The interaction of testosterone and breeding phase on the reproductive behavior and use of space of male zebra finches

It is well known that androgens play a critical role in mediating the reproductive behavior of males. However, many laboratory experiments that examined the effects of testosterone in male songbirds typically limited their investigations to the early phase of breeding. We sought to determine the influence of testosterone on social behavior, pair bonding, nesting, and use of space in captive zebra finch (Taeniopygia guttata) males as a function of breeding stage (pre-laying, incubation, and nestling phases). Fourteen males were released into an aviary with 14 females and allowed to breed for 7 weeks. Half of the males were given testosterone implants and half were given control implants. During the pre-laying phase, testosterone-implanted males spent significantly more time in nesting activities than control birds and more time elapsed from starting to build a nest to when their mates initiated egg-laying. During the incubation phase, testosterone-implanted subjects spent significantly more time in female-directed and undirected singing. Use of space varied between hormone conditions depending upon breeding phase: there was no difference during pre-laying, but during the incubation and nestling phases, testosterone-implanted subjects used significantly more space. This significant increase in "home range" during the latter phase of the breeding cycle coincides with results from field studies on other species. These results underscore the importance of considering breeding phase in assessing the behavioral sensitivity to hormones.     

Repeatability of basal metabolism in breeding female kittiwakes Rissa tridactyla

We studied kittiwakes (Rissa tridactyla) breeding near Ny-Ålesund (79° N, 12° E) on Svalbard. In 1997, the basal metabolic rates (BMRs) of 17 breeding females were measured during the incubation and chick-rearing periods. The mean body mass of the kittiwakes decreased significantly (by 10%) between the incubation and chick-rearing periods. At the same time, both the whole-body and mass-specific BMRs decreased significantly. There was a positive and significant relationship between the BMR residuals from the incubation period and those from the chick-rearing period. Thus, the BMR of incubating female kittiwakes is a significant predictor of their BMR during the chick-rearing period. New BMR data were collected in 1998 from ten of these females, measured around the chick-hatching date. Repeatability values were calculated using either (i) the data for eight individuals for which three BMR measurements existed, or (ii) all the data from both years, yielding significant repeatabilities of 0.52 and 0.35, respectively. These values indicate that between 48 and 65% of the observed variation in BMR is due to intraindividual variability, while between-individual variability accounts for 35 to 52% of the variation in the BMR. This is the first report of a significant repeatability of the BMR of an endothermic organism across an elapsed time of more than one day.     

Sex differences in Little Auk Alle alle parental care: transition from biparental to paternal-only care

Understanding differences in male and female care in biparental care systems can help interpret the selective pressures that shape parental strategies. We examined Little Auk Alle alle parental care at a breeding colony during the chick-rearing and fledging periods by conducting observations on marked, known-sex pairs, and by examining the sex ratio of birds carrying food to the colony. Little Auks transitioned from biparental to mostly paternal-only care during late chick-rearing. Males delivered more meals and spent more time at the colony than females during late chick-rearing. Very few females were present at the colony by the end of chick-rearing and through the fledging period, and all marked parents observed accompanying their chick to sea were male. Chick mass loss prior to fledging was associated with the lack of provisioning by the female parent, rather than a reduction in feeding frequency by both parents. The occurrence of paternal-only care during and after fledging is discussed in relation to physiological, ecological and phylogenetic constraints.     

Activity budgets of a marked common loon (Gavia immer) nesting population

A newly devised nightlighting technique was used to capture breeding adult common loons (Gavia immer) at the Seney National Wildlife Refuge in northern Michigan in 1989. The behaviors of 6 pairs of known-sex, color-marked common loons were subsequently quantified during the breeding cycle in 1990. Collected observational data indicate that foraging, resting, locomotion, and preening were frequent throughout the breeding cycle. Time spent foraging was greatest during the pre-nesting period (53 to 57%), but declined significantly during the nesting and post-nesting periods (p<0.05). Time spent foraging during the pre-nesting period was similar to that of fall and winter studies. During the pre-nesting period adult loons spent about 15% of the time in locomotion; this was significantly greater than the other time periods (p<0.05) and is attributed to selecting a nest site. During the nesting cycle, almost half of each bird's time spent was nest-sitting. Sexual differences were negligible during nest-sitting. Resting and chick-rearing were the predominant behaviors during the post-nesting period and were responsible for the biggest difference in parental duties. Time spent preening declined from 8% during the pre-nesting period to 4 to 5% during the post-nesting period. Time spent by nesting pairs to produce chicks is approximately 10% during pre-nesting, 48 to 49% during nesting, and between 38 to 44% during post-nesting. By quantifying and establishing behavioral standards, subtle abnormalities or changes can be detected to better manage for viable common loon populations.     

