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Ecology of a partially migratory elk population Woods, John G. 1991

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ECOLOGY OF A PARTIALLY MIGRATORY ELK POPULATION by JOHN G. WOODS B.Sc, University of Guelph, Guelph, 1972 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES (Department of Zoology) We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA July 1991 ® John G. Woods 1991 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of The University of British Columbia Vancouver, Canada Date /if/ - J ^ Z T 3 0 DE-6 (2/88) ABSTRACT In this thesis I investigate the ecology of a p a r t i a l l y migratory (<100% of the animals migrate) population of elk (Cervus elaphus) in the Canadian Rockies. I radio-tagged elk in a 330 km2 area of the Bow River v a l l e y (BRV) in Banff National Park, Alberta and followed them for 36 months. Elk movements to adjacent watersheds expanded the study area to 2900 km2. My goals were to describe the seasonal movements made by BRV elk and to reach some understanding of the causes of these movements. The M/R (migrant/resident) ratios for adult b u l l s and adult cows were 4 . 3 and 0 . 5 respectively. C l a s s i f i e d ground counts revealed that adult bulls made up only 11% of the population and that the overall M/R r a t i o for the population was 0 . 6 . Migrations did not take elk beyond the foraging range of timber wolves (Canis lupus), their p r i n c i p a l predator. Three cows changed migration status between years and some migrants were sympatric with residents during the rutting (breeding) season. These observations are consistent with the hypothesis that migration in elk i s a conditional ESS (evolutionarily stable strategy). Although 1 adult radio-tagged b u l l dispersed, individual annual home-ranges of the remaining elk overlapped from year-to-year. Elk were strongly p h i l o p a t r i c to winter, summer, and rutting ranges. There were no differences detected in the r e l a t i v e philopatry of bulls and cows, or of migrants and residents. Bulls had average 1-way migrations of 30 km ho r i z o n t a l l y and 840 m v e r t i c a l l y . Cows had average 1-way migrations of 36 km horizontally and 1079 m v e r t i c a l l y . The net energy and time investments for these movements were calculated and judged to be t r i v i a l . Elk on both high and low elevation ranges ate primarily grasses and sedges during the autumn, winter, and spring, and the leaves of deciduous shrubs during the summer. S i m i l a r i t i e s outweighed differences between high and low elevation ranges. Elk population c h a r a c t e r i s t i c s (numbers, composition, survival, recruitment, predation, parasites, animal morphology) were measured during 1 9 8 5 - 9 1 and compared with similar data gathered during 1 9 4 4 - 6 9 . In most respects, the population has not changed over these years and l i t t l e i s known of density-dependent processes. i i Table of Contents Abstract i i Table of Contents i i i L i s t of Tables v L i s t of Figures v i i Acknowledgements v i i i CHAPTER 1 . GENERAL INTRODUCTION 1 Study Area 5 CHAPTER 2 . PARTIAL MIGRATION 11 Introduction 11 Methods 15 Radio-tagging 15 Data analysis 18 Results 20 Seasonal movement patterns 20 Migrant-resident ratio and sex biases 24 Survival 24 Discussion 33 Seasonal movement patterns 33 Mixed ESS 34 Learning 36 Migrant/resident ratio 37 CHAPTER 3 . PHILOPATRY 41 Introduction 41 Methods 44 Results. 46 Overlap of consecutive annual home-ranges 46 Distances between inter-annual seasonal ranges.... 46 Discussion 55 Seasonal ranges 55 Definitions of philopatry 57 CHAPTER 4 . NET COSTS OF MIGRATION 60 Introduction. 60 Methods 62 Energy calculations 62 Time and distance estimates 63 Body weights and sex/age classes 64 Results 65 Migration time and speed 65 Body weights 65 Net costs 65 Discussion 70 Speed of travel 71 Relative energy costs 72 i i i CHAPTER 5. DIET COMPOSITION AND QUALITY 75 Introduction . 75 Methods 78 Fecal c o l l e c t i o n s 78 Forage co l l e c t i o n s 80 Estimates of diet quality 81 Results 82 Diet composition 82 Forage quality indicators 86 Discussion 93 Diet composition 93 Forage quality 94 Diet quality 95 Nut r i t i o n a l advantage hypothesis.. 96 CHAPTER 6. POPULATION CHARACTERISTICS... 98 Introduction 98 Methods 101 H i s t o r i c a l data 101 Population estimates 101 Recruitment 103 Mortality 104 Animal condition indices 105 Results 107 Population estimates 107 Recruitment 107 Survival 109 Parasites 110 Body and antler weights .111 Discussion 122 Population estimates 122 Sex ratios and longevity 123 Body and antler weights 124 Parasites 125 Recruitment and survival 126 The future of the BRV elk population 128 CHAPTER 7. CONCLUSIONS 131 LITERATURE CITED 137 APPENDIX I 148 iv L i s t of Tables Table 1. Migrant/resident ratio for radio-tracked adult elk in Banff National Park, 1986-88... 26 Table 2. Horizontal and v e r t i c a l movements of migrant and resident radio-tracked adult elk in Banff National Park, 1986-89 27 Table 3. Median migration dates for radio-tacked adult elk in Banff National Park, 1986-88... 28 Table 4. Annual survival rates and 95% confidence intervals for migrant and resident radio-tagged adult elk in the BRV, 1986-89.. 29 Table 5. Migrant/resident ratios, hunting status, and timber wolf status for various elk populations in the Rocky Mountains 30 Table 6. Numbers of migrant and resident radio-tracked elk in Banff National Park, 1986-88 48 Table 7. Overlap of annual home-ranges during consecutive years for radio-tracked elk in Banff National Park, 1986-88 49 Table 8. Distances between inter-annual home-range centres during consecutive years for radio-tagged elk in Banff National Park, 1986-88 50 Table 9. Number of radio-tagged elk within and outside the BRV during the rutting seasons, 1986-88 51 Table 10. Daily travel distances of resident elk compared to d a i l y travel distances of migrant elk during migration, May and June, 1988 67 Table 11. Net 1-way travel times and energy requirements for migrant elk based on annual movements of migrant and resident radio-tracked elk 68 Table 12. Horizontal and v e r t i c a l distances reported for migrating elk and other North Temperate cervids. . 74 Table 13. Major components of elk diets during 4 seasons on subalpine and montane ranges 84 Table 14. Comparison of 6 measures of forage qu a l i t y means during the period May-September, 1988 for 1 subalpine and 2 montane sit e s 89 Table 15. Early spring elk population estimates based on ground counts, a e r i a l counts, and mark-recapture, BRV, 1985-90 112 Table 16. Autumn elk population estimates based on ground counts corrected for ground-aerial bias during 3 periods: 1944-53, 1959-68, and 1985-90 112 v Table 17. C l a s s i f i e d elk count ratios and 95% confidence intervals of c a l f , 1 year-old b u l l , and mature b u l l elk in the BRV expressed per 100 adult cows, 1943-90 113 Table 18. Bull/cow ratios of BRV elk during 1985-89, 1957-64, and 1944-54 114 Table 19. Deaths of adult elk reported from a l l sources and from radio-tagged animals only, February 1986 - A p r i l 1989 115 Table 20. Annual survival rates and 95% confidence intervals for radio-tagged adult elk i n the BRV, 1986-89 116 Table 21. Cumulative survival estimates from b i r t h to age 24 months for 6 cohorts of b u l l elk based on c l a s s i f i e d counts, 1985-90 117 Table 22. Prevalence of giant l i v e r flukes, lungworms, and hydatid cysts in BRV elk during 3 periods: 1944-54, 1958-66, and 1985-89 118 Table 23. Mean body weights and 95% confidence intervals for r a i l - k i l l e d and road-killed elk, BRV, 1985-89 119 Table 24. Antler weights and 95% confidence inte r v a l s for r a i l - k i l l e d and road-killed elk, BRV, 1985-89 120 v i L i s t of Figures Figure 1. The study area 10 Figure 2. Examples of elevational movements of 8 radio-tagged elk, BRV, 1986-88 31 Figure 3. Mean elevation of resident and migrant radio-tagged elk during winter, summer, and rut, BRV, 1986-88 32 Figure 4. Distances between centres of consecutive winter ranges 52 Figure 5. Distances between centres of consecutive summer ranges 53 Figure 6. Distances between centres of consecutive rutting ranges 54 Figure 7. Net cost of an upslope migration for various weight classes of BRV elk 69 Figure 8. Grass and deciduous shrubs in elk diets collected in subalpine and montane habitats, 1988 85 Figure 9. Nitrogen in elk fecal samples from montane and subalpine s i t e s in Banff National Park, 1985-88 90 Figure 10. Nitrogen, c e l l solubles, and l i g n i n in montane and subalpine forages and estimated diets, Banff National Park, 1988 91 Figure 11. Calcium and phosphorus in montane and subalpine forages and estimated elk d i e t s , Banff National Park, 1988 92 Figure 12. Corrected population estimates, 1944-89.... 121 v i i Acknowledgements This research was supported f i n a n c i a l l y by Public Works Canada, Edmonton; the Canadian Parks Service, Calgary, Revelstoke, and Banff; and the National Science and Engineering Research Council, Ottawa. I would l i k e to thank Bruce Leeson of the Canadian Parks Service for suggesting the project and A.R.E. S i n c l a i r , C. Walters, J. Smith, and D. Shackleton of the University of B r i t i s h Columbia and D. Houston of the US National Park Service, for p a r t i c i p a t i n g on the author's graduate committee. H. Flygare, C. Elverum, and P. Harris,, braved the elements at a l l seasons and provided the technical assistance necessary to follow the elk. The Banff warden service assisted in every aspect of the f i e l d work and in p a r t i c u l a r I would l i k e to thank the senior w i l d l i f e wardens: R. Kunelius, T. Skjonsberg, and B. Browne. P. Jacobson of the Banff Warden Service made the f i e l d data from a previous elk telemetry study in the Bow River va l l e y available to me. Park Superintendents R. Beardmore and W. Gallacher encouraged me to pursue these studies. Mas Matshushita of the Canadian Parks Service drafted the f i n a l figures. v i i i CHAPTER 1. GENERAL INTRODUCTION Migration in birds and mammals i s generally defined as the annual return movement of individuals between 2 geographically separate areas. It i m p l i c i t l y follows that migratory animals establish a degree of philopatry to seasonal ranges and make r e l a t i v e l y l i t t l e use of areas between these ranges. In contrast, dispersers make one-way movements and nomads are not ph i l o p a t r i c to seasonal ranges. Although many species of animals have been c l a s s i f i e d as either migrants or non-migrants, i n t r a s p e c i f i c v a r i a t i o n in seasonal movement patterns i s common (Baker 1978). P a r t i a l migration occurs when a population contains both migrant and sedentary individuals, and d i f f e r e n t i a l migration when individuals within a population migrate varying distances (Ketterson and Nolan 1985, Lundberg 1988). P a r t i a l migration has received considerable attention in bird migration studies in the context of evo l u t i o n a r i l y stable strategies (ESS's). Maynard Smith and Price (1973) defined an ESS as: "a strategy such that, i f most of the members of a population adopt i t , there.is no 'mutant' strategy that would give higher reproductive f i t n e s s . " However, i f seasonal movement behavior i s an ESS, how i s that both migrants and residents can coexist within a single population? Three solutions have been advanced: 1) a mixed ESS at the population l e v e l (with equal fitness of migrant and resident morphs and i n f l e x i b l e individual behavior); 2) a conditional ESS in which morphs need not have equal fi t n e s s and can change their behavior (Lundberg 1988, Adriaensen and Dhondt 1990); and, 3) an unstable movement strategy ( i . e . in transition) (Swingland 1983). The observation that migratory restlessness i s a heritable t r a i t in some birds (Berthold and Querner 1981, Biebach 1983) suggests the existence of resident and migrant genetic morphs and i s consistent with a mixed ESS explanation. However, a conditional ESS solution i s supported by the fact that timing and distance of migration i s facultative in some birds ( T e r r i l l and Ohmart 1984), and that individuals may switch between migration and residency (Baker 1978). Adriaensen and Dhondt (1990) explained this range of behaviors by suggesting that a within a population, individuals may have varying genetic biases. Those strongly predisposed to migrate, always do so; those with l i t t l e migratory i n c l i n a t i o n , never migrate; and, those between these extremes, make a conditional assessment of the environment and decide to migrate or not. A genetic basis for migration has not been demonstrated in mammals. In large mammals, explanations of p a r t i a l migration emphasize the importance of learning in the development of seasonal movement 2 behavior (McCullough 1985). In this view, seasonal movement strategies of individuals would not be "closed program" ESS's in a genetic sense, but rather the product of cu l t u r a l inheritance and individual experience. As Mayr (1974) argued, retention of phenotypic f l e x i b i l i t y enabled by "open programs" would be of selective advantage for non-communicative behaviors such as habitat use in a varying environment. Elk are often c l a s s i f i e d as migrants, despite the observation of coexisting non-migrant populations and populations with a mix of movement strategies. Morgantini (1988) suggested that v a r i a b i l i t y in movement behavior by elk i s an exhibition of the species's f l e x i b i l i t y in coping with a varied environment and Boyce (1991) showed that seasonal fluctuations in the a v a i l a b i l i t y of food resources could determine the fitness of migratory and non-migratory elk populations. To-date, no studies have d i r e c t l y compared the ecology of migrant and sedentary individuals from a single population. In this thesis, I examine a p a r t i a l l y migrant elk population in the Canadian Rockies and address the general hypothesis that elk seasonal movements are conditional strategies ( i . e . individually-based decisions). I f i r s t introduce the problem and study area (Chapter 1 ) , I then examine the movement cha r a c t e r i s t i c s of the population (Chapter 2 ) , the 3 r e l a t e d t o p i c of p h i l o p a t r y (Chapter 3), the net c o s t o f m i g r a t i o n (Chapter 4), and the d i e t of migrants and r e s i d e n t s (Chapter 5). I conc lude by d e s c r i b i n g some p o p u l a t i o n parameters (Chapter 6) t h a t might be i n f l u e n c e d by movement s t r a t e g i e s . 4 Study area This study was centred in a portion of the Bow River v a l l e y (BRV) between Lake Louise and Canmore, Alberta in the Continental Ranges of the Canadian Rocky Mountains (Figure 1 ) . The valley bottom elevation increases from 1341 m at Canmore to 1646 m at the Continental Divide. Approximately 330 km2 of the BRV i s below 1700 m elevation and i s used by elk year-round. Migratory movements of elk from the BRV expanded the study area to 2900 kra2. The BRV valley floor runs for approximately 80 km within Banff National Park and 4 km outside the park, and varies in width from 2 -6 km. Mountains rise to elevations of 3000+ m on either side. This area of the Rockies has a continental climate characterized by r e l a t i v e l y long winters and short summers (Holland and Coen 1 9 8 3 ) . Below 1600 m, low snowfall coupled with sudden warming periods i n winter, results i n intermittent snow-free periods. Above 1600 m, snowpack i s t y p i c a l l y continuous from November through May or June except on very steep slopes or windswept ridges. During the study ( 1 9 8 5 - 8 9 ) , snowfall in the BRV was below average (58-83% of the long-term average of 2 4 8 . 9 cm). The vegetation in and adjacent to the BRV has d i s t i n c t a l t i t u d i n a l zonation (Holland and Coen 1 9 8 3 ) . The montane zone ( <1600 m) i s characterized by mixed forests of evergreen trees (Pinus contorta, Pseudotsuga 5 menziesii, Picea glauca) and aspen (Populus tremuloides) interspersed with natural grasslands. Approximately 200 km2 of the BRV i s in the montane. The lower subalpine zone (1600-2000 m) has closed evergreen forests (Pinus  contorta, Pseudotsuga menziesii, Abies lasiocarpa, Picea  Engelmannii). Higher in this zone (2000-2300 m), the evergreen forest i s interspersed with subalpine meadows. The alpine begins above 2300 m. In many places, avalanches descending from the alpine and upper subalpine have created openings through the lower forests. The Trans-Canada Highway and Canadian P a c i f i c Railway roughly p a r a l l e l the Bow River through the study area. Both are major transcontinental transportation routes with large t r a f f i c volumes (Woods 1990). There are approximately 116 km of additional roads within the BRV. During 1983-87, 26 km of the easternmost section of the highway within the park were fenced on either side with 2.4 m high, ungulate-proof fencing. The remainder of the highway (58 km) was not fenced. Underpasses and bridges provided ungulates opportunities to cross beneath the fenced highway at 12 locations. During the study, highway t r a f f i c peaked during July and August (26% of annual total) and was lowest during November (5% of annual total)(Woods 1990). A town (Banff) and a small service centre (Lake Louise) are located within the BRV and the town of Canmore i s 6 located at the eastern edge. In addition to developments within the towns, there are numerous hiking t r a i l s , p i c n i c areas, campgrounds, a grass-surfaced a i r f i e l d , a golf course, and several lodges. Three downhill ski resorts (Mount Norquay, Sunshine V i l l a g e , Lake Louise) are located d i r e c t l y above the BRV i n the subalpine and alpine zones. Overviews of the elk biology in the park have been presented by Flook (1967, 1970), Holroyd and Van Tighem (1983), Morgantini (1988), Woods (1990) and Chapter 6. Although native elk were found in BRV when the park was established in 1885, they had disappeared by 1906. In 1915, native elk reappeared in the park as summer migrants from B r i t i s h Columbia. During 1918-20, 235 elk from Yellowstone National Park were released at 2 locations within the BRV (Lloyd 1927). From 1941-69, nearly annual elk slaughters were conducted in and around the BRV. Since the 1970's, management interest i n BRV elk has focused on elk-vehicle c o l l i s i o n s on park roadways and on the railway (Woods 1990 and Chapter 6 ) . BRV elk were v i s u a l l y tagged as early as the 1960's. In the early 1980's, park wardens radio-tagged 7 adult cows and observed both resident and migrant elk (P. Jacobson, pers. comm.). Dispersal from the BRV and other remnant populations i s believed to have established the Red Deer River population in the north-east of the park during the 1930 fs (Morgantini 7 1 9 8 8 ) . Radio-tracking and visua l tagging programs in the adjacent Kootenay National Park documented dispersal into the BRV from the Kootenay River valley (Gibbons 1 9 7 8 ) . One of these dispersers subsequently moved to the Red Deer valley, a distance of approximately 150 km (Morgantini 1 9 8 8 ) . Elk were the most abundant ungulate in the BRV during the study ( 1 9 8 5 - 9 0 ) with an average population of approximately 900 animals (Woods 1 9 9 0 , Chapter 6 ) . Based on c l a s s i f i e d count data and the observation that few, i f any 1-year-olds breed (Chapter 6 ) , the ef f e c t i v e population size (Ne) was approximately 450 (Chepko-Sade and Shields 1 9 8 7 , sex-ratio method). An additional population of 300 elk were estimated to winter in the area immediately east of the BRV (T. Nette, Alberta W i l d l i f e , unpubl. data). Since the majority of the BRV i s within the national park, and since most migrant elk move to other etreas of the park, this population i s largely unhunted. Bow-hunting for elk i s allowed during annual open seasons on prov i n c i a l lands in the portion of the BRV between the park and Canmore. Those elk migrating out of the BRV via the Spray River watershed pass through an area of prov i n c i a l land with an annual r i f l e - h u n t i n g season for elk. Predation by timber wolves and c o l l i s i o n s with trains and automobiles are the major mortality sources 8 for this population (Weaver 1979, Woods 1990, Chapter 6 ) . Coincident with the start of this research, the wolf population apparently increased within the BRV (R. Kunelius, CPS, pers. comm.). Since the 1940's, habituation to people has been a notable feature of BRV elk (Green 1949). Although both sexes are r e l a t i v e l y i n d i f f e r e n t to human presence, cows are noticeably more nervous. Some adult b u l l s regularly spend summer and winter within the urban boundaries of Banff townsite. Between the park and Canmore, elk are noticeably more wary of humans and adult elk are rarely found within the town of Canmore. 9 ALBERTA Figure 1. The study area. (l=village of Lake Louise; 2=town of Banff; 3=town of Canmore) 10 CHAPTER 2. PARTIAL MIGRATION Introduction The study of animal movements has lacked continuous observations of individuals over s i g n i f i c a n t fractions of their l i f e t i m e s (Wiens 1976, Baker 1978, Swingland 1983). Advances in technology, including radio-telemetry, a e r i a l tracking, and satellite-based tracking, now make i t possible to Obtain detailed individual movement data for medium to large vertebrates. This allows us to re-examine animal movement, giving greater attention to i n d i v i d u a l i t y . Individual animals are often either migratory or resident. Migrations in vertebrates can be defined as "the r e p e t i t i v e seasonal movements of individuals between d i s t i n c t areas" ( S i n c l a i r 1983). Residents use the same area throughout the seasons, in " p a r t i a l " migrant species, not a l l individuals migrate (Baker 1978). P a r t i a l migration i s widespread amongst vertebrates (marine mammals, Wursig and Wursig 1980; cervids and bovids, Swingland 1983; wild horses, Berger 1983; bears, McLellan 1989; bats, Swingland 1983; passerine birds, Adriaensen and Dhondt 1990; f i s h and t u r t l e s , S i n c l a i r 1983). How can 2 or more strategies coexist within 1 species or population? Lack (1943) suggested that p a r t i a l migration in the European robin (Erithacus  rubecula) could be explained by the presence of 2 11 genetically d i s t i n c t raorphs whose average long-term pay-offs balance ( i . e . a mixed ESS). Although migratory restlessness i s a heritable t r a i t in birds (Berthold and Querner 1981), the genetic polymorphism hypothesis i s weakened by the fact that individuals may change their migratory status between years (Baker 1978), and by the observation of facultative migration in species such as the yellow-rumped warbler (Dendroica coronata) ( T e r r i l l and Ohmart 1984 ) . Recently, Adriaensen and Dhondt (1990) i l l u s t r a t e d unequal pay-offs between residents and migrants (residents did better) in the European robin in Belgium. They presented a "conditional" explanation for the 2 behaviors. They suggest that individuals with strong genetic tendencies to migrate or to remain resident, w i l l always do so. Between these extremes, individuals make a "conditional" assessment of their environment and choose to migrate/stay accordingly. This suggests that the choice might be density-dependent or that i t might change with variation in the quality of the environment. Unequal pay-offs are suggested by work on African ungulates (Fryxell and S i n c l a i r 1988, F r y x e l l et a l . 