UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

Social behaviour in a non-pupping colony of steller sea-lion (Eumetopias jubata) Harestad, Alton Sidney 1973

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Notice for Google Chrome users:
If you are having trouble viewing or searching the PDF with Google Chrome, please download it here instead.

Item Metadata

Download

Media
831-UBC_1973_A6_7 H36_4.pdf [ 3.57MB ]
Metadata
JSON: 831-1.0101262.json
JSON-LD: 831-1.0101262-ld.json
RDF/XML (Pretty): 831-1.0101262-rdf.xml
RDF/JSON: 831-1.0101262-rdf.json
Turtle: 831-1.0101262-turtle.txt
N-Triples: 831-1.0101262-rdf-ntriples.txt
Original Record: 831-1.0101262-source.json
Full Text
831-1.0101262-fulltext.txt
Citation
831-1.0101262.ris

Full Text

SOCIAL BEHAVIOUR IN A NON-PUPPING COLONY OF STELLER SEA-LION (EUMETOPIAS JUB ATA) by A l t o n S i d n e y H a r e s t a d B . S c , U n i v e r s i t y o f B r i t i s h C o l u m b i a , 1971 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n t h e De p a r t m e n t o f Z o o l o g y We a c c e p t t h i s t h e s i s a s c o n f o r m i n g t o t h e r e q u i r e d s t a n d a r d THE UNIVERSITY OF BRITISH COLUMBIA A u g u s t , 1973 i I n p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r t h e a d v a n c e d d e g r e e a t t h e U n i v e r s i t y o f f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r a g r e e t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by t h e Head o f my D e p a r t m e n t o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n , s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l make i t The U n i v e r s i t y o f B r i t i s h C o l u m b i a D e p a r t m e n t o f V a n c o u v e r 8, Canada i i ABSTRACT J j J l ^ i o c i i a s j u b a t a i s p o l y g y n c u s . The s e a - l i o n s s e g r e g a t e d u r i n g t h e b r e e d i n g s e a s o n i n t o p u p p i n g c o l o n i e s c o n s i s t i n g o f a d u l t males and f e m a l e s , and n o n - p u p p i n g c o l o n i e s c o n s i s t i n g m a i n l y o f s u b - a d u l t m a l e s . A l l p r e v i o u s s t u d i e s o f t h e s o c i a l b e h a v i o u r 1 o f E_. j u b a t a have been c o n c e r n e d o n l y w i t h p u p p i n g c o l o n i e s . T h i s s t u d y d e s c r i b e s t h e s o c i a l b e h a v i o u r and o r g a n i z a t i o n o f a n o n - p u p p i n g c o l o n y . The c o l o n y i s l o c a t e d a t M c l n n e s I s l a n d , B r i t i s h C o l u m b i a ( L a t . 52016* B. , Long. 128<>43» H.). I t c o n s i s t s o f 100-150 a n i m a l s . They a r e m a i n l y s u b - a d u l t males t h o u g h s e v e r a l a d u l t s e a - l i o n s o f b o t h s e x e s were p r e s e n t . Data c o n c e r n i n g s i x a g e - s e x c l a s s e s were t a k e n u s i n g two methods. One method i n v o l v e d t h e d e v e l o p m e n t o f an ethogram c o n s i s t i n g o f 34 b e h a v i o u r p a t t e r n s and t h e o b s e r v a t i o n o f s o c i a l i n t e r a c t i o n s . The o t h e r method i n v o l v e d s p a c i a l o r g a n i z a t i o n and a c t i v i t y . Q u a l i t a t i v e n o t e s on t e r r i t o r i a l and r e p r o d u c t i v e b e h a v i o u r a s w e l l as p o p u l a t i o n s t r u c t u r e were a l s o r e c o r d e d . A p e c k - d o m i n a n c e h i e r a r c h y e x i s t s between t h e c l a s s e s and i s r e l a t e d t o many o f t h e b e h a v i o u r s measured. Males a r e more s o c i a l l y i n v o l v e d t h a n f e m a l e s . T h i s i s more v o l u n t a r y f o r m a les t h a n i t i s f o r f e m a l e s . As males mature t h e i r b e h a v i o u r becomes more complex t h a n t h a t o f f e m a l e s . F e m a l e s a r e n o t a g g r e s s i v e . T h i s i s r e f l e c t e d by t h e i r l a c k o f Body C o n t a c t b e h a v i o u r . M a l e s a r e a g g r e s s i v e t o w a r d s o t h e r males I i i i and use more Body C o n t a c t b e h a v i o u r t h a n f e m a l e s . Body C o n t a c t b e h a v i o u r i s m a i n l y p h y s i c a l a g g r e s s i v e i n t e r a c t i o n . As males mature i t i s r e p l a c e d w i t h Non-body C o n t a c t b e h a v i o u r which i s l a r g e l y t h r e a t s and d i s p l a y s . The i n t e n s e s o c i a l c o n d i t i o n i n g o f s u b - a d u l t males r e s u l t s i n more s o c i a l l y a d e p t a d u l t s t h a t a r e b e t t e r a b l e t o cope w i t h the complex s o c i e t y a t p u p p i n g c o l o n i e s . S u b - a d u l t males a r e n o t t e r r i t o r i a l . However, some a d u l t males a r e . T h i s i s r e l a t e d t o t h e p r e s e n c e o f f e m a l e s . T e r r i t o r i e s a c t a s r e f u g e s f o r f e m a l e s who a v o i d a r e a s o f a c t i v i t y and h a r a s s m e n t by s e x u a l l y mature s u b - a d u l t males. T h i s c o n t r i b u t e s t o t h e g r o u p i n g o f f e m a l e s a r o u n d t e r r i t o r i a l a d u l t m a les. Non-pupping c o l o n i e s a r e s p a t i a l l y o r g a n i z e d s i m i l a r t o p u p p i n g c o l o n i e s e x c e p t f o r t h e r e l a t i v e p r o p o r t i o n o f age-sex c l a s s e s . S o c i a l l y , n o n - p u p p i n g c o l o n i e s a r e l e s s o r g a n i z e d t h a n p u p p i n g c o l o n i e s . S o c i a l o r g a n i z a t i o n i n I s . infes!3. i s promoted by t h e t r a n q u i l b e h a v i o u r o f a d u l t s and i n h i b i t e d by t h e d i s r u p t i v e b e h a v i o u r o f s u b - a d u l t males. i v TABLE OF CONTENTS A b s t r a c t i i L i s t Of T a b l e s v L i s t Of F i g u r e s , 1 v i i i I n t r o d u c t i o n 1 M a t e r i a l s and methods 2 R e s u l t s 16 T e r r i t o r i a l B e h a v i o u r 17 R e p r o d u c t i v e B e h a v i o u r 19 H i e r a r c h y 20 D u r a t i o n 24 I n i t i a t i o n 26 V a r i e t y Of B e h a v i o u r P a t t e r n s 27 B e g i n n i n g and T e r m i n a t i o n E e h a v i o u r 29 I n v e s t i g a t i v e B e h a v i o u r 39 Non-Body and Body C o n t a c t B e h a v i o u r 41 P o p u l a t i o n S t r u c t u r e 47 C l a s s A s s o c i a t i o n 49 A c t i v i t y 50 T r a n s i e n c y 52 I n t e r a c t i o n R a t e 53 I n t e r a c t i o n F r e q u e n c y 57 D i s c u s s i o n 59 Summary 76 Acknowledgements 79 L i t e r a t u r e C i t e d 80 V L I S T OF TABLES T a b l e Page, 1. D e t e r m i n a t i o n o f h i e r a r c h y u s i n g t h e r e t r e a t , 21 2. D e t e r m i n a t i o n d f h i e r a r c h y u s i n g i n i t i a t i o n o f i n t e r a c t i o n s . 23 3. V a r i e t y o f b e h a v i o u r p a t t e r n s used i n i n t r a - c l a s s b e h a v i o u r . 28 4. Use o f t h e Advance by a l l c l a s s e s . 30 5. Use o f t h e Advance by males t o o t h e r m a l e s . 31 6. Use o f t h e Advance by m a l e s t o L a r g e Cows. 32 7. Use o f t h e Advance by males t o Y e a r l i n g s . 32 8. Use o f t h e Advance by males t o S m a l l Cows. 33 9. The r e l a t i o n s h i p o f t h e Advance t o h i e r a r c h y . 33 10. Use o f t h e A t t a c k by m a l e s , f e m a l e s and Y e a r l i n g s . 35 11. Use o f t h e A t t a c k by males t o m a l e s . Y e a r l i n g s and f e m a l e s . 35 12. Use o f t h e A t t a c k by m a l e s . 36 13. Use o f t h e R e t r e a t by m a l e s , f e m a l e s and Y e a r l i n g s i n i n t r a - c l a s s b e h a v i o u r . 36 14. R e l a t i o n s h i p o f t h e R e t r e a t t o h i e r a r c h y . 37 v i 15- Use o f t h e Chase by m a l e s , f e m a l e s , and Y e a r l i n g s . 37 16. Use o f t h e Chase by males t o L a r g e Cows. 39 17. Use o f t h e Chase by males t o S m a l l Cows. 39 18. Use o f I n v e s t i g a t i v e b e h a v i o u r by m a l e s , f e m a l e s and Y e a r l i n g s i n i n t r a - c l a s s b e h a v i o u r . 40 19. Use o f I n v e s t i g a t i v e by males t o m a l e s , f e m a l e s and Y e a r l i n g s . 41 20. Use o f I n v e s t i g a t i v e b e h a v i o u r by f e m a l e s and Y e a r l i n g s t o m a l e s , f e m a l e s and Y e a r l i n g s . 41 21. Use o f Non-body and Body C o n t a c t b e h a v i o u r by a l l c l a s s e s i n i n t r a - c l a s s b e h a v i o u r . 43 22. Use o f Non-body and Body C o n t a c t b e h a v i o u r by males t o o t h e r c l a s s e s . 43 23. Use o f Non-body and Body C o n t a c t b e h a v i o u r by f e m a l e s t o o t h e r c l a s s e s . - 45 24. Use o f Non-body and Body C o n t a c t b e h a v i o u r by Y e a r l i n g s t o o t h e r c l a s s e s . 45 25. Use o f mouth by m a l e s , f e m a l e s and Y e a r l i n g s t o a l l c l a s s e s . 46 26. Use o f Mouth by male c l a s s e s t o a l l c l a s s e s . 46 27. Use o f M o u t h i n g by males and f e m a l e s t o a l l c l a s s e s . 47 28. Use o f M o u t h i n g by males t o a l l c l a s s e s . 47 v i i 29. Age-sex s t r u c t u r e o f a non-p u p p i n g c o l o n y . 48 30. I n t e r a c t i o n r a t e g i v e n a s i n t e r a c t i o n s p e r m i n u t e . 54 31. Time e a c h c l a s s s p e n d s i n t e r a c t i n g w i t h o t h e r s e a -l i o n s . 56 32. A c t i v e t i m e e a c h c l a s s s p e n d s i n t e r a c t i n g w i t h o t h e r s e a -l i o n s . 57 33. F r e q u e n c y o f male, f e m a l e , and Y e a r l i n g i n t e r a c t i o n s 59 34. I n t e r a c t i o n f r e q u e n c y o f L a r g e Cows w i t h m a l e s . 59 v i i i L I S T OF FIGURES F i g u r e Page 1. P e c k - d o m i n a n c e h i e r a r c h y o f c l a s s e s 24 2. Mean i n t e r a c t i o n d u r a t i o n ( i n s e c o n d s ) . 25 3. P e r c e n t o f i n t e r a c t i o n s i n i t i a t e d by t h e c l a s s on t h e o r d i n a t e when i n t e r a c t i n g w i t h t h e c l a s s e s on t h e a b s c i s s a . 27 4. A s s o c i a t i o n . 50 5. A c t i v i t y . 52 6. T r a n s i e n c y . 53 1 INTRODUCTION Euf f l e t q p i a s j u b a t a i s d i s t r i b u t e d from J a p a n , t h r o u g h t h e A l e u t i a n I s l a n d s , and down t h e c o a s t o f N o r t h A m e r i c a a s f a r s o u t h as C a l i f o r n i a . Members o f t h e s p e c i e s a r e g r e g a r i o u s and f o r m c o l o n i e s when t h e y h a u l o u t on l a n d . The c o l o n i e s a r e u s u a l l y l o c a t e d on remote and e x p o s e d i s l a n d s . I i j u b a t a i s p o l y g y n o u s . ftt c e r t a i n c o l o n i e s d u r i n g t h e b r e e d i n g s e a s o n t h e a d u l t m a l e s e s t a b l i s h t e r r i t o r i e s which t h e y m a i n t a i n d u r i n g l a t e s p r i n g and e a r l y summer. O n l y f e m a l e s and y e a r l i n g s c a n o c c u p y t h e s e a r e a s w i t h o u t h a r a s s m e n t from t h e t e r r i t o r y h o l d e r . Most o f t h e b r e e d i n g o c c u r s i n c o l o n i e s o r g a n i z e d i n t h i s way. These g r o u p s o f s e a - l i o n s w i l l be r e f e r r e d t o as p u p p i n g c o l o n i e s . The immature and a d u l t males u n a b l e t o e s t a b l i s h and m a i n t a i n t e r r i t o r i e s a r e n o t t o l e r a t e d w i t h i n p u p p i n g c o l o n i e s . These e x c l u d e d a n i m a l s f o r m n o n - p u p p i n g c o l o n i e s ( B e l k i n , 1966; P i k e , 1966) which c o n s i s t o f s u b - a d u l t males and a few a d u l t m a les and f e m a l e s . T h e r e have been s e v e r a l s t u d i e s on t h e b e h a v i o u r and s o c i a l o r g a n i z a t i o n o f E.. j u b a t a . I t s b e h a v i o u r and d i s t r i b u t i o n i n t h e K u r i l e I s l a n d s i s d e s c r i b e d by B e l k i n ( 1 9 6 6 ) . O r r and P o u l t e r (1966) g i v e a more d e t a i l e d a c c o u n t o f t h e s o c i a l o r g a n i z a t i o n o f a p u p p i n g c o l o n y i n C a l i f o r n i a . S a n d e g r e n (1970) d e s c r i b e s t h e b e h a v i o u r o f t h e f e m a l e s and young. t G e n t r y (1970) e m p h a s i z e s t h e s i g n i f i c a n c e o f male t e r r i t o r i a l i t y and f e m a l e d i s t r i b u t i o n i n the o r g a n i z a t i o n o f a p u p p i n g c o l o n y . 2 The a g g r e g a t i o n s o f n o n - b r e e d i n g males and y e a r l i n g s have o n l y been m e n t i o n e d a n e c d o t a l l y . The o b j e c t i v e o f t h i s s t u d y i s t o d e s c r i b e t h e b e h a v i o u r and s o c i a l o r g a n i z a t i o n o f a n o n - p u p p i n g c o l o n y . C o m p a r i s o n o f a n o n - p u p p i n g c o l o n y w i t h a p u p p i n g c o l o n y may f a c i l i t a t e a b e t t e r u n d e r s t a n d i n g s p e c i f i c a l l y o f t h e s o c i a l o r g a n i z a t i o n o f E. j u b a t a , and more g e n e r a l l y o f p o l y g y n y . The s o c i a l b e h a v i o u r o c c u r i n g i n i n t e r a c t i o n s between c l a s s e s o f JB. j u b a t a shows s i m i l a r i t i e s as w e l l as d i s t i n c t d i f f e r e n c e s . Much o f t h i s b e h a v i o u r c h a n g e s p r o g r e s s i v e l y a s t h e a n i m a l s g e t o l d e r . F e m ale b e h a v i o u r i s f o l l o w e d from Y e a r l i n g s t o S m a l l Cows and t h e n t o l a r g e Cows. Male b e h a v i o u r i s f o l l o w e d f r o m Y e a r l i n g s t o Young B u l l s and t h e n t h r o u g h t o O l d B u l l s . T r e n d s a r e d i s c u s s e d as t h e o n t o g e n y o f s o c i a l b e h a v i o u r . A number o f t h i n g s o c c u r a l o n g w i t h the o n t o g e n y o f s o c i a l b e h a v i o u r . The s e a - l i o n s g e t l a r g e r , s e x u a l l y m a t u r e , and become more e x p e r i e n c e d . These p r o c e s s e s a r e r e l a t e d t o and a f f e c t t h e b e h a v i o u r s measured. The b e h a v i o u r o f t h e c l a s s e s and t h e i r d e v e l o p m e n t g i v e p a t t e r n s o f s o c i a l b e h a v i o u r . T h e s e p a t t e r n s p r o v i d e a b a s i s f o r t h e d i s c u s s i o n o f s o c i a l o r g a n i z a t i o n . 3 MATERIALS AND METHODS j u b i i i i ! w i l l be r e f e r r e d t o as " s e a - l i o n " . T h i s t e r m i n o l o g y w i l l be used i n t h e r e m a i n i n g s e c t i o n s . The s e l e c t i o n o f a n o n - p u p p i n g c o l o n y was b a s e d on i n f o r m a t i o n f r o m a e r i a l s u r v e y s and Canada Department o f F i s h e r i e s r e c o r d s . The c o l o n y i s i n an a r e a c o n t a i n i n g many s m a l l e x p o s e d i s l a n d s and l o c a t e d n e a r M c l n n e s I s l a n d , B r i t i s h C o l u m b i a ( L a t . 5 2 « 1 6 » N . , Long. 1 2 8 ° 4 3 ' W . ) . The c o l o n y c o n s i s t s m a i n l y o f immature males and Y e a r l i n g s . M a t u r e males and f e m a l e s f o r m a minor p o r t i o n o f t h e c o l o n y . The summer p o p u l a t i o n s i n 1971 and 1972 were between 120 and 150 s e a - l i o n s . D u r i n g c e r t a i n p e r i o d s o f t h e y e a r most o f t h e c o l o n y l e a v e s and o n l y a few s e a - l i o n s r e m a i n . T h r e e b l i n d s were e r e c t e d on i s l a n d s n e i g h b o u r i n g t h e c o l o n y . The b l i n d s were from 50 t o 250 m e t e r s from t h e c o l o n y . O b s e r v a t i o n s were made f r o m t h e s e b l i n d s u s i n g b i n o c u l a r s (8x30 and 20x60) and a s p o t t i n g s c o p e (20 t o 60 po w e r ) . A l t h o u g h s ounds were e a s i l y h e a r d , t h e l o n g d i s t a n c e made i t d i f f i c u l t t o r e a d i l y l o c a t e t h e s o u r c e s . V o c a l b e h a v i o u r was n e g l e c t e d f o r t h i s r e a s o n . A p r e l i m i n a r y s t u d y c o n s i s t i n g o f g e n e r a l o b s e r v a t i o n s was s t a r t e d on Hay 9, 1972 and l a s t e d u n t i l May 27, 1972. D u r i n g t h i s t i m e , 6 a g e - s e x c l a s s e s and 34 b e h a v i o u r p a t t e r n s were d e t e r m i n e d . From May 28, t o J u l y 9, 1972, t e c h n i q u e s were d e v i s e d and more s t r u c t u r e d o b s e r v a t i o n s were t a k e n . 4 C l a s s e s The s e a - l i o n s i n t h e c o l o n y a r e d i v i d e d i n t o s i x c l a s s e s . A b r i e f d e s c r i p t i o n o f e a c h c l a s s i s g i v e n below. The age o f t h e a n i m a l s c o m p r i s i n g e a c h c l a s s i s e s t i m a t e d by c o m p a r i n g t h e r e l a t i v e s i z e s o f t h e c l a s s e s t o the growth c u r v e s c o m p i l e d by P i k e (1966). 