FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page i FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page ii about island press Island Press is the only nonprofit organization in the United States whose principal purpose is the publication of books on environmental issues and natural resource management We provide solutions-oriented information to professionals, public officials, business and community leaders, and concerned citizens who are shaping responses to environmental problems Since 1984, Island Press has been the leading provider of timely and practical books that take a multidisciplinary approach to critical environmental concerns Our growing list of titles reflects our commitment to bringing the best of an expanding body of literature to the environmental community throughout North America and the world Support for Island Press is provided by the Agua Fund, The Geraldine R Dodge Foundation, Doris Duke Charitable Foundation, The Ford Foundation, The William and Flora Hewlett Foundation, The Joyce Foundation, Kendeda Sustainability Fund of the Tides Foundation, The Forrest & Frances Lattner Foundation, The Henry Luce Foundation, The John D and Catherine T MacArthur Foundation, The Marisla Foundation, The Andrew W Mellon Foundation, Gordon and Betty Moore Foundation, The Curtis and Edith Munson Foundation, Oak Foundation, The Overbrook Foundation, The David and Lucile Packard Foundation, Wallace Global Fund, The Winslow Foundation, and other generous donors The opinions expressed in this book are those of the authors and not necessarily reflect the views of these foundations FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page iii Mountain Goats FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page iv FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page v Mountain Goats Ecology, Behavior, and Conservation of an Alpine Ungulate Marco Festa-Bianchet Steeve D Cơté Washington • Covelo • London FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page vi Copyright © 2008 Island Press All rights reserved under International and Pan-American Copyright Conventions No part of this book may be reproduced in any form or by any means without permission in writing from the publisher: Island Press, 1718 Connecticut Avenue NW, Suite 300, Washington, DC 20009, USA Island Press is a trademark of The Center for Resource Economics Library of Congress Cataloging-in-Publication Data Festa-Bianchet, Marco Mountain goats : ecology, behavior, and conservation of an alpine ungulate / Marco Festa-Bianchet, Steeve Côté p cm Includes bibliographical references and index ISBN 978-1-59726-170-8 (hardcover : alk paper) -ISBN 978-1-59726-171-5 (pbk : alk paper) Mountain goat I Côté, Steeve D II Title QL737.U53F48 2007 639.97'96475 dc22 2007025958 Printed on recycled, acid-free paper Manufactured in the United States of America 10 FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page vii À Wendy et Mélanie, qui ont toujours accepté et respecté nos longs séjours sur la montagne E per Alberto, a cui sarebbe piaciuto vedere le capre FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page viii FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page ix Contents Acknowledgments xi Ecological Questions, Conservation Challenges, and Long-Term Research The Study Area and the Goat Population 17 Caw Ridge Study Methods and Limitations 36 Home Ranges, Forage Availability, and Habitat Use 48 Social Organization 71 Body and Horn Growth 91 Individual Variability in Yearly and Lifetime Reproductive Success of Females 118 Female Reproductive Strategy 137 Survival and Dispersal 157 10 Density-Dependence and the Question of Population Regulation 177 11 Female Reproductive Strategy and Ungulate Population Dynamics 191 12 Management and Conservation of Mountain Goats 204 13 Long-Term Monitoring of Marked Individuals and Advances in Ecology and Conservation 219 Appendix 231 Literature Cited 235 Index 257 ix Lit Cited:IP_Festa-Bianchet 9/7/07 1:34 PM Page 253 Literature Cited 253 turity in female moose (Alces alces): the effects of environmental variability Journal of Animal Ecology 62:482–489 Sæther, B E., M Lillegard, V Grøtan, F Filli, and S Engen 2007 Predicting fluctuations of reintroduced ibex populations: the importance of density dependence, environmental stochasticity and uncertain population estimates Journal of Animal Ecology 76:326–336 Sæther, B E., E J Solberg, and M Heim 2003 Effects of altering sex ratio structure on the demography of an isolated moose population Journal of Wildlife Management 67:455– 466 Saltz, D., and B P Kotler 2003 Maternal age is a predominant determinant of progeny sex ratio variation in ungulates: a reply to Hewison et al Oikos 101:646–648 San José, C., S Lovari, and N Ferrari 1997 Grouping in roe deer: an effect of habitat openness or cover distribution? 