Open Collections

UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

A comparison of the reproductive and behavioural differences in feral and domestic Japanese quail Nichols, Cathleen Rose 1991

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

Item Metadata

Download

Media
831-UBC_1991_A6_7 N49.pdf [ 7.46MB ]
Metadata
JSON: 831-1.0098657.json
JSON-LD: 831-1.0098657-ld.json
RDF/XML (Pretty): 831-1.0098657-rdf.xml
RDF/JSON: 831-1.0098657-rdf.json
Turtle: 831-1.0098657-turtle.txt
N-Triples: 831-1.0098657-rdf-ntriples.txt
Original Record: 831-1.0098657-source.json
Full Text
831-1.0098657-fulltext.txt
Citation
831-1.0098657.ris

Full Text

COMPARISON OF THE REPRODUCTIVE AND BEHAVIOURAL DIFFERENCES IN FERAL AND DOMESTIC JAPANESE QUAIL By CATHLEEN ROSE NICHOLS B . S c , U n i v e r s i t y o f B r i t i s h Columbia, 1983 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES DEPARTMENT OF POULTRY SCIENCE We accept t h i s t h e s i s as conforming t o the r e q u i r e d standard August, 1991 (c) Cathleen Rose N i c h o l s , 1991 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of ? o O c m g ^ Sofc jQ<L£ The University of British Columbia Vancouver, Canada DE-6 (2/88) A B S T R A C T Aspects of r e p r o d u c t i o n o f f e r a l and domestic Japanese q u a i l ( C o t u r n i x j a p o n i c a ) were compared under a s e m i - n a t u r a l s e t t i n g : 1) the f r e q u e n c i e s of c o u r t s h i p d i s p l a y s and mating behaviour with r e l a t i o n t o age, time o f day, and the female's l a y i n g c y c l e ; 2) p a i r f o r m a t i o n and maintenance; mate guarding, and e x t r a - p a i r a c t i v i t i e s ; 3) f e c u n d i t y , h a t c h a b i l i t y , f e r t i l i t y , c h i c k m o r t a l i t y , and d u r a t i o n of p a r e n t - c h i c k a s s o c i a t i o n . A l l but two of the c o u r t s h i p d i s p l a y s and b e h a v i o u r a l components r e p o r t e d i n the l i t e r a t u r e were e x h i b i t e d by both domestic Japanese q u a i l and f e r a l q u a i l i n t h i s study. Two h i t h e r t o u n r eported d i s p l a y s (Dancing and Leading) were a l s o observed performed by f o r both domestic and f e r a l q u a i l . F e r a l males crowed l e s s than domestic males, but u n l i k e the domestic males, v a r i e d t h e i r crowing frequency d u r i n g the female * s l a y i n g c y c l e . Domestic males performed fewer c o u r t s h i p d i s p l a y s , but c o p u l a t e d more f r e q u e n t l y than f e r a l males. F e r a l males were more a g g r e s s i v e towards both males and females than were domestic males. Domestic females, however, were s i g n i f i c a n t l y more a g g r e s s i v e towards males than were f e r a l females. D i s p l a y f r e q u e n c i e s and mating behaviour were h i g h e r i n the morning than a f t e r n o o n , and h i g h e r i n two-year o l d b i r d s than i n y e a r l i n g s f o r both f e r a l and domestic q u a i l . G e n e r a l l y , f e r a l males were p a i r e d w i t h one female f o r the whole season, whereas domestic males had s h o r t e r p a i r bonds and f r e q u e n t l y switched mates. Domestic males attempted p r o p o r t i o n a l l y more f o r c e d e x t r a - p a i r c o p u l a t i o n s than f e r a l males, and o n l y a s s o c i a t e d c l o s e l y w i t h females when t h e i r eggs c o u l d be f e r t i l i z e d . F e r a l females produced one s u c c e s s f u l c l u t c h per season and r e n e s t e d o n l y i f n e s t i n g was d i s r u p t e d p r i o r t o i n c u b a t i o n , w h i l e domestic females produced up t o t h r e e c l u t c h e s per season. C l u t c h s i z e , h a t c h a b i l i t y , f e r t i l i t y , and c h i c k m o r t a l i t y were s i m i l a r f o r both f e r a l and domestic q u a i l . F a c u l t a t i v e n e s t p a r a s i t i s m was seen f o r both f e r a l and domestic q u a i l . In t h i s study domestic Japanese q u a i l showed a b e h a v i o u r a l r e p e r t o i r e much l i k e f e r a l q u a i l . T h i s suggests d o m e s t i c a t i o n was not " d e g e n e r a t i v e " ; i n s t e a d b e h a v i o u r a l components may not be expressed i n domestic c o n d i t i o n s t h a t l a c k the a p p r o p r i a t e e l i c i t i n g s t i m u l i . i v TABLE OF CONTENTS ABSTRACT i i TABLE OF CONTENTS i v LIST OF TABLES v i i i LIST OF FIGURES i x ACKNOWLEDGEMENTS X CHAPTER 1: GENERAL INTRODUCTION 1 CHAPTER 2: GENERAL METHODS 5 MAIN STUDY 5 EXPERIMENTAL BIRDS 5 1. H i s t o r y o f f e r a l q u a i l p a r e n t a l s t o c k 5 2. H i s t o r y o f domestic q u a i l p a r e n t a l s t o c k . . . . . 7 PEN DESIGN 7 EXPERIMENTAL DESIGN 9 1. Experimental b i r d s - Group 1 10 2. Experimental b i r d s - Group 2 12 BIRD IDENTIFICATION 12 BEHAVIOURAL OBSERVATIONS 12 SUPPLEMENTARY STUDY 13 DATA ANALYSIS 14 CHAPTER 3: COURTSHIP AND BEHAVIOURAL DISPLAYS OF FERAL AND DOMESTIC JAPANESE QUAIL INTRODUCTION 15 METHODS 16 BEHAVIOURAL OBSERVATIONS 16 V 1. C o u r t s h i p and s o c i a l d i s p l a y s 17 2. Mating behaviour 18 3. A g g r e s s i v e and escape behaviour 19 4. Male-female i n t e r a c t i o n s 19 BIRD LOCATIONS 20 WHOLE DAY OBSERVATIONS 20 DATA ANALYSES 20 RESULTS 24 NEW DISPLAYS 24 DISPLAY FREQUENCIES OF FERAL AND DOMESTIC QUAIL 25 1. C o u r t s h i p and s o c i a l d i s p l a y s 25 2. Mating behaviour 25 3. A g g r e s s i v e and escape 29 4. Male-female i n t e r a c t i o n s 29 MALE DISPLAY FREQUENCIES DURING THE LAYING CYCLE...31 DISPLAY FREQUENCIES DURING MORNING AND AFTERNOON OBSERVATION PERIODS 31 DISPLAY FREQUENCIES DURING 1987 AND 1988.. 31 INDIVIDUAL ASSOCIATIONS 35 DISCUSSION 35 COURTSHIP DISPLAYS 35 DEVIATIONS OF DOMESTIC QUAIL FROM FERAL QUAIL 40 CHAPTER 4: PAIR BONDS OF FERAL AND DOMESTIC JAPANESE QUAIL INTRODUCTION 44 METHODS 45 DEFINITION 45 BEHAVIOURAL OBSERVATIONS 46 DATA ANALYSES 47 RESULTS 49 PAIR FORMATION IN JAPANESE QUAIL 49 WITHIN-PAIR AND EXTRA-PAIR ACTIVITIES 54 1. C o u r t s h i p d i s p l a y s 54 2. Mating a c t i v i t i e s 54 3. Mate guarding 59 DISCUSSION 60 CHAPTER 5: REPRODUCTION OF FERAL AND DOMESTIC JAPANESE QUAIL INTRODUCTION 66 METHODS 69 CLUTCH DATA 69 FERTILITY AND HATCHABILITY 70 CHICK MORTALITY AND PARENT-YOUNG ASSOCIATION 70 DATA ANALYSES 71 RESULTS 71 AGE OF SEXUAL MATURITY AND INCIDENCE OF NON-BREEDING 71 CLUTCH SIZE 71 1. C l u t c h e s incubated 71 2. Eggs l a i d 73 TIME AND FREQUENCY OF RENESTING 75 FERTILITY, HATCHABILITY AND INCUBATION OF EGGS 76 NEST PARASITISM 76 CHICK MORTALITY 77 v i i " PARENT-YOUNG ASSOCIATION 78 EGG PRODUCTION 78 DISCUSSION 79 FERAL FEMALES 81 1. Onset of bree d i n g 81 2. Nest p a r a s i t i s m 82 3. In c u b a t i o n and Brooding 83 DOMESTIC FEMALES 85 1. E a r l i e r age of onset of b r e e d i n g 8 6 2. Extent of the b r e e d i n g season and number of eggs produced 87 3. P a r e n t a l behaviour 88 IMPLICATIONS 89 CHAPTER 6: GENERAL DISCUSSION 91 IMPLICATIONS 92 RECOMMENDATIONS 96 FUTURE STUDIES 97 LITERATURE CITED. 98 Appendix 1. Data sheets f o r r e c o r d i n g b e h a v i o r a l d i s p l a y s . 1 0 8 Appendix 2. G r i d maps f o r r e c o r d i n g b i r d l o c a t i o n s 109 Appendix 3. Observations of the dancing d i s p l a y (DN) i n Japanese q u a i l . . . 110 Appendix 4. Male - Female a s s o c i a t i o n s and Female - Male a s s o c i a t i o n s I l l v i i i LIST OF TABLES Tabl e 3.1 Q u a n t i t a t i v e d i f f e r e n c e s i n c o u r t s h i p d i s p l a y s and mating behaviour between f e r a l and domestic Japanese q u a i l .26 T a b l e 3.2 Q u a n t i t a t i v e d i f f e r e n c e s i n c o u r t s h i p d i s p l a y s and mating behaviour between 1987 and 1988 36 T a b l e 4.1 P a i r formation i n Japanese q u a i l kept i n the f l i g h t - p e n s 50 T a b l e 4.2 The d u r a t i o n o f pair-bonds i n days 53 T a b l e 4.3 R e l a t i v e frequency o f c o u r t s h i p d i s p l a y s and mating behaviour performed by p a i r e d male t o h i s mate (I P ) , by the p a i r e d male t o another female not h i s mate (EP), and by a non-paired male t o a female (OP) 55 T a b l e 4.4 The d i s t r i b u t i o n o f a male's mating a c t i v i t i e s ( w i t h i n - p a i r vs e x t r a - p a i r ) a t the t h r e e stages of the l a y i n g c y c l e o f h i s own mate 56 T a b l e 4.5 Success r a t e of w i t h i n - p a i r and e x t r a - p a i r c o p u l a t i o n s by f e r a l and domestic males d u r i n g the t h r e e stages of t h e i r mates' l a y i n g c y c l e 58 T a b l e 5.1 D e s c r i p t i o n of i n c u b a t e d n e s t s found i n the two y e a r s of the study 72 T a b l e 5.2 C l u t c h s i z e of females i n the f l i g h t - p e n 74 i x LIST OF FIGURES F i g . 1.1. Number of w i l d Japanese q u a i l c a p t u r e d by l i c e n s e d hunters 3 F i g . 2.1. Pen Design 8 F i g . 2.2. Experimental d e s i g n 11 F i g . 3.1. D i u r n a l p a t t e r n o f males r e t u r n i n g and s t r u t t i n g t o females d u r i n g f i v e days o f o b s e r v a t i o n s 27 F i g . 3.2. D i f f e r e n c e s i n the frequency of male crowing between f e r a l and domestic males w i t h r e l a t i o n s h i p t o the female's l a y i n g c y c l e 28 F i g . 3.3. D i u r n a l p a t t e r n o f a g g r e s s i v e behaviour between males and females d u r i n g f i v e days o f o b s e r v a t i o n 30 F i g . 3.4. Frequencies of male-female i n t e r a c t i o n s w i t h r e l a t i o n t o the female's l a y i n g c y c l e 32 F i g . 3.5. Frequencies of male a g g r e s s i v e and mating behaviour w i t h r e l a t i o n s h i p t o the female's l a y i n g c y c l e 33 F i g . 3.6. D i f f e r e n c e s i n c o u r t s h i p d i s p l a y s and mating behaviour between morning and a f t e r n o o n o b s e r v a t i o n s .34 F i g . 3.7. FYG2-1988: Male - female a s s o c i a t i o n s 37 F i g . 3.8. DYG2-1988: Male - female a s s o c i a t i o n s 38 F i g . 4.1. The average l e n g t h of time (seconds) f o r a male t o r e t u r n t o h i s mate a f t e r l e a v i n g her s i d e d u r i n g the f o u r stages o f her l a y i n g c y c l e 61 F i g . 5.1. T o t a l egg p r o d u c t i o n 80 X ACKNOWLEDGEMENTS T h i s study would never have m a t e r i a l i z e d without my i n i t i a l i n t r o d u c t i o n t o the world o f Japanese q u a i l p r o v i d e d by the l a t e CW. Roberts. H i s enthusiasm when working w i t h t h i s s p e c i e s c a p t u r e d my i n t e r e s t . T h i s i n t e r e s t c o n t i n u e d t o be n u r t u r e d by K. M. Cheng, my t h e s i s a d v i s o r , who i n t r o d u c e d me t o the f a s c i n a t i n g s c i e n c e of a v i a n r e p r o d u c t i v e b i o l o g y . To both of them I am g r e a t l y indebted. The success o f t h i s p r o j e c t depended upon my r e t u r n i n g from Hawaii w i t h some trapped q u a i l . To t h a t end I r e c e i v e d h e l p from many. I thank Ronald Walker, Ron Bachman, Nelson Santos and Jon G i f f i n of the S t a t e of Hawaii, Department of Land and N a t u r a l Resources, D i v i s i o n o f F o r e s t r y and W i l d l i f e - Ron was a constant source of h e l p and i n f o r m a t i o n , and taught us how t o throw the throw-net, Nelson was the f i r s t t o l e a r n o f t h e q u a i l on the macadamia nut p l a n t a t i o n . Dale Anderson, the manager, granted p e r m i s s i o n t o t r a p on the Puna Macadamia Nut P l a n t a t i o n ; employees Jerome S a s a k i and D a n i e l V i c t u r o p r o v i d e d maps and i n f o r m a t i o n . Kathy and Mannix M i l l e r , Jimmy Wright, Fred Yamashiro and Monte Ric h a r d s allowed us t o t r a p on t h e i r p r o p e r t y , and B i l l y Yasuda and James Watt a s s i s t e d us i n the f i e l d . A s s i s t a n c e was a l s o p r o v i d e d by Masa Takaoka, Reggie Lee and Steve Mountainspring. V e t e r i n a r y a s s i s t a n c e was p r o v i d e d by Drs. B. B e l f r i d g e and Jason Moniz. Important p r e l i m i n a r y i n f o r m a t i o n was p r o v i d e d by V i c x i Lewin, Robert P y l e and Andrew J . Berger. To complete a graduate degree p a r t - t i m e , the support o f one's employer i s a n e c e s s i t y . I wish t o thank Ted C a t h c a r t and the U.B.C. Department o f Animal S c i e n c e f o r t h e i r support and f l e x i b i l i t y , and a l s o f o r the encouragement they g i v e t e c h n i c a l s t a f f t o f u r t h e r t h e i r e d u c a t i o n . To the s t a f f a t the U.B.C. Q u a i l G e n e t i c Stock Centre ( L o i s Enns, A p r i l Peasgood, Pamela D e t t w i l e r , Karen F o r r e s t , Sue Smith, John Baah, Rosemarie F a b i j a n , M a mie Ford, Sandie Morrow, E l l e n Teng, Jim Cox, Sarah C a r l s o n , and Joanne McLarty), I can o n l y say how f o r t u n a t e I have been t o be a s s o c i a t e d w i t h so many hard working people. T h e i r e f f o r t s r e s u l t e d i n the e f f i c i e n t running o f the Q u a i l U n i t , a l l o w i n g me t o take time t o do my t h e s i s . For t h e i r t e c h n i c a l a s s i s t a n c e , I must thank Andre B r e a u l t and Yolanda Leung f o r the a d d i t i o n a l hours o f o b s e r v a t i o n s they p r o v i d e d . To the members of my s u p e r v i s o r y committee, K. Cheng, D. S h a c k l e t o n and J . Smith, a s p e c i a l thanks f o r t h e i r many c o n s t r u c t i v e c r i t i c i s m s and h e l p f u l s u g g e s t i o n s . To complete t h i s t h e s i s i t was necessary t o become a r e c l u s e , so I would l i k e t o thank my parents (Annette and C h a r l e s N i c h o l s ) and p a r e n t s - i n - l a w (Daphne and the l a t e Roland Robinson) f o r t h e i r p a t i e n c e and understanding d u r i n g t h i s time. F i n a l l y , I am g r a t e f u l f o r the support and l o v e shown by my husband C h r i s t o p h e r Robinson. He helped t o c a p t u r e the q u a i l , a p p l i e d h i s computer programming s k i l l s towards h e l p i n g me x i i c o n s o l i d a t e my volumes o f data f o r a n a l y s i s , and a l s o p r o v i d e d many u s e f u l c r i t i c i s m s d u r i n g the w r i t i n g o f t h i s t h e s i s . 1 CHAPTER 1 GENERAL INTRODUCTION Many s p e c i e s o f g a l l i f o r m e s a re c u r r e n t l y f a c i n g d r a s t i c p o p u l a t i o n d e c l i n e s (Cramp and Simmons 1980; J e n k i n s 1990) as a r e s u l t o f h a b i t a t d e s t r u c t i o n and over e x p l o i t a t i o n . There have been many attempts t o r e s t o c k d e p l e t e d areas o r i n t r o d u c e g a l l i f o r m e s i n t o new areas w i t h c a p t i v e r e a r e d b i r d s and/or w i t h domestic c o n s p e c i f i c s (Potts 1986; J e n k i n s 1990). De s p i t e these e f f o r t s , c a p t i v e - b r e d b i r d s have d i f f i c u l t y e s t a b l i s h i n g themselves i n the w i l d ; f o r example, bobwhite q u a i l ( C o l i n u s v i r q i n i a n u s ) (Roseberry e t a l . 1987), gray p a r t r i d g e s (Perdix perdix) (Beani and D e s s i - F u l g h e r i 1985) and ring-necked pheasants (Phasianus c o l c h i c u s ) (Krauss e t a l . 1987) . While f a c t o r s such as i n a p p r o p r i a t e h a b i t a t and poor t i m i n g o f r e l e a s e can be s i g n i f i c a n t , r e s t o c k i n g and i n t r o d u c t i o n f a i l u r e s may be due t o a l a c k o f "wildness" and/or " n a t u r a l " behaviour ( K l i m s t r a 1975; P o t t s 1986). Both g e n e t i c " d e f i c i e n c i e s " (Backs 1982) and environmental e f f e c t s of b e i n g r a i s e d i n c a p t i v i t y have been blamed (Roseberry e t a l . 1987). Lorenz (1965) s t a t e d , " G e n e t i c i s t s have c o r r e c t l y emphasized t h a t w i l d animals cannot, s t r i c t l y speaking be propagated i n c a p t i v i t y , because c a p t i v i t y changes a l l h i t h e r t o e f f e c t i v e s e l e c t i v e f a c t o r s i n so profound a manner t h a t s e r i o u s changes must be expected i n the genome of the 2 s t o c k a f t e r o n l y a few g e n e r a t i o n s " . Lorenz's i m p l i c a t i o n t h a t domestic animals are "degenerate" s t i l l p e r s i s t s (see review i n P r i c e 1984). However, a r e c e n t comparison of p o s t -r e l e a s e behaviour and s u r v i v a l among t h r e e types o f pen-reared and w i l d bobwhite q u a i l (Roseberry e t a l . 1987) i n d i c a t e d t h a t the r e a r i n g environment was more important than g e n e t i c s i n d e t e r m i n i n g t r a i t s such as "wildness" among the c a p t i v e p o p u l a t i o n s . R e s t o c k i n g areas where g a l l i f o r m e s are d e c l i n i n g or b eing e x t i r p a t e d i s a p r a c t i c e t h a t i s c u r r e n t l y b e i n g e v a l u a t e d f o r i t s successes and f a i l u r e s (Krauss e t a l . 1987; Rands and Hayward 1987). A d d i t i o n a l s t u d i e s comparing domesticated g a l l i f o r m e s w i t h t h e i r w i l d c o n s p e c i f i c s can p r o v i d e i n f o r m a t i o n t h a t may h e l p i n c r e a s e f u t u r e r e s t o c k i n g success r a t e s . Japanese q u a i l (Coturnix i a p o n i c a ) are c l o s e l y r e l a t e d t o c h i c k e n s and p a r t r i d g e s , and belong t o the pheasant f a m i l y . They are n a t i v e t o e a s t A s i a and, i n p a r t i c u l a r , t o the i s l a n d s of Japan where p o p u l a t i o n s are d r a s t i c a l l y d e c l i n i n g (Kimura 1991) ( F i g . 1.1). L i t t l e i s known about t h e i r r e p r o d u c t i v e b i o l o g y . In c a p t i v i t y , Japanese q u a i l have been bred f o r meat and eggs throughout the world. They have been domesticated s i n c e a t l e a s t the e l e v e n t h c e n t u r y (Wetherbee 1961). S i n c e domestic Japanese q u a i l may be the o n l y source f o r r e p l e n i s h i n g d e p l e t e d w i l d s t o c k s , i t i s important t h a t we understand the d i f f e r e n c e s i n the r e p r o d u c t i v e behaviour of Figure 1.1: Number of wild Japanese quail captured by l icensed hunters in Japan. 700 , 000 600 , 000 -5 00 , 0 00 4 00 , 0 00 300 , 000 2 00 , 0 00 100 ,000 0 i i i i i i i I I i i i i i i i i i i i i i i i i i i i i i i i | i i i i i i i 1925 1935 1945 1955 1965 1975 1985 (Reproduced from K imura (1991)), 4 w i l d and domes t i c Japanese q u a i l . In the w i l d , Japanese q u a i l h a b i t a t i s i n g r a s s l a n d s , c r o p l a n d s , r i v e r s i d e s , a l p i n e meadows and g r a s s s t eppes (Long 1981) . They a r e d i f f i c u l t t o s t u d y i n t h e w i l d due t o t h e i r s m a l l s i z e and s e c r e t i v e h a b i t s . W i t h t h i s i n mind , I chose t o compare t h e r e p r o d u c t i v e and b e h a v i o u r a l d i f f e r e n c e s o f w i l d and domes t i c Japanese q u a i l i n l a r g e o u t d o o r f l i g h t - p e n s t o de termine i f t h e r e are enough d i f f e r e n c e s t o make the domes t i c s t r a i n s u n s u i t a b l e f o r r e s t o c k i n g p u r p o s e s . In C h a p t e r 3 o f t h i s t h e s i s , I compare the c o u r t s h i p and b e h a v i o u r a l d i s p l a y s o f w i l d and domes t i c q u a i l t o de termine i f the domes t i c q u a i l have r e t a i n e d i m p o r t a n t communicat ion s i g n a l s . In C h a p t e r 4, I examine the d i f f e r e n c e between the w i l d and domest i c q u a i l i n male - f emale i n t e r a c t i o n s both w i t h i n and o u t s i d e the p a i r bond. In C h a p t e r 5, I compare the r e p r o d u c t i o n o f the two t y p e s o f q u a i l . 5 CHAPTER 2 GENERAL METHODS MAIN STUDY EXPERIMENTAL BIRDS 1. H i s t o r y o f f e r a l q u a i l p a r e n t a l s t o c k : Japanese q u a i l were o r i g i n a l l y i n t r o d u c e d t o North America as gamebirds i n 1870, and r e l e a s e s c o n t i n u e d i n t o the l a t e 1950's ( S t a n f o r d 1957). The m a j o r i t y o f b i r d s r e l e a s e d o r i g i n a t e d from domestic Japanese q u a i l imported from Japan by the U.S. F i s h and W i l d l i f e S e r v i c e . A l l q u a i l s t o c k s r e l e a s e d i n North America p e r i s h e d w i t h i n 1 year. Although a l l s t o c k i n g attempts f a i l e d on the North American c o n t i n e n t , r e l e a s e attempts on the Hawaiian i s l a n d s were s u c c e s s f u l and a p o p u l a t i o n o f f e r a l Japanese q u a i l s u r v i v e d and has been r e g u l a r l y hunted. T h i s p o p u l a t i o n o r i g i n a t e d from domestic Japanese q u a i l r e l e a s e d on Maui and Lan a i i n 1921 (Munro 1960). Other domestic Japanese q u a i l were r e l e a s e d on Kauai i n 1944 (Munro 1960) and are p r e s e n t l y w e l l e s t a b l i s h e d on Kauai, Molokai, L a n a i , Maui, and Hawaii (Peterson 1961). The l a r g e s t p o p u l a t i o n o f f e r a l Japanese q u a i l on the Hawaiian I s l a n d s i n 1985, was on the i s l a n d of Hawaii. S i n c e import r e g u l a t i o n s ban the i m p o r t a t i o n o f animals from A s i a t o North America, I trapped my study b i r d s 6 from t h i s p o p u l a t i o n . Although d e s c r i b e d here as f e r a l , P r i c e (1984) suggests t h a t a f t e r s u r v i v i n g many g e n e r a t i o n s i n the w i l d , domestic animals l o s e t h e i r " d o m e s t i c i t y " and can be t r e a t e d as w i l d . In December 1985, f o u r male and e i g h t female f e r a l q u a i l were trapped and t r a n s p o r t e d t o the Q u a i l G e n e t i c Stock Center of the U n i v e r s i t y o f B r i t i s h Columbia. They were housed i n a qua r a n t i n e shed (3 x 4m) f o r 4 months. The f l o o r was covered w i t h shavings, and p i l e s o f a l f a l f a hay were p l a c e d around the pe r i m e t e r t o p r o v i d e h i d i n g p l a c e s f o r the b i r d s . The b i r d s were maintained on 9h of l i g h t and 15h of darkness (9L/15D). The l i g h t i n g was g r a d u a l l y i n c r e a s e d (1 h/week) s t a r t i n g 3 weeks p r i o r t o r e l e a s i n g them i n t o an outdoor f l i g h t pen. During the summer of 1986, a t l e a s t f i v e o f the e i g h t females produced c l u t c h e s from which 34 f i r s t g e n e r a t i o n o f f s p r i n g were hatched a r t i f i c a l l y d u r i n g J u l y and August of 1986. A f t e r b e i n g housed i n brooders f o r the f i r s t 6 weeks, 26 b i r d s (nine males and 17 females) remained a f t e r j u v e n i l e m o r t a l i t y . These b i r d s were moved t o ind o o r f l o o r pens (1.21 by 2.44m) and were maintained t h e r e from September 1986 t o A p r i l 1987 on 9L/15D. In March, the l i g h t i n g p e r i o d was i n c r e a s e d 1 hour/week so t h a t by A p r i l 22 the b i r d s were on 14L/10D. 7 2. H i s t o r y o f the domestic q u a i l p a r e n t a l s t o c k : The s t r a i n o f domestic q u a i l used f o r t h i s study was the "U.B.C. w i l d type" s t r a i n ( w i l d type i n terms o f plumage c o l o u r and p a t t e r n ) . I t o r i g i n a t e d i n 1964 from q u a i l s t o c k s a t the Washington S t a t e U n i v e r s i t y (USA). These i n t u r n came from imported domestic b i r d s from Korea (1958) o u t c r o s s e d t o a domestic s t r a i n from the U n i v e r s i t y o f C a l i f o r n i a a t Davis (USA) i n 1961. The U.B.C. p o p u l a t i o n has been c l o s e d t o the immigration of new genes s i n c e 1968 and has been maintained by random-bred matings of 100 females and 50 males per g e n e r a t i o n . There had been 63 g e n e r a t i o n s by the s p r i n g o f 1986 when I began t h i s study. S i x t y c h i c k s from t h i s s t r a i n were hatched near or a t the same time as the f e r a l c h i c k s and were housed i n brooders ( i d e n t i c a l t o those used f o r the f e r a l c h i c k s ) . The domestic b i r d s were c u l l e d t o nine males and 17 females a t 6 weeks of age and p l a c e d i n a separate but i d e n t i c a l f l o o r pen next t o the f e r a l q u a i l . P E N D E S I G N The f e r a l and domestic b i r d s were e n c l o s e d i n each of two i d e n t i c a l f l i g h t - p e n s ( F i g . 