Den sharing behaviour of captive brushtail possums (Trichosurus vulpecula)

Abstract

Den sharing among wild brushtail possums (Trichosurus vulpecula) has important implications for disease transmission. This study investigated den sharing in captive possums, and measured interactions between possums sharing dens. Thirty‐four sexually mature possums (16 female, 18 male) were housed in single‐sex or mixed‐sex pairs in large enclosures that contained two dens. Daily patterns of den sharing were recorded for each pair over a 69 day period in the breeding or non‐breeding season. Social behaviour within shared dens was sampled using miniature infrared cameras. Male pairs rarely shared dens in the breeding or non‐breeding seasons (4% and 1% of days respectively) and usually engaged in ‘threats’ and ‘fights’ associated with den defence. Pairs of female possums (in both seasons) and mixed‐sex pairs housed together in the breeding season shared dens most frequently (between 84% and 91% of days), and also spent the most time together in dens each night. While sharing dens, affiliative interactions were frequent, including long periods of ‘touching’, and also ‘food sharing’ and ‘allogrooming’. The preference for den sharing and close contact shown by captive possums highlights the importance of den sharing as a potential route for disease transmission.     

Hierarchical Den Selection of Canada Lynx in Western Montana

Abstract We studied den selection of Canada lynx (Lynx canadensis; hereafter lynx) at multiple ecological scales based on 57 dens from 19 females located in western Montana, USA, between 1999 and 2006. We considered 3 spatial scales in this analysis, including den site (11-m-radius circle surrounding dens), den area (100-m-radius circle), and den environ (1-km radius surrounding dens). Lynx denned in preexisting sheltered spaces created by downed logs (62%), root-wads from wind-thrown trees (19%), boulder fields (10%), slash piles (6%), and live trees (4%). Lynx preferentially selected den sites with northeasterly aspects that averaged 248. Average distance between dens of 13 females monitored in consecutive years was 2,248 m, indicating low den site fidelity. Lynx exhibited habitat selection at all 3 spatial scales. Based on logistic regression, den sites differed from the surrounding den areas in having higher horizontal cover and log volume. Abundant woody debris from piled logs was the dominant habitat feature at den sites. Lynx generally denned in mature spruce-fir (Picea-Abies) forests with high horizontal cover and abundant coarse woody debris. Eighty percent of dens were in mature forest stands and 13% in mid-seral regenerating stands; young regenerating (5%) and thinned (either naturally sparse or mechanically thinned) stands with discontinuous canopies (2%) were seldom used. Female lynx selected den areas with greater spruce-fir tree basal area, higher horizontal cover, and larger-diameter trees compared to random locations within their home range. Lynx selected den environs in topographically concave or drainage-like areas, and farther from forest edges than random expectation. Maintaining mature and mid-seral regenerating spruce-fir forests with high horizontal cover and abundant woody debris would be most valuable for denning when located in drainages or in concave, drainage-like basins. Management actions that alter spruce-fir forests to a condition that is sparsely stocked (e.g., mechanically thinned) and with low canopy closure (<50%) would create forest conditions that are poorly suitable for lynx denning.     

Denning ecology of the striped skunk in the Canadian prairies: implications for waterfowl nest predation

1. From 1993 to 1995, we studied the summer denning ecology of 40 radio-collared striped skunks Mephitis mephitis (32 females, eight males) during the waterfowl nesting season in Saskatchewan, Canada.
2. Forty-seven natal dens were used by 30 females. Natal den sites were either underground burrows (55%) or holes underneath buildings (45%), and were preferably located in farmsteads.
3. Male and female striped skunks used similar resting sites which consisted of above-ground retreats (57%), underground burrows (23%), buildings (17%) and culverts (3%).
4. Habitat preferences for resting sites did not differ between males and females: both sexes preferred farmsteads and wetlands, whereas managed waterfowl nesting areas, woodland and miscellanous habitats were avoided. Cropland was avoided for both natal dens and resting sites.
5. Females did not simultaneously share natal dens or resting sites, but both types of retreats were occupied by different females over time. Natal dens were occupied for 47 ± 3 days for females using a single den vs. 60 ± 3 days for females using >1 natal dens. Den switching may have been induced by a build-up of faeces.
6. Management of natal den sites such as abandoned farmsteads and buildings may provide an avenue of non-destructive predator management that is less expensive and controversial than exclusion or control.     