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Oikos 61:401–411 Smith, B L 1976 Ecology of the Rocky Mountain goat in the Bitterroot Mountains, Montana M.Sc thesis, University of Montana, Missoula ——— 1988a Criteria for determining age and sex of American mountain goats in the field Journal of Mammalogy 69:395–402 Smith, B L., and S H Anderson 1998 Juvenile survival and population regulation of the Jackson elk herd Journal of Wildlife Management 62:1036–1045 Smith, C A 1986 Rates and causes of mortality in mountain goats in southeast Alaska Journal of Wildlife Management 50:743–746 Smith, K G 1988b Factors affecting the population dynamics of mountain goats in westcentral Alberta Proceedings of the Biennial Symposium of the Northern Wild Sheep and Goat Council 6:308–329 ——— 2004 Woodland caribou demography and persistence relative to landscape change in westcentral Alberta M.Sc thesis, University of Alberta, Edmonton Smith, K G., E J Ficht, D Hobson, T C Sorensen, and D Hervieux 2000 Winter distribution of woodland caribou in relation to clear-cut logging in west-central Alberta Canadian Journal of Zoology 78:1433–1440 Sokal, R R., and F J Rohlf 1981 Biometry 2nd ed San Francisco: Freeman Lit Cited:IP_Festa-Bianchet 254 9/7/07 1:34 PM Page 254 Literature Cited Solberg, E J., A Loison, B E Sæther, and O Strand 2000 Age-specific harvest mortality in a Norwegian moose Alces alces population Wildlife Biology 6:41–52 Stearns, S C 1992 The evolution of life histories Oxford: Oxford University Press Steele, B M., and J T Hogg 2003 Measuring individual quality in conservation and behavior 243–270 in Animal behavior and wildlife conservation, ed M Festa-Bianchet, M Apollonio Washington, DC: Island Press Steen, W J v d 1983 Methodological problems in evolutionary biology, I: Testability and tautologies Acta Biotheoretica 32:207–215 Stelfox, J B., 1993 Hoofed mammals of Alberta Edmonton: Lone Pine Publishing Stevens, V 1983 The dynamics of dispersal in an introduced mountain goat population Ph.D thesis, University of Washington, Pullman Stevens, V., and D B Houston 1989 Reliability of age determination of mountain goats Wildlife Society Bulletin 17:72–74 Stockwell, C A., G C Bateman, and J Berger 1991 Conflicts in National Parks: a case study of helicopters and bighorn sheep time budgets at the Grand Canyon Biological Conservation 56:317–328 Swenson, J E 1985 Compensatory reproduction in an introduced mountain goat population in the Absaroka Mountains, Montana Journal of Wildlife Management 49:837–843 Swihart, R K., H P Weeks, A L Easter-Pilcher, and A J DeNicola 1998 Nutritional condition and fertility of white-tailed deer (Odocoileus virginianus) from areas with contrasting histories of hunting Canadian Journal of Zoology 76:1932–1941 Taillon, J., D G Sauvé, and S D Côté 2006 The effects of decreasing winter diet quality on foraging behavior and life-history traits of white-tailed deer fawns Journal of Wildlife Management 70:1445–1454 Tavecchia, G., T N Coulson, B J T Morgan, J M Pemberton, J Pilkington, F M D Gulland, and T H Clutton-Brock 2005 Predictors of reproductive cost in female Soay sheep Journal of Animal Ecology 74:201–213 Terry, E L., B N McLellan, and G S Watts 2000 Winter habitat ecology of mountain caribou in relation to forest management Journal of Applied Ecology 37:589–602 Testa, J W., and G P Adams 1998 Body condition and adjustments to reproductive effort in female moose (Alces alces) Journal of Mammalogy 79:1345–1354 Thomas, C D., A Cameron, R E Green, M Bakkenes, L J Beaumont, Y C Collingham, B F N Erasmus, M F de Siqueira, A Grainger, L Hannah, L Hughes, B Huntley, A S van Jaarsveld, G F Midgley, L Miles, M A Ortega-Huerta, A T Peterson, O L Phillips, and S E Williams 2004 Extinction risk from climate change Nature 427:145–148 Thomas, D C., S J Barry, and H P Kiliaan 1989 Fetal sex ratios in caribou: maternal age and condition effects Journal of Wildlife Management 53:885–890 Thomas, D C., and D R Grey, 2002 Update COSEWIC Status Report on the woodland caribou Rangifer tarandus caribou in Canada In COSEWIC assessment and update Status Report on the woodland caribou Rangifer tarandus caribou in Canada Ottawa: Committee on the Status of Endangered Wildlife in Canada, 98 Thompson, K V 1995 Flehmen and birth synchrony among female sable antelope, Hyppotragus niger Animal Behaviour 50:475–484 Thompson, R W., and J C Turner 1982 Temporal geographic variation in the lambing season of bighorn sheep Canadian Journal of Zoology 60:1781–1793 Toïgo, C., J.-M Gaillard, D Gauthier, I Girard, J P Martinot, and J Michallet 2002 Female reproductive success and costs in an alpine capital breeder under contrasting environments Ecoscience 9:427–433 Toïgo, C., J.