The Paleoecology of Lower Magdalenian Cantabrian Spain 9780860549529, 9781407350295

Table of contents :
Front Cover
Copyright
TABLE OF CONTENTS
LIST OF TABLES
LIST OF FIGURES
ACKNOWLEDGEMENTS
DEDICATION
ABSTRACT
CHAPTER 1: INTRODUCTION
CHAPTER 2: TAPHONOMY
CHAPTER 3: SPECIES CHARACTERISTICS AND ECOLOGIES
CHAPTER 4: MODERN MICROFAUNA OF CANTABRIAN SPAIN
CHAPTER 5: EL JUYO MICROFAUNA AND PALEOENVIRONMENTAL INTERPRETATION
CHAPTER 6: BIOMETRICAL ANALYSIS
CHAPTER 7: OTHER CLIMATIC INDICATORS FROM EL JUYO
CHAPTER 8: MICROFAUNA FROM CANTABRIAN REGION SITES
CHAPTER 9: EXPLOITATION OF FUR-BEARING MAMMALS
CHAPTER 10: HUMAN ENVIRONMENTAL INTERACTIONS
CHAPTER 11: SUMMARY AND CONCLUSIONS
APPENDIX A
REFERENCES

Citation preview

The Paleoecology of Lower Magdalenian Cantabrian Spain

James T. Pokines

BAR International Series 713 1998

Published in 2019 by BAR Publishing, Oxford BAR International Series 713 The Paleoecology of Lower Magdalenian Cantabrian Spain © James T. Pokines and the Publisher 1998 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 9780860549529 paperback ISBN 9781407350295 e-book DOI https://doi.org/10.30861/9780860549529 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 1998. This present volume is published by BAR Publishing, 2019.

BAR

PUBLISHING BAR titles are available from:

E MAIL P HONE F AX

BAR Publishing 122 Banbury Rd, Oxford, OX2 7BP, UK [email protected] +44 (0)1865 310431 +44 (0)1865 316916 www.barpublishing.com

TABLE OF CONTENTS

L IST O F TABLES

i

L IST O FF IGURES ACKNO WLEDGE MENTS

v i i

ABSTRACT

i x

CHAPTER : I NTRODUCT ION

1

I

TAPHONO MY

1 3

I I

S PEC IES CHARACTER IST ICS AND E COLOG IES

3 3

I V

MODERN M ICROFAUNA OF CANTABR IAN S PA IN

6 1

V

E LJ UYO M ICROFAUNA AND P ALEOENV IRON MENTAL I NTERPRETAT ION

7 1

V I

B IOMETR ICAL ANALYS IS

8 5

V I I

OTHER CL I MAT IC I ND ICATORS F ROM E LJ UYO

1 03

V I I I

M ICROFAUNA F ROM CANTABR IAN R EG ION S ITES

13

I X

EXPLO ITAT ION O F FUR -BEAR ING MAM MALS

1 37

X

HU MAN ENV IRON MENTAL I NTERACT IONS

X I

S UM MARY AND CONCLUS IONS

APPEND IX A : METR ICAL DATA REFERENCES

. .

1 51 1 59

. .

1 67 1 73

L IST OF TABLES

T ab le 2 . 1 : M ic rof auna lR ema ins f r om E lJ uyo b y S i z e C la s sa nd L eve l T ab le 2 . 2 : C ompa r i son o fB a rn Ow l P r ey S ke le ta l R ep re sen ta t ion ( % )

1 8 1 9

T ab le 2 . 3 : E lJ uyo S ke le ta l E lemen tB reakage -Sma l l

2 0

T ab le 2 . 4 : E lJ uyo S ke le ta l E lemen tB reakage -Med ium T ab le 2 . 5 : E lJ uyo S ke le ta lE lemen tB reakage -La rge

2 1 2 2

T ab le T ab le T ab le T ab le T ab le

2 3 2 8 3 0 3 1 3 2

2 . 6 : E lJ uyo S ke le ta l E lemen tB reakage -TaI p a 2 . 7 : S umma ry o fM ic rover teb ra te T aphonomy 2 . 8 : P o s tc ran ia lE p iphy sea l F u s ion f o rA l lL eve l s o fE lJ uyo 2 . 9 :A pod emu sA ge C la s se s B a sed u pon T oo th Wea r 2 . 10 : E r inaceu se uropaeu sR ema in s f r om E lJ uyo

T ab le 3 . 1 : S umma ry o fE co log ica lC ha rac te r i s t ic s

4 6

T ab le 4 . 1 : D i s t r ibu t ion o fT rap N igh t s( Can tab r ia ) T ab le 4 . 2 : S pec ie sD imen s ion s o fC an tab r ian Sma l l Mamma l s

6 1 6 2

T ab le 4 . 2 .1 :A pod emu ss y lva t icu sD imen s ion s T ab le 4 . 2 .2 :A pod emu sf lav ico l l i sD imen s ion s

. .

6 2 6 2

T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le

4 . 2 .3 : R a t tu sr a t tu sD imen s ion s 4 . 2 .4 : M icro tu sa g re s t i sD imen s ion s 4 . 2 .5 :S orex c o rona tu sD imen s ion s 4 . 2 .6 :S orex m inu tu sD imen s ion s 4 . 2 .7 :N eomy sf od ien s D imen s ion s 4 . 2 .8 : C roc idu ra r u s su la D imens ion s 4 . 3 : S pec ie sD i s t r ibu t ion a mong T rapp ing L oca t ion s( Can tab r ia ) 4 . 4 : Moder nS ma l lF auna o ft h eE lJ uyo A r ea f r om T y to a l ba a nd B u teo b u teo P e l le t s 4 . 5 : M ic rof auna f r om B ar n Ow l P e l le t s , C an tab r ian C o rd i l le ra 4 . 6 : T y to a l ba Mamma lD ie tf r om C a l if or n ia , U .S .A 4 . 7 : Modem T e r r e s t r ia l Sma l l Mamma l P rey o fB a rn Ow l s f r om t h eE nv i rons o fE lJ uyo

6 2 6 2 6 2 6 3 6 3 6 3 6 4 6 7 6 9 6 9 7 0

T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le

5 . 1 : I d en t if i ed M ic rof auna l T ee th f r om E lJ uyo 5 . 2 : Ow l In t roduced M ic rof auna f r om L eve l s 1 a nd 1 0 5 . 3 : Ow l In t roduced M ic rof auna f r om L eve l s 8a nd 9 5 . 4 : Ow l In t roduced M ic rof auna f r om L eve l 7 5 . 5 : Ow l In t roduced M ic rof auna f r om L eve l 6 / 9 5 . 6 : Ow l In t roduced M ic rof auna f r om L eve l6 5 . 7 : Ow l In t roduced M ic rof auna f r om L eve l 5 5 . 8 : Ow l In t roduced M ic rof auna f r om L eve l4 R 5 . 9 : Ow l In t roduced M ic rof auna f r om L eve l4 / 4S 5 . 10 : Ow l In t roduced M ic rof auna f r om L eve l3 5 . 11 : Ow l In t roduced M ic rof auna f r om L eve l s2 / 4 a nd 2 5 . 12 : Ow l In t roduced M ic rof auna f r om L eve l 1 5 . 13 : C ompa r i son o fE lJ uyo E nv i ronmen t sb y B ioma s sU n i t s

7 2 7 4 7 5 7 6 7 7 7 8 7 7 7 9 8 0 8 1 8 1 8 2 8 2

. .

T ab le 6 . 1 : Max imum L eng th ( mm )o f, M , o fA r v i co la s pp . f r om E lJ uyo T ab le 6 . 2 : Max imum L eng th ( mm )o f 3M 3 o fA rv ico la s pp . f r om E lJ uyo T ab le 6 . 3 : C ompa r i son o f, M ,L eng th ( mm )o fA r v i co la

8 5 8 5 8 8

T ab le 6 . 4 : C ompa r i son o f 3M 3L eng th ( mm )o fA rv ico la T ab le 6 . 5 : Max imum L eng th ( mm )o f, M , o fM icro tu so economu sf r om E lJ uyo

8 8 8 9

T ab le 6 . 6 : C ompa r i son o f, M , L eng th ( mm )o fM icro tu so economu s T ab le 6 . 7 : Max imum L eng th ( mm )o f, M , o fP i tymy ss pp . f r om E lJ uyo T ab le 6 . 8 : Max imum L eng th ( mm )o f, M , o fP i tymy s s pp . f r om E lJ uyo

9 0 9 0 9 0

T ab le 6 . 9 : C ompa r i son o f, M , L eng th ( mm )o fP i tymy ss pp T ab le 6 . 10 : Max imum L eng th ( mm ) o f 3M 3 o fP i tymy sl u s i tan icu sf r om E lJ uyo

9 0 9 1

T ab le 6 . 11 : C ompa r i son o f 3M 3 L eng th ( mm ) o fP i tymy sl u s i tan icu s

9 1

T ab le T ab le T ab le T ab le

9 1 9 1 9 2 9 2

6 . 12 : 6 . 13 : 6 . 14 : 6 . 15 :

Max imum L eng th ( mm )o f 3M 3 o fP i tymy sp y rena icu sf r om E lJ uyo C ompa r i son o f 3M 3 L eng th ( mm )o fP i tymy sp y rena icu s Max imum L eng th ( mm )o f, M , o fM icro tu sa g re s t i s la rva l i sf r om E lJ uyo C ompa r i son o f, M , L eng th ( mm )o fM i cro tu sa g re s t i s larva l i s

T ab le 6 . 16 : Max imum L eng th ( mm )o f 2M 2 o fM icro tu sa g re s t i sf r om E lJ uyo T ab le 6 . 17 : C ompa r ison o f 2M 2L eng th ( mm )o fM icro tu sa g re s t i s T ab le 6 . 18 : C ompa r ison o f, M , L eng th ( mm )o fM icro tu sn i va l i s T ab le 6 . 19 .1 : Max imum L eng th ( mm )o f, M , o fA pod emu sf r om E lJ uyo

9 2 . .

9 2 9 3 9 3

T ab le 6 . 19 .2 : Max imum L eng th ( mm )o f, M , o fA pod emu sf r om E lJ uyo

9 3

T ab le 6 . 19 .3 : Max imum L eng th ( mm )o f' M ' o fA pod emu sf r om E lJ uyo T ab le 6 . 19 .4 : Max imum L eng th ( mm )o f 2M 2 o fA pod emu sf r om E lJ uyo T ab le 6 . 20 .1 : C ompa r ison o f, M , L eng th ( mm )o fA pod emu s

9 4 9 4 9 4

T ab le 6 . 20 .2 : C ompa r i son o f, M ,L eng th ( mm )o fA pod emu s T ab le 6 . 20 .3 : C ompa r i son o f' M IL eng th ( mm ) o fA pod emu s

9 4 9 4

T ab le 6 . 20 .4 : C ompa r ison o f 2M 2 L eng th ( mm )o fA pod emu s T ab le 6 . 2 1 .1 : Max imum R amu sH e igh t( mm ) o fS orex c orona tu sf r om E lJ uyo

9 4 9 5

T ab le 6 . 2 1 .2 : Max imum L eng th ( mm )o f, M , o fS orex c orona tu sf r om E lJ uyo T ab le 6 . 21 .3 : Max imum L eng th ( mm )o f, M ,o fS o rex c orona tu sf r om E lJ uyo

9 5 9 5

T ab le 6 . 2 1 .4 : Max imum L eng th ( mm )o f 3M 3 o fS orex c orona tu sf r om E lJ uyo

9 5

T ab le 6 . 22 .1 : C ompa r ison o fR amu sH e igh t( mm )o fS o rex c o rona tu s

9 6

T ab le 6 . 22 .2 : C ompa r i son o f, M , L eng th ( mm )o fS orex c o rona tu s T ab le 6 . 22 .3 : C ompa r i son o f 2M 2 L eng th ( mm )o fS orex c o rona tu s

9 6 9 6

T ab le T ab le T ab le T ab le

6 . 22 .4 : 6 . 23 .1 : 6 . 23 .2 : 6 . 23 .3 :

9 6 9 6 9 6 9 6

T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le T ab le

6 . 23 .4 : Max imum L eng th ( mm )o f, M , o fS orex m inu tu sf r om E lJ uyo 6 . 24 .1 : C ompa r i son o fR amu sH e igh t( mm )o fN eomy sf od ien s 6 . 24 .2 : Max imum L eng th ( mm )o f, M , o fN eomy sf od ien sf r om E lJ uyo 6 . 24 .3 : Max imum L eng th ( mm )o f, M , o fN eomy sf od ien sf r om E lJ uyo 6 . 25 : H ume ru sL eng th ( mm )o fT a lpa f r om E lJ uyo 6 . 26 .1 : C ompa r ison o f Mand ibu la rL eng th ( mm )o fE r inaceu se u ropaeu s 6 . 26 .2 : C ompa r i son o f, M , L eng th ( mm ) o fE r inaceu se u ropaeu s 6 . 26 .3 : C ompa r i son o f' M ' L eng th ( mm )o fE r inaceu se u ropaeu s 6 . 27 .1 : C ompa r i son o f, M , L eng th ( mm )o f Mu s te la n i va l i s 6 . 27 .2 : Max imum L eng th ( mm )o f 4P4 o f Mu s te la n i va l i sf r om E lJ uyo 6 . 27 .3 : C ompa r i son o f 4P4 L eng th ( mm )o f Mu s te la n i va l i s

C ompa r i son o f, M , L eng th ( mm ) o fS orex c o rona tu s C ompa r i son o fR amu sH e igh t( mm )o fS o rex m inu tu s Max imum L eng th ( mm )o f, M , o fS orex m inu tu sf r om E lJ uyo Max imum L eng th ( mm )o f, M , o fS orex m inu tu sf r om E lJ uyo

. .

9 6 9 6 9 6 9 7 9 7 9 8 9 8 9 8 9 8 9 8 9 8

T ab le 7 . 1 : L a rge F auna f r om E lJ uyo T ab le 7 . 2 : Mo l lu scan R ema in s f r om E lJ uyo T ab le 7 . 3 : Mac robo tan ica lR ema in sf r om E lJ uyo

1 05 1 06 10

T ab le 8 . 1 : M icrof auna f r om C an tab r ian Mou s te r ian L eve l s T ab le 8 . 2 : M ic rof auna f r om C an tab r ian E a r ly U ppe rP a leo l i th ic L eve l s

1 26 1 27

T ab le 8 . 3 : M ic rof auna f r om C an tab r ian S o lu t rean L eve l s T ab le 8 . 4 : M ic rof auna f r om Magda len ian L eve l s T ab le 8 . 5 : L owe r Magda len ian L eve l Vo fE r ra l la

1 29 1 3 1 1 25

T ab le 8 . 6 : M ic rof auna f r om C an tab r ian A z i l ian L eve l s T ab le 8 . 7 : M ic rof auna f r om C an tab r ian P o s t -Pa leo l i th ic L eve l s T ab le T ab le T ab le T ab le T ab le

.

9 . 1 : F u r -bea r ing T axa f r om Mous te r ian L eve l s 9 . 2 : F u r -bea r ing t a xa f r om E a r ly U ppe rP a leo l i th ic L eve l s 9 . 3 : F u r -bea r ing T axa f r om S o lu t rean L eve ls 9 . 4 : F u r -bea r ing T axa f r om Magda len ian L eve ls 9 . 5 .1 : T ee th o fF u r -bea r ing T axa f r om E lJ uyo

1 34 1 35 1 43 1 44 1 46 1 47 1 4 1

T ab le 9 . 5 .2 : P o s tc ran ia lR ema ins o fF u r -bea r ing T axa f r om E lJ uyo

1 4 1

T ab le 9 . 6 :F u r -bea r ing T axa f r om A z i l ian L eve l s

1 50

1 1

T ab le A .1 : Max imum L eng th ( mm ) o f, M , o fA r v ico la T ab le A .2 : Max imum L eng th ( mm ) o f 3M 3 o fA r v ico la

1 67 1 67

T ab le A .3 : Max imum L eng th ( mm ) o f, M , o fM ic ro tu so economu s T ab le A .4 : Max imum L eng th ( mm )o f, M , o fP i tymy ss pp

1 68 1 68

T ab le A .5 : Max imum L eng th ( mm )o f, M2o fP i tymy ss pp

1 68

T ab le A .6 : Max imum L eng th ( mm )o f 3M 3 o fP i tymy sl u s i tan icu s T ab le A .7 : Max imum L eng th ( mm )o f 3M 3 o fP i tymy sp y rena icu s

1 69 1 69

T ab le A .8 : Max imum L eng th ( mm )o f, M , o fM icro tu sa g re s t i s lar va l i s T ab le A .9 : Max imum L eng th ( mm )o f 2M 2 o fM icro tu sa g re s t i s

1 69 1 70

T ab le A .10 : Max imum L eng th ( mm )o f, M , o fM icro tu sn i va l i s T ab le A .11 : Max imum L eng th ( mm )o f, M , , 2M 2,1M la nd 2M 2 o fA pod emu s

1 70 1 70

T ab le A .12 : Max imum R amu s H e igh ta nd L eng th o f M a nd 3M 3( mm )o fS orex c orona tu s T ab le A .13 : Max imum R amu sH e igh ta nd L eng th o f a nd , M 3( mm ) o fS o rex m inu tu s T ab le A .14 : Max imum L eng th ( mm )o f, M , a nd ' M ' o fE r inaceu se u ropaeu s

1 7 1 1 7 1 1 72

T ab le A .15 : Max imum L eng th ( mm ) o f M 1, P — a nd 4P4o f Mu s te la n i va l i s

1 72

1

1 , 2 M2

1 M1e 2 M2

1

4

4

i v

L IST OF F IGURES

F igu re 1 . 1 : S i te sD i scu s sed i nC hap te r 1 F igu re 1 . 2 : T opog raph ic S u r round ing so fE lJ uyo

2 7

F igu re 1 . 3 : A r ea lE x ten to fE lJ uyo E xcava t ion s

9

F igu re 1 . 4 : S t ra t ig raph ic P rof i le o fE lJ uyo

1 0

F igu r e 3 . 1 : Mode rn R ange o fG l i sg l i s F igu re 3 . 2 : Mode rn R ange o fE l iomy sq ue rc inu s

4 7 4 7

F igu re 3 . 3 : Mode rn R ange o fM icro tu sa g re s t i s F igu re 3 . 4 : Mode rn R ange o fM icro tu sa rva l i s

4 7 4 8

F igu re 3 . 5 : Mode rn R ange o fM icro tu sn i va l i s F igu r e 3 . 6 : Moder nP a lea r c t ic R ange o fM icro tu s o economu s

4 8 4 8

F igu re 3 . 7 : Mode rn R ange o fM icro tu sg rega l i s F igu re 3 . 8 : Moder nR ange o fC le th r ionomy sg lareo lu s F igu re 3 . 9 : Moder nR ange o fP i tymy sl u s i tan icu s

4 9 4 9 4 9

F igu re F igu r e F igu r e F igu r e

3 . 10 : 3 . 11 : 3 . 12 : 3 . 13 :

Modern Moder n Moder n Moder n

5 0 5 0 5 0 5 1

F igu re F igu re F igu re F igu r e F igu r e F igu re F igu r e F igu r e F igu re F igu re F igu r e F igu re F igu re F igu re F igu r e F igu r e F i gu r e

3 . 14 : 3 . 15 : 3 . 16 : 3 . 17 : 3 . 18 : 3 . 19 : 3 . 20 : 3 . 2 1 : 3 . 22 : 3 . 23 : 3 . 24 : 3 . 25 : 3 . 26 : 3 . 27 : 3 . 28 : 3 . 29 : 3 . 30 :

Moder nR ange o fA pod emu sf la v ico l l i s Moder nR ange o fM ic romy sm inu tu s Mode rn R ange o fC roc id ura r u s su la Moder nR ange o fS o rex m inu tu s Moder nR ange o fS o rex c orona tu s Moder nR ange o fN eomy sa noma lu s Moder nR ange o fN eomy sf od ien s Moder nR ange o fT a lpa e uropaea Moder nR ange o fT a lpa o cc id en ta l i s Moder nR ange o fG a l emy sp y rena icu s Moder nR ange o fE r inaceu se uropaeu s Moder nR ange o f Ma rmo ta ma rmo ta Mode rn R ange o fO ryc to lagu sc un icu lu s Moder nR ange o fL epu sc apen s i s Moder nR ange o fL epu st i m idu s Moder nP a lea rc t ic R ang eo f Mu s te la n i va l i s Moder nP a lea rc t ic R ange o f Mu s te la e rm inea

F igu r e F igu re F igu re F igu r e

3 . 31 : 3 . 32 : 3 . 33 : 3 . 34 :

Moder n Moder n Moder n Moder n

R ange R ange R ange R ange

R ange R ange R ange R ange

o fP i tymy sp y rena icu s o fA r v i co la t e r re s t r i s o fA rv i co la s ap idu s o fA pod emu ss y lva t icu s

o f Mu s te la p u tor iu s o f Mu s te la l u t reo la o f Mane sm ar te s o f Man e sf o ina

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

5 7 5 7 5 7 5 8

F igu re 3 . 35 : Mode rn R ange o fL u t ra l u t ra F igu r e 3 . 36 : Modern R ange o f Me /e sm e /e s F igu re 3 . 37 : Moder nP a lea rc t ic R ang eo fG u lo g u lo

5 8 5 8 5 9

F igu re 3 . 38 : Moder nP a lea rc t ic R ang eo fV u /pe sv u /pe s F igu re 3 . 39 : Moder nP a lea rc t ic R ang eo fA lopex l agopu s F igu re 3 . 40 : Moder nR ange o fF e / i ss i lve s t r i s

5 9 5 9 6 0

F igu re 3 . 4 1 : Moder nR ange o fF e / i sl y nx F igu re 3 . 42 : Moder nR ange o fG ene t ta g ene t ta

6 0 6 0

F igu re 4 . 1 : Map o fC an tab r ia S how ing T rapp ing L oca t ion s 1 5 a nd R oo s tL oca t ion s A , Ba nd C

6 3

F igu re 4 . 2 : E lJ uyo Q ua r ry S i te # 2( Roo s tA )

6 6

F igu re 4 . 3 : E lJ uyo Q ua r ry S i te 3a nd 3 A ( Roo s tB ) F igu re 4 . 4 : E nv i rons o fE lJ uyo O ve r look ing N ea rby Q ua r ry F igu re 4 . 5 : A rea l E x ten t( % ) o fH ab i ta tT ype s i nt h e Moder nE nv i rons o fE lJ uyo D e r ived f r om B ar n Ow l D ie t . . . . .

6 6 6 6 6 7

F igu r e 5 . 1 : P a leoenv i ronmen to fE lJ uyo L eve l s 1 0a nd 1 F igu r e 5 . 2 : P a leoenv i ronmen to fE lJ uyo L eve l s 8a nd 9

7 4 7 5

F igu re 5 . 3 : P a leoenv i ronmen to fE lJ uyo L eve l7 F igu r e 5 . 4 : P a leoenv i ronmen to fE LJ uyo L eve l6

7 6 7 8

F igu r e 5 . 5 : P a leoenv i ronmen to fE lJ uyo L eve l4 R F igu r e 5 . 6 : P a leoenv i ronmen to fE LJ uyo L eve l s4 / 4S

7 9 8 0

F igu re 5 . 7 : C ompa r i son o fE lJ uyo E nv i ronmen t sb y B ioma s sU n i t s

8 3

Max imum Max imum Max imum Max imum

L eng th Mea su remen t s T aken o nM ic ro t ine Mo la r s

F igu re F igu re F igu re F igu re

6 . 1 .1 : 6 . 1 .2 : 6 . 1 .3 : 6 . 1 .4 :

F igu re F igu re F igu re F igu r e F igu re F igu r e F igu r e F igu r e F igu re F igu re F igu r e F igu re F igu r e F igu re

fA r v ico la 3M 3 L eng th ( mm ) f r om E lJ uyo 6 . 3 : D i s t r ibu t ion o fM icro tu so economu s , M , L eng th ( mm )f r om E lJ uyo 6 . 4 : D i s t r ibu t ion o fP i tymy s , M , L eng th ( mm )f r om E lJ uyo 6 . 5 : D i s t r ibu t ion o fP i tymy s, M ,Leng th ( mm )f r om E lJ uyo 6 . 6 : D i s t r ibu t ion o fP i tymy sl u s i tan icu s 3M 3 L eng th ( mm ) f r om E lJ uyo 6 . 7 : D i s t r ibu t ion o fP i tymy sp y rena icu s 3M 3 L eng th ( mm )f r om E lJ uyo 6 . 8 : D i s t r ibu t ion o 6 . 9 : D i s t r ibu t ion o fM icro tu sa g re s t i s lar va l i s , M ,L eng th ( mm ) f r om E lJ uyo 6 . 10 : D i s t r ibu t ion o fM icro tu sa g re s t i s 2M 2 L eng th ( mm ) f r om E lJ uyo 6 . 11 : D i s t r ibu t ion o fA pod emu s , M , L eng th ( mm ) f r om E lJ uyo 6 . 12 : D i s t r ibu t ion o fA pod emu s, M ,L eng th ( mm )f r om E lJ uyo 6 . 13 : D i s t r ibu t ion o fA pod emu s 1M 1L eng th ( mm ) f r om E lJ uyo 6 . 14 : D i s t r ibu t ion o fA pod emu s 2M 2 L eng th ( mm ) f r om E lJ uyo 6 . 15 : D i s t r ibu t ion o fS orex c o rona tu sR amu sH e igh t( mm ) f r om E lJ uyo 6 . 16 : D i s t r ibu t ion o fS orex c orona tu s , M , L eng th ( mm )f r om E lJ uyo

L eng th Mea su remen t sT aken o nA pod emu s mo la r s L eng th Mea su remen t sT aken o n S h rew S pec ies ( Mo la rL eng th a nd R amu s H e igh t )

8 6 8 6 8 6

F igu re 6 . 17 : D i s t r ibu t ion o fS orex c o rona tu s, M ,L eng th ( mm ) f r om E lJ uyo F igu r e6 . 18 : D i s t r ibu t ion o fS orex c orona tu s 3M ,L eng th ( mm )f r om E lJ uyo

8 7 8 7 8 7 9 9 9 9 9 9 9 9 9 9 9 9 1 00 1 00 1 00 1 00 1 00 1 00 1 0 1 1 0 1 1 0 1 1 0 1 1 0 1

F igu re 7 . 1 : R e la t ive S ea L eve lD u r ing t h eU ppe rP a leo l i th ic

1 04

F igu re 8 . 1 : C an tab r ian M ic rof auna l S i te sD i scu s sed i nt h eT ex t

14

F igu re 8 . 2 : P a leoenv i ronmen to fE r ra l la L eve lV

1 25

F igu r e9 . 1 : C an tab r ian M idd le a nd U ppe rP a leo l i th ic S i te sY ie ld ing t h e R ema in so fF u r -Bea r ing S pec ie s

1 38

L eng th Mea su remen t sT aken o n T a lpa H umerus eng th Mea su r emen t sT aken o nt h e Mo la r s a nd Mand ib le o fE r inaceu se u ropaeu s F igu re 6 . 1 .5 : Max imum L eng th Mea su remen t sT aken o nt h e Mo la r so f Mu s te la n i va l i s F igu re 6 . 1 .6 : Max imum L fA rv ico la , M , L eng th ( mm )f r om E lJ uyo F igu re 6 . 2 : D i s t r ibu t ion o

v i

ACKNO WLEDGE MENTS

T h i s wo rk i sd rawn l a rge ly f r om my P h .D . d i s se r ta t ion a tt h e D epa r tmen t o f A n th ropo logy o f t h e U n ive r s i ty o f C h icago . T hank s g o t o my d i s se r ta t ion c omm i t tee f o r t h e i r g u idance a nd f o rbea rance : D r . L e s l ie G . F reeman a nd D r . M ichae l D ie t le r ( Un ive r s i ty o f C h icago ) , D r . J u l ian K e rb i s P e te rhans ( F ie ld Mu seum o f N a tu ra l H is to ry , C h icago ) a nd D r . J ane B u ik s t ra ( Un ive r s i ty o fA r izona ) . D r . J oaqu in G onz ä lez E chega ray ( I n s t i tu to p a ra I nve s t iga c ione s P reh i s tö r ica s , S an tande r , S pa in ) a l so p rov ided i n va luab le s uppo r t , y ea r so fi n s t ruc t ion , a nd a cce ss t od a ta . C ompu t ing s uppo r tw a s s upp l ied b yL .R eed A s soc ia te s . S eve ra l s e a son s o fi n ve s t iga to r s a nd f i e ld s c hoo l s t uden t s s pen t w eek s i nt h e c a ref u l c o l lec t ion o ft h i s d a ta . T he se h ave b een l e d b yC a r lo sF e rnändez R od r iguez , Mon ica A ngu lo I ba r lez , S t even K lu sken s , H ea the r S t e t t le r , Ma r cy K rupa a nd L ance R eed . Imu s ta l so t h ank t h eD epa r tmen to f Mamma l s o ft h e F ie ld Mu seum o fN a tu ra lH i s to ry a nd e spec ia l ly D r . W i l l iam S t an ley , D r . B ruce P a t te r son , Me l i s sa Mo ra le s , M ich i S o lomon a nd D r . S t even G oodman f o rt h e i ra dv ice , h e lp a nd s pace . A l so i n va luab le w a s t h e h e lp , a cce s s t o d a ta a nd g ene ra l s av ing g race o f t h e s t af f o f t h e C en t ro d e I nve s t igac iön y Museo d e A l tam i ra ( San t i l lana d e l Ma r , S pa in ) : C a rmen d e l a sH e ra s Ma r t in a nd D r . J o sé A . L a she ra sC o r ruchaga . Many c o l league s h ave c on t r ibu ted i n d i rec t ly t ot h i s wo rk , t h rough t h e i r y ea r s o fa dv ice a nd e ncouragemen t . T he se i n c lude A p r i l Nowe l l , D r . G a i lL a r sen P e t e rk in , D r . H ea the rP r ice , D r . A nne P ike -Tay , a nd D r . H e id iK nech t . F und ing f o r t h i s r e sea r ch w a s s upp l ied b y N a t iona l S c ience F ounda t ion D i s se r ta t ion I mp rovemen t G ran t S BR -9408920 , t h e I n s t i tu to p a ra I nve s t igac ione s P reh i s tö r ica s( San tande r , S pa in ) , a nd b y t h e D epa r tmen t o f Mamma l s , F i e ld Mu seum o fN a tu ra l H i s to ry ( Ch icago ,U .S .A . ) .

v i i

ABSTRACT

T he t e r re s t r ia l mamma l ian m ic rof auna f r om t h e L owe r Magda len ian s i t e o fE l J uyo ( San tande r , S pa in ) a nd r e la ted s i te s a r e u sed t or e con s t ruc tt h e p a leoenv i ronmen t C an tab r ian S pa in . T h i s l ow land c ave s i te i sl o ca ted n ea r t h e mode rn c oa s t l ine a nd p rov ides 3 m o f s t ra t if ied , u nd i s tu rbed L owe r Magda len ian d epo s i t s . O ccupa t ion s panned a pp rox ima te ly 1 , 000 y ea r s c a . 1 4 ,000 B .P . F ine s c reen ing h a s y i e lded al a rge s amp le o fs ma l l m amma l r ema in s . T aphonom ic a na ly s i s i n d ica te s t h a t t h e se r ema in s w e re a ccumu la ted a s ar e su l t o f b a rn o w l ( T y to a l ba ) p e l le t r egu rg i ta t ion w i th in t h e c ave v e s t ibu le . T he s ma l l m amma l ian f a una i sd om ina ted b y m ic ro t ine s , p r ima r i ly o f t h e t a xa A rv ico la c f . t e r re s t r i s a nd M icro tu s o economu s , a nd i n c ludes M icro tu sn i va l i s ,M a g re s t i s , M a r va l i s , P i tymy sl u s i tan icu s , a nd P .p y rena icu s . T he l a te s u rv iva l o ft h en ow e x t inc t m ic ro t ine s pec ie s P l iomy s l e nk i i sa l so i n d ica ted . A l so p r e sen t a r e t h e mu r id s A pod emu sf lav ico l l i s a nd A . s y lva t icu s , t h e i n sec t ivo re s T a lpa e u ropaea , Tc aeca , S o rex c orona tu s , S .m inu tu s , N eomy sf od ien s , N . a noma lu s , a nd E r inaceu s e uropaeu s . A s i ng le l a gomo rph s pec ie s 07c to lagu s c un icu lu s i sp r e sen t . Sma l l c a rn ivo r e s a r e r ep re sen ted b y Mu s te la n i va l i s , M . p u tor ius a nd V u lpe s v u lpe s .

E l J uyo 's l a rge f a una l s pec ie s i n c lude ( i n d e scend ing o rde r o f a bundance ) C er vu s e l aphu s ,

C apreo lu s c apreo lu s , B i son b ona su s , E quu s c aba l /u s , C apra p y rena ica , S u s s c rof a , P an the ra l e o , C an i s l u pu s , a nd U r su s a rc to s .

H uman f ood r ema in s a l so i n c lude t h e a bundan t r ema in s o f m a r ine mo l lu sc s pec ie s P a te l la v u lga ta a nd L i t tor ina

l i t torea a nd A t lan t ic s a lmon ( Sa lmo s a la r ) . An ew m e thod o fp a leoenv i ronmen ta lr e con s t ruc t ion i sp r e sen ted , b a sed u pon t h er e la t ive p ropor t ion s o ft e r r e s t r ia l m amma l ian m ic rof auna . A na ly s i s o ft h e modem b ar n o w l d i e t i n d ica te s t h a t t h e r e la t ive b i oma ss c on t r ibu t ion s o fm ic romamma l ian p rey a r e h igh ly i n d ica t ive o f t h e r e la t ive h ab i ta t p ropor t ion s w i th in t h e o w l 's h un t ing r ad ius : w e t land s , f o re s t , h um id m eadow , h ea th land , d ry m eadow , e t c . T he se d a ta a r e c ompa red w i th o t he r s ou r ce s ( po l len , l a rge mamma l ian f a una , a qua t ic f a una , s ea l e ve l c hange s a nd m ac robo tan ica l r ema in s ) t op roduce ac l ea r e r p i c tu re o ft h e L owe r Magda len ian s e t t ing . T h i s a na ly s is i s e x tended t ot h er ema in ing p ha se s o ft h eU ppe rP a leo l i th ic . T he se s ou rce s i n d ica te t h a tt h e c l ima te d u r ing t h e o ccupa t ion o fE lJ uyo w a s v e ry h um id a nd c o ld , w i th o pen c oa s ta l h ea th t h e d om inan tt e r ra in t y pe . Sma l la r ea so ff o r e s ta nd g ra s s land w e r e a l so p r e sen t . R ap id c l ima t ic d e te r io ra t ion o ccu r r ed d u r ing t h e d epo s i t ion o ft h e u ppe ro ccupa t ion l e ve l s . T he m ed ium m ic rof auna l s pec ie s E r inaceu s e u ropaeu s ( h edgehog ) e n te red t h e s i te a s ac ommen sa l a nd w a s n o t o w l o rh uman p r ey . T he l a rge r m ic rof auna ( p r ima r i ly mu s te l id s ) a r e p o s i ted t oh ave e n te red t h e c ave a st h e r e su l t o fh uman f u r t r app ing a c t iv i ty , s i nce t h e se a r e r e p r e sen ted a lmo s t e xc lu s ive ly b y f oo t e l emen t s . A l a rge r p a t ter no ff u ra cqu is i t ion i sd e tec ted f o rt h eC an tab r ian U ppe rP a leo l i th ic . T he U ppe r P a leo l i th ic h uman e co log ica l a dap ta t ion s i nC an tab r ian S pa in a r e e xam ined a st h ey r e la te t ot h i s i n te rp r e ta t ion o f t h e p a leoenv i ronmen t . T he p a t te rn i sa n i n c rea s ing s epa ra t ion o fh uman a dap ta t ion s f r om t h e c ons t ra in t s o f e nv i ronmen t . T he se a dap ta t ion sf o cu sed u pon t h e " w i ld h a rve s t ing" o fc oncen t ra ted r e sou rce s , e spec ia l ly r ed d ee r , i b ex , s a lmon , a nd ma r ine mo l lu sc s . T he se s pec ie s w e re h a rve s ted w i th i n c rea s ing e ff ic iency , d e sp i te m a jo r f l uc tua t ion s i nl o ca l c l ima te a nd s e a l e ve l . T he se c hange s i nw i ld f ood p rocu remen ti nm any a spec t sp r e sage t h e a dap ta t ion s l e ad ing t of ood p roduc t ion

i x

CHAPTER 1 I NTRODUCT ION

T h i s r e s ea r ch r e con s t ruc t s t h e p a leoenv i ronmen t o f t h e L owe r Magda len ian i n hab i tan t s o f C an tab r ian S pa in , t h e s t r ip o fl a nd c ompo sed o ft h e f o u r n o r the rn c oa s ta l r e g ion s a nd p rov ince s o f A s tu r ia s , C an tab r ia , V iz caya a nd G u ipüzcoa ( F igu re 1 . 1) . T he d a ta p r e sen t ed a r e t h e t e r r e s t r ia l m amma l ian m ic rof auna l r ema in s ( d ef ined a s s p ec ie s w i th a n a du l t b ody w e igh t l e s s t h an 5k g ) f r om E l J uyo , a n e x t r eme ly r i ch L owe r Magda len ian c ave s i t e . T he m ic rof au n a l d a ta f r om E l J uyo a nd o t he r U pp e r P a leo l i th ic s i t e s a r e c ompa red w i th t h e o t he r s ou rce s o f p a laeoenv i ronmen ta l d a ta . T he se i n c lude e nv i ronmen ta l d a ta d e r ived f r om l a rge t e r r e s t r ia l f a una , m a r ine f a una , s ed imen to logy , p a lyno logy , a nd c h rono log ica l s ynch ron iza t ion w i th c hange s i n w o r ld o cean l e ve l . E a ch s ou r ce p o s se s se s i n he r en tb i a se st h a tm u s t b e a c coun t ed f o r w hen e x t rapo la t ing p a s t e nv i ronmen ta l c ond i t ion s f r om t h em ( Kun iho lm 1 990 ) . T he a u tho r m a in t a in s t h a t i nb o th t h e r e so lu t ion o fi n f o rma t ion c on ta ined i n t h em a nd i nt h ep o ten t ia lt oa c commoda t et h eb i a se s i n cu r r ed i nt h e i r i n t roduc t ion t o as i t e , t e r r e s t r ia l m amma l ian m ic ro f a una a r et h em o s t a c cu ra t e i n f o rma t ion s ou r ce o n p a s t e nv i r onmen ta l c ond i t ion s . T he se b i a se s a r e b e s t e va lua ted a nd c on t ro l led f o ru s ing at a phonom ic a pp roa ch : d e te rm in ing t h e p a th t h e se o r gan i sm s t o ok f r om t h e b i o sphe r e t o t h e l i t ho s phe r e( Ef r emov 1 940 ) . A ne ven h i ghe rd eg r ee o fr e so lu t ion m ay b e a ch ieved u s ing t h e se mu l t ip le s ou r ce s t o ge the r ( Te s s i e re ta l . 1 993 ) .

w i th a b rup t ly r i s ing moun ta in s , C an tab r ian S pa in h a s m u l t i p l e e co log ica l z one s w i th in e a sy a c ce s s o f o ne a no the r , a l l ow ing f o r av a r ie ty o fa dap t ive r e spon se s a nd l o g i s t ic s t ra te g i e sb yi t sh uman o ccupan t s . I t s c u rren t b i ogeog raph ic p o s i t ion i sd i s t inc t f r om t h e r e m a inde r o f t h e I b e r ian P en in su la , w he r e ad r ie r , w a rme r Med i te r ranean c l ima t e n ow p r eva i l s . T he l o ca l s p ec t rum o f e xp lo i tab le s pec ie s i nC an tab r ian S pa in i sa nd w a s d i s t inc t i ve . D u r ing t h ec o lde rp ha se so fg l a c ia t ion , c l ima t ew a sa l so d i s t inc tf r om n e ighbo r ing F r ance , w he r et r ue t u nd ra a nd t u n d r a -dwe l l ing s pec ie s c ou ld b e f o und i na bundance ( r e indee r , a r c t ic f ox , s nowy o w l s , w o lve r ine , e t c . ) . C an tab r ian S pa in t h u s o f f e r ed ad i s t inc t s e t o ff o rag ing o p t ion s t oi t s i n hab i t a n t s . T he a r ea ' s k a r s t t e r ra in h a s a lowed e xce l len t p r e se r v a t ion o fa r chaeo log ica l m a te r ia l s , a l be i t i n as e lec t ive d epo s i t iona l s e t t ing : a lmo s ta l l o fi t sP a leo l i th ic o ccupa t ion l e ve l s h ave b een f ound i nl i me s tone c ave s . H ISTORY OF I NVEST IGAT ION P a leoenv i ronmen ta l r e sea r ch t r ace s i t s o r ig in s t o t h e v e ry b eg inn ing s o fg eo log ica l i n ve s t iga t ion i nt h e l a s t h a lf o ft h e 1 8 th c en tu ry . U n if o rm i ta r ian t h ough t tha t t h e s ame g eo l o g ica l p roce s se s v i s ib le t o day w e r e a l so i no pe ra t ion l o ng i n to t h e p a s t -pu shed t h e c on jec tu r ed a ge o f t h e E a r th f a r b eyond t h a t h e ld b y B ib l ica l s c ho la r s . T he s l ow d epo s i t ion o fs e d imen t s mu s t i n d ica t e t h a t l o ng s e quence s o fs e d imen t a ry r o ck t o ok m i l l ion s o fy ea r s t oa c cumu la t e , c ompa c t a nd e r ode . T hough p r eceded b y o t he r e a r ly g eo log i s t s , J ame s H u t ton i nh i s T he T heo ry o ft h e E a r th ( 1 795 ) f i r s tp ub l i shed t h e p r inc ip le o f u n if o rm i ta r ian i sm : t h e E a r th 's s e d imen ta ry r o ck s w e r e a ccumu la t ed b y t h e s ame p ro ce s se s o fs e d imen t d epo s i t ion v i s ib le i nc on tempo ra ry r i ve r s a nd s e a s . C ha r le s L ye l l l a te rc od if i ed t h e se i d ea s o fg eo log ica l p r oce s se s i nh i s P r inc ip le so fG eo logy ( 1 830 -33 ) .

P a leoenv i ronmen ta l r e sea r ch i sr e levan t t o mo r e t h an t h e n a tu ra l h i s to ry a nd e vo lu t ion o ft h e s ma l l m amma l s t h em s e lve s . I t s r e levance t oa r chaeo log ica l s t udy i si nt h e e s tab l i shmen t o fa n a r ena f o r h uman e co log i ca l i n te ra c t ion . O ne c anno t a s se s s h ow h uman r e spon se s t o t h e i r e nv i ronmen t c hang ed t h roughou t h om in id e vo lu t ion w i thou t e s tab l i sh ing w ha tt h o se e nv i ronmen t sw e r e . C hange si ne nv i ronmen ta l so m ay h ave p r omp ted c hange s i n h uman f o rag ing a c t iv i ty , f o od e xp lo i ta t ion t e chno logy a nd l a nd scape u s e a nd mu s tb e u nde r s tood b ef o r e c hange s i nt h e se a s pec t s m ay b e a s s igned t oo t he r c au se s . H uman s e pa ra t ion f r om t h e c on s t ra in t s o f e nv i ronmen t d u r ing t h e e nd o ft h e U ppe r P a leo l i th ic i sa l so r e levan tt ol a t e r m a jo r c hange s , i n c lud ing s e den t i sm , u rban i sm , s o c ia ls p ec ia l iza t ion a nd a g r icu l tu r e .

H u t ton a nd many o t he r n a scen t g eo log i s t s o ft h i s e a r ly p e r i od a l so n o ted t h a t g l ac ia l t r an spo r t w a s ap l au s ib le e xp la n a t ion f o rl a rge b ou lde r s d i sp la ced f a rf r om t h e i rp a r en tb ed r o ck ( g lac ia l e r ra t ic s ) . O the rr e la t ed p henomena t h a tb egged e xp lana t ion w e r e s t r ia t ion s o n e xpo sed r o ck s u rf a ce s a nd m o ra ine d epo s i t s f a r f r om e x tan t a l p ine g l ac ie r s . T he m a in c ompe t ing t h eo ry w a s t h a t b ou lde r s a nd o t he r d eb r i s m e l ted o u t o f d r if t ing i c ebe rg s i n au n ive r sa l d e luge ( i nc r ea s ing ly d i sa s soc ia ted t h roughou t t h i s p e r iod w i th t h e n o t ion o f a B ib l ica l f l ood ) . H o lde r s o ft h i s v i ewpo in t i n c luded W i l l iam B uck land , w ho p ub l i shed m any i n s igh tf u l o b se rva t ion s i n s uppo r t o ft h i s t h eo ry i nh i s R e l iqu iae D i lu v ianae i n 1 823 . E v idence f r om t h e s t udy o f moder n a l p ine g l a c ia t ion i n S w i tze r land , h oweve r , b egan t o c onv ince m u l t ip le i n ve s t i g a to r s t h a t t h e se p henomena r e su l t ed f r om t h e a c t ion o f mov ing g l ac ie r s .

C an tab r ian S pa in i s ap a r t icu la r ly r i ch r e g ion i n w h ich t o s t udy p a s t h uman a dap ta t ion s t o c hang ing e nv i ronmen t , w he r e h uman o ccupa t ion i s d ocumen t ed t h roughou t t h e L owe r , M idd le , a nd U ppe r P a leo l i th ic a nd Me so l i th ic p e r i o d s ( Gonz a lez E chega ray a nd F r eeman , 1 998 ; S t rau s 1 985 , 1 992 ) . T he r e g ion i sp hy s ica l ly c i r cum sc r ibed , b o rde r ed b y t h e B ay o fB i scay ( Can tab r ian S ea ) o n t h e n o r th , t h e P y r en e s t ot h e e a s ta nd t h e C an tab r ian C o rd i l le ra a l ong i t s s ou th e r n e dge . T h rough m uch o fi t s h i s to ry t h e se n a tu ra l o b s ta c l e s h ave p a r t ia l ly i s o la ted i tf r om a d ja cen t a r ea s . O ne m ay t h u s s t udy r e g iona l h uman e co log ica l a dap ta t ion s ( hun t ing /f o rag ing b ehav io r , l a nd scap e u s e , a nd r e sou r ce a c q u i s i t ion ) w i thou t p l a c ing n ece s sa r i ly a r t if i c ia l b o rde r s a r ound a n a r ea o f i n t e ra c t ion . D ue t o i t s c oa s ta l l o ca t ion

T he i d ea t h a t v a s t i c e s h ee t s c ove r ed E u rope i nt h e p a s t w a s l o ng i ng a in ing a c cep tance a mong t h e s c ien t i f ic c ommun i ty ( s umma r iz ed b yH a l lam 1 989 ) . I g na ce V ene tz r e po r t ed t oa

1

C an tabr ian S ea

S i te sD i scu s sed i nC hap ter L

c / a

0 . ) d c u A

•

. 0«

T . :

8? "

e s —

o • W▪° ' ,t X o 1 c " A r om ' a4 3 4 2 L I e s 0

e tZ

• ••

.-

' '

C m . .

t e

,.

. .

_ . ' 'a l

. ..

e z i. -e 0

6 a , — • " " 0 3 — m e t r 2

c l i

6

0 .r i e e MP In e N 0 . 1e i 1 . . t1 4e . t‘ •. •e 4 0 v

. r

O

e n

E i gi 7 1

.

9 3

s u

•

e v

o

e . Z . . t e 1 1

e n". •

4 0 1

e 4

> o

C . )

Ö ' V ( i .

e i s c e 4

=1 50 ,3 E E r : . . o ,-a2 e i .0 :• : . u . . . .nu o u g e 0 AM o 0 4 0

t

e ne al ' i oe n

.e ie n' e tt A

2

° A. c

0 3

; 7 . 'G2 , 4 0 .I N0 0 %

g a t ion s n a tu ra l ly f o cu sed u pon t h e s t ra t ig raph ic s ucce s s ion o fa r chaeo log ica lt e chno -comp lexe s a nd t h e p a r ie ta la r ta s so c i a ted w i th t h em . I tw a s a tt h i s s i te t h a t t h e v e rac i ty o fP a l e o l i th ic c ave a r t w a s e s tab l i shed a mong t h e s c ien t if ic c om -

S w i s s n a tu ra l h i s to ry s oc ie ty i n 1 82 1 t h e t h eo ry t h a t g l ac ie r s h ad o nce c ove red much mo re o f E u rope t h an p rev iou s ly p o s i ted . H e w a s t o e xpand u pon t h i s i d ea t h roughou t t h e d ecade , i n c lud ing t h e s cope o fh ow f a r t h e se a l p ine g l ac ie r s h ad s p read . T he n o t ion o f a n E i s ze i t ( I ce A ge ) w a s f i r s t

mun i ty ( Ca r ta i lha c 1 902 ) .

p ub l i shed b y t h e b o tan i s t K a r l S ch impe r i n 1 837 . T h i s t h e o ry w a s f u r the r e xpanded u pon a nd p opu la r ized t h rough t h e

I n t h e e a r ly h a lf o f t h e 2 0 th c en tu ry a r cha eo log ica l f o cu s r ema ined u pon e s tab l i sh ing t h e s ucce s s ion o ft h e s t one a ge c u l tu re s t ra t ig raph ic u n i t s o f We s te rn E u rope . Many i mpo r t a n ta nd f amou s s i te sw e re e xcava ted w i th v a ry ing d eg ree s o f

wo rk o f t wo n a tu ra l i s t s , J ean C ha rpen t ie r a nd ( mo re f a mou s ly ) L ou i s A ga s s iz . A ga s s iz p ub l i shed a s ub s tan t ia l w o rk i n 1 840 c onc lud ing t h a t av a s t i c e s hee t o nce c ove red E u rope , a nd t h rough much e f f o r th e s u cceeded i np e r sona l ly

p ub l ica t ion s ucce s s , s uch a st h e i mmen se d epo s i t s o fE l C a s t i l lo ( Cab re ra 1 984 ; O be rma ie r 1 924 ) . T h i s s i t e h a s y i e lded 1 8 m o fd epo s i t s w i th a r chaeo log ica l l e ve l s s p ann ing v i r tu -

c onv inc ing many c o l league s o fg l ac ia l t h eory . T h is t h eo ry w a s t og a in a ccep tance t h roughou t t h e e a r ly h a lf o ft h e 1 9"

a l ly t h e e n t i re p reh i s tory o f S pa in : A cheu lean , Mou s te r ian , A u r ignac ian , P e r igo rd ian , S o lu t rean , Magda len ian , A z i l ian , a nd N eo l i th ic . T yp ica l t o t h e e xcava t ion s t anda rd s o f t h e

c en tu ry , w i th p e rhap s c on suma t ing p roof i nt h e d i scove ry o f t h e G r een land i c es hee ti n1 852 . T he n o t ion h ad a r r ived t h a t t h e E a r th ' s e nv i ronmen t s w e re mu tab le , a nd i nt h e p a s t v a s t

d ay , t h e o n ly p a leoenv i ronmen ta l d a ta c o l lec ted w a s t h a t i n f e r red f r om t h e l a rge f a una l r ema in s a nd i n f requen t ly r e c ove red s ma l l f a una . R e sea rch f ocu s l a y u pon t h e t empo ra l s ucce s s ion o fs t one a nd o s seou s t oo l i n du s t r ie s . A t E l C a s t i l o t h e c o r rec t t empo ra l p l acemen t o f t h e S o lu t r ean i n dus t r ie s -be tween t h e A u r ignac ian a nd Magda len ian i n du s t r ie s w a s e s tab l i shed . P a lyno log ica l s t ud ie s w e re u nknown , a nd

s hee t so fi c ec ove r ed w ha t i sn ow E u rope . A l so n o tab le f r om t h i s p e r iod w e re s ome e a r ly p ub l i shed o b se rva t ion s c once rn ing s i te f o rma t ion a nd t a phonom ic p roce s se s . L ye 1 1( 1 830 -33 : 1 1 :219 -227 )d i scu s sed h ow a n ima l r ema in s c ou ld a ccumu la te i nt h e d epo s i t s o f c ave s a nd f i s s u r e sb yt h e i ra c t ing a sn a tu ra lp i tt r ap so ve rl o ng p e r iod s . I n t h i s m anne r , t h e se r ema in s o f t en b ecame a s soc ia ted w i th t ho se o fh uman s . B uck land ( 1823 :93 ) h ad p rev iou s ly n o ted t h a t " t he b one s o fv a r iou s b i rd s , o f mo le s , w a te r ra t s , m ice"

t h e r e t r ieva l o fs ma l l f a una l r ema ins w a s s ub jec t t ot h e l ow p robab i l i ty o f b e ing s po t ted d u r ing r a the r h a s ty e xcava t ion a nd n o t ap a r to fa ny s y s tema t ic a t temp ta tt h e i rc o l lec t ion .

h ad b een a ccumu la ted i nl i me s tone c ave s b y t h e " agency o f h awk s" , w h i le f i sh b one s h ad b een a ccumu la ted i nt h e s ame s i te s b y g u l l s . T h i s r e la t ion sh ip b e tween t h e t ype o f s ma l l f a una l r ema in s r e cove r ed a nd t h e p r eda to ry a gency o fa ccu mu la t ion w a st ob ecome c en t ra l i n2 0' c en tury i n ve s t iga t ions o ft a phonomy ( i nc lud ing t h ep re sen tw o rk ) .

P a lyno log ica l i n ves t iga t ion i nC an tab r ian S pa in b egan w i th t h e a na ly s i s o f t h e l o ng p a leo l i th ic s equence a t E l P endo ( Mou s te r ian , A u r ignac ian , G rave t t ian , Magda len ian a nd A z i l ian L eve l s ) . D ug o r ig ina l ly b y S au tuo la i n 1 878 a nd m any o t he r i n ve s t iga to r s i n s ub sequen t y ea r s ( i nc lud ing J e sü s C a rba l lo i n 1 926 ) , e x ten s ive r e -excava t ion s w e re c a r r i ed o u t i n 1 953 1957 , t hough p ub l ica t ion w a i ted u n t i l 1 980 ( Gonza lez E chega ray ) . A r lene L e ro i -Gou rhan ( 1980 ) p ro d uced a b r ief a na ly s i s o ft h e c hang ing p o l len c oun t s . T he p a lyno log ica l a na ly s is f o r t h e s i te o f I s tu r i tz w a s p ub l ished

A r cha eo log i s t s o ft h e 1 9 th c en tu ry b egan t o t i e h uman h i s t o ry i n to t h e se p a s t g l ac ia l e nv i ronmen t s ( s umma r ized b y G ray son 1 983 ) . T he a n t iqu i ty o fs t one t o o l s t h em se lve s w a s v e r if i ed b yt h e i r a s soc ia t ion s w i th e x t inc t f a una . Mo s t n o ta b l y , d e Mo r t i l le t ( 1883 ) p ub l i shed as t age s y s tem o fh uman d eve lopmen t . H i s s y s tem i sr e levan t t o t h e o r ig in s o f p a l e oenv i ronmen ta l t hough t i n i t s c o r r e la t ion o f s t one t oo l s w i th s uppo sed u n ique o rd om inan t f a una l a s soc ia t ion s . O u r S t one A g e a nce s to r s w e re t h eo r ized t oh ave p a s sed t h rough mu l t ip le s t age s : G r ea t C ave B ea r , E lephan t /Rh inoce ro s , R e ind ee r , A u roch s , a nd R ecen t . Wh i le l a te r r e jec ted , d e

much e a r l ie r ( Le ro i -Gou rhan 1 959 ) , t hough t h i s s i t e e x i s t s j u s t o u t s ide o fC an tab r ian S pa in i nF r ench B a sque c oun t ry . P a lyno log ica l a na ly s is b ecame s t anda rd p r ac t ice f o r s ub se q uen te xcava t ion s . T he se i n c lude s uch s i te s a sO te ro , Mo r in , a nd L a R ie ra ( A r l . L e ro i -Gou rhan 1 966 , 1 978 a nd 1 986 , r e spec t ive ly ) ; T i to B us t i l lo , R a scar io , C huf in , a nd E ra l la ( Boye r -K le in 1 976 , 1 981 , 1 984 , 1 985 ) ; L o s A zu le s ( Lopez 1 981) ; E ka in a nd Ama lda ( Dup r é 1 984 , 1 990 ) , a nd

Mor t i l le t ' s s cheme d i d l i nk S tone A ge p eop le s w i th f a una t o ta l ly o r l o ca l ly e x t inc t i nc on tempo ra ry t empe ra te E u rope .

L eze tx ik i , L abeko a nd U r t iaga ( Sanchez G or i i 1 993 ) .

P eop le w e r e p o s i ted t o h ave p a s sed t h rough a R e indee r S t age , a dap t ing t o au n ique a nd much mo r e c l ima t ica l ly r i g -

e nv i ronmen ta lc ond i t ion sh ad b een e s tab l i shed .

T h i s c once rn f o r p a lyno log ica l a na ly s i s i so ne o f as heaf o f n ew i n t erp re t ive t e chn ique s a pp l ied t op a leo l i th ic i n ve s t iga t i on . A na ly s i so fs ed imen t sa nd t h e i rc hange sb etween l e ve l s a s a n i n d ica t ion o f p a leoenv i ronmen t h a s a l so b een u nde r -

S e r iou s P a leo l i th ic i n ve s t iga t ion i n C an tab r ian S pa in c om m enced a s e a r ly a s 1 872 w i th t h e e xcava t ion s b y E . d e l a P r ed ra ja i nt h e s i te o fC ova le jo s , w h ich y i e lded a s semb lage s

t a ken a tl e a s t s i nce t h e 1 953 -57 r e -excava t ion s a tE l P endo ( Bu tze r 1 980 ) . I n teg ra ted r e sea rch o ft h i s k i nd h a s g rown d u r ing t h e p a s t t h ree d ecade s w i th t h e a dven t o fc on t ro l led , moder n e xcava t ion s b eg inn ing w i th t h e r e -excava t ion o f

( l a te r i d en t if i ed a s ) Mou s te r ian ( Gonza lez E chega ray a nd

C ueva Mo r in ( Gonza lez E chega ray a nd F r eeman

F reeman , 1 998 ) . T he f i r s t e xp lo ra t ion s o f A l tam i ra w e re c a r r ied o u t i n 1 876 b y Ma rce l ino S anz d e S au tuo la , al o ca l a ma teu r p r eh i s to r ian . T h i s w o r ld renowned c ave s i te h a s

1 973 , 1 978 ) d u r ing t h e l a te 1 960s . N ew e xcava t ion s a tt h e s i te o fL a R ie ra ( C la rk a nd R icha rd s 1 978 ; S t rau s a nd C la rk

o rou s e nv i ronmen ta l s i tua t ion . T he i d ea , h oweve r h e s i tan t , t h a t h uman c u l tu r e w a s mu tab le a nd a dap tab le t o c hang ing

1 971 ,

1 978 , 1 986 ) w e re u nder taken w i th t h e s pec if i c p ro jec t f o cu s o f p a leoeco logy . O ngo ing i n ve s t iga t ion s a t E l C a s t i l lo ( Cab re ra 1 984 ; B i schof fe ta l . 1 991) a nd E l P endo a r e u n d er taken w i th t h e s pec if ic g oa l s o f mo re p r ec i se e xcava t ion

y i e lded o ccupa t ion l a ye r s f r om t h e S o lu t rean a nd Mag d a len ian p e r iod s , a s w e l l a s magn if i cen t p a r ie ta l a r t d a t ing f r om t h e l a t te r p e r iod ( F reeman 1 987 ) . T he se e a r ly i n ve s t i 3

o fr emnan ts t ra t ig raphy a nd t og a the re v idence m i ssed d u r ing t h e e a r l ie r , l e s s c on t ro l led d i g s . S uch mode rn t e chn iques a r e

l a rge f a una ( Ave ry 1 982a ; B ou r l ie re 1 975 ; D e l ibe s d eC a s t ro 1 985 ; F a e ta L 1 982 ) . C hange s i nl a t i tud ina l ( o r c l i se r ia l -

a pp l ied t o t h e moder n e xcava t ion s o f s uch s i te s a s T i to B u s t i l lo ( Mou r e R oman i l lo 1 975 ) , E r ra l la ( A l tuna 1 985a ) ,

G raham 1 979 ) , a l t i tud ina l ( Mo reno a nd B a rbo sa 1 992 ) , d e g r ee o fp rec ip i ta t ion , a nd v ege ta t ion a s soc ia t ive makeup o f s ma l l mamma l c ommun i t ie s a r e n o ted . T he se c hang ing m i -

Ama lda ( A l tuna e t a l . 1 990 ) , E ka in ( A l tuna a nd Me r ino 1 984 ) , a nd R a sca r lo ( Gonza lez E chega ray a nd B a rand ia rän 1 981) . Much c on tempo ra ry r e sea rch c en te r s u pon u t i l iza t ion o fl a nd scape , s i te l o ca t ion , a nd s i te t e r r i to r ie s w i th r ega rd t o

c r of auna l p a rame te r s i n d ica te t h e c hang ing L a te P l e i s tocene e nv i ronmen t .

f ood p rocu remen t ( Ba i ley 1 983 ; B a i ley a nd D av id son 1 983 ;

When a ccumu la ted b y g ene ra l is t p reda to r s a s i si n d ica ted a t

C la rk 1 985 ) . I nve s t iga t ion o fd eco ra t ive mo t if s o n b one i m p l emen t s h a s i n d ica t ed t h e p o s s ib le s ea sona l a gg rega t ion o f s oc ia lg roup sa tc e r ta in l o c i ( A l tam i ra a nd C ue to d e l aM ina ) ,

E lJ uyo ( s ome o w l s pec ies i np a r t icu la r ) , m ic rof auna l a s sem b l age s c an g i ve al e ss n ebu lou s p i c tu re o fl o ca l h ab i ta t s a nd t h e i r r e la t ive p ropo r t ion s ( And rew s

1 990 ; A ve ry 1 982a ,

1 982b , 1 987 , 1 992 ; B ra in 1 974 ; B ra in a nd B ra in 1 977 ; L o v a n ie t a L 1 976 ; T hacke ray 1 987 ; T ha cke ray a nd A ve ry 1 990 ) . B ia se s d ue t op rey s e lec t iv i ty b y p r eda to r s o rp r ef e r -

t hu s d i rec t ly t y ing t oge the r t h e e conom ic a c t iv i t ie s o f mu l t i p l e s i te s ( Conkey 1 978 , 1 980 ) . T he t r end i n c on tempo ra ry C an tab r ian p a leo l i th ic i n ve s t iga t ion s h a s t hu s b een i n teg ra t i ve r e sea rch c omb in ing mu l t ip le s pec ia l t ie s , w i th s i te s s t ud -

e nce f o r c e r ta in h un t ing h ab i ta t s a r e u nde r s tood b y c ompa r i s on o ff o s s i l a s semb lage s w i th moder n o ne s a c cumu la ted b y p r eda to r s i n av a r ie ty o f e co log ica l z one s . P a leoenv i ron m en ta l r e con s t ruc t ion t h e refo r e c ome s f r om c ompa r i son b e tween m ic rof auna l s i gna ls d e r ived f r om g i ven e nv i ron m en t s ( Ave ry 1 993 ) , a nd n o t f r om t h e a s sump t ion t h a t r e la t i ve s pec ie s p ropor t ions d i rec t ly p red ic t r e la t ive h ab i ta t p r o p or t ion s . Al a rge b ody o fr e sea r ch f r om h ab i ta t s t h roughou t t h e H o la rc t ic e x i s t s f o r c ompa r i son s o f t h i s t y pe ( Br une t L ecom te a nd D e l ibe s 1 984 ; B uck ley a nd G o ld sm i th 1 975 ; C ampo s 1 978 ; D e l ibe s e ta L 1 984 ; G lue 1 967 , 1 970 , 1 974 , 1 975 ; G o sz czyn sk i 1 974 ; J ak s ic e ta L 1 982 ; P i te lka e ta l . 1 955 ; S ou ther n 1 954 ) . T he p re sen t r e sea r ch , h oweve r , p r e s e n t s an ew m e thod f o r c a lcu la t ion d i r ec t ly f r om t h e a r c haeo log ica lr ema in s ( Chap te r4 ) .

i e d i nt h ec on tex to ft h e i rr e g iona ls e t t ing a nd i n te rac t ion s . Mo s t r e sea rch p r io r t ot h e p a s t t wo d ecade s t e nded t o l ump m ic rof auna w i th l a rge f a una , t h e l a t te r o fw h ich w a s g iven t h e g r ea t e rp r io r i ty o fa na ly s i s a nd i n te rp re ta t ion . T he w o rk o f J e sü s A l tuna ( 1972 , 1 976 , 1 978 , 1 986 ; A l tuna a nd Ma r i ezku r rena 1 980 , 1 985 ) s pan s t h i s s epa ra t ion p e r iod o fl a rge a nd s ma l l f a una l a na ly se s . A s ad i rec tr e su l t o f mo re r ef ined e xcava t ion t e chn ique s , r e sea r ch s pec if ica l ly c once rn ing m amma l ian m ic rof auna ( a sw e l l a so t he rs ma l l f auna s uch a s f i sh , r ep t i le s , a mph ib ian s a nd b i rd s ) b ecome s p rac t ica l . Wh i le s ome o f t h e se f a una l e l emen t s w e re r e cove red e ven u nde r e a r l ie r , l e s s r i go rou s e xcava t ion r eg imes ( Cab r e ra 1 984 ) , t h e i r t o ta l s w e r e f a r s ma l le r t h an t h e mo re a r chae o l og ica l ly v i s ib le a nd e conom ica l ly i mpo r tan t l a rge f a una ( A l tuna 1 972 ) . F ine -me sh s c reen ing o fs ed imen t s i s am in i mum r equ i remen t f o rt h e i r r e cove ry . S ome mode rn e xcava t i on s h ave y i e lded s i gn if ican t ly l a rge m ic rof auna l s amp le s , n o tab ly E ka in ( n =1 , 008 i d en t if ied r ema in s [ Zaba la 1 984 ] ) , Ama lda ( n =2 , 346 [ P emän 1 990 ] ) , a nd E ra l la ( n =1 5 ,710 [ P emän 1 985 ] ) . Moder ne xcava t ion t e chn iques a r e n o g ua r a n tee o f s ub s tan t ia l m ic rof auna l s amp le s , a s a t L a R ie ra w he re mode rn r e -excava t ion o fr emnan ts t ra t ig raphy y i e lded as ma l l m ic rof auna l s amp le ( n =1 52 ) ( A l tuna 1 986 ) . Mod er n r e covery t e chn ique s h ave t h e a dd i t iona l b enef i t o f p ro -

T he s e cond r e sea rch p rob lem i s t h e c ompa r i son o f t h i s s ou rce o f e nv i ronmen ta l d a ta w i th o t he r s ou r ce s a l r eady a va i lab le f o r E l J uyo a nd o the r s i t es . S ed imen to log i ca l a na ly s i s s ugge s t s i n c rea s ing i n s tab i l i ty a nd a n e ven tua l r e t u rn t oc o lde rc ond i t ions d u r ing t h e t ime o fd epo s i t ion o ft h e u ppe r h ab i ta t ion l e ve ls ( F reeman e ta L 1 988 :11) . Mea su r e m en t o ft h e d eg r ee o fc ryoc la s t ic s pa l l ing c an g i ve ar e la t ive i n d ica t ion o fa n i n c rea se i nc o ld c ond i t ion s . A ny a c tua l p a l e oenv i ronmen ta l r e con s t ruc t ion , h oweve r , i s b a sed u pon e x t rapo la t ion f r om p a leo tempe ra tu re r e g ime s a nd a na logy w i th p re sen t -day c o ld c l ima te s ( Bu tze r 1 986 ) . A dd i t iona l ly ,

d uc ing u np recenden ted l a rge s amp le s o f s t one a nd o s seou s m ic ro too l s s u ch a s m ic rob lade s , f i ne p o in t s , a nd f r agmen t s o fb one n eed le s( F reeman e ta l . 1 998 ) .

h uman a c t iv i ty m ay p o ten t ia l ly a l te r s ed imen t d epo s i t ion i n w ay s t h a t m ay b e m i s taken f o r c hange s i n e nv i ronmen t . T he se i n c lude c hanges i n f l uv ia l d epo s i t ion d ue t o t h e i n -

RESEARCH OBJECT IVES T h ree p rob lem s a r e a dd re s sed b y t h i s s t udy o f p a leoenv i -

c r ea sed r unof f a lowed b y t r amp led v eg e ta t ion o u t s ide a n o ccupa t ion s i t e , a nd c hange s t o t h e f r eez e / thaw r eg ime o fa c ave o r r ock she l te r c e i l ing b y h uman -cau sed t empe ra t u r e

r onmen t . O ne i st h e r e con s t ruc t ion o f t h e L owe r Mag d a len ian h ab i ta t s u r round ing E l J uyo a nd b y e x ten s ion t h e

r e gu la t ion : e r ec t ion o fb a r r ie r s t oa i rf l ow a nd u se o ff i r e i na

h ab i ta t o ft h e n ume rou s o t he r s i te s e x is t ing i nt h e s ame t ime r ange a nd c oa s ta l l ow land e nv i ronmen ta l z one . T he se s i te s i n c lude t h e Magda len ian o ccupa t ion s o f A l tam i ra ( A l tuna

c ave . S ed imen to logy t hu s i ss i gn if ican t ly l i m i ted i ni t s p o t e n t ia lf o ri n f e r r ing c hange s i nc l ima te .

a nd S t rau s 1 976 ; S t rau s 1 976 -77 ) , L a R ie ra ( S t rau s a nd C la rk 1 986 ) , E l C a s t i l lo ( Cab re ra 1 984 ) , T i to B us t i l lo ( Mou re R oman i l lo 1 975 ; Mou re R oman i l lo a nd C ano H e r -

t h e P l e i s tocene ( Oxygen I s o tope S t age 2 ) , w i th s e a l e ve l s ,

O ccupa t ion a tE l J uyo o ccu r red d u r ing t h e c l o s ing s t age s o f

r e ra 1 976 , 1 978 , 1 979 ) , C ueva Mo r in ( A l tuna 1 978 ) , a nd

d ay ' s t empe ra te a nd h um id c l ima te ( CL IMAP 1 976 ; Mc In -

E ka in ( Zaba la 1 984 ) . B y f u r the r e x t rapo la t ion , t h e e co log i c a l s e t t ing o fu p land s i t e s s uch a sR a sca i lo may b e s u rm ised .

t y re e ta l . 1 976 ; R udd iman a nd Mc In ty r e 1 976 ; S ha ck le ton

o cean c i r cu la t ion , a nd t empe ra tu re s v e ry d i f f e r en t f r om t o -

a nd O pdyke 1 976 ; v an D onk 1 976 ) . G loba l s e a l e ve l s a tt h e l a s t g l ac ia l m ax imum w e re a tl e a s t 8 5 m l owe rt han p r e sen t -

T he b a s ic p a rame te r s o ft h i s r e con s t ruc t ion a r e t h e v a ry ing c omponen t sa nd p ropo r t ion s o ft e r re s t r ia l mamma l ian m ic ro -

d ay l e ve l s ( CL IMAP 1 976 ) , w i th o t he r e s t ima te s p l ac ing s e a l e ve l f r om 9 0 mt o 1 30 m l owe rt h an p r e s en t( f o rs umma r ie s s e e B a i ley 1 983 ; B u tze r 1 983) . E v idence d e r ived f r om d eep

f a una . T he se t a xa a r e r e p re sen ta t ive o fc e r ta in m ic rohab i ta t s a nd t e nd t ob e f a r mo r e s pec ia l ized i nt h e i r c ho ice s t h an a r e 4

c a to r s p ec ie s , h oweve r , a nd c omp lemen t o t he r s ou r ce s o f d a ta . O ne mu s t n o te t h a t t h e p r inc ipa l p r ey s p ec ie s a tE l J uyo a nd mo s t o t he r C an tab r ian s i t e s , r e d d ee r ( C er vu s e l aphu s ) , c an t h r ive a c ro s s ar a nge o fe nv i ronmen t st h rough o u tt h e H o la r c t ic a nd i si n d ica t ive o fn o o ne h ab i ta t . L a rge r s p ec ie s o fm ic rof auna ( s uch a s l a gomo rph s o rs ma l l m u s te l i d s ) a l so m ay h ave b een f ood o r f u r r e sou r ce s f o r h uman s a nd mu s t b e c r i t ica l ly e va lua t ed f o r e v id ence o ft h e i r i n c lu s i on i nt h es i t eh av ing b een c u l tu r a l ly a f f ec ted .

s e ac o r e f o ram in if e ra s ugge s t st h a tt h eN o r th A t lan t ic s u r f a ce o cean w a t e rt empe r a tu r e s a t4 2° N o r th l a t i tude w e r e a r ound 1 0° C c o lde ro na ve rage c a . 1 8 ,000 B .P .t h an a tp r e sen t , i n d i c a t ing p o la r w a t e r c ond i t ion s ( Mc In ty r e e ta L 1 976 ) . T he l owe s t e l eva t ion o ft e rm ina l mo ra ine s l e f tb y a l p ine g l a c ie r s d u r ing t h e l a s tg l a c ia l m ax imum i nt h eC an t ab r ian C o rd i l le ra i s 50 m a bove s e a l e ve l ( C la rk 1 981 : 7 ) . A i rt empe ra tu r e s a tg l a c ia l m ax imum f o rt h e c oa s t n ea r E lJ uyo h ad aF eb ru a r y a ve rage o f a r ound 1 .6 ° C a nd a n A ugu s t a ve rage o f 1 0 .4 ° C . T he n ea r e s t m od e rn a na log f o rt h e se w ea the r c on d i t ion s i st h e t r ee le s s c oa s t o f I c e land a t 6 6° N l a t i tude ( Bu tze r 1 986 :215 ) . D u r ing t h e h a r she s t c l ima t ic c ond i t ion s o fE l J uyo ' s l a t e r o ccupa t ion , t h e t e r ra in a r ound t h e s i t e h ad

A f u r the r s ou rce o f e v idence u t i l iz ed i n c on junc t ion w i th m ic rof auna a r e t h e ma c robo tan ica l r ema in s r e cove r ed f r om E l J uyo . E xce l len t p r e serva t ion ( d ue i n l a rge p a r t t o t h e s e a l ing o ft h e d epo s i t s b y f l ow s tone s ho r t ly a f t e r o ccupa t ion c ea sed )c omb ined w i th c omp r ehen s ive r e t r ieva lp r a c t ice sh a s a lowed u np receden ted r e cove ry o f c a rbon iz ed a nd u nca r b on iz ed p l an t p a r t s a nd s e ed s ( C rowe 1 987 ; F r eeman e ta l . 1 988 ; F r eeman e t a l . 1 998 ; I b af i ez -Angu lo 1 99 1 ; S t e t t le r 1 995 ) . T h i s b o tan ica l d a ta d oe s s u f f e r f r om s ome o f t h e s ame b i a se s a s p o l len f r om t h e s i t e : p a s sage t h rough ac u l t u ra l f i l te r i nt h e p r e sence a nd r e la t ive a bundance o ft a xa , d i f f e r en t ia l p o ten t ia l f o r t r an spo r t , a nd d i f f e r en t ia l s u scep t i b i l i ty t o d e s t ruc t ion . L oca l p l an t t a xa r e p r e sen ted t h rough b o tan ica l r ema in s r e conf i rm t h e l o ca l o r ig in o fp o l len i n t ro d uced t ot h e s i te a nd a s i n d ica to r s p ec ie s i nt h em se lve s . I n t h i sr o le t h ey c omp l imen tt h em ic rof auna ld a ta f r om t h es i t e .

m uch l e s s f o r e s t c ove r a nd much mo r e o pen g r a s s land t h an m ode rn c l ima t ew ou ld a low . O ne m ay i n f e rt h a tt h e c l ima te w a s s i gn if i can t ly c o lde rt h an t o day i n g ene ra l t h roughou t E u rope d u r ing t h e l a te r ( c a . 1 4 ,000 B .P . ) o ccupa t ion o f E l J uyo , w i th a me l io ra t ing c l i m a t e c ond i t ion s f r om t h e g l ac ia l m ax imum c a . 2 2 ,000 1 8 ,000 B .P . ( S t rau s 1 99 1b ) . L oca lc ond i t ion s , h oweve r ,m ay h ave v a r ied g r ea t ly f r om r e g ion t or e g ion . S uch i sp a r t icu l a r ly t r ue f o r C an tab r ian S pa in , w h ich o we s i nl a rge p a r t i t s m oder n t empe ra te c l ima t e t o i t s c oa s t a l p r ox im i ty a nd t h e w a rm w a t e r s o ft h e G u lf S t ream . S i n ce t h e s ou thwa rd a d v ance o ft h e p o la r i c e c ap i n to t h e N o r th A t lan t i c m ay h ave d e f lec t ed t h e G u lf S t r eam a brup t ly s o u thwa rd ( Rudd iman a nd Mc In ty r e 1 976 ) , t h i s s ou rce o f l o ca l c l ima t ic w a rm ing m ay h ave b een s w i t ched o n o ro f fo ve rr e la t ive ly s ho r tp e r i o d s . C hange s i n l o ca l c l ima te m ay h ave b een s im i la r ly a b rup t .

T he f i na l p rob lem u nde r c on s ide ra t ion i st h e n a tu r e o ft h e c hang ing a dap ta t ion s o f t h e h uman g roup s i n hab i t ing C an tab r ian S pa in a tt h e e nd o ft h e P l e i s tocene i nr e la t ion t o t h e i r e nv i ronmen t . B road c hange s i n f ood p rocu r emen t s t ra tegy o ve r t i me a r e e v iden t i nt h e a r cha eo log ica l l e ve l s a l r eady e xcava t ed . Mou s te r ian a nd E a r ly U ppe r P a leo l i th ic l e ve l s a r e c ha ra c t e r iz ed b y o ppo r tun i s t ic h un t ing a nd /o r s c aveng ing o f l e s s d ange rou s l a rge m amma l t a xa , s u ch a s h o r se ( Equu s c aba l lu s ) , l a rge b ov id s a nd r e d d ee r ( F r eeman 1 973 , 1 975 , 1 98 1 ; S t rau s 1 977 , 1 992 , 1 993 ) . B eg inn ing w i th t h e L a te U ppe r P a leo l i th ic ( So lu t r ean , Magda len ian , a nd A z i l ian c u l tu r e s t ra t ig raph ic u n i t s ) a nd a c ce le ra t ing i n to Me so l i th ic ( A s tu r ian )t ime s , f ood p r ocu r emen t i nC an tab r ian S pa in i n t en s if ied i nt h e e xp lo i ta t ion o fk ey s p ec ie s , p r ima r i ly r e dd ee r , A t lan t ic s a lmon ( Sa lmo s a la r ) a nd m a r ine mo l lu sc s ( Pa te l la v u lga ta a nd L i t to r ina l i t to rea m o s tc ommon ly ) . S i t e l o ca t ion /f unc t ion b egan t od i ve r s if y i nw ay s t h a t s ugge s t t h e e xp lo i ta t ion o f o t he r mo re a r ea l ly r e s t r ic ted / spec ia l iz ed r e s ou r ce s , s u ch a s i b ex ( Cap ra p y rena i ca ) h un t ing a tt h e u p l a nd s i t e o fR a scaf io ( Gonza lez E chega ray a nd B a rand ia rän 1 98 1) . T he L owe r Magda len ian l e ve l s o fE l J uyo t h u s s p an

P a lyno log ica l a na ly s i s h a s b een u nde r taken f o r E l J uyo a s w e l l ( Boye r -K le in a nd A r l . L e ro i -Gou rhan 1 987 ; A r l . L e ro i G ou rhan 1 994 ) . T he p r ob lem s o fp o l len t a phonomy , d i f f e r e n t ia lp roduc t ion a nd b uoyancy h ave i n c r ea s ing ly d r awn i n to q ue s t ion t h ef a ce v a lue o fa ny p o l len d i ag ram ( S anchez G of i i 1 993) . T he p ropen s i ty o fs ome w ind -po l l ina t ed t a xa ( s uch a s P inu s ) t o p r oduce a bundan t a nd f a r t r ave l ing p o l len w i l l c au se t h e i ro ve r r ep r e sen ta t ion i n ap o l len p r of i le ( F aeg r i a nd I v e r sen 1 989 ; P r en t ice 1 986 ) . P o l len f r om g r ound p l an t s w i l l a l so b e i n t roduced t o as i te o nt h eb od ie s o fh uman sa nd a n ima l s , c au s ing o ve r r ep r e sen ta t ion o f s h rubby a nd h e rba c eou s f o rm s . F o l iage t r an spo r ted i n to t h e s i t e f o r f o od , b ed d i ng , m ed ic ine , o r a sa n i n c iden ta l i n c lu s ion m ay a l so c on c en t ra t e t h e p o l len o fs e lec ted s pec ie s i np ropo r t ion s u n l ike t h o se i nt h e l o ca l s u r round ing s . R e s ea r ch h a s s hown t h a t p o l len i sr ewo rked a nd t r an spo r ted b y w a t e r ( Dav i s 1 968 ; T rave r se 1 990 ) a nd t h a t c on sequen t ly s o r t ing b y s i z e m ay o ccu r( Fa l l 1 987 ; G oodw in 1 988 ; H o lme s 1 990 ) . P o l len a nd s po r e s o fd i f f e r en t s p ec ie s a r e a l so d i f f e r en t ia l ly s u scep t ib le t od e s t ru c t ion b ym ic rob ia la t ta ck , o x ida t ion , a nd m echan ica l f o r ce s o fc o r ro s ion ( Hav inga 1 967 ) . Wh i le b r oad ly s ugge s t i ve o fp a s t e nv i ronmen ta l s e t t ing s a nd g r o s s c hange s i nc l i m a t ic r e g ime s , p a lyno logy a l one c anno tg i ve ac omp le t e a nd c onv inc ing p i c tu r e o f t h e p a leoenv i ronmen t o f C an tab r ian S pa in ,a ny m o r et h an c an s e d imen to logy .

af a sc ina t ing p e r iod w he r e i n t en s if i ca t ion o fs ome a s pec t s o f f o od p rocu r emen t ( s laugh te r o fw ho le h e rd s o fr e d d ee r , i n c r ea sed u s eo fm a r ine r e sou rce s , b r oaden ing o ft h e f o od b a se t o i n c lude p r ev iou s ly u nde ru t i l iz ed s p ec ie s ) o ccu r r ed . A n ece s sa ry c omponen ti nu nde r s tand ing t h e se c hange s i nf o od p rocu r emen t s t ra t egy i st h e r e con s t ruc t ion o ft h e i r c hang ing e nv i ronmen ta ls e t t ing . ARCHAEOLOG ICAL CONTEXT T he E l J uyo m a t e r ia l s f i t w i th in t h e o ve ra l l c on t ex t o f t h e L a te U ppe r P a leo l i th ic a s r e p r e sen t ed s e qu en t ia l ly b y t h e S o lu t r ean ( 20 ,500 t o 1 6 ,000 B .P . ) , L owe r Magda len ian ( 1 6 ,500 t o 1 3 ,500 B .P . ) , U ppe r Magda len ian ( 1 3 ,500 t o 1 ,000 B .P . ) , a nd A z i l ian ( 1 1 ,000 t o9 , 000 B .P . ) t o o l i n du s -

T he l a rge f a una o fE lJ uyo a nd o t he r Magda len ian s i t e sh ave b een a f f ec t ed b y c u l tu ra l b i a se s i nt h e i r s e lec t ive h a rve s t , m ak ing t h e i rr e la t ive p r opo r t ion s g ene ra l ly u n sa t i sf ac to ry f o r p a leoenv i ronmen ta l r e con s t ruc t ion . T h ey a r e u s ef u l a si n d i 5

t r ie s ( f o l low ing S t rau s 1 992 ) . S ome r e sea r che r s r e cogn ize f u r the r s ubd iv i s ion s s uch a s aM idd le Magda len ian , w h ich c o r re spond s t o Magda len ian I V o f F rance ( ma rked b y t h e p r e sence o f" p ro to -ha rpoon s " ) . T h i s p ha se h a s b een d ema r c a ted a t s i te s i n c lud ing L a P a loma ( Ma r t inez a nd C hapa

T he S o lu t rean i ss ucceeded w i th s ome t empo ra l o ve r lap b y t h e L owe r Magda len ian , a ga in a bundan t ly r e p re sen ted a mong C an tab r ian s i te s . T o ta l s i te n umbe r s v a ry a cco rd ing t o h ow r e sea r che r s c l a s s if y e qu ivoca l a s semb lage s . U t r i l la ( 1981) c omp i le s 1 8 d ef in i te L owe r Magda len ian s i te s i n C an tab r ia S pa in , w i th m any mo re Magda len ian s i t e s i n de -

1 980 ) a nd L a s C a lda s ( Co r chön 1 981) . O the r f i ne r s ubd iv i s i on s a k in t ot ho se c ommon ly d eno ted i nF rench L a te Mag -

t e rm ina te a st oL owe r o r U ppe r p ha se a nd many s h e d e s ig -

d a len ian s i te s b a sed u pon c hange s i nh a rpoon s t y le ( Bo rde s

n a te s M idd le Magda len ian . S t rau s ( 1992 :129 ) u pda te s t h i s f i gu re t oa sh i gh a s 5 2 , b u t n o te s t h e p robab le i n f la t ion d ue

1 968 :16 1-166 ) h ave b een a pp l ied w i th l i m i ted s ucce s s t o C an tab r ian S pa in ( s ee S t rau s 1 992 :135 138 f o r d i scu s s ion ) .

t o t h e i n c lu s ion o f e qu ivoca l Magda len ian p ha se s i t e s a nd

F ine r g rada t ion s , w he the r v a l id o r i n va l id a s a pp l ied t o C an tab r ian S pa in , a r en o tr e levan tt ot h ep r e sen tt op ic .

d oe s n o t r e cogn ize ad i s t inc t M idd le Magda len ian p ha se i n C an tab r ian S pa in . L owe r Magda len ian s i tes i n c lude E l C a s t i l lo , A l tam i ra , L a R ie ra , R a sca l lo , L a P a loma , C ue to d e l a

H uman h ab i ta t ion i nt h i s r eg ion d a te s f r om t h eA cheu lean a s r e p re sen ted b yt h e b a sa l d epo s i t s a tE l C a s t i l lo , a n i mpo r tan t

M ina , C ueva d e l a L lo se ta ( J o rdä 1 958 ) , a nd E l J uyo ( Ba rand ia rän e ta l . 1 987 ; F reeman a nd G onza lez E chega ray

c ave s i te a l so y i e ld ing Mou s te r ian , A u r igna c ian , G rave t t ian , S o lu t rean , L owe ra nd Uppe r Magda len ian ,A z i l ian , a nd p o s t N eo l i th ic l e ve l s , a sw e l l a sa bundan tc ave a r t ( B i schof fe ta l . 1 99 1 ; C ab r e ra 1 984 ; Mou re R oman i l lo e ta l . 1 996 ) . O the r A cheu lean o ccu r r ence s c on s i s t o n ly o f ( mo s t ly e qu ivoca l ) o pen a i r s i te s , t hu s mak ing t h is e a r ly t o o l c omp lex p oo r ly r ep re sen ted a tt h e p re sen t t ime . T he Mou s te r ian i sb e t te r r ep re sen ted i n C an tab r ian S pa in , w i th l e ve l s e xcava ted a t s i te s s uch a s E l C a s t i l lo ( Cab re ra 1 984 ) , C ueva Mo r in ( Gonza lez E chega ray a nd F reeman 1 978 ) , a nd E l P endo ( Gonza lez E chega ray 1 980 )a mong o t he r s .

1 981 , 1 982 , 1 995 ; F reeman e ta l . 1 988 ; J an ssen s a nd G on z ä lez E chega ray 1 958 ; K rupa 1 994 ; P ok ines 1 993 , 1 997 ; S t e t t le r 1 995 ) . T he U ppe r Magda len ian i se qua l ly w e l l r ep r e sen ted a ts uch s i te s a s E l C a s t i l lo , L a R ie ra , R a sca f io , L a P a loma , a nd T i to B u s t i l lo ( Mou re R oman i l lo 1 975 ; Mou re R oman i l lo a nd C ano H e r r e ra 1 976 , 1 978 , 1 979 ) . T he A z i l i a n f o l low s , w i th l e ve l s a tL o s A zu le s , L a R ie ra , R a scaf io , a nd L aP a loma a mong o t he rs i te s . Wh i le e ach o f t h e se a r chaeo log ica l t oo l c omp lexe s d e r ive s i t sn ame f r om al o ca l i ty i nF rance ( So lu t rd , L a Made le ine , S t . A cheu l , e t c . ) , t h e se p ha se s a s r ep re sen ted i n C an tab r ian S pa in d o n o t c o r re la t e p r ec i se ly w i th e x tan t F r ench -ba sed

E a r ly U ppe r P a leo l i th ic s i te s i nC an tab r ian S pa in t e nd t ob e mo r e n ume rou s a nd may i n d ica te ac on t inu ing t r end o fs l ow p opu la t ion i n c rea se ( S t rau s 1 992 ) . Wh i le C ha te lpe r ron ian l e ve l s h ave b een f ound a t o n ly a f ew s i t e s , a s a t C ueva Mo r in , E l P endo a nd p o s s ib ly E ka in ( A l tuna a nd Me r ino 1 984 ) , A u r ignac ian l e ve l s o ccu r a t s eve ra l s i te s , n o tab ly C ue to d e l aM ina ( Vega d e l S e l la 1 9 16 ) , R a scaf io ( Gonza lez E chega ray a nd B a rand ia rän 1 981) a nd a ga in a tE l C a s t i l lo , E l P endo , a nd C ueva Mo r in . T he E a r ly U ppe r P a leo l i th ic i s c on t inued b y U ppe r P e r igo rd ian /G rave t t ian l e ve l s a t s i t e s s uch a sE l C a s t i l lo , Ama lda ( A l tuna e ta l . 1 990 ) a nd C ueva Mo r in .

c h rono log ie s . T he S o lu t rean i so f t en d iv ided i n to p ha se s b a sed u pon t h e s ucce s s ive a ppea rance o ff o l ia te p o in t t ype s . T he se p ha se s d o n o t a ppea r a pp l icab le t o C an tab r ian s i t e s , s uch a s a t L a R ie ra w he re s hou lde red S o lu t r ean p o in t s ( s uppo sed ly am a rke r o ft h e Uppe r S o lu t rean ) w e re f ound i n e a r l ie r l e ve l s ( S t rau s 1 990 :437 ) . S im i la r ly , t h ough t h e C an tab r ian L owe r Magda len ian g ene ra l ly c o r re spond s t ot h e Magda len ian I I p ha se o f t h e n o rma t ive F r ench s y s tem , a va i lab le r ad ioca rbon d a te s i n d ica te a l a te r p e r s is t ence t h an i nF rance , w i th t h e f i r s t t wo p ha se s ( Magda len ian Ia nd I ) a b sen tl o ca l ly ( Gonza lez E chega ray 1 960 ) .

T he p e r iod e xam ined i n t h e p re sen t w o rk , t h e L a te U ppe r P a leo l i th ic ( s pec if ica l ly t h e L owe r Magda len ian ) , s how s a s i gn if ican t i n c rea se i ns i te a bundance . T he S o lu t rean i sr e p r e sen ted p r e sen t ly b y mo r et h an 4 4 s i te s i nC an tab r ian S pa in ( S t rau s 1 992 :96 ) , a l though ma te r ia l s f r om s ome e a r ly e xca v a t ion s h ave b een l o s t o r d i spe r sed o ve r t h e y ea r s . T h i s l i s t

T he p r ima ry d i v i s ion o ft h e C an tab r ian Magda len ian o ccu r s a mong a s semb lage s b a sed u pon t h e p re sence o r a b sence o f l a te ra l ly b a rbed h a rpoon h ead s made o fa n t le r . L owe r Mag d a len ian t oo l a s semb lage s a r e p r ima r i ly c ha rac te r ized b y u nba rbed p ro jec t i le p o in t so fa n t le r , w i th q uad rangu la r f o rms ( r ec tangu la r , s qua re , t r apezo ida l , r hombo id ) b ecom ing f a r mo re p r eva len t i n r e la t ion t o o va te f o rms ( c i rcu la r , o va l ,

o f s i te s i n c lude s E l P endo , E l C a s t i l lo , C ueva Mo r in , R a s C ue to d e l aM ina , A l tam i ra ( A l tuna a nd S t rau s 1 976 ; C ab r e ra a nd G imenez 1 99 1 ; F reeman 1 988 ; G onzä lez E che -

h a lfmoon )t h an w a st y p ica l o ft h ep reced ing S o lu t r ean p ha se . B y d ef in i t ion , L owe r Magda len ian s i te s l a ck h a rpoon s a nd

g a ray 1 988 ; S t raus 1 976 -77 ) , L a R ie ra ( C la rk a nd R icha rd s

o ccu r b ef o re U ppe r Magda len ian s i te s , b u t t h e r e i ss ome o ve r lap i nd a t ing b e tween t h e t wo p ha se sj us t a so ve r lap o c -

1 978 ; C la rk a nd S t rau s 1 983 ; G onza lez Mo ra le s 1 986 ; S t rau s e t a l . 1 981 ; S t rau s a nd C la rk 1 978 , 1 986 ) , L a s C a lda s ( Co rchön 1 981) , a nd B a lmo r i ( C la rk 1 974 -75 ) .

c u r s b e tween t h e S o lu t rean a nd t h e L owe r Magda len ian . E lemen t s o fL owe r Magda len ian l i t h ic t oo l k i t s i n c lude t h e u sua l U ppe r P a leo l i th ic f o rm s , w i th a n i n c r ea se i nt h e i m -

T he S o lu t -

r e an t oo l i n du s t ry i sc ha rac te r ized b y f o l ia te s t one p o in t s i n w i l low l e af , l a u re l l e af , o r s hou lde red f o rm s . Add i t iona l c omponen t s o f t h e l i t h ic i n du s t ry i n c lude e nd sc rape r s , b u -

p o r tance o f n uc le if o rm e nd sc rape r s a nd ( a t s ome s i te s ) o f b acked b l ade le t s .

r i n s , d en t icu la te s a nd n o tche s , a l ong w i th l ow p e r cen tage s o f s i de sc rape r s . L i th ic e l emen t s g ene ra l ly l a ck ing a r e b a cked b l ade le t s , w i th e xcep t ion s o ccu r r ing a ts ome s i te s . I t s o s se o u s i n du s t ry i sw e l l d eve loped , w i th a n t le r p ro jec t i le p o in t s g ene ra l ly h av ing c i rcu la r o r o va l ( ova te ) c r o s s s e c t ion s , a nd

THE LO WER MAGDALEN IAN S ITE OF E L J UYO E lJ uyo i s ac ave s i t e l o ca ted a pp rox ima te ly 8km w e s to ft h e c i ty o fS an tande ra nd 5k m s ou th o ft h ep resen t -day c oa s t l ine o ft h e C an tab r ian S ea ( Ba rand ia rän e ta l . 1 987 ) . T he p r e sen t

o t he ri t ems i n c lud ing e yed n eed le so fb one o ra n t le r .

( a r t if ic ia l )c ave e n t rance i sl o ca ted i n as i nkho le i nt h e l o ca l 6

F igure 1 . 2 : T opog raph ic S u r round ing so fE lJ uyo ( c on tou r s a t 2m ) ( af t e rB a rand ia rän e ta l . 1 987 :20 ) .

7

l i me s tone t o pog raphy , n ea r t h e t o p o f ah i l l o f f e r ing a n e x c e l len t v i ew o f t h e s u r round ing t e r ra in ( F igu r e 1 . 2 ) . T he c o l lap sed p r eh i s to r ic e n t r an ce t o E l J uyo , l i ke t h e m oder n o ne , i sa pp rox ima t e ly 6 0 ma bove s e a l e ve l , w h i le t h e a d ja c en t h i l l s r e a ch o ve r 1 00 m . D u r ing t h e l a s t g l a c ia l m ax i mum , t h e s i t e w a s s l igh t ly f u r the r f r om t h e c oa s t l ine ( t hen l e s s t h an 1 5 k m a way ) . T he s i t e w a s s im i la r ly c l o se t ot h e mon tane r e g ion p a ra l le l t o t h e n o r th c oa s t o f S pa in ( t he C an tab r ian C o rd i l le ra ) , w h ich i s aw e s twa rd e x ten s ion o ft h e P y r enee s . T he se u p land s b eg in t h e i rc l imb n ea r ly a tt h ep r e s e n t -day c oa s t l ine i t s e lf , s e rv ing t o c i r cum sc r ibe t h e a p p rox ima t e ly 6 0 k m w ide c oa s ta l s t r ip k nown i n f o rma l ly a s C an tab r ian S pa in ( s ee S t rau sa nd C la rk 1 978 ; S t rau s 1 992 ) .

( p r ima r i ly s a lmon id s ) a r e a l so a bundan t . L eve l 9i ss im i la r i nc ha ra c t e rt oL eve l 8 ,t h ough mo l lu sc c on t en t i sg ene ra l ly l owe r . L eve l s 1 0a nd 1 l a ck t h e s h e l l a bundance o ft h ep r e v i ou s t wo l e ve l s a nd s o f a r c on ta in d en se c oncen t ra t ion s o f l a rge f a una lr ema in sa nd ( i nt h e l a t te rl e ve l ) al a rge p i t . METHODOLOGY T he m ic rof auna l r ema in s r e po r ted h e r e w e r e r e t r ieved b y w e t sc r een ing a f t e rd i spe r sa l o fc l ay s f r om t h e s i t e ' s d epo s i t s a s p a r t o f ap r oce s s o f f u l l r e cove ry o f a l l a r cha eo log ica l r ema in s ( F r eeman e ta l . 1 988 ) , i n c lud ing f l o ta t ion f o r t h e r e cove ry o fb o tan ica l r ema in s f r om a l s e d imen t s e xcava ted . S c reen m e sh s i z e e mp loyed f o rt h e f i ne f r ac t ion i s0 . 25 mm , o ne o rde r o fm agn i tude f i ne r t h an t h e 0 . 125 i n ch ( 3 .2 mm ) m e sh c ommon ly e mp loyed i n o t he r m od e rn e x cava t ion s ( Shaf f e r a nd S änchez 1 994 ) . T he i n t roduc t ion o f b i a s t h rough t h e l o s s o f r ema in s o f s ma l le r s p ec ie s t h e r ef o r e i s i n s ign if i can t . T h i s s ma l le rs i ze m e sh u s ed d u r ing e x cava t ion i sa l so n ece s sa ry t oa s su r e t h e f u l l r e cove ry o fb o tan ica l r e m a in s . E a ch s e d imen ts ub samp le w a sp l o t t ed t os e c to r( 9p e r s qua r e m e te r ) a nd a n o rma l max imum d ep th o f 5c m . I n l a te r p roce s s ing b y t h e a u tho r , d r ied s e d imen t s w e r e r e s haken t h rough a1mm m e sh s c reen p r io rt os o r t ing , s i nce i t w a s d e t e rm ined t h a tt h e f i ne rd u s t s e r ved t oo b scu r et h et i n i e s t r ema in s , n o r c ou ld r ema in s t h i s s ma l l ( > 0 . 25 mm a nd < 1 . 0 mm ) b e i d en t if i ed u nde r n o rma l c i r cum s tance s . S ed i m en t s w e r e m an ipu la ted u s ing a f i ne b ru sh . T e r r e s t r ia l m amma l ian m ic rof auna l r ema in s a nd t h e r ema in s o f o t he r t a xonom ic g r oup s ( f i sh , b i rd s , r e p t i le s a nd a mph ib ian s ) a nd b one a nd l i t h ic m ic ro too l s w e re s e pa ra t ed f r om t h e f i ne s e d imen t s u nde r c on s tan t i l um ina t ion . T he m ic rof auna r e c ove r ed i nt h el a rge rs c r een m e sh f r a c t ion ( 2 -3 mm )w a sa l so i n co rpo ra t ed i n to t h i s a na ly s i s , a sw e r e t h e r a re i n s tance s o f m apped m ic rof auna lr ema in ss po t t ed d u r ing e xcava t ion .

E xcep t f o rt r a ce s o fs po rad ic u s e , i n c lud ing t h e f ew a r t if a c t s l ef t b y w ha t m ay h ave b een as i ng le B ronz e A ge e xp lo r e r a nd al a t e r( V i s igo th ic ) b u r ia l i no n e o ft h e n a r row b a ck g a l l e r ie s o f t h e c ave , t h e d epo s i t s i nE l J uyo c on s i s t o n ly o f C an tab r ian L owe r Magda len ian m a t e r ia l s ( Ba rand ia rän e ta l . 1 987 ; J a n s sen s a nd G onza lez E chega ray 1 958 ) . T he se m a in d epo s i t s i nt h e c ave ' s f o rme rv e s t ibu le w e r e p r o t ec t ed b y t h e c o l lap se o ft h e m a in e n t rance a r ound 1 4 ,000 B .P . a nd s ub se q uen t s e a l ing b y al a ye r o f f l ow s tone ( F r eeman e t a l . 1 988 :6 ) . T he s i t e w a s r e d i s cove r ed o n ly i nt h e e a r ly 1 950 s a nd t e s t ed a r cha eo log ica l ly f o r t h e f i r s t t i me f r om 1 9 55 t o 1 957 b y J a n s sen s a nd G onza lez E chega r ay ( 1 958 ; s e e a l so G onzä lez E chega ray 1 987 ) . N ew e x cava t ion s w e r e b egun i n 1 978 a nd 1 979 u nd e r t h e j o in t d i r ec t ion o f B a rand ia rän , F reeman , G onz a lez E chega ray a nd K le in , a nd i ns ub sequen t s e a son s ( 1 982 -83 , a nd 1 987 -96 ) b y F r eeman a nd G onzä lez E chega ray ( Ba rand ia rän e ta l . 1 987 ; F r eeman a nd G onza lez E chega r ay 1 9 8 1 ; F r eeman e ta l . 1 9 88 ;K le in e ta l . 1 98 1) . T he o ngo ing e x cava t ion s i nt h em a in v e s t ibu le c ove ra n a r ea o f s omewha t m o r e t h an 4 0 s qua r e m e t e r s ( s ee F i gu r e 1 . 3 ) . A bou t a3 . 5 md ep th o f s t r a t if i ed d epo s i t s e x i s t s a tt h e s i t e ( s ee F igu r e 1 . 4 ) , w i th a n e s t ima t ed d u ra t ion o fo ccupa t ion o f a bou t 1 , 000 y ea r s ( F r eeman e ta l . 1 988 ) . C ha r coa l f r om L eve l s 4a nd 7h a s b een r a d ioca rbon d a ted t o 1 3 ,920+ / -240 B .P . a nd 1 4 ,440+ / 180 B .P . , r e sp ec t ive ly ( F r eeman e t a l . 1 988 :18 ) . T he u ppe rmo s t L owe r Magda len ian d epo s i t i s L eve l 4 , at h in l a ye r o fo ccupa t ion d eb r i s a bove t h e n ea r ly s t e r i le f l ow s tone o fL eve l 5 . I t s l a t e ra l v a r ian t L eve l4 S ( t h e s a nc tua ry c on s t ruc t ed w i th in E lJ uyo ) i n t rude s i n to t h e l owe r d epo s i t sf r om L eve l 4d ownwa rd ( s ee F r eeman a nd G onza lez E chega r ay 1 9 8 1 f o r ad e ta i led d e sc r ip t ion o ft h i s c omp lex ly c on s t ruc t ed s e r ie s o fm ound s , w a l l s , o f f e r ing l a ye r s , a nd r e l a ted f e a tu r e s ) . L eve l 4R amp r e f e r s t oa n a r ea o ff i l l c on t a in ing m a t e r ia l s m ixed f r om o t he r l e ve l s , r e su l t ing a ppa r e n t ly f r om a c cumu la t ion s o f d eb r i s r emoved d u r ing t h e b u i ld ing o ft h e s a nc t ua ry . L eve l s 6 ,7 ,8 ,a nd 9a r et h e l ow e rmo s t l e ve l s w h ich t od a t e h ave b een e xpo sed o ve r aw ide a r ea . S ma l l e xpo su r e s o fL eve l s 1 0a nd 1 h ave b een m ade t od a t e . L ev e l 6i sn o tab le f o r ad ug o u t , s t one l ined , m u l t i r oomed s t ruc tu r e c omp lex y i e ld ing m uch w o rked a nd u n w o rk ed a n t le r , al a rg e l amp i no ne s ma l l l a te ra lr o om , a nd a n a bundan ce o fr e dd ee rr ema in si nt h em a in r oom o ft h e c om p l ex ( K l e in a nd C ruz U r ibe 1 987 ;K le in e ta l . 1 98 1) . L eve l7 c on ta ined t wo b a s in shaped h ea r th s . L eve l 8i sd i s t ingu i shed b y i t sn ea rm idden l ik e c ha ra c t e r : i th a sy i e lded t e n so ft h ou s a nd s o f s h e l l s , m a in ly o f l i mpe t s ( Pa te l la v u lga ta ) a nd p e r iw ink le s ( L i t to r ina l i t to r ea ) ( K r upa 1 994 ) . F i sh r ema in s

Mo s t s p ec ie s w e r e i d en t if ied f r om i s o la t ed t e e th , s i nce c om p l e te o rp a r t ia l t o o th r ow s w e re r a r e ly r e cove r ed . T he m a j o r i ty o fp o s t c ran ia l r ema in s c ou ld o n ly b e i d en t if i ed t og en e r a l m ic rof auna l s i z e c l a s s ( Sma l l , Med ium a nd L a rge ) a nd a r e a na l iz ed t a phonom ica l ly i nC hap t e r 2 . A n e x cep t ion t o t h i s p a t t er n a r e t h e p o s tc ran ia l r ema in s o f mo le s ( T a lpa ) , w h ich a r eh i gh ly r obu s ta nd i d en t if iab le . I s o la ted p o s t c ran ia l r ema in s o f l a rge r s p ec ie s o fm ic rof auna ( mu s t e l id s , h edg e h og , e t c . ) w e r e a l so s ome t ime s i d en t if i ed . T he b a s ic e nu m e ra t ion o f m ic rof auna l s p ec ie s t h e r ef o r e i se xp r e s sed i n t e rm s o f t h e N umbe r o f I d en t if i ed T ee th ( N IT ) , w i th t h e M in imum N umbe r o f I n d iv idua l s ( MN I ) d e r ived f r om ( s ided ) N IT c oun t s . S i nce t e e th a r e g ene r a l ly a mong t h e mo s t d u rab le e l emen t s o ft h e v e r t eb ra t e s k e le ton , n o m a jo r l o s s o f i n f o rma t ion i si n cu r red b y t h i s f a i lu r e t o i d en t if y p o s tc ran ia lr ema in st os p ec ie sl e ve l . C hap t e r 2a l so c ove r s t h e o t he r a spec t s o ft a phonomy r e le v an t t ot h i s a na ly s i s . T he se i n c lude t h e t a phonom i c s i gna t u re s o f v a r iou s p r eda to r / scavenge r t y pe s t h a t p r ey u pon s ma l lm amma l s . T he se s p ec ie s i n c lude d i ur na l a nd n o c tur na l r a p to r s , p a ck r a t s , p o r cup ine s , f e l id s , m u s te l id s , c an id s , v i v e r r id s a nd h e rpe s t id s . T he p owe r o fn a tu ra l f o r ce s s u ch a s mov ing w a te r ,t r ap s f o rmed b y s i nkho le s , a nd i n -d en m o r ta l i t y t oc oncen t ra t e b one s i sa l so b e d i scu s sed . T h e s e k nown moder np a t t er n s a r e r e la t ed t ot h e E lJ uyo s amp le i no r d e rt o d efme t h ep o s s ib le w ay si nw h ich t h eb one sw e r ei n t rodu ced

8

/ 1

I n t e r i o rG a l l e r i e s II i f!, /JO ' , //if

II

1 50 0

1

2I d

2 00

•

\% N .

0 c r n1 00

5 0

4 9 %

6

1 2

q • #

.

-2 -. ; ‘ • ' , '

\ . _

11 0 , . . „ . y \' i r e,, ; #/ ) ,f r . „ . : , . , . , . % . ‹ . . )F i r,, , 2 1i li 10 d 0 0 4/ >

.

i

.

11

8

1 0 1

A

' 1 ,I A \i

7

9

V it

•

C o l lapsed E n t r ance

( d i

1 50 ( 1

W

6'

‘ N , I(

( .-4 " . . . ' 1 4" / 4' 1 -w i

r-

r / 1ir )

b i l i l i / i ) tr ei i f u,

H

lf f

( 7 / 1[ 140 i i' 1 \1 \ oc m 1d i iJ I

I n t e r io rG al e r i e s

Ä I

\ 1\1 4 / ,

i iL

/

\,

/ 1 j

5 0

r, J an s sp

)/

T rench

Mode rn E n tr ance

I / H . 2J I/ 5 0

1 00

I

MI NI 01 PI QI RI S I

E lJ uyo P r inc ipa l Excava t ion s

F igure 1 . 3 : A r ea lE x t en to fE lJ uyo E xcava t ion s( af t e rF r eeman e ta l . 1 988 :23 ) .

9

/ C u l -de -sac

in g •W

3

DEPTH ( cm)

1 00

1 50 • it : r .t e , . . : . .. . r. , . .. • •• • • : . . . . . . . . . .. . •z • > ,, . ., • • • • . . .

e.'.I %. „ ,. . . . .. 7% .,

. . . . ..r ., . . . 4 , . . . . . . „ ., . . . . • •• • • • • e . , ,, . 4 i t 7. . " "i "" 4 2 V ` e be v

4 9" . . .

C4

% ._,

* - ,

• • "% b r% , . . e . % . ./ . . ., : • .

. . . . . • % . . . . . . . . .• Ja i r%

: •

r i ei ` • •• • • • • • _ _' s N . 4. • • • •• . , .. . . . .• , . . • • „ .% t ‘ , . . ,

. . . %

,h ., C " ' . . . . .e% . ., . ." . . .r . L . . e •

• : • ' .• :• \• •• • •: -•::• • -• •

, . . . . " ' ' . %1 . -%• • "‘ % \ . 1 /". 4 . s ., . ,

y* e m •" % * . . e •N % . 2 .• % • • • .1

••; . * : •" A',••••

W '1, 2

. ' "1 % .

•

9

2 00 • • •

• .•

•• :

.•

• • • • . .

•

•

•

•

•

f e

••

•

• • •

•. •

•• • , . • f.•

:

•. •

L f

J UYO 1 978 F igure 1 . 4 : S t ra t ig raph ic P rof i l eo fE l by° ( af t e rB a rand ia rän e ta l . 1 987 :50 ) .

1 0

t ot h e s i t e . T he se i nt u rn s hape t h e i n t e rp r e ta t ion o ft h e p a l e oenv i ronmen ta l d a ta a nd a low f o r g r ea t e r r e so lu t ion o f r e con s t ru c t ion . T he se d a ta a l so i l um ina t et h ec ompe t ing u s e o ft h e c ave b y h uman s , r a p to r s , a nd o t he r s p ec ie s s u ch a s h edgehog o r d enn ing c a rn ivo r e s . S pa t ia l d a ta r ef lec t s i t e u sage a nd s u r f ace a l t e ra t ion p a t te rn s ( i . e . s t ruc tu r e b u i ld ing ) b y E l J uyo ' s Magda len ian o ccupan t s , s i nce m o s t o f t h e se r ema in s w e r e i n t roduced i n dependen t ly o f h uman a c t ion . T he i r n a t u r a l t e ndency t o c o l lec t i n l ow p o in t s o n t h e s i t e s u rf a ce i l um ina te s i ns ome c a se s t h e d u ra t ion o fc on s t ru c t i on ,p a r t i cu la r ly i nt h eS anc tua ry a r ea o fL eve l4 . S pec ie s i d en t if ica t ion a l so a i d s i nd e t e rm in ing t h e p robab le a c cumu la to r s o f t h e m ic rof auna l a s semb lage . N oc tur na l p r ey s p ec ie s t e nd t o i n d ica te n oc tu rna l p r eda to r s a s a n a gency o fa c cumu la t ion . T he s i ze c l a s se so ft h ea c cumu la t ed s p ec ie s a l so a i d s i n p r ima ry p r eda to r i d en t if i ca t ion , s i n ce p r eda to r s o f t en s p ec ia l iz e , h un t ing w i th in ag i ven s i ze r a nge o fp r ey ( Go sz czyn sk i 1 974 , 1 977 ) . D e t e rm ina t ion o fa gen c i e s o f a c cumu la t ion i s o f g r ea t i mpo r tance t owa rd s t h e e va lua t ion o fp o s s ib le b i a se s i nr e la t ive s p ec ie s a bundan ce . E v idence o f d i g e s t ive c o r ro s ion a nd w ea the r ing ( And r ew s 1 990 ; B eh r en smeye r 1 978 ) a l so h e lp s t o i d en t if y t h e t a p honom ic h i s tor y o f t h e a s semb lage . S ome p r eda to r s o f s ma l lf a una ( c an id sa nd d i ur na lr a p to r sm o s tr e levan t ly )h ave as uf f i c ien t ly l ow s t omach p H t oc o r rod e t h e b one s o ft h e i r p r ey ( And r ew s 1 990 ; C la rk 1 972 ; C umm ing s e ta l . 1 976 ; D uke e ta l . 1 975 ; H of fman 1 988 ; K o r th 1 9 79 ) . S uch c o r ro s i on d oe s n o t n o rma l ly o ccu r i nt h e s t oma ch s o fn oc tu rna l r a p to r s( ow l s ) , w h ich a s ag roup t e nd t oe g e s tv e ry c omp le te a nd u ndamag ed o s seou sr ema in s i nt h e i rp e l le t s( Dod son a nd Wex la r 1 9 79 ; L ev in son 1 982 ; P ok ine s a nd K e rb i s P e te rhan s 1 998 ) . I na dd i t ion , a ny c o r ro s ion o n ab r eak s u rf ace w ou ld i n d ica t e t h a t b r eakage o ccu r red a s ar e su l t o fp r eda t ion a nd n o t a s ar e su l t o ft r amp l ing w h i le i nt h e g r ound o rt h rough o t he r p o s tmo r t em d e s t ruc t ive f o rce s . T he se d a ta a i d i nt h e c ha ra c te r iz a t ion o f t h e r e la t ive e f f ec t s o f t r amp l ing , w a t e r t r an spo r t a nd i n i t ia l d epo s i t ion u pon t h e d i spe r sa l p a t ter n o f t h e m ic rof auna i nE l J uyo . T h i s i n f o rma t ion i nt u rn a i d s i n t h e d e t e rm ina t ion o ft h e o r ig in ( s ) o ft h e m ic rof auna w i th in t h es i t ea nd t h e i rr e l iab i l i ty a sp a leoenv i ronmen ta l i n d ica to r s .

t r oduc t ion i n to t h e s i t e . C oncen t ra t ion a r ound a n a r ea o f f o od s t o rage o rd i spo sa l m ay i n d ica t e c ommen sa l i sm b y m i c r omamma l ian s p ec ie so re ven c on sump t ion b yh uman s ,t h u s i n d ica t ing ad i r ec th uman -env i ronmen ti n t e ra c t ion . C hap te r 3d e sc r ibe s t h e mo rpho log ica l a nd e co log ica l c ha r a c te r i s t ic so ft h e s p ec ie s e ncoun t e r ed i nt h eP a leo l i th ic l e ve l s f r om C an tab r ian S pa in , a sw e l l a st h o se f r om moder nr a p to r p e l le t - a nd t r app ing -de r ived c o l lec t ion s . T h i s i n f o rma t ion i s p r e sen ted a nd s umma r iz ed t o a low t h e r e con s t ruc t ion o f P l e i s tocene c l ima t e b a sed u pon t h e r e la t ive a bundance o f t h e se t a xa . T he r e p roduc t ive c y c l e so ft h e se p r ey s p ec ie s a r e a l so i n f o rma t ive c once rn ing t h e p a t t er n s o f s i t e u sage b y p r eda to r sa nd h uman s( d i scu s sed i nC hap t e r 1 0 ) . C hap te r 4p r e sen t s n ew d a ta o n t h e m oder n m ic rof auna o f C an tab r ian S pa in a l ong w i th ar e v iew o fs t ud ie s i nt h e I b e r i an P en in su la . As amp le o fo w l a nd h awk p e l le t s f r om t h e r u ra l e nv i ron so fE lJ uyo h a sb een c o l lec t ed a s ab a se l ine f o r c ompa r i son o fl o ca l e nv i ronmen t a l c hange . T he a r ea h o ld s m any i d ea l r oo s t ing s i t e s , a nd t h e a u tho rh a s o b se rved m any s p ec ie s o fp e l le t -p roduc ing r a p to r( b a rn o w l T y to a l ba , p e re g r ine f a lcon F a l co p e reg r inu s a nd b uzza rd B u teo b u teo ) h un t ing n ea r E l J uyo . O the r s p ec ie s o f o w l c ommon t o n o r ther n S pa in i n c lude S cop s o w l ( O tu s s cop s ) , e ag le o w l ( Bubo b ubo ) , l i t t le o w l ( A thene n oc tua ) , t awny o w l ( S t r ix a l uco ) , l o ng -ea red o w l ( O tu s a s io ) , a nd s ho r t -ea red o w l ( As io f l ammeu s ) ( Bu r ton 1 973 ) . T he o s seou s r ema in s c on t a ined i nt h e r e gu rg i ta ted p e l le t so ft h e se s p ec ie s g i ve at h o r o ugh p i c tu r e o ft h e moder n s ma l l f a una c ommun i ty i nt h e i mmed ia t e s i t e a r ea . T hey a l so i l u s t ra t e t h e r a d ica l c hange s t h e s ma l l m amma l c ommun i ty h a s u nde rgone a s ar e su l t o f f o r e ign s p ec ie s i n t roduc t ion a nd l a nd scape a l te ra t ion d ue t o a g r icu l tu re , p a s to ra l i sm , u rban iza t ion , a nd f o r e s t ry . T he mode rn s ma l l m amma l c ommun i ty i n t h e i mmed ia t e a r ea a r ound E lJ uyo a nd o t he rC an tab r ian l o ca t ion s h a s a l so b een s amp led b y t r app ing t o i n d ica t e t h e c hange s i ni t s m akeup . T he m o r e c omp le t e f a una l c ompo s i t ion a nd l a rge r s amp le a c cumu la t ed f r om o w l p e l le t s a low s d i r ec t c ompa r i son w i th t h e m ic rof auna r ema in s f r om E lJ uyo , s i nce o w l s w e r e t h e i r p r ima ry a c cumu la to r s d u r ing L owe r Magda len ian o ccupa t i on .

A ge p rof i l e sw e r ed e r ived f o re ach s p e c i e ss u ff i c ien t ly r e p r e s e n ted t oh e lp d e te rm ine p robab le a genc ie s o fa c cumu la t ion , s i nce p r eda to r s p ec ie s p r ey u pon d i f f e r en t a ge s e gmen t s o f s ome p r ey p opu la t ion s . A g ing f o r s ome s p ec ie s o f s ma l l m amma l i s a c comp l i shed u s ing t o o th w ea r s t age s ( Adamcz ew ska -And rz e jew ska 1 967 ; D e lany a nd D av i s 1 96 1 ; T rou t 1 978 ) o r e s t ima ted f r om d eg r ee o fp o s tc ran ia l f u s ion . D e t e rm ina t ion o fa ge u s ing l e ng th o ft h e max i l la ry d i a s t ema f o r s p ec ie s w i th c on t inuou s ly g r ow ing t e e th ( Sa in t G i ron s 1 9 73 ) w a s n o t p r ac t ica l , d ue t ot h e l ow n umbe r s o f t h e se e l emen t s t h a t s u rv ived a nd c ou ld b e a s s igned t o s p e c i e s . V e r yy oung s ma l l mamma l ss t i l li n ab u r row o rd en a r e u n l ik e ly t ob e t a ken b y r a p to r s a nd m ay b e i n t roduced i n to a n a r cha eo log ica l s i te b y t e r re s t r ia l p r eda to r s o r t h rough n a tu ra ld ea th i ns i tu .

A s w i l l b e d i scu s sed , n oc tur na l r a p to r s a r e f a r b e t t e r a ccu mu la to r s o fm ic rof auna w i th in t h e i rp r eda to ry r a nge t h an a r e t r ap a r ray s a nd a ss u ch g i ve f a r m o r e c omp le t e i n f o rma t ion o nt h er e la t ive a bundance so ft h es p ec ie sp r e sen t . T he se d a ta s e rve a s ab a se l ine f o r c ompa r i son w i th t h e P l e i s tocene d a ta a nd a s as ou r ce o fb i ome t r ic d a ta . T hey a l so d emon s t ra te h ow t h e m od er na s semb lage sd e r ived f r om o w l p e l le t s r e la t e t o t h e mod er n v ege ta t ion a nd c l ima t e , a nd t h e r ef o r e h ow a s semb lage si n t roduced t oa r cha eo log ica ls i t e sb yo w l s i nt h e p a s tc an b eu s ed t or e con s t r uc tp a leoenv i ronmen t s . C hap t e r 5d e ta i l s t h e m ic rof auna r e cove red f r om t h e n ew e xcava t ion sa tE lJ uyo . T he se s p e c ie s i n c lud e as i ng le mo la r i d en t if i ed t oP l iomy s l e nk i , an ow -ex t inc t m ic ro t ine . T h i s r ema in r e p r e sen t s p e rhap s t h e l a t e s t s u rv iva l o ft h i s s p ec ie s i nC an tab r ian S pa in . C hange s i ns p ec ie s c ompo s i t ion a nd d i ve r s it y a mong t h e l e ve l s i si n t e rp r e t ed i n at a phonom ic f r amewo rk , i n c lud ing s u ch f a c to r s a s t h e e co log ica l h ab i t s ( s e lec t iv i ty o f p r eda t ion , c ho ice o f p r ey , d enn ing / roo s t ing

T he l o ca t ion o f t h e m ic rof auna l r ema in s w i th in t h e s i t e i s a l so d i scu s sed i nC hap t e r 2 . C oncen t r a t ion n ea r t h e f o rme r c ave e n t r an ce o r u nde r al i ke ly r oo s t ing p l a ce f o r ar a p to r w i th in t h e t w i l igh t z one h e lp s t o i n d ica t e t h e a gency o fi n 1

( r oden t s a nd i n sec t ivo re s ) , a st h e i r i n t roduc t ion t ot h e s i te i n mo s t c a se s w a s p robab ly t h e r e su l t o fd i r ec t h uman a c t ion . T he p a t ter n o f t h e se r ema in s ( ove rabundan ce o f f oo t e l e m en t s ) s ugge s t s t h a t t h e se t a xa w e r e e xp lo i ted f o r f u r s . T he se n ew d a ta f r om E l J uyo a r e d i s cu s sed w i th in t h e f r amewo rk o f s ma l l c a rn ivo ran / lagomo rph r e cove ry f r om

h ab i t s , e t c . ) o ft h e a ccumu la t ing s pec ie s . I nt h i s m anne r t h e p a leoenv i ronmen t o fE l J uyo i se xam ined . T h i s a na ly s i s i s l a te r e x tended w he re p o s s ib le t o o t he r C an tab r ian s i te s y i e ld ing s uff ic ien ta moun t so fm ic rof auna , i n c lud ing Ama lda ( Pemän 1 990 ) , A i tzb i ta r te ( A l tuna 1 972 ) , E ka in ( Zaba la 1 984 )a nd E r ra l la ( Pemän 1 985 ) .

o t he r s i tes t h roughou t t h e C an tab r ian P a leo l i th ic . A n h y p o thes iz ed s pec ia l ized e xp lo i ta t ion o ft h e se s ma l l m amma l s f o r t h e i r f u r s i sr e la ted t o t h e g ene ra l p a t te rn o f f a una l e x -

C hap te r 6( a nd A ppend ix A ) p r e sen tt h e r e su l t s o fb i ome t r i c a l a na ly s is o ft h e m ic rof auna f r om E l J uyo . T he se d a ta a l l ow c ompa r i son w i th t ho se d e r ived f r om o t he r s i te s o f

p l o i ta t ion i nC an tab r ian S pa in .