Denning ecology of Molina's hog-nosed skunk in a farmland area in the Pampas grassland of Argentina

This paper reports the very first data on the denning habits of the little-studied Molina’s hog-nosed skunk (Conepatus chinga). We identified and characterized 240 (males: n = 92, females: n = 148) den sites used by nine radio-tracked skunks from November 2002 to October 2003 and from December 2005 to October 2007. Most dens were found in underground burrows and we found no variation in dimensions of den entrances across seasons. On average, 24.1% of the den sites were reused by C. chinga. Den sites were not homogeneously distributed within an animal's home range, being the density in core areas greater than in proximity to the borders of home ranges. C. chinga selected specific habitat characteristics for their den sites, such as high shrub and grass cover. Accordingly, the habitats preferred by skunks were native grasslands and pastures. Also, C. chinga preferred den sites located close to fences, roads, and grass patches. We argue that these preferences for den sites may provide two, not mutually exclusive, advantages: access to consistent and predictable sources of food, and protection from predators. Our results suggest that availability of semi-natural grassland and pasture areas may favor the conservation of C. chinga in the human-modified landscapes of the Pampas.     

Survival,density and population composition of bellbirds (Anthornis melanura) on the Poor Knights Islands,New Zealand

Abstract

Annual survival by age, sex and population size and structure were estimated for bellbirds (Anthornis melanura) from a banding study undertaken on 66 ha Aorangi Island, the Poor Knights Islands, during the period 1978–86. The most parsimonious model showed that survival varied little with age and between years, and survival of males was significantly greater than that of females. Recapture probabilities of males and females were similar, but did vary with age and among years. There was a greater proportion of adult males than adult females and sub‐adult males and females in the population, and among adults the male bias in the population was greater during the breeding season than at other times of the year. The high density of bellbirds on the Poor Knights and larger size of males may explain the reduced survival of females relative to that of males.     

Polar bear (<Emphasis Type="Italic">Ursus maritimus</Emphasis>) maternity den distribution in Svalbard,Norway

The aim of the current study was to identify maternity denning areas and to describe aspects of denning behaviour in polar bears (Ursus maritimus) in Svalbard, Norway. Den observations made during aerial and ground surveys (n = 553), observations of recently emerged family groups made during capture and tagging of bears (n = 206), and dens recorded by satellite telemetry (n = 76 dens) were used to describe the distribution of dens in the Barents Sea region. Data were collected during March, April, and May from 1972 to 2010, and while most dens were abandoned during the first 2 weeks of April, there was substantial variation. The highest number of dens was recorded on the islands in the eastern and northern parts of Svalbard with fewer dens found further west on the island Spitsbergen. Females with cubs-of-the-year or their tracks indicated some denning in southern, eastern, and northern parts of Spitsbergen. Telemetry also revealed that females captured in Svalbard and the Barents Sea den in Franz Josef Land, Russia. The majority of dens (62%) in Svalbard were located on land within ca. 1 km of the shore with the most inland den 19 km from shore. Heavy exploitation of polar bears before 1973, when a hunting moratorium was implemented, likely led to reduced denning in some areas. Our observations of den distribution indicate that denning is now more widespread in the archipelago than 50 years ago and reflects a reestablishment of denning areas following years of protection.     

