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Antipredator Strategies of Alaskan Moose: Are Maternal Trade-Offs Influenced by Offspring Activity?

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Antipredator Strategies of Alaskan Moose: Are Maternal Trade-Offs Influenced by Offspring Activity? Color profile: Disabled Composite Default screen 2055 Antipredator strategies of Alaskan moose: are maternal trade-offs influenced by offspring activity? Kevin S. White and Joel Berger Abstract: To maximize fitness, mothers must both provision and protect neonates, demands that may be in conflict, particularly in systems that still experience high levels of natural predation. Whether variation in offspring behaviour alters this putative conflict is not known. The objective of this study was to test hypotheses about the extent to which neonatal activity and ecological variables mediate trade-offs between maternal vigilance and foraging. To address these questions we contrasted data from behavioural observations on female moose (Alces alces) that differed in parity, calf activity, and habitat use at a site in south-central Alaska where they are subject to high levels of grizzly bear (Ursus arctos) and wolf (Canis lupus) predation. Our analyses revealed that females with active juveniles were more vigilant (and as a consequence spent less time feeding) than those with inactive young; vigilance of females without attendant young was intermediate. Distance to apparent protective refugia (e.g., vegetative cover) was positively related to vigilance for all calf-status categories, but lactating females spent more time closer to thick vegetation than did nonlactating females. These results suggest that (i) mothers adjust vigilance when young are inactive to compensate for the loss of foraging opportunities during periods of neonate activity, thereby reducing juvenile vulnerability and in- creasing the overall feeding rate, and (ii) females with young reduce foraging compromises and, presumably, predation risk by spending more time close to protective cover than do nonlactating females. We conclude that maternal trade- offs can be highly labile and that mothers are able to adjust rapidly to environment-specific situations. Résumé : Pour maximiser le fitness, les mères doivent à la fois approvisionner et protéger leurs nouveau-nés, deux exigences qui peuvent être en conflit, particulièrement dans les systèmes où il se fait beaucoup de prédation naturelle. Le comportement de la progéniture peut-il modifier cet éventuel conflit? Nous avons éprouvé les hypothèses selon les- quelles l’activité des nouveau-nés et certaines variables écologiques forcent les mères à faire des compromis entre la vigilance et la recherche de nourriture. Pour résoudre ces problèmes, nous avons comparé les données de comportement obtenues chez des femelles de l’Orignal (Alces alces) distinctes par le nombre de leurs mise-bas, l’activité de leurs petits et leur utilisation de l’habitat à un site du centre-sud de l’Alaska où il se fait une importante prédation par les Grizzlis (Ursus arctos) et les Loups (Canis lupus). Nos analyses ont démontré que les femelles dont les jeunes sont ac- tifs sont plus vigilantes (et, conséquemment, vouent moins de temps à l’alimentation) que les femelles dont les jeunes sont inactifs; les femelles sans petit occupent une position intermédiaire. La distance jusqu’à un refuge apparent (e.g., la couverture végétale) est en corrélation positive avec la vigilance, quel que soit le statut des nouveau-nés, mais les femelles nourricières passent plus de temps dans le voisinage d’une végétation épaisse que les femelles non nourriciè- res. Ces résultats indiquent (i) que les mères ajustent leur vigilance quand les petits sont inactifs de façon à compenser la perte des occasions de recherche de nourriture durant les périodes d’activité des nouveau-nés, réduisant de cette fa- çon la vulnérabilité des jeunes, tout en augmentant la fréquence des périodes d’alimentation et (ii) que les femelles avec des petits réduisent les compromis et, présumément, les risques de prédation en passant plus de temps dans le voisinage de couvertures protectrices que les femelles non nourricières. Les compromis envisagés par les mères sont donc très labiles et les mères sont en mesure de s’ajuster rapidement à des situations environnementales spécifiques. [Traduit par la Rédaction] 2062 Introduction White and Bergertrient gain while minimizing costs associated with acquiring energetic resources (Krebs 1973; Charnov 1976). However, a Predictions of “optimal foraging” indicate that individuals more complete assessment of fitness correlates results when develop foraging strategies that maximize energetic and nu- such factors as predation, disease, and abiotic variability are Received November 1, 2000. Accepted September 7, 2001. Published on the NRC Research Press Web site at http://cjz.nrc.ca on November 23, 2001. K.S. White.1,2 Department of Environmental and Natural Resource Sciences, University of Nevada, 1000 Valley Road, Reno, NV 89512, U.S.A. J. Berger. Program in Ecology, Evolution, and Conservation Biology, University of Nevada, 1000 Valley Road, Reno, NV 89512, U.S.A., and Wildlife Conservation Society, P.O. Box 340, Moose, WY 83012, U.S.A. 1Corresponding author (e-mail: [email protected]). 