The Effects of Breeder Loss on Wolves

Abstract Managers of recovering wolf (Canis lupus) populations require knowledge regarding the potential impacts caused by the loss of territorial, breeding wolves when devising plans that aim to balance population goals with human concerns. Although ecologists have studied wolves extensively, we lack an understanding of this phenomenon as published records are sparse. Therefore, we pooled data (n = 134 cases) on 148 territorial breeding wolves (75 M and 73 F) from our research and published accounts to assess the impacts of breeder loss on wolf pup survival, reproduction, and territorial social groups. In 58 of 71 cases (84%), ≥1 pup survived, and the number or sex of remaining breeders (including multiple breeders) did not influence pup survival. Pups survived more frequently in groups of ≥6 wolves (90%) compared with smaller groups (68%). Auxiliary nonbreeders benefited pup survival, with pups surviving in 92% of cases where auxiliaries were present and 64% where they were absent. Logistic regression analysis indicated that the number of adult-sized wolves remaining after breeder loss, along with pup age, had the greatest influence on pup survival. Territorial wolves reproduced the following season in 47% of cases, and a greater proportion reproduced where one breeder had to be replaced (56%) versus cases where both breeders had to be replaced (9%). Group size was greater for wolves that reproduced the following season compared with those that did not reproduce. Large recolonizing (>75 wolves) and saturated wolf populations had similar times to breeder replacement and next reproduction, which was about half that for small recolonizing (≤75 wolves) populations. We found inverse relationships between recolonizing population size and time to breeder replacement (r= —0.37) and time to next reproduction (r= —0.36). Time to breeder replacement correlated strongly with time to next reproduction (r=0.97). Wolf social groups dissolved and abandoned their territories subsequent to breeder loss in 38% of cases. Where groups dissolved, wolves reestablished territories in 53% of cases, and neighboring wolves usurped territories in an additional 21% of cases. Fewer groups dissolved where breeders remained (26%) versus cases where breeders were absent (85%). Group size after breeder loss was smaller where groups dissolved versus cases where groups did not dissolve. To minimize negative impacts, we recommend that managers of recolonizing wolf populations limit lethal control to solitary individuals or territorial pairs where possible, because selective removal of pack members can be difficult. When reproductive packs are to be managed, we recommend that managers only remove wolves from reproductive packs when pups are ≥6 months old and packs contain ≥6 members (including ≥3 ad-sized wolves). Ideally, such packs should be close to neighboring packs and occur within larger (≥75 wolves) recolonizing populations.     

Estimation of Successful Breeding Pairs for Wolves in the Northern Rocky Mountains,USA

Abstract: Under the Endangered Species Act, documenting recovery and federally mandated population levels of wolves (Canis lupus) in the Northern Rocky Mountains (NRM) requires monitoring wolf packs that successfully recruit young. United States Fish and Wildlife Service regulations define successful breeding pairs as packs estimated to contain an adult male and female, accompanied by ≥2 pups on 31 December of a given year. Monitoring successful breeding pairs will become more difficult following proposed delisting of NRM wolves; alternatives to historically intensive methods, appropriate to the different ecological and regulatory context following delisting, are required. Because pack size is easier to monitor than pack composition, we estimated probability a pack would contain a successful breeding pair based on its size for wolf populations inhabiting 6 areas in the NRM. We also evaluated the extent to which differences in demography of wolves and levels of human-caused mortality among the areas influenced the probability of packs of different sizes would contain successful breeding pairs. Probability curves differed among analysis areas, depending primarily on levels of human-caused mortality, secondarily on annual population growth rate, and little on annual population density. Probabilities that packs contained successful breeding pairs were more uniformly distributed across pack sizes in areas with low levels of human mortality and stable populations. Large packs in areas with high levels of human-caused mortality and high annual growth rates had relatively high probabilities of containing breeding pairs whereas those for small packs were relatively low. Our approach can be used by managers to estimate number of successful breeding pairs in a population where number of packs and their sizes are known. Following delisting of NRM wolves, human-caused mortality is likely to increase, resulting in more small packs with low probabilities of containing breeding pairs. Differing contributions of packs to wolf population growth based on their size suggests monitoring successful breeding pairs will provide more accurate insights into population dynamics of wolves than will monitoring number of packs or individuals only.     

