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Reproduction biology in grey wolves Canis lupus in Belarus: Common beliefs versus reality
Abstract
This scientific monograph gives a detailed information about reproduction biology in the grey wolf Canis lupus in Belarus. This topic includes the wolf breeding (mating and denning) behavior, fertility of the species and mortality of its pups. The initial material was not collected occasionally from wolf hunters and wolf pup searchers, but mainly gained by authors first-hand according to a well-set research design and long-term. By analyzing the gathered data, we became convinced that in the wolf reproduction biology there are more exceptions than rules. Therefore, the standard patterns of reproduction biology in wolves that are wide-spread in the published literature about the species we call as common beliefs that are given versus the wolf reality that we have found in Belarus. Concerning the non-standard features in the wolf reproduction biology, we revealed that multiple breeding in wolf pack is a common phenomenon, breeding of yearling females and wolf-dog hybridization were found to be irrespective the food base and strongly depending on the species population density i.e. they are reproduction regulations. Wolf pup mortality was investigated and the crucial role of deliberate predation of lynxes on wolf pups was revealed.
... As such, the size of a typical wolf pack at peak size consists of two breeders, a few yearlings (< 2 year old) and pups (< 1 year old). Occasionally, there are multiple breeders in a pack, which typically happens when the resident male is replaced and daughters of the former male are still pack members (Sidorovich and Rotenko 2019;Pacheco et al. 2024). Depending on the timing of mortality, birth, and dispersal, the size of a single pack can vary within a year from 3 to > 15 individuals, typically including no more than two adult individuals. ...
Molecular methods are routinely used to estimate the effective size of populations (Ne). However, underlying model assumptions are frequently violated to an unknown extent. Although simulations can detect sources of bias and help to adjust sampling strategies and analyses methods, additional information from empirical data can also be used to calibrate methods and improve molecular Ne estimation methods. Here, we take advantage of long‐term genetic and ecological monitoring data of the grey wolf (Canis lupus) in Germany, and detailed population genetic studies in Poland, Spain and Portugal to improve Ne estimation strategies in this species, and species with similar life history traits. We first calculated Ne from average lifetime reproductive success and detailed census data from the German population, which served as a baseline to compare to molecular estimates based on linkage disequilibrium and sibship frequency. This yielded a robust Ne/Nc estimation that we used to calibrate molecular estimates of German, Polish and Iberian wolf populations. The linkage disequilibrium method was strongly influenced by spatial genetic structure, much more than the sibship frequency method. When Ne was estimated in local neighbourhoods, both methods yielded comparable results. Estimates of the metapopulation effective size seemed to correspond generally well with the sum of the estimates of local neighbourhoods. Overall, we found that the number of packs is a good proxy of the effective population size. Using this as a rule of thumb, we evaluated for all European wolf populations the Ne 500 indicator and concluded that half of the European wolf populations do not yet fulfil this criterion.
... Op basis van waarden uit Europese literatuur gebruiken we in het default model voor welpen een jaarlijkse overleving van 0.58(Merli et al. (2023) 36 . Sidorovich & Rotenko (2019)37 noemen een veel hogere sterfte, met slechts 27% tot 4% overleving. Oorzaak van deze hoge sterfte is predatie door lynx, iets wat in Nederland nog niet speelt, aangezien de lynx Nederland (nog) niet heeft bereikt. ...
De auteurs hebben een analyse uitgevoerd naar wat kan worden verwacht omtrent de verdere ontwikkeling van de in Nederland voorkomende wolvenpopulatie in groei en ook verdere verspreiding.
Meer specifiek heeft de analyse zich gericht op het verkrijgen van inzicht in:
1. De huidige en toekomstige ontwikkeling van de wolvenpopulatie in Nederland en de factoren die hierop van invloed kunnen zijn (populatie en toekomstperspectief);
2. De huidige en toekomstige ontwikkeling van de verspreiding van de wolf in Nederland en de factoren die hierop van invloed kunnen zijn (verspreiding en toekomstperspectief);
3. De samenhang tussen de in Nederland voorkomende wolven(sub)populatie met de binnen Europa te onderscheiden wolvenpopulaties én de schaal waarop deze gedefinieerd kan worden (samenhang van de populatie).