1988). F r y x e l l et al.(1988) observed that migrants were more abundant than residents and proposed a model that suggested that migrants might escape population regulation by sedentary predators by moving away from them for part of the year. At the individual l e v e l , 12 movements that reduce encounters with predators could increase f i t n e s s . Boyce (1991) used a model to i l l u s t r a t e that seasonal fluctuations in food a v a i l a b i l i t y on summer (high elevation) elk ranges could determine the fitn e s s of migratory and non-migratory elk. In areas with low seasonality, residents had the highest fitness because of the cost of migration. At moderate levels of seasonality, fitness of both morphs decreased, although fit n e s s of residents decreased more steeply. The model did not address the problem of coexisting migrant and non-migrant elk within a single population. Female-biased ratios of migrants to residents have been found i n several species of birds (Adriaensen and Dhondt 1990). Although female-biased migration has been shown in 1-year-old elk (Boyce 1989), p a r t i a l migration biases in mammals have received l i t t l e attention. Greenwood (1980) reviewed sex-biased dispersal (non-return movement) and i d e n t i f i e d an apparent dichotomy: female-biased dispersal in the majority of birds and male-biased dispersal in the majority of mammals. He suggested that this i s related to mating strategy. In many birds, males defend resources ( t e r r i t o r i e s ) and are monogamous. In many mammals, males defend females and are polygynous. Dispersal in mammals i s thus most l i k e l y in the sex which defends mates rather than resources. 13 Elk have the widest natural d i s t r i b u t i o n of any wild ungulate in the world (Clutton-Brock et a l . 1982) and have a range of movement behaviors including migration, residency, and p a r t i a l migration (Adams 1982). In mountainous areas, elk often make a l t i t u d i n a l migrations featuring considerable climatic and vegetation change over r e l a t i v e l y short horizontal distances. They are seasonal breeders with a polygynous mating system. In t h i s study, I radio-tracked individual elk from a p a r t i a l l y migrant population and described their seasonal movement patterns. I addressed the following questions: Is the observed pattern of migration consistent with a mixed ESS explanation?; Is p a r t i a l migration in elk sex-biased? Are migrants more abundant than residents?; and, Do migrant elk escape the foraging range of their predators?. P a r t i a l migration as a mixed ESS predicts that (1) resident and migratory individuals remain f a i t h f u l to their movement strategies and that the pay-offs to migrants and residents must be equal. A conditional ESS predicts that (2) the movement strategy depends on status (e.g. sex, age) or an individual's assessment of environmental s u i t a b i l i t y . Fryxell's hypothesis of predator avoidance ( i . e . migration takes elk out of the range of sedentary predators) predicts that (3) migrants should outnumber residents. 14 Methods Radio-tagging From January 1986 to A p r i l 1989, 53 adult elk were radio-tracked successfully within the BRV for more than 6 months. This sample consisted of 18 adult b u l l s (older than 2-year-olds) and 35 adult cows (older than calves) The cow sample included 1 radio-tagged mother-daughter pair. The daughter was radio-tagged as a c a l f and followed u n t i l she was a 3-year-old. Radio-transmitters were distributed across the winter (October-May) d i s t r i b u t i o n of elk as determined by previous a e r i a l surveys conducted by park wardens (Woods 1990). Park wardens chemically immobilized elk by darting them from patrol trucks or after approach on foot. One animal was darted from a helicopter. The f i r s t elk o f f e r i n g a safe darting opportunity was selected. No more than 2 elk were captured from any group (animals within v i s u a l contact of each other) at a time. Two cows radio-tagged in previous studies within the BRV were re-tagged. Most animals were captured during the winter but 2 adult b u l l s and an adult cow were tagged in August. Most transmitters placed on adult elk were attached to c o l l a r s . These radios had a battery l i f e of about 4 years and had mortality sensors that were triggered by 3 hours of i n a c t i v i t y . A numbered tag on the c o l l a r allowed individual v i s u a l recognition. A l l c a l f and most 1- year-old elk were f i t t e d with solar powered ear-tag transmitters. Two of these ear-tag transmitters were la t e r replaced with r a d i o - c o l l a r s . From January 1986 u n t i l February 1989, the presence of radio-tagged elk within the BRV was determined by ground searches during daylight hours at intervals of 7 days or le s s . Observers drove the entire length of the highway and made frequent stops (usually <3 km) to scan radio frequencies using a portable radio receiver and hand-held antenna. During a ty p i c a l week, 100% of the radio-transmitters within the BRV were located during 2- 3 days of ground searches. When a radio-transmitter was heard, the observer drove as close as possible to the animal's location and took 3 or more bearings from points approximately 0.5 km apart. These bearings were then plotted on study area maps (1:50,000) in the f i e l d . If the resulting polygon was greater than 0.2 km in any dimension, additional bearings were made to reduce the size of the polygon. The centre of the f i n a l polygon was converted to a grid reference with a resolution of 0.1 x 0.1 km. The elevation of this point was estimated to the nearest 30 m contour in t e r v a l from maps of the study area. If a radio-tagged elk was seen during a survey, i t s i d e n t i t y was determined from the number on the c o l l a r and confirmed by checking i t s radio frequency. In addition to ground triangulation, major 16 t r i b u t a r i e s of the Bow River and passes connecting the BRV to neighboring valleys were searched from the a i r during daylight hours at monthly intervals in 1986, and biweekly thereafter, to locate animals absent from the BRV. A mobile receiver in a helicopter with 2 externally mounted antennae allowed a l l functioning radios to be located throughout the study. Repeated low-elevation passes with the helicopter allowed point estimates of the elk's position. Elevations were estimated from study area maps and cross-checked with the helicopter's altimeter. The animal's position was converted to a grid reference. Radio signal "bounce" was encountered frequently but these spurious signals became obvious when multiple bearings were plotted in the f i e l d . Additional bearings were taken u n t i l the bounce problem was eliminated. F i e l d t r i a l s suggested grid reference positions were t y p i c a l l y within 0.2 km of the animal's true position. The actual error of each mapped ground observation was not measured but believed to be a complex function of position i n the valley and observer experience. Concurrent radio-tracking of 2 timber wolf packs by the park warden service and by D. Huggard (pers. comm.), allowed general comparisons of wolf and elk movements. 17 Data analysis Individual locations were transcribed into a computer f i l e and plotted sequentially by date and elevation, and by date and horizontal position. On an annual basis, a l l radio-tagged animals within the BRV were c l a s s i f i e d as either migrants or residents. Sequential location plots were examined and animals c l a s s i f i e d as migrants i f they made at least 1 return movement l a s t i n g more than 2 weeks between 2 separate areas. This d e f i n i t i o n of migration implies philopatry to seasonal ranges and r e l a t i v e l y l i t t l e use of areas between these ranges. The remaining elk were classed as residents. Horizontal movements were estimated for each individual by calculating the difference between extreme observation points for the entire period an individual was tracked. These were then adjusted to represent the shortest possible movement route for an elk when topographical features (e.g. c l i f f s ) barred s t r a i g h t - l i n e t r a v e l . V e r t i c a l movements were estimated as the maximum elevation minus the minimum elevation point. Departure and return dates of migrating animals were estimated as the l a s t day of radio contact within the BRV and the f i r s t day of contact when that animal returned. Seasonal ranges were i d e n t i f i e d by p a r t i t i o n i n g the data into 3 periods: winter, October 1 to May 30; 18 summer, June 1 to August 31; and, rut (breeding), September 1 to September 30. In winter, most elk were within the BRV, and in summer most migratory animals were out of the valley (Woods 1990). The rut season embraces the height of breeding a c t i v i t y for this population (Struhsaker 1967). Average elevations were calculated from a l l available observation points for an individual for each season. The migrant/resident (M/R) ratio for each sex was based on the number of radio-tagged elk that were la t e r defined as either migrants or residents. Individuals that switched movement status between years, or that were captured in the summer, were excluded from these calculations. An estimate of the overall composition of the population was calculated by summing the number of elk observed during 6 autumn and 6 spring c l a s s i f i e d counts (Chapter 6) into 2 categories: adult b u l l s and adult cows (calves and 1-year-olds are believed to migrate with adult cows). The M/R ratio for each sex was then used to estimate the number of migrants and residents in the overall population. The fate of a l l radio-tagged adult elk was known. Therefore, dispersal could be separated from mortality. Survival rates and 95% confidence intervals were calculated for radio-tagged elk using the Kaplan-Meier procedure as modified for staggered entry and censored (Pollock et a l . 1989). 19 Results Seasonal movement patterns Elk were located on 7640 occasions for periods of 6-36 months. Sequential plots of locations demonstrated that both sexes had resident and migrant individuals (Table 1, Figure 2) . One adult b u l l made a return migration in 1986 and then dispersed about 150 km west. Migrants moved over a s i g n i f i c a n t l y greater horizontal and v e r t i c a l range than residents (Table 2) . Three elk had narrow home-ranges along the valley floor with dimensions comparable to the travel distances of v e r t i c a l migrants. Two of these (cows) did not make discernible return movements between segments of the vall e y and were c l a s s i f i e d as residents. They produced the extreme ranges reported for residents in Table 2. The t h i r d , a b u l l , made repetitive movements between d i s t i n c t areas in the valley and was classed as a migrant. Twenty-five elk did not migrate (Figure 2a,c) and 25 did (Figure 2b,d,h). Fifteen migrants made 1 migration cycle per year and 10 made 2-3 cycles within a year (Figure 2h). These additional cycles included returns to low elevation during the summer, returns to high elevation during the winter, and repetitive o s c i l l a t i o n s between low and high elevations throughout the year. Two bul l s spent most of the summer at high elevations, returned to low elevations in advance of the 20 rut, moved back to high elevations after the rut, and then descended again in the early winter (Figure 2h). The elevational d i s t r i b u t i o n of migrant and resident elk d i f f e r e d markedly between seasons (Figure 3). In winter, both migrants and residents were i n the montane zone of the BRV. In summer, most migrants moved to the subalpine and their mean elevation was s i g n i f i c a n t l y higher than that of residents (Wilcoxon pairs test, P<0.05). In the rut, many migrant b u l l s returned to the BRV and were sympatric with residents. However, the mean elevation of migrant cows during the rut was s i g n i f i c a n t l y higher than that of resident cows. Most migratory individuals i n i t i a t e d their migrations in early June and returned in September (Table 3) . Although median departure dates were similar for both sexes, bulls returned to the BRV s l i g h t l y e a r l i e r than cows. Fewer than 25% of the radio-tracked elk migrated during any weekly period. During any year, most individuals migrated within 17 days of their migration date the previous year. The presence of timber wolves was suspected of delaying the migration of 1 adult cow elk. In June 1988 t h i s elk was radio-tracked 12 km from the BRV part way along the migration route she had used during 1986 and 1987. At this point she encountered a wolf pack containing a radio-tagged individual (D. Huggard, pers. comm.). The next day the elk returned to the BRV. 21 F i f t y - f o u r days l a t e r she r e t r a c e d the route and continued 70 km to her summer range. Seven mature b u l l s (2 r e s i d e n t s , 5 migrants) moved to s p e c i f i c areas of t h e i r annual home-ranges only used d u r i n g the r u t . Both of the r e s i d e n t b u l l s moved from w i t h i n the Banff townsite to p e r i p h e r a l areas. For 1 of the 5 migrant b u l l s , a r u t t i n g season e x c u r s i o n to low e l e v a t i o n was the only r e t u r n movement observed. Most migrant cows moved to low e l e v a t i o n s before or d u r i n g the r u t . They d i d not have s p e c i f i c areas used onl y d u r i n g the r u t t i n g season. A d u l t cow elk are s e c r e t i v e at c a l v i n g time, and i t was thus d i f f i c u l t to determine whether or not a cow had a young c a l f . There was only 1 case of an apparent movement to a s p e c i f i c c a l v i n g area. T h i s r e s i d e n t returned to small fenced b i s o n (Bison bison) enclosure to bear her young duri n g 3 consecutive y e a r s . The remaining r e s i d e n t s apparently c a l v e d i n a v a r i e t y of l o c a t i o n s w i t h i n t h e i r annual home-ranges. The median departure dates f o r migrant elk c o i n c i d e d with the c a l v i n g season ( l a t e May-early June, Table 3). I n c i d e n t a l o b s e r v a t i o n s of r a d i o - t r a c k e d migratory cows i n d i c a t e d that c a l v i n g could take p l a c e on e i t h e r winter or summer ranges, or i n the case of 1 radio-tagged cow, between ranges. A f t e r 3 weeks i n t h i s l o c a t i o n , she completed her m i g r a t i o n . The m a j o r i t y (94%) of elk showed the same movement 22 s t r a t e g i e s from ye a r - t o - y e a r . A cow r a d i o - t r a c k e d i n the BRV i n 1981 and determined to be r e s i d e n t ( P . Jacobson, pe r s . comm.), was a s t i l l a r e s i d e n t d u r i n g t h i s study (1986-89). However, another p r e v i o u s l y s t u d i e d a d u l t cow made a b r i e f d u r a t i o n v e r t i c a l movement i n 1981 ( P . Jacobson, pers. comm.), but was r e s i d e n t throughout 1987-89. During the study, 3 cows switched s t r a t e g i e s between y e a r s . One a l t e r n a t e d between m i g r a t i o n , r e s i d e n c y , and m i g r a t i o n . (Figure 2e). A 1-year-old e l k , r e s i d e n t i n 1986, migrated up an adjacent slope as a 2-year-old (Figure 2 f ) . An a d u l t radio-tagged i n August 1986, remained i n the BRV throughout 1986, but made mig r a t i o n s d u r i n g 1987 and 1988 (Figure 2g). None of these e l k had to cross the fenced highway c o r r i d o r to migrate. In 3 cases, c a l v e s of radio-tagged cows were captured and followed f o r up to 29 months. The female c a l f of 1 migrant accompanied her mother f o r 2 y e a r s . In her 3rd year, she separated from her mother but migrated to the same areas she had t r a v e l l e d to as a c a l f and as a 1-year-old. One female and 1 male c a l f of r e s i d e n t e l k stayed with t h e i r mothers f o r at l e a s t 9 months a f t e r b i r t h . Although both where observed the f o l l o w i n g winter w i t h i n t h e i r mother's annual home-ranges, both of t h e i r t r a n s m i t t e r s had f a i l e d . 23 M i g r a n t - r e s i d e n t r a t i o a n d s e x - b i a s e s The c o r r e c t e d m i g r a n t / r e s i d e n t r a t i o f o r t h e e n t i r e BRV p o p u l a t i o n was 0 . 6 ( T a b l e 1 ) . When t h e s e d a t a w e r e s u b - d i v i d e d i n t o t h e a r e a s w e s t a n d e a s t o f B a n f f t o w n s i t e ( W e s t - B R V , E a s t - B R V ) , t h e y d e m o n s t r a t e d a m i g r a n t / r e s i d e n t r a t i o <1 i n t h e E a s t - B R V a n d >1 i n t h e W e s t - B R V . I n t h e BRV as a w h o l e a n d i n b o t h s u b - d i v i s i o n s , more b u l l s m i g r a t e d t h a n c o w s . T h i s d i f f e r e n c e was c o n s i s t e n t , a n d s t a t i s t i c a l l y s i g n i f i c a n t i n t h e e n t i r e a r e a a n d i n E a s t - B R V ( T a b l e 1 ) . I n W e s t - B R V , t h e s a m p l e s i z e ( 1 5 ) was t o o s m a l l t o r e j e c t t h e n u l l h y p o t h e s i s o f e q u a l n u m b e r s o f m i g r a n t s a n d r e s i d e n t s , b u t 1 2 o f 1 5 a n i m a l s m i g r a t e d . S u r v i v a l T h i r t e e n r a d i o - t a g g e d a d u l t e l k o f known m i g r a t o r y - s t a t u s e l k d i e d d u r i n g t h e s t u d y . A l l m o r t a l i t i e s o c c u r r e d a t l o w e l e v a t i o n s w i t h i n o r a d j a c e n t t o t h e B R V . T h e s e d e a t h s i n c l u d e d e q u a l n u m b e r s o f r e s i d e n t s a n d m i g r a n t s . H u m a n - r e l a t e d e l k m o r t a l i t i e s o u t n u m b e r e d n a t u r a l m o r t a l i t i e s 7 : 6 . H u m a n - c a u s e d m o r t a l i t i e s i n c l u d e d r o a d - k i l l s ( 4 ) , r a i l w a y - k i l l s ( 1 ) , h u n t i n g ( 1 ) , a n d m i s c e l l a n e o u s ( 1 ) . N a t u r a l d e a t h s i n c l u d e d p a r a s i t e s ( 2 ) , unknown ( 2 ) , a n d w o l f p r e d a t i o n ( 2 ) . A l t h o u g h s u r v i v a l r a t e s o f r a d i o - t a g g e d a d u l t e l k w e r e h i g h e s t f o r m i g r a n t s ( 0 . 9 7 cow, 0 . 8 6 b u l l ) , 24 confidence intervals on these estimates were too broad to reject the n u l l hypothesis (Table 4). Concurrent radio-tracking studies of timber wolves showed that wolves rarely used East-BRV (D. Huggard, pers. comm.). One pack used a den in the subalpine zone within the West-BRV and the other used a den approximately 20 km south of the BRV beside the Spray River and along an elk migration route. Movements by individual wolves equaled or exceeded elk migration distances. Migration could take individual elk into the range of other wolf packs. The farthest elk migration (74 km) was across the Continental Divide and into an area where wolves were seen or heard throughout this study. Although there were small areas of high elevation range where wolves were not seen, in general, elk and wolves were sympatric at a l l seasons. Other potential predators preying on elk, and observed during the study included: g r i z z l y bears (Ursus arctos), black bears (Ursus americanus), and cougar ( F e l i s concolor). Of these, g r i z z l i e s were the most commonly seen and were sympatric with resident and migrant elk during the spring, summer, and autumn. 25 Table 1. Migrant/resident r a t i o 8 for radio-tracked adult elk in Banff National Park, 1986 -88. Sex Area BRV BRV-Westb BRV-Eastc Bulls .4.3 7.0 3.0 (13/3) d (7/1) (6/2) Cows 0.5s 2.5 0.3f (10/21) (5/2) (5/19) Population* 0.6 2.7 0.4 a excludes 3 cow elk that switched movement strategy between years and 3 elk captured in the summer b radio-tagged west of Banff townsite in area frequented by timber wolves c radio-tagged east of Banff townsite in area rarely used by timber wolves d numbers in brackets indicate observed number of individuals • s i g n i f i c a n t l y smaller than the corresponding b u l l migrant/resident proportion, 2x2 Contingency table with Yates' correction, P=0.003; a l l s t a t i s t i c a l tests were made on the STATGRAPHICS software ( S t a t i s t i c a l Graphics Corporation) run on a microcomputer. f s i g n i f i c a n t l y smaller than the corresponding b u l l migrant/resident proportion, 2x2 Contingency table with Yates' correction, P=0.018 9 weighted by the estimated population composition (11% adult b u l l s , 89% adult cows, calves, and spike bulls) 26 Table 2 . Horizontal and v e r t i c a l movements8 of migrant and resident radio-tracked adult e l k b in Banff National Park, 1 9 8 6 - 8 9 . Category Migrants Residents Sex Average Range Average Range (N) c (N) Horizontal (km) Bulls 3 0 d ( 1 4 ) 1 8 - 7 0 11 ( 4) 5 -20 Cows 3 6 d ( 1 0 ) 18 -74 14 (22) 7 -36 V e r t i c a l (m) Bulls 8 4 0 d ( 1 4 ) 396 -1147 419 ( 4) 1 2 2 - 9 4 5 Cows 1 0 7 9 d ( 1 0 ) 7 0 1 - 1 4 3 2 296 (22) 1 2 2 - 8 8 4 a calculated as the difference of the extreme observed locations during entire study period b excludes three cows that switched movement classes between years c numbers in brackets indicate observed number of individuals d value for migrants s i g n i f i c a n t l y d i f f e r e n t from corresponding values for residents (Wilcoxon pairs test, P<0.05). 27 T a b l e 3. M e d i a n m i g r a t i o n d a t e s f o r r a d i o - t r a c k e d a d u l t e l k i n B a n f f N a t i o n a l P a r k , 1986-88. D e p a r t / R e t u r n Y e a r Sex 1986 (N) a 1987 (N) 1988 (N) D e p a r t B u l l s J u n e 2 (8) J u n e 7 (7) J u n e 8 (8) Cows J u n e 6 (9) May 26 (12) J u n e 17 (13) R e t u r n B u l l s S e p t 5 (6) S e p t 9 (7) S e p t 13 (8) Cows O c t 3 (9) O c t 2 (12) S e p t 19 (13) a n u m b e r s i n b r a c k e t s i n d i c a t e o b s e r v e d number o f i n d i v i d u a l s 28 Table 4. Annual survival rates (S) and 95% confidence intervals (CI) for migrant 8 and resident radio-tagged adult elk in the BRV, 1986-89. Class Year N S CI b Migrant cows 1.00-1.00 1986-87 9 1.00 1987-88 10 1.00 1.00-1.00 1988-89 10 0.90 0.72-1.08 Average 10 0.97 0.91-1.03 Migrant b u l l 0.44-0.94 1986-87 9 0.69 1987-88 11 0.89 0.71-1.07 1988-89 11 1.00 1.00-1.00 Average 10 0.86 0.72-1.00 Resident cow 1986-87 20 0.89 0.75-1.02 1987-88 20 0.94 0.84-1.05 1988-89 20 0.90 0.76-1.03 Average 20 0.91 0.78-1.03 Resident b u l l 1986-87 3 1.00 1.00-1.00 1987-88 4 0.75 0.38-1.12 1988-89 3 0.67 0.13-1.20 Average 3 0.81 0.50-1.08 a excludes 3 cows that switched strategies between years b calculated following Pollock et a l . (1989) 29 Table 5. Migrant/resident ratios (M/R), hunting status 8, and timber wolf b status for various elk populations in the Rocky Mountains Area M/R Hunting Wolves Reference BRV, ABC <1 N Y this study Red Deer, AB >1 Y Y Morgantini (1988) East Kootenays, BCC >1 Y Y Woods and Hlady (1988) Sun River, Mont >1 Y N Knight (1970) Jackson Hole,WY >1 Y N Boyce (1989) a c l a s s i f i e d as hunted i f most of the population in a sport hunting zone during the hunting season b timber wolf status at time of study, a l l populations had mountain lions present c AB=Alberta; BC=British Columbia 30 2300 ^ 1400 0-RESIDENT FEMALE 86 87 88 ©-RESIDENT MALE —I 1 1 86 87 88 ©-SWITCHER FEMALE H h 86 87 88 ©-SWITCHER FEMALE Alpine 86 87 88 ©-SWITCHER FEMALE 86 87 88 ©-2 CYCLE MIGRANT MALE (WO* 86 87 88 86 87 88 Figure 2. Examples of elevational movements of 8 radio-tagged elk, BRV, 1986-88. 