Individual variation in field metabolic rate of kittiwakes (Rissa tridactyla) during the chick-rearing period

Field metabolic rate (FMR), using the doubly labelled water (DLW) method, was measured in free-ranging adult kittiwakes (Rissa tridactyla) early and late in the chick-rearing period at Svalbard, Norway. Individual variation in FMR was analysed by comparing FMR with body mass, sex, nest attendance, chick age, brood size, and basal metabolic rate (BMR). Mean FMR of kittiwakes during the chick-rearing period was 27.0+/-0.9 (SE) W kg(-1), while the individual variation (calculated as coefficient of variation [CV]) in FMR was 24%. Sex, time spent away from the nest, age of the chicks, and brood size contributed significantly to FMR and explained 65% of the variation in FMR. The FMR increased by 32% from early until late in the chick-rearing period. This occurred simultaneously with an increase in the time spent away from the nest. In 15 of 20 pairs, one of the mates had 15% or higher (mean of the 15 pairs, 22%+/-8%) FMR (W kg(-1)) than their partner, even though the mates spent equal amounts of time away from the nest. This indicates an intrapair conflict in FMR. The variation in total FMR of pairs was 40% less than the individual variation, and total FMR of pairs increased with age of the chicks. This indicates that the mates adjust their energy expenditure within a relatively constant FMR determined by the energy needs of the chicks. Individual variation in FMR could not be explained by variation in body mass or BMR. BMR measured late in the chick-rearing period was 26% lower than previous measurements of BMR from the prebreeding and incubation periods. The increase in FMR and simultaneous decrease in BMR caused a 40% increase in metabolic intensity (FMR/BMR) of kittiwakes during the chick-rearing period. It is suggested that the metabolic intensity is not a proper measure of the metabolic load in seabirds.     

Diving behaviour differs between incubating and brooding Brünnich's guillemots, Uria lomvia

Bird-borne data loggers were used to investigate the foraging strategies of Brünnich's guillemots breeding in a colony in the North Water Polynya: the flight performance and diving activity of incubating birds were compared to those of chick-rearing individuals. No significant differences were recorded between the bird groups in the potential foraging range. Conversely, clear differences were revealed between incubating and chick-rearing birds in diving behaviour. Chick-rearing birds were generally foraging at a significantly greater depth, and spent significantly higher proportions of the time submerged, than brooding individuals. Despite these differences, the estimated average daily energy expenditure of chick-rearing Brünnich's guillemots was only about 6% higher than that during incubation.     

高原鼠兔领域行为时间分配格局及其对风险环境适应的探讨

采用直接观察法测定了高原鼠兔地面、洞道活动时间及进、出洞频率。结果表明,在07：00-18：00,雄性和雌性高原鼠兔平均地面活动时间分别占总活动时间比例的87．09％和85．22％。地面活动时间具有明显的季节性,但两性个体间无显著的差异。繁殖早期,成年雌体地面活动时问显著高于繁殖后期,第2胎幼体地面活动时间显著低于第1胎幼体。不同年龄和性别的高原鼠兔进洞频率存在极显著差异,5月成年雌体进洞频率最高,而成年雄体在4月进洞频率最低。研究结果验证了捕食风险可制约鼠兔属动物领域活动时间分配,高风险环境能增加其利用洞道的时间及频率的特定假设。     

Sex-specific patterns in baseline corticosterone and body condition changes in breeding Red-footed Boobies Sula sula