1. O l d B u l l s - much l a r g e r t h a n a l l e t h e r c l a s s e s . 6 y e a r s and o l d e r . - l a r g e f o r e h e a d and v e r y s t o u t neck. t h i c k l o w e r jaw. l o n g mane. t h e t e s t e s a r e o b v i o u s . 2. I n t e r m e d i a t e B u l l s - l a r g e r t h a n Young B u l l s b u t s m a l l e r than O l d B u l l s . a t y p i c a l I n t e r m e d i a t e B u l l i s 2/3 the s i z e o f an O l d B u l l and 1 1/2 t o 2 ' t i m e s t h e s i z e o f a Young B u l l . 4 t o 6 y e a r s o l d . a c c e n t u a t e d f o r e h e a d and s t o u t n e c k . t h e l o w e r jaw i s n o t as t h i c k o r m a s s i v e a s t h a t o f an O l d B u l l , n o t i c e a b l e mane. t e s t e s a r e n o t as o b v i o u s as cn an 5 3. Young B u l l s 4. L a r g e Cows 5. S m a l l Cows O l d B u l l . 1/2 the s i z e o f I n t e r m e d i a t e B u l l s . 2 t o 4 y e a r s o l d . some i n d i c a t i o n o f an a c c e n t u a t e d f o r e h e a d . neck i s n o t s t o u t . t e s t e s a r e not r e a d i l y s e e n . mane i s s h o r t o r n o n - e x i s t a n t . l o w e r jaw i s n o t t h i c k . p e n i s i s v i s i b l e when t h e a n i m a l i s moving. a b o u t t h e s i z e o f Young B u l l s a l t h o u g h some L a r g e Cows a r e s l i g h t l y l a r g e r . 4 t o 5 y e a r s and o l d e r . s l i m neck. s l i m l o w e r jaw. no mane. smooth p r o f i l e . t e a t s n o t i c e a b l e , e s p e c i a l l y i f s u c k l i n g a young. 1/2 t h e s i z e o f L a r g e Cows. 2 t o 4 y e a r s o l d . s l i m n e c k . s l i m l o w e r jaw. no mane. 6 smooth p r o f i l e . 6. Y e a r l i n g s - 1/4 t o 1/3 t h e s i z e o f Young B u l l s o r L a r g e Cows. 1 y e a r o l d . no o b v i o u s s e c o n d sex c h a r a c -t e r i s t i c s . The above c l a s s e s do n o t form d i s t i n c t g r o u p s . The Young B u l l c l a s s g r a d e s i n t o t h e I n t e r m e d i a t e B u l l c l a s s . S e a - l i o n s t h a t a r e o r d i n a r i l y t o o l a r g e t o be Young B u l l s b u t t o o s m a l l t o be I n t e r m e d i a t e B u l l s a r e c l a s s e d as Young B u l l s . T h i s scheme i s a l s o a p p l i e d t o t h e o t h e r c l a s s e s . The Young B u l l and I n t e r m e d i a t e B u l l c l a s s e s a r e t h e most d i f f i c u l t t o s e p a r a t e . I f t h e sex o f a s e a - l i o n c a n n o t be d e t e r m i n e d , t h e n i t i s c l a s s e d as U n c l a s s i f i e d L a r g e o r U n c l a s s i f i e d S m a l l . O c c a s i o n a l l y n e i t h e r t h e sex nor s i z e o f a s e a - l i o n i s known, In t h e s e i n s t a n c e s i t i s r e c o r d e d as U n c l a s s i f i e d . O l d B u l l s and L a r g e Cows a r e s e x u a l l y mature, f e m a l e Y e a r l i n g s and S m a l l Cows a r e n o t ( P i k e , 1966). The age a t w hich males become s e x u a l l y mature i s not known. T e s t e s w e i g h t s a r e t h e same and sperm i s p r e s e n t i n t h e e p i d i d y m i d e s o f males aged s i x y e a r s and o l d e r ( T h o r s t e i n s o n and L e n s i k , 1962). T h e r e were no males y o u n g e r t h a n s i x y e a r s e l d i n t h e sample c o l l e c t e d . T h i s meant t h a t a t l e a s t t h e o l d e r I n t e r m m e d i a t e B u l l s a r e c a p a b l e o f p r o d u c i n g sperm. P e r l o v (1971) f o u n d i n d i v i d u a l s p e r m a t o z o a i n smears t a k e n from t h e e p i d i d y m i d e s o f males f i v e y e a r s and o l d e r . He a l s o f o u n d 7 t h a t i n f o u r y e a r o l d males, t h e s p e r m a t o z o a were a b s e n t . W h i l e t h e p r e s e n c e o f s p e r m a t o z o a i s n o t d e f i n a t i v e , t h i s , c o u p l e d w i t h t h e o b v i o u s s e x u a l d i m o r p h i s m , s u g g e s t s t h a t I n t e r m e d i a t e B u l l s a r e s e x u a l l y mature. Male Y e a r l i n g s and Young B u l l s a r e p r o b a b l y s e x u a l l y immature. The sex o f Y e a r l i n g s i s n o t d i s t i n g u i s h a b l e by o b v i o u s m o r p h o l o g i c a l c h a r a c t e r i s t i c s . The b e h a v i o u r s o f i n d i v i d u a l Y e a r l i n g s do n o t i n t u i t i v e l y seem d i f f e r e n t . U n t i l a t l e a s t two months o f a g e , pups o f b o t h s e x e s behave s i m i l a r l y ( F a r e n t i n o s , 1971). On t h e s e c r i t e r i a , t h e b e h a v i o u r o f male and f e m a l e Y e a r l i n g s i s assumed t o be t h e same. Y e a r l i n g b e h a v i o u r i s used as a b a s e from which male and f e m a l e b e h a v i o u r o r i g i n a t e s . B e h a v i o u r P a t t e r n s A t o t a l o f 34 b e h a v i o u r p a t t e r n s were d e t e r m i n e d . T h e s e a r e d i v i d e d i n t o f i v e g r o u p s ; B e g i n n i n g and T e r m i n a t i o n , I n v e s t i g a t i v e , Non-body C o n t a c t , Body C o n t a c t , and M i s c e l l a n e o u s . A l l t h e b e h a v i o u r p a t t e r n s were o b s e r v e d e x c e p t C o p u l a t i o n . B e g i n n i n g And T e r m i n a t i o n 1. Advance - A s e a - l i o n a p p r o a c h e s a n o t h e r . I t seems a s i f i t i s a p p r o a c h i n g w i t h i n t e n t . I t o c c u r s a t a s l e w o r medium s h u f f l e . 2. A t t a c k - T h i s i s s i m i l a r t o t h e Advance 3. Chase 4. fietreat I n v e s t i g a t i v e 5. N o s e i n g A 6. N o s e i n g B 7. N o s e i n g C 8. L i c k i n g A 9. L i c k i n g B Non-Body C o n t a c t 10. Mouth 11. S n o b b i n g A 8 e x c e p t t h a t i t o c c u r s v e r y q u i c k l y . - A s e a - l i o n c h a s e s o r f o l l o w s a n o t h e r . T h i s o c c u r s a t e i t h e r a f a s t r u n o r slow t o medium s h u f f l e . - A s e a - l i o n moves away f r o m a n o t h e r . T h i s a p p e a r s t o be a r e s u l t o f t h e o t h e r ' s b e h a v i o u r . - A s e a - l i o n p u t s i t s nose on o r n e a r t h e head, a n t e r i o r body o r back o f a n o t h e r s e a - l i o n . - A s e a - l i o n p u t s i t s n o s e cn o r n e a r t h e p e r i - a n a l r e g i o n o f a n o t h e r . - A s e a - l i o n p u t s i t s nose cn o r n e a r t h e g r o u n d where a n o t h e r has r e c e n t l y p a s t . - A s e a - l i o n l i c k s a n o t h e r cn t h e head, a n t e r i o r body, o r back. - A s e a - l i o n l i c k s a n o t h e r i n t h e p e r i - a n a l r e g i o n . - The mouth o f a s e a - l i o n i s opened and d i r e c t e d t o w a r d s a n o t h e r . - A s e a - l i o n s i t s w i t h i t s head on a 45 d e g r e e a n g l e . The e y e s a r e 9 12. S n o b b i n g B 13. T u r n Away 14. N o d d i n g Body C o n t a c t 15. H o u t h i n g 16. N e c k i n g A 17. N e c k i n g B 18. N e c k i n g C 19. N e c k i n g D sometimes c l o s e d , A s e a - l i o n s i t s w i t h i t s head on a 90 d e g r e e a n g l e . The e y e s a r e o f t e n c l o s e d . A s e a - l i o n moves s l i g h t l y back from t h e o t h e r s e a - l i o n and t h e n t u r n s i t s head away. A s e a - l i o n nods i t s h e a d , u s u a l l y s u c c e s s i v e l y . T h i s may be g u i t e a v i o l e n t o r r e l a t i v e l y s l o w a c t i o n . The mouths o f two i n t e r a c t i n g s e a -l i o n s a r e open and v e r y c l o s e t o g e t h e r . The v i b r i s s a e a r e e r e c t and sometimes d i r e c t e d f o r w a r d . T h e r e i s s h a k i n g and moving o f t h e head. T h e s e movements may be q u i t e r a p i d . A s e a - l i o n u s e s i t s own neck t o s t r o k e t h e neck o f a n o t h e r . A s e a - l i o n l i g h t l y p u s h e s a n o t h e r w i t h i t s neck. A s e a - l i o n v i g o r o u s l y p u shes a n o t h e r w i t h i t s neck and s h o u l d e r . A s e a - l i o n v i g o r o u s l y p u shes a n o t h e r w i t h i t s neck and 10 f r e q u e n t l y t r i e s to b i t e t h e o p p o n e n t s f o r e - f l i p p e r s . 20. B i t i n g A - A s e a - l i o n l i g h t l y b i t e s a n o t h e r . 21. B i t i n g B - A s e a - l i o n v i g o r o u s l y b i t e s a n o t h e r . T h i s a p p e a r s q u i t e v i c i o u s . 22., L u n g i n g - a s e a - l i o n , w i t h i t s mouth open, l u n g e s a t a n o t h e r . The jaws may snap s h u t . The v i b r i s s a e a r e e r e c t and e x t e n d e d f o r w a r d . M i s c e l l a n e o u s 23. Down 24. Ground 25. Moving 26. C h i n n i n g - A s e a - l i o n l e i s u r e l y l i e s down. I t s body does n o t have a s t r e t c h e d a p p e a r a n c e . The a n i m a l does n o t seem a l e r t . - A s e a - l i o n l i e s down w i t h t h e body w e i g h t m o s t l y i n t h e f o r e f U p p e r s . The neck i s e x t e n d e d and t o u c h i n g t h e g r o u n d . The body a p p e a r s t o be s t r e t c h e d . The c h i n may t o u c h the gr o u n d o r be r a i s e d s l i g h t l y . The a n i m a l seems a l e r t . - A s e a - l i o n i s w a l k i n g a t a slow o r medium r a t e . - A s e a - l i o n s u c c e s i v e l y n i p s t h e c h i n , t h r o a t , and neck o f a n o t h e r . 11 27. B a c k i n g 28. B e h i n d 29. S c r a t c h i n g 30. L o o k i n g 31. S u c k l i n g 32. a t t e m p t e d Mount 33. Mount 34. C o p u l a t i o n - D u r i n g an i n t e r a c t i o n a s e a - l i o n g oes t o t h e s i d e o f t h e e t h e r a n i m a l and t r i e s , t o l a y i t s head and neck on t h e o t h e r ' s b a c k . - D u r i n g an i n t e r a c t i o n one s e a - l i o n d e l i b e r a t e l y goes b e h i n d a n o t h e r . - A s e a - l i o n s c r a t c h e s i t s e l f . - A s e a - l i o n i s l o o k i n g a r o u n d . - A s e a - l i o n s u c k s t h e t e a t o f a n o t h e r . - A s e a - l i o n t r i e s t o mount a n o t h e r b u t f a l l s o f f a l m o s t i m m e d i a t e l y . - A s e a - l i o n mounts a n o t h e r . - A s e a - l i o n c o p u l a t e s w i t h a n o t h e r . T y p e s o f Data C o l l e c t e d A f t e r t h e p r e l i m i n a r y o b s e r v a t i o n s , methods were d e v i s e d t o c o l l e c t d a t a p e r t i n e n t t o t h e s t u d y . The t e c h n i q u e s and t h e i n f o r m a t i o n t h e y y i e l d e d a r e d e s c r i b e d i n t h i s s e c t i o n . 1. E n c o u n t e r Method The b e h a v i o u r s o f t h e v a r i o u s c l a s s e s a r e d e t e r m i n e d u s i n g what w i l l be r e f e r r e d t o as t h e E n c o u n t e r Method. A s e a - l i o n i s s e l e c t e d from t h e c o l o n y and watched f o r a p p r o x i m a t e l y 30 m i n u t e s . The f o l l o w i n g i n f o r m a t i i c n i s 12 r e c o r d e d : 1. C l a s s o f t h e s e a - l i o n b e i n g o b s e r v e d . 2. C l a s s e s o f t h e s e a - l i o n s i t i n t e r a c t e d w i t h . 3. D u r a t i o n o f t h e i n t e r a c t i o n s . 4. Which s e a - l i o n i n i t i a t e d t h e i n t e r a c t i o n . 5. T y p e s o f b e h a v i o u r p a t t e r n s used i n t h e i n t e r -a c t i o n s and t h e i r s e q u e n c e o f use. The E n c o u n t e r Method g i v e s a v a r i e t y o f i n f o r m a t i o n . T h i s i s o u t l i n e d b e l o w . H i e r a r c h y A c l a s s h i e r a r c h y c a n be d e t e r m i n e d by u s i n g c r i t e r i a w hich s u g g e s t d o m i n a n t - s u b o r d i n a n t r e l a t i o n s h i p s . D u r a t i o n The mean d u r a t i o n o f i n t e r a c t i o n s c a n be d e t e r m i n e d . T h i s a l l o w s c o m p a r i s o n s between t h e l e n g t h o f t h e t i m e t h e c l a s s e s t e n d to i n t e r a c t . The l e n g t h o f an i n t e r a c t i o n i s i n d i c a t i v e o f t h e amount o r k i n d c f i n f o r m a t i o n b e i n g e x c h a n g e d ( C a l d w e l l , 1 9 72). The d u r a t i o n o f i n t e r a c t i o n s can be u s e d t o c a l c u l a t e t h e p r o p o r t i o n o f t i m e a c l a s s s p e n d s i n t e r a c t i n g w i t h o t h e r c l a s s e s . I n i t i a t i o n The c l a s s e s o f t h e s e a - l i o n s which i n i t i a t e d i n t e r a c t i o n s a r e r e c o r d e d . T h i s i s used as a c r i t e r i o n t o d e t e r m i n e c l a s s h i e r a r c h y . I t may a l s o i n d i c a t e i f c l a s s e s 13 have p r e f e r e n c e s w i t h whom t h e y i n t e r a c t . I n t r a And I n t e r C l a s s B e h a v i o u r The t y p e s o f b e h a v i o u r p a t t e r n s employed and t h e number o f t i m e s t h e y were used a r e r e c o r d e d . T h i s i n f o r m a t i o n d e s c r i b e s t h e b e h a v i o u r o f one c l a s s t o w a r d s a n o t h e r . The i n t e r - c l a s s and i n t r a - c l a s s b e h a v i o u r s a r e c o n s i d e r e d f o r a l l b u t one g r o u p o f b e h a v i o u r p a t t e r n s . The M i s c e l l a n e o u s b e h a v i o u r p a t t e r n s a r e n o t d i s c u s s e d . T h i s i s b e c a u s e many a r e s e l d o m used and a r e c f minor s i g n i f i c a n c e . The d a t a a r e p r e s e n t e d i n a way t h a t b e s t shows t h e o n t o g e n y o f s o c i a l b e h a v i o u r . The t y p e s and r e l a t i v e f r e q u e n c y o f use o f c e r t a i n b e h a v i o u r p a t t e r n s i n d i c a t e the r e l a t i o n s h i p s e x i s t i n g between t h e c l a s s e s . T h e s e r e l a t i o n s h i p s w i l l h e l p d e s c r i b e t h e c o l o n y ' s s o c i a l o r g a n i z a t i o n . I n t e r a c t i o n R a t e S e a - l i o n s o f a p a r t i c u l a r c l a s s were o b s e r v e d and t h e number o f i n t e r a c t i o n s t h e y had were r e c o r d e d . An i n t e r a c t i o n r a t e i s c a l c u l a t e d f o r each c l a s s . T hese d a t a combined w i t h t h e D u r a t i o n d a t a a r e used t o c a l c u l a t e t h e p r o p o r t i o n o f t i m e a c l a s s i s a c t i v e . The p r o p o r t i o n o f t h i s t i m e t h a t a c l a s s i s i n t e r a c t i n g i s c a l c u l a t e d . The r e m a i n d e r o f t h e a c t i v e t i m e c a n be used as an e s t i m a t e o f t h e t i m e s p e n t d i s p l a y i n g a t no s p e c i f i c s o c i a l p a r t n e r o r u s i n g t h e L o o k i n g b e h a v i o u r p a t t e r n . 11 I n t e r a c t i o n F r e q u e n c y The number o f i n t e r a c t i o n s and c l a s s e s o f s e a - l i o n s w i t h which t h e o b s e r v e d c l a s s i n t e r a c t e d a r e known. The f r e q u e n c y w i t h which c l a s s e s i n t e r a c t i s c a l c u l a t e d . T h i s i s compared t o t h e e x p e c t e d f r e q u e n c y o f i n t e r a c t i o n b a s e d cn t h e a s s o c i a t i o n o f c l a s s e s (page 4 9 ) . F r e q u e n c y d a t a g i v e i n f o r m a t i o n on c l a s s s e l e c t i o n o f s o c i a l p a r t n e r s . 2. I n d i v i d u a l D i s t r i b u t i o n Method A s e r i e s o f o b s e r v a t i o n s were made c o n c u r r e n t l y w i t h t h e E n c o u n t e r Method. T h e s e o b s e r v a t i o n s w i l l be r e f e r r e d t o a s t h e I n d i v i d u a l D i s t r i b u t i o n Method. The t e c h n i q u e was d e s i g n e d t o d e t e r m i n e t h e a s s o c i a t i o n , a c t i v i t y , and t r a n s i e n c y o f t h e c l a s s e s . S e a - l i o n s were s e l e c t e d f r o m each c l a s s . T h e s e were a n i m a l s t h a t c o u l d be e a s i l y and r e p e a t e d l y r e c o g n i z e d . A c a r d f i l e was used i n which s c a r s and o t h e r i d e n t i f y i n g f e a t u r e s were r e c o r d e d . The c o l o n y was s e a r c h e d f o r t h e s e s e a - l i o n s a p p r o x i m a t e l y e v e r y h o u r . The f o l l o w i n g i n f o r m a t i o n was r e c o r d e d ; 1. Date and t i m e . 2. I n d i v i d u a l s l o o k e d f o r and t h o s e f o u n d . When a s e a - l i o n was f o u n d I f u r t h e r r e c o r d e d ; 3. I t s l o c a t i o n . 4. The c l a s s e s o f i t s f i v e n e a r e s t n e i g h b o u r s . N e i g h b o u r s g r e a t e r t h a n 3 body l e n g t h s away were not c o n s i d e r e d . In o n l y a few i n s t a n c e s d i d t h i s s i t u a t i o n a r i s e . 