31 2000 UJ CO 1600 LU < 21 11 0 t—i GO UJ o r , CD 1300 M 14 I 11 M 'i i - A-i 1 1 11 / -21/ 1 M 13 0 / / / i WINTER SUMMER SEASON RUT Figure 3. Mean elevation of resident and migrant radio-tagged elk during winter, summer, and rut, BRV, 1986-88. Letter above bars indicate sex (F/M), number indicates sample size, and asterisk indicates s i g n i f i c a n t difference between migrants and residents. 32 Discussion Seasonal movement patterns Although elk at Jackson Hole, Wyoming moved in large groups (Boyce 1991), elk in a Montana population migrated singly or in matriarchical groups (Knight 1970). The staggered dates of migrating BRV elk suggest that elk in this population migrated singly or i n small groups. Further, although most BRV migrants wintered at low elevation and summered at high elevations, individual behaviors such as multiple migration cycles per year, and s p e c i f i c migrations to rutting areas, i l l u s t r a t e d that migration patterns varied widely between individuals. Because of the generally forested nature of the BRV and surrounding area, t h i s behavioral v a r i a t i o n could not have been i d e n t i f i e d without radio-tracking. The f a i l u r e to detect "calving grounds" (places where cow elk return to bear their young) agrees with research by Knight (1970) in the American Rockies and by Morgantini (1988) in an adjacent area of the Canadian Rockies. However, calving areas were i d e n t i f i e d by Boyce (1989) for the Jackson elk herd. A clear d e f i n i t i o n of what constitutes a calving area i s lacking in these studies and i t i s possible that the discrepancy i s semantic. "Rutting grounds" (places where elk return to breed each autumn), have been reported for resident b u l l elk 33 in C a l i f o r n i a (Franklin et a l . 1975). In the BRV, cows were concentrated in the valleys during the rut (resident cows plus some of the returning migrants) but rarely entered the townsite. Movement by bul l s to rutting grounds within the valley, but outside the townsite, probably gave these b u l l s access to more cows than would have been available on their summer ranges. Migrant b u l l s generally returned to the val l e y at the onset of rut (early September), before the peak in mating opportunities (mid to late September). This movement placed them in the sit e s with the most mating opportunities at the onset of the sexual r e c e p t i v i t y of cows. Since migrant cows generally returned at the end of the rut, I presume that most of them mated with migrant b u l l s that remained on summer ranges or that they mated late in the rut. Mixed ESS Although p a r t i a l migration has been observed i n many elk populations (Scotland, Darling 1937; Scandinavia, Langvatn and Albon 1986; coastal and mountainous areas of the United States, Adams 1982, McCullough 1985; Canadian Rockies, Morgantini 1988), there i s l i t t l e information on f i d e l i t y to movement strategy. The existence of "switchers", such as demonstrated in thi s study, requires r e l a t i v e l y long-term, continuous tracking and therefore may be more common than i s reported in the l i t e r a t u r e . 34 On the basis of high ratios of 1-year-old b u l l s spending the summer on the "winter" range of the Jackson Hole elk, Martinka (1969) and Boyce (1989) suggested that 1-year-old b u l l elk are less l i k e l y to migrate than are other sex/age classes. The Jackson Hole herd has a migrant/resident ratio >1. Therefore, most of these young bu l l s presumably migrated during their f i r s t summer with their mothers but f a i l e d to do so during the following summer (=switching). There are no data to suggest a disproportionate number of 1-year-old b u l l s at low elevation in the BRV during the summer. Since young elk accompany their mothers at least through their f i r s t summer and autumn, the female-bias in resident BRV elk should result in more calves being exposed to the residents than to migratory individuals. However, by the time they are adults, there are fewer b u l l s in the population (Flook 1970, Chapter 6), and most of these are migrants. Either more bul l s "switch" to migration as a strategy, or unbalanced mortality (or dispersal) results in the change of r a t i o . Between the ages of 1 and 2, the proportion of bulls to adult cows decreases sharply in this population (Flook 1970, Chapter 6) . Flook (1970) speculated that this might be due to greater dispersal by 1- or 2-year-old males. If this i s the case, then bulls in these age classes with resident mothers should be the most frequent dispersers. This could be tested by radio-tracking b u l l calves of 35 resident mothers from b i r t h through to their 3rd birthday. Since most bulls and many cows return to the v a l l e y during September, many migrants are sympatric with residents during the rut. This makes the existence of 2 genetically d i s t i n c t sub-populations u n l i k e l y . Learning An alternative explanation for choice of movement strategies based on learning, has been presented by Murie (1951), McCullough (1985), and Baker (1978), who suggest that young animals develop patterns of home-range use through association with others (especially their mothers) and by individual experience. Strategies proven to work (the c a l f survived to breeding age) should be preferred over unknown strategies in the absence of intervening variables (climatic extremes, the appearance of predators, shortage of mates). In the single instance where a mother-young pair was continuously tracked in the present study, the daughter adopted the movement strategy of her mother. Although Clutton-Brock et a l . (1982) demonstrated a close association between mother and daughter red deer (also Cervus elaphus) during the f i r s t years of the daughter's l i f e , t his t r a i t has not been extensively documented in elk or other ungulates. More empirical data are needed of tracked mother/young pairs from p a r t i a l migration populations to investigate i f most 36 young adopt the movement pattern of their mothers. Morgantini (1988) stressed that f l e x i b i l i t y in movement patterns allows elk to adjust to a variable environment and Boyce (1991) noted that such phenotypic p l a s t i c i t y can be shaped by natural selection. Bergerud (1974) developed much the same argument to explain v a r i a t i o n i n caribou (Rangifer tarandus) movements, and suggested that adaptability was a major adaptation in that species. The patterns and variations observed in this study support these views. F l e x i b i l i t y would allow elk to make a conditional assessment of density and environmental variables, and to either migrate or stay accordingly. Migrant/resident ratio The excess of migrants over residents in African ungulates (Fryxell et a l . 1988) contrasts to my results for the BRV as a whole (M/R= 0 .6) . However, as Fry x e l l et a l . (1988) noted, the short distances involved in a l t i t u d i n a l migration may not take migrants beyond the foraging range of their p r i n c i p a l predators. Although migrants outnumbered residents (M/R= 2.7) within the West-BRV, this difference 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 . In addition to being the primary area used by wolves in the BRV, West-BRV d i f f e r s from East-BRV in elevation, snowfall, vegetation zones, and human densities. A larger sample of radio-tagged elk studied over a range wolf densities i s required to 37 establish a relationship between migrant/resident ratios and predation in this population. Predators can migrate too. This appears to be the case in the mountain lion-elk population studied by Seidensticker et a l . (1973). These lions made a l t i t u d i n a l movements in concert with migrating elk and deer (Odocoileus). Harestad (1979) suggested that risk of predation might be less for migrating b l a c k - t a i l e d deer (Odocoileus hemionus columbianus) during the period when wolves are feeding young at den s i t e s . However, the deer he studied migrated on average less than 5 km from their wintering areas. In my study area, wolves moved over much larger distances and wolf mobility equalled or exceeded the elk migratory range during a l l seasons. The hypothesis that wolves are less l i k e l y to capture elk on summer ranges remains to be tested. The BRV elk population i s unusual today in that wolves are p r i n c i p a l predators, few BRV elk are subject to sport hunting, and migrant/resident ratio i s <1 (Table 5 ) . These differences are l i k e l y to confound comparisons with other populations. As Lima and D i l l (1990) observed in their review of behavioral decisions made under the risk of predation, the f a i l u r e to avoid predators has both immediate and permanent consequences on individual f i t n e s s . Hornocker (1970) noted that elk and deer reacted to k i l l s by mountain lions by moving away from the k i l l s i t e s , 38 sometime to adjacent drainages. Numerous authors have demonstrated that sport hunting can disturb elk movement patterns (Altmann 1956, Morgantini and Hudson 1985, Edge et a l . 1985, M e r r i l l et a l . 1988, Boyce 1989). It seems l i k e l y that, where elk are hunted, they avoid human contact as much as possible. This could result in a di f f e r e n t ratio of residents to migrants and discourage return to low elevations. The potential for human influence on the migrant/resident ratio i s i l l u s t r a t e d by the seasonal movements of elk in the Red Deer River drainage immediately north of the BRV (Morgantini 1988). "Most" animals in this population were migratory and had a two-stage migration from winter range outside the park, to an intermediate range inside the park, and then to summer range farther within the park. They reversed this pattern in the autumn. Although timber wolves were p r i n c i p a l predators of elk in both the BRV and the Red Deer River valley, Red Deer elk were hunted and were not habituated to humans. Although they l e f t the winter range at about the same time as BRV elk, they remained within the protection of the park for about a month longer in the autumn. During the 1940's, disturbance by humans and timber wolves in the BRV may have influenced the timing of the autumn elk migration and the migrant/resident r a t i o . In 1943-44, an intensive e f f o r t to reduce the elk population began and prescribed numbers were shot during 39 the late autumn or early winter (Chapter 6) . Coincidentally, wolves reappeared in Banff and were r e l a t i v e l y common by 1947 (Flook 1970). At the star t of this period, migration was believed to be r e s t r i c t e d to adult b u l l s and migrants were thought to return in September (K. Mi t c h e l l , CPS, 1944, unpubl. data). By autumn 1948, the park was unable to provide an accurate estimate of elk numbers because most of them had not returned to the valley (K. Mi t c h e l l , CPS, 1948, unpubl. data). Green (1949) speculated that this was due to either shooting disturbance or the presence of timber wolves. Like movements of mountain l i o n (Seidensticker et a l . 1973), use of space by elk i s f l e x i b l e . Simple categorization of populations as either migrant or resident may mask the relationships driving the system. As Morgantini (1988) and Boyce (1991) suggested, phenotypic p l a s t i c i t y ( f l e x i b i l i t y ) in elk appears to be an adaptation to cope with a variable environment. 40 CHAPTER 3. PHILOPATRY Introduction Movements of individuals within and among populations are of central importance to genetics, ecology, and conservation. However, analysis of these movements i s complicated by problems of consistency in de f i n i t i o n s and inequ a l i t i e s of scale (Chepko-Sade and Halpin 1987). For example, migration i n the context of genetics refers to the dispersal of genes between populations (Chepko-Sade and Shields 1987). In vertebrate ecology, migration describes the seasonal return movements of individuals between alternate ranges ( S i n c l a i r 1983). Therefore, even though migrant individuals might undertake extensive movements, i f they return to the same area to breed, they remain part of a re s t r i c t e d population in a genetic sense. The tenacity with which an organism returns to s p e c i f i c areas or groups, i s generally referred to as philopatry (Greenwood 1980). In contrast, dispersers either move away from their place of b i r t h (natal dispersers) or move between breeding s i t e s (breeding dispersers) (Shields 1987). In practice, the d i s t i n c t i o n between philopatry and dispersal i s less clear. A bird which might be philopat r i c on a regional scale, might not be phi l o p a t r i c at some other scale (e.g. patch). In response to this problem, d e f i n i t i o n s of "dispersers" have become numerous in f i e l d studies. 41 Dispersers have been defined as those animals which l e f t the study area and never returned (Mech 1987, Boonstra et a l . 1987), or as those that moved some fixed distance away from where they were born or where they previously bred. These distances have been established on the basis of the maximum movements of migrant individuals (Bunnell and Harestad 1983), numbers of t e r r i t o r i e s or home-ranges traversed (Shields 1987, Arcese 1989) or simply whether or not consecutive home-ranges overlapped (Edge et a l . 1985). Detailed studies of several large t e r r e s t r i a l mammals have suggested high degrees of adult philopatry (Knight 1970, Adams 1982, Nelson and Mech 1987, Morgantini 1988) and have related inbreeding to conservation (Weishampel 1990). However, theoretical c r i t e r i a for defining philopatry (or dispersal) are lacking. Many studies have noted that elk habitually return to seasonal ranges (Knight 1970, Adams 1982, Edge et a l . 1985, Morgantini 1988, Boyce 1989). Elk are polygynous, seasonal breeders, with female-biased adult sex ratios (Flook 1970). Populations may be resident, migratory, or p a r t i a l l y migratory. According to Greenwood's hypothesis, polygynous species such as elk should have male-biased dispersal (Greenwood 1980), and females should show the greatest s i t e attachment. While there i s evidence that 1-year-old bulls (spikes) are the 42 most frequent dispersers in elk (Boyce 1989), good evidence of male-biased dispersal i s lacking. There also are few comparative data on philopatry of resident versus migrant animals within a population. Since residents move over a smaller t o t a l range than migrants, they might be more ph i l o p a t r i c . In this study, I measured the consistency with which adult elk in a p a r t i a l l y migratory population returned to seasonal ranges in the BRV. Strong philopatry predicts that individual elk use the same areas r e p e t i t i v e l y , such that the distances separating seasonal ranges during consecutive years are less than predicted by a random model. I also compared the degree of philopatry to test whether cows and residents were less p h i l o p a t r i c than b u l l s and migrants. 43 Methods I examined philopatry using the radio-tagged sample of 53 elk, f i e l d methods, and de f i n i t i o n s described in Chapter 2. Consistency in use of annual home-ranges was measured by comparing successive annual home-ranges of individual elk. Annual home-ranges were estimated using McPAAL software (Stuwe and Blohowiak 1985) to generate minimum convex polygons. These polygons are formed by connecting the outermost locations without creating any in t e r i o r angle greater than 180 degrees. Annual polygons were generated from February to February, for 3 years (1986-89). Successive annual home-ranges for an individual were superimposed on a d i g i t a l plotter and the area of overlap was calculated as a percent of the to t a l area. Consistency in use of seasonal ranges was measured by p a r t i t i o n i n g the data into 7 periods sta r t i n g with winter 1986 and ending with winter 1989. Seasons were defined as winter, summer, and rut as described in Chapter 2 (p.18). If only 1 point was available per season (as was the case for some migrants in the rutting season), that point became the centre of a c t i v i t y . Otherwise, for each radio-tagged elk, I determined the harmonic centre of i t s observed locations on a 1000 m grid for each season (Dixon and Chapman 1980). I then calculated the s t r a i g h t - l i n e distances between these centres i n that season in consecutive years (e.g. the distance between individual's rutting season a c t i v i t y centres in 1986 and 1987) . The frequency d i s t r i b u t i o n s of observed seasonal centres of a c t i v i t y were then compared to a frequency d i s t r i b u t i o n derived from a model simulating random location of successive a c t i v i t y centres. The random model was generated by calculating the point-to-point distances for 1000 random pairs in a hypothetical square of 196 km2. This area approximates the montane zone of the BRV which i s available to elk on a year-round basis. The model generated a unimodal d i s t r i b u t i o n of distances between seasonal ranges in di f f e r e n t years with a mean of 10.62 km and a range of distance classes from 0-1 to 19-20 km. The actual dimensions of the available area are approximately 40 x 5 km in winter and 90 x 40 km i n summer and would have generated frequency d i s t r i b u t i o n s with higher means and ranges. The Kolomogrov-Smirnoff 2-sample test on was used to compare d i s t r i b u t i o n s . 45 0 Results Overlap of consecutive annual home-ranges One b u l l l e f t the BRV and did not return subsequently (emigration rate 0.008). The remaining animals either stayed in the BRV throughout the year, or made seasonal migrations from the BRV with travel distances of up to 74 km (Tables 2,6)* A l l radio-tracked resident and migrant elk had overlapping annual home-ranges in consecutive years (Table 7) . Annual home ranges of 26 elk tracked for 3 years, also had overlapping ranges. The 2 radio-tagged elk with known home-ranges in 1981 had ranges that overlapped with the 1981 ranges in 1986-89. Two cows o r i g i n a l l y marked in Kootenay National Park were recovered within the BRV during the study. One was radio-tagged 13 years previously, when she wintered 73 km to the west along the Kootenay River. The other was ear-tagged 9 years e a r l i e r just west of the Continental Divide approximately 5 km from where she was recovered. Both cows were 2-year-olds when o r i g i n a l l y marked. Distances between inter-annual seasonal ranges Average inter-annual distances between consecutive a c t i v i t y centres for winter, summer, and rutting ranges were from 3.0-4.2 km. The frequency d i s t r i b u t i o n s were skewed to the right (Table 8, Figures 4 ,5 ,6 ) . There were no s i g n i f i c a n t differences in the frequency 46 d i s t r i b u t i o n s of cows and b u l l s , or of migrants and residents. The mean inter-annual distances between seasonal centres of a c t i v i t y were s i g n i f i c a n t l y less than predicted by the random model in a l l cases (Table 8, Figures 4 ,5 ,6 ) . The majority (75%) of migrant elk returned to the BRV before or during the rut (Table 9) . Therefore, residents and migrants were sympatric at this time and most (87%) radio-tracked elk were in the BRV. Of the 6 cows which remained out of the BRV during 1 or more rutting seasons, 3 returned to the BRV for at least 1 rutting season. The 3 which never returned during the rut had the longest migration distances recorded. Of the 3 b u l l s not in the BRV during 1 or more ruts, 1 died after 1 year. Another migrated during the l a s t year of the study to an adjacent elk population 13 km west of the Continental Divide during the rut and la t e r returned to the BRV. The l a s t animal was out of the BRV during the rut during the f i r s t 2 years of the study, and then returned to the BRV during the l a s t rutting season. 47 Table 6. Numbers of migrant (MIG) and resident (RES) radio-tracked elk a in Banff National Park, 1986-88. Status 1986 1987 1988 MIG RES MIG RES MIG RES Bull 9 3 8 4 9 3 Cow 9 17 13 18 13 18 Both 18 20 21 22 22 21 a includes three cows which switched between migrant and resident status between years 48 Table 7. Overlap (%) a of annual home-ranges during consecutive years for radio-tracked elk in Banff National Park, 1986-88. Elk group Nb Overlap  Years Mean Min. Max. Cows 1986- 87 21 54 16 80 1987- 88 28 55 24 78 1986-88 20 40 15 63 Bulls 1986- 87 8 55 43 71 1987- 88 10 69 42 90 1986-88 6 44 25 70 Both sexes 1986- 87 29 54 16 80 1987- 88 38 59 24 90 1986-88 26 40 15 . 70 a percentage of aggregate area overlapped by consecutive ranges b sample size Table 8. Distances (km) between inter-annual home-range centres 8 during consecutive years for radio-tagged elk in Banff National Park, 1986-88b. Elk group Nb Distances between seasonal centres 0 Season Mean Min. Max. A l l elk Winter 117 3.0 0.0 24.6 Summer 73 4.2 0.1 33.7 Rut 69 3.0 0.0 12.1 A l l residents Winter 60 2.7 0.0 18.8 Summe r 37 2.3 0.1 6.6 Rut 33 2.6 0.0 8.7 A l l migrants Winter 50 3.5 0.3 24.6 Summe r 31 6.8 0.2 33.8 Rut 31 3.3 0.6 12.1 A l l GOWS Winter 88 2.7 0.0 18.8 Summe r 55 4.7 0.1 33.7 Rut 51 3.2 0.5 12.1 A l l b u l l s Winter 29 3.9 0.1 24.6 Summe r 18 2.9 0.2 12.1 Rut 18 2.1 0.0 9.8 Model d 1000 10.8 0.2 19.3 8 winter-winter, summer-summer, rut-rut b data for a l l years pooled c harmonic centres d random pairs in a 196 km2 square 50 Table 9. Number of radio-tagged elk within and outside the BRV during the rutting seasons, 1986-88. Location Year  Class 1986 1987 1988 Within BRV Resident cow 17 18 18 Migrant 8 cow 4 9 10 Resident b u l l 3 4 3 Migrant 8 b u l l 7 7 8 Total in BRV 31 38 39 Outside BRV Migrant cow 5 4 3 Migrant b u l l 2 1 1 Total out of BRV 7 5 4 Total Tracked 38 43 43 Percent in BRV 82 88 91 returned to the BRV before or during the rut F i g u r e 4. D i s t a n c e s b e t w e e n c e n t r e s o f c o n s e c u t i v e w i n t e r r a n g e s . 52 DISTANCE CLASS (KM) F i g u r e 5. Distances between centres of consecutive summer ranges. 53 0.5 T -0.4 U 0.3 UJ => o UJ Cd ^ 0 2 Cd 0.1 +-0.0 -•-MALES •0--FEMALES -A—RESIDENTS •MIGRANTS -MODEL 0 1—I I 1 I—I—I—I—r-8 12 16 DISTANCE CLASS (KM) 20 F i g u r e 6. D i s t a n c e s between c e n t r e s of c o n s e c u t i v e r u t t i n g r a n g e s . 