2.1), each measuring 14.6 x 14.6 x 2m h i g h and covered w i t h 2cm wire c h i c k e n mesh. Both o u t e r and c e n t r e w a l l s were c o n s t r u c t e d o f plywood t o s t o p the b i r d s i n each pen from s e e i n g each o t h e r and t o reduce d i s t u r b a n c e from o u t s i d e a c t i v i t i e s . A 4m h i g h o b s e r v a t i o n tower was Figure 2.1: Pen design. 14.6m F E E D <P> WATER / MIRROR 9 p l a c e d t o g i v e an equal view o f both s i d e s ( F i g . 2.1). The f l o o r s of each of the pens were composed o f l e v e l crushed limestone upon which t h r e e rows of g r a s s , lm wide, r a d i a t e d out from the tower. T h i s minimized any v i s u a l r e s t r i c t i o n of pen a c t i v i t i e s f o r an observer i n the tower. A m i r r o r was p l a c e d a t the end of the middle g r a s s row " f u r t h e s t from the tower t o reduce b l i n d s p o t s . Two feeder l o c a t i o n s were p l a c e d i n each pen a l o n g the p e r i m e t e r w a l l s , these l o c a t i o n s had a 1 x 0.5m plywood r o o f p l a c e d above them. A water j a r was p l a c e d 2.8m from each feeder. Each pen was d i v i d e d i n t o 36 quadrants measuring 2.4 x 2.4m. Posts of the p e r i m e t e r w a l l , 2.4m a p a r t , were used as quadrant guides and s m a l l markers were p l a c e d on the ground a t the p o i n t where the c o - o r d i n a t e s met t o h e l p the observer i d e n t i f y the quadrants. Each quandrant c o u l d be i n s t a n t l y i d e n t i f i e d from the o b s e r v a t i o n tower by r e a d i n g the a p p r o p r i a t e numbers on the w a l l s between the p o s t s . EXPERIMENTAL DESIGN S i x t e e n experimental b i r d s ( e i g h t males and e i g h t females) were taken from each of the above two groups of 26 f e r a l and 2 6 domestic i n d i v i d u a l s . These were then d i v i d e d i n t o two separate groups of f o u r males and f o u r females t h a t were deployed i n the f o l l o w i n g way: 10 Group 1: 1987 Pen 1: 4 f e r a l males / 4 f e r a l females (10 months old) Pen 2: 4 domestic males / 4 domestic females (10 months old) Group 2: 1988 Pen 1: 4 domestic males / 4 domestic females (22 months old) Pen 2: 4 f e r a l males / 4 f e r a l females (22 months old) In each year the f e r a l and domestic Japanese q u a i l were housed i n the separate outdoor f l i g h t - p e n s . The pens were entered on a weekly b a s i s t o r e p l a c e feed (commercial t u r k e y s t a r t e r d i e t w i t h 26% p r o t e i n ) and water. An a d d i t i o n a l weekly check was done f o r n e s t s and eggs. The 16 experimental b i r d s not used i n the f l i g h t - p e n s i n a p a r t i c u l a r y e a r were housed i n out-door h o l d i n g pens ( F i g . 2.2, see Supplementary Study, p.13). 1. Experimental b i r d s - Group 1: On A p r i l 22, 1987 f o u r males of each s t r a i n were moved from the ind o o r f l o o r pens t o the f l i g h t - p e n s . The males were moved o u t s i d e e a r l i e r than the females because s t u d i e s of the c l o s e l y r e l a t e d Common q u a i l ( C o t u r n i x c o t u r n i x ) i n d i c a t e d t h a t males a r r i v e a t t h e i r b r e e d i n g t e r r i t o r i e s p r i o r t o the females (Moreau 1951). Females were added t o the outdoor pens two weeks l a t e r on May 6 and s y s t e m a t i c o b s e r v a t i o n s began 13 days l a t e r . The number of males was reduced on J u l y 29 by two i n each pen when i t appeared the f e r a l females were unable t o 11 Figure 2.2: Experimental Design. Feral Experimental Birds Domestic Experimental Birds Group HOLDING PENS HOLDING PENS 1987 PEN 1 PEN 2 <C 1988 > PEN 2 PEN 1 HOLDING PENS HOLDING PENS 12 n e s t because of d i s t u r b a n c e by the males. 2. Experimental b i r d s - Group 2: On A p r i l 18, 1988, f o u r males o f each s t r a i n and on A p r i l 29, f o u r females of each s t r a i n were p l a c e d i n the f l i g h t -pens, o b s e r v a t i o n s began f o u r days l a t e r . As i n 1987, i t appeared t h a t the f e r a l females were unable t o n e s t p r o p e r l y because of d i s t u r b a n c e by the males, t h e r e f o r e one male was removed from each pen on J u l y 8. A domestic female (DB2) was removed on J u l y 15 when i t was apparent t h a t she had become b l i n d . I r e p l a c e d her w i t h another female, but d i d not use data from the replacement. BIRD IDENTIFICATION Each b i r d was f i t t e d w i t h c o l o u r e d l e g bands and c o r r e s p o n d i n g c o l o u r e d p l a s t i c r i b b o n s (approx. 0.2 x 2cm) were t i e d t o t h e i r wing tags f o r q u i c k i d e n t i f i c a t i o n of i n d i v i d u a l s and sex. P r i o r t o b e i n g p l a c e d i n the outdoor pens, a l l b i r d s had t h e i r primary wing f e a t h e r s c l i p p e d (by approx. 2cm) t o prevent them from f l y i n g i n t o the wire r o o f and i n j u r i n g themselves. BEHAVIOURAL OBSERVATIONS Each year 1 hour per day of o b s e r v a t i o n s was performed on each pen of Japanese q u a i l from Monday t o F r i d a y . The hour of o b s e r v a t i o n c o n s i s t e d of h a l f an hour between 0800 and 1000 13 hours, and h a l f an hour between 1430 and 1630 hours. In a d d i t i o n t o these scheduled o b s e r v a t i o n s , one hour of c a s u a l o b s e r v a t i o n s was made on the Saturday o r Sunday of each week. Once each month a f u l l day o f o b s e r v a t i o n s was performed, s t a r t i n g a t dawn and ending a t dusk. On these days, the o b s e r v a t i o n s were c a r r i e d out f o r both pens s i m u l t a n e o u s l y every o t h e r hour. During each o b s e r v a t i o n p e r i o d the l o c a t i o n s o f a l l v i s i b l e b i r d s were reco r d e d on maps a t the s t a r t o f o b s e r v a t i o n s , 15 minutes l a t e r , and a t the end of o b s e r v a t i o n s . A l l o b s e r v a t i o n s were d i v i d e d e q u a l l y between two observers who observed on a l t e r n a t e days. The same two i n d i v i d u a l s made o b s e r v a t i o n s d u r i n g both years of the study. The o b s e r v e r s used b i n o c u l a r s (7x50), data sheets, and stop watches f o r r e c o r d i n g b e h a v i o u r a l d i s p l a y s (Appendix 1) . The l o c a t i o n o f a l l b i r d s were recorded on maps o f each pen (Appendix 2) . SUPPLEMENTARY STUDY B i r d s not used f o r the main study i n 1987 were maintained i n s m a l l outdoor " h o l d i n g pens". These b i r d s were switched w i t h the main study b i r d s i n 1988. One male and two females o f the same s t r a i n were p l a c e d i n a pen measuring 2.4 x 2.4 x 2.4m. A 3m h i g h s t r i p o f p l a s t i c was p l a c e d a t the base of the p e r i m e t e r w a l l s t o minimize d i s t u r b a n c e t o the b i r d s . The f l o o r o f each pen was d i v i d e d i n t o q u a r t e r s . Two d i a g o n a l 14 q u a r t e r s had t u r f , the remaining two had lim e s t o n e . Egg p r o d u c t i o n data from these b i r d s were i n c l u d e d w i t h data gathered from the main study and are d i s c u s s e d i n Chapter 5. DATA ANALYSIS Data were ana l y s e d by A n a l y s i s o f V a r i a n c e , t - t e s t , o r c h i - s q u a r e t e s t wherever a p p r o p r i a t e (see s p e c i f i c c h a p t e r s f o r d e t a i l ) . D i f f e r e n c e s are c o n s i d e r e d s i g n i f i c a n t i f the p r o b a b i l i t y i s l e s s than 0.05. Values t h a t f o l l o w means i n the f o l l o w i n g t e x t and bars on graphs r e p r e s e n t standard e r r o r s . 15 CHAPTER THREE COURTSHIP AND BEHAVIOURAL DISPLAYS OF FERAL AND DOMESTIC JAPANESE QUAIL INTRODUCTION Bannerman (1963) wrote of the c l o s e l y r e l a t e d European q u a i l ( C o t u r n i x c o t u r n i x ) , "There are c e r t a i n l y no s o - c a l l e d game-birds about which i t i s so d i f f i c u l t t o l e a r n t h e i r l i f e -h i s t o r y . . . To o b t a i n i n f o r m a t i o n r e g a r d i n g i t s c o u r t s h i p and br e e d i n g behaviour g e n e r a l l y when i n a w i l d s t a t e r e q u i r e s the p a t i e n c e o f Job and no l i t t l e i n g e n u i t y . " O p p o r t u n i s t i c o b s e r v a t i o n s o f w i l d Japanese q u a i l have produced c o n f l i c t i n g r e p o r t s (Kawahara 1967; Dement'ev e t a l . 1967) o f i t s mating system r a n g i n g from monogamy t o polygamy. Stevens (1961), o b s e r v i n g domestic Japanese q u a i l i n an a v i a r y , even suggested t h a t some males may share i n c u b a t i o n . The d i s p l a y s a s s o c i a t e d w i t h c o u r t s h i p , mating and ot h e r s o c i a l i n t e r a c t i o n s have not been s y s t e m a t i c a l l y observed i n w i l d Japanese q u a i l . Although s o c i a l and c o u r t s h i p d i s p l a y s have been d e s c r i b e d f o r domestic Japanese q u a i l ( F a r r i s 1964; Eynon 1968; Wilson and Bermant 1972), i t i s not known how a c c u r a t e l y they r e f l e c t those of w i l d c o n s p e c i f i c s . A comparison of p a t t e r n s o f sexual behaviour among descendants of w i l d r e d j u n g l e f o w l ( G a l l u s g a l l u s ) and domestic j u n g l e f o w l has i n d i c a t e d q u a n t i t a t i v e d i f f e r e n c e s among the c o u r t s h i p 16 d i s p l a y s ( S t a d i e 1969, c i t e d by Soss i n k a 1982). In t h i s chapter, I examine the s o c i a l i n t e r a c t i o n s , c o u r t s h i p , and mating behaviour of Japanese q u a i l i n f o u r ways. I t e s t i f d i s p l a y s r e p o r t e d f o r domestic q u a i l a re a l s o e x h i b i t e d by f e r a l q u a i l and d e s c r i b e any new d i s p l a y t h a t has not been r e p o r t e d p r e v i o u s l y . I a l s o comment on whether the d i s p l a y s are b r o a d l y s i m i l a r , and compare f r e q u e n c i e s o f the same d i s p l a y i n f e r a l and domestic q u a i l . I then determine i f the c o n t e x t under which the d i s p l a y s are e x h i b i t e d i s d i f f e r e n t by r e l a t i n g the frequency o f these d i s p l a y s t o the female's egg l a y i n g c y c l e . F i n a l l y , I t e s t i f f e r a l q u a i l form p a i r bonds d u r i n g the br e e d i n g season by n o t i n g the frequency t h a t i n d i v i d u a l males and females are "t o g e t h e r " i n the same l o c a t i o n . M E T H O D S B E H A V I O U R A L O B S E R V A T I O N S During the o b s e r v a t i o n p e r i o d s , s o c i a l i n t e r a c t i o n s and behaviours o f the b i r d s i n the pen were recorded. In p a r t i c u l a r , f r e q u e n c i e s o f occurrence were re c o r d e d f o r each d i s p l a y and behaviour p r e v i o u s l y d e s c r i b e d i n the l i t e r a t u r e (the m a j o r i t y o f which were d e s c r i b e d by F a r r i s 1964 and Eynon 1968; see a l s o P r e s t o n 1961; Beach and Inman 1965; Sachs 1966; S e l i n g e r and Bermant 1967; Wilson and Bermant 1972; Potash 17 1975; R a m e n o f s k y 1984): 1. C o u r t s h i p and s o c i a l displays: S t r u t t i n g ( S T ) : A m a l e h o l d s i t s h e a d c o c k e d i n w a r d t o w a r d s t h e f e m a l e , t h e n e c k i s f u l l y e x t e n d e d , w i n g s h e l d i n a s l i g h t d r o o p i n g p o s i t i o n , l e g s a r e f u l l y e x t e n d e d a n d h e l d i n a s t i f f p o s i t i o n . W h i l e i n t h i s p o s i t i o n t h e m a l e made a r a p i d s e r i e s o f s t e p s t o w a r d t h e f e m a l e . O c c a s i o n a l l y a m a l e w i l l p e r f o r m t h i s d i s p l a y t o a n o t h e r m a l e . Dancing (DN1) : " T h e m a l e j u m p s u p a n d down e x c i t e d l y , h i g h e r a n d h i g h e r t o 3 o r 4 f t . " ( P f e i f e r 1954, c i t e d b y E y n o n 1968) . T h i s o b s e r v a t i o n was made o n w i l d c o t u r n i x q u a i l . Dancing ( D N 2 ) : B a i l l e y (1854) d e s c r i b e d a q u a i l d a n c e a s "a m a l e r u n n i n g q u i c k l y t o t h e f e m a l e , s t o p p i n g b e s i d e h e r t u r n i n g a r o u n d h e r , d r a g g i n g h i s w i n g s , a n d s t r e t c h i n g h i s n e c k w i t h t h e f e a t h e r s o f h i s t h r o a t a n d b r e a s t p u f f e d " . T i d b i t t i n g ( T T ) : A m a l e s c r a t c h e s t h e f l o o r a n d h o l d s a p a r t i c l e o f f o o d o r d e b r i s i n h i s b e a k a n d c a l l s . U s u a l l y f e m a l e s r e s p o n d b y a p p r o a c h i n g , b u t m a l e s a l s o may r e s p o n d . N e s t C e r e m o n y ( N E ) : A r i t u a l i z e d n e s t d i s p l a y t h a t i n v o l v e s e i t h e r t h e m a l e o r f e m a l e s c r a p i n g a s h a l l o w d e p r e s s i o n i n t h e n e s t i n g s u b s t r a t e a n d s q u a t t i n g i n t h e m i d d l e o f i t . S q u a t t i n g ( S Q ) : A m a l e s q u a t s f l a t o n t h e g r o u n d w i t h h i s h e a d t u r n e d t o one s i d e f a c i n g a f e m a l e . I t i s a s u b m i s s i v e p o s t u r e d i s p l a y e d b y a m a l e t o t h e f e m a l e o r a n o t h e r m a l e . 18 Crowing (CW): A t h r e e - s y l l a b l e c a l l ("chip-per-cherr") where "the male drew i t s e l f up t o f u l l h e i g h t w i t h the neck and body extended as i t reached f o r even g r e a t e r h e i g h t by s t a n d i n g on the t i p s o f the f o r e t o e s " . 2» Mating Behaviour: For ease of q u a n t i f i c a t i o n , I have broken the mating behaviours i n t o the f o l l o w i n g s e q u e n t i a l components: Sexual Crouching (CR): A female o f t e n responds t o a male's advances by s q u a t t i n g where she i s o r by running a s h o r t d i s t a n c e i n f r o n t of the male and then s q u a t t i n g . F o l l o w i n g (FO): A r i t u a l i z e d male walk; he walks a f t e r a female w i t h a t a l l body po s t u r e and neck s t r e t c h e d out h o r i z o n t a l l y w i t h the i n t e n t t o p o s i t i o n h i m s e l f f o r mounting. B i l l - o n - n a p e (BN): A male cranes h i s neck above a female and grasps h e r by the back of the head or neck. Mounting (MO) : One o r both f e e t are p l a c e d on the female's back. T a i l b e n d i n g (TB): The male lowers h i s t a i l w h i l e p u l l i n g backwards on the neck of the female, bends h i s t a i l under the female's, i n an attempt t o achieve c l o a c a l c o n t a c t . The wings are spread f o r balance but f r e q u e n t l y both b i r d s f a l l t o one s i d e b e f o r e c l o a c a l c o n t a c t i s achieved. C l o a c a l c o n t a c t (CC): The male t h r u s t s h i s body downward and f r e e z e s momentarily when c l o a c a l c o n t a c t i s achieved. T h i s i s taken as a s i g n of a s u c c e s s f u l c o p u l a t i o n . 19 3. A g g r e s s i v e and Escape Behaviour: Chasing (CH): A male charges a f t e r a male o r female w i t h neck s t r e t c h e d and body lowered. Pecking (PK): A male o r female pecks a t another b i r d , u s u a l l y i n the head r e g i o n , w i t h a v i g o u r t h a t o f t e n e l i c i t s avoidance by the pecked b i r d . B l o c k i n g (BL): A male stands between an i n t r u d i n g male and a female. The i n t r u d e r male e i t h e r circumvents the b l o c k i n g male, r e t r e a t s or stands h i s ground. A g g r e s s i v e Crouching (AG): The female lowers her head and body t o the ground w i t h beak and t a i l p o i n t i n g upwards, forming a sharp body angle p r i o r t o c h a r g i n g a t an i n t r u d i n g male. Jumping ( J P ) : A female jumps and runs t o a v o i d c o p u l a t i n g w i t h a p e r s i s t e n t male. 4 . Male-female I n t e r a c t i o n s : S e a r c h i n g (SE): A male searches f o r a female a f t e r l o s i n g c o n t a c t w i t h her. R e t u r n i n g (RT) : A male r e t u r n i n g t o the female t h a t he l e f t t o chase a male or t o attempt c o p u l a t i o n w i t h another female. Casual F o l l o w i n g (CF): A male or female f o l l o w i n g a b i r d of the o p p o s i t e sex, o f t e n w i t h s o f t c o n t a c t v o c a l i z a t i o n . P r o s p e c t i n g (NP): A male and female walking through the g r a s s cover t o g e t h e r a p p a r e n t l y l o o k i n g f o r a s u i t a b l e n e st s i t e . Frequent stops are made when the female p u l l s and bends gr a s s stems or l o o k s about w h i l e s t a n d i n g t a l l . The two b i r d s may r e t u r n t o the same spot r e p e a t e d l y . T h i s i s an i n d i c a t i o n t h a t the female i s c l o s e t o egg l a y i n g . In a d d i t i o n , a t t e n t i o n was p a i d t o any d i s p l a y s (seemingly r i t u a l i z e d behaviour seen r e p e a t e d l y under s i m i l a r context) t h a t have not been p r e v i o u s l y d e s c r i b e d . B I R D L O C A T I O N S Each 15 min. t h a t a male and female were re c o r d e d i n the same quadrant they were g i v e n a s c o r e o f one. The a s s o c i a t i o n s c o r e f o r a p a r t i c u l a r p a i r o f b i r d s t h e r e f o r e would range from zero t o t h r e e f o r each o b s e r v a t i o n p e r i o d . On the days where o b s e r v a t i o n s were not made, the mean v a l u e s o f the score from the p r e v i o u s and the f o l l o w i n g days were a s s i g n e d . W H O L E D A Y O B S E R V A T I O N S F u l l day o b s e r v a t i o n s were conducted t o i n v e s t i g a t e the d i u r n a l p a t t e r n o f the r e p r o d u c t i v e d i s p l a y s (see Chapter 2) . D A T A A N A L Y S E S 1. Accumulated f r e q u e n c i e s o f the d i s p l a y s p e r h a l f hour of o b s e r v a t i o n f o r the p e r i o d when a l l males and females were i n the pen were used. The frequency o f each d i s p l a y was anal y s e d s e p a r a t e l y by 21 A n a l y s i s o f V a r i a n c e w i t h Repeated Measures (ANOVA, SAS 1985) u s i n g the f o l l o w i n g s t a t i s t i c a l model. Y i j k l m = U + G- + Y , + Ok + ( G Y ) f J + ( G O ) f k + (Y0) j k + E l i j k + T L + (YT)^ + (OT) k l + ( G T ) U + ( G Y T ) f J l + E2 i j k l m where i = 1,2; j = 1,2; k = 1,2; and 1 = 1,2. Vijkim = ^ e frequency o f the d i s p l a y by b i r d s o f the i t h genotype i n the j t h year, recorded by the k t h obse r v e r d u r i n g the 1th h a l f - h o u r p e r i o d o f the day. U = the p o p u l a t i o n mean; G- = the e f f e c t o f whether the b i r d s i n v o l v e d were f e r a l o r domestic; Y} = the e f f e c t o f year (1987 or 1988); 0 k = the e f f e c t o f observer; T t = the e f f e c t o f morning v e r s e s a f t e r n o o n o b s e r v a t i o n s ; (YT)^, ( G T ) U , ( 0 T ) k l , (G0) j k, (GY)^-, (YO)j k = 2-way i n t e r a c t i o n s , (GYT) i j t = 3-way i n t e r a c t i o n s , E l . . k = e r r o r term f o r the main e f f e c t s comparisons and i n t e r a c t i o n s i n v o l v i n g the main e f f e c t s , and E2.-kl|n = s u b - p l o t e r r o r term. Because the observed f r e q u e n c i e s f o r some d i s p l a y s were low, data were transformed ((Y+0.5) 0 - 5) t o s t a b i l i z e v a r i a n c e s (Snedecor and Cochran 1980) p r i o r t o a n a l y s i s . For data which had unequal v a r i a n c e s d e s p i t e the t r a n s f o r m a t i o n , a t - t e s t (SAS 1985) was used t o s e p a r a t e l y compare genotype, year, o b s e r v e r and time e f f e c t s . The Duncan's M u l t i p l e Range T e s t was used t o t e s t f o r mean s e p a r a t i o n f o r a l l the f a c t o r s t h a t were s i g n i f i c a n t by the ANOVA. 2. Accumulated f r e q u e n c i e s o f the male d i s p l a y s per male per h a l f - h o u r o b s e r v a t i o n were s o r t e d a c c o r d i n g t o the f i v e p e r i o d s o f a female's l a y i n g c y c l e : N o n - l a y i n g (NL) : 8-14 days p r i o r t o egg l a y i n g P r e - l a y i n g (PL) : 1-7 days p r i o r t o egg l a y i n g L a y i n g (L) : egg l a y i n g - i n c l u d e s days f o r f i r s t and l a s t egg of each c l u t c h I n c u b a t i n g (I) : i n c l u d e s f i r s t day o f i n c u b a t i o n t o h a t c h i n g . Brooding (B) : from the f i r s t day c h i c k s l e a v e nest u n t i l t h e i r a c t i v i t i e s a re independent of the female. Data not f a l l i n g i n t o one of these f i v e p e r i o d s were not used. The mean v a l u e s f o r the v a r i o u s a c t i v i t i e s were lower than those i n the f i r s t a n a l y s es (1) because the means were f r e q u e n c i e s p e r male per o b s e r v a t i o n i n s t e a d o f per pen per o b s e r v a t i o n , and a l s o because t h i s s e t o f data i n c l u d e d the l a t e r p a r t o f the br e e d i n g season when some males were removed from the pens. Furthermore, d u r i n g the i n c u b a t i o n and brooding p e r i o d s , the l e v e l o f a c t i v i t i e s were much reduced. The s e l e c t e d data were analysed by the A n a l y s i s o f V a r i a n c e (ANOVA, SAS 1985)) u s i n g the f o l l o w i n g s t a t i s t i c a l model: YUkl = U + G i + Y j + P k + (GY) f J + (GP),-k + (YP) j k + (GYP) j j k + E i j k l where i = 1,2; j = 1,2; and k = 1,2,..5 Y i j k i = t h e cumulative frequency o f the d i s p l a y d i r e c t e d a t females of the i t h genotype d u r i n g the k t h p e r i o d o f her l a y i n g c y c l e i n the j t h year. U = the p o p u l a t i o n mean; G. = the e f f e c t o f whether i t was a f e r a l o r domestic female; Yj = the e f f e c t o f year (1987 and 1988) ; P k = t h e e f f e c t o f d i f f e r e n t p e r i o d s o f the female's l a y i n g c y c l e ; (GP) 1 k, (GY)^., and (YP)j k = 2-way i n t e r a c t i o n s , ( G Y P ) ^ = 3-way i n t e r a c t i o n , and Eij.|cl = e r r o r term. The same t r a n s f o r m a t i o n as i n (1) t o s t a b i l i z e v a r i a n c e s was a p p l i e d t o the data, and data which had unequal v a r i a n c e s were a n a l y s e d as i n (1). S i n c e o b s e r v e r e f f e c t s were s i g n i f i c a n t i n o n l y one of the d i s p l a y s a n a l y s e d i n (1), t h i s f a c t o r was dropped from the s t a t i s t i c a l model. 3. Data from the f i v e f u l l days (1987: 2 days; 1988: 3 days) of o b s e r v a t i o n s were combined. Data were summarized as means per male per h a l f - h o u r o f o b s e r v a t i o n . Student's t - t e s t s (Snedecor and Cochran 1980) were used t o t e s t f o r d i f f e r e n c e s i n frequency by the f e r a l and domestic males i n each d i s p l a y a t each of the seven time p o i n t s . P a i r e d t - t e s t s were conducted t o determine i f the o v e r a l l frequency o f each d i s p l a y was d i f f e r e n t f o r f e r a l and domestic q u a i l . A l l data were l o g transformed (Y = LnX) t o normalize the data (Snedecor and Cochran 1980). 