P l e i s tocene a nd H o locene E u rope f o r m ic roevo lu t iona ry c hange s . T he se s ma l l mamma l m ic roevo lu t iona ry c hange s a r e c r i t ica l t oP l e is tocene c h rono logy a s b u i l t u pon t h e s uc c e s s ion o fe vo lu t iona ry f o rms ( Cha l ine 1 974 , 1 985 ; H e in r ich 1 982 , 1 987 ; H in ton 1 9 11 , 1 926 ; J ano s sy 1 975 ; S t ua r t 1 980 , 1 981 , 1 982 ; T chemov 1 968 , 1 975 , 1 979 , 1 982 ; V on K oen ig s wa ld 1 973 ) . S i ze c hange sm ay a l so r e f lec tc hange s i ne nv i r onmen ta l p a rame te r s s uch a sr a inf a l l o r t empe ra tu re ( K le in 1 99 1) a nd w i l l b e c ompa r ed w i th mode rn d a ta f r om s ma l l f a una p opu la t ion s e x i s t ing a c ro s s v a ry ing c l ima t ic r e g ime s . B iome t r ica l a na ly s i s w a s a l so p e rf o rmed , i ns ome c a se s , t o s epa ra te t h ec l o se ly r e la ted s pec ie so f ag enu sw he r et h is w a s n o t p o s s ib le o n mo rpho log ica l g round s a l one . S pec ie s i d en t i f ica t ion w i th in ag enu s may a f f ec t p a leoenv i ronmen ta l i n t e rp re ta t ion . C hap te r 7d i scu s se s t h e o the ra va i lab le s ou r ce so fp a leoenv i -

T he c hang ing h uman e co log ica l i n te ra c t ion s ( f auna l e xp lo i t a t ion , l a nd scape u sage , s i te u sage , t e chno logy a nd p opu la t i on movemen t ) f o rt h e C an tab r ian P a leo l i th ic i ng ene ra l a nd t h e L owe r Magda len ian p e r iod i np a r t icu la r a r e o u t l ined i n C hap te r 1 0 . T he se p a t te rn s a r e r e la ted t ot h e p a leoenv i ron m en ta l r e con s t ruc t ion p o s i ted i n t h e p r ev iou s c hap te r s . I t w i l l b e s hown t h a t t h e se p a rame te r s i n d ica t e a n i n c rea s ing i n ten s i ty a nd o rgan iza t ion o fe nv i ronmen ta l e xp lo i ta t ion w i th i n c rea s ing s epa ra t ion f r om t h e c on s t ra in t s o ft h e f l uc tua t ing L a te P l e i s tocene c l ima te a nd e nv i ronmen t t h rough t h e m e d i a t ion o fc u l tu ra l m echan i sms . B y t h e t ime o f t h e L owe r Magda len ian o ccupa t ion o fE l J uyo a nd t h e s ub sequen t U p p e r Magda len ian a nd A z i l ian p e r iod s , r e sou r ce s w e r e b e ing " w i ld h a rve s ted" ( F reeman 1 973 , 1 975 ) w i th a n e ff i c iency a nd r egu la r i ty t h a t w ou ld p r esage t h e l a t e r c hange s i ne nv i r onmen ta l e xp lo i ta t ion o ft h eN eo l i th ic p e r iod .

r onmen ta l d a ta f r om E l J uyo . T he se a r e c o r re la t ion w i th g l oba l o cean l e ve l c h rono logy , s ed imen to logy , p a lyno logy , l a rge t e r r e s t r ia l mamma l s ( r ed d ee r , b i son , h o r se , e t c . ) , a qua t ic r e sou rce s ( f i sh a nd mo l lu sc s ) , a nd m a c robo tan ica l r ema in s ( c a rbon ized s e ed s a nd o t he r p l an t r ema in s ) . T he l i m i ta t ion s a nd b i a se s o f t h e se m e thod s a r e a l so d i scu s sed . T he se mu l t ip le s ou rces a l ong w i th t h e m ic rof auna l d a ta a r e s yn the s ized i nC hap te r 8f o r E l J uyo a nd t h e C an tab r ian r e g i on t o g i ve a mo re c omp le te p a leoenv i ronmen ta l r e con s t ruc t ion o f t h e L owe r Magda len ian a nd t h e s u r round ing p e r iod s . C hap te r 9r e po r t s t h e f i nd s o f l a rge r m ic rof auna f r om E l J uyo a nd o t he r C an tab r ian S pan i sh s i te s . Wh i le mo s t o f t h e se t a xa f a l lw i th in t h e 5k gu ppe rb ounda ry t h a ta r b i t ra r i ly i su sed t od e l im i tm ic rof auna f r om m ac rof auna , t h i s g roup i s c ha ra c te r ized b y ag ene ra l s u i tab i l i ty f o r f u r e xp lo i ta t ion . T he r e p re sen ted s pec ie s i n c lude w ea se l ( M u s te la n i va l i s ) , e rm ine (M e rm inea ) , p o leca t (M p u tor iu s ) , m ink (M l u t reo la ) , f ox ( Vu lpe s v u lpe s ) , l y nx ( F e l l s l y nx ) , r abb i t

F ina l c onc lu s ion s a r e p re sen ted i nC hap t e r 1 . T he se i n c l ude a s umma ry o ft h e p r e sen ted r e sea r ch a l ong w i th s ug g e s t ion s f o r f u tu re U ppe r P a leo l i th ic r e sea r ch f r om a p a l e oenv i ronmen ta lp e r spec t ive . NOTES ON T ERM INOLOGY F o r t h e s ake o f c l a r i ty , a l l t e e th w i l l b e d e s igna ted b y t h e f o l low ing s hor thand s y s tem . T ee th l i s t ed w i th a s i ng le s u p e r sc r ip t ( uppe r ) o r s ub sc r ip t ( l owe r ) n umb e r w i l l r e f e r t o t h a t s i de a l one , a s i n "M ," r e f e r r ing t ot h e l owe r r i gh t t h i rd mo la r . T he d e s igna t ion " 3M 3"r e f e r st ot h eu ppe rt h i rd mo la r i ng ene ra l , b o th l ef t a nd r i gh t . T h i s s y s t em i sa to dd s w i th o t he r a u tho r s ( H i l l son 1 986 ) , who u se " M 3"t or e f e rt oe i the r l e f t / r igh t u ppe r t h i rd mo la r o r s pec if i ca l ly t h e r i gh t u ppe r t h i rd mo la r a nd t h e reby may i n t roduce c onf u s ion . T he c on v en t ion s o fl e t te r a bb rev ia t ion s s t i l l h o ld : " M " r e f e r s t o mo l a r , " P " r e f e r s t op r emo la r , " C" r e f e r s t oc an ine , a nd " I " r e f e r s t oi n c i so r . U ppe rca se l e t te r s w i l l a lway sr e f e rt op e rma n en t a du l t d en t i t ion , w h i le l owe rca se l e t te r s r e f e r t oj uven i le d ec iduou sd en t i t ion .

( O ryc to lagu sc un icu lu s )a nd h a re ( Lepu ss pp . ) . T h i sg roup i s t h e ref o r e e xam ined s epa ra te ly f r om t h e s ma l le r m ic rof auna

1 2

CHAPTER 2 TAPHONO MY

w i th in a n i n d iv idua l 's h un t ing r a ng e . A s s u ch , c o r r ec t ion s f o r a ny b i a se s i n cu r red w hen u s ing i t s p r ey t oi n te rp r e t p a l e oenv i ronmen t a r e r e la t ive ly s t ra igh tf o rwa rd ( Chap t e r 4 ) . O the r p r eda tory a genc ie s o fm ic rof auna l a ccumu la t ion h ave d i f f e r ing a nd u sua l ly mo re i n vo lved b i a se s f o r w h ich t oa c c oun t ( g rea t e r d i f f e ren t ia l d e s t ruc t ion o fp r ey b one , d i e ta ry s p ec ia l iza t ion o n af ew s p ec ie s , o rf e ed ing h ab i t s t h a t d i f f e r e n t ia l ly c oncen t ra te t h e r ema in s o fp r ey t a xa w i th in a n a r c ha eo log ica l s i te ) . T he se o t he r p o s s ib le a genc ie s a r e e l im i n a t ed ( b e low ) o n mu l t ip le g round s a st h e p r ima ry f o r ce s r e s pon s ib le f o r t h e E l J uyo m ic rof auna l s amp le . T he p r e sen t i n ve s t iga t ion t h e ref o r e f o cu se s u pon b a rn o w l s a s t a p honom ic a genc ie s .

A s e r ie s o f f a c to r s i nf luence s t h e p a s sage o f m ic rof auna l r ema in s f r om l i v ing c r ea tu r e s t o t h e i r f i na l p ub l ica t ion i n a r chaeo log ica l monog raph s a nd a r t ic l e s . T he t a phonom ic s t a tu s a nd h i s to ry o ft h e se r ema in s a r e p r e sen t ed h e r e . T he i n te rmed ia t e s t age sp roceed f r om l i v ing a n ima l st h a t a r e p a r t o ft h e l o ca l p opu la t ion , t ot h e d ead i n d iv idua l s c u l led f r om t h a t p opu la t ion , t o d e f le shed /d ige s t ed s k e le ton s , t ot h e i r i n t r oduc t ion i n to ac ave a nd b u r ia l i ns e d imen t s , t ot h e p r e se r v a t ion o f as ub se to ft h e se e l emen t s , a nd t ot h e i rf i na l r e cov e r y b y f i ne s c r een ing a nd s o r t ing . E a ch o ft h e se s t age s i n v o lve s s ome l o s s o f e l emen t s a nd h ence p o t en t ia l l o s s o f i n f o rma t ion f r om t h ep r eced ing s t ag e( Dav i s 1 987 ;K le in a nd C ruz -U r ibe 1 984 ; S h ipman 1 981) . S i gn if ican t ly , m ovemen t b e tween t h e se s t age s a l so m ay a dd i n f o rma t ion i nt e rm s o f m a rk su pon t h e b one s t h em se lve s , t h e i rp a t te rn a nd l o ca t ion , p r eda to ry a gency o f a c cumu la t ion , a nd d a ta i n he r en t t ot h e s p ec ie s r e p r e sen ted s uch a s mo r ta l i ty p r of i le ( S t ine r 1 994 ) . F a c to r s a f f ec t ing t h e f i na l s t a te o fm i c rof auna l r ema in s a l so l e ave t h e i r m a rk s u pon t h em , f r equen t ly a low ing t h e d e t e r m ina t ion o ft h e s t age s i nt h e w innow ing p roce s s . T he g oa l o ft a phonom ic r e sea rch t h e r ef o r e i st ob a ck t ra ck u pt h i s t r a i l o fi n te rmed ia t e s t age s i no r de rt od e r ive a sn ea r ly a sp o s s ib le ad e sc r ip t ion o f t h e o r ig ina l l i v ing p opu la t ion s o f a n ima l s a nd t h e i r i n t e rac t ion s ( o r l a ck t h e r eo f ) w i th t h e h uman p opu la t ion o f as i te . E ven a p a r t ia l r e con s t r uc t ion o f t h i s p a s sage a low s af i ne r a s se s smen t o ft h e b i a se s a f f ec t ing t h e p ropo r t ion s o f am ic rof auna l a s semb lage a nd i t sr e la t ion sh ip t ot h ee nv i ronmen tf r om w h ich i tw a sd e r iv ed .

M ic rover t eb ra te t a phonomy h a s o n ly a b r ief h i s to ry , s i nce t h e f i e ld w a s f o rma l ly d ef ined f i ve d ecade s a go a s " t he s c i e nce o f t h e l aw s o f e mbedd ing " ( Ef r emov 1 940 :93 ) . I n b roade r t e rm s t a phonomy i st h e s t udy o f t h e p r oce s se s a f f e c t ing t h e r ema in s o f l i v ing c r ea tu r e s f r om t h e p o in t o f d ea th t h rough t h e i r e xcava t ion o rd i scove ry a nd s ub sequen t m an ipu la t ion . M ic rove r teb ra t e s ( d ef ined a ss p ec ie sn o l a rge r t h an 5k g ) o f t en m ake al a rge p o r t ion o ft h i sj ou rney -f rom d ea th u n t i l d epo s i t ion i nt h e p l a ce w he r e t h ey w i l l b e i n co r p o ra ted i n to s ed imen t s in s ide a n o w l . T h i s a bb rev ia t ed j ou rney i sa tt h e o ppo s i te e nd o f as p ec t r um f r om u ngu la t e s t h a t a r e o f t en s l a in b y o ne p r eda to r , s t o len a nd c on sumed o t he r p r eda to r s o r s c avenge r s , d i sa r t icu la ted a nd d i f f e r en t i a l ly t r an spo r ted i n p i ece s t o p r ov i s ion y oung , f e d a nd g nawed u pon a nd d i spe r sed b y as u cce s s ion o f s c avenge r s p ec ie s ( B lumen sch ine 1 986 ; H ayne s 1 980 , 1 9 82 ) , s ome t i me s r e gu rg i ta ted a s m a s se s o f b one b y h yaen id s ( Ke rb i s P e t e rhan s 1 990 ; K e rb i s P e t e rhan s a nd H o rw i tz 1 992 ; S u t c l if f e 1 970 ) , a nd l ef t e xpo sed t ot h e e l emen t s f o r e x tended p e r iod s o f t i me b ef o re c oncen t ra t ion o r b u r ia l o f t h e i r r e m a in s b y g eomo rph ic a genc ie s ( Beh r en smey e r 1 978 , 1 99 1) . Ow l st h u s e ncompa s s mu l t ip le , s u cce s s ive t a phonom ic p roc e s se s i no ne o rgan i sm : s e lec t ion a nd m o r ta l i t y o fp r ey , p r ey d i smembe rmen t , d i ge s t ion o fs of tt i s sue s , i n i t ia l b one b r eak a g e a nd d i ge s t ion , a nd t h e t r an spo r t a nd c oncen t ra t ion o f v e r teb ra te r ema in s t o t h e s i t e o f t h e i r i n co rpo ra t ion i n to a p a leon to log ica lo ra r chaeo log ica ls e quence .

T he c onc lu s ion s o ft h e a na ly s i s p r e sen t ed h e r e a r e t h a t o w l s w e r e o ve rwhe lm ing ly r e spon s ib le f o r t h e i n t roduc t ion o f m ic rof auna lr ema ins i n to t h e s i t e o fE lJ uyo a nd a r et h em o s t l i ke ly v e c to r o fi n t roduc t ion f o r m o s t C an tab r ian c ave s i t e s . Mo re s p ec if i ca l ly , b a rn o w l s ( T y to a l ba ) a r e b y f a rt h e m o s t l i ke ly s ou r ce a mong t h ep o s s ib le s p ec i e s o fn oc tur na l r a p to r . T he e v idence f o r t h i s c onc lu s ion c ome s f r om m u l t ip le s ou rce s , w h ich i n c lude t h e s i t e s e t t ing , t h e a moun t a nd d i s t r ibu t ion o fm ic rof auna i nt h e c ave , t h e i r a ge p r of i le s , s i z e d i s t r ibu t ion , h ab i ta t p r ef e r ence s , a c t iv i ty p a t t e rn s , a nd t h e d eg ree o f b r eakage ( l igh t ) a nd d i ge s t ive c o r ro s ion ( a lmo s t n one ) u pon t h e i n d iv idua l r ema in s . A n e xcep t ion t o t h i s p a t ter n i sf o und w i th t h e r ema in s o fh edgehog ( E r inaceu s e u ropaeu s ) , al a rge r s p ec ie s o fm ic rof auna w h ich p robab ly f ound i t sw ay i n to E lJ uyo a s ac ommen sa l . A l so e xcep t iona l a r e t h e r ema in s o fs ma l l f u r -bea r ing m amma l s s u ch a sw ea s e l ( M u s t e la n i va l i s ) , w h ich m ay h ave e n t e r ed E lJ uyo f r om h uman t r app ing a c t iv i ty ( Chap t e r 9 ) . T he m a jo r i ty o fo t he r m ic rof auna lr ema ins c on s i s t s o fr o den t sa nd i n sec t ivo r e s a nd w e re i n t roduced a st h ep r oduc t s o fb a rn o w lp e l le tr e gu rg i ta t i on .

B a rn o w l s ( T y to a l ba ) a r e a mong t h e mo s t i mpo r tan t a c cu m u la to r s o f m ic rove r teb ra t e a s semb lage s . T he g enu s T y to h ad e vo lved b y t h e M iocene P e r iod a bou t 2 5 m i l l ion y ea r s a go ( Bu r ton 1 973 ) , t hough t h e e vo lu t iona ry h i s to ry o fb i rd s i sl e s s w e l l k nown d ue t o t h e g ene ra l f r ag i l i ty o ft h e i r r e m a in s . A s sum ing c on t inu i ty o f m od er n b ehav io r i n to t h e p a s t ,t h et a phonom ic p roce s se se ncompa s s ed b yb a rn o w l s o r t h e i r n ea r r e la t ive s h ave o pe ra t ed c on tempo raneou s ly w i th h om in id e vo lu t ion f o rm i l l ion so fy ea r s . W i th in t h e s c ope o f t h e p r e sen t a na ly s i s ( l e s s t h an 2 0 ,000 y ea r s ) , b a rn o w l b e h av io r i sa s sumed n o t t o h ave u nde rgone a ny s i gn if i can t c hange .

T he i d en t if ica t ion o ft h e p r inc ipa l m ic rof auna l t a phonom ic a gency a tE l J uyo a s b a rn o w l s i ss i gn if i can t . A s d i scu s sed b e low , t h i s p a r t icu la r p r eda to r i s aw e l l r e sea rched d i e ta ry g ene ra l i s t w ho se p rey a c cu ra t e ly r ef l ec t s t h e e nv i ronmen t

1 3

OWL D IGEST ION Many b i rd s p ec ie s p roduce p e l le t s ( Bunn e ta l . 1 982 ) , t h e u nd ige s t ed r ema in so f mea l s . Among a v ian p r eda to rs p e c ie s , t h o se k nown t oe ge s t p e l le t s a r e d i s t r ibu t ed a mong 6 0 f am i l i e s a nd 30 s p ec ie s a c co rd ing t o as u rvey b y G lue ( 1 970 :54 ) . T he m embe r s o f b o th o w l f am i l ie s ( Ty ton ida e a nd S t r ig ida e , o r de r S t r ig if o rme s ) e ge s t p e l le t s . T he se a r e p r ima r i ly c ompo sed o fb one , t e e th , f u r , f e a the r s a nd a r th ro p od e xo ske le ton . D iu rna l r a p to r s ( f am i l ie s A cc ip i t r ida e a nd P and ion ida e o f t h e o rde r A cc ip i t r if o rme s ; a nd F a lcon ida e , o r d e rF a lcon if o rme s )p roduce p e l le t s , a l though t h e i r o s s eou s c on t en t t e nd s t ob e much l owe r t h an t h o se o fo w l s . M em b e r so fo t he rb i rd f am i l ie ss u ch a sA rde ida e( h e ron s ,b i t t e rn s ) p r odu ce p e l le t s , b u t t h e se t e nd t o l e ave f ew r e cogn iz ab le b one sa nd c on s i s t mo s t ly o ff u r( G lue 1 970 :54 ) .

I n te rp r e ta t ion o fm ic rof auna l a s semb lage s i sc r uc ia l f o r t h e r e con s t ruc t ion o ft h e e nv i ronmen ta l s e t t ing o fh uman e vo lu t i on . B a rn o w l s t e nd t or e gu rg i ta te l a rge ly i n tac t b one s o f t h e i rs ma l l p r ey , g i v ing av e ry c omp le te r e co rd o fw ha tt h ey h ave e a t en a nd m ak ing i d en t if i ca t ion o f s k e le ta l e l emen t s r e la t ive ly e a sy . I nc on t ra s t w i th o t he r a c cumu la to r s o fm i c r of auna l r ema in s , b a rn o w l s r e ad i ly r oo s t a nd n e s t i na ban d oned o rh uman -occup ied s t ruc tu r e s a nd l i v ing s i t e s , s ome t i me s i nc l o se p rox im i ty t oo ngo ing h uman a c t iv i t ie s . T h i s n ea r -commen sa l i sm m ean st h a tr ema in sa c cumu la t ed b yb ar n o w l s w i l l o f t en b e i n te rm ixed w i th a r cha eo log ica l m a t e r ia l s ( B ra in 1 974 , 1 980 , 1 98 1 ; B r a in a nd B ra in 1 977 ) . O ne c an t h e r ef o r e c o r r e la te e v id ence o fh uman a c t iv i ty w i th a n i n d e p enden t s ou r ce o f p a leoenv i ronmen ta l i n f o rma t ion .

R e -

s e a r ch i n to t h e t a phonom ic s i gna tu r e o fb a rn o w l s the t y pe o f b one a s semb lage t h a t t h ey c r ea t e w h i le s amp l ing s ma l l a n ima l p opu la t ion s i s o f i mpo r tance r e ga rd ing t h e p a l e oeco log ica ls i gn if i cance o ft h o se r ema in s . Mo s t p r ev iou s b a rn o w l p e l le t r e sea r ch h a s f o cu sed u pon d i e ta ry d i ve r s i ty a nd e co log ica l i n te r a c t ion s b e tween p r eda t o r a nd p r ey . B ar n o w l p r ey s amp le s h ave b een u s ed t o i d en t if y a nd q uan t if y t h e s ma l l m amma l p opu la t ion s o fr e g i on s s u ch a sB r i ta in ( G lue 1 967 , 1 970 , 1 974 , 1 975 ) , C o lo r a do ( Ma r t i 1 973a ) a nd S pa in ( Br une t -Lecom t e a nd D e l ibe s 1 984 ; D e l ibe s e ta l . 1 984 ) . T he se s t ud ie s a r e v a luab le t ot h e t a phonom i s ti nt h e i ra na ly se so ft h ep r eda t ion h ab i t s , h un t ing t e r r i to r ie s ,a nd s i z er a ng eo fp r ey t a ken b yo w l s . T aphonom ic s t ud ie s o f m ic rover t eb ra te a s semb lage s h ave a l lowed ag r ea t e r u nde r s tand ing o ft h e r e la t ive d epo s i t iona l b i a se s a f f ec t ing t h e p a leoeco log ica l i n f o rma t ion t h a t t h ey m ay y i e ld . Many s t ud ie s h ave f o cu sed u pon t h e v a ry ing t a phonom ic s i gna tu r e s t h a t ap a r t icu la r p r eda to r t y pe m ay l e ave a si tc oncen t ra te s t h e b one s o fm ic rove r t eb ra te p r ey i n o rn ea r ad en , r oo s t , o rn e s ts i t e . C oncen t ra to r s o fm ic rove r t e b ra te b one s i n c lude n oc tu rna l r a p to r s ( And r ew s 1 990 ; A ve ry 1 982a , 1 982b , 1 987 , 1 992 , 1 993 , 1 995a , 1 995b ; B ra in 1 980 , 1 98 1 ; D eny sa nd Mahboud i 1 992 ; D od son a nd Wex la r 1 979 ;H of fman 1 988 ;K u sme r 1 992 ;L ev in son 1 982 ; P ok ine s a nd K e rb i s P e t e rhan s 1 998 ; R aczyn sk i a nd R up r ech t 1 974 ; T ha cke ray 1 987 ; T ha cke ray a nd A ve ry 1 990 ) , d i u rna l r a p t o r s ( Mayhew 1 977 ) , a qua t ic b i rd s ( W i l son 1 987 ) , s ma l l m amma l ian c ar n ivo r e s ( And r ew s a nd N e sb i t E van s 1 983 ; Me l le t t 1 974 ; O k sanen 1 983 ) , a nd c r ocod i l ian s ( F i she r 1 981a , 1 98 1b , 1 981c ) . O the rt a phonom ic a genc ie s t h a tm ay c oncen t ra te m ic rove r t eb ra t e b one s i n c lude p a ck ra t s ( E l ia s 1 987 ) , h a rve s t e r a n t s ( Sh ipman a nd Wa lke r 1 980 ) , n a tu ra l s i nkho le t r ap s ( And r ew s 1 990 ; B ra in 1 980 , 1 981) a nd m ov i n gw a te r( Dod son 1 973 ; K o r th 1 979 ; V oo rh ie s 1 969 ; Wo lf f 1 973 ) . S i gna tu r e s o fa c cumu la t ing a genc ie s i n c lud e a s sem b l age l o ca t ion , d eg r ee a nd t y pe o fs k e le ta l b r eakage , d i ge s t i ve c o r ro s ion , s k e le ta l p a r t r e p r e sen ta t ion , s i z e d i s t r ibu t ion a mong p r ey s p ec ie s ,a nd t h es e xa nd a g ed i s t r ibu t ion w i th in a p r ey s p ec ie s . I d en t if i ca t ion o ft h e p r inc ipa l a gency o fa c cu m u la t ion o f am ic rove r t eb ra t e a s semb lag e w i l l a i d i nt h e a s s e s smen t o ft h e r e la t ive b i a se s w h ich o ccu r red i nt h e s am p l ing o ft h el i v ing p opu la t ion s .

1 4

T he m echan i sm o fp e l le t f o rma t ion i no w l s i si t s e lf a n i m p o r tan t t a phonom ic p roce s s . I n o w l s t omach s r e cen t ly d i s p a t ched p r ey a r ec onver ted i n to c ompac tl ump s o fs oon t ob e e ge s t ed b one a nd f u r . Wh i le b a rn o w l s t e nd t os wa l low p r ey i n ta c t , o t he r s p e c ie s s u ch a sg rea t h or n ed o w l ( Bubo v i rg in i a nu s ) o f t en t a k e p r ey t o o l a rge t o c on sum e w ho le ( J one s 1 9 52 ) . T he se p r ey i t em s mu s t e i the r b e t o rn i n to p i ece s b e f o r e c on sump t ion o r t h e i r f l e sh e a ten o f f t h e i r b one s ( E ina r sen 1 9 56 ; H owa rd 1 958 ) . T h i s b ehav io r n a tu ra l ly a l t e r sr e la t ive s k e le ta l r e p re sen ta t ion a nd b r eakage p e r cen t age s i np e l le t s a mong p r eda to r s p ec ie s . S ome o w l s p ec ie s a bou t t h e s i z e o fb a rn o w l s , s u ch a s s hor t -ea r ed o w l s ( As io / l am m eu s )h ab i tua l ly d i smembe rp rey b ef o r ec on sump t ion e v en i f t h e se i t em sa r er e la t ive ly s ma l l ( Sho r ta nd D r ew 1 962 ) . Af t e rs wa l low ing , f ood p a s se st ot h e g l andu la rp or t ion o ft h e s t oma ch ( p roven t r icu lu s ) , t h en t h rough a s ph inc te r t o t h e m u s cu la r p o r t ion ( v en t r icu lu s o r g i zza rd ) . T h i s l a t t e r s t om a c h c omponen t t e nd s t o b e b ag l ike a nd e x t en s ib le i no w l s a nd o t he r a v ian c ar n ivo r e s ( Fa r r ie r 1 960 :428 -432 ) . I ti sn o t h eav i ly mu sc led i no w l s , t h u s c on t r ibu t ing t ob one p r e se rva t i on . T he p y lo r ic o u t le t i sv e ry s ma l l ( 1 mm o r l e s s ) a nd l o ca ted o nt h e s upe r io rs u rf ace o ft h e v en t r icu lu s a tt h e s ame l e ve l a st h e e s ophagea l o pen ing ( R eed a nd R eed 1 928 ) , t h u s p r even t ing m o s to u tf low o fs o l id i n d ige s t ib le e l emen t s . I na r o en tgenog raph ic s t udy o fp e l le t f o rma t ion i n ag r ea th or n ed o w l , G r imm a nd Wh i tehou se ( 1963 ) f o und t h a t f o l low ing a m ea l t h e v en t r icu lu s r ema in s e x tended f o r a bou t o ne h ou r . T he p e l le tc onden se s o ve rt h en ex t2 8h ou r sa nd i sc omp l e t e b y 1 0 h ou r s a f t e r e a t ing . P e l le t s w e r e t y p ica l ly e xpe l led a bou t 1 6h ou r s a f t e r e a t ing . V en t r icu la r c on t ra c t ion a nd a c c e s so ry c on t r a c t ion s o ft h e a bdom ina l w a l l a nd p roven t r i cu l u s f o r ce t h e p e l le t u pwa rd s i n s ma l l s t ep s , t h e e n t i r e f i na l e ge s t ion p ha se l a s t ing a bou t 4m inu te s . L ow s t oma ch a c id i ty a mong o w l sr e la t iv et oo t he rc a rn ivo r e s c on t r ibu te st ob one p r e se rva t ion ( Duke e ta l . 1 976 ; D uk ea nd R hoad e s 1 977 ) . B a sa l s t omach p H f o r F a lcon if o rme s i s a bou t 1 . 6 , w h i le f o r o w l s t h i s f i gu r e i sa bou t 2 . 35 ( Duk e e t a l . 1 975 ) . P e r cen tage w e igh t o fb one p e r p e l le t i st h e r ef o r e m u ch l owe r a mong d i ur na l r ap to r s . A ve r ag e s a r ound 6 . 5% o fd r y p e l le t m a s s w e r e o b ta ined f o r s ome s p ec ie s o fh awk ( Bu teo ) ,f a lcon ( Fa l co )a nd e ag le ( H a l iae tu s )a sc ompa r ed t o a n a ve rage o f4 5 .8% a mong t wo S t r ig ida e : g r ea t h o rned o w l ( Bubo v i rg in ianu s ) a nd s nowy o w l ( N yc tea s cand ia ca )

a mph ib ian s a nd b a t s a r e t a ken , t h e b u lk o ft h e d i e t i sm ade u p o f n oc tu rna l t e r r e s t r ia l m amma l s . B a rn o w l s e xce l a t s amp l ing l o ca l s ma l l m amma l f a una , s u ch t h a t s ome s p ec ie s h ave b een f ound i n o w l p e l le t s b ef o r e t h e i r p r e sence i n a g i ven r a nge w a s d i scove r ed t h rough t r app ing ( Ande r son a nd L ong 1 96 1 ; H anney 1 963 ; H e im d e B a l sa c a nd d e B eauf o r t 1 969 ) . S ome t a xa h ave b een f i r s t d i s cove r ed t h rough t h e a na ly s i s o fb a rn o w l p e l le t s ( e .g . A nd e r son a nd L ong 1 96 1) . T he k nown r a nge o fs ma l l m amma l s p ec ie s i sa l so o f t en e x t e nded t h rough t h i sa na ly s i s( Bunn e ta l . 1 982 ) .

( Duke e ta l . 1 975 :653 ) . S i nce t h e i n ve s t iga to r s f ound t h a t a ve rage m ea l t op e l le t i n t erva l s a mong a l t h e s p ec ie s t e s ted ( e xcep t f o r H a l iae tu s ) f e l l b e tween 1 8 t o 2 2 h ou r s , d i f f e r e nce s i np e l le t b one c on t en t w e re n o t ar e su l t o fd i f f e r ence s i nd i ge s t ion t i me ( Duke e ta l . 1 975 :654 ) . C la rk ( 1972 )f o und t h a tp r ey b one m ade u po n ly 1 7% o fp e l le tw e igh t f o rm a r sh h awk s ( C i r cu s c yaneu s ) a nd 4 4% f o r s h o r t -ea r ed o w l s ( A s io f l ammeu s )f e ed ing u pon t h es ame p r ey s p ec ie s . S t oma ch p H a ppea r s t ob e t h e m a in i n d ica to r o fp r ey b one s u rv ivab i l i ty b e tween d i u rna l a nd n o c tu rna l r a p to r p e l le t s , a l though f o od h ab i t s p l ay a n o bv iou s r o le . C umm ing s e ta l . ( 1 976 ) m ea su r ed l o s s f r om c l eaned b one s o f h ou se m ou se ( M u s m u s cu lu s ) i ns o lu t ion s o f HC 1 i nc oncen t ra t ion s s imu l a t ing f a l con if o rm a nd s t r ig if o rm g a s t r ic j u ice ( pH 1 . 66 a nd 2 . 35 , r e sp ec t ive ly ) . B ov id p ep s in w a s a l so a dded i nv a ry ing c oncen t r a t ion s . B one l o s sb yw e igh tw a ss i gn if i can t ly h i ghe r t h roughou tt h el owe rp Ht r i a l s . L o s se sa ve raged o ve r7 0% a t t h e f a lcon if o rm c oncen t ra t ion , w h i le b one s t r ea t ed w i th t h e s t r ig if o rm c oncen t ra t ion a ve raged b e low 3 5% l o s sb yw e igh t ( Cumm ing s e ta l . 1 976 :57 ) . I n c r ea s ing t h e s upp lemen ta l b ov ine p ep s in c oncen t ra t ion t e nded t oi n c r ea se b one l o s se s s l igh t ly a te a ch HC lc oncen t ra t ion . O the r f a c to r s h e lp t o e xp la in h ow m o s t o w l s e ge s t i n ta c t b one s o fp r ey . L ow m ix ing o fs t oma ch c on t en t s ( R eed a nd R eed 1 928 ) c on t r ibu te st ol ow l e ve l s o fb one b r eakag e . T he h i gh p ep s in a c t iv i ty i nt h e s t oma ch s o fS t r ig if o rme s c on t r ib u t e s t oe f f ec t ive d i ge s t ion o fp r ey . R eed a nd R eed ( 1 928 ) f o und t h e p ep t ic a c t iv i ty o f B ubo v i rg in ianu s t o b e a bou t t h r ee t im e st h a to f ad og ' s g a s t r ic j u ic e a tc ompa rab le s t age s o f d i ge s t ion . I n a dd i t ion , g a s t r ic p H d oe s n o t r ema in a ta c on s tan t l ow l e ve l . S m i th a nd R ichmond ( 1 972 ) f o und t h a t b ar no w ls t oma ch p H r i se s a f t e r f e ed ing u n t i l p e l le t e ge s t ion t o am ax imum o f a r ound 6 . 0 . T h i s a c id i ty l e ve l i n c r ea se s p r ec ip i tou s ly w i th in o ne h ou r t o a r ound 2 . 0 . Sm i th a nd R ichmond ( 1 972 :183 184 )a l so f ound t h a t ac ap t ive b a rn o w l c ou ld b e i n duced t oe ge s t ap e l le t b y l e t t ing i ts e e l i ve p r ey a f t e r as u ff i c ien tt ime l a p se f r om i t s l a s t f e ed ing . T he m in i mum t i m e f o r i n duced e ge s t ion w a s 6 . 5 h ou r s . P r ey b one s t h e ref o r ec an s p end r e la t ive ly s ho r tp e r iod so ft i me i na n o w l s t oma ch u nde rc e r ta in c i r cum s tance s . BARN OWL B EHAV IOR T y to a l ba i s ap a r t icu la r ly p ro l if i c a c cumu la to r o fs ma l l a n i m a l r ema in s ( Dav i s 1 959 ; T ay lo r 1 994 ) . I t s n ea r c o smopo l i tan d i s t r ibu t ion ( Bu r ton 1 973 ) s pan s ar a nge o f h ab i ta t s , f r om t empe ra t e f o r e s t s t ot r op ica l d e ser t s a nd r a in f o r e s t . T h roughou t t h i s r a nge , b a rn o w l d i e t i sg ene r a l i s t w i th in t h e s i z e r a nge f r om l a rge a r th ropod s t os ubadu l t R a t t u s a nd A r v i co la ( a bou t 1 00 g )( Ave ry 1 993 ; C ampo s 1 978 ; G lue 1 967 , 1 970 , 1 974 , 1 975 ; H e r r e ra a nd J ak s ic 1 980 ; Web s te r 1 973 ) . T he r ef o r e , b a rn o w l d i e t t e nd s t o i n d ica te l o ca l a va i lab i l i ty o fp r ey s p ec ie s . B a rn o w l s t e nd n o t t ob e s e lec t ive i nt h e i rp rey c ho ice s , a l though t h e t e r ra in o ccup ied b y ap r ey s p ec ie s ( t he o w l s p r ef e r t o h un t i n mo r e o pen h ab i ta t s ) a nd t h e a c t iv i ty p a t ter n s o f p r ey ( a t e ndency t o f r eez e w hen s t a r t led o rt os p end mo r et i me o n o pen g r ound , m ak ing c a p tu r e e a s ie r ) w i l l i nf luence a g i ven s p e c ie s ' c hance s o fb e ing t a ken ( De r t ing a nd C ranf o rd 1 989 ) . Wh i le i n sec t s , o t he r a r th ropod s s u ch a s s c o rp ion s , b i rd s , r e p t i le s ,

1 5

A l though o cca s iona l ly o b se rved t oh un t i nd ay l igh tj u s t b e f o r e s un se t o rj u s t a f t e r d awn ( Bunn 1 972 ) , b a rn o w l s a r e p r ima r i ly n oc tur na l p r eda to r s ( Bu r ton 1 973 ; G i l l ia rd 1 958 ) . S pec ia l a dap ta t ion s i n c lude t h e a b i l i ty t ol o ca t e p r ey i nv e ry l ow l i gh t l e ve l s ( D ice 1 945 ) o r i n t o ta l d a rkne s s t h rough h ea r ing a l one ( Payne a nd D ru ry 1 958 ; P ayne 1 97 1) . Mo s t h un t ing i sd one o n t h e w ing w i th in ar a d iu s o f af ew k i lo m e te r s f r om ad ay t ime r o o s t o rn e s t ing s i t e . E van s a nd Em l e n ( 1 947 :3 ) f o und t h a t t h e h un t ing r a nge o f ab ar n o w l s t ud ied i n n o r ther n C a l if o rn ia ( U .S .A . ) c omp r i sed r ough ly 1 65 a c r e s , o r l e s s t h an a0 . 5 km r a d iu s i na ve rage d i s tance f r om t h e r oo s t . Web s te r( 1 973 :186 ) f o und t h eh un t ing r a nge o f an e s t ing p a i r o f b a rn o w l s i n S ou th We s tmoo r land ( Eng land ) t oc ove r 1 47 a c r e s , a ga in l e s s t h an a0 . 5 k m r a d i u s . Mo s t r e sea r che r s e s t ima t e al a rge rr a d iu s , s u ch a sL o v a n ie ta l . ( 1 976 :173 )i nc hoo s ing 0 . 8k mt o 5k m f o rh un t ing r a d i if o rt h eb a rn o w lc o l lec t ion st h ey s t ud ied i nc en t ra l I t a ly . L og ica l ly , s u ch v a r iab le s a s p r ey d en s i ty a nd c ompe t i t ion w i th o t he rr a p to r s m ay e f f ec tt h e h un t ing r a nge f o ra ny i n d i v i dua l b a rn o w l . E ven t h e se l a rge r e s t ima te s o fh un t ing r a d i u s , h oweve r , i n d ica t e e x t r eme ly l o ca l iz ed c oncen t ra t ion o f p r ey r ema in sa tt h ec en t r a lr oo s t . P r ey i sc on sumed w ho le , w h ich c on t r ibu t e s t ot h e c omp le t e n e s s o f d i e ta ry r e p r e sen ta t ion i np e l le t s . G lue ( 1967 :170 ) n o te s t h a t b a rn o w l s a r e v e ry r a re ly o b se rved t o d ecap i ta te s ma l l m amma l p r ey b ef o r e c on sum ing i t . B i rd d ecap i ta t ion i s mo r e f r equen t , e s pec ia l ly b ef o re f e ed ing t o n e s t l ing s . S i nce a v ian MN I ' s a r e u sua l ly b a sed u pon p o s tc ran ia l e l e m en t s , a v ian p r ey c oun t ss hou ld n o tb es i gn if i can t ly a f f ec t ed . Mo s tn i gh t s , ap e l le tc on ta in ing t h e r ema in s o ft h e e a r ly e ve n i ng ' s h un t ing i se ge s t ed a way f r om t h e d ay r oo s t . L a t e n i gh t / ea r ly mor n ing p r ey t y p ica l ly c ompo se t h e p e l le t t h a t a b a rn o w l e ge s t s a ti t s d ay r oo s t b ef o r e l e av ing f o r t h e f o l l ow ing n i gh t ' s h un t ing ( Sm i th a nd R ichmond 1 972 ) . P e l le t s a t an e s t ing s i t ew i l la l so i n c lude t h o se e g e s t ed b yt h e c h ick s . A bou th a lf o ft h e d i e tw i l lt h e r ef o r e b ee ge s t ed a t ad ay r oo s t s i t e . I no rde rt ot e s t w he the rb ar no w l p e l le t s a r e a n u nb ia sed r e f l ec t ion o fd i e t , G lad ing e ta l . ( 1 943 )f e dc ap t ive m embe r so f t h i sa nd o t he rr a p to rs p ec ie s an a tu ra l d i e to fs ma l l mamma l s a nd b i rd s . T he p e l le t s e g e s t ed w e re t h en b l ind t e s ted f o rt h e c on ta ined s p ec ie s . T he h awk s p ec ie s f a r ed p oo r ly : t h e i rp e l l e t s c on ta ined a lmo s t n o i d en t if iab le b one . T he o w l s g ene r a ly a nd b a rn o w l s i np a r t icu la r e ge s t ed p e l le t s w h ich c on t a ined ah i gh ly a c cu ra t e r e co rd o ft h e i r d i e t s . Y oung b i rd p r ey d i d t e nd t owa rd s u nde r rep r e sen ta t ion i np e l le t s o fb ar n o w l s , i n d ica t ing t h a t t h e i r b one s t e nd t o b e c omp le te ly d i g e s ted mo r eo f t en o rr e nde r ed o t he rw i se u n iden t if iab le . T h i s

i n c r ea sed a v ian b one l o s si smo s tl i ke ly d ue t ot h ef r ag i l i ty o f t h e b one st h em se lve s . I n ap a r a l le l s t udy , S e l leck a nd G lad i n g ( 1943 ) e nc lo sed b a rn o w l n e s ts i t e s w i th w i r e i no rde rt o e nume ra t e t h e p r ey c a r r ied t ot h e n e s t l ing s b y t h e i r p a r en t s . T hey f ound t h a tb i rd sm ade u po n ly 6 . 4% b yb u lk o ft h ed i e t , n o t d i f f e r en t s i gn if i can t ly f r om t h e a moun t o f b i rd p r ey i n t h e d i e t ( 6 .7% ) i n d ica t ed b y p e l le t s f r om t h e s ame s i t e . S pa r se a v ian p r ey i np e l le t s t h e r ef o r e s e em s t o i n d ica t e t h e a c tua l h un t ing p r a c t ice s o ft h e o w l s a nd n o t d i f f e r en t ia l s u r v i vab i l i ty o fb i rd r ema in s . Af i na lo b se rva t ion o nt h er e la t ion sh ip b e tween b a rn o w ld i e t a nd l o ca l p r ey a va i lab i l i ty i st h e o cca s iona l h ype rabundance o fo ne s p ec ie s . A n e xamp le o ft h i s s i t ua t ion i s ap e l le t s am p l e f r om I r aq t h a ty i e lded al a rge p r opo r t ion o fb a t s ( n =5 3 , 2 7 .0% o f t o ta l p r ey i t em s ) . T he se s p ec ie s r o o s ted i n t h e s ame l a rge s t ruc tu r e a s t h e o w l s a nd w e r e t h e r ef o re h y p e rabundan t l o ca l ly a nd c o r re spond ing ly e a sy t op r ey u pon ( Nade r 1 969 ) . T ay lo r ( 1 994 ) a l so n o t e s i n s tance s o f b ar n o w l p r eda t ion u pon l o ca l ly a bundan t f i sh ( b eached g run ion ) o rb i rd s o na ni s land w i th f ew t e r r e s t r ia ls ma l lp r ey s p ec ie s . B a rn o w l s a nd o t he r o w l s p ec ie s w i th s im i la r d i e t s a nd r oo s t ing h ab i t s c an r a p id ly a c cumu la t e l a rge n umbe r s o f p e l le t s w i th in ac ave o rh uman s t r uc t u r e . B a rn o w l p e l le t s a r er e s i s tan tt od ecay a nd c an r ema in i n ta c tf o rm any y ea r s i n ad ry d epo s i t iona l s e t t ing . T ho se d epo s i ted i nm o i s t e r s e t t i ng s a r eb r oken d own r a p id ly b yb a c t e r ia , f u ng i , a nd v a r iou s i n sec t l a rvae f e ed ing o n t h e f u r m a t r ix o f t h e p e l le t s ( Lev in son 1 982 ) , t h u s d i spe r s ing t h e i r b one c on t en t s . T he s phe r ica l s h ape o fp e l le t s a low s t h em t ot umb le i n to a nd b e d i spe r sed a r ound ac ave f l oo r w i th s l igh t h i nd rance . T he i r c on ten t s c an t h en b e d i spe r sed i n to c ave d epo s i t s b y t h e d e c ay o f t h e i r f u r m a t r ix a nd t h e p hy s ica l t r amp l ing b y a ny s ub sequen to rs imu l taneou sh uman o ccupan t s . TAPHONOM IC ANALYS IS O FE LJ UY0 T he p hy s ica l l a you t a nd s e t t ing o ft h e c ave o fE l J uyo n a r r ow s t h e p o s s ib le v enue s f o r e n t ry o f m ic rof auna i n l a rge n umbe r s . T he f o rme re n t rance ( r ef e rb ack t oF i gu r e 1 . 3 )w a s w ide a nd o pened d i r ec t ly o n to t h e m a in i n hab i t ed v e s t ibu le a r ea b ef o r et h e p a r t ia l c o l lap se a nd s e a l ing o ft h e c ave . T h i s e n t rance w a s s i tua t ed a tt h e b a se o f av e ry s t eep s i nkho le l e ad ing d i r ec t ly i n to t h e c ave . T he e n t rance o pened o n to t h e b o t tom o f al a rge , b r oad s i nkho le i nt h e l i me s tone b ed rock . S ome s ma l l a n ima l s c ou ld b e e xpe c t ed t od i e o rb e k i l led i n t h i s f unne l shaped a r ea a r ound t h e m ou th o ft h e c ave . T he i r b one s c ou ld h ave b een t umb led , w a shed o rt r a cked i n to t h e c ave h ab i ta t ion a r ea a nd i n co rpo r a ted i n to t h e s e d imen t s . B one s d e r ived f r om c a rn ivo r e s c a t s o r o w l p e l le t s d r opped i n to t h i s a r ea c ou ld a l so h ave c on t r ibu t ed t ot h i s s ou r ce o f b one s . S eve ra l f a c to r s a r gue a ga in s t t h i s s ou r ce o fm ic rof auna r e m a in s b e ing am a jo r c on t r ibu to rt ot h e a s semb lage f r om t h e c ave . P r ima ry i st h e l a rge s i z e o ft h e a s semb lage i t s e lf , c u r r e n t ly c omp r i sed o f7 , 707 i d en t if i ed t e e th , 1 7 ,770 p o s tc ran ia l a nd f r agmen ta ry t e e th r ema in s i d en t if i ed t o s i z e c l a s s , a nd 1 9 ,976 u n iden t if ied m ic rof auna l f r agmen t s . A n a dd i t iona l 1 55 t e e th w e re i d en t if i ed t o l a rg e r s p ec ie s o f m ic rof auna ( h edgehog s , e t c . ) d i scu s sed s e pa r a te ly . Wh i le e ven v e ry

1 6

s l ow w a te r f l ow m ay t r an spor t m ic rof auna l r ema in s ( Ko r th 1 979 ) , i ti sd i ff i cu l tt oe nv i s ion mo re t h an as ma l l p r opo r t ion o ft h e se r ema in sb e ing w a shed i n to E lJ uyo f r om i t si mmed i a t e n e ighbo rhood . A nd r ew s ( 1990 ) c on c lud e s t h a t w a t e r i s mo r e l i ke ly t od i spe r se s ma l lb one st h an t oc oncen t ra t et h em . Wh i le t h e p r o ce s se s o fm ic rof auna l s k e le t a l e xpo su r e d eg r a d a t ion a r e l e s sw e l lr e sea r ched t h an t h o se o fl a rge rf a una ,t h e t wo p roce s se sm u s tb ea tl e a s ti np a r tc ompa r ab le d u et ot h e i r i d en t ica l b one c hem i s t ry . K or th ( 1 979 ) f o und t h rough e x p e r imen ta t ion t h a t a e r ia l ly e xpo sed s ma l l m amma l c a r ca s se s mu s tb e b u r ied q u ick ly o rb one s l y ing o nt h e s u rf a ce w i l l b e d e s t royed . N one o ft h e E l J uyo r ema in s e xh ib i t s s i gn s o f e xpo su r e t y p i ca l ly c au sed b y e x tend ed a e r ia l e xpo su r e o n l a rge f a una l r ema in s ( And r ew s 1 990 ; B eh ren smeye r 1 978 ) , s u ch a sc r ack ing o fe xpo sed s u rf a ce s , f l ak ing i n to p i e ce s , o r b l ea ch ing . T he r obu s t s t a te o f t h i s a s semb lage ( e sp ec ia l ly w hen c ompa r ed w i th t h e f i sh r ema in sr e cove r ed i na n i d en t i c a l m anne r f r om E l J uyo ) a l so a r gue s a g a in s t w a sh ing i n to t h ec ave f r om o u t s ide . Many d e l ica te p o s t c ran ia lr ema in sa r e f o und i na bundance , i n c lud ing r oden t a nd i n sec t ivo r e p ha l a nge s , m e tapod ia l s , a nd v e r teb rae . S uch t i ny r ema in sc ou ld n o th ave b een t r an spo r t ed i nb u lk w i thou t as i gn if i can t , c l ea r f l ow o fw a t e r , s i nce t h e i r s ma l l s i ze p r ec lud e s t umb l ing a c t i on . N o s i gn if ican t f l ow o fw a te r w a s e v e r e n t ran t i n to t h e mou th o fE l J uyo . I ti sp o s s ib le t h a t s ome r ema in s m ak ing t h i s d e s t ruc t iv e j ou rney f r om o u t s ide t h e c ave d i d e n t e r i n p r ev iou s ly g r ea te r n umbe r s , o n ly t o b e d e s t royed l a t e r b y p o s tdepo s i t iona l f o r ce s o rt h e e xcava t ion a nd r e cove ry p r oc e s se s . I n t rodu c t ion i nt h i s m anne r d oe s n o t c ha ra c t e r iz e t h e E lJ uyo a s semb lage . L ime s tone c ave s a l so c an a c t a s n a tu ra l s i nkho le t r ap s f o r s ma l l a n ima l s ( And r ew s 1 990 ; B ra in 1 9 80 , 1 98 1) . S ma l l m amma l s f a l l ing i n to ap i t q u ick ly d i e o fs t a rva t ion d u e t o t h e i r v e ry h i gh m e tabo l ic r e qu i remen t s , w i th t h e i r l a rg e ly i n ta c t r ema in s b ecom ing i n co rpo ra ted i n to s i t e s e d imen t s . T he p hy s ica l l a you t o fE lJ uyo p r ec lude s t h i s s c ena r io , s i nce t h e w ide f o rme r e n t rance a lowed e a sy e g r e s s , n o r d o e s a ny n a tu ra l c h imn ey f e a tu r e e x t end t ot h e s u r f ac e a bove t h e i n h ab i ted v e s t ibu le a r ea . T he p o ten t ia l f o r t h i s p roce s s t oa c c umu la te m i c rof auna m u s t b e e xam ined i nt h e c a s e o fe a ch c ave s i t e s e t t ing . T he c u r r en t n ea r ly s e a l ed e n t ranc e o fE l J uyo d oe s a low t h e o cca s iona l t r app ing o fm oder n m ic ro f a una w h ich c anno t f i nd a n e x i t f r om t h e b a ck g a l le r ie s o f t h e c ave . S ome r e cen tr ema in s o fr a t ( Ra t tu sr a t tu s ) a nd r e d f o x( Vu lpe sv u lpe s )h ave b een a ccumu la t ed i nt h i sm anne r . W i th t h e e x cep t ion o fs now v o le (M i cro tu s n i va l i s ) t h e b e h av io ro ft h e se s ma l l m amma l s pec ie s ( Ch ap t e r 3 )p r e c lud e s t h e i r u s ing t h e c ave a s ad enn ing s i t e a nd t h e r eby a dd ing t h e i r r ema in s t h rough n a tu ra l mo r ta l i ty ( Ave ry 1 982a ) . M n i va l i s , h owev e r , p r ef e r s r o cky c r ev ice s a mong t a lu s s l ope s a nd o t he ra r ea so fe xpo sed r o ck o ve rn a t u r a l c ave rn s . Wh i le m any o ft h e l o ca lr oden ta nd i n sec t ivo r e s p ec i e sa r en a t u ra l ly f o s so r ia l , t h e i r b u r row s y s tem s a r e c on s t ruc t ed i n w e l l d r a ined ,e a s i ly t u nne led t o p so i l . T he se t u nne l s i ns ome c a se s a l so s upp ly f o od : e a r thwo rm s a nd a r th ropod s i nt h e c a s e o f mo le s ( T a lpa s pp . ) a nd p l an t r oo t s i nt h e c a s e o fa r c t i c v o le ( M i c ro tu s o e conomu s ) . N one o ft h e se c r i t e r ia a r em e tb yt h e d en se ly t r amp led , o b jec t -den se , p oo r ly -d r a in ed a nd v ege ta t i on -f r ee s e d imen t s o fa n i n hab i ted l i me s tone c ave v e s t ibu le .

A lmo s tn ob u r row ing d i s tu rbance h a sb een d e tec ted i nt h eE l J uyo d epo s i t s . N o C h i rop t e ra ( b a t ) r ema in s h ave y e t b een r e cove red , t h ough t h e se a ppea r i n l ow a moun t s i n o t he r C an tab r ian s i t e s d ue t o t h e i r n a tu ra l c ave roo s t ing t e nd en c i e s . T he r ema in so ft h el a rge rc ommen sa l s p ec ie sh edgehog ( E r ina ceu s e u ropaeu s ) m ay h ave e n te r ed t h e E lJ uyo d epo s i t s t h rough t h e i r n a tu ra l i n s i tu m o r ta l i ty a nd s ub sequen t b u r ia l o fs k e le ta l e l emen t s . W i th t h e e xcep t ion o ft h i s l a t t e r s p ec ie s , d enn ing o f m ic rof auna s p ec ie s w i th in t h e c ave i n b e tween e p i sode so fh uman o ccupa t ion i sn o ti n d ica ted .

d a len ian d epo s i t s a l so s ugge s t s t h a t h uman p r e sence w a s e nough t od e t e r s imu l taneou s s i t e o ccupa t ion b y t h e se s p e c i e s a nd p e rhap s o t he r s . T he se s p ec ie s , h oweve r , a r e m o r e l i ke ly t oh ave r oo s ted a nd d i ed i ns i tu i nt h e ( unexcava ted ) i n te r io rg a l le r ie so ft h ec ave . P e l le t sa c cumu la ted i nt h ec ave d u r ing e p i sode s o fo w l o ccupa t ion m ay t h en b e i n co rpo r a ted i n to s ub sequen t h uman o ccupa t ion d eb r i s , i n t e rm ix ing t h e t wo e ven t h ough e ach s pec ie s o ccup ie s t h e c ave a td i f f e r en t t ime s . D i s t r ibu t ion p a t te rn s o fm ic rof auna m ay a l so i l um i n a te t h e a l te ra t ion o ft h e s i t e s u rf a ce a nd s t r uc tu r e b u i ld ing b y h uman s , s i nce r ound o b jec t s r a ndom ly t r amp led o ve rt h e s u rf a ce o ft h es i t ew i l ln a tu ra l ly t e nd t os e t t le i n to o pen , l ow l y ing s pa ce s . O pen p i t s t ruc tu r e s , s u ch a st h e s t one r oom o f L eve l 6a nd e s pec ia l ly t h e s a n c tua ry s t ruc tu r e o f L eve l 4 S w ou ld c oncen t ra t e m ic rof auna e ge s ted b y t h e o w l s . T he se p a t ter n s a nd t h e i r i mp l ica t ion s f o r h uman s i t e u sage w i l l b e d i scu s sed i nC hap t e r 1 0 .

H uman u t i l iz a t ion a nd d i spo sa l o f t h e se s p ec ie s , w i th t h e i mpo r tan te xc ep t ion o fs ma l l f u r -bea r ing c a rn ivo r e s( Chap t e r 9 ) , i sn o t as i gn if ican t f a c to r i nt h e i r a c cumu la t ion . G iven t h e i r l ow y i e ld o f m ea t r e la t ive t ot h e e f f o r t n ece s sa ry f o r t h e i r c ap t u r e , s ma l l r od en t s a nd i n s ec t ivo r e s ( mo s t u nde r 4 0 gi nt o ta l a du l t m a s s ) h ave r a r e ly b een as i gn if i can t h uman f o od r e sou r ce . T he i r u s e a s af o od s ou r ce i se s pec ia l ly u n l i ke ly i n ar i ch s ub s i s tence e nv i ronmen t l i ke C an tab r ian S pa in , w he r e aw ide r a nge o fl a rge r o r mo r e e ff i c ien t ly h a r v e s t ed s p ec ie s e x i s ted ( s ee C hap t e r 7 ) . Mo r e d i r ec t ly , t h e p a t te rn o fb one r e cove ry a nd b r eakage a r gue s a ga in s t t h e i r u se b y h uman s . S uch c omp le t e b one s a r e a n u n l ike ly r e su l t o f a ny c ook ing p roce s s , w h ich w ou ld w eak en b one s a nd c au se g r ea t e r p o s tdepo s i t iona l d e s t ruc t ion t h an h a s b een t h e c a se . I na dd i t ion , t h e s pa t ia l d i s t r ibu t ion o fm ic rof auna i nt h e s i t e i n d ica te s t h e i r i n t roduc t ion b y n on -human m ean s . T hough n o t a l a r ea s o ft h e u ppe r h ab i ta t ion l e ve l s h ave b een c om p l e te ly e xcava t ed ( l eav ing g ap s i nd i s t r ibu t ion ) , p r e l im ina ry a na ly s i s i n d i ca t e s t h a t m ic rof auna l r ema in s c l u s te r t owa rd s t h e n o r thea s t c or ne r o ft h e c av e v e s t ibu le , d i r ec t ly u nde r a n a tu ra l l a rg ef i s su r e i nt h ec ave c e i l ing o ri nn a tu ra l l ow spo t s d own s lope f r om t h i s l o ca t ion . T h i sc e i l ing c hambe r i so u to f u na ided h uman r e ach , a nd i sb yf a rt h em o s ts u i tab le r oo s t ing p l a ce w i th in t h i s c hambe r f o r ar a p to r . I ti sa l so w i th in t h e t w i l igh tz one o ft h e c ave , w h ich i sp r ef e r r ed b yr a p to r s . T he r oo s t ' s s h ap e a l so a l low s e a sy s p i l lage o f e ge s ted p e l l e t s d own i n to t h em a in f l oo ro ft h ec ave . O the rl o ca t ion s w i th in t h e v e s t ibu le o f f e r n o a dequa te p l a tf o rm s p a ce o ra r e w i th in e a sy a c ce s s t o t e r r e s t r ia l p r eda to r s o f r a p to r s : h uman s , c an id s , f e l id s a nd o t he r s . S i nce h uman s t r uc tu ra l f ounda t i on s h av e b een d i scove r ed , i tr ema in s s l igh t ly p o s s ib le t h a t o w l s r oo s t ed o n a bandoned h u t s o r s im i la r c on s t ruc t ion s w i th in t h e v e s t ibu le b ef o re t h e se c o l lap sed o rw e r e d i sp la ced b y s ub sequen t h uman o ccupa t ion s . U nf o r t una t e ly f o r m od e r n i n ve s t iga t ion , t h e l a you t o f t h e c ave w ou ld a l so h ave s p i l led m uch o w l e ge s ta t ion i n to a n i n t e r io r , u n inhab i ted a nd u nexcava ted g a l le ry o fE lJ uyo , w he r et h ey a r e l o s tt or e cov e r y a mong s t ra t if i ed h uman o ccupa t ion r ema in s . T he E l J uyo s amp le i st h e ref o r e ar a ndom s ub se t o ft h e c omp l e te m ic rof auna a c cumu la t ed a tt h es i t e . T he s pa t ia l a r rangemen t o f t h e c ave a l so s ugge s t s t h a t h u m an s a nd b i rd s w ou ld n o t h ave b een o ccupan t s a tt h e s ame t i me . T h e p r ev iou s e n t rance w a s s ma l l e nough t op l a ce a ny h uman so rh uman s t ruc tu r e s( s uch a si nL ev e l6 )i nt h e d i r ec t f l igh t p a th o ft h e b i rd s . T he f r equen tu s e o ff i r e i nt h e c ave m igh t a l so d e t e r r oo s t ing . T he c omp le t e a b sence t od a te o f a ny i d en t if iab le c h i rop t e ran r ema in s w i th in t h e L owe r Mag -

1 7

E LJ UYO SKELETAL R ECOVERY AND BREAKAGE T ab le 2 . 1 l i s t s m ic rof auna l r ema in s f r om E l J uyo i n c lud ing i d en t if i ed t e e th ( d e sc r ibed i nC hap te r 5 )t h a t m ay h ave b een i n t roduced b y b a rn o w l s . T he s e h ave b een g rouped b y g en e r a l s i z e c l a s s , s i nce f o r t h e m o s t p a r t t h e se r ema in s c anno t b e a s c r ibed t os p ec ie s o rg enu s . T he m a in e xcep t ion t ot h i s r u le a r e t h e p o s t c ran ia l r ema in s o f T a lpa , w h ich a r e h i gh ly i d en t if iab le ( d i scu s sed b e low ) . T he m a in s i ze c l a s se s( Sma l l , Med ium a nd L a rge ) g ene ra l ly c onf o rm t ot h e f o l low ing t a xa w ho se p r e sence h a sb een d e t e rm ined b yt h e i rd en ta lr ema in s : Sma l l : T h i s c l a s s i n c lude st h e p o s t c ran ia l r ema in s o fs h r ew s . T ho se i d en t if i ed f r om E l J uyo a r e N eomy s f od ien s , N . a noma lu s , S o rex m inu tu s a nd S .c o rona tu s . E ach o ft h e se s p ec ie s h a s a n a ve rage a du l t m a s s o fl e s s t h an 1 5 g( Chap te r 3 ) . T he se h ave b een i d en t if i ed i nl ow n umbe r s f r om t h e s i t e , w h ich i s r ef lec t ed b y t h e r e la t ive ly l ow t o ta l o f n on i d en t if i ed t o o th r ema in s a s s ignab le t ot h i s g ene ra l c l a s s ( n= 2 6 1) . T he se t o ta l s i n c lude i n c i so r a nd f r agmen ta ry m o la r r ema in s , s i nce ( u n l ik e m ic ro t ine s ) t h e se w e r e i d en t if iab le t o s p ec ie s . Max i l la ry a nd m and ibu la rc oun t si n c lude t h o se w i th i d en t if iab le t e e th t h a t w e r e a s s ignab le t os p ec ie s , a sw e l l a s u n iden t if iab le f r agmen t sa s s ignab le o n ly t os i z ec l a s s . Med ium : T h i s c l a s s i n c lude s t h e r ema in s o t he r t h an i d en t i f i ed t e e th o f s ma l l r oden t s . T he se h ave a n a ve rage a du l t b ody m a s s o f2 0 -50 g . T he t a xa w ho se p r e sence h a s b een d e te rm ined f r om t h e i r mo la r s a nd t h a t c onf o rm t ot h i s c l a s s a r e A pod emu s ( Mu r ida e ) a nd P l iomy s l e nk i , P i tymy s p y r ena i cu s , P .l u s i tan i cu s , M i c ro tu s n i va l i s , M . o economu s , M . a g re s t i s a nd M . a r va l i s ( M ic ro t ida e ) . T he p o s s ib le p r e sence o ft h e p o s t c ran ia l r ema in s o fo t he r t a xa c onf o rm ing t ot h i s g ene ra l s i z e c l a s s ( s uch a s C l e th r ionomy s g la reo lu s ) c anno t b e d i scoun t ed . T he se r ema in s a l so i n c lude r oden t i n c i so r s , w h ich w e re n o ta s s ignab le t os p ec ie s . Mand ibu la r a nd m ax i l a ry r ema in s a r e s ummed t o g e the r e ven w he r e t h e se w e re i d en t if iab le t o s p ec ie s , s i nce t h i s w a s p o s s ib le o n ly i nr a r e c a se s . M ic ro t ine mo la r f r agmen t s a r e a l so i n c luded . T he l a rge t o ta lo ft h e se n on iden t if i ed t o o th r ema in s ( n =1 ,287 ) c o r re spond s w i th t h e l a rge n umbe r s o ft h e se t a xa i d en t if i ed f r om t h e i rt e e th .

Mi c ro f a una l SMALL Max i l la Mand ib le

e m am sf r om E lJ uyo b yS i z eC l a s sa nd L evel

S capu l a H um e ru s R ad iu s U ln a P e lv i s F emu r T ib-i ö -f ibu la ri lcan eu s T a lu s T o ta l , MED IUM

Mo la rf r ag S capu la H um e ru s R ad iu s U lna P e lv i s F emu r T ib io f ibu l ä C a lcan eu s T a lu s T o ta l LARGE Max i l la Mand ib l e I n c i so r Mo la r Mo la rf r ag S capu l a H um e ru s R ad iu s U ln a T ra i i i s F emu r T ib io -f ibu l a r i lcan eu s T a lu s TRT t I—

0

9 0

1 2

0 0

0 0 0 0

1 0

T th

Max i l la Mand ib l e I n c i so r Mo l a r

0

1

8 / 9

8

7

6 / 9

5

6

0

0

0

0

1 2

5 1 7

0 0

0 1

0 0

0

0

0

0

0

0

1 0 0

1 0

0

3

0

0 0-

0 0

0

1

0

5 0 1

I 0

0 0

1 3

0 6

0

1

0 0

0

1

1 0 0

0 0 0.

1

3 0 1

0 0 0

5

1

0

1

1 0 0 0

1 3 1 2 4 0 2 0 0 0

3 3 4 1

1 0 0 0

1 00

0

1 6

1 2 0 4 3 6

1 / 3 0 0 2 23 1 0 1 1 0

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

9 0 0 2 3

/ 0 3 1 0 0 0 0 0 0 II

3 8 8

2 0

4 3

0 1

1

1

1 0 9

0 0

1 2 0 3 5

1 0

1 3 1 2 3 9 3 2 6 1 4 2 2 2 2 2 8 1

8 / 9 0 0

1 05 2 30

2 0

0 0

0

0

0

2

2 3

0 0 0

1 1 0

2 5-

3

2

0

7

0

0

0

0

0

0

0 2

t s 0

8 3

1 8 7

0

1

1 3

5

8

0

0 0

1 6

1 6 8

6 / 9

2 2 6 4

0 0' 0 0

2 3

0

4

1 4 4 6 5 09

0 0

3 4

0 2

2 1 33

2 5 3 4 12 1 2 5 9 2 2 2 8 1 2 5 -3 2 13

8 / 9

9 1 9 1 6 2

7 0 0 a

0 0 0 0 0 " 0 0 0 0 0 0

4

0 0

3 4 2 2 6 1 3 1 5 -

8

4 S 0

0 6

0 0

4 R 0

7

1 2 2 4

4 2 5 9 4 I 1 4

1 67 . 6

5

0

4 0

0 2 7

8 5 4 10 7 28

1 0 1 1 1 0

6 0 7 2 1 57 4 2 9 7 3 5

0 0 0 0 1 3 6 / 9

0 3 6 4 6

0 0 0 1

1 1 3 0 3 0 1 3 0 1 6 8

0 0 0 ' 0 0 0 0 1 0 0 2

0 1 0 0

r

3 0

9 S 9 22 0

1 4 1 5 5 2 3 0 3 0 2 5 5 6 3 6 1 3 1 0 9 65

1 8

2 1 6 5

2 1 3 4 5

0

4 S

2 26 1

1 5

5 2 4 1 2

7 88 6 1 1 82

0

0

0

0

0

0 0

0 0

0 -ö -

0 0

0

0

0

0

6-

9 7

0 0

0 0

0

0

8 4 4

1 21 6 67

0

0

2 7

1

0

1

0

0 0

0

0

1 9 1 9

4 09 2 88

1 8 4

0 0

0

0

0

4 87

2 / 4

3

2

0

I

1 5 7 6

3 22 1 5 36

1 3 8-

2 340

7 8 3 4 1 7 4 1

0 1 7

1 1 1 4

1 8 1 0 3 1 7 1

3 73 15 3 1 1 9 0 1 99

1 2 1 4 9 1 696 5 1 06

6 6 3 1 0

4 7

4 4

0

40 6, 3 57

3 1 6 066

1 07 9 7 4 0 2 3 2 029

2 2 0 0 2 2

4

0 9 4 5 8 9

3 8 1 0 7 5 3 1 1 583

3 0 1

5 5 1 0 S r i 4 1 7 8

2 00 4 0 1 2 3 7 4 8 9 3 2 6 2 5 5 6 1 3 3 003

1 6 2 5 3 6 2 4 2 1 1 7 3 9 3 7 1 6 8 6 79

7 3 2 0 1

1 00 0

0

1 99 1 7 8 4 5

0 0 0 0 0 0 2 0 0 0 2

0

0

0

0

0

0

4

4 S

WI W I ) I 3 -6 -

9 6 6

4 8

4 R

T o ta l 0 6

8 0

8

1 0 0

0

2 2 1 3 4

0 0 0

2 0 0 0

3

1

5

3

0

2 3

9

2 / 4— 0 0 0

0 0 0

1 6 1 947 6 1 7 4 2 1 66 4 59

4 R 0 0

3

1 1 3 9 8 9

2 / 4

1 0 0 I 1 0 0 1 0 0 0 9

T o ta l

I 0 2

0 0 2 1

3 1 0 0 0

0 0 0 0 0

60 0 0 0 2 0

0 1

0 0 1 0 0 0 0 0 0 0 4

3 1 0 4

1

r

0 0 0 1 2

1

0 0 0 6

0 0 1 2

1 0 0 0 0 0 0 0 ' 0 0 7

0 0 0 ' 0 0 0 0 0 1 0 4

2 88

%R ep 1 50 4 1 6 5 3 1 1 00 0 2 550 1 9 1

4 534 6 40 2 50 7 23 2 23 4 35 1 37 4 59 4 08 1 22 8 4 1287 T o ta l

0 0 2 2 0 0 0 0 0 0 0 0 0 0 4

6 76 * 1 96 5 67 1 75 S 4 1 1 07 3 60 3 20 96 6 6

%R ep

7 0 2 08

1 45 4 30

9 02 2 647 2 54 9 4 2 43 1 1 4 1 1 5 3 7 9 1 78 1 56 1 00 4 0 5 265

9 3 3 1 00 0 * 1 94 5 03 2 92 3 16 1 63 3 68 3 23 2 07 8 3

T ab le 2 . 1 : ( Con t . ) 1 0

1

T a lpa

8

9

6 / 9

7

4 2

1

6 9

5 7

1

1 02

3-

0

Maxi l la

0

0

Mand ib l e

3

0

1

S capu la

0

0

0

0

0

0

2

0

1

U lna T er v i s

0 0 0 0 0

0

0

0

0

3

0

0

0

0

F emu r

0

0

1

0

0

0

2

0

0

0 0

0

0 0

0

0 0 1 0

1

8

R ad iu s

1

C a l can eu s T a lu s

4

P ha l /Ca rp /M e ta

onal t

1 0

1

0 0

0

0

3

0

0

0

2

0

2

0

1

1 2

T o ta l

1 0

2 4 9

8

1

0

7

8 / 9

1 07

4 64-

3 1 0

0

2

0 0 1 0 0 0 1 0 0

0 0

5 4

0 0 0 0 0 0

0 6 8 3

1 0 1

0

1

2 3 06

1 5

3 2 4 7 4 R

4

4 S

4 R 1

0

9

6

6 / 9

0

2 66

1 20

4

4 49-

1 95-

4-

4

1 1 0

3

0

0

4 S 2 93

6 1 2 4 3 5

1 3 1 3 8 1 4 5

1 0 0

2 6

4

5 34

3

8 3

1 0 37

2 6 3 4

3 2

2 85 4 76

0

5 7 8

4 38 1 -1 5 3

4

4 12

Me tapod ia l P h a lange s

5 1

T o ta lI d en t .

9 2

3 0

3 65

173

U n id en t . F rag

1 09

3 8

4 63

163

1 4

2 46

T o ta lA l l

2 12

7 8

8 37 '

2 344

1 8

6 65

6

0

6

6

5 2

5 3

3 1 3

5

0 0 0

5

0

4

4

1 9

2 28

4

1 1 0

5 7

0

V e r t . /S te rn .

6

2 6

1 5

R ib

3

4 1 72 ' 6 10 1 24

5 1

2 / 4

c i 8 4 6

T o ta l

2

3

0 0 0 0 0 0

0 0 0 0 0 1 0 1 0 0 0 1

0

7

2 / 4

3 2

2

4 2

0

N ot

5

6

1 0 1 0 0 0 0 0 0 0 0 0 1

2 2

3

' Hum e ru s

1

8 / 9

T ee th

0 0 0 0 0 1 0 0 0 0 0 0 0 1 2

3

1

0 1

0 0 1

4

4

5 066

1 2

8 92

1 2607

4 404

5 3

3 6

2 1

806 6 3

1 2

6 30

8 69 1

4 660

6 5

3 8

3 0

868

2 9

1 522

2 1298

q 0 6 8

18

7 7

5 3

2 7

3

0 0 0 0 0 0 0 0 0 0 0 0 0 0

3 06 1 0 5 2 3 3 1 8 1 9 1 8 3 7 1 4 5 8 56 1 365 T o ta l

0 0 1 1 1 0

8 14 2 506 1 0-0 7 2 473 2 ' 5204 1 9976

5 1 6

4 5180

c a lcu la t ed d ue t od if e rence si ns p ec i e st o ta it e e th .

* I n c luded w i th m o la rc a lcu la t ion s

T ab le 2 . 2 : C ompa r i son o fB a rn Ow lP rey S ke le ta l R ep re sen ta t ion ( / 0 ) . N egev '

K u sme r ( 1990 )

D od son 8 z , Wex la r

K or th ( 1979 )

H of fman ( 1988 )

A nd rew s ( 1990 )

MEAN

( 1979 ) MN I= Max i l la Mand ib le

2 39 9 2 .2 8 8 .4

9 4 9 4 9 4

3 4 7 4 9 0

1 7 1 00 .0 9 4 .1

2 0 8 7 .5 9 2 .5

5 0 8 6 9 8

6 0 5 4 .2 7 1 .6

7 1 6 0 .5 1 00 .0

S capu la H ume ru s

8 2 .9 9 1 .1

6 1 8 2

6 5 8 7

8 8 .2 8 2 .4

6 2 .5 8 2 .5

8 8 9 9

5 8 .3 6 8 .3

5 2 .8 7 2 .5

7 2 .1 8 4 .9

R ad iu s U lna P e lv i s F emu r

7 8 .5 9 1 .8 8 8 .0 8 7 .8

6 8 7 9 6 5 8 1

7 6 8 1 8 9 1

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

6 2 .5 8 2 .5 7 2 .5 8 5 .0

7 5 9 2 9 2 9 4

5 0 .0 5 6 .7 9 0 .8 1 00 .0

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

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

T ib io -

9 1 .1

8 3

9 7

6 7 .7

8 7 .5

1 0 1

1 00 .0

1 00 .0

9 2 .1

F ibu la MEAN

8 8 .0

7 9

8 3

8 2 .7

7 9 .4

9 2

7 2 .2

7 2 .5

8 2 .5

( F rom P ok ine sa nd K e rb i sP e te rhan s 1 998 ) . 2

3

T o ta l s amp le MN I . P e rp a r tm ean sa r ew e igh ted a cco rd ing t os amp le s i ze .

1 9

( 585 )2 8 3 .2 3 9 0 .2

uyo S ke le ta lE lemen tB r eakage—Sma l l . T ab le 2 . 3 : E lJ

T o ta l

Max i a

Mand ib l e _

S capu la

H ume ru s

0

0

4

0

Ö

3

2

0

0

0

0

6-

0

T o ta l

1

0

1

5

0

0

0

C omp

0

F rag

1

T o ta l

0

0 0

0 1

1 4

0 0

0

0

3

2 5 3

0

0

0

0 0

0 22

0

0

0 0

3

2

1 3

2 1 5 1 6

1 0

0

0

0

0

0

0

0

0

0

0

1 2

0

0

0

0

5 4

P rox

0

0

0

0

0

0

0

0

0

0

D i s t

o 0 0 0 0 0 0 1 0 0 0

T o ta l

0

1

1

5

0

3

0

1 4-

0

2

0 0

1 0

0

1 0

2 0

0 0

0

1 0

4 - 0

0

0

0 0 0

P rox E p i

0

0

0

0

0

1

0

0

S ha f t

0

0

0

2

0

1

0

2

0

0

0

8

6

0

1

0

0

1

-

0

2 3

6 6

0

0

0

0 0

0

0

0

0

0

3

0

0

0

0

5 2

3

0

0

0

0

2 1

0

9

0

2

0

0

0

0

0

0 0

0 2

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

4

0

2 2

2

1 0

D i s t

0

T o ta l

a o o o o 0 o 1 o o 1 0 0 0 0

0

1

8

0

C omp

0

9

0

o 0 0 0 1 0 0 o 0 0 o 0 o 0 o o 6 0 0 0 0 0 0 0 0 0 0

1

2

0

0

0

0

0

o

0

0

0

0

0

0

0

1

1

0

2 ' 7 2 1

0

D i s t

0 0

0

0

0

0

0

0

0

0

0

T o ta l

0

0

6

1

0

0

0

2

0

1

0

0

0

0

1

0

0

0

1

0

0

3 2

0

0

5 0

0

C omp

0

0

0

0 -

‘ t

P rox

0

0

0

0

0

0

0

1

0

1

5

1

0

0

0

0

8

S haf t

0

0

0

0

0

0

0

0

0

0

0

0

T o ta l

1

1 0

0

0

1

0

3

0

0

0

0

0

0

0

0 0

8

0

1

0

0

0 0 0

0

0

2 0

0

0

0

7

3

6

C omp

1

0 0

F rag

0

0

T o ta l

0 0

C omp P rox P rox e p i S haf t

0

D i s tE p i

0 1

0

0

0

0 T

3

0

0 0

0

r

0

0 18

1

0

0

7 1

0

0

0

0

0

4

0 '

1

0

6

0

0

0

0

0

6-

0

0

0

0

0-

1

1

0

0

0

0

0

0

0 0-

1

0

0

0

0

0

0

0

0

0 0 0 0

D i s t

T ib io f ibu la

0

0

' P rox

F emu r

0

F rag

C omp

1 5 -g v i s

0

T o ta l

1

2

3

2 / 4

4

4 S

4 R

5

6

C omp

P rox

Ul n a

0

6 / 9

7

8 / 9

o 0 1 5 0 0 I 1 o o 0 0 o 0 0 0

C omp

R ad iu s

0

8

9

1 0

1

I I

0

1

2

0 0

0

0

0

0

0

0

0

n

4

0

0

0

0

2 7

0

0

0

0

0

0

0

1

0

0

0

0

0

3

1

1

0

0

0

3

1 2

0

0

1

3

0

7

7

0

0

0

0

C omp

0

0

0

0

0

0

1

0

0

0

1

0

0

0

0

P rox

1

0

1

1

0

0

0

1

0

0

4

4

0

0

0

0

P rox E p i

0

0

0

0

0

0

0

0

0

3

2

0

0

0

0

, D i s t /Shaf t

0 0

4 4

9 4

4

0

T o ta l

1 2

0 0

0 2

8

0

0 0

0

2 0

6

0

2 7 2 1 2 0 1 3

T ab le 2 . 4 : E lJ uyo S ke le ta lE l emen tB reak 1 0

1

Med ium . 9

7

8 / 9

8

6 / 9

6

4 R

5

T ax a

Mand ib l e

I

o t a

S capu la

H um e ru s

U lna

P e lv i s

0

0

5 19

4 1 7 6 1 5 36' 3 8 9 57 ' 2 8 3 7

1

1

2

1

0 1

0

550 2 2 6 55 2 I :•

1 7 1 4 3

7 7 0

1 5 174 1 4 534 0 6 40

1 0

0 0

0 0

0

0 0

7 3 6 4 9

8 7 88 7 7 28 1 6 0

0 0

1 0

2 0

1 6 1 2 4

4

5 1 4 1 1 0

2 64

0

2 30 3 4

T o ta l C omp P r ox Di s t

0 0

0 0

T o ta l C omp P rox P rox E p i

2 I 0 0

0 0 0 0

9 2 2 2

0 0

16

7 2 0

1 1 57 4 1 2 5 1 0 0 1 7-

1 1 0 0

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

0 0 0 0

0 0 0 0

1 3 0 1 0 3

0 0 0 0

1 1 0 0

1 1 0 0

4 2 1 0 3 1 1

3 2 1 0

T o ta l C omp ' r ox S haf t

0 0

0 0

4 0

1 2

0 0

6 1

1 1

9 75

0 0

0

I

0

0

0

0

0

3

0

T o ta l

0 0 0

3 0 3

5 1 4

0 0 0

2 0 2

0 0 0

3 5 1 3 4

0 0 0

. 1 s t

mig i mi n g 3 0 1 1

0 0 0 0

0 0 0 0

1 0 0 0

minn i mi n i

mg

0

i

0

0 0

0 0

0 0

4 35 3 0

0

0

0

0

2 3

0 I

0 0 0

0 0 0

0 0 0

1 40 2 1 38

06 1 1 9 9 1 8 1 2 1 ,1 6 i t ) 1 4

0 0

4 4 0 4 4

4 7 6 4 7 "

•

0 0

0 0

0 0

0 0

0

0

2

0

o

r 5

l i

2

1

1

7

0

1 0'

3 2

0

1 4 3 1

0

3

0

9 2

0

1

0

2 7

0

0

0

4 .

a• ' 1s t

2 1

0

0

I le In

mi u maii 1

[ C omp P rox P rox E p i

7 23 9 1 1 92 7 9

0 0

0 0

I

! T o ta l

I

2 23 4 6 1 7 3 4

0 0 0 0

9 0 1 7 9 0

na me

1 2

I

0 0 0 0

1 0 1 0

3 1 6 3 i

2 50 0

1 . 6 5 2 0 4 5 0

7 1 6 0

p i

T ib io -f i bu la

1 6 9 3 1 3 6 2 6

0 0

1 5

0 0 0 0

i s t

4 9 0

Eimmi n i

T o ta l C omp P rox , D i s t

S haf t

3 17-

5 1 0 1 0 0

0 0

1

T o ta l C omp I P rox P rox E p i

8 0

913 . 8 62 1 1 56 2 4 1 54 0 7 1 1 38 2 1 2 1 0 1 8 3 73

15 2 1 1 1 11 1 1 1 1 1E I MM IL

C omp F r ag T emu r

1

0

1 0 5 3 8

r

0

2

3 8

2 6 3 7 3

7 8

2 4 16

4 0

1 2 0 3 2

5 31 1 2

2 2 9

0

0 0 0 0

0 0

1 5

2 1

2 0

0 0

0

0

6 0 6

0 0

1 0

8 1

0 0

3 1

0 0

0

2 4 3

0

0 0 0

T o ta l I d en t if i ed U n iden tF r ag

7 8 0

22 1 5 3 5 0

0 0

6 1 8 5

0 0

0 0

0 0

•1 0 0

0 0

gr 4

T o ta l

I

I

0 4 8

0 0

0

1 7

0

3 7

0 0

,:

0 1

0

0 0

5

: l

0

2 1 0

1 3

:

0

0 2

T o ta l

I

l 3

0 0

D i s t R ad iu s

I

0 0

C omp T rag Mo la r

I

C omp F rag

C omp F r ag I n c i so r

I

I

T o ta l

1

2

3

2 / 4

4

4 S

1

1 3 4

1 7 8 5 7 4

9 7

2

3 5 4

0 0

0

min d

I

I

I

i

. .

1 0

4 08 1 3 5 1

i

1 5 4

T ab le 2 . 5 : E lJ uyo S ke l e ta lE l emen tB r eakage -La rge . 1 0

1 Max i l la

,

Mand ib l e

T o ta l C omp

S capu la

H um e ru s

U lna

P e lv i s

F emu r

1

0 0

0 0

0 0

0

1 7 2

0

0 0

1

3 8 0

0

0

0

1

0

0

0

1 5

0

0

3 8

T o ta l ; Comp

0

0

3 1

5

0

3

0

4 2

0

9

1 07

3 9

0

1

3 9

0

8

T o ta l C omp

0 0

0, 0 2

1 0 0 1 0

2 1 2

0

0

5 31 8 4

8 9

4

0

1

2

3 0

1

0

0

2

1 7 6

4 47

5 9

3

0

1

0 !

7 26

1

7

2

2 2 90 1

3 0 1

1

6

2

2 00

1 6

0

1

0

647 2 2 54 0 2

1 2

0

T o ta l

1 0

0 0

2 0

5

C omp

0

P rox

1

0

D i s t

0

0

0 2

T o ta l

0

3

6

4 7 4 6

0

4 5 6

1

0

1 4

0

5

0

1

0

1 5

0

5

0

0

0

0

0

0

2 5 0

0

0

4 0 0

0

0

0

0

0

0

0

1 4

0

0

0

0

0

0

1

0

0

0

1

0

1 5

0

5

0 0

0 0

0 0

3 9 1 ,

9

0

3

0

5 2

0

0 0

0 0

0

0

5 8

1 0

3 9

2 5

1 23

3 6 '

1

0

1 2 3 2

5 6

0 0

0 0

4

0 0

8 1 3

0 0

1 0

1 2 2 1 7

0 0

2 0

2 9 4

1 4

0 0

0 0

0 0

0 0

6 1 1

0

3

4 6

1 0

1

0

0

0

8 3

3 0 1

0

8

7 4

2 4

1

0

0

0

1 4 1

0 0 0

1 7 0

5 6 9 0

6 1 6 2

0 0

0 0

0

1

0

0 0

0 0 0

1 3 1 2 3 5

0

4

8 9

2 1

0 0

0 4

1 8 4

0

0

4

3 0

0

3 1 1

1

0

2

1 0 1

1

2

0

0

0

! P rox

0 0 0

0 1

0 2

! D i s t

0

0

0

0

0 0 0

0 0 0

0 0 0

T o ta l C omp P rox

0 0 0

0 0 0

4 0 4

2

0

0 2

0 0

3 0

0 0

3 0 1

3

0

2 7

S haf t

0

0

0

0

0

0

0

1 1

0

0

2

2

0

0

0

2 5

0

1

3 2

C omp F r ag

0

0

0

0

0

2

0 2

0 0

0 0

0 b-

0 2 5

0 0

0 1

0 3 2

T o ta l

0

0

2

8

0

1

0

5 6

2

7

6 2

0

J

0

0

0

0

2

3

0

1

0

0

1

0

0

0

0

0

0

0

0

D i s t

0

3 -

0

0

0

0

0 0

0

D i s tE p i

0-

2

0

0

0

T o ta l

2

0

2

0

3

1

f P rox

P rox E p i S haf t /D i s t

0 1 0 1

0

0

1 2

f

1 3 4 0 0 1 5 6 3 6

1

0

0

2 2

7

1 7

0

0

0

0

7 9

0

0

0

1 7

0

0

0

0 0

7 9

3 9

r

6

0

0

1 7 8

0

0

0

0

1

0

0

0

0

2 2 4

0

9 1 00

0

0

0

0

0

0

0

0

0

2

7

0

0

0

0

4 0

1

1

4

0

0

0

0

2 1

0

0

0

0

1 56

0

0

0

0

0

0

0

0

4 1

0

0

0

0

2 4

0

0

0

0

8 4

0

8

0

0

2

2 5

0

0

3

0

0

0

0

0

1

0

0

0

1

0

2

4-

0

1 5 3 4 1 4 4

1

3

0

0 0 0

4

0

5

0

0 0 2

1

7

0 '

1 8 1

0 0

3

1

1

3 2

0

0

0

0

1

5

0

2

0

0

0

6

2

0

0

0

2 7 2

0

1

0

3 0

0 0

0 0 2

C omp

2 43

0 0

1

0 0 0

0

0

9 4

4 1

0 0

0

9 02

0

3 17

3

0

1 9 5

0

1 783

2

S haf t

0

3 7

1 583

0

0 0

1 3

0

1 0 1

8 9

I d en t i f i ed

P rox E p i

2 08

0

9 4

U n iden tF r ag

P rox

0

0

0

0 0

0

0

0

0

12

0

0

0

73 1 4 1 4 59

1 24

0

0

4

3 9

3 9

T o ta l C omp

6

0

4 2

D i s t

0

0 1 36

5

0 0

0

4

3

P rox E p i S haf t

0

0

3

1

1

66 0 1 0 0 3

3

1

0

2

2

0

0

0

6

0

0

0

0

0

1 5

0

0

3 4 1 9

0

7 0

1 0

0

1 0

2

3

2

T o ta l

1

3

0

0

0

6

2

0

5

T o ta l

C omp

T ib io -f ibu la

1

2

0

C omp P rox

R ad iu s

0

1

F r ag

F r ag Mo la r

0

0

3

2 / 4

4

4 S

4 R

5

6

6 / 9

7

8 / 9

1

F r ag I n c i so r

0

8

9

1 3 7 5 5 1 0 6 3 4

3 7 1 1 2 9 1 5

4 4

7

T ab le 2 . 6 : E lJ uyo S ke le ta lE lemen tB r eakage -Ta lpa . 1 Max i l la

T o ta l C omp

S capu la

H ume ru s

U lna

P e lv i s

F emu r

T ib io -F ibu la

0

0 0

1 0

7

8 / 9

8 0 0

0 0 0

6 / 9 1

6

0

0

0

1

0

0

0

1

0

0

3

1

0

0

0

0

0

0

3

0

1

0

0

3

1

T o ta l

0

0

0

0

0

0

0

C omp

0

0

0

0

0

0

0

0

0

0

0

2

0

2

0

0

0

T o ta l

3

C omp

0

F rag

F rag

0

0

0

T o ta l

0

0

0

C omp

R ad iu s

0

9

1

F rag Mand ib l e

1 0

0

0

0

0

0

1

0

0

0

0

0

0

C o

0

0

0

0

3

0

0

0

0

1 0 5 2

0

0

2

0

0

0

2

1

0

0

0

0

0

1

0

0

1

9

0

0

1 0 0 1 0

3

0

0

2

1 0 0

6

0

0

0

0

0

0

0

0

0

6

0

0

0

0

1

1

0

0

0

0

3

0

0

0

0

0

0

0

1

0

0

0

0

3

0

0

0

0

0

0

0

0

1

0

0

2 2 1 8

2 6 1 2 5

1 0 4

1 0 3

1 5 1

3 3

0

1

5

3

0

0

1

0

0

0

0

0

0

0

0

1

5

3

0

0

1

0

1 8

0 ,

0

L 7

4

0

1

0

0

1 9

0

0

0

0

0

0

1

0

0

0

4

0

0

7

4

0

0

0

0

0

0

0

0

0

0

0

0 1

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

6

0

0

5

5

0

1

0

0

0

0

0

0

1

0

0

2

0

0

0

0

0

1

0

0

0

0

5

0

0

3

5

0

1

0

0

1 5

2

0

0

1

0

8

0

1

3 7

0

0

1

0

1

0

0

0

1

0

1

0

0

0

5

0

0

0

0

2

0

1

0

5

0 1

0

0

0

3

0

4

0

0

0

0

0

0

0

0

0

0

0

3

T o ta l

0

0

0

C omp

0

0

0

0-'

0

0

0

F rag

0

0

0

0

T o ta l

0

0

C omp

0

0

1 0

F rag

0

0

T o ta l

0

0

0

T o ta l

0

0

0

2

C omp

0

0

0

0

F rag

0

0

0

T o ta l

0

0

C omp

0

F rag

0

3

0

0

2

0

2

0

6

0

F rag

0

7

0

0

0

0 0

3

T o ta l

1

2

6

1

0

0

0

3

2 /4

0

F rag

C omp

3

4

4 S

4 R

5

0 2

0

0

0

0

2 3

7

0

1 9 2 1 7

6

0

0

0

0

0

0

0

0

0

6

0

0

0

0

0

1 1 8

1 8 3

4 3 3

L arge : T he r ema in s i nt h i s c l a s s c onf o rm i na lmo s t a l l c a se s t oA rv ico la , al a rge m ic ro t ine g enu s w i th am ax imum a du l t ma s s o f2 00 -300 g . C onf u s ion w i th r ema in s o fs ma l le rt a xa , e ven o t he rm ic ro t ine s , i st h e ref o re u n l ike ly . N o o t he rp r ey i n t h i s s i ze c l a s s h a s s im i la rp o s tc ran ia l r ema in s . R e jec ted p o s -

Max imum E lemen t MN I %R ep re sen ta t ion —

X1 00 E lemen t MN I

s i b i l i t ie s i n c lude Mu s te la n i va l i s( wea se l ) . Wh i le t h i ss pec ie s

P e r cen tage r e p re sen ta t ion f o r t h e r e la t ive ly mo s t a bundan t

f a l l s i nt h i s s i ze c l a s s , i ti s as ma l l c ar n ivo ran a nd h a s d i s t in -

e l emen t w i l l t h e ref o re e qua l 1 00% , w i th o t he r mo r e f r ag i le e l emen t s f a l l ing b e tween 1 00 % a nd 0% . D if f e ren t ia l b r eak -

g u i shab le s ke le ta l c ha rac te r i s t ics . Wh i le a du l t T a lpa c an e xceed 1 00 g ,t h e i rr ema in sa r eh igh ly d i s t inc t ive d ue t ot h e i r s pec ia l ized f o s so r ia l l i f e s ty le . I ti s , h oweve r , p o s s ib le t h a t s ome f r agmen ta ry T a lpa p e lv ic r ema in s m ay h ave b een i n -

r ough g u ide t or e la t ive a bundance . S i nce t h e se b i a se s a pp ly

c l uded i nt h e t o ta l s f o r t h e L a rge s i ze c l a s s s i nce n one w e re d i r ec t ly a s s ignab le t ot h i s g enu s . A l l t o ta l s f o r mo la r f r ag -

t o a l l a s semb lage s , i n te ra s semb lage c ompa r ison o n t h e se c r i te r ia r ema ins v a l id . T he g ro s s c ha ra c t e r i s t ic s o fr e la t ive

m en t s i nt h e L a rge c l a s s a r e d i rec t ly a t t r ibu tab le t oA r v ico la ,

e l emen tr e p re sen ta t ion a r e s uf f ic ien tt oe l im ina te mo s tp o s s i b l e a ccumu la t ing p r eda to r s a sd i scu s sed b e low .

a ge o fe l emen t s i n to mu l t ip le p i ece s a nd d i f f e ren t ia l i d en t if i a b i l i ty o f t h e se f r agmen t s make s t h i s c a lcu la t ion o n ly a

s i nce i ti st h e o n ly m ic ro t ine g enu s w i th in t h i s c l a s s i n E u rope . T he l a rge i n c i so r r ema in s m ay s im i la r ly b e a t t r ib u t ed t oA rv i co la , s i nce t h e re i sg ene ra l ly n o o t he r r oden t o f

T ab le 2 . 2 c omp i le s p e r cen tage r ep re sen ta t ion f o r k ey m ic ro f a una l s ke le ta l e l emen t s d e r ived f r om s t ud ies o f b a rn o w l

t h i s s i ze p o s s ib le . T he l a rge n umbe r o fn on iden t if ied t oo th r ema in s i nt h i s c l a s s ( n =2 6 18 ) c o r re spond s w i th t h e l a rge

p e l le t s . T he se f i gu re s m us t b e c on s ide r ed a s m ax ima l a moun t s f o rt h e s u rv ivab i l i ty o ft h e se e l emen t s . I ne a ch c a se t h ey a r e d e r ived f r om u nd i s tu rbed p e l le tc o l lec t ion s , a nd t wo o f t h e s t ud ie s ( Dod son a nd Wex la r 1 979 ; H of fman 1 988 ) u t i l ized p e l le t s e ge s ted b y c ap t ive o w l s f e d ac on t ro l led d i e t . B reakage i na n a r chaeo log ica l o rp a leon to log ica l s e t t ing w i l l n a tu ra l ly b e much g r ea te r , a nd f u r the r i n c rea sed b y t h e m e c han ica l f o rce s o fe xcava t ion a nd r e cove ry . T he se v a lue s d o r e f lec t t h e e xceed ing ly l i gh t d amage i n cu r red b y b ar n o w l s u pon m ic rof auna l b one , a nd t h e r e la t ive d u rab i l i ty a mong t h e se e l emen t s .

n umbe r so f mo la r si d en t if ied t ot h i sg enu s . T a lpa : A s m en t ioned a bove , t h e r ema ins a t t r ibu tab le t ot h i s h i gh ly d i s t inc t ive g enu s a r e l i s ted s epa ra te ly f r om t h e o t he r l a rge m ic rof auna . T he se a r ea l so n o t i n c luded i nc a lcu la t ion s f o r b reakag e , s i nce t h e p o s tc ran ia l e l emen t s o f T a lpa a r e r obu s t , h igh ly s pec ia l ized a nd d o n o t c onf o rm i ng ene ra l t o t h e b r eakage a nd p a r t r e p re sen ta t ion o ft h e r oden t a nd s ma l l i n sec t ivo re t a xa d i scu s sed h e re . A t o ta l o f 1 065 n on i d en t if ied t o o th r ema in s w e r e i d en t if ied t o T a lpa . T he se b e long t ot h es pec ie s To cc id en ta l i sa nd T e uropaea .

No t s u rp r i s ing ly , t h e mo re d u rab le e l emen t s a r e t h e t i b io f i bu la ( mean r e p r e sen ta t ion o f 9 2 .1 % ) , m and ib le ( 90 .2% ) , f emu r ( 86 .1 % ) a nd h umeru s ( 84 .9% ) . T he se e l emen t s a r e mo re r obu s t a nd i d en t if iab le when i n ad amaged s t a te . T he s e cond s u rv ivo r sh ip t i e r i so ccup ied b y t h e m ax i l la ( 8 3 .2% ) , u l na ( 80 .75 ) a nd p e lv i s( 80 .2 % ) . T he l owe rr e p resen ta t ion o f t h e max i l la i se xp l icab le b y t h e f r equency a t wh ich t h e c r a n i a l e l emen t s o fy oung m ic rof auna f a l l a pa r t a l ong t h e i r s u t u re l i ne s w hen c on sumed b y o w l s . T he u l na i sb o th mo re g rac i le t h an t h e f emu r o r h umerus a nd i sl e s s r e cogn izab le . T he j uven i le p e lv i s , w h i le s upe rf ic ia l ly r obu s t , o f t en f a l l s a pa r t i n to r e la t ive ly u n iden t if iab le i s ch ia l , p ub ic a nd i l ac p o r t ion s w hen a n i n d iv idua l i sc on sumed b efo re t h e se e l e m en t s h ave f u l ly f u sed . T he r ad iu s ( 7 3 .4% ) a nd s c apu la ( 72 .1 % )a r et h e l e a s tr e cove r ed e l emen t s . T he r ad iu s i sd i ff i c u l t t os epa ra te f r om s ma l l mamma l m e tapod ia ls , w h ich a r e s im i la r ly l o ng a nd t h in . L ow p e r cen tage r e p re sen ta t ion o f

O ther : T o ta l n umbe r s o fr i b s , v er teb rae / s ter neb rae , m e tapo d i a l s a nd p ha lange s w e re t a l l ied r ega rd le s s o f s i ze c l a s s , s i nce t h i s c ou ld b e d e t e rm ined w i th c e r ta in ty o n ly r a r e ly . S ome o f t h e se e l emen t s w e re a s s ignab le t o T a lpa a nd a r e i n c luded i nt h a t c a tego ry 's t o ta ls , s i nce t h e p ha lange s , c a r p a l s a nd m e ta ca rpa l s o f t h e f r on t l imb s o f t h i s g enu s a r e h igh ly d i s t inc t ive . D e sp i te t h e i r s ma l l s i ze , t h e se e l emen t s make u p t h e b u lk o ft h e i d en t if ied T a lpa r ema in s . F o r n on T a lpa c a tego r ie s ,t h e se e l emen t st o ta l6 800 . N o t L i s ted : O the r n on -den ta l r ema in s a r e n o t i n c luded i n t h i s t a bu la t ion . F o r l a rge rt a xa s uch a sl a gomo rph s , p r e l im i n a ry a na ly s i s i n d ica te s t h a t t h e se o ccu r i n e x t r eme ly s ma l l n umbe r s i nE lJ uyo . T h i s i n c lude s a na ly s i s o fr ema in s f r om s ma l l s c reen s , l a rge s c reen s a nd mapped i t ems . T h i s p a t t e rn o fr e cove ry i sd i scu s sed i ng r ea te rd ep th e l sewhe re ( Chap te r 9 ) . N on -den ta l r ema in s o fo ne m ed ium m ic rof auna s pec ie s , E r inaceu s e u ropaeu s ( h edgehog ) a r e d i scu s sed s epa ra te ly a t t h ee nd o ft h i sc hap te r .

t h i s e l emen t t h e ref o r e m ay b e d ue l e s s t og r ac i l i ty a nd mo re t o al a ck o fd i s t ingu i sh ing i d en t ify ing f e a tu re s . T he l ow p e r c en tage r e p r e sen ta t ion o f t h e s capu la i sa t t r ibu tab le t o i t s f r ag i l i ty .

P ercen tage r epre sen ta t ion : T h i s c a lcu la t ion ( f o l low ing A nd rew s 1 990 ) a l low s c ompa r i son a mong t h e v a r iou s

A pp l ica t ion o ft h e se b r eakage g u ide l ine s t oa n a r chaeo log i -

s ke le ta l e l emen t s .

c a l s amp le c anno t b e d i r ec t . S pec if i ca l ly , b r eakage p a t ter n s

T he MN I p e r s i ze c l a s s i sd e r ived f r om

t h e t o ta l n umbe r ( f ragmen ta ry o r w ho le ) o fe ach e l emen t i s d i v ided b y t h e n umbe r o ft h a t e l emen t e xpec ted p e r i n d iv id -

a mong r oden t i n c i so r s a r e l i ke ly t o b i a s r e la t ive p a r t r e p r e s en ta t ion p e r cen tage s ( Tab le s 2 . 3 -2 .6 ) . T he se e l emen t s a r e

u a l . T h i s a moun t i s1 2 f o r mo la r s , 4f o r i n c i so r s , a nd 2 f o r

h igh ly d u rab le a nd i d en t if iab le , y e tm ino rb r eakage w i l l s im p l yp roduce mo re f r agmen t st h a ta r ed u rab le a nd i d en t if iab le . O the rt oo th f r agmen t s a l so f i tt h is p a t ter n , i nt ha t t h ey m ake

a l l r ema in ing m a jo r a ppend icu la r e l emen t s ( hume r i , s c apu l a e , e t c . ) . T he l a rge s t t o ta l a mong a l l t h e se e l emen t s i st h e MN I . E ach e l emen t i st h en s ca led a ga in s t t h e MN I b a sed u pon t h e mo s ta bundan te l emen t :

u p av e ry s ma l l p o r t ion o f u n iden t if i ed m ic rof auna l f r ag m en t s : at i ny f r agmen t c an a tl e a s t b e i d en t if ied a s at o o th .

2 4

Many t i ny n on -den ta l f r agmen t s r ema in u n iden t if ied . E l e m en t sw i th l a rg e , d u rab le a nd i d en t if iab le u nf u sed e p iphy se s o r at e ndency t of r agmen t i n to mo r e t h an o ne ( p rox ima l a nd d i s ta l ) i d en t if iab le p i ece ( hume r i , f emo ra a nd t i b io -f ibu la e ) a l so i n c r ea se t h e i r r e la t ive p a r t r e p r e sen ta t ion . E l emen t s l a ck ing d i agno s t ic c ha rac te r s ( u lnae , r a d i i ) b i a s t h e i r p a r t r e p re sen t a t ion n ega t ive ly . T he se d i f f e r ence s a r e c on s ide r ed i ni n te rp r e t a t ion o fr e la t ive p a r tr e p r e sen ta t ion .

m ic ro t ine m and ib le s . M ic ro t ine t o o th l o s s s im i la r ly m ake s m ax i l lae mo re s u scep t ib le t oc r u sh ing . I nt h e Med ium s i ze c l a s s ( t he r ema in s o fw h ich a r e d om ina ted b ym ic ro t ine s p e c i e s ) , m and ib le s a r e r e p r e sen ted b y 5 3 1 t o ta l p i ece s . O n ly 1 2 ( 2 .3 % o ft h e t o ta l ) w e r e c omp le t e a nd u nb roken . I nt h e L a rge s i z e c l a s s ( c o r re spond ing t ot h em ic ro t ine g enu sA r v i c o la ) , 1 3 o f2 08 ( 6 .3% ) w e r e c omp le t e . Mand ib le s a mong t h e Sma l l s p ec ie s ( c o r r e spond ing t o s h r ew s , w h ich h ave r oo ted t e e th ) 1 2o f6 6 ( 18 .2% )w e r e c omp le te . O n ly 1o f5 2 ( 1 .9% )o fT a lpa m and ib le sw e r e c omp le te . Wh i le t h i s g enu s h a sr oo ted t e e th , i t sm and ib le sa r er e la t ive ly l o ng ,g r a c i le a nd h ence e a s i ly f r agmen ted .

T he s k e le ta le l emen tr e p r e sen ta t ion p a t te rn a mong t h e Sma l l , Med ium a nd L a rge c l a s se s i n d ica t e s h i gh r a t e s o f s u r v iva l a mong e v en t h e mo s t f r ag i le e l emen t s . T he h umeru s i s h i gh ly r e p r e s en t ed ( 78 .8% , 5 6 .7% a nd 5 0 .3% , r e spec t ive ly ) a mong a l t h r ee c l a s se s . T he e x t r eme ly f r ag i le a nd l e s s i d en t if iab l e s c apu la w a s f ound i ns i gn if i can t a moun t s i nt h e M ed ium a nd L a rge c l a s se s ( 1 9 .6% a nd 1 9 .4% ) . T h i s r e la t i ve ly h i gh r e p r e sen ta t ion i si n d ica t ive o fv e ry l i gh t d amag e t os c apu la e . I na lmo s t a l c a se s t h e mo r e d u rab le d i s ta l e x t r em i ty o f t h e s c apu la w a s i d en t if i ed , c ompa r ed w i th t h e p r ox ima l e p iphy s i s , p rox ima l e nd , s haf t , a nd d i s ta l e nd o f f r agmen t ed h ume r i . T he h umer u s t h e r ef o re i sr e la t ive ly o ve r r ep r e sen t ed a s c ompa red w i th t h e s c apu la , s i nce t h e f o rme r c a n s e pa ra te /f ragmen t i n to mu l t ip le c omponen t p i ece s ,a l c om ing f r om as i ng le o r ig ina le l emen t .

P e lve s d i d n o t f a r e w e l l g ene ra l ly .

Al a ck o f f u s ion c om -

b i ned w i th l e s se r i d en t if iab i l i ty o ff r agmen t s m ade t h i s e l e m en t l e s s v i s ib le . B reakage a tt h e a c e tabu lum w a s c ommon , t h ough i ft h i s a r ea h ad f u sed s ome t ime s t h e a c e tabu la r p o r t i on a l one s u rv ived . T h i s e l emen t w a s r a r e ly r e cove r ed i na c omp le te s t a t e : 1o f 8( 1 2 .5% ) i nt h e S ma l l c l a s s , 2o f 1 40 ( 1 .4% ) i nt h e Med ium c l a s s , a nd 0o f7 9 i nt h e L a rge c l a s s w e r er e cove r ed c omp le te . Sma l l e l emen t s ( r ib s , v e r teb ra e / s t e rneb ra e , m e tapod ia l s a nd p ha lange s ) w e r e r e cove r ed i nr e la t ive ly l a rge a moun t s . T h i s m ay b e a t t r ibu tab le t o t h e s i z e o ft h e s c r een s e mp loyed i n l a bo ra to ry p roce s s ing ( 0 .25 mm ) . T he se e l emen t s a r e t y p i c a l ly r e cove r ed i n l a rge a moun t s i nt h e a s semb lage s p ro d uced b ym any s p ec ie s o fo w la nd a r er a r e ly p r e se rved i nt h e a s semb lage s p r oduced b y d i u rna l r a p to r s a nd c ar n ivo r e s ( d i scu s sed b e low ) .

C ompa r i son o f h ow f r equen t ly t h e m o r e d e l ica te s c apu la s u rv ived w i th t h e r obu s th ume ru s i s mo r e i l um ina t ing w hen c on s id e r ing t h e se f a c to r s o fi d en t if iab i l i ty , r e cove rab i l i ty a nd s u rv ivab i l i ty o ff r agmen t s . When c ompa r ing t h e t o ta l n um b e r o fd i s ta l s c apu lae ( n =2 50 ) w i th p rox ima l h ume r i ( n= 2 83 , d i s r ega rd ing u nf u sed e p iphy se s ) i nt h e Med ium c l a s s , t h i s d e l ica t e e l emen t s u rv ive s a t8 .3% t h e r a te o ft h e l a t te r . T h i s p a t t er n i sn o t r e pea ted i nt h e S ma l l c l a s s , p e rhap s b e c au se e ven t h e d i s ta l e x t r em i t ie s o fs h r ew s c apu lae w e r e t o o d e l ica t e t os u rv iv e i na n i d en t if iab le s t a te . T he se m igh t a l so b e s ma l l e nough w hen f r agmen t ed t op a s s t h rough a 1mm m e sh s c r een a nd b e l o s td ue t ot h e a na ly t ica lp rocedu re u s ed . T h i s p a t t er n i s , h oweve r , r e pea t ed w i th t h e d i s ta l s c apu la e a nd p rox ima l h ume r i o ft h e L a rg e s i z e c l a s s , w h ich c o r r e s pond t ot h et a xon A r v i co la . I nt h i s c a se , t h en umbe ro fd i s t a l s c apu la e( n =9 1) e xceed st h en umbe ro fp r ox ima lh ume r i ( n = 8) , o r 1 0 3 .4% . G r ea t d e s t ruc t ive f o r ce s u pon t h i s m i c r of auna l a s semb lage w ou ld h ave c au sed a m uch g r ea t e r d i sc repancy b e tween i n he r en t ly m o r e d u rab le a nd l e s s d u ra b l e e l em en t s . T he t a phonom ic a g ency r e spon s ib le f o r t h e i r a c cumu la t ion t h e r ef o r e c au sed v e ry l i t t le d amage d u r ing t h i s p r oce s s .

D IGEST IVE CORROS ION E v idence o f d i ge s t ive c o r ro s ion w a s e xam ined f o r o n a l e l emen t s i d en t if i ed b yt h e a u tho r . S u r f a ce c o r ro s ion m ay b e a t t r ibu t ed a lmo s t e n t i re ly t op r eda to r a c t ion , s i nce t h e p H o f e a ch s ed imen t l o t i nt h i s l i me s tone c ave v e s t ibu le w a s a bove 7 . 0 . Among a l t e e th i d en t if i ed i nt h i ss t udy ( n =7 486 ) , o n ly 2 77 ( 3 .7% ) h ad t r ace s o f d i ge s t ive c oro s ion , t y p ica l ly r o und ing t h e o cc lu sa l s u rf ace a nd t h e s i d e s e xpo sed a bove t h e s o cke t s . T h i s t y pe o fd amage i sm o r e l i k e ly t oo ccu r o n t e e th , s i nce t h e se h ave t h e h i ghe s t m ine ra l c on t en t a mong s k e le ta l e l emen t s ( H i l l son 1 986 ) . T hey a r e a l so t h e m o s t e xpo sed s ke le ta l e l emen t s : t h ey e n t e r ap r eda to r ' s s t oma ch e xpo sed t o d i ge s t ive a c id , w h i le o n ly a f t e r i n i t ia l d i ge s t ion d o o t he r e l emen t s ( o the r t h an t h o se e xpo sed t h rough b r eak a ge ) b ecome e xpo sed t oa c id c o r ro s ion . Among t h e i d en t i f i ed l a rge n on -den ta l r ema in s ( t he r eby e xc lud ing f o o t e l e m en t s , v e r teb ra e a nd r i b s ) , o n ly 1 4 o f 9 84 1 ( 0 .14% ) h ave v i s ib le a c id c o r ro s ion . T h i s a ppa r en t l a ck o f c o r ro s ion o f c ou r se m ay h ave b een n ega t ive ly b i a sed , s i nce c o r roded e l emen t sa r em o r e l i ke ly t ob ed e s t royed o rr e nd e r ed u n iden t i f iab le t h an n on -co r roded e l emen t s . C l ea r ly , h oweve r , d i g e s t ive c o r ro s ion i sn o t as i gn if i can t f e a tu r e o ft h i s m ic ro f a una la s semb lag e .

T he g en e r a l b r eakage c ha rac t e r i sv e ry l i gh t d amage , e ven w hen c on s id e r ing p o s tdepo s i t iona l t r amp l ing a nd t h e e f f ec t s o fe xcava t ion a nd w e ts c r een ing . P rox ima la nd d i s ta lh ume r i a nd f emo r a a nd p rox ima l t i b io -f ibu la e s u rv ived w e l l . O the r e l emen t s s u rv ived l e s s w e l l . Mand ib le s , t h ough n ume rou s , w e r e r a r e ly c omp le te . T h i sb r eakage o fs u ch n o rma l ly d u ra b l e e l emen t s i se xp la ined b y t h e p ropen s i ty o fr oo t le s s m i c r o t ine m o la r s t o f a l l f r om t h e i r s o cke t s . M ic ro t ine t o o th l o s s i sf r equ en te ven a mong moder n o w l p e l le t a s semb lage s . T h i s l o s s l e ave s al a rge g ap a l ong t h e m id l ine o ft h e m and i b l e a nd m ak e s i tmo re s u scep t ib le t op o s td epo s i t iona l c r u sh i n g . T h e s ame i sn o t t r ue f o r mu r id o r s o r ic id m and ib le s , w h ich ( t hough f ew i nn umbe r ) w e r e l e s s o f t en c r u shed t h an

P REY ACT IV ITY P ATTERNS T he a c t iv i ty p a t ter n s o ft h e r oden t s a nd i n sec t ivo re s r e cov e r ed ( Chap te r 3 ) a r e o ve rwhe lm ing ly n oc tu rna l , c o inc id ing w i th mo s t o w l h un t ing a c t iv i ty . T h i s p a t t e rn i st y p ica l a mong s ma l l m amma l a c cumu la t ion s , s i nce n oc tur na l b e h av io r a mong s ma l l m amma l s i s ag ene r a l e vo lu t iona ry a d -

2 5

a p ta t ion f o r p reda to r a vo idance . Many o ft h e se s pec ie s f a l l p r ey t od i urna l p reda to r s a sw e l l w hen t h e ir a c t iv i ty p a t te rn s o ve r lap d u r ing t h e c r epu scu la r h ou r s . T he se c r epu scu la r s pec ie s i n c lude C roc idura r u s su la , S orex c o rona tu s , M ic ro t u sa g re s t i sa nd P i tymy s l u s i tan icu sa l o fw h ich a r e c ommon

COMPAR ISON OF P REDATOR TYPES T he d amage p a t ter n o f t h e se r ema in s p r ec lude s t h e i r c on c en t ra t ion b y t e r re s t r ia l c ar n ivo re s i n c lud ing c an id s , m u s te l i d s o rf e l id s . Many s pec ie s o f mamma l ian c a rn ivo r eh ab i tu a l ly a ccumu la te s ca t s i n l a t r ine a r ea s , s ome t ime s n ea r t h e i r

p r ey o fb uzza rd s ( Bu teo b u teo ) a nd b a rn o w l s i nt h e mode rn e nv i ron s o fE l J uyo ( Chap te r 4 ) . P a t te rn s o fb one r e cove ry a nd d amage a r e much mo re c ompe l l ing e v idence o fo w l a c -

d en s

o r t o ma rk

t e r r i to r ies

1 983 :290 ; S tua r t 1 977 ) .

( s ee A nd rews

a nd E van s

Me l le t ( 1974 ) c o ined t h e t e rm

" cop rocoeno s is " t oi n d ica te af o s s i l a s semb lage d e r ived f r om a n a ccumu la t ion o f mamma l ian c a rn ivo r e s c a t s . A nd r ew s

c umu la t ion t h an n oc tu rna l i ty o fp rey .

a nd E van s ( 1983 ) e xam ined t h e s ca t c on ten t s o f ab a t te ry o f s pec ie s f r om t h ree f am i l ie s o f mamma l ian c a rn ivo r e . T he se w e re t h e V ive r r idae ( r ep re sen ted b y mongoo se khneum ia

A n a ppa ren t e xcep t ion t o t h e c o r re spondence o f o w l a nd p r ey a c t iv i ty i st h e p re sence o f s ma l l a moun t s o f mo le s ( Ta lpa s pp . ) i nt h i s a s semb lage . Mo le s , b e ing f o s so r ia l , t e nd

a l b icauda a nd g ene t G ene t ta g ene t ta ) , C an idae ( r ed f ox V u /pe s v u /pe s , b a t -ea red f ox O tocyon m ega lo t i s , a r c t ic f ox

t o e xc lude t h emse lve s f r om p reda t ion b y a v ian p r eda to r s . T he moder nb a rn o w l p e l le t s amp le s e xam ined f r om t h e i m -

A lopex l agopu s a nd c oyo te C an i s l a t ran s ) a nd Mu s t e l idae ( p ine m a r ten Mar te s ma r te s ; w ea se l , s t oa t a nd p o leca t Mu s te la s pp . ) . O ve ra l l b one b r eakage w a s much g r ea te r f o r a l l t h e se p r eda to r s pec ie s a s c ompa r ed t oo w l -p roduced a s -

m ed ia te e nv i ron so fE lJ uyo ( Chap te r4 )y i e lded o n ly aT a lpa e u ropaea MN I o f 3o u to f at o ta l MN I o f9 34 v e r teb ra te p r ey i t em s . G lue ( 1974 ) f ound t ha t T a lpa e u ropaea m ade u p l e s s t h an 1 % o ft h e d i e t ( 26 o f4 7 ,865 v e r teb ra te p r ey i t em s ) o f t h e b a rn o w lt a ken f r om 1 88 p e l le t s amp le s f r om B r i ta in a nd I r e land . I nc on t ra s t , at o ta l o f3 00 t e e th o fT a lpa s pp . w e r e i n c luded i nt h e t o ta l s amp le o f7 , 862 i d en t if i ed t e e th ( 3 .8% o ft h et o ta l ) . P a r to ft h i s a ppa ren t o ve rabundance i se xp l ica b l e i nt e rm s o ft h e g rea te r n umbe r o ft e e th p e r i n d iv idua l i n t h i s g enu s o f mo le . T a lpa h a s t h e f u l l p l acen ta l m amma l ian c omp lemen to f1t e e th p e rd em i -mand ib le /max i l la ( 3 / 1/4 /3 ) .

s emb lage s . P ropor t ion s o f c omp le te h ume r i r e la t ive t ot h e t o ta l r e cove red a mong t h e se p reda to r s r a nged f r om 0% t o 4 3% , u l nae r a nged 0% t o 5 7% , f emo ra 0% t o2 5% , a nd t i b i a e 0% t o 5 7% ( And rew s a nd E van s 1 983 :295) . O ve ra l l b one b r eakage r a nged f r om mode ra te ( g ene t ) t o v e ry g r ea t ( c an id s ) , a nd r ound ing o rp o l ish ing o fe l emen t s r a nged f r om l i t t le ( ba t -ea red f ox )t og rea t( r ema in ing c an id s ) . Mamma l ian p reda to r s a l so d amage a nd d i ges t s ma l l p r ey b one s much mo re t h an d o o w l s ( Day 1 966 ; G r eenwood 1 979 ; L ock ie 1 959 ; R ober t son a nd Whe lan 1 987 ; S t uar t 1 977 ; Weave r a nd H of fman 1 979 ) . K o r th ( 1979 ) c ompa red s ma l l mamma l r ema ins f r om c oyo te s c a t s t ot ho se e x t ra c ted f r om s amp le s o f g rea t h o rned a nd b a rn o w l p e l le t s . T he c oyo te -de r ived s amp le i n c luded many s ha rp ly b roken b one f r agmen t s , a l though ma jo r l imb b one s o f t en s u rv ived w ho le : 6 2% o fa l r e cogn izab le h ume r i a nd f emo ra w e re u nb roken ( Ko r th 1 979 :248 ) . Sma l le r mamma l s p ec ie s s ke le ta l e l e -

A l l t h ese t e e th a r e g ene ra l ly i d en t if iab le , e ven t h e t i ny i n c i s o r s i ff ound i n tac t i nt h e i r s ocke t s . T h i s c on t ra s t s w i th t h e t e e th o fa l t h er oden t sr e cove red a tE lJ uyo , w h ich h ave o n ly t h e e vo lu t iona r i ly r educed p a t te rn o f 1 / 0 /0 /3 , o r 1 6 t e e th p e r i nd iv idua l . T h i s a moun t i sw innowed t o 1 2 mo la r s p e r i n d i v idua l w hen o ne c on s ide r s t h e g ene ra l u n iden t i f iab i l i ty o f r oden t i n c i so r s b eyond s i ze c l a s s . T a lpa t h e r ef o r eh a s n ea r ly 3t ime s a sm any p o ten t ia l ly i d en t if iab le t e e th w hen c ompa red w i th am embe r o ft h e g enu s M icro tu s , w ho se t e e th a r e r e c ove r ed i ng r ea tn umbe r s f r om E lJ uyo a nd o t he rC an tab r ian s i te s .

m en t s g ene ra l ly w e re mo re l i ke ly t oe s cape d amage , p roba b l y b ecau se l e s s ma s t ica t ion w a s n ece s sa ry f o r t h e i r c on s ump t ion . Mand ib le s w e re n o ted r a r e ly t or e ta in t h e i r o r ig i n a l c omp lemen t o f t e e th , i n c on t ra s t t o m and ib le s d e r ived f r om o w lp e l le t s .

A n a dd i t iona l 1 , 065 T a lpa r ema in s b e s ide s i d en t if i ed t e e th w e re r e cove r ed f r om at o ta l o f2 5 ,204 f r om a l l e ve l s o fE l J uyo . A n o ve r rep re sen ta t ion o f mo le p o s tc ran ia l e l emen t s i s e a s i ly e xp l icab le : t h e se e l emen t s a r e h igh ly d i s t inc t ive , i d en t i f iab le , a nd i nt h e c a se o ft h e f o re l imb e l emen t s ( pha lange s ,

D eg ree o f d i ge s t ive c o r ro s ion o n s k e le ta l r ema in s i sg r ea t a mong C a rn ivo ra -p roduced a s semb lage s . A nd r ew s a nd E van s ( 1983 ) f ound c o r ro s ion o fb one s a nd t e e th b y d i ge s t i ve a c id w a s c ommon a mong t h e m amma l ian c ar n ivo r e s pe -

c a rpa l s , m e taca rpa l s , r ad iu s , u lna a nd h ume ru s ) e x t reme ly r obu s t d ue t o t h e i r e vo lu t ion a s s p ec ia l iz ed e xcava t ion a p p endage s . P o s tc ran ia l e l emen t s w e re i d en t if i ed o n ly t os i z e c l a s s f o r t h e r ema inde r o ft h e s ma l l i n sec t ivo r e a nd r oden t s pec ie s r e cove red , w i th t h e e xcep t ion o f w a te r v o le s

c i e s e xam ined . Wh i le mongoo se -con sumed p rey b one s a nd t e e th s howed n o s uch a l te ra t ion , t h e r ema in ing s p ec ie s

( Ar v ico la t e r re s t r i s ) wh ich a r e mo s t ly d i s t ingu i shab le b y t h e i rl a rge rs i ze .

r a nged f r om l i t t le t o s eve re d amage .

