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Studies of the holding, behaviour and nutrition of captive blue grouse Stirling, Ian Grote 1965

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STUDIES OF THE HOLDING, BEHAVIOUR AND NUTRITION OF CAPTIVE BLUE GROUSE by IAN GROTE STIRLING B.Sc., University of B r i t i s h Columbia, 1963 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department of Zoology We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA November, 1965 In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f th e r e q u i r e m e n t s f o r an advanced degree a t t h e 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 , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r a g r e e t h a t p e r -m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by the Head o f my Department o r by h i s r e p r e s e n t a t i v e s , , I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i -c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Department o f The U n i v e r s i t y o f B r i t i s h Columbia Vancouver 8, Canada Date D*?<? S'Z&i''  i ABSTRACT The primary purpose of keeping blue grouse was to l e a r n to keep them i n good h e a l t h and breed them. New data were obtained on the weights, n u t r i t i o n , s u r v i v a l , d i s e a s e s f a n d behaviour of blue grouse i n c a p t i v i t y . They were kept i n 3 s i z e s of pens, 2 f e e t wide by 4 f e e t high by 8 f e e t l o n g , 6 f e e t wide by 4 f e e t h i g h by 8 f e e t long> and 20 f e e t wide by 10 f e e t high by 20 f e e t l o n g . On the b a s i s of plumage c o n d i t i o n and r e p r o d u c t i v e behaviour the medium s i z e d pens were b e s t . The l e v e l of p r o t e i n of the d i e t of the hens appeared to a f f e c t s u r v i v a l and r e p r o d u c t i v e behaviour. Grouse that were f e d a d i e t w i t h IS percent p r o t e i n had b e t t e r s u r v i v a l and ex-h i b i t e d more r e p r o d u c t i v e behaviour than grouse f e d a d i e t w i t h 24 or 28 percent p r o t e i n . Grouse were unable to s u r v i v e on a d i e t of d r i e d and p e l l e t e d Douglas f i r needles. None of the c o n c l u s i o n s reached appear a p p l i c a b l e t o c o n d i t i o n s observed i n the f i e l d . Hens e x h i b i t i n g r e p r o d u c t i v e behaviour were s e x u a l l y i m p r i n t e d upon humans. The only s u c c e s s f u l method of mating grouse was t o take a male e x h i b i t i n g s e x u a l d i s p l a y t o a squat-t i n g female. A r t i f i c i a l i n s e m i n a t i o n was as s u c c e s s f u l as n a t u r a l matings i n the a v i a r y . The apparent d i g e s t i b i l i t y of the commercial chicken breeder r a t i o n v a r i e d from 51.2 t o 64.9 percent. There was a relationship between the daily consumption of water and body weight. The study of behaviour indicated there was a relation-ship between the hooting of males and the squatting and egg laying of females. Adult males hooted more than yearlings. • The female usually gave a pre-copulatory cry when ready to mate Males became more aggressive during the breeding season and less aggressive through the summer. The females appeared to have two peaks of aggressive behaviour during the reproductive season. Aggressive behaviour of females may serve to space them in the f i e l d during the period of nesting. There may be a period of aggressive behaviour in males during the winter. A partial catalogue of grouse behaviour was made. Photo graphs of postures and sonographs of calls supplement descrip-tion. i i i TABLE OF CONTENTS Page ABSTRACT i TABLE OF CONTENTS . i i LIST OF TABLES v LIST OF FIGURES v i i LIST OF PLATES v i i i ACKNOWLEDGEMENTS x PART 1. INTRODUCTION Chapter 1. I n t r o d u c t i o n 1 PART 2. HOLDING AND BREEDING Chapter 2. Holding Blue Grouse i n C a p t i v i t y . . . 3 Chapter 3- M o r t a l i t y of Blue Grouse i n C a p t i v i t y . 16 Chapter 4« Breeding Blue Grouse i n C a p t i v i t y . . 25 Chapter 5 . A r t i f i c i a l Insemination of Blue Grouse . 35 Chapter 6. Incubation and Brooding 41 PART 3- NUTRITION Chapter 1. Experiments on D i e t s and V i a b i l i t y . . 44 Chapter 8. Ba s i c N u t r i t i o n 54 PART 4- BEHAVIOUR Chapter 9 . Reproductive Behaviour 61 Chapter 10. Aggression and the Reproductive Cycle . 73 Chapter 11. Winter Aggression 84 PART 5- SUMMARY Chapter 12. Summary 89 LIST OF REFERENCES APPENDIX 1 . . APPENDIX 2 APPENDIX 3 V LIST OF TABLES TABLE Page 1. Sizes and number of pens i n U.B.C. aviary . . . 5 2. Survival of grouse i n each size of pen from December to June of 1963-4 and 1964-5 . . . . 9 3 . Numbers, age, and sex of grouse held i n each size of pen and the proportion that showed reproductive display 12 4. Numbers, sex,and s p e c i f i c l o c a l i t y of capture of each cohort received i n the U.B.C. aviary each August 17 5. Causes of mortality of blue grouse of each age class at the U.B.C. aviary from August 1962 through 'August 1965 19 6. Number of blue grouse at U.B.C. aviary showing reproductive behaviour from 1963 through 1965 . 27 7. Results of natural and forced mating attempts i n U.B.C. aviary 29 8. F e r t i l i t y and hat c h a b i l i t y of eggs l a i d at the U.B.C. aviary 30 9. Comparative f e r t i l i t y of blue grouse eggs . . . 3 3 10. The effect of handling hens f o r a r t i f i c i a l i n -semination, upon squatting and egg laying . . . 3 7 11. Comparison of mean clutch s i z e of blue grouse on Vancouver Island to blue grouse that were naturally mated and a r t i f i c i a l l y inseminated in the U.B.C. aviary 3$ 12. Number of hen blue grouse i n U.B.C. aviary that showed reproductive behaviour and laying on each rat i o n 48 13. Survival of hen blue grouse held on each rat i o n at U.B.C. aviary from September, 1963 through December, 1964 50 v i TABLE Page 14. C a l o r i c v a l u e s and r a t i o o f c a l o r i e s t o p e r c e n t p r o t e i n of r a t i o n s f e d t o grouse i n t h e U.B.C. a v i a r y f rom September, 1963 t h r o u g h December, 1964 52 15. Body w e i g h t s , d i g e s t i b i l i t y and c a l o r i c and wate r r e q u i r e m e n t s o f b l u e g r o u s e h e l d a t U.B.C. v i v a r i u m . 56 l6o Weights o f g r o u s e , t h e i r d a i l y c a l o r i c i n t a k e and t h e i r a pproximate maintenance energy r e q u i r e m e n t 59 17. P e r c e n t o f t o t a l o b s e r v a t i o n s on a d u l t and y e a r l i n g male b l u e g r o u s e upon wh i c h males h o o t e d , May 1 t h r o u g h June 20, 1963-65 64 v i i LIST OF FIGURES FIGURE Page 1. Plan view of the aviary 6 2. Comparison of body weights of blue grouse in the f i e l d and in the aviary 11 3. Deaths and their cause by age and number . . 21 4. Survivorship of each age-class of chicks brought to U.B.C. aviary 22 5. The relationship between the occurrence of male hooting to female squatting and egg laying 65 6. Indicies of female activity during the breeding season 78 7. Indicies of male activity during the breeding season 80 v i i i LIST OF PLATES PLATE Page 1. U.B.C. a v i a r y from s o u t h 117 2. I n s i d e o f a pen a t U.B.C. a v i a r y t o show p o s i -t i o n o f water f a u n t s , f e e d e r s , d i v i d e r s and r o o s t 117 3« E l e c t r i c w e i g h i n g s c a l e w i t h basket . . . . 117 4. Humidaire i n c u b a t o r 118 5. B a t t e r y i n c u b a t o r s . 118 6. Brooder a t U.B.C. a v i a r y 118 7. P o s t u r e o f a s i c k g r ouse 118 8. Normal s t a n c e o f a b l u e g rouse 119 9. Head t u r n p o s t u r e 119 10. N e c k - s t r e t c h p o s t u r e , v e r t i c a l 119 11. N e c k - s t r e t c h p o s t u r e , h o r i z o n t a l 120 12. Backward s t a n c e p o s t u r e . 120 13. Male s e x u a l d i s p l a y 120 14. Female g i v i n g male s e x u a l d i s p l a y 120 15. Male p r e s e n t a t i o n o f a i r sac 121 16. Male f a n n i n g t a i l w h i l e e x h i b i t i n g s e x u a l d i s p l a y 121 17. Male w a l t z i n g 121 18. Male g i v i n g p r e - c o p u l a t o r y whoot a t t e r m i n a t i o n o f p r e - c o p u l a t o r y r u n . 121 19. Female s q u a t t i n g 122 20. Male e x h i b i t i n g s q u a t t i n g b e h a v i o u r . . . . 122 21; Sonogram o f male h o o t i n g 122 TABLE Page 22. Male i n h o o t i n g p o s t u r e 123 23. Sonogram o f male c a c k l e a g g r e s s i o n c a l l . . 123 24 • Male g i v i n g a g g r e s s i o n c a l l 123 25. Sonogram o f male cough a g g r e s s i o n c r y . . . 124 26o Sonogram o f female b l e a t 124 27. Sonogram of female whinny . 124 28. Sonogram o f female c r y . . . . . . . . 125 29. Sonogram o f female c l u c k 125 30. Sonogram o f female h o o t i n g 125 X ACKNOWLEDGEMENTS I am p a r t i c u l a r l y g r a t e f u l t o my s u p e r v i s o r , Dr. J.F. B e n d e l l , who made t h i s s t u d y p o s s i b l e . He o b t a i n e d f i n a n c i a l s u p p o r t and made h i m s e l f c o n s t a n t l y a v a i l a b l e f o r d i s c u s s i o n , c r i t i c i s m o f t h e t h e s i s and g e n e r a l s u g g e s t i o n s . Dr. C.W. R o b e r t s o f t h e Department o f P o u l t r y S c i e n c e gave f r e e l y of h i s t i m e f o r c o n s u l t a t i o n on problems o f a r t i f i c i a l i n s e m i n a -t i o n and s t a t i s t i c s . Dr. A . J . Wood p r o v i d e d t h e f a c i l i t i e s a t t h e v i v a r i u m and Dr. H.C. Norden gave h e l p f u l c r i t i c i s m on t h e c h a p t e r s on n u t r i t i o n . Other members o f my committee, Dr. J.R. Adams, Dr. D.H. C h i t t y , Dr. I . McT. Cowan and Dr. D.J. R a n d a l l gave h e l p f u l c r i t i c i s m o f t h e t h e s i s . T h i s work was s u p p o r t e d by a g r a n t from t h e N a t i o n a l R e s e a r c h C o u n c i l o f Canada and a s c h o l a r s h i p from t h e Canadian W i l d l i f e S e r v i c e . I am a l s o g r a t e f u l t o my f e l l o w s t u d e n t s , R.B. Dobson, A.N. Lance, Dr. B.R. Simard and Dr. F.C. Z w i c k e l f o r t h e i r a s s i s t a n c e and a d v i c e t h r o u g h o u t t h e s t u d y . CHAPTER 1 INTRODUCTION This work i s part of a long range study of the natural regulation of numbers of blue grouse (Dendragapus obscurus  fuliginosus) on Vancouver Island, conducted by Dr. J.F. Bendell. In the study of vertebrates i n the f i e l d i t i s often d i f -f i c u l t i f not impossible to manipulate environmental conditions for experimental purposes. It may also be d i f f i c u l t to obtain quantitative observations. Alternately, the unnatural environment of animals i n captiv-i t y may have unobserved effects upon t h e i r reactions to a given stimulus. For t h i s reason, conclusions drawn from a study upon captive animals should not be used to explain observations upon the same species i n the f i e l d without q u a l i f i c a t i o n . There were several objectives i n keeping blue grouse i n the U . B o C . aviary. One was to learn to keep them a l i v e and i n good health. Another was to learn to breed them in c a p t i v i t y . I f these two objectives were reached, further studies could provide information applicable to problems encountered in the f i e l d . In the course of holding and breeding grouse, many data were gath-ered that contributed to the understanding of t h e i r biology. F i e l d observations show that the mortality of chicks i s higher within some broods than the mean mortality of broods (Zwickel, 1965). This suggests a maternal influence upon the v i a b i l i t y of the eggs. One possible explanation i s that the quality of the diet of a hen a f f e c t s the v i a b i l i t y of her chicks. 2 Another possible explanation i s that the sperm of some males are inviable, but the importance of this suggestion i s diminished by the fact that the hens are polyandrous. If blue grouse could be held and bred successfully in captivity, hens could be fed diets varying in quality and the subsequent v i a b i l i t y of their chicks observed. Successful methods of holding and breeding would also permit a study of the basic nutritional requirements of blue grouse. Holding blue grouse would f a c i l i t a t e the study of behaviour by providing a set of constant conditions under which controlled experiments could be done. Additional descriptive data on pos-tures, displays and calls , d i f f i c u l t to obtain in the f i e l d , could be collected. This thesis describes the methods of holding and rearing blue grouse at the U.B.C. aviary from 1 9 6 2 to I 9 6 5 and the results of nutritional and behavioural studies done there. The following terms are defined here for use throughout the text. U.B.C. - University of British Columbia, Vancouver, British Columbia, Canada. Spring March, April, May, June Summer July, August Fa l l September, October Winter November, December, January, February Juvenile bird less than 1 year of age Yearling bird more than 1 year but less than 2 years of age bird over 2 years of age. Adult CHAPTER 2 ' METHODS OF HOLDING BLUE GROUSE IN CAPTIVITY Introduction Stocks of a l l North American galliforms have been held in captivity at some time. The Phasianidae and Perdicidae have reproduced many generations in captivity. Hatchability and sur-vival of over 80 percent are recorded for several varieties of pheasants, quail, and partridges (Greenberg, 1949; Delacour, 1959). Hatchability and survival has not generally been as high in the Tetraonidae. After 11 generations of ruffed grouse (Bonasa  umbellus) were raised in New York, Hatchability of eggs laid was only 65 percent and survival of chicks was 30 to 50 percent (Bump, 1949)» Fay (1963) got 76 percent survival of ruffed grouse chicks raised from eggs collected in the f i e l d . Evans (1964) stated that greater prairie chicken (Tympanuchus cupido pinnatus) did not survive well in captivity but no data were given. Sage grouse (Centrocercus urophasianus) raised in captivity had less than 30 percent survival (Pyrah, 1964). Blue grouse have been held in captivity several time before (Batterson, 1957; Gibson, 1965; Lacher, 1965; Hansen, 1961; Simp-son, 1935; Smith, 1963; and Wood, 1957). Most of these attempts were with open air pens 10 to 15 feet wide by 20 to 30 feet long and built on the ground. Gibson (1965) used small wire pens 19 inches square and 16 inches high. None of these attempts produced many grouse compared to productivity in the f i e l d . 4 Disease is the biggest problem with keeping gamebirds. Bump ( 1 9 4 9 ) and Delacour (1959) state the only way to prevent disease i s to raise birds in pens with wire mesh floors. Thus birds cannot feed in the feces and become infected with diseases such as enteritis, aspergillosis or coccidiosis. Smith (I960) and Lacher ( 1 9 6 5 ) had disease in their natural-floored pens. Gibson (personal communication) had disease in his wire-floored pens. Sage grouse kept in wire-floored pens had better survival than sage grouse kept on the ground (Pyrah, 1 9 6 4 ) . Materials and Methods Plate 1 i s a photograph of the avairy. Note the spaces in the sides to let in light but lack of spaces in the end. Trans-lucent fibreglass sheets were put on the roof to allow extra light and a r t i f i c i a l lighting was occasionally used to extend the day length during the reproductive season. The whole unit, with the exception of the outer half of the big pen, was roofed for protec-tion against the excessive rain. Figure 1 gives a plan view of the U.B.C. aviary. Table 1 gives the size and number of pens in the U.B.C. avi-ary. The pens of rows A, B, C, and D had a wire mesh (1 inch dia-meter) floor 2 feet above the ground. There were 2 foot high ply-wood dividers along the wire mesh walls between pens to prevent interaction (Plate 2 ) . There was a perch in each pen made of a piece of 2 inch square wood (Plate 2 ) . The big pen, E, had a crushed rock ( 1 / 2 inch diameter) floor. A few larger rocks (1 foot diameter) and logs were put into the big pen to simulate the natural environment. There was no growing vegetation. 