24 R E S U L T S NEW D I S P L A Y S A l l except two of the p r e v i o u s l y r e p o r t e d c o u r t s h i p d i s p l a y s and b e h a v i o u r a l components e x h i b i t e d by domestic Japanese q u a i l were performed by both the domestic and the f e r a l q u a i l . Two c o u r t s h i p dancing d i s p l a y s (DN1 and DN2), d e s c r i b e d by P f e i f e r (1954) and B a i l l e y (1854) r e s p e c t i v e l y , were not seen. However two r a r e and h i t h e r t o unreported behaviours were observed. The f i r s t was a c o u r t s h i p dancing d i s p l a y (DN). I t was performed by both f e r a l (n=10) and domestic q u a i l (n=3) (Appendix 3) . A male o r female would dash i n a z i g - z a g p a t t e r n , c r o u c h i n g b r i e f l y a t the t u r n s , o r run w i t h wings f l a p p i n g and i n t e r m i t t e n t l y hop i n a f i g u r e - 8 p a t t e r n , around one or two o t h e r i n d i v i d u a l s of e i t h e r sex. T h i s dancing d i s p l a y d i d not resemble e i t h e r DN1 o r DN2. I t was u s u a l l y but not e x c l u s i v e l y performed by a p a i r e d male i n the presence of h i s mate. The second unreported d i s p l a y , " l e a d i n g " (LD), was performed by both f e r a l (n=12) and domestic males (n=8). In t h i s d i s p l a y , a male s t r e t c h e d h i s neck forward and walked i n f r o n t o f the female, who u s u a l l y f o l l o w e d him. I f the female f a i l e d t o f o l l o w the male performing t h i s d i s p l a y , the male sometimes became a g g r e s s i v e and pecked the female. 25 DISPLAY FREQUENCIES OF FERAL AND DOMESTIC QUAIL 1. C o u r t s h i p and s o c i a l d i s p l a y s : F e r a l males performed the c o u r t s h i p d i s p l a y t i d b i t t i n g (TT) s i g n i f i c a n t l y more f r e q u e n t l y than domestic males (Table 3.1) . During f u l l - d a y o b s e r v a t i o n s , f e r a l males a l s o s t r u t t e d (ST) t o the females s i g n i f i c a n t l y more f r e q u e n t l y than domestic males ( F i g . 3.1). When a l l f o u r males were i n the f l i g h t pen, domestic males crowed (CW) more f r e q u e n t l y than f e r a l males (Table 3.1). However, when the data o n l y i n c l u d e d the d i f f e r e n t p e r i o d s of the female's l a y i n g c y c l e , f e r a l males crowed s i g n i f i c a n t l y more f r e q u e n t l y than domestic males d u r i n g t h e i r mate's n o n - l a y i n g , p r e - l a y i n g and l a y i n g p e r i o d s ( F i g . 3.2). There were no s i g n i f i c a n t d i f f e r e n c e s between f e r a l and domestic males i n o t h e r c o u r t s h i p d i s p l a y s (Table 3.1) . 2. Mating Behaviour: T a i l b e n d i n g (TB) and c l o a c a l c o n t a c t s (CC) by domestic males and sexual crouch (CR) by domestic females were s i g n i f i c a n t l y h i g h e r than by f e r a l males and females r e s p e c t i v e l y (Table 3.1). The f e r a l males showed a s i g n i f i c a n t i n c r e a s e i n TB from 1987 and 1988 (n=36; X=0.24+0.09 and n=36; X=0.73±0.12 r e s p e c t i v e l y ) , w h i l e domestics showed no d i f f e r e n c e over the two y e a r s (n=36; X=1.00+0.12 and n=36; X=1.16+0.21 26 T a b l e 3.1: Q u a n t i t a t i v e d i f f e r e n c e s i n c o u r t s h i p d i s p l a y s and mating behaviour between f e r a l and domestic Japanese q u a i l . Behaviour Frequency 1 N F e r a l N Domestic C o u r t s h i p d i s p l a y s TT ST NE CW 72 72 72 72 0.12+0.04 0.72+0.11 0.02+0.01 5.92+1.04 72 72 72 72 0.03+0.01 0.77+0.09 0.04+0.02 8.62+1.03 (*) Mating behaviour FO CR BN MO TB CC 72 72 72 72 72 72 0.68+0.09 0.37+0.06 1.88+0.22 1.50+0.18 0.48+0.08 0.26+0.05 72 72 72 72 72 72 1.22+0.16 0.52+0.09 83+0.31 35+0.25 08+0.12 52+0.07 2. 2. 1. 0. ** Male-female i n t e r a c t i o n s CF RT SE 72 72 72 2.37+0.30 0.81+0.11 0.48+0.09 72 72 72 2.95+0.28 0.51+0.08 0.39+0.08 Ag g r e s i v e behaviour CH BL 72 72 2.91+0.46 0.88+0.16 72 72 2.43+0.36 0.73+0.09 mean frequency per pen per h a l f - h o u r o f o b s e r v a t i o n . 2 number o f samples ** P<0.01; * P<0.05 ANOVA (*) P<0.05 t - t e s t Figure 3.1: Diurnal pattern of males returning and strutting to females during 5 full days of observation. 0.75 Male returning to female 0.50 0.25. 0 0.50 0.25 630 830 1030 1230 1530 1700 1900 2100 Time of day Feral Domestic Figure 3.2: Differences in frequencies of male crowing between feral and domestic males with relation to the mate's laying cycle. Crowing Frequency m e a n f r e q u e n c y p e r m a l e p e r h a l f - h o u r of o b s e r v a t i o n M e a n s w i t h t h e s a m e l e t t e r a r e n o t s i g n i f i c a n t l y d i f f e r e n t M e a n s w i t h d i f f e r e n t l e t t e r s a re s i g n i f i c a n t l y d i f f e r e n t by D u n c a n ' s M u l t i p l e R a n g e Tes t . Laying Cycle 29 r e s p e c t i v e l y ) . There was a s i g n i f i c a n t genotype by o b s e r v e r e f f e c t f o r TB, i n d i c a t i n g t h a t w h i l e the two observers were c o n s i s t e n t i n r e c o r d i n g the TB frequency f o r f e r a l males, they d i f f e r e d i n r e c o r d i n g the TB frequency f o r domestic males. T h i s i s p r o b a b l y because o f the h i g h e r frequency of f o r c e d e x t r a - p a i r c o p u l a t i o n s performed by domestic males (see Chapter 4) , which, because the female was s t r u g g l i n g , l e a d t o bouts of m u l t i p l e TB b e f o r e CC. T h i s made i t d i f f i c u l t t o r e c o r d the exact number of TB. Frequencies of f o l l o w i n g (FO), b i l l - o n - n a p e (BN) and mounting (MO) were not s i g n i f i c a n t l y d i f f e r e n t between f e r a l and domestic males (Table 3.1). 3. A g g r e s s i v e and escape behaviour: F e r a l and domestic q u a i l d i d not d i f f e r i n any a g g r e s s i v e or escape behaviours. However, when c h a s i n g and pecking behaviours from the whole-day o b s e r v a t i o n s were analyzed f o r male c h a s i n g male, male c h a s i n g female, and female c h a s i n g female, f e r a l males were s i g n i f i c a n t l y more a g g r e s s i v e towards an i n d i v i d u a l o f e i t h e r sex than domestic males ( F i g . 3.3). On the o t h e r hand, domestic females were s i g n i f i c a n t l y more a g g r e s s i v e towards males than f e r a l females ( F i g . 3.3). 4 . Male-female i n t e r a c t i o n s : More c a s u a l f o l l o w i n g (CF) was performed by domestic 30 Figure 3.3: Diurnal pattern of aggressive behaviour between males and females during 5 days of observation. M e a n f r e q u e n c y P e r h a h o u r P e r m a Male pecking male T Male chasing male o 0 . 3 „ 0.2 0.1 0 1 0.5 Male pecking female Female pecking male I 1.5^ 0.5 0.5 0 . 2 5 J Female chasing male 630 830 1030 1230 1530 1700 1900 630 8 3 0 1030 1230 1530 1700 1900 Feral _ . , , Time of day Domest ic q u a i l than f e r a l q u a i l (Table 3.1). A f t e r s e p a r a t i n g from t h e i r mate, f e r a l males r e t u r n e d (RT) t o t h e i r mate s i g n i f i c a n t l y more o f t e n than domestic males (Table 3.1). The d i f f e r e n c e was most apparent i n the morning between 0830 and 1030 hours as w e l l as i n the l a t e a f t e r n o o n between 1900 and 2100 hours. F e r a l and domestic q u a i l d i d not d i f f e r i n s e a r c h i n g (SE) behaviour (Table 3.1). MALE DISPLAY FREQUENCIES DURING THE LAYING CYCLE Summed over both the f e r a l and domestic males, t h e r e were s i g n i f i c a n t d i f f e r e n c e s i n d i s p l a y s and behaviours (CF, SE, RT, CH and CC) d u r i n g the females' l a y i n g c y c l e ( F i g . 3.4 and 3 . 5 ) . A c t i v i t i e s peaked d u r i n g egg l a y i n g and dropped d u r i n g the brooding p e r i o d . DISPLAY FREQUENCIES DURING THE MORNING AND AFTERNOON OBSERVATION PERIODS Most d i s p l a y s and a c t i v i t i e s decreased s i g n i f i c a n t l y i n frequency from morning t o a f t e r n o o n ( F i g . 3.6). Only one c o u r t s h i p (CW) and one a c t i v i t y (NP) showed the o p p o s i t e t r e n d (Table 3.1). DISPLAY FREQUENCIES DURING 1987 AND 1988 Summing over both f e r a l and domestic q u a i l , t h e r e were h i g h e r f r e q u e n c i e s of mating behaviours (CR, TB) , male-female Figure 3.4: Frequencies of Male-Female interactions with relation to the mate's laying cycle (feral and domestic combined). Display Frequency 1 N Casual Following Returning To Searching 1 m e a n f r e q u e n c y p e r m a l e p e r h a l f - h o u r of o b s e r v a t i o n For e a c h b e h a v i o r , m e a n s w i t h t h e s a m e l e t t e r a r e n o t s i g n i f i c a n t l y d i f f e r e n t . M e a n s w i t h d i f f e r e n t l e t t e r s a r e s i g n i f i c a n t l y d i f f e r e n t by D u n c a n ' s M u l t i p l e R a n g e Te s t. Laying Cycle Figure 3.5: Frequencies of male aggressive and mating behaviour with relation to the mate's laying cycle (feral and domestic combined). Display Frequency m e a n f r e q u e n c y p e r m a l e p e r h a l f - h o u r of o b s e r v a t i o n For e a c h b e h a v i o u r , m e a n s w i t h t h e s a m e l e t t e r are n o t s i g n i f i c a n t l y d i f f e r e n t . M e a n s w i t h d i f f e r e n t l e t t e r s a r e s i g n i f i c a n t l y d i f f e r e n t by D u n c a n ' s M u l t i p l e R a n g e T e s t . Laying Cycle Figure 3.6: Differences in courtship displays and mating behaviour between morning and afternoon (feral and domestic combined). Display Frequency 1 10 i * P < 0 . 0 5 P<0.01 N = 72 s a m p l e s i n t e r a c t i o n s (CF, RT) , and c o u r t s h i p (CW) i n 1988 than 1987 (Table 3.2). While NE showed the o p p o s i t e t r e n d , c a u t i o n i s needed i n i n t e r p r e t i n g t h i s g i v e n the low frequency. The frequency o f mounting (MO) was s i g n i f i c a n t l y h i g h e r i n the morning (n=36; X=3.02+0.40) than i n the a f t e r n o o n (n=36; X=0.73+0.14) d u r i n g 1987, and showed a s i m i l a r t r e n d i n 1988 (n=36; X=2.24+0.28 and n=36; X=1.70+0.25 r e s p e c t i v e l y ) . INDIVIDUAL ASSOCIATIONS The a s s o c i a t i o n s c o r e s o f each male were p l o t t e d a g a i n s t the f o u r females i n the f l i g h t - p e n f o r each b r e e d i n g season ( f o r e.g. see F i g s . 3.7 & 3.8; the remaining a s s o c i a t i o n p r o f i l e s can be found i n Appendix 4) . S i x o f e i g h t f e r a l males showed c l e a r t r e n d s o f a s s o c i a t i n g w i t h the same female through the br e e d i n g season ( F i g . 3.7). Domestic males a l s o a s s o c i a t e d w i t h one female a t a time, but had s e v e r a l a s s o c i a t i o n s throughout the season each u s u a l l y i n i t i a t e d j u s t p r i o r t o egg l a y i n g . DISCUSSION COURTSHIP DISPLAYS Most d e s c r i p t i o n s of the c o u r t s h i p d i s p l a y s o f domestic Japanese q u a i l i n the l i t e r a t u r e a re u s u a l l y b r i e f o r i n c o n s i s t e n t and the context i n which the d i s p l a y s were performed i s not always c l e a r . N e v e r t h e l e s s , the d i s p l a y s I 36 T a b l e 3 . 2 : Q u a n t i t a t i v e d i f f e r e n c e s i n c o u r t s h i p d i s p l a y s and mating behaviour between 1987 and 1988 ( f e r a l and domestic combined). Behaviour Frequency 1 N 1987 N 1988 C o u r t s h i p d i s p l a y s NE CW 72 72 0.05+0.02 6.31+0.89 72 72 0.01+0.01 8.22+1.89 (*) Mating behaviour CR TB 72 72 0.26+0.06 0.62+0.09 72 72 0.63±0.09 0.94+0.12 ** ** Male-female i n t e r a c t i o n s CF RT 72 72 1.91+0.21 0.53+0.09 72 72 3.40+0.33 0.79+0.10 ** * frequency p e r pen per h a l f - h o u r o f o b s e r v a t i o n . N =number of samples ** P<0.01; * P<0.05 ANOVA (*) P<0.05 t - t e s t F E M A L E FERAL MALE - FYG2 - 1988 Male - Female Associat ion (Total Score / Observat ion period) FB2 3 2 1 0 FG2 3 2 1 0 F Y 2 3 2 1 0 FP2 3 2 1 A A A A A A A A May 2 May 17 Jun 1 Jun 16 • PAIR B O N D (see Chapter 4) Jul 1 Ju l 16 Jul 31 Aug 15 EGG Aug 30 C Y C L E ^ I N C U B A T E O Y O L b | H B R O O D LAY F E M A L E DB2 3 DOMESTIC MALE - DYG2 - 1988 Male - Female Associat ion (Total S c o r e / Observat ion per iod) May 2 May 17 Jun 1 Jun 16 • PAIR B O N D (see Chapter 4) Ju l 1 Ju l 16 J u l 31 Aug 15 EGG CYCLE Aug 30 LAY INCUBATE Hi BROOD observed i n t h i s study match those r e p o r t e d i n the l i t e r a t u r e except f o r the two c o u r t s h i p dances ( P f e i f e r 1954; B a i l l e y 1854). P f e i f e r (1954) made h i s o b s e r v a t i o n (DN1) on w i l d c o t u r n i x q u a i l (C. c o t u r n i x ) and t h e i r d i s p l a y s c o u l d d i f f e r from Japanese q u a i l (C. i a p o n i c a ) . Eynon (1968) a l s o commented t h a t t h i s d i s p l a y was not observed i n h i s study. A f t e r o b s e r v i n g the Japanese q u a i l i n the f l i g h t - p e n s , I c o n s i d e r t h a t what B a i l l e y (1854) d e s c r i b e d as dancing (DN2) i s j u s t a v a r i a t i o n of s t r u t t i n g (ST). Cheng (unpubl. data) observed mating behaviour of domestic Japanese q u a i l u s i n g two-male, s i x - f e m a l e mating groups i n 2.5 x 3.1m indoor f l o o r - p e n s . Under t h i s s i t u a t i o n , a male c o u l d dominate the o t h e r male and attempt t o keep the s u b o r d i n a t e male from mating w i t h the females. Often the s u b o r d i n a t e male would dash i n t o a group o f females i n a z i g -zag manner and the females responded by s c a t t e r i n g i n d i f f e r e n t d i r e c t i o n s , c r o u c h i n g and hopping up i n the a i r . The s u b o r d i n a t e male, and sometimes both males, would s t a r t t o b i l l - o n - n a p e and mount females. The c o n t e x t and the behaviour of the b i r d s were s i m i l a r t o the dancing (DN) d i s p l a y s observed i n the f l i g h t - p e n s . Cheng (pers. comm.) hyp o t h e s i z e d t h a t f u n c t i o n a l l y the male "dance" was an a t t e n t i o n d i v e r t i n g t a c t i c and the female "dance" seemed t o be e v a s i v e a c t i o n . My o b s e r v a t i o n s suggest t h a t t h i s i s a means by which a b i r d can get the a t t e n t i o n of a p o s s i b l e mate. However, more o b s e r v a t i o n s are needed t o see i f t h i s d i s p l a y i s performed c o n s i s t e n t l y under the same con t e x t t o c o n f i r m t h a t i t i s a c o u r t s h i p dance. DEVIATIONS OF DOMESTIC QUAIL FROM FERAL QUAIL So s s i n k a (1982) commented t h a t f o r most g a l l i n a c e o u s s p e c i e s s t u d i e d , d o m e s t i c a t i o n caused q u a n t i t a t i v e ( f r e q u e n c i e s ) but no obvious q u a l i t a t i v e (body p o s i t i o n s ) changes i n c o u r t s h i p and sexual behaviour. S i m i l a r l y , i n my study of Japanese q u a i l a l l of the d i s p l a y s observed i n domestic q u a i l were observed i n f e r a l q u a i l , and v i c e v e r s a . The d i f f e r e n c e between f e r a l and domestic q u a i l were i n f r e q u e n c i e s of occurrence and, i n some cases, i n the context under which the d i s p l a y s were performed. Domestic males performed l e s s c o u r t s h i p d i s p l a y s , but c o p u l a t e d more f r e q u e n t l y than f e r a l males. T h i s i s c o n s i s t e n t with Sossinka's (1982) review of domestic g a l l i f o r m and a n s e r i f o r m s p e c i e s . The crowing of the males i s of s p e c i a l i n t e r e s t . Japanese q u a i l were o r i g i n a l l y domesticated i n Japan as a song b i r d , and many v a r i e t i e s were bred by the samurais f o r t h e i r d i f f e r e n t crows (Yamashina 1961). A l l t h e s e domestic v a r i e t i e s d isappeared d u r i n g World War I I . S i m i l a r l y , many breeds of domestic chickens d i f f e r i n the l e n g t h , a c c e n t u a t i o n , p i t c h , and timbre of the t h e i r crows (Sossinka 1982) . Observations from my f l i g h t - p e n study i n d i c a t e d t h a t the domestic males produce a more u n i f o r m l y l o u d crow, w h i l e the f e r a l males v a r i e d the loudness o f the crow and the l e n g t h o f the l a s t note depending upon the d i s t a n c e between them and t h e i r mates. F e r a l males u s u a l l y crowed w h i l e separated from t h e i r mates, probably t o s t a y i n c o n t a c t w i t h them. T h i s peaked d u r i n g the p r e - l a y and l a y p e r i o d when the males most f r e q u e n t l y separated form t h e i r mates w h i l e c h a s i n g away i n t r u d i n g males. Domestic males, however, crowed f r e q u e n t l y i n the b e g i n n i n g of the b r e e d i n g season b e f o r e the i n d i v i d u a l s were p a i r e d , and the frequency of t h e i r crows d i d not v a r y a c c o r d i n g t o the l a y i n g p e r i o d s of t h e i r mate. I t seems t h a t not o n l y was the frequency of the crow d i f f e r e n t between the f e r a l and domestic males, but so was the c o n t e x t under which they crowed. I t i s no wonder t h a t r e s e a r c h e r s s t u d y i n g the f u n c t i o n of the crow of Japanese q u a i l u s i n g domestic s t r a i n s under l a b o r a t o r y c o n d i t i o n s drew c o n f l i c t i n g c o n c l u s i o n s (e.g. Potash 1975; O t t i n g e r e t a l . 1982; Wada 1986). A h i g h frequency of a g g r e s s i o n (chasing/pecking) by males t o females o c c u r r e d p r i o r t o p a i r f ormation. As p a i r s formed, a g g r e s s i o n decreased i n frequency and was r e p l a c e d w i t h c o u r t s h i p and mating behaviour. S t u d i e s o f o t h e r g a l l i n a c e o u s b i r d s (Wiley 1973; Menzdorf 1982) showed t h a t s u c c e s s f u l p a i r f o r m a t i o n i n v o l v e d a degree of i n i t i a l f e a r and h o s t i l i t y . I found t h a t f e r a l males were s i g n i f i c a n t l y more a g g r e s s i v e t o females than were domestic males and i t appears t h i s a g g r e s s i o n p l a y s a s i g n i f i c a n t r o l e , not o n l y i n p a i r f ormation, but a l s o i t s maintenance. 42 Male-male a g g r e s s i o n was s i g n i f i c a n t l y h i g h e r i n the f e r a l q u a i l than domestic q u a i l . F e r a l males were v e r y a l e r t t o i n t r u d i n g males and responded by b l o c k i n g and, i f t h a t f a i l e d , c h a s i n g and pecking. T h i s not o n l y p r o t e c t e d t h e i r mates from the advances o f i n t r u d i n g males, but c o u l d have allowed females t o e v a l u a t e the males as p o t e n t i a l mates (Wiley 1973). S i n c e domestic males p r o t e c t e d t h e i r females s i g n i f i c a n t l y l e s s than f e r a l males, the domestic females had t o spend more time f i g h t i n g o f f male i n t r u d e r s . From the a s s o c i a t i o n i n d i c e s of the f e r a l males w i t h the d i f f e r e n t f e r a l females, and my b e h a v i o r a l o b s e r v a t i o n s , i t i s apparent t h a t male and female f e r a l Japanese q u a i l form a p a i r bond d u r i n g the b r e e d i n g season. During the f i r s t two weeks a f t e r the b i r d s were p l a c e d i n the f l i g h t - p e n s , the e i g h t b i r d s mingled as a group and showed no s i g n s o f p a i r f o r m a t i o n . The i n i t i a l p a i r i n g took s e v e r a l weeks t o e s t a b l i s h i n 1987, whereas p a i r i n g was i n i t i a t e d w i t h i n a few days i n 1988. Most f e r a l males a s s o c i a t e d w i t h o n l y one female through the whole b r e e d i n g season. They were q u i c k t o r e t u r n t o t h e i r mate when separated, even d u r i n g the non-l a y i n g p e r i o d of the mate. T h i s was not t r u e f o r domestic males. They spent more time c a s u a l l y f o l l o w i n g females and l o o k i n g f o r an o p p o r t u n i t y f o r c o p u l a t i o n (see Chapter 4). The d i f f e r e n c e i n p a i r i n g t a c t i c s between f e r a l and domestic q u a i l w i l l be examined i n more d e t a i l i n Chapter 4. In summary, I documented the temporal and d i u r n a l p a t t e r n o f mating a c t i v i t i e s and d i s p l a y s o f f e r a l Japanese q u a i l i n a s e m i - n a t u r a l s e t t i n g . A l l c o u r t s h i p d i s p l a y s observed i n domestic Japanese q u a i l were observed i n f e r a l q u a i l . However, q u a n t i t a t i v e and c o n t e x t u a l d i f f e r e n c e s suggest t h a t some communication s i g n a l s (e.g. crowing) became l e s s s p e c i f i c i n domestic q u a i l because the s i g n a l s were no l o n g e r e s s e n t i a l f o r the s u r v i v a l and/or r e p r o d u c t i o n o f these i n d i v i d u a l s i n c a p t i v i t y . 44 CHAPTER FOUR THE PAIR BONDS OF FERAL AND DOMESTIC JAPANESE QUAIL INTRODUCTION Recent advances i n the study o f a v i a n mating systems r e v e a l t h a t they are much more f l e x i b l e than had been p r e v i o u s l y thought (Gibbs e t a l . 1990; Mock and F u j i o k a 1990). S t u d i e s o f "monogamous s p e c i e s " i n d i c a t e t h a t both sexes p r a c t i c e v a r i o u s s t r a t e g i e s o f mate i n f i d e l i t y t o i n c r e a s e r e p r o d u c t i v e success (Gladstone 1979; McKinney e t a l . 1983). However, the o r i g i n a l d e f i n i t i o n f o r monogamy, "one male forms a p a i r bond w i t h one female" does not take i n t o account e x t r a -p a i r mating by e i t h e r o f the p a i r . Two o f these s t r a t e g i e s , e x t r a - p a i r c o p u l a t i o n s (Ford 1983; McKinney e t a l . 