Molecular determination of paternity in a natural population of the multiply mating polygynous lizard Eulamprus heatwolei

We studied the mating system of the southern water skink, Eulamprus heatwolei, during spring and summer (encompassing the breeding season) in a population in southeastern Australia. We examined potential attributes that might influence the mating system and male reproductive success including home range size, physical proximity of adults and body size, and then genotyped all mothers, offspring and potential sires. Home range overlap of both sexes was extensive, with adult females sharing the greatest amount of space with each other and adult males the least amount of space with each other. However, not all adults hold home ranges. We classified approximately one quarter of adult males as home range holders and the rest as 'floaters'. Adult females occupy home ranges more than males, with approximately three-quarters classified as home range holders. Home range ownership is not correlated with body size for either sex, however, male body size is positively correlated with the number of adult female home ranges that his home range overlaps and adult male home ranges are larger than those of females. We used microsatellite genotyping to assign paternities to 55 offspring from 17 litters and then compared this data with our home range and behavioural observations. This species displays extreme levels of multiple paternity given the small mean clutch size of three. Multiple paternity was confirmed in 11 (64.7%) of 17 clutches but three other clutches (for a total of 82.4%) also may display multiple paternity. A total of 30 offspring from 12 litters were assigned to 10 of the 32 genotyped adult males from our study site. Of these 10 adult males, half were home range holders. Five complete clutches and a total of 25 out of the 55 offspring could not be positively assigned to any male surveyed as part of the study and were attributed to floater males or resident males adjacent to our study site that had not been genotyped. While sample sizes are small, neither male home range ownership nor body size is significantly correlated with the number of paternities a male obtained. Our study suggests a polygynous mating system for this species.     

Size and overlap of home range inCalomys musculinus (Muridae: Sigmodontinae)

Home range size and overlap ofCalomys musculinus (Thomas, 1913) was examined in relation to sex and breeding periods. The study was carried out in four 0.25 ha enclosures, in a natural pasture, between October 2002 and July 2003. The four enclosures functioned as independent populations and each was considered a replicate. The capture, mark and recapture method was used. Home range size and overlap inC. musculinus depended on sex and period. Home range size of males was always larger than that of females. Females showed a small degree of intra-sexual home range overlap during the breeding period. In general, male home ranges overlapped with females. We conclude that differences in home range size inC. musculinus is determined by sex and breeding period. Moreover, the degree of inter- and intra-sexual home range overlap during the breeding period suggests that both males and females ofC. musculinus use space differently. Females did not share their home range with other females, while males fully shared it with both sexes, and male spacing is influenced by the distribution of females. A promiscuous-polygynous mating system is suggested forC. musculinus.     

Space use of common genets<Emphasis Type="Italic">Genetta genetta</Emphasis> in a Mediterranean habitat of northeastern Spain: differences between sexes and seasons

The seasonal home range size and spatial relationships of 16 adult genetsGenetta genetta Linnaeus, 1758 (6 males and 10 females) were estimated in a Mediterranean habitat of northeastern Spain. Genets minimum density was estimated as 0.98/km². Mean annual home range was 113.1 ha in males and of 72.0 ha in females. Males had larger home ranges than females in all seasons, but differences were only significant in winter. Home range size changed seasonally and showed a similar pattern in both sexes, with lower values in summer (males — 41.2 ha, females — 29.0 ha) and maximum ones in spring (males — 78.8 ha, females — 56.1 ha). Animals displayed spatial fidelity throughout the year. Core areas (MCP50) represented 27% and 19% of total home range size for males and females, respectively. Resting home ranges (based on locations of inactive animals) were 9 times lower than overall home range size. Individuals of the same sex overlapped less than individuals of different sexes, especially with regard to core areas, which showed almost no overlap. The results obtained suggest that (1) different factors are likely to affect the space use of genets, such as body mass, food abundance and reproductive cycle; (2) genets use space in a heterogeneous way, with areas of greater activity than others within their home range; (3) there was intrasexual segregation with regard to space use.     