54 Discussion Seasonal ranges During 4 consecutive years in the Upper Red Deer River valley, Morgantini (1988) observed that 11 of 18 radio-tagged elk returned to the same summer ranges and 16 of the 18 returned to the same winter ranges. However his c r i t e r i a for defining "return" were not presented. Si m i l a r l y , Knight (1970) found that most (but not a l l ) adult elk in his Montana study area, returned to the same winter and summer ranges. The results obtained in my study confirm that most adult elk pe r s i s t e n t l y centre their seasonal a c t i v i t i e s on the same ranges from year-to-year. Greenwood's (1980) hypothesis suggests that elk have male-biased dispersal, and female- and resident-biased r e l a t i v e philopatry. Although the only radio-tagged elk to permanently leave the BRV during the study was an adult b u l l , adult cow dispersal was demonstrated for 2 animals immigrating to the BRV from Kootenay National Park. These data are too meagre to support or refute the hypothesis of sex-biased adult dis p e r s a l . There were neither female- or resident-biases in r e l a t i v e philopatry of adult elk, as predicted by Greenwood's hypothesis. Based on their work on white-tailed deer in Michigan, Nelson and Mech (1987) suggested that the re s t r i c t e d movements made by many large mammals could lead to the development of demes with l i t t l e reproductive contact. Amongst the radio-tagged elk followed in this study, most animals centred their rutting season a c t i v i t i e s within a few km of their previous rutting season range and most of the population was sympatric during the rut. However, a few radio-tagged elk did not return to the BRV during the rut and could have bred either with other BRV migrants, or with elk from other populations sharing a common summer/rut range. This i s i l l u s t r a t e d by radio-tracked elk sharing a summer/rut range in the Kananaskis Lakes area of Alberta and returning to winter ranges 200 km apart near Sparwood, B r i t i s h Columbia and Canmore Alberta (Woods and Hlady 1 9 8 8 ; T. Nette, Alberta Fish and W i l d l i f e , unpubl. data). Since some elk alternated between rutting in the BRV and other areas, and some alternated between resident and migrant status (Chapter 2 ) , return migrations may promote gene flow. Although breeding dispersal was rare amongst these elk, natal dispersal rates in elk have not yet been studied. Amongst most mammals, young bulls are thought to be the predominant dispersal group. In the case of elk, young of both sexes t y p i c a l l y stay with their mothers for the f i r s t year and become sexually mature after 2 years. Therefore, future research on philopatry should focus on 1 - , 2 - , and 3-year-olds, e s p e c i a l l y males. 56 Definitions of philopatry In most studies mentioning persistent use of annual or seasonal home-ranges by elk, home-range overlap i s the i m p l i c i t d e f i n i t i o n of philopatry. Edge et a l . (1985) e x p l i c i t l y defined s i t e f i d e l i t y (-philopatry) as any overlap. However, since a l l of their study animals (31 cows) had overlapping ranges, the issue of adjacent, non-overlapping ranges and dispersers was not addressed. In the present study, of 68 cases where year-to-year overlap of annual home-ranges could be determined, 67 overlapped. The problem with the overlap d e f i n i t i o n i s that i t disregards the distance moved. Clearly, animals moving to adjacent home-ranges have di f f e r e n t consequences for both population ecology and gene flow than do long-distance dispersers. Both the modified Cole's association index (values 0 to 1) used by Edge et al . (1985) and the percent overlap (values 0 to 100) used in the present study, are d i f f i c u l t to use in assessing the degree of philopatry. Indices of overlap of successive home-ranges are highly dependent on the shape of the home-range and the method used to calculate the range. Although the 100 % minimum convex polygon i s commonly employed to map home-ranges, i t i s also very susceptible to d i s t o r t i o n due to outlying points (Dixon and Chapman 1980). A single outlying point can greatly a l t e r the calculated 57 area of overlap. It i s also possible that the i d e n t i f i e d area of overlap f a l l s in an area not used during either year and i s an a r t i f a c t of the mapping technique. Another problem i s that of scale. In a small study area, home-range could overlap by chance, whereas in a large area this i s unlikel y . Bunnell and Harestad's (1985) d e f i n i t i o n of dispersal as a distance greater than any observed return migration (5 km) for black-tailed deer on Vancouver Island, i s arbit r a r y and l i m i t i n g . In the present study, migrants t r a v e l l e d up to 74 km; and in other areas of the western mountains, migrations of up to 100 km are quite common (Boyce 1989) . In Europe, migratory red deer travel up to 150 km (Langvatn and Albon 1986) . Of the 3 elk that changed their winter ranges during t h i s study (1 emigrated from the BRV and 2 immigrated to the BRV), a l l were within the known maximum migratory range for the species. Application of distance rules based on home-range diameters (Shields 1987) are subject to problems of mapping technique and determination of the number of diameters home-range diameters which create the s i g n i f i c a n t distance. As Arcese (1989) pointed out, home-range sizes may vary with population size, and therefore do not provide a universal scale for i n t e r - s p e c i f i c comparisons. In addition, empirical data on song sparrow (Melospiza melodia) dispersal did not 58 demonstrate that relatedness decreased with numbers of t e r r i t o r i e s traversed in a study of short-range dispersal (Arcese 1989). Approaching the problem of philopatry from a p r o b a b i l i s t i c point of view accepts that philopatry and dispersal are relat i v e rather than absolute terms. Therefore, an observed d i s t r i b u t i o n can be tested against another observed or hypothetical d i s t r i b u t i o n . In the present study, this allows the comparison of residents with migrants, bulls with cows, and the population as a whole with a random model. I conclude that r e l a t i v e to a random d i s t r i b u t i o n , this population of adult elk i s p h i l o p a t r i c . 59 CHAPTER 4. NET COST OF MIGRATION Introduction Migration should be favored when the benefits of moving outweigh the benefits of not moving (Baker 1978). Potential benefits of seasonal migration include: 1) access to better quality and quantity of food; 2) reduced competition for food; 3) escape from predation and other forms of harassment; 4) access to increased mating opportunities; and, 5) avoidance of severe seasonal weather ( S i n c l a i r 1983, Festa-Bianchet 1988, S i n c l a i r and Fryxell 1988, Chapter 3). Potential costs of migration include: 1) energy requirements for travel between ranges; 2) time l o s t from other a c t i v i t i e s ; and, 3) increased mortality due to travel hazards and predation. Within a species, a l l individuals may not face the same costs and benefits. For example, the cost/benefit ratio evaluation may vary with the size of the individual (Tucker 1975) and individuals may vary in their genetic tendencies to move (Berthold and Querner 1981). Migrations by t e r r e s t r i a l mammals are generally less extensive (<100 km) than migrations by birds, and are assumed to be less costly energetically (Baker 1978), although these migration costs have rarely been quantified. However, there i s a growing body of information on the cost of locomotion, p a r t i c u l a r l y for large ungulates (Parker et a l . 1984, Gates and Hudson 60 1 9 7 8 ) so i t i s now possible to combine empirical data on migration with physiological information to estimate the energetic costs of migration. P a r t i a l migrants (species that include both residents and migrants) offer the opportunity to compare net costs and benefits of migration. In this study, I used data on movements of residents and migrants to calculate energy and travel time costs and asked the following questions: 1 ) Are energy and time costs for migration s i g n i f i c a n t r e l a t i v e to other l i f e - h i s t o r y costs?, 2 ) How do energetic costs of migration vary between age/sex classes? and, 3 ) Do the r e l a t i v e costs of migration allow predictions on the r e l a t i v e mobility of age/sex classes? 6 1 Methods Energy calculations The net energy cost of migration was calculated as the estimated locomotion cost for migration less the estimated locomotion cost expended by non-migrating elk during the same time i n t e r v a l . Energy expenditures (kcal-kg - 1-km - 1) for locomotion on lev e l and up-sloping ground were computed using the equations for elk presented by Parker et a l . (1984): E l e v e l =2.97 x M<-">(kg) (1); E u p =3.57 x M<-">(kg) (2). Resident cow elk made only i n s i g n i f i c a n t v e r t i c a l movements during the spring migration period (results to follow), and (1) was used to estimate their movement energy expenditures. For migrant elk, I considered migrations to consist of a 4 km upslope component resulting in a 1 km v e r t i c a l rise (Chapter 2), and a lev e l component for the remainder of the migration distance: =(E u p x 4) + ( ( E l e v e l x (Dist - 4 ) ) (3), where Dist i s the estimated 1-way travel distance in km. I scaled the net energy expenditures for migration by d a i l y energy expenditures estimated for resting elk using the equation presented by Bobek et a l . (1983) to estimate kcal-kg _ 1-day _ 1 : E r e s t = 90 x M<°-75»(kg) (4). 62 Time and distance estimates On an annual basis, average horizontal distances traveled by migrant and resident elk (Chapter 2, Table 2) were used to calculate average time requirements for movements. Average hourly speeds were not determined but assumed to be 3.0 km-h-1, a rate observed by Gates and Hudson (1978) in an experimental si t u a t i o n . This i s s l i g h t l y faster than the 2.5 km-h-1. rate observed by Craighead et a l . (1973) for free-ranging elk. During May and June 1988, elk with previous h i s t o r i e s of migratory behavior were monitored on a da i l y basis within the valle y . When their radio signals disappeared, an attempt was made to track them each day u n t i l they reached their summer range. Incidental observations of radio-tracked elk allowed additional calculations of travel times. During May 1988, I followed 10 resident cows (1 switched to migrant status l a t e r i n the summer) on a da i l y basis and estimated their net d a i l y movements. For both migrants and residents, the d a i l y movement distance was assumed to be a straight l i n e between relocation points. Since observations were made at comparable times during consecutive days, observed travel times are expressed i n km-day-1. The net time cost of migration was calculated as the estimated time for migration less the estimated time spent moving by non-migrating elk during the same time 63 i n t e r v a l . Body weights and sex/age classes Body weights were obtained from f r e s h l y - k i l l e d , intact elk which had died in c o l l i s i o n s with automobiles and trains within the BRV. In most cases, the seasonal movement status of the dead elk was unknown. These data were partitioned into seven categories based on age and sex: newborn calves (June), July calves, September calves, 1-year-olds, 2-year-olds, >2-year-old cows, and >2-year-old b u l l s . The newborn weight was that of the largest fetus measured during the study. July c a l f weight estimated the mass of a young elk shortly after making i t s f i r s t upslope migration (in June) and data from September weights estimated the masses of calves during their f i r s t return journey. 64 Results Migration time and speed Most migrations between winter and summer ranges took 3 days or less. An exception was a cow elk that calved between ranges. She did not move for 3 weeks after calving and then moved to her summer range. in May and June 1988, migration speed of b u l l s averaged 7.1 km-day-1 and cows averaged 9.2 kmday - 1 (Table 1 0 ) . The farthest movement in a single day recorded during the study was 24.2 km (an adult cow). Travel speed in km-h-1 could be determined in a single case where an adult cow moved 21.43 km in 4 h (5.1 km-h"1) . Body weights The largest elk fetus examined in the study weighed 21 kg. July and September weights for calves averaged 51 kg (N=7) and 104 kg (N=8) respectively, while 1- and 2-year-old elk averaged 165 kg (N=>21) and 202 kg (N=ll). Cows and bu l l s >2 years old weighed an average of 238 kg (N=29) and 339 kg (N=22) respectively, and were s i g n i f i c a n t l y heavier than other age classes ( t - t e s t , P<0.05). Net costs Net energy costs for migration based on annual movements averaged 12.84 kcal-kg- 1 for bulls and 15.33 kcal-kg- 1 for adult cows (Table 11). The r e l a t i v e net energy requirement for migration decreased with 65 increasing body size (Figure 7) . The net energy cost for migration based on observed cow elk movements during the spring of 1988 was 8.86 kcal-kg _ 1-day _ 1 (Table 10). Making the assumption that resident b u l l s moved similar distances as resident cows, the net energy cost for migration by b u l l elk was 7.47 kcal-kg _ 1-day - 1. The calculated d a i l y resting energy requirements for adult cow and b u l l elk were 22.91 and 20.98 kcal-kg - 1-day - 1 respectively. Net time requirements for migration based on annual movements averaged 6.3 h for bulls and 7.3 h for cows (Table 11). 66 Table 10. Daily travel distances of resident 8 elk compared to d a i l y travel distances of migrant 1 5 elk during migration, May and June, 1988. Sex Status Sample km/day (S.E.) Cow Resident 10 1.2 (0.15) Migrant 8 9.2C (1.28) Bull Migrant 7 7.1c (1.31) 8 average straight l i n e distance between locations measured d a i l y May 16-22, 1988 b average straight l i n e distance between locations measured while the elk was migrating, May 18-June 18, 1988 c s i g n i f i c a n t l y d i f f e r e n t from resident average (Wilcoxon pairs test, P<0.05) 67 Table 11. Net 1-way travel times (h) and energy requirements (kcal-kg - 1) for migrant elk based on annual movements of migrant and resident radio-tracked elk. Net requirement Bull Cow Time 8 Average 6.3 7.3 Minimum 5.1 2.4 Maximum 16.6 12.7 Energy15 Average 12.84 15.33 Minimum 10.38 10.25 Maximum 25.54 22.72 8 computed from data presented in Table 2 Chapter 2 and an estimated walking speed of 3 km-h-1 b computed from data presented in Table 2 Chapter 2 and formulae presented in the text 68 30 T . A . B 2 0 -. C . D ci . E . F . G 10 0 0 100 200 KILOGRAMS 300 400 Figure 7. Net cost (Real.kg) of an upslope migration (36 km) for various weight classes of BRV elk. (A=new-born c a l f , B=July c a l f , C=September c a l f , D=l-year-old, E=2-year-old, F=>2-year-old cow, G=>2-year-old bull) 69 Discussion Bobek et al . (1983) modeled the energy budget of BRV elk and calculated the annual energy requirements to be 3.09 x 106 kcal for adult bulls and 3.57 x 106 kcal for pregnant cows. Based on annual movements, the average 1- way net cost of upslope migration estimated in thi s study represents 0.14% and 0.10% respectively of these requirements. Based on the dire c t comparison of migrant and resident elk movements in the spring of 1988, the net migration cost for cow elk was 0.06% of the estimated annual energy requirement. In comparison, Bobek et al . (1983) estimated costs of reproduction for adult cows as 0.67 x 106 kcal or 18.8% of the annual energy budget of pregnant cows. Nelson and Leege (1982) presented data suggesting that the annual energy expenditure for a pregnant cow elk would be approximately 2.77 x 106 kcal, of which 14.5% (0.40 x 106 kcal) would be required for pregnancy and la c t a t i o n . Relative to the costs of reproduction, I conclude that net energy costs for migration in BRV elk are t r i v i a l . The horizontal and v e r t i c a l distances traveled by migrating BRV elk are within the observed ranges for other elk populations and other North Temperate Zone cervids (Table 12). North Temperate Zone cervids have reported average migration distances in the range of 2- 67 km horizontally and 200-1000 m v e r t i c a l l y . With the 70 exception of tundra caribou which travel up to 2500 km, reported maximum migration travel distances for other cervids ranged 3-150 km. For the maximum reported migration distance in Cervus elaphus (150 km, Norway, Table 12), the energy investment would be less than 1% of the species' annual energy budget. Most migrations would require much less then 1%. Speed of travel Descriptive accounts of the speed of migration i n elk present c o n f l i c t i n g views. Brazda (1953) described elk migration as "movements to higher elevations as the season progressed", while Peek and Lovaas (1968) reported migration started in August and ended i n November and December. Other authors report migration as a rapid event, sometimes accomplished overnight (Knight 1970, Hudson and Morgantini 1988). S i m i l a r l y , reported speeds for other North American cervids vary from "slow" (e.g. mountain caribou, Edwards and Ritcey 1956) to "fast" (black-tailed deer, Harestad 1979; tundra caribou, Fancy et a l . 1989). While some of this discrepancy may be due to i n t e r - s p e c i f i c and inter-population differences, i t i s also possible that i t i s in part an a r t i f a c t of observation technique. Actual calculation of speed of travel requires the type of continuous data usually only available from i n d i v i d u a l l y radio-tracked animals. The fastest cervid migration speed I have found in 71 the l i t e r a t u r e i s 40 km-day-1 for tundra caribou (Fancy et a l . 1989). They reported pregnant eows averaged 7-24 km-day-1 while migrating. This i s similar to the speeds observed for elk during my study (2-25 km-day-1). The rapid migrations made by some cervids are not consistent with the view that migrants follow the growth of their food plants upslope in the spring (Harestad 1979). However, i f f a m i l i a r i t y with s p e c i f i c ranges (summer and winter) i s an advantage to individuals (knowledge of food patches, escape routes etc . ) , then rapid migrations may serve to minimize time spent in r e l a t i v e l y unfamiliar or unfavorable t e r r i t o r y . Relative energy costs Migrant cows may give b i r t h on either summer or winter ranges, or on route between ranges. Newborn elk do not migrate immediately after b i r t h (Knight 1970, Adams 1982, Boyce 1989). By late June or July, calves have more than doubled their weight and reduced their r e l a t i v e energy investment in migration (Figure 7) . If they are born on the winter range or between ranges, they then accompany their mothers to the summer range. Energy investment in migration decreases r e l a t i v e to body size (Figure 7) suggesting that r e l a t i v e mobility might increase with body size. However, adult cows are usually accompanied by young (Woods 1990) and their mobility should be thus constrained. Therefore, mobility as predicted by rel a t i v e energy investments, would predict adult b u l l s , non-lactating adult cows, 2-year-olds, and 1-year-olds to be the most mobile groups. In BRV elk, the majority of adult b u l l s followed i n thi s study were migrants whereas most adult cows were residents (Chapter 2). Furthermore, the most complex migration patterns (up to 3 cycles per year) were seen in adult b u l l s . Migration distance did not vary s i g n i f i c a n t l y between the sexes. Although i t i s poorly documented, dispersal in elk i s thought to be most prevalent in 1- and 2-year-old males (Flook 1970). 73 Table 12. Horizontal and v e r t i c a l distances* reported for migrating elk and other North Temperate cervids. Species Lateral(km) Vertical(m) Source Area Av. Max. Min. Av. Max. Min. Elk CND Rockies 32 74 CND Rockies 50 69 CND Rockies - 100 USA Rockies - 129 Norway - 150 Mule Deer USA Nevada - 150 Black-tailed Deer CND BC 2 3 White-tailed Deer USA, Various 6 89 Caribou USA, Alaska, Tundra - 2500 CND, AB, woodland 15 26 CND, AB, Mountain 67 134 CND, BC, Mountain Moose CND, AB CND, BC USA Alaska Europe Sweden - 20+ 64 - 100 - 150 14 14 951 1432 392 26 1000 2 800 1067 610 - 1219 1 200 400 13 5 33 60 1067 - 1372 t h i s s t u d y M o r g a n t i n i l 9 8 8 Woods and Hlady 1988 Adams 1982 Langvatn and A l b o n 1986 G r u e l l and Papez 1963 H a r e s t a d 1979 Marchinton and Hirth 1984 Fancy et a l . 1989 Edmonds 1988 Edmonds 1988 Edwards and Ritcey 1959 Hauge and Keith 1981 Edwards and Ritcey 1959 Van Ballenberghe 1977 Pullianen 1974 Sandegren and Bergstrom 1983 one-way 74 CHAPTER 5. DIET COMPOSITION AND QUALITY Introduction A l t i t u d i n a l migration i s a common behavior amongst North American ungulates l i v i n g in mountainous areas (Fryxell and S i n c l a i r 1988). Several possible benefits have been proposed for these migrations, the most favored being a n u t r i t i o n a l advantage over residency (Klein 1965, Hebert 1973, Morgantini and Hudson 1983). Since forage quality generally peaks in young, rapidly growing plants and then declines as the plants mature (Johnston et a l . 1968, Nelson and Leege 1982), animals could benefit from tracking early growth stages as spring advances from valley bottom to alpine. In thi s case, then at any time during the growing season, migrant animals should have access to more nutritious forage than i f they had remained on low elevation ranges. Empirical data for various North American ungulates on forage quality, animal condition, and animal movements, do not consistently support the hypothesis that migrants have access to better quality food. In bighorn sheep ranges in Western North America, forage q u a l i t y may be higher (Hebert 1973), lower (Festa-Bianchet 1988), or similar (Krausman et a l . 1989) in high elevation s i t e s compared to low elevation s i t e s at the same time. On elk summer ranges d i f f e r i n g substantially in al t i t u d e , plant species, and vegetation 75 structure, Baker and Hobbs (1982) found consistency in diet quality and suggested that elk can generalize their choice of feeding habitats without incurring a loss in diet quality. Langvatn and Albon (1986) observed that migrant elk returning to low elevation ranges were heavier than residents in a p a r t i a l l y migratory Scandinavian population. However, as Morgantini (1988) noted i n his study of elk in the Canadian Rockies, even in populations where a n u t r i t i o n a l advantage for migrants i s suspected, the coexistence of healthy non-migratory elk suggests that non-migratory individuals may be able to compensate (e.g. dietary changes, increased intake, increased s e l e c t i v i t y ) . Poor correlation with n u t r i t i o n a l quality i s suggested by Brazda's (1953) observation that most migrant elk in a Rocky Mountain population did not arrive on high elevation ranges u n t i l plants were in lat e r stages of development. Radio-tracking studies showing rapid (1-3 days) spring migration by blac k - t a i l e d deer on Vancouver Island (Harestad 1979) and by elk in this study (Chapter 4), do not suggest that these species follow the phenological progression of green-up between low and high elevation ranges. Demonstration of a n u t r i t i o n a l advantage for migrant individuals requires some estimation of what their diet composition and quality would have been, had 76 they not migrated. However, i f a l l animals migrate (e.g. Hebert 1973), or i f only the migratory component i s studied (e.g. Morgantini 1988), then comparisons rely on assumed diets of "residents". Simultaneous comparison of the diet composition and quality for migrant and resident elk within a p a r t i a l l y migrant population, offers a more dire c t method of investigating the n u t r i t i o n a l advantage hypothesis. In this study, I followed migrant and resident elk from a single population and compared diets and forage quality on low and high elevation ranges. I addressed the question: during the part of year when migrants are separate from residents, do migrants have higher qua l i t y forage i n their diets? 