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Social structure of Andean deer (Hippocamelus antisensis) in southern Peru Merkt, Juan R. 1985

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SOCIAL STRUCTURE OF ANDEAN DEER (HI PPOCAMELUS ANTI SENS-IS) IN SOUTHERN PERU by JUAN R. MERKT B . S c , U n i v e r s i t y of B r i t i s h C olumbia, 1978 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES (Department of Zoology) We ac c e p t t h i s t h e s i s as c o n f o r m i n g t o the r e a u i r e d s t a n d a r d THE UNIVERSITY OF BRITISH September 1985 COLUMBIA © Juan R. Me r k t , 1985 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of \^&Ot-0C<f The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date OCJT 7 , ^ f DE-6(3/81) i i ABSTRACT The t a r u c a (Hippocamelus a n t i s e n s i s ) i s the o n l y deer s p e c i e s found permanently i n rugged mountainous h a b i t a t above the t r e e l i n e . I s t u d i e d the s o c i a l o r g a n i z a t i o n of t h i s deer i n r e l a t i o n t o i t s r e p r o d u c t i v e c y c l e and h a b i t a t use i n the h i g h Andes of s o u t h e r n P e r u . Tarucas bred s e a s o n a l l y . Most fawns were obse r v e d towards the end of the r a i n y season between F e b r u a r y and A p r i l . M a t i n g was most common i n June, d u r i n g the d r y season, and a n t l e r - s h e d d i n g i n males o c c u r r e d i n September/October, a t the onset of the r a i n y season. The deer l i v e d i n s o c i a l groups and, u n l i k e most s e a s o n a l l y b r e e d i n g c e r v i d s , formed l a r g e mixed-sex groups n e a r l y a l l y e a r . D u r i n g the b i r t h season, however, a l l pregnant females s e g r e g a t e d t o form female a s s o c i a t i o n s . At t h i s t i m e , a d u l t males were found e q u a l l y i n mixed-sex groups or i n s m a l l a l l - m a l e groups. These groups d i f f e r e d i n t h e i r h a b i t a t use. Female groups used a r e a s of h i g h e r e l e v a t i o n , s t e e p e r s l o p e s , and g r e a t e r r o c k - c o v e r than e i t h e r male or mixed-sex groups. I suggest t h a t s e l e c t i o n of more rugged and c o n c e a l e d h a b i t a t s by l a c t a t i n g females i s p r i m a r i l y an a n t i p r e d a t o r s t r a t e g y t o reduce r i s k of p r e d a t i o n on fawns. Tarucas a r e compared w i t h o t h e r s o c i a l C e r v i d a e and w i t h t h e i r e c o l o g i c a l c o u n t e r p a r t : the mountain C a p r i n a e . The s o c i a l s t r u c t u r e of Hippocamelus resembles t h a t of w i l d g o a t s (Capra spp) and o t h e r C a p r i n a e of s i m i l a r e c o l o g y but i t d i f f e r s from t h a t of w i l d sheep ( O v i s s p p ) . i i i TABLE OF CONTENTS ABSTRACT i i LIST OF TABLES v LIST OF FIGURES v i ACKNOWLEDGEMENTS v i i GENERAL INTRODUCTION 1 CHAPTER 1. REPRODUCTIVE CYCLE AND GROUPING PATTERNS 3 1.1 INTRODUCTION 3 1.2 MATERIALS AND METHODS 5 1.2.1 Study Area 5 1.2.2 Study A n i m a l s and Data A c q u i s i t i o n 12 1.2.3 Data A n a l y s i s 16 1 .3 RESULTS 17 1.3.1 R e p r o d u c t i o n 17 Fawning 17 M a t i n g a c t i v i t y 17 A n t l e r c y c l e 25 1.3.2 Gr o u p i n g P a t t e r n s 28 Mean group s i z e 30 P r o p o r t i o n of t y p e s of groups 33 D i s t r i b u t i o n of deer c l a s s e s among groups 34 S e g r e g a t i o n of pregnant females and a d u l t males .. 37 Group s t a b i l i t y 48 1 .4 DISCUSSION 54 R e p r o d u c t i v e C y c l e 54 G r e g a r i o u s n e s s of Tarucas and T h e i r "Open" H a b i t a t .. 58 CHAPTER 2. HABITAT USE BY MALES AND FEMALES 61 2.1 INTRODUCTION 61 2.2 MATERIALS AND METHODS 65 2.2.1 G r i d System and E n v i r o n m e n t a l F a c t o r s Recorded 65 2.2.2 Deer O b s e r v a t i o n s and Data Recorded 66 2.2.3 Data A n a l y s i s 69 2.3 RESULTS 72 2.3.1 H a b i t a t and R e l a t i o n s between E n v i r o n m e n t a l F a c t o r s 72 G e n e r a l d e s c r i p t i o n of the a r e a 72 C o r r e l a t i o n s between h a b i t a t v a r i a b l e s 75 A l t i t u d i n a l l e v e l s 77 2.3.2 H a b i t a t Use by the Deer 79 G e n e r a l 79 H a b i t a t use comparisons 79 (a) Fawning-season comparisons 82 (b) I n t e r - s e a s o n a l comparisons 84 2.3.3 H a b i t a t S e l e c t i o n , B i a s e s , and C o r r e l a t i o n s ... 87 2.3.4 A n t i p r e d a t o r b e h a v i o r ... 92 2.3.5 C r y p t i c c o l o r a t i o n and h i d i n g b e h a v i o r of fawns 96 2.4 DISCUSSION 98 D i f f e r e n t i a l F o o d - s e l e c t i o n 98 S o c i a l Antagonism 100 P r e d a t o r a v o i d a n c e 101 Comparison w i t h Mountain C a p r i n a e 107 i v APPENDIX A 109 APPENDIX B 113 APPENDIX C 122 REFERENCES 127 V LIST OF TABLES Table 1. Mean number of days w i t h n i g h t f r o s t s 11 Tab l e 2. Mean group s i z e throughout the year 28 Tab l e 3. Group types encountered d u r i n g study 29 Tab l e 4. Number of groups w i t h and w i t h o u t young 30 Tab l e 5. S t r u c t u r e of mixed groups 33 Tab l e 6. S t r u c t u r e of female groups 34 Table 7. Changes i n group s i z e / c o m p o s i t i o n w i t h i n o b s e r v a t i o n p e r i o d s 49 Tab l e 8. Long-term s t a b i l i t y of groups 52 Table 9. S i z e of s u b u n i t s i n main study a r e a 67 Table 10. Number of v i s i t s t o and number of groups i n each s u b u n i t i n the main study area 68 Table 11. H a b i t a t d e s c r i p t i o n of main study a r e a 73 Table 12. P a r t i a l c o r r e l a t i o n s between h a b i t a t v a r i a b l e s . . 76 Tab l e 13. The f i v e a l t i t u d i n a l l e v e l s i n the main study a r e a 80 Tab l e 14. Summary of h a b i t a t comparisons 88 Table 15. H a b i t a t v a r i a b l e f r e q u e n c i e s d u r i n g the fawning season 90 Tab l e 16. Escape cover 94 Tab l e A1. P l a n t s p e c i e s i n l o n g - g r a s s v e g e t a t i o n - t y p e 110 Table A2. P l a n t s p e c i e s i n s h o r t - g r a s s v e g e t a t i o n - t y p e . ...111 Table A3. P l a n t s p e c i e s i n marsh v e g e t a t i o n - t y p e 112 Tab l e B1. V e r t i c a l d i s t r i b u t i o n : f emale, male, and mixed groups 114 Table B2. Changes i n v e r t i c a l d i s t r i b u t i o n w i t h time of day 115 Table B3. S l o p e : female, male and mixed groups 116 Table B4. Rock-cover: female, male, and mixed groups 116 Table B5. T o p o g r a p h i c a l r e l i e f : f emale, male, and mixed groups 117 Table B6. T e r r a c e s : female, male, and mixed groups 117 Table B7. V e g e t a t i o n t y p e : female, male, and mixed groups. 118 Tab l e B8. D i s t a n c e t o water: f e m a l e , male, and mixed groups 118 Table B9. H a b i t a t comparisons between groups d u r i n g fawning season 119 Tab l e B10. I n t e r - s e a s o n a l h a b i t a t comparisons. 1 120 Table B1 1 . I n t e r - s e a s o n a l h a b i t a t comparisons.2 121 Tab l e C1. V a r i a b l e s used f o r d i s c r i m i n a n t a n a l y s e s .123 Tab l e C2. W i t h i n - s e a s o n r e s u l t s from d i s c r i m i n a n t a n a l y s e s . 124 T a b l e C3. Between-season r e s u l t s from d i s c r i m i n a n t a n a l y s e s 125 LIST OF FIGURES F i g u r e 1 . Map of Pe r u and the study a r e a 6 F i g u r e 2. Mean monthly p r e c i p i t a t i o n , and minimum and maximum t e m p e r a t u r e s 9 F i g u r e 3. F a c i a l markings of e i g h t known i n d i v i d u a l s 14 F i g u r e 4. R e l a t i v e number of young fawns 18 F i g u r e 5. Number of a d u l t male/female i n t e r a c t i o n s 21 F i g u r e 6. P e r c e n t a g e of i n t e r a c t i o n s l e a d i n g t o mounting .. 23 F i g u r e 7. A n t l e r c y c l e of males 26 F i g u r e 8. Mean s i z e of deer groups 31 F i g u r e 9. P r o p o r t i o n of d i f f e r e n t t y p e s of groups: 35 F i g u r e 10. R e l a t i v e number of a d u l t males i n mixed-sex and male groups 38 F i g u r e 11. R e l a t i v e number of a d u l t females i n mixed-sex and female groups 40 F i g u r e 12. P e r c e n t a g e of a d u l t males and females o b s e r v e d as s o l i t a r y i n d i v i d u a l s 42 F i g u r e 13. R e l a t i v e number of y e a r l i n g s i n mixed-sex, male, and female groups 44 F i g u r e 14. R e l a t i v e number of fawns i n female and mixed-sex groups 46 F i g u r e 15. L o c a t i o n s of known a d u l t males and females 50 v i i ACKNOWLEDGEMENTS I wi s h t o thank my a d v i s o r Dr. W i l l i a m E. Rees f o r h i s s u p p o r t , a d v i c e , and encouragement throughout the s t u d y , and D r s . F r e d L. B u n n e l l , A.R.E. S i n c l a i r , and J.N.M. Smith f o r a d v i c e and h e l p f u l s u g g e s t i o n s d u r i n g the w r i t e - u p s t a g e . I e x t e n d my g r a t i t u d e t o Dr. C a r l W a l t e r s f o r m a t e r i a l s u p p o r t d u r i n g my s t a y at the I.A.R.E. Many p e o p l e and s e v e r a l i n s t i t u t i o n s made t h i s study p o s s i b l e . I am d e e p l y i n d e b t e d t o Dr. A l b e r t o S a t o , D i r e c t o r , I n s t i t u t o V e t e r i n a r i o de I n v e s t i g a c i o n e s T r o p i c a l e s y de A l t u r a ( I . V . I . T . A . ) , U n i v e r s i d a d N a c i o n a l Mayor de San Marcos, Lima, f o r k i n d l y a l l o w i n g me t o work a t La Raya, Cusco. I am most t h a n k f u l t o Dr. J u l i o Sumar, C o o r d i n a d o r T e c n i c o , I.V.I.T.A., and a l l h i s c o l l e a g u e s a t La Raya f o r g i v i n g me t h e i r f r i e n d s h i p and i n v a l u a b l e m a t e r i a l s u p p o r t . I n g . Juan A l p a c a , A d m i n i s t r a d o r of La Raya, was a l s o most h e l p f u l . Renato M a r i n i n t r o d u c e d me t o the s p e c t a c u l a r mountains of La Raya, and p r o v i d e d h e l p f u l i n f o r m a t i o n on t a r u c a s i n the Cusco a r e a . Domingo J a r a t a u g h t me where t o l o o k f o r the e l u s i v e deer and, a l o n g w i t h o t h e r p e r s o n n e l from I.V.I.T.A. and the M i n i s t e r i o de A g r i c u l t u r a , a s s i s t e d me on many o c c a s i o n s . Dr. M a r i o T a p i a and L u c r e c i a A g u i r r e , from U n i v e r s i d a d de Cusco, k i n d l y i d e n t i f i e d many p l a n t s p e c i e s from the study a r e a . For f i e l d s u p p o r t d u r i n g a p r e l i m i n a r y study a t the V i c u n a Reserve of Pampa G a l e r a s , Ayacucho, I thank Dr. A n t o n i o B r a c k , former D i r e c t o r of the V i c u n a P r o j e c t , and Dr. Rudolph v i i i Hofmann, former head of the German M i s s i o n f o r T e c h n i c a l C o o p e r a t i o n a t the R e s e r v e . I a l s o thank Dr. Marc D o u r o j e a n n i , former D i r e c t o r of the D i r e c c i o n G e n e r a l F o r e s t a l y de Fauna, M i n i s t e r i o de A g r i c u l t u r a , Lima, f o r g i v i n g me moral support and a d v i c e d u r i n g i n i t i a l s t a g e s of the s t u d y . My deepest thanks go t o my r e l a t i v e s i n Lima and Cusco, e s p e c i a l l y W i l l i a m and L u i s a L e o n a r d , ' f o r t h e i r more than generous h o s p i t a l i t y d u r i n g my s t a y i n P e r u . I am f o r e v e r g r a t e f u l t o Peggy S t e r n f o r her f r i e n d s h i p , encouragement, and eagerness t o h e l p me many times d u r i n g my s t u d y . L a s t but not l e a s t , I thank my f r i e n d s and f e l l o w g r aduate s t u d e n t s a t t h e I n s t i t u t e of Animal Resource E c o l o g y , f o r making my time spent i n Vancouver most e n j o y a b l e , f o r t h e i r p a t i e n c e i n l i s t e n i n g t o the t a r u c a s t o r y over and o v e r , and f o r h e l p d u r i n g the f i n a l h e c t i c s t a g e s of t h e s i s p r e p a r a t i o n . I am e s p e c i a l l y t h a n k f u l t o C a r l o s G a l i n d o , Kaaren L e w i s , Laura Jamieson, Marco R o d r i g u e z , A l i s t a i r B l a c h f o r d , John F r y x e l l , H o l l y D u b l i n , P e t e r A r c e s e , A l i s o n S o m e r v i l l e , and Joyce Andrew. T h i s work was p a r t i a l l y funded by the F a c u l t y of Graduate S t u d i e s , S c h o o l of Community and R e g i o n a l P l a n n i n g , and I n s t i t u t e of Animal Resource E c o l o g y a t t h e U n i v e r s i t y of B r i t i s h C o lumbia, B.C., Canada. 1 GENERAL INTRODUCTION Most of our knowledge on r e l a t i o n s h i p s between s o c i a l o r g a n i z a t i o n and h a b i t a t among the C e r v i d a e comes from s t u d i e s of s p e c i e s i n temperate and s u b - t r o p i c a l h a b i t a t s i n the N o r t h e r n hemisphere ( C l u t t o n - B r o c k et a l . , 1982; E i s e n b e r g , 1981; H i r t h , 1977; LaGory, 1985; Maublanc e t a l . , 1985; S c h a l l e r , 1967; T a k a t s u k i , 1983). An i m p o r t a n t group of n e o t r o p i c a l s p e c i e s has been l a r g e l y n e g l e c t e d (Roe, 1974). Of the 16 genera of c e r v i d s i n the w o r l d , 6 (11 e x t a n t s p e c i e s ) a re found i n South America ( H e r s h k o v i t z , 1982). N e o t r o p i c a l deer r e p r e s e n t one of the most d i v e r s i f i e d a s s e m b l i e s of c e r v i d s , both i n terms of e x t e r n a l c h a r a c t e r i s t i c s and range of h a b i t a t s o c c u p i e d . I n f o r m a t i o n on t h e i r b e h a v i o r and e c o l o g y s h o u l d t h e r e f o r e a i d our g e n e r a l u n d e r s t a n d i n g of s o c i a l systems among the C e r v i d a e . T h i s study f o c u s e s on a n e o t r o p i c a l s p e c i e s , the Andean deer or t a r u c a (Hippocamelus a n t i s e n s i s ) . The t a r u c a i s a medium-sized deer t h a t i n h a b i t s the h i g h Andes between n o r t h e r n A r g e n t i n a and C h i l e and n o r t h e r n P e r u ( C a b r e r a , 1961; Grimwood, 1969; IUCN, 1982). D e s c r i p t i o n s of i t s e x t e r n a l c h a r a c t e r i s t i c s a re found i n F r a d r i c h (1978), Pearson (1951), Roe (1974), Roe and Rees (1976), and Whitehead (1972). Found a t e l e v a t i o n s of up t o 5,200 m, the t a r u c a i s the o n l y deer t h a t l i v e s p ermanently i n h i g h mountainous h a b i t a t above the t r e e l i n e . I t s e c o l o g i c a l range does not o v e r l a p t h a t of o t h e r c e r v i d s . I t s c l o s e s t r e l a t i v e , the huemul, Hippocamelus  b i s u l c u s , i s found a t lower e l e v a t i o n s i n c e n t r a l C h i l e and 2 n e i g h b o r i n g a r e a s of A r g e n t i n a ( P o v i l i t i s , 1978). Thus, H. b i s u l c u s and H. a n t i s e n s i s have a l l o p a t r i c d i s t r i b u t i o n s . Only i n some a r e a s does the lower l i m i t of t a r u c a h a b i t a t appear t o be c o n t i g u o u s w i t h t h a t of w h i t e - t a i l e d d e e r , O d o c o i l e u s  v i r g i n i a n u s ( J u n g i u s , 1974; R. M a r i n , p e r s . comm.). The t a r u c a i s thus e c o l o g i c a l l y unique among c e r v i d s , i n h a b i t i n g a n i c h e u s u a l l y o c c u p i e d by mountain sheep and goats i n o t h e r p a r t s of the w o r l d ( G e i s t , 1971; S c h a l l e r , 1977). I t s unusual h a b i t a t may r e f l e c t t he f a c t t h a t mountain C a p r i n a e have never become e s t a b l i s h e d i n the n e o t r o p i c s . Indeed, t a r u c a s seem s u p e r b l y adapted t o rugged mountainous t e r r a i n . The a n i m a l s a re s t o c k i l y b u i l t and have s h o r t l e g s and v e r y s h o r t hooves ( F r a d r i c h , 1978; Roe and Rees, 1976). Some o b s e r v e r s have noted the d e e r ' s a g i l i t y on r o c k y t e r r a i n (Roe and Rees, 1976) and o t h e r s have remarked on the p h y s i c a l resemblance of Hippocamelus t o w i l d mountain sheep ( F r a d r i c h , 1978). These c h a r a c t e r i s t i c s s u g g e s t t h a t Hippocamelus i s the e c o l o g i c a l c o u n t e r p a r t of C a p r i n a e . D e s p i t e the unusual n i c h e of the t a r u c a , no d e t a i l e d s t u d i e s of the s o c i a l o r g a n i z a t i o n of the s p e c i e s have ever been made. In t h i s study I d e s c r i b e some a s p e c t s of the s o c i a l o r g a n i z a t i o n of a p o p u l a t i o n of Hippocamelus a n t i s e n s i s i n the Andes of s o u t h e r n Peru and examine e c o l o g i c a l f a c t o r s i n f l u e n c i n g i t s g r o u p i n g p a t t e r n s . I c o n s i d e r t a r u c a group dynamics i n r e l a t i o n t o (1) r e p r o d u c t i v e phenology (Chapter 1 ) , and (2) h a b i t a t use (Chapter 2 ) . 3 CHAPTER 1. REPRODUCTIVE CYCLE AND GROUPING PATTERNS J_.j_ INTRODUCTION T h i s s t u d y d e a l s w i t h some a s p e c t s of the s o c i a l o r g a n i z a t i o n of Hippocamelus a n t i s e n s i s . For the purposes of t h i s s t u d y , the term s o c i a l o r g a n i z a t i o n r e f e r s t o the manner i n which members of a s i n g l e - s p e c i e s p o p u l a t i o n a r e p o s i t i o n e d i n time and space i n r e l a t i o n t o one ano t h e r ( H i r t h , 1977; L e u t h o l d , 1977). Thus, s o c i a l o r g a n i z a t i o n i n c l u d e s group s i z e s , sex-age group c o m p o s i t i o n s (group t y p e s ) , and t e m p o r a l and s p a t i a l arrangements among the d i f f e r e n t i n d i v i d u a l s or groups. I s p e c i f i c a l l y do not emphasize the b e h a v i o r a l mechanisms u n d e r l y i n g s o c i a l o r g a n i z a t i o n . T h e r e f o r e , I w i l l be concerned w i t h the consequences of s o c i a l or i n d i v i d u a l - h a b i t a t i n t e r a c t i o n s . Among u n g u l a t e s , s e a s o n a l changes i n s o c i a l o r g a n i z a t i o n a r e g r e a t l y i n f l u e n c e d by the r e p r o d u c t i v e c y c l e , which i n t u r n i s d e t e r m i n e d by s e a s o n a l f l u c t u a t i o n s of the environment ( B u n n e l l , 1982; C l u t t o n - B r o c k e t a l . 1982; H i r t h , 1977; L e u t h o l d , 1977; N i e v e r g e l t , 1974; S c h a l l e r , 1977; Thompson and T u r n e r , 1982). A l t h o u g h t h e r e i s i n f o r m a t i o n on s e a s o n a l f l u c t u a t i o n s of the environment i n n e o t r o p i c a l h i g h l a n d s ( C a r p e n t e r , 1976; F r a n k l i n , 1983; O.N.E.R.N., 1965; Pe a r s o n , 1951; T r o l l , 1968), l i t t l e i s known about ann u a l r e p r o d u c t i v e or s o c i a l p a t t e r n s of Hippocamelus a n t i s e n s i s (Roe, 1974; Roe and Rees, 1976). 4 In t h i s c h a p t e r I f i r s t p r e s e n t b a s i c i n f o r m a t i o n on t h e a n n u a l r e p r o d u c t i v e c y c l e of the deer. Three a s p e c t s a r e examined: (1) the time and s p r e a d of the b i r t h season; (2) the m a t i n g p e r i o d ; and (3) the a n t l e r c y c l e i n males. Second, I examine the g r o u p i n g p a t t e r n s of Hippocamelus a s s o c i a t e d w i t h the r e p r o d u c t i v e c y c l e . Four g r o u p i n g f e a t u r e s a r e c o n s i d e r e d : (1) group s i z e s ; (2) t y p e s of g r o u p i n g ; (3) d i s t r i b u t i o n of sex-age c l a s s e s among the d i f f e r e n t groups; and (4) s t a b i l i t y of the groups e n c o u n t e r e d d u r i n g the s t u d y . a 5 1.2 MATERIALS AND METHODS _J_.2.J_ Study Area T h i s study was c a r r i e d out a t La Raya (about 14° 30'S, 70° 58'W) i n s o u t h e r n P e r u ( F i g . 1 ) . T h i s s i t e i s a c c e s s i b l e , has a r e l a t i v e l y l a r g e deer p o p u l a t i o n , and has a l a r g e and w e l l e quipped r e s e a r c h and range management s t a t i o n , the C e n t r o N a c i o n a l de Camelidos Sudamericanos. In c o n t r a s t t o o t h e r a r e a s t h a t I v i s i t e d , La Raya shows almost no si.gns of o v e r g r a z i n g by d o m e s t i c s t o c k . My o b s e r v a t i o n s were made w i t h i n the s t a t i o n ' s 120 km 2 a r e a , on a main study a r e a of 79 km 2 . The r e g i o n i s p a r t of the Puna, a t r e e l e s s , h i g h Andean f o r m a t i o n dominated by p e r e n n i a l bunch g r a s s e s . Here, the E a s t e r n and C e n t r a l C o r d i l l e r a s of the Andes come t o g e t h e r , f o r m i n g a knot a t the d i v i d e between the Amazon and Lake T i t i c a c a B a s i n d r a i n a g e s . The t e r r a i n i s v e r y rugged w i t h s t e e p - s i d e d mountains s u r r o u n d i n g a broad main v a l l e y and s m a l l e r s i d e v a l l e y s . Rocky o u t c r o p s or permanent i c e f i e l d s c over most of the upper s l o p e s and mountain t o p s . Mean e l e v a t i o n of the main study a r e a i s 4,630 m and w i t h i n i t s a l t i t u d i n a l range (4,070 t o 5,470 m) t h r e e e c o l o g i c a l zones o c c u r : s u b a l p i n e meadow, a l p i n e t u n d r a , and g l a c i a l (O.N.E.R.N., 1965). The v e g e t a t i o n c o n s i s t s m a i n l y of t a l l bunch-grasses ( F e s t u c a , S t i p a , and C a l a m a g r o s t i s spp) which g i v e cover t o s m a l l e r g r a m i n o i d s , f o r b s , mosses, and l i c h e n s . Marshy v e g e t a t i o n , dominated by D i s t i c h i a muscoides , i s common a l o n g the m o i s t v a l l e y bottoms and on some s l o p e s where water f i l t e r s 6 F i g u r e 1. Map of P e r u and the s t u d y a r e a a t La Raya. 8 out from the m e l t i n g g l a c i e r s . S h o r t shrubs ( S e n e c i o , B a c c h a r i s spp) a r e found o n l y i n a few s i t e s a l o n g the upper l i m i t of the v e g e t a t i o n . A more d e t a i l e d d e s c r i p t i o n of the st u d y a r e a i s g i v e n i n S e c t i o n 2.3.1 where t h e r e s u l t s of h a b i t a t a n a l y s i s a r e p r e s e n t e d . The c l i m a t e i n t h e h i g h l a n d s of s o u t h e r n P e r u has been d e s c r i b e d by C a r p e n t e r (1976), F r a n k l i n (1983), O.N.E.R.N. (1965), Pearson (1951), and T r o l l (1968). Annual p r e c i p i t a t i o n of about 900 mm i s s t r o n g l y s e a s o n a l , w i t h a wet season i n October t h r o u g h A p r i l and a d r y season i n May t h r o u g h September ( F i g . 