Space use in the root vole: basic patterns and variability

Home range characteristics and spatial organization of the root vole Microtus oeconomus were assessed by radio-tracking The aim of this paper is to test the assumption that each microtine species employs a certain modal spatial system around which there is considerable flexibility In the studied population the basic modes of spatial organization were female territoriality and male use of overlapping home ranges Female territories were significantly smaller and less diversified in size than male ranges The size of male home ranges was positively correlated with body mass Each female territory was overlapped by several male ranges, larger male ranges usually overlapped more female territories The above spatial organization suggests that the prevailing mating system in the root vole is promiscuity, and larger males have an access to more receptive females than have small-sized males
Departures from these basic modes were found among young mature females from summer cohort, forming clusters even at low density, and occasionally among females in locally crowded habitat patches, where they held overlapping home ranges A single male usually monopolized such a cluster, excluding other males from the area used by the females     

Woodland fragmentation affects space use of Eurasian red squirrels

When habitats become fragmented, variation in patch size and quality are expected to impose changes on the spacing pattern and social organization of animal populations. General theory predicts different possible responses including shrinking home ranges (fission response), increasing range overlap (fusion) and incorporation of multiple patches in the home range (expansion response) as fragmentation increases. We studied space use and social organization in a metapopulation of red squirrels (Sciurus vulgaris) in 15 woodland fragments differing in size and tree species composition. Home ranges and core areas of males were larger than females, and fragmentation had different and complex effects on the spacing pattern of both sexes. In food-supplemented patches, high densities led to increased intra-sexual overlap. In linear-shaped patches, squirrels used smaller home ranges and core areas and had lower male–male and male–female overlap levels, independent of patch quality or size. Home range and core area size of males increased with patch size, and male core areas overlapped extensively those of other males and females. Hence males seemed to show a fission response only in some patches. In contrast, home range and core area size of females was not related with patch size, but decreased with habitat quality, supporting predictions of a fusion response and intra-sexual defense of food-based core areas. Hence, where patch size and shape strongly affected space use of male red squirrels, social organization of females was only affected in small, food-supplemented patches, suggesting that the basic spatio-social organization of adult females is very resistant to fragmentation.     

Pattern of space use by female black bears in the White River National Wildlife Refuge,Arkansas, USA

The manner in which space is used by animals may influence several aspects of biology, including the pattern of resource use and intra-specific competition. We monitored 16 radio-collared female black bears (Ursus americanus) for 9,216 radio days during 1993–1995 in the White River National Wildlife Refuge (WRNWR), Arkansas, U.S.A. to investigate space use patterns. Annual home ranges (95% convex polygon) ranged from 2.10 to 11.34 km² with a mean (± SD) size of 4.90 (± 2.09) km² (n = 16). Largest home ranges were occupied by 2 females with yearlings during one year of study. Home ranges among neighbouring bears overlapped considerably. Although bears maintained larger home ranges during summer, the size of home range did not differ among seasons (P > 0.50). Our estimates of home range size for female black bears were smaller than those obtained in a study of the same population during 1979–1982. Because the size of the bear population at WRNWR was substantially smaller (about 130 bears) during 1979–1982 compared to the present population of ≥348 bears, these results suggested that population density and size of female black bear home ranges may be negatively correlated. Conservation implications of density-dependent space use pattern are also discussed.     

Home ranges and movement patterns in a vulnerable polecat<Emphasis Type="Italic">Mustela putorius</Emphasis> population

Home range characteristics and movement patterns of four female and six male polecatsMustela putorius Linnaeus, 1758 were studied in Luxembourg using radiotelemetry. Home range size of polecats ranged from 42 to 428 ha with an average of 181 ha. The mean (± SE) home range size of males of (246±45 ha) was significantly larger than that of females (84±17 ha). Polecats concentrated 50% of their space use in only 15% of their home range possibly indicating a patchy environment. Comparing our data with other studies in Europe, polecats seem to occupy approximately the same home range size (except in Switzerland) regardless of population density. Average distance traveled per night by males was 3.6 times greater than that of females. Also, seasonal variation in movements was observed in males but not in females.     