77 Methods Fecal c o l l e c t i o n s Fresh elk feces were collected at 2-4 week interv a l s from October 1985 through December 1988. Each c o l l e c t i o n consisted of at least 3 p e l l e t s from 15 p e l l e t groups. Collection sites included 5 locations within the montane zone (representing ranges used by resident elk) and 2 areas in the upper subalpine (representing ranges used by migrant e l k ) . Samples for diet analysis were a i r dried and diet composition was assessed through fragment analysis at the Composition Laboratory, Colorado State University, Fort C o l l i n s , Colorado. Estimates of percent dry weight were obtained using the technique described by Sparks and Malechek (1968). Each f i e l d sample was sub-sampled by making 3-5 slides and examining 20 f i e l d s per s l i d e . Although this technique may under-represent forbs and leaves (Leslie et a l . 1983, Putnam 1984), I assume that any bias i s constant for comparison of diets of migrant and resident elk. This analysis could not discriminate between Shepherdia (buffaloberry) and Eleagnus (wolf-willow). However, Eleagnus was re s t r i c t e d to mesic si t e s in the lowest areas of the montane within the BRV. Shepherdia  canadensis was widespread in dry sit e s in the montane and occurred through to upper subalpine elevations. Since elk were frequently observed eating Shepherdia 78 leaves and since Shepherdia/Eleagnus occurred as a major diet item in montane and subalpine s i t e s , a l l c o l l e c t i o n s were assumed to be Shepherdia rather than Eleagnus. Forages were grouped into 4 classes: 1) grasses (grasses and sedges), 2) deciduous trees and shrubs, 3) evergreen trees and shrubs, and 4) forbs and miscellaneous items. Major forage items were i d e n t i f i e d as any genus (or group) constituting more than 5% of a sample (Collins et a l . 1978). Seasonal differences in diet composition in the montane and subalpine were examined by pa r t i t i o n i n g the data into 4 seasons: 1) winter (December-February), 2) spring (March-May), 3) summer (June-August); and, autumn (September-November). Crude nitrogen was used as an estimate of protein content in fecal samples (crude protein = 6.25 x nitrogen, Nelson and Leege 1982). Samples for chemical analysis were frozen and shipped to Norwest Laboratories, Edmonton, Alberta or to Cro s s f i e l d Laboratories, C r o s s f i e l d , Alberta. Organic nitrogen was determined on a dry weight basis using the semi-automated Kjeldahl technique (A.O.A.C. 1965). Although fecal nitrogen has been c r i t i c i z e d as an indicator of.diet quality (Robbins et a l . 1987), i t has been widely used in elk studies (e.g. Gates and Hudson 1981, M e r r i l l et a l . 1987, Morgantini 1988) and i s po s i t i v e l y correlated with dietary nitrogen. Holechek et 79 a l . (1982) reviewed the use of fecal indices and concluded that they are useful in the study of re l a t i v e differences (such as my study), rather than absolute values. Forage co l l e c t i o n s Based on major diet items i d e n t i f i e d in fecal samples during 1986-87, 3 diet items were selected for forage analysis in 1988: 1) mixed grasses (Gramineae); 2) Salix spp.; and 3) Shepherdia. At 1 subalpine and 2 montane areas (A and B), habitat patches a c t i v e l y used by elk were i d e n t i f i e d by dire c t observation of feeding elk or fresh feces, or by the presence of radio-tagged elk (Chapter 2) . Forage samples were collected as close as possible to these patches. Observers walked across the patch ( t y p i c a l l y less than 100 m) and hand-plucked 50 sub-samples (approximately 1 gm each). These samples were fresh frozen and sent to Cro s s f i e l d Laboratories, C r o s s f i e l d , Alberta for analysis. For grasses, cured stems and leaves were picked only i f green material was not available. For shrubs, current year stem growth was picked only i f leaves were not available. Although the observers may not have been as selective as feeding elk, the same observers made a l l collections and I assume that t h i s bias was constant between areas. Forage quality was assessed on the basis of parameters commonly used in range studies: nitrogen, 80 c e l l solubles, l i g n i n , phosphorus, and calcium (Nagy and Haufler 1980). Nitrogen was used as an index of crude protein and i s recognized as a c r i t i c a l nutrient for herbivores. C e l l solubles are 98% digestible and l i g n i n i n h i b i t s c e l l u l o s e d i g e s t i b i l i t y . Phosphorus and calcium are important for growth (especially of antlers) and phosphorus i s often a l i m i t i n g nutrient. Organic nitrogen, phosphorus, and calcium were determined using the semi-automated Kjeldahl technique (A.O.A.C. 1965). Neutral detergent fiber (NDF), acid detergent fiber (ADF), and l i g n i n were assayed as described by Robbins and Moen (1975). C e l l solubles were calculated as 1-NDF. Differences in forage and fecal quality parameters were compared using the Wilcoxon signed ranks matched pairs test . Estimates of diet quality Diet quality i n the subalpine and montane was estimated by assuming that 100% of the diets were composed of Shepherdia, Salix spp., and grasses. These forage components constituted approximately 92% of a l l subalpine elk diets and 94% of a l l montane di e t s . For each s i t e and forage component, a monthly diet q u a l i t y estimate was calculated by multiplying the percentage occurrence of a forage type represented in elk di e t s , by the diet quality parameter for that s i t e . The results for the 3 forage components were then added together. 81 Results Diet composition Grasses and sedges were major components of elk diets in both subalpine and montane habitats throughout the year (Table 13). However, in both habitats grasses and sedges were eaten more during the winter, spring, and autumn, than during the summer (Figure 8) . There was no s i g n i f i c a n t difference in the relati v e use of grasses and shrubs between the habitats at any season. The rel a t i v e use of grasses and sedges in the subalpine (mean=38.5%, S.E.=4.75) during the summer was considerably greater than rela t i v e use in the montane (mean=13.8%, S.E.=4.14), although this was not quite s t a t i s t i c a l l y s i g n i f i c a n t (P=0.06). Deciduous shrubs were major components of elk diets during spring, summer, and autumn in the subalpine and during a l l seasons in the montane (Table 13). Use of Salix spp. was high in both habitats during the summer and moderate in the montane during the autumn. Shepherdia was used frequently in the subalpine during the summer and in the montane during the spring, summer and autumn. In both habitats, use of shrubs peaked during the summer months when shrubs were in leaf (Figure 8) . There were no si g n i f i c a n t differences in the rel a t i v e use of shrubs between habitats during any season. The relati v e use of shrub leaves in the subalpine (mean=55.4%, S.E.=8.34) during the summer 82 months, was considerably less than the re l a t i v e use i n the montane (mean=83.1%, S.E . =5 .02 ) , although this was not quite s t a t i s t i c a l l y s i g n i f i c a n t (P= 0 . 0 6 ) . Coniferous trees and shrubs were not eaten by elk in the subalpine during any season, and were only l i g h t l y used by elk in the montane during winter, spring, and autumn (Table 13). Equisetum use was l i g h t to moderate during spring and autumn in the subalpine, and during winter, spring, and autumn in the montane. Legumes and mosses were l i g h t l y used during the spring in the montane, but their use did not exceed 5% during other seasons in the montane, or during any season in the subalpine. 83 Table 1 3 . Major 8 components of elk diets during 4 seasons b on subalpine and montane ranges. Plant Subalpine Montane W Sp Su A W Sp Su A Grasses and sedges Agropyron L L L Carex H H M M H H M Festuca L M L H M M L M Juncus L Koeleria L L . . L . . L . . . Poa M L . . M L M L L Stipa/Oryopsis • • L • • • • L H L L Deciduous trees and shrubs Populus Rubus Salix Shepherdia H H L H H H M H Coniferous trees and shrubs Picea L L . . . . Pinus L L . . L Pseudotsuga L L . . L Other plants Equisetum .. L .. M L L .. L Medicago-Melilotus .. L . . . , Moss . . L . . . 8 a food component constituting >5% of any sample b symbols: W=winter, Sp=spring, Su=summer, A=autumn, H=maximum occurrence 75 -100%, M=maximum occurrence 25 -74%, L=maximum occurrence 6-24% 84 Figure 8. Grass and deciduous shrubs in elk diets collected in subalpine and montane habitats, 1988. (•—• subalpine grass;* •montane grass;A—Asubalpine deciduous shrubs;0—O montane deciduous shrubs; horizontal bars indicate period between median migration dates) 85 Forage quality indicators  Nitrogen Fecal nitrogen values in both the subalpine and the montane peaked during the summer (Figure 9) . In 1988, paired samples from the subalpine and montane tracked each other closely (Figure 9) and were not s i g n i f i c a n t l y d i f f e r e n t . Forage nitrogen in Shepherdia > Salix spp., and grass in the subalpine and the montane peaked in June (Figure 10). Mean forage nitrogen values during the period May-September in Shepherdia and Salix spp. were similar in subalpine and montane habitats (Table 14). At Montane B s i t e , the forage nitrogen in grass was s i g n i f i c a n t l y less than the corresponding value from the subalpine s i t e in this period but not s i g n i f i c a n t l y d i f f e r e n t from Montane A. Estimated nitrogen in elk diets in subalpine and montane environments peaked in June and was lowest in A p r i l and November (Figure 10). Phosphorus Phosphorus content in Shepherdia, Salix spp., and grass in the subalpine and the montane peaked in June and declined through November (Figure 11). Mean forage phosphorus during the period May-September for Shepherdia was similar in subalpine and montane habitats (Table 14). Mean forage phosphorus during this period for Salix spp. and grass was s i g n i f i c a n t l y lower at s i t e 86 Montane A than in the subalpine s i t e . Phosphorus i n Montane s i t e B was similar to Montane s i t e A, but not s t a t i s t i c a l l y d i f f e r e n t from phosphorus in the subalpine s i t e (P -0 .05) . Estimated phosphorus in elk diets peaked in June-July and was lowest in A p r i l and November in both subalpine and montane environments. Phosphorus levels were generally higher in estimated subalpine diets (Figure 11). Calcium Forage calcium in Shepherdia, Salix spp., and grasses in the subalpine and the montane generally increased from spring through autumn (Figure 11). Mean forage calcium during the period May-September for Salix spp. and grass was similar in subalpine and montane habitats (Table 14). Shepherdia calcium estimates were s i g n i f i c a n t l y greater in si t e s Montane A and B then in the subalpine s i t e . Estimated calcium in elk diets in the subalpine and montane were lowest in May (Figure 11). C e l l Solubles Estimates of c e l l solubles (1-NDF) in Shepherdia were lowest in the early spring twigs and generally higher in leaves throughout the summer and early autumn in the subalpine and montane (Figure 10). Salix spp. c e l l soluble estimates had less obvious seasonal fluctuation than other nutrient parameters (Figures 87 10,11). In grasses, c e l l solubles were generally highest in mid-June. Mean forage c e l l solubles were s i g n i f i c a n t l y higher in Salix spp. samples from s i t e Montane A and s i g n i f i c a n t l y less in grass samples from s i t e Montane B compared to the subalpine s i t e . Estimated c e l l solubles in elk diets were consistently higher in montane diets than in subalpine diets throughout the period June-September (Figure 10). Lignin Lignin in 3 forage classes d i f f e r e d markedly in i t s seasonal pattern of occurrence. In Shepherdia, twigs were r e l a t i v e l y high in l i g n i n (mid-May, early June) but low in l i g n i n throughout the leaf period (Figure 10). In Salix spp., l i g n i n concentrations were variable with no consistent difference between twig and leaf samples. In grasses, l i g n i n was consistently low throughout the sampling period. Mean forage l i g n i n was similar for a l l 3 forage classes in the subalpine and montane (Table 14) . In estimated elk diets, l i g n i n peaked in mid-July in both the subalpine and montane diets (Figure 10). 88 Table 14. Comparison of 6 measures of forage quality means during the period May-September, 1988a for 1 subalpine (A) and 2 montane (B,C) s i t e s . Plant Subalpine Montane B Montane A Parameter (Elk Meadows) (Duthil) ( H i l l s d a l e ) Salix spp. Nitrogen 2, .49 (0, .36) 2, .67 (0, .20) 2, .59 (0, .34) Phosphorus 0, .40 (0. .06) 0. .28 (0, .05) 0, ,29b (0. .05) Calcium 1, .02 (0. .13) 1, .14 (0, .10) 1, .08 (0, .11) NDF 46, .28 (2, .63) 37, .99 (2, .75) 36, .88 (3, .57) ADF 29, .80 (2. • 77) 29, .77 (1. .79) 34, .81 (3, .12) Lignin 14, .60 (1. .36) 14, .48 (1, .16) 16, .72 (1. .17) Shepherdia Nitrogen 3, .51 (0. .20) 3, .27 (0, .23) 3, .33 (0, .19) Phosphorus 0. .25 (0, .03) 0, .21 (0, .04) 0, .24 (0, .03) Calcium 0, .77 (0, .11) 0, .91b (0, .08) 0, ,87b (0, .11) NDF 36, .79 (4. .96) 33, .68 (3, .61) 30, .37 (2, .79) ADF 25, .32 (2, .61) 23, .61 (1. .99) 24, .03 (2, .08) Lignin 8. .75 (1. .77) 7, .58 (1. .15) 6, .79b (0. .94) Grasses Nitrogen 2, .25 (0. .27) 1, ,43b (0, .13) 2, .13 (0, .31) Phosphorus 0, .23 (0, .03) 0, .13b (0, .02) 0, .17" (0, .03) Calcium 0, .70 (0. ,05) 0, .63 (0. .07) 0, .60 (0, .04) NDF 56, .16 (3, .99) 67, ,07b (1. .62) 60, .01 (4, .65) ADF 31, .66 (1. .26) 34, .59 (1. .96) 34, .96 (1. .94) Lignin 3, .58 (0. .47) 4, .14 (0, .45) 4, .00 (0, .30) a mean % dry weight (standard error) of 9 paired samples from each s i t e , May 15, June 1, June 15, July 1, July 15, August 1, August 15, September 1, September 15 b s i g n i f i c a n t l y d i f f e r e n t from corresponding subalpine measurement (Wilcoxon signed rank test, P<0.05); no s i g n i f i c a n t differences for any Montane A - Montane B compari son 89 Figure 9. Nitrogen in elk fecal samples from montane and subalpine s i t e s in Banff National Park, 1985-88. (horizontal bars indicate period between median migration dates) 90 NITROGEN SHEPHERDIA CELL SOLUBLES 5 1 , /v-Subalpine 1 0 0 n 1 -4 " -^O^V^Montane A 8 0 ; *, 3 H y~>xZ^^ 60 / Mon1 2 - £ tane B 40 0 I I I 1 l I l l I l J J A S H F 20 H M SALIX CD t — I >- I I I I I I I I I GRASS i 11 i 11 i 11 ESTIMATED ELK DIETS Subalpine Montane I I l M l I I I -»M I I I I I I I I I J J A S I I I I I I I I I I I I I I I I I I I I I I 1 I I I I MONTH (1988) LIGNIN 20-i 16-12-8-4-0-1 F 1 M I I I I I I I I 1 J J A S A -I i i l i I i i i i i I l l l I i I I l - A W i i i i i i i i i F i g u r e 10. N i t r o g e n , c e l l s o l u b l e s , and l i g n i n i n montane and s u b a l p i n e forages and es t imated d i e t s , Banf f N a t i o n a l P a r k , 1988. ( h o r i z o n t a l bars i n d i c a t e p e r i o d between median m i g r a t i o n dates of females (F) and males (M)) 91 CALCIUM PHOSPHORUS SHEPHERDIA 2.0-j 1.6-1 .2-0.8-0.4-o.o : 1.0' r-Montane B \ -^Montane A 0.8-0.6-- 1 F 0.2-^ • ~\ M I I I I I I I I i J J A S SALIX CD »——1 >-It! I I 1 I I I I I I GRASS i i 11 11 111 ESTIMATED ELK DIETS Subalpine Montane I I i i M 1 l l 0.0-- t F H M I I I I I I I I I J J A S I I I I i i I I I I I I I I I I I I i ) I I I I I i i MONTH (1988) Figure 11. Calcium and phosphorus in montane and subalpine forages and estimated elk diets, Banff National Park, 1988. (horizontal bars indicate period between > median migration dates of females (F) and males (M)) 92 Discussion Diet composition BRV elk in both subalpine and montane environments r e l i e d on grasses, sedges, and shrubs during 1 or more seasons, consistent with the view that elk are "generalist" (Gates and Hudson 1981) or "intermediate" (Hobbs et a l . 1983) feeders. Although some elk populations (migrants) eat a high percentage of forbs during the summer and grass during the winter (Schuerholz 1984, M e r r i l l et a l . 1987), others eat primarily shrubs in summer and grass in the winter (Morgantini 1988). The importance of forbs in Banff elk summer diets suggested by Flook (1970) was not confirmed by my study or by Morgantini (1988). The prevalence of Shepherdia canadensis in elk diets was surprising since plants rarely appear browsed and the twigs are known to contain high levels of phenolics ( S i n c l a i r et a l . 1982). However, elk use was almost e n t i r e l y r e s t r i c t e d to the period when Shepherdia was i n leaf in the summer and autumn. The phenolic content of these leaves i s unknown as i s the re l a t i v e phenolic content of montane and subalpine plants. High use of Shepherdia has recently been found in Kootenay National Park elk (D. P o l l , Canadian Parks Service, pers. comm.) and i s noted in summer elk diets by Murie (1951) and Martinka (1969). 93 Forage quality As found by Morgantini (1988) and M e r r i l l et al . ( 1 9 8 7 ) , fecal nitrogen of elk in my study generally tracked forage nitrogen. In a l l 3 studies, estimated spring and summer peaks in fecal nitrogen were approximately 4%. Fecal nitrogen winter minima were lowest in Morgantini's study along the Red Deer river (<1%), highest in M e r r i l l et al.'s Mt. St. Helens study (1.6%), and intermediate in my study (1%). Although the Red Deer River population also was p a r t i a l l y migratory, most animals were migrants and fecal nitrogen values were not obtained for resident animals during the summer (Morgantini 1988). Fecal data thus described the diets of migrant elk as they moved from low elevation winter ranges to high elevation summer ranges. Fecal nitrogen on these summer ranges peaked at the same time (July) as did those on both montane and subalpine ranges in the nearby BRV. Nitrogen levels in grass forage samples from high elevation ranges of Red Deer River elk were s i g n i f i c a n t l y higher than in grasses on low elevation ranges at corresponding times (Morgantini 1988). This agrees with my comparison of Montane si t e B and the subalpine. However, unlike Morgantini (1988), I could detect no differences in mean Salix spp. nitrogen l e v e l s during the summer between the low and high elevations. Summer values for Salix spp. nitrogen in the BRV montane 94 were higher than the corresponding low elevation Red Deer River values, perhaps due to drier conditions i n that area. Differences between the n u t r i t i v e values of forage classes appeared to be more s i g n i f i c a n t than differences between ranges within a forage class. This i s i l l u s t r a t e d by the generally low protein content of grasses throughout the summer compared to both Shepherdia and Salix spp. leaves. Forage l i g n i n was lowest in grasses and Shepherdia leaves, and much higher in Salix spp. leaves. Since l i g n i n decreases the d i g e s t i b i l i t y of hemicellulose and cellulose (Nagy and Haufler 1980), this may explain Morgantini's (1988) observation that summer samples of Salix spp. were higher in protein but less digestible than grass samples. Diet quality Although Bobek et al . (1983) suggested that non-browse food was higher in n u t r i t i o n a l value for BRV elk than browse during the growing season, they assumed a hypothetical elk diet that contained many species (e.g. Astragulus, Artemisia) which have not been i d e n t i f i e d as major foods in BRV elk. When actual diet data are combined with n u t r i t i o n a l estimates, the leaves of shrubs are superior foods to grasses in several ways (e.g. nitrogen content). Levels of nitrogen, calcium, and l i g n i n in the 95 estimated diets of migrant and resident elk were sim i l a r . Migrant elk had consistently more phosphorus in their d i e t s . A notable exception to the s i m i l a r i t i e s i s the higher l e v e l of c e l l solubles estimated for low elevation elk during the summer. Since these solubles are estimated to be 98% digestible (Robbins and Moen 1975), resident elk diets were superior to diets of migrant elk in this respect. In a study of summer food composition and diet q u a l i t y of elk in Colorado, Baker and Hobbs (1982) noted consistency in elk diet quality from year-to-year, and between plant communities which varied substantially in al t i t u d e , plant species composition, and vegetation structure. Their hypothesis, that elk can generalize their choice of foraging habitats without suffering n u t r i t i o n a l disadvantage, i s consistent with the results I obtained for migrant and resident BRV elk. N u t r i t i o n a l advantage hypothesis These results do not support the hypothesis that migrants universally gain access to better quality food, even by moving to higher elevational ranges. Considering the extremely large geographical range of the species and the diverse habitats occupied by elk (Geist 1982), i t i s l i k e l y that that the n u t r i t i o n a l pay-offs for migration w i l l vary widely between populations. In areas where summers are hot and dry on the winter ranges, large differences in forage quality may exist between 96 low and high elevation s i t e s . In more mesic and cooler situations such as the BRV, the n u t r i t i o n a l benefits of migration could be reduced or non-existent. Although Geist (1982) considered elk in the Rockies to be migratory and to follow melt lines of snow, more recent evidence has shown that a substantial portion of several populations, p a r t i c u l a r l y cows, may neither migrate nor follow snow melt lines i f they do migrate (Morgantini 1988, Woods and Hlady 1988, Chapter 2) . As Morgantini (1988) pointed out, migration as such may not be a species c h a r a c t e r i s t i c of elk, as has been implied by many authors, but rather a " v e r s a t i l e " response to a given environmental si t u a t i o n . This i s consistent with Geist's (1982) "opportunism" theory of elk behavior and the observation that no one variable (e.g. diet quality, previous experience, harassment by predators) i s l i k e l y to explain the range of foraging and movement behaviors seen in the species. 