Hunting Strategies in Central Europe During the Last Glacial Maximum 9780860549123, 9781407349831

Table of contents :
Front Cover
Copyright
Table of Contents
List of Tables
List of Figures
Chapter 1: Introduction
Chapter 2: EVALUATING THE DATA
Chapter 3: Biology and Ecology of Fauna at Grubgraben
Chapter 4: Hunters and Their Prey
Chapter 5: Horse Hunting, Processing, and Transport
Chapter 6: Reindeer Hunting, Processing, and Transport
Chapter 7: Secondary Faunal Resources
Chapter 8: Conclusions
References Cited
Appendices

Citation preview

Hunting Strategies in Central Europe During the Last Glacial Maximum

Dixie West

BAR International Series 672 1997

Published in 2019 by BAR Publishing, Oxford BAR International Series 672 Hunting Strategies in Central Europe During the Last Glacial Maximum © Dixie West and the Publisher 1997 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 9780860549123 paperback ISBN 9781407349831 e-book DOI https://doi.org/10.30861/9780860549123 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 John and Erica Hedges in conjunction with British Archaeological Reports (Oxford) Ltd / Hadrian Books Ltd, the Series principal publisher, in 1997. This present volume is published by BAR Publishing, 2019.

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T ab le o fC on ten t s T ab le o fC on ten ts L i s t o fT ab le s L is t o fF igu re s

i i v

G rubg raben : L eve l 1 . A na tom ica l F requenc ie sa nd MN I . H eads _ B od ie s . G rubg raben : L eve l 2 .

C hap te r 1 :i n t roduc t ion . Gene ra l S ta temen ta nd H is to r ica lP e r spec t ive .

1

P e rspec t ive R esea rch Goa ls P rev ious E nv ironmen ta l a nd A rchaeo log ica l I nve s t iga t ion . App roach a nd O rgan iza t ion o fT h is I nve s t iga t ion . .

1

H eads .

3

B od ies . G rubg raben : L eve l 3 . A na tom ica l F requenc ie sa nd MN I . H eads . B od ies .

C hap te r2 :E va lua t ing t h eD a ta . . R eco rd ing t h eD a ta . . I nf orma t ion R eco rded i nt he D a ta B a se . C a ta logue Numbe ra nd P roven ience . T axon a nd E lemen t I den t if ica t ion . T he G rubg raben A ssemb lage . C u l tura l Mod if ica t ions . C u t t ing , C hopp ing , a nd A b rad ing o nB one S u rf ace s . B reaks a nd Ma r row E x t rac t ion . B one G rea se Manuf ac tu re . B u rn ing . Na tu ra l Mod if ica t ions .

A na tom ica lF requenc ie sa nd M IN I .

3

5 7 5 7 5 7 6 1 6 1 6 1 6 1 6 1 6 4 6 4 6 4

G rubg raben : L eve l 4 .

6 4 6 4

7

A na tom ica lF requenc ie sa nd M IN I

6 4

7 7

H eads B od ie s

6 4

7 7

G rubg raben Ho r se s N a tu ra l a nd C u l tu ra l Mod if ica t ions o f

1 1

t h eG rubg raben Ho r se s C ompa ra t ive D a ta A na tom ica l F requenc ie so f Hor se B ones a t

6 9 6 9

1 1 4 1 4 1 4

C amps S t ran ska S ka la I V H eads B od ies

7 6 7 6 7 6

1 8 1 8

B reaks C u t Ma rks a nd S ur f ace E ros ion

8 2 8 2

5

6 9 6 9

7 6 8 2

Wea ther ing . Gnaw ing .

2 0

S ea son o fD ea th

Roo t le tE tch ing .

2 4

P a t te rns o fP os tc ran ia l P roce ss ing a nd

Mea suremen ts a nd We igh t .

2 6

T ran spor t T he P a t te rn o fH ead T ran spor t a nd P roces s ing A na tom ica l F requency a nd N a tura l

8 2

3 2

A t tr i t ion

3 5 3 6

D iscuss ion o fH ypo the se s

8 5 8 5

C hap te r 3 :B io logy a nd E co logy o fF auna a t G rubg raben R ang if er t a randus : R e indee r /Ca r ibou Equus c f .p r zewa l sk i /f erus : W i ld h o r se Capra i b ex : Moun ta in g oa t A lopex l agop ia : A rc t ic f ox . L epus t i m idus : A rc t ic h a re . A dap ta t ion , M ig ra t ion , a nd t he L a s t G lac ia l Max imum . C hap te r4 :H un te r sa nd T he ir P rey .

3 2

3 7

8 4

C hap te r6R e indee rH un t ing , P roce ss ing , a nd

3 7

T r anspor t

8 8 8

3 7

S ea sona l i ty A na tom ica l F requenc ie s , M IN I , a nd

4 3

S ubs is tence S t ra teg ie sa nd B ody S i ze . B ody C ond i t ion . Ma r row /G rea se R e sources .

4 3 4 3 4 3

S ubs is tence D ec is ions Ma sked b yA t t r i t ion

4 4 4 4

D emog raphy a tG rubg raben

8 8

Mor ta l i ty P a t terns o fH e rd P opu la t ions P roce ss ing S pec if ic C a rca s sP ar ts o f R e indee r H eads a nd Mand ib le s

9 4 1 0 1 1 0 1 1 0 1

4 4

R ibs P ha lange s B one F ragmen ta t ion a nd P roce ss ing o ft h e

Equus c f .p r zewa l sk if erus : Ho r se .

4 8

G rubg raben R e indee r

1 02

Capra i b ex : I b ex .

5 0

Q uan t if y ing F ragmen ta t ion

1 03

F urbea re r s .

5 2

Ma r row P roces s ing

S ea sona l S t ra teg ie s . S pr ing .

5 4 5 4

G rea se Manuf ac tu re C ompa ra t ive D a ta

1 03 1 09

S umme r . A u tumn . . W in te r .

5 4 5 6

C onc lus ion s

F ur the r S ubs is tence S t ra teg ie s . Rang if er t a randus : R e indee r .

C hap te r5 : Ho r se H un t ing , P roce ss ing , a nd T ranspor t E p ig rave t t ian H or se H un t ing , P roce ss ing a nd T ran spor t .

5 6

C hap te r 7-S econda ry F auna l R esources Mammo th

1 02

1 1 1 2 1 1 22 1 22

5 7

I b ex B ov ids

1 22 1 23

5 7

F ox H a re

1 23

1 2 3

B ea r S umma ry

1 /7 1 27

C hap te r 8 :C onc lus ions

1 28

R ef e rence s C i ted

1 32

A ppend ix 2 . 1 : T he d a ta c od ing s y s tem u sed t o r e cord a t tr ibu te so fe ach s pec imen i nt h e G rubg raben a na lys is A ppend ix 2 .2 : G nawed b ones r e cove red f r om t h ef our c u l tu ra ll e ve ls a tG rubg raben

1 46

1 49

A ppend ix 5 . 1 : Numbe ro fi d en t if ied s pec imen s a nd m in imum n umbe ro fe l emen ts o fh o r se f r om t he f ou rc u l tu ra l l e ve ls a tG rubg raben

1 50

A ppend ix 5 . 2 : Numbe ro fi d en t if ied s pec imens a nd m in imum n umbe ro fe l emen ts o fh o r se r e cove red f r om S agva r , P i l isszan to R ock she l te r a nd S t ranska S ka la I V A ppend ix 6 . 1 : N umbe ro fi d en t if ied S pec imen s a nd m in imum n umbe ro fe l emen ts o fr e indee r r e cove red f r om t h ef our c u l tura l l e ve ls a t G rubg raben

1 5 1

1 52

A ppend ix 6 . 2 : Numbe ro fi d en t i f ied s pec imens a nd m in imum n umbe ro fe l emen ts o fr e indee r r e cove red f r om P i l is szan to R ock she l te r , S agva r , a nd P i l isma ro t

1 53

1

ab les L i s to fT 2 . 1 . N ISP a nd p e rcen tages o ff a una r e cove red f r om t he f ou rc u l tu ra l l e ve ls a tG rubg raben

5 . 6 . Mean w e t w e igh ts o fm a r row i nm a jo rl o ng b one sf o r a du l t c a r ibou a nd z eb ra

1 2

6 . 1 . Age p rof i les o fr e indee rm and ibu la rt e e th r e cove red

2 . 2 . T ype so fc u l tura l mod if ica t ion sf o und o n r e indee r b one s r e cove red f r om t he f ou rc u l tu ra l l e ve ls a t G rubg raben

f r om t h e 1 990 e xcava t ion sa tG rubg raben c ompa red 1 5

w i th S p ie s s( 1979 ) d a ta f r om A br i P a taud a nd L ogan ( 1990b )d a ta f r om 1 986 -87 e xcava t ion s a tG rubg raben

2 . 3 . Numbe r sa nd p e rcen tages o ft y pe so fb reaks o n r e indee rb one sr ecove red f r om t h ef o ur c u l tura l l e ve ls a tG rubg raben

8 4

9 0

6 . 2 . Age r a t io o ft h eG rubg raben s amp le o fm and ib le s v e r sus at h eo re t ica l s amp le o f1 000 c a r ibou f r om t h e

1 7

9 4

N e lch ina h e rd d u r ing a u tumn 2 . 4 . Numbe r sa nd p e rcen tage so ft y pe so fb reaks o nh or se b ones r e cove red f r om t h ef o u rc u l tu ra l l e ve ls a tG rubg raben

6 . 3 . Ma r row c av i ty v o lume so fa na tom ica l e l emen ts f o r 1 7

c ar ibou

2 . 5 . C oun t o fb one t ype sa nd b u rn ing r e cove red f r om t h e f ou rc u l tu ra l l e ve ls a tG rubg raben 2 . 6 . Wea the r ing s t age so fr e indee rb one s r e cove red f r om G rubg raben

6 . 4 . Numbe ro fa r t icu la rh eads a nd s haf tf r agmen ts o f 1 9

r i bs f r om t h ef ou rc u l tu ra l l e ve ls a tG rubg raben 6 . 5 . Mean l e ng th a nd r a nge o fr i b f r agmen ts f r om t he f ou rc u l tu ra l l e ve ls a tG rubg raben

2 1

6 . 6 . Numbe ra nd k i nds o fb reaks o n p ha lange so fr e indee r f r om t he f ou rc u l tu ra l l e ve ls a tG rubg raben 1 02

2 . 8 . C ompa r ison o ft a phonom ic t r a i ts b e tween h uman a nd a n ima la c t iv i ty o nb one

1 02

2 1

2 . 7 . Wea the r ing s t age so fh or se b one sr e cove red f r om G rubg raben a nd S t ranska S ka la

1 0 1

1 02

6 . 7 . P e rcen tage so fi d en t if ied a nd u n iden t if ied r e indee r 2 5

b ones r e cove red f r om t h ef ou rc u l tu ra l l e ve ls a t G rubg raben

1 03

2 . 9 . T ype so fg naw ing f ound o nb one sr e cove red f r om G rubg raben

2 4

6 . 8 . T ype s , n umbe r sa nd ° Ao fc omp le te a nd i n comp le te r e indee rb ones r e cove red f r om t h ef our c u l tu ra l l e ve ls a tG rubg raben

2 . 10 . R oo t e t ch ing s t ages o n r e indee rb one s r e cove red f r om t h ef ou rc u l tura l l e ve ls a tG rubg raben

1 04

2 7 6 . 9 . Numbe ro fi d en t if ied s pec imens a nd w e igh t i ng rams

2 . 11 . R oo t e t ch ing s t age s o n h o r se b ones r e cove red f r om G rubg raben a nd S t ranska S ka la

o fr e indee rb one sr e cove red f r om t h ef ou rc u l tu ra l l e ve ls 2 7

a tG rubg raben

2 . 12 . B one w e igh ts a nd p e rcen tage so fi n d iv idua l t a xa r e cove red f r om t h e f ou rc u l tu ra l l e ve ls a t G rubg raben

1 05

6 . 10 . Numbe ro fi d en t if ied s pec imens o fr e indee rb one f r agmen t s p e rk i log ram o fw e igh t

10 6

3 0 6 . 1 1.Means o fl e ng th sa nd w id ths ( i n c m ) o fl o ng b one

3 . 1 . D eve lopmen t a nd c a s t ing o fR e indee r /Ca r ibou a n t le r s

3 4

4 . 1 . E thnog raph ic r e indee r h un t ing s t ra teg ies

4 7

s haf t s p l in te r s r e cove red f r om t h e f ou rc u l tu ra l l e ve ls a tG rubg raben

1 03

6 . 12 . N ISP o fb one e nds a nd b one s haf ts o fr e indee rf r om t h e f ou rc u l tu ra l l e ve ls a tG rubg raben

11

5 . 1 . Numbe r o fi d en t if ied s pec imen sa nd m in imum n umbe ro fh o r se e l emen ts f r om t h ef o u rc u l tu ra l l e ve l sa tG rubg raben 5 . 2 . Wea the r ing s t age so fh o r se b one s r e cove red f r om G rubg raben a nd S t ranska S ka la

7 . 1 . Numbe ra nd w e igh t i ng rams o fi v ory f r agmen ts 5 8

r e cove red f r om t h ef ou rc u l tu ra l l e ve ls a tG rubg raben 7 . 2 . N ISP a nd MNE o fa na tom ica l p ar t so fi b ex r e cove red

7 0

f r om G rubg raben

5 . 3 . R oo t e t ch ing s t age so n h or se b one sr e cove red f r om G rubg raben a nd S t ranska S ka la

5 . 5 M in imum n umbe ro fh o r se b one s f r om G rubg raben , S t ranska S ka la I V , a nd S agva rd i v ided b y t h e n umbe r o fl i ke e l emen ts f ound i no ne h o r se

1 22

7 . 3 . N ISP a nd MNE o fa na tom ica l p a r t s o fb ov id r e cove red 7 0

f r om t h ef ou rc u l tu ra l l e ve ls a tG rubg raben

5 . 4 . Numbe ro fi d en t if ied s pec imen sa nd m in imum n umbe ro fe l emen ts o fh o r se f r om S agva r , P i l isszan to R ockshe l te r s Ia nd H , a nd S t ran ska S ka la I V

1 22

1 23

7 . 4 . N ISP a nd MNE o fa na tom ica l p ar ts o ff o x r e cove red f r om t h ef ou rc u l tu ra l l e ve ls a tG rubg raben

1 23

7 2 7 . 5 . N ISP a nd MNE o fa na tom ica l p ar ts o fh a re r e cove red f r om G rubg raben 174 8 3 1 1

L is to fF igures F igu re

P age

1 . 1 G eog r aph ic d i s t r ibu t ion o ft h eC en t ra l E u ropean s i t e si nt h es t udy samp le

2

1 . 2 Map o fC en t ra lE u rope s how ing t h e l o ca t ion o fm a jo rs i te sd i scu s sed i nt h et e x t

4

F i gu re

P age

4 . 5 S ea sona la tr ibu te so fa r c t ic h a re u s ef u li n r e con s t ruc t ing P a leo l i th ic f o od / techno logy m anagemen ts y s tem s

5

5 . 1 N umbe ro fi d en t if ied s p e c imen so fh o r s e f r om l e ve l 1a tG rubg raben

5 9

6 0

2 . 1 S t ra t ig raph ic s e c t ion o fG r ubg raben s how ing c u l tu ra ll e ve l s

8

2 . 2 Map o fG rubg raben AL I

9

5 . 2 H i s tog ram so fn umbe ro fi d en t if i ed s p e c imen s ( N ISP )and m in imum n umbe ro fe l emen t s( MNE ) o fh o r se f r om l e ve l 1a tG rubg raben

2 . 3 P l an o ft h es t one p avemen tI d le i nG rubg raben AL2

1 0

5 . 3 N umbe ro fi d en t if ied s pe c imen so fh o r s e f r om l e ve l 2a tG rubg raben

6 2

2 . 4 P e rcen tage so ft a xa r e cove red f r om t h e f ou rc u l tu ra ll e ve la tG rubg r aben

1 3

5 . 4 H is tog ram so fn umbe ro fi d en t if ied s p e c imen s ( N ISP )a nd m in imum n umbe ro fe l emen t s( MNE ) o fh o r se f r om l e ve l 2a tG r ubg raben

6 3

5 . 5 N umbe ro fi d en t if ied s p e c imen so fh o r s e f r om l e ve l 3a tG rubg raben

6 5

5 . 6 H is tog ram so fn umbe ro fi d en t if ied s p e c imen s ( N ISP )a nd m in imum n umbe ro fe l emen t s( MNE ) o fh o r se f r om l e ve l 3a tG r ubg raben

6

2 . 5 L oca t ion so fc u l tu ra lm od i f ica t ion so n r e indee rb one sf r om t h ef o u rc u l tu ra ll e ve l s a tG rubg raben

1 6

2 . 6 Wea the r ing s t age so fr e indee rb one sf r om t h ef ou rc u l tu ra ll e ve l sa tG rubg raben

2 2

2 . 7 Wea the r ing s t age so fh o r se b one sf r om t h ef ou r c u l tu r a ll e ve l sa tG rubg raben

2 3

2 . 8 R oo te t ch ing o nr e indee rb one sf r om t h e f ou rc u l tu ra ll e ve l sa tG rubg raben

2 8

2 . 9 R oo te t ch ing o nh o r se b one sf r om t h ef o u r c u l tu ra ll e ve l sa tG rubg raben

2 9

2 . 10 B one w e igh to ft a xa f r om t h ef o u rc u l tu ra l l e ve l sa tG rubg raben

3 1

3 . 1 A nnua la n t le rc y c le si nr e indee r /ca r ibou

3

3 . 2 R um inan ta nd m onoga s t r ic d i ge s t ive s y s tem s

3 9

3 . 3 C ompa r i son o fr um ina n ta nd m onoga s t r ic f e ed ing s t r a teg ie s 4 . 1 S ea sona la t t r ibu te so fr e indee ru s ef u li n r e con s t ruc t ing P a leo l i th ic f o od / techno logy m anagemen ts y s tem s 4 . 2 S ea sona la t t r ibu te so fh o r se su sef u li n r e con s t ruc t ing P a leo l i th ic f o od / te chno logy m anagemen ts y s tem s 4 . 3 S ea sona la t t r ibu te so fi b ex u s ef u li n r e con s t ruc t ing P a leo l i th ic f o od / techno logy m anagemen ts y s tem s 4 . 4 S ea sona la t t r ibu te so fa r c t ic f ox u s ef u li n r e con s tr uc t ing P a leo l i th ic f o od / te chno log y m anagemen t s y s tem s

. 5 . 7

4 0

4 5

4 9

5 1

5 3 i v

N umbe ro fi d en t if ied s p e c imen so fh o r s ef r om l e ve l 4a tG rubg r aben

6 7

5 . 8 H i s tog ram so fn umbe ro fi d en t if ied s p e c imen s ( N ISP )a nd m in imum n umbe ro fe l emen t s( MNE ) o fh o r se f r om l e ve l 4a tG rubg raben

6 8

5 . 9 L oca t ion o fc u tm a rk so nh o r se sf r om G rubg r aben

7 1

5 . 10 N umbe ro fi d en t if ied s p ec imen so fh o r s ef r om S agva r( 1930 -31) e xcava t ion s

7 3

5 . 11 N umbe ro fi d en t if ied s p e c imen so fh o r s ef r om S agva r( 1937 38 )e xcava t ion s

7 4

5 . 12 H is tog ram so fn umbe ro fi d en t if ied s p e c imen s ( N ISP ) a nd m in imum n umbe ro fe l emen t s( MNE ) o fh o r se f r om S agva r

7 5

5 . 13 H is tog ram so fn umbe ro fi d en t if ied s p e c imen s ( N ISP )a nd m in imum n umbe ro fe l emen t s( MNE ) o fh o r se f r om P i l i s szan to R ock she l te r

7

5 . 14 S t ran ska S ka la I V , o b l ique v i ew o fp l an o ft h e e a s te rn c oncen t ra t ion o fa r t if ac ts a nd h o r se b one s , e xcava t ion s , 1 985 a nd 1 986

7 8

5 . 15 S t ran ska S ka la I V , o b l ique v i ew o fp l a no ft h e w e s te rn c oncen t r a t ion o fa r t if ac ts a nd h o r s e b one s , e xcava t ion s 1 985 a nd 1 986

7 9

5 . 16 N umbe ro fi d en t if ied s p e c imen so fh o r s ef r om S t ran ska S ka la I V

8 0

F igu re 51 7H i s tog ram s o fn umbe ro fi d en t if i ed s pec imens ( N ISP )a nd m in imum n umbe ro fe l emen ts (MNE ) o fh or se f r om S t ranska S ka la I V

P age

8 1

t e e th o nt h el ef ts how s how n a tu ra lb r eaks . T he 8 6

6 . 15 C ompa ra t ive p e rcen tage so fr e indee rb one s f r om t h er e con s truc ted K ak inya S i te a nd G rubg raben : L eve l 1 13

L eve l 2

14

6 . 17 C ompa ra t ive p e rcen tage so fr e indee rb one s

5 . 19 J aw o fh or se w i th s hea red c heek t oo th embedded w i th in a l ve loa

P age

6 . 16 C ompa ra t ive p e rcen tage so fr e indee rb one s f r om t h er e con s truc ted K ak inya S i te a nd G rubg raben :

5 . 18 Ho r se c heek t e e th f r om G rubg raben . T he t wo f ou rt e e th o nt h er i gh t s how c u l tu ra lb reaks

F igu re

8 6

f r om t h er e con s truc ted K ak inya S i te a nd G rubg raben : L eve l 3 15

6 . 1 A ge p rof i le so fr e indee r mand ibu la rt ee th r e cove red f r om t he f ou rc u l tu ra l l e ve ls a t G rubg raben

8 9

6 . 18 C ompa ra t ive p e rcen tage so fr e indee rb one s f r om t h er e con s t ruc ted K ak inya S i te a nd G rubg raben : L eve l4

16

6 . 2 E rup t ion p a t te rns o fm and ibu la rt e e th o f d ee r

9 1

6 . 19 N umbe ro fi d en t if ied s pec imen so fr e indee r f r om S agva r( 1930 -31) e xcava t ion s2

17

6 . 3 Numbe ro fi d en t if ied s pec imens o fr e indee r f r om L eve l 1a tG rubg raben . S haded a r ea sr ep re sen t a bsen t b ones 6 . 4 Numbe ro fi d en t if ied s pec imen s( N ISP ) a nd m in imum n umbe ro fe l emen ts (MNE ) o fr e indee r f r om L eve l 1a tG rubg raben 6 . 5 N umbe ro fi d en t if ied s pec imen so fr e indee r f r om L eve l 2a tG rubg raben

9 2

6 . 20 Numbe ro fi d en t if ied s pec imens o fr e indee r f r om S agva r( 1937 -38 ) e xcava t ions

18

6 . 21 Numbe ro fi d en t if ied s pec imen so fr e indee r f r om P i l isma ro t

19

9 3 6 . 22 Numbe ro fi d en t if ied s pec imen so fr e indee r f r om P i l isszan to R ockshe l te r

1 20

9 5 -7 . 1 Numbe ro fi d en t if ied s pec imens o fh a re f r om

6 . 6 N umbe ro fi d en t if ied s pec imens ( N ISP ) a nd m in imum n umbe ro fe l emen ts (MNE )o fr e indee r f r om L eve l 2a tG rubg raben

G rubg raben . ( a ) L eve l 1 .( b )L eve l 2 9 6

6 . 7 Numbe ro fi d en t if ied s pec imens o fr e indee r f r om L eve l 3a tG rubg raben

9 7

6 . 8 Numbe ro fi d en t if ied s pec imens ( N 1SP ) a nd m in imum n umbe ro fe l emen ts (MNE )o fr e indee r f r om L eve l 3a tG rubg raben

9 8

6 . 9 N umbe ro fi d en t if ied s pec imens o fr e indee r f r om L eve l 4a tG rubg raben

9 9

6 . 10 N umbe ro fi d en t if ied s pec imen s( N ISP ) a nd m in imum n umbe ro fe l emen ts (MNE ) o fr e indee r f r om L eve l 4a tG rubg raben

1 00

6 . 11 F ragmen ta t ion r a t io o fr e indee rb ones f r om t h ef our c u l tu ra l l e ve ls a tG rubg raben

1 07

6 . 12 P e rcen tages o ff emo ra l ma r row f a t f ound i n c a r ibou t h roughou t t he y ea r

1 08

6 . 13 B ack f a t d ep th ( i n mm ) f ound i nc a r ibou t h roughou t t h ey ea r

10

6 . 14 C ompa ra t ive p e rcen tage s o fr e indee rb one s f r om t h e r e con s truc ted K ak inya S i te a nd t he N e t S i te

12

7 . 2 N umbe ro fi d en t if ied s pec imens o fh a re f r om G rubg raben . ( a ) L eve l 3 .( b )L eve l4

1 25

1 26

Chapter 1 Introduction

GENERAL AND HISTORICAL PERSPECTIVE: This study explores the roles of hunting and animal processing during the Last Glacial Maximum (LGM; 20,000-15,000 BP) in the long term evolutionary development of prehistoric societies. The data base is derived primarily from the open air campsite Grubgraben, Lower Austria. Current methods of zooarchaeology are used to reconstruct the socioeconomics of Epigravettian hunting. An ecological/behavioral model for prey species adaptation to the glacial environment of Central Europe is developed; and Epigravettian decisions, reflected by the types, parts, and quantities of prey carcasses transported, processed, and discarded, are defined. A basic assumption of this study is that the behavior of wild animals from Paleolithic aged archaeological sites resembles that of their modem descendents. Thus, the behavior of extant ungulates and furbearers provides analogs for behavior of their Pleistocene counterparts which can be used to reconstruct mobility, breeding, and birthing patterns; fluctuations in body and coat condition; food preferences; and digestion. Identification and quantification of animal bones documents the economic importance of animal species from any site, given consideration for preservation and recovery. Seasonality, coupled with animal behavior, kinds of animals represented, and representation of parts of carcasses provides infonnation about procurement/processing strategies within a regional framework. Age profiles of recovered ungulate bones can be used to infer hunting strategies and the organizational behavior of human groups. Comparisons of frequencies of prey body parts at residential camps with those at kill sites provide information on field butchering and transport strategies. Green bone fractures, cut marks, and bone frequencies yield valuable insights into butchering practices, marrow extraction, and bone grease processing. Quantification of these characteristics for horse and reindeer reveal the extent to which carcasses of each were manipulated to obtain the greatest return with the least effort. Differential species use reflects both risk reducing and energy maximizing behavior, as well as technological needs and limits. Fragmentation, a reflection of the extent of bone manipulation, and evidence of hearths and/or cooking pits provides infonnation on processing--roasting, drying, and/or increased use of marrow and bone grease. Two sites are the core of the study: Grubgraben, in Lower Austria and Stanska Skala IV in the Czech Republic (Fig. 1.1 ). Grubgraben is an Epigravettian multilevel site, now buried in loess, lies in a protected ravine flanked by two promontories and overlooks the Kamp Valley of the Danube Plain three km to the southwest. This location was an ideal campsite for ancient hunters as it: 1) was situated between an upper plateau and river valley ecotones, 2) was protected from the elements, 3) provided a wide overview of the Kamp Valley below, and 4) was near active springs.

Grubgraben was discovered and initially investigated in the l 870's with occasional, limited excavations from the early years of the twentieth century to the l 960's (Urbanek 1990). Large scale excavations, directed by Anta Montet-White, University of Kansas, between 1985 and 1990) revealed a long stratigraphic sequence beginning with the Last Glacial Maximum (LGM) and including three of the four cultural levels subsequent to that event (Montet- White 1990, Montet­ White and Williams 1994). Four cultural levels, AL1-AL4, were recognized beneath a deep loess cover. These levels were dated between 18,500 and 16,500 BP. The oldest cultural levels, AL3 and AL4, are stratified within a noticeable humic zone marking an interruption in loess deposition, suggesting a milder, but certainly wetter climate between 18,000 and 17,500 BP. AL3 and AL4 contain dense beds of reindeer and horse bones, the apparent result of processing, consuming, and discarding animal carcasses at the site. Younger levels, AL I and AL2, in loess deposited under drier conditions, contain possible storage features, pits, but relatively few bones (Montet-White 1990). Montet-White (1990: 3)) noted, "The first two seasons of excavations have contributed a body of new data about a little · known time period. First and foremost, the discovery of a series of Epigravettian levels posterior to 19,000 BP demonstrated that there were times during the last glacial advance when the area offered sufficient natural resources to sustain groups of Paleolithic hunters." A series of radiocarbon samples from the lowest archaeological level dates between 18,980 and 18,200 BP. People inhabited, if only sporadically, the Middle Danube region during the Last Glacial Maximum. Sedimentological and geomorphological investigations by Haessaerts (1990) and Pawlikowsky (1990) suggested the environment at Grubgraben was relatively humid circa 18,000 based on a series of well developed paleosols. After 18,000 BP, the environment at Grubgraben became increasingly arid. Logan (1990a), investigating adaptability of paleolithic hunting economies, showed a shift from a broad hunting economy of the Gravettian to a narrower economy of the Epigravettian. By comparing the numbers and types of bones from Gravettian aged Willendorf to those recovered from Grubgraben, he showed that humans at Willendorf hunted a diversity of animals. These included: horse, red deer, reindeer, mammoth, bison, aurochs, wolves, foxes, bear, and hares. Epigravettian hunters at Grubgraben specialized in reindeer and horse. Stranska Skala IV, located in the outskirts of Bmo, Czech Republic dates between 17,740 BP and 18,220 BP (Svoboda 1990). This rare site has been interpreted as a specialized horse hunting station where under twenty horses were either driven upslope and over a limestone cliff or were backed against a rock wall (Svoboda 1990, 1991 ). Excavations of nearly 150 square meters between 1985 and 1986, revealed concentrations of limestone blocks, horse bones and lithic artifacts in the uppermost part of the glacial loess. Cultural

Mo rav ian H i l ls

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F ig .1 .1 . G eog raph ic d i s trubu t ion o ft h eC en t ra l E u ropean s i tes i nt h e s t udy s amp le . * —G rubg raben ; 1 =S transka S ka la I V ; 2 =Sagva r ; 3 =P i l is szan to R ockshe l te r , 4 = P i l isma ro t

d h i t mt.%

e l '1

•

i ti ng e

r ema ins w e re c oncen t ra ted i nt wo c i rcu la rf e a tu re s f i ve m e te r s

C ha rac te r ized b y r e la t ive ly l a rge s e t t lemen ts , s i te f u rn i tu re ,

a pa r t . T he w es te rn a ccumu la t ion c ons is ted o f ac i rcu la rc l us te r o f l i me s tone b lock s , h o r se b one s , a nd l i th ic s . Hor se b one s r edepos i ted l o ng i tud ina l ly a l ong t h e s l ope w e re a s soc ia ted

d i s t inc t ive a r t a nd o rnamen t , a nd a n a bundance o fd i sca rded

w i th t h e w e s tern f e a tu re . L a rge , l i me s tone b l ocks a nd h o r se b one sa l so p redom ina ted i nt h ee a s ter na ccumu la t ion .

ar i ch a nd v a r ied f a una l b i oma s s i n c lud ing : mammo th , r ed d ee r , r e indee r , h o r se , i b ex , a u roch , b i son , a nd f u rbea re r s

RESEARCH GOALS :

a nd K l ima

r e f use , t h ey h ave e me rged t o s ymbo l ize t h e U ppe r P a leo l i th ic o fC en tra l E u rope . T he e conom ic b a se o ft h e se s i te s i n c luded

i n c lud ing b ea r , a r c t ic f ox , wo lf a nd h a re ( Mus i l 1 959 , A bso lon 1 977 ,

K l ima

1 969 ,

T hen iu s

1 956 1959 ,

a nd

Koz low sk i 1 986 ) . T h is s t udy f o cuse s o n t h e a dap t ive p roces sess a s soc ia ted w i th E p ig rave t t ian r e cove red

b ehav io r

f r om

r e f lec ted

G rubg raben .

i n t he

a n ima l

P rocu remen t ,

F o l low ing

r ema in s

t h e

l a s t

g l ac ia l

a dvance ,

a r ound

1 8 ,000

B P

( CL IMAP 1 976 , 1 981) , t h e c l ima te i nC en t ra l E u rope b ecame

t r anspor t ,

mo re a r id wh ich d ep le ted a va i lab le a n ima l r e sou rce s .