Spatial relationship among individuals of the Japanese lacertidTakydromus tachydromoides (Sauria,Lacertidae)

The home range ofTakydromus tachydromoides was studied in a grassland area from April 1977 to November 1978. The mean size of home range did not differ markedly between sexes; 136.5 m² for males and 130.8 m² for females. Home ranges of adults overlapped greatly in each sex, and the lizard was considered to be non-territorial. Individuals showed return movement to a definite area (sleeping site) within the home range, and the home range did not shift within a year or between years. Characteristics of the home range of this grassland-inhabiting lizard were discussed in relation to resource abundance and predation pressure.     

Spatial and temporal habitat use of kob antelopes (Kobus kob kob, Erxleben 1777) in the Comoé National Park, Ivory Coast as revealed by radio tracking

Spatial and temporal habitat use of kob antelopes (Kobus kob kob) have been investigated in the Comoé National Park (Ivory Coast, West Africa) by use of radio telemetry. A total of 23 kob were equipped with radio collars and radio tracked for up to 15 months. Home ranges of males were smaller and those of females larger than expected from theoretical models. Adult males used smaller areas than adult females and did not show seasonal home range shifts. Daily distances travelled did not differ between sexes. Kob walked less during the night than by day and covered shorter distances in the wet season. Whereas an increase in home range overlap between females resulted in higher rates of association among individuals, association of adults of mixed sexes was not correlated with the degree of home range overlap. Territorial behaviour of males and predator avoidance by females are suggested to explain the sex‐specific differences in home range size of adults and the deviation from the predicted sizes. Predator avoidance is presumed as the main reason for the reduced walking distances at night as well as in the wet season. Reproductive behaviour and feeding ecology are assumed to determine the degree of association of conspecifics.     

Home ranges in adult Scandinavian brown bears (Ursus arctos): effect of mass, sex, reproductive category, population density and habitat type

Annual home-range size indices for 36 male and 52 female adult brown bears Ursus arctos in two study areas in central and northern Scandinavia were estimated to evaluate factors believed to influence home-range size. Male home ranges were larger than home ranges of lone females after controlling for the sexual size dimorphism acting on metabolic needs. Further, home ranges of females with cubs were smaller than home ranges of lone females and females with yearlings. Thus, differences in metabolic need were not able to explain the variation in range size among females of different reproductive categories or between males and females, suggesting roaming behaviour of males in this promiscuous species. Home-range size in both males and females was inversely related to population density along a density gradient that was not linked to food availability. This contradicts the hypothesis that females use the minimum areas that sustain their energy requirements. However, on a large geographical scale a negative relationship between range size and food availability was evident. The annual home ranges in inland boreal environments in Scandinavia are the largest reported for brown bears in Eurasia, and similar to those in inland boreal and montane environments in North America.     

Staying close to home: Ecological constraints on space use and range fidelity in a mountain ungulate

Understanding patterns of animal space use and range fidelity has important implications for species and habitat conservation. For species that live in highly seasonal environments, such as mountain goats (Oreamnos americanus), spatial use patterns are expected to vary in relation to seasonal changes in environmental conditions and sex‐ or age‐specific selection pressures. To address hypotheses about sex, age, and seasonality influence on space‐use ecology, we collected GPS location data from 263 radio‐collared mountain goats (males, n = 140; females, n = 123) in coastal Alaska during 2005–2016. Location data were analyzed to derive seasonal and sex‐specific fixed‐kernel home range estimates and to quantify the degree of seasonal range and utilization distribution overlap. Overall, we determined that home range size was smallest during winter, expanded coincident with the onset of green‐up and parturition, and was largest during summer. Home range size of males and females did not differ significantly during winter, but females had larger home ranges than males during summer, a relationship that was switched during the mating season. Pairwise comparisons involving individual females across subsequent years indicated home ranges were significantly smaller during years when they gave birth to offspring. Mountain goats exhibited a strong degree of range fidelity, and 99% (n = 138) of individual animals returned to their previous year''s seasonal range with an average annual Bhattacharyya''s affinity utilization distribution overlap index of 68%. Similarity of seasonal home range utilization distributions varied in relation to sex and season in some respects. Home range overlap was highest during the summer vegetation growing season, particularly among females. These findings advance our understanding about how environmental variation and sex‐ and age‐related reproductive constraints influence space use and range fidelity among alpine ungulates. Documentation of the high degree of range fidelity among mountain goats has important conservation implications in landscapes increasingly altered by anthropogenic activities.     