2 ) . Most of t h i s p r e c i p i t a t i o n i s i n the form of r a i n or h a i l (below 4,800 m) and snow or h a i l (above 4,800 m). Both the a l t i t u d e and the p r e c i p i t a t i o n regime g r e a t l y i n f l u e n c e temperature p a t t e r n s i n the s e t r o p i c a l h i g h l a n d s . Mean annua l t e m p e r a t u r e ranges from 6° C a t 4,100 m t o 0° C above 4,800 m (O.N.E.R.N., 1965) but f l u c t u a t e s l i t t l e w i t h season. Large d i u r n a l f l u c t u a t i o n s do o c c u r . The i n t e n s e s o l a r r a d i a t i o n a t these e l e v a t i o n s r e s u l t s i n r e l a t i v e l y warm days and c o l d n i g h t s . T h i s i s e s p e c i a l l y t r u e d u r i n g the dry season from May t h r o u g h September, a time of c l e a r s k i e s and heavy n i g h t f r o s t s (Table 1 ) . D u r i n g the r a i n y season, d i u r n a l f l u c t u a t i o n s i n t e m p e r a t u r e a r e l e s s marked. N i g h t t e m p e r a t u r e s a r e u s u a l l y above f r e e z i n g , but days t e n d t o be c o l d because of o v e r c a s t s k i e s . F i g u r e 2 shows mean monthly maximum and minimum t e m p e r a t u r e s a t La Raya. 9 F i g u r e 2. Mean monthly p r e c i p i t a t i o n , and minimum and maximum t e m p e r a t u r e s a t La Raya. Means were e s t i m a t e d from 1980 and 1981 d a i l y r e c o r d s . P r e c i p i t a t i o n i s shown by b a r s . 10 300-1 g 200 -c o CO •4—' Q. O 2 100-Q_ +2 0 . 0 / ° \ ^ . o -o -o -+1 5 -4-1 0 \ O m e a n max t e m p • m e a n min t e m p -+ 5 H 0 3 CD —s £> «-t-C - s CD \ / I - o o - - 5 J F M A M J J A S O N D 11 The main human a c t i v i t y a t La Raya i n v o l v e s r e s e a r c h Table 1. Mean number of days w i t h f r e e z i n g n i g h t t e m p e r a t u r e s (<= 0 C) a t La Raya, P e r u . Means were c a l c u l a t e d from 1980 and 1981 d a i l y r e c o r d s . MONTH NUMBER OF DAYS WITH NIGHT FROSTS January 5 F e b r u a r y 4 March 5 A p r i l 26 May 31 June 30 J u l y 31 August 30 September 23 October 1 7 November 1 3 December 1 6 and management of the a l p a c a (Lama pacos) . Some 5,000 a l p a c a s a r e m a i n t a i n e d i n herds a v e r a g i n g 340 a n i m a l s . The s t a t i o n a l s o s u p p o r t s about 400 l l a m a s (Lama glama) and a few d o m e s t i c sheep, c a t t l e , and h o r s e s . Most of the h i g h l a n d p a s t u r e s of Peru a r e used f o r e x t e n s i v e a l p a c a g r a z i n g , but u n l i k e La Raya, th e s e l a n d s are community owned and i n some a r e a s domestic sheep 12 outnumber a l p a c a s . B e s i d e Hippocamelus , the o n l y o t h e r w i l d u n g u l a t e i n the area i s the v i c u n a (Lama v i c u g n a ) . P o t e n t i a l p r e d a t o r s of t a r u c a s i n c l u d e the cougar ( F e l i s c o n c o l o r ) , s e m i - f e r a l dogs, and Andean f o x e s (Dusicyon c u l p a e u s ) . More w i l l be s a i d about p r e d a t o r s i n Chapter 2. J..2.2 Study Animals and Data A c q u i s i t i o n Most f i e l d work was done from November 1980 t o December 1981, a l t h o u g h some p r e l i m i n a r y o b s e r v a t i o n s were made between August and October 1980. I c o v e r e d the ar e a on horseback, and observ e d t a r u c a s m a i n l y from r e g u l a r s e a r c h paths and l o o k - o u t p o i n t s . When weather and time p e r m i t t e d , groups were obser v e d f o r as l o n g as they remained i n s i g h t w i t h the a i d of b i n o c u l a r s and a 20-45X s p o t t i n g scope. A l l o b s e r v a t i o n s were made i n d a y l i g h t between 0500 and 1800 h. Tarucas were observed a t d i s t a n c e s of 30 t o 1,500 m, though most o b s e r v a t i o n s were w i t h i n a range of 100 - 1,000 m. An i m a l s were c l a s s i f i e d i n t o 5 broad age/sex c l a s s e s : a d u l t males, a d u l t f e m a l e s , y e a r l i n g males, y e a r l i n g f e m a l e s , and fawns. A d u l t males were d i s t i n g u i s h e d by t h e i r l a r g e body s i z e , l a r g e a n t l e r s ( r e a r t i n e up t o t w i c e the ear l e n g t h ) , l o n g r o s t r u m , and t h i c k neck. A d u l t females were s m a l l e r than a d u l t males, l a c k e d a n t l e r s , and had l o n g r o s t r a but t h i n necks. Y e a r l i n g s , w i t h s h o r t r o s t r a and t h i n n e cks, were s m a l l e r than a d u l t males and f e m a l e s . Though most y e a r l i n g males had o n l y a 13 p a i r of s h o r t s p i k e s , some had s m a l l f o r k e d a n t l e r s . Fawns were d i s t i n g u i s h e d by t h e i r s m a l l s i z e and t h e i r d a r k - g r e y c o a t s . About a month a f t e r they were f i r s t o b s e r v e d , fawns s t a r t e d t o a c q u i r e the sandy-brown p e l a g e c h a r a c t e r i s t i c of o l d e r a n i m a l s . I was e v e n t u a l l y a b l e t o i d e n t i f y s p e c i f i c i n d i v i d u a l s by the markings on t h e i r heads, and i n the case of males, a l s o by the s i z e and shape of t h e i r a n t l e r s . F i g u r e 3 i l l u s t r a t e s t he f a c i a l p a t t e r n s and a n t l e r s of s e l e c t e d known i n d i v i d u a l s . Once a group was l o c a t e d , I r e c o r d e d the presence of any known i n d i v i d u a l , the number and c l a s s e s of deer w i t h i n the group, t h e i r a c t i v i t i e s a t 5-min i n t e r v a l s , any b e h a v i o r a l i n t e r a c t i o n between two a n i m a l s , and a range of e n v i r o n m e n t a l v a r i a b l e s a t the b e g i n n i n g , end, and every hour d u r i n g the o b s e r v a t i o n p e r i o d . A d e t a i l e d account of h a b i t a t d a t a r e c o r d e d i s g i v e n i n S e c t i o n 2.2. Throughout the s t u d y I conducted 175 o b s e r v a t i o n p e r i o d s r a n g i n g from l e s s than 5-min t o 8-hrs i n d u r a t i o n . On average o b s e r v a t i o n s l a s t e d a p p r o x i m a t e l y 1 hour. S i n c e deer d e n s i t y was r e l a t i v e l y low (about 1.2 animals/km 2) groups were found w i d e l y spaced, and i t was r a r e t o have more than one group i n s i g h t . However, i f a group b e i n g m o n i t o r e d changed i n s i z e and/or c o m p o s i t i o n (because of a n i m a l s j o i n i n g or l e a v i n g the group) i t was t r e a t e d as a new group. Thus, I had 208 group s i g h t i n g s which i n c l u d e d 310 o b s e r v a t i o n s of a d u l t males, 500 of a d u l t f e m a l e s , 264 of y e a r l i n g s , and 112 of fawns. In a d d i t i o n , I o b s e r v e d 19 groups whose age/sex c o m p o s i t i o n c o u l d not be d e t e r m i n e d . 14 F i g u r e 3. F a c i a l markings of e i g h t known i n d i v i d u a l s . A l s o note a n t l e r shapes i n a d u l t males. 16 J_.2.3 Data A n a l y s i s Data from 1980 and 1981 showed s i m i l a r p a t t e r n s and, u n l e s s o t h e r w i s e s p e c i f i e d , have been combined. Sample s i z e s were s m a l l i n some months, and a l l group d a t a have been p o o l e d i n t o 2-month p e r i o d s . From J u l y 1981 on, I c o u l d not d i s t i n g u i s h y e a r l i n g females born i n 1980 from a d u l t f e m a l e s . Y e a r l i n g males, however, remained d i s t i n c t u n t i l the end of the study i n December 1981. Fawns born i n e a r l y 1981 were c o n s i d e r e d y e a r l i n g s from September 1981 o r , i f males, when they grew t h e i r f i r s t s e t of a n t l e r s . 17 1.3 RESULTS J_.3._1_ R e p r o d u c t i o n Here, I c o n s i d e r t h r e e e v e n t s r e l a t e d t o r e p r o d u c t i o n : 1) fawning ( b i r t h s ) , d e t e r m i n e d by the r e l a t i v e number of fawn s i g h t i n g s ; 2) mating a c t i v i t y , d e f i n e d by the f r e q u e n c y and n a t u r e of male/female i n t e r a c t i o n s ; and 3) the a n t l e r c y c l e i n males. Fawn i n g To e s t i m a t e the r e l a t i v e number of fawns born per month I used the number of fawn s i g h t i n g s / 1 0 0 a d u l t f e m a l e s . I d i s t i n g u i s h e d s m a l l fawns from o l d e r fawns by p e l a g e c o l o r and s i z e r e l a t i v e t o t h e i r mothers'. In 1981, fawns appeared between F e b r u a r y and A p r i l , w i t h a s h a r p peak i n March, l a t e i n the r a i n y season ( F i g . 4 ) . Of a l l s m a l l fawn s i g h t i n g s , 83% (29/35) o c c u r r e d between 20 F e b r u a r y and 31 March. These " b i r t h " d a t e s , however, a r e l i k e l y t o be b i a s e d , and peak number of b i r t h s may have o c c u r r e d e a r l i e r i n F e b r u a r y (see b e l o w ) . Only 2 fawns were o b s e r v e d o u t s i d e t h e main fawning season i n November 1980. M a t i n g a c t i v i t y In the a n a l y s i s of male/female i n t e r a c t i o n s , o n l y o b s e r v a t i o n s of 5-min or l o n g e r were used. I d e f i n e d a 18 F i g u r e 4. R e l a t i v e number of young fawns e x p r e s s e d as fawn s i g h t i n g s per 100 a d u l t f e m a l e s . November s i g h t i n g s c o r r e s p o n d t o 1980 o n l y , the remainder t o 1981. 1 OCH CO E d) 80 -L L 6 0 -4 0 -2 0 -r ~ i i i i ~i—1—i 1 1 1 1 1——i——r~ J F M A M J J A S O N D 20 male/female i n t e r a c t i o n as any sequence of b e h a v i o r a l d i s p l a y s or c o n t a c t b e h a v i o r s between a d u l t s of d i f f e r e n t sex. They u s u a l l y were i n i t i a t e d by a male a p p r o a c h i n g a female and were o f t e n ended by the female moving away or by the male mounting t h e female. I r e c o r d e d 70 i n t e r a c t i o n s d u r i n g 64 o b s e r v a t i o n p e r i o d s t o t a l l i n g 107 h 51 min. The h i g h e s t number of i n t e r a c t i o n s took p l a c e i n June i n the m i d d l e of the dry season ( F i g . 5 ) . The number of i n t e r a c t i o n s was lowest i n March and A p r i l when the sexes were most s e g r e g a t e d . Male/female i n t e r a c t i o n s were a l s o r a r e i n October and November when males had dropped t h e i r a n t l e r s (see b e l o w ) . The p r o p o r t i o n of male/female i n t e r a c t i o n s t h a t l e d t o mounting was a l s o h i g h e s t i n June ( F i g . 6 ) . Most of the time I c o u l d not d i s t i n g u i s h between s u c c e s s f u l and a t t e m p t e d mounts. The former might a l s o have been r e l a t i v e l y more f r e q u e n t i n June. I d i d not observe any male/female i n t e r a c t i o n s i n J u l y . However, none of my o b s e r v a t i o n s of mixed-sex groups d u r i n g t h i s month l a s t e d more than 5 m i n u t e s . Thus b r e e d i n g may have c o n t i n u e d i n t o J u l y . E x t r a p o l a t i o n from peak of mating a c t i v i t y i n June t o peak of r e l a t i v e number of s i g h t i n g s of s m a l l - f a w n s i n March g i v e s an approximate g e s t a t i o n p e r i o d of 270 days. T h i s f i g u r e i s h i g h e r than t h a t f o r any o t h e r c e r v i d s p e c i e s ( E i s e n b e r g , 1981: Appendix 4 ) . The most l i k e l y e x p l a n a t i o n i s t h a t b i r t h s a c t u a l l y o c c u r r e d b e f o r e t a r u c a fawns were f i r s t o b s e r v e d . In most c e r v i d s , young remain h i d d e n f o r some time a f t e r b i r t h (De 21 F i g u r e 5. Number of a d u l t male/female i n t e r a c t i o n s /hr of o b s e r v . Only obs. of 5-min or l o n g e r a r e i n c l u d e d . R a t i o s r e p r e s e n t num. of i n t e r a c t i o n s / t o t a l monthly obs. time ( m i n . ) . No d a t a were o b t a i n e d i n J u l y and no i n t e r a c t i o n s o c c u r r e d i n November. 1980/1981 data are combined. Number of Interactions / Hour 23 F i g u r e 6. P e r c e n t a g e of i n t e r a c t i o n s l e a d i n g t o mounting of a d u l t females by a d u l t males. No d a t a were o b t a i n e d i n J u l y and no i n t e r a c t i o n s o c c u r r e d i n November. 1980/1981 d a t a have been combined. H Int. l ead ing to mount ing D Int. not leading to mounting 1 0 0 | 1 j 1 1 1 1 | 8 0 -J F M A M J J A S O N D 25 Vos, 1967; L e n t , 1974). From e v i d e n c e p r e s e n t e d l a t e r ( s e c t i o n s 1.3.2 and 2.3.5), I e s t i m a t e peak number of b i r t h s i n F e b r u a r y , w i t h the b i r t h season e x t e n d i n g from January t o March. T h i s c o r r e c t i o n y i e l d s an approximate g e s t a t i o n p e r i o d of 240 days, a f i g u r e more c o m p a t i b l e w i t h those known f o r o t h e r deer ( E i s e n b e r g , 1981). A n t l e r c y c l e Data on a n t l e r c o n d i t i o n was based on 453 s i g h t i n g s of males ( a d u l t s and y e a r l i n g s p o o l e d t o g e t h e r ) . The a n t l e r c y c l e was h i g h l y s y n c h r o n i z e d and s e a s o n a l ( F i g . 7 ) . The p r o p o r t i o n of males w i t h c l e a n a n t l e r s i n c r e a s e d from 30% i n J a n u a r y t o 100% i n May. A l l a d u l t males, however, had shed t h e i r v e l v e t as e a r l y as F e b r u a r y . D u r i n g the d r y season, when m a t i n g a c t i v i t y was most i n t e n s e ( F i g s . 5 and 6 ) , a l l males had c l e a n a n t l e r s ( F i g . 7 ) . By September males were d r o p p i n g a n t l e r s , and some new a n t l e r s were i n v e l v e t . By December a l l males were i n v e l v e t ( F i g . 7 ) . In summary, 94% of b i r t h s o c c u r r e d towards the end of the r a i n y season; mating took p l a c e d u r i n g the d r i e s t months; and a n t l e r s h e d d i n g i n males was o b s e r v e d d u r i n g t h e onset of the r a i n y season. T h e r e f o r e , the r e p r o d u c t i v e c y c l e i n Hippocamelus a n t i s e n s i s i s markedly a s s o c i a t e d w i t h s e a s o n a l c l i m a t i c p a t t e r n s . 26 F i g u r e 7. A n t l e r c y c l e of males. P e r c e n t a g e of males ( a d u l t s and y e a r l i n g s p o o l e d ) w i t h no a n t l e r s (NA), a n t l e r s i n v e l v e t ( V ) , or f r e e of v e l v e t (NV). N= number of male s i g h t i n g s . 1980/1981 d a t a have been combined. J F M A M J J A S O N D 54 43 58 20 89 37 4 32 17 51 35 13 28 _1_._3.2_ Grouping P a t t e r n s Mean group s i z e f o r the e n t i r e s t u d y was 6.4 ( + 0.36 SE) d e e r , and groups ranged from 1 t o 31 a n i m a l s . Mean s i z e v a r i e d l i t t l e s e a s o n a l l y but was s m a l l e r (5.0-5.3) j u s t p r i o r t o and d u r i n g fawning ( J a n u a r y - A p r i l ) and l a r g e r (7.3-7.6) d u r i n g the r e s t of the year (Table 2 ) . Ta b l e 2. Mean group s i z e throughout the y e a r . Data from 1980 and 1981 have been combined and p o o l e d i n t o 2-month p e r i o d s . MONTH MEAN SIZE ( +.SE) No. GROUPS JAN/FEB 5.3 ( + 0.60) 61 MAR/APR 5.0 ( + 0.59) 43 MAY/JUN 7.4 ( + 0.89) 57 JUL/AUG 7.3 ( + 1 .50) 20 SEP/OCT 7.3 ( + 1.01) 21 NOV/DEC 7.6 ( + 1.16) 26 Groups were c l a s s e d i n t o 3 group t y p e s of v a r i o u s age/sex c o m p o s i t i o n s (Table 3 ) . Mixed groups c o n t a i n e d both a d u l t males and a d u l t females and a l s o i n c l u d e d y e a r l i n g s and/or fawns most of the time (Table 4 ) . Male groups c o n s i s t e d of one or more a d u l t males and r a r e l y i n c l u d e d y e a r l i n g s of e i t h e r sex. Fawns never o c c u r r e d i n male groups ( T a b l e 4 ) . Female groups 29 c o n t a i n e d one or more a d u l t females and o f t e n i n c l u d e d fawns and/or y e a r l i n g s ( T a b l e 4 ) . Y e a r l i n g s never a s s o c i a t e d t o form a l l - y e a r l i n g groups. Lone y e a r l i n g s were o b s e r v e d on o n l y 4 o c c a s i o n s d u r i n g the e n t i r e study and were p r o b a b l y a n i m a l s b r i e f l y s e p a r a t e d from o t h e r groups. T a b l e 3. Group types e n c o u n t e r e d a t La Raya d u r i n g e n t i r e s t u d y . GROUP TYPE MEAN SIZE ( + SE) % (n) MIXED-SEX 9.5 ( + 0.54) 52.9 (108) MALE 1.8 ( + 0.21) 22.5 (46) FEMALE 4.2 ( + 0.46) 24.5 (50) I c o n s i d e r f o u r a s p e c t s of g r o u p i n g p a t t e r n s : 1) the mean s i z e of the t h r e e t y p e s of groups d u r i n g the y e a r ; 2) changes i n the p r o p o r t i o n of t y p e s of groups over the y e a r ; 3) the d i s t r i b u t i o n of i n d i v i d u a l deer c l a s s e s among the d i f f e r e n t g r oups; and 4) the t e m p o r a l s t a b i l i t y of these groups. 30 T a b l e 4. T o t a l number of groups w i t h and w i t h o u t young ( y e a r -l i n g s and fawns p o o l e d t o g e t h e r ) f o r the e n t i r e s t u d y . GROUP TYPE WITH YOUNG WITHOUT YOUNG MIXED-SEX 81 26 MALE 4 * 42 FEMALE 28 22 * These groups c o n t a i n e d no fawns. Mean group s i z e M i x e d groups were by f a r the l a r g e s t t hroughout the year though they were r e l a t i v e l y s m a l l e r (mean = 7.7-8.8) b e f o r e and d u r i n g fawning ( J a n u a r y - A p r i l ) than (mean = 10.4-11.7) d u r i n g the d r y season when ma t i n g took p l a c e ( F i g . 8 ) . On average mixed groups c o n s i s t e d of 2.4 a d u l t males, 3.9 a d u l t f e m a l e s , and, i f p r e s e n t , 2.8 young (fawns and y e a r l i n g s p o o l e d t o g e t h e r ) . T a b l e 5 p r e s e n t s the d e t a i l e d c o m p o s i t i o n of mixed groups throughout the y e a r . Male groups were v e r y s t a b l e i n s i z e and c o n s i s t e d m a i n l y of one or two a d u l t males ( F i g . 8 ) . The l a r g e s t male group c o n t a i n e d 5 a d u l t s and 2 y e a r l i n g s . Y e a r l i n g s of e i t h e r sex o c c u r r e d w i t h a d u l t males on o n l y f o u r o c c a s i o n s between January and A p r i l . Female groups were l a r g e r (mean = 5.4) d u r i n g and a f t e r fawning (March-June) and s m a l l e r (mean = 2.5-4.5) d u r i n g the r e s t of the year ( F i g . 8 ) . On average they 31 F i g u r e 8. Mean s i z e of deer groups : mixed-sex (MX), female ( F ) , and male (M). V e r t i c a l l i n e s r e p r e s e n t +/- SE. 1980/1981 d a t a have been combined and p o o l e d i n t o 2-month p e r i o d s . 32 14H 12^ 104 0 N CO CL 8 O O c CO CD 6H 44 2H • M X o F • M / / i - 0 - 4 — i J /F — i 1 1 1 1 — M / A M/J J / A S / O N/D 33 T a ble 5. S t r u c t u r e of mixed-sex groups d u r i n g e n t i r e s t u d y . Only groups f o r which a l l i n d i v i d u a l s were c l a s s e d a r e i n c l u d e d . F o r each c l a s s , the mean number of i n d i v i d u a l s per group ( + SE) was o b t a i n e d by p o o l i n g data f o r 1980 and 1981 i n t o 2-month p e r i o d s . MONTH No.GROUPS AD.MALES AD.FEMALES YEARLINGS FAWNS JAN/FEB 22 2 .3 ( ± 0 . 2 8 ) 2 .7 ( +0 . 32 ) 2.1 ( +0 . 49 ) 0 .0 MAR/APR 1 2 2 .0 ( +0 .33 ) 2 . 9 ( +0 . 47 ) 3 .5 ( + 0 . 51 ) 0 .3 ( ± 0 . 2 2 ) MAY/JUN 27 2.6 ( +0 .23 ) 5. 1 ( + 0 . 64 ) 2 . 2 ( ± 0 . 3 6 ) 1 .8 ( +0 .42 ) JUL/AUG 9 2 .7 ( +o !53 ) 5.0 ( ± 1 . 4 7 ) 0 . 3 ( ± 0 . 1 7 ) 0 . 0 SEP/OCT 1 3 2 .6 (+0.31 ) 3 .9 ( + 0 . 47 ) 3 .2 ( ± 0 . 4 4 ) * NOV/DEC 1 3 1 .8 ( +0 .25 ) 3 .0 ( + 0 . 45 ) 1 . 1 ( + 0 . 33 ) * * Y e a r l i n g s and o l d e r fawns p o o l e d t o g e t h e r . c o n s i s t e d of 2.5 a d u l t f e m a l e s , and i f p r e s e n t , 3.2 young (fawns and y e a r l i n g s p o o l e d ) . T able 6 g i v e s the c o m p o s i t i o n of female groups t h r o u g h o u t the y e a r . P r o p o r t i o n of t y p e s of groups M i x e d groups were p r e s e n t y e a r - r o u n d and, w i t h the e x c e p t i o n of J a n u a r y - A p r i l , were the most common group-type o b s e r v e d ( F i g . 9 ) . The low p r o p o r t i o n of mixed groups i n the sample b e f o r e and d u r i n g f a w n i ng was a s s o c i a t e d w i t h a s l i g h t 34 T a b l e 6. S t r u c t u r e of female groups d u r i n g e n t i r e s t u d y . Only groups f o r which a l l i n d i v i d u a l s were c l a s s e d a r e i n c l u d e d . For each c l a s s , mean number of i n d i v i d u a l s per group ( + SE) was o b t a i n e d by p o o l i n g d a t a f o r 1980 and 1981 i n t o 2-month p e r i o d s . MONTH No.GROUPS AD. FEMALES YEARLINGS FAWNS JAN/FEB 1 7 1 .8 ( +0 .22 ) 0 .9 ( +0 .44 ) 0 . 2 (+o. 14) MAR/APR 1 4 2.6 (+0.33) 0 .7 ( +0 .27 ) 2.1 (+0. 44) MAY/JUN 1 2 3.1 ( ± 0 . 5 1 ) 1 .2 ( +0 .49 ) 1 .2 (+0. 68) JUL/AUG 2 2 . 5 ( +0 .50 ) 0 .0 0 . 0 SEP/OCT 3 2.3 (+0.88) 0.3 (+0.33) 0 . 0 NOV/DEC 2 4 . 0 (+3 .00 ) 0 . 0 0 . 5 (+0. 50) i n c r e a s e i n the p r o p o r t i o n of male groups and a marked i n c r e a s e i n t he p r o p o r t i o n of female groups. A f t e r f awning, mixed groups became p r o g r e s s i v e l y more common and by November/December, when a l l males had dropped t h e i r a n t l e r s or were i n v e l v e t , 80% of a l l groups were mixed. D i s t r i b u t i o n of deer c l a s s e s among groups The manner i n which d i f f e r e n t sex/age c l a s s e s a s s o c i a t e d w i t h d i f f e r e n t t y p e s of groups v a r i e d s e a s o n a l l y ( F i g s . 10-14). A d u l t males spent most of the time i n mixed g r o u p s , except d u r i n g f a wning, when more than 50% of them were found i n male groups. The p a t t e r n f o r a d u l t females was s i m i l a r t o t h a t of a d u l t males. They spent most of the ti m e i n mixed 35 F i g u r e 9. P r o p o r t i o n of d i f f e r e n t t y p e s of g r o u p s : : mixed-sex (MX), male (M), and female ( F ) . N= number of groups s i g h t e d . 1980/1981 d a t a have been combined and p o o l e d i n t o 2-month p e r i o d s . 37 g r o u p s , except d u r i n g f a w n i n g , when 50% of them were a s s o c i a t e d w i t h t h e i r fawns, y e a r l i n g s , or o t h e r f e m a l e s . Both a d u l t males and females were a s s o c i a t e d i n mixed groups or w i t h a n i m a l s of t h e i r own sex throughout the y e a r . S o l i t a r y a n i m a l s were i n f r e q u e n t l y o b s e r v e d . A d u l t males, however, were s o l i t a r y more o f t e n than a d u l t females ( F i g . 