Despotism and risk of infanticide influence grizzly bear den-site selection

Given documented social dominance and intraspecific predation in bear populations, the ideal despotic distribution model and sex hypothesis of sexual segregation predict adult female grizzly bears (Ursus arctos) will avoid areas occupied by adult males to reduce risk of infanticide. Under ideal despotic distribution, juveniles should similarly avoid adult males to reduce predation risk. Den-site selection and use is an important component of grizzly bear ecology and may be influenced by multiple factors, including risk from conspecifics. To test the role of predation risk and the sex hypothesis of sexual segregation, we compared adult female (n = 142), adult male (n = 36), and juvenile (n = 35) den locations in Denali National Park and Preserve, Alaska, USA. We measured elevation, aspect, slope, and dominant land cover for each den site, and used maximum entropy modeling to determine which variables best predicted den sites. We identified the global model as the best-fitting model for adult female (area under curve (AUC) = 0.926) and elevation as the best predictive variable for adult male (AUC = 0.880) den sites. The model containing land cover and elevation best-predicted juvenile (AUC = 0.841) den sites. Adult females spatially segregated from adult males, with dens characterized by higher elevations ( = 1,412 m, SE = 52) and steeper slopes ( = 21.9°, SE = 1.1) than adult male (elevation:  = 1,209 m, SE = 76; slope:  = 15.6°, SE = 1.9) den sites. Juveniles used a broad range of landscape attributes but did not avoid adult male denning areas. Observed spatial segregation by adult females supports the sex hypothesis of sexual segregation and we suggest is a mechanism to reduce risk of infanticide. Den site selection of adult males is likely related to distribution of food resources during spring.     

Kin‐bias,breeding site selection and female fitness in a cannibalistic Neotropical frog

Resource availability influences sexual selection within populations and determines whether behaviours such as territoriality or resource sharing are adaptive. In Thoropa taophora, a frog endemic to the Atlantic Coastal Rainforest of Brazil, males compete for and defend limited breeding sites while females often share breeding sites with other females; however, sharing breeding sites may involve costs due to cannibalism by conspecific tadpoles. We studied a breeding population of T. taophora to determine (i) whether this species exhibits polygynous mating involving female choice for territorial males and limited breeding resources; (ii) whether limited breeding resources create the potential for male–male cooperation in defence of neighbouring territories; and (iii) whether females sharing breeding sites exhibit kin‐biased breeding site choice, possibly driven by fitness losses due to cannibalism among offspring of females sharing sites. We used microsatellites to reconstruct parentage and quantify relatedness at eight breeding sites in our focal population, where these sites are scarce, and in a second population, where sites are abundant. We found that at localities where the appropriate sites for reproduction are spatially limited, the mating system for this species is polygynous, with typically two females sharing a breeding site with a male. We also found that females exhibit negative kin‐bias in their choice of breeding sites, potentially to maximize their inclusive fitness by avoiding tadpole cannibalism of highly related kin. Our results indicate that male territorial defence and female site sharing are likely important components of this mating system, and we propose that kinship‐dependent avoidance in mating strategies may be more general than previously realized.     

Habitat selection by dispersing yellow-headed blackbirds: evidence of prospecting and the use of public information

In migratory birds individuals may prospect for potential breeding sites months before they attempt to breed and should use the cues most predictive of future reproductive success when selecting a breeding site. However, what cues individuals use when prospecting and which cues are used in selecting a breeding site are unknown for most species. I investigated whether yellow-headed blackbirds (Xanthocephalus xanthocephalus) prospect for future breeding sites and whether they select breeding habitats based on food availability, male or female density, or the average number of young produced per female in the previous year. Although it is often assumed that migratory birds prospect for potential breeding sites at the end of the breeding season, I investigated this by recording all visits to sites early and late in the breeding season. I found that males and females who visited sites other than the site at which they bred were more likely to disperse than individuals only observed at the site where they bred, and that males and females were more likely to prospect late in the breeding season. Both food availability and density in year ^x were not predictive of the number of young per female in year ^x+1; however, the number of young produced per female at a site in year ^x was predictive of the number of young per female in year ^x+1. As expected, dispersers used the most informative cue, the number of young per female and moved to sites with a relatively high number of young per female. This study suggests that individuals prospect for potential breeding sites late in the breeding season when they can use information gathered from the reproductive success of other individuals (i.e., public information) to select a breeding site.     