A nd r ew s a nd E van s

( 1983 :301) a l so s umma r ize t h ec ha ra c te r i s t ic so ff e l id -a l t e red p r ey b one s , w h ich w e re f ound t oi n cu re x t r eme b reakage a nd c o r ro s ion a nd g r ea t r ound ing a nd p o l i sh ing . Al e s se r d eg ree

Mo le s d o f a l l t o o w l s i n s i gn if ican t n umbe r s d u r ing o ne p ha se o ft h e i r y ea r ly c yc le : t h e l a te s umme r d i spe r sa l o fj u -

o fb one b reakage a nd c o r ro s ion a mong s ma l le r mamma l ian p r ey w a sn o ted .

v en i le s w ho mu s t l e ave t h e i r n a ta l b u r rows t oe s tab l i sh t h e i r o wn . A t t h i s t ime mo le s t r ave l ing o n t h e s u rf ace a r e e a sy

T he d amage p a t te rn o f t h e E l J uyo r ema in s a l so p r ec lude s

p r ey f o r o w l s . T he p resence o f as l igh t o ve rabundance o f mo le s m ay i n d ica te t h a t t h e s ea son o fo w l o ccupa t ion o ft h e c ave t e nded t oi n c lude l a te s umme r .

t h e i r a ccumu la t ion b y d iurna l r ap to r s s u ch a sh awk s , f a lcon s o r e ag le s . D iur na l r ap to r s l e ave f a r f ewe r r e cogn izab le b one s p e r c on sumed p rey i t em i nt h e i r p e l le t s t han d o o w l s . E a t ing h ab i t s a r e p a r t ia l ly r e spon s ib le f o rt h i s l ow b one r ep r e sen ta t ion . H awk s a nd o t he r d iu rna l r a p to r s t e nd t ob e f a r

2 6

c omp le t ene s s m ean ( 7 5% ) t h an t h e b ar n o w l s s h e s t ud ied . T he se r e su l t s m ay i n d ica te t h a t s e pa ra t ion o f s ho r t -ea r ed f r om b a rn o w l a s semb lage s b a sed o n ly u pon s k e le ta l p a r t r e p r e sen ta t ion a nd b r eakage m ay n o t b e p o s s ib le . F eed ing b ehav io r i ss im i la r t o b a rn o w l s , w i th s ma l l m amma l p r ey u sua l ly s wa l lowed w ho le a nd l a rge r p r ey t or n i n to p i ece s f i r s t ( Ch i t ty 1 938 :270 ) . S ho r t -ea r ed o w l s a r e o pen -coun t ry d we l l ing , h oweve r , a nd t h ey i n va r iab ly r oo s t i ng r a s s n e s t s o n t h e g round n ea r ma r shy a r ea s ( Bu r ton 1 973 ) . T hey a r e a l so p r ima r i ly d i ur na l i nt h e i ra c t iv i ty .

m o r e f a s t id iou s e a te r s t h an mo s t o w l s , o f t en c a ref u l ly f l en s i n gp rey o ft h e i re d ib le p o r t ion o rt e a r ing i ti n to s ma l lc hunk s ( C la rk 1 972 ;E ina r sen 1 956 ) . T he se m e thod so ff e ed ing t e nd t o l e ave p r ey b ones b eh ind a tt h e s i t e o f e a t ing ( t hu s n o t l e ad ing t o a c cumu la t ion i n as i ng le l o ca l i ty ) o r t o b r eak b one s t o am uch g r ea te rd eg r ee t h an w ou ld s wa l low ing p r ey w ho le . T he l owe r d i u rna l r a p to r g a s t r ic p H l e ad s t o i n c r ea sed d i ge s t ion o f b one ( Duke e t a l . 1 975 ) . H of fman ( 1 988 ) f e d c ap t ive s pec ie s o fo w l a nd d i u rna l r a p to r ac on t r o l led d i e t o fh ou se mou se (M u s m u s cu lu s ) . P e l le t s o ft h e h awk s p ec ie s t e s ted ( r ed ta i led h awk B u teo j ama i cen s i s , r o ugh legged h awk B u teo l a gopu s , a nd s p a r row h awk F a l co s par ve r iu s )a ve raged l e s st h an o ne i d en t if iab le b one f o re a ch m ou se e a ten . N o t as i ng le c omp le t e s k e le ta l e l emen t w a s r e cove red f r om t h e p e l le t s o fa ny o ft h e se s p ec ie s , a nd m o s t o f t h e f r agmen t s w e r e i n v e ry p oo r c ond i t ion . A nd r ew s ( 1 990 :38 ) e xam ined t h e p e l le t s o f s e v e ra l c ommon d i ur na l r a p to r s a nd f o und s ke le ta l p a r t r e la t ive a bundance t of a l l t o l e ve ls m uch l owe rt h an a mong b ar no w l s , g ene ra l ly l e s st h an h a lf .

E ag le o w l s c an l e ave b eh ind b one s w i th s im i la r d eg r ee s o f b r eakage , t h ough t h i s s p ec ie s t e nd s t o t e a r i t s p r ey i n to s ma l le r p i ece s b ef o r e c on sump t ion ( E ina r sen 1 9 56 ; H owa rd 1 958 ) . T he p e l le t s o f t h e c l o se ly r e la ted a nd e co log ica l ly e qu iva len tN ew Wo r ld g r ea t h or n ed o w l ( Bubo v i rg in ianu s ) h ave b een s t ud ied i n mo re d e ta i l . K u sme r( 1 990 ) f o und t h a t g r ea t h or n ed o w l p e l le t s y i e ld p r ey b one s w i th v ery l i t t le b r eakage a nd h i gh r e p re sen ta t ion o f e l emen t s . Mean p a r t r e p r e sen ta t ion f o r n i ne s ke le ta l e l emen t s w a s 8 2% a nd 8 7% f o r t wo s amp le s o fg r ea t h o rned o w l p e l le t s . O ve ra l l b ar n o w l p e l le t p a r t r e p re sen ta t ion f o r as amp le f r om t h e N egev D e se r t ( I s ra e l ) w a s 8 7 .9% ( Pok ine s a nd K e rb i s P e t e rhan s 1 998 ) , w i th ag r and m ean o f8 2 .5% f o r t h e b a rn o w l p e l le t s t ud ie s s umma r iz ed b e low . K o r th ' s ( 1 979 ) r e su l t s i n d ica te a n o ve ra l l p a r t r e p r e sen ta t ion m ean o f 8 4 .4% f o r t h e s ame n i ne e l emen t s i nh i s s t udy o fg r ea t h o rned o w l p e l le t s , a nd H off inan ' s ( 1 988 ) r e su l t s i n d i ca t e am ean o f 8 7 .3% f o r h i s s amp le f r om t h i s s p ec ie s . D od son a nd Wex la r ( 1 979 ) n o ted as im i la r ly h i gh p a r t r e p r e sen ta t ion m ean ( 8 5 .2% ) f o r t h e g r ea t h or n ed o w l p e l le t s t h a t t h ey e xam ined , b u t f o und a much l owe r o ve ra l l m ean o f c omp le t e e l emen t s ( 3 4 .0% ) . Moon ( 1940 :46 1) f ound s im i la rb one w e igh tp e r cen tage s f o r p e l le t s o fb ar n o w l s a nd g r ea t h or n ed o w l s : 3 8% a nd 4 0% , r e spec t ive ly .

T he r e cove ry a nd d amage p a t t e rn s o ft h e E lJ uyo m ic rof au n a l a s semb lage a r e i ns t a rk c on t ra s t t oa s semb lage s d e r ived f r om t e r r e s t r ia l c a rn ivo r e s o r d i ur na l r a p to r s . T he l a rge a moun t o f i d en t if iab le p o s t c ran ia l r ema in s , m any o f t h em l a rge ly i n ta c t , i si nc omp le t e c on t r a s tt ot h e c ha ra c te r i s t ic so f t h e se p r eda to rt y pe s d i scu s sed a bove . G a s t r ic c o r ro s ion w a s a l so v e ry l ow a mong t h e se r ema in s . T h i s p a t te rn o fd amage a nd r e cove ry i se n t i re ly c on s i s ten t w i th m any o w l -p rodu ced m ic rof auna la s semb lage s . S ome o w l s p ec ie s may b e q u ick ly e l im ina t ed a s p o t en t ia l a c cumu la to r s o f t h e E l J uyo a s semb lage . R a czyn sk i a nd R up rech t ( 1 974 ) f ound t h a t l o ng -ea r ed o w l s ( A s io o t u s ) d e s t royed p r ey b one t o am uch g r ea te re x ten tt h an d i dt h e b a rn o w l s s t ud ied . B one d e s t ruc t ion w a s m uch h i ghe r a mong t h e j uven i le s o ft h i s s pec ie s . L ong -ea r ed o w l s , l i k e t awny o w l s , h un t a nd r oo s t i nt r ee s ( Bu r ton 1 973 ) , m ak ing ac ave a tt h e b a se o f as i nk h o le av e ry u n l ike ly r o o s t ing s i t e . L i t t le o w l s ( A thene n oc tua ) d o n o t c r ea te l a rg e b one a c cumu la t ion s d ue t o t h e i r d i e ta ry f o cu s u pon i n ve r t eb r a t e s . S nowy o w l ( N y c tea s cand iaca ) , g r ea t g r ey o w l ( S t r ix n ebu lo sa ) , U r a l o w l ( S t r ix u ra l en s i s ) , T engma lm ' s o w l ( Aego l iu sf une reu s ) , S cop so w l( O tu ss cop s ) ,h awk o w l( S u rn ia u l u la ) a nd p ygmy o w l( G lauc id ium p a s se r inum )m ay b e e l im ina t ed b ym u l t ip le f a c to r s s u ch a ss pec ia l iza t ion o fd i e t , r a nge , h ab i ta t r e qu i r e m en t s ,a nd b one a ccumu la t ion h ab i t s .

B e ing much l a rge r t h an b a rn o w l s ( 3 ,000 -4 ,000 g v s . 3 50 6 00 g , r e spec t ive ly ) , e ag le o w l s h ave a d i e t a ve rag ing a much h i ghe r b ody m a s s p e r i n d iv idua l t a ken . H edgehog s , l a gomo rph s a nd l a rge b i rd s a r e t a ken f r equen t ly ( Bu r ton 1 973 ; D onäza r a nd C eba l lo s 1 989 ; D onazä r e t a l . 1 989 ; E ina r sen 1 956 ; F rey a nd Wa l t e r 1 986 ; G oodman 1 990 ; O r s i n i 1 985 ) . B ar n o w l s d o n o t f o cu s u pon p r ey g r ea t e r t h an 1 00 gi ns i z e a nd c oncen t ra t et h e i ra t t en t ion u pon p r ey i nt h e 1 -50 g r a nge ( Go sz czyn sk i 1 977 :409 ) . A n a s semb lage d om ina ted b y s ma l l r oden t s a nd i n sec t ivo r e s i st h e r ef o r e u n l ik e ly t oh ave b een a c cumu la t ed b ys u ch al a rge s p ec ie s o f o w l .

C e r ta in c ha ra c t e r i s t ic s o f t h e E l J uyo a s semb lage p o in t t o b a rn o w l s a s i t sp r inc ipa l a c cumu la to r s . S ome o w l s p ec ie s w i th E u ropean r a nge sp roduce a s semb lage sw i th c ompa rab le a moun t so fp r ey b one b r eakage : b a rn o w l s( T y to a l ba ) , e ag le o w l s ( Bubo b ubo ) a nd s ho r t -ea r ed o w l s ( A s io f l ammeu s ) . Wh i le s ome d i f f e rence s i nb r eakage a nd c o r ro s ion p a t t er n a r e e v id en t a mong m od e rn p e l le t c o l lec t ion s f r om t h e se p r eda to r s p ec ie s , t h e se n uance s a r e l i ke ly s ubme rged i na n a r cha eo log ica l m ic rof auna l a s semb lage t h a t h a s u nde rgone p o s tdepo s i t iona lb r eakage . T he se t h r ee s p ec ie s ,h oweve r ,a r e d i s t ingu i shab le b yo the ra s pec t s o fb ehav io r . K u sme r( 1 990 ) f o und t h a t s ho r t -ea red o w l ( A s io f l ammeu s ) p e l le t s ( MN ! 1 9 )h ad as l igh t ly g rea t e rp a r tr e p r e sen ta t ion m ean ( 9 1% )a nd

I na dd i t ion ,t h es e t t ing o fE lJ uyo a r gue s a ga in s ti t su s ea sa n e ag le o w l r o o s t . T h i s ( l a rge s t ) s p ec ie s o f o w l w ou ld h ave f o und g r ea td i ff i cu l ty f i t t ing i t sl a rge w ing span i n to t h es ma l l c onf ine s o ft h e a va i lab le r oo s t ing s p a ce : i t s w ing span i s1 . 5 m , w i th ab ody l e ng th o f0 . 5 m . I nc ompa r i son , E u ropean b a rn o w l s ( T y to a l ba a l ba ) h ave aw ing span a bou t 7 60 mm a nd ab ody l e ng th u nde r4 00 mm ( Bu r ton 1 973 ) . Wh i le t h ey d o r oo s t i nc ave s a nd f i s su re s , s u ch l ow p l a ce s a s E l J uyo ( l oca ted a tt h e b a se o f as i nkho le ) a r e a vo ided . E ag le o w l s c hoo se r oo s t s f o r ac l ea r f i e ld o f v i ew a nd h i gh l o ca t ion . T h i ss p ec ie sa l so s hun sh uman c on ta c ta nd a vo id sr oo s t ing i n h uman h ab i ta t ion s( M i ld co la 1 983 ) .

2 7

T ab le 2 . 7 : S umma ry o fM ic rove r t eb ra te T aphonomy . A ccumu la t ing P reda tor

A s semb lage

Sma l l

D iu rna l

T awny

L ong -

E ag le

S ho r t -

B a rn

E l

C a rn ivo r e

R ap to r

Ow l

e a red

Ow l

e a red

Ow l

J uyo

V e ry L ow

Charac ter

Ow l

D eg ree o f B reakage

V e ry H igh

V e ry H igh

H igh

H igh

C omp le tene s so f

V e ry

V e ry

L ow

L ow

S ke le ta lE lemen t s

L ow

L ow

D eg ree o fD ige s t ive C o r ro s ion

V e ry H igh

V e ry H igh

L ow

L ow t o H igh

Mode ra te

S i ze C la s so f Mo s tP r ey

V a r iab le

A c t iv i ty P a t te rn o fP rey H ab i ta to f

Ow l V e ry L ow

V ery L ow

V e ry L ow

V e ry

V e ry

V e ry

V e ry

H igh

H igh

H igh

H igh

L ow

V e ry L ow

V e ry L ow

V e ry L ow

V e ry L ow

L ow

Mode ra te

V e ry H igh

V ery H igh

V e ry H igh

H igh

V a r iab le

5 100 g

5 100 g

3 0t o 3 000+ g

5 100 g

5 100 g

5 100 g

V a r iab le

D ium .

N oc t .

Noc t .

N oc t .

D ium .

N oc t .

N oc t .

V a r iab le

V a r iab le

F o re s t

F o re s t

Mo s t

O pen

Mo s t

O pen

P rey S pec ia l iz ed

V a r iab le

V a r iab le

N o

N o

N o

No

N o

N o

D en s /Roo s t s i nC ave s

S ome

S ome

N o

N o

Y e s

No

Y e s

Y e s

D ens i ty o fM ic rof auna l A ccumu la t ion

P reda to r

2 8

s t oa t s (M u s t e la s pp . ) d o s ome t ime s c a che u nea ten s ma l l v e r t e b ra te p r ey i n ad en f o r l a te r c on sump t ion ( Ok sanen 1 983 ) . I fn o t r e cove r ed b y t h e a ccumu la t ing p r eda to r s u ch c o l le c t i on s c ou ld l e ave a c oncen t ra t ion o f h i gh ly i n ta c t b one s w i th in t h e a r ea o fa n a r chaeo log ica l s i te . S uch b ehav io r i s h i gh ly u n l ike ly t o h ave c on t r ibu t ed s i gn if i can t ly t o t h e E l J uyo a s semb lage d ue t ot h e l a ck o f s u i tab le d enn ing a r ea s w i th in t h e v e s t ibu le o ft h e c ave , a nd t h e l a ck o fa ny f i nd s o f h i gh ly c omp le t e s ma l l v e r teb ra t e s k e le ton s c oncen t ra t ed i na v e ry s ma l l a r ea . T he l a rge m ic rof auna l c oncen t ra t ion a tt h e b a se o fT umu lu s 4i nt h e S anc t ua ry f e a tu re o fL eve l 4w a s a c cumu la t ed a tt h e b a se o f al a rge o pen p i t . T h i s l o ca t ion w ou ld n o th ave b een as u i tab le s ma l l c ar n ivo r ed enn ing a r ea , n o ra r ev e ry l a rge c on cen t ra t ion sp roduced i nt h i sm anne r .

O the r s p ec ie s o fo w l f o und i nE u rop e c an b e e l im ina ted a s a c cumu la to r s o fm ic rof auna i nE lJ uyo o n mu l t ip le g round s . T awny o w l s ( S t r ix a l uco ) t e nd t o e g e s t t h e i r p e l le t s a way f r om t h e i r r o o s t s ( Sou the rn 1 954 ) , t h u s g r ea t ly d im in i sh ing t h e i r p o t en t ia l t oa ccumu la te l a rge n umbe r s o fm ic rof auna l r ema in sb enea th ar oo s t . T awny o w l sd e s t roy p r ey b one t oa much g r ea t e r e x t en t t h an d o b a rn o w l s ( Ra czyn sk i a nd R u p r ech t 1 974 ) . T hey a r e a l so t y p ica l ly a s soc ia t ed w i th a nd h un t w h i l e p e r ched a mong t h e b r an che s i n af o r e s t , e i the r c on if e rou s o rd ec iduou s ( Bu r ton 1 973 ) . A l l e nv i ronmen ta l r e con s t ru c t ion s o f C an t ab r ian S pa in d u r ing t h e l a s t g l a c ia l p e r iod b a s ed u pon n on -m ic rof auna l e v idence ( Chap te r 7 ) i n d ica t e s p a r se f o re s t c ove r . T awny o w l s a l so r oo s t i nt r ee s , w h ich m ak e s t h e c on s i s t en t s e le c t ion o f al ow ly ing c ave s u ch a s E l J uyo a n u n l ike ly c ho ice f o r i n d iv idua l s o f t h i s s p ec ie s . T he c oncen t r a t ion a nd d i s t r ibu t ion o f m ic rof auna w i th in t h e c ave r u le s o u ta c cumu la t ion b y o w l s r oo s t ing i na t r ee j u s to u t s id et h e c ave mou th . O th e ra s pec t s o ft awny o w l b ehav io r d o c o inc ide w i th t h o se o f b a rn o w l s ( non s p ec ia l iz ed d i e t , r a nge i n c lud ing E u rope , s i ze r a nge a mong p r ey s p e c ie s ) , s o o ne m u s t c on s id e r t h e g ene ra l p o s s ib i l i ty t h a tt awny o w l s m ay h ave c on t r ibu t ed t ot h e E l J uyo m ic ro f a una l a s semb lage . H oweve r , t h e o v e ra l l e v idence i n d ica te s t h a t t h ey a r e n o t s i gn if i can t a c cumu la to r s o ft h e E l J uyo r e m a in s .

D iu rna lr a p to r s ( e ag le s , h awk s , o rf a lcon s ) c an b e e l im ina t ed a ss i gn if i can tc on t r ibu to r s b y m any f a c to r s . T he se p r eda to r s c au se much g a s t r ic c o r ro s ion o n p r ey b one s a nd t y p ica l ly b r eak t h em m uch mo r e g r ea t ly t h an i st h e c a se f o r t h e E l J uyo a s semb lage . T h i s b r eakage d i sc repancy i st r ue e ven t h ough p o s tdepo s i t iona l a nd e xcava t ion / sc r een ing f o r ce s h ave c au sed f u r the r d amage t o t h i s s amp le . Wh i le m any s p ec ie s o fr a p to r w i l l r oo s t i nr o ck f i s su re s , ad eepe r c ave s e t t ing l i k eE lJ uyo i sl e s s l i ke ly . A l l a spec t s o ft h e E lJ uyo a s semb lage i n d ica t e i t sa c cumu la t i on b yb a rn o w l s . O the rs p ec ie s o fo w l c ommon t o We s t er n E u rope c an b e e l im ina ted b y g r o s s c ha ra c t e r i s t ic s . L ong e a r ed o w l s a nd t awny o w l s d e s t roy b one s o ft h e i r p r ey t oa much g r ea t e re x ten tt h an d oe ag le o w l s , s ho r t -ea r ed o w l sa nd b a rn o w l s . L ong -ea red o w l s a nd t awny o w l s a r e a l so f o r e s t d we l l ing a nd r o o s t i nt r ee s . T awny o w l s a l so h ave am uch l owe rp o t en t ia lt oc oncen t ra te p r ey b one s a t as i ng le p o in t i n t h e i r r a nge , s i nce p e l le t s a r e e g e s ted mo s t o f t en a way f r om t h e i r r oo s t s . S ho r t -ea r ed o w l s a nd e ag le o w l s c an b e e l im i n a t ed b ya s pec t so ft h e i rb ehav io r : c ho ice o fr o o s t ing s i t ea nd i nt h ec a se o ft h el a t t e rs i ze r a nge o fp r ey .

B a rn o w l s l e ave a h i gh p ropo r t ion o f c omp le te e l emen t s , m uch h i ghe rt h an g r ea th or n ed o w l s d o . D e l ica te l o ngbone s a r e r a r e ly d amaged . T he h i ghe s t d eg r ee o f d amage i st o s c apu lae ,b u te ven t h e se a r en o rma l ly r e cove r ed i na n i d en t i f i ab le s t a t e . O n ly af ew i n s tance s o fs l igh t g a s t r ic c o r ro s ion a r e t y p ica l . G roup s o f a r t icu la t ed e l emen t s ( v e r teb rae o r m e tapod ia l s ) a r e o cca s iona l ly r e cove r ed f r om p e l le t s . T ee th o f t en r ema in i nt h e i r s o cke t s , b u t c ou ld b e e a s i ly r emoved . B ar n o w l d amage t ob one a s semb lag e s i sm in ima l , i nk eep i n gw i th t h e i rs wa l low ing o fw ho le p r ey , r e la t ive ly h i gh g a s t r i c p H , s l igh t s t oma ch mu scu la tu r e , a nd f r equen t p e l le t e ge s t ion .

P OSTCRAN IAL F US ION T he r e la t ive e x ten t o fp o s t c ran ia l e p iphy sea l f u s ion w a s r e c o rd ed a mong t h e t h r ee m a in s i ze c l a s se s o fe l emen t s r e cov e r ed ( sma l l , m ed ium a nd l a rge ) . T he se d a ta a r e i n f o rma t ive d ue t ot h ed i ff i cu l ty o fa ge d e t e rm ina t ion u s ing t e e th o fm o s t m ic ro t ine s , w h ich g row c on t inuou s ly t h roughou t t h e l i f e o f t h e se s p ec ie s . M ic ro t ine s make u p t h e m a jo r i ty o f s ma l l mamma l r ema in s f r om t h e s i t e . T he se r e su l t s a r e l i s t ed i n T ab le 2 . 8 . T he r e co rded a moun t s i n d ica t e t h e t o ta l n umbe r ( r igh t a nd l e f t ) o f u nf u sed p rox ima l o r d i s ta l e l emen t s f o r t h r ee mo s t v i s ib le a nd r obu s t p o s t c ran ia l e l emen t s : p rox ima l h umeru s , p r ox ima l f emu r , d i s t a l f emu r a nd p r ox ima l t i b io f i bu la . Wh i le m any o f t h e u nf u sed e p iphy se s t h em se lve s w e r e r e cove r ed , t h ey w e re n o t i n c luded i nt h e se c a lcu la t ion s s i nce t h ey w ou ld i nf la te t h ep ropo r t ion so fu nf u sed e l emen t s : t h e se w ou ld i n s ome c a se s b e r e p r e sen t ed t w ice . T h e f u sed /unf u sed d i aphy se sw e r et a bu la ted , w h ich i nt h e c a se o f s ome f emo ra i n c luded b o th e nd s . T he se w e r e t a bu la t ed s epa ra t e ly f o r p rox ima l a nd d i s ta l f u s ion . T he se d a ta a r e p r e sen t ed c umu la t ive ly f o r a l l l e ve l s o f E l J uyo , s i nce i n g ene ra l s i ng le l e ve lt o ta l s w e r et o o s ma l l a nd i nn o c a se w a s as i gn if i can t d i f f e rence n o t ed b e tween l e ve l s f o r a ny o ft h e s i ze c l a s se s . T h i s d a ta d oe s n o t i n c lud e t h e r ema in s o f

P REDATOR S UMMARY T ab le 2 . 7 ( a bove ) s umma r ize s t h e c ha ra c te r i s t ic s o f t ho se p r eda to r s t h a t m u s t b e s e r iou s ly c on s ide red a s a mong t h e p o s s ib le a c cumu la to r s o f t h e E l J uyo a s semb lage . Non p r eda to ry a g enc ie sh ave b een r u led o u to nt h eg round s o ft h e s i z e a nd s t a t e o f t h e m ic rof auna l a c cumu la t ion a nd t h e p hy s i ca l l a you t o ft h e c ave v e s t ibu l e . Mov ing w a te r c ou ld n o t h ave p r oduced t h e p a t t er n o fr ema in s f ound a nd wou ld h ave l e f tt r a ce s o fd amage o n t h e b one s . N a tu ra l s i nkho le t r app ing i sn o tp o s s ib le , s i nce t h ev e s t ibu le o ft h e c ave i sn o t as u i t ab l e d e s ign . N a tu ra l d enn ing d ea th s o f m ic rof auna w ou ld n o t h ave o ccu r r ed i n t h i s d epo s i t iona l s e t t ing n o r w ou ld i th ave p roduced t h es p ec t rum o fs p ec ie sa ccumu la t ed . S ome a n ima l a gency t h e r ef o r e m u s t h ave b een t h e p r ima ry a c cumu la to rt h e se r ema in s . A ccumu la t ion i n as ma l l c a rn ivo r e ( mu s te l id , c an id o r f e l id ) l a t r ine a r ea i se l im ina t ed b y t h e v e ry h i gh d eg r ee o f p r ey s k e le t a lb r eakage , l ow d eg r ee o fs k e le ta lc omp le t ene s s( mo r e d u rab l e e l emen t s g r ea t ly o u tnumbe r l e s s d u rab le e l emen t s ) a nd v e ry h i gh d eg ree o fg a s t r ic c o r ro s ion c au sed b y p r eda t o r s i nt h i sc l a s s . S ome s ma l lc a rn ivo r e ss u ch a sw ea se l s a nd

2 9

T ab le 2 . 8 : P o s tc ran ia lE p iphy sea lF u s ion f o rA l lL eve l so fE lJ uyo . Sma l l

Med ium x /y

L a rge

x / y

%

%

x /y

%

P rox H umeru s

9 / 23

3 9 .1 %

2 34 /283

8 2 .7%

7 2 /88

8 1 .8%

P rox F emu r D i s tF emu r P rox T ib ia

2 / 38 1 / 14 9 / 14

5 . 3% 7 . 1% 6 4 .3 %

4 5 /353 6 3 /98 9 2 / 146

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

1 8 / 109 3 3 /49 3 3 /48

1 6 .5% 6 7 .3% 6 8 .8%

x=#o fu nf u sed e l emen t s( doe sn o ti n c lude e p iphy se s ) y=t o ta l #o fe l emen t s %= % u nfu sed T a lpa , w h ich h a s v e ry d i f f e r en t p o s tc ran ia l e l emen t s f r om t h e o t he rs ma l l mamma ls p ec ie st a bu la ted .

D ickman e ta l . 1 99 1 ; S a in t G i ron s 1 973 ) . Mo s t i n d iv idua l s t a ken f a l l i n to t h e s ubadu l t a nd j uven i le c l a s se s . T h i s a ge p ref e rence i sa t t r ibu tab le t o t he r e s t r ic t ion o f s ma l le r i n d i -

Sma l l p o s tc ran ia l e l emen t s g ene ra l ly mu s t c onf orm t o t h e s ma l le r t a xa i d en t if ied f r om t e e th . T he se a r e N eomy s a nd S orex s pp . , t hough t h e p r e sence o fs ome r ema in s o fs pec ie s p re sen t i ns uch l ow a moun t s t h a t t h ey c ou ld n o t b e d e tec ted f r om d en ta lr ema in sc anno tb ed i scoun ted .

v idua ls ( bo th w i th in a nd a mong s pec ie s ) t o f o rag ing a r ea s w i th p oo r e r c ove r ( Ko t le r e t a l . 1 99 1 ; K o t le r e t a l . 1 988 ; L ong land a nd J enk ins 1 987 ) . T h is c ompe t i t ive e xc lu s ion r e su l t s f r om d om inance h i e ra rch ie s t y p ica l t o s ma l l r oden t s ( C rowc rof t 1 966 ; N oye s e ta l . 1 982 ; L loyd 1 975 ) . A du l t ma le s t e nd t o e xc lude n on -ma ted f ema le s a nd a l l s ma l le r ma le s f r om h ome t e r r i to r ie s . Sma l le r i n d iv idua l s o f o t he r s pec ie s a r e s im i la r ly e xc luded . T h is e xc lu s ion f r om t h e b e s t a r ea s o fc ove r o r d en se s t f ood f o rce s t h e se i n d iv idua l s i n to mo re o pen m ic rohab i ta t s a nd /o r s i tua t ion s w he re mo r e t ime i sr equ i red a way f r om b u r rows t o a cqu i r e f ood . Ow l s t a ke t h e se i n d iv idua l s i ng r ea te r n umbe r s d ue t ot h e i r g r ea te rv u l n e rab i l i ty . A ge a nd s ex e f f ec t s m ay b e l e s s p ronounced a mong t e r re s t r ia l c a rn ivo re s b e t te r a b le t o p u r sue s ma l l m amma l s i n to d en se c ove ro rb ur row s .

T he med ium r ema in s c o r re spond t o t h e ma jo r i ty o f t a xa i d en t if ied f r om t e e th . T he se i n c lude A pod emu s ( Mu r idae ) a nd M icro tu s , P l iomy s a nd P i tymy s ( M ic ro t idae ) . Mo s t o f t h e se t a xa c anno t b e a ged b y d en ta l a t t r i t ion , s i nce M icro tu s a nd P i tymy s h ave c on t inuou s ly g row ing mo la r s . Me thod s o f d e te rm in ing t h e se g row th r a te s ( Von K oen ig swa ld a nd G o l e n i shev 1 979 )a r en o ta pp l icab le t oa r chaeo log ica l ma te r ia l s . L eng th o f mand ibu la r d i a s tema ( Sa in t G i rons 1 973 ) c ou ld n o t b e u sed s i nce s o f ew m and ib le s w e re r e cove r ed i n a m ea su rab le s t a te . A s s ignmen t t o as pec ie s w i th in t h e se t wo g ene ra f o r t oo th le s s m and ib le s w a s g ene ra l ly n o t p o s s ib le . S pec ie s d i f f e rence s w ou ld a l te r r e la t ive d i a s tema l e ng th , f u r the r c omp l ica t ing r e su l t s . T he mo la r s o f P l iomy s g row c on t inuou s ly b u t d eve lop r oo t s a nd c an b e a ged u s ing t h i s c r i te r ion . T h i s s pec ie s i sr e p r e sen t ed , h oweve r , b y as i ng le t oo th i nL eve l 4 S . T he mo la r s o fA pod emu s h ave b een a s s i gned r e la t ive a ge s b a sed u pon t o o th w ea r s t age s . T he se r e su l t s a r e i n c luded i nC hap te r 6 . T h i s g enu s , b a sed u pon i d en t if ied t oo th c oun t s , c on s t i tu te s as ma l l p ropo r t ion o fm e d ium r oden t r ema in s . H ence , t h e m a jo r i ty o f i nd iv idua l s t a ken i nt h e m ed ium s i ze c l a s s mu s t b e g i ven r e la t ive a ge

T he r ema in s i nt h e L a rge s i ze c l a s s c o r r e spond a lmo s t e n t i re ly t o A rv i co la . A g ing t h is l a rge m ic ro t ine s pec ie s b y d en ta l a t t r i t ion a ga in i s i mposs ib le d ue t o i t s c on t inua l ly g row ing mo la r s . Mo s t p o s tcran ia l r ema in s o f t h e o t he r l a rge r s ma l l m amma l t a xon d e tec ted f r om t e e th ( Ta lpa ) a r e d i s t ingu i shab le a nd h ave b een t a l l ied s e pa ra t e ly . A ge c l a s se s o ft h e A rv i co la i n d iv idua ls t a ken a r e p a r t icu la r ly i mpo r tan t , s i nce t h e se s p ec ie s ( bo th A . t e rre s t r i s a nd A . s ap idu s ) f a l l a t t h e u ppe r r a nge o f p rey n o rma l ly t a ken b y ab a rn o w l o r o t he r m ed ium o w l s pec ie s . Adu l t m a s s i nt h e se s pec ie s a p p roa che s 3 00 g( Chap te r 3 ) . Wh i le A r v ico la i sr egu la r b a rn o w l p rey w he r e t h e i r r a nge s o ve r lap , i ti su sua l ly p r eyed

b a sed u pon p o s tc ran ia lf u s ion a tl a rge .

u pon i ns ma l le rn umbe r s( Chap te r4 ) . R e la t ive ly l i t t le i sk nown a bou t t h e t ime s o fe p iphy sea l f u s i on f o r s ma l l m amma l s ( e ven i n o t herw ise e x ten s ive ly

B a rn o w l p r eda t ion u pon s ma l l mamma l s e xceed ing 1 00 g

s t ud ied

r e su l t s ma in ly f r om t h er e la t ive a va i lab i l i ty o fs ma l lp r ey i na

m ice

a nd

r a t s ) ,

s o a ge

c l a s se s

b eyond

f u l ly

g rown / subadu l t c anno tb e a s s igned ( Pucek a nd L owe 1 975 ) . P o s tc ran ia l f u s ion i n d ica te s t h a t t h e ma jo r i ty o f i n d iv idua l

g i ven e nv i ronmen t . O ve rt h i s p reda to r ' sc o smopo l i tan r a nge , l a rge r s pec ie s c an m ake u p as i gn if i can t p ropo r t ion o f d i e t ( Ave ry 1 993 ; D av i s 1 958 ; E vans a nd Em len 1 947 ; H e r re ra

p rey i t ems t a ken i n t h e Med ium s i ze c l a s s w e re n o t f u l ly g rown a du l t s . Wh i le t h e f emo ra l h ead w a s u nfu sed o n ly i n

a nd J ak s ic 1 980 ; Ma r t i 1 969 ; P e r r in 1 982 ) . Mo s t s ma l l mamma l s ( p r ima r i ly r oden t s a nd i n sec t ivo re s ) t e nd t oh ave

4 5 o f 3 53 c a se s ( 12 .7% ) , t h e r ema in ing e l emen t s w e re u n f u sed i n6 3 .0% t o8 2 .7% o fa l lc a se s .

a ve rage a du l tm a s sw e l lb e low 1 00 g . T h i sp ropen s i ty b yt h e b a rn o w ls t oc on sume t h e s ma l le r t a xa r e f lec t s t h e i r g r ea t e r g ene ra l a va i lab i l i ty . I ns i tua t ions whe r e l a rge rt a xa a r e mo re a bundan t , t h ey t e nd t ob e t a ken i ng r ea t e r n umbe r s . T he se

T he se d a ta c o r r e spond w i th r e po r t ed a ge p rof i les o f s ma l l mamma l p rey t a ken b y o w l s ( B rown a nd Tw igg 1 971 ; B uck ley 1 977 ; D ean 1 973 ; D e r t ing a nd C ranf o rd 1 989 ;

3 0

l a rge rt a xa , h oweve r , a r e p r ef e r en t ia l ly p r eyed u pon i nt h e i r p r e -adu l ta ge c l a s se s .

w ea r s t age a s s ignmen t f o r A pod emu s i sp r e sen t ed i nT ab le 2 . 9 a nd f o l low s t h e g ene ra l c r i te r ia o f A damcz ew ska A nd rz e jew ska ( 1 967 ) , B i shop a nd D e lany ( 1 963 ) , a nd D e l a ny a nd D av i s ( 196 1) . B o th u ppe r a nd l owe r m o la r s w e r e a ged f o l low ing t h e se g ene ra l s t age sb a sed u pon A .s y l va t i cu s a nd A .f l a v i co l l i s , b o th o f w h ich m ay b e p r e sen t i nt h e E l J uyo s amp le . S i nce t h e se a ge c l a s se s a r e o n ly a pp rox ima te , v a r ia t ion sb ys p ec ie sa r eu n impo r tan t .

T he e p iphy sea l d a ta s uppo r t s t h e i n t e rp r e ta t ion t h a t mo s t ly j uven i le A r v i co la w e r e t a ken . W i th t h e e xcep t ion o f t h e p rox ima l f emu r w h ich f u se s a t ay ounge r a ge , n onf u s ion a mong t h e e l emen t s r a nged f r om 6 7 .3% t o 8 1 .8% . A ge c l a s se sf o rt h em a jo r i ty o fA r v ico la t a k en mu s th ave f a l len i n t h e j uven i le a nd s ubadu l t c l a s se s . T he se i n d iv idua l s a r e much s ma l le r t h an a du l t s a nd c onf o rm mo r e c l o se ly t o a n u ppe r l i m i t o f1 00 gp e r i n d iv idua l p r ey i t em f o r b a rn o w l s . T hey w e r e a l so l i ke ly e xc luded f r om a r ea s o fp r ime f o rage a nd c ov e r b y l a rge r , m o r e d om inan t i n d iv idua l s a nd t h e r e f o r ew e r ea va i lab le i ng r ea te rr e la t ive n umbe r s .

T ab le 2 . 9 : E lJ uyo A pod emu sA ge C la s se sB a sed u pon T oo th Wea r . Wea r S t age

T he r e la t ive s t a te o f e p iphy sea l f u s ion p e r s i z e c l a s s w a s c ompa red . N o s t a t i s t ica l d i f f e r ence w a s f o und c ompa r ing S ma l l , M ed ium a nd L a rge . I n c r ea s ing s amp le s i z e a mong t h e S ma l l s i z e c l a s s m igh t c hang e t h e se r e su l t s , s i nce t h e se r ema in s s h ow ah i ghe r d eg r ee o f f u s ion . R e la t ive ly m o r e a du l t sa mong t h i s s i z ec l a s s ( s h r ew s )m ay f a l lp r ey , s i nce t h e s ubadu l t s a nd j uven i le s o ft h e se s p ec ie s m ay b e s o s ma l l a s t o b e al e s s d e s i rab le f o od c ho ice a mong t h e o w l s . S i nce b a rn o w l s w i l l a l so c on sume a r th ropod s i ns ma l l b u t r e gu la r a moun t s , t h i s i n te rp r e ta t ion i s l e s s t e nab le . T he s t r ik ing a g r eemen t b e tween i n d iv idua l s o f m ed ium a nd l a rge s i z e c l a s s m ay r e f lec t s im i la r g row th p a t t er n s a mong m ic ro t ine s , s i nce e f f ec t ive ly a l o ft h e l a rge r ema in sa nd a lmo s ta l o ft h e m ed ium r ema in s a r e a s s ignab le t ot h i s r oden t f am i ly . A l so i n d ica ted a r e s im i la r p a t t e rn s o fb ar n o w l p r eda t ion , w he r e y ounge ri n d iv idua l sa r ep r eyed u pon p r ef e r en t ia l ly t oa du l t s .

I I I V T o ta l

A ge C la s s

A pp rox ima te A ge

N umbe ro f T ee th

N eona te J uven i le S ubadu l t A du l t

4w eek s 2mon th s 5mon th s 9mon th s

0 2 7 2 7 1 4 6 8

0% 3 9 .7% 3 9 .7% 2 0 .6% 1 00 .0%

( Fo l low ing : A damcz ew ska -And rz e jew ska 1 967 ; B i shop a nd D e lany 1 963 ; D e lany a nd D av i s 1 96 1) T he m a jo r i ty o fi n d iv idua l s t a ken ( 7 9 .4% ) f a l l i n to t h ej uve n i le a nd s ubadu l t c a tego r ie s , p l a c ing t h em w i th in t h e f i r s t h a lf -yea r o fl i f e . A s d i scu s sed a bove , t h e m a jo r i ty o fs ma l l r od en t sa r et a ken b yp r eda to r sb ef o r er e ach ing a du l thood . I n t h e c a se o fA pod emu s a nd b ar n o w l s , t h i s p r ef e r ence r e p r e s e n t s g r ea t e r v u lne rab i l i ty o fy ounge r i n d iv idua l s t op r eda t i on a nd n o t s e lec t ion b y t h e o w l s f o r s ma l le r p r ey : a du l t A pod emu s a r e e a s i ly w i th in t h e p r ey s i z e l i m i t s o f t h i s p r eda to r . N o n eona t e s w e r e t a k en , u n l ike a mong A r v ico la . T h i s a ge c l a s s a mong A pod emu s i sn o rma l ly u nava i lab le t o a v ian p r eda to r s , s i nce t h e y oung h ave n o ty e td i spe r sed f r om t h e i rn a ta l n e s t . I nd iv idua l st h i sy oung m ay f a l lt ot e r r e s t r ia l p r eda to r s , e s pec ia l ly s ma l l mu s te l id s t h a t c an h un t p r ey i n s ma l lb u r row s .

TOOTH E RUPT ION T oo th e r up t ion f o rA rv i co la i n d ica t e s t h a t s ome v e ry y oung i n d iv idua l s w e r e t a ken . Among A r v i co la mo la r s i d en t if i ed b y t h e a u tho r ( n =2 486 ) , 7 3 ( 2 .9% ) w e r e n o t f u l ly e r up t ed ( t he o cc lu sa l s u rf ace h ad n o t c ome i n to c omp le te w ea r ) . I n t h e c a se o f t h i rd mo la r s o n ly ( 3M 3 a nd 3M 3) ,2 7 o f 8 33 ( 3 .2% )h ad n o tf u l ly e r up t ed a tt h et i me o fd ea th . F i na lt o o th e r up t ion o ccu r s d eve lopmen ta l ly v e ry e a r ly i nr oden t s , g en e r a l ly w i th in t h e f i r s tf ew w eek so fl i f e a nd c o r re spond sw i th w ean ing a nd e me rgence o f n ew l i t t e r s f r om m a t er na l c a r e . T he se i n d iv idua l sa r ev e ry v u lne rab le t op r eda t ion a tt h et i me o fi n i t ia ld i sp e r sa l . I nd iv idua l so n ly s l igh t ly o l de rw ou ld n o t b e d e t ec tab le b y t o o th e r up t ion , s i nce t h e i r o cc lu sa l s u rf a ce s w ou ld h ave c ome i n to w ea r a l ong t h e e n t i r e s u rf a ce . C on t i nuou s m o la rg r ow th w ou ld r e p la ce s ub sequen t d en ta l a t t r i t i on .

S ea son o fo ccupa t ion b y t h e o w l s i ss ugge s t ed b y t h e se r e m a in s . L a te s p r ing t oa u t umn m u s th ave b een a tl e a s tp a r to f t h e o ccupa t ion s pan d ue t ot h e p r e sence o fj uven i le s . T he b r eed ing s e a son f o rA pod emu s e nd s i ne a r ly a u tumn , s oj u v en i le s c ou ld n o t b e p r e sen t i n ap opu la t ion p a s t t h e e nd o f t h i s s e a son . T he p r e sence o f s ome a du l t mo la r s ( 2 0 .6% ) i n d ica te s o ccupa t ion m ay h ave c on t inued d u r ing t h e w in te r mon th s ,t h ough t h i sa ge c l a s si sp r e sen tt h roughou tt h ey ea r . TAPHONOMY O F HEDGEHOG R EMA INS T he s k e le ta l e l emen t r e cove ry p a t ter n f o r t h e r ema in s o f h edgehog ( E r inaceu s e u ropaeu s ) d oe s n o t c onf o rm t o t h e l a rge r , f u r -b ea r ing t a xa ( Chap t e r 9 ) , n o r d oe s i tc onf o rm t o t h e p a t t er n f ound f o r s ma l le r i n sec t ivo r e s a nd r od en t s d i s c u s sed a bove . T h i ss p ec ie s( Chap t e r3 )i smuch l a rge r( a bou t 1k g ) t h an t h e t y p ica l m ic rof auna o t he rw i se r e cove r ed ( mo s t

T he p r e s ence o ft h e se v e ry y oung i n d iv idua l s i n d ica te s o c c upa t ion o ft h e c ave b y b a rn o w l s a tl e a s t p a r t ly d u r ing l a t e s p r ing , s umme r , o r e a r ly a u tumn , s i nce t h i s i st h e b r eed ing s e a son f o rA r v i co la ( Chap t e r 3 ) . I n d iv idua l s w i th u ne rup ted mo la r s m u s t h ave e n t e r ed t h e p opu la t ion d u r ing t h i s p e r iod , t h ough o l d e r i n d iv idua l s c ou ld h ave b een t a ken a ta ny t i me o ft h ey ea r .

b e low 4 0 g ,w i th s ome r e a ch ing u p t o3 00 g ) . I ti sr a r e ly p r ey o fs p ec ie s s u ch a sb a rn o w l s . H edgehog s w e r e e n t i r e ly a b sen t f r om at o ta l s amp le o f4 7 ,865 v e r teb ra te p r ey i t em s t a ken f r om 1 88 b a rn o w l p e l le t s amp le s f r om B r i ta in a nd I r e land ( G lue 1 974 ) . Wh i le i ti sp o s s ib le t h a t j uven i le

T he mo la r s o fA pod emu s ( Mu r ida e ) d o n o t g row c on t inu o u s ly a nd a low a s s ignmen tt og ene ra l a ge c l a s s . T h i s g enu s b e long s t ot h e Med ium s i z e c l a s s , w h ich i sd om ina ted b y m ic ro t ine s t h a t c anno t b e a ged b y t h i s c r i t e r ion . G ene r a l

3 1

d e s t royed . N o g a s t r ic c o r ro s ion w a s n o t ed o n a ny o ft h e se e l emen t s . S i nce t h ey a r e g ene ra l ly o fl ow m ea t u t i l i ty , t h ey m ay r e p re sen t t h e r ema in s l e f t b eh ind b y al a rge r p r eda to r i n hab i t ing t h e a r ea o ft h e c ave t h a t h ad c on sumed t h e m i s s i n ge l emen t s .

h edgehog c ou ld b et a ken b y ab a rn o w l( a sa r et h eo cca s iona l j uven i le r a bb i t ) , t h i s t a xa i sg ene ra l ly o u t s ide t h e s i z e o ft h i s p r eda to r . H ypo the t ica l j uven i le p r ey , b e ing t o o l a rge t o s wa l low w ho le , a r e l i ke ly t o h ave t h e i r e d ib le p o r t ion s f l en sed f r om t h e i r b one s a nd s p iny h i d e s . T h i s f e ed ing p a t t e rn w ou ld l e ave mo s t b one s b eh ind a tt h e k i l l s i t e a nd p r e v en t t h e i r r e tu rn t o ar oo s t ing s i t e . E ag le o w l s w i l l t a ke h edgehog s ,b u tt h i sp r ey w a sr e cove red i nv e ry l ow n umbe r s . P r ey i nt h i s s i ze r a nge i st y p ica l ly am uch l a rge r p ropo r t ion o f e ag le o w l d i e t . T he p r eponde rance o f c r an ia l e l emen t s a l so a r gue s a ga in s t o w l p r eda t ion u pon t h i s s p ec ie s , s i nce t h e se b one sa r ed i ff i cu l tt os wa l low a nd a r et h em o s tl i ke ly t o b e l e f tb eh ind a t ak i l l s i t e . L ow m ea t u t i l i ty e l emen t s s u ch a sf o o t b one s a r e a l so l e s s l i ke ly t ob e s wa l lowed , y e t t h e se a ppea ri nh i gh r e la t ive a bundance a tE lJ uyo .

S UMMARY T aphonom ic a na ly s i s o ft h e m ic rof auna f r om E l J uyo i sa n ece s sa ry p r ecu r so rt op a leoenv i ronmen ta l a na ly s i s , s i nce i n t h i s m anne r t h e p r ima ry a c cumu la to r o ft h e a s semb lage c an b ed e t e rm ined . A na ly s i s i n d ica te st h a tb a rn o w l s( T y to a l ba ) a r e r e spon s ib le f o rn ea r ly a l t h e m ic rof auna l r ema in s i nt h e s i t e . T h i s c onc lu s ion t a ke s i n to c on s id e ra t ion t h e s pa t ia l l a y o u t o ft h e c ave : i t sa va i lab le d enn ing i roo s t ing p l a ce s , e x te r n a l a c ce s s , r e la t ive l o ca t ion a nd u t i l iza t ion b y h uman s . T he s i z er a nge , a c t iv i ty p a t t e rn sa nd a ge p r of i le s o ft h ep r ey w e r e a l so c on s ide r ed , a sw e r e t h e p e r e l emen t r e cove ry p a t ter n , d i ge s t ive c o r ro s ion , a nd b r eakage o f t h e s k e le ta l r ema in s . T h i s d a ta w a s c ompa red w i th t h e k nown e co log ica l a c t iv i ty a nd t a phonom i c s i gna tu r e s o f t h e p o s s ib l e a c cumu la t ing a genc ie s . A l l o ft h e se e xcep t b a rn o w l s w e r e e l im ina t ed o n m u l t ip le g round s . T he r e la t ion sh ip b e tween b a rn o w l s a nd t h e e co log ica l s i gna l t h a t i sr e co rded b yt h e i r s ma l l m amma l p r ey i sd i scu s sed i nC hap t e r s 4a nd 5 ,w he re i nt h e l a t te r c hap t e r p a leoenv i ronmen ta l r e con s t ru c t ion f o r E l J uyo i s p r e sen ted .

T he r e cove ry p a t t e rn o ft h e se r ema in s ( Tab le 2 . 10 ) i sc on s i s ten t w i th n a tu ra l i n s i te mo r ta l i ty o ft h i s s p ec ie s . H edge h og s d o n o t s hun h uman h ab i ta t ion a r ea s ( Chap te r 3 ) a nd m ay h ave e n te red t h e c ave e ven w h i le o ccup ied . W i th t h e e xcep t ion o f as i ng le f r agmen ta ry t i b io -f ibu la d i aphy s i s , a l e l emen t s r e cove r ed f r om t h i s s p ec ie s w e r e o n ly t h e mo s t d u rab le : t e e th , m and ib le /max i l la f r agmen t s o r w ho le e l e m en t s , a nd f o o t e l emen t s . T h i s p a t ter n i n d ica t e s ah i gh d e g r ee o fp o s tmor t em d amage . T h i s c ou ld h ave o ccu r r ed a sa r e su l t o ft r amp l ing u pon p r ev iou s ly i n ta c t s k e le ton s l e f t o n t h es i te s u rf a ce , w i th t h e mo r e d e l ica te b one sg ene ra l ly b e ing

T ab le 2 . 10 : E r ina ceu se u ropaeu sR ema in sf r om E lJ uyo .

E lemen t t e e th 1 "p ha l 2 " 'p ha l p ha l m e tapod ia l c a rpa l a s t raga lu s t i b ia f r ag . m and ib le /f rag . m ax i l la /f r ag . T o ta l

9 1 0

8 3 1

7 2 0

2 2

L eve l 6 4 R 1 8 3 5 1 1

4 S 2 3

2

1 1 1 1 1 2

3 8

3 2 0

2 5

1 9

3 2

2 6

4 1 7

T o ta l 1 22 4 5 3 3 2 1 1 1 6 1 1 8 1 48

CHAPTER 3 S PEC IES CHARACTER IST ICS AND ECOLOG IES

T h i s c hap t e r d e sc r ibe s t h e b ehav io ra l , mo rpho log ica l , a nd e co log ica l c ha ra c te r i s t ic s o ft h e s ma l l m amma l s p ec ie s p r e s e n t i n C an t ab r ian S pa in d u r ing t h e L a te P l e i s tocene a nd H o locene . T he a u tho r a s sume s t h a t t h e b ehav io r o f t h e se s p ec ie s h a s n o t s i gn if i can t ly c hanged i nt h e f ew t h ou sand i n te rven ing y ea r s . I na lmo s ta l c a s e s ,t h i s a s sump t ion i sn o t r a d ica l s i nce t h e se s pec ie s h ave u nd e rgone n o m a jo r e vo lu t i ona ry c hange s . O ne p o s s ib le e xcep t ion i st h e e vo lu t ion o f c ommen sa lb ehav io ro nt h ep a r to fs ome o ft h e se t a xa ,w h ich n ece s sa r i ly c anno t p r eda te t h e r i se o fm a jo r h uman induced i mpa c t s u pon t h e l a nd scape ( f o r e s t ry , a g r icu l tu r e , p u rpo se f u l ly s e t f i r e s ) a nd t h e e x t en s ive c on s t ruc t ion o fh ab i ta t ion s . Mo s t o ft h i s c oevo lu t iona r y p roce s s t h e r ef o r e m u s t h ave o c c u r r ed b eg inn ing w i th t h e N eo l i th i c P e r iod a nd i t s r e su l tan t c hange s i nl a nd scape u sage . O ne m ay i n f e rt h a tt h i s p roce s s w a s i na n i n c ip ien t s t age d u r ing t h e U ppe r P a leo l i th ic , p a r t i cu la r ly w i th t h e b a rn o w l s r e spon s ib le f o r t h e i n t roduc t ion o fs ma l lm amma lr ema in st ot h es i t e .

w a l laby (M acropu s r uf og r i seu s ) , Ame r ican m ink (M u s t e la v i son ) , C anad ian b eave r ( Ca s to r c anad en s i s ) , m u sk ra t ( Onda t ra z i be th i cu s ) , r a coon ( P rocyon l o tor ) , r a coon d og s ( N yc t e reu te s p rocyono id e s ) , a nd f e ra l a nd d ome s t ica t ed f o rm s o f mu l t ip le s p ec ie s . O the r a l te ra t ion s i n c lude t h e r a nge e x ten s ion o fs ome s p ec i e s a l ready ap a r t o ft h e E u ro p ean f a una s u ch a sh a rve s t m ice (M ic romy s m in i tu s ) , r a bb i t ( O ly c to lagu sc un icu lu s )a nd e d ib le d o rmou se ( G l i sg l i s ) , a nd t h e r a nge c on t ra c t ion / loca l e x t i rpa t ion o f mo s t l a rge p r eda t o r s . T h i s r a nge o fp r ob lem s m u s t b e t a ken i n to c on s ide ra t i on w hen u s ing t h e c on t empo ra ry f a una o f E u rope t o d e s c r ibe e x t inc te co log ica lc ond i t ion s .

T he m e t r ica l c ha rac t e r i s t ic s o f t h e se s p ec i e s h ave v a r ied s omewha t . T h i s v a r ia t ion i sd e sc r ibed i nC hap te r 6 . F o l l ow ing B e rgmann ' s R u le , s ome s p ec ie s m ay h ave h ad a l a rg e r m ean s i ze d u r ing t h e g ene ra l ly c o ld e r P l e i s tocene . T he p o s s ib i l i ty o fs u ch m ic roevo lu t iona ry c hange w a s t a ken i n to c on s ide ra t ion i nt h ei d en t if ica t ion o ft h e se t a xa .

G l i sg l i s( e d ib le d o rmou se ,f a td o rmou se ) ( F igu r e3 . 1 ,e nd o fc hap t e r ) G .g l i si sf o und t oday i nE u rope , r a ng ing f r om n o r ther n S pa in t o U k ra ine . S ou the rn r a nge i n c lude s t h e Med i t e r ranean r e g ion a nd T u rkey , a nd i s o la t ed p opu la t ion s a r e f o und i n t h e C aucu su s r eg ion , s ou ther n E ng land , a nd s ome Med i te r ranean i s land s . T h i s s p ec ie s ' E ng l i sh n ame d e r ive s f r om t h e f a c t t h a t i tw a s r a i sed a nd e a t en b y R oman s . I t s r a nge m ay t h e r ef o re h ave b een a r t if i c i a l ly e x tended t o p r ev iou s ly u n inhab i ted r e g ion s ( a s i n E ng land : C o rbe t a nd O venden 1 982 :153 ) . Mean a du l t m a s s i s1 40 g ,t hough p r io rt oh i ber n a t ion t h i s may e x tend u pwa rd t o2 50 g( Co rbe ta nd H a r r i s 1 99 1 :264 ) . G . g l i s i st h e l a rge s t m embe r o f t h e f am i ly G l i r ida e . I t s d en ta l p a t te rn i s 1 . 0 .1 .3 / 1 .0 .1 .3 , w i th r o o t ed m o la r s e xh ib i t ing d i s t inc t ive t r an sve r se r i dge s . T h i s s p ec ie s i st h o rough ly a r bo r ea l a nd p r ef e r s d ec iduou s w ood land s o fv a ry ing a ge s , a nd w i l l a l so o ccupy o r cha rd s a nd g a rden s . S ome c ommen sa l i sm h a s b een n o ted ( Co rbe ta nd H a r r i s 1 99 1 :265 ) . I t s a l t i tude r a nge s u p t ot h e l i m i t s o ft h e t r ee l i ne , i nt h e P y renee s a bou t 2 , 000 m( Co rbe t a nd O v enden 1 982 :153 ) . T he d i e t o fG . g l i s . i s v a r ied a nd i n c lude s f r u i t s , n u t s , b ud s , b a rk , i n ve r teb ra t e s , f u ng i ,e gg s , a nd n e s t l ing s .

RODENT IA , F AM ILY GL IR IDAE T h i sr oden tf am i ly i n c lude s 6g ene ra i nt h e P a lea r c t ic r e g ion , 5o fw h ich a r e mono spec i f ic , a nd h a s r ema ined t a xonom i c a l ly s t ab le f o r s ome t i me ( Co rbe t 1 978 , 1 984 ) . T he se s p e c i e sa r ec ha ra c t e r i s t ica l ly a r bo r ea la nd h i be rna t ing .

E co log ica l b ehav io rp r e sen t s amo r e d i ff i cu l tp r ob lem , s i nce t h e P l e i s tocene e co logy o fC an tab r ian S pa in d oe s n o t n ece s s a r i ly c onf o rm t oa ny e x tan t l a nd scape . T he m ode rn c l ima te o fE u rop e o we s i t s t empe ra t e n a t u r e t ot h e w a rm ing i nf lu e nce o ft h e G u lf S t r eam a nd g ene ra l o cean p r ox im i ty . T he d ef l ec t ion o ft h e G u lf S t r eam a way f r om E u rope b y t h e i n t r u s ion o fp o la r i c e s t r e t ch ing s ou thwa rd s m ay h ave r e su l ted i nc l ima t ic c hange u np r ed ic tab le b yt h es imp le g l a c ia lr e g ime o fr i s ing / f a l l ing o cean l e ve l . I na dd i t ion , s imp le a na logy t o m o r e n o r the r ly r eg ion s o f E u rop e f a i l s t o c on s ide r d i f f e r e nce s i ns o la ri n f lux , w h ich w a s m a in ta ined e ven d u r ing g l a c i a l c ond i t ion s . T opog raphy f o rt h e n o r the rn c oa s t o fS pa in i sa l so au n ique c on s ide ra t ion , d ue t ot h e p r ox im i ty o ft h e h i gh land z one s o c l o se t ot h e A t lan t ic c oa s t . S pec ie s i n te r a c t ion s t h a t n o l o nge r o ccu r , s u ch a s M i c ro tu s o economu s ( a r c t ic v o le ) w i th e ndem ic I b e r ian o rP y r enean v o le s p ec ie s s u ch a sP i tymy s l u s i tan icu s a nd P .p y rena i cu s m ay h ave a f f e c ted t h e i r e co log ica l b ehav io r a nd h ab i ta t e xp lo i ta t ion . S ome e co log ica l c ond i t ion s o f t h e L a t e P l e i s tocene a r e n o l o ng e rd i r ec t ly o b se rvab le e ven i nm oder nn o r ther nc l ima t e s , w h ich r e duce s t h e u sef u lne s s o f d i r ec t a na logy t o mod er n c ond i t ion s( And r ew se ta l . 1 979 ) .

G . g l i s l i ve s i ng roup s w i th s t rong h i e ra r ch ie s . B o th y oung a nd t h e m a le m ay r ema in w i th t h e a du l t f ema le s d u r ing w in t e r h i ber na t ion . T h i s o ccu r s i nn e s t s b o th a bove a nd b e low g r ound , a s t h i s s p ec ie s c an a l so b u r row . H ome r a ng e s a r e t i ny ( c a . 1 00 m d i ame te r ) a nd d i spe r sa l i so n ly 1 2 k m . A c t i v i ty i se n t i re ly n oc t u rna l , w i th t h e d ay l igh t h ou r s s p en t h o led u p i n at r ee c r ev ice o r an e s tc l o se t ot h et r unk ( Co rbe t a nd H a r r i s 1 99 1 :265 -266 ) .

T he l a rg e sca le h uman i n duced c hange s t o t h e E u ropean l a nd scap e a luded t oa bove h ad n o t y e t o ccu r r ed , s o o u r u n d e r s tand ing o ft h e P l e i s tocene f a una i sa l so p a r t ia l ly l i m i ted b y h av ing a n a l te red m oder n f a una f o r s t udy a nd c ompa r i -

S exua l ma tu r i ty t y p ica l ly i sn o t r e ached u n t i l a f t e r t h e f i r s t w in t e r . L i t t e r s i z e s o f4 6 a r e c ommon , t h rough t h e k nown r a nge i s1 11 ( Co rbe t a nd H a r r i s 1 99 1 :263 -266 ) . G e s ta t ion l a s t s2 2 -24 d ay s ,w i th w ean ing o ccu r r ing a ta bou t3 0 d ay s , a t w h ich t ime t h e y oung l e ave t h e n e s t . T he b r eed ing s e a son

s on . T h i sa l te ra t ion i n c lude st h e i n t roduc t ion o fn ew s p ec ie s s u ch a s g r ay s qu i r r e l ( S c iu ru s c a ro l inen s i s ) , h ou se mou se ( M u s m u scu lu s ) , r a t ( Ra t tu s r a t tu s ) a nd ( R . n or veg i cu s ) ,

3

P i tymy s i n to s ubgene r a o fM i c ro tu s . T h e a u tho r w i l l i nt h i s c a se r e ta in t h e u s age o f s p l i t t ing i n to d i s t inc t , b u t c l o se ly r e la t ed g ene ra . O the r m embe r s o ft h i s f am i ly i n c lude l em m ing s ( L emmu s l emmu s a nd D i cro s tony xt o rqua tu s i nn o r th er n E u rope ) , t r ue t u nd r a -dwe l l ing s p ec ie s n eve r y e t r e cov e r ed f r om a ny S pan i sh s i t e . T he s t epp e l emm ing ( Lagu ru s l agu ru s ) , t o day e ndem ic t ot h e s t eppe s o fE u ropean R u s s ia , h a s b een i d en t if i ed v e ry r a r e ly f r om C an tab r ian s i t e s . T he t u nd ra v o le ( M i c ro tu s g reg ra l i s ) h a s b een f o und i no ne s i t e o n ly a nd i nv e ry s ma l l n umbe r s . Memb e r s o ft h e m ic ro t id f am i ly a r e t y p ica l ly o pen g r a s s land o rt u nd ra d we l le r s , a nd b u r row e x ten s iv e ly b u ts h a l low ly i n to s u i t ab le t o p so i l .

e x tend s o n ly f r om m id June t h rough A ugu s t . L if e e xpec t a ncy i nt h ew i ld i s3 -4 y ea r sa nd c an b eh i ghe ri nc ap t iv i ty . E l iomy sq ue rc inu s( g a rden d o rmou se ) ( F igu r e 3 . 2 ) E . q ue r c inu s o ccup ie s t empe ra te E u rope f r om t h e Med i t e r ranean t oF i n land , i n c lud ing S pa in a nd E u ropean R u s s ia . T h i ss pec ie so fd o rmou se i ss ma l le rt h an G .g l i s , b u t h a s t h e s ame d en ta l f o rmu la ( 1 .0 . 1 .3 / 1 .0 .1 .3 ) a nd l ame l la r p a t te rn t o m o la r r i dge s . A du l t m a s s i se x t reme ly v a r iab le a c ro s s i t sr a nge . Mean sr a ng ef r om 6 0 gi nc en t ra lE u rope t o 1 70 go nF o rmen t e ra I s land , S pa in . 1 00 gi smo re t y p ica lf o r t h i s s p ec ie s ( s umma r iz ed b y Mo r eno a nd C o l lado 1 989 ) . A l though i t s p r ima ry h ab i ta t i sc on if e rou s a nd d ec iduou s f o r e s t , E . q ue rc inu s i sl e s s a r bo r ea l t h an o t he r s p ec ie s o f d o rmou se a nd c an b e e ncoun t e r ed i nl owe rf o l iage z one s , o n t h e g r ound , a nd i so cca s iona l ly c ommen sa l . T h i s p r ef e r ence f o r t r ee s t e nd s t o m ake t h i s s p ec ie s l e s s v u lne rab le t o o w l p r eda t ion , t h ough t h i s m ay i n c r ea se d u r ing t i me s o fj uven i le d i spe r sa l . I n hab i ted a l t i t ud e s r a nge u p t o 2 , 000 m i nt h e A lp sa nd P y r enee s( Co rbe ta nd O venden 1 982 :152 ) .

M ic ro tu sa g r e s t i s( f i e ld v o le ; s ho r t ta i led v o le ) ( F igu r e 3 . 3 ) M . a g r e s t i s r a nge s a c ro s s t empe ra t e a nd b o r ea l E u rope t ot h e U r a l s , w i th d i scon t inuou s d i s t r ibu t ion e x tend i n g e a s twa rd t oL ak e B a ika l i nS i be r ia . Moder nd i s t r ibu t ion d oe s i n c lude n o r the rn S pa in , b u t d oe s n o t i n c lude t h e Med i t e r ranean r e g ion . C o rbe t a nd H a r r i s ( 1 99 1 :204 ) f o und a n a du l t a ve rage o f3 9 .7 g f o r m a le s ( n =2 0 ) a nd 3 0 .9 g f o r f ema le s( n =2 5 )f o rs e cond y ea r( f u l ly g r own )a du l t sa tL och T ay , S co t land . M ean m a s s f o r ap opu la t ion i n c lud ing j uve n i le s a nd s ubadu l t s w ou ld t h e ref o r e b e s omewha t l owe r . P r ef e r r ed h ab i ta t s a r e r o ugh , u ng raz ed g r a s s land s a nd m a r g i na l ly i n to w ood land , h edge row s , b og s , d une s , s c ree , a nd moo r land s ( Co rbe t a nd H a r r i s 1 99 1 :205 ) . F eed ing i sh e r b i vo rou s ,p r ima r i ly o ng r een l e ave sa nd g r a s ss t ems .

E . q ue rc inu s b r eed ing s e a son i nS pa in r u n s f r om F eb rua ry t h rough O c tobe r . S exua l m a tu r i ty i sr e ached a ta bou t 6 0 g a du l t b ody m a s s . Two l i t t e r s p e r a du l t f ema le e a ch y ea r a r e t y p ica l , w i th r e cen t ly w eaned y oung a ppea r ing i n Ma rch J une a nd S ep tembe r -Oc tobe r . G e s ta t ion l a s t s f o r a bou t 2 5 d ay s . Wean ing o ccu r s a t4 0 -45 d ay s , a tw h ich p o in t t h ej u v en i le s h ave r e ached ab ody m a s s o fa bou t 3 2 g . A ve rage l i t t e rs i z ei s5 . 5 , w i th ar a nge o f4 8( Mo r eno 1 988 ) .

T h i s s p ec ie s i sp r ima r i ly n oc tu rna l i n s umme r , b ecom ing mo r e d i u rna l i nw in te r . J uven i le s t e nd t ob e mo re n oc tu rna l t h an a du l t s ( Co rbe t a nd H a r r i s 1 991) , p e rhap s d ue t ot h e i r c ompe t i t ive e xc lu s ion f r om f a vo red m ic rohab i ta t s w i th d en se r c ove r b y l a rge r a du l t s . B r eed ing o c cu r s f r om Ma r ch t h rough O c tobe r , w i th m u l t ip le l i t te r s o fg ene ra l ly 2 7 e a ch . Ag e s ta t ion p e r iod o f 1 8 -20 d ay s i st h e n o rm . We igh t a t b i r th i s2g ,a nd w ean ing o ccu r sb y2 8 d ay s( Co rbe ta nd H a r r i s 1 99 1 :206 ) b y w h ich p o in t a l t e e th a r e f u l ly e r up ted a nd h ave c ome i n to w ea r . V e ry f ew i n d iv idua l s s u rv ive t h rough t wo w in t e r s i nt h e w i ld . T h i s s pec ie s i sc ommon p r ey o f d i ur na lr a p to r s , n oc tur n a lr a p to r s , a nd m u s t e l id s a nd i s ap r e f e r red p r ey i t em o ff o xe s .

RODENT IA ,F AM ILY M ICROT IDAE Membe r s o ft h i s r od en t f am i ly d om ina t e t h e m ic rof auna o f E l J uyo a nd o t he r C an tab r ian s i t e s . T h i s t a xon i sv a r iou s ly l i s t ed a s t h e f am i ly M ic ro t ida e d ue t o p opu la r u sage , b u t t a xonom ic r u le s g i ve p r eced ence t oA rv ico l ida e . T o a vo id c onf u s ion , t h e a u tho r w i l l r e ta in t h e p opu la r u sage o f t h e t e rm m ic ro t ine s t od e sc r ibe t h i s g r oup . T h i s f am i ly i sa l so v a r iab ly d e sc r ibed a s as ubf am i ly o fC r ice t idae ( w i th h am s t e r s c omp r i s ing C r ice t inae see f o r e xamp le C o rbe t 1 978 ) . T he m embe r s o ft h i sr od en t f am i ly a r e c ha ra c te r ized b yt h e i r p r i sma t ic , c on t inuou s ly g r ow ing m o la r s w h ich f o rm r oo t s o n ly i no ne e x tan t g enu s , C l e th r ionomy s . Mo la r r oo t s a l so f o rmed i no n e s p ec ie s ( P l iomy s l e nk i ) , w h ich w en t e x t inc t a t t h e e nd o ft h e P l e i s tocene a nd i sf o und i ns ma l l n umbe r s i n C an tab r ian s i t e s . As ho r t , a b r up t f a ce , s ma l l e x t er na lp i nna e , a nd r e la t ive ly s ho r t t a i l s a l so c ha r a c t e r iz e t h i s t a xon . A s i s t h e c a se f o r a l r oden t s , t h e u ppe r a nd l owe r i n c i so r s g row c on t inuou s ly . T he d en ta lf o rmu la f o ra l s p ec ie so fm ic ro t ine i s1 . 0 .0 .3 / 1 .0 .0 .3 .

M ic ro tu sa r va l i s ( c ommon v o le ) ( F igu r e3 . 4 ) Mod e rn r a nge e x tend sa c ro s st empe ra te E u rope , f r om n o r ther n S pa in a nd i n to S i be r ia n ea r ly a s f a r a s L ak e B a ika l . Med i t e r ranean a nd f a r n o r ther n r e g ion s a r e n o t i n c l uded i st h i s d i s t r ibu t ion . C o rbe t a nd H a r r i s ( 199 1 :209 ) i n d ica t e am ean o f4 2 gf o r m a le s ( n =3 9 , r a nge 2 9 -67 g ) a nd 3 6 gf o rf ema le s ( n =3 8 , r a nge 2 2 -55 g )o n O rkney ( t he C hanne l I s land s ) . M . a r va l i s i s ah ab i t a t g ene ra l i s t a nd c an b e f o und i nd ec iduou s a nd c on if e rou s f o r e s t , m a r sh , h ea the r moo r land , h ay m eadow s , a nd g a rden s b u t i s mo s t ly a b sen t f r om a r ab le l a nd a nd s ho r t p a s tu r e s ( Co rbe t a nd H a r r i s 1 99 1 :210 ) . I td oe s p r ef e r m eadow o ve rt h e se o t he r h ab i ta t s ( Co rbe t a nd O venden 1 982 :160 ) , a nd c a n b e f ound i n i t s h i ghe s t d en s i t ie s t h e r e . Mo s t f e ed ing i sh e rb ivo rou s : l e ave s

M i c ro t in e e vo lu t iona ry h i s to ry b eg in s b y t h e P l iocene , a t w h ich t i me a l g ene ra h ad c l o sed r o o t s . T h i s s h if t t owa rd s h yp sodon ty o ccu r r ed i nm u l t ip le p a ra l le l l i neage st h roughou t t h e i r s ub sequen t e vo lu t ion ( Brune t -Lecom t e 1 990 ; K owa l sk i 1 966 ) . T he g enu s P i tymy s e vo lved d u r ing t h e M idd le P l e i s t o cene , b u t w a s l a te r s upp lan ted b y M i c ro tu s , w h i ch i ss t i l l d om inan t t o day a nd h a s u nde rgon e r a d ia t ion i n to d oz en s o f H o la r c t i c t a xa ( s ee G romov a nd P o lyakov 1 992 f o r ar e cen t s umma ry ) . A r v ico la e vo lved i nt h e M idd le P l e i s tocene f r om M imomy s , a nd h a s t wo e x tan t f o rm s t h a t d i f f e rc y to log ica l ly ( A . s ap id u s a nd A . t e r re s t r i s ) . N omenc la tu r e i sf u r the r c on f u sed i nt h i s c l ade , s i nce s ome a u tho r s l ump C h ionomy s a nd

a nd s t em o f g r a s se s a nd a v a r ie ty o f d i co ty ledon s ( Co rb e t a nd H a r r i s 1 99 1 :2 10 ) . T h i s s p ec ie s i sa c t ive b o th n i gh t a nd d ay . B reed ing o ccu r s s omewha t e a r l ie r t h an o t he r s p ec ie s o fm i c r o t id , s t a r t ing i nF eb r ua ry a nd r unn ing t h rough S ep t embe r , 3 4

( i nc lud ing p a s tu r e a nd c r op land ) f a vo r ed b y o t he r s pec ie s o f M i c ro tu s .

w i th mu l t ip le l i t t e r s i n as e a son b e ing t h e n o rm . F ema le s b o rn e a r ly e nough i n as e a son m ay e v en r e p roduce b ef o r e i t s e nd . Two t of i ve a r e t y p ica l f o r o ne l i t t e r , w i th 3 . 3 b e ing a ve rage . G e s ta t ion a nd l a c ta t ion b o th l a s t a bou t 2 0 d ay s ( Co rbe ta nd H a r r i s 1 99 1 :2 11) .

B r eed ing f o r M o economu s i nE u rope e x t end s f r om e a r ly May t h rough l a te S ep tembe r . G e s ta t ion l a s t s f o r a bou t 2 0 d ay s ( Nadeau 1 985 ) . L i t te r s i z e r a nge s f r om 2 8 , w i th a n a ve rage o f 5i nE u rope , w i th h i ghe r v a lu e s i nt h e N ea r c t ic ( K e l le r 1 985 ;L ig tvoe ta nd V an W i jngaa rd en 1 994 ) .

M i cro tu sn i va l i s( a lp ine v o le , s now v o l e ) ( F igu r e 3 . 5 ) M . n i va l i s h a s ar e l i c t d i s t r ibu t ion a c ro s s t h e a l p ine r e g ion s o f E u rope , i n c lud ing S pa in , F rance , I t a ly , G reece , S w i tz e r land , a nd t h e f o rme r Y ugo s lav ia . N ie tham m e ra nd K r app ( 1 982 :264 ) f o und am ean s i z e o f4 1 .3 g( n2 0 , r a ng e 1 8 -59 ) f o r E u ropean m u seum s p ec imen s o f t h i s s p ec ie s . I t s p r ef e r red h ab i ta t i sl o o s e r o ck a nd g r a s s land a bove t h e t r e e l i ne a nd m o r e o pen m on tane w ood land . I ti s f o und u pt os now l ine , a nd i nS w i tz e r land i smo s tp opu lou sa t 1 600 -2600 m ( Co rbe t 1 966 :247 ) . M . n i va l i s l i ve s u p t o 4 , 000 m o n Moun t B lanc ( Co rbe t a nd O venden 1 982 :16 1) , b u tc an b ef o und a sl ow a s3 0ma . s . l . i nt h e f o rme rY ugo s la v i a ( K ry s t uf ek 1 990 ; K ry s tuf ek a nd K ova c ic 1 989 ) . I t s d i s t r ibu t ion t h e r ef o r e s e em s mo re d ependen t u pon t h e a va i l a b i l i ty o fr o cky / f i s su re h ab i ta t st h an a l t i tude i t s e lf . T h i s s p e c i e s a l so w i l l e n te r u nde rg round c av e rn s , ap o ten t ia l ly i m p o r tan t c on s ide r a t ion w hen d ea l ing w i th a r cha eo log ica l c ave d epo s i t s .

M ic ro tu sg rega l i s( t und ra v o le ,n a r row sku l led v o le ) ( F igu r e 3 . 7 ) M . g rega l i s h a s ad i scon t inuou s r a nge a c ro s s t h et u nd ra a nd f o r e s t ed s t eppe z one s o fA s ia . I t s p ene t ra t ion i n to n o r the rn S pa in d u r ing ac o ld e p i sode t h e r ef o r e i n d ica te s s i gn if i can t a l p ine t u nd r a / s teppe i nt h e C an tab r ian C o rd i l le ra . T yp ica l a du l t m a s s i sa bou t 4 0 g( Ognev 1 964 :404 ) . T h i s s p ec ie sh a sb een i d en t if i ed f r om o n ly o ne S pan i sh P a leo l i th ic s i te : E r ra l la L owe r Magda len ian L eve l V ( P emän 1 985 ) . S i nce t h i s s i t e i sl o ca t ed w e l l a way f r om t h e moder n c oa s t l i ne a nd i sw i th in t h eh un t ing r a d iu s o fu p land h ab i ta t s , t h e r e i sn on eed t op o s tu la t et h a tt h i sh ab i ta tp ene t ra ted t ol ow land c oa s ta lC an tab r ian S pa in e ven d u r ing g l a c ia lm ax ima . C omp lex b u r row c o lon ie s w i th n ume rou s e x i t s ( 30 -40 ) a r e e xcava t ed a nd i n hab i t ed p e r en ia l ly . A c t iv i ty i sn oc tu rna l a nd p eak s a f t e r s un se t . T he b r eed ing s e a son i sr e la t ive ly s ho r t f o rM g rega l i si nt h en o r the rn p a r to fi t sr a nge , c ommenc ing i nl a t eJ une / ea r ly J u ly a nd l a s t ing a tl e a s tt h rough S ep tembe r . Two l i t te r s a r e n o rma l p e r a du l t f ema le e a ch s e a son . L i t t e r s i z e i sv e ry h i gh , a ve rag ing 8 9 . F a r the r s ou th , l i t te r s a r e p roduced a s e a r ly a s May , w i th 5 12 n eona te s t y p ica l . Membe r s o f t h e f i r s t l i t t e r e a ch b r eed ing s e a son t y p ica l ly r e a ch s e xua l m a tu r i ty a nd r e p roduce t h a t s ame y ea r ( Ognev 1 964 ) .

B r eed ing b eg in s a tt h e e nd o f A p r i l , w i th l i t t e r s p roduced f r om May t h rough A ugu s t . T h i s s e a son c an e x tend s l igh t ly a tt h e l owe r a l t i tud ina l e x t r eme s o fM i c ro tu s n i va l i s r a nge , b u t i ti sr e la t ive ly s hor t w hen c ompa r ed w i th t h o se o fo t he r s p ec ie s o fE u ropean m ic ro t ine s . F ema le s b o rn e a r ly e nough i nt h e y ea r a r e a b le t o r e p roduce b ef o r e t h e s e a son ' s e nd . A du l tm a s s i so b ta ined a ta bou t 3m on th s o fa ge . L i t t e r s i z e r a nge s f r om 2 4 , w i th am ean o f3 ,a nd a n a du l t f ema le n o r m a l ly p r oduce s t wo l i t t e r s p e r y ea r . G e s t a t ion l a s t s f o r l e s s t h an o ne m on th ( Le L oua rn a nd J a neau 1 975 ) .

C l e thr ionomy sg lareo lu s( r ed -ba cked v o le ;b ank v o le ) ( F igu re 3 . 8 ) C . g la reo lu s r a nge s a c ro s s t empe ra te E u rope i n to c en t ra lA s ia a sf a ra sL ake B a ika l . N o r the rn S pa in , B r i t a i n a nd S weden a r e i n c luded i nt h i s r a nge , b u t t h e Med i t e r r a nean z one g ene ra l ly i sn o t . T h i s v o le g enu s d i f f e r s f r om t h e o t he r s d e sc r ibed h e r e i n t h e p r e sence o f mo la r r oo t s , w h ich d eve lop a f t e r b i r th a nd g r ow t h roughou t t h e i n d iv id u a l ' s l i f e t ime . T he mo la r s h ave t h e t y p ica l m ic ro t ine mo r p ho logy o f d i v i s ion i n to p r i sm s w h ich a r e w or n d own i n c r o s s sec t ion . T he i r r oo t s b ecome v i s ib le b y a ge 3 m on th s ( A l iba i 1 980 ; B i shop a nd D e lany 1 963 ) . D if f e rence s i n p r i sma t ic mo rpho logy a l so m ake t h e mo la r s o ft h i s s p ec ie s d i s t ingu i shab le f r om t h o se o ft h eo t he rm ic ro t ine sp r e sen t .

F o r s imp l ic i t y , t h e a u tho rh a s r e ta ined t h e g enu s d e s igna t ion M ic ro tu s f o r t h i s s pec ie s . I ti ss ome t ime s c l a s s if ied i n a s e pa ra te g enu s C h ionomy s , o r i n as ubgenu s M ic ro tu s ( Ch ionomy s )n i va l i s . M i cro tu so e conomu s( a r c t ic v o le ;r o o tv o le ) ( F igu r e 3 . 6 ) T h i s a bundan t s p ec ie s t o day h a s aH o la r c t ic d i s t r ibu t ion , f r om E u rope i n to C anad a . I t s s ou thwa rd r a nge d oe sn o ti n c lud eS pa in ,w i th i t sp r e sen ce i n d ica t ing ad ef in i te c o ld c l ima t e . T h i sm ed ium v o le s p ec i e sa ve raged 3 1 .8 g( n= 3 06 , r a ng e 1 0 -65 g ) f o r ap opu la t ion i n c lud ing j uven i le s i n t h eN e th e r land s( L ig tvoe ta nd W i jnga a rden 1 994 ) . I tp r ef e r s w e t , m a r shy a r ea s a nd h um id g r a s s land ( Co rbe t 1 966 :246 ; C o rbe ta nd O venden 1 982 :16 1) . T und ra a nd w ood ed s t eppe a r e a l so o c cup ied . I ta l so p r ef e r s m o i s t g r a s s land m ic ro h ab i ta t s s u ch a s r e edbed s , b u t c an b e f o und i n d rye r d i ke s l ope s a nd a mong d une s , h edge row s a nd f i e ld s d u r ing p eak y ea r s a nd i nt h e a b sence o fd i r ec t c ompe t i t ion f r om s p ec ie s s u ch a sM i c ro tu s a r va l i s ( L ig tvoe t a nd W i jngaa rden 1 994 ) . A l though s l i gh t ly l a rg e r o n a ve rag e t h an M . a r va l i s , M . o economu st e nd s t ob e o u tcompe t ed d ue t oi t s mo r e s o l i ta ry l i f e s ty le : M . a r va l i s t e nd s t o l i ve i ns ma l l g r oup s a nd t h u s d om ina t e s a gon i s t ic e ncoun t e r s b e tween t h e t wo s p ec ie s . Whe r e m od e rn r a nge s o ve r lap ( s uch a s i nt h e N e the r land s ) M . o economu s h a s t h e a dvan tage i nh um id m ic rohab i ta t s , w h ich a r e s i gn if i can t ly c oo le r t h an t h e d ry g r a s s land

B ody s i ze o f t h i s s p ec ie s i ss ma l le r a mong t h e m ic ro t ine s r e levan t h e r e . N ie thamme r a nd K rapp ( 1 982 :110 ) r e po r t a m ean m a s so f1 6 .4 g( n =3 0 , r a nge : 1 3 .2 -23 .2g ) f o r mu seum s p ec imen s f r om F i n land . T h i s s amp le i n c lude s ma le s , f e m a le s a nd j uven i le s a nd t h u s r ef l ec t s mo r e c l o se ly t h e a ve r a g e ma s s o ft h o se i n d iv idua l s l i ke ly t o f a l l p r ey t o as ma l l p r eda to r . Wh i le C . g la reo lu s p r ef e r s m a tu r e d ec iduou s f o re s t s w i th t h ick g round c ove r , i t s h ab i ta t s a l so e x tend m a rg ina l ly i n to g r a s s land , h edge row s a nd c on if e rou s f o r e s t s . T h i s s pec ie s i s n o rma l ly c r epu scu la r , s op r eda t ion b y n oc tu rna l a nd d i ur na l p r eda to r s i sc ommon . T he b r eed ing s e a son e x tend s f r om A p r i l t h rough O c tobe r . G e s ta t ion p e r iod s r a nge f r om 1 8 t o 3 5

a r e i n c luded . I ti sf ound mo s t f r equen t ly i nl ow land s , a l ong t h e b ank s o fs l ow w a te r a va i lab le y ea r r ound . A . t e rre s t r i s s ome t ime s l i ve s a way f r om w a te r a nd f o rm s e x ten s ive b u r r ow s ( Co rbe t a nd H a r r i s 1 991 :214 ) . T h i s b imoda l l i f e s ty le l e d t oo r ig ina l d e s igna t ion o fw a te r -dwe l l ing ( A . a mph ib iu s )

2 7 d ay s , w i th ap o s i t ive c o r r e la t ion b e tween l e ng th a nd l i t te r s i ze . T h is s i ze r ange s f r om 1 7 y oung , w i th 3 -4 a ve rage . Wean ing o ccu r s a t 1 7 18 d ay s , a nd a l l mo la r s h ave e r up ted b y2 8 -30 d ay sC o rbe ta nd H a r r i s 1 99 1 :195 199 ) . P i tymy sp yrena icu s( Py r enean p i ne v o le )

a nd b u r row ing ( A . t e r re s t r i s ) a s s epa r a t e s pec ie s . T he n ame

( F igu re 3 . 9 ) P .p y rena icu s h a s ar e la t ive ly r e s t r ic ted r a nge

A . a mph ib iu s h a s t e chn ica l p recedence , b u t A . t e r re s t r i s h a s b een r e ta ined d ue t ol o ng t ime u sage .

a nd t oday i sf ound o n ly i nn or thea s ter nS pa in a nd s ou thwe s t er n F rance . N ie thamme r a nd K rapp ( 1982 :443 ) i n d ica te a m ean s i ze o f2 1 g( n =8 ,r a nge 1 7 -25 g )f o rs pec imen s f r om

A . t e r re s t r i s p u r sue s amo s t ly h e rb ivo rou s l i f e s ty le . I nver te -

T h i s s pec ie s i sf ound i nt h e u ppe r r e a che s o fb eech

b ra te s a nd f i sh a r e r a re d i e ta ry i n c lu s ion s ( Co rbe t a nd H a r r is

f o re s t t o 1 500 m i nt h e F rench P y renee s a nd a l so i nS pan i sh p a rk land ( N ie thamme r a nd K rapp 1 982 :446 ) . B o rgh i e ta l .

1 991 :217 ) . A c t iv i ty i sb o th a tn igh t a nd d ay . B reed ing n o r ma l ly o ccu r s f r om A p r i lt h rough S ep tembe r , w i th a n a ve rage o fa bou t 6y oung e ach . G e s ta t ion r a nge s f r om 2 0 -30 d ay s .

S pa in .

( 1994 ) r epo r t t h i s s pec ie s a to ve r 2 , 000 m i n t h e S pan i sh P y renee s .

B ir th w e igh t i sf r om 3 . 5 t o7 . 5 ga nd i n c r ea se s t oa bou t 2 2 g b y 1 4 d ay s . Wean ing o ccu r s a ta round 2 2 d ay s ( Co rbe t a nd

P .p y rena icu s i sn ow c on s ide r ed t ob e ad i s t inc ts p ec ie s f r om P . s a v i i ( Sav i ' s p ine v o le ) , w h ich i sd i s t r ibu ted d i scon t inu o us ly f r om t h e f o rme r s pec ie s ' r a nge i nI t a ly a nd S i c i ly . P .

H a r r i s 1 991 :2 17 ; V en tu ra a nd Go sä lbez 1 990 ) , a t wh ich t ime t h e t h i rd mo la r h a s f u l ly e r up ted a nd s t a r ted t o c ome i n to w ea r . Ma le sd i spe r se b yt h ea ge o f 4m on ths f r om t h e i rn a ta l t e r r i to ry . D om inan t f ema le s s e t t le i nt h e i r n a ta l t e r r i to r ie s , w i th t h e r ema inde r d i spe r s ing ( Co rbe t a nd H a r r i s 1 99 1 :2 15 ) . T he se d i spe r s ing j uven i le s a r e t h e m o s t f r equen t p rey o f r ap to r s a nd c a rn ivo r e s , a st h ey t e nd t ob e e xc luded t o mo re o pen m ic rohab i ta t s a nd mu s t f o rage o ve r g r ea te r d i s tance s f o rf ood .

p y rena icu s i sf o s so r ia l a nd h e rb ivo rous , p r ef e r r ing d eep s o i ls f o rt unne l ing a nd d en se v ege ta t ion c ove r . I na r ea so fo ve r lap w i th P . l u s i tan icu s , t h e l a t te r s pec ie s i sc ompe t i t ive ly e x c l uded t o a r ea s o f s pa r se r v ege ta t ive c ove r ( Bo rgh i e t a l . 1 994 ) . B reed ing o ccu r s f r om A p r i l t h rough N ovembe r . L i t te r s i ze i ss omewha ts ma l lf o r ar oden to ft h i s s i ze ,w i th ar a nge o f 1 3a nd am ean o f2 . 5 ( Ca s t idn a nd G o sä lbez 1 995b ) .

A rv ico la s ap idu s( s ou thwe s ter nw a t e rv o le ) ( F igu re 3 . 12 )T h i s s pec ie s i sf ound o n ly i nt h e I b e r ian p en in s u la a nd s ou thwe s te rn a nd c en t ra l F rance . T he r ange o fA . s ap idu s o ve r lap s w i th t h a t o f A . t e r r e s t r i s o n ly i n c en t ra l F rance , C an tab r ian S pa in a nd t h e P y renee s . N ie thamme ra nd K rapp ( 1982 :212 ) i n d ica te a mean s i z e o f 2 0 1 g( n =2 8 , r ange : 1 65 -275 g ) f o r s pec imen s f r om F rance , w h ich i n c l ude s n o t f u l ly g rown i n d iv idua l s . T he i r h ab i ta t i sa lway s a l ong w a te r s u ch a sr i ve r s , c ana l s a nd l a ke s w he re t h e c ove r v ege ta t ion i s s uff ic ien t ( Co rbe t a nd O venden 1 982 :166 ) . T he e co logy o ft h i s s pec ie s i sv e ry s im i la r t ot ha t o fA . t e r r e s t r i sd i scu s sed a bove .

P i tymy sl u s i tan icu s( Lu s i tan ian p i ne v o le ) ( F igu re 3 . 10 ) P . l u s i tan icu s i sf ound o n ly i nt h e n o r th a nd w es t I be r ian p en in su la , i n c lud ing n or the rn P or tuga l a nd S pa in a s f a r a s t h e w e s te rn P y r enee s . N ie thamme r a nd K rapp ( 1982 :478 ) r epo r t am ean s i ze o f 1 6 .3 g ( n =2 0 , r ange : 1 0 .0 -23 .0 g ) f o r s pec imen s f r om b o th S pa in a nd P o r t uga l . P ref e r red h ab i ta t s a r e o pen m eadow s , a g r icu l tu ra l l a nd a nd s h rub land ( N ie thamme r a nd K rapp 1 982 :482 ) . P . l u s i tan icu s i sf o s so r ia l , p r ef e r r ing s ha l low s o i l s w i th l ow p ropo r t ions o fr o cky o u tc rop s ( Bo rgh i e ta l . 1 994 ) . R ecen t o w l p e l le t a na ly s i s f r om S pa in ( s ee C hap te r 4 ) c o r robo ra te s t h e se r e su l t s , w i th P . l u s i tan icu s c omp r is ing a s i gn if i can t p e rcen tage o fp rey i n ac oa s ta l l ow land z one o fm ixed p a s t u re , meadow , a nd s ma l lp a tche so fd ec iduou sf o r e s t .

P regnan t f ema le s a ppea r i n t h e p opu la t ion f r om F eb rua ry t h rough O c tobe r . T he se r e ach s exua lm a tu r i ty a ta bou t 1 70 g b ody ma s s . L i t te r s i ze r ange i s2 -5 , w i th a n a ve rage o f3 . 3 ( Ven tu ra a nd G o sä lbez 1 987 ) .

T he r ep roduc t ive b i o logy o ft h i s s pec ie s i sl e s sw e l lr e po r ted t han f o ro t he rm ic ro t ine s pec ie s( wh ich g ene ra l ly h ave l a rge r r ange s ) . T he b reed ing s e a son e x tend s f r om s p r ing t h rough

RODENT IA ,F AM ILY MUR IDAE T he r oden t f am i ly Mu r idae i sc omp r i sed o fO ld Wo r ld m ice a nd r a t s . S ome a u tho r s ( Co rbe t a nd O vend en 1 982 ) p l ace

f a l l , w i th t h e h ighe s t r e p roduc t iv i ty i ns p r ing ( Ve iga 1 986 ) .

t h e s ubf am i l i e s M ic ro t inae ( vo le s a nd l emm ing s , a bove ) ,

G e s ta t ion l a s t s f o r a bou t 2 1 d ay s , w i th mu l t ip le l i t te r s i na s ea son b e ing t yp ica l .

S pa lac inae ( mo le r a t s ) , a nd C r ice t ina e ( h am s te r s ) u nde r t h e f am i ly Mu r idae w i th t h e Mu r inae . F o r t h e s ake o fc l a r i ty , e ach o ft h e se g roup ing s w i l l b e t r ea ted a ss epa ra te f am i l ie s

A rv ico la t e r re s t r i s( no r ther nw a t e rv o le )

u n t i lt h e i re vo lu t iona ry r e la t ion sh ip s c an b e b e t te r s or ted o u t .

( F igu re 3 . 11) A . t e r re s t r i s t o day o ccup ie s al a rge r a nge , i n -

T he f am i ly Mu r idae i sn o tab le f o r i t s c ommensa l i sm a nd r e su l tan t c osmopo l i tan r ange s p read o f h ou se mou se (Mu s

c l ud ing n or the rn t empe ra t e E u rope a nd s t re tch ing i n to S ibe r i a e a s t o fL ake B a ika l a nd s ou th t oI r aq . I t s d i scon t inuou s r ange i n c lude s p opu la t ion s i nB r i ta in a nd C an tab r ian S pa in ,

m u scu lu s ) , b l a ck r a t s ( Ra t tu s r a t tu s ) a nd b rown r a t s ( R . n or v eg icu s ) . T he e a r ly s p read o ft h e se s p e c ie s may b e l i nked t o

b u t e xc lude s t h e Med i te r ranean a nd w e s te rn F rance . C o rbe t

t h e s p read o f n a tu ra l ly a nd a r t if i c ia l ly c r ea ted g ra s s land s

a nd H a r r is ( 1991 :214 ) i n d ica te a m ean s i ze o f 3 11 g f o r ma le s ( n =1 8 ) a nd 2 72 gf o rf ema le s ( n =1 7 ) , i nb o th c a se s f o r o ve rw in te red a du l t s f r om O xf o rd sh i re , E ng land . T he se

( ag r icu l tu ra l f i e ld s a nd o t he r a l te ra t ion o f l a nd scape ) . F o r e xamp le , M . m u scu lu s s how s u p i nI s r ae l f o r t h e f i r s t t ime a round 1 20 ,000 B .P . ( Tche rnov 1 975 ) a nd R . r a t tu s d u r ing t h e l a te Mou s te r ian ( Tcher nov 1 979 ) . A r r iva l i n S pa in o f t h e se s pec ie s p robab ly d id n o t o ccu r b ef o re N eo l i th ic t imes

s i zes may t h e ref o re b e t a ken a s max ima l a du l t w e igh t s , w i th al owe r p opu la t ion m ean e xpec ted i fj uven i le s a nd s ubadu l t s 3 6

T h i s s pec ie s i sa c t ive m o s t ly a tn i gh t . A .f l a v i co l l i s f o rm s e x ten s ive b u r row s y s t em s , i nw h ich i tm ay s t o r e f o od . I n w in te r , n e s t ing i ns ma l l g roup sm ay o ccu r . I t s d i e ti ss im i la r t oA . s y lva t icu s a nd i n c lude s av a r ie ty o f p l an t a nd s ome a n ima l ma te r ia l s . L i t t e r s i z e r a nge s f r om 2 t o 1 , a nd b r eed ing o ccu r s f r om F eb rua ry t h rough O c tobe r . G e s ta t ion a nd w ean ing a r e s im i la r t ot h o se o fA . s y l va t i cu s , w i th b o th a r ound 2 0d ay s( Co rbe ta nd H a r r i s 1 99 1 :232 ) .

a nd h ence t h e i r a ppea rance i np a leo l i th ic l e v e l s t h e r e w ou ld b es t rong i n d ica t ion o fm ix ing o fd epo s i t s . P hy s i ca l c ha ra c t e r i s t ic s i n c lude l a rge e x te rna l p i nna e , ar e la t i ve ly l o ng t a i l , a nd al o nge r , t h inne rb ody t h an v o le s . D en t a l f o rmu la r ema in s 1 . 0 .0 .3 / 1 .0 .0 .3 , w i th c on t inuou s ly g r ow ing i n c i so r s a nd r oo t ed mo la r s w i th m any s ma l l c u sp s a r rang ed i ns t agge r ed r ow s .

M ic romy sm inu tu s( h a rve s tm ou se ) ( F igu re 3 . 15 ) M . m inu tu s r a nge s a c ro s s t h e t empe ra t e P a lea r c t ic z one f r om S pa in t oK o r ea . T h i s r a ng e m ay h ave b een e x tended i ns ome a r ea s w i th t h e s p r ead o fa g r icu l tu r e , s i nce t h i s s pec ie s i n hab i t s t h e s t a lk z one i ng r a s s land s a nd h a s r e ad i ly a dap t ed t oc r op land s ( T rou t 1 978 ) . T a l l , d en se g r a s s land s a r e p r ef e r r ed , t h ough s ome s p i l love r i n to m a tu r e f o re s t s h a s b een r e po r ted . T he g round l e ve l r u nway s o f o t he r r oden t s a r e i n hab i t ed i n w in t e r ( Co rbe t a nd H a r r i s 1 99 1 :236 ) . P reda t ion t h e ref o r e s e em st oi n c r ea se i na u tumn w hen l e s s c ove r i sa va i lab le a nd j uven i le s d i spe r se t ot h e i r n ew t e r r i to r ie s . A c t iv i ty i sn o rma l ly n oc tu rna l , w i th p eak s a f t e rs un se t a nd b ef o r e d awn . T h i s e l u s iv e s p ec ie s i s ar e gu l a rb u tn ume r ica l ly l ow p r ey i t em ,n eve rr e ach ing > 1% o ft h e d i e t o fB r i t i sh r a p to r s ( G lue 1 975 ) . I t s i mpo r tance t or a p to r d i e t i sf u r the r r e duced b y i t s s ma l l a ve rage b ody m a s s o f o n ly 4 10 g .

A pod emu ss y l va t icu s( wood mou se ) ( F igu r e3 . 13 ) A .s y lva t i cu st oday h a sa n e x t en s ive r a nge t h a t h a s p e rhap s b een e x tended b y c ommen sa l c on ta c t w i th h u m an s , a se v id enced b y t h e p r e sence o ft h i s s p ec ie s o n I c e l a nd . I t sr a nge a l so i n c lud e s a l l o ft empe ra t e a nd Med i t e r r a n ean w e s t er n E u rope , a sw e l l a s p a r t o fN o r th A f r ica , T u r k ey ,a nd e a s twa rd t h rough R u s s ia p a s tt h eU ra l s . C o rbe ta nd H a r r i s( 1 99 1 :222 )r e por t am ean s i z eo f1 9 .1 gf o rm a le s( n= 2 0 ) a nd 1 7 .8 g f o r f ema le s ( n =1 3 ) f r om P e r th sh i r e , E ng l a nd . T r app ing i nC an tab r ian S pa in ( s ee C hap t e r4 )r e su l t ed i na n a ve rage s i ze o f2 1 .3 gf o r m a le s ( n =2 6 , r a nge : 1 4 -32 g ) a nd 1 9 .1 g f o r f ema le s ( n =2 1 , r a nge : 1 2 -25 g ) f o r a m o s t ly a du l ts amp le . A .s y l va t icu s i s ah ab i ta tg ene ra l i s ta nd c an b e f o und i nm o s tt y pe s i ft h ey a r en o tt o ow e t , i n c lud ing w ood land , c r op land , g r a s s land , h ea the r , b l anke t b og a nd s a nd d une s ( Co rbe ta nd H a r r i s 1 99 1 :224 ) . D ec iduou s f o r e s t , h oweve r , i st h e i r f a vo r ed h ab i ta t . T h i s s p ec ie s i sa l so c om m en sa l a nd w i l l f o l low h uman s h e l t e r s t o h i ghe r a l t i tude s t h an t h e i r n o rma l t r ee l ine l i m i t ( Co rbe t a nd O vend en 1 982 :170 ) .

T he b r eed ing s e a son o f h a rve s t m ice e x tend s f r om May t h rough O c tobe r ,w i th m o s tl i t t e r sp r oduced a tt h ee nd o ft h i s p e r iod . L i t te r s r a nge f r om 4 t o8 ,w i th a n a ve rage o f 5 . 4 . G e s ta t ion l a s t 1 7 19 d ay s , a nd t h e y oung a r e w eaned a t1 5 1 6 d ay s . L i t t e r s a r e r a i sed i nn e s t s i nt h e s t a lk z one , w h ich f u r the r r e duce s t h e i r s u scep t ib i l i ty t o p r eda t ion b y r a p to r s u n t i ld i spe r sa lo ccu r s( Co rbe ta nd H a r r i s 1 99 1 :237 ) .

A c t iv i ty i sm a in ly n oc tu rna l , t h ough t h i s t e nd s t o b ecome b i pha s ic d u r ing l o ng n i gh t s . A . s y l va t i cu s f o rm s e x t en s ive b u r row s y s t em s a nd u sua l ly n e s t s b e low g round . D ie t i so p p o r tun i s t ic : s e ed s , v ege ta t ion , f r u i t , a nd e ven a n ima l f o od s a r e a l c on sumed ( Co rbe t a nd H a r r i s 1 99 1 :226 -227 ) . L i t t e r s o f4 7 a r e c ommon a nd a r e p roduced f r om Ma r ch t h rough O c tob e r . G e s ta t ion l a s t s 1 9 -20 d ay s ,w i th w ean ing o ccu r r ing a t1 8 -22 d ay s . F ew a du l t s s u rv ive f o rm o r et h an o ne w in te r . T h i s s p ec ie s i sc ommon o w l p r ey , e s pec ia l ly d ue t o t h e o ve r lap o fa c t iv i ty .

I NSECT IVORA ,F AM ILY S OR IC IDAE T he t e e th o f t h e se s p ec ie s a r e d i s t ingu i shab le o n m u l t ip le g round s . T he t e e th o fS o rex h ave ar e d c o lo r o n t h e i r c u sp s w h ich i sh i gh ly d i s t inc t ive u n le s s s e ve r e a t t r i t ion h a s o c c u r red . D en ta l f o rmu la f o l low s 3 . 1 .3 .3 / 1 . 1 .1 .3 . S .m inu tu s i s much s ma l le r t h an S .c o rona tu s . S .a raneu s m ay b e d i s t i ngu i shed f r om t h e l a t t e r s p ec ie s b y s l igh t d i f f e r ence s i n m and ibu la r mo rpho logy ( Hau s se r a nd J ammo t 1 974 ; N ie th a mme ra nd K rapp 1 990 ; T u rn ia nd Mü l le r 1 996 )w he r et h e se f e a tu r e s h ave b een r e ta ined i n a r cha eo log ica l a nd p a leon t o log ica l m a te r ia l s . T he t e e th o f N eomy s e xh ib i t t h i s r e d s t a in ing t o al e s se r d eg r ee a nd f o l low a3 . 1 .2 .3 / 1 .1 .1 .3 f o r mu la . T ee th o fN .f od ien s a nd N . a noma lu s a r e d i s t ingu i sh a b le f r om e a ch o t he r p r ima r i ly u pon m e t r ica l g round s ( Rup rech t 1 97 1) . T he t e e th o f C roc id u ra f o l low a 3 . 1 .1 .3 / 1 .1 .1 .3 f o rmu la a nd l a ck r e ds t a in ing o ft h e i rc u sp s .

A pod emu sf l av i co l l i s( y e l low -neck ed m ou se ) ( F igu r e 3 . 14 ) A . f la v i co l l i s h a s ad i scon t inuou s r a nge t h roughou tt empe ra te E u rope , i n c lud ing i s o la t ed p opu la t ion s i n n o r th er n S pa in a nd t h e P y r enee s , s ou ther n E ng land , s o u the rn N o rway a nd S weden , a nd t h eL evan t . T he Med i t e r r a n ean a r ea i sp opu la t ed i nI t a ly a nd G r eece . S i z e r a nge s u p t o 1 . 5 t i me s t h a t o f A . s y l va t i cu s ( Co rbe t a nd H a r r i s 1 99 1 :229 ) . T rapp ing d a ta f r om C an tab r ian S pa in ( s ee C hap t e r4 )y i e lded m a s se so f2 5 .1 gf o rm a le s( n =1 6 ,r a ng e : 1 7 -34 g )a nd 2 3 .4 gf o r f ema le s ( n =1 5 , r a nge : 1 6 -32 g )f o r ap opu la t ion i n c lud ing j uven i le s . T he se m ean s a r e o n ly s l igh t ly a bov e t h o se o fA . s y lva t icu s t r apped i nt h e s ame l o c a t ion s i nC an tab r ian S pa in ( s ee a bove ) . A .f la v i co l l i s i n h ab i t s m a in ly d ec iduou s w ood land s a nd m a rg ina l ly e x tend s i n to h edge s , r u ra l g a rden s a nd b u i ld ing s , b u t i sn o t at r ue c ommen sa l . I ti sm o r e c ommon t h an A .s y l va t icu s i na l p ine a nd c on if e rou sf o r e s t sa nd l i ve su pt ot h et r ee l ine , a nd i sl e s s c ommon i ns c rub a nd f i e ld s ( Co rbe t a nd H a r r i s 1 99 1 :230 -

C roc idura r u s su la ( g r ea te r w h i t e too thed s h r ew ; h ou se s h rew ) ( F igu re 3 . 16 ) T h i s s p ec ie s i sf o und t o day o n ly i n We s t er n E u rope a nd N o r th A f r ica , i n c lud ing t h e c oun t r ie s o f S pa in , P o r tuga l , F rance , a nd Mo r rocco . B ody m a s s f o r as amp le o f m a le a nd f ema le a du l t s f r om w e s te rn S w i tz e r land a ve raged 1 .4 g ( n =2 1 , r a nge : 9 . 0 16 .1 g ) ( N ie thamme r a nd K rapp 1 990 :432 ) . S even i n d iv idua l s t r apped i n C an tab r ia ( s ee C hap te r4 )y i e lded am ean m a s s o f7 . 9 g( 6m a le s , 1f ema le , r a nge : 6 . 5 10 g ) . C .r u s su la i ss e pa rab le f r om C .s u c r veo len s

2 31) .

3 7

s umme r s now l ine ( Co rbe t a nd O venden 1 982 :124 ) .

( l e s se r w h i te too thed s h rew ) o n ly b y d en t i t ion a nd i so t he r w i se s l igh t ly l a rge r i n mo s t d imen s ion s . C . r u s su la i sa h ab i ta t g ene ra l i s t a nd c an b e f ound i n f o re s t s , h edge row s , g ra s s land , a g r icu l tu ra l l a nd s , maqu is s c rub land a nd a mong s t

T he

p r ima ry a s soc ia t ion o fS .c o rona tu si sw i th h um id m eadow s . T he b r eed ing s ea son f o r t h i s s pec ie s e x tend s f r om l a te J anu -

h uman h ab i ta t ion s( Co rbe ta nd H a r r i s 1 99 1 :74 ) .

a r y u n t i l m id -Novembe r ( Löpez -Fu s te r 1 989 ; L öpez -Fu s te r e ta l . 1 988 ) , s oi n d iv idua l sb or ne a r ly i nt h e y ea ra r e l i ke ly t o r e ach s exua l ma tu r i ty a nd r ep roduce b ef o r e t h e y ea r e nd s .

T h is e s sen t ia l ly s o l i ta ry s pec ie s i sa c t ive b o th d ay a nd n i gh t a nd p u r sue st h e t yp ica l s h rew d i e t o fp r ima r i ly i n ve r teb ra te s a nd af ew s ma l l v er teb ra tes . A ve rage l i t te r s i ze i s4 ,w i th a

L i t te r s i ze r ange s f r om 2 -6 w i th am ean o f3 . 7 . G e s ta t ion l a s t s f o r 2 4 -25 d ay s . J uven i le s l e ave t h e n e s t a f t e r 2 0 -25

r a nge o f2 11 , a nd b r eed ing o ccu r sf r om a se a r ly a sF eb rua ry

d ay sa nd h ave at o ta l l i f e e xpec tancy o f1 4 -2 1 m on th s .

t h rough O c tobe r . G e s ta t ion l a s t s f r om 2 8 -33 d ay s . B i r th w e igh t i s0 . 8 -0 .9 g ,w i th w ean ing o ccu r r ing a t2 8 d ay s . S ex u a l ma tu r i ty i sr e ached a t5 8 -71 d ay s , s o b r eed ing d u r ing t h e

N eomy sa noma lus ( M i l le r ' sw a te rs h rew ) ( F igu r e 3 . 19 ) N . a noma lu s h a s ad i scon t inuou s d i s t r ibu t ion

s e a son o f b i r th i sc ommon ( Co rbe t a nd H a r r i s 1 991 :74 ) . D en s i t ie sh ave b een r e co rded t op eak a t7 7 100 /ha i nE u rope

t h rough E u rope , f r om S pa in i n to R u s s ia . I t s n o r the rn r ange

d u r ing t h e s umme r b r eed ing s e a son ( Co rbe t a nd H a r r i s

i n to G reece . N ie thamme r a nd K rapp ( 1990 :318 ) r e por t a m ean s i ze o f 1 0 .9 g ( n =9 ,r ange : 1 0 .1-12 .3 g ) f o r mo s t ly j uven i le mu seum s pec imen s f r om G e rmany . T he p r ima ry h ab i ta t o ft h i s s pec ie s i sa l ong w a te r c ou r se s , b u t i ta l so c an b e f o und a way f r om t h em i n h um id f o r es t s a nd m eadow s ( Co rbe t a nd O venden 1 982 :127 ) . N a noma lu s i sf o und b e t ween 1 80 -1 ,850 m a . s . l . a nd n o rma l ly b e low 1 , 000 m t oday i nn o r thea s te rn S pa in ( To r re a nd T e l la 1 994 ) .

e x tend s a s f a r a s G e rmany , a nd i t s s ou the rn r a nge e x tend s

1 991 :75 ) . T h i s s p ec ie s i sc ommon r ap to r p rey i nS pa in , b u t w a s am uch r a r e rc omponen to fP l e i s tocene f a una s . S orex m inu tus ( pygmy s h rew ) ( F igu re 3 . 17 ) S .m inu tu s t oday r a nge s t h roughou t t empe ra te E u rope , f r om n or the rn P o r tuga l t o L ake B a ika l i n S i be r ia , a nd e x tend s f r om s ou the rn G reece t o n or ther n S cand inav ia . I t s a ve rage s i ze i so ne o ft h e s ma l le s ta mong mamma l s , w i th am ean o f3 . 6 g( n =2 0 ,r ange : 2 . 6 -5 .2 g )f o rs pec imen sf r om G e rmany a nd F rance ( N ie thamme ra nd K rapp 1 990 :193 ) . I t i sf ound i na l l h ab i ta t t ype s , t hough p r ima r i ly i nt ho se w i th a bundan t g round c ove r . S . m inu tu s p ref e r s g ra s s land o ve r w ood land , a nd w h i le l a rge ly g round -dwe l l ing , i tw i l l c l imb f o l iage ( Co rbe t a nd H a r r i s 1 99 1 :62 ) . I tt o le ra te s l owe r g round c ove r t h an S .c o rona tu s , a nd i t s r ange e x tend s e ven t o mon tane h ea the r( Co rbe ta nd O venden 1 982 :125 ) .

T he b r eed ing s ea son o f N . a noma lu s e x tend s f r om Ma rch t h rough S ep tembe r , w i th t h e f i r s t l i t te r s p roduced b y t h e e nd o f May . L i t te r s i ze r ange i sf r om 5 13 ( N ie thamme r a nd K rapp 1 990 :332 ) . Wean ing o ccu r s a t2 9 -30 d ay s ( M icha lak 1 987 ) . N eomy sj od iens ( wa te rs h rew ) ( F igu r e 3 . 20 ) N .f od ien sr anges f r om n or ther nS pa in t oL ake B a ika l i nS ibe r ia . I t s r ange i n c lude s E ng land a nd e x tend s t o n o r ther n S cand inav ia a nd i n to t h e Med i ter ranean r e g ion i n I t a ly . I s o la ted p opu la t ion s c an a l so b e f ound i nt h e f a r E a s t , i n c lud ing t h e K o rean p en in su la a nd S akha l in I s land . C o rbe t a nd H a r r i s ( 199 1 :65 ) i n d ica te at yp ica l s i z e r a nge o f 1 2 18 g f o r a du l t s o f t h i s s pec ie s . F ou r s pec imen s t r apped i n C an tab r ia ( s ee C hap te r 4 ) h ad a n a ve rage ma s s o f 1 2 .5 g ( 1 m a le , 3f ema les , r a nge : 1 0 -17 g ) . I t s h ab i ta tp r ef e r ence s a r e m a in ly t h e b ank s o f c l ea r , f a s t -f low ing w a t e r s a nd a l so b y p ond s . I ti se spec ia l ly n ume rou s i nw a te rc re s sb ed s . N .f od i e n s i sa l so f ound a mong b ou lde r s o n r o cky b ea che s a nd s po -

T h is s pec ie s i sa c t ive b o th d ay a nd n igh t , w i th p eak s a r ound 0 800 -1000 a nd 2 100 -2300 ( Co rbe t a nd H a r r i s 1 991 :62 ) . S . m inu tu s i sb o th s o l i ta ry a nd a gg re s s ive , a nd t hu s r e ache s t yp ica l d en s i t ie so fo n ly 1 2 /ha , w i th am ean o f6 /ha f ound i n E ng land ( Co rbe t a nd H a r r i s 1 99 1 :63) . T he d i e t c on s is t s p r i m a r i ly o f i n ver teb ra te s . T h i s s pec ie s ' e ne rge t ic n eed s a r e s uch t h a t i t mu s t c on sume 1 . 25 t ime s i t s o wn b ody w e igh t e a ch d ay . B reed ing o ccu r s f r om Ma rch t h rough O c tobe ra nd p eak s i nJ une / Ju ly . F ou rt os even p e r l i t te r a r e t yp ica l , a f t e r ag e s ta t ion p e r iod o f 2 2 -25 d ay s . T he se n ewbo rns w e igh

r ad ica l ly a way f r om w a te r i n d ec iduou s w ood land , h edge r ow s , g r a s s land , a nd o cca s iona l ly i n c on if e rou s w ood land ( Co rbe t a nd H a r r i s 1 991 :65 ) . I ti st h e mo s t a qua t ic o f t h e E u ropean s h rew s pec ie s , a nd i tc an b e f ound u pt o2 500 m i n

o n ly 0 . 25 ga nd a r ew eaned a ta bou t2 2 d ay s . T hey g ene ra l ly d o n o t o b ta in s e xua l m a tu r i ty u n t i l a f te r t h e i r f i r s t w in te r . L if e span i nt h ew i ld i sa pp rox ima te ly 1 3 mon th s . S .m inu tu s i sc ommon p rey o fb a rn o w l s a nd t awny o w l s , t hough i nl ow n umbe r sr ef lec t ive o fi t sl ow o ve ra l ld en s i ty .

t h e S w i s sA lp s( Co rbe ta nd O venden 1 982 :126 ) .

S orex c orona tu s( M i l le t ' s s h rew ) ( F igu re 3 . 18 ) S .c o rona tu s i sf ound o n ly i nn o r the rn S pa in

N .f od ien s i sa c t ive b o th b y d ay a nd n i gh t , t h ough n i gh t a c t i v i ty p r edom ina te s w i th ap eak j u s t b ef o r e d awn . I n d iv idu -

a nd t h roughou t F rance , t h e L ow C oun t r ie s , a nd i n to S w i t z e r land . I ti s as i b l ing s pec ie st oS .a raneu s , w ho se n on sym -

a l s a r e s o l i ta ry , b u t l i ve i nc l o se p rox im i ty . T he i r d i e t c on -

p a t r ic r a nge e x tend s e a s twa rd t h rough E u rope . As amp le o f mo s t ly a du l t S . c o rona tu s f r om Sw i tz e r land ( N ie thamme r a nd K rapp 1 990 :238 ) a ve raged 7 . 8 g( n =1 9 , r ange : 6 . 5 -

a qua t ic v e r teb ra te s . F o rag ing o ccu r s i nt h e w a t e r , a nd t h i s s pec ie s w i l l a l so f o rm e x ten s ive b u r row s ( Co rbe t a nd H a r r i s

1 .75 g ) .

t h an o t he r s pec ies o f t h i s s i ze d i scu s sed a bove . B reed ing o ccu r s f r om A p r i l t o S ep tembe r a nd p eak s i n May / June . A ve rage l i t te rs i ze i s6 ,r ang ing f r om 3 15 . G e s ta t ion i st yp i c a l f o r amamma l s pec ie s t h i s s i ze a nd l a s t s f o r 1 9 -2 1 d ay s , b u tw ean ing t ime i sl o nge r , a t3 8 -40 d ay s ( Löpez -Fu s te r a nd

s i s t s p r ima r i ly o f i n ver teb ra te s a nd a l so i n c lude s s ma l l

1 99 1 :65 -67 ) .

F ive s pec imen s t r apped i n n o r the rn S pa in ( s ee

C hap t e r 4 ) h ad a n a ve rage ma s s o f 7 . 4 g( 2 ma le s , 3f e m a le s ) . S .c orona tu s c an b e f ound i na l l h ab i ta t s w i th s uf f i c i en t c ove r , i n c lud ing f o re s t s , h edge row s , d une s , f i e ld s a nd h i gh , b a r ren p l a in s . I t s mon tane h ab i ta t s e x tend a sf a ra st h e

T he b reed ing p a t ter n i ss omewha t d i f f e ren t

V en tu ra 1 995 ) . A max imum o f 3l i t te r s a r e t h e ref o r e p o s s i 3 8

b l e d u r ing ab r eed ing s e a son , a nd f ema le s n o rma l ly b r eed o n ly a f t e ro ve rw in te r ing . L if e span i st y p ica l ly n om o r et h an 1 4 19 mon th si nt h ew i ld ( Co rbe ta nd H a r r i s 1 99 1 :67 ) . T h i s s p ec ie s d oe s n o t n o rma l ly o b ta in l a rge d en s i t ie s , a nd i s r e gu la r( b u tn o ta bundan t )p r ey o fb a rn o w l sa nd t awny o w l s .

i nt h ep opu la t ion f r om O c tobe rt h rough A p r i l , w h i le n u r s ing f ema le s a r e f o und f r om N ovembe rt h rough May ( J im6nez e t a l . 1 990 ) . S exua lm a tu r i ty i nb o th s e xe s i sr e a ched a f t e ro ne y ea r . T he l o ng b r eed ing s e a son p e rm i t s mu l t ip le l i t te r s p e r y ea rf o r s ome f ema le s . T he d u ra t ion o fg e s t a t ion a nd l a c t a t i on i n t h i s s p ec ie s i sp robab ly s im i la r t o t h o se o f T . e u r opaea ,t h ough t h e se r ema in u n r epo r ted .

I NSECT IVORA ,F AM ILY TALP IDAE T a lpa e uropaea ( no r ther n mo le ) ( F igu r e 3 . 2 1) 7 ' .e u ropaea c an b ef o und f r om n o r the rn S pa in t oE u ropean R u s s ia , b u t i t s r a nge d oe s n o t i n c lude n o r th e rn S cand inav ia o rt h e Med i t e r ranean r e g ion ,e xcep tf o rs ou the rn F rance . B r i t a in i si n c luded i nt h i s r a ng e . S i z e i ss omewha t v a r iab le . C o rbe t a nd H a r r i s ( 199 1 :46 ) r e po r t am ean s i z e o f 10 gf o rm a l e s( n =4 2 ) a nd 8 5 gf o rf ema le s ( n =5 7 ) f r om S uf f o lk ,E ng land . Mau r iz io a nd H au s se r( 1 990 :225 )i n d ica t e am ean s i z eo f6 3 g( n =3 6 , 1 8m a le sa nd 1 8f ema le s ,r a nge : 5 1 -88 g )f o r ap opu la t ion f r om V a l B r egag l ia , I t a ly . T .e u r opaea i sp r e sen ti n mo s th ab i ta t sw he r et h e s o i lp e rm i t st u n n e l ing . I tw a so r ig ina l ly ad ec iduou sw ood land s p ec ie sb u ti s n ow f ound i np a s tu re s a nd c r op land a nd i su ncommon i n c on if e rou sf o r e s t s , moo r land , a nd d une s .

G a l emy sp yrena icu s( Py renean d e sman ) ( F igu re 3 . 23 ) G . p y rena icu s i se ndem ic t o t h e P y r enee s , t h e i r w e s twa rd e x ten s ion ( t he C an tab r ian C o rd i l le ra ) , a nd h i gh land s f u r the rs ou th i nn o r the rn P o r tuga l a nd t h e S pan i sh m e se ta . T h i s s p ec ie s b e long s t ot h e f am i ly T a lp idae w i th in t h e s ubf am i ly D e sman inae . T h i s s ubf am i ly c on ta in s o n ly t wo w ide ly s e pa ra te t a xa , G . p y rena i cu s a nd D e smana m o scha ta f ound o n ly i nt h e a r ea n o r th o ft h e B la ck a nd C a s p i an s e a s i nt h e f o rme rE u ropean U .S .S .R . T h i s l a rge i n sec t i vo r eh a s ac r an ium g ene ra l ly s im i la rt ot h a to fT a lpa ,w i th a l o ng r o s t rum c on ta in ing t h e m ax imum p l a cen ta l m amma l ian t o o th p a t te rn ( 3 .1 .4 .3 / 3 .1 .4 .3 ) . T he se t e e th a r e mo rpho log i c a l ly a nd m e t r ica l ly d i s t ingu i shab le f r om t h o se o f T a lpa . B ody m a s s r a nge s f r om 3 5 .5 gt o8 0 g( s umma r iz ed b y P a l m e i r im a nd H of fman 1 983 :1) . T h i s s p ec ie s i st h e r ef o r e i n t h e p rope r s i z e r a nge t ob e t a ken b y m any s p ec ie s o fE u ro p ean o w l s ,b u ti su sua l ly an ume r ica l ly s ma l lp r ey i t em .

Mo s to ft h el i f e span o ft h i ss p ec ie si ss p en tw i th in i t st u nne l s , s o p r eda t ion b y r a p to r s mu s t o ccu r d u r ing t h o se r a r e t i me s w hen t h i s s p ec ie s i sa bove g round . P r eda t ion r i sk t h u s i n c r ea se sd r ama t ica l ly d u r ing t h et ime o fj uven i le d i spe r sa la nd a l so d u r ing m a t ing s e a son w hen m a le s l e ave t h e i r t e r r i to r ia l b u r row st oe ncoun te r f ema le s w i th o ve r lapp ing r a nge s . D ie t c on s i s t s a lmo s t e n t i re ly o fi n ve r t eb r a t e s o b ta ined b y t h e p i t f a l l t r ap e f f ec t o ft unne l e n t rance s o r t h o se o b ta ined w h i le t u nne l ing ,e s p ec ia l ly e a r thwo rm s .