12). The p r o p o r t i o n of s i n g l e males was above 9% between January and August, and reached a peak of 14% i n M a r c h / A p r i l d u r i n g f a w n i n g . Only 3% of a l l a d u l t males were found a l o n e between September and December, d u r i n g a n t l e r shedding and growth. D u r i n g most of the year s i n g l e females were e x t r e m e l y r a r e . Only i m m e d i a t e l y b e f o r e fawning d i d the p r o p o r t i o n of l o n e females i n c r e a s e t o 8%. Most y e a r l i n g s remained i n mixed groups throughout the y e a r ( F i g . 13). Young fawns were o b s e r v e d e x c l u s i v e l y i n female groups between Fe b r u a r y and A p r i l ( F i g . 14). A few o l d e r fawns were found i n mixed groups i n A p r i l when some l a c t a t i n g females began t o r e t u r n t o mixed groups. By May/June almost a l l fawns were w i t h t h e i r mothers i n the l a r g e mixed groups. S e g r e g a t i o n of pregnant females and a d u l t males The o c c u r r e n c e of s m a l l fawns o n l y i n female groups from F e b r u a r y t o A p r i l , c l e a r l y i n d i c a t e s t h a t a l l pregnant females s e g r e g a t e d from mixed groups and t h u s from a d u l t males. No female groups were obse r v e d i n December 1980 or 1981. Females began t o s e g r e g a t e i n mid J a n u a r y , when 6 (29% of a l l 38 F i g u r e 10. R e l a t i v e number of a d u l t males i n mixed-sex and male groups. MX= mixed-sex g r o u p s ; M= male groups; n= number of a d u l t male s i g h t i n g s . 1980/1981 d a t a have been combined and p o o l e d i n t o 2-month p e r i o d s . 39 too I 1 1 1 • J / F M / A M / J J / A S / O N / D n 69 5 0 8 8 35 37 31 40 F i g u r e 11. R e l a t i v e number of a d u l t females i n mixed-sex and female groups. MX= mixed-sex groups; F= female groups; n= number of a d u l t female s i g h t i n g s . 1980/1981 d a t a have been combined and p o o l e d i n t o 2-month p e r i o d s . 41 i I | 1 • J /F M / A M / J J / A S/0 N / D n 92 71 176 50 58 53 42 F i g u r e 12. P e r c e n t a g e of a d u l t males and females o b s e r v e d as s o l i t a r y i n d i v i d u a l s . S i n g l e females accompanied by fawns have been i n c l u d e d . SM= s o l i t a r y males; SF= s o l i t a r y f e m a l e s . 1980/1981 d a t a have been combined and p o o l e d i n t o 2-month p e r i o d s . 4 3 44 F i g u r e 13. R e l a t i v e number of y e a r l i n g s i n mixed-sex, male, and female groups. MX= mixed-sex groups; M= male g r o u p s ; F= female groups; Y= l o n e y e a r l i n g s ; n= number of y e a r l i n g s i g h t i n g s . 1980/1981 d a t a have been combined and p o o l e d i n t o 2-month p e r i o d s . 46 F i g u r e 14. R e l a t i v e number of fawns i n female and mixed-sex groups. MX= mixed-sex groups; F= female groups; n= number of fawn s i g h t i n g s . For t h i s f i g u r e , fawns born i n 1981 have been c o n s i d e r e d as such u n t i l t he end of the study (December 1981) t o show t h e i r y e a r l y a s s o c i a t i o n p a t t e r n . 48 groups i n t h i s month) female groups were encou n t e r e d . Some l a c t a t i n g females were f i r s t seen i n mixed groups i n e a r l y A p r i l . These group o b s e r v a t i o n s and the r e c o r d s of two known pregnant females i n d i c a t e t h a t the s e x u a l s e g r e g a t i o n may have l a s t e d from 5 t o 10 weeks. My o b s e r v a t i o n s a l s o suggest t h a t once a d u l t females became s e g r e g a t e d , they may a l s o a v o i d c o n t a c t w i t h male or mixed groups. A d i f f e r e n t p a t t e r n was found among a d u l t males. Records of known a d u l t males (5 of which were the l a r g e s t males d u r i n g the s t u d y ) i n d i c a t e t h a t a l t h o u g h up t o 50% of a d u l t male s i g h t i n g s d u r i n g fawning were i n male groups, some r e j o i n e d mixed groups s e v e r a l t i m e s w h i l e o t h e r s r a r e l y l e f t mixed groups. T h i s f i n d i n g i n d i c a t e s t h a t a d u l t males do not move away from mixed-sex groups; r a t h e r , females s e g r e g a t e t h e m s e l v e s . Group s t a b i l i t y From the a n a l y s i s so f a r , i t i s not c l e a r how s t a b l e or " c o h e s i v e " the v a r i o u s t y p e s of groups d e s c r i b e d were a t d i f f e r e n t t i m e s of the y e a r . A measure of r e l a t i v e s t a b i l i t y can be o b t a i n e d by comparing changes i n group s i z e and c o m p o s i t i o n w i t h i n o b s e r v a t i o n p e r i o d s w i t h l o n g e r - t e r m changes o b t a i n e d from d a t a on known i n d i v i d u a l s . On the whole, a l l g r o u p - t y p e s remained v e r y s t a b l e i n s i z e and c o m p o s i t i o n w i t h i n o b s e r v a t i o n p e r i o d s (Table 7) both d u r i n g and o u t s i d e the fawning season. Even l a r g e groups (9-18 49 a n i m a l s ) r e t a i n e d t h e i r i d e n t i t y d u r i n g o b s e r v a t i o n s t h a t l a s t e d up t o 6 h o u r s . T h i s s u g g e s t s t h a t t a r u c a s formed c o h e s i v e s o c i a l a g g r e g a t i o n s a t l e a s t d u r i n g d a i l y a c t i v i t i e s . T a b l e 7. Changes i n group s i z e / c o m p o s i t i o n w i t h i n o b s e r v -a t i o n p e r i o d s . Only o b s e r v a t i o n s of 30 min or l o n g e r have been i n c l u d e d . GROUP-TYPE TOTAL NUMBER NUMBER OF CHANGES OBSERVATIONS PER OBSERVATION/HOUR FEMALE 10 0.02 MALE 12 0.02 MIXED-SEX 43 0.01 Data on known i n d i v i d u a l s , however, i n d i c a t e * t h a t s p e c i f i c groups d i d not remain s t a b l e over s e v e r a l days. Known a d u l t males and females were b o t h o b s e r v e d t o be i n groups of d i f f e r e n t s i z e or c o m p o s i t i o n on a l m o s t a l l o c c a s i o n s i n which t h e s e i n d i v i d u a l s were i d e n t i f i e d ( T a b l e 8 ) . L o c a t i o n a l d a t a from t h e s e known a n i m a l s i n d i c a t e t h a t b o t h sexes remained i n p a r t i c u l a r a r e a s d u r i n g the study ( F i g . 15) and s t r o n g l y suggest t h a t a d u l t s have y e a r - r o u n d home range s , though s e a s o n a l s h i f t s i n h a b i t a t use a r e marked (see 50 F i g u r e 15. L o c a t i o n s of known a d u l t males and fe m a l e s . Only i n d i v i d u a l s w i t h 5 or more s i g h t i n g s and w i t h more than 30 days between f i r s t and l a s t s i g h t i n g have been i n c l u d e d . 52 Tab l e 8. Number of s i g h t i n g s of known i n d i v i d u a l s and number of d i f f e r e n t groups i n which the former were found. Only a n i m a l s w i t h 5 or more s i g h t i n g s a r e i n c l u d e d . Mean time-span between s i g h t i n g s was 15 days. Mean time-span between f i r s t and l a s t s i g h t i n g was 135 days. CLASS NO. ANIMALS TOTAL SIGHTINGS TOTAL NO. OF DIFFERENT GROUPS ADULT MALES 8 80 74 ADULT FEMALES 4 27 27 Chapter 2 ) . T h i s r e s u l t e d i n d i f f e r e n t a n i m a l s a s s o c i a t i n g w i t h p a r t i c u l a r i n d i v i d u a l s or groups but not w i t h o t h e r s w i t h i n the study a r e a . I t appears t h a t a l t h o u g h p a r t i c u l a r groups were not s t a b l e on a l o n g - t e r m b a s i s , d i f f e r e n t i n d i v i d u a l s seemed t o be p a r t of l a r g e r " s u b - p o p u l a t i o n " u n i t s . To summarize, t a r u c a s d i s p l a y e d a s t r o n g tendency t o form s o c i a l a g g r e g a t i o n s t h r o u g h o u t the y e a r . S o l i t a r y a n i m a l s were i n f r e q u e n t . Group s i z e and s t r u c t u r e , however, changed s e a s o n a l l y and t h e s e changes were h i g h l y c o r r e l a t e d w i t h the r e p r o d u c t i v e c y c l e of the d e e r . Mixed-sex groups were the l a r g e s t and most common, except d u r i n g the fawning season when s m a l l e r s i n g l e - s e x groups were more f r e q u e n t . At t h i s t i m e , a l l pregnant females (and perhaps some b a r r e n females as w e l l ) were c o m p l e t e l y s e g r e g a t e d from mixed-sex groups. A l t h o u g h a d u l t males were f r e q u e n t l y seen i n male groups d u r i n g t h i s p e r i o d , d a t a on known i n d i v i d u a l s i n d i c a t e t h a t male groups a r e 53 t r a n s i e n t . My o b s e r v a t i o n s s t r o n g l y suggest t h a t pregnant females move away from and s u b s e q u e n t l y a v o i d c o n t a c t w i t h male or mixed groups. F i n a l l y , the s e a s o n a l g r o u p i n g s of t a r u c a s are n e i t h e r random a g g r e g a t i o n s of i n d i v i d u a l s nor s t a b l e s o c i a l u n i t s . Both sexes formed s h o r t - t e r m c o h e s i v e s o c i a l groups w h i l e f o r a g i n g and engaging i n o t h e r a c t i v i t i e s d u r i n g the day but d i d not aggregate always w i t h the same i n d i v i d u a l s on a l o n g e r - t e r m b a s i s . A l t h o u g h u n s t a b l e , t h e s e groups were com p r i s e d of i n d i v i d u a l s who remained i n r e s t r i c t e d a r e a s y e a r -round. Thus, these "open" groups appeared t o be s u b s e t s of l a r g e r and perhaps more s t a b l e s u b - p o p u l a t i o n u n i t s . 54 J_.4 DISCUSSION R e p r o d u c t i v e C y c l e Tarucas breed s e a s o n a l l y i n an a r e a c h a r a c t e r i z e d by marked s e a s o n a l f l u c t u a t i o n s i n the e n v i r o n m e n t . M a t i n g o c c u r s i n w i n t e r d u r i n g the d r i e s t months, a n t l e r shedding i n males o c c u r s i n s p r i n g d u r i n g the onset of t h e r a i n y season, and fawning towards the end of the r a i n y season i n l a t e summer. The s p a r s e i n f o r m a t i o n a v a i l a b l e f o r t h i s s p e c i e s i n o t h e r l o c a t i o n s i n s o u t h e r n Peru a g r e e s w i t h my f i n d i n g s . Pearson (1951) examined s i x specimens from A r e q u i p a and Puno i n December. Three a d u l t males had t h e i r a n t l e r s i n v e l v e t and one young male had s h o r t s p i k e a n t l e r s and l a c k e d the l a s t m o l a r s . The r e m a i n i n g two were pregnant f e m a l e s . From the s i z e of one f e t u s (410 mm), he e s t i m a t e d b i r t h i n F e b r u a r y or March. Whitehead (1972) s t a t e d t h a t the r u t i n P e r u o c c u r s between June and August and Roe and Rees (1976) r e c o r d e d i n s t a n c e s of c o u r t s h i p and mounting b e h a v i o r d u r i n g t h e i r o b s e r v a t i o n s between 8 and 15 June i n Puno. They n o t e d t h a t a l l a d u l t males had a n t l e r s f r e e of v e l v e t . A l t h o u g h t h e y observed no fawns, some o l d e r fawns may have been m i s t a k e n f o r y e a r l i n g f e m a l e s . D u r i n g a p r e l i m i n a r y study of t a r u c a s i n Pampa G a l e r a s (15° 40'S, 74° 40'W), Ayacucho, between 24 F e b r u a r y and 20 June 1980, I observed fawns i n A p r i l . I was u n a b l e , however, t o f i n d deer i n March. In the c o u r s e of making 20 group o b s e r v a t i o n s i n t h i s l o c a t i o n , I observed o n l y one i n s t a n c e of mounting b e h a v i o r on 17 May 1980. I c o u l d not f i n d any deer i n June 1980. A l l males 55 had c l e a n a n t l e r s throughout t h i s p i l o t s t u d y . The r e l a t i o n s h i p between b r e e d i n g c y c l e s and s e a s o n a l environments i s w e l l documented f o r many u n g u l a t e s i n n o r t h e r n l a t i t u d e s (e.g. Anderson, 1981; B u n n e l l , 1982; Chapman and Feldhamer, 1982; S c h a l l e r , 1977; Thompson and T u r n e r , 1982) where s e a s o n a l b r e e d i n g presumably o c c u r s i n response t o p e r i o d i c f l u c t u a t i o n s i n f o o d a v a i l a b i l i t y ( B u n n e l l , 1982; G e i s t , 1974a; Thompson and T u r n e r , 1982). The c l i m a t i c p a t t e r n s i n n o r t h e r n l a t i t u d e s , however, a r e d i f f e r e n t from those i n t r o p i c a l h i g h l a n d s ( F r a n k l i n , 1983; T r o l l , 1968) and the d i f f e r e n c e s may e x p l a i n why the t i m i n g of r e p r o d u c t i v e e v e nts i n Hippocamelus a n t i s e n s i s d i f f e r s from t h a t i n n o r t h e r n l a t i t u d e u n g u l a t e s . Temperate c l i m a t e s a r e c h a r a c t e r i z e d by s e a s o n a l v a r i a t i o n s i n day l e n g t h and t e m p e r a t u r e . T h i s p a t t e r n r e s u l t s i n most young b e i n g born d u r i n g s p r i n g and e a r l y summer, a p e r i o d of r a p i d p l a n t growth. The onset and l e n g t h of the b i r t h season i n any p a r t i c u l a r l o c a t i o n i s r e l a t e d t o the d u r a t i o n and p r e d i c t a b i l i t y of t h i s p l a n t - g r o w i n g season ( B u n n e l l , 1982; Thompson and T u r n e r , 1982). On the o t h e r hand, i n t r o p i c a l h i g h l a n d s c l i m a t e i s c h a r a c t e r i z e d by l a r g e d i u r n a l f l u c t u a t i o n s i n temperature and season i s i n f l u e n c e d by summer r a i n s . Changes i n day l e n g t h throughout the y e a r a r e r e l a t i v e l y s m a l l . Drought and subzero n i g h t t e m p e r a t u r e s i n the w i n t e r l i m i t the growing season t o one p a r t of the year ( F r a n k l i n , 1983; T r o l l , 1968). D u r i n g the d r y w i n t e r months, most a r e a s a r e d e v o i d of green v e g e t a t i o n and 56 many annual f o r b s and s m a l l g r a s s e s d i s a p p e a r . At the onset of the wet season ( i n s p r i n g ) i n c r e a s i n g r a i n s b r i n g m o i s t u r e t o the d r i e d - o u t s o i l s , but the p e r s i s t e n t n i g h t f r o s t s (see Table 1) r e s u l t i n a slow r e c o v e r y of the v e g e t a t i o n . A l t h o u g h I d i d not m o n i t o r p l a n t phenology q u a n t i t a t i v e l y , my o b s e r v a t i o n s i n d i c a t e t h a t p l a n t growth a t La Raya, as i n o t h e r s i t e s i n s o u t h e r n P e r u ( F r a n k l i n , 1983; Menard, 1984), reaches i t s peak sometime i n February/March. T h i s time appears t o be the o p t i m a l fawning time i n Hippocamelus . The v i c u n a (Lama v i c u g n a ) i s the o n l y o t h e r l o c a l w i l d u n g u l a t e l i v i n g as h i g h as the t a r u c a . I t too has a r e s t r i c t e d b r e e d i n g season i n s o u t h e r n Peru and g i v e s b i r t h t o young between F e b r u a r y and A p r i l ( F r a n k l i n , 1974), w i t h a peak of b i r t h s i n March ( K o f o r d , 1957; p e r s . o b s . ) . Another w i l d u n g u l a t e of comparable s i z e t h a t i n h a b i t s t r o p i c a l h i g h l a n d s i s the W a l i a i b e x (Capra i b e x w a l i a ) . I t i s found i n the Simen Mountains of E t h i o p i a , a t about the same l a t i t u d e as La Raya but n o r t h from the e q u a t o r . Seasons a r e thus r e v e r s e d . A l t h o u g h W a l i a i b e x e s breed throughout the y e a r , they have d i s t i n c t r u t t i n g and b i r t h peaks ( N i e v e r g e l t , 1974, 1981). B i r t h s a r e most common i n September/October, a t the end of the r a i n y s e a s o n , and t h i s time c o i n c i d e s w i t h g r e a t e r a v a i l a b i l i t y of f r e s h f o o d ( N i e v e r g e l t , 1974). The f a c t t h a t W a l i a i b e x e s can b r e e d y e a r - r o u n d presumably r e f l e c t s the more c o n s t a n t e n v i r o n m e n t a l c o n d i t i o n s t h a t e x i s t i n the Simen Mountains. I b e x e s a r e found a t an average e l e v a t i o n of 3,390 m ( N i e v e r g e l t , 1974) which i s lower than t h a t f o r Hippocamelus a t La Raya (mean = 4,661 m). A l t h o u g h the Simen Mountains have a w e l l - d e f i n e d 57 r a i n y season (from May t o September/October) and l a r g e d i u r n a l t e m p e r a t u r e f l u c t u a t i o n s (mean temperature remains f a i r l y c o n s t a n t throughout the y e a r ) , mean minimum t e m p e r a t u r e s never drop below 0° C and even a b s o l u t e minimum t e m p e r a t u r e s never f a l l below -2.5° C ( N i e v e r g e l t , 1981: T a b l e 1 ) . The m i l d e r c l i m a t i c c o n d i t i o n s and the f a c t t h a t numerous water s o u r c e s o u t l a s t the d r y season i n the Simen Mountains r e s u l t i n green v e g e t a t i o n b e i n g a v a i l a b l e y e a r - r o u n d ( N i e v e r g e l t , 1974). I c o n c l u d e t h a t the b r e e d i n g c y c l e i n Hippocamelus i s s t r o n g l y s e a s o n a l and i s d e t e r m i n e d by the p r e c i p i t a t i o n and temperature p a t t e r n s c h a r a c t e r i s t i c of the h i g h l a n d s of s o u t h e r n P e r u . A q u a n t i t a t i v e s t u d y of the e f f e c t s of r a i n f a l l and t e m p e r a t u r e on p l a n t growth and a comparison of t a r u c a b r e e d i n g c y c l e s under d i f f e r e n t c l i m a t i c c o n d i t i o n s a r e needed t o t e s t t h i s h y p o t h e s i s . D i f f e r i n g e n v i r o n m e n t a l c o n d i t i o n s seem t o e x i s t w i t h i n the g e o g r a p h i c range of Hippocamelus a n t i s e n s i s T a r u c a s have been r e p o r t e d i n a number of h i g h Andean h a b i t a t s r a n g i n g from more e q u a t o r i a l environments i n La L i b e r t a d (about 8° S ) , P e r u , t o more temperate l a t i t u d e s i n Catamarca (about 28° S ) , A r g e n t i n a , and from w e t t e r and more c o n s t a n t c l i m a t e of the e a s t e r n Andean s l o p e s t o d r i e r and more u n p r e d i c t a b l e environments of the western s l o p e s of the Andes (I.U.C.N., 1982). My o b s e r v a t i o n s of two fawns o u t s i d e the main fawning season a t La Raya, suggest t h a t t a r u c a s c o u l d breed a t o t h e r t i m e s of the year and t h u s be a b l e t o adapt t o l o c a l c o n d i t i o n s w i t h i n t h e i r e n t i r e range. 58 G r e q a r i o u s n e s s of Tarucas and T h e i r "Open" H a b i t a t At La Raya, t a r u c a s l i v e i n open h a b i t a t and a r e g r e g a r i o u s . The l a r g e s t group I o b s e r v e d c o n s i s t e d of 31 a n i m a l s a l t h o u g h groups of over 40 deer have been r e p o r t e d o c c a s i o n a l l y by p e r s o n n e l of La Raya Research S t a t i o n . S o l i t a r y d e er were uncommon. I n f o r m a t i o n on the d i s t r i b u t i o n of t a r u c a s suggest t h a t they l i v e i n open mountainous t e r r a i n throughout t h e i r range (I.U.C.N., 1982; J u n g i u s , 1974; P e a r s o n , 1951; Roe and Rees, 1976; Whitehead, 1972). Z o o a r c h a e o l o g i c a l f i n d i n g s i n d i c a t e t h a t n a t i v e s have hunted t a r u c a s i n the h i g h Andes s i n c e a t l e a s t 6,000 b.c. (Wheeler, 1982). Grimwood's (1969) c l a i m t h a t the d i s t r i b u t i o n of Hippocamelus a n t i s e n s i s i s d e t e r m i n e d by the a v a i l a b i l i t y of P o l y l e p i s f o r e s t s i s thus unfounded (Roe, 1974; Roe and Rees, 1976; p e r s . o b s . ) . Data on group s i z e e lsewhere a r e meager, but they suggest t h a t t a r u c a s a r e a l s o g r e g a r i o u s i n o t h e r l o c a t i o n s . Hofmann and Ponce d e l Prado (1970) r e p o r t e d a group of 15 deer and obser v e d a l o n e f e m a l e t a r u c a on h i g h , s t e e p s l o p e s , i n a l p i n e t u n d r a h a b i t a t near Abra Malaga, Cusco. In the same a r e a , " l a r g e h e r d s " were common i n p r e v i o u s y e a r s (Hofmann and Ponce d e l Prado, 1970). J u n g i u s (1974) o b s e r v e d some s o l i t a r y males and groups of 3 t o 13 a n i m a l s i n rugged t e r r a i n above t h e t r e e l i n e i n n o r t h w e s t e r n B o l i v i a , and Pearson (1951) found t a r u c a s i n groups of 2 t o 8 a n i m a l s among g r a s s y h i l l s i n t e r s p e r s e d by rugged r o c k o u t c r o p p i n g s i n s o u t h e r n P e r u . Roe and Rees (1976) observed s i x grou p s r a n g i n g from 3 t o 14 deer i n the same t y p e of h a b i t a t i n 59 s o u t h e r n P e r u . From t h e i r d a t a ( T a b l e 1, p. 726) I computed a mean group s i z e of 6.8 ( + 1.58 SE) dee r . I n Pampa G a l e r a s , Ayacucho, I found t a r u c a s at v e r y low d e n s i t i e s (approx. 