A

b u tche r ing , ma r row a nd b one g rea se manuf ac tu re , a nd p roce s s ing s t ra teg ies a r e i n ve s t iga ted /ana lyzed . T he f ou r

p auc i ty o f E p ig rave t t ian a nd e a r ly Magda len ian

c u l tu ra l l e ve ls a tG rubg raben c on ta ined t housands o fa r t if ac ts i n c lud ing a pp rox ima te ly 4 1 ,000 c omp le te a nd f r agmen ted

d ur ing t h e m ax imum g l ac ia l a dvance , t h e r eg ion w a s e i the r

s i te s i n

C en t ra l E u rope h a s b een i n terp re ted a s a n i n d ica t ion t h a t

a n ima l b one s , a ppa ren t r e f use o f h uman t r anspor ta t ion o f

o n ly i n te rm i t ten t ly t r ave l led o r c omp le te ly a bandoned . H uman g roup s a nd t h e ir p rey s ough t mo re f a vo rab le e nv i ronmen ts t o

a n ima l r e sou rce s . F requenc ies o f b ones s how r e indee r a nd h o r se w e re t h ec ons is ten t ly f a vo red p rey o fh un te r sa tt h is s i te . O n t h e o t he r h and , r ema ins o fm ammo th , b i son , a uroch , a nd

t h e s ou th a nd e a s t ( Hahn 1 976 , G amb le 1 983 , K oz low sk i

i b ex a r e r e la t ive ly s pa r se , s ugges t ing t h a t t h e se a n ima ls w e re

a nd Wen ige r 1 990 ) . K oz low sk i a nd Koz low sk i ( 1979 : 3 9-40 )

e i the r s e a sona l ly o r who l ly a bsen t i nt h e a r ea d ur ing h uman

c ompa red t h e e a r l ie r G rave t t ian e conomy t o t ha t o f t h e

o ccupa t ion . F urbea re r s , s uch a s f ox a nd h a re , a r e f ew i n n umbe r , b u t t h e ir n umbe r sv a ry f r om l e ve l t ol e ve l .

E p ig rave t t ian . D ur ing t h e G rave t t ian , l a rge h e rds o f s t eppe a n ima ls " . .p rov ided a n e a s i ly a ces s ib le s ou rce o f f ood a nd t hus p roduced l a rge s e t t lemen ts w i th a s em i -nomad ic w ay o f

PREV IOUS ENV IRON MENTAL AND

l i f e ." A s c l ima te d e te r io ra ted , " . . the r ep lacemen t o ft h e s t eppe

1 986 , J och im 1 987 , S oi f e r a nd G amb le 1 990 , S voboda 1 990 ,

ARCHAEOLOG ICAL I NVEST IGAT ION

b y t und ra o r e ven p o la r d e ser t " a l ong w i th " . .exces s ive

E nv i ronmen t p l ayed a k ey r o le i nd e te rm in ing s t ra teg ie s a nd

u t i l iza t ion o fa n ima l r e sou rce s " g ene ra ted a n e m ig ra t ion f r om

a dap ta t ion s o f h un te r /ga the re r s i n E u rope d ur ing t h e P a leo l i th ic . I n t he l a rge s t s ense , c l ima te s t ruc tured h uman

C en t ra l E u rope t ot h ee a s t .

d ec is ion s . I tc on t ro l led t h e t ype a nd a moun t o ff ood a va i lab le

G amb le ( 1983 : 2 06 -207 ) a rgued t h a t u ng lac ia ted a r ea s o f

a nd e xp lo i ta t ion p a t ter n s . E nv i ronmen t a l so l i m i ted c ho ices o f

n or ther na nd c en t ra l E u rope , e spec ia l ly b e tween a dvanc ing i c e

h uman o ccupa t ion s i te s , s e a sona l p opu la t ion s i ze , movemen t s ,

f r on ts , s uf e red i n tense p e r ig lac ia l c ond i t ions . Ac omb ina t ion

a nd m ig ra t ions d ur ing p er iods o fm a jo rc l ima t ic c hange s . A t a

o f i n c rea s ing c o ld ,

s ma l le r

g row ing s e a sons , l a te s now c ove r , a nd p e rmaf ros t a l te red t h e

s c a le ,

v ege ta t ion ,

s pa t ia l

d i s tr ibu t ion

t empe ra ture ,

o f

p rec ip i ta t ion ,

s o i ls ,

e l eva t ion ,

d ra inage ,

l e s s d i rec t

s o la r r a d ia t ion ,

s hor tened

w ind

v ege ta t ion u pon wh ich h e rb ivo re s d epended . T he se f a c to r s

d i rec t ion , a nd r ock t ype s d ef ined t h e i mmed ia te u se o f t h e l a ndscape ( J och im 1 976 ) .

r a nge s , a nd i n f luenced t h e a nnua l movemen ts o f a n ima ls .

F o l low ing t h e D enekamp I n te r s tad ia l , ( 29 ,000 BP ) t h e c l ima te

t h e se mo re s pec ia l ized e nv i ronmen ta l c ond i t ions b egan t o

i nE u rope d e te r io ra ted t o t h e d ry , c o ld e p isode r e f e r red t o a s t h e LGM . I c e s hee ts a dvanced s ou th a c ross t h e Nor ther n

a n ima l r e sou rce s b ecame l e s s p red ic tab le . A l though s ou thern

r e duced b i oma s s p e r u n i t a r ea , a l te red t h e s i ze o f a n ima l D ive r s i ty o fl o ca l a nd r eg iona l a n ima l c ommun i t ie s c hanged a s e me rge . E xp lo i ta t ion r i sks f o r g roups o fh un te r s i n c rea sed a s

E u ropean P l a in a nd d own t h e s l ope s f r om t he A lps . Ef ec ts o f

a nd w e s ter n E urope s uf e red o n ly mod if ied e f ec ts ,

t h is g l ac ia l p er iod v ar ied a c ros s E u rope w i th t h e g rea te s t

wou ld e xpec t , i nc en t ra l E u rope , a s t h e c l ima te d e te r io ra ted

i mpac t o n C en t ra l

E u rope ,

G e rmany ,

g eog raph ic a r ea e x is ted a s an a r row ,

a nd

A us tr ia .

e a s t -we s t ,

"We

T h is

w i th t h e a dvance o ft h e c on t inen ta l i c e s hee ts , a nd r e sou rce s

i c e -f ree

b ecame m o re u npred ic tab le d ue t ot h e ir g rea te r mob i l i ty . . . tha t

c o r r ido r b e tween t h e S cand inav ian a nd A lp ine i c e s hee ts

s e t t lemen t wou ld c ea se " ( Gamb le 1 983 : 2 06 -207 ) .

( Pose r 1 948 , K a ise r 1 960 , CL I MAP 1 976 ) . E x treme c ond i t ions a s soc ia ted w i th t h e L a s t G lac ia l Max imum

K oz lowsk i a nd K oz lowsk i ( 1979 : 4 2 ) a dded t h a t s hor t t e rm

p r ed ica ted s i gn if ican t s h i f ts i ns e t t lemen ts t h roughou t E u rope

f l uc tua t ions

w i th t h em o s t n o tab le c hange so ccur r ing i nC en tra l E u rope .

a me l io ra ted c l ima te i n E a s ter n a nd We s te rn E u rope , b u t i n t roduced " . . l i t t le , i f a ny , c l ima t ica l i mp rovemen t t o t h e

T he a r chaeo log ica l b a se f o r t h e p r eh is tory o fC en t ra l E u rope h a s b een d om ina ted b y as e r ie s o f s t ra t i f ied s i tes ( F ig . 12 )

s uch

r eg ion o f C en t ra l

a s

t h e

L auge r ie -La scaux

i n te r s tad ia ls

E u rope . . a lmos t t h e w ho le o f C en t ra l

E urope n or th o ft h e D anube c ons t i tu ted a n u n inhab i ted a r ea ,

d a ted t o t h e e a r l ie s t p ha se s o f t h e l a s t g l ac ia l ( 28 ,000 a nd 2 0 ,000 B P ) ; i n c lud ing W i l lendorf ( Fe lgenhaue r 1 956 -1959 ) ,

w h ich i s o la ted We s tern f r om E a s ter n E u rope ."

D o ln i

V es ton ice

( K l ima

1 963 ,

1 969 ,

1 981) ,

A l though

c l ima t ic a me l io ra t ions f a vo rab le t o s e t t lemen t o ccur red i nt h e

P redmos t

C a rpa th ian B a s in ( Sagva r ian c u l tu re ) , s i te s w e re n e i the r a s

( Ab so lon a nd K l ima 1 977 ) , a nd P av lov ( Sk lena r 1 977 ) . T hese

s t ab le n o ra sn ume rous a st h e i r e a r l ie r G rave t t ian c oun te rpa r ts

s i te s a r e d e scr ibed i n e ve ry i n troduc tory t e x tbook o n E u ropean p r eh is tory , a nd h av ing b een e x tens ive ly e xcava ted

i n Mo rav ia a nd A us t r ia .

a nd s t ud ied , r e co rd t h e l o ng s t ra t ig raph ic c h rono logy o f t h e

R ecen t r e sea rch i sc hang ing c l as s ic v i ew s o ft h e e nv ironmen t

G rave t t ian . T he ir p resence i n f e r s as t ab i l i ty o f s e t t lemen t

a nd p r eh is to ry o f C en t ra l E u rope d ur ing t h e L a s t G lac ia l

d u r ing

e a r l ie s t

p ha se s

o f

t h e

S econd

P len ig lac ia l .

Max imum . Wo i l la rd ( 1978 ) , a na lyz ing p o l len f r om G rande 3

e fej t oe „ , •

lev e l 101

e 00 ) A Z d in l y t i l l t ue e in `P R " l eT i n •

U i j i

W il

'n

4" v i r

i ' F I

F ig . 1 . 2 . Map o fC en tra l E u rope s how ing t h e l o ca t ion o fm a jo r s i te sd i scus sed i nt h e t e x t 1 =W i l lendor f ; 2 =Do ln i V e s ton ic ; 3 =P redmos t ; 4 =Pav lov ; 5 =Ku lna ; 6 =Mada ra s ; 7 =Kada r ; 8 =Zupanov S podmao l ; 9 =A rka ; 1 0=Bodrogke re s tu r ; 11 =Spadz is ta ; 1 2=Temna ta , 1 3=Ovca J ama ; 1 4= M i lov ice

4

P i le , g ene ra l ly i d en t if ied g l ac ia l e p isode s a s c o ld , d ry p ha se s

e t hnoa rchaeo logy the s t udy o f mode rn h un te r /ga the re r s t o

p r ima r i ly d om ina ted b y g ra s se s a nd s edges a tt h e e xpense o f

p rov ide c ompa ra t ive f r amewo rks f o r i n terp re t ing p a s t h uman

a rbo rea l s pec ie s . V a loch ( 1989 ) , e s t ima ted t h a t t h e g l ac ia l a dvance b egan a pp rox ima te ly 2 7 ,000 BP b a sed o n c ha rcoa l a nd p o l len s amp le s f r om Mo rav ia . C ha rcoa l o f p i ne a nd f i r

b ehav io r , 2 )env ironmen ta l s t ud ie s , 3 )eco log ica l a nd a dap t ive s t ra teg ies o ft h e g ame a n ima ls u sed b y h un te rs a tt h e s i te , a nd

r e cove red

a l te r b one s a nd o the r m a ter ia ls a f te r t h e d ea th o f a n a n ima l . T he r e sea rch a nd a na lys is i so rgan ized a s f o l lows . C hap te r

f r om

K u Ina

C ave

i n d ica te

a m i ld

b e tween 2 3 ,000 a nd 2 2 ,000 BP ( Va loch i n terp re ta t ion i s s uppor ted b y e v idence d eve lopmen t ,

o sc i l la t ion

1 989 ) . T h is o f p a leoso l

d a ted a t 2 3 ,000 B P , a t S padz is ta ,

P o land

4 )t aphonomy the s t udy o fc u l tu ra l a nd n a tura l p roce s se st h a t

Two r ev iew s r e sea rch m e thodo logy . F i e ldwo rk , i n c lud ing e xcava t ion t e chn ique s , b one r e cove ry , p roven iance l a be l l ing ,

( Koz low sk i e t a l . 1 974 ) . A s e cond m i ld o sc i l la t ion , d a t ing

a nd l a bora to ry p repa ra t ion o ff a una l e l emen ts f o r a na ly s is a r e

b e tween 1 9 ,000 a nd

1 8 ,000 BP , c ha rac te r ized b y h um ic

d escr ibed . A na ly t ica l p rocedu re s , i n c lud ing a s ta nda rd ized

h or izons a nd /o r p o l len d om ina ted b y p i ne , j un ipe r , l i nden , s edge s a nd g ra s s , w e re i d en t if ied a t l o e ss s i te s . T hese s i te s i n c lude : G rubg raben , L owe r A us tr ia ( Hae s sae r ts 1 990 a nd A .

Mo rpho log ic a t t r ibu te s i n c lude i d en t if ica t ion o ft axon , s ke le ta l

r e cord ing

s y s tem

o f a t t r ibu te s ,

a r e

e xp la ined

i n d e ta i l .

e l emen t , p a r t , s egmen t , s i de , a nd f u s ion r a te so fe p iphy se s .

L e ro i -Gou rhan c i ted i n Mon te t - Wh i te 1 990 ) , S agva r i n t h e w es t c en t ra l s e c t ion o ft h e C en t ra l E u ropean B a s in ( Gabo r i

C u l tu ra l a nd n a tu ra l t a phono rn ic p roce s se s wh ich i n f luenced

a nd G abor i 1 957 , G abo r i -Csank 1 978 ) , P i l isma ro t n ea r t h e

s i te

Danube B end ( Dobos i e t a l . 1 983) , Mada ra s i n C en tra l Hunga ry ( Dobos i 1 989 ) , l e ve l 2a tK ada r ( Mon te t - Wh i te e ta l .

mod if ica t ion s : t h e r e cogn i t ion o fe l emen t r e p re sen ta t ion , c u t

f o rma t ion

a t

G rubg raben

a r e

d ef ined .

H uman

ma rks , t ypes o fb reaks , a nd e v idence o fb u rn ing a r e d e scr ibed .

1 986 ) , Z upanov S podmo l ( Oso le 1 97 1) n ea r t h e S ava R ive r ,

A ma jo r t a sk o ft h is a na ly s i s i st o " pee l a way " f r om i n d iv idua l

a nd S t ran ska S ka la I V , t o t he n o r th i n Mo rav ia ( Svoboda

b ones , a nd t he b one a s semb lage a s aw ho le , t he t a phonom ic

1 990 ) . A t h i rd c l ima t ic a me l iora t ion , d a ted b e tween 1 7 ,500

o ve rp r in t t o e xpo se h uman a c t iv i t ie s a t G rubg raben . T he se

a nd 1 7 ,000 BP i s a na logous t o t h e L a scaux i n w e s tern E u rope . E v idence o f t h is e ven t o ccu r s i n u ppe r l e ve ls a t

n a tura l mod if ica t ions i n c lude w ea ther ing , g naw ing , a nd r oo t

S agva r ( Gabor i a nd G abor i 1 957 , G abo r i -Csank 1 978 ) , a nd l e ve l 2a t G rubg raben ( Hae ssaer ts 1 990 ) . P o l len o f p i ne ,

p os s ib le ,

b eech , a nd a sh o ccu r a ts i te s i n S loven ia ( Mon te t - Wh i te a nd

b y p l ac ing t hem o n ag r id .

K oz low sk i 2 1 ,000

1 983) .

a nd

e t ch ing . A dd i t iona l ly , a l b one s w e re w e ighed , a nd w hen m ea su red ,

f o l low ing

v on

d en

D r ie sch

( 1976 ) .

U n iden t if ied b one f r agmen t s w e re m ea su red t ot h e n ea re s t c m

T he f a una l a s semb lage , d a ted b e tween

1 6 ,000

B P ,

f r om

P i l is szan to

R ockshe l te r ,

Ad i scuss ion o ft h ee co logy o fa n ima ls r e cove red f r om t h e s i te

H unga ry , s ugge s ts t h ee nv i ronmen t o fC en tra l E u rope w a s n o t

i s p re sen ted i n C hap te r T h ree . S ea sona l m ig ra t ion a nd /o r

a mono l i th ic " co ld s pe l l ," b u t r a the r ac omp lex o f v a r ied

movemen t

c l ima t ic p ha se s d ur ing t h e LG M . A t t h is s i te , f a una i n c lude a m ix tu re o fa r c t ic , s uba rc t ic , b o rea l , a l p ine , f o re s t , a nd s t eppe

m echan isms , a nd f ood p r ef e rence s a nd d i ge s t ion a f ec ted

s pec ie s . I n ar eg iona l r e eva lua t ion , Mon te t - Wh i te ( 1994 : 4 91) d e sc r ibed t h e g l ac ia l o fC en t ra l E urope a s " . .one o fl o ca l ized

r e indee r , h o r se , a nd i b ex a dap ted t o t h e i n c reas ing a r id i ty o f

p a t ter n s ,

b reed ing ,

b i r th ing

a c t iv i t ie s ,

s pec ies a dap tab i l i ty t o ac hang ing e nv i ronmen t

d ef ense A l though

t h e L a te G lac ia l , m ammo th s a nd l a rge b ov ids w e re f o rced t o

a nd s h if t ing , r e la t ive ly h um id n i che s w i th in a g ene ra l ly d r ie r

e m ig ra te f r om C en tra l E u rope . A mode l o f a dap ta t ion b y

r e g ion ."

r e indee r , h or se , a nd i b ex t o t h e i n c rea s ing a r id i ty o ft h e L a te G lac ia l i nC en t ra l E u rope i sp roposed .

D om inance o f af ew e x ten s ive ly s t ud ied s i te s i nt h e l i t e ra tu re , s amp l ing b i a s i n r eg iona l s t ud ie s , a nd e ro s iona l a c t iv i t ie s l e ad ing t os t ra t ig raph ic h i a tuse s i nk ey a r ea s , h ave s kewed o u r i n te rp re ta t ion o ft h e a r chaeo log ica l r e co rd i nt h is l i t t le k now a r ea ( Mon te t - Wh i te 1 994 ) . Ma ss em ig ra t ion f r om t h e a r ea h a s b een c ha l lenged a nd r e eva lua ted i nl i gh t o fr e cen t e xcava t ion s a nd r e g iona l s u rveys i n C en tra l E u rope . A s e r ie s o f s i tes , d a t ing t o t h e h a r she s t p ha se s o f t h e P l en ig lac ia l ( 2 3 ,000 -

C hap te rF ou r i sd evo ted t o ad i scus s ion o fh uman a dap ta t ions t o t h e s h if t ing f a una l b i oma s s d e scr ibed i n C hap te r T h ree T heore t ica l a spec t s o fh un t ing e conomy a r e d i scus sed i nl i gh t o f . 1 ) a n ima l s i ze a nd d i s tance f r om k i l l t o p l ace o f c onsump t ion (Wh i te 1 952 , 1 953 , 1 954 ; P e rk ins a nd D a ly 1 968 ; O 'Conne l l e t a l . 1 988 ; Me tca lf e a nd J one s 1 988 ) , 2 ) a n ima l c ond i t ion b a sed o n s e x , a ge , a nd s e ason ( Spe th a nd S p ie lmann 1 983 , S pe th 1 987 ) , 3 ) a va i lab i l i ty o f m a r row

1 5 ,000 B P ) , h ave b een r e co rded . T he se i n c lude : P i l isma ro t P a la re t , S agva r , A rka , a nd B od rogke re s tu r i n H unga ry

/ g rease r e sou rce s ( B inf o rd 1 978 , Me tca lf e a nd J one s 1 988 )

( Dobo s i

a nd , 4 ) h i ghe s t r e tu rns f o r l e a s t e f o r t /r isk ( Z ipf 1 949 , J och im

1 981 ,

1 991a ,

1 99 1b ;

Dobos i a nd V o ros

1 986 ;

D obos i e t a l . 1 983) ; S padz is ta i n P o land ( Koz low sk i e t a l .

1 976 , Sm i th a nd W in te rha lde r 1 992 ) .

1 974 ) ; T emna ta i nB u lga r ia ( Koz lowsk i e ta l . 1 992 ) ; K ada r i n B o sn ia ( Mon te t - Wh i te e ta l1 986 ) ; Z upanov ( Oso le 1 97 1) a nd

A r e shuf f l ing

O vca J ama i nS loven ia ( O so le 1 977 ) ; a nd S t ranska S ka la I V

i mpac ted

( Svoboda 1 990 ) a nd M i lov ice ( O l iva 1 988 ) i n t h e C zech R epub l ic .

mob i le

AND

ORGAN IZAT ION

OF

TH IS

t o

e xp lo re

I NVEST IGAT ION : p rov ide s

a u n ique

o ppor tun i ty

e conom ic l i f eways o fh un te r s a t ac amp r e pea ted ly o ccup ied d u r ing

t h e

m e thodo log ica l

L a te

G lac ia l

c r i ter ia

f o r

p er iod . t h is

T heo re t ica l s t udy

i n c lude :

f a una l

b i oma ss

n ece s sa r i ly

u ngu la te s

w hose

m ig ra t ion

A dd i t iona l ly ,

s e a sona l ly

p a t te rns b ecame f l uc tua t ing

l e s s

p hy s ica l

f e a tu re s : h a ir a nd s k in q ua l i ty , f a t /mar row d epos i t ion a nd q ua l i ty , a nd a va i lab i l i ty o fs e a sona l a n t le r s p l ayed ak ey r o le i n t h e

G rubg raben

a nd

a dop ted a mob i le l i f e s ty le a s t h ey p reyed u pon t h e m o re p r ed ic tab le .

A PPROACH

i n f o rage

h un te r s i n C en t ra l E u rope . E p igrave t t ian h un te r s

s e lec t ion

o f s i te s ,

a nd

p rocu remen t

a nd

p roce s s ing

s t ra teg ie s ( B inf o rd 1 978 , S pe th 1 983 , S pe th a nd S p ie lmann 1 983) .

a nd 1 )

S ubsequen t c hap te r s c on ta in t h e r e su l ts o f t h e a na lys is o f 5

h o r se ( Chap te r F i ve ) , r e indee r ( Chap te r S i x ) , a nd a n ima ls l e s s c ommon ly f ound a tt h es i t e( Chap te r S even ) .

s t ra teg ie s a s soc ia ted w i th i b ex , a nd t h e f u r a nd m ea t b ea r e r s : f ox a nd h a re . E xp lo i ta t ion o f f u rbea re r s r e qu i red e qu ipmen t s uch a s s na re s a nd /o r t r ap s a s w e l l a s t r ad i t iona l h un t ing

I nC hap te rF i ve , m eage r e t hnog raph ic s ou rce s c omb ined w i th i n f o rma t ion a bou tt h eb ehav io ro fw i ld h o r se s p rov ide d a ta f o r r e cons t ruc t ing a nc ien t h o r se h un t ing /p roce s s ing t e chn ique s .

e qu ipmen t a nd w a s ah i gh ly s e a sona l a c t iv i ty . I b ex h un t ing m ay h ave b een i n teg ra ted i n to t h e a nnua l e xp lo i ta t ion o f r e indee r . A s r e indee r m ig ra ted t o c a lv ing g rounds i nh i ghe r a l t i tude s , h un te r s i nt h e v i c in i ty m ay h ave u nder taken s ho r t h un t ing f o ray s t o p rocu re i b ex a th i ghe r e l eva t ion s . P r io r t o t h e LGM , m ammo th h ad e m ig ra ted f r om C en t ra l E u rope .

F requenc ie s o f b one s r e cove red f r om t h e k i l l s i te S t ranska S ka la I V ( Mo rav ia ) a r e c ompa red w i th h o r se b one s f r om E p ig rave t t ian c amps . F rac tu red b one s , wh ich m ay b e

I vo ry f ound a t G rubg raben p robab ly s c avenged f r om a n ima ls l o ng d ead .

i n d ica t ive o fe i the r d i smembe rmen t a nd /o r m a r row e x t rac t ion a r e c ompa red a mong t h e se s i te s . A mode l f o r h o r se h un t ing , f i e ld p roce s s ing , a nd t r an spor tt oc amp i sd eve loped .

r e p re sen t s

t u sks

F i na l ly , i n C hap te r E igh t , r e su l ts o f t h e a na ly se s a r e s umma r ized a nd r ev iewed . A g ene ra l ized mode l f o r E p ig rave t t ian h un t ing /p roce s s ing s t ra teg ie s i s p ropo sed i n l i gh t o ft h e e nv i ronmen ta l i mpac t s u s ta ined i nC en t ra l E u rope d u r ing t h eL a s tG lac ia l Max imum .

I n C hap te r S i x , r e cen t e t hnog raph ic s t ud ie s o ft h e N unam iu t , mode rn c a r ibou h un te r s ,(B inf o rd 1 978 , S p ie s s 1 979 , E n loe 1 993) p rov ided t e s tab le m ode l s f o r r e indee r h un t ing a nd p roce s s ing s t ra teg ie s d u r ing t h e U ppe r P a leo l i th ic . T e s ts i n c lude : 1 ) a na ly s is a nd q uan t if ica t ion o f b one s p l in te r s , p roduc t s o fb u tche r ing a nd m a r row e x t rac t ion ; 2 ) p roce s s ing f o r b one g rea se i n wh ich a r t icu la r e nd s o f l o ng b one s a r e e f ec t ive ly r emoved f r om t h eb one a s semb lage ; a nd 3 )s t o rage s t ra teg ie s , i n c lud ing m ea t d ry ing a nd c ach ing . T he l a rge n umbe r o fr e indee r b one f r agmen ts r e cove red f r om l e ve ls 3 a nd 4 m ay r e p re sen t e i the r an umbe r o f s e pa ra te h un t ing e p i sode s o r af ew m a s s ive k i l l s . E thnog raph ic wo rk ( En loe 1 99 1) s ugge s ted t h a tt wo s ubs is tence s t ra teg ie sm ay b e r e f lec ted b y t h e s i ze s a nd s h ape s o fb one f r agmen t s : 1 )m a ss p roce s s ing o fm a r row f o r s t o rage a nd f u tu re u se ( a s soc ia ted w i th m a s s k i l l s ) a nd 2 ) b one c r ack ing a nd m a r row i n c iden ta l t om ea t c onsump t ion a tm ea ls ( a s soc ia ted w i th s epa ra te , s ma l l h un t ing e p i sode s ) . A no the r c on s ide ra t ion i st h e v a r iab i l i ty o f t h e t o ta l m a ss o f b one b e tween o l de r l e ve ls a nd y ounge r l e ve ls B one c oncen t ra t ion s a r e h i gh i n o l de r l e ve ls b u t u p t o 1 0 t ime s l owe r i ny oung l e ve ls _ T h is c ou ld r e su l t f r om : 1 )c hange s i n r e indee r p opu la t ion s a nd m ig ra t ion h ab i t s a s soc ia ted w i th a n i n c rea s ing c o ld /dry c l ima te m ak ing t h em l e s s a cce s s ib le t o h un te r s , 2 ) d if e ren t s pa t ia l u se o f t h e s i te o ve r t i me , 3 ) a s e a sona l c hange i nt h eu se o ft h e c amp s i te t h rough t ime , o r4 ) ac omb ina t ion o ft wo o r mo re o ft h ea bove f a c to r s . R e indee r a ge a nd s i te s e a sona l i ty a r e s t ud ied b y a na lyz ing t oo th e r up t ion p a t te rn s a nd w ea r ( Ha r low a nd D eFoo r 1 962 a nd S p ie s s 1 979 ) Q uan t i ta t ive a na ly s i s o fb one s p l in te r s a nd d i agno s t ic b one f r agmen ts m igh t i n d ica te a d if e ren t p roce s s ing s t ra tegy o fr e indee r c a rca s se s a s c l ima te c hanged . I fm ea t a nd m a r row w e re s t o red a st h e c l ima te d e te r io ra ted o r a s s e a sona l o ccupa t ion c hanged , t h is m igh t b e r e f lec ted i n d if e ren tb u tche r ing a nd m a r row e x t rac t ion p a t te rn s . I n C hap te r s F i ve a nd S i x , h un t ing /p roce s s ing s t ra teg ie s a t G rubg raben a r e c ompa red w i th b ehav io r s p ropo sed f o r c on tempo raneous s i te s i n C en t ra l E u rope F requenc ie s o f d if e ren t a na tom ica l p a r t s o f r e indee r a nd h o r se f r om G rubg raben a r e c ompa red w i th t h e s ame f r equenc ie s o b ta ined f r om H unga r ian o pen a i r s i te s S agva r ( Dobo s i a nd V o ros 1 986 ) , P i l isma ro t ( Dobo s i e t a l . 1 983 ) , a nd t h e P i l is szan to R ock S he l te r( Dobos i a nd V o ro s 1 986 , 1 987 ) . C hap te r S even i s ad i scu s s ion o fp rocu remen t a nd p roce s s ing 6

Chap ter 2 EVALUAT ING THE DATA

F rom 1 986 1990 , t h e a r chaeo log ica l t e am o f Mon te t - Wh i te e xcava ted 9 2 s qua re m e te r s a t G rubg raben ( Mon te t - Wh i te

o fo ve r4 0 ,000 a n ima lb one sr e cove red f r om G rubg raben .

1 990 , Mon te t - Wh i te a nd W i l l iams 1 994 ) . F ou r c u l tu ra l l e ve ls

I NFOR MAT ION RECORDED I N THE DATA BASE :

w e re r e cogn ized

( f rom y ounges t ,

AL1

t o

o l de s t ,

AL4 )

b enea th ad eep l o es s c ove r( F ig . 2 . 1) . T he se l e ve ls w e re d a ted b e tween 1 8 ,500 a nd 1 6 ,500 BP . T he o l de s t c u l tu ra l l e ve ls ,

C a ta logue Numbe ra nd P roven ience :

AL3 a nd AL4 , a r e s t ra t if ied w i th in a n o t iceab le h um ic z one

T he c a ta logue n umbe r ( obse rva t ion A ; s e e A ppend ix 2 . 1) o f

ma rk ing a n i n ter rup t ion i n l o es s d epos i t ion ,

s uge s t ing a

e ach s pec iman wa s f i r s t r e co rded . T h is i n c luded t he n ame o f

m i lde r , b u t c e r ta in ly w e t te r c l ima te b e tween

1 8 ,000 a nd

t h e s qua re a nd t h e n umbe r o n t h e i n d iv idua l f a una l e l emen t .

1 7 ,500 BP . AL3 a nd AL4 c on ta ined t h e g rea te s t d en s i ty o f a n ima l b one s a nd a r t if ac ts . F ea tu re s a s soc ia ted w i th t h ese

O b se rva t ion B w a s t h e r e co rd o f t he a r chaeo log ica l l e ve l i n w h ich t he f a una l e l emen t w as r e cove red . 1 , 2 , 3 , o r 4 .

c u l tu ra l l e ve ls i n c lude b one -f i l led , b a s in s haped p i ts c u t i n to t h e u nde r ly ing l o e ss ( Mon te t - Wh i te 1 990 : 6 3 ; Mon te t - Wh i te a nd W i l l iam s 1 994 ) . D ense b eds o fr e indee r a nd h o r se b one s i nt h e se e a r l ie s t l e ve ls a r e t he a ppa ren t r e su l t o f p roce ss ing ,

P roven ience i n f o rma t ion w a s r e co rded f o r o bserva t ion s C

c on sum ing , a nd d i sca rd ing mu l t ip le a n ima l c a rca sse s a t t h e

T axon a nd E lemen tI den t if ica t ion :

t h rough E . C o lumns f o r Xa nd Y r ep re sen t h or izon ta l p o in t p roven ience a nd Z i st he e l eva t ion f r om d a tum .

s i te . I nc on tras t , y ounge rl e ve ls , ( AL I a nd AL2 ) a re l o ca ted i n l o es s d epos i ted u nde r d r ie r c ond i t ion s . T he se c u l tu ra l l e ve ls

B one s w e re i d en t if ied u s ing c ompa ra t ive c o l lec t ions h oused i n

c on ta in s t one p avemen ts ; w e l l d ef ined h ea r th s ; c oncen t ra t ions

t h e Museum o fA n th ropo logy a nd Museum o fN a tu ra l H is tory

o f o ch re ; p os s ib le s t o rage f e a tu re s ; a nd m anupor ts u sed a s

a tt h e U n ive r s i ty o f K ansa s , a nd t he O s teo logy L abora tory ,

h ea r th s t one s , p o t b o i le r s , a nv i ls , h amme r sa nd g r ind ing s t one s

K ansa s S ta te U n ive r s i ty . As ke le ton o fa n a du l t m a le c a r ibou

( F ig s . 2 . 2 a nd 2 . 3) . R e la t ive ly f ew b one s s ugge s t al e s s i n tense

w a s

u t i l iza t ion o fa n ima l c a rca s ses i nt h e y ounge r l e ve ls ( Mon te t -

U n ive r s i ty o f A la ska . Ia l so u sed c ompa ra t ive s ke le ton s o f

Wh i te 1 990 , Mon te t - Wh i te a nd W i l l iams 1 994 ) .

c on t r ibu ted

b y

t h e

D epar tmen t

o f

A n th ropo logy ,

h o r se , f ox a nd d ee r i np e r sona l c o l lec t ion s . L i te ra ture u sef u l i n " i n i t ia l i d en t if ica t ion o fb one s i n c luded : G i lbe r t ( 1980 ) , S chm id

T he s i te w a s d i v ided i n to 2X 2m e te r s qua re s . A l l l e ve ls w e re e xcava ted u s ing t r owe ls a nd s ma l l h and h e ld t oo ls f o r f i ne

( 1972 ) , a nd G e t ty ( 1975 ) .

wo rk . I nd iv idua l b one s w e re a s s igned p roven ience n umbe r s

E lemen ts w e re a na lyzed b y l e ve l b eg inn ing w i th t he mos t

b a sed o n s pec if ic c oo rd ina te s i n l e ve ls 1 a nd 2 . A s b one

r e cen t l e ve l 1 .A l l b one s w e re a na lyzed u nde r b r igh t l i gh t a nd

d ens i ty i n c rea sed i n L eve ls 3 , e ach 2 X 2 m s qua re w as

w i th a t l e a s t 1 0X m agn if ica t ion u s ing aj ewe le r 's l e ns . E ach

s ubd iv ided i n to s ma l le r 5 0 X 5 0cm s qua re s . B one s r e cove red

e l emen t

f r om e ach 5 0 X5 0cm u n i t w e re b agged t oge the r a nd g i ven a

( observa t ion

p roven ience n umbe r . A l l p roven ience i n f o rma t ion w a s r e co rded o n l a p top c ompu te rs i nt h e f i e ld . F r iab le e l emen ts ,

R ang if e r ) , t y pe o fb one ( ob serva t ion H ; e . g . h umerus ) , p or t ion o f b one

i n c lud ing

b l ade s ,

( observa t ion J ; e . g . a n te r io r ) a nd s i de ( ob se rva t ion K ; l e f t ,

m and ib le s , m ax i l lar ie s w i th t e e th , a nd a n t le r s w e re p l a s te r j a cke ted a nd l a be l led .

r i gh t , a x ia l , o r n onde te rm inan t ) . R e la t ive m a tur i ty o ft h e l o ng

m ammo th

i v ory ,

s capu lae

w i th

i n tac t

w a s

a s s igned F ;

e . g .