-M Gaillard, and J Michallet 1997 Adult survival of the sexually dimorphic Alpine ibex (Capra ibex ibex) Canadian Journal of Zoology 75:75–79 Toïgo, C., and J M Gaillard 2003 Causes of sex-biased adult survival in ungulates: sexual size dimorphism, mating tactic or environment harshness? Oikos 101:376–384 Lit Cited:IP_Festa-Bianchet 9/7/07 1:34 PM Page 255 Literature Cited 255 Toïgo, C., J M Gaillard, and J Michallet 1999 Cohort affects growth of males but not females in alpine ibex (Capra ibex ibex) Journal of Mammalogy 80:1021–1027 Trivers, R L., and D E Willard 1973 Natural selection of parental ability to vary the sex ratio of offspring Science 179:90–92 Van Ballenberghe, V., and W B Ballard 1994 Limitation and regulation of moose populations: the role of predation Canadian Journal of Zoology 72:2071–2077 VanderWerf, E 1992 Lack’s clutch size hypothesis: an examination of the evidence using meta-analysis Ecology 73:1699–1705 Veitch, A., E Simmons, M Promislow, D Tate, M Swallow, and R Popko 2002 The status of mountain goats in Canada’s Northwest Territories Proceedings of the Biennial Symposium of the Northern Wild Sheep and Goat Council 13:49–62 Verme, L J 1985 Progeny sex ratio relationships in deer: theoretical vs observed Journal of Wildlife Management 49:134–136 von Hardenberg, A., B Bassano, M Z Arranz, and G Bogliani 2004 Horn growth but not asymmetry heralds the onset of senescence in Alpine ibex males Journal of Zoology 263:425–432 von Hardenberg, A., B Bassano, A Peracino, and S Lovari 2000 Male alpine chamois occupy territories at hotspots before the mating season Ethology 106:617–630 Wade, M J., and S Kalisz 1990 The causes of natural selection Evolution 44:1947–1955 Wauters, L A., S A d Crombrugghe, N Nour, and E Matthysen 1995 Do female roe deer in good condition produce more sons than daughters? Behavioral Ecology and Sociobiology 37:189–193 Weatherhead, P J 1986 How unusual are unusual events? American Naturalist 128:150–154 Wehausen, J D 1995 Fecal measures of diet quality in wild and domestic ruminants Journal of Wildlife Management 59:816–823 ——— 1996 Effects of mountain lion predation on bighorn sheep in the Sierra Nevada and Granite Mountains of California Wildlife Society Bulletin 24:471–479 White, G C., and R M Bartmann 1998 Effect of density reduction on overwinter survival of free-ranging mule deer fawns Journal of Wildlife Management 62:214–225 White, R G 1983 Foraging patterns and their multiplier effects on productivity of northern ungulates Oikos 40:377–384 Williams, J S 1999 Compensatory reproduction and dispersal in an introduced mountain goat population in central Montana Wildlife Society Bulletin 27:1019–1024 Wittmer, H U., A R E Sinclair, and B N McLellan 2005 The role of predation in the decline and extirpation of woodland caribou Oecologia 114:257–267 Wright, G J., R O Peterson, D W Smith, and T O Lemke 2006 Selection of Northern Yellowstone elk by gray wolves and hunters Journal of Wildlife Management 70:1070– 1078 Yoccoz, N G., A Mysterud, R Langvatn, and N C Stenseth 2002 Age- and density-dependent reproductive effort in male red deer Proceedings of the Royal Society of London B 269:1523–1528 Lit Cited:IP_Festa-Bianchet 9/7/07 1:34 PM Page 256 index:IP_Festa-Bianchet 9/7/07 1:35 PM Page 257 Index Age: dominance and, 71–72; horn annuli and, 100, 100, 171 Aggression: antiparallel fighting style, 99, 99; displays, 99, 99, 105; of females, 72, 76–77, 82, 98, 99; horn use, 98–99, 99, 103; measuring, 76; submissive behavior to, 45, 99; in traps, 38; weighing scales and, 42 Alberta Conservation Association, 226 Alberta Fish and Wildlife Division, 26, 29, 212, 226 All-occurrence sampling, 45 Antelope, sable, 128 Antiparallel fighting style, 99, 99 ArcView GIS, Animal Movement Analyst, 49 Ash content (forage), 58 ATVs, 31, 230 Avens, mountain (Dryas spp.), 18, 46 Bears, black, 24 Bears, grizzly, 24, 64, 65, 68, 127 Behavior of goats: observing (overview), 45 See also specific behavior Birthdate synchronization: breeding cycle synchronization and, 127–128; kid mass and, 129–130; maternal age/condition factors, 128; median birthdate (Caw Ridge), 129; neonatal mortality and, 127, 130–131; overwinter mortality and, 127, 128, 130, 131; plant phenology hypothesis on, 126, 127, 131; predation hypothesis on, 126–127, 127–128, 130–131; statistics on (1993–1999/Caw Ridge), 128–129, 128, 129; in ungulates/examples, 126 Birthing: description, 55–56; gestation time, 56, 57; isolation and, 45, 56– 57, 72, 73; timing factors, 48 Bison, 152 Black Hills, South Dakota, 13 Body condition and reproductive success, 125–126 Body mass: average adult weights, 79; birthweight, 93; female reproductive success and, 120, 125, 126; kid survival and, 162–163, 162; longevity correlation and, 171–172; monitoring, 91, 92; mother/offspring at weaning, 147–148, 147; sexual dimorphism in, 79–81, 91, 94, 95, 106– 