Effects of high density on spacing behaviour and reproduction in Akodon azarae: A fencing experiment

We studied the short term spacing behavioural responses of Pampean grassland mouse (Akodon azarae) with regard to population density in four 0.25 ha enclosures (two control and two experimental) in the 2011 breeding season. Based on the hypothesis that A. azarae breeding females exhibit spacing behaviour, and breeding males show a fusion spatial response, we tested the following predictions: (1) home range size and intrasexual overlap degree of females are independent of population density values; (2) at high population density, home range size of males decreases and the intrasexual home range overlap degree increases. To determine if female reproductive success decreases at high population density, we analyzed pregnancy rate, size and weight of litters, and period until fecundation in both low and high enclosure population density. We found that both males and females varied their home range size in relation to population density. Although male home ranges were always bigger than those of females in populations with high density, home range sizes of both sexes decreased. Females kept exclusive home ranges independent of density values meanwhile males decreased home range overlap in high breeding density populations. Although females produced litters of similar size in both treatments, weight of litter, pregnant rate and period until fecundation varied in relation to population density. Our results did not support the hypothesis that at high density females of A. azarae exhibit spacing behaviour neither that males exhibit a fusion spatial response.     

Ranging behaviour of little bustard males, Tetrax tetrax, in the lekking grounds

We investigated the ranging behaviour during the breeding season of 18 radiotracked little bustard (Tetrax tetrax) males, a disperse-lekking species inhabiting the cereal pseudo-steppes. The average kernel 95% home range was 60±50 ha and the average cluster 85% area was 17±17 ha. Range structure was as relevant as home range size for explaining the variation in the ranging behaviour of males, which could be partially explained by age, habitat quality and site. Ranging behaviour varied from males defending small and concentrated home ranges with high habitat quality, to males holding larger home ranges composed by several arenas. Our results suggest that social dominance and resource availability may affect ranging behaviour of males during the breeding season. Also, mating systems constraints may play a role on the use of space of males within the lekking ground. The ranging behaviour of a given male may be determined by a tendency to reduce and concentrate the home range as age and social status increase, and several fine-tuning mechanisms adjusting the ranging behaviour to the prevailing environmental or social factors on a given site and year.     

小型啮齿动物种群系统调节复合因子理论的野外实验研究：食物可利用性和捕食对根田鼠种群空间行为的作用模式及其对种群调节的探讨

本项研究在野外围栏条件下,采用析因实验设计,测定食物可利用性和捕食对根田鼠（Ｍｉｃｒｏｔｕｓｏｅｃｏｎｏｍｕｓ）种群空间行为的作用模式。检验的特定假设为,高质量食物较大的可利用性能降低田鼠的攻击行为和活动;捕食能减少田鼠的活动。研究结果表明,食物可利用性能间接地和直接地影响根田鼠的空间行为。附加食物种群具有较高的密度和较小的巢区,且在诱捕期间具有较少的长距离活动和较低的攻击水平。捕食者的存在不直接影响攻击行为,但能影响诱捕期间的长距离活动,此为根田鼠对捕食者存在作出的直接反应。在阐明田鼠种群动态时,应仔细考虑上述因子相互作用的效应。     

Spring home range, spatial organisation and activity of stoats Mustela erminea in a South Island Nothofagus forest, New Zealand

This radio-tracking study reports the spring home range, spatial organisation and activity of 11 stoats Mustela erminea in a New Zealand Nothofagus forest, 1,5 yr after significant seedfall when rodent density was low, but stoat density was high. The average home range of 4 male stoats was 223 (SE = 45) ha, significantly larger than the average range area of 94 (SE = 13) ha recorded for 7 female stoats. Stoats were generally tolerant of sharing space and did not maintain intra- or intersexual territorial spacing systems. There was no evidence of temporal avoidance with several stoats of the same sex showing slight attraction to one another. However, stoats still may avoid one another when in close proximity. Long-term radio-tracking studies are required to determine the general patterns of spacing behaviour in stoats, Male stoats showed higher levels of activity during daylight than females.     