B r eed ing l a s t f r om J anua ry t h rough J u ly , w i th a g e s ta t ion p e r iod o fa bou t 3 0 d ay s . L i t te r s i ze r a nge s f r om 1 5 , w i th a m ean o f3 . 6( P a lme i r im a nd H of fman 1 983 :3 ) . G .p y rena icu s i sa n a l p ine a nd r i ve r ine s p ec ie s ( Ca s t iin a nd G o sä lbez 1 992b ; P a lme i r im a nd H of fman 1 983 ) . I t s a l t i tude z one i sf r om 3 00 t o 2 200 m , t h ough t h e l ow e r e nd o ft h i s r a nge i s mo r e t y p ica l . I t s p r e sence i nn ea r s e a l e ve l i n a c oa s ta l l ow land s i t et h e r ef o re i n d ica te ss i gn if i can td ownwa rd a l t i tud ina l z one s h if t d u r ing c o ld p ha se s , a tl e a s t p l ac ing i t s h ab i ta tw i th in t h e h un t ing r a d iu s o f ap r eda to ri n hab i t ing t h e l ow land z one . T h i s s pec ie s i sw e l l a dap t ed f o r as em i a qua t ic l i f e s ty le ,p o s se s s ing f o u rw ebbed f e e t , ak ee led t a i l , a mob i le s e n so ry s nou t f o r p rob ing s t r eam b o t tom s , a nd i n su l a t ing p e lage . I ti sa s soc ia ted w i th c o ld , f a s t -r u ni ng o xy g ena ted ( t rou t -abundan t ) s t ream s f r om w h i ch i td e r ive s i t s m a in d i e t o fc r u s ta cean s a nd o t he r a r th ropod s . A c t iv i ty i s t y p ica l ly n oc tur na l ,t h ough t h i sm ay e x tend p a s td awn s ome t ime s( p e r sona lo b se rva t ion ) .

B r eed ing i ss l owe r i nt h i s s p ec ie s t h an i no t he r i n sec t ivo r e s . Al i t t e r o f3 4 i st y p ica l ly p roduced i nt h e s p r ing , w i th o ne a dd i t iona l l i t t e r p roduced i nt h e m o r e s ou the r ly p o r t ion s o f t h i s s p ec i e s ' r a nge . B i r th w e igh t r u n s a r ound 3 . 5 g ,a nd t h e n e s t i sl e f ta f t e r o ne mon th . D i spe r sa l o fj uven i le s t h en o c c u r s a ta g e 5 6 w eek s , w i th n ea r ly f u l l g r ow th o b ta ined b y a ge 3 m on th s . S exua l m a tu r i ty i sr e ached t h e f o l low ing s p r ing . L i f e span i nt h ew i ld i su pt o 6y ea r s , b u t l e s s t h an 3 y ea r si sm o r eu sua l( Co rbe ta nd H a r r i s 1 99 1 :47 -48 ) . T a lpa o cc id en ta l i s( I be r ian mo le ) ( F igu r e 3 . 22 ) T h i ss pec ie s 'd i s t r ibu t ion i sl i m i ted t oa l o ft h e I b e r ian P en in su la a nd t h e s ou ther n e dge o f F rance . T he r a nge o f To cc id en ta l i s t h e r ef o r e o ve r lap s w i th t h a t o f T . e u ropaea t h roughou t mo s t o ft h e f o rme r ' s r a nge , e s pec ia l ly i nS pa in . N ie thamme ra nd K rapp ( 1 990 :158 ) r e po r t am ean m a s s o f4 8 .9 g( n =2 0 , r a nge : 3 4 -63g ) f o r ap opu la t ion o f m a le s a nd f ema le s f r om S pa in . I t s h ab i ta t p r ef e r ence s a r e s im i la rt ot h o se o fT .e u ropaea : f i e ld s a nd d ec iduou s f o r e s t s w he r e s u i tab le s o i l s f o r t unne l ing a r e f o und , a nd r a r e ly i n to c on if e rou s f o r e s t s a nd h ea th . T o cc id en ta l i s h a s d emon s t ra t ed k a ryo typ ica ld i f f e r ence sf r om T .c aeca ( J imdnez e ta l . 1 984 ) , w i th w h ich i tw a s p r ev iou s ly c on s ide red t ob e c on s p ec if ic . 7 ' .c aeca h a s ad i scon t inuou s r a nge f r om t h a t o fT . o cc id en ta l i s a nd i sf ound f r om s ou ther n F rance t oI t a ly a nd G r eece . T he r a nge o fb o th o ft h e se s p ec ie s o ve r lap s w i th t h a to f( t he g ene ra l ly l a rge r ) T .e u ropaea .

I NSECT IVORA , F AM ILY ER INACE IDAE E r inaceu se uropaeu s( h edgehog ,w e s t e rn h edgehog ) ( F igu re 3 . 24 ) T h i s s p ec ie s i sd i s t r ibu ted t h roughou t We s te rn E u rope , i n c lud ing s ou ther n S cand inav ia , b u ti ti sr e p laced i n E a s t er n E u rope b y t h e e a s te rn h edgehog ( E . c onco lo r ) . E . e u ropaeu sw e igh sc a . 1 20 ga tw ean ing a nd i n c rea se st oo ve r 1 . 1 k g a s a n a du l t ( Co rbe t a nd H a r r i s 1 99 1 :38 ) . H ab i ta t p r ef e r ence s a r e g r a s s land /wood land e dge s o f a l t y pe s , i n c l ud ing h edge row s . T hey a r e f ound r a r e ly i n c on if e rou s w ood land , m a r she s o r moo r land ( Co rbe t a nd H a r r i s 1 99 1 :40 ) . T h i s s p ec ie s i sp a r t ia l ly c ommen sa l , r e ad i ly e n t e r ing c ampg round s a nd o t he r o pen h uman h ab i ta t ion s a nd r a p id ly l o s ing i t s f e a r o f h uman s ( p e r sona l o b se rva t ion ) . S uch b ehav io r i sa n i mpo r tan t c on s ide ra t ion i nt h e i n te rp r e t a t ion o fa r chaeo log ica lc ave f a una s .

A du l t m a le a nd f ema le T o cc id en ta l i s a r e s e xua l ly a c t ive f r om S ep t embe r t h rough May . P r egnan t f ema le s a r e f o und 3 9

T h i s s p ec ie s i sa lmo s t e n t i r e ly n oc tu rna l , w i th s ome a c t iv i ty a t d awn a nd d u sk . W in t e r h i be rna t ion i sa l so c ommon , t h ough i ti sf r equen t ly i n te r rup ted a nd s ome t ime s b u r row s a r e c hanged . D ie t i s mo s t ly i n ve r t eb ra te a nd a l so i n c lude s s ome c a r r ion a nd b i rd e gg s . O ne l i t t e r o f4 -6 i sn o rma l ly b o rn i nJ une , b u t as e cond o ne m ay b e b o rn l a t e r i nt h e s e a s on . B i r th w e igh t i s1 0 -25 g . D ec iduou s d en t i t ion r e p lace m en t b eg in s b y 2 4 d ay s a nd w ean ing o ccu r s a t4 -6 w eek s , w i th d i spe r sa l h appen ing t h e r eaf t e r . Y oung d o n o t b r eed i n t h e s e a son o ft h e i r b i r th . C o rbe t a nd H a r r i s ( 1 99 1 :42 ) a l so r e po r tt h a td en s i t ie sa r ea bou t 1 / ha ,w h i le n o t ing t h a tn o c on c l u s ive d a ta e x i s t s . T he a u tho r h a s n o t ed g r ea te r d en s i t ie s t h an t h i s i nn o r the rn F rance , w he r e m any i n d iv idua l s m ay g a the r a r ound a h uman e ncampmen t t o s c avenge f ood . L i f e span i nt h e w i ld d oe s n o t u sua l ly e xceed 7y ea r s , w i th 6 0 -70% o fm o r ta l i ty o ccu r r ing d u r ing t h e f i r s t y ea r o f l i f e ( Co rbe t a nd H a r r i s 1 99 1 :43 ) . D ue t ot h e i r s i z e a nd p ro tec t i ve s p ine s , E . e u ropaeu s i sv e ry r a r e t awny o w l p r ey a nd e ven r a r e r b a rn o w l p r ey . D e sp i t e t h e i r p r o t ec t ive s p ine s , t h ey a r eo f t en t a ken b ye ag le o w l s .

i t s b u r row s y s t em , i t s a cqu i s i t ion b y h uman s mu s t o ccu r d u r ing t h ew a rme rp a r to ft h ey ea r . B r eed ing o ccu r s s ho r t ly a f t e rt h e a du l t f ema le s e me rge f r om h i ber na t ion i n Ma r ch . G e s ta t ion l a s t s f o r 3 0 d ay s , a nd w ean ing o ccu r s a f t e r a n a dd i t iona l 3 0 d ay s . T he j uven i le s e me rge f r om t h e i rn a ta l b u r row s i n May . Mean l i t t e r s i z e i s a bou t 2 . 4 , b u t t h i s s pec ie s r e p roduce s e ve ry y ea r , u n l ike s ome s p ec ie s o f Ma rmo ta t h a t r e p roduce b i enn ia l ly ( Ba ra sh 1 989 ) . L AGOMORPHA , F AM ILY L EPOR IDAE T he t h r ee s p ec ie s d i scu s sed b e low a l h av e ad en ta l f o rmu la f o l low ing 2 . 0 .3 .3 / 1 .0 .2 .3 . T he se d en ta l r ema in s a r e d i s t in g u i shab le o n s l igh t mo rpho log ica l g r ound s , e spec ia l ly w i th t h e 2P2,, P , a nd 3M 3 ( H i l l son 1 986 :80 ) . L epu s i sa l so s i gn if i c an t ly l a rge rt h an O ryc to lagu s . O r yc to lagu sc un icu lu s( r abb i t ) ( F igu r e3 . 26 ) T he o r ig ina lr a nge o ft h i ss p ec ie s i st h ough tt o h ave b een c onf ined t o N o r thwe s t A f r ica a nd t h e I b e r ian p en in su la ( Co rbe t 1 978 ) . H uman h ab i ta t a l te r a t ion a nd d e l i b e ra t e i n t roduc t ion s / e scape s f r om f a rm s h a s l e d t o h uge r a nge e xpan s ion t ot h e B r i t i sh I s le s , F ran ce , G e rmany , I t a ly , s ome Med i t e r ranean i s land s , a nd n o tab ly t o A u s t ra l ia a nd N ew Z ea land . T h i s r a nge e xpan s ion i n c lude s n o r ther n S pa in , w he r et h i s s p ec ie sh ad n o t b een p r ev iou s ly n ume rou s . P r ef e r r ed h ab i ta t s a r e n a tu ra l o r a r t if i c ia l a r ea s o f s ho r t g r a s se s p rov id ing g ood c ove r o n o r i n c l o se p rox im i ty t o b u r rowab le t o p so i l( Co rbe ta nd H a r r i s 1 99 1 :149 150 ) .

F UR -BEAR ING S PEC IES T he se s p ec ie sa r e g r ouped s e pa ra t e ly n o to n ly f o rt a xonom ic d i f f e r ence s , b u t a l so f o r d i f f e r ence s i nt h e i r p r obab le mode o fp r ocu r emen t a nd p r e sumed d epo s i t ion w i th in C an tab r ian U ppe rP a leo l i th ic s i te s : t h rough h uman h un t ing . T h i s a cqu i s i t ion m ay r e p r e sen t f u r t r app ing b y h uman s ( Chap t e r 9 ) . T he m a jo r i ty o ft h e se s p ec ie s a r e t o o l a rge t o b e c ommon b ar n o w l p r ey i nm o s t i n s tance s , t h ough e ag le o w l s c om m on ly f e ed u pon t h e s ma l le r t a xa o rj uven i le s o ft h e l a rge r . A n e xcep t ion i st h e w ea se l ( M u s t e la n i va l i s ) w h ich i ss ma l l e nough t ob ep r eyed u pon b yb a rn o w l s , y e ts t i l l m ay o f f e ra u sab le p e l tt oh uman t r appe r s f o rt h e l a rge r m embe r s o ft h i s s p ec ie s . S ome o ft h e se s pec ie s m ay a l so h ave b een a cqu i r ed f o r t h e i r m ea t , t h ough a mong t h e c a rn ivo r e s t h i s i s al e s s d e s i r ed r e sou r ce f o r h uman s ( d ue t o t h e g ene ra l ly p oo r e r t a s t ing m ea t ) . T he se t a xa g ene ra l ly o ccu r i nl ow n umbe r s a mong C an tab r ian U ppe rP a leo l i th ic s i t e s .

B r eed ing b eg in s a s e a r ly a s J anua ry a nd e x t end s t h rough A ugu s t , t h ough t h e o n se t o ft h e b r eed ing s e a son c an b e d e l a yed b y s e ve r e w ea the r . L i t te r s i z e s o f 3 7 a r e t y p ica l . G e s ta t ion l a s t sa bou t3 0 d ay s , a f t e rw h ich t h ef ema le i so f t en p r egnan t a ga in w i th in o ne d ay . We igh t a tb i r th i s3 0 -35 g , w h ich i n c r ea se st o 1 50 -200 ga tw ean ing ( c a . 2 1 d ay s ) , t h en 1 200 -2000 ga ta du l thood ( Co rbe ta nd H a r r i s 1 99 1 :149 152 ) .

F AM ILY S C IUR IDAE Ma rmo ta m armo ta ( a lp ine m a rmo t ) T h i s l a rg e r oden t s p ec ie s i sf ound t o day i nt h e a l p ine r e g ion o f E u rope ( F igu r e 3 . 25 ) a nd h a s b een r e in t roduced t o t h e P y r enee sa nd t h eB a lkan s( He r r e ro e ta l . 1 992 ) . T he m a s so f t h i s s p ec ie s i se x t r eme ly v a r iab le t h roughou tt h e y ea r d ue t o s e a sona l f a t s t o rage f o r h i ber na t ion . M m a rmo ta r e a che s 3 00 -400 g b y s i x w eek s o fa ge , o v e r 2 000 gb y 2y ea r s o f a ge a nd o ve r 3 000 g b y 3y ea r s o f a ge w hen f u l l s i z e i s r e a ched . W i th s e a sona lf a tg a in ,a du l tm a s sc an r e a ch 4 500 g i na u tumn ( N ie thamme ra nd K rapp 1 978 :174 ) .

L epu sc apen s i s( b rown h a r e ;c ommon h a r e ) ( F igu r e 3 . 27 ) L . c apen s i s i sf ound i n t h e t empe ra t e a nd Med i t e r ranean z one o ft h eP a lea r c t ic , i n to t h eA rab ian p en in s u la a nd a l o fA f r ica . T h i s w ide rang ing s p ec ie s i sd i v ided b ys ome a u tho r s i n to L .c apen s i s f o rt h e s ou the r ly p o r t ion o f t h i s r a nge a nd L . e u ropaeu s f o r t h e n o r the r ly p o r t ion , i n c l ud ing E u rope . T h i ss e pa r a t ion i sp r ima r i ly b a sed u pon s i z e ( Co rbe t 1 978 :72 ) , v a r ia t ion o fw h ich a c ro s s s u ch al a rge a nd v a r ied r a ng e i st y p ica l w i th in m any s p ec ie s . T he a u tho r t h e r ef o r e r e ta in s f o r t h e p r e sen t t h e m ono spec if i c d e s igna t i on , a l though t h i s i sn o t a lway s t h e c a se f o r E u ropean P a l e o l i th ic r e sea r che r s .

Ma rmo t s i n hab i tt h e a l p ine z one g ene ra l ly a bove t h et r ee l ine i no pen p a s tu r e s . A l t i tude s f r om 1 800 t o2 400 m a r e mo s t t y p ica l , b u tt h i s s p ec ie s c an b e f o und d own t o 1 450 m i nt h e P y r enee s ( He r r e ro e ta l . 1 992 ) . S o i l d ep th s u ff ic ien t f o rt h e c on s t ruc t ion o f t h e i r e x t en s ive b u r row s y s t em s i sa no the r r e qu i r emen t ( Co rbe t a nd O venden 1 982 :148 ) . L a rge f am i ly g r oup s o ccupy t h e se s y s tem s . G roup c oope ra t ion f o rp r eda t o r a vo idance m ake s t h i s s p ec ie s d i f f i cu l t t oh un t , b u t t h ey m ay b e t r apped o r s na r ed b y d ev ice s p l a ced n ea r t h e i r b u r r ow e n t rance s . S i nce t h i s s p ec ie s h i b er na te s e a ch w in te r i n

C o rbe t a nd H a r r i s ( 1991 :155 157 ) r e po r tm ean a du l tm a s se s o f3 . 23 k g f o rm a le s ( n —7 23 ) a nd 3 . 43 k g f o r f ema le s ( n= 6 89 ) f o r ap opu la t ion f r om E a s t A ng l ia ( Eng land ) . T he i r s i z em ake st h em u n l ike ly p r ey e xcep ta sj uven i le sf o rs ma l le r s p ec ie s o fo w l ( b a rn , t awny o r l o ng -ea r ed ) , s i nce w e igh t a t b i r th i sn o rma l ly o ve r 1 00 g . T hey a r e c ommon p r ey o ft h e m uch l a rge r e ag le o w l a nd p a r t icu la r ly t h a t o f r e d f o xe s . B r eed ing c on t inue s f r om F eb rua ry t h rough O c tobe r a nd e x p and si nm i ld c l ima t e , w i th l i t te rs i z e so f 3t h e mo s tc ommon

4 0

( r ange : 1 4 ) . G e s ta t ion l a s t s f o r 4 1 -42 d ay s , w i th w ean ing o ccu r r ing a f t e r2 3 d ay s .

n ea r ly a l o ft h eH o la r c t ic r eg ion a nd a l so p a r t s o fN o r th A f r i ca . T h i sr a nge m ay h ave b een a l te r ed s omewha tb yh uman a c t iv i ty a nd f o r e xamp le s p r ead b y R oman s t o E gyp t , b u t D ayan a nd T che rnov ( 1988 ) s ugge s t t h a t t h i s E gyp t ian p opu la t ion i s aP l e i s tocene r e l ic t . S i z e a c ro s s t h i s r a nge i s v a r iab le , w i th a v e ry h i gh d eg r ee o f s exua l d imo rph i sm . P opu la t ion s t h roughou t B r i ta in ( s umma r iz ed b y C o rbe t a nd H a r r i s 1 99 1 :39 1) h ad m ean s i ze sr a ng ing f r om 1 06 131 gf o r m a le sa nd 5-63 gf o rf ema le s . S heff i e ld a nd K ing ( 1994 :2 ) s umma r iz em ean a du l tm a le b ody m a s se s , r a ng ing : 3 0 100 g i nN o r th Ame r ica , 5 -202 gi nt h e U n i ted K ingdom , 3108 gi nS weden , 4 0 -220 g i n( t he f o rme r ) C z echo s lovak ia , 3 6 2 50 g i n( t he f o rme r ) U .S .S .R . , 7 2 185 g i n( t he i n t roduced p opu la t ion o f ) N ew Z ea land , a nd 1 25 -250 g i nA f r ica . I n d i r ec t o ppo s i t ion t oB e rgmann 's r u le , s i z e t e nd s t oi n c rea se s i gn if i can t ly t owa rd s t h e s ou the rn e nd o ft h i s s p ec ie s ' r a nge ( Co rbe t a nd H a r r i s 1 991 :388 ) . R educ t ion s i ns i z e i ns ome a r ea s m ay a l so r e f lec ts ympa t ry w i th M . e rm inea a nd M . .f r e n a ta : w he r e t h e se t wo s p ec ie s o ve r lap w i th i t , M . n i va l i s t e nd s t ob e s ma l le r . I n N o r th Ame r ica , t h i s r e la t ion sh ip i s l e s s c l ea r , a nd s i z e v a r ia t ion d oe s n o t s e em t ob e r e la ted t o s ympa t ry o rB e rgmann 's r u le ( Sheff i e ld a nd K ing 1 994 :2 ) .

L .c apen s i si so r ig ina l ly as p ec ie s o fo p en g r a s s land s , t h ough a si st y p i ca l ly t h e c a se t h ey h ave a dap t ed t oa nd s p read t h e i r o r ig ina l r a nge w i th t h e e xpan s ion o f a g r icu l tu re a nd t h e o pen ing o f f o rme r ly f o r e s t ed h ab i ta t s . T h i s s pec ie s m ay l e ave v i s ib le t r a i l s o n o pen m eadow s a nd c ha ra c te r i s t ic g ap s i nd en se r v ege ta t ion , w h ich m ake s t h e p l a c ing o fs na r e s o r o t he r t r ap t y pe s ap r ac t ica l m e thod o f h a rve s t b y h uman s . T ra ck ing t h rough s now i sa l so p o s s ib l e . Wh i le w in te rp e lage d oe s n o t t u rn w h i t e i nt h i s s p ec ie s , t h e i r f u r i ss t i l l p r iz ed . T hey a r e a l so h un t ed f o r t h e i r m ea t , t h ough mo s t mode rn h un t ing i sf o rp u rpo se so fa g r icu l tu ra lp e s tc on t ro l . L epu st i m idu s( a r c t ic h a r e ; moun ta in h a r e ) ( F igu r e 3 . 28 ) L .t i m id u s i n hab i t s t h e t u nd ra a nd c on if e rou s f o re s tz one o ft h eH o la r c t ic , f r om t h en or the rn B r i t i sh i s land s t oG r e en land a nd L ab rado r . A n i s o la t ed p opu la t ion a l so e x i s t s i na nd a r ound S w i tz e r land a nd m ay b e aP l e i s tocene r e l i c t . T h i s s p ec ie s i sr e p la ced e co log i ca l ly t ot h e s ou th o fi t s r a nge b y L .c apen s i s i nE u rope . L .t i m idu s t e nd s t ob e t h e s ma l le r o ft h e t wo . A du l t s a v e raged 2 . 8 k g ( n =4 5 ) i na S co t t i sh p opu la t ion a nd 3 . 2 k g ( n =2 7 ) i n a n I r i sh o ne ( Co rb e ta nd H a r r i s 1 99 1 :164 ) . T h i s s p ec ie s s e em s t of o l low B e rgmann 's r u le i nE u rope , a sn o r th e rn i n d iv idua l s t e nd t o b e l a rge r t h an s ou ther n i n d iv idua l s ( Ange rb jö rn a nd F l ux 1 995 ) . A s w i th L .c apen s i s ,j uven i le s c ou ld b e t a ken o cca s i ona l ly b y s ma l le r o w l s p ec ie s , w i th a du l t p r eda t ion f a l l ing m o s t ly t ol a rge rp r eda to r sl i ke r e df o x .

H ab i ta t p r ef e r ence s a r e e x t r eme ly v a r ied a nd h ave i nc om mon a d equa t e c ove r t o p ro tec t t h i s s p ec ie s f r om t h e m any t h a t p r ey u pon i t( mo s t l a rge r c ar n ivo re s , o w l s , h awk s , h u m an s , e t c . ) . H en ce , M . n i va l i s m ay b e f o und w he re i t s ( t yp ica l ly s ma l l r o den t ) p r ey m ay b e f ound : o pen f o re s t s , a g r icu l t u ra l f i e ld s , m eadow s , h edge row s , a l p ine m eadow s a nd w ood s , s c rub land s , s t eppe a nd s em i -de se r t , p r a i r ie s , a nd c oa s ta l d une s . D en se f o r e s t a nd t r ue s a ndy d e se r t s a r e a vo ided ( Sheff i e ld a nd K ing 1 994 :5 ) . Wa t e r v o le s a r e af r e q uen t c omponen t o ft h e d i e t w he r e t h e r a nge s o ft h e se t wo s p ec ie so ve r lap i nt h eP a lea r c t ic ( De la t t re 1 983 ;J ed rze jew sk i e t a l . 1 995 ) . S ma l l p r ey i sf r equen t ly c a ched ( Ok sanen 1 983 ) . M . n i va l i si sa c t ive b o th d ay a nd n i gh t , a nd i st r apped mo r e f r equen t ly d u r ing t h e d ay ( Sheff i e ld a nd K ing 1 994 :6 ) . T h i s b road a c t iv i ty p a t ter n c omb ined w i th s ma l l s i ze m ake s t h i s s p e c ie s v u lne rab le t o mo s t l a rge r p r eda to r s , i n c lud ing d i ur na l a nd n o c tur na l r a p to r s , a nd m o s t l a rge r c a rn ivo r e s . T h i s s p ec ie s i sr a r e p r ey ( we l l b e low 1 % o fd i e t ) o fs ma l le r s p ec ie s o fo w l , s u ch a sb a rn o rt awny ( G lue 1 974 ; S ou the rn 1 954 ) .

T h i s s p ec ie s i sa s soc ia ted w i th o pen h ab i ta t s , p a r t icu la r ly h ea th e r m oo r land , a nd h a s n o t s u rp r i s ing ly c o lon ized f a rm l a nd a s w e l l . H ighe s t d en s i t ie s a r e r e ached i n m ixed o pen / f o r e s t h ab i ta t s a l ong h edg e row s a nd c l ea r ing s ( Ange rb jö rn a nd F l ux 1 995 :5 ) . Mo s t a c t iv i ty i sn oc tur na l a nd i ng r oup s w h ich m ea su r ing u p t o1 00 i n d iv idua l s . S im p l e , s ho r t b u r row s 1 2 m l o ng a r e e xcava ted f o r d ay t ime s h e l te r , t h ough l e s s p e rmanen t s u r f a ce s c rape s a r e a l so c om m on . B u r row s c ou ld h ave d r awn t h e t r app ing a t ten t ion o f P a leo l i th ic h un te r s . T he p e lage o f L . t i m idu s t u rn s f r om d u sky b r own t o mo s t ly w h i t e i nw in t e r , p e rhap s i n c r ea s ing i t sv a lue ( i . e . s upp ly ing n a tu ra lc amouf lage t oh uman h un t e r s a ga in s t as now b a ckd rop ) .

O ne o rt wo l i t t e r s a r e p roduced e a ch y ea r , w i thou t d e layed i mp lan ta t ion o fe mb ryo s ( Co rbe t a nd H a r r i s 1 99 1 :393 -394 ) . P r egnancy t y p i ca l ly o ccu r s d u r ing l a te F eb rua ry t h rough J u ly , w i th b i r th s b eg inn ing i nA p r i l . G e s ta t ion l a s t s 3 4 -37 d ay s . L i t t e r s i z e v a r ie s ( Sheff i e ld a nd K ing 1 994 :3 -4 ) , b u t a n a ve rage o f a bou t 5i st y p ica l f o r m any p o r t ion s o f t h i s s p ec ie s 'r a nge . N ewbor n sw e igh l e s st h an 2g . T he i rp e rma n en t d en t i t ion i sc omp le te ly i np l a ce b y d ay 4 2 , w i th f u l l a du l t b ody m a s s b e ing o b ta ined b y w eek s 1 2 15 . L if e e x p ec tancy i sl i t t le o ve r ay ea r i nt h e w i ld , t h ough u p t ot h r ee y ea r s i sp o s s ib le . T he i rp e lage t ur n sw h i te i nw in t e r i na r ea s o fs i gn if i can t s now c ove r , w h ich m ay i n c r ea se t h e i r v a lue t o h uman t r appe r s .

B reed ing c on t inue s f r om J a nua ry t h rough O c tobe r , w i th t h r ee l i t t e r s p e r s e a son p e r a du l t f ema le t y p ica l . L i t te r s i z e r a nge s f r om 1 8 , w i th m ean s f o r f i r s t l i t t e r s ( 2 .15 ) s ma l le r t h an t h o se f o rt h es e cond l i t t e r s( 3 .24 )i n as e a son . G e s ta t ion a ve rage s a bou t 5 0 d ay s , w i th w ean ing o ccu r r ing a f t e r 4 w eek s( Ange rb jö rn a nd F l ux 1 995 :4 -5 ) . CARN IVORA ,F AM ILY MUSTEL IDAE T h i s c a rn ivo ran f am i ly i sa bundan t a nd d i ve r se i nE u rope , w i th u p t o 8s p ec ie s f o und n a tu ra l ly t o day i n t h e I b e r ian p en in su la ( Co rbe t a nd O venden 1 9 82 ; S an to s R e i s 1 983 ) . Among Mu s t e la a nd G u lo , d en ta l f o rmu la i s3 . 1 .3 .1/ 3 . 1 .3 .2 , w h i le Ma r te s ,M e /e sa nd L u t ra h ave a dd i t iona lt e e th .

Mu s t e la e rm inea ( e rm ine , s t oa t ) ( F igu r e 3 . 30 ) M . e rm inea a l so h a s aH o la rc t ic d i s t r ibu t ion , t h ough p r ima r i ly i nt h e t u nd ra , b o r ea l , a nd d ec iduou s z one s ( Co rbe t 1 978 ) . S exua l d imo rph i sm i sl e s s p ronounced t h an

Mu s t e la n i va l i s ( wea se l , l e a s tw ea se l ) ( F igu re 3 . 29 ) T h i s s ma l le s tm embe ro ft h e mu s te l id s ( a nd o f t h eo rde rC a rn ivo ra )h a s aw ide sp r ead d i s t r ibu t ion , i n c lud ing 4 1

i nM . n i va l i s , w i th m a le s a bou t 1 . 5 t ime s t h e s i ze o ff ema le s . Adu l t ma le m ean w e igh t ( s umma r ized b y C o rbe t a nd H a r r i s

c onf u sed b y t h e p a s t i n t roduc t ion o f Ame r ican m ink (M . v i son ) t oE u rope i no rde r t or ep len ish l o ca l s upp l ie s o ft r ap p ab le a n ima l s . F ur the rc onf u s ion r e su l t s f r om t h e i n te rf er t i l -

1 991 :378 ) f o rE u ropean c oun t r ie s r a nge s f r om 2 08 -283 g ,i n R u ss ia 1 34 -191 g ,i nN o r th Ame r ica 5 6 -206 g ,a nd i nB r i ta in

i t y ( i n c ap t iv i ty ) o f t h e se t wo s pec ie s ( Co rbe t a nd H a r r i s 1 991 :408 ) a nd t h e i r v ery s im i la r a ppea ran ce . T h i s r a nge

c a . 3 20 g . T h i sd i s t r ibu t ion d oe sn o tf o l low B e rgmann 's r u le a nd may r e f lec t v a r ia t ion i nl o ca l p rey s i ze ( E r l inge 1 987 ;

m ay a l so h ave e xpanded i n to s ome a r ea s , a s i n to F rance w i th in t h e p a s t f ew c en tu r ie s a nd i n to C an tab r ian S pa in a s

K ing 1 991) . T he h ab i ta t r e qu i remen t s o f t h i s s pec ie s a r e s im i la r t o t ho se o f M . n i va l i s : mo s t t h a t o f f e r s uff ic ien t c ove r , e i the rf o ri t se lf o rf o ri t sp rey t oe x i s t( S leeman 1 993) .

r e cen t ly a s 1 951 ( Youngman 1 982 :12 -13 ) .

A s amp le o f 7

m a le s f r om F rance a ve raged 8 71 g( r ange : 6 50 -1005 g ) , w i th

F a rm land , f o r e s t s , w e t land s , a nd a l p ine z one s a r e a l l o ccu p i ed , b u t ma tu re f o re s t s l a ck ing i nc ove r a nd s ma l l mamma l p rey a r e a vo ided ( Co rbe ta nd H a r r i s 1 99 1 :379 ) . D ep reda t ion

t wo f ema le s w e igh ing 5 05 ga nd 5 80 g( Youngman 1 990 :1) . T he p r ef e rred h ab i ta t s o ft h i s s pec ie s a r e m a r shy a r ea s a nd t h e b o rde r s o f r i ve r s a nd l a ke s ( Co rbe t a nd O venden 1 982 :182 ; Y oungman 1 990 :2 ) . T h i sp ref e r ence i si nk eep ing w i th i t ss em i -aqua t ic l i f e s ty le a nd d i e t .

u pon p ou l t ry r a i s ing o pe ra t ion sa r ea l so c ommon ,b u tn ot r ue c ommen sa l r e la t ion sh ip w i th h uman s e x i s t s . T h i s s pec ie s a l so c ache s s ma l l p r ey i nl a rge n umbe r s ( Ok sanen 1 983 ) , a b ehav io r w h ich may l e ave p a t te rn s d e tec tab le i nt h e f o s s i l r e co rd .

Ma t ing o ccu r s i n t h e e a r ly s p r ing ( Youngman 1 990 :1-2 ) , w i th a l i t te r o f2 -7 b or n 4 0 -43 d ay s l a te r . I mp lan ta t ion o f

Ma t ing i nt h i s s pec ie s o ccu r s i ns umme r , b u t i mp lan ta t ion o f t h e e mb ryo s i sd e layed u n t i l t h e f o l low ing s p r ing w hen a s i ng le l i t te ro f6 12 i sp roduced . A c t ive p r egnancy t h e ref o re l a s t s a bou t 4w eek s . T he y oung a r e w eaned a t7 12 w eek s ( Co rbe ta nd H a r r i s 1 99 1 :384 -385 ) .

e mb ryo s i ss l igh t ly d e layed , s o a c t ive p r egnancy l a s t s o n ly a bou t 2 8 d ay s ( Co rbe t a nd H a r r i s 1 99 1 :4 13 ) . S i ze a tb i r th r un s a bou t 8g . F u l l a du l t d en t i t ion i si n p l ace b y 6 0 -67 d ay s , a nd t h e y oung b eg in t h e i r b reakup f r om t h e l i t te r a nd d i spe r sa l b y a ge 4 mon th s . Ma le s a nd f ema le s c an b r eed i n t h ey ea rf o l low ing t h e i rb i r th .

T h i s s pec ie s i sg ene ra l ly t o o l a rge t of a l l p r ey e xcep t a sj u v en i le s t os ma l le r s pec ie s o fo w l , t hough p reda t ion b y e ag le o w ls i sn o t u ncommon . W in te r p e lage o f M . e rm inea i s mos t ly w h i te , w h ich h i s to r ica l ly h a s i n c r ea sed i t s v a lue t o h uman h un te r s .

T he p e l t s o f t h i s s pec ie s a r e s t i l l c ove ted f o r t h e i r b eau ty , w a rm th , a nd w a te r r epe l lence , t hough mos t t oday a r e o b t a ined f r om c omme rc ia lm ink f a rms . T rap s o rs na r e sm ay b e p l aced a tl ow p o in t s a l ong r i ve rb ank s w he re t h e se a r e l i ke ly t op a s s .

Mus te la p u tor ius ( Eu ropean p o leca t )

Man te s man te s( p ine ma r ten ) ( F igu re 3 . 33 ) T h i s s pec ie s i s f ound t h roughou t mo s t o f E u rope a nd i n to E u ropean R u s s ia . No r ther n S pa in i si n c l uded i n t h i s r a nge , a s a r e S cand inav ia a nd t h e c en t ra l Med i te rranean r e g ion . A s w i th mo s t l a rge p reda to r s , i t s r a nge m ay h ave b een s i gn if ican t ly r educed f r om s ome a r ea s o fE u rope . S i nce i ti s av a lued f u r -bea r ing a n ima l , i t sr a nge m ay h ave b een a r t if ic ia l ly o ra cc iden ta l ly e x tended i n to o t he r a r ea s . T he g enu s Ma r te s h a s mo re t e e th t h an o t he r m u s te l i d s , w i th a f o rmu la o f 3 . 1 .4 .1/3 .1 .4 .2 . C o rbe t a nd H a r r i s ( 199 1 :373 ) r e por t b o th s i gn if ican t s exua l d imo rph i sm a nd v a r ia t ion b e tween p opu la t ion s . I nI r e land , m a le s r a nge f r om 1 500 -1950 g( n =3 1) , w i th f ema le s r ang ing 110 -1450 g( n =3 8 ) . Wh i le p opu la t ion s i n S co t land a r e s im i la r i n s i ze , s pec imen s f r om ap opu la t ion i nt h e ( f o rme r ) U .S .S .R . h ave a m a le r a nge o f6 70 -1050 g ( n =5 9 ) a nd a f ema le r a nge o f 4 84 -850 g ( n =4 9 ) . H ab i ta t p ref e rence i sc on if e rou s o r

( F igu re 3 . 31) T he r a nge o ft h i s s pec ie s c ove r s a l l o ft empe r a t e a nd mo s t o f Med i te r ranean E u rope , e xc lud ing G reece . T h is r ange , h oweve r , m ay h ave b een a l te red d ue t o l o ca l e x t i rpa t ion b y h uman s i n s ome a r ea s a nd e xpan s ion i n to o the r a r ea s w he re t h e l a nd scape h a s b een a l te red . B ody s i ze i nB r i ta in ( Co rbe t a nd H a r r i s 1 991 :399 ) a ve rage s 111 gf o r ma le s( n =2 8 ,r a nge : 8 00 17 10 g )a nd 6 89 gf o rf ema le s ( n= 1 4 , r ange : 5 30 -9 15 g ) . M .p u to r iu s i st h ep robab le w i ld p ro g en i to r o fd ome s t ica ted f e r re t s (M .f uro ) . O ccup ied h ab i ta t s a r e v a r ied , a ga in w i th t ho se o f f e r ing s uff ic ien t c ove r b e ing f avo red . T he se i n c lude f o re s t s , h edge row s , a g r icu l tu ra l l a nd s , w e t land s , a nd w a te r m a rg in s . U ppe r a l t i tude s t e nd t o b e mo re t h in ly p opu la ted ( Co rbe ta nd H a r r i s 1 991 :401) . B o th ma le s a nd f ema le s b ecome s exua l ly ma tu re a nd b r eed f o r t h e f i r s t t ime i nt h e y ea r f o l low ing t h e i r b i r th . Ma t ing t yp ica l ly t a ke s p l ace i nA p r i l , a nd a l i t te r o f u sua l ly 5 10

m ixed f o re s t , b u t t h i s s pec ie s may a l so b e f o und a tt ime s i n

y oung i sp roduced a f t e r ag e s ta t ion p e r iod o f4 2 d ay s . Mo s t

d ec iduou s f o r e s t o r mo r e o pen v ege ta t ion ( Co rbe t a nd 0 ve nden 1 982 :184 ) . I t may t h u s b e f ound u p t ot h e t r ee l i ne ,

l i t te r s a r e t hu s p roduced i n May o r J une , w i th n ewbor n s w e igh ing 9 10 g . P e rmanen t d en t i t ion i sp l ace b y 1 13

w h ich i nt h e P y renee s a nd A lp s i sa pp rox ima te ly 2 , 000 m .

w eek s , a nd j uven i le s r e ach a du l t d imen s ion s b y t h e a u tumn

M . m a r te s i sa r bo rea l a nd h igh ly a g i le .

f o l low ing t h e i r b i r th . T he y oung a r e f u l ly i n dependen t b y l a te s umme r a nd d i spe r se i nt h e e a r ly a u tumn ( Co rbe t a nd H a r r i s 1 991 :403 -404 ) .

a r e n o t a vo id ed a nd s ome s c aveng ing o r l i ve s tock r a id ing

H uman h ab i ta t ion s

m ay o ccu r . Mo s t a c t iv i ty i ss o l i ta ry a nd n oc tur na l , w i th d ay t ime s s p en t

Mus te la l u t reo la ( Eu ropean m ink )

i n ad en o f v a r iou s c onf igu ra t ion s ( r ockf a l l , t r ee c r ev ice ,

( F igu re 3 . 32 ) T h i ss pec ie sm ay b e f ound c u r ren t ly i nE a s te rn

h uman c on s t r uc t ion s , e t c . ) . D ie t i s mo s t ly c a rn ivo rou s u pon s ma l l mamma l sa nd b i rd s ,b u ta l so i n c lude si n ver teb ra te sa nd

E u rope a nd R u s s ia , a nd i ns ou thwe s ter n F rance . D ue t ot h e c on t inu ing v a lue o fm ink p e l t s , t h i s r ange i ns ome p opu la ted a r ea s mu s t b e as i gn if i can t c on t rac t ion f r om i t s f o rme r s t a te ( Youngman 1 982 :3) .

b e r r ie s ( Go sz czyn sk i 1 976 ) . D e layed e mb ryo i mp lan ta t ion i n c rea se st h e g e s ta t ion t ime b e tween t h e s umme r ma t ing s e a s on a nd b i r th t h e f o l low ing Ma rch t o A p r i l . A c t ive p r eg -

T he r a nge o f t h i s s pec ie s i sf u r the r 4 2

a bandoned b u r row s . T rapp ing a ts u ch j unc tu r e s i st h e m o s t v i ab le m e thod o f c ap tu r e b y h uman s , s i nce t h i s s p ec ie s i s s o l i ta ry ,n oc tur na la nd v e ry e l u s ive ( San to sR e i s 1 983 ) .

n ancy t h e r ef o r e l a s t s o n ly a bou t 3 0 d ay s . L i t te r s i ze i sf r om 1 5 , w i th a n a ve rage s i z e o f 3 0 ge a ch a tb i r th . Wean ing o ccu r s a t5 6 1 2 w / eek s . A du l tw e igh ti sr e a ched b yt h e s um m e rf o l low ing b i r th ( Co rbe ta nd H a r r i s 1 99 1 :374 ) .

Me le s me l e s( b adg e r ) ( F igu r e 3 . 36 ) M . m e / e s i sf ound f r om P o r tuga l t oJ apan a nd C h ina , i n c lud ing t h e B r i t i sh I s le s , S cand inav ia , A s ia M ino r a nd t h e Med i t e r ranean r e g ion i ni t sr a nge . I t sd en ta lf o rmu la i s3 . 1 .3 -4 .1 / 3 .1 .3 -4 .1 , w i th m inu t e v e s t ig ia l f i r s t p r emo la r s s ome t ime s o ccu r r ing , mo r e o f t en m and ibu la r ly ( Co rbe t a nd H a r r i s 1 99 1 :4 15 ) . Mean m a s s f o r i n d iv idua l s a tl e a s t o ne y ea ro l d f r om s ou thwe s tE ng land ( ma le s ) i s1 .6 k g ( n =3 , r a nge : 9 . 116 .7 k g ) a nd ( f ema le s ) 1 0 .1 k g ( n =8 4 , r a nge : 6 . 6 13 .9 k g )( Co rbe ta nd H a r r i s 1 99 1 :419 ) .

Ma r te sf o ina ( b eech m a r t en , s t one m a r t en ) ( F igu r e 3 . 34 ) C u r ren t r a nge o ft h i s s p ec ie s i sp r ima r i ly i n w e s t e rn a nd s ou thea s t e rn E u rope , i n c lud ing n o r the rn S pa in a nd t h e c en t ra l Med i te r ranean r e g ion , b u t e xc lud ing S cand i n av ia . I s o la t ed p opu la t ion s a r e a l so f o und i nC en t ra l A s ia . A sw i th M . m a r t e s , t h i s r a nge m ay h av eb een a l te red b y h u m an a c t ion s . M .f o ina i n hab i t s d ec iduou s f o r e s t s w i th in t h i s z one , a nd m ay t h e ref o r eb ef ound t ot h el i m i t so ft h et r ee l ine . H uman h ab i ta t ion s a r e n o t a vo id ed a nd f ood i so f t en s c av e ng ed f r om t h e se s e t t ing s ( Co rbe t a nd O venden 1 982 :185 ; D e l ibe s 1 983 ) . T he i r l i f e s ty le i sm a in ly a r bo r ea l . T he i rd i e t i se c l ec t i c a nd o ppor tun i s t ic : s ma l l m amma l s , b i rd s , e gg s , r e p t i le s , a mph ib ian s , f i sh , i n ve r teb ra te s ,v ege tab le m a t te ra nd c a r r ion a r e a l c ommon ly c on sumed ( Amo re s 1 980 ; G o sz c z yn sk i 1 976 ) . T he c omb ina t ion o fa s soc ia t ion w i th t h e b i o t ope s a r ound h uman d we l l ing s a nd t h e i r a t t rac tab i l i ty t oa w ide v a r ie ty o fp o s s ib le b a i t s w ou ld h ave m ade t h i s s p ec ie s a na t t r ac t ive f u rt r app ing t a rge t .

M . m e / e s p r ef e r s d ec iduou s f o r e s t a nd o pen p a s tu re s , a nd p r ef e ren t ia l ly h ab i ta t s w h ich o f f e r am ix tu r e o f t h e se t wo v ege ta t ion s . T h i s s p ec ie s i s mo r e g r ega r iou s t h an o t he r mu s te l id s , l i v ing i n l a rge b u r row s w i th mu l t ip le e n t rance s o ccup ied f o rm u l t ip le g ene ra t ion s . A n a ve rage g roup s i z e o f 6i su sua l ly p r e s ided o ve r b y ad om inan t m a le ( Co rbe t a nd H a r r i s 1 99 1 :4 19 ) . A dap ta t ion s f o r as i gn if ican t ly f o s so r ia l l i f e s ty le i n c lude e y e s igh t b e s t i np oo r l i gh t a nd f r on t l i mb s a nd c l aw s a dap t ed f o r t u nne l ing . T he d i e t i so ppo r tun i s t ic a nd o mn ivo rou s , w i th e a r thwo rm s a s ap r ima ry f ood s ou r ce a nd i n c lud ing o t he r i n ve r teb ra t e s , s ma l l m amma l s , f r u i t s , t u be r s ,b i rd s , f u ng ia nd c a r r ion . A c t iv i ty i sp r ima r i ly n oc tu r n a la nd c r epu scu la r .

M f o ina r e a ch e s r e p roduc t ive m a tu r i t y i nt h e s e cond y ea ro f l i f e . Ma t ing o ccu r s b e tween J u ly a nd m id -Augu s t , w i th d e l a y o f e mb ryo i mp lan ta t ion u n t i l t h e f o l low ing J a nu a r y /Februa ry . B i r th t a ke s p l a ce i n Ma r ch /Ap r i l a f t e r a n a c t i ve p r egnan cy o fj u s t o ve r o ne m on th . L i t te r s i z e r a nge s f r om 1 5 , w i th a n a ve rage o fa r ound 3 . Wean ing o ccu r s n o l a t e rt h an J u ly ( Mad sen a nd R a smu s sen 1 985 ) .

L i t te r s o f 1 5 a r e b o rn a r ound F eb rua ry , t h ough m a t ing i s d u r ing t h e p r ev iou s s p r ing a nd d e layed i mp lan ta t ion o fe m b ryo s o ccu r s . A c t ive p r egnancy l a s t s a bou t 7w eek s . B i r th w e igh t r a nge s f r om 7 5 132 g ,a nd n ea r a du l t w e igh t i so b t a ined b yt h ee nd o ft h ef i r s tc a lenda ry ea ro fl i f e . P e rmanen t d en t i t ion i si np l a ce b y a ge 4 mon th s , w h ich c o inc ide s w i th w ean ing . L i f e span i nt h e w i ld i su n l ike ly t oe xceed 6y ea r s ( Co rbe ta nd H a r r i s 1 99 1 :422 ) .

L u t ra l i t t ra ( o t t e r ) ( F igu r e 3 . 35 ) L . l u t ra r a nge c ove r s t h e e xpan se o f t h e P a lea r c t i c r e g ion f r om t h e B r i t i sh t ot h e J apane se i s land s . S pa in a nd t h eM ed i te r ranean r e g ion a r ei n c luded i nt h i s a r ea , a sa r eC h ina a nd S ou thea s t A s ia . T h et u nd ra z one i sn o t i n h ab i ted . T h i sh i gh ly a qua t ic s p ec ie s i sa s soc ia t ed w i th r i ve r , l a ke , s t r eam a nd ma r sh h ab i ta t s . T h i s s p ec ie s ' a du l t m a s s a ve raged 1 0 . 1 k g f o rm a le s( n =4 33 )a nd 7 . 0 k g f o r f ema le s ( n =2 20 )a ss umma r iz ed f o rp opu la t ion s i nt h e G r ea tB r i ta in ( Co rb e t a nd H a r r i s 1 99 1 : 4 27 ) . D en ta l f o rmu la f o l low s 3 . 1 .4 . 1 /3 . 1 .3 .2 .

T he d enn ing b ehav io r o f M . m e / e s m ake s i t s t r app ing o r s na r ing l e s s d i ff i cu l t , s i nce t h e se m ay b e p l a ce s o a st oi n t e r s e c te n t rance so ro t he rc hokepo in t s . G u lo g u lo ( wo lve r ine ,g l u t ton ) ( F igu r e 3 . 37 ) T h i s l a rge s tm embe r o ft h e Mu s te l idae r a nge s t h roughou t t h e f o r e s t t u nd ra a nd b o r ea l r e g ion s o f t h e H o la r c t ic ,t h ough c ompe t i t ion w i th h uman sm ay h ave c au sed c on t rac t ion f r om t h em o r es ou the r ly p a r t so fi t sr a nge . A du l t m a s s n o rma l ly r a nge s f r om 1 0 .9 18 .1 k g , b u t may r e a ch u p t o3 2 .0 k g ( s umma r iz ed b y P a s i t schn iak -A r t s a nd L a r iv ie r e 1 995 :1-2 ) . T he f a vo r ed h ab i ta to ft h i s s p ec ie s i s ma tu r et i m b e r s t and s , e s pec ia l ly c on if e r s . S h rub , r o ck , t u nd ra , a nd i c e a r e a l so o ccup ied ( P a s i t schn iak -A r t s a nd L a r iv ie re 1 995 :6 ) . V a r ia t ion s i nh ab i ta t u sage p r ima r i ly r ef lec t c hange s , b o th s e a sona la nd c yc l i ca l ,i np r ey a va i lab i l i ty .

B r eed ing i n t h i s s pec ie s i s a s ea son a l , t h ough b o th mo re b i r th s a nd m o r e s ucce s sf u l b i r th s a r e r e co rded f o r s umme r m on th s , e s pec ia l ly i nt h o se p o r t ion s o f i t s e x ten s ive r a nge w he r ew in te r s a r e s eve r e . Ma t ing t h e r ef o r e i s mo s t l i ke ly t o o ccu r i ns p r ing . T he r e i sn o d e lay ed i mp lan ta t ion o f e m b ryo s ,a nd g e s ta t ion l a s t s a bou t 6 2 -63 d ay s . L i t t e r s a r et y p i c a l ly 2 3 n ewbor n , t h ough t h e k nown r a nge i s 1 5 . T he y oung a r e w eaned a t1 4w eek s , b u t s t ay w i th t h e mo the r f o r 7 12 m on th s a nd d o n o t r e a ch s e xua l m a tu r i ty u n t i l d u r ing t h e i rs e cond y ea ro fl i f e( Co rbe ta nd H a r r i s 1 991 :429 ) .

T he i r d i e t i sc a rn ivo rou s . Sma l le r p r ey a r e u sua l ly h un t ed , w h i le l a rg e ra n ima l ss u ch a sc a r ibou a nd e l k a r e u sua l ly p ro c u red t h rough s c aveng ing . D eep s now t os l ow o rt r ap l a rge r m amma l s s e em s t o b e ar e qu i r emen t f o r t h e i r d i rec t p ro c u r emen t ( P a s i t s chn iak -A r t s a nd L a r iv ie re 1 995 :5 ) . I n d i v i dua l s a r e s o l i ta ry a nd h ave l a rge h ome r a nge s , b u t t h e se

O t t e r f u r i sp a r t icu la r ly p r iz ed f o r i t s i n su la t ing a nd w a t e r p roof c ha r ac t e r i s t ic s , a sw e l l a sl a rge s i z e mak ing f ewe r f u r s n ece s sa ry t oc omp le te au s ef u l g a rmen t . Ma le s a nd f ema le s i n hab i t s o l i ta ry r ange s , w i th mu ch o ve r lap b e tween t h em . T h i s s p ec ie s l e ave s v i s ib le t r a i l s a nd h au l ing -ou t p l a ce s n ea r w a te r e dge s . D u r ing t h e d ay t ime , s h e l t e r i sf ound a bove g round u nd e r c ove r o r b e low g r ound i nn a tu ra l f i s su re s o r 4 3

f r ee ly o v e r lap w i th t h o se o fo t he r s . A c t iv i ty i s mo s t ly n oc t u rna l ,b u tt h i sm ay v a ry .

c ompan ied b y o t he r e x t reme c o ld i n d ica t ive s p ec ie s l i k e r e indee ra nd s nowy o w l .

S i nce e mb ryo i mp lan ta t ion i sd e layed , m a t ing o ccu r s i n s umme ra nd b i r th i sn o tu n t i lt h ee nd o ft h ef o l low ing w in t e r , u sua l ly b y l a t e Ma r ch . A c t ive p r egnancy r a nge s f r om 3 0 -50 d ay s . L i t te r s i z e v a r ie s b y r e g ion , b u t2 3 i sa ve rage . B i r th w e igh t a ve rage s 8 4 g , w ean ing o ccu r s a t7 8 w eek s , a nd a du l t s i z e i sr e a ched b y t h e w in t e r f o l low ing b i r th w hen j u v en i le d i spe r sa l o ccu r s . S exua l m a tu r i ty i sr e a ched f o r b o th s exe s a tt h e a g e ay ea r a nd a h a lf ( Pa s i t schn iak -A r t s a nd L a r iv ie r e 1 995 :5 ) . G . g u lo f u r i sp a r t icu la r ly l o ng , l u sh a nd w a rm , a nd t h e i r p ropen s i ty f o r s c aveng ing m ake s t h em a t t r a c tab le t ot r ap sb a i t ed w i th c a r r ion .

Ma t ing o ccu r s a bou t A p r i l , w i th a6 w eek g e s ta t ion p e r iod a nd b i r th o fp up s i n May o rJ une . O lde rf ema le s s ome t ime s h ave as e cond l i t t e r i nt h e s ame s umme r . L i t te r s i z e i sq u i t e l a rge : 5 8a nd 4 10 a r er e po r ted a st y p ica l ,a nd s ome e x t r eme c a se s o fu pt o2 3 i n as i ng le l i t t e r a r e a l so k nown . B o th p a r e n t s p a r t ic ipa te i nr a i s ing t h e y oung , w i th t h e m a le o f t en p rov i s ion ing t h el a c ta t ing f ema le ( Ognev 1 962 ) . A . l agopu s f u r i sl o ng , l u sh , w a rm a nd w h i te , m ak ing i ta v a luab le a cqu i s i t ion f o rh un te r sa nd t r app e r s i n to t h i sc e n tu ry ( Ognev 1 962 )a nd i d ea l ly s u i ted f o rw in t e rc l o th ing i ng l a c ia l a ge E u ra s ia .

CARN IVORA ,F AM ILY CAN IDAE T he t e e th o ft h e t wo s p ec ie s d e sc r ibed b e low a r e s im i la r i n m o rpho logy ,w i th t h o se o fA lopex l a gopu sb e ing s ma l le ra nd h av ing r e du ced d i s ta lm o la r sa sc ompa r ed w i th V u lpe sv u lpe s ( H i l l son 1 986 ) . T he i rd en ta lf o rmu la f o l low s3 . 1 .4 .2 / 3 .1 .4 .3 .

CARN IVORA , F AM ILY F EL IDAE B o th o f t h e t wo s p ec ie s d i scu s sed b e low h a s t h e r e du ced d en ta l f o rmu la c ha ra c te r i s t ic o ft h e F e l idae : 3 . 1 .2 .1 / 3 . 1 .2 . 1 a nd a r e c l ea r ly d i s t ingu i shab le f r om o t he r s ma l l c a rn ivo r e s o n t h e se g round s . S i ze d i f f e rence s b e tween t h e se t wo s p e c i e sa low st h e i rs e pa ra t ion b yt oo th s i ze .

V u lpe sv u lpe s( r ed f o x ) ( F igu r e 3 . 38 ) T h i s u b iqu i tou s H o la r c t ic s p ec ie s h a s a n e x p an s ive r a nge e x t end ing a l so i n to N o r th A f r ica a nd t h e A ra b i an p en in su la . C o rbe t a nd H a r r i s ( 1 99 1 :355 ) r e por t b ody s i z em ean sr a ng ing f r om 6 . 4 -7 .3 k g f o rm a le s a nd 5 . 4 -6 .2 k g f o r f ema le s f r om p opu la t ion s i nt h e U n i t ed K ingdom . I t s h ab i ta t r e qu i r emen t s a r e p a r t o ft h e r e a son f o r i t s a dap t ive s u cce s s : i ta dap t sr e ad i ly t om o s th ab i ta t s , e ven u rban , w i th in t h i s r a nge . F ragmen t ed , h i gh ly v a r iab le h ab i ta t s s upp ly ing a n a bundan t a r ray o ff o rag ing c ho ice s a r e p r ef e r r ed ( Co rbe t a nd H a r r i s 1 99 1 :357 -358 ) . S ma l l m amma l s , e s pec ia l ly r a b b i t s , m ak e u p t h e m a jo r i ty o f p r ey i t em s , b u t a dd i t iona l ly b i rd s , a mph ib ian s , i n ve r t eb ra t e s , a nd p l an tm a t e r ia l s a r e c on s umed ( Go sz czyn sk i 1 974 , H owe s 1 978 , L ock ie 1 959 , R ob e r t son a nd Whe lan 1 987 , S co t t 1 94 1) . H uman supp l ied r e s ou r ce s s u ch a s f a rm p ou l t ry o rg a rbage d ump s a r e r e ad i ly e xp lo i ted . S uch c amp -f o l lowe r b ehav io r m ay h ave b egun e a r ly i np r eh i s to ry , a nd i tm ay b e r ef l ec t ed i nt h e p a t te rn s o f s ke le ta l r ema in s f r om p r eh i s to r ic s i t e s . A c t iv i ty i sm a in ly n oc tu rna l a nd c r epu scu la r , b u t e ven t h i s i sm u tab le . V . v u lpe si st e r r i to r ia la nd l i ve si nf am i ly g roup s .

F e l l ss i l ve s t r i s( w i ld ca t ) ( F igu r e 3 . 40 ) F .s i lve s t r i s c an t oday b e f ound f r om S pa in , t h rough t h e c i r cum -Med i te r ranean r e g ion , A rab ia , I n d ia , A f r ica , a nd t ow e s ter nC h ina . P opu la t ion sa l so e x i s ti n af ew n o r ther n E u ropean a r ea s , s u ch a s S co t land a nd n o r ther n F rance . Med i t e r r anean f o rm s o ft h i s s p e c ie s a r e b e l ieved t o b e t h e w i ld p r ogen i to r s f o r d ome s t ic c a t s , F .c a tu s . A s w i th a ny d ome s t i ca ted s p ec ie s , o ne may e xpec tt h a tt h er a ng ea nd c ha rac te r i s t ic s o ft h e w i ld f o rm may h ave b een g r ea t ly a l t e r ed b yh uman i n t e rven t ion . T he e x i s tence o ff e ra l a nd h y b r id f o rm s f u r the r b l u r s u nde r s tand ing a bou t F . s i l ve s t r i s i n i t sp r edome s t i ca t ed s t a te . T yp ica l ly , F .s i l ve s t r i s i sl a rge r a nd l e s s g r a c i le t h an i t sd o m e s t ica t ed c oun t e rpa r t . C o rbe t a nd H a r r i s( 1 99 1 :434 )r e po r t m ean w e igh t s o f5 . 2 , 4 . 7 a nd 4 . 4 k g f o r m a le s a nd 4 . 5 , 3 . 9 a nd 4 . 0 k g f o r f ema le s o ft h r ee S co t t i sh ( a du l t ) p opu la t ion s . D ie t i se c lec t i c , r ef l ec t ing t h e l a rge a nd d i ve r se r a nge o ft h i s s p ec ie s w i th i t s m any f o od o p t ion s . Mo s t s ma l l m amma l s a nd b i rd s a r e t a ken a nd a r e s upp lemen ted w i th s ma l le r a moun t s o fa mph ib ian , r e p t i le s , a nd i n sec t s . P r ef e r r ed h ab i t a t s a r e f o r e s t a nd b ru sh a r ea s ( Co rbe t a nd O vend en 1 982 :188 ) . A c t iv i ty i sp r ima r i ly s o l i ta ry a nd n oc tu rna l , a nd t e r r i to r ie sa r em a in ta ined .

Ma t ing o ccu r s m a in ly i nJ a nua ry , w i th b i r th s f o l low ing i n Ma r ch . G e s ta t ion l a s t s 5 2 -53 d ay s , a nd l i t t e r s o f 4 5 a r e t y p ica l . B i r th w e igh t r a ng e s f r om 8 0 120 g a nd w ean ing o ccu r s a ta bou t 7w eek s . Y oung a r e b or n i n ab u r row a nd e me rge f o rt h e f i r s t t i me a r ound 4w eek s , a tw h ich a ge t h ey t h e ref o re a r e m o r e s u scep t ib le t op r eda t ion . A du l t s i z e a nd i n depend ence i sr e a ched b y S ep t embe r o f t h e b i r th y ea r ( Co rbe ta nd H a r r i s 1 99 1 :363 -364 ) .

O ne l i t te rw i t h a n a ve rage s i ze o f3 . 7 i sp roduced ( r ange : 2 6 ) ,n o rma l ly i n May a f t e r ag e s ta t ion p e r iod o fa bou t6 d ay s ( Co rbe ta nd H a r r i s 1 99 1 :436 ) . B ir th w e igh ti sf r om 1 0 0 163 g , w i th p e rmanen t d en t i t ion a r r iv ing a t a bou t 6m on th s . Mo s t a du l t m a s s i so b ta ined b y 1 0 mon th s , b u t e p iphy sea l f u s ion i sn o t c omp le t e u n t i l 1 8 19 mon th s . Y oung l e ave t h e i r mo the r s i nt h e a u t umn a nd d i spe r se t of o rm t h e i r o wn h ome t e r r i to r i e s , a nd t h e r ef o r e t h ey m ay b e m o r e v u lne r ab le t op r eda t ion a tt h a tt i me .

A lopex l a gopu s( a r c t ic f o x ) ( F igu re 3 . 39 ) T h i s a r c t ic s p e c ie sh a s ac i rcumpo la r d i s t r ibu t i on , e x t end ing i nt h e P a lea r c t ic f r om S weden t ot h e K am c ha tka p en in su la . T h i sr a nge e x t end ed f a rs ou thwa rd sd u r ing g l a c ia l e p i sode s a nd i n c lud ed i n t e rm i t t en t p ene t ra t ion i n to n o r ther nS pa in . I t sn umbe r st h e r ew e r e n eve rn ume rou s a nd p robab ly r ef l ec t a r ea s o fa l p in e t u nd ra e xpan s ion d u r ing t h e c o lde s t p e r iod s . A bundance i n F rance w he r e t r ue t u nd ra c ond i t ion s w e r e o b ta ined w a s m o r e c ommon a nd w a s a c -

F e l l sl y nx ( l ynx ) ( F igu r e 3 . 4 1) T h i s s p ec ie sh a s aH o la r c t i c , b o rea l f o r e s td i s t r ibu t ion t h a ts t r e t che s f r om t h e mon tane r e g ion s o ft h e I b e r i an p en in su la t h rough S i be r ia . I t s r a nge i ss im i la r ly e x t en 4 4

CARN IVORA ,F AM ILY V IVERR IDAE G ene t ta g ene t ta ( g ene t ) ( F igu r e 3 . 42 ) T he m embe r s o ft h i s g enu s ( numbe r ing a bou t 1 0 ) a l h ave as ub -Saha ran A f r ican d i s t r ibu t ion , w i th t h e e x c ep t ion o f G . g ene t ta w h ich e x t end s f r om t h roughou t t h i s a r ea i n to t h e I b e r ian p en in su la , F rance , N o r th A f r ica a nd A rab ia . C o rbe t( 1 978 :177 ) t h e ref o r e s ugge s t st h a tt h e E u ro p ean r a nge o fg ene t sr e su l t ed f r om h uman i n t roduc t ion . T h i s s p ec ie s , h oweve r , h a s b een i d en t if i ed f r om o ne n o r ther n S pan i sh p a leo l i th ic c on t ex t : 2r ema in s f r om A z i l ian L eve l I a tE l P endo ( Fuen t e s V ida r te 1 980 ) . S ign if i can t n o r ther n s p r ead o f t h i s t r op ica l / sub t rop ica l s p ec ie s w a s u n l ike ly t o h ave o ccu r r ed p r ev iou st on ea r -comp le t eg l ac ia lr e t r ea t .

s i ve i n t h i s z one o f N o r th Ame r ica . I t s c u r r en t r a nge i n E u rope i ss e ve r e ly r educed t oi s o la t ed a l p ine r e g ion s . T h i s s p ec ie s i ss ome t ime s p l aced i n to a s e pa ra te g enu s ( Lynx ) a l ong w i th F . r uf u s ( t he N o r th Ame r ican b obca t ) , t h ough C o rbe t ( 1 978 ) r e ta in s t h i s d i s t inc t ion o n ly a tt h e s ubgenu s l e ve l . T he I b e r ian v a r ie ty i sa l so s ome t ime s g i ven s epa ra te s p ec ie ss t a tu s( F .p a rd ina ) , b u tt h ea u tho rd oe sn o tr e cogn iz e t h i s s e pa ra t ion a sv a l id . A dd i t iona l ly , t h e N ea rc t ic l y nx i s s ome t ime s d e s igna ted a s as e pa ra t e s p ec ie s ( F e l i s lLynx C a n ad en s i s ) f r om t h e P a lea r c t ic v a r ie ty , b u t t h i s d i s t inc t ion m u s ta wa i tf u r the ri n ve s t iga t ion . A du l tb ody m a s s f o r ma le s i nS weden a v e rage s 1 7 .9 k g a nd i sg ene ra l ly l a rge r i nt h e P a lea r c t ic t h an i n t h e N ea r c t ic ( s umma r iz ed b y T um l i son 1 987 ) . S i z e si nS pa in t e nd t ob e much s ma l le r , a ve rag ing 5 . 7 k g ( n =1 0 ) f o rj uven i le s l e s s t h an 1y ea r o l d a nd > 9 .3 12 .8 k g ( n =1 3 ) f o r s ubadu l t s /adu l t s i nD o f iana N a t iona l P a rk ( Be l t rän a nd D e l ibe s 1 993 ) .

G . g ene t ta i s as ma l l c ar n ivo r e r a ng ing f r om 1 360 -2250 g a du l tb ody m a s s( N ie thamme ra nd K rapp 1 993 ) . I t st e e th a r e i d en t if iab le mo rpho log ica l ly f r om o t he r s ma l l c a rn ivo r e s o f s im i la r s i ze ( s uch a s Ma r te s o r Mu s te la ) a nd f o l low a 3 . 1 .4 .2 / 3 .1 .4 .2 f o rmu la . I ti ss o l i ta ry , s t r ic t ly n oc tur na l a nd s u rv ive so n ag ene ra l i s td i e to f mo s ts ma l la n ima l s , i n c lud ing b i rd s , f r u i t s , m amma l s , a nd i n ve r teb r a te s ( L e J a cque s a nd L ode 1 994 ; V i rgö s e ta l . 1 996 ) . T h i s s p ec ie s i sa s soc ia ted w i th f o r e s t a nd s h r ub i n i t s E u ropean r a nge . I t s b ehav io r make s i te a s ie s tt oa cqu i re b y t r app ing . I t s a ppea rance i n d i c a t e s s i gn if i can t w a rm ing a nd c l o s ing o fe nv i ronmen t s d u r i n gt h eH o locene i nC an tab r ian S pa in .

Ma t ing o ccu r s i nF eb rua ry a nd Ma r ch i nt h e P a lea r c t ic r e g i on , a nd s ome t ime se a r l ie ri nt h em o r e s ou the r ly p o r t ion so f t h i s r a ng e . E s t ima t ion o f g e s ta t ion t i me s pan 6 3 -74 d ay s , w i th b i r th s o f l i t te r s i nl a t e May o r e a r ly J une . L i t t e r s i z e r a nge s f r om 1 6 , w i th 2 3 t y p ica l i nt h e P a lea r c t ic r e g ion . D ec iduou s d en t i t ion b eg in s r e p la c emen t a t 4 mon th s . Wean ing i sc omp le te b y s i x mon th s o f a ge , t h ough s ome l i t t e r s r ema in w i th t h e mo the r f o r af u l l y ea r a f t e r b i r th . C omp le t e e p iphy sea l f u s ion i s r e a ched b y 3 2 mon th s ( Tum l i son 1 9 87 :3 -4 ) .

L i t te r s a r e p roduced d u r ing t h e s umme r mon th s , r ough ly f r om A p r i l t h rough A ugu s t ( t hough t h e b r eed ing s e a son i s r e ve r sed i nt h e S ou ther n H em i sphe r e p o r t ion o f i t s r a nge ) . L i t t e r s i z e i sf r om 2 4 , w i th 2 . 6 a ve rage . G e s ta t ion l a s t s f o r 1 0 11 w eek s , w i th w ean ing o ccu r r ing a ta bou t 9w eek s . T he y oung a r e b or n a nd w eaned i n ar e u sed b u r row o r n a tu ra l h o l low ( Sk inne ra nd S m i the r s 1 990 :477 ) .

F avo r ed h ab i t a t i sc on if e rou s f o r e s t , b u t i n S pa in a r ea s o f t h ick b ru sh a r e s ome t ime s i n hab i t ed ( Co rbe t a nd O vend en 1 9 82 :189 ) . A c t iv i ty p a t te rn s a r e s o l i t a ry a nd n oc tu rna l , a nd p r inc ip le p r ey i n c lude s s ma l l m amma l s s u ch a s l a gomo rph s a nd r oden t s( S aunde r s 1 963 ) .

4 5

T ab le 3 . 1 : S umma ry o fE co log ica l C ha r ac te r is t ics . F o r e s t

G l i sg l i s E l iomy sq ue rc inu s A pod emu sf l av ico l l i s

X X X

A pod emu ss y lva t icu s C le th r ionomy sg la reo lu s

X X

Mar te sm ar te s

X

Ma r te sf o ina F e l l ss i lve s t r i s

X X

F e l l sl y nx G ene t ta g ene t ta E r inaceu se u ropaeu s

X X

Me /e sm e /e s M icro tu sn i va l i s

S h rub s

F o res t A lp ine Wa te r H um id E dge / E dge / Meadow Meadow We t land s

D ry B o rea l / Meadow T und ra

G ene ra l

X X

G a lemy sp y rena icu s Ma rmo ta m armo ta

X X X

X

A rv ico la t e r re s t r i s A rv ico la s ap idu s M icro tu so economu s

X X X

N eomy sf od ien s N eomy sa noma lu s Mu s te la l u t reo la

X X X

L u t ra l u t ra P i tymy sl u s i tan icu s T a lpa e uropaea T a lpa o cc id en ta l i s S o rex m inu tu s S o rex c orona tu s M i cro tu sa r va l i s M icro tu sa g re s t i s M icromy sm inu tu s C roc idura r u s su la L epu sc apen s i s L epu st i m idu s O tyc to lagu sc un icu lu s M icro tu sg rega l i s

X

X X X

X X X X X

X X X

X X X X X X X

X

X

G u lo g u lo A lopex l agopu s P i tymy sp y rena icu s Mu s te la n i va l i s Mu s te la e rm inea

X X X X X

Mu s te la p u to r iu s V u lpe sv u lpe s

X X

X= p r ima ry a s soc ia t ion x =s e conda ry a s soc ia t ion

4 6

F igure 3 . 1 : Mode rn R ange o fG lä g l i s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 99 1 ; C o rbe ta nd O venden 1 982 ) .

F igure 3 . 2 : Mode rn R ange o fE l iomy sq uerc inu s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 991 ; C o rbe ta nd O venden 1 982 ) .

F igure 3 . 3 : Mode rn R ange o fM i c ro tu sa g re s t i s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 991 ; C o rbe ta nd O venden 1 982 ) .

o l t z fo i s te r s 4 N e a r A r i i g e k \4 e m l i t t e e e 4 r i a ie ey 4 e4 " / 41

tor t

F igure 3 . 4 : Mode rn R ange o fM ic ro tu sa r va l i s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 99 1 ; C o rbe t a nd O venden 1 982 ) .

F igure 3 . 5 : Mode rn R ange o fM icro tu sn i va l i s r e d rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd O venden 1 982 ; N ie thamme ra nd K rapp 1 982 ) .

pop wo rw o in w w _ i lr e i

p r o p r

ai l

e r . 40 1 1 1 1 K

mee , „ I4 rJ44 e . ep i t i l l g w P A T Veive -

A

l

Ä , ) ‘ ,Ä rt e a o de ) L A r a i d r i l i k d i r 4t. 4

l y r

i

F igure 3 . 6 : Modem P a lea r c t ic R ange o fM icro tu so economu s( e x tend ing i n to t h eN ea rc t ic a sf a ra sC anada ) ( r ed rawn a f t e rC o rbe t 1 978 ;C o rbe ta nd O venden 1 982 ) . 4 8

F igure 3 . 8 : Mode rn R ange o fC le thr ionomy sg lareo lu s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 991; C o rbe ta nd O venden 1 982 ;N ie thamme ra nd K rapp 1 982 ) .

F igure 3 . 9 : Mode rn R ange o fP i ty r ny sp yrena icu s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd O venden 1 982 ; N ie thamme ra nd K rapp 1 982 ) .

F igure 3 . 10 : Mode rn R ange o fP i tymy sl u s i tan i cu s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd O venden 1 982 ; N ie thamme ra nd K rapp 1 982 ) .

F igure 3 . 11 : Mode rn R ange o fA rv ico la t e r re s t r i s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 991 ; C o rbe ta nd O venden 1 982 ) .

F igure 3 . 12 : Mode rn R ange o fA rv ico la s ap idu s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd O venden 1 982 ) .

l t

d(

a f t

‘ .

%Ä

z r i t g

F igure 3 . 13 : Moder nR ang eo fA pod emu ss y lva t icu s r e d rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 991 ; C o rbe ta nd O venden 1 982 ) .

F igure 3 . 14 : Mode rn R ange o fA pod emu sf la v ico l l i s r e d rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 99 1 ; C o rbe ta nd O venden 1 982 ) .

F igure 3 . 15 : Moder nR ang eo fM ic romy sm inu tu s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 99 1 ; C o rbe ta nd O venden 1 982 ) . 5 1

F igure 3 . 16 : Mode rn R ange o fC roc idura r u s su la ( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd O venden 1 982 ; N ie thamme ra nd K rapp 1 990 ) .

F igure 3 . 17 : Mode rn R ange o fS orex m inu tu s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 99 1 ; C o rbe t a nd O venden 1 982 ;N ie thamme ra nd K rapp 1 990 ) .

F igure 3 . 18 : Mode rn R ange o fS orex c o rona tu s r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd O venden 1 982 ; N ie thamme ra nd K rapp 1 990 ) . 5 2

F igure 3 . 19 : Mode rn R ange o fN eomy sa noma lu s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd O venden 1 982 ; N ie thamme ra nd K rapp 1 990 ) .

F igure 3 . 21 : Mode rn R ange o fT a lpa e u ropaea ( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 991 ; C o rbe ta nd O venden 1 982 ) .

F igure 3 . 22 : Modern R ange o fT a lpa o cc id en ta l i s r ed rawn a f te rC o rbe t 1 978 ; C o rbe ta nd O venden 1 982 ; N ie thammer a nd K rapp 1 990 ) .

F igure 3 . 23 : Modern R ange o fG a lemy sp y rena icus ( r ed rawn a f te rC o rbe t 1 978 ; C o rbe ta nd O venden 1 982 ; P a lme i r im a nd H of fman 1 983) .

F igure 3 . 24 : Moder nR ange o fE r inaceu se uropaeus ( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r is 1 991 ; C o rbe ta nd O venden 1 982 ) .

5 4

F igure 3 . 25 : Mode rn R ange o f Ma rmo ta marmo ta ( r ed rawn a f te rC o rbe t 1 978 ; C o rbe ta nd O venden 1 982 ; N ie thamme ra nd K rapp 1 978 ) .

' 1 F igure 3 . 26 : Modern R ange o fO ryc to lagu sc un icu lu s( a l so i n t roduced i n to A u s t ra l ia a nd N ew Z ea land ) ( r ed rawn a f te rC o rbe t 1 978 ;C o rbe ta nd H a r r i s 1 991 ; C o rbe ta nd O venden 1 982 ) .

o r am m er i l‚ ree 44‘ 4 1 1 16?"1 1 r

4 , 1 1L 1i k e t w i r

4

F igure 3 .27 : Mode rn R ange o fL epu sc apen s i s( ex tend ing t h rough A f r ica )( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r i s 1 99 1 ; C o rbe ta nd O venden 1 982 ) . 5

i n p oP m , A • o v iae l1 1 1 , 40

‚' m e i m ie s i , e -.

elel e 14

12

.t Ir 4 %

e re

m g

* »

4 h 4 1

ri ä 4 r o k i r i= l ti F igure 3 . 28 : Moder nP a lea r c t ic R ange o fL epu st i m idu s( e x tend ing a l so i n to t h eN ea rc t ic )( r ed rawn a f t e r

C o rbe t 1 978 ;C o rbe ta nd H a r r i s 1 99 1 ;C o rbe ta nd O venden 1 982 ) .

1 0 1 r ' e V 7 ' P m . e i l g zi f ir a t w ' 4 1 1 , A ht ie l e „

, 1e 1 Ww e n11 11 rw e e

kA 1 (.

* t ' I ‚ 4 ) P 4 °\ e ‚ l

g e R i le i r 1a d d m y o u1 .1 1 1 1

F igure 3 . 29 : Mode rn P a lea r c t ic R ange o f Mu s te la n i va l i s( e x tend ing a l so i n to t h eN ea rc t ic )( r ed rawn a f t e r C o rbe t 1 978 ;C o rbe ta nd H a r r i s 1 991 ; S heff ie ld a nd K ing 1 994 ) .

111 111 1

,

21

m , me w r o d oi ru

1

0i l

>

,

)

m

4 1

4 P411p Os

41 * * 1 L ‘ eh d i th ' an I l L4 ,

F igure 3 . 30 : Mode rn P a lea r c t ic R ange o f Mu s te la e rm inea ( ex tend ing a l so i n to t h eN ea rc t ic )( r ed rawn a f t e r

C o rbe t 1 978 ;C o rbe ta nd H a r r i s 1 99 1) . 5 6

F igure 3 .31 : Modem R ange o f Mu s te la p u tor iu s( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe t a nd H a r r i s 1 991 ; C o rbe ta nd O venden 1 982 ) .

F igure 3 .32 : Modem R ange o f Mu s te la l u treo la ( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe t a nd Ovenden 1 982 ; Y oungman 1 982 , 1 990 ) .

F igure 3 .33 : Modem R ange o f Ma r te s mar tes ( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r is 1 991 ; C o rbe t a nd O venden 1 982 ) .

F igure 3 . 35 : Mode rn R ange o fL u t ra l u t ra ( e x tend ing a l so i n to S ou thea s tA s ia )( r ed rawn a f t e rC o rbe t 1 978 ; C o rbe ta nd H a r r is 1 99 1 ;C o rbe ta nd O venden 1 982 ) .

F igure 3 . 36 : Mode rn R ange o f Me le sm e le s( r ed rawn a f t e rC o rbe t 1 978 ;C o rbe ta nd H a r r i s 1 99 1 ;C o rbe t a nd O venden 1 982 ) .

F igure 3 .37 : Modern P a lea r c t ic R ange o fG u lo g u lo ( e x tend ing a l so i n to t h eN ea rc t ic )( r ed rawn a f t e r C o rbe t 1 978 ; P a s i t schn iak -A r t s a nd L a r iv ie re 1 995 ) .

F igure 3 .38 : Mode rn P a lea r c t ic R ange o fV u lpe sv u lpe s( ex tend ing a l so i n to t h eN ea rc t ic )( r ed rawn a f t e r C o rbe t 1 978 ; C o rbe ta nd H a r r is 1 991) .

F igure 3 . 42 : Moder nR ange o fG ene t ta g ene t ta ( who se p r ima ry r ang e i sS ub -Saha ran A f r ica )( r ed rawn a f t e r C o rbe t 1 978 ; S k inne ra nd Sm i the r s 1 990 ) .

CHAPTER 4 MODERN MICROFAUNA OF CANTABR IAN S PA IN

T he mod e rn m ic rof auna o f t h e C an t ab r ian r e g ion a nd t h e s u r round ing I b e r ian p en in su la i sp r e s en ted h e r e i no rde r t o s e rve a s ab a se l ine f o r c ompa r i son t ot h e p a leom ic rof auna . T he se d a ta c once rn ing t h e c on t empo r a ry f a una c ome s f r om b o th t r app ing s t ud ie s a nd t h e a na ly s i s o fr a p to r p e l le t s am p l e s . N ew d a ta f r om e a ch o ft h e se s o u r ce s c o l l ec t ed b y t h e a u tho r i sa l so p r e sen ted . B e s ide s t h e s t r ic t ly p r a c t ica l t a sk s o fr e p r e sen t ing t h er e la t ive m ic rof auna lp opu la t ion d en s i t ie s , c ompo s i t ion s , a nd v a r ia t ion a c ro s s c l ima t ic , a l t i t ud ina l a nd l a t i tud ina l c l ine s , t h e r e la t ive u t i l i ty a nd b i a s e s o ft h e se t wo s ou rce s o fd a t aw i l l a l so b e m ade a ppa r en t ( Pu lk 1 976 ) . A s d i scu s sed i nC hap te r 2 , t h e s ou r ce o f m o r ta l i ty f o r a n a r c haeo log ica l o rp a leon to log ica l s amp le o fs ma l lm amma l s i s o fe x t r eme i mpo r tance i ni t si n te rp r e ta t ion . T h i s s ame s i tua t i on i sl o g ica l ly t r ue f o r am oder n s amp le , s i nce t h e o n ly e s sen t ia l d i f f e r ence i si nt h e r e cen tne s s o fm o r ta l i ty . A dd i t i ona l ly , e xam ina t ion o ft h e mode rn s ma l l m amma l f a una o f ar e g ion a nd i t sr e la t ion sh ip t ot h e c on t empo ra ry c l ima t e a nd e nv i ronmen t w i l l i l u s t ra te t h e r e la t ion sh ip b e tween P l e i s to c ene f a una sa nd t h e i ra s soc ia ted p a leo env i ronmen t s (W e s t er n 1 980 ) .

Wh i le t r ap a r ray s o ft h i s c onf igu r a t ion a r e e f f ec t ive i nc ap t u r ing ab road s pec t rum o fs ma l lm amma l s , s ome e xcep t ion s d o o ccu r ( Boon s t ra a nd K r eb s 1 978 ; Maddock 1 992 ) . S h rew s n o rma l ly f e ed u pon ad i e to fm ixed i n ve r teb ra te s , s o p eanu t b u t te r / ro l led o a t s m ay b e al e s s e f f ec t ive e n t icemen t t o t h em . S im i la r ly , t h e i r s ma l l s i z e m ake s t r app ing t h em w i th s t anda rd s nap t r ap s d i ff icu l t . T he se s pec ie s s e emed t o b et r apped mo re f r equen t ly d u r ing e p i sode s o fp oo rw ea the r , i n d ica t ing p e rhap s t h a t t h ey m u s t e xpand t h e i r f e ed ing c ho ice s w hen u nde r s t r e s s ( p e r sona l o b se rva t ion ) . M i c r o t ine s ( vo le s , e t c . )a r e a l so mo r e d i f f i cu l tt ot r ap u s ing t h i s t y pe o f b a i t ( s ee r e su l t s b e low ) . L ig tvoe t a nd V an W i jngaa rden ( 1994 ) r e commend c a r ro t a sb a i t f o rt h e se s p e c i e s . Mu r id s( O ld Wo r ld m ice )h ave amo re e c lec t ic d i e ta nd a r e r e ad i ly t r apped u s ing p eanu t b u t t e r / ro l led o a t s b a i t . A r b o r ea l s p ec ie s s u ch a sd o rm ice ( G l i r idae ) a nd f o s so r ia l s p e c i e s s u ch a s mo le s ( Ta lp ida e )t e nd t ob e o ve r looked b y t r ap s p l a ced a tg round l e ve l . A l l c ap tu r ed s pec imen s w e r e w e ighed t o t h e n ea r e s t g r am a nd t h e f o l low ing m ea su r emen t s w e r e t a k en ( f o l low ing N a g o r sen a nd P e te r son 1 980 ) : t o ta l l e ng th , t a i l l e ng th , e a r l e ng th a nd h i ndf oo tl e ng th i n c lud ing n a i l ( Tab le 4 . 2 .1-4 .2 .8 ) . S ex , r e p roduc t ive s t a t e a nd /o r h i s to ry w a s r e co rded f o r e a ch i n d iv idua l w he r e p o s s ib le . S ome s c aveng ing f r om t r ap s d i d o ccu r , p a r t icu la r ly a tL oca t ion 4 b y l o ca l d ome s t ic c a t s a nd i no ne c a se b y amu s t e l id . S caveng ing a l so o ccu r r ed a tL o c a t ion s Ia nd 2 . T he se i n d iv idua l sw e r e n o t i n c luded i ns p e c i e st a l l ie s , s i nce t h e se c ou ld n o tb ed e t e rm ined i nm o s tc a se s a nd f ew m ean ingf u l m ea su r emen t s c ou ld b e t a ken . Mo s t s c avenged i n d iv idua l s a ppea r ed i n a ny c a se t o b e long t o A pod emu s , w h ich w a s b y f a r t h e m o s t f r equen t ly t r apped g enu s . I n a dd i t ion , s ome A pod emu s c ou ld n o t b e a s s igned r e l iab ly t oA . s y l va t i cu s o rA .f l a v i co l l i s , i n mo s t c a se s b e c au se t h ey w e re j uven i le s .

M ic romamma l ian s t ud ie s i n t h e I b e r ian p en in su la h ave a s ub s tan t ia l m ode rn h i s to ry . T he se h ad b egun a tl e a s t w i th t h e w o rk e a r l ie rt h i s c en tu ry b y C ab r e ra ( 1 907 , 1 908 , 1 9 14 ) . Mo re r e cen tw o rk i n c lude s s t ud ie s o nt h e f a una l d i s t r ibu t ion o ft h e n o r ther np en in su la rr e g ion b y H e im d e B a l sa c a nd d e B eauf or t ( 1 969 ) , G a rzon -Heyd t a nd C a s t rov ie jo ( 1 970 ) , C a s t idn a nd G o sä lbez ( 1 992a ) a nd A lmaa ( 1 973 ) . Much c u r r en t w o rk c once rn s d i s t r ibu t iona l v a r ia t ion s r e la t ed t o a l t i tude a nd /o rl a t i tude ( De l ibe sd eC a s t ro 1 985 ; Mo r eno a nd B a rbo sa 1 9 92 ; P a lomo e ta l . 1 994 ) o rh ab i ta t ( Bo rgh i e ta l . 1 994 ; F ae ta l . 1 992 ) . Ow l d i e ta ry s t ud ie so fr e la t ing t oI b e r i an s ma l l m amma l d i s t r ibu t ion a r e a l so t h e f o cu s o fm uch m ode rn r e sea r ch ( B rune t -Lecom te a nd D e l ibe s 1 984 ; C ama cho Muf toz 1 975 ; C ampo s 1 978 ;H e r r e r a 1 9 73 ;H e r r e ra a nd J ak s ic 1 980 ; J ak s ic e ta l . 1 982 ;L öpez -Go rdo e ta l . 1 977 ; V e iga 1 980 ) .

T ab le 4 . 1 : D i s t r ibu t ion o fT rap N igh t s( Can tab r ia ) . # of T rap N igh t s :

T RAPP ING S TUDY O FI BER IAN F AUNA T he a u tho rs amp led t h et e r r e s t r ia ls ma l lm amma l s i nt h ea r ea s u r round ing t h e s i te o f E l J uyo a nd o t he r l o ca t ion s i nt h e p rov ince o f C an tab r ia ( F igu r e 4 . 1) . At o ta l o f 2 000 t r ap n i gh t sw e r e a ma s sed ( Tab le 4 . 1) . T h i s t r app ing w a s s upp le m en ted b yt h e a na ly s i s o fo w la nd h awk p e l le ts amp le s f r om t h e s ame a r ea . Mo s t t r app ing w a s d one w i th s t anda rd Mu s e um S pec ia l s nap t r ap s ( 7 c m x1 4 c m ) , s u pp lemen t ed w he r e i n d ica t ed b e low w i th t h e l a rg e r ( 8 .3 c m x1 7 .6 c m ) V ic to r R a t s n ap t r ap s ( bo th manuf a c tu r ed b y t h e Wood s t r eam C o rpo ra t ion , L i t iz , P enn sy lvan ia , U SA ) . B a i t w a s u n if o rm ly am ix tu r e o f p eanu t b u t t e r a nd r o l led o a t s , f o l l ow ing N ago r sen a nd P e t e r son ( 1 980 ) . T rap s t a t ion sw e r e a t 1 0 m i n t e rva l s , w i th t wo t r ap s p e r s t a t ion . Whe r e p o s s ib le , p i tf a l l t r ap s s upp lemen t ed t h i s a r ray , b u t t h e i r u s e w a s l i m i t ed b y s h a l low s o i l a nd i n s ome i n s tance s , c on s tan t r a in . T he t r ap sw e r ec hecked e ve ry mor n ing .

L oc . 1

L oc .2

L oc . 3

L oc .4

L oc . 5

Mu seum S pec ia l

3 57

1 20

3 53

7 73

2 92

V ic to r R a t

0

3 9

2 6

0

0

P i tf a l l

0

0

1 6

2 4

0

TOTAL

3 57

1 59

3 95

7 97

2 92

( To ta l=2000 )

6 1

T ab le 4 . 2 : S pec ie sD imen s ions o fC an tab r ian S ma l l Mamma l s T ab le 4 . 2 .1 :A pod emu ss y lva t icu sD imen s ion s . # S ex l o c . L T a i l F oo t E a r

T ab le 4 . 2 .2 :A pod emu sf la v ico l l i sD imen s ion s . # S ex l o c . L T a i l F oo t E a r

W t .

mm 7 3

mm 2 2

mm 1 7

g 1 8

9 4001

M

8 6 8 7 8 6

2 3 2 4 2 1

1 7 1 6 1 6

1 3 1 8 1 7

9 4002 9 4003 9 4021

F F F

W t .

9 4004

F

1

mm 1 64

1

mm 2 13

mm 1 04

mm 2 4

mm 1 7

g 3 2

9 4005 9 4006 9 4007

F F M

1 1 1

1 67 1 74 1 70

1 1 1

1 63 1 59 2 08

8 1 6 0 1 02

2 2 2 3 2 4

1 7 1 7 1 7

1 7 2 9 2 9

9 4008

M

1

1 81

9 1

2 3

1 7

1 7

9 4031

M

1

1 94

1 02

2 4

1 8

2 6

9 4009 9 4010

M F

1 1

1 89 1 64

9 8 6 5

2 3 2 4

1 5 1 8

2 6 2 3

9 4032 9 4035

M F

1 2

1 6 1 1 53

8 6 6 2

2 4 2 3

1 7 1 7

2 4 2 2

9 40 11 9 4012

M F

1 1

1 62 1 62

7 5 6 9

2 1 2 3

1 5 1 6

1 4 1 6

9 4036 9 4037

F F

3 3

2 02 1 94

1 03 9 2

2 2 2 4

1 7 1 5

2 8 2 7

9 4013 9 40 14 9 4015

F M M

1 1 1

1 56 1 73 1 82

7 9 8 7 8 5

2 3 2 2 2 5

1 4 1 5 1 8

1 2 1 9 2 4

9 4039 9 4040 9 4041

M M M

3 3 3

2 00 2 17 2 07

1 02 1 09 1 07

2 3 2 5 2 4

1 7 1 8 1 8

2 6 3 4 2 9

9 40 16

F

1

1 74

8 8

2 1

1 7

1 5

9 4044

M

3

2 10

1 04

2 5

1 7

2 9

9 4017 9 40 18 9 4019 9 4020 9 4022 9 4023

F M F M F M

1 1 1 1 1 1

1 65 1 74 1 80 1 83 1 77 2 08

8 2 8 5 8 9 8 8 2 1 05

2 3 2 4 2 4 2 3 2 1 2 4

1 6 1 6 1 7 1 7 1 7 1 8

1 5 2 0 2 5 2 0 2 3 2 9

9 4045 9 4046 9 4061 9 4062 9 4063 9 4064

F M M M F F

3 3 4 4 4 4

2 11 1 9 1 1 9 1 1 86 1 76 1 78

1 08 1 0 1 1 00 9 2 9 1 8 9

2 4 2 5 2 4 2 4 2 3 2 3

1 8 1 7 1 7 1 6 1 6 1 7

3 2 2 2 2 3 2 3 1 6 1 9

9 4024 9 4025 9 4027 9 4030 9 4034 9 4038

F F M M M F

1 1 1 1 2 3

1 35 1 46 1 81 1 60 1 92 1 76

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

2 4 2 3 2 4 2 2 2 2 2 2

1 7 1 6 1 8 1 8 1 8 1 7

2 1 1 2 1 7 2 0 3 2 1 4

9 4065 9 4068 9 4069 9 4074 9 4075 9 4076

F M M M M F

4 4 4 4 4 4

1 71 1 40 1 74 1 87 1 83 1 83

8 7 5 4 9 1 9 7 9 2 8 9

2 3 2 4 2 4 2 5 2 5 2 4

1 7 1 7 1 6 1 6 1 8 1 7

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

9 4050 9 4051

F F

4 4

1 79 1 85

9 0 9 5

2 1 2 4

1 7 1 6

2 5 2 4

9 4084 9 4085

F F

4 4

1 97 1 65

1 0 1 8 7

2 5 2 2

1 7 1 5

2 7 1 6

9 4052 9 4053 9 4054 9 4055 9 4056 9 4059 9 4060 9 4066 9 4067 9 407 1

M M M F F F M M M M

4 4 4 4 4 4 4 4 4 4

1 80 1 72 1 64 1 51 1 80 1 65 1 74 1 65 1 85 1 95

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

2 4 2 3 2 4 2 2 2 3 2 2 2 4 2 3 2 3 2 4

1 6 1 6 1 6 1 6 1 6 1 7 1 6 1 7 1 7 1 7

1 8 1 8 1 6 2 2 2 2 2 2 1 8 1 8 2 0 3 2

9 4091 9 4105 9 5002 9 5007

F M M F

4 4 5 5

1 7 1 1 60 1 90 1 55

7 8 6 7 9 5 7 3

2 4 2 4 2 5 2 2

1 8 1 7 1 7 1 7

3 2 2 5 2 9 1 8

T ab le 4 . 2 .3 : R a t tu sr a t tu sD imen s ion s . # S ex l o c . L T a i l F oo t mm mm mm 9 4086 M 4 2 64 1 47 3 1

E a r mm 2 2

W t . g 5 6

9 4072 9 4073 9 4081 9 4090

M F F M

4 4 4 4

1 83 1 70 1 80 1 83

9 6 8 8 8 5 9 7

2 4 2 4 2 3 2 4

1 7 1 6 1 6 1 9

2 1 1 8 2 0 1 8

9 4093 9 4094

F M

4 4

2 00 1 83

1 03 9 6

2 3 2 3

1 7 1 6

2 4 2 4

9 500 1

M

5

1 89

1 00

2 4

1 7

2 2

9 5004 9 5005

M M

5 5

1 9 1 1 90

9 8 1 04

2 4 2 4

1 6 1 7

2 4 2 6

9 5008

M

5

1 88

9 2

2 3

1 9

2 4

T ab le 4 . 2 .4 : M ic ro tu sa g re s t i s D imen s ion s . # 9 5003

S ex M

l o c . 5

L

T a i l

F oo t

E a r

W t .

mm 1 40

mm 4 1

mm 2 0

mm 1 4

g 3 4

E a r

W t .

T ab le 4 . 2 .5 :S o rex c orona tu sD imen s ion s . # S ex l o c . L T a i l F oo t mm

6 2

mm

mm

mm

9 4042

F

3

17

4 6

1 3

7

7

9 4043

F

3

14

4 6

1 3

7

7

9 4092

M

4

16

4 0

1 3

7

9

9 4098 9 4103

F M

4 4

11 10

4 8 4 1

1 3 1 2

6 5

7 7

T ab le 4 . 2 .6 :S orex m inu tu sD imen s ion s . # S ex l o c . L T a i l F oo t mm mm mm 9 4047 F ? 3 1 05 4 1 1 9 4048

F ?

3

1 00

4 0

E a r mm 7

1

T ab le 4 . 2 .7 :N eomy sf od ien sD imen s ion s . # S ex l o c . L T a i l F oo t

7

E a r

T ab le 4 . 2 .8 : C roc idura r u s su la D imen s ion s . # S ex l o c . L T a i l F oo t Mm mm mm 9 4079 F 4 18 3 7 1 2

W t . g 3 . 5 3

E a r mm 9

9 4080 9 4 102

M M

4 4

9 4 1 0 1

3 7 3 9

8 1 3

8 8

W t .

9 4106 9 4107

M M

4 4

12 1 02

3 9 3 8

1 3 1 2

9 7

1 0 7

9 4108 9 4109

M M

4 4

1 05 9 9

3 6 3 6

1 3 1 3

8 9

9 7

9 4070

F

4

mm 1 37

mm 4 8

mm 1 7

mm 7

g 1 7

9 4099 9 4 100 9 4 104

F F M

4 4 4

1 28 1 38 1 32

5 2 5 0 5 2

1 8 1 8 1 7

8 7 6

1 2 1 1 0

L oc . =T r app ing l o ca t ion L =B ody l e ng th f r om n o se t oe nd o fl a s tt a i lv e r t eb ra ( mm ) T a i l =L eng th o ft a i lf r om b a se t oe nd o fl a s tv er teb ra ( mm ) F oo t =L eng th o fh ind f oo t , i n c lud ing n a i l( mm ) E a r =L eng th o fp innae ,e xc lud ing f u r( mm ) W t . = Ma s si ng rams I t a l ic s d eno t e t ha t am ea su remen t i si n comp le te d ue t op r e mo r tem i n ju ry o rp o s tmo r tem s c aveng ing .

F igure 4 . 1 : Map o fC an tab r ia S how ing T rapp ing L oca t ion s 1 5 a nd R oo s tL oca t ion sA -C .

C an tabr ian S ea

Roos t

T rapp ing L oc .4 ,5 R oos t sA ,B

P icos d e Europa

W t . g 9

S antande

C an tabr ia

T rapp ing L oc . 1 ,2 ,3

g f r

"

6 3

5 0 km

•

6 . 5 6 . 5

T ab le 4 . 3 : S p ec ie sD i s t r ibu t ion a mong T rapp ing L oca t ion s( Can tab r ia ) . L oc .1 S cr ub O ak F o r e s t T axon A pod emu s( i nde te rm ina te ) A pod emu sf l a v i co l l i s A pod emu ss y l va t i cu s R a t tu sr a t tu s S o r ex c o rona tu s S o rex m inu tu s N eomy sf od ien s C roc id u ra r u s su la T o ta l

3 6 2 3 0 0 0 0 0 3 2

L oc .2 S t r eam E dge

0 1 1 0 0 0 0 0 2

T rapp ing l o ca t ion 1 : L i ebäna , C an tab r ia , S pa in ; a bou t 3 k m W -N W o f P o t e s , a l t i tude 5 50 m , A ugu s t 2 7 -30 , 1 994 . T he s amp led h ab i ta t w a s s c rub o ak f o r e s t o n av e ry s t eep s l ope , w i th s h a l low t o p so i l a nd m any p a t che s o f e xpo sed b ed rock . G round c ove r w a s s pa r se . S hor t p a s tu r e w a s n ea rby . N oo pen w a t e rw a s n ea rby , a nd c ond i t ion s p r ev iou s t oa nd d u r ing t h i sc yc le o ft r app ing w e re v e ry d ry . T rapp ing l o ca t ion 2 : L i ebäna , C an tab r ia , S pa in ; a bou t 0 . 3 k m W o f T u r ieno , a l t i tude 3 40 m , S ep t embe r 2 10 , 1 994 . T he h ab i ta tw a s a l ong t h e R io D eva , w h ich a tt h i s p o in t i sa s h a l low s t r eam l i ned w i th g a l le ry d ec iduou sf o r e s t . S ome b ru shy g r ound c ove rw a s a va i lab le . N ea rby h ab i ta t s i n c lude h uman h ab i ta t ion sa nd s ho r tp a s tu r e s . N o mo le t u nne l sw e r e n o t ed , t h ough t h e t o p so i l a low s b u r row ing . O ne P y r enean d e sman ( Ga l emy s p y rena icu s ; I n sec t ivo ra , T a lp ida e , D e s m an ina e ) w a s o b se r ved i nt h e s t r eam h ab i ta t b u t n one w e r e t r apped . T rapp ing l o ca t ion 3 : L i ebäna , C an tab r ia , S pa in ; a bou t 1 k m No fA güebane s , a l t i tude 8 20 m , S ep t embe r 6 10 , 1 994 . T he t r ap sw e r e p l a ced i n ama tu r e o ak f o r e s t , w i th mode ra t e g r ound c ove r a nd s l ope . N o o pen w a t e r i sp r e sen t a tt h i s l o ca t ion ,a nd c ond i t ion sh ad b een d r y i ng ene ra l . T he t o p so i l a low s b u r row ing , b u t e n t rance s t o mo le t unne l s w e r e o n ly i n f r equen t ly e n coun te r ed .

L oc .3 Ma tu r e O ak F o r e s t

L oc .4 F o r e s t / Meadow / P a s tu r e

L oc .5 Meadow

T o ta l

0 8 1 0 2 2 0 0 1 3

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

0 2 4 0 0 0 0 1 8

1 6 3 1 4 7 1 5 2 4 7 14

S ome p a t che so fl ow b ru sh , s c ree s l ope ,a nd o pen s ho r tg r a s s a r ea l so p r e sen t . T he s ha l low t o p so i l i sg ene r a l ly u n su i t ed t o b u r row ing . T he w ea the r w a s d ry t om o i s t p r ev iou s t oa nd d u r ing t h i st i me o ft r app ing . T he r e su l t s o ft h e se t r app ing s e s s ion s a r e i l u s t ra t ive o ft h e c on t empo ra ry C an tab r ian m ic rof auna a sw e l la sp roce s se s o f m ic rof auna l mo r ta l i ty . F o remo s t i st h e h i gh p ropo r t ion o f A pod emu ss y l va t i cu sa nd A .f la v i co l l i s , t wo s p ec ie s n o rma l ly a s so c ia t ed w i th s i gn if ican t f o r e s tc ove r( T ab l e4 . 3 ) . E ven i n t h e se m ixed f o r e s t / c rop land s e t t ing s d om ina t ed b y t h e l a t t e r c ov e r t y pe , t h e se t wo s p ec ie s d om ina t ed a l t r apped t a xa . B a s ing e nv i ronmen ta l r e con s t ruc t ion u pon t r app ing r e su l t s a l one f o r a l t h e l o ca t ion s s amp led w ou ld l e ad t o t h e f a l se p i c tu r et h a t a l a r ea s h ad p r ima r i ly f o r e s tc ov e r , e ven t h ough t h i s v a r ied g r ea t ly a mong l o ca t ion s . M i c rof auna l s amp le s f r om P l e i s tocene S pan i sh s i te s ( Chap t e r s 5a nd 8 )h ave o n ly t i ny p r opo r t ion s o f A pod emu s . T h i s p r opo r t ion g ene ra l ly i n c r ea se s t h rough t h e A z i l ian p e r iod w h en t h e a me l io ra t ed c l ima t e o ft h e H o locene w a s r e a ched . M i c rof auna l d i ve r s i ty a sd i sc lo sed b y t r app ing i sa t ah i gh v a r iance f r om t h a t i n d i c a t ed f r om r a p to rp e l le ts t ud ie s( d i scu s s ed b e low ) . RAPTOR P ELLET S TUD IES O FI BER IAN F AUNA R oo s t ing l o ca t ion sw e r e s c ou t ed f o rp r inc ipa l ly i nt h e i mme d i a t e a r ea o ft h e s i t e o fE lJ uyo i t s e lf . T h i s a r ea w a s c ho sen f o r i t s p r ox im i ty t o t h e s i te , t h e r eby r emov ing a ny d i f f e r e n ce si nt e r ra in o ra l t i tude ( e xcep tf o rr e la t ive s e al e ve l )f r om a f f ec t ing ad i r ec t c ompa r i son w i th t h e P l e i s tocene f a una r e c ov e r ed . T he i mmed ia te e nv i ron s o ft h e s i t ea l so o f f e rr e la t i ve ly u ndeve loped t e r r i to ry . T he a u tho r h a s a l so o b se rved b a rn o w l s ( T y to a l ba ) , P e r eg r ine f a lcon ( Fa l co p e reg r inu s ) a nd e ve r -p r e sen t b uzza rd s ( Bu t eo b u t eo ) a c t ive ly h un t ing i n t h i sa r ea . R oo s t l o ca t ion s A , Ba nd C t h e r ef o r e i mmed ia te ly s u r round T rapp ing L oca t ion 5 a nd a r e n ea rby t o T rapp ing L oca t ion 4 .

T rapp ing l o ca t ion 4 : I g o l lo , C an tab r ia , S pa in ; a d ja cen t t o c ave o fE lJ uyo , a l t i tude 6 0 m , S ep tembe r2 3 -30 , 1 994 . T h i s h ab i ta ti si s o la t ed p ock e t so ff o r e s t( p r ima r i ly e uca lyp tu sa nd o ak ) s u r rounded b y t a l lp a s tu r e a nd m eadow s . A r ea s o fl ow b r u sh a r ea l so p r e sen t . As ma l l s ou r ce o fo pen w a t e ri sp r e s e n t . T he t o p so i l i sv e ry s u i tab le f o r b u r row ing , a nd t h e e n t r ance s t o m o le t u nne l s h ave b een f r equen t ly e ncoun te r ed ( t hough n one o ft h e se a n ima l s w e r e t r apped ) . Wea the r c on d i t ion sd u r ing t h ec yc le o ft r app ing w e r ee x t r eme ly w e t , w i th m ode ra t et oh eavy r a in e a ch d ay .

R oo s tl o ca t ion A : E lJ uyo Q ua r ry S i t e -2 S amp l ing o ccu r r ed o n J u ly 2 1 , 1 994 . T h i s n a r row f i s su r e ( F igu r e 4 . 2 ) w a s v e ry d i ff i cu l t t o a c ce s s . R oo s t ing b i rd s p e r ch h i gh a bove t h eb a se o ft h e f i s su r e , w h i ch i ss l oped a nd a low s p e l le t s t os p i l l e ven tua l ly o n to t h e s c r ee s l ope b e low .

T rapp ing l o ca t ion 5 : I g o l lo ,C an tab r ia , S pa in ;a bou t0 . 2k m No ft h e c ave o fE l J uyo , a l t i tud e 5 0 m , A p r i l 6 14 , 1 995 . T he a r ea t r apped l a y w i th in t h e c onf ine s o f al a rg e , l o ng a bandoned l i me s tone q ua r ry n ow c onver t ed t o m eadow .

6 4

S amp l ing o ft h e a c cumu la ted p e l le t s i n d ica ted t h a t t wo s p e c i e s o fr a p to r u t i l iz ed t h i s r oo s t s u cce s s ive ly . Al a rg e m a s s ( o ve r 1k g ) o fd i sagg rega t ing b ar n o w l p e l le t s w a s f ound t o u nde r lay af ew f r e sh ly d epo s i t ed , d i s t inc t ive p e l le t s o f t h e b uzza rd s( Bu t eo b u t eo )t h a th ad a ppa r en t ly j u s to ccup ied t h i s s i t e . A bou t 1 50 b a rn o w l p e l le t s a r e e s t ima ted t om ake u p t h i s s amp le . I n ta c t p e l le t s h ad t h e t y p ica l b l a ck , p o l i shed e x te r io ra nd s i z e( a ga in 4 . 5 c ml o ng a nd 2 . 5 c mw ide )t y p ica l t ot h i s s p e c i e s . C a r ew a st a ken t os e pa ra t e t h e se t wo s ou r ce s o fp e l le t s .

o fo pen t e r ra in w i th av a r ie ty o fc ove r t y pe s : h i gh p a s tu re , m eadow s , c r op land , a nd c oa s ta l s c rub a nd e xpo sed r o ck s . S ca t te r ed b u i ld ing s a nd i s o la ted p a t che s o ft r ee s a r e f o und t h roughou t t h i s a r ea . N o s i gn if i can t o pen f r e shwa te r l i e s w i th in t h i sz one . T he h i ghe s tp o in ti s4 9ma . s . l . S amp l ing o ccu r r ed o n S ep tembe r 1 5 , 1 994 . T h i s s i t e w a s u s ed f o r n e s t ing t h e p r ev iou s s umme r , s i nce s ome o f t h e p e l le t s w e r e s t i l l r e la t ive ly f r e sh . T h i s s amp le t h e r ef o r e i s t h eo n ly o ne d e r ived f r om b a rn o w l s f o rw h ich t h e s e a son o f a c cumu la t ion c an b e s t a t ed . E v idence t h a t b a rn o w l s o ccu p i ed t h i s r oo s t c ome s f r om t h e s i ze a nd a ppea rance o ft h e i n ta c tp e l le t s a nd t h e r e cove ry o ff r e sh f e a the r s i d en t if ied t o t h i s s p ec ie s . T he p e l le tt o ta l i sa bou t 1 0 . N o a dd i t iona l u se o ft h i sr oo s th a sy e tb een m ade b yt h eo w l s .

S ub sequen t r e tu rn t ot h i s s i te ( s amp led a ga in o n J u ly 4 a nd A ugu s t 1 6 , 1 995 ) y i e lded 4 3 f r e sh p e l le t s o f B u t eo b u t eo . T he se p e l le t sw e re t h e r ef o re a c cumu la t ed d u r ing t h i s s e a son , w h i le t h es e a son o fo ccupa t ion f o rt h eb a rn o w l sa tt h i s s ame r oo s t i su nknown . N o e v idence o fa dd i t iona l o w l h ab i ta t ion a tt h i s s i t ew a s d e tec t ed . T h i s s p ec ie s o fh awk i su b iqu i tou s i nt h e d ay t ime s ky a bove t h e E l J uyo c ave s i t e , a nd i st h e mo s t c ommon ly o b se rved d i ur na l r a p to r i n t h e C an tab r ian r e g ion .

T he f a i lu r e o f t r app ing a l one t o r e p re sen t f u l ly t h e s ma l l m amma l f a una o f C an tab r ia i sa ppa ren t ( Tab le 4 . 4 ) . T he t a xa M i cromy s m inu tu s , Mu s m u scu lu s , A r v i co la t e r re s t r i s , P i tymy s l u s i tan i cu s , T a lpa e u ropaea a nd C roc idu ra s ua v eo len s w e re n o t a cqu i r ed i n2 000 t r ap n i gh t s , a nd t h e t a xa R a t tu s r a t tu s a nd M ic ro tu s a g re s t i s a r e r e p r e sen ted b y o ne i n d iv idua l e ach . T he se t a xa a c coun t f o r 3 4 1 ( 3 6 .5% ) o ft h e t o ta l p r ey i t em st a ken b y t h e l o ca l r a p to r s . T he o n ly s p ec ie s o f s ma l l t e r re s t r ia l m amma l k nown t o b e p r e sen t i n C an tab r ian l ow land s t h a t w e re n o t t a k en b y t h e se r a p to r s s amp led a r e T a lpa o cc id en ta l i s , N eomy s a noma lu s , C l e th r ionomy sg la reo lu s , M i c ro tu s a rva l i s , A r v ico la s ap idu s a nd R a t tu s n o rveg icu s . B a rn o w l s p rove f a r mo r e e f f ec t ive a ts amp l ing t h e l o ca l C an tab r ian f a una t h an d o s t anda rd t r ap p i ng t e chn ique s .

A l l t h e p e l le t s amp le s f r om t h e E l J uyo q ua r ry c ome f r om r oo s t s i nn a t u ra l f i s su re s e xpo sed b yt h e l i me s tone q ua r ry ing t h a t h ad g one o n a tt h i s l o ca t ion ( s ee T rapp ing L oca t ion 5 ) . T he h ab i ta t s a va i lab le w i th in a 1k m r a d iu s f o rt h e se p r eda t o r s a r e p r ima r i ly a g r icu l tu ra l , w i th am ix o fh i gh p a s tu r e s a nd s ome c r op land . I s o la t ed p a t che s o f d ec iduou s f o r e s t , h edge row s , a nd b ru sh m ake u p l e s s t h an 2 0% o ft h e t o ta l a r ea . B a r e r o ck , e ven i nt h e q ua r ry , m ake s u p o n ly at i ny p e r cen tag e o ft h e o ve ra l l a r ea . H uman h ab i ta t ion s c on s i s t p r ima r i ly o ff a rm b u i ld ing s a nd w e l l spa ced h ome s . N o s i g n if i can t b od ie s o fw a te r e x i s t w i th in t h i s r a d iu s , e xcep t ing s ome s ma l l s p r ing s . A l t i tud ew i th in t h i s a r ea d oe s n o tc l imb a bove 1 2 1 m .

T y to a l ba a l so p roved t ob e amo r e e f f ec t ive s amp le r o ft h e l o ca l s ma l l f a una t h an t h ed i ur na l r a p to rB u teo b u t eo . D i rec t c ompa r i son o ft h et wo p r eda to r s a tR oo s t L oca t ion A i sp o s s i b le i n as e t t ing w he r e t h e e xa c t h un t ing r a nge w a s u t i l ized i ns u cce s s ion . Among s ma l lt e r r e s t r ia lm amma l s , Ta l ba l e f t b eh ind 1 2 p r ey s p ec ie s a so ppo sed t o 5f o rB .b u teo . Wh i le t o ta l s amp le MN I a tt h i s l o ca t ion w a s m uch h i ghe r f o r t h e f o rme rp r eda to rt h an t h e l a t t e r( 5 54 v s . 6 9 ) , t h e se p r ey i t em s c ome f r om mo r e c l o se ly a l ied n umbe r s o fp e l le t s ( c a . 1 50 v s . 4 3 ) . Wh i le b o th s p ec ie sh un t o n t h ew ing w i th in c ompa r a b le h un t ing r a d i i , B . b u teo h a s f ewe r s ma l l m amma l s p e c i e s f r om w h ich t oc hoo se : o n ly t h o se s p ec ie s a c t ive d u r ing t h e d ay t ime c an b e t a ken . F a r mo r e s ma l l m amma l s p ec ie s a r e p r ima r i ly n oc tur na l , t h u s h av ing a c t iv i ty p a t te rn s c o in c i d ing w i th T a l ba . B . b u t eo i sa l so ag r ea t e r d e s t ruc to r o f s ma l l b one s ( Chap te r2 ) , s o s ome p r ey i t em s m ay h ave b een r e nde r ed u n iden t if iab le a nd t h e r ef o r e r e duced a ppa r en t d i e t a ry b i od ive r s i ty .

R oo s tl o ca t ion B : E lJ uyo Q ua r ry S i t e -3 & 3 A T h i s t i ny c ave / la rge f i s su r e ( F igu r e 4 . 3 ) i sl o ca ted h i gh u p t h e q ua r ry w a l l a nd c on ta in s t wo s e pa ra te r o o s t ing s i t e s . S amp l ing o c cu r r ed o n J u ly 2 2 , 1 994 . A l l i n ta c t p e l le t s h ad t h e e x t e r io r b l a ck s h een t y p ica l o ft h o se e ge s t ed b y ab a rn o w l . S i t e 3w a st h e mo r e i n te r io r , w h i ch a lowed mo s to fi t s p e l le t st ot umb le f r om ah i gh l e dge i n to ad eep , b a r e ly a c ce s s i b le f i s su r e . O n ly o ne i n ta c t p e l le t w a s r e cove r ed , w h ich m ea su red 3 . 8 c m x2 . 1 c m , p l u s d i sagg rega t ing b one s a nd f u r . L e s st h an 2 0p e l le t sw e r eo r ig ina l ly p r e sen t . S i te 3 A c on s i s t s o f as e cu r e l e dge n ea r t h e e n t rance o ft h e c av e , f r om w h ich p e l le t s d rop t o as ma l l , s h e l t e r ed a r ea o f s c r ee . Many w ho le a nd p a r t ia l ly i n ta c t p e l le t s w e r e r e cov e r ed , at y p ica l o ne o fw h ich m ea su r ed 3 . 7 c m x2 . 5 c m . R e t u rn v i s i t st ot h i s l o ca t ion h ave y i e lded n o a dd i t iona l s i gn so f o w l o ccupa t ion . A l l i n d ica t ion s p o in t t ot h e se t wo a d ja cen t r oo s t sb e ing u s ed a tt h es ame t ime ,p e rhap sa l t er na t e ly .

OTHER M ICROFAUNAL S TUD IES O F NORTHERN S PAIN A l t i tud ina l g r ada t ion o ft h e s ma l l f a una o fC an tab r ian S pa in h a s b een s t ud ied f u r the r b y D e l ibe s d e C a s t ro ( 1 985 ) . T he s t udy a r ea i nt h i s c a se w a s a n 1 8 b y 3 0 k m r e g ion c ove r ing t h e n o r th a nd s ou th s l ope s o ft h e e a s t er n C an tab r ian m oun t a in s . T he se a l t i tud ina l g r ad ien t s w e r e d i v id ed i n to 2 50 m s e c t ion s , f r om b e low 5 00 m t o 1 700 m . At o ta l o f 115 s ma l l m amma l s w e r e t r apped f r om J u ly t h rough O c tobe r . T he n o r the rn s l ope h a sa nA t lan t ic c l ima te a nd y i e lded at o ta l o f2 4s p ec ie s ,w h i le t h es ou ther ns l ope h a s am o r ec on t inen -

R oo s tl o ca t ion C : C eme te ry S i t e 1 T h i s i s o la t ed s i t e i sl o ca t ed i nt h e u ppe r i n t e r io r o f al a rge m au so leum ,a bou t 4m a bove t h ef l oo ro n as t one l e dge . T he c eme t e ry i sl o ca t ed a bou t0 . 5 k m f r om t h e C an tab r ian c oa s t , o n t h e t o p o f a4 0 mh i l l , 4km N -NE o ft h e s i t e o fE lJ uyo . A 1k m h un t ing r a d iu s w ou ld e ncompa s s as i gn if i can t p o r t i on o fo pen s e a , s o t h i s z one m ay h ave b een s h if t ed s ou th w a rd s t oi n c r ea se t h e u sab le a r ea . T h i s e n t i r e z one c on s i s t s 6 5

F igure 4 . 2 : E lJ uyo Q ua rry S i te 2( Roo s t A ) .

F igure 4 . 3 : E l J uyo Q uar ry S i te 3a nd 3 A ( Roo s t B ) .

F igure 4 . 4 : E nv i rons o fE lJ uyo O ve r look ing N ea rby Quar ry .

6 6

T ab le 4 . 4 : Moder n Sma l l F auna o ft h eE lJ uyo A rea f r om T y to a l ba a nd B u teo b u teo P e l le t s . L oca t ion

Mur idae A pod emu ss pp .

A

A

B( 3 )

B( 3A )

C

( T y to a lba )

( Bu teo b u teo )

( T y to a l ba )

( T y to a l ba )

( T y to a l ba )

T o ta l

6 7 '

1

2

3 8

3

11

M . m inu tu s R .r a t tu s

8 4

0 0

1 0

4 5

1 0

1 4 9

M . mu scu /u s M a g re s t i s

7 2 7

0 6

0 4

1 1 7

0 5

8 5 9

M icro t idae A .t e r re s t r i s P .l u s i tan icu s

5 2 9 0

0 3 0

1 5 2

8 4 2

8 0

8 3 1 64

I nsec t ivora T .e uropaea

3

0

0

0

0

3

7 7 2 1 96

4 0 2 7

5 1 4

1 7 0 10

1 0 1 0

1 04 3 3 47

C .s ua veo len s

1

0

0

0

0

1

O ther C h i rop te ra Amph ib ia A ye s

2 2 1 6

0 0 1

0 0 0

0 0 7

0 0 0

2 2 2 4

I nver teb ra tes

X '

X

TOTAL

5 54

6 9

2 8

9 34

S .c orona tu s N .f od ien s C .r u s su la

X 3 4

2 49

1 MN I

c oun t sf o r mamma l sa nd a mph ib ians b a sed u pon s i ded m and ib le so r max i l lae ;A yes b a sed u pon s yn sac ra . d eno t e sp r e sence . No ti n c luded i nt o ta l s .

F igure 4 . 5 : A r ea lE x ten t( % )o fH ab i ta tT ypes i nt h e Moder nE nv i ron s o fE lJ uyo D e r ived f r om B a rn Ow l D ie t

% A rea

F o r e s t 1% D ry M eadow 2%

H um id M eadow /Wa t e r 6 1%

C ommen sa l 6 %

t a l c l ima t ew i th s ome Med i te r r an ean f o r e s ta r ea s a nd y i e ld ed

3 . } S pec ie sd i s t r ibu t ion r e la ted t on e i th e r : E l iomy s q ue r c inu s , R a t tu s r a t tu s , R . n o rv eg i cu s , M ic ro tu s n i va l i s , M c ab re rae , P i tymy s d uod ec im c o s ta tu s ,S o rex m inu tu s .

2 1 s pec ie s . F ou ra l t i tud ina lg roup sw e re i n d ica t ed : 1 . } S pec ie sd ec r ea s ing i na bundan ce a sa l t i tude i n c r ea se s : A pod emu s s pp . , Mu s m u scu lu s , R a t tu s r a t tu s , R . n or veg icu s , A r v i co la s ap id u s , C le th r ionomy s g l a reo lu s , P i tymy s d uod ec imco s ta tu s , C roc id u ra r u s su la .

Wh i le s ome o ft h e se s p ec ie sa r ef o und o n ly i nt h e Med i t e r ra n ean z one o fS pa in , m any o ft h e se s p e c ie s a r e d i s t r ibu t ed i n C an tab r ian S pa in . A . s y l va t icu s d i s t r ibu t ion d i r ec t ly c o r re l a t ed w i th l a t i t ud e , w i th ac l ea rp o s i t iv e g r ad ien tt ot h e n o r th o ft h i s p rov ince ( Mo r eno a nd B a rbo sa 1 992 : 172 173 ) . T h i s d i s t r ibu t ion a l so y i e ld s ap o s i t ive c o r r e la t ion w i th p r ec ip i ta t i on a nd w a s u n r e la t ed t o t empe ra tu r e . T h i s r e su l t i si n k eep ing w i th t h e o ve r a l l r a nge o ft h i s s p ec ie s a c ro s s m o s t o f t empe r a t e E u rope a nd i n to A s ia ( s ee C hap te r 3 ) . A .s y lva t i c u s d i s t r ibu t ion w a s a l so r e la ted t os t r uc tu ra l f e a tu r e s o ft h e v ege ta t ion , i . e . c ove r . P .p y rena i cu s h a s ar a nge c en te r ing u pon t h e P y r enee s , s o ap o s i t ive c o r r e la t ion w i th l a t i tude r ef l ec t s p ene t ra t ion n o r thwa rd b ack i n to i t s m a in r a nge . S . c o rona tu s a nd N .f od i en s f a l l i n to as im i la r c a t ego ry . E a ch h a st h e P y renee s /Can tab r ian C o rd i l le ra a st h e s ou the r ly l i m i t t o i t s m oder n r a nge , s o i n c r ea s ing d en s i ty c o r r e la t ing w i th mo r e n o r the r ly l a t i tud e i nt h i s r e g ion s imp ly m ean s am ove m en ti n to t h ep r ima ry r a nge . M . a g re s t i si n c r ea sed i nd en s i ty w i th i n c r ea s ing a l t i tud e , w h ich i st h e g en e ra l t r end f o r t h i s s p ec ie s i nS pa in ( Mo r eno a nd B a rbo sa 1 992 :173 ) . C . r u s s u la a l so i n c r ea sed i nd en s i ty w i th i n c r ea s ing a l t i tude , w h ich g oe s a ga in s t t h e g ene ra l t r end f o r t h e C ro c idu r ina e . I nt h i s c a se h oweve r , ap o s i t ive c o r r e la t ion w i th h uman h ab i ta t ion s my b er e f lec t ed i n s tead o f ap r ef e r ence f o rh i ghe r a l t i tud ina l z one s . S .m inu tu s p r ef e r s h i ghe r a l t i tude s ( > 1000 m ) i nt h e P y r enean p o r t ion o f i t s r a nge . S i nce a l s t udy s i t e s w e r e l owe r t h an t h i s l i m i t , i t s d i s t r ibu t ion w a s l e s s l i k e ly t o b e a f f ec t ed b y l a t i tude a nd a l t i tude a nd m o r e l i ke ly t ob e p o s i t i ve ly c o r r e la t ed w i th h um id i ty . T he a u tho r , t h ough d ea l ing w i th m uch s ma l le r t r app ing s amp le s , d i d n o t e ncoun t e r t h i s s p ec ie s u n t i l t r app ing a bove 8 00 m i nt h e C an tab r ian C o r d i l le r a . S im i la r ly , M ic ro tu s n i va l i s w ou ld b e e xpec ted t o s how ap o s i t ive c o r r e la t ion w i th a l t i tud e h ad t h i s s t udy b een a b le t o e ncompa s s h i ghe r m on tane z on e s a nd t h e i r m uch g r ea t e rp ropo r t ion o fs u i tab le r o cky s ub s t ra t e s . O th e rs p ec ie s s u ch a sR .r a t tu s a nd R .n o rveg i cu sa r ea l so f ound i nm ode rn C an tab r ian S pa in , b u t w i th t h e i r r e la t ive ly r e cen t i n t roduc t i on a sc ommen sa l s ,t h e i rd i s t r ibu t iona lh ab i t sa r eo fl i t t le u s e i nt h ei n te rp r e ta t ion o fP l e i s tocene f a una s .