Als bijlage is een uitgebreide samenvatting in Engels en Nederlands bewschikbaar (zie Supplementary materials)
... This was observed for the Scandinavian and the Finnish wolf populations following a severe bottleneck (Jansson et al., 2012). Within wolves, parent-offspring mating events are known to occur (Sidorovich and Rotenko, 2019) and were documented for the German wolves as well (Jarausch et al., 2021). Hence, while quite good levels of genetic diversity within the German wolves of 2015 (12 years after their first reappearance) were found (Jarausch et al., 2021), high uncertainty about their future genetic diversity remained. ...
Population viability analyses (PVA) are important tools for decision-making and planning of adaptive wildlife management actions. While earlier approaches on individual based PVAs have often been age-based, analyses of species with strong social structure might benefit from a stage-based model approach. In this study, we designed an individual-based and stage-based PVA within the software Vortex. As a case study, we applied our model to the German part of the European wolf population, making use of comprehensive data sets originating from the German monitoring regime including individual genotypes. Genetic diversity and inbreeding were important considerations in our analysis, as they could greatly impact population dynamics. We aimed to assess the pop-ulation's trajectory, extinction risk, and genetic integrity under different scenarios while analyzing factors that could affect its survival. We found that mortality rates at different life cycle stages had varying effects on population growth. Higher mortality rates among pups and dispersers led to negative growth and increased the risk of extinction. In comparison, higher mortality rates among yearlings and subdominant wolves still resulted in positive growth but at a slower rate. Mortality among the breeding individuals within the pack (territorials) had the highest impact on population trajectory. Although the German wolves represent a rather recently founded population, our simulations predicted generally good genetic diversity as long as the population was not held at artificially low numbers. Ultimately, we present a generic, easy-to-use-and-adapt model built within the Vortex environment, that after appropriate modification, calibration and testing could be used within conservation practice and management in collaboration with scientific research. Our study highlights the importance of stage-based modeling for understanding the demographic traits of social species like wolves.
... The role of wolves as agents responsible for archaeofaunal assemblages has generated great interest among researchers [41][42][43][44][45][46][47] . In fact, the large number of taphonomic studies dedicated to wolves, along with the results obtained by studies about wolves' behaviour demonstrating their capacity to accumulate prey remains through carcass transport and scats [37][38][39] , are proof of the important role that this carnivore may have played on archaeological assemblages. This is also confirmed by the archaeological evidence, a good example is the case of Denisova Cave where wolf fossil scats were recovered from different levels and chambers 48 . ...
The study of human subsistence strategies in prehistoric hunter-gatherer communities is essential to understanding the evolution of human behaviour. An important topic of interest is the expansion of dietary breadth, resulting in the procurement of a larger number of small game species. However, to make accurate interpretations of human subsistence, the correct identification of the agents responsible for archaeofaunal assemblages is crucial, and actualistic studies that establish the taphonomic signature of the different predators are indispensable. Despite being one of the most ubiquitous carnivores in prehistoric archaeological sites, the role of wolves (Canis lupus) as agents responsible for small-prey accumulations has never been examined. The aims of this study are to analyse the taphonomic patterns left by wolves on rabbit remains and to put forward a series of criteria that can help distinguish assemblages produced by this carnivore from those accumulated by people or by other predators. Our results reveal that wolves ingest and consume the whole rabbit carcass, with the consequence that all rabbit remains accumulated by wolves come from the scats. The referential framework provided in this study will make it possible to discriminate wolves as agents of fossil rabbit accumulations.
Predation, one of the most dramatic interactions in animals' lives, has long fascinated ecologists.