107, 108, 113, 115; sexual segregation and, 75, 78–80, 89–90, 89 257 index:IP_Festa-Bianchet 258 9/7/07 1:35 PM Page 258 Index Body mass growth: climate effects, 112–113; dormant forage and, 112– 113; goats vs bighorn sheep, 94, 106–107, 107, 108; kids, 93–94, 93; over spring/summer, 92–93, 93; population density and, 108–113, 110, 111, 112, 115, 116–117; as resource availability indicator, 92; seasonal/age-specific description, 94– 95, 96, 96, 97; statistical notes on, 115–117; summary, 91, 115; yearlings, 94, 95 Body mass loss: aging, 113; winter, 95– 96 Capture–mark–recapture technique, 165 Capturing/handling goats: immobilization description, 38; immobilization effects, 41–42, 47, 119, 141, 142; kid abandonment and, 41 See also Traps Caribou: barren-ground, 154; helicopter harassment of, 33; woodland, 23– 24, 230 Caw Ridge mountain goat study (overview): buildings, 30; conservation and, 3, 5–7, 216–217; field season, 36–37; future research, 228– 229; goals of, 2–7; history, 27, 29; limitations overview, 46–47; need for long-term study, 1–2, 5–6; previous information on goats, 2–3, 15; technique summary, 36, 47; winter and, 46–47 See also Long-term studies/ marked individuals Caw Ridge study area: bighorn sheep study locations, 18; biodiversity, 22; cliffs and, 18, 20; components of long-term studies, 17; description/ overview, 6–7, 7, 17–25, 20, 21; fauna (overview), 22, 23–25, 231– 233; location, 17, 18; mammals/birds list, 231–233; map, 22; protection of, 230; seasons, 19–20, 29; treeline, 18– 19, 23; vegetation (overview), 18; weather, 17–18, 19–21, 19, 46–47 Caw Ridge study area/mountain goats: future of, 229–230; isolation and, 19, 31; recent history, 25–26, 27; threats to goats (overview), 31–35 Censusing goats: description, 42–45; determining sex–age class, 205; helicopter surveys/problems, 205–206, 212, 217; hunters/game wardens and, 206; management and, 205– 206, 207, 208; nursery groups, 74; recommendations on, 206 See also Population size/goats Chamois: age-specific body growth, 167; dispersal, 175; home ranges of, 55; horn growth, 103; mountain goats and, 16; paragliders effects, 34; photo, 10; yearling survival, 163 Chamois, Alpine, 175 Climate change See Global warming Coal exploration history, 25, 31–32 Coal mine pit/area: bighorn sheep and, 22; mountain goats and, 22, 228; photo, 24 Compensatory growth (horns), 99, 103–104, 104, 105, 112 Conservation: habitat protection, 3, 216; habitat-use patterns and, 69, 216; long-term studies and, 2, 227; metapopulation approach to, 217; motorized vehicles and, 31, 217, 218, 230; overview, 216–217; reproduction variability factor and, 118; research and, 2, 227 See also specific threats Côté, Steeve, 19, 20, 29, 38, 46 Cougars, 24, 64, 65, 68, 88, 134–135, 186, 188 Coyotes, 24 Deer, fallow: Phoenix Park study, 227; predator avoidance, 74; reproduction, 92, 119, 126, 152; survival, 171 Deer, mule, 23 Deer, red: hunting effects, 227; population density studies, 109; reproduc- index:IP_Festa-Bianchet 9/7/07 1:35 PM Page 259 Index tion, 119, 128, 144, 149–150, 195, 202; survival, 167 Deer, roe: predator avoidance, 74; reproduction, 154; survival, 167, 168, 171; yearling survival, 163 Deer, white-tailed, 22, 23, 195, 200 Deer social organization, 84 Density dependence/effects: goat studies and, 177, 179–183, 180, 182; lack of (Caw Ridge), 185–186, 188–189, 189; management implications, 178 Density independence/effects: management implications, 178, 189; overview, 177 Dispersal of goats: description, 172– 175; emigration summary table, 164; at Olympic National Park, 172, 173, 185; sex/age differences, 173–174, 175, 176 Domestic goats, Dominance relations: adult females, 71–72, 88, 155; dyadic relationships, 88; hunting effects on, 211–212 Dryas spp (mountain avens), 18, 46 Eagles, golden, 24, 127, 130–131 Elasticity in survival, 214 Elk, 199 Emigration from Caw Ridge (summary table), 164 Energy costs: lactation, 56, 60, 127, 143–144, 145; primiparity timing, 143; reproduction (overview), 139 Escape terrain, 48, 64, 69, 84, 85, 87 Fauna at Caw Ridge, 22, 23–25, 231– 233 Fecal ash, 58 Fecal crude protein: horn growth and, 101; mass gain and, 93–94; measuring forage quality and, 58–61, 59, 60, 62, 70, 93; population density and, 109, 110, 110 Fecal sampling method, 46 Females, adult: aggression of, 72, 76– 77, 82, 98, 99; dominance relation- 259 ships in, 71–72, 88, 155; dyadic relationships, 88; home range examples, 50–51; kin groups and, 84, 85–86, 87; lactation status determination, 40; marking of, 38, 39; survival overview, 166–168, 169–170, 169, 170, 175; urination posture, 43 See also Birthing; Nursery groups; reproduction Females, adult-offspring association, 81; description, 81–84, 82, 83, 84; mother’s age and, 83, 83; offspring gender and, 82, 82; offspring mass gain and, 83, 89–90, 89; offspring