15 5. Which o f t h e s e a - l i o n s were a c t i v e . T h e s e d a t a r e s u l t i n i n f o r m a t i o n as o u t l i n e d below. A s s o c i a t i o n The a s s o c i a t i o n o f one c l a s s t o a n o t h e r i s f o u n d by t o t a l l i n g t h e number o f n e i g h b o u r s r e c o r d e d n e a r t h e i n d i v i d u a l s b e i n g o b s e r v e d and t h e n c a l c u l a t i n g the p r o p o r t i o n o f t h e s e t h a t b e l o n g t o e a c h c l a s s . T h i s d e t e r m i n e d whether t h e c l a s s e s were clumped o r r a n d o m l y d i s t r i b u t e d . A c t i v i t y The p r o p o r t i o n o f t i m e a c l a s s i s a c t i v e i s c a l c u l a t e d by d i v i d i n g t h e number o f a c t i v e n e i g h b o u r s i n a c l a s s w i t h t h e t o t a l number o f n e i g h b o u r s i n t h a t c l a s s . A s e a - l i o n i s c o n s i d e r e d a c t i v e i f when o b s e r v e d , i t had i t s head r a i s e d . T r a n s i e n c y T r a n s i e n c y i s a measure o f a s e a - l i o n , s movements w i t h i n t h e c o l o n y . The i s l a n d was d i v i d e d i n t o 20 a r e a s . These a r e a s were e i t h e r n a t u r a l p l a t f o r m s o r b o r d e r e d by r o c k f a c e s and c r a c k s . I f a s e a - l i o n moved t o a d i f f e r e n t a r e a between o b s e r v a t i o n s t h i s was c o u n t e d a s one movement. T r a n s i e n c y i s c a l c u l a t e d by d i v i d i n g t h e number o f t i m e s a s e a - l i o n c l a s s moved by t h e number o f t i m e s i t was s i g h t e d . 16 3 . O t h e r D e s c r i p t i v e n o t e s . c o n c e r n i n g t e r r i t o r i a l and r e p r o d u c t i v e b e h a v i o u r were t a k e n . The a g e - s e x s t r u c t u r e o f t h e c o l o n y was a l s o r e c o r d e d . P r e s e n t a t i o n o f t h e Data The f o l l o w i n g a b b r e v i a t i o n s f o r t h e c l a s s e s a r e used i n f i g u r e s and t a b l e s : OB - O l d B u l l s IB - I n t e r m e d i a t e B u l l s YB - Young B u l l s YY - Y e a r l i n g s SC - S m a l l Cows LC - L a r g e Cows Most o f t h e d a t a a r e p r e s e n t e d i n c o n t i n g e n c y t a b l e s and h i s t o g r a m s . The c l a s s e s a r e o r d e r e d a c c o r d i n g t o age and sex i n d a t a where c l a s s i n t e r a c t i o n s a r e n e t i n v o l v e d . Data t h a t i n v o l v e c l a s s i n t e r a c t i o n s a r e shown one o f two ways. In t h e s e c t i o n on h i e r a r c h y , t h e c l a s s e s a r e o r d e r e d t o show t h e i r r e l a t i v e s t a t u s most e a s i l y . F o r t h e r e m a i n i n g d a t a a d i f f e r e n t f o r m a t i s u s e d . The c l a s s e s a r e g r o u p e d by s e x , i n c r e a s i n g i n age f r o m Y e a r l i n g s . T h i s makes d i f f e r e n c e s i n male and f e m a l e b e h a v i o u r more a p p a r e n t . 17 RESULTS T e r r i t o r i a l B e h a v i o u r T e r r i t o r i a l b o u n d a r y d i s p l a y s s i m i l a r t o t h o s e d e s c r i b e d by G e n t r y (1970) o c c u r i n t h e i n t e r a c t i o n s between t h e male c l a s s e s . I n t e r m e d i a t e and Young B u l l s o c c a s i o n a l l y e x h i b i t t h i s b e h a v i o u r b u t no t e r r i t o r y i s d e f e n d e d and no b o u n d a r i e s a r e r e c o g n i z e d . Some O l d B u l l s d e f e n d t e r r i t o r i e s . Two o f t h e O l d B u l l s f r e q u e n t i n g t h e c o l o n y were t e r r i t o r i a l . O l d B u l l #2 was more t e r r i t o r i a l l y i n c l i n e d t h a n O l d B u l l #1. O r i g i n a l l y O l d B u l l #2 o c c u p i e d an a r e a on t o p o f t h e i s l a n d and m a i n t a i n e d a t e r r i t o r y t h e r e . When O l d B u l l #2 was n o t on t h e i s l a n d . O l d B u l l #1 e v e n t u a l l y a c g u i r e d O l d B u l l #2's t e r r i t o r y . When O l d B u l l #2 r e t u r n e d . O l d B u l l #1 l e f t t h e a r e a u s u a l l y w i t h o u t any p h y s i c a l c o n t a c t w i t h O l d B u l l #2. T e r r i t o r i e s a r e l o o s e l y m a i n t a i n e d . Young and I n t e r m e d i a t e B u l l s a r e u s u a l l y e x c l u d e d . However on many o c c a s i o n s , t h e O l d B u l l l i e s down and t h e s u b - a d u l t males f i l t e r i n among t h e f e m a l e s and Y e a r l i n g s . The O l d B u l l i s o b l i v i o u s t o t h e i r p r e s e n c e . E v e n t u a l l y t h e O l d B u l l g e t s up, nods h i s head, v o c a l i z e s and a p p r o a c h e s t h e o t h e r m a l e s . The o t h e r m a l e s q u i c k l y l e a v e t h e t e r r i t o r y . Any male t h a t e n t e r s t h e t e r r i t o r y and s u b s e q u e n t l y l i e s down a l s o l e a v e s . F e m a l e s and Y e a r l i n g s a r e n o t a g g r e s s i v e l y c h a s e d from t h e t e r r i t o r y by t h e O l d B u l l . 18 F e m a l e s were s e e n e a r l y i n t h e s t u d y on t h e t o p o f the i s l a n d where O l d B u l l #2 had e s t a b l i s h e d a t e r r i t o r y . L a t e r f e m a l e s were s e e n i n a s m a l l a r e a on t h e i s l a n d ' s n o r t h - e a s t end where O l d B u l l #2, a f t e r a b a n d o n i n g h i s p r e v i o u s t e r r i t o r y , e s t a b l i s h e d a new t e r r i t o r y . O l d B u l l #1 o c c a s i o n a l l y o c c u p i e d t h e s e a r e a s as p r e v i o u s l y d e s c r i b e d . When O l d B u l l #1 was not on e i t h e r o f t h e above a r e a s , he d i d not e s t a b l i s h a t e r r i t o r y e l s e w h e r e , b u t j o i n e d the g r o u p s o f Young and I n t e r m e d i a t e B u l l s . D u r i n g t h e s e t i m e s O l d B u l l #1 d i d n o t c h a s e t h e s u b a d u l t males from around him a l t h o u g h he o c c a s i o n a l l y i n t e r a c t e d w i t h them. F e m a l e s and Y e a r l i n g s o c c u p y i n g t h e b e s t m a i n t a i n e d t e r r i t o r y , a r e n o t a g g r e s s i v e l y h a r a s s e d , v b u t males a r e . T h i s t e n d s t o c r e a t e a r e f u g e f o r f e m a l e s and Y e a r l i n g s . F o r example on June 25, a L a r g e Cow was l y i n g down nea r t h e i s l a n d ' s n o r t h - e a s t end. An I n t e r m e d i a t e B u l l a p p r o a c h e d h e r and p u t h i s nose n e a r h e r p e r i - a n a l r e g i o n . She moved away as he p e r s i s t e d . The L a r g e Cow a f t e r b e i n g s u b j e c t e d t o t h i s b e h a v i o u r f o r s e v e r a l m i n u t e s moved 10 f e e t c l o s e r t o t h e O l d B u l l . She was now w i t h i n 10 t o 15 f e e t f r o m t h e O l d B u l l . The I n t e r m e d i a t e B u l l f o l l o w e d t h e f e m a l e f o r s e v e r a l f e e t . He t h e n l a y down and d i d n o t a p p r o a c h any f u r t h e r . I t i s p o s s i b l e t h a t t h e I n t e r m e d i a t e B u l l r e c o g n i z e d t h e t e r r i t o r i a l b o u n d a r y and so d i d n o t p u r s u e t h e f e m a l e . The L a r g e Cow l a y down a g a i n . 19 A s i m i l a r i n s t a n c e o c c u r e d on June 20. An I n t e r m e d i a t e B u l l a t t e m p t e d t o mount an U n c l a s s i f i e d S m a l l n e a r the t e r r i t o r y o f O l d B u l l #2. The U n c l a s s i f i e d S m a l l moved away and went c l o s e r t o w a r d s t h e O l d B u l l . The I n t e r m e d i a t e B u l l d i d not a t t e m p t t o f o l l o w i t . R e p r o d u c t i v e B e h a v i o u r No pups were b o r n a t M c l n n e s I s l a n d i n 1972. However s e v e r a l f e m a l e s s u c k l e d Y e a r l i n g s . I n v o l v e d s e x u a l b e h a v i o u r was n o t common. No c o p u l a t i o n s were o b s e r v e d , a l t h o u g h p r e c o p u l a t o r y d i s p l a y s s i m i l a r t o t h o s e d e s c r i b e d by S a n d e g r e n (1970) o c c u r e d . In s e v e r a l i n s t a n c e s s e a - l i o n s mounted o r a t t e m p t e d t o mount o t h e r s on t h e c o l o n y . On June 16, an I n t e r m e d i a t e B u l l a t t e m p t e d t o mount an U n c l a s s i f i e d S m a l l . The U n c l a s s i f i e d S m a l l moved i n t o an O l d B u l l ' s t e r r i t o r y . The I n t e r m e d i a t e B u l l l a y down and d i d n o t p u r s u e i t . a l s o on J une 16, an I n t e r m e d i a t e B u l l mounted a S m a l l Cow. He was on t o p and h e l d h e r down w i t h h i s f o r e f l i p p e r s and neck as d e s c r i b e d by S a n d e g r e n (1970). T h e r e was some movements o f t h e male's p e l v i c r e g i o n . A f t e r a few m i n u t e s t h e S m a l l Cow s g u i r m e d away. The I n t e r m e d i a t e B u l l t h e n mounted a n e a r b y U n c l a s s i f i e d S m a l l . A s i m i l a r b ut l e s s i n v o l v e d s e q u e n c e o c c u r e d . On J une 20, an I n t e r m e d i a t e B u l l a t t e m p t e d t o mount an U n c l a s s i f i e d S m a l l . The U n c l a s s i f i e d S m a l l i m m e d i a t e l y s q u i r m e d away. The I n t e r m e d i a t e B u l l a p p r o a c h e d i t a g a i n . 20 The U n c l a s s i f i e d S m a l l moved i n t o O l d B u l l #2's t e r r i t o r y . The I n t e r m e d i a t e B u l l d i d n o t a t t e m p t t o f o l l o w i t but t r i e d t o mount a n o t h e r U n c l a s s i f i e d S m a l l i n t h e same a r e a . The U n c l a s s i f i e d S m a l l r e s i s t e d . A s e c o n d I n t e r m e d i a t e B u l l a p p r o a c h e d and t h e two males began i n t e r a c t i n g . The U n c l a s s i f i e d S m a l l was f o r g o t t e n . The f i r s t I n t e r m e d i a t e B u l l , w i t h h i s p e n i s e x t e n d e d , t h e n a t t e m p t e d t o mount a Young B u l l l y i n g n e a r b y but t h e Young B u l l moved away. H i e r a r c h y The R e t r e a t b e h a v i o u r p a t t e r n i s an i n d e x t o t h e r e l a t i v e s t a t u s o f t h e c l a s s e s . L e E o e u f and P e t e r s o n (1968) use t h e R e t r e a t f o r d e t e r m i n i n g dominance i n t h e n o r t h e r n e l e p h a n t s e a l ( H i r g u n g a a n ^ u s t i r o s t r i s ) . S u b o r d i n a n t s e a -l i o n s s h o u l d g i v e t h e R e t r e a t p a t t e r n more o f t e n t h a n t h e y r e c i e v e i t when t h e y i n t e r a c t w i t h dominant a n i m a l s ( T a b l e 1 ) . D u r i n g t h e I n t e r a c t i o n s o f two c l a s s e s , i f one c l a s s u s e s t h e R e t r e a t ; 1. More t h a n t h e o t h e r , t h e n i t i s c o n s i d e r e d s u b o r d i n a n t . 2. L e s s t h a n t h e o t h e r , t h e n i t i s c o n s i d e r e d d o m i n a n t . 3. E q u a l t o t h e o t h e r , t h e n i t i s c o n s i d e r e d a p e e r . A s e c o n d c r i t e r i o n c a n be u s e d t o d e t e r m i n e c l a s s h i e r a r c h y . S e a - l i o n s s h o u l d t e n d t c i n i t i a t e i n t e r a c t i o n s w i t h s u b o r d i n a n t s more t h a n t h e y do w i t h d o m i n a n t s . In 21 IB LC YB SC YY OB 2/23 0/35 0/8 0/1 0/14 IB 3/6 5/39 0/11 3/35 LC 0/1 0/0 0/2 YB 0/0 0/12 SC 0/2 T a b l e 1. D e t e r m i n a t i o n o f h i e r a r c h y u s i n g t h e r e t r e a t . The r a t i o i s t h e number o f i n t e r a c t i o n s where t h e R e t r e a t was used by t h e v e r t i c a l l y l i s t e d c l a s s e s o v e r t h a t used by t h e h o r i z o n t a l l y l i s t e d c l a s s e s . 22 T a b l e 2, i f one c l a s s i n i t i a t e d ; 1. Wore i n t e r a c t i o n s t h a n a n o t h e r , t h e n i t i s c o n s i d e r e d d o m i n a n t . 2. Fewer i n t e r a c t i o n s t h a n a n o t h e r , t h e n i t i s c o n s i d e r e d s u b o r d i n a n t . 3. An e q u a l number, t h e n i t i s c o n s i d e r e d a p e e r . O l d B u l l s a r e d o m i n a n t t o a l l c l a s s e s . I n t e r m e d i a t e B u l l s a r e s u b o r d i n a n t o n l y t o O l d B u l l s . Young B u l l s , S m a l l Cows, and L a r g e Cows a r e s u b o r d i n a n t t o O l d B u l l s and I n t e r m e d i a t e B u l l s . A l l c l a s s e s a r e d o m i n a n t t o Y e a r l i n g s . The h i e r a r c h y o f Young B u l l s , S m a l l Cows, and L a r g e Cows a r e n o t as w e l l d e l i n e a t e d as t h e o t h e r c l a s s e s . P a r t c f t h i s i s a r e s u l t o f s m a l l s a m p l e s . From t h e a v a i l a b l e d a t a . L a r g e Cows a r e s l i g h t l y d o minant t o Young B u l l s and p e e r s w i t h S m a l l Cows. Young B u l l s and S m a l l Cows a r e o f s i m i l a r s t a t u s . The c l a s s h i e r a r c h y i s r e p r e s e n t e d i n F i g u r e 1 . D u r a t i o n The male c l a s s e s i n t e r a c t f o r l o n g e r p e r i o d s t h a n do f e m a l e and Y e a r l i n g c l a s s e s ( F i g u r e 2 ) . I n t e r a c t i o n s i n v o l v i n g m ales o f t h e same c l a s s a r e l o n g e r t h a n t h o s e i n v o l v i n g o t h e r male c l a s s e s o r Y e a r l i n g s . O l d B u l l s i n t e r a c t w i t h s u b o r d i n a n t a n i m a l s f o r s h o r t e r i n t e r v a l s t h a n t h e y do w i t h p e e r s . I n t e r m e d i a t e B u l l s i n t e r a c t f o r s h o r t e r p e r i o d s w i t h s u b o r d i n a n t and dominant c l a s s e s t h a n t h e y do w i t h p e e r s . Young B u l l s show a t r e n d s i m i l a r t o O l d and I n t e r m e d i a t e B u l l s . T h a t i s , t h e d u r a t i o n d e c r e a s e s as t h e 23 IB LC YB SC YY OB 25/6 38/5 12/2 3/2 17/0 IB 16/4 80/20 32/1 71/6 LC 4/0 11/3 21/11 YB 1/0 40/5 SC 6/0 T a b l e 2. D e t e r m i n a t i o n o f h i e r a r c h y u s i n g i n i t i a t i o n o f i n t e r a c t i o n s . The r a t i o i s t h e number o f i n t e r a c -t i o n s i n i t i a t e d by t h e v e r t i c a l l y l i s t e d c l a s s e s o v e r t h o s e i n i t i a t e d by t h e h o r i z o n t a l y l i s t e d c l a s s e s . 24 l OB I B LC YY YY SC dominant s u b o r d i n a n t F i g . 1. P e c k - d o m i n a n c e h i e r a r c h y o f c l a s s e s 25 OB IB "D C o « YB C o 3 Q c o YY SC LC 100 0 0 n= 100 n~= 1 0 2 6 1 9 3 6 100 n= 12 100 192 47 100 n= 21 82 46 2 7 100 3 7 I 4 6 0= 27 167 101 84 37 23 100 2 8 2 0 13 n= 46 23 4 28 12 8 OB IB YB YY SC LC H= 66.6 p< .05 df= 5 H= 34.3 p< .05 rti df= 5 H= 89. 1 p< .05 df= 5 H= 17.6 p< .05 df= 5 H= 13.0 p< .05 df= 5 H= 30.9 p< .05 df= 5 F i g . 2. Mean i n t e r a c t i o n d u r a t i o n ( i n s e c o n d s ) . The i n t e r a c t i o n d u r a t i o n i s g i v e n f o r t h e c l a s s on t h e o r d i n a t e when i n t e r a c t i n g w i t h t h e c l a s s on t h e a b s c i s s a . 26 d i f f e r e n c e i n t h e s t a t u s between t h e i n t e r a c t i n g c l a s s e s i n c r e a s e s . The L a r g e Cow, S m a l l Cow, and Y e a r l i n g c l a s s e s do n o t show t h e t r e n d m e n t i o n e d a b o v e . O l d B u l l s and I n t e r m e d i a t e B u l l s i n t e r a c t f o r f a i r l y l o n g p e r i o d s w i t h L a r g e Cows. They i n t e r a c t f o r s h o r t p e r i o d s w i t h S m a l l Cows and Y e a r l i n g s . L a r g e Cows i n t e r a c t f o r l o n g e r p e r i o d s w i t h O l d and I n t e r m e d i a t e B u l l s t h a n t h e y do w i t h o t h e r c l a s s e s . I n i t i a t i o n The p r o p o r t i o n o f t h e i n t e r a c t i o n s i n i t i a t e d by each c l a s s i s g i v e n i n F i g u r e 3. H i e r a r c h y i s d e f i n e d u s i n g t h i s b e h a v i o u r and so t h e c l a s s o f t h e i n i t i a t o r i s r e l a t e d t o t h e r e l a t i v e s t a t u s o f t h e i n t e r a c t i n g c l a s s e s . S e a - l i o n s i n i t i a t e more o f t h e i r i n t e r a c t i o n s w i t h s u b o r d i n a n t s t h a n w i t h d o m i n a n t s . Young B u l l s do n o t i n i t i a t e any i n t e r a c t i o n s w i t h L a r g e Cows. T h i s c o n t r a s t s w i t h w i t h b e h a v i o u r o f O l d and I n t e r m e d i a t e B u l l s who i n i t i a t e upwards o f 15% o f t h e i r i n t e r a c t i o n s w i t h L a r g e Cows. V a r i e t y Of B e h a v i o u r P a t t e r n s M a l e s use a g r e a t e r v a r i e t y o f b e h a v i o u r p a t t e r n s t h a n f e m a l e s and Y e a r l i n g s ( T a b l e 3 ) . Y e a r l i n g s use a g r e a t e r v a r i e t y t h a n f e m a l e s . O n l y i n t r a - c l a s s b e h a v i o u r s a r e c o n s i d e r e d . T h i s removes any b i a s c a u s e d by c l a s s e s m i m i c i n g e a c h o t h e r . P a t t e r n s used more than 1% o f t h e t o t a l p a t t e r n s o b s e r v e d a r e used i n t h e c o n s t r u c t i o n o f T a b l e 3. 