97 CHAPTER 6. CHANGES IN THE ELK POPULATION I n t r o d u c t i o n I n t h e s t u d y o f w i l d l i f e p o p u l a t i o n s , a n y management a c t i o n c a n be t r e a t e d as an e x p e r i m e n t a l p e r t u r b a t i o n . M a c n a b (1983) d i s c u s s e d how t h e i n f r e q u e n t a p p l i c a t i o n o f t h i s a p p r o a c h h a s i m p e d e d p r o g r e s s i n s e p a r a t i n g e d u c a t e d g u e s s e s f r o m t e s t e d h y p o t h e s e s . The e x p e r i m e n t a l a p p r o a c h r e q u i r e s t h a t t h e a p p r o p r i a t e d a t a be c o l l e c t e d s y s t e m a t i c a l l y , w h i c h i s d i f f i c u l t a n d e x p e n s i v e f o r a l l l a r g e mammals . H o w e v e r , as B u r k (1973) o u t l i n e d i n h i s r e v i e w o f p r e d a t o r - p r e y l e s s o n s f r o m t h e K a i b a b d e e r h e r d , t h e a b s e n c e o f e x p e r i m e n t a l r i g o r c a n l e a d t o t h e a c c e p t a n c e o f i d e a s n o t s u p p o r t e d by f a c t s . The BRV e l k p o p u l a t i o n o f B a n f f N a t i o n a l P a r k h a s b e e n o f management i n t e r e s t f o r more t h a n a c e n t u r y . A l t h o u g h t h e p a r k was e s t a b l i s h e d i n 1885, e l k d i s a p p e a r e d f r o m t h e BRV e a r l y i n t h e 1900' s . The c a u s e o f t h e d i s a p p e a r a n c e was n o t i d e n t i f i e d . C a n d i d a t e e x p l a n a t i o n s i n c l u d e d d i s e a s e , s e v e r e w i n t e r s , a n d o v e r - h a r v e s t ( B a n f i e l d 1958, H o l r o y d a n d V a n T i g h e m 1983) . D u r i n g 1915-20, n a t i v e e l k r e a p p e a r e d i n t h e p a r k a r e a a n d a d d i t i o n a l e l k were t r a n s p l a n t e d i n t o t h e BRV f r o m Y e l l o w s t o n e N a t i o n a l P a r k , Wyoming ( L l o y d 1927, B a n f i e l d 1958, H o l r o y d a n d V a n T i g h e m 1983). By t h e 1 9 4 0 ' s , t h e BRV was c o n s i d e r e d t o be " o v e r - s t o c k e d " a n d a n e l k p o p u l a t i o n c o n t r o l p r o g r a m i n s t i t u t e d , w h i c h 98 l a s t e d u n t i l 1969 ( H o l r o y d a n d V a n T i g h e m 1983). The C a n a d i a n P a r k s S e r v i c e ( C P S ) , i n c o n j u n c t i o n w i t h t h e C a n a d i a n W i l d l i f e S e r v i c e , s t u d i e d t h e e l k p o p u l a t i o n t h r o u g h o u t t h e c u l l p e r i o d (Cowan 1950, G r e e n 1950, G r e e n 1957, B a n f i e l d 1958, F l o o k 1967, F l o o k a n d S t e n t o n 1969, F l o o k 1970, H o l r o y d a n d V a n T i g h e m 1983). I n t h e 1970's, management a t t e n t i o n s h i f t e d t o t h e n u m b e r s o f e l k k i l l e d i n c o l l i s i o n s w i t h a u t o m o b i l e s a n d t r a i n s w i t h i n t h e B R V . H i g h w a y t r a f f i c a n d e l k r o a d - k i l l s i n c r e a s e d d r a m a t i c a l l y t h r o u g h o u t t h e 1970's a n d t h e r e was c o n c e r n t h a t t h e a d d i t i v e e f f e c t s o f m a n - c a u s e d e l k m o r t a l i t i e s w o u l d r e s u l t i n t h e c o l l a p s e o f t h e p o p u l a t i o n ( H o l r o y d a n d V a n T i g h e m 1983). I n a d d i t i o n , a c c i d e n t s c a u s e d by e l k - v e h i c l e c o l l i s i o n s p o s e d a human s a f e t y p r o b l e m . F o r t h e s e r e a s o n s , an u n g u l a t e - p r o o f f e n c e was c o n s t r u c t e d a l o n g an e x p a n d e d p o r t i o n o f t h e h i g h w a y b e t w e e n 1983-87 ( K l e n a v i c 1979, P a r a d i n e 1982, Woods 1990). D u r i n g t h e p e r i o d 1983-89, C P S , i n c o n j u n c t i o n w i t h P u b l i c W o r k s C a n a d a , s t u d i e d t h e e l k p o p u l a t i o n a n d t h e i m p a c t o f t h e f e n c e , a n d c o n c l u d e d t h a t t h e f e n c e h a d s u c c e s s f u l l y r e d u c e d t h e number o f e l k r o a d - k i l l s (Woods 1990). The h i s t o r y o f t h e BRV e l k p o p u l a t i o n s u g g e s t s s e v e r a l i m p l i c i t management h y p o t h e s e s , i n c l u d i n g : 1) t h a t t h e t r a n s p l a n t was n e c e s s a r y i n o r d e r t o r e - e s t a b l i s h t h e p o p u l a t i o n (1917-20); 2) t h a t t h e r a n g e was " o v e r - s t o c k e d " (1940-69); a n d , 3) t h a t p a r t i a l 99 h i g h w a y f e n c i n g w o u l d p r e v e n t t h e c o l l a p s e o f t h e e l k p o p u l a t i o n a n d a l l o w i t t o i n c r e a s e (1983-87). D e s p i t e t h e h i s t o r i c a n d c o n t i n u i n g i n t e r e s t i n BRV e l k , d a t a on p o p u l a t i o n s i z e a r e s c a n t . T h i s i s o f i n t e r e s t b o t h f o r t h e management o f e l k i n t h e p a r k , a n d f o r t h e a n a l y s i s o f d e n s i t y - d e p e n d e n c e . I n t h i s s t u d y , I m e a s u r e d s e v e r a l p o p u l a t i o n c h a r a c t e r i s t i c s ( e . g . s i z e , c o n d i t i o n , p a r a s i t e s ) a n d c o m p a r e d them w i t h s i m i l a r d a t a g a t h e r e d d u r i n g t h e p o p u l a t i o n c o n t r o l p e r i o d (1940-69). I a d d r e s s t h e g e n e r a l n u l l h y p o t h e s i s t h a t t h e s e c h a r a c t e r i s t i c s h a v e n o t c h a n g e d . 100 M e t h o d s H i s t o r i c a l d a t a The d a t a f i l e s m a i n t a i n e d b y t h e p a r k w a r d e n s e r v i c e i n B a n f f were r e v i e w e d a n d s u m m a r i z e d b y y e a r . U n l e s s o t h e r w i s e n o t e d , d a t a f o r t h e p e r i o d 1944-54 a r e f r o m H . U . G r e e n ( G r e e n 1949, 1957) a n d d a t a f o r t h e p e r i o d 1958-66 f r o m F l o o k (1967, 1970). P o p u l a t i o n e s t i m a t e s I u s e d 3 m e t h o d s t o e s t i m a t e t h e e l k p o p u l a t i o n : d i r e c t g r o u n d c o u n t s , m a r k - r e c a p t u r e e s t i m a t e s , a n d a e r i a l c o u n t s . G r o u n d c o u n t s were c o n d u c t e d e a c h a u t u m n ( l a t e O c t o b e r - e a r l y N o v e m b e r ) a n d s p r i n g ( l a t e A p r i l - e a r l y M a y ) . A u t u m n c o u n t s c o i n c i d e d w i t h p o s t - r u t h e r d f o r m a t i o n a n d s p r i n g c o u n t s w i t h t h e f i r s t v i s i b l e " g r e e n - u p " o f g r a s s e s i n t h e B R V . A e r i a l c o u n t s a n d m a r k - r e c a p t u r e e s t i m a t e s were made a t t h e same t i m e a s t h e s p r i n g g r o u n d c o u n t s . T h i s r e f e r e n c e t i m e was c h o s e n b e c a u s e e l k were more v i s i b l e t h a n a t a n y o t h e r t i m e o f t h e y e a r a n d b e c a u s e i t p r e c e d e s r e c r u i t m e n t i n t o t h e p o p u l a t i o n ( l a t e M a y - e a r l y J u n e ) ( R . K u n e l i u s , C P S , p e r s . c o m m u . ) . G r o u n d c o u n t s were made f r o m r o a d w a y s ( e s t i m a t e d l e n g t h 200 km) d u r i n g t h e e a r l y m o r n i n g a n d l a t e a f t e r n o o n . To m i n i m i z e d u p l i c a t i o n c a u s e d b y e l k m o v e m e n t , a d j a c e n t a r e a s were s u r v e y e d e i t h e r a t t h e same t i m e (2 t e a m s ) o r as s h o r t l y (<1 d a y ) a f t e r one a n o t h e r a s p o s s i b l e . G r o u n d c o u n t s a p p r o x i m a t e d t h e s u r v e y 101 t e c h n i q u e u s e d i n e l k c o u n t s b e f o r e 1985 e x c e p t t h a t p r e v i o u s BRV c o u n t s were c o n f i n e d t o t h e p a r k a n d o c c a s i o n a l l y w e r e c o n d u c t e d as e a r l y as l a t e S e p t e m b e r . D u r i n g t h e g r o u n d c o u n t s , r a d i o t r a n s m i t t e r s w e r e n o t u s e d t o l o c a t e e l k , b u t were n o t e d i n t h e h e r d s s u r v e y e d . A f t e r t h e g r o u n d c o u n t was c o m p l e t e d , a l l r a d i o - t a g g e d e l k were l o c a t e d ( r a d i o - t a g g i n g m e t h o d s d e s c r i b e d i n C h a p t e r 2). U s i n g t h e number known t o be w i t h i n t h e BRV u s e d as t h e m a r k e d s a m p l e a n d t h e number s e e n d u r i n g t h e c o u n t as t h e " r e c a p t u r e " s a m p l e , a Chapman e s t i m a t e was c a l c u l a t e d u s i n g t h e m e t h o d s p r e s e n t e d b y P o l l o c k e t a l.(1990). A e r i a l c o u n t s w e r e made b y t h e p a r k w a r d e n s e r v i c e u s i n g a s t a n d a r d i z e d m e t h o d o l o g y a n d e x p e r i e n c e d o b s e r v e r s ( R . K u n e l i u s , C P S , p e r s . c o m m . ) . E a c h y e a r f r o m 1985-90, t h e BRV was s u r v e y e d by h e l i c o p t e r f o l l o w i n g a p r e d e t e r m i n e d f l i g h t p a t h w h i c h c o v e r e d t h e e n t i r e BRV f r o m Canmore t o L a k e L o u i s e . The f l i g h t s were s c h e d u l e d t o c o m p l e t e t h e c o u n t s on t h r e e c o n s e c u t i v e m o r n i n g s , c o v e r i n g a p p r o x i m a t e l y o n e - t h i r d o f t h e a r e a p e r f l i g h t . The r e s u l t i n g c o u n t g a v e a minimum number o f e l k w i t h i n t h e B R V . I n 1988, t h e a e r i a l c o u n t was r e p e a t e d 3 t i m e s . R a d i o t r a n s m i t t e r s were n o t u s e d t o l o c a t e e l k d u r i n g t h e s e f l i g h t s . B e c a u s e c o l l a r s c o u l d n o t be r e a d i l y o b s e r v e d f r o m t h e a i r , Chapman i n d i c e s o f p o p u l a t i o n s i z e a n d a e r i a l v i s i b i l i t y b i a s e s b a s e d on c o l l a r e d e l k w e r e n o t d e t e r m i n e d . 102 Samuel e t a l . ( 1 9 8 7 ) found t h a t p e r c e n t v e g e t a t i o n cover and group s i z e were p r i m a r y f a c t o r s i n f l u e n c i n g o b s e r v a b i l i t y i n a e r i a l e l k c o u n t s . Group s i z e b i a s i n a e r i a l and ground counts was m i n i m i z e d i n s p r i n g counts by s t a n d a r d i z i n g the count date a t the s t a r t o f the green-up. Group s i z e s e s t i m a t e d from 5 s p r i n g ground counts averaged 11.98 (S.E.=1.03) and group s i z e s e s t i m a t e d from 6 autumn ground counts averaged 14.45 (S.E.=1 .35) . P e r c e n t v e g e t a t i o n cover was a c o n s t a n t . C o n c u r r e n t ground counts and a e r i a l counts d u r i n g 5 s p r i n g s (1986-90) a l l o w e d an e s t i m a t e of the g r o u n d - a e r i a l b i a s . The assumption t h a t s i m i l a r b i a s e s occur i n the autumn a l l o w e d the autumn ground counts f o r a l l y e a r s (1944-90) t o be c o r r e c t e d and d i r e c t l y compared. I d i d not use h i s t o r i c p o p u l a t i o n e s t i m a t e s and c u l l d a t a t o c a l c u l a t e an i n d e x - m a n i p u l a t i o n - i n d e x e s t i m a t e (Caughley 1977) because the i n t e r v a l between r e p e a t e d i n d i c e s (6-12 months) v i o l a t e d the assumption of a c l o s e d p o p u l a t i o n . R e c r u i t m e n t The pregnancy r a t e was e s t i m a t e d by d e t e r m i n i n g i n u t e r o p r e g n a n c i e s of 45 s a l v a g e d road- or r a i l - k i l l e d e l k d u r i n g the p e r i o d November-May. C l a s s i f i e d counts were made each autumn and s p r i n g i n c o n j u n c t i o n w i t h ground c o u n t s . E l k were observed e i t h e r a t v e r y c l o s e range (<100 m) through b i n o c u l a r s , or t h r ough a 20-25x s p o t t i n g scope. A n i m a l s were 103 c l a s s i f i e d i n t o 4 c a t e g o r i e s : c a l f ( e i t h e r sex < l - y e a r - o l d ) , a d u l t cow ( 1 - y e a r - o l d a n d o l d e r ) , s p i k e b u l l ( 1 - y e a r - o l d ) , a n d a d u l t b u l l ( 2 - y e a r s - o l d a n d o l d e r ) . C a l f , s p i k e b u l l , a n d a d u l t b u l l r e s u l t s w e r e e x p r e s s e d a s a r a t i o p e r 100 a d u l t c o w s . C o n f i d e n c e i n t e r v a l s w e r e e s t a b l i s h e d u s i n g t h e f o r m u l a p r e s e n t e d b y S i n c l a i r a n d G r i m s d e l l (1983) w i t h e l k h e r d s as s u b - s a m p l e s . S u r v i v a l o f c o h o r t s o f y o u n g b u l l e l k f r o m c o n c e p t i o n t o 2 - y e a r - o l d s was e s t i m a t e d f r o m t h e i n u t e r o p r e g n a n c y r a t e a n d t h e s u c c e s s i v e c h a n g e s i n r a t i o s o b s e r v e d i n s e q u e n t i a l c l a s s i f i e d c o u n t s . T h i s e s t i m a t e a s s u m e d a 1:1 sex r a t i o in u t e r o a n d n e g l i g i b l e a d u l t cow m o r t a l i t y . M o r t a l i t y M o s t e l k r o a d - k i l l s were l o c a t e d by members o f t h e p a r k w a r d e n s e r v i c e a n d c o o p e r a t i n g a g e n c i e s . A n i m a l s h i t on r o a d w a y s b u t d y i n g o u t o f s i g h t were l o c a t e d d u r i n g o t h e r w o r k , o r by m o r t a l i t y mode s i g n a l s f r o m r a d i o - t a g g e d e l k ( C h a p t e r 2). I t h i n k t h a t m o s t r o a d - k i l l s w e r e f o u n d b e c a u s e w a r d e n p a t r o l s w e r e f r e q u e n t a n d b e c a u s e 4 o f 5 r o a d - k i l l e d r a d i o - t a g g e d e l k w e r e f o u n d b y p a r k w a r d e n s w i t h o u t t h e u s e o f r a d i o s i g n a l s . E l k k i l l e d i n c o l l i s i o n s w i t h t r a i n s w e r e much more d i f f i c u l t t o l o c a t e b e c a u s e much o f t h e r a i l - l i n e was n o t o b s e r v a b l e f r o m p a r k r o a d s . R a i l - k i l l s w e r e l o c a t e d b y f o l l o w i n g up i n c i d e n t a l r e p o r t s , a n d b y 104 b i m o n t h l y h e l i c o p t e r f l i g h t s a l o n g t h e e n t i r e r a i l - l i n e f r o m M a r c h 1987 t o M a r c h 1989. A c o n c u r r e n t s t u d y o f r a d i o - t a g g e d w o l v e s s t a r t e d i n F e b r u a r y 1987 a n d p r o v i d e d i n f o r m a t i o n on e l k k i l l e d by w o l v e s . E l k d e a t h s were c l a s s i f i e d as h i g h w a y o r r a i l w a y m o r t a l i t i e s i f e v i d e n c e was f o u n d i n d i c a t i n g a c o l l i s i o n , a n d w o l f k i l l s i f t h e r e was e v i d e n c e o f s t r u g g l e . O t h e r c a t e g o r i e s ( e . g . p a r a s i t e s , h u n t i n g ) were d e t e r m i n e d f r o m e i t h e r i n c i d e n t a l r e p o r t s o r n e c r o p s i e s . C a l f a n d 1 - y e a r - o l d e l k were a g e d b y t o o t h r e p l a c e m e n t p a t t e r n s ( L a r s o n a n d T a b e r 1980). O l d e r e l k w e r e a g e d f r o m cementum a n a l y s i s o f e i t h e r t h e l o w e r c e n t r a l i n c i s o r o r t h e u p p e r c a n i n e by M a t s o n ' s L a b o r a t o r y , M i l l t o w n , M o n t a n a . May 1st was t h e a s s u m e d b i r t h d a y f o r a l l e l k . R e l o c a t i n g r a d i o - t a g s b r o a d c a s t i n g on m o r t a l i t y mode a l l o w e d a n i n d e p e n d e n t e s t i m a t e o f m o r t a l i t y . S u r v i v a l r a t e s w e r e c a l c u l a t e d f r o m t h e number o f d e a t h s i n t h e r a d i o - t a g g e d s a m p l e u s i n g t h e K a p l a n - M e i e r p r o c e d u r e a s m o d i f i e d f o r s t a g g e r e d e n t r y a n d c e n s o r e d d a t a ( P o l l o c k e t a l . 1989). D i f f e r e n c e s i n s u r v i v a l b e t w e e n y e a r s a n d b e t w e e n s e x e s were c o m p a r e d u s i n g t h e l o g - r a n k t e s t a s d e s c r i b e d by P o l l o c k e t a l . ( 1 9 9 0 ) . A n i m a l c o n d i t i o n i n d i c e s D a t a on m o r p h o l o g y a n d p a r a s i t e l o a d s w e r e g a t h e r e d f r o m s a l v a g e d c a r c a s s e s o f e l k k i l l e d i n c o l l i s i o n s w i t h v e h i c l e s on r o a d s a n d t h e r a i l - l i n e . The l u n g s a n d l i v e r 105 w e r e f r o z e n a n d s e n t t o A l b e r t a F i s h a n d W i l d l i f e i n E d m o n t o n , A l b e r t a . , where t h e y were e x a m i n e d b y M . P y b u s f o r t h e p r e s e n c e o f g i a n t l i v e r f l u k e ( F a s c i o l o i d e s  m a g n a ) , l u n g w o r m s ( D i c t y o c a u l u s v i v i p a r u s ) a n d h y d a t i d c y s t s ( E c h i n o c o c c u s g r a n u l o u s ) . L i v e r s were t h i n l y s e c t i o n e d i n e n t i r e t y a n d t h e m a j o r l u n g p a s s a g e w a y s c u t o p e n a n d e x a m i n e d v i s u a l l y . The p r e v a l e n c e o f p a r a s i t e s was e x p r e s s e d as a p e r c e n t a g e o f a n i m a l s w i t h a p a r t i c u l a r p a r a s i t e w i t h i n a s p e c i f i c age c l a s s . B o d y w e i g h t s w e r e o b t a i n e d f r o m i n t a c t c a r c a s s e s w e i g h e d w h o l e on a s p r i n g b a l a n c e l o c a t e d i n a c e n t r a l a b a t t o i r . B u l l w e i g h t s i n c l u d e d a n t l e r s . A n t l e r w e i g h t s w e r e o b t a i n e d b y s a w i n g one s i d e o f a n an i n t a c t a n t l e r f r o m t h e s k u l l j u s t b e l o w t h e b u r r a n d d r y i n g t h e a n t l e r a t room t e m p e r a t u r e f o r s i x m o n t h s ( C l u t t o n - B r o c k e t a l . 1982) . I n a t r i a l c o m p a r i n g f r e s h t o d r y a n t l e r s , t h e a v e r a g e w e i g h t l o s s was 7% (N=21). The t w o - s i d e d , f r e s h a n t l e r w e i g h t s p r e s e n t e d b y F l o o k (1967) w e r e r e d u c e d b y 7% a n d d i v i d e d i n h a l f t o p r o v i d e a c o m p a r a b l e d r y , s i n g l e - s i d e d a n t l e r w e i g h t . 106 R e s u l t s P o p u l a t i o n e s t i m a t e s D u r i n g 1985-90, a e r i a l c o u n t s i n s p r i n g r a n g e d f r o m 795-955 e l k a n d were w i t h i n c o n t e m p o r a n e o u s m a r k - r e c a p t u r e c o n f i d e n c e i n t e r v a l s ( T a b l e 1 5 ) . G r o u n d c o u n t s i n s p r i n g were c o n s i s t e n t l y s m a l l e r t h a n a e r i a l c o u n t s a n d n o t w i t h i n t h e m a r k - r e c a p t u r e c o n f i d e n c e i n t e r v a l s ( T a b l e 15). R e s u l t s f r o m a l l 3 c e n s u s m e t h o d s s u g g e s t e d l i t t l e c h a n g e i n e l k n u m b e r s f r o m 1985-90. A u t u m n c o u n t s c o r r e c t e d f o r a e r i a l - g r o u n d b i a s w e r e h i g h e r a n d more c o n s i s t e n t f o r 1985-90 t h a n d u r i n g 1944-53 o r 1959-68 ( T a b l e 16, F i g u r e 12, A p p e n d i x I ) . D u r i n g t h e s e e a r l i e r p e r i o d s , autumn c e n s u s d a t a s u g g e s t e d u n l i k e l y y e a r l y i n c r e a s e s d u r i n g 3 i n t e r v a l s (57%, 1952-53; 80%, 1962-63; a n d 180%, 1967-68). D u r i n g 1944-48, s p r i n g c o u n t s c o m b i n e d w i t h known n u m b e r s o f e l k c u l l e d d u r i n g t h e w i n t e r , c o n s i s t e n t l y e x c e e d e d t h e p r e c e d i n g autumn c o u n t ( A p p e n d i x I ) . D u r i n g t h e 18 y e a r s w i t h p o p u l a t i o n c o n t r o l p r o g r a m s a n d a u t u m n p o p u l a t i o n e s t i m a t e s , t h e a v e r a g e c u l l was 21.7% o f t h e c o r r e c t e d autumn c o u n t ( r a n g e 0.2-55.8%)( F i g u r e 12, A p p e n d i x I ) . R e c r u i t m e n t A l l 29 p r e g n a n t e l k e x a m i n e d i n t h i s s t u d y c a r r i e d a s i n g l e f e t u s . One c a s e o f t w i n s was r e c o r d e d d u r i n g t h e 1944-54 e l k c u l l s (N-571). The p r e g n a n c y r a t e d u r i n g 1985-89 was 67% (N=45). 107 T h i s was s i g n i f i c a n t l y l o w e r ( C h i - s q u a r e=7.449, P=0.006, d . f . = l ) t h a n t h e 1958-67 r a t e o f 83% (N=736) b u t n o t s i g n i f i c a n t l y d i f f e r e n t ( C h i - s q u a r e=0.103, P=0.748, d . f . = l ) f r o m t h e 1944-54 r a t e o f 69% (N=823). F l o o k (1970) r e p o r t e d t h a t 21% o f 1 - y e a r - o l d cows (N=82) w e r e p r e g n a n t d u r i n g 1958-67. T h e r e were no p r e g n a n t 1 - y e a r - o l d s i n t h i s s t u d y (N=4). The n u m b e r s o f c a l v e s p e r 100 a d u l t cows i n a u t u m n d u r i n g 1985-90 were h i g h e r a n d more u n i f o r m (42-57) t h a n t h o s e o b s e r v e d d u r i n g 1944-54 (13-45) ( T a b l e 1 7 ) . The s i n g l e r a t i o r e p o r t e d f o r 1958-66 (55) was w i t h i n t h e r a n g e o f r e c e n t c o u n t s . The h i g h e s t r a t i o r e p o r t e d f o r t h e p o p u l a t i o n was 63 (1976) a n d t h e l o w e s t 18 b a s e d o n p o o l e d o b s e r v a t i o n s 1975-80 ( H o l r o y d a n d V a n T i g h e m 1983) . C l a s s i f i e d c o u n t s showed a s i g n i f i c a n t r e d u c t i o n i n t h e n u m b e r s o f c a l f e l k b e t w e e n autumn a n d s p r i n g c o u n t s d u r i n g 1985-90 ( T a b l e 1 7 ) . A s s u m i n g a 1:1 r a t i o o f b u l l t o cow c a l v e s , t h e r e was a f u r t h e r s i g n i f i c a n t r e d u c t i o n i n t h e r e l a t i v e number o f b u l l s t o cows b e t w e e n t h e i r 2nd a n d 3rd w i n t e r s ( T a b l e 1 8 ) . The e f f e c t i v e r e c r u i t m e n t o f b u l l e l k e n t e r i n g t h e i r 3rd y e a r was 8 p e r 100 a d u l t cows ( r a n g e 7-12 ) . The sex r a t i o s o f c a l f a n d 1 - y e a r - o l d e l k k i l l e d i n r o a d a n d r a i l c o l l i s i o n s were n o t s i g n i f i c a n t l y d i f f e r e n t f r o m 1:1 ( T a b l e 18). H o w e v e r , t h e b u l l / c o w r a t i o f o r age c l a s s e s o l d e r t h a n 1 - y e a r - o l d s was s i g n i f i c a n t l y l o w e r 108 (45). T h e s e r e s u l t s p a r a l l e l t h o s e o b s e r v e d d u r i n g 1944-54 a n d 1957-64 ( T a b l e 18). S u r v i v a l D u r i n g t h e p e r i o d F e b r u a r y 1986 t o A p r i l 1989, 300 d e a t h s o f a d u l t e l k were r e p o r t e d , i n c l u d i n g 17 r a d i o - t a g g e d e l k ( T a b l e 19). S i g n i f i c a n t l y more a d u l t e l k w e r e c l a s s i f i e d a s n a t u r a l d e a t h s i n t h e r a d i o - t a g g e d s a m p l e t h a n i n t h e t o t a l s a m p l e . O f t h e r a d i o - t a g g e d s a m p l e , r o a d - k i l l s w e r e t h e l e a d i n g c a u s e o f d e a t h (N=5), f o l l o w e d b y w o l f - k i l l s (N=3), u n i d e n t i f i e d n a t u r a l c a u s e s (N=3), r a i l - k i l l s (N=2), p a r a s i t e s (N=2), h u n t i n g ( N = l ) , a n d h u m a n - r e l a t e d a c c i d e n t s ( N = l ) . A l t h o u g h a v e r a g e a d u l t cow s u r v i v a l (0.90) e x c e e d e d b u l l s u r v i v a l (0.83) d u r i n g 1986-89, t h i s d i f f e r e n c e was n o t s i g n i f i c a n t ( T a b l e 20). C u m u l a t i v e s u r v i v a l o f y o u n g e l k f r o m c o n c e p t i o n t h r o u g h 23 m o n t h s a v e r a g e d 0.23 ( T a b l e 21). The c a l f : c o w r a t i o o f h i g h w a y k i l l s was s i g n i f i c a n t l y h i g h e r (106:100) t h a n t h e p r e g n a n c y r a t e (67:100). The c a l f : c o w r a t i o i n r a i l - k i l l s (27:100) was s l i g h t l y l a r g e r t h a n t h e l a t e w i n t e r c l a s s i f i e d c o u n t s f o r t h e p o p u l a t i o n (21:100). I n a s a m p l e o f 53 w o l f k i l l s , t h e r a t i o o f c a l v e s p e r 100 a d u l t cows (154) was s i g n i f i c a n t l y g r e a t e r t h a n t h e e l k p r e g n a n c y r a t e , a n d t h e r a t i o o f b u l l e l k p e r 100 a d u l t cows (227) i n w o l f k i l l s was s i g n i f i c a n t l y g r e a t e r t h a n t h e a s s u m e d 1:1 r a t i o _in u t e r o . 