survival and, 83–84; signs of, 43; with/without kid, 83, 84 See also Nursery groups Festa-Bianchet, Marco, 27, 29 Fighting style of goats, 99, 99 Fitness: complexity of, 137; measuring, 137 Fitness costs: definition/description, 139; female/male kids, 148–149; individual variability and, 139–140, 146–147; measuring, 137, 139–140; primiparity timing and, 142, 143, 144, 147, 155, 193; reproductive potential and, 140; reproductive strategy/female goats, 143–146 Focal sampling, 45 Foot length/indicator: body condition measurements, 126; population density study and, 110, 111, 111; skeletal development and, 97–98, 97, 113 Forage/foraging: altitudinal migration, 48, 57; ash content, 58; biomass/ seasonal changes, 62–63, 63; crude protein in graminoids/willows, 61, 62; goats vs bighorn sheep, 58, 60– 61, 60; grasses, 58; measuring quality, 58–61, 59, 60, 62, 70, 93; phenolic compounds, 58; seasonal quality changes, 56–62, 59, 60; topography effects on, 48, 57; winter weather arrival and, 57; yearly quality changes, 59–61, 59, 60 index:IP_Festa-Bianchet 9/7/07 1:35 PM 260 Page 260 Index Forestry effects, 24, 229–230 Forest use by males, 5, 47, 64–66, 65, 66, 75 Gas exploration/effects, 24, 25 Gazelle, Cuvier’s, 154 Generalized linear mixed models (GLMMs), 151 Genetic analysis of goats, 40 Gestation time, 56, 57 GIS, ArcView Animal Movement Analyst, 49 GLMMs (generalized linear mixed models), 151 Global warming: alpine habitat and, 2, 229; conservation and, 217, 218; future research on, 229 Goats, domestic goats, Goral, 16 Grande Cache weather, 21 Habitat: group size and, 74, 84, 86; harvesting implications, 88; loss effects, 69; protecting, 3, 216 Hamell, Mount, 19, 25 Hares, snowshoe, 24 Haviernick, Martine, 61 Helicopters: goat reactions to, 32–35, 33; harm from, 7, 26, 33–35, 216, 217, 218; recommendations on, 34– 35; surveys/problems, 205–206, 212, 217 Herd-specificity approach See Metapopulation approach Home ranges: of adult males, 49, 52– 53, 55, 55; comparison to other species, 49, 55; escape terrain and, 69; examples, 50–54; of nursery groups, 49, 53–54, 55; population density and, 109; seasonality of, 49, 52–54, 55 Horn annuli, 100, 100, 102, 103, 104 Horn growth: by age, 102; compensatory growth, 99, 103–104, 104, 105, 112; description, 99, 100, 101–105, 113; fecal crude protein and, 101; goats vs bighorn sheep, 107–108; lactation energy costs and, 99, 105; population density and, 109–110, 111, 111, 112, 115; as resource availability indicator, 92; by sex, 102; statistical notes on, 116; summary, 91, 115 Horns: description, 3, 4, 8, 10, 98; as display organ, 105; functions, 38, 98–99, 99, 103; intraspecific interactions and, 98–99, 99, 103; longevity correlation and, 171–172; long horns, 102, 103, 104–105, 113; measuring, 100–101, 100; sexual dimorphism and, 8, 98, 102, 103, 107, 108, 113, 115; trophy hunting/effects, 114, 211; wearing of tips, 105 Horn threat display, 99 Horses, feral, 150 Human disturbance and goats: compared to other ungulates, 33, 41; conservation and, 216, 217 See also specific disturbances Hunting: decrease in, 202; densitydependent populations and, 178; density-independent populations and, 178; female ungulates reproductive strategy and, 196, 198–203, 201; recommendations on, 202–203; of young, 202–203, 209–211 Hunting ethics, 210 Hunting/goats: accessible populations and, 213–214; closing of, 2, 27, 209, 214; double-quota system, 209; elasticity in survival and, 214; extirpation, 4, 11, 204; female age/kid sex ratio and, 155; female harvesting/effects, 5, 114, 209; goat sensitivity, 2– 3, 4; harvest program examples, 208; harvest rate/introduced goats, 214; harvest rate policies, 207, 208; harvest rate recommendations, 214; herd specificity and, 208, 212–213, 213, 216; identification of sex and, 4– 5, 5, 114, 209; interest in, 204, 205; introduced goats, 3, 214–215; of index:IP_Festa-Bianchet 9/7/07 1:35 PM Page 261 Index kids/yearlings, 203, 209–210; limited male goats and, 209; male-biased harvest/problems, 208–209, 211– 212, 214; management of introduced goats, 214–215; management overview, 204, 206–212, 208; natural mortality vs., 196–197, 197; as not needed, 209; overview, 14; population decline detection and, 214; population dynamics and, 26, 27, 114, 196, 196–197, 197, 214; population ecology and, 208; recent history, 26; recommendations, 203, 209–210, 214; reproductive strategy and, 200, 201, 201; summary, 217–218; trophy hunting/effects, 114, 209; of twoyear-old males, 210–211 Ibex: horns/fighting, 103–104; Nubian, 79; photo, 9; sexual segregation, 79; Spanish, 103–104; survival, 171 Ibex, Alpine: horn growth, 104; sexual segregation, 79; survival rates, 79; weighing, 42 Immobilization of goats: description, 38; effects, 41, 119, 141, 142 Inbreeding, 212, 228 Industry threats, 6, 7, 31–32, 69, 217, 229 See also