih a s b een e s tab l ished , e ven i ft h e e xac t m igra t ion p a t ter n s

r e sea rch h a s p re sen ted e v idence o f e x tended , o r mu l t i -

h ave n o t b een i d en t if ied . I f h uman s w e re d ependan t o n

s e a sona l s i te u se f o r t h e A qu i ta ine B a s in d ur ing t h e U ppe r

r e inde e r d ur ing t h is p er iod , t h ey w i l l h ave h ad t oa dop t a

P a leo l i th ic , f r om 1 8 ,000 t o 1 4 ,000 B .P . . T he c onc lu s ion

s im i la rp a t ter no fs e a sona l movemen t . T he refore , t h er e cord s hou ld s how c l ea r d i f f e ren t ia t ion b e tween w in te r a nd

t oa c comoda te s e a sona l s h if t si nt h e s ubs is tence b ase , r a the r

s umme r o ccupa t ion s , a s w e l l a s w i th s pr ing a nd f a l l s i te s .

t h an a dop t amo re mob i le p a t ter n o f r e s idence .

s uppor t t h e h ypo the s is o f h e rd f o l low ing . N or d o t h ey

r e ached h e re i st h a th umans u sed d i f f e ren th un t ing s t ra teg ie s

T he s ea sona l d a ta f o r b o th h or se a nd

T he s oc ia l c or re la te s o f t h e se s e a sona l

r e inde e r d o n o ts uppor t ap a t tern o fr e s iden t ia l mob i l i ty t i ed

s h if t s i nh un t ing s t ra tegy a r e s ub t le . I n te rcep t s t ra teg ie s a r e

t o

T he

n o t , o f t h em se lve s , e v idence o f s oc ia l i n terac t ion o n t h e

e x tended

s c a le o f t h e c ompos i te , r e g iona l b and a s d ef ined b y L e e &

a

s i ng le

r e source

s ke le tochrono log ica l

d a ta

f o r

( i .e ., h or se

r e indeer ) . i n d ica te

s e a sona l r e s idence a tt h e s i te s s amp led , a l though m any o f

D evo re

t h e f a una l a s semb lage s a r e d om ina ted b y r e indee r r ema in s .

m igra t ion p a ths c an b e v ar iab le , o ne o ft h e r e qu iremen t s o f

T he p a t ter no fh o r se mor ta l i ty c an b e u sed t os uppo r t e i the r

i n te rcep t h un t ing i sa ccura te i n forma t ion d ur ing t h e h e rd

y ea r r ound s i te -use , o r a more l i m i ted p a t tern o f s e a sona l

m on i tor ing

7 7

( 1968 )

a nd

s t age s .

D ama s

T h is

( 1969 ) .

m eans

t h a t

H oweve r ,

l o ca l

s i nce

b ands

i n

I ft h e c ompos i te b and mode l h a sn o tb e en v a l ida ted f o r t h e s t udy r eg ion , t h is i sp ar t ly d ue t o t h e l oo se ly d ef ined a s so c ia t ion b e tween c ommuna l a nd

S ou thwe s te rn F rance w e re p robab ly i n te rac t ing o n a r eg iona lb a s is . T h is i n te rac t ion w i l lh ave r e su l ted i nc u l tu ra l l i nk s a nd s im i la r i t ie s r eg iona l ly , a s p red ic ted b y t h e c ompo s i te b and mode l , w i thou t r equ ir ing t h e l o ng -d i s tance

i nd iv idua l h un t ing t a c t ic s a nd s oc ia l o rgan isa t ion . E thnograph ic mode ls o f h un te r -ga the re r s ys tem s ( s uch a s t h e c ompo s i te b and mode l ) mu s t c on t inue t ob e p u t t ot h e

p opu la t ion movemen t s p red ic ted b y t h e h e rd -fo l low ing mode l .

t e s t . T he " t e s t s " u sed b y a r chaeo log is t s mu s tb em ade mo re e xp l ic i t t h an h e re tofo re , i fp roof t h a t t h e se mode ls a r e a pp l icab le t ot h e P a laeo l i th ic i st ob e f ound .

7 .8 .0 S ugge s t ion s f or F ur ther R e search . 7 .8 .1 T heo re t ica l R e sea r ch .

7 .8 .2 F u tu re R e sea r ch i nC emen tum A na lys is .

T he d i f f e ren th un t ing s t ra teg ie s a luded t o a bove , n ame ly i n te r cep t a nd e ncoun te r b a sed h un t ing , a r e

T he

u sua l ly c ha rac ter ised a sc ommuna la nd i n d iv idua ls t ra teg ie s , r e spe c t ive ly . T he n umbe r o f a n ima ls i n te r cep ted a nd p roce ss ed a t ag iven s i te i sa n i nd ica t ion o f t h e n umbe r o f p eop le p re sen to n ly w hen f i ne s t ra t ig raph ic c on t ro l p e rm i t s

p re sen t

s t udy

h a s

e x tended

t h e

a pp l ica t ion o f c emen tum a na lys is t o an ew s pec ie s , E . c aba l lus . T h is w a s d one t h rough c a re fu l c o l lec t ion o f s pe c imen s o fk nown p roven ience a nd k nown d a te o fd ea th . T he s pe c imen sw e re e xam ined u s ing av ar ie ty o fe s tab l ished t e chn ique s , t o d e term ine 1 ) t h e n a tu re o f t h e g row th

u st od i s t ingu ish s epe ra te h un t ing /p roce s s ing e p i sode s . F ew B adegou l ian a nd L owe r Magda len ian c ave s i te s c an c l a im s uch f i ne -g ra ined c on t ro l o ve r d i scre te h un t ing a nd

s t ruc ture s o b served a nd 2 ) t h e b e s t m eans o f 7 ob ser v ing t h em . T he r e su l t s h ave b een l a rge ly p o s i t ive .

p roce s s ing e p isode s" . We a r e l e f t w i th a n i n tu i t ive i n te rpre ta t ion o f r e la t ive r ep re sen ta t ion s o f m ig ra to ry s pe c ie si nt o ta la s semb lage s wh ich c ou ld r ep re sen t an umbe r o f d i sc r e te h un t ing e ven t s , o r e ven a n umbe r o f d i sc re te s ea sona le ven t s . T he p rob lem w i th i n te rp re t ing s uch r e la t ive r ep re sen ta t ion f i gu re s ( e .g . , N ISP s , MN I s ) i st h a t d i f f e ren t s ea sona l h un t ing s t ra teg ie s , y i e ld ing d if er en tp a t tern ing i n t he f auna l a s semb lage s , a r e e mp loyed f o r d i f f e ren t p rey ,

F u r the r r e sear ch i n to t h e p o s s ib i l i ty o f e s tab l ish ing t h e r a te o f d epos i t ion o f g row th l a ye r s a s a f unc t ion o fa ge r equ ire s t h ee xpan s ion o ft h ep re sen tc on t ro l s amp le . T h is r a ise s t h e p os s ib i l i ty o f s ub -d iv id ing t h e s i x mon th p e r iod d ur ing wh ich g row th z one s a r e d epo s i ted . F ur thermo re , t h e p o ss ib i l i ty o f e s tab l ish ing m a rg in s f o r

i . e . , t h e v a lue s o b ta ined a r e n o t c ompa rab le .

e r ro r i n m ea su remen t s , i . e . , s t a t is t ica l p robab i l i t ie s f o r s e a sona l d e te rm ina t ion s , m ake s e xpans ion o f t h e c on t ro l

T he re a r e d i f f e rence so fs c a le w h ich mu s t b e c ons ide red w hen i n te rp re t ing h un t ing s t ra teg ie s i nt e rm s o f h uman s oc ia l o rgan isa t ion . I n te rcep t h un t ing c an b e

s amp le a v a luab le d i rec t ion c emen tum g row th l a ye r s .

f o r

f u tu re

r e sea r ch

i n to

b one

T he a pp l icab i l i ty o fs ke le toch rono logy t o

f r agmen ta t ion p a t te rn s f o r r e indee r i nt h e A qu i ta ine B a s in d o n o t s ugge s t c a ch ing o r s mok ing ( Boy le 1 990 ) , i ts e em s

f o s s i l h o r se t e e th i s d emon s tra ted h e re . Howeve r , i ti s s ugge s ted t h a tt r ia ls s hou ld b e c onduc ted p r io r t oe x ten s ive t o o th s e c t ion ing t o e n su re l e g ib i l i ty o f t h e p ropos ed c o l le c t ion s , i . e . , e n sur ing t h a td i agene t ic c hange s o bscur ing t h e g row th l a ye r s h ave n o t o ccur red .

c ha rac ter ised

a s

c ommuna l .

G iven

t h a t

t h e

l i ke ly t h a t ar e la t ive ly l a rge n umbe ro fp eop le w e re e ngaged i n " bu lk " p roce s s ing a c t iv i t ie s . I f i n te r cep t h un t ing o f r e indee r i nt h e A qu i ta ine w a s i ndeed ac ommuna l s t ra tegy , h oweve r , t h is d oe s n o t n ece s sar i ly i mp ly r e g iona l s oc ia l i n te rac t ion a t t h e l e ve l o f t h e c ompo s i te b and - w he re

A s a l ready s t a ted , r e sea rch i n to s e a sona l i n d ica to r s mu s t a l so b e e xpanded t oi n c lude n o t o n ly m o re s i te t ype s ( i .e . , o pen a i r a s w e l l a s c ave s i te l o ca t ion s ) b u t

s ea sona l i n te r cep t h un t s f o rmed t h e e conom ic b a s is f o r l a rge , c o re s iden t ia l g roup ing s . I t s e em s l i ke ly t h a t c ommuna l , i n te r cep t h un t ing d ur ing t h e e a r ly U ppe r P a laeo l i th ic i n vo lved ad eg ree o fr e g iona l i n teg ra t ion . T h is

a l so mo re s pec ie s o fh uman p rey . T he f a c t t h a t m any l o ca l e conom ie s i nt h e A qu i ta ine a r e d om ina ted b y s pe c ie s o t he r t h an t ho se s t ud ied h e re i nd ica te s t h a t w e d o n o t h ave e nough d a ta f o r t h e r e con s truc t ion o f r e g iona l p a t ter n s o f

l i ke l ihood i s b a sed o n v a r iou s t h eor ie s o f i n forma t ion s har ing a nd r i sk r educ t ion . Howeve r , t h e a r gumen t s a r e

E ncoun te r -ba sed h un t ing c an a l so s uppor t

s ubs is tence a nd s e t t lemen t y e t . T he i n c lu s ion o f o t he r s pe c ie s i n as ke le toch rono log ica l s t udy o f s e a sona l s i t e o ccupa t ion s r e qu i re s t h e c o l lec t ion a nd s t udy o f c on t ro l

ar eg iona l ly i n tegra ted , a ggrega ted g roup u nde r c e r ta in , a lbe i tr a re , c i rcum s tance s . T he ref o re ,e v idence o fe ncoun te r

s amp le s a nd t h e c a refu l r e cons truc t ion o f t h e p a t tern o f c emen t d epo s i t ion f o r e ach s pe c ie s .

h igh ly i n f e ren t ia l .

h un t ing o f d i spe r sed r e indee r i n w in te r i sn o t n ece s sar i ly e v idence o f a n i nd iv idua l s t ra tegy -j u s t a s e v idence o f i n te rcep t h un t s d oe s n o t n e ce ssar i ly c on s t i tu te p roof o ft h e e x is tence o f ac ompo s i te b and . We m ay o n ly i n f e r t h e e x is tence o f ar e g iona l ly i n tegr a ted b and , b a sed o n t h e e v idence f o rs e a sona l ly a l terna t ing , e ncoun te ra nd i n te rcep t h un t ing s t ra teg ie s .

7 8

E ndno te s LAG , o r L ine s o f A r re s ted G row th , s e n su C a s tane t ( 1 981 ; 1 982 ) a r e t h in d i scon t inu i t ie s o ccu r ing w hen c emen togene s is r e sume s a f te r t empo ra ry g row th c e ssa t ion ( F ran c i l lon— V ie i l lo te ta l . 1 991 :505 ) . S ee d ef in i t ion s ,b e low . " G round s ub s tance " r e f er s t o t h e a mo rphou s , o rgan ic c omponen to fc emen tum . 3 "M ine ra l d en s i ty " , i n t h is c a se , r e f e r s t o : " t he m a s s o f m ine ra l d epo s i ted p e rv o lume o ft i s sue ." ( F ranc i l lon— V ie i l lo t e ta l . 1 991 :513 ) a nd i st h e re fo re a f f ec ted b y o rgan ic c on ten t a nd p o ro s i ty . I nd ica t ing h ypo the t ica l ly t h a t p r ima ry , s e conda r y a nd e ven a c ce ssory a nnu l ia r ei n d i s t ingu i shab le i nd ec iduou s t e e th . 5T he c a lc ium i o n si nc emen tr eac tw i th t h ep ho spho r ic a c id t o f o rm d i ca lc ium p ho spha te ( Mo skow i tz e t a l . 1 973 ) a nd i s r i n sed a way . 6 S ee J one s & B oyde ( 1974 ) f o r ad e scr ip t ion o f e name l r e so rp t ion p r io rt od epo s i t ion o fc emen t . 7H oweve r , Wa r r en ( 1963 )c on te s t s t h is h ypo the s is . 8E xcep ti nt h er oo t a r ea w he re c emen ti sd epo s i ted o nd en t in ( ame logene s is d oe sn o to c cu ri nt h er ad icu la rr e g ion ) . 9 P o ss ib le c o r r e la t ion b e tween t h e n umbe r o f a syme t r ica l c emen tum l a ye r s l o ca ted o n t h e c r own , t h e r i dge s v i s ib le o n t h e c emen t s u r f ace o f a n u nsec t ioned t o o th j u s t a bove t h e g um l ine a nd t h ea ge o ft h ei n d iv idua ln eed st ob ei n ve s t iga ted . N .b . a r chaeo log ica l s amp le s a r e o f ten d amaged i n t h e r e la t ive ly d e l ica te a p ica lr oo ta r ea . 1 F er n s E quu s c aba l lu s f o rms t wo d i s t in c t k i nd s o f s oc ia l g roup , t h e h a r em a nd t h e b a che lo r g roup ( S imp son 1 950 ; B e rge r 1 986 ; Wa r ing 1 983 ) a nd a r chaeo log ica l a s semb lages m ay r e f lec t as amp le o fe i the r o rb o th t y pe s o fg roup ( Lev ine 1 983 :30 ) . 1 2 I n t h e c a se o f E quu s f er n s , a n a s semb lage m ay r ef lec t s amp l ing o f ab ache lo r g roup a nd b e n a tu ra l ly s kewed i n f a vou r o f y oung a du l t ( c a .4 y ea r s ) a nd s ene scen t m a le s ( Lev ine 1 983 :30 ) . 1 3 T he p rac t ice o fc o r r e la t ing r e g iona l s t ad ia l s equen ce s w i th t h e c l a ss ic A lp ine g l ac ia l s equence a se s tab l ished b y P enck & B ruckne r ( 1909 ) h a s b een t h o rough ly c r i t ic i sed b y K uk la ( 1977 :307 )a nd B owen ( 1 978 :17-20 ) , a mong o t he r s . 1 4 Af ew d eep p o l len c o res p rov ide r a re t e r re s tr ia l r e f e r ence s ca les , s uch a st h eo ne a tG rande P i le ,c . f . Wo i l la rd ( 1 978 ; a s q uo ted i nP e te r son e ta l . 1 979 ; B u tze r 1 982 ) . 1 5 T he m e thodo logy u sed b y L av i l le e ta l . ( 1980 ) t oe s tab l i sh r e la t ive c o r re la t ion s b e tween s egmen t s o f l o ca l p ro f i le s , i . e . , ' c oun t f r om t h e t o p ' ,h a sb een c r i t ic ised i nt h e l i t e ra tu re f o r i t s ' f a i lu re t o s t and u p t oa b so lu te d a ta ( e .g . , K uk la 1 977 ) . T h is c r i t ic ism c an b e e x tended t o t h e s im i la r u se o f t h e ' f o ss i le d i rec teu r ' . 1 6 S ea sona l d a ta f r om G o rdon 's ( 1988 ) s t udy o f r e indee r t e e th a r e i n c luded i nt h is s e c t ion . O n ly t h o se t e e th r a ted a s h i gh ly l e g ib le b y t h e a u tho r ( 4 & 5o n as ca le o f 5 , w he re 5 =h igh ly l e g ib le , 1 = i l leg ib le ) ( p e r s c omm . 1 991 ) w e r e u sed . R esu l t s o f t h e p r e sen t s t udy a r e s umma r ized i n t a b le 4 . 4 ( above ) . 1 7 L aco r r e ( 1953 ) a nd B ouchud ( 1 966 )p r esume t h ee x i s ten ce o fb o th f o re s t a nd t u nd ra s ub— sp ec ie s o fr e indee r . T he r e e x i s t m e tr ica l d i f f e rence s b e tween t h e t wo s ub— spec ie s ( Ke l sa l l 1 968 :29 ) . T he e x is tence o ft wo s ub— spec ies o fr e indee r i nt h e D o rdogne d u r ing t h e l a s t g l ac ia l i sn o t g ene ra l ly a ccep ted t oday , h oweve r , s i nce D e lpech 's s t udy o f r e indee r r ema in s a t L auge r ie h a s s hown t h a t t h e o b served m e tr ica l v a r ia t ion s o ccu rt h rough t i me ( c .f . D e lpech 1 986 ) . 1 8 S ome e qu id s a r e t e r r i to r ia l , e . g . E quu s g r eyv i ( a z eb ra ) a nd t h e w i ld A f r ican a s s , E . a f r icanu s ( We l sh 1 973 :187 ; C lu t ton—B rock 1 987 :85 ) a s w e l l a s t h e f e ra l b u r ro ( We l sh , i b id . ) . T he h o r se i st y p ica l ly n on— te r r i to r ia l , e x i s t ing i nf am i ly h e rd s ( i . e .h a rem s ) ( i b id . ) .