This volume presents carnivores, raptors and their prey in the complicated net of interrelationships, and shows them against the background of their biotic and abiotic settings. It is based on long-term research conducted in the best preserved woodland of Europe's temperate zone. The role of predation, whether limiting or regulating prey (ungulate, rodent, shrew, bird, and amphibian) populations, is quantified and compared to parts played by other factors: climate, food resources for prey, and availability of other potential resources for predators.
We conducted a 15-year study (1999–2014) of wolf denning behavior in Naliboki Forest, Belarus, to determine the extent to which some external factors (rain, mosquitoes, human disturbances) and litter size are negatively correlated with pup survival until autumn.
Information is sparse about aspects of female wolf (Canis lupus) breeding in the wild, including age of first reproduction, mean age of primiparity, generation time, and proportion of each age that breeds in any given year. We studied these subjects in 86 wolves (113 captures) in the Superior National Forest (SNF), Minnesota (MN), during 1972–2013 where wolves were legally protected for most of the period, and in 159 harvested wolves from throughout MN wolf range during 2012–2014. Breeding status of SNF wolves were assessed via nipple measurements, and wolves from throughout MN wolf range, by placental scars. In the SNF, proportions of currently breeding females (those breeding in the year sampled) ranged from 19% at age 2 to 80% at age 5, and from throughout wolf range, from 33% at age 2 to 100% at age 7. Excluding pups and yearlings, only 33% to 36% of SNF females and 58% of females from throughout MN wolf range bred in any given year. Generation time for SNF wolves was 4.3 years and for MN wolf range, 4.7 years. These findings will be useful in modeling wolf population dynamics and in wolf genetic and dog-domestication studies.
This text focuses on the principles and methods of using growth layers formed in teeth and bones of mammals to make a judgement on essential traits of the animal's life history. In nearly all mammalian species, including man, the age of individuals can be determined from the number of growth layers and, at least in some of them, it is possible to estimate the season of an animal's birth and death, age of sexual maturation, periodicity of reproduction, certain feeding habits and other aspects of the individual's biology. It is also possible, from tooth-enamel analysis, to assess doses of radiation accumulated by animals and human beings during their lifetime.;This book is intended for zoologists, wild-game biologists and zoo archaeologists, but some of the sections could also be of interest for anthropologists, radioecologists and conservation biologists.
European wolves (Canis lupus) show population genetic structure in the absence of geographic barriers, and across relatively short distances for this highly mobile species. Additional information on the location of and divergence between population clusters is required, particularly because wolves are currently recolonizing parts of Europe. We evaluated genetic structure in 177 wolves from 11 countries using over 67K single nucleotide polymorphism (SNP) loci. The results supported previous findings of an isolated Italian population with lower genetic diversity than that observed across other areas of Europe. Wolves from the remaining countries were primarily structured in a north-south axis, with Croatia, Bulgaria, and Greece (Dinaric-Balkan) differentiated from northcentral wolves that included individuals from Finland, Latvia, Belarus, Poland and Russia. Carpathian Mountain wolves in central Europe had genotypes intermediate between those identified in northcentral Europe and the Dinaric-Balkan cluster. Overall, individual genotypes from northcentral Europe suggested high levels of admixture. We observed high diversity within Belarus, with wolves from western and northern Belarus representing the two most differentiated groups within northcentral Europe. Our results support the presence of at least three major clusters (Italy, Carpathians, Dinaric-Balkan) in southern and central Europe. Individuals from Croatia also appeared differentiated from wolves in Greece and Bulgaria. Expansion from glacial refugia, adaptation to local environments, and human-related factors such as landscape fragmentation and frequent killing of wolves in some areas may have contributed to the observed patterns. Our findings can help inform conservation management of these apex predators and the ecosystems of which they are part.