CHORUS HOWLING BY WOLVES: ACOUSTIC STRUCTURE,PACK SIZE AND THE BEAU GESTE EFFECT

ABSTRACT

A variety of structural parameters were measured from wolf choruses recorded in the Superior National Forest, Minnesota, USA. Mean duration of 60s did not vary with pack size or composition. Packs replied to simulated howling after an average of 40s, often interrupting the stimulus howls. Choruses began with simply-structured howls, which became increasingly modulated as the chorus progressed. Little difference in mean fundamental frequency or other howl parameters was found among the choruses from packs of various sizes and compositions. In particular, choruses produced by single adult pairs did not differ from those of larger packs accompanied by pups. The lack of relationship between chorus parameters and pack size or composition indicates there is little useful information concerning a pack's size to be found in its chorus howling.

The observation that chorus howling by adult pairs is often perceived as that of larger groups with pups suggests that chorus structure has evolved to exaggerate the apparent size of the pack, especially those newly-established or otherwise reduced in number. If so, wolf howling choruses may represent a mammalian example of the Beau Geste effect, made particularly viable because of the relative immunity of the signal to probing.     

Space and Habitat Use by a Red Wolf Pack and Their Pups During Pup-Rearing

ABSTRACT During summer 2005, we evaluated space and habitat use by red wolves (Canis rufus) during pup-rearing. Home-range sizes for red wolves (3 ad, 3 juv, and 4 pups) varied from 3.48 km² to 12.24 km². Red wolves selected agricultural fields over adjacent forested areas and used less space during pup-rearing than we expected based on prior knowledge of the species. Attending pack members rarely left pups alone, pack members shared pup-rearing duties, and male red wolves appeared to play a significant role in pup-rearing.     

The process of a wolf pack splitting in Białowieża Primeval Forest,Poland

In 1998, the pack of 7 wolvesCanis lupus Linnaeus, 1758, radio-tracked in Białowieża Primeval Forest, East Poland, split into 2 packs (2 and 5 wolves), when an 8-year-old alpha female ceased breeding. The two sister-packs subdivided their original territory, but their ranges overlapped extensively (49%) for one year after the split, except for May-June, when both new packs reared pups. We propose that food related factors could have been the ultimate cause of splitting of a large pack. In European temperate forests, pack size of 5–6 wolves is optimal for the consumption of the red deerCervus elaphus.     

Some Aspects of the Population Ecology of Wolves, Alaska

Information on Alaskan wolf populations was obtained from examinationof bounty records, 4,150 wolf radii and ulnae, 1,262 wolf carcasses,and from observations of wolves inhabiting an area of 20,000square miles where wolves were protected. Pregnant adult female wolves averaged 6.5 fetuses; two-year-oldfemales averaged 5.3 fetuses;female pups were not sexually mature. In Alaska, wolves conceive from late February through earlyApril but most females breed in March. Multiparous females breedearlier than first breeders. Multiparous females produce anaverage of 7.3 ova and 6.5 implanted fetuses. The loss of ovafrom ovulation to implantation is significant. Multiparous femalesproduce more ova than first breeders; the difference is highlysignificant. Mortality of pups rather than the lack of initial productionof pups is believed to be the reason for the observed variationsin the proportion of pups in wolf populations. Wolf packs includemembers in all categories of sex and age during the breedingseason. Size of the pack is an indicator of abundance. Wolves in an area where they were protected increased at anaverage rate of 20–30% per year during an 11-year period.     

Differences in winter activity,courtship, and social behavior of two captive family groups of Mexican wolves (Canis lupus baileyi)

The purpose of this study was to determine differences in activity patterns and social behavior of two groups of endangered Mexican wolves maintained at two quite different facilities and to determine some of the variables that should be considered when making specific behavioral comparisons among wolves in this binational captive breeding program. Quantitative measurements of an Activity Index and social behaviors were obtained for three individuals in each pack. Within each age/sex category, activity, aggression, and play were more frequent in the pack at a zoo facility, compared to the pack at a field station facility. Frequency of courtship interactions and scent marking were significantly higher in the field station pack. The packs were similar in the frequency of active submission, but differed significantly in the pattern of this behavior. Given the large number of interacting variables and small number of individuals in this study, we recommend caution in generalizing results to other packs or facilities. Zoo Biol 16:435–443, 1997. © 1997 Wiley-Liss, Inc.     