2Present address: Division of Wildlife Conservation, Alaska Department of Fish and Game, P.O. Box 240020, Douglas, AK 99824, U.S.A. Can. J. Zool. 79: 2055–2062 (2001) DOI: 10.1139/cjz-79-11-2055 © 2001 NRC Canada J:\cjz\cjz79\cjz-11\Z01-170.vp Tuesday, November 20, 2001 8:25:19 AM Color profile: Disabled Composite Default screen 2056 Can. J. Zool. Vol. 79, 2001 considered (Stephens and Krebs 1986). In this context, may also be used in concert with vigilance to achieve an mammalian systems can offer unique insights. In ungulates, optimal offspring protection/provisioning strategy. Use of for example, mothers face challenges because they must protective cover is one behavioural means by which preda- balance increased foraging demands associated with the high tion risk is reduced (Mech 1966; Caraco et al. 1980; Lazarus energetic costs of lactation (Bunnell and Gillingham 1985; and Symonds 1992; Molvar and Bowyer 1994; Kunkel and Oftedal 1985; Carl and Robbins 1988) with the protection of Pletscher 2000). Protective cover (after Lazarus and Symonds vulnerable neonates from predators, a trade-off that may be 1992) inhibits prey detection, facilitates escape, and reduces especially important in populations which still experience the capture efficiency of visually oriented predators (Daly et intense natural predation. al. 1990; Longland and Price 1991). If prey with vulnerable Not only can predation be an important source of mortal- young confine their foraging activity to areas in or near veg- ity in terrestrial and aquatic systems (Frame 1974; Gasaway etative refugia, foraging compromises might be minimized et al. 1992; Sih et al. 1992), but variation in the intensity of (Molvar and Bowyer 1994; Weixelman et al. 1998). predation can modify antipredator responses (Berger 1998; Here we assess behavioural patterns of maternal invest- Hunter and Skinner 1998). Since behaviour can be linked to ment, specifically the trade-off between offspring protection predation in numerous organisms (Lima and Dill 1990), and subsequent provisioning, by using moose in a system study of trade-offs between foraging and antipredator behav- where offspring are subjected to high mortality due to preda- iour can be enhanced with knowledge of actual predation tion. This issue is of interest for two principal reasons. First, risk. Few studies, however, have relied on direct measures of most studies investigating the trade-off between predation predation pressure (White 1999), and the resulting lack of risk and foraging were carried out at sites where rates of direct knowledge has precluded more precise assessments of prey mortality are unknown and therefore the risk of predation how predation shapes behaviour. Acquiring an understanding has not been directly assessed (White 1999). Second, al- of such relationships in the case of some types of organisms, though lactating females may be more vigilant than non- such as large mammals in human-dominated landscapes, mothers (Lipetz and Bekoff 1982; FitzGibbon 1993; Molvar may be impossible. For example, grizzly bears (Ursus arctos) and Bowyer 1994; Burger and Gochfield 1994), the extent to and wolves have been extirpated from about 98% of North which offspring activity affects maternal antipredator behav- America from the Canadian border south, which means that iour is unclear and to our knowledge has never been investi- predation on some herbivores as a selective force has been gated. It may be possible to gauge whether risk perception lost (e.g., on bison (Bison bison) and moose (Alces alces); by mothers changes when offspring vulnerability to preda- Berger 1998, 1999). tion is exacerbated by activity of the offspring, and, further, Investigation of maternal antipredator behaviour in mam- to determine whether variation in offspring behaviour modu- mals has focused on habitat shifts (Festa-Bianchet 1988; lates the trade-off between food-acquisition and predator- Berger 1991; Kohlmann et al. 1996; Rachlow and Bowyer avoidance behaviour. 1998), presence or absence of young (Lipetz and Bekoff To examine these issues we tested two principal hypothe- 1982; Burger and Gochfield 1994; Molvar and Bowyer 1994), ses: (1) lactating females adjust vigilance and foraging in effects of predator–scavenger associations (Berger 1999), relation to putative neonate vulnerability to predation, and offspring age (Caro 1994), mother–young spatial relation- (2) females with young spend more time in close proximity ships (Byers and Byers 1983; FitzGibbon 1993; Byers
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