Wolves (Canis lupus) (9 females and 2 males from 4 packs), were radio-tracked in a large Polish woodland in Bialowieża Primeval Forest in 1996-1999. Based on 360 days of radio tracking with locations taken at 30- or 15-min intervals, daily movement distances (DMDs) of wolves and their utilization of territories were analyzed. DMDs averaged 22.1 km for females and 27.6 km for males. In reproductive and subadult females, DMDs varied seasonally, with the shortest daily routes in May and the longest in autumn-winter. Little seasonal variation was observed in nonbreeding and unsuccessfully breeding adult females. An adult male covered the longest DMDs in February (mating season). The mean speed of travelling wolves was 2.2 km/h. Wolves' hunting activity affected the length and speed of their movements, both of which were higher before than after a kill was made. With growing abundance of prey, DMDs of wolves became shorter. Snow cover and rainfall had a negligible effect on wolf travel. The mean straight-line distance between consecutive daily locations (SLD) was 4.4 km, i.e., on average, 21% of the actual route covered by wolves. Daily ranges utilized by wolves averaged 21.4 km2, or 9% of the whole territory. Variation in SLDs and daily ranges was shaped predominantly by mating, breeding, and pup rearing. The pattern of territory use by wolves differed between seasons. In spring-summer, their movements concentrated around the breeding den and rendezvous sites, and the areas used on consecutive days overlapped extensively. In autumn-winter, wolves moved widely and utilized their territory in a rotational way, returning to the same parts every 6 days, on average. Rotational use is related to intense patrolling and defense of territory, but may also help wolves to avoid behavioral depression of prey availability.
Relationships of wolves (Canis lupus) and ungulates were studied in the Polish part of Biaowieza Primeval Forest with high densities of prey. The number of wolves ranged from 7 to 19, and the number of packs ranged from 2 to 4. Average densities were 2.3 wolves/ 100 km 2. Red deer (Cervus elaphus) was the main prey of wolves. Roe deer (Capreolus capreolus), wild boar (Sus scrofa), moose (Alces alces), and European bison (Bison bon- asus) were hunted less than expected based on their abundance. Mean mass of ungulates killed by wolves was 55 kg. Prey were consumed quickly, with 57% of kills completely eaten on the 1st day after killing. Average killing rate by wolves was 0.78 ungulate per wolf pack per day (0.14 prey item per wolf per day). Results of this study combined with the data obtained in the Belarussian part of Biaowieza Primeval Forest in 1946-1985 allowed for analysis of dietary response of wolves to changes in densities of ungulates. Wolves showed a response to abundance of red deer. The amount of other ungulates in their diet depended on the densities of red deer. From 1991 to 1996, wolves annually removed 57-105 red deer, 19-38 wild boar, 19-25 roe deer, and 0-2 moose per 100 km 2. Those amounts were equivalent to 9-13% of spring-summer densities of red deer, 4-8% of wild boar, 3-4% of roe deer, and 0-29% of moose. Additionally, hunters annually harvested 131-140 red deer, 44-114 roe deer, 1-7 moose, and 45-142 wild boar per 100 km 2. Effects of predation and harvest by hunters on ungulate mortality were likely additive and caused declines in ungulate populations during our study.
Males comprised 66 percent of wild wolf (Canis lupus) pups from a saturated, high-density wolf range in northeastern Minnesota, possibly reflecting disproportionate conception of males. Packs from areas of lower wolf density in other areas of Minnesota had equal sex ratios of pups or a disproportionate number of female pups. Captive wolves showed a slight preponderance of male pups.
We evaluated the accuracy and precision of tooth wear for aging gray wolves (Canis lupus) from Alaska, Minnesota, and Ontario based on 47 known-age or known-minimum-age skulls. Estimates of age using tooth wear and a commercial cementum annuli-aging service were useful for wolves up to 14 years old. The precision of estimates from cementum annuli was greater than estimates from tooth wear, but tooth wear estimates are more applicable in the field. We tended to overestimate age by 1-2 years and occasionally by 3 or 4 years. The commercial service aged young wolves with cementum annuli to within ±1 year of actual age, but under estimated ages of wolves ≥9 years old by 1-3 years. No differences were detected in tooth wear patterns for wild wolves from Alaska, Minnesota, and Ontario, nor between captive and wild wolves. Tooth wear was not appropriate for aging wolves with an underbite that prevented normal wear or severely broken and missing teeth.