27 100 c CD u a. O B I B Y B Y Y SC LC X 2= 23.0 p< . 05 df= 5 5 4 3 X2 = P< df= 65. 4 05 4 OB IB YB YY SC LC F i g . 3. P e r c e n t o f i n t e r a c t i o n s i n i t i a t e d by t h e c l a s s on t h e o r d i n a t e when i n t e r a c t i n g w i t h t h e c l a s s e s on t h e a b s c i s s a . & The sample s i z e i s t o o s m a l l s o t h e d a t a a r e not i n c l u d e d . 28 OB IB YB YY SC LC B e g i n n i n g and 3 4 2 2 0 0 T e r m i n a t i o n I n v e s t i g a t i v e 0 0 1 1 1 1 Non-body C o n t a c t 5 4 3 3 3 2 Body C o n t a c t 5 7 6 0 1 1 M i s c e l l a n e o u s 2 4 4 4 2 2 T o t a l p a t t e r n s 15 19 16 10 7 6 us e d T a b l e 3. V a r i e t y o f b e h a v i o u r p a t t e r n s used i n i n t r a - c l a s s b e h a v i o u r . X2= 17.7, p<.05, df= 5 29 M a l e s and f e m a l e s d i f f e r m o s t l y i n B e g i n n i n g and T e r m i n a t i o n , Non-body C o n t a c t and Body C o n t a c t b e h a v i o u r . B e g i n n i n g and T e r m i n a t i o n B e h a v i o u r Advance The u s e o f t h e Advance b e h a v i o u r p a t t e r n i n a l l c l a s s e s a r e compared ( T a b l e 4 ) . M a l e s use t h e Advance much more t h a n f e m a l e s and Y e a r l i n g s . M a l e s a r e more s e l e c t i v e i n t h i s ( T a b l e 5 ) . O l d B u l l s use t h e Advance most f r e q u e n t l y w i t h O l d B u l l s and I n t e r m e d i a t e B u l l s , and s e l d o m wi t h Young B u l l s . I n t e r m e d i a t e B u l l s use t h e Advance most w i t h Young B u l l s but u s e i t a b o u t h a l f as o f t e n w i t h o t h e r I n t e r m e d i a t e B u l l s and s e l d o m w i t h O l d B u l l s . I t i s o c c a s i o n a l l y used i n i n t e r a c t i o n s w i t h I n t e r m e d i a t e B u l l s and most o f t e n i n t h e i n t r a - c l a s s b e h a v i o u r o f Young B u l l s . The f r e g u e n c y w i t h which males use t h e Advance w i t h L a r g e Cows i s g i v e n i n T a b l e 6. O l d B u l l s use t h i s p a t t e r n o n l y s l i g h t l y more t h a n I n t e r m e d i a t e B u l l s . Young B u l l s s e l d o m i f e v e r use t h e Advance w i t h L a r g e Cows. T h e r e i s no s i g n i f i c a n t d i f f e r e n c e i n t h e use o f t h i s b e h a v i o u r p a t t e r n by t h e male c l a s s e s when t h e y i n t e r a c t w i t h Y e a r l i n g s ( T a b l e 7) o r S m a l l Cows ( T a b l e 8). Use o f t h e Advance i s r e l a t e d t o t h e r e l a t i v e s t a t u s o f t h e a n i m a l s i n v o l v e d ( T a b l e 9 ) . A s e a - l i o n u s e s t h i s b e h a v i o u r p a t t e r n more when i n t e r a c t i n g w i t h a s u b o r d i n a n t t h a n i t does w i t h a d o m i n a n t . 30 OB IB YB YY SC LC advance 50 149 105 7 3 4 no Advance 116 343 292 227 35 135 f r e g u e n c y .301 . 302 .264 .030 .024 .029 o f use T a b l e 4. Use o f t h e Advance by a l l c l a s s e s . The number o f i n t e r a c t i o n s where t h e Advance d i d and d i d n o t o c c u r i s g i v e n f o r each c l a s s . X 2= 130.5, p<.05, df= 5 O l d B u l l s t o males OB IB YB Advance 5 12 1 no Advance 11 25 20 f r e q u e n c y .312 . 324 .050 o f use X2= 6.1, p<.05, df= 2 I n t e r m e d i a t e B u l l s t o males OB IB YB Advance 1 45 62 no Advance 35 151 46 f r e q u e n c y .027 .229 .574 o f use X2= 53.6, p<.05, df= 2 Young B u l l s t o males OB IB YB Advance 0 14 84 no Advance 14 97 134 f r e q u e n c y .0 .126 .385 o f use X 2= 32.5, p<.05, df= 2 T a b l e 5. Use o f t h e Advance by m a l e s t o o t h e r m a l e s . The number o f i n t e r a c t i o n s where t h e Advance d i d and d i d n o t o c c u r . 32 OB IB YB Advance 30 12 0 no Advance 18 11 4 f r e q u e n c y .625 .521 .0 o f use T a b l e 6. Use o f t h e Advance by males t o L a r g e Cows. The number o f i n t e r a c t i o n s when t h e Advance d i d and d i d n o t o c c u r i s g i v e n . X 2= 6.0, p<.05, af= 2 OB IB YB Advance 1 18 7 no Advance 36 74 42 f r e q u e n c y .027 .24.3 . 166 o f use T a b l e 7. Use o f t h e Advance by males t o Y e a r l i n q s . The number o f i n t e r a c t i o n s when t h e Advance d i d and d i d n o t o c c u r i s g i v e n . X 2= 5.64, p>.05, df= 2 ( 33 OB IB Advance 1 11 no Advance 6 26 f r e q u e n c y .142 .297 o f use T a b l e 8. -Use o f t h e Advance by males t o S m a l l Cows. The number o f i n t e r a c t i o n s when t h e Advance d i d and d i d n o t o c c u r i s g i v e n . p>.05 d o m i n a n t t o s u b o r d i n a n t s u b o r d i n a n t t o dominant Advance 157 25 no Advance 359 477 f r e q u e n c y .304 .049 o f use T a b l e 9. The r e l a t i o n s h i p o f t h e Advance t o h i e r a r c h y . The number o f i n t e r a c t i o n s when t h e Advance d i d and d i d n o t o c c u r i s g i v e n . X2= 112.2, p<.05, df= 1 3U A t t a c k The A t t a c k b e h a v i o u r p a t t e r n i s used o n l y by males ( T a b l e 1 0 ) . I t i s l e s s f r e q u e n t l y employed t h a n t h e Advance ( T a b l e 4 ) . The use o f t h e A t t a c k depends on t h e c l a s s o f t h e a n i m a l t h e male i s i n t e r a c t i n g w i t h ( T a b l e 1 1 ) . Males d i r e c t t h i s b e h a v i o u r p a t t e r n most f r e q u e n t l y t o o t h e r m a l e s , but n e v e r t o f e m a l e s and o n l y o c c a s i o n a l l y t o Y e a r l i n g s . The male c l a s s e s use t h e A t t a c k i n d i f f e r e n t f r e q u e n c i e s ( T a b l e 1 2 ) . O l d B u l l s use t h i s b e h a v i o u r p a t t e r n more f r e q u e n t l y t h a n I n t e r m e d i a t e B u l l s , and I n t e r m e d i a t e B u l l s use i t more f r e q u e n t l y t h a n Young B u l l s . R e t r e a t The R e t r e a t b e h a v i o u r p a t t e r n i n i n t r a - c l a s s b e h a v i o u r i s shown i n T a b l e 13. O n l y i n t r a - c l a s s b e h a v i o u r i s c o n s i d e r e d i n o r d e r t o r e d u c e b i a s t h a t would be i n t r o d u c e d by h i e r a r c h i c a l r e l a t i o n s h i p s . M a l e s use t h i s b e h a v i o u r p a t t e r n more t h a n f e m a l e s and Y e a r l i n g s . The R e t r e a t i s a c r i t e r i o n used t o d e t e r m i n e c l a s s h i e r a r c h y (page 20) and i s r e l a t e d t o s t a t u s ( T a b l e 1 4 ) . S u b o r d i n a n t s use t h i s a c t i o n more o f t e n than d o m i n a n t s . P e e r s use i t more t h a n d o m i n a n t s but l e s s t h a n s u b o r d i n a n t s . Chase The Chase b e h a v i o u r p a t t e r n i s used most f r e q u e n t l y by m a l e s ( T a b l e 1 5 ) . Male c l a s s e s use t h i s p a t t e r n d i f f e r e n t l y when t h e y i n t e r a c t w i t h L a r g e Cows ( T a b l e 1 6 ) . O l d and 35 f e m a l e s and males Y e a r l i n g s A t t a c k 73 0 no A t t a c k ' 982 461 f r e q u e n c y .074 .0 o f use T a b l e 10. Ose o f t h e A t t a c k by males, f e m a l e s and Y e a r l i n g s . The number o f i n t e r a c t i o n s when t h e A t t a c k d i d and d i d not o c c u r i s g i v e n . p<.05 A t t a c k no A t t a c k males 70 687 YY 3 175 f e m a l e s 0 120 f r e q u e n c y o f use 092 017 .0 T a b l e 11. Ose o f t h e A t t a c k by males t o m a l e s . Y e a r l i n g s and f e m a l e s . The number o f i n t e r a c t i o n s when t h e A t t a c k d i d and d i d not o c c u r i s g i v e n . X 2= 22.8, p<.05, df= 2 OB IB YB A t t a c k 18 42 13 no A t t a c k 93 390 379 f r e q u e n c y . 162 .097 .033 o f u s e T a b l e 12. Ose o f t h e A t t a c k by ma l e s . The number o f i n t e r a c t i o n s when t h e A t t a c k d i d and d i d n o t o c c u r i s g i v e n . X*= 24.3, p<.05, df= 2 f e m a l e s males Y e a r l i n g s R e t r e a t 86 1 no R e t r e a t 430 54 f r e q u e n c y .166 .018 o f use T a b l e 13. Use o f t h e R e t r e a t by mal e s , f e m a l e s and Y e a r l i n g s i n i n t r a - c l a s s b e h a v i o u r . The number o f i n t e r a c t i o n s when t h e R e t r e a t d i d and d i d n o t o c c u r i s g i v e n . p<.05 V 37 dominant t o p e e r s u b o r d i n a n t s u b o r d i n a n t to dom i n a n t R e t r e a t 13 87 189 no R e t r e a t 503 398 313 f r e q u e n c y .025 .179 .376 o f use T a b l e 14. R e l a t i o n s h i p o f t h e R e t r e a t t o h i e r a r c h y . The number o f i n t e r a c t i o n s when t h e R e t r e a t d i d and d i d not o c c u r i s g i v e n . X 2= 203.0, p<.05, df= 2 f e m a l e s and males Y e a r l i n g s Chase 94 4 no Chase 961 457 f r e g u e n c y .089 .008 o f use T a b l e 15. Use o f t h e Chase by m a l e s , f e m a l e s , and Y e a r l i n g s . The number o f i n t e r a c t i o n s when t h e Chase d i d and d i d n o t o c c u r i s g i v e n . X*= 34.3, p<.05, df= 1 38 I n t e r m e d i a t e B u l l s use i t , w h i l e Young B u l l s do n o t . O l d B u l l s use t h e Chase much more t h a n I n t e r m e d i a t e B u l l s . The male c l a s s e s do n o t show t h i s d i f f e r e n c e when t h e y i n t e r a c t w i t h S m a l l Cows ( T a b l e 1 7 ) . I n v e s t i g a t i v e B e h a v i o u r The o c c u r e n c e f r e g u e n c y o f I n v e s t i g a t i v e b e h a v i o u r i s c a l c u l a t e d ( T a b l e 1 8 ) . I n t r a - c l a s s b e h a v i o u r i s c o n s i d e r e d t o a v o i d b i a s . F e m a l e s and Y e a r l i n g s use more I n v e s t i g a t i v e b e h a v i o u r t h a n males. The i n t r a - c l a s s use o f t h i s b e h a v i o u r d e c r e a s e s as males mature. I n v e s t i g a t i v e b e h a v i o u r i s used by m a l e s more f r e q u e n t l y i n i n t e r a c t i o n s w i t h f e m a l e s and Y e a r l i n g s t h a n t h o s e w i t h o t h e r males ( T a b l e 1 9 ) . T h i s b e h a v i o u r i s d i r e c t e d more t o f e m a l e s t h a n i t i s t o Y e a r l i n g s . F emales and Y e a r l i n g s show a t r e n d s i m i l a r t o males ( T a b l e 2 0 ) . I n v e s t i g a t i v e b e h a v i o u r i s u s e d most f r e q u e n t l y i n i n t e r a c t i o n s w i t h f e m a l e s and Y e a r l i n g s t h a n w i t h males. Non-Body and Body C o n t a c t B e h a v i o u r The v a r i e t y o f Non-body and Body c o n t a c t b e h a v i o u r p a t t e r n s u s e d i s shown i n T a b l e 3. I f we c o n s i d e r Y e a r l i n g s as a b a s e , t h e o n t o g e n y o f s e a - l i o n b e h a v i o u r i s a p p a r e n t . Non-body C o n t a c t b e h a v i o u r p a t t e r n s a r e a b o u t t h e same f o r Y e a r l i n g s , f e m a l e s and Young B u l l s but as males mature the v a r i e t y i n c r e a s e s . OB IB YB Chase 28 3 0 no Chase 20 20 4 f r e q u e n c y .583 .130 .0 o f use T a b l e 16. Use o f t h e Chase by males t o L a r g e Cows. The number o f i n t e r a c t i o n s when t h e Chase d i d and d i d not o c c u r i s g i v e n . X2= 73.3, p<.05, df= 2 OB IB Chase 1 2 no Chase 6 35 f r e q u e n c y .166 .057 o f use T a b l e 17. Use o f t h e Chase by males t o S m a l l Cows. The number o f i n t e r a c t i o n s when t h e Chase d i d and d i d not o c c u r i s g i v e n . p>.05 40 OB IB YB Fem a l e s and Y e a r l i n g s I n v e s t i g a t i v e 0 11 32 14 N o n - i n v e s t i g a t i v e 79 834 1694 68 f r e g u e n c y .0 .013 .018 ,206 o f use T a b l e 18. Use o f I n v e s t i g a t i v e b e h a v i o u r by ma l e s , f e m a l e s and Y e a r l i n g s i n i n t r a -c l a s s b e h a v i o u r . The number o f I n v e s t i g a t i v e and N o n - i n v e s t i g a t i v e b e h a v i o u r p a t t e r n s a r e compared. X2= 94.9, p<.05, df= 3 4 1 i males YY f e m a l e s I n v e s t i g a t i v e 81 80 137 N o n - i n v e s t i g a t i v e 3729 301 286 f r e q u e n c y .021 .209 .323 o f use T a b l e 19. Use o f I n v e s t i g a t i v e by males t o males, f e m a l e s and Y e a r l i n g s . The number o f I n v e s t i g a t i v e and Non-i n v e s t i g a t i v e b e h a v i o u r p a t t e r n s a r e compared. X 2= 723.0, p<.05, df= 2 I n v e s t i g a t i v e N o n - i n v e s t i g a t i v e f r e q u e n c y o f use males f e m a l e s and Y e a r l i n g s 37 27 599 184 .058 .128 T a b l e 20. Use o f I n v e s t i g a t i v e b e h a v i o u r by f e m a l e s and Y e a r l i n g s t o m a l e s , f e m a l e s and Y e a r l i n g s . The number o f I n v e s t i g a t i v e and N o n - i n v e s t i g a t i v e b e h a v i o u r p a t t e r n s a r e compared. X2= 11.04, p<.05, df= 1 42 Y e a r l i n g s do not use Body C o n t a c t b e h a v i o u r . F e m a l e s use o n l y t h e t h e M o u t h i n g b e h a v i o u r p a t t e r n . Body C o n t a c t b e h a v i o u r i s a c q u i r e d by Young B u l l s and i s r e t a i n e d as t h e male m a t u r e s . The use o f Non-body and Body C o n t a c t b e h a v i o u r i n i n t r a -c l a s s b e h a v i o u r i s compared ( T a b l e 2 1 ) . Fe m a l e s and Y e a r l i n g s u s e more Non-body C o n t a c t t h a n Body C o n t a c t b e h a v i o u r . They a l s o use more Non-body C o n t a c t b e h a v i o u r t h a n do m a l e s . Young and I n t e r m e d i a t e B u l l s use l e s s Non-body C o n t a c t t h a n Body C o n t a c t b e h a v i o u r . T h i s c o n t r a s t s w i t h O l d B u l l s , who s i m i l a r t o f e m a l e s use more Non-body C o n t a c t b e h a v i o u r . As males mature, t h e amount o f Non-body C o n t a c t b e h a v i o u r i n c r e a s e s . M a l e s a r e s e l e c t i v e i n r e g a r d s t o which s e a - l i o n s t h e y d i r e c t Body C o n t a c t and Non-body C o n t a c t b e h a v i o u r ( T a b l e 2 2 ) . M a l e s use more Non-body C o n t a c t t h a n Body C o n t a c t b e h a v i o u r i n i n t e r a c t i o n s w i t h f e m a l e s and Y e a r l i n g s . They us e l e s s i n i n t e r a c t i o n s w i t h m ales. M a l e s use more Non-body C o n t a c t b e h a v i o u r when i n t e r a c t i n g w i t h f e m a l e s t h a n t h e y do w i t h Y e a r l i n g s . F e m a l e s use more Non-body C o n t a c t t h a n Body C o n t a c t b e h a v i o u r ( T a b l e 2 3 ) . T h e r e i s no d i f f e r e n c e i n t h e p r o p o r t i o n s o f Non-body C o n t a c t b e h a v i o u r d i r e c t e d t o d i f f e r e n t c l a s s e s . T h i s c o n t r a s t s w i t h t h e b e h a v i o u r o f t h e m a l e s . 