SEXUAL SELECTION AND FITNESS VARIATION IN A POPULATION OF SMALLMOUTH BASS,MICROPTERUS DOLOMIEUI (PISCES: CENTRARCHIDAE)

Monogamy is often presumed to constrain mating variance and restrict the action of sexual selection. We examined the reproductive patterns of a monogamous population of smallmouth bass (Micropterus dolomieui), and attempted to identify sources of within-season fitness variation among females and known-age males. Many males did not acquire a nest site, and many territorial males were unsuccessful in acquiring a mate. The likelihood that territorial males mated depended on several aspects of nest sites. Mated males of age three were larger than the average size of age-three males in the population. The mean sizes of age-four and age-five mated males were not different from the average of same-age males in the population. Thus, selection resulting from the acquisition of a mate favored large size among only age-three males. Timing of nest construction and breeding among territorial males was negatively related to male size and did not depend on male age after taking male size into account. Indirect evidence (numbers of eggs deposited in nests) suggests that the timing of spawning among females was also negatively related to female size. Fertility selection favored early reproduction within the season by males of all ages, but large male size was favored among only age-four males. The combined early breeding of fecund females and female mate choice of large males may explain the positive correlation between the size of age-four males and the number of eggs acquired. Despite large differences of female fecundity, however, the variance of relative mate number contributed about two times more than the variance of relative fertility among females to the total variance of relative fitness within each sex.     

Reproductive behaviour of free-ranging rural dogs in West Bengal,India

Reproductive behaviour of free-ranging dogsCanis familiaris Linnaeus, 1758 was studied in a village in the state of West Bengal, India. Increased synchronized breeding was the most striking feature of this study. October (late monsoon) represented the peak period of mating for the feral dogs. Of all courting males, only 41% were observed to mount and copulate. On average, each male mounted 5.47 ± 2.49 (mean ± SD) times per hour. Of all mountings, only 10% were successful matings, ie copulatory ties. There was a negative correlation between the number of courting males and the number of successful copulations. The average duration of copulatory tie was 15.73 ± 7.75 min. Several factors interrupting the duration of copulatory ties were identified. December was the peak period of pup rearing. Mean litter size was 5.70 ± 2.03 with a male-biased sex ratio 1.41:1. Only a single annual breeding cycle recorded here differed from the previous studies on European and American dogs. Mothers spent most of the time with their pups at the dens during the early stage of rearing. The duration of time spent at dens by mothers was minimum when the pups were highly mobile at the age of 10 weeks. The lactating mothers were observed to be more aggressive immediately following litter production. Typically, an old adult male remained near the den as a ‘guard’.     

Co-existence of subadult males and females as alternative tactics of breeding post acquisition in a monogamous and protandrous anemonefish

Summary The monogamous and protandrous anemonefish,Amphiprion clarkii, utilizes only large host sea anemones as breeding sites. Under conditions of breeding site shortage, an important strategy for unmated fish may be to prepare themselves to enter quickly a breeding site from which a breeding male or female has recently disappeared. There are two types of subadults. Some juveniles become subadult males, which can change either to male or female afterwards, but others become subadult females, which can change only to female. Subadult females seem to acquire female posts more easily than subadult males. We develop an ESS model which evaluates the future reproductive success of subadult females and subadult males and test whether coexistence of subadult males and females can be interpreted as adaptive alternative tactics, using field data from two populations. The model predicts that if mortality does not differ between adult males and females the frequency for juveniles to become subadult females should be equal to the proportion of breeding post acquisition by subadult females among that by all subadults. The model also predicts that if mortality does not differ between subadult males and females, nor in adults, the frequency of subadult females among all subadults should be equal to the proportion of post acquisition. The model, with data from two study sites, suggests that alternative life-history pathways found inA. clarkii have evolved through adaptive tactics of breeding post acquisition.     

Male mate-locating behavior inEuphydryas chalcedona (Lepidoptera: Nymphalidae) related to pupation site preferences

Males of the checkerspot butterfly (Euphydryas chalcedona)patrol and perch near but not on the larval foodplant in search of females. Experiments with tethered butterflies show that searching males chase virgin females for longer times than they do mated females or males. The larvae leave the larval food-plant to pupate. The correspondence between the distance from the larval foodplant to pupation sites and where males search for females suggests that male mate-locating behavior has evolved to maximize the rate of encounters with newly emerged, virgin females. These conclusions are compared to a recent analysis of butterfly mating systems by Odendaal et al. (Am. Nat. 125: 673–678, 1985).     

