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Wolf behavior: reproductive, social and intelligent

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Jane M. Packard at Texas A&M University
Jane M. Packard
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... Being a K-strategist, the wolf is one of the longest-lived canids, with a moderate litter size and considerable investment in rearing the young. Thus, the population size is constantly close to the capacity limit of its habitat or available prey (Mech and Boitani, 2010;Packard, 2003) and will be oriented toward wildlife density (Guimarães et al., 2022;Mech and Boitani, 2010) unless livestock are available as easy prey (Ansorge and Balzer, 2019;Smith et al., 2000;Treves et al., 2004). In addition, socially active animals impart knowledge and experience within the pack to perfect specific prey-catching methods (Mech and Boitani, 2010;Packard, 2003). ...
... Thus, the population size is constantly close to the capacity limit of its habitat or available prey (Mech and Boitani, 2010;Packard, 2003) and will be oriented toward wildlife density (Guimarães et al., 2022;Mech and Boitani, 2010) unless livestock are available as easy prey (Ansorge and Balzer, 2019;Smith et al., 2000;Treves et al., 2004). In addition, socially active animals impart knowledge and experience within the pack to perfect specific prey-catching methods (Mech and Boitani, 2010;Packard, 2003). This means that once a pack has successfully habituated to livestock prey, it is destined to capture it (Boitani, 2000;Treves et al., 2004). ...
... As expected, the risk of an attack on livestock decreased with the maximum distance to a wolf territory. However, if wolves roam through an area not yet occupied by a wolf pack, there is increased pressure on the livestock from these dispersed wolves (Imbert et al., 2016;Mech and Boitani, 2010;Packard, 2003). In newly occupied regions, the risk of a wolf attack on livestock was considerable (Müller, 2020;Reinhardt and Kluth, 2007). ...
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... roads and human settlements (Hebblewhite et al. 2005, Kaartinen et al. 2005, Karlsson et al. 2007, Ordiz et al. 2015, Carricondo-Sanchez et al. 2020. However, the strength of the effects depends on season and time of day, and it seems likely that wolves are the most vulnerable to disturbance during the denning and rendezvous season (Houle et al. 2010), when movements of breeding wolves and their pack members are concentrated to the denning area due to the restricted mobility of the pups (Jedrzejewski et al. 2001, Packard 2003. A study on wolf area use during summer in Scandinavia showed that wolves chose daytime resting sites at intermediate distances to gravel roads and human settlements, at large distances to main roads, and that they avoided open areas (Zimmermann et al. 2014). ...
... There are several features of wolf behavioural ecology that might have influenced these results. During the denning and rendezvous seasons, movements of breeding wolves are concentrated near the sites where pups are located (Jedrzejewski et al. 2001, Packard 2003, with a successive increase in area use over time as the pups become more mobile. As a result, breeding pairs typically have smaller activity centres, relative to their annual home ranges, than non-breeding pairs. ...
... In contrast, the seasonal activity centres are formed by processes at a much wider spatio-temporal scale than the diel activity patterns of wolves. Breeding behaviour (Jedrzejewski et al. 2001, Packard 2003 and seasonally changing weather conditions, which can lead to a change in prey distribution and therefore change of wolf movement patterns (Fuller 1991), could be drivers of such processes. ...
Will future wind power development in Scandinavia have an impact on wolves?
Article
Full-text available
  • May 2024
The global energy demand is growing, and the world is shifting towards using more renewable energy, like increased onshore wind power development. We used Global Positioning System (GPS) and Very High Frequency (VHF) location data from adult, territorial wolves Canis lupus in Scandinavia (Sweden and Norway; 1999–2021), to examine the potential for wind power development to affect wolf behavioural ecology. We examined the spatial overlap of areas proposed for wind power development with wolf territory activity centres prior to construction, to test to what extent overlap varies with season, time of day and social status (breeding versus non‐breeding wolves). Measures of overlap were the distance between wolf activity centre points and nearest proposed wind turbine, the probability of proposed wind turbines being within the activity centre, and the density of proposed wind turbines within the activity centre. The wolf activity centre points were closer to sites of proposed turbines in early summer than in late winter and the density of proposed turbines in the activity centre was higher in early summer than in late winter. These findings probably result from an altitudinal shift in wolf area use between summer and winter. We also found that the probability for proposed turbines to be within the activity centre was higher for non‐breeding than for breeding wolves during early summer, whereas it was higher for breeding compared to non‐breeding wolves during late winter. This difference might be an effect of that breeding wolves have a restricted area use during the early summer season (denning period), resulting in a lower probability of turbines being inside their activity centre as compared to late winter. There was no clear pattern for other seasonal and social status differences. The results should be viewed as a starting point for further research and supplemented with before‐after studies.
