Mortality and reproductive patterns of wild European polecats<Emphasis Type="Italic">Mustela putorius</Emphasis> in Denmark

Despite its relative abundance and wide geographical range, the population dynamics and reproductive biology of the European polecatMustela putorius Linnaeus, 1758 are largely unknown as to the wild living. We therefore investigated age and reproductive status of 239 Danish polecats primarily killed in traffic or trapped during 1998–2004. Males comprised two third of all individuals in all age groups. Based on a static life table, apparent annual mortality was 68% during the first year of life, 33% during the second year and 65% from the third year in both sexes. The mean (± SE) litter size of 5.95 ± 0.62 (n = 18), estimated from placental scar counts, was significantly lower than litter sizes at birth reported for captive individuals but consistent with litter sizes reported for wild polecats in Russia. Female yearlings conceived at the same rate and produced litters of the same size as older individuals.Males had spermatozoa in their testes from February through August. Testes mass peaked in April and May, ie the same period when most females conceive. A lower prevalence of individuals with spermatozoa in yearlings suggests that most males postponed sexual maturity to two years of age.     

Breeding performance and survival in the peregrine falcon Falco peregrinus support an age‐related competence improvement hypothesis mediated via an age threshold

We studied the effects of age on breeding performance and survival probability in a peregrine falcon population, using data from a long term monitoring programme (carried out over 16 yr), in which we were able to identify individual birds. We compared the breeding performance and survival of yearling breeders, first‐time adult breeders and adult breeders. We found significant differences in breeding performance but not in survival. Yearling breeders had lower breeding success than older individuals but the breeding performance of inexperienced adults did not differ from that of experienced adults. We did not find changes in terminal breeding success since peregrines in their last year of life sustained the performance levels shown in previous years although with increasing variability. We found no evidence that attempting to breed affected survival probability in any age group. We argue that differences in breeding performance are related to age, not to breeding experience, and that there is an age threshold, coincident with the development of adult plumage, after which breeding performance is not affected either by age or experience. Peregrines that start breeding as yearlings are likely to have greater lifetime reproductive success than birds entering the breeding pool as adults. Consequently, such birds may represent a set of high quality individuals. Our results support the age‐related competence improvement hypothesis as being the relevant explanation for the increase in breeding performance with age.     

Age-related patterns of reproductive success in house sparrows Passer domesticus

A common life history pattern in many organisms is that reproductive success increases with age. We report a similar pattern in house sparrows Passer domesticus , older individuals performed better than yearlings for most measures of reproductive success. Older males and females began breeding earlier in a given season and fledged more young than their yearling counterparts. Individual males also fledged more young in their second breeding season than they did in their first, but individual females did not show consistent improvement in reproductive success from year one to two. A path analysis indicated that age in both sexes acted primarily through the timing of breeding; earlier nesters laid more eggs and hence fledged more young but did not have more nesting attempts. We tested whether the increased reproductive success with age arose from high quality individuals surviving to be older (selection hypothesis). In contrast to the main prediction of this hypothesis that reproductive success and survival should be positively related, we found that survival from one year of age to two years of age was negatively related to reproductive success in the first year for males and females combined. Additionally, individuals that survived to breed as two-year-olds did not differ in total young fledged in their first year from those that did not survive to their second season of breeding. Our results indicate that fledgling production increases with age due to improvements in timing of breeding, particularly in females, and not because of the loss of poor breeders or increased output. Mechanisms producing age-related differences in timing of breeding warrant further study.     

Elevated mortality among birds in Chernobyl as judged from skewed age and sex ratios

Background

Radiation has negative effects on survival of animals including humans, although the generality of this claim is poorly documented under low-dose field conditions. Because females may suffer disproportionately from the effects of radiation on survival due to differences in sex roles during reproduction, radiation-induced mortality may result in male-skewed adult sex ratios.

Methodology/Principal Finding

We estimated the effects of low-dose radiation on adult survival rates in birds by determining age ratios of adults captured in mist nets during the breeding season in relation to background radiation levels around Chernobyl and in nearby uncontaminated control areas. Age ratios were skewed towards yearlings, especially in the most contaminated areas, implying that adult survival rates were reduced in contaminated areas, and that populations in such areas could only be maintained through immigration from nearby uncontaminated areas. Differential mortality in females resulted in a strongly male-skewed sex ratio in the most contaminated areas. In addition, males sang disproportionately commonly in the most contaminated areas where the sex ratio was male skewed presumably because males had difficulty finding and acquiring mates when females were rare. The results were not caused by permanent emigration by females from the most contaminated areas because none of the recaptured birds had changed breeding site, and the proportion of individuals with morphological abnormalities did not differ significantly between the sexes for areas with normal and higher levels of contamination.

Conclusions/Significance

These findings are consistent with the hypothesis that the adult survival rate of female birds is particularly susceptible to the effects of low-dose radiation, resulting in male skewed sex ratios at high levels of radiation. Such skewed age ratios towards yearlings in contaminated areas are consistent with the hypothesis that an area exceeding 30,000 km² in Chernobyl’s surroundings constitutes an ecological trap that causes dramatic excess mortality.     

Using dynamic N‐mixture models to test cavity limitation on northern flying squirrel demographic parameters using experimental nest box supplementation

Dynamic N‐mixture models have been recently developed to estimate demographic parameters of unmarked individuals while accounting for imperfect detection. We propose an application of the Dail and Madsen ( 2011 : Biometrics, 67 , 577–587) dynamic N‐mixture model in a manipulative experiment using a before‐after control‐impact design (BACI). Specifically, we tested the hypothesis of cavity limitation of a cavity specialist species, the northern flying squirrel, using nest box supplementation on half of 56 trapping sites. Our main purpose was to evaluate the impact of an increase in cavity availability on flying squirrel population dynamics in deciduous stands in northwestern Québec with the dynamic N‐mixture model. We compared abundance estimates from this recent approach with those from classic capture–mark–recapture models and generalized linear models. We compared apparent survival estimates with those from Cormack–Jolly–Seber (CJS) models. Average recruitment rate was 6 individuals per site after 4 years. Nevertheless, we found no effect of cavity supplementation on apparent survival and recruitment rates of flying squirrels. Contrary to our expectations, initial abundance was not affected by conifer basal area (food availability) and was negatively affected by snag basal area (cavity availability). Northern flying squirrel population dynamics are not influenced by cavity availability at our deciduous sites. Consequently, we suggest that this species should not be considered an indicator of old forest attributes in our study area, especially in view of apparent wide population fluctuations across years. Abundance estimates from N‐mixture models were similar to those from capture–mark–recapture models, although the latter had greater precision. Generalized linear mixed models produced lower abundance estimates, but revealed the same relationship between abundance and snag basal area. Apparent survival estimates from N‐mixture models were higher and less precise than those from CJS models. However, N‐mixture models can be particularly useful to evaluate management effects on animal populations, especially for species that are difficult to detect in situations where individuals cannot be uniquely identified. They also allow investigating the effects of covariates at the site level, when low recapture rates would require restricting classic CMR analyses to a subset of sites with the most captures.     

How has the remnant population of the threatened frog Leiopelma pakeka (Anura: Leiopelmatidae) fared on Maud Island,New Zealand,over the past 25 years?

Despite widespread global reports of declining amphibian populations, supporting long‐term census data are few, limiting opportunities to study changes in numbers and survival over time. However, in New Zealand, for the past 25 years (1983–2008), we studied Leiopelma pakeka, a threatened, terrestrial frog that inhabits rocky boulder banks under forest on Maud Island. Using night sampling at least annually on two 12 × 12 m plots, we had 5390 captures of 1000