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Wolf behavior: reproductive, social and intelligent

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... Wolves are renowned for their behavioral and dietary plasticity (Peterson & Ciucci, 2003), allowing them to acquire resources in a variety of habitats (Mech & Peterson, 2003). However, wolf activity patterns and space use become focused on natal sites during denning season as pups have reduced mobility (Mech & Boitani, 2003;Packard, 2003). Pups are dependent on milk during the first 5 weeks of life after which time they begin to develop teeth and the ability to digest food regurgitated and carried to them by adults (Packard, 2003). ...
... However, wolf activity patterns and space use become focused on natal sites during denning season as pups have reduced mobility (Mech & Boitani, 2003;Packard, 2003). Pups are dependent on milk during the first 5 weeks of life after which time they begin to develop teeth and the ability to digest food regurgitated and carried to them by adults (Packard, 2003). Because adult wolves, and especially breeding wolves (Mech & Boitani, 2003;Packard, 2003), need to make frequent trips to the den site to provision pups, their movements become constricted and their core use areas around den sites are typically smaller than during the rest of the year . ...
... Pups are dependent on milk during the first 5 weeks of life after which time they begin to develop teeth and the ability to digest food regurgitated and carried to them by adults (Packard, 2003). Because adult wolves, and especially breeding wolves (Mech & Boitani, 2003;Packard, 2003), need to make frequent trips to the den site to provision pups, their movements become constricted and their core use areas around den sites are typically smaller than during the rest of the year . Non-breeding wolves also play an important role in both attending and provisioning the pups especially before weaning due to the limited ability of the breeding female to leave the den for extended periods (Packard, 2003;Ruprecht et al., 2012). ...
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Although wolves are wide-ranging generalist carnivores throughout their life cycle, during the pup-rearing season wolf activity is focused on natal den sites where pup survival depends upon pack members provisioning food. Because prey availability is influenced by habitat quality within the home range, we investigated the relative importance of prey species for adults and pups and further examined the relationship between habitat characteristics, wolf diet, and litter size on Prince of Wales Island (POW) in Southeast Alaska. During 2012-2020, we detected 13 active den sites within the home ranges of nine wolf packs. We estimated minimum pup counts using motion-detecting cameras and individual genotypes from noninvasive samples (hair: n = 322; scat: n = 227) and quantified wolf diet composition using fecal DNA metabarcoding (n = 538). We assessed habitat composition, configuration, and connectivity within denning and annual home ranges estimated using wolf GPS-collar data. Contrary to expectations, wolves had a more constricted diet during denning season (April 15-July 31), and within this season pups had a narrower dietary niche (species richness [S] = 4) focused more on deer (relative frequency of occurrence [O/I] = 0.924) than adults (S = 15; deer O/I = 0.591). Litter size had a positive relationship with the relative frequency of deer in a wolf pack's diet. Wolf consumption of deer was positively associated with the proportion of young-growth forest (≤25 years old) within denning and annual home ranges. High levels of vegetation patch interspersion, and the density of closed logging roads were also important predictors, suggesting these habitat qualities were influential for increasing the availability of deer to wolves. Our results contrast with previous research indicating wolf pup diets included more alternate prey (i.e., beaver) than adults and emphasize the importance of deer to wolf viability on POW, especially during denning season.
... This may be an example of social learning between mother and pups, something that has previously been recorded only when hunting rabbits (Corbett and Newsome 1975), but has been extensively studied in other predators such as the wolf . In wolves, not all components of hunting and foraging behaviour are innate; for example, where to find food and how to kill (Packard 2003). These behaviours are learned over time via observation and experience (Mech 1988(Mech , 1991Fentress 1992). ...
... These behaviours are learned over time via observation and experience (Mech 1988(Mech , 1991Fentress 1992). Wolf pups often join hunts and are able to learn the more subtle aspects of hunting and killing (Packard 2003). It is possible that Observations 1, 2, and 3 give a glimpse into the social habits of hunting dingoes; however, more data are necessary to explore this further. ...