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... Breeding females will directly care for and feed the pups during the early months after birth, and the male breeder will contribute indirectly to pup care by provisioning the lactating mother with food and through territory defense Boitani 2003, Boyd et al. 2023). Similarly, nonbreeding individuals may also help rear young through behaviors such as provisioning and pup-guarding (Packard 2003). Studies of gray and red wolves (C. ...
... By contrast, during the dispersal season, pups are moving with the pack outside of their rendezvous sites for the first time and may be naïve to the dangers presented by hunters and trappers (Packard 2003). Additionally, some nonbreeding adults from a group will be dispersing and using roads and trails to travel longer distances during hunting and trapping season (Fritts 1983, Pusey 1987, Mech and Boitani 2003. ...
Gray wolf breeders are more vulnerable to harvest during the breeding season
Article
Full-text available
  • Nov 2024
In cooperatively breeding carnivores, breeders are vital to perpetuating the group; the death or removal of an individual breeder can greatly affect group composition, genetic content, and short‐term population growth. Understanding the number of breeders harvested and timing of harvest can increase our knowledge of how mortality affects groups of cooperative breeders. Gray wolves (Canis lupus) in Idaho, USA, are exposed to annual harvest and are an ideal species for studying the effects of harvest on breeder turnover. We combined genotypes from tissue samples of harvested wolves with parentage analyses and cementum annuli ages and estimated when and how many breeding wolves were harvested. We genotyped and aged 229 adults and 203 pups using tissue and tooth samples from wolves harvested between 2014 and 2016. We identified a minimum count of 33 breeders in the harvest and found that they were disproportionately harvested more during the breeding season. We estimated that a minimum of ~14.5% of adult wolves harvested annually, or approximately 1 in 7, were breeders. We posit their behavior during breeding season may increase their vulnerability to harvest. By linking animal life history with vulnerability to human‐caused mortality we show that managers could structure harvest seasons so there is less overlap with wolves’ breeding season if there is concern about the demographic consequences of harvesting breeders.
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... Males and females can disperse at any age, though most commonly as yearlings or subadults (Gese and Mech 1991, Kojola et al. 2006, Morales-González et al. 2022, after which they may disperse over vast distances to acquire suitable territory (Wabakken et al. 2001, Mech and Boitani 2003, Ražen et al. 2016. Once suitable territory is found, colonisation success is increased by their highly flexible diet (Newsome et al. 2016) and rapid reproduction (Packard 2003). In the Northern Hemisphere, wolves have historically occurred in almost all habitats with sufficient food resources, though they generally display a preference for two complementary features, presence of forests and an absence of human settlements and roads (Kaartinen et al. 2005, Jedrzejewski et al. 2008, Ronnenberg et al. 2017. ...
... Features such as choice of prey (Fuller et al. 2003), landscape topography (Kauffman et al. 2007, Kittle et al. 2015 and phase of colonisation (Mech and Boitani 2003) can also act as important determinants. Finally, the HR of each individual in the pack may show substantial variation throughout the year, with reproduction status, which generally peaks in late spring (Packard 2003), being the main factor influencing such variation, with HRs reducing in size as individuals stay closer to the den (Kusak et al. 2005, Jedrzejewski et al. 2007, Roffler and Gregovich 2018. Once the pups gain mobility in late summer, HRs usually increase once again, reaching their maximum size in winter (Ciucci et al. 1997). ...