0.3 deer/km 2) o c c u p y i n g the same mountainous, open t e r r a i n . Group s i z e ranged from 1 t o 11 a n i m a l s , but mean s i z e was lower (mean = 2.65 + 0.55 SE) than i t was a t La Raya (mean = 5.8 + 0.49 SE) f o r the same months. The r e l a t i o n s h i p between group s i z e and h a b i t a t s t r u c t u r e has been e x t e n s i v e l y examined f o r A f r i c a n u n g u l a t e s and has l e d t o development of a g e n e r a l model. A c c o r d i n g t o E s t e s (1974), G e i s t (1974b), Jarman (1974), and L e u t h o l d (1977) u n g u l a t e s p e c i e s or p o p u l a t i o n s o c c u p y i n g open h a b i t a t s l i v e i n s o c i a l g roups, whereas those i n h a b i t i n g dense bush or f o r e s t t e n d t o l i v e s i n g l y or i n v e r y s m a l l groups. Groups i n open h a b i t a t presumably form i n response t o g r e a t e r a v a i l a b i l i t y and more even d i s t r i b u t i o n of food r e s o u r c e s and t o p r e d a t i o n p r e s s u r e s d e r i v e d from t h e i r c o n s p i c u o u s n e s s i n open t e r r a i n . A l t h o u g h t h i s model was de v e l o p e d m a i n l y f o r A f r i c a n b o v i d s , i t seems t o a p p l y t o c e r v i d s as w e l l ( C l u t t o n - B r o c k e t a l . , 1982; E i s e n b e r g , 1981; H i r t h , 1977; Maublanc et a l . , 1985; T a k a t s u k i , 1983). As p r e d i c t e d by the above model, the s o c i a l s t r u c t u r e of Hippocamelus a n t i s e n s i s d i f f e r s from t h a t of i t s c l o s e s t r e l a t i v e , the C h i l e a n huemul (H. b i s u l c u s ) . In t h e Nevados de C h i l i a n and R i o C l a r o a r e a s , P o v i l i t i s (1978, 1983) found huemuls m a i n l y i n dense shrub or f o r e s t h a b i t a t s a t e l e v a t i o n s between 1,450 and 1,700 m. Mean group s i z e was v e r y s m a l l (mean 60 = 1.6 a n i m a l s ) and huemuls were seen s i n g l y or i n a d u l t male-female p a i r s w i t h or w i t h o u t young. Another c l o s e r e l a t i v e of Hippocamelus, the w h i t e - t a i l e d deer ( O d o c o i l e u s v i r g i n i a n u s ) , l i v e s i n a v a r i e t y of h a b i t a t s , but tends t o form l a r g e s o c i a l groups o n l y i n open h a b i t a t s ( H i r t h , 1977). S i m i l a r phenomena have been obse r v e d i n Roe deer, C a p r e o l u s c a p r e o l u s (Maublanc e t a l . , 1985), and S i k a d e e r , Cervus nippon ( T a k a t s u k i , 1983). A l t h o u g h the g e n e r a l tendency of t a r u c a s t o form s o c i a l a g g r e g a t i o n s can be e x p l a i n e d by the above model, a c l o s e r e x a m i n a t i o n of the h a b i t a t i s r e q u i r e d t o e x p l a i n s e a s o n a l changes i n s o c i a l s t r u c t u r e of the s p e c i e s . For example, i t i s u n c l e a r why a d u l t sexes s e g r e g a t e and females form s m a l l e r groups d u r i n g t h e fawning season, when foo d s u p p l i e s a r e presumably most abundant and e v e n l y d i s t r i b u t e d . A l s o , i f p r e d a t i o n f a v o r s the f o r m a t i o n of l a r g e groups i n open h a b i t a t , why do females not form even l a r g e r groups a t a time when r i s k of p r e d a t i o n of o f f s p r i n g i s presumably g r e a t e s t ? In the next c h a p t e r , I seek t o answer these q u e s t i o n s by examining the h a b i t a t use and a n t i p r e d a t o r b e h a v i o r of Hippocamelus a n t i s e n s i s . 61 CHAPTER 2. HABITAT USE BY MALES AND FEMALES 2.J_ INTRODUCTION In many s p e c i e s , males occupy d i f f e r e n t h a b i t a t s than females ( f i s h : K e a s t , 1977; r e p t i l e s : Schoener, 1967; b i r d s : P e t e r s and Grubb, 1983; S e l a n d e r , 1966; and mammals: Bowers and Smith, 1979; Bowyer, 1984; H a r r i s o n , 1983; N i e v e r g e l t , 1981; Shank, 1982). S e x u a l d i f f e r e n c e s i n h a b i t a t use can i n f l u e n c e the s o c i a l dynamics of a s p e c i e s , s i n c e annual changes i n the h a b i t a t r e q u i r e m e n t s of each sex may determine t h e i r p r o x i m i t y . C o n s e q u e n t l y , one way t o i n v e s t i g a t e the i n f l u e n c e of the environment on s o c i a l s t r u c t u r e i s t o d e t e r m i n e and compare h a b i t a t use by males and f e m a l e s . S p a t i a l s e g r e g a t i o n of males and females i s wi d e s p r e a d among s o c i a l c e r v i d s , and t y p i c a l l y t a k e s p l a c e soon a f t e r the mating season (Bowyer, 1984; Brown, 1974; Chapman and Chapman, 1975; C l u t t o n - B r o c k et a l . , 1982; H i r t h , 1977; T a k a t s u k i , 1983). The extreme s e x u a l dimorphism of the s e s p e c i e s may f a c i l i t a t e h a b i t a t s e g r e g a t i o n of the sexes (Bowyers, 1984; McC u l l o u g h , 1979). Some sex d i f f e r e n c e s i n h a b i t a t use have been documented (e.g. S t a i n e s e t a l . 1982). However, the r o l e of n i c h e s e p a r a t i o n of males and females i s s t i l l p o o r l y u n d e r s t o o d ( M c C u l l o u g h , 1979; Shank, 1982). A l e a d i n g h y p o t h e s i s t o e x p l a i n s e x u a l d i f f e r e n c e s i n h a b i t a t use i n s o c i a l u n g u l a t e s i s t h a t d i f f e r e n t i a l m e t a b o l i c and n u t r i t i o n a l r e q u i r e m e n t s of males and females r e s u l t i n 62 d i f f e r e n t i a l f o o d - h a b i t a t s e l e c t i o n ( C l u t t o n - B r o c k e t a l . , 1982; G e i s t , 1974a,b; M c C u l l o u g h , 1979; N i e v e r g e l t , 1981). T h i s h y p o t h e s i s assumes t h a t t h e h i g h c o s t s of g e s t a t i o n and l a c t a t i o n , c o u p l e d w i t h s m a l l e r body s i z e , r e s u l t i n females h a v i n g g r e a t e r e n e r g e t i c and n u t r i t i o n a l r e q u i r e m e n t s per u n i t of body-weight than do males. These d i f f e r e n c e s then l e a d females t o s e l e c t h a b i t a t s w i t h q u a l i t a t i v e l y b e t t e r f o r a g e , and a l l o w males t o be l e s s s e l e c t i v e and s u b s i s t on p o o r e r foods ( C l u t t o n - B r o c k and Harvey, 1983; C l u t t o n - B r o c k e t a l . , 1982; G e i s t , 1974a,b; M c C u l l o u g h , 1979). F u r t h e r m o r e , n i c h e s e g r e g a t i o n may a l s o l e s s e n i n t e r - s e x u a l c o m p e t i t i o n , e s p e c i a l l y d u r i n g t i m e s c r i t i c a l f o r females ( G e i s t , 1974a,b; G e i s t and P e t o c z , 1977). Some s t u d i e s have shown t h a t s p a t i a l s e g r e g a t i o n of t h e sexes i s a s s o c i a t e d w i t h d i f f e r e n c e s i n food h a b i t s ( C h a r l e s e t a l , 1977; S t a i n e s e t a l . , 1982; T a k a t s u k i , 1980; Watson and S t a i n e s , 1978) or o t h e r m e t a b o l i c r e q u i r e m e n t s such as water needs (Bowyer, 1984). Other s t u d i e s have produced ambiguous ( C l u t t o n - B r o c k et a l . , 1982; M c C u l l o u g h , 1979) and o p p o s i t e (LaGory, 1985; Shank, 1982) r e s u l t s t o those p r e d i c t e d by t h i s h y p o t h e s i s , i n d i c a t i n g t h a t o t h e r f a c t o r s can cause s e x u a l s e g r e g a t i o n . An a l t e r n a t i v e h y p o t h e s i s i s t h a t females s e g r e g a t e from males t o reduce r i s k s of p r e d a t i o n t o t h e i r young (LaGory, 1985; M c C u l l o u g h , 1979). A l t h o u g h t h i s h y p o t h e s i s has not been e x p l i c i t l y f o r m u l a t e d t o e x p l a i n h a b i t a t s e g r e g a t i o n of the sexes i n u n g u l a t e s , i t appears r e l e v a n t f o r a number of i m p o r t a n t r e a s o n s : (1) s e x u a l s e g r e g a t i o n i s most pronounced d u r i n g the season of b i r t h s ( B u n n e l l and F a r r , 1983; De Vos e t 63 a l . , 1967; P o v i l i t i s , 1983; S c h a l l e r , 1977); (2) w i t h the e x c e p t i o n of the s i c k and v e r y o l d , young a r e the most v u l n e r a b l e i n d i v i d u a l s t o p r e d a t i o n ( C u r i o , 1976; Mech, 1970; M u r i e , 1944; S c h a l l e r , 1972; S i n c l a i r , 1977; W i l s o n , 1984); and (3) u n l i k e a d u l t s , young of many u n g u l a t e s r e l y on h i d i n g r a t h e r than on f l i g h t t o escape p r e d a t i o n d u r i n g t h e i r f i r s t weeks or months of l i f e (De Vos et a l . , 1967; E s t e s , 1974; Jac o b s e n , 1984; L e n t , 1974; L e u t h o l d , 1977). In many s p e c i e s , young go t h r o u g h a " h i d i n g " phase t h a t may l a s t up t o 4 months ( L e n t , 1974; E s t e s , 1974). Among c e r v i d s , h i d i n g of young appears t o be n e a r l y u n i v e r s a l (De Vos et a l . , 1967; L e n t , 1974). The e f f e c t i v e n e s s of h i d i n g b e h a v i o r as an a n t i p r e d a t o r s t r a t e g y depends on the a b i l i t y of young t o f i n d adequate concealment or c r y p t i c background. C e r t a i n p h y s i c a l c h a r a c t e r i s t i c s of open, . mountainous h a b i t a t s , such as rock o u t c r o p s and broken t e r r a i n , can o f f e r c o n s i d e r a b l e concealment. These f e a t u r e s a r e used f o r p r o t e c t i o n a g a i n s t p r e d a t i o n i n some s p e c i e s ( M u r i e , 1944; M c F e t r i d g e , 1977; S c h a l l e r , 1977; Shannon e t a l . , 1975; Smith, 1983). Thus, a major p r e d i c t i o n of the p r e d a t i o n h y p o t h e s i s i s t h a t females s h o u l d s e l e c t h a b i t a t s t h a t p r o v i d e good h i d i n g p l a c e s f o r t h e i r young d u r i n g the b i r t h s e ason. In the t a r u c a , males and females s e p a r a t e most s t r i k i n g l y d u r i n g t h e fawning season (Chapter 1 ) . The s p e c i e s i s m arkedly s e x u a l l y d i m o r p h i c . Males a r e l a r g e r than females and o n l y males grow a n t l e r s . T a r ucas a l s o l i v e i n a t o p o g r a p h i c a l l y d i v e r s e environment t h a t o f f e r s a c h o i c e of 64 d i f f e r e n t h a b i t a t s . However, i t i s not known i f the sexes s e g r e g a t e by h a b i t a t or what f a c t o r s cause t h e i r s e p a r a t i o n . In t h i s C h a p t e r , I d e s c r i b e h a b i t a t s i n the study s i t e a t La Raya, P e r u , and a s s e s s the e x t e n t and r o l e of h a b i t a t s e p a r a t i o n of the sexes i n Hippocamelus a n t i s e n s i s . I e v a l u a t e h a b i t a t use by female, male, and mixed-sex groups i n r e l a t i o n t o : (1) e l e v a t i o n ; (2) s l o p e of the t e r r a i n ; (3) v e g e t a t i o n -t y p e ; (4) d i s t a n c e t o water s u p p l i e s ; (5) concealment ( p r o v i d e d by r o c k - c o v e r and r e l i e f f e a t u r e s ) ; and (6) a n t i p r e d a t o r b e h a v i o r and escape t e r r a i n . A l t h o u g h a d d i t i o n a l hypotheses a r e c o n s i d e r e d , I d i s c u s s e c o l o g i c a l s e p a r a t i o n of the sexes m a i n l y i n l i g h t of the hypotheses t h a t : (1) females s e l e c t the be s t f o r a g i n g h a b i t a t s t o meet h i g h e r m e t a b o l i c demands of l a c t a t i o n ; and (2) females s e l e c t h a b i t a t s t h a t p r o v i d e the g r e a t e s t concealment f o r t h e i r fawns. 65 2.2 MATERIALS AND METHODS 2.2._1_ G r i d System and E n v i r o n m e n t a l F a c t o r s Recorded A l l of the a n a l y s e s r e f e r t o the main study a r e a ( F i g . 1). I d i v i d e d t h i s a r e a i n t o 250x250-m q u a d r a t s and drew a g r i d system on a t o p o g r a p h i c a l map of La Raya ( s c a l e 1:25,000). A t o t a l of 1,264 q u a d r a t s were c l a s s i f i e d a c c o r d i n g t o 8 e c o l o g i c a l v a r i a b l e s w i t h the a i d of the map, a e r i a l and panoramic ph o t o g r a p h s , and t e r r e s t r i a l s u r v e y s of the a r e a . The f o l l o w i n g e n v i r o n m e n t a l f a c t o r s were r e c o r d e d a t m i d - p o i n t of each q u a d r a t . (1) E l e v a t i o n , t o t h e n e a r e s t 25-m c o n t o u r i n t e r v a l . (2) V e g e t a t i o n - t y p e , w i t h i n a 50-m r a d i u s . Three broad c l a s s e s were c o n s i d e r e d : "Long-Grass", dominated by p e r e n n i a l s t r a w g r a s s e s ( C a l a m a g r o s t i s , S t i p a , F e s t u c a ) r e a c h i n g h e i g h t s of over 15 cm (but r a r e l y measuring over 50 cm); " S h o r t - G r a s s " , l a c k i n g l o n g g r a s s e s and dominated m a i n l y by annual s h o r t g r a s s e s (Hordeum , Bromus) and f o r b s ( A l c h e m i l l a , Gnaphalium , H i p o c h o e r i s ) m e a s u r i n g 15 cm or l e s s i n h e i g h t , and; "Marsh", dominated by p e r e n n i a l and annual f o r b s ( D i s t i c h i a , H i p o c h o e r i s , L u z u l a , P l a n t a g o ) and g r a s s e s ( C a l a m a g r o s t i s , A g r o s t i s ) g rowing i n m o i s t a r e a s . Areas d e v o i d of any v e g e t a t i o n were c l a s s e d as "Bare". (3) D i s t a n c e t o n e a r e s t  water s u p p l y , measured o n l y i f water l o c a t i o n f e l l o u t s i d e q u a d r a t . O t h e r w i s e i t was r e c o r d e d as " p r e s e n t " . In a d d i t i o n , the f o l l o w i n g f a c t o r s were r e c o r d e d f o r the e n t i r e q u a d r a t . (4) Average g r a d i e n t of t h e s l o p e , i n 66 d e g r e e s . (5) R o c k i n e s s of t e r r a i n , i n p e r c e n t c o v e r . (6) Amount of permanent snow/ice cover , i n p e r c e n t c o v e r . (7) Number of tr o u g h s and/or r i d q e s , o n l y r e l i e f f e a t u r e s l a r g e enough t o c o n c e a l a d e e r - s i z e d o b j e c t were c o n s i d e r e d . (8) Presence/absence of t e r r a c e ( s ) . A l i s t of the s e e n v i r o n m e n t a l f a c t o r s and t h e i r r e l a t i v e f r e q u e n c i e s a re g i v e n i n s e c t i o n 2.3.1 (Table 11). 2.2.2 Deer O b s e r v a t i o n s and Data Recorded C o l l e c t i o n of d a t a on deer d i s t r i b u t i o n and h a b i t a t use was g r e a t l y i n f l u e n c e d by the t o p o g r a p h i c a l f e a t u r e s of the a r e a . The rugged t e r r a i n made s y s t e m a t i c t r a n s e c t s a m p l i n g ( e . g . Western , 1976) i m p o s s i b l e . A l s o , because of i t s s i z e (79 km 2), I c o u l d not cov e r the e n t i r e a r e a i n one day. I , t h e r e f o r e d i v i d e d the a r e a i n t o 8 s u b u n i t s (Table 9) each of which c o u l d be c o v e r e d i n a day. T h i s p a r t i t i o n i n g was based on geomorphology. Tarucas were se a r c h e d from f i x e d s e a r c h r o u t e s and from n a t u r a l vantage p o i n t s . Once a deer group was found, I would s t a y a t the l o c a t i o n f o r as l o n g as the group remained i n s i g h t . Time p e r m i t t i n g , I c o n t i n u e d t o s e a r c h f o r o t h e r groups i n the same a r e a , or moved on t o another s u b u n i t . The b i a s e s i n h e r e n t i n t h i s method and t h e i r p o s s i b l e i n f l u e n c e on my r e s u l t s a r e d i s c u s s e d below (see s e c t i o n s 2.2.3 and 2.3.3). 67 Ta b l e 9. S i z e of the 8 s u b u n i t s i n the main st u d y a r e a . SUBUNIT No. Of QUADRATS km2 % STUDY AREA CHIMBOLLA 134 8.4 10.6 HUARIPINA 1 56 9.8 12.3 JATUNCUCHO 240 15.0 19.0 JUCHUYCUCHO 1 19 7.4 9.4 PULPERAPATA 58 3.6 4.6 TAMBO 1 90 11.9 15.0 VISCACHANE 124 7.8 9.8 YAHUARCOTA 243 15.2 19.2 TOTAL 1 ,264 79. 1 100.0 T a b l e 10 p r e s e n t s the t o t a l number of v i s i t s and the number of groups obse r v e d i n each s u b u n i t d u r i n g the e n t i r e s t u d y . Only group o b s e r v a t i o n s made w h i l e s e a r c h i n g f o r deer i n a l l r e p r e s e n t a t i v e h a b i t a t s a r e i n c l u d e d ( i e . c a s u a l s i g h t i n g s have been o m i t t e d ) . When a group of deer was l o c a t e d , the f o l l o w i n g a d d i t i o n a l e n v i r o n m e n t a l f a c t o r s were r e c o r d e d a t the b e g i n n i n g , end, and e v e r y hour d u r i n g the o b s e r v a t i o n : (1) date ; (2) time ; (3) l o c a t i o n : r e c o r d e d on the map; (4) e l e v a t i o n : a t the 68 Table 10. Number of v i s i t s t o each s u b u n i t w i t h i n the main study a r e a and number of groups o b s e r v e d d u r i n g the e n t i r e s t u d y . SUBUNIT No. VISITS No. GROUPS CHIMBOLLA 41 41 HUARIPINA 17 10 JATUNCUCHO 35 55 JUCHUYCUCHO 1 1 8 PULPERAPATA 32 5 TAMBO 6 0 VISCACHANE 49 50 YAHUARCOTA 19 19 TOTAL 210 188 group's l o c a t i o n t o the n e a r e s t 25-m c o n t o u r i n t e r v a l ; and (5) v e g e t a t i o n - t y p e : w i t h i n 50-m r a d i u s around a n i m a l s . The remainder f a c t o r s ( T able 11, s e c t i o n 2.3.1) were g i v e n t h e c l a s s of the q u a d r a t i n which the deer were o b s e r v e d . On o c c a s i o n s when I d i s t u r b e d and f l u s h e d the d e e r , the t o p o g r a p h i c f e a t u r e (e.g. t r o u g h , r i d g e , mountain-top, r o c k - o u t c r o p ) t h a t caused me to l o s e s i g h t of the a n i m a l s was r e c o r d e d as "escape c o v e r " . Some r e c o r d s were not used i n h a b i t a t a n a l y s e s . These were: (1) o b s e r v a t i o n s a f t e r a group was d i s t u r b e d by the o b s e r v e r ; (2) the f i n a l r e c o r d of o b s e r v a t i o n s l e s s than 1/2 hr i n d u r a t i o n ; (3) the f i n a l r e c o r d of o b s e r v a t i o n s more than 1/2 69 hr but l e s s than 1 hr i n d u r a t i o n i f the r e c o r d f e l l w i t h i n the same h o u r - p e r i o d as the f i r s t r e c o r d (e.g. f i r s t r e c o r d a t 9:00 and f i n a l r e c o r d a t 9:45); and (4) c a s u a l o b s e r v a t i o n s (see above). A t o t a l of 310 o b s e r v a t i o n s were r e c o r d e d d u r i n g the e n t i r e s t u d y . 2.2.3 Data A n a l y s i s In most a n a l y s e s t h a t f o l l o w I have used the MIDAS (M i c h i g a n I n t e r a c t i v e Data A n a l y s i s System) s t a t i s t i c a l package. H a b i t a t d a t a i n c l u d e d i n t e r v a l , o r d i n a l and nominal v a r i a b l e s . T h i s p r e c l u d e d the use of some v a r i a b l e s i n a few a n a l y s e s and r e q u i r e d d a t a t r a n s f o r m a t i o n i n o t h e r s . A d e s c r i p t i o n of the main study a r e a was o b t a i n e d from i n f o r m a t i o n g a t h e r e d f o r a l l q u a d r a t s . Three t y p e s of a n a l y s e s were used: 1) fr e q u e n c y d i s t r i b u t i o n of the 8 e n v i r o n m e n t a l v a r i a b l e s ( i n t e r v a l d a t a were d i v i d e d i n t o d i s c r e t e c a t e g o r i e s ) ; 2) p a r t i a l c o r r e l a t i o n s among i n t e r v a l and o r d i n a l v a r i a b l e s ; and 3) f r e q u e n c y d i s t r i b u t i o n of v a r i a b l e s w i t h i n f i v e a l t i t u d i n a l l e v e l s . My s a m p l i n g t e c h n i q u e s d i d not account f o r h a b i t a t a v a i l a b i l i t y f o r two r e a s o n s . F i r s t , r e p e a t e d v i s i t s t o a g i v e n s u b u n i t were not always comparable s i n c e I o f t e n s e a r c h e d an a r e a o n l y u n t i l a deer group was found. Thus on o c c a s i o n s I d i d s e a r c h a whole s u b u n i t . Second, q u a d r a t s c o u l d not be se a r c h e d e q u a l l y , s i n c e v i s i b i l i t y was i n f l u e n c e d by d i s t a n c e and by d i f f e r i n g t o p o g r a p h i c f e a t u r e s (e.g. amount of r o c k i n e s s , 70 d i r e c t i o n of s l o p e , and r i c h n e s s of r e l i e f f e a t u r e s ) . A l t h o u g h my a n a l y s e s cannot a c c u r a t e l y r e v e a l h a b i t a t s e l e c t i o n by t a r u c a s , comparisons of h a b i t a t use between sexes or t y p e s of groups a r e p o s s i b l e . For the s e comparisons I o n l y used d a t a from q u a d r a t s where deer were o b s e r v e d . Observed f r e q u e n c i e s were then compared t o e x p e c t e d h a b i t a t f r e q u e n c i e s . S i n c e v i s i b i l i t y c o u l d i n f l u e n c e t h e s e c o m p a r i s o n s , the r e s u l t s may s t i l be b i a s e d . I e v a l u a t e d t h i s b i a s by c h e c k i n g f o r l a r g e d e v i a t i o n s from e x p e c t e d h a b i t a t f r e q u e n c i e s and d e t e r m i n i n g the d i r e c t i o n of the v i s i b i l i t y b i a s f o r a p a r t i c u l a r v a r i a b l e . In g e n e r a l , however, deer were found more f r e q u e n t l y i n s i t e s w i t h reduced v i s i b i l i t y . I t h e r e f o r e p r o b a b l y m i s s e d some groups, and thus may have u n d e r e s t i m a t e d a c t u a l use of the p r e f e r r e d h a b i t a t s . I a s s e s s e d h a b i t a t use by groups r a t h e r than i n d i v i d u a l s f o r two r e a s o n s . F i r s t , p r e l i m i n a r y t e s t s of a s s o c i a t i o n showed the same t r e n d s when i n d i v i d u a l s or groups were compared. Second, i t i s v e r y u n l i k e l y t h a t a l l i n d i v i d u a l s i n a group a c t i v e l y s e l e c t e d a g i v e n s i t e . I n d i v i d u a l s i n a group moved c o h e s i v e l y , o f t e n i n s i n g l e f i l e , between f o r a g i n g or r e s t i n g s i t e s . An a d u l t female u s u a l l y l e d female or mixed-sex groups. To compare the h a b i t a t use by female, male, and mixed-sex groups I d i v i d e d the y e a r i n t o t h r e e p e r i o d s . The f i r s t one, January t h r o u g h A p r i l , spanned the fawning season and had the h i g h e s t monthly p r o p o r t i o n of s i n g l e - s e x g r o u p s . The second p e r i o d , May th r o u g h August, i n c l u d e d the mating season, which 71 c o i n c i d e d w i t h the d r i e s t time of the y e a r . D u r i n g the l a s t p e r i o d , September through December, males shed t h e i r a n t l e r s and the r a i n y season s e t s i n . In t h e l a s t two p e r i o d s the p r o p o r t i o n of s i n g l e - s e x groups was v e r y low. These t h r e e p e r i o d s I r e f e r t o as "fawning", "mating", and " a n t l e r - s h e d d i n g " seasons. Comparisons of h a b i t a t use were made as f o l l o w s : 1) comparisons among female, male, and mixed-sex groups e n c o u n t e r e d d u r i n g the fawning season; and 2) comparisons of each of the above group-types i n the fawning season w i t h mixed-sex groups found d u r i n g the o t h e r two seasons. 