43 OB Non-body C o n t a c t 44 Body C o n t a c t 35 p r o p o r t i o n o f .556 Non-body C o n t a c t IB YB YY F e m a l e s 346 671 26 30 475 992 0 5 .421 .403 1.0 .857 T a b l e 21. Use o f Non-body and Body C o n t a c t b e h a v i o u r by a l l c l a s s e s i n i n t r a -c l a s s b e h a v i o u r . The number o f Non-body C o n t a c t and Body C o n t a c t b e h a v i o u r p a t t e r n s used a r e compared. X*= 70.5, p<.05, df= 4 m a l e s YY f e m a l e s Non-body C o n t a c t 1707 185 201 Body C o n t a c t 1935 96 72 p r o p o r t i o n o f .468 .658 .736 Non-body C o n t a c t T a b l e 22. Use o f Non-body and Body C o n t a c t b e h a v i o u r by males t o o t h e r c l a s s e s . The number o f Non-body C o n t a c t and Body C o n t a c t b e h a v i o u r p a t t e r n s used a r e compared. X2= 103.2, p<.05, df= 2 44 Y e a r l i n g s ose a l l Non-body C o n t a c t b e h a v i o u r i n i n t r a -c l a s s b e h a v i o u r ( T a b l e 2 4 ) . They do n o t d i s c r i m i n a t e between t h e s e x e s i n t h e use o f Non-body and Body C o n t a c t b e h a v i o u r . I n e a c h o f t h e Non-body C o n t a c t and Body C o n t a c t b e h a v i o u r s , one b e h a v i o u r p a t t e r n i s v e r y common. T h i s i s t h e Mouth f o r Non-body C o n t a c t b e h a v i o u r and t h e M o u t h i n g f o r Body C o n t a c t b e h a v i o u r . The use o f t h e Mouth by males i s compared t o t h a t o f f e m a l e s and Y e a r l i n g s ( T a b l e 2 5 ) . F e m a l e s and Y e a r l i n g s u s e t h e Mouth more t h a n t h e o t h e r Non-body C o n t a c t b e h a v i o u r p a t t e r n s . Males use l e s s . The male c l a s s e s use t h i s b e h a v i o u r p a t t e r n i n d i f f e r e n t f r e q u e n c i e s ( T a b l e 2 6 ) . The use o f t h e Mouth d e c r e a s e s as t h e males mature. The M o u t h i n g i s t h e o n l y Body C o n t a c t b e h a v i o u r p a t t e r n u s e d by f e m a l e s ( T a b l e 2 7 ) . T h i s c o n t r a s t s w i t h male b e h a v i o u r where t h e M o u t h i n g f o r m s o n l y a s m a l l p r o p o r t i o n o f t h e Body C o n t a c t b e h a v i o u r . T h e r e i s no d i f f e r e n c e i n t h e use o f t h e M o u t h i n g by t h e male c l a s s e s ( T a b l e 2 8 ) . P o p u l a t i o n S t r u c t u r e The number o f s e a - l i o n s i n t h e d i f f e r e n t c l a s s e s i s not known, however e s t i m a t e s c a n be made. T h i s i s based cn my f a m i l i a r i t y w i t h t h e c o l o n y ( T a b l e 2 9 ) . The p o p u l a t i o n s t r u c t u r e v a r i e d w i t h i m m i g r a t i o n t o and e m m i g r a t i o n from t h e c o l o n y . At t h e t i m e t h e I n d i v i d u a l D i s t r i b u t i o n Method d a t a a s w e l l a s most o f t h e E n c o u n t e r Method d a t a were c o l l e c t e d , t h e a c t u a l p o p u l a t i o n s t r u c t u r e was p r o b a b l y c l o s e t o t h a t 45 males YY f e m a l e s Non-body C o n t a c t 177 23 54 Body C o n t a c t 37 9 10 p r o p o r t i o n o f .823 .718 .843 Non-body C o n t a c t T a b l e 23. Use o f Non-body and Body C o n t a c t b e h a v i o u r by f e m a l e s t o o t h e r c l a s s e s . The number o f Non-body C o n t a c t and Body C o n t a c t b e h a v i o u r p a t t e r n s used a r e compared. X2= 2.5, p>.05, df= 2 males YY f e m a l e s Non-body C o n t a c t 154 26 19 Body C o n t a c t 84 0 9 p r o p o r t i o n o f .647 1.0 .678 Non-body C o n t a c t T a b l e 24. Use o f Non-body and Body C o n t a c t b e h a v i o u r by Y e a r l i n g s t o o t h e r c l a s s e s . The number o f Non-body C o n t a c t and Body C o n t a c t b e h a v i o u r p a t t e r n s used a r e compared. X*= 13.4, p<.05, df= 2 46 f e m a l e s and males Y e a r l i n g s Mouth 378 39 o t h e r Non-body 683 17 C o n t a c t p r o p o r t i o n .356 .696 o f Mouth T a b l e 25. Use o f mouth by m a l e s , f e m a l e s and Y e a r l i n g s t o a l l c l a s s e s . The number o f Mouth and o t h e r Non-body C o n t a c t b e h a v i o u r p a t t e r n s a r e compared. xz= 26.3, p<.05, df= 1 OB IB YB Mouth 5 89 284 o t h e r Non-body 39 257 387 C o n t a c t p r o p o r t i o n .113 .257 .423 o f Mouth T a b l e 26. Use o f Mouth by male c l a s s e s t o a l l c l a s s e s . The number o f Mouth and o t h e r Non-body C o n t a c t b e h a v i o u r p a t t e r n s a r e compared. X2= 39.2, p<.05, df= 2 47 males f e m a l e s M o u t h i n g 280 5 o t h e r Body 1222 0 C o n t a c t p r o p o r t i o n .186 1.0 o f M o u t h i n g T a b l e 27. Ose o f M o u t h i n g by males and f e m a l e s t o a l l c l a s s e s . The number o f M o u t h i n g and o t h e r Body C o n t a c t b e h a v i o u r p a t t e r n s a r e compared. p<.05 OB IB YB M o u t h i n g 6 104 170 o t h e r Body 26 341 822 C o n t a c t p r o p o r t i o n . 171 .218 . 171 o f M o u t h i n g T a b l e 28. Use o f M o u t h i n g by males t o a l l c l a s s e s . The number o f M o u t h i n g and o t h e r Body C o n t a c t b e h a v i o u r p a t t e r n s a r e compared. X 2= 4.8, p>.05, df= 2 C l a s s P r o p o r t i o n o f c o l o n y OB .05 IB \ .30 YB .40 YY . 15 SC .05 LC . 05 T a b l e 29. ftge-sex s t r u c t u r e o f a n o n - p u p p i n g c o l o n y . 49 e s t i m a t e d . C l a s s A s s o c i a t i o n F o r e a c h c l a s s t h e p r o p o r t i o n o f n e i g h b o u r i n g s e a - l i o n s t h a t b e l o n g t o t h e d i f f e r e n t c l a s s e s i s c a l c u l a t e d ( F i g . 4 ) . I t would be e x p e c t e d t h a t i f t h e r e were no a s s o r t i t i v e a s s o c i a t i o n , t h e n t h e p r o p o r t i o n s o f c l a s s e s f o u n d as n e i g h b o u r s would be s i m i l a r t o t h o s e c h a r a c t e r i s t i c o f t h e whole c o l o n y ( T a b l e 2 9 ) . T h a t i s , most n e i g h b o u r s would be s u b - a d u l t m a l e s and Y e a r l i n g s , w i t h o n l y a few a d u l t males and f e m a l e s . T e r r i t o r i a l O l d B u l l s a s s o c i a t e m a i n l y w i t h f e m a l e s and Y e a r l i n g s . I n t e r m e d i a t e and Young B u l l s a s s o c i a t e w i t h t h e m s e l v e s and Y e a r l i n g s a s e x p e c t e d . L a r g e Cows t e n d t o a s s o c i a t e w i t h O l d B u l l s , Y e a r l i n g s and S m a l l Cows. They a s s o c i a t e w i t h o t h e r L a r g e Cows as e x p e c t e d , b u t do n o t a s s o c i a t e w i t h I n t e r m e d i a t e and Young B u l l s . S m a l l Cows a s s o c i a t e more w i t h L a r g e Cows and S m a l l Cows and l e s s w i t h s u b - a d u l t m a l e s t h a n e x p e c t e d . They a s s o c i a t e w i t h O l d E u l l s and Y e a r l i n g s a t t h e l e v e l s e x p e c t e d . Y e a r l i n g s a s s o c i a t e a s e x p e c t e d w i t h t h e o t h e r c l a s s e s e x c e p t Y e a r l i n g s and I n t e r m e d i a t e B u l l s . They a s s o c i a t e more w i t h t h e m s e l v e s and l e s s w i t h I n t e r m e d i a t e B u l l s t h a n e x p e c t e d . 50 OB IB YB c u « YY SC LC 100 0 n = 100 100 0 n= 100 100 0 n= 11 37 14 13 52 67 23 IS 3 6 4 7 1 3 4 2 2 0 OB IB YB YY SC LC F i g . H. A s s o c i a t i o n . F o r e a c h c l a s s on the o r d i n a t e , i t s n e i g h b o u r s t h a t b e l o n g t o d i f f e r e n t c l a s s e s i s g i v e n i n p e r c e n t . Sample s i z e s o f l e s s t h a n f i v e were n o t i n c l u d e d i n t h e c a l c u l a t i o n o f X 2 . X 2= 371.7, p<.05, df= 15 . The e x p e c t e d v a l u e s a r e c a l c u l a t e d u s i n g T a b l e 29 and a r e s i m i l a r t o t h e Young B u l l and IB f r e g u e n c y d i s t r i b u t i o n s . 51 A c t i v i t y The p r o p o r t i o n o f a c t i v e t i m e f o r e a c h c l a s s i s c a l c u l a t e d ( F i g . 5}. H a l e s and Y e a r l i n g s a r e mere a c t i v e t h a n f e m a l e s . As s e a - l i o n s mature t h e y become l e s s a c t i v e . T h a t i s , o l d e r males a r e l e s s a c t i v e t h a n y o u n g e r males, and L a r g e Cows a r e l e s s a c t i v e t h a n S m a l l Cows. T r a n s i e n c y Y e a r l i n g s a r e v e r y t r a n s i e n t o r i n o t h e r words n o t s i t e s p e c i f i c ( F i g . 6 ) . They f r e q u e n t l y move from one a r e a on the c o l o n y t o a n o t h e r . M a l e s and f e m a l e s a r e l e s s t r a n s i e n t t h a n Y e a r l i n g s . M a l e s become more s i t e s p e c i f i c a s t h e y mature. F e m a l e s r e m a i n i n one a r e a i n t h e n o n - p u p p i n g c o l o n y . U s i n g Y e a r l i n g s a s a c o m p a r i s o n , f e m a l e s become more s i t e s p e c i f i c a s t h e y mature. I n t e r a c t i o n B a t e The i n t e r a c t i o n r a t e i s g i v e n as t h e mean number o f i n t e r a c t i o n s p e r minute ( T a b l e 3 0 ) . An e x p e c t e d number o f i n t e r a c t i o n s i s c a l c u l a t e d u s i n g an o v e r a l l i n t e r a c t i o n r a t e d e r i v e d from t h e t o t a l number o f i n t e r a c t i o n s and m i n u t e s o b s e r v e d . T h i s i s compared t o t h e o b s e r v e d r a t e . I n t e r m e d i a t e and Young B u l l s i n t e r a c t w i t h o t h e r s e a - l i o n s t w i c e as o f t e n as O l d B u l l s and L a r g e Cows, and one and one h a l f t i m e s a s o f t e n a s Y e a r l i n g s and S m a l l Cows. T h e r e i s a g r a d u a l d e c l i n e a s f e m a l e s mature. I n males i t i n c r e a s e s f r o m a low r a t e i n Y e a r l i n g s t o a h i g h r a t e i n Young and 50 18 49 28 18 3 19 OB IB YB YY SC LC Fig. 6. T r a n s i e n c y . The p r o p o r t i o n of m o v e m e n t s to the number of s i g h t i n g s . X2=33.0, P<.05. df = 4. Jfrlhe sample s i z e is too small so the data are not used. Cn CO 54 # o f m i n u t e s i n t e r a c t i o n c l a s s i n t e r a c t i o n s o b s e r v e d r a t e OB 149 1188 .125 IB 283 1051 .269 YB 165 654 .252 YY 118 665 .177 SC 60 353 .169 LC 129 1003 .128 T a b l e 30. I n t e r a c t i o n r a t e g i v e n as i n t e r a c t i o n s per m i n u t e . X2= 101.5, p<.05, df= 5 ' i 55 I n t e r m e d i a t e B u l l s . I t t h e n d e c r e a s e s i n O l d B u l l s . The p r o p o r t i o n o f t i m e s p e n t i n t e r a c t i n g i n c r e a s e s s h a r p l y between t h e Y e a r l i n g and Young B u l l s t a g e s . I t t h e n d e c r e a s e s as males mature. T h e r e i s nc d i s t i n c t t r e n d i n t h e o n t o g e n y o f f e m a l e b e h a v i o u r . Y e a r l i n g s , S m a l l Cows and L a r g e Cows spend a b o u t e q u a l t i m e i n t e r a c t i n g ( T a b l e 3 1 ) . The p r o p o r t i o n o f a c t i v e t i m e t h a t a s e a - l i o n s p e n d s i n t e r a c t i n g w i t h o t h e r s e a - l i o n s i s c a l c u l a t e d ( T a b l e 3 2 ) . Y e a r l i n g s i n t e r a c t f o r o n l y a s m a l l p o r t i o n o f t h e r e a c t i v e t i m e . Young B u l l s spend a l l t h e i r a c t i v e t i m e i n i n t e r a c t i o n s . As males m a t u r e , l e s s o f t h e i r a c t i v e t i m e i s used i n t h i s manner. As f e m a l e s mature, t h e p r o p r o t i o n o f a c t i v e t i m e s p e n t i n t e r a c t i n g i n c r e a s e s . L a r g e Cows i n t e r a c t a l l t h e t i m e t h a t t h e y a r e a c t i v e . A c t i v i t y i s d e f i n e d i n such a way t h a t , i n most c a s e s , i f two a n i m a l s a r e i n t e r a c t i n g , t h e n t h e y a r e a l s o a c t i v e . The v a l u e s c a l c u l a t e d , f o r Young B u l l s and L a r g e Cows show t h a t t h e r e i s a minimum o f 6% e r r o r i n v o l v e d ( T a b l e 3 2 ) . T h i s e r r o r i s m i n o r . Z e r o s were used i n p l a c e o f t h e n e g a t i v e v a l u e s i n T a b l e 32 when X 2 i s c a l c u l a t e d . S i n c e X 2 i s v e r y l a r g e , i t i s u n l i k e l y t h a t t h e s e a d j u s t m e n t s a f f e c t t h e s i g n i f i c a n c e o f t h e d a t a . I n t e r a c t i o n F r e q u e n c y I f s e a - l i o n s i n t e r a c t r a n d o m l y w i t h s u r r o u n d i n g a n i m a l s t h e f r e q u e n c y o f i n t e r a c t i o n w i t h a p a r t i c u l a r c l a s s w i l l be t h e same as t h e f r e q u e n c y t h a t t h e c l a s s e s a r e a s s o c i a t e d 56 OB IB YB YY SC LC m i n u t e s 48.9 208.3 252.2 27.6 9.6 60.4 i n t e r a c t i n g m i n u t e s n o t 1139.1 842.7 401.8 637.4 343.4 942.6 i n t e r a c t i n g p r o p o r t i o n o f t i m e s p e n t .041 .198 .386 .042 .027 .060 i n t e r a c t i n g T a b l e 31. Time e a c h c l a s s s p e n d s i n t e r a c t i n g w i t h o t h e r s e a - l i o n s . X 2= 651.6, p<.05, df= 5 i 57 OB IB m i n u t e s 48.9 208.3 i n t e r a c t i n g m i n u t e s a c t i v e b u t not 137.6 138.5 i n t e r a c t i n g p r o p o r t i o n o f a c t i v e t i m e .261 .600 s p e n t i n t e r a c t i n g YB YY SC LC 252.2 27.8 9.6 60.4 -16.*T 153.3 32.4 -2.2 1.072 .154 .227 1.034 T a b l e 32. A c t i v e t i m e e a c h c l a s s s p e n d s i n t e r a c t i n g w i t h o t h e r s e a - l i o n s . X 2= 457.2, p<.05, df= 5 Th e s e e r r o r s a r e p r o b a b l y c a u s e d by an o v e r e s t i m a t e o f a c t i v e t i m e . '0* was used i n the c a l c u l a t i o n o f X 2 . 58 ( F i g . 4 ) . The o b s e r v e d i n t e r a c t i o n f r e q u e n c i e s o f t h e c l a s s e s a r e compared t o t h e e x p e c t e d f r e q u e n c i e s (based on c l a s s a s s o c i a t i o n ) ( T a b l e 3 3 ) . S e a - l i o n s t e n d t o i n t e r a c t more w i t h t h e male c l a s s e s and l e s s w i t h t h e f e m a l e and Y e a r l i n g c l a s s e s . The i n t e r a c t i o n f r e q u e n c y o f L a r g e Cows i s c o n s i d e r e d i n more d e t a i l ( T a b l e 3 4 ) . The e x p e c t e d i n t e r a c t i o n f r e q u e n c i e s o f t h e male c l a s s e s i s c a l c u l a t e d o n l y on t h e b a s i s c f male a s s o c i a t i o n . O l d and Young B u l l s i n t e r a c t w i t h L a r g e Cows l e s s t h a n e x p e c t e d , w h i l e I n t e r m e d i a t e B u l l s i n t e r a c t more. DISCUSSION H i e r a r c h y A p e c k - d o m i n a n c e h i e r a r c h y ( A l l e e , 1951) e x i s t s between t h e c l a s s e s i n n o n - p u p p i n g c o l o n i e s o f E u m e t o p i a s j u b a t a and C a l i f o r n i a s e a - l i o n ( Z a l o p h u s c a l i f o r n i a n u s ) ( P e t e r s o n and Bartholomew, 1967). The l a r g e r and o l d e r c l a s s e s a r e do m i n a n t t o t h e s m a l l e r and you n g e r o n e s . I n d o m e s t i c c h i c k e n s , d o m i n a n t - s u b o r d i n a n t r e l a t i o n s h i p s r e d u c e t h e amount o f f i g h t i n g t h a t would o t h e r w i s e o c c u r ( G u h l , 1953). T h i s i s a l s o s u g g e s t e d f o r New Z e a l a n d f u r s e a l ( A r c t g c e p h a l u s f p r s t e r i ) ( S t i r l i n g , 1970). The e s t a b l i s h m e n t o f a c l a s s h i e r a r c h y i n 12. j u b a t a may f u n c t i o n i n c a u s i n g l e s s f i g h t i n g and t h e r e b y r e d u c e s o c i a l l y i n d u c e d pup m o r t a l i t y (Boyd and C a m p b e l l , 1971; Bartholomew, 1953; S a n d e g r e n , 1970). ' 59 O b s e r v e d E x p e c t e d M a l e s 597 485 YY 145 194 F e m a l e s 47 108 T a b l e 33. F r e g u e n c y o f male, f e m a l e , and Y e a r l i n g i n t e r a c t i o n s e x p e c t e d i s c a l c u l a t e d on t h e b a s i s o f random i n t e r a c t i o n w i t h n e i g h b o u r s . X*= 72.7, p<.05, df= 2 O b s e r v e d E x p e c t e d OB 37 51 IB 22 4 YB ,4 8 T a b l e 34. I n t e r a c t i o n f r e g u e n c y o f L a r g e Cons w i t h m a l e s . E x p e c t e d i s c a l c u l a t e d on t h e b a s i s o f random i n t e r a c t i o n s w i t h male n e i g h b o u r s . X«= 86.8, p<.05, df= 2 60 H i e r a r c h y i s r e l a t e d t o many o f t h e b e h a v i o u r s measured and i s d i s c u s s e d where a p p r o p r i a t e . D u r a t i o n Hany b e h a v i o u r p a t t e r n s used by f e m a l e s and Y e a r l i n g s a r e I n v e s t i g a t i v e . T h i s s u g g e s t s t h a t f e m a l e s a r e o n l y k e e p i n g aware o f t h e i r i m m e d i a t e s o c i a l e n v i r o n m e n t . By t h e Young B u l l s t a g e , males a r e d o i n g t h i s as w e l l as e s t a b l i s h i n g h i e r a r c h i c a l r e l a t i o n s h i p s w i t h t h e i r n e i g h b o u r s and d e v e l o p i n g c o m m u n i c a t i v e and motor s k i l l s (Mason, 1960). H a l e i n t e r a c t i o n s a r e complex and so i n v o l v e more i n f o r m a t i o n e x c h a n g e t h a n t h o s e o f f e m a l e s o r Y e a r l i n g s . The d u r a t i o n o f t h e i n t e r a c t i o n depends on t h e amount and t y p e o f i n f o r m a t i o n b e i n g communicated ( C a l d w e l l , 1972). Y e a r l i n g and f e m a l e i n t e r a c t i o n s a r e u s u a l l y v e r y s h o r t . T h i s r e s u l t s f r o m t h e s m a l l amount o f i n f o r m a t i o n i n v o l v e d . Males i n t e r a c t l o n g e r b e c a u s e more i n f o r m a t i o n i s b e i n g e x c h a n g e d . The d u r a t i o n o f i n t e r a c t i o n s i n c r e a s e s a s t h e d i f f e r e n c e between t h e a n i m a l s * s t a t u s d e c r e a s e s . S i m i l a r b e h a v i o u r i s r e p o r t e d i n s t o n e s h e e p ( O v i s d a l l i s t o n e i ) ( G e i s t , 1968, 1971) and m a n t i s s h r i m p ( G o n o d a c t y l u s z a c a e ) ( D i n g l e and C a l d w e l l , 1969). The c l o s e r t h e a n i m a l s a r e i n t h e h i e r a r c h y , t h e more i n f o r m a t i o n i s e x c h a n g e d i n o r d e r f o r one t o a s s e r t i t ' s dominance o r b o t h t o a c c e p t e a c h o t h e r as p e e r s . 