( observa t ion

t o

b one ) ,

a g ene ra l ized t a xon

I ; e . g .

c l a ss

( obse rva t ion

p rox ima l

e nd ) ,

c a tegory G ;

e . g .

s egmen t

b one s b a sed o ne p iphy sea l u n ion a tt h e p rox ima l o r d i s ta l e nds w a sr e co rded i no bserva t ion s La nd M ( s ee A ppend ix 2 . 1) .

F o l low ing e ach d ay 's e xcava t ion , b one s w e re w a shed , d r ied , a nd l a be l led i nt h e f i e ld l a b a s t ime a lowed . Wa shed , l a be l led

C e r ta in b ones o f t h e s ke le ton : r i b s haf t s , f r agmen ts o f l o ng

b one s , a l ong w i th u nwashed b one s , w e re s h ipped t o t h e

b one s ha f ts , s c apu la f r agmen t s w i thou t t h e g l eno id , v er teb ra l

Museum o fA n th ropo logy , U n ive r s i ty o fK ansa s w he re b one

c en tra l a ck ing t h e n eu ra l a r ch , v er teb ra l s p ine s , i mma tu re

p r epa ra t ion w a sc omp le ted . J acke ted b one s w e re c l eaned w i th

b one s m inus e p iphy se s d o n o t l e nd t h emse lve s t o g ene r ic

s of t t oo thbrushe s , t oo thp icks , a nd c l ea r w a te r , a nd w e re k ep t a s soc ia ted w i th t h e ir c as ts .

p ar t icu la r g enus , w e re a s s igned t ot o t h e g ene ra l ized c a tegory

RECORD ING THE DATA

w a s l i m i ted b y a VERY h i gh d eg ree o f a nc ien t b reakage .

F ou r b road c a tegor ie s o f a t tr ibu te s w e re r e co rded f o r b one

b one , s ome b reakage a nd c r ush ing w as s u s ta ined d ur ing

i d en t if ica t ion .

S uch e l emen ts , i f p resumed t o r e p re sen t a

c fR ang if er , c fE quus , e t c . I nm any c a se s , b one i d en t if ica t ion A dd i t iona l ly , d ue t o e x t reme f r i ab i l i ty o f t h e G rubg raben e l emen ts . T he se c a tegor ie s i n c luded c r i te r ia f o r : 1 )t a xon a nd

s h ipp ing . When p os s ib le , b roken e l emen ts w e re i d en t if ied t o

e l emen t i d en t if i ca t ion , 2 ) c u l tu ra l mod if ica t ions , 3 ) n a tura l

g enus

mod if ica t ions , a nd 4 ) m ea suremen ts a nd w e igh t . A l l b one s a nd

A dd i t iona l ly , i d en t if ica t ion o f m idsha f t f r agmen ts o f l o ng

b one f r agmen ts a nd t h e ir a s soc ia ted a t tr ibu te s f o l lowed t h e

b one s o f t h e h o r se , a nd t o s ome d eg ree t h e r e indee r , w a s f a c i l i ta ted b y s aw ing l o ng b one s f r om c ompa ra t ive c o l lec t ions .

c od ing s y s tem d eve loped b y T odd ( 1987 ) a nd a r e p r e sen ted i n A ppend ix 2 . 1 . A l l a t tr ibu te s w e re e n te red i n to a P a radox 3 . 5 p rog ram ( Bo r land I n te rna t iona l 1 985 1990 )f i l ea sn umbe r c odes . T h is r e co rd ing p rocedu re f a c i l i ta ted d a ta m an ipu la t ion

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F i g . 2 . 2 Map o fG rubg raben , A L I . S tone f e a tu re s i n c lude ah ea r th c i rc le i nu n i t L f , d i ff use h ea r th i nu n i t L g , s t one p i le i nu n i ts L f /Mf , a nd al a rge s t one c l us te re x tend ing f r om u n i t L h t ou n i ts Mg , Ng , a nd N h ( f rom Mon te t - Wh i te a nd W i l l iams 1 994 :128 ; f i g . 3 ) .

9

F ig . 2 . 3 . P lan o ft h e s t one p avemen t I d le i nG rubg raben , AL2 . B lackened a r ea s i n d ica te b urned s t one s . S tr iped l i ne s d e l inea te t h e o che rc oncen t ra t ion w i th in t h e s t one s t ruc tu re . T he d i scon t inuous l i ne i nJ eo u t l ine s as econd s t one p avemen t ( f rom Mon te t - Wh i te 1 990 :54 ; f i g . V .6 ) .

1 0

a r chaeo log ica l b one c ou ld b e m o re e a s i ly d e te rm ined . S haf t e l emen t s s u ch a s m e tapod ia l s , p o s se s s ing h i gh ly d i s t inc t ive t n id shaf tf e a tu re s , w e re r e ad i ly i d en t if iab le . O nt h eo the rh and , b one s s uch a s h ume r i a nd f emo ra w e re l a rge ly i n d i s t ingu i shab le a s b one f r agmen t s i ze d ec rea sed . S uch p rob lema t ic b one e l emen t s w e re a s s igned t o t h e g ene r a l ized c a tego r ie s : l o ng b one , f l a t b one , o r c ancelou s b one . E nd s o f l ong b one s w e re i n spec ted f o r p r e sence /ab sence o f f u sed e p iphy se st o d e te rm ine g ene ra l m a tu r i ty o fa n ima ls r e cove red f r om t h es i te .

o f 5 6 ( 2 .9% ) . F ox w a s r a re i n l e ve l 1 ,r ep re sen ted o n ly b y p e rf o ra ted c an ine s u sed a s o rnamen ta t ion . I b ex , b ea r , a nd b ov id w e re r a re o ra b sen t . C u l tu ra l l e ve l 2 c on ta ined n ea r ly t w ice a s m any b one s ( N=59 13) a s l e ve l 1a nd r e p re sen ted 1 4 .2% o ft h e b one s a t t h es i te . R e indee rb one sa ga in d om ina ted , r e p re sen t ing 9 7% o f i d en t if iab le b one s . H o r se w a s n o t a s c ommon i nl e ve l 2w i th o n ly 8 9 b one s ( 2 .4% ) r e cove red . Numbe r o f h a re b one s d ec rea sed d rama t ica l ly f r om l e ve l 1 ,a nd i b ex w a s r e p re sen ted o n ly b y o ne b one . B ov id s , b ea r , a nd f ox w e re a b sen t i nl e ve l

B one s w e re q uan t if ied b y c a lcu la t ing N ISP ( numbe r o f i d en t if ied s pe c imen s ) d ef ined b y G rayson ( 1984 ) a nd MN I ( m in imum n umbe r o f i n d iv idua l s ) d ef ined b y Wh i te ( 1953 ) .

2 .

U s ing Wh i te ' sm e thod , t h eg rea te s tn umbe ro fa ny s i ng le b one e l emen t ( e .g . l e f t d i s ta l h ume ru s , p rox ima l r i gh t f emu r ) i d en t if i ed f o r a ny g i ven s pec ie s r e p re sen t s a tl e a s t t h a t s ame n umbe r o f i n d iv idua l s o f t h a t a n ima l . Wheneve r p o s s ib le ,

9 8% o f i d en t if iab le b one s ( N ISP= 16 ,009 ) b e long ing t o r e indee r . Mo re b one s w e re r e cove red i n l e ve l 3t h an i n a l o the rl e ve ls o ft h e s i t ec omb ined . A l though t he l a rge s tn umbe r o f h o r se b one s ( N ISP=262 ) w e re f ound i n l e ve l 3 , t h ey r e p re sen ted o n ly 1 . 6% o fi d en t if iab le b one s . B one so fa lo the r a n ima ls w e re n eg l ig ib le i nt h is l e ve l a l though b one s o f i b ex

c r i ter ia f o r a g ing a nd s exua l d imo rph i sm w e re t a ken i n to a ccoun t w hen c a lcu la t ing MN I v a lue s , f o l low ing t h e m e thod s o fC hap l in ( 197 1) , D a ly ( 1969 ) , K le in a nd C ruz -U r ibe ( 1984 ) , a nd U e rpman ( 1973 ) . F o r a pp rop r ia te s t a t is t ica l c ompa r ison s w i th k nown e t hnog raph ic d a ta , MN Iw a sc a lcu la ted a sd ef ined b y B inf o rd ( 1978 : 7 0 ) whe re a l MN I s w e re c a lcu la ted b y " . .d iv id ing t h eo b ser ved b one c oun t f o r ag i ven i d en t if i ca t ion u n i t b y t h e n umbe r o fb one s i nt h e a na tomy o f ac omp le te a n ima lf o rt h a tu n i t ." A dd i t iona l ly , MNE ( m in imum n umbe ro f a na tom ica l e l emen ts ) w a s c a lcu la ted f r om t h e mo s t c ommon p o r t ion o f e ach s ke le ta l e l emen t f o r e ach t a xa ; MNE r e p re sen t s t h e s um o fl e f t a nd r i gh t s i de s . " Ske le ta l e l emen t c oun t s (MNE ) p rov ide t h e b e s t a pp rox ima t ion o ft r an spo r ted e l emen t s . . . ." ( S t ine r 1 990 : 4 13 ) .

C u l tu ra l l e ve l 3w a s ad en se b one b ed ( N ISP=21 ,745 ) w i th

a nd h a re i n c rea sed f r om l e ve l 2 . F ox e l emen t s w e re c ompa ra t ive ly h i gh ( N ISP= 12 ) . B ov id r ema in sw e re f ound f o r t h ef i r s tt ime i nl e ve l3 . L eve l 4 ,t h e d eepe s t a nd o l de s t c u l tu ra l l e ve l , c on ta ined l e s s b one s t h an l e ve l 3( N ISP= 11 ,049 ) , b u t f a r mo re t h an e i the r l e ve l s 1o r 2 . R e indee r a ga in d om ina ted t h e a s semb lage ( N ISP=9243) , b u t h o r se b one s w e re n ume rou s ( N ISP=431) w hen c ompa red t o o the r l e ve l s . A l though i b ex b one s n umbe red h i ghe s t i nt h i s l e ve l , h a re a nd f ox b one s w e re r a re . B ov id sr e ached t h e i rh i ghe s tn umbe r si nt h i se a r l ie s tl e ve l . T he se g ene ra l d a ta s how t h a t G r ubg raben w a s ac amps i te w he re h un te r s s pec ia l ized o n r e indee r p rocu remen t . Ho r se s w e re a l so p roce s sed d u r ing e ach o ccupa t ion o ft h i s s i te . I b ex w e re r a re i na l l e ve l s . N umbe r s o fb one s o ff u rbea re r s , h a re a nd f ox , v a r ied a mong l e ve l s , a nd s ugge s t t h a t h un t ing / t rapp ing w a s n o t u nder taken o n al a rge s c a le a tt h i s s i te . B one s o fb ov id s i n d ica te t h a t t h e se a n ima l s w e re p re sen t i nt h e C en t ra l E u ropean B a s in f r om 1 8 19 ,000 B P , b u t m ay h ave m ig ra ted f r om t h e a r ea a s d e s s ica t ion i n t h e a r ea i n c rea sed . Mammo th a nd b ea r , r ep re sen ted b y i v o ry a nd t e e th , s ugge s t h un t ing o f t h e se a n ima l s d i d n o t o ccu r a tt h e s i te . S caveng ing o f i v o ry a nd t e e th f o r o rnamen ta t ion i s al i ke ly s c ena r io f o rt h ep re sence o ft h e se r aw m a te r ia l sa tt h es i te .

T he G rubg raben A s semb lage : At o ta l o f4 1 ,768 b one e l emen t s , t e e th , a n t le r , a nd i v o ry w e re a na ly sed i nt h i s s t udy ( Tab le 2 . 1) . I n c luded i nt h i s t o ta l a r e 2 , 355 b one s a nd t e e th f r om t h e 1 987 e xcava t ion s i d en t if ied b y L ogan ( 1990a a nd 1 990b ) . A pp rox ima te ly 2 4% ( 10 ,155 b one f r agmen t s )w e re e i the rs ob ad ly f r agmen ted o ro the rw ise l a ck ing i n d i agno s t ic c ha rac te r i s t ic s t h a t t h ey c ou ld n o t b e i d en t if ied b eyond g ene ra l ized m anuna l ia c a tego r ie s . Mammo th w a sr ep re sen ted o n ly b yt u sk ( i vo ry )f r agmen t s . N ob i rd s , f i sh , a mph ib ian so rr e p t i le sw e re r e cogn ized i nt h ea s semb lage . T he f ou r c u l tu ra l l e ve ls a t G rubg raben r e f lec t ar e la t ive ly n a r row s pec t rum o fa n ima le conomy a tt h es i t ew i th o n ly e i gh t g ene ra r ep re sen ted ( Tab le 2 . 1 , F igu re 2 . 4 ) . R e indee r r ema in s o ve rwhe lmed t h e f ou r c u l tu ra l l e ve l s . T he h o r se a l so f i gu red c on s is ten t ly a s am a jo rf ood /h ide s ou rce i na l l e ve l s a l though t ol e sse r ad eg ree t h an r e indee r . D epend ing o n l e ve l , b one so f i b ex , f ox , h a re , b ov id a nd b ea r w e r e e i the r a bsen t o r i nf a r f ewe r n umbe r s t h an e i the r r e indee r o r h o r se . A l though t h e se a n ima l s w e re p robab ly u t i l ized b y a nc ien t h un te r /ga the re r s a t G rubg raben , t h e s ma l l s amp le s i ze s ugge s t s t h ey w e re n eve r h eav i ly e xp lo i ted b yE p ig rave t t ian i n hab i tan t sa tt h is p a r t icu la r

CULTURAL MODIF ICAT IONS : C u l tu ra l mod if ica t ion s : c u t m a rk s , c hopp ing m a rk s ( Append ix 2 . 1 ; o bse rva t ion N ) a nd t y pe s o fb reak s ( e .g . s p i ra l , d r y , e t c . ; o b se rva t ion P ) , w e re r e co rded f o r r e indee r a nd h o r se . C ha r ac te r is t ic s o f b u rn ing : l o ca l ized c a rbon iza t ion , c a rbon i za t ion , a nd c a lc ined , w e re r e co rded w hen o bse rved ( s ee A ppend ix 2 . 1 ; o bse rva t ion 0 ) . Mod if ica t ion s a r e d i scu s sed b r ie f ly h e re a nd a r e e xp la ined i n m o re d e ta i l f o r h o r se , r e indee r , a nd l e s s u t i l ized f a una i nC hap te r s F i ve , S i x , a nd S even .

s i t e . V a r iou s l e ve l s o ft h e s i te p roduced d if e ren t s amp le s i ze s f o r a ny p a r t icu la r g enus , i n r aw q uan t i ty ( N ISP ) . R e indee r d om ina ted t h el e ve l 1a s semb lage , r e p re sen t ing 9 0% o ff a una l e l emen t s . H o r se r ema in s r e p re sen ted o n ly 6 . 6% o f b one s . H a re b one sw e re i ng rea te s t q uan t i t ie si nl e ve l lw i th a nN ISP

C u t t ing , C hopp ing a nd A b rad ing o nB one S u r f ace s :

1

I no rde rf o r a n a n ima l t op rov ide h uman s w i th f ood , c l o th ing , a nd t oo l s f o r t h e i r p a r t icu la r n eed s , t h e a n ima l m u s t b e e i the r s c avenged a s a c a rca s s o r c ap tu red , s l augh te red a nd

T ab le 2 . 1 . N ISP a nd p e rcen tage so ff a una r e cove red f r om t h ef o u rc u l tu ra ll e ve ls a tG rubg raben . I v o ry c oun t sa nd u n iden t if ied e l emen t sa r ee x c luded f r om p e rcen tage s ) .

T axon

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1 , 323

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GRUBGRABEN

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F ig . 3 . 1 . A nnua l a n t le rc yc le s i nr e indee r /ca r ibou ( der ived f r om B an f ie ld 1 962 ) .

3

T ab le 3 . 1 . D eve lopmen ta nd C a s t ing o fR e indee rA n t le r s( c omp i led f r om S t u rdy 1 975 ) . C yc le V e lve tc ommence s V e lve tc omp le ted C ompac t ing c omp le te A n t le r sp o l i shed C a s t ing c ommence s A n t le rc a s t

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A du l t Ma le s

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May

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J une m id O c tobe r e a r ly May

3 4

r ou te s a r e m a rked b y t r a i l s 6 12 i n ch e s w ide a nd v a ry ing i n d ep th b e twe en 4 i n che s t o n e a r ly a m e te r ( Banf i e ld 1 962 , N owak a nd P a rad iso 1 983 ) . D u r ing s p r ing m ig ra t ion s , b ul s t r a i l c ow s a n d y ea r l ing s a sp r egnan t f ema le s a r e i n s t inc t ive ly " d r iven " t ow a rd b i r th ing g round s . D u r ing f a l l m ig ra t ion s , b u l l s , y ea r l ing s a nd b a r r en c ow s a r r iv e a tw ooded w in te r ing g round sp r io rt oc ow ss l owed b yc a lve s . D u r ing s umme r , b u l l s f a t ten f o r t h e r i go r s o f t h e r u t a nd f ema le s , o f b r eed ing a ge , a t ta in c ond i t ion n ece s sa ry f o r s u cce s sf u lp r egnanc ie sd u r ing h a r sh w in te r s . S umme rf o rag ei s c r i t ica l t os u r v iva l a nd r e p roduc t ion . R e indee r a r e a dap ted t o m a rg ina l s u b s i s tence d u r ing w in te r b y r e ly ing o n af u l l n u t r i t iona l r e cove ry d u r ing s umme rf o rm a in tenance , b r e ed ing a nd , r e poduc t ion ( Dauph ine 1 976 ) . F a ti ss t o red v i sce ra l ly a nd s u bcu tan eou s ly w i th m o s t f a t s t o r ed o ve r t h e s a dd le a nd r ump . O n ah ea l thy a n ima l , f a t m ay b et h ree i n che s t h ick a nd w e igh 3 0 l b s ( Banf i e ld 1 962 ) . B o th b u l l s a nd c ow s a r e h eav ie s tp r io rt ot h ef a l lr u t . Af te rr u t , m a le s h ave l o s t a lmo s t a lo ft h e i rf a ta nd a r ei ng ene ra l ly p oo rc ond i t ion . I nc on t ra s t , c ow s ,f a t tened o ve rt h es umme rs l ow ly d e te r io ra te t h roughou t w in te r a nd a r e i nt h e i r p oo re s t b ody c ond i t ion f o l low ing t h e b i r th o fy oung i nl a te s p r ing /ea r ly s umme r .

E quu sc f .p r z ewa l sk i / f er u s :H o r se T he e co logy o f l a te P l e i s tocene h o r se s i s d e r ived f r om l i t e ra tu r e o n t h e P r zewa l sk i h o r se ( Moh r 1 97 1) , a nd m oder n f e ra l h o r se s i ng r a s s land /de se r t e nv i ronmen t s ( Be rge r 1 986 , T y le r 1 972 , D uncan 1 992 ) . D u r ing t h e P l e i s tocene , w i ld h o r se s r a nged f r om S ou thwe s t E u rope a c ro s s C en t ra l E u rope t o N or th a nd C en t ra l A s ia ( Moh r 1 97 1) . O ve rhun t ing a nd e n c roa chmen t i n to t h e i r s p a r se e nv i ronmen t h a s l e d t o t h e e x t inc t ion o ft h e A s ia t ic h o r se i nt h e w i ld . T he l a s t s u rv ivo r s i nt h i s H o locene r a nge l i ved a l ong t h e w e s te rn a nd n o r the rn b o rde r s o ft h e G ob iD e se r tf r om t h e D zunga r ian g a te s a c ro s s t h e n o r the rn p a r t o fS i nk i ang . As a l ine h i gh s t eppe b e tween 1 , 000 a nd 1 , 400 m e te r s a bove s e a l e ve l , t h e a r ea 's v ege ta t ion c on s i s t s o f s a l t t o le ran t p l an t s i n c lud ing s a xau l , t ama r i sk , w o rmwood , a nd t u s sock ga s se s . A s ia t ic w i ld h o r se s a dap ted t ot h i su nf avo r ab le , l ow p roduc t ive e nv i ronmen tb yg r az ing i n l a te e ven ing a nd e a r ly m o rn ing w hen p l an t s c on ta in m o r e w a te r . T h i s f e ed ing b ehav io ri so b se rved i no the r w i ld h o r se s ( Ty le r 1 972 , B e rge r 1 986 , D unc an 1 992 ) . R e s i l ian t p r zewa l s l c i s d i g h o le s w i th t h e i r h oov e s t o g a in a c ce s s t o s a l ine w a te r i n s umme r a nd t o f i nd f o rage u nde r s now i n w in te r . W i ld h o r se s c an s u rv ive w i thou t w a te r f o r u p t o f ou r d ay s( Moh r 1 97 1) .

B reed ing s e a son o ccu r s i n O c tobe r /Novembe r w hen b u l l s b a t t le t o e s tabl i sh h a rem s o f 1 0 15 c ow s o r i n d i sc r im inan t ly m a te w i th a n y a ccep t ing f ema le . F o l low ing t h e r u t , t h in a nd e xhau s ted b u l l s s hed t h e i r a n t le r s . A f te r g e s ta t ion o f e i gh t m on th s , s i ng le c a lve s , w e igh ing b e tw e en f i ve a nd n i ne k g , a r e b o rn w i th in a t wo w eek p e r iod d u r ing May June . P r io r t o p a r tru i t ion , c ow s s hed t h e i ra n t le r s , s p ec ia l ized m ine ra l s t o r e s , e a t ing t h em t oi n c rea se c a lc ium a nd m ine ra l sr e qu i red f o rt h e i r m i lk . P r e co c iou s c a lv e s c an s t and w i th in 3 0 m inu te s o fb i r th a nd c an r u n a f te r 9 0 . W i th in 2 4 h ou r s , c a lve s a r e s t rong e nough t o f o l low t h e h e rd . C a lve s b eg in t o e a t s o l id f o od w i th in t wo w e ek s , c on t inue t on u r se t h roughou t t h e s umme r , b u t c an b e w eaned w i th in am on th o fb i r th i ft h e i r m o the r s c anno t f e ed t h em . Y oung r e a ch s e xua l m a tu r i ty b e tween 2 9 a nd 4 1 m on th s ; a ve rage l o ngev i ty i nt h e w i ld i sb e tween f o u r a nd f i ve y e a r s .

P rzewa l sk i s a r e m ed ium s i zed a n ima l s . A ve rage h e igh t f o r m a le s i s 1 38 146 c m a nd 1 34 140 c m f o r f ema le s . T hey t y p ica l ly w e igh 2 50 -350 k g ( Moh r 1 97 1) . P i c tu red i n Magda len ian p a r ie ta l a r t a nd a r t m ob i l ie r ( Le ro i -Gou rhan 1 982 ) , t h ey a r e l a rg e h eaded , d un c o lo red a n ima l sw i th s ho r t , u pwa rd ly t ur n ed m ane s l a ck ing f o re lock s . D u r ing f a l l a nd w in te r , P rzewa l s l c i s g row a d en se w oo l ly u nde rcoa t o f h a i r w h ich t h ey s h ed , a l ong w i th t h e i r m ane a nd t a i l , b e tween Ma r ch a nd May ( Moh r 1 97 1) . U n l ike r e indee r , h e rd c ompo s i t ion o fw i ld h o r se si ss t ab le a nd n o rma l ly c on s i s t s o f s ma l l f am i ly u n i t s o ff i ve o r s i x m a re s , t h e i r c o l t s a nd a d e spo t ic s t al ion . B e cau se t h e d om inan t s t a l l ion i si n to le ran t o f a no the r a du l t m a le a s p a r t o f t h e m a re /co l t g roup , m a le o f sp r ing , 2y ea r s o r o l de r , a r e d r iv en f r om t h e i rp a r en ta l h e rd t ol i ve a l one o rf o rm b a che lo rg roup s o f t h re e t o f i f te en m a le s . A s t h ey g row a nd g a in s t reng th , y oung s t a l l ion sa t temp tt os t ea l y oung m a r e s f r om e s tab l i shed f am i ly g roup st of o rm t h e i ro wn h a r em s . U n l ike r e inde e r , w i ld h o r se s d o n o t h ave as e a sona l r u t t o e xhau s t b a t tl ing m a le s ; s t a l l ion sp ro te c t as ma l ln umbe ro ff ema le sy ea rr ound .

C h ief p r ed a to r s i n c lude w o lve s , b e a r s , w o lve r ine s , r a p to r s , a nd f o x e s . When t h rea tened , r e inde e r f l ee , r unn ing u p t o 8 0 k m /h ri n ac l o se g roup t oc onf u s e t h e i r p u r sue r s . T hey r a r e ly u se t h e i r i mp re s s ive a n t le r s a s am e an s o f d ef en se . F ema le s m ay d o s ot o d i s suade p r eda to r s a tack ing t h e i r n ew ly b o rn c a lve s a nd w ounded b u l l s m ay u s e t h e i r r a ck s a ga in s t p reda to r s . I n sec t p e s t s i n f luen ce b ody a nd c o a t c ond i t ion s . T o e s cape s wa rm s o f s umme r m o squ i to s , r e indee r m u s t c on t inua l ly move , r e su l t ing i n d e te r io r a t ion o f o p t ima l b ody c ond i t ion ( Remm e r t 1 9 80 ) . Wa rb le f l i e sl a ye gg so nt h ec oa t so fa n ima l s i nJ u ly /Aug u s t ; d u r ing w in te r , e gg s h a tch a nd l a rvae b o r e t h e s k in o fr e indee r , e n la rg ing b r e a th ing h o le s i nt h e s k in a st h e i n se c t s g r ow . F rom N ovembe r t o J une , w a rb le f l y l a r va e c r ea te s i g n if i can t d amage t ot h e h i d e s . D u r ing s umme r , l a rva e s c a r s h e a l . S ub sequen t ly , s umme r / ea r ly f a l l h i de s , l a ck ing w a rb le f l y b o r ing s , a r e m o s t s u i ta b le f o r c l o th ing ; h eav i ly f u r r ed w in te r c oa t s , a l though p e rm e a ted w i th l a rva h o le s , a r e n eve r the le s s , u sef u lf o rb edd ing a nd r o be s( Ban f ie ld 1 962 ) .

3 5

H o r se s l i ve w i th in u ndef ended c o r e t e r r i to r ie s w h ich o f t en o ve r lap . T h i s a r e a i sd ef ined a st h e u s e a r ea w he re t h ey s e ek s h e l te r , f ood , w a te r , a nd p o ten t ia l m a te s ( Bu r t 1 943 , T y le r 1 972 ) . H ome r a nge v a r ie s d epend ing u pon r e sou r ce a bundance ; i fs ou r ce s a r e p l en t if u l , h ome r a nge s t e nd t o b e s male r a nd c an b e u s ed b o th s umme r a nd w in te r w i th h o r se s s e a sona l ly u s ing d if e ren t s e c t ion s o ft h e r a nge . W i th s c a r ce r e sou r ce s , t h e i r r a nge s i n c rea se ( Ty le r 1 972 , B e rge r 1 986 ) . R ange s o f N ew F o re s t p on ie s c an v a ry f r om 2 00 t o 2 500 a c re s d epend ing o n a va i lab le r e sou r ce s ( Ty le r 1 972 ) . W i ld p r ezwa l sk i sc on t inua l ly w ande red t oe conom ize s pa r se f e ed ing g r ound s i nt h e G ob i ( Moh r 1 97 1) . I nt h e G rea t B a s in , e qu id h ome r a nge v a r ie s a l t i tud inaly a nd s e a sona l ly w i th s umme r r a nge s a bove 2 000 m a nd a l o t he r s e a son s b e low 2 000 m ( Be rge r 1 986 ) . B a che lo rg roup sa r em o s t u np red ic tab le , u s ing al a rge r c o re a r ea t h an h a r em g roup s ( An thony 1 99 1 , B e rge r

1 986 ) . H a rem g roup s , s l ow ed b y y oung s te r s , t r ave l l e s s d i s tance o na ny g i ven d ay ( Be rge r 1 986 ) .

r ugged t e r r a in . L iv ing i np r ec ip i tou sa r e a s , i b ex d i spe r se l i t t le , o ccupy ing l i m i ted t e r r i to r ie s t o w h ich t h ey a r e l o ya l . T hey r a re ly i n ha b i t a r ea s w i th s pa r se s h rub a nd f o r e s t c ove r p r ef e r r ing l a rge , h i gh a l p ine m eadow s w he re t h ey f e ed o n a m ixed d i e t o fg r a s se s , s e dge s , f o rb s , l e ave s o fd wa r fs h rub s , a nd l i chen s ( S cha l le r 1 977 ) . I bex d o n o t s e ek w a te r a st h ey e x t rac te nough l i qu id f r om p l an t st os a t i sf yt h e i rn eed s .

S t a l l ion sc hoo se t h ec o r ea r ea t ob ee xp lo i ted i na ny o ne y ea r , a nd h a rem g roup s a r e f a i thf u l t ot h e i r t e r r i to ry ( Ty le r 1 972 , B e rge r 1 986 ) . F o r e xamp le , m u s tang h a r em s i n t h e G rea t B a s in r e tu rned t o t h e c o re a r ea e xp lo i ted t h e p r ev iou s y ea r 8 1 -94% o ft h e t i me ( Be rge r 1 986 : 5 2 -53 ) . Y oung b a che lo r g roup s v a ry i n t h e i r c o r e a r ea s e lec t ion w i th o n ly 1 9% r e tu rn ing t o t e r r i to ry e xp lo i ted d u r ing t h e p r ev iou s y ea r ( Be rge r 1 986 : 5 3 ) . I n hab i t ing a s ma l le r c o re a r ea , t o w h ich t h ey r ema in f a i thf u l , h a rem g roup sm u s t h ave n ece s sa r i ly b een m o re p r ed ic tab le a nd e a s i ly e xp lo i ted , b y p r eh i s to r ic h un te r s , t h an b a ch e lo rg roup s .

I b ex s e a son a l ly m ove u p a nd d own m oun ta in s l ope s . I n w in te r , i b ex p r ef e r h i gh w ind s wep t v a l ley s f r e e o f s now w he re t h ey c a nm o s t e a s i ly f i nd f o rag e . I nc o ld w ea the r , t h ey f a vo r a r ea s e xpo sed t o s un l igh t a nd s e ek s he l te r f r om t h e w ind , p r ef e r ing t h ew a rme s t a r ea so ft h em oun ta in s a bove t h e t h e rmoc l ine . I n l a te w in te r , i b ex m ov e t o l owe r a l t i tude s w he re s now f i r s tm e l t sa nd t e nde rn ew g r a s sa ppea r s . P oo re s t b ody c ond i t ion o ccu r s i ne a r ly s p r ing w hen t h ey a r e m ov ing d own s lope t o l ow ly ing s p r ing p a s tu r e s . A s t h e s e a son p rog re s se s , t h e se c ap r id s m ove u pwa rd s a s t h e i r f o r age , f e d b y m e l t ing s n ow , b eg in s t o g reen . I b ex r e ach t h e i r h i ghe s t p a s tu re sd u r ing l a te s umme r . B ye a r ly f a l li b ex a r ei nt h e i rb e s t b ody c ond i t ion .