1 9 A l though h i s tor ica l a ccoun t s f r om t h e P ampa s r eg ion d ocumen t l a rge n umbe r s o f a n ima l s mov ing t oge the r , p r e sumeab ly d u r ing t h e l a te f a l l a s h e rd s move t op ro tec ted l o ca t ion s ( Fa lkne r 1 935 :37 a s q uo ted i nA n thony 1 991 :253 ) , t h is m ay b e a n e xc ep t iona l p henomenon i n f luenced b y l o ca l t o pog raphy . 2 0 T he c a lv ing d a te f o r r e indee ri sg ene ra l ly c on s ide red t ob e ag i ven . Howeve r , K e lsa l l ( 1968 :177 ) d escr ibe s v a r ia t ion i n c a lv ing a s so c ia ted w i th w ea the r— induced c hanges i nt h e t empo o f r u t t ing . H o r ses g ene ra l ly f oa l i n May / June (Wa r ing 1 983 :134 ; G roves 1 974 :63 ) a l though b i r th s may t a ke p lace y ea rr ound ,e ven i nt h e w i ld (We l sh 1 975 :27 ) . 2 1 B ouchud ( 1966 ) u sed e rup t ion t a b les t oa ge a n ima ls u p t o 2 9 mon th s , u s ing s i ng le t e e th w he re n ec essa ry , i n va l ida t ing h i s a na ly s is a cco rd ing t oB in fo rd ( 1973 :239 ) . 2 2 B in fo rd ( 1978 ) c a lcu la ted t h e se v a lues u s ing v a r iou s b ody p a r t u t i l i ty i n dexe s i n c lud ing M .G .U . I . ( t he mod if ied g ene ra l u t i l i ty i n dex ) . T he se i n dexe s h ave b een c r i t ic ised a s t oo s pec if ic t o t h e d a ta c o l lec t ing c on tex t , i . e . t h e Nunam iu t e conomy , a nd o t he r mo re g ene ra l ised i n dexe s h ave b een p ropo sed ,e . g . , Me tca lf & J ones ( 1988 ) . 2 3 No te t h a t t h i s i n fo rma t ion c an b e u sed t o s uppor t l a te s p r ing ,e a r ly f a l l a nd /o rs umme rd ea th s . 2 4 G o rdon h imse lf s t a tes t h a t : "W in te r a nd e a r ly s p r ing p rov ide t h e mo s t c r ed ib le r e su l t s [ . . ]" ( 1988 :29 ; c . f . a l so 1 990 :69 ; Wh i te 1 990 :70 ) . P resumab ly , c on fu s ion w ou ld o ccu r b e tween s umme r a nd f a l l i n c remen t s . Howeve r , f ew s umme r i n c remen t s w e re i d en t if ied b yG o rdon . 2 5 No t a l l r e indee r m ig ra te e ve ry y ea r , i n d iv idua ls m ay r ema in i n o r n ea r t h e w in te r r anges d ue t o av a r ie ty o f c i rcum s tance s s uch a s i lnes s ( Laco r re 1 953 , 1 958 ) . F u r the rmo re , b u l l a nd j uven i le g roup s t yp ica l ly l a g b eh ind t h e c ow sd ur ing s p r ing m ig ra t ion , n o t a dvanc ing a sd eep ly i n to t h e s umme r r a nges ( Ke lsa l l 1 957 :29 ; M i l le r 1 974 :77 ; Go rdon 1 988 :8 ,14 ) . 2 6 N .b . t h i s c ou ld b e e v idence o ff a l l a nd /o rs p r ing , a s w e l l a s w in te re xp lo i ta t ion o fh o r se i nt h is h o r izon . 2 7 I ti si n te re s t ing t on o te t h a t r e indee r movemen t s i nt h e f a l l h ave b een d e sc r ibed a s c on fu sed , i . e . l e ss p u rpo sefu l t h an s p r ing movemen t s ( Ke lsa l l 1 968 :174 ) . R e indee r a r e a l so i n c rea s ing ly w a ry d u r ing t h is s ea son , a nd a pp roached w i th m o re d i f f icu l ty t han i n t h e s pr ing ( Ke lsa l l 1 957 :44 ) . A lso , s p r ing a nd f a l l m ig ra t ion c o r r ido r s d o n o t a lway s c o inc ide ( Go rdon 1 988 ) . I t may t h e refo r e n o t b e a s s t range a s i ts eem s a tf i r s t t h a t a tn one o ft h e s i te s s u rveyed b y G o rdon ( 1988 ) d o s pr ing a nd f a l le xp lo i ta t ion p a t te rn s f o rr e indee rc o inc ide . 2 8 T he g ene ra l mode lp resen ted i st h a t o ft h e c ompo s i te b and ( Lee & D evo r e 1 969 ) . C .f . L ee & D evo re ( 1969 :7 ) a nd c on t r ibu to r s t ot h e i rv o lume f o rc r i t ic ism s o fe a r l ie r mode l s o f t h e p a tr i lo ca l b and o f fe red b y R adc l if f e—B rown ( 1931) , S t ewa rd ( 1936 ) a nd S er v ice ( 1962 ) . F o r ac r i t ic i sm o f b and t h eo ry a s an o rma t ive mode l , s e e B e t t inge r( 1983 ) . F o r amo re a b s t rac t c oncep t o f t h e r eg iona l b and , e ven g r ea te r e mpha s is o n f l u id i ty a t t h e l o ca l l e ve l a nd e mpha s is o n i n d iv idua l r e la t ion s r a the r t h an r e la t ion s b e tween l o ca l g roup s a s ab a s is f o rr e g iona l o rgan isa t ion , s ee Mye r s ( 1989 :73 ,9 7 ) . 2 9 E .g . G amb le ( 1986 :39 ) s t a tes t h a t " Hun te r—ga the r e r o rgan isa t ion i s as y s tem t r an sforma t ion o f t h e u nde r ly ing e co log ica l s t ruc tu res " . A nd c . f . S t ewa rd ( 1977 :43 ) . 3 0 B in fo rd , af r equen t c r i t ic o f o p t ima l f o rag ing t h eo r ies ( 1982 ; 1 983 ) , e ngages i n m idd le r ange r e sea rch , b u t c . f . B e t t inge r( 1991 :107 ) f o rd i scus s ion o fo p t ima l f o rag ing t h eo ry a sm idd le r a nge r e sea rch a nd ( 1991 :108 ) f o r ar app ro chemen t b e tween B in fo rd 's ( 1978 ) w o rk a nd g ene ra l s ub s is tence t h eory , o rc u l tu ra le co logy . 3 1 N .b . N on— con f igu ra t iona l o p t ima l f o rag ing mode ls , s u ch a s t h e l i nea r p rog ramm ing mode l , f o r e xamp le , h ave a dded a f u r the r d imen s ion o f l im i t ing f a c to r s , i . e . b i nd ing c on s t ra in t s

2

7 9

( e .g .c r i t ica ln u t r ien t s ; K eene 1 981) . 3 2 A no the rr e f inemen to ft h i s e qua t ion i st h e Ma rg ina l V a lue T heo rem , i n t roduc ed b y C hamov ( 1976 ) , w h ich p red ic t s t h e a moun to ft i me s pen te xp lo i t ing ag i ven p a tch . 3 H oweve r , a s S t ewa rd ( 1 977 :43 ) s t a te s , c u l tu ra l e co logy " do e s n o t e qua te s oc ia l f e a tu r es w i th b i o log ica l s pec ie s o r a s sume t h a tc omp e t i t ion i st h em a jo rp ro ce ss ." 3 4 C .f . a l so F r eeman ( 1 973 :40 ) o n t h e f a i lu r e t o i d en t i fy h un t ing c amp sa nd b a se c amp si nt h eP y r enean r e co rd . 3 5 T he p r ec i se d ef in i t ion o ft h e t e rm " h e rd f o l low ing " i ss t i l l u n r e so lved , h oweve r , a s i s e v iden t i n B leh r ' s ( 1991 :444 ) c ommen t : " ( . . . ) i tw a s n o t t h e c ha rac te r o f t h e c on ta c t b u t i t s ' e x t en s ion t o m o s t o f t h e a nnua l c yc le t h a t g ave ( t h e C h ipewyan t h e i r ) s t a tu s a s h e rd f o l lowe r s " . ( Pa r en these s a dd ed ) . 3 6 P y r enean h e rd s a r e c on s ide r ed l e s s mob i le t h an P e r igo rd ian h e rd sd ue t ot h ec omp re s sed z ona t ion o ft h er e g ion ( 1 977 :247 ) . 3 7 T h i s p ropo r t ion d rop s t o 4 0% w hen l i ve w e igh t i s c on s ide r ed , c . f . S p ie ss ' d i scu s s ion o f % t o ta l u ngu la te MN I , v e r su s % o f t o ta l e s t ima ted l i ve w e igh t , i n t a b le 6 . 16 a nd f i g .6-7 ( i nS p ie s s 1 979 ) . 3 8 T he e x i s tenc e o f 3 , 000 r e indee r r ema in s a t G ou rdan , f o u i l le sP ie t te ,e v idence o ft h ei n te r cep t ion o fm ig ra to ry h e rd s , i st a ken a s s uppo r t ive o ft h e h e rd f o l low ing mode l i nC lo t t e s ( 1 987 :30 5 ) . 3 9 T he e x i s tence o f L es E g l ise s , a" h a l te d e c ha sse " , a nd " c amp s d e b a se " s u ch a s E n lene a nd G ou rdan , i s u sed t o s uppo r tt h es e a sona lr ound s mode l ( C lo t te s 1 987 :324 ) . 4 0 N o t e t h a t t h i s s ma l l s amp le s i ze i sn o t ar e su l t o f t h e S p ie s s ' s amp l ing d e s ign ,b u to fe i the rp oo rp r ese rva t ion o ft h e t e e th o rm e thodo log ica ld i f f icu l t ie s ,s i nce 1 7 1 s pec imen s w e r e p r epa r ed ( 1 979 :t a b le 6 . 2 ) . 4 1 R e s iden t ia l s h i f t s a r e f r equen t ly c o r re la ted w i th s ea sona l a va i lab i l i ty o f r e indee r i n t h e mode l s d i scu s sed , w he the r e conom ic s pe c ia l i sa t ion i sa s sumed o rn o t . 4 2 I tm ay e ven b e c on s t rued a s e v idence o f y ea r r o und o c cupa t ion , s i nce i ti sn o tp o s s ib le t ok now t h e d i s t r ibu t ion o f h o r se d ea th sw i th in t h e May—O c t . a t t r ibu t ion . 4 3 E ncoun te r—ba sed h un t ing c an a l so b e ac ommuna ls t ra tegy , t a k ing p l ac e d u r ing a " pub l ic " , o r a gg lome ra ted p ha se o f s o c ia l o rgan i sa t ion , e . g . w in te r s ea l h un t ing a mong t h e N e t s i l ik I n u i t . 4 4 E spec ia l ly w i th r e ga rd s t o d i ff i cu l t ie s e ncoun te red i d en t ify ing s i t e f u n c t ion w i th in a r e g iona l s y s tem ( c .f . F r eeman 1 973 ;K oe t je 1 987 ) . 4 5 A ge i ss ugge s ted e l sewhe re a s af a c to r e xp la in ing t h e d i f f e r ence s i n r a te o f g row th d u r ing d eve lopmen t ( e .g . K leveza l 1 987 ) . 4 6 T he e x i s tence o f t wo s ympa t r ic s pec ie s o f r e indee r ( R . t a r andu s s y lva t icu s a nd R . t a randu s ) i n t h e S ou thwe s t i s r e jec ted i n f a vou r o f t h e t u nd ra d we l l ing f o rm , R . t a randu s ( D e lpech 1 986 ) . 4 7 C . f . D e lpech 1 983 , 1 987 . 4 8 A l though n o t i nt h e s t udy r e g ion , t h e s i te o fS o lu t r e , i nt h e e a s t e rn f oo th i l l s o ft h e Ma s s if C en t ra l ,b ea r s m en t ion . S o lu t r e i ss i tua ted i n at r an s i t ion z one b e tween h i l l a nd p l a in b i o tope s a nd a pp ea r s t oh av e b een ab a se f o r t h e i n te r cep t ion o fs ma l l , m ig ra to ry h e rd so fh o r se ( O lsen 1 989 :316 ) . 4 9 I nt e rm s o fc a lo r ic e xpend i tu re ( du r ing s ea r ch , p u r su i t a nd h and l ing )r e la t ive t oe ne rg e t ic g a in sp e ru n i to ft i me . 5 0 U n fo r tuna t e ly , g ood c h rono s t ra t ig raph ic c on t ro l i sr a r e ly c omb ined w i th g ood f a una lp r e se rva t ion ,e . g . o pen—a i rs i te s .

8 0

B IBL IOGRAPHY A a jane ,A .

a nd C o .

1 986 C on t r ibu t ion a L 'E tude d e s F aunes d u Wu rm R ecen t e n H au te—Lo i re : l e s N iveaux Magda len ien s d u R ond d u B a r ry . U n ive r s i te d e B o rdeaux : T he se 3 e C yc le .

A rambou rou ,R . 1 976 P au—Agen : G ro t te D u ru thy . L iv re t—Gu ide d e L 'Excu r s ion A 4 , S ud—Oue s t . I Xe C ong res d e l ' U ISPP , N ice .

A dam s , L .a nd S .G . Wa tk in s 1 967 A nnu l i i n T oo th C emen tum I n d ica te a ge i n C a l ifo rn ia G round S k i r re l s . J ou rna l o f W i ld l i f e B io logy 3 1 :836-839 .

1 979 L a f i n d e s T emp s G lac ia i re s äD u ru thy , C ommune d e S o rde—L 'Abbaye ( Landes ) . I nL a F in d e s T emp s G lac ia i res e n E u rope . C o l lec t ion I n ter na t iona le d uC .N .R .S . n o .271 ,e d . C .N .R .S . .

A ge r ,D .V . 1 980 T he G eo logy o fE u rope . N ew Y o rk : H a l s tead P res s .

A r to i s , M . , a nd D . S a lmon 1 981 D e te rm ina t ion E xp er imen ta le d u S exe e t d e L ' äge C hez l eR ena rd R oux ( Vu lpes V u lpes ) : V a l id i te e t R ep roduc t ib i l i te d es T echn ique s C ho i s ies .

A i tken ,R . J . 1 975 C emen tum L aye r s a nd T oo th Wea r a sC r i te r ia

Mamma l ia 4 5 (3 ) :373-378 .

f o r A ge ing R oe D ee r ( Cap reo lu s C ap r eo lu s ) . J ou rna l o fZ oo logy ( London ) 1 75 :15-28 .

A sch n d T he S lavey I n d ian s : t h eR e levance o fE thnoh is tory t oD eve lopmen t .