1984; Birkhead 1987) and mate-switching (Carey and Nowlan 1975; P r i c e 1984) , make i t d i f f i c u l t , and i n some cases i n a p p r o p r i a t e , t o c l a s s i f y the mating system o f a s p e c i e s as monogamous or polygamous a c c o r d i n g t o t r a d i t i o n a l d e f i n i t i o n s (Lack 1968; Wittenberger 1979). Thus i n c o n s i s t e n c i e s i n the l i t e r a t u r e become apparent: Ford (1983) notes t h a t w h i l e many authors l a b e l mate-switching between b r e e d i n g attempts as polygamy (Fraga 1972; Burns 1982), o t h e r s l a b e l them as s u c c e s s i v e monogamy (Carey and Nolan 1975). In t h i s chapter, I compare the mating system o f f e r a l and domestic Japanese q u a i l . I do t h i s by f i r s t d e v e l o p i n g a working d e f i n i t i o n o f a p a i r bond, s p e c i f i c t o the c o n d i t i o n s of t h i s s p e c i e s . Under t h i s framework, I compare f e r a l and domestic q u a i l i n : 1) t i m i n g and d u r a t i o n o f a s s o c i a t i o n s ; 2) b e h a v i o u r a l d i s p l a y s ; 3) mate quarding; and 4) r e l a t i o n s h i p t o female's l a y i n g c y c l e . I t i s not my i n t e n t i o n t o r e - e v a l u a t e the concept o f p a i r bond o r t o examine the e v o l u t i o n o f a v i a n mating systems. METHODS DEFINITION Eynon (19 68), i n h i s s t u d i e s o f domestic Japanese q u a i l , d e f i n e d p a i r bond as "when a male [ i s ] r e p e a t e d l y s t a y i n g c l o s e t o a female and p o s i t i o n i n g h i m s e l f between her and othe r males". T h i s d e f i n i t i o n p r o v i d e s a u s e f u l s t a r t i n g p o i n t , however, i t does not d e f i n e when a p a i r bond begins and ends. As a working d e f i n i t i o n o f p a i r i n g I use the f o l l o w i n g t h r e e c r i t e r i a t o estimate the l e n g t h o f a p a i r bond. While p a i r bonds do not a c t u a l l y s t a r t and end suddenly, these c r i t e r i a p r o v i d e d me w i t h measurable end p o i n t s : 1. A seven-day p e r i o d i n which a p a i r had an a s s o c i a t i o n s c o r e (see Chapter 3 and Appendix 4) g r e a t e r than zero f o r a t l e a s t f i v e days, and no c o n s e c u t i v e days of zero. B l o c k i n g (BL) by the male must occur d u r i n g t h i s time. T h e r e f o r e , the minimum d u r a t i o n o f a p a i r bond would be seven days. 2. A f t e r the f i r s t day o f the bond has been as s i g n e d , the c o n t i n u a t i o n o f the bond beyond the seventh day i s i n d i c a t e d by c o n s e c u t i v e days of an a s s o c i a t i o n s c o r e o f not l e s s than one. 3. The end of the p a i r bond i s d e f i n e d by t h e f o l l o w i n g c r i t e r i a : (a) Two c o n s e c u t i v e days where the s c o r e was zero, or (b) f o r more than t h r e e c o n s e c u t i v e days, the s c o r e was l e s s than one and none o f the f o l l o w i n g behaviours were observed; ( i ) b l o c k i n g by the male f o r h i s mate, and the female d i d not run away or h i d e from the male, ( i i ) the two b i r d s f e e d i n g t o g e t h e r , or ( i i i ) the male searched f o r and r e t u r n e d t o h i s mate a f t e r he l e f t her s i d e . BEHAVIOURAL OBSERVATIONS Be h a v i o u r a l o b s e r v a t i o n s were c a r r i e d out as o u t l i n e d i n Chapter 3. The data f o r c o u r t s h i p d i s p l a y s and mating a c t i v i t i e s i n c l u d e o n l y the p e r i o d i n each y e a r when a l l f o u r males and f o u r females o f each type were i n the pens (1987: May 20-July 28, 1988: May 2 - J u l y 7 ) . Otherwise, data were taken from the whole b r e e d i n g season (see a l s o Chapter 2: General Methods). The l a y i n g c y c l e was d i v i d e d i n t o t h r e e p e r i o d s : non-lay (NL = 8-14 days p r i o r t o l a y i n g ) ; p r e - l a y (PL = 1-7 days p r i o r t o l a y i n g ) and l a y ( L ) . In some i n s t a n c e s l a y was s u b d i v i d e d i n t o l a y 1 ( L l = f i r s t 5 days o f l a y i n g ) and l a y 2 (L2 = remaining days of l a y i n g ) . 47 DATA ANALYSES 1. The d u r a t i o n s of p a i r bonds were a n a l y s e d by a t -t e s t . Log t r a n s f o r m a t i o n (Y = Ln X) was done t o n o r m a l i z e the data (Snedecor and Cochran 1980). 2. The d u r a t i o n o f the p a i r bond p r i o r t o e g g - l a y i n g was c a l c u l a t e d by c o u n t i n g the f i r s t day o f e g g - l a y i n g as zero. Thus, f o r example, a male t h a t f i r s t p a i r e d w i t h a female two days a f t e r her f i r s t e g g - l a y i n g was g i v e n a v a l u e of minus two and a male t h a t p a i r e d two days p r i o r t o e g g - l a y i n g was g i v e n a two. The data were analyzed by t - t e s t . 3. The d i s t r i b u t i o n of the p a i r e d males• mating a c t i v i t i e s was analysed by Chi-square t e s t s ( 2 x 2 and 2 x 3 contingency t a b l e s ) . 4. The r a t e of s u c c e s s f u l c o p u l a t i o n was c a l c u l a t e d by d i v i d i n g the observed frequency of c l o a c a l c o n t a c t s (an i n d i c a t i o n of s u c c e s s f u l c o p u l a t i o n ) by the observed frequency of b i l l - o n - n a p e s (an i n d i c a t i o n o f a male i n t e n d i n g t o mount the female) and expressed as a percentage. The percentages were a r c - s i n transformed b e f o r e a n a l y s i s (Snedecor and Cochran 1980). The f o l l o w i n g s t a t i s t i c a l model was used i n an a n a l y s i s of v a r i a n c e (PROC GLM; SAS 1985): Y f J k l = U + Gf + A j + P k + (GA),, + (GP), k + (AP) j k + ( G A P ) | J k + E,.jkl where i = 1,2; j = 1,2; and k = 1,2,3 Y.jk l = the r a t e of s u c c e s s f u l c o p u l a t i o n of a male of the i t h genotype performing the j t h type of c o p u l a t i o n d u r i n g the 48 k t h p e r i o d of the l a y i n g c y c l e o f h i s mate. U = the p o p u l a t i o n mean; G. = the e f f e c t o f whether the male i n v o l v e d was f e r a l o r domestic; Aj = the e f f e c t o f whether i t was a w i t h i n - p a i r or e x t r a - p a i r c o p u l a t i o n attempt; P k = the e f f e c t o f mate's l a y i n g c y c l e ; (GP) ] k, (GA).j, ( p A ) j k = 2-way i n t e r a c t i o n s , (GPA) j j k = 3-way i n t e r a c t i o n ; and E1-J-kl = the e r r o r term. 5. The average l e n g t h of time f o r a male t o r e t u r n t o h i s mate (mean l a t e n c y ) a f t e r he l e f t her ( u s u a l l y f o r : 1. mate-guarding, i.e.male-male b l o c k i n g and c h a s i n g ; 2. e x t r a -p a i r compulations; and 3. maintenance a c t i v i t i e s , e.g. f e e d i n g and d r i n k i n g ) were analysed by a n a l y s i s o f v a r i a n c e (PROC GLM, SAS 1985) w i t h the f o l l o w i n g s t a t i s t i c a l model: Y i j k l - U + Gf + P J + A k + (GP),, + (GA) I k + (PA) j k + (GPA) 1 j k + E,-jkl where i = 1,2; j = 1,2,3,4; and k = 1,2,3 Y i j k i = t n e l a t e n c y ( i n seconds) o f r e t u r n i n g t o the female by male of the i t h genotype a f t e r the k t h type of a c t i v i t i e s d u r i n g the j t h p e r i o d o f her l a y i n g c y c l e . U = the p o p u l a t i o n mean; Gi = the e f f e c t of whether the male i n v o l v e d was f e r a l o r domestic; Pj = the e f f e c t of the female's l a y i n g c y c l e ; A k = the e f f e c t of type o f a c t i v i t y the male was engaged i n w h i l e away from the female; (GP)^., (GA) j k, (PA)j k = 2-way i n t e r a c t i o n s , (GPA) i j k = 3-way i n t e r a c t i o n ; and E i j- k l = the e r r o r term. 6. The amount of time ( i n seconds) a male l e f t h i s mate unattended d u r i n g a h a l f - h o u r o b s e r v a t i o n p e r i o d was c a l c u l a t e d by m u l t i p l y i n g the mean l a t e n c y o f r e t u r n i n g t o the female by the mean frequency of t h a t male l e a v i n g the female d u r i n g a h a l f - h o u r o b s e r v a t i o n . The f o l l o w i n g s t a t i s t i c a l model was used i n an a n a l y s i s o f v a r i a n c e (PROC GLM; SAS 1985): Y,-jk = U + Gj + Pj + (GP),, + E,.jk where i = 1,2 and j = 1,2,3,4 Y..k = the mean amount of time a male of the i t h genotype l e f t h i s mate unattended d u r i n g a h a l f hour i n the j t h p e r i o d of her l a y i n g c y c l e . U = the p o p u l a t i o n mean; G, = the e f f e c t o f whether the male was f e r a l o r domestic; P, = the e f f e c t o f h i s mate's l a y i n g c y c l e ; ( G p ) , j = genotype by p e r i o d i n t e r a c t i o n ; and E.-k = the e r r o r term. RESULTS PAIR FORMATION IN JAPANESE QUAIL A p p l y i n g the c r i t e r i a I s e t t o determine p a i r bonding, I i d e n t i f i e d 39 cases of p a i r i n g ( e x c l u d i n g 3 " p a i r bonds" by FB2 a f t e r she became b l i n d ) between a male and a female Japanese q u a i l i n the two years of o b s e r v a t i o n s (Table 4.1). In 35 cases, the s i t u a t i o n c l e a r l y i d e n t i f i e d the b e g i n n i n g and end of the p a i r i n g . In f o u r cases t h e r e were doubts t h a t the s t a r t o f the p a i r bond was p r o p e r l y i d e n t i f i e d . TABLE 4 . 1 : P a i r f o r m a t i o n i n Japanese q u a i l k e p t i n the f l i g h t - p e n s . MALE FEMALE FP1 FY1 FG1 FBI 1987 FERAL FYB1* 6/5-7/281 FYP1* 6/19-7/20 FYG1 7/17-8/4 6/24-7/20 FPB1 6/20-8/9 DPI DY1 DG1 DB1 TT DOMES DYB1 5/20-6/5 6/4-6/17 6/27-7/7 6/16-6/29 (7/06-7/24) DYP1 5/22-6/5 (6/5-6/22) 7/6-7/17 DYG1 6/11-8/19 DPB1 (7/3-7/15) (6/8-7/5) 7/14-8/29 FP2 FY2 FG2 FB2 1988 FERAL FYB2 5/2-5/22 5/19-6/10 (6/2-7/3) 7/04-7/23 (7/18-8/17) FYP2 6/21-8/5 FYG2 (5/22-8/5) FPB2* * 5/2-7/6 DP2 DY2 DG2 DB2Z DOMES DYB2 5/3-5/15 5/31-6/16 (5/16-6/1) DYP2 (5/20-5/30) 5/16-5/22 5/8-5/17 6/6-6/21 (5/24-6/8) 7/27-8/17 DYG2 6/3-7/17 (5/13-6/2) 8/11-8/30 7/19-8/1 DPB2* * 5/9-6/5 1 Month/day 2 Three p a i r bonds a s s o c i a t e d w i t h DB2 excluded because of onset o f b l i n d n e s s (see Appendix 4 ) . ( ) Female p a i r e d t o two males s i m u l t a n e o u s l y . * Males removed from f l i g h t pen on J u l y 29, 1987. ** Males removed from f l i g h t pen on J u l y 8, 1988. Except f o r one male i n each year (FYG1 and FYB2), f e r a l males p a i r e d w i t h o n l y one female f o r the whole b r e e d i n g season (Table 4.1). In 1987, FYG1 p a i r e d w i t h FG1 from June 24 t o J u l y 20, and switched t o FP1, t a k i n g her away from FYP1, f o r the r e s t o f the season. In 1988, FYB2 switched r e p e a t e d l y between FY2 and FB2, although FY2 a l s o m aintained a s t a b l e p a i r bond w i t h FYG2 f o r most of the season. Except f o r v e r y b r i e f p e r i o d s w i t h o v e r l a p p i n g p a i r bonds w i t h two d i f f e r e n t females, these two f e r a l males p a i r e d w i t h one female a t any one time. F e r a l females a l s o p a i r e d w i t h o n l y one male f o r the season, w i t h the e x c e p t i o n o f FP1 i n 1987 and FY2 i n 1988. Other than FY2 who maintained simultaneous p a i r bonds with FYB2 and FYG2 from June 2 t o J u l y 3 and J u l y 18 t o Aug. 5 i n 1988 f e r a l females a l s o formed p a i r bonds w i t h o n l y one male at any one time. On the oth e r hand, domestic males switched mates throughout the season i n both y e a r s . Most males formed s e q u e n t i a l p a i r bonds w i t h t h r e e o f the f o u r females i n the pen. Except f o r b r i e f o v e r l a p p i n g p e r i o d s , domestic males a l s o p a i r e d w i t h o n l y one female a t any one time. Two domestic females i n 1987 (DPI and DG1) formed simultaneous p a i r bonds w i t h two males from J u l y 6 t o J u l y 15, and June 8 t o June 22, r e s p e c t i v e l y . Two females i n 1988 (DY2 and DB2) a l s o p a i r e d w i t h two males a t the same time d u r i n g the l a s t week o f May. Female (DB2) a l s o p a i r e d w i t h two males from May 24 t o June 1. P a i r bonds f o r both f e r a l and domestic q u a i l 52 were g e n e r a l l y formed p r i o r t o egg l a y i n g and t e r m i n a t e d d u r i n g egg l a y i n g o r e a r l y i n c u b a t i o n . In 1987, the f i r s t p a i r bond i n y e a r l i n g domestic q u a i l was formed on May 20 and the l a s t p a i r bond ended on August 29. P a i r i n g f o r the f e r a l q u a i l d i d not s t a r t u n t i l June 5 and the l a s t p a i r bond was d i s s o l v e d on August 9. In 1988, the f i r s t p a i r bond f o r the two y e a r - o l d q u a i l was formed on May 2 f o r f e r a l and May 3 f o r domestic. The l a s t p a i r bond between f e r a l q u a i l was t e r m i n a t e d on August 17 w h i l e the l a s t p a i r bond f o r domestic q u a i l ended on August 30. The p a i r bonds f o r f e r a l q u a i l were formed 22 days (mean) p r i o r t o egg l a y i n g compared t o a mean of f o u r days f o r the domestic q u a i l . The d i f f e r e n c e was s i g n i f i c a n t (t=2.88, df=28, P<0.01). The d u r a t i o n o f p a i r bonds (Table 4.2) i n f e r a l q u a i l (X=38 days) was a l s o s i g n i f i c a n t l y l o n g e r than p a i r bonds i n domestic q u a i l (X=19 days; t=4.17, df=40, p<.01). On f o u r o c c a s i o n s domestic males st a y e d c l o s e t o the female w h i l e she was i n c u b a t i n g . Such behaviour was not observed i n the f e r a l males. In two o f f i v e cases i n 1987 and two of s i x cases i n 1988 when a female was i n c u b a t i n g a c l u t c h , a domestic male was seen e i t h e r b e s i d e the i n c u b a t i n g female o r s i t t i n g on the n e st w i t h her d u r i n g the i n c u b a t i o n p e r i o d . At no time was the male seen i n c u b a t i n g a l o n e . In two of the f o u r cases, the male was the one t h a t was p a i r e d t o the i n c u b a t i n g female b e f o r e the s t a r t of i n c u b a t i o n . In the o t h e r two cases, the p a i r bond was formed d u r i n g the 53 TABLE 4.2: Mean d u r a t i o n s o f pair-bonds i n days YEAR N 1987* N 1988** N Tot a l * * FERAL 5 36.60 + 6.83 8 39.12 ± 7.51 13 X=38.15 + 5.14 DOMESTIC 12 2 0 . 5 8 + 3 . 3 2 14 1 8 . 2 8 + 2 . 4 5 26 X=19.35 + 1.99 N =number o f p a i r b o n d s . * p < 0.05 ** p < 0.01 54 i n c u b a t i o n s t a g e . N e i t h e r t h e f e r a l n o r t h e d o m e s t i c m a l e s p a r t i c i p a t e d i n b r o o d i n g t h e c h i c k s . WITHIN-PAIR AND EXTRA-PAIR ACTIVITIES: C o u r t s h i p D i s p l a y s C o u r t s h i p d i s p l a y s e x h i b i t e d by p a i r e d a n d u n p a i r e d m a l e s a r e shown i n t h e s e c o n d g r o u p o f d i s p l a y s i n T a b l e 4.3 ( s t r u t t i n g ST, l e a d i n g LD, c r o w i n g CW, t i d b i t t i n g TT, and n e s t ceremony N E ) . S i m i l a r f r e q u e n c i e s o f s t r u t t i n g ( a g g r e s s i v e c o u r t s h i p ) and n e s t ceremony were e x h i b i t e d b y b o t h f e r a l and d o m e s t i c m a l e s . L e a d i n g was p e r f o r m e d e x c l u s i v e l y b y p a i r e d f e r a l m a l e s and a l w a y s t o w a r d s t h e i r m ates, b u t was p e r f o r m e d by b o t h p a i r e d and u n p a i r e d d o m e s t i c m a l e s . F e r a l m a l e s c r o w e d w h i l e s e a r c h i n g f o r t h e i r mates a f t e r t h e y h a d b e e n s e p a r a t e d , however, c r o w i n g by d o m e s t i c m a l e s seemed t o be p e r f o r m e d i n a l e s s s p e c i f i c c o n t e x t ( s e e a l s o C h a p t e r 3) . F e r a l m a l e s p e r f o r m e d t i d b i t t i n g t o t h e i r mate more o f t e n t h a n d i d d o m e s t i c m a l e s . M a t i n g A c t i v i t i e s T h e r e was a s i g n i f i c a n t d i f f e r e n c e (P<0.01) between p a i r e d f e r a l m a l e s and p a i r e d d o m e s t i c m a l e s i n t h e d i s t r i b u t i o n o f e x t r a - p a i r v e r s u s w i t h i n - p a i r c o p u l a t i o n s . P a i r e d d o m e s t i c m a l e s d i d p r o p o r t i o n a l l y more e x t r a - p a i r c o p u l a t i o n s t h a n p a i r e d f e r a l m a l e s ( T a b l e 4.4) d u r i n g t h e i r m a t e s ' n o n - l a y i n g and l a y i n g p e r i o d s . P a i r e d d o m e s t i c m a l e s s e a r c h e d more f r e q u e n t l y f o r f e m a l e s o t h e r t h a n t h e i r mates 55 TABLE 4 . 3 : R e l a t i v e f r e q u e n c y o f c o u r t s h i p d i s p l a y s and mat ing b e h a v i o u r per formed by a p a i r e d male t o h i s mate ( I P ) , by the p a i r e d male t o a n o t h e r female no t h i s mate ( E P ) , and by a n o n - p a i r e d male t o a female (OP) . DISPLAY 1 FERAL MALES DOMESTIC MALES "1? IP EP OP z N IP EP OP" GROUP IS BL 243 89% 11% 0% 185 85% 3% 12 SE 139 94 4 2 91 62 20 18 RT 197 94 4 2 122 80 5 15 GROUP 2: LD 12 100 0 0 6 67 0 33 NE 1 100 0 0 5 100 0 0 TT 30 90 0 10 7 57 14 29 CW 648 89 6 5 453 58 21 21 ST 176 68 14 18 154 62 8 30 GROUP 3: BN 394 55 32 13 604 32 40 28 MO 321 59 31 10 553 34 36 30 TB 114 68 31 1 253 38 30 32 CC 63 70 29 1 125 42 33 25 The f i r s t group of d i s p l a y s was p a r t o f the c r i t e r i a f o r dete r m i n i n g p a i r - b o n d d u r a t i o n , the second group of d i s p l a y s was not used as p a r t o f c r i t e r i a t o determine p a i r -bonds, and the t h i r d group c o n s i s t s o f mating behaviour components. See Chapter 3 f o r a b r e v i a t i o n s . There were 404 male-days of o b s e r v a t i o n s over the two year p e r i o d , f e r a l males were un p a i r e d f o r 129 male-days and domestic males were un p a i r e d f o r 56 male-days. T o t a l number of d i s p l a y s by a l l males over the two y e a r s . TABLE 4 . 4 : The d i s t r i b u t i o n o f a m a l e ' s m a t i n g a c t i v i t i e s ( w i t h i n - p a i r v s e x t r a - p a i r ) a t the t h r e e s tages o f the l a y i n g c y c l e o f h i s own mate . MALE FERAL DOMESTIC A C T I V I T I E S N* IP EP IP EP FO (NL) 17 71% 29% 45 20% 80%** (PL) 4 75 25 24 42 58 ( L) 12 100 0 33 36 e A** 64 BN (NL) 37 76 24 78 23 77 (PL) 14 57 43 40 38 62 ( L) 43 70 30 115 48 52** MO (NL) 36 75 25 71 21 79** (PL) 10 80 20 32 41 59 ( L) 39 69 31 104 54 46 TB (NL) 15 67 33 36 22 _ _ *» 78 (PL) 5 80 20 15 67 33 ( L) 24 83 17 48 50 50 CC (NL) 9 67 33 18 44 56 (PL) 4 100 0 6 83 17 ( L) 15 93 7 26 46 C A ** 54 1 IP = w i t h i n - p a i r , EP = e x t r a - p a i r 2 NL = Non-laying p e r i o d (8-14 days p r i o r t o l a y i n g ) , PL = P r e - l a y i n g p r i o d ( w i t h i n 7 days b e f o r e l a y i n g ) , and L = L a y i n g p e r i o d . 3 t o t a l number of d i s p l a y s over two y e a r s . P<0.05, s i g n i f i c a n t d i f f e r e n c e between f e r a l and domestic males i n the perc e n t o f e x t r a - p a i r a c t i v i t i e s d u r i n g t h a t p e r i o d . 57 (Table 4.3), presumably seeking e x t r a - p a i r c o p u l a t i o n s , but p a i r e d f e r a l males searched mainly f o r t h e i r mates. P a i r e d f e r a l males had a lower success r a t e f o r e x t r a - p a i r c o p u l a t i o n s than w i t h i n - p a i r c o p u l a t i o n s d u r i n g t h e i r mate's p r e - l a y i n g and l a y i n g p e r i o d (Table 4.5). There was no d i f f e r e n c e i n success r a t e between w i t h i n - p a i r and e x t r a - p a i r c o p u l a t i o n s by p a i r e d domestic males. C o n s i d e r i n g t h a t domesticated males were p a i r e d f o r 348/404 male-days and the f e r a l males o n l y 275/404 male-days (see f o o t n o t e 2 of Table 4.3), i t i s i n t e r e s t i n g t h a t 25% of the c o p u l a t i o n s of domestic males were done w h i l e unpaired, compared t o o n l y 1% by nonpaired f e r a l males. E x t r a - p a i r c o p u l a t i o n attempts by males o f t e n r e s u l t e d i n the d i s r u p t i o n t o the females' l a y i n g and n e s t i n g a c t i v i t i e s . B e h a v i o u r a l o b s e r v a t i o n s showed t h a t f e r a l females were v e r y aware o f approaching males w i t h the i n t e n t o f f o r c e d e x t r a -p a i r c o p u l a t i o n s . They o f t e n dashed t o g r a s s cover t o a v o i d f u r t h e r d e t e c t i o n by the males and sometimes h i d and d i d not respond t o the crowing and s e a r c h i n g of t h e i r mate. Such behaviour was seldom observed i n domestic females. During i n c u b a t i o n , 57% (n=37) of the time a domestic female spent o f f her n e s t was because o f d i s t u r b a n c e d u r i n g c o p u l a t i o n attempts by her mate o r by o t h e r males. (The r e s t o f the time she l e f t her n e s t t o feed and d r i n k . ) The c o r r e s p o n d i n g f i g u r e f o r f e r a l females (20%; n=15) was s i g n i f i c a n t l y l e s s (X2=5.75, d f = l , p<0.05). 58 TABLE 4 .5: Success r a t e 1 o f w i t h i n - p a i r and e x t r a - p a i r c o p u l a t i o n s by f e r a l and domes t i c males d u r i n g the t h r e e s tages o f t h e i r mates* l a y i n g c y c l e . H A T E ' S FERAL MALES DOMESTIC MALES CYCLE N 2 I P 3 EP N IP EP NON-LAYING 4 7 11+ 7.6bc 7+ 7.3c 7 43+20.5ab 16+ 7.5abc PRE-LAYING 6 39±15.6ab 0+ 0.0c 8 21+ 8.2abc 6+ 6.3c LAYING 9 49+12.2a 2+ 1.6c 11 16+ 5.8abc 18+ 9.4abc 1 Success r a t e (¥) i s c a l c u l a t e d by the frequency o f c l o a c a l c o n t a c t s d i v i d e d by the frequency of b i l l - o n - n a p e s f o r t h a t p e r i o d m u l t i p l i e d by 100. 2 N = Number of p a i r s sampled i n each p e r i o d over two y e a r s . 3 IP = w i t h i n - p a i r , EP = e x t r a - p a i r 4 Non-laying p e r i o d = 8-14 days p r i o r t o l a y i n g , P r e - l a y i n g p r i o d = w i t h i n 7 days b e f o r e l a y i n g . Means t h a t are f o l l o w e d by the same l e t t e r a re not s i g n i f i c a n t l y d i f f e r e n t . Means t h a t are f o l l o w e d by d i f f e r e n t l e t t e r s are s i g n i f i c a n t l y d i f f e r e n t (P<0.05) by Duncan's M u l t i p l e Range T e s t . 59 Domestic females appeared t o be l e s s s e l e c t i v e i n t h e i r s e x u a l crouch. Of the f e r a l female's crouches, o n l y 18% (n=34) were not d i r e c t e d towards a mate, s i g n i f i c a n t l y l e s s than domestic females (45%; n=51, X2=6.83, d f = l , p<0.05). Mate Guarding A p a i r e d f e r a l male o f t e n p o s i t i o n e d h i m s e l f between h i s mate and an i n t r u d i n g male t r y i n g t o prevent the i n t r u d i n g male from g e t t i n g t o h i s female (BL) . When the i n t r u d i n g male p e r s i s t e d a f i g h t ensued, w i t h the def e n d i n g male u s u a l l y winning and c h a s i n g the i n t r u d i n g male away. In t h i s way p a i r e d males defended t h e i r mates and, i n some cases, o t h e r females (Table 4.3). T h i s behaviour, however, was a l s o observed i n unp a i r e d domestic males i n defence o f females they were i n t e r e s t e d i n . A p a i r e d male sometimes l e f t h i s mate unattended when the male was c h a s i n g an i n t r u d i n g m a l e ( s ) , was attempting e x t r a -p a i r c o p u l a t i o n , o r was f e e d i n g , d r i n k i n g o r d u s t - b a t h i n g . P a i r e d males r e t u r n e d t o t h e i r mates a f t e r these a c t i v i t i e s and i f the female was not i n s i g h t , o f t e n searched f o r her u n t i l f i n d i n g ' h e r and r e t u r n i n g t o her s i d e . On a per f o r a y b a s i s , the mean l a t e n c y o f a male r e t u r n i n g t o h i s mate a f t e r c h a s i n g o f f an i n t r u d e r was 97.54+7.00s (n=210). The mean l a t e n c y a f t e r f e e d i n g and d r i n k i n g a c t i v i t i e s was a l s o s h o r t (137.43+18.32s, n=94) and not d i f f e r e n t , but p a i r e d males were away from mates s i g n i f i c a n t l y (P<0.01) l o n g e r when they attempted e x t r a - p a i r c o p u l a t i o n s (190. 64+23 .14s, n=125) . T h i s was t r u e f o r both f e r a l and domestic males (no s i g n i f i c a n t genotype by a c t i v i t y i n t e r a c t i o n ) , but domestic males had s i g n i f i c a n t l y (P<0.01) l o n g e r l a t e n c i e s (193.82+18.90s, n=159) t o r e t u r n t o t h e i r females than f e r a l males (97.83+7.64s, n=270). F e r a l males r e t u r n e d t o t h e i r mate q u i c k l y through a l l f o u r p e r i o d s o f her l a y i n g c y c l e ( F i g . 4.1). Domestic males r e t u r n e d t o t h e i r mate as q u i c k l y as the f e r a l males d u r i n g the p r e - l a y i n g and l a t t e r p a r t o f the l a y i n g p e r i o d , but they took much lo n g e r t o r e t u r n d u r i n g the n o n - l a y i n g and e a r l y l a y i n g p e r i o d s of her l a y i n g c y c l e . When c o n s i d e r i n g both the l a t e n c y t o r e t u r n and the frequency o f these f o r a y s by the male d u r i n g a h a l f - h o u r o b s e r v a t i o n , f e r a l males l e f t t h e i r mate unattended f o r a mean of 178.70+15.48s per o b s e r v a t i o n (n=178) , s i g n i f i c a n t l y s h o r t e r than domestic males (299.39+48.03s, n=31). There were, however, no s i g n i f i c a n t d i f f e r e n c e s among the f o u r p e r i o d s of the l a y i n g c y c l e . DISCUSSION G e n e r a l l y , f e r a l Japanese q u a i l demonstrated "monogamy" (Wittenberger 1979) w i t h some o p p o r t u n i s t i c f o r c e d e x t r a - p a i r c o p u l a t i o n s (FEPC) (McKinney e t a l . 