specific industries Introduced goat populations: harvest rates, 214; hunting of, 3, 214–215; in Olympic Mountains National Park, 11–12, 172, 173, 185, 215; overview, 11–13, 13; problems with, 11–12, 215; reproduction of, 215 Kids: anogenital patch, 43, 43; birthdate determination, 44–45; birthweight, 93; following birth, 56; horn growth, 101, 102; marking of, 38, 39; radio collar effects, 41; sexing of, 43, 43; survival, 120–121, 121, 125, 155, 159–163, 161, 162, 175; weighing, 42 See also Females, adultoffspring association; Nursery groups 261 Kids/body mass growth: description, 93–94, 93; maternal age and, 94 Kin groups: adult females and, 84, 85– 86, 87; analyzing, 85–86 Kudu, greater, 223, 225 Lactation energy costs, 56, 60, 99, 105, 127, 143–144, 145 Local resource competition hypothesis, 150, 153 Logging effects, 24, 229–230 Longevity: female reproductive success and, 122–123, 131–134, 132, 133, 134, 192, 195; survival, 166, 170–172 Long-term studies/marked individuals: access to study area, 226; components of, 17; conservation value of, 2, 227–228; contributions from, 1–2, 219, 222, 224–225; examples, 6, 223; funding, 2, 226; geographical distribution of, 225; hunted populations and, 198–199, 202, 227–228; limits to, 226–227; limits with, 222, 224– 225; management value, 227; pooling data from, 225; predator absence and, 225; short-term studies vs., 221, 226, 227 See also Caw Ridge mountain goat study (overview) Males, adult: associations and, 5, 43; daily movements of, 66–67; forest use by, 5, 47, 64–66, 65, 66, 75; future research on, 228; home range of, 49, 52–53, 55, 55; marking of, 38, 39; survival, 168–169; urination posture, 43 Managing goats: density dependence/ effects, 178; density independence/ effects, 178, 189; dispersal and, 175; herd specificity and, 208, 212– 213, 213; introduced goats, 214– 215; metapopulation dynamics and, 212–214, 213; overview, 204, 206–212, 208; reproductive senescence and, 123 See also Hunting/goats index:IP_Festa-Bianchet 262 9/7/07 1:35 PM Page 262 Index Marking goats, 38, 39, 40 Marmots, hoary, 24–25, 25 Maternal care: kid sex ratio and, 149, 153, 155; kid survival and, 134; measuring difficulties, 194; yearling growth/survival and, 193 Mean group size vs typical group size, 73–74 Metapopulation approach: conservation, 216; hunting policies, 207–208, 212–213, 213, 216; managing goats, 212–214, 213 Mice, deer, 24 Moose: hunting of, 200, 210, 211; predator avoidance, 74; reproduction, 126; in study area, 23 Mortality: causes, 158, 159, 175; hunting vs natural mortality/effects, 196–197, 197; overview, 157–158; sex-specific mortality, 106, 157, 165– 166, 166, 175 See also Survival Motorized vehicles threat, 7, 31, 216– 217, 218, 229 Mountain goats: ancestors, 8, 11; description, 8, 9, 10, 11, 12; distribution, 11–13, 13; estimated numbers of, 14; habitats of, 13–14; names for, 7; relationship with bighorn sheep, 14, 22–23; sex–age class names, 8; taxonomy, 7–8 Muskox, 33 Natural Sciences and Engineering Research Council of Canada (NSERC), 29, 225 Nisga’a People, 7, 14 NSERC (Natural Sciences and Engineering Research Council of Canada), 29, 225 Nursery groups: daily movements of, 66–67; definition, 49; forest use by, 64–65, 65, 66; home ranges, 49, 53– 54, 55; as single group, 74; size of, 72–74, 73; summer movement, 67– 68, 67, 69, 84 Oil exploration/effects, 17, 24, 25 Olympic National Park: goat dispersal at, 172, 173, 185; introduced goats/problems, 11–12, 215 Orientation avoidance display, 99 Phenolic compounds (forage), 58 Phoenix Park deer study, 225 Photoperiod and rut, 57 Phylopatry, 87 Pikas, 24 Plant phenology hypothesis (birthdate synchronization), 126, 127, 131 Population density: body mass growth and, 108–113, 110, 111, 112, 115, 116–117; density independence/effects, 177, 178, 189; fecal crude protein and, 109, 110, 110; foot length/ indicator study, 110, 111, 111; home ranges and, 109; horn growth and, 109–110, 111, 111, 112, 115; primiparity timing and, 142 See also Density dependence/effects Population dynamics/goats: age–sex structure/numbers (1990–2003), 29; density dependence absence, 185– 186, 188–189, 189; hunting effects, 26, 27, 114, 196–197, 197, 214; kid survival/adult female numbers, 179, 180, 182, 188, 189; population regulation and, 184–186, 188–189; recent history (study area), 26; removal/transplant program, 26, 27; resource availability and, 178–180, 181, 182, 185, 186, 188; sex–age class variability/effects, 29, 177, 183–184; statistical notes, 190; summary, 189–190; weather/snowmelt and, 180–181, 183, 184–185, 189; yearling mass and, 181–182, 182, 189 Population dynamics/ungulates: female reproductive strategy and, 194–195, 198, 203; hunting effects, 198–203, 201; predation and, 224 index:IP_Festa-Bianchet 9/7/07 1:35 PM Page 263 Index Population regulation, 184–186, 188– 189 