A lb r igh t ,D . 1 989 P r ec i s ion o f S ea sona l i ty D e te rm ina t ion i n

A l la in ,J . 1 976 L es C iv i l isa t ion s d u P a leo l i th ique S upe r ieu r

B ag l in ie r e , J .L . , J . C a s tane t , F . C onand , a nd F . J . Meun ie r 1 991 T e rm ino log ie e n S c le ro ch rono log ie c hez l e s V er teb re s . A c te s d e l aC on f e rence T issu s D u r s e tA ge L n d iv idue l c hez l e s V er teb res . B ondy , 1 991 .

D an s l e s ud—Oue s t d u B a s s in P a r is ien . I n L a P reh is to i re F rança ise . H . d e L um ley , e d . p p .

B ahn ,P .

R inged S ea l s ( Phoca H i sp ida ) . P ap e r P re sen ted a tt h e A nnua l S .A .A . Mee t ing s ,A t lan ta , 1 989 .

1 315-1320 .P a r i s : E d i t ion sd uC .N .R .S . .

1 977

S easona l M ig ra t ion i n S ou thwe s t F rance

Amp r ino ,R .

D u r ing t h e L a te G lac ia l P er iod . A r chaeo log ica l S c ience . 4 : 245-257 .

1 947 L a S t ruc tu r e d u T i s su O s seux E nv isa ge C omme E xp re ss ion d e D i f f e rences D an s l aV i tes se d e L 'acc ro issemen t . A rch ive s B io log ique s5 8 :315-330 .

1 983 L a te P le is to cene E conom ies o f t h e F rench P y renee s . I nH un te r—Ga the re r E conomy i nP reh is to ry . G . B a i ley ,

J ou rna l

o f

e d . p p .168-186 . C amb r idge :

1 967 B one H is tophy s io logy . G uy 's H o sp i ta l R epo r t 16 :51-69 .

C amb r idge U n ive r s i ty P ress .

A nde r son ,J .G .

t h e F rench S i tes . Wa rm in s te r : Ans a nd P h i l l ip s L td .

1 984 P y renean P reh is tory . AP a laeo l i th ic S u rvey o f 1 978 T he S t ruc tu re o f We s ter n E u rope . O x fo rd : P e rgamon P re s s .

B a l ick i ,A . 1 970 T he N e t s i l ik E sk imo . N ew Y o rk : Na tu ra l

A nd r ieu ,P . 1 975 O b se rva t ion s a u S u je t d e l aP ub l ica t ion d eJ . P .

H is to ry P r e ss .

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B on f ie ld , A .W .F . 1 954 P r e l im ina ry I n ves t iga t ion o ft h e B a r ren—g round C a r ibou . P a r t s 1a nd 2 .W i ld l if e Managemen t B u l le t in ,

A n thony , D .

O t tawa .

1 991 T he D ome s t ica t ion o f t h e H o r se . I nE qu id s i n t h e A nc ien t Wo r ld , V o l . I I . R . Meadow a nd H .—P .

1 96 1 AR ev is ion o ft h e R e indee r a nd C a r ibou , G enu s R ang if e r . B io logy s e r ies n o .66 . N a t iona l Mu seums o f

U e rpmann e d .s p p .250— 2 77 . G ö t t ingen : H uber t

8 1

C anada B u l le t in 1 77 . 1 982 T he A r cha eo logy o f P lace . J ou rna l o f A n th ropo log ica lA r cha eo logy 1 : 5-31 .

B ay le d e sH e rmen s ,R .d e 1 974 V ue D 'en semb le s u rl e sN iveaux P reh i s to r iques d e l a G ro t te d u R ond d u B a r ry , F ou i l le s 1 966-73 . B u l le t in d e l a S oc ie te P r eh i s to r ique F ran 9a i se . 7 1 ( 5 ) :131 — f f .

1 983 I nP u r su i t o f t h e P a s t . G e rman D emo c ra t ic R epub l ic :T hame sa nd H ud son . 1 987 S ea r ch ing f o r C amp s a nd M i ss ing t h e E v idenc e? I n T h e P l e i s to cene o f t h e O ld Wo r ld . 0 . S o f f e r ,e d .p p . 1 7-320 .N ew Y o rk :P l enum P re s s .

1 977 L e s N iveaux S upe r ieu r s d u Magda len ien d e l a G ro t te d u R ond d u B a r ry , P o l ignac ( Hau te L o i r e ) . I n L aF i n d e s T emp s G lac ia i r e s . D . d e S onnev i l le— B o rd e s ,e d .p p .60 1-6 14 .E d i t ion sd u CNRS .

B lan cha rd ,H . ,e ta l . 1 972 L e G i semen t e t l e S que le t te d e S a in t—Ge rma in— la—R iv ie r e . A r ch ive s d e L ' In s t i tu t d e P a leon to log ie H uma in e n o .34 .P a r i s .

B eaune—Rome ra ,S . ,a nd A .R ou s so t 1 982 L e s l amp e s d e S a in t—Ge rma in— la—R iv ie r e . R evue H i s to r ique e tA r cheo log ique d u L ibou rna i s e t d el aV a l lee d el aD o rdogne 5 0 (185 ) :10 1-10 9 .

B leh r ,0 . 1 99 1 R ep ly t o B u r ch . I N C u r ren t A n th ropo logy . 3 2 (4 ) :444— 4 45 .

B egouen ,R . ,a nd J .C lo t te s 1 981 N ouve l le s F ou i l le s D an s l aS a l le d e s Mo r t sd e l aC aveme D 'En lene äMon te squ ieu—Avan te s ( A r iege ) . E x t ra i t d u C ong re s P r eh i s to r ique d e F ran ce XX Ie S e s s ion .Q ue r cy 1 979 .

B ob r in sk i i ,N .A . ,B .A .K ur ie t sov ,a nd A .P . K uz iak in 1 944 K ey t ot h e Mamma l s o ft h e U .S .S .R . Mo scow : S ta te P ub l i sh ing H ou se . ( T ran s la ted b y J .D . J ack son , 1 948 , f o rt h e B u r eau o fA n ima l P opu la t ion , D ep t . o f S oo logy ,F i e ld S t ud ie s ,O xfo rd .T ran s la t ion n o .200 ) .

B e rge r ,J . 1 986 W i ld H o r ses o f t h e G r ea t B a s in . C h icago : U n ive r s i ty o fC h icago P re s s .

B o rde s ,F . 1 958 N ouve l le s F ou i l le s aL auge r ie—Hau te E s t . P rem ie r sR e su l ta t s . L 'An th ropo log ie 6 2 (3 ) :205-242 .

B e t t inge r ,R .L . 1 982 E xp lana to ry /p r ed ic t ive Mode l s o f H un te r—Ga the r e r A dap ta t ion . I n A dvance s i n A r chaeo log ica l Me thod a nd T heo ry , S e lec t ion f o r S t uden t s V o l . 1T h rough 4 . M .B . S ch i f f e r , e d . p p . 1 5 7-2 10 .N ew Y o rk :A cadem ic P r e s s . 1 99 1 H un te r—Ga the re r s . A r cha e log ica l E vo lu t iona ry T heo ry .N ew Y o rk :P lenum P re ss .

1 972 C i r con sc r ip t ion D 'Aqu i ta in e . G a l l ia P r eh i s to i r e 1 5 :487-497 . 1 973 O n t h e C h rono logy a nd C on t empo rane i ty o f D i f f e r en t P a leo l i th ic C u l tu r e s i n F ran ce . I n T h e E xp lana t ion o f C u l tu r e C hang e . C . R en f r ew , e d . p p . 2 17-226 .L ondon : D u ckwo r th .

a nd

B o rde sF . ,a nd D . D eS onnev i l le—Bo rde s 1 970 T h e S i gn if i cance o f V a r iab i l i ty i nP a leo l i th ic A s semb lag e s . Wo r ld A r chaeo logy 2 : 6 1-73 .

B in fo rd ,L .R . 1 973 I n te ra s semb lage V a r iab i l i ty— t h e Mou s te r ian a nd t h e " Func t iona l A rgumen t " . I nT he E xp lana t ion o f C u l tu r e c hange : Mode l s i nP r eh i s to ry . C . R en f r ew , e d . D u ckwo r th P r e s s .

B o rde sF . ,a nd H .P ra t 1 96 5 O b se rva t ion s s u r l e s F aune s d u R i s s e t d u Wu rm Ie nD o rdogne .L 'An th ropo log ie 6 9 :31-46 .

1 978 N unam iu t E thnoa rchaeo logy . N ew Y o rk : A cadem ic P re s s .

B o rde s , 1 980 W i l low Smoke a nd D og s ' T a i ls : H un te r—Ga the r e r S e t t lemen t S y s tem s a nd A r ch eo log ica l S i te F o rma t ion . Ame r ican A n t iqu i ty 4 5 :4-20 .

F . ,

J . —Ph .

R igaud ,

a nd

D .

D e

S onnev i l le—Bo rde s 1 972 D es B u t s , P rob leme s e t L im i tes d e L 'a r ch eo log ie P a leo l i th ique .Q ua tema r ia 1 6 :15-35 . B ouchud ,J . 1 966 E s sa i s u r l e R enn e e t l a C l ima to log ie d u P a leo l i th iqu e Moyen e t S up e r ieu r . P e r igueux : I mp .

1 98 1 B one s : A n c ien t Men a nd Modem My th s . N ew Y o rk : A cadem ic P r e s s .

8 2

Ma sse .

O don to ce te s .

R epo r t

C omm i ss ion .

o f t h e

I n terna t iona l Wha l ing

S p ec ia l I s sue n o .3 :65-87 .

B ouchud ,J . ,A . ,C hey rüe r ,a nd Y . G u i l l ien 1 953 D en t s d e R enne e tM ig ra t ion s . B u l le t in d e l a S o c ie te P reh is tor ique F rança i se 5 0 (9-10 ) :127132 . B ou rde l le ,Y .

1 987 D eve lopmen ta l I n te rp re ta t ion s o f D en ta l M ic ro s t ruc tu r e . P ape r p r e sen ted a t t h e Wenne r—G ren C on f e rence " P r ima te L i f e H i s to ry a nd E vo lu t ion " , C abo S an L uca s , B a ja , Mex ico ,O c tobe r 1 987 .

1 977 L 'Ab r i D ur if ä E nva l : C ommune d e V ic— le— C om te . I N L a F in d e s T emp s G lac ia i res . D . d e S onnev i l le— B o rdes e d . p p .523-530 . E d . d u CNRS .

B oyde , A . , a nd S . J .J one s 1 972 M ine ra l ized T i ssues : S EM S tud ie s I n D eve lopmen ta l A spec t s o fO ra l B io logy . H .C . S lavk in , L .A . B ave t ta A cadem ic P re ss .

B ou rde l le ,Y . ,H . , D e lpo r te , a nd J .V irmon t 1 971 L eG isemen t Magda len ien e tl aV enu s D 'Enva l , C ommune d e V ic— le—Com te ( Puy—de—Döme ) . L 'An th ropo log ie 7 5 (1) :119128 .

1 67-174 . N ew Y o rk :

B oy le , K . 1 990 Uppe r P a laeo l i th ic F auna s F rom S ou th Wes t F rance . O xfo rd : B r i t ish A r chaeo log ica l R epo r t s , n o . S 557 .

B ou rde l le ,Y ,a nd J . —C . Me r le t 1 987 Nouve l les D ecouver te s D 'a r t Mob i l ie r aE nva l ( Puy—de—Döme ) . i L 'A r t Mob i l ie r e t s on C on tex te . A c te s 2 55-259 .

e d .s p p .

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d u C o l loque d e F o ix— le Mas D 'Az i l . p p .

1 987 T he c a se o f t h e r e g iona l p e r sp ec t ive : an ew w o r ld v i ew . I nT he P le i s to cene o f t h e O ld Wo r ld . 0 . S of f e r , e d . p p .365-372 . N ew Y o rk : P l enum P re ss .

B ouv ie r ,J . —M . , a nd G . T reco l le 1 966 L a F r i se G ray & d e S a in t—Germa in— la— R iv ie re . l ' An th ropo log ie 7 0 (5-6 ) :535-540 .

B uf f ren i l , V . d e 1 980 M i se e n

B ou rque ,B . J . ,K . Mo r r i s ,a nd A . S p iess

C ond i t ion s d e M i l ieu s u r l aC ro issance d e C ro cody lu s S iamen s is ( Schne ide r , 1 80 1) e t V a leu r d e s Ma rques .

E v id ence d e

L ' inc idence

A r ch ives Z oo log ique s 1 21 :63-76 .

1 978 D e term in ing t h e S eason o f D ea th o f Mamma l T ee th F rom A r cheo log ica l S i te s : an ew S ec t ion ing T echn ique . S c ience 1 99 :530-531 .

B u r ch , E .S . 1 972 T he C a r ibou /w i ld R e indee r a s aH uman

B ouv ie r ,J .M .

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1 969 L a C ha i re—ä—Ca lv in ( Mou th ie r s , C ha r en te ) D onnee s e t P rob leme s . L iv re t—Gu ide d e L ' excu r s ion A 4 ,V I I Ie C ong res d eL ' INQUA .p p .95102 .

1 991 H e rd F o l low ing R econ s ide red . A n th ropo logy 3 2 (4 ) : 4 39-445 .

1 976

L e G isemen t d e l a Made le ine , C ommune d e

I n c remen ta l S t r uc tu res . P ape r P resen ted a t t h e S AA A nnua l

L a Made le ine : A cqu is R ecen ts . G lac ia i re s e n E u rop e .

L a F in d es C o l loques

D u r ing t h e U ppe r P a laeo l i th ic i n S ou ther n F rance . P h .d . d i sser ta t ion , N ew Y o rk Un ive r s i ty , May 1 992 . 1 993 O b ser va t ion o fi n c r emen ta l g row th s t ruc tu re s i n

A S t ra t ig , raph ic Wo rk . O xfo rd :

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e l ec t ron

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1 979 Q ua ter na ry G eo logy . N a tu r e2 77 :171-172 . B oyde ,A . 1 980

Mee t ing , A t lan ta , 1 988 .

1 992 P rey Movemen t s a nd S e t t lemen t P a t te rn s

I n tema t ionaux :e d i t ion sd uC .N .R .S . B owen ,D .Q . 1 978 Q ua te rna ry G eo logy : F ramewo rk f o r Mu l t id isc ip l ina r y

C u r r en t

B u rke , A . 1 988 A n S .E .M . E va lua t ion o f D en ta l C emen tum

T u r sac . L iv r e t—Gu ide d e L ' excu r s ion A 4 , I Xe C ong res d e L 'U ISSP ,N ice .P p .108-109 . 1 977 T emp s

d e s

C amb r idge U n iv . P r ess . H is to log ica l S t ud ie s o f D en ta l T issue s o f

8 3

C ae tano M .H . ,J .C a s tane t ,a nd H . F ranc i l lon 1 985 D e te rm ina t ion d e L ' äg e d e T r i tu ru s Ma rmo ra tu s Ma rmo ra tu s ( La t r e i l le 1 800 ) d u P a r c N a t iona l d e P eneda G e r e s ( Po r tuga l ) p a r S que le t toch rono log ie . Amph ib ia—Rep t i l ia 6 : 117132 .

a nd P opu la t ion S t ru c tu re o f t h e T ua ta ra , S phenodon P u n c ta tu s , o n S t ephen s a nd L ady A l ice I s land s . H e rp e to log ia 4 4 (1 ) :25-37 . C ha l in e ,J . 1 977 L e s Mod if ica t ion s d e P ay sages e td e C l ima t s d e l af i n d e s T emp s G lac ia i r e s e n F rance ( doma ine B o r ea l ) R eve lee sp a rl e sM ig ra t ion sd eR ongeu r s . I n L a F i n d e s T emp s G lac ia i r e s e n E u rope . D . d e S onn ev i l le—Bo rde s ,e d .p p . 9 7103 .P a r is :C .N .R .S . .

C ap i tan ,L .a nd D . P ey rony 1 928 L a Made le ine — s on G i semen t , s on I n du s t r ie , s e s O euv re s d 'A r t . P ub l ica t ion s d e l ' I n s t i tu t i n ter na t iona ld 'An th ropo log ie n o .2 .P a r i s . C a s tane t ,J . 1 979 D onn ee s C ompa ra t ive s s u r l aM ine ra l i sa t ion d e s Ma rque s d e C ro i s sance S que le t t ique s C hez l e s V e r teb re s . P a r i s : C omp te sR endu sd e l ' Academ ie d e sS c ience sd eP a r i s2 89 :405-408 .