61 The a s s o c i a t i o n o f d u r a t i o n t o i n f o r m a t i o n c o n t e n t o f i n t e r a c t i o n s i s f u r t h e r d e m o n s t r a t e d by t h e b e h a v i o u r o f m a l e s t o w a r d s f e m a l e s and Y e a r l i n g s . O l d and I n t e r m e d i a t e B u l l s i n t e r a c t f o r s h o r t p e r i o d s w i t h Y e a r l i n g s and S m a l l Cows and l o n g p e r i o d s w i t h L a r g e Cows. T h i s i s s i m i l a r t o s t o n e s h e e p ( G e i s t , 1968, 1971). The t y p e o f i n f o r m a t i o n communicated w i t h L a r g e Cows i s s e x u a l l y o r i e n t e d whereas t h a t w i t h Y e a r l i n g s and S m a l l Cows i s n o t . I n i t i a t i o n A s J i j u b a t a and s t o n e s h e e p ( G e i s t , 1968) mature, t h e y i n i t i a t e more i n t e r a c t i o n s w i t h an i n c r e a s i n g v a r i e t y o f c l a s s e s . O l d e r a n i m a l s a r e more i n c l i n e d t oward v o l u n t a r y s o c i a l b e h a v i o u r . I n i t i a t i o n i s r e l a t e d t o h i e r a r c h y . The yo u n g e r s u b o r d i n a n t c l a s s e s do n o t s t a r t i n t e r a c t i o n s w i t h o l d e r dominant c l a s s e s . I f t h e i n t e r a c t i o n i s a h i e r a r c h i c a l c o n t e s t , t h e s u b o r d i n a n t would l o s e o r be i n j u r e d . The o l d e r c l a s s e s a r e n o t p r e v e n t e d from s t a r t i n g i n t e r a c t i o n s w i t h y o u n g e r c l a s s e s b e c a u s e t h e y c a n a s s e r t t h e i r dominance. D i f f e r e n c e s i n s e x u a l m a t u r i t y o f t h e male c l a s s e s a r e e v i d e n c e d by i n i t i a t i o n o f i n t e r a c t i o n s w i t h L a r g e Cows. O l d B u l l s , I n t e r m e d i a t e B u l l s and L a r g e Cows a r e s e x u a l l y mature. The o l d e r male c l a s s e s may i n i t i a t e i n t e r a c t i o n s w i t h L a r g e Cows f o r s e x u a l r e a s o n s . Young B u l l s do n o t i n i t i a t e i n t e r a c t i o n s w i t h L a r g e Cows. T h i s r e f l e c t s t h e Young B u l l s ' s e x u a l i m m a t u r i t y . 62 V a r i e t y o f B e h a v i o u r P a t t e r n s The v a r i e t y o f b e h a v i o u r p a t t e r n s i n t h e r e p e r t o i r e o f ma l e s i s g r e a t e r t h a n t h a t o f Y e a r l i n g s and f e m a l e s . T h i s a r i s e s f r o m d i f f e r e n t i n t r a - c l a s s r e l a t i o n s h i p s . Male i n t e r a c t i o n s i n v o l v e h i e r a r c h i c a l c o n t e s t s and mock f i g h t i n g ( F a r e n t i n o s , 1971). The t y p e o f i n f o r m a t i o n t r a n s f e r r e d between t h e p a r t i c i p a n t s i s more c o m p l i c a t e d t h a n t h e s i m p l e i n v e s t i g a t i v e and minor t h r e a t b e h a v i o u r o f f e m a l e s and Y e a r l i n g s . The v a r i e t y o f male s i g n a l s a s s i s t s i n c o m m u n i c a t i n g t h e i r more complex messages. B e g i n n i n g and T e r m i n a t i o n B e h a v i o u r The Advance b e h a v i o u r p a t t e r n i s a d e f i n i t e a p p r o a c h o f one s e a - l i o n t o a n o t h e r . W h i l e b e i n g s i m i l a r t o i n i t i a t i o n o f an i n t e r a c t i o n i t i s more i n t e n s e . The s e n d e r i s making an e f f o r t t o i n t e r a c t w i t h a p a r t i c u l a r a n i m a l . M a l e s use t h i s a c t i o n more f r e q u e n t l y t h a n do Y e a r l i n g s and f e m a l e s . T h i s i n d i c a t e s t h a t males a r e more w i l l i n g t o be s o c i a l l y i n v o l v e d . The f r e q u e n c y o f t h e Advance i n i n t e r a c t i o n s w i t h L a r g e Cows i n c r e a s e s w i t h t h e male's age and r e f l e c t s t h e male's s e x u a l m a t u r a t i o n . When males i n t e r a c t w i t h Y e a r l i n g s and S m a l l Cows, t h e r e a r e no s i g n i f i c a n t d i f f e r e n c e s i n t h e i r use o f t h e Advance. The males may r e c o g n i z e t h e s e c l a s s e s a s s e x u a l l y immature (page 66) and so do not a p p r o a c h them. 63 M a l e s a r e s e l e c t i v e o f o t h e r males i n t h e i r use o f t h e A dvance. T h i s s e l e c t i o n i s b a s e d on h i e r a r c h y . The Advance i s used most f r e q u e n t l y w i t h t h e c l a s s t h a t i s i m m e d i a t e l y s u b o r d i n a t e and n e x t most o f t e n w i t h p e e r s . S i n c e most male i n t e r a c t i o n s a r e q u i t e a g g r e s s i v e , s m a l l e r males a v o i d i n t e r a c t i o n s w i t h l a r g e r o n e s . I n t e r a c t i o n s w i t h v e r y s u b o r d i n a n t a n i m a l s do not y i e l d new e x p e r i e n c e s . I f t h e i n t e n s e b e h a v i o u r o f t h e male c l a s s e s f u n c t i o n s i n d e v e l o p i n g c o m m u n i c a t i v e and motor s k i l l s as w e l l a s l e a r n i n g s o c i a l b e h a v i o u r (Mason, 1960; S c h a l l e r , 1968; F a r e n t i n o s , 1971; B e k o f f , 1972) t h e n t h e v a l u e o f i n t e r a c t i o n w i t h e x t r e m e s u b o r d i n a n t s i s m i n i m a l . S e l e c t i o n would f a v o u r males t h a t i n t e r a c t w i t h i m m e d i a t e l y s u b o r d i n a n t a n i m a l s and p e e r s . T h e r e i s l i t t l e c h a n c e o f b e i n g i n j u r e d by s u b o r d i n a n t s and an e g u a l c h a n c e by p e e r s . I m m e d i a t e l y s u b o r d i n a n t a n i m a l s a r e b e h a v i o u r a l l y s i m i l a r t o t h e d o m i n a n t . L i t t l e new b e h a v i o u r i s e n c o u n t e r e d i n t h e s e i n t e r a c t i o n s . However t h e b e h a v i o u r a l r e a d y a c q u i r e d c a n be p r a c t i s e d . I n t e r a c t i o n s w i t h p e e r s p r o v i d e o p p o r t u n i t y t o be e x p o s e d t o new b e h a v i o u r a s w e l l as t o use t h a t a l r e a d y a c q u i r e d . A n i m a l s t h a t c o n t i n u a l l y a p p r o a c h d o m i n a n t s would be s e l e c t e d a g a i n s t . A l t h o u g h more e x p e r i e n c e would be g a i n e d , t h e s u b o r d i n a n t would be d e f e a t e d and r i s k i n j u r y . The A t t a c k b e h a v i o u r p a t t e r n d e m o n s t r a t e s one o f t h e major d i f f e r e n c e s between male and f e m a l e b e h a v i o u r . Males become o b v i o u s l y more a g g r e s s i v e between t h e Y e a r l i n g and Young B u l l s t a g e s , as i n d i c a t e d by t h e f i r s t use o f t h e 64 A t t a c k . The use o f t h i s p a t t e r n ana a l s o a g g r e s s i v e n e s s i n c r e a s e as t h e males mature. A s i m i l a r d e v e l o p m e n t a l s e q u e n c e i s e v i d e n t i n f e r a l g o a t s (Cajjra h i r c u s ) (Shank, 1972). The use o f t h e A t t a c k i n f e m a l e s does n o t change as t h e y mature. The a d u l t f e m a l e s a t a n o n - p u p p i n g c o l o n y a r e a b o u t as a g g r e s s i v e as Y e a r l i n g s . M a l e s a r e s e l e c t i v e o f c l a s s e s t o which t h e y d i r e c t the A t t a c k . I n E i j u b a t a t h e A t t a c k i s n o t used a g a i n s t f e m a l e s . T h i s i s a l s o t r u e o f f e r a l g o a t s (Shank, 1972), Uganda kob ( A d e n o t a kob) (Buechner and S c h l o e t h , 1956) and mountain s h e e p ( G e i s t , 1971). O t h e r w i s e f e m a l e s would withdraw and p o t e n t i a l mates would be l o s t . T h i s i s p a r t i c u l a r l y i m p o r t a n t i n a p i n n i p e d c o l o n y where u n a t t e n d e d f e m a l e s a r e a l m o s t i m m e d i a t e l y a p p r o a c h e d by n e i g h b o u r i n g males ( P a u l i a n , 1964; Bartholomew, 1952; C a r r i c k e t a l , 1962b; S a n d e g r e n , 1970). The A t t a c k i s o n l y o c c a s i o n a l l y used on Y e a r l i n g s . I n a p u p p i n g c o l o n y , between 7% ( S a n d e g r e n , 1970) and 255? (Matheson, Badde and Lopp, 1962) o f t h e f e m a l e s a r e s u c k l i n g Y e a r l i n g s . I f males were a g g r e s s i v e t o w a r d s Y e a r l i n g s i t would f o r c e , them away and t h e i r mothers might f o l l o w . T h i s would r e d u c e t h e r e p r o d u c t i v e s u c c e s s o f t h e m a l e s . The use o f t h e A t t a c k on o t h e r m a l e s , e s p e c i a l l y s u b o r d i n a n t s , i n d i c a t e s c h a n g e s i n t h e a g g r e s s i v e n e s s o f m a l e s a s t h e y mature. A g g r e s s i o n c a u s e s s p a c i n g o u t o f male n o r t h e r n e l e p h a n t s e a l s ( M i r o u n g a g n g u s t i r o s t r i g ) ( L e B o e u f , 1972) and n o n - b r e e d i n g a d u l t male E^ j u b a t a (Orr and P o u l t e r , 65 1967). I n E._ j u b a t a , t h e i n c r e a s e i n a g g r e s s i v e n e s s w i t h m a t u r i t y may accomodate t h e d e v e l o p m e n t o f t e r r i t o r i a l b e h a v i o u r . The l a c k o f i n t e n s e a g g r e s s i o n i n f e m a l e s and Y e a r l i n g s c o u l d be as a d a p t i v e as t h e a g o n i s t i c b e h a v i o u r o f m a les. I f f e m a l e s were a g g r e s s i v e t h e y c o u l d not f o r m c l o s e g r o u p s a r o u n d t e r r i t o r i a l m a l e s . The male would have fewer mates, t h e r e b y r e d u c i n g h i s r e p r o d u c t i v e s u c c e s s , and a l s o t h a t o f t h e f e m a l e s a s a r e s u l t o f t h e p e r i p h e r a l male e f f e c t ( M c L aren, 1967). H i g h l y a g g r e s s i v e f e m a l e s and Y e a r l i n g s would a l s o be s e l e c t e d a g a i n s t b e c a u s e t h e i r b e h a v i o u r would t e n d t o d i s r u p t t h e p u p p i n g c o l o n y , i n c r e a s i n g t h e c h a n c e s o f l o s t , t r a m p l e d o r n e g l e c t e d p u p s . The R e t r e a t b e h a v i o u r p a t t e r n i s u s e d by a s e a - l i o n a t t e m p t i n g t o t e r m i n a t e an i n t e r a c t i o n . M a l e s use the R e t r e a t i n a g r e a t e r p r o p o r t i o n o f i n t r a - c l a s s i n t e r a c t i o n s t h a n do f e m a l e s o r Y e a r l i n g s . T h i s r e s u l t s from the a g g r e s s i o n c h a r a c t e r i s t i c o f male i n t e r a c t i o n s . C l a s s h i e r a r c h y i s d e f i n e d u s i n g t h e R e t r e a t . S e a - l i o n s use t h i s b e h a v i o u r p a t t e r n most o f t e n when t h e y i n t e r a c t w i t h a d o m i n a n t . I f an i n t e r a c t i o n i s a g o n i s t i c , t h e n s u b o r d i n a n t a n i m a l s a r e s u r e l y g o i n g t o be d e f e a t e d and so t r y t o e s c a p e . The C h a s e b e h a v i o u r p a t t e r n c a n be i n t e r p r e t e d t o mean t h a t a s e a - l i o n i s t r y i n g t o p r o l o n g o r r e i n f o r c e t h e r e s u l t o f an i n t e r a c t i o n . T h i s p a t t e r n i s used by males more t h a n by f e m a l e s and Y e a r l i n g s . I t has a l r e a d y been s u g g e s t e d b e c a u s e o f I n i t i a t i o n b e h a v i o u r and t h e a d v a n c e , t h a t males 66 a r e more v o l u n t a r i l y s o c i a l l y i n c l i n e d t h a n t h e e t h e r c l a s s e s . ~ The use o f t h e Chase complements t h i s c o n c l u s i o n . The s e x u a l r e l a t i o n s h i p s between t h e c l a s s e s i s r e f l e c t e d by use o f t h e C h a s e . O l d and I n t e r m e d i a t e B u l l s use t h e p a t t e r n more f r e q u e n t l y t h a n Young B u l l s when i n t e r a c t i n g w i t h L a r g e Cows. A l t h o u g h t h e s a m p l e s a r e s m a l l , t h e r e a r e no s i g n i f i c a n t d i f f e r e n c e s i n t h e use o f t h i s a c t i o n by a l l males c l a s s e s t o w a r d s S m a l l Cows. As w i t h t h e Advance, t h e o l d e r males a r e p r o b a b l y p r o l o n g i n g t h e i r i n t e r a c t i o n s w i t h L a r g e Cows f o r s e x u a l r e a s o n s . I n v e s t i g a t i v e B e h a v i o u r M a t u r i n g males use a d e c r e a s i n g amount o f I n v e s t i g a t i v e b e h a v i o u r i n i n t r a - c l a s s i n t e r a c t i o n s . F e m a l e s and Y e a r l i n g s use i t more t h a n do m a l e s . T h i s does n o t change as t h e f e m a l e s mature. Male d i m o r p h i s m becomes more o b v i o u s w i t h age. I n f e m a l e s s i z e i s t h e o n l y c h a r a c t e r i s t i c t h a t r e a d i l y d i s t i n g u i s h e s young from o l d a n i m a l s . I n v e s t i g a t i v e b e h a v i o u r may be u s e d t o i d e n t i f y t h e m a t u r i t y and s e x o f s e a - l i o n s when p h y s i c a l f e a t u r e s a r e n o t v i s i b l e . T h i s e x p l a i n s t h e d e c r e a s i n g use by m a t u r i n g males and t h e g r e a t e r b u t c o n s t a n t use by m a t u r i n g f e m a l e s . I t i s a l s o p o s s i b l e t h a t t h e l e s s a g g r e s s i v e n a t u r e o f f e m a l e s and Y e a r l i n g s p e r m i t s c l o s e r c o n t a c t between i n t e r a c t i n g a n i m a l s and s u b s e q u e n t l y t h e use o f I n v e s t i g a t i v e b e h a v i o u r . M a l e s , b e i n g i n c r e a s i n g l y a g g r e s s i v e w i t h age, a r e s e p a r a t e d when i n t e r a c t i n g i n o r d e r t o d e f e n d t h e m s e l v e s . T h i s s e p a r a t i o n 67 d o e s n o t p e r m i t t h e use o f I n v e s t i g a t i v e b e h a v i o u r . Body C o n t a c t and Son-Body C o n t a c t B e h a v i o u r Body C o n t a c t b e h a v i o u r p a t t e r n s a r e p h y s i c a l l y e x p r e s s e d a g g r e s s i o n . Non-body C o n t a c t b e h a v i o u r p a t t e r n s a r e a l s o a g g r e s s i v e b u t l i m i t e d t o t h r e a t s and d i s p l a y s . Many o f t h e s e p a t t e r n s a r e f o u n d i n t h e b e h a v i o u r r e p e r t o i r e s o f o t h e r p o l y g y n o u s p i n n i p e d s p e c i e s ( S t i r l i n g , 1971; P e t e r s o n and Bartholomew, 1967; C a r r i c k e t a l , 1962b; Laws, 1956; P a u l i a n , 1964; P e t e r s o n , 1965; Bartholomew, 1952). The v a r i e t y o f b e h a v i o u r p a t t e r n s used by t h e v a r i o u s c l a s s e s was d i s c u s s e d p r e v i o u s l y . D i f f e r e n c e s between male and f e m a l e c l a s s e s r e s u l t f r o m two major s o u r c e s : 1. the d e v e l o p m e n t o f s e v e r a l Body C o n t a c t b e h a v i o u r p a t t e r n s between t h e Y e a r l i n g and Young B u l l s t a g e s , and 2. t h e use o f an i n c r e a s i n g v a r i e t y o f Non-body C o n t a c t b e h a v i o u r p a t t e r n s as males mature. V a r i e t y o f b e h a v i o u r p a t t e r n s i s r e l a t e d t o t h e t y p e o f s o c i a l i n t e r a c t i o n s (page 6 2 ) . F e m a l e s and Y e a r l i n g s a r e n o t as a g g r e s s i v e as males and t h e r e f o r e l a c k many o f t h e s e p a t t e r n s i n t h e i r r e p e r t o i r e . The h i g h l y a g g r e s s i v e males use t h e s e p a t t e r n s i n t h e i r complex h i e r a r c h i c a l and mock f i g h t i n g i n t e r a c t i o n s . The l i f e o f a d u l t males i s s o c i a l l y complex. The e s t a b l i s h m e n t and m a i n t e n a n c e o f t e r r i t o r i e s i n v o l v e s i n t e n s e i n t e r a c t i o n s . Most o f t h e m a l e s ' b e h a v i o u r i n a n o n - p u p p i n g c o l o n y i s mock f i g h t i n g . T h i s term i s used by F a r e n t i n o s (1971) who d i f f e r e n t i a t e s i t from t r u e f i g h t i n g b e c a u s e 68 wounds a c e n o t i n f l i c t e d . B e k o f f (1972) s t a t e s t h a t mock o r p l a y f i g h t i n g c a n be d i s t i n g u i s h e d f r o m t r u e f i g h t i n g by t h e low i n t e n s i t y o f i n t e r a c t i o n s . The i n t e r a c t i o n s o f 13. j u b a t a s u b - a d u l t males a t n o n - p u p p i n g c o l o n i e s a r e n o t as v i c i o u s o r i n t e n s e a s t h o s e d e s c r i b e d f o r a d u l t m a l e s a t a p u p p i n g c o l o n i e s by G e n t r y (1970) and S a n d e g r e n (1970). These d i f f e r e n c e s between a d u l t and s u b - a d u l t male i n t e r a c t i o n s a r e a l s o r e c o g n i z e d by G e n t r y (1970) and O r r and P o u l t e r (1967). Mock f i g h t i n g a s s i s t s i n t h e d e v e l o p m e n t o f c o m m u n i c a t i v e and motor s k i l l s (Mason, 1960; S c h a l l e r , 1968; Shank, 1972). A n i m a l s , s o c i a l l y r e s t r i c t e d i n e a r l y l i f e , show marked i m p a i r m e n t o f s o c i a l a b i l i t i e s ( G i l b e r t , 1968; Mason, 1960; Fox, 1971). The i m p o r t a n c e o f s o c i a l i n t e r a c t i o n s and p l a y i s r e v i e w e d by B e k o f f ( 1 9 7 2 ) . S e a - l i o n s g a i n i n g e a r l y s o c i a l e x p e r i e n c e would be b e t t e r