W i ld h o r se s m ove t o p ro tec ted v a l ley s i nw in te r f o r s h e l te r a nd f o rage ( Sp ie s s 1 979 , A n thony 1 99 1) . A s s e pa ra te f am i ly u n i t s m ig ra te , t h ey m ay a gg r ega te i n to l a rge r h e rd s . D u r ing t h e 1 9 th a nd e a r ly 2 0 th c en tu ry , m ig ra t ing m u s tang h e rd s c on ta ined b e twe en 5 000 t o 7 000 i n d iv idua l s , a nd l a rge e qu id h e rd s o f t h ou sand s o f a n ima l s o ccu r r ed o n t h e P ampa s o f S ou th Ame r ica ( Dob ie 1 952 ) . S o lu t r e m ay r e p re sen t as e r ie s o f k i l s o f h o r se s a s t h ey u nde rw en t s e a sona l m ig ra t ion s ( O l sen 1 989 ) .

I b ex f r equen t ly l i ve i nh e rd s o f3 -40 i n d iv idua ls ; r a r e ly d oe s a h e rd c on s is t o f2 00 i n d iv idua l s . I b ex s o c i a l s t ruc tu r e c on s i s t s o f t h ree f o rm s : n u r se ry b and s ; a l m a le h e rd s ; a nd m ixed h e rd s , d u r ing t h e r u t ( N iev e rge l t 1 966 , 1 9 81 ; S cha l le r 1 977 ) . T he se h e rd s a r e d if e ren t ia l ly d i spe r sed a l ong a n a l t i tud ina l g r ad ien tw i th m a le g roup sl i v ing i nh i ghe r , mo re r ugged a r ea s t h an f ema le /k id g roup s . Ma le s b row s e w i th in a l a rge r , l e s s p roduc t ive , c o re a r ea t h an f ema le s . F ema le s , e xpend ing e ne rgy o n p r egnancy , l a c ta t ion , a nd e ncumbe red w i th o f sp r ing i n expe r ienced i n c l imb ing , r e duce e n e rgy e xpend i tu re b y u s ing l owe r , l e s s r ugg ed a r ea s t h a t s u ppo r t h i ghe r n u t r i t iona l f ood s ( N ieve rge l t 1 966 , 1 98 1) . Y oung m a le s a r e v e ry a gg re s s ive . When a temp t ing b a t t le w i th i n d iv idua l s f r om a m o the r /k id g roup , f ema le s r e spond b y m ov ing a way . A s ar e su l t , y oung m a le s a s soc ia te w i th e a ch o t he r , c ha l leng ing a nd c l a sh ing . F ema le s a vo id c on f l ic t s t o p ro tec t t h e i r y oung a nd b e cau se t h ey a r e m uch s ma l le r t h an m a le s . F ema le sa ve rage 3 2k g( 24 -68 k g ) i nw e igh t ; m a le s a r e m uch l a rge r , a v e rag ing 8 4k g( 47 117 k g ) .

S t a l l ion s b eg in b r eed ing b e tween f o u r a nd f i ve y ea r s o fa ge , r ema in ing s e xua l ly a c t ive u n t i l 3 1 y ea r s ; m a r e sb eg in b r eed ing a t f ou r y ea r s a nd p roduce f o a l s u n t i l 2 4 y ea r s . G e s ta t ion a ve rage s 35 d ay s ( Hammond 1 960 , T y le r 1 972 ) . B i r th ing s e a son i si n May a nd J une ; d u r ing p a r t ru i t ion , m a re s d rop t h e i r f o a l s a pa r t f r om t h e h e rd a tn i gh t o r e a r ly d awn . F oa l b i r thwe igh t a ve rage sb e tween 2 5 a nd 3 0 k g . W i th in n i ne d ay s m a re s r e jo in t h e h e rd ( Moh r 1 97 1) . C o l t s r ema in w i th t h e i r m o the r s b e tween t wo a nd t h r ee y ea r s . N ea r ing s exua l m a tu r i ty , t h e m a le c o l t s a r e d r iven f r om t h e h e rd b y t h e s t a l l ion ; f ema le s a r e a lowed t os t ay u n t i l t h ey a r e " a bduc ted " b yy oung s t a l l ion s . T he d om inan t m a r e d e te rm ine s t i me a nd d i re c t ion s o f m ovemen t , l e ad ing h e r h e rd i ns i ng le f i l e ( Ty le r 1 972 , B e rge r 1 986 , A n thony 1 99 1) . Wh en t h r ea tened , w i ld h o r se s f l ee i n s i ng le f i l e w i th t h e f o a l s i n t h e m idd le a nd t h e s t a l l ion f o l low ing o nt h e s i de o fc l o se s t d ange r . I fn o y oung f o a l s a r e p r e sen t , s t a l l ion s l e ad t h e h e rd w i th m a re s c l o se ly b eh ind . When c o rne r ed , t h e s t a ll ion w i l l f a ce a nd a t ta ck , u s e b ru te f o r ce , s h a rp h oove s a nd t e e th , t o p ro tec t t h e h a rem . Mu s tang s , w hen a t ta cked b y w o lv e s , a r e k nown t o f o rm a c i rc le w i th h ead s o u t a nd f o a l s i nt h e c en te r . A s p r eda to r s c l o se i n , h o r se s g rab w o lve s w i th t h e i r t e e th a nd b ea t t h em w i th t h e i r h oove s . T h i s p ha lanx d e f en se p o s tu re c ove r s t h e u np ro tec ted f l ank s o f h o r se s a nd p r even t s h am s t r ing ing ( Dob ie 1 952 ) . S t a l l ion s s uf e r g r ea te s t l e tha l t h rea t s a nd y oung s t a l l ion s h ave t h e h i ghe s t d ea th r a te s d ue t o f o r ced e xpu l s ion f r om t h e i rm o the rh e rd . T he n o rma ll i f e span f o rw i ld h o r se s i s 2 0 -25 y ea r s w i th 3 4 b e ing t h e o l de s t k nown p r zewa l s l c i h o r se ( Moh r 1 97 1) . C ap ra i b ex : i b ex I b ex a r e a l p ine h e rb ivo re s , a dap ted t o s t e ep , r o cky , a nd h e te rogenou ss l ope sa bove t r eel ine b e twe en 2 000 a nd 2 800 m ( N ieve rge l t 1 966 , 1 98 1) . T he se r o bu s t , s u re -f oo ted g oa t s a r e e qu ipped w i th l a rge h oove s w e l l a dap ted t o n ego t ia t ing

T he r u t ,w he r em a le sc l a sh t og a in a cce s st of ema le s , o ccu r si n D ecembe r Janua r y . F o l low ing a r e la t ive ly l o ng 5 6 m on th g e s ta t ion p e r iod , k i d s a r e b o rn i n e a r ly s umme r ( S cha l le r 1 977 ) . F ema le s s e ek al o ne ly p l ace w h e r et h ey g i ve b i r th t oa s i ng le k i d ; t w in b i r th s b e ing r a re . D u r ing , a nd i mm ed ia te ly a f te r , t h e b i r th ing s e a son , f ema le s w i th y oung f o rm n u r s ing b and s . As e n t ina l , u sua l ly af ema le , p o s t sw a tch o ve rt h eh e rd i n ap rom inen tp o s i t ion . I b ex l i ve a n a v e rage o fs e ven y ea r s , a l thoug h s ome h ave l i ved a sl o ng a s1 8 . O n ly 5% o fi b ex r e ach t h ea g eo ff i ve . Ma le s , i n p a r t icu la r , s uf e r ad i sp ropo r t iona te ly h i gh m or ta l i ty i nw in te r i ft h ey h ave b e en v e ry a c t ive i nt h e p re ced ing r u t . Wo lv e sa r e t h e p r ima ry p r eda to r s o fi b ex ; s o l i ta ry m a le s , o l de r t h an f o u r t o f i ve y ea r s , a ppea r t o b e m o s t v u lne r ab le t o p r eda t ion ( Hep tne r e ta l1 9 66 ) . When t h rea tened , i b ex o f ten s e ek s a f e ty b y m ov ing u p s lope i n to r u gged t e r ra in , b u t t h ey d o o cca s iona l ly c r owd a nd m i l l t o ge the r t oc on f u se t h e i r e n emy . When a t tacked , t h e se g oa t s r e a r o n t h e i r h i nd l e g s a nd l u nge a tp r eda to r s . 3 6

T he e l u s iv e i b ex h a s b een h un ted i n av a r ie ty o f w ay s :

L epus t im id us : A r c t ic h a re

1 ) ambu sh e s a t s a l t l i ck s , 2 ) ca ref u l ly p l anned d r ive s w he re h un te r s h e rd f r igh tened a n ima l s u p s lope t owa rd a wa i t ing h un te r s , a nd 3 ) s ta l l c ing f r om u ps lope ( S t rau s 1 987 ) .

A rc t ic h a re s a r e e x t reme ly h ea r ty a n ima l s t h a t t h r ive o n g ra s se s , h e rb s , a nd n u t r i ton a l ly d ef ic ien t v ege ta t ion i n c lud ing dwa rf b i r ch a nd w i l low t w ig s ( Ha rpe r 1 956 ) . T he se l a gomo rph s a r e e qu ipped w i th v e ry s t ra igh t i n c i so r s w h ich

A lopex l a gopus : A rc t ic f ox

t hey u se t o e x t rac t t i ny p l an t s f r om c r ev ice s ( Nowak a nd P a rad iso 1 983 ) . H a re s , a s o ppo r tun i s t s , h ave b een k now t o f e ed o n t h e s t oma ch c on ten t s o fe v i sce ra ted c a r ibou ( Ha rpe r 1 956 ) .

E x tan t a r c t ic f oxes l i ve i n a r c t ic a nd a l p ine t und ra , o pen wood land , a nd t he i c e c ove red w a te r s o f n o r ther n E u ra s ia , No r th Ame r ica , G reen land , a nd I c e land ( Co rbe t 1 980 , H a l l 1 981 ,C h e semo re 1 975 ,B anf ie ld 1 974 ,P re s t rud 1 992 ) .

A rc t ic h a re s p ref e r r ocky p l ace s a nd h i gh r ugged e l eva t ion s a vo id ing m a r shy a r ea s a nd w ide , f l a t p l a in s ( Ha rpe r 1 956 ,

T he se s ma l l f oxe s a r e d i ur na l a nd o ppor tun i s t ic h un te r s / s cavenge r s t ha t f e ed o n l emm ing s , v o le s , l e ve re t s , b i rd s a nd e gg s , s ea l p ups , r e inde e r c a lve s , a nd r e f u se t h ey s cavenge f r om k i l ls o f o t he r p reda to r s ( Che semo re 1 969 , Macphe r son 1 969 , C o rbe t 1 980 , Nowak a nd P a rad i so 1 983 , B rae s tn ip 1 941 , a nd P re s t rud 1 992 ) . I np repa ra t ion f o rw in te r

P ede r son 1 966 ) . T h i s i s p robab ly a d ef ense t a c t ic ; when t h rea tened b yp reda to r s ,b o th y oung a nd a du l ts f r eeze t ob l end i nw i th s u r round ing s t one s . N oc tu rna l ly a c t ive , h a re s s he l te ri n r ock c r ev ice s a nd u nde r b ou lde r s d u r ing t h e d ay ( Pede r son 1 966 ) . Du r ing w in te r , h a re s p ref e r t o l i ve i n w indswep t , moun ta inou st e r ra in ( Pede r son 1 966 ) , w he re , t r oop s o f1 5 -20 a n ima l sg a the r . I ns p r ing ,t h ey move t ol owe re l eva t ion s .

h a rd sh ip s ,f o xe sc ache d ead b i rd s , b i rd e gg sa nd s ma l l r oden t s u nde r b ou lde r s , i n c r ev ice s i ns haded a r ea s , o r i nh o le s d ug d own t o p e rmaf ros t ( Pede r son 1 966 , Nowak a nd P a rad i so 1 983 , P r e s t rud 1 992 ) . A rc t ic f ox p opu la t ion s f l uc tua te w i th

A tt h ee nd o fA p r i l , t r oop sb r eak u p a si n d iv idua l s s e ek m a te s ( Pede r son 1 966 ) . C oup le s r ema in i n dependen t o fo the r s u n t i l a u tumn w hen t r oop s a ga in f o rm . A f ema le h a re b ea r s a n a ve rage o f s e ven l e ve re t s p e r l i t te r n ea r t he e nd o f J une

t he t h re e t o f i ve y ea r l emm ing c r a sh c yc le s ( Bee a nd H a l l 1 956 ,P ede r son 1 966 ,C he semo re 1 975 ) . A s y oung a st e n mon th s , f oxe s m a te b e tween F eb rua ry a nd May w i th m os t ma t ing o ccu r r ing d u r ing Ma rch ( Pede r son 1 966 , N owak a nd P a rad i so 1 983 ) . T hey a r e v e ry d if i cu l t t o t r ap d u r ing m a t ing s e a son a s t h ey a r e p reoccup ied w i th f i nd ing m a te sa nd p ay l i t t le a t ten t ion t of ood ( Pede r son 1 966 ) .

( Pede r son 1 966 ) . N ew ly b or n h a re s , u n l ike r a bb i t s , a r e b o rn w i th g ray f u r a nd w i th e ye s o pen . T he f ema le s t ay s w i th h e r b rood f o r t wo t o t h ree d ay s d ef end ing t h em f r om p reda to r s ;

F o l low ing a n a ve rage g e s ta t ion p e r iod o f 5 2 d ay s , am a ted p a i r d i g s ah o le i ns of t s o i l o r s and o ro ccup ie s ad en f r om a p rev iou sy e a r , o f ten b e ing l o ca ted i nl ow m ound s o ne t of ou r m e te r s h i gh ( Corbe t 1 980 , N owak a nd P a rad iso 1 983) . B ecau s e a r c t ic f oxe s a r e m o re s oc ia l t h an o the r f ox s pec ie s , b reed ing b u r row s m ay b ec l ose t oge the r . S ome f ox d en s h ave b een u s ed f o ro ve r 3 00 y ea r s a nd m ay c on ta in f ou rt ot we lve e n t ranc e s ( Macphe r son 1 969 , E be rha rd t e ta l . 1 983 ) . T he se a nc ien t d en s a r e e a s i ly s po t ted o n t h e l a nd scape b ecause o rgan ic d eb r i s a ccumu la te s i nt h e i mmed ia te v i c in i ty s pu r r ing v ege ta t ive g row th ( Sm i t s e t a l . 1 988 ) . Ma t ing a nd b i r th ing s ea son i st h e o n ly t ime t h a t f oxe s p o s se s s af i xed h ome s i te . A s ar u le , k i t s a r e b or n a tt h e e nd o f May o r f i r s t o fJ une ( Pede r son 1 966 ) . Av i xen m ay b ea ru p t o2 5 k i t sa l though s i x t o t w e lve a r e mo re c ommon ( Pede r son 1 966 ) . K i t s a r e w eaned w hen t wo t of ou rw eek s o l da nd y oung a r e c a red f o r b y b o th p a ren ts u n t i l t h e f am i ly d i spe r se s f r om m id A ugus t t h rough a u tumn ( Nowak a nd P a rad i so 1 983 ) . W i th t h eo n se t o fc o ld w in te r m on th s , f oxe s m ig ra te , o f ten i n s oc ia l g r oup s a nd o ve r l o ng d i s tan ce s , s ou thwa rd o r o n to p ack i c e d epend ing o n t h e a va i lab i l i ty o f f ood . L onge s t m ig ra t ion s a r e s pu r red b y d rop s i nl emm ing p opu la t ion s i na g iven a r ea ( Nowak a nd P a rad iso 1 983 ) . A rc t ic f oxe s h ave b een d ocumen ted t ot r ave l a sf a ra s2 000 k i lome te r s i nw in te r m ig ra t ion ss e ek ing f ood ( Nowak a nd P a rad i so 1 983) . O nce a t t h e i rd e s t ina t ion s , f oxe s d i spe r se a nd b ecome s o l i ta ry h un te r s . A l though w i ly a nd t e nac iou s , f oxe so n ly l i ve f o r s e ve ra l y ea r s d ue t ot h e i rp reca r iou se nv i ronmen t . Mo s te x tan tf oxe s d o n o t s urv iv et h e i rf i r s t y ea r , s t a rv ing t h ef i r s t f ew m on th s o fw in te r ( Che s emo re 1 975 ) .

t h e m a le o f ten a bandon s t h e f am i ly g roup a tt h i s t ime . Af te r t h ree d ay s , p r ecoc iou s , y oung h a re s a vo id p reda t ion b y c amo f laug ing t h em se lve s a mong s ma l l r o ck s( Pede r son 1 966 ) . L eve re t s a r e n u r sed b y t h e i r mo the r s f o r t wo t ot h ree w eek s ; a tt h i st ime t h ey a r e s uf i c ien t ly m a tu re t o l e ave t h e i r mo the r . L i t te rm a te s r ema in t oge the rt h roughou t t h e f o l low ing w in te r ; b yw in te r , a la r ef u l ly a du l t . I n a u tumn , c on t ro l led b y d ec rea s ing d ay l igh t h ou r s , t h e c oa t l e ng then s a nd t u rn s p u r ew h i te . U n l ike t h a t o ft h e f ox , t h e f u r o fh a re s i sn o t ap r ized c ommod i ty . A l though t h e f u r i sl o ng a nd v e ry s o f t , t h e s k in i sw eak a nd i so n ly u sef u l f o rt r imm ing o r l i n ing g a rmen t s ( Pede r son 1 966 , Wa lke r 1 968 ) . H a re s a r e f a t te s t i n l a te s umme r /ea r ly f a l l , a nd a r e p rey f o r f oxe s , wo lve s , e rm ine s , a nd r a p to r s( Remme r t 1 980 ) . A dap ta t ion , M ig ra t ion , a nd t h eL a s tG lac ia l Max imum : A sd e ta i led i nC hap te rO ne , moder na r chaeo log ica lt e chn ique s c oup led w i th r ef ined e nv i ronmen ta l s t ud ie s r evea l t h a t r a the r t h an b e ing a v a s t p o la r d e se r t d u r ing t h e LGM , C en t ra l E u rope c on ta ined " . . loca l ized a nd s h if t ing , r e la t ive ly h um id n i che s w i th in ag ene ra l ly d r ie r r eg ion " ( Mon te t - Wh i te 1 994 : 4 9 1) . C o ld , a nd i n c rea s ing a r id i ty , a s soc ia ted w i th t h e L a s t G lac ia l Max imum i nC en t ra l E u rope , m u s t h ave n ece s sa r i ly i mpac ted p l an t c ommun i t ie s a nd t h e h e rb ivo re s w h ich f o raged u pon t h em . T h i s c l ima t ic d e te r io ra t ion r e su l ted i np a tche s o f r e sou r ce s o f r e duced p r ima ry p roduc t iv i ty , w i th a r e su l tan t d ec rea se i nt h e r i ch a nd v a r ied f a una wh ich h ad p r ev iou s ly b een a va i lab le t o G rave t t ia n h un te r s ( Logan 1 990a a nd

1 990b ) . T he c a ta logue o f g ame a n ima l s a t G rave t t ian s i te s s uch a sW i l lendo rf ( Then iu s 1 956 1959 ) a nd D o ln i V e s ton ice ( K l ima 1 963 ) w hen c ompa red t o t ho se f r om E p ig rave t t ian s i te s s uch a s C r rubg raben ( Logan 1 990a , 1 990b ) , S agva r ( Vo ros 1 982 ) , a nd P i l i sma ro t ( Dobo s i e ta l . 1 983) , r ef lec t s a 3 7

r e duced d i ve r s i ty o f a n ima l s e xp lo i ted t h rough t i me i n t h e C en t ra l E u ropean P l a in . I ti su n l ike ly t h a t h un te r s i n ag i ven a r ea w i l l ing ly c ho se t o i g no re g ame s p ec ie s w h ich t h ey h ad f o rme r ly u t i l i zed . T he l i m i ted f a una l l i s t o f g ame a n ima l s r e cov e red f r om E p ig rave t t ian a s semb lage s r e f lec t a d epopu la t ion a nd /o r m ig ra t ion o f c e r ta in s p ec ie s f r om a d e te r io ra t ing , p a tchy , a nd l e s sp rodu c t ive e nv i ronmen t . C en t r a l E u ropean s pe c ie s i n c lud ing b i son , a u roch s , a nd m ammo th s w e r e a ppa r en t ly i n to le ran t o fv ege ta t ive p a t te rn s w h ich r e su l ted f r om t h e c o ld a nd i n c r ea s ing a r id i ty a s soc ia ted w i th t h eL a s t G lac ia l Max imum . C onve r se ly , h o r se s , r e indee r , a nd i b ex w e r e s uf i c ien t ly f l ex ib le t o p r eva i l a nd b e come p r ima ry f o od /f a t /h ide r e sou r ce s f o r P a leo l i th ic h un te r s . T h i s e co log ica l r e shu f f l ing m ay b e r e la ted t o a c omp lex c omb ina t ion o f p hy s io log ic a l a nd b ehav io ra l p roce s se s i n c lud ing f e ed ing b ehav io r , r e p roduc t ive p hy s io logy , m a t ing , a nd d e f en s em ech an i sm sa dap t ive f o rs ome s p ec ie sa nd n o tf o r o t he r s . G roup s i ze , d a i ly a nd s e a sona lb ehav io r , a nd d i ge s t ive c apab i l i t ie s o fh o r se , r e indee r , a nd i b ex p r e sumab ly e qu ipped t h em w i th a c ompe t i t iv e e dge o ve r l a rge r b ov id s a nd m ammo th s . G ene ra l ly , u ng u la te s , h oof ed m amma l s , a r et h ep r ey o fc ho ice b y m o s t l a nd locked t empe ra te h un te r /ga the r e r s . U ngu la te s , e qu ipped w i th s p e c ia l ized d i ge s t ive c hambe r s , c onve r t l a rge ly u nd ige s t ib le l e ave sa nd s t em so fp l an t si n to t h ep ro te in a nd f a t n ece s sa ry f o r h uman c on sump t ion ( Van S oe s t 1 980 , F oose 1 982 , D un c an 1 992 ) . P l an t s c on s i s t o ft wo m a jo r f un c t iona l c omponen t s : c e l l c on ten t s a nd c e l l w a l l s . C e l l c on ten t s : p ro te in , s t a r ch , s uga r s , l i p id s , a nd o rg an ic a c id s a r e l a rge ly d i ges t ib le b y a l m amma l s . C e l l w a l l s o fp l an t s a r e c ompo sed o f f i b rou s , c omp lex c a rbohyd ra te s : c e l lu lo se , h em icelu lo se , a nd l i gn in ( Van S oe s t 1 980 , J an i s 1 976 ) . C e l l w a l l s p rov ide p l an t s w i th s t ruc tu r e a nd d ef end t h e o rgan i sm a ga in s t p r eda t ion . Mamma ls c anno tb r eak d own t ough c omponen t so f c el w a l l sf o rn u t r i t ion . T he u ngu la te g a s t ro in te s t ina l t r a c t h a s e vo lved t o i n c lude s p ec ia l ized f e rmen ta t ion c hambe r s w i th e nzyme -p roduc ing m ic roo rgan i sm s w h ich ef i c ien t ly b r eak d own f i b rou sc omponen t s o fp l an t c e l l w a l l s . U ngu la te s a l one c an u t i l ize t h e se m ic roo rgan i sm s a nd t h e i r b y -p roduc t s , v o la t i le f a t ty a c id s , f o r n u t r i t ion a nd g row th . T hu s , u ngu la te s h ave e n te red i n to a s ymb io t ic r e la t ion sh ip w i th c elu la se p roduc ing b a c te r ia . T he u ngu la te h o s t p rov ide s t h e se m ic roo rgan i sm sw i th af e rmen ta t ion c hambe ri nw h ich t ol i ve ; i n r e tu rn , t h e b ac te r ia b r eak d own u nd ige s t ib le c e l lu lo s ic s u b s tance s f o rt h e h o s t ' s u s e ( Van S oe s t 1 980 , D uncan 1 992 , J a n i s 1 974 , 1 976 ) .

H e rb ivo re s g a in t h e i r n u t r i t iona l r e qu i r em en t s t h rough a v a r ie ty o fs t r a teg ie s . G r aze r s s u ch a sc a t t le ,b i son , a nd h o r se s , e qu ipped w i th l a rge m ou th s a nd h yp sodon tt e e th , c l ip g r a s se s a nd f o rb s . T h e i r l a rge s i ze d i c ta te s t h a t t h ey m u s t t a ke i na s u b s tan t ia l a moun t o f g r a s se s t o m a in ta in t h e i r n u t r i t iona l n eed s . L a rge r um inan t g r a ze r s s uch a sb i son a nd c a t t le a r e e x t reme ly e f i c ien t a te x t rac t ing n u t r ien t s f r om p l an t s i nt h e m ode ra te ly s i zed f i be rc a tego ry ( Van S oe s t 1 982 , J a n i s 1 9 76 , D emmen t a nd V an S oe s t 1 985 ) . Howev e r , a sf i be r i n c r ea se s , m a s t ica ted v ege ta t ion m u s t r ema in i nt h e r um en f e rmen ta t ion c hambe r l o ng e r s ot h a t i tm ay b e d i ge s t ed t o as ma l l e nough s i ze t o p a s s t h rough t h e r e t icu lo rumen o r if i ce i n to t h e t h i rd s t oma ch f o ra b so rp t ion . A s f i be r c on ten t i nt h e d i e t i n c r ea se s , t h er umen s oon f i ls a nd d i ge s t ion r a te sa r en e ce s sa r i ly s l owed . A c oa r se f i be r d i e t , t h e re f o re , p re sen t s ad i lemma f o r t h e g r a z ing r um in an t a dap ted t o am ed ium f i be rd i e t ( F igu r e 3 . 3 ) . B e cau se o fi t s l a rge s i ze , t h e a u roch /b i son , s p ec ia l iz ing i na m ode ra te f i be r /h igh p ro te in d i e t , m u s t i n ge s t av e ry l a rge q uan t i ty o fh i gh f i be rv ege ta t ion t om a in ta in i t s e lf ; h owev e r , i t m u s tw a i tf o re x ten s ive r um ina t ion a nd s u b s equen t p a s sag et o o ccu rb ef o r e i tc an i n ge s t m o r e f ood . F o r ag r az ing r um inan t , t h e r ei s ad ef in i te c u t o fi nt h e p e rcen tage o ff i be rt h a t i tc a n t o le r a te . B eyond t h a t p o in t i tc anno t m a in ta in i t se lf ( J an i s 1 976 , F oo se 1 9 82 ) . F oo se ( 1 982 : 1 3 3 ) , s t udy ing d i ge s t iv e r a te s o fw i ld a nd d ome s t ic u ngu la te s , c on c luded , " . .on v e ry f i b rou s f o rage s r um inan t s c an l i t e ra l ly s t a r ve t o d ea th w i th t h e i rg u t sf i l l . "

Two b a s ic u ngu la te d i ge s t ive s y s tem s e x i s t : a r t ioda c ty l ( f o r egu t )a nd p e r i s sodac ty l ( h indgu t ) f e rmen te r s ( Foo se 1 982 , J an i s 1 976 , D uncan 1 992 , V an S oe s t 1 980 ; F igu re 3 . 2 ) . A r t ioda c-t y l s u s e t h e l a rge c hambe r o f t h e f o regu t , r umen / r e t icu lum , a nd i t s a s soc ia ted b ac te r ia t o e f i c ien t ly b r eak d ow n a nd f e rmen t p l an t c el w a l l s p r io r t o t h e i r a b so rp t ion i n t h e l owe r d i ge s t ive t r a c t . T he r umen a l so d e tox if ie s " s e conda ry p l an t s u b s tance s " a low ing t h ea n ima lt o c hoo se w ha t w ou ld o t he rw i se b e p o ten t ia l ly p o i sonou s f o rage s i n i t s d i e t ( Foo se a nd L l oyd 1 974 , F r ee land a nd J an zen 1 974 ) . A r t ioda c ty l s i n c lude : b i son , c a t t le , g oa t , a nd d ee rs p ec ie s . O nt h eo t he rh and , m onoga s t r ic d i ge s te r su se t h e e n la rged c e cum , ab l ind s a c a tt h ej unc t ion o ft h e s ma l l a nd l a rge i n te s t ine s , t o b r eak d own a nd f e rmen t f i b rou s p l an t

t i s sue s . Monoga s t r ic d i ge s te r s i n c lud e t h e h o r se a nd t h e m ammo th . I nt r u th , b o th t h e r umen a n d c e cum p e rf o rm t h e s ame d u t ie s , h o s t ing c elu la s ic b ac te r ia w h ich b r eak d own n ond ige s t ib l et i s sue s ; i ti st h e i rd if e r en tl o ca t ion sw h ich a low f o rd if e ren tt y pe so fe f f i c ien ce si np roce s s ing p l an ts u b s tan ce s ( J an i s 1 976 ) . B o th u ngu la te f o rm s a r o s ed u r ing t h e P a leocene ( Scha ef f e r 1 948 , R ome r 1 966 ) . T hey e v o lved i np a r a l le l u n t i l t h e O l igocene w hen a r t iodac ty ls , i n p a r t icu la r t h e b ov id s , u nde rw en t a t r emendou s r ad ia t ion f a r o u td i s t an c ing p e r i s sodac ty l s i nb o th s p ec ie s a nd n umb e r s ( J an i s 1 976 ) . T he e xpan s ion o fr um inan t s a tt h e e xpen se o fm onoga s t r ic f o rm s w a s o nce a tr ibu ted t o t h e mo re h i gh ly e vo lved , e f i c ien t d i ge s t iv e s y s tem a s soc ia ted w i th a f o u r c h ambe red s t oma ch ( Mo i r 1 967 ) .F i e ld s t ud ie si nA f r ica ( Be l l 1 970 , 1 97 1) s h owed t h a t r um inan t s a nd m onoga s t r ic s , r a th e r t h an c ompe t ing f o r t h e s ame f o od s ou r ce s , c omp lem en t e a ch o t he r b y : 1 ) oc cupy ing s e pa ra te s u bhab i ta t s , 2 ) s e le c t ing d if e ren t p l an t s a nd / d if e r en t p a r t s o f p l an t s w i th in ag i ven h ab i ta t , o r 3 ) s e lec t ing s e pa ra te g row th s t age s o f ap l an t w i th in a g i v en p l an t t a xon ( Lamp r ey 1 963 , V e sey -F i tzg e ra ld 1 960 , Ma ck ie 1 970 ) . A s ar e su l t , t h e c ompe t i t ion m od e lf o ra r t iodac ty l sa nd p e r i s soda c ty l sh a s l a rge ly b een d i sm i s sed ( J an i s 1 976 , D un can 1 992 ) .