Plasma levels of corticosterone often mirror changes in body condition and parental effort. In seabirds, the brooding of young chicks is often paralleled by a marked decline in adult body condition. This may reflect the trade-off between the needs of the chick to be fed regularly and brooded, and that of the adult to spend enough time at sea to restore its body reserves and find food for the chick. In this paper, we describe changes in body condition and baseline corticosterone levels in a tropical seabird, the Red-footed Booby Sula sula , in which the chick is continuously brooded for 5–6 weeks. Body condition did not decline during the brooding period. Female body condition remained stable throughout breeding whereas male body condition declined significantly during the late chick-rearing period. Baseline corticosterone levels were low during the prelaying phase, increased significantly during incubation and were highest during the brooding period. Later in the chick-rearing period, baseline corticosterone levels decreased markedly in females, but stayed elevated in males. There was no correlation between baseline corticosterone and body condition. Because of the chick's slow growth and reduced daily need for food, brooding in Red-footed Boobies may not require a marked increase in parental effort, and the rise in corticosterone levels is probably more a reflection of slight changes in foraging activity. Red-footed Booby males are 15% smaller than females and we suggest that the decline in male body condition during the late chick-rearing period is likely to result from higher energetic flight costs or a lower foraging efficiency imposed by a smaller body size.     

Sex roles, parental experience and reproductive success of eastern kingbirds, Tyrannus tyrannus

We quantified parental behaviour of eastern kingbirds during the incubation and nestling periods to determine parental roles, and to examine the impact of previous breeding experience (defined as having bred on the territory in the past) on behaviour and reproductive success. Females performed all incubation, while males spent more than 60% of their time in vigilant or nest guarding behaviour during incubation. Parental roles were not defined as sharply during the nestling period. Females spent more time vigilant, but males provisioned young at only 54% of the rate of females. Vigilance and nest watching were still primarily male duties. Male and female behaviour did not vary with the pair's combination of experience (e.g. experienced-experienced versus inexperienced-inexperienced in previous-current breeding season, respectively) during either phase of reproduction, but experienced males were more vigilant during incubation and fed young relatively more than inexperienced males. Experienced females were also more efficient foragers. Although behaviour did not differ among the four combinations of pair experience, inexperienced pairs none the less lost the most young to starvation and predation. Consequently, inexperienced pairs fledged one less nestling per nesting attempt than did pairs with at least one experienced breeder. Our results suggest that having at least one experienced breeder substantially improved a pair's reproductive success. We propose that female site fidelity is a safeguard to avoid the lower breeding success a female would incur if she were to move to a new territory and breed with an inexperienced male. Copyright 1999 The Association for the Study of Animal Behaviour.     

Parental sex roles of Malaysian plovers during territory acquisition,incubation and chick-rearing

Shorebirds show high variability in parental care strategies among species, populations, and environments. Research on shorebird parental sex roles can help to understand the selective pressures that shape avian breeding strategies. Although several studies have examined parental care strategies in holarctic shorebirds, very little research has been conducted in the tropics. Here we examined parental sex roles during territorial defence, incubation, and chick-rearing in Malaysian plovers Charadrius peronii in the Gulf of Thailand. The costs and gains of particular parental behaviour may vary between the sexes and can be affected differently by environmental factors and chick age. Thus we also examined how temperature, prey availability, chick or embryo age, and time of day affected parental sex roles. Males spent more time defending territories and were further away from chicks whereas females spent more time incubating eggs. Both adults contributed to chick defence during disturbances throughout the entire chick-rearing period. Total nest attendance (sum of both sexes) was affected by the modelled temperature of an unincubated egg. Prey availability, embryo age, and time of day had no effect on total nest attendance. Males adjusted incubation effort in response to temperature only at high temperatures (>36°C) whereas females adjusted nest attendance at high and low temperatures. Chick age had no effect on the proportion of time adults spent defending territories or responding to disturbance. Pairs were more likely to fledge chicks if both the male and female spent more time defending territories. For Malaysian plovers, high cooperation between the sexes during parental care may help to achieve high quality breeding territories, maintain body conditions during hot days, protect offspring from predators and attacking conspecifics, and contribute to high lifetime reproductive success.     

BREEDING BIOLOGY OF THE MANX SHEARWATER PUFFINUS PUFFINUS

Studies on the breeding biology of Puffinus puffinus were carried out in 1963 and 1964 at the large colony on Skokholm, Wales. During the six weeks before laying the birds spent up to a quarter of the days in the burrows, but the ten days immediately prior to laying were normally spent at sea. There is a prolonged laying period, with a marked peak in the first half of May. Details are given of a second egg being laid when the first was deserted immediately after being laid. The male took the first incubation spell. The incubation spells ranged from one to 26 days and averaged six. The incubation period was about 51 days. The frequency of visits to land by breeding birds, unlike those by non-breeders, was not affected by the moon. On hatching, the chicks grew rapidly and reached maximum weights of between 505 and 755 gm. sometime between 39 and 61 days. There was a variable desertion period, usually eight or nine days, before the chicks left the island about 70 days after hatching. During the feeding period the chick received about two feeds every three days. There is evidence that adults visited the chicks more frequently than this. There was no correlation between growth of the chicks, their feeding rates or fledging weights and the time of laying. There was a high survival (about 95 %) of chicks during the fledging period but some eggs were lost in disputes for burrows. Nine pairs in 1964 were unable to raise two young simultaneously. Parents altered their feeding rhythms to try to feed two young but did not themselves lose weight. It is suggested that the critical factor in the production of young is the availability of food for the young immediately after they leave the colonies.     