a d a p t e d t o t h e complex s o c i a l s y s t e m o f t h e i r a d u l t l i f e . M a l e s use more Body C o n t a c t b e h a v i o u r t h a n do f e m a l e s and Y e a r l i n g s . However, a s m a l e s mature, t h e amount o f Body C o n t a c t t o Non-body C o n t a c t b e h a v i o u r d e c r e a s e s . M a l e s a r e n o t b ecoming l e s s a g g r e s s i v e , b u t r a t h e r , t h e i r a g g r e s s i o n i s b e i n g e x p r e s s e d i n a d i f f e r e n t manner. O l d e r males a r e l a r g e r and more e x p e r i e n c e d t h a n younger o n e s and s o t h e i r p o t e n t i a l f o r i n f l i c t i n g i n j u r i e s i s g r e a t e r . The younger m a l e s a r e a b l e t o f i g h t v e r y c l o s e t o g e t h e r b e c a u s e p h y s i c a l and b e h a v i o u r a l r e s t r i c t i o n s p r e v e n t s e r i o u s i n j u r i e s . The o l d e r males must r e m a i n s e p a r a t e d b e c a u s e t h e i r i n t e n s e a g g r e s s i o n would r e s u l t i n i n j u r y . From remote p o s i t i o n s 69 t h e y i n t e r a c t and communicate u s i n g Non-body C o n t a c t b e h a v i o u r . I f t h i s f a i l s t o p e r m i t one male from a s s e r t i n g i t s e l f o r e a c h male a c c e p t i n g t h e o t h e r as p e e r , t h e n Body C o n t a c t b e h a v i o u r may be u s e d . T h i s i s o f t e n t h e c a s e d u r i n g t h e e a r l y p a r t o f t h e b r e e d i n g s e a s o n when males a r e u n a c c u s t o m e d t o t h e i r n e i g h b o u r s and s o c i a l r e l a t i o n s h i p s have not been e s t a b l i s h e d ( G e n t r y , 1970; Rand, 1967; P e t e r s o n and Bartholomew, 1967; S t i r l i n g , 1 9 7 1). i s m ales o f b o t h E. -jubata and s t o n e s h e e p ( G e i s t , 1971) m a ture, t h e y i n c r e a s e t h e amount o f Non-body C o n t a c t o r d i s p l a y b e h a v i o u r . E.. j u b a t a a d u l t males r e m a i n f o r l o n g p e r i o d s i n a p u p p i n g c o l o n y a f t e r e s t a b l i s h i n g a t e r r i t o r y ( G e n t r y , 1 9 7 0 ) . They do n o t f e e d d u r i n g t h i s t i m e but r e l y i n f a t r e s e r v e s ( G e n t r y , 1970). Non-body C o n t a c t b e h a v i o u r i s l e s s s t r e n u o u s t h a n Body C o n t a c t b e h a v i o u r and i t s use c o n s e r v e s e n e r g y ( S t i r l i n g , 1970; G e i s t , 1966; G e n t r y , 1970). The e n e r g y s a v e d c o u l d p r o l o n g t h e males* s t a y a t a p u p p i n g c o l o n y . L o n g e r t e n e n c y i n c r e a s e s t h e m a l e s ' r e p r o d u c t i v e s u c c e s s ( G e n t r y , 1970) and so t h e use o f Non-body C o n t a c t b e h a v i o u r i n p l a c e o f Body C o n t a c t b e h a v i o u r i s s e l e c t i v e l y a d v a n t a g e o u s . M a l e s u s e more s Non-body C o n t a c t b e h a v i o u r w i t h f e m a l e s , l e s s w i t h Y e a r l i n g s and l e a s t w i t h m a l e s , The Non-body C o n t a c t b e h a v i o u r p a t t e r n s a r e n o t p h y s i c a l l y a g g r e s s i v e and so may be i n t e r p r e t e d d i f f e r e n t l y by males and f e m a l e s . M a l e s may use Non-body C o n t a c t b e h a v i o u r more w i t h f e m a l e s i n o r d e r t o a t t r a c t them and promote f e m a l e g r e g a r i o u s n e s s . 70 T h i s would t e n d t o i n c r e a s e t h e males* r e p r o d u c t i v e s u c c e s s . Body C o n t a c t b e h a v i o u r would d i s r u p t t h e f e m a l e s and c a u s e them t o d i s p e r s e . F e m a l e s t r e a t a l l c l a s s e s s i m i l a r l y when r e l a t i v e amounts o f Body and Non-body C o n t a c t b e h a v i o u r s a r e c o n s i d e r e d . T h i s i n d i c a t e s t h e s i m p l i c i t y o f f e m a l e b e h a v i o u r and t h e i r r e l a t i o n s h i p s w i t h t h e o t h e r c l a s s e s . Y e a r l i n g s use some Body C o n t a c t b e h a v i o u r w i t h males and f e m a l e s , b u t none w i t h o t h e r Y e a r l i n g s . T h i s s u g g e s t s t h a t Y e a r l i n g s a r e m i m i c i n g t h e b e h a v i o u r s o f t h e a n i m a l s t h e y i n t e r a c t w i t h . I t adds f u r t h e r t o t h e argument o f e a r l y s o c i a l i n v o l v e m e n t b e i n g i m p o r t a n t i n g a i n i n g e x p e r i e n c e . The M o u t h i n g i s a s i m p l e p a t t e r n compared t o L u n g i n g o r N e c k i n g . I t i s i n f r e q u e n t l y used by f e m a l e s and Y e a r l i n g s and i s t h e o n l y Body C o n t a c t b e h a v i o u r p a t t e r n i n t h e i r r e p e r t o i r e . T h i s r e f l e c t s t h e s i m p l e and n o n - a g g r e s s i v e b e h a v i o u r o f t h e s e c l a s s e s . The use o f M o u t h i n g r e l a t i v e t o o t h e r Body C o n t a c t b e h a v i o u r r e m a i n s c o n s t a n t a s males mature. T h i s i n d i c a t e s t h a t most Body C o n t a c t b e h a v i o u r i s a c q u i r e d b e f o r e t h e Young B u l l s t a g e and does n o t d e v e l o p f u r t h e r . The Mouth i s a s i m p l e t h r e a t d i s p l a y ( Sandegren, 1970). As males mature, i t s use d e c r e a s e s r e l a t i v e t o o t h e r Non-body C o n t a c t b e h a v i o u r . Males a r e t h e r e f o r e n o t o n l y i n c r e a s i n g t h e use and v a r i e t y o f Non-body C o n t a c t b e h a v i o u r but a r e a l s o i n c r e a s i n g t h e i r use o f more complex b e h a v i o u r p a t t e r n s . F e m a l e s use t h e Mouth more t h a n males do. As w i t h t h e 71 M o u t h i n g , t h i s d e m o n s t r a t e s t h e s i m p l i s t i c n a t u r e c f f e m a l e b e h a v i o u r . A s s o c i a t i o n and I n t e r a c t i o n F r e q u e n c y S e a - l i o n s a r e n o t r a n d o m l y d i s t r i b u t e d i n a n c n - p u p p i n g c o l o n y . F e m a l e s , a t t h e S m a l l Cow s t a g e , b e g i n t o g r o u p a r o u n d O l d B u l l s and o t h e r f e m a l e s . L a r g e Cows a r e even more s e l e c t i v e . Immature m a l e s show no d i s t i n c t c l u m p i n g . When males r e a c h t h e O l d B u l l s t a g e and become t e r r i t o r i a l , t h e y b e g i n t o f o r m g r o u p s w i t h f e m a l e s . The same d i s t r i b u t i o n p a t t e r n i s e v i d e n t a t p u p p i n g c o l o n i e s ( G e n t r y , 1970; O r r and P o u l t e r , 1967) b u t b e c a u s e t h e r e l a t i v e p r o p o r t i o n s o f t h e c l a s s e s a r e d i f f e r e n t : i t i s n o t o b v i o u s . The clumped d i s t r i b u t i o n o f t h e s e a - l i o n s r e s t r i c t s t h e i r a v a i l a b l e s o c i a l p a r t n e r s . S e l e c t i v e i n t e r a c t i o n r e s u l t s i n f u r t h e r r e s t r i c t i o n s . In E_. j u b a t a , i n t e r a c t i o n s i n v o l v e males more t h a n f e m a l e s . T h i s i s a l s o t r u e o f b i g h o r n s h e e p ( G e i s t , 1 971). M a l e s a r e more s o c i a l l y i n c l i n e d t h a n f e m a l e s . T h i s i s e v i d e n t i n many a s p e c t s o f male b e h a v i o u r . S e l e c t i v e i n t e r a c t i o n s a r e an i m p o r t a n t f a c t o r i n t h e d i s t r i b u t i o n o f c l a s s e s . I n t e r m e d i a t e B u l l s i n t e r a c t w i t h f e m a l e s more t h a n i f t h e y c h o s e t h e i r s o c i a l p a r t n e r s r a n d o m l y from s u r r o u n d i n g a n i m a l s . A l t h o u g h f e m a l e s s e l d o m i n i t i a t e i n t e r a c t i o n s w i t h m a l e s , t h e y a r e h a r a s s e d by I n t e r m e d i a t e B u l l s and a t t e m p t t o a v o i d i n t e r a c t i o n s w i t h them. H a r a s s m e n t o f f e m a l e s by s u b - a d u l t males i s r e p o r t e d i n t h e f o l l o w i n g s p e c i e s : E A j u b a t a ( G e n t r y , 1970), A l a s k a 72 f u r s e a l ( C a l l o r h i n u s u r s i n u s ) (Bartholomew, 1953), s o u t h e r n e l e p h a n t s e a l ( M i r o u n g a l e o n i n a ) ( C a r r i c k e t a l , 1 9 6 2 a , b ) , n o r t h e r n e l e p h a n t s e a l ( L e B o e u f , 1972) and p r o n g h o r n ( i J S i i l p j s I p r a a m e r i c a n a ) (Bromley and K i t c h e n , 1973). O n l y B r o m l e y and K i t c h e n (1973) q u a n t i f y t h i s h a r a s s m e n t . They show t h a t f e m a l e s o u t s i d e t e r r i t o r i e s a r e h a r a s s e d by t h e o v e r z e a l o u s c o u r t s h i p o f n o n - t e r r i t o r i a l m ales and c o n c l u d e t h a t t e r r i t o r i e s a c t as s a n c t u a r i e s . I n E. j u b a t a , s u b - a d u l t males a r e e x c l u d e d from t e r r i t o r i e s . T h i s r e s t r i c t s t h e i r a c c e s s t o f e m a l e s . S i n c e f e m a l e s n e a r O l d B u l l s a r e h a r a s s e d l e s s t h a n i n a g g r e g a t i o n s w i t h s u b - a d u l t m a l e s , t h e y t e n d t o l e a v e t h e a r e a s o f h a r a s s m e n t and c o l l e c t i n t h e r e f u g e s c r e a t e d by t e r r i t o r i e s . T e r r i t o r i a l B e h a v i o u r and T r a n s i e n c y Some a d u l t males a t n o n - p u p p i n g c o l o n i e s a r e t e r r i t o r i a l w h i l e o t h e r s a r e n o t . T h i s i s r e l a t e d t o t h e p r e s e n c e o f f e m a l e s . F e m a l e s may be a r e l e a s i n g mechanism e l i c i t i n g t e r r i t o r i a l i t y s i n c e t h e y i n f l u e n c e t e r r i t o r i a l b e h a v i o u r i n many p i n n i p e d s ( G e n t r y , 1970; Kenycn, 1960; P e t e r s o n and Bartholomew, 1967; Rand, 1967; P a u l i a n , 1967; Bartholomew, 1952; Laws, 1954; Cameron, 1971; Hewer and B a c k h o u s e , 1960). E. j u b a t a s u b - a d u l t males a r e n o t t e r r i t o r i a l a t p u p p i n g c o l o n i e s ( G e n t r y , 1970) o r n o n - p u p p i n g c o l o n i e s . S i n c e Young B u l l s and male Y e a r l i n g s a r e s e x u a l l y immature, t h e r e would be no r e p r o d u c t i v e a d v a n t a g e i f t h e y e s t a b l i s h e d and m a i n t a i n e d t e r r i t o r i e s . I n t e r m e d i a t e B u l l s a r e s e x u a l l y 73 mature. However, s i n c e t h e r e a r e few f e m a l e s a t n o n - p u p p i n g c o l o n i e s i t may be s e l e c t i v e l y a d v a n t a g e o u s f o r them t o r e f r a i n f r o m t e r r i t o r i a l b e h a v i o u r . M a l e s i n t h e I n t e r m e d i a t e B u l l s t a g e a r e i n an a c c e l e r a t e d g r o w t h phase ( P i k e , 1966). T h o s e t h a t m a i n t a i n a t e r r i t o r y have t o s t a y on t h e c o l o n y . T h i s r e s t r i c t s t h e i r f e e d i n g and c o n s e q u e n t l y g r o w t h . N o n - t e r r i t o r i a l I n t e r m e d i a t e B u l l s , f r e e t c f e e d any t i m e , would be l a r g e r and s o o n e r a b l e t o e s t a b l i s h a t e r r i t o r y a t a p u p p i n g c o l o n y . I n t e r m e d i a t e B u l l s t h a t a r e t e r r i t o r i a l would be s o c i a l l y d e p r i v e d s i n c e t h e y g a i n e x p e r i e n c e f r o m i n t e r a c t i o n s w i t h a s m a l l number o f f a m i l i a r n e i g h b o u r s . N o n - t e r r i t o r i a l I n t e r m e d i a t e B u l l s a r e f r e e t o move a r o u n d and i n t e r a c t w i t h u n f a m i l i a r n e i g h b o u r s o f t h e c o l o n y , t h e r e b y g a i n i n g new s o c i a l e x p e r i e n c e s . A n i m a l s w i t h more s o c i a l e x p e r i e n c e would b e t t e r be a b l e t o communicate and a d a p t t o t h e complex s y s t e m a t p u p p i n g c o l o n i e s . J i JJjbtSta become more s i t e s p e c i f i c as t h e y mature. T h i s may accommodate t h e d e v e l o p m e n t o f male t e r r i t o r i a l b e h a v i o u r . G e n t r y (1970) d i s c u s s e s t h e i m p o r t a n c e o f h a b i t u a t i o n i n t h e s t a b i l i t y o f t e r r i t o r i a l s y s t e m s . M a l e s become s o c i a l l y f a m i l i a r w i t h t h e i r n e i g h b o u r s by s t a y i n g i n one p l a c e . T h i s a l l o w s m utual b o u n d a r i e s and h i e r a r c h i c a l r e l a t i o n s h i p s t o be e s t a b l i s h e d . F e m a l e s a r e more s i t e s p e c i f i c t h a n m a l e s . T h i s r e f l e c t s t h e i r a d j u s t m e n t t o t h e l i m i t e d t e r r i t o r i a l s y s t e m i n n o n - p u p p i n g c o l o n i e s . H arassment by s u b - a d u l t males 74 f o r c e s f e m a l e s t o o c c u p y and r e m a i n i n wh a t e v e r t e r r i t o r i e s a r e a v a i l a b l e . In p u p p i n g c o l o n i e s where t h e r e a r e many a d j a c e n t t e r r i t o r i e s , f e m a l e s a r e n o t s t a t i o n a r y ( S a n d e g r e n , 1970; P e t e r s o n and Bartholomew, 1967). A c t i v i t y , I n t e r a c t i o n R a t e s and I n t e r a c t i o n Time The h i g h l e v e l s o f a c t i v i t y and i n t e r a c t i o n r a t e s o f s u b - a d u l t males a f f e c t c l a s s d i s t r i b u t i o n . T e r r i t o r i e s a r e l e s s d i s r u p t e d t h a n o t h e r a r e a s i n a n o n - p u p p i n g c o l o n y b e c a u s e o f t h e e x c l u s i o n o f s u b - a d u l t m a l e s . The f e m a l e s e n t e r and r e m a i n i n t h e t e r r i t o r i e s i n o r d e r t o a v o i d a r e a s o f a c t i v i t y (Bartholomew, 1953; C a r r i c k e t a l , 1962b). T h i s c o n t r i b u t e s t o t h e clumped d i s t r i b u t i o n o f O l d B u l l s and f e m a l e s . The h i g h i n t e r a c t i o n r a t e s and l o n g e r t i m e s s p e n t i n t e r a c t i n g o f s u b - a d u l t males r e s u l t s i n t h e i r r e c e i v i n g more s o c i a l c o n t a c t w i t h a b r o a d e r r a n g e o f c o l o n y members t h a n f e m a l e s and Y e a r l i n g s . The a d a p t i v e n e s s o f s o c i a l e x p e r i e n c e t o t h e s u b - a d u l t males has a l r e a d y been d i s c u s s e d (page 6 3 ) . T r a n s i e n c y may be a f a c t o r c o n t r i b u t i n g t o t h e t r e n d s o b s e r v e d i n a c t i v i t y and i n t e r a c t i o n r a t e . The o l d e r a n i m a l s move a r o u n d l e s s and so become h a b i t u a t e d t o t h e i r n e i g h b o u r s . G e n t r y ' s (1970) d a t a s u g g e s t t h a t s o c i a l l y f a m i l i a r a n i m a l s i n t e r a c t l e s s f r e q u e n t l y . Younger s e a - l i o n s move a r o u n d t h e c o l o n y and a r e a l w a y s b e i n g c o n f r o n t e d w i t h u n f a m i l i a r a n i m a l s whose sex and s t a t u s a r e unknown. 