Home range and habitat selection of wolves recolonising central European human‐dominated landscapes
Article
Full-text available
  • Jun 2024
Decades of persecution has resulted in the long‐term absence of grey wolves Canis lupus from most European countries. However, recent changes in both legislation and public attitudes toward wolves has eased the pressure, allowing wolves to rapidly re‐establish territories in their previous central European habitats over the last 20 years. Unfortunately, these habitats are now heavily altered by humans. Understanding the spatial ecology of wolves in such highly modified environments is crucial, given the high potential for conflict and the need to reconcile their return with multiple human concerns. We equipped 20 wolves, originating from seven packs in six central European regions, with GPS collars, allowing us to calculate monthly average home range sizes for 14 of the animals of 213.3 km² using autocorrelated kernel density estimation. We then used ESA WorldCover data to assess the mosaic of available habitats used within each home range. Our data confirmed a general seasonal pattern for breeding individuals, with smaller apparent home ranges during the reproduction phase, and no specific pattern for non‐breeders. Predictably, our wolves showed a general preference for remote areas, and especially forests, though some wolves within military training areas also showed a broader preference for grassland, possibly influenced by local land use and high availability of prey. Our results provide a comprehensive insight into the ecology of wolves during their re‐colonisation of central Europe. Though wolves are spreading relatively quickly across central European landscapes, their permanent reoccupation remains uncertain due to conflicts with the human population. To secure the restoration of European wolf populations, further robust biological data, including data on spatial ecology, will be needed to clearly identify any management implications.
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... In Mexico, females usually frequent steep sites with vegetation cover that probably provides them with resting places with privacy and the ease of hiding their burrows and cubs (Packard 2003, Trapp 2004. Males, for their part, tend to move greater distances through open terrain with moderate relief , Lara-Díaz et al. 2022. ...
The importance of livestock in the diet of Mexican wolf Canis lupus baileyi in northwestern Mexico
Article
Full-text available
  • Nov 2024
The reintroduction of the Mexican wolf in the wilds of northwestern Mexico has allowed us to address its trophic ecology and elucidate conflicts with livestock producers: their main mortality factor. Our objective was to determine the feeding habits of wolves in Mexico, as well as the quantity and frequency of livestock predation in relation to seasonal and individual variables, through the analysis of genetically identified scats. During 2012–2022 we collected 1171 Mexican wolf scats. We extracted and sequenced DNAm and identified individuals and their sex using microsatellite analysis. We washed the scat and separated the undigested components for taxonomic identification. We estimated the frequency of prey items, the biomass it contributed to the diet, and compared prey consumption between sexes and between the birth and dispersal seasons. We constructed generalized linear models to identify the relationship between livestock presence in the diet and dietary prey richness with respect to environmental and individual variables. We identified 68 wolves that had consumed 30 species of vertebrates. Of these, white‐tailed deer (36.12%), diversionary feeding (22.79%), and cattle (25.56%) had the highest contribution to biomass. The ingestion of items was independent of the sex of the wolves but was dependent on the season. The presence of deer and diversionary feeding decreased the likelihood of cattle being ingested but also decreased the richness of items of wild species in the wolf diet. Wolves in northwestern Mexico fed mainly on large prey available in the reintroduction area, including livestock. As wolves consume livestock, it increases the risk of retaliatory actions from ranchers. Our results serve as a basis for the implementation of strategies to reduce human–wolf conflicts and set a baseline for coexistence in northwestern Mexico.
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... The dimensions of the body and skull of mammalian predators have a profound impact on various characteristics, such as their basal metabolism, communication, social hierarchy, reproductive characteristics, and reproductive success (Geffen et al. 1996;Harrington and Asa 2003;Packard 2003;MacNulty et al. 2009). ...