109 D u r i n g b o t h 1957-64 a n d 1985-90, t h e o l d e s t cows ( 2 1 - a n d 2 2 - y e a r - o l d s r e s p e c t i v e l y ) o u t l i v e d t h e o l d e s t b u l l s (14- a n d 1 3 - y e a r - o l d s r e s p e c t i v e l y ) . I n t h i s s t u d y , t h e a v e r a g e age f o r e l k o l d e r t h a n c a l v e s k i l l e d on t h e h i g h w a y a n d r a i l w a y was 5.8 y e a r s (N=130) f o r cows a n d 3.4 y e a r s (N=82) f o r b u l l s . A l t h o u g h i t was n o t p o s s i b l e t o c a l c u l a t e a v e r a g e a g e s f r o m d a t a r e p o r t e d b y F l o o k (1967, 1970), 4 - y e a r - o l d a n d o l d e r cow e l k c o m p r i s e d 57.3% (N=131) o f t h e a d u l t cows i n t h i s s t u d y a n d 55.7% (N=531) o f t h e 1957-64 s a m p l e . F o u r - y e a r - o l d a n d o l d e r b u l l e l k c o m p r i s e d 73.8% (N=42) o f a l l b u l l s o l d e r t h a n 1 - y e a r - o l d s i n t h i s s t u d y , a n d 66.7% (N=225) d u r i n g 1957-64. P a r a s i t e s G i a n t l i v e r f l u k e s , l u n g w o r m s , a n d h y d a t i d c y s t s i n BRV w e r e f o u n d i n t h e BRV e l k p o p u l a t i o n ( T a b l e 2 2 ) . B o t h g i a n t l i v e r f l u k e s a n d h y d a t i d c y s t s h a d p e a k p r e v a l e n c e i n o l d e r age c l a s s e l k ( > 2 - y e a r - o l d s ) , w h e r e a s l u n g w o r m p r e v a l e n c e was h i g h e s t i n age c l a s s 1 - y e a r - o l d s . L i v e r f l u k e s were f i r s t r e p o r t e d i n BRV e l k i n 1959 ( F l o o k 1967). B e t w e e n 1959-89 t h e y i n c r e a s e d f r o m 0-89% p r e v a l e n c e i n e l k > 2 - y e a r - o l d s ( T a b l e 2 2 ) . The p r e v a l e n c e o f l i v e r f l u k e s i n e l k > 2 - y e a r - o l d s d u r i n g 1985-89 was s i g n i f i c a n t l y g r e a t e r t h a n r e p o r t e d f o r t h i s age c l a s s d u r i n g 1959-65 ( T a b l e 2 2 ) . The o n l y e l k d e a t h s a t t r i b u t e d t o p a r a s i t e s were t h e 2 a d u l t r a d i o - t a g g e d e l k f o u n d d e a d d u r i n g my s t u d y . T h e y h a d m a s s i v e l i v e r damage r e s u l t i n g 110 f r o m g i a n t l i v e r f l u k e i n f e s t a t i o n s . A l t h o u g h b o t h h y d a t i d c y s t s a n d l u n g w o r m s a r e known f r o m t h i s p o p u l a t i o n s i n c e t h e 1 9 4 0 ' s , t h e p r e v a l e n c e s I o b s e r v e d i n > 2 - y e a r - o l d e l k were s i g n i f i c a n t l y g r e a t e r t h a n i n e a r l i e r y e a r s ( T a b l e 22). B o d y a n d a n t l e r w e i g h t s A g e s p e c i f i c b o d y w e i g h t s were o b t a i n e d f r o m 129 i n t a c t r o a d a n d r a i l ^ k i l l e d e l k ( T a b l e 2 3 ) . B u l l s > 2 - y e a r - o l d s were s i g n i f i c a n t l y h e a v i e r (mean=339 k g , N=22, range=230-471 ) t h a n cows i n t h e same age c l a s s (mean=238 k g , N=29, r a n g e=168-311 ) . C o n f i d e n c e i n t e r v a l s f o r mean w e i g h t s o f b u l l a n d cow e l k 3- t o 1 0 - y e a r s o l d d u r i n g 1985-89 o v e r l a p p e d w i t h t h e c o r r e s p o n d i n g d a t a f r o m B a n f f c u l l s i n 1960-63 w i t h e x c e p t i o n o f 6 - y e a r - o l d s . D r i e d a n t l e r w e i g h t s were o b t a i n e d f r o m 68 r o a d - a n d r a i l - k i l l e d e l k ( T a b l e 2 4 ) . A n t l e r s f r o m 4 - y e a r - o l d e l k w e r e s i g n i f i c a n t l y h e a v i e r t h o s e t h a n f r o m 1- t o 3 - y e a r - o l d s a n d s i g n i f i c a n t l y l i g h t e r t h a n a n t l e r s f r o m 8- t o 9 - y e a r - o l d e l k . C o n f i d e n c e i n t e r v a l s f o r 1960-63 o v e r l a p p e d 1985-89 c o n f i d e n c e i n t e r v a l s i n a l l c a s e s . I l l Table 15. E a r l y s p r i n g e lk p o p u l a t i o n e s t imates based on ground counts (GC) , a e r i a l counts ( A C ) , and m a r k - r e c a p t u r e (MR), BRV, 1985-90 Year GC A C a MRb 95% C I b 1984-85 n . a . 795 n . a . n . a 1985-86 488 955 926 652-1200 1986-87 487 891 951 673-1229 1987-88 440 942 = 860 620-1100 1988-89 531 860 979 694-1264 1989-90 523 894 1222 776-1668 a e a r l y s p r i n g t o t a l count from h e l i c o p t e r b e a r l y s p r i n g Chapman es t imate and conf idence i n t e r v a l s based on r a d i o - t a g g e d a d u l t e l k ( P o l l o c k e t a l . 1990) c h i g h e s t of three counts (933, 939, 942) T a b l e 16. Autumn e l k p o p u l a t i o n e s t imates based on ground counts c o r r e c t e d for g r o u n d - a e r i a l b i a s d u r i n g 3 p e r i o d s : 1944-53, 1959-68, and 1985-90. P e r i o d N(years ) Average C o r r e c t e d 8 Count S . E . 1944-53 10 648 66 1959-68 10 725 92 1985-90 6 978b 54 8 g r o u n d - a e r i a l b i a s 0 . 5 5 based on 5 p a i r e d s p r i n g c o u n t s , 1985 -89 b s i g n i f i c a n t l y g r e a t e r than 1944 -53 average ( t - t e s t , P=0 .004) 112 T a b l e 17. C l a s s i f i e d e l k c o u n t r a t i o s a n d 95% c o n f i d e n c e i n t e r v a l s ( C l ) o f c a l f ( C ) , 1 - y e a r - o l d b u l l ( Y B ) , a n d m a t u r e b u l l (MB) e l k i n t h e BRV e x p r e s s e d p e r 100 a d u l t c o w s , 1943-90. Y e a r S e a s o n C l a s s r a t i o s ( C l ) C YB MB N a 1990- 91 A u t u m n 47 .1 (6 .6) 11 .5 (2 .9) 22 .5 (13 .4) 442 1989- 90 A u t u m n 43 .1 (3 .3) 13 .1 (4 .8) 18 .3 (10 .3) 506 S p r i n g 28 .2 (4 .2) 7 .9 (1 .3) 17 .6 (3 .8) 523 1988- 89 A u t u m n 42 .2 (2 .4) 9 .1 (1 .9) 14 .8 (3 .3) 618 S p r i n g 24 .5 (4 • 5) 7 .7 (2 .3) 7 .9 (3 .9) 531 1987- 88 A u t u m n 44 .4 (2 .6) 7 .4 (2 .2) 20 .1 (4 • 8) 624 S p r i n g 21 .8 (3 .6) 7 .4 (2 .0) 11 .9 (5 .0) 440 1986- 87 A u t u m n 42 .8 (4 .0) 13 .6 (3 .5) 24 .0 (8 .5) 489 S p r i n g 29 .8 (4 .2) 7 .0 (1 .6) 17 .4 (5 .2) 487 1985- 86 A u t u m n 57 .4 (2 .1) 13 .6 (1 .5) 35 .1 (5 .0) 546 S p r i n g 23 .8 (3 .0) 11 .9 (2 .0) 17 .2 (5 .3) 488 1976- 77 A u t u m n b 63 .0 n . a . n . a . n . a . 1963- 64 A u t u m n c 54 .6 12 .4 28 .5 567 1953- 54 A u t u m n d 28 .8 12 .5 16 .3 126 1951- 52 A u t u m n d 35 .1 5 .2 20 .1 215 1950- •51 A u t u m n d 13 .0 10 .2 26 .9 162 1949- •50 A u t u m n d 44 .9 5 .1 25 .0 273 1948- 49 A u t u m n d 13 .3 5 .1 13 .9 496 1947- •48 A u t u m n d 36 .8 3 .2 12 .9 523 1946- •47 A u t u m n d 14 .5 4 .8 15 .8 387 1945- •46 A u t u m n d 34 .6 3 .7 18 .9 341 S p r i n g 8 15 n . a . n . a . n . a . 1944- •45 A u t u m n d 39 .9 5 .5 21 .6 456 S p r i n g 6 21 n . a . n . a . n . a . 1943- •44 S p r i n g 6 20 n . a . n . a . n . a . a s a m p l e s i z e b H o l r o y d a n d V a n T i g h e m (1983) c F l o o k (1967) d G r e e n (1957) e Cowan (1950) 113 T a b l e 1 8 . B u l l / c o w r a t i o s o f BRV e l k d u r i n g 1 9 8 5 - 8 9 a , 1 9 5 7 - 6 4 " , a n d 1 9 4 4 - 5 4 = . A g e T i m e P e r i o d  1 9 4 4 - 5 4 ( N ) 1 9 5 7 - 6 4 ( N ) 1 9 8 5 - 8 9 ( N ) C a l f 1 . 1 0 ( 1 9 7 ) 1 . 2 5 ( 1 4 6 ) 0 . 7 7 ( 1 0 6 ) 1 - y e a r 1 . 1 5 ( 1 5 1 ) 1 . 0 4 ( 1 4 5 ) 1 . 5 7 ( 5 9 ) 2 - y e a r n . a . n . a . 0 . 4 7 d ( 1 3 4 ) 0 . 3 5 d ( 2 3 ) > l - y e a r 0 . 2 2 d ( 9 2 8 ) 0 . 4 9 d ' e ( 6 8 5 ) 0 . 4 5 d ' f ( 1 5 4 ) a p r e s e n t s t u d y , r a i l - a n d r o a d - k i l l s b c a l c u l a t e d f r o m F l o o k ( 1 9 6 7 ) , c u l l e d e l k = c a l c u l a t e d f r o m G r e e n ( 1 9 5 7 ) , c u l l e d e l k d s i g n i f i c a n t l y d i f f e r e n t f r o m 1 : 1 e o l d e s t b u l l 14 y e a r s , o l d e s t cow 21 y e a r s f o l d e s t b u l l 1 3 y e a r s , o l d e s t cow 2 2 y e a r s 1 1 4 T a b l e 19. D e a t h s 8 o f a d u l t e l k r e p o r t e d f r o m a l l s o u r c e s a n d f r o m r a d i o - t a g g e d a n i m a l s o n l y , F e b r u a r y 1986 - A p r i l 1989. C a u s e A l l s o u r c e s R a d i o - t e l e m e t r y o n l y Number % Number % M a n - r e l a t e d 212 80 8a 47 b N a t u r a l 53 20 9 53 b T o t a l 265 100 17 100 8 e x c l u d e s d e a t h s o f unknown c a u s e (N=19) a n d o f u n c e r t a i n age (N=16) b s i g n i f i c a n t l y d i f f e r e n t f r o m d e a t h s r e p o r t e d f r o m a l l s o u r c e s 115 T a b l e 2 0 . A n n u a l s u r v i v a l r a t e s (S) a n d 95% c o n f i d e n c e i n t e r v a l s ( C I ) f o r r a d i o - t a g g e d a d u l t e l k i n t h e B R V , 1 9 8 6 - 8 9 . C l a s s N S C I a Y e a r A d u l t cow 1 9 8 6 - 87 34 0 .86 0 . 7 5 - 0 . 9 8 1 9 8 7 - 88 33 0 .94 0 . 8 6 - 1 . 0 2 1 9 8 8 - 89 31 0 .90 0 . 8 0 - 1 . 0 1 A v e r a g e 33 0 . 9 0 0 . 8 0 - 1 . 0 0 A d u l t b u l l 1 9 8 6 - 87 16 0 .74 0 . 5 2 - 0 . 9 7 1 9 8 7 - 88 14 0 . 8 5 0 . 6 6 - 1 . 0 3 1 9 8 8 - 89 12 0 .91 0 . 7 4 - 1 . 0 8 A v e r a g e 14 0 . 8 3 0 . 6 4 - 1 . 0 3 P o o l e d cow a n d b u l l 1 9 8 6 - 87 50 0 . 8 3 0 . 7 2 - 0 . 9 3 1 9 8 7 - 88 47 0 .91 0 . 8 3 - 0 . 9 9 1 9 8 8 - 89 43 0 .91 0 . 8 2 - 0 . 9 9 A v e r a g e 47 0 .88 0 . 7 9 - 0 . 9 7 c a l c u l a t e d f o l l o w i n g P o l l o c k e t a l . (1989) 116 Table 21. Cumulative s u r v i v a l e s t i m a t e s from b i r t h t o age 24 months f o r 6 c o h o r t s of b u l l e l k based on c l a s s i f i e d c o u n t s , 1985-90°. Year Age (months) 0-6 7-12 13-18 19-24 1985-86 0.86 0.36 0.41 0.21 1986-87 0.64 0.45 0.22 0.22 1987-88 0.66 0.33 0.27 0.23 1988-89 0.63 0.37 0.39 0.24 1989-90 0.65 0.42 0.34 -1990-91 0.71 - - -a based on d a t a p r e s e n t e d i n Table 17 117 T a b l e 22. P r e v a l e n c e o f g i a n t l i v e r f l u k e s , l u n g w o r m s , a n d h y d a t i d c y s t s i n BRV e l k d u r i n g 3 p e r i o d s : 1 9 4 4 - 5 4 ° , 1958-66b, a n d 1985-89 c. P a r a s i t e P r e v a l e n c e % by age c l a s s ( s a m p l e s i z e ) 00 01 02 >02 L i v e r f l u k e s 1985-89 3 ( 69) 57 ( 30) 69 ( 16) 89d( 70) 1958-66 0 (138) 6 (126) 13 (117) 13 (462) 1944-54 0 (197) 0 (147) n . a . 0 (906) H y d a t i d c y s t s 1985-89 2 (51) 0 ( 24) 0 ( 12) 31e( 52) 1958-66 0 (154) 4 (142) 18 (130) 24 (523) 1944-54 0 (197) 4 (147) n . a . 9 (906) Lungworms 1985-89 8 ( 51) 44 ( 25) 25 ( 12) 10f( 52) 1958-66 1 (204) 6 (192) 5 (181) 2 (517) 1944-54 15 ( 83) 8 ( 72) n . a . 2 (228) a c a l c u l a t e d f r o m G r e e n (1957) b c a l c u l a t e d f r o m d a t a p r e s e n t e d by F l o o k (1967); l i v e r f l u k e s a m p l e p e r i o d 1959-65; l u n g w o r m d a t a f r o m t h e B R V , K o o t e n a y N a t i o n a l P a r k , a n d t h e Y a - H a T i n d a R a n c h , A l b e r t a c s o u r c e p r e s e n t s t u d y , r o a d - a n d r a i l - k i l l e d e l k d s i g n i f i c a n t l y d i f f e r e n t f r o m a n d 1944-54 a n d 1958-66 p r e v a l e n c e (2x2 C o n t i n g e n c y t a b l e P<0.001) e s i g n i f i c a n t l y d i f f e r e n t f r o m 1944-54 p r e v a l e n c e (P<0.001) b u t n o t t h e 1958-66 p r e v a l e n c e f s i g n i f i c a n t l y d i f f e r e n t f r o m 1944-54 a n d 1958-66 p r e v a l e n c e (P=0.003 a n d P=0.030 r e s p e c t i v e l y ) 118 T a b l e 23. Mean b o d y w e i g h t s (BW) (kg) a n d 95% c o n f i d e n c e i n t e r v a l s ( C I ) f o r r a i l - k i l l e d a n d r o a d - k i l l e d e l k , B R V , 1985-89. A g e B u l l Cows N BW CI N BW CI 0 23 105 88-122 23 94 80-1088 1 16 170 150-190a 5 147 118-1768 2 5 221 185-257 6 186 159-2138 3 7 280 250-310 4 221 188-254 4 0 — — 5 234 197-271 5 3 351 305-397 5 225 195-255 6 4 403 363-443a 4 255 222-288 7 5 351 315-387 3 234 196-272 8 2 341 285-397 3 209 171-247 9 1 401 - 2 265 218-312 10 0 — — 3 285 247-323 a C I ' s do n o t o v e r l a p w i t h c o r r e s p o n d i n g C I ' s p r e s e n t e d b y F l o o k (1970) 119 T a b l e 24. A n t l e r w e i g h t s (kg) a n d 95% c o n f i d e n c e i n t e r v a l s ( C I ) f o r r a i l - k i l l e d a n d r o a d - k i l l e d e l k , B R V , 1985-89. A g e N A v e r a g e W e i g h t CI 1 14 0.20 0.00-0.48 2 2 0.61 0.00-1.35 3 12 1.01 0.71-1.31 4 3 2.28 1.68-2.88 5 7 2.66 2.27-3.05 6 8 2.65 2.28-3.04 7 8 2.98 2.61-3.35 8 7 3.41 3.01-3.81 9 5 3.60 3.13-4.07 10 2 3.19 2.45-3.93 120 1250 F i g u r e 12. A u t u m n e l k p o p u l a t i o n e s t i m a t e s b a s e d o n g r o u n d c o u n t s c o r r e c t e d f o r g r o u n d - a i r b i a s a n d p e r c e n t o f t h i s e s t i m a t e s u b s e q u e n t l y c u l l e d d u r i n g t h e a u t u m n o r e a r l y w i n t e r , B V E R , 1944-89. 121 D i s c u s s i o n P o p u l a t i o n e s t i m a t e s A c c u r a t e e s t i m a t e s o f p o p u l a t i o n s i z e o r t r e n d a r e r e q u i r e d t o t e s t d e n s i t y h y p o t h e s e s f o r BRV e l k . The autumn g r o u n d c o u n t s e m p l o y e d d u r i n g m o s t o f t h e c u l l i n g p e r i o d (1944-69), c o n s i s t e n t l y u n d e r e s t i m a t e d t h e p o p u l a t i o n a n d i n c l u d e d b i o l o g i c a l l y u n l i k e l y i n c r e a s e s ( e . g . 180%, 1967-68). Some o f t h e e s t i m a t e s o f p o s t c e n s u s a n n u a l c u l l r a t e s were a l s o i m p o s s i b l y l a r g e (>100%). D u r i n g y e a r s when b o t h autumn a n d s p r i n g c o u n t s w e r e a v a i l a b l e , f e w e r e l k were s e e n i n t h e autumn t h a n c a n be a c c o u n t e d f o r b y a d d i n g t h e w i n t e r c u l l s t o t h e c o u n t s t h e f o l l o w i n g s p r i n g . F u r t h e r , s i m u l t a n e o u s g r o u n d a n d a e r i a l c o u n t s i n t h i s s t u d y i l l u s t r a t e d t h a t g r o u n d c o u n t s c o n s i s t e n t l y u n d e r e s t i m a t e d t h e p o p u l a t i o n . R e c e n t r a d i o - t e l e m e t r y d a t a f o r BRV e l k ( C h a p t e r 2) c o n f i r m e a r l i e r o b s e r v a t i o n s t h a t o n l y p a r t o f t h e w i n t e r i n g p o p u l a t i o n i s i n t h e BRV d u r i n g t h e p e r i o d o f t h e l a t e autumn c e n s u s e s ( C P S , u n p u b l . d a t a , F l o o k 1970). A f t e r 5 y e a r s o f i n t e n s i v e c u l l i n g (1944-48), e l k d e l a y e d t h e i r r e t u r n t o t h e v a l l e y . T h i s may h a v e c o n t r i b u t e d t h e l a r g e (45%) d r o p i n autumn c o u n t s b e t w e e n 1947-48 ( F i g u r e 12). The e n h a n c e d o b s e r v a b i l i t y o f e l k d u r i n g t h e e a r l y s p r i n g g r e e n - u p a n d t h e c o n c e n t r a t i o n o f e l k w i t h i n t h e BRV a t t h a t t i m e l i k e l y e x p l a i n t h e p a r a d o x o f e l k 122 n u m b e r s i n c r e a s i n g o v e r t h e w i n t e r , a n d show t h a t e a r l y s p r i n g i s t h e b e s t c e n s u s t i m e . The g e n e r a l a g r e e m e n t b e t w e e n m a r k - r e c a p t u r e e s t i m a t e s a n d a e r i a l c o u n t s , a n d t h e c l o s e a g r e e m e n t o f t h e r e p l i c a t e d a e r i a l c o u n t s i n 1987, s u g g e s t t h a t t h e more r e c e n t a e r i a l c o u n t p r o c e d u r e s e m p l o y e d b y t h e p a r k w a r d e n s e r v i c e h a v e p r o d u c e d more c o n s i s t e n t e l k p o p u l a t i o n d a t a . H o w e v e r , c o n f i d e n c e i n t e r v a l s c a n n o t be a p p l i e d t o t h e s e d i r e c t c o u n t s / a n d l i k e g r o u n d c o u n t s , t h e y r e p r e s e n t a minimum p o p u l a t i o n e s t i m a t e . I c o n c l u d e t h a t t h e autumn i s an u n r e l i a b l e c e n s u s t i m e f o r BRV e l k a n d t h u s , t h a t t h e a p p a r e n t i n c r e a s e i n t h e e l k p o p u l a t i o n f r o m 1944-53 t o 1985-90 i s s u s p e c t . Sex r a t i o s a n d l o n g e v i t y F l o o k (1970) c o n s i d e r e d t h e a p p a r e n t d i s p a r i t y i n e l k sex r a t i o s i n BRV (1957-64) a n d i n o t h e r e l k p o p u l a t i o n s i n t h e C a n a d i a n R o c k i e s , a n d n o t e d an a b r u p t c h a n g e i n t h e sex r a t i o b e t w e e n 1 - y e a r - o l d s a n d 2 - y e a r - o l d s . D a t a p r e s e n t e d i n t h i s p a p e r f o r t h e p e r i o d s 1944-54 a n d 1985-90, a l s o showed t h i s r a t i o c h a n g e a t t h e same age i n t e r v a l . F u r t h e r m o r e , d u r i n g b o t h 1957-64 a n d 1985-90, t h e o l d e s t e l k s a m p l e d w e r e c o w s . I n an a n a l y s i s o f p o p u l a t i o n d y n a m i c s o f t h e J a c k s o n H o l e e l k h e r d , B o y c e (1989) d e m o n s t r a t e d c o w - b i a s e d s e x r a t i o s a n d g r e a t e r l o n g e v i t y a m o n g s t c o w s . T h e s e o b s e r v a t i o n s were c o n s i s t e n t w i t h a h i g h e r 123 o v e r - w i n t e r s u r v i v a l o f a d u l t cows i n t h a t p o p u l a t i o n . F l o o k (1970) s u g g e s t e d t h a t u n e q u a l d i s p e r s a l , u n e q u a l s u r v i v a l , o r b o t h c o u l d a c c o u n t f o r t h e u n b a l a n c e d s e x r a t i o s o b s e r v e d i n t h e B R V . A l t h o u g h t h e p r e s e n t s t u d y d i d n o t d e m o n s t r a t e s i g n i f i c a n t d i f f e r e n c e s i n s u r v i v a l o f a d u l t b u l l a n d a d u l t cow e l k , t h e s u r v i v a l e s t i m a t e s h a d w i d e c o n f i d e n c e i n t e r v a l s . I n a d d i t i o n , t h e r a d i o - t a g g e d s a m p l e c o n s i s t e d p r i m a r i l y o f a d u l t e l k a n d t h e r e f o r e w o u l d n o t h a v e d e t e c t e d d i f f e r e n c e s i n s u r v i v a l b e t w e e n t h e a g e s o f 1 a n d 2 y e a r s . S i n c e more a d u l t b u l l s m i g r a t e t h a n a d u l t cows i n t h e BRV p o p u l a t i o n , a n d s i n c e t h e sex r a t i o d i s p a r i t y d o e s n o t a p p e a r u n t i l a f t e r age 2, d e c r e a s e d s u r v i v a l a s s o c i a t e d w i t h m i g r a t i o n o r d i s p e r s a l p r o v i d e s a r e a s o n a b l e , b u t u n t e s t e d e x p l a n a t i o n o f t h e d a t a . B o d y a n d a n t l e r w e i g h t s C a u g h l e y (1979) n o t e d t h a t a l t h o u g h p o p u l a t i o n c o n d i t i o n p a r a m e t e r s s u c h as b o d y s i z e a r e n o t d i r e c t l y r e l a t e d t o " o v e r - p o p u l a t i o n " , t h e y w o u l d be e x p e c t e d be c h a n g e w i t h p o p u l a t i o n d e n s i t i e s . M c C o r q u o d a l e e t a l . (1989) f o u n d t h a t a n t l e r w e i g h t s f r o m a c o l o n i z i n g ( l o w d e n s i t y ) e l k p o p u l a t i o n i n W a s h i n g t o n S t a t e w e r e s i g n i f i c a n t l y h e a v i e r t h a n t h o s e o b s e r v e d e i t h e r b y F l o o k (1970) o r i n t h e p r e s e n t s t u d y i n t h e B R V . W h i l e t h i s b e t w e e n - p o p u l a t i o n c o m p a r i s o n i s c o n s i s t e n t w i t h t h e d e n s i t y - d e p e n d e n c e h y p o t h e s i s t h a t a n t l e r s i z e d e c r e a s e s w i t h i n c r e a s i n g e l k d e n s i t y , more c o n v i n c i n g e v i d e n c e w o u l d come f r o m 1 p o p u l a t i o n o b s e r v e d a t d i f f e r e n t d e n s i t i e s . T h i s may be p o s s i b l e i n t h e f u t u r e i f BRV e l k n u m b e r s i n c r e a s e . P a r a s i t e s The s i g n i f i c a n t i n c r e a s e i n g i a n t l i v e r f l u k e p r e v a l e n c e was t h e o n l y c l e a r c h a n g e i n t h e e l k p o p u l a t i o n c o n d i t i o n b e t w e e n t h e s t a r t o f t h e e l k p o p u l a t i o n c o n t r o l p r o g r a m i n t h e 1940's a n d t h e p r e s e n t s t u d y . S i n c e s p e c i f i c e f f o r t s were made f i n d p a r a s i t e s t h r o u g h o u t t h e c u l l p r o g r a m ( G r e e n 1957, F l o o k 1967, 1970), a n d g i a n t l i v e r f l u k e s a r e c o n s p i c u o u s p a r a s i t e s , I b e l i e v e t h a t t h i s i n c r e a s e i s r e a l . H y d a t i d c y s t s a n d l u n g w o r m s a r e more d i f f i c u l t t o d e t e c t , a n d f o r t h e s e s p e c i e s I c o n s i d e r t h e s t a t i s t i c a l l y s i g n i f i c a n t i n c r e a s e s s u s p e c t . P y b u s (1990) d i s c u s s e d t h e r e c e n t d i s t r i b u t i o n o f t h e g i a n t l i v e r f l u k e i n A l b e r t a , a n d n o t e d t h a t i t h a d a h i g h e r p r e v a l e n c e (89%) i n e l k i n B a n f f N a t i o n a l P a r k ( A l b e r t a ) a n d a d j a c e n t K o o t e n a y N a t i o n a l P a r k ( B r i t i s h C o l u m b i a ) t h a n i n A l b e r t a g e n e r a l l y (29%). T h i s a b u n d a n c e p a t t e r n may be r e l a t e d t o t h e o r i g i n a l d i s p e r s a l p a t t e r n o f t h e p a r a s i t e f r o m B r i t i s h C o l u m b i a ( P y b u s 1990). The two p a r a s i t e - r e l a t e d m o r t a l i t i e s o b s e r v e d i n t h i s s t u d y were t h e f i r s t d e a t h s o f f r e e - r a n g i n g e l k a t t r i b u t e d t o g i a n t l i v e r f l u k e s i n t h e p a r k a n d 125 p o s s i b l y t h e f i r s t d e a t h s o f f r e e - r a n g i n g e l k a t t r i b u t e d t o g i a n t l i v e r f l u k e s a n y w h e r e ( M . P y b u s , p e r s . c o m m . ) . H o w e v e r , t h e s e d e a t h s (2/17) may u n d e r - r e p r e s e n t t h e t r u e m o r t a l i t y f r o m t h e s e p a r a s i t e s b e c a u s e i n t a c t l i v e r s f r o m n a t u r a l d e a t h s a r e r a r e l y a v a i l a b l e f o r n e c r o p s y . R e c r u i t m e n t a n d s u r v i v a l I n a r e c e n t s t u d y o f in . u t e r o p r e g n a n c i e s a m o n g s t e l k o n r a n g e s i m m e d i a t e l y n o r t h o f t h e B R V , t h e r a t e f o r a l l e l k o l d e r t h a n c a l v e s was 0.66 (N=417) ( M o r g a n t i n i 1988). T h i s r a t e was w i t h i n t h e v a l u e s o b s e r v e d f o r BRV e l k d u r i n g t h e p a s t 50 y e a r s a n d w i t h i n v a l u e s o b s e r v e d f o r many N o r t h A m e r i c a n e l k p o p u l a t i o n s ( T a b e r e t a l . 1982). No s i m p l e c o n c l u s i o n c a n be d r a w n f r o m BRV p r e g n a n c y r a t e s o t h e r t h a n t h e y a p p e a r t o h a v e i n c r e a s e d a n d more r e c e n t l y d e c l i n e d a g a i n . The p e r c e n t o f o l d e r cows ( > 3 - y e a r - o l d s ) was s i m i l a r i n t h e 1960's a n d i n t h i s s t u d y . H o w e v e r , t h e l a r g e number (21%) o f b r e e d i n g 1 - y e a r - o l d s d u r i n g t h e 1958-67 s u g g e s t s t h a t t h e h i g h p r e g n a n c y r a t e may h a v e b e e n t h e c o n s e q u e n c e o f more b r e e d i n g p a r t i c i p a t i o n by y o u n g e r age c l a s s e s . R e l i a b l e c a l f : c o w e s t i m a t e s t a k e n o v e r a v a r i e t y o f p o p u l a t i o n d e n s i t i e s h a v e b e e n u s e d t o d e m o n s t r a t e d e n s i t y - d e p e n d e n t r e c r u i t m e n t i n e l k p o p u l a t i o n s ( H o u s t o n 1982, B o y c e 1989). A p p r o p r i a t e d a t a f o r t h i s a n a l y s i s i n t h e BRV were l a c k i n g . 