2 . } S pec ie sp r ef e r r ing m ed ian a l t i t ud e s : G l i s g l i s , E l iomy s q ue r c inu s , P i tymy s l u s i tan i cu s , M i c ro tu sa g re s t i s . 3 . } S pec ie si n c r ea s ing i na bundan ce a sa l t i tude i n c r ea se s : P i tymy s s a v i i ( =P . p y rena i cu s ) , M ic ro tu s n i va l i s , a nd p o s s ib ly S o rex m inu tu s a nd N eomy s f od ien s ( b u ts amp le s i ze sf o rt h e s et wo t a xa w e r es ma l l ) . 4 . } R ema in ing s p ec ie s : T a lpa e u ropaea , 7 ' .c ae ca ( = To cc id en ta l i s ) , N eo m y s a noma lu s , G a l emy s p y rena icu s a nd A r v i co la t e r r e s t r i s . D e l ibe s d e C a s t ro ( 1 985 ) a l so n o t ed t h a t n on -a l t i tud ina l f a c t o r s m ay a f f ec tt h e se s p ec ie s ' d i s t r ibu t ion s . M i c ro tu s n i va l i s w a s a lway s a s soc ia t ed w i th i t s t y p ica l b i o tope ( r ocky s ub s t ra te s ) e ven a t l owe r a l t i tude s . C hange s i n a l t i tude m ay a f f ec t s p ec ie s d i s t r ibu t ion o n ly i n d i r ec t ly , i nt h a t c hang ing p l an t c ommun i t ie s o ve r a n a l t i tud ina l c l ine m ay b e t h e p rox ima t e c au se f o r as p ec ie s ' p r e sence o r a b sence . O ne w ou ld t h e ref o r e n o t e xpec t T .e u ropaea a nd T .c aeca ( = T o cc id en ta l i s )t ob e a f f ec t ed a sg r ea t ly b y c hange s i na l t i tude , g i ven t h e i r f o s so r ia l l i f e s ty le . S im i la r ly , s p ec ie s a s soc ia t ed w i th o pen w a te r ( Arv i co la t e r r e s t r i s a nd G a l emy s p y r ena i c u s )m ay b e l e s sa s soc ia ted w i th c hange s i na l t i tude . D e l ibe s d e C a s t ro ( 1 985 :55 ) a l so n o t e s t h a t t h e se r e su l t s a r e o n ly s e a sona l ly a pp l icab le , s i nce c oo le rc ond i t ion s c ou ld p u sh t h e a l t i tud ina l r a nge o fs ome s p ec i e s d ownwa rd s f o r p a r t o ft h e y ea r . L a t i tude a l so a f f ec t s t h e d i s t r ibu t ion p a t t e rn o fs ma l l m am m a l s , e v en o ve r r e la t ive ly s ma l l c hange s i nr a nge . Mo r eno a nd B a rbo sa ( 1 992 ) e xam ined at o ta l o f3 000 b a rn o w l p e l l e t s f r om 2 0 l o ca l i t ie s i nt h e p r ov ince o f H ue sca ( Spa in ) . A l t i tude s r a nged f r om 4 20 t o 9 8 1 m c ove r ing s i t e s i nt h e E b ro V a l ley , t h e P y r enean f o o th i l l s a nd t h e P y r enee s t h em s e lve s . At o ta lo f9 744 s ma l lm amma l sw e r er e cove r ed f r om t h e s e a sona l ly -poo led p e l le t s , s o y ea r ly c yc l ic c hange s i n d i s t r ibu t ion w e r e a ve raged o u t . At o ta l o f 1 7 ( 1 2 r o den t , 5 i n sec t ivo r e ) s p ec ie s w e r e f o und . T he se f e l l i n to t h r ee g roup ing s :

B rune t -Lecom te a nd D e l ibe s ( 1984 ) p r e sen t b a rn o w l p e l le t d a ta ( Tab le 4 . 5 ) f o r t h e D ue ro B a s in ( Ca s t i l la -L6on , S pa in ) w h ich w a s d i v id ed i n to mu l t ip le b i ome s ( mon tane , t r an s i t i ona l a nd f l a t land s ) . T he mo s t n o r the r ly ( t empe ra t e , m on t a ne ) z one c on ta in s m u l t ip le r oo s t s i t e s f r om t h e C an t ab r ian C o rd i l le ra , a l though m o s t o f t h e se a r e f r om t h e s ou the rn s l ope s o ft h i s r a nge . T he C an tab r ian s u b samp le c ame f r om 3 l o ca l i t ie s a nd y i e lded 5 684 i n d iv idua l p r ey i t em s . T he p eak s i nt h i s z one r e a ch o ve r 2 000 m , w i th r e gu la rp eak s o f 1 000 m . P r ec ip i ta t ion a ve rage s 6 00 mm /y ea r , w i th s ome y ea r sr e a ch ing 1 000 mm . A nnua lt empe ra tu r e a ve r age s< 10 ° C . L e s s t h an 3 0% o ft h i s s t ud ied z on e i su nd e r c u l t iva t ion , w i th m o s to ft h e r ema inde rc on s i s t ing o fp a s t u r e a nd d ec idu o u s f o r e s t ( mo s t ly P y r enean o ak Q ue rqu sp y rena i ca , b eech F agu s s y l va t i ca a nd S wee t c he s tnu t C a s tanea s a t /va ) , a nd s ome S co t s p i ne ( P inu s s y l ve s t r i s ) . Mo s t r oo s t s i t e s w e r e

1 . } S pec ie sd i s t r ibu t ion r e la t ed t ol a t i tude : A pod emu s s y lva t icu s , Mu s s p re tu s , P i tymy s p y r e na i cu s , C l e th r ionomy sg l a r eo lu s ,S o r ex c o rona tu s a nd N eomy sf od ien s . 2 . } S pec ie sd i s t r ibu t ion r e la t ed t oa l t i tude : M i c ro tu s a g re s t i s , S uncu s e t ru scu s a nd C roc id u ra r u s su la .

6 8

c hu rche s , a bandoned b u i ld ing s a nd n a tu ra l s he l te r s ( Brune t L ecom te a nd D e l ibe s 1 984 :214 -2 15 ) .

( Fa s t a nd H a rr i son 1 975 ; G ray son 1 981) . T hey , h oweve r , f a i l t o c on s ide r t h e r e la t ive s i ze s o ft h e p rey s pec ie s t a ken .

an tab r ian T ab le 4 . 5 : M ic rof auna f r om B a rn Ow l P e l le t s , C

r e f lec t t h e r e la t ive p ropo r t ion s o fw ood land a nd o pen f i e ld s f o rt h e b i rd s t ud ied a t as i te i nC a l if or n ia , U .S .A . T he h un t -

E vans a nd Em len ( 1947 ) f ound t h a t b a rn o w l d i e t d i d n o t C o rd i l le ra . T axon

i ng r ange o f t h i s b i rd w a s d i rec t ly o b se rved t o c ove r 1 65 a c res , 1 40 ( 84 .8% ) o fw h ich w e r e o pen g ra in /a lf a lf a f i e ld s

#o f

%T o ta l

L oc . 4

B ioma s s 0 . 1

T o ta l 0 . 4

3 0

1 4 .3

2 1 .3

Mu ss pp . R a t tu ss pp . M ic ro tu sa r va l i s

2 3 1 2 1

6 . 7 < 0 .1 5 . 5

6 . 0 0 . 2 8 . 3

M . a g re s t i s A rv ico la t e r re s t r i s

1 8 4

5 . 1 0 . 5

7 . 7 1 . 7

A .s ap idu s P i tymy s s pp .

5 2 9 7 1 9 4 2

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

1 . 3 2 1 .3 1 . 6 7 . 6 < 0 .1 < 0 .1

6 1 3 0 2 1 4 1 5 5

0 . 4 < 0 .1 3 3 .1 0 . 1 0 . 8 < 0 .1 0 . 3 0 . 1

0 . 3 < 0 .1 1 9 .7 0 . 1 2 . 2 < 0 .1 0 . 2 < 0 .1

E l iomy s q ue rc inu s A pod emu ss y lva t icu s

T a lpa s pp . S o rex s pp . S orex m inu tu s N eomy sf od ien s N . a noma lu s C roc idura s uaveo len s C .r u s su la C h i rop te ra A ye s R ep t i le s Amph ib ia A r th ropod s

a nd 2 5 ( 1 5 .2% ) w e re c o t tonwood /oak /w i l low f o re s t . T he m ic romamma l ian p r ey s pec ie s r e cove r ed f r om t h e p e l le t s a r e s hown i nT ab le 4 . 6 . Wh i le t h e n umbe r o fp r ey i t em s r e cov e r ed f r om t h e t wo m a jo r h ab i ta t t ype s d o n o t r ef lec t t h e i r r e la t ive a bundance s , t h e i r t o ta l b ioma s se s d o . P rey s pec ie s a s soc ia ted w i th o pen h ab i ta t s c on t r ibu ted 3 07 p rey i t em s ( 42 .9% ) a nd f o re s t -a s soc ia ted s pec ie s c on t r ibu ted 4 08 p r ey i t ems ( 57 .1 % ) t ot h e t o ta l d i e t . T h i s r e la t ion sh ip i si n ver ted when p r ey b i oma s s i st a ken i n to a ccoun t . F o re s t s pec ie s c on t r ibu ted 6 , 920 g ( 24 .7% ) t ot h e d i e t , w h i le o pen s pec ie s c on t r ibu ted 2 1 ,105 g ( 84 .8% ) . T h i s r e la t ion sh ip c l o se ly ma tche s t h e p ropo r t ion s o ft h e se t wo ma jo r b i o tope s i nt h e h un t ing r a nge . T h i s d i sc repancy b e tween n umbe r s o fp rey i t ems a nd a s soc i a t ed h ab i ta t a r ea i se xp la ined b y t h e d i f f e rence s i na ve rage p rey ma s s . T he l a rge s t n ume r ic c on t r ibu to r t od i e t ( 283 i n d iv idua l s ) i s Mu s ( hou se mou se ) , w h ich a ve rage s o n ly 1 5 g e ach . T he l a rge s t b i oma s s c on t r ibu to r t o d i e t ( 16 ,405 g ) i s T homomy s ( pocke t g ophe r ) , wh ich a ve rage s 8 5 ge ach . A s i ng le i n d iv idua l o fT homomy s i sa n a ve rage n igh t 's h un t ing t a ke . O ve r f i ve Mu s n eed b e t a ken t o ma tch t h is t o ta l , s o c on t inued h un t ing i sn ece s sa ry o n a ve rage t o f e ed t h e o w l a dequa t e ly ( Ma r t i 1 973b ) . P rov i s ion ing o f n es t l ing s w i l l o n ly i n c rea se t h i st o ta l . T he h igh n umbe r o f Mu s t a ken c an n o t b e e xp la ined t h rough g rea te r e a se o fc ap tu re , s i nce t h ey l i ve i n t h e mo r e d i ff icu l t ( c lo sed ) h un t ing h ab i ta t a nd o ne wou ld n o rma l ly e xpec t t h e i r u nde r rep resen ta t ion i nt h e o w l d i e t .

#o fL oc . = #o fl o ca l i t ie s o ft o ta l ( 3 3 )a tw h ich ag i ven t a xa w a s f ound . %T o ta l =% o fn ume r ica lt o ta lo fp r ey i t em s . %T o ta l =% o ft o ta lb ioma s so fp r ey i t ems . ( Fo l low ing B rune t -Lecom te a nd D e l ibe s 1 984 :216 )

T ab le 4 . 6 : T y to a l ba Mamma l D ie tf r om C a l if o rn ia , U .S .A .

PALEOENV IRON MENTAL I NTERPRETAT ION FRO M BARN OWL D IET P a leoenv i ronmen ta l i n te rp r e ta t ion f r om m ic rof auna l r ema in s f o l low s f r om t h e t a phonom ic a gency r e spon s ib le f o r t h e i r i n t roduc t ion i n to E l J uyo : t h e e ge s ted p e l le t s o f T y to a lba . R e sea rch b y p rev iou s a u tho r s a l so i n d ica te s t ha t t h e d i e t o f t h i s p reda to r i s ar e la t ive ly u nb ia sed s amp le o f t h e s ma l l mamma l ian p r ey w i th in a n a r ea a nd h ence ac l ea r p i c tu re o f

T axa

H ab i ta t P ref .

T o ta l P rey I t ems

A ve . Ma s s ( g )

B ioma s s i n D ie t( g )

t h e e nv i ronmen t a tt h e t ime o fd epo s i t ion . S ome e xcep t ion s d ue t op r ey b ehav io r d o o ccu r b u t a r e c o r rec tab le g iven t h e

Mu s T homomy s M icro tu s P eromy scu s

Wood s O pen O pen

2 83 1 93 10

1 5 8 5 4 0

4 , 245 1 6 ,405 4 , 400

Wood s

9 2

2 0

1 , 840

p re sen t s t a te o fi n f o rma t ion a bou tb a rn o w l b ehav io r . F o re s t s pec ie s t e nd t ob e u nde r rep r e sen ted , e spec ia l ly t ho se d we l l -

R e i throdon tomy s R a t tu s

Wood s Wood s Wood s

1 9 8 6

1 5 6 5 5

2 85 5 20 3 0

O pen

4

7 5

3 00

S orex L epu s

i n g a r bo r ea l ly s uch a s m embe r s o f G l i r idae (M u sca rd inu s a ve l lana r iu s , E l iomy s q ue rc inu s , e t c . ) . A n u ppe r l im i t o f a bou t 1 00 gf o r i nd iv idua l s t a ken a l so l im i t s t a xa i nt h e d i e t ( Mar t i 1 973b ) . T he l owe r s i ze l i m i t i n c lude s a r th ropod s . W i th in t h i s s i ze r ange , t h i s p r eda to rt a ke s p rey i nc l o se p ro p or t ion t ow ha t i sa va i lab le .

O pen

Wood s

S pec if i ca l ly , t h e p rey b ioma s s

t a ken f r om t h e h ab i ta t s a va i lab le w i th in t h e h un t ing r ange o f

T o ta l

P rey

4 08

( 5 7 .1 % )

3 07

( 4 2 .9% )

I t ems P rey

6 , 920 g

( 2 4 .7% )

2 1 , 105 g

( 7 5 .3% )

2 8 ,025 g

2 5 a .

( 1 5 .2% )

1 40 a .

( 8 4 .8% )

1 6 5 a .

7 15

B ioma s s

t h e o w l ' s d ay t ime r oo s t i sr ef lec t ive o f t h e i r r e la t ive a bun -

H ab i ta t A rea

d ance . O the r a u tho r s d i sag ree w i th t h i s a s se s smen t o fb ar n o w l d i e t a nd i t s r e p r e sen ta t ion o ft h e s u r round ing h ab i ta t p ropor t ion s

( Sou rce : E van sa nd Em len 1 947 :9 )

6 9

O the rr e sea r che r sh ave d e t ec t ed t h ec o r r e la t ion b e tween b a rn o w ld i e ta nd t h er e la t ive a moun t so fp r ey h ab i ta tt y pe sw i th in t h e i r h un t ing r a nge s . L ova r i e ta l . ( 1 976 ) s t ud ied b a rn o w l d i e t f r om s e ven r o o s t s i t e s i n ac en t ra l I t a l ian Med i t e r ranean e nv i ronmen t . P ropo r t ion s o ft r ee c ove rr a ng ed f r om 6 0% t o 2% , w i th v a ry ing p ropo r t ion s o f s c rub land , c r op land a nd g r a s s land m ak ing u pt h er ema ind e r s . Two o ft h e mo s ta bun d an t ly t a ken p r ey t a xa , A pod emu s s pp . a nd P i tymy s s a v i i s how , r e spec t ive ly , s i gn if i can tp o s i t ive a nd n ega t ive c o r r e la t i on s w i th f o r e s t c ove r . T he b i oma s s o fA pod emu s r a nged f r om 5 0 .9% t o2 7 .6% a c ro s s t h e r a nge f r om mo s t t o l e a s t f o re s tc ove r ,w h i le P .s a v i ir a ng ed i na n i n ve r se p a t t e rn f r om 8 . 0% t o5 6 .8% o ft h e b i oma s s t a k en . T he r e su l t s o ft h i s r e s e a rch a ga in s how t h a t t h e b i oma s se s o fp r ey s p ec ie s t a ken c an b e h i gh ly r e f lec t ive o ft h eh ab i ta tt y pe s i nt h e b ar no w l ' s h un t ing r a nge . O ne m u s t c on s ide r a dd i t iona l ly t h a t s ome s pec ie s h ave p r ima ry a nd s e conda ry h ab i ta t c ho ice s . T he se i n c lude mo s t i mpo r tan t ly A pod emu s s pp . ( A . s y l va t i cu s a nd A .f la v ico l l i s i nt h i s a r ea ) w h ich c an a l so b e f o und i no pen a r ea s , s oi n t e rp r e ta t ion m u s ti n c lude tt h i sv a r iab i l i ty .

T ab le 4 . 7 : Moder n T e r r e s t r ia l M ic romamma l P r ey o fB ar n Ow l sf r om t h eE nv i ron so fE lJ uyo . P r ey U n i t s '

MN I

B i oma s s

F o r e s t A pod emu ss pp . T o ta l

1 07

1 3 .2

1 . 0

1 . 1 1 . 0

50 1 . 0

1 07

1 .3 % 1 .3 %

C ommen s a l R .r a t tu s M m uscu lu s T o ta l

9 8

4 5 8

4 . 8% 0 . 8% 5 . 6%

H um id M e adow /Wa t e rE dg e A .t e r r es t r i s P .l u s i tan icu s S .c orona tu s T .e uropaea N .f od ien s T o ta l

7 5 1 34 9 9 3 3

9 . 3 1 6 .5 1 2 .2 0 . 4 0 . 4

5 . 0 1 . 0 0 . 5 5 . 0 0 . 75

3 75 1 34 4 9 .5 1 5 2 . 25

3 9 .7% 1 4 .2% 5 . 2% 1 . 6% 0 . 2% 6 0 .9 %

C .r us su la M . a g r es t i s M . m inu tu s C .s ua veo l en s T o ta l

3 10 4 8 1 3 1

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

0 . 5 1 . 0 0 . 5 0 . 5

1 55 4 8 6 . 5 0 5

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

T o ta l

8 10

1 00 .0%

9 45 .75

1 00 .0 %

D ry M eadow

R ecen t a na ly s i s o fo w l p e l le t s f r om t h e v i c in i ty o f E l J uyo ( b e low ) a l so i n d ica t e s as t rong c o r r e la t ion b e tween t h e b i o m a s s o fb ar n o w l p r ey a nd t h e a moun t s o fv ege ta t ion t y pe s p r e sen t . T ab le 4 . 7 i n d ica t e s t h e t o ta l t e r r e s t r ia l s ma l l m am m a lp r ey o fb a rn o w l s f r om p e l le tL o ca t ion s Aa nd B . T he se b a rn o w lr oo s t ing s i t e s a r e w i th in 1 50 m o ft h e E lJ uyo c ave e n t rance . T he s u r round ing a r ea i sp r ima r i ly f a rm land a nd p a s tu re ,w i th f ew h ou se sa nd l i t t le o p en w a t e r( F igu r e4 . 4 ) .

1P r ey U n i t sa r eb a sed u pon 1U n i t =o ne 2 0 gi n d iv idua la nd c on s ide r sj uv en i le p r ey c on t r ibu t ion t od i e t( Sou ther n 1 9 54 ) .

T h i s e nv i ronmen t ( w i th in a n a r b i t ra ry 1k m h un t ing r a d iu s ) i sr e f lec t ed b y t h e t o ta l b i oma s s ( a s a pp rox ima ted b y P r ey U n i t s ) d e r ived f r om e a ch h ab i ta tt y pe . T he s l igh t d ec iduou s f o r e s t c ove r ( p r ima r i ly i n t roduced e uca lyp tu s s t and s a nd h edge row s ) m ake s u p l e s s t h an 2 0% o ft o ta l c ove r . F o re s t s pec ie s ( Apod emu s ) m ake u p 1 .3% o fp r ey b i oma s s( F igu r e 4 . 5 ) . T he l ow b i oma s s o fc ommen sa l s p ec ie s ( 5 .6% ) i si n k eep ing w i th t h e s p a r se s t ruc tu r e s i nt h e a r ea . B o th h um id a nd d ry m eadow f a vo r ing s p ec ie s m ake u p t h e b u lk o fd i e t ( 8 3 .2% ) . T h i s r ef l ec t s c l o se ly t h e a r ea l e x t en t o f c l ea red a g r icu l tu ra l /pa s to ra l l a nd i nt h e a r ea . T he m ix o fh um id a nd d ry m eadow f a vo r ing s p ec ie s i se xp l icab le i nt e rm s o f mod e r n l a nd u se p r ac t ice s . T h i s c oa s ta l r e g ion i sq u i t e h um id a nd r e ce ive s o ve ro ne m e t e ro fr a in a nnua l ly . A r ea s o fd r ie r m eadow ( o rw i th v ege ta t ion m im ick ing t h a to fd ry m eadow ) i sc r ea ted b y f r equen t mow ing f o r h ay a nd g r az ing . O pen w a te rw i th in 1k m i sr e s t r ic t ed t os ma l l s p r ing s a nd i n te rm i t -

I n c ompa r i son , m ode rn b ar n o w l p r ey f r om s upe rf i c ia l d e p o s i t s a t Ama lda , a n u p land s i te i n t h e B a sque C oun ty ( Chap t e r8 )a r e q u i te d i f f e r en t f r om t h o se n ea rE lJ uyo . T h i s f a una ( P emän 1 990 :229 ) i sd om ina ted b yA pod emu s a nd h a s p r e sen t t h e a r bo r ea l G l i s g l i s a nd f o r e s t -dwe l l ing C l e th r ion o my s g la reo lu s . T he r e g ion s u r round ing Ama lda h a s f o r e s t c ove ra pp roa ch ing 4 0% . T h i s u p land s i t e a l so h a s f ew A r v i c o la . P i tymy sp y rena i cu s i sa bundan t , r e f lec t ing t h e h i ghe r a l t i tude o ft h i s s e t t ing . T he o pen m eadow c omponen t i sr e f l ec ted mo s t a bundan t ly b y C roc idura s p p . , S o rex s pp . , a nd P i tymy s l u s i tan icu s . T rue a l p ine s pec ie s ( G .p y rena i cu s a nd M i c ro tu sn i va l i s )a r ea b sen t . I n s umma ry , t h e m oder n m ic rof auna o f n o r ther n S pa in a s s amp led b yb ar no w l sp r ove st ob e ar e l iab le i n d ica to ro ft h e s ma l l t e r r e s t r ia l m amma l s p ec ie s p r e sen t a nd t h e i r r e la t ive b i oma s s . T he se r e la t ive b i oma s se s i nt ur na r e ar e l iab le i n d i c a to r o ft h e l o ca l b i o tope s w i th in a n o w l 's h un t ing r a nge , f a l l ing b road ly i n to f o r e s t c ove r , d r y g r a s s land , h um id g r a s s land , w e t land s , w a t e r e dge s , s c rub l a nd , h ea th land , a l p i ne m eadow , o pen r o ck / sc ree , a nd c ommen sa l s e t t ing s . When e xam in ed i nt h i s f r amewo rk ( Chap te r 5 ) , t h e U ppe r P a leo l i th ic s ma l l m amma l f a una o fE l J uyo w i l l b e u s ed t o i n t e rp r e t t h e L owe r Magda len ian p a leo env i ronmen t . T h i s a na ly s i s w i l l b e e x t ended t o o t he r s i t e s w he re p o s s ib le i n C hap t e r 8 . I n a dd i t ion , t h e c on tempo ra ry s ma l l m amma l f a una o f S pa in h ave b e en i l u s t ra t ed t o s how t h e l a rge c hange st h a th ave o ccu r r ed a f t e rt h eH o lo cene t r an s i t ion .

t e n t s ma l l i n unda t ed a r ea s . T he se a r e r ef l ec t ed b y t h e p r e s e nce o fA r v i co la t e r re s t r i s a nd N eomy sf od i en s . T he f o rme r s pec ie s c an a l so b e f o und a way f r om o pen w a t e r i nh um id m eadow s a nd i s af r equen t c omponen t o f t h e s ma l l f a una a l ong t h i sh um id c oa s ta ls t r ip . A l so s i gn if i can t i st h e l a ck o fs ome s p ec ie s . A rbo r ea l d o r m ice s pe c ie s s u ch a sG l i s g l i s a r e a b sen t . T he se c an b e e x p ec ted t o s how u p i nl ow n umbe r s i nb a rn o w l d i e t w he re t r ee c ove r i sm o r e e x ten s ive . C l e th r ionomy s g la reo lu s , a f o r e s t -dwe l l ing s p ec ie s , i sa l so a b sen t . T he l ow land s e t t ing i sr e f lec t ed b y t h e a b sence o fP i tymy sp y rena i cu s , G a l emy s p y rena icu sa nd M i c ro tu sn i va l i sw h ich a r ef o und a tg ene ra l ly h ighe ra l t i tude si nt h eC an tab r ian Moun ta in s( Chap t e r3 ) .

7 0

CHAPTER 5 EL J UYO MICROFAUNA AND PALEOENV IRON MENTAL I NTERPRETAT ION

s e pa rab le t o s p ec ie s l e ve l e ven w i th m e t r ica l a na ly s i s ( Chap te r 6 ) . MN I c oun t s f o r e a ch l e ve l w e r e t h e ref o r e d e r i ved f o rP i tymy s , w i th s epa ra t e MN I c oun t s f o r P . l u s i tan i c u sa nd P .p y rena i cu s . T he se l a t t e rc oun t sw e r en o ti n c luded i nl e ve l t o ta l s a nd c a lcu la t ion s d e r ived f r om t h em , s i nce t h i s p rocedu re w ou ld h ave c oun ted s ome i n d iv idua l s mo r e t h an o nce . P i tymy s g ene r ic c oun t s w e r e u s ed i nb i oma s s c a lcu la t i on s , w i th t h e r e la t ive p ropo r t ion s o ft h e s e t wo s p ec ie s e s t i m a ted f r om t h e i r i n d iv idua l c oun t s . I no ne c a se o n ly ( Leve l 4 R ) d i d t h e MN I f o r o ne o f t h e se s p ec i e s ( P . p y rena i cu s ) e xceed t h a td e r ived f o rP i tymy sa tl a rge . I nt h i s o ne c a se t h e MN I f o r P .p y r ena i cu s w a s u s ed i np l ace o f u nde s igna ted P i tymy s , s i nce n o P . l u s i tan i cu s w a s d e tec t ed f o r t h a t l e ve l . S i nce t h e se t wo s p ec ie sh ave s im i la ra du l tm a s s , e s t ima t e s o f t h e i rr e la t ive p r opo r t ion sd on o ta f f ec tb i oma s sc a lcu la t ion s .

T h i s c hap t e r d e ta i l s t h e s ma l le r m ic rof auna l r ema in s ( Roden t ia , I n sec t ivo ra a nd Mu s te la n i va l i s o f C a rn ivo ra ) f r om E l J u yo a nd t h e i n t e rp r e ta t ion o f p a leoenv i ronmen t f r om t h e i rr e la t ive p ropo r t ion s . T he r ema in so fl a rge rm ic ro f a una ( l a rg e r C a rn ivo ra a nd L agomo rpha ) a r e p r e sen t ed e l sewh e r e( Chap te r9 ) . T h i s a na ly t ica ls e pa ra t ion i sn o to n ly d ue t ot h e s i z e a nd t a xonom ic d i f f e r ence s o ft h e se s pec ie s , b u tt ot h e p r obab le manne r o fi n t rodu c t ion i n to s i te s o fe a ch s i z e c a tego ry . T he s ma l le r t a xa u sua l ly e n te r ed s i te s a st h e b one s i ne g e s t ed n oc tu rna l r a p to r p e l le t s . T he l a rge r t a xa u sua l ly e n t e r ed a s ar e su l to fh uman h un t ing f o rf ood o rf u r s o ra sl a rg e t e r r e s t r ia l c a rn ivo re p r ey r ema in s . A n e xcep t ion t o t h e se p a t t er n s a r e t h e r ema in s o f h edgehog ( E r inaceu s e u ropaeu s ) ,w h ich f a l l i n to n e i the rs i t ei n t roduc t ion c a t ego ry . T he se i n d iv idua l s mo s t l i ke ly e n t e r ed s i t e s o n t h e i r o wn , o w ing t ot h e i rc ommen sa l h ab i t s ( Chap t e r 3 ) . T he i n c lu s ion o f t h e i r r ema in s t h e ref o r e mo s t l i ke ly r e su l t s f r om n a tu ra l m o r ta l i ty . T h i s i n te rp re ta t ion i sr e inf o r ced b y t h e p a t te rn o f t h e i r s k e l e ta l r e cove ry a tE l J uyo ( Chap t e r 2 ) , w h ich m ake s t h e i ri n t rodu c t ion t ot h e c ave a sh uman p r ey l e s s l i ke ly . T he r ema in s a n a lyzed b e low a r e o n ly t h o s e w i th in t h e p r ey s i z e r a nge o fb a rn o w l s .

Among t h e g enu s M i c ro tu s , t wo o ft h e f o u r s p ec ie s p r e sen t a r e i d en t if iab le b y t h e i r , M , : M i c ro tu s n i va l i s a nd M . o economu s . N o s p ec ia l m an ipu la t ion w a s n ece s sa ry t oc a l c u la t ed t h e r e la t ive p ropor t ion s o ft h e se t wo t a xa . T he , M , o fM . a r va l i s a nd M . a g re s t i s i sn o t c on s i s t en t ly s e pa rab le mo rpho log ica l ly a nd mu s t b e g r ouped t o g e the r . T he 2M 2 o f M . a g re s t i s i ss e pa rab le f r om a l t h eM i cro tu ss p ec ie sp r e sen t a nd a low s e s t ima t ion o ft h e r e la t ive p r opo r t ion s o fM . a r v a l i sa nd M . a g re s t i sw i th in t h ec oun t so ft h e i rg rouped s am p l e . T he r e la t ive b i oma s s o fM a g re s t i s w a s e s t ima t ed b y s ub t ra c t ing i t s MN I f r om t h a t o ft h e c omb ined MN I f o r M . a g re s t i s a nd M a r va l i s . T h i s MN ! f o r M a g re s t i s w a s n o t i n c luded i no ve ra l l t o ta l s , s i nce t h i s w ou ld h ave r e su l t ed i n s ome i n d iv idua l sb e ing c oun t ed t w ice ( a sp e rc a lcu la t ion s f o r P i tymy s ) . M a g re s t i s a nd M a r va l i s a r e o f s im i la r a du l t m a s s , s o e s t ima t ion o f t h e i r r e la t ive p ropo r t ion s d oe s n o t a f f ec to ve ra l lb i oma s sc a lcu la t ion s .

E LJ UYO T he m ic rof auna l r ema in s f r om E l J uyo a r e l i s ted i n T ab le 5 . 1 . T he e l emen t s t a l l ied a r e t h e i d en t if i ed t e e th f o r e a ch t a xa , w i th n = 7 862 . T h i s t o ta l i n c lud e s t e e th ( n =3 76 ) i d en t if ied i na n e a r l ie r s t udy o fE lJ uyo b y C a s t ro B e rna rdez ( 1 986 ) . T he se t e e th h ave b een i d en t if i ed t os p ec ie s l e ve l i n m o s t c a s e s . Whe re t h i s w a s n o t p o s s ib le ( a s f o r t h e t e e th o f A pod emu s n o t b e ing s epa rab le i n to A .s y l va t i cu s o r Af la v i c o l l i s ) , m e t r i ca l a na ly s i s w a s a t temp t ed i no rd e r t oe s t ima t e t h e p r opo r t ion s o f t h e p o s s ib le s p e c ie s p r e sen t w i th in t h e g ene r ic t o t a l s . T h i s b i ome t r ica l a na ly s i s i sp r e sen ted i n C hap t e r 6 . I n mo s t c a se s p o s tc ran ia l m ic rof auna l r ema in s c ou ld n o tb ei d en t if ied t os p ec ie so re v en g enu s l e ve l , s ot h ey c ou ld n o tb eu sed i np a leoenv i ronmen ta l i n t e rp re ta t ion . T he p o s tc ran ia l r ema in s o fT a lpa h ave a l so b een a na lyzed m e t r i c a l ly t o s u pp lemen t s pec ie s leve l i d en t if ica t ion t o T . e u r opaea a nd T .o cc id en ta l i s( Chap t e r6 ) .

T he o t he r m o la r s o fM ic ro tu s a r e n o t s e pa rab le a mong t h i s g enu sa nd i ns ome c a se sn o tf r om P i tymy s . T he se mo la r sa r e g r ouped t o ge the ru nd e r M ic ro tu s lP i tymy s f o rt h e p u rpo se o f e s t ima t ing r e la t ive a bundance o ft h e se s ma l le r t a xa t oA rv i c o la a nd t ot h e o t he r s p ec ie s p r e sen t . T he se i d en t if ied t e e th a r e n o t i n c luded i n MN I t o ta l s , e xcep t f o r as i ngu la r m o la r f r om L eve l 1w ho se p r e sence i n d ica t e st h e e x i s tence w i thou t d oub le -coun t ing o fo ne i n d iv idua l M i c ro tu s /P i tymy s . T he se o t he r t a xa ( N eomy sf od ien s , N . a noma lu s , S o rex c o rona tu s , S . m inu tu s , A pod emu s , P l iomy s l e nk i , T a lpa a nd Mu s te la n i va l i s ) g ene ra l ly h ave a l l t h e i r m o la r s ( a nd o t he r t e e th i f p r e sen t ) i d en t if i ab le t ot h et a xonom ic l e ve l i n d ica ted . L a rge r t a xa ( E r inaceu s e u ropaeu s , C hy c to lagu s c un icu lu s , Mu s te la p u to r iu s a nd V u lpe s v u lpe s ) a r e e xc lud ed f r om t h i s a na ly s i s s i nce t h ey a r et o ol a rge i ng ene r a lt ob eb a rn o w lp r ey .

T he n umbe ro ft e e th i d en t if iab le t oe a ch t a xa a l so v a r ie s a nd i s af a c to r i np a leoenv i ronmen ta l i n t e rp r e ta t ion . E ach mo la r o f A r v ico la i si d en t if iab le t o t h a t g enu s . C onf u s ion w i th o t he rm i c ro t ine s pec ie s i si mpo s s ib le d ue t ot h e much g r ea te r s i z e o ft h i s g enu s : a du l t b ody m a s s a pp roa che s 3 00 g ,w h i le t h e o t he rm i c ro t ine s i nq ue s t ion a pp roa ch o n ly 4 0 -60 g . I n r aw n umbe r s o f i d en t if iab le mo la r s , A r v i co la i so ve r r ep r e s e n t ed i nc ompa r i son t oo t he rm ic ro t ine s p ec ie s . T h i s p rob l em i sc i r cumven ted b y u s ing t h e MN I ( M in imum N umbe r o fI n d iv idua l s ) f o r e ach t a xa , w he r e c oun t s a r e b a sed u pon t h em o s ta bundan t( s ided )t o o th f o re a ch s p ec ie s .

P r ey u n i t s ( f o l low ing S ou the rn 1 9 54 ) a r e b a sed u pon o ne u n i te qua l ing o ne 2 0 gi n d iv idua l . T h i s s omewha ta b s t ra c ted f i gu r e c on s ide r s t h e n o rma l p r opo r t ion o f j uven i le s ma l l m amma l s i nb a rn o w l d i e t ( wh ich n o rma l ly m ake s u p t h e b u lk ) . H ence ,m o s tr oden t s( M i c ro tu ss pp . a nd P i tymy ss pp . ) e qua l o ne p r ey u n i t . L a rge r t a xa ( Arv i co la c f . t e r re s t r i s ,

T h i s i d en t if i e r t oo th v a r ied a mong s p ec ie s . B o th P i tymy s p y rena icu s a nd P . l u s i tan i cu s a r e i d en t if iab le b y 'W . B o th , M , a nd , M ,a r e i d en t if iab le t ot h eg enu sP i tymy s ,b u ta r en o t

7 1

9,

. 0 . . 2

e .b

6 ° .z

2 x

, e z i u ,

0

—

0, 9 . 0, 00

—

— .

e l .

0

, C , r n

0 , e n

, . .

C h I r . . .

N

N C r -

2

. _ . , . . .

X

e_ R .z Le . e n

n i N

0 , i r z i

0, e n e t —

— • —

' 0

e n — — ,

. . .

—

N

e . 1

• •

e l

. ._ . . ,- . K r )

. . .. . .

e l

e l

( ( Z. ,

e l

: . : 7 .

( -

N

e n

I A

N

e n

. , * — •

0 ,

o o

o o

s

r 4

U ' .

nN e i i

_ ,". _. 0, . z , . . *. i . . „

N

i r ' l

, . . . , . 4

0 ,

e r e n c g, e 9 , ;, ) 1 91 a u i r J

N

e N

— .

7 ,—

C T

r ,

( 4 -

_2

7 , , 0 72 .r -

—. .

I .. .

e l e n .. .

> r s 1

e

1 " «, ^ .

e l e n

h i —

, . t : )

i f-.

0 ,

. . . . .

C 7. , . c :, e n

—

—

N

—

• •

e r

o o

z r . . o r 2 — .

4 .

. . 1

u p

F .

o o e n

N

— •

— , 0

0 ,

N

m N

un e n

F .

e n

—

c h

. t : i i n e r 0 0

.— , ,— , v-) e n

p ig

N

N i n e l , . k r ,

r . - ,— , *

1 - i = *

— ,

4 — .

L

e r 2

d t -

C, N

. . . r ,.., _.. 0

e r z

. 1 -2

C D • e •

" e r

. . .

• •i =. o i 1 ,0 . .

x

0 —

— , e n 0

o o e l . . . — . o o e r i

N e r v-, e l ,0

e l ( 4

—

—

z . • • e s 1

F .

• •

e n e s 1

e l

e l

O N

•

0

k e n e t

; en

0 1 v n

N

N

0, i l l

^

• •

*

—

0

— •

o 0

N

T N

0..4

•

C e,

e N

'.. "

t n

M r

. .. .

— ,

— ,

, c ) e r

k e n

e l

r -

T -

( V — • e r

r . .

0

' .

z e, 2

— z . . K . . .

. . . _.

. . „

•

r
30 ,600 B .P .

X I -X I I S t e r i le

o fE rm i t t ia a nd 1 . 5 km f r om t h e p r e sen t d ay s ho re . I t s e l e v a t ion i s1 30 m a . s . 1 . ,t hough t h ep rox im i ty o fl ow h i l l st ot h e c oa s t p lace e l eva t ion s o f6 00 m w i th in e a sy d i s tance . T he c ave i t se lf i s av e ry n a r row -mou thed ( a bou t 1m ) t unne l t h a t , w h i le d eep , d oe s n o t w iden b eyond 4 m e te r s i n i t s m a in c hambe r . I t s o ccupa t ion d u r ing t h e P a leo l i th ic a s as em i p e rmanen tc amp ing s i te i tt h u sr e s t r ic ted , w i th u se a st empo r a ry h un t ing o r e me rgency s he l te r b y s ma l l g roup s mo re l i ke ly . E xcava t ion w a sc a r r ied o u t i nt wo p ha se s , f r om 1 928 t o 1 936 b y T . d e A ranzad i a nd J .M . B a rand ia rän a nd 1 954 , 1 955 a nd 1 959 b y t h e l a t te r r e sea r che r . T he se w e re c a r r ied o u t i n t h e n a r row e n t ranceway a nd y i e lded ma te r ia l s f r om F ina l Magda len ian , A z i l ian a nd B ronze A ge o ccupa t ion s ( A l tuna 1 972 ) . Mar izu lo L eve l /Too l I ndu s t ry

1 2 ,750+ / -250 B .P . 9 , 460+ / 185 B .P . 1 3 ,350+ 1 -250 B .P .

V I Ia L owe r Magda len ian V I Ib L owe r Magda len ian V I Ic L owe r Magda len ian

v a ted a tL ez e tx ik i . U r t iaga L eve l /Too l I ndu s t ry

IA z i l ian I I A z i l ian I V A z i l ian

E ka in ( 4 3 °1 4 '10" N ,1 °2 4 '4 1" W) i sl o ca t ed 9 0 m a . s . l . a nd l e s st h an 7k m f r om t h e mode rn s ho re l ine . T he c ave i sa d ja c en tt o as ma l lt r ibu ta ry o ft h e R io U ro la , t h e v a l ley o fw h ich a l low s e a sy p a s sage b e tween t h e c oa s t a nd t h e r a p id ly r i s ing h i gh land s o ft h e i n te r io r . F oo th i l l s o ft h e C an tab r ian C o r d i l le ra r i se t oo ve r 8 00 m w i th in 3km s ou th o ft h e s i te . S i te p l a cemen t , a s w a s t yp ica l a mong U ppe r P a leo l i th ic C an tab r ian o ccupa t ion s , a l lowed a cce s s t o ar a nge o fh ab i ta t z one s w i th in a n e a sy w a lk ing r ad iu s . T he c ave i t s e lf b i f u r c a te s t o f o rm i n ne r g a l le r ie s , w i th o n ly a s ma l l i n hab i ted v e s t ibu le a r ea i ns ome p l ace s o n ly 2 m w ide . Wh i le s ma l l a nd l ow ly ing , t h e c ave e n t rance h a s s u i tab le s he lf a r ea s f o r r a p to r s t or oo s t , t hough t h e se wou ld h ave b een w i th in e a sy d i s tu rbance a c ce s st oh uman s . T he s i te i sa l so s i gn if i can t f o r i t sp a r ie ta l a r t , d ep ic t ing mo s t ly h o r se s b u t a l so ( i n d e scend i n g o rde r ) b i son , i b ex , r ed d ee r , b ea r , f i sh , a nd p o s s ib le r h i n oce ro s( A l tuna a nd Me r ino 1 984 ) .

A s soc ia ted ' 4C D a te s 5 , 285+ / -65

E xcava t ion s a tE ka in b egan i n 1 969 u nde rt h e d i r ec t ion o fJ . M . d e B a rand ia rän a nd c on t inued f r om 1 971-1975 . D i rec t i on f o rt h e f i na l t h ree c ampa ign s w a s b y J e sü s A l tuna . A l l s ed imen t s w e r e s c reened t h rough 2 mm m esh . T h i s s i z e i s

Ma r izu lo ( 4 3 °1 3 ' 10 " Na nd 1 °5 8 '10 " W) i sf ound 2 60 m a . s . l . a nd 1 2 km f r om t h e moder n c oa s t l ine . T h i s s i te l i e s o n t h e e dge o f t h e C an tab r ian f oo th i l l s , w i th h e igh t s o ve r 8 00 m l e s st han 2km t ot h es ou thea s t . T he R io L e iza ran , at r ibu ta ry o ft h e R io O r ia , i sj u s to ve r 2km t ot h ew e s t . Ma r izu lo i t s e lf i s av ery s ma l l c ave , t h e e n t rance o fw h ich c on s i s t s mu l t ip le n a r row i n te r l inked c hambe r s n o mo re t h an 2 m e te r s w ide .

s uf f ic ien t t or e t r ieve mo s t m ic rof auna l r ema in s , t h ough i s o l a ted t e e th o fs ma l le r s pec ie s ( s h r ew s i np a r t icu la r ) m ay b e l o s t . E rra l la L eve l /Too l I n du s t ry

A s soc ia ted ' 4C D a te s

IS t e r i le

I t s s ma l lnes s p e rhap s e xp la in s w hy i ta ppa r en t ly w a s n eve r

I F ina l Magda len ian

u sed a s ah ab i ta t ion s i te d u r ing t h e P a leo l i th ic , t hough i ti s p o s s ib le t h a t i tw a s u sed s po rad ica l ly a s e me rgency s he l te r .

I IF ina l Magda len ian

1 2 ,310+ / -190

I V S t e r i le

1 4 ,570+ / -300

V L owe r Magda len ian

1 5 ,800+ / -230 1 5 ,740+ / -240

A r t if ac t s f r om F ina l Me so l i th ic , N eo l i th ic a nd B ronze A ge l e ve l s h ave b een f ound t h e re .

T h i s s i te w a s f i r s t s amp led

1 6 ,200+ / -240 1 0 ,580+ / -270

u nde r t he d i r ec t ion o fJ .M . Me r ino i n 1 962 a nd e xcava ted f r om 1 962 t h rough 1 967 ( A l tuna 1 972 ) .

1 6 ,270+ / -240 E ka in L eve l /Too lI ndu s t ry

V I S t e r i le A ssoc ia ted ' 4C D a te s

V I ' S t e r i le

IS t e r i le 16

E r ra l la ( 43 °1 2 '32" N , 1 °3 0 '20 " W ) i sl o ca ted 2 30 m a . s . 1 . , a bou t 9km f r om t h e mode rn s ho r e l ine ( A l tuna 1 985a ) . T he c ave i sa d ja cen t t o t h e A lzo la ra s R ive r , at r ibu ta ry o f t h e U ro la . T he s i t e o f Ama lda l i e s a bou t 3 km d own s t ream . A l t i tude s o fo ve r 1 000 m a r e r e a ched l e s s t h an 3 km t ot h e s ou th i nt h e C an tab r ian f oo th i l l s . T h i s l o ca t ion i st hu s i n te r m ed ia te b e tween t h e c oa s ta l a nd h i gh land z one s a nd w i th in e a sy t r ave l o fb o th . T he c ave i t se lf h a st wo e n t rance s o pen i n gt o av e s t ibu le a r ea a pp rox ima te ly 6m b y 1 8 m . S u i tab i l i t y f o r r oo s t ing b y r ap to r s i si n d ica ted b y t h e v e ry l a rge m i c r of auna s amp le d e r ived f r om t h i s s i t e , f r om b o th s t e r i le a nd a r chaeo log ica l l e ve ls . E r ra l la h a s y i e lded L owe r a nd F ina l Magda len ian o ccupa t ion s . E xcava t ion s a tt h i s s i te a r e v ery r e cen ta nd w e r ed i rec ted b yJ .A l tuna i n1 976 a nd 1 977 .

T hough t h e e a r ly c o l lec t ion s f r om t h e s i te h ave s uf f e r ed g rea t c u ra t iona l l o s se s ( A l tuna a nd S t raus 1 976 ) , h uge d en s i t i e s o fa r t if ac t s a nd l i v ing d eb r i s w e re d epo s i ted t h e re d u r ing t h e S o lu t rean a nd L owe r Magda len ian p e r iod s . T he a r t i s t ic s t y le s r e co rded o n o s seou s i mp lemen t s t h e re h ave b een u sed ( Conkey 1 978 , 1 980 ) t os ugge s t t h a t A l tam i ra f unc t ioned a s as ea sona l a gg r ega t ion s i te f o r d i spe r sed g roup s l i v ing i nt h e l o ca l r eg ion . W i th i t s a r two rk a nd l a rge s i ze , A l tam i ra m ay h ave f unc t ioned a s am ee t ing p l ace f o rg roup r i tua l a c t iv i t ie s d u r ing t h eL owe r Magda len ian p e r iod . E lC a s t i l lo L eve l /Too l I ndu s t ry

A s soc ia ted ' 4C D a te s

1 a Moder n l b S t a lagm i te 2N eo l i th ic

CANTABR IA A l tam ira L eve l /Too lI n du s t ry

A s soc ia ted ' 4C D a te s

2L owe r Magda len ian

1 5 ,500+ / -700 1 4 ,520+4260 1 4 ,480+ / -250

3S o lu t rean

1 8 ,540+ / -320

3S t a lagm i te 4A z i l ian 5 a -b S t a lagm i te 6 Magda len ian A

A l tam i ra i sl o ca ted 1 53 m a . s . l . n ea r t h e t o p o f as ma l l h i l l o ve r look ing t h ec oa s ta l l ow land s . T he moder n s ho re l ine i s4 k m d i s tan t . T he c ave i t s e lf o r ig ina l ly o pened o n to a l a rge v e s t ibu le , w i th s ubs tan t ia l t u nne l s l e ad ing i nwa rd s t o mu l t i p l e c hambe r s i n c lud ing t h e a r ea s o f p a in t ing s a nd e ng rav i n g s . B o th S o lu t rean a nd L owe r Magda len ian l e ve l s h ave b een e xcava t ed t h e re , t h ough t h e m a jo r a r two rk e p i sode s h ave b een a s s igned a Magda len ian a ge ( Mo re R oman i l lo e t a l . 1 996 ;V a l lada se ta l . 1 992 ) . T he e xcava t ion s a t A l tam i ra a r e a mong t h e e a r l ie s t P a leo l i t h ic i n ve s t iga t ions i nC an tab r ian S pa in ( A l tuna a nd S t rau s 1 976 ; F r eeman 1 987 , 1 988 ; S t rau s 1 976 -77 , 1 992 ) . T h i s c ave w a sd i scove red o n ly i n1 868 a nd w a s e xcava ted f o rt h e f i r s t t ime i n 1 876 b y Ma r ce l ino S anz d e S au tuo la , al o ca l a ma teu rn a tu r a l s c ien t i s t . H e c on t inued e xcava t ion s i n 1 879 i nt h e v e s t ibu le a r ea , a tw h ich t ime ( a cco rd ing p e rhap s t o l e gend ) h i s y oung d augh t e r d i scove r ed j u s t i n s ide t h e v e s t i b u le t h e w o r ld renowned b i ch rome c e i l ing c ompo s i t ion o f b i son a nd d ee r . T hese f i nd s w e re m e t w i th s kep t ic ism o ve r t h e a s soc ia t ion o ft h e p a r ie ta l a r t w i th p r im i t ive a r t if ac t s b y t h e r e ign ing F r ench p reh i s to r ian s o ft h e d ay , n o tab ly G . d e

7 Magda len ian A /B 8 Magda len ian B 9S t e r i le 1 0 S o lu t rean 1 1 2a -b A u r ignac ian A 1 3 A u r ignac ian A /B 1 4a 14c A u r ignac ian B 1 5 A u r ignac ian B /G 1 6a R oden t s I 1 6b -c A u r ignac ian G 1 6d R oden t s H 1 7 S t e r i le 1 8a -b A u r ignac ian D

1 0 ,310+ / 120 1 2 ,390+ / 130 1 6 ,850+4220

3 9 ,000+ / -2000 ( a ve . o f3 ) ( B i schof fe ta l . 1 991)

1 9 S t a lagm i te 2 0a -b Mou s te r ian A u ppe r& lowe r 2 1 S t e r i le 2 2a -c Mou s te r ian B 8 9 ,000+ 11 ,000 / -10 ,000 2 3a -c S t e r i le U se r ie s i s och ron d a te o n f l ows tone ( B i schof fe ta l . 1 99 1) 2 4 A cheu lean

Mo r t i l le ta nd E .C a r ta i lha c . I n 1 881 t h e c ave w a s v i s i ted b y E . H a r lé a tt h e i r b ehe s t a nd d ec la r ed a p robab le f o rge ry .

2 5a -b ,26 B e low A cheu lean

O ve rt ime a nd i nt h e f a ce o fg row ing e v idence a to the rn ew ly d i scove r ed p a r ie ta l a r t c ave s i t e s , C a r ta i lhac l a te r r e can ted

E l C a s t i l lo i sl o ca ted 1 75 m a . s . l . o n as t eep , c on ica l h i l l w h ich i t se lf r i se s t o3 75 m . T he s i te o ve r look s t h e R io P a s

( 1 902 ) . S ome o the rr e sea r che r s e xcava ted i nt h e c ave i nt h e

v a l ley , w h ich d e lve s s ou thwa rd i n to t h e f oo th i l l s o f t h e

i n te rven ing y ea r s b ef o re mo r e s ub s tan t ia l wo rk b y H . A l c a lde d e lR io i n1 902 1903 ,d igg ing i nt h e v e s t ibu le a r ea a nd

C an tab r ian C o rd i l le ra . I t s l o ca t ion a lows a cce ss t o t h e u p l a nd a nd c oa s ta l l ow land z one s , a nd e a sy c ommun ica t ion t o

i n t he

b i ch rome s .

b o th a l ong t h er i ve rv a l ley . T he c ave n ow c on s i s t s o f ah igh -

A l tam i ra w a s s ub sequen t ly e xcava t ed b y H . O be rma ie r i n 1 924 a nd 1 925 . H i s w o rk w a s c a r r ied o u t p r ima r i ly i nt h e

c e i l inged o pen a br ia tt h e e n t rance a nd ac avern l e ad ing b ack i n to t h e h i l l many h und r ed s o f m e te r s , w i th s pec tacu la r

v e s t ibu le a r ea . Mode rn e xcava t ion s u nde r t h e o ve ra l l d i rec t i on o fJ .G onza lez E chega ray t o ok p l ace i n 1 980 a nd 1 98 1

c hambe r s a nd f l ow s tone f o rma t ion s . T h i s a nd o the r c aver n s i nt h e C a s t i l lo h i l l h ave mu l t ip le i n s tances o fp a r ie ta l a r t l o c a ted ‚ t h roughou t , e ven d eep w i th in i ns ome c a se s . L oca ted i nt h e s ame h i l l r ad ia l ly a l ong t h e s ame a pp rox ima te a l t i tude a r e t h e c ave s o fL ago , L a s C h imenea s ( Gonza lez E chega ray

" G r ea t

H a l l"

u nder nea th

t h e

b i son

i n t h e " k i tchen m idden " a r ea o f t h e v e s t ibu le ( F reeman 1 988 ) .

17

1 953 , 1 974 ) , L aF l echa , P a s iega a nd L a s Moneda s . T h ree o f t h e se ( Ch imenea s , P a s iega a nd Moneda s ) a r e ma jo r p a r ie ta l a r ts i te s a dd i t iona lt oE lC a s t i l lo i t s e lf . L aF l echa h a sy i e lded Mous te r ian a r t if ac t s . I n a dd i t ion t o i t s a r t , E l C a s t i l lo h a s y i e lded l e ve l s c ove r ing v i r tua l ly t h e e n t i re P a leo l i th ic s e q uence o f We s te rn E u rope : A cheu lean , Mou s te r ian , A u r ig n a c ian , S o lu t rean , Magda len ian , A z i l ian a nd a l so N eo l i th ic l e ve l sh ave b een d i scove red t h e r e . J u s t a s t h e e xcava t ion s a tA l tam i ra a r e a n e a r ly e xamp le o f s e r iou s P a leo l i th ic i n ve s t iga t ion i n C an tab r ian S pa in , t h e e xcava t ion s o fE l C a s t i l lo c ha r t m uch o fi t s s ub sequen t h i s t o ry i n to t h e mode rn p e r iod ( Cab r e ra 1 984 ; S t rau s 1 992 ) . H . A lca lde d e l R io d i scove red t h e p a r ie ta l a r t o fE l C a s t i l lo i n 1 903 , a nd s ubsequen t ly d ug i nt h e u ppe r l e ve l s o ft h e ma s s i ve d epo s i t s a tt h e o pen a br i a tt h e f r on t o ft h e s i te .

T he

n ew ly f o rmed I n s t i tu t d e P a leon to log ie H uma ine a nd i t s p a t r on , P r ince A lbe r t Io f Monaco , s pon so r ed m a s s ive e xcava t i ons i n t h e 1 8 m d eep s equence i nt h e f r on t o f t h e c ave . T he se w e re c a r r ied o u t f r om 1 9 10 t h rough 1 9 14 u nde r t h e d i rec t ion o fH . O be rma ie r a nd i n c luded t h e p a r t ic ipa t ion o f s uch n o tab le s a s P . We rne r t a nd t h e C onde d e l aV ega d e l S e l la . U nf o r tuna te ly ( a nd f a r t o o f r equen t ly ) t h e se r e su l t s w e re o n ly s pa r se ly p ub l i shed a nd p oo r ly c u ra ted , w i th much l o s s o f i rep laceab le i n f o rma t ion ( Cab re ra 1 984 ) . E xcava t i on o f r emnan t s t ra t ig raphy c on t inue s a t E l C a s t i l lo c u r r en t ly , u nde r t h e d i r ec t ion o fF .B e rna ldo d e Q u i rö s a nd V . C ab re ra . Cueva Mor in L eve l /Too lI ndu s t ry

t h e ma in v e s t ibu le a r ea , w h ich i t se lf r un s a bou t 8m b y 8 m , t hough t h ec ave c on t inue s i nwa rd s f r om t h ere . E l P endo L eve l /Too l I ndu s t ry IA z i l ian I F ina l Magda len ian I ab F ina l Magda len ian I cd F ina l Magda len ian I I L a te A u r igna c ian I V L a te A u r ignac ian V ,V a G rave t t ian V b , V I E vo lved A u r ignac ian V I I A u r igna c ian I V I I I C ha te lpe r ron ian V I T Ia ,b A r cha ic A u r ignac ian V I I Ic S t e r i le V I I Id Mou s t e r ian I X Mou s te r ian X I -X I I I Mou s te r ian X IV Mou s te r ian XV Mou s te r ian XV I a c Mou s te r ian

A s soc ia ted ' 4C D a t e s E l P endo i sl o ca ted l e s s t h an 3 km i n land f r om E l J uyo , o n t h e c oa s ta l l ow land s a bou t 1 0 km f r om t h e mod e rn c oa s t . T he n ea re s ts i gn if ican tr i ve ri st he R io P a s , a bou t 3k m t ot h e w e s t . T he c ave i t s e lf i se xpan s ive , w i th t h e ma in v e s t ibu le c hambe r a bou t 3 0 mh igh , u p t o2 0 mw ide a nd m any t ime s t ha t a moun t d eep . T h i s s i te h a s y i e lded al o ng s t ra t if ied s e q uence w i th Mou s te r ian , A u r igna c ian , C ha te lpe r ron ian , G rave t t ian , F i na l Magda len ian a nd A z i l ian l e ve l s . I ta l so c on ta ins p a r ie ta la r t i nt h e f o rm o fb i rd e ng rav ing s , ar a r i ty i n P a leo l i th ic a r t .

1A z i l ian 2F ina l Magda len ian 3S o lu t rean 4G rave t t ian 5G rave t t ian /Au r ignac ian I ( Sup . )2 0 ,710+4350 6A u r ignac ian I( 7 /6 ) 3 2 ,4 15+ / -90 1 7 Au r ignac ian I 2 9 ,515+ 1 -865 2 8 ,055+4 1 ,535 8A u r ignac ian 0( 8a ) 2 8 ,435+ 1 -556 2 8 ,155+ 1 -757 2 8 ,515+ / -1 ,324 9A u r ignac ian 0 1 0C ha te lpe r ron ian

e x ten s ive ly s amp led i n 1 966 . R e -excava t ion o ft h e c ave i n 1 968 a nd 1 969 w a s u nde rt h e d i rec t ion o fJ .G onza lez E che g a ray a nd L .G . F reeman ( 1971 , 1 973 , 1 978 ) . T he ' 4C d a te s f o r l e ve l s d ownwa rd f r om 8 a a r e s u spec ted b y t h ee xcava to r s t ob e u n re l iab ly y oung . T he se e xcava t ions c ove r ed mo s t o f

T he e xcava t ion h i s to ry o fE l P endo i sl o ng a nd c omp lex . E l P endo w a s f i r s t e xcava ted i n 1 878 b y M . S anz d e S au tuo la a nd a ga in i n 1 880 w i th J .V i lanova y P i e ra . S ome e ng rav i ng s i nt h e s i t ew e re d e sc r ibed b y H . A lca lde d e l R io i n1 907 . J . C a rba l lo v i s i ted t h e s i te i n 1 908 a ccompan ied b y H . O be rma ie r , a nd i tw a s a l so r e v i s i ted t h e A lca lde d e l R io a nd

2 8 ,6 10+4577 3 6 ,950+46 ,777

1 2 Mou s te r ian C ueva Mo r in i sl o ca ted l e s s t h an 2k m s ou th o ft h e s ou the rn

t h eA bbé B reu i l i nt h a ts ame y ea r . J .C a rba l lo r e tur n ed t ot h e

s ho re o ft h e B ay o fS an tande r . I t s p rox im i ty t ot h e g l a c ia l a ge s ho re l ine w ou ld h ave b een s i gn if ican t ly g r ea te r ( ove r 1 5

c ave i n 1 910 a ccompan ied b y W . B ea t ty a nd c a r r ied o u t s ome a r chaeo log ica l t e s t ing o ft h e d epo s i t s . R . S ha l lc ra s , 0 .

km ) , s i nce t h i s s ha l low b ay w ou ld h ave b een d ry l a nd . S i te s e t t ing i sw i th in t h e n a r row c oa s ta l l ow land s . A t C ueva

C end re ro , G ömez L lueca a nd C . P e rez B u s tamen t e a l so s am -

Mo r in ,n ea r ly t h ee n t i re l a te rP a leo l i th ic c u l tu ra l s t ra t ig raph ic s equence h a s b een r e cove red : Mou s t e r ian , C ha t e lpe r ron ian ,

o ft h e se f i nd ing s ( Cend re ro 1 9 15 ) .

Au r ignac ian , G rave t t ian , F ina l Magda len ian , a nd A z i l ian

i n c luded v i s i t s b y t h e Ame r ican S choo l o f P reh i s to r ic R e -

p l ed t h e d epo s i t s i n 1 9 14 a nd s ub sequen t ly p ub l i shed s ome J .C a rba l lo c a r r ied o u t

s eve ra l b r ief c ampa ign s i nt h e 1 920 ' s , 3 0 ' s a nd 4 0 ' s . T he se

l e ve l sh ave b een d i scove red . C ha te lpe r ron ian L eve l 1 0 i st h e

s ea rch i n 1 930 .

mo s t

t h e d i rec t ion o fJ . Ma r t inez S an ta O la l la ( t hen d i r ec to ro ft h e S em ina r io d e H i s to r ia P r im i t iva d e l aU n ive r s idad d e Ma -

s e cu re

i d en t if ica t ion

o f t h i s

t o o l

c omp lex

f o r

C an tab r ian S pa in . T he e a r l ie r e xcava t ion s w e re c a r r ied o u t t h e re f r om 1 9 12 t o1 9 19 i na l ter na t ing s e a son s b yJ .C a rba l lo a nd b y t h e C onde d e l aV ega d e l S e l la . B r ief t e s t s w e re made a tt h e s i te i n 1 962 a nd t h e w i tne s s s e c t ion w a s mo re

Ma jo r e xcava t ion s w e re u nde r taken u nde r

d r id ) w i th A . C he in ie r , B . S aez Ma r t in , V . R u is A rg i ld s , J . G onza lez E chega ray a nd C . B o rden i n1 953 i nc o l labo ra t ion w i th t h e D ipu tac iön P rov inc ia l a nd A . G a r c ia L o r enzo . D i 18

r e c t ion i n 1 954 a nd 1 955 w a s b y R u i s A rg i le s i nc o l labo ra t i on w i th F rance sca M ine lono . E xcava t ion i n1 956 a nd 1 957 w a s d i rec t ed b y Ma r t inez S an ta -O la l la a nd A nd re L e ro i G ou rhan . P ub l ica t ion o ft h e r e su l t s f r om t h e e xcava t ion s o f t h e 1 950 's w a s d e layed many y ea r s ( Gonza lez E chega ray

L a s C a lda s ( 43 °2 0 '10" N , 2°1 4 '05" W ) i sl o ca ted 1 60 m a . s . l . n ea r t h e c ap i ta l o f A s tu r ia s ( Ov iedo ) . T h i s s i te i sl o ca ted a bou t2 5 km f r om t h e modem s ho r e l ine a nd s im i la r ly d i s tan t f r om t h e C an tab r ian C o rd i l le ra t ot h e s ou th . N ea rby f oo th i l l s d o n o t r e ach a bove 4 50 m , b u tw i th in 8km o ft h e s i te p eak s

1 980 ) . C u r r en t s amp l ing o f t h e r emnan t d epo s i t s i sb e ing u nde r taken u nde rt he d i rec t ion o fR . Mon te sa nd V . C ab re ra .

u p t o 1 712 m ( P ico G amona l ) a r e r e ached . T he R io N a lon r uns j u s t s ou th t h e e n t rance s , t h e v a l ley o fw h ich a l low s e a sy

R a scaf io L eve l /Too lI ndu s t ry

c ommun ica t ion t o t h e i n te r io r a nd c oa s t . T he c ave i t se lf c on s i s t s o ft wo s epa ra te , n a r row c hambe r s r unn ing p a ra l le l i n to ah i l l s ide . E ach o ft h e se c ave s i sn o mo re t h an 5m w ide

A s soc ia t ed ' 4C D a te s

a nd u sua l ly much n a r rowe r , a nd e x tend s a bou t 3 5 mi nwa rd s 1A z i l ian

1 0 ,486+ / -90 1 0 ,558+ / -244

( Co rchön 1 981) .

2U ppe r Magda len ian

1 2 ,896+ / 137 1 2 ,282+ / 164

L a s C a lda s w a s f i r s t m apped i n 1 958 b y l o ca l s pe leo log is t s

3L owe r Magda len ian

1 5 , 173+ 1 160 1 5 ,988+ / 193 1 6 ,433+ / 131

4L owe r Magda len ian 5I n i t ia l Magda len ian 6N ea r ly S t e r i le 7N ea r ly S t e r i le

2 7 ,240+950 -810

8N ea r ly S t e r i le 9A ur igna c ian?

R . Q ue ro l a nd J .C a r ra sco . L oo t ing p i t s s ub sequen t ly d e s t royed al a rge p o r t ion o fv e s t ibu le d epo s i t s , w h ich h ad b een r educed p rev iou s ly b y n a tu ra l w a te r f l ow . T he i n ta c t d epo s i t s w e re f i r s t e xcava ted f r om 1 971 t o 1 973 u nde r t h e d i rec t i on o fM . S .C o r chön , w h ich i n c luded a tl e a s t s ome s c reen i n g o fs i te s ed imen t s . T he f a una l r ema in s f r om L a s C a lda s w e re h eav i ly f r agmen ted a nd i n c luded v e ry f ew m ic rof auna l s pec ie s .

1 0S t e r i le T he s i te o fR a sca t io ( 43 °1 7 '38 "N , 3°4 144 " W ) i sl o ca ted 2 75 ma . s . l . a t ad i s tance f r om t h e modem s ho re l ine o fo ve r 2 5 k m . T he c ave i sn ea r ly a d jacen tt ot h e R io M ie ra a nd l i es o n t h e e dge o ft h e f oo th i l l s c l imb ing t ot h e C an tab r ian C o rd i l l e ra af ew k i lome te r s t ot h e s ou th . R a scaf io i st h u s s i tua ted t oa low a c ce s st ot h e a d ja cen t h igh land z one a nd e a sy c om m un ica t ion w i th t h e c oa s t a l ong t h e r i ve r v a l ley . U t i l iza t ion o ft h is s i te o ccu r red p o s s ib ly d u r ing t h e A u r ignac ian p e r iod , a nd c l ea r ly d u r ing t h e L owe r Magda len ian , U ppe r Mag d a len ian a nd A z i l ian p e r iod s . T h is l o ca t ion w a s p robab ly c ho sen f o r u se a s as pec ia l ized i b ex ( Capra p y rena ica ) h un t ing c amp , a s i si n d ica ted b y t h e a bundance o ft h i s s pe c i e s ' r ema in s i nt h e s i te d epo s i t s . T he f i r s te xcava t ion o ft h is s i te o ccu r r ed i n 1 912 u nde r t h e d i r ec t ion o f J .R . G ömez R iaf io a nd J .C a rba l lo i nc on junc t ion w i th t h e C om i s iön d e I n ves t iga c ione s P a leon to lög ica s yP r eh is tö r ica s . S ome e xca v a t ion c on t inued i ns ub sequen t y ea r s u nde r t h e d i rec t ion o f P .S i e r ra a nd G ömez R iaf io . S ome i n ve s t iga t ion s w e re a l so c a r r ied o u t t h e re b y H ugo O be rma ie r a nd t h e C onde d e l a V ega d e lS e l la i n1 921 , a nd a ga in i n1 923 b y O be rma ie rw i th J .a nd F .R ia f io . Modem e xcava t ion o fR a scaf io w a i ted u n t i l 1 974 a nd w a s c onduc ted u nde r t h e d i rec t ion o fJ .G onza lez E chega ray a nd I .B a rand ia rän Mae s tu ( 1981) . ASTUR IAS

L a R iera L eve l /Too l I ndu s t ry 2 6 U ppe r Mag . /Az i l ian 2 4 U ppe r Magda len ian 2 1-23 Uppe r Magda len ian 2 0 L owe r Magda len ian

1 9 /20 L owe r Magda len ian 1 9 L owe r Magda len ian

1 8 L owe r Magda len ian 1 7 S o lu t rean 1 6 S o lu t rean 1 5 S o lu t rean 1 4 S o lu t r ean 1 3 S o lu t rean 1 S o lu t rean 1 0 S o lu t rean 9S o lu t rean 8S o lu t rean

A s soc ia ted ' 4C D a te s

1 0 ,890+ / -430 1 0 ,340+ / -560 1 2 ,620+ / -300 9 , 090+ / -570 1 2 ,360+ / -670 1 7 , 160+ / -440 1 5 ,230+ / -300 1 5 ,520+ 1 -350 1 6 ,420+ / -430 1 6 ,900+ / -200 1 7 ,070+ / -230 1 8 ,200+ / -6 10 1 5 ,600+ / -570 1 7 ,225+ / -350 1 5 ,690+ / -310

1 9 ,820+ / -390 1 5 ,860+ / -330 2 0 ,690+ / -810

L a sC a lda s L eve l /Too lI ndus t ry

7S o lu t rean 5S o lu t rean 4S o lu t rean

2 0 ,970+ / -620

1P os tg la c ia l 2M idd le Magda len ian

L a R ie ra i sl o ca ted a t4 3 °2 5 '31" N a nd a bou t 4°5 2 ' W . T he

3 -6 F ina l S o lu t rean 7 9 U ppe rS o lu t rean

c ave e n t rance i s3 0 m a . s . 1 . , a nd i s1 . 5 km f r om t h e R io

1 0 13 S t e r i le 1 4 M idd le S o lu t rean -B

L a R ie ra i s s i tua ted i n t h e c oa s ta l l ow land s o f A s tu r ia s , t hough f oo th i l l s o fo ve r6 00 m a l t i tude c an b e r e ached j u s t 4

1 5 S t e r i le 1 6 -18 M idd le S o lu t rean -A

km t ot h e s ou th . I t s l o ca t ion made f o r e a sy c oa s ta l r e sou rce e xp lo i ta t ion , e ven d u r ing t ime s o f l owe r ed s e a l e ve l . T he c ave i t se lf h a s a n e n t rance a bou t 5m w ide a nd am a in v e s t i -

1 9 S t er i le

C a lab res t ot h e w e s ta nd 1 . 75 km s ou th o ft h e mode rn s ho re .

19

b u le e x tend ing o ve r 1 5 m i nwa rd s . E xcava t ion s h e re b egan w i th t h e C onde d e l aV ega d e l S e l la i n1 917 t o 1 9 18 . Af ew

i nt he f o rm o f mu l t ip le g eome t r ic s a l ong t h e m a in c ave rn , e ng rav ing s i n as ma l l s i de c hambe r , a nd t h e R e ind ee r P ane l

e xamp le s o fp a r ie ta l a r t w e re d i scove red i nt h e m a in c ham b e r b y M . Ma l lo , M . P e rez a nd J .A . A lva r ez i n 1 968 . I n 1 969 , G . C la rk t e s ted t h e d epo s i t s i nf r on to ft h e c ave , w h ich

c on ta in ing mu l t ip le s upe r impo sed e ng rav ing s a nd p o ly c h rome i mage s o fr e indee ra nd h o r se d eep w i th in t h e c aver n .

w a s r epea ted i n1 972 b y J .M . G ömez -Tabane ra a nd M . P e r e zw i th t wo s ma l l p i t s . T he c ave a l so s uf f e r ed m any d ep r e d a t ion s b y l o o te r s i nt h e many y ea r s s i nce i t s o r ig ina l e xca -

t r ance a nd a tt h e f oo t o ft h e R e indee r P ane l .

v a t ion a nd t h e mo s t r e cen t p ro jec t d i rec ted b y L . G . S t rau s a nd G . C la rk f r om 1 976 t h rough 1 979 ( S t rau s a nd C la rk 1 986 ; S t rau s e ta l . 1 981) . T h i s s i te h a s y i e lded l e ve l s f r om t h e S o lu t r ean , L owe r a nd U ppe r Magda len ian , a nd A z i l ian p e r iod s a sw e l la ss ome A s tu r ian p e r iod c onche ro d epo s i t s . C ue to d el aM ina

aA s tu r ian bU ppe r Magda len ian cM idd le Magda len ian dM idd le Magda len ian eL owe r Magda len ian fU ppe rS o lu t r ean gS t e r i le hM idd le S o lu t rean IS t e r i le jU ppe rA u r ignac ian T he s i te o fC ue to d e l aM ina i sf ound i na no the r v e s t ibu le o pen ing o ft h e s ame c ave rn c omp lex t h a t h ou se s t h e s i te o f L a R ie ra . C ue to d e l aM ina i s3 5 ma . s . 1 . , a nd a bou t 1 . 5 km f r om t h e R io C a lab re s t ot h e w e s t a nd 1 . 75 k m s ou th o ft h e moder n s ho r e l ine . I ti ss i tua ted i nt h e n a r row c oa s ta l l ow l a nd s o fA s t u r ia s , w i th f oo th i l l s o fo ve r6 00 m a l t i tude j u s t4 km t ot h e s ou th . I t s l o ca t ion m ade f o r e a sy c oa s ta l r e sou r ce e xp lo i ta t ion , e ven d u r ing t ime s o f l owe red s e a l e ve l . T he o ccup ied v e s t ibu le a r ea i sa bou t 1 0 mb y 1 0m . A u r igna c ian , S o lu t rean , L owe r a nd U ppe r Magda len ian , a nd A s t u r ian d e p o s i t s h ave b een d i scove red h e re . T h i s s i te w a s f i r s td i scov e r ed a nd e xcava ted b y C onde d e l aV ega d e l S e l la i n 1 9 14 a nd t e s ted b yF .J o rdä i n1 953 ( Ca s t ro B e rna rdez 1 986 ) .

l a /b U ppe r Magda len ian

A l l t h e l e ve l s

e xcava ted t o d a te h ave b een U ppe r Magda len ian , t h e o l de s t d a ted o ccu r rence o ft h i st o o lc omp lex i nC an tab r ian S pa in . T he f i r s t t e s t s o fi t s a r chaeo log ica l d epo s i t s w e r e c a r r ied o u t b y M . A . G a r c ia G u inea i n1 970 , w ho d ue t ot h e a b sence o f a n t le rh a rpoon s a nd t h e p r e sence o fo t he rt ype s a s s igned t h i s s i te t ot h e L owe r Magda len ian . Mo r e e x t en s ive e xcava t ion s i n 1 972 a nd 1 974 -75 u nde rt h e d i rec t ion o fJ .A . Mou re R o man i l lo u nea r thed h a rpoon s , l e ad ing t or e a s s ignmen t t ot h e U ppe r Magda len ian ( Mou re R oman i l lo 1 975 ; Mou r e R o man i l lo a nd C ano H e r re ra 1 976 , 1 978 , 1 979 ) .

L eve l /Too lI ndu s t ry

T i to B us t i l lo L eve l /Too l I n du s t ry

O ccupa t ion d eb r i s h a s b een d e tec ted n ea r t h e o l d m a in e n -

A s soc ia ted ' 4C D a t e s 1 4 ,220+4 180 1 4 ,250+ / -300

MOUSTER IAN ENV IRONMENTS V e ry l i t t le c an b e s a id a bou t v a ry ing Mou s te r ian c l ima te f r om t h e m ic rof auna l d a ta , s i nce n o l a rge s amp le s h ave y e t b een r e cover ed . O n ly t h ree s i te s ( Ama lda , Mo r in a nd L eze tx ik i ) h ave y i e lded m ic rof auna f r om t h e i r Mou s te r ian l e ve l s , a nd e a ch h a s as ma l l s amp le s i z e ( Tab le 8 . 1 , e nd o f c hap te r ) . I n a dd i t ion , t h e Mou s te r ian p e r iod c ove r s al o ng p e r iod f r om p e rhap s 1 30 ,000 B .P . t o4 0 ,000 B .P . a nd s pan s much o fO xygen I s o tope S t ages 5 , 4a nd 3 :f r om f u l l i n te r g l ac ia l t o i n te r s tad ia l c ond i t ion s . Mo s t p a leoenv i ronmen ta l i n f orma t ion f r om t h is p e r iod c omes f r om c o r r e la t ion w i th s e a l e ve l c hanges ,p a lyno logy a nd mac romamma l ian f a una . Ama lda L eve l s X a nd V I I I ( S te r i le o ve r ly ing Mou s te r ian ) a nd V I I ( Mou s te r ian ) h ave y i e lded s ma l l m ic rof auna l s am p l e s ( Pemän 1 990 ) . T he m ic ro t ine s p r e sen t i nL eve l Xa r e M icro tu s a rva l i s ( n =1 33 ) , M . o economu s ( n =4 4 ) , P i tymy s p y rena icu s ( n =2 4 ) , M . n i va l i s ( n =2 )a nd A r v i co la c f . t e r r e s t r i s ( n =1 ) , w h ich i n d ica t e ac o ld , t r ee le s s e nv i ronmen t w i th o pen a r ea s o fg ra s s land a nd p e rhap s s ome m o i s t h ea th . L eve l s V I I I a nd V I I a r e l e s s i n f o rma t ive , w i th s ma l l a moun t s o fA rv ico la c f . t e r re s t r i s , P l iomy s l e nk i a nd S o rex a raneu s p re sen t . C ueva Mo r in h a s y i e lded o n ly as i ng le e l emen t o f A r v ico la a nd o ne a dd i t iona l u n iden t if i ed r o den t r ema in f r om i t s Mou s te r ian L eve l 1 5 ( A l tuna 1 978 ) . Mou s te r ian a ge m i c rof auna f r om L eze tx ik i L eve l s V I I , V i a nd V b i ss im i la r ly u n inf o rma t ive , w i th M icro tu s a g re s t i s /ar va l i s , A r v i co la , t h e e x t inc t m ic ro t ine P l iomy s l e nk i , a nd S o r ex p re sen t i ns ma l l n umbe r s .

1 5 ,180+ / -300 1 5 ,400+4300

EARLY UPPER P ALEOL ITH IC ENV IRON MENTS

1 3 ,520+4300

Wh i le a l a rge r b ody o f m ic rof auna l d a ta e x i s t s f r om t h i s s hor te r p e r iod , g ene ra l ly s ma l l s amp le s i z e s l i m i t i n t e rp re ta -

1 3 ,870+4300

t i on t ot h e p re sence /ab sence o fi nd ica to r s p ec ie s ( Tab le 8 . 2 ) .

l b -c U ppe r Magda len ian 1 cU ppe r Magda len ian

T he c ave o fT i to B u s t i l lo i sl o ca ted n ea r ly a ts e a l e ve l , 5 00 m f r om t h e mou th o ft h e S e l la R ive r a nd 7 50 m f r om t h e mod er nc oa s t l ine . T i to B u s t i l lo w a sd i scove red o n ly i n1 968 , t h i s d e lay c au sed b y t h e s ea l ing o f i t s o r ig ina l e n t rance . T he c ave i t se lf i sq u i te ma s s ive , e x tend ing i n o ne m a in t unne l t e n s o fm e te r st a l l f o rh und red s o fm e te r s i nwa rd , w i th s pec t a cu la r l ime s tone f o rma t ion s a nd mu l t ip le s i de c ave rn s . T he s i te o fL a L lo se ta w a s d i scove red a ta no the r e n t rance t ot h i s

M ic rof auna f r om E a r ly U ppe r P a leo l i th i c l e ve l s h a s b een r epor ted f r om Ama lda ( Pemän 1 984 ) , A i tzb i ta r t e ( A l tuna 1 972 ) , E l C a s t i l lo ( Cab r e ra 1 984 ) , E ka in ( Zaba la 1 984 ) , L eze tx ik i ( A l tuna 1 972 ) a nd C ueva Mo r in ( A l tuna 1 978 ) . C u l tu re s t ra t ig r aph ic u n i t s c ove r A u r igna c ian a nd C ha t e lpe r r on ian ( 40 ,000 t o 2 7 ,000 B .P .) a nd G rave t t ian /Uppe r P e r igo rd ian ( 27 ,000 t o2 0 ,500 B .P . ) l e ve l s . O xygen I s o tope S t age s s pan l a t e 3a nd e a r ly 2 :f r om i n t e r s tad ia l c ond i t ion st o g l ac ia l max imum .

c aver nc omp lex . T i to B u s t i l lo a l so h a s e x ten s ive p a r ie ta l a r t 1 20

T he m i c rof auna l s equence f r om L ez e tx ik i ( Leve l s I Vc , I Vb , I Va , I Ib a nd I Ia ) s pan s P ro toau r igna c ian t oU ppe r A u r ig n a c ian d epo s i t s ( A l tuna 1 972 ) . S amp le s i z e i ss ma l l , b u tt h e m i c rof auna i n d ica te s a mo s t ly t r ee le s s e nv i ronmen t b a sed u pon t h e d om inance o fm ic ro t ine s . T he u n ce r ta in ly a t t r ib u t ed L eve l s X a ( Cha te lpe r ron ian ) a nd I Xb ( Au r igna c ian ) o f E ka in a r e s im i la r ly d om ina ted b y m ic ro t ine s , i n d ica t ing t r ee le s s c ond i t ion s ( Zaba la 1 984 ) . O n ly A r v i co la a nd T a lpa e u ropa ea ( b o th r e la t ive ly l a rge a nd e a s i ly i d en t if iab le ) h ave b een r e po r t ed f r om t h e S upe rf i c ia l a nd B a sa lA u r igna c ian G L eve l s f r om E l C a s t i l lo ( Cab r e ra 1 984 ) . T he se t wo t a xa h ave a l so b e en i d en t if i ed f r om t h eG rav e t t ian l e ve l so fC ueva Mo r in ( A l t una 1 978 ) . Mo re c omp le t e i n f o rma t ion i s a va i lab le f r om t h e A u r ig n a c ian l e v e l o f A i tzb i ta r t e ( A l tuna 1 972 ) . T r ee le s s c ond i t i on s a r e i n d ica ted h e r e a sw e l l , w i th m ic ro t ine s d om ina t ing t h e s amp l e : A . t e r re s t r i s ( n =5 9 ) , M i c ro tu s o e conomu s ( n= 4 8 ) ,M .a g r e s t i s /a rva l i s( n =3 2 ) , P i tymy ss p p . ( n —4 )a nd M . n i va l i s ( n = 1 ) h ave b een r e cove r ed a l ong w i th S o rex a raneu s a nd T a lpa e u ropaea . H um id c ond i t ion s a r e a l so i n d ica t ed b yt h ep re sence o fA . t e r r e s t r i s a nd M . o economu s , w i th s ome b a re r o cky a r ea s a tl e a s tw i th in ap r eda to r ' s h un t i n gr a d iu sa si n d ica ted b yM . n i va l i s . O pen , g r a s sy a r ea s a r e i n d ica t ed b y M . a g re s t i s /a rva l i s a nd P i tymy s . T he h ab i ta t i n d ica t ed b yt h i sm ic rof auna i ss im i la rt oE lJ uyo L eve l4 / 4S , t h ough A i t zb i ta r te e x i s t s i n as l igh t ly h i ghe r a l t i tud ina l z one ( 2 20 m a . s . 1 . )w i th n ea rby f o o th i l l sc l imb ing t oo ve r5 00 m . C ue to d e l aM ina h a s y i e lded a l a rg e m ic rof auna l s amp le f r om i t s A u r ignac ian L eve l 7( Ca s t ro B e rna rdez 1 986 ) . S ome p a r t ia lt r ee c ove ri si n d ica ted f o rt h et i me o fd epo s i t ion o f t h i s a s semb lage b y t h e p r e sence o f a r bo r ea l E l iomy s q ue rc inu s a nd f o re s t -f avo r ing A pod emu s s y l va t icu s a nd A . f l a v ico l l i s . O the r f a una i n d ica t e s i gn if i can t ly c o lde r a nd m o re o p en c ond i t ion s t h an t o day . S i gn if ican t d ownwa rd s h if to ft h e a l p ine z one i si n d ica t ed b yt h e p r e sence o fG a l e m y s p y r ena i cu s a nd M i c ro tu s n i va l i s . S i gn if i can t o pen g r a s sy a r ea s a r e i n d ica ted b y M . a g re s t i s /a r va l i s a nd o pen w a te r /hum id m eadow b y A rv i co la , S o rex c o rona tu s , S . m inu tu sa nd M . o economu s . L eve l V( Evo lved P e r igo rd ian ) f r om Ama lda ( P emän 1 990 ) h a s b een r a d ioca rbon d a ted t o at i me o f i n t e r s tad ia l o cean l e ve l ( 2 7 ,000+ 1 1 ,000 a nd 2 7 ,000+ / 1 , 100 B .P . ) . T he s ma l l m ic rof auna l s amp le i si n d ica t ive o fo pen , g r a s sy c ond i t ion s w i th M i c ro tu sa g re s t i s( n 6 9 )a nd M . a r va l i s( n = 12 )p r e d om ina t ing o ve r M . o economu s ( n =3 9 ) , M . n i va l i s ( n =1 ) a nd A . c f .t e r re s t r i s ( n =2 2 ) . L eve l V I ( P e r igo rd ian V ) f r om Ama lda h a sb een r ad ioca rbon d a t ed t oa r ound t h e l a s tg l a c ia l m ax imum ( 1 7 ,880+ / -390 a nd 1 9 ,000+ 1 -340 B .P . ) a nd c o in c i de sw i th m o r e e x t reme e nv i ronmen t . T he s ma l l m ic rof au n a l s amp l e i se ven mo r e d om ina ted b y M a r va l i s ( n =9 6 ) a nd i si n d ica t ive o ft h e s p r ead o fd r ie r , o pen g r a s s e nv i ron m en t . S OLUTREAN ENV IRONMENTS O f s ho r t e r d u ra t ion t h an t h e Mou s t e r ian a nd E a r ly U ppe r P a leo l i th ic , t h e S o lu t r ean i n C an tab r ian S pa in ( 2 0 ,500 t o 1 6 ,500 B .P . ) s pann ing r ough ly t h e l a s tg l a c ia lm ax imum ( c a . 2 2 ,000 18 ,000 B .P . ) i s u nde r r ep r e sen ted b y m ic rof auna l d a ta . S ome i sr epor ted f r om t h e s i t e s o f Ama lda ( P emän

1 990 ) , A i tzb i ta r t e ( A l tuna 1 972 ) , L a s C a lda s ( Co r chön 1 981) , E rm i t t ia ( A l tuna 1 972 ) , Mo r in ( A l tuna 1 978 ) a nd L a R ie ra ( A l tuna 1 986 ) . N o l a rge s amp le sh ave b een r e cove red , s o p a leoenv i ronmen ta l i n f e r ence s mu s t b e d r awn b y t h e p r e sence /ab sence o fi n d ica to rs p ec ie s . T he s ma l l a moun t s o f m ic rof auna r e cov e r ed m ay a l so b e ap a r t ia lr e su l to fa n e nv i r onmen t u nf avo rab le t o b a rn o w l s ( v e ry c o ld ) , w h ich r e d uced t h e p opu la t ion o f t h i s a c cumu la t ing s p ec ie s i n C an tab r ian S pa in . C o ld a nd t r ee le s s c ond i t ion s a r e i n d ica ted b y t h e s amp le f r om t h eU ppe rS o lu t r ean l e ve l s( IVb a nd I Va ) f r om Ama lda ( Pemän 1 990 ) . T he se h ave b een r a d ioca rbon d a t ed t oa f t e r t h e g l a c ia l m ax imum : d a te s o f 1 7 ,580+ / -440 , 1 6 ,200+ / -300 a nd 1 6 ,090+ / -240 B .P . i n d ica t e s ome o ve r lap w i th t h e s u c c eed ing L owe r Magda len ian p e r iod . T he m ic rof auna l s am p l e s f r om t h e se t wo l e ve l s a r e d om ina ted b y o pen m eadow m ic ro t ine s ( M i c ro tu s a r va l i s , M . a g re s t i s , P i tymy s p y rena i c u s a nd P . l u s i tan i cu s ) i n d ica t ive o fd r ie r a nd c oo le r c ond i t i on s . M . n i va l i s i sa l so f o und i nh i gh a moun t s f o r ac oa s ta l C an tab r ian s i t e ( t o ta l n=2 2 ) a nd i n d i ca te s o p en r o cky a r ea s n ea r Ama lda . H um id i n d ica to r s p ec ie s a r e r e duced , i n c lud i n g A . c f . t e r re s t r i s a nd M . o e conomu s . S ome t r ee c ove r i s i n d ica ted b yt h ep r e sence o fA pod emu s . T he S o lu t r ean l e ve l m ic rof auna l s amp le o f A i tzb i ta r te e x h i b i t s ad ef in i te c o ld c ha ra c t e r( A l t una 1 972 ) . Wh i le A . t e r r e s t r i s h a s t h e h i ghe s t f r equency a mong t h e m ic ro t ine s ( n= 1 20 ) , M . o economu s i ss im i la r ly a bundan t ( n =1 09 ) a nd f a r o u twe igh sb o th M . a g r e s t i s /a r va l i s( n =3 6 ) a nd P i tymy s( n= 1 3 ) . A lp ine -dwe l l ing M . n i va l i s i sa l so p r e sen t( n =3 ) ,w h i le n o A pod emu s h a s b een r e cove r ed f r om t h i s l e ve l . T a lpa i s t h e mo s t a bundan tt a xon ( n =6 16 ) d ue t oi t s g r ea te r e a se o f i d en t if i ca t ion f r om m u l t ip le d u rab le e l emen t s . T h i s m ic ro f a una a s semb lage i n d ica te s ah um id , c o ld , o pen e nv i ronmen t w i th p e rhap s s p a r se t r ee s a nd o pen w a te r n ea rby a nd i s s im i la rt oE lJ uyo L eve l4 / 4S . T he M idd le S o lu t r ean l e ve l s ( 1 5 , 1 6 , 1 7 ) o fL a s C a lda s a r e n o ted o n ly f o r t h e p r e sence o fA pod emu s c f .f i c r v ico l l i s , M . a r va l i s , M i c ro tu s s p p . , A r v i co la s ap id u s a nd P i tymy s c f . l u s i tan icu s( Co r chön 1 981) . T he se s p ec ie s a r e a l so c ommon t o t h e mode rn C an tab r ian f a una , t h ough t h e m ic ro t ine s f a vo r o pen e nv i ronmen t s a nd A pod emu s f a vo r s w ood land . Ma te r i a l f r om S o lu t r ean L eve l 3a tC ueva Mo r in i ss im i la r ly u n in f o rma t ive , w i th o n ly a s i ng le r ema in e ach o fA r v ico la a nd T a lpa r e po r ted ( A l tuna 1 978 ) . T he S o lu t r ean l e ve l a tE rm i t t i a ( A l tuna 1 972 ) h a sy i e lded o n ly M i c ro tu s a g re s t i s /ar va l i s , M . o economu s ,A r v i co la t e r re s t r i sa nd T a lpa e u ropaea . T he p r e sence o fM . o economu s a g r ee s w i th t h e c o ld p ha se a si n d i ca ted b y l owe r ed o cean l e ve l . L aR ie ra h a sy i e lded as ma l l m ic rof auna l s amp le f r om i t sS o lu t r ean l e ve l s ( 4 , 5 ,7 11 a nd 1 4 -17 ) t h a t c on s i s t s m o s t ly o fA r v i co la t e r re s t r i s . T he p r e s e nce o fG a l emy sp y rena i cu s i nL eve l 7i s as t rong i n d ica to r o f d ownwa rd a l t i tud ina l z one s h if t , s i nce t h i s s p ec ie s ( Py renean d e sman ) i st o day f o und o n ly a d ja cen t t ow a te r i n t h e a l p ine z one a bove 3 00 m ( Chap t e r 3 ) , w h i le L a R ie ra i s a lmo s tu pon t h em oder nc oa s t . MAGDALEN IAN ENV IRONMENTS T he Magda len ian p e r iod h a s s u f f ic ien ta moun t s o fr e cove r ed m ic rof auna l r ema in st oa low p a leoenv i ronmen ta l i n te rp r e ta 1 2 1

t i on b eyond i n d ica to rs pec ie s f o rp or t ion so ft h i s t ime ( Tab le 8 . 4 ) . T h i s c u l tu re s t ra t ig raph ic p ha se s pan s 1 6 ,500 t o1 3 ,500 B .P . ( Lowe r Magda len ian )a nd 1 3 ,500 t o1 ,000 B .P . ( Uppe r Magda len ian ) i n C an tab r ian S pa in , w i th s ome o ve r lap b e -

Mo re c omp le te i n f o rma t ion c ome s f r om t h e L owe r Mag d a len ian L eve l Vo fE r ra l la , wh ich o ve r l ies c u l tu ra l ly s te r i le L eve l I V ( P emän 1 985 ) . L eve l V h a s y i e lded d a te s o f 1 5 ,740+ / -240 a nd 1 6 ,200+4240 B .P . ( A l tuna 1 985b :26 ) .

t ween t h e se t wo ma jo r d iv i s ion s . I n a dd i t ion t o E l J uyo ( Chap te r 5 ) , m ic rof auna l d a ta h a s b een r e por ted f o r Mag -

T h i s d epo s i t c o r re spond s w i th t empo ra r i ly l owe red o cean

d a len ian l e ve l s f r om A i tzb i ta r te ( A l tuna 1 972 ) , A l tam i ra ( Ca s t ro B er na rdez 1 986 ) , E ka in ( Zaba la 1 984 ) , E rm i t t ia ( A l tuna 1 972 ) , E r ra l la ( Pemän 1 985 ) , E l P endo ( Fuen te s

t i on s o f n ea r ly t h o se o b ta ined a tt h e l a s t g l ac ia l m ax imum ( c a . 2 2 ,000 18 ,000 B .P . ) .

V ida r te 1 980 ) , L eze tx ik i ( A l tuna 1 972 ) , C ueva Mo r in ( A l tuna 1 978 ) , R a scaho ( A l tuna 1 981) , L a R ie ra ( A l tuna

A s d i scu s sed i nC hap te r 5 ,t h e L owe r Magda len ian l e ve l s o f

1 986 ) , T i to B u s t i l lo ( A l tuna 1 976 ) a nd U r t iaga ( A l tuna 1 972 ) . T h i s a r chaeo log ica l p e r iod c ove r s t h e l a te O xygen I s o tope S tage 2 a nd t h e o n se t o f S t age 1 ,r ep resen t ing s e a l e ve l r i se f r om t h e g l ac ia l m ax imum d u r ing t h e S o lu t r ean p e r iod t owa rd st h eh igh l e ve l s o ft h eH o locene . F luc tua t ion s b e tween t h e se t wo e x t reme s c omp l ica te i n te rp re ta t ion , s i nce o cean l e ve l r i se w a s n o t s t eady a nd w en t t h rough r e pea ted t empo ra ry l owe r ing s b ef o re o b ta in ing H o locene l e ve l s c a . 1 0 ,000 B .P . I n a dd i t ion , w a rm ing t empe ra tu re s c an b e e x p ec ted t op recede a n i n c r ea se i no cean l e ve l , s i nce m e l t ing o f c on t inen ta l a nd a l p ine g l ac ie r s i sn o t i n s tan taneou s . S im i l a r ly , t h e o n se t o fc oo l ing t empe ra tu re s w i l l b e f o l lowed b y l owe r ing o cean l e ve la si c ea ccumu la te so n l a nd . L owe r Magda len ian e nv i ronmen t s i nC an tab r ian S pa in c o r r e spond t og ene ra l ly h ighe r o cean l e ve l s a nd c o ld c ond i t ion s s omewha t a me l io ra ted f r om t h e g l a c ia l m ax imum . L owe r Magda len ian l e ve l s j u s t p o s tda t ing t h e g l ac ia l m ax imum h ave b een e xcava ted a tt h e u p land s i t e o fR a sca l ( ) ( A l tuna 1 981) . I n i t ia l Magda len ian L eve l 5h a s b een d a ted t o 1 6 ,433+ / 131 B .P . , a nd L owe r Magda len ian L eve l s 4a nd 3 h ave y i e lded d a te s r e spec t ive ly o f 1 5 ,988+ / 193 B .P . a nd 1 5 ,173+4 160 B .P . ( Gonza lez E chega ray a nd B a rand ia rän 1 98 1 :213 ) . T he s ma l l m ic rof auna l s amp le f r om L eve l 4 b i n c ludes s omewha t a noma lou s ly t h e p r e sence o f G l i s g l i s , i n d ica t ing s ome f o re s t c ove r . O the r s pec ie s p re sen t r ef lec t o pen a nd h um id c ond i t ion s . T he L owe r Magda len ian l e ve l s o fL a R ie ra ( 18 , 1 9 a nd 2 0 ) a l so i n d ica te o pen a nd h um id c ond i t ion s a nd i n c lude o ne a l p ine i n d ica to r ( M icro tu s n i va l i s ) a t as i t e a d jacen t t o t h e moder n c oa s t l ine ( A l tuna 1 986 ) . T he se l e ve l s h ave y i e lded c ompa rab le r ad ioca rbon d e te rm ina t ion so f1 6 ,420+4430 , 1 5 ,520+ / -350 a nd 1 5 ,230+ / 3 00 B .P . ( S t rau sa nd C la rk 1 986 ) . T he L owe r Magda len ian L eve l s V I Ia -f a t E ka in s how a s im i la r p a t te rn o fo pen , c o ld a nd h um id c ond i t ion s ( Zaba la 1 984 ) . M ic ro t ine s a r e d om ina ted b y A rv ico la a nd M i cro tu s o economu s w i th M . n i va l i s p r e sen t . P i tymy s a nd M . a g re s t i s /ar va l i s ( i nd ica t ive o f d rye r g r a s s land s ) a r e p r e sen t i n

l e ve l a nd t h e D rya s Ip o l len z one , a nd h ence c oo le r c ond i -

E l J uyo s how g radua l c l ima t ic d e t e r io ra t ion . M ic rof auna f r om L eve l s 9 ,8 , 7( d a ted t o 1 4 ,400+4 180 B .P . ) a nd 6s how ac o ld , v e ry h um id a nd o pen e nv i ronmen t . P a lyno log ica l a nd ma c robo tan ica l a na lys i s i nd ica te s t h a t h ea th p r edom i n a ted , a l ong w i th m any w e t land -p ref e r r ing t a xa . S ome s l igh t f o re s t c ove ra nd a r ea s o fg ra s s land a r e i n d ica ted . T he se l e v e l s c o r re spond t ot h e r e la t ive ly t empe ra t e P r e -Bö l l ing p o l len z one . M ic rof auna f r om L eve l 4 /4S ( d a t ed t o 1 3 ,920+4240 B .P . ) s how s d e te r io ra t ion o ft h i s e nv i ronmen t c o r re spond ing t ot empo ra ry l owe r ing o fo cean l e ve l a nd t h e D rya s Ip o l len z one . A n i n c r ea se i na l p ine s pec ie s i sn o ted a mong t h e m i c r of auna l , p o l len a nd m ac robo tan ica l r ema ins w h ich i n d i c a te s s i gn if ican t l owe r ing o f a l p ine z one s t o w i th in a f ew k i lome te r s o ft h i sc oa s ta l l ow land s i te . T he u nde s igna ted Magda len ian l e ve l a t E rm i t t ia ( A l tuna 1 972 ) h a s t o o s ma l l as amp le t o a l low i n te rp re ta t ion , w i th o n ly t h e l a rge r t a xa A . t e r re s t r i s a nd T .e u ropaea p r e sen t . B o th i n d ica te o pen a nd h um id c ond i t ion s . T he m ic rof auna l s amp le o ft h e u nde s igna ted Magda len ian l e ve l a tA i tzb i ta r te ( A l tuna 1 972 ) r e f lec t s ah a r she r e nv i ronmen t a nd i sh ence mo s t p robab ly L owe r Magda len ian i na g e . C o ld , o pen a nd w e t c ond i t ion s a r e i n d ica ted b y t h e d om inance o f A . t e r r e s t r i s ( n =1 95 ) a nd M . o economu s ( n =1 44 ) a nd t h e p r e s e nce o fM . n i va l i s ( n =5 ) , w h i le n o A pod emu s h ave b een r e cove red . S ome d rye r g ra s s land i sp r e sen t b a sed u pon t h e p re sence i ns i gn if ican t a moun t s o f M . a g re s t i s lar va l i s ( n = 6 5 )a nd P i tymy s( n =1 8 ) . T he m ic rof auna l s amp le f r om L owe r M agda len ian L eve l V a tE r ra l la i sl a rge e nough t oa l low c l o se rs t a t i s t ica l e xam ina t i on . T h i s s amp le i si n te rp re ted b y P emän ( 1985 ) a sh av ing r e su l ted f r om o w l p e l le t e ge s t ion i nt h e c a ve . Wh i le h e d oe s n o t s pec if y w h ich s pec ie s , e xam ina t ion o ft h e b ody s i ze p ro f i l e o ft h e m ic rof auna l p rey i n d ica te s p r obab le a ccumu la t ion b y am ed ium o w l . T he a ccumu la t ion o fs u ch al a rge s amp le i n ac ave s e t t ing a nd t h e v e ry l i gh t d amage u pon t h e i n d iv id u a l e l emen t s r u le s o u t o t he r m ed ium s i z ed o w l s pec ie s o t he r t h an b a rn o w l s ( Chap te r 2 ) . A na ly s i s o f t h i s s amp le w i l l p roceed w i th t h e a s sump t ion t h a t b ar n o w l s w e re i t sp r ima ry

much s ma l le rn umbe r s . A l tam i ra L owe r Magda len ian L eve l

a ccumu la to r s , t h ough s ub sequen t t a phonom ic e xam ina t ion

2 +3 r e f lec t s as im i la r ly o pen a nd h um id e nv i ronmen t , b u t w i th mo re i n d ica t ion s o f g ra s s land a nd a f o re s t s pec ie s

may a l te rt h i s v i ew a nd n ece s s i ta te a d ju s tmen t o fp a leoenv i -

( Apod emu s s y lva t icu s ) ( Ca s t ro B e rna rdez 1 986 ) .

r onmen ta l i n te rp r e ta t ion .

G rea te r

t r ee c ove r i si n d ica ted b y t h e Magda len ian ( Leve l 6 ) a nd

A na ly s i s o fE r ra l la L eve l Vi sp resen ted i nT ab le 8 . 5 . T h i s

M idd le Magda len ian ( Leve l 5 ) f r om D uf au re i n n ea rby F rance , w he r e A pod emu s i s as i gn if i can t p o r t ion o ft h e m i -

o ccupa t ion h a s b een r a d ioca rbon d a ted t o1 5 ,740+ / -240 B .P .

c r of auna l a s semb lage ( Ea s tham 1 995 ) .

L owe r Magda len ian o ccupa t ion o fE l J uyo . T h i s t ime c o r r e s pond s w i th r i s ing o cean l e ve l s j u s t p o s tda t ing t h e g l a c ia l max imum c a . 2 2 ,000 18 ,000 B .P . , s o c l ima t ica l ly r i go rou s c ond i t ion s w e r e l i ke ly . T h i s l e ve l h a s t h e l a rge s t t o ta l M IN I ( 7469 ) f o r a ny C an tab r ian U ppe r P a leo l i th ic l e ve l p ub l i shed

A c o ld , o pen a nd

h um id e nv i ronmen t i sa l so i n d ica ted b y t h e d om inance o f A r v ico la t e r re s t r i s ,M icro tu sr a t t icep s( =o economu s ) a nd M . a g re s t i s .

a nd 1 6 ,200+4240 B .P . ( A l tuna 1 985b :26 ) , p reda t ing t h e

1 22

t od a te . T h i s s amp le h a s b een a na lyz ed i nt h e s ame m anne r a s t h e E l J uyo m ic rof auna ( Chap t e r 5 ) : i n d iv idua l s p ec ie s MN I h ave b een c onver ted t ot o ta l p r ey u n i t s , w h ich i nt u rn r e p re sen t t h e t o ta l b i oma s s d e r ived f r om e a ch h ab i ta t t y pe a nd i t sp ropo r t iona lr e p r e sen ta t ion o fl a nd scape a r ea . S im i la r t o t h e L owe r Magda len ian o c cupa t ion o f E l J uyo , h ab i ta t a r ea ( F igu re 8 . 2 ) i s d om ina t ed b y h um id m eadow /wa t e re dge s pec ie s ( 7 2 .2 1% ) . T h i s t o ta l i n c lude s a v e ry h i gh p r opo r t ion o fM i c ro tu s o e conomu s , i n d ica t ing s i g n if i can t ly c o ld c l ima te a sw e l l a sh um id c l ima te . T h i s o pen l a nd scape a l so h ad s i gn if i can t a r ea s ( 1 0 .8% ) o fd ry m eadow ( g ra s se s ) . T he i n d iv idua l s o f t h e g enu s P i tymy s w e r e n o t i d en t if i ed t os p ec ie s . I nt h i sc o lde rr e g ime ,t h e se mo s t l i k e ly b e longed t oP .p y rena i cu s , ag ene r a l i s t s p ec ie s . S ome p o r t i on o ft h eP i tymy st o ta lm ay h ave b e longed t oP . l u s i tan i cu s , w h ich w ou ld i n c r ea se t h e t o ta l a r ea o fh um id m eadow p r e s e n t . F o r e s t c ove r w a s m in ima l w i th in t h e b a rn o w l h un t ing r a d iu s o ft h e s i t e , a moun t ing t ow e l l u nde r 1 % o ft h e l a nd a r ea . T ha t s ome f o r e s t w a s p r e sen t i sa l so i n d ica t ed b y t h e p r e sence o fG l i s g l i s , t h e p r e sence o fw h ich i nt h i s l e ve l i s s omewha ta noma lou st ot h eg ene ra le nv i ronmen ta lp i c tu r e . A ny t r ee c ove r w a s f ound i n s h e l t e r ed a r ea s o n t h e l a nd s c ape . H a r sh c ond i t ion s a r e r e inf o r ced b y t h e p r e sence o f t h e a l p ine s p ec ie s M ic ro tu s n i va l i s a nd t h e o n ly C an tab r ian U ppe r P a leo l i th ic o ccu r rence o f M i c ro tu s g rega l i s ( s t eppe l emm ing ) . T h i s l a t te r s p ec ie s i sf o und t o day i nt h e t u nd ra a nd s t epp e b i ome s o fA s ia ( Chap t e r 3 ) . T he se t wo s p ec ie s i n d ica t e s i gn if ican t a l t i t ude z ona l s h if td ownwa rd s t ow i th in ar a d iu s o f af ew k i lome t e r s o ft h i s l ow land s i te . T he p ro p o r t ion o ft h em m u s th ave b een g r ea t e r i nt h e a l p ine z one o f t h e C an tab r ian C o rd i l le ra t ot h e s ou th d u r ing t h i s o ccupa t ion o fE r ra l la . T h i s s i t e d oe s n o t i n c lude a ny r ema in s o ft h e a l p i ne s p ec ie sG a lemy sp y rena icu s . C an tab r ian U ppe r Magda len ian e nv i ronmen t s c hange i n a f l uc tua t ing m anne r f r om r e la t ive ly h a r sh i n te r s tad ia l t o w a rme r a nd m o r e f o r e s ted c ond i t ion s . O n ly i n conc lu s ive s amp le s a r e a va i lab le f r om U ppe r Magda len ian L eve l 2a t R a sca r lo ( A l tuna 1 981) a nd L eve l Ia t L ez e tx ik i ( A l tuna 1 972 ) . T he U ppe r Magda len ian L eve l s 1 a c o f T i to B u s t i l lo h ave b een d a t ed t ot h e v e ry e a r ly U ppe r M agda len ian p e r iod a nd i n f a c t o ve r lap w i th t h e e nd o f t h e L owe r Magda len ian . T he se r a d ioca rbon d e te rm ina t ion s r a nge f r om 1 3 ,520+ 1 -300 t o ( p e rhap s a noma lou s ) 1 5 ,400+ / -300 B .P . ( Mou r e R o m an i l lo 1 976 ) . A d a t e j u s t p r eced ing 1 3 ,000 B .P . w ou ld p l a ce T i to B u s t i l lo a m id s t t h e r e la t iv e ly t empe ra t e B o l l ing p o l len z one . T he m ic rof auna l s amp le f r om t h i s s i t e ( A l t una 1 976 ) s ugg e s t s t h a t s omewha t e a r l ie rd a t ing i sc o r r ec t , p l a c i n g t h e t i me o fo ccupa t ion d u r ing t h e e nd o ft h e c o lde r p o l l e np ha se D rya s I . Wh i le s ma l l ,t h e s amp le h a so n ly as i ng le r ema in o f af o r e s t s pec ie s ( Apod emu s ) a nd i sd om ina ted b y o pen -env i ronmen t m ic ro t ine s : A . t e r re s t r i s ( n = 6 1) , M . o economu s( n =7 ) , a nd M . a g re s t i s la r va l i s ( n =6 ) . Ac o ld p ha se a t t r ibu t ion f o r t h i s o ccupa t ion i sf u r the r s uppo r t ed b y t h e p r e sence o f as i ng le r ema in o fr e indee r ( Rang if e r t a ran d u s ) f r om l e ve l l b ( A l tuna 1 976 :153 ) . T h i s s pec ie s r a r e ly p ene t ra t ed a sf a rs ou th a sC an tab r ian S pa in a nd i sa s soc ia t ed w i th t r ue t u nd ra e nv i ronmen t s . O r ig ina l i n te rp r e ta t ion o ft h i s

s i t e i n d ica ted a L owe r Magda len ian i n du s t r ia l a t t r ibu t ion , u n t i l t h e s ub sequen t d i scove ry o f a n t le r h a rpoon h ead s p l a ced i tb yd ef in i t ion w i th in t h eU ppe r Magda len ian ( Mou r e R oman i l lo 1 975 ; Mou r e R oman i l lo a nd C ano H e r r e ra 1 976 , 1 978 , 1 979 ) . T i to B u s t i l lo i sa d ja cen t t o am a jo r r i ve r a nd n ea r t h e moder n c oa s t l ine . T he e a r ly p r e sence i ni to fh a r p oon h ead s( p re sumab ly f o rf i sh ing )m ay b em o r ed ue t os i t e f u nc t ion t h an t oc hang ing a r t if a c ts t y le . As im i la rm ic rof au n a l s amp le h a s b een r e cove red f r om t h e U ppe r Magda len ian L eve lV Ib o fE ka in ( Zaba la 1 984 ) . T he l a te ro ccur r ence so ft h eU ppe r Magda len ian s how m ixed a moun t s o fe nv i ronmen ta l a me l io ra t ion a nd m ay c o r r e spond t ot h e B o l l ing ( wa rme r ) , D rya s I( c oo le r ) , A l le röd ( wa rme r ) a nd D rya s I I ( c oo le r ) p o l len z one s . L eve l s 2 4 a nd 2 1-23 o f L aR ie ra h ave b een d a ted t ot h e e nd o ft h i sp ha se ( S t rau sa nd C la rk 1 986 ) . T h i s s i te r e ta in s a l p ine v o le M i c ro tu sn i va l i s i n L eve l2 4 a nd ( A l tuna 1 986 ) . T he s amp le s i z e i st o o s ma l lt o c onc lude m o r e a bou tt h e e nv i ronmen to t he rt h an m o r e p r ob a b le a s s ignmen t t o ac oo le r p ha se . T he F i na l Magda len ian L eve l V Ia a tE ka in h a s av e ry s im i la r m ic rof auna l s amp le ( o the r t h an t h e r e la t ive r e duc t ion i nA r v i co la ) a st h e U ppe r Magda len ian L eve l V Ib a tt h i ss i t e( Zaba la 1 984 ) . M ic rof auna l s amp le s f r om t h e F ina l Magda len ian l e ve l o f U r t iaga ( A l tuna 1 972 ) a nd L eve l 2o fC ueva Mo r in ( A l t una 1 978 ) a r e u n inf o rma t ive , a se a ch i sd om ina ted b y l a rge r m i c r of auna a s t h e p robab le r e su l t o f e xcava t ion t e chn ique . A pod emu s s y l va t icu s d oe s s how u p i n F i na l Magda len ian L eve l s I ab a nd I a tE l P endo ( Fuen te s V ida r t e 1 980 ) , i n d i c a t ing s ome r e tur no ff o re s tc ove r . T he F i na l Magda len ian l e ve l s a tE r ra l la a r e m o r e i l um ina t i n gd ue t ot h e i rg r ea te rs amp le s i z e . L eve l I h a sb een d a ted t o1 2 ,310+ / 190 B .P . ( A l tuna 1 985b :26 ) , c o inc id ing w i th t h e D rya s Io rA l le röd p o l len p ha se s . A s s ignmen tt ot h et empo r a ry w a rm ing c ond i t ion s o ft h e A l le röd i s mo r e l i ke ly b a sed u pon t h e m ic rof auna l s amp le ( Pemän 1 985 ) . T he a r bo rea l G l i sg l i s s how s u p f o rt h e f i r s tt ime a tE r ra l la i nt h e se l e ve l s , a sd oe s f o r e s t -dwe l l ing C l e th r ionomy sg la reo lu s . A pod emu s i sa l so a bundan t , w h i le M ic ro tu s n i va l i s d ec l ine s f r om p r ev i o u s ly h i ghe r l e ve l s . T he o t he r a l p ine s p ec ie s ( M . g rega l i s ) d oe s n o t r e cu r . Wh i le M . o economu s i ss t i l l p r e sen t , i tn o l o nge r d om ina te s t h e m ic rof auna l s amp le . O the r i n d ica to r s o f h um id o p en c ove r h ave d ec l ined ( Arv i co la , T a lpa e u r opaea a nd S o rex s pp . ) . T he c hanged m ic rof auna l p r of i le i n d ica te s t h e c oo l , h um id h ea th w i th s p a r s e t r ee s o f t h e L owe r Magda len ian h ave s h if t ed t owa rd s d r ye r g r a s s land w i th i n c r ea s ing t r ee c ove r a nd a r e t r ea t o ft h e a l p ine z one a way f r om t h e h un t ing r a d iu s o f E r ra l la . I n c on t ra s t , t h e n ea rby F r ench s i te o fD uf au r e s how s r e la t ive ly l i t t le m ic ro f a una l t ur nove r f r om i t s Magda len ian l e ve l ( 6 ) t h rough i t s M idd le ( 5 ) a nd F ina l Magda len ian l e ve l s ( 4 i n f . t h rough 4 s up . )( Ea s tham 1 995 ) . AZ IL IAN ENV IRONMENTS T he A z i l ian a r cha eo log ica l p e r iod i n C an tab r ian S pa in ( 1 1 ,000 t o9 , 000 B .P . ) o ccu r r ed d u r ing t h e T e rm ina l P l e i s to c ene / In i t ia l H o locene a nd t h e r ef o r e s pan s t h e t i me o fa b rup t o cean l e ve l r i se a nd g ene ra l c l ima t ic a me l io r a t ion . T h i s p e r i od b eg in s d u r ing t h e ( c oo le r ) D rya s I I p o l len z one a nd c on t inue s i n to P rebo r ea l . G loba l s e a l e ve l i sr e co rded a s 1 2 3

e a r ly O xygen I s o tope S t age 1 ,w he re l e ve l s r a p id ly r o se t o t h e a pp rox ima te mode rn l e ve l b y 1 0 ,000 B .P . M ic rof auna l s amp le s ( Tab le 8 . 6 ) a r e l e s s a dequa te t h an t ho se f o r t h e Magda len ian p e r iod a nd c ome f r om t h e s i te s o fA i tzb i ta r te ( A l tuna 1 972 ) , E ka in ( Zaba la 1 984 ) , E rm i t t ia ( A l tuna 1 972 ) ,

e f f ec t s o fa g r icu l tu re , p a s to ra l i sm , f o re s t ry a nd u rban iza t ion a nd d oe sn o tr ef l ec ts ub sequen tc l ima t ic c hange . M ic rof auna f r om t h e se l e ve ls i sl i s ted i nT ab le 8 . 7 . T he F ina l Me so l i th ic i sr e p re sen ted a t Ma r izu lo ( A l tuna 1 972 ) , w he re

E lP endo ( Fuen te sV ida r te 1 980 ) , R a scaf io ( A l tuna 1 981) , L a R ie ra ( A l tuna 1 986 ) ,a nd U r t iaga ( A l tuna 1 972 ) .

t h e o n ly s ma l le rm ic rof auna l s pec ie s r e p re sen ted i sG lä g l i s . T he E neo l i th ic L eve l b 2 -b1 a tA baun tz h a s y i e lded o n ly t h e r ema in s o f l a rge r m ic rof auna i n c lud ing E r inaceu s e u -

T he c hang ing C an tab r ian A z i l ian e nv i ronmen t r ef lec t s t h e

r opaeu s . T h i s l e ve l h a s y i e lded a r a d ioca rbon d a te o f 4 , 240+ / -140 B .P . T he N eo l i th ic i sr e p r e sen ted a t Ma r izu lo ,

w a rm ing c l ima te w i th as p read o f f o re s t s , t h e c l i se r ia l s h if t n o r thwa rd s o f c o ld -adap ted s pec ies r ange s a nd t h e r e t r ea t u pwa rd s o fa l p ine b io t ic z one s ( Pef ia lba 1 994 ) . T h i s t r an s i t i on i sn o ted a tL a R ie ra , whe re t h e s ma l l m ic rof auna l s am p l ef r om A z i l ian l e ve l s2 7 -30 i n c lude st h ea r bo rea l d o rmou se s pec ie s G l i s g l i s f o r t h e f i r s t t ime ( A l tuna 1 986 ) . D o rm ice t e nd t ob er a re o w lp rey , s i nce t h ey d we l l a mong t r ee c r own s a nd a r e t hu s mo re d i ff icu l t t oh un t . T he i rp r e sence i n as i te t h e ref o r e s t rong ly i n d ica te s f o re s ted c ond i t ion s . A s im i la r p a t te rn i sf ound a tE l P endo ( Fuen te s V ida r t e 1 980 ) . T he m ic rof auna l s amp le s f r om t h e A z i l ian l e ve l s o f R a scaf io ( A l tuna 1 98 1) , E rm i t t ia a nd U r t iaga ( A l tuna 1 972 ) , a nd E ka in ( Zaba la 1 984 ) a r e u n inf o rma t ive a st h ey y i e ld p r ima r i l y A r v ico la a nd T a lpa ( p e rhap s n o t c o inc iden ta l ly t h e t wo mo s tv i s ib le t a xa a mong t h e s ma l le rm ic rof auna ) . T he s i tua t i on i ss im i la r f o r A z i l ian L eve l 3f r om t h e n ea rby F rench s i te o fD uf au re , t hough t h e a r c t ic v o le M icro tu s o economu s p e r s i s t s l o ca l ly ( Ea s tham 1 995 ) . A i tzb i ta r t e h a s y i e lded amo re s ub s tan t ia l m ic rof auna l s am p l e f r om i t s A z i l ian l e ve l ( A l tuna 1 972 ) . A pod emu s s how s u p f o rt h e f i r s tt ime i nt h i s s i te . A i tzb i ta r te h a s al a te p e r s i s t e nce o f M i cro tu s o economu s a nd M . n i va l i s , t hough t h e f o rme r s pec ie s d ec rea se s r e la t ive t o o the r m ic ro t ine s i nt h e Magda len ian l e ve la tt h i s s i te .

w he re r ema in s o f u b iqu i tous A r v ico la t e r re s t r i s a nd T a lpa a r er e por ted . T he C ha lco l i th ic l e ve l s I Ib a nd I Ia a t Ama lda ( Pemän 1 990 ) h ave y i e lded s ome r ema in s o f a r bo rea l G l i s g l i s a s w e l l a s f o re s t -f avo r ing A pod emu s a nd C le thr ionomy s g la reo lu s . A l so c ommon a r e M icro tu s a g re s t i s a nd P i tymy s p y rena icu s , w h ich f a vo r mo re o pen e nv i ronmen t s . Ama lda a l so s hows t h e l a te s u rv iva l o f al o ca l ly r e l ic t p opu la t ion o f M ic ro tu s o economu s . T he p re sence o f t h i s s pec ie s i n a w a rme r c l ima te i nc ompe t i t ion w i th M . a g re s t i s a nd M . a r v a l i s i n d ica te s t h e p re sence o f aw e t land /hum id m ic rohab i ta t n ea r t h e s i te . A l so p re sen t i sM . n i va l i s , a n a l p ine s pec ie s w h ich i n hab i t s e xpo sed r ocky a r ea s . T he l o ca t ion o f Ama lda i nt h eC an tab r ian C o rd i l le ra f oo th i l l s( w i th h i l l s a bove 6 00 m w i th in 2 km o f t h e s i te , w e l l w i th in t h e h un t ing r a nge o f m any o w l s pec ie s ) e xp la in s t h e p e r s i s tence o fM . n i va l i s t o t h i s l a te d a te a f t e r t h e g ene ra l c l ima t ic a me l io ra t ion . T he B ronze A ge i sr e p re sen ted b y L eve l Ba t U r t iaga ( A l tuna 1 972 ) . S ign if ican t f o re s t c ove r i nt h e v i c in i ty o ft h i s s i te i s i n d ica ted b y t h e p r e sence ( n =1 0 ) o fG l i s g l i s , w i th A . t e r r e s t r i s , T a lpa e u ropaea a nd t h ree t a xa o fb a t sa l so p r e sen t .

1 982 ) , Ama lda ( Pemän 1 990 ) , Ma r izu lo a nd U r t iaga ( A l tuna 1 972 ) . T he m ic rof auna f r om t h ese l e ve l sr ef l ec t st h e g ene ra l w a rm ing t h a th ad o ccu r r ed a nd w i th i tt h e r e tur no ff o re s ted

T he L a te R oman p e r iod ( Leve l I ) s amp le f r om Ama lda i s mo r e s im i la r t ot h e c on tempo ra ry m ic rof auna o fC an tab r ian S pa in t h an t ot h a t o f t h e L a te P l e i s tocene o r e a r l ie r H o lo c ene . T h i s l e ve l h a s h igh p ropor t ion s ( n =1 56 ) o ft h e f o res t t a xon A pod emu sa nd s ome ( n =1 ) C l e th r ionomy sg la reo lu s a nd ( n =3 )G lä g l i s . A rv ico la c f . t e r re s t r i sh a s n ea r ly d i sap p ea r ed f r om t h e m ic rof auna l p rof i le . Wh i le o t he r h um id m eadow /wa te r e dge t a xa a r e p re sen t ( P i tymy s l u s i tan icu s , S o rex a raneu s a nd N eomy s ) , d ry m eadow s p ec ie s (M icro tu s a g re s t i s a nd C roc idura s pp . ) a r e f ound i ng r ea te r a moun t s . T he i r g r ea t a bundance i n d ica te s t h e s i gn if ican t l o ca l a l te ra t i on o f l a nd scape f o r a g r icu l tu re , a nd i sr e inf o rced b y t h e a r r iva l o ft h e mu r id s M icromy s m inu tu s a nd Mu s m u scu lu s . T he f o rme r s p ec ie s , wh i le n o t at r ue c ommen sa l , i sa s soc i -

c ond i t ion s t o t h e C an tab r ian c oa s t . T he mode rn o pen C an tab r ian e nv i ronmen t ( Chap te r 4 ) i sl a rge ly d ue t o t h e

a t ed w i th t h e s p read o fa g r icu l tu ra l f i e ld s . T he l a t te r s pec ie s i so fc ou r se a s soc ia ted w i th t h e s p read o fh uman h ab i ta t ion s .