C har nov ,E .L . 1 9 76 O p t ima l F o rag ing : T he Ma rg ina l V a lue T h eo r em .T heo re t ica lP opu la t ion B io logy 9 : 129-136 . C h er nov ,I .U . I . 1 985 T he L iv ing T und ra . C amb r idge : C amb r idge U n ive r s i ty P r e s s .

1 980 Q ue lque s R ema rque s s u r l a Me thode S que le t toch rono log ique C hez l e s V e r teb re s S upe r ieu r s ( o iseaux e t Mamm if e re s ) . B u l le t in S o c ie te Z oo log ique d eF rance 1 05 (2 ) :371-376 .

C h eyn ie r ,A . 1 960 L e s C ouch e s I n f e r ieu r s d e L auge r ie—Hau te . B u l le t in d e l aS o c ie te H i s to r ique e tA r cheo log ique d u P e r igo rd 8 7 (3 ) :132136 .

1 981 Nouve l le s D onnee s s u rl e s L igne s C imen tan t e s d eL 'o s . A r ch ives B io log iques . ( Br uxe l le s )9 2 :1-24 .

C h eyn ie r ,A . ,J .B ou chud ,a nd Y .G u i l l ien 1 9 52 L e sB o i sd eR enne d eB adegou le . B u l le t in d el a S o c ie te P r eh i s to r ique F r ança i se 4 9 :53-55 .

1 982 R eche r ch e s s u r l aC ro i s san ce d u T i s su O s seux d e s R ep t i le s . A pp l ica t ion : l a Me thode S que le t toch rono log ique . T h ese d e D o c to ra t d 'E ta t , U n ive r s i te d eP a r i sV I I ,J u in 1 982 .

C la rk ,G .A . 1 9 80 C ommen t o n C onkey : P reh i s to r ic H un te r— G a th e r e r A gg rega t ion S i te s C u r ren t A n th ropo logy 2 1 (5 ) :62 1-622 .

1 986-87 L a S que le t toch rono log ie C hez l e s R ep t i le s I I— A pp l ica t ion . P a r i s : A nna le s d e s S c ien ce s N a tu re I le s , Z oo log ie , 1 3e S e r ie 8 : 157172 .

C .L . I .M .A .P . 1 976 T h e S u r f ace o f I c e—Age E a r th . S c ience 1 9 1(4232 ) :11311137 .

C a s tane tJ . ,a nd M . B a ez 1 988 D a ta o n a ge a nd L ongev i ty i nG a lo t ia G a l lo t i ( Sauna , L ace r t idae ) A s se s sed b y S ke le to ch rono logy . H e rp e to log ica lJ ou rna l . 1 : 218-222 .

C lo t te s ,J . 1 987 L e Magda len ien d e s P y renees . A c te s d u

C a s tane tJ . ,a nd A .d eR icq le s 1 986-87 S u r l aR e la t iv i te d e l aN o t ion D 'o s teone s P r ima i re s e t S econda i r e s e t d e T i s su s O s seux P r ima i r e s e t S econda i re s e n G en e ra l . A rm ie s d e S c ience sN a tu r e l le s ,Z oo log ie ,P a r i s8 : 103109 .

C o l loque d e Mayence . E RAUL 3 8 :281-360 .

C a s tane tJ . ,F . J . Meun ie r ,a nd A .d eR icq le s 1 976 L ' en reg i s t r emen td e l aC ro i s san ce C yc l ique p a r l eT i s su O s seux C hez l e s V e r teb r e s P o ik i lo the r rne s : D onnee s C ompa ra t ive s e t E s sa i d e S yn th e se . P ap e r P r e sen ted t o t h e S oc ie te Z oo log ique d e F rance , S t ruc tu r e s e tF onc t ion s ,P a r i s , 1S ep t . 1 976 .

C .O .H .M .A .P . 1 988 C l ima t ic C hange s o f t h e L a s t 1 8000 Y ea r s : O b se r va t io n a nd Mode l S imu la t ion . S c ience 2 4 1 : 104310 50 .

C lu t ton—B ro ck ,J . 1 9 89 A N a tu ra l H is to ry o f D ome s t ic A n ima l s . L ondon :C .U .P . B r i t i sh Mu seum ( Na tu ra lH i s to ry ) .

C o l l in son ,A .S . 1 988 I n t roduc t ion t o Wo r ld V ege ta t ion . L ondon : U nw in H yman .

C a s tane t ,J . ,D .G .N ewman ,a nd H .S a in tG i ron s 1 988 S ke le toch rono log ica l D a ta o n t h e G row th , a g e

8 4

C onkey ,M . 1 980 T he I d en t if ica t ion o f P reh i s to r ic H un te r—Ga the r e r A gg rega t ion S i te s : t h e C a se o f A l tam i ra . C u r ren t A n th ropo logy 2 1 (5 ) :609-630 .

1 988 R ev iew o f Men a nd R e indee r H e rd s i nF rench Magda len ian P reh i s tory , b y B . G o rdon . B u l le t in d e l a S o c ie te P reh i s to r ique F rança ise ( 8 5 )9 : 280-282 .

C oucha rd ,J . 1 966 L a S t ra t ig raph ie d u G isemen t B adegou le—Oues t . L 'An th ropo log ie 7 0 (1) :17-28 .

D e lp ech F . , E . D ona rd , A . G i lber t , J . —L . G uade l l i , 0 . L eGa l l , A . Ma r t in i— Jaqu in , M .— M . P aque reau , F . P ra t , a nd J . —F . T ou rnep iche 1 983 C on t r ibu t ion ä l a L ec tu re d es P a leoc l ima t s Q ua ter na i re s D 'ap ie s l e s D onnees d e l aP a leon to log ie e nM i l ieu C on t inen ta l . A c tes d u C o l loque AGSO , B o rdeaux , Ma i 1 983 . B u l le t in d e l ' I n s t i tu t G eo log ique

d e

C oy ,J . P . ,e ta l . 1 982 A b so lu te A ge ing o fC a t t le F rom T oo th S ec t ion s a nd i t s R e levance t o A rchaeo logy . I n A ge ing a nd S ex ing A n ima l B one s F rom A r chaeo log ica l S i te s . W i lson , G r ig son a nd P ayne e d .s . B r i t i sh A rchaeo log ica lR epor t s , B r i t i sh S e r ie sn o .109 .

d uB a ss in

D 'Aqu i ta ine ,n o . s pec ia l ,p p .165-177 .

D e lpech ,F . , a nd D . d e S onnev i l le—Bo rde s 1 977 L ' Indu s t r ie d e L 'o s ä L auger ie—Hau te ,

D 'Hua r t ,J . P . 1 980 D e term ina t ion d e L age p a r l e s A nneaux d e C ro i s sance d e s T i ssus D en ta i res : u ne N ouve l le Me thode d e P r epa ra t ion s u r H y locho erus ( Su idae ) . Mamma l ia 4 4 (1) :128-136 .

D o rdogne ( f ou i l les F . B o rde s ) : D eb i tage e tO u t i ls d e F o r tune . C o l lec t ion I n te rna t iona le d u C .N .R .S . 5 68 :6 1-68 . D e lpor te , H . 1 980 B ra ssempouy , l a G ro t te d u P ape , S ta t ion P reh i s to r ique . E d i t ion sP ica rd : P a r is .

Dama s ,D . 1 969 C ha rac ter i s t ic s o f C en t ra l E sk imo B and S t ruc tu re . I n C on t r ibu t ion s t o A n th ropo logy : B and S oc ie t ie s . D . D ama s , e d . N a t iona l Mu seum s o f C anada B u l le t in 2 28 :11614 1 . O t tawa : N a t iona l

D e L um ley , H . 1 976 C ad re C h rono log ique A b so lu , P a leomagne t i sme , C h rono log ie P a leon to log ique e t B o tan ique , E squ isse P a leoc l ima t ique , S equence s

Mu seum so fC anada .

C u l tu re l le s . I n L a P reh i s to i re L um ley , e d . p p . 5 -23 .

1 969 C on t r ibu t ion s t oA n th ropo logy : B and S o c ie t ie s . Na t iona l Mu seums o fC anada B u l le t in 2 28 . D . D ama s e d . O t tawa : N a t iona l Mu seum so fC anada .

F ranQa i se . H . d e P a r is : E d i t ion s d u

C .N .R .S . . D ema r s ,P .—Y . 1 982 U t i l i sa t ion d u S i lex a u P a leo l i th ique S uper ieu r .

D av id ,N .

P a r i s : e d i t ion sd u CNRS .

1 973 O n Uppe r P a laeo l i th ic S oc ie ty , E co logy , a nd T echno log ica l C hange : t h e N oa i l l ian C a se . I n T he E xp lana t ion o fC u l tu r e C hange . C . R en f r ew , e d .

1 989 L ' Ind ice L am ina i re d e l ' Ou t i l lage d an s l e P a leo l i th ique S upe r ieu re nP er igo rd . P a le ( ) 1 : 17-21 .

p p . 2 77-383 . C ambr idge : D uckwo r th .

D e S onnev i l le—Bo rdes , D . 1 960 L e P a leo l i th ique S upe r ieu r e n P er igo rd .

D e lpech ,F . 1 970 L 'Ab r i Magda len ien d u F lageo le t I : P a leon to log ie ( Bezenac—Do rdogne ) . B .S .P .F .

B o rdeaux : D e lma s .

6 7 (2 ) :494-499 . D up les sy ,J —C . ,e ta l . 1 976 P a leo c l ima to log ie d es T emp s Q ua te rna i re s a

1 983 L e s F aune s d u P a leo l i th ique S uper ieu r d u S ud O ues t d e l aF rance . C o l lec t ion s d u Q ua ter na i r e , n o .6 . P a r i s :

l ' A ide d e s Me thodes N uc lea i res . I n L a P reh is to i r e F rança ise . H . d e L um ley , e d . p p .352-361 . P ar i s :

C .N .R .S . .

C .N .R .S . . 1 986 L es R enne s d u G rand A b r i d e L auger ie—Hau te D u rham ,W .H . 1 981 O ver v iew : O p t ima l F o rag ing A na lys is i n H uman E co logy . I n H un te r—Ga the re r F o rag ing S t ra teg ie s . B . W in te rha lde r a nd E . Sm i th , e d .s p p .

e nD o rdogne . A r cheo log ia 1 3 :66-70 . 1 987

L ' env i ronnemen t A n ima l d es Magda len ien s .

A c te sd uC o l loque s u r l e Magda len ien . Mayence 1 987 . ERAUL 3 8 .

2 18-232 . C h icago : U n ive r s i ty o fC h icago P r ess .

8 5

E a r le ,T .K .

G au s sen ,J . 1 980 L e P a leo l i th ique S up e r ieu r d e P l e in A i r e n P e r igo rd . X IVe S upp lemen t aG a l l ia P reh i s to i r e .P a r i s : C .N .R .S . .

1 980 A Mode l o fS ub s i s tenc e C hange . I n Mode l l ing C hange i n P r eh i s to r ic S ub s i s t enc e E conom ie s . T .K . E a r le a nd C h r i s ten sen e d .s . N ew Y o rk : A cadem ic P r e s s . E n low ,D .H . ,a nd S .O . B rown 1 958 A C ompa ra t ive H i s to log ica l S t udy o f F o s s i l a nd R ec en t B on e T i ssue s . T exa s J ou rna l o f S c ien ce 1 0 (2 ) :1 87— 2 30 .

1 987 L eP eup lemen t Magda len ien D an sl aV a l lee d e L ' I s le ( s ec teu r Mu s s idan—S t .—A s t ie r ) . F o rme d e s H ab i ta t s e tC ho ix d e s Emp lacemen t s . I N L a F in d e s T emp s G lac ia i r e s e n E u rope . C o l loque s I n ter na t ionaux d u C .N .R .S .n o .271 .

E n r igh t ,J .T . 1 970 E co log ica l A spec t s o f E ndogenou s R hy thm ic i ty . A nnua l R ev iew o f E co logy a nd S y s tema t ic s 1 : 22 1-238 .

G i lbe r t ,F .F . ,a nd S .L . S to l t 1 970 V a r iab i l i ty i n A ge ing Ma in e Wh i te—Ta i led D ee r b y T oo th Wea r C ha rac t e r i s t ic s . J ou rna l o f W i ld l i f e Managemen t3 4 :532-535 .

F ranc i l lon ,H . 1 979 E tude E xpe r imen ta le d e s Ma rque s d e s C ro i s sanc e s u r l e s H ume ru s e tl e s F emu r s d e T r i ron s C re te s ( T r i tu r u s c r i s ta tu s c r i s ta tu s L au r en t i ) e n R e la t ion a vec l a D e te rm ina t ion d e l ' Age I n d iv idue l . A c ta Z oo log ica 6 0 :223-232 .

G l imche r ,M . J . 1 976 C ompo s i t ion , S t ruc tu r e , a nd O rgan i sa t ion o f B one a nd O the r M ine ra l iz ed T i s sue s a nd t h e Mechan i sm o f C a lc i f ica t ion . I n H andbook o f P hy s io logy—Endo c r ino logy V I I . Am . P hy s . S o c . , e d . B a l t imo re , MA : W i l l iam sa nd W i lk in sC o . . G o rdon ,B . 1 975 O f Men a nd H e rd s i nB a r r en land P r eh i s to ry . Me r cu ry S e r ie s n o .28 : A r chaeo log ica l S u r vey s o f C anada .

F ran c i l lon—V ie i l lo t H . , V . d e B u f f ren i l , J .C a s tane t , J . G e raud ie , F . J . Meun ie r ,J . —Y . S i r e , L .Z y lb e rbe rg , a nd A .d eR icq le s . 1 99 1 M ic ro s t ruc tu re a nd M in e ra l iza t ion o f V e r teb ra te S ke le ta l T i s sue s . I n S ke le ta l B iom ine ra l iza t ion : P a t ter n s , P roces se s a nd E vo lu t iona ry T r end s . J o seph G . C a r te r , e d . p p . 4 71-530 .N ew Y o rk : V an No s t rand R e inho ld .

1 981 Man—Env i ronmen tR e la t ion sh ip s i nB a r r en land P r eh is to ry . Mu sk—Ox 2 8 :1-19 . 1 988 O f Men a nd R e indee r i nF r ench Magda len ian P r eh is to ry . O x fo rd : B r i t i sh A r chaeo log ica l R epo r t s , I n te rna t iona l s e r ie sn o .390 .

F reeman ,L .G . 1 973 T he S i gn if icance o f Mamma l ian F auna s F rom P a leo l i th ic O c cupa t ion s o f C an tab r ian S pa in . Ame r ican A n t iqu i ty 3 8 (1) :3-44 .

G ran t ,A . 1 982 T he u se o fT oo th Wea ra s aG u ide t ot h e a ge o f D ome s t ic U ngu la te s . I n A ge ing a nd S ex ing A n ima l B on es F rom A r chaeo log ica l S i te s . W i l son , G r ig son , a nd P ayne , e d .s p p . 9 1107 . B r i t i sh A r chaeo log ica l R epo r t s .

F r i son ,G .C . 1 978 P reh i s to r ic H un te r s o f t h e H igh P la in s . A cadem ic P re s s :N ew Y o rk .

G ray son ,D . 1 988 D ange rC ave ,L a s tS uppe rC ave ,H ang ing R ock S he l te r . A n th ropo log ica l P ape r s o f t h e Mu seum o f

G amb le ,C .S . 1 983 C u l tu r ea nd S oc ie ty i nt h eU pp e rP a laeo l i th ic i n E u rop e . ln H un te r—Ga the r e r E conomy i nP r eh is to ry . G . B a i ley , e d . p p . 2 01-2 11 . C amb r idge : C amb r idge U n ive r s i ty P r e s s .

N a tu ra l

H is to ry ,n o .6 6 .

1 986 T he P a laeo l i th ic S e t t lemen t o f E u rop e . C amb . U n iv .P re ss :C amb r idge .

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8 6

W ie sbaden : D r . L udw ig V e r lag .

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H ayden , B . 1 980 C ommen t o n C onkey : P reh i s to r ic Hun te r— G a the re r A gg r ega t ion S i te s . C u r r en t A n th ropo logy 2 1(5 ) :623-624 .

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H ef f ley , S . 1 981 T he R e la t ion sh ip B e tween No r ther n A thapa skan S e t t lemen t P a t te rn s a nd R e sou rc e D is t r ibu t ion : a n App l ica t ion o f H or n 's Mode l . I n H un te r—Ga the re r F o rag ing S t ra teg ie s . B . W in te rha lde r a nd E . Sm i th , e d .s p p . 1 26147 . C h icago : U n ive r s i ty o fC h icago P re ss .