Population size/goats: study area 1989– 2003, 28; study area current population, 17, 26; variability in, 177 See also Censusing goats Porcupine, 24 Predation: cursorial vs stalking predators, 88, 187, 188; events summary tables, 65, 159; forest use and, 64– 66, 65, 66, 74; habitat loss and, 69; major predators, 7, 64, 68; monitoring, 68, 158; on neonates/kids, 130– 131; prey risk tolerance, 68, 75; specialist/consequences, 186, 188, 224; ungulate population dynamics and, 224 Predation avoidance: area-use behavior, 68, 69; birthing isolation, 45, 56– 57, 72, 73; escape terrain, 48, 64, 69, 84, 85, 87; female gregariousness, 84, 86, 87, 187; group defense, 86; group size, 74; horns, 98; primiparity timing, 143 Predation hypothesis (birthdate synchronization), 126–127, 127–128, 130–131 Predator-prey equilibrium, 187–188 Present threat display, 99 Primiparity timing: body mass and, 141–142, 141, 143; chemical immobilization effects, 141, 142; competing factors, 142–143; conception timing and, 143; energy costs, 143; fitness costs, 142, 143, 144, 147, 155, 193; horn growth and, 141, 141; population density and, 142; predation risks and, 143; reproductive success and, 119–120, 119, 123–124, 135; weather and, 143 Radio collars: description/use, 40, 40; harmful/potential effects, 41, 42; mortality switch on, 40; numbers by sex/age/year, 40 263 Ram Mountain, Alberta bighorn sheep study, 18, 106, 220, 222, 225 See also Sheep, bighorn Reindeer: hunting, 211; reproduction, 140, 154; studies, 223 Remote sensing, 40, 42 Reproduction/goats: conception timing, 56; litter size, 132, 192, 194; monitoring status, 43, 44–45 Reproductive effort: mother’s age and, 150; reproductive strategy and, 148– 149, 155, 192–193, 203 Reproductive senescence, 122–123 Reproductive strategy: definition/description, 137–138; offspring sex ratio, 149–152, 151, 152, 154–155 Reproductive strategy/female goats: environmental harshness and, 193; fitness costs and, 143–146; hunting effects on, 200, 201, 201; maternal effort, 148–149, 155, 192–193, 203; maternal/offspring mass at weaning, 147–148, 147; maximizing maternal survival, 146–149, 147, 192–194, 195, 203; mother’s age/kid sex ratio, 151–152, 151, 152, 154–155, 155– 156; overview, 137–138; primiparity age and, 138, 141–143, 141, 147, 155, 193; statistical notes on, 156; twinning, 194 Reproductive success/female goats: bighorn sheep longevity comparison, 132, 133–135; birthdate/kid survival, 126–131, 128, 129, 135; Caw Ridge survival and, 119, 123; chemical immobilization effects, 41, 119, 141, 142; emigration before reproduction, 119; female age and, 118–125, 119, 121, 123, 124; female age/kid mass, 124–125, 124; female body condition/energy reserves, 125–126; female body mass and, 120, 125, 126; individual variability importance, 118; kid survival, 120–121, 121, 125; lactation energy costs, 56, 60, 99, index:IP_Festa-Bianchet 264 9/7/07 1:35 PM Page 264 Index Reproductive success (continued) 105, 127, 143–144, 145; longevity and, 122–123, 131–134, 132, 133, 134, 192, 195; of mature females, 122; phenotypic quality and, 122– 123; primiparity age, 119–120, 119, 123–124, 135; primiparity and female body mass, 120; reproductive senescence and, 122–123; statistical notes on, 136; summary, 135 Research: future research on Caw Ridge, 228–229 See also Caw Ridge mountain goat study (overview); Long-term studies/marked individuals Rush avoidance display, 99 Rush threat display, 99 Rut/rutting: behavior, 47; photoperiod and, 57; physical effects of, 56 Scan sampling, 45 Scientific names list, 233 Serow, Japanese: dispersal, 175; horn growth/lactation, 99–100; mountain goats and, 16; photo, 10 Sex differences: in aggression, 72, 76– 77, 82, 98, 99; in daily movement, 66–67; in forest use, 47, 64–66, 65, 66, 75; group size, 72–74, 73; predation risk tolerance, 68, 75; predation vulnerability, 88; summer habitat/ movement, 67–68, 67, 69, 80, 84 Sexual dimorphism: body mass, 79–81, 91, 94, 95, 106–107, 108, 113, 115; development in goats, 8, 155; evolutionary causes, 106; horns and, 8, 98, 102, 103, 107, 108, 113, 115; male mass/effects, 106; in polygynous species, 106 Sexual segregation: activity/time budgets and, 75, 78, 79, 80; among ungulates/mammals, 75; body mass and, 75, 78–80, 89–90, 89; description, 72–74, 73; explanations of, 75–81; predation risk and, 75; summary, 88– 89; temporal vs spatial, 72; young males and, 76–80, 77 Sheep: female kin groups and, 87; home range/area-use pattern of, 49; reproduction, 140 Sheep, bighorn: body mass effects, 91, 92, 96; body mass/growth, 92, 93, 94, 106–107, 107, 108; capture/handling of, 41, 47; density dependence of, 135; dispersal, 174, 175; female kin groups and, 84, 85, 87; forage/ foraging, 58, 60–61, 60; helicopters and, 33; home range/area-use pattern of, 49, 109; horns/horn growth, 100, 103, 104, 