75 E s t a b l i s h m e n t o f s o c i a l r e l a t i o n s h i p s r e s u l t s i n h i g h e r i n t e r a c t i o n r a t e s and a c t i v i t y . Y e a r l i n g s s p e n d l i t t l e a c t i v e t i m e i n t e r a c t i n g and do n o t d i s p l a y f o r l o n g p e r i o d s a s do O l d B u l l s . However t h e y do spend t i m e u s i n g t h e L o o k i n g b e h a v i o u r p a t t e r n . T h i s e x p l a i n s t h e i r l a r g e amount o f a c t i v e t i m e w i t h no d i r e c t s o c i a l i n v o l v e m e n t . as m ales g e t o l d e r , t h e a c t i v e t i m e s p e n t i n t e r a c t i n g d e c r e a s e s and more t i m e i s s p e n t d i s p l a y i n g w i t h o u t s p e c i f i c s o c i a l p a r t n e r s . M a t u r i n g f e m a l e s spend an i n c r e a s i n g p r o p o r t i o n o f t h e i r a c t i v e t i m e i n t e r a c t i n g . S i n c e t h e s e i n t e r a c t i o n s a r e i n i t i a t e d by t h e o l d e r male c l a s s e s , i n a c t i v e f e m a l e s a r e d i s t u r b e d and f o r c e d t o i n t e r a c t w i t h t h e m a l e s . T h i s complements t h e o b s e r v a t i o n o f m a les h a r a s s i n g f e m a l e s . In p u p p i n g c o l o n i e s o f t h e New Z e a l a n d f u r s e a l , a d u l t males a r e a c t i v e f o r 21.9$ and i n t e r a c t i n g f o r 3.1% o f t h e t i m e ( S t i r l i n g , 1971). The v a l u e s f o r t e r r i t o r i a l a d u l t male Ii. j u b a t a i n n o n - p u p p i n g c o l o n i e s a r e 15.7$ and 4.19? r e s p e c t i v e l y . The c l o s e a greement o f t h e s e d a t a s u g g e s t t h a t t h e two s p e c i e s a r e o p e r a t i n g under s i m i l a r b e h a v i o u r a l s y s t e m s . S t i r l i n g (1971) a t t r i b u t e s t h e r e l a t i v e l y low a c t i v i t y o f t h e a d u l t males t o t h e a d a p t i v e n e s s o f c o n s e r v i n g e n e r g y . T h i s p r o l o n g s t h e m a l e s ' p r e s e n c e a t t h e c o l o n y and i n c r e a s e s t h e i r r e p r o d u c t i v e p o t e n t i a l . An e q u a l l y p l a u s i b l e e x p l a n a t i o n i s t h e s t a b i l i z i n g e f f e c t r e d u c e d a c t i v i t y would have a t a p u p p i n g c o l o n y . I n t e r a c t i n g males d i s r u p t f e m a l e s 76 and c a u s e them t o move away (Bartholomew, 1953; C a r r i c k e t a l , 1962b) and t h e r e b y t h e y l o s e p o t e n t i a l mates. S e l e c t i o n would f a v o u r a d u l t males who c o u l d m a i n t a i n t h e i r t e r r i t o r y w i t h a minimum amount o f d i s r u p t i o n . T h i s i s a c c o m p l i s h e d by males becoming l e s s a c t i v e and i n t e r a c t i v e and by u s i n g t h e more p a s s i v e Non-body C o n t a c t b e h a v i o u r . SUMMARY Non- p u p p i n g c o l o n i e s p f E± j u b a t a a r e s o c i a l l y o r g a n i z e d i n t o a p e c k - d o m i n a n c e h i e r a r c h y , w i t h t h e l a r g e r and o l d e r c l a s s e s d ominant t o t h e s m a l l e r and y o u n g e r o n e s . The s t a t u s o f t h e o l d e r male c l a s s e s a r e d i s t i n c t whereas t h o s e o f f e m a l e s and y o u n g e r males a r e n o t . C l a s s e s behave d i f f e r e n t l y t o w a r d s o t h e r s . T h e s e d i f f e r e n c e s r e s u l t f r o m s e x u a l , h i e r a r c h i c a l and t e r r i t o r i a l r e l a t i o n s h i p s . I n t e r a c t i o n s between s e x u a l l y mature males and f e m a l e s a r e n o n - a g o n i s t i c . M a l e s i n c r e a s e t h e i r use o f Non-body C o n t a c t b e h a v i o u r when t h e y i n t e r a c t w i t h f e m a l e s . T h i s and t h e a l m o s t e x c l u s i v e use o f Non-body C o n t a c t b e h a v i o u r by f e m a l e s r e s u l t i n more t r a n g u i l i n t e r a c t i o n s t h a n i n t e r a c t i o n s i n v o l v i n g o n l y males where th e v i o l e n t Body C o n t a c t b e h a v i o u r p r e d o m i n a t e s . I n t e r a c t i o n s between males a r e a g o n i s t i c . As males mature t h e y become more a g g r e s s i v e and use an i n c r e a s i n g amount o f Non-body C o n t a c t b e h a v i o u r i n t h e i r mock f i g h t s . The Non-body C o n t a c t b e h a v i o u r r e p l a c e s Body C o n t a c t b e h a v i o u r and a l l o w s a g o n i s t i c c o m m u n i c a t i o n f r o m a d i s t a n c e . T h i s would be a d a p t i v e i n t h e t e r r i t o r i a l 77 s i t u a t i o n a t a t p u p p i n g c o l o n y where males a r e s e p a r a t e d . M a l e s i n t e r a c t l o n g e r and more f r e q u e n t l y t h a n f e m a l e s and so a r e more s o c i a l l y i n v o l v e d . T h i s s o c i a l i n v o l v e m e n t i s v o l u n t a r y f o r males but l e s s s o f o r f e m a l e s . T h i s i s shown by t h e m a l e s ' t e n d e n c y t o i n i t i a t e i n t e r a c t i o n s and use t h e Advance and C h a s e b e h a v i o u r p a t t e r n s . The s o c i a l c o n d i t i o n i n g o f males by mock f i g h t s c o n t r i b u t e s t o t h e d e v e l o p m e n t o f t h e i r c o m m u n i c a t i v e and motor s k i l l s . T h i s r e s u l t s i n a n i m a l s a d e p t i n s o c i a l a b i l i t i e s and so b e t t e r a b l e t o cope w i t h t h e c o m p l e x i t i e s o f a d u l t s o c i e t y . A d u l t male and f e m a l e b e h a v i o u r s a r e l e s s v i g o r o u s v t h a n t h a t o f s u b - a d u l t males. The a d u l t m a l e s and f e m a l e s use l a r g e r amounts o f t h e p a s s i v e Non-body C o n t a c t b e h a v i o u r , a r e l e s s a c t i v e , i n t e r a c t l e s s f r e g u e n t l y and spend l e s s t i m e i n t e r a c t i n g t h a n s u b - a d u l t males. T e r r i t o r i e s where a d u l t m a l e s and f e m a l e s a r e c o n c e n t r a t e d a r e more t r a n q u i l t h a n a r e a s where s u b - a d u l t males a g g r e g a t e . S i n c e f e m a l e s a v o i d a r e a s o f a c t i v i t y and h a r a s s m e n t by s e x u a l l y mature s u b - a d u l t m a l e s , t h e y group i n t e r r i t o r i e s which p r o v i d e r e f u g e s . T h i s c o n t r i b u t e s t o t h e c o l o n y ' s s p a c i a l o r g a n i z a t i o n . S u b - a d u l t males a r e not t e r r i t o r i a l . Some a d u l t males e s t a b l i s h and m a i n t a i n t e r r i t o r i e s b u t o n l y when t h e y a r e n e a r f e m a l e s . T h i s t e r r i t o r i a l b e h a v i o u r i s n o t as i n t e n s e as t h a t o f a d u l t males a t p u p p i n g c o l o n i e s . T e r r i t o r i a l i t y r e s u l t s i n t h e e x p u l s i o n o f s u b - a d u l t males from c e r t a i n a r e a s and t h e s u b s e q u e n t a g g r e g a t i o n o f f e m a l e s . S u b - a d u l t m a l e s mix t o g e t h e r o u t s i d e t e r r i t o r i e s and do n o t s e g r e g a t e 78 i n t o age s p e c i f i c g r o u p s . S u b - a d u l t males do n o t m a i n t a i n t e r r i t o r i e s and s o a r e f r e e t o move a b o u t t h e c o l o n y . S i n c e t h e y a r e v e r y t r a n s i e n t , i n d i v i d u a l h i e r a r c h i c a l and o t h e r s o c i a l r e l a t i o n s h i p s a r e c o n t i n u a l l y b e i n g b r o k e n and new ones e s t a b l i s h e d . T h i s r e s u l t s i n s o c i a l d i s r u p t i o n , whereas s o c i a l . s t a b i l i t y i s a ccomodated by a d u l t m ales and f e m a l e s who a r e more s t a t i o n a r y and so m a i n t a i n p r o l o n g e d s o c i a l r e l a t i o n s h i p s . N o n - p u p p i n g c o l o n i e s a r e s p a t i a l l y o r g a n i z e d s i m i l a r t o p u p p i n g c o l o n i e s e x c e p t f o r t h e r e l a t i v e p r o p o r t i o n s o f age-sex c l a s s e s . N on-pupping c o l o n i e s a r e l e s s s o c i a l l y o r g a n i z e d t h a n p u p p i n g c o l o n i e s . S o c i a l o r g a n i z a t i o n i n J i j u b a t a c o l o n i e s i s promoted by t h e t r a n g u i l b e h a v i o u r o f a d u l t s and i n h i b i t e d by t h e d i s r u p t i v e b e h a v i o u r a f s u b - a d u l t m a l e s . 79 ACKNOWLEDGEMENTS I am v e r y g r a t e f u l t o Dr. H. Dean F i s h e r f o r h i s s u p e r v i s i o n and c r i t i c i s m o f t h i s t h e s i s . I t h a n k Dr. J.R. K r e b s , Dr. I . M c T a g g a r t Cowan, B r . H.C. Nordan, P e t e r Maser, T s i t i k a H a r e s t a d , M u r r a y Kennedy, D o l o r e s S t . A r n a u d , A l l a n E d i e and t h e f a m i l y o f M a u r i c e and F r a n c i s C c l l e t t e f o r t h e i r a s s i s t a n c e . The Canada M i n i s t r y o f T r a n s p o r t p r o v i d e d l o g i s t i c s u p p o r t . The r e s e a r c h was f i n a n c e d by t h e N a t i o n a l R e s e a r c h C o u n c i l , F i s h e r i e s R e s e a r c h E o a r d o f Canada and t h e U n i t e d S t a t e s N a t i o n a l G e o g r a p h i c S o c i e t y . 80 LITERATURE CITED A l l e e , S.C. 1951. C o - o p e r a t i o n Among A n i m a l s , With Human I m p l i c a t i o n s . Henry Schuman, New Y o r k . 233 p. Bartholomew, G.A. 1952. R e p r o d u c t i o n and t h e s o c i a l b e h a v i o u r o f t h e n o r t h e r n e l e p h a n t s e a l . U n i v . C a l i f . P u b l . Z o o l . 47: 369-472. Bartholomew, G.A. 1953. B e h a v i o u r a l f a c t o r s a f f e c t i n g s o c i a l s t r u c t u r e i n t h e A l a s k a f u r s e a l . T r a n s . 18th N o r t h Amer. W i l d l . C o n f . , 481-502. B e k o f f , M. 1972. The d e v e l o p m e n t o f s o c i a l i n t e r a c t i o n , p l a y and m e t a c o m m u n i c a t i o n i n mammals: an e t i o l o g i c a l p e r s p e c t i v e . Q u a r t . Rev. o f B i o l . 47: 412-434. B e l k i n , A.N. 1966. Summer d i s t r i b u t i o n , s t o c k s , p r o s p e c t s f o r c o m m e r c i a l u t i l i z a t i o n and c e r t a i n f e a t u r e s o f t h e b i o l o g y o f s e a - l i o n s i n h a b i t i n g t h e K u r i l e I s l a n d s . I z v e s t i y a o f t h e P a c i f i c S c i e n t i f i c R e s e a r c h I n s t i t u t e o f F i s h e r i e s and O c e a n o g r a p h y (T.I.N.R.O.) 58: 69-95. T r a n s l a t e d by F i s h . Res. Bd. Canada, S e r . #720. Boyd, J.M. and R.N. C a m p b e l l . 1971. The g r e y s e a l , H a l i c h o e r u s j r j j a u s * a t N o r t h Rona, 1959-1968. J . Z o o l . 164:~469-512. B r o m l e y , P.T. and D.W. K i t c h e n . 1973. C o u r t s h i p i n p r o n g h o r n . Symposium on t h e B e h a v i o u r o f U n g u l a t e Mammals. In p r e s s . I.U.C.N. 1973. B u e c h n e r , H.K. and R. S c h o l e t h . 1965. C e r e m o n i a l m a t i n g b e h a v i o u r i n Uganda kob d d e n o t a kob thomasi,Neumann). Z. T i e r p s y c h o l . 22: 209-225. C a l d w e l l , R . I . 1972. The making o f a winner - i n d i v i d u a l v a r i a t i o n i n t h e a g g r e s s i v e b e h a v i o u r o f t h e m a n t i s s h r i m p , G g n g d a c t y l u s z a c a e . Amer. Z o o l . 12: 651. Cameron, A.W. 1971. T e r r i t o r i a l b e h a v i o u r i n t h e w e s t e r n A t l a n t i c g r e y s e a l ( H a l i c h o e r u s g r y p u s ) . J . Z o o l . 164: 443-449. C a r r i c k , R., S.E. C s o r d a s , S.E. Ingham and K. K e i t h . 1962a. S t u d i e s on t h e s o u t h e r n e l e p h a n t s e a l , Mitcjjncja l e o n i n a ( L . ) . I I I . The a n n u a l c y c l e i n r e l a t i o n t o age and s e x . C.S.I.R.O. W i l d l . Res. 7: 119-160. 81 C a r r i c k , R., S.E. C s o r d a s , S.E. Ingham and K. K e i t h . 1962b. S t u d i e s on t h e s o u t h e r n e l e p h a n t s e a l , M i r c u i u j a l e o n i n a ( L . ) . I V . B r e e d i n g and d e v e l o p m e n t . C.S.I.B.O. W i l d l . Res. 7: 161-197. D i n g l e , H. and R.I. C a l d w e l l . 1969. t h e a g g r e s s i v e and t e r r i t o r i a l b e h a v i o u r o f t h e m a n t i s s h r i m p . B e h a v i o u r 33: 115-136. E d i e , A.G. 1973. P e r s o n a l c o m m u n i c a t i o n . F a r e n t i n o s , B.C. 1971. Some o b s e r v a t i o n s on t h e p l a y b e h a v i o u r o f t h e S t e l l e r s e a - l i o n ( J u m e t o j j i a s j u b a t a ) . Z. T i e r p s y c h o l . 28: 428-438. Fox, M.W. 1971. I n t e g r a t i v e Development o f t h e B r a i n and B e h a v i o u r i n t h e Dog. U n i v . C h i c a g o P r e s s , C h i c a g o . 348 G e i s t , V. 1968. The e v o l u t i o n a r y s i g n i f i c a n c e o f mountain s h e e p h o r n s . E v o l . 20: 558-566. G e i s t , V. x 1968. On t h e i n t e r r e l a t i o n o f e x t e r n a l a p p e a r a n c e , s o c i a l b e h a v i o u r and s o c i a l s t r u c t u r e o f m o u n t a i n s h e e p . Z. T i e r p s y c h o l . 25: 199-215. G e i s t , V. 1971. M o u n t a i n Sheep, a S t u d y o f B e h a v i o u r and E v o l u t i o n . U n i v . o f C h i c a g o P r e s s , C h i c a g o and London. 383 p. G e n t r y , R.L. 1970. S o c i a l b e h a v i o u r o f t h e S t e l l e r s e a -l i o n . Ph.d. T h e s i s , U n i v . o f C a l i f . , S a n t a C r u z . 113 p. G i l b e r t , B.K. 1968. D evelopment o f s o c i a l b e h a v i o u r i n t h e f a l l o w d e e r (Dama dama). Z. T i e r p s y c h o l . 25: 867-876. G u h l , A.M. 1953. S o c i a l b e h a v i o u r o f t h e d o m e s t i c f o w l . A g r i c . Exp. S t n . T e c h . B u l l . 73. K a n s a s S t a t e C o l l e g e , M a n h a t t a n , K a n s a s . i n E t k i n , W. 1964. S o c i a l B e h a v i o u r and o r g a n i z a t i o n Among V e r t e b r a t e s . U n i v . C h i c a g o P r e s s , C h i c a g o and L ondon. 307 p. Hewer, H.R. and K.M. B a c k h o u s e . 1960. A p r e l i m i n a r y a c c o u n t o f a c o l o n y o f g r e y s e a l s i n t h e S o u t h e r n I n n e r H e b r i d e s . P r o c . Z o o l . S o c . Lond. 134: 157-195. Kenyon, K.W. 1962. H i s t o r y o f t h e S t e l l e r s e a - l i o n a t t h e P r i b o l o f I s l a n d s , A l a s k a . 0. Mammal. 43: 68-75. 82 L e B o e u f , B . J . 1972. S e x u a l b e h a v i o u r i n t h e n o r t h e r n e l e p h a n t s e a l , M i r p u n g a a n g u s t i r o s t r i g . B e h a v i o u r 41: 1-26. L e B o e u f , B . J . and 8.S. P e t e r s o n . 1968. S o c i a l s t a t u s and ma t i n g a c t i v i t y i n e l e p h a n t s e a l s . S c i e n c e 163: 91-93. Laws, R.M. 1956. The e l e p h a n t s e a l ( M i r p u n g a l e o n i n a L i n n . ) . I I . G e n e r a l , s o c i a l and r e p r o d u c t i v e b e h a v i o u r . F a l k l a n d I s . D e p e n d e n c i e s S u r v e y , London, S c i . Rep. 13. 88 p. M c L a r e n , I.A. 1967. S e a l s and g r o u p s e l e c t i o n . E c o l o g y 48: 104-110. Mason, W.A. 1960. The e f f e c t s o f s o c i a l r e s t r i c t i o n on t h e b e h a v i o u r o f r h e s u s monkeys: I . F r e e s o c i a l b e h a v i o u r . J . Comp. P h y s i o l . P s y c h o l . 53: 582-589. H a t h i s e n , O.A., R.T. Baade and R.J. Lopp. 1962. B r e e d i n g h a b i t s , g r o w t h and stomach c o n t e n t s o f t h e S t e l l e r s e a -l i o n i n A l a s k a . J . Mammal. 43: 469-477. O r r , R.T. and T.C. P o u l t e r . 1967. Some o b s e r v a t i o n s on r e p r o d u c t i o n , g r o w t h and s o c i a l b e h a v i o u r i n t h e S t e l l e r s e a - l i o n . P r o c . C a l i f . A c a d . S c i . 35: 193-226. P a u l i a n , P. 1964. C o n t r i b u t i o n a l»etude de l ' o t a r i e de l ' l l e Amsterdam. Mammalia 28: S u p p l . 1: 1-146. P e r l o v , A.S. 1971. The o n s e t o f s e x u a l m a t u r i t y i n s e a -l i o n s . A l l - U n i o n R e s e a r c h I n s t i t u t e o f M a r i n e F i s h e r i e s and O c e a n o g r a p h y (V.N.I.B.O.) , P r o c . 80: 174-189. P e t e r s o n , R.S. 1965. B e h a v i o u r o f t h e n o r t h e r n f u r s e a l . Dr. S c i . T h e s i s . J o h n H o p k i n s U n i v . , B a l t i m o r e , M a r y l a n d . 214 p. P e t e r s o n , R.S. and G.A. Bartholomew. 1967. The n a t u r a l h i s t o r y and b e h a v i o u r o f t h e C a l i f o r n i a s e a - l i o n . Amer. Soc. Mammal. S p e c . P u b l . 1: 1-79. P i k e , G.C. 1966. The n o r t h e r n s e a - l i o n , E u m e t o p i a s j u b a t a , on t h e c o a s t o f B r i t i s h C o l u m b i a . M a n u s c r i p t c f F i s h . Res. Bd. Canada B u l l . 51 p. Rand, R.W. 1967. The cape f u r s e a l ( A r c t p c e p h a l u s j g u s i l l u s ) 3. G e n e r a l b e h a v i o u r on l a n d and s e a . I n v e s t . Rep. D i v . S ea. F i s h . S. A f r . 60: 1-39. 83 S a n d e g r e n , F . E . 1970. B r e e d i n g and m a t e r n a l b e h a v i o u r o f t h e S t e l l e r s e a - l i o n ( E u m e t g g i a s j u b a t a ) i n A l a s k a . M.Sc. T h e s i s , U n i v . o f A l a s k a . 138 p. S c h a l l e r , G. 1967. The Deer and t h e T i g e r ; a S t u d y o f W i l d l i f e i n I n d i a . U n i v . C h i c a g o P r e s s , C h i c a g o . 370 p. Shank. C C . 1972. Some a s p e c t s o f s o c i a l b e h a v i o u r i n a p o p u l a t i o n o f f e r a l g o a t s (Cajsra h i r c u s I . ) . Z. T i e r -p s y c h o l . 30: 488-528. S t i r l i n g , I . 1970. O b s e r v a t i o n s o f t h e New Z e a l a n d f u r s e a l ( l££t°cephalus f o r s t e r i ) . J . Nammal. 51: 766-778. S t i r l i n g , I . 1971. S t u d i e s on t h e b e h a v i o u r o f t h e s o u t h A u s t r a l i a n f u r s e a l , A r c t o c e p h a l u s f o r s t e r i ( L e s s o n ) . I . A n n u a l c y c l e , p o s t u r e s and c a l l s , and a d u l t males d u r i n g t h e b r e e d i n g s e a s o n . A u s t . J . Z o o l . 19: 243-266. T h o r s t e i n s e n , F.V. And C . J . L e n s i n k . 1962. B i o l o g i c a l o b s e r v a t i o n s o f t h e S t e l l e r s e a - l i o n s t a k e n d u r i n g an e x p e r i m e n t a l h a r v e s t . J . W i l d l . Hgmt. 26: 353-359. 

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            data-media="{[{embed.selectedMedia}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
https://iiif.library.ubc.ca/presentation/dsp.831.1-0101262/manifest

Comment

Related Items