3 8

T he h o r se , b e a r ing a h i ndgu t f e rmen ta t ion s y s tem , i sn o t l i m i ted b y r a te o f p a r t ic le s i ze r e duc t ion t o p a s s h i gh f i be r p l an t st h rough i t sd i ge s t ive t r ac t . A sf i be ri n c r ea se sa nd u s ab le p ro te in d e c rea s e s , t h e e qu id i n c rea se s i t si n take o fv ege ta t ion w h ich i n t u rn , p a s se s t h rough t h e d i ge s t ive s y s tem i n a r e la t ive ly s ho r t t i me ( Foo se 1 982 , G u th r ie 1 990 ) . F o r e xamp le , p l an t p a r t ic le s i n al a rge b ov id p a s s t h rough t h e s y s tem b e tween 7 0 a nd 9 0 h ou r s ( Ba lch a n d C amp l ing 1 965 ) ; t h es ame t y pe o fd i ge s ted m a t te rp a s se st h rough ah o r se i n4 8 h ou r s ( A lexand e r 1 946 , H e in le in e ta l . 1 966 , V ande r N oo t e t a l . 1 967 ) . Ad i e ta ry s t ra tegy o fq u ick p a s sag ea low s e qu id st o

3 9

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e x i s to n ah i gh f i be rd i e tu pon wh ich r um in an ts o fs im i la rb ody s i ze s imp ly c anno t ma in ta in t h em se lve s . S uch ad i e t s t ra tegy m ay h ave p rov ided e qu id s w i th a s e le c t ive a dvan tage i nt he

B ecau se o ft h e i r s ma l le r s i ze , h e rb ivo re s s uch a sr e indee r a nd i b ex d o n o t p o s se s s g u t s l a rge e nough t ou se l a rge q uan t i t ie s o fh i gh f i be r g r a s se s . T he i r i n take c apa c i ty , d i c ta ted b y t h e i r s ma l l s i ze , i s n ece s sa r i ly l owe r t h an t h a t o f t h e i r l a rge r r um inan t c ou s in s . T o c ompensa te , t h e se s male r b od ied r um inan t s , e qu ipped w i th s ma l le r m ou th s a nd l ow

d e s s ica t ing c l ima te a nd a s soc ia ted l ow p roduc t iv i ty o fC en t ra l E u rope d u r ing t he L a s t G lac ia l Max imum . E x tan t e qu id p opu la t ion s c an f l ou r i sh i n n or the rn l a t i tude s , w he re s now c ove r l a s t s s i x mon th s a nd t empe ra tu re s f a l l a s l ow a s 40 d eg ree s C . , t o d e ser t r eg ion s w he r e f ood i s s pa r se ly a nd u neven ly d i s t r ibu ted a nd t empe ra tu re s e xceed 4 2 d eg ree s C . ( Be rge r 1 986 ) . C a t t le a nd b i son a r e u nab le t o s u rv ive i n e nv i ronmen t so fs uch l ow p l an tp roduc t iv i ty .

b rachyodon tt e e th , a vo id t ough f i be rb y s e lec t ive ly f e ed ing o n c e l l c on ten t s a nd e a s i ly d i ge s t ib le c e l l w a l l c omponen t s ( J an i s 1 976 , V an S oe s t 1 982 ) . B row se r s c ompen sa ted f o r s ma l le r r umen s i ze w i th a l owe r a b i l i ty t o d i ge s t c e l lu lose , b y c on sum ing d en se r , m o re h i gh ly s e le c ted , l e s s f i b rou s p l an t s . C onve r se ly , l a rge gaz ing r um inan t s s uch a sc a t t le a nd b i son c anno t a dap t t o ab row s ing s t ra tegy d ue t o t h e i r s i ze a nd mou th d e s ign . T hey a r e s imp ly u nab le t o s e lec t ive ly c hoose

E lephan t s , u s ing p rehen s i le t r unk s , h e igh t , a nd s hea r m a s s , f e ed o n aw ide v a r ie ty o f f o rage s f r om r oo t s t o h i gh t r ee f o l iage . " The i r d i e ts a r e t h e mo s t i n c lu s ive p o s s ib le i n a ny h ab i ta t " ( Hayne s 1 99 1 : 5 9 ) . E lephan t s a r e a b le t o d i ge s t l ow f i be r f r e sh g ra s se s a s w e l l a s h i gh f i be r wood . A h i ndgu t f e rmen te rl i ke e qu id s , t h e e l ephan t c an e a t g rea t q uan t i t ie s o f v ege ta ton , d i ge s ti tq u ick ly i ni t sl a rge c e cwn , a nd r a p id ly p a s s

s egmen t so ff o rage w h ich p rov ide t h eg r ea te s t b enef i t ( Owen Sm i th 1 980 ) . D u r ing t h e L a s t G lac ia l Max imum , a st h e c l ima te c oo led a nd b ecame m o re a r id , p l an t g row th w ou ld h ave b ecome mo re f i b rou s d u r ing s ome p a r t s o f t h e y ea r . " I n d e se r t r eg ion s , a dap ted p e renn ia lp l an t sm ay r eve r tt od ormancy , t r an spor t ing n u t r ien t s t ot h e r oo t s a nd l e av ing a n a e r ia l p a r t o fd ec rea sed n u t r i t ive v a lue . A nnua l p l an t st e nd t or ema in o fh i gh q ua l i ty a t m a tu r i ty b ecau se s t o rage c a rbohyd ra te i sl e f ti nt h e s e eds o ve r t h e d ry p e r iod s . . .The q ua l i ty o ft h is f o rage i su sua l ly n o t s o l i m i t ing a si si t s s upp ly " ( Van S oe s t 1 982 : 7 1 -72 ) . T he r a te o f f i be rc on ten t m u s t h ave i n c rea sed t ot h e c r i t ica l l e ve l w he re i t wou ld h ave m ade f o rage s u nava i lab le t o s ome s pec ie s o f g raz ing u ngu la te s . L a rge , g ra z ing b ov id s s u ch a sa u roch s a nd b i son w i th t h e i r s l ow r um ina t ion s y s tem s w ou ld h ave b een u nab le t o d i ge s t h i gh f i be r v ege ta t ion q u ick ly e nough t o m a in ta in t h e i r d i e ta ry n eed s d u r ing t h e d r ie s t s e a son s o ft h e y ea r . T h i s " bo t t lene ck " w ou ld h ave f o rced t h e se g raze r s t o move i n to a r ea s o fE u rope w he re t h ey c ou ld f i nd mo re e a s i ly d i ge s t ib le , l e s s d i spe r sed p l an t s . T he h o r se , a dap ted t o ah i gh f i be r / low p ro te in d i e t a nd p o s se s s ing ar e la t ive ly r a p id f o rage p a s sage r a te s u rv ived i nC en t ra l E u rope d u r ing t h e h a r she s t o f t ime s .

u nd iges t ib le f r ac t ion a s i tc on t inue s f e ed ing ( Hayne s 1 991) . D ige s t ion r a te s o fe l ephan ts v a ry b e tween 2 4 a nd 5 4 h ou r s d epend ing o n f i be r c on ten t o ft h e f o r age ( Bax a nd S he ld r ick 1 963 ,B ened ic t 1 936 ) . P re se rv ed d ung , s t oma ch c on ten t s , a nd v i sce ra i n d ica te t ha t a nc ien t e l eph an ts p o s se s sed a d i ge s t ive s t ra tegy m uch l i ke t he i r m ode rn a na logue s . F ood f r agm en t s a nd p o l len f ound i n w e l l p r e se rved S i be r ian m ammo th s i n d ica te a d i e t o f l a rch , b i rch , w i l low , s edge , m o s se s , a nd g r a s se s . T he p ref e r red d i e t w a s g ra s se s , a l though w oody p l an t s a nd s ma l l f o rbs w e re a l so e a ten ( Ve ra shchag in 1 975 , V e ra sh chag in a nd B a ry shn ikov 1 982 , Y ud ichev a nd A ve r ikh in 1 975 , H e rz 1 902 , 1 904 , Gor lov a 1 982 ) . Howeve r , R a t t ray , C o rmack a nd S tap le s ( 1953) n o ted t ha t g ra s s w ou ld h ave b een t h e p ref e r red f ood f o r o n ly p a r t o f t he y ea r ; u pon m a tu r ing a nd d ry ing , i t s n u tr i t iona l q ua l i ty wou ld d im in i sh t o s u ch a n e x ten t t h a t i t wou ld n o t b e u sed . G uy ( 1976 ) a g r eed t ha t g raz ing wou ld h ave o ccu r red o n ly i nt h ew e t s e a son , a nd b row se w ou ld h ave b een p r ef e r red i nd ry s e a son .

B row se r s , s u ch a s r e indee r a nd i b ex , e qu ipped w i th s ma l l mou th s a nd s ma l l s t omach s a dap ted t o as po rad ic f o rage q ua l i ty b ys e lec t iv e ly f e ed ing o nt h e mo s t h i gh ly n u tr i t iou sa nd u n l ig n if ied p a r t s o fp l an t s . T he h i gh ly m ob i le r e indee r , l i ke i t s mode rn c oun te rpa r t s , a dap ted a s t ra tegy o f q u ick ly mov ing a c ros s t h e l a nd scape a s i t s ough t f o rage . Mov ing m any

A s e v idenced a tK rakow S pa c lz i s ta ( Koz low sk i e ta l . 1 974 ) , Do ln i V e s ton ice ( K l ima 1 963 , 1 98 1) , a nd M i lov ice ( O l iva 1 988 ) , m ammo th i n hab i ted C en t ra l E u rope b e tween 2 8 ,000 a nd 2 1 ,800 B P . B y 2 2 ,000 B P s i ze r e duc t ion a nd t w is ted mo la r si n d ica te t ha t m ammo th sw e re u nde rgo ing s t re s sr e la ted t oe nv i ronmen ta ld e te r io ra t ion ( Kub iak a nd Z a l c rzew ska 1 974 , We s t 1 990 ) . B y 1 8 ,000 B P r e sou r ce p a tche s d ec rea sed t o s uch a ne x ten tt ha tt h ey c ou ld n o t s u ppor t s uch l a rge a n ima l s . G uy ( 1976 ) s howed t h a t m a le e l ephan t s c on sume b e tween 1 75 a nd 2 00 k g o f v ege ta t ion p e r d ay ; V e ra shchag in a nd B a ry shn ikov ( 1982 ) p r ed ic ted t h a t a du l t m ammo th s wou ld

h und red s o fk i lome te r s , t h e r e indee r c ou ld s e ek a v a r ie ty o f f o rage s mov ing b e tween p a tche s d i spe r sed o ve r w ide a r ea s . A dd i t iona l ly , r e indee r p o s se s s s pe c if ic , s pec ia l ized r umen b ac ter ia c apab le o fd i ge s t ing av a r ie ty o fl i chen s -f o rb s w h ich a r e l a rge ly u nd ige s t ib le b y o the r u ngu la te s ( Deho r i ty 1 975 , P e r son e t a l . 1 975 ) . R a the r t h an m ov ing a c ro s s ab road l a nd scape , t h e i b ex a dap t ive ly moved a l t i tud inaly , s e ek ing y oung b row se a nd g ra s se s a st h ey b ecame a va i lab le o n s t eep

h ave r e qu i red 2 00 t o 3 00 k g o f " s uccu len t " f e ed d a i ly . I n c on t ra s t , t h e h o r se , ah i ndgu t f e r tnen te r s pec ia l iz ing i n h i gh f i be rf o rage s , r equ i re sl e s st han 1 0k go fv ege ta t ion p e rd ay t o m a in ta in i t se lf (W iloughby 1 975 :142 ; T ab le 2 2 a nd B radbu ry

m oun ta in s l ope s . A lp ine v ege ta t ion g ene ra l ly i s o f i n termed ia te f i b ro s i ty a nd o f r e la t ive ly h i gh q ua l i ty ( Foo se 1 982 : 1 33) .

a nd We rk 1 974 :46 ; T ab le 6 ) . A dd i t iona l ly , " . .s to rage c apab i l i t ie s o f mamma l s a r e i s ome t r ic w i th b ody s i ze wh i le m e tabo l ic r e qu i remen t s a r e f r ac t iona l ly e xponen t ia l " ( Foose 1 982 : 1 4 3) . I n ad e s s ica t ing e nv i ronmen t , m ammo th s , l i ke e l eph an t s , w ou ld h ave b een f o r ced t o t e the r t h em se lve s t o w a te rho le s o r r i ve r s , t hu s d ep le t ing v ege ta t ion i n t h e i mmed ia te a r ea o ft h e se w a te rs ou r ce s . 4 1

T he m ammo th , a l though m o re ef i c ien t t h an r um inan t s , r e qu i red a t r emendou s a moun t o f f o rage t o m a in ta in i t se lf Mammo th sw e re a t ad i sadvan tage i na nu n s tab le e nv i ronmen t w he re p l an t r e sou rce s m ay h ave b een o f l ow q ua l i ty a nd s po rad ica l ly d i s t r ibu ted . W i th l a rge m a s s a nd a wkwa rd l i mbs , t h e m ammo th c ou ld n o t f i nd f o rage i nr ugged t e r ra in l i ke t h e

n imb le i b ex . N o rc ou ld i tm ig ra te o ve rl o ng d i s tance s l i ke t h e r e inde e r . F o r t h e h eavy e l ephan t , w a lk ing r e qu i re s t w ice a s m uch e ne rgy a ss t and ing . T he c a r ibou , o nt h eo t he rh and , u se s o n ly 2 0% m o r e e ne rgy w hen i tw a lk s a f te r s t and ing ( Gu th r ie 1 990 ) . T he r ef o re , i t w a s e ne rge t ica l ly i n ef f i c ien t f o r m ammo th s t o m ig ra te l o ng d i s tan ce s a nnua l ly i nt h e i r q ue s t f o r r e sou r ce s i n t h e C en t ra l E u rope an P l a in . T he se m onoga s t r ic h e rb ivo re s , l i ke t h e s ma l le r r um inan t g r aze r s , c ou ld n o ta dap tt o ar e la t ive ly a r id , l e s sp roduc t ive w o r ld , a nd o p ted t om ig ra te t ot h e s ou th a nd e a s t w he re e a s i ly d i ge s t ib le r e sou r ce sw e r em o re a bundan t . E nv i ronmen ta l d e te r io ra t ion , b e tween 2 0 ,000 a nd 1 5 ,000 B P p roba b ly r e su l ted i np a tchy r e sou r ce s w i th in am o re g ene r a l ly a r id e nv i ronmen t . I n c rea s ing a r id i ty r e su l ted i n p l an t c ommun i t ie sw h ich w e r em o r eh i gh ly f i b rou sf o r ag r ea te rp a r t o ft h ey ea r . D ec rea sed p roduc t iv i ty a s soc ia ted w i th d ec rea s ing q ua l i ty o ff o rage s p roduced as e a sona l " bo t t leneck " t ow h ich l a rge , g r az ing r um inan t s a nd m onoga s t r ic m ammo th s w e re i n to le ran t . L ow e red p l an t a va i lab i l i ty a nd p roduc t iv i ty f o r ced t h e se a n ima l st om ig ra te t oa r ea se a s ta nd s ou th o ft h eC en t ra l E u ropean B a s in . H o r se s , l a rge m onoga s t r ic g r aze r s , a dap ted t o d e te r io ra t ing c ond i t ion s b y i n c rea s ing f e ed ing t ime , i n ge s t ing l a rge q u an t i t ie s o fh i gh f i be r f o rage s , a nd i n c r ea s ing t h e s i z e o f e xp lo i ted c o r e t e r r i to r ie s . R e indee r , p o s se s s ing s p ec ia l ized r umen b ac te r ia , f e d o nf o rage sl a rge ly u nd ige s t ib le b y o t he r u ngu la te s . T hey o p ted t o q u ick ly m ove a c ro s s t h e l a nd scape b e tween p a tche s o f v ege ta t ion l o ca ted i n r i ve r v a l ley s . I b exe s , f e ed ing o n l e s s f i b rou s b row se i n r ugged t e r ra in , m oved a l t i tud ina l ly b e tween s e a sona l p a tche s . E p ig rav e t t ian h un te r s , u n l ike t h e i r G rave t t ian f o rebea re r s , s h if t ed t o am o re m ob i le l i f e s ty le a st h ey p r eyed u pon u ngu la te s p ec ie s w h ich w e re t h em s e lve s b ecom ing m o re m ob i le a nd s ub sequen t ly u np red ic tab le . C hap te r F ou r d i scu s se s h un t ing s t ra teg ie sa nd r e sou r ce m ob i l i ty .

4 2

Chap ter 4 Hun ter sa nd T he ir P rey m oved t oc amp s i te s .

A s a n ima l s b ecame m o re m ob i le a nd r e sou r ce s m o re u np red ic tab le d u r ing t h e L a s t G la c ia l Max imum , C en t r a l E u ropean h un te r s i n c rea s ing ly d epend ed o n s k i l l , t e amwo rk , s c hedu l ing , a nd c r uc ia l t e chno log ica l i n ven t ion s t o s u rv ive . Y e t , l i t t le i sk nown a bou t s p ec if i c s t ra t eg ie st h a tE p ig rave t t ian h un te r s u s ed t o r e duce e n e rgy / r i sk s a nd i n c rea se f ood / te chno log ica lr e tur n s .

A l though as c ena r io o fd i f f e r en t ia lt r an spo r to fb one sb a sed o n b ody s i ze a nd d i s tan ce t o p l ace o f c on sump t ion i sc l a s s ic , o f ten c i ted , a nd u ndoub ted ly t r ue , i ti s o ve r ly s imp l i s t ic . D e c i s ion s b y a nc ien t h un te r s , a st o w h ich p a r t s o fl a rge a nd m ed ium s i zed u ngu la te st ot r an spo r tt oc amp s , g ow el b eyond m e re ly l e av ing b one sb eh ind t ol i gh ten t h el o ad .

E nv i ronm en ta l s h if t s o f t h e LGM n e ce s sa r i ly i mpa c ted t h e d en s i ty a nd v a r ie ty o f a n ima l r e sou r ce s a va i lab le t o E p ig rave t t ian h un te r s( s ee C hap te rT h r ee ) . C o s t si n c rea sed a s p r ey p opu la t ion s d e c l ined a nd s e a r ch t ime i n c rea sed . E p ig r ave t t ian h un te r s r e sponded t o t h i s r e sou r ce s h i f t b y k i l ing , b u tche r ing , t r an spo r t ing , p roc e s s ing , a nd s t o r ing p r ey s p ec ie s d if e ren t ly t o e x t r ac t t h e m o s t r e tur n s i n f ood a nd t e chno log ica l r aw m a te r ia l s f o r t h e l e a s t r i sk a nd e ne rgy o u tpu t .

B ody C ond i t ion : T he c ond i t ion o ft h e a n im a l b a sed o n s e x , a ge , a nd s e a son w e re a l i mpo r tan t i nd ec id ing w h ich a n ima l s t o p rocu r e a nd w h ich b ody p a r t st ot r an spor tt oc amp sf o rf u r the rp roce s s ing . I n n o r ther n l a t i tude s , p l an t r e sou r ce s d ec rea se i na va i lab i l i ty a nd n u t r i t ion v a lue d u r ing w in te r a nd e a r ly s p r ing . H o la r c t ic h e rb ivo re s h ave a dap ted t o t h i s s e a sona l s ho r tage b y c on sum ing l a rge q uan t i t ie s o f f o rage d u r ing s e a son s o f a bundance a nd c onve r t ing t h i s f o rage t o s t o red e ne rgy i nt h e f o rm o f s u bcu taneou s a nd v i sce ra l f a t a nd m a r row i n m edu l la ry c av i t ie s o ft h e b one s . H un te r s a nd o t he r p r eda to r s p r ef e r f a t tened a n ima l s a s f a t i s n ece s sa ry t o m e tabo l iz e p ro te in a nd t o o b ta in c a lo r ic e ne rgy ( Spe th a nd S p ie lmann 1 983 ) . I ff a t i si ns hor t s u pp ly , l a rge q uan t i t ie s o fm ea t m u s t b e c on sumed t o o b ta in t h e s ame a moun t o fe ne rgy f o und i n s ma l le ra moun t s o ff a t . I ff a t i st o ta l ly a b sen t i nt h e d i e t , v e ry l e an m ea ts h ou ld b ea vo ided a si tt a ke s ah i ghe rm e tabo l i sm t o d i ge s t p u re ly l e an m ea t a nd p r eda to r s , i n c lud ing h um an s , c an r e ad i ly l o se w e igh t o n al e an m ea t d i e t ( Spe th a nd S p ie lmann 1 983 ; S pe th 1 987 ) . D u r ing r e sou rce s t re s s , a cqu i s i t ion o ff a t i sm o r ec r i t ica lt h an a cqu i s i t ion o fm ea t( Spe th 1 987 ) .

P a t ter n s o f h uman b ehav io r d u r ing t h e E p ig rave t t ian a r e r e f lec ted i n t y pe s , p a r t s , a nd q uan t i t ie s o f p r ey c a r ca s se s h um an s t r an spor ted , p roce s sed , a nd d i sca rded a t c amp s . S t ra teg ie s i n a n ima l s l augh te r ing , b u tche r ing , t r an spo r t ing , a nd p roc e s s ing a r e r e la ted t o t h e i n t r in s ic n a tu re o ft h e f o od s uppl i e r -agg rega t ion , d en s i ty o f p opu la t ion , m ob i l i ty ( Chap te r T h ree ) , a s w e l l a s p r ey s i ze , b ody a nd b one f a t c ond i t ion , a nd p o ten t i a l f o r t e chno log ica l ( n on f ood ) r e sou r ce s : h i de , a n t le r , s i new , a nd b one . P r ey s i ze , b ody c ond i t ion , a nd m a r row /g r ea se p o ten t ia l a r e d i scu s sed b e low a nd a r e s p ec if ica l ly a dd re s sed f o r h o r se s a nd r e indee r i n C hap te r sF i ve a nd S i x .

S ub s i s ten ce S t ra teg ie sa nd B ody S i ze : P rocu r emen t , p roce s s ing , a nd t r an spor t c o s t s a r e o f ten a s soc ia ted w i th a n ima l s i ze a nd d i s tan ce f r om k i l l t op l a ce o f c on sump t ion . Wh i te ( 1 952 , 1 953 , 1 9 54 ; O 'Conne l le ta l . 1 988 ; Me t ca lf e a nd J one s 1 9 88 ) s ugge s ted t h a t b one s o fl a rge p r ey w e re f r equen t ly s t r ipped o fm ea t a t ak i l l t or e duce t r an spo r t c o s t s . U ppe r l i mb e l emen t s w e re m o r e l i ke ly t o b e r emoved f r om ak i l l a st h ey b o r e g r ea t a moun t s o fe d ib le m ea t a nd f a t r e la t iv e t o t o ta l e l emen t w e igh t . P e rk in s a nd D a ly ( 1 968 ) p ropo sed a" s ch lepp e f ec t " t oa c coun tf o rt h en e tb enef i t sa nd c o s t si n vo lved i nd e c id ing w he the rt oc a r ry o rd i sca rd s e lec ted e l emen t sa t ak i l l s i t e . T he m o s t f r equen t ly u s ed a rg umen t , t h e " s ch lepp e f ec t , " f o r d if e ren t ia l t r an spo r t o fb one s f r om k i l l s i t e s i si l u s t ra ted b y O l sen ( 1 989 ) . S tudy ing t h e b on e s f r om S o lu t ri , s h e ( 1 989 : 2 69 ) s t a ted , " L ike b i son , a u roch s , a n d m ammo th , h o r se s m ay b e f r equen t ly u nde r r ep re sen ted i n o ccupa t ion s i t e s , i n c ompa r i son t o r e indee r a nd o t h e r s ma l le r g ame , b e cau se o fa r e lu c ta n ce o nt h ep a r t o fh un te r st od r ag h eavy b on e so fl a rge g ame b a ck t o t h e b a se c amp u nne ce s sa r i ly ." S uch s c ena r io s h av e b e en c r i t ic ized b y B in f o rd ( 1 984 ) a nd d o n o t h o ld t r ue f o r t h e H adza ( O 'Conne l l e t a l . 1 9 88 ) w he r e a ppend icu la r e l emen t sa l ong w i th t h er e s to ft h ec a rca s sa r ef r equen t ly 4 3

W in te r a nd e a r ly s p r ing a r e t i me s o ff ood s t re s s a nd w e igh t l o s s f o r u ngu la te s ( Spe th 1 987 ) . I nu ngu la te s , f a t i sl o s t i na f i xed s e quence . F a t i sf i r s t m ob i l ized f r om t h e b ack , v i sce ra , a nd f i na l ly t h em a r row ( Spe th 1 983 , 1 987 ) . Ma r row f a t i nt h e l owe rl i mb sa nd m and ib le i st h el a s td epo s i tt ob ed ep le ted a nd i n as t a rv ing a n ima l m ay b e t h e o n ly r ema in ing p a r t o ft h e c a r ca s s w i th f a t w o r th e a t ing ( Spe th 1 983 , 1 987 ) . A s n u t r i t iona l s t r e s s v a r ied s e a sona l ly , h un te r s m ay h ave r e duced r i sk s a nd g a ined t h e m o s t r e tu rn b y : 1 ) se lec t ing s p ec ie s t h a t t e nd t om a in ta in h i gh f a t l e ve ls d u r ing s t re s s p e r iod s , 2 ) se lec t i n d iv idua l s o f ag i ven s p ec ie s , b e cau se o f a ge , s e x , a nd r e p rodu c t iv e s t a tu s , t h a tw e r el e s s s u scep t ib le t of a t d ep le t ion d u r ing ag i ven s e a son , 3 )s e lec t f o r s k e le ta l p a r t s l e a s t p r one t of a tm ob i l iza t ion , o r4 ) ac omb ina t ion o ft wo o rm o r e o ft h e a bov e ( Spe th 1 987 : 2 2 ) . S uch s e lec t ion m ay b e r e f lec ted i n t h e b u tche r ed p o r t ion s o f p r ey t h a t E p ig rave t t ian h un te r s o p ted t ot r an spo r tf r om k i l l st oc amp s i te s . Ma r row /G rea se R e sou r ce s : H un te r s w e re a wa re o f t h e m a r row q uan t i t ie s o f d if e r en t b one sw i th in as i ng le c a r ca s s a nd s t ra teg ica l ly l e f t b eh ind l e an c a r ca s s p a r t s a nd a s soc ia ted b one s t h a t w e re p oo r i nm a r row a nd c o s t ly t o b r eak a nd t r an spo r t . C onve r se ly , h un te r s c on s i s ten t ly t r an spo r ted t h o se b ody p a r ts t h a tp o s se s sed s t ab le s ou rce s o f s t ruc tu ra l f a t . T he s e p a r t s c ou ld b e r e la t ive ly

c omp lex a nd c o s t ly t o p roce s s . T ran spo r t t o c amp s a nd a s soc ia ted p roce s s ing s t ra teg ie s o p t im ized e x t ra c t ion f o r a m ax imum r e tu rn i nm ea t a nd f a t . B inf o rd ( 1978 ) d eve loped a s e r ie s o f e conom ic u t i l i ty i n d ice s t o r a nk t h e c ompa ra t ive v a lue o fs h e ep a nd c a r ibou b ody p a r t s . H et e s ted t h i sm ode lo f u t i l i ty b y d ocumen t ing t h e k i nd s o fb ody p a r t s t h a t E sk imo s a b andoned a tk i l l s i te s . H igh r a nked c a rca s s p a r t s w e re m o re l i ke ly t o b e t r an spo r ted f r om k i l l s t h an l owe r r a nked p a r t s . Me tca lf e a nd J one s ( 1 988 ) s howed t h a t e l emen t r a nk w a s a s soc ia ted w i th n o t o n ly m ea t v a lue s , b u t a l so t h e a moun t o f m a r row a nd g rea se a t ta ched t o ab one . T ho se b one s w h ich c omb in ed m ea t q uan t i ty w i th h i gh m a r row a nd g r ea s e c on ten t w e r e m o r e l i ke ly t o b e t r an spo r ted t o c amp . Me tca lf e a nd J one s( 1 988 )a l so t ook i n to a c coun tt h e e conom ic a na tomy o f d if e ren ts p ec ie sa nd t h ep o ten t ia lf o rb one e l emen t st os u rv iv e i nt h ea r cha eo log ica lr e co rd .

t h i s c hap te r p r e sen t s am ode l s ugge s t ing t h e m o s t e f i c ien t m e thod so fp r ocu r ing a nd p r oce s s ing u ngu la te s a nd f i rbea r e r s d u r ing a n a nnua l c yc le o f t h e E p ig r ave t t ian h un te r . F i r s t , p rocu remen t a nd p roce s s ing s t ra teg ie s o f i n d iv idua l p r ey s p ec ie s a r e d i scu s sed . T hen , s e a sona l m ode l s f o r h un t ing , / t rapp ing a lp r ey s p e c ie sa r ep r e s en ted . R anpf e rt a rand u s : R e indee r L a r te t a nd C h r i s ty ( 1 875 ) r e por t ing a n a bundan ce o fr e indee r b one s i nc ave s o ft h e D o rdogne , d e s igna ted t h e Magda len ian " L 'Age d u R enne . " B u r ch ( 1 972 : 39 ) s t a ted t h a tt h er e indee r " . .may w el b et h es pec ie so fs i ng le g r ea te s ti mpo r tance i nt h e e n t i re a n th ropo log ica l l i t e ra tu re o n h un t ing ." R e indee r c a n p rov ide n e a r ly a l h uman s u b s i s ten ce r e qu i remen t s : m ea t , e n t ra i l s , s t om a ch c on ten t s , b l ood , f a t , m a r row , a l ong w i th m i lk f r om s l augh te red , l a c ta t ing c ow s . Moder n c a r ibou h un te r s p rov ide e v idence t h a t a nc ien t h uman s c ou ld h ave r e l ied o n p a r t ia l ly d i ge s ted g u t c on ten t s a nd f e ce s o fr e indee r t o f u lf i l l n u t r i t iona l r e qu i r emen t s (W i lk in son 1 975 ) . A n t le r s c an b ee a ten w h i le i ns umme rv e lve t . Of a lc an b eu s ed t ob a i t t r aps f o r f u r bea re r s . R e indee r f a t c an b e b ur ned f o r l i gh t . L ong s t ra igh t m e tapod ia a nd a n t le r s c a n b e f a sh ion ed i n to e f i c ien t h un t ing /p roce s s ing t oo l s . A r t icu la r e nd s o f l o ng b one s c an b e c r u shed a nd b o i led t o e x t r a c t g rea se ( B in f o rd 1 978 , S p ie s s 1 979 ) . Af te r t h e J u ly m ou l t , t h e l i gh t ly h a i red h i de s o fA ug u s t /Sep tembe r c an b e t a nn ed a nd s ewn t om ake l i gh twe igh t u nde rga rmen t s a nd s umm e r c l o th ing . L a te a u tumn /ea r ly w in te r h i de s w i th d en se f u r c an b e m ade i n to b l anke t s , r o be s , b oo t s , h eav yw in te r o v e rga rmen t s , a nd t e n t s . S k in s c an a l so b e e a ten a s s t a rva t ion f o od s . S i new s a nd t e ndon s p rov ide c o rdage . H ide s , g u t s a nd b l adde r s w ou ld h ave b een u se f u lc on ta ine r s .

S ub s i s ten ce D ec i s ion s Ma sked b yA t t r i t ion : D if f e r en t ia l r e p re sen ta t ion o f b one e l em en t s a t ag i v en s i t e o f t en m ay b e b i a sed b y a t t r i t iona l p roce s se s w h ich s e le c t ive ly r emov e s ome b one s f r om t h e a s semb lage . S uch a t t r i t iona l p roce s se s m ay m a sk e v idence o f d ec i s ion s b y p r eh i s to r ic h un te r s . L yman ( 1984 , 1 994 ) c r i t ic ized e a r l ie r a pp roache s t o b ody t r an spo r t s t ra teg ie s b y i d en t if y ing b one s , b a sed o n d en s i ty v a lue s , t h a t d if e ren t ia l ly s u rv ive i nt h e a r chaeo log ica l r e co rd . B one s r e cove red f r om k i l l o r r e s iden t ia l a r ea s a r e a f ec ted b ya t t r i t iona lp roce s se sa s soc ia ted w i th b one d en s i ty . F u r the rS ub s i s tence S t r a teg ie s : G ene ra l ly , h un te r s t e nd t o c hoo se t h e h i ghe s t r e tu rn f o r t h e l e a s t e f o r t a nd r i sk ( Z ipf 1 949 , J och im 1 976 ) . I n a ny h un t ing , /f o rag ing s t ra tegy , b a lance s m u s t b e m e t b e tween c o s t s / r i sk a nd r e tu rn s . C o s t s c an b e a s soc ia ted w i th : 1 ) techno logy , 2 ) t ran spo r ta t ion , 3 ) co l le c t ion , 4 ) p roce s s ing , a nd 5 ) s to rage ( Ea r le 1 980 ) . R educ ing c o s t s c an b e f a c i l i ta ted b y ar e o rgan iza t ion i ns u b s i s tence p a t te rn s . H un te r /ga the r e r s , l i v ing i nt empe ra te o r a r c t ic e nv i ronmen t s , w he re r e sou r ce s a r e c l umped a nd t empo ra l ly d i spe r sed , r e qu i re i n c rea sed o rg an iza t ion o fs u b s i s ten ce s t ra teg ie s . E thnog r aph ic e v idence s how s t h a t t h e se h un te r s p l an w el o rgan ized h un t s o fl a rge b od ied a n ima l s a nd s t o r e f o od f o r s e a son s o f l owe r ed a bundan ce . R e sou rce s a r et a k en i nq uan t i ty d u r ing s e a son s o f a bundan ce a nd a r es t o r ed ( B inf o rd 1 980 ) .

T he p a leoeconomy o fr e indee r h un t ing v a r ie s s e a sona l ly w i th a n ima l s o fd if e r en t a ge s a nd s e xe s . B o th c ow s a nd b u l l s g a in w e igh t t h roughou t s umm e r a nd a r e f a t te s t d u r ing S ep tembe r /e a r ly O c tobe r ( F ig . 4 . 1) . R u t t ing a c t iv i t ie s q u i ck ly d ep le te f a t r e s e rve s a nd t h eb ody c ond i t ion o fb u l l s s ot h a tb y N ovembe r , b ul s a r e i nt h e i rp oo re s t a nnua l c ond i t ion . C ow s , o n t h e o t he r h a nd , s l ow ly d e te r io ra te o v e r w in te r a nd a r e i n p oo re s t c ond i t ion i n l a te w in te r a nd s p r ing a f te r b i r th s o f c a lve s . Ma r row y i e ld s , a l ong w i th f a t a nd g en e ra l b ody c ond i t ion s , v a ry w i th t h e s e a son s . A s a n a n ima l b e come s i n c r ea s ing ly m a lnou r i shed , i t s b ody r e se rve s d im in i sh i n r a nked o r de r . S ubcu taneou s f a t d ec rea se s f i r s t , f o l low ed b y v i s ce ra l f a t , m a r row , a nd f i na l ly b r a in f a t ( Ha r r i s 1 945 , R iney 1 9 55 , R an som 1 965 , S pe th 1 983 , S t ine r 1 990 ) . Ma r row d ep le t ion c ommence sf r om t h ep rox ima le nd o ft h el i mb sw i th f r on t l e g s m o re s u scep t ib le t od ep le t ion t h an h i nd l im b s ( Ra tcl if e 1 9 80 , B rook s e ta l . 1 977 ) . B e cau se m a r row i so ne o ft h e l a s t f a t s ou rce s t o b e d ep le ted , i t c a n b e p r ed ic ted t h a t b u l l s , e xhau s ted f r om t h er u t , w ou ld b ea rl i t t le u s ab le m ar row i nl a t e f a l l /ea r ly w in t e r . C ow s , a t t h e i r p oo r e s t c ond i t ion a f t e r b i r th ing , w ou ld p r ov ide p oo r e s t m a r row y i e ld s i nt h e s p r ing . A s a n ima l s w e r e t a ken i nd if e r en t s e a son s , s p ec if ic e l em en t s o fm a le s a nd f ema le sw e re t e s ted f o rm a r row q ua l i ty a nd u s ed o re l im ina ted . T he N unam iu tp r a c t ice s u ch as t ra tegy ( B in f o rd 1 978 ) .