... The macropods that dingoes target also employ it (Behrendorff 2018); just like wolves, dingoes will enter the water in pursuit of prey (Purcell 2010b) and have even overcome and drowned their prey on occasion (Behrendorff 2018). Moreover, young wolves often accompany older wolves on hunts to learn aspects of predatory behaviour (Mech 1991;Packard 2003). We also observed evidence of social learning of hunting behaviour in the dingoes (mother: Dingo 1; pups: Dingoes 3, 4, and 5; Observations 2, 3, and especially 1). ...
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Context. The behaviours used by mammalian predators to track, kill, and consume prey are some of the most dynamic interspecific interactions in nature. However, they are often challenging to follow through the landscape and observe directly without disturbing the animals being watched. Aims. We describe the behaviours used by wild dingoes while hunting macropods in Namadgi National Park, Australian Capital Territory, Australia. Methods. Footage was initially captured by wildlife cinematographers on behalf of documentary programs and was made available for viewing after production. Hunting events were filmed from an altitude of >50 m by using a 'long lens' fitted to either a drone or helicopter. Results. We recorded a suite of hunting behaviours that would have been extremely challenging to observe from the ground via traditional methods. This includes some of the first video records published in the scientific literature of the behaviours used by dingoes to hunt and kill macropod prey, as well as some rare observations of mother and pup hunting dynamics. We did not observe any signs of disturbance as a result of filming for either predator or prey. Conclusions. The varied repertoire of predatory behaviours displayed by dingoes is similar to that documented in wolves and asserts them as a behaviourally complex top predator in the Australian landscape. In addition, we highlight the use of drones as a valuable approach for directly observing wild behaviours. They offer a minimally invasive and relatively inexpensive and accessible alternative to helicopters. This project is also a case study exemplifying the value of collaborations between filmmakers and researchers that enable the sharing of archival documentary footage for the study of wild animal behaviour. Implications. Future studies of wild animal behaviour should consider employing drones (at a safe distance and in accordance with published best practices and guidelines) as an additional tool to collect types of data that would be challenging using other methods.
... Although much is known about the behavior and development of young, hand-raised or captive wolf pups (synthesized by Packard, 2003), much less such information has been recorded for free-ranging wolf pups during their first several weeks of age. General descriptions of early wolf pup behavior during life around wolf dens are available (Murie, 1944;Clark, 1971;Mech, 1988;Packard, 2003) as well as more detailed information about critical aspects of early pup behavior such as nursing (Packard et al., 1992) and provisioning by adults (Mech et al., 1999). ...
... Although much is known about the behavior and development of young, hand-raised or captive wolf pups (synthesized by Packard, 2003), much less such information has been recorded for free-ranging wolf pups during their first several weeks of age. General descriptions of early wolf pup behavior during life around wolf dens are available (Murie, 1944;Clark, 1971;Mech, 1988;Packard, 2003) as well as more detailed information about critical aspects of early pup behavior such as nursing (Packard et al., 1992) and provisioning by adults (Mech et al., 1999). However, the only specific information for various early ages that is available for free-ranging wolf pups is contained in descriptions of pup behavior during different days when pups were 9 to 74 days old on Baffin Island, Nunavut, Canada (Clark, 1971). ...
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Whereas much is known about the behavior and development of captive young wolf (Canis lupus) pups, less detail has been published about some aspects of free-ranging wolf pup behavior. This article synthesizes 42 observations of free-ranging Arctic wolf pups from ages 13 through 52 days made during 10 summers from 1987 through 2006 on Ellesmere Island, Nunavut, Canada. Besides listing key behaviors such as howling and caching, I record unique observations of ages of pup urination without adult stimulation (22, 33, 42, 52 days), knowledge of which is important to studies of wolf domestication, and of a 48-day-old pup that traveled 39 km during a 12 h and 19 min round trip between the den and a prey carcass, including a 26.5 km trek in 5 h. These observations should lead to a deeper and more complete understanding of this critical period of pup growth and development.