R ev iew o fG ame B io logy 1 0 (3 ) . G uen the r ,M .G . 1 980 C ommen t o n C onkey : P reh is to r ic H un te r— G a the re r A gg rega t ion S i te s . C u r ren t A n th ropo logy 2 1 (5 ) :6 22 . G u i l l ien ,Y . 1 977 C ro i s sance d u R enne e tC ha s se P a leo l i th ique . I n App ro che E co log ique d e L 'Homme F o ss i le . H . L av i l le a nd J .R enau l t—M ikov sky , e d .s p p . 1 37-139 . P a r is : A .F .E .Q . .

H e lm 1 968 T he N a tu re o f D ogr i b T e r r i tor ia l G roup s . I N P rob lem s i n t h e S tudy o f H un te r—Ga ther e r s . R . L ee a nd D evo re e d .s p p .118-125 . C h icago : A ld ine .

1 983 C ro i ssance d u R enne e tC ha sse P a leo l i th ique . P a leoc l ima t s . B u l le t in d e L ' In s t i tu t d e G eo log ie d u B a ss in D 'Aqu i ta ine 3 4 .

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1 987 A u r ignac ian a nd G rave t t ian S e t t lemen tP a t tern s i nC en t ra l E u rop e . I nP le i s to cene o ft h e O ld Wo r ld . 0 .

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8 7

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1 968 T he M ig ra to ry B a r r en—g round C a r ibou o f C anada . C anad ia n W i ld l i f e S e r v ice Monog raph n o .3 , O t tawa .

1 987 L a te P l e i s tocene R e f ug ia i nE u rop e . I nT h e P l e i s to cene o f t h e O ld Wo r ld . 0 . S o f f e r , e d . p p . 3 17-332 .N ew Y o rk :P lenum P r e ss .

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K le in ,R . ,e ta l . 1 981 T he u s e o f D en ta l C rown H e igh t s f o r R econ s t ru c t ing a ge P ro f i le s o f r e d D ee r a nd S im i la r S p ec ie s i n A r cha eo log ica l S amp le s . J ou rna l o f A r chaeo log ica lS c ien ce 8 : 1-31 .

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1 980 L aye r s i nt h e H a rd T i s sue s o f Mamma l s a s a R eco rd o fG row th R hy thm s o fI n d iv idua ls . R epo r t o f t h e I n te rna t iona l Wha l ing C omm i s s ion . C amb r idge . S p ec ia lI s sue 3 : 89-84 . 1 988 R eco rd ing S t ruc tu r e s o f Mamma l s i n Z oo log ica l I n ve s t iga t ion s . Mo scow : N auka . ( I n R u s s ian ) .

1 983 B io logy , B ehav iou r a nd B o r row ing : aC r i t ica l E xam ina t ion o f O p t ima l F o rag ing T h eo r y . I n A r chaeo log ica l H amme r s a nd T heo r ie s . J .A . Moo re a nd A .S .Keen e , e d . p p . 1 37155 . N ew Y o rk : A cadem ic P r e ss .

K leveza l ,G .A . a nd S .E . K le inenb e rg 1 967 A ge D e te rm ina t ion o f Mamma l s F rom A nnua l L aye r s i n T ee th a nd B one s . A cademy o f S c ience s U .S .S .R . T ran s la t ion D ep t . o f t h e I n ter io r a nd N a t iona l S c ien c e F ounda t ion , U .S . D ep t . o f C omme r ce , C lea r inghou se f o r F ed era l S c ien t if ic a nd T echn ica lI n fo rma t ion ,S p r ing f ie ld .

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8 8

1 984 T e tracyc l ine L ab e l l ing a s aMe thod o f F ie ld S t ud ie so fI nd iv idua l G row th a nd P opu la t ion S t ruc tu re i n R oden t s &L ynx .

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K oh le r—Ro l lef son , I 1 989 C hange s i nG oa t E xp lo i ta t ion a t' AM G haza l B e tween t h e E a r ly a nd L a te N eo l i th ic : a Me t r ica l A na ly s i s .P a leo r ien t 1 5( 1 ) :4145 .

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1 979 T he ! Kung S an : Men Women a nd Wo rk i na F o rag ing S o c ie ty . C ambr idge : C amb r idge U n ive r s i ty P res s .

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L iebe rman ,D .E . 1 993 Mob i l i ty a nd S t ra in : t h e B io logy o f C emen togene s i sa nd i t sA pp l ica t ion t ot h e E vo lu t ion o f H un te r—Ga the r e r S ea sona l Mob i l i ty D u r ing t h e L a te Q ua ter na ry i nt h e S ou ther n L evan t . P h .d d i sse r ta t ion , H a r va rd U n ive r s i ty .J anua ry , 1 993 .

McKennan ,R .A . 1 969 A thapa skan G roup ing sa nd S o c ia lO rgan iza t ion i nC en t ra l A la ska . I n C on t r ibu t ion s t oA n th ropo logy : B and S oc ie t ie s . D . D ama s , e d . p p . 9 30-115 . O t tawa :Q ueen ' sP r in te r .

L iebe rman ,D .E . ,T .W . D eacon , a nd R . Meadow 1 990 C ompu te rI mage E nhancemen t a nd A na ly s i s o f C emen tum I n c remen t s a sA pp l ied t oT ee th o fG aze I la G aze I la . J ou rna l o f A r chaeo log ica l S c ienc e 1 7 :5195 33 .

Me l la r s ,P . 1 985 T he e co log ica lb a s i so fs o c ia lc omp lex i ty i nt h e

9 0

Uppe r P a laeo l i th ic o f S ou thwe s tern F rance . I n P reh i s to r ic H un te r—Ga the re r s , t h e Eme rgence o f C u l tu ra l C omp lex i ty . T . P r ice a nd J .B rown , e d .s p p . 2 71-297 .N ew

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Y o rk : A cadem ic P ress . Mye r s ,F . 1 986 P in tup i C oun try , P in tup i S e lf : S en t imen t , P lace a nd P o l i t ic s a mong We s te rn D e ser t A bo r ig ine s . Wa sh ing ton : Sm i th son ian I n s t i tu t ion P re ss .

Menn ick ,P . n . d . O b ser va t ion s o f t h e W i ld H o r se . Ho r se N ew s ( 4 ) :4-46 . Me tca lf e , D . , a nd K .T . J ones 1 988 A R econ s ide ra t ion o f A n ima l B ody—Pa r t U t i l i ty I n dece s . Ame r ican A n t iqu i ty 5 3 (3 ) :486-504 .

My r ick , A .C . 1 980 S ome App roaches t ot h e C a l l ib ra t ion o f a ge i n O don toce te s U s ing L aye red H a rd T is sue s . I n R epo r t o f t h e I n ter na t iona l Wha l ing C omm iss ion . C ambr idge . S pec ia l I s sue n o .3 :95-97 . A .J . My r ick a nd W .F . P e r r in ,e d .s .

M i l le r ,F . 1 974 B io logy o f t h e K am inu r iak P opu la t ion o f B a r ren G round C a r ibou . C anad ian W i ld l if e S er v ice R epo r t s . 3 1(Pa r tI ) .O t tawa .

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M i tche l l ,B . 1 969 G row th L aye r s i n D en ta l C emen t f o r D e te rm in ing t h e a ge o fr ed D ee r ( Cervus E laphu s L . ) . J our na lo fA n ima l E co logy . 3 6 (2 ) :279-293 .

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S ea sona l i ty

S t ud ie s .

I n A dvance s

i n

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P aque reau , M .M . 1 970 F lo res e t C l ima t s P a leo l i th ique s D an s l e s ud—Oue s t d e l a F rance . B u l le t in d e l a S o c ie te P r eh is to r ique F ran p ise . 6 7 :489-493 .

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9 1

1 981 C ompa ra i son d e D i f f e r en te s Mi thode s d e D e te rm ina t ion d e L ' äge I n d iv idue l C hez l e V i son ( Mu s te la V ison S ch r e ib e r ) . C anad ian J ou rna l o f Z oo logy .5 9 (2 ) :202-211 .

S u r vey o fC anada . P r i t cha rd ,J . J . 1 956 G ene ra l H i s to logy o f B one . I n T he H i s to ch em i s t r y a nd P hy s io logy o fB one . G .H . B ou rne e d .pp .1-20 .N ew Y o rk : A cadem ic P r e ss .

P a sca l ,M . ,a nd G . Kova l c s 1 983 L a D e te rm ina t ion d e L 'äge I n d iv idue l C h ez l e L iev re E u rop een p a r l a T echn ique S que le t to ch rono log ique . R evue E co log ique ( Te r r e V ie ) .N o .37 .

Q ue r eJ . P . ,a nd M .P a sca l 1 983 C ompa ra i son d e P lu s ieu r s Me thode s d e l a D e te rm ina t ion d e l ' äge I n d iv idue l c h ez l eC e r fE laphe ( Ce rvu s e l aphu s L . ) . A rm ie s d e S c iences N a tu r e I le s , Z oo log ie e t B io log ie A n ima le 5 (4 ) :2352 52 .

P eabody ,F .E . 1 958 A K an sa s D rough t R eco rded i nG row th Z on e s o ft h eB u l l snake .C ope ia 2 : 99104 .

R ayna l ,J . P . 1 975 L a G ro t te D 'E sc lauzu r ( c ommune d e L i s sac , C o r r eze ) , N ouveau S i te S o lu t r een . B u l le t in d e l a S oc ie te P r eh i s to r ique F ran ça i se .7 2 (2 ) :50-53 .

1 96 1 A nnua l G row th Z one s i n L iv ing a nd F o s s i l V e r teb ra te s .J ou rna lo f Mo rpho logy . 1 08 :11-62 . P enck ,A . ,a nd E .B rückn e r 1 909 D ie A lpen i nE i sze i ta l te r .L e ipz ig .

1 977 L e Q ua tema i r e e n C o r r eze : E ta t s d e s R eche r ch e s . B u l le t in d e l a S oc ie te S c ien t if ique , H i s to r iqu ee tA r cheo log ique d el aC o r rez e .9 9 :47-71 .

P e te r son ,G .M .e ta l . 1 979 T he C on t inen ta l R eco rd o f E nv i ronmen ta l C ond i t ion s a t 1 8 ,000 y r . B .P . : a n I n i t ia l E va lua t ion . Q ua ter na ry R e sea r ch . 1 2 :47-82 .

R e ime r ,E .a nd 0 . No rdby 1 968 R e la t ion sh ip B e tween a ge a nd T oo th C emen tum L aye r s i nN o rweg ian R e indee r . J ou rna l o f W i ld l if e Managemen t3 2 (4 ) :957-961 .

P e t i t ,M . 1 939 A na tom ie d e s Mo la i r e sd e s E qu ide s : C h eva le t Me . I mp r ime r ie T ou lou sa ine ,L ion e tF i l s :T ou lou se .

R emy ,J .A . 1 974 O b se r va t ion ss u rl e sR eman iemen t sS t ruc tu raux P o s t—Mo r t em D an sl e sD en t sd e Mamm i f e re sF o s s i le s P rovenan t d e s P ho spho r e s d u Q ue r cy . P a leove r teb ra ta 6 : 163176 .

P ey rony ,D . 1 950 N o te s s u r Q ue lque s P e t i t s G i semen t s P reh i s to r ique s . V I . A b r i d u C ap B lan c . B u l le t in d e l a S o c ie te H i s to r ique e tA r cheo log ique d u P e r igo rd 7 7 :63-64 .

R iche s ,D . 1 981 T he O b l iga t ion t o G ive— a n I n te rac t iona l S ke tch . I nT he S t ruc tu r e o fF o lk Mode l s . L . H o ly a nd M . S t uch l ik , e d . s p p .209-231 . L ondon : A cadem ic P r e s s .

P ey rony ,D . ,a nd E .P ey rony 1 938 L auge r ie—Hau te , p r e s d e s E yz ie s ( Do rdogne ) . A rch ive s d e l ' I n s t i tu t d e P a leon to log ie H uma ine , Memo i r e 1 9 .

R icq les ,A .d e 1 975 R e che r che s P a leoh i s to log ique s s u rl e so s l o ng s d e s T e t rapode s . V I I— su r l a C la s s i f ica t ion , l a S i gn i f ica t ion F on c t ione l le e tL 'h i s to i r e . A nn& P a leon to log ique .6 1 :51129 .

P ike—Tay ,A . 1 991 R ed D ee r H un t ing i nt h e U pp e rP a laeo l i th ic o f S ou thwes t F ran ce : A S t udy i n S ea sona l i ty . O x fo rd : B r i t i sh A r chaeo log ica l R epo r t s , I n t er n a t iona l S e r ie s 5 69 . P ra t t ,F . 1 976 L e sP e r i ssoda c ty le s : E qu ide s . I nL aP r eh i s to i r e F rança i se .p p .409-4 15 .P a r i s : C ah ie r sd uC .N .R .S . .

1 979 Q ue lque s R ema rque s s u r l ' H i s to i r e E vo lu t ive d e s T i s su s S que le t t ique s c h ez l e s V e r teb r e s e t p l u s P a r t icu l ie r emen t c hez l e s T e t rapod e s . A nnie B io log ique 1 8 :1-35 .

P res ley ,S .L . 1 987 D e term ina t ion o fS i te S ea sona l i ty U s ing T oo th A nnu lu s T echn ique . I n te rna l R epo r t . A r chaeo log ica l

R icq le s , A . d e , F . J . Meun ie r , J . C a s tane t , a nd H . F ranc i l lon—V ie i l lo t 1 992 C ompa ra t ive M ic ro s t ruc tu r e o fB one . I nB one

9 2

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1 990 O n t h e O s teo log ica l E v idence f o r P a laeo l i th ic D ome s t ica t ion : B a rk ing u p t h e Wrong T ree . C u r ren t A n th ropo logy . 3 1(5 ) :543-547 .

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S ah l in s , M . 1 972 S tone A ge E conom ic s . N ew Y o rk : A ld ine . S ave I le ,J . ,a nd O .W . B ea t t ie 1 983 A na ly s is o f D en ta l A nnu l i i n Muskoxen ( Ov ibo s Mo scha tu s ) a sa n a i d t ot h e D e term ina t ion o f A r chaeo log ica l S i te S ea sona l i ty . C anad ian J ou rna l o f A r chaeo logy . 3 ( 1) :123129 .

R oge r sE .S . 1 969 B and O rgan isa t ion Among t h e I n d ian s o f E a s ter n S uba rc t ic C anada . I n C on t r ibu t ion s t o A n th ropo logy : B and S o c ie t ie s . D . D ama s , e d . p p . 2 1-55 . O t tawa :Q ueen 's P r es s .

S axon ,A . , a nd C .F . H igham 1 968 I d en t if ica t ion o fG row th R ing s i nt h e S econda ry D en ta l C emen tum o f O v is A r ie s L . . N a tu re

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S axon ,A . , a nd C .F . H igham 1 969 A n ew R esea rch Me thod f o r E conom ic P r eh is to r ian s . Ame r ican A n t iqu i ty 3 4 (3 ) :303-311 .

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S ha rp ,H .S . 1 978 C ompa ra t ive E tho logy o f t h e Wo lf a nd C h ip ewyan . R ep r in ted F rom : Wo l f a nd Man . p p .55-79 . A cadem ic P r e s s :N ew Y o rk .

S p iess ,A .E . 1 990 D ee r T oo th S e c t ion ing , E r up t ion a nd S ea sona l i ty o fD ee rH un t ing i nP reh i s to r ic Ma ine . Man i nt h e No r th ea s t . 3 9 :29-44 .

S i lb e rbaue r ,G eo rg eB . 1 981 H un te r a nd H ab i ta t i nt h e C en t ra l K a laha r i D e se r t .C amb r idge :C amb r idg eU n ive r s i ty P r e s s .

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1 979 R e indee r a nd C a r ibou H un te r s . N ew Y o rk : A cadem ic P r e s s . S p inage ,C .A . 1 976 A ge D e te rm ina t ion i nF ema le G ran t ' sG a ze l le . E a s tA f r ican W i ld l if eJ ou rna l . 1 4 :121134 .

S imp son ,G .G . 1 951 H o r se s .O x fo rd U n ive r s i ty P r e s s .