Den attendance patterns in swift foxes during pup rearing: varying degrees of parental investment within the breeding pair

Parental investment varies in mammalian species, with male care of young being more common in social and monogamous species. Monogamy is commonly observed in canid species, with both males and females, and often “helper” individuals, providing some degree of care for the young. Social units of the swift fox (Vulpes velox), a small North American canid species, usually consist of a male–female pair and occasionally helpers. The role of parental investment and behavior in swift fox society is currently poorly understood. We observed swift fox dens during the pup-rearing season in each of 2 years to evaluate attendance and frequency of visits to natal dens by adult males and females. Female foxes remained at dens longer and visited them more frequently than did male foxes. Female attendance and visitation decreased throughout the pup-rearing season as pups became older and more independent. Environmental factors, including climate and its effect on prey, appeared to contribute to differences in fox behavior between the 2 years. We observed only one fox outside of the breeding pair attending a den in each of the 2 years, both of which were males. We concluded that each of these two foxes were living within the social unit of the male–female pair as a trio, but not serving as a helper and contributing to the care of the pups. Our results increased knowledge of the ecology and behavior of the swift fox, a species of conservation concern in the Great Plains of North America.     

Flagellar setae of the second antennae in decapod shrimps: sexual dimorphism and possible role in detection of contact sex pheromones

Summary

The antenna 2 (antennal) flagella of decapod shrimps are chemotactile, and their setae are proposed as sensilla involved in recognition of females by males via a contact sex pheromone on the surface of the female. Male recognition of females receptive to mating occurs in many caridean species upon contact of male antennal flagella with the surface of a newly molted parturial female. The hypothesis of sexual dimorphism in the number and kind of setae on the antennal flagella of four caridean and one penaeid shrimp species was tested with setal counts and observations on setal morphology. Unique male antennal setae (“male-specific sensilla”) were not identified in any of the species investigated. However, the abundance of antennal setae was significantly greater in males than in breeding females in the palaemonid carideans Palemonetes pugio and Macrobrachium ohione. In the hippolytid caridean Thor manningi and alpheid caridean Alpheus normanni, no sexual dimorphism in setal abundance was demonstrated. In the penaeoid Rimapenaeus similis, males had a higher abundance of antennal setae than the larger breeding females but so did juvenile females, similar in size to males. The sexual dimorphism in antennal sensilla in the palaemonid species and its absence in A. normanni might be related to their different mating systems, but no such association is suggested for T. manningi and R. similis. Setal morphology suggestive of chemoreceptive function (a terminal pore) was observed in all species.     

Wolf denning behaviour in response to external disturbances and implications for pup survival

We conducted a 15-year study (1999–2014) of wolf denning behavior in Naliboki Forest, Belarus, to determine the extent to which some external factors (rain, mosquitoes, human disturbances) and litter size are negatively correlated with pup survival until autumn.For this purpose we entirely documented denning areas of 30 breeding wolf pairs. Initial denning area includes the natal den site and (ii) secondary den sites between which the parent wolves transported sucessively the litter during the rearing season.The number of den sites per denning area ranged from 1 to 4. The distances in kilometres differed significantly between den sites. The total number of dens (surface dens and burrows) was higher in the natal den site compared with the other den sites. The number of surface dens per den site was higher compared with burrows in den sites 1, 2 and 3 but not den site 4. A total of 168 pups was found over the 15 years. Litter sizes ranged from 1 to 11 (average 5.6 ± 1.9). The rate of pup survival ranged from 0 to 100% and averaged 54.5 ± 29%.We examined the bivariate association between environmental conditions (mosquitoes, rainfall and human presence) and number of surface dens, total number of burrows and number of den sites. Environmental conditions were assessed from spring to mid-summer. Results show that there was a positive relationship between environmental conditions and number of den sites, and also that more difficult environmental conditions (rainy weather, many mosquitoes) decreased pup survival both in absolute and in relative terms. A cluster analysis showed that pup survival was also influenced by human presence. Wolves probably have a more complicated denning behaviour following stressful event which may lead to less effectiveness for feeding or protecting pups. The number of pups surviving was also found to be lower when there were many (8–10) pups because den shifting is more risky in this case. Burrows improve pup survival. Our results tend to suggest that, in our study area, a high mortality rate of wolf pups could be related to external and internal factors of stress.     