... Lastly, we pooled all estimated change-points and defined the birthing period as the range from the earliest to the latest date of change (Appendix Table A1). In addition, since the gestation period of wolves ranges from 61 to 64 days (Mech, 1970;Packard, 2003;Pulliainen, 1965) we subtracted 64 days from the earliest and latest birth range dates to estimate the mating period. ...
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Social organization in animals is a fundamental factor driving population dynamics and individual spatial distribution. Affiliation among kin is common in social groups, but kinship is no safeguard against intraspecific competition. Within social groups, the closest competitors are often related. In this study, we present 14 years of GPS-position movement data from 65 pairs (i.e. breeding pair, pup–adult breeder and pup–pup) of Scandinavian wolves, Canis lupus. We investigated social behaviour by examining spatial and temporal patterns of cohesion through winter to early summer (January to July) including two key reproductive periods, mating and birth, using mixed regressive nonlinear time series analysis (GAMM). During the mating period, breeding wolf pairs remained cohesive, while offspring increasingly dissociated. Offspring became even more solitary until the next birth period, when most permanently left their natal pack and territory. We suggest that the social organization of wolves may be modulated by intraspecific competition related to social aggression during the mating period and food competition around the birthing period. Early independence followed by immediate natal dispersal may be advantageous to both offspring and parents if the chances of finding food and a breeding partner and settling in a vacant area are high.
... We calculated daily vocal rates (solo-howl rates and chorus-howl rates) of wolf packs for RS periods (number of vocalization events/number of sampling days), considering only days with all the scheduled hours recorded. Central daylight hours generally correspond to wolf inactivity (Packard 2003). Thus, to estimate daily vocal rates, we defined "days" as 24 h beginning at noon local time (wolves commonly begin activity at dusk and end at dawn). ...
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We used automatic sound recorders to study spontaneous vocalizations of wild wolves during the pup-rearing season around rendezvous sites from 24 wolf packs in six study areas across North America, Asia, and Europe. Between 2018 and 2021, for a total of 1225 pack-days, we recorded 605 spontaneous wolf chorus howls and 224 solo-howl series. Howling occurrence varied across areas, from 12.50 to 94.12% days with howling. Daily howling ranged from 0.00 to 3.47 solo howls/day and 0.13 to 5.29 chorus howls/day. Generally, spontaneous chorus howls peaked between sunset and sunrise. Howling rate depended on area, pack size, and density of people living nearby, being greater where fewer people lived. High rates in Yellowstone National Park, (800,000+ visitors during the study) could reflect accommodation to human activities such as wolf watching. One to six automatic recorders per site within 1000 m of rendezvous sites needed 4-15 days to detect the pack (average 9.5) and 5-21 days (average 11.3) to detect pups, both with a probability of 95%. Our results may guide wolf-monitoring programs using automatic sound recorders, a promising method offering advantages over howling surveys, especially in human-dominated landscapes.
... Fathers of feral domestic dogs Canis lupus familiaris reportedly guard the nest site and regurgitate food for the pups (Pal 2005). Gray wolf fathers Canis lupus also reportedly participate in guarding the den (Ruprecht et al. 2012) and feed the nursing mother, and when the cubs leave the den around weaning, the breeding male as well as other adults regurgitate food for them (Packard 2010). Even the maned wolf Chrysocyon brachyurus, thought to be solitary, has been observed to spend time sleeping together with its mate, and after birth of the cubs, the male stays in the vicinity, reducing his home range and activity levels, suggesting that he may contribute to parental care. ...