The ecology of 3-D space use in a sexually dimorphic mammal

The distribution of animals is the result of habitat selection according to sex, reproductive status and resource availability. Little is known about how marine predators investigate their 3-dimensional space along both the horizontal and vertical axes and how temporal variation affects space use. In this study, we assessed the spatio-temporal movement of a sexually dimorphic marine mammal, the grey seal Halichoerus grypus by 1) determining seasonal home range size, 2) testing whether space use of seals was affected by water depth, and 3) investigating the vertical movement of seals according to the maximum depth of each dive. Between 1993 and 2005, we fitted 49 grey seals in the Gulf of St. Lawrence with satellite transmitters. We estimated seasonal 95% fixed-kernel home ranges for each individual. For each seal, we tested for selectivity and preference for 4 water depth classes at the home range scale and within the home range. We also evaluated the proportional number of dives made in each water depth classes according to the maximum depth of each dive. Home ranges were 10 times larger in winter than in summer. Seals generally selected habitats <50 m deep. They also mainly dove to depths of 40 m or less. At both scales of selection, preference for shallow areas decreased in winter. We also observed that adults used shallow habitats more than juveniles to establish their home range. A spatial segregation based on sex also occurred at the finer scale of selection where females were more concentrated in the shallowest parts of their home range than males. Segregation in space use according to age and sex classes occurred at both the horizontal and vertical scales. Our results emphasise the importance of studying habitat selection of marine predators in 3-dimensional space, in addition to the temporal scale.     

Space use and site fidelity of wintering whooping cranes on the Texas Gulf Coast

The Aransas-Wood Buffalo population (the only non-reintroduced, migratory population) of endangered whooping cranes (Grus americana) overwinters along the Texas Gulf Coast, USA. Understanding whooping crane space use on the wintering grounds reveals essential aspects of this species' ecology, which subsequently assists with conservation. Using global positioning system telemetry data from marked whooping cranes during 2009–2017, we fit continuous-time stochastic process models to describe movement and home range using autocorrelated kernel density estimation (AKDE) and explored variation in home range size in relation to age, sex, reproductive status, and drought conditions. We used the Bhattacharyya coefficient of overlap and distance between home range centroids to quantify site fidelity. We examined the effects of time between winter home ranges and the sex of the crane on site fidelity using Bayesian mixed-effects beta regression. Winter whooping crane 95% AKDE home range size averaged 30.1 ± 45.2 (SD) km² (median = 14.3, range = 1.1–308.6). Home ranges of sub-adult females were approximately 2 times larger than those of sub-adult males or families. As drought worsened, home ranges typically expanded. Between consecutive years, the home ranges of an adult crane exhibited 68 ± 31% overlap (site fidelity), but fidelity to winter sites declined in subsequent winters. The overlap of adult home ranges with the nearest unrelated family averaged 33 ± 28%. As a whooping crane aged, overlap with its winter home range as a juvenile declined, regardless of sex. By 4 years of age, a whooping crane had approximately 14 ± 28% overlap with its juvenile winter home range. Limited evidence suggested male whooping cranes return to within 2 km of their juvenile home range by their fifth winter. Previous data obtained from aerial surveys led ecologists to assume that whooping crane families normally used small areas (~2 km²) and expressed persistent site fidelity. Our analyses showed <8% of families had home ranges ≤2 km², with the average area 15 times greater, and waning site fidelity over time. Our work represents an analysis of whooping crane home ranges for this population, identifying past misconceptions of winter space use and resulting in better estimates of space requirements for future conservation efforts.     