C l ima t ic d e te r io ra t ion o ft h e L a s t G la c ia l Max imum c au sed h un te r st oi n c rea se t h e i r u s e o fe x i s t ing s t ra teg ie s a nd /o r a dd n ew s t ra teg ie s . P o s s ib le i n c rea sed c o s t s a s soc ia ted w i th l owe red a bundance d u r ing t h eL a s t G la c ia l Max imum r e su l ted i n : 1 )mo re f r equen t m ovemen t s o f ac amp u n i t d u r ing t h e c ou r se o f as e a son , 2 ) t rave l l ing f a r the rf r om ag i ven c amp f o r r e sou r ce s , 3 ) inc rea sed s e a r ch t i me f o r ag i ven r e sou r ce , 4 )po s s ib le p rocu remen t o fr e sou r ce s o fl e s se r q uan t i ty a nd /o r s i ze , o r5 ) ac omb ina t ion o ft wo o rm o r eo ft h ea bove ( B inf o rd 1 980 , W in te rha lde r a nd S m i th 1 98 1 , K e l ly 1 992 ) . T he se i n c rea sed c o s t s c ou ld b e p o ten t ia l ly o f se t b y o rgan iz ing s e a sona l s t ra teg ie s ( ma s s k i l l s , s t a lk ing , a nd n e t t ing o r t r app ing ) s ot h a t d e c rea sed a nd /o r d i spe r sed s p e c ie s c ou ld b e m o re e f i c ien t ly e xp lo i ted . C ha rac te r i s t ic s o f p r ey s p ec ie s : d i s t r ibu t ion , d en s i ty , s o c ia l b ehav io r , c ond i t ion , a nd s e a sona l i ty i n f luenced h ow s p e c if i cr e sou r ce sw e re c ho sen a nd p rocu red . U s ing c l ima t ic d e te r io ra t ion a nd u ngu la te e co logy ,

H ide q ua l i ty o fr e indee r a l so v a r ie s s e a sona l ly . Af te r as p r ing 4 4

J u ly Aug .

C oa tC ond i t ion

J une

F ema le C ond i t ion

S ept .

May

Apr .

Oct .

Nov .

Mar .

D ec .

F eb .

J an .

F i g . 4 . 1 . S easona l a t t r ibu te s o fr e indee ru sef u l i nr e con s truc t ing P a leo l i th ic f ood / te chno logy m anagemen t s y s tem s .

4 5

mou l t , r e indee rg row s of t , s hor t h a i r st h roughou t s umme r . A s w in te r a pp roache s , g ua rd h a i r s c on t inue t o l e ng then a nd a n u nde r coa t d eve lop s . B y m idw in te r , a g ing h a i r sb eg in t o b reak a nd g i ve t h e a n ima l am o t t led a ppea rance . P a ra s i te s a l so s e a sona l ly d amage h i de s . Wa rb le f i e s l a y e gg s o n c oa t s o f c a r ibou i nJ u ly /Augu s t . E gg s h a tch d u r ing w in te r a nd l a rvae b o re i n to t h e s k in o f t h e r e indee r . A s l a rvae g row , t h ey i n c rea se t h e s i ze o f b rea th ing h o le s , s i gn if ican t ly d amag ing s k in s f r om Novembe rt oJ une . Af te rw a rb le f l y a du l t s e me rge

a l ong t h es ame r ou te sf o rs eve ra l s e a son s . A sf o rage sd ec rea se i n ag i ven a r ea , r e indee r m ay c omp le te ly s h if t t h e i r m ig ra t ion r ou te s a nd d i sappea r . E thnog raph ic e v idence s ugge s t s t ha t s uch s h if t si nm ig ra t ion r ou te sa re t o ta l ly u np red ic tab le . I fr e indee r p opu la t ion s d ec l ined d ue t od ec rea sed f o rage o r i f m ig ra t ing a n ima ls c hanged r ou te s , s ub s i s tence w ou ld h ave b ecome i n c rea s ing ly p reca r ious . U nde r t he se c ond i t ion s , h un te r s c ou ld o p t t o c hange l o ca t ion s t h em se lve s o r t u rn t o o the r s ou rce s o fg ame . C la rke ( 1940 ) s ugge s ted t h a t c a r ibou

f r om t h e s k in , i tq u ick ly h ea l s . S ca r red , b u t h ea led , l a te s umme r /e a r ly f a l ls k in s , s po r t ing s of t , s hor t h a i r s , a r ee xcelen t f o r c l o th ing ( Banf ie ld 1 962 ) . H eav i ly f u r red , b u t i n sec t d amaged , w in te r h i de s p rov ide b l anke t s , r obe s a nd t e n ts . T h rough i n te rv iew s w i th A thapa skan c a r ibou h un te r s , B anf ie ld ( 1962 ) c a lcu la ted t h a t t we lve c a r ibou h i de s a r e r equ ir ed t o c l o the o ne p e r son e ach s e a son : t h ree f o r i n ne r

u nde rgo c y c l ic f l uc tua t ion s . S ubsequen t c oun t s o f c a r ibou h e rd s i nN o r th Ame r ica b y B an f ie ld ( 1 954a , 1 954b ) , K e lsa l l ( 1968 , 1 970 ) a nd S koog c on t inua l ly m ove , s h if t ing a nnua l ly , a nd o ve r l o nge r B ecau se o f t h i s s h if t ing ,

p arka , f ou r f o r o u te r p a rka , t wo f o r i n ne r t r ou se r s , t wo f o r o u te r t r ou se r s , o ne f o r m i t t s , s t ock ing s , a nd b oo t s . Twen ty -

( 1968 ) s uppor t t h i s . R e indee r t he i r l o ca t ion d a i ly , s e a sona l ly , p e r iod s o f t ime ( Bu r ch 1 972 ) . B u rch ( 1 972 : 3 56 ) s t a ted t h a t

" . .hum an p opu la t ion s l a rge ly d ependen t u pon c a r ibou w i l l b e f a ced w i th a m a jo r r e sou r ce c r is i s a t l e a s t e ve ry 2 t o 3 g ene ra t ion s . " T h i si sr e la ted t oL ieb ing ' sL aw o ft h eM in imum ( c i ted i nO dum 1 971 :106 ) b a sed o n t h e p rem i se t h a t " . .an

f ou r h i de s a r e r equ i red p e r p e r son p e r y ea r f o r t he t wo s ea son s . Twen ty o r mo re h i de s a r e r e qu i red t o m ake a t e n t ( Banf i e ld 1 962 ) .

o rgan i sm i sn o s t ronge rt h an t he weake s t l i nk i ni t se co log ica l c ha in o fr e qu i remen t s ." I tm ake s n o d if e r ence i nt h e l o ng r un i fr e inde e ra ppea rb yt h et hou sand sf o r at i me , b ecau se i ti st h e p oo re s t o r l e ane s t y ea r s w h ich s e t t h e l i m i t t o l o ng te rm

R e indee r f i gu re h eav i ly i n to t h e e conomy o f p reh i s to r ic h un te r s i ng l ac ia l e nv i ronmen t s b ecause , w hen e ncoun te red , t he a n ima l sa r ee a s i ly k i led ( Zeune r 1 963 , C la rk 1 967 , K e l sal

r e l iab i l i ty o fr e indee rh un t ing . B u rch ( 1 972 )p o in ted o u tt h a t i t o n ly t a ke s s e ve ra l w eek st os t a rve t od ea th . Unde r s u ch s t re s s s i tua t ion s , p r eda to r s h ave t h ree a l ter n a t ive s : 1 ) popu la t ion d ec rea se t h rough s t a rva t ion a nd /o r e m ig ra t ion , 2 ) exp lo i t d if e ren t r e sou rce s , o r 3 ) a c omb ina t ion o f t he a bove t wo ( S lobodk in 1 96 1) . N ine teen th c en tu ry N ua t iqm iu t e sk imo s , o f t h e w e s te rn B rook s R ange , s t a rv ing d u r ing c r i se s a s soc ia ted w i th d e c l in ing a nd s h if t ing r e indee r p opu la t ion s , e m ig ra ted 6 00km i n land a t temp t ing t o f i nd t he m i s s ing r e inde e r ; o the r , s ma l le r g roup s moved t o t h e c oa s t t o e xp lo i t s e a m amma ls ; s t i l l o the r sr ema ined b eh ind t o ek o u t al i v ing i nt h e i ro r ig ina l t e r r i to ry ( Bu r ch 1 972 ) . S uch ap a t te rn o fd i r th a nd p l en ty , a nd r e sou r ce s w i tch ing , m ay b e r ef lec ted i nC en t ra l E u rope c i rca 1 8 ,000 B P .

1 968 , B u rch 1 972 ) . R e indee r : 1 ) no rma l ly o ccu r i n g roups ; 2 )move i nd ef in i te d i re c t ion s w h ich c an b e e a s i ly i n te r cep ted b yp reda to r s ; 3 )a re d oc i le , r e sor t ing t of l i gh t r a the rt h an f i gh t , a nd a r e n o t d ange rou s t o t h e ir p reda to r s ; 4 )pos se s s p oo r e ye s igh t a nd c anno t d i s t ingu ish d ange rous f r om h a rm le s s movemen t s ; 5 )a re n o t a l e r t u nde rs t re s s c ond i t ion s a nd d u r ing m ig ra t ion s ; 6 )a re c u r iou sa nd a t t rac ted t ou nu sua l o b jec t s a nd movemen t s ; a nd 7 )f o l low p red ic tab le r ou te s o fm ig ra t ion f o r s eve ra ls e a son s . Wo r ldw ide c a r ibou / re indee r h un te r s h ave d eve loped , s ome t ime s i n gen iou s , s t ra teg ie s t o d i rec t , c ap tu re a nd s l augh te r t h e i r p rey ( Tab le 4 . 1) . D e sc r ibed t e chn ique s a r e h i gh ly d ependen t u pon w ind d i rec t ion , t opog raphy , p resence o ra bsence o fw a te r , s i ze o ft h e r e indee rb and , t h e s ea son ( on

S tu rdy ( 1975 )p ropo sed t h a ta nc ien t r e indee rh un te r sp robab ly r e l ied u pon o ne o ft h ree s t ra teg ie s : 1 ) they f o lowed t h e h e rd s , 2 ) they l i ved o n t h e r ange o ccup ied b y t h e h e rd f o r o n ly o ne s e a son , o r3 )t h ey p l a ced t h em se lve s o n m ig ra t ion r ou te s a nd a t temp ted t o k i l l e nough a n ima ls d u r ing s p r ing a nd a u tumn

m ig ra t ion r ou te so rd i spe r sed a nd f e ed ing q u ie t ly ) , p re sence o f s now , a nd a v a i lab i l i ty o ft e chno logy o ft h e h un te r ( i nc lud ing s k i s , s l ed s , d ome s t ica ted a n ima ls ) . P reh i s to r ic r e indee rh un te r s w e re p robab ly a sr e sou rcef u l .

m ig ra t ion s t o s uppor t t h em se lve s y ea r r ound . C la rk ( 1967 ) b e l ieved t h a t r e indee r h un te r s mus t h ave f o l lowed t h e i r p r ey

T h ree a spec t s o f t h e d i s t r ibu t ion o f a n ima l r e sou rce s a r e r e levan t t o h un t ing s t ra teg ie s : 1 ) change s t h rough t ime i nt h e

d u r ing t h e i r a nnua l c yc le . U s ing r e indee r a n t le r d eve lopmen t , B ouchud ( 1966 ) c onc luded t h a t a l though r e indee rw ou ld h ave o n ly l i ved i nt h e v i c in i ty d u r ing t he s p r ing a nd a u tumn , s i te s

d i s t r ibu t ion o f as pec ie s , 2 )va r ia t ion s i nr eg iona l a bundance o f a n ima l s a t a ny g i ven t ime , a nd 3 ) loca l ized d i s t r ibu t ion o f i n d iv idua l p opu la t ion u n i t s o n a s c a le r e levan t t o t h e i r e xp lo i ta t ion b y h uman p opu la t ion s . " The f r equency ,

n ea r t h e V eze re w e re i n hab i ted y ea r r ound w i th a mb i t iou s h un te r s t r ekk ing t o f a r a way r e indee r p a s tu re s w hen h e rd s m ig ra ted . S tu rdy ( 1975 ) s ugge s ted t ha t s u ch a c t iv i t ie s w ou ld l e ad t ou l t ima te s t a rva t ion a sc a lor ies e xpended wou ld e xceed

r e gu la r i ty , a nd e x ten t o f m ovemen t s b y a n ima l s p rof ound ly i n f luence t h e m anne r i n w h ich t h ey c an b e e xp lo i ted " ( W i lk in son 1 975 : 1 0 -11) . R e indee r a r e h i gh ly m ob i le a n ima l s t h a t move t o m a in ta in h e rd h ea l th a nd t o a vo id o ve rg r az ing h i gh ly s e n s i t ive l i chen s . S ea sona l movemen t s o fr e indee r a r e r e la ted t o t h e a moun t o fa va i lab le f odde r , s he l te r a nd s now c ove r ( Dege rbo l a nd K rog 1 959 , W i lk in son 1 975 , B anf ie ld 1 951 , K e l sal 1 957 , 1 968 , 1 969 ) . I f ag i ven a r ea s uppor ts a r i ch h e rb b i oma s so ri fr e indee rp opu la t ion s a r e r e la t ive ly l ow , r e indee rd o n o tt r ave l f a r . I ff o rage i ss c a r ce a nd /o r h e rd s a r e l a rge , r e indee r w i l l t r ave l f a r the r t o f i nd a dequa te f oods . A s l o ng a s f o rage i ss uf i c ien t , r e indee r c an p r ed ic tab ly m ig ra te 4 6

c a lo r ic i n take . T hu s , s e a sona l , r a the r t h an y ea r r ound , e xp lo i ta t ion i sr e p re sen ted a t a ny g iven s i t e ( Bu rch 1 972 ) . S tud ie s o f m ode rn r e indee r /ca r ibou b ehav io r s ugge s t t h a t t h e se a n ima ls c an e a s i ly o u td i s tance t h e i r p r eda to r s ( Cr i s le r 1 956 , K e lsa l l 1 968 ) . T rave l l ing a s m uch a s 6 5 k m /day , r e indee r c an m ove a s m uch a s 8 00km i ns i x w eek s ( Ke l sa l l 1 968 ) . B u r ch ( 1 972 ) a rgued t h a t n o h un t ing g roup , b u rdened w i th p roce s s ing , a nd s l owed b y t he o l d a nd y oung o f t h e g roup , c ou ld h ope t o f o l low r e indee r . H e ( 1972 : 3 51) s ugge s ted t h a t h un t ing g roup s , b y s t ra teg ica l ly p l ac ing

Table 4.1. Ethnographic reindeer hunting strategies. HUNTING STRATEGY

HUNTER

SOURCE

Kutchin

McKennon 1965 Leechman 1954 Osgood 1937 Vorren and Manker 1962 Vorren and Manker 1962 Stefannson 1914 Balikci 1970 Speck 1940 McKennon 1965 Steensby 1910 McKennon 1959 Speck 1940 Forde 1934 Vorren and Manker 1962 Parry 1824 Balikci 1970 Binford 1978 Stefannson 1914 MacNeish 1956 Steensby 1910 Nelleman 1969-70 Gubser 1965 Binford 1978 Jeness 1937 Spiess 1979 Speck 1940 Stefannson 1914 Balikci 1970 Gubser 1965 Binford 1978 McKennon 1965 J. Smith 1976 Osgood 1937 Howley 1974 Vorren and Manker 1962 Popov 1966

Land based: Snare

Pitfall Stalking of foot with disguise Pursuit with special footgear (snowshoes, skis)

Shooting from manmade blinds Drives using guides of stone, moss, snow, wood, leather

Corralling within manmade structures of stone or wood

Fence/snare combination

"

" Lapp Lapp - Copper Eskimo Netsilingmiut Penobscot Indian Chandalar Kutchin Polar Eskimo Upper Tenana Indian Penobscot Indian Yukaghir Lapp Igoolik Eskimo Netsilingmiut Nunamiut Tikkerarrniut Pelly River Indian Polar Eskimo W. Greenland Eskimo Nunamiut Sekani Kaska Indian Penobscot Indian Copper Eskimo Netsilingmiut Nunamiut " Kutchin Chipewyan Upper Inlet Eskimo Newfoundland Beothuk Lapp Nganasan

Water Based: Spearing from kayaks Fence/kayak combination

lgoolik Eskimo Nain-area Eskimo Caribou Eskimo

Drives into waterfalls

Great Slave Lake Indian

Parry 1824 Turner in Taylor 1969 Birket-Smith 1929 Arima 1975 Mason 1946

Tanaina Lapp Nganasan

Osgood 1937 Vorren and Manker 1962 Popov 1966

Using Domesticated Animals: Driving by dogs Domesticated reindeer used as decoys

47

d ome s t ica ted h e rd f o r t h e i r t r ip b a ck t o c a p t iv i ty . H oweve r , S i be r ian s c a p tu red w i ld h o r se s b y d i gg ing p i t s c l o se t o t h e i r w a te rho le s( El ia s 1 873 i n Moh r 1 97 1) a nd m oder nH adza k i l l z eb r a o n f o o t ( O 'Conne l l e t a l . 1 990 , 1 992 ) . I tn e ce s sa r i ly r e qu i red ah i gh d eg ree o fp l ann ing , o rgan iz a t ion , a nd c ou rage t oh un tt h e s es w if t , a l e r t , a nd d ange rou sa n ima l s .

t h em se lve si nt h e" . .bounda ry b e tween t h ea n ima l s 'w in te ra nd s umm e r r a nge s " c ou ld l i ve o n r e indee r y ea r r ound t h rough s l i gh t s h if t s i nl o ca t ion . I n ar e eva lua t ion o fh e rd f o low ing , B u rch ( 1 99 1 : 4 40 ) n o ted , " Peop le w ho ' f o l low t h e h e rd s ' d o n o ta t temp tt ok eep u pw i th as pe c if i c s e to fa n ima l sb u tm ov e i n at i me ly m anne r f r om t h e w in te r r a nge o f t h e b r e ed ing p opu la t ion i nt h et a iga t oi t ss umme rr a nge o nt h et u nd ra ( a nd v i ce v e r sa ) . " U s ing d i a r ie s a nd r e co rd s o fe a r ly m i s s iona r ie s a nd e xp lo r e r s i n N o r th Ame r ica , B u r ch ( 1 99 1) d i s cove red C h ipewyan f am i ly g roup s , b u rdened w i th g ood s , a ve raged 1 3 k m /day w i th am ax imum o f2 3 .5 k m t r ave l led i no ne d ay a s t h e g roup f o lowed r e inde e r h e rd s d u r ing s p r ing m ig r a t ion s . C h ipewyan m en , u nencumbe red w i th f am i ly g roup s a nd p e r sona l g ood s , " a ve raged 3 0 k m p e r d ay , w i th c on se cu t iv e d a i ly s p r in t s o f 6 8 a nd 7 2 k m . " ( Bu r ch 1 99 1 : 4 43 ) . B u rch ( 1 99 1 : 4 43 ) c onc luded : " a l though C h ipewyan m en c ou ld n o t l i t e ra l ly f o l low a b and o f c a r ibou d u r ing t h e i r r a p id J u ly m ovemen t , a t l e a s t s ome o f t h em c ame c l o se t o i t . " T he C h ipewyan i sa ppa ren t ly t h e o n ly e t hnog raph ic g roup t h a t m ove sa t af a s t e nough p a ce t om a in ta in c on tac tw i th r e indee r h e rd s . H e rd f o l low ing , i nt h e s e n se o f C la rk ( 1967 ) a nd B ouchud ( 1 966 ) m ay h ave b e en a v i ab le , a l though e x t reme ly c o s t ly o p t ion , f o rE p ig rave t t ian h un te r s . H e rd f o l low ing i nt h e s e n se o fB u r ch ( 1 99 1) , w he r e h uman g roup s m ig ra ted i n at i me ly f a sh ion t o i n te rcep t r e inde e r h e rd s , w ou ld p rove m o r e c o s t e f ec t ive . A udouze a nd E n loe ( 1 99 1 : 6 4 ) , s t udy ing r e indee r b one s f r om Magda len ian s i t e s i nt h e P a r i s B a s in , d e sc r ibed p o s s ib le s t ra teg ie s t h a t r e indee r h un te r s a dop ted d u r ing d if e ren t s e a son s o f a bundance . U s ing B in f o rd ' s c o l lec to r / f o rage rm ode l , t h ea u tho r s s howed t h a t Magda len ian h un te r s l i v ing i nt h e P a r i s B a s in p r ac t iced a n o rgan iza t iona l m ix o fs u b s i s tence a c t iv i t ie s . A v a i lab i l i ty o fr e indee r ( e n m a s s o r d i spe r sed ) l e d t o s t ra teg ic p o s i t ion ing o fh un te r s a c ro s s a l a nd scape w h ich r e su l ted i n ap a t te rn o fc o l lec t ing o rf o rag ing . D u r ing a u tumn m ig ra t ion s , f a t tened a n ima l si nt h eP a r i sB a s in c ou ld b et a ken e n m a s s , p r oce s sed , a nd s t o red . T h i s l o g i s t ica l b ehav io ri sr e f le c ted a tP i nceven t a nd V e rbe r ie . When a n ima l s w e re s c a r ce , d i spe r sa l o f i n d iv idua l h un te r s a c ro s s t h e l a nd scape i n a" s eek o u t a nd d e s t roy " f a sh ion i n c rea sed t h e p robab i l i ty o f e ncoun ter ing a nd k i l l ing i n d iv idua l s o r s ma l l g roup s o fr e indee r a sw e l l a s al a rge v a r ie ty o fo t he r a n ima l s w h ich w e re i mmed ia te ly s h a red a nd c on sumed b y h un t ing g roup s . T h i s b ehav io r i sr e f l ec ted a tE t io l le s a nd Ma r s angy N 19 . T he s h a r ing o f af ew a n ima l sw ou ld r e su l ti nt h es u rv iva l o f t h e g roup . T h i s b ehav io r i ss uppo r ted b y e xamp le s o f m ode rn c a r ibou h un t ing t e chn ique s w he re a n ima l s c a n b e t a ken e n m a s s d u r ing m ig ra t ion s , b u t t a ken s i ng ly o r i ns ma l l g roup sd u r ing o t he rs e a son s . E quu sc f .p r zewa l sk ig erus : H o r se C ompa red w i th r e indee r h un te r s , f ew e t hnog raph ic e xamp le s p rov ide m ode l s f o r p r eh i s to r ic h o r se h un t ing a nd p roce s s ing t e chn ique s . N ea r ly a l h i s to r ic e qu id c ap tu re i n vo lved t h e u s e o f d ome s t ica ted h o r se s a nd m ode rn h un t ing e qu ipmen t . H i s to r ica l ly , t h e S i be r ian K a lmuck s , h un ted t h e p r zewa l sk i b y r i d ing i n to ah e rd a nd k i l l ing t h em w i th l a nce s ( Moh r 1 97 1) . N ine te en th c e n tu ry E u ropean s , a nx iou st oa t ta in a s ia t ic h o r se s f o r z oo s , u s ed r e lay s o f d ome s t ica ted h o r se s t o t i r e P rzewa l s l c i s u n t i l a du l t s w e r e w i th in r a nge t o b e s h o t . O rphaned c o l t s w e re p rov ided s u r roga te m o the r s f r om t h e

4 8

Meage r e t hnog raph ic s ou r ce s , w h en c omb ined w i th i n f o rma t ion g l eaned f r om t h e d ocumen ted b ehav io r o f w i ld h o r se s , p rov ide d a ta f o r r e con s t ruc t ing a n c ien t h o r s e h un t ing t e chn ique s ( F ig . 4 . 2 ) . A s d e sc r ibed i nC hap te r T h ree , h o r se s a r e g ene r a l ly f o und w i th in t wo d emog r aph ic g roup s : h a r em s a nd b a che lo r h e rd s . H a r em s a r e s ma l l f am i ly u n i t s o fm a r e s , t h e i rf o a l s , a nd ad om inan ts t a l l ion . D a i ly a nd a nnua l a c t iv i t ie s o fh a rem s a r e e x t reme ly p a t te rned . G en e ra l ly , h a rem g roup s l i ve i n s ma l le r h ome r a nge s t o w h ich t h ey a nnua l ly r em a in f a i thf u l . W i th in t h e se r a nge s , t h e g roup t r ave l s a l ong w e l l d ef ined , d ung l i ned t r a i l s . H a rem sd on o tm ove l o ng d i s tance s e a ch d ay o r t h roughou t t h e y ea r a s t h ey a r e s l owed b y t h e y oung . When c o l t sa r es ma l l ,t h eh a rem u s e st h es ame s l eep ing a r ea e a ch e v en ing . S l eep ing a r ea sa r ee a s i ly r e cogn iz ab le f r om a c cumu la t ion s o f d ung . H o r se s r e tur n e a ch d ay o r t wo t o p r ed ic tab le w a te r s ou r ce s . T he d om inan t m a r e a nd h e r f o a l d e te rm ine t i me a nd d i rec t ion o f m ovem en t ; w hen t h e m a re l e ad s , a lf am i ly m embe r si n c lud ing t h es t a l l ion f o low i ns i ng le f i l e . When t h r ea tened , w i ld h o r se s f l ee i ns i ng le f i l ew i th f o a l s i nt h e m idd le a nd t h e s t a ll ion f a l ing t o t h e s i de o f c l o se s t d ange r . I fn o y oung f o a l s a r e p r e sen t , t h e s t a l l ion t a ke s t h e l e ad w i th m a r e s f o low i ng . I fc or ne red , s t a l l ion sw i l l f a ce a nd a t tack p r eda to r s , k i ck ing , b i t ing , a nd s t omp ing . B a che lo r g roup s , s i gn if i can t ly l e s s p r ed ic tab le t h an h a rem g roup s i nb o th d a i ly a nd a nnua l b ehav io r , u se a l a rge r c o re a r ea a nd t r av e l l o nge r d i s tance s o n a ny g i ven d ay . Y oung b a che lo r g roup s a l so v a ry i n t h e i r c o r e a r ea s e lec t ion , r e tu rn ing l e s st h an 2 0% o ft h et i me t o at e r r i to ry e xp lo i ted t h e p r ev iou s y ea r . U nh inde r ed b y y oung c o l t s , b a che lo r g roup s a r e b e t te r a b le t oo u t run p r eda to r s . Y oung s t a l l ion s , s h ow ing n oa l t ru i sm f o rc o l league s , f l ee d ange rr a th e rt h an f i gh t . A nc ien t h un t e r s p robab ly f o cu sed o n h a r em g roup s w h ich w ou ld h ave b e en e a s ie r t o l o ca te d ue t o t h e i r p r ed ic tab le b ehav io r , s h o r te r m ovemen t s , a nd s ma l le r h ome r a nge ( An thony 1 99 1) . I n a mbu sh f a sh ion , h un te r s c ou ld l o ca te h o r se t r a i l s a nd w a i t f o r t h e d om inan t m a r e t o a pp roa ch f o l lowed b yo t he rm embe r s . T he l e ad m a re a nd h e rf o a lw ou ld b et h e f i r s t t wo a n im a l sw ounded . A la rmed b y s c ream s a tt h e f r on t o ft h e h e rd , t h e s t a l l ion w ou ld r u sh t ot h e d e f en se a nd w ou ld b e d i spa tched . M i l l ing m a re s a nd f o a l s c ou ld b e w ounded a tt h i s p o in t . I fa n a t tack w a s l a unched f r om t h e r e a r , as u bo rd ina te m a re a nd h e ro f sp r ing w ou ld b ew ounded f i r s tb ef o re t h es t a l l ion r u shed a t tacke r s . H o r se so fb o th h a r em a nd b ache lo r h e rd s c ou ld a l so b e a mbu shed a t w a te rho le s w he r e t h ey r e gu la r ly d r ank . I f m anmade b l ind s w e r e n o t c on s t ruc ted , a mbu she s w ou ld n ece s sa r i ly r e qu i r e a dequa te v ege ta t ion o rt o pog raph ic r e l ief t oc oncea l h un te r s . I fh un te r s d r ove t h e h e rd i n to a c u l d e s a c , h un te r s w e r e r e qu i r ed t o a t tack a nd k i l l t h e s t a l l ion b ef o re m a r e s a nd c o l t s w e re d i spa tched . I f a h a rem g roup w a s a ta cked , o ne c an h ypo the s ize t h a tb one sa nd t e e th o fi mma tu r ec o l t sa nd m a tu re m a re sa nd s t a l l ion w ou ld b er e p r e sen ted a tt h ep roce s s ing s i t e . H o r se b one sf u se r a p id ly a nd a t ak nown r a te ; t o o th e r up t ion

J u ly

S umme r C oa tC ond i t ion

Aug .

J une

F ema le C ond i t ion Ma le C ond i t ion S ept .

May

Apr .

Oct .

Nov .

Mar .

—V i c

C

D ec .

4 /V .

F eb .

` 9) J an .

F i g . 4 . 2 . S ea sona l a t t r ibu tes o fh o r se s u sef u l i nr e cons tr uc t ing P a leo l i th ic f ood / techno logy m anagemen t s y s tem s .

4 9

s e quen ce s a r e a l so w e l l k nown . T he ref o r e , i t s h ou ld b e p o s s ib le t od e te rm ine i fh o r se e l emen t sr e cove red f r om ag i ven a r cha eo log ica l s i t e r e p r e s en t as l augh te red h a rem f am i ly . S t a l l ion s i n ab a che lo r h e rd a r e s t ronge r , m o re v i c iou s , a nd u np r ed ic tab le o n ad a i ly a nd a nnua lb a s i st h an t h ef ema le so fa h a rem . H ence , s u cce s sf u l ly s t a lk ing a nd k i l l ing a b a che lo r g roup w ou ld h ave b e en c on s ide rab ly m o re r i sky f o r p r eh i s to r ic h un te r s . Ma le s c ou ld b e p i cked o f s i ng ly i na n a mbu sh . O n ly o ne o rt wo i n d iv idua l s c ou ld p robab ly b ek i l led i nt h i s w ay a s o t he r m a le s w ou ld f l ee r a the r t h an f i gh t . T he m o s t e f ec t ive w ay t o d i spa tch a b a che lo r h e rd w ou ld b e a d r ive o rc o r ra l l ing t h em i n ac u l d e s a c . B ecau se a l m embe r s o f ab a che lo rg roup a r e a du l t m a le s o l de rt h an t wo y ea r s , t h e m o s t r e cogn izab le f e a tu r e o f ab ache lo r k i l l i st h e p r e sence o f n ea r ly o rf u l ly m a tu re b one s a nd t e e th . B ecau se o n ly s t a l l ion s ( b u tn o ta ls t a l l ion s )h av ec an ine s , ar e la t ive ly l a rge n umbe ro f t h e se t e e th a t as i te w ou ld i n f e rs l augh te ro f ab a che lo rg roup . H o r se s e xh ib i t l i t t le s e xua l d imo rph i sm . Ma le s c anno t b e d i s t ingu ished f r om f ema le s b y e i the r m ea su remen t s o fb one s o r t e e th . T he r ef o re , ac omb ina t ion o f a ge s t ruc tu re a nd p r e sen ce /ab sen ce o f s e xua l ly d imo rph ic c an ine s , i st h e b e s t m e thod i nd e te rm in ing t h ed emog raph ic g roup h un ted .

S ma l l h e rd s c ou ld a l so b e d i spa tched b y d r iv ing a nd /o r c o r ra l l ing . P r e requ i s i te s f o r s u ch s t ra teg ie s w ou ld i n c lude a dequa te l o ca l t opog raphy i n c lud ing r a v ine s a nd d une s . Mo s t p roduc t ive h un t ing w ou ld h ave o ccu r r ed d u r ing l a te s umme r / ea r ly f a l l a s h o r s e s m ig ra ted t owa rd w in te r ing g round s i n s h e l te red a r ea s . D u r ing m ig ra t ion s , h o r se s a gg r ega te i n to l a rge r b and s n umbe r ing i nt h e h und r ed s o r t h ou sand s . S uch l a rge n umbe r s c ou ld b e s t b e p rocu red b y d r ive s . A ga in , t opog raphy w ou ld p l ay ak ey r o le . H o r s e sm ay h ave b een s im i la r ly h un ted d u r ing s p r ing m ig r a t ion s t o o pen s t eppe a r ea s , b u t b o th m a re s a nd s t a l l ion s w ou ld b e i nt h e i r p oo r e s t c ond i t ion d u r ing t h i s s e a son . H a rem h e rd s , p r ed ic ta b le , a l t ru i s t ic , a nd s l owed b y y oung s te r s , w e r e t h e m o s te a s i ly h un ted . T h i s d emog raph ic g roup w ou ld h ave b een a c t iv e ly s ough t b y h un te r s . B ache lo r g roup s , f a s te r , u np r ed ic tab le , a nd m o re h o s t i le , w ou ld n o th ave b een a c t iv e ly s ough t ,b u tw ou ld h ave b een a t ta cked i fe n coun te red . A s d e sc r ibed i n C hap te r Two , d u r ing t h e L a s t G la c ia l Max imum , c o ld a nd i n c rea s ing a r id i ty i mpa c ted p l an t c ommun i t ie s a nd o ve ra l l p l an t b i oma s s . W i ld h o r se s p r obab ly a dap ted b y i n c rea s ing t h e s i ze s o ft h e i r h ome r a nge s . O n a g i ven d ay , h un te r s w ou ld h ave h ad l e s s c han ce o f e n coun te r ing ag i ven h e rd .