Flight speeds of two seabirds: a test of Norberg's hypothesis

Norberg suggested that birds should increase their flight speed when rearing chicks in order to maximize chick energy intake by reducing commuting time. We measured the incubation and chick-rearing flight speeds of a medium-range (Brünnich's Guillemot Uria lomvia ) and long-range (Northern Fulmar Fulmarus glacialis ) forager near the Prince Leopold Island colony, Nunavut, Canada. The mean flight speed for the long-range forager was significantly higher during chick-rearing than during incubation. The medium-range forager showed no difference in mean flight speed during the two periods. We suggest that because petrels fly close to their minimum power velocity and have a low wing-loading, whereas alcids fly close to their maximum range velocity and have a high wing-loading, petrels have a greater ability than alcids to alter their flight speed according to changes in the demands of different breeding stages. Consequently, whereas Northern Fulmars adapt to the additional cost of chick-rearing partially by altering flight speed, Brünnich's Guillemots can only do so by reducing mass.     

Low incubation investment in the burrow‐nesting Crab Plover Dromas ardeola permits extended foraging on a tidal food resource

We used GPS data‐loggers, video‐recordings and dummy eggs to assess whether foraging needs may force the low incubation attentiveness (< 55%) of the Crab Plover Dromas ardeola, a crab‐eating wader of the Indian Ocean that nests colonially in burrows. The tidal cycle was the major determinant of the time budget and some foraging trips were more distant from the colony than previously known (up to 26 km away and lasting up to 45 h). The longest trips were mostly made by off‐duty parents, but on‐duty parents also frequently left the nest unattended while foraging for 1–7 h. However, the time spent at the colony area (47%) and the time spent roosting on the foraging grounds (16%) would have allowed almost continuous incubation, as in other species with shared incubation. Therefore, the low incubation attentiveness is not explained by the need for long foraging trips but is largely dependent on a high intermittent rhythm of incubation with many short recesses (5.8 ± 2.6 recesses/h) that were not spent foraging but just outside the burrow or thermoregulating at the seashore. As a result, the eggs were warmed on average only 1.7 °C above burrow temperature, slightly more during high tide periods and when burrow temperature was lower between 20:00 and 10:00 h, only partly counteracting the temperature fluctuations of the incubation chamber. These results suggest that low incubation attentiveness is due to the favourable thermal conditions provided by safe nesting burrows and by the hot tropical breeding season, a combination that allows simultaneous foraging by parents and the exploitation of distant foraging grounds. Why Crab Plovers engage in many short recesses from incubation still remains to be clarified but the need to thermoregulate at the seashore and to watch for predators may play a role.     

Foraging behavior of Adélie penguins during incubation period in Lützow-Holm Bay

The diving behavior of Adélie penguins Pygoscelis adeliae was investigated using time–depth recorders during the incubation period in the fast sea-ice area of Lützow-Holm Bay, Antarctica. Dive profiles and activity/time allocation suggested that penguins were obligated to walk on the fast-ice for 90–100 km until a polynya, which they used as an access to the pack-ice zone. Dive depth did not differ between males and females, though males’ dive duration was longer than that of females. Dive depth was slightly shallower and dive duration was shorter during the incubation than during the chick-rearing phase. Birds dove throughout the day, although less frequently around midnight, and there was no clear diel change in dive depth. This daily dive pattern during incubation period was similar to that previously observed during the chick-rearing period in a fast sea-ice area, but differed from that observed in sea-ice-free area. Variations in diving behavior resulted from different environmental conditions, such as foraging area with different sea-ice condition, as well as from different life history strategies.     