107–108; horn size effects, 92; hunting/effects, 211, 224, 227; lamb survival, 160, 162; mother–offspring association, 82; mountain goat relationship with, 14, 22–23; pneumonia epizootics, 135, 144; population density studies, 109; predation/avoidance, 88, 186, 187, 188, 224; predation on, 134–135; ram “breeding commutes,” 174; reproduction, 92, 120, 126, 130, 131, 132, 133–135, 144, 145, 147, 149, 150, 152, 154, 174, 195; reproductive success factors, 92; resource availability indicator, 181; sex differences/ summer movement, 68; sexual segregation, 78; study areas, 18; survival, 167, 168, 198; weighing, 42 Sheep, Dall’s, 103 Sheep River study area, 18, 226 Size of groups: mean group size vs typical group size, 73–74; monthly variation in, 72–74, 73; nursery vs bachelor groups, 72–74, 73; predation and, 74 Skeletal development, 97–98, 97, 113 Smith, Kirby, 27, 173 Snow/snowmelt: description, 18, 19, 19, 20–21, 23; mortality and, 57; reproduction and, 57 See also Weather index:IP_Festa-Bianchet 9/7/07 1:35 PM Page 265 Index 265 Social organization: description overview, 71; implications overview, 88; “natural selection” and, 71, 84 See also specific traits Squirrels: Columbian ground, 24; golden-mantled ground, 24 Submissive behavior, 45, 99 SURGE program, 165 Survival: elasticity in, 214; overview, 157–158; statistical notes on, 176; of yearlings (overview), 163–164, 175 See also Mortality Survival of adult goats: age structure of females, 169–170, 170; analysis methods, 165; female survival, 166– 168, 169–170, 169, 170, 175; longevity, 166, 170–172; male survival, 168–169; sex-specific survival, 106, 157, 165–166, 166, 175; survival senescence, 168, 171 Survival of kids: body mass and, 162– 163, 162; maternal age and, 155; maternal reproductive strategy, 120– 121, 121, 125; overview, 159–163, 161, 162, 175 Survival senescence, 168, 171 Typical group size vs mean group size, 73-74 Taxonomy of mountain goats, 7–8 Traps: aggression in, 38; deaths from, 158, 159; description, 37–38, 37, 43; visitation frequency, 38 See also Capturing/handling goats Trap site map, 22 Trivers-Willard hypothesis, 149, 150, 153-154, 155, 156 Trophy hunting/effects: bighorn sheep, 211, 224; goats, 114, 211 Yearlings: associations with mother, 81–84, 81, 82, 83, 84; body mass growth, 94, 95; horn growth, 101, 102; marking of, 38, 39; survival of, 163–164, 175 See also Nursery groups Yellowstone National Park and goats, 12–13, 208, 215 Urquhart, Martin, 27, 29 Vegetative sampling: biomass/seasonal changes, 62–63, 63; description, 46, 46, 62–63 Wapiti, 23 Weaning success: defining, 45; future success and, 145–146; maternal/offspring body mass, 147–148, 147; sons vs daughters, 146 Weather: Caw Ridge study area, 17–18, 19–21, 19, 46–47; population dynamics and, 180–181, 183, 184–185, 189; primiparity timing and, 143 See also Snow/snowmelt Weighing goats: description, 40, 42, 43, 47, 92; future research and, 227– 229 Willmore Wilderness Park, 25–26, 230 Wolverines, 24 Wolves, 64, 65, 68, 88, 188, 199 Woodrats, bushy-tailed, 24 index:IP_Festa-Bianchet 9/7/07 1:35 PM Page 266 bd of dir:IP_Festa-Bianchet 9/7/07 1:36 PM Page 267 Island Press Board of Directors Victor M Sher, Esq (Chair) Sher & Leff San Francisco, CA Dane A Nichols (Vice-Chair) Washington, DC Carolyn Peachey (Secretary) Campbell, Peachey & Associates Washington, DC Drummond Pike (Treasurer) President The Tides Foundation San Francisco, CA Robert Baensch Director, Center for Publishing New York University New York, NY Charles C Savitt President Island Press Washington, DC William H Meadows President The Wilderness Society Washington, DC Susan E Sechler Senior Advisor The German Marshall Fund Washington, DC Merloyd Ludington Lawrence Merloyd Lawrence Inc Boston, MA Nancy Sidamon-Eristoff Washington, DC Henry Reath Princeton, NJ Will Rogers President The Trust for Public Land San Francisco, CA Alexis G Sant Trustee and Treasurer Summit Foundation Washington, DC Peter R Stein General Partner LTC Conservation Advisory Services The Lyme Timber Company Hanover, NH Diana Wall, Ph.D Director and Professor Natural Resource Ecology Laboratory Colorado State University Fort Collins, CO Wren Wirth Washington, DC ... iii Mountain Goats FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page iv FM:IP_Festa-Bianchet 9/7/07 1:19 PM Page v Mountain Goats Ecology, Behavior, and Conservation of an Alpine Ungulate Marco Festa-Bianchet... results of other long-term studies of ungulates, including other mountain ungulates such as bighorn sheep in Alberta, ibex in the Italian and French Alps, and both Alpine and Pyrenean chamois in France... Survival and Dispersal 157 10 Density-Dependence and the Question of Population Regulation 177 11 Female Reproductive Strategy and Ungulate Population Dynamics 191 12 Management and Conservation of Mountain