1983, see a l s o Mock & F u j i o k a 1990). While one female (FY2) seemingly maintained Figure 4.1: Average length of time (seconds) for a male to return to his mate after leaving her side during four stages of her laying cycle (latency). Latency Domestic J H ^ 1 N u m b e r o f f o r a y s over t w o y e a r s . M e a n s w i t h t h e s a m e l e t t e r a r e n o t s i g n i f i c a n t l y d i f f e r e n t M e a n s w i t h d i f f e r e n t l e t t e r s a re s i g n i f i c a n t l y ( P < 0 . 0 5 ) d i f f e r e n t by D u n c a n ' s M u l t i p l e R a n g e Tes t . Laying Cycle 62 p a i r bonds w i t h two males s i m u l t a n e o u s l y , t h i s c o u l d have been an a r t i f a c t o f my d e f i n i t i o n of the p a i r bond, which was mostly based on male behaviour. In waterfowl s p e c i e s , acceptance o f the male by the female p l a y s an important r o l e i n p r e s e r v i n g the bond (McKinney 1985). In Japanese q u a i l , an i n d i c a t i o n o f female acceptance of a male c o u l d be s o l i c i t a t i o n of c o p u l a t i o n by the female. A female would walk i n f r o n t of her mate and crouch, i n v i t i n g the male t o mount her. Mates a l s o keep i n c l o s e v o c a l c o n t a c t w i t h each other, g i v i n g v e r y q u i e t c a l l s as they move around t o g e t h e r . These are, however, qu i c k s u b t l e events t h a t can be d i f f i c u l t f o r an o bserver t o see or hear. I b e l i e v e t h a t the female p l a y s a r o l e i n the f o r m a t i o n and the maintenance of the bond by choosing and s t a y i n g c l o s e t o her mate. Japanese q u a i l females are b i g g e r than males and can r e s i s t c o p u l a t i o n attempts by males e f f e c t i v e l y . With the h e l p of her mate, a p a i r e d female should not be an easy t a r g e t f o r a male attempting f o r c e d c o p u l a t i o n u n l e s s more than one male i s advancing on the female a t the same time. Given t h a t t h e i r n a t u r a l d e n s i t y i s about a p a i r per h e c t a r e (Schwartz & Schwartz 1949, Robinson e t a l . i n press) and the f a c t t h a t these b i r d s are mostly a c t i v e on the ground and seldom f l y over l o n g d i s t a n c e s , the s i t u a t i o n where m u l t i p l e males descend on a female may be r a r e . Even i n the f l i g h t - p e n s where b i r d d e n s i t i e s were hi g h , the success r a t e o f FEPC's by f e r a l males was s t i l l low. 63 P a i r e d males c o n t r i b u t e t o t h e i r mate's b r e e d i n g e f f o r t by p r o t e c t i n g her from d i s t u r b a n c e by o t h e r males and a l e r t i n g her t o the presence o f p r e d a t o r s (Wittenberger and T i l s o n 1980). Such v i g i l a n c e a l l o w s the female t o f e e d more e f f i c i e n t l y and a l s o minimizes the chance t h a t she w i l l l o s e her mate t o another male or be inseminated by o t h e r males (McKinney 1985). Under t y p i c a l domestic r e a r i n g c o n d i t i o n s , feed i s p r o v i d e d , p r e d a t o r s are kept away, n e s t i n g i s p r o h i b i t e d , and h i g h d e n s i t y r e a r i n g makes mate-guarding i n e f f e c t i v e . No b e n e f i t s of a prolonged p a i r bond, e s p e c i a l l y a monogamous one, e x i s t and the behaviour i s a c t u a l l y d i s c o u r a g e d . Only i n a l t r i c i a l s p e c i e s , such as doves, f i n c h e s , and c a n a r i e s , where the male has t o h e l p r e a r the young, i s the p a i r bond maintained under d o m e s t i c a t i o n . Reports of breakdown of the p a i r bond and q u a n t i t a t i v e changes i n c o u r t s h i p d i s p l a y s and i n s e x u a l behaviour are common f o r domestic p r e c o c i a l s p e c i e s o f b i r d s (Sossinka 1982). However, documentation o f the q u a l i t a t i v e and q u a n t i t a t i v e changes of the p a i r bond of domestic b i r d s under a more n a t u r a l environment has not been done u n t i l t h i s study. Compared t o f e r a l males under s i m i l a r environmental c o n d i t i o n s , domestic males showed l e s s c o u r t s h i p , had s h o r t e r p a i r bonds, attempted more FEPC's, and were o n l y c l o s e l y a s s o c i a t e d w i t h the female d u r i n g the p e r i o d when she was f e r t i l i z a b l e ( p r e - l a y i n g and l a y i n g ) . Under domestic c o n d i t i o n s , the premium i s t o f e r t i l i z e as many eggs as 64 p o s s i b l e (Haase and Donham 1980) and the r e s t i s i n c o n s e q u e n t i a l . Stevens (1961) r e p o r t e d t h a t domestic male Japanese q u a i l may share i n c u b a t i o n w i t h the female. Under the f l i g h t - p e n c o n d i t i o n s I confirmed t h a t domestic males s a t on eggs a l o n g s i d e females. However, these males were not " s h a r i n g i n c u b a t i o n " , as they never s a t on the eggs alon e . I am a l s o not convinced t h a t they were mate guarding, as i n two o f the f o u r cases, p a i r bonds o n l y formed a f t e r i n c u b a t i o n had begun. I t i s more l i k e l y t h a t the accompanying male was p r i m a r i l y i n t e r e s t e d i n mating w i t h the female ( S t a n f o r d 1957, McKinney 1985). One f a c t o r promoting monogamy i s the i n a b i l i t y o f any male t o monopolize more than one female under a synchronous b r e e d i n g c o n d i t i o n (Wittenberger & T i l s o n 1980). McKinney (1985) p r e d i c t s t h a t a l t e r n a t i v e mating s t r a t e g i e s such as polygyny may evolve i n some s p e c i e s because an extended b r e e d i n g season can g i v e r i s e t o m u l t i p l e b rooding and asynchronous b r e e d i n g . T h i s h y p o t h e s i s has not been t e s t e d e x p e r i m e n t a l l y . Under the f l i g h t - p e n s i t u a t i o n , domestic females brought o f f m u l t i p l e broods (see Chapter 5) and were a l s o l e s s s e l e c t i v e than f e r a l females i n t h e i r s e x u a l crouch. I t i s not c l e a r how important a r o l e t h i s p l a y s i n the shortened p a i r i n g , more frequent mate s w i t c h i n g , and i n c r e a s e d FEPC observed i n the domestic males. I t w i l l be v e r y i n f o r m a t i v e t o p l a c e f e r a l males and domestic females, and the 65 r e c i p r o c a l p a i r i n g s i n the f l i g h t - p e n s , o r a l t e r the sex r a t i o , and study t h e i r mating behaviour. In summary, both f e r a l and domestic Japanese q u a i l had a monogamous mating system, however domestic q u a i l had s h o r t e r p a i r bonds and switched mates f r e q u e n t l y d u r i n g a b r e e d i n g season. F e r a l q u a i l l e f t t h e i r mates unattended f o r s h o r t e r p e r i o d s and performed fewer e x t r a - p a i r c o p u l a t i o n s . 66 CHAPTER F I V E REPRODUCTION OF FERAL AND DOMESTIC JAPANESE QUAIL INTRODUCTION A v i a n r e p r o d u c t i o n i s p e r i o d i c . When the a p p r o p r i a t e c o n d i t i o n s a re met, a b i r d throws much o f i t s p h y s i o l o g i c a l r e s o u r c e s i n t o a c o n c e n t r a t e d e f f o r t t o reproduce. The r e p r o d u c t i v e c y c l e i s r e g u l a t e d by the neuro-endocrine system which i s s e n s i t i v e t o a v a r i e t y o f environmental s t i m u l i ; f o r example, day l e n g t h (Blank and Ash 1960; Howes 1964; Klomp 1970), p r e c i p i t a t i o n ( F o s t e r 1974; Yom-Tov and H i l b o r n 1981), temperature (Arad e t a l . 1981), and food a v a i l a b i l i t y (Lack 1968; R a v e l i n g 1979; R i c k l e f s 1980). Thus a v i a n r e p r o d u c t i o n u s u a l l y occurs when environmental c o n d i t i o n s p r o v i d e enough n u t r i e n t s f o r the female t o produce eggs and r a i s e young (Klomp 1970; P e r r i n s 1970; Daan e t a l . 1988). By m a n i p u l a t i n g environmental c o n d i t i o n s , mankind has succeeded i n extending the r e p r o d u c t i v e c y c l e o f domestic s p e c i e s (Wilson e t a l . 1961). W i l d Japanese q u a i l l a y seven t o 14 eggs per c l u t c h and they a p p a r e n t l y l a y o n l y one c l u t c h i n a b r e e d i n g season i f the eggs are s u c c e s s f u l l y hatched (Demente'ev 1967). T h e i r domestic c o n s p e c i f i c s can produce 250 - 3 65 eggs per year under the a p p r o p r i a t e manangement scheme ( A u s t i n and Kuroda 1953; McGrew 1958, Kawahara 1971; Abplanalp e t a l . 1963) and males can s t a y f e r t i l e y e a r round. T h i s i n c r e a s e i n r e p r o d u c t i v e c a p a b i l i t y i n domestic animals 67 may i n v o l v e h e r i t a b l e changes i n t h e i r p h y s i o l o g y , anatomy, and behaviour (Hale 1969; Kawahara 1972; Boice 1972, 1973). C l a y t o n (1972), however, argued t h a t t h e r e have not been d r a s t i c changes i n the g e n e t i c c o n t r o l over r e p r o d u c t i o n i n domestic animals. Rather domestic animals express t h e i r n a t u r a l f e c u n d i t y more s t r o n g l y when i n h i b i t i n g s t i m u l i are removed and enhancing s t i m u l i are p r o v i d e d . A c c o r d i n g t o t h i s argument, when a domestic animal i s r e t u r n e d t o a n a t u r a l s e t t i n g , i t should e x h i b i t a r e p r o d u c t i v e p a t t e r n s i m i l a r t o i t s w i l d c o n s p e c i f i c . The two hypotheses are not e x c l u s i v e but d i f f e r i n the degree of expected g e n e t i c changes. P h y s i o l o g i c a l comparisons (Kawahara 1971, 1972; Kawahara and M i t a 1968) and r e l e a s e and r e s t o c k i n g p r o j e c t s (Backs 1982; Krauss e t a l . 1987) have f a i l e d t o d i s t i n g u i s h the two hypotheses. While Roseberry e t a l . (1987) p r o v i d e d some q u a l i t a t i v e b e h a v i o u r a l i n d i c a t i o n s and q u a n t i t a t i v e post r e l e a s e data t o suggest t h a t r e a r i n g c o n d i t i o n s may be more important than g e n e t i c s , a c o n t r o l l e d experiment t h a t examines these hypotheses i s s t i l l needed. In f o r m a t i o n r e g a r d i n g r e p r o d u c t i o n of w i l d Japanese q u a i l i s s c a r c e (Wakasugi 1984), and the few s t u d i e s of domestic Japanese q u a i l i n a v i a r i e s , a more n a t u r a l s e t t i n g than crowded cages, have produced c o n f l i c t i n g c o n c l u s i o n s . Eynon (1968), R o t h s t e i n (1967), and F a r r i s (1964) concluded t h a t females would l a y eggs s c a t t e r e d throughout the a v i a r y without any i n t e n t i o n t o incubate the eggs. Others ( S t a n f o r d 1957; 68 S t e v e n s 1961; and O r c u t t and O r c u t t 1976) r e p o r t e d t h a t d o m e s t i c f e m a l e s w o u l d n e s t i f " c o n d i t i o n s a r e f a v o u r a b l e " , b u t t h e y f a i l e d t o p r o v i d e i n f o r m a t i o n a b o u t what t h e a p p r o p r i a t e e n v i r o n m e n t a l s t i m u l i f o r n e s t i n g a n d i n c u b a t i o n i n J a p a n e s e q u a i l a r e . I n t h i s c h a p t e r I i n v e s t i g a t e d i f f e r e n c e s i n r e p r o d u c t i o n b e tween f e r a l and d o m e s t i c J a p a n e s e q u a i l k e p t i n t h e same s e m i - n a t u r a l e n v i r o n m e n t . T h e s e d a t a p r o v i d e some i n s i g h t s i n t o w h i c h a s p e c t s o f r e p r o d u c t i o n h a v e b e e n c h a n g e d by d o m e s t i c a t i o n . C o m p a r i s o n u n d e r s i m i l a r e n v i r o n m e n t a l c o n d i t i o n s s h o u l d r e v e a l g e n e t i c d i f f e r e n c e s ( S o s s i n k a 1982) and how t h e s e a s p e c t s may be r e l a t e d t o one a n o t h e r . I compare t h e f o l l o w i n g : 1. o n s e t o f egg l a y i n g 2. c l u t c h s i z e a. number o f e g g s i n a n e s t b. number o f e g g s a f e m a l e l a i d i n c o n s e c u t i v e d a y s 3. number o f c l u t c h e s p e r s e a s o n 4. t i m e and f r e q u e n c y o f r e n e s t i n g 5. number o f e g g s l a i d p a r a s i t i c a l l y 6. f e r t i l i t y a nd h a t c h a b i l i t y o f e g g s 7. m o r t a l i t y o f c h i c k s , and 8. d u r a t i o n o f p a r e n t - y o u n g a s s o c i a t i o n s 69 METHODS Data were c o l l e c t e d i n the l a r g e f l i g h t - p e n s d u r i n g the two b r e e d i n g seasons, except f o r egg p r o d u c t i o n data t h a t i n c l u d e data taken from the s m a l l pens (see Chapter 2 -General Methods). CLUTCH DATA B e h a v i o u r a l o b s e r v a t i o n s were used t o i d e n t i f y c l u t c h e s l a i d by i n d i v i d u a l females. S h e l l c o l o u r p a t t e r n , egg s i z e , and shape were a l s o used t o i d e n t i f y m a t e r n i t y o f the eggs w i t h i n a n e s t (Jones e t . a l . 1964). No attempt was made t o i d e n t i f y odd eggs found i n v a r i o u s l o c a t i o n s o u t s i d e the ne s t s s i n c e these eggs were o f t e n l a i d i n exposed areas where t h e i r c o l o u r p a t t e r n would q u i c k l y be bleached by the sun. Nest checks were c a r r i e d out tw i c e a week w h i l e the fee d e r s and water jugs were being s e r v i c e d . For each nest examined, the f o l l o w i n g i n f o r m a t i o n was reco r d e d or e x t r a p o l a t e d : the l o c a t i o n ; the i d e n t i t y o f the female(s) l a y i n g and i n c u b a t i n g eggs; the d u r a t i o n o f i n c u b a t i o n ; and the date o f h a t c h i n g (or the date o f nest d e s e r t i o n ) . Because females l a i d i n d i v i d u a l l y i d e n t i f i a b l e eggs, the date when each female l a i d was known even i f she l a i d i n d i f f e r e n t n e s t s . 7 0 FERTILITY AND HATCHABILITY F e r t i l i t y i s the p e r c e n t o f the t o t a l number o f eggs incubated (by the female) t h a t were f e r t i l e , and h a t c h a b i l i t y i s t he p e r c e n t of f e r t i l e eggs t h a t hatched. Data on h a t c h a b i l i t y were c o l l e c t e d by c o u n t i n g the number o f c h i c k s hatched. Unhatched eggs were removed and examined f o r embryonic development. Deserted n e s t s were monitored f o r two weeks and i f the female d i d not r e t u r n t o the n e s t s , the eggs were removed and r e p l a c e d w i t h "dummy" eggs t o minimize a l t e r a t i o n o f the n e s t i n g environment i n the f l i g h t - p e n s . I f i n c u b a t i o n had s t a r t e d p r i o r t o d e s e r t i o n , the eggs were checked f o r f e r t i l i t y . I f o l l o w e d Rosin's (1944) method of examining the germinal d i s c m a c r o s c o p i c a l l y t o determine f e r t i l i t y . CHICK MORTALITY AND PARENT-YOUNG ASSOCIATION Once the c h i c k s hatched, I rec o r d e d the number of c h i c k s f o l l o w i n g the maternal or f o s t e r female d u r i n g the r e g u l a r o b s e r v a t i o n p e r i o d s . C h icks t h a t d i s a p p e a r e d were assumed t o be dead. D e s p i t e frequent mixing of broods and c h i c k swapping among the domestic females, t h e r e was always s u f f i c i e n t age d i f f e r e n c e among broods f o r proper i d e n t i f i c a t i o n and t o t a l c h i c k counts. The time when c h i c k s showed l i t t l e attachment t o the female and were moving about the pen independently of the female and/or broodmates was noted. A l l c h i c k s were then removed from the pen when they were t h r e e t o f o u r weeks o l d 71 (or a f t e r they showed no attachment t o t h e i r parent, whichever time came l a t e r ) t o minimize b i r d d e n s i t y i n the pens. DATA ANALYSIS Data were analyzed by Chi-squares o r Student's t - t e s t (SAS 1985) and were l o g transformed (Y = Ln X) f o r t h e t - t e s t s (Snedecor and Cochran 1980). RESULTS AGE OF SEXUAL MATURITY AND INCIDENCE OF NON-BREEDING A l l f o u r domestic females i n each o f the two years ( i . e . both age groups) l a i d a t l e a s t one c l u t c h o f eggs. While each o f the f o u r f e r a l females (2 year o l d s ) l a i d a t l e a s t one c l u t c h i n 1988, o n l y one of the f o u r y e a r l i n g f e r a l females l a i d i n 1987 (Table 5.1). CLUTCH SIZE I . C l u t c h e s Incubated (Table 5 . 1 ) : In t he two y e a r s , females i n both pens s t a r t e d i n c u b a t i o n on 19 n e s t s ; f o u r o f these were by f e r a l females and 15 by domestic females. Of the f o u r n e s t s o f f e r a l females, t h r e e (75%) were incubated t o h a t c h i n g . These n e s t s c o n t a i n e d 10, I I , and 11 eggs (X=10.67+0.33) . E l e v e n o f the 15 n e s t s (73%) 72 TABLE 5 . 1 : D e s c r i p t i o n Of Incubated N e s t s 1 Found i n the Two Years o f the S t u d y . YEAR FERAL NESTS DOMESTIC NESTS FP1 10(10) [34] DPI 13(11),13(12),6(6) [29] FY1 [ 0] DY1 20(A),7(7) [31] FG1 [ 0] DG1 10(8) [21] FBI [ 0] DB1 [12] UNIDENTIFIED [ 2] UNIDENTIFIED [21] FY2 8(A) [14] DP2 9(A),8(7) [28] FG2 11(11) [11] DY2 16(A).8(7).10(7).6(3) [35] FB2 11(3) [ 5] DG2 14(6),7(A),8(1) [34] FP2 [ 4] DB2 [23] UNIDENTIFIED [29] UNIDENTIFIED [37] sequence. Number i n d i c a t e s number of eggs found i n t h e n e s t . Number i n () i n d i c a t e s number of c h i c k s hatched from t h a t n e s t . Nests d e s e r t e d b e f o r e h a t c h i n g are i n d i c a t e d by "A" i n the ( ) . Number u n d e r l i n e d i n d i c a t e s eggs from more than one female were found i n the nest . Number i n [] i n d i c a t e s t o t a l number of eggs l a i d by t h a t female from May t o September. These i n c l u d e her own eggs i n the incubated n e s t s , eggs l a i d i n n e s t s d e s e r t e d b e f o r e s t a r t o f i n c u b a t i o n , and/or eggs l a i d i n o t h e r females' n e s t s . of the domestic females were incubated t o h a t c h i n g . The mean number o f eggs found i n these n e s t s was s i m i l a r but the standard e r r o r was much l a r g e r (X=10.33+1.05). The nest d e s e r t e d by a f e r a l female (FY2) had e i g h t eggs, w h i l e the f o u r n e s t s d e s e r t e d by domestic females had seven, n i n e , 16, and 20 eggs r e s p e c t i v e l y . The mean number o f eggs i n the f i v e n e s t s t h a t had eggs from more than one female was 12.20+1.43 (see s e c t i o n on nest p a r a s i t i s m below). 2. Eggs l a i d (Table 5 . 2 ) : The mean c l u t c h s i z e of f e r a l females i n a l l n e s t s f o r the 2 y e a r p e r i o d was 7.55+0.87 (n=9), and t h a t of domestic females was 8.52+0.77 (n=25) and was not s i g n i f i c a n t l y d i f f e r e n t (t=-0.40, df=32, P=0.69) from t h a t o f the f e r a l females. For domestic females, the mean c l u t c h s i z e of y e a r l i n g s i n 1987, 11.62+1.43 (n=8), was s i g n i f i c a n t l y l a r g e r (t=2.90, df=23, P=0.01) than t h a t f o r the 2 y e a r - o l d s i n 1988 (7.06+0.68) (n=17). However, the mean c l u t c h s i z e i s b i a s e d downward as a l a r g e number of odd eggs were found o u t s i d e n e s t s , and these were not i d e n t i f i a b l e by female. T h i s was p a r t i c u l a r l y t r u e f o r the domestic females i n the second year. I compare f e r a l females a c r o s s years s i n c e o n l y one female l a i d i n the f i r s t y e ar. For the domestic females t h a t l a i d m u l t i p l e c l u t c h e s i n a season, the c l u t c h s i z e d i d not change through the season. Although the l a s t c l u t c h e s (n=7, X=7.00+0.98) were s l i g h t l y s m a l l e r , they were not 74 TABLE 5 . 2 : C l u t c h S i z e 1 o f Females i n the F l i g h t - p e n . YEAR FERAL FEMALES DOMESTIC FEMALES 1987 FP1 5,10,9,10 [10] DPI 13,10,6 [27] FY1 0 [ 0] DY1 20,11 [ 9] FG1 0 C 0] DG1 12,9 [ 8] FBI 0 t 0] DB1 12 [ 0] UNIDENTIFIED (2) UNIDENTIFIED (21) 1988 FY2 6,8 [ 1] DP2 5,13,2,8 t 7] FG2 11 [11] DY2 11,8,10,6 [17] FB2 5 [ 2] DG2 7,9,7,5,6 [ 7] FP2 4 [ 0] DB2 5,6,9,3 C 0] UNIDENTIFIED (29) UNIDENTIFIED (37) 1 Number of eggs i n c l u d e d i n a c l u t c h were those l a i d by a female: 1. i n her own nest 2. elsewhere w h i l e l a y i n g i n her nest 3. elsewhere when her own n e s t was complete so l o n g as no more than one day was skipped. C l u t c h e s from the same female are l i s t e d i n temporal sequence. Number i n [] i n d i c a t e s number of her eggs t h a t hatched. Number i n () i n d i c a t e s t o t a l number of u n i d e n t i f i a b l e eggs found i n t h a t pen. s i g n i f i c a n t l y d i f f e r e n t (t=1.00, df=23, P=0.33) from the ot h e r s (n=18, X=9.11+0.98). TIME AND FREQUENCY OF RENESTING I n c l u d i n g n e s t s t h a t were d e s e r t e d b e f o r e the i n i t i a t i o n o f i n c u b a t i o n , f i v e o f e i g h t n e s t i n g attempts by f e r a l females f a i l e d . In the f i r s t year, when o n l y one female was l a y i n g , the female (FP1) d e s e r t e d t h r e e n e s t s b e f o r e she s t a r t e d i n c u b a t i o n . F o l l o w i n g each f a i l e d attempt, she s t a r t e d l a y i n g i n a new ne s t w i t h i n t h r e e days o f nest d e s e r t i o n . The f o u r t h n e s t was incubated t o h a t c h i n g . In the second year, one nest (FP2) was d e s e r t e d b e f o r e i n c u b a t i o n s t a r t e d and another nest (FY2) was incubated f o r 3 days by the female b e f o r e she d e s e r t e d . There were no r e n e s t i n g attempts a f t e r these f a i l u r e s . During both b r e e d i n g seasons, f e r a l females o n l y incubated once and d i d not r e n e s t a f t e r a c l u t c h was s u c c e s s f u l l y hatched (n=3). The domestic females f a i l e d i n 10 of 21 n e s t i n g attempts. In the f i r s t year, o n l y one female (DPI) i n c u b a t e d her f i r s t c l u t c h t o h a t c h i n g . Most were s u c c e s s f u l i n h a t c h i n g c h i c k s i n t h e i r second attempt (see Table 5.1). In the second year, a l l f o u r females d e s e r t e d t h e i r f i r s t c l u t c h . Most were s u c c e s s f u l i n subsequent attempts. In the f o u r cases, over the 2 y e a r s , where a c l u t c h was d e s e r t e d p r i o r t o i n c u b a t i o n , the female s t a r t e d l a y i n g a new c l u t c h i n a mean of 6 days. In the f o u r i n s t a n c e s where d e s e r t i o n o c c u r r e d d u r i n g i n c u b a t i o n , r e n e s t i n g began i n a 76 mean o f 9 days. FERTILITY, HATCHABILITY, AND INCUBATION OF EGGS In c u b a t i o n p e r i o d s ranged from 16 t o 17 days f o r the f e r a l n e s t s and 17 t o 19 days f o r the domestic n e s t s . Data f o r h a t c h a b i l i t y and f e r t i l i t y were based on n e s t s where a t l e a s t one egg hatched. While eggs l e f t i n d e s e r t e d n e s t s were examined f o r f e r t i l i t y , I found i t d i f f i c u l t t o assess f e r t i l i t y a c c u r a t e l y because of the s h o r t i n c u b a t i o n p e r i o d p l u s the d e t e r i o r a t i o n o f the egg content by the time the eggs were examined (2 weeks a f t e r d e s e r t i o n ) . F e r t i l i t y of eggs from both f e r a l and domestic females was over 90% and d i d not d i f f e r between y e a r s . However, h a t c h a b i l i t y was s i g n i f i c a n t l y lower i n 1988 compared t o 1987 f o r both the f e r a l and domestic q u a i l (95%, n=59 vs 63%, n=76; X2 = 