Non-invasive acoustic detection of wolves

Monitoring wolves (Canis lupus) is a difficult and often expensive task due to high mobility, pack dynamic, shyness and nocturnal activity of this species. Wolves communicate acoustically through howling, within pack and with packs of the neighbourhood. A wolf howl is a low-frequency vocalization that can be transmitted over long distances and thus it can be used for monitoring. Elicited howling survey is a current method to monitor wolves in different areas all over the world. Elicited howling, however, may be invasive to residential wolf packs and could create possible negative reactions from the human population. Here we show that it is possible to detect wolves by recording spontaneous howling events. We measured the sound pressure level of wolf howls by captive individuals and we further found that elicited howling may be recorded and clearly identified up to a distance of 3 km. We finally conducted a non-invasive acoustic detection of wolves in a free-ranging population. The use of passive sound recorders may provide a powerful non-invasive tool for future wolf monitoring and could help to establish sustainable management plans for this species.     

Severe inbreeding depression in a wild wolf (Canis lupus) population

The difficulty of obtaining pedigrees for wild populations has hampered the possibility of demonstrating inbreeding depression in nature. In a small, naturally restored, wild population of grey wolves in Scandinavia, founded in 1983, we constructed a pedigree for 24 of the 28 breeding pairs established in the period 1983-2002. Ancestry for the breeding animals was determined through a combination of field data (snow tracking and radio telemetry) and DNA microsatellite analysis. The population was founded by only three individuals. The inbreeding coefficient F varied between 0.00 and 0.41 for wolves born during the study period. The number of surviving pups per litter during their first winter after birth was strongly correlated with inbreeding coefficients of pups (R2=0.39, p<0.001). This inbreeding depression was recalculated to match standard estimates of lethal equivalents (2B), corresponding to 6.04 (2.58-9.48, 95% CI) litter-size-reducing equivalents in this wolf population.     

BIOACOUSTIC PUBLICATIONS, 1987

The use of amplitudes to identify individuals has historically been ignored by bioacoustic researchers due to problems of attenuation. However, recent studies have shown that amplitudes encode identity in a variety of mammal species. Previously, individuality has been demonstrated in both fundamental frequency (F₀) and amplitude changes of captive Eastern wolf (Canis lupus lycaon) howls with 100% accuracy where attenuation of amplitude due to distance was controlled in a captive environment. In this study, we aim to determine whether both fundamental frequency and amplitude data collected from vocalizations of wild wolves recorded over unknown distances, in variable conditions and with different recording equipment, can still encode identity. We used a bespoke code, developed in Matlab, to extract simple scalar variables from 67 high-quality solo howls from 10 wild individuals and 112 chorus howls from another 109 individuals, including lower quality howls with wind or water noise. Principal component analysis (PCA) was carried out on the fundamental frequency and normalized amplitude of harmonic 1, yielding histogram-derived PCA values on which discriminant function analysis was applied. An accuracy of 100% was achieved when assigning solo howls to individuals, and for the chorus howls a best accuracy of 97.4% was achieved. We suggest that individual recognition using our new extraction and analysis methods involving fundamental frequency and amplitudes together can identify wild wolves with high accuracy, and that this method should be applied to surveys of individuals in capture–mark–recapture and presence–absence studies of canid species.     

Wolf monitoring in Scandinavia: evaluating counts of packs and reproduction events