Activity patterns, feeding and burrowing behaviour of the crab Ucides cordatus (Ucididae) in a high intertidal mangrove forest in North Brazil

Activity patterns, feeding and burrowing behaviour of the economically important semi-terrestrial mangrove crab Ucides cordatus (Ucididae, L. 1763) was studied in a high intertidal Rhizophora mangle forest stand in Bragança, North Brazil. Video observations in the rainy and dry season were conducted over 24 h cycles at different lunar phases to investigate the behaviour of these litter-feeding crabs outside their burrows. During the rainy season, crabs stayed inside their burrows for 79% and 92% of the time during day and night, respectively. Time spent for feeding, burrowing and other activities outside their burrows was significantly longer during the day with 9.9% (night: 1.7%) and at waning and waxing moon with 9% (full and new moon: 0.9%). At neap tides (no tidal inundation) foraging and feeding activities outside burrows were clearly light-dependent, increasing at dawn and decreasing at dusk. Highest activities during daytime relate to the visual localisation of food. During the dry season, crabs spent less time inside burrows at neap tides than during the rainy season (80% and 91%, respectively). However, time spent for feeding activities was similar during both seasons. During almost all observation periods crabs collected leaf litter, but rarely fed on it outside burrows. At neap tides nearly all available litter was collected, suggesting that the U. cordatus population is litter-limited during these times. At spring tides (regular tidal inundation) the surface activity of U. cordatus was tide-dependent. Crabs closed their burrow entrances 2-3 h before flooding and re-emerged as soon as the tide retreated. During the day, burrow maintenance was the second most frequent behaviour after feeding. Agonistic interactions were regularly observed and were mainly related to burrow defence. The mean foraging radius of the crabs was only 19 cm (max: 1 m) underneath high Rhizophora mangle trees where crab densities were high. The results point to a high competition for burrows and show that U. cordatus is territorial. It is concluded that several exogenous factors, in particular light, leaf litter availability, flooding of burrows and the presence of conspecifics are important in controlling the crabs' activity patterns.     

Tracking reveals behavioural coordination driven by environmental constraints in the Black-vented Shearwater Puffinus opisthomelas

During the breeding season, seabird foraging trips are constrained by nest attendance schedule and are necessarily colony centred. Oceanographic cues play a major role in the choice of foraging areas to minimize the time spent away from the nest. Here, we analysed the foraging tracks of Black-vented Shearwaters Puffinus opisthomelas during the incubation and chick-rearing periods of 2016 and 2017 at Isla Natividad (Mexico). We applied expectation-maximization binary clustering to track data to clusterize different behaviour patterns during foraging flights. We then applied binary generalized linear mixed models to characterize of foraging areas based on of environmental variables. We finally used kernel estimation techniques to describe main foraging areas. In 2016, breeding shearwaters used two core areas for foraging and resting on the water; the core area delineated by males was located northward from the colony in the Vizcaino Bay and the core area for females was located southward from the colony at the entrance of San Ignacio Lagoon. In 2017, males and females used the same areas with no evident segregation. Our study provided the first information on Black-vented Shearwater foraging areas during the breeding season and indicated that sexual segregation within coastal waters off the central Baja California Peninsula might be a foraging strategy during years of warmer ocean, likely less productive regimes. Factors including ocean-climate-mediated sexual segregation at sea, leading to interannual variation in foraging areas, should be considered when evaluating management actions intended to protect critical foraging habitats for Black-vented Shearwaters.     

Pre-incubation feeding activities and energy budgets of Snow Geese: can food on the breeding grounds influence fecundity?

The potential contribution of early spring feeding conditions in the Arctic to clutch size variation was examined in a population of Lesser Snow Geese Anser caerulescens caerulescens. Behavioural observations were combined with energetic analyses of food material to derive an estimate of the energy budgets of pre-laying and laying females. Food intake of females between arrival on the breeding grounds and incubation was considerable; estimated energy gains in this period were in the same magnitude as the cost of one or several eggs. The pre-laying period spent on the breeding grounds can thus be energetically beneficial rather than costly. Accumulation of resources for reproduction in Snow Geese is a continual process including the breeding grounds, and nutrient limitation after arrival in the Arctic cannot sufficiently explain the environmental component of clutch size variation. The timing of migration and follicle development is such that clutch size decisions are sometimes made during the late stages of migration and some-times after arrival. In the latter case food conditions on the breeding grounds may greatly influence clutch size; in the former case they may still influence readjustments of clutch size after the initial decision. The universal negative correlation between clutch size and laying date in Snow Geese can be explained by negative fitness consequences of late hatching, which outweigh the benefits of delayed laying and further nutrient accumulation. Food shortage on the breeding grounds may sometimes be a secondary factor contributing to seasonal clutch size decline.     