Intra-group home ranges in a female-biased group of sex-changing fish

Simultaneous loss of multiple males from large, multi-male social groups of the coral reef fish, Anthias squamipinnis, induces an equivalent number of females to change sex. One-to-one replacement could be produced in large groups if the group was organized according to either of two hypotheses. The group could consist of an aggregation of single-male, multi-female units with the home ranges of particular females falling within the home ranges of each male, and with sex change controlled separately within each unit. Alternatively, females might range freely throughout the group, with sex change controlled by differential interaction rates among males and females. In the latter case, female home ranges would overlap substantially with the home ranges of several males. To choose between these hypotheses, the three-dimensional, intra-group home ranges of five males and five adult females were measured in a group containing five males and 32 adult females. All 10 fish had definable home ranges. Home ranges of females were larger than home ranges of males. All females ranged much farther outside of the home range of individual males than inside, and most females spent most of the observation time outside of the home range of any one male. These results effectively disproved the hypothesis of spatially-defined, single-male, multi-female units. Rather, large groups consist of females ranging freely over the discrete home ranges of several males.     

The Breeding System of the Acorn Woodpecker in Central Coastal California

Adult spatial relationships and social organization in a marked population of alpine mammals, the pika (Ochotona princeps), were studied over a 3-year period in the Rocky Mountains of Colorado. Home range size, distances between centers of activity of dyads, and weighted overlaps of home ranges of dyads were used to define space use patterns. Disappearance and establishment of individuals reflected the temporal component of space use. Relative frequencies of foraging (haying and feeding), surveillance, and communication (short calling, long calling, cheek rubbing) behaviors were recorded. Social relationships among adults were defined by agonistic interactions (aggression) and affiliative behaviors (social tolerance, vocal duets, copulations). Males and females occupied individual home ranges of equal size on talus, their obligate habitat type. Adjacent home ranges were normally occupied by pikas of the opposite sex. Replacement of home ranges was always by a member of the same sex as the previous occupant. A high degree of affiliative behavior was expressed between spatially contiguous heterosexual dyads. Aggression was greater among intrasexual than heterosexual dyads, and spatial overlaps among intrasexual conspecifics were less than among heterosexual conspecifics. Ecological constraints, such as the distribution of food (primarily located in meadows adjacent to the talus) and the short summer reproductive season (placing a premium on early appropriately timed litter;;) have apparently led to a facultatively monogamous mating system. Males can neither monopolize essential resources sufficiently to attract several females, nor defend groups of females as social repulsion among females further increases their dispersion.     

Spatial organization in the Ethiopian wolf Canis simensis: large packs and small stable home ranges

The spatial organization of the rare Ethiopian wolf ( Canis simensis ) was studied in the Afroalpine heathlands of Bale Mountains National Park, southern Ethiopia, between 1988 and 1992. Nineteen animals were radio-tracked, 48 ear-tagged and 64 others recognized by coat patterns and observed directly. Dry season (October—March) home ranges of resident wolves covered between 2 and 15 km². The ranges of adult males were slightly larger than those of females, and subadults' home ranges were slightly smaller than those of adults. The population density of the wolves was correlated with prey biomass. In optimal habitat, wolves lived in packs of 3—13 adults (mean 5.9 wolves > 1year old) containing several close-kin males; adult sex ratio favoured males 1.88: 1 and combined pack home ranges averaged 6.0km². In an area of lower prey productivity, wolves lived in pairs or small groups (mean 2.7), adult sex ratio was 1:1 and home ranges averaged 13.4 km².
Home ranges overlapped extensively (mean 85%) between members of the same pack. Four to seven percent of the population was additionally composed of non-resident females, inhabiting larger ranges (mean 11.1 km²). Home ranges of neighbouring packs were largely discrete, forming a tessellating mosaic of packs occupying all available habitat. Pack home ranges were stable in time, drifting only during major pack readjustment after the disappearance of a pack or significant demographic changes. Ethiopian wolf home ranges were smaller than would be expected for a carnivore of its size and sociality, presumably as a result of the high rodent productivity of the Afroalpine ecosystem.     