126 B o b e k e t al . (1983) u s e d c l a s s i f i e d c o u n t d a t a f r o m t h e BRV (1944-70) t o i l l u s t r a t e h i g h l y v a r i a b l e c a l f t c o w r a t i o s , an i n v e r s e r e l a t i o n s h i p b e t w e e n s n o w f a l l a n d c a l f p r o d u c t i o n , a n d an i n v e r s e r e l a t i o n s h i p b e t w e e n s n o w f a l l a n d b u l l t c o w r a t i o s . H o w e v e r , t h e s e d a t a s h o u l d be u s e d w i t h c a u t i o n b e c a u s e c o n f i d e n c e i n t e r v a l s w e r e n o t d e t e r m i n e d p r i o r t o 1985. D u r i n g t h e p r e s e n t s t u d y , I f o u n d t h a t 1 - y e a r - o l d b u l l s w i t h s l e n d e r s p i k e a n t l e r s c o u l d be e a s i l y m i s t a k e n f o r a d u l t c o w s , a s c o u l d l a r g e b u l l c a l v e s . F o r t h i s r e a s o n , e l k c l a s s i f i c a t i o n s w e r e o n l y made b y p r a c t i c e d o b s e r v e r s w i t h t h e a i d o f a s p o t t i n g s c o p e o r b i n o c u l a r s a t c l o s e r a n g e . A l t h o u g h G r e e n (1957) was an e x p e r i e n c e d o b s e r v e r , d e t a i l s on t h e 1944-54 c l a s s i f i c a t i o n m e t h o d s a r e l a c k i n g a n d t h e h i g h l y v a r i a b l e a n n u a l r e s u l t s a t t h a t t i m e s h o u l d be v i e w e d w i t h some c a u t i o n . F o r t h e same r e a s o n , I l a c k c o n f i d e n c e i n t h e e x t r e m e l y l o w r a t i o (18:100) f o r B a n f f e l k c a l c u l a t e d f r o m p o o l e d i n c i d e n t a l o b s e r v a t i o n s made o v e r t h r e e m o n t h s b y v a r i o u s o b s e r v e r s d u r i n g 1975-80 ( H o l r o y d a n d V a n T i g h e m 1983). D u r i n g t h e same p e r i o d (November 1976), a s i n g l e c l a s s i f i e d c o u n t i n t h e BRV e s t i m a t e d 63 c a l v e s : 1 0 0 c o w s , t h e h i g h e s t r a t i o e v e r r e p o r t e d ( H o l r o y d a n d V a n T i g h e m 1983). P a i r e d a u t u m n / s p r i n g c l a s s i f i e d c o u n t s f o r t h e BRV d u r i n g 1985-90 i l l u s t r a t e d t h e i m p o r t a n c e o f c o u n t t i m i n g . C l a s s i f i e d c o u n t s d e s c r i b e d as " w i n t e r " c o u n t s c o u l d c o n t a i n a g r e a t d e a l o f v a r i a b i l i t y due t o c o u n t 127 d a t e . Low 1st y e a r s u r v i v a l r a t e s h a v e b e e n p r e v i o u s l y d o c u m e n t e d i n t h i s e l k p o p u l a t i o n (Cowan 1950, G r e e n 1957, a n d F l o o k 1967) a n d i n t h e U p p e r Red D e e r R i v e r v a l l e y e l k p o p u l a t i o n ( M o r g a n t i n i 1988). S i m i l a r r e s u l t s w e r e f o u n d i n my s t u d y . Cowan (1950) p r e s e n t e d l o w j u v e n i l e s u r v i v a l as a c h a r a c t e r i s t i c o f t h i s p o p u l a t i o n o n " o v e r - s t o c k e d " r a n g e . S i n c e c u r r e n t j u v e n i l e s u r v i v a l r a t e s a r e s i m i l a r t o t h o s e p r e s e n t e d b y Cowan (1950), a n d s i n c e r a t e s h a v e n o t b e e n o b s e r v e d o v e r a r a n g e o f d e n s i t i e s , I d o u b t t h a t t h e s e s u r v i v a l r a t e s i n d i c a t e a r a n g e a t o r a b o v e " c a r r y i n g c a p a c i t y " . H o l r o y d a n d V a n T i g h e m (1983) r e v i e w e d m o r t a l i t y s o u r c e s f o r B a n f f e l k , s u g g e s t i n g a p r e d o m i n a n c e o f h u m a n - c a u s e d m o r t a l i t i e s . My c o m p a r i s o n o f r e p o r t s f r o m a l l s o u r c e s , w i t h d e a t h s o f r a d i o - t a g g e d e l k , s u g g e s t s t h a t i n c i d e n t a l m o r t a l i t y r e p o r t s a r e h i g h l y b i a s e d . E l k d y i n g on t h e h i g h w a y a n d r a i l w a y a r e much e a s i e r t o f i n d t h a n n a t u r a l d e a t h s . T h e r e h a v e b e e n no p r e v i o u s e s t i m a t e s o f s u r v i v a l r a t e s f o r a d u l t e l k i n t h e B R V . The F u t u r e o f t h e BRV E l k P o p u l a t i o n A l t h o u g h " c a r r y i n g c a p a c i t y " i s a f r e q u e n t l y u s e d t e r m i n w i l d l i f e e c o l o g y , much a m b i g u i t y s u r r o u n d s t h e c o n c e p t . C a u g h l e y (1979) d i s t i n g u i s h e d b e t w e e n " e c o l o g i c a l " c a r r y i n g c a p a c i t y ( K A ) , t h e u n a i d e d e q u i l i b r i u m r e s u l t i n g f r o m t h e i n t e r a c t i o n o f a p o p u l a t i o n a n d i t s e n v i r o n m e n t , a n d a l t e r n a t i v e c a r r y i n g 128 c a p a c i t i e s d e f i n e d b y human o b j e c t i v e s ( e . g . K e , maximum s u s t a i n e d y i e l d ) . I t f o l l o w s t h a t t h e c o n c e p t o f " o v e r - p o p u l a t i o n " ( i . e . e x c e e d i n g c a r r y i n g c a p a c i t y ) m u s t be e x p l i c i t l y d e f i n e d . M a c n a b (1985) n o t e d t h a t a l t h o u g h u n g u l a t e n u m b e r s may n a t u r a l l y f l u c t u a t e a b o u t Kit c h r o n i c a n d e p i s o d i c m o r t a l i t y may be v i e w e d a s " a b a d t h i n g " b y some p e o p l e . F o r some 30 y e a r s , t h e BRV e l k p o p u l a t i o n was c o n s i d e r e d t o be " o v e r - s t o c k e d " o r " e x c e s s i v e " , b a s e d o n t h e o b s e r v e d c o n d i t i o n o f t h e r a n g e , t h e s u s p e c t e d d e p r e s s i o n o f o t h e r u n g u l a t e s p e c i e s , a n d t h e p o t e n t i a l f o r a m a j o r e l k d i e - o f f i n t h e e v e n t o f a h a r s h w i n t e r (Cowan 1950, B a n f i e l d 1958, F l o o k 1970, H o l r o y d a n d V a n T i g h e m 1983). S i n c e r e c e n t p o p u l a t i o n e s t i m a t e s a r e a t o r a b o v e v a l u e s o b s e r v e d d u r i n g t h e p o p u l a t i o n c o n t r o l p e r i o d , a n d s i n c e t h e f e n c e may r e s u l t i n a s t a b l e o r i n c r e a s i n g e l k p o p u l a t i o n , t h e t o p i c o f " o v e r - a b u n d a n c e " i s o f c o n t i n u i n g i n t e r e s t . E l k a n d o t h e r l a r g e u n g u l a t e s a r e f r e q u e n t l y i n v o l v e d i n K A d i s c u s s i o n s , a n d c o n c e r n f o r t h e w e l f a r e o f t h e s e p o p u l a t i o n s h a s r e s u l t e d i n a n t i c i p a t o r y i n t e r v e n t i o n s u c h as c u l l p r o g r a m s . H o u s t o n (1985) o u t l i n e d s u c h a c a s e i n Y e l l o w s t o n e N a t i o n a l P a r k a n d t r a c k e d p o p u l a t i o n c h a r a c t e r i s t i c s a f t e r p o p u l a t i o n c o n t r o l was d i s c o n t i n u e d . O b s e r v a t i o n s o f r e c r u i t m e n t r a t e s a t c o m p a r a t i v e l y l o w a n d h i g h d e n s i t i e s , i l l u s t r a t e d d e n s i t y - d e p e n d e n t r e c r u i t m e n t f o r c a l f a n d 129 1 - y e a r - o l d b u l l s i n t h e Y e l l o w s t o n e p o p u l a t i o n a f t e r t h e c u l l i n g s t o p p e d . I n t h e B R V , c e s s a t i o n o f t h e c u l l p r o g r a m was c o i n c i d e n t a l l y f o l l o w e d by a s u b s t a n t i a l i n c r e a s e i n h i g h w a y t r a f f i c . E l k r o a d - k i l l s a l s o i n c r e a s e d a n d t h e p o p u l a t i o n d i d n o t e x p a n d m a r k e d l y . T h e r e f o r e , r e c r u i t m e n t , m i g r a t i o n r a t e s a n d o t h e r p o t e n t i a l l y d e n s i t y - d e p e n d e n t c h a r a c t e r i s t i c s ( B o y c e 1984) h a v e n o t b e e n m e a s u r e d o v e r a w i d e r a n g e o f p o p u l a t i o n s i z e s . I f management i n t e r v e n t i o n s s u c h as h i g h w a y f e n c i n g r e d u c e m o r t a l i t y , e l k d e n s i t i e s may i n c r e a s e . H o w e v e r , i s o l a t i o n o f t h e c a u s e a n d e f f e c t may be c o m p l i c a t e d b y s i m u l t a n e o u s c h a n g e s i n s e v e r a l f a c t o r s s u c h a s m o r t a l i t y f r o m o t h e r s o u r c e s ( r o a d - k i l l s on u n f e n c e d p o r t i o n s o f t h e h i g h w a y , r a i l - k i l l s , p r e d a t i o n , p a r a s i t e s , h u n t i n g ) a n d a c h a n g e i n t h e Kt o f t h e BRV r e s u l t i n g f r o m f o r e s t s u c c e s s i o n , l a n d s c a p e a l i e n a t i o n , o r c l i m a t e c h a n g e . 130 CHAPTER 7. CONCLUSION In this study I set out to understand the nature of seasonal movements in a population of elk. My goals were to describe these movements and to reach some understanding of their causes. I found a complex set of annual movement behaviors including migration and residency (Chapter 2). If these movement behaviors are considered to be alternate strategies, then demonstration of a mixed ESS would require that migrant and resident morphs have equal fit n e s s (Maynard Smith and Price 1973) and that individuals would have fixed strategies. Although I did not measure fi t n e s s , I documented individuals "switching" between resident and migrant status from year-to-year, and a mixture of residents and migrants during the rut. Qualitative observations of this population during former years also i l l u s t r a t e d f l e x i b i l i t y in seasonal movement behavior related to disturbance by humans. These data support the hypothesis (1) that p a r t i a l migration in elk i s a conditional ESS and ate consistent with the conclusions of Morgantini (1988) and Boyce (1991) for elk, Sandegren and Bergstrom (1983) for moose, and McCullough (1985) for long range movements of t e r r e s t r i a l mammals in general. Given the ecological p l a s t i c i t y of elk, and the d i v e r s i t y of documented movement behaviors, i t would be surprising to find a s t r i c t genetic linkage with 131 a n n u a l movement b e h a v i o r . V e r s a t i l i t y w o u l d a l l o w a s p e c i e s t o o p t i m i z e i t s r e s p o n s e t o a g i v e n s e t o f e n v i r o n m e n t a l v a r i a b l e s a n d t o c h a n g e t h i s r e s p o n s e a s c o n d i t i o n s d i c t a t e . N o t o n l y w o u l d e l k p o p u l a t i o n s f r o m e n v i r o n m e n t a l l y d i f f e r e n t a r e a s be e x p e c t e d t o v a r y i n movement p a t t e r n s , b u t w i t h i n an a r e a , movement p a t t e r n s c o u l d v a r y w i t h e n v i r o n m e n t a l f l u c t u a t i o n ( e . g . d e n s i t y , p r e d a t o r d i s t u r b a n c e , snow d e p t h ) . The c o m p l e x i t y o f e l k s e a s o n a l movement b e h a v i o r i s f u r t h e r i l l u s t r a t e d b y t h e p r e p o n d e r a n c e o f m a l e m i g r a n t s w h i c h c o n t r a d i c t s t h e h y p o t h e s i s (2) t h a t a d u l t m a l e s a n d a d u l t f e m a l e s i n t h i s p o p u l a t i o n a r e e q u a l l y l i k e l y t o m i g r a t e . The a s y m m e t r i e s i n m o r p h o l o g y a n d e c o l o g y b e t w e e n t h e s e x e s i n t h i s s p e c i e s ( C l u t t o n - B r o c k e t a l . 1982), s u g g e s t t h a t m a l e a n d f e m a l e e l k e v a l u a t e t h e i r e n v i r o n m e n t d i f f e r e n t l y , a n d t h e r e f o r e m i g h t h a v e d i f f e r e n t m i g r a n t / r e s i d e n t r a t i o s . I n a d d i t i o n , i f j u v e n i l e m a l e s a r e t h e p r e d o m i n a n t d i s p e r s e r s i n e l k ( c u r r e n t l y u n p r o v e n ) , t h e y w o u l d be t h e sex m o s t l i k e l y t o d e v e l o p e x p e r i e n c e w i t h a l t e r n a t e r a n g e s . R e t u r n m i g r a t i o n b e h a v i o r c o u l d d e v e l o p f r o m " e x p l o r a t o r y " d i s p e r s a l s . The h y p o t h e s i s (3) o f e q u a l n u m b e r s o f m i g r a n t s a n d r e s i d e n t s was r e j e c t e d by t h e s i g n i f i c a n t l y g r e a t e r number o f r e s i d e n t s i n t h e BRV p o p u l a t i o n . I n c o n t r a s t , m i g r a n t s o u t n u m b e r r e s i d e n t s i n most o t h e r e l k p o p u l a t i o n s i n t h e R o c k i e s . The BRV p o p u l a t i o n a l s o i s 132 u n u s u a l i n b e i n g l a r g e l y u n h u n t e d . I t i s p o s s i b l e ( b u t u n t e s t e d ) t h a t i f t h e y were h u n t e d , t h e m i g r a n t / r e s i d e n t r a t i o w o u l d c h a n g e i n f a v o r o f m i g r a n t s . S i n c e s e a s o n a l movements o f e l k a r e p o p u l a t i o n - s p e c i f i c , i t i s i m p o r t a n t t o r e c o g n i z e i n t e r - p o p u l a t i o n d i f f e r e n c e s . F o r e x a m p l e , a l t h o u g h t h e BRV a n d Red D e e r R i v e r v a l l e y e l k p o p u l a t i o n s i n B a n f f N a t i o n a l P a r k o c c u p y a d j a c e n t d r a i n a g e s o f t h e C a n a d i a n R o c k i e s , t h e s e p o p u l a t i o n s h a v e d i f f e r e n t m i g r a n t / r e s i d e n t r a t i o s a n d d i s t i n c t m i g r a t i o n p a t t e r n s a n d t i m i n g ( M o r g a n t i n i 1988, t h i s s t u d y C h a p t e r 1) . A s B o y c e (1991) d i s c u s s e d f o r t h e J a c k s o n e l k h e r d , k n o w l e d g e o f t h e s e f a c t o r s c a n p l a y an i m p o r t a n t r o l e i n s p e c i e s m a n a g e m e n t . A l t h o u g h m i g r a t i o n h a s t h e p o t e n t i a l o f u n c o u p l i n g p r e d a t o r - p r e y r e l a t i o n s h i p s ( F r y x e l l a n d S i n c l a i r 1988), t h i s was u n l i k e l y t o be t h e c a s e f o r e l k a n d w o l v e s i n t h e B R V . The h y p o t h e s i s (4) t h a t m i g r a t i o n t a k e s p r e y b e y o n d t h e f o r a g i n g r a n g e o f t h e i r p r e d a t o r s was i n c o n s i s t e n t w i t h t h e o b s e r v a t i o n t h a t m o s t BRV e l k a n d w o l v e s w e r e s y m p a t r i c t h r o u g h o u t t h e y e a r . D e s p i t e i n d i v i d u a l v e r s a t i l i t y i n movement b e h a v i o r w i t h i n BRV e l k , t h e y were n o t n o m a d i c , a n d most a d u l t s showed p h i l o p a t r y t o b o t h t h e i r movement s t r a t e g y a n d t h e i r o c c u p a n c y o f w i n t e r , summer, a n d r u t t i n g r a n g e s ( C h a p t e r 3). A l t h o u g h BRV e l k were p h i l o p a t r i c r e l a t i v e t o a random m o d e l , I c o u l d n o t r e j e c t t h e h y p o t h e s i s (5) 133 t h a t t h e r e l a t i v e p h i l o p a t r y o f cows a n d r e s i d e n t s was e q u a l t o t h a t o f b u l l s a n d m i g r a n t s . M o s t a d u l t s u s e d t h e same a r e a s a t t h e same t i m e s f r o m y e a r - t o - y e a r . E x c e p t i o n s t o t h i s p a t t e r n were shown b y t h e few a n i m a l s t h a t s w i t c h e d m i g r a t i o n s t r a t e g i e s ( a l t h o u g h t h e i r u s e o f w i n t e r r a n g e r e m a i n e d c o n s t a n t ) a n d t h e s i n g l e i n s t a n c e o f a d u l t d i s p e r s a l . D u r i n g t h i s s t u d y e n v i r o n m e n t a l c o n d i t i o n s i n t h e BRV w e r e r e l a t i v e l y c o n s t a n t (no e x t r e m e snow y e a r s , no c u l l p r o g r a m , p r e s e n c e o f t i m b e r w o l v e s ) a n d t h e e l k w e r e p h i l o p a t r i c . H o w e v e r , i n t h e m i d - 1 9 4 0 ' s , t h e s i m u l t a n e o u s i n i t i a t i o n o f an e l k c u l l p r o g r a m a n d t h e r e a p p e a r a n c e o f t i m b e r w o l v e s may h a v e c a u s e d b o t h t h e m i g r a n t / r e s i d e n t r a t i o a n d m i g r a t i o n t i m i n g t o c h a n g e r e l a t i v e t o e a r l i e r y e a r s . T h i s s u g g e s t s t h a t w h i l e e l k a r e g e n e r a l l y p h i l o p a t r i c , t h e y r e m a i n v e r s a t i l e e n o u g h t o a d o p t new movement p a t t e r n s a n d r a n g e s i f f a c e d w i t h m a j o r c h a n g e s i n t h e i r e n v i r o n m e n t ( e . g . i n i t i a t i o n o f a h u n t i n g s e a s o n , a p p e a r a n c e o f a m a j o r p r e d a t o r , e x t r e m e s n o w f a l l ) . T h e r e i s n e e d f o r a d d i t i o n a l r e s e a r c h on m i g r a t i o n a n d d i s p e r s a l o f y o u n g e l k ( f r o m c a l v e s t h r o u g h t o 3 - y e a r - o l d s ) , p a r t i c u l a r l y o f y o u n g f r o m cows o f known movement h i s t o r y . The d r a m a t i c d r o p i n t h e p r o p o r t i o n o f 2 - y e a r - o l d b u l l s f i r s t o b s e r v e d b y F l o o k (1967), c o n t i n u e s t o be shown b y t h i s p o p u l a t i o n ( C h a p t e r 6) . A s F l o o k (1967) s u g g e s t e d , t h i s may be t h e r e s u l t o f 134 d i f f e r e n t i a l d i s p e r s a l , d i f f e r e n t i a l m o r t a l i t y , o r b o t h . C a l c u l a t i o n o f n e t t i m e a n d e n e r g y c o s t s o f m i g r a t i o n b y BRV e l k d i d n o t s u p p o r t t h e h y p o t h e s i s (6) t h a t t h e s e i n v e s t m e n t s o f t i m e a n d e n e r g y a r e s i g n i f i c a n t r e l a t i v e t o o t h e r l i f e h i s t o r y c o s t s ( C h a p t e r 4). T h i s r e s u l t c o u l d s u g g e s t t h a t n e t r e w a r d s r e s u l t i n g f r o m m i g r a t i o n ( e . g . a v o i d a n c e o f p r e d a t i o n , b e t t e r q u a l i t y f o o d ) n e e d n o t be l a r g e t o p r o d u c e n e t b e n e f i t s . The s h o r t - t e r m v i s i t s t o a l t e r n a t e r a n g e s ( e . g . w i n t e r r a n g e s d u r i n g t h e summer) made b y s e v e r a l r a d i o - t a g g e d BRV e l k d u r i n g t h i s s t u d y , c o u l d be i n t e r p r e t e d a s l o w - c o s t " t e s t s " o f e n v i r o n m e n t a l q u a l i t y o n t h e s e a l t e r n a t e r a n g e s . D a t a on f o r a g e q u a l i t y on r e s i d e n t a n d m i g r a t o r y r a n g e s o f BRV e l k d i d n o t d e m o n s t r a t e a c o n s i s t e n t d i f f e r e n c e b e t w e e n r a n g e s . H o w e v e r , I c o u l d n o t r e j e c t t h e h y p o t h e s i s (7) t h a t t h e n u t r i t i o n a l q u a l i t y o f l o w a n d h i g h e l e v a t i o n r a n g e s was e q u a l ( C h a p t e r 5). T h i s s i m i l a r i t y i n t h e f o o d q u a l i t y o f d i f f e r e n t r a n g e s c o r r o b o r a t e s e v i d e n c e f r o m o t h e r a r e a s t h a t h i g h e l e v a t i o n r a n g e s c a n be h i g h , l o w , o r e q u a l i n n u t r i t i o n a l q u a l i t y c o m p a r e d t o l o w e l e v a t i o n r a n g e s u s e d b y t h e same p o p u l a t i o n s . S u c h v a r i a b i l i t y i n f o o d q u a l i t y u n d e r s c o r e s t h e v a l u e o f f l e x i b i l i t y i n s e a s o n a l m o v e m e n t s . I n m o s t r e s p e c t s , t h e h y p o t h e s i s (8) t h a t p o p u l a t i o n c h a r a c t e r i s t i c s h a v e n o t c h a n g e d s i n c e t h e 135 c u l l p r o g r a m , was u p h e l d . A l t h o u g h t h e i n c i d e n c e o f g i a n t l i v e r f l u k e s i n c r e a s e d , i n most o t h e r r e s p e c t s ( e . g . p o p u l a t i o n s i z e , b o d y w e i g h t , a n t l e r w e i g h t , sex r a t i o s , m a j o r p r e d a t o r s ) , t h e e l k p o p u l a t i o n e i t h e r h a s c h a n g e d r e l a t i v e l y l i t t l e , o r t h e d a t a a r e i n a d e q u a t e t o d e m o n s t r a t e a c h a n g e ( C h a p t e r 6 ) . U n f o r t u n a t e l y , m e a s u r e m e n t s o f t h e s e c h a r a c t e r i s t i c s h a v e n o t b e e n made a t d i f f e r e n t p o p u l a t i o n s l e v e l s . F i n a l l y , f u r t h e r work a l s o i s n e e d e d on n a t a l d i s p e r s a l , d e n s i t y - d e p e n d e n c e p r o c e s s e s , a n d c a r r y i n g c a p a c i t y . 136 L I T E R A T U R E C I T E D A . O . A . C . 1965. O f f i c i a l m e t h o d s o f a n a l y s i s . 10th e d . W a s h i n g t o n . A s s o c i a t i o n o f O f f i c i a l A g r i c u l t u r a l C h e m i s t s . A D A M S , A . W. 1982. M i g r a t i o n . I n E l k o f N o r t h A m e r i c a : e c o l o g y a n d m a n a g e m e n t . E d i t e d bj£ J . W. Thomas a n d D . E . T o w e i l l . H a r r i s b u r g . S t a c k p o l e B o o k s , p p . 301-321. A D R I A E N S E N , F . AND A . A . DHONDT. 1990. P o p u l a t i o n d y n a m i c s a n d p a r t i a l m i g r a t i o n o f t h e E u r o p e a n r o b i n ( E r i t h a c u s r u b e c u l a ) i n d i f f e r e n t h a b i t a t s . J . A n i m . E c o l . 59: 1077-1090. A L T M A N N , M . 1956. P a t t e r n s o f h e r d b e h a v i o r i n f r e e - r a n g i n g e l k o f W y o m i n g , C e r v u s c a n a d e n s i s  n e l s o n i i . Z o o l o g i c a 41: 65-71. A R C E S E , P . 1989. I n t r a s e x u a l c o m p e t i t i o n , m a t i n g s y s t e m a n d n a t a l d i s p e r s a l i n s o n g s p a r r o w s . A n i m . 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P e r i o d AUT SPR C U L L R A I L ROAD S o u r c e 1885-1903 + + 0 ? 0 C P S a 1903-14 0 0 0 0 0 CPS 1915-16 +b + 0 ? 0 CPS 1918-19 41+c ? 0 ? 0 L l o y d (1927) 1920-21 194+c ? 0 ? 0 L l o y d (1927) 1923-24 + 282 0 ? 0 L l o y d (1927) 1929-30 + + 0 18 0 H o l r o y d a n d V a n T i g h e m (1983) 1930-31 + + 0 12 0 H o l r o y d a n d V a n T i g h e m (1983) 1931-32 + + 0 10 0 H o l r o y d a n d V a n T i g h e m (1983) 1932-33 + + 0 121 0 H o l r o y d a n d V a n T i g h e m (1983) 1933-34 + + 0 18 0 H o l r o y d a n d V a n T i g h e m (1983) 1940-41 + + + ? ? CPS 1941-42 + + + ? ? CPS 1942-43 + + + ? ? CPS 1943-44 + 570 175d ? ? CPS 1944-45 456 592e 200 50 4 CPS 1945-46 341 535 352 50 4 C P S , F l o o k ( 1970) 1946-47 387 343 309 50 4 C P S , F l o o k ( 1970) 1947-48 523 394 254 50 4 C P S , F l o o k ( 1970) 1948-49 496 + 103f 50 4 C P S , F l o o k ( 1970) 1949-50 273 + 270f 50 4 C P S , F l o o k ( 1970) 1950-51 190 + 143f 50 4 C P S , F l o o k ( 1970) 1951-52 244 + 102f 50 4 C P S , F l o o k ( 1970) 1952-53 255 + 0 50 4 C P S , F l o o k ( 1970) 1953-54 400 + 53 50 4 C P S , F l o o k ( 1970) 1954-55 + + 7 7 ? C P S , F l o o k ( 1970) 1955-56 + + 10 ? ? C P S , F l o o k ( 1970) 1956-57 + + ? 13 ? CPS 1957-58 + + 91 26 ? CPS 1958-59 + + 99 9 ? CPS 148 A p p e n d i x I . ( p a g e 2) A u t u m n (AUT) a n d s p r i n g (SPR) g r o u n d c o u n t s , n u m b e r s r e m o v e d f o r p o p u l a t i o n c o n t r o l ( C U L L ) , n u m b e r s o f r a i l - k i l l s ( R A I L ) , a n d n u m b e r s o f r o a d - k i l l s (ROAD) f o r e l k i n t h e B R V , 1885-1990. P e r i o d AUT SPR C U L L R A I L ROAD S o u r c e 1959-60 669 + 300 ? ? CPS 1960-61 459 + 151 ? ? CPS 1961-62 434 + 199 ? ? CPS 1962-63 340 + 100 7 ? CPS 1963-64 613 + 73 ? ? F l o o k 1964-65 360 + 67 ? ? F l o o k 1965-66 250 + 1 ? ? F l o o k 1966-67 260 + 32 ? ? F l o o k 1967-68 158 + 0 ? ? CPS 1968-69 442 + 9 ? ? CPS 1969-70 + + 132 ? 20 CPS 1970-71 + + 0 3h 12h CPS 1971-72 + + 0 25h 21h CPS 1972-73 + + 0 80h 8h CPS 1973-74 + + 0 6h 7h CPS 1974-75 + 0 10h 1 7 h CPS 1975-76 + + 0 17h 29h CPS 1976-77 372 + 0 4h 26h CPS 1977-78 + + 0 6h 37h CPS 1978-79 + + 0 1 7 h 46h CPS 1979-80 + + 0 13h 59h CPS 1980-81 + + 0 41h 60h CPS 1981-82 + + 0 18h 44h CPS 1982-83 + + 0 l l h 48h CPS 1983-84 + + 0 9 h 48h CPS 1984-85 + + 0 25h 56h CPS 1985-86 546 488 0 30 90 CPS 1986-87 489 487 0 49 88 CPS 1987-88 624 440 0 33 45 CPS 1988-89 618 531 0 26 32 CPS 1989-90 506 523 0 11 + 39 CPS 1990-91 442 559 0 79 53 CPS a C a n a d i a n P a r k s S e r v i c e u n p u b l i s h e d d a t a , W a r d e n O f f i c e , B a n f f , d a t a f o r 1944-54 f r o m G r e e n (1957) b n a t i v e e l k n o t e d i n p a r k s u p e r i n t e n d e n t r e p o r t s c number o f e l k t r a n s p l a n t e d i n BRV f r o m Y e l l o w s t o n e N a t i o n a l P a r k d a g g r e g a t e c u l l e d 1940-43 e w a r d e n e s t i m a t e o f 1500 c i t e d b y B a n f i e l d (1958) f i n c l u d e d BRV a n d C a s c a d e R i v e r v a l l e y * e s t i m a t e p u b l i s h e d i n Bobek e t al . (1982) o f 30 e x c l u d e d a s p r o b a b l e e r r o r h b a s e d o n c a l e n d a r y e a r o f f i r s t y e a r 149 

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