POST -PALEOL ITH IC ENV IRON MENTS Wh i le n o t t h e f o cu s o f t h is r e sea rch , d a ta f r om p o s t p a leo l i th ic l e ve l s o fs i te s a l so y i e ld ing p a leo l i th ic l e ve l s a r e p r e sen ted t oi l u s t ra t e t h e e nv i ronmen ta l c hange st h a te n sued a f t e r t h e P l e i s tocene /Ho locene t r ans i t ion c a . 1 0 ,000 B .P . We s ter n E u ropean p o l len z one s p roceeded t h rough t h e B o r e a l( 9 ,000 -8 ,000 B .P . ) ,A t lan t ic ( 8 ,000 -5 ,000 B .P . )a nd S ub B o rea l ( 5 ,000 -3 ,000 B .P . ) . M ic rof auna l d a ta i sp r e sen ted h e re f r om t h e s i te s o f A baun tz ( A l tuna a nd Ma r iezku r r ena

1 24

T ab le 8 .5 : L ower Magda len ian L eve l Vo fE r ra l la .

MN I M i cro tu sn i va l i s

%MN I 1 29

M i cro tu sg r ega l i s

2

G l i sg l i s

X

A pod emu s

8

T a lpa c f . e uropaea S o r ex a raneu s S o r ex m inu tu s

*

%B ioma s s

1 29

1

H ab i ta t

%Hab i ta t

A s soc ia t ion

A rea

1 . 4

A lp ine

2

0 . 02

T und ra

8

0 . 1

F o re s t

1 370

1 5 .3

H um id Meadow

7 4 1

8 . 3

H um id Meadow

0 . 0 1

H um id Meadow

[ 4 ]

2 74

3 . 7 1 9 .8

0 . 5

0 . 07

0 . 2 0 . 75

1 . 42

F o re s t

1

1 482 5

N eomy sf od i en s

T o ta l B ioma s s

1

1 . 7 0 . 03

0 . 1

P r ey U n i t s

5

1

0 . 1

2 9

0 . 4

2 1 .75

0 . 2

We t land s / Wa te r

2 9 10

3 9 .0

1

2 9 10

3 2 .5

H um id Meadow

A rv ico la

2 85

3 . 8

5

1 425

1 5 .9

H um id Mead / Wa te r

M ic ro tu sa g r e s t i s -a rva l i s

9 68

1 3 .0

1

9 68

1 0 .8

D ry Meadow

1 0 .8

P i tymy s

1 377

1 8 .4

1

1 377

1 5 .4

G ene ra l i s t?

1 5 .4

T o ta l

7 469

1 00 .0

8 952 .75

9 9 .9

M ic ro tu so economu s

F igure 8 .2 : P a leoenv ironmen t o fE r ra l la L eve l V .

% A rea G ene ra l is t /O the r 1 5%

A lp ine /Tund ra 1 %

( Fo r e s t 0 . 1% )

H um id Meadow /Wa te r 7 3%

1 25

7 2 .2 1

9 9 .9

> 1 3

r n

e s i

( ,1

g o». I

C I D

,

r n

X

C . )

0 2 . .

0

C . ) •C . ) •

0 •C . )

•

• » ‘

0 • E l i

u t

1 26

S o r ex m inu tus

0

S o r ex c o rona tus

0 2 . C . 3

S or ex a raneus

O 2 . .

C . ) •

A r v ico la t e r r e s t r i s

u so economus

R a t tu sr a t tu s

e • I

3: . 3 .

o o

E

e

oo z e : , e uc ue u b e .

z e m

. z. .. . , r . e . =

c u

U . 1

• —

X

' 1 2 1 X

t a . 3

= . . e c t

. b e L . 2

r y

a .

7 )

r -

g b e • 3 7 .

0 3 . . . . • , .

• • 1

m t

' 3 . — P. M • mb d

1

.

0

V I I . .

)

C, 1

=

z c o t le ' E s a s . 1 C L J

5

0 0

e r

4 0 1 . )

g .5.

c d e = s o

r ‘ s

. .

V D 0 0 C V

r n

0

r - " r c . _

e v

r

C P

• •

, .. .

• — • v-)

C * 93

• c . , =
2 .

E

0 2

z E 0

e 3 0

-G 1 3 >

0

2 t . )

. 2

. . • .

22

›,

Z

O

z s

r e.

2 en

. . 2 .."Z " . . i . :

t 2

• . .

. >

e e

0 • . J2

•.4

k • . .4

Z • . . . .

ee . 3

In

e ? : . « . e 0 1

2 2

7l ä

e, e,e,. ' : ." .c ' •G . 1L e e l c 3

S o r ex c orona tus

r . ; z

. . e

S or ex a raneus

. 2

A r v i co la t e r res t r i s

1 . 1

R a t tus r a t tu s

T ab le 8 . 3 : ( Con t . )

0 0

V Z ,

S o r ex m inu tu s

-c r d

0

e r ' ä

1 , 1 . 1 1

en

1 , 1 1 1

0

4 1 2 ^ r ,

1 . ) 0

▪ L ▪ L . 1

7 3 '

0 0 0 0

0

•.••

0

u . 3 c i

L L 2

• • z , L

1 . )

t 5:

>

e ä ).

• •••/

0 .

0 0

4 / 1 . 0

e r

ee l

r e )

▪0 0

0 0

i r )

• — •

I t

e m e

e t •

• • • 1

c h

e z n

2 . 5 ( 2 1

• "

a . ) c • e = = = C l . ) >

c d

•

= l•• e . — . . . = 3 ) •c u e

c o

l e )

ee l

g . ,

b . + N

c v

4 / 1 . 0

en

. 0 . )

o z e c e2 , > 4 e

t A

0 \ M r

'J D

t r ) c 7 N

0

es

X

. . . . . • C U

4 2

e l

t 30

. 2

E

E

E

E

z " e s l o -t 3 L

X

N • —. , — a .r z .c z .• . 4 . 1 . 1 0

0

0

o v )

1 3 1

C roc idura r u s su la

. z L . z . )

S o rex m inu tus

, . * . z 5 . . . . _ , . e . . . —

S o r ex c orona tus

. . 0 >

z— . — .2

' 2

S o r ex a raneus

c o , c n

, =

A rv i co la t e r r es t r i s

2

R a t tu sr a t tus

* c e = = « 3 t 4 .

t U

r g r-

• •••4

‚ 1 )

VD

1 •••• 1

0 0

' 1 1

( e l

0

2

N

e r e r

• — • e n

00

le t

00 CT +

c 7 p >

0 0 0

en

>
g . j . 2

R a t tus r a t tu s

c e

2 c 4

0 " 0 1 1 ,

7 . 2

" ä ". L I Z

G L 3

> r s 1

• — z u L U

, N t 3 • • • •

0 0

t r ä

/• • •

L aR ie ra

L aR ie ra

L aR ie ra

L aR ie ra U ppe r Mag /Az i l

B ronze Age

c o

F ina l Me so l i th ic

c v

e l •

f9 1

1 . .. . 1

v-)

N

N

N

e«. I

VI

• e r

e l-

CT

. .. .

0

C1

1 1 1

—

0 en

S

e n

• ••• •

N

S

en

c v

X

e r r N

X

C T , —

en

e f 00

S

N

00

00 . . .

N

CN

N

el N

en . -.

S en

V D e t '

39 N —

en

. 3 =

0 L . C . )

0 L . C . )

. . .

. ' . ' '

0 s . C . )

0 ) . C . 3

0 l . Z . )

0 I . C . 3

0 4 . C . )

' .

. . .

. ' . .

0 0 6 2 L

. 1

. 3 •. .

z l› e . . . . , 0 .E c s,

' zm e z - u, ' b

' t l i

, e , „ o

c u

• • •

. 2

,

Z

• ••

1 .

1 . 3

tn

M

c . >

o .r . z . )• e E eE er iu ue l ' S i e • / ' ' ' c % ` ' c i '2 o • , mL . )c l .t a . . Or Q., F. E-. E- cn t e

0 2

1 1

0

•

t

1 3 5

2

2

. S

‘

2

-

-

C roc idura r us su la

r n

3 4 / 1

S or ex c orona tu s

N

N

S or ex m inu tus

X

S

N

e r

S or ex a raneu s

0

A r v i co la t e r r e s t r i s

E
-0

e d

a < ; # N C I J

C Z

-0

. 2

1 . -

0 0 0

b

c h 0 r • • t-

c • 1 N

,, E S. 'c — c uc h u 0 \ , c z i U ? eJ ,— — 6 .. . , •0 — N

0

2 .

=

s 0 . .

h e

C • e i

c " e

C7'

--

e l

' e.

6 "

c >

,

. t e L )

c e

I D

c : i

6 ' 3
' , = • . ,. — : . , )O . ) • •c eW 2 V GEa . )< < ' L . . *2 4 4 C< 1 t . . k i • • o . . c l , , .•.-s .o r t— • —1 : 3 i t t . I ,k i' . 2 ° > MI ,= . z . . "0

6 .

E

C A
L H7 1.

. J

1 3 6

CHAPTER 9 EXPLO ITAT ION OF FUR -BEAR ING MA M MALS

Ar e la t ed s ubca t ego ry o f s ma l l m amma l w a s r e cove r ed i n s i gn if i can ta moun t s f r om t h e f i ne s c r een d epo s i t so fE lJ uyo : t h e r ema in s o f m ed ium s i z ed f u r -bea r ing s p ec ie s , mo s t o f w h ich b e long t o O rde r C a rn ivo ra . Wh i le m any o f t h e se s p ec ie s f a l l b e low t h e u ppe r b ounda ry o f 5k g u s ed i n d e l im i t ing m ic rof auna , m any d o n o t ( a t l e a s t a s a du l t s ) . T he s e l a rge r s p ec ie s a r e i n c luded i nd i scu s s ion h e r e , s i n ce t h e i r i n c lu s ion i n t h e s i t e s how s t h e s ame p a t t e rn a s t h e s ma l le r r ema in s w i th in t h i s g roup . E v en t h e s ma l le r o ft h e t a xa d i scu s sed h e re , w h i le f i t t ing u nde r t h e c a t ego ry o f m i c romamma l s , a r e g ene ra l ly l a rge r t h an t h e t y p ica l r oden t a nd i n sec t ivo r ef a una d e sc r ibed i nt h ep r ev iou sc hap te r s . A s s u ch , t h e s e t a xa e xpe r ience d i f f e r en t t a phonom ic p roce s se s , i n c lud ing s ou r ce o f mo r ta l i ty , manne ro ft r an spo r tt ot h e s i t e , p a t t e rn a nd d eg r ee o f s k e le ta l e l emen t d e s t ruc t ion , a nd d epo s i t ion w i th in t h e s i t e . Wh i le i ti sp o s s ib le t h a t s ome o f t h e s e i n d iv idua l s ( e spec ia l ly t h e s ma l le s t s p ec ie s s u ch a s Mu s te la n i va l i s a nd M . e rm inea o rj uven i le s o f t h e l a rge r s p ec ie s ) w e r e i n t roduced a st h e p r ey o fo w l s , t h e p a t t er n o f s k e l e ta l e l emen t r e cove ry s ugge s t s t h a t t h e se w e r e b rough t i n to t h e c ave b y t h e d i rec t a c t ion o f h uman s . T he d epo s i t iona l p a t t er n o ft h e se f u r -bea r ing m amma l s i sa l so i n d i r ec t c on t ra s t t o t h a t o f h edgehog ( E r inaceu s e u ropaeu s ) r ema in s r e cov e r ed f r om E l J uyo ( Chap t e r 2 ) . Wh i le t h i s s p ec ie s i so f as im i la r s i ze t o s ome f u r -bea r e r s ( a bou t 1k g a du l t m a s s ) , i ti sn o t au sab le s ou r ce o f f u r d ue t o i t s p r o t ec t ive s p ine s . T he p a t ter n o f f u r -bea r e r r e p re sen ta t ion a nd i t si mp l ica t ion sw i l lb ed i scu s sed b e low . E l J uyo h a s y i e lded t o d a te 7 2 f o o t e l emen t s , p r ima r i ly p ha lange s a nd m e tapod ia l s o fc ar n ivo r e s i nt h e s i z e r a ng e o f e rm ine ( M u s t e la e rm inea ) , f e r re t (M p u to r iu s ) , a nd m a r t en ( M a r t e s s pp . ) . T he p r e sence o ft h e se e l emen t s i si ns t r ik ing c on t ra s t t ot h e v i r tua l l a ck o fa ny o t h e r e l emen t s , i n c lud ing t e e th , f r om t h e se t a xa ( Chap te r 5 ) . N on -ca rn ivo re s o ft h i s s i z e r a ng e ( s u ch a s r a bb i t o r h edgehog ) d o n o t s how t h i s s ame e l emen t r e p re sen ta t ion d i sc r epancy , n o r d o s ma l le r , m o r e f r ag i le r o den t a nd s ma l l i n sec t ivo r e t a xa . D if f e r en t ia l d e s t ruc t ion o fn on -f oo t e l emen t s o ft h e f u r -bea r ing s p ec ie s t h e r ef o re m ay b e r u led o u t . T he a u tho r a t t r ibu t e s t h e p r eponde rance o ff u r -bea r ing mamma l f o o te l emen t st ot h e i r s e lec t ive i n t roduc t ion t ot h e s i te b y ah uman a gency . S uch a p a t t er n i st h e l o g ica l r e su l t o f f oo t b one s l e f t i n ap e l t t h a t w a s s k inned f r om a n a n ima l t r apped o r h un t ed a way f r om t h e s i te , w i th t h e r ema in ing c a r ca s s i n c lud ing a l o t he r s k e le ta l e l emen t sd i sca rd ed i nt h e f i e ld . Mu s t e l id m ea th a s a s t rong , u np lea san t f l avo r d ue t o t h e i r s t rong s c en t g l and s ( p e r sona l o b s e rva t ion ) a nd may h ave b een t h e r ef o re a l e s s d e s i rab le f o od c ho ice . T he se s pec ie sw e r ep r obab ly a cqu i r ed w ho l ly f o r t h e i r p e l t s , w i th n o o t he r a r cha eo log ica l ly r e cove rab le p o r t ion o ft h em r e a ch ing t h es i t e . D ue t ot h e n a t u r e o ft h e e v idence i n vo lved , t h e e xp lo i ta t ion o f a n ima l s f o r t h e i r p e l t s i s d i ff i cu l t t o d e t ec t a r cha eo log ica l ly . F i r s t , i tp r e suppo s e s ap a t t er n o f f a una l p r o cu r emen t a nd p roce s s ing t h a t , w h i le al o g ica l e xped ien t , i sb y n o m ean s t h e o n ly s y s tem p o s s ib le f o r f u r a cqu i s i t ion .

Med ium c a rn ivo r e s m ay h ave b een u sed f o r t h e i r m ea t a l so ( s uch a s f ox o r l y nx ) , a nd i ns u ch c a se s a n e n t i r e c a r ca s s w ou ld b e r e tur ned t oa nd d i spo s ed o fi n as i te . E n t i r e s ma l l c a rn ivo ran c a r ca s se s m igh t a l so h ave b een i n t roduced t o a s i t ep r io rt os k inn ing , s imp ly b ecau se t h e c o s to ft r an spo r t ing t h em w a s n o t d eemed s i gn if i can t o rw a s l e s s t h an t h e e f f o r t o f s k inn ing t h em i n t h e f i e ld . Mamma l f oo t e l emen t s ( c a rpa l s / ta r sa l s , m e tapod ia l s a nd p ha lange s ) a r e r e la t ive ly s ma l l a nd d u rab le ( K le in a nd C ruz -U r ibe 1 984 ) a nd h ave a r e la t ive ly l ow u t i l i ty i na s so c ia t ed m ea t , c on ta ined m a r row , o ra s ar aw ma t e r ia lf o rt o o lm anuf a c tu r e . T hey a r et h u s l e s s l i ke ly t h an many o t he r e l emen t s ( l ongbone s , v e r t eb ra e , r i b s ) t ob e d e s t royed b y p o s tdepo s i t iona l f o r ce s o rt ob e d amaged a s a p a r t o f b u t che ry , m a r row e x t ra c t ion , o r a r t if ac t m anuf ac t u r e . T he i r o ve r rep r e sen ta t ion i ns ome s i t e s i sd ue t o t h e se t a phonom ic f a c to r s . I n a dd i t ion , c ar n ivo re s o f t en d en i nc ave s a l t e rna t e ly w i th a ny h uman o ccupan t s o r m ay e nc roa ch u pon h uman o ccupa t ion i no rde rt os c avenge f ood . C a rn ivo r e r ema in s m ay b e p r e sen t i n as i te t h e r ef o r e a s a r e su l t o f n a tu ra l mo r ta l i ty ( Ku r t 6n 1 972 , 1 976 ) a nd h ave l i t t le t od ow i th h uman a c t iv i ty . Ap a t ter n o ff u r e xp lo i ta t ion h a s b een p r ev iou s ly p ropo sed f o rU ppe rP a leo l i th ic s i t e s i nC an tab r ian S pa in ( S t rau s 1 977 , 1 982 ) . S pec ie sr e cove r ed f r om C an tab r ian s i t e s r a nge a c ro s s t h e e n t i r e c a rn ivo r e s p ec t rum f r om t h e s ma l le s t c a rn ivo ran (M u s te la n i va l i s ) t h rough m ed ium s iz ed mu s te l id s , f o xe s , a nd l a rge r s p ec ie s s u ch a s w o lf a nd b ea r . T he se a r e s umma r iz ed b yS t rau s( 1 992 ) . T he m a jo r i ty o ft h e se r ema in s m ay b e a t t r ibu t ed mo re e conom ica l ly t o t h e u t i l iza t ion o f c ave s b y c a rn ivo re s i n d epend en t o f h uman a c t ion s . A i tzb i ta r te I V i nB a sque c oun t ry h a s y i e lded s e ve ra l r ema in s o f Mu s te la n i va l i s , M e rm inea , a nd M . p u to r iu s i n i t s S o lu t rean a nd Magda len ian l e ve l s ( A l tuna 1 972 ) . B o l inkoba , aB a sque s i t e , h a s a l so y i e lded t h e r ema in s o f Mu s t e l id sa nd r e df o x( Vu lpe sv u lpe s ) . THE LO WER P ALEOL ITH IC T he A cheu lean i nC an tab r ian S pa in i sp oo r ly r e p r e sen ted , s o m ak ing m ean ingf u l s t a t emen t s a bou t s ma l l f a una l e xp lo i ta t ion i sd i ff i cu l t( F igu r e9 . 1) . O n ly t h eb a sa l l e ve l s o f E l C a s t i l lo h ave y i e lded s t r a t if i ed d epo s i t s f r om t h i s p e r iod ( Cab r e ra 1 984 ) . S ome e qu ivoca l o pen a i r s i t e s a l so h ave b een f ound , s u ch a s B a f iugue s ( Rod r iguez a nd F l o r 1 979 ) , a nd t h e r e a r e s ome i s o la t ed s u rf a ce f i nd s . F auna l i n f o rma t ion i st h e ref o re s c a r ce . Sma l l mamma l f i nd s i n o t he rt h an g eo log ica l s e t t ing s c on s i s to n ly o ft h e p r e sence o f Ma rmo ta m a rmo ta i n E l C a s t i l lo L eve l 2 5a ( Cab re ra 1 984 :134 ) . T yp ica l l a rg e f a una ( Can i s l u pu s , E quu s c aba l lu s , U r su s s pe laeu s a nd C er vu s e l aphu s ) a r e a l so r e p r e sen ted i ng ene ra l ly much g r ea t e r a moun t s . T he r e i sn o d i r ec t e v idence t h a t M m a rmo ta w a s i n t roduced t ot h e E l C a s t i l lo c ave b y h om in id a c t ion , w i th n a tu ra l mo r ta l i ty o r a n ima lp r eda t ion b e ing l i k e ly a l t er na t ive s .

1 37

41 .

+ + + 4+

C an tabr ian S ea

F igure 9 . 1 : C an tabr ian M idd le a nd Upper P a leo l i th ic S i tes Y ie ld ing t he R ema ins o fF ur -bear ing S pec ies .

4+ +

. 0a o n

AZ E

e g N •C

4 1 4 . 0 1 0 L )

O

E 0 3

c 0 3

g

-a

r -

A

C 0 3 U 5 0 7 g . .

, 2,

: = C

0 3

L A

0

e t o .4 > 1 : 3 ' ; ) ' c

. . o

oE c o o

3 4e .c 1 3c / .

C

0 3 0 1 J 2 . 1

4 A 0 —

E4 , , , ,3 4'i E 7 7 . K i e r x ea 0e lz " gz r w ,r . 2 E 3= ,-0 oc 4 1 , 0 . 0 • • fA › , . . . . 4 1 4 C . . . g o w C + .. , 1 • A .. 7 5 'C O ,

t 3 .

4

i l :

o

i e

g

3

Z

c

0

4

4

4 ' 4

4

4

, 1 ( 4 1 e n e t I i ) % J D 0 . . .

1 38

P4 c c " . 4 e l M I N e t f ) s o r - c o o n

Q CC . %0 . 11 . 11 . . 1l •1 . 1v lv . 1v . 1v , 1/ . 1 C J

THE M IDDLE P ALEOL ITH IC T he Mou s t e r ian p e r iod i nC an tab r ian S pa in ( Tab le 9 . 1 , e nd o fc hap te r ) i sf a r b e t te r d ocumen t ed , a nd mu l t ip le s t ra t if i ed c ave s i t es h ave b een e xcava ted . L a rge f a una e xp lo i ta t ion b y t h e N eand e r tha l i n hab i tan t s d u r ing t h i s p e r iod e xh ib i t s a m o r e e c l ec t ic a cqu i s i t ion p a t t e rn ( F r eeman 1 973 , 1 98 1 ; S t rau s 1 982 , 1 9 92 ) t h an i su sua l ly t h e c a se f o r t h e U ppe r P a leo l i th i c . T yp ica l l a rge p r ey s p ec i e s s u ch a s r e d d ee r , h o r s e , a nd b i son w e re t a ken . N o s p ec ia l iza t ion u pon r e d d ee ro ri b ex , s u ch a se v idence f o rm a s sd r ive so fe n t i r eh e rd s t h a t o ccu r s l a t e r ( S t rau s 1 977 ; K le in e ta l . 1 98 1) , w a s y e t e xh ib i t ed d u r ing t h i se a r l ie rp e r iod . A ne c l ec t ic p a t t e rn a l so h o ld s f o rs ma l lm amma le xp lo i ta t ion d u r ing t h e M idd le P a l eo l i th ic i n S pa in a nd e l sewhe re i n E u rope ( S t ine r 1 993 , 1 994 ) . S ome s ma l l m amma l s a r e f o und i ns ma l la moun t s i ns ome l e ve l s . E lC a s t i l lo L eve l 2 0 i n c lud e s t h e p r e sence o f V u /pe s v u /pe s , a nd L eve l V I I a t Ama lda y i e ld ed 2 9 r ema in s o f t h i s s p ec ie s ( P emän 1 990 ) . L eve l V I a tL ez e tx ik i ( A l tuna 1 972 ) y i e lded f o x , p l u s Me / e s m e / e s , F e l l s l y nx a nd O ty c to lagu s c un i cu lu s i n s ma l l a moun t s , w h i l e b o th f o x a nd F e / i s s i l ve s t r i s w e r e f o und a t A x lo r ( S t rau s 1 982 ; A l t una 1 989 ) . T h i s l ow r e cove ry l e ve l o f s ma l l f u r -b ea r ing m amma l s f o r t h e Mou s te r ian m ay b e d ue i ns ome c a se st ot h ea n t iqu i ty o ft h ee xcava t ion sy i e ld ing t h ed a ta . S uch i sn o tt h ec a se f o r Ama lda ,w h i ch p e rhap sn o t c o inc iden ta l ly y i e lded t h e l a rge s t n umbe r o f i d en t if i ed r ema in s o f a ny o f t h e se t a xa . C on t ra r iw i se , t h e mod e rn e xcava t ion s o f C ueva Mo r in w i th i t s m u l t ip le Mou s te r ian l e ve l s( Gonza l ez E chega r ay a nd F r eeman 1 978 )d i dn o ty i e ld a ny o ft h e s et a xa , e ven t h ough s ma l le rr oden ta nd i n sec t ivo re s p ec ie s w e r e r e cove r ed ( A l tuna 1 978 ) . T h i s s i tua t ion i s m a in ta ined a t E l P endo , w ho se l e s s mod er n e xcava t ion ( 1 953 -57 ) d i d y i e ld r o den t , i n sec t ivo r e , a nd s ma l l f u r b ea r ing m amma l t a xa f r om i t s U ppe r P a leo l i th ic l e ve l s ( Fuen t e s V ida r te 1 980 ) , b u t n one o f t h e se t a xa f r om i t s Mou s t e r ian l e ve l s . L ez e tx ik i a l so y i e ld ed a g r ea t e r a bundan ce a nd d i ve r s i ty o fs ma l l f u r -bea r ing m amma l s f r om i t s U ppe r P a l eo l i th ic l e ve l s ( A l tuna 1 972 ) . T h i s v a r iance w i th in s i t e s b e tween t h e i r M idd le a nd U ppe r P a leo l i th ic l e ve l s w ou ld t e nd t o i n d ica te t h a t t h e se t a xa w e r e n o t e xp lo i ted a s i n ten s ive ly b y C an tab r ian S pa in ' s N eande r tha l i n hab i tan t s . T he se g ene r a l ly l ow l e ve l s o fr ema in sa r ea l so i na c co rdance w i th t h e i n t e rp r e ta t ion t h a t n a tu ra l m o r ta l i ty i nt h e se t a xa o r t h ea c t ion so fn on -human p r eda to r sc au sed t h e i r i n t roduc t ion i n to t h e se s i t e s . T he r e i sn o e v id en ce t os ugge s t t h a t t h e se t a xa w e r e e xp lo i ted b y h om in id s p r inc ipa l ly f o r t h e i r f u r , o t he r t h an t h e i r g ene ra l s u i tab i l i ty f o r t h i s p u rpo se . O the r f a una l n i che s t h a t w e r e u n - o r u nd e r -exp lo i t ed d u r ing t h i s p e r iod i n c lud e g ame b i rd s , mo l lu sc s , a nd f i sh . T he l a ck o f s p ec ia l iz a t ion i nf u r t rapp ing t h e r ef o r ef i t st h eo ve ra l lp a t t er n o f n on spec ia l iza t ion v i s ib le a mong o t he r r e sou r ce s r e p r e sen t ed a r chaeo log i ca l ly . THE E ARLY U PPER P ALEOL ITH IC T he E a r ly U ppe r P a leo l i th ic o f C an tab r ian S pa in ( Au r igna c ian , C ha t e lpe r ron ian , P e r igo rd ian /G rave t t ian ) , w i th t h e a r r iva l o f a na tom i ca l ly mod e rn H omo s ap ien s , m a rk s s omewha t o f as h if t i nf a una l a cqu i s i t ion . T h i s t r an s i t ion a l so m a rk sm a jo rc hang e st h roughou tt h eO ld Wo r ld i ns t one

a nd o s seou s t o o l m anuf a c t u r e , l a nd scape u sage , a nd s ymbo l ic b ehav io r ( Cha se a nd D ibb le 1 987 ; H a r ro ld 1 980 ; K le in 1 989 ; Me l la r s 1 990 ; Me l la r s a nd S t r inge r 1 989 ) . Wh i le t h e s ame s u i t e o fl a rge f a una ( r ed d ee r , h o r se , b i son a nd i b ex ) w e r e t h e p r ima ry p r ey s p ec ie s , s ome w iden ing o f t h e f o od b a se h a s b een d e t ec t ed ( F r eeman 1 973 ; S t rau s 1 977 ) . T h i s s h if ti s mo re a ppa r en t a mong s ma l l f u r -bea r ing m amma l t a xa , w he r e t h e re i sa n i n c r ea se i n q uan t i ty a nd d i ve r s i ty o fr ema in sr e cove r ed f r om t h e se l e ve l s . S eve ra l t a xa a r e r e cove r ed f o r t h e f i r s t t ime i nC an tab r ian U ppe r P a leo l i th ic a r chaeo log ica l c on tex t s ( Tab le 9 . 2 ) . Mu s te l id s b eg in t o a s sume s ome s i gn if i cance a mong t h e se a s semb lage s . Mu s te la n i va l i s i s p r e sen t i n E l C a s t i l lo ( Au r igna c ian ) L eve l 1 6 ( Cab r e r a 1 984 ) , L ez e tx ik i ( Au r igna c ian ) L eve l I Va ( A l tuna 1 972 ) , Ama lda ( P e r igo rd ian ) L eve l V ( A l tuna 1 990 ) a nd t h e A u r ignac ian l e ve l o f A i tzb i ta r te ( A l tuna 1 972 ) . Mu s t e la e rm inea i s r e p r e sen t ed i n mu l t ip le A u r igna c ian l e ve l s a tE lC a s t i l lo a nd t h e A u r ignac ian o f A i tzb i ta r t e , a s w e l l a s ( Au r igna c ian ) L eve l 7o f R a scaf io ( A l tuna 1 981) . Mu s te la p u to r iu s a l so a ppea r s a nd i sf ound a tE l C a s t i l lo , L ez e tx ik i a nd Ama lda . S i gn if i can t ly , t h e Mou s te r ian l e ve l s f r om e ach o ft h e se s i t e s d i d n o t y i e ld a ny r e po r t ed r ema in s f r om t h e se t a xa . Mu s t e l id s a r e f u r the r r e p re sen t ed b y 2r ema in s o f Ma r t e s a t L eve l X a ( Cha te lpe r ron ian? ) o f E ka in ( A l tuna a nd Ma r r iezku r r ena 1 984 ) a nd o ne e l emen t f r om L eve l Ma ( Au r ignac ian ) a tL ez e tx ik i ( A l t una 1 972 ) , a nd t h e p r e sence o f Me / e s m e / e s i nt h i s s ame l e ve l a nd a tE l C a s t i l lo . G u lo g u lo i su n ique ly r e p r e sen ted i n aC an tab r ian a r chaeo log ica l c on t ex t a tL ez e tx ik i L eve l I( G rave t t ian /So lu t rean ) w i th a s i ng le i d en t if i ed e l emen t . S i nce w o lve r ine i sb o r ea l / tund ra z one s p ec ie s , t h i s l e ve l m ay b e mo r e l i ke ly o f S o lu t r ean a ff i l ia t ion g i ven t h e c o inc id ence o f t h i s t o o l c omp lex a nd g l ac ia lm ax imum c ond i t ion s . Wh i le p r e sen t i n Mou s te r ian l e ve l s , V u /pe s v u lpe s r ema in s b ecome f r equen t ly r e p re sen t ed , t h ough i n g ene ra l ly s ma l l a moun t s a mong t h e se l e ve l s . F ox h a s b een i d en t if i ed a t S an t imam i t le ( A l tuna 1 972 ) , S an t ian ( A l tuna 1 972 ) , O t e ro ( Gonza lez E chega ray e t a l . 1 966 ) , Mo r in ( A l tuna 1 978 ) , G a tza r r ia ( A l tuna 1 972 ) , E ka in ( A l tuna a nd Ma r r iezku r r ena 1 984 ) , E lC a s t i l lo , L ez e tx ik i , B o l inkoba a nd Ama lda . A tt h i s l a s t s i t e , 1 3 3 r ema in s w e re r e cove r ed f r om ( P e r igo rd ian V ) L eve l V I ( A l tuna 1 990 ) . T h i s l e ve l a l so y i e lded t h e o n ly i n s tance o f a r c t ic f o x ( A lopex l a gopu s ) f r om a C an tab r ian c on tex t . L agomo rph s ( O tyc to lagu s c un icu lu s a nd L epu s s pp . ) a l so c on t inue t os how u p i ns ma l l a moun t s , s u ch a sa t E l C a s t i l lo , Mo r in , R a scaf io a nd Ama lda . Sma l l - a nd m ed ium s iz ed f e l id s ( F e / i s s i l ve s t r i s a nd F e l i s l y nx ) a l so s how u p mo re f r equen t ly , a s a t t h e s i t e s o f E l C a s t i l lo , G a tza r r ia , Mo r in a nd L eze tx ik i . W i th t h e e xcep t ion o f t h e f o x r ema in s f r om P e r igo rd ian L eve l s Va nd V I a t Ama lda ,t h e se s ma l l f u r -bea r ing m amma l t a xa h ave n o tb een r e cove red i nl a rge a moun t s . T h i s p a t t e rn m ay r e f lec t t h e g ene ra l ly l ow u t i l iz a t ion b y E a r ly U ppe r P a leo l i th ic g roup so ft h i sf a una ln i che . T h i sp a t ter nm ay a l so r e f lec t l e s s s t r enuou s e xcava t ion / r ecove ry t e chn ique s , w h ich a r e b i a sed a ga in s t t h e r e cove ry o ft h e se s ma l l r ema in s e ven w he r e t h e se w e r e p r e sen t i n l a rge r n umbe r s . T h i s i n te rp r e ta t ion i ss t r eng thened b y n o t ing t h a t t h e mo s t r e cen t 1 39

e xcava t ion s w e r e t h o se o f Ama lda c a r r ied o u t f r om 1 979 1 984 ( A l tuna e ta l . 1 990 ) . H oweve r ,t h e moder ne xcava t ion s o f Mo r in ( Gonza lez E chega ray a nd F r eeman 1 978 ) y i e ld ed

e xpan s ion i nt h i s o r a ny o t he r a r ea . As h if tt os ma l le r f u r b ea re r s m ay t h e r ef o re h ave b een i n p a r t ia l r e spon se t o u nava i lab i l i ty o f l a rge r s p ec ie s . T he se s ma l le r s p ec ie s a r e t o day f a r m o r e a va i lab le i n E u rope t h an t h e i r l a rge r c oun te rpa r t s ( Co rbe t a nd O venden 1 982 ) a nd i n h i s to r ica l t i me s w e re t h e o ne s t h a t m ade u p t h e b u lk o ft h e f u r t r ade ( Youngman 1 982 , 1 990 ) . N e i the r m ay o ne d i scoun t t h e s t a tu s a nd /o r d eco ra t ive v a lue o f p r eda to r s k in s , g i ven t h e f r equen t u s e o f c a rn ivo r e t e e th a s p endan t s ( Co r chön R od r iguez 1 986 ) a nd o t he r s ymbo l ic a s soc ia t ion s w i th p r eda to r s . E xamp le s o ft h e l a t te r c a t ego ry i n c lude p a r ie ta l a r two rk d ep ic t ing p r eda to r s a nd t h e c a rved f e l ine /human h ead f r om E l J uyo ( F r eeman e t a l . 1 988 ; F r eeman a nd G onza lez E chega ray 1 98 1) .

o n ly 1 0 r ema in s a mong t h e se t a xa f r om i t s E a r ly U ppe r P a leo l i th ic l e ve l s ( A l tuna 1 978 ) . T he se d epo s i t s w e r e w e t s c r eened w i th a2mm m e sh , w i th s ome s amp le s f l oa t ed f o r t h e r e cove ry o f f i ne r ema in s . L ow l e ve l s o f s ma l l f a una r ema in s m ay t h e r ef o r e a c cu ra te ly r ef l ec t l ow l e ve l s o f e xp lo i ta t ion i n a t l e a s t s ome c a se s . Wh i le t h e i n c r ea sed d i ve r s i ty a nd a bundance o f t h e se t a xa d oe s s ugge s t t h e i r e xp lo i ta t ion b y h uman s i n s t ead o f o t he r p r eda to r s , t h e i r e xp lo i ta t ion m ay h av e b een a r e su l t o f r a ndom e ncoun t e r h un t ing a nd n o to f at r ue s h if tt owa rd i n ten s ive u t i l iza t ion . W i th o ne e x cep t ion , t h e i r u se s p ec if i ca l ly a s f u r -bea r ing m amma l s c anno tb e d e te rm ined f r om t h e c u r r en t d a ta , g i ven t h e l ow p e rp a r t a bundance s . T he f o x s amp le f r om Ama lda L eve l V I i sl a rge e nough t oe xam ine m ean ingf u l ly i nt e rm s o fp a r t r e p r e sen ta t ion . T he se r ema in s , w h i le i n c lud ing 3 2 ( 2 4 .1 % ) 1 s t , 2 nd a nd 3 r d p ha lange s , a l so i n c lude a w ide s p r ead a mong a l b ody p a r t s w i th c r an ia l e l emen t s , v e r t eb ra e a nd r i b s , a nd l o ng b one s r e p r e sen t ed ( A l tuna 1 990 :174 ) . T hu sa no v e r abundance o ff oo te l emen t s i so n ly s ugge s t ed i n t h i s c a se . H oweve r , t h e p r e sence o fm u s te l id r ema in s ( t he m ea t o fw h i ch i sg ene ra l ly u npa la tab le t oh uman s ) i nE a r ly U ppe r P a leo l i th ic l e ve l s d oe s a r gue f o r f u r e xp lo i ta t ion d u r ing t h i sp e r iod . THE LATE U PPER P ALEOL ITH IC T he L a t e U ppe r P a leo l i th ic o f C an tab r ian S pa in , b eg inn ing w i th t h e S o lu t r ean a nd c on t inu ing w i th t h e Magda len ian a nd A z i l ian p e r iod s m a rk s af u r the r s h if t i n t h e s p ec t rum o f f a una l e xp lo i ta t ion . S o lu t r ean l e ve l s i n C an tab r ian S pa in ( Tab le 9 . 3 ) s h ow ap a t ter n s im i la rt ot h a t o ft h e E a r ly U ppe r P a leo l i th i c , w i th g ene ra l ly l ow n umbe r s o f r ema in s f r om m any s p ec ie s . T h i s p a t ter n m ay i n t h i s c a se b e m o r e a t t r ibu tab le t om e thod s o fa r chaeo log ica l r e cove ry , g i ven t h e a n t iqu i ty o fm o s t o ft h e se e xcava t ion s . Ama lda a ga in i sa n e xcep t ion , t h e U ppe r S o lu t r ean l e ve l ( IV ) o fw h ich y i e ld ed 2 7 V .v u lpe sr ema in s( A l tuna 1 990 ) . T he r e cen tp ro jec ta tL a R ie r a C ave f a i led t o y i e ld l a rge a moun t s o f t h e se t a xa ( A l tuna 1 986 ) , b u t i nt h i s c a se o n ly r emnan t s t ra t ig raphy i n t h e r e a ro ft h e c ave w a s a va i lab le f o re xcava t ion ( S t rau s a nd C la rk 1 986 ) . V . v u lpe s c on t inue s t o s how u p i n s ma l l a moun t s a tn ea r ly a l s i t e s , a l ong w i th f e l id s a nd l a gomo rph s i n s ome l e ve l s . A l so s i gn if i can t i st h e S o lu t r ean l e ve l o f A i tzb i ta r t ew i th t h et a xa Mu s te la n i va l i s ,M . e rm inea , a nd M . p u to r iu s( A l tuna 1 972 ) . T he L ow e r a nd U ppe r Magda len ian l e ve l s f r om C an tab r ian S pa in ( Tab le 9 . 4 ) s how am a rked i n c r ea se i nt h e i n t en s i ty o f u t i l iz a t ion o ft h e se t a xa a sw e l la sd ef in i t ei n d ica t ion st h a tf u r a cqu i s i t ion w a s as p ec if i co b jec t ive . T h i sa ppa r en tc hang e i n f a una l e xp lo i ta t ion m ay a l so b e e xp la ined i nt e rm s o f t h e i n c r ea s ing s c a r c i ty o f l a rge c a rn ivo r e s , w h ich h ad b een i n d i r ec t c ompe t i t ion w i th h uman s f o r d enn ing c ave s , p r ey s p ec ie s a nd o ccupa t iona l r a nge . Ah ypo the t ica l d ec l ine o r l o ca l / to ta l e x t inc t ion t h roughou t t h e U ppe r P a leo l i th ic o f s p ec ie s s u ch a s c av e b ea r ( U r su s s pe laeu s ) , c ave l i on ( Pan the ra l e o ) o r l e opa rd ( P . p a rd u s ) a s w e l l a s t h e m o r e c ommonp la ce w o lf a nd E u ropean b rown b ea r w ou ld ( s eem ing ly i n ev i tab ly ) a c company h uman p opu la t ion 1 40

T he s ame s p ec ie s e xp lo i t ed d u r ing t h e S o lu t r ean p e r iod c on t inue t ob e i mpo r tan t d u r ing t h e Magda len ian p e r iod . V . v u lpe sa ppea r s i nl a rge a moun t sa tU r t iaga ( A l tuna 1 9 72 )a nd R a scaho ( A l tuna 1 98 1) . S ke le ta l p a r t r e p r e sen ta t ion a tt h e f o rme r s i t e i n d ica te s al a rge p ropo r t ion o ff oo t e l emen t s , b u t t e e th a r e a l so h i gh ly r e p r e sen t ed ( A l t una 1 972 :246 ) . R ema in ing e l emen t s a r e g ene ra l ly p r e sen t i nl owe ra moun t s . T h i s p a t te rn o f b one r e p r e sen ta t ion r ef le c t s t h e r e la t ive s u rv ivab i l i ty o ft h e i n d iv idua l e l emen t s , w i th t e e th a nd t h e s ma l l , d en se f o o t e l emen t s p r edom ina t ing . N o o ve r r e p r e sen ta t ion o f f oo t e l emen t s c an b e a s se s sed h e r e w i th c e r ta in ty . F ox s k e le ta l p a r tr e p re sen t a t ion a tR a scaho s h ow s as l igh t e mpha s i s u pon f o o t e l emen t s , t h ough t h e r ema inde r o ft h e s ke le ton i sr e p r e sen ted w e l l e nough t o i n d ica t e t h a t w ho le i n d iv idua l s a t l e a s t a tt ime s w e r e i n t rodu ced t ot h e s i t e . I n ( I n i t ia l Magda len ian ) L ev e l 5o f R a scaho , f o o t e l emen t s m ake u p 1 4 o f2 1 ( 66 .7% ) i d en t if ied f o x s k e le ta l e l emen t s : 1c a lcaneu s , 1t a lu s , 8m e tapod ia l s a nd 4 f i r s t p ha lange s . I n( Lowe r Magda len ian )L ev e l4 b , f o o t e l emen t s m ake u p 6o f 2 9 ( 2 0 .7% ) i d en t if i ed e l emen t s ( A l tuna 1 98 1 :255 ) . T he n a tu ra l d i spo s i t ion o f t h e se s ma l l , d en se e l emen t st os u rv ive i na n i d en t if iab le s t a t ew ou ld e xp la in t h i s p a r t r e p r e sen ta t ion p a t t er n w i thou t t h e i r h av ing b een i n t roduced i ng r ea te rn umbe r st oR a s ca f lo . T h i ss i t e s e em st o h ave b een u t i l iz ed a s as p ec ia l iz ed h i gh land i b ex h un t ing c amp , s o o t he r s p ec ia l iz ed h un t ing p r a c t ice s w e r e p e rhap s n o t e n t i r e ly c ompa t ib le w i th t h e m a in s i t e f u nc t ion . Wh i le f o x w a s e xp lo i t ed i n s i gn if i can t n umbe r s a te a ch o f t h e se s i t e s , t h e d i f f e r en t ia l i n t roduc t ion o f f oo t e l emen t s t r an spo r ted i nf u r s , w i th t h e r ema in ing e l emen t s l e f t b eh ind i nt h e f i e ld ,c anno tb ed e t ec t ed . S ma l l mu s te l id s a l so s how a n i n c r ea se i ne xp lo i ta t ion . I n p a r t icu la r , t h e Magda len ian l e ve l o fA i tzb i ta r t e y i e ld ed 3 1 r ema in s o f Mu s t e la n i va l i s ( A l tuna 1 9 72 :276 ) . T h i s a moun t i s p a r t icu la r ly s i gn if i can t g i ven i t s s i z e , t h e a ge o f t h e e xcava t ion ( b egun a new i n1 960 ) , a nd t h e s ma l l s i z e o ft h i s s p ec ie s w i th t h e c oncom i tan t d i ff icu l ty o fi t s r e cove ry . N o f oo t e l emen t s w e re i d en t if i ed , b u t t h e s e e l emen t s o f M . n i va l i s a r e u n l ik e ly t o b e r e t r ieved w i thou t t h e u s e o fv e ry f i ne me sh s c r een s . O ne s hou ld a l so n o t e t h a t t h e s ma l l s i z e o ft h i s s pec ie s m ake s t h e t r an spo r t o fi t se n t i re c a r ca s s t oa s i t e ,o n ly t od i spo se o f mo s to fi t sm a s s , l e s sc umbe r some . A s im i la r s i tua t ion i s e ncoun te red a t ( Lowe r Magda len ian ) L eve l V o f E r r a l la , w he r e 1 35 r ema in s o f M . n i va l i s w e r e r e cove red , i n c lud ing n o f oo t e l emen t s ( A l tuna a nd Ma r iezku r r ena 1 985 :112 ) . T he se m o r er e cen te xcava t ion s a t

E r ra l la ( c onc luded i n 1 983 ) p e rhap s n o t c o inc iden ta l ly h ave p r oduced t h e l a rge s t s amp le o f t h i s s p ec ie s f r om a C an tab r ian P a leo l i th ic o ccupa t ion t od a t e . O ne mu s t a lway s s u spec t t h a t t h e a b sence o r l ow r e cove ry o f t h e s ma l le r c a rn ivo ran t a xa f r om c e r ta in o ccupa t ion s r e su l t s f r om t h e a n t iqu i ty o ft h e i re xcava t ion .

T ab le 9 . 5 .2 : P o s tc ran ia l R ema in s o fF u r -Bea r ing T axa f r om E lJ uyo . L eve l

T ab le 9 . 5 .1 : T ee th o fF u r -Bea r ing T axa f r om E lJ uyo . M u s t e la n i va l i s L eve l 1 2 1 8 6 4 +4S

Mu s te la p u to r iu s

V u lpe s v u lpe s

O tyc to lagu s c un icu lu s

1 0 6 3 4 9

2

4 1 5

1

I d en t if i ca t ion o f t h e se m ed ium c a rn ivo r e f o o t e l emen t s i s e x t r eme ly d i f f icu l t , s o mo s t c an o n ly b e a s s igned t o a p r obab le t a xon b a sed u pon mo rpho logy a nd g ene ra l s i z e c l a s s . T he m a jo r i ty o f t h e se r ema in s c ome f r om t h e Mu s t e l id a e f am i ly , f a l l ing i n to t h e s i z e r a nge o f Mu s te la p u tor iu s( f e r r e t ) w i th s ome s ma l le rM e rm inea ( e rm ine ) a nd l a rge r Ma r t e s ( ma r ten ) . S pec if i c i d en t if i ca t ion c anno t b e m ade i nm o s t c a se s , s i nce Mu s te la l u t r eo la ( m ink ) f a l l s i n to t h e s ame s i z er a nge a sM p u to r iu s . C onf u s ion w i th t h e f oo t e l emen t s o fC an idae o r F e l idae i sv e ry u n l ike ly d ue t os i z e a nd m o rpho log ica l d i f f e rence s . T he f o o t e l emen t s o f E r inaceu s e u ropaeu s a r e d i s t inc t ive m o rpho log ica l ly f r om t h o se o fm ed ium c a rn ivo r e s pec ie s , s o c onf u s ion w i th t h i s s p ec ie s ' r ema in sm ay b er u led o u t( Chap t e r3 ) . T he U pp e r P a leo l i th ic o f C an tab r ian S pa in c l o se s w i th t h e A z i l ian p e r iod ( Tab le 9 . 6 , e nd o fc hap t e r ) . I na dd i t ion t ot h e u sua l s p ec t r um o f s pec ie s G ene t ta g ene t ta a ppea r s f o r t h e f i r s t t i me , i nL eve l Io f E l P endo ( F uen t e s V ida r t e 1 980 ) . S i nce t h e m od er n n o r ther nmo s t r a ng e o f t h i s g ene ra l ly A f r ican s p ec ie s e x tend s t oI b e r ia a nd F r an ce , i t s a ppea rance i nt h eA z i l ian p robab ly r e f lec t sc l ima t e a me l io ra t ion a nd t h e

S i z eC la s s /Taxon

1 2 : (7 1)

r i gh tf r on t 1 "p ha lanx ( I I )

V u lp e sv u lpe s

1 0 : (7 1)

m id shaf tp ha lanx

c f . Mu s t e la p u to r iu s

9 : ( n= 4 )

m e tapod ia lf r agmen t 3 "p ha lanx 2 'p ha lanx r i gh tf r on t2 "p ha lanx I

c f . Mu s t e la e rm inea c f . Mu s t e la p u to r iu s

3 "p ha lanx p r ox . 1 " /2" p ha lanx Vp ha lanx l e f tm e tapod ia l ( I ) 3 'p ha lanx ( 8 )2 'p ha lanx ( 4 ) Vp ha lanx r i gh t l 'p ha lanx ( I ) 1 "p ha lanx ( 1 /1 1 ) ( 2 )l e f tf r on tm e tapod ia l ( I ) r i gh tf r on tm e tapod ia l ( I / I I I / IV ) ( 2 ) 1 "p ha lanx c a rpa lf r agm en t r i gh tf r on tm e tapod ia l ( I ) I "p ha la nx ( I / I I I ) r i gh tf r on t 1 "p ha lanx ( V ) r i gh tf r on t2 "p ha lanx ( I I / IV ) u nf u s ed 2 'p ha lanx

c f . Mu s t e la e rm inea

7 : ( n= 1)

2 "p ha lanx

c f . Mu s t e la p u to r iu s

6 : ( n= 18 )

p r ox .3 'p ha lanx ( 2 )2 "p ha la nx 3 "p ha lanx ( 3 ) rdp ha lanx l e f tf r on t 1 "p ha lanx ( 1 ) 1' p ha lanx Vp ha lanx ( I / I I / IV ) u nf u sed p ha lanx f r ag . ( 2 )c a rpa l s r i gh tp r ox . r a d iu s ( 1 )2 "p ha lanx f r ag . 1 "p ha lanx ( I I / IV ) r i gh tf r on tm e tapod ia l ( I ) 1 "p ha lanx

c f . Mu s t e la e rm inea

( 3 )3" p ha lanx d p ha lanx 1 "p ha lanx ( 3 )3 "p ha lanx ( 3 )2 " "p ha lanx f r ag . V /2" p ha lanx f r ag p ha lanx r i gh tf r on tm e tapod . ( I V ) r i gh t Vp ha lanx ( I ) r i gh t Vp ha lanx ( V ) r i gh tp r ox . u nf u s ed f emu r

c f . Mu s t e la e rm inea

8 : ( n1. 3 0 )

T he o ngo ing e xcava t ion s a tE lJ uyo , t h rough t h e u se o fv e ry f i ne m e sh s c r e en s ( 0 .25 mm ) h ave a lowed t h e r e cove ry o f m u l t ip le f o o te l emen t so ft h e se t a xa ( Tab le s 9 . 5 .1 a nd 9 . 5 .2 ) . T h i s p a t t e rn o fr e cove ry i ss t r ik ing i ni t sc on t ra s t t ot h e n ea r c omp le te l a ck o fo t he r e l emen t s f r om t h e se t a xa , i n c lud ing t e e th . D if f e r en t ia l s u rv ivab i l i ty t h e r ef o r e i s a n u n l ike ly e xp lana t ion f o rt h i s p a t t e rn . T h i s i n t e rp re ta t ion i sr e inf o r ced b y c ompa r i son w i th r e cove ry o f h edgehog ( E r inaceu s e u ropaeu s ) , as p ec ie s f o r w h ich m any t e e th w e r e r e cove r ed a s w e l l a s n on -f oo t p o s tc ran ia l e l emen t s ( Chap te r 2 ) . H edgehog f a l l s i n to t h e s ame s i z e r a nge a s t h e se s ma l le r c a rn ivo r an s p ec ie s , y e t d ue t o i t s s p iny p e lage i tc anno t b e u t i l iz ed f o r f u r . S ince t h e s k e le ton s o f h edgehog a nd t h e o t he r t a xa d i s cu s sed h e r e a r e e qu iva len t ly s u scep t ib le t o d e s t ruc t ion , p o s tdepo s i t iona l p roce s se s c anno tb e b l amed f o r l o s so fh ypo the t ica lc a rn ivo ran t e e th a nd n on -f oo tp o s tc ran ia l e l emen t s . T h e se e l emen t sp robab ly w e r en eve ri n t roduced t o E l J uyo , t h e m o s t l o g ica l e xp lana t ion f o r w h ich i st h e i r a bandonmen ta tt h es i te o fk i l l ing /p roce s s ing .

E lemen t

4 / 4 s : ( n= 17 )

r

4 4

V u lp e sv u lp e s

c f . Mu s t e la p u to r iu s

t t

c f . Mar t e s s ma l lc a rn ivo r e 4 6

V u lp e sv u lpe s 4 4

t 6

4 4

c f . Mu s t e la p u to r iu s i t

4 4

l t

‘ , 4 4

c f . Ma n t e s V u lp e sv u lpe s c f .F e l l ss i l v e s t r i s

6 4

t 6

c f . Mu s t e la p u to r iu s

4 4

V u lp e sv u lpe s O ry c to lagu sc un icu lu s

T o t a l : n =72

r e tur n o f a va i lab i l i t y o f t h i s s p ec ie s r a the r t h an a s h if t i n f a una le xp lo i ta t ion . R a the rt h an a n i n c rea sed i n ten s i ty o ff u r a cqu i s i t ion b e ing i n d ica ted , t h e p a t t e rn o ff a una l r e cove ry i s mo r e s im i la r t o t h e S o lu t r ean t h an t o t h e Magda len ian p e r iod . Wh i le Me l e s m e l e s ( b adge r ) i sr e p r e sen ted b y 3 0 1 4 1

r ema in s a tU r t iaga ( A l tuna 1 972 ) a nd 1 8 r ema in s a tE ka in L eve l s I I t o V ( A l tuna a nd Ma r r iezku r r ena 1 9 84 ) , f a una l r e p re sen ta t ion i so t herw i se s p a r se . V .v u lpe s a ppea r s i nl ow n umbe r s a tm any s i t e s : S an t imam if i e , U r t iaga , E lP endo , L a R ie ra , E ka in , a nd A i tzb i ta r t e . S ma l le r a moun t s o f f e l id s , m u s t e l id s a nd l a gomo tph s a r e a l so p r e sen t . T h i s l e s sened r e t r ieva l o f f u r -bea r ing s p ec ie s f r om A z i l ian l e ve l s m ay r e f lec t , a s i n t h e S o lu t rean , t h e a r cha eo log ica l r e cove ry m e thod s e mp loyed a tm o s t o ft h e e xcava t ion s . Ar e a l d e e mpha s i s o ff u r e xp lo i ta t ion w ou ld a l so f i tw i th t h e g ene ra l c l ima t ic a me l io ra t ion o ccu r r ing d u r ing t h i s t e rm ina l P l e i s tocene t o o l i n du s t ry . I n ac l ima t e m o r e t empe ra te t h an g l a c ia l S pa in f u r s w ou ld n o l o ng e r b e a s n ece s sa ry , a nd c l o th ing c ou ld h ave b een m ade e n t i r e ly f r om t e x t i le s o rt h e h i de so fl a rge g ame a n ima l s . HUNT ING /TRAPP ING METHODS Many m e thod s c ou ld h ave b een e mp loyed t oa cqu i r e t h e se s p ec ie s , s i nce t h e i n genu i ty o f U ppe r P a leo l i t h ic p eop le s s e em s l i m i ted o n ly b y t e chno logy i no t he r a r ea s o f f a una l a cqu i s i t ion . T he se a r ea s i n c lude t h e i n ven t ion o f t h e b ow a nd a r row . A n i n nova t ion o f t h i s i mpo r tance c ou ld h ave s p read f r om o ne p a r t o fE u rope t oa no the r i n as ho r t p e r iod o ft i me ( Be rgman 1 993 ) . T he C an tab r ian r e g ion i t s e lf d oe s h ave s ma l l S o lu t r ean s t one p o in t s a nd s ma l l a n t le r p o in t s s u i tab le f o r u se a sa r row a rma tu re s , s u ch a st h o se f r om t h e Magda len ian o ccupa t ion so fL aP a loma ( Ma r t inez a nd C hapa 1 980 ) , R a sca f io ( Ba rand ia rän 1 98 1) a nd E l J uyo ( Ba r and ia rän 1 987 ; P ok ine s 1 993 ) . I nt e rm so fh un t ing l a rge g ame , av a r ie ty o fp r o jec t i le t i p s ( Pok ine s 1 99 3 ) a sw e l l a s d r ive a nd s u r round t e chn ique s ( K le in e ta l . 1 98 1 ; F reeman 1 98 1 ; S t rau s 1 977 , 1 987a , 1 987b , 1 992 ) w e re a lmo s t c e r ta in ly e mp loyed . P ro jec t i le i n nova t ion a l so e x t ended t o r e a r -f a c ing b a rbed p o in t s d u r ing t h e S o lu t r ean a nd L owe r Magda len ian ( Pok ine s a nd K r upa , i n p r ep . ) a nd t r ue h a rpoon sd u r ing t h e U ppe r Magda len ian a nd A z i l ian p e r iod s ( Co r chön R od r iguez 1 986 ; Mou r e R oman i l lo 1 9 75 ; Mou re R oman i l lo a nd C ano H e r r e ra 1 976 , 1 978 , 1 979 ) m o s t l i ke ly u s ed f o r f i sh ing . T he r ef o r e , t h e a b i l i ty o ft h e se p eop le s t o c on t r ive e ff i c ien t m e thod s f o r f u r h un t ing a nd t r app ing i s u nque s t ioned . Wha t t h e se m e thod s w e r e , h oweve r , i ss ub je c t t o d eba t e . T he se t a xa p r e sen t t h e s imu l taneou s p r ob lem s i n t h e i r a cqu i s i t ion o f r e la t ive ly s ma l l t a rge t s i z e , t h e a b i l i ty t o d ef end a nd c oncea l t h em se lve s , a nd g ene ra l ly n oc tu rna l a c t iv i ty p a t t e rn s i nw h ich t h e i rs e n so ry a b i l i t ie sg i ve t h em a n a dvan tag e o ve r h uman s . T hey a l so r e p r e s en t as ma l le r p ayof fi nt e rm s o fu sab le m ea t o rh i de t h an l a rge rp r ey s u ch a sr e d d e e r , h o r se , o ri b ex . T he l a rge s p ea r p o in t s t y p ica l t o U ppe r P a leo l i th ic o ccupa t ion s w ou ld b e u n su i tab le f o r u s e u pon a l l b u t t h e l a rge s t o f t h e se t a xa ( f oxe s , b adge r s a nd l y nx ) , t h ough a r row s i fu s ed c ou ld h ave f u nc t ioned i nt h i s

1 42

r o le . S ma l le r t a xa r ema in mo r e p r ob l ema t ic . A l o g ica l e xped ien t w ou ld h ave b een t h e e mp loym en t o ft r ap s , w h ich o pe ra te i n dependen t ly o f h uman a c t iv i ty a nd t h u s c an a cqu i r en oc t ur na ls p ec ie s . H igh s p ec ie sd i ve r s i ty a l so a r gue s f o r t h e e mp loymen t o f t r ap s , i n t h a t m any t y pe s o f s ma l l c a rn ivo r ew ou ld b ea t t ra c ted t ot h es ame b a i ta nd c ap tu r ed i n t h e s ame m anne r . Mo r ta l i ty w ou ld e x t end a c ro s s ar a nge o f s p ec ie s , s i nce s p ec ia l iza t ion u pon o ne w ou ld p rove d i ff i cu l t . T he n umbe ro fp e r son -hou r s n ece s sa ry t oa cqu i re t h e se p r ey i t em s i sa l so g r ea t ly r e duced , s i nce o nc e p l aced t h e t r ap s n eed b e c hecked o n ly o nce p e r d ay . A du l t l a bo r l o ad m ay h ave b een f u r the r r e duced , s i nce c h i ld r en c ou ld h ave m a in ta ined t r ap l i ne s w i th l i t t le s u p e rv i s ion ( p e r sona l o b se rva t ion ) . T he p a t ter n o ff u r e xp lo i t a t ion d e t ec t ed a tE l J uyo a nd r e f le c t ed a t l a rge i n t h e C an tab r ian U ppe r P a leo l i th ic r e co rd s how s at r end o f i n c rea s ing h uman e xp lo i ta t ive e ff i c iency o ve r t h e P l e i s to cen e e nv i ronmen t ( d i scu s sed i nC hap t e r 1 0 ) . T r ap s c ou ld h ave t a ken m any f o rm s , n on e o f w h ich a r e o rd ina r i ly r e t r ievab le a r chaeo log ica l ly d ue t ot h e i rp l a cemen t a way f r om h ab i ta t ion s i te s a nd p e r i shab le c on s t r uc t ion . S na r e s , p i tf a l l s a nd d eadf a l l s o r a ny v a r ia t ion u pon t h e se m a in t h eme s c ou ld h ave b een e mp loy ed ( Coon 1 97 1) . N ece s sa ry r aw m a t e r ia l s a r e m in ima l ly w ood , h i de t h ong s o r s ome t y pe o f f i be r o r s i new f o r b i nd ing , a nd s t one t o o l s s u i tab le f o rs h a rp en ing s t ake s . I mp lemen t sf o rd i gg ing c ou ld h ave b een i mp rov i sed ( s imp le s t ick s ) , o r t h e m u l t ip le r i b d i gge r s r e t r ieved f r om E l J uyo m ay h av e b e en u s ed i nt h i s r o le ( F reeman e ta l . 1 988 ) . I tr ema in s p o s s ib le t h a tt r ap s a r e d ep ic ted i ns ome o ft h e g eome t r ic mo t if s i np a r ie ta l a r t , b u t u nequ ivoca lr e p r e sen ta t ion sa r e l a ck ing . S UMMARY T he C an tab r ian r e g ion s how s a p a t t er n o f i n c r ea s ing u t i l iza t ion o f s ma l l f u r -bea r ing m amma l s t h roughou t t h e P a leo l i th ic . N ea r ly n on -ex i s ten t i nt h e p oo r ly r e p r e sen ted A cheu lean l e ve l s , t h e se t a xa a pp ea r s po rad ica l ly i n Mou s te r ian l e ve l s . T he E a r ly U ppe r P a leo l i th ic , w i th t h e p r e sumed a r r iva l o fa na tom ica l ly mod er nh uman s , s how s a n i n c r ea se i na bundance a nd d i ve r s i ty o ft h e se t a xa , i n c lud ing f o r t h e f i r s t t i me Mu s te la s pp . S ke le ta l r e p re sen ta t ion o f r ema in s f r om t h i s p e r iod d oe s n o t i n d i ca t e ap a t t e rn o f s k inn ing i n t h e f i e ld a nd t r an spo r t ing f oo t e l emen t s ( p ha lange s , m e tapod ia l s , c a rpa l s / ta r sa l s ) b ack t o c ave s i t e s a l ong w i th f u r s . Af u r the r s h if tb ecome s e v iden t l a t e r i nt h e Magda len ian p e r iod , w he r e s pec ie s a bundan ce i n c r ea se s i n s e ve ra l s i t e s , a nd e v idence o f d i f f e r en t ia l t r an spo r t o f f oo t e l emen t s i n to s i t e s ( E l J uyo ) h a s b een r e cove r ed . T he A z i l ian p e r iod s how s a n a ppa ren td ec l ine i nh un t ed m amma l s p ec ie sd i ve r s i ty a nd a bundance . T h i st r end p e rhap sr e su l ted f r om ad e -empha s i s o ff u r a cqu i s i t ion r e la t ing t ot h e g ene ra l c l ima t ic a me l io ra t ion a tt h ec l o se o ft h eP l e i s tocene .

V Z ) 0 0 C T

V ' ) 0 0 C T

N a )

=

$ . e s E

o c s

c r , , N a . ) ' 2

r q 0 0 C. -

, , 0 0 0 , -

0 . )

C D•

" 1 4 • C ) • L . _ .

o - , , e , ( 9

) ' F d

C )

t o t -, 0 ., -

' E = . 1 ) e u c o

e • l i • Z ` C . > o , i

N c r , -

c e

0

, ,

E a . ) a .

0 r q - 0 .

m r o 0 0 b c h . , 4 . ) c u e t 0 1 5 . -

2

› . .Ei • i sc )

a z ; 3 . .a . ) s = M ) ( › . e C I OX . 'e u C D m u

c o c h c u

< c i )

K i

=

‘ ... < . . ..- 3 g. c i 3)

c e =

r \ 1 N .

c s e

c : = . :=

e =

c ue ,

—

c h

w ) t - . . .

r '' r '1 2= ' 0 3

0,

5 1 ,




( r )

•• • ••

2 2 2

O s ,

e l e u

e l 0 . >

e

e l

c i

1

T ab le 9 . 3 : F u r -Bea r ing T axa f r om S o lu t r ean L eve l s .

2 S g , F 3

e

2 2

%

I .s

Z

G en e t ta g en e t ta

A lop ex l a gopu s

•

2

•

Ma rmo ta m armo ta

F e l l ss i l v e s t r ts

C .d el aM ina

N

4.

0

L U

C o

C O

r e t

S an t imam if t e

—

0.

C O

= — .

00

r 4

V ' )

• •

C ue to d el aM ina

e , 4

0

0 C ) 0 0

r a C O

: E )

. 3

e z

0

b .

z e

c z

0

22 IZ S I 7 ), . 5 .C S 4,

E

e , : . % , , : , .% . , % ) .1 V , .

b e e

e › . ,

' 4) b 4

c u

F e l l ss i l v es in s

T ab le 9 . 4 : F u r -Bear ing T axa f r om Magda len ian L eve ls .

2

-

••••

4 -

1 47

› e.

r • 1

s . r s

4 •

( " • . 1

e u e n

o o

0, 0

r a

e n

0 0

c o

0 C . ) —

4 . ) V D

.r e

T i

e n

0

e Ü ' : z. c i ä e

e

. " 2 , .e e

_ • ) 4

e e . .

O

t e i . 2. 2 .

q .

e

V

e e u

21

e e

e e

e

e

t '° " • '

F e !

e C s

G en e t ta g en e t ta

T ab le 9 . 4 : ( Con t . ) .

e n

T ab le 9 . 4 : ( Con t . ) .

. 0

e

eeee

U

e

b

.. I . , 2 2

e

G en e t ta g en e t ta

T i to B u s t i l lo

' — '

z

0 .

c . )

r e

1 49

T ab le 9 . 6 : F u r -Bea r ing T axa f r om A z i l ian L eve l s .

. 2 t o

2 2 2

. 3 t o

e t

F e l l ss i lv es t ra

G ene t ta g ene t ta

Me t e sm e t e s

S an t imam iße

0 0 t o n e l »

e s 4

0 0

l e ». 1 1

t ' s

1 50

CHAPTER 1 0 HU MAN ENV IRON MENTAL I NTERACT IONS

Uppe r P a leo l i th ic h uman -env i ronmen t i n te ra c t ion i n C an tab r ian S pa in e xh ib i t s a n a c ce le ra t ing p a t te rn o fs epa r a t i on o fh uman a c t ion s f r om t h e i mmed ia te c on s t ra in t s o fe n v i ronmen t . T he se c hange s a r e v i s ib le a se a r ly a st h e E a r ly Uppe r P a leo l i th ic ( Au r igna c ian , C ha t e lpe r ron ian , G rave t t i an /Upp e r P e r igo rd ian ) a nd g r ea t ly i n c r ea se t h roughou t t h e L a te U pp e r P a l eo l i th ic ( So lu t r ean , Magda len ian a nd A z i l i a n ) . T h e se c hange s a r e mo r e r ema rkab le c on s ide r ing t h e ma jo r a nd m ino r f l uc tua t ion s o f t h e c l ima te . T he se p r o c eeded f r om v a c i l la t ing i n t e r s tad ia l c ond i t ion s , t h rough t h e g l a c ia l m ax imum c a . 2 2 ,000 18 ,000 B .P . a nd t h e s t epw i se r e t r ea t / readvance o fg l a c ie r s b ef o r e t h e w a rm c l ima te o ft h e H o locen e w a s o b ta ined b y 1 0 ,000 B .P . T he mu l t ip le e nv i r onmen t a l z one s o fc oa s ta l C an tab r ian S pa in a r e a l so s t r ik i n g , w i th l i t to ra l , r i ve r ine , c oa s ta l l ow land a nd mon tane z one s a l w i th in e a sy m u tua l r e a ch . V a r iab le e nv i ronmen t w a s t h u s o b ta inab le b o th o ve r t i me a nd t h rough p hy s ica l t r ave lo v e rt h e l a nd scape . Mou s t e r ian o ccupa t ion o fC an tab r ian S pa in w a s s pa r se a nd l e s s i n t ru s ive u pon t h e l o ca l r e sou r ce s . Wh i le s u rf ace f i nd s a r en ume rou s , Mou s t e r ian s i t e sn umb e rj u s to ve r ad oz en i na r e cen ts umma ry b y S t rau s ( 1 992 ) . T h e se i n c lude l e ve l sa tE l C a s t i l lo ( Cab r e ra 1 984 ) , Ama lda ( A l t una e ta l . 1 990 ) , C ueva Mo r in ( Gonza lez E chega r ay a nd F r eeman 1 978 ) a nd E l P endo ( Gonza lez E chega ray 1 980 ) . F auna l e xp lo i ta t ion f o r t h i s p e r iod i sm o r e o ppo r tun i s t ic t h an s e em s t ob e t h e c a se d u r ing a tl e a s t t h e L a te U ppe r P a leo l i th ic . Wh i le r e d d ee r s how u p i nm o s t Mou s t e r ian l e ve l s a nd a r e o f t en t h e m o s t n ume rou s s p ec ie s , t h e i rn umbe r s a r e s ome t ime s l e s s t h an o r c ompa rab le t oo t he rf o od t a xa . T h i ss i t ua t ion i si lu s t ra t ed a t C ueva Mo r in L eve l 1 7 , w he r e C e r vu se l aphu s ( N ISP = 1 42 ) i sf o und i ns im i la ra moun t sa sl a rg eb ov id s( N ISP =2 15 )a nd E quu s C aba l lu s ( N ISP = 1 24 ) ( A l tuna 1 978 :203 ) . I n Ama lda L ev e l V I I , C er vu se l aphu s( N ISP = 1 50 ) a r e f a ro u t n umb e r ed b y R up i cap ra r up i cap ra ( N ISP = 5 36 ) ( A l tuna 1 990 :151 ) . Wh i le C . e l aphu s i s a bundan t a mong t h e Mou s t e r ian l e ve l s o fA x lo r , l a rg e b ov id s , C ap ra p y rena i ca , a nd E quu s c aba l lu s a l o u tnumbe r i ti n a t l e a s t o ne l e v e l ( A l tuna 1 9 89 c i t ed i nS t rau s 1 992 ) . T he p a t t e rn o f Mou s t e r ian p r ey a cqu i s i t ion i n d ica te s m o r e o ppo r tun i s t ic h un t ing w a s t h e n o rm f o r m any l a te r o ccupa t i on s ( F r eeman 1 973 , 1 981 ; S t rau s 1 9 77 ) , w i th s pec ie st a ken o n a n i n d iv idua l e ncoun te r b a s i s . S u ch e ncoun te r h un t ing i mp l ie s al e s se r n eed f o r g roup c oop e ra t ion : f ewe r h un t e r s a r en e ce s sa ry f o rt h ek i l l ,n o ra r et h e r el a rge a moun t so fm ea t t o p ro ce s s , t r an spo r t a nd s t o r e a to n e t i me . L e s s r ewa rd i s b a lan ced b y l owe r ed r i sk . H un t ing s u cce s s i sb a sed u pon a s e r ie s o ff r equen t e ncoun te r s , a ny o ne o fw h ich may p rove s ucce s sf u l , r a the rt h an l a rge , p l anned g ame d r ive s i n vo lv ing many h un t e r s a tk ey s e a sona l / topog r aph ic a gg r ega t ion s o f a n ima l s .

w he re U r su s s pe laeu s ( c ave b ea r ) , U . a rc to s ( b rown b ea r ) , C an i s l u pu s ( wo l f ) a nd P an the ra s pe laea ( c ave l i on ) a r e n u m e rou s , a nd t h e i r r ema in s s ome t ime s o u tnumbe r t h o se o f mo r e p robab le h uman f o od t a xa ( A l tuna 1 972 :14 1142 ) . T he se e l eva ted l e ve l s o fl a rge c ar n ivo r e s i n d ica t e p e rhap s a l a rge r d eg r ee o f d i r ec t c ompe t i t ion w i th h uman s , b o th f o r c ave d enn ing s i t e s a nd u ngu la te p r ey ( S t ine r 1 993 , 1 994 ) . H om in id s s e em t oh ave a c ted i nw ay s a na logou s t ot h o se o f o t he rl a rge p r eda to r sp r e sen t . T he n a tu r e o f t h e M idd le /Uppe r P a leo l i th ic t r an s i t ion i n E u rope i sa n e no rmou s t o p ic ( Cha se a nd D ibb l e 1 987 ; H a r r o ld 1 980 ; G a rge t t 1 989 ; K le in 1 989 ; Me l la r s 1 990 ; Me l la r s a nd S t r inge r 1 989 ) , o fw h ich o n ly o ne a s pec t i se co log ica l . E a r ly U ppe r P a leo l i th ic C an tab r ian s b egan t os h if t t h i s e co l o g ica l p a t t er n a nd s how i n c r ea s ing c on t ro l o ve r t h e l a nd s c ape . L a rge p r eda to r s n o l o nge r a ppea r i ns u ch g r ea tn um b e r s a mong h uman h ab i ta t ion l e ve l s . S ome s p ec ie s r e a ch n ea r e x t inc t ion d u r ing t h i s p e r iod , i n c lud ing U r su s s pe laeu s a nd P an the ra s pe laeu s ( Ku r t 6n 1 972 , 1 976 ) . O the r s r e a ch n ea r l o ca l e x t inc t ion , s u ch a s P an the ra l e o a nd P .p a rd u s . S ome v e ry l a rge p r ey s p ec ie s a l so d ec l ined t oo rn ea re x t inc t i on , ap a t t er n t h a t w ou ld r e pea t a c ro s s E u ra s ia . T he se i n c l ude n ow e x t inc t f o rm s o fr h inoce ro s ( D i ce ro rh inu s k i r ch b e rg en s i s a nd C oe lod on ta a n t iqu i ta tu s ) , e l ephan t (M ammu thu s p r im ig en iu s a nd E l epha s a n t iquu s ) a nd d ee r (M ega ce ro s g igan teu s ) . A l though d eba t e c on t inue s o n t h e p rox ima te c au se o fm egaf auna l e x t inc t ion s a r ound t h i s t i me , t h et h eo ry o fc l ima te c hange a sc au se h a s l e s sc r edence i nt h e f a ce o f t h e r e gu la r , r a p id f l uc t ua t ion s i nc l ima t e o ccu r r ing t h roughou t t h e P l e i s tocene ( CL IMAP 1 976 ; Mc In ty r e e ta l . 1 976 ; R udd iman a nd Mc In ty r e 1 976 ; S ha ck le ton a nd O p d yke 1 976 ; v an D onk 1 976 ) . S i nce t h e se s p ec ie s r e gu la r ly a dap ted t om a jo ra nd m ino rg l a c ia l f l uc tua t ion s ,t h e i rd em i se o n t h e g round s o ft h e se c hange s a l one i si mp lau s ib le . T h i s i n t e rp r e ta t ion i sr e inf o r ced b yt h e i rd ec l ine a nd i nm o s tc a se s e x t inc t ion i n We s t er n E u rope b ef o re t h e s ub s tan t ia l ly w a rme r c ond i t ion s o ft h e H o locene w e r e o b ta ined . H uman h un t ing ( a nd d i r ec t c ompe t i t ion f o r r e sou rce s i nt h e c a se o f l a rge c a rn ivo r e s )t h e r ef o r e i st h e mo re l o g ica l c au se o ft h e se s pec ie s ' d em i se ( Ma r t in a nd K le in 1 984 ) . T he e v idence f o r C an tab r ian S pa in d u r ing t h eE a r ly U ppe rP a leo l i th ic s ugge s t s t h a t h uman s h ad a s sumed ag r ea t e r i mpa c t u pon t h e e co sy s t em t h an o t he rl a rge p r eda to r s . E a r ly U ppe r P a leo l i th ic s i t e n umbe r s i n c r ea sed o ve r t h e Mou s t e r ian , e ven t h ough amuch s ho r te rp e r iod w a s i n vo lved ( 40 ,000 t o2 0 ,500 B .P . ) . I n ar e cen t s umma ry S t rau s ( 1 992 : 7 5 ) i n d ica t e s a s m any a s 2 6 s i t e s c on ta in ing A u r igna c ian , C ha te lpe r ron ian , a nd /o rG rave t t ian /Uppe rP e r igo rd ian l e ve l s . T he se o ccupa t ion s i n c lude E l P endo ( Gonz a lez E chega ray 1 980 ) ,E lC a s t i l lo ( Cab r e r a 1 984 ) , E ka in ( A l t una a nd Me r ino 1 984 ) , Ama lda ( A l tuna e ta l . 1 990 ) a nd A i tzb i ta r t e ( A l tuna 1 972 ) .

Mou s te r ian l e ve l s a l so t e nd t oy i e ld s ub s tan t ia l n umbe r s o f l a rge c a rn ivo r er ema in s . T h i s i se s p e c ia l ly t r ue o fL eze tx ik i ,

1 5 1

E a r ly U ppe rP a leo l i th ic f a una le xp lo i ta t ion c en t e r ed u pon t h e s ame u ngu la te s p ec ie s a sd u r ing t h e Mou s te r ian p e r iod . T he b a lance a mong l a rge p r ey s p ec ie s i si l u s t ra ted a tL a R ie ra L eve l 1 ,w he re 4 t a xa h ave a n MN I o f 4o r mo r e : E quu s c aba l /u s( 5 ) , C ap ra p y rena ica ( 4 ) ,B o s IB i son ( 4 ) a nd C er vu s e l aphu s ( 7 ) ( A l tuna 1 986 ) . T h i s b a lance a mong l a rge u ngu l a t e s i sa l so i l u s t ra t ed a mong t h e A u r igna c ian a nd G rave t t i an l e ve l s o fC ueva Mo r in , w he r e C e rvu s e l aphu s , C ap r eo l u s c ap reo lu s , B o s IB i son , C ap ra p y rena i ca a nd E quu s c ab a l /u s a l w e r e t a ken i n s ub s tan t ia l a moun t s ( A l tuna 1 978 :203 ) . O the r l e ve l s f r om t h i s t i me p e r iod , h oweve r , s how t h e b eg inn ing s o f mo r e i n t en s ive u s e o f o ne o r t wo u ngu la te s p ec ie s . A t E lC a s t i l lo A u r igna c ian L eve l 1 8 , C e r v u s e l aphu s ( MN ! —2 16 ) a nd B o s ( M IN I =2 9 ) d om ina t e t h e f a una l p r of i le ( Cab r e r a 1 984 :4 14 -4 15 ) . A t B o l inkoba G rave t t ian L eve l Fi sd om ina ted b y t h e r ema in s o fR up i ca p ra r up i cap ra ( N ISP = 6 34 ) , w h i le t h e n ex t h i ghe s t r e p r e s e n ted t a xon i sB o s tB i son ( N ISP = 3 7 ) ( S t rau s 1 992 :262 ) . T h i s t r end w a s t o c on t inue a nd b e g r ea t ly e xpanded u pon d u r ing t h ee n su ing o ccupa t ion o fC an tab r ian S pa in . T he L a t e U ppe r P a leo l i th ic ( So lu t r ean , Magda len ian a nd A z i l ian ) C an tab r ian s c ho se b o th t o h a r ve s t p r ev iou s ly e x p l o i t ed s p ec ie s w i th i n c r ea s ing i n t en s i ty a nd t ob r oaden t h e i r r e sou r ce b a se d u r ing t h i s t ime o f f l uc t ua t ing b u t g ene ra l ly a me l io ra t ing c l ima te t h a t u l t ima t e ly l e d t o t h e H o locene . P opu la t ion a l so s e em s t oh ave e xpand ed , w i th a bou t 4 4 S o l u t rean , 5 2 L owe r Magda len ian , 4 5 U ppe r Magda len ian a nd 3 5 A z i l ian s i t e s ( S t rau s 1 992 ) . T he se s i t e s w e r e a l so a c cu m u la ted o ve r s ho r t e r p e r iod s t h an d u r ing t h e Mou s te r ian o r E a r ly U ppe rP a leo l i th i c : 4 , 000 y ea r sf o rt h e S o lu t rean , 3 , 500 f o rt h e L owe r Magda len ian , 3 , 000 y ea r s f o rt h e U ppe r Mag d a len ian a nd 2 , 000 y ea r s f o r t h e A z i l ian . I n c r ea s ing s p e c i a l iza t ion o fs i t ef u nc t ion w ou ld a l so i n c r ea se t h en umbe ro f s i t e s , b u t t h e i n c r ea se i ss o g r ea t t h a t i tm u s t a l so r ef l ec t p opu la t ion i n c r ea se . T he se f i gu r e s c an o n ly s ugge s t c hang i n g p opu la t ion l e ve l s . S i nce n ea r ly a l k nown P a leo l i th ic s i t e s i nC an tab r ian S pa in a r e i nc ave s , d e s t royed o r u nd i s c ove r ed o pen -a i r s i t e s w ou ld i fa ny th ing r a i se p opu la t ion e s t ima te s . T he se f i gu r e ss uppo r tb o th a nt h e i n c r ea sed i n t en s i ty o fc ave s i t e u s age , g i ven t h e i n c r ea s ed n umbe r o fs i t e s a nd a moun t o f e xcava ted m a te r ia l s , a nd a n i n c r ea se i n p opu la t ion . R ed d ee rw e r et h e m o s ta bundan tl a rge m amma l ian p r ey a nd w e re e xp lo i t ed w i th i n c r ea s ing i n t en s i ty ( F r eeman 1 973 , 1 975 , 1 98 1 ; S t rau s 1 974a , 1 974b , 1 975 , 1 976 , 1 977 ) . T h i s p a t te rn b eg in s i nS o lu t r ean t ime s . I n s u rvey s o f S o lu t r ean f a una s b y S t rau s ( 1 977 , 1 992 ) , r e d d ee r a r e t h e m o s t a bun d an t l a rge m amma l t a ken i nn ea r ly a l s i t e s a nd l e ve l s . T he e xcep t ion s i n c lud e B o l inkoba a nd E rm i t t ia , w he r e i b ex r a nk a head o fr e dd ee ra nd B ux i ta nd Ama lda w he r ec hamo i sr a nk a head . T h i s t r end w a s i n t en s if i ed d u r ing t h e Magda len ian , w hen l a rge k i l l s o f r e d d ee r b ecome m o r e p r onounced . A s d i s c u s sed i n C hap t e r 7 , r e d d ee r d om ina t e t h e L owe r Mag d a len ian l e ve l s o fE lJ uyo . O the rL owe r Magda len ian o ccu p a t ion s w i th e l eva t ed p ropo r t ion s o f r e d d ee r i n c lude A l tam i ra ( S t rau s 1 977 ) , L a P a loma ( Ca s ta f io s 1 980 ) , C ue to d e l a M ina ( Ca s ta f io s 1 982 ) , L a R ie ra ( A l tuna 1 986 ) , E l P endo ( Fuen te s 1 980 ) , U r t iaga ( A l tuna 1 9 72 ) , C ueva Mo r in

( A l tuna 1 978 ) , E ka in ( A l tuna a nd Ma r iezku r r ena 1 984 ) a nd T i to B u s t i l lo ( A l tuna 1 976 ) . A no the r c hange i nt h e f a una l e xp lo i ta t ion p a t ter n i st h e u s e o fs p ec ia l iz ed h un t ing c amp s , a s a tt h e u p land s i te o fR a s c af to , w he r e Magda len ian a nd A z i l ian l e ve l s a r e d om ina t ed b y i b ex ( A l t una 1 981) . C o l lub i l a l so s e em s t oh ave s e rv ed t h i s p u rpo se d u r ing t h i s t ime ( Gonzä lez Mo ra le s 1 974 c i t ed i n S t rau s 1 9 92 :132 ) . I b ex a r e a l so p l en t if u l a to t he r s i t e s s u ch a s E r r a l la ( A l tuna a nd Ma r iezku r r ena 1 985 ) , E rm i t t ia a nd U r t iaga ( A l tuna 1 972 ) ,a nd L aR ie ra ( A l tuna 1 986 ) . A g e e v idence f o r i b ex f r om R a scaf to i n d ica t e s m o s t w e r e a du l t s o ve r 2y ea r s o l d ( A l tuna 1 98 1 :251) . T hough b r eakdown b y a ge c l a s s w a s n o t i n d ica ted , ah igh p ropo r t ion o f a du l t s i s mo r e l i ke ly t or ef l ec t ac a ta s t roph ic k i l l . I b ex w e r e m o s t l i ke ly h a rve s t ed u s ing d r ive a nd s u r round t e chn ique s , s i n ce t h e i r a b i l i ty t oe s cape o ve r r ugged t e r ra in m ake s t h e i r i n d i v i dua l s t a lk ing b y h un te r s w i thou t m ode rn f i rea rm s e x t r eme ly d i ff i cu l t ( S t rau s 1 987b ) . T h i s s p ec ie s a l so c ome s f r om t h e h i gh land a l t i tud ina l z one n o tn o rma l ly o ccup ied b y t h e se h un te r -ga the r e r s , e s pec ia l ly d u r ing e p i sode s o fa l p ine g l a c ia t ion . T h i s c oncen t ra t ed e x t ra c t ive e xp lo i ta t ion o f a m on tane r e sou r ce i n d ica t e s f u r the r i n t en s i t y o f l a nd s cap e u sage . T he m e thod s o f d ee r h a r ve s t a l so s e em t o h ave c hang ed d u r ing t h e L a t e U ppe r P a leo l i th ic i nC an t ab r ian S pa in . T he r e d d ee r t a ken a tL eve l 4 / 4S a nd 6 a tE l J uyo s how ac a t a s t roph ic a ge p r of i le , ap robab le i n d ica t ion t h a td r ive a nd s u r r ound t e chn ique s t a k ing w ho le h e rd s w e r e p r a c t iced ( K l e in a nd C ruz U r ib e 1 987 ; K le in e ta l . 1 98 1) . R andom e ncoun t e r h un t ing t e nd st op r oduce ap auc i ty o fp r ime a ge a du l t s , s i nce t h e se i n d iv idua l s a r e t h e b e s t a b le t oa vo id p r eda to r s , w h i l e t h e o l de s t a nd y ounge s t a r e mo re v u lne rab le . T he t a k ing o f p r ime a ge a du l t s i np ropo r t ion w i th t h e i r n a tu ra l a bundanc e i n ap opu la t ion i n d ica te s t h a t t h e t e chn ique u s ed t oh a r v e s t t h em n eu t ra l iz e s t h e b enef i t s o f t h e i r m o r e r obu s t s t a t e . P r eda to ra vo idance e qua l iz e s a mong i n d iv idua l s i n at r app ed h e rd . L og ica l p l ace s f o r a mbu she s t o o ccu r a r e i n l a k e s , s t r eam s o r w e t land s w he r e mob i l i ty i sl e s sened , i nn a tu r a l t r ap s s u ch a s r a v ine s , o r i n a r t if i c ia l t r ap s s u ch a s p en s . T rapp ing a nd d i spa t ch ing a n e n t i re d ee r h e rd a l so r e qu i r e s a l a rge c oope ra t ing g roup o fh un te r s t ob l o ck a venue s o fe s c ape a nd t op r oce s sa nd t r an spor tt h ek i l l . T h i se mpha s i su pon " w i ld h a rve s t ing "( F r eeman 1 973 :39 -4 1 , 1 975 :254 -255 ) a ts e lec ted s e a sona l t i me s a ppea r s t ob e r e f l ec t ed a s w e l l i nt h e p ro jec t i le t e chno logy o f t h i s p e r iod . T he S o lu t r ean p e r iod i sd ef ined i no ne r e sp ec t b y i t s l a rg e l i t h ic p ro jec t i l e p o in t s f o und i n mu l t ip le f o rm s : l a u r e l l e a f , w i l low l e af a nd s h ou lde r ed . F ound a l ong w i th t h e se p o in t s a r es i gn if i can tn umbe r s o fo va l -p rof i led a n t le rp ro jec t i le t i p s . B yt h e L owe r Magda len ian p e r iod , l a rge l i t h i c p ro jec t i le t i p s h ad d i sappea r ed a nd a n t le r t i p s h ad a s sumed t h i s r o l e . T he d om inan tp rof i le s h ape o ft h e se p o in t s s h if t ed t oq uad rangu l a r , as h ape t h a t i s ah a l lma rk o ft h e C an tab r ian L owe rM ag d a len ian . T h e i n t roduc t ion o ft h e b ow a nd a r row m ay h ave a l so o ccu r r ed a ts ome p o in t i nt h i s p e r iod ( Be rgman 1 9 93 ) . P ro jec t i le t e chno log ie s c hanged a ga in l a t e r w i th t h e a n t l e r h a rpoon f o rm s t h a t p a r t ia l ly d e f ine t h e U pp e r Magda l en ian a nd A z i l ian p e r iod s .

1 52

A n t le rp r o jec t i le t i p s a r e m o r e r e l iab le t h an l i t h ic o ne s i na n e ng inee r ing s e n se ( B leed 1 986 ) . E xpe r imen ta t ion w i th r e p l i ca t ed p o in t s ( Pok ine s 1 993 , i np r e s s ) i n d ica te s t h a t a n t le r p ro jec t i le t i p s c an w i th s tand r e pea ted i mpa c t s w i th m in ima l d amag e . T hey a r ea l so m u ch m o r el i k e ly t ow i th s tand l a t e ra l s t r e s se s a f t e r i mpa c t w h i le b e ing d r agged b y aw ounded a n ima l . R epa i r i sp o s s ib le m o r e o f t en a nd i s much q u icke r , u sua l ly i n vo lv ing o n ly t h e r e sha rp en ing o f ad amaged t i p . A n t le rp r o jec t i le st h roughou tt h eL a te rU ppe rP a leo l i th ic a l so s how ap a t t e rn o f mo r e s y s tema t i c m anuf a c tu r e . Q uad ran g u la r f o rm s c an b e p roduced m o r e q u ick ly t h an o va l f o rm s , s i nce m o r e o f t h e o r ig ina l s h ape o ft h e a n t le r b l ank i sr e t a ined i nt h e f o rme r . E v idence f r om E lJ uyo a l so s how st h a t l o ng b l ank s ( l a rge e nough f o r m u l t ip le p o in t s ) w e r e c u t i n s t r ip s f r om a n t le r , a low ing mo r e e ff i c ien tp ro jec t i le p roduc t i on . T h i s c hange i np r o jec t i le t i p s c omp lemen t s o t he r c hange s i n h un t ing t e chn ique . I fh un t ing s u cce s s i sb a sed u pon af ew i s o la ted , l a bo r i n ten s ive i n s tance s w he r e a n e n t i re h e rd i s d i spa t ched , t h en t h e c on sequence s o f f a i lu r e a r e much g r ea te r . F a i lu r e d u r ing r a ndom e ncoun t e r h un t ing l e ave s s e ve ra lh un t e r si no t he rl o ca t ion sw ho m ay s t i l lb e s u cce s sf u l o n t h a t s ame d ay . G roup d r ive s h ave ag r ea t e r p ena l ty f o r f a i lu r e , s i n ce m o s to f ag r oup s ' r e sou r ce s h ave b een i n ve s t ed i no ne a r ea . P ro jec t i le t i p s t h a t h ave b een mo r e r e l iab ly e n g i nee r ed ( mo r e l i ke ly t ow i th s tand i mpa c t , l e s s l i ke ly t ob e d amag ed a c c iden ta l ly , a b le t o w i th s tand r e pea t ed u se , a nd e a s ie rt or e pa i ri fd amaged )a r ew o r th a ny a dd i t iona l i n ve s ted m anuf ac tu r et i me . A n t le rt i p st h e r ef o r eb ecome amo r e l o g i c a l c ho i c e i n s i tua t ion s w he r e r e l iab i l i ty i sr e qu i red a nd l o nge r p e r iod s o fd own t i me a r e a va i lab le b e tween h un t ing e p i sod e s . A n a dd i t iona l c hange i np r o jec t i le t e chno logy o ccu r r ed i n h a rpoon s t y le s . L a te ra l ly b a rbed h a rpoon h ead s m ade o f a n t le r a ppea r b y d ef in i t ion a tt h e o u t se t o ft h e U ppe r Mag d a len ian p e r iod . T he u sua l f u nc t ion a s s igned t ot h e se t i p s i s f i sh ing , s i nce t h ey s o lve t h e p r ob lem o f s e cu r ing a qua t ic p r ey t h a tm igh t o t he rw i se f l oa t a way i fs p ea r ed w i th a n u n b a rbed p r o jec t i le t i p . T hey a l so a pp ea r w hen e v idence o f f i sh ing t e nd s t oi n c r ea se . H a rpoon s , h oweve r , a r e p r eceded b y a n u nba rbed p ro jec t i le t i p t h a t c ou ld h ave f u nc t ioned i n t h e s ame r o le ( Pok ine s a nd K rupa 1 997 ) . T h i s a n t le r p o in t t y pe , w h ich o ccu r s i n S o lu t r ean a nd L owe r Magda len ian c on tex t s , h a s al a te ra l h af t ing s u rf a ce a nd ac urved r e a rp o r t i on e nd ing i n as h a rp p o in t . When h af t ed l a t e ra l ly t ot h ee nd o f as h af t ,t h er e a rt i pp ro jec t s o u twa rd a nd f o rm s ar e a r f a c i n g b a rb . T he se t i p s f u nc t ion w e l l a sf i sh s p ea r s i n moder n e xpe r im en t s . T he i r p r e sence i n s i t e s p r eda t ing t h e U ppe r Magda len ian p e r iod i n d ica te s t h a t s p ea r f i sh ing m ay h ave g one o n m o r e i n t en s ive ly d u r ing t h i s e a r l ie r p e r iod . T h i s i n te rp re t a t ion i ss uppo r t ed b y n ew e v idence f r om E l J uyo , w he r e v e ry a bundan t f i sh r ema in s o ccu r i n aL owe r Mag d a len ian c on t ex t . Mo r ei n t en s ive " w i ld h a rve s t ing" o fd en se , s e a sona l r e sou r ce s m ay h ave o ccu r r ed e a r l ie r i nt h i s p e r iod ( Dav id son 1 976a , 1 976b , 1 983 ) . L and scape u sage c hanged i no t he r m anne r s . R e sea r ch b y C onk ey ( 1 978 , 1 980 ) o n e ng rav ed m ob i le a r t mo t if s f r om L owe r M agda len ian s i t e s s ugge s t s t h a t s ome s i t e s may h ave s e rved a ss e a sona l a gg r ega t ion l o c i f o r b and s t h a t r ema ined

d i spe r sed f o rt h e r e s t o ft h e y ea r . A l tam i ra a nd C ue to d e l a M ina a r e t wo s i t e s i d en t if i ed f o r t h i s p o s s ib le r o le . T he i r mob i le a r th a s ag r ea t e r v a r ie ty o fe ng raved mo t if st h an s u r r ound ing s i t e s , s ugge s t ing t h a t p e r iod ic v i s i to r s f r om m any p a r t s o f t h e s u r round ing r e g ion l e f t t h e se b eh ind . T ha t s ma l le r n ea rby s i t e s h ave l owe r d eco ra t ive d i ve r s i ty s h e i n t e rp re t s a sr ef l ec t ing t h e f a c tt h a tt h ey w e r e e xc lu s ive ly u s ed b y o ne o r af ew g roup s w ho se d eco ra t ive r e pe r to i r e s w e re l i m i ted . T h i sp a t t er nm ay ,h oweve r ,a l so b ee xp la ined b yt h e l ump ing t o g e the r o fo l d e xcava t ed c o l lec t ion s f r om m u l t ip le l e ve l s a t p r opo sed " agg r ega t ion s i t e s" w ho se d epo s i t ion s pan s mo r e t i me ( S t rau s 1 980 ) . T empo ra l i n s tead o fs pa t ia l s t y l i s t ic d i ve r s i ty w ou ld t h u s b e r e f lec t ed . T he s e a sona l a g g r ega t ion h ypo the s i s i sr e inf o rced a tl e a s ta tA l tam i ra b yt h a t s i t e ' s s t a tu s a s am a jo r p a in ted c av e w ho se mo t if s a r e c en t e r ed u pon m a t ing b i son ( F reeman 1 987 ) a nd t h rough g eo g r aph ic a na ly s i s o ft h e r e g ion ( F reeman 1 994 ) . T he t empo r a ry j o in ing o f s ma l l d i spe r sed b and s f o r t h e p u rpo se s o f h un t ing m a s sed h e rd s o f u ngu la te s m e she s w i th t h e o t he r e v idence o fh un t ing b ehav io r i n t e rp r e t ed h e r e . A t A l tam i ra t h e re i sa l so e v idence t h a t Magda len ian o ccupa t ion sw e re n o t e s pec ia l ly l o ng la s t ing ( Gonza lez E chega ray 1 988 ; F r eeman 1 988 ) . F u r the r a na ly s i s o fn ew ly e xcava t ed m a t e r ia l s w i th b e t te rt empo ra lc on t ro l sm ay p rov id e ac onc lu s ive r e so lu t ion . Ma r ine mo l lu s c s w e r e a l so s ub jec t t oi n c r ea s ing ly s e lec t ive e xp lo i ta t ion t h roughou tt h i sp e r iod ( C la rk 1 983 ; K rupa 1 994 ; S t rau s e ta l . 1 98 1) . Wh i le o fe x t r eme ly l i m i ted d i e ta ry i m p o r tance d u r ing b o th Mou s te r ian a nd E a r ly U ppe rP a leo l i th ic o ccupa t ion s ,t h e se a s sume as i gn if i can tr o le d u r ing t h e S o lu t r e an a nd w e r e a c cumu la t ed i nt r ue s h e l l m idden s b eg inn ing w i th t h e L owe r Magda len ian . T he m a jo r o ccupa t ion l e ve l s a tE lJ uyo ( 4 , 6 ,a nd 8 )h ave e a ch y i e lded i ne xce s s o f8 , 000 w ho le s he l l s a nd m any t i me s t h a t q uan t i ty o ff r agmen t s , a l mo s t e n t i r e ly o f l i mpe t s ( Pa te l la v u lga ta ) a nd p e r iw ink le s ( L i t to r ina l i t to rea ) . S i gn if i can t q uan t i t ie s h ave a l so b een f ound a ts i t e s s u ch a sA l tam i ra ( S t rau s 1 976 -77 ) , E l C a s t i l lo ( Cab r e ra 1 984 ) , a nd O t e ro ( Gonza lez E chega r ay e t a l . 1 966 ) , a nd T i to B u s t i l lo ( Mou r e R oman i l lo 1 975 ) d u r ing t h e U ppe r Magda len ian . T h i s t r end w a s t o a c ce le ra t e , c u lm i n a t ing i nt h e t r ue c onche ro s o f t h e A s tu r ian p e r iod , ar e g i ona l man if e s ta t ion o f t h e Me so l i th ic ( C la rk 1 97 1 , 1 983 ; S t rau s 1 99 1a ) . A s d i scu s sed i nC hap te r 7 , f i sh s how t h e s ame p a t te rn o f i n c r ea s ing a nd c oncen t ra t ed e xp lo i ta t ion d u r ing t h i s p e r iod , p r ima r i ly A t lan t ic s a lmon ( Sa lmo s a la r )( K rupa 1 994 , 1 996 ) . Many o t he r C an tab r ian s i t e s h ave y i e lded f i sh r ema in s , t h ough i ns ma l le r a moun t s ( p robab ly d ue t o d i f f e r ence s i n r e cove ry s y s t em ) . T he mode rn s y s tema t ic e xcava t ion so ft h e r emnan ts t ra t ig raphy a tL aR ie ra y i e lded 8 1 i d en t if i ed S a lmo r ema in sf r om i t sL a te U ppe rP a leo l i th ic s e quence ( S t rau sa nd C la rk 1 986 ) . Wa te rf ow lw e r ea no the rr e sou r ce w ho se e xp lo i ta t ion i n ten s i f i ed d u r ing t h e L a t e U ppe r P a leo l i th ic i nC an tab r ian S pa in . Magda len ian l e ve l s y i e ld ing t h e se c ome f r om B a lmo r i , C ue to d e l aM ina , L a R ie ra , C obe r iza s , A l tam i ra , E l P endo , O te ro , L a C ho ra , L umen txa , T o r r e , E r ra l la , E ka in , U r t iaga a nd A i tzb i ta r t e ( s umma r iz ed b y S t rau s 1 992 :151) . T he m a j o r i ty o fb i rd r ema in s f r om E l J uyo a wa i t f u r the r i d en t if i ca t i on ,b u tr e p r e sen t as i gn if ican ts amp le . M ig ra to ry w a t e r f ow l 1 53

a r e as e a sona l r e sou r ce t h a tw a sp e rhap s h a rve s ted w hen t h i s h um id c oa s ta l r e g ion w a s as t opp ing p o in t o n m ig ra to ry f l y w ay so r as e a sona lr e f uge a r ea .

" w i ld h ar ve s t ing " . Ma jo rp r o t e in r e sou r ce s , w i th t h e e xcep t i on o fm a r ine mo l lu sc s , b ecame i n c rea s ing ly c oncen t ra t ed i n t empo ra ry ,p e r i shab le s u rp lu se s .

F auna l e xp lo i ta t ion i nt h i s p e r iod i n c lud e s h un t ing o r t r ap p i ng o fs ma l lt om ed ium s i z ed f u r -bea r ing m amma l s s p ec if i c a l ly f o r t h e i r p e l t s ( Chap t e r 9 ) . E l J uyo s o f a r h a s y i e lded 7 2 i d en t if i ed f o o t e l emen t s , p r ima r i ly p ha lange s a nd m e ta p od ia l s , o f c a rn ivo r e s i nt h e s i z e r a ng e o f f e r r e t ( M u s t e la

C hange s i nf o od s t o rage n eed s r e p r e sen t al o g i ca l n ew p r ob l em f o r t h e p r a c t i t ione r s o f m a s s h a rve s t t e chn ique s . E n c oun t e r h un t ing l a ck s t h e s ame u rgency f o r m ea t s t o rag e , s i nce i n d iv idua l a n ima l s p rocu red o n as em i -da i ly b a s i s c an b e c on sumed b ef o re t h ey s po i l a nd k i l led o n ly w hen n o s u r p l u s a l r eady e x i s t s . T he g ene ra l ly c oo l c l ima te a l so r e duced t h e u rgency f o r a mo re p l anned r e spon se . Mea t s h a r ing a mong g roup m embe r s m ay a l so h ave r e duc ed a ny n eed f o r s t o rage : at empo ra ry p e r sona l s u rp lu s t h a t c anno t b e c on s umed b ef o r e s po i l ing c an b e t r aded a mong m embe r s o ft h e g roup o rb e tween g r oup s f o r f u tu r e r e c ip roca t ion w he r e t h e s i tua t ion i sr e ve r sed . O ne t h u sg i ve s u p o n ly w ha tc anno tb e u s ed i nr e tur nf o rf u t u r er e sou r ce s .

p u to r iu s ) , E u ropean m ink ( M u s te la l u t reo la ) , m a r t en (M a r te s s pp . ) a nd p o s s ib ly w o lve r ine ( Gu lo g u lo ) . T he p r e sence o f t h e se b one s i si ns t r ik ing c on t ra s t t ot h e v i r tua l l a ck o fa ny o t he r s ke le ta l e l emen t s , i n c lud ing t e e th , f r om t h e se t a xa . D if f e r en t ia l s u rv iva l d ue t ot h e d u rab i l i ty o ft h e se f o o t e l e m en t s c anno t e xp la in t h i s p a t t e rn , s i nce t h e h ypo the t ica l ly " m i s s ing" t e e th a r et h e mo s td u rab le s k e le ta l e l emen t s o fa l . N on -ca rn ivo r e s o ft h i s s i z e r a nge ( s u ch a sr a bb i t o r h edge h og ) a nd t h e s ma l le r , m o r e f r ag i le m amma l ian t a xa d o n o t s how t h i s s ame e l emen t r e p r e sen ta t ion d i sc repancy . T he a u tho r t h e r ef o r e a t r ibu te s t h e p r eponde rance o ff u r -bea r ing m amma l f o o t e l emen t s t ot h e i r s e lec t ive i n t roduc t ion t ot h e s i te . T h i s p a t t e rn w ou ld b e t h e l o g ica l r e su l t o ff o o t b one s l e f t i n ap e l tr emoved f r om at r app ed o rh un t ed a n ima l a way f r om t h e s i t e , w i th t h e r ema in ing c a r ca s s d i sca rded i nt h e f i e ld . Mu s t e l id m ea th a s as t rong ,u np lea san tf l avo ra nd m ay h ave b een t h e r ef o r e ar e la t iv e ly l e s s d e s i rab le f o od c ho ice , u s ed o n ly i nt i me s o fh a rd sh ip . T h i s p a t t e rn o ff u r e xp lo i ta t i on h a s b een s ugg e s ted f o r o t he r C an tab r ian s i t e s ( S t rau s 1 977 , 1 992 ) . A i tzb i ta r te I V i nB a sque c oun t ry h a s y i e lded s e ve ra l r ema in s f r om Mu s t e la n i va l i s , M . e rm inea , a nd M . p u to r iu s i n i t s S o lu t rean a nd Magda len ian l e ve l s ( A l tuna 1 972 ) . B o l inkoba , aB a sque s i t e , h a s a l so y i e lded t h e r e m a in s o f Mu s t e l id s a nd r e d f o x ( V u lpe s v u lpe s ) . T he s ug g e s t ed p a t t er n o f s p ec ia l iz ed f u r e x t ra c t ion e me rge s d u r ing t h eL a te U ppe rP a leo l i th ic , i nc on junc t ion w i th o t he rc hange s i nf a una le xp lo i ta t ion n o ted . T he e xp lo i ta t ion o fp l an tr e sou r ce s i sn a tu ra l ly l e s s v i s ib le i n t h e p a leo l i th ic a r cha eo log ica l r e co rd d ue t op oo r c hance s o f p r e serva t ion . W i th t h e a dven t o fn ew f l o ta t ion p rocedu r e s s u ch a s t h o se i np r a c t ice a tE l J uyo ( F r eeman e ta l . 1 998 ; S t e t t le r 1 995 ) a nd a t F ranch th i C ave i n G r eece ( Han sen 1 99 1) , f u tu r ee x cava t ion so fe a r l ie ro ccupa t ion sm ay i n d ica te w he the r p l an t r e sou r ce u s e c hanged o ve r t h i s p e r iod . P l an t r e sou rce s d u r ing s umme r i nt h i s i n t e r s tad ia l c l ima te w ou ld h ave b een m uch m o r e a bundan t a nd w ou ld s e a sona l ly c om p l emen tw in t e rm ea tr e sou r ce s . I ns hor t , s i t e s f r om t h e C an tab r ian r e g ion e xh ib i t i n te r r e la t ed t r end s i n h uman / env i ronmen t r e la t ion sh ip s t h roughou t t h e L a t e U ppe r P a leo l i th ic , ap e r iod o f g ene ra l ly r i go rou s a nd f l uc tua t ing c l ima t e . L a rge m amma l e xp lo i ta t ion b ecame f o cu sed u pon r e la t ive ly r a p id ly r e newab l e ,h i gh ly p roduc t ive c ong rega t ing s p ec ie s w h ich w e r e t a k en w i th i n c rea sed e ff i c i ency . E v id ence o ft h i st r end i n c lud e st h em a s s s l augh t e ro f r e d d ee r a nd i b ex h e rd s , a sw e l l a st h e s p ec if i c u s e o fs ome s i t e s ( Ra scaf io )a sp r ima r i ly s i ng le spec ie se x t ra c t ive h un t ing c amp s . S a lmon r u n s w e r e a l so s ub jec tt oi n c r ea s ing e xp lo i t a t ion t h rough Magda len ian t i me sa nd i n to t h e E p ipa leo l i th ic ( Pok ine s a nd K rupa 1 997 ) . T he se r u n s b y t h e i r n a tu r e p ro v i de s e a sona l a nd c oncen t ra t ed s ou r ce s o ff o od , a nd a ss u ch r e qu i re g roup c oope ra t ion a nd c oo rd ina t ion f o rt h e i re ff i c ien t

Ma s s h a rve s t ing o fp e r i shab le r e sou r ce s p r e s en t s a n e n t i r e ly n ew p rob lem t o s o lve a nd w i th i tt h e p o ten t ia l f o r g r ea t ly c hanged g roup b ehav io r . Wh i le ma r ine m o l lu sc s c ou ld b e h a rve s ted i nl a rge a moun t s o n ly a sn eeded , am a s sh a r ve s to f r e d d ee r , i b ex , o r s a lmon p r oduce s ah i gh ly p e r i shab le r e s ou r ce t h a t w i l l r a p id ly s po i l u n le s s m ea su r e s a r e t a ken f o r i t s p r e se rva t ion . T h e o p t ion o f t r ad ing a way a p e r i shab le r e sou r ce f o r f u tu r e c on s ide ra t ion s i sg r ea t ly r e duced . S i nce g roup c oope ra t ion w a s n ece s sa ry f r om t h e o u t se t , e a ch m embe r h a s t h e s imu l taneou s p rob lem o fi mmed ia te s u rp lu s a nd f u tu r e n eed . S i nce t h i s f a una l e xp lo i ta t ion p a t ter n h o ld s f o r t h e w ho le C an tab r ian r e g ion , t r ad ing w i th o ne ' s n e igh b o r s i sl e s s o f a n o p t ion ( t hey a r e l i ke ly t oh ave t h e s ame p rob lem a nd a tt h e s ame t i me ) . T he r ef o r e , m ea t s t o rage b e c ome s ap r e s s ing n eed i n ah um id c l ima t e i l s u i t ed f o r i t d u r ing mo s to ft h ey ea r . T he s e a son o fo ccupa t ion f o r h uman s a tE l J uyo i sp a r t ia l ly c l a r if i ed b y t h e n ew d a ta p r e sen ted h e r e ( Chap te r 2 ) c on c er n ing t h e s e a son o f o ccupa t ion o f b a rn o w l s i nt h e s i t e . Wh i le c oncu r r en t o ccupa t ion i sp o s s ib le ( a nd o ccu r s c om mon ly i no t he r s e t t ing s ) , t h e s p a t ia l l a you to ft h e c ave v e s t i b u le m ake s t h e s imu l taneou s h ab i ta t ion b y r oo s t ing b a rn o w l s a nd a ny s ub s tan t ia l , s em i -pe rmanen t h uman e ncamp m en tt h e r e u n l ike ly . O ccupa t ion b y b ar no w l s f o rp a r t icu la r s e a son st h e r ef o re s ugge s t st h a th uman st e nded n o tt oo ccupy E lJ uyo a tt h o se s ame t ime s . T he a ge p rof i le s , w he r e d e t e r m inab le , f o r b ar n o w l s ma l l m amma l p r ey i n d ica t e s t h a t m any o ft h e se i n d iv idua l s w e r e v e ry y oung j uven i le s , p e r h ap s o n ly d ay s p a s tw ean ing . T he se v e ry y oung i n d iv idua l s w ou ld h ave b een p r e sen ti nt h e l o ca lp opu la t ion so n ly d u r ing t h e l a t e s p r ing / summe r /ea r ly a u tumn mon th s , r ough ly A p r i l t h rough S ep t emb e r e a ch y ea r ( s ee C hap t e r 3f o r s umma r ie s o f t h e b r eed ing s e a son o f t h e se s ma l l m amma l s p ec ie s ) . B a rn o w l s m u s th ave b een p r e sen t i nE l J uyo a tl e a s tp a r to f t h e t ime d u r ing t h e se mon th s , t hough t h e i r p r e sence t h e r e d u r ing t h e r ema inde r o f t h e y ea r c anno t b e d i scoun t ed b y a na ly s i so ft h e i rd i e t . H uman o ccupa t ion a tE lJ uyo t h e r ef o r e m ay h ave b een c on c en t ra t ed f r om l a t e a u tumn t h rough l a te s p r ing . N o c l ea r s e a son o f h uman o ccupa t ion w a s d e t e rm inab le t h rough i n v e s t iga t ion o fd en ta la t t r i t ion m ode sa nd c emen tum a nnu l i o f t h e r e d d ee r t e e th ( F r eeman e ta l . 1 988 ) . T he p r e sence o f 1 54

c e rv id f o e ta l b one s i n t h e L eve l 4S anc tua ry f e a tu r e a tE l J uyo s how s t h a t h uman o ccupa t ion l a s t ed a tt i me s u n t i l l a te s p r ing / ea r ly s umme r when f u l l te rm ( o r n ea r ly s o ) p r egnan t f ema le s w e r e p r e sen t i nt h e r e d d ee r p opu la t ion . T he p r e s e nce o f mo s t ly u n shed a n t le r i n d ica t e s t h a t s i gn if i can t n um b e r s o fa du l t m a le r ed d ee r w e r e t a ken d u r ing a u tumn /ea r ly w in t e r , s i n ce t h e i r a n t le r s a r e s h ed s h o r t ly a f t e r t h e ma t ing s e a son e nd s ( i n J a nua ry ) . A no the r i n d i ca t ion o fh uman o c c upa t ion o fE l J uyo i nt h e a u tumn i st h e p r e sence o fm any s e ed s p ec ie s , m o s to fw h ich a r e p rodu c ed ( a nd a r e o p t ima l ly h a rve s t ed ) i nt h e a u tumn mon th s . H uman a cqu i s i t ion , h ow e ve r , i sn o t t h e o n ly e xp lana t ion f o rt h e i re n t ry i n to t h e s i te . S a lmon h a rve s t m o s t l i k e ly o ccu r r ed d u r ing t h e l a t e a u tumn s p awn ing r un , w h ich f u r the r i n d ica t e s h uman o ccupa t ion o f E l J uyo a tt h i s t ime o f y ea r . H uman s e a sona l c yc le s i n L owe r Magda len ian C an tab r ian S pa in t h e r ef o r e m ay h ave i n c luded a u tumn a nd w in t e r o ccupa t ion o fl ow land s i t e s l i ke E lJ uyo , c omb ined w i th s umme ro ccupa t ion o fw a rm s e a son a c ce s s ib le u p land s i te ss u ch a sR a sca f io . T he e a s ie s t m e thod o fm ea t p r e se rva t ion i ss imp le f r eez ing d u r ing w in t e r . S i nce i n t e r s tad ia l c ond i t ion s e x i s ted t h rough o u tm o s to ft h i sp e r iod , s ub s tan t ia lf r e ez ing d u r ing t h ew in te r m on th s b ecome s av i ab le p a r t ia l s o lu t ion . A n i d ea l t ime t o h a rve s tr e dd ee ri s when t h ey c ong r ega t e f o rt h e a u tumn r u t t i ng s e a son , s o l a rge m ea t s upp ly c ou ld b e a cqu i r ed j u s t a s t empe r a tu re s d i pb e low f r eez ing . N o s p ec ia l e f f o r t i sn ece s s a ry o t he rt h an a n e ven g r ea te rn eed t or e duce l o ca l s c aven g e r / ca rn ivo r e p opu la t ion s . S i nce m any o ft h e r e d d ee r r e m a in s f r om E l J uyo w e r e t a ken i nl a t e s p r ing / ea r ly s umme r w hen f r eez ing o ft h e m ea t a s soc ia t ed w i th t h e se s ke le ta l r e m a in s w ou ld n o t b e f e a s ib le , s ome o t he r m e thod o fp r e se r v a t ion m u s t h ave b een p r a c t iced a tt h a t t i me . A n o p t ion i n t h i ss i tua t ion i sd r y ing / smok ing m ea t . T h i sp rocedu re i sn ec e s sa ry t op r oce s s l a rge a moun t s o fm ea t a cqu i r ed d u r ing t h e w a rme r mon th s o ft h ey ea ra nd p r e sen t sn o g r ea tt e chno log i c a l o b s ta c le s( Coon 1 971) . T h in s e c t ion s o fm ea tm ay b e a i r d r ied i ns un l igh t , o rl a rge rp i ece s m ay b e c u r ed b y s mok ing . Mea t p roce s sed t h i s w ay c an b e k ep t f o r l o ng p e r iod s o f t i me . D i rec t a r chaeo log ica l e v idence o ft h e se p rocedu r e s i s mo re d i ff i cu l tt oo b ta in . E v idence f o rt h es y s t ema t ic p roce s s ing o f f a una t h a t i s an ece s sa ry p r e l im ina ry s t ep i nt h e se p roce d u r e s , h oweve r , h a s b een o b ta ined f r om E l J uyo . A l a rge p r opo r t ion o fr e d d ee rr ema in s w e r e d i s ca rded i nt h e d ugou t s t ruc tu re i nL eve l 6 ,w i th i n d ica t ion s o fs e lec t iv i ty i nb ody p a r tr e p r e sen ta t ion a nd d i spo sa l ( F r eeman e ta l . 1 988 ) . T h i s p a t t e rn s ugge s t s m e thod ica l , w ho le sa le p roce s s ing o f r e d d ee r . T he o rde r ly p a t te rn o fs k e le ta l d i spo sa l i sd i ff i cu l t t o e xp la in i fi n d iv idua l d ee rw e r e a cqu i r ed i n mu l t ip le e p i sode s a nd i sr e inf o r ced b y t h e c a ta s t roph i c a ge p ro f i le o b ta ined f r om c r own h e igh t s . R ed d ee rw e r e n o t o n ly h a rve s ted m e t h od ica l ly , b u t a l so p roce s sed m e thod ica l ly . T he r e la t ive l a ck o f b one s f r om m ea t ie r b ody p o r t ion s i nL eve l 6i sa n i n d ica t ion t h a t s ub s tan t ia l p roce s s ing o fc a r ca s se s t o ok p l ace a way f r om t h e c ave v e s t ibu le . O ngo ing r e sea r ch i n to t h e f a una l r ema in s o f E l J uyo may d i sc lo se f u r the r r e inf o r c ing p a t te rn so fb u t che ry o rp a r tr e p r e s en t a t ion . F i sh r ema in s f r om E l J uyo a l so s ugg e s tp a t ter n s o fm e thod i c a l p roce s s ing o fl a rge a moun t s o ff i sh e n m a s se . T h i s e v i -

d ence i so b ta ined f r om s k e le ta lp a r tr e p r e sen ta t ion . Wh i le a l l p a leo l i th ic l e ve l s a nd f e a tu r e s o ft h e s i t e h ave y i e lded f r ag m en ta ry f i sh r ema in s , a lmo s t n one o ft h e se i s ac r an ia l e l e m en t . N o o t o l i th s h ave b een d i scove r ed . T he e l emen t s t h a t h av e b een r e cov e r ed i ng r ea te s t a moun t s f r om s c r een ing s i te d epo s i t s a r e f r agmen ta ry r i b s a nd f m s p ine s , w i th v e r teb ra l e l emen t s af a r d i s tan t s e cond . T he se e l emen t s a r e t h e mo s t f r ag i le , s o t h e i r o ve r r ep re sen ta t ion c anno t b e e xp la ined i n t e rm s o fd i f f e r en t ia l p r e ser va t ion . T h i s p a t te rn s ugge s t s t h a t mo s t c r an ia l e l emen t s w e r e d i sca rded a way f r om t h e s i te , p e rhap s a tt h e s i t e o fc ap tu r e . T h i s p r a c t ice w ou ld b e e ff i c i en t a nd n ece s sa ry i fl a rge a moun t s o f f i sh w e r e t a ken a t o nce a nd t h e f l e sh w a s d eboned a nd p r e se rved . S i nce p r e l i m ina ry i n ve s t iga t ion i n d ica t e s t h a t mo s t o f t h e se r ema in s a r e t h o se o f A t lan t ic s a lmon , h a rve s t ing mu s t h ave t a ken p l ace d u r ing s e a sona ls p awn ing r u n s i nn ea rby r i ve r s . T he se o ccu rb eg inn ing i nl a t ea u tumn ,t h ough s ome f i sh c ou ld h ave b een t a ken i nt h e p r in c ip le s p awn ing m on th s o fN ovembe r t h rough J a nua ry ( Ne tboy 1 980 ) . T he n ece s s i ty f o r f o od s t o rage a l so a f f ec t s p a t ter n s o fl a nd s c ape u sage ( Ba i ley a nd D av id son 1 983 ) . L a rge a moun t s o f s t o r ed m ea t d ec r ea se t h e mob i l i ty o f ag r oup , e spec ia l ly i n t h i s g ene ra l ly r u gged t e r ra in . T h i s d ec r ea sed mob i l i ty i s r e f lec t ed i nt h e u s e o f s p ec ia l iz ed r e sou r ce c amp s , s u ch a s t h e Magda len ian i b ex h un t ing c amp o fR a scaf io . I n s tead o f m ov ing t h e b a s e c amp t o w he r e t h e r e sou r ce s a r e , t h e r e s ou r ce s a r e b r ough t t o ac en t r a l ized l o ca t ion . E l J uyo h a s y i e lded i b ex r ema in s f r om L eve l s 4 / 4S a nd 6 ( K le in a nd C r uz U r ibe 1 987 ) , e ven t h ough t h i s mon tane s pec ie s w ou ld h ave b een m any k i lome t e r s d i s tan t f r om t h e s i t e . Wh i le r e d d ee ra nd b i son w e r e a va i lab le i nt h e e nv i ron s o ft h e s i te , m a r i ne mo l lu sc s w e r e t r an spo r t ed u p t o 1 0 km t o t h e s i te i n l a rge n umbe r s . T he n ea r e s t s a lmon r i ve r( R io P a s ) i so ve r 5 k m d i s tan t . Ma jo r f o od r e sou r ce s t h a t w e r e c oncen t ra ted a t t h e i rp o in to fh a rve s tw e r et r an spo r ted t oE lJ uyo . O ne mu s t a l so c on s ide r , h oweve r , t h a t i nC an tab r ian S pa in d i s tance s b e tween a l v e r t ica l ly z oned h ab i ta t s a r e s ma l l , a nd mob i l i ty n eed n o t h ave b e en g r ea t f o r as i ng le g r oup e xp lo i t ing a l o f t h e se . I na dd i t ion , t r ue s t o rage p i t s ( t ha t m igh t b e e xpec ted u nde r t h e c i r cum s tance s s ugge s t ed ) a r e v i r tua l ly a b sen t . O the r m ean s o f f o od s t o rage m ay h ave b een p r ac t iced . T he se i n c lud e t h e s t ruc tu ra l r emnan t s f r om L eve l 6o f E l J uyo , o ra r cha eo log ica l ly l e s s v i s ib le m ean s s u ch a ss u spen s i on o np l a tf o rm so rp o le sa way f r om p r eda to r s . D u r ing t h i ss ame p e r iod w es e e ag r adua l i n c r ea se i np opu la t i on , a ss hown b y t h e i n c r ea s ing n umbe r o fs i t e s a nd i nt h e i n t en s i ty o f t h e i r o ccupa t ion ( S t rau s 1 977 , 1 992 ) . A n i n c r ea s ing p opu la t ion l o g ica l ly c on ta in s o n a ve rage a h i ghe r p ropo r t ion o f c h i ld r en . Two o t he r i n c r ea s ing ly i mpo r tan t a r ea s o ff a una l e xp lo i ta t ion d u r ing t h i s p e r iod i nC an tab r ian S pa in a r et h e g a the r ing o fm a r ine mo l lu sc s a nd f u rt r app ing . B o th a r e h i gh ly s u i tab le a c t iv i t ie s f o r c h i ld ren , a s t h ey r e s p ec t ive ly i n vo lve i mmob i le , e a s i ly -ha rve s ted mo l lu sc s o r t h e m a in t enance o f at r ap l i ne w he re s ma l l t om ed ium f u r b ea r e r s a r e a l r eady d ead o r c ap tu r ed . T he i r h a rve s t a tt h a t p o in t s imp ly i n vo lve s c o l lec t ion f r om t h e t r ap s . F a r l e s s s t r eng th , s p eed o rc r i t ica lj udgmen t a r e n ece s sa ry a tt h e t ime o fh a rve s to fs u ch s p ec ie st h an d u r ing t h e d r iv ing , s u r round i n g ,a nd s l ay ing o fl a rge f a una . We igh tl o ad sf o rt r an spo r tt o ah ab i ta t ion s i t e a r e a l so e a s i ly c on t ro l led b y l i m i t ing mo l 1 55

l u sc h a r ve s t a nd s k inn ing t r apped a n ima l s i nt h e f i e ld , t h e p roce s s ing p a t te rn s ugge s ted b y t h e i r p ropo r t iona l s k e le ta l r e p r e sen ta t ion . T he c o l lec t ion o fm o l lu sc s b y c h i ld r en e s pe c i a l ly f r ee s a du l t s f o rm o r e d i ff i cu l tt a sk s , w h i le s upp ly ing a l a rge a nd r e l iab le s upp ly o fp r o t e in . D ISCUSS ION T he c onf l a t ion o fp opu la t ion i n c r ea se a nd r e sou r ce m anage m en t a tt h i s t i me o fc l ima t ic o s c i l la t ion s -gene ra l a me l io ra t i on w i th i n t e rm i t t en t p e r iod s o f d e te r io ra t ion ind i ca t e s a g row ing s e pa ra t ion o fh uman b ehav io r f r om t h e d i r ec t c on s t ra in t s o fa nu np r ed i c tab le n a tu ra l s e t t ing . M ic rof auna l a nd o t he rp a leoenv i ronmen ti n d ica to r sp o r t ray ah um id , c o ld a nd o pen l a nd scape m ov ing t h rough m u l t ip le c yc le s o fp oo r e r o r mo r e f a vo r ab le c ond i t ion s , a pp roa ch ing t h e t empe ra t e a nd mo r e f o r e s t ed H o locene . R ed d ee r , i b ex , m o l lu sc s , s a lmon , a nd p e rhap s f u r s w e r e o b ta ined i na n i n c r ea s ing ly e ff i c ien t a nd c oope ra t ive m anne r . T he se t r end s a r e r ef l ec t ed i nt h e f a una l r ema in s t h em se lve s , a sw e l l a st h e c hang ing p a t ter n s o fs i te u s ea nd t e chno logy . I n s t ead o fh uman r e sou r ce a cqu i s i t ion f o l low ing f l uc tua t ion s c au sed b y c hang ing e nv i ron m en t , h uman c on t ro l o ve r t h e h a rve s t o ft h e se r e sou r ce s a c c e le ra te si n dependen to fe nv i ronmen ta lf l uc tua t ion s . Wh i le t h e a u tho r i nn o w ay s ugge s t s t h a t C an tab r ian S pa in w a s ah ea r th a r ea f o r a g r icu l tu r e ( wh ich c ame t ot h i s r e g ion r a the r l a t e ) , t h i s c hang ing h uman r e la t ion sh ip w i th e nv i ron m en t i nm any w ay sp r e sage st h e c hange s n ece s sa ry f o ra g r i c u l tu ra l s y s t em s t o d eve lop ( F r eeman 1 973 , 1 975 ) . F ood p roduc t ion , w he the r b y h e rd ing o rc r op g r ow ing , r e p r e sen t s s e pa ra t ion f r om t h e c on s t ra in t s o f n a tu ra l ly h a rve s tab le r e s ou r ce s . E nv i ronmen ta l f a c to r s i nt h e f o rm o fc l ima t e , t e r r a in a nd b i og eog raphy o fs p ec ie sa va i lab le f o rd ome s t ica t ion s t i l l a f f ec ta g r icu l tu ra l s y s t em s . A g r icu l tu r i s t s i nm any c a se s r e ly u pon w i ld s p ec ie st os upp lemen tt h e i ro wn f o od p r oduc t i on , a nd s o r ema in t i ed t ot h e i r n a tu ra l e nv i ronmen t r a the r c l o se ly . T hey a l so c r ea t e a r t if i c ia l e co sy s t em s w ho se i n t e r n a lw o rk ing sa pp roa ch t h ec omp lex i ty o fn a t u ra l e co sy s t em s . C ro s s p o l l ina t ion a nd h ence g ene f l ow o c cu r s b e tween d o m e s t ica t ed a nd w i ld a nce s t ra l f o rm s ( a s w i th m a iz e a nd t e o s in te i nC en t ra l Ame r ica ) , f u r the rl i nk ing t h i sr e la t ion sh ip . U l t ima te r e sou r ce h a rve s t , h oweve r , i sl a rge ly d e te rm ined b y d i r ec th uman a c t ion a nd d oe sn o tr e ly o n ly u pon n a tu r a l p ro d uc t iv i ty t oe s tab l i sh r e sou r ce y i e ld s . W i ld h a r ve s t ing o fr e sou r ce s p l a ce s h uman e co log ica l i n t e r a c t ion a l ong t h e s p ec t rum c l o se r t oa g r icu l t u r e . T he o t he r e nd o ft h i s s p ec t r um i sr a ndom -encoun t e r c o l lec t ion , w he r e f ood r e sou r ce s a r e a cqu i r ed m o r e i np r opo r t ion w i th t h e i r n a tu ra l a bundance s a sf ound o n t h e l a nd scape . T he e x t r eme e nd o ft h i s s p ec t rum i sn o t o b ta inab le , s i nce h uman s m ake c ho ice s i nt h e i r f o od a cqu i s i t ion e ven t ot h e p o in t o fs t a rva t i on i n t h e p r e sence o f c u l tu ra l ly u na ccep tab le f o od r e s ou rce s . N o r i st h e o t he r e nd o ft h i s s p ec t rum o b ta inab le ( t o ta l i n d ep endence o fe nv i ronmen ta l c on s t ra in t s u pon f o od p roduc t ion ) , s i nce t h e se f a c to r s p l a ce b ounda r ie s t h a t l i m i t t h e b r ead th o fh uman d ec i s ion m ak ing . N o e v id ence e x i s t s t h a t U pp e r P a leo l i th ic C an tab r ian s w e r e a c t ing t or e p len i sh n a tu ra l r e sou r ce s o ra l t e r e nv i ronmen t s t h a t w ou ld f a vo rt h e g row th o fp r ef e r red r e sou r ce s p ec ie s , o t he r t h an t h e r e duc t i on o fc ompe t i t ion f r om o t he r l a rg e p r eda to r sa nd t h ep o s s i b l ed e l ib e ra t e c u l t iva t ion o fm a r ine m o l lu sc s( F r eeman 1 973 ,

1 975 ) . O the rp o s s ib le a c t ion s i nt h i sr e ga rd i n c lud et e r r i to r i a l i sm i n o r d e r t o p ro tec t i mpo r tan t r e sou r ce s f r om o t he r h un te r -ga the r e r g roup s . L and scape a l t e r a t ion u s ing f i r e t o f a vo rt h e h ab i ta t s o fc e r ta in u ngu la t e p r ey s p ec ie s i sa no the r p o s s ib i l i ty , o r t h e s e lec t ive p r eda t ion w he r e p o s s ib le u pon w eakened o rn on r ep roduc ing i n d iv idua l s . T he a lowance o f " f a l low " p e r iod s f o ra r ea s o f mo l lu sc h a rve s t i sa no the rc on c e ivab le p r a c t ice . C hang e s i nr e sou r ce a cqu i s i t ion t h e r ef o re l i e n o t s o m uch i nc on t ro l o f p roduc t ion b u t i nc on t ro l o f h a rve s t . I n c r ea sed e ff i c iency i nl a nd scape u s age , t e chno logy a nd i t s e mp loymen t , a nd s t ag ing o ft h e e xp lo i ta t ion a nd p r oce s s ing o f r e sou rce s p l a ce s ah uman g roup i na n e n t i r e ly d i f f e r en t r e la t ion sh ip w i th i t s e nv i ronmen t t h an h ad p r ev iou s ly b een t h e p a t t er n f o r C an tab r ian S pa in . A s m en t ioned a bove , h u m an s h ad b egun i ng ene ra l d u r ing t h e U pp e r P a leo l i th ic t o r e duce c omp e t i t ion f r om p r eda to r s a nd g r ea t ly d ec r ea se t h e n umbe r s a nd r a nge s o ra i d i nt h e e x t inc t ion o fm any m ega f a una s p ec ie s ( Ma r t in a nd K le in 1 984 ) . T he se a c t ion s p l a ce h uman s d i s tan t f r om o t he r l a rge p r eda to r s , w ho se n umbe r s a r e l a rge ly d e te rm ined b yt h ea va i lab i l i ty o fp r ey s p ec ie s . A no the r p a r a l le l t o e a r ly c hange s r e la t ed e l sewhe r e t ot h e o r ig in s o f a g r icu l tu r e ( Cohen 1 977 ; R eed 1 977 ) i st h e i n c r ea sed i mpo r tance o fs t o r ed f ood r e sou r ce s a nd t h e i r r e la t i on sh ip t os e d en t i sm . A g r icu l tu r e l i e sa to ne e x t r eme ,w he re mo s t f o od r e sou r ce s a r e l i t e ra l ly r o o ted i np l a ce a nd a r e h a r v e s ted d u r ing o ne o r af ew b r ief p e r iod s . T h e r e su l t s o ft h i s h a rve s tm u s tb e s t o r ed t h roughou tt h e f o l low ing y ea rt op r o v i de m o s t f o od r e sou r ce s a nd s e ed s t ock f o r t h e f o l low ing y ea r ' s c r op s . H un t ing a nd g a the r ing r e l ie s f a r l e s s u pon s t o r ed r e sou r ce s , s i nce t h e s e a r e i ne s sence s t o r ed d i spe r sed t h roughou t al a rg e t e r r i to ry , w i th m any n a tu ra l b a ckup s a va i lab le i nt h e e ven t o ft h e f a i lu r e o fo ne r e sou r ce . C on t r o l led w i ld h a rve s t ing t h a t p roduce s as t o r ed r e sou r ce p u she s ah un t ing -ga the r ing g roup t owa rd s t h e c on s t ra in t s o f s e den t i sm . Mo r e o ft h e i r f ood r e sou r ce s a r e s e a sona l ly o r p e r iod ica l ly c oncen t ra t ed . T h i s e ff i c iency a nd r e sou r ce c on c en t ra t ion m ay i nt ur n a low p opu la t ion t og r ow , f o rm ing a p o s i t ive f e edback r e la t ion sh ip b e tween i n c r ea sed l a bo r a nd i n c r ea sed r e t u rn s . S eden t i sm m ay a l so a f f e c t t h i s r e la t ion s h ip , s i nce r e duced m ob i l i ty a l low s s ho r te r b i r th i n t e rva l s a nd a n i n c r ea se i np opu la t ion g row th . T he se f a c to r s b ecame v e ry i mpo r tan t i n l a t e r c hange s a s soc ia t ed w i th a dvan ced a g r icu l tu ra l s o c ie t ie s : t h e g r ow th o fu r ban i sm , c r af ts p ec ia l i z a t ion a nd s o c ie ta ls t ra t if i ca t ion . E ff i c ien t w i ld h a rve s t ing a l so i mpa c t s t h e f o rm o f e t hnog r a ph ica l ly k nown h un t e r -ga the re r s o c ie t ie s , s u ch a s t h e c h ief dom so ft h eN o r thwe s tC oa s tI n d ian so fN o r th Ame r ica . T he se r i ch ly c omp lex , s e den ta ry a nd h i gh ly t e r r i to r ia l t r ibe s h ad a st h e i r s u cce s sf u l f o od e ne rgy b a se t h e r i ch s a lmon r e s ou r ce s o f t h e N o r th P a c if i c O cean ( Ne tboy 1 980 ) . T h i s c oa s t a l r e g ion h a rd ly a low s s ucce s sf u l a g r icu l tu r e a nd p a r a le l s L a te P l e i s tocene C an tab r ian S pa in i nt h e s u i tab i l i ty o f i t s r e sou r ce s . I n ten se , c on t ro l led s e a sona l h a rve s t ing a nd s t o rage o f aw i ld r e sou r ce a lowed t h e g r ow th o fs e t t led v i l l a ge s , d en se p opu la t ion , e l abo ra te a r t , i n t en s e c r af t s p ec ia l i z a t ion . T e r r i to r ia l i sm a l so b ecame v i ab le , s i nce t h i s n on d i spe r sed r e sou r ce w a sw o r th p ro te c t ing . E labo ra te c e r emo n i a l s a t l e a s t p a r t ly r e la ted t o t empo ra ry s u rp lu s r e sou r ce 1 56

r e d i s t r ibu t ion ( t he p o t la t ch ) w e re a n a dd i t iona l c u l tu ra l r e s pon se t ot h ep a r ame te r so ft h i se nv i ronmen t . O ne f a c to rt h a t m ay h ave s haped t h e g r ow th o ft h e c u l tu ra l s y s tem s o ft h e P ac if ic N o r thwe s t i st h e g eog raphy o fm a jo r s a lmon r i ve r d r a inage s . T he C an tab r ian c oa s t h a s m any s ho r tr i ve r s f l ow ing d i r ec t ly f r om t h e c o a s ta l m oun ta in sw i th e a ch d r a in ing ar e la t ive ly t i ny a r ea . T he a moun t o fs a lmon r e -en te r ing t h e s er i ve r se a ch y ea rt os p awn i sp r opo r t iona lt o t h i s d ra inage a nd i t s a r ea o f s p awn ing s t r eam s . T he C an tab r ian s a lmon r e sou r ce i st h e ref o r e d i spe r s ed a mong m any e n t ry p o in t s t o i n d iv idua l v a l l ey s . L a rge r d r a inage s y s tem s i nt h e P ac if i cN o r thwe s tm ean t h a t ag r ea t e rn umbe r o f s a lmon m u s t p a s s t h rough t h e b o t t leneck p o in t s o f k ey r i ve r mou th s . T h i sr eg ion a l so h a s ar i che rv a r ie ty o fs a lmon s p ec ie s ( 6 s p ec ie s i n t h e g enu s O nco rhynchu s i n hab i t t h e N o r th P a c if i c v s . o ne s a lmon s pec i e s i nt h e g enu sS a lmo i n h ab i t ing t h eN o r th A t lan t ic )w ho se r u n s , s i gn if ican t ly , d on o t c o inc ide . A t lan t ic s a lmon a l so e n t e r t h e f r e shwa te r s y s t em o n a mo r e d i spe r sed t i me f r ame ( M i l l s 1 989 ) . T he s a lmon r e sou r ce s o ft h e P ac if i cN o r thwe s tt h e r ef o r e a lowed s i gn if i c an t ly m o r e c oncen t ra t ion o f mu l t ip l e f i sh s p ec ie s a tk ey g eog raph i c p o in t s d u r ing s e ve ra l s e a sona l r u n s a nd am o re

p ro longed h a rve s t . T he C an tab r ian r e g ion d u r ing t h e L a te U ppe rP a leo l i th ic h ad amo r e d i spe r sed a nd l e s s r i ch s a lmon r e sou rce . T he C an tab r ian r e g ion a l so h ad o t he r r i ch r e s ou r ce s t o o f f e r ( r ed d ee r , i b ex , m a r ine mo l lu sc s ) , s o t h a t w h i le p r ima ry r e l iance u pon s a lmon w a s n o tp o s s ib le , i tw a s a l so u nnece s sa ry . I n s umma ry , t h e p a leoenv i ronmen ta l r e la t ion sh ip s o f t h e U ppe r P a leo l i th ic C an tab r ian s s how a n i n c r ea s ing p a t te rn o f s e pa ra t ion o fh uman a c t ion f r om t h e i mmed ia t e c on s t ra in t s o ft h e i r e nv i ronmen t ( F r eeman 1 9 73 , 1 975 ) . H a rve s t ing o f w i ld r e sou r ce s i n c r ea sed i ni n t en s i ty t h roughou t t h i s p e r iod d e sp i t e m a jo r c l ima t ic f l uc tua t ion s i n t h i s o pen , h um id c oa s ta l z one . I n s t ead o fa bandon ing t h i s r e g ion d u r ing t h e g l ac ia l max imum a nd o t he r c o ld p e r iod s , t h e C an tab r ian s s t ead i ly i n t en s if i ed t h e i r u t i l iza t ion o ft h e l a nd scape . S ub s i s tence f o cu sed u pon af ew h i gh ly p r oduc t ive a nd c oncen t r a ted r e sou r ce s , w h i le c on t inu ing t oe xpand i n to o ne sp r ev i o u s ly u nde r exp lo i t ed . T he se c hange s i n l a nd scape u sage ( away f r om mo r e r a ndom r e sou r ce e xp lo i ta t ion a nd t owa rd s p l anned , i n ten s ive h a rve s t ing a te f f i c ien tl o ca t ion sa nd t ime s ) i nm any w ay s p r e sage t h e c hange s n ece s sa ry f o r a g r icu l tu ra l s y s tem st oa r i se .