5 PEN SIZE PEN ROW NUMBER HEIGHT1 WIDTH LENGTH Small A 12 4 2 10 B 10 4 2 8 Medium C 3 4 6 8 D 4 4 6 10 Large E 1 10 20 24 Table 1. Sizes and number of "*"A11 dimensions in feet. pens in U.B.C. aviary Laboratory, water and washing faciliti O Figure I. Plan view of the aviary (dimensions given in text) Legend: A,B,C,D,E, pen rows discussed in text permanent wire removable wire a water and feed W weighing scale F feed storage WB work bench O electrical outlet 7 Combinations of grouse of different ages, sexes and at dif-ferent densities were tested in the different sizes of pens. One grouse was usually kept in each small pen although 2 grouse were sometimes put in when there was an excess of birds as there was every f a l l . Two to 3 grouse were normally kept in the medium pens although 6 to & grouse were held in the f a l l . Six grouse were held in the big pen in one year and 4 the second year. Grouse were fed a commercial chicken breeder ration and #2 crushed g r i t . Feed was replaced every 2 or 3 days. Plate 2 shows the feeders and water-faunts used. Feed was stored in the Zoology building where i t was dry and mold did not form. Nest boxes were provided for females during the breeding season. Several sanitary measures were used to try to reduce the incidence of disease. Plastic sheets were put underneath the pens with wire floors to catch the droppings. These sheets were re-placed every 2 months. Feeders and water-faunts were washed and sterilized with bleach every 2 weeks. A spray of bactericide (brand name Izal) and fungicide (brand name Kleenzade) was used every month on the whole unit. Terramycin, polysol, baciferm, cod-liver o i l and aspirin were added to the drinking water of sick birds. Sick birds that stopped eating were force-fed a mix-ture of water, feed, and medicine to ensure they received some nutrients and medicine. Birds were weighed once a week during the f a l l and winter. They were caught by hand and weighed in a basket on an electric scale (Plate 3). They were not usually weighed during the spring or summer. Four grouse were held in individual wire cages in the vivar-8 ium. This vivarium was inside a building and was warm and dry. The pens were 21+ inches long, 18 inches wide and 16 inches high. Water and feed were provided in small pans which were weighed daily to measure intake. These birds were weighed daily. The room received natural light from a window on the wall. Results Feather picking developed i f more than 3 chicks were kept in the medium sized pens, or more than 1 chick in the small pens. Crowding resulted in several deaths due to picking. When Douglas f i r (Pseudosuga menziesii) boughs were put into the pens, the picking stopped. The plumage of birds kept in the small pens was frayed more than birds kept in larger pens. The plumage of birds held in the vivarium was frayed. The survival of grouse in each size of pen, from December to June of 1963-4 and 1964-5 i s given in Table 2. Birds that were moved from one size of pen to another between December and June were not counted. There was no significant difference in the sur-vival of grouse held in any size of pen during either year or be-tween the pooled survival of each group for 1964 and 1965• (Chi-square test at the 5 percent level of probability). A l l four grouse held in the vivarium survived from February through August. Survival of grouse in the aviary could only be compared to survival of grouse in the vivarium from February through June because there were no birds in the vivarium during December or January. There was no significant difference in sur-vival during this time (Chi-square test at the 5 percent level of probability). However, the time of heaviest mortality in the 9 1963-4 1964-5 LIVE IN LIVE IN LIVE IN LIVE IN raw o i i r , DECEMBER JUNE DECEMBER JUNE Small 46 22 46 13 Medium 8 4 13 3 Large 6 3 - 4 2 Table 2. Survival of grouse in each size of pen from December to June of 1963-64 and 1964-65 r 10 aviary was over. Disease was always present. It was d i f f i c u l t to determine whether medicine in the drinking water was of any benefit. Some sick birds taking medicine returned to good health. These birds were a minority. Figure 2 compares the body weights of grouse in the aviary to body weights of grouse from Vancouver Island for each month. Data from the same month of different years are considered toget-her. The Vancouver Island data are from Bendell (1955) and Simard (1964). The average weights of grouse from the f i e l d and from the aviary are about the same at the same time of the year. The weights of grouse held at the vivarium were comparable. One factor which could not be controlled in the aviary was moisture. The unit i s located in an area of poor drainage. Dur-ing the rainy winter season, the moisture of the ground gives a damp cold atmosphere to the unit. Air circulation i s poor because of the partitions and screens between the pens. The air tempera-ture was noticably lower inside the aviary than outside and the humidity was higher, but no quantitative data on the microclimate of the aviary was collected. A second unpredicted factor in aviary conditions was the very high population of house mice (Mus muscuius). They ate food from the bird feeders and le f t a mess of powdered feed and feces. Poisoning and trapping were done extensively but with l i t t l e ob-served effect. Table 3 gives the numbers, age and sex of grouse held in each size of pen and the proportion that showed reproductive 1500 IOOOH 500 0 adult males — i — i — i i — i — i — i — i — i — i — i j f m a m j j a s o n d 15001 1000 £ 500-j o o» 0 c adult females . I I — i — i — i — — i — i — i — i — i — r j f m a m j j a s o n d "5 1500 l 1000 m 5001 yearling males — i — i — i — i — i — i — i — i — i — i — i j f m a m j j a s o n d I500-I000 -500-0 yearling females — i — i — i — i — i — i — i — i — i — i — i j f m a m j j a s o n d Time Figure 2. Comparison of body weights of blue grouse in the aviary and the field Legendi weights of grouse in the aviary weights of grouse in the field 12 SMALL NO. OF NO. OF BIRDS BIRDS CLASS HELD SIZE OF PEN MEDIUM NO. OF NO. OF BIRDS BIRDS DISPLAYED HELD LARGE NO. OF NO. OF BIRDS BIRDS DISPLAYED HELD DISPLAYED Adult male Adult female Yearling male Yearling female Total 15 21 25 27 88 6 9 7 8 30 3 0 5 5 13 3 0 4 3 10 3 3 3 10 1 0 2 1 4 Table 3 . Numbers, age and sex of grouse held in each size pen and the proportion that showed reproductive display 13 display. Display here is defined as squatting for females and hooting or display for males (Appendix 2). The proportion of grouse that showed reproductive display in the medium pens was significantly larger than the proportion of grouse that showed reproductive display in the small pens (Chi-square test at the five percent level of probability done on the totals). There was no significant difference between the proportion of grouse that displayed in the small and large pens, or the medium and large pens. No grouse at the vivarium showed reproductive behaviour. The time that reproductive behaviour began each spring varied. In 19^5, hens began to squat and lay eggs 2 to 3 weeks before they did so in 1963 or I964. The peak of hooting was also earlier in 1965. In a l l years the beginning of reproductive behaviour in the aviary was later than in the f i e l d . Discussion The mortality rate of birds in a l l sizes of pens was equal (Table 2). The fact that birds frayed their plumage badly in the small pens suggests that larger pens are necessary to main-tain good plumage. The weights of aviary birds were comparable to the weights of birds in the f i e l d (Figure 2). This suggests that commercial chicken breeder ration met the nutritional requirements of blue grouse for maintenance of body weight. Conversely, i t suggests that birds in the f i e l d have no d i f f i c u l t y meeting their nutri-tional requirements. 14 The damp cool atmosphere of the aviary in the winter apparently provided an ideal environment for the growth of molds. Spraying was of l i t t l e lasting value. The mice may have acted as mechanical vectors of disease, since they were observed in the feeders many times. It is possible these two factors com-bined to aggravate the problem of disease. The pens at the vivarium were adequate for keeping birds alive but plumage was badly frayed. Daily handling for weighing caused additional fraying. Despite the extra handling, plus some sickness, no birds died. The major difference between the vivarium and the aviary was the climate. The aviary was cool and damp while the vivarium was warm and dry. It is lik e l y the environment of the vivarium was less conducive to the growth of mold and bacteria. It was free of mice that might have mechan-ic a l l y vectored diseases from the feces of one grouse to the feed of another. The small size of pens at the vivarium was probably not the sole reason that none of the grouse exhibited reproductive behaviour. Grouse held in small cages by Gibson, but not han-dled daily, exhibited reproductive behaviour. By comparison, when weighing of birds at the aviary was stopped well ahead of the breeding season, the reproductive behaviour did not seem to affected. This suggests that the stress of handling of grouse during the breeding season inhibits reproductive display. How-ever, there was no control group at the aviary which was not weighed during the same period. 15 The f a c t t h a t b i r d s b e g i n t o e x h i b i t r e p r o d u c t i v e b e h a v i o u r i n t h e a v i a r y l a t e r t h a n t h e y do i n t h e f i e l d s u g g e s t s t h a t some f a c t o r o r c o m b i n a t i o n o f f a c t o r s may i n h i b i t r e p r o d u c t i v e be-h a v i o u r i n t h e a v i a r y . The p h o t o p e r i o d i s about t h e same be-tween t h e a v i a r y and t h e f i e l d . One p o s s i b i l i t y i s t h a t t h e l o w e r t e m p e r a t u r e o f t h e a v i a r y may a c t t o i n h i b i t r e p r o d u c t i v e b e h a v i o u r . I n 1965, when t h e grouse i n t h e a v i a r y began t o d i s -p l a y two weeks e a r l i e r t h a n i n 1963 and 1964, t h e average number of hours of s u n l i g h t per day i n March was about double t h a t o f t h e p r e v i o u s 2 y e a r s . The mean monthly maximum and minimum te m p e r a t u r e from January t h r o u g h May showed l i t t l e v a r i a t i o n be-tween y e a r s . I t i s p o s s i b l e t h a t t h i s e x t r a amount of s u n l i g h t might have had an e f f e c t s i m i l a r t o t h a t of a greenhouse and made t h e a v i a r y warmer. Summary and C o n c l u s i o n s Of t h e pen s i z e s used, t h e medium s i z e d pens were best f o r h o l d i n g grouse on t h e b a s i s o f plumage c o n d i t i o n and r e p r o d u c -t i v e performance. Weights and death r a t e s were t h e same i n a l l s i z e s of pens. The c o l d damp environment o f t h e a v i a r y d u r i n g t h e w i n t e r may have been l e s s s u i t a b l e f o r h o l d i n g b l u e g r o u s e t h a n t h e warm d r y environment o f t h e v i v a r i u m because i t was more con-d u c i v e t o t h e growth o f d i s e a s e organisms. H a n d l i n g b l u e grouse d u r i n g t h e b r e e d i n g p e r i o d may have i n h i b i t e d r e p r o d u c t i v e d i s p l a y . 1 CHAPTER 3 MORTALITY OF BLUE GROUSE IN CAPTIVITY Introduction Mortality was high in grouse held at the aviary. This section is to discuss the pattern of mortality in each age class, the variation in the pattern and the causes of mortality. The mean annual mortality of adult blue grouse in the f i e l d is about 30 percent, and most of this occurs during the winter (Zwickel, 1965). Juveniles have a mortality rate of about 70 to 80 percent during the period from spring through f a l l and about 40 to 50 percent from f a l l through spring (Zwickel, I965). From these data i t i s clear that the highest mortality of a cohort of blue grouse in the f i e l d occurs during the f i r s t year of l i f e . I wanted to see i f this pattern of mortality was repeated in the aviary. Mortality in the aviary may be divided into two categories, natural and unnatural. A natural death i s one caused by disease or the aggressive actions of one bird on another. An unnatural death i s one caused by an artifact of captivity such as preda-tors, birds that suffocated in transport, sacrificed birds, birds donated to parasitology, escapes; and birds k i l l e d because of physical deformities. Materials and Methods Grouse chicks were brought to the aviary from Vancouver 17 I s l a n d each y e a r a t the end o f August. The numbers, sex and p l a c e o f c a p t u r e o f each c o h o r t o f c h i c k s i s g i v e n i n Table 4. D a i l y c h e c k i n g o f t h e a v i a r y gave a c c u r a t e d a t a on t h e t i m e o f m o r t a l i t y . B i r d s were e i t h e r a u t o p s i e d on t h e day t h e y were found dead, or f r o z e n and a u t o p s i e d i m m e d i a t e l y upon th a w i n g . Two g r a p h i c p r e s e n t a t i o n s o f m o r t a l i t y a r e g i v e n . To show t i m e d i s t r i b u t i o n and causes o f d e a t h s , a h i s t o g r a m i s g i v e n f o r a l l deaths o f each age c l a s s . A s u r v i v o r s h i p curve f o r each age c l a s s i s a l s o g i v e n . R e s u l t s Of t h e 192 b i r d s r e c e i v e d , 140 d i e d o f n a t u r a l c a u s e s , 31 o f u n n a t u r a l causes and 21 were a l i v e a t t h e t e r m i n a t i o n o f t h e p r o j e c t . T a b l e 5 g i v e s t h e causes o f m o r t a l i t y o f b l u e g rouse o f each age c l a s s a t t h e U.B.C. a v i a r y from August 1962 t h r o u g h August 1965. The c l a s s " o t h e r " i n c l u d e s deaths i n t r a m s p o r t , p r e d a t o r k i l l s , s a c r i f i c e s , e s c a p e s , b i r d s g i v e n t o p a r a s i t o l o g y and b i r d s k i l l e d because o f p h y s i c a l d e f o r m i t i e s . I n many (49) cases t h e r e were s e v e r a l symptoms o f s i c k n e s s f o u n d upon p o s t -mortem e x a m i n a t i o n but t h e exact cause of death c o u l d not be p o s i t i v e l y d e t ermined. The p r o p o r t i o n o f b i r d s t h a t d i e d o f each cause o f death was about t h e same f o r each sex and age c l a s s ( T able 5). One e x c e p t i o n was a s p e r g i l l o s i s w h i c h ap-peared t o be more p r e v a l e n t i n males, but t h i s was not s t a t i s -t i c a l l y s i g n i f i c a n t ( C h i - s q u a r e a t t h e 5 p e r c e n t l e v e l o f prob-a b i l i t y ) . The number o f c h i c k s t h a t d i e d o f p i c k i n g was h i g h e s t 18 YEAR •NO. OF MALES NO. OF FEMALES SOURCE TOTAL 1962 1963 1964 Total 21 37 37 95 22 32 43 97 Middle Quinsara Lake, Vancouver Island 16 mi. N.W. Courtenay, Vancouver Island 16 mi. N.W. Courtenay, Vancouver Island 43 69 80 192 Table 4 . Numbers, sex and place of capture of each cohort of blue grouse chicks received in the U.B.C. aviary each August 19 CAUSE OF DEATH 1962 MALES CHICKS FEMALES 1963 MALES CHICKS FEMALES 1964 MALES CHICKS FEMALES Unknown 5 6 8 10 9 11 Gizzard erosion 4 1 7 5 1 7 Aspergillosis 8 3 3 4 5 1 Enteritis 0 • 1 7 3 3 4 Picking and aggression 0 0 7 5 3 0 Starved on f i r 0 5 0 0 0 0 Predator 0 0 2 0 0 3 Internal hemorrhage 0 1 1 0 0 0 Escaped 0 0 0 0 2 0 Ulcerative gizzard 1 0 0 0 1 0 Proventriculitis 0 1 0 0 0 0 Other 2 1 2 4 S 6 Number starting 21 22 37 32 37 43 Total deaths 20 19 37 31 32 32 Total survivors 1 3 1 0 5 11 Table 5. Causes of mortality of blue grouse of each age class at the aviary from August, 1962 through August, 1965 20 in the 1963 cohort. Figure 3 is a histogram of the number of deaths and their causes for each age class from the time they arrived at the aviary through August 1965. Figure 4 gives the survivorship curves of each cohort of grouse from the time they arrived in the aviary through August I965. The mortality was generally higher during the winter than the summer in each age class (Figures 3 and 4 ) . The exception was the cohort of chicks from 1963 which had low mortality dur-ing the f i r s t winter and high mortality during the following summer. The period of greatest mortality for the whole popula-tion was during the winter of 1964-5. The survival of the 1962 and I963 chicks through the f i r s t year of l i f e was significantly better than the survival of the I964 chicks (Chi-square test at the 5 percent level of prob-ability) . The 1963 chicks were held in the medium pens at densities of & to 10 with no cover. They pecked each other until f i r boughs were put in for cover. Picking then stopped immediately. Boughs were put in the pens when holding the 1964 chicks and no deaths resulted from picking caused by the stress of crowding. Discussion The incidence of disease was usually higher during the winter than the summer. Normally the winters are cool and wet while summers are warm and dry. As discussed in Chapter 2, the warm dry environment is less suitable for growth of mold or 21 10; 6-2-0 1962 chicks asondj fmamjj asond j fmamjjasondj f T — I — I — I . I . I I mam j j a 1962 1963 1964 1965 to JC o <u •o o a> Time I0n 6 0 1963 chicks asondj fmamj j asondj fmamj j a 1963 1964 1965 101 6 2- $ 0 11 1964 chicks asond j fmamj ja 1964 1965 Figure 3. Deaths and their cause by age and number Legend: mm picking and aggression aspergillosis gizzard erosion s enteritis starved on fir m ulcerated gizzard m internal hemorrhage s unknown ca proventriculitis 22 40 20 1962 chicks 0 t r r r r b ' s ' o ' n ' d ' j ' f ' m ' a ' m ' j ' j ' a ' s ' o ' n ' d j ' f ' m ' a ' r n ' j ' j V s V n ' d ' j ' f W a r n 1 j ' j ' a 1 1962 1963 co o lc o «4— o a> £ 3 60 40 20-1964 1965 1963 chicks asondj fmamj j asondj f ma m j j a 1963 1964 6 0 1965 Time Figure 4. Survivorship of each age-class of chicks brought to U.B.C. aviary 40 20 -1964 chicks i—i—i—i , i . i , i—i i I . I . I i asondjfmamj j a 1964 1965 23 bacteria. November through January of 1964-65 was particularly cold and wet. Several feet of snow f e l l in December and remained through half of January. I think this period of excessive cold and wet climate in the aviary was a factor in the particularly high mortality of a l l age classes during that period. I cannot explain why the I963 chicks had a low winter mortality and high summer mortality during their f i r s t year in the aviary. They later followed the normal pattern of mortality. The survival of the 1962 chicks was significantly better than that of the 1964 chicks for the f i r s t year of l i f e and significantly better than the 1963 chicks for the f i r s t 2 years of l i f e . The 1962 chicks were from Middle Quinsam Lake, Vancou-ver Island, while the I963 and I964 chicks were from 16 miles N.W. of Courtenay, Vancouver Island. Differences in survival may be a reflection of a difference in the stock. However, I think the higher mortality of the 1964 chicks was more lik e l y due to the bad environmental conditions during their f i r s t year in the aviary. I do not think the high incidence of disease affected the subsequent observations upon "healthy" birds. However, none of the "healthy" birds were sacrificed to check for disease that might have inhibited reproduction or behaviour. Summary and Conclusions The time of heaviest mortality of grouse in the aviary was usually during the winter. This was probably because the damp 24 climate during that period was conducive to the growth of dis-ease organisms. The most common diseases of grouse at the aviary were giz-zard erosion, aspergillosis and enteritis. The differences in survival were more dependent upon c l i -matic conditions of the aviary than upon some difference between the cohorts. Thus, i t was unlikely that the 1962 birds had bet-ter survival because they were of a more resistant stock. The