Large carnivores are elusive and use large areas, which causes monitoring to be challenging and costly. Moreover, management to reduce conflicts and simultaneously ensure long-term population viability require precise population estimates. In Scandinavia, the monitoring of wolves (Canis lupus) is primarily based on counting packs, identifying reproduction, and genetically identifying territorial wolves from noninvasive DNA samples. We assessed the reliability of wolf monitoring in Scandinavia by estimating the detectability of territorial pairs, packs, and reproduction. Our data, comprising snow-tracking data and DNA-identified individuals from 2005–2016, covered 11 consecutive winter monitoring seasons (Oct–Mar). Among 343 cases where we identified a wolf pack, territorial wolves were also detected in the same area during the previous season in 323 (94.2%) cases. In only 6 of the remaining 20 cases, there was no prior knowledge of territorial wolves in the area. Among the 328 detected reproduction events (litter born to a pack), we detected 97% during the monitoring period and identified the rest ≥1 year later from kinship assessments of all DNA-detected individuals. These results suggest that we failed to detect only few packs with reproduction events during the monitoring season that followed breeding. Yearly monitoring of territorial individuals and continuous updates of the pedigree allowed us to retrospectively identify reproduction events and packs that were not identified earlier.     

Howling activity of free-ranging wolves (<Emphasis Type="Italic">Canis lupus</Emphasis>) in the Białowieża Primeval Forest and the Western Beskidy Mountains (Poland)

We investigated spontaneous howling by radio-collared wolves Canis lupus inhabiting the Białowieża Primeval Forest (BPF), eastern Poland, and elicited howling behavior in wolves of the BPF and the Western Beskidy Mountains, southern Poland. Over half (58%) of all spontaneous howls recorded throughout a year occurred in the period from July to October, with a peak in August. The daily pattern of vocal activity by wolves was characterised by a peak between 1800 and 0000 hours, which coincided with the first (dusk) peak of wolf mobility. Wolves howled from the core areas of their territories and not from the peripheries. Howls served as communication between temporarily separated pack mates (43% of cases), after re-union (18%) and before setting out for a hunt (22%). Very few spontaneous howls (2%) were targeted at a neighbouring pack. Wolves responded to human-simulated howling in June–September, with a peak in August (reply rate: 39%). The duration of elicited howling increased significantly with group size: howls by single wolves or pairs lasted, on average, 34–40 s, whereas those of five to seven wolves (including pups) had an average duration of 67–95 s, with a maximum length of nearly 4 min. In the populations of Polish wolves studied here, spontaneous howling served primarily for intra-pack communication. Nonetheless, the high reply rate to howling simulation showed that – if necessary – wolves readily advertised their presence in a territory to strangers.     

Movements and Home Sites of Timber Wolves in Algonquin Park

Ecological studies of timber wolves (Canis lupus) in a forestedenvironment have always been difficult to undertake in the past,particularly during the summer months, because of the lack ofa suitable technique. In 1960 Pimlott devised a technique whichemployed the use of broadcast wolf howls in locating wolvesin the field. This report reviews the success of this techniquein studying the movements of two packs of wolves, and theiruse of home sites in Algonquin Park, Ontario. Wolves responded by howling a total of 476 occasions or approximately13% of the occasions that broadcast howls were given. Humanimitations were more successful than tape-recordings in inducingresponses. Wolves responded at any time of day, but dusk wasthe most favorable period. They also responded more frequentlyin July and August than in May and June. Two types of home sites were found: the den site, occupied duringthe early life of the pups, and the rendezvous site, a placeoccupied by wolves during later development of pups. All ofhe home sites were adjacent to some immediate source of water.The movement of wolves appeared also to be concentrated alongthe water courses. The locations of the home sites and the evidence obtained fromhowling responses, tracks, and scats suggested that the summerrange comprised a minimum area of eight square miles. The samerange was utilized by a pack in 1961 and 1963.     

Response of Wolves to Experimental Disturbance at Homesites

ABSTRACT Events during the denning period (parturition to first autumn) often determine the reproductive success of wolves (Canis lupus). Consequently, there is concern about the potential adverse effects of human-caused disturbance at wolf den and rendezvous sites (homesites), but relatively little information on this subject is available. We conducted standardized experimental disturbance treatments at 12 unique wolf homesites in the Northwest Territories, Canada, during summers 2002 and 2003. The treatment consisted of an intruder approaching a homesite once per day for 3 consecutive days and recording behavioral responses, response distance, and response intensity of wolves. We counted pups and estimated their ages prior to the initial treatment at each site. Adult wolves moved pups at 3 of 6 treated homesites in each year. The amount and type of known human activity within a pack's home range did not influence whether adults moved pups in response to the treatment. The response intensity of wolves to the treatment was inversely related to the amount of human activity near a homesite. There was no relationship between the distance at which wolves responded to the intruder and the amount or type of human activity. There was a positive relationship between increasing age of pups and their relocation in response to the treatment. Reproductive success was not influenced by the treatment or by the amount and type of human activity. Treated sites were used by wolves the following year in the same proportion as untreated sites. It appears that pups are most vulnerable early in the year when less mobile; therefore, managers should consider age of pups before human activity at or near wolf homesites occurs.     