1 57

CHAPTER 1 SU M MARY AND CONCLUS IONS

T he g oa l o f t h i s r e sea r ch i st h e r e con s t ruc t ion o f t h e p a l e oenv i ronmen ta l s e t t ing o ft h e L owe r Magda len ian s i t e o f E lJ uyo a nd r e la t ed s i te s i nC an tab r ian S pa in . T he d a ta c en t r a l t o t h i s p r o jec t a r e t e r r e s t r ia l m amma l ian m ic rof auna l r ema in s . T h e se h ave b een c ompa r ed w i th o t he r s ou r ce s o f p a leoenv i ronmen ta ld a ta : c h rono log ica ls ynch ron iza t ion w i th g l oba l o cean l e ve l , s e d imen to logy , l a rge t e r re s t r ia l f a una , a qua t ic f a una , p a lyno logy , a nd m a c robo tan ica l r ema in s . E ach o ft h e s e s ou rce s p o s se s se s i n he r en tb i a se s t h a t mu s tb e t a ken i n to a c coun t w hen i n te rp r e t ing p a s t e nv i ronmen ta l c ond i t ion s f r om t h em . T he a u tho r h a s d emon s t ra ted t h a t i n b o th t h e r e so lu t ion o fi n f o rma t ion c on ta ined a nd t h e p o t en t i a l t o c o r rec t t h e b i a se s i n cu r r ed i nt h e i r i n t roduc t ion t oa s i t e , t e r r e s t r ia l m amma l ian m ic rof auna , w heneve r t h e se a r e r e cove red , a r e am uch m o r e s a t i sf a c to ry s ou rce o f i n f o rma t i on o n p a s t e nv i ronmen ta l c ond i t ion s . T he t a phonom ic j our n ey t r ave led b y t h e se r ema in s i sf a r mo r e d e te rm inab le t h an t h a to fp o l len g r a in s , a nd t h e i ri n t roduc t ion t o as i t e h a s n o t p a ssed t h rough a c u l t u ra l f i l t e r a sh ave t h e r ema in s o f l a rge t e r r e s t r ia l f a una , a qua t ic f a una , o r m ac robo tan ica l r e m a in s . M ic rof auna a r e m o r e d i r e c t ly r e levan t t ot h e r e con s t ruc t ion o fl o ca l p a leo env i ronmen t t h an a r e s ed imen to log i c a l o r g l oba l o c ean l e ve l e x t rapo la t ion s . A n e ven h i ghe r d eg ree o f p a leo env i ronmen ta l r e so lu t ion o f c ou r se m ay b e a ch ieved u s ing t h e se m u l t ip le s ou r ce s i n c on junc t ion , b u t m ic rof auna lr ema in sm u s tb e c on s ide r ed i fi n f o rma t ion y i e ld a nd r e l iab i l i ty a r et ob em ax im iz ed . T h i s s t udy o fp a leoenv i ronmen t h a s a dd r e s sed t h ree r e la t ed q ue s t ion s . T he f i r s t i st h e r e con s t r uc t ion o ft h e L owe r Mag d a len ian h ab i ta t s s u r round ing E l J uyo a nd t h e e x t rapo la t ion o ft h e se e nv i ronmen ta l r e con s t ruc t ion s t o t h e c oa s ta l z one , i n c lud ing t h e n ume rou s o t he r s i t e s i nt h e s ame t ime r a nge . T he se Magda len ian s i t e s i n c lude t h e o c cupa t ion s a tA l tam i ra ( A l tuna a nd S t rau s 1 976 ; S t rau s 1 976 -77 ; F reeman 1 988 ; G onza lez E ch ega ray 1 988 ) , L a R ie ra ( S t rau s a nd C la rk 1 986 ) , E l C a s t i l lo ( Cab r e r a 1 984 ) , T i to B u s t i l lo ( Mou r e R o m an i l lo 1 975 ; Mou re R oman i l lo a nd C ano H e r r e ra 1 976 , 1 978 , 1 979 ) , C ueva Mo r in ( A l tuna 1 9 78 ) , a nd E ka in ( Zaba la 1 984 ) . T he d a ta f o r t h e p r e sen t r e con s t ruc t ion s w e r e t h e v a ry ing p r opo r t ion s o f t e r r e s t r ia l m amma l ian m ic rof auna f r om E lJ uyo a nd r e la t ed s i t e s . Whe r ep o s s ib le , a na ly s i sw a s e x t ended b eyond t h e p r e sence /ab s en c e o f m ic rof auna l s p e c i e s t ot a ke i n to a ccoun tt h e r o le o fb ar n o w l s a nd t h e i r p a t t e rn o fp r ey a c cumu la t ion . T he s e cond r e sea r ch p r ob lem i st h e c ompa r i son o f t h i s s ou r ce o fe nv i ronmen ta l d a ta w i th o t h e r s ou r ce s a l ready e m p l oyed a tE lJ uyo a nd o t he r s i t e s . T he m ic rof auna l t a xa a r e r e p re sen ta t ive o f c e r ta in m ic rohab i t a t s a nd t e nd t o b e f a r mo re s p ec ia l iz ed i nt h e i rc ho ice st h an a r e l a rge f a una ( Av e ry 1 982a ; B ou r l ie r e 1 975 ; D e l ibe s d e C a s t ro 1 985 ; F a e ta l . 1 982 ) . Ac u l t u ra l f i l te rw a s n o t i n vo lved i nt h e i n t roduc t ion o ft h e m ic rof auna l r ema in s , a s i st h e c a se w i th l a rge t e r r e s t r i a l f a una , a qua t ic f a una , c ave s e d imen t s , p o l len a nd m a c ro b o tan ica l r ema in s . A dd i t iona l ly , t h e t a phonom ic t r a jec to ry

f o l lowed b y m ic rof auna l r ema in s i sb e t te r u nde r s tood a nd h ence mo r e c on t ro l lab le t h an o t he r s ou r ce s o fp a leoenv i ron m en ta l d a ta . T hu s , m ic rof auna a r e i n he r en t ly mo re r e l iab le p a leoenv i ronmen ta l i n d ica to r st h an a r e o t he rd a ta c a t ego r ie s , d i s r ega rd ing t h ep opu la r i ty o fu s eo ft h el a t te r . T he t h i rd p rob lem i st h e n a tu r e o ft h e a dap ta t ion o ft h e h u man g roup s i n hab i t ing C an tab r ian S pa in a tt h e e nd o f t h e P l e i s tocene i nr e la t ion t ot h i s c hang ing e nv i ronmen t . B road c hange s i nf ood p ro cu r emen ts t ra teg ie so ve rt i me a r e e v iden t i nt h e a r chaeo log i ca l l e ve l s a l r eady e xcava ted . Mou s te r ian , C ha t e lpe r ron ian , A u r ignac ian a nd P e r igo rd ian /G rave t t ian l e ve l s a r e c ha ra c t e r iz ed b y o ppo r tun i s t ic h un t ing a nd /o r s c aveng ing o fl e s s d ange rou s l a rg e m amma l t a xa : h o r se , b i s on a nd r e d d ee r ( F r eeman 1 973 , 1 975 , 1 98 1 ; S t rau s 1 977 , 1 98 1 , 1 992 ) . B eg inn ing w i th t h e S o lu t r ean a nd a c ce le ra t ing t h rough t h e Magda len ian , A z i l ian a nd A s tu r ian ( Me so l i th ic ) p e r iod s , w i ld f o od p r ocu remen t i nC an tab r ian S pa in s how s a n i n t en s if ica t ion i ne xp lo i ta t ion o f c e r ta in s p ec ie s . T he se w e re p r ima r i ly r e d d ee r , A t lan t ic s a lmon a nd m a r ine m o l l u sc s ( l impe t s a nd p e r iw ink le s ) . S i te l o ca t ion a nd f u nc t ion w e re d i ve r s if i ed a nd t a i lo r ed t o e xp lo i t o t he r s p ec if i c r e s ou r ce s , s u ch a st h e i b ex h un ted a tR a scaho . O the r n ew ly e xp lo i t ed r e sou r ce s i n c lude b i rd s a nd s maW m ed ium m am m a l p e l t s . T he L owe r Magda len ian o ccupa t ion o f E l J uyo s panned a p e r iod o f i n ten s if i ca t ion o f s ome a s pec t s o f r e s ou r ce p rocu r emen ta nd ab r oaden ing o ft h e r e sou r ce b a se t o i n c lude s ome t h a tw e r ep r ev iou s ly u nde ru t i l iz ed . T he se q ue s t ion s w e r e f i r s t a dd r e s sed w i th a t a phonom ic a na ly s i s o f a l m ic rof auna l r ema in s f r om t h e c ave . T h i s a na ly s i s w a s p e r f o rmed t o d e t e rm ine t h e s e r ie s o f f a c to r s i nf luen c ing t h e p a s sage o f t h e se r ema in s f r om l i v ing i n d i v i dua l so ft h e l o ca lp opu la t ion t ot h e i rf i na l r e cove ry t h rough t h e s o r t ing o fu l t ra f ine sc r eened s i t e d epo s i t s . T he i n te rme d i a te s t age s p roceed t h rough t h e f r e sh ly d ead i n d iv idua l s c u l led f r om t h e l o ca l p opu la t ion , t ot h e i r d ef l e shed /d ige s t ed s ke le ton s , t h e i r i n t roduc t ion t oE lJ uyo a nd b u r ia l a mong t h e a r chaeo log ica l s e d imen t s , t h e p r e se rva t ion o f as ub se t o f t h e se e l emen t s , t h e e xcava t ion o f ap o r t ion o ft h e c ave v e s t i b u le , a nd t h e p roce s s ing o ft h e se s e d imen t s b y f l o ta t ion a nd w e t s c r een ing . E ach o ft h e se s t age s r e p r e sen t s s ome l o s s o f e l emen t s a nd h en ce p o t en t ia l l o s s o f i n f o rma t ion f r om t h e p roceed ing s t age ( Dav i s 1 987 ; K le in a nd C r uz -U r ibe 1 984 ) . S i gn if i can t ly , movemen tb e tween t h e se s t age s i nm any c a se s a dd s i n f o rma t ion i nt e rm s o f m a rk s u pon t h e b one s t h em s e lve s , t h e i r p a t t e rn a nd l o ca t ion , a nd d a ta i n he r en t t o t h e s p ec ie sr e p r e sen t ed s u ch a sa ge p rof i le , a c t iv i ty p a t t er n s , a nd b ody s i z e . F a c to r sa f f ec t ing t h ef i na ls t a t eo ft h em ic rof auna l r ema in s a l so l e ave t h e i rm a rk s u pon t h em , f r equen t ly a low i n gt h ed e t e rm ina t ion o ft h es t age si nt h ew innow ing p roce s s . T he g oa l o ft a phonom ic r e sea r ch t h e r ef o r e i st ob a ck t rack u p t h i s t r a i l o f i n t e rmed ia te s t age s i n o rd e r t o r e con s t ruc t a s n ea r ly a sp o s s ib le t h e o r ig ina l l i v ing p opu la t ion s o fa n ima l s a nd t h e i r i n te ra c t ion s w i th t h e h uman p opu la t ion o f as i t e . E ven ap a r t ia l r e con s t ruc t ion o ft h i s p a s sage a low s af i ne r

1 59

a s se s smen to ft h eb i a se sa f f ec t ing t h ep ropo r t ion so f am i c ro f a una la s semb lage . T he c onc lu s ion s o ft h e a na ly s i s p r e sen t ed h e r e a r e t h a t o w l s w e re o ve rwhe lm ing ly r e spon s ib le f o r t h e i n t roduc t ion o f m ic rof auna l r ema in s i n to E l J uyo a nd a r e t h e mo s t l i ke ly v ec to ro fi n t roduc t ion f o r mo s t C an tab r ian c ave s i te s . Mo re s p ec i f ica l ly , b a rn o w l s( T y to a l ba ) a r et h em o s t l i ke ly s ou r ce a mong t h e p o s s ib le s p ec ie s o f n oc tu rna l r a p to r . T he e v i d ence f o r t h i s c onc lu s ion c ame f r om m u l t ip le s ou r ce s : s i t e s e t t ing , t h e a moun t a nd d i s t r ibu t ion o f m ic rof auna i n E l J uyo , p r ey a ge p rof i le s a nd s i z e d i s t r ibu t ion , t h e i r h ab i ta t p r ef e r ence s ,a c t iv i ty p a t t e rn s , a nd t h ed eg r ee o fb r eakage a nd d i ge s t ive c o r ro s ion u pon t h ei n d iv idua lr ema in s . A l lo ft h e se c on s ide ra t ion s c o r r e spond w i th k nown b a rn o w l b ehav io r a nd p hy s io logy , w h ich a r e a s sumed n o tt oh ave c hanged s i g n if i can t ly s i nce t h e L owe r Magda len ian p e r iod . O the r t a p honom ic a genc ie s c ou ld a l b e r u led o u t o n mu l t ip le g round s . T he se i n c lud e p hy s ica l a genc ie s s u ch a s n a t u ra l p i tf a l lt r app ing b y al i me s tone c ave , w h ich w a s n o t p l au s ib le d ue t ot h e s pa t ia l l a you t o ft h e E l J uyo v e s t ibu le . R unn ing w a te r w a s s im i la r ly d i sm i s sed . O the r p r eda to r s w e re d i s m i s sed b yt h e p a t ter no fb one r e cove ry a nd b r eakage . S ma l l c a rn ivo ran s l e ave b eh ind p r ey s k e le ta l r ema in s w i th r e la t i ve ly h i gh b r eakage a nd d eg r ee s o f d i ge s t ive c o r ro s ion . D iur na l r a p to r s f o l low a s im i la r p a t te rn . O the r s pec ie s o f o w l ( Bubo b ubo , A s io o t u s , A .f l ammeu s a nd S t r ix a l uco ) w e re e l im ina ted o n t h e g round s o ft h e i r u sua l r oo s t ing p a t t er n s ,h un t ing p a t t e rn sa nd p r ey p r of i le s . A n e xcep t ion t ot h i s p a t t er n w a s f o und w i th t h e r ema in s o f h edgehog ( E r inaceu s e u ropa eu s ) , al a rge r s p ec ie s o fm ic ro f a una w h ich p robab ly f o und i t sw ay i n to E l J uyo a s as em i c ommen sa l . A l so e xcep t iona lw e r et h er ema in s o fs ma l l f u r b ea r ing m amma l s s u ch a sw ea se l( M u s t e la n i va l i s ) a nd f e r r e t (M p u to r iu s ) , w h ich t h ea u tho rs ugge s t se n t e r ed E lJ uyo a sa r e su l t o fh uman t r app ing a c t iv i ty . T he m a jo r i ty o fr ema in s c on s i s t o fr oden t s a nd i n sec t ivo r e s a nd w e r e i n t roduced a s t h e p roduc t s o fb a rn o w l p e l le t r e gu rg i ta t ion . T h i s i d en t if i c a t ion o ft h ea c cumu la t ing a gency a low s ag r ea te rr e so lu t ion t op a leoenv i ronmen ta li n te rp r e ta t ion t h an w ou ld t h e p r e sence o r a b sence o fi n d ica to r s p ec ie s . B ar n o w l s a r e ag ene r a l i s t p r eda to r w ho se d i e t c l o se ly r e f lec t s t h e r e la t ive p ropo r t ion s o fp r ey s p ec ie s ' a s soc ia t ed h ab i ta t s . R e la t ive p ropo r t ion s o f s ma l l m amma l p r ey s p ec ie s t h e r ef o r e c l o se ly r e f lec t r e la t ive p ropo r t ion s o f h ab i ta t t y pe s p r e sen t i n t h e L owe r Mag d a len ian l a nd scape . T he b ehav io ra l , m o rpho log ica l , a nd e co log ica l c ha rac t e r i s t i c so ft h es ma l lm amma ls p ec ie sp r e sen ti nC an tab r ian S pa in d u r ing t h e L a te P l e i s tocene a nd H o locene w e r e p r e sen ted i n C hap t e r3 . I ti sf a i rt oa s sume t h a tt h eb ehav io ro ft h e se s p e c i e s h a s n o t c hanged s i gn if ican t ly i nt h e f ew t h ou sand i n t e r v en ing y ea r s . I n a lmo s t a l c a se s , t h i s a s sump t ion i sn o t r a d ica ls i nce t h e se s p ec ie sh ave u nde rgone n om a jo rp hy s ica l c hange si nt h a tt i me . O ne p o s s ib le e xcep t ion i st h e e vo lu t ion o fc ommen sa l b ehav io r o n t h e p a r t o f s ome o ft h e se t a xa , w h ich n ece s sa r i ly c anno t p r eda t e t h e e vo lu t ion o f h uman i n duced i mpa c t s u pon t h e l a nd scape : d ef o r e s ta t ion , a g r icu l t u r e a nd t h e e x ten s ive c on s t ruc t ion o f h ab i ta t ion s . T h i s c oevo lu t iona ry p roce s sc ou ld n o th ave b egun o n al a rge s c a le b ef o r et h eN eo l i th ic P e r iod a nd i t sr e su l tan tc hange s i nl a nd -

s c ape u sage . O ne m ay t h e ref o re i n f e rt h a t t h i s p r oce s s w a s i na n i n c ip i en t s t age d u r ing t h e U ppe r P a l eo l i th i c , p a r t icu l a r ly w i th t h e b a rn o w l s r e spon s ib le f o r t h e i n t roduc t ion o f s ma l l m amma l r ema in s t ot h e s i te . H edg ehog b ehav io rm ay a l so f a l li n to t h i sc a t ego ry . I na dd i t ion , o ne s p ec ie s o fv o le p r e sen t i nP l e i s tocen e S pa in ( P l iomy s l e nk i ) h a s g one e x t inc t , a nd i t sp r e sen ce i nE lJ uyo r e p r e sen t s a nu n expec ted ly l a te l o ca l s u rv iva l . I t sb ehav io r a l c ha ra c te r i s t i c s w e r e t h e ref o re s u rm i sed f r om i t s l i neage , mo rpho logy ,a nd a s soc ia ted s pec ie s /hab i t a t s . R econ s t ruc t ing p a s t e co sy s tem s p r e sen t ed amo r e d i ff i cu l t p rob lem , s i n ce t h e P l e i s tocene e nv i ronm en t s o f C an tab r ian S pa in d o n o th ave d i rec t mode rn a na log s . A na logy t om od e r n n o r the rn E u rope f a i l s t o c on s ide r d i f f e r ence s i n s o la r r a d ia t ion , w h ich w a s h i gh a nd d ay l e ng th m o re r e gu la r e v en d u r ing g l a c ia l c ond i t ion s i nC an tab r ian S pa in . T opog raphy f o rt h en o r ther nc oa s to fS pa in i sa l so ac omp l ica t ing c on s id e r a t ion , d ue t ot h ep r ox im i ty o ft h eh i gh land z one s oc l o s et o t h e A t lan t ic c oa s t . S ome s p ec ie s i n t e ra c t ion s n o l o ng e r o c c u r : t h er a ng eo fM i c ro tu s o economu s( a r c t ic v o le )n o l o nge r o ve r lap sw i th e ndem ic I b e r ian o rP y r enean v o le s p ec i e ss u ch a sP i tymy s l u s i tan i cu s a nd P .p y rena i cu s . T he p r e s en ce o f o t he r s p ec ie s o fv o le m ay h ave a f f ec t ed e co log ica l b ehav io r a nd h ab i ta t e xp lo i ta t ion , a s i st h e c a se t o day w i th c e r ta in s p ec ie s i n te r a c t ion s ( L ig tvoe t a nd V an W i jngaa rden 1 9 94 ) . S ome e co log ica l c ond i t ion s o f t h e L a t e P l e i s tocene a r e n o l o nge rd i rec t ly o b se rvab le e ven i nm oder nn o r ther nc l ima t e s , w h ich r e duce s t h e p o s s ib i l i ty a nd u s ef u lne s s o fd i r ec t a na l o gy w i th mod er nc ond i t ion s . T he m ode rn m ic rof auna o f t h e C an tab r ian r e g ion a nd t h e s u r round ing I b e r ian p en in su la w e r e p r e s en t ed t os e rve a sa b a se l ine f o r c ompa r i son t ot h e p a leom i c rof auna . D a t a c on c er n ing t h e c on t empo ra ry f a una c ame f r om b o th t r app ing s t ud ie s a nd t h e a na ly s i s o f r a p to r p e l le t s amp le s , w i th n ew d a ta f r om e a ch o ft h e se s ou r ce s d e r ived f r om t h e i mmed ia t e e nv i ron so fE lJ uyo . R e la t ive p opu la t ion d en s i t ie s , c ompo s i t i on s ,a nd v a r ia t ion a c ro s sc l ima t ic , a l t i t ud ina l a nd l a t i t ud ina l c l ine so ft h e s em ic rof auna l s pec ie sw e r ep r e s en t ed f o rS pa in . T he t y pe o ft r ap su s ed o rt h e s pec ie s o fp r eda to rw ho s em ea l r ema in s a r e a na lyz ed g r ea t ly a f f ec t s t h e r e la t ive p r opo r t ion s o fp r ey s p ec i e s . T he s ou rce o f mo r ta l i ty f o ra n a r cha eo log i c a l o r p a leon to log ica l s amp le o f s ma l l m amma l s i so f e x t r eme i mpo r tance i ni t si n t e rp r e ta t ion . T he s ame i sl o g ica l ly t r ue f o r amod er ns amp le , s i nce t h em a jo rd i f f e r ence b e tw een t h em i st h e r e cen tne s s o f mo r ta l i ty . A dd i t iona l ly , e xam ina t i on o ft h e m oder n s ma l l m amma l f a una o f ar e g ion a nd i t s r e la t ion sh ip t o t h e c on tempo ra ry c l ima t e a nd e nv i ronmen t c an i l um ina t e t h e r e la t ion sh ip b e twe en P l e i s tocene f a una s a nd t h e i ra s so c ia t ed p a leoenv i ronmen t s . T h i s a na ly s i s i n d ica t ed t h a t b a rn o w l s t o day a r e e x t r eme ly e f f ec t ive s amp le r s o f t h e t e r re s t r ia l s ma l l m amma l s w i th in t h e i rh un t ing t e r r i to ry . T he b i oma s so fs ma l lm amma l st a k en f r om e a ch m a jo r h ab i ta t t y pe w i th in t h i s r a nge ( f o r e s t , w e t l a nd s , h uman h ab i ta t ion s , s h rub land , d r y g r a s s land , h um id g r a s s land , e t c . ) w a s s hown t o r ef l ec t t h e g ene ra l a r ea l p r o p o r t ion s o ft h e se h ab i ta t s . Wh i le s ome s ma l l m amma l s p e c i e s w i l l b e u nde r r ep r e sen ted d ue t ot h e i r a c t iv i ty p a t t er n s ( e spec ia l ly a r bo r ea la nd f o s so r ia l s p ec ie s ) ,o t he rs p ec ie sf r om 1 60

t h e i rg ene r a lh ab i ta t( f o r e s to rh um id m eadow )w i l lr e p r e sen t t h e p r e sen ce o ft h a t b i ome . T he p a t te rn s d e tec t ed w e r e u s ed t oi n te rp r e tt h e m ic rof auna f r om t h e P l e i s tocene l e ve l s o fE l J uyo .

t h e h eav i ly d ef o r e s ted l a nd scape o f moder n C an tab r ian S pa in . T he mo s t s i gn if ican t s h if t i si nt h e a ppea r an ce o ft h e a l p ine s p ec ie s M i c ro tu s n i va l i s i nL eve l 4 / 4S , i n d ica t ing d e t e r io ra t ion o fc l ima te d u r ing i t sa c cumu la t ion .

T he s ma l le r m ic rof auna l r ema in s ( Rod en t ia , I n sec t ivo ra a nd Mu s te la n i va l i s o fC a rn ivo ra ) f r om E l J uyo a nd t h e r e con s t ruc t ion o fp a l eoenv i ronmen tf r om t h e i rr e la t ive p ropo r t ion s w a s p r e sen t ed i nC hap t e r 5 . T he L owe r Magda len ian l e ve l s a r e n o t c ha ra c t e r ized b y ac omp le te m ic rof auna l t ur nove r w hen c ompa r ed w i th m oder n C an tab r ian S pa in : t h e c hange s r e f lec t as h if t i n e mpha s i s . Wh i le m ic ro t ine s (M i c ro tu s , A r v ico la a nd P i tymy s ) a r e as i gn if i can t c omponen t o f t h e m odem m i c rof auna , t h ey d om ina t e t h e L owe r Magda len ian l e ve l s . T h i s d om inance c ome s a tt h e e xpen se o f m u r id s ( Apod emu s a nd M icromy s , p l u s t h e i n t roduced modem c ommen sa l s Mu sa nd R a t tu s ) .

S ome e l emen t s o ft h e E l J uyo m ic rof auna l s amp le w e r e e x a m ined m e t r ica l ly ( Chap te r 6 ) . Mea su r emen t o f s e lec t ed e l em en t s a i d s t a xonom ic i d en t if i ca t ion ( Rup r ech t 1 97 1) , s e pa ra t ion a mong t h e s pec ie s o f ag enu s , a nd q uan t if i e s m i c r oevo lu t iona ry c hange s i np opu la t ion s ( Be r ry 1 978 ) . B io m e t r ica l a na ly s i s i n d ica ted t h a tt h eA r v i co la r ema in s p r e sen t i nE lJ uyo p r ima r i ly b e long t oA . t e r re s t r i s , t h ough t h e p r e s e nce o fs ome i n d iv idua l so fA .s ap id u sc anno tb ed i scoun t ed . T he f o rme r s pec ie s h a s s ma l le r t o o th d imen s ion s t h an t h e l a t t e r a mong P l e i s tocene a nd H o locene s amp le s . T he p r e s e nce o fA .t e r re s t r i si sa l so mo r e l i ke ly d ue t oi t sm o re n o r th e r ly m ode rn r a nge . A na ly s i s a l so i n d ica ted t h a tt h e r ema in s o fA pod emu s mo r e l i ke ly b e long t ot h e s ma l le rA .s y lva t i cu s t h an t oA .f la v i co l l i s . B o th s p ec ie sa r e a s soc ia t ed w i th f o re s t h ab i ta t s . T he p r e sence o f b o th T e u ropaeu s a nd T o cc i d en ta l i s a mong t h e r ema in s a s s igned t oT a lpa w a s i n d ica t ed b ym ea su remen to fh umeru s l e ng th . T h i se l emen ti sl a rge ri n t h e f o rme rs p ec ie s , w i th n o o ve r lap w i th t h e l a t t e r s p ec ie s i n m o s t modem s amp le s . Mea su r emen t o f , M , i d en t if i ed t o M ic ro tu s a g re s t i s la rva l i s r e conf i rmed t h a t t h e l a t t e r s p ec ie s w a s m o re p r eva len t a tE l J uyo . E s t ima t ion f r om i d en t if i ca t i on o ft h e 2M 2 o fM a g re s t i s w a s t h a t t h i s s p ec ie s c on t r ib u t ed a bou t2 5% o ft h et o o th r ema in sa t r ibu tab le t ot h e se t wo s p ec ie sc omb ined .

S h if t s a l so o c cu r red a mong t h e m ic ro t ine s p ec ie s . Wh i le M i c ro tu s n i va l i s ( s now v o le ) i sp r e sen t i n t h e C an tab r ian C o rd i l le r at o day , i t sr a nge n o rma l ly d oe s n o t e x t end i n to t h e c oa s ta l l ow land s . T he p r e sence o ft h i s a l p ine s p ec ie s i nt h e l ow land z one i n d ica te s as i gn if i can t d ownwa rd a l t i tud ina l s h if t c on cu r r en t w i th c l ima t ic d e t e r io ra t ion a t t h e e nd o f L owe r Magda l en ian o ccupa t ion a tE lJ uyo ( Leve l 4 / 4S ) . M o economu s i sn o t f ound t o day i nS pa in , y e t d u r ing t h e P l e i s t o cene i tw a s ar e gu la r p a r t o ft h e m i c rof auna . I t s p r e sence t h e r e i n d ica t e s c l i se r ia l s h if t o f h ab i ta t z one s s ou thwa rd t h rough E u rope . T h i s s h if ta l so i n d ica t e s s i gn if i can t c oo l ing o ft h e l o ca l c l ima te , a sw e l l a s h um id a nd o pen c ond i t ion s . A r v ico la t e r r e s t r i s d om ina t e st h e L owe r Magda len ian m ic ro f a una , b u t i sp r e sen t i nl owe r p ropo r t ion s i nb a rn o w l p e l le t s amp le s f r om c on tempo ra ry C an tab r ian S pa in . C l e thr iono m y s g la r eo lu s w a s n o t p r e sen t i nt h e L owe r Magda len ian m ic rof auna . T h i s m ic ro t ine s p ec ie s i s c ommon i n C an tab r ian S pa in t oday a nd i sa s so c ia t ed w i th f o r e s th ab i ta t s . T he C an tab r ian l ow land s a r e mo r e d en se ly p opu la t ed b y P i tymy s l u s i tan i cu s , w h i le t h e E l J uyo m ic rof auna i sd om i n a ted b y P .p y rena icu s . T h i s i n d ica te s a no the r a l t i tud ina l s h if t i n s p ec ie s r e p re sen ta t ion , s i nce P .p y rena i cu s d en s i ty c o r re la te sw i th i n c rea s ing a l t i tude ( De l ibe sd eC a s t ro 1 985 ) . T he i n se c t ivo r e s p r e sen t i nt h e L owe r Magda len ian l e ve l s o f E l J uyo ( S o re x m inu tu s , S . c orona tu s , N eomy sf od ien s , N a noma lu s , T a lpa e uropaeu s , T o c c id en ta l i s , a nd E r inaceu s e u ropaeu s ) a r e c ommon t o mode rn C an tab r ian S pa in . C ro c i d ura r u s su la a nd C . s ua veo l en s w e r e n o t f o und a mong t h e E lJ uyo L owe r Magda len ian m ic rof auna , a nd b o th a r e f o und r a r e ly i nC an tab r ian U ppe r P a leo l i th ic l e ve l s . E a ch s p ec ie s i sa s soc ia t ed w i th d ry m eadow c ond i t ion s , a nd t h e i r c om p l e te a b s ence f r om E l J uyo f u r the r r e inf o r ce s d i r ec t i n d ica t i on s o fh um id c ond i t ion s . G a l emy sp y rena icu s ( Py r enean d e sman ) i sa l so a b sen t f r om t h e E l J uyo m ic rof auna . T h i s a l p ine s p ec ie si sn o tf ound t o day i nt h e c oa s ta l l ow land s , b u t i sf ound i ns ome P l e i s to cene l e ve l s o f o t he r l ow land s i t e s ( Chap te r8 ) . M ic rof auna a mong t h eL owe r Magda l en ian l e ve l so fE lJ uyo c hanged v e ry l i t t le o ve rt h e i ra ccumu la t ion . O pen , c o ld , a nd h um id h ab i ta ts p ec ie s d om ina te t h et o ta lm ic rof auna b i oma s s t h rough a l l e v e l s . Sma l la moun t so ff o r e s ta nd d ry g r a s s land s p ec ie s b i oma s s a r e p r e sen t . F o r e s t c ove r ed much mo r e o f

T oo th l e ng th a l so r e conf i rmed i d en t if i ca t ion i ns ome c a se s , s u ch a st h o se o f Mu s te la n i va l i s , w ho s e t e e th a r e s im i la r t o mo rpho log ica l ly b u t s ma l le rt h an t h o se o fM e rm inea . T he i d en t if i ca t ion o ft h e e x t inc t s p ec i e s P l iomy s l e nk i w a s a l so r e conf i rmed f r om t o o th l e ng th . T he mu l t ip le s ou rce s o f p a leoenv i ronmen ta l d a ta f r om E l J uyo w e r e c on s id e r ed t oge the r i nC hap te r 7t og i ve am o r e c omp le te p i c tu r e o ft h e w an ing P l e i s tocene l a nd s cape . S ea c o r e f o ram in if e ra ( CL I MAP 1 976 ; Mc In ty r e e t a l . 1 976 ; R udd iman a nd Mc In ty re 1 976 ; S ha ck le ton 1 987 ; S ha ck le ton a nd O pdyke 1 976 ; v an D onk 1 976 ) i n d ica t e t h a t t h e N o r th A t lan t ic s u r f a ce o cean w a te r t empe r a tu re s a t4 2 ° N o r th l a t i t u de ( t he C an tab r ian c oa s t ) a ve raged 1 0° C c o ld e r a t1 8 ,000 B .P . t h an a tp r e sen t . T h i s t empe ra tu r e i n d ica te s p o la r w a t e r c ond i t ion s w i th a t tendan t p ack i c e . B y 1 4 ,000 B .P . ( t he o c c upa t ion o f E l J uyo ) , o cean l e ve l s w e r e a ga in l owe r ing a s c on t inen ta l i c e c ap s e xpanded a ga in a f t e r ap e r iod o fr e t r ea t . O cean l e ve l f l uc tua ted s omewhe r e i n t e rmed ia t e t ot h e 1 00 mg l a c ia l m ax imum l owpo in t a nd m od e rn s e a l e ve l t h a tw a s r e a ched s ho r t ly a f t e r t h e H o locene b egan . S u rf a ce o cean w a t e r t empe ra tu r e s a nd a ccompany ing c l ima t e w e re t h e r e f o r e s omewhe re b e tween t h o se o ft h e g l a c ia l m ax imum a nd t h e w a rme r A t lan t ic c l ima te o f t h e H o locene . L oca l C an tab r ian c l ima t en eed n o th ave f o l lowed p r ec i se ly c hange s i n g l oba l s e a l e ve l . C hange s i n o cean c i r cu la t ion a nd w ea the r p a t ter n s a l so a l low g r ea t v a r ia t ion i nl o ca l c l ima t e : h ence t h e n eed t o e xam ine o t he r m o r e d i r ec t e v idence o f C an tab r ian P l e i s tocene c ond i t ion s . S ed imen to log ica l a na ly s i s o ft h e E l J uyo d epo s i t s s ugge s t s t h a tc o lde r c ond i t ion s r e tu rned d u r ing t h e t i me o fd epo s i t ion 1 6 1

o ft h e u ppe rh ab i ta t ion L eve l s 6 4 ( F reeman e ta l . 1 988 : 111 2 ) , f o l low ing a n e a r l ie rp e r iod o fr e la t ive ly d r ie r , t empe r a te c ond i t ion s i nL eve l s 7 13 . H um id i ty o r t h e s e a sona l c on c en t ra t ion o fp r ec ip i ta t ion h ad i n c r ea sed b yt h et i me o fd epo s i t ion o fL eve l 6s u ch t h a tt h ew a t e rt a b le o ft h ec ave r e ached t h i s l e ve l , d epo s i t ing c l ay s . B o th L eve l 5( as t e r i le f l ow s tone l a ye r ) a nd L eve l 4s how a n i n c r ea se i ng bou l i s , w h ich m ay i n d ica te a n i n c r ea se i nc o ld r e g ime d ue t o s pa l l ing o f t h e c ave c e i l ing . L eve l 3w a sf o rmed b yt h ec o l lap se o ft h em a in e n t rance f o l lowed b yf u r the rs e a l ing o ft h ev e s t ibu le d epo s i t s w i th a l a ye r o f t r av e r t ine , s ugge s t ing a c l ima t e r e g ime o f h i gh h um id i ty f o l lowed b y ap e r iod o fc o ld ,d ry c ond i t ion s . T he l a rge m amma l , m a r ine mo l lu sc a nd f i sh r ema in s i n t ro d uced t oE lJ uyo d o n o t g i ve ad ef in i te p i c tu r e o fp a leoenv i r o nmen t . T h i s s i tua t ion i sd ue t o t h e w ide e nv i ronmen ta l t o le rance s o f t h e se s p ec ie s , a lmo s t a l o f w h ich a r e f o und t o day ( o r w e r e e x t i rpa t ed l o ca l ly i n h i s to r ica l t i me s ) i n C an tab r ian S pa in . E l J uyo ' s l a rge m amma l f a una ( i n d e s c end ing o r de r o fi mpo r tance ) c on s i s t o fr e d d ee r , r o e d ee r , b i son , h o r se , i b ex , b oa r , l i on a nd w o lf . T he se s p ec ie s a r e n ow o r w e r e f o rme r ly f o und t h roughou t E u rope . R ed d e e r t h r ive o n m ixed f o r e s t /open h ab i ta t s w i th t h e i r e dge e f f ec t s a nd r e su l tan t b r ow s ing f o l iage . B o th r o e d ee r a nd b oa r , f o und i nm uch s ma l le r a moun t s t h an r e d d ee r , p r ef e r f o r e s t h ab i ta t s . H o r se s a nd b i son a r e o pen g r a s s land /pa rk land s p e c i e s , a nd e a ch i sf o und i ns ma l l a moun t s t h roughou t t h e s i t e l e ve l s . I b ex a r e a n u p land s p ec ie s t h a t w e re w i th in t r an s p o r tab le r a nge o fE lJ uyo e ven d u r ing w a rme r c l ima t ic e p i s ode s . T he l a rge f a una t h e r ef o r e i n d ica t e ar a nge o fh ab i ta t s p r e sen t i n u nknown p r opor t ion s . Mo l lu scan f a una w e r e s im i la r ly i n conc lu s ive , s i nce t h e t wo p r ima ry h a rve s t s p ec ie s P a te l la v u lga ta a nd L i t to r ina l i t to rea a r e f ound t h roughou ta w ide r a nge o f A t lan t ic c oa s t l ine s , b o th n o r th a nd s ou th o f C an tab r ian S pa in . Wa rm a nd c o ld i n d ica to r s p ec ie s , r e t r i eved i nv e ry s ma l l a moun t s , p r ov ide c onf l ic t ing o ri n con c l u s ive e v id ence . T he m any s a lmon id r ema in sr e cove red a r e a l so e qu ivoca l i n d ica to r s o fp a leoenv i ronmen t , s i nce moder n S pa in a nd P o r tuga l a r e a tt h e s ou the rn e dge o ft h e i r s pawn i n gr i ve rr a nge ( wh i ch e x tend st om oder nS cand inav ia ) . P a lyno log ica l a na ly se s o f f e r af i ne r r e so lu t ion o f t h e p a l e oenv i ronmen t o f E l J uyo , b u t s u f f e r f r om t h e l i m i ta t ion s i n he r en t t o t h i s t y pe o f r e sea r ch . P r ev iou s i n t e rp re ta t ion ( Boye r -K le in a nd A r . L e ro i -Gou rhan 1 987 ; B oye r -K le in 1 988 ; A r . L e ro i -Gou rhan 1 994 ) i n d ica ted t h a tt h e l owe r l e v e l s ( 1 2 -7 ) o fE l J uyo c ou ld b e c o r r e la t ed w i th t h e r e la t ive ly t empe ra t e P r e -Bö l l ing c l ima t ic p ha se . T h i s i n t e rp r e ta t ion i s b a sed u pon t h e p r e sence o f mode ra t e l e ve l s o f t r ee a nd s t eppe p o l len a nd t h e d om inance o fE r icaceae ( h ea the r ) p o l l e n . T he u ppe rl e ve l so fE lJ uyo ' sL owe r Magda len ian o ccu p a t ion w e re a s s igned t ot h e D rya s Ic l ima t ic p ha se , s i nce a m a jo rd e t e r io ra t ion o fc ond i t ion sw a s i n d ica ted b yt h e d om i n ance o fs t epp ic p o l len s a tt h e e xpen se o ft r ee a nd s h rub s . S anchez G of i i ( 1 993 ) r e in te rp r e t ed t h i s p o l len d a ta a s s t i l l r e f lec t ing t h e c o ld , d r y c ond i t ion s o f D rya s Ii nt h e u pp e r l e ve l s ( 6 -3 ) o ft h e s i t e , w i th t h e l owe r L eve l s 1 2 -7 n o t a s s i gnab le t ot h e P r e -Bö l l ing . T he l owe r l e ve l s t h e r ef o r e a l so m ay h av e b een d epo s i ted d u r ing D rya s c ond i t ion s , w h ich i s i na c co rdance w i th r a d ioca rbon d a t ing f r om t h es i t e .

T he m a c robo tan ica l r ema in s f r om E l J uyo g i ve s im i la r ly m ixed p a leoenv i ronmen ta l s i gna l s , w i th a v a r ied g r oup o f s p ec ie s p r e s en t . L eve l 8h a s y i e ld ed ap ropo r t iona l ly h i gh a bundance o fm a r sh -f avo r ing p l an t s , i n d ica t ing w e t t e r c on d i t ion s d u r ing i t s f o rma t ion o r t h e m o r e i n t en s iv e e xp lo i ta t i on o ft h e n ea rby m a r she s a nd r i a s ( e s tua r ie s ) . S uba lp ine t a xa a ppea rd u r ing t h e d epo s i t ion o fL eve l4 ,r e inf o r c ing t h e i n t e rp r e ta t ion t h a t c o ld e r c ond i t ion s r e t ur n ed i nt h e l a t e r o c c upa t ion o fE l J uyo ( t hough t h e mo r e i n t en s ive e xp lo i ta t ion o fa l p ine r e sou r ce s i sa n a l ter na t ive p o s s ib i l i ty f o rt h e i n t ro d uc t ion o fs ome t a xa ) . T h roughou t a l l e ve l s , t a xa f a vo r ing d i s tu rbed h ab i ta t s a r e a bundan t . T he p r e sence o ft h e se t a xa i n d ica te s t h e s e lec t ed n a tu r e o ft h e m a c robo tan ica l r ema in s i n t roduced t o t h e s i t e b y d e l ibe ra t e a nd a c c iden t a l h uman a c t iv i t ie s , s i nce h uman o ccupa t ion i t s e lf p roduce s d i s tu rbed l o ca l h ab i ta t s . T hu s , ac u l tu ra l f i l t e r h a s i n l a rge p a r t a f f e c ted t h e r e la t ive a bundance s o ft h e se r ema in s a nd l e s sened t h er e so lu t ion o ft h e i rp a leoenv i ronmen ta li n f o rma t ion . U s ing t h e se m any d a ta s ou r ce s i nc on junc t ion w i th t h e m i c r of auna s amp le , t h e p a leoenv i ronmen t a l p i c tu r e o fE l J uyo i sf a r c l ea r e r . T empe ra tu r e s w e re g ene r a l ly c oo l , i n t e rmed i a t eb e tween t h e g l a c ia lm ax imum o f2 ,000 18 ,000 B .P . a nd t h e t empe ra t e H o locene p e r iod . L a rg e m amma l i n d ica to r s p ec ie s a g re e w i th t h e i n t e rp r e ta t ion t h a t o pen c ond i t ion s w e r e m ixed w i th a t l e a s t s ome a r ea s o f f o re s t c ove r a nd g r a s s land a mong h ea th . T he r e la t iv e a moun t s o f t h e se v ege ta t ion t y pe s a r e b e s tr ef l ec ted b yt h e m ic rof auna , w ho se a c cumu la t ion d i d n o t p a s s t h rough a c u l tu ra l f i l t e r . H igh h um id i ty i sr e f lec ted b y t h e m ic rof auna t h rough a l l e ve l s . Mo i s t h ab i ta t m i c rof auna a r e a ccompan ied b y m a c robo tan i c a l r ema in s o fm a r sh p l an t s i n o ccupa t ion s b e low L eve l 4 . S ed imen to logy , p a lyno logy a nd m a c robo tan ica l a na ly s i s a l a g r ee w i th t h e m ic rof auna l d a ta t h a t s ome d e te r io r a t ion o f c l ima t e o ccu r r ed d u r ing t h e d epo s i t ion o fL eve l4 ,i n d ica t ing t h a t s ome c oo l ing a nd d r y ing o ft h e c l ima te o ccu r r ed . D u r i n gt h i st i me t h e a l p ine z one i n t ruded t ol owe ra l t i tude s ,w i th s ome p l an ta nd m ic rof auna t a xa p r e s en ti nt h i s l ow land s i t e . T he s ma l le rm ic rof auna l r ema in s ( Rod en t ia , I n sec t ivo ra , a nd C a rn ivo ra ) f r om C an tab r ian s i te s a nd t h e i n te rp r e ta t ion o f p a leoenv i ronmen t f r om t h e i r r e la t iv e p r opo r t ion s w e r e p r e s e n ted i nC hap te r8 . S ome l e ve l s f r om t h e Me so l i th ic p e r iod a nd o t he r l a t e r e nv i ronmen t s w e re p r e sen t ed f o rp u rpo se s o f p a leoenv i ronmen ta l c ompa r i son a c ro s s t h e P l e i s to c ene /Ho locene t r an s i t ion . S ome s i t e s f r om j u s t o u t s ide t h e C an tab r ian r e g ion ( Py r enean F rance , N ava r ra p r ov ince i n S pa in )w e r es im i la r ly i n c luded . T he r ema in sf r om t h e se s i te s r e inf o r ce i n t e rp r e ta t ion so ft h ep a t ter nd e r ived f r om E lJ uyo . T he M idd le a nd U ppe r P a leo l i th ic e nv i ronmen t s o f C an tab r ian S pa in w e r ed om ina ted b yo p en ,h um id -a s soc ia t ed s p ec ie s . M ic ro t ine s a r et h e mo s tn ume rou sr oden t s i nn ea r ly e ve ry l e ve l f r om a l s i t e s , p r ima r i ly A r v i co la c f . t e r re s t r i s a nd M ic ro tu s o e conomu s . A t many o l d e r e xcava t ion s , o n ly t h e r e la t ive ly l a rge t a xa A r v ico la a nd T a lpa w e r e r e cove red . S ome d ry g r a s s land m ic ro t ine s w e r e p r e sen t (M a r va l i s a nd M a g re s t i s ) ,t h ough g ene ra l ly i nl owe rp r opo r t ion s . I n sec t i v o r e s w e r e a l so t y p ica l ly d om ina ted b y h um id m eadow a nd w a t e r e dge s p ec ie s ( So r ex c orona tu s , S . m inu tu s , N eomy s f od ien s a nd N . a noma lu s ) a tt h e e xpen se o f d r y g r a s s land s p ec ie s ( C roc id u ra r u s su la a nd C . s ua veo l en s ) . C o lde r e p i 1 62

s ode s a r e i n d ica ted b y t h e a ppea rance o fa l p ine / t und ra t a xa ( M ic ro tu s n i va l i s , G a lemy sp y rena i cu s a nd M i c ro tu s g rega l i s ) i nt h e c oa s ta l l ow land z one . Wa rme r e p i sode s a nd t h e r e tur n o ff o r e s t ed c ond i t ion s w i th t h e o n se t o ft h e H o locene a r e i n d ica t ed b y t h e i n c rea sed p ropo r t ion o f A pod emu s a t m any s i t e s , r e inf o r ced b y t h e p r e sence o f G l i s g l i s a nd C l e thr ionomy sg la reo lu s . Mode rn C an tab r ian r e g ion f a una s a r e c ha ra c te r iz ed b y a n i n c rea sed i mpo r tance o f mu r id s p e c i e s , i n c lud ing t h e i n t roduced c ommen sa l /ag r icu l tu ra l ly a s s o c ia ted t a xa ( Ra t tu s , Mu sm u scu lu sa nd M i c romy sm inu tu s ) . A n i mpo r tan t s ubca tego ry o fs ma l l m amma l w a s r e cove r ed i ns i gn if i can t a moun t s f r om t h e f i ne s c r een d epo s i t s o f E l J uyo : t h er ema in s o fm ed ium s i z ed f u r -bea r ing s p ec ie s , mo s t o fw h ich b e long t o O rd e r C a rn ivo ra . Wh i le s ome o ft h e se s p ec ie sf a l lb e low t h e u ppe rb ounda ry o f 5k gu s ed i nd e l im i t i ng m ic rof auna , many d o n o t ( a t l e a s t a s a du l t s ) . T he se l a rge r s p ec ie s , w h i le f i t t ing u nde r t h e c a t ego ry o f m ic ro m amma l s , a r e g ene ra l ly l a rge r t h an t h e t y p ica l r oden t a nd i n sec t ivo r e f a una f a l l ing p r ey t ob a rn o w l s . A s s u ch , t h e se t a xa e xpe r ience d i f f e r en t t a phonom i c p ro ce s se s , i n c lud ing s ou rce o fm or ta l i ty , m anne r o ft r an spo r t t ot h e s i t e , p a t t e rn a nd d eg r ee o f s k e le ta l e l emen t d e s t ruc t ion , a nd d epo s i t ion w i th in t h e s i t e . Wh i le i ti sp o s s ib le t h a t s ome o ft h e se i n d i v i dua l s ( e spec ia l ly t h e s ma l le s t s p e c ie s s u ch a s Mu s te la n i va l i sa nd M . e rm inea o rj uven i le so ft h e l a rge rs p ec ie ss u ch a s 07c to lagu s c un icu lu s ) w e r e i n t roduced a s t h e p r ey o f o w l s , t h e p a t t e rn o fs k e le ta l e l emen t r e cov e ry s ugge s t s t h a t t h e se w e r e b r ough t i n to t h e c ave a st h e d i r ec t a c t ion o fh u m an s . T he d epo s i t iona l p a t te rn o f t h e se f u r -bea r ing m am m a l s i sa l so i nd i rec t c on t ra s tt ot h a t o fh edg ehog ( E r inaceu s e u ropaeu s )r ema in sr e cove r ed f r om E lJ uyo . Wh i le t h i s s p e c i e s i so f as im i la rs i ze t os ome f u r -bea r e r s ( a bou t 1k g a du l t m a s s ) , i ti sn o t au sab le s ou rce o ff u r d ue t o i t s p ro t ec t ive s p ine s . E lJ uyo h a s y i e lded t od a t e 7 2 f oo t e l emen t s , p r ima r i ly p ha l a nge s a nd m e tapod ia l s o f c a rn ivo r e s i nt h e s i ze r a nge o f e rm ine ( M u s t e la e rm inea ) , f e r re t (M p u to r iu s ) , a nd m a r t en ( M ar te s s p p . ) . T he p r e sence o ft h e se e l emen t s i si ns t r ik ing c on t ra s t t ot h e v i r tua l l a ck o fa ny o t h e r e l emen t s , i n c lud ing t e e th , f r om t h e se t a xa ( Chap te r 5 ) . N on -ca rn ivo r e s o ft h i s s i ze r a nge ( s uch a s r a bb i t o r h edgehog ) d o n o t s how t h i s s ame e l emen t r e p re sen ta t ion d i sc repancy , n o r d o s ma l le r , m o re f r ag i le r oden t a nd s ma l l i n sec t ivo r e t a xa . D if f e r en t ia l d e s t ruc t ion o fn on -f oo t e l emen t s o ft h e f u r -bea r ing s p ec ie s t h e ref o r e m ay b e r u led o u t . T he a u tho r a t t r ibu t e s t h e p r e p onde rance o f f u r -bea r ing mamma l f o o t e l emen t s t o t h e i r s e lec t ive i n t roduc t ion t ot h e s i te b y ah uman a gency . S uch a p a t ter n i st h e l o g ica l r e su l t o f f o o t b one s l e f t i n ap e l t t h a t w a s s k inned f r om a n a n ima l t r apped o r h un t ed a way f r om t h e s i te , w i th t h e r ema in ing c a r ca s s i n c lud ing a l o t he r s k e le ta l e l emen t s d i sca rded i nt h e f i e ld . Mu s t e l id m ea th a s a s t rong , u np lea san t f l avo r d ue t o t h e i r s t rong s c en t g l and s ( p e r sona l o b se rva t ion ) a nd may h ave b een t h e r ef o r e al e s s d e s i rab le f o od c ho ice . T he se s p ec i e s w e r e t h u s p robab ly a cqu i red w ho l ly f o r t h e i r p e l t s , w i th n o o t he r a r cha eo log i c a l ly r e cove rab le p o r t ion o ft h em r e a ch ing t h es i t e . T h i s p a t t e rn o fi n c r ea s ing e xp lo i ta t ion o ff u r -bea r ing s p ec ie s i sr e f lec ted t h roughou t t h e C an tab r ian U ppe r P a leo l i th ic . T he se t a xa a r en ea r ly n on -ex i s ten t i nt h e f ew A cheu lean l e v -

e l s a nd a ppea r o n ly s po rad ica l ly i n Mou s te r ian l e ve l s . T he E a r ly U ppe r P a leo l i th ic ( Au r igna c ian , C ha t e lpe r ron ian a nd U ppe r P e r igo rd ian /G rave t t ian ) w i th i t s p r e sumed a r r iva l o f a na tom ica l ly mode rn h uman s h a s a n i n c rea sed a bundance a nd d i ve r s i ty o f t h e se s p ec ie s , i n c lud ing f o r t h e f i r s t t i me Mu s te la s pp . S ke le ta lp a r tr e p re sen ta t ion f o rt h i sp e r iod d oe s n o ti n d ica t e ap a t ter no fs k inn ing i nt h ef i e ld a nd t r an spo r t ing f oo t e l emen t s ( pha lange s , m e tapod ia l s , c a rpa l s / ta r sa l s ) b a ck t oc ave s i t e s a l ong w i th f u r s , t h ough s amp le s i z e s a r e g ene r a ly q u i te s ma l l . T he p r e sence o f t h e se s p ec ie s i nh uman h ab i ta t ion l e ve l s d u r ing t h i s p e r iod t h e ref o re m ay b e d ue a s much t o i n t e rm i t ten t o ccupa t ion o f s i te s b y t h e se s ma l l p r eda to r s o rt h e i ri n t roduc t ion a st h ep r ey o fl a rge rp r eda to r s a st os po rad ic u se b yh uman s . T he L a te U ppe r P a leo l i th ic ( So lu t rean , Magda len ian a nd A z i l ian ) i n C an tab r ian S pa in i sc ha ra c te r iz ed b y af u r the r i n c r ea se i nf u re xp lo i ta t ion . T h i s p a t ter nb ecome s mo s t e v i d en t i n t h e Magda len ian p e r iod , w he r e s pec ie s a bundance a nd d i ve r s i ty i n c r ea se s i ns eve ra l s i t e s , a nd b ody p a r t r e p r e s e n ta t ion ( a t E l J uyo ) v i r tua l ly r u le s o u t o t he r e xp lana t ion s . T he A z i l ian p e r iod s how s a n a ppa r en t d ec l ine i n h un ted / t rapped s p ec ie s d i ve r s i ty a nd a bundance . T h i s t r end p e rhap s r e su l t ed f r om a d e -empha s i s o f f u r a cqu i s i t ion r e l a t ing t ot h e g ene ra l c l ima t ic a me l io ra t ion a tt h e c l o se o ft h e P l e i s tocene . I tm ay a l so r ef l ec t d ec r ea sed d en s i t ie s o fs ma l l a nd m ed ium p r eda to r s i na n i n c r ea s ing ly h uman -a l t e r ed e n v i ronmen t . U ppe r P a leo l i th ic h uman -env i ronmen t i n t e ra c t ion i n C an tab r ian S pa in e xh ib i t s a n a c ce le ra t ing p a t ter n o fs e pa ra t i on o fh uman a c t ion s f r om t h e d i r ec t a nd i mmed ia t e c on s t ra in t s o fe nv i ronmen t . T h i s i n te ra c t ion i n c lude s t h e c on t r o l led h a r ve s to fr e dd ee r , i b ex , s a lmon a nd m a r ine m o l lu sc s b y c oope ra t ive h uman a c t ion . O the r c hange s i n c lude t h e e xpan s ion i n to p r ev iou s ly l e s s i mpo r tan t r e sou r ce s ( a v if auna a nd f u r t r app ing ) a nd mo re o rgan iz ed a nd s p ec ia l iz ed l a nd s c ape u s e . T he se c hange s a r e v i s ib le a s e a r ly a st h e E a r ly U ppe r P a leo l i th ic a nd g r ea t ly i n c r ea se t h roughou t t h e L a t e U ppe rP a leo l i th ic . T h i ss h if ti smo re r ema rkab le c on s ide r ing t h e m a jo r a nd m ino r f l uc tua t ion s o ft h e P l e i s tocene c l ima t e , p r oceeded f r om v a c i l la t ing i n t e r s tad ia l c ond i t ion s , t h rough t h e i n t en se c o ld o ft h e g l ac ia l m ax imum c a . 2 2 ,000 18 ,000 B .P . a nd t h e s t epw i se r e t rea t / r eadvance o fg l a c ie r sb ef o r et h e t empe ra t e c ond i t ion s o f t h e H o locene , w h ich b egan a bou t 1 0 ,000 B .P . T he p r ox im i ty o fC an tab r ian S pa in 's m u l t ip le a nd d i ve r se e nv i ronmen ta l z one s i sa l so s t r ik ing : l i t to ra l , r i ve r ine , c oa s ta l l ow land a nd mon tane z one s a r e w i th in e a sy m u tua l r e a ch . A cce s s t o v e ry d i f f e r en t e nv i ronmen ta l r e s ou r ce s w a s a va i lab le o ve r s pa ce b y p hy s ica l t r ave l o ve rt h e l a nd scape a nd t h rough c hange si ne nv i ronmen to ve rt i me . T he U ppe r P a leo l i th ic C an tab r ian s s how a n a b i l i ty o r p ro p en s i ty t of r ee t h em se lve sp rog r e s s ive ly f r om t h e v aga r ie s o f c hang ing e nv i ronmen ta l c ond i t ion s . W i ld r e sou r ce h a rve s t i n g i n c r ea sed i ni n t en s i ty t h roughou t t h i s p e r iod d e sp i t e m a j o r c l ima t ic f l uc tua t ion s i n t h i s o pen , h um id c oa s ta l z one . I n s tead o f a bandon ing t h e r e g ion d u r ing t h e g l a c ia l m ax i mum a nd o t he r c o ld p eak s , t h e C an tab r ian s s t ead i ly i m p roved a nd r ef ined t h e i r l a nd scape u t i l iza t ion . S ub s i s t ence f o cu sed u pon c on t inua l ly o r s e a sona l ly c oncen t ra t ed r e s ou r ce s a nd e xpand ed i n to o ne s p r ev iou s ly u nd e r exp lo i t ed , 1 6 3

a nd s ub s i s tence a nd s e t t lemen t s y s t em s s h if t ed w i th l o ca l o r s e a sona l s h if t s i n r e sou r ce a bundance . T he se c hange s i n r e sou r ce u s e ( away f r om mo r e r a ndom e xp lo i ta t ion a nd t o w a rd sp l anned , i n ten s ive h a rve s t ing a te ff i c ien t l o ca t ion sa nd t i me s w i th a pp rop r ia te ly e ng inee r ed t e chno logy ) i n m any w ay sp r e sage st h e c hange sn ece s sa ry f o ra g r icu l tu ra l s y s t em s t oa r i se . T he se c hange s i nC an tab r ian S pa in , t h ough n o t d i r e c t ly l e ad ing t oa g r icu l tu r e , d o i n d ica te l o ca l ly t h e p o s s ib le e vo lu t ion o fi n c r ea sed s o c ia l c omp lex i ty . F u r the r e v idence o ft h i s p a t te rn i sf ound i nt h e a r t ( Conkey 1 978 , 1 980 ; F r ee m an e ta l . 1 987 ) a nd l a nd scape u s e p a t ter n s ( F r eeman 1 994 ) o ft h ep e r iod . P a leo l i th ic e nv i ronmen ta ls t udy mu s tb e p l anned a sp a r to fa r e sea r ch p rog ram b ef o re e x cava t ion b eg in s . R ecove ry o f n ece s sa ry d a ta , e spec ia l ly i nt h e c a se s o fm ic rof auna , p o l len a nd m a c robo tan ica l r ema in s c anno t t a ke p l a ce r e t roa c t iv e ly ( F reeman e ta l . 1 998 ) . S ed imen ts amp le s f o rp o l len a na ly s i s m u s t b e d e r ived f r om f r e sh ly e xpo sed d epo s i t s t om in im iz e t h e r i sk o f c on tam ina t ion a nd p r ef e r ab ly t a ken a t mu l t ip le p o in t so ve rt h ee xpo sed s u rf a ce o f al e ve lt oa d ju s tf o rs p a t ia l v a r ia t ion s i np o l len f l ux . Ma c robo tan ica l s mu s t b e o b ta ined t h rough f l o ta t ion o f s e d imen t s a nd t h e r ef o r e a r e o b ta ined d u r ing t h e i n i t ia l p roce s s ing o f e xcava ted s e d imen t l o t s . M ic rof auna a r e s im i la r ly o b ta ined t h rough f i ne s c r een ing . T he se s t ud ie s t h e r ef o r e c anno t b e c a r r ied o u t r e t roa c t ive ly u pon a l r eady e xcava ted c o l lec t ion s , a s c an m ean ingf u l a na ly se s o fl a rge f a una o rm o l lu sc s . Moder n e xcava t ion s i n C an tab r ian S pa in h ave b egun t oi n co rpo ra t et h e se p rocedu r e s f r om t h e o u t se t o f e xcava t ion p l ann ing , a s a t E l J uyo ( F reeman e ta l . 1 988 ; F r eeman e ta l . 1 998 ) a nd t os ome e x t e n t a tL a R ie ra ( S t rau s a nd C la rk 1 986 ) , E ka in ( A l tuna a nd Me r ino 1 984 ) , Ama lda ( A l tuna e t a l . 1 990 ) a nd E r a l la ( A l tuna e ta l . 1 995 ) . F u tu r e r e sea r ch s hou ld f o l low s e ve ra l p a th s ( B ro thwe l l a nd J one s 1 978 ) . O ne a r ea i st h ee s tab l i shmen to ff u r the rc r i t e r ia f o ri d en t if y ing c l o se ly r e la t ed s p ec ie sb y i s o la t ed t e e th a l one . S uch s e pa ra t ion i s n ece s sa ry f o r a r cha eo log ica l s amp le s , w he re c omp le te c r an ia a r er a r ea nd mo s tm ea su r emen t st y p i c a l ly u s ed f o r s pec ie s i d en t if i ca t ion a r e n o t p o s s ib le . T axa t h a t s hou ld b e r e sea r ched i nt h i s m anne ra r eA rv i co la , T a lpa a nd A pod emu s . Sma l l m amma l s a r ea l so p o t en t ia l ly i d en t if i a b le b y t h e i r p o s tc ran ia l r ema in s ( G raham a nd S aund e r s 1 978 ) . T he se d a ta m ay h e lp t o i d en t if y a c cumu la t ing p r eda to r s b y t h e i r p a t t er n s o f p r ey p o s t c ran ia l d e s t ruc t ion ( Pok ine s a nd K e rb i s P e t e rhan s 1 998 ) , w h ich i nt ur n a i d s i n p a leoenv i ronmen ta l i n t e rp r e ta t ion . A ge ing c r i te r ia f o r s ma l l m amma ls p ec ie sb yt o o th w ea ro rp o s t c ran ia lf u s ion w i l l a l so a i d i np r eda to r i d en t if i ca t ion , y e t r e sea r ch i n to t h i s t o p i c h a s b een h i gh ly s po rad ic . T he c on t inua l ly g row ing mo la r s o f m ic ro t ine s r e p r e sen t ap a r t i cu la r p rob lem i n t h i s r e ga rd . P r ey a ge p rof i le s a r e af u r the r c r i t e r ion i nt h e i d en t if i ca t ion o fa c cumu la t ing p r eda to r . A dd i t iona lo w lp e l le ta nd t r app ing s t ud ie s o ft h e a l t i tud ina l r e la t ion sh ip s o fC an tab r ian m ic ro f a una w ou ld a l so b e t t e r i l um ina t e t h e a l t i t ud ina l s h if t s i n s p ec ie s r a nge s t h a t a ccompany e p i sode s o fa l p ine g l ac ia t ion . F u r the r i n ve s t iga t ion o ft h e r e la t ion sh ip s b e tween l o ca l s p e c i e s b i ome t r ic s o ve r d i f f e r en t h ab i ta t s a nd c l ima te s i nt h e i r m oder nr a nge s( K le in 1 99 1) m ay a l so a low i n t e rp r e ta t ion o f p a s t c hange s i n e nv i ronmen ta l p a rame t e r s ( t empe ra tu r e o r r a inf a l l ) .

Wh i le b a rn o w l b ehav io r i sw e l l r e sea r ched , t h a t o f o t he r p o ten t ia l a c cumu la t ing a genc ie s h a s b een l e s s s o . Mo r e r e s e a r ch i n to t h e b i a se s t h ey m ay c au se i np a leoeco log ica l i n t e rp r e ta t ion b ya f f ec t ing r e la t ive p ropo r t ion s o fp r ey r ema in s i sn eeded . T aphonom ic r e ana ly s i s o fa l r eady e xcava t ed m i c r of auna lc o l lec t ion sm igh ta l so p rove i l um ina t ing , a low ing af i ne rd eg r e eo fp a leoenv i ronmen ta l i n te rp re ta t ion . S amp le s t h a t m ay b en ef i t f r om s u ch a na ly s i s e x i s t f r om t h e s i t e s o f E r ra l la ( P emän 1 985 ) , A i tzb i ta r te ( A l tuna 1 972 ) , Ama lda ( P emän 1 990 )a nd E ka in ( Zaba la 1 984 ) . A t E l J uyo , m ic rof auna l a ge p rof i le a na ly s i s i n d ica t e s t h a t b a rn o w l s w e r e r e gu la r ly p r e sen t d u r ing t h e w a rme r m on th s o ft h e y ea r , t h ough t h e i r p r e sence d u r ing t h e c o lde r m on th s c anno tb ed i s coun ted . B a rn o w l s a nd h uman s a r eu n l ike ly t o h ave o ccup i ed t h ec ave a tt h e s ame t i me . T h i s p a t t er n , a l ong w i th i n d ica t ion so fh uman f ood a cqu i s i t ion d u r ing t h e c o lde r m on th s o ft h e y ea r r e inf o r ce s t h e i n t e rp r e ta t ion o fs e a sona l h uman o ccupa t ion o ft h i s l ow land s i t e . S im i la r a na ly se s o f m ic rof auna f r om o t he r s i t e s m ay f u r the r i l um ina t e p a t te rn s o fh uman s i t e u t i l iza t ion a nd s e a sona l m ovemen t , a s c oun t e rpo i sed a ga in s tp a t t er n so fn on -human a n ima la c t iv i t ie s . F u tu r er e sea r ch a tE lJ uyo w i l l c oncen t r a t eu pon t h e a cqu i s i t i on o fl a rge rm ic rof auna l s amp le s f r om t h e l owe rs i t e l e ve l s ( 1 1-8 ) , a sw e l l a st h e i d en t i f ica t ion o fb i rd , r e p t i le a nd a m p h ib ian r ema in sa l r eady e xcava ted . T h i s i n f o rma t ion w i l l b e c omb ined w i th n ew l a rge f a una , f i sh , s e d imen to log ica l , a nd m a c robo tan i ca l d a ta a s t h ey a c cumu la t e c on t inua l ly d u r ing t h eo ngo ing e x cava t ion s . P a ra l le l p a th s o f r e sea r ch i n c lude a na ly s i s o f t h e r e la t ion s h ip s b e twe en a n ima l d ep ic t ion s a nd f a una l r ema in s ( Gonza lez S a inz 1 988 ) o r a n ima l d ep ic t ion s a nd s i t e f u nc t i on /h ie ra r chy ( Conkey 1 978 , 1 980 ; F r eeman 1 994 ; F r eeman e ta l . 1 987 ) . R e sea r ch o ft h i s t y pe e xam ine s t h e c ogn i t ive r o le i nh uman p a leoeco log ica l r e la t ion sh ip s a nd h ow h uman p e r cep t ion so ft h ee nv i ronmen tm ay h ave a f f ec ted p a t t e rn s o f r e sou r ce a nd l a nd scape u s e . C on t inu ing r e sea r ch i n to c hang ing p a t t er n s o f P a leo l i th ic r e sou r ce e xp lo i ta t ion o r o ve r exp lo i ta t ion ( K le in a nd C ruz U r ibe 1 987 ; K rupa 1 994 ) w i l l a low f u r th e r i n t e rp re ta t ion o f h uman i n te ra c t ion w i th t h ec hang ing C an tab r ian e nv i ronmen t . P a leoenv i ronmen ta l r e sea r ch i sr e levan t t o mo r e t h an t h e n a tu ra l h i s to ry a nd e vo lu t ion o ft h e s ma l l mamma l s t h em s e lve s , o rt oh uman e co log ica l i n t e ra c t ion s p ec if i c t oe x t inc t h un t e r -ga the r e r g r oup s i n ag l ac ia l e nv i ronmen t t h a t n o l o nge r e x i s t s . T he e vo lu t iona ry p a th t h a t w e a r e o n d i d n o t b eg in w i th t h e I n du s t r ia l A ge o r e ven w i th t h e " Neo l i th ic R evo lu t ion" , b u t t e n s o f t h ou sand s o f y ea r s e a r l ie r a s o u r e nv i ronmen t a l r e la t ion sh ip s b egan t od ev ia te f r om t h o se o f o t he rl a rge s o c ia lp r eda to r s . T h i sd ev ia t ion c an e ven b e s e en t oh ave b egun m uch e a r l ie rw i th t h e e vo lu t ion o fH omo a nd p r eced ing /con cu r r en t d eve lopmen t s s u ch a s b i peda l i sm , i n c r ea s ing r e la t ive b r a in s i z e , a nd t o o l u s e . T he h uman e vo lu t i ona ry n i che t h e r ef o r e m ay b e s e en a st h e d eve lopmen t o f t h e a b i l i ty t oa dap t t h e n a tu ra l e nv i ronmen t t oh uman n eed s ( t he c on s t ru c t ion o f ac u l tu ra l e nv i ronmen t ) r a the r t h an a dap t ion t ot h en a tu ra le nv i ronmen ti t s e lf . T h i s e vo lu t iona r y t r end d oe sh ave l i m i ta t ion s . H uman sm u s to pe ra te w i th in t h e p a rame t e r s s e tb yt h e i rc on t empo ra ry e nv i ronmen t , a l t e ri ta s 1 64

t h ey w i l l , a nd a r e d i scove r ing t h i s f a c t w i th i n c rea s ing i m m ed iacy a sb i od ive r s i ty s h r ink s( Conne l l 1 978 ; H us ton 1 979 ;

c r ea s ing a b i l i ty t oa l te r t h e b i o sphe re b ecome s d i r ec t ly r e le v an t t o u nde r s tand ing t h i s i n c rea sed modern p ropen s i ty t o

S t o rk 1 988 ; W i l son 1 989 ) a nd e nv i ronmen ta l c r ise s g row ( Sou l6 1 992 ) . T he o r ig in s a nd t r a jec to ry o ft h i s p a th o fi n -

d amage t h eb i o sphe re a nd l e a rn ing h ow t oc on t ro l i t .

1 65

APPENDIX A METR ICAL DATA

T ab le A .1 : M ax imum L eng th ( mm )o f, M , o fA r v i co la . E lJ uyo L eve l 1

9

4 .02

6

E lJ uyo

8

7

4 R

4 S

4

Quarry S i te 2

3 . 44

3 .74

3 . 60

3 . 77

3 . 81

4 . 11

3 . 38

3 . 67

3 . 55

3 . 64

3 . 78

3 . 9 1

3 . 90

3 . 46

3 . 49

3 .94

3 . 61

3 . 36

3 . 62

3 . 47

3 . 23

3 . 76

3 . 22

3 . 7 1

3 . 68

3 . 69

3 . 62

3 . 40

3 . 67

3 . 45

3 , 97

3 . 67

3 . 36

3 . 76

3 . 58

3 . 89

3 . 68

3 . 60

3 , 44

3 . 90

3 . 73

3 . 79

3 . 55

3 . 45

4 . 0 1

3 . 56

4 .09

3 . 46

3 . 55

3 . 89

3 . 9 1

3 . 34

3 . 90

3 . 55

3 . 85

4 . 24

3 . 73

3 . 63

3 . 24

3 . 66

4 .65

3 . 61

3 . 75

3 . 23

3 . 78

3 . 51

3 . 6 1

3 . 43

3 . 69

3 . 78

3 . 46

3 . 56

4 . 21

4 . 03

3 . 85

3 . 70

3 . 28

3 . 83

3 . 97

3 . 95

3 . 40

3 . 69

3 . 6 1

3 . 47

3 . 96

4 . 05

3 . 82

3 . 75

3 . 91

3 . 56

3 . 27

3 . 67

4 . 06

3 . 87

3 . 60

3 . 44

3 . 51

3 . 91

3 . 56

4 . 07

3 . 85

3 . 69

4 .01

3 . 73

3 . 70

4 . 16

3 . 87

4 .03

3 . 47

3 . 86

3 . 56

3 . 96

3 . 83

3 . 57

4 . 16

4 .0 1

3 . 96

3 . 70

3 . 73

3 . 35

3 . 57

3 . 79

3 . 62

4 . 05

3 . 62

3 . 65

3 . 84

3 . 59

3 . 47

3 . 52

3 . 83

3 . 50

3 . 67

4 . 0 1

4 . 24

3 . 74

3 . 87

3 . 58

3 . 63

3 . 77

3 . 51

3 . 59

3 . 47

3 . 6 1

4 . 42

3 . 60

3 . 75

3 . 85

3 . 68

3 . 85

3 . 85

3 . 52

3 . 55

3 . 65

3 . 57

3 . 98

3 . 66

3 . 52

3 . 50

3 . 63

4 . 0 1

3 . 9 1

3 . 55

3 . 67

3 . 41

3 . 60

3 . 96

3 . 92

3 . 7 1

3 . 60

3 . 47

3 . 26

3 . 31

3 . 66

3 . 77

3 . 84

3 . 83

3 . 65

3 . 47

3 . 51

3 . 39

3 . 66

3 . 8 1

4 . 4 1

3 . 76

3 . 45

3 . 35

3 . 31

3 . 47 3 . 31

3 . 60

3 . 60

3 . 11

3 . 58

3 . 86

3 . 6 1

3 . 59

4 . 18

3 . 8 1

3 . 85

3 . 51

3 . 97

4 . 43

3 . 31

3 . 54

3 . 6 1

3 . 42

3 . 32

3 . 82

3 . 84

3 . 52

3 . 98

3 . 68

3 . 75

4 . 11

3 . 6 1

4 . 00

3 . 7 1

T ab le A .2 : M ax imum L eng th ( mm )o f 3M 3o fA r v i co la . E l J uyo L eve l 1 2 . 80

9

8

7

6

E l J uyo 4 S

4 R

4

2 / 4 2 . 14

Quarry S i te 2

2 . 38

2 .22

2 . 52

2 . 33

2 . 59

2 . 54

2 . 33

2 . 65

2 . 20

2 . 39

2 . 87

2 . 18

2 . 56

2 . 39

2 . 22

2 . 31

2 . 43

2 .29

2 . 32

2 . 40

2 . 47

2 . 45

2 . 25

2 .07

2 . 31

2 .69

2 . 25

2 . 30

2 . 56

2 . 66

2 . 57

2 . 35

2 .48

2 . 6 1

2 . 2 1

2 . 26

2 . 55

2 . 42

2 . 14

2 . 50

2 . 34

2 . 20

2 . 51

1 . 92

2 . 09

2 . 28

2 . 31

2 . 37

2 . 19

1 . 97

2 . 16

2 . 42

2 . 46

2 . 58

2 .26

2 . 21

2 . 53

2 . 46

2 . 15

2 . 28

2 .26

2 . 28

2 . 26

2 .22

2 . 55

2 . 29

2 .06

2 . 21

2 . 45

2 .60

2 . 13

2 . 43

2 .61

2 . 28

2 . 24

2 . 33

2 . 27

2 . 10

2 .71

2 . 70

2 . 26

2 . 06

2 .96

2 . 41

2 .47

2 .09

2 .61

2 . 05

2 . 26

2 .20

2 . 72

2 .30

2 .04

1 . 92

2 . 41

2 . 44

2 . 57

2 . 31

2 .65

2 . 36

2 .26

2 . 08

2 . 50

2 . 37

2 .48

2 . 29

2 . 43

2 . 39

2 . 53

2 . 45

2 .45

2 . 46

2 .44

1 . 98

2 .03

2 .4 1

1 . 94

2 . 4 1

2 . 16

2 . 37

2 .26

2 .60

2 . 37

2 . 52

2 . 19

2 . 0 1

2 .02

2 .57

2 . 32

2 . 51

2 . 62

2 . 21

2 . 11

2 . 36

2 . 47

2 . 16

2 . 39

2 . 31

2 . 11

2 .26

2 .66

2 . 23

2 . 36

2 . 51

2 . 19

2 . 26

2 . 39

2 . 40

2 .26

2 . 11

2 .05

2 . 12

2 . 58

2 . 45

2 . 20

2 . 55

2 .60

2 .52

2 . 51

2 .40

2 . 11

2 . 13

2 . 27

2 . 30

2 31

2 . 57

2 . 51

2 .44

2 . 26

2 . 30

2 . 57

2 .09

2 .02

2 . 13

2 . 16

2 .63

2 .99

2 . 57

2 . 23

2 .59

2 . 37

2 .22

2 . 22

2 . 23

2 . 32

2 . 24

2 . 17

2 .29

2 .06

2 .25

2 . 86

1 . 96

2 .50

2 . 25

2 . 16

2 . 26

2 . 23

2 . 33

2 . 11

2 .55

2 . 45

2 . 10

2 .60

2 . 61

2 . 27

2 .25

2 .52

1 . 98

2 .50

2 . 21

2 .09

2 .43

2 . 57

2 . 41

2 . 36

2 .52

2 . 28

2 . 34

2 . 82

2 . 15

2 .72

2 . 32

2 . 18

2 . 31

2 . 50

2 . 41

2 .66

2 . 04

1 . 85

2 . 39

2 . 40

2 . 34

2 .65

2 .62

2 . 21

2 .65

2 . 42

2 .26

2 .76

2 . 30

2 . 26

2 .07

2 .70

2 . 21

2 .65

2 . 00

2 . 35

2 .42

2 . 50

2 . 35

2 . 33

2 . 31

2 . 37

1 . 89

2 . 70

2 .23

2 . 54

2 . 40

2 . 23

2 . 45

2 . 18

2 . 35

2 .06

2 . 39

2 . 13

2 . 54

2 . 56

2 . 45

2 .21

2 .77

2 . 27

2 .40

2 . 40

2 .43

2 . 12

2 . 23

2 . 15

2 . 42

2 .62

2 . 31

2 . 15

2 . 80

2 . 32

2 .79

2 . 46

2 . 16

2 .63

2 .76

2 . 19

2 . 34

2 .25

1 67

T ab le A .3 : Max imum L eng th ( mm )o f, M , o fM icro tu s o economu s . E lJ uyo L eve l 1

9

8

7

6 /8

2 .40

2 . 27

2 . 52

2 . 57

2 .57

6

4R

4 S

4

2 . 51

2 .84

2 . 59

2 . 42

2 . 86

2 . 65

2 . 92

2 .46

2 . 56

2 . 53

2 .22

2 . 50

2 .70

2 . 53

2 .82

2 . 55

2 . 47

2 .59

2 .72

2 .70

2 . 45

2 . 46

2 . 54

2 . 59

2 .48

2 ,74

2 . 53

2 . 36

2 . 44

2 .77

2 .58

2 . 74

2 . 65

2 . 56

2 . 55

2 . 38

2 .63

2 . 70

2 .46

2 . 37

2 . 43

2 . 35

2 . 56

2 . 38

2 . 35

2 . 20

2 . 53

2 .81

2 . 54

2 .61

2 .72

172

2 . 54

2 . 55

2 . 37

2 .6 1

2 .60

2 .70

2 . 57

2 . 56

2 . 37

2 . 58

2 .70

2 . 66

2 . 57

2 .7 1

2 . 52

2 . 47

2 . 48

2 / 4

2 .26

2 . 62

2 . 47

2 . 59

2 . 45

2 . 51

2 . 55

2 . 58

2 .23

2 . 67

2 . 45

2 . 37

2 . 81

2 . 67

2 .49

2 .41

2 . 46

2 . 69

2 . 59

2 . 65

2 .67

2 . 68

2 . 08

2 . 59

2 . 68

2 . 50

2 . 48

2 . 61

2 ,72

2 . 55

2 . 50

2 . 49

2 . 79

2 .42

2 . 62

2 . 57

2 .29

2 . 55

2 . 56

2 .47

2 . 45

2 . 61

2 . 33

2 . 26

2 . 41

2 . 50

2 .61

2 . 57

2 . 46

2 . 51

2 . 58

2 .71

2 . 66

2 . 48

2 . 65

2 . 88

2 . 33

2 . 37

2 . 46

2 . 50

2 . 51

2 . 35

2 . 65

2 . 72

2 . 54

2 .43 2 . 43

2 . 60

2 . 46

2 . 58

2 .66

2 . 58

2 . 82

2 . 50

2 .40

2 . 30

2 .46

2 . 57

2 . 51

2 .61

2 . 39

2 .63

2 . 37

2 .76

2 . 45

2 . 36

2 .66

2 . 42

2 . 40

2 .70

2 . 48

2 . 52

2 . 56

2 . 46

2 . 59

2 . 52

2 . 51

2 . 58

2 . 53 2 .74

2 . 55

2 . 51

2 . 84

2 . 25

2 . 44

2 . 56

2 . 65

2 . 81

2 . 47

2 . 44

2 . 62

2 . 36

2 . 55

2 . 47

2 . 69

2 .70

2 . 58

2 . 37

2 . 52

2 . 62

2 . 57

2 . 67

2 . 45

T ab le A .4 : Max imum L eng th ( mm )o f, M , o fP i tymy ss pp . E lJ uyo L eve l 1

9

8

7

2 . 63

2 . 36

2 . 57

6

2 . 4 1

2 . 51

2 .73

4 R

4 S

2 . 22

2 .70

2 . 30

2 . 30

2 . 63

2 . 38

2 . 52

2 .53

2 . 48

2 . 25

2 . 56

2 . 47

2 . 52

2 . 35

2 .35

2 . 66

2 .63

2 .72

2 . 56

2 . 56

2 . 6 1

2 .72

2 . 42

2 .69

2 . 65

2 . 42

2 . 60

2 . 54

2 .43

2 . 51

2 . 31

4

2 . 38

2 . 39

2 .68

2 . 40

2 . 50

2 . 56

2 . 60

2 . 76

2 . 71

2 .64

2 . 42

2 . 55

2 . 38

2 . 64

2 . 48

2 . 46

2 .44

2 . 50

2 .60

2 . 58

2 . 65

2 . 50

2 . 40

2 .72

2 . 55

2 . 72

2 . 27

2 . 64

2 .60

2 . 62

2 . 60

2 . 45

2 . 54

2 .58

2 . 57

2 .65

2 . 37

2 . 52

2 .47

2 .7 1

2 . 50

2 . 46

2 . 52

T ab le A .5 : Max imum L eng th ( mm )o f 2M 2 o fP i tymy s s pp . E lJ uyo L eve l 1

8

7

6

1 . 42

1 . 37

1 . 38

1 . 40

1 . 46

1 . 39

4 S 1 . 48

1 . 38

1 . 44

4 1 . 37

1 . 44

1 . 49

1 . 40

1 . 33

1 . 25

1 . 25

1 . 3 1

1 . 34

1 . 47

1 . 39

1 . 40

1 . 30

1 . 47

1 . 35

1 . 43

1 . 26

1 . 24

1 . 5 1

1 . 42

1 . 45

1 . 3 1

1 . 29

1 . 39

1 . 30

1 . 40

1 . 45

1 . 42

1 . 3 1

1 . 38

1 . 38

1 . 45

1 . 42

1 . 35

1 . 48

1 . 29

1 . 39

1 . 37

1 . 25

1 . 40

1 . 44

1 . 29

1 . 40

1 . 22

1 . 37

1 . 4 1

1 . 43

1 . 40

1 . 20

1 . 43

1 . 37

1 . 30

1 . 44

1 . 30

1 . 46

1 . 39

1 . 50

1 . 26

1 . 36

1 . 50

1 . 29

1 . 38

T ab le A .6 : Max imum L eng th ( mm ) o f 3M 3 o fP i tymy s l u s i -

T ab le A .7 : Max imum L eng th ( mm ) o f 3M 3 o fP i tymy s p y -

t a n icu s .

r ena icu s .

E l J uyo L eve l

E lJ uyo

8

4 R

4 S

1 . 84

1 , 83

1 . 97

1 . 66

1 . 87 1 . 75

E lJ uyo L eve l

Quarry S i te 2

9

8

1 . 76

1 . 66

1 . 5 1

1 . 53

1 . 52

1 . 44

1 . 59

1 . 51

1 . 4 1

1 . 62

1 . 35

1 . 51

1 . 6 1

1 . 42

1 . 3 1

1 . 37

1 . 68

1 . 82

1 . 44

1 . 43

1 . 51

1 . 57

1 . 42

1 . 46

1 . 88

1 . 43

1 . 42

1 . 75

1 . 5 1

1 . 71

1 . 4 1

1 . 53

1 . 46

1 . 59

1 . 5 1

1 . 4 1

1 . 56

1 . 47

1 . 50

1 . 45

1 . 64

1 . 47

1 . 42

1 . 47

1 . 60

1 . 44

1 . 60

1 . 64

1 . 40

1 . 69

1 . 53

1 . 38

1 . 68

1 . 44

1 . 50

1 . 48

1 . 55

1 . 64

6 1 . 81

4 S

4

1 . 62

1 . 59

1 . 75

1 . 60

1 . 65

1 . 67

1 . 66

1 . 76

1 . 75

1 . 62

1 . 93

1 . 65

1 . 78

1 . 67

1 . 98

1 . 85

1 . 59

1 . 67

1 . 77

1 . 87

1 . 84

1 . 75

1 . 64

1 . 76

1 . 64

1 . 75

1 . 72

1 . 45

1 . 69

1 . 90

1 . 81

1 . 6 1

1 . 52

1 . 59

1 . 80

1 . 71

1 . 6 1

1 . 40

1 . 70

1 . 45

1 . 55

1 . 48

1 . 76

1 . 42

1 . 46

1 . 69

1 . 47

1 . 85

1 . 76

1 . 67

1 . 62

1 . 7 1

1 . 45

1 . 92

1 . 38

1 . 58

1 . 42

1 . 77

1 . 9 1

1 . 70

1 . 85

1 . 84

1 . 54

1 . 74

1 . 76

1 . 65

1 . 79

1 . 66

1 . 50

1 . 77

1 . 70

1 . 86

1 . 65

1 . 74

1 . 6 1

1 . 64

1 . 64

1 . 70

1 . 90

1 . 50

1 . 55

1 . 75

1 . 49

T ab le A .8 : Max imum L eng th ( mm )o f, M , o fM icro tu sa g re s t i s larva l i s . E lJ uyo

E lJ uyo L eve l 1 0 2 . 47

9

8

7

6

4 R

4 S

4

2 /4

Quarry S i te 2

2 . 42

2 . 84

2 . 58

2 . 59

2 . 35

2 . 7 1

2 . 66

2 . 74

2 . 6 1

2 . 82

2 . 6 1

2 . 71

2 . 64

2 . 6 1

2 . 40

2 . 46

2 . 79

2 . 32

2 . 34

2 . 58

2 . 46

2 . 53

2 . 8 1

2 . 57

2 . 79

2 . 76

2 . 74

2 . 47

2 . 62

2 . 36

3 . 02

2 . 50

2 . 42

2 . 65

2 . 79

2 . 4 1

2 . 4 1

2 . 67

2 . 26

2 . 76

2 . 55

2 . 48

2 . 69

2 . 69

2 . 69

2 . 64

2 . 73

2 . 8 1

2 . 60

2 . 35

2 . 57

2 . 78

2 . 77

2 . 50

2 . 37

2 . 86

2 . 57

2 . 51

2 . 72

2 . 86

2 . 73

2 . 73

2 . 67

2 . 90

2 . 8 1

2 . 37

2 . 75

2 . 54

2 . 70

2 . 59

2 . 55

3 . 0 1

2 . 86

2 . 38

2 . 82

2 . 74

2 . 78

2 . 36

2 . 67

2 . 90

2 . 64

2 . 73

2 . 74

2 . 7 1

2 . 65

2 . 56

2 . 75

2 . 65

2 . 90

2 . 59

2 . 77

2 . 4 1

2 . 54

2 . 78

2 . 74

2 . 37

3 . 03

1 59

2 . 47

2 . 74

2 . 82

2 . 76

2 . 72

2 . 68

2 . 62

2 . 63

2 . 54

2 . 82

2 . 74

2 . 4 1

2 . 74

2 . 66

2 . 55

2 . 65

2 . 63

2 . 59

2 . 8 1

2 . 75

2 . 68

2 . 64

2 . 44

2 . 70

2 . 73

2 . 77

2 . 87

2 . 93

2 . 72

2 . 60

2 . 9 1

2 . 51

2 . 77

2 . 76

2 . 92

2 . 53

2 . 75

2 . 57

2 . 68

2 . 69

2 . 94

2 . 76

2 . 74

2 . 73

2 . 65

2 . 63

2 . 57

2 . 78

2 . 7 1

2 . 70

2 . 70

2 . 84

2 . 68

2 . 80

2 . 70

2 . 56

2 . 81

2 . 86

2 . 73

2 . 47

2 . 68

2 . 76

3 . 16

2 . 62

2 . 69

2 . 84

3 . 22

2 . 82

2 . 74

2 . 81

2 . 73

2 . 68

2 . 70

2 . 57

2 . 78

2 . 44

2 . 76

2 . 90

2 . 78

2 . 70

2 . 79

2 . 86

2 . 82

2 . 78

2 . 67

2 . 88

2 . 84

2 . 89

2 . 77

2 . 9 1

3 . 23

2 . 8 1

3 . 00

3 . 07

2 . 27

2 . 74

2 . 77

2 . 71

2 . 45

2 . 67

2 . 76

2 . 87

2 . 60

2 . 74

1 69

2 . 44 2 . 37

1 . 63 1 . 75

T ab le A .9 : Max imum L eng th ( mm )o f 2M 2 o fM icro tus a g re s t i s . E l J uyo L eve l 9 1 . 59

8

7

6

4 14

1 . 6 1

1 . 55

1 .59

1 . 59

E l J uyo 4 S

4

2 1 . 56

Quarry S ite 2

1 . 61

1 . 60

1 .62

1 . 71

6 6

1 .64

1 . 59

1 . 50

1 . 54

1 . 76

1 . 63

1 ,69

1 . 52

1 . 45

1 , 65

1 . 60

1 . 57

1 . 65

1 .70

1 . 69

1 . 72

1 . 66

1 .68

1 . 66

1 . 56

1 .59 1 .61

1 . 52

1 . 70

1 . 66

1 .6 1

1 . 46

1 . 45

1 . 61

1 . 62

1 . 59

1 . 56

1 . 61

1 . 65

1 . 62

1 . 58

1 . 53

1 . 72

1 . 66

1 .60

1 . 60

1 . 62

1 . 64

1 .60

1 . 64

1 . 56

1 . 6 1

1 . 63

T ab le A .10 : Max imum L eng th ( mm ) o f, M , o fM icro tu s n i va l i s . E l J uyo ( A l l Leve ls) 2 .61 2 .96 3 .27

T ab le A .11 : Max imum L eng th ( mm ) o f, M , ,

2 M2,

M la nd

I

2M

2

o fA pod e r nu s .

E lJ uyo L eve l 8

6

M I l ow

M I l ow

1 . 7 1

1 . 84

I . 90

4 S M1 l ow

4

8

4 S

4

3

4 S

4

4 S

4

MI l ow

M2 l ow

M2 l ow

M2 l ow

M2 l ow

M 1u p

M Iu p

M2 u p

M 2 u p

1 . 26

1 . 33

1 . 17

1 . 38

1 . 96

2 . 00

1 . 70

1 . 90

1 . 87

2 . 10

1 . 41

1 . 27

1 . 86

1 . 90

1 . 95

1 . 28

1 . 95

1 . 99

1 . 25

1 . 28

1 . 74

1 . 99

1 . 90

1 . 28

1 . 83

1 . 59

1 . 11

1 . 85

1 . 75

1 . 86

1 . 25

1 . 85

1 . 41

. 7 1

1 . 80

1 . 82

1 . 20

1 . 78

1 . 23

. 7 3

1 . 78

1 . 26

1 . 95

1 . 25

. 7 6

1 . 24

1 . 96

1 . 27

. 7 5

1 . 19

1 . 95

1 . 17

. 7 2

1 . 75

1 . 23

1 . 66

1 . 80

1 . 71

1 . 96

1 . 72

1 . 81

E l J uyo Quarry S i te 2 A . f l av ico l l is

. 1 s y lva t icus

MI l ow

M2 l ow

M Iu p

M2 l ow

M1 l ow

M2 l ow

MI u p

M2 l ow

1 . 92

1 . 25

1 . 91

1 . 22

1 . 78

1 . 22

1 . 89

1 . 22

1 . 77

1 . 26

1 . 97

1 . 31

1 . 70

1 . 18

1 . 76

1 . 11

1 . 74

1 . 17

1 . 76

1 . 14

1 . 81

1 . 26

1 . 78

1 . 16

1 . 82

1 . 26

1 . 95

1 . 20

1 . 68

1 . 18

1 . 77

1 . 26 1 . 19

1 . 89

1 . 37

1 . 94

1 . 32

1 . 76

1 . 23

1 . 90

1 . 85

1 . 34

2 .02

1 . 31

1 . 80

1 . 22

1 . 98

1 . 32

1 . 81

1 . 26

1 . 89

1 . 28

1 . 82

1 . 27

1 . 91

1 . 27

1 . 73

1 . 17

1 . 72

1 . 20

1 . 74

1 . 17

1 . 79

1 . 19

1 . 84

1 . 28

1 . 86

1 . 26

1 . 72

1 . 11

1 . 76

1 . 13

1 . 75

1 . 23

1 . 80

1 . 22

T ab le A .12 : Max imum R amu sH e igh ta nd L eng th o f 1M 1 , 2M2 a nd 3M 3( mm ) o fS orex c orona tu s . E lJ uyo L eve l 8

6

4 S

4

8

6

r amus

r amus

r amu s

4 S

4

8

6

4 S

4

8

6

4 S

4

r amu s

M I

M I

M I

M I

M2

M2

M 3

M2

M 3

M 3

M 3

M 3

3 . 88

4 . 44

4 . 36

4 . 18

4 . 11

4 . 14

1 . 67

1 . 50

1 . 45

1 . 34

1 . 32

1 . 16

1 . 23

1 . 27

1 . 06

0 . 98

1 . 05

1 . 04

4 . 34

4 . 25

1 . 40

1 . 42

1 . 45

1 . 27

1 . 27

1 . 22

1 . 24

1 . 08

1 . 0 1

0 . 90

4 . 38

1 . 43

1 . 48

1 . 37

1 . 30

1 . 24

1 . 24

4 . 00

1 . 55

1 . 48

1 . 42

1 . 42

1 . 26

1 . 47

1 . 50

4 . 26

1 . 0 1 0 . 99

1 . 28

1 . 49

1 . 3 1

1 . 44

1 . 22

I . 39

1 . 07

1 . 39 1 . 28

E lJ uyo Quarry S i te 2 R amus

MI

M3

M3

4 . 27

4 . 50

1 . 38

1 . 32

1 . 22

1 . 20

0 . 97

4 . 50

4 . 26

1 . 40

1 . 30

1 . 27

1 . 24

1 . 07

1 . 06 1 . 08

4 . 44

4 . 65

1 . 36

1 . 44

1 . 30

1 . 31

1 . 00

1 . 05

4 . 41

4 . 52

1 . 35

1 . 40

1 . 25

1 . 34

0 . 9 1

1 . 05

4 . 57

4 . 50

1 . 50

1 . 50

1 . 33

1 . 28

1 . 03

1 . 01

4 . 28

4 . 40

1 . 42

1 . 43

1 . 25

1 . 21

1 , 04

1 . 05

4 . 43

4 . 47

1 . 45

1 . 46

1 . 31

1 . 30

1 . 05

1 . 10

4 . 50

4 . 32

1 . 40

1 . 45

1 . 32

1 . 28

1 . 06

1 . 00

4 . 65

4 . 54

1 . 38

1 . 45

1 . 27

1 . 32

1 . 00

1 . 06

4 . 46

4 . 55

1 . 41

1 . 36

1 . 28

1 . 26

1 . 03

1 . 10

4 . 56

4 . 53

1 . 42

1 . 42

1 . 31

1 . 25

1 . 02

1 . 02

4 . 42

4 . 52

1 . 49

1 . 45

1 . 31

1 . 26

1 . 00

1 . 00

4 . 69

4 . 56

1 . 42

1 , 38

1 . 31

1 . 25

1 . 03

1 . 04

4 . 50

4 . 78

1 . 46

1 . 46

1 . 31

1 . 35

1 . 09

0 . 97

4 . 45

4 . 56

1 . 42

1 . 43

1 . 30

1 . 29

1 . 0 1

0 . 98

4 . 36

4 . 45

1 . 50

1 . 37

1 . 34

1 . 31

1 . 00

1 . 00

4 . 50

4 . 39

1 . 41

1 . 46

1 . 29

1 . 27

1 . 08

4 . 54

4 . 56

1 . 45

1 . 45

1 . 29

1 . 30

1 . 00

4 . 57

4 . 40

1 . 48

1 . 40

1 . 31

1 . 32

1 . 05

4 . 47

4 . 37

1 . 49

1 . 36

1 . 39

1 . 23

1 . 03

4 . 57

4 . 40

1 . 40

1 . 32

1 . 25

1 . 21

1 . 00

4 . 50

4 . 42

1 . 40

1 . 40

1 . 32

1 . 25

1 . 00

4 . 56

4 . 13

1 . 44

1 . 42

1 . 30

1 . 34

0 . 95

4 . 43

4 . 59

1 . 40

1 . 51

1 . 28

1 . 29

1 . 04

4 . 54

4 . 26

1 . 40

1 . 42

1 . 28

1 . 30

1 . 00

4 . 74

4 . 21

1 . 43

1 . 43

1 . 25

1 . 27

1 . 01

1 . 31

4 . 55

4 . 54

1 . 42

1 . 37

1 . 23

4 . 42

4 . 50

1 . 50

1 . 35

1 . 28

1 . 04

1 . 00

4 . 73

4 . 20

1 . 41

1 . 32

1 . 28

0 . 95

4 . 45

4 . 26

1 . 43

1 . 46

1 . 32

0 . 97

4 . 44

4 . 32

1 , 41

1 . 25

1 . 10

4 . 35

4 . 43

1 . 45

1 . 30

1 . 00

T ab le A .I3 : Max imum R amu s H e igh ta nd L eng th o f 1M 1 , 2M2 a nd , M 3( mm )o fS o rex m inu tu s . E l J uyo ( A l l L eve ls ) R amus

MI

M2

M3

2 .97

1 . 28

1 . 10

0 .92

3 . 12

1 . 27

1 . 14

0 .95

3 .00

1 . 41

1 . 15

0 .93

3 .26

1 . 27

1 . 13

0 .89

3 .20

1 . 24

1 . 14

0 .92

3 . 15

1 . 09

0 .98

2 .71

1 . 04

0 .92

3 . 14

T ab le A .14 : Max imum L eng th ( mm ) o f, M , a nd ' M ' o fE r i n aceu se uropaeu s .

T ab le A .15 : Max imum L eng th ( mm ) o f , M , „P , a nd 41 }4o f Mu s te la n iva l i s .

E lJ uyo ( A l l L eve l s )

E lJ uyo ( A l l L eve ls )

1l owe r

1u ppe r

5 . 84

M 1 l ow

P 4 u p

5 . 60

4 . 60

3 . 39

6 . 00

5 . 74

3 . 90

6 . 53

6 . 00

4 . 56

6 . 84

6 . 34

7 . 25

1 72

P 4 l ow 1 . 89 2 . 02

REFERENCES

A damczew ska -And rze jew ska , K . A . , 1 967 . Age r e f e rence

A l tuna , J ,& K . Ma r iezku r r ena , 1 982 . R e s to s o seo s d e l y a -

E k -

c im ien to p r eh i s tö r ico d e A baun tz ( Ar ra iz , N ava r ra ) . T raba jo sd eA rqueo log ia Na va r ra 3 ,3 47 -353 .

A l ibha i , S .K . , 1 980 . A n X ray t e chn ique f o r a ge ing b ank v o le s ( C l e th r ionomy sg la reo lu s ) u s ing t h e f i r s t mand ibu -

A l tuna , J ,& K . Ma r iezku r rena , 1 984 . B a ses d e s ub s i s tenc ia

mode l f o r A pod emu sf la v ico l l i s ( Me lch io r , 1 834 ) . o log ia P o l ska S er ia A 1 5 , 7 87 -790 .

d e o r igen a n ima l e n e ly ac im ien to d e E ka in . I n E l Y a c im ien to d el aC ue va d eE ka in ( Deba , G u ipüzcoa ) , e d s . J .

l a r mo la r . J ourna lo fZ oo logy 1 9 1 ,4 18 -423 .

A l tuna & J .M . Me r ino . A lma9a , C . , 1 973 . S u r l a s t ruc tu re d e s p opu la t ion s d e s P i tymy s I b e r ique s . R e v i s ta d a F a cu l tad d e C ienc ia s d e L i sboa 1 7 ,3 83-426 .

S an S eba s t ian : S oc iedad d e

E s tud io sV a sco s , S e r ie B i ,p p . 2 11-280 . A l tuna , J ,& K . Ma r iezku r r ena , 1 985 . B a ses d e s ub s i s tenc ia d e l o s p ob lado re s d e E r ra l la : Mac romam ff e ro s . Mun ibe 3 7 , 8 7 117 .

A l tuna , J . , 1 972 . F auna d e mam if e ro s d e l a s y ac im ien to s p reh i s tö r ico sd eG u ipüz coa . Mun ibe 2 4 , 1 464 . A l tuna , 1 , 1 976 . L o s m am if e ro s d e l y ac im ien to p reh i s tö r ico d e T i to B u s t i l lo ( A s tu r ia s ) . I nE xcavac ione se nl aC ueva d e " T i to B u s t i l lo" ( A s tu r ia s ) , e d s . J .A . Mou re R oman i l lo a nd M . C ano -He r re ra . O v iedo : I n s t i tu to d e E s tud io s A s t u r iano s ,p p . 1 51193 . A l tuna , J . , 1 978 . L os m am ff e ro s d e C ueva Mo r in . I n V ida y Mue r t e e n C ueva Mo r in , e d s . J .G onza lez E chega ray & L . G . F r eeman . S an tande r : I n s t i tuc iön C u l tu ra l d e C an t ab r ia ,p p . 2 01-209 . A l tuna , J . , 1 981 . R e s to s o seo s d e l y a c im ien to p reh i s tö r ico d e l R a sca f io . I nE lP a leo l i t ico S upe r ior d e l aC ueva d e l R a sca i io ( San tand e r ) , e d s . J .G onza lez E chega ray & I . B a rand ia rän Mae s tu . S an tande r : C en t ro d e I nve s t igac ion yMu seo d eA l tam i ra , Monog raf ia N o . 3 ,p p . 2 21-269 .

A l tuna , J .& J .M . Me r ino , e d s . , 1 980 . E l Y ac im ien to d e l a C ueva d e E ka in ( Deba , G u ipüzcoa ) . S an S eba s t ian : S o c i edad d eE s tud io sV a sco s , S e r ie B l . A l tuna , J .& L . G . S t rau s , 1 976 . T he S o lu t rean o fA l tam i ra : T he a r t if ac tua l a nd f auna l e v idence . Z ephy ru s 2 6 -27 , 1 75 -182 . Amo re s , F . , 1 980 . F eed ing h ab i t s o f t h e s t one m a r ten s , Mane sf o ina ( E rx leben , 1 777 ) , i n s ou thwe s te rn S pa in . S äuge t ie r -k und l ichen M i t te i lungen 5 4 , 3 16 -322 . A nde r son , S .& C .A . L ong , 1 961 . Sma l l mamma l s i np e l le t s o f b a rn o w l s f r om M i f iaca , C h ihuahua . Ame r ican Mu s eum N ov i ta te s2 052 , 1 3 . A nd rew s , P . , 1 990 , Ow l s , Ca ve s a nd F o s s i l s . C h icago : U n i v e r s i ty o fC h icago P re s s .

A l tuna , J . ,1 9 85a . I n t roducc iön . Mun ibe , 3 7 :7 -9 . A l tuna , J . ,1 9 85b . D a ta c ione s d e l C 14 . Mun ibe 3 7 , 2 5 -28 . A l tuna , J . ,1 986 . T he m amma l ian f a una s f r om t h ep r eh i s to r ic s i te o fL a R ie ra . I nL a R ie ra C a ve , e d s . L . G . S t rau s & G . A . C la rk . A r izona S t a te U n iv e r s i ty , An th ropo log ica l R e sea r ch P ape r sN o . 3 6 , p p . 2 37 -274

A nd rew s , P . , J .M . L o rd , & E . M . N e sb i t E van s , 1 979 . P a t t e rn s o fe co log ica l d i ve r s i ty i nf o s s i l a nd mode rn m am ma l ian F auna s . B io log ica lJ ourna l o ft h e L innean S oc i e t y 1 , 1 77 -205 . A nd rew s , P . & E . M . N e sb i t E van s , 1 983 . Sma l l m amma l b one a ccumu la t ion s p roduced b y m amma l ian c a rn ivo re s . P a leob io log y9 ,2 89 -307 .

A l tuna , J . , 1 989 . S ub s i s tence d 'o r ig ine a n ima le p endan t l e Mou s t e r ien d ans l ar e g ion c an tab r ique . I n L 'Homme d e Mand e r tha l , V o lume 6 ,e d . M . O t te . E tudes e tR eche r c hes A r cheo log ique sd e l 'Un ive r s i t ed eL iege ,p p . 3 1-44 . A l tuna , J . , 1 990 . C aza y a l imen ta c iön p roceden te d e mac ro -

A nge r sb jö rn , A . &J .E .C . F lux , 1 995 . L epu s t i m idu s . Mam m a l ian S pec ie s4 95 , 1 11 . A ve ry , D . M . , 1 982a . M ic romamma l s a sp a leoenv i ronmen ta l i n d ica to r s a nd a n i n te rp r e ta t ion o ft h e L a te Q ua ter na ry i n

mam if e ro s d u ran te e l P a leo l i t ico d e Ama lda . I n L a C ue va d e Ama lda ( Z e s toa , P a i s V a sco ) , e d s . J .A l tuna , A . B a ld eön & K . Ma r iezku r rena . S an S eba s t ian : S oc iedad d eE s tud io sV a sco s ,p p . 1 49 192 .

t h e S ou ther nC ape P rov ince , S ou th A f r ica . A nna l s o ft h e S ou th Af r ican Mu seum 8 5 , 1 83 -374 . A ve ry , D . M . , 1 982b . T he m ic romamma l ian f a una f r om B o rde r C ave , Kwazu lu , S ou th A f r ica . c haeo log ica lS c ience 9 ,1 87 -204 .

A l tuna , J . ,A . B a ldeön & K . Ma r iezku r rena , e d s . , 1 990 . L a C ue va d e Ama lda ( Z e s toa , P a i s V a sco ) . S oc iedad d eE s tud io sV a sco s .

J ourna l o fA r -

S an S eba s t ian : A ve ry , D . M . , 1 987 . L a te P l e i s tocene c oa s ta l e nv i ronmen to f t h e S ou ther n C ape P rov ince o f S ou th A f r ica : M ic ro mamma l s f r om K la s ie s R ive r Mou th . J ou rna l o fA r c haeo log ica lS c ience 1 4 ,4 05 -421 . 1 73

Ave ry , D . M . , 1 992 . T he e nv i ronmen t o fe a r ly mode rn h u man s a t B o rde r C ave , S ou th A f r ica : M ic romamma l ian e v idence . P a laeogeog raphy , l a eoeco logy 9 1 ,7 1-87 .

P a laeoc l ima to logy ,

F os s i l R ecord , e d s . P . A . A l l i son & D . E . G . B r igg s . N ew Y o rk : P l enum P re s s ,p p . 2 91-335 .