proportion of birds of each cohort that died of each natural cause of mortality was about even between years and sexes. When f i r boughs were put into pens in which several chicks were held, they did not peck each other. CHAPTER 4 BREEDING BLUE GROUSE IN CAPTIVITY Introduction Several d i f f i c u l t i e s are involved in breeding game birds in captivity. The stress of an unnatural environment may in-hibit reproductive behaviour. It is d i f f i c u l t to assess the importance of various factors in the natural environment for the stimulation of breeding behaviour. Lighting, external noise, size of pen, age ratios, feed, stock and synchronization of the reproductive behaviour of both sexes may have influenced breeding behaviour in the aviary. There are two basic methods of breeding game birds in captivity, natural and forced. The natural method involves selection of breeding pairs in the f a l l or about 2 months before the breeding season occurs and penning the pairs together. Courtship and mating then occurs without human interference. The sex ratio may be monogamous or polygamous depending upon the social structure of the species. The forced matings, the male and female are put together at the time mating should occur. The female may be taken to the male, the male to the female, or they may both be put into a strange pen at the same time. They may be l e f t together for only a few minutes or for a few days. 26 Materials and Methods Al l 3 sizes of pens described in Chapter 2 were used for breeding experiments. Several variations of the natural method of mating grouse ' were tried. In December 1963, 4 pairs of blue grouse were put in medium pens and held there through the breeding season of 1964- Six birds (1 adult male, 2 adult females, 1 juvenile male and 2 juvenile females) were put in the big pen (E, Fig. 1) in February, 1964 and held through the summer. The following year 4 grouse (1 adult male, 1 adult female, 1 juvenile male and 1 juvenile female) were held in the same pen through the breed-ing season. Pen rows A and B (Fig. 1) faced each other on a common aisle. During the breeding season, doors were le f t open over-night to allow breeding pairs to meet in the aisle without human interference. Pens that held males and females on oppo-site sides of the aisle were joined together and l e f t over-night . Forced matings were tried several times. Females were taken to males and males to females for short periods of time. Males and females were also penned together overnight. Results The number of grouse showing reproductive behaviour in the aviary is given in Table 6. Of the 4 pairs of grouse penned together in 1964, 3 males exhibited reproductive behaviour. No females were observed 27 (a) Males CATEGORY NUMBER DISPLAYING1 HOOTING 2 Ad. D.o. fuliginosus 19 10 11 Ad. D.o. pallidus 1 1 1 3 Yr. D.o. fuliginosus 33 13 9 Total 53 24 21 (b) Females CATEGORY NUMBER SQUATTING BRED LAYING Ad. D.o. fuliginosus 24 9 4 g Ad. D.o. pallidus 6 4 0 0 Yr. D.o. fuliginosus 36 12 5 7 Total 66 25 9 15 Table 6. Number of blue grouse at U.B.C. aviary showing reproductive behaviour from 1963 through 1965 Appendix 2 2Adult Yearling 28 exhibiting reproductive behaviour, no matings were observed and no eggs were la i d . One male k i l l e d his pen mate. In 1964 the dominant adult male in the big pen displayed and hooted. He was observed displaying to females on several occasions. However no females were observed exhibiting repro-ductive behaviour, no matings were observed and no eggs were laid . In 1965 the adult male in the big pen again displayed and became dominant. However he k i l l e d 2 yearling males put succes-sively into the big pen and had to be removed because of a shortage of males. Of yearling males put into the big pen in February, one became dominant by the end of April. He hooted and displayed to the females. One female was observed squatting but no matings were seen and no eggs were lai d . The only successful method of getting grouse to mate in the aviary was to take a male to a female that was already squatting (Table 7). Eggs were laid each year. Several were la i d by hens that were not mated. F e r t i l i t y and hatchability of these eggs were low (Table 8). Discussion Almost a l l hens on the breeding range, including yearlings, breed (Zwickel, I 9 6 5 ) . Yet at the aviary, only 38 percent of the hens exhibited reproductive display and only 13 percent bred. A slightly higher percent of the males displayed and hooted. Even though several grouse exhibited reproductive display in the aviary, they did not breed readily. Hens did not squat i f 29 1963 1964 1965 METHOD TIMES TRIED TIMES MATED TIMES TIMES TRIED MATED TIMES TIMES TRIED MATED Joining pens 7 0 - - — — Loose in aisle 4 0 - -Penned overnight 8 0 - -Female to male before squatting 16 0 - - -Female to male after squatting 4 0 - -Male to female before squatting 11 0 - -Male to female after squatting 2 3 a 24 1 27 3 Table 7 . Results of natural and forced mating attempts in U.B.C. aviary 30 SOURCE OF EGGS NO. OF EGGS NO. POSSIBLY FERTILE NO. FERTILE NO. HATCHED NO. SURVIVED UNTIL FALL Naturally mated hens 47 39 18 4 2 A r t i f i c i a l l y inseminated 59 41 10 5 2 hens Table 8. F e r t i l i t y and hatchability of eggs laid at the U.B.C. aviary .31 they were handled to be put into the pen of a male. There were only 1 or 2 males each year that would display after being han-dled, but these males would do so at any time during the repro-ductive period. The mating of grouse in the aviary was complicated by 2 additional problems that were d i f f i c u l t to overcome. The f i r s t was the fact that females were sexually imprinted upon humans and squatted only in the presence of humans. Males did not become sexually imprinted upon humans. The second was that the males often broke their primaries in the pens and were not able to balance on a squatting female long enough to copulate. In 1965, 5 potential matings were unsuccessful because the male f e l l off the female. External noise also seemed to affect the behaviour of grouse. The caribou unit was constructed during the period of reproductive display at the aviary. The birds became extremely restless and very easily frightened. It has been observed since, that the behaviour of the grouse i s unsettled and unpredictable upon days of excessive noise made by the caribou and subsequent construction. It i s d i f f i c u l t to quantify the effect of such a factor upon the reproductive behaviour of grouse in captivity. However, I feel the influence of external noise was detrimental to breeding success. Several factors contributed to the low f e r t i l i t y and hatchability of the eggs from the aviary. A l l embryos in 1963 were k i l l e d at about 1 day of age by the overheating of the incubator. As a result, the determination of f e r t i l i t y was 32 d i f f i c u l t because t h e r e was l i t t l e o r no development. Thus t h e f i g u r e f o r t h e number o f f e r t i l e eggs i n I963 i s a minimum. These eggs a r e i n c l u d e d i n t h e f i g u r e s f o r n a t u r a l l y mated hens (Table 8). The f e r t i l i t y and h a t c h a b i l i t y r e c o r d e d by o t h e r w o r k e r s has a l s o been low (Sm i t h , 1963; L a c h e r , 1965). However, n e i t h e r of t h e s e workers examined unhatched eggs and t h u s c o n s i d e r e d h a t c h a b i l i t y as f e r t i l i t y . T a b l e 9 compares t h e h a t c h a b i l i t y of b l u e g r o u s e eggs from t h e d a t a o f Smith ( i 9 6 0 ) , L a c h e r (1965) and t h e a v i a r y a t U.B.C. The h a t c h a b i l i t y r e c o r d e d by Smith i s h i g h e r t h a n t h a t o f U.B.C. o r La c h e r (Table 8). T h i s d i f f e r e n c e was s t a t i s t i c a l l y s i g n i f i c a n t w h i l e the d i f f e r e n c e i n h a t c h a b i l i t y between t h e da t a o f Lache r and U.B.C. was not ( C h i - s q u a r e a t t h e 5 p e r c e n t l e v e l o f p r o b a b i l i t y ) . The f e r t i l i t y and h a t c h a b i l i t y of eggs i n t h e f i e l d a r e about 90 p e r c e n t and 80 p e r c e n t r e s p e c t i v e l y ( Z w i c k e l , I 9 6 5 ) . Thus o n l y t h e h a t c h a b i l i t y r e c o r d e d by Smith (I960) approached t h a t r e c o r d e d i n t h e f i e l d . However, d e s p i t e t h e h i g h e r h a t c h -a b i l i t y , t h e s u r v i v a l o f c h i c k s u n t i l f a l l was poor. Summary and C o n c l u s i o n s A l o w e r p r o p o r t i o n o f male and female grouse i n t h e a v i a r y e x h i b i t e d r e p r o d u c t i v e d i s p l a y t h a n appeared t o do so i n t h e f i e l d . T h i s was p r o b a b l y due t o t h e s t r e s s o f c a p t i v e c o n d i t i o n s . The o n l y s u c c e s s f u l method o f mati n g grouse a t U.B.C. was t o put a d i s p l a y i n g male i n t o t h e pen o f / s q u a t t i n g f e m a l e . The hens SOURCE OF DATA NO. OF EGGS NO. HATCHED % HATCH Smith 30 23 77 Lacher 23 7 30 U.B.C.1 54 9 17 Table 9. Comparative hatchability of blue grouse eggs "^ The clutch of eggs accidentally k i l l e d in 1963 is not included in this figure. 34 were sexually imprinted on humans so that for the female to remain squatting, a human s t i l l had to be in sight. The f e r t i l i t y and hatchability of eggs was low at U.B.C. The data of other workers also indicated d i f f i c u l t y breeding and rearing blue grouse successfully. CHAPTER 5 ARTIFICIAL INSEMINATION Introduction The breeding of blue grouse at U.B.C. aviary was relatively unsuccessful, as discussed in Chapter 4« In 1964, 3 hens began to lay i n f e r t i l e eggs. These hens had not previously squatted or otherwise indicated arrival into reproductive condition. Attempts to breed them failed. It was decided to attempt a r t i f i c i a l insemination as a last possible method to get f e r t i l e eggs. Squatting and egg laying were taken as indicative of the time of ovulation and thus the correct time to inseminate. Materials and Methods The techniques developed for insemination of chickens and turkeys were used (Taylor, 1949). Dr. C'.W. Roberts, Dept. of Poultry Science, U.B.C, further modified the technique as necessary for blue grouse. For collection of sperm, the male was held with his head and shoulders under the arm of one worker. A second worker massaged the region around the base of the t a i l . This stimulated the eversion of copulatory protrusions upon which the sperm flowed. The sperm was collected upon a clean slide and trans-ferred to a small syringe. Sperm was collected from males that hooted or showed 36 reproductive display. The response varied greatly. The volume collected was always small, varying from . 1 to . 5 ml. An aver-age collection from a chicken is . 5 to 1 . 0 c.c. (Taylor, 1 9 4 9 ) ' Although no quantitative data were collected, the adults gener-ally produced more seminal f l u i d than did yearlings and the response of both improved after 1 or 2 collections. For insemination, the female was held between one worker's knees. Pressure applied to the abdomen with the hands forced the uterus to evert through the cloaca. The syringe was in-serted into the uterus and sperm injected. Touching a squatting female on the back sometimes caused the eversion of the uterus. One squatting hen was stimulated to evert her uterus and then was inseminated. To prevent contamination of sperm by feces, food was re-moved from males one day prior to collection of sperm. Care was also taken to prevent fecal contamination of sperm when everting the uterus. Results Handling the grouse for a r t i f i c i a l insemination caused excitement and stress. The number of hens squatting and laying eggs was reduced after handling for insemination (Table 1 0 ) . Only hens that lai d 2 or more eggs prior to handling for insemi-nation continued to lay. Table 11 compares the mean clutch size of hen blue grouse from Vancouver Island to the clutch size of hens that were naturally mated and a r t i f i c i a l l y inseminated in the U.B.C. aviary. 37 NO. OF HENS NO. OF HENS NO. OF HENS NO. OF HENS SQUATTING SQUATTING LAYING EGGS' LAYING EGGS BEFORE AFTER BEFORE AFTER YEAR INSEMINATION INSEMINATION INSEMINATION INSEMINATION 1964 4 2 4 3 1965 4 0 6 2 Table 10. The effect of handling hens for a r t i f i c i a l insemi-nation upon squatting and egg laying 3$ MEAN CLUTCH GROUP SAMPLE SIZE SIZE RANGE Vancouver Island 1964 15 7.00 $-9 A r t i f i c i a l l y inseminated hens in the aviary 5 9.60 7-13 Naturally mated hens in the aviary 4 10.25 7-14 Table 11. Comparison of mean clutch size of blue grouse on Vancouver Island to blue grouse that were naturally mated and a r t i f i c i a l l y inseminated in the U.B.C. aviary r 39 The clutch sizes from hens in the aviary were larger. The f e r t i l i t y of eggs l a id by a r t i f i c i a l l y inseminated hens was 24 percent. The f e r t i l i t y of eggs l a id in the aviary by naturally mated hens was 46 percent (Table 8 ) . The differ-ence was not s ignif icantly different (Chi-square at the 5 per-cent level of probabil ity) . The f e r t i l i t y of eggs from a r t i f i c i a l l y inseminated grouse is low compared with 83 percent f e r t i l i t y obtained by a r t i f i c i -a l ly inseminating Japanese quail (Coturnix japonica) (Marks and Lepore, 1965). Discussion The stress of handling hens for insemination seemed to affect subsequent squatting and egg laying. Hens usually dis-continued laying i f handled before dropping 2 or 3 eggs, but continued i f handled later . This suggests there may be a c r i t i -cal period after which laying w i l l continue despite a certain amount of stress. The technique of a r t i f i c i a l insemination overcomes the d i f f i cu l t i e s of trying t o get grouse to display and mate natu-r a l l y . Table 11 suggests that blue grouse are indeterminate layers so that larger clutches than found in the f i e l d may be expected i f eggs are taken from the nest box dai ly. It may be possible t o maintain an aviary population by selecting males which give a good collection of sperm, selecting females that are good layers and allowing them to lay 2-3 eggs prior to in -semination . 40 Summary and C o n c l u s i o n s B l u e grouse can be a r t i f i c i a l l y i n s e m i n a t e d by u s i n g t h e t e c h n i q u e s d e v e l o p e d f o r domestic f o w l . For b e s t r e s u l t s i n terms o f c o n t i n u e d egg l a y i n g , hens s h o u l d be a l l o w e d t o l a y 2-3 eggs b e f o r e i n s e m i n a t i o n . The f e r t i l i t y and h a t c h a b i l i t y of eggs from hens t h a t were a r t i f i c i a l l y i n s e m i n a t e d i n t h e a v i a r y were comparable t o t h a t of hens t h a t were n a t u r a l l y b r e d i n t h e a v i a r y . The f e r t i l i t y and h a t c h a b i l i t y o f both t h e s e s e t s o f eggs was 1/3 t o 1/2 o f t h a t r e c o r d e d i n t h e f i e l d . CHAPTER 6 INCUBATION AND BROODING Introduction A l l incubation and brooding were done a r t i f i c i a l l y so that a l l eggs and a l l chicks would receive identical treatment. This eliminated possible variation in maternal care. Materials and Methods Thirty-four eggs were incubated in a Humidaire 150 egg gravity air incubator, model 10 (Plate 4). This was kept in a small plywood shack. Each egg was collected, individually marked and set the day i t was la i d . Eggs were incubated as close to 100 degrees F. and 70 percent humidity as possible. They were hand-turned 3 times a day at angle of about 30 degrees from the vertical. Eggs were not turned after the twenty-second day of incubation. The incubation period of blue grouse eggs is about 25 days (Smith, I960). Sixty-three eggs were incubated in a large Jamesway incu-bator, model 2940, with capacity for several thousand eggs (Plate 5). The temperature was kept around 101 degrees. Humid-ity was high but was not measured. Eggs were automatically turned about 30 degrees from vertical 5 times a day. After 22 days of incubation each egg was put in an individual wire hatch-ing basket and set to hatch in another compartment of the incu-bator at the same temperature and humidity. Eggs set to hatch 42 were not turned. Chicks were l e f t to dry in the incubator for one day after hatching., They were then transferred to the brooder. The brooder was 3 feet square and 1 1/2 feet deep with a wire floor (Plate 6 ) . The mesh was 1/4 inch by 3/4 inch which was large enough for droppings to pass through. The brooder was heated by an infra-red heat lamp suspended so that the temperature at the level of the back of a chick was 98 degrees F. Chicks were fed for 6 to 8 weeks on Buckerfield Ts turkey poult ration plus a daily supplement of chopped lettuce and tomato fed in excess. Chicks were then fed Buckerfield Ts tur-key starter ration up to about 3 months of age. They were fi n a l l y put on Buckerfield's chicken breeder ration for holding. Results No eggs hatched in the small incubator. A maximum-minimum thermometer showed that the temperature in the hut varied between 96 and 108 degrees F. The internal temperature of the incubator varied between 98 and 101 degrees F. Examination of the eggs showed that a l l embryos died after about 1 day in the incubator. It was later found that the thermometer inside the incubator read about 3 degrees low. Thus a l l the embryos likely had been k i l l e d by overheating. Conditions remained constant between 99 and 100 degrees F. in the battery incubator. Nine chicks hatched. Conditions in the brooder remained constant. Five chicks died while their 4 pen-mates and brood-mates flourished under 43 i d e n t i c a l c o n d i t i o n s at the same time. The dead c h i c k s were not pic k e d and no s i g n of s i c k n e s s was found i n t e r n a l l y . D i s c u s s i o n The v a r i a t i o n i n temperature of the s m a l l i n c u b a t o r was due to v a r i a t i o n i n e x t e r n a l temperature