Habitat selection and home range size of field voles<Emphasis Type="Italic">Microtus agrestis</Emphasis> in Słowiński National Park,Poland

Habitat preference, home range size and intra-specific overlap were investigated in summer 1998 among field volesMicrotus agrestis (Linnaeus, 1761) in Słowiński National Park (N Poland). Eight individuals (2 females, 6 males) were radio-tracked for one week in August. Field voles were shown to exhibit a marked preference for meadow and the ecotone between grassland and habitats with common reed, while avoiding alder forest and proper reedbeds. No significant differences between night and day in habitat-use of voles were noted. The results suggest that, at the end of the breeding season, it was food resources, rather than the risk of predation, played an important role in the voles’ utilisation of space. The home ranges of males were larger and more diverse than those of females; their sizes being correlated with body mass, such that heavier males had larger home ranges. This further suggests that intra-sexual competition exists between males for females. The low number of females influenced their spatial behaviour, as females had completely exclusive home ranges. Four males (out of six) had overlapping home ranges with other males; three of the overlaps were of less than 20%. Attributes of promiscuity (such as a 3.5:1 operational sex ratio of males to females, intra-sexual competition between males and the territorial exclusivity of females) influenced the social system. However, the period of radio-tracking during this study was too short to define accurately the social system in the field vole population.     

Space use patterns of mountain hare (<Emphasis Type="Italic">Lepus timidus</Emphasis>) on the Alps

Populations on the limits of species’ distribution can show different behavioral adaptations to strong ecological pressure than in the central part of the range. We investigated space use patterns of alpine mountain hare (Lepus timidus) at two areas on the southern edge of the species’ range. We monitored 34 hares between 2005 and 2008, estimating home range size, overlap, and site fidelity, and compared our results with space use in Scottish and North-European populations. Home ranges of mountain hares did not differ between two study areas with different habitat types. Subadult animals used larger ranges than adults and both age groups reduced home range size in autumn, a period that might be critical for hares due to changes in diet and/or high energy expenditure during the previous breeding season. Home ranges in these alpine populations were smaller than in Scandinavian populations but within the range of populations in different habitat types in Scotland. Seasonal home ranges overlapped considerably, but differed among the sexes: male–female overlap was higher than same sex (male–male and female–female) spatial overlap. Seasonal shifts of home ranges were small, and site fidelity remained high over the seasons, suggesting that resource distribution remained constant throughout the year and that the knowledge of an intensively frequented area is an important element of habitat quality. We concluded that habitat structure and availability of mates interact in affecting mountain hare space use in alpine habitats.     

Home range size and activity of the wood lemming, Myopus schisticolor

We present the results from a radiotelemetric study on space use and activity in a natural population of the wood lemming Myopus schisticolor. Male home ranges were larger than females, and the ratio female home range to male home range was smaller than expected compared to other small rodents. Males moved distances 4-12 times longer than females. We argue that this difference in mobility gives a higher probability of capturing males than females in snap trap studies. If there is a sex biased trappability, this might explain the increasing female biased sex ratio during the snow free season (1:1 in spring, 3:1 [female:male| in autumn), as a higher mortality of males during the reproductive season.     

Home range size and distribution in a wild population of Cape pangolins, Manis temminckii, in north‐west Zimbabwe

The aim of this study was to estimate the size of the home range used by individual Cape pangolins ( Manis temminckii ). The study was conducted in 1991–95 at Sengwa Wildlife Research Institute, Gokwe, Zimbabwe. Radio‐telemetry was used to repeatedly locate study pangolins. Home range area was estimated from the location of the burrows used by each pangolin. It was established that pangolins must be tracked for 85+ days to generate a reliable estimate of home range. The size of home ranges, determined from 1141 tracking days of data for 10 pangolins (3·0–15·8 kg body mass) that were each tracked for at least 85 days, was from 0·17 to 11·07 km². Larger (older) pangolins used more burrows and had larger home ranges than smaller (younger) pangolins. The data indicate that large adult males had larger home ranges than large adult females. Within each sex, the home ranges were adjacent to each other with only slight overlap at the boundaries. There was clear overlap of home ranges between males and females.