72 2.3 RESULTS 2.3.1 H a b i t a t and R e l a t i o n s between E n v i r o n m e n t a l F a c t o r s In a rugged, h i g h mountainous t e r r a i n w i t h open v e g e t a t i o n such as t h a t found a t La Raya, topography and o t h e r g e o m o r p h o l o g i c a l f e a t u r e s c o n t r i b u t e g r e a t l y t o t h e t h r e e -d i m e n s i o n a l s t r u c t u r e and h e t e r o g e n e i t y of the h a b i t a t . The r e s u l t i n g s p a t i a l c o m p l e x i t y i s a mosaic of h a b i t a t c o n d i t i o n s r a t h e r than w e l l - d e l i m i t e d h a b i t a t - t y p e s . I f i r s t d e s c r i b e the f e a t u r e s of the main study a r e a : a) e l e v a t i o n , b) permanent snow-cover, c) s l o p e , d) r o c k - c o v e r , e) number of r e l i e f f e a t u r e s , f ) d i s t a n c e t o water s o u r c e s , and g) v e g e t a t i o n - t y p e s . Second, I a s s e s s r e l a t i o n s h i p s between p a i r s of e n v i r o n m e n t a l v a r i a b l e s . F i n a l l y , I compare the h a b i t a t s i n f i v e a l t i t u d i n a l l e v e l s . G e n e r a l d e s c r i p t i o n of the a r e a T a b l e 11 g i v e s a summary of the h a b i t a t . The 1,400-m a l t i t u d i n a l drop from the h i g h e s t peak t o the l o w e s t v a l l e y bottom i n the a r e a encompassed both a permanent ice/snow l i n e (around 5,200 m) t h a t changed l i t t l e s e a s o n a l l y , and the uppermost l i m i t of the v e g e t a t i o n (between 4,900 and 5,000 m). A p p r o x i m a t e l y 90% of the a r e a was f r e e of permanent snow c o v e r . D u r i n g the wet, summer season, snow r a r e l y a c c u mulated below the permanent snow l i n e , and i t never s t a y e d on the ground below the v e g e t a t i o n ' s upper l i m i t f o r more than 3 days a t a t i m e . Thus 73 T a b l e 11. H a b i t a t d e s c r i p t i o n of the main study a r e a a c c o r d i n g t o the e n v i r o n m e n t a l v a r i a b l e s r e c o r d e d f o r each qua d r a t (N=1,264). ELEVATION (m) 4,075-4,275 16.4 4,300-4,575 31.4 4,600-4,875 27.1 4,900-5,175 19.8 5,200-5,450 5.4 SLOPE (degrees) <15 22.3 >=15-<30 48.3 >=30 29.4 RELIEF (sum of r i d g e s <=2 70.1 and t r o u g h s ) 3-4 23.3 >=5 6.6 ROCK COVER (%) 0 43.8 <50 37.3 >=50 18.8 PERMANENT SNOW COVER (%) 0 90.4 <50 3.4 >=50 6.2 TERRACE absent 91.1 p r e s e n t 8.9 VEGETATION TYPE l o n g g r a s s 53.6 marsh 11.2 s h o r t g r a s s 4.1 bare ground 31.1 DISTANCE TO NEAREST p r e s e n t i n quadrat 28.0/21.8 * OPEN WATER SOURCE (m) ab s e n t ; <=500 41.6/37.3 ab s e n t ; >500-<=l000 25.6/29.5 ab s e n t ; >1000 4.8/11.4 * Nov-May/Jun-Oct 74 snow c o v e r was p r o b a b l y not a major f a c t o r a f f e c t i n g deer d u r i n g the s t u d y . Most water s u p p l i e s i n the a r e a (see b e l o w ) , however, o r i g i n a t e d from the permanent snow f i e l d s . The s l o p e of the t e r r a i n v a r i e d from 0° t o 50° (mean = 22° ). F l a t or g e n t l y s l o p i n g t e r r a i n was m o s t l y a s s o c i a t e d w i t h v a l l e y bottoms w h i l e s t e e p t e r r a i n was c h a r a c t e r i s t i c of mid and upper mountain s i d e s . About 70% of the a r e a had s l o p e s of 15° or s t e e p e r . R o c k - o u t c r o p s , b o u l d e r f i e l d s , or rock escarpments (some massive) i n t e r s p e r s e d g r a s s meadows and were p r e s e n t i n 56% of a l l q u a d r a t s , m a i n l y on mountain s i d e s and t o p s . Rock-cover was one of the few f e a t u r e s i n t h i s "open" environment t h a t c o u l d p r o v i d e good concealment f o r deer. Other f a c t o r s t h a t p r o v i d e d some concealment were t r o u g h s , r a v i n e s , s h o u l d e r s , and r i d g e s . About 30% of the q u a d r a t s had 3 or more of t h e s e r e l i e f f e a t u r e s . Other r e l i e f f e a t u r e s c o n s i s t e d of s m a l l t e r r a c e s (few were l a r g e r than the 250x250-m q u a d r a t s ) on mountain s i d e s . These were uncommon ( p r e s e n t i n 9% of the q u a d r a t s ) but seemed i m p o r t a n t f o r c e r t a i n deer groups. B e i n g f l a t , exposed a r e a s , t e r r a c e s o f f e r e d l i t t l e concealment except from below. Open water s o u r c e s were common. D u r i n g the r a i n y season, 70% of the q u a d r a t s were l o c a t e d w i t h i n 500 m of a water s u p p l y . D u r i n g the d r y season, t h i s f i g u r e dropped t o about 60%. Most water s o u r c e s c o n s i s t e d of streams, but a few ponds and s e a s o n a l w a t e r - h o l e s were a l s o p r e s e n t . F i n a l l y , Long-grass type was p r e s e n t i n 54% of a l l q u a d r a t s . Next were Marsh ( 1 1 % ) , and S h o r t - g r a s s ( 4 % ) . Appendix A l i s t s and g i v e s the mean number of p l a n t s p e c i e s found i n each of t h e s e v e g e t a t i o n - t y p e s . The remainder of the a r e a (30%) l a c k e d any v e g e t a t i o n . 75 C o r r e l a t i o n s between h a b i t a t v a r i a b l e s As e x p e c t e d , t h e r e were c o r r e l a t i o n s between p a i r s of t h e s e v a r i a b l e s (Table 12). The h i g h e s t p o s i t i v e p a r t i a l c o r r e l a t i o n s o c c u r r e d between amount of r o c k - c o v e r and s l o p e ; d i s t a n c e t o water and s l o p e ; and e l e v a t i o n and amount of r o c k -c o v e r . There was no c o r r e l a t i o n between d i s t a n c e t o water and amount of r e l i e f . A n e g a t i v e c o r r e l a t i o n e x i s t e d d u r i n g the r a i n y season between d i s t a n c e t o water and amount of r o c k - c o v e r but t h e s e v a r i a b l e s were not c o r r e l a t e d d u r i n g the dry season. S i n c e v e g e t a t i o n - t y p e s and t e r r a c e s were r e c o r d e d as p r e s e n t or a b s e n t , they were e x c l u d e d from th e c o r r e l a t i o n a n a l y s e s . C h i - s q u a r e t e s t s , however, r e v e a l e d some i m p o r t a n t r e l a t i o n s h i p s . The p r o p o r t i o n of q u a d r a t s l a c k i n g v e g e t a t i o n s h a r p l y i n c r e a s e d w i t h g r e a t e r e l e v a t i o n , r o c k i n e s s , s l o p e , and r e l i e f (p<0.000l i n a l l c a s e s ) . The d e c r e a s e of p l a n t c o v e r w i t h e l e v a t i o n and the f a c t t h a t the a r e a c o n t a i n e d the uppermost a l t i t u d i n a l l i m i t of v e g e t a t i o n , suggest a l s o a n e g a t i v e r e l a t i o n s h i p between p l a n t biomass per u n i t a r e a and e l e v a t i o n w i t h i n the study a r e a . T h i s was shown i n a p r e v i o u s s t u d y i n t h i s a r e a (Holgado et a l . , 1979). The p r o p o r t i o n of a r e a s w i t h bare ground a l s o i n c r e a s e d w i t h d i s t a n c e t o w a t e r , d u r i n g b o t h the r a i n y and dry seasons (p<0.0001 i n both c a s e s ) . The p r e s ence of v e g e t a t i o n was independent of the presence of t e r r a c e s (p>0.05). There were a l s o a s s o c i a t i o n s between p a r t i c u l a r v e g e t a t i o n - t y p e s and o t h e r v a r i a b l e s . The p r o p o r t i o n of Table 12. P a r t i a l c o r r e l a t i o n s between h a b i t a t v a r i a b l e s . Only i n t e r v a l and o r d i n a l v a r i a b l e s were c o n s i d e r e d . The p a r t i a l c o r r e l a t i o n c o e f f i c i e n t f o r a g i v e n v a r i a b l e - p a i r was e s t i m a t e d w i t h a l l o t h e r v a r i a b l e s h e l d c o n s t a n t . The c o r r e l a t i o n c o -e f f i c i e n t (R) and p v a l u e s a r e shown. S i n c e the d i s t r i b u t i o n of f r e e water s o u r c e s v a r i e s w i t h season, p a r t i a l c o r r e l a t i o n s f o r the r a i n y (November-May) and the dry (June-October) seasons a r e p r e s e n t e d s e p a r a t e l y . NOVEMBER-MAY JUNE-OCTOBER VARIABLE-PAIR R P VARIABLE-PAIR R P ROCK/SLOPE .40 <.01 ROCK/SLOPE .40 <.01 DISWATER/SLOPE .32 <.01 ROCK/ELEV .27 <.01 ROCK/ELEV .28 <.01 DISWATER/SLOPE .25 <.01 SLOPE/ELEV .20 <.01 SLOPE/ELEV .21 <.01 RELIEF/ROCK .19 <.01 RELIEF/ROCK .20 <.01 RELIEF/ELEV .17 <.01 RELIEF/ELEV .17 <.01 RELIEF/SLOPE .14 <.01 RELIEF/SLOPE .12 <.01 DISWATER/ELEV .08 <.01 DISWATER/ELEV .07 <.05 DISWATER/ROCK - . 10 <.01 ' DISWATER/RELIEF .01 N.S DISWATER/RELIEF - .05 N.S. DISWATER/ROCK .00 N.S N=1264; df=1259 f o r a l l c a s e s . 77 q u a d r a t s w i t h Long-grass or Marsh t y p e s d e c r e a s e d w i t h i n c r e a s i n g e l e v a t i o n (p<0 .000 l i n both c a s e s ) , r o c k i n e s s (p< 0 . 0 0 0 l i n both c a s e s ) , and r e l i e f (p< 0 . 0 5 , and p<0 .0 l r e s p e c t i v e l y ) . The p r o p o r t i o n of q u a d r a t s w i t h S h o r t - g r a s s t y p e , i n c o n t r a s t , i n c r e a s e d w i t h e l e v a t i o n (p< 0 . 0 0 0 l ) and w i t h g r e a t e r r o c k i n e s s of the t e r r a i n ( p < 0 . 0 l ) . S h o r t - g r a s s h a b i t a t was a l s o a s s o c i a t e d w i t h t e r r a c e s more than e x p e c t e d (p < 0 . 0 5 ) . Long-grass and Marsh h a b i t a t s o c c u r r e d i n d e p e n d e n t l y of presence of t e r r a c e s (p> 0 . 0 5 , i n both c a s e s ) . As exp e c t e d , the p r o p o r t i o n of marshy a r e a s d e c r e a s e d w i t h i n c r e a s i n g d i s t a n c e t o water y e a r - r o u n d ( p < 0 . 0 0 0 l ) . D u r i n g the r a i n y season, t h e r e was no a s s o c i a t i o n between the p r o p o r t i o n of e i t h e r Long-grass or S h o r t - g r a s s and d i s t a n c e t o water (p>0.05 i n both c a s e s ) . D u r i n g the dry season, however, the p r o p o r t i o n of both these two v e g e t a t i o n - t y p e s i n c r e a s e d w i t h i n c r e a s i n g d i s t a n c e t o water s o u r c e s ( p < 0 . 0 l , and p<0 .00 l r e s p e c t i v e l y ) . A l t i t u d i n a l l e v e l s To i l l u s t r a t e the r e l a t i o n s h i p between e l e v a t i o n and o t h e r h a b i t a t v a r i a b l e s , I d i v i d e d the main study a r e a i n t o 5 a l t i t u d i n a l l e v e l s ( T a b l e 1 3 ) . L e v e l 1 ( 4 , 0 7 5 - 4 , 2 7 5 m) c o n s i s t e d m a i n l y of v a l l e y bottom and had good d r a i n a g e , g e n t l y s l o p i n g t e r r a i n , and e x t e n s i v e c o v e r by v e g e t a t i o n . T h i s l e v e l had the h i g h e s t p r o p o r t i o n (30%) of q u a d r a t s w i t h marshy v e g e t a t i o n . However, r o c k - c o v e r and r e l i e f f e a t u r e s were r a r e . L e v e l 2 ( 4 , 3 0 0 - 4 , 5 7 5 m) was c h a r a c t e r i z e d by h i g h v a l l e y - b o t t o m s and low mountain s i d e , good d r a i n a g e , t e r r a i n of 78 v a r y i n g s l o p e s , and good co v e r by v e g e t a t i o n . The p r o p o r t i o n of Long-grass v e g e t a t i o n reached i t s h i g h e s t v a l u e (83%) her e . T h i s l e v e l had l i t t l e r o c k - c o v e r and o n l y a few r e l i e f f e a t u r e s . L e v e l 3 (4,600-4,875 m) i n c l u d e d m a i n l y mid mountain s i d e s . T h i s l e v e l had a h i g h p e r c e n t of r o c k - c o v e r (82% of q u a d r a t s had a t l e a s t some r o c k - c o v e r ) and r e l i e f f e a t u r e s were common (43% of q u a d r a t s had 3 or more t r o u g h s and/or r i d g e s ) . I t was a l s o c h a r a c t e r i z e d by v e r y s t e e p t e r r a i n ( s l o p e s of 30° or s t e e p e r g r a d i e n t s were p r e s e n t i n 44% of q u a d r a t s i n t h i s l e v e l ) . Water s u p p l i e s were l e s s common than they were i n lower l e v e l s . About 64% of q u a d r a t s i n t h i s l e v e l l a y w i t h i n 500 m of water d u r i n g the r a i n y season. T h i s p r o p o r t i o n dropped t o 42% d u r i n g the d r y se a s o n , the lo w e s t v a l u e among a l l l e v e l s . Some v e g e t a t i o n was p r e s e n t i n 70% of the q u a d r a t s . The p r o p o r t i o n of q u a d r a t s w i t h S h o r t - g r a s s was h i g h e s t (10%) i n t h i s l e v e l . L e v e l 4 (4,900-5,175 m) c o n s i s t e d of upper mountain s i d e s and some mountain t o p s . As was the case f o r l e v e l 3, i t had a h i g h p r o p o r t i o n of r o c k - c o v e r , r e l a t i v e l y h i g h number of r e l i e f f e a t u r e s , and v e r y s t e e p s l o p e s . An i m p o r t a n t f e a t u r e of t h i s l e v e l i s t h a t i t c o n t a i n e d the uppermost l i m i t of the v e g e t a t i o n . Only s p a r s e v e g e t a t i o n , dominated by t a l l s t r a w -g r a s s e s , and a few s m a l l p a t c h e s of marshy v e g e t a t i o n were p r e s e n t . Here, a few s c a t t e r e d low s h r u b s , r a r e l y measuring over 50 cm i n h e i g h t , were a l s o found. E i g h t y - e i g h t p e r c e n t of the q u a d r a t s l a c k e d v e g e t a t i o n . L e v e l 5 (5,200-5,450 m) i n c l u d e d the h i g h e s t mountain peaks i n the a r e a . S l o p e s were i n t e r m e d i a t e i n st e e p n e s s 79 between l e v e l s 2 and 3. Snow/ice f i e l d s c o v e r e d most of the ground. T h i s l e v e l had " l i t t l e r o c k - c o v e r , few r e l i e f f e a t u r e s , and l a c k e d v e g e t a t i o n . 2.3.2 H a b i t a t Use by the Deer G e n e r a l I o b s e r v e d groups of t a r u c a most f r e q u e n t l y (96%) on mountain s i d e s and t o p s a t a mean e l e v a t i o n of 4,661 m ( + 10.7 SE). Deer were l a r g e l y absent from two of the f i v e a l t i t u d i n a l l e v e l s d e s c r i b e d above. I never o b s e r v e d t a r u c a s i n l e v e l 1 (main v a l l e y - b o t t o m ; 4,075-4,275 m), a l t h o u g h they have o c c a s i o n a l l y been seen here a f t e r dusk or a t dawn by La Raya p e r s o n n e l . T h i s l e v e l was e x t e n s i v e l y used by a l p a c a s and humans. T a r u c a s were seldom o b s e r v e d a t l e v e l 5 (5,200-5,450 m) e i t h e r . I d i d observe one group a t 5,200 m on a snow-free r o c k -o u t c r o p o u t s i d e the main study a r e a . Deer were seen a t the permanent snow l i n e once. H a b i t a t use comparisons There were s i g n i f i c a n t d i f f e r e n c e s i n h a b i t a t use among the d i f f e r e n t t y p e s of groups. I used two approaches t o determine t h e e x t e n t and d i r e c t i o n of t h e s e d i f f e r e n c e s . F i r s t , I c a r r i e d out C h i - s q u a r e t e s t s t o compare use of each i n d i v i d u a l h a b i t a t f a c t o r by groups of d i f f e r e n t t y p e s . T a b l e s B1 t h r o u g h B8 g i v e p e r c e n t use of each h a b i t a t v a r i a b l e and T a b l e s B9 T a b l e 13. Frequency d i s t r i b u t i o n ( i n p e r c e n t ) of h a b i t a t v a r i a b l e s w i t h i n 5 a l t i t u d i n a l l e v e l s i n the main study a r e a . LEVEL 1= 4,075-4,275m; LEVEL 2= 4,300"4575m; LEVEL 3= 4,600-4875m; LEVEL 4= 4,900-5,175m; LEVEL 5= 5,200-5,450m. RELIEF= Sum of r i d g e s and t r o u g h s ; ROCK= P e r c e n t cover by r o c k ; SNOW P e r c e n t c o v e r by permanent snow/ice; DISWAT= D i s t a n c e t o n e a r e s t open water s o u r c e . VARIABLE CLASS LEVEL 1 N=207 LEVEL 2 N=397 LEVEL 3 N=342 LEVEL 4 N=250 LEVEL N=68 100.0% 100.0% 100.0% 100.0% 100.0% SLOPE <1 5 68.6 23.9 8.2 4.0 10.3 >=15<30 25.6 49.4 58.2 48.8 60.3 > = 30 5.8 26.7 43.6 47.2 29.4 RELIEF <2 92.8 74.8 57.0 59.6 77.9 3-4 6.3 22.2 31.6 29.6 16.2 > = 5 1.0 3.0 11.4 10.8 5.9 ROCK 0 % 97. 1 56.7 17.8 10.0 61 .8 <50 % 2.9 36.5 49.7 51 .6 32.4 > = 50 % 0.0 6.8 32.5 38.4 5.9 SNOW 0 % 100.0 100.0 100.0 78.0 2.9 <50 % 0.0 0.0 0.0 12.8 16.2 > = 50 % 0.0 0.0 0.0 9.2 80.9 TERRACE absent 100.0 90.4 86.8 90.0 94. 1 p r e s e n t 0.0 9.6 13.2 10.0 5.9 VEGET l o n g g r a s s 67.6 83.4 54. 1 8.4 0.0 marsh 30.4 11.8 7.0 3.2 0.0 s h o r t g r a s s 2.0 3.8 9.6 0.0 0.0 bare ground 0.0 1.0 29.2 88.4 100.0 DISWAT <=500m 84.5 70.5 64.0 67.2 55.9 NOV-MAY >500m 15.5 29.5 36.0 32.8 44. 1 DISWAT <=500m 84.5 61 .7 41.8 59.2 52.9 JUN-OCT >500m 15.5 38.3 58.2 40.8 47. 1 81 t h r o u g h B11 p r e s e n t s t a t i s t i c a l r e s u l t s of p a i r e d - w i s e group c o m p a r i s o n s . I a l s o used C h i - s q u a r e a n a l y s e s t o t e s t f o r a s s o c i a t i o n s between p a i r s of h a b i t a t v a r i a b l e s i n the ar e a s where d i f f e r e n t t y p e s of groups were o b s e r v e d . I g i v e t h e s e r e s u l t s i n the t e x t below ( o n l y p v a l u e s of the C h i - s q u a r e t e s t s are shown). I compare these a s s o c i a t i o n s t o c o r r e l a t i o n s between the same p a i r s of v a r i a b l e s i n the study a r e a i n s e c t i o n 2.3.3. The second approach uses m u l t i v a r i a t e d i s c r i m i n a n t a n a l y s i s t o determine which h a b i t a t f a c t o r s b e s t d i s c r i m i n a t e the d i f f e r e n t t y p e s of groups and t o o b t a i n an o v e r a l l index of h a b i t a t use o v e r l a p . S i n c e my d a t a c o n t a i n e d i n t e r v a l , o r d i n a l , and nominal v a r i a b l e s , a l l of thes e were t r a n s f o r m e d i n t o dummy v a r i a b l e s (Green, 1976; N i e v e r g e l t , 1981). There was good agreement between d i s c r i m i n a n t a n a l y s e s and C h i - s q u a r e t e s t s ; i e . the bes t v a r i a b l e s f o r d i s c r i m i n a t i n g groups had i n g e n e r a l the h i g h e s t C h i - s q u a r e v a l u e s , and the d i r e c t i o n of d i f f e r e n c e s ( g i v e n by the d i s c r i m i n a n t f u n c t i o n s ) was the same as t h a t from c o n t i n g e n c y t a b l e s . T a b l e C1 l i s t s the dummy v a r i a b l e s used, and T a b l e s C2 and C3 p r e s e n t s the r e s u l t s of the d i s c r i m i n a n t a n a l y s e s . I f i r s t examine h a b i t a t use d i f f e r e n c e s d u r i n g the fawning season (Jan-Apr) when s e x u a l s e g r e g a t i o n took p l a c e . I then g i v e r e s u l t s f o r i n t e r - s e a s o n a l c o m p a r i s o n s . 82 (a) Fawning-season comparisons D u r i n g the fawning season, female groups were found a t s i g n i f i c a n t l y h i g h e r e l e v a t i o n s (mean = 4,684 m + 28.2 SE) than were e i t h e r male (mean = 4,573 m + 26.2 SE) or mixed-sex groups (mean = 4,550 m + 19.2 S E ) . These two l a t t e r groups d i d not d i f f e r i n t h i s r e s p e c t ( T a b l e s B1, B 9 ) . These d i f f e r e n c e s were p a r t l y e x p l a i n e d by d i f f e r e n c e s i n d a i l y v e r t i c a l movement of the groups (Table B2). Mixed-sex groups showed a s t r o n g tendency t o move v e r t i c a l l y d u r i n g the day, u s i n g lower e l e v a t i o n s a t dawn, then moving upwards as the day p r o g r e s s e d (p<0.05). A l t h o u g h male groups showed no s i g n i f i c a n t d i f f e r e n c e s (X 2= 6.6, p>0.05) i n v e r t i c a l d i s t r i b u t i o n d u r i n g the day, i n d i v i d u a l males showed the same v e r t i c a l movement ( X 2 = 13.3, p<0.0l) as d i d mixed-sex groups. Female groups, however, remained a t r e l a t i v e l y h i g h e l e v a t i o n s a l l day (p>0.05). Groups a l s o d i f f e r e d w i t h r e s p e c t t o the s l o p e of the t e r r a i n ( T a b l e s B3, B 9 ) . Female groups used s i g n i f i c a n t l y s t e e p e r s l o p e s than e i t h e r male or mixed-sex groups. No d i f f e r e n c e s i n t h i s r e g a r d were found between male and mixed-sex groups. Female groups always used s t e e p t e r r a i n r e g a r d l e s s of e l e v a t i o n (p>0.05). In c o n t r a s t , f o r male and mixed-sex groups the degree of s l o p e used was dependent on e l e v a t i o n (p<0.000l i n bo t h c a s e s ) . Male and mixed-sex groups o c c u p i e d s t e e p e r t e r r a i n a t h i g h e r e l e v a t i o n s . Groups were found i n a r e a s t h a t d i f f e r e d s i g n i f i c a n t l y 83 i n amount of r o c k - c o v e r ( T a b l e s B 4 , B S ) . Female groups were always found i n t e r r a i n w i t h r o c k - c o v e r , and used r o c k i e r a r e a s than d i d e i t h e r male or mixed-sex groups. Male groups a l s o used a r e a s w i t h more r o c k - c o v e r than d i d mixed-sex groups. The f r e q u e n c i e s of use of roc k y t e r r a i n were independent of e l e v a t i o n f o r a l l t h r e e g r o u p - t y p e s ( p > 0 . 0 5 i n a l l c a s e s ) . For a l l t h r e e t y p e s of groups, t h e r e was a p o s i t i v e r e l a t i o n s h i p between r o c k i n e s s and s l o p e of the t e r r a i n t h a t they used ( p < 0 . 0 l f o r female and male groups, p < 0 . 0 5 f o r mixed-sex g r o u p s ) . Male groups used a r e a s w i t h the l e a s t amount of r e l i e f (number of t r o u g h s and r i d g e s i n a q u a d r a t ) ( T a b l e s B 5 , B 9 ) . Female and mixed-sex groups d i d not d i f f e r i n t h i s r e s p e c t The use of a r e a s w i t h d i f f e r e n t amounts of r e l i e f was r e l a t e d t o e l e v a t i o n f o r a l l t h r e e groups.. The a r e a s used by female groups had more r e l i e f f e a t u r e s a t lower e l e v a t i o n s than they had a t h i g h e r l e v e l s ( p < 0 . 0 5 ) , w h i l e the t e r r a i n used by male and mixed-sex groups i n c r e a s e d i n amount of r e l i e f as they moved t o h i g h e r e l e v a t i o n s ( p < 0 . 0 l and p < 0 . 0 0 l r e s p e c t i v e l y ) . A . s i m i l a r r e l a t i o n s h i p e x i s t e d f o r the use of t e r r a i n w i t h v a r y i n g degrees of r e l i e f and s l o p e . Female groups o c c u p i e d a r e a s w i t h g r e a t e r amount of r e l i e f i f the t e r r a i n was not s t e e p and used a r e a s w i t h l e s s r e l i e f on s t e e p e r t e r r a i n ( p < 0 . 0 0 " l ) . Male and mixed-sex groups, on the o t h e r hand, used a r e a s w i t h g r e a t e r r e l i e f as they moved onto s t e e p e r t e r r a i n ( p < 0 . 0 5 and p < 0 . 