18.03, df = 1, P<0.05). For f e r a l females, 24% o f a l l eggs l a i d hatched. The c o r r e s p o n d i n g v a l u e f o r domestic females was s i m i l a r (29%). NEST PARASITISM Nest p a r a s i t i s m can o n l y occur i f the l a y i n g o r n e s t i n g p e r i o d of two or more females o v e r l a p s . In 1987, o n l y one f e r a l female was l a y i n g . In 1988, when the l a y i n g p e r i o d of the two f e r a l females overlapped, FY2 l a i d s i x eggs i n the n e s t of FB2 (with f i v e o f her own eggs). Only one o f these s i x eggs hatched compared w i t h two o f f i v e from the host female. The i n c i d e n c e o f more than one female l a y i n g o r n e s t i n g a t the same time o r o v e r l a p p i n g i n time was much h i g h e r f o r the domestic females (15 o f 21 n e s t s ) . Consequently, e i g h t o f the 21 n e s t s examined ( i n c l u d i n g non-inc u b a t e d ones) c o n t a i n e d eggs from two females. Only two c h i c k s hatched from 32 eggs l a i d i n o t h e r s ' n e s t s . Taking i n t o account both the f e r a l and domestic females, two cases o f nest p a r a s i t i s m o c c u r r e d a f t e r a female had d e s e r t e d her own nest and s t a r t e d l a y i n g i n another female's n e s t . In t h r e e o t h e r cases a female a l s o l a i d i n her own nest w h i l e she was l a y i n g i n another female's n e s t . In f o u r cases a female ( i n c l u d i n g one f e r a l female) l a i d eggs i n another female's n e s t p r i o r t o s t a r t i n g her own. In the 2 ye a r s of the study, no female l a i d eggs i n the n e s t o f another female without s t a r t i n g one of her own. The n e s t s w i t h eggs from two females had s i g n i f i c a n t l y lower h a t c h i n g success (X2=5.82, d f = l , P<0.05) than n e s t s c o n t a i n i n g eggs from o n l y one female. Four p a r a s i t i z e d n e s t s were d e s e r t e d e i t h e r b e f o r e o r a f t e r i n c u b a t i o n had s t a r t e d . CHICK MORTALITY Of the 99 c h i c k s hatched i n the f l i g h t - p e n s , 28% disap p e a r e d o r were found dead w i t h i n 21 days o f h a t c h i n g . M o r t a l i t y f o r f e r a l c h i c k s was h i g h e r (10/24, 42%) but not s t a t i s t i c a l l y d i f f e r e n t (X2=2.77, df=l) from t h a t f o r domestic c h i c k s (18/75, 24%). The m o r t a l i t y o f f e r a l c h i c k s was h i g h e r 78 i n 1988 (7/14, o r 50%) than 1987 (3/10), but was not t e s t e d s t a t i s t i c a l l y because o f s m a l l numbers. PARENT-YOUNG ASSOCIATION Domestic c h i c k s began t o move around the pen independently o f t h e i r p a rents (or f o s t e r parents) a f t e r a mean o f 16 (n=ll) days p o s t - h a t c h . F e r a l c h i c k s remained w i t h t h e i r p a r e n t s s i g n i f i c a n t l y (t=3.26, df=12, P=0.01) l o n g e r (23 days)(n=3). Domestic c h i c k s o f one brood f r e q u e n t l y mixed wit h those o f another. Except f o r one female, DG1, who had not y e t l a i d i n a nest, f o s t e r i n g was always done between broody females. Only two f e r a l broods overlapped and no c r o s s f o s t e r i n g was observed. The f e r a l females were v e r y a g g r e s s i v e t o ot h e r females and t o males t h a t approached t h e i r c h i c k s . T h i s l e v e l o f a g g r e s s i o n was seen i n o n l y one domestic female, DP2. EGG PRODUCTION Egg p r o d u c t i o n was measured by d i v i d i n g the t o t a l number of eggs found i n the pen (from May 1 t o September 30) by the number of females i n the pen. Data were a l s o c o l l e c t e d from the 12 s m a l l pens t h a t each housed a male and two females. L a y i n g began e a r l i e r i n the second year than the f i r s t f o r both f e r a l and domestic females. Egg l a y i n g peaked f o r domestic females i n J u l y d u r i n g the f i r s t y e ar. Corresponding t o the e a r l i e r s t a r t i n g date, the peak came i n June i n the second year. Egg l a y i n g peaked f o r f e r a l females i n J u l y f o r both y e a r s d e s p i t e the e a r l i e r s t a r t i n g date i n 1988 ( F i g . 5.1). In 1987, the f o u r domestic females from the f l i g h t - p e n produced an average of 28 eggs and i n 1988 an average of 39 eggs. The c o r r e s p o n d i n g r e c o r d s f o r f e r a l females were 9 and 16 eggs r e s p e c t i v e l y (Table 5.1). Domestic females i n the s m a l l pens produced c o n s i d e r a b l y more eggs than females i n the f l i g h t - p e n . Each female l a i d 70 and 80 eggs r e s p e c t i v e l y i n 1987 and 1988. On the o t h e r hand, f e r a l females i n the s m a l l pens each l a i d o n l y s i x eggs i n 1987, compared t o 46 eggs i n 1988. Very l i t t l e i n c u b a t i o n behaviour was observed i n the s m a l l pens. DISCUSSION Japanese q u a i l i n Hawaii, the source p o p u l a t i o n of the f e r a l q u a i l used i n t h i s experiment, have 5-8 egg c l u t c h e s (Schwarz and Schwarz 1949). F e r a l females i n my f l i g h t - p e n study l a i d c l u t c h e s o f 4-11 eggs. While t h i s range o f c l u t c h s i z e i s s l i g h t l y wider than t h a t r e p o r t e d by Schwarz and Schwarz (1949), i t i s s i m i l a r t o ranges r e p o r t e d f o r w i l d q u a i l i n Japan by Kawahara (1967) and by Wakasugi (1984) (7-12 and 7-14 eggs r e s p e c t i v e l y ) . Japanese q u a i l i n n o r t h e r l y l a t i t u d e s g e n e r a l l y have l a r g e r c l u t c h s i z e s than those Figure 5.1: Total egg production. Feral 50 -i May Jun Jul Aug Sep 1987 • 1988 Domestic 70 -i May Jun Jul Aug Sep 1987 • 1988 81 n e s t i n g n e a r e r the t r o p i c s (Dement'ev e t a l . 1967, Cramp and Simmons 1980). The f e r a l q u a i l i n the f l i g h t - p e n s , l i k e w i l d Japanese q u a i l , r a i s e d o n l y one brood per season (Dement'ev e t a l . 1967). S i m i l a r l y , the i n c i d e n c e of i n f e r t i l e eggs i n w i l d q a l l i f o r m p o p u l a t i o n s i s l e s s than 10% (Johnsgard 1973); my f e r a l q u a i l produced no i n f e r t i l e eggs. Thus, the f l i g h t - p e n c o n d i t i o n s i n t h i s study appeared t o s t i m u l a t e s i m i l a r l e v e l s o f r e p r o d u c t i o n i n f e r a l q u a i l as those found i n the w i l d . F E R A L F E M A L E S l . o nset o f b r e e d i n g : In the f l i g h t - p e n s , o n l y one y e a r l i n g female attempted t o breed and none of the y e a r l i n g females d i s p l a y e d r e p r o d u c t i v e behaviour ( i . e . , a sequence of a g g r e s s i v e , c o u r t s h i p , and mating a c t i v i t i e s , see Chapter 3) u n t i l mid June. In a v i a n s p e c i e s where y e a r l i n g s attempt t o breed, many do not have h i g h r e p r o d u c t i v e success compared w i t h o l d e r i n d i v i d u a l s (Crawford 1977; Blus and Keahey 1978), although age d i f f e r e n c e s were not seen f o r w i l l o w ptarmigan (Hannon and Smith 1984) . Where d i f f e r e n c e s o c c u r r e d , a p o s s i b l e e x p l a n a t i o n i s t h a t y e a r l i n g s do not compete w e l l w i t h 2 or 3 y e a r - o l d s f o r r e s o u r c e s (Lack 1968; Jamieson 1985) and t e r r i t o r y (Wittenberger 1978; Jamieson 1985) through l a c k of e x p e r i e n c e . However, y e a r l i n g female q u a i l were g i v e n ample n u t r i t i o n and s i m i l a r space t o 2 y e a r - o l d s , and they were not 82 competing w i t h o l d e r females. The f o u r 2 y e a r - o l d s a l l attempted t o breed soon a f t e r they were put i n the f l i g h t - p e n . T h i s suggests t h a t y e a r l i n g females were not q u i t e ready t o breed; they may have been more s e n s i t i v e t o d e n s i t y o r p h y s i o l o g i c a l l y a t a stage where b r e e d i n g attempts were more e a s i l y thwarted by unfavourable f a c t o r s (Sossinka 1982). High p o p u l a t i o n d e n s i t y e x i s t e d i n the s m a l l pens, where one of s i x y e a r l i n g s and seven o f e i g h t 2 y e a r - o l d s l a i d eggs. And although e x p e r i e n c e w i t h the pen environment has t o be c o n s i d e r e d , 60% of the females o f the o r i g i n a l Hawaiian s t o c k l a i d w i t h i n 2 months when p l a c e d i n the f l i g h t - p e n . While I cannot separate age e f f e c t s from year e f f e c t s , age i s the o n l y known d i f f e r e n c e between the two y e a r s . Johnsgard (1988) concluded t h a t most s p e c i e s o f q u a i l , p a r t r i d g e s , and f r a n c o l i n s mature d u r i n g t h e i r f i r s t y ear and are a b l e t o breed as y e a r l i n g s . He based h i s c o n c l u s i o n on the e a r l y attainment of a d u l t plumages, the apparent r a r i t y o f non-breeding females i n w i l d p o p u l a t i o n s , and e a r l y b r e e d i n g under c a p t i v e c o n d i t i o n s . I b e l i e v e t h a t w h i l e f e r a l (or wild) females can reproduce as y e a r l i n g s , they may not do so u n t i l they are 2 years o l d i f c o n d i t i o n s are unfavourable (e.g. h i g h p o p u l a t i o n d e n s i t y ) . 2. Nest Parasitism: F a c u l t a t i v e n est p a r a s i t i s m was seen f o r both f e r a l and domestic females. I t c o u l d be argued t h a t t h i s i s an a r t i f a c t 83 o f u n u s u a l l y h i g h b i r d d e n s i t y i n the f l i g h t - p e n s . However, both i n t e r - and i n t r a s p e c i f i c n e st p a r a s i t i s m has been documented f o r Bobwhite q u a i l ( C o l i n u s v i r o i n i a n u s ) (Holcomb 1968; P i a t t 1968), C a l i f o r n i a q u a i l (Lophortyx c a l i f o r n i c u s ) ( G e n e l l y 1955), Ring-necked pheasants (Phasianus c o l c h i c u s ) (Westemeier and Esker 1989), and S h a r p - t a i l e d grouse (Pedioecetes p h a s i a n e l l u s ) (Gratson 1989). The success of nest p a r a s i t i s m was poor i n the f l i g h t -pens, and few c h i c k s hatched from p a r a s i t i c eggs. However, nest p a r a s i t i s m may not have been a c c i d e n t a l or abnormal ( S i e g f r i e d 1976). Rather i t appears t o be an a l t e r n a t i v e r e p r o d u c t i v e behaviour, p o s s i b l y e i t h e r as a c o n d i t i o n a l or mixed s t r a t e g y (Eadie 1989). In t h i s study, most examples of nest p a r a s i t i s m were a response t o n e s t d i s r u p t i o n or e x c e s s i v e male d i s t u r b a n c e . However, on two o c c a s i o n s a female l a i d both i n her nest and the nest of another female f o r no obvious reason. T h i s may have been a mixed s t r a t e g y t h a t c o u l d i n c r e a s e her r e p r o d u c t i v e success (Evans 1988). 3 . I n c u b a t i o n and B r o o d i n g : In both y e a r s , o n l y one f e r a l female o f f o u r reproduced w i t h any success (1987:FP1, 1988:FG2). The b r e e d i n g d e n s i t y r e p o r t e d f o r f e r a l q u a i l i n Hawaii i s approximately one p a i r per h e c t a r e (Schwarz and Schwarz 1949; Robinson e t a l . i n p r e s s ) . Many s p e c i e s which normally e s t a b l i s h b r e e d i n g t e r r i t o r i e s become or g a n i z e d i n t o dominance o r d e r s when f o r c e d 84 t o g e t h e r i n s m a l l spaces (see review by Wilson 1975) . B e h a v i o u r a l o b s e r v a t i o n s i n d i c a t e d t h a t the f l i g h t - p e n c o n d i t i o n s d i d not cause the f e r a l q u a i l t o form o r g a n i z e d dominance h i e r a r c h i e s , but dominance and a g g r e s s i o n may a f f e c t who breeds s u c c e s s f u l l y under these c o n d i t i o n s . A more a g g r e s s i v e female may be a b l e t o d e t e r male advances d u r i n g i n c u b a t i o n o r d i s r u p t o t h e r females' n e s t i n g attempts (Hannon 1983; M a r t i n e t a l . 1990). However, t h e r e i s no evidence (Chapter 3, F i g . 3.3) t h a t t h i s o c c u r r e d i n the f l i g h t - p e n s . I t may be important f o r a female t o p a i r w i t h a more dominant male who can p r o t e c t her from d i s t u r b a n c e by o t h e r males d u r i n g egg l a y i n g and e a r l y i n c u b a t i o n (see Chapter 3). S t a n f o r d (1957) noted, "The male C o t u r n i x i s the most a v i d s u i t o r of any b i r d t h a t has been observed, a t times pushing the female o f f the nest t o c o p u l a t e . " He then s p e c u l a t e d t h a t i n many cases the female had t o h i d e from males t o brood the c l u t c h . In 1988, when more than one f e r a l female attempted t o breed, almost 50% of t h e eggs produced were l a i d o u t s i d e the n e s t s , s u g g e s t i n g t h a t many n e s t i n g attempts were d i s r u p t e d d u r i n g l a y i n g . F e r a l females i n the f l i g h t - p e n s o n l y i n i t i a t e d i n c u b a t i o n once, and when i n c u b a t i o n was d i s r u p t e d and the n e s t d e s e r t e d , t h e r e was no attempt t o r e n e s t . Renesting a f t e r n e s t d e s t r u c t i o n i s common i n many a v i a n s p e c i e s (Angelstam 1979; Parker 1981, 1985; Dumke and P i l s 1979). I t i s l i k e l y t h a t the h i g h d e n s i t y i n the f l i g h t - p e n s d e l a y e d the 85 f i r s t n e s t i n g attempts of the females, thus n e s t f a i l u r e s ( i n cases where i n c u b a t i o n had s t a r t e d ) a l s o o c c u r r e d q u i t e l a t e i n the season. The same h i g h d e n s i t y may a l s o have di s c o u r a g e d r e n e s t i n g . D O M E S T I C F E M A L E S In t h i s study domestic Japanese q u a i l stopped l a y i n g i n d e t e r m i n a t e l y and l a i d i n c l u t c h e s , b u i l t n e s t s , became broody, and r a i s e d c h i c k s i n the f l i g h t - p e n environment. C l u t c h s i z e (number of eggs i n a nest) and l e v e l s o f f e r t i l i t y and h a t c h a b i l i t y of domestic and f e r a l q u a i l were a l s o s i m i l a r . A t t h i s l e v e l of comparison, C l a y t o n ' s (1972) h y p o t h e s i s i s supported. P r e v i o u s r e p o r t s ( R o t h s t e i n 1967; Eynon 1968; and F a r r i s 1964) of f a i l u r e t o induce n e s t i n g i n outdoor a v i a r i e s probably r e s u l t e d from l a c k of space ( i . e . a l l o w i n g f o r d i s t u r b a n c e s by males) o r l a c k o f proper n e s t i n g cover. T h i s was the case i n the s m a l l h o l d i n g pens. Both f e r a l and domestic q u a i l i n the s m a l l pens f a i l e d t o nest, but s c a t t e r e d eggs throughout the pens. In the few cases where n e s t s were s t a r t e d , many eggs were found o u t s i d e the nest d u r i n g the next few days and no c l u t c h e s were completed. Kennedy (1991) c i t e d two mechanisms f o r l i m i t i n g c l u t c h s i z e : t a c t i l e s t i m u l a t i o n of eggs a g a i n s t the brood p a t c h i n g u l l s (Winkler and Walters 1983) and the p o s s i b l e response t o the v i s u a l s t i m u l i i of a c o l l e c t i o n of eggs by g a l l i f o r m e s (Steen and Parker 1981). As l o n g as eggs are s c a t t e r e d , the female 86 may keep on l a y i n g . D e s s i - F u l g h e r i (pers. comm.) suggested t h a t , compared t o those allowed t o form p a i r s n a t u r a l l y , c a p t i v e female gray p a r t r i d g e s t h a t have a mate s e l e c t e d f o r them are more l i k e l y t o l a y eggs s c a t t e r e d and l e s s l i k e l y t o nest and i n c u b a t e . T h i s may apply t o domestic Japanese q u a i l g i v e n t h a t the females i n the s m a l l pens had t h e i r mates s e l e c t e d f o r them and were more l i k e l y t o l a y s c a t t e r e d eggs than females i n the l a r g e pen. While the r e p r o d u c t i o n of domestic and f e r a l Japanese q u a i l i s s i m i l a r , t h e r e are some d i f f e r e n c e s . I t i s common f o r domestic b i r d s t o demonstrate advanced maturation, an, extended b r e e d i n g season and an i n c r e a s e d number of eggs (Sossinka 1982). l . E a r l i e r age of onset of b r e e d i n g : U n l i k e f e r a l females, a l l domestic females were r e p r o d u c t i v e l y f u n c t i o n a l as y e a r l i n g s . A l l y e a r l i n g females l a i d eggs i n the f l i g h t - p e n i n 1987, and many eggs were found i n the s m a l l pens t h a t year. B e h a v i o u r a l o b s e r v a t i o n s a l s o i n d i c a t e d t h a t y e a r l i n g domestic q u a i l formed p a i r s sooner than f e r a l q u a i l i n 1987 under the same c o n d i t i o n s (Chapter 3), but t h e r e was no d i f f e r e n c e i n t i m i n g f o r them as 2-year o l d s i n 1988. In the domestic turkey, s e x u a l m a t u r i t y i s reached a t 5 t o 7 months of age (Hale e t a l . 1969), whereas w i l d t u r k e y s are not mature u n t i l t h e i r second y e a r (Leopold 1944). P r e c o c i o u s n e s s i s a l r e a d y r e p o r t e d i n domestic q u a i l 87 (Meise 1954), and i s a l s o known i n domestic c h i c k e n s (Romanoff 1960) and geese (Lorenz 1940; Schmidt 1976). 2. E x t e n t o f the b r e e d i n g season and number o f eggs p r o d u c e d : Given the same amount of space and t h e same plane of n u t r i t i o n , domestic q u a i l i n the f l i g h t - p e n had a l o n g e r b r e e d i n g season, nested r e p e a t e d l y , and produced many more eggs than f e r a l q u a i l . P r e v i o u s s t u d i e s have r e v e a l e d d i f f e r e n c e s i n t e r r i t o r i a l behaviour between w i l d and domestic c o n s p e c i f i c s (Sossinka 1982). T e r r i t o r i a l i t y i s much l e s s pronounced i n domestic c h i c k e n s than i n w i l d r e d j u n g l e fowl and f e r a l c h i c k e n s (Hughes e t a l . 1974; L i l l 1968; McBride e t a l . 1969; S i e g e l 1976). The average i n d i v i d u a l d i s t a n c e i n w i l d m a l l a r d s i s a l s o l a r g e r than t h a t o f domestic A y l e s b u r y ducks (Deforges and Wood-Gush 1975). Given the same amount of space, reduced t e r r i t o r i a l i t y enhances r e p r o d u c t i o n i n domestic q u a i l . W i l d b i r d s r a i s e d i n c a p t i v i t y c o u l d s t i l l e x p e r i e n c e g r e a t e r s t r e s s (e.g., more a g i t a t e d when human c o n t a c t occurs) than domestic b i r d s , t h i s would a f f e c t t h e i r b r e e d i n g success ( P h i l l i p s 1964). So although both w i l d and domestic c o n s p e c i f i c s were exposed t o the "same" environment, the i n t e r a c t i o n o f genotype and environment does not a l l o w the c l e a r s e p a r a t i o n o f environmental and g e n e t i c e f f e c t s needed to a s s e s s the v a l i d i t y o f C l a y t o n ' s (1972) c l a i m s . 88 3 . P a r e n t a l Behaviour: In t h i s study domestic q u a i l c h i c k s took 16 days t o break the parent-young a s s o c i a t i o n , compared t o 23 days f o r f e r a l c h i c k s . A s i m i l a r f i g u r e f o r domestic Japanese q u a i l (12 days) was noted by S t a n f o r d (1957) . G e n e t i c s t u d i e s o f domesticated w i l d Japanese q u a i l (Kawahara and M i t a 1968; Kawahara 1972) have shown a r a p i d i n c r e a s e i n growth and egg p r o d u c t i o n over t h r e e g e n e r a t i o n s . Domestic animals g e n e r a l l y have a f a s t e r growth r a t e compared t o t h e i r w i l d c o n s p e c i f i c s ( P r i c e 1984). The younger f l e d g i n g age o f domestic q u a i l c h i c k s may be due, i n p a r t , t o t h e i r f a s t e r growth r a t e . On the o t h e r hand, domestic females may be l e s s broody because of t h e i r s t r o n g e r urge t o l a y more eggs. A l l except one domestic female d e s e r t e d t h e i r f i r s t n e s t of eggs. Many breeds of domestic c h i c k e n s no l o n g e r develop b r o o d i n e s s . In g e n e r a l , a t r e n d towards a d i m i n i s h e d brooding and c a r e - t a k i n g a c t i v i t y o c curs i n domestic b i r d s (Guhl and F i s h e r 1969; Hale 1969). In the f l i g h t - p e n study, domestic females w i t h younger c h i c k s always took over o l d e r broods from t h e i r parent, i n d i c a t i n g t h a t b r o o d i n e s s i n domestic q u a i l i s s t r o n g soon a f t e r the c h i c k s hatched, but drops o f f more q u i c k l y than f o r f e r a l females. I m p r i n t i n g was a l s o weaker i n domestic q u a i l compared t o f e r a l q u a i l . D e s p i t e d i m i n i s h e d b r o o d i n e s s of domestic females, c h i c k m o r t a l i t y was not s i g n i f i c a n t l y d i f f e r e n t from t h a t o f f e r a l q u a i l . C h ick m o r t a l i t y o f f e r a l q u a i l was h i g h i n 1988 when more than one female was 89 brooding. IMPLICATIONS T h i s study compares r e p r o d u c t i o n i n domestic and f e r a l Japanese q u a i l under the same s e t o f environmental c o n d i t i o n s . The d i f f e r e n c e between the two should r e f l e c t g e n e t i c d i f f e r e n c e s and thus the e f f e c t s o f d o m e s t i c a t i o n on r e p r o d u c t i o n (Sossinka 1982). However, because o f genotype x environment i n t e r a c t i o n s , one should compare the two types of q u a i l i n more than one s e t o f environmental c o n d i t i o n s t o understand the e f f e c t s of d o m e s t i c a t i o n more f u l l y and t o get b e t t e r i n f o r m a t i o n t h a t can be a p p l i e d t o c a p t i v e b r e e d i n g and r e s t o c k i n g p r o j e c t s . For example, i f a v a i l a b l e n u t r i e n t s are l i m i t e d , w i l l the performance of the f e r a l and domestic q u a i l change i n the same d i r e c t i o n and magnitude? The same genotype x environment i n t e r a c t i o n a l s o make a s s e s s i n g C l a y t o n ' s (1972) h y p o t h e s i s d i f f i c u l t . The "removal of p r o h i b i t i n g and a d d i t i o n o f enhancing s t i m u l i i " ( C l a y t o n 1972) i n v o l v e s not o n l y changes i n environmental c o n d i t i o n s but a l s o p a r t s o f the genotype not d i r e c t l y a f f e c t i n g r e p r o d u c t i o n . N e v e r t h e l e s s , i f one l o o k s a t each component independently, one can argue t h a t the change i s not d r a s t i c ( r e : C l a y t o n 1972), but t o g e t h e r , the consequence c o u l d be s e r i o u s . L i t t l e changes i n each component r e s u l t s i n a s e r i o u s l a c k of cohesiveness of these components and the necessary " f i n e t u n i n g " t o the environment. 