Article
The mammalian order Carnivora is generally defined as species that feed exclusively or to some degree by eating other animals. The Carnivora comprise around 280 species, divided into 16 families, 13 of which are terrestrial and 3 aquatic. Carnivores are spread across the entire planet, including the two polar regions and on land and sea. Consistent with such diverse ecologies, there is no typical pattern of parental care distinguishing carnivores from other mammals. Using examples from different taxonomic families, our aim is to illustrate the diversity of parental care in Carnivora. Major topics include parental care before and after birth of the young, paternal, and alloparental care and the process of weaning. Given the position of many carnivores at the apex of food chains, a greater understanding of their patterns of parental care as a vital part of reproductive biology is essential to conservation programs.
... Fathers of feral domestic dogs Canis lupus familiaris reportedly guard the nest site and regurgitate food for the pups (Pal 2005). Gray wolf fathers Canis lupus also reportedly participate in guarding the den (Ruprecht et al. 2012) and feed the nursing mother, and when the cubs leave the den around weaning, the breeding male as well as other adults regurgitate food for them (Packard 2010). Even the maned wolf Chrysocyon brachyurus, thought to be solitary, has been observed to spend time sleeping together with its mate, and after birth of the cubs, the male stays in the vicinity, reducing his home range and activity levels, suggesting that he may contribute to parental care. ...
Article
Direct care of offspring by the father (sire) is relatively rare in primates. Besides humans, there are a number of species where the male is essential for the survival of offspring: marmosets, tamarins, titis and owl monkeys, some lemurs, and siamangs. All these species show reduced sexual dimorphism, territoriality, and biparental care. However, timing and levels of direct care may vary among these species. Here, relying on both lab and field data, we address the variability found in father's involvement with his infants, the behavioral, neuroendocrine and sensory systems that are a cause and consequence of paternal care, and social bonds between the breeding pair. We integrate studies of laboratory animals (where detailed observations and experimentation are possible) with field studies (which illuminate the ecological and evolutionary functions of paternal care) and discuss the future directions for examining the proximate and ultimate mechanisms of paternal care in nonhuman primates.
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Abstract. Wolves are currently recolonising their historic range in France. The collection of scats is a widely used a non-invasive survey method to monitor wolf population size. However, seasonal changes in wolf faecal deposition patterns might affect the results of surveys. We used a detection dog and camera trapping (CT) to compare wolf scat detectability during winter and the nursing season. We collected 113 scats deposited by adult wolves at 29 marking sites on forest roads in the Sainte-Baume Regional Park, Provence, France. After parturition, the mean number of adult wolf scats increased by 160% inside the nursing territory and decreased by 80% outside of it. Around the time the pups are born, changes in faecal deposition patterns of adults make it easier to find scats around the wolf den (87% probability per wolf marking site) and harder to find scats outside the nursing territory (11% probability). During winter, the chance to find scats is equal (38 to 40% probability per wolf marking site) inside vs. outside the nursing territory. The combined use of a detection dog and camera traps allowed us to gather data on wolf defecation patterns non-invasively. Detectability of adult wolf scats during the nursing season is highly variable compared to winter due to seasonal behavioural changes affecting scat location. We conclude that surveys to collect samples and estimate wolf population size should be conducted exclusively during winter to avoid sampling biases.
Chapter
Several factors may underlie the ability of animals to solve problems in the physical environment including motivation, attention and inhibitory control. Here we review studies comparing wolves and dogs on these aspects. The data available to date suggest that while wolves are more neophobic than dogs, they are also more explorative and manipulative when encountering new objects in their environment. These are important differences since they are likely to affect results in many (social and non-social) tasks. Indeed, these results would suggest that, even if wolves and dogs do not differ in their cognitive abilities, wolves would outperform dogs in all sorts of physical tasks just because they have a better chance to stumble on the solution during their more persistent exploration. In combination with being more sensitive towards details of a situation and maybe better working memory capacities, wolves should be in general better problem-solvers. This is indeed what emerges, although a number of aspects still require further investigation. With regard to inhibitory control, dogs outperform wolves in situations where humans are involved, potentially because they are more sensitive to human social inhibition than wolves. But in contexts where humans are not involved, wolves and dogs inhibitory control abilities do not differ.
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