S ta l l ib ra s s ,S . 1 982 T h eu s eo fC emen t L aye r s f o rA b so lu te A ge ing o f Mamma l ian T ee th : aS e lec t ive R ev iew o f t h e L i te ra tu r e , W i th S ugge s t ion s f o r F u r the r S t ud ie s . I n A ge ing a nd S ex ing A n ima l B one s F rom A r chaeo log ica l S i te s . B . W i lson , C . G r ig son , a nd S . P ayne , e d .s p p . 1 09-126 . L ondon : B r i t i sh A r chaeo log ica lR epo r t s , 1 09 .

S lobod in ,R . 1 969 L ead e r sh ip a nd P a r t ic ipa t ion i n aK u tch in T rapp ing P a r ty . I n C on t r ibu t ion s i n A n th ropo logy : B and S oc ie t ie s . D . D ama s ,e d . p p .56-92 . O t tawa : Q ueen ' sP re ss . S chm ide r ,B . 1 97 1 L e s I n du s t r ie s L i th ique d u P a leo l i th ique S up e r ieu r e n I l e—de F ran ce .V le s upp lemen t t oG a l l ia P reh i s to i r e . P a r i s :e d i t ion sd u CNRS .

S t ewa rd ,J .H . 1 938 B a s in—P la teau A bo r ig ina l S oc iopo l i t ica l G roup s . Wa sh ing ton : U .S . G over n emen t P r in t ing O f f ice .

Sm i th ,P .E .L . 1 965 S ome S o lu t r ean P rob lem s a nd S ugge s t ion s f o r F u r the rR e sea r ch . I N M i sc e l len ia e nH omage a lA ba te H en r i B r eu i l .T ome 1 . B a rce lona .

1 977 T h eC oncep t a nd Me thod s o fC u l tu ra l E co logy I N E vo lu t ion a nd E co logy : E s say s o n S o c ia l T ran sfo rma t ion . J .C . S t ewa rd a nd R .F . Mu rphy e d .s . U n ive r s i ty o fI l ino i sP r e ss .p p .43-57 .

1 966 L eS o lu t r een e nF rance . B o rdeaux : D e lma s . Sm i th ,E .A . 1 981 T he A pp l ica t ion o fO p t ima lF o rag ing T heo ry t o t h e A na ly s is o f H un te r—Ga the r e r G roup S i ze . I n H un te r—Ga the r e r F o rag ing S t ra teg ie s . B . W in te rha lde r a nd E . S m i th e d .s p p . 3 6-65 . C h icago : U n ive r s i ty o f C h icago P re s s .

S t o t t ,G .G . ,R .F . S i s , a nd B .M . L evy 1 982 C emen ta l A nnu la t ion a s a n a ge C r i te r ion i n F o ren s ic D en t is t ry . J ou rna l o f D en ta l R e sea r ch . 6 1 (6 ) :814-8 17 . S t rau s ,L .G . 1 979 C an tab r ia a nd V a scongada s , 2 1 ,00017 ,000 B P : T owa rd s aS o lu t rean S e t t lemen tP a t te rn . Mun ibe S oc iedad d e C ienc ia s N a tu ra le s A razad i , S an S eba s t ian ,A no XX I# 3-4 .

S o f f e r ,0 . 1 987 T he P l e i s toc en e o ft h e O ld Wo r ld . N ew Y o rk : P lenum P re ss .

1 980

1 987b U ppe rP a laeo l i th ic c onnub ia , r e f ug ia , a nd t h e a r chaeo log ica l r e co rd f r om E a s ter n E u rop e . I N T he P le i s to cene o f t h e O ld Wo r ld . 0 . S o f f e r , e d . p p . 3 333 46 .N ew Y o rk : P l enum P r e s s .

C ommen t o n C onkey : P r eh i s to r ic H un t e r—

G a the re r A gg r ega t ion S i te s . C u r ren t A n th ropo logy 2 1 (5 ) :624-625 .

9 4

1 983

T e rm ina l P le is tocene F auna l E xp lo i ta t ion i n

V i l la ,P . 1 983 T e r ra Ama ta a nd t h e M idd le P le i s to cene A r chaeo log ica l R eco rd o f S ou ther n F rance . B e rke ley : U n ive r s i ty o fC a l ifor n ia P res s .

C an tab r ia a nd G a scony . I nA n ima ls a nd A r chaeo logy . J .C lu t ton—Brock a nd C . G r ig son , e d . p p . 2 09-224 . O xfo rd . : B r i t i sh A r chaeo log ica l R epor t s . I n te rna t iona l S er ie s 1 63 .

V i ta—F in z i ,C . 1 978 A r chaeo log ica l S i te s a nd T he i r S e t t ing s . L ondon : T hame s a nd H ud son .

1 987 H un t ing i nL a te Uppe r P a laeo l i th ic We s ter n E u rope . I n T he E vo lu t ion o f H uman H un t ing . M .H . N i teck i a nd D . V . N i teck i ,e d . p p . 1 47-175 . N ew Y o rk :P l enum P ub l i sh ing .

V i ta—F inz i ,C . , a nd E .H igg s 1 970 P reh is to r ic E conomy i nt h e Moun tC a rme lA rea o fP a les t ine , S i te C a tchmen t A na lys is . I N P roceed ing s

S t rau s ,L .G . a nd C .H e l le r 1 988 E xp lo ra t ion s o f t h e Tw i l igh t Z one : t h e E a r ly U ppe r P a leo l i th ic o f V a sco—Can tab r ian S pa in a nd G a scony . h i T he E a r ly U pp e r P a leo l i th ic . J . F . H o f fe cke r a nd C .A . Wo lf , e d . B r i t i sh A r chaeo log ica l R epo r t s ,I n te rna t iona lS er ie s4 37 .

o ft h eP reh is to r ic S o c ie ty . 3 6 :1-37 . Wa r ing , G .H . 1 983 Ho r se B ehav io r . P a rk R idge , N ew J e r sey : Noye sP ub l ica t ion s .

S t rau s ,L .G . a nd A . S p ies s 1 985 L e Magda len ien F ina ld e l ' Ab r i D ufau re ( So rde d e L 'abbaye , L ande s ) : u n Ap e r çu d e l aC h rono log ie e t d e l aS a i son d 'Hab i ta t ion H uma ine . B u l le t in d e l a S oc ie te P reh is to r ique d el ' A r iege . XXXX :169-184 .

Wa r r en ,J .W . 1 963 G row th Z one s i nt h e S ke le ton o f R ecen t a nd F o ss i l V er teb ra te s . P h .D . T he s is . M ich igan : U n ive r s i ty M ic ro f i lm s .

A nn

A rbo r ,

Wa shbu rn , A .L . 1 973 P er ig lac ia l P roc esses a nd E nv i ronmen t s .

S tu rdy ,D . 1 972 T he E xp lo i ta t ion P a t te rn s o f a Moder n R e indee r E conomy i n Wes t G reen land . I n P ap e r s i n E conom ic P reh i s to ry . E .S . H igg s , e d . p p . 1 6 1168 . C amb r idge : C amb r idge U n ive r s i ty P r es s .

L ondon : E dwa rd A rno ld p ub l ishe r s . 1 985

P er ig lac ia l P rob lem s . I n F ie ld a nd T heo ry .

M .A . C hu rch a nd 0 . S laymake r , e d .s p p . 1 66-197 . U n ive r s i ty o fB r i t ish C o lumb ia P r es s .

1 975 S ome R e indee r E conom ies i n P reh is to r ic E u rope . I iP a laeo economy . E . H igg s , e d . p p . 5-95 . C amb r idge : C amb r idge Un ive r s i ty P res s .

Wa t t s ,P .a nd D .E . G a sk in 1 989 A C ompa r i son o f a ge D e term ina t ion T echn iques f o r t h e H a rbou r P o rpo i se , P ho co ena P hocoena

T ex ie r , J . —P . , J . —P . R ayna l , H . L av i l le , M .—M . P aque reau , F .P ra t ,A . D eb ena th ,a nd F .D e lpech . 1 983 H is to i r e P a leo c l ima t ique d e L 'Aqu i ta ine d u

L .

C anad ian

J ou rna l

Z oo logy

6 7 :1832-1836 .

P l e i s tocene A nc ien a u D er n ie r I n te rg lac ia i re . B u l le t in

We l l s , S .M . , a nd B .v on G o ld schm id t—Ro th sch i ld 1 979 S o c ia l B ehav iou r a nd R e la t ion sh ip s i n aH e rd

d e l ' I n s t i tu tG eo log ique d u B a s s in D 'Aqu i ta ine 3 4 :207 .

o fC ama rgue H o r ses .Z .T ierp sycho log ie .4 9 :363-380 .

T r ica r t ,J . 1 967 G eomo rpho logy o f C o ld E nv i ronmen t s . P a r is :

We l sh , D .A . 1 975 P opu la t ion , B ehav iou ra l a nd G ra z ing E co logy

S oc ie te d 'Ed i t ion sd 'En se ignemen tS uper ieu r s .

o f t h e H o r se s U npub l ished

T u rnbu l l ,P .F . 1 986 Mea su remen t s o f E quu s h em ionu s f r om

U n ive r s i ty .

P a legaw ra C ave ( Za r z ian , I r aq ) . I N E qu id s i n t h e A nc ien t Wo r ld . R .H . Meadow a nd H .—P . U erpmann

Wh i te , R . 1 985 U ppe rP a laeo l i th ic L and—use i nt h eP er igo rd . A T opog raph ic A pp roach t oS ub s is tence . L ondon : B r i t ish

e d .s p p .319-365 .

o f S ab le I s land , N ova S co t ia . P h .D . D i sser ta t ion , D a lhou s ie

A rchaeo log ica lR epo r t s , I n ter na t iona l s e r ie s 2 53 . V igneaux . M . 1 975 A qu i ta ine O cc iden ta le . P a r i s : Ma sson .

1 987

G l imp se s o f L ong—Term S h if t s i n L a te

P a laeo l i th ic L and u se i nt h eP er igo rd . I nP l e i s tocene o f

9 5

t h e O ld Wo r ld . 0 . S o f f e r , e d . p p . 2 63-277 . N ew Y o rk :P l enum P r es s . 1 990 H u sband ry a nd H e rd C on t ro l i nt h e U pp e r P a laeo l i th ic : a C r i t ica l R ev iew o f t h e E v iden ce . C u r r en tA n th ropo logy 3 0 :609-6 16 . 1 990b P roduc t ion C omp lex i ty a nd S t anda rd i sa t ion i n e a r ly A u r ignac ian B ead a nd P endan t Manu fa c tu r e : E vo lu t iona ry I mp l ica t ion s . I N T h e O r ig in s a nd D i sp e r sa l o f Mod er n H uman s : B ehav iou ra l a nd B io log ica l P e r spec t ive s . C . S t r inge r a nd P . Me l la r se d .s . E d inbu rgh : E d inbu rgh U n ive r s i ty P r e s s . W in te rha lde r ,B . 1 981 Op t ima l F o rag ing S t ra teg ie s a nd H un te r—Ga the r e r R e sea r ch i n A n th ropo logy : T h eo ry a nd Mode ls . I nH un te r—Ga the r e r F o rag ing S t ra t eg ie s . B . W in t e rha lde r a nd E . Sm i th , e d . p p .13-35 . C h icago : U n ive r s i ty o fC h icago P r e ss . 1 981b F o rag ing S t ra teg ie s i nt h e B o r ea l F o r e s t : a n A na ly s i s o f C r ee H un t ing a nd G a the r ing . I n H un te r—Ga the r e F o rag ing S t ra teg ie s . B . W in te rha lde r a nd E . S m i th , e d . p p . 6 6-98 . C h icago : U n ive r s i ty o f C h icago P re ss . 1 986 D ie t C ho ic e , R isk a nd F ood S ha r ing i n a S tocha s t ic E nv i ronmen t . J ou rna l o f A n th ropo log ica l A r chaeo logy 5 : 369-392 . Wob s t ,H .M . 1 977 S t y l is t ic B ehav iou r a nd I n fo rma t ion E xchange . I NP ape r sf o rt h e D i rec to r : R esea r ch E s say s i nH onou r o f J ame s B . G r if f in . C .E . C le land e d . p p .317-342 . M ich igan : A n th ropo log ica l P ape r s , Mu seum o f A n th ropo logy , U n ive r s i ty o fM ich igan . Y e sne r ,D .R . 1 981 A rchaeo log ica l App l ica t ion s o f O p t ima l F o rag ing T heo ry : H a r ve s t S t ra teg ie s o f A leu t H un te r—Ga the r e r s . I nH un te r—Ga th e r e r F o rag ing S t ra teg ie s . B . W in te rha lde r a nd E . S m i th , e d . p p . 1 48-170 .C h icago : U n ive r s i ty o fC h icago P r e s s .

9 6

A PPENDIX A M easurements: f ossil

I dent B 01 B 02 B 03 B 04 B 05 B 06 B 07 B 08 B 09 B 10 B ll B 12 B 13 B 14 B 15 B 16 B 17 B 18 B 19 B 20 B 21 B 22 B 23 B 24 B 25 B 26 B 27 B 28 B 29 B 30 B 31 B 32 B 33 B 34 B 35 B 36 B 37 B 38 B 39

i n i n i n i n i n i n i n i n i n i n i n i n i n i n i n i n i n s u s u s u s u s u s u s u s u s u s u s u s u s u s u s u s u s u s u s u s u i n i n

t eeth.

H eight

M eas.1

M eas.2

Meas.3

M eas.4

M eas.5

6 .85

2 .85 2 .52 2 .80

1 .65 1 .42 1 .49 1 .50 1 .43 1 .32 1 .45 1 .40 1 .30 1 .61 1 .51 1 .40 1 .52 1 .61 1 .55 1 .60 1 .53 2 .60 2 .50

. 25 . 16 . 26 . 26 . 35 . 25 . 43 . 26 . 31 . 29 . 25 . 22 . 24 . 53 . 57 . 65 . 49 1 .35 1 .30 1 .35 1 .45 1 .25 1 .40 1 .20 1 .40

1 .20 1 .06 1 .16 1 .20 1 .25 1 .06 1 .25 1 .24 1 .22 1 .31 1 .20 1 .11 1 .20 1 .40 1 .64 1 .66 1 .37

1 .49 1 .50 1 .45 1 .42 1 .30 1 .41 1 .30 1 .46 1 .39 1 .32 1 .49 1 .49

3 .06 3 .22 3 .15 2 .70 3 .21 2 .84 2 .72

4 .20 3 .30 4 .10 3 .70 4 .70 6 .50 5 .15 5 .70 6 .90 5 .50 5 .80 5 .25 2 .20 4 .00 3 .25 4 .40 4 .55

2 .80 2 .94 2 .90 3 .06 2 .45 2 .40 2 .48 2 .60 2 .55 2 .50 2 .55 2 .50 2 .55 2 .50 2 .60 2 .50 2 .35 2 .45 2 .35 2 .55 2 .50

4 .00 4 .30

3 .65 4 .00

6 .16

2 .80 2 .70 2 .60 2 .60 2 .40 2 .70 2 .60 2 .70 2 .55 2 .70 2 .70 2 .50 2 .40 2 .40 1 .55

9 7

1 .15 1 .40 1 .35 1 .20 1 .35 1 .30 1 .30 1 .25 1 .25 . 85 . 90 . 50

1 .74 1 .59 1 .80 1 .60

0 .00

. 85 . 85

1 .45

APPENDIX A Measurements: f ossil

I dent B 40 s u B 41 s u B 42 i n B 43 i n B 44 i n B 45 i n B 46 i n B 47 i n B 48 s u B 49 su B 50 s u B 51 s u B 52 s u B 53 su B 54 s u B 55 s u B 56 s u B 57 i n B 58 i n B 59 i n B 60 i n B 61 i n B 62 i n B 63 i n B 64 i n B 65 i n B 66 i n B 67 i n B 68 i n B 69 i n B 70 i n B 71 i n B 72 i n B 73 i n B 75 i n B 76 i n B 77 i n B 78 s u B 79 i n B 80 i n

H eight

Meas.1

M eas.2

M eas.4

M eas.5

1 .20 1 .20 1 .20 . 50 . 70 1 .00

1 .35 1 .45 1 .25 1 .50 1 .50 1 .10

2 .30 2 .50 2 .50 2 .40 2 .60

. 20 . 20 . 30 . 11 . 20 . 30 1 .40 1 .50 1 .45 1 .45 1 .40 1 .40 1 .30

3 .00 3 .10 2 .70 2 .70 2 .60 2 .80

2 .40 1 .30 1 .20 1 .40 1 .20 1 .50

. 20 . 20 . 30 . 10 . 20

1 .35 1 .15 1 .10 . 90 1 .20

1 .25 1 .50 1 .50 1 .50 1 .50

2 .55 2 .65 3 .10 2 .80 2 .90 3 .00 2 .90 2 . 90 2 .70 2 .80 3 .00 2 .75 2 .75 2 .75 2 .60 2 .80 2 .60

1 .40 1 .30 1 .40 1 .50 1 .60 1 .50 1 .70 1 .60 1 .60

. 10 . 15 . 30 . 30 . 30 . 50 . 50 . 50 . 40

1 .00 . 90 1 .25 1 .20 1 .25 1 .30 1 .30 1 .50 1 .35

1 .50 1 .30 1 .40 1 .40 2 .70 1 .60 1 .40

. 60 . 10 . 15 . 15 1 .50 . 40 . 20

1 .40 . 90 1 .20 1 .20

1 .40 1 .50 1 .30 1 .50 1 .40 1 .40 1 .60 1 .70 1 .80 1 .60 1 .60

4 .50

2 .50 2 .25

0 .00 0 .00

7 .80

4 .90

5 .70 7 .60 5 .80 7 .80

t eeth.