91 C H A P T E R S I X G E N E R A L D I S C U S S I O N Once removed from a domestic s e t t i n g and p l a c e d i n a s e m i n a t u r a l environment, domestic Japanese q u a i l e x h i b i t e d a b e h a v i o u r a l r e p e r t o i r e as r i c h as t h a t shown by the f e r a l q u a i l taken from the w i l d . T h i s c o n t r a d i c t s the view t h a t d o m e s t i c a t i o n has had a " d e g e n e r a t i v e " e f f e c t on behaviour (Leopold 1944; Lorenz 1965) and supports the i d e a t h a t the absence o f these d i s p l a y s i n a domestic s e t t i n g i s a f u n c t i o n of environmental c o n s t r a i n t s ( C l a y t o n 1972). F a i l u r e s i n v o l v i n g r e s t o c k i n g o f domestic o r c a p t i v e b i r d s have o f t e n l e d observers t o conclude t h a t the b i r d s have l o s t the a b i l i t y t o behave and reproduce "normally". An example of these f a i l u r e s i s the i n t e n s i v e i n t r o d u c t i o n program f o r Japanese q u a i l throughout North America conducted i n the e a r l y 1900's ( P h i l l i p s 1928; McAtee 1944; S t a n f o r d 1957; L a b i s k y 1961). A l l s t o c k i n g attempts f a i l e d , and L a b i s k y (1961) blamed i t on g e n e t i c l o s s o f " w i l d n e s s " and v i g o r due t o d o m e s t i c a t i o n . T h i s was r e i n f o r c e d by repeated f a i l u r e s t o get domestic Japanese q u a i l t o brood c h i c k s i n the l a b o r a t o r y ( R o t h s t e i n 1967; F a r r i s 1964). My study however, has demonstrated how a domestic s t r a i n of Japanese q u a i l t h a t has shown no broody behaviour i n 63 g e n e r a t i o n s , i s not o n l y capable of becoming broody, but can s u c c e s s f u l l y r a i s e a brood t o f l e d g i n g . Comparisons between the f e r a l and domestic q u a i l a l s o r e v e a l e d d i f f e r e n c e s i n t h e i r r e p r o d u c t i v e behaviour. These d i f f e r e n c e s were p r i m a r i l y a matter o f degree. For example, although p a i r bonds and p arent c h i c k a s s o c i a t i o n s formed, they were found t o be weaker i n the domestic q u a i l . These v a r i a t i o n s might e f f e c t the s u r v i v a b i l i t y o f domestic q u a i l r e t u r n e d t o the w i l d . A c o n t r o l l e d study such as t h i s a l l o w s one t o determine d i f f e r e n c e s t h a t might go undetected i n a f i e l d study. Given an understanding of these v a r i a t i o n s , s t e p s (presented below) can be taken t o reduce t h e i r impact when r e l e a s i n g b i r d s i n t o the w i l d . IMPLICATIONS I n v e s t i g a t i o n s o f r e s t o c k i n g f a i l u r e s i n d i c a t e t h a t c a p t i v e r e a r e d and r e l o c a t e d b i r d s are more s u s c e p t i b l e t o p r e d a t i o n ( H e s s l e r e t a l . 1970; P o t t s 1986; and Krauss, e t a l . 1987), poor w i n t e r s u r v i v a l (Bouchner and Temmlova 1974, 1975) and have lower n e s t i n g success (Rands and Hayward 1987). They a l s o i n d i c a t e t h a t the p e r i o d immediately a f t e r r e l e a s e accounts f o r h i g h numbers of b i r d m o r t a l i t y ( S c h l a d w e i l e r and T e s t e r 1972; Krauss e t a l . 1987). These f a c t o r s are not m u t u a l l y e x c l u s i v e but are interwoven. During r e l e a s e s the h i g h i n i t i a l m o r t a l i t y , i n c r e a s e d s u s c e p t i b i l i t y t o p r e d a t i o n , and lower n e s t i n g success can be a t t r i b u t e d t o a lower waryness of r e l e a s e d b i r d s compared t o t h e i r w i l d c o u n t e r p a r t s ( S c h l a d w e i l e r and T e s t e r 1972; E l l s w o r t h e t a l . 1988). My study, although not designed t o l o o k a t "tameness", d i d show b e h a v i o u r a l d i f f e r e n c e s between domestic and f e r a l q u a i l i n t h i s r e g a r d . I noted t h a t domestic q u a i l i n open areas o r when moving from one l o c a t i o n t o another r a r e l y demonstrated c a u t i o u s n e s s . In c o n t r a s t , f e r a l q u a i l under s i m i l a r c o n d i t i o n s were always a l e r t t o b i r d s f l y i n g overhead and r e a c t e d immediately t o sudden n o i c e s by d a r t i n g f o r cover o r f r e e z i n g . S c h l a d w e i l e r and T e s t e r (1972) showed t h a t hand-reared m a l l a r d s can l e a r n t o be more wary by a s s o c i a t i n g w i t h w i l d c o n s p e c i f i c s and consequently have b e t t e r s u r v i v a l . I n c r e a s e d v o c a l i z a t i o n by domestic q u a i l c o u l d a l s o make them more s u s c e p t i b l e t o p r e d a t i o n . F e r a l q u a i l o n l y crowed when l o o k i n g f o r p o t e n t i a l mates o r r e u n i t i n g w i t h t h e i r mates upon s e p a r a t i o n . T h i s was done by u s i n g d i f f e r e n t loudness of crows (see Chapter 3 ). Thus when a male was near h i s mate, but had l o s t v i s u a l c o n t a c t , he would crow s o f t l y . The domestic q u a i l , however, crowed more o f t e n , i n a l o u d volume, and w i t h l e s s v a r i a t i o n . Although t h i s c a l l i s important i n a l l o w i n g a male t o s t a y i n c o n t a c t w i t h h i s mate, i t a l s o p r o v i d e s p r e d a t o r s a t a r g e t . The n o n - d i s c r e t e use of v o c a l i z a t i o n by domestic q u a i l would i n c r e a s e t h e i r exposure t o p r e d a t i o n . S u c c e s s f u l r e s t o c k i n g depends upon the a b i l i t y o f the i n t r o d u c e d b i r d s t o breed. Although i n the f l i g h t - p e n s , the domestic q u a i l were as s u c c e s s f u l a t b r e e d i n g as the f e r a l , t h e i r reduced l e n g t h of p a i r bonds, g r e a t e r number o f FEPC's, decreased mate guarding and the i n c r e a s e d d u r a t i o n of the l a y i n g p e r i o d c o u l d reduce t h e i r s u r v i v a b i l i t y . K u r z e j e s k i and Root (1988) found t h a t grouse w i t h l a r g e d a i l y movements had s i g n i f i c a n t l y lower s u r v i v a l r a t e s than more sedentary i n d i v i d u a l s . Female's e x c u r s i o n s are more e x t e n s i v e when s e a r c h i n g f o r nest s i t e s ( s t a r t i n g approximately two weeks p r i o r t o l a y ) . S i n c e domestic q u a i l p a i r an average o f f o u r days p r i o r t o l a y i n g , compared t o 22 f o r f e r a l , the domestic females l a c k a male s e n t r y f o r d e t e c t i n g p r e d a t o r s and guarding a g a i n s t i n t r u d i n g males d u r i n g t h i s p e r i o d (Martin 1984). The weaker p a i r bonds and i n c r e a s e d e x c u r s i o n s f o r FEPC's by domestic males exposes both the male and h i s mate t o i n c r e a s e d p r e d a t i o n . However, g i v e n t h a t t h i s study found a h i g h degree o f v a r i a b i l i t y between domestic p a i r s f o r p a i r bond l e n g t h and the number of male FEPC e x c u r s i o n s , i t seems reasonable t o assume t h a t n a t u r a l s e l e c t i o n , i f g i v e n time, w i l l be a b l e t o use these v a r i a t i o n s i n r e l e a s e d p o p u l a t i o n s t o complete the p r o c e s s o f f e r a l i z a t i o n . S t u d i e s of d i f f e r e n t g a l l i f o r m s p e c i e s have shown t h a t female s u r v i v a b i l i t y was a t i t s lowest d u r i n g the n e s t i n g p e r i o d (Vander Haegen e t a l . 1988; P o t t s 1986). Nest s i t e s chosen by some domestic q u a i l were not as w e l l hidden as f o r f e r a l s , and they were l e s s s e c r e t i v e when l e a v i n g and r e t u r n i n g t o the n e s t . T h i s c o u l d expose the females and/or t h e i r n e s t s t o h i g h e r l e v e l s o f p r e d a t i o n and thus lower n e s t i n g s u c c e s s . P r e d a t i o n on n e s t i n g females and t h e i r 95 c l u t c h e s i s a major cause f o r low n e s t i n g success i n gray p a r t r i d g e s ( P o t t s 1986; Rands and Hayward 1987) and w i l d t u r k e y s ( M e l e a g r i s g a l l o p a v o s i l v e s t r i s ) ( V a n g i l d e r e t a l . 1987) . The l o n g e r b r e e d i n g season o f domestic q u a i l c o u l d have both a p o s i t i v e and n e g a t i v e e f f e c t on t h e i r r e p r o d u c t i v e s u c c e s s . By p r o d u c i n g s e v e r a l c l u t c h e s i n a season the domestic q u a i l stand t o i n c r e a s e t h e i r r e p r o d u c t i v e s u c c e s s . However, b r e e d i n g too e a r l y or too l a t e i n the season i n c r e a s e s the chance of h a t c h i n g a c l u t c h when the weather c o n d i t i o n s a r e u n f a v o u r able and food i s s c a r c e . S t u d i e s of Hungarian p a r t r i d g e (Weigand 1980) and C a l i f o r n i a q u a i l (Leopold e t a l . 1976) have shown t h a t e x c e s s i v e r a i n f a l l d u r i n g the n e s t i n g and h a t c h i n g p e r i o d s s e r i o u s l y reduce b r e e d i n g s u c c e s s . The c h i c k s u r v i v a l o f gray p a r t r i d g e ( P o t t s 1973) i n the f i r s t s i x weeks a f t e r h a t c h i n g was h i g h l y c o r r e l a t e d t o i n s e c t food a v a i l a b i l i t y . By i n c r e a s i n g the b r e e d i n g season, the p a i r s , and p a r t i c u l a r l y the females, may not have enough time t o c o n d i t i o n themselves f o r the w i n t e r months. S t u d i e s of r e l e a s e s of hand-reared p a r t r i d g e have shown t h a t h i g h m o r t a l i t y i n the f i r s t y ear i s due t o h i g h w i n t e r l o s s e s (Paludan 1963; Bouchner and Temmlova 1974, 1975). Domestic q u a i l are v e r y adaptable; I b e l i e v e t h a t under n a t u r a l c o n d i t i o n s the l e n g t h of t h e i r b r e e d i n g season c o u l d be a f f e c t e d by food a v a i l a b i l i t y and, as a r e s u l t , be s h o r t e r than i t was i n the f l i g h t pen. 96 RECOMMENDATIONS Domestic Japanese q u a i l have the f l e x i b i l i t y t o adapt t o a " w i l d " environment, as i n d i c a t e d by the f e r a l p o p u l a t i o n o f q u a i l t h a t e x i s t s today on the Hawaiian I s l a n d s . Why d i d these q u a i l s u r v i v e when so many o t h e r i n t r o d u c t i o n s f a i l e d ? Many have s p e c u l a t e d , but a l a c k o f knowledge of t h e i r r e p r o d u c t i v e b i o l o g y has made i t d i f f i c u l t t o draw c o n c l u s i o n s . So too has a l a c k o f d e t a i l e d i n f o r m a t i o n about the h i s t o r y o f q u a i l r e l e a s e s ( i . e . o r i g i n o f s t r a i n s , r e a r i n g c o n d i t i o n s , age of b i r d s a t time o f r e l e a s e ) . I b e l i e v e t h a t domestic Japanese q u a i l , g i v e n an environment where p r e d a t o r d e n s i t y i s c o n t r o l l e d , have a good chance o f s u r v i v i n g ; they demonstrated h i g h l e v e l s o f v a r i a b i l i t y f o r many r e p r o d u c t i v e t r a i t s t h a t under n a t u r a l s e l e c t i o n c o u l d be enhanced t o the l e v e l o f w i l d q u a i l . For r e s t o c k i n g w i t h c a p t i v e r e a r e d o r domestic q u a i l t o be s u c c e s s f u l , a t t e n t i o n must be g i v e n t o t h e i r r e a r i n g environment. Thus b i r d s should be r a i s e d i n a n a t u r a l environment where they have t o se a r c h f o r food and water. A minimum o f human c o n t a c t i s e s s e n t i a l and a d d i t i o n a l b e n e f i t s would be gained by r a i s i n g domestic s t o c k w i t h w i l d b i r d s . Bouchner and Temmlova (1974) found t h a t p r e d a t o r avoidance behaviour q u i c k l y developed i n young r e l e a s e d p a r t r i d g e s a f t e r they were adopted by w i l d a d u l t s . P e r i o d i c exposure t o p r e d a t o r s (e.g. foxes) , o r even mock p r e d a t o r s , may be u s e f u l . F i n a l l y , t h e r e are i n d i c a t i o n s t h a t the success o f 97 i n t r o d u c t i o n s might be i n c r e a s e d by u s i n g o l d e r b i r d s . K u r z e j e s k i and Root (1988) found t h e r e was a lower s u r v i v a l r a t e f o r r e l o c a t e d immature grouse than a d u l t grouse. FUTURE STUDIES T h i s study has l a i d a f o u n d a t i o n f o r f u t u r e r e s e a r c h i n t o the e f f e c t s o f d o m e s t i c a t i o n on the r e p r o d u c t i v e behaviour of Japanese q u a i l and p o s s i b l y o t h e r g a l l i f o r m e s . I t would be i n t e r e s t i n g t o study the r e p r o d u c t i o n o f domestic q u a i l when they have t o forage f o r t h e i r own food and water, and see how t h i s a f f e c t s t h e i r p h y s i c a l c o n d i t i o n and a b i l i t y t o s u r v i v e the w i n t e r . I t would a l s o be u s e f u l t o see how b i r d d e n s i t y a f f e c t s the mating system o f w i l d and domestic Japanese q u a i l . F i n a l l y , more c o n t r o l l e d and comparative s t u d i e s a re needed t o look a t the g e n e t i c and environmental a f f e c t s on r e p r o d u c t i o n of domestic Japanese q u a i l . These s t u d i e s should be conducted w i t h d i f f e r e n t p o p u l a t i o n s o f domestic and w i l d Japanese q u a i l , s i n c e s u b t l e o r major d i f f e r e n c e s may e x i s t between p o p u l a t i o n s . 98 LITERATURE CITED Abplanalp, H., Woodard, A.E. and W. 0. Wilson. 1963. The e f f e c t s o f u n n a t u r a l day l e n g t h s upon m a t u r a t i o n and egg p r o d u c t i o n o f Japanese q u a i l C o t u r n i x c o t u r n i x j a p o n i c a . P o u l t r y S c i . 41:1963-68. Angelstam, P. 1979. Black grouse (Lyrurus t e t r i x ) r e p r o d u c t i v e success and s u r v i v a l r a t e i n peak and c r a s h s m a l l rodent years i n c e n t r a l Sweden. World Pheasant Assoc. Woodland Grouse Symp. pplOl-111. Arad, Z., Marder, J . and M. S o l l e r . 1981. E f f e c t o f g r a d u a l a c c l i m a t i o n t o temperatures up t o 44°C on p r o d u c t i v e performance o f the d e s e r t Bedouin fowl, the commercial White Leghorn and the two r e c i p r o c a l c r o s s b r e d s . B r i t . P o u l t r y S c i . 22:511-520. A u s t i n , O.L. and N. Kuroda. 1953. The b i r d s o f Japan: t h e i r s t a t u s and d i s t r i b u t i o n . B u l l . Mus. Comp. Z o o l . 109:277-612. Backs, S.E. 1982. An e v a l u t a t i o n o f r e l e a s i n g f i r s t g e n e r a t i o n (F l ) bobwhite q u a i l produced from w i l d s t o c k . Ind. Dep. Nat. Resour., Pittman-Robertson B u l l . 14. 17pp. B a i l l e y , J.B. 1854. O r n i t h o l o g i e de l a Savoie. B a i l l y , P a r i s . 201p. Bannerman, D.A. 1963. C o t u r n i x q u a i l . In: B i r d s o f the B r i t i s h I s l e s . V o l . 12. O l i v e r and Boyd, Edinburgh. Beach, F.A. and N.G. Inman. 1965. E f f e c t s o f c a s t r a t i o n and androgen replacement on mating i n male q u a i l . Proc. N a t l . Acad. S c i . U.S.A. 54:1426-31. Beani, L. and F. D e s s i - F u l g h e r i . 1985. Gray p a r t r i d g e r e s t o c k i n g : A problem o f a p p l i e d ethology. Monitore Z o o l . I t a l . 19:144-145. Birkhead, T.R. 1987. Sperm c o m p e t i t i o n i n b i r d s . Trends E c o l . E v o l . 2:268-272. Blank, T.H. and J.S. Ash. 1960. Some asp e c t s o f c l u t c h s i z e i n the p a r t r i d g e ( P e r d i x p e r d i x ) . Proceedings o f the 12th I n t e r n a t i o n a l O r n i t h o l g i c a l Congress, p p l l 8 - 2 6 . Blus, L . J . and J.A. Keahey. 1978. V a r i a t i o n i n r e p r o d u c t i v i t y w i t h age i n the Brown P e l i c a n . Auk 95:128-134. Boice, R. 1972. Some b e h a v i o u r a l t e s t s o f d o m e s t i c a t i o n i n 99 Norway r a t s . Behaviour 42:198-231. Boice, R. 1973. Domestication. Psych. B u l l . 80:215-30. Bouchner, M. and B. Temmlova. 1974. F a c t o r s i n f l u e n c i n g the s u r v i v a l o f p a r t r i d g e s from h a n d - r e a r i n g a f t e r the r e l e a s e i n t o f r e e h u n t i n g grounds. Prace Vulhm 45:73-92. Bouchner, M. and B. Temmlova 1975. S o c i a l behaviour o f young p a r t r i d g e s from h a n d - r e a r i n g a f t e r the r e l e a s e . Prace Vulhm 47:43-53. (In C z e c h o s l o v a k i a n w i t h summary i n E n g l i s h ) . Burns, J.T. 1982. Nests, t e r r i t o r i e s and r e p r o d u c t i o n o f Sedge Wrens ( C i s t o t h o r u s p l a t e n s i s ) . Wilson B u l l . 94:338-349. Carey, M. and V. Nolan J r . 1975. Polygyny i n Indigo Buntings: A h y p o t h e s i s t e s t e d . S c i ence 190:1296-1297. C l a y t o n , G.A. 1972. E f f e c t s of s e l e c t i o n on r e p r o d u c t i o n i n a v i a n s p e c i e s . J . Reprod. F e r t . Suppl. 15:1-21. Cramp, S. and K.E.L. Simmons (eds). 1980. The b i r d s of the western P a l e a r c t i c V o l . 2. Oxford U n i v e r s i t y P r e s s , Oxford. Crawford, R.D. 1977. Breeding b i o l o g y o f year o l d and o l d e r female Red-winged and Yellow-headed b l a c k b i r d s . W ilson B u l l . 89:73-80. Daan, S., C. D i j k s t r a , R. Drent and T. M e i j e r . 1988. Food supply and the annual t i m i n g o f a v i a n r e p r o d u c t i o n . A c t a XIX Cong. Proc. I n t e r n . O r n i t h o l . Univ. o f Ottawa P r e s s . Ottawa, Canada. Desforges, M.F. and D.G.M. Wood-Gush. 1975. A b e h a v i o u r a l comparison o f domestic and m a l l a r d ducks: h a b i t u a t i o n and f l i g h t r e a c t i o n s . Animal Behaviour 23: 692-697. Dement'ev, G.P., N.A. Gladkov, Y. Isakov, N.M. Kartashev, S.V. K i r i k o v , A.V, Mikheev and E.S. Ptushenko. 1967. In: B i r d s of the S o v i e t Union, V o l . IV (G.P. Dement*ev and N.A.Gladkov, e d s . ) , . I s r a e l Program f o r S c i e n t i f i c T r a n s l a t i o n s , Jerusalem. ppl45-162. Dumke, R.T. and CM. P i l s . 1979. Renesting and dynamics o f nest s i t e s e l e c t i o n by Wisconsin pheasants. J . W i l d l . Manage. 43:705-716. Eadie, J.M. 1989. A l t e r n a t i v e r e p r o d u c t i v e t a c t i c s i n a p r e c o c i a l b i r d : the ecology and e v o l u t i o n o f brood p a r a s i t i s m i n goldeneyes. Ph.D. T h e s i s , The Univ. o f B r i t i s h Columbia. E l l s w o r t h , D. L., Roseberry, J . L., and W. D. K l i m s t r a . 1988. 100 B i o c h e m i c a l g e n e t i c s o f w i l d , semi-wild and game-farm n o r t h e r n bobwhites. J . W i l d l . Manage. 52:138-144. E r i c k s o n , C.J. and P.G. Zenone. 1976. C o u r t s h i p d i f f e r e n c e s i n male r i n g doves: Avoidance o f cu c k o l d r y ? S c i e n c e 192:1353-1354. Evans, P.G. 1988. I n t r a s p e c i f i c n e st p a r a s i t i s m i n t h e European s t a r l i n g . Sturnus v u l g a r i s . Anim. Behav. 36:1282-1294. Eynon, A.E. 1968. The a g o n i s t i c and se x u a l b e h a v i o r o f c a p t i v e Japanese Q u a i l ( C o t u r n i x c o t u r n i x i a p o n i c a ) . Ph.D. T h e s i s , Univ. o f Wisconsin. F a r r i s H.E. 1964. B e h a v i o r a l development, s o c i a l o r g a n i z a t i o n , and c o n d i t i o n i n g o f c o u r t i n g b e h a v i o r i n the Japanese Q u a i l C o t u r n i x c o t u r n i x j a p o n i c a . Ph.D. T h e s i s , Mich. S t a t e Univ., E a s t Lansing. Ford,. N.L. 1983. V a r i a t i o n i n mate f i d e l i t y i n monogamous b i r d s . Curr. O r n i t h o l . 1:329-356. F o s t e r , M.S. 1974. Rain, f e e d i n g behavior, and c l u t c h s i z e i n t r o p i c a l b i r d s . Auk 91:722-726. Fraga, Rj.M. 1972. Coopera t i v e b r e e d i n g and a case o f s u c c e s s i v e polyandry i n the Bay-winged Cowbird. Auk 89:447-449. G e n e l l y , R.E. 1955. Annual c y c l e i n a p o p u l a t i o n o f C a l i f o r n i a q u a i l . Condor 57:263-285. Gibbs, H.L., P.J. Weatherhead, P.T. Borg, B.N. White, L.M. Tabak and D.J. Hoysak. 1990. R e a l i z e d r e p r o d u c t i v e success o f polygynous red-winged b l a c k b i r d s r e v e a l e d by DNA markers. S c i e n c e 250:1394-1397. Gladstone, D.E. 1979. P r o m i s c u i t y i n c o l o n i a l monogamous b i r d s . Am. Nat. 114:545-557. Gratson, M.W. 1989. I n t r a s p e c i f i c n e s t p a r a s i t i s m by S h a r p - t a i l e d grouse. Wilson B u l l . 101:126-127. Guhl, A.M. and G.J. F i s c h e r . 1969. The behaviour o f c h i c k e n s . In: The Behaviour o f Domestic Animals. (E.S.E. Hafez, ed.), B a i l l i e r e , T i n d a l l and C a s s e l l , London. pp515-553. Haase, E. and R.S. Donham. 1980. Hormones and d o m e s t i c a t i o n . In: A v i a n e n d o c r i n o l o g y (A. Epple and M. S t e t s o n , e d s . ) . Academic p r e s s , New York. Hale, E.B. 1969. Domestication and e v o l u t i o n o f behaviour. In: The Behaviour o f Domestic Animals. (E.S.E. Hafez, e d .), 101 B a i l l i e r e , T i n d a l l and C a s s e l l , London. pp22-42. Hale, E.B., W.M. S c h l e i d t and Schein, M.W. 1969. The behaviour of t u r k e y s . In: The Behaviour o f Domestic Animals (E.S.E. Hafez, e d.), B a i l l i e r e , T i n d a l l and C a s s e l l , London. pp554-592. Hannon, S.J> 1983. Spacing and b r e e d i n g d e n s i t y o f w i l l o w ptarmigan i n response t o experimental a l t e r a t i o n o f sex r a t i o . J . Anim. E c o l . 52:807-820. Hannon, S.J. and J.N.M. Smith. 1984. F a c t o r s i n f l u e n c i n g age-r e l a t e d r e p r o d u c t i v e success i n the w i l l o w ptarmigan. Auk 101:848-854. H e s s l e r , E., T e s t e r , J.R., S i n i f f , D.B. and M.M. Nelson. 1970. A b i o t e l e m e t r y study o f s u r v i v a l o f pen-reared pheasants r e l e a s e d i n s e l e c t e d h a b i t a t s . J . W i l d l . Manage. 34:267-274. Holcomb, L.C. 1968. F u r t h e r o b s e r v a t i o n on Ring-necked Pheasant n e s t i n g . Bird-Banding 39:133. Howes, J.R. 1964. The e f f e c t s o f season on o v u l a t i o n r a t e and egg composition of C o t u r n i x q u a i l s . Proc. Ass. 61st A g r i c . Wkrs. P o u l t r y s e c t i o n , A t l a n t a . pp283-284. Hughes, D.G., D.G.M. Wood-Gush and R.M. Jones. 1974. S p a t i a l o r g a n i z a t i o n i n f l o c k s o f domestic f o w l . Anim. Behav. 22, 438-445. Jamieson, I. 1985. Behavior of y e a r l i n g male b l u e grouse and i t s r e l a t i o n t o delayed b r e e d i n g . Wilson B u l l . 97: 71-77. J e n k i n s , D. (ed.) 1990. A c o n s e r v a t i o n s t r a t e g y f o r the g a l l i f o r m e s 1990-1992. World Pheasant A s s o c i a t i o n , Reading. Johnsgard, P.A. 1973. Grouse and q u a i l s of North America. U n i v e r s i t y o f Nebraska Press, L i n c o l n . Johnsgard, P.A. 1988. The q u a i l s , p a r t r i d g e s , and f r a n c o l i n s o f the world. Oxford U n i v e r s i t y P r e s s . Jones, J.M., M.A. Maloney and J . G i l b r e a t h . 1964. S i z e shape and c o l o u r p a t t e r n as c r i t e r i a f o r i d e n t i f y i n g C o t u r n i x eggs. P o u l t . S c i . 43:1292-1294. Kawahara, T. 1967. Wi l d C o t u r n i x q u a i l i n Japan. Q u a i l Quart. 4:62-63. Kawahara, T. 1971. Comparative study of q u a n t i t a t i v e t r a i t s between w i l d and domestic Japanese q u a i l ( C o t u r n i x c o t u r n i x j a p o n i c a ) . Exp. Animals, v o l . 22 Suppl., 1973: "Proceedings 102 o f the ICLA A s i a n P a c i f i c Meeting o f Labo r a t o r y Animals Sept. 20-25, 1971, Tokyo and Inuyama". Kawahara, T. 1972. G e n e t i c changes o c c u r i n g i n w i l d q u a i l s due t o " n a t u r a l s e l e c t i o n " under d o m e s t i c a t i o n . Ann. Rep. Nat. I n s t . G e n e t i c s (Japan) 22:111-112. Kawahara, T. and A. M i t a . 1968. A comparative a n a l y s i s o f p r o d u c t i v e t r a i t s i n w i l d and domesticated Japanese q u a i l s . Ann. Rep. Nat. I n s t . G e n e t i c s , Japan, 19:97-98. Kennedy, E.D. 1991. Determinate and in d e t e r m i n a t e e g g - l a y i n g p a t t e r n s : A review. Condor 93:106-124. Kimura, M. 1991. A t r a d i t i o n a l method f o r c a p t u r i n g w i l d Japanese q u a i l and p r o f i l e o f the numbers o f q u a i l c a p t u r e d i n Japan d u r i n g the p a s t 60 y e a r s . Jap. P o u l t r y S c i . 28:166-172. K l i m s t r a , W.D. 1975. Harvest r e t u r n s o f pen-reared bobwhite q u a i l . Trans. 111. S t a t e Acad. S c i . 68:278-284. Klomp, H. 1970. The d e t e r m i n a t i o n o f c l u t c h s i z e i n b i r d s . A review. Ardea, 58:1-24. Kosin, I. 1944. Macro- and m i c r o s c o p i c methods o f d e t e c t i n g f e r t i l i t y i n unincubated eggs. P o u l t r y S c i . 