Annual dynamics of den use by red foxesVulpes vulpes and badgersMeles meles in central Poland

This study on den use by red foxesVulpes vulpes (Linnaeus, 1758) and badgersMeles meles (Linnaeus, 1758) was conducted in central Poland (51°48’N, 19°53’E) over two years. We found a strong negative correlation between monthly den utilisation rate by foxes and air temperature (r=−0.83,p=0.0001). Correlation between temperature and den use by badgers was positive but less distinct (r=0.40,p=0.036). During the whole study period, a negative correlation between the monthly intensity of den use by badgers and by foxes was found (r=−0.51,p=0.008). The seasonal pattern of den utilisation differed between the species. Foxes used the dens mainly in winter and spring whereas in summer and autumn the intensity of den utilisation sharply decreased to a minimum in August-September. A peak in den use by foxes was observed in March and coincided with rut and selection of future breeding dens. In badgers, two peaks in den use were observed: the first in April and the second in August-September. An individual badger used more dens and used them more evenly throughout the year than an individual fox. The same dens were visited by foxes and badgers mostly from January to April, with a maximum in March.     

Mating Behavior and Reproductive Potential in the Turnip Moth Agrotis segetum (Lepidoptera: Noctuidae)

We investigated the lifetime mating potential and the reproductive behavior of male and female turnip moths Agrotis segetum (Schiff.) under field and laboratory conditions. The sex ratio was 1 : 1 in a lab-reared population as well as in two wild populations. Males were capable of mating repetitively a relatively large number of times (mean of 6.7 ± 2.7 matings) when given access to new virgin females throughout their lifetimes. Females seldom mated more than once (mean ± 1.3 ± 0.6 matings), indicating a male-biased operational sex ratio. The mean potential lifetime mating was five times higher in males, while the coefficient of variance was lower in males. There was no differences in longevity between animals that were allowed to mate and animals not allowed to mate, indicating no direct costs or benefits of mating in physiological terms. In males, the number of matings was positively correlated with longevity, but this was not the case in females. Nor was there a correlation between the number of female matings and the number of fertilized eggs. There was a negative correlation between the number of eggs fertilized and the number of times males had previously mated, indicating that male ejaculates were limited. Male spermatophore size also decreased with number of achieved matings. Laboratory-reared females attracted males in the field throughout their lifetimes, with a peak at 3–7 days of age. Wild males, allowed to choose between pairs of caged females in the field, were attracted in equal numbers to females of different ages. Females did not show any mate-rejection behavior in the field. They mated with the first male that courted them. No incidence of mate replacement by males arriving later to already courted females were recorded.     

Den distribution and den use by the red foxVulpes vulpes japonica in Kyushu

The distribution of dens and den use by the red fox was studied in Kumamoto Prefecture between 1968 and 1982. A total of 80 dens were classified into natal, residential, and temporarily-visited dens. Natal dens comprised 12.4% of all dens. Relative proportions were quite stable for a long period. This indicates a constant number of reproducing females in the area for a considerable length of time. Seasonally the utilization of den showed three peaks in February, May and October. The peaks coincided well with the peaks of reproductive activity of female foxes. The results of multiple regression analysis of distribution of dens with respect to selected environmental parameters around densites indicated that the fox selectively utilize open land, rather than densely vegetated areas. The persistence of dens, the selective utilization by foxes of open land and the sizes of dens were all related to the breeding activities of female foxes inhabiting the area. Reproductive females usually used plural dens (“den group”) in one reproductive season. The numbers of those females occuring in the area were maintained at a steady level for fairy long periods. These indicate that the existence of a stable social relationship between females mediated through occupancy of “den groups.”