Sexual dimorphism in the Hungarian golden jackal population: analysing body and skull size and shape
Article
Full-text available
  • Jul 2024
Defining species within the Canidae family is challenging due to morphological convergence, behavioral plasticity, traditional taxonomic limitations, and possible hybridisation. This challenge is timely with the recent population and range expansion of the golden jackal ( Canis aureus ). Exploring their morphological data and sexual dimorphism is essential for identifying factors driving their success in new habitats. The proven hybridization of golden jackals with dogs and wolves may affect species description, population dynamics, and genetic diversity, impacting conservation strategies. This study, for the first time, conducts a morphometric analysis of golden jackals in Somogy County, Hungary, to prove sexual size dimorphism (SSD) in body and skull and sexual shape dimorphism (SShD) in skull across juvenile and adult age groups. 719 golden jackals (362 females and 357 males) were collected between January 2021 and January 2023. Descriptive statistics revealed significant SSD in body and skull measurements among both age groups, with males generally larger than females, particularly in body mass (11.72% in juveniles and 13.37% in adults). Most skull dimensions differed significantly between sexes and age groups, except for foramen magnum height, foramen magnum width, and postorbital breadth among juveniles and foramen magnum height and postorbital breadth among adults. We used principal component analyses (PCA) on raw dimension data and the log shape ratio method to extract shape information. Linear discriminant analysis (LDA) explored skull SShD between sexes. Notably, our study achieved over 71% accuracy in sex classification, illustrating the clear presence of SShD of the skull in golden jackals across both age groups. Our study provides a comprehensive database of golden jackals in the overpopulated Hungarian habitat, which will be helpful for further research on ecology, behavior, and conservation management.
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Helpers show plasticity in their responses to breeder turnover
Article
Full-text available
  • Jun 2024
Nonbreeding helpers can greatly improve the survival of young and the reproductive fitness of breeders in many cooperatively breeding species. Breeder turnover, in turn, can have profound effects on dispersal decisions made by helpers. Despite its importance in explaining group size and predicting the population demography of cooperative breeders, our current understanding of how individual traits influence animal behavior after disruptions to social structure is incomplete particularly for terrestrial mammals. We used 12 yr of genetic sampling and group pedigrees of gray wolves (Canis lupus) in Idaho, USA, to ask questions about how breeder turnover affected the apparent decisions by mature helpers (≥2-yr-old) to stay or leave a group over a 1-yr time interval. We found that helpers showed plasticity in their responses to breeder turnover. Most notably, helpers varied by sex and appeared to base dispersal decisions on the sex of the breeder that was lost as well. Male and female helpers stayed in a group slightly more often when there was breeder turnover of the same sex, although males that stayed were often recent adoptees in the group. Males, however, appeared to remain in a group less often when there was breeding female turnover likely because such vacancies were typically filled by related females from the males' natal group (i.e. inbreeding avoidance). We show that helpers exploit instability in the breeding pair to secure future breeding opportunities for themselves. The confluence of breeder turnover, helper sex, and dispersal and breeding strategies merge to influence group composition in gray wolves.
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Many faces of dominance: the manifestation of cohabiting companion dogs' rank in competitive and non-competitive scenarios
Article
Full-text available
  • Mar 2024
  • ANIM COGN
There are indications that companion dogs of multi-dog households form a hierarchy, maintained by formal and agonistic dominance. Although it was found that the behaviour of dogs depends on their rank in several contexts, so far, the assessment of their rank itself has been based on owner-completed questionnaires. With this research we endeavoured to find associations between rank scores from the Dog Rank Assessment Questionnaire (DRA-Q) and cohabiting dogs’ behaviour in a competitive test (Toy Possession test—32 dog pairs) and a non-competitive, citizen science scenario (Greeting test—20 dog pairs). Based on the grabbing the toy first and keeping the toy at the end variables, the dogs’ rank score provided a reliable indication of the dominant and subordinate dogs’ behaviour in the Toy Possession test. Similarly, the occurrence of dominant and submissive behaviours in the Greeting Test showed a good match with the agonistic and leadership subscores of the composite rank score from the DRA-Q. Our results provide a pioneering case for validating a questionnaire-based rank scoring method with biologically meaningful behavioural tests in the case of companion dogs. The finer analysis of the results highlighted that in the case of a multi-question scoring system, some components might provide more effective prediction of the dogs’ rank-related behaviour in some situations, while other components are more relevant in others, with traits related to agonistic dominance having relevance across contexts. Supplementary Information The online version contains supplementary material available at 10.1007/s10071-024-01842-0.
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