M eas.3 . 90

3 .00 2 .40 2 .50 2 .70 2 .70 2 .70 2 .90 2 .65 2 .80 2 .90 2 .60

1 .55 1 .30 1 .40 1 .40 1 .35 1 .50 1 .45 2 .40

9 8

1 .10 1 .10

1 .70 1 .70 1 .50 1 .50

APPENDIX A M easurements: f ossil

I dent B 81 B 82 B 83 B 84 B 85 B 86 B 87 B 88 B 89 B 90 B 91 B 92 B 93 B 94 B 95 B 96 B 97 B 98 B 99 B 100 B 101 B 102

H eight i n i n i n i n i n i n i n i n i n i n i n s u s u s u i n i n i n i n i n i n i n i n

8 .10 7 .85 3 .10 6 .50 6 .00 4 .15

4 .20

t eeth.

M eas.1

M eas.2

M eas.3

M eas.4

M eas.5

2 .50 3 .00

1 .25 1 .50 1 .40 1 .30 1 .50 1 .55 1 .50 1 .35 1 .45 1 .25 1 .30 2 .85

. 20 . 40 . 10 . 20 . 40 . 35 . 30 . 35 . 20 . 25 . 35 1 .50

. 90 1 .20 1 .10 . 85 1 .20 1 .30 1 .00 1 .20 . 95 . 90 1 .30

1 .30 1 .50 1 .30 1 .50 1 .70 1 .70 1 .65 1 .15 1 .60 1 .40 1 .15

1 .10 1 .30

1 .55 1 .35 . 90 1 .25 1 .50 1 .45 1 .55 1 .23

2 .40 2 .60 2 .70 2 .50 3 .10

2 .50 3 .10 3 .00 3 .00 3 .05 2 .60 1 .30 2 .80 1 .45 . 20 3 .10 1 .43 . 25 2 .90 1 .27 . 30 3 .30 1 .40 . 15 2 .90 1 .55 . 25 3 .05 2 .50 1 .30 . 10 2 .80 1 .45 . 20 3 .15 1 .35 . 40 * su=maxillary,in=mandibular

9 9

1 .30 1 .05 . 90 1 .10 1 .30

A ppend ix B Lum inance P rof i le s f or c on tro l s amp le s pec imen s v i ewed u nder t r an sm i t ted p o lar ised l i gh t .

T he k nown mon th o fd ea th o fe a ch a n ima li sp r in ted i nt h eu ppe rl e f tc or ne ro fe ach p age . T he i d en t if ica t ion o ft h ef i na lc emen tum l a ye ri sp r in ted o n t h eu ppe rr i gh to ft h ep age . E ach p age c on ta in st wo t ot h ree p rof i le sc or re spond ing t o 8 -p ixe l w ide t r an sec t s o f as i ng le s l ide v i ewed a t2 5X m agn if ica t ion . T r an sec t s a r e o rde red f r om t op t ob o t tom a nd a r e s paced a bou t 1 00 p i xe ls a par t . D ec reas e s i nl um inance v a lue s , o r " t rough s " , i nt h ep rof i le sc or re spond t o LAGs , i n c rea se s , o r " pe ak s " , t og row th z one s . I n c rea se s i nl um inance v a lue sa tt h et oo th m a rg in a re n o tc oun ted whe re : 1 )t h em ax imum v a lue o f t h e p eak i sl e s s t h an 2 5% o f t h e m ean v a lue o f t h e p re ceed ing p eak a nd /o r 2 ) t h e p eak c or re spond s t o as epa ra t ion o ccur ing b e tween t h e t o o th m a rg in a nd t h e r e s in . E a ch p rof i le i sl a be l led i nt h e u ppe r l ef tc or ne r a cco rd ing t ot h e m and ib le a nd t h e a na tom ica l p o s i t ion o f t he t oo th s amp led , e . g . , CON21P2 i st h e s e cond p r emo la r o fm and ib le n umbe r CON21 . LAGs a r en umbe red c on se cu t ive ly f r om t h e e name l /cemen tj unc t ion ( " 0 " ) t ot h e t o o th m a rg in ( "ma rg in " o r " m " ) .

1 00

MAY

G row th z one f i na l

a sc la 2 55 .00

r i t

I mars 0. 00 N =227

M ean=84 . 48

W idth=9

C a l ibrated()

• a sc 1b

0. C ID N =204

Mean=

1 0 9 .26

W id th=8

Cal i br at ed()

2 55 .00

r iI I l l ,

I I1

i l

2 .

i i,1 1 % 1

I I

1

I I I

pL,7 . 2 \

\fi e

I

1

I I . 1 ) L I

II 1

i k

I '

f ‘

1

. _ .

i

I

I )

I M ean= 1 02 .04

' . l id th=2 »

1 01

.

1 i i I1 i l j I I I I I ' l i \ I ° ii i1 L i I p i'

1 I

I1

3

N = 26o

1 1 ' 1II

i l

I I

C a l ibrated()

7

I .

n-a rg in

MAY

G row th z one f i na l

A SC2 I 2 55 .00

0 .00

2 55 .00

p i = 11 9

Mean= 14 5 .55

W id th=8

C a l ibra ted ()

a sc2 b

1

0. 00

2

N = 111

Mean=1 2 9 .59

1 02

W idth=8

C a l ibra ted( )

G row th z one f i na l

AUGUST

2 55 .00

a sc5a

0 .00 N =1 5 4

Mean= 13 5 .4 1

V idth= : 3

C a l ibra ted°

a sc5b 2 55 .00

( 8 . 1 1 ` . " ‘

' u s

1 1 . 1 1 1 1 * . 1

I I I I

3

I

5 1 L I

0 .00

N =1 3 7

M argin

Mean= 1 4 .50

' i r e idth=8

I

C a l ibra ted()

SC 5 . . .

' 7' 55 .00 r — f . . 1 ( s . I ' ' ,

. 1 ' 1, J , I

t I

.k

I

f

I I—( )

0

1

I ' s

. . . I I

I I

I 1

. 1' 13

i l l

I

4

Mean= 16 0 .71

V idth=8

1 03

I i i t i 1I 1

I I /

5

0. 00

N =1 : 3 1

i i I I

f•

3

• )

r. 1 I f i t

C a l ibra ted°

NOVE MBER

LAG f i na l a

c on20m 1 2 55 .00

0 .00 N =253

Mean= 116 .86

W idth=8

C a l ibrated()

c on2Orn 1 2 55 .00

, . , „ ,

i \ ?.

V s '

\ PI ' 1 . , )1

f

\ I I I

. ( . , .I I i 1 . 1 I I' 1 . i I I' ` ' .r i ar . 1

. 1 i. 1 1 i i \

IL . '

I % . 01 ' t , _ , , , , t 1 . 2

1

f t. 00

N =2 19

3

4

Mean= 10 6. 15

W idth=8

C a l ibrated()

Mean= 15 0 .1 9

W id th=8

C a l ibrated()

c on20m 1 2 55 .00

0 .0n N =256

1 04

i l

.

I I I ' t I

I.

NOVE MBER

G row th z one f i na l

c on20p2 2 55 .00

r n

1 1 5

. 4

7

4

1

0. I . D0

6

N =644

N 1 E -an= 14 2. 45

' y e ld th=8

C a l ibrated (. )

c on20p2 2 55 .00

d

L I

' ' I \N

A ( -\ 1 ' 1 1 (4(

)

I A )

Ll

14h . ,

1

II i f1 /I „ p i t I i l 9. / r n

L I. 1 : 10

N = i ;n7 ,

. 00

Mean= 141. 73

W id th=8

C a l ibrated()

c o n20p2

c : l ' ' , i r . o i ld i ro . .

. y, 111 , . o g % l i e! 1

1

' l i ' ' ) 1 . i ‘ IV I I I )) A I ( ) 1 I i I

1 1 , 1

0c I 0

1

2

N = E .1 2

3

4

t y l ear l= 157 . 83

l 'i

I' j h i f

1 1

1 05

f ,

1 , I ,

i

9 r n 7

5 W id th=8

1 11' n ^

C a l ibrated()

LAG f i na l

NOVE MBER

a

c on20p3 2 55 .00

0 . n0

N = 1 1 4

Mean=89 . 69

W idth=8

C a l ibrated( )

c on20p3 2 55 .00

1 1 .' 1 *

I

I

• 1

7-* ' I t

I

I

I

-)I I I

• • •I I

I I

i t

1 1

1

A

2

1

3 /4

5

0. 00

p i = 114

Mean= 14 4. 6E ,

W id th=8

C a l ibrated( )

on 2Op 3

( ' . , — I Ii , i

e name l

N =99

(\ . .

I I I

I 1 , 1 I I t t l I II I I I 1. I L . I 1 0

• % .

1

Mean= 17 0 .1 9

• . )

(

3

4 4 /marg in

W idth=8

1 06

C a l ibrated ()

i n a 1 1 )

f

LAG f i na l

DECE MBER c on21 r n1 2 55 .00

0 .00 N =359

Mean= 13 2

C a1 i b r t ed0

W idth=2

c on2 1m 1 2 55 .00

,

I , Al l ' I

" .1

• ;

' I .

d I • i . 1 I . I )

1

I. I0

N =730

2

7

4

3

V l ean= 113. 22

W id th=8

9 1 0

C a l ibrated ()

n2 1 n-1

. 7 2 55 .00

1i I

1 iI

r 1 i A l ,.

I

1

I

i

I m i c i N =321

I II I

0

i ' ll

I I

i

. 11 il

(

1

2

3

Mean= 12 4. 73

1 07

1

I

, . t s i „ 1 I

l' / \.

7 : 1

1

LAG f i na l

DECE MBER

c on2 1p 3 2 55 .00 in 11 1 1

t i

r y,f , ,

mar Q

' 111 , I I

5 1

0 . no

2

3

N =7A5

4

6

Mean= 111. 60

W id th=8

7

1 9 I

C a l ibra ted()

L i

C ON21 P 3 ' 7 ' 95 .00

p

P i l l )

I

Jul r

I i r . V \

e

I .

0. 0 o

0

1 N =31 9

2 .

4

N l ean=99 . 1

1 08

7 C a l ibrated()

2

( f ina l) •

LAG f i na l

DECE MBER

2 55 .00

a

c on22p2

I ,

ji

' I.

.

I

1 A % • , .

. 1

I i

(

I

I

0m % #I i I 9 10 I ,

i i I 4 , 1

,

. 1

. 4 .

N =250

2 55 .00

„1.1 i

I

M ean= 10 5 .54

W id th=8

C a l ibrated ()

c on22p2

e name l

I t o .

• • •\ ./ • , /

2

3

. JI I

I I

/I

/

L (

1 4 01

. 1. 1

• (

IAI I I N I

4

0 . '1

e

r

7

I I I I

I

r n

8

CI , 00

1

N =255

' 7 ' 55 .00

M ean= 11n . 22

W id th=8

0

C a l ibrated ()

c on22p 2

in.

enar i le1 ' -. ,2,.1' 1

. , ",

' a1

1

'

l ' . , . ./)

i'l

k1 . /L/ 11 , . ,\. . . ,

2

3

4

I

I II

i,

I .. . . .1

.y

1 .4

1 1

Ii 1

7 L i 8

' e l id th=8

1 09

. , 1

• 11 , . "

, C I 0 M ean= 10 9 .82

. . I

C a l ibrated ()

I ' I I

. 1i i. . , i ,1

II

I

A

N =264

. . .

' I

i i -AP' t

c )

I I

I I )I 10/r n

END J ANUARY

LAG f i na l

ma rg in + s epa ra t ion

2 55 .00

c on23m1 _ _h ie .t o-

0 .00 N =405

Mea n =101 . 69

W id th=12

C a l ibrated()

marg in

• 7 • iF i. 00

c on23rn1

9

0. 00

Me an =133 . 90

10

' e l idth=12

C a l ibrated()

END J ANUARY

LAG f i na l

a

C ON23 P 2 2 55 .00

CI

1

I It . , )

1 0 . no

0

N =278

2 55 .00

1

V ' r . 1 / \t , 'I I I4 dr \(

r ,

, \ rs . , .

2

3

Mean=80 .16

1 1

i

,. . .. , ' . , , . ; 4

W id th=8

5

6

7

: 3 mar

C a l ibra ted( )

C ON23 P 2

g i k . r

/

I \II N

1, 1

'

I I

0 .00

0 N =253

1

2

Mear i=9 1-1 . 50

11

3

4

W id th=8

I I I I

c-

C a l ibrated ()

i d

LI

P

I

7

1

I .

I

1 4I

: 3r i a

END J ANUARY

LAG f i na l a

C ON2 4M1 2 55 .00

marg in

1 3

N =346

Mean=199 . 80

' e l id th=8

C ON24M1 2 55 .00

C a l ibra ted ()

'/ W s

4 r i

; E :

3

1

7

0. 0c I N =332

Mea t-1 = 1 '29 .29

W idth=3

C a l ibra ted()

C ON24M1

C :

2 55 .00

I ' d

2 1 : .

3 1

0. oc i N =423

Mean=186 . 8 : E :

W idth=8

12

C a l ibra ted°

0

LAG f i na l

END APR IL c on25mla 2 55 .00

0 .00 N =194

Mean=159 .22

c on25m1

2 55 .00

p i ,I •V f e . 4 I

i

1 1 i I ?

' W idth=8

C a l ibra ted( )

h isto-

› I ) I

. 61

i t .. r' 41 N I % .

I

A l

II

• I h

I ' .1 -,

' . \/

L. 2

I

,

1

r i

2

1 \

I I /

I

p . , ' 1 4 1 4

I i .

I . i

I III v. :

. 2 i i

. , .i.

z

0

I II I

e

t s

r

0 . 1 10 N =141

` ,5 5 .0n

Mean=104 . 90

W id th=8

C a l ibrated()

--con25rhl h i sto-

( \A 1 , 1

, ,, 1 2

r 1 = 155

Mean=90 . 8 1

13

W idth=8

. I .l i i i I ,

C a l ibrated()

i l I I

G row th z one f i na l

J UNE

2 55 .00

0. 00

N =377

M ean=136 . 74

' y f idth=8

C a l ibrated°

M ean=129 .92

W idth=8

C a l ibra ted°

N E : 1 11 1 2 55 .00

0 .00 N =358 r e 1 . 1 M 1 2 55 .00

' It . r

1)

I :

/ I l o .

R

di

1 I I ,

o

1

' .

j

( .

I I

' 1.i, . 1

I 1I 1 I 'm 1 I J I f f l e i ' I I

1 , 2 , j i ,

. j \ ' . t i o\ r‘i )

I

p d l

' 1I

. ,

2

3

4

I I

. . . , , . . ,,

5

1I1 . 4 1 ) II 1I r n' LI

0 .00 N =382

M ean=125 . 02

V idth=8

14

C a l ibrated()

J UNE

G row th z one f i na l N B 1 . 1 M2

a

2 55 .00

0. CI 0 N =28E .

Mean=128 .23

V id th=8

C a l ibrated()

N B 1 . 1 M2

b

h

i r i

(

4

V

I

I

I IP ‚I

1

kJ

IA

1 _ 1

(

I

i

i

\

I de t\ I\

(11

1

FdI j

3

0. 1 . 1 1_ 1 N =291

I 1 1

( I

1 ? . r94 . 54

' e l i t i th=2

1 1 ,1 1

I

(

I I ; I

5 e E .

4

C a l ibrated°

N B 1 . 1 1 .2 ' 7 ' 55 . 00

I

,.