and thus i n d i c a t e d poor i n -s u l a t i o n . Zwickel (1965) used s m a l l i n c u b a t o r s i n a shaded te n t where e x t e r n a l temperature had minimal f l u c t u a t i o n and found they h e l d f a i r l y constant temperature. Thus the main problem i n using the s m a l l i n c u b a t o r i s keeping i t at a r e l a t i v e l y constant temperature. The overhe a t i n g of the i n c u b a t o r probably k i l l e d the embryos i n 19&3• Summary and Conclusions The combination of i n c u b a t i n g grouse eggs i n the b a t t e r y i n c u b a t o r s and brooding c h i c k s by Z w i c k e l T s procedures was s a t i s f a c t o r y . CHAPTER 7 EXPERIMENTS ON DIETS AND VIABILITY Introduction This chapter presents data from experiments designed to test the effect of the diet of the hen upon the v i a b i l i t y of her chicks. As discussed in the introduction, f i e l d observations on blue grouse have shown that the mortality of chicks is higher within some broods, that the mean mortality of broods (Zwickel, 1 9 6 5 ) . One possible explanation i s that a difference in the quality of the diet of the hen may be reflected in the v i a b i l i t y of her chicks. The quality of the diet of the hen apparently determines the v i a b i l i t y of the chicks of the red grouse (Lagopus  lagopus scoticus) and the peafowl (Legumus g a l l i ) (Allen, 1964; Jenkins, 1963). The protein level of the diet of the hen i s important be-cause of the heavy drain upon protein reserves for egg produc-tion. The protein level of the diet of female rats i s c r i t i c a l to reproductive success in the laboratory (Nelson and Evans, 1953). The main part of the diet of blue grouse on the winter range i s f i r needles (Beer, 1943; Hoffmann, 1961; Marshall, 1946; Stewart, 1944). A chemical analysis of Douglas f i r (Pseudosuga  menziesii) needles at U.B.C. indicated a crude protein level of 6 percent. A similar analysis by Hoffmann of needles of white 45 f i r (Abies concolor) indicated a protein l e v e l of 5 to 7 percent. By comparison to recommendations i n the l i t e r a t u r e f o r poultry di e t s , 6 percent would seem to be f a r too low f o r maintenance l e t alone successful breeding. After a r r i v a l on the breeding range i n the spring, hens eat succulent greens almost exclusively. Cocks s t i l l eat f i r needles as well as greens (Bendell, personal communication). This sug-gests that the f i r diet of the winter range i s not suitable for egg production and a supplement of greens, higher in protein, i s needed. Materials and Methods The plan i n 1°63 was to hold females over the winter on diets that d i f f e r e d i n l e v e l s of protein. In the spring, these hens were to be bred and the subsequent v i a b i l i t y of t h e i r chicks compared. Two groups of 12 hens were selected at random. The f i r s t group was held on a commercial chicken breeder rati o n which was t h e o r e t i c a l l y as complete i n n u t r i t i v e requirements as possible. The protein l e v e l was 18 percent. The second group was held on a diet of dried and pelleted f i r needles. In 1962, Zwickel held a group of blue grouse for two months on a diet of dried and pe l -leted f i r needles (personal communication). I hoped to do the same and breed the hens to study the effect of t h i s low quality diet on the v i a b i l i t y of the chicks. Four diets were designed to condition the grouse to a grad-ual change from chicken breeder ration to f i r . These diets con-46 tained 25 percent, 50 percent, 75 percent and 100 percent f i r . The remainder of the diet was made up of the commercial chicken breeder ration. The level of f i r in the diet was increased at two week intervals to allow time for the hens to adjust to the change. To compare the effect of different l o c a l i t i e s on the qual-i t y of the f i r , pellets were made from stands at U.B.C. and the Haney forest. A ration was also made with needles from a l l parts of the tree to compare with one from only the growing tips of the branches. In the spring, when greens were available to hens in the f i e l d , 1/2 of the group of hens s t i l l on the f i r diet and 1/2 the group of hens on the breeder ration diet received a supple-ment of greens which was in excess at a l l times. This was to test i f the availability of succulent greens had a significant effect upon the reproductive success of the hens. The grouse were weighed twice a week to follow weight changes. In I 9 6 4 , 3 randomly selected groups of hen blue grouse were selected to be held on 3 commercial rations with varying levels of protein. The percent protein of the 3 rations fed to hen blue grouse at U.B.C. aviary from September, I963 through Decem-ber, I964 was as follows. RATION PERCENT PROTEIN Chicken breeder ration Turkey starter ration Turkey poult ration 18 24 28 47 Hens were h e l d on t h i s d i e t t h r o u g h t h e w i n t e r o f 1963-4 t o be b r e d i n t h e s p r i n g o f 1964. I n 1965, hens were h e l d on c h i c k e n b r e e d e r r a t i o n o n l y . The s o l e o b j e c t i v e was t o h o l d them over t h e w i n t e r under t h e most f a v o u r a b l e c o n d i t i o n s and b r e e d them i n t h e s p r i n g . R e s u l t s The grouse s u r v i v e d and m a i n t a i n e d weight on a d i e t o f up t o and i n c l u d i n g 75 p e r c e n t f i r and 25 p e r c e n t commercial b r e e d e r r a t i o n . By o b s e r v a t i o n , t h e f o o d i n t a k e was h i g h e r on t h i s d i e t t h a n on o t h e r r a t i o n s . When t h e b i r d s were p l a c e d on a pure f i r d i e t t h e y began t o l o s e weight i n 3 days. W i t h i n 2 weeks t h e y r e a c h e d a w e i g h t o f 750 grams from a h e a l t h y w e i ght of 85O t o 900 grams o r more. At t h i s p o i n t t h e y were a g a i n f e d the c h i c k e n b r e e d e r r a t i o n . However, 5 o f t h e s e hens c o n t i n u e d t o l o s e weight and e v e n t u a l l y d i e d . No b i r d s e x h i b i t e d r e p r o d u c t i v e b e h a v i o u r w h i l e on any of t h e f o u r d i e t s , and a l l b i r d s were back on c h i c k e n b r e e d e r r a t i o n by t h e time t h e b r e e d i n g season was ov e r . No a d d i t i o n a l b r e e d i n g b e h a v i o u r was e x h i b i t e d by hens w i t h a d i e t supplement of g r e e n s . I n 1963, a l l embryos were k i l l e d i n t h e i n c u b a t o r as d i s c u s s e d i n Chapter 6, so t h a t t h e r e was no i n f o r m a t i o n on v i a b i l i t y . I n 1964, o n l y f e m a l e s h e l d on c h i c k e n b r e e d e r r a t i o n showed a s i g n i f i c a n t amount o f r e p r o d u c t i v e b e h a v i o u r ( T a b l e 12). T h i s d i f f e r e n c e i s s t a t i s t i c a l l y s i g n i f i c a n t ( C h i - s q u a r e t e s t a t t h e 5 p e r c e n t l e v e l o f s i g n i f i c a n c e ) . I n a d d i t i o n , 2 a d u l t 4 8 RATION NO. OF HENS ON RATION DURING BREEDING SEASON NO. OF HENS EXHIBITING REPRODUCTIVE BEHAVIOUR NO. OF LAYING HENS EGGS Chicken breeder ration 13 6 5 Turkey starter ration 14 0 1 Turkey poult ration 9 1 0 Table 1 2 . Number of hen blue grouse in U.B.C. aviary that showed reproductive behaviour and laying on each ration 49 females that displayed and bred while held on the chicken breeder ration in 1963 did not display or breed while held on turkey poult ration in 1964. One female that displayed and bred in 1963 while held on chicken breeder ration did so again in 1964 while on the same ration. The mortality rates of birds held on the turkey starter and poult rations are higher (Table 13) . These are not statis-t i c a l l y significant (Chi-square test at the 5 percent level of probability). However, I feel the differences are s t i l l bio-logically significant. Discussion I do not know why the grouse lived on Zwickel's pelleted f i r but not on mine. It was suggested by Buckerfield's agent who supervised on the making of the pellets that they might have been contaminated with other feed residues that were s t i l l in the pelleting machine. It may have been due to variation in processing and pelleting procedures, the time of year, portion of tree used, location of stand, annual v a r i a b i l i t y > o r another factor of which I am not aware. The 1964 experiments compared 3 high quality diets. They did not provide any information upon the v i a b i l i t y of chicks from hens on different diets. The excessively high protein content of the turkey starter and poult rations was the most important difference between them and the chicken breeder ration. Bump (1949) stated that exces-sive protein in the diet w i l l stop laying and bring on moult. NO. BIRDS ON NO.BIRDS DIED RATION EACH RATION ON EACH RATION Chicken breeder ration 23 13 Turkey breeder ration 14 12 Turkey poult ration 9 7 Table 13. Survival of hen blue grouse held on each ration at U.B.C. aviary from September, 1963 through December, 1964 51 The recommended dietary protein level for chickens is 18 percent (Ewing, 1963); turkeys, 20 percent (Ewing, 1963); and pheasants, 18 percent (Greenberg, 1949). Delacour (1959) recommends 24 percent protein for pheasants. However, animals eat to meet a caloric requirement so that percent protein is.of less importance. The caloric values of rations used to hold female blue grouse at U.B.C. aviary, September, 1963 through December, I964 are given in Table 14. Note the ratio of calories to percent protein. By simple ratio i t may be seen that a hen on turkey starter ration is taking in a greater amount of protein than a hen on chicken breeder ration, to meet the same caloric require-ment, by not 1/3 greater. The same applies to the turkey poult ration. Thus, comparison of the effect of different dietary levels of protein was invalid. However, a difference in repro-ductive behaviour and laying was s t i l l demonstrated between groups of hens on different diets. The hypothesis was that the quality of the diet of the hen would be reflected in the v i a b i l i t y of her chicks. Thus, the quality of the diet of the hen would ultimately control the num-ber of grouse produced. However, to .date there is no evidence from the f i e l d to indicate that birds arrive onto the breeding range in poor condition. There is no evidence of any shortage of nutritious food for hen grouse on the breeding range. The aviary experiments in 1963 and 1964 showed a difference in repro-duction and survival between hens on different diets. However, I cannot conclude that these experiments have produced any re-sults which are meaningful to natural conditions. 52 RATION % PROTEIN CALORIC VALUE CAL./% PROTEIN Chicken breeder ration 1172 cal./gr. 63.O Turkey starter ration 24 1404 cal./gr. 56.8 Turkey poult ration 28 1371 cal./gr. 46.7 Table 14. Caloric values and ratio of calories to percent protein of rations fed to grouse in the U.B.C. aviary from September, 1 9 6 3 through December, 1 9 6 4 . 53 Summary and Conclusions Blue grouse were not able to survive in the U.B.C. aviary on a diet of dried and pelleted f i r needles for longer than 2 weeks. Hen blue grouse held on chicken breeder ration (18 percent protein) had better survival and exhibited more reproductive behaviour than hen blue grouse held on turkey starter ration (24 percent protein) or turkey poult ration (28 percent protein. No conclusions applicable to natural conditions were made. CHAPTER 8 BASIC Nl/TRITION Introduction Once the basic parameters of the nutritional requirements of blue grouse are known, then experiments may be designed to test the effect of the quality of the diet upon growth, survival, egg production, and v i a b i l i t y of chicks. Materials and Methods Four grouse (3 females and 1 male) were held in the vivar-ium for nutritional studies. The conditions of holding were described in Chapter 2. The grouse were fed Buckerfield 1s chicken breeder ration. This ration has a protein level of 18 percent and a caloric value of 2578 calories per kilogram, or 2.58 calories per gram. Feed and water intake were measured daily. Two water pans were kept in empty cages and weighed daily to record evaporation. Droppings were collected on plastic sheets underneath the pens, dried in an oven and weighed un t i l the weight was constant. This point occurred after 2 days in the oven. The apparent diges t i b i l i t y of the feed was calculated as follows: Apparent % dig e s t i b i l i t y equals dry weight of feed consumed - dry weight of feces dry weight of feed consumed 55 The energy for maintenance was the number of digestible calories required by a bird of a particular weight per day at a particular temperature. The temperature at the vivarium when these data were collected was 72 to 75 degrees F. The basic water requirement was calculated as the number of grams of water consumed per digestible calorie. The daily intake of dry weight of feed and water for individual body weights was also calculated. Results Table 15 presents a summary of the data collected on body weights, diges t i b i l i t y of feed and the daily caloric and water consumption of grouse held at the vivarium. This data was col-lected in four series July 9 to 16, July 20 to 24, July 28 to 30, and August 2 to 6, 1965. The variation in d i g e s t i b i l i t y from 51.2 to 64 .9 percent was not s t a t i s t i c a l l y significant (analysis of variance at the 5 percent level of probability). There was a 6 percent variation from the mean body weight but a 30 percent variation from the mean digestible calories. The daily intake of digestible calories was not significantly related to body weight (linear regression at the 5 percent level of probability). Analysis of the daily water consumption showed i t was not directly related to the intake of digestible calories except for yearling male 1215 (linear regression at the 5 percent level of probability). The daily consumption of grams of water was d i -rectly related to body weight in a l l birds except adult female 1002 (linear regression at the 5 percent level of probability). BIRD AVERAGE AVERAGE AVERAGE RATIO OF AVERAGE DIGEST- DAILY DIGESTIBLE WATER CON-AVERAGE DAILY IBILITY WATER CALORIES SUMPTION TO NO. OF BODY -, CALORIC OF FEED CONSUMP- RECEIVED DIGESTIBLE SAMPLES WEIGHT INTAKE {%) TION DAILY CALORIES Yearling female 1105 Yearling female 1109 Adult female 1002 Yearling male 1215 19 20 864.2 946.1 69.7 82.0 19 915.3 74.3 51.8 4 5 . 8 51.2 64.9 19 965.5 98,5 61,9 54.8 37.9 57.2 36.2 41 .8 48.3 61.1 1.25 1.30 .79 .86 Table 15. Body weights, digestibility and caloric and water requirements of blue grouse held at U.B.C. vivarium "*"A11 weights in grams, o> 57 Discussion There was variation in the results of the diges t i b i l i t y study. Although the methods were f a i r l y crude, they were con-stant. Birds showed some variation in daily food consumption. They appeared healthy at a l l times during the collection of these data. There were no diarrhetic feces which might indicate sickness although some days they consumed only a few grams of feed. The average diges t i b i l i t y of 57-4 percent i s f a i r l y low compared to the 71 percent diges t i b i l i t y calculated using the same technique on rats (Schurch, Lloyd,and Crampton, 1950). The daily water requirement varied. I do not feel that error was incurred by the technique of measurement. If birds were at a l l sick, water consumption might change. There i s also the possibility that the water consumption may not be directly related to the daily intake of apparent digestible calories since only 1 of 4 showed a s t a t i s t i c a l l y significant relation-ship. The significant relationship between the number of grams of feed and the number of grams of water taken in per day, but the lack of a significant relationship between the intake of water and apparent digestive calories or the intake of apparent digestible calories to body weight suggest an error in calcula-tion of the dig e s t i b i l i t y . I do not know what caused the error, since as nearly as I was able to determine, the methods were constant. From Brody (1945:375) the basal metabolic rates for chic-kens of weights comparable to grouse are as follows: 53 WEIGHT BASAL METABOLISM, CAL/24 HR. .85 Kg. 65 .90 Kg. 70 .95 Kg. 75 The basal metabolic rate is equivalent to 85 percent of the maintenance energy (Brody, 1945:470). The weights of grouse, their daily caloric intake and their approximate maintenance energy requirement (as calculated from BrodyTs metabolic require-ments) are given in Table 16. From Table 16, the measured caloric intake for the grouse was lower than that required for maintenance according to Brody. As discussed before, the grouse in the vivarium were kept in small pens and were inactive. Possibly due to this lack of activity, they did not require as many calories for maintenance as BrodyTs figures suggest. It is also possible that the chic-kens tested might have had a higher metabolic rate. This com-parison suggests that the calculated dige s t i b i l i t y was low. Summary and Conclusions The apparent diges t i b i l i t y of the chicken breeder ration by the 4 grouse ranged from 51.2 to 64.9 percent. The mean was 57.4 percent. The variation was not s t a t i s t i c a l l y significant. The range of the daily requirements of apparent digestible calories at about 72 to 75 degrees F. was 35-3 to 6 6 . 0 . The range in body weight was 864.2 to 965.0 grams. The range of the calculated daily requirement of water was 37.9 to 57.2 grams. The consumption of water was directly AVERAGE WEIGHT OF GROUSE (in grams) AVERAGE MEASURED CALORIC INTAKE CALCULATED MAINTENANCE REQUIREMENT 864.2 69.7 70.5 915-3 74-3 82.3 946.1 8 2 . 