Evidence of wolf dispersal in anthropogenic habitats of the Polish Carpathian Mountains

In the course of their maturation, most young wolves leave their natal pack and disperse in search for mating partners, improved food availability and new territories. We investigated whether this dispersal is affected by anthropogenic infrastructure in a 5,000 km2 area of the eastern region of the Polish Carpathian Mountains occupied by wolves. A radio-collared male wolf covered 230 km while dispersing through forested hills and densely populated valleys. To test if such dispersal is common in the population we analysed by microsatellite genotyping 39 samples taken from live-trapped wolves or wolves found dead in the study area. Although the obtained genotypes were assigned to different clusters in Bayesian tests, we could not ascribe this structure to landscape features, but rather to shared ancestry of wolf individuals found in distant locations. Moreover, we could not detect a spatial genetic structure in the wolf population, indicating a random occurrence of genotypes within the study area. Observation of the dispersing wolf and the absence of spatial genetic structure imply that wolves are still able to roam the entire area despite high densities of roads and a dense human population. Thus, we concluded that the existing anthropogenic infrastructure does not restrict wolf dispersal in the area and the studied wolves represent a coherent part of the Polish Carpathian wolf population.     

Numbers, Turnover, and Social Structure of the Isle Royale Wolf Population

Observations of wolves on Isle Royale are reported for 1961–66,with interpretations including the earlier 3-year period describedby Mech (1966). On this 210-square-mile island the fully protectedwolf population varied from approximately 22 to 28 in midwinter.The major and minor foods were moose and beaver, respectively. The main pack varied in number between 11 and 22 with aboutthree breeding pairs believed present. The population remainedrelatively stable; mating occurred every winter; and adult mortalityappeared to be low. High mortality among pups seemed to be thepoint of population control. Socio-economic factors may havecontrolled the size of the large pack. Availability of foodduring the period of parturition and rearing probably was criticalto survival of young. Recruitment of young appeared to take place in years of highproduction of moose calves. Numbers in the large pack probablywere curtailed through the progressive exclusion of aged andsocially subordinate individuals. Under harassment these animalsseparated and became pack-following scavengers, then probablytrue loners ranging outside the area used by the pack. Smalleraggregations of two or three non-breeders were seen each winter,as were the loners, some of which appeared thin and weak. The only known breeding outside the big pack was in a groupof five present in the winter of 1965. This group was probablya family unit which separated from the main pack. A year laterthe male had disappeared, and remains of a pup, probably theirs,were found. In the winter of 1966 the alpha male of the largepack became lame and apparently was killed. This backgroundappears favorable for further changes in social organization.     

Den shifting by wolves in semi-wild landscapes in the Deccan Plateau, Maharashtra, India

Reproductive success is crucial for the survival and persistence of any species. The Deccan Biogeographic Zone of India is the stronghold of a population of the Indian wolf Canis lupus pallipes . Gaining a better understanding of the den-use pattern of wolves in different areas in this zone is thus vital for their conservation and management. The wolves excavated multiple dens in our study sites and kept shifting their litters among them. Major disturbance factors around denning sites were active stone quarries, traffic, crop harvesting and livestock movement. One wolf pack used 14 dens in four breeding seasons. Discriminant function analysis indicated that den shifting by wolves was not entirely governed by disturbance levels at den sites. Increasing age of pups was one of the main factors associated with den shifting rather than the magnitude of disturbance ( χ ²=34.26, d.f.=12, P <0.001). Tolerance to disturbance around dens during the early stages of pup development was negatively correlated ( r =−0.519, P <0.05) with availability of water and age of pups ( r =−0.613, P <0.01). Excavation of multiple dens by wolves was apparently related to den shifting, which seemingly is a survival strategy of wolves in these semi-wild human-dominated landscapes in the Deccan Biogeographic Zone. Principal components analysis indicated that during the initial stages of pup development, nature of the land, den orientation and distance of the den from roads were important cues in addition to age category of pups for den shifting. The analysis also suggested that factors such as distance of the new den where pups are to be transferred, distance from water source and availability of fox Vulpes bengalensis holes for den use were also important.     