0 0 0 l r e s p e c t i v e l y ) . There was a l s o a dependency between r o c k i n e s s and amount of r e l i e f i n the t e r r a i n used by male and female groups. Female groups o c c u p i e d t e r r a i n w i t h g r e a t e r r o c k - c o v e r 84 i n a r e a s w i t h l i t t l e r e l i e f and v i c e v e r s a (p<0.0"l), w h i l e male groups used a r e a s w i t h i n c r e a s i n g p e r c e n t of r o c k - c o v e r as they o c c u p i e d t e r r a i n w i t h g r e a t e r amounts of r e l i e f (p<0.05). Mixed groups used a r e a s i n which r o c k i n e s s of the t e r r a i n was independent of amount of r e l i e f (p>0.05). F i n a l l y , female groups were found on t e r r a c e s s i g n i f i c a n t l y l e s s f r e q u e n t l y than o t h e r groups ( T a b l e s B6, B9). Male and mixed-sex groups d i d not d i f f e r i n t h i s r e g a r d ( T a b l e B 9 ) . No d i f f e r e n c e s e x i s t e d i n the use of v e g e t a t i o n - t y p e s between female and male or female and mixed-sex groups ( T a b l e s B7, B9). Female and mixed-sex groups were found f a r t h e r away from f r e e water s o u r c e s than were male groups ( T a b l e s B8, B9). No d i f f e r e n c e s i n t h i s r e g a r d were d e t e c t e d between female and mixed-sex groups. (b) I n t e r - s e a s o n a l comparisons To a n a l y z e s e a s o n a l s h i f t s i n h a b i t a t use I compared the t h r e e g r o u p - t y p e s o b s e r v e d d u r i n g the fawning season w i t h mixed-sex groups found d u r i n g the mating and a n t l e r - s h e d d i n g seasons. O u t s i d e of the fawning season female and male groups were uncommon. Thus, t h e y are not i n c l u d e d i n the comparisons. S t a t i s t i c a l r e s u l t s f o r the comparisons are g i v e n i n T a b l e s B10 and B11. D u r i n g the mating season (May-Aug), mixed-sex groups o c c u p i e d h i g h e r e l e v a t i o n s (mean = 4,802 m + 21.5 SE) than any o t h e r groups a t any t i m e d u r i n g t h e y e a r . Use of h i g h e r t e r r a i n 85 was a s s o c i a t e d w i t h h i g h e s t use (22.4%) of marshy v e g e t a t i o n , i n d i c a t i n g a s h i f t from d r i e r t o m o i s t e r h a b i t a t s d u r i n g the dry season. T h i s f i n d i n g i s i m p o r t a n t because a s h i f t i n r e l a t i v e use of t y p e s of v e g e t a t i o n took p l a c e when the sexes a g g r e g a t e d . Compared t o female groups i n the fawning season, these mixed-sex groups used s i g n i f i c a n t l y l e s s s t e e p and l e s s r o c k y t e r r a i n . Mixed-sex groups i n the mating season d i d , however, use s t e e p e r t e r r a i n than male or mixed-sex groups i n the fawning season. The a r e a s used by mixed-sex groups i n the mating season and those o c c u p i e d by female or mixed-sex groups i n the fawning season d i d not d i f f e r w i t h r e g a r d t o amount of r e l i e f or d i s t a n c e t o water. Mixed-sex groups used t e r r a i n w i t h g r e a t e r r e l i e f and were found f a r t h e r from water s o u r c e s d u r i n g the mating season than were male groups d u r i n g the fawning season. F i n a l l y , m ating season mixed-sex groups, l i k e fawning season female g r o u p s , made l i t t l e use of t e r r a c e s . Mixed-sex groups d u r i n g the a n t l e r - s h e d d i n g season were found a t h i g h e l e v a t i o n (mean = 4,721 m + 20.8 S E ) , but not as h i g h as t h a t of mixed-sex groups i n the mating season. They were, however, found a t g r e a t e r d i s t a n c e s from water than any o t h e r group-type d u r i n g the y e a r . D u r i n g t h i s season, mixed-sex groups d e c r e a s e d t h e i r use of marshy h a b i t a t s , i n d i c a t i n g a r e t u r n t o u s i n g more x e r i c v e g e t a t i o n . D u r i n g the a n t l e r -s hedding season, mixed-sex groups used v e g e t a t i o n h a b i t a t s w i t h the same freq u e n c y as t h a t of groups d u r i n g the fawning season, i n d i c a t i n g an absence of s h i f t s i n use of v e g e t a t i o n - t y p e s as the sexes s e g r e g a t e d . Groups from the a n t l e r - s h e d d i n g season d i d not d i f f e r from female groups i n the fawning season i n use 86 of s t e e p t e r r a i n , but used s i g n i f i c a n t l y l e s s r o c k y t e r r a i n than d i d the l a t t e r ( T a b l e s B4, B11). Compared t o mixed-sex and male groups i n the fawning season, a n t l e r - s h e d d i n g groups used s t e e p e r s l o p e s . F i n a l l y , a n t l e r - s h e d d i n g groups used a r e a s w i t h l i t t l e r e l i e f , as d i d male groups i n the f a w n i n g season. R e s u l t s from s t e p w i s e m u l t i p l e d i s c r i m i n a n t a n a l y s e s ( T a b l e s C2-C3) agree b r o a d l y w i t h the above r e s u l t s . In a d d i t i o n , they suggest t h a t female and mixed-sex groups d i f f e r e d most d u r i n g the fawning season, w h i l e male and mixed-sex groups were c l o s e s t i n o v e r a l l h a b i t a t use. D u r i n g the mating season, mixed-sex groups (which c o n t a i n e d p r e v i o u s l y s e g r e g a t e d females and t h e i r fawns) appeared c l o s e r i n o v e r a l l h a b i t a t use t o female groups i n the fawning season than t o any o t h e r group. Mixed-sex groups i n the a n t l e r - s h e d d i n g season resembled mating season groups i n h a b i t a t use, w h i l e they were most d i f f e r e n t from male groups i n the fawning season. To summarize, d u r i n g the fawning season p r e g n a n t / l a c t a t i n g females o c c u p i e d d i f f e r e n t h a b i t a t s than groups c o n t a i n i n g a d u l t males. In c o n t r a s t , the h a b i t a t s of male and mixed-sex groups were r a t h e r s i m i l a r . Female groups were r e s t r i c t e d t o h i g h e l e v a t i o n s , w h i l e male and mixed-sex groups o c c u p i e d lower e l e v a t i o n s . Some mixed-sex and male groups, however, moved v e r t i c a l l y each day. Thus, they used a wi d e r range of h a b i t a t s than d i d female g r o u p s . The most s t r i k i n g h a b i t a t d i f f e r e n c e l a y i n the use o f v e r y s t e e p , r o c k y t e r r a i n by female groups. No o t h e r groups, i n s i d e or o u t s i d e the fawning season, used r o c k y h a b i t a t t o the same e x t e n t . 87 Female groups a l s o used t e r r a i n w i t h more t r o u g h s and r i d g e s than d i d contemporary male groups, or mixed-sex groups o u t s i d e the fawning season. Compared t o o t h e r g r o u p s , female groups made l i t t l e use of t e r r a c e s . Groups c o n t a i n i n g females o c c u r r e d f a r t h e r away from water s o u r c e s than d i d male groups. At the time of s e x u a l s e g r e g a t i o n I d e t e c t e d no d i f f e r e n c e s between female and o t h e r groups i n the use of v e g e t a t i o n t y p e s , a l t h o u g h t h e r e was a c l e a r s h i f t from x e r i c t o m o i s t v e g e t a t i o n when the sexes a g g r e g a t e d d u r i n g the dry season. In c o n t r a s t , no s h i f t s i n use of v e g e t a t i o n - t y p e s o c c u r r e d as mixed-sex groups i n the a n t l e r - s h e d d i n g season broke a p a r t i n t o d i f f e r e n t g roup-types i n the fawning season. T a b l e 14 summarizes h a b i t a t comparisons f o r the fawning season. 2.3.3 H a b i t a t S e l e c t i o n , B i a s e s , and C o r r e l a t i o n s I have shown t h a t d i f f e r e n t g r o u p - t y p e s used the h a b i t a t d i f f e r e n t l y . Three i m p o r t a n t q u e s t i o n s remain: 1) were the d i f f e r e n t groups " s e l e c t i n g " or " a v o i d i n g " c e r t a i n h a b i t a t s ? ; 2) how d i d v i s i b i l i t y a f f e c t my comparisons of h a b i t a t use?; and 3) were a s s o c i a t i o n s between h a b i t a t f a c t o r s used by t h e deer i n the d i r e c t i o n p r e d i c t e d from h a b i t a t c o r r e l a t i o n s f o r the whole s t u d y a r e a ? To answer these q u e s t i o n s I r e l a t e h a b i t a t use t o the a c t u a l f requency d i s t r i b u t i o n of h a b i t a t v a r i a b l e s and c o r r e l a t i o n s among them. I r e s t r i c t t h i s a n a l y s i s t o f e m a l e , male, and mixed-sex groups i n t h e fawning season because d u r i n g t h i s p e r i o d a l l group-types were o b s e r v a b l e . 88 T a b l e 14. Summary of h a b i t a t comparisons among femal e , male, and mixed-sex groups d u r i n g the fawning season ( J a n - A p r ) . The symbols ">" and "=" mean s i g n i f i c a n t e l y g r e a t e r (at .05 l e v e l or l e s s ) and not s i g n i f i c a n t r e s p e c t i v e l y . V a r i a b l e s t h a t were b e s t d i s c r i m i n a n t between female/male, and female/mixed-sex group comparisons a r e a l s o i n d i c a t e d . HABITAT VARIABLE GROUPS COMPARED BEST DISCRIMINANT? ELEVATION FEMALE > MALE = MIXED YES SLOPE FEMALE > MALE = MIXED NO ROCK-COVER FEMALE > MALE > MIXED YES RELIEF FEMALE = MIXED > MALE YES* TERRACE MALE = MIXED > FEMALE YES** VEGETATION-TYPES FEMALE = MALE; FEMALE = MIXED NO MALE ( S h o r t - G r a s s ) > MIXED DISTANCE TO WATER FEMALE = MIXED > MALE NO * Only i n female/mixed comparison ** Only i n female/male comparison 89 I e a r l i e r noted t h a t my a n a l y s e s c o u l d not f u l l y a c count f o r h a b i t a t a v a i l a b i l i t y because of samp l i n g t e c h n i q u e s . T a b l e 15 g i v e s approximate e x p e c t e d h a b i t a t v a l u e s f o r the a r e a s m o n i t o r e d d u r i n g the fawning season ( J a n - A p r ) . I e s t i m a t e d e x p e c t e d v a l u e s by c o r r e c t i n g f o r the number of v i s i t s t o each s u b u n i t and then t a l l y i n g the c o r r e c t e d f r e q u e n c i e s from a l l s u b u n i t s . These v a l u e s a r e o n l y a p p r o x i m a t i o n s s i n c e q u a d r a t s i n a g i v e n s u b u n i t were not s e a r c h e d e q u a l l y (see s e c t i o n 2.2.3), though I searched a l l r e p r e s e n t a t i v e h a b i t a t s i n an a r e a d u r i n g each v i s i t . Compared t o ex p e c t e d v a l u e s i n Ta b l e 15, v a l u e s f o r a r e a s where deer were observed showed some l a r g e d e v i a t i o n s . Female groups were found more f r e q u e n t l y than e x p e c t e d on v e r y s t e e p and r o c k y t e r r a i n w i t h h i g h number of r e l i e f f e a t u r e s between 4,600 and 4,875 m. These groups d i d not s e l e c t or a v o i d t e r r a c e s , and were i n d i f f e r e n t t o d i s t a n c e from water s o u r c e s . Male groups o c c u p i e d t e r r a c e s more f r e q u e n t l y than e x p e c t e d , and p r e f e r r e d a r e a s w i t h moderate r o c k - c o v e r and near water s u p p l i e s between 4,300 and 4,575 m. Males were i n d i f f e r e n t t o amount of r e l i e f and s t e e p n e s s of the t e r r a i n they used. Mixed-sex groups o c c u p i e d more o f t e n than e x p e c t e d t e r r a c e s , and t e r r a i n w i t h moderate r o c k - c o v e r , moderate s l o p e , and many r e l i e f f e a t u r e s a t e l e v a t i o n s between 4,300 and 4,575 m. Mixed-sex groups were u n s e l e c t i v e w i t h r e g a r d t o d i s t a n c e t o water. F i n a l l y , a l l groups appeared t o s e l e c t s h o r t - g r a s s h a b i t a t over the o t h e r two v e g e t a t i o n - t y p e s . Were d e v i a t i o n s from e x p e c t e d h a b i t a t v a l u e s a 90 Table 15. H a b i t a t v a r i a b l e f r e q u e n c i e s d u r i n g the fawning season ( J a n - A p r ) . A c t u a l v a l u e s have been a d j u s t e d t o number of v i s i t s t o d i f f e r e n t s u b u n i t s i n the main study a r e a (N=8,842). Permanent snow co v e r has not been i n c l u d e d . VARIABLE CLASSES % of N ELEVATION (m) 4,075-4,275 4.6 4,300-4,575 35.7 4,600-4,875 28. 1 4,900-5,175 24.4 5,200-5,450 7.2 SLOPE (degrees) <15 18.3 >=15-<30 47.7 > = 30 34.0 ROCK-COVER ( p e r c e n t ) 0 32.3 <50 40.8 > = 50 27.0 RELIEF (# of throughs < = 2 59.3 and/or r i d g e s ) 3-4 29.7 >=5 10.9 TERRACE absent 88.2 p r e s e n t 11.8 VEGETATION-TYPE Long g r a s s 43.4 S h o r t g r a s s 6.4 Marsh 11.0 Bare ground 39.3 DISTANCE TO NEAREST <=500 77.3 WATER SOURCE (m) >500 22.7 91 consequence of unequal v i s i b i l i t y of deer i n d i f f e r e n t a r e a s ? T h i s i s u n l i k e l y . A r e a s w i t h good v i s i b i l i t y ( f l a t v a l l e y bottoms and low s l o p e s l a c k i n g r o c k - c o v e r and t o p o g r a p h i c r e l i e f ) were sear c h e d a t l e a s t as i n t e n s i v e l y as h i g h e r t e r r a i n . Though deer were e a s i l y s p o t t e d i n the former a r e a s , they were ob s e r v e d l e s s f r e q u e n t l y t h e r e . Thus, the d e v i a t i o n s of female groups toward more br o k e n , r o c k i e r t e r r a i n were p r o b a b l y r e a l and the v a l u e s o b t a i n e d s h o u l d be c o n s e r v a t i v e . Group-type d i f f e r e n c e s were p r o b a b l y a l s o r e a l . There i s no reason t o expect mixed-sex or male groups t o be l e s s v i s i b l e than female groups i n rugged t e r r a i n . The o p p o s i t e was l i k e l y , s i n c e mixed-sex groups were l a r g e r i n group s i z e , and a d u l t males, owing t o t h e i r l a r g e body s i z e and a n t l e r s , were g e n e r a l l y more c o n s p i c u o u s than a d u l t f e m a l e s . Were a s s o c i a t i o n s i n the use of d i f f e r e n t h a b i t a t f a c t o r s i n the d i r e c t i o n e x p e c t e d from c o r r e l a t i o n s between v a r i a b l e s i n the st u d y a r e a ? T h i s seemed t o be the case o n l y f o r male and mixed-sex groups. There were i n t e r e s t i n g e x c e p t i o n s i n the case of female groups. R o c k i n e s s , r e l i e f , s l o p e , and e l e v a t i o n were a l l p o s i t i v e l y c o r r e l a t e d i n the study a r e a ( T a b l e 12). D e s p i t e t h i s , the use of v e r y s t e e p t e r r a i n by female groups was independent of e l e v a t i o n (see page 8 2 ) . In the t e s t s of a s s o c i a t i o n of r e l i e f / e l e v a t i o n , r e l i e f / s l o p e , and r e l i e f / r o c k - c o v e r , female groups showed n e g a t i v e a s s o c i a t i o n s , not p o s i t i v e as e x p e c t e d and found f o r male and mixed-sex groups (see pages 83,84). T h i s s t r o n g l y s u g g e s t s t h a t female groups were not o n l y h i g h l y s e l e c t i v e i n t h e i r h a b i t a t use, but were a l s o a b l e t o compensate f o r the l a c k of some t o p o g r a p h i c a l 92 f a c t o r ( i e . by c h o o s i n g a r e a s of g r e a t e r r e l i e f i f r o c k - c o v e r was o n l y m o d e r a t e ) . To summarize, d u r i n g the fawning season, groups of t a r u c a s used some h a b i t a t s more f r e q u e n t l y than e x p e c t e d . T h i s was p a r t i c u l a r l y t r u e f o r female groups i n t h e i r use of s t e e p , r o c k y , and broken t e r r a i n . D e v i a t i o n s i n h a b i t a t use from e x p e c t e d h a b i t a t v a l u e s d i d not r e s u l t from s e a r c h i n g e r r o r or v i s i b i l i t y b i a s , but r e f l e c t e d s e l e c t i v i t y i n the use of p a r t i c u l a r h a b i t a t s by d e e r . When I t e s t e d f o r a s s o c i a t i o n s between p a i r s of v a r i a b l e s i n a r e a s used by each g r o u p - t y p e , the t r e n d s f o r male and mixed-sex groups were l a r g e l y those e x p e c t e d from h a b i t a t c o r r e l a t i o n s i n the whole study a r e a . I n c o n t r a s t , female groups responded t o c e r t a i n h a b i t a t f e a t u r e s i n d e p e n d e n t l y or i n o p p o s i t e d i r e c t i o n t o t h a t e x p e c t e d from h a b i t a t c o r r e l a t i o n s . I n c o n c l u s i o n , i n a d d i t i o n t o d i f f e r i n g from o t h e r groups i n h a b i t a t use, female groups were h i g h l y s e l e c t i v e i n t h e i r use of h a b i t a t s . 2.3.4 A n t i p r e d a t o r b e h a v i o r P r e d a t o r s were p r e s e n t i n the study a r e a . In 1981, an a l p a c a h e r d e r found the h a l f - b u r i e d body of a t a r u c a fawn i n a s i t e above the v e g e t a t i o n l i n e . Puma ( F e l i s c o n c o l o r ) t r a c k s were i d e n t i f i e d around the c a r c a s s . Pumas a l s o k i l l e d some a l p a c a s and c a p t i v e v i c u n a s d u r i n g my s t a y a t La Raya. I found puma t r a c k s on two h e a v i l y used d e e r - f e e d i n g a r e a s . Though I never o b s e r v e d pumas, they have o c c a s i o n a l l y been seen by h e r d e r s d u r i n g the day. One puma was k i l l e d by l o c a l p e o p l e i n 93 1981 . Other l i k e l y p r e d a t o r s of deer were s e m i - f e r a l dogs t h a t belonged t o a l p a c a h e r d e r s , and were seen roaming i n the s t u d y a r e a up t o s e v e r a l m i l e s from t h e i r owners' shacks. L o c a l p e o p l e a s s u r e d me t h a t dogs chase and k i l l even a d u l t d e e r . I w i t n e s s e d one chase of a s i n g l e male t a r u c a by two dogs. The dogs gave up the p u r s u i t and came back t o the v a l l e y bottom a f t e r the deer d i s a p p e a r e d over a h i g h r i d g e . T a rucas responded t o the s i g h t or b a r k s of a dog by s t o p p i n g f e e d i n g or s t a n d i n g up i f they were r e s t i n g , and l o o k i n g a t t e n t i v e l y i n the d i r e c t i o n of the dog. A f t e r a s h o r t t i m e , the deer would r a p i d l y c l i m b up t o s t e e p e r , r o c k i e r t e r r a i n or d i s a p p e a r over t h e n e a r e s t r i d g e or mountain t o p . Andean f o x e s (Dusicyon c u l p a e u s ) were a l s o abundant. I o f t e n h e a r d t h e i r c a l l s a t dawn and saw them on f o u r o c c a s i o n s d u r i n g the day i n a r e a s f r e q u e n t e d by t a r u c a s . I saw no i n t e r a c t i o n s between f o x e s and t a r u c a s . Foxes p r o b a b l y r e p r e s e n t no danger f o r grown d e e r , though t h e y may have been c a p a b l e of k i l l i n g newborns (Whitehead, 1972). Andean f o x e s do p r e y on young v i c u n a s ( K o f o r d , 1957; F r a n k l i n , 1983). One way t o study the response of deer t o p r e d a t o r s was t o observe the r e a c t i o n of t a r u c a s t o c l o s e e n c o u n t e r s w i t h the o b s e r v e r . N o r m a l l y , as l o n g as I kept more than 100 meters away and below the d e e r , t a r u c a s would remain u n d i s t u r b e d and would resume t h e i r p r e v i o u s a c t i v i t y soon a f t e r s p o t t i n g me. I approached deer a t c l o s e range on 46 o c c a s i o n s . When I approached them s l o w l y and i n f u l l v i e w , the d e e r ' s r e a c t i o n was 94 t o walk away t o a h i g h e r l o c a t i o n and o f t e n out of s i g h t . However, when s t a r t l e d a t a d i s t a n c e of 50 m or l e s s , t hey would i m m e d i a t e l y f l e e u p h i l l , q u i c k l y d i s a p p e a r i n g from view. When a group was approached from above, they would sometimes go f u r t h e r down or run a l o n g the mountain s i d e a t f i r s t , but they always moved t o h i g h e r t e r r a i n than the o b s e r v e r e v e n t u a l l y . The t o p o g r a p h i c f e a t u r e s t h a t caused me t o l o s e s i g h t of the deer a f t e r a d i s t u r b a n c e a r e shown i n Table 16. For a l l groups, r i d g e s or mountain t o p s were the most common (71%) escape c o v e r , f o l l o w e d by r o c k - o u t c r o p s ( 2 7 % ) . T a b l e 16. Escape c o v e r used by deer groups. In i n s t a n c e s when deer where d i s t u r b e d or " f l u s h e d " by the o b s e r v e r , escape c o v e r was d e f i n e d as the t o p o g r a p h i c a l f e a t u r e t h a t caused the o b s e r v e r t o l o s e s i g h t of the a n i m a l s . ESCAPE COVER GROUP--TYPE FEMALE MALE MIXED-SEX ALL GROUPS %(n) %(n) %(n) %(n) RIDGE/ 58.3(7) 66.7(6) 81 .0(17) 71.4(30) MOUNTAIN TOP TROUGH 8.0(1) (0) (0) 2.4(1) ROCK OUTCROP 33.3(4) 33.3(3) 19.0(4) 26.8(11) Tarucas e x h i b i t e d a number of b e h a v i o r s when al a r m e d . O c c a s i o n a l l y , j u s t p r i o r t o e s c a p i n g , one a n i m a l ( e i t h e r male or 95 female) would g i v e a "sneeze" sound t h a t would i m m e d i a t e l y b r i n g the a t t e n t i o n of any i n d i v i d u a l not a l r e a d y a l e r t e d i n the group. The sneezer c o u l d u s u a l l y be i d e n t i f i e d s i n c e the b e h a v i o r was accompanied by a r a p i d downward motion of the head. The a n i m a l s ' a l e r t n e s s , p r i o r t o f l e e i n g , was o f t e n c h a r a c t e r i z e d by " s t i f f " w a l k i n g , i n which movements of the neck and l e g s were slow and e x a g g e r a t e d , and by t a i l - f l i c k i n g . W h i l e f l e e i n g , deer g e n e r a l l y kept t h e i r t a i l s up and rump h a i r e r e c t e d . T h i s made the rump-patch appear w h i t e r and b r o a d e r . A l t h o u g h f l e e i n g t o h i g h e r and more rugged t e r r a i n was the most common response t o d i s t u r b a n c e s among t a r u c a s , t h e r e were some e x c e p t i o n s . On f o u r o c c a s i o n s , a l l i n v o l v i n g l a r g e s o l i t a r y males, the a n i m a l s d i d not escape but e i t h e r walked a s h o r t d i s t a n c e away or s i m p l y d i d not move. In a l l c a s e s the males e v e n t u a l l y l a y down.in f u l l v i e w , s t a r i n g c a l m l y a t me. I d i d not attempt t o approach t h e s e males c l o s e r than 50 m. On f o u r o c c a s i o n s , females r e a c t e d i n a v e r y d i f f e r e n t manner. U n l i k e l a r g e s o l i t a r y males, the females i n q u e s t i o n e v e n t u a l l y f l e d but o n l y a f t e r much s t i f f - w a l k i n g , t a i l -f l i c k i n g , and s n e e z i n g . A l t h o u g h two of t h e s e females were d i s t u r b e d d u r i n g the fawning season, I was unable t o d i s c o v e r any fawns i n the v i c i n i t y . V e r y young fawns were never observed among groups t h a t f l e d t o h i g h e r t e r r a i n . The r e a c t i o n of a s m a l l fawn and her mother was o b s e r v e d once. Both were found i n an opening a few meters from a r o c k - o u t c r o p . As soon as they s p o t t e d me they ran t o t h e r o c k s . S h o r t l y a f t e r e n t e r i n g the o u t c r o p , the fawn h i d 96 b e h i n d a l a r g e r o c k , w h i l e the female stopped and s t a r e d a t me f o r s e v e r a l m i n u t e s . I d i d not attempt t o f o l l o w c l o s e r . Soon the female a l s o d i s a p p e a r e d i n the r o c k - o u t c r o p . A l t h o u g h I never approached a fawn a t v e r y c l o s e range, i t appears t h a t t h r e a t e n e d young fawns c r o u c h down and " f r e e z e " (D. J a r a , p e r s . c omm.). 