0 88.2 965.5 88.2 98.5 Table 16. Weights of grouse, their daily-caloric intake and their approxi mate maintenance energy require-ment 60 related to the body weight of 3 of the 4 chickens. The consumption of water per digestible calorie varied from .79 to 1.30 grams. The consumption of water per digestible calorie was significantly related in only 1 of 4 birds. That ratio was .86 grams water per digestible calorie. CHAPTER 9 REPRODUCTIVE BEHAVIOUR Introduction This chapter is to present data on the breeding behaviour of blue grouse in the aviary and compare i t to the behaviour of blue grouse in the f i e l d . The 3 main areas of investigation were visual communication, auditory communication, and the mating display. Materials and Methods Observations on wild and captive blue grouse were recorded descriptively. The aviary was visited daily for at least 2 hours from May through August. It was visited at least every 2 days through the remainder of the year. Fred Gornall made films of the mating display of grouse in the f i e l d which were used for comparison to the display of captive birds. Vocaliza-tions were recorded on a Wollensack tape recorder at a tape speed of 7 1/2 feet per minute. The Wollensack microphone was either held by hand in the pen of the grouse, or fixed to the wall. The recordings were played to captive and wild grouse to observe their response. Results Visual Communication The sexual display (Appendix 2) of the male blue grouse is described in the literature (Bendell, 1954; Bendell and E l l i o t t , 62 1965; and Smith, I 9 6 0 ) . Plate 7 i s a photograph of the display. The sexual display was given only during the spring and early summer. There was no difference observed between the sexual display of adults and yearlings, between different years, between varying daily weather conditions, or between the sexual display in the aviary and that described in the literature. Males in the aviary sometimes displayed on the roost. However, most display was on the floor of the pen. On a few occasions females were observed giving the same sexual display as the males. Plate 8 is a photograph of the female display. The caruncle of the eye was yellow. The air sacs were not visible although the feathers on the neck were turned back. Auditory Communication Several vocalizations of blue grouse were recorded in the aviary. Te r r i t o r i a l males hoot during the reproductive season (Bendell and Elliot. 1 9 6 5). The males in the aviary also hooted during the spring. There was no hierarchy observed between the hooting males, in that one male did not stop hooting i f a second male began. If anything, the hooting of one male stimulated the hooting of other males since groups of males were often heard hooting together in the aviary. Bendell and E l l i o t t (1965) also observed in the f i e l d that the hooting of one male seemed to stimulate males on adjacent territories to hoot. For the period of May 1 through June 20 of each year the 63 aviary was visited daily, and at different times of the day. The number of males hooting each day were counted and summed. Table 17 presents the total number of observations of hooting by adult and yearling birds as a percent of the possible obser-vations of adults and yearlings hooting. From Table 17, the adult males hooted more frequently than yearlings. This difference was s t a t i s t i c a l l y significant (Chi-square test at the 5 percent level of probability). The incidence of hooting was calculated by summing the number of males hooting each day for 5 days and dividing by 5« This was done through the period April 20 through June 30 for 1963-65. The relationship of the occurrence of male hooting and female squatting and egg laying during the spring is given in Figure 5« Note that the period of squatting coincides with the period of peak hooting each year. In 1963 and 1965, an adult male began to hoot about a week before yearling males, while in 1964, a yearling male hooted for a week before the f i r s t adult. The f i r s t aggressive calls of males (Appendix 2) were ob-served about a week before the f i r s t hooting was heard each spring. The sexual display began and ended at the same time that hooting began and ended. The flutter-jump described for males in the f i e l d (Bendell and E l l i o t t , 1965; Blackford, 1963; and Wing, 1946) was not ob-served in the aviary. In the spring, yearling and adult hens usually gave a dis-tinct pre-copulatory cry, the whinny (Appendix 2 ) , when ready to 64 1963 1964 1965 TOTAL AD1 YR2 AD YR AD YR AD YR No. males available No„ of v i s i t s No. possible observations No. actual observations Percent of observations on which males hooted 3 15 51 51 153 765 46 33 30 4 14 9 51 51 714 459 53 13 7 2 2 9 51 51 102 459 36 5 35 1 19 33 153 153 969 1683 135 51 14 3 Table 17. Percent of total observations on adult and year-ling male blue grouse upon which males hooted, May 1 through June 20 , 1963-65 1Adults 2 Yearlings 65 c o o -C «/> o E c 0) u a> a. 20 10 21 April 20 10 21 20 1 0 0 2 May April May I April May 10 10 10 20 20 20 1963 31 10 June 20 30 1964 7 «•»."*:»'•.*•  vJv-.i-l? 31 10 June 20 3 0 1965 — —i h 31 10 20 30 June Figure 5. The relationship between the occurence of male hooting to female squatting and egg laying Legend- percent of males in aviary population hooting n m period of squatting of females f&nrzi period of egg laying 6 6 squat for copulation. It could be heard at any time during day-light hours. Whenever a female gave the whinny, reproductively active males immediately began to hoot or display or both. The whinny was played on the tape recorder to aggressive males in the aviary. In every instance, reproductively active males began to hoot or give sexual display or both. One male was continuously aggressive. However, every time the whinny was played during the display of intense of intense aggressive be-haviour, the male exhibited sexual behaviour and hooted. Hens in the aviary immediately stopped a l l movement and listened whenever a hen gave a whinny, or a whinny was played on the tape recorder. They looked in the direction of the source but rarely replied. Male grouse in the f i e l d responded to the whinny by hoot-ing and advancing in sexual display toward the speaker (Stirling and Bendell, 1 9 6 6 ) . It did not cause known individually banded males to leave their previously observed territories. Adult and yearling females that squatted or la i d eggs with-out squatting often gave a staccato cry during the breeding sea-son. It was called the female cry (Appendix 2). It was some-times heard when females were squatting, but more often when females exhibited aggressive behaviour. Females in reproductive condition quickly gave the cry in response to another hen or a tape recording of the cry. Males responded to the female cry by hooting, but with less intensity than they responded to the whinny. 67 No other calls related to breeding were observed. Mating Displays The sequence of acts of the male during the actual mating display observed in the U.B.C. aviary was essentially the same as described by Bendell and E l l i o t t (1965). In the aviary, the female either remains squatting through the series of male acts leading up to copulation, or walks around the pen very slowly. If the hen squats later, she does so when the male makes the short pre-copulatory run and gives a distinct pre-copulatory hoot. The pre-copulatory hoot sounds a l i t t l e like a single syllable of the hoot, but because the function is completely different i t is onomatopoetically called the "whoot" (Appendix 2) . Otherwise she runs or f l i e s away from the male when he be-gins the pre-copulatory run. If the female remains squatting after the male gives the pre-copulatory whoot, the male begins to tread her. He may f i r s t bob his head 1 to 10 times over a period of 5 seconds to 3 or 4 minutes, or he may tread immediately. The male takes 4 to 8 or 9 steps and settles with his feet approximately over the scapulae. The nape of the hen's neck i s taken in the cock's b i l l . The wings of the male clasp the sides of the female for balance. The male tips his t a i l to one side, the female tips her t a i l to the other side. In the matings ob-served, the t a i l of the male was always tipped to the le f t and the t a i l of the female to the right. The male lowers himself so that the cloacae may come together for copulation. The uterus 68 protrudes from the cloaca of the female and from the cloaca of the male protrude two copulatory papillae upon which sperm is carried. These are applied together. Copulation lasts about 2 to 5 seconds. Then, either the male steps off the back of the female or the female runs from underneath. Both birds stand s t i l l and ruffle their feathers for about 30 seconds. The male then be-comes extremely aggressive and viciously attacks the female. In the aviary, the grouse were always separated at this point. About half the naturally mated hens squatted again after copula-tion. Two naturally mated hens laid 9 f e r t i l e eggs after only one mating. One of these hens la i d a f e r t i l e egg 17 days after mating. Some of the following variations and peculiarities were observed. The male did not always give the pre-copulatory whoot prior to mating i f the female was already squatting at the time the male was introduced to the pen to begin his display. Females squatted in the presence of humans. Males in single pens some-times gave the pre-copulatory whoot when alone. Some pairs mated after a minute of display or less while some required forty min-utes of display or more before copulation took place. One female was selective about which male she mated with. Consistently she squatted only when a particular male was put in to her pen. In successive years the males were both yearlings. Discussion The reproductive behaviour observed in the aviary was 69 essentially the same as reproductive behaviour observed in the f i e l d . This suggests that the basic pattern of displays and calls related to reproductive behaviour are innate. The main function of the display of birds that come toget-her only for mating i s sex recognition (Marler, 1961). Display is also indicative of the bird's physiological condition (Marler, 196l ) o Blue grouse associate during the spring only for mating. Thus the primary function of the display would seem to be for sex recognition. The display also indicates the physiological states of both sexes. It has been demonstrated that the size of the eye caruncle of a male chicken partially determines the success of i n i t i a l encounters, that i s , the male with the larger comb i s more suc-cessful (Guhl, 1953). The eye caruncle of blue grouse changes from yellow to scarlet and expands as reproductive display be-comes intense. This suggests that the intensity of the color of the caruncle and the size may function in reproductive display to indicate the physiological condition of the male. Hamerstrom and Hamerstrom (I960) suggest that male t e r r i -t o r i a l calls in the prairie chicken (Tympanuchus cupido pinnatus) serve 3 functions, to advertise the general territory, to threat-en other rnales^and to attract females. Observations on the hoot-ing of male blue grouse support this general suggestion. The hooting of male grouse is of low frequency and d i f f i c u l t to l o -cate precisely by ear. The assumption i s made that i t i s equally d i f f i c u l t for a grouse to locate exactly by ear. The hooting of an individual male has been observed to affect the hooting and 70 movements of males in adjacent territories (Bendell and E l l i o t t , 1965). The fact that males begin to hoot immediately upon hear-ing the female whinny supports the suggestion that one function of hooting is to attract females. The whinny i s a c a l l of high frequency and easy to locate accurately by ear. The variation in pitch may also aid location. When the whinny was played in the f i e l d , males responded quickly and were able to locate the source of the sound accurately. This suggests the function of i t may be to direct the male to the fe-male when she i s ready to mate. Thus I suggest the following relationship between the hoot-ing of the male and the whinny of the female. Hooting advertises the presence and general location of the male. The female is at-tracted to the non-specific location of the male when she is ready to mate. The hen then gives the whinny, which directs the male to her. The function of the female cry i s uncertain. It was heard in the aviary when hens were ready to squat and also immediately prior to the attack of a hen upon i t s mirror image. It may be given as a result of conflicting drives or i t may have a func-tion in reproductive and aggressive behaviour. The pattern of the mating display was essentially the same as recorded by Bendell and E l l i o t t (1965) and filmed by Gornall. Some of the peculiarities such as a male treading a female with-out giving the pre-copulatory whoot may have been artifacts of captivity since i t has not been observed in the f i e l d . There was no difference observed between treading with or without the whoot. 71 Male chickens with a high reproductive drive and no re-leaser sometimes display and ejaculate while penned alone (C.V. Roberts, personal communication). Similarly, male grouse giving the pre-copulatory whoot while alone in their pens probably had a high reproductive drive and no releaser. This behaviour has not been observed in the f i e l d . The squatting of hen grouse for a human i s similar to sexual imprinting described for domestic fowl (Guiton, 1959; Guiton, 1961; and Klopfer, 1965). Some of these imprinted grouse mated with a cock i f a human was present. Hens squatted in response to the waving of a human hand. Imitation of the oot was not as effective for stimulating the female to squat. Possibly because the hen faces forward while squatting, any move-ment sufficient to be seen from the corner of the eye i s suf-ficient to act as a releaser for squatting behaviour. The hooting of males always began before any female repro-ductive behaviour was observed. In I965 the f i r s t hooting and squatting were both observed on May 1 (Figure 5 ) . Male aggres-sive behaviour was observed before May 1 in 1965 and i t i s pos-sible that hooting may have occurred earlier but was not heard. It is also possible that males may not hoot for extended periods at f i r s t and thus have less chance of being heard. Most squatting behaviour i s synchronized with the peak of hooting. This, plus the fact that hooting was usually observed f i r s t , suggests that hooting may be an ultimate stimulus to female reproductive behaviour. 72 Summary and Conclusions The data suggest that the reproductive displays are in-nate. The data suggest the hooting of the male serves to attract females to the territory of the male. Once attracted to the general area, the female gives a whinny c a l l to direct the male to her. Hooting may be an ultimate stimulus that releases repro-ductive behaviour in hens. Hen blue grouse imprint sexually upon humans but w i l l often mate with a cock grouse i f a human is present. Movement immediately behind a hen may be the stimulus necessary to cause that hen to squat. CHAPTER 10 AGGRESSION AND THE REPRODUCTIVE CYCLE Introduction The behaviour of t e r r i t o r i a l male blue grouse has been described in detail (Bendell and E l l i o t t , 1965). In short, males advertise their territories by hooting and defend them by threat displays and physical attack. Most non-territorial males are yearlings and do not contribute significantly to the breed-ing population. Females occupy a home range during the pre-nesting and nest-in periods and these home ranges are spaced (Bendell and E l l i o t t , 1965). It is uncertain whether or not these home ranges are mutually exclusive. If the home range of the hen i s exclusive and thus homologous to the territory of the cock, then I would make the hypothesis that hens would show some aggressive behaviour homologous to the aggressive behaviour of the cocks. Thus, I wanted to test hens in the aviary for aggressive be-haviour that might function in the f i e l d to space them during the reproductive season. The function of spacing of the home ranges of hens is discussed. Materials and Methods Data on aggressive behaviour was collected in several ways. Photographs, tape recordings and written descriptions were made. As a test for aggressive behaviour, male and female grouse 74 in single pens were presented with a mirror in which they could see their own image. Grouse reacted as much to their own mirror image as to a dummy, and dummies were too quickly mutilated to be used for extended periods. Live birds were not used because they were too frightened after being handled to act normally. The test were done in mid-morning once a week during the late winter and spring and twice a week from May through August. Since grouse are most active during the early morning and even-ing (Bendell, 1955) the tests were done in mid-morning to avoid recording this diurnal movement. Activity for the remainder of this chapter i s defined as any movement, posture, or c a l l given by a grouse in response to the mirror image. The number of hens tested was as follows. YEAR NO. OF ADULTS NO. OF YEARLINGS 1964 10 15 1965 9 11 The number of cocks tested was as follows. YEAR NO. OF ADULTS NO. OF YEARLINGS 1964 14 9 1965 2 9 A shorthand system of assigning a 1 or 2 letter symbol to 75 each posture, movement or c a l l was used to record each action the bird made while the mirror was in the pen. Every 5 seconds during the tests, an electric interval timer buzzed and a check-mark was made in the notes. Thus a l l movements were recorded chronologically and at known intervals of time. Data was collected for 3 minutes after the mirror was pre-sented to the grouse. This length of time was chosen after sev-eral extended tests indicated that a grouse reacted to i t s image within a maximum of 3 minutes. Usually grouse reacted within a few seconds. A l l activity was recorded. The symbols used for females were as follows. They are li s t e d in functions I think they serve. There is some duplica-tion. A detailed description of each is included in Appendix 2. Class Act Symbol Investigative head turn ht Threat neck stretch ns backward stance bs croak c a l l cr peck mirror image pk cluck c a l l c l Conflict male sexual display dis walk wa bleat c a l l bl puk-puk c a l l puk scratch scr stretch str 76 Class Act Symbol conflict pecking pkc Fear walk wa bleat c a l l bl Reproductive whinny c a l l wh cry c a l l cy squat sq The symbols used to record male behaviour w< Class Act Symbol Investigative head turn ht Threat neck stretch ns backward stance bs cackle c a l l ca cough c a l l CO peck mirror image pk Conflict female sexual display dis-f walk wa scratch scr stretch str conflict pecking pkc Fear walk wa Reproductive display dis hoot c a l l ho The data were analyzed by scoring one point for each sym-bol recorded. The total number of observations upon each type of movement, for a l l hens (or cocks) on each day was divided by 77 the number of grouse tested. This gave a numerical average that could be graphed from day to day. The total of a l l observations of a l l movements on one day was divided by the total number of birds tested, to give an overall mean number of movements for each grouse in the tested population. This totalled mean was used because a l l movement represented a reaction to the mirror image, whether i t was investigative, threat, conflict, fear, or reproductive. For convenience the totalled mean of the activity of the population is referred to as the "index" in the following discussion. The daily indicies were graphed. The values were tested with an analysis of variance and Duncan's Multiple Range Test for significant differences between points at the 5 percent level of significance. The periods during which hens squatted and l a i d eggs were plotted on the graph to check for correlation of periods of activity in reaction to the mirror image. The data for males was treated identically. Results Figure 6 gives the graphs of the daily indicies for the females through the breeding seasons of 1964 and 1965• The level of activity changed through the breeding season and i t was highest when the females squatted and l a i d eggs. The range of higher values (marked by horizontal dashed line) were s i g n i f i -cantly different from the range of lower values (marked by hori-zontal dashed li n e ) . The values between these lines were not significantly different from each other. 