I J 1

11. 1 1 J i

I

I II I l j

e name ll

I j i

. ,

[ 41 I .

II ,

I

I I( I l

1

/ ( I d

1 I

I I

I

I . , w , i

i • A ' 1 , 1 I 1

I i

4

1

N =295

2

r l ean=87 .38

15

41 . 1

1 0 ir

i jI

.1 , .i

.i

. 4

1 ‘ '

C a l ibrated ()

i

r n3 r -Q in ,

G row th z one f i na l

J UNE

a

( i nverted)

N B 1 .2 M 1 . 0 0

r . J = 2 16

Mean=149 .46

' , . ' idth=8

C a l ibra ted()

N E : 1 .2 M 1

t u

2 55 .00 . e •— ,

A

. 1

I ' I

I t . .

i

/

I

' * I I I

2

1

II

1 \.f e t I V

3

4

1 -1 1 L . . 1 r

? II. I I I

)

I I 1 ,

I I

. I

r i argin

0 .00

N =224

Mean = 167 .77

s e i id th=8

C a l ibrated ()

N B 1 .2 M 1 2 55 .00 , P\21 n # II

I I i i

I V "

I

;

i j

1

2

0. C 1 0 N =227

Mean=147 . : 3 0

W id th=2 :

16

C a l ibrated()

marg in

J UNE

G row th z one f i na l

n b 1.2m2a 2 55 .00

A 1 , II

0. 00

0 N =344

,I

I I I‚ 1 .1 k .

•

I j

1

I .

3

1 Mean=133 . 35

. . i 'dth=8

i m . 9rd in

C a l ibrated°

n b1.2m2

t i

2 55 .00

1 4(1 1 (11 1 1' i l l

I

N = 7 . t '. . 73

2 55 .00

1

I

I

11 4

1

2

( I

1 ean=118

1

V1 4 1

4

Sr i

3

. . . ' idth=8

C a l ibr 3ted( f )

n b1

I 1 II ' l

IN

I

rd

r

4

5

1 4

6. 1r n r qi n

N =344

Mean=142

' y l idth=8

17

C a l ibrated()

J UNE

G row th z one f i na l

N B 1 .3 P 3 2 55 .00

l a g

l ag

0 .00 N =366

Mean=142 . 15

W id th=8

C a l ibrated ()

N E : 1. 3P 3 ' 7 2 5 .00

• „

1-1

'

I I I I I

I j

l ag

0. C IO N =338

Mean=157 . 69

\ % / id th=8

18

C a l ibrated()

G row th z one f i na l

J UNE

2 55 .00

, N 13 1 . 3P4

I A )

HO ,

1 1 34 5

' 1 01 2 1 (1

V4 1 , el

I ji

0. CI 0 N =437

Mean=106 . 16

W id th=8

C a l ibrated ()

n t i 1. 3 p4t , 2 55 .00

I )

r i l

7 ' 1 4 . 00 N =37 : 1 :

t l ean=105 . 49

19

C : a l it ir t e . in

G row th z one f i na l

OCTOBER

2 55 .00

n b2 . 1ml a

f 4 5

N =213

2 55 .00

Mean=139 . 50

C a l ibrated()

W id t h=8

N B2 1M1 b

1 A T \ •i t " ) I (A I

)

' P I

I I f

V 1 /

\

' II

4

0

3 N =236

Mean= 1 39 . 16

W idth=8

C a l ibrated()

n b2 . 1r n 1 I

I

(P .1 1 \ 4\

I

)

p i l

p i

I

. 2

I I

4

I ' I

I

Ir , 4

II

1 , 1I L P

o no

( . 1

3

1

r l ean=142 .83

r a l ibra tedr . . )

1 20

. 9

? I

f ij

0 N =235

1 . 1 1

I 4

I

marg i n, I

OCTOBER

G row th z one f i na l

N =30 i .

Mean= 1 76 . n3

W id th=8

C a l ibrated()

n b2 . 1D 4b 2 55 .00

I I L .

i h r l

I d i l dr P 1 I II I f ' " 1

1

I

1 4 1

1 ) 1

2

3

r(

i d

1 \ , 1 (

II

I

j \

( ' 7

1 _ 1 . 1 _ 1 1

N =317

Mean=156 .79

W id th=8

1 21

C a l ibrated()

II

1. 1

I ' l l

NOVE MBER

LAG f i na l ( a= g row th z one 1 00% e xpec ted )

N B 3 .1 M 1

a

2 55 .00

0

marg

4

0 .00 N =293

Mean= 14 6 .90

% . ' idth=8

C a l ibra ted()

N E : 3 . 1 M 1 2 55 .00

4

A P . CI

t c . 2

0 .00 N =251

2 55 .00

Mean= 1 56 .37

W idth=8

C a l ibrated()

N B3 .1 M 1 CI

, t

l

r I f r Y s il i I

\ I

I ' , I i I i

1 I '

I I i i i I , 1 I

0 N =267

V ' dI 1 4 I b : 41 41 /

I r 1 1

I I ,

0 d (1 I 1 i

r .,

h; 1

A . 0

hi

2

Mean=1 1 3 .83

1

' I"

z , W idth=8

1 22

I r j' ri 4

1

I I 1

h

I ' l I I

1 )

li l ' l' I d°A d 1 P

(

1

I ' I . 5r i C a l ibrated()

I

' I I1

•

6

I I

LAG f i na l

NOVE MBER

a

N B 3 . 1 M2 2 55 .00

0. C IO

Mean=1

N =219

2 6. 2

W id th=8

C a l ibrated()

N B 3 . 1 M2

1 \ e I I I

I ( \ II I ) ,

II

Ii i

I

9

. 1 I

I I I

Al

I _ I l

\ ii

I

I )

› ) 4

• I

CI N =223

5

n-l a rg in M ean=137 . 66

' . / idth=8

C a l ibrated()

N E : 3 . 1 M2 ' 7 55 . 00

F I J I

•

I.1 1 I

I

I I I•

II

i l

I I I . i lI I

L . ,

. »II

.

I

k

1 2, t\ (\ I

1 ,1

,

' 1, 1

'

, I

0 N =230

• f ' i

1 M ean=100 . 0 1

W id th=8

1 23

I .1

i i Ii I

II

I / I1 i el l4I L i1

L i Ii ,

3

1 C a l ibrated()

1 1

4

1

1 . I

4

5

marg in

A PPENDIX C R esults o f t he F ossil S amples.

I dent

S ite

C ouche

D ate

A nat.

P os.

I dent.

% z one S eason

B 01

L augerie H E

M ag .III

M 2G

i nf

I LLEG

0

B 02 B 03 B 04 B 05 B 06 B 07

L augerie L augerie L augerie L augerie L augerie L augerie

H H H H H H

E E E E E E

M ag .III M ag.III M ag .III M ag.III M ag .III M ag .III

M 2G M 2G M 2G M 2G M 2G M 2G

i nf i nf. i nf i nf i nf i nf

L AG I LLEG I LLEG L AG I LLEG I LLEG

3 0 0 3 0 0

B 08 B 09 B 10 B 11 B 12 B 13 B 14 B 15 B 16 B 17 B 18 B 19 B 20 B 21 B 22 B 23 B 24 B 25 B 26 B 27 B 28 B 29

L augerie H L augerie H L augerie H L augerie H L augerie H L augerie H B adegoule B adegoule B adegoule B adegoule D uruthy D uruthy D uruthy D uruthy D uruthy D uruthy D uruthy D uruthy D uruthy D uruthy D uruthy D uruthy

E E E E E E

M ag .III M ag.III M ag .III M ag.III M ag .III M ag .III M ag . M ag. M ag. M ag. M ag .IV M ag.IV M ag .IV M ag .IV M ag .IV M ag .IV M ag .IV M ag .IV M ag .IV M ag.IV M ag.IV M ag.IV

M 2G M 2G M 2G M 2G M 2G M 2G P 3D P 3D P 3D P 3D M 1G M 1G M 1G M 1G M 1G M 1G M 1G M 1G M 1G M 1G M 1G M 1G

i nf i nf i nf i nf i nf i nf i nf i nf i nf 1 nf s up s up s up s up s up s up s up s up s up s up s up s up

I LLEG Z ONE I LLEG * ZONE Z ONE * ZONE Z ONE L AG L AG Z ONE Z ONE * ZONE Z ONE I LLEG L AG I LLEG Z ONE ? ZONE * ZONE Z ONE Z ONE

2 5 4 3 3 1 3 7

8 30 B 31 B 32 B 33 B 34 B 35 B 36 B 37 B 38

D uruthy D uruthy D uruthy D uruthy D uruthy D uruthy L augerie H E L augerle H E L augerie H E

M ag .IV M ag.III M ag .III M ag .III M ag.III M ag .III M ag .III M ag .III M ag.I

M 1G M 1D M 1D M 1D M 1D M 1D P 2D P 2D P 2G

s up s up s up s up s up s up s up s up i nf

Z ONE Z ONE Z ONE Z ONE

2 5 1 0 2 4 1 2

Z ONE Z ONE I LLEG Z ONE

4 0 4 2 4 6

1 1 1 1 0 1 1 0 1

8 39

L augerie H E

M ag .I

P 2G

i nf

Z ONE

3 3

1

4 4 4 4 4 4 4 4 4 4 4 4 4 5 5 5 5 5

1 24

4 0

7 5 4 1 1 9 8 1

5 6 7 8 7 3 2 6 6

0 1 0 0 4 1 4 1 3 3 1 1 4 1 0 3 0 1 0 4 1 1

A PPENDIX C R esults o f t he F ossil S amples.

I dent B 40 B 41 B 42 B 43 B 44 B 45 B 46 B 47 B 48 B 49 B 50 B 51 8 52 B 53 B 54 B 55 B 56 B 57 8 58 B 59 B 60 B 61 B 62 B 63 B 64 B 65 B 66 B 67 B 68 B 69 B 70 B 71 B 72 B 73 B 75 B 76 B 77 8 78 8 79 B 80

S ite L augerie H E L augerie H E L augerie H E L augerie H E L augerie H E L a M adeleine L a M adeleine L a M adeleine R ondduBarry R ondduBarry R ondduBarry R ondduBarry R ondduBarry R ondduBarry R ondduBarry R ondduBarry R ondduBarry R ondduBarry R ondduBarry R ondduBarry R ondduBarry R ondduBarry E nval E nval E nval E sclauzur E sclauzur E sclauzur E sclauzur S t G ermain S t G ermain S t G ermain S t G ermain S t G ermain S t G ermain S t G ermain S t G ermain S t G ermain M oulin N euf M oulin N euf

C ouche

2 7 2 7 2 7 F 2 F 2 F 2 F 2 F 2 F 2 F 2 F 2 F 2 B 3 E 3 E 3 E 3 E 3 f f f

D ate

A nat.

P os.

I dent .

H .I II M ag .III M ag .0 M ag .0 M ag .0

P 2D P 2D M 2G M 2G M 2G M 2D M 1G MO M 2D M 2D M 2D M 2D M 2D M 2D M 2D M 2D M 2D M 2D M 2D M 2G M 2G M 1D M 1D M 1D M 1D M 2D M 2D M 2D M 2D P 4G P 4G P 4G P 4G P 4G M 1G M 1G M 1G

s up s up i nf i nf i nf i nf I nf i nf s up s up s up s up s up s up s up s up s up i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf i nf s up i nf i nf

I LLEG Z ONE Z ONE Z ONE L AG I LLEG Z ONE Z ONE L AG Z ONE Z ONE Z ONE Z ONE Z ONE L AG L AG Z ONE Z ONE Z ONE Z ONE L AG Z ONE Z ONE Z ONE L AG Z ONE ? ZONE Z ONE Z ONE Z ONE

M ag . M ag . M ag. M ag . M ag . M ag . M ag . M ag . M ag . M ag . M ag . M ag . M ag . M ag . M ag . M ag .

3 3 3 3 3 1 1 1 4 2

M 2G M 2G

1 2 5

I LLEG Z ONE I LLEG Z ONE Z ONE Z ONE L AG Z ONE

% z one S eason

8 6 7 1 6 5

5 7 5 8 7 2 4 9 7 9 7 0

9 1 2 1 1 04 1 8 3 3 6 5 4 5 6 3 2 7 3 5 7 7

7 3

1 00 9 2 1 5

C 1 1 1 3 0 1 1 3 1 1 1 0 1 3 3 1 1 2 4 3 1 1 1 3 1 0 1 1 1 0 0 1 0 0 3 1 3 4

A PPENDIX C R esults o f t he F ossil S amples .

I dent B 81 8 82 B 83 B 84 B 85 B 86 B 87 B 88

C ouche

S ite M oulin M oulin M oulin R oc d e R oc d e R oc d e R oc d e R oc d e

N euf N euf N euf M cpsl M cpsl M cpsl M cpsl M cpsl

D ate

2 b 2 b 2 2 2 b 2 2 3

B 89 R oc d e M cpsl 3 B 90 R oc d e M cpsl 4 B 91 R oc d e M cpsl 4 d B 92 R oc d e M cps2 2 B 93 R oc d e M cps2 1 B 94 R oc d e M cps2 3 c B 95 R eignac B 96 R eignac R h B 97 R eignac R h B 98 R eignac R h B 99 R eignac R h B 100 C ap B lanc 1 B 101 C ap B lanc i B 102 C ap B lanc i * Seasonal a ttributions: 0 =none ,

1 26

A nat .

P os.

I dent .

M 2G M 2G M 2G M 1D M 1D M 1D M 1D M 2G

i nf i nf i nf i nf i nf i nf i nf i nf

Z ONE Z ONE Z ONE Z ONE L AG L AG I LLEG Z ONE

M 2G i nf M 2 i nf M 2 i nf P 4G s up P 4G s up P 4G s up M 2G i nf M 2G i nf M 2G i nf M 2G i nf M 2G i nf M 2D i nf M 2D i nf M 2D i nf 1 =summer , 2 =fall ,

% z one S eason

* LAG Z ONE ? LAG L AG I LLEG Z ONE I LLEG Z ONE I LLEG I LLEG I LLEG Z ONE L AG I LLEG 3 =winter ,

8 3 1 00 6 0 2 6

6 4 5 3

1 9 3 2

6 4

4 =spring

1 2 1 1 3 3 0 1 2 1 0 3 0 4 0 2 1 1 1 2 4 0

P late 3 .1 Mandibular tooth row, buccal and

sagittal

views.

Plate LAGS

3 .2

visible

Top:

on a polished, etched tooth surface under S . E. M. Sample no. NB 1 .2 ( P2). Magnification 1 80X. Width of field 0 .64 mm.

Bottom:

Magnification

8 25X.

W . O. F.

0 . 14

mm.

P late

3 .3

Histological preparation. Sample no. CON 2 2 ( M2). Magnification

P late

Sample

2 5X.

3 .3b

Histological preparation. no. CON 2 2 ( M2). Magnification

6 3X.

Plate

3 .4

Microradiograph of Control Sample Specimen. Sample no. CON 2 2 ( M1). Magnification: 25X.

Plate

3 .5a

Growth Layers in Cement Viewed Under Transmitted, Ordinary Light. Sample no. CON 2 2 ( P2). Magnification 25X.

Plate 3 .5b Transmitted, Polarised Light. Sample no. CON 25 ( Ml). Magnification:

25X.

Plate 3 .5c Transmitted, Polarised Light. Sample no.

CON 25

( Ml).

Magnification:

6 3X.

E namel/ist growth

Periodontal

z one

l igament

I

I

Plate

3 .6

Primary Cementum Layer. The f irst l ayer of cementum deposited ( on the l eft o f this photograph o f a z ebra third i ncisor), when viewed under tramsmitted l ight, i s c learly distinguishable f rom succeeding growth l ayers. Resorption bays clearly demarcate enamel and cementum regions i n this photograph.

Plate

3 .7

Resorption Bays in Enamel Viewed Under SEM. Specimen: NB 1 .3 ( P4) Magnification: 2 210 X . Width of Field: 5 .52 X 1 02.

Plate

5 .1

Fossil Teeth Observed Using Microradiography.

Top: Sample no. B-58. Magnification 25X. Bottom: Sample no. B-3. Magnification 2 5X.

Plate

5 .2

Fossil Tooth Observed Under Transmitted Polarised Light. Sample no. B50 Magnification: 1 00 X .

Plate Effects

of

5 .3

Poor Preservation on Observation of Cementum Growth Layers i n Fossil Teeth. Sample no. B 44. Magnification: 6 3 X .