2.3.5 C r y p t i c c o l o r a t i o n and h i d i n g b e h a v i o r of fawns B e s i d e from b e i n g the s m a l l e s t c l a s s , t h e i r u n s p o t t e d d a r k - g r e y c o a t s made fawns d i f f i c u l t t o spot among r o c k s or ro c k y background. Moreover, Hippocamelus fawns appeared t o have a " h i d i n g o u t " p e r i o d . D u r i n g o b s e r v a t i o n s of female groups i n the b i r t h season, fawns remained h i d d e n from view more o f t e n than d i d a d u l t s (X 2=31.2, p<0.00l) or y e a r l i n g s (X 2=14.0, P<0.001). I f h i d i n g b e h a v i o r i s pronounced d u r i n g the f i r s t days or weeks a f t e r b i r t h ( L e n t , 1974), then i t i s l i k e l y t h a t I was unable t o d e t e c t v e r y s m a l l fawns. Females began t o se g r e g a t e on 11 January 1981. B e f o r e 25 F e b r u a r y 1981, o n l y two fawns were s i g h t e d i n 2 out of 15 female groups observed d u r i n g t h i s p e r i o d . T h e r e a f t e r , fawns were obse r v e d i n most (79%) female groups. In c o n c l u s i o n , c o u g a r s , dogs, and f o x e s were p r e s e n t i n t he st u d y a r e a . The remains of a fawn suggested t h a t i t had been k i l l e d a cougar. The response of deer t o dogs i n d i c a t e d t h a t a d u l t t a r u c a s f l e e i n the presence of p r e d a t o r s . G e n e r a l l y , a l l t y p e s of groups c l i m b e d t o h i g h e r t e r r a i n when I approached them c l o s e l y ( e . g . l e s s than 50 m), but I never 97 o b s e r v e d v e r y young fawns among e s c a p i n g i n d i v i d u a l s . When f l e e i n g , the a n i m a l s d a r t e d toward r i d g e s or roc k y t e r r a i n where they u s u a l l y d i s a p p e a r e d from view. B e h a v i o r s such as s n e e z i n g , s t i f f - w a l k i n g , t a i l - f l i c k i n g , and w h i t e rump-patch exposure by t a r u c a s were commonly a s s o c i a t e d w i t h escape. These b e h a v i o r s may have been used t o a l e r t o t h e r s i n the group. In a few o c c a s i o n s , l a r g e males d i d not escape when I approached them. I t i s p o s s i b l e t h a t they d i d not r e g a r d me as a p o t e n t i a l p r e d a t o r , or t h a t they c o u l d have r e p e l l e d my approach had I come any c l o s e r . My o b s e r v a t i o n s of s m a l l fawns were l i m i t e d but suggest t h a t young h i d e i n response t o impeding danger. Fawns a l s o appeared t o go th r o u g h a " h i d i n g o u t " p e r i o d t h a t c o i n c i d e d w i t h the s e g r e g a t i o n of t h e i r mothers from groups c o n t a i n i n g a d u l t males. 98 2.4 DISCUSSION H a b i t a t s e g r e g a t i o n of the sexes i n Hippocamelus  a n t i s e n s i s took p l a c e d u r i n g the fawning season, a p e r i o d i n which a l l pregnant and l a c t a t i n g females s e p a r a t e d from groups t h a t c o n t a i n e d males. At t h i s t i m e , females moved away from l a r g e mixed-sex groups t h a t o c c u p i e d r e l a t i v e l y open h a b i t a t . Once s e g r e g a t e d , they formed s m a l l e r groups w i t h o t h e r females and t h e i r young and o c c u p i e d s t e e p e r , r o c k i e r , and more broken h a b i t a t than t h a t of male or mixed-sex groups. Here, I f i r s t c o n s i d e r s e v e r a l hypotheses t o e x p l a i n why the sexes s e g r e g a t e d . Second, I b r i e f l y compare the t i m i n g of s e x u a l s e g r e g a t i o n of t a r u c a s t o t h a t of o t h e r c e r v i d s and t o mountain c a p r i d s . D i f f e r e n t i a l F o o d - s e l e c t i o n Most s t u d i e s on d i f f e r e n t i a l r e s o u r c e e x p l o i t a t i o n by the sexes i n u n g u l a t e s have f o c u s e d on food u t i l i z a t i o n (Bowyer, 1984; C h a r l e s e t a l . , 1977; C l u t t o n - B r o c k e t a l . , 1982; S t a i n e s et a l . , 1982; T a k a t s u k i , 1980; Watson and S t a i n e s , 1978). Some a u t h o r s have c o n c l u d e d t h a t s e x u a l s e g r e g a t i o n i s the r e s u l t of d i f f e r e n c e s i n food h a b i t s , and t h a t females occur i n a r e a s of b e t t e r f o r a g e q u a l i t y ( C l u t t o n - B r o c k and Harvey, 1983; C l u t t o n -Brock et a l . , 1982; G e i s t , 1974 a , b ) . D e s p i t e some e v i d e n c e a g a i n s t t h i s h y p o t h e s i s (Shank, 1982; McCullough, 1979), l i t t l e a t t e n t i o n has been p a i d t o o t h e r f a c t o r s . My r e s u l t s suggest t h a t the s h i f t i n h a b i t a t use by l a c t a t i n g t a r u c a s was not d e t e r m i n e d by f o r a g e q u a l i t y . F i r s t , 99 d u r i n g s e x u a l s e g r e g a t i o n , groups d i d not d i f f e r i n f r e q u e n c y of use of d i f f e r e n t v e g e t a t i o n t y p e s . Second, s h i f t s i n use of v e g e t a t i o n h a b i t a t s o c c u r r e d as t h e sexes a g g r e g a t e d , not when they s e g r e g a t e d . T h i r d , p l a n t biomass and p r o d u c t i v i t y a r e n e g a t i v e l y c o r r e l a t e d w i t h e l e v a t i o n (Holgado e t a l . , 1979), and l a c t a t i n g females p r e f e r r e d h i g h e r e l e v a t i o n s . F o u r t h , p l a n t c o v e r d e c r e a s e d w i t h g r e a t e r e l e v a t i o n , s l o p e , r o c k - c o v e r , number of r e l i e f f e a t u r e s , and d i s t a n c e t o water. I f females were s e l e c t i n g more p r o d u c t i v e a r e a s than those used by o t h e r groups, they s h o u l d have used l o w e r e l e v a t i o n s and l e s s s t e e p , l e s s r o c k y , and l e s s broken a r e a s t h a t l a y c l o s e r t o water s o u r c e s . T h i s s u g g e s t s t h a t f e m a l e s a c t u a l l y used a r e a s of poo r e r f o r a g e q u a l i t y . F i n a l l y , i t i s u n l i k e l y t h a t food s h o r t a g e or a r e d u c t i o n i n number and s i z e of h i g h q u a l i t y p a t c h e s were r e s p o n s i b l e f o r s p a t i a l s e g r e g a t i o n , s i n c e s e x u a l s e p a r a t i o n took p l a c e d u r i n g t h e growing season when food was most abundant and presumably more e v e n l y d i s t r i b u t e d . I t c o u l d be argued t h a t h i g h q u a l i t y v e g e t a t i o n , a l t h o u g h l e s s abundant, grew among r o c k s , but t h e r e a r e no d a t a t o support t h i s . Female groups d i d s e l e c t , as d i d o t h e r groups, s h o r t - g r a s s h a b i t a t over o t h e r v e g e t a t i o n - t y p e s . S h o r t - g r a s s h a b i t a t i n c r e a s e d w i t h g r e a t e r e l e v a t i o n and r o c k - c o v e r , but i t was a l s o a s s o c i a t e d w i t h t e r r a c e s . S i n c e male and mixed-sex groups, i n c o n t r a s t t o female g r o u p s , s e l e c t e d t e r r a c e s , one c o u l d a l s o argue t h a t t h i s s e l e c t i o n was the r e s u l t of p r e f e r e n c e f o r a r e a s w i t h s h o r t - g r a s s . To suggest t h a t f e m a l e s used h a b i t a t s of i n f e r i o r 100 f o r a g e r e l a t i v e t o males i s not f a r - f e t c h e d . Shank (1982), s t u d y i n g b i g h o r n sheep ( O v i s c a n a d e n s i s ) , and LaGory (1985), w o r k i n g on w h i t e - t a i l e d deer ( O d o c o i l e u s v i r g i n i a n u s ) , found t h a t females and young o c c u p i e d h a b i t a t s of p o o r e r food q u a l i t y than d i d a d u l t males. In b i g h o r n s , p o o r e r v e g e t a t i o n i n female ranges was a s s o c i a t e d w i t h b r o k e n , r o c k y c l i f f s . H i g h er q u a l i t y f o r a g e i n male ranges was a s s o c i a t e d w i t h more open s l o p e s . Shank (1982) c o n c l u d e d t h a t d i f f e r e n c e s i n sheep d i e t s s i m p l y r e f l e c t e d f o r a g e d i f f e r e n c e s i n the a r e a s used by the s e x e s , and t h a t s e x u a l s e g r e g a t i o n was caused by some unknown f a c t o r . From a study of h a b i t a t s t r u c t u r e and d i e t s i n mule deer ( O d o c o i l e u s  hemionus) , Mahgoub (1984) c o n c l u d e d t h a t o t h e r f e a t u r e s , such as topography, were more c r i t i c a l than f o o d r e s o u r c e s i n d e t e r m i n i n g s u i t a b l e deer h a b i t a t . S o c i a l Antagonism A second e x p l a n a t i o n i s t h a t s o c i a l antagonism between the sexes causes them t o s e g r e g a t e (LaGory, 1985; T a k a t s u k i , 1983). My o b s e r v a t i o n s do not support t h i s . I never o b s e r v e d a d u l t males a g g r e s s i v e l y e x c l u d e or d r i v e females away from a r e a s used by males. Females a l s o d i d not behave a g g r e s s i v e l y toward males. I b e l i e v e t h a t p r e g n a n t / l a c t a t i n g females chose t o move away from the a r e a s used by mixed-sex groups a t the time of s e g r e g a t i o n . There i s disagreement i n the l i t e r a t u r e r e g a r d i n g the r o l e of male/female i n t e r a c t i o n s i n s e x u a l s e g r e g a t i o n i n u n g u l a t e s . I f males use h a b i t a t s w i t h p o o r e r foods than 101 f e m a l e s , s o c i a l l y dominant males may move away from or a v o i d prime a r e a s so t h a t t h e i r mates and o f f s p r i n g can enjoy reduced c o m p e t i t i o n f o r food r e s o u r c e s ( G e i s t and P e t o c z , 1977). Where males occupy b e t t e r f o r a g i n g h a b i t a t s than females w i t h young, dominant males may be e x c l u d i n g females (LaGory, 1985). T a k a t s u k i (1983) proposed t h a t l a c t a t i n g females e x c l u d e a d u l t males from p r e f e r r e d a r e a s , w h i l e Bowyer (1984) and McCullough (1979) c o n c l u d e d t h a t male/female i n t e r a c t i o n s d i d not i n f l u e n c e s p a t i a l s e g r e g a t i o n of the sex e s . C l e a r l y , we need d e t a i l e d s t u d i e s of food p r e f e r e n c e s , f o r a g e q u a l i t y and a v a i l a b i l i t y , and s o c i a l b e h a v i o r of Hippocamelus t o a s s e s s p r o p e r l y the f o o d - s e l e c t i o n and s o c i a l -i n t e r a c t i o n h y p o theses. My s t u d y , however, s u g g e s t s t h a t a n o ther f a c t o r was i n v o l v e d . P r e d a t o r a v o i d a n c e An a l t e r n a t i v e h y p o t h e s i s i s t h a t l a c t a t i n g females s e l e c t e d a r e a s t h a t p r o v i d e d concealment t o t h e i r young d u r i n g the b i r t h season. One assumption of t h i s h y p o t h e s i s i s t h a t h a b i t a t s d i f f e r i n the ease of d e t e c t i o n of t a r u c a s by p r e d a t o r s . The t u n d r a - l i k e v e g e t a t i o n of La Raya p r o v i d e s l i t t l e or no c o v e r , but t o p o g r a p h i c f e a t u r e s o f f e r v a r y i n g degrees of concealment. Among t h e s e , r o c k - c o v e r seemed t o be most c r i t i c a l . F i r s t , i t p r o v i d e d h i d i n g s i t e s and c r y p t i c background f o r the fawns. Second, i t made u n d e t e c t e d approach v e r y d i f f i c u l t . T a r u c a s c o u l d scan t h e i r s u r r o u n d i n g s from r o c k - o u t c r o p s and thus spot a p p r o a c h i n g p r e d a t o r s . In c o n t r a s t , 1 02 r i d g e s and t r o u g h s , though they c o n c e a l e d the deer from c e r t a i n a n g l e s and l o n g d i s t a n c e s , a c t u a l l y made t h e deer v u l n e r a b l e t o c l o s e u n d e t e c t e d approaches. My c l o s e s t o b s e r v a t i o n s , w i t h o u t b e i n g d e t e c t e d by the a n i m a l s , were of deer i n s m a l l r a v i n e s or r i d g e - s i d e s . T e r r a c e s a l s o p r o v i d e d concealment f o r the deer from an o b s e r v e r l o c a t e d below. However, b e i n g exposed and f l a t a r e a s , t e r r a c e s made deer v e r y c o n s p i c u o u s from above. My r e s u l t s a r e c o n s i s t e n t w i t h the assumption t h a t h a b i t a t s d i f f e r i n p r o t e c t i o n o f f e r e d . Females w i t h young p r e f e r r e d v e r y r o c k y t e r r a i n , w h i l e they d i d not d i f f e r from mixed-sex groups i n r e g a r d t o r e l i e f of the t e r r a i n . In a d d i t i o n , female groups were found o c c u p y i n g t e r r a c e s l e s s f r e q u e n t l y than o t h e r groups. Another assumption of the p r e d a t o r a v o i d a n c e h y p o t h e s i s i s t h a t young fawns are more v u l n e r a b l e t o p r e d a t i o n than a r e a d u l t s , once t h e y have been d e t e c t e d . Though I have no d i r e c t e v i d e n c e , t h i s assumption seems r e a s o n a b l e . F i r s t , young fawns l a c k the stamina and a g i l i t y of o l d e r i n d i v i d u a l s t o f l e e from p r e d a t o r s ( C u r i o , 1976; S c h a l l e r , 1972) because neuromuscular development i s i n c o m p l e t e ( J a c o b s e n , 1984). Second, s t a m i n a i n f l e e i n g i s p r o b a b l y c r i t i c a l a t h i g h e l e v a t i o n s , where c o n c e n t r a t i o n of a t m o s p h e r i c oxygen i s s e v e r e l y reduced and a e r o b i c c a p a c i t y f o r extreme muscular a c t i v i t y i s l i m i t e d (Hochachka et a l . , 1983). F i n a l l y , young l a c k e x p e r i e n c e of t h e i r s u r r o u n d i n g s , and a c c i d e n t s i n mountainous t e r r a i n a r e common ( M u r i e , 1944; S c h a l l e r , 1977). Knowledge of the a r e a may be u s e f u l t o t r a v e r s e rugged t e r r a i n , 103 e s p e c i a l l y when an i n d i v i d u a l i s b e i n g chased by a p r e d a t o r . S e l e c t i v e p r e d a t i o n on young has been documented f o r many u n g u l a t e s ( r e v i e w by C u r i o , 1976). A t h i r d assumption i s t h a t d i f f e r e n t i a l v u l n e r a b i l i t y among age c l a s s e s has s e l e c t e d f o r d i f f e r e n t a n t i p r e d a t o r b e h a v i o r s . Taruca a d u l t s f l e e i n the pr e s e n c e of p r e d a t o r s , w h i l e I found t h a t young fawns p r o b a b l y h i d e and f r e e z e , l i k e most o t h e r c e r v i d s (De Vos, 1967; L e n t , 1974). The prone response of newborns t o a p p r o a c h i n g p r e d a t o r s i s uncommon a f t e r one week i n r e d d e e r , Cervus e l a p h u s ( C l u t t o n - B r o c k , 1982) , though i t may s t i l l be used by 3-week o l d e l k , Cervus c a n a d e n s i s (Altmann, 1963). Young c e r v i d s a l s o go t h r o u g h a h i d i n g phase c h a r a c t e r i z e d by l o n g p e r i o d s of s e p a r a t i o n from t h e i r mothers, d u r i n g which the young remain h i g h l y immobile ( L e n t , 1974). T h i s p e r i o d l a s t s about 10 days i n w h i t e - t a i l e d d e e r , O d o c o i l e u s  v i r g i n i a n u s ( J a c o b s e n , 1984), and 3 weeks i n muntjac, M u n t i a c u s  r e e v e s i , and roe d e e r , C a p r e o l u s c a p r e o l u s ( L e n t , 1974). P r e d a t o r a v o i d a n c e may be the most i m p o r t a n t f u n c t i o n of t h i s h i d i n g p e r i o d ( C l u t t o n - B r o c k e t a l . , 1982; L e n t , 1974). A l t h o u g h young may a c t i v e l y s e l e c t h i d i n g s i t e s ( C l u t t o n - B r o c k e t a l . , 1982; L e n t , 1974; S c h a l l e r , 1977), the h a b i t a t s chosen by females i n f l u e n c e the a v a i l a b i l i t y of co v e r f o r t h e i r young ( R i l e y and Dood, 1984). Among most mountain C a p r i n a e , females s e l e c t b r o k e n , r o c k y c l i f f s a t the time of p a r t u r i t i o n and remain i n t h i s t e r r a i n when " n u r s e r y " groups of females and young a r e formed ( G e i s t , 1971; M c F e t r i d g e , 1977; M u r i e , 1944). 1 04 The o n l y c e r v i d s p e c i e s where young do not h i d e i s c a r i b o u , R a n g i f e r t a r a n d u s (De Vos, 1967; L e n t , 1974), a h i g h l y m o b i l e and g r e g a r i o u s s p e c i e s t h a t l i v e s i n v e r y open h a b i t a t (Whitehead, 1972). Young c a r i b o u a r e v e r y p r e c o c i a l and f o l l o w t h e i r mothers from the f i r s t day. They cannot h i d e and do not f r e e z e i n response t o p r e d a t o r s (De Vos e t a l . , 1967), but use l a r g e groups of c o n s p e c i f i c s as p r o t e c t i o n ( L e n t , 1974). I n t e r e s t i n g l y , the c a r i b o u i s a l s o unique i n t h a t f e m a l e s , l i k e males, grow a n t l e r s (Whitehead, 1972). S i n c e c a r i b o u females do not s e g r e g a t e from males d u r i n g the c a l v i n g season, reduced s e x u a l dimorphism may have d e v e l o p e d t o a l l o w females t o compete f o r food s u c c e s s f u l l y w i t h males ( C l u t t o n - B r o c k and Harvey, 1978). I t i s e q u a l l y l i k e l y , however, t h a t a n t l e r s have deve l o p e d i n female c a r i b o u t o improve t h e i r a b i l i t y t o defend t h e i r young ( C l u t t o n - B r o c k et a l . , 1982). Some s p e c i e s where young do not h i d e from p r e d a t o r s have d e v e l o p e d group defence (e.g . muskox, Ovibos moschotus: L e n t , 1974; b u f f a l o , Syncerus  c a f f e r : S i n c l a i r , 1977). The l a s t assumption i s t h a t p r e d a t i o n has been a major r i s k f a c t o r f o r t a r u c a s . I b e l i e v e t h i s t o be the c a s e , but have l i t t l e d i r e c t e v i d e n c e . Cougars were p r e s e n t i n low numbers i n my study a r e a , and d i d k i l l one fawn. Andean f o x e s were common i n the study a r e a , and have been known t o p r e y on young v i c u n a s ( F r a n k l i n , 1974; K o f o r d , 1957). There were a l s o some 15 t o 20 s t r a y dogs a t La Raya, and t h e s e may be more imp o r t a n t p r e d a t o r s (Gavin e t a l . , 1984; Roe and Rees, 1975). E v i d e n c e l i n k i n g t he use of rugged, r o c k y t e r r a i n by 1 05 female t a r u c a s t o r e d u c t i o n i n p r e d a t i o n r i s k f o r fawns i s c i r c u m s t a n t i a l . Tarucas used t h i s t e r r a i n t o evade me when I approached or s t a r t l e d them from c l o s e d i s t a n c e , and a l s o t o escape from dogs. In mountain C a p r i n a e , rugged, rock c l i f f s and o u t c r o p s a r e used by a l l age c l a s s e s t o escape from p u r s u i n g p r e d a t o r s ( S c h a l l e r , 1977; Smith, 1983). G e i s t (1971) s t a t e s t h a t t h i s t e r r a i n i s f r e q u e n t l y used by s i c k and i n j u r e d a n i m a l s , presumably because th e s e a n i m a l s a r e more v u l n e r a b l e t o p r e d a t i o n . A d d i t i o n a l e v i d e n c e comes from M u r i e ' s (1944) s t u d i e s on D a l l sheep, O v i s d a l l i , and w o l v e s , C a n i s l u p u s . In the presence of heavy wolf p r e d a t i o n , sheep were r e s t r i c t e d t o rugged c l i f f a r e a s , and i n the absence of w o l v e s , sheep moved onto more open s l o p e s . A comparison between v i c u n a s and t a r u c a s can f u r t h e r i l l u s t r a t e the r e l a t i o n s h i p between a n t i p r e d a t o r b e h a v i o r and h a b i t a t s t r u c t u r e . On o c c a s i o n s , both s p e c i e s can be found i n a d j a c e n t a r e a s ( p e r s . o b s . ) , but t h e i r b e h a v i o r and h a b i t a t d i f f e r . V i c u n a s p r e f e r v e r y open and f l a t o r u n d u l a t i n g t e r r a i n ( F r a n k l i n , 1983). Though I found them on mountain s l o p e s a t La Raya, v i c u n a s d i d not occupy rugged and r o c k y t e r r a i n as much as t a r u c a s d i d . T h i s was p a r t i c u l a r l y o b v i o u s when the a n i m a l s escaped from d i s t u r b a n c e s . V i c u n a s r e l y on speed t o a v o i d p r e d a t o r s ( K o f o r d , 1957). Thus, v i c u n a s f l e d a l o n g mountain-s i d e s , a v o i d i n g r o c k - c l i f f s and o u t c r o p s , w h i l e t a r u c a s c l i m b e d up t o rugged t e r r a i n . The b e h a v i o r of young and h a b i t a t d u r i n g b i r t h a l s o d i f f e r i n both s p e c i e s . Young v i c u n a s can run as f a s t as t h e i r mothers w i t h i n a few hours a f t e r b i r t h , do not f r e e z e i n response t o p r e d a t o r s , and l a c k a h i d i n g p e r i o d 1 06 ( K o f o r d , 1957). I f n e c e s s a r y , female v i c u n a s mob and chase p r e d a t o r s away from young d u r i n g the b i r t h season ( F r a n k l i n , 1983). F i n a l l y , pregnant v i c u n a s do not seek rugged, r o c k y t e r r a i n d u r i n g p a r t u r i t i o n but g i v e b i r t h i n the open ( K o f o r d , 1957; p e r s . o b s . ) . I c o n c l u d e t h a t d u r i n g s e x u a l s e g r e g a t i o n , l a c t a t i n g t a r u c a females s e l e c t e d h a b i t a t s o f f e r i n g adequate concealment. I suggest t h a t s e l e c t i o n of these h a b i t a t s i s a s t r a t e g y t o reduce p r e d a t i o n r i s k f o r fawns. T h i s h y p o t h e s i s i l l u s t r a t e s how a s i n g l e f a c t o r , p r e d a t i o n r i s k , may r e s u l t i n d i f f e r e n t p a t t e r n s of h a b i t a t use i f v u l n e r a b i l i t y t o p r e d a t i o n v a r i e s w i t h age i n d i f f e r e n t h a b i t a t s (Werner e t a l . , 1983). T h i s h y p o t h e s i s a l s o e x p l a i n s the f o r m a t i o n of s m a l l e r groups d u r i n g p e r i o d s of food abundance, when g e n e r a l t h e o r y ( e.g. E s t e s , 1974) p r e d i c t s f o r m a t i o n of l a r g e r groups. D u r i n g s e x u a l s e g r e g a t i o n , l a c t a t i n g females might f u r t h e r d e c r e a s e t h e chances of b e i n g d e t e c t e d by p r e d a t o r s by f o r m i n g s m a l l e r groups as w e l l as s e l e c t i n g more c o n c e a l i n g h a b i t a t s . E x p e r i m e n t s i n v o l v i n g m a n i p u l a t i o n of r e s o u r c e s , t a r u c a s , or p r e d a t o r s a r e i m p r a c t i c a l or u n a d v i s a b l e s i n c e we a r e d e a l i n g w i t h a l a r g e and endangered s p e c i e s . However, t h e r