78 > o o X 80 60" 40-20 1964 0 1 — I 1 1 1 r -i 1 r 1 1 1 1 — 20 30 10 20 31 10 20 30 10 20 31 10 20 80 60 40 20 2 0 30 10 20 31 10 2 0 30 10 20 31 10 20 Figure 6. Indices of female activity during the breeding season Legend: mmm period of squatting period of egg laying 79 The mirror test was f i r s t developed and used to test the activity level of the population 3 days after squatting behav-iour began in 1964. Note the rise in activity in August/ 1964 compared to August,1965. The tests done in August, 1964 were done early in the morning when the grouse were more active. Be-cause the August 1964 data was not collected in the standard manner i t was not considered in a comparison of the 2 graphs. Thus the 2 graphs were compared for the period from 3 days after squatting behaviour began through 30 days after egg laying ceased (68 days). They were not significantly different (Sign test at the 5 percent level of probability). A l l behavioural characters were analyzed to determine which ones occurred in a repeatable pattern. The characters that showed a pattern a l l showed the same pattern as the graph. These characters were: head turn, neck stretch, backward stance, walk, and cluck. The incidence of the rest of the characters varied at random. Figure 7 gives the graphs of the daily indicies for the males through the breeding seasons of 1964 and 1965. The August, 1964 data was not considered because i t was collected at a dif-ferent time of day than the earlier data of that year. As in Figure 6, the horizontal dashed lines indicate the ranges of significant difference between indicies. The graphs of male indicies of 1964 and 1965 were tested over the same period as the graphs of female indicies were compared. They were s t a t i s t i c a l l y different (Sign test at the 5 percent level of probability). 1964 20 30 10 20 31 10 20 30 10 20 31 10 20 Figure 7. Indices of male activity during the breeding season Sl Discussion Hens show f i d e l i t y to a specific area for nesting (Zwickel, 1965)° As previously mentioned, hens occupy a home range during the pre-nesting and incubating period but i t is uncertain whether these home ranges are exclusive or defended. After the clutch hatches, hens and broods are observed at widely separated points and at times mixed with other broods (Bendell, 1955). The greatest reaction of hens to their mirror image was during the time that hens in the f i e l d would occupy a home range (Figure 6 ) . The index decreased past the end of egg laying to the low-est point 25 to 30 days after the last egg was la i d . This co-incided with the time the last egg would have hatched. After that period, the index fluctuated slightly but remained low. The possibility remains however that the index might have remained high i f hens had been allowed to incubate and hatch clutches. The index may have dropped from the lack of stimulus. I cannot deny this possibility as there was no control, but the evidence from the f i e l d suggests that interaction between hens at this time i s low. A strong dip in the value of the index during the squatting and laying period of I964 was repeated in I965 . There was no change in weather that might effect activity. As mentioned be-fore, these graphs are not different s t a t i s t i c a l l y . This sug-gests there may be a biological function for a lack of aggres-sive behaviour at this time. The only suggestion I can make i s that hens may be aggressive during the period of selecting a 82 nesting site, then become more docile for a period to allow a male to copulate, and then become more aggressive when laying and defending the nest site. There are data contrary to this reasoning. First, a l l hens do not exhibit the same display at the same time, that i s , some hens may be laying eggs when others begin to squat. Thus, one would not expect the population to give such a low index, secondly, the one natural mating that occurred in 1964 and 2 in 1965 a l l took place during the f i r s t peak of activity, not during the depression. The patterns of aggressive behaviour exhibited by cocks in the aviary are observed in the f i e l d at the corresponding time in the reproductive cycle. Correspondingly, the aggressive behaviour exhibited by hens in the aviary during the reproduc-tive period may also occur in the f i e l d . If i t does, then the home ranges of hens may be mutually exclusive during the nesting period and possibly defended. The nests of hens in the f i e l d are spaced rather than clumped. It has been shown in the wood pigeon (Columba palumbus) that spacing of nests functions for protection against predation (Murton, 1962). Thus the spacing of nests of blue grouse may serve the same function. The range of indicies of male behaviour show l i t t l e statis-t i c a l significance. This i s because there i s more individual variation between males than there i s between females. The dif -ference between the graphs of the male indicies of 1964 and 1965 is thus partly caused by individual males that were exceptionally aggressive. I cannot explain why the index for males in July, 83 1964 was so high. It was not as high in 1965. The July, 1964 index for females was not high. The lack of s t a t i s t i c a l l y signi-ficant results and lack of repeatable data between years reduced the value of the male aggression tests. Descriptive data on aggressive behaviour of cocks was the same as observed in the f i e l d and thus acted as a partial con-t r o l . This was valuable since i t is d i f f i c u l t to obtain observa-tions on female behaviour in the f i e l d for comparison to behav-iour observed in the aviary. Summary and Conclusions A relationship between the aggressive behaviour of females and the reproductive cycle was demonstrated in the aviary. The aggressive behaviour of males in the aviary and in the f i e l d was the same. Thus, the data suggest that the behaviour of females in the f i e l d may be the same as observed in the aviary. On the basis of aggressive behaviour observed in the aviary, t i seems possible that the home ranges of females in the f i e l d are mutually exclusive and may also be defended u n t i l the clutch has hatched. CHAPTER 11 WINTER AGGRESSION Introduction Blue grouse have a seasonal migration. They winter in the subalpine forests and breed on open alpine areas and lowland burns and logging slashes. Present data on the natural regula-tion of numbers of blue grouse show that more chicks leave the breeding range at the end of the summer than are required to compensate for adult mortality (Zwickel, 1965). This suggests either heavy mortality of chicks on the winter range or high-dispersal, since a surplus of yearlings is not observed on the breeding range in the spring. It has been suggested that winter t e r r i t o r i a l i t y of red grouse (Lagopus lagopus scoticus) regulates the number of red grouse available for breeding in the spring. Surplus birds are forced into marginal habitat and die there (Watson, 1964). The behaviour of blue grouse was studied at the aviary during the winters of 1963 through 1965 for evidence of winter aggression. Materials and Methods The big pen described in chapter 2 (Fig. 1, E) was used to study interactions of blue grouse. The number, age, and sex of birds put into the big pen in February 1964, were as follows. 85 SEX AGE NUMBER female yearling 2 female adult 2 male yearling 1 male yearling 1 Birds which died were replaced as required. The number, age, and sex of birds put into the big pen in October, 1964 for the winter of 1965 were as follows. SEX AGE NUMBER female yearling 1 female adult 1 male yearling 1 male adult 1 Birds which died were replaced as required. Results In a l l 3 years, 1 or 2 adult males in the aviary popula-tion showed aggressive behaviour and hooted from about the end of January through February of each year. After this period, there was no hooting until April or May, prior to the normal breeding season. In 1965, the adult male in the big pen quickly became 86 dominant. He was unable to f l y so that the yearling male was safe from attack while roosting. The yearling male was subordi-nate although he gave aggression calls from the roost, where he was most frequently seen. His sperm was viable but the adult male prevented him from displaying to the females. This year-ling male was dead in September, 1964. He was emaciated and there was no food in the crop or stomach. He probably starved to death. Yearling male 1286 was introduced to the big pen in October, 1964. The adult male followed the yearling for a few minutes but made no attack. The yearling was submissive. The adult remained dominant but allowed the yearling to feed. The adult male was f i r s t observed giving aggression calls on January 5, 1965. This was the last day the yearling was ob-served on the ground. By the end of January, the yearling was so hungry he flew from the roost to my arm to feed from my hand. He was dead of starvation on February 2, I 9 6 5 . His crop and stomach were empty and a l l body fat was gone. He weighed 998 grams. A second yearling male was placed in the big pen on Feb-ruary 6. He was selected because of large size. The adult male attacked the yearling male immediately. He was badly pecked during the next 2 days and was dead on February 9« Males were never observed to interact with the females during the winter. No interaction between females was observed during the winter. There was also no interaction observed during the f a l l . 87 Discussion Aggressive behaviour exhibited by males in the aviary dur-ing the spring has been observed in the f i e l d . Correspondingly, the aggressive behaviour observed in the aviary during the win-ter may also occur in the f i e l d . In the aviary, winter aggressive behaviour was only ex-hibited by adult males. Male red grouse are t e r r i t o r i a l during the winter. Non-territorial males are excluded to marginal habitat and die (Watson, I 9 6 4 ) . However, i t i s d i f f i c u l t to see how juvenile male blue grouse could be excluded to marginal habitat during the winter since there i s no evidence of any shortage of winter range. The fact there was no interaction during the f a l l suggests that simple crowding did not cause the interactions between grouse in the spring. The hens showed no aggressive behaviour during the winter. In late spring, 1964, one hen became very aggressive for several days, chased the adult male around the pen and viciously pecked him. No other interaction between the hens in the big pen was observed. Summary and Conclusions Aggressive behaviour was studied during the winters from I963 through 1965. Each year adult males were observed giving aggression calls or hooting during late January to February. One adult male in the big pen k i l l e d 2 yearling males in early February. BB This suggests the possibility of a period of winter aggres-sion similar to that observed in red grouse. No similar behaviour was observed in females. CHAPTER 12 SUMMARY 1. Blue grouse were h e l d i n c a p t i v i t y over a p e r i o d of 3 years t o f i n d the best methods of h o l d i n g and r e a r i n g them. In a d d i t i o n , data were obtained on the weights, s u r v i v a l , be-ha v i o u r , n u t r i t i o n , and d i s e a s e s of c a p t i v e blue grouse. 2. Grouse were h e l d i n wire pens t h a t measured 2 f e e t by 4 f e e t by 8 f e e t , 6 f e e t by 4 f e e t by 8 f e e t , and 20 f e e t by 10 f e e t by 24 f e e t . On the b a s i s of plumage c o n d i t i o n and r e p r o d u c t i v e performance the medium s i z e d pens were most s u i t a b l e . Weights and death r a t e s were the same i n a l l s i z e s of pens. 3. Captive grouse f e d on commercial chicken breeder r a t i o n l i v e d and maintained weight i n d e f i n i t e l y . The weights of the grouse i n the a v i a r y were comparable throughout the year t o weights of grouse i n the f i e l d . 4. The c o l d wet weather of win t e r a f f e c t e d the s u r v i v a l of grouse i n t h a t the number of deaths i n the win t e r was double the number of deaths i n summer. 5. The m o r t a l i t y of 1965 was e x c e p t i o n a l l y severe and c o r r e l a t e d w i t h the e x c e p t i o n a l l y c o l d wet w i n t e r of t h a t year. 6. Death r a t e s were the same among, (a) the t h r e e groups of c h i c k s brought t o the a v i a r y , (b) b i r d s of both sexes, and (c) b i r d s of each age group. 90 ? u The diseases that caused the greatest mortality were gizzard erosion, aspergillosis, and enteritis. 8. Feather-pecking by chicks was stopped by placing f i r boughs in the pens. 9. Reproductive display of males and females was less in the aviary than in the f i e l d . 10. The only successful method of mating grouse in the aviary was to put a displaying male into the pen of a squatting female. 11. Hens that exhibited reproductive behaviour were sexually imprinted upon humans. 12. The f e r t i l i t y and hatchability of blue grouse eggs in the aviary was 1/3 to 1/2 of that recorded from the f i e l d . 13» The mean clutch size of hens in the aviary was larger than the mean clutch size of hens in the f i e l d . 14. Blue grouse can be a r t i f i c i a l l y inseminated by using tech-niques developed for domestic fowl. Hens should be allowed to lay 2 or 3 eggs before insemination for best results in terms of continued egg laying. 15. The f e r t i l i t y and hatchability of eggs from hens that were a r t i f i c i a l l y inseminated were comparable to that of eggs from hens that were naturally bred in the aviary. 16. The combination of incubating grouse eggs in the battery incubators and brooding chicks by the methods of Zwickel (196$) was satisfactory. 17. Blue grouse were unable to survive in the aviary on a diet of dried and pelleted Douglas f i r needles. 18. Grouse t h a t were f e d a d i e t w i t h 18 p e r c e n t p r o t e i n had b e t t e r s u r v i v a l and e x h i b i t e d more r e p r o d u c t i v e b e h a v i o u r t h a n g rouse f e d a d i e t w i t h 24 or 28 p e r c e n t p r o t e i n . 19» The apparent d i g e s t i b i l i t y o f commercial c h i c k e n b r e e d e r r a t i o n by f o u r g r o u s e v a r i e d from 51.2 t o 64-9 p e r c e n t . The mean was 57.4 p e r c e n t . The v a r i a t i o n was not s t a t i s t i -c a l l y s i g n i f i c a n t . 20. The range o f t h e c a l c u l a t e d d a i l y w a t e r r e q u i r e m e n t was 37«9 t o 57.2 grams. The consumption o f wa t e r was d i r e c t l y r e -l a t e d t o body weight i n 3 of t h e 4 b i r d s . 21. A d u l t males i n t h e a v i a r y hooted more t h a n . y e a r l i n g males. 22. The f a c t t h a t males i n t h e f i e l d a r e a t t r a c t e d so q u i c k l y and a c c u r a t e l y t o t h e whinny su g g e s t s i t d i r e c t s t h e male t o t h e fe m a l e . 23. The s t i m u l u s t o squat i n t h e female i s l i k e l y movement be-h i n d and a t t h e same l e v e l as t h e hen. 24. A r e l a t i o n s h i p between t h e a g g r e s s i v e b e h a v i o u r o f f e m a l e s and t h e r e p r o d u c t i v e c y c l e was demonstrated i n t h e a v i a r y . T h i s p a t t e r n of b e h a v i o u r may a l s o o c c u r i n t h e f i e l d as a mechanism o f s p a c i n g d u r i n g t h e p e r i o d o f n e s t i n g . 25. Males became more a g g r e s s i v e d u r i n g t h e b r e e d i n g season and l e s s a g g r e s s i v e t h r o u g h t h e summer. 