23:266-269. Krauss, G.D., H.B. Graves and S.M. Zervanos. 1987. S u r v i v a l o f w i l d and game-farm cock pheasants r e l e a s e d i n Pe n n s y l v a n i a . J . W i l d l . Manage. 51:555-559. K u r z e j e s k i , E. W. and B. G. Root. 1988. S u r v i v a l o f r e i n t r o d u c e d R u f f e d grouse i n M i s s o u r i . J . W i l d l . Mgmt. 52:248-252. Labisky, R.F. 1961. Report of attempts t o e s t a b l i s h Japanese q u a i l i n I l l i n o i s . J . W i l d l . Manage. 25:290-295. Lack, D. 1968. E c o l o g i c a l a d a p t a t i o n s f o r b r e e d i n g i n b i r d s . Methuen. London. Leopold, A.S. 1944. The nature of h e r i t a b l e w i l d n e s s i n t u r k e y s . Condor 64:133-197. Leopold, A.S., M. Erwin, J . Oh and B. Browning. 1976. Phytoestrogens: adverse e f f e c t s on r e p r o d u c t i o n i n C a l i f o r n i a q u a i l . S c i e n c e 191:98-99. L i l l , A. 1968. S p a t i a l o r g a n i z a t i o n i n s m a l l f l o c k s o f domestic f o w l . Behaviour 32:258-290. Long, J . 1981. Introduced B i r d s of the World. U n i v e r s e Books, N. Y. Lorenz, K. 1940. Durch Domestikation v e r u r s a c h t e Storungen a r t e i g e n e n V e r h a l t e n s . Z e i t s c h r . angewandte P s y c h o l , und Charakter. 59:2-82. Lorenz, K. 1965. E v o l u t i o n and M o d i f i c a t i o n of Behavior. Univ. o f Chicago Press, Chicago. M a r t i n , K. 1984. Reproductive defense p r i o r i t i e s o f male w i l l o w ptarmigans (Lagopus i a g o p u s ) : enhancing mate s u r v i v a l o r exte n d i n g p a t e r n i t y o p t i o n s ? Behav. E c o l . S o c i o b i o l . 16:57-63. M a r t i n , K., S.J. Hannon and S. Lord. 1990. Female-female a g g r e s s i o n i n W h i t e - t a i l e d Ptarmigan and Willow Ptarmigan d u r i n g the p r e - i n c u b a t i o n p e r i o d . Wilson B u l l . 102:532-536. McAtee, W.L. 1944. Game b i r d s s u i t a b l e f o r n a t u r a l i z i n g i n the U n i t e d S t a t e s . U.S.D.A. C i r c . No. 96. McBride, G., I.P. P a r e r and F. Foenander. 1969. The s o c i a l o r g a n i z a t i o n and behaviour o f the f e r a l domestic f o w l . Anim. Behav. Monog. 2:127-181. McGrew, D. 1958. A l l about C o t u r n i x . Outdoor Nebraska 36:17-19. McKinney, F. 1985. Primary and secondary male r e p r o d u c t i v e s t r a t e g i e s o f d a b b l i n g ducks. O r n i t h o l . Monogr. 37:57-67. McKinney, F., S.R. D e r r i c k s o n and P. Mineau. 1983. Forced c o p u l a t i o n i n waterfowl. Behaviour 86:250-294. McKinney, F. K.M. Cheng and D.J. Bruggers. 1984. Sperm c o m p e t i t i o n i n a p p a r e n t l y monogamous b i r d s . In: Sperm c o m p e t i t i o n and the e v o l u t i o n o f animal mating systems. (R.L. Smith, ed). Academic Press, Orlando, F l a . pp523-545. Meise, W. 1954. Uber Zucht, E i n t r i t t der G e s c h l e c t s r e i f e , Zwischenund Weiterzug der Wachtel (C. c o t u r n i x ) . Vogelwarte 17:211-215. Menzdorf, A. 1982. S o c i a l behaviour o f rock p a r t r i d g e s ( A l e c t o r i s g r a e c a ) . W i l d Pheasant Assoc. J . 7:70-89. Mock, D.W. and M. F u j i o k a . 1990. Monogamy and long-term p a i r bonding i n v e r t e b r a t e s . Tree 5:39-43. Moreau, R.E. 1951. The B r i t i s h s t a t u s of the q u a i l and some problems o f i t s b i o l o g y . B r i t i s h B i r d s 44:257-276. 104 Munro, G.C. 1960. B i r d s o f Hawaii. C h a r l e s E. T u t t l e , Rutland, Vermont. O r c u t t , F. and A. O r c u t t . 1976. N e s t i n g and P a r e n t a l Behavior i n domestic common q u a i l . Auk 93:135-141. O t t i n g e r , M.A., W.M. S c h l e i d t and E. Russek. 1982. D a i l y p a t t e r n s o f c o u r t s h i p and mating be h a v i o r i n the male Japanese q u a i l . Behav. Proc. 7:223-233. Paludan, K. 1963. P a r t r i d g e markings i n Denmark. Dan. Rev. Game B i o l . 4:25-60. Parker, H. 1981. Renesting b i o l o g y o f Norwegian w i l l o w ptarmigan. J . Wi l d . Manage. 45:858-864. Parker, H. 1985. Compensatory r e p r o d u c t i o n through r e n e s t i n g i n w i l l o w ptarmigan. J . W i l d l . Manage. 49:599-604. P e r r i n s , CM. 1970. The t i m i n g o f b i r d s * b r e e d i n g seasons. I b i s 112:242-255. Peterson, R.T. 1961. A f i e l d guide t o western b i r d s . Houghton M i f f l i n Co., Boston. P f e i f e r , S. 1954. Vergewaltigung e i n e s v e r l e t z e n Wachtelweibchens durch e i n Wachtelmannchen ( C o t u r n i x c o t u r n i x ) . O r n i t h . M i t t . 6:174-175. P h i l l i p s , J.C. 1928. W i l d b i r d s i n t r o d u c e d o r t r a n s p l a n t e d i n North America. U.S.D.A. Tech. B u l l . 61. P h i l l i p s , R.E. 1964. "Wildness" i n the m a l l a r d duck: e f f e c t s o f b r a i n l e s i o n s and s t i m u l a t i o n on "escape behaviour" and r e p r o d u c t i o n . J . Comp. Neurol. 122:139-155. P i a t t , D.R. 1968. Nest p a r a s i t i s m by the bobwhite. Kans. O r n i t h . Soc. B u l l . 19:18. Potash, L.M. 1975. An experimental a n a l y s i s o f the use o f l o c a t i o n c a l l s by Japanese q u a i l ( C o t u r n i x c o t u r n i x i a p o n i c a ) . Behaviour 54:153-179. P o t t s , G.R. 1973. F a c t o r s governing the c h i c k s u r v i v a l r a t e o f the grey p a r t r i d g e . Proceedings o f the 10th I n t e r n a t i o n a l Congress on Game B i o l o g y , pp85-96. P o t t s , G.R. 1986. The p a r t r i d g e - p e s t i c i d e s , p r e d a t i o n and c o n s e r v a t i o n . C o l l i n s P r o f , and Tech. Books, London, U.K. Presto n , J.R. 1961. The beha v i o r i n l a b o r a t o r y C o t u r n i x c o t u r n i x j a p o n i c a Temminck and S c h l e g e l , from h a t c h i n g t o the 105 appearance o f a s o c i a l h i e r a r c h y . M.Sc. t h e s i s , Univ. o f Arkansas. P r i c e , E.O. 1984. B e h a v i o r a l a s p e c t s of Animal d o m e s t i c a t i o n . Quart. Rev. B i o l . 59:1-32. Ramenofsky, M. 1984. A g o n i s t i c behaviour and endogenous plasma hormones i n male Japanese q u a i l . Anim. Behav. 32:698-708. Rands, M. and P. Hayward. 1987. S u r v i v a l and c h i c k p r o d u c t i o n o f hand-reared gray p a r t r i d g e s i n the w i l d . W i l d l . Soc. B u l l . 15:456-457. R a v e l i n g , D.C. 1979. The annual c y c l e of body composition of Canada Geese w i t h s p e c i a l r e f e r e n c e t o c o n t r o l o f r e p r o d u c t i o n . Auk 96:234-252. R i c k l e f s , R.E. 1980. Geo g r a p h i c a l v a r i a t i o n i n c l u t c h s i z e among p a s s e r i n e b i r d s : Ashmole's h y p o t h e s i s . Auk 97:38-49. Robinson, C.A., C R . N i c h o l s and K.M. Cheng. 1991. The use o f a throw-net t o capture w i l d Japanese q u a i l . G i b i e r Faune Sauvage ( i n p r e s s ) . Romanoff, A.L. 1960. The a v i a n embryo. S t r u c t u r a l and F u n c t i o n a l Development. Macmillan, New York, N.Y. Roseberry, J.L., D.L. E l l s w o r t h and W.D. K l i m s t r a . 1987. Comparative p o s t - r e l e a s e b e h a v i o r and s u r v i v a l o f w i l d , semi-w i l d , and game farm bobwhites. W i l d l . Soc. B u l l . 15: 449-455. R o t h s t e i n , R. 1967. Some o b s e r v a t i o n s on the n e s t i n g b e h a v i o r o f Japanese q u a i l ( C o t u r n i x c o t u r n i x j a p o n i c a ) i n pseudonatural c o n d i t i o n s . P o u l t r y S c i . 46:260-262. Sachs, B.D. 1966. S e x u a l - a g g r e s s i v e i n t e r a c t i o n s among p a i r s of q u a i l ( C o t u r n i x c o t u r n i x j a p o n i c a ) . Am. Z o o l . 6:559. S c h l a d w e i l e r , J.L. and J.R. T e s t e r . 1972. S u r v i v a l and beh a v i o r o f hand-reared m a l l a r d s r e l e a s e d i n the w i l d . J . W i l d l . Manage. 36:1118-1127. Schmidt, W. 1976. Q u a l i t a t i v e und q u a n t i t a t i v e Untersuchungen am V e r h a l t e n von Haus-und Graugansen T h e s i s , D u s s e l d o r f . Schwarz, CW. and E.R. Schwartz. 1949. A Reconnaissance o f the Game B i r d s i n Hawaii. H i l o , Hawaii: Hawaii Board o f Commissioners of A g r i c u l t u r e and F o r e s t r y . S e l i n g e r , H. and G. Bermant. 1967. Hormonal c o n t r o l o f a g g r e s s i v e b e h a v i o r i n Japanese q u a i l ( C o t u r n i x c o t u r n i x 106 i a p o n i c a ) . Behaviour 28:255-268. S i e g e l , P.B. 1976. S o c i a l b e h a v i o r o f the fo w l . P o u l t . S c i . 55:5-13. S i e g f r i e d , W.R. 1976. Breeding b i o l o g y and p a r a s i t i s m i n the ruddy duck. Wilson B u l l . 88:566-574. Snedecor, G.W. and W.G. Cochran. 1980. S t a t i s t i c a l Methods. Iowa S t a t e U n i v e r s i t y P ress, Ames, Iowa. So s s i n k a , R. 1982. Domestication i n b i r d s . In "Avian B i o l o g y " . (Farner, D.S., J.R. King and K.C. Parkes, e d s ) , V o l . 6, pp373-403. S t a d i e , C. 1969. V e r g l e i c h e n d e Beobachtungen an Ver h a l t e n s w e i s e n v e r s c h i e d e n e r Wildhuhnarten der Gattung. G a l l u s . Z o o l . Anz. 183:13-30. S t a n f o r d , J.A. 1957. A p r o g r e s s r e p o r t o f C o t u r n i x q u a i l i n v e s t i g a t i o n s i n M i s s o u r i . Twenty-Second North American W i l d l i f e Conference, pp316-359. Steen, J.B. and H. Parker. 1981. The egg numerostat-a new concept i n the r e g u l a t i o n o f c l u t c h s i z e . O r n i s Scand. 12:109-110. Stevens, V.C. 1961. Experimental study o f n e s t i n g by C o t u r n i x q u a i l . J . W i l d l . Mgmt. 25:99-101. Vander Haegen, M., W. Dodge and M. Sayre. 1988. F a c t o r s a f f e c t i n g p r o d u c t i v i t y i n a n o r t h e r n w i l d t u r k e y p o p u l a t i o n . J . W i l d l . Manag. 52:127-133. V a n g i l d e r , L.D., E.W. K u r z e j e s k i , V.L. Kimmel-Truitt and J.B. * Lewis. 1987. Reproductive parameters o f w i l d t u r k e y hens i n North M i s s o u r i . J . W i l d l . Manage. 51::535-540. Wada, M. 1986. C i r c a d i a n rhythms o f testosterone-dependent b e h a v i o r s , crowing and locomotor a c t i v i t y , i n male Japanese q u a i l . J . Comp. P h y s i o l . A 158:17-25. Wakasugi, N. 1984. Japanese q u a i l . In: E v o l u t i o n o f domesticated animals. ( I . L . Mason, ed). Longman Inc., New York. Weigand, J.P. 1980. Ecology of the Hungarian p a r t r i d g e i n n o r t h - c e n t r a l Montana. W i l d l . Monogr. No. 74. Westemeier, R.L. and T.L. Esker. 1989. An u n s u c c e s s f u l c l u t c h of Northern Bobwhites w i t h hatched pheasant eggs. Wilson B u l l . 101:640-642. Wetherbee, D.K. 1961. I n v e s t i g a t i o n s i n the l i f e h i s t o r y o f the Common C o t u r n i x . Am. M i d i . Nat. 65:168-86. Wiley, R.H. 1973. T e r r i t o r i a l i t y and non-random mating i n sage grouse C e n t r o c e r c u s urophasianus. Anim. Behav. Mongr. 6:85-169. Wilson, E.O. 1975. S o c i o b i o l o g y : the new s y n t h e s i s . Cambridge, Massachusetts, Belknap P r e s s . Wilson, W.O., Abbott, U.K. and Abplanalp, H. 1961. E v a l u a t i o n of C o t u r n i x (Japanese q u a i l ) as p i l o t animal f o r p o u l t r y . P o u l t r y S c i . 40:651-657. Wilson, M.I. and Bermant, G. 1972. An a n a l y s i s o f s o c i a l i n t e r a c t i o n s i n Japanese q u a i l , C o t u r n i x c o t u r n i x j a p o n i c a . Anim. Behav. 20:252-258. Winkler, D.W. and J.R. Walters. 1983. The d e t e r m i n a t i o n o f c l u t c h s i z e i n p r e c o c i a l b i r d s . In: Current o r n i t h o l o g y . V o l . 1 (R.F. Jownston ed.). Plenum Press, New York. pp33-68. Wittenberger, J.F. 1978. The e v o l u t i o n o f mating systems i n grouse. Condor 80:126-137. Wittenberger, J.F. 1979. The e v o l u t i o n o f mating systems i n b i r d s and mammals. In "Handbook o f B e h a v i o r a l Neurobiology", V o l . 3 : S o c i a l Behavior and Communication (P. M a r l e r and J . Vandenbergh, e d s . ) . Plenum, New York. pp271-349. Wittenberger, J.F. and R.L. T i l s o n . 1980. The e v o l u t i o n o f monogamy: Hypotheses and evidence. Annu. Rev. E c o l . S y s t . 11:197-232. Yamashina, Y. 1961. Q u a i l b r e e d i n g i n Japan. J . Bombay Nat. H i s t . Soc. 58:216-222. Yom-Tov. Y., and R. H i l b o r n . 1981. E n e r g e t i c c o n s t r a i n t s on c l u t c h s i z e and time o f br e e d i n g i n temperate zone b i r d s . O e c o l o g i a 48:234-243. time p a t I n i t i a t o r j 0 r e a c t o r 5 % c r o u c h S t r u t t i n g F o l l o w i n a B i l l on Nape M o u n t i n g T a i l b e n d i n g C l o a c a l C-T i t - b i d d i n g P e c k i n g C r o w i n g J umo I,»fti hv ft . w i n g f l a D D e d R u n n i n g P a c i n g Chasing F e e d i n c S l e e p i n g P r e e n i n g R e l a x e d -P e c k i n o Gr. to •-3 D M a. o o 01 > CD a t—-X 0 Q *• Q in & CD CD •—* cn 1—fc o »-l CD O O a »—•• CQ cr CD D* Q < o d Q to n »—* G *< to 80T Appendix 2. Grid maps lor recording bird locations. 110 Appendix 3 . O b s e r v a t i o n s o f the d a n c i n g d i s p l a y (DN) i n Japanese q u a i l . Y e a r Genotype Obs 1 1987 f e r a l 1 2 2 2 2 1 domestic 1 1988 f e r a l domestic 1 Context* FY1 WF 3-DN FJTB1/FYP1; FYP1 WF F Y 1 ; FYB1 ST F Y 1 ; FY1 WF FYB1 . FYG1 CF-DN-ST FG1. FPB1 AP4-CW-DN-BN-MO-DN F B I . FYG1 AP-DN F G 1 / F Y B 1 ; FYB1 CH FYG1. FYG1 CF FG1; FG1 AP-CF F Y B 1 / F Y 1 ; YG1 DN YG1. FPB1 AP-DN-CW F B I . FG1 DN F Y 1 . DY1 CR DYB1; DYB1 DN-BN-MO-TB-CC DY1. FYB2 TT-ST FY2; FY2 CF FYB2; FYB2 BN-MO-TB-CC F Y 2 ; FY2 AP-CF FYP2; FYP2 DN F Y 2 . FYB2 BN-MO-TB-CC-TT-BN-TT-BN-TT F Y 2 ; FY2 DN-CR FYB2; FYB2 BN, MO, TB, CC, F Y 2 . FYB2 CF F P 2 / F Y P 2 ; FYP2 PK FYB2; FYB2 BN-MO FYP2; FYP2 DN-CH FYB2. DPB2 MO-TB-CC DY2; DYG2 CH DPB2; DPB2 CW DY2 DN. DYG2 MO-TB DB2; DY2 DN DYG2 1 Observer 2 S i t u a t i o n under which the d i s p l a y was observed. See methods s e c t i o n f o r a b b r e v i a t i o n s o f v a r i o u s d i s p l a y s . U n d e r l i n i n g i n d i c a t e s the members of a p a i r (see Chapter 4) . e.g. L i n e one should read: FY1 wing-flapped and danced around her mate FYB1 and another male FYP1; FYP1 wing-flapped f a c i n g FY1; FYB1 s t r u t t e d t o FY1; FY1 wing-flapped f a c i n g FYB1. 3 W i n g - f l a p p i n g 4 Approaching Appendix 4 . Male - Female A s s o c i a t i o n s and Female - Male A s s o c i a t i o n s . F E M A L E FERAL MALE - FYB1 - 1987 Male - Female Associat ion (Total Score / Observation period) F B 1 3 F E M A L E FERAL MALE - FYP1 - 1987 Male - Female Associat ion (Total Score / Observation period) A . A A 20 Jun 2 Jun 15 Jun 28 I PAIR BOND (see Chapter 4) Ju l 11 Jul 24 Aug 6 Male Removed Aug 19 Aug 30 EGG CYCLE | H LAY I N C U B A T E B R O O D F E M A L E FERAL MALE - FYG1 - 1987 Male - Female Associat ion (Total Score / Observat ion period) FB1 3 2 1 0 FG1 3 2 1 0 FY1 3 2 1 0 FP1 3 2 1 May 20 Jun 2 Jun 15 Jun 28 I PAIR BOND (see Chapter 4) Jul 11 Ju l 24 Aug 6 Aug 19 EGG CYCLE H Aug 30 LAY INCUBATE BROOD F E M A L E FERAL MALE - FPB1 - 1987 Male - Female Associat ion (Total S c o r e / Observat ion per iod) A A 20 Jun 2 Jun 15 Jun 28 I PAIR BOND (see Chapter 4) Ju l 11 Ju l 24 Aug 6 Aug 19 Aug 30 EGG r v r , c ujgid INCUBATE H I BROOD LAY F E M A L E FERAL MALE - FYB2 - 1988 Male - Female Associat ion (Total Score / Observat ion per iod) FB2 3 2 1 0 FG2 3 A A A A F Y 2 3 FP2 3 1 May 2 May 17 Jun 1 Jun 16 • PAIR B O N D (see Chapter 4) Ju l 1 Ju l 16 J u l 31 Aug 15 EGG Aug 30 C Y C L E ^ I N C U B A T E O Y U L t Hi BROOD LAY F E M A L E FERAL MALE - FYP2 - 1988 Male - Female Associat ion (Total Score / Observat ion per iod) /A A A A A A A A A A A A A A A A May 2 May 17 Jun 1 Jun 16 I PAIR BOND (see Chapter 4) Aug 30 r v n C W>& INCUBATE 11 BROOD FEMALE FERAL MALE - FPB2 - 1988 Male - Female Association (Total S c o r e / Observat ion period) ^ m , ^ ^ ^ ^ ^ ^ h KrT\ A A A A . M ^ ^ ^ ^ ^ ^ ^ I Al -iik A luwliJMiiliiJl -H H I p . , pa, A MHHl A l\ f\K AAA A A FB2 3 2 1 0 FG2 3 2 1 0 FY2 3 2 1 0 FP2 3 2 1 0 May 2 May 17 Jun 1 Jun 16 • PAIR BOND (see Chapter 4) Ju l 1 Ju l 16 Male Removed Ju l 31 Aug 15 EGG CYCLE Aug 30 LAY INCUBATE Hi BROOD MALE F Y B 1 3 FERAL FEMALE - FP1 - 1987 Female - Male Associat ion (Total S c o r e / Observat ion per iod) May 20 Jun 2 Jun 15 Jun 28 • PAIR BOND (see Chapter 4) Jul 11 Ju l 24 Aug 6 Aug 19 Aug 30 EGG m LAY r v r i H | i INCUBATE O Y O L t HH BROOD MALE FERAL FEMALE - FY1 - 1987 Female - Male Assoc iat ion (Total Score / Observat ion period) F Y B 1 3 2 1 0 F Y P 1 3 F Y G 1 3 F P B 1 3 2 1 A . May 20 Jun 2 Jun 15 Jun 28 • PAIR B O N D (see Chapter 4) Jul 11 Ju l 24 Aug 6 Aug 19 EGG CYCLE ^ Aug 30 LAY INCUBATE BROOD M A L E FERAL FEMALE - FG1 - 1987 Female - Male Associat ion (Total S c o r e / Observat ion per iod) A A May 20 Jun 2 Jun 15 Jun 28 • PAIR BOND (see Chapter 4) Jul 11 J u l 24 Aug 6 Aug 19 Aug 30 E G G C Y f L E ^ I N C U B A T E HI BROOD LAY MALE FERAL FEMALE - FB1 - 1987 Female - Male Associat ion (Total S c o r e / Observat ion per iod) May 20 Jun 2 Jun 15 • PAIR BOND (see Chapter 4) Jun 28 Jul 11 J u l 24 Aug 6 Aug 19 Aug 30 EGG m LAY <~WM c IP INCUBATE HI BROOD MALE FERAL FEMALE - FP2 - 1988 Female - Male Associat ion (Total S c o r e / Observat ion period) A AA A A May 2 May 17 Jun 1 Jun 16 • PAIR BOND (see Chapter 4) Ju l 1 Ju l 16 Ju l 31 Aug 15 EGG Aug 30 C Y C L E ^ I N C U B A T E O Y O L b m BROOD LAY MALE FYB2 3 FYP2 3 2 FYG2 3 2 1 0 FPB2 3 2 1 A A FERAL FEMALE - FY2 - 1988 Female - Male Associat ion (Total S c o r e / Observat ion per iod) A A A A A A A A A A May 2 May 17 Jun 1 Jun 16 • PAIR BOND (see Chapter 4) Ju l 1 Ju l 16 Ju l 31 Aug 15 EGG Aug 30 C Y C L E ^ I N C U B A T E O Y O L b m BROOD LAY MALE FERAL FEMALE - FG2 - 1988 Female - Male Associat ion (Total S c o r e / Observat ion period) A A A A A A A A A A A A May 17 Jun 1 Jun 16 PAIR BOND (see Chapter 4) Ju l 1 Ju l 16 J u l 31 Aug 15 EGG CYCLE Aug 30 LAY INCUBATE HI BROOD MALE FYB2 3 2 1 0 FYP2 3 FERAL FEMALE - FB2 - 1988 Female - Male Associat ion (Total Score / Observation period) o A, FYG2 3 2 1 FPB2 3 1 / \ A A A A A A A A A , A May 2 May 17 Jun 1 Jun 16 • PAIR BOND (see Chapter 4) Jul 1 Jul 16 Jul 31 Aug 15 EGG Aug 30 r v r i P ^ I N C U B A T E O Y O L t | H B R O O D L A Y DOMESTIC MALE - DYB1 - 1987 Male - Female Associat ion F E M A L E (Total Score / Observation period) F E M A L E DOMESTIC MALE - DYP1 - 1987 Male - Female Associat ion (Total Score / Observation period) DB1 3 2 1 0 DG1 3 2 1 0 DY1 3 2 1 0 DP1 3 2 1 A A A A A May 20 Jun 2 Jun 15 Jun 28 • PAIR BOND (see Chapter 4) Jul 11 Jul 24 Aug 6 Male Removed Aug 19 Aug 30 EGG r v r , c ^ INCUBATE U Y O L b H| BROOD LAY F E M A L E DOMESTIC MALE - DYG1 - 1987 Male - Female Associat ion (Total S c o r e / Observat ion period) A A A . A AA A A A A A May 20 Jun 2 Jun 15 • PAIR BOND (see Chapter 4) Jun 28 Jul 11 J u l 24 Aug 6 Aug 19 Aug 30 EGG CYC I F ^ INCUBATE HI BROOD LAY F E M A L E DOMESTIC MALE - DPB1 - 1987 Male - Female Associat ion (Total S c o r e / Observat ion period) May 20 Jun 2 Jun 15 Jun 28 • PAIR BOND (see Chapter 4) Jul 11 Ju l 24 Aug 6 Aug 19 EGG CYCLE Aug 30 LAY INCUBATE Hi BROOD F E M A L E DB2 3 DG2 DP2 DOMESTIC MALE - DYB2 - 1988 Male - Female Associat ion (Total S c o r e / Observat ion per iod) May 2 May 17 Jun 1 Jun 16 • PAIR B O N D ( s e e C h a p t e r 4) Jul 1 J u l 16 Ju l 31 Aug 15 EGG Aug 30 C Y C L E ^ I N C U B A T E H B R O O D LAY DOMESTIC MALE - DYP2 - 1988 Male - Female Associat ion F E M A L E (Total Score / Observation period) F E M A L E D B 2 3 2 1 0 D G 2 3 2 1 0 D Y 2 3 DOMESTIC MALE - DPB2 - 1988 Male - Female Associat ion (Total Score / Observat ion per iod) A A A _ D P 2 3 1 A A A A M A May 2 May 17 Jun 1 Jun 16 I PAIR BOND (see Chapter 4 ) Ju l 1 Ju l 16 Male Removed Ju l 31 Aug 15 EGG 1 Aug 30 r v n P fesS INCUBATE O Y O L t m BROOD LAY MALE DYB1 3 DYP1 3 2 1 0 DYG1 3 2 1 0 DPB1 3 2 1 A DOMESTIC FEMALE - DP1 - 1987 Female - Male Associat ion (Total Score / Observation period) A A A A A A A A A. A May 20 Jun 2 Jun 15 Jun 28 • PAIR BOND (see Chapter 4) Jul 11 Jul 24 Aug 6 Aug 19 EGG CYCLE Aug 30 LAY INCUBATE HI BROOD MALE DOMESTIC FEMALE - DY1 - 1987 Female - Male Associat ion (Total Score / Observat ion per iod) May 20 Jun 2 Jun 15 Jun 28 • PAIR BOND (see Chapter 4) Jul 11 Ju l 24 Aug 6 Aug 19 Aug 30 EGG r v r , c uvjHi IN CU BAT E HH BROOD LAY M A L E DOMESTIC FEMALE - DG1 - 1987 Female - Male Assoc ia t ion (Total Score / Observation period) A . A A A A A A A A A May 20 Jun 2 Jun 15 • PAIR BOND (see Chapter 4) Jun 28 Jul 11 Jul 24 Aug 6 Aug 19 Aug 30 E G G C Y C L E INCUBATE HI BROOD LAY MALE D P B 1 3 DOMESTIC FEMALE - DB1 - 1987 Female - Male Associat ion (Total Score / Observat ion per iod) May 20 Jun 2 Jun 15 Jun 28 • PAIR BOND (see Chapter 4) Jul 11 Ju l 24 Aug 6 Aug 19 Aug 30 EGG LAY C Y C L E ^ ' N C U B A T E O Y O L t |H BROOD MALE DOMESTIC FEMALE - DP2 - 1988 Female - Male Associat ion (Total Score / Observat ion period) A A. A- A A..A May 2 May 17 Jun 1 Jun 16 • PAIR BOND (see Chapter 4) Jul 1 Ju l 16 Ju l 31 Aug 15 EGG Aug 30 r v r i P SSS INCUBATE CYCLE HH BROOD LAY DOMESTIC FEMALE - DY2 - 1988 Female - Male Associat ion MALE (Total Score / Observat ion per iod) May 2 May 17 Jun 1 Jun 16 Ju l 1 Ju l 16 Ju l 31 Aug 15 Aug 30 • PAIR BOND (see Chapter 4) MALE DOMESTIC FEMALE - DG2 - 1988 Female - Male Assoc iat ion (Total Score / Observat ion per iod) A A A . A A , A May 2 May 17 Jun 1 Jun 16 • PAIR BOND (see Chapter 4) Jul 1 Ju l 16 Ju l 31 Aug 15 EGG Aug 30 r v r i n issaa INCUBATE HH BROOD LAY MALE DOMESTIC FEMALE - DB2 - 1988 Female - Male Associat ion (Total Score / Observat ion period) May 2 May 17 Jun 1 Jun 16 • PAIR BOND (see Chapter 4) Ju l 1 Ju l 16 Jul 31 Aug 15 EGG Aug 30 LAY INCUBATE ILL i 1 s 3 1 8 7 8 5 6 ! B o r r o w e r s IUL Re c o r d 1 o f 1 20001£01 sReqDate: £ 0 0 0 1 0 2 3 : N e e d B e f o r e : :RecDate: :RenewalReq: s DueDat e : £ 0 0 0 1 1 £ 6 : NewDueDat e s : I L L : ;3187856 : S t a t us: SHIPPED :OCLC: £ 8 0 © 7 4 £ £ s S o u r c e : C l i o -.Lender: UBC, *UBC, UBC sAUTHOR: N i c h o l s , C a t h l e e n Rose, 1958-: T I T L E : A c o m p a r i s o n o f t h e r e p r o d u c t i v e and b e h a v i o u r a l d i f f e r e n c e s i n f e r a l and d o m e s t i c J a p a n e s e q u a i l / SEDITION: M a s t e r ' s t h e s i s , 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 , 1991 :IMPRINT s 1991. :DISSERTATION: T h e s i s (M. 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 , 1991. s VERIFIED s OCLC s PATRON: ( B r i n d s t a f f , J e n n i f e r / J o r d a n Hall Room 1 4 £ / 3 £ 1 6 4 7 6 0 7 / j g r i n d s t sSHIP TO: DDS/1320 E. 10TH ST. RM E065/IND.U.LIB/BLOOMINGTON IN USA 47405 :BILL TO s SAME CIC sSHIP VIA: 1 2 9 . 7 9 . 3 2 . 1 0 0 / A r i e l :MAXCOST: 25IFM :COPYRT COMPLIANCE: sFAX: <812)855-3701 :E-MAIL: l i b i l l @ i n d i a n a . e d u sBORROWING NOTES: C l i o I D s Z 0 0 1 7 5 7 9 / / / / T h i s E d i t i o n O n l y : yes S o u r c e o f C i t a t i o n s R e f s o f p a p e r sPATRON IDs sPDEPT: B i o l o g y sPSTATUSs Grad :PATRON PHONE: 855-1096 :PATRON E-MAIL: sPATRON FAX: :LENDING CHARGES: *16 i f m :SHIPPED: £ 0 0 0 1 0 2 7 :SHIP INSURANCE: :RETURN TO s RSS-KOERNER LIBRARY/UBC, P.O. BOX £139, VAN. BC, CANADA, V6B 3T1 :RETURN VIA: BOOK RATE sRETURNED VIA: :RETURNED: sINSURANCE: 

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}]}"
                            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:
http://iiif.library.ubc.ca/presentation/dsp.831.1-0098657/manifest

Comment

Related Items