e a r e a number of ways i n which d i f f e r e n t h y p otheses c o u l d be t e s t e d under n a t u r a l c o n d i t i o n s . For i n s t a n c e , i f f o o d p r e f e r e n c e s d i f f e r i n male and f e m a l e s , then t h e degree of s p a t i a l s e g r e g a t i o n of the sexes s h o u l d be a f u n c t i o n of t h e d i s p e r s i o n p a t t e r n of p r e f e r r e d p l a n t s . Thus, s e x u a l s e g r e g a t i o n ought t o be more pronounced i n a r e a s where p r e f e r r e d 1 07 items of each sex grow s e p a r a t e l y on d i f f e r e n t p o r t i o n s of the range. I f on the o t h e r hand, the sexes have s i m i l a r d i e t s but segr e g a t e t o reduce c o m p e t i t i o n , then s e p a r a t i o n s h o u l d be a f u n c t i o n of food abundance. T h e r e f o r e , s e x u a l s e g r e g a t i o n s h o u l d be more pronounced i n y e a r s or a r e a s where food i s s c a r c e or p o p u l a t i o n d e n s i t y h i g h . S i n c e a n i m a l s may not f o r a g e o p t i m a l l y i n the presence of p r e d a t o r s (Werner, e t a l . , 1983), p r e d a t o r abundance i n the stu d y a r e a s s h o u l d a l s o be m o n i t o r e d . I f p r e d a t i o n r i s k f o r young i s a major f a c t o r r e s p o n s i b l e f o r s e g r e g a t i o n of the s e x e s , the degree of s e p a r a t i o n s h o u l d be a f u n c t i o n of d i s p e r s i o n of c o v e r , v a r i a b i l i t y i n the concealment t h a t each h a b i t a t can o f f e r , and p r e d a t o r abundance. S e g r e g a t i o n of males and females s h o u l d be more pronounced i n a r e a s w i t h g r e a t e r h e t e r o g e n e i t y of cov e r h a b i t a t s and g r e a t e r p r e d a t o r abundance. R e s u l t s from d i f f e r e n t a r e a s and y e a r s c o u l d be used t o t e s t f o r c o n s i s t e n c y among p r e d i c t i o n s . F i n a l l y , s i n c e t h e s e s t u d i e s would be t e s t i n g a l t e r n a t i v e consequences of i n d i v i d u a l - h a b i t a t i n t e r a c t i o n s , q u a n t i t a t i v e i n f o r m a t i o n on y e a r l y e n e r g e t i c and n u t r i t i o n a l r e q u i r e m e n t s of males and f e m a l e s , and on s o c i a l , f o r a g i n g , and a n t i p r e d a t o r b e h a v i o r s of d i f f e r e n t age-sex c l a s s e s a re n e c e s s a r y t o un d e r s t a n d the mechanisms of s e x u a l s e g r e g a t i o n . Comparison w i t h Mountain C a p r i n a e T a r u c a s a r e unique among c e r v i d s but resemble mountain C a p r i n a e i n s o c i a l o r g a n i z a t i o n and h a b i t a t u t i l i z a t i o n . A major f e a t u r e of t a r u c a s o c i a l dynamics i s t h a t s e x u a l 108 s e g r e g a t i o n i s c o n f i n e d t o the b i r t h season. Thus, a d u l t males and females a r e found t o g e t h e r n e a r l y a l l y e a r , as a r e some mountain c a p r i d s ( e . g . b h a r a l , P s e u d o i s nayaur, A s i a t i c i b e x , Capra i b e x , w i l d g o a t , Capra nayaur, and chamois, R u p r i c a p r a  r u p r i c a p r a ; S c h a l l e r , 1977). T h i s p a t t e r n c o n t r a s t s t h a t of most s e a s o n a l b r e e d i n g c e r v i d s , where males and females r a r e l y i n t e r m i x o u t s i d e of the mating season (Bowyer, 1984; C l u t t o n -Brock et a l . , 1982; De Vos, 1967; H i r t h , 1977; T a k a t s u k i , 1983). The f a c t t h a t no o t h e r c e r v i d resembles Hippocamelus i n i t s c a p r i d - l i k e use of rugged and roc k y mountainous t e r r a i n , s u g g e s t s t h a t the s o c i a l o r g a n i z a t i o n of t h i s s p e c i e s has been i n f l u e n c e d by i t s u n u s u a l environment. The use of r e s t r i c t e d and l o c a l i z e d a r e a s by t a r u c a s y e a r - r o u n d may have f a c i l i t a t e d the f o r m a t i o n of mixed-sex groups o u t s i d e the ma t i n g season. A l t h o u g h more d e t a i l e d i n f o r m a t i o n i s needed f o r comparisons a c r o s s s p e c i e s of mountain C a p r i n a e , some e v i d e n c e suggest t h a t Capra s p e c i e s a r e more r e s t r i c t e d i n t h e i r y e a r - r o u n d movements and use a more s p e c i a l i z e d h a b i t a t than do O v i s s p e c i e s ( S c h a l l e r , 1977). In c o n t r a s t t o g o a t s , O v i s s p e c i e s show marked s e x u a l s e g r e g a t i o n n e a r l y a l l year ( G e i s t , 1971; S c h a l l e r , 1977). Thus, t a r u c a s resemble g o a t s more than sheep i n h a b i t a t and s o c i a l s t r u c t u r e . 109 APPENDIX A D u r i n g the s t u d y , 30 t a r u c a f e e d i n g - s i t e s were s e l e c t e d a t random from a l a r g e r sample. Each s i t e was c l a s s i f i e d as e i t h e r LONG-GRASS, SHORT-GRASS, or MARSH v e g e t a t i o n - t y p e (see s e c t i o n 2.2) On each s i t e , v e g e t a t i o n was sampled a l o n g two 15-m t r a n s e c t l i n e s which c r o s s e d at m i d - p o i n t a t r i g h t a n g l e s . The f o l l o w i n g t a b l e s (A1, A2, and A3) g i v e the mean number of s p e c i e s and l i s t i d e n t i f i e d s p e c i e s c o l l e c t e d from 11 LONG-GRASS , 12 SHORT-GRASS, and 7 MARSH s i t e s . U n l e s s o t h e r w i s e s p e c i f i e d , p l a n t s f o r which o n l y g e n e r i c name i s g i v e n r e p r e s e n t s i n g l e s p e c i e s . A l t h o u g h moss and l i c h e n s p e c i e s were not c o l l e c t e d , t h e i r p resence i n a s i t e i s n o t e d . T a b l e A1. P l a n t s p e c i e s i n LONG-GRASS v e g e t a t i o n - t y p e . SPECIES NAME FAMILY # SI Juncaceae 8 Gramineae 6 Ge r a n i a c e a e 6 Gramineae 6 Compositae g* Ge n t i a n a c e a e 6* Gramineae 4 Rosaceae 3 Juncaceae 3 Gramineae 3 Gramineae 3 C a r y o p h y l l a c e a e 2 Compositae 2 Gramineae 2 Gramineae 2 Gramineae 2 Cyperaceae 2 Gramineae 2 Gramineae 1 U m b e l l i f e r a e 1 Compositae 1 Gramineae 1 Gramineae 1 C a r y o p h y l l a c e a e 1 Gramineae 1 Compositae 1 Compositae 1 Compositae 1 Cactaceae 1 Leguminosae 1 Gramineae 1 Gramineae 1 P l a n t a g i a n a c e a e 1 Gramineae 1 C a r y o p h y l l a c e a e 1 I r i d a c e a e 1 Solanaceae 1 Leguminosae 1 V a l e r a n a c e a e 1 L u z u l a p e r u v i a n a  C a l a m a g r o s t i s a n t o n i a n a  Geranium sp S t i p a sp Gnaphalium spp G e n t i a n a spp C a l a m a g r o s t i s h e t e r o p h y l l a Alchemi11a p i n n a t a L u z u l a sp Poa gimnanta Poa p e a r s o n i i A r e n a r i a l a n u g i n o s a B a c c h a r i s sp Bromus l a n a t u s C a l a m a g r o s t i s vicunarum F e s t u c a d o l i c h o p h y l l a S c i r p u s r i g i d u s S t i p a i c h u A c i a c h n e p u l v i n a t a A z o r e l l a sp B a c c h a r i s sp C a l a m a g r o s t i s b r e v i f o l i a C a l a m a g r o s t i s c h r y s a n t h a C e r a s t i u m sp F e s t u c a r i g e s c e n s H i p o c h o e r i s s o n c h o i d e s H i p o c h o e r i s t a r a x a c o i d e s Liabum sp L o b i v i a sp Lu p i n u s m i c r o p h y l l u s Muhlembergia p e r u v i a n a O r y z o p s i s sp P l a n t a g o sp Poa annua P y c n o p h y l l u m sp S i s y r i n c h i u m j a m e s o n i i Solanum sp T r i f o l i u m sp V a l e r i a n a sp LICHEN MOSS 3 7 * Two s p e c i e s . MEAN # SPECIES PER SITE = 15.6+ 1.41 SE MEAN # UNIDENTIFIED SPP PER SITE = 5.3 + 1.09 SE T a b l e A2. P l a n t s p e c i e s i n SHORT-GRASS v e g e t a t i o n - t y p e . SPECIES NAME FAMILY # s Rosaceae 8 Juncaceae 6 Cyperaceae 6 Gramineae 5 Gramineae 5 Gramineae 4 Gramineae 3 Compositae 3 Compositae 3 Gramineae 3 Gramineae 2 Gramineae 2 Ge r a n i a c e a e 2 Gramineae 1 U m b e l l i f e r a e 1 Gramineae 1 Gramineae 1 Junacaceae 1 Gent ianaceae 1 Juncaceae 1 Gramineae 1 Gramineae 1 C a r y o p h y l l a c e a e 1 P a p i l i o n a c e a e 1 U r t i c a c e a e 1 A l c h e m i l l a p i n n a t a  L u z u l a p e r u v i a n a  S c i r p u s r i q i d u s  C a l a m a g r o s t i s vicunarum  Hordeum muticum  Bromus l a n a t u s  C a l a m a g r o s t i s h e t e r o p h y l l a  Gnaphalium sp H i p o c h o e r i s t a r a x i c o i d e s  S t i p a sp A c i a c h n e p u l v i n a t a  C a l a m a g r o s t i s a n t o n i a n a  Geranium s e s s i l i f l o r u m  Avena sp A z o r e l l a compacta  Bromus u n i o l o i d e s  D i s s a n t h e l i u m sp D i s t i c h i a muscoides  G e n t i a n a sp L u z u l a sp Poa annua  Poa gimnanta  Py c n o p h y l l u m m o l l e  T r i f o l i u m sp U r t i c a sp LICHEN MOSS 4 5 MEAN # SPECIES PER SITE = 11.8 + 1.28 SE MEAN # UNIDENTIFIED SPP PER SITE = 5.6 + 1.06 SE T a b l e A3. P l a n t s p e c i e s i n MARSH v e g e t a t i o n - t y p e . SPECIES NAME D i s t i c h i a muscoides  H i p o c h o e r i s sp C a l a m a g r o s t i s b r e v i f o l i a  A q r o s t i s sp  F e s t u c a d o l i c h o p h y l l a  L u z u l a p e r u v i a n a  P l a n t a g o sp C a l a m a g r o s t i s a n t o n i a n a  C a l a m a g r o s t i s c h r y s a n t h a  Poa sp D i s s a n t h e l i u m sp A z o r e l l a compacta C a l a m a g r o s t i s r i g e s c e n s Carex sp G e n t i a n a sp Poa gimnanta MOSS FAMILY # SITES Juncaceae 7 Compositae 6 Gramineae 5 Gramineae 4 Gramineae 4 Juncaceae 4 P l a n t a g i a n a c e a e 4 Gramineae 3 Gramineae 3 Gramineae 3 Gramineae 2 U m b e l l i f e r a e 1 Gramineae 1 Cyperaceae 1 Gentianaceae 1 Gramineae 1 7 MEAN # SPECIES PER SITE = 12.7 + 1.60 SE MEAN # UNIDENTIFIED SPP PER SITE = 5.4 + 1.07 SE 1 1 3 APPENDIX B T a b l e B1. P e r c e n t v e r t i c a l d i s t r i b u t i o n of fema l e , male, and mixed-sex groups (FEMALE, MALE, and MIXED-F) from January t h r o u g h A p r i l ; mixed-sex groups from May t h r o u g h August (MIXED-M); and mixed-sex groups from September t h r o u g h December (MIXED-S) i n the main study a r e a . The t h r e e 4-month p e r i o d s g i v e n above c o n t a i n the fa w n i n g , m a t i n g , and a n t l e r -s h e d d i n g seasons r e s p e c t i v e l y . Out of the f i v e a l t i t u d i n a l l e v e l s p r e s e n t , deer o c c u r r e d o n l y i n t h r e e : :LEVEL 2= 4,300-4,575m; LEVEL 3= 4,600-4,875m; and LEVEL 4= 4,900-5,175m. LEVEL GROUP-TYPE FEMALE MALE MIXED-F MIXED-M MIXED-S 100.0% 100.0% 100.0% 100.0% 100.0% N= 47 N= 42 N= 72 N= 49 N= 51 LEVEL 2 31.9 69.0 76.4 10.2 19.6 LEVEL 3 51.1 21.4 15.3 59.2 68.6 LEVEL 4 17.0 9.5 8.3 30.6 11.8 T a b l e B2. P e r c e n t v e r t i c a l d i s t r i b u t i o n of d i f f e r e n t deer group-types a t 3 time p e r i o d s d u r i n g the day. Out of the 5 a l t i t u d i n a l l e v e l s p r e s e n t i n the main study a r e a , deer groups o c c u r r e d i n t h r e e : LEVEL 2= 4,300-4,575m; LEVEL 3= 4,600-4,875m; and LEVEL 4= 4,900-5,175m. The time p e r i o d s c o n s i d e r e d a r e : PERIOD 1= 500-759hrs; PERIOD 2= 800-1059hrs; and PERIOD 3= 1100-1259hrs. Group-types a r e : FEMALE; MALE; MIXED-F ( f e m a l e , male, and mixed- sex groups from January t h r o u g h A p r i l ) ; MIXED-M (mixed-sex groups from May t h r o u g h A u g u s t ) ; and MIXED-S (mixed-sex groups from September t h r o u g h December). The t h r e e 4-month seasons g i v e n above c o n t a i n the f a w n i n g , m a t i n g , and a n t l e r - s h e d d i n g seasons r e s p e c t i v e l y . LEVEL GROUP-TYPE FEMALE MALE MIXED-F MIXED-M MIXED-S 100.0% 100.0% 100.0% 100.0% 100.0% PERIOD 1: N=1 1 N=7 N=20 N=2 N=1 4 LEVEL 2 27.3 100.0 95.0 0.0 28.6 LEVEL 3 63.6 0.0 0.0 100.0 71 .4 LEVEL 4 9.1 0.0 5.0 0.0 0.0 PERIOD 2: N=1 8 N=1 4 N=29 N=21 N=1 5 LEVEL 2 22.2 78.6 79.3 4.8 13.3 LEVEL 3 50.0 14.3 10.3 61 .9 66.7 LEVEL 4 27.8 7.1 10.3 33.3 20.0 PERIOD 3: N=1 5 N=1 6 N=1 7 N=1 9 N=1 2 LEVEL 2 33.3 50.0 52.9 5.3 8.3 LEVEL 3 53.3 31.3 35.3 52.6 66.7 LEVEL 4 13.3 18.8 1 1 .8 42. 1 25.0 116 T a b l e B3. P e r c e n t use of q u a d r a t s w i t h d i f f e r e n t s l o p e s by female, male and mixed-sex groups (FEMALE, MALE, and MIXED-F) from January t h r o u g h A p r i l ; mixed-sex groups from September th r o u g h August (MIXED-M); and mixed-sex groups from September th r o u g h December (MIXED-S) i n the main study a r e a . SLOPE FEMALE GROUP-MALE •TYPE MIXED-F MIXED-M MIXED-S 100.0% N= 47 1 00.0% N= 42 100.0% N= 72 100.0% N= 49 100.0% N= 51 <1 5 6.4 21.4 6.9 0.0 3.9 1 5-30 48.9 57. 1 75.0 69.4 41.2 30-45 44.7 21.4 18.1 30.6 54.9 T a b l e B4. P e r c e n t use of q u a d r a t s w i t h 0%, <50%, and >=50% r o c k - c o v e r by female, male, and mixed-sex groups (FEMALE, MALE, and MIXED-F) from January t h r o u g h A p r i l ; mixed-sex groups from May t h r o u g h August (MIXED-M); and mixed-sex groups from Septem-ber t h r o u g h December (MIXED-S) i n the main study a r e a . ROCK-COVER GROUP-TYPE FEMALE MALE MIXED-F MIXED-M MIXED-S 100.0% 100.0% 100.0% 100.0% 100.0.% N= 47 N= 42 N= 72 N=49 N= 51 0 0.0 9.5 11.1 2.0 5.9 <50 34.0 59.5 81.9 63.3 70.6 >=50 66.0 31.0 6.9 34.7 • 23.5 T a b l e B5. P e r c e n t use of q u a d r a t s w i t h v a r i o u s amounts of t o p o g r a p h i c a l r e l i e f by female, male, and mixed-sex groups (FEMALE, MALE, and MIXED-F) from January t h r o u g h A p r i l ; mixed-sex groups (MIXED-M) from May through August; and mixed-sex groups (MIXED-S) from September t h r o u g h December i n the main s t u d y a r e a . T o p o g r a p h i c a l r e l i e f was measured by c o u n t i n g the number of r i d g e s and t r o u g h s i n each q u a d r a t . RELIEF GROUP-•TYPE FEMALE MALE MIXED-F MIXED-M MIXED-S 100.0% 100.0% 100.0% 100.0% 100.0% N= 47 N= 42 N= 72 N= 49 N= 51 0-2 21 .3 38. 1 31.9 18.4 41.2 3-4 53.2 54.8 37.5 38.8 43. 1 >4 25.5 7.1 30.6 42.9 15.7 T a b l e B6. P e r c e n t use of t e r r a c e s (YES) by f e m a l e , male and mixed-sex groups (FEMALE, MALE, and MIXED-F) from J a n u a r y t h r o u g h A p r i l ; mixed-sex groups from May t h r o u g h August (MIXED-M); and mixed-sex groups from September t h r o u g h December (MIXED-S) i n the main study a r e a . TERRACE GROUP-TYPE FEMALE MALE MIXED-F MIXED-M MIXED-S 1 00.0% 100.0% 100.0% 100.0% 1 00.0% N= 47 N= 42 N= 72 N= 49 N= 51 NO 91.5 61 .9 63.9 85.7 62.7 YES 8.5 38. 1 36. 1 14.3 37.3 118 Table B7. P e r c e n t use of d i f f e r e n t v e g e t a t i o n t y p e s ( i n c l u d i n g a r e a s w i t h no v e g e t a t i o n ) by fe m a l e , male, and mixed-sex groups (FEMALE, MALE, and MIXED-F) from January through A p r i l ; mixed- sex groups from May t h r o u g h August (MIXED-M); and mixed-sex groups from September t h r o u g h December (MIXED-S) i n the main s t u d y a r e a . VEGETATION GROUP-TYPE TYPE FEMALE MALE MIXED-F MIXED-M MIXED-S 100.0% 100.0% 100.0% 100.0% 100.0% N= 47 N= 42 N= 72 N= 49 N= 51 LONG GRASS 68. 1 50.0 76.4 51 .0 76.5 MARSH 4.3 4.8 2.8 22.4 2.0 SHORT GRASS 25.5 35.7 18.1 4. 1 17.6 BARE GROUND 2.1 9.5 2.8 22.4 3.9 Table B8. P e r c e n t use of q u a d r a t s w i t h water s o u r c e s <=500m or >500m away by female, male, and mixed-sex groups (FEMALE, MALE, and MIXED-F) from January t h r o u g h A p r i l ; mixed-sex groups from May th r o u g h August (MIXED-M); and mixed-sex groups from September t h r o u g h December (MIXED-S) i n the main study a r e a . DISTANCE TO WATER FEMALE MALE GROUP-TYPE MIXED-F MIXED-M MIXED-S 100.0% N= 47 100.0% N= 42 100.0% N= 72 100.0% N= 49 100.0% N= 51 <=500 72.3 95.2 81.9 73.5 51.0 >500 27.7 4.8 18.1 26.5 49.0 T a b l e B9. H a b i t a t comparisons between female, male and mixed-sex groups d u r i n g the fawning season ( J a n - A p r i l ) i n the main study a r e a . VARIABLE COMPARISONS FEMALE-MALE FEMALE-MIXED MALE-MIXED ELEVATION X2 12.36 23.77 0.81 df = 2 P <.01 <. 0001 N.S. SLOPE X2 7.56 1 0.06 5.92 df = 2 P <.05 <.01 N.S. ROCK-COVER X2 13.10 48.31 1 1 .56 df= 2 P <.01 <.0001 <.01 RELIEF X2 6.61 3.02 8.73 df= 2 P <.05 N.S. <.05 TERRACE X2 11.14 1 1 .49 0.04 df= 1 P <. 00 1 <. 00 1 N.S. VEGET-TYPE X2 4.15 1 .26 8.73 df= 3 P N.S. N.S. <.05 DIST-WATER X2 8.30 1 .54 4.10 df= 1 P <.01 N.S. <.05 120 T a b l e B10. H a b i t a t comparisons between mixed-sex groups (MIXM) ob s e r v e d d u r i n g the mating season (May-Aug) and groups (FEMALE, MALE, MIXF) o b s e r v e d d u r i n g the fawning season (Jan-Apr) and mixed-sex groups (MIXS) d u r i n g the a n t l e r - s h e d d i n g season (Sep-Dec). VARIABLE COMPARISONS MIXM-FEMALE MIXM-MALE MIXM-MIXF MIXM-MIXS ELEVATION X2 7. 56 33 .50 51 .10 6. 05 df= 2 P < .05 < .0001 < .0001 < .05 SLOPE X2 6. 08 1 1 .76 5. 52 8. 97 df= 2 P < .05 < .01 N .S. < .05 ROCK-COVER X2 9. 83 2. 45 16 .54 2. 19 df= 2 P < .01 N .s. < .001 N .S. RELIEF X2 3. 29 1 5 .39 3. 29 10 .81 df= 2 P N .s. < .001 N .S. < .01 TERRACE X2 0. 79 6. 79 7. 00 6. 85 df= 1 P N .s. < .01 < .01 < .01 VEGET-TYPE X2 22 .54 19 .36 28 .43 22 .05 df= 3 P < .001 < .001 < .0001 < .001 DIST-WATER X2 0. 02 7. 78 1 . 24 5. 36 df= 1 P N .s. < .01 N .s. < .05 1 2 1 T a b l e B11. H a b i t a t comparisons between mixed-sex groups (MIXS) obser v e d d u r i n g the a n t l e r - s h e d d i n g season (Sep-Dec) and groups (FEMALE, MALE, MIXF) ob s e r v e d d u r i n g the fawning season ( J a n - A p r ) . VARIABLE COMPARISONS MIXS-FEMALE MIXS-MALE MIXS-MIXF ELEVATION X2 3.18 24.38 41 .29 df= 2 P N.S. <.0001 <. 0001 SLOPE X2 1.13 1 3.67 18.24 df= 2 P N.S. <.0l <. 001 ROCK-COVER X2 18.96 1.31 7.35 df= 2 P <. 00 1 N.S. <.05 RELIEF X2 4.74 2.12 3.66 df= 2 P N.S. N.S. N.S. TERRACE X2 1 1 .25 0.01 0.02 df= 1 P <. 00 1 N.S. N.S. VEGET-TYPE X2 1 .62 7.10 0.20 df= 3 P N.S. N.S. N.S. DIST-WATER X2 4.70 21 .90 13.41 df= 1 P <.05 <.0001 <. 001 1 22 APPENDIX C 123 T a b l e C1. L i s t of e n v i r o n m e n t a l v a r i a b l e s and the c l a s s e s used as "dummy" v a r i a b l e s f o r the s t e p w i s e m u l t i p l e d i s c r i m i n a n t a n a l y s i s . For each e n v i r o n m e n t a l v a r i a b l e one c l a s s does not appear as dummy v a r i a b l e and i s l i s t e d w i t h o u t a v a r i a b l e number. E n v i r o n m e n t a l v a r i a b l e s c o r r e s p o n d o n l y t o q u a d r a t s were deer were obs e r v e d . Thus, two e l e v a t i o n c l a s s e s (LEVEL 1= 4,075-4,275m; and LEVEL 5= 5,200-5,450m) do not appear i n the l i s t s i n c e no deer groups were observed i n t h e s e a l t i t u d i n a l l e v e l s i n the main study a r e a . ENVIRONMENTAL No. OF DUMMY CLASSES: DUMMY VARIABLES VARIABLE VARIABLE ELEVATION — 4,300-4,575 (m) 1 4,600-4,875 2 4,900-5,175 SLOPE -- <15 (degrees) 3 15-30 4 30-45 RELIEF (sum r i d g e s / t r o u g h s ) — 0-2 5 3-4 6 >4 ROCK-COVER -- 0 (%) 7 <50 8 > = 50 TERRACE — no 9 yes VEGETATION TYPE ~ bare ground 10 l o n g 11 marsh 12 s h o r t DISTANCE TO NEAREST — <500 WATER SOURCE (m) 13 >=500 1 2 4 T a b l e C2. W i t h i n - s e a s o n r e s u l t s from two-group s t e p w i s e d i s c r i m i n a n t a n a l y s e s c a r r i e d out w i t h "dummy" v a r i a b l e s . I n c l u d e d a r e p e r c e n t c o r r e c t c l a s s i f i c a t i o n , M a h a l a n o b i s d i s t a n c e (D-SQUARE) between groups, and b e s t d i s c r i m i n a n t v a r i a b l e s f o r each two-group a n a l y s i s . Groups a r e female (FEM), male (MAL), and mixed-sex (MIXF) observed d u r i n g the fawning season ( J a n - A p r ) . Best d i s c r i m i n a n t v a r i a b l e s a r e those t h a t were i n c l u d e d i n the d i s c r i m i n a n t f u n c t i o n s on the f i n a l s t e p of each a n a l y s i s . GROUPS USED IN ANALYSIS % CORRECT CLASSIFICATION D-SQUARE BEST DISCRIMINANTS * FEM/MAL 76.4 1 .66 ROCK-COVER [8] ELEVATION [1] TERRACE [9] ROCK-COVER [7] (11 (7 (5 (5 .61 ) .68) .69) .50) FEM/MIXF 84.9 3.98 ROCK-COVER [8] ELEVATION [1] RELIEF [6] (78 (19 (4 .03) .42) .74) MAL/MIXF 67.5 0.84 ROCK-COVER [8] RELIEF [6] (12 (8 .23) .72) * For each e n v i r o n m e n t a l v a r i a b l e , the c o r r e s p o n d i n g number, i n [ ] , and t h e F v a l u e , i n ( ) , f o r the dummy d i s c r i m i n a n t v a r i a b l e s a r e a l s o shown. 125 T a b l e C3. Between-season r e s u l t s from two-group s t e p w i s e d i s c r i m i n a n t a n a l y s e s c a r r i e d out w i t h "dummy" v a r i a b l e s . I n c l u d e d ar e p e r c e n t c o r r e c t c l a s s i f i c a t i o n , M a h a l a n o b i s d i s t a n c e (D-SQUARE) between group means, and be s t d i s c r i m i n a n t v a r i a b l e s f o r each two-group a n a l y s i s . Each group-type (female= FEM, male= MAL, and mixed-sex= MIXF) from the fawning season (Jan-Apr) i s d i s c r i m i n a t e d a g a i n s t mixed-sex groups (MIXM) from the mating season (May-Aug), and mixed-sex groups (MIXS) from the a n t l e r - s h e d d i n g season (Sep-Dec). GROUPS USED % CORRECT D-SQUARE BEST DISCRIMINANTS * IN ANALYSIS CLASSIFICATION FEM/MIXM 71 . 9 1 . 42 VEGETATION [12] (18.68) VEGETATION [10] (13.19) ROCK-COVER [8] (4.15) MAL/MIXM 86. 8 4. 30 ELEVATION | 1] (57.03) ELEVATION | 2] (29.13) VEGETATION [12] (24.65) ROCK-COVER [7] (5.29) MIXF/MIXM 81 . 8 3. 75 ELEVATION | 1] (46.66) ELEVATION I 2] (33.75) ROCK-COVER [8] (6.61 ) VEGETATION [11] (6.16) FEM/MIXS 78. 6 2. 73 ROCK-COVER [8] (16.63) TERRACE [9! (13.57) RELIEF [6] (12.10) SLOPE [4] (7.40) DISTWATER | 13] (5.85) RELIEF [5] (4.74) MAL/MIXS 89. 2 5. 79 ELEVATION [ 1] (54.25) SLOPE [4] (28.78) DISTWATER | 13] (25.24) SLOPE [3] (17.35) RELIEF [6] (14.81) ELEVATION I 2] (13.26) VEGETATION [11] (11.19) MIXF/MIXS 79. 7 2. 57 ELEVATION | 1] (46.95) DISTWATER | 13] (10.68) ROCK-COVER [8] (5.08) MIXM/MIXS 75. 0 1 . 89 RELIEF [6] (16.81 ) TERRACE [9\ (15.71 ) SLOPE [3] (8.58) RELIEF [5] (5.53) ELEVATION I 2] (4.81 ) For each e n v i r o n m e n t a l v a r i a b l e , the c o r r e s p o n d i n g number, i n [ ] , and the F v a l u e , i n ( ) , f o r the dummy v a r i a b l e s a r e a l s o shown. 1 2 6 127 REFERENCES Altmann, M. 1963. N a t u r a l i s t i c s t u d i e s of m a t e r n a l c a r e i n moose and e l k . In M a t e r n a l B e h a v i o r i n Mamma1s, e d i t e d by L.H. R h e i n g o l d , pp. 233-253. New York: W i l e y . Anderson, A.E. 1981. 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