26. A g g r e s s i v e b e h a v i o u r was observed d u r i n g t h e w i n t e r i n a d u l t males. T h i s s u g g e s t s t h e p o s s i b i l i t y o f a p e r i o d o f w i n t e r a g g r e s s i o n s i m i l a r t o t h a t observed i n r e d gro u s e . 27. The f o l l o w i n g p o s t u r e s and c a l l s were more f r e q u e n t d u r i n g t h e r e p r o d u c t i v e season; f e m a l e s , h e a d - t u r n , n e c k - s t r e t c h , 92 backward stance, walk, cluck, whinny, cry, and squat; males, head-turn, neck-stretch, backward stance, peck, walk, cackle, cough, hoot, and whoot. 28. The following grouse calls were analyzed with sonograms: male hoot, female hoot, male cackle, male cough, female bleat, female whinny, female cry, and female cluck. LIST OF REFERENCES Allen, A.A., 1929, Ten years experiments in the rearing of ruf-fed grouse in captivity, Trans. Am. Game Conf., 16:3-21. Allen, G.A., 1964, Green Peafowl (Pavo muticus), Game Bird Breeder's Gazette, 8(1):10-14. Armstrong, E.A., 1950, The nature and function of displacement ac t i v i t i e s , Symp. Soc. Exp, Biol., 4:361-384. _____•> 1963, A Study of Bird Song, Oxford University Press, New York, 335 pp. Banks, E.M., 1954, Social organization in red junglefowl hens, Ecology, 37:239-248'. Batterson, W.M., 1957, The propagation of grouse, Game Bird Breeder's Gazette, 7(10):12-17, 50-51. Beer, J.B., 1943, Food habits of the blue grouse, Jour. Wildl. Mgmt., 7 (D :32-43 . Biely, J., E.A. Lloyd and I.W. Moynihan, 1950, Poultry Diseases, Province of British Columbia, Dept. of Agriculture, Bull. No. PB 108, 109 pp. Bendell, J.F. 1955a, Age, molt and weight characteristics of blue grouse, Condor, 57(6):354-36l. . , 19556, Age, breeding behaviour and migration of sooty grouse, Dendragapus obscurus fuliginosus (Ridgway) Trans. N. Am. Wildl. Conf. 20:367-38X ., and P.W. E l l i o t t , 1965, T e r r i t o r i a l behaviour and the regulation of breeding density in blue grouse, Can. Wildl. Service Series (in press). Blackford, J.L., 195$, T e r r i t o r i a l i t y and breeding behaviour of a population of blue grouse in Montana, Condor, 60(3):145-158. . , 1963, Further observations on the breeding behaviour of a blue grouse population in Montana, Condor, 65(6) :485-Borror, D.J., I960, The analysis of animal sounds, Animal sounds and communication, A.I.B.S. publication No. fm 26-37. 94 . , and C R . Reese, 1953, The a n a l y s i s o f b i r d songs by means o f a v i b r i l i z e r , W i l s o n B u l l e t i n , 65(4):271-276. Brody, S. , 1945, B i o e n e r g e t i c s and Growth, Hafner P u b l i s h i n g Co., New Yo r k , 1023 pp. Bump. G., R.W. Darrow, F.C. E d m i n s t e r , and W.P. C r i s s y , 1947, The R u f f e d Grouse, H o l l i n g P r e s s I n c . , B u f f a l o , New Y o r k , W5 PP. Buss, I.O., and B.A. S c h o t t e l i u s , 1954, B r e e d i n g age o f b l u e g r o u s e , J o u r . W i l d l . Mgmt., 18(1):137-138. R.D. Conrad and J.R. R e i l l y , 1958, U l c e r a t i v e e n t e r -i t i s i n t h e pheasant, b l u e g r o u s e and C a l i f o r n i a q u a i l , J o u r . Wildl. Mgmt., 22(4):446-449. C o l l i a s , N.E. and M. J o o s , 1953, The s p e c t r o g r a p h i c a n a l y s i s o f sound s i g n a l s o f t h e domestic f o w l , B e h a v i o u r , 5:175-187. Cowan, J.M., 1959, R e a r i n g g r o u s e , a c h a l l e n g e t o t h e game b r e e d e r , Game B i r d B r e e d e r ' s G a z e t t e , 10(2):49-54. D a l k e , P.D. et a l . , I960, S e a s o n a l movements and b r e e d i n g behav-i o u r of sage grouse i n Idaho, T r a n s . N. Am. W i l d l . Conf. 25:396-407. D a v i s , D.E., 1963, The hormonal c o n t r o l of a g g r e s s i v e b e h a v i o u r , P r o c . I n t . O r n i t h . Cong., 13:994-1003. D a v i s , I . L . , 1964, B i o l o g i c a l a c o u s t i c s and t h e use o f t h e sound s p e c t r o g r a p h , Southwestern N a t u r a l i s t , 9(3):118-145. D e l a c o u r , J . , 1959, Pheasant B r e e d i n g and Care, A l l - P e t s Books, Fond du L a c , W i s c o n s i n , 108 pp. E d m i n s t e r , F.C., 1947, The R u f f e d Grouse, I t s L i f e H i s t o r y , E c o l o g y and Management, The' M a c M i l l a n Co. , New Y o r k , 385 pp. E t k i n , E.T., 1964, S o c i a l B e h a v i o u r and O r g a n i z a t i o n Among V e r t e - b r a t e s , U n i v e r s i t y o f Chicago P r e s s , C h i c a g o , 307 pp. Evans, K.E., 1964, H a b i t a t r e q u i r e m e n t s of t h e g r e a t e r p r a i r i e c h i c k e n i n C o l o r a d o , U n p u b l i s h e d M.Sc. T h e s i s , C o l o r a d o S t a t e U n i v e r s i t y , 50 pp. Ewing, W.R., 1963, P o u l t r y N u t r i t i o n , 5th E d i t i o n , Ray Ewing Co., Pasadena C a l i f o r n i a , 1475 pp. Fay, L.D., 1963, Recent s u c c e s s i n r a i s i n g r u f f e d g r ouse i n cap-t i v i t y , J o u r . W i l d l . Mgmt., 27(4):642-647-F o w l e r , R.G. and D.I. Meyer, 1961, P h y s i c s f o r t h e E n g i n e e r and 95 Scientist, Allyn and Bacon, Inc., Boston, 553 pp. Gibson, G.G„, 1965, The taxonomy and biology of Splendidofilari-ine Nematodes of the Tetraonidae of British Columbia, Un-published Ph.D. Thesis, University of British Columbia, 235 pp. Grange, W.B., 1948, Wisconsin Grouse Problems, Wisconsin Con-servation Department, Madison, Wisconsin, 318 pp. Greenberg, D.B., 1949, Raising Gamebirds in Captivity, D. Van Nostrad Co. Inc., New York, 224 pp. Guhl, A.M. and L.L. Ortmann, 1953, Visual patterns of recog-nition of individuals among chickens, Condor, 55(4): 287-298. . , 1962, The Behaviour of Chickens, CHapter 17 of The Behaviour of Domestic Animals, E.S.E. Hafez, London, 857 pp. Guiton, P., 1959, Socialization and imprinting in brown leghorn chicks, Animal Behaviour, 7(1-2):26-34. ., 1961, The influence of imprinting on the agonistic and courtship responses of the brown leghorn cock, Animal Behaviour, 9(3-4):167-177-Hamerstrom, F. and Frances Hamerstrom, I960, Comparability of some social displays of grouse, Proc. 7th Ornith. Cong. 274-293. Hansen, N., 1961, Raising blue grouse, Game Bird Breeder's Gazette,-10(2):49-54. Hart, D., I964, Propagation of game birds, Gamebird Breeder's Gazette, 13(l):l8-22. H i l l , R., 1962, Raising prairie chickens, Game Bird Breeder's Gazette; 11(6):11-13 Hoffman, R.S., I 9 6 I , The quality of the winterfood of the blue grouse, Jour. Wildl. Mgmt., 25(2):209-210. Jenkins, D., 1961, Social behaviour in the partridge, Ibis 103: 155-188. •, A. Watson and G.R. Miller, Population studies on red grouse, Lagopus lagopus scoticus (Lath.) in Northeast Scotland, Jour. Anim. Ecol., 32:317-376. Kirkpatrick, CM., 1964, Age vs. environment as conditions for 96 reproduction i n caged bobwhites, Jour. Wildl . Mgmt., 28 (2) : 240-243. Klopfer, P.H., 1965. Imprinting: a reassessment, Science, 147(3655):302-303. Koivisto, I., 1965, Behaviour of the black grouse Lyrurus t e t r i x (L.) during the spring display, Finnish Game Paper 26, H e l s i n k i , 60 pp. Lacher, J.R. and D.D. Lacher, 1965, Blue grouse i n c a p t i v i t y , Jour. Wildl . Mgmt., 2 9 ( 2 ) : 4 0 4 - 4 0 5 . Lancaster, D.A., 1964, Biology of the brushland tinamou, Notho-procta cinerascens, B u l l . Am. Mus. Nat. Hist., Vol. 127, A r t i c l e 6, 314 pp. Lehmann, V.W., 1941, Attwater's p r a i r i e chicken, i t s l i f e h i s -tory and management, N. Am. Fauna 57, U.S. Govt. Printing Off i c e , 63 pp. L i , J.C., 1964, S t a t i s t i c a l Inference, Edwards Brothers, Inc., Ann Arbour, Michigan, 657 pp. Lumsden, H., 1961, Displays of the spruce grouse, Can. F i e l d Nat., 75(3):152-160. Marks, H.L., and P.D. Lepore, 1965, A procedure f o r a r t i f i c i a l insemination of Japanese q u a i l , Poultry Science, 4 4 ( 4 ) : 1001-1003. Marler, P., 1955, Characteristics of some animal c a l l s , Nature, 176:6-8. . , 1961, The evolution of v i s u a l communication, A Univer-s i t y of Texas Symposium, pp. 96-121. , and D. Isaac, I 9 6 0 , Physical analysis of a simple bird song as exemplified by the chipping sparrow, Condor, 6 2 ( 2 ) : 124-135. Marshall, W.H., 1946, Cover preferences, seasonal movements and food habits of Richardson's grouse and ruffed grouse i n southern Idaho, Wilson B u l l e t i n , 58(1) :42-51 . Moynihan, M. , 1955a, Remarks on the o r i g i n a l sources of displays, Auk, 72(3):240-246. . , 1955b, Types of h o s t i l e display, Auk, 72(3) :247-259-Murton, R.K. and A.J. Isaacson, 1962, The functional basis of some behaviour i n the wood pigeon Columba palumbus, Ibis , 104:503-521. 9 7 Nelson, M.M. and H.M. Evans, 1953, Relation of dietary protein levels to reproduction in the rat, Journal of Nutrition, 32:71-83. Pyrah, D., 1961, Sage grouse investigations, Job completion report, Idaho Dept. of Fish and Game, mimeo. 20 pp. Ralston Purina Co., 1963, Commercial game bird raising, Game Bird Breeder's Gazette, 1 2 ( 5 ) : 1 0 . . , 1964, Ulcerative enteritis in quail and grouse, Game Bird Breeder's Gazette, 13(2) :10-11 . Ripley, S.D., 1962, Important news about aspergillosis, Game Bird Breeder's Gazette, 1 1 ( 7 ) : 2 0 . ; Schaller, G.B. and J.T. Emlen, 1962, The ontogeny of avoidance : behaviour in some precocial birds, Animal Behaviour, ; 10(3-4):370-381. Schurch, A.F., L.E. Lloyd and E.N. Crampton, 1950, The use of chromic oxide as an index for determining the di g e s t i b i l i t y of a diet, Journal of Nutrition, 43:629-636. Simard, B.R., 1964, The testicular cycle of blue grouse (Den- dragapus obscurus fuliginosis) and i t s relation to age, breeding behaviour and migration, Unpublished M.Sc. Thesis, University of British Columbia, 113 pp. Simpson, G., 1935, Breeding blue grouse in captivity, Trans. Am. Game Conf., 21:218-219. Smith, N., I960, Molt patterns and breeding behaviour in cap-tive blue grouse, M.S. Thesis, Washington State University, 93 pp. Smith, N., and 1 .0 . Buss, I963, Age determination and plumage observations of blue grouse, Jour. Wildl. Mgmt., 2 7 ( 4 ) : 566-578. '~~'-Smith, W.J., 1963, Vocal communication of information in birds, American Naturalist, 47(893):117-125. Stewart, R.E., 1944, Food habits of blue grouse, Auk, 4 6 ( 2 ) : 112-120. S t i r l i n g , I. and J.F. Bendell, 1966, Census of blue grouse with recorded calls of a female, Jour, of Wildl. Mgmt., 30(1): (in press). Stokes, A.W., 1961, Voice and social behaviour of the chukar partridge, Condor, 63(2):111-127. 98 . , 1963, Agonistic and sexual behaviour in the chukar partridge, Animal Behaviour, 1 1 ( 1 ) : ; 2 ; - ; 3 4 . Taylor, W.L., 1949, F e r t i l i t y and Hatchability of Chicken and  Turkey Eggs, John Wiley and Sons, Inc., New York, 423 pp. Tinbergen, N., 1953, Social Behaviour in Animals, Methuen and Co. Ltd., London, 150 pp. . , 1959, Comparative studies of the behaviour of gulls, Behaviour, 1 5 ( 1 - 2):l - 7 0 . Watson, A., 1964, Aggression and population regulation in red grouse, Nature, 202:506-7. ., and D. Jenkins, 1964, Notes on the behaviour of the red grouse, British Birds, 57:137-170. White, J o C . , 1957, The American Grouse, Game Bird Breeder's Gazette, 6(2):14-18, 53-Wing, L., 1946, Drumming flight in the blue grouse and courtship characters of the Tetraonidae, Condor, 48(2)-.154-157. . , 1947, Seasonal movements of the blue grouse, North Am. Wildl. Conf., 12:504-510. Wood, E.D., 1956, Blue grouse, Game Bird Breeder's Gazette, 6(4):16-17. Wood-Gush, D.G.M., 1955, The behaviour of the domestic chicken: a review of the literature, British Journal of Animal Be-haviour, 111(3):81-110. . , 1956, The agonistic and courtship behaviour of the brown leghorn cock, British Journal of Animal Behaviour, 4:133-142. . , 1957, Aggression and sexual activity in the brown leg-horn cock, British Journal of Animal Behaviour, 5 :1-6. Zwickel, F.C., 1964, Early mortality and the numbers of blue grouse, Unpublished Ph.D. Thesis, University of British Columbia, 153 pp. APPENDIX 1 DISEASES OF GROUSE IN U.B.C. AVIARY Introduction This appendix is to catalogue the diseases identified in blue grouse held at the U.B.C. aviary. The signs and treatments for each disease are discussed. Sick grouse exhibited several external signs. A hunched posture was most characteristic (Plate 7). Sick grouse exhibited l i t t l e movement. Breathing was often forced and had a rasping sound. White diarrhetic feces were characteristic. Post mortem internal examination of many grouse showed several signs of disorder but none severe enough to have caused death. Several birds were taken to the Animal Pathology Labora-tory of the Federal Department of Agriculture but nothing defi-nite was isolated. The following i s a l i s t of the diseases recognised in the aviary. Aspergillosis Aspergillosis i s a fungal disease of the lungs and air sacs, caused by the fungus Aspergillus fumigatus or, rarely, A. niger. In extreme cases i t spreads throughout the viscera also. External symptoms are a hunched stance, loud rasping breath-ing and white diarrhetic droppings. Internally, cream coloured nodules are found throughout the lungs and air sacs. The air 100 sacs may become a solid lump of mold and nodules of fungus may occur throughout the viscera. Death occurs by suffocation when the fungi block the trachea. Treatment of aspergillosis has not been very successful. Biely et a l . (1950) state treatment i s of no value. Ripley (1962) used Amphoteracin B (Squibb) mixed with Alevaire in a spray to cur waterfowl. One gosling was successfully treated by spraying i t four times a day for 10 days in a plastic tent with an auxiliary supply of oxygen. This method was impracti-cal for use at U.B.C. W.G. Ford, Supervisor of Washington State Game Farms, found that zinc bacitracin, marketed under the brand name of Baciferm, reduced the incidence of aspergillosis on the State Game Farms (personal communication). This product was used at U.B.C. although no difference in the number of deaths due to aspergillosis was observed. The mechanism of infection i s inhalation of spores from moldy feces and feed. The aviary was sprayed regularly to reduce the mold. Gizzard Erosion Gizzard erosion i s a degeneration of the lining of the gizzard. The gizzard stones eventually go into the muscle. The bird stops eating and death i s by starvation. The causa-tive organism has not been isolated. External signs are hunched stance, white diarrhetic droppings and bad smelling breath. Internal examination reveals brown pussy scars of earlier infections that have healed. The 101 g i z z a r d s m e l l s f o u l . Treatment was w i t h T e r r a m y c i n and P o l y s o l i n d r i n k i n g w a t e r . U l c e r a t i v e E n t e r i t i s U l c e r a t i v e e n t e r i t i s i s an i n f l a m m a t i o n and hemorrhaging of t h e mucous membrane o f t h e i n t e s t i n e . I t i s not a s p e c i f i c d i s e a s e i n i t s e l f but i s t h e most common g e n e r a l c o n d i t i o n ob-s e r v e d i n p o u l t r y d i s e a s e s ( B i e l y et a l . , 1950). I t has been r e c o r d e d p r e v i o u s l y f rom c a p t i v e b l u e g rouse but no c a u s a t i v e o rganism has been i s o l a t e d (Buss, et a l . , 1958). The same e x t e r n a l s i g n s d e s c r i b e d f o r o t h e r d i s e a s e s a r e obs e r v e d . I n t e r n a l l y , t h e i n t e s t i n e s a r e f l a c c i d and wate r y . Upon s e c t i o n i n g , t h e l i n i n g i s found s l o u g h i n g and o f t e n b l e e d -i n g . T e r r a m y c i n and P o l y s o l were used i n t h e d r i n k i n g w a t e r . Success was f a i r where b i r d s were kept i n a l a b o r a t o r y f r e e o f "mechanical v e c t o r s o f d i s e a s e , o t h e r w i s e r e i n f e c t i o n o c c u r r e d . U l c e r a t i v e G i z z a r d T h i s i s t h e f o r m a t i o n o f a l a r g e p u s - f i l l e d u l c e r i n t h e g i z z a r d . The b i r d s t o p s e a t i n g and death i s by s t a r v a t i o n . I have o n l y f o u n d i t t w i c e (Table 3). No mention o f t h i s c o n d i -t i o n i s made i n t h e l i t e r a t u r e on p o u l t r y s c i e n c e . E x t e r n a l s i g n s were t h e same as f o r e n t e r i t i s and I c o u l d not d i a g n o s e i t s p e c i f i c a l l y . C o n s e q u e n t l y t h e t r e a t m e n t was th e same as d e s c r i b e d b e f o r e . 