D u r ing f a l l a nd w in te r , h o r se s move t o p ro tec ted v a l ley s w h e re s h e l te r a nd b row se i sm o r e p l en t if u l . B ecau s e t h ey d o n o t d e f end t e r r i to r ie s , m any s ma l l h e rd s o f h o r se s c ome t oge the r o n t h e se m ig ra t ion r ou te s . T hou sand s o f s umme r f a t tened h o r se s , a gg r ega t ing d u r ing l a te s umme r /ea r ly f a l l m ig ra t ion s , c ou ld h ave b e en d r iven e n m a s s i fa pp rop r ia te t opog raph ic r e l ief w a s n e a rby . We l l e xecu ted s e a sona l d r ive s w ou ld h av e p rov ided l a rg e a moun t s o f m ea t a nd e x celen t h i de s . I nc onf in ed c ond i t ion s , b rough t o n b y w in te r s ho r tage s o f f ood , s t a l l ion s r e lax v i g i lance o f t h e i r h a rem s ( An thony 1 99 1) . A s ar e su l t , s e pa ra te f am i ly g roup s a r e f o und i nv e ry c l o se p rox im i ty t oe a ch o t h e r a nd m ay h ave b een m o re e a s i ly e ncoun te red d u r ing c o lde rm on th s .

C ap ra i b ex : i b ex L iv ing i n r ugged , m oun ta inou s t e r r a in , i b ex w e r e c ompa ra t ive ly d if i cu l t a n im a l s t o h un t a nd k i l l . P r ef e r r ing h i gh , w ind swep t m eadow s d u r ing w in te r , i b ex d e s cend t o l owe r p a s tu r e s a s s now s m e l t d u r ing e a r ly s p r ing ( F ig . 4 . 3 ) . I b ex a r e i nt h e i r p oo re s t c ond i t ion d u r ing t h i s s e a son a nd a r e m o s t v u lne rab le t o p r eda t ion b y h un te r s . D u r ing s p r ing a nd s umme r , i b ex m ove u pwa rd a sn ew gaz ing a r ea sa r ef r eed b y m e l t ing s n ow . B y a u tumn t h ey a r e a ga in a th i gh a l t i tude s a nd i np r ime c ond i t ion .

L ev ine ( 1 983 ) , u s ing m ea su remen t sa nd e r up t ion s e quence so f h o r se t e e th f r om S o lu tM , p r opo sed t h a t h un te r s d r ove a nd k i led l a rge n umbe r s o f h o r se s d u r ing e ach k i l l ing e p i sode . O l sen ( 1 989 ) , a na lyz ing b one s a nd t e e th , s ugge s ted t h e S o lu t rean h o r se s w e re r e pea ted ly d i ve r ted i n to a c u l d e s a c a nd k i l led i nl a te s umm e r a s t h ey m ig ra ted i nl a rge b and s . E qu id s a tS o lu t r é p rov ide e v idence t h a t w el o rg an ized a nd e xecu ted h un t s o fl a rge b od ied u ngu la te s r e pea ted ly o ccu r red t h roughou t t h e U ppe r P a leo l i th ic ( O l sen 1 989 ) . S t ran ska

F o r m o s t o ft h e y ea r , i b ex l i v e i nt wo b a s ic g roup s : f ema le s w i th t h e i r y oung a nd m a le h e rd s . T he s e g roup s c omb in e d u r ing t h er u ti nD e cembe r Janua ry . Ma le h e rd sg ene ra l ly f e ed i nh i ghe r , m o r e r ugged t e r ra in , a nd f ema le s w i th k i d s p r e f e r l owe r , l e s sp r e c ip i tou sa r ea s . T he e l u s ive i b ex m ay h ave b een h un ted i n av a r ie ty o fw ay s : 1 ) ambu sh e s a t s a l t l i ck s , 2 ) ca ref u l ly p l ann ed d r iv e s w he r e h un te r s d r ove a n ima l s u p s lope t owa rd a wa i t ing h un te r s , a nd 3 ) s ta lk ing f r om u p s lope ( S t rau s 1 987 ) . S pec ia l ized i b ex h un t ing s t a t ion sw e re d i scove red i nh i gh , r ugged t e r ra in o ft h e F r en ch P y r ene e s a nd t h e C an tab r ian C o rd i l le ra ( S t rau s 1 987 ) . S t rau s ( 1 987 ) d ocumen ted b ody p a r t f r equenc ie s f o r i b ex a t s i t e s l o ca ted d if e ren t d i s tance s f r om k i l l s . H e c on c luded t h a t b ody p a r t so fm a rg ina l u t i l i ty -head s a nd f e e t -we r e d i sca rded a th un t ing s t a t ion s c l o se t ok i l l s . H ighe ru t i l i ty p a r t s— tho r a ce s a nd l i mb s -we r et r an spo r ted t or e s iden t ia lb a se sl o ca ted w i th in at wo h ou r w a lk f r om t h e k i l l . S ea sona l i ty s t ud ie s , b a sed o n t oo th e r up t ion s e quence sa nd w ea r , s ugge s ted i b ex w e re k i l led d u r ing a ls e a son s .

S ka la I V ( Czech R epubl i c ) , af o cu s o f t h i s s t udy ( Chap te r F i ve ) , r e p re sen t s as i ng le k i l l ing e p i sode w he r e am in imum n umbe r o f 1 2 h o r se s w e re d r iven i n to ad une . Ambu she s o f o ne o rs e ve ra li n d iv idua l sw ou ld l e ave l i t t le o rn o s i gna tu re o n t h e l a nd s cape . U nde r s tandab ly , t h e se s ma l l e ven t s m ay n eve r b ed e tec t ed i nt h ea r chaeo log ica lr e co rd . I n s umma ry , d u r ing s umme r o r w in te r s ma l l h e rd s o f b a che lo r s o r h a rem s , a ve rag ing 5 12 i n d iv idua l s c ou ld h ave b een d i spa tched f r om a mbu sh a l ong w el t r aveled t r a i l s o r n ea r w a te rho le s . I fo ne o r s e ve ra l h o r se s w e r e k i l led , t h i s w ou ld h ave p rov ided b e twe en 5 00 k g t o s e ve ra l t h ou sand k g o fu sab le m ea t a nd f a t p e r k i l l ing e p i sode . F a t tened h o r se s , k i l led d u r ing s umme r w ou ld p rov ide m o re u sab le f o od s tuf f s t h an t h o s ek i led i nl a te w in te r . B ecau se s t a l l ion sf o rm h a rem s a nd d o n o t s uf e rt h e e xhau s t ive r i go r s o ft h e r u t , b o th m a le s a nd f ema le sw ou ld s l ow ly l o se c ond i t ion a sw in te rp r oceeded .

5 0

B e cau se f ema le g roup sg r a z e a tl owe r a l t i tude sw i th in s ma l le r c o r e a r ea s , a nd a r e s l owed b y k i d s , t h e se g r oup s w e r e m o s t e a s i ly d i spa tched u s ing a l h un t ing t e chn ique s . P ref e r red s e a son o fk i l lt oo b ta in t h em o s tm ea ta nd f a tw ou ld h ave b een s umme r a nd e a r ly f a l l w hen i b ex w e re m o s t i n a cce s s ib le .

J u ly

Aug .

F ema le &M a le C ond i t ion

J une

S ept .

May

Oct .

Apr .

Nov .

Mar .

D ec .

F eb . J an .

F ig . 4 . 3 . S ea sona l a t tr ibu te so fi b ex u sef u l i nr e cons t ruc t ing P a leo l i th ic f ood / techno logy m anagemen t s y s tems .

5 1

F u rbea re r s

a s soc ia ted t h e se m a rk s w i th m ea t r emova l , a s t ra tegy i n d ica t ive o fs e a sona l f o od s t re s s a nd /o r r i tua l c on sump t ion . T he p r ef e r r ed s e a son f o r q ua l i ty h i de , m a tu r e t e e th , a nd h i ghe s t f a t c on ten t w ou ld h ave b een l a te f a l l /ea r ly w in te r . H un te r /ga th e r e r sd u r ing t h eL a s t G la c ia l Max imum i nC en t ra l E u rope p ro ba b ly t ook a dv an tage o f a l p roduc t s o f t h e i r c a rn ivo re , a sw e l la sh e rb ivo re , q ua r ry .

F oxe s p r e sen t ly , a nd p r e sumab ly i n p r eh i s to ry , a r e h i gh ly v a lued f o r t h e i r f u r . W i th t h e o n se t o f c o ld w ea the r i n O c tobe r , t h ew in te rp e lage d eve lop sa nd i ss h ed i nA p r i l ( F ig . 4 . 4 ) . I ti sl i ke ly t h a t f u l ly a du l t f oxe s w e re h un ted d u r ing l a te a u tumn /w in te rw hen t h e i rc oa tw a sm o s tf u l ly d eve loped . F ox b one sr e cove red f r om E u ropean a r chaeo log ica l s i t e sa r er a re ly r e p re sen ted b y e p iphy se s , e p iphy sea l l i ne s o r i mma tu r e t e e th ( Pop l in 1 968 , P i dop l ichko 1 976 , S oi f e r 1 985 , 1 990 , We s t 1 990 ) . D o lu ld ianov ( 1 982 ) n o ted t h a t a s e a r ly a s t h e Mou s te r ian i nt h e R u s s ian P l a in , f u r b ea r ing a n ima l s w ou ld h ave b een h un ted i nw in te r , a nd S oi f e r ( 1 985 ) s t a ted t h a t a l r ema in s o f f o x f ound i n a l U ppe r P a leo l i th ic s i t e s i n t h e R u s s ian P l a in w e re f u ly a du l t , i n d ica t ing t h ey w e re h un ted i n w in te r .

A l though P i dop l ichko ( 1 976 ) s ugge s ted t h a tf oxe sc ou ld h ave b een s p ea r ed a st h ey w e re a t t rac ted t oh ou seho ld g a rbage a t c amp s i te so nt h eC en t ra lR u s s ian P l a in , a c t iv e ly s t a lk ing t h e se s ma l l a n ima l so n ar e gu la rb a s i sm u s th av eb een c o s t ly i nt i me a nd e ne rgy r e la t ive t ot h e r e tu rn . F ox p r ocu remen t w a s m o s t e f i c ien t b y i n teg ra t ing i ti n to al a rge r f o od / f at /f u r e x t ra c t ion s t r a tegy . S n a re so rt r ap sc ou ld b es e ta nd p e r iod ica l ly c hecked a s h un te r s r e tu rned f r om a n e xped i t ion f o r o the r r e sou r ce s . C r ea ted o fs t one sa nd /o rw ood , d eadf a l l sc ou ld k i lt h ea n ima l a nd p ro tec t t h e c a rca s s f r om s c avenge r s u n t i l h un te r s r e tu rned . T he se l i gh twe igh t a n ima l s w e re t r an spo r ted a s w ho le b ody u n i t s t oc amp s i te s w he re t h ey w e re s u b sequen t ly s k inned , b u t che r ed ( ? ) , a nd t h e i rt e e th e x t r ac ted ( Pop l in 1 968 ; S oi f e r 1 985 , 1 990 ; We s t 1 990 ) .

D u r ing t h e w in te r , f o xe s m ay h ave b een m o r e a cce s s ib le a nd p r ed ic tab le . F oxe sf o l low s p e c if i cm ig r a t ion r ou te sa l ong r i ve r v a l ley s a st h ey s e ek g am e ( Nowak a nd P a rad i so 1 983 , S of f e r 1 985 ) . F ox p opu la t ion d en s i t ie sv a ry s e a sona l ly b e ing g r ea te s t d u r ing b i r th ing s e a son a nd o n f a l l m ig ra t ion r ou te s ( Sa f e r 1 985 ) . H oweve r , f o xe s a r e d if i cu l t t ot r ap i ns p r ing a st h ey a r ep r eoccup ied s e ek ing m a te sa nd p ay l i t t le a t ten t ion t of ood ( Pede r son 1 966 ) . A dd i t iona l ly , s p r ing m ou l t ing w ou ld d e t rac t f r om t h e v a lue o ff u r s . F o r t h e s a ke o fp r a c t ica l i ty , t h e f ox , w ou ld h ave b een m o s te a s i ly c ap tu red b ys n a re so rd eadf a l l si n w in te r m on th s w h en f r e sh s now s w ou ld c oncea l h uman s c en t a nd w hen k ey t r a i l s c ou ld b e e a s i ly d e tec ted b y h un te r s . W in te r p e l t p rocu remen t s t ra teg ie s m ay h ave a l so b een a dvan tageou s f o r t h e e conom ic b enef i t s o f t a nn ing . S k in s a cqu i red i nw a rm s e a son s m u s t b e n ece s sa r i ly t a nned q u ick ly a sh a i r t e nd s t o f a l l f r om r a p id ly d e te r io ra t ing c a r ca s se s . O n t h e o t he r h and , p e l t s w i l l n o t a sr e ad i ly l o se t h e i r h a i r i ft h e f o x w e r e k i l led i n a u tumn /w in te r w hen t h e b ody c oo l s r e la t ive ly r a p id ly ( D r . L a r ry Ma r t in , p e r s . c omm ) .

A r c t ic f ox p opu la t ion s f l uc tua te w i th t h e t h ree t o f i ve y ea r l emm ing c r a she s ( Bee a nd H a l l 1 9 56 , P ede r son 1 966 , C he semo re 1 9 75 ) . S uppl i e s o ff ox a s af u r s ou r ce , m u s t h ave p e r iod ica l ly w axed a nd w aned i n a ny g i v en a r ea . C on sequen t ly , ac amp s i te m ay h av e b een o ccup ied d u r ing f a l l /w in te r a nd b ea rn o e v idence o ff o xp r ocu remen t a c t iv i t ie s i ff o xe s w e r e i ns h o r t s upp ly . O n t h e o t h e r h and , s i t e s l i ke W i l lendo r fL eve l N ine i nL owe r A u s t r ia ( Then iu s 1 9 56 -59 ) a nd E l i s ivee ch i a nd Mez in o n t h e C en t ra l R u s s ian P l a in ( Sof f e r 1 985 ) , w he r e f o x b one s a bound , s ugge s t t h e se s i t e s w e re p e r iod ic a l ly i n hab i ted a l ong m ig ra t ion r ou te s w hen f o xe s a nd t h e i rl emm ing p r ey w e r ea tp opu la t ion p eak s .

D u r ing l a te w in te r /ea r ly s p r ing i b ex w e re m o s t a c ce s s ib le i n l owe r e l eva t ion s , b u t w e re i n p oo re s t c ond i t ion . A r chaeo log ica l e v iden ce f r om S ou thwe s t E u rope s ugge s t s t h a tn ea r ly a lb one so fi b ex w ou ld b ef o und w i th in at wo h ou r w a lk o ft h ek i l l .

U n l ike f o xe s , f u r bea r ing h a re s a r e n oc tu rna l . H a re sb ea r d a rk f u r w h ich c amof l auge s t h em w i th s u r round ing s t one s d u r ing s umme r , W i th o n se t o f c o ld w ea the r t h e i r f u r t u rn s w h i te , a low ing t h em t oh i de i ns n ow . A s ar e su l t , h a re sw ou ld h ave b een d if i cu l t t o h un t w i th h and h e ld w eapon s . I n a n a r c t ic e xped i t ion , S t ef an s son ( 1 9 13 , 2 13 ) d e s c r ibed t h e r a r i ty o f h a re s : " The se a n ima l sa r er e a l ly n o t s ov e r yr a re o nt h eb a r ren g round , a s o n e m ay s e e b y t h e i r t r a ce s l e f t o n t h e s n ow i n w in te r , b u t i nm y e n t i r e A rc t ic e xpe r ien ce Ih ave s e en o n ly f o u r o r f i ve , a n d h ave n eve r s ho t as i ng l e o ne ." H a r e s w e re m o s t ef i c ien t l y o b ta ined b y t r app ing t h em l i ke f oxe s , o r i n s e a sona l d r iv e s . B ea r ing l i t le f a t a nd m e a t ( Spe th 1 983 , J och im 1 976 ) , h a r e s w ou ld h ave p rov ided s ma l l , w eak , b u t l u xu r iou s p e l t s u s e f u l i n l i n ing o r t r imm ing g a rm en t s ( P ede r son 1 966 , Wa lke r 1 968 ) . Af te r p e l t s w e re r emoved , h a re m ea t c ou ld a l so b e u sed a s b a i t i nf o x a nd w o lf t r ap s . H igh f r equen c ie s o f a r t icu la ted b one s o f h a re p aw s a t Mezh i r ich s ugg e s t t h a t t h e se f u zzy a ppendage s w e r e u s ed a s b r u she s i np r eh i s to ry ( P idop l ichko 1 976 ) , S pe th ( 1 983 ) a nd J och im ( 1 976 ) a g r ee t h a t h a r e s p o s se s s l i t le b ody f a t a nd r e ly ing o n t h em a s ac h ief s ou rce o fp r o te in w ou ld l e ad t o s t a rva t ion . Is u gge s t t h a t l e an h a re s w ou ld h ave b een e a ten i f h un te r s e x t rac t ed a dequa te f a t s upp l ie s f r om o t he r p r ey s p e c ie s . N e i th e r f o x n o r h a re p rov ided a dequa te k ca l . t o m a in ta in a b and o f h un te r g a the re r s , b u t t h e i r m ea t w a s c on sumed r a th e r t h an d i sca rded . T h e s e f u rbea r e r s w e re u ndoub ted ly m o s t v a lued f o rt h e i r or namen ta l p e l t s a nd t e e th .

Sma l l f u rbe a r ing a n ima l s a r e u sua l ly c on s ide r ed " s ta rva t ion " f ood s b ecau se t h ey s u ppo sed ly b ea r l i t t le f a t ( Spe th 1 983 , J och im 1 976 ) . P re s t rud a nd N i l son ( 1 992 ) d ocumen ted t h a t f o xe s e a t a sm uch a sp o s s ib le d u r ing s umme r m on th s a nd a r e f a t te s t , b o th v i sce r a l ly a nd s u bcu taneou s ly , i nN ovembe rw hen f a t c on s t i tu te s o ve r 2 0% o f t h e i r s k inned c a r ca s s w e igh t . B ody w e igh ta nd f a tc on ten td on o td im in i sh u n t i le a r ly s p r ing . F rom Ma rch t o May , b ody f a t d ec rea se s w i th t h e r i go r s o f m a t ing , g e s ta t ion , l a c ta t ion a nd g ene ra lc a re o fy oung , w i th f a t a ti t sl owe s t ( 6% ) i nt h e s umme r . P re s t r ud a nd N i l s sen ( 1 992 ) s howed t h a t s ubcu taneou s f a t o ve rt h e r ump a nd b a ck i sf i r s t d epo s i ted a nd l a s t t ob e d ep le ted i na r c t ic f o xe s . I nh ea l th ie r i n d iv idua l s , f a t i sa l so d epo s i ted o n f o re - a nd h i nd l imb s , f l ank sa nd t h eb e l ly . P o s s ib ly , t h ef o xw a sp rocu red i nf a l l a nd e a r ly w in te r m on th s f o rt h eb enef i t o fi t sf a t a nd m ea t a sw el a sf o ri t sp l u sh h i de a nd d eco ra t ive t e e th . D if e r en t ia lr e cove ry o fb one s o ff o requa r te r s a nd h i ndqua r te r s o fa r c t ic f o x f r om K rakow S padz i s ta s ugge s t s h aun che s o ff o xe s w e re u s ed f o r m ea t ( We s t 1 990 ) . S oi f e r ( 1 990 ) d ocumen ted b u tche r ing a nd f i le t ing m a rk s o n f o x b one s f r om D o ln i V e s ton ice . S he 5 2

J u ly Aug .

C oa tC ond i t ion

J une

F ema le & Ma le C ond i t ion S ept .

May

Apr .

Oct .

Nov .

Mar .

B e s tC oa t D ec .

F eb .

J an .

F i g . 4 . 4 . S easona l a t tr ibu te so fa r c t ic f ox u sef u l i nr e cons truc t ing P a leo l i th ic f ood / techno logy m anagemen t s y s tems .

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R e indee r w e re p robab ly n o t v a lued f o r t h e i r h i de d u r ing t h e s p r ing . G ua rd h a i r s w e re b roken a nd a n ima ls w e re s h edd ing t h e i r b r i t t le c oa t s . Wa r b le d amage w a s m o s t s ub s tan t ia l a nd h i de s w e re r i dd led w i th l a rge h o le s . T he b l ood e ngo rged a n t le r s i nv e lv e t o fb u l l s c ou ld b e e a ten , a nd t h e h a rdened a n t le r so fc ow sf a sh ioned i n to t oo l s .

L ong b one s m ay h ave b een f a sh ioned i n to h ou seho ld t oo l s i n c lud ing n eed le sa nd a w l s . T h e h a re , l i ke t h ef o x , w a s p robab ly o n ly s e a sona l ly a va i lab le t o h un te r s o f t h e C en t r a l E u ropean B a s in . Du r ing w in te r , h a r e sm ove t oh i gh e l eva t ion sw he r et h ey l i ve i nt r oop s o f1 0 2 0 , f e ed ing o nv ege ta ted s l ope ss wep tf r ee o fs now ( F ig . 4 . 5 ) . D u r ing s p r ing , t r oop s b r e ak u p a s h a re s m ove t o l owe r e l eva t ion s t o b r eed a nd r a i se t h e i r y oung . T hey r ema in i n l owe r e l eva t ion s d u r ing s umme r a nd b y a u tumn , a r e i np r ime c ond i t ion a nd p o s se s sl o ng , w h i te s of tf u r . L a te s umme r /ea r ly a u tumn w a s t h e o p t ima l s e a son t o t r ap h a re s b ef o r e t h ey m ig ra ted t o h i gh e l eva t ion s . S ea sona l f o r ay s t o h i ghe r e l eva t ion st o s n a re o r d r ive h a r e s i nw in te r p e lage m ay h ave o ccu r r ed , b u t m ay n o t h ave b een c o s t e f ec t ive u n le s s i n teg ra ted w i th al a rge rr e indee r / ibex p rocu r emen ts t ra tegy .

H o r se sw e r ea l so a va i lab le t oh un te r s a tt h i s s i te a si n d iv idua l h e rd s a gg r ega ted a nd m ig ra ted f r om s h e l te red v a l ley s t o u p land p l a te au s . L a rge g roup s o fh o r se s c ou ld b e d i spa tched b y w el o rg an ized d r ive s o r t aken i n a mbu sh i fa dequa te t o pog raphy a lowed . G rubg raben i sl o c a ted a d ja cen t t o a r a v ine w h ich c ou ld s e rv ea s at r ap f o rh o r se s . Al a rg e n umbe r o fw e l l o rga n ized h un te r s c ou ld d r iv e h o r se s i n to t h i s c u l d e s a cw he re t h ey c ou ld b es p ea red . I nt h ep a s t , G rubg ra ben w a s l o ca ted a d ja cen t t o a c t iv e s p r ing s a t rac t ive t o h o r se s . A l though e t hnog raph ic e v idence s how sh o r se sc an b et a ken b y d i gg ing p i t s n ea r w a te r ing h o le s ( Moh r 1 97 1) , Is ugge s t h o r s e s c ou ld b ea mbu sh ed f r om m a nmade o rn a tu ra l c ove r a s t h ey c ame t od r ink . I na mbu sh , h un te r sc ou ld e xpec tt ot a ke a t l e a s t o ne o r s e ve ra l a n ima l si n as i ng le k i l l ing e p i sode . H o r se s w e re a t t h e i r p oo re s t c ond i t ion i n s p r ing ; b o th m a re s a nd s t a l l ion s h ad s l ow ly l o s t w e igh t o ve r w in te r . O ne p a r t icu la r d emog r aph ic g roup w ou ld n o t b e f a vo r ed f o r h un t ing a l though h a r em g r oup sw e r el e s sd ange rou sw hen a t t a cked .

S ea sona l S t ra teg ie s : P re c i se s c hedu l ing f o r t empo ra l ly d i spe r sed a nd l i m i ted r e sou r ce s a lowed E p ig rave t t ia n h un te r s t o m a in ta in a n e f i c ien t b a lance b e tween c o s t s a nd r e tu rn s d u r ing t h e L a s t G la c ia l Max imum . A n o rgan iza t iona l m ode l b a sed o n s c hedu l ing t or e ce iv et h eg r ea te s tf o od / techno log ica lr e tu rn a t t h el e a s tc o s ti sp r e sen ted f o rG rubg raben :

I b ex , a l p ine s p e c ie s , n eve r l i ved i nt h e i mmed ia te v i c in i ty o f G rubg raben , b u tw e re l o ca ted i nt h eu p land s n or th o ft h e s i t e , t h e Wachau a nd A lp st ot h es ou thwe s t , a nd t h eC a rpa th ian st o t h e e a s t . D u r i ng t h e s p r ing , i b ex moved t ol owe s t e l eva t ion s t o t a ke a dvan tage o f g r a s s g reen ing f r om r e cen t ly m e l ted s now s , a nd a l though m o s ta c ce s s ib le t oh un te r s , t h ey p r ov ided t h e l owe s t m ea t / f a t y i e ld a tt h a t t ime o fy ea r . I fi b ex w e re h un ted f o r t h e i r l a te w in te r p e l t s a nd o rb one s , s p r ing w ou ld h ave b een a n i d ea l t i me t o p r ocu re p e l t s o fa n ima l s a tl owe r e l eva t ion s o ft h e Wachau . H a re s , mov ing t ol owe r e l eva t ion s i ns p r ing c ou ld b e i n teg ra ted i n to t h e l a rg e ri b ex p rocu r emen t s t ra tegy . B o th i b ex a nd h a re sc ou ld b em o s t p rof i tab ly h un ted i nd r ive s w he r e l a rge n umbe r o f a n im a l s c ou ld b e t a k en a t o nce . As i ng le o rs e ve ra l i b ex c ou ld a l so b et a ken i na mbu sh , b u t n oc tu rna l a nd c amo f lauged h a r e s w ou ld b e s i ng ly p rocu red t h rough t r ap sa nd /o rs na re s . S p r ing f o ray st or u gged t e r ra in i mp lemen ted s u ch a c t iv i t ie s , b u t m u s t h av e b een c on s ide red r e l a t iv e ly c o s t ly e xped i t ion s w h en l a rge r r e indee r a nd h o r se s w e r e a c ce s s ib le i n t h e i mmed ia te a r ea o f G rubg raben . When c ompa red t o r e indee r a nd h o r se s , w in te r t h inned i b ex p r ov ided l i t t le m ea t a nd w e r e v a lued f o r t h e i r h i de s , h or n s , a nd s t ra igh tm e tapod ia l s—ex ce l len tr aw m a te r ia l s f o rh un t ing w e apon s .

S p r ing : I n t h e v i c in i ty o f G rubg r aben , d u r ing e a r ly s p r ing , r e indee r m ig ra ted f r om w in te rf e ed ing g round s i ns h e l te red m a jo rr i ve r d r a inage so ft h eD anube . F olow ing a r ea so fl e a s tt opog raph ic r e l ie f , r e indee r a gg rega ted a s t h ey m oved t o h i ghe r , l e s s s h e l te red s umme rf e ed ing t e r r i to r ie s . S ome h e rd sm oved f r om t h e b road r e g ion o ft h e A u s t r i an P l a in a l ong t h e D anube , u p t h eK amp t owa rd t h e Mo rav ian H i l l s . O the r sm ay h ave m oved t o t h e f oo th i l l s o f t h e T a t ry a nd /o r t h e n o r thea s t A lp s . R e inde e r d i d n o t m ove a s as i ng le m a s s ive u n i t , b u t f i l te red t h rough p a s se s a nd a c ro s s r i ve r s i n s ma l l h e rd s . P r egnan t f ema le s t o ok t h e l e ad , s pu r red t o a r r ive a tc a lv ing g round s . Ma le s a nd b a r r en f ema le s f o l lowed , m ov ing a t as l owe rp ace . M ig ra t ion s p r obab ly o ccu r r ed o v e r s e ve ra l w eek s t o am on th a st h ea n im a l sf i l te red a l ong a l ong t h e se r ou te s . T he o p t ima l s t ra tegy f o r t a k ing l a rge n umbe r s o f r e indee r i n vo lved s t ra teg ica l ly p o s i t ion ing h un te r s a l ong p a th s o ft h e m ig ra t ing a n ima l s j u s t t o t h e n or th o r s ou thea s t o f G rubg raben . N a r row r i ve r c r o s s ing s a nd t o pog raph ic " bo t t lene ck s " c on cen t ra ted a nd s l owed m ig ra t ing a n ima l s . L a rge n umbe r so fa n ima l sc ou ld b ed i spa tched a st h ey c r o s sed f o rd s i nt h e K amp o rw e re d r iven i n to " bo t t lene ck s " n or th o f G rubg raben . R e inde e r c ou ld b e i n d iv idua l ly s t a lked a s t h ey p a s sed . D epend ing o nd en s i ty o fa n ima lh e rd s , al a rge q uan t i ty o f m ea t a nd s ome m a r row c ou ld b e a cqu i red , d r ied , a nd s t o red d u r ing t h i ss e a son .

A l though f ox e s p r obab ly l i ved i n t h e i mm ed ia te a r ea o f G rubg raben , b o th i nt h e v a l ley s a nd u p land p l a teau s , s p r ing w a s n o t a n o p t ima l t i me t o t r ap t h e se a n ima l s a s t h ey w e re s h edd ing t h e i rp l u sh w in te rc oa t sa nd w ou ld h ave l o s tm uch o f t h e i r b ody f a t . B e cau se f o xe s w e re p r eoc cup ied w i th m a t ing a nd b i r th ing , s p r ing w a s ap a r t icu la r ly d if i cu l t t ime t oh un t , t r ap , o rs n a r et h e se a n ima l s .

D u r ing t h e s p r ing , p r egnan t f ema le s a r e h eavy , p rov id ing m ea t , b u t l i t t le f a t a nd m a r row . P regnan t f ema le s w e r e t h e f i r s tk i l led b e cau se t h ey w e re t h ef i r s tt ob ei n te r cep ted d u r ing m ig ra t ion s . F a s t m ov ing f ema le s c ou ld t i de t h e h un t ing g roup o ve r u n t i l l a gga rd m a le s c ou ld b e e xp lo i ted . L a rge r m a le s , h av ing p a r t ia l ly r e bu i l tf a ts upp l ie so v e rw in te r , w e re t h ef o cu s o fh un t ing p a r t ie sd u r ing t h es p r ing m ig ra t ion s .

S umme r : D u r ing a nd i mmed ia te ly a f te r c a lv ing , r e indee r w e re h i gh ly c oncen t ra ted a tc a lv ing g round s i nt h e f o o th i l l s o f Mo rav ia , t h e T a t ry , a nd t h e A lp s . T h i s d i s tan ce p r obab ly c on s t i tu ted 5 4

J u ly Aug .

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D i spe r sed S ept .

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t i91' e s t Nov .

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c l e s

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C once n t ra t ed

B e s tC oa t D ec .

F eb . J an .

F ig . 4 . 5 . S ea sona l a t tr ibu te so fa r c t ic h a re u sef u l i nr e con s truc t ing P a leo l i th ic f ood / techno logy m anagemen t s y s tems .