P a B e l t rän , J .F .& M . D e l ibe s , 1 993 . P hy s ica l c ha ra c t e r i s t ic s o f I b e r ian l y nxe s( Lynx p a rd inus ) f r om D ohana , S ou thwe s t -

A ve ry , D . M . , 1 993 . E co log ica l d a ta o n m ic romamma l s c o l l e c ted b yb a rn o w l s T y to a l ba i nt h e We s tC oa s tN a t iona l P a rk , S ou th A f r ica . I s rae l J ourna l o fZ oo logy 3 8 , 3 85 -

er nS pa in . J ourna lo f Mamma logy 7 4 ,8 52 -862 . B e rgman , C .A . , 1 993 . T he d eve lopmen to ft h e b ow i n We s t er n E u rope : At e chno log ica l a nd f u n c t iona l p e r spec t ive . I n Hun t ing a nd A n ima lE xp lo i ta t ion i nt h eL a te rP a laeo -

3 97 . A ve ry , D . M . , 1 995a . M ic romamma l ian s t ud ie s : I nf o rma t ion

l i th ic a nd Me so l i th ic o fE ura s ia , e d s . G . L a r sen P e t e rk in , H . M . B r icke r&P . Me l la r s . A rcheo log ica l P ape r so ft h e Ame r ican A n th ropo log ica l A s soc ia t ion , N umbe r 4 ,

f r om t h e p a s t o fr e levance t ot h e f u tu re . T ran sac t ion s o f t h eR oya lS oc ie ty o fS ou th Af r i ca 5 0 ,4 1-47 .

p p .95 105 . Ave ry , D . M . , 1 995b . P hy s ica l e nv i ronmen t a nd s i te c ho ice i nS ou th A f r ica . J ou rna l o fA rchaeo log ica l S c ience 2 2 , 3 43 -353 . B a i ley , G . , 1 983 . E conom ic c hange i n L a t e P l e i s tocene C an tab r ia . I n H un te r -Ga the re r E conomy i n P reh i s tory e d . G . B a i ley . C amb r idge : C amb r idge U n ive r s i ty P re s s , p p . 1 49 165 .

B e r ry , R . J . , 1 978 . M ic ro -evo lu t iona ry s t ud ie s i na n ima l s : T he i rr e levance t oa r chaeo logy . I nR e sea rch P rob l em s i n Z ooa rchaeo logy , e d s . D . R . B ro thwe l l , K . D . T homa s & J .C lu t ton -B rock . L ondon : I n s t i tu t e o fA rchaeo logy , O c c a s iona lP ub l ica t ion N o . 3 ,p p . 1 8 . B i schof f , J .L . , J .F ranc i sco G a rc ia & L . G . S t rau s , 1 99 1 . U ran ium se r ie s i s och ron d a t ing a t E l C a s t i l lo c ave ( Can tab r ia , S pa in ) : T he " Acheu lean " / " Mou s te r ian " q ue s t i on . J ou rna lo fA rchaeo log ica lS c i ence 1 9 ,4 9 -62 .

B a i ley , G . & I .D av id son , 1 983 . S i te e xp lo i ta t ion t e r r i to r ie s a nd t opog raphy : Two c a se s t ud ie s f r om P a laeo l i th ic S pa in . J ourna lo fA rchaeo log ica lS c ience 1 0 , 8 7 115 . B a rand ia rän Mae s tu , I . , 1 981 . I ndu s t r ia ° sea . I nE l P a leo l i t ico S uper ior d el aC ueva d e lR a scaho ( San tand er ) , e d s . J .G onza lez E chega ray & I .B a rand ia rän Mae s tu . S an t a nde r : C en t ro d e I nve s t igac iön y Mu seo d e A l tam i ra , Monog raf ia sN o .3 ,p p . 9 7 164 . B a rand ia rän Mae s tu , I . ,1 987 . I ndu s t r ia o sea p a leo l i t ica d e l a C ueva d e l J uyo , e xcavac ione s d e 1 978 y 1 979 . I n E x c a vac ione s e n l a C ueva D e l J uyo , e d s . I .B a rand ia rän Mae s tu , L . G . F r eeman , J .G onza lez E chega ray & R . G . K le in . S an tande r : C en t ro d e I nve s t igac iön y Mu seo d e A l tam i ra , Monog raf ia s No . 1 4 ,p p . 1 6 1-194 B a rand ia rän , I . ,L .G . F reeman , J .G onza lez E chega ray & R . G . K le in ,e d s . , 1 987 .E xca vac ione se nl aC ue va D e lJ uyo . S an tande r : C en t ro d e I nve s t igac iön yMu seo d e A l tam i ra , Monog raf ia s No . 1 4 . B a ra sh , D . P . , 1 989 . Ma rmo t s : S oc ia lB eha v io ra nd E co logy . S t anf o rd : S t anf o rd U n ive r s i ty P re s s .

B i shop , I .R . & M . J .D e lany , 1 963 . L i f e h i s to r ie s o fs ma l l m amma l s i nt h e C hanne l I s land s i n 1 960 -61 . P roceed i ng so ft h eZ oo log ica lS oc ie ty o fL ond on 1 41 , 5 15 -526 . B leed , P . , 1 986 . T he o p t ima l d es ign o f h un t ing w eapon s : Ma in ta inab i l i ty o r r e l iab i l i ty . Ame r i can A n t iqu i ty 5 1 , 7 37 -747 . B lumen sch ine , R . J . , 1 986 . C a rca s s c on sump t ion s e quence s a nd t h e a r chaeo log ica l d i s t inc t ion o f s c aveng ing a nd h un t ing . J ou rna lo fH uman E vo lu t ion 1 5 ,6 39 -659 . B oon s t ra , R . &C . J .K r eb s , 1 978 . P i tf a l l t r app ing o fM i cro t u st own send i i . J ourna lo f Mamma log y5 9 , 1 36 148 . B o rde s , F . , 1 968 . T he O ld S tone Age . N ew Y o rk : M cG raw H i l l . B o rgh i , C .E . , S . M . G iannon i & J . P . Ma r t inez -R ica , 1 994 . H ab i ta t s e g rega t ion o ft h ree s ympa t r ic f o sso r ia l r oden t s i nt h e S pan i sh P y renee s . Z e i t schr i f if ür S äug e t ie rkund e 5 9 , 5 2 -57 .

B a r to lome i , G . ,J .C ha l ine , 0 . F e jf a r , D . J äno s sy ,M . J eanne t , W . v on K oen ig swa ld & K . K owa l sk i , 1 975 . P l iomy s l e nk i ( He l le r 1 930 ) ( Roden t ia , Mamma l ia ) e n E u rope . A c ta Z oo l ig ica C racov ien s ia 2 0 ,3 93 -467 .

B ou r l ie re , F . , 1 975 . Mamma l s , s ma l l a nd l a rge : T he e co log i c a l i mp l ica t ion s o fs i z e . I n Sma l l Mamma l s : T he i r P ro d uc t iv i ty a nd P opu la t ion D ynam ic s , e d s . F .B . G o l ley , K . P e t ru sew icz & L . R y szkowsk i . C amb r idge : C amb r idge

B eh re , K . -E . , 1 986 . A n th ropogen ic I nd ica to r s i nP o l len D ia g ram s . R o t te rdam :A .A . B a lkema . B eh ren smeye r , A . K . , 1 978 . T aphonom ic a nd e co log ic i n f o rma t ion f r om b one w ea the r ing . P a leob io logy 4 , 1 50 1 62 .

U n ive r s i ty P re s s ,p p . 1 8 . B oye r -K le in , A . 1 976 . A nä l i s i s p o l in ico d e l aC ueva d e T i to B u s t i l lo ( A s tu r ia s ) . I nE xcavac ione se nl aC ue va d e T i to B u s t i l lo ( A s tu r ia s ) . E xcavac ione s d e I9 75 , . e d s . A . Mou r e R oman i l lo & M . C ano H e r r e ra . O v iedo : I n s t i tu to d eE s tud io sA s tu r iano s ,p p . 2 03 -206

B eh ren smeye r , A . K . , 1 99 1 . T e r r e s t r ia l v e r teb ra te a ccumu la t i on s . I n T aphonomy : R e lea s ing t h e Da ta L ocked i nt h e 1 74

B oye r -K le in , A . , 1 981 . A nä l i s i s p o l in ico d e l a c ueva d e l R a sca f lo . I n E l P a leo l i t ico S upe r ior d e l a C ueva d e l R a scah 'o ( San tand er ) , e d s . I .B a rand ia rän Mae s tu & J . G onza lez E chega ray . C en t ro d e I nve s t igac iön y Mu seo d eA l tam i ra , Monog raf ia s No . 3 ,p p . 2 17 -220 .

B rune t -Lecom te , P .&M . D e l ibe s , 1 984 . A l imen tac iön d e l a l e chuza c omun T y to a lba e n l a C uenca d e l D ue ro , E spa f ia . D o l iana A c ta V e r tebra ta 1 ,2 13 -229 . B rune t -Lecom te , P . , T . L ode & P . P a i l ley ,

1 995 . Mo r -

p home t r ie c ompa ree d e l ap rem ie re mo la i re i n f e r ieu r e d u c ampagno l d e G e rbe , M icro tu s ( T er r ico la ) p y rena icu s g e rbe i( Roden t ia ,A rv ico l idae ) . Mamma l ia 5 9 ,2 49 -254 .

B oye r -K le in , A . , 1 984 . A na ly se s p o l l in ique s c an tab r ique s a u T a rd ig la c ia i r e . R evue d eP a ldob io log ie , V o l . s pec ia l , p p . 3 3 -39 .

B rune t -Lecom te , P . , P . T houy & J .C ha l ine , 1 994 . E tude B oye r -K le in , A . , 1 985 . A na ly se p o l l in ique d e l ag ro t te d ' E r -

c ompa rde d e p opu la t ion s a c tue l le s e tf o ss i le s d e M icro tu s ( T e r r ico la )p y rena icu s ( Roden t ia , A rv ico l idae ) . B u l le t in

r a l la . Mun ibe 3 7 ,4 5 -48 .

d el aS oc id td P rdh i s tor ique F ranva i se 19 , 3 7 -49 . B oye r -K le in , A . & A r . L e ro i -Gou rhan , 1 987 . A nä l i s i s p a l i n o log ico d e l aC ueva d e l J uyo . I n E xca vac ione s e n l a

B ucha lczyk , T . & J .R aczyn sk i , 1 96 1 . T ak sonom iczna w a r t o se n i ek to rych p om ia row c za szk i l c ra jowych p rzed s t aw ic ic l i r odza ju S orex L innaeu s , 1 758 iN eomy s K aup ,

C ueva D e lJ uyo , e d s . I .B a rand ia r än Mae s tu , L . G . F ree m an , J .G onza lez E chega ray & R . G . K le in . S an tande r : C en t ro d e I nves t igac iön y Mu seo d e A l tam i ra , Mono g raf ia sN o . 1 4 ,p p . 5-61

1 829 . A c ta T he r io log ica 5 , 15 124 . B uck land , W . , 1 823 . R e l iqu iae D i luv ianae .

B ra in , C . K . , 1 974 . T he u se o fm ic rof auna l r ema in s a sh ab i t a t i nd ica to r s i nt he N am ib . S ou th Af r ican A rchaeo log i c a lS oc ie ty G oodw in S e r ie s 2 ,5 5 -60 .

L ondon : J ohn

Mu r ray . B uck ley , J . , 1 977 . B a rn o w l p reda t ion o n t h e h a rve s t mouse . Mamma lR e v iew 7,17 -121 .

B ra in , C . K . , ( 1980 ) S ome c r i t e r ia f o r t h e r e cogn i t ion o f b one -co l lec t ing a genc ie s i nA f r ican C ave s . I nF o s s i l s i n t h e Mak ing : V er tebra te T aphonomy a nd P a leoeco logy , e d s . A . K . B eh ren smeye r & A . P . H i l l . C h icago : T he U n ive r s i ty o fC h icago P re s s , p p . 1 07 130 . B ra in , C . K . , 1 981 . T he H un te r s o r t h e Hun ted ? A n I n t ro d uc t ion t o Af r ican C a ve T aphonomy . C h icago : T he U n i v e r s i ty o fC h icago P re s s . B ra in , C .K . & V . B ra in , 1 977 . M ic rof auna l r ema ins f r om M i rab ib : S ome e v idence o fp a laeo -eco log ica l c hange s i n t h eN am ib . Madoqua 1 0 ,2 85 -293 . B ro thwe l l , D . R . & R . J one s , 1 978 . T he r e levance o fs ma l l mamma l s t ud ies t o a r chaeo logy . I n R e sea rch P rob lems i nZ ooa rchaeo logy , e d s . D . R . B ro thwe l l , K . D . T homa s & J .C lu t ton -B rock . L ondon : I n s t i tu te o f A rchaeo logy , O cca s iona lP ub l ica t ion N o . 3 ,p p .4 7 -57 B rown , J .C . &G . J . Tw igg , 1 969 . S t ud ie s o n t h e p e lv i s i n B r i t i sh Mu r idae a nd C r ice t idae ( Roden t ia ) . J ourna l o f

B uck ley , J .& J .G . G o ld sm i th , 1 975 . T he p r ey o ft h e b ar n o w l ( T y to a l ba a lba )i nE a s t No rf o lk . Mamma lR ev iew 5 , 1 3 16 . B uh le r , P . , 1 964 . Z u r G a t tung s - u nd A r tbe s t immung v on N eomy sS chode ln . Z e i t sch r i f if ü rS äuge t ie rkund e2 9 , 6 5 9 3 . B unn , D . S . , 1 972 . R egu la r d ay l igh t h un t ing b y b a rn o w l s . B r i t i sh B i rd s6 5 , 2 6 -30 . B unn , D . S . , A . B . Wa rbur ton & R . D . S .W i l son , 1 982 . T he B a rn Ow l . V e rm i l l ion , SD : B u teo B ook s . B u r ton , J .A . , e d . , 1 973 . Ow l s o ft h e W or ld . L ondon : P e te r L owe . B u tze r , K . W . , 1 980 . I nve s t igac iön p r e l i rn ina r d e l ag eo log ia d e l ac ueva d e E l P endo . I nE l Y ac im ien to d e l aC ueva d e " E l P end o" ( Exca vac ione s 1 953 -1957 ) , e d . J . Gon z ä lez E chega ray . Mad r id : B ib l io theca P raeh is to r ica H i s p ana ,V o l . 1 7 ,p p . 1 99 -213 .

Z oo logy ,L ondon 1 58 , 8 1132 . B rown , J .C . &G . J . Tw igg , 1 97 1 . Mamma l ian p rey o ft h e b a rn o w l ( T y to a lba ) o n S kome r I s land , P emb roke sh i re .

B u tze r , K . W . , 1 983 . G loba l s ea l e ve l s t ra t ig raphy :

A n a p -

p ra i sa l . Q ua te rna ry S c ience R e v iew s2 ,1 15

J ourna lo fZ oo logy ,L ond on 1 65 ,5 27 -530 . B u tze r , K . W . , 1 986 . P a leo l i th ic a dap ta t ion s a nd s e t t lemen t B rune t -Lecom te , P . , 1 990 . E vo lu t ion mo rpho log ique d e l a p rem ie re mo la i re i n f e r ieu re d e s c ampagno l s s ou te r ra in s d 'Eu rope

( Arv ico l idae ,

R oden t ia ) .

i nC an tab r ian S pa in . I n Ad vance s i n Wor ld A rchaeo log y , Vo l . 5 ,e d s . F . Wendo rf & A . E . C lo se . O r lando : A ca d em ic P r e s s ,p p . 2 01-252 .

Z e i t schr if t f ür

S äug e t ie r -kund e5 5 , 3 71-382 . B rune t -Lecom te , P . ,G . B roche t , J .C ha l ine & M . D e l ibe s , 1 987 . Mo rpho log ie d en ta i re c ompa ree d e P i tymy s l u s i t an i cu s e t P i tymy s d uod ec imco s ta tu s ( A rv ico l idae , R o d en t ia ) d an s l en o rd -oue s t d e l ' E spagne . Mamma l ia 5 1 , 1 45 158 .

CL I MAP , 1 976 . T he S u rf ace o ft h e I c e Age E a r th . S c ience 1 9 1 , 1311137 . C ab re ra , A . , 1 907 . T h ree n ew S pan i sh i n sec t ivo re s . A nna l s a nd Maga z ine o fN a tura lH i s tory S e r . 7 ,2 0 ,2 12 .

1 75

C ab re ra , A . , 1 908 . O n Mu sca rd in idae f r om t h e I b e r ian P en i n su la . A nna l s a nd Maga z ine o fN a tura l H i s tory S e r . 8 ,

t a nd e r , yn o t ic ia s obre u no n ue vo d e l a C a ve rna " E l P endo" . Mad r id : C om i s iön d e I n ve s t igac iöne s P a leon t o lög ica s yP reh i s tö r ica s , No . 1 .

1 ,1 88 . C ab re ra , A . , 1 9 14 . F auna l bd r ica : Mam if e ro s .

Mad r id :

Mu seo N ac iona ld eC ienc ia sN a tu ra le s .

C ha l ine , J . 1 972 . L e s R ongeur s d u P ld i s tocdne Moyen e t S updr ieu r d e F rance . P a r i s : C en t re Na t iona l d e l aR e c he rche S c ien t if ique .

C ab re ra V a lde s , V . , 1 984 . E l Y ac im ien to d e l aC ueva d e " E l C a s t i l lo" ( Puen te V i e sgo , S an tand e r ) .

Mad r id : B ib -

C ha l ine , J . , 1 974 . E squ is se d e l ' evo lu t ion mo rpho log ique ,

l i o theca P raeh i s to r ica H i span ica ,N o .2 2 . C ab re ra V a lde s , V . &M . G imenez L a R o sa , 1 981 . S ob r e l a i ndu s t r ia o sea d eA l tam i ra . I nE spac io , T i empo yF orma , S e r ie s 1 ,P reh i s to r ia , V o l . 4 . Mad r id : U n ive r s idad N a c i ona ld eE ducac iön aD is tanc ia ,p p . 9 3 -110 C amacho Muf ioz , I . , 1 975 . L a a l imen t ic iön d e T y to a l ba ( Scop . ) e n l aV ega d e G ranada . C uad erno s d e C ienc ia s B io log ica s , U n iver s idad d eG ranada 4 , 11-121 .

b i ome t r ique e t c h romosom ique d u g en re M icro tu s ( A rv ico l idae , R oden t ia ) d ans l eP l e i s tocene d e l ' hem i s phe re N o rd . B u l le t in d e l a S oc id td G do log ique d e F rance s e r . 7 ,1 6 ,4 40 -450 . C ha l ine , J . , 1 985 . H i s to i re d e l ' homme e t d e s c l ima t s a u Q ua terna i re . P a r i s : D o in E d i teu r s . C ha l ine , J .& J . -D . G raf , 1 988 . P hy logeny o ft h e A rv ico l idae ( Roden t ia ) : B iochem ica l a nd p a leon to log ica l e v idence . J ourna lo f Mamma logy 6 9 , 2 2 -33 .

C ampo s , F . , 1 978 . R eg imen a l imen t ic io d e T y to a l ba e n l a s P rovenc ia s d e S a lamanca y Z amo ra . 19 .

A rd eo la 2 4 , 1 05 -

C ha se , P .G . &H . L .D ibb le , 1 987 . M idd le P a leo l i th ic s ym b o l i sm : Ar ev iew o fc u r ren te v idence a nd i n te rp r e ta t ion s . J ou rna lo fA n th ropo log ica lA rchaeo logy 6 ,2 62 -296 .

C andau , P . & F . J .G onza lez M ine ro , 1 992 . R e la t ion sh ip b e tween v ege ta t ion a nd p o l len s p ec t rum i n S ou thwe s t S pa in . I s rae lJ ourna lo fB o tany 4 1 ,2 85 -295 . C a r ta i lhac , E . , 1 902 . L e s c ave rne s o rnee s d e d e s s in s . L a g ro t te d 'A l tam i ra , E spagne : ` Mea C u lpa d 'un s c ep t ique ' . L 'An thropo log ie 1 3 , 3 48 -354 . C a s ta i lo s , P . , 1 980 . L a mac rof auna d e l a C ueva d e L a P a loma ( P le i s toceno T e rm ina l d e A s tu r ia s ) . I nL a C ueva d e l aP a loma , e d s . M . H oyo s , M . I . Ma r t inez N ava r r e te , T . C hapa B r une t , P .C a s taf lo s &F .B . S anch iz . E xcava c i one sA rqueo lög ica se nE spaf ia N o . 16 ,p p . 6 5 -100 . C a s t ien , E . & J .G o sä lbez , 1 992a . D i s t r ibuc iön d e m ic ro mam if e ro s ( I n sec t ivo ra y R oden t ia ) e nN ava r ra . M i sce l l anea Z oo log ica 1 6 , 1 83 -195 . C a s t ien , E .&J .G o sä lbez , (1992b ) D i s t r ibuc iön g eog raf ica y h ab i ta t s o cupado s p o r G a lemy s p y rena icu s ( Geof f rey , 1 811) ( I n sec t ivo ra : T a lp idae ) e n l o s P i r ineo s O cc iden ta l e s . D ona r ia , A c ta V e r tebra ta 1 9 , 3 7 -44 . C a s t ien , E .&J .G o sä lbez , 1 995a . D ie to fS o rex c orona tu s i n t h ew e s te rn P y renee s . A c ta T he r io log ica 4 0 , 13 -121 .

C h i t ty , D . , 1 938 . A l a bo ra to ry s t udy o f p e l le t f o rma t ion i n t h e s hor t -ea r ed o w l ( As iof lammeu s ) . P roceed ing s o ft h e Z oo log ica lS oc ie ty o fL ondon S e r . A , 1 08 ,2 67 -287 . C la rk , G . A . , 1 971 . T he A s tur ian o f C an tabr ia : A R e e va lua t ion . U npub l i shed P h .D . D i s se r ta t ion , D epa r tmen t o fA n th ropo logy , U n ive r s i ty o fC h icago . C la rk , G . A . , 1 974 -75 . E xcava t ion s i nt h e L a te P l e i s tocene c ave s i te o fB a lmo r i , A s tu r ia s ( Spa in ) . Q ua te rna r ia 1 8 , 3 83 -426 . C la rk , G . A . , 1 983 . T he A s tur ian o fC an tabr ia : E a r l y H o lo c ene Hun te r -Ga there r s i nN or the rn S pa in . T u scan : U n i v e r s i ty o fA r izona P r e s s . An th ropo log ica l P ape r s o ft h e U n ive r s i ty o fA r izona ,N o . 4 1 . C la rk , G . A . & L . R . R icha rd s , 1 978 . L a te a nd p o s t P l e i s tocene i n du s t r ie s a nd f auna f r om t h e c ave s i t e o fL a R ie ra ( P rov ince o f A s tu r ia s , S pa in ) . I n V i ew s o f t h e P a s t , . e d . L . G . F reeman . T he H ague : Mou ton P ub l i sh e r s ,p p . 17 -152 C la rk , G . A . &L . G . S t rau s , 1 983 . L a t e P l e i s tocene h un te r g a the re r a dap ta t ion s i n C an tab r ian S pa in . I n H un te r G a therer E conomy i nP reh i s tor y , e d . G . B a i ley . C am -

C a s t ien , E . & J .G o sä lbez , 1 995b . R ep roduc t ive c yc le o f M icro tu s p y rena icu s D e S e ly s -Longchamp s , 1 847 ( Mamma l ia , R oden t ia ) i n t h e We s te rn P y renee s . s chr if tf ür S äuge t ie rkund e6 0 , 1 70 -175 .

C la rk , R . 1 981 . C o ld c l ima te f e a tu r e s o f t h e C o rd i l le ra C an tab r ica , n o r ther n S pa in . B iu le tyn P e tyg la c ja lny 2 8 ,

C a s t ro B e rna rdez , D . , 1 986 . E s tud io d el am icrof auna P l e i s t o cena S upe r ior y ac im ien to s d e A l tam i ra , E l J uyo y C ue to d e l a M ina ( Reg ion C an tabr ica ) . M .S . T he s i s ,U n ive r s idad d e S an t iago .

b r idge : C amb r idge U n ive r s i ty P re s s ,p p . 1 31-148 .

Z e i t -

5 13 . C la rk , R . J . , 1 972 . P e l le t s o ft h e S ho r t -ea r ed o w l a nd Ma r sh

U npub l ished

h awk c ompa r ed . 9 62 -964 .

C end r e ro , 0 . , 1 9 15 . R e sumen d e l o s b a s tone s p e rf o rado s ( ba s tone s d e m ando ) h a l lado s e n l ap rov inc ia d e S an 1 76

J ou rna l o f W i ld l if e Managemen t 3 6 ,

C lu t ton -B ro ck , T . H . , F . E . G u ine s s & S . D . A lban , 1 982 . R ed D ee r :B ehav ior a nd E co logy o fT wo S exe s . C h icago : T he U n ive r s i ty o fC h icago P re s s .

C umm ing s , J .H . , G . E . D uke & A . A . J ege r s , 1 976 . C o r ro -

C ohen , M . N . , 1 977 . T he F ood C r i s i s i nP reh i s tory .

D a L uz Madu r e i ra , M . 1 983 . O n t h e i d en t if i ca t ion o f We s t -

N ew

s i on o fb one b y s o lu t ion s s imu la t ing r ap to r g a s t r ic j u ice . R ap tor R e sea rch 1 0 , 5 5 -57 .

E u ropean M ic ro tu s o f t h e s ub -gene ra M icro tu s a nd

Haven :Y a le U n ive r s i ty P re s s .

C h ionomy s( Mamma l ia , R oden t ia ) ap r inc ipa l c omponen t a na ly s i s . R ev i s ta d eB io log ia 1 2 , 6 15 -632 .

C onkey , M . W . , 1 978 . A n A na ly s i s o f D e s ign S t ruc tu re : V a r iab i l i ty a mong Magda len ian E ng ra ved B one s f rom Nor th C oa s ta l S pa in . U npub l i shed P h .D . D i s ser ta t ion , D epa r tmen to f An th ropo logy ,U n ive r s i ty o fC h icago .

D av id son , I . ,1 976a . L e s Ma lae t e s a nd Monduve r : T he e con omy o f ah uman g roup i np r eh i s to r ic S pa in . I nP rob l em s i n E conom ic a nd S oc ia l A rchaeo logy , e d s . G . d e G . S i evek ing , I .H . L ongwo r th & K . E . W i l son . L ondon :

C onkey , M . W . , 1 980 . T he i d en t if i ca t ion o f p reh i s to r ic h un te r -ga the re r a gg rega t ion s i t e s : T he c a se o fA l tam i ra .

Duckwor th ,p p . 4 83 -499 .

C u rren tA n th ropo logy 2 1 ,6 09 -630 . C onne l l , J .H . , 1 978 . D ive r s i ty i nt r op ica l r a in f o re s t s a nd

D av id son , I . , 1 976b . S ea sona l i ty i nS pa in . Z ephy ru s 2 6 -27 , 1 75 -182 .

c o ra lr e ef s . S c ience 1 99 , 1 302 1310 . C oon , C . S . , 1 971 . T he Hun t ing P eop le s . N ew Y o rk : N ick L yon s .

D av id son , I . , 1 983 . S i te v a r iab i l i ty a nd p reh i s to r ic e conomy i nL evan te . I nH un te r -Ga there r E conomy i nP reh i s to ry , e d . G . B a i ley . C amb r idge : C amb r idge Un ive r s i ty P re s s , p p . 7 9 -95 .

C o rchön , M . S o ledad , 1 981 . C ueva d e l a s C a lda s .

Mad r id :

E xcava c ione sA rqueo lög ica se nE spaha No . 15 . C o rchön , M . S o ledad , 1 986 . E l A r te Mueb le P a leo l i t ico C an täb r i co . S an tande r : C en t ro d eI nve s t igac iön yMu seo d eA l tam i ra , Monog raf ia s No . 1 6 . C o rbe t , G . B . , 1 966 . T he T e r re s t r ia l Mamma l s o f W e s te rn E u rope . L ondon : G .T . F ou l i s &C o . C o rbe t , G . B . , 1 978 . T he Mamma l s o ft h eP a l ea rc t ic R eg ion : AT a xonom ic R ev iew . L ondon : C o rne l l U n ive r s i ty P re s s .

D av i s , D . H . S . , 1 958 . N o te s o n s ome s ma l l mamma l s i nt h e K a laha r i G em sbok N a t iona l P a rk , w i th s pec ia l r e f e rence t ot ho se p r eyed u pon b yb a rn o w l s . K oedoe 1 ,1 84 -188 . D av i s , D . H . S . , 1 959 . T he b a rn o w l ' sc on t r ibu t ion t oe co logy a nd p a laeoeco logy . I n P roceed ing s o f t h e F i r s t P an Af r ican O rn i tho log ica l C onf e rence , e d s . M . K . R owan . O s t r ich , S upp lemen t No . 3 ,p p . 1 43 -153 . D av i s , M . B . , 1 968 . P o l len g ra in s i ns ake s ed imen t s : R ede p o s i t ion c au sed b y s ea sona l w a te r c i rcu la t ion . S c ience 1 62 , 7 96 -799 .

C o rbe t , G . B . , 1 984 . T he Mamma l so ft h eP a lea rc t ic R eg ion : A T axonom ic R ev iew ( Supp lemen t ) . D o rche s te r : H en ry L ing .

D av i s , S . J .M . , 1 987 . T he A rchaeo logy o fA n ima l s . H aven : Y a le U n ive r s i ty P re s s .

C o rbe t , G . B . , J .C umm in s , S .R . H edge s & W . K rzanow sk i , 1 970 . T he t a xonom ic s t a tu s o fB r i t i sh Wa te rv o le s , g enu s A rv ico la . J ourna lo fZ oo logy , L ondon 1 6 1 , 3 01-316 .

D ay , M . G . , 1 966 . I d en t if ica t ion o fh a i r a nd f e a the r r ema in s i nt h e g u t a nd f a ece s o f s t oa t s a nd w ea se l s . J ourna l o f Z oo logy 1 48 ,2 0 1-217 .

C o rbe t , G . B . & S . H a r r i s , e d s . , 1 991 . T he Hand book o f B r i t i sh Mamma l s . O xf o rd : B lackwe l l S c ien t if ic P ub l ica t i on s .

D ayan , T . & E . T che rnov , 1 988 . O n t h e f i r s t o ccu r rence o f t h e c ommon w ea se l (Mu s te la n i va l i s ) i nt h e f o s s i l r e co rd

C o rbe t , G . B . & D . O venden , 1 982 . Manua l d e l o s Mam i f e ro s d e E spar ta y d e E u ropa . B a rce lona : E d ic io -

d e P az , 0 . &J .B enza l , 1 989 . C lave p a ra l ai d en t i f icac iön d e

o fI s rae l . Mamma l ia 5 2 , 1 65 168 .

l o s mu rc ie lago s d e l aP en in su la I b e r ica ( Mamma l ia , C h i -

n e s Omega . C rowc rof t ,P .1 966 .M ice A l lO ve r . L ondon : G .T . F ou l i s .

N ew

r op te ra ) . M i sce l lan ia Z oo log i ca 1 3 , 1 53-176 . D ean , W . R .J , . 1 973 . A ge d i s t r ibu t ion o fP raomy sn a ta len s i s p rey i nT y to a l ba p e l le t s . Z oo log ica Af r icana 8 ,1 40 .

C rowe , W .T . , 1 987 . T ecn ica sd er e cupe rac iön i n teg ra ld e l o s d a to s o b ten ido s e n l o s s ed imen to s d e y ac im ien to s p re -

D e lany , M . J .& P . E . D av i s , 1 961 . O b se rva t ion s o n t h e

h i s tö r ico s . I nE xca vac ione se ll aC ue va d e lJ uyo , . e d s . I . B a rand ia rän ,L .G . F reeman , J .G onza lez E chega ray & R .

e co logy a nd l i f e h i s to ry o ft h e F a i r I s le f i e ld -mou se A po d emu s s y lva t icu sf r ida r ien s i s ( K innea r ) . P roceed ing s o f

G . K le in . C en t ro d e I nve s t iga c iön yMu seo d e A l tam i ra , Monog ra f ia s No . 1 4 , p p . 6 3 -74 .

t h eZ oo log i ca lS oc ie ty o fL ond on 1 36 ,4 39 -452 . D e la t t re , P . , 1 983 . D en s i ty o fw ea se l (M u s te la n i va l i s L . ) a nd s t oa t (M u s t e la n i va l i s L . ) i nr e la t ion t ow a te r v o le a bun d ance . A c ta Z oo log ica F enn i ca 1 74 , 2 21-222 . 1 77

D e l ibe s , M . , 1 983 . I n te r spec if i c c ompe t i t ion a nd t h e h ab i ta t o f t h e s t one m ar ten Ma r te s f o ina ( E rx leben 1 777 ) i n E u rope . A c ta Z oo log ica F enn i ca 1 74 ,2 29 -231 .

D uke , G . E . , 0 . A . E van son , &A . A . J e ger s , 1 976 . Mea l t o p e l le t i n t erva l s i n 1 4 s pec ie s o f c ap t ive r ap to r s . C om p a ra t ive B iochem i s t ry a nd P hy s io logy 5 3A , 1 6 .

D e l ibes , M . , P . B rune t -Lecom t e & M . Manez , 1 984 . D a to s

D uke , G . E . , A . A . J ege r s , G . L of f& 0 . A . E van son , 1 975 . G a s t r ic d i ge s t ion i n s ome r ap to r s . C ompa ra t ive B io -

s ob re l aa l imen tac iön d e l al e chuza c omun ( T y to a l ba ) , e l

c hem i s t ry a nd P hy s io logy 5 0A , 6 49 -656 .

b uho c h ico ( As io o tu s ) ye l mochue lo ( A thene n oc tua )e n u na m i sma l o ca l idad d e C a s t i l la l aV ie ja . A rd eo la 3 0 ,

D uke , G . E . &D . D . R hoade s , 1 977 . F a c to r s a f f ec t ing m ea l

5 7 -63 . D e l ibe s d e C a s t ro , J . , 1 985 . D i s t r ibu t ion a nd a bundance o f s ma l l mamma l s i n ag rad ien to fa l t i tude . A c ta Z oo log ica F enn ica 1 73 , 5 3 -56 .

t o p e l le t i n terva l s i n G rea t h or ned o w l s . C ompa ra t ive B iochem i s tr ya nd P hy s io logy 5 6A ,2 83 -286 . D unm i re , W . W . , 1 955 . S ex d imo rph i sm i nt h e p e lv i s o fr o d en t s . J ou rna lo f Mamma logy 3 6 , 3 56 -361 .

d e Mor t i l le t , G . 1 883 . L e P rjh i s to r ique .

P a r i s : R e inwa ld .

B ib l io theque d e sS c ience sC on tempo ra ine , No . 8 . D eny s , C . &M . Mahboub i . 1 992 . A l te ra t ion s s t ruc tu ra le s e t c h im ique s d e s e l emen t s s que le t t ique s d e p e lo te s d e r e g u rg i ta t ion sd ' un r apace d i u rne . B u l le t in d u Mu seum N a t i ona l d 'H i s to i re N a ture l le 4 t h s e r ie s , s e c t ion A , n o . 1 , 1 4 , 2 29 -249 . D er t ing , T . L . &J .A . C ranf o rd , 1 989 . P hy s ica l a nd b ehav i o ra l c o r r e la te s o f p rey v u lne rab i l i ty t o b a rn o w l ( T y to a lba ) p r eda t ion . Amer ican M id land N a tura l i s t 1 2 1 , 1 2 0 . D ice , L .R . , 1 945 . M in imum i n ten s i t ie so fi lum ina t ion u nde r w h ich o w l sc an f i nd d ead p r ey b y s i gh t . Amer ican N a tu r a l i s t7 9 ,3 85 -416 . D ickman , C . R . , M . P redavec & A . J .L yman , 1 99 1 . D if f e r e n t ia l p r eda t ion o f s i ze a nd s e x c l a s se s o f m ice b y t h e b a rn Ow l , T y to a lba . O iko s6 2 , 6 7 -76 . D ien ske , H . , 1 969 . No te s o n d i f f e r ence s b e tween s ome e x t er na l a nd s ku l l c ha rac te r i s t ic s o f M icro tu s a rva l i s ( Pa l la s , 1 778 ) a nd o fM ic ro tu s a g re s t i s ( L innaeu s , f r om t h eN e the r land s . Z oo log i sche Med ed e l ingen 4 4 , 8 3 108 . D ipu tac idn R eg iona l d e C an tab r ia , 1 994 . D i spo s ic ione s g en e r a le s . No .2 60 , D ecembe r3 0 . C on se je r ia d e G anade r ia , Ag r icu l tu ra yP e sca , S an tande r , S pa in . D od son , P . , 1 973 . T he s i gn if icance o f s ma l l b one s i n p a l e oeco log ica l i n te rp re ta t ion . C on t r ibu t ion s t o G eo logy ,

D unw idd ie , P .W . , 1 987 . Mac rof o s s i l a nd p o l len r e p r e sen ta t i on o f c on if e rou s t r ee s i n mode rn s ed imen t s f r om Wa sh ing ton . E co logy 6 8 , 1 11 . D up re , M . , 1 984 . P a l ino log ia d e l o s n i ve le s V I I aI d e l a c ueva d e E ka in . I n E l Y ac im ien to P reh i s tör ico d e l a C ueva d e E ka in ( Deva , G u ipü zcoa ) , e ds . J .A l tuna & J . M . Me r ino . S an S eba s t iän : S oc iedad d e E s tud io s V a sco s S e r ie B 1 ,p p . 6 1-63 . D up re , M . , 1 990 . A nä l i s i s p o l in ico d e l ac ueva d e Ama lda . I n L a C ueva d e Ama lda ( Z e s toa , P a i s V a sco ) , . e d s . J . A l tuna , A . B a ldeön & K . Ma r iezku r r ena . S an S eba s t ian : S oc iedad d eE s tud io sV a sco s , p p . 4 9 -51 E a s tham , A . , 1 995 . L a m ic rof aune . I nL e s D e rn ie r s C ha s s eu r s d e R enne s d u Mond e P y rdneen . L 'Abr i D uf au re : U n G i semen t T a rd ig lac ia re e n G a scogne , e d . L . G . S t rau s . P a r i s : S oc ie te P reh i s to r ique F rança i se , Memo i re 2 2 , p p . 2 35 -245 . E f remov , J .A . , 1 940 . T aphonomy : N ew b ranch o fp a leon t o logy . P an -Ame r ican G eo log i s t7 4 , 8 1 -93 . E ina r sen , A . S . , 1 956 . D e te rm ina t ion o fs ome p reda to r s pe c i e s b y f i e ld s i gn s . O regon S ta te Monog raph s S tud ie s i n Z oo lo gy 1 0 , 1 34 . E l ia s , S .A . , 1 987 . P a leoenv i ronmen ta l s i gn if icance o fL a te Q ua te rna ry i n sec t f o s s i l s f r om p ack ra tm idden s i nS ou th C en t ra l N ew Mex ico . T he S ou thw e s t e rn Na tura l i s t 3 2 , 3 83 -390 .

U n ive r s i ty o fW yom ing 1 2 , 1 5 19 . D od son , P . & D . Wex la r , 1 979 . T aphonom ic i n ve s t iga t ion s o fo w lp e l le t s . P a leob io logy 5 ,2 75 -284 .

E ppe r son , C . , 1 976 . F ood r ema in s f r om ab a rn o w l n e s t i n N eb ra ska . N ebra ska B i rd R ev iew 3 4 ,5 4 -57 . E r l inge , S . , 1 987 . Why d o E u ropean s t oa t s Mus te la e rm inea

D onazä r , J .A . &0 . C eba l lo s , 1 989 . S e lec t ive p reda t ion b y e ag le o w l s B ubo b ubo o n r a bb i t s O ryc to lagu s c un icu lu s : Age a nd s ex p ref e rence s . O rn i s S cand inav ica 2 0 , 17 1 22 .

n o t f o l low B e rgmann ' s r u le? Ho /a r c t i c E co logy 1 0 , 3 3 3 9 . E van s , F . C . & J .T . Em len , J r . , 1 947 . E co log ica l n o t e s o n t h ep rey s e lec ted b y ab a rn o w l . C ondo r4 9 , 3 -9 .

D onazä r , J .A . & F . H i ra ldo , M . D e l ibe s & R . R . E s t r e l la , 1 989 . C ompa ra t ive f ood h ab i t s o f t h e e ag le o w l B ubo b ubo a nd t h e g r ea t h or ned o w l B ubo v i rg in ianu s i ns i x P a lea rc t ic a nd N ea rc t ic b iome s . O rn i s S cand ina v ica 2 0 , 2 98 -306 .

F a , J .E . , E . S haw & A . S an tana , 1 992 . H ab i ta t a s soc ia t ion s o f s ma l l m amma l s i n as ou the rn S pan ish f i r f o r e s t . Mamma l ia 5 6 ,4 78 -48 1 .

1 78

F aeg r i , K . & J .I v e r sen , 1 989 . T ex tbook o f Mod e rn P o l len A na ly s i s( 4 th e d . ) . C h iche s te r : J ohn W i ley & S on s . F a l l , P . L . , 1 987 . P o l len t a phonomy i n ac anyon s t ream . Qua te rna ry R e search 2 8 ,3 93 -406 .

F reeman , L . G . , e d . , 1 987 . A l tam i ra R ev i s i ted . I n s t i tu te f o rP reh i s to r ic S tud ie s .

C h icago :

F r eeman , L . G . , 1 988 . T he s t ra t ig raph ic s equence a t A l tam i ra , 1 980 -1981 . E spac io , T i empo y F o rma , S e r ie s 1 ,P r eh i s to r ia , V o l . 1 , . Mad r id : U n ive r s idad N ac iona l d e E duca c iön aD i s tanc ia ,p p . 1 49 163

F a rne r , D . S . , 1 960 . D ige s t ion a nd t h e d ige s t ive s y s tem . I n B io logy a nd C ompara t ive P hy s io logy o fB i rd s , Vo l . 1 , e d s . A . J . Ma r sha l l .

N ew Y o rk : A cadem ic P re s s , p p .

4 11-467 .

F reeman ,L .G . , 1 994 . P a leo l i th ic p o lygon s : V o rono it e s se rae a nd s e t t lemen t h i e ra rch ies i nC an tab r ian S pa in . E spac io , T i empo y F orma , S e r ie s 1 ,P r eh i s to r ia , V o l . 7 , Mad r id : Un ive r s idad N ac iona l d e E ducac iön aD i s tanc ia , p p . 4 3 -

F a r rand , W .R . , 1 975 .A na ly s i so ft h e A b r iP a taud s ed imen t s . I nE xca va t ion o ft h eA br iP a taud , L e sE y z ie s ( Do rdogne ) , e d . H . L . Mov iu s , J r .

6 1 .

C amb r idge : H a rva rd U n ive r s i ty

P re ss ,p p .2 7 -68 .

F reeman , L . G . & J . Gonza lez E chega ray , 1 981 . E l J uyo : A 1 4 ,000 -yea r -o ld s anc tua ry f r om n o r the rn S pa in . H i s tory

F a s t , S . J .& W . A . H a r r i son , 1 975 . P rey p r ef e rence a nd h un t ing h ab i ta t s e lec t ion i n t h e b a rn o w l . Ame r i can M id land N a tura l i s t9 6 , 5 03 -507 .

o fR e l ig ion s2 1 , 1 19 . F reeman , L . G . & J . G onza lez E chega ray , 1 982 . Mag d a len ian mob i le a r t f r om E l J uyo ( Can tab r ia ) . A r s P rae h i s to r ica 1 ,1 61167 .

F edyk , A . &A . L . R up rech t , 1 971 . T axonom ic v a lue o f MI mea su remen t s i nM ic ro tu s a g re s t i s ( L innaeu s , 1 761) a nd M icro tu s a r va l i s ( Pa l la s , 1 779 ) . A c ta T he r io log ica 1 6 , 4 08 -4 12 . F e l ten , H . , A . H e lf r ich t&G . S to r ch , 1 973 . D ie B e s t immung d e re u ropa i schen F l ede rmau se n ach d e rd i s ta len E p iphy se d e sH ume ru s . S enckenbe rg iana B io log ica 5 4 ,2 9 1-297 . F er nändez -T re sgue r re s , J . , 1 981 . E lA z i l ien se e n l a s P rov in c i a s d e A s tur ia s yS an tand e r . S an tande r : C en t ro d e I n v e s t igac iön yMu seo d eA l tam i ra , Monog raf ia sN o . 2 .

F r eeman , L . G . & J .G onzä lez E chega ray , 1 995 . T he Mag d a len ian s i te o f E l J uyo ( Can tab r ia , S pa in ) : A r t i s t ic d ocumen t s i nc on tex t . B o l le t t ino d e l C en t ro C amuno d e S tud iP re i s tor ic i2 7 , 2 5 -42 . F reeman , L .G . , J .G onza lez E chega ray , R . G . K le in & W . T . C rowe , 1 988 . D imen s ions o fr e sea rch a tE l J uyo , a n e a r l i e r Magda len ian s i te i n C an tab r ian S pa in . I n U pper P le i s tocene P reh i s tory o f W e s te rn E ura s ia , e d s . H . L . D ibb le & A . Mon te t - Wh i te . P h i lade lph ia : U n ive r s i ty o f P enn sy lvan ia Mu seum , p p . 1 39 .

F i she r , D . C . , 1 981a . C rocod i l ian s c a to logy , m ic rover teb ra te c oncen t ra t ion s , a nd e name l le s s t e e th . P a leob io logy 7 , 2 62 -275 . F i she r , D . C . , 1 98 1 b . Mode o fp re se rva t ion o ft h e S ho tgun L oca l F auna ( Pa leocene , Wyom ing ) a nd i t s i mp l ica t ion f o r t h e t a phonomy o f am ic rove r teb ra te c oncen t ra t ion . C on t r ibu t ion s f rom t h e Mu seum o f P a leon to logy , t h e U n ive r s i ty o fM ich igan 2 5 ,2 47 -257 . F i she r , D . C . , 1 981c . T aphonom ic i n te rp re ta t ion o fe name l l e s s t e e th i nt h e S ho tgun L oca l F auna ( Pa leocene , Wyo m ing ) . C on t r ibu t ion sf rom t h e Mu seum o fP a leon to logy , T he U n ive r s i ty o fM ich igan 2 5 ,2 59 -275 .

F reeman , L . G . , J .G onza lez E chega ray , J .P ok ine s , H . S t e t t i e r&M . K rupa , 1 998 . T am i sage u l t ra f m e tr e cupe ra t ion d e l ' ou t i l lage : O b se rva t ion s / t an sie s aE l J uyo ( E spagne C an tab r ique ) . L' An thropo log ie 1 02 , 3 5 -44 . F r ey , H . &W . Wa l te r , 1 986 . Z u rE rnährung d e s U hu s , B ubo b ubo ( L innaeu s 1 758 ) , A ye s , a n e i nem a l p inen B ru tp la tz i nd en H ohen T auer n( Sa lzbe rg , Ö s te r re ich ) . A nna len d i e Na turh i s to r i schen Mu seums e n W i en 8 8 /89 , 9 1-99 . F uen te s V ida r te , C . , 1 980 . E s tud io d e l aF auna d e E l P endo . I n E l Y ac im ien to d e l a C ueva d e " E l P endo" ( Exca vac ione s 1 953 -1957 ) , e d . J .G onza lez E chega ray . Mad r id : B ib l io theca P raeh i s to r ica H i span ica , No . 1 7 , p p . 2 15 -237 .

F reeman , L . G . , 1 973 . T he s i gn if icance o fm amma l ian f a u n a s f r om p a leo l i th ic o ccupa t ion s i n C an tab r ian S pa in . Ame r i can A n t iqu i ty 3 8 , 3 -44 .

F u lk , G . K . , 1 976 . Ow l p reda t ion a nd r oden t mo r ta l i ty : A c a se s t udy . Mamma l ia 4 0 ,4 23 -427 .

F reeman , L .G . , 1 975 . B y t h e i r w o rk s y ou s ha l l k now t h em : C u l tu ra l d eve lopmen t s i nt h e P a leo l i th ic . I n H om in i sa t i on u nd V e rha l ten , e d s . G . K ur th & I .E ib l -E ibe sf e ld t .

G a rge t t , R . H . , 1 989 . G rave s ho r tcom ing s : T he e v idence f o r N eander tha lb u r ia l . C ur ren tA n th ropo logy 3 0 , 1 57 -190 .

S tu t tga r t :G .F i sche r ,p p .2 34 -26 1 . F reeman , L .G . , 1 981 . T he F a t o ft h e l a nd : N o t e s o n P a leo l i th ic d i e t i nI b e r ia . I n Omn ivorou s P r ima te s , e d s . R . S . 0 . H a rd ing & G . T e lek i . N ew Y o rk : C o lumb ia U n ive r -

G a rzon -Heyd t , J .& J .C a s t rov ie jo , 1 970 . No ta s p re l im ina re s s ob re l ad i s t r ibuc iön d e a l guno s m ic ro -mam if e ro s e n e l n o r te d e E spa f ia . S äuge t ierkund l iche M i t te lungen 1 9 , 2 17 -222 .

s i ty P r e s s ,p p . 1 04 -165 .

1 79

G i l l ia rd , E . T . , 1 958 . T y to a lba , P a r t I . N a tu ra lH i s to ry 6 7 , 2 38 -245 . G lad ing , B . , D . F .T i l lo t son & D . M . S e l leck , 1 943 . R ap to r p e l le t s a si n d ica to r s o ff ood h ab i t s . C a l if orn ia F i sh a nd G ame 2 9 ,9 2 121 .

G onza lez E chega ray , J . & L . G . F reeman , 1 998 . L e P a läo l i th ique I nf ä r ieur e t Moyen d 'E spagne . P reh i s to i r e d 'Eu rope 6 .G renob le : E d i t ion sJ e rome M i l lon . G onza lez E chega ray , J . , M . A . G a r c ia G u inea , A . B eg ine s R am i rez & B . Mada r iaga d e l aC ampa , e d s . , 1 963 . C ueva d el aC ho ra . E xcavac ione sA rqueo lög ica s e nE spaha No .

G lue , D . E . , 1 967 . P rey t a ken b y t h e b a rn o w l i nE ng land a nd Wa le s . B i rd S tud y 1 4 , 1 69 183 .

2 6 . G onza lez E chega ray , J . , M . A . G a rc ia G u inea & A . B eg ine s

G lue , D . E . , 1 970 . A v ian p reda to r p e l le t a na ly s i s a nd t h e

R am i rez , e d s . , 1 966 . C ueva d e lO t e ro . E xcavac ione s A r -

mamma log i s t . Mamma lR ev iew 1 ,5 3 -62 .

q ueo lög ica se nE spaha No . 5 3 .

G lue , D . E . , 1 974 . F ood o ft h e b a rn o w l i nB r i ta in a nd I r e l a nd . B i rd S tud y2 1 ,2 00 -210 .

G onzä lez Mo ra le s , M . R . , 1 974 . L a C ue va d e C o l lub i l . L i c enc ia tu ra t h e s i s , D epa r tam ien to d e P r eh i s to r ia y A r q ueo log ia ,U n ive r s idad d eO v iedo .

G lue , D . E . , 1 975 . H a rve s t m ice a s b a rn o w l p r ey i n t h e B r i t i sh I s le s . Mamma lR ev iew 5 ,9 12 .

G onza lez Mo ra le s , M . R . , 1 986 . L a R ie ra b one a nd a n t le r a r t if ac ta s semb lage s . I nL aR iera C a ve , e d s . L .G . S t rau s

G onza lez E chega ray , J . , 1 953 . L e s o euv re s d 'a r td e l aG ro t te d e l a s C h imenea s . B u l le t in d e l aS oc iä tä P räh i s to r ique d el ' Ar iäge 8 ,7 5 -77 .

& G eof f r ey A . C la rk . A r izona S t a t e U n ive r s i ty A n th ro p o log ica lR e sea r ch P ape r s No . 3 6 ,p p .2 09 -218 . G onza lez S a inz , C . , 1 988 . L e f a i t a r t i s t ique a l a f i n d u

G onza lez E chega ray , J . , 1 960 . E l Magda len ien se I I d e l a c o s ta C an täb r ica . B o le t in d e S em ina r io d e E s tud io s d e A r te y A rqueo log ia , U n ive r s idad d e V a l ledo l id 2 6 , 6 9 1 00 . G onza lez E chega ray , J . , 1 974 . C ueva d e l a s C h imenea s . Mad r id : E xcavac ione sA rqueo lög ica se nE spaha No 2 1 . G onza lez E chega ray , J . , e d . , 1 980 . E l Y ac im ien to d e l a C ueva d e " E lP endo" ( Exca vac ione s 1 953 -57 ) . Mad r id : B ib l io theca P ra eh i s to r ica H i spana ,N o . 1 7 .

P a leo l i th ique : Q ue lque sr e f lex ion s . B u l l e t in d el aS oc iä tä P räh i s tor ique A r iäge -Py ränäe s4 3 , 3 5 -62 . G oodman , S . M . , 1 990 . T he f ood h ab i t s o f t h e e ag le o w l ( Bubo b ubo a sca laphu s ) i n K ha rga O a s i s , E gyp t ian We s te rn D e ser t . J ourna l o fA r id E nv i ronmen t s 1 8 , 2 17 2 20 . G oodw in , R . G . , 1 988 . P o l len t a phonomy i nH o locene g l a c i o lacu s t r ine s ed imen t s , G lac ie r B ay , A la ska : Ac au t ion a r yn o te . P a la io s3 ,6 06 -611 .

G onzä lez E chega ray , J . , 1 987 . I n t roducc iön g eog raf ica e h i s tö r ica . I nE xca vac ione s e n l aC ueva D e lJ uyo , e d s . I . B a rand ia rän Mae s tu , L .G . F reeman , J .G onza lez E chega r ay & R . G . K le in . S an tande r : C en t ro d e I nve s t igac iön y Mu seo d eA l tam i ra , Monog raf ia s No . 1 4 , p p . 7 14 .

G o szczyn sk i , J . , 1 974 . S t ud ie s o n t h e f o od o f f oxe s . T her io log ica 1 9 , 1 18 .

G onza lez

G o szczynsk i , J . , 1 977 . C onnec t ion s b e tween p r eda to ry b i rd s a nd mamma l s a nd t h e i rp rey . A c ta T her io log ica 2 2 , 3 99 4 30 .

E chega ray ,

J . ,

1 988 .

E l

Magda len ien se

d e

A l tam i ra . E spac io , T i empo y F orma , S e r ie s 1 ,P reh i s to r i a , V o l . 1 , Mad r id : U n ive r s idad N ac iona l d e E ducac iön a D i s tanc ia ,p p . 1 65 175 . G onzä lez E chega ray , J .& I .B a rand ia rän Mae s tu , e d s , . 1 981 . E l P a l eo l i t i co S upe r ior d e

l a C ueva

d e l R a scaho

G o szczyn sk i , J . , 1 976 . C ompo s i t ion o ft h e f ood o f ma r ten s . A c ta T he r io log ica 2 1 , 5 27 -534 .

G raham , I .& A . S aunde r s , 1 978 . A m u l t iva r ia te s t a t i s t ica l a na ly s i s o fs ma l l mamma l b one s . I nR e sea rch P rob lem s i nZ ooa rchaeo logy , e d s . D . R . B ro thwe l l , K . D . T homa s

( San tand e r ) . S an tande r : C en t ro d e I nve s t igac iön y Mu seo d eA l tam i ra , Monog raf ia s No . 3 . G onza lez E chega ray , J . & L . G . F reeman , 1 971 . C ueva Mor in : E xca vac ione s 1 966 -1968 . d e l a sC ueva sP reh i s tö r ica s .

& J .C lu t ton -B rock .

G raham , R . W . , 1 979 . P a leoc l ima te s a nd L a t e P l e i s tocene f a una l p rov ince s i n No r th Ame r ica . I n P re -L lano C u l t u re s o ft h e Ame r ica s : P rob lems a nd P a radoxe s , e d s . R . L . H umph rey & D . S t anf o rd . Wa sh ing ton , D .C . : Wa sh -

G onza lez E chega ray , J . & L . G . F reeman , 1 973 . C ue va

G onza lez E chega ray , J .& L . G . F reeman , 1 978 . V ida y

L ondon : I n s t i tu te o f A rchaeo logy ,

O cca s iona lP ub l ica t ion N o . 3 ,p p . 5 9 -67 .

S an tande r : P a t rona to

Mor in : E xca va c ione s 1 969 . S an tande r : P a t rona to d e l a s C ueva sP reh i s tö r ica s .

A c ta

i n g ton A n th ropo log ica l S oc ie ty ,p p .4 9 -69 . G ray , J .A . , 1 989 . E f f ec t s o fe nv i ronmen ta l s t re s s o n s pec ie s r i ch a s semb lage s . B io log ica lJ ou rna l o ft h eL innean S o c i e ty 3 7 , 1 9 -32 .

Mue r te e n C ueva Mor in . S an tande r : I n s t i tuc iön C u l tu ra l d eC an tab r ia . 1 80

G ray son , D . K . , 1 981 . Ac r i t ica l v i ew o ft h e u se o fa r chaeo l o g i ca l v e r teb ra tes i np a leoenv i ronmen ta l r e con s t ruc t ion . J ou rna lo fE thnob io log y 1 ,2 8 -38 .

H e r re ra , C . M . , 1 973 . R eg imen a l imen t ic io d e T y to a l ba e n E spa i la s udocc iden ta l . A rd eo la 1 9 , 3 59 -394 . H e r re ra , C . M . &F .M . J ak s ic , 1 980 )F eed ing e co logy o ft h e

G ray son , D . K . , 1 983 . T he E s tab l i shmen t o fH uman A n t iq u i ty . N ew Y o rk : A cadem ic P re s s .

b ar n o w l i n c en t ra l C h i le a nd s ou ther n S pa in : A c om -

G reenwood ,R .J . , 1 979 . R e la t ing r e s idue i nr a ccoon f e ce st o

H e r re ro , J . ,J .C anu t , D . G a rc ia -Fe r r e , R . G a r c ia -Gonza lez & R . H ida lgo , 1 992 . T he a l p ine m a rmo t (M a rmo ta m a rmo ta

f ood c on sumed . Ame r ican M id land N a tu ra l i s t 1 02 , 1 9 1-

p a ra t ive s t udy . A uk 9 7 , 7 60 -767 .

L . ) i n t h e S pan i sh P y renee s . i e rkund e5 7 ,2 11-215 .

1 93 . G r imm , R .J .& W . M . Wh i tehou se , 1 963 . P e l le tf o rma t ion i n aG rea th o rned ow l : Ar oen tgenog raph ic s t udy . A uk 8 0 , 3 01-306 .

Z e i t sch r if t f ü r S äuge t -

H ev ly , R . H . , 1 981 . P o l len p roduc t ion , t r an spo r t a nd p r e se r v a t ion : P o ten t ia l s a nd l im i ta t ion s i na r chaeo log ica l p a ly n o logy . J ou rna lo fE thnob io logy 1 ,3 9 -54 .

G romov , I .M . &I .Y a . P o lyakov , 1 992 V o le s ( M icro t inae ) . F auna o ft h e USSR , Mamma l s , V o l . 3 ,N o . 8 . Wa sh ing t on : Sm i th son ian I n s t i tu t ion .

H i l l son , S . 1 986 . T ee th . C amb r idge : C amb r idge U n ive r s i ty

H a l lam , A . , 1 989 . G rea t G eo log ica l C on t rove r s ie s ( 2 O xf o rd : O xf o rd Un ive r s i ty P re s s .

e d . ) .

H in ton , M . A . C . , 1 9 11 . T he B r i t i sh f o s s i l s h rew s . G eo log i c a l Maga z ine ,N ew S e r ie s 8 ,5 29 -529 .

H anney , P . , 1 963 . Ob se rva t ion s u pon t h e f ood o ft h e b a rn o w l ( T y to a l ba ) i ns ou ther nN ya sa land , w i th ame thod o f a s cer ta in ing p opu la t ion d ynam ics o fr oden tp r ey . A nna l s a nd Maga z ine o fNa tu ra lH i s to ry S e r ie s 1 3 ,6 ,3 05 -313 .

H in ton , M . A . C . , 1 926 . Monog raph o fV o l e sa nd L emm ing s , V o l . I .L ondon : T ru s tee so ft h eB r i t i sh Mu seum .

H an sen , J .M . , 1 99 1 . T he P a laeoe thnobo tany o fF ranch th i C a ve . B loom ing ton : I nd iana U n ive r s i ty P r e s s .

P re s s .

H of fman , R . , 1 983 . T he c on t r ibu t ion o f r a p to r ia l b i rd s t o p a t te rn ing i ns ma l l mamma l a s semb lage s . P a leob io log y 1 4 , 8 1-90 .

H a r ro ld , F . B . , 1 980 . A c ompa ra t ive a na ly s i s o f E u ra s ian P a leo l i th ic b u r ia l s . W or ld A rchaeo logy 1 2 , 1 95 -211 .

H o lme s , P . L . , 1 990 . D if f e ren t ia l t r an spor t o f s po re s a nd p o l len : Al a bo ra to ry s t udy . R ev iew o fP a laeobo tany a nd P a lyno logy 6 4 , 2 89 -296 .

H av inga , A . J . , 1 967 . P a lyno logy a nd p o l len p re se rva t ion . R ev iew o fP a laeobo tany a nd P a lyno logy 2 ,8 1 -98 .

H owa rd , W . A . , 1 958 . F ood i n take a nd p e l le t f o rma t ion o fa H or ned o w l . W i l son B u l le t in 7 0 , 1 45 150 .

H au s se r , J .& D . J ammo t , 1 974 . ü tude b i ome t r ique d e s m a cho i r e s c hez l e sS orex d ue g roupe a raneu s e n E u rope c on t inen ta le (Mamma l ia , I n sec t ivo ra ) . Mamma l ia 3 ,

H owe s , C . A . , 1 978 . No te s o n t h e f ood o ff oxe s a tG ib ra l ta r P o in t , L inco ln sh i r e . N a tura l i s t , L eed s 1 03 , 2 5 -26 . H u s ton , M . , 1 979 . Ag ene ra l h ypo the s i s o fs pec ie s d ive r s i ty . T he Ame r ican N a tura l i s t 13 , 8 1 10 1 .

6 30 -658 . H ayne s , G . , 1 980 . P rey b one s a nd p r eda to r s : P o ten t ia l e co log ic i n f o rma t ion f r om a na ly s i so fb one s i te s . O s sa 7 , 7 5 -97 . H ayne s , G . , 1 982 . U t i l iza t ion a nd s k e le ta l d i s tu rbance s o f N o r th Ame r ican p rey c a rca s se s . A rc t ic 3 5 ,2 66 -281 .

H u t ton , J . , 1 795 . T he T heory o ft h e E a r th . L ondon : Me ss r s . C ade l l , J r . R ep r in ted i n1 959 , e d . J .C rame r . We inhe im : H . R . E nge lmann . I b a i lez -Angu lo , M . ,

1 99 1 . P a leoe thnobo tan ica l A na ly s i s :

P rob lems a nd Me thodo logy . AC a se S tud y : T he Ca ve o f E lJ uyo ( Leve l s 4a nd 8 ) . U npub l i shed M .A . T he s i s , D e p a r tmen to fA n th ropo logy , U n ive r s i ty o fC h icago .

H e im d e B a l sa c , H . & F .d e B eauf o r t , 1 969 . C on t r ibu t ion ä l ' t tud ed e sM ic romam if e re sd uN o rd -Oue s td e l ' E spange . Mamma l ia 3 3 , 6 30 -658 .

J ak s ic , F .M . , R . L . S e ib & C . M . H e r re ra , 1 982 . P reda t ion H e in r ich , W . -D . , 1 982 . Z u r E vo lu t ion u nd B io s t ra t ig raph ie v on A r v ico la ( Roden t ia , Mamma l ia ) i m P l e i s tozan E u r opa s . Z e i t schr if tf ür G eo log i sche W i s sen schraf t en 1 0 ,

b y t h e b a rn o w l ( T y to a l ba ) i n Med i te r ranean h ab i ta t s o f C h i le , S pa in a nd C a l if or n ia : A c ompa ra t ive a pp roach . Amer ican M id land N a tu ra l i s t 1 07 , 1 51162 .

6 83 -735 . H e in r ich , W . -D . , 1 987 . N eue E rgebn i s se z u r E vo lu t ion u nd B ios t ra t ig raph ie v on A r v ico la ( Roden t ia , Mamma l ia ) i m Q ua r ta r E u ropa s .

J ano s sy , D . , 1 975 . M id -P le i s tocene m ic rof auna s o f c on t i n en ta l E u rope a nd a d jo in ing a r ea s . I nA f t er t h eA u s t ra lo p i thec ine s , . e d s . K . W . B u tz e r & G . L I . I s aac . H ague : Mou ton P ub l ishe r s ,p p . 3 75 -397 .

Z e i t sch r if tf u r G eo log i sche W i s sen -

s chraf i en 1 5 , 3 89 -406 .

1 8 1

T he

J ano s sy , D . &E . S chm id t , 1 975 . E x t r eme V a r ian ten d e s MI i n U nga rn .

K le in , R . G . &K . C ruz -U r ibe , 1 984 T he A na ly s i s o fA n ima l B one sf rom A rcheo log i ca lS i te s . C h icago : U n iv e r s i ty o f C h icago P re s s .

J an s sen s , P .&J .G onza lez E chega ray , 1 958 . Memor ia d el a s

K le in , R . G . &K . C ruz -U r ibe , 1 987 . L af a una m am if e ra d e l

d e r F e ldmau s (M icro tu s a r va l i s P a l la s ) Z e i t schr if tf ür S äuge t ie rkund e4 0 , 3 4 -36 .

E xca vac ione s d e l a C ueva d e l J uyo .

y ac im ien to d e l aC ueva d e " E lJ uyo " . C ampa f ia sd e 1 978 y1 979 . I nE xcavac ione s e n l aC ue va D e lJ uyo , e d s . I . B a rand ia rän Mae s tu , L .G . F reeman ,J .G onza lez E chega -

S an tande r : P a t ro -

n a to d e l a s C ueva s P reh i s tö r ica s d e l aP rov inc ia d e S an t a nde r( E spa f ia ) .

r ay & R . G . K le in . S an tande r : C en t ro d e I nve s t iga c iön y J ed rze jew sk i , W . , B . J ed rze jew sk i & L . S zymu ra , 1 995 .

Mu seo d eA l tam i ra , Monog raf ia sN o . 1 4 ,p p . 9 7 120 .

Wea se l p opu la t ion r e spon se , h ome r a nge , a nd p r eda t ion o n r oden t s i n ad ec iduou s f o re s t i nP o land . E co logy 7 6 , 1 79 195 .

K le in , R . G . , C . Wo lf , L .G . F reeman & K . A l lwa rd en , 1 981 . T he u se o fd en ta l c rown h e igh t s f o rc ons t ruc t ing a ge p ro f i l e s o f r e d d ee r a nd s im i la r s pec ie s i n a r cha eo log ica l

J imenez , R . , M . B u rgo s & R . D iaz d e l a G ua rd ia , 1 984 .

s amp le s . J ourna lo fA rchaeo log i ca lS c ience 8 ,1 31 .

K a ryo type a nd c h romo some b and ing i nt h e mo le ( Ta lpa o cc id en ta l i s ) f r om t h e s ou thea s t o ft h e I b e r ian P en in su la : I mp l ica t ion s o n i t s t a xonom ic p o s i t ion . C a lyo log ia 3 7 , 2 53 -258 . J imenez , R . , M . B u rgo s , A . S änchez & R . D iaz d e l a G ua rd ia , 1 990 . T he r ep roduc t ive c yc le o f T a lpa o cc i -

K o r th , W . W . , 1 979 . T aphonomy o f m ic rover teb ra te f o s s i l a s semb lage s . A nna l s o ft h e C a rneg i e Mu seum 4 8 , 2 35 2 85 .

A c ta

K o t le r , B . P . , J .S . B rown & 0 . H a s son , 1 99 1 . F a c to r s a f f e c t ing g e rb i l f o rag ing b ehav io r a nd r a t e s o fo w l p r eda t i on . E co logy 7 2 , 2 249 -2260 .

J one s , J .K . , J r . , 1 952 . N o te s o n t h e f ood h ab i t s o ft h e g r ea t h or ned o w l i nC he r ry C oun ty , N eb ra ska . N ebra ska B i rd

K o t le r , B . P . , J .S . B rown , R . J . Sm i th & W . 0 . W i r tz I , 1 988 . T he e f f ec t s o f mo rpho logy a nd b ody s i ze o n r a te s o fo w lp r eda t ion o nd e ser tr oden t s . O iko s5 3 , 1 45 152 .

d en ta l i s i n t h e s ou thea s te rn I b e r ian P en in su la . T he r io log ica 3 5 , 1 65 169 .

R e v iew 2 0 , 1 0 11 . J o rdä C e rdä , F . , 1 958 . A vance a lE s tud io d e l aC ueva d e l a L lo se ta . O v iedo : D ipu tac iön P rov inc ia ld eA s tu r ia s . K e l le r , B . L . , 1 985 . R ep roduc t ive p a t te rn s . I n B io log y o f N ew W or ld M ic ro tu s , e d . R . H . T ama r in . T he Ame r ican S oc ie ty o f Mamma log i s t s , S pec ia l P ub l ica t ion N o . 8 ,p p . 7 25 -778 . K e rb i s P e te rhan s , J . C . , 1 990 . T he R o le s o f P o rcup ine s , L eopa rd s , a nd H yena s i n U ngu la te C a rca s s D i spe r sa l : I mp l ica t ion sf or P a leoan th ropo logy . U npub l i shed P h .D . D i s ser ta t ion , D epa r tmen t o fA n th ropo logy , U n ive r s i ty o f C h icago . K e rb i s P e te rhan s , J .C . & L . K o l ska H o rw i tz , 1 992 . Ab one a s semb lage f r om as t r iped h yaena ( H yaena h yaena ) d en i nt h e N egev D e se r t , I s rae l . I s rae l J ourna l o fZ oo logy

K owa l sk i , K . , 1 966 . T he s t ra t ig raph ic i mpo r tance o fr od en t s i nt h e s t ud ie s o n t h e E u ropean Q ua t er na ry . F o l ia Q ua t e rna r ia 2 2 , 1 16 . I C ra tochv i l , J . , 1 970 . P i tymy s -A r ten a u s d e r H ohen T a t ra ( Mamma l ia ,R oden t ia ) . A c ta S c . N a t . B rno 4 ,1 63 . K rupa , M . , 1 994 . L impe t s a nd O the r Ma r ine Mo l lu sc s : A n E va lua t ion o f T he i r A rcheo log ica l S ign if icance a t E l J uyo a nd O ther P a leo l i th ic S i te s i nN o r thern S pa in . U n p ub l ished M . A . T he s i s , D epa r tmen t o f A n th ropo logy , U n ive r s i ty o fC h icago . K rupa , M . , 1 996 . E f f ec t s o fp roce s se s o fd epo s i t ion , t a phon omy a nd e xcava t ion o n t h e i n t e rp r e ta t ion o ff i sh r ema in s f r om U ppe r P a leo l i th ic s i te s i n w e s t e rn E u rope . P ape r p r e sen ted a tt h e 6 1' Annua l Mee t ing o ft h e S oc ie ty f o r Ame r ican A r chaeo logy , Ap r i l 1 ,N ew O r lean s , LA .

3 7 ,2 25 -245 . K ry s tuf ek , B . , 1 986 . V a r iab i l i ty o fe name l t oo th p a t t er n s i n K ing , C . M . , 1 991 . B ody s i z e -p rey s i ze r e la t ion sh ip s i n E u ropean

s t oa t s

i n Mu s te la e rm inea :

A t e s t

M icro tu s a r va l i s f r om D a lma t ia ( Roden t ia , Mamma l ia ) .

c a se .

B io lo sk i V e s tn ik 3 4 , 3 7 -42 .

Ho la rc t i cE co log y 1 4 , 1 73 185 . K ry s tuf ek ,

B . ,

1 990 .

G eog raph ic v a r ia t ion

i n M ic ro tu s

K le in , R . G . , 1 989 . T he H uman C a ree r . C h icago : T he U n i v e r s i ty o fC h icago P re s s .

n i va l i s ( Ma r t in s , 1 842 ) f r om A u s t r ia a nd Y ugo s lav ia . B onne rZ oo log i sche B e i t räge 4 1 , 1 2 1139 .

K le in , R . G . , 1 991 . S i z e v a r ia t ion i nt h e C ape D une mo le ra t

K ry s tuf ek , B . & B . K ova c ic , 1 989 . V e r t ica l d i s t r ibu t ion o f

( Ba thye rgu ss u i l lu s )a nd L a te Q ua te rna ry c l ima t ic c hange

t h e S now v o le M icro tu s n i va l i s ( Ma r t in s ,

i nt h e S ou thwe s ter n C ape P rov ince , S ou th A f r ica . Q ua t e rna ry R e sea rch 3 6 ,2 43 -256 .

N or thwe s ter n Yugo s lav ia . k und e5 4 , 1 53 -156 .

1 82

1 842 ) i n

Z e i t sch r if t f ür S äug e t ie r -

K un iho lm , P . I . , 1 990 . A rchaeo log ica l e v idence a nd n on e v idence f o r c l ima t ic c hange . P h i lo soph ica l T ran sac t i ons o ft h eR oya lS oc ie ty o fL ondon A 30 , 6 45 -655 .

L ig tvoe t , W . &A . V an W i jngaa rden , 1 994 . T he c o lon iza t ion o f t h e I s land o f Noo rd -Beve land ( The N e the r land s ) b y t h e C ommon v o le M icro tu sa r va l i s , a nd i t sc on sequence s f o rt h eR oo tv o le M . o economu s . L u t ra 3 7 , 1 28 .

K u r ten , B . , 1 972 . T he C ave b ea r . S c ien t i f ic Amer ican 2 26 , L loyd , J .A . , 1 975 . S oc ia l s t ruc tu re a nd r ep roduc t ion i nt wo

6 0 -72 .

f r ee ly -g row ing p opu la t ion s o f h ou se mou se (M u s m u s c u /u sL . ) . A n ima lB eha v ior 2 3 ,4 13 -424 .

Ku r ten , B . , 1 976 . T he Ca ve B ea rS to ry . N ew Y o rk : C o lum b ia U n ive r s i ty P re s s . K u sme r , K . D . , 1 990 T aphonomy o fo w l p e l le t d epo s i t ion .

L ock ie , J .D . , 1 959 . T he e s t ima t ion o f t h e f ood o f f oxe s . J ourna lo fW i ld l if e Managemen t2 3 ,2 24 -227 .

J ourna lo fP a leon to logy 6 4 ,6 29 -637 . L ong land , W . S .& S .H . J enk in s , 1 987 . S ex a nd a ge a f f ec t Kw ia tow sk i , A . & K . L ub l ine r -M ianow ska , 1 957 . B adan ie s k ladu c hem ic mego p u lku , 2 . B adan ia b l on p y lkow ich .

v u lne rab i l i ty o fd e se r t r oden t s t oo w l p r eda t ion . J ourna l o f Mamma logy 6 8 , 7 46 -754 .

A c ta S oc . B o tan . P o lon .2 6 , 5 01-514 . L opez , P . , 1 981 . Anä l i s i s p o l in ico d e l y ac im ien to d e L o s L e J acque s , D . & T . L ode , 1 994 . L 'a l imen ta t ion d e l a G ene t t e d 'Eu rope , G ene t ta g ene t ta L . , 1 758 , d an s u n b o c age d el ' oue s td el aF rance . Mamma l ia 5 8 . 3 83 -389 .

A zu le s ( Canga s d e O n i s , O v iedo ) . r ond s ica 8 -9 ,2 43 -248 .

B o tän ica Maca -

L öpez -Be rge s , M . A . &M . V a l le , 1 985 . E s tud io o s t eo lög ico L e L oua rn , H . &G . J aneau , 1 975 . R epa r t i t ion e tb io log ie d u c ampagno l d e s n e ige s M icro tu s n i va l i s Ma r t in s d an s l a r eg ion d eB r iancon . Mamma l ia 3 9 , 5 89 -604 . L e ro i -Gou rhan , A r . , 1 959 . R e su l ta t s d e l ' a na ly se p o l l in ique d e l ag ro t te d ' Is tu r i tz . B u l le t in d e l aS oc ié té P rjh i s to r i q ue F ranva i se 5 6 ,6 19 -624 . L e ro i -Gou rhan , A r . , 1 966 . A nä l i s i s p o l in ico d e l aC ueva d e l O te ro . I nC ueva d e l O te ro , e d s . J .G onza lez E chega ray , M . A .G a rc ia G u inea & A . B eg ine sR am i rez . E xcava c io n e s A rqueo lög ico se nE spa i la , No .5 3 , p p . 8 3 -85 . L e ro i -Gou rhan , A r . , 1 978 . P a l ino log ia . I n V ida y Mue r te e n C ueva Mo r in , e d s . J .G onzä lez E chega ray & L . G . F ree man . S an tande r : I n s t i tuc iön C u l tu ra l d e C an tab r ia , p p . 1 99 -200 . L e ro i -Gou rhan , A r . , 1 980 . A nä l i s i s p o l in ico d e E l P endo . I n E l Y a c im ien to d el aC ueva d e " E lP endo" ( Exca vac ione s 1 953 -1957 ) , e d . J .G onz ä lez E chega ray . Mad r id : B ib l i o theca P raeh i s to r ica H i spana ,V o l . 1 7 ,p p .2 63 -266 L e ro i -Gou rhan , A r . , 1 986 . T he p a lyno logy o fL aR ie ra c ave . I nL aR ie ra Ca ve , e d s . L . G . S t rau s &G . A . C la rk . A r i z ona S t a te U n ive r s i ty , A n th ropo log ica l P ape r s No . 3 6 ,

d e l aC ueva d eP i e lago I . S au tuo la 4 , 13 -121 . L öpez -Fu s te r , M . J . , E . C a s t ien & J .G o sä lbez , 1 988 . R ep ro d uc t ive c yc le a nd p opu la t ion s t ruc tu re o fS orex c o rona tu s M i l le t , 1 828 ( I nsec t ivo ra , S o r ic idae ) i nt h e n o r the rn I b e r i an P en in su la . B onne r Z oo log i sche B e ig t rage 3 9 , 1 63 1 70 . L öpez -Fu s te r , M . J . , 1 989 . R ep roduc t ive s t ra tegy o f t h e M i l le t 's s h rew ( Sorex c o rona tu s M i l le t , 1 928 ) v e r su s t h e c ommon s h rew ( Sorex a raneu s L . , 1 758 ) i nt h e n or thea s t o ft h e I b e r ian P en in su la . Z oo log i sche A bhand lungen 4 4 , 1 43 -148 . L öpez -Fu s te r , M . J .& J .V en tu ra , 1 995 . S oma tome t ry a nd r ep roduc t ive d a ta o n t h e n o r ther n w a te r s h rew , N eomy s f od ien s ( Pennan t , 1 77 1) f r om t h e A ran V a l ley ( Span ish P y renee s ) . Mamma l ia 5 9 , 1 56 159 . L opez -Go rdo , J .L . , E . L aza ro & A . F er nändez Jo rge , 1 977 . C ompa rac ion d e l a s d i e ta s d e S t r ix a luco , A s io o t u s y T y to a lba e n u n m i smo b i o topo d e l aP rov inc ia d e Ma d r id . A rd eo la 2 3 , 1 89 -221 . L ova r i , S . , A . R enzon i & R . F ond i , 1 976 . T he p r eda to ry h ab i t s o ft h e b a rn o w l ( T y to a l ba S copo l i ) i nr e la t ion t o

p p . 5 9 -64 .

t h e v ege ta t ion c ove r . 1 91 .

B o l le t t ino d e Z oo log ia 4 3 , 1 73 -

L e ro i -Gou rhan , A r . , 1 989 . S t ra t ig raph ie d u Wü rm d epu i s 3 5 ,000 a n s p a r l ap a lyno log ie . I nQ ua te rna ry T ype S ec t i on s , e d s . J .R o se & C . S ch luch te r . B a lkema ,p p . 9 1-92 .

R o t te rdam : A . A .

L e ro i -Gou rhan ,A r . , 1 994 . L et emp sd ' E lJ uyo . I n Homena je a lD r . J oaqu in G onza le z E chega ray , e d . J .A . L a she ra s .

L ub in sk i , P .M . , 1 996 . F i sh h ead s , f i sh h ead s : A n e xpe r imen t o n d i f f e ren t ia l b one p re serva t ion i n as a lmon id f i sh . J ou rna lo fA rchaeo log ica lS c ience 2 3 , 1 75 -181 . L ye l l , C . , 1 830 -33 .P r inc ip le so fG eo logy ( 3V o l s . ) . L ondon : J . Mu r ray . R ep r in ted f a c s im i le e d i t ion ( 1991) , e d . M . J .

Mu seo y C en t ro d e I nve s t iga c ion d e A l tam i ra , Mono g raf ia sN o . 1 7 ,p p . 1 23 -129 . L ev in son , M . , 1 982 . T aphonomy o f m ic rover teb ra te s f r om o w lp e l le t st oc ave b recc ia . A nna l s o ft h e T ransvaa l Mu s eum 3 , 15 120 .

S .R udw ick . C h icago : U n ive r s i ty o fC h icago P r e s s . Maddock , A . H . , 1 992 . C ompa r i son o ft wo m e thod s f o rt r ap p ing r oden t s a nd s h rew s . I s rae lJ ou rna l o fZ oo logy 3 8 , 3 33 -340 .

1 83

Mad sen , A . B . &A . M . R a smu s sen , 1 985 . R ep roduc t ion o f t h e s t one ma r ten Mar te sf o ina i nD enma rk . Na tura J u t l and i ca 2 1 , 1 45 148 . Ma r t i , C . D . , 1 969 . S ome c ompa r i son s o ft h e f e ed ing e co l o gy o ff ou r o w l s i n No r th -Cen t ra l C o lo rado . T he S ou th -

Mo r eno , S . , 1 988 . R ep roduc t ion o f g a rden d o rmou se E l io m y s q ue rc inu s l u s i tan icu s , i n s ou thwes t S pa in . Mam m a l ia 5 2 ,4 0 1-407 . Mo reno , S .&E . C o l lado , 1 989 G row th o ft he g a rden d o r mou se ( E l iomy s q uerc inus L innaeu s , 1 766 ) i ns ou thwe s t e r n S pa in .Z e i t schr if tf ü rS äuger t ie rkund e5 4 , 1 00 106 .

w e s te rn N a tura l i s t 1 4 , 1 63 170 . Ma r t i , C .D . , 1 973a . T en y ea r so fb a rn o w lp rey f r om aC o lo r ado n e s ts i te . W i l son B u l l e t in 8 5 , 8 5 -86 .

Mou re R oman i l lo , A . , 1 975 . E xca vac iones e n l a C ueva d e " T i to B u s t i l lo" ( A s tur ia s ) . Ov iedo : I n s t i tu to d e E s tud io s A s tu r iano s .

Ma r t i , C . D . , 1 973b . F ood c on sump t ion a nd p e l le t f o rma t ion r a te si nf ou ro w ls pec ie s . W i l son B u l le t in 8 5 , 1 78 -181 . Ma r t in , P .S .&R . G . K le in , e d s . , 1 984 . Q ua terna ry E x t inc -

Mou re R oman i l lo , A . &M . C ano H e r re ra , 1 976 . E xca vac io n e s e n l aC ueva d e " T i to B u s t i l lo" ( A s tur ia s ) . O v iedo : I n s t i tu to d eE s tud io sA s tu r iano s .

t i on s . T u scon : U n ive r s i ty o fA r izona P re s s . Ma r t inez N ava re t te , M . I .& Y . T . C hapa B rune t , 1 980 . L a i n du s t r ia p reh i s tö r ica d e l ac ueva d e l aP a loma ( So to d e l a s R egue ra s , A s tu r ia s ) . I nL a C ueva d e l aP a loma , e d s . M . H oyo s , M . I .Ma r t inez N ava r e t te , T .C hapa B rune t , P .

Mou re R oman i l lo , A . & M . C ano H e r re ra , 1 978 . Mag d a len ian h ab i ta t ion s t ruc tu re a t T i to B u s t i l lo C ave ( A s tu r ia s , S pa in ) . C ur ren tA n th ropo logy 1 9 , 3 92 -394 . Mou r e R oman i l lo , A . & M . C ano H e r re ra , 1 979 . T i to B u s t i -

C a s taf lo s &F .B . S anchez . E xcavac ione s A rqueo lög ica s e nE spa t ia ,N o . 16 ,p p . 15 -204 .

l o C ave ( A s tu r ia s , S pa in ) a nd t h e Magda len ian o f C an tab r ia . W or ld A rchaeo logy 1 0 ,2 80 -289 .

Mayhew , D . F . , 1 977 . A v ian p r eda to r s a s a ccumu la to r s o f f o ss i l mamma lm a te r ia l . B orea s6 ,2 5 -31 .

Mou r e R oman i l lo , A . , C . G onza lez S a inz , F . B e rna ldo d e Q u i rö s &V . C ab re ra V a ldis , 1 996 . D a ta c ione s a b so lu ta s d e p igmen to s e n c ueva s C an täb r ica s : A l tam i ra , E l C a s t i l o , C h imenea s yL a s Moneda s . I n " E l Hombre F ö s i l" 8 0 Mo s s D e spuä s , e d . A . Mou re R oman i l lo . S an tande r : U n ive r s idad d e C an tab r ia , F undac iön B o t in , a nd I n s t i tu to p a ra I nve s t igac ione sP reh i s tö r ica s ,p p . 2 95 -324 .

Mc In ty r e , A . ,N . G . K ipp , A . W . H . B e , T .C row ley , T .K e l l o gg , J .V . G a rdne r , W . P r e l l , &W . F .R udd iman , 1 976 . G lac ia l N o r th A t lan t ic 1 8 ,000 y ea r s a go : A CL I MAP r e c on s t ruc t ion . I nI nve s t iga t ion o fL a te Q ua te rnary P a leo c eanog raphy a nd P a l eoc l ima to logy , e d s . R . M . C l ine & J .D . H ay s . T he G eo log ica l S oc ie ty o f Ame r ica , Memo i r 1 45 ,p p .4 3 -76 . Me l la r s , P . , 1 990 . T he Eme rgence o f Mod e rn H uman s . a ca : C or ne l lU n ive r s i ty P re s s .

N adachow sk i , A . , 1 982 . L a te Qua te rna ry r oden t s o fP o land w i th s pec ia l r e f e rence t o mo rpho type d en t i t ion a na ly s i s o fv o le s . P o l ska A kad em ia Nauk 1 982 , 1 108 .

I t h -

Me l la r s , P . & C . S t r inge r , e d s . , 1 989 . T he H uman R evo lu t i on . E d inbu rgh : U n ive r s i ty o fE d inbu rgh P re s s . Me l le t , J .S . , 1 974 . S ca to log ica l o r ig in o f m ic rover teb ra te f o s s i la ccumu la t ion s . S c ience 1 85 , 3 49 -350 . M icha lak , I . ,1 987 . G row th a nd p o s tna ta l d eve lopmen t o ft h e E u ropean w a te rs h rew . A c ta T her io log ica 3 2 , 2 61-288 .

N adeau , J .H . , 1 985 . O n togeny . I n B io lo gy o fN ew W o r ld M icro tu s , e d . R . H . T ama r in . T he Ame r ican S oc ie ty o f Mamma log i s t s , S pec ia l P ub l ica t ion N o . 8 ,p p . 2 54 -285 . N ade r , I .A . , 1 969 . A n ima l r ema in s i np e l le t s o ft h eb a rn o w l T y to a l ba f r om t h ev i c in i ty o f An -Na ja f , I r aq . B u l le t in o f t h eI r aq N a tu ra lH i s tory Museum 4 ,1 7 . N ago r sen , D . W . & R . L . P e ter son , 1 980 . Mamma l C o l l ec t o r s ' Manua l . L if e S c ience s M isce l laneous P ub l ica t ion s , R oya lO n ta r io Mu seum . T o ron to : A lg e rP re ss .

M ikko la ,H . , 1 983 . Ow l so fE u rope . C a l ton : P oy se rP re s s . N ee t , C . R . &J .H au s se r , 1 990 . H ab i ta t s e lec t ion i nz one s o f M i l l s , D . , 1 989 . E co log y a nd Managemen t o f A t lan t ic S a lmon . L ondon , C hapman a nd H a l l .

p a rapa t r ic c on ta c t b e tween t h e c ommon s h rew S or ex a raneu s a nd M i l le t 's s h rew S . c o rona tu s . A n ima lE co lo gy 5 9 , 2 35 -250 .

J ou rna l o f

Moon , E .L . , 1 940 . N o te s o n h awk a nd o w l p e l le t f o rma t ion a nd i d en t if ica t ion . T ran sac t ion s o ft h e K an sa s A cad emy o fS c ience 4 3 ,4 57 -466 .

N e tboy ,A . , 1 980 .S a lmon . T u lsa : W inche s te rP re s s . N ie thamme r , J . & F . K rapp , e d s . , 1 978 . Hand buch d e r

Mo reno , E . & A . B a rbo sa , 1 992 . D i s t r ibu t ion p a t ter n s o f

S äuge t ie r e E u ropa s .

s ma l l mamma l f a una a l ong g rad ien t s o fl a t i tude a nd a l t i t ude i nn o r ther n S pa in . 5 7 , 1 69 175 .

B and I :N age t ie re I . W ie sbaden :

A kadem i sche V e r lag sge se l l schaf t .

Z e i t sch r if tf ü r S äuge r t ierkund e N ie thamme r , J . & F . K rapp , e d s . , 1 982 . Hand buch d e r S äuge t ie re E u ropa s . B and 2 / I : Nage t i e re I . W ie sbaden : A kadem i sche V e r lag sge se l lschaf t . 1 84

N ie thamme r , J . & F . K rapp , e d s . , 1 990 . Hand buch d er S äuge t ie re E uropa s . B and 3 / 1 : I n sek tenf re s ser , H e r ren t i ere . V e r lag W ie sbaden : AULA .

P e r r in , M . R . , 1 982 . P rey s pec if ic i ty o f t h e b ar n o w l , T y to a l ba , i nt h e G rea t F i sh R ive r V a l ley o ft h e E a s ter nC ape P rov ince . S ou th Af r ican J ourna lo fW i ld l if eR e sea rch 1 2 , 1 4 -26 .

N ie thamme r , J . & F . K rapp , e d s . , 1 993 . Hand buch d er S äuge t ie re E uropa s .

B and 5 / 1 1 : R aub säuge r .

V e r lag

W ie sbaden : AULA .

P i te lka , F .A . , P . Q . T om ich & G . W . T re iche l , 1 955 . E co l o g ica l r e la t ion s o fj aege r s a nd o w l s a s l emm ing p r eda t o r s n ea r B a r row , A la ska .

Noyes , R . F . , G . W . B a r re t t & D . H . T ay lo r , 1 982 . S oc ia l s t ruc tu re o ff e ra l h ou se mou se (M u s m u scu lu s L . ) p opu l a t ion s : E f f ec t s o f r e sou rce p a r t i t ion ing . B eha v io ra l E co lo gy a nd S oc iob io logy 1 0 , 1 57 163 . O be rma ie r , H . , 1 924 . F o s s i l Man i nS pa in . Y a le U n ive r s i ty P re s s .

E co log ica l Monog raph s 2 5 ,

8 5 -117 . P ok ine s , J .T . , 1 993 . A n t ler P o in t sf rom E lJ uyo ( San tand e r , S pa in ) : F o rm , Manuf ac ture a nd P a rame te r s o fU se . U n p ub l i shed M .A . T he s is , D epa r tmen t o f A n th ropo logy , U n ive r s i ty o fC h icago .

N ew H aven :

P ok ine s , J .T . , 1 997 . M icrof auna l I nd ica to r s o fP a leoenv i r onmen tf rom E lJ uyo ( San tand er , S pa in ) . U npub l i shed P h .D . D i s ser ta t ion , D epa r tmen t o f A n th ropo logy , U n i -

O gnev , S .I . ,1 962 . Mamma l s o fE a s te rn E u rope a nd N o r th e rn A s ia . V o l . I : C a rn ivora ( F i s s iped ia ) . J e ru sa lem : I s r a e lP rog ram f o rS c ien t if ic T ran s la t ion s .

v e r s i ty o fC h icago .

O gnev , S .I . ,1 964 . Mamma l s o fE a s te rn E u rope a nd N o r th e rn A s ia . V o l . V I I : R od en t s . J e ru sa lem : I s rae l P rog ram f o rS c ien t if i cT rans la t ion s .

P ok ine s , J .T . , i np re s s . E xpe r imen ta l r ep l ica t ion a nd u se o f

O k sanen , T . , 1 983 . P rey -cach ing i nt h e h un t ing s t ra tegy o f s ma l lm amma l s . A c ta Z oo log i ca F enn ica 1 74 , 1 97 199 .

P ok ine s , J .T . & J .K e rb i s P e te rhan s , 1 998 . B ar n o w l ( T y to a l ba ) t a phonomy i n t h e N egev D e ser t , I s rae l . I s rae l J ou rna lo fZ oo logy 4 4 .

C an tab r ian L owe r Magda len ian a n t le rp ro jec t i le p o in t s . J ourna lo fA rchaeo log ica lS c ience .

O r s in i , P . , 1 985 . L e r eg ime a l imen ta i r e d u h ibou g rand -duc B ubo b ubo e nP rovence . A lauda 5 3 , 1 -28 . P a lme i r im , J .M . & R . S .H offman , 1 983 . G a lemy sp y rena i c u s . Mamma l ian S pec ie s2 07 , 1 5 . P a lomo , L .J . ,J .M . V a rga s &M . P .J imenez -Gömez , 1 994 . D i s t r ibu t ion p a t te rn s i nI b e r ian P en in su la r oden t s . P o l i sh E co log i ca lS tud ie s2 0 ,4 97 -502 . P a s i t schn iak -A r t s , M . & S . L a r iv ie r e ,

1 995 . G u lo g u lo .

Mamma l ian S pec ie s4 99 , 1 10 . P ayne , R . S . , 1 971 . A cou s t ic l o ca t ion o fp r ey b y b a rn o w l s ( T y to a l ba ) . J ourna l o fE xpe r imen ta l B io logy 5 4 , 5 35 5 73 .

P ok ine s , J .T . & M . K rupa , 1 997 . S e lf -ba rbed a n t le r s pea r p o in t s a nd e v idence o f f i sh ing i nt h e L a te U ppe r P a leo l i t h ic o fC an tab r ian S pa in . I nP ro jec t i le T echno logy , e d . H . K nech t . N ew Y o rk , P l enum P ub l ish ing , p p . 2 4 1-262 . P o lun in , 0 . & M . Wa l te r s , 1 985 . AG u id e t ot h e V ege ta t ion o fB r i ta in a nd E urope . O xf o rd : O xf o rd U n ive r s i ty P r e s s . P ren t ice , I .C . , 1 986 . F o res t -compo s i t ion c a l ib ra t ion o fp o l l e n d a ta . I n Handbook o f Ho locene P a laeoeco log y a nd P a laeohyd ro logy ,e d . B . E .B e rg lund . N ew Y o rk : W i ley I n te r sc ience , p p . 7 99 -816 .

P ayne , R . S .& W . H . D ru ry , J r . , 1 958 . Ma rk sman o f t h e d a rkne s s : T y to a lba P a r tI . N a tu ra lH i s to ry 6 7 ,3 16 -323 .

P ucek , Z . & V . P . W . L owe , 1 975 . A ge c r i te r ia i n s ma l l mamma l s . I n S ma l l Mamma l s : T he i r P roduc t iv i ty a nd P opu la t ion D ynam ic s , e d s . F . B . G o l ley , K . P e t ru sew icz & L . R y szkow sk i . C amb r idge : C amb r idge U n ive r s i ty P re s s ,p p . 5 5 -72 .

P emän , E . , 1 985 . A spec to s c l imä t ico s ye co lög ico s d e l o s

R eed , C . A . , e d . , 1 977 . O r ig in s o fA g r icu l tu re . T he H ague :

m icromam if e ro s d e l y ac im ien to d e E r ra l la .

Mun ibe 3 7 ,

Mou ton P re s s .

4 9 -57 . R aczyn sk i , J .& A . L . R up rech t , 1 974 . T he e f f ec t o fd i ge s P emän , E . , 1 990 . L o s m ic romam if e ro s d e l a C ueva d e

t i on o nt h e o s t eo log ica l c ompo s i t ion o fo w lp e l le t s . A c ta

Ama lda y s u s i gn if icado . I n L a C ue va d e Ama lda ( Z e s toa , P a i s V a sco ) , e d s . J .A l tuna , A . B a ldeön & K . Ma r iezku r r ena .

S an S eba s t ian : S oc iedad d e E s tud io s

V a sco s ,p p .2 25 -238 .

O rn i tho log ica 1 4 ,2 6 -38 . R eed , C . I .& B . P . R eed , 1 928 . T he m echan i sm o f p e l le t f o rma t ion i n t h e g rea t h or ned o w l ( Bubo v i rg in ianu s ) . S c ience 6 8 , 3 59 -360 .

Paa lba , M . C . , 1 994 . T he h i s to ry o ft h e H o locene v ege ta t i on i nn o r the rn S pa in f r om p o l len a na ly s i s . J ourna l o f E co logy 8 2 , 8 15 -832 .

R e ich s te in , H . , 1 963 . B e i t rag z u r s y s tema t ischen G l iederung d e s G enu s A r v ico la L acepede 1 799 . Z e i t schr if tf ü r z o o log i sche S y s tema t ik u nd E vo lu t ion s -f o r schung 1 , 1 55 2 04 . 1 85

R obe r t son ,P .A . &J . Whe lan , 1 987 . T he f ood o ft h eR ed f ox ( Vu lpe sv u lpe s ) i nC o . K i lda re , I r e land . J ourna lo fZ oo l o g y , L ondon 2 13 , 7 40 -743 . R od r iguez , J .& G . F lo r , 1 979 . E s tud io d e l y ac im ien to p r e h i s tö r ico

d e

B a r iugue s

y s u

m dd io

d e

S h ipman , P . , 1 981 .L if eH i s tory o f aF o s s i l : A nI n t rod uc t ion t o T aphonomy a nd P a leoeco log y . C amb r idge : H a rva rd U n ive r s i ty P re s s . S h ipman , P . & A . Wa lke r , 1 980 . B one -co l lec t ing b y h a r v e s te ra n t s . P a leob io logy 6 ,4 96 -502 .

d epo s i to .

Z ephy ru s ,2 9 :205 -222 . R udd iman , W . F . & A . Mc In ty re , 1 976 . No r thea s t A t lan t ic

S ho r t , H . L . & L . C . D rew , 1 962 . O b serva t ion s c once rn ing b ehav io r , f e ed ing , a nd p e l le t s o f s ho r t -ea red o w l s . Ame r i can M id land Na tura l i s t6 7 ,4 24 -433 .

p a leoc lüna t ic c hange s o ve r t h e p a s t 6 00 ,000 Y ea r s . I n I nve s t iga t ion o fL a te Q ua terna ry P a leoceanog raphy a nd P a leoc l ima to logy , e d s . R . M . C l ine & J .D . H ay s .

T he

G eo log ica l S oc ie ty o f Ame r ica , Memo i r 1 45 . p p . 111 46 . R up rech t , A . L . , 1 971 . T axonom ic v a lue o f mand ib le m ea s u r emen t s i n S o r ic idae ( I n sec t ivo ra ) .

A c ta T he r io log ica

S k inne r , J .D . &R . H . N . Sm i the i s , 1 990 . T he Mamma l s o f t h e S ou the rn Af r ican S ubreg ion . P re to r ia : U n ive r s i ty o f P re to r ia . S l eeman , D . P . , 1 993 . H ab i ta t s o ft h e I r i sh s t oa t . I r i sh N a tu r a l i s t ' sJ ou rna l2 4 , 3 18 -321 .

1 6 , 3 4 1-357 . S a in tG i ron s ,M . -C . , 1 973 . L 'age d e sm ic romam if e r e sd an s l e r eg ime d ed eux r apace sn oc tu rne s , T y to a l ba e tA s io o t u s . Mamma l ia 3 7 ,4 39 -456 . S anchez G o f i i , M . F . , 1 993 . D e l a T aphonom ie P o l l in ique ä

Sm i th , C . R . & M . E . R ichmond , 1 972 . F a c to r s i nf luenc ing p e l le t e ge s t ion a nd g a s t r ic p H i nt h e b a rn o w l . W i l son B u l le t in 8 4 , 1 79 -186 . S ou le , M . E . , 1 992 . C on se rva t ion : T a c t ic s f o r ac on s tan t c r i s i s . S c ience 2 53 , 7 44 -750 .

l aR econ s i tu t ion d e l ' Env i ronmen t : L 'Exemp le d e l aR e g ion C an tabr ique . B r i t i sh A r chaeo log ica l R epor t s , I n te r n a t iona l S e r ie s5 86 .

S ou ther n , H . N . , 1 954 . T awny o w l s a nd t h e i r p r ey . I b i s 9 6 , 3 84 -4 10 .

S an to s R e i s , M . , 1 983 . S t a tu s a nd d i s t r ibu t ion o ft h e P o r tu g uese mu s te l id s . A c ta Z oo log ica F enn i ca 1 74 , 2 13 -216 .

S p i tz , F . , 1 978 . ü tude c r än iomd t r ique d u g en re P i tymy s . Mamma l ia 4 2 ,2 67 -304 .

S aunde r s , J r . , J .K . , 1 963 . F ood h ab i t s o ft h e l y nx i nN ew f ound land . J ou rna l o f W i ld l if e Managemen t 2 7 , 3 84 3 90 .

S t e t t le r , H . K . , 1 995 . Macrobo tan i ca lR ema in sf rom t h e S i te o fE l J uyo ( San tand e r ) , S pa in S pa t ia l A na ly s i s a nd P a l e oenv i ronmen ta l I mp l ica t ion s . U npub l i shed M .A . T he s i s , D epa r tmen to fA n th ropo logy , U n iver s i ty o f C h icago .

S co t t , T .G . , 1 941 . Me thod s a nd c ompu ta t ion i nf e ca l a na ly s i s w i th r e f e r ence t o t h e r ed f ox . I owa S ta te C o l lege J ourna lo fS c ience 1 5 ,2 79 -285 . S e l leck , D . M . & B . G lad ing , 1 943 . F ood h ab i t s o fn e s t ing b a rn o w l sa nd Ma r sh h awk s a tD une L ake s , C a l if o rn ia , a s d e t e rm ined b y t h e " Cage N e s t " m e thod . C a l if orn ia F i sh a nd G ame 2 9 , 1 22 -131 .

S t ine r , M . C . , 1 993 . Sma l l a n ima l e xp lo i ta t ion a nd i t s r e la t i on t oh un t ing , s caveng ing , a nd g a the r ing i nt h e I t a l ian Mou s te r ian . I n Hun t ing a nd A n ima l E xp lo i ta t ion i n t h e L a te r P a laeo l i th i c a nd Me so l i th i c o f E ura s ia , e d s . G . L a r sen P e te rk in , H . M . B r icke r & P . Me l la r s . A r cheo l o g ica l P ape r s o ft h e Ame r ican A n th ropo log ica l A s soc ia t i on ,N o .4 ,p p . 1 07 -125 .

S hack le ton , N . J . , 1 987 . O xygen i s o tope s , i c e v o lume a nd s ea l e ve l . Q ua te rnary S c ience R ev iew s , 6 : 183 -190 .

S t ine r , M . C . , 1 994 . H ono r Among T h ieve s . P r ince ton : P r in c e ton U n ive r s i ty P re s s .

S hack le ton , N . J .& N . D . O pdyke , 1 976 . O xygen i so tope

S to rk , N . E . , 1 988 . I n sec t d ive r s i ty : F ac t s , f i c t ion a nd s pecu l a t ion .

a nd p a leomagne t ic s t ra t ig raphy o fP ac if ic c o re V 28 -239 L a te P l iocene t o L a te s t P l e i s tocene . I n I nve s t iga t ion o f L a te Q ua te rna ry P a leoceanog raphy a nd P a leoc l ima to l o gy ,e d s . R .M . C l ine & J .D . H ay s . G eo log ica l S oc ie ty o f Ame r ica , Memo i r 1 45 , p p .4 49 -464 . S haf f e r , B . S .&J .L .J .S anchez , 1 994 . C ompa r i son o f 1 / 8" a nd 1 / 4" -me sh r e cove ry o f c on t ro l led s amp le s o fs ma l l -

S t rau s , L .G . , 1 974a . L e s o lu t reen d u P ay s B a sque E spagno l : U ne e squ i s se d e sd onnee s . Mun ibe 2 6 , 1 73 -181 . S t rau s , L . G . , 1 974b . No ta s p re l im ina r e s s ob re e lS o lu t r en se d e A s tu r ia s . B o le t in d e l I ns t i tu to d e E s tud io s A s tu r iano s

t o -med ium s ized mamma l s . Ame r ican A n t iqu i ty 5 9 , 5 25 5 30 . S heff ie ld , S . R . & C . M . K ing , 1 994 . Mu s te la n i va l i s . Mamma l ian S pec ie s4 54 , 1 3 .

J ou rna l o ft h e L innean S oc ie ty o fL ond on 2 0 ,

3 21-338 .

8 2 , 4 83 -504 . S t rau s , L .G . , 1 975 E l S o lu t ren se d e l a s c ueva s d e l C a s t i l lo y H omo sd e l aP ena ( San tande r )e n e l Mu seo A rqueo lög ico N ac iona ld e Mad r id . T raba jo sd eP reh i s tor ia 3 2 , 9 19 .

1 86

S t raus , L . G . , 1 976 . Anä l i s i s a rqueo lög ico d e l af a una p a l e o l i t ica d e l n one d e l a P en i su la I b e r ica . Mun ibe 2 8 ,

S t rau s , L .G . &G . A . C la rk , 1 978 . P reh i s to r ic i n ve s t iga t ion s i n C an tab r ian S pa in . 2 89 -317 .

2 27 -285 . S t raus , L . G . , 1 976 -77 . T he U ppe r P a leo l i th ic c ave s i te o f A l tam i ra ( San tande r , S pa in ) . Q ua te rna r ia 1 9 , 1 35 148 .

J ourna l o fF ie ld A rchaeo lo gy 5 ,

S t rau s , L .G . &G . A . C la rk , e d s . , 1 986 . L aR ie ra Ca ve . A r i z ona S t a te U n ive r s i ty A n th ropo log ica l R e sea rch P ape r s No . 3 6 .

S t raus , L . G . , 1 977 . O fd ee r s laye r s a nd moun ta in men : P a l e o l i th ic f a una l e xp lo i ta t ion i nC an tab r ian S pa in . I n F or T heory B u i ld ing i nA rchaeo logy , e d . L .R .B inf o rd . N ew Y o rk :A cadem ic P re s s ,p p .4 1-76 .

S t rau s ,L .G . , G .A . C la rk , J .A l tuna & J .A . O r tea , 1 980 . I c e A ge s ub s i s tence i nn or the rn S pa in . S c ien t i f ic Amer ican

S t rau s ,L .G . , 1 980 . R ep ly t oC onkey . C u r ren tA n thropo lo gy

S tua r t , A. J . , 1 980 . T he v er teb ra te f auna f r om t h e i n te rg la c ia l d epo s i t s a tS ugwo r th ,n ea rO xfo rd . P h i lo soph ica l T ran s -

2 1 ,6 24 -625 .

2 42 , 1 42 -152 .

a c t ion so ft h eR oya lS oc ie ty o fL ondon , B2 89 , 8 7 -97 . S t rau s , L . G . , 1 982 . C a rn ivo re s a nd c ave s i t e s i nC an tab r ian S pa in . J ou rna lo fA n th ropo log ica lR e sea r ch 3 8 , 7 5 -98 . S t rau s , L .G . , 1 985 . S tone A ge p r eh i s to ry o fn or the rn S pa in . S c i ence 2 30 , 5 01-507 .

S tua r t , A. J . , 1 981 . Ac ompa r i son o ft h e M idd le P l e i s tocene mamma l f auna s o fV o ig s ted t( Thu r ing ia , GDR ) a nd We s t R un ton ( No rf o lk , E ng land ) . Q ua r ta rpa laon to log ie 4 , 1 55 163 .

S t rau s , L . G . , 1 987a . H un t ing i n L a te U ppe r P a leo l i th ic We s t e rn E u rope . I n T he E vo lu t ion o fH uman Hun t ing ,

S tua r t , A . J . , 1 982 . P l e i s tocene V er tebra te s i n t h e B r i t i sh I s le s . L ondon : L ongman .

e d s . M . H .N i teck i&D . V . N i teck i . N ew Y o rk : P lenum P re ss ,p p . 1 47 176 .

S tua r t , C . T . , 1 977 . A na ly s i s o f F e l l s l i by ca a nd G ene t ta g ene t ta s c a t s f r om t h e C en t ra l Nam ib D e se r t , S ou th We s t A f r ica . Z oo log ica Af r icana 1 2 , 2 39 -24 1 .

S t rau s , L . G . ,1987b . U ppe r P a leo l i th ic i b ex h un t ing i n s ou thwe s t E u rope . J ourna l o fA rchaeo log ica l S c ience

S u tc l if f e , A . J . , 1 970 . S po t ted h yaena : C rushe r , g nawe r , d i g e s te ra nd c o l lec to ro fb one s . Na tu re 2 27 , 110 -1113 .

1 4 , 1 63 178 . S t rau s , L .G . , 1 990 . T he o r ig ina l a rms r a ce : I b e r ian p e r spec t i ves o n t h e S o lu t rean p henomenon . I nF eu i l le s d e P i e r re , e d . J . K oz low sk i . E tude s e t R eche r che s A r c heo log ique sd e l 'Un ive r s i te d eL iege ,p p .4 25 -447 .

T ay lo r , I . , 1 994 . B a rn Ow l s .

C amb r idge :

C amb r idge U n i -

v e r s i ty P re s s . T cher nov , E . , 1 968 . S ucce s s ion o fR od en tF auna s d ur ing t h e U ppe r P le i s tocene o f I s rae l . P a rey .

S t rau s , L . G . , 1 99 1a . E p ipa leo l i th ic a nd Me so l i th ic a dap ta t i ons i nC an tab r ian S pa in a nd P y r enean F rance . J ourna l o fW o r ld P reh i s to ry 5 ,8 3 104 . S t rau s , L .G . , 1 99 1b . S ou thwe s ter nE u rope a tt h e l a s t g l ac ia l max imum . C ur ren tA n th ropo logy 3 2 , 1 89 199 . S t rau s , L . G . , 1 992 . I be r ia b ef ore t h e I be r ian s . q ue : U n ive r s i ty o fN ew Mex ico P re s s .

A lbuque r -

S t rau s , L . G . , 1 993 . U ppe r P a leo l i th ic h un t ing t a c t ics a nd w eapon si n We s te rn E u rope . I nH un t ing a nd A n ima lE x p lo i ta t ion i n t h e L a ter P a laeo l i th ic a nd Me so l i th ic o f E ura s ia , e d s . G . L a r sen P e te rk in , H . M . B r icke r & P . Me l la r s . A r cheo log ica l P ape r s o ft h e Ame r ican An th ro -

H ambu rg : V e r lag P au l

T cher nov , E . , 1 975 . R oden t f auna s a nd e nv i ronmen ta l c hange s i nt h e P l e i s tocene o fI s rae l . I nR od en t s i nD e ser t E nv i ronmen t s , e d s . I . P raka sh & P . K . G ho sh . Monog raph iae B io log icae , V o l . 2 8 . T he H ague : D r . W . J unk ,p p . 3 31-362 . T che rnov , E . , 1 979 . Q ua te rna ry f auna . I nT he Q ua terna ry o f I s rae l , e d . A . H o row i tz . N ew Yo rk : A cadem ic P re s s , p p . 2 47 -290 . T che rnov , E . ,

1 982 . F auna l r e spon se s t o e nv i ronmen ta l

c hange s i n t h e e a s te rn Med i ter ranean d u r ing t h e l a s t 2 0 ,000 y ea r s . I n P a laeoc l ima te s , P a laeoenv i ronmen t s a nd H uman C ommun i t ie s i n t h e E a s te rn Med i te r ranean R eg ion i nL a te rP reh i s tory , e d s . J .L . B in t l if f& W . V an Z e i s t . O xf o rd : B r i t i sh A rchaeo log ica l R epo r t s I n te rna -

p o log ica lA s soc ia t ion , No . 4 ,p p . 8 3 -93 . S t rau s , L aw r ence G . , e d . , 1 995 . L e s D ern ie r s C ha s seur s d e R enne s d u Mond e P y rdnden . L 'Abr i D uf aure : U n G i se -

t i ona l S e r ie s , 1 33 1 ,p p . 1 05 -127 .

m en t T a rd ig lac ia re e n G a scogne . P a r i s : S oc ie te P reh i s t o r ique F r an p i se , Memo i re 2 2 .

T e s s ie r , L . ,J . -L . d eB eau l ieu , M . C on teaux , J . -L . E doua rd , P . P ane l , C . R o lando , M . T h inon , A . T homa s & K . T obo l s k i , 1 993 ) . H o locene p a laeoenv i ronmen t s a tt h e t imbe r l i ne i n t h e F rench A lp s -a mu l t id i sc ip l ina ry a pp roach . B orea s2 2 , 2 44 -254 .

S t rau s , L .G . , J .A l tuna , G . A . C la rk , M . G onza lez , H . L av i l e , A r . L e ro i -Gou rhan , M . Menendez & J .A . O r tea , 1 981 . P a leoeco logy a tL a R ie ra ( A s tu r ia s , S pa in ) . r en tA n th ropo lo gy 2 2 ,6 55 -674 .

C ur -

1 87

T hacke ray , J . F . , 1 987 . L a te Q ua te rna ry e nv i ronmen ta l c hange s i n f e r red f r om s ma l l m amma l ian f a una , S ou the rn A f r ica . C l ima t ic C hang e 1 0 ,2 85 -305 .

V en tu ra , J . , 1 99 1 . Mo rpho log ica l c ha rac ter i s t ic s o ft h e mo l a r s o fA rv ico la t e r re s t r i s ( Roden t ia , A rv ico l ida e ) i n i t s s ou thwe s t e rn d i s t r ibu t ion a r ea . Z oo log i sche A n ze ige r 2 26 , 6 4 -70 .

T hacke ray , J .F . & D . M . A ve ry , 1 990 . A c ompa r i son b e t ween t empe ra tu re i n d ice s f o rL a te P l e i s tocene s equence s a t K la s ie s R ive r a nd B o rde r C ave , S ou th A f r ica .

P a -

l a eoeco logy o fAf r ica 2 1 , 3 11-316 .

V en tu ra , J . , 1 992 . Mo rphome t r ic d a ta o n t h e s c apu la a nd l i mb l o ng b one s o fA r v ico la t e r re s t r i s ( L innaeu s , 1 758 ) ( Roden t ia , A rv ico l idae ) . R evue S u i s se Z oo log ie 9 9 , 6 29 6 36 .

T o r re , I .& J .L .T e l la , 1 994 . D i s t r ibu t ion o ft h e C ab re ra w a t e r s h rew ( N eomy s a noma lu s ) i n n o r thea s ter n S pa in . Z e i t schr if tf ü rS äuge t ie rkund e5 9 ,2 82 -288 .

V en tu ra , J .& J .G o sä lbez , 1 986 . C a ra c t e r i s t ica s mo rf o l6g i c a s yc r i t e r io s d e d i sc r im inac iön s exua l an ive l d e l c oxa l e n A r v ico la s ap idu s ( Roden t ia , A rv ico l idae ) .

T rave r se ,A . , 1 990 . S tud ie so fp o l len a nd s po re si nr i ve r s a nd

P ub l ica -

c i one s d e l D epa r tmen to d e Z oo log ia d e B a rce lona 1 2 , 17 128 .

o t he r b od ie s o fw a te r i nt e rm s o fs ou rce -vege ta t ion a nd s ed imen ta t ion ,w i th s pec ia lr e f e rence t oT r in i ty R ive ra nd B ay , T exa s . R ev iew o fP a laeobo tany a nd P a lyno logy 6 4 ,

V en tu ra , J .& J .G o sä lbez , 1 987 . R ep roduc t ive b i o logy o f A r v ico la s ap idu s ( Roden t ia , A rv ico l idae ) i n t h e E b ro d e l ta , S pa in . Z e i t schr if tf ü rS äuge t ie rkund e5 2 , 3 64 -37 1 .

2 97 -303 . T rou t , R . C . , 1 978 . A r ev iew o f s t ud ie s o n p opu la t ion s o f w i ld h a rve s t m ice (M ic romy s m inu tu s P a l la s ) . Mamma l R ev iew 8 ,1 43 -158 .

V en tu ra , J .& J .G o sä lbez , 1 989 . T axonom ic r ev iew o fA r v i c o la t e r re s t r i s ( L innaeu s , 1 758 ) ( Roden t ia , A rv ico l idae ) i nt h e I b e r ian P en in su la . 4 0 ,2 27 -242 .

T um l i son , R . , 1 987 . F e l i s l y nx . Mamma l ian S pec ie s 2 69 , 1 8 .

B onne r Z oo log i sche B e i t räge

V en t u ra , J .& J .G o sä lbez , 1 990 . R ep roduc t ive c yc le o fA r v i T u rn i , H . & E . F . Mü l le r , 1 996 . U n te r sche idung d e r S p i tz mau sa r ten S orex a raneu s L . , 1 758 u nd S o rex c orona tu s M i l le t , 1 828 m i tH i lf e e i ne rn euen D i s l c r im inanzf unk t ion . Z e i t schr if tf ü rS äug e t ie rkund e6 1 , 7 3 -92 .

c o la t e r re s t r i s ( Roden t ia , A rv ico l idae ) i nt h e A ran V a l l e y , S pa in . Z e i t schr if tf ü rS äuge t ie rkund e5 5 , 3 83 -391 . V i rgö s , E . , J .G . C a sanova s & T . B läzquez , 1 994 . G ene t ( Gene t ta g ene t ta L . , 1 758 ) d i e t s h if t i n moun ta in s o f

U t r i l la , P . , 1 981 . E l Magda l en ien se I nf er ior y Mgd io e n l a C o s ta C an täbr ica . S an tande r : C en t ro d e I nve s t igac ion y Museo d eA l tam i ra , Monog ra f ia N o . 4 . V a l lada s ,H . , H . C a ch ie r , P . Mau r ice , F .B er na ldo d e Q u i rö s , J .C lo t te s , V . C ab re ra V a ldd s , P .U zqu iano & M . Ar no ld , 1 992 . D i rec t r ad ioca rbon d a te s f o r p reh i s to r ic p a in t ing s a tt h eA l tam i ra , E lC a s t i l lo a nd N iaux c ave s . N a ture 3 57 , 6 8 -70 . v an D onk , J . , 1 976 . O ' sR eco rd o ft h e A t lan t ic O cean f o rt h e e n t i r e P l e i s tocene e poch . I n I nve s t iga t ion o fL a te Q ua t e rna ry P a leoceanog raphy a nd P a leoc l ima to logy , e d s . R . M . C l ine & J .D . H ay s . G eo log ica l S oc ie ty o f Ame r ica , Memo i r 1 45 ,p p . 1 47 -163 . V ega d e lS e l la ,C onde d e l a , 1 9 16 .P a leo l i t ico d eC ue to d el a M ina . Mad r id : C om i s iön d e I nve s t iga c ione s P a leon -

c en t ra l S pa in . 2 27 .

Von K oen ig swa ld , W . , 1 973 . V e randerungen i n d e r K le in s auge rf auna v on M i t te leu ropa z w i schen C rome ru nd E em ( P le i s to san ) . E i sze i ta l te ru nd G eg enwa r t2 3 -24 , 1 59 167 . V on K oen ig swa ld , W . & F .N ico la ev ich G o len i shev , 1 979 . Am e thod f o r d e te rm in ing g row th r a t es i n c on t inuou s ly g row ing mo la r s . J ourna lo f Mamma logy 6 0 , 3 97 -400 . Voo rh ie s , M . R . , 1 969 . T aphonomy a nd p opu la t ion d ynam i c s o fa n E a r ly P l iocene v er teb ra te f a una , Knox C oun ty , N eb ra ska . U n ive r s i ty o fW yom ing , C on t r ibu t ion s t oG e o l ogy , S pec ia lP ape r s , No . 1 . Weave r , J .L .& S .W . H of fman , 1 979 . D if f e ren t ia l d e tec ta b i l i ty o f r oden t s i n c oyo te s ca t s . Managemen t4 3 , 7 83 -786 .

t o l6g ica s yP reh i s tö r ica s , Memo i r No . 1 3 . V e iga ,J .P . , 1 980 .A l imen ta c ion yr e lac ione st r of ica s e n t r e l a l u chuza c omun ( T y to a l ba ) ye lb uho c h ico ( As io o t u s )e n

Z e i t schr if tf ür S äuge t ie rkund e 6 1 , 2 2 1-

J ou rna l o f W i ld l if e

Web s te r , J .A . , 1 973 . S ea sona l v a r ia t ion i n mamma l c on ten t s o fb a rn o w lc a s t ing s . B i rd S tud y2 0 , 1 85 196 .

l aS i e r ra d eG uada r rama ( E spana ) . A rd eo la 2 5 , 13 -142 . We s ter n , D . , 1 980 . L ink ing t h e e co logy o fp a s t a nd p r e sen t V e iga ,J .P . , 1 986 . I n te rannua l f l uc tua t ion so ft h ree m ic ro t ine

m amma l c ommun i t ie s . I nF o s s i l s i nt h e Mak ing : V er te -

p opu la t ion s i nm ed i te r r eanean e nv i ronmen t s : T he e f f ec t

b ra te T aphonomy a nd P a leoeco logy , e d s . A . K . B e -

o fr a inf a l l . Mamma l ia 5 0 , 14 116 .

h r en smeye r&A . P .H i l l . C h icago : U n ive r s i ty o fC h icago P re s s ,p p .4 1-54 .

V en tu ra , J . , 1 988 . R ev i s ion d e l ac o ro log ia d e A r v ico la t e r r e s t r i s ( L innaeu s , 1 758 ) ( Roden t ia , A rv ico l idae ) e n l a P en in su la I b e r ica . M i sce l lan ia Z oo log ica 1 2 ,3 19 -328 .

W i l l i s , J. C . , 1 973 . AD ic t iona ry o ft h eF low er ing P lan t sa nd F e rn s( 8' e d . ) . C amb r idge : C amb r idge U n ive r s i ty P re s s . 1 88

W i l son , E . 0 . , 1 989 . T h r ea t s t o b i od ive r s i ty . S c ien t if i c Amer i can 3 0 , 1 08 -116 .

Y oungman , P .M . , 1 990 . Mu s t e la l u t reo la . Mamma l ian S pec i e s3 62 , 1 3 .

W i l son , M . V . H . , 1 987 . P r eda t ion a s as ou r ce o ff i sh f o s s i l s i nE ocene l a ke s e d imen t s . P a la io s2 ,4 97 -504 .

Z aba la , J . , 1 983 . P r ime ra c i ta d e A r v i co la s ap idu s ( M i l le r 1 908 , Mamma l ia , R oden t ia ) p a ra l a p rov inc ia d e G u ipüz coa . Mun ibe 3 5 , 1 09 114 .

Wo lf f , R . G . , 1 973 . H yd rodynam ic s o r t ing a nd e co logy o fa P l e i s tocene m amma l ian a s semb lage f r om C a l if o rn ia ( U .S .A . ) . P a laeog eog raphy , P a laeoc l ima to log y , P a l a eoe co logy 1 3 ,9 1 10 1 . Y oungman , P .M . , 1 982 . D i s t r ibu t ion a nd s y s tema t ic s o ft h e E u rop ean m ink Mu s te la l u t r eo la L inna eu s 1 76 1 . A c ta Z oo log i ca F enn i ca 1 66 , 1 48 .

Z aba la , J . , 1 984 . L o s m ic romam if e ro s d e l y ac im ien to p r e h i s tö r ico d e E ka in ( Gu ipüz coa ) . I nE l Y ac im ien to P re h i s tö r i co d e l aC ue va d e E ka in ( Deba , G u ipü zcoa ) , e d s . J .A l tuna & J .M . Me r ino . S an S eba s t ian : S oc iedad d e E s tud io sV a sco s , S e r ie B i ,p p . 3 17 -330 .

1 89