102 P r o v e n t r l c u l i t i s P r o v e n t r i c u l i t i s i s c h a r a c t e r i z e d i n t e r n a l l y by e x c e s s i v e s w e l l i n g o f t h e p r o v e n t r i c u l u s . The membranes and g l a n d s a r e i n f l a m e d . I t i s p r e v a l e n t i n c h i c k e n s r a i s e d i n confinement ( B i e l y et a l . , 1950). No c a u s a t i v e organism has been i s o l a t e d . T h i s c o n d i t i o n r e s u l t s i n poor d i g e s t i o n and t h e b i r d becomes em a c i a t e d . No d e f i n i t i v e e x t e r n a l symptoms ar e o bserved so t h a t d i a g n o s i s can o n l y be post-mortem. P r o v e n t r i c u l i t i s was o b s e r v e d s e v e r a l t i m e s , p a r t i c u l a r l y i n c h i c k s which had been b a d l y pecked when h e l d under crowded c o n d i t i o n s . Death was o n l y a t t r i b u t e d t o p r o v e n t r i c u l i t i s once. The s t r e s s which seems t o cause p r o v e n t r i c u l i t i s may be p r e v e n t e d by r a i s i n g b i r d s i n l e s s c o nfinement. I n t e r n a l Hemorrhage I n t e r n a l hemorrhage i s d i a g n o s e d by l o o s e c l o t s o f b l o o d i n t h e body c a v i t y . I t cannot be d i a g n o s e d e x t e r n a l l y . Hemor-r h a g i n g i s u s u a l l y due t o t h e r u p t u r e of b l o o d v e s s e l s i n t h e l i v e r , s p l e e n ; o v a r i e s , o r k i d n e y ( B i e l y et a l . , 1950). I t o c c u r s i n b i r d s w h i c h a r e nervous and f l y i n t o t h i n g s when s t a r t l e d . P r e v e n t i o n i s t w o f o l d , t o a v o i d s t a r t l i n g t h e b i r d s and t o h a n d l e them g e n t l y . APPENDIX 2 BEHAVIOUR Introduction The purpose of this appendix i s to describe the displays and cal l s of blue grouse observed in the U.B.C. aviary. This i s not intended to be a complete catalogue of blue grouse behav-iour. Only the behaviour discussed in the thesis i s explained. Photographs and sonographs are used for i l lu s t ra t ion . The sono-graphy were made on a Kay missilyzer, Audio and Sub-audio Spec-trograph. The methods are given by Borror ( I 9 6 0 ) , Davis (1964) and Marler and Isaac ( I 9 6 0 ) . An attempt to understand the func-tion i s given whenever possible. Investigative, agonistic and reproductive displays  Normal Stance (Plate 8) In the normal or neutral posture of blue grouse the body is upright, head s l ight ly forward, b i l l horizontal, wings beside the body and t a i l down. There is no display of a i r sacs or eye caruncle. This i s the posture blue grouse are observed in most of the time. It is not associated with any particular movements and' ho sounds are emitted." Head turning (Plate 9) Head turning is done by both sexes. The grouse stands in an upright position with the wings beside the body and t a i l down while watching the mirror image or other grouse. The head is 104 moved up, down or to the side and held there. It may occur as rapidly as one turn per second. The bird continues to watch. Usually there i s no sound given although males may give the cackle and females may bleat or cluck. It i s the most common movement seen in the aviary besides the normal stance and is seen at a l l times of the year. It seems to be mainly investigative. Neck stretching (Plates 10 & 11) This is the most common threat posture. The body may be upright with the neck vertical (Plate 9) or i t may be tipped forward with the neck horizontal (Plate 10). The t a i l i s down and the wings are held close to the body. No sound is given while this posture i s held. Male aggression cries and female clucks may be associated with neck stretching. It is often ex-hibited immediately prior to direct attack. It i s observed most often during spring and.early summer. The posture functions as a threat to other grouse. Simi-lar postures are taken by red grouse (Lagopus lagopus scoticus) Watson and Jenkins, 1964) and several species of gull (Larus sp. ) (Tinbergen, 1959). Backward stance (Plate 12) This is a very intense threat display used by both sexes. The body is upright and the neck i s arched over the back. The b i l l i s level, t a i l down, wings close to the body and neck feathers flattened. This threat display is used less than others. It i s most often seen during spring and early summer. No sounds 105 are given while standing in this posture. Male aggression calls and the female cluck are associated with i t . Sexual display (Plate 13) The male display i s mainly for sex recognition. It is only seen during the spring and early summer. It i s given dur-ing the courting of a female. The head is erect, wings slightly drooped, t a i l fanned, and air sacs expanded. The skin of the air sac i s orange and the feathers are white giving the general ef-fect of a fried egg. The eye caruncle i s expanded and yellow, changing to bright red as display becomes more intense. No sound is given. Males stand on sexual display for several min-utes at a time. Very similar displays have been described for the prairie chicken (Tympanuchus cupido pinnatus) (Hamerstrom and Hamerstrom, I960; Lehman, 1941) and sage grouse (Centro-cercus urophasianus) (Dalke, et a l . , I960). Display has been observed twice in female grouse (Plate 14). I do not understand the function in this instance. Head nodding This display i s given by the male during courtship. The male stands on f u l l display with his head in an upright position and looks at the female. He then quickly nods his head almost to the ground and back to the upright position. No sound is given. Individual nods may be from 5 seconds to a minute apart. A similar behaviour pattern was observed in the brushland tinamou (Nothoprocta cinerascens) and i t was suggested that i t might be a visual stimulatory mechanism for the female (Lancaster, 1964). 106 The function may be similar in the blue grouse since the nod is often given laterally or on an angle to the hen and displays the air sac. Presentation of air sac (Plate 15) The male displays laterally to the hen. The head and neck lower and go slightly forward. The eye i s just visible above the feathers surrounding the air sac. The air sac is then slowly expanded and retracted. No sound is emitted. It is a display of presentation and functions in sex recognition. Prai-rie chicken give a similar display (Hamerstrom and Hamerstrom, I960). Ta i l fanning (Plate 16) While displaying, the male sometimes turns and stands with his t a i l fanned toward the hen. This makes the black and white pattern very striking. No sound is emitted. The function seems to be sex recognition. It exemplifies the observation of Stokes (1961) that most of the male reproductive display is presentation. Strutting Strutting is a series of slow deliberate steps- taken by the male while in f u l l sexual display. The male may strut when alone but more frequently does so before a female. The feet are raised about 2 Inches off the ground and set down firmly. No sound is given. The wings and t a i l do not usually move from the normal display position. The cock may look around while 107 strutting. The function of strutting would seem to be sex recog-nition and possibly to stimulate the female to exhibit reproduc-tive behaviour. Waltzing (Plate 17) This is a term used to describe chicken behaviour (Wood-Gush, 1955). It refers to a displaying male drooping his wings slightly and making short runs of a few feet, toward or past a female. No vocal sounds are emitted but the primaries rattle as they drag on the ground. The function seems to be to stimu-late the female to begin reproductive display. The waltz usually ends behind the female so that i t may be to stimulate her to squat. It i s only seen during the spring and early sum-mer. Waltzing has been noted in chukar partridge (Alectoris  graeca) (Stokes, 1961) and prairie chicken (Hamerstrom and Hamerstrom, I960). Pre-copulatory run (Plate 18) This is a short run the displaying male makes toward the female, immediately prior to or while the female is squatting. The run i s usually on a curved path to behind the female. The male suddenly stops behind the female, tips forward with the momentum, and a cry called the pre-copulatory whoot is given. The cry i t s e l f i s described later. The eye caruncles are red. Prairie chickens have a similar run and c a l l (Hamerstrom and Hamerstrom, I960). The function seems to be to stimulate the female to squat. 108 Sgaut (Plate 19) This i s the posture the female assumes when ready to mate. It i s only seen during the spring. The legs are bent causing the body to lower, the wings are lowered with the t i p s on the ground f o r balance and the neck i s extended s l i g h t l y f o r -ward. No sounds are given. The function seems to be to stimu-l a t e the male to mount. A male was observed squatting once (Plate 20). I do not know what the function of t h i s was. Tread This i s the advance of the male upon the back of the female to copulate. The male takes 2 to 5 steps onto the center of the back of the female, takes the nape of her neck i n his b i l l and clasps her sides with his wingtips f o r balance. The t a i l s of the male and female are tipped to opposite sides to allow the cloacae to meet f o r copulation. Pecking This i s an act of direct aggression exhibited by grouse of both sexes, but more by males than females. The pecks are usually delivered on the head or neck. The neck stretch and backward stance are usually associated. The male usually gives aggression c a l l s . The female may cluck or give the croak c a l l . The function i s to drive the opponent away. Con f l i c t pecking This was exhibited by grouse of both sexes when there was no stimulus f o r a s p e c i f i c behaviour pattern. They would stand 109 e r e c t o r walk s l o w l y around t h e cage p e c k i n g t h e w a l l s , f l o o r or w a t e r f a u n t . No sounds a r e e m i t t e d . T h i s seems t o be d i s -placement b e h a v i o u r . Wing a t t a c k T h i s i s an a c t o f d i r e c t a g g r e s s i o n which may be g i v e n by grouse o f e i t h e r sex but p r e d o m i n a n t l y by males. The grouse u s u a l l y does a v e r t i c a l neck s t r e t c h , t a k e s a l i t t l e hop i n t o t h e a i r and d e l i v e r s t h e blow w i t h t h e wing. I t i s o f s u f f i -c i e n t s t r e n g t h t o be p a i n f u l t o a human. No v o c a l sounds accom-pany t h i s a c t i o n . I t may be preceded i n t h e case o f t h e male by a g g r e s s i o n c a l l s . No sounds have been h e a r d e m i t t e d by t h e female at t h i s t i m e . Grouse c a l l s and c r i e s A l l f r e q u e n c i e s a r e i n c y c l e s per second and a l l t i m e l e n g t h s a r e i n seconds. The fundamental and harmonics a r e de-f i n e d as f o l l o w s : "The l o w e s t f r e q u e n c y mode i s c a l l e d t h e fu n d a m e n t a l . H i g h e r modes a r e r e f e r r e d t o as ha r m o n i c s , " ( F o w l e r and Meyer, 1 9 6 1 : 3 7 6 ) . Male hoot ( P l a t e 21) The hoot i s h e a r d m a i n l y i n t h e s p r i n g d u r i n g t h e r e p r o -d u c t i v e p e r i o d . The male assumes an u p r i g h t s t a n c e , t a i l down, head f o r w a r d and a i r s a c s f i l l e d ( P l a t e 2 2 ) . The suggested f u n c t i o n i s t o a d v e r t i s e t h e t e r r i t o r y t o o t h e r males and a t -t r a c t f e m a l e s , as d i s c u s s e d i n c h a p t e r 9 . The song c o n s i s t s o f 7 s y l l a b l e s and l a s t about 2 .90 110 seconds (mean of 5 samples). The f i r s t s y l l a b l e i s the loudest and longest. The l a s t 2 s y l l a b l e s are run together and are d i f f i c u l t to distinguish. The fundamental and harmonics are run together so only the maximum and minimum frequencies of each s y l l a b l e are given. These are as follows. SYLLABLE 1 2 3 4 5 6-7 Frequency range 50-708 50-913 50-1000 50-831 50-675 50-530 Length of s y l l a b l e i n seconds .43 .20 .20 .20 .21 .23 Length of time between .46 .44 • 35 .27 • 41 s y l l a b l e s Male pre-copulatory cry The male pre-copulatory cry i s heard only i n the spring. It i s given to a female by a male exhibiting reproductive be-haviour. The male makes a short run toward the female, stops suddenly just behind the female, t i p s forward with the momentum and gives a low pitched "whoot." The function seems to be to stimulate the female to squat. It was not recorded on a tape so that no sonogram was made. Male cackle (Plate 23) The male cackle i s heard only during the spring and sum-mer. It sounds l i k e a staccato r e p e t i t i o n of "ca-ca-ca-. . . . " It i s given from an upright position (Plate 24) or while walking, I l l and functions as a threat c a l l . The c a l l has 8 to 10 syllables and lasts for about .87 seconds (mean of 20 samples). The frequency of the fundamental and f i r s t harmonic, length of time of each syllable and length of time between syllables are given on page 112. Male cough (Plate 25) The male cough i s heard only during the spring and summer. It i s a throaty, rough sounding, slower form of the cackle. It is also given as a threat. It i s not given as often as the cackle. It i s given from the same stance ; as the cackle. Upon analysis of the 1965 aggression tests i t was found given in a ratio of 1 :6.27 to the cackle. It has 7 to 8 syllables and lasts about 11.29 seconds ( mean of 11 samples). The frequency of the fundamental and f i r s t harmonic, length of time of each syllable and length of time between syllables are given on page 13. Female bleat (Plate 26) The bleat may be given by a hen in any stance except ag-gressive and at any time of the day or year.- Very intense bleating may occur prior to egg laying or when startled. Largely, I feel i t is a result of conflicting drives and possibly fear. The bleat i s about .26 seconds long (mean of 25 samples) and may occur about .47 to .56 seconds apart. The fundamental and four harmonics are consistently plain. The frequency of these in cycles per second are as follows (mean of 25 samples). SYLLABLE 1 2 3 4 5 6 7 8 9 10 Frequency of fundamental 50-650 50-680 50-640 50-635 50-630 50-635 50-620 50-595 50-543 50-453 Frequency of harmonic 1 1347-2512 1640-2667 1710-2707 1802-2822 1885-2795 1895-2850 1897-2780 2077-2692 1950-2517 1650-1850 Time length Of each syllable .16 .08 .06 .06 .05 .05 .05 .05 .05 .04 Time length between syllables .11 .02 .02 .02 .02 .02 .02 .01 .01 The frequency of the fundamental and f i r s t harmonic, length of time of each each syllable, and length of time between syllables of male cackle. SYLLABLE 1 2 3 4 5 6 7 8 Frequency' of fundamental 50-600 50-620 50-620 50-597 50-607 50-600 50-596 50-500 Frequency of harmonic 1 807-1432 812-1595 992-1585 825-1515 817-1474 805-1442 827-1452 900-1425 Time length of each syllable .18 .07 .07 .07 .06 .06 .06 .05 Time length of period between syllables .06 .05 .05 .06 .07 .09 .09 The frequency of the fundamental and f i r s t harmonic, length of time of each syllable, and length of time between syllables of male cough. 114 NOTE CYCLES/SECOND Fundamental 460-955 Harmonic 1 1 1 0 7 - 1 3 5 2 2 1592-2060 3 2322-2660 4 2859-3217 Female whinny (Plate 27) The whinny i s heard only during the spring or early sum-mer. It may be given at any time within a day of squatting and often immediately prior to squatting. Ths sound resembles the whinny of a horse but i s much higher pitched. Males in the aviary and in the f i e l d immediately respond to the whinny by hooting and giving reproductive display. A possible function is suggested in Chapter 10. The fundamental and at least 4 harmonics are broken up into several subnotes which make the sonograph d i f f i c u l t to measure. Thus only the length of the song, minimum and maximum frequencies and a photograph of the sonograph are given. The frequency ranges from 88 to 2105 cycles per second and the cry is about 10.5 seconds long (mean of 10 samples. Female cry (Plate 2 8 ) The female cry is only heard during the spring and early summer. It i s a staccato repetition of one basic note and sounds like "ka-ka-ka-. . . . " Note from Plate 26 that the intensity rises and f a l l s . It is sometimes associated with reproductive behaviour and sometimes with aggressive behaviour. I am uncertain 115 of i t s function. It i s of no definite length and has been recorded from 9 to 32 syllables. The length of the syllables is about .7 to .8 seconds and the time between syllables about .06 and .12 seconds (based on means of 7 c a l l s ) . The frequency of the fundamental i s between 450 to 85O cycles per second and the f i r s t harmonic is between 1375 to 2100 cycles per second. Female cluck (Plate 29) The cluck i s heard mainly during the spring and summer but has occasionally been recorded during the winter. It i s given by a female during aggressive display. It is often given prior to a direct attack. It sounds like the written word "cluck." It has at least 4 harmonics above the fundamental but be-cause the peculiar shape of the note makes a normal measurement impossible, only the minimum and maximum frequencies and a photograph of the sonograph are given. The c a l l ranges between 50 and 2270 cycles per second and i s about 2.34 seconds long (mean of 10 samples). Female puk The puk c a l l may be heard at any time of year. It i s given by a female sitting or standing s t i l l . It i s a low c a l l which sounds like "puk-puk-puk-. . ." rapidly repeated for several seconds. It has been recorded from 5 to 20 seconds. I was unable to make a tape recording. It was normally only heard when someone looked into the pen or f i l l e d the feeder. This suggests i t may be a fear reaction or possibly conflict. 116 Female croak The female croak was only heard during the spring. It i s low pitched sound much like the purr of a cat. It was only given during periods of aggressive behaviour when a female at-tacked her mirror image. It was not recorded on tape. Female hoot (Plate 28) Hooting by a female was only observed twice. Both oc-casions were in the spring. It sounded like the male hooting but had a higher pitch. One female did i t with no observed stimulus and one did i t on 3 separate occasions when she saw her mirror image. The female takes the same stance as a male but the eye caruncles do not become inflated and the air sacs are not exposed. Only 2 series of female hooting were recorded. It lasted about 1.7 to 2.1 seconds. The frequency range i s from about 50 to 2250 cycles per second. The syllables were from .09 to .39 seconds long and from .20 to .39 seconds apart (mean of 2 samples). 117 APPENDIX 3 • i r B i n m t n m i i i m m m m n n n f ^  n n P l a t e 1. U.B.C. a v i a r y from south, P l a t e 2. I n s i d e o f a pen a t U.B.C. a v i a r y t o show p o s i t i o n o f water f a u n t s , f e e d e r s , d i v i d e r s and r o o s t . P l a t e 3. E l e c t r i c w e i g h i n g s c a l e w i t h b asket 118 119 Plate 9. Head turn posture. Plate 10. vertical. Neck-stretch posture, 120 Plate 13. Male sexual display. Plate 14. Female giving male sexual display. 121 Plate 17- Male waltzing. Plate 18. Male giving pre-copu-latory whoot at termination of pre-copulatory run. P l a t e 21. Sonogram of male hooting. P l a t e 23. Sonogram o f male c a c k l e a g g r e s s i o n c a l l . SECONDS P l a t e 26. Sonogram of female bleat. 4000 a 2 S 4-.00O a: 3,000 1/5 2,000 lOOO IT /»</ Plate 27. Sonogram of female whinny, SECONDS Plate 29. Sonogram of female cluck. ' v ' fntf ' O £0001 48 -urf" T ^ 1 3 4,000 SECONDS Plate 30 . Sonogram of female hooting. 

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