5

m o re t h an a w eek 's w a lk f r om G rubg raben . Af ew a n ima l s p robab ly l i nge red a l ong m ig ra t ion r ou te s , b u t m o s t d ee rw e re p robab ly f a r e nough a way f r om G rubg raben t h a t h un t ing g roup s w e r e f o rced t o m ov e i n e i the r a n n o r thea s ter n o r s ou the rn d i re c t ion i ft h ey h oped t oe ncoun te rr e indee ri nt h e i r s umme r r a nge . H un te r s m ay h av e o p ted t o k i l l a s m any r e indee r a s p o s s ib le d u r ing t h e i r s p r ing m ig ra t ion s n ea r t h e s i t e . D ry ing a nd c ach ing m ea t , t h ey c ou ld l i ve o ft h e s t o r e s u n t i lr e indee rr e tu rned d u r ing t h ef a l l . H oweve r , t h i si s ar i sky a nd i n ef f i c ien t s t ra tegy . I fh un te r s c ho se t o r ema in a t t h e G rubg r aben s i t e d u r ing s umme r t h ey m i s sed t h e o ppo r tun i ty t o h un t a n ima l s w i th n ew ly h a i red s umme r c oa t s w i th n o w a rb le d amage . A dd i t iona l ly , t h ey w ou ld m i s s o ppor tun i t ie s t o k i l l a n ima l s f a t ten ing d u r ing t h e s umme r . C ow s s h ed t h e i r a n t le r s i mmed ia te ly a f te r p a r tu r i t ion a nd t h e s e a l ong w i th m a s sa c red a n t le r sf r om b u l l sp rov ided r aw m a te r ia l sf o rt oo l s . I mmed ia te ly a f te r b i r th ing , c ow s a r e i n t h e i r p oo re s t c ond i t ion ,b u tb ul s , c on t inu ing t of a t ten f r om t h ep r ev iou sf a l l r u t , w ou ld p rov ide l a rge a moun t s o fm ea t , f a t , a nd m a r row . I b ex a nd h a r e s a l though f o und i nm o re r u gged t e r ra in t h an s umm e r u p land t e r r i to r ie s o f r e indee r , c ou ld b e h un ted i n f o r ay sf r om b a se c amp sl o ca ted i nr e indee rs umme rt e r r i to r ie s . A ss now sm e l ted a nd f o rage g r eened i nh i ghe re l eva t ion s , i b ex m oved h i ghe r b ecom ing i n c rea s ing ly f a t te r , b u t m o r e i n a cce s s ib le t oh un te r si nt h ef o o th i l l s . H un te r sw e r ef o r ced t o m ake l o nge r e xcu r s ion s t o h i ghe r e l eva t ion s t o e xp lo i t t h e se r e sou r ce s . S uch e ndeavo r sm ay h ave p roved i mpo s s ib le i ft h a t s umme rc amp w a sl o ca ted a tG rubg raben . S ma l l a nd w ide ly d i spe r sed h o r s eh e rd sf o raged i nt h eu p land s s u r round ing G rubg raben d u r ing s umme r . L im i ted a nd w ide ly d i s t r ibu ted p l an t s d u r ing t h e LGM i n c rea sed s umme r t e r r i to r ie so fb o th h a rem sa nd b a che lo rg roup s . B o th c ou ld b e e ncoun te red a nd d i spa tched i na mbu she s o r s ma l l d r ive s , b u t e ncoun te r sm ay h ave b een r a r e . Ab a lance o fc o s t a nd r e tu rn s i mp roved a st h e s e a son p rog r e s sed a nd a n ima l s f a t tened . O ne o r s e ve ra l h o r se s p rov ided l a rg e q uan t i t ie s o f m ea t a nd f a t . Y e t , b e cau se o ft h e i ru se o fl a rge t e r r i to r ie s , h o r se s w e re t o o u np r ed ic tab le f o ra n t ic ipa ted w indf a l lr e tu rn so f ak i l l .

D u r ing e a r ly f a l l , h i de s w i th h eav ie r f u r , p rov ided e x ce l len t m a te r ia l s f o rw a rm c l o th ing . H a rd a n t le r s o fm a s sa c r ed c ow s a nd b u l l s m ade u sef u l w eapon s a nd h ou seho ld t oo l s . I nl a te f a l l , a f te rt h er u t , s h ed a n t le r so fb u l l sc ou ld b ec o l lec ted . L a rge n umbe r so ff a t tened , m ig ra t ing r e indee r c ou ld b et a ken o v e r as pan o f t h ree o r f ou r w eek s d u r ing a u tumn . S i te a c t iv i t ie s a tG rubg raben w ou ld h ave i n c luded p r epa r ing h i de s f o r t a nn ing , b u tche r ing m ea t f o r d r y ing / smok ing , b r eak ing b one sf o rm a r row , a nd c r u sh ing b one sf o rg r ea se e x t r a c t ion . H o r se s w e r e p robab ly a l so o n t h e m ov e a s t h ey m ig r a ted t owa rd s h e l te red r i ve r v a l ley s . H o r se m e a t a nd h i de w a s o f p r em ium q ua l i ty d u r ing t h e f a l l a nd s e v e r a l a n ima l s p r ov ided m ea t e qua l ing a tl e a s t e i gh t t o t e n r e ind ee r . L a rge h i de s o f h o r se s c ou ld b e t a nned a nd u sed a s w in te r t e n t s o r f l oo r c ove r ing s . I b ex w e r e f a ro u t o fr e ach i nr u gged a l p ine t e r ra in a nd h a re w e re m ig ra t ing f r om l owe r t o h i ghe r e l eva t ion s . F oxe s w e re i np r ime c ond i t ion i nl a te r f a l l w hen t h e i r c oa t s w e re i nt h em o s tl u xu r iou sc ond i t ion a nd t h ey b o re ar e la t ive ly h i gh p e r cen tage o fy ea r ly f a t . A l l f oxe sw e re a du l t b ya u tumn a nd i n f u l l c oa t . I n expe r ienced y oung s te r s c ou ld b e e a s i ly e n t iced b yb a i ti n to t r ap sa nd s n a r e s . W in te r : A sr e indee rr e a ched s h e l te red a r ea si nt h eA u s t r ian P l a in , t h ey d i spe r sed , l o ok ing f o r w in te r f o rage . D u r ing w in te r , l a rge n umbe r s o fa gg r ega ted a n ima l s c ou ld n o t b e s imu l taneou s ly e ncoun te red . H un te r s a tG rubg raben m u s t h ave s t a lked a nd c ap tu red r e indee r a s t h ey w e re e ncoun te r ed . B y m idw in te r , a f t e r t h e r u t , b u l l s w e re i n e x t reme ly p oo r c ond i t ion , b u t w ou ld h ave p r ov ided t h e l a rge s t , h eav i ly f u r r ed h i de s . C ow s a nd y oung a n ima l s w e r e f a t d u r ing e a r ly f a l l a nd t h e i r b ody c ond i t ion s l ow ly d im in i shed t h roughou tw in te r . H eav i ly f u r r ed w in te r h i de s w e r e t a nned f o r b l anke t s , r o be s a nd t e n t s . F ema le s s t i l l r e ta ined t h e i r h a rdened a n t l e r s , b u t b u l l s w e re a c t iv e ly g row ing t h e i rr a ck s . B u l la n t le r s , i ne a r ly w in te r ,w e r e p r ef e r red f o rf o od , r a the rt h an t h e i rt e chno log ica lv a lue . H o r se s , f o rag ing i ns h e l te red v a l ley s , w e r e m o re a gg r ega ted t h an i n t h e p r ev iou s s umme r a nd m ay h ave b een m o re f r equen t ly e n coun te r ed a nd h un ted . D u r ing e a r ly a nd m idw in te r , b o th m a re s a nd s t a l l ion s p r ov ided a t r emendou s a moun to fm ea ti ft h ey c ou ld b et a ken , b u tt h er i sk sa s soc ia ted w i th h un t ing t h e se d ange rou s a n ima l s w e r e a lway s p r e s en t . I b ex a nd h a re sw e r e l i v ing i nd i s tan t , r ugged t e r r a in a nd w e re n o ta sa cce s s ib le t oh un te r sa tG rubg raben . H un te r sc on t inued t oh un t a nd /o rt r ap f o xe st h roughou tw in te rf o rf u r , t e e th , a nd f o od .

F ox e s i nl i gh t , s umme r c oa t s p r obab ly m oved t h rough t h e G rubg raben a r ea , a nd c ou ld b e e a s i ly t r apped a st h ey s ough t f ood . H oweve r , t h in f oxe s w ea r ing l i gh t s umme r c oa t s w e re n o t v a lued . I n expe r ienced , g row ing p up s , c ou ld b e e a s i ly t r apped , b u t b o re d e c iduou s c an ine s o f l i t t le v a lue f o r or namen t . A u tumn : A u tumn w a sp robab ly a no p t ima lt i me o fy ea ri nt h ev i c in i ty o f G rubg raben . R e indee r a nd h o r s e s , m ov ing f r om p l a teau s a nd f oo th i l l r e g ion s , p a s sed a l ong t h e D anube t o r e a ch s h e l te red a r ea s . L ike i ns p r ing , i n d iv idua l s o fr e indee r h e rd s c ou ld b e p i cked o f a s t h ey g r azed b e low i n t h e K amp V a l ley a nd m oved t h rough " bo t t lene ck s " o f D anube t r i bu ta r ie s . We l l e xecu ted d r ive s o fa n ima l s t h rough " bo t t lene ck s " a nd a c ro s s r i ve r s l i ke t h e K amp o r D anube w ou ld p rov ide aw indf a l l o f m ea t , f a t , m a r row , h i de s , a nd a n t le r s . A n ima l s p robab ly m ig ra ted b ef o r et h eO c tobe rr u t ; s u b s equen t ly , b o th m a le sa nd f ema le s w e r e i np r ime c ond i t ion f o r m ea t , f a t , a nd m a r row . B ul s w e ighed u p t o 2 0% m o r e t h an c ow s w e r e p r ef e r red i n e a r ly a u tumn . A f te rt h e r u t , i nN ovembe r , b u l l s l o s t n ea r ly a l f a tr e sou r ce s ;a s ar e su l t , c ow sw e r ep r ef e r red i nt h el a te f a l l .

5 6

Chap ter 5 Hor se Hun t ing ,P roce s s ing ,a nd T ran spor t T he h o r se w a s am a jo r f o od /h ide s ou rce f o r E p ig rave t t ian h un te r s . A tG rubg raben ,t e e th a nd b one so fh o r se sw e r ef o und i na l l e ve l s . Am ode l f o r E p ig rave t t ian s t ra teg ie s f o r h o r s e p rocu r emen t ,f i e ld b u tche ry ,t r an spo r t , a nd c amp p roce s s ing i s d eve loped i nt h i s c hap te r . T h i si sa c compl i shed b y c ompa r ing a ge s t ruc tu r e o f t e e th a nd a na tom ica l f r equenc ie s o f h o r s e b one s f r om c amps w i th t h o se a s soc ia ted w i th a c on tempo raneou s k i l l . A s e r ie s o f h ypo the se s c on ce rn ing c ul ing p r a c t ice s o f h e rd s , d i sa r t icu la t ion a nd t r an spo r t o f b one s , a nd m a r row e x t ra c t ion s t r a teg ie sa r et e s ted . B o th l a rg e a nd s ma l l t y pe s o f h o r se s c oex i s ted i n w e s ter n E u rope d u r ing t h e P l e i s tocene ( P ra t 1 976 , D e lpe ch 1 983 ) . T he se t wo h o r se s a r e f e a tu r ed i np ar ie ta l a r t . P rzewa l sk i t y pe a n ima l s w i th u p r igh t m ane s w e re p o r t rayed a t N iaux , L e P o r te l , L a s caux , a nd T ro i s F re re s a nd t h e s ma l le r p ony a t L a scaux ( Le ro i -Gou rhan 1 965 , 1 982 ; S anda r s 1 985 ) . V ö rö s ( 1982 ) c onf i rmed t h a t t wo t y pe s o f h o r se s w e re s ympa t r ic d u r ing t h eE p ig rave t t ia ni nH unga ry . H ed e s ig na ted t h e l a rge r P r zewa l sk i s i zed a n ima l a sT ype Ia nd as ma l le rp ony a sT ype I .B one so fb o th t y pe sw e r er e cove red f r om G rubg raben . P a leol i th i c h un te r s r e a l ized t h et r ade -of fb e tween e a sy k i l so f r e indee r sw h ich p e r a n ima lp rov ide r e la t ive ly l i t t le p r o te in a nd t h em o r ed ange rou sh o r se w h ich w a sh a rde rt ok i l lb u ty i e lded " w indf a l l " p r of i t s i nm ea t ( Sp ie s s 1 979 ) . B ecau se o fi t s l a rg e s i ze , t h e h o r se w a s e x ten s ive ly d i smembe red a tt h e k i l l s i t e b ef o r ei tc ou ld b et r an spo r ted t o ac amp . I na ni d ea l s i t ua t ion , h o r s e b one s r e cove red f r om ak i l l a nd r e s iden t ia l s i t e s h ou ld b ep ropo r t iona l i n ve r se s ( B inf o rd 1 98 1 : 2 18 ) . A l though b ody p a r t r e p r e sen ta t ion o fh o r se s a tc amp s i te s h a sb een d i scu s sed , f ew a r chaeo log i s t s h ave p ropo sed a r a t iona le f o r a n c ien t h un te r sw ho t r an spo r ted c e r ta in p a r t s o ft h e c a r ca s s f r om k i l l t o r e s iden t ia l b a se s , b u t o p ted t o l e ave o t he r p a r t s b eh ind . V ö rö s ( 1 982 : 5 0 ) , c a lcu la t ing a na tom ica l p a r t s o f e qu id s r e cove r ed f r om t h e E p ig rave t t ian o pen a i r s i t e o f S agva r , n o ted t h a to ve r8 1% o ft h er e cove red r ema in sr e p re sen ted t h e h ead r e g ion f o l lowed b y e l emen t s o f t h e l owe r , n onmea t b ea r ing r e g ion o f t h e l e g s . H e c on c luded t h a t h o r se s w e re p a r t i t ioned e l sewhe re a nd b one s o ft h e h ead a nd f e e t w e r e t r an spo r ted t o t h e c amp . H e d i d n o t e xp l a in h ow s u ch a na tom ica lf r equenc ie sm igh t r e f lec t c on sc iou s p roce s s ing a nd t r an spo r t s t ra teg ie s o f a nc ien t h un te r s . D obo s i a nd V ö rö s ( 1986 , 1 987 ) a l so r e cogn ized a h i gh f r equency o f h ead e l emen t s o f h o r se a t P i l i s szan to R ock she l te r s Ia nd H . A na lyz ing h o r se r ema in s f r om t h e Magda len ian s i t e o f G onne r sdo r f , P op l in ( 1 968 ) s aw a s omewha t d if e ren t a na tom ica l p a t te rn . H e n o ted a r e la t ive s c a r c i ty o f h ead e l emen t s a nd m ea ty e l emen t s o ft h e c a rca s s , b u t av e ry h i gh r a t io o f m e tapod ia l s a nd p ha lange s . P op l in c onc luded t h a t d if e ren t ia l p r e se rva t ion , b a sed o n b r eakage a nd b one d en s i ty v a lue s , m igh t a c coun t f o r t h e f r equen c ie s o fa na tom ica l p a r t s a tt h e s i t e . S p ie s s ( 1 979 ) , a na lyz ing h o r se f r om A b r i P a taud , d i d n o t a dd re s s t r an spo r t i s sue s . H o r se b one s w e r e h i gh ly f r agm en ted a nd w e r e i ng ene ra l ly p oo r c ond i t ion a tt h i s s i t e . O l sen ( 1 989 ) , s t udy ing a na tom ica l f r eqenc ie s o fb one s f r om S o lu tM , p r opo sed t h a t h o r se c a r ca s se s w e re f i le ted a nd t h e i r h eavy b one sl e f ta tt h ek i l lt oc u tt r an spo r t ing c o s t s . 5 7

E p ig rave t t ia nH o r se H un t ing ,P roce s s ing , a nd T ran spor t : A ge a nd a na tom ica l f r equenc ie s o fh o r se b on e s a r e d e sc r ibed f r om t h e f ou r c u l tu ra l l e ve l s a t G rubg raben , A u s t r ia . T h e se f r equenc ie s a r e c ompa red w i th t y pe s o f h o r se e l emen t s r e cove red f r om o t he r C en t ra l E u ropean r e s iden t ia l c amp s : S agva ra nd P i l i s szan to R ock she l te r s Ia nd I LH unga ry , t os e e i fp a t ter n s o f h o r se e l emen t t r an spo r t a r e c on s i s ten t a c ro s s s p a ce a nd o ve r t i me . T h e t y pe s o f h o r se b one s f o und a t r e s iden t ia l s i t e s a r e c ompa red w i th t h o se r e cove r ed f r om a c on tempo raneou s h o r se k i l l s i te : S t ran ska S ka la I V i nt h e C zech R epub l ic . T he f o l low ing h ypo the se sa r et e s ted : 1 .I n ah a r sh e nv i ronmen t , h un te r s s e le c t iv e ly c u l led p r im e a du l t e qu id s . T he se i n d iv idua l s p rov ided t h e m o s t m ea t , f a t , a nd m a r row t h a t o f se t t h e r i sk s o f h un t ing a nd k i l l ing d ange rou sq ua r ry . 2 .B ecau se h o r se sa r el a rge b od ied u ngu la te s , t h ec a rca s sw a s d i smembe r ed a t t h e k i l l s i t e t o f a c i l i ta te t r an spo r ta t ion o f s k in s ,b one s , a nd m ea tu n i t st or e s iden t ia lc amp s . 3 . I fb one s w e r e t r an spo r ted , t h ey s h ou ld r e p re sen t t h o s e c a r ca s s p a r t s t h a t p r om i sed t h e g rea te s t r e tu rn i nm ea t , f a t a nd /o r m a r row f o r t h e a moun t o f e n e rgy e xpended i n p roce s s ing a nd c a r ry ing . T he se e l emen t s s h ou ld i n c lude m ea ty p a r t so fu ppe rl i mb s ,v e r teb rae , r i b s , a nd p e lv e s . 4 .H un te r sd i dn o tt r an spo r tb one sa s soc ia ted w i th l owe rm ea t a nd m a r row y i e ld s . T he se e l emen t s s h ou ld i n c lude n onmea ty p a r t so fs ome l owe re x t rem i t ie sa nd h ead s . 5 . I n ad e te r io r a t ing e nv i ronmen t , h un te r s , s e ek ing f a t , e x ten s ive ly f r agmen ted h o r se b one st oe x t ra c tm a r row . G rubg raben : L eve l 1 A na tom ica lF requen c ie sa nd MN I : At o ta l o f 1 26 s k e le ta l f r agmen t s , r e p re sen t ing a m in imum n umbe r o ff ou r h o r se s w e r e r e cove r ed f r om A L 1 ( Tab le 5 . 1 , F i gu re s5 . 1 a nd 5 . 2 ) . Al a rge a nd s ma l le ra du l t , aj uv en i le , a nd o ne p ony w e r e i d en t if i ed . T he h o r se i sr e p re sen ted m o s t f r equen t ly b y b on e s o ft h e l owe r l e g / f oo t a nd h ead . B one s a s soc ia ted w i th m ea t b ea r ing r e g ion s o ft h e c a r ca s s a r e m uch r a re r . H ead s F ou r h o r se s a r e r e p re sen ted b y 1 2 i s o la ted t e e th a nd 2 a l veo lae f r agmen t s i nl e ve l 1a tG rubg raben . T h ree m and ib le s a nd o ne c r an ium r e p re s en ted t h e se f o u r a n ima l s . O ne o ft h e f ou rh o r se si sas ma l l , p ony l ike a n ima lw i th s ma l l c he ek t e e th p o s se s s ing a d i s t inc t ive p l icaba l l ine c u sp w h ich d if e r s s i gn if i can t ly f r om t h a to ft h eP rzewa l sk it y pe . T h i s s ma l le qu id w a s a l so f o und a t S agva r ( Vö rö s 1 982 ) . T h i s p ony i s r e p r e sen ted b y r i gh t m ax i l la ry M1 a nd M2 . T he M 1 i sm o r e w or n t h an t h e M2 w h ich i s e xpe c ted a s t h e M 1 e r up t s b e tween n i ne a nd t we lve m on th s a nd t h e M2 e r up t s a t2 4 m on th s . T h i si n d iv idua lr e p re sen t s ap r ime a du l ta tt h et i me o f d ea th .

T ab le 5 . 1 . N umbe ro fi d en t if ied s pe c imen sa nd m in imum n umbe ro fh o r se e l emen t sf r om t h ef o u rc u l tu ra l l e ve ls a t G rubg raben .

A na tom ica le l emen t

N ISP

AL I MNE

N ISP

AL2 MNE

N ISP

AL3 MNE

AL 4 N ISP

MNE

H ead : T ee th S ku l l Mand ib le

1 2 2

A x ia l : C e rv ica l T ho rac ic L umba r P e lv i s R ib

2 2

M ea ty F o r e leg : S capu la H ume ru s R ad iu s /U lna

1

Mea ty H ind leg : F emu r T ib ia L owe rL eg s : C a rpa l M e taca rpa l T a r sa l M e ta ta r sa l P ha lanx I P ha lanx I P ha lan xI l M e tapod ia l

1 6 1 3

3

2 1

-

1 1

3 4 2 8

3 5

1

4 5 8 3 5

3 3

6 1

2 1

3 2 6 5

1 3 1

1 5 1

3 1

1

1

1

1 4 2

1 2 1

1 4 6 2 0

2 2 2

1 8 1 8 2 3

4 2 2

3 2

2 2

1 8

1 3

7 1 8

2 2

1 0 4 1

2 3

3 4 5 4 3 3

2 2 4 3 3 2

2 1

2 1 1 2 1

6 5 I 1 1

1 1 1 2 1

2

1 3

4

5 8

1

1

1

1

1 2 I

2

1

6

F i g . 5 .1 . Numbe r o fi d en t if ied s pec imens o fh o rse f r om L eve l 1a tG r ubg raben . U nshaded a r eas r e pre sen t b one sa bsen t .

5 9

S KELETAL E LEMENTS

G RUBGRABEN

A L 1 -N ISP 1 2

T EETH S KULL « .0 MAND M O M 2 C ERV 0 1 -OR 0 L UM 0 P ELV » MR 2 R IB 2 SCAP 0 1 I UM «0 R /ULNA »I I I 1 F EM T IB IA » M i l 2 CARP 0 MC TARS 0 MT I I MM IM E P HAL 1 P HAL 2 P HAL 3 MTP

3

3

M I

4

5

4

3 3

0

5

1 0

1 5

NUMBER

S KELETAL E LEMENTS

GRUBGRABEN T EETH 0 S KULL » MI ME MAND C ERV 0 T HoR 1 0 L UM 0 P EL R IB V SCAP 0 H UM R /ULNA F EM T IB IA CARP 0 MC T ARS 0 MT P HAL 1 P HAL 2 P HAL 3 MTP

AL 1 - MNE

1

3

1

1 2 2

2 2 4 3

3 2 r -

1

2

3

4

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NUMBER

F ig . 5 . 2 . H is tog rams o fn umbe ro fi d en t if ied s pec imens ( N 1SP ) a nd m in imum n umbe ro fe l emen ts (MNE ) o f h o r se f r om L eve l 1a tG rubg raben . F o r MNE , t e e th h ave b een c omb ined w i th h eads .

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AP rzewa l sk it y pe h o r se i sr e p re sen ted b y am ode ra te ly w o rn r i gh t m and ibu la r M3 . T h i s a n ima l w a s ay oung a du l t a s t h e M3 e r up t sb e tween t h ree a nd ah a l ft of o u ry ea r s . A s e cond P rzewa l sk i t y pe h o r se i s r e p r e sen ted b y a n e x t rem e ly w o rn r i gh t m a x i l la ry M3 . T he M3 e r up t s b e tween t h ree a nd ah a lf a nd f o u r y ea r s a nd t h i s i n d iv idua l w a s q u i te o l dw h en i td i ed . Ay oung a n ima l i sr e p re sen ted b y ar i gh t , m and ibu la r t h i rd d e c iduou s i n c i so r . T h i s t o o th i nt h e d ome s t ic h o r se e r up t s b e tween s i x a nd t e n m on th s w i th t h e p e rman en t t h i rd i n c i so r e r up t ing b e twe en f ou r a nd ah a lf a nd f i ve y ea r s (W i l loughby 1 974 ) . T h i s i n c i so r s h owed l i t t le t o m ode ra te w ea r a nd i n d i ca t e s ar e la t ive ly y oung i n d iv idua l , c e r ta in ly l e s s t h an t wo y ea r s o l d a tt h e t i me o fd ea th . I ti si mpo s s ib le t o t e l l i ft h i s i n d iv idua lw a s aP r zewa l sk io rp ony t y pe h o r se . T h r ee p e rmanen t i n c i so r s r e cove r ed c anno t b e i d en t if ied t o e i the rP r zewa l sk io rT ype I h o r se . F ou rt e e th f r agm en t sw e r e b roken a nd t h ey c ou ld n o tb ea c cu r a t e ly i d en t if ied . B od ie s Two h o r se s a r e r e p re sen ted b y 7 0 b one f r agmen t s . Two f r agmen t s o fi l i a , t wo r i gh t f emu r s h af t f r agmen t s ( MNE= 1) a nd al e f tf emu rs h af tf r agmen tw e r ei d en t if i ed ;t h e se r e p re sen t r ump m ea t . Two t i b iae s h af t f r agm en t s a r e a l so a s soc ia ted w i th t h e h i nd q ua r te r . T h i s s ugge s t st wo h i nd h aun che s , w i th b one s i n c luded , w e re t a ken t o t h e c amp . As h ou lde r c u t i s r e p r e sen ted b ya nu n s ided s h af tf r agmen t o f ar a d iu s /u lna . T he h ead o f ar i gh t r i b , t o o l a rge t or e p r e sen tr e indee r , i sa s s igned t oh o r s e ; t h i s s i ng le r i b m ay b e r e p r e sen ta t ive o fa n a x ia l c u t o f m ea t . B one s f r om n onmea t b ea r ing p a r t s o f t h e h o r se c a r ca s s a r e m o s t h i gh ly r e p re sen ted , b o th a sN ISP a nd MNE a t t h e s i t e . A r i gh t h i nd leg , r e p r e sen ted b y ar i gh t d i s ta l m e ta ta r sa l a nd p ha lange s 1 ,2 ,a nd 3m ay b e a s soc ia ted w i th t h e t wo s h af t f r agmen t s o ft h e r i gh t f emu r m en t ioned a bove . T he d i s ta l e nd o f al e f tm e ta ta r sa l m ay b e a s soc ia ted w i th t h e l e f tf emo ra ls h af tf r agmen t . Ar i gh tf o re leg , r e p r e sen ted b yt h e d i s ta l e nd o f ar i gh tm e ta ca rpa la nd p ha lange s 1 ,2 ,a nd 3m ay b e a s soc ia ted w i th t h e u n s ided r a d iu s /u lna s h af t f r agmen t . A l l o f t h e l e g b one s o f t h i s m ed ium s i zed a n ima l h ave f u sed e p iphy s ea l j o in t s . I n h o r se s , t h e d i s ta l e p iphy s i s o f t h e m e taca rpa l u n i te sw i th t h e d i aphy s i sb e twe en s i x a nd e i gh teen m on th s . T he p rox ima l e p iphy sea l l i n e o ft h e p r ox ima l p ha lanx c l o s e s d u r ing t h e s i x th t h rough f i f teen th m on th s a nd t h e e p iphy s e s o fp ha lanx 1u n i te s p r io rt ob i r th a tt h e d i s ta l e nd a nd o ccu r s d u r ing t h e s i x th t h rough t we lf th m on th s a t t h e p rox ima le nd ( G e t ty 1 975 ) . As e cond l a rge ra n ima li sr e p r e s en ted b y ac omp le te p ha lanx 2 a nd p r ox im a l e nd o f ap ha lanx 3 .T h i s a n ima l i sl a rge r t h an a n ima l 1a nd m ay p o s s ib ly b e af u l ly g rown m a re o rs t a l l ion . B a sed o n c ompa r i son o fm ea su r em en t s o ff i r s t p ha lange s , t h e p o s tc ran ia l b one s r e cove red f r om G rubgaben AL I c ompa re f a vo r ab ly w i th t h o se r e cove r ed f r om G onne r sdo r f , A b r i P a taud , a nd l a rge r e qu id s a t S agva r . L eng th s o f t h e m e a su rab le p rox ima l p ha lange s f r om G rubgaben m ea su r e b e twe en 7 1 a nd 7 7 mm w h ich f a l l s w e l l w i th in t h e 6 8 t o 8 2 mm r a nge f o und a tG onne r sdo r f( Pop l in 1 968 : 1 58 ) . L ike t h e h o r se s r e cov e red a tA b r i P a taud , t h e A u s t r ian h o r se s " . .f i t a 6 1

s i ze r a nge o f mode ra te ly s i zed m ode rn f a rm h o r se . . .a m ed ium s i zed c hunky h o r se l i ke E . p r zew a l sk i " ( Sp ie s s 1 979 : 2 73 ) . Moder n P r zewa l s l c i s w e igh b e tween 2 50 a nd 3 00 k g ; a s im i la r ly s i zed a n ima l w ou ld h ave p rov ided 5 % o fl i v e b ody w e igh t i ne d ib le p ro te in ( Spe th 1 983 ) . T h e t h ree P rzewa l sk i t y pe h o r s e s w ou ld h ave , t h e ref o r e , p rov ided b e twe en 4 12 t o 4 95 k g o fm ea t f o r i n ha b i tan t s a tt h e s i t e . T he p ony s i zed a n ima l , r e p re sen ted b y c heek t e e th , w a s p robab ly s omewha t s ma l le r . At y p ica l We l sh p ony w e igh s a r ound 2 50 k g . I ft h e p ony r e p r e sen ted i n c u l tu ra l l e ve l 1w a s c ompa r ab le , t h e m ax imum m ea ta va i lab le f r om t h i sa n ima lw ou ld e qua l 1 37 k g . G rubg raben : L eve l2 A na tom ica lF requen c ie sa nd MN I : T h ree P r zewa l sk it y pe h o r se s a r e r e p re sen ted b y 8 9 t e e th a nd b one f r agmen t s ( Tab le 5 . 1 a nd F i gu re s 5 . 3 a nd 5 . 4 ) . A x ia l e l emen t s a r e r a re . Mea ty f o r e - a nd h i nd l imb s a r e r e p r e sen ted b y as c apu la , h ume ru s , f emu r , a nd t i b ia ; b one s o ft h e l owe r l i mb s a r e r a r e . T he t h ree h o r se s f r om l e ve l 2w ou ld h ave p rov ided 4 12 t o 4 95 k g o f u sab le m ea t i fa l m ea t w a s t r an spo r ted . H ead s T h ree h o r se h ead s a r e r e p re sen ted b y 2 1 t e e th f r agmen t s i n l e ve l 2 .Ay oung a n ima l i sr e p r e sen ted b y ar i gh t d e c iduou s m and ibu la rp r n2 . T h i st oo th e r up t s w i th in t h e f i r s t t wo w eek s o fb i r th . T he l i gh t a moun t o fw ea r o nt h i st oo th s ugge s t st h a t t h i s w a s av e ry y oung a n ima l a tt h e t i me o fd ea th . T he s l i gh t w ea r i n d ica te s t h a t i tw a s c e r t a in ly l e s s t h an o ne y ea r o l d w hen i td i ed . I t s y ou th a t te s t s t ot h e f a c t t h a t i tw a s s e le c ted f r om ah a r em g roup ; s t i l la ti t sm o the r ' ss i de w hen i td i ed . Af u ly a du l t h o r se i sr e p re sen ted b y aw o rn , l e f t m ax i l la ry PM2 . T h e a du l t p r emo la r e r up t s a tt wo a nd ah a lf y ea r s . A s im i la r ly w or n , l e f t m and ibu la r M3 r e p r e s en t s as e cond a du l t i n d iv idua l . T h i st oo th e r up t s w hen t h e h o r se i sb e tween t h re e a nd ah a lf a nd f ou r y ea r s o l d . I ti su n l ik e ly t h a t t h e se t wo c he ek t e e th c ame f r om t h e s ame i n d iv idua l . T he p r emo la r s hou ld s h ow mo re w ea rt h an t h e mo la r b e cau se i te r up ted a t l e a s t o ne y ea r b ef o r e t h e M3 . B o th i n d iv idua l s r e p re sen t P rzewa l sk it y pe a n ima l s . Ab roken p e rmanen t i n c i so rm ay a l so h ave b e en f r om o ne o ft h e se i n d iv idua l s . F ou r teen b roken a nd u n iden t if i ab le t oo th f r agmen t sw e re f o und i nl e ve l2 . B od ie s T he a x ia ls k e le ton i sr e p re sen ted b y af r agmen t o f al e f tp e lv i s a nd as a c rum . No r i b sw e re r e cove red a nd o n ly 1u n iden t if ied v e r teb r a f r agmen t w a s f o und . F o re l imb s a r e r e p re sen ted b y a l e f ts c apu la , l e f ta nd r i gh th ume r i , a nd al e f tr a d iu s /u lna . T h re e h aunch e s a r e r e p re sen ted b y af r agm en t o f al e f t f emu r a nd o ne r i gh t a nd t wo l e f t t i b iae . L owe r l i mb e l emen t s a r e q u i te r a re i nl e ve l 2 .As i ng le m e ta ta ta r sa l , p robab ly a s soc ia ted w i th o ne o ft h et h ree t i b iae w a sr e cove red . U n f o r tuna te ly , a lb one s w e r et o o f r agmen ted t op e rm i t a na tom ica l r e f i t t ing . O n ly o ne p ha la nx , as e cond , w a s r e cove red a nd n o s e samo id s w e re i d en t if i ed . U n l ike l e ve l 1 ,m ea ty p a r t so ft h eh aunch o fa tl e a s t t h ree h i ndqua r te r sa r er e p r e sen ted b yt h et i bae .

F i g . 5 . 3 N umbe ro fi d en t if ied s pec imens o fh o r se f r om L eve l 2a tG rubg raben . U nshaded a r ea s r e p re sen t b ones a bsen t

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