Estimation of pack density in grey wolf (<Emphasis Type="Italic">Canis lupus</Emphasis>) by applying spatially explicit capture-recapture models to camera trap data supported by genetic monitoring

Background

Density estimation is a key issue in wildlife management but is particularly challenging and labour-intensive for elusive species. Recently developed approaches based on remotely collected data and capture-recapture models, though representing a valid alternative to more traditional methods, have found little application to species with limited morphological variation. We implemented a camera trap capture-recapture study to survey wolf packs in a 560-km² area of Central Italy. Individual recognition of focal animals (alpha) in the packs was possible by relying on morphological and behavioural traits and was validated by non-invasive genotyping and inter-observer agreement tests. Two types (Bayesian and likelihood-based) of spatially explicit capture-recapture (SCR) models were fitted on wolf pack capture histories, thus obtaining an estimation of pack density in the area.

Results

In two sessions of camera trapping surveys (2014 and 2015), we detected a maximum of 12 wolf packs. A Bayesian model implementing a half-normal detection function without a trap-specific response provided the most robust result, corresponding to a density of 1.21?±?0.27 packs/100 km² in 2015. Average pack size varied from 3.40 (summer 2014, excluding pups and lone-transient wolves) to 4.17 (late winter-spring 2015, excluding lone-transient wolves).

Conclusions

We applied for the first time a camera-based SCR approach in wolves, providing the first robust estimate of wolf pack density for an area of Italy. We showed that this method is applicable to wolves under the following conditions: i) the existence of sufficient phenotypic/behavioural variation and the recognition of focal individuals (i.e. alpha, verified by non-invasive genotyping); ii) the investigated area is sufficiently large to include a minimum number of packs (ideally 10); iii) a pilot study is carried out to pursue an adequate sampling design and to train operators on individual wolf recognition. We believe that replicating this approach in other areas can allow for an assessment of density variation across the wolf range and would provide a reliable reference parameter for ecological studies.

     

Social living mitigates the costs of a chronic illness in a cooperative carnivore

Infection risk is assumed to increase with social group size, and thus be a cost of group living. We assess infection risk and costs with respect to group size using data from an epidemic of sarcoptic mange (Sarcoptes scabiei) among grey wolves (Canis lupus). We demonstrate that group size does not predict infection risk and that individual costs of infection, in terms of reduced survival, can be entirely offset by having sufficient numbers of pack‐mates. Infected individuals experience increased mortality hazards with increasing proportions of infected pack‐mates, but healthy individuals remain unaffected. The social support of group hunting and territory defence are two possible mechanisms mediating infection costs. This is likely a common phenomenon among other social species and chronic infections, but difficult to detect in systems where infection status cannot be measured continuously over time.     

Direct Estimation of Early Survival and Movements in Eastern Wolf Pups

Abstract: Determining juvenile survival and recruitment rates is essential to assess status and viability of animal populations. Currently, the demographic attributes of juvenile carnivores, specifically wolves (Canis lycaon), are poorly known but of considerable conservation interest. We measured survival and dispersal rates for 51 juvenile (age 3.5–31 weeks) wolves in Algonquin Provincial Park, Canada, from 2004 to 2005, using implantable very high frequency transmitters. Monthly pup survival was high (0.970, 95% CI = 0.951–0.990) and constant from June to November, and most pup mortality was from natural causes. Pups dispersed as early as age 15 weeks, and monthly dispersal rates were high for young pups (min. = 0.008, 95% CI = 0.000–0.019; max. = 0.030, 95% CI = 0.010–0.050). We failed to detect any influence of pack or litter size on pup survival or probability of dispersal. Radiotelemetry offers an individual-based monitoring technique capable of providing direct assessment of wolf pup survival and movements, with rigorous estimation of survival and dispersal rates and quantification of cause-specific mortality.