Patterns of colony attendance in the Cape Petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica

The annual cycle of colony attendance of the Cape Petrel Daption capense at Nelson Island, Antarctica, covered about 9 months from the start of September until the end of May. Large numbers of birds attended the colony continuously for 6 months from arrival until fledging, except for a 2-week incomplete pre-laying exodus when attendance was reduced to 10% of that during incubation. After 1 month (March) of post-fledging absence, birds returned to the colony for 2 months in numbers comparable to the level in the breeding season. This was followed by a winter absence that lasted only 3 months and was interrupted by two short visits by a small number of birds. Mean attendance was highest during incubation, but short peaks of maximum attendance occurred during the pre-breeding period. Except for the incubation and brooding periods, colony attendance showed high variability both within and between days. Large numbers of visiting birds occurred on the sea and on islets in front of the colony until the end of October. Unemployed birds constituted an almost constant proportion of up to 50% of colony-attending birds during the incubation period. Diurnal variability did not show a consistent pattern but may represent preferential feeding during early morning hours, with additional short departures depending on the length of the daylight period. Colony counts were the most consistent during the second half of incubation, when the lowest counts gave an estimate of the number of breeding pairs and peak numbers an impression of the additional number of attending nonbreeders. Pre-breeding counts revealed the maximum numbers of birds associated with a particular colony. However, for both pre- and post-breeding surveys, several counts a day over a period of at least a week are recommended because of variability in attendance.     

Influence of nest seclusion and nesting material on pre-laying behaviour of laying hens

Provision of nest sites is beneficial for the welfare of laying hens in intensive production systems. The design of these nest sites has a direct effect on pre-laying behaviour. The aim of this study was to investigate the effects of screening off the entrances of the nest boxes with non-transparent flaps and providing nesting or flooring material on pre-laying behaviour. Eighteen individual Hisex brown laying hens and 18 groups of five hens were tested for 2 days in a test cage containing three nest boxes each with a different nesting material (peat, artificial turf and plastic-coated wire mesh). On one of the days the nest boxes were equipped with plastic flaps at the entrances. Pre-laying behaviour was analysed using focal sampling. Both degree of seclusion and nesting material had an effect on pre-laying behaviour of hens. Nest boxes with flaps were visited less frequently but for a longer duration per visit than nest boxes without flaps. Hens spent more time sitting and scratching, and less time standing and moving when flaps were present. Hens also received fewer pecks (group test) when nest boxes were equipped with flaps. Major differences were found between pre-laying behaviour on coated wire mesh compared to peat and artificial turf with shorter duration of nest visits (group test), more standing (group test) and moving (individual and group test), and less sitting (individual and group test) and object pecking (group test) on coated wire mesh. Differences in pre-laying behaviour on peat versus artificial turf were observed for the behaviours moving (less on peat), sitting (more on peat in the individual test) and object pecking (more on peat in the group test).

It is concluded that hens show more settled pre-laying behaviour and more nest-building behaviour in the presence of flaps at the entrances of nest boxes. These results indicate that seclusion of nest boxes with non-transparent flaps is beneficial to laying hen welfare. Differences in pre-laying behaviour also suggest that plastic-coated wire mesh is less suitable as nesting material than peat and artificial turf.     

高原鼠兔间断性移动模式与反捕食对策分析

本文采用直接观测法, 测定了高原鼠兔地面移动频率及每次移动距离。结果表明, 高原鼠兔地面移动频率和每次移动距离依繁殖时期、年龄及性别而存在极显著的差异。繁殖早期, 成年雄体地面活动频率大于成年雌体。繁殖后期, 第1 胎雄性幼体大于同年龄的雌体。繁殖早期, 成年雌体地面活动频率高于繁殖后期, 而幼体与成体之间无显著的差异。繁殖初期雄体每次移动距离大于雌体, 其它时期雌、雄体之间则无明显的差别。雄体每次移动距离逐月降低。雌体每次移动距离无显著的季节性差异。研究结果进一步验证了暴露于风险环境中的植食性小哺乳动物, 主要采取间断性移动模式, 增加反捕食时间投入, 降低风险的假设。