Individual and spatio-temporal variations in the home range behaviour of a long-lived, territorial species

Despite the fact that investigations of home range behaviour have exponentially evolved on theoretical, analytical and technological grounds, the factors that shape animal home range behaviour still represent an unsolved question and a challenging field of research. However, home range studies have recently begun to be approached under a new integrated conceptual framework, considering home range behaviour as the result of the simultaneous influences of temporal, spatial and individual-level processes, with potential consequences at the population level. Following an integrated approach, we studied the influence of both external and internal factors on variations in the home range behaviour of 34 radiotagged eagle owl (Bubo bubo) breeders. Home range behaviour was characterised through complementary analysis of space use, movement patterns and rhythms of activity at multiple spatio-temporal scales. The effects of the different phases of the biological cycle became considerably evident at the level of movement patterns, with males travelling longer distances than females during incubation and nestling periods. Both external (i.e. habitat structure and composition) and internal (i.e. sex and health state) factors explained a substantial amount of the variation in home range behaviour. At the broader temporal scale, home range and core area size were negatively correlated with landscape heterogeneity. Males showed (1) smaller home range and core area sizes, (2) more complex home range internal structure and (3) higher rates of movement. The better the physiological condition of the individuals, the simpler the internal home range structure. Finally, inter- and intra-individual effects contributed to shaping space use and movement patterns during the biological cycle. Because of the plurality of behavioural and ecological processes simultaneously involved in home range behaviour, we claim that an integrative approach is required for adequate investigation of its temporal and spatial variation.     

Female–female aggression is linked to food defence in a poison frog

Habitat occupancy by territorial animals is expected to depend on the distribution of critical resources. Knowledge on female territoriality is scarce, but it has been suggested as a mechanism to defend limited resources for reproduction. A previous study showed female intrasexual aggression to be associated with territorial behaviour in the strawberry poison frog Oophaga pumilio, a diurnal aposematic species with complex maternal care. Here, we investigate the link between spatial distribution of resources important for reproduction and female distribution and behaviour. We observed focal females in their natural habitat in Costa Rica, and recorded the distribution of ecological predictor variables in a grid system. We used the data for calculating home range and territory sizes and for connecting female habitat use to the distribution of potential resources by computing spatial habitat occupancy models. Even though we found females to occupy large home ranges, they were highly aggressive towards other females only inside a small part of their home range, here termed core area. Among the ecological factors, the sustained abundance of ants (main food item of the frogs), the presence of leaf litter and suitable rearing sites for tadpoles predicted female site occupancy patterns. The number of ants per grid was twice as high in the core areas compared to the rest of the female home ranges. Our results suggest that female spacing behaviour is principally driven by the spatial distribution of its main food resource, but that hiding places (leaf litter) and tadpole‐rearing sites also play a role. The defence of areas with sustainably high abundance of ants could be relevant, as egg production and maternal care are energetically highly demanding in this prolonged‐breeding species. Regarding the link between resource defence and maternal care, the reproductive strategy of female strawberry poison frogs resembles that of the females of small mammals comprising same‐sex competition for food and high investment in producing and rearing young.     

Use of space and habitat selection by roe deer Capreolus capreolus in a Mediterranean coastal area: how does woods landscape affect home range?

The home-range dynamics and habitat selection of nine roe deer were studied from March 1994 to August 1994 in the Maremma Natural Park along the Tyrrhenian coast of Italy. The habitat was highly fragmented, with open agricultural fields prevailing in the study area (57%); the climate was Mediterranean. Data on spatial behaviour were collected by radio-tracking techniques. Habitat selection and structure were investigated by compositional and landscape analysis, both within the study area and within the home ranges. Animals of our sample showed spatial-use patterns varying from stationary to roaming. Stationary individuals used small home ranges while roaming ones moved, especially during the reproductive period in July and August. The percentage and structure of woodlands influenced the size of home ranges and the behaviour of males: stationary males used large amounts of woodlands within their home ranges and showed a territorial behaviour whereas males that used a high percentage of fields showed wider home ranges even during the territorial period. Females seemed to be less influenced by the presence and patch-structure of woodland within their home range. Landscape structure and habitat composition seemed to be important factors influencing the spatial behaviour of this roe deer population.     

Does risk of predation by mammalian predators affect the spacing behaviour of rodents? Two large-scale experiments

Predator-prey interactions between small mammals and their avian and mammalian predators have attracted much attention. However, large-scale field experiments examining small-mammal antipredatory responses under the risk of predation by mammals are rare. As recently pointed out, the scale of experiments may cause misleading results in studies of decision-making under predation risk. We studied the effect of small mustelid predators on the spacing behaviour of the gray-tailed vole (Microtus canicaudus) and the bank vole (Clethrionomys glareolus) in two separate field enclosure experiments. The experiments were conducted during the breeding season in North America and northern Europe, where small mustelids have been suggested to be important mammalian predators of voles. As in most of the earlier laboratory studies, predation risk was simulated using fresh mustelid faeces and urine. This made it possible to compare the results from experiments at different spatial scales. We did not find any effect of increased predation risk on spacing behaviour (mean and/or maximum distance moved and home range size) or trappability in either vole species. Simulated predation risk did not affect the breeding of females in gray-tailed voles, as has previously been shown in bank voles. The results disagree with most of the studies conducted in laboratory conditions with small mammals. We discuss whether this discrepancy could be caused by differences in the scale of the experiments. Received: 12 April 1999 / Accepted: 7 October 1999     

Effects of spatial variation in food availability on spacing behaviour and demography of Eurasian red squirrels

In heterogeneous habitats with limited resources, spacing behaviour will affect individual variation in breeding success and density of populations, and is thus of general interest to ecologists. We investigated how red squirrels Sciurus vulgaris adapt their social organisation to tine‐grained heterogeneity in habitat quality, studying spacing behaviour, habitat use and population dynamics in a forest in north Italy, characterised by a mosaic of high‐quality (chestnut‐pine) and poor‐quality patches. We compared the data with those from more homogeneous broadleaf and mixed woodlands with similar overall tree seed abundance (“stable” habitats). Squirrels lived at lower densities (pre‐breeding density 0.39‐0.58 ha^‐1) than in “stable” habitats, although breeding rate was not reduced. Female breeding success was related to being primiparous as yearlings, and increased with body mass and proportion high‐quality habitat in the home range. Persistency rate of females was as in stable habitats, It was higher than male persistency, but immigration and recruitment rates were male biased, resulting in even sex‐ratio. All residents occupied high‐quality patches, and no subadults established a permanent home range in poor‐quality habitat. Home range and core‐area size was typically larger in males than in females and a male's core‐area overlapped those of other males and of females. Female core‐areas were overlapped by males but not. or very little, by other females (intrasexual territoriality). Home ranges, or core‐areas, were not smaller than in “stable” habitats, nor did we find a higher degree of core‐area overlap. We conclude that in patchy habitats dominant, resident red squirrels exclude dispersing animals from preferred, high‐quality habitat, producing a spacing pattern referred to as ideal despotic distribution, and that poor patches were only used temporarily by transient individuals, resulting in a reduction of density in comparison to populations in “stable” habitats.     

Ranging behaviour of feral cattle (Bos taurus) in Doñana National Park, S.W. Spain

The cattle of Doñana (139 individuals in four social groups in 1989) have lived under free-ranging conditions for centuries. Their ranging behaviour was analysed during a three-year period. A total of 17,603 locations corresponding to 247 different animals allowed both for the estimation of global and seasonal home ranges of individuals and social groups and for the comparison of movement patterns. Cattle ranging behaviour was not affected by human interference, and was shown to be regulated by a complex interaction of environment, individual and social factors. Habitat structure and seasonal fluctuations in abundance and distribution of resources determined general patterns of ranging behaviour: the greater the concentration of resources, the smaller the home ranges of individuals and social groups. These patterns were modified at an individual level by the sex of the animal and its reproductive status if male. Social influences on ranging behaviour were important because these implied the segregation of home ranges among dominant bulls and among social groups. As a result, there was a great variability in space use and home-range behaviour.     

Spacing pattern in a social group of stray cats: effects on male reproductive success

Reproductive consequences of male spacing patterns have received relatively little attention in nonterritorial mammals, in particular in group-living species, where most studies have focused on the relation between social rank and reproductive success. We investigated the effects of spacing pattern on male reproductive success within a social, nonterritorial, promiscuous population of stray cats, Felis catus. Male home ranges overlapped home ranges of many females, consistent with a promiscuous mating system. Furthermore, males with the largest home ranges included the most female home ranges; they successfully reproduced with these females and had the highest reproductive success. Home range size predicted male reproductive success even when controlling for the effect of social rank. However, males also reproduced with females whose home range did not overlap their home range, suggesting that males can make quick excursions outside their home range to find new mating opportunities. We conclude that, in group-living situations, a male's ability to maintain a large home range may be one of the principal causes of variation in mating success in the stray cat.     

Territory and group living in the polygynandrous Alpine Accentor Prunella collaris

The spacing system of Alpine Accentors Prunella collaris was studied on the summit of Mt. Norikura in central Japan for five breeding seasons. This species lived in groups (mean 7.2 individuals), sharing large areas of their individual home ranges within which all activities occurred. Membership of a group was closed and stable within a breeding season. The group home ranges overlapped little with each other, and antagonistic behaviour, including communal defence, was observed at the boundaries. Each female established an exclusive area around her own nest which she defended against other females (but not males) within the shared home range, but her activities (feeding, singing and mating) were observed over the whole of the group home range. Members of the same group moved around and fed together within the home range during the prebreeding season, but individual birds tended to become more solitary as the breeding season progressed. These results suggest that the primary breeding unit of Alpine Accentors is a group consisting of five to ten members who share a group territory which contains all the resources necessary for living and breeding, but this species is not a typical social one in which all members move around together within their group territory.     

Spacing of Golden Eagle Aquila chrysaetos nests in relation to nest site and food availability

A regular nest spacing of Golden Eagle pairs was found in every carefully searched area within the species range in Sweden. The spacing regularity of nests used by pairs in forests (mostly tree-nesting) was similar to that in mountain areas (cliff-nesting), indicating that regular nest spacing is affected more by spacing behaviour than by shortage of nest sites. Distances to nearest neighbours were shorter in the mountain region (mean distance 102 km) than in the forests (170) of northern Sweden. Densities of the most important prey species for Golden Eagle were apparently higher in the mountains than in the forests. Differences in topography and habitat diversity between the two regions may also have contributed to the difference in breeding density.     

When Top-Down Becomes Bottom Up: Behaviour of Hyperdense Howler Monkeys (Alouatta seniculus) Trapped on a 0.6 Ha Island

Predators are a ubiquitous presence in most natural environments. Opportunities to contrast the behaviour of a species in the presence and absence of predators are thus rare. Here we report on the behaviour of howler monkey groups living under radically different conditions on two land-bridge islands in Lago Guri, Venezuela. One group of 6 adults inhabited a 190-ha island (Danto) where they were exposed to multiple potential predators. This group, the control, occupied a home range of 23 ha and contested access to food resources with neighbouring groups in typical fashion. The second group, containing 6 adults, was isolated on a remote, predator-free 0.6 ha islet (Iguana) offering limited food resources. Howlers living on the large island moved, fed and rested in a coherent group, frequently engaged in affiliative activities, rarely displayed agonistic behaviour and maintained intergroup spacing through howling. In contrast, the howlers on Iguana showed repulsion, as individuals spent most of their time spaced widely around the perimeter of the island. Iguana howlers rarely engaged in affiliative behaviour, often chased or fought with one another and were not observed to howl. These behaviors are interpreted as adjustments to the unrelenting deprivation associated with bottom-up limitation in a predator-free environment.     

Reproduction and spacing behaviour of females in a peak density population of Clethrionomys glareolus

A peak density population of Clethrionomys glareolus was studied by snap-trapping to determine by experimentation whether spacing behaviour regulated the breeding density. On control plots onset of reproduction was asynchronous, litter size of overwintered females decreased significantly during the summer and no year-born animals matured during the reproductive period. A removal experiment in the early summer resulted in sexual maturation of year-born females. A second removal in late summer indicated an inhibition of maturation in year-born females despite continued intense reproduction of overwintered females. We conclude that spacing behaviour regulated maturation of females early in the summer but other factors, such as food quality, limited maturation in late summer.     

Weaning behaviour in human evolution

Human life history incorporates early weaning, a prolonged period of post-weaning dependency and slow somatic growth, late onset of female reproduction, reduced birth spacing and a significant post-reproductive female lifespan, combined with rapid early brain growth. Weaned human offspring lack the cognitive skills and physical capacity required to locate, procure and prepare foods that are appropriate for their immature state and sufficient for their high energy requirements. During the weaning process and throughout childhood human offspring are supported by the provision of energy dense and easily digestible foods. Changes in weaning behaviour during human evolution imply a shift in the balance between maternal costs of lactation and the risk of poor offspring outcome, and may have been driven by an increase in infant nutritional and metabolic requirements, a reduction reproductive lifespan resulting in selection for reduced birth spacing or a change in other factors affecting offspring survival and fitness.     

Spacing behaviour and habitat use of rock ptarmigan (Lagopus mutus) at low density in the Italian Alps

Studies of rock ptarmigan Lagopus mutus in northern latitudes have shown that, in the breeding season, the majority of cocks pair monogamously and defend small territories, whilst in most populations, a small proportion of cocks are polygynous or remain unmated. Little is known, however, on spacing behaviour and habitat use of alpine rock ptarmigan populations occurring at low densities at the southern edge of the species’ range. From 1995 to 1997, we trapped, radio-tracked and observed birds in the Central Italian Alps (elevation 2,000–3,000 m) in order to investigate spacing behaviour and habitat use in alpine landscapes where habitats offering rich food and cover are patchy. Both sexes were selective in their habitat use, and cocks did not establish territories on bare ground, artificial or nutrient rich grasslands or bogs. In the breeding season, cocks had larger home ranges than hens (cocks 99 ± 57 ha, hens 50 ± 25 ha) that overlapped with the ranges of neighbouring cocks and hens. Cocks were monogamous and defended relatively large territories (core areas of home ranges: cocks 37 ± 26 ha, hens 24 ± 12 ha), which corresponded with low spring densities (0.47–2.29 cocks km⁻² and 0.35–1.60 pairs km⁻²). Territory size of individual cocks was negatively correlated with the amount of high quality habitat in the core-area. Our results suggest increased home ranges and large breeding territories in low density alpine rock ptarmigan populations, compared to populations occurring at higher densities in the central and northern alps, and on subarctic and arctic grounds at northern latitudes, confirming the predictions of models on food-based territoriality.     

A Study of the Behaviour of the Australian Field Cricket Teleogryllus commodus (Walker) (Orthoptera: Gryllidae) in the Field and in Habitat Simulations

The behaviour of T. commodus (Walker) in the field and in simulated natural habitats is described. Regular predictable sequences of behaviour are represented by an ethogram. Most behaviour was centred around burrows or sheltered sites. Burrows appeared to be a limiting resource and there may be behavioural dimorphism in ♂ ♂ (burrower and non-burrower strategies). Individuals were found to form gregarious clusters and appeared to be attracted to areas already occupied by crickets. Territorial spacing and other social interactions occurred within these clusters. ♂ ♂ and ♀ ♀ were found to be mobile, and moved continually between sites. The movement of ♂ ♂ and ♀ ♀ between burrows (calling sites) was measured. Adult ♂ ♂ never remained at a burrow for longer than two days. In simulated natural habitats ♂ ♂ moved in response to two factors: first, and most common, as a result of eviction by other ♂ ♂ second, as a response to the absence of ♀ ♀. When cricket density was increased in habitat simulations crickets clustered at calling sites. The adaptive significance of T. commodus behaviour is discussed.     

Space use and the evolution of social monogamy in eastern rock sengis

Limiting resources often differ between males and females. Reproductive success in females is constrained by resources such as food and shelter, while the availability of receptive females determines male reproductive success. In addition to limiting resources, low intra-sexual tolerances among females can affect their spacing. High intolerances coinciding with low population densities have been suggested as a cause for the evolution of monogamy in mammals, but the evidence for female intolerance is limited. We investigated long-term space use patterns (measured as home range sizes and centers of activity) in a wild population of eastern rock sengis (Elephantulus myurus) from Limpopo, South Africa. Between March 2012 and March 2016, we recorded capture locations for 93 sengis and home range size for 22 sengis and evaluated the contributions of study year, sex, and season on these measures. Sex had no significant effect on home range size, but ranges were significantly greater during the breeding compared to the non-breeding season, consistent with the increased energetic demands during reproduction. We found corroborative support for the role of energetic demands on home range size fluctuations. The activity centers of female–female dyads were further apart than those of male–male or mixed-sex dyads, suggesting lower female–female tolerances in the study species. Our results offer evidence that intolerances may affect female spacing behavior and may have contributed to the evolution of monogamy in sengis.     

Wide home ranges for widely foraging lizards

Space usage by animals may be influenced by a range of factors. In this study we investigate whether foraging behaviour affects the home range size of lizards. Two distinct tactics of foraging have been recognized in predators: sit-and-wait foraging (SW) and active foraging (AF). Foraging activity level of a data set of lizard species, mainly compiled from literature, is compared with their home range sizes. Two opposite predictions can be made about foraging in connection with home range area: on the one hand, SW species may exhibit larger home ranges due to their mating system; on the other hand, AF species have higher metabolic energy and thus food requirements and can be expected to have larger home ranges that have to yield this food. This study shows that percentage of the time moving (as an index of foraging mode) correlates positively with home range, even after correcting for body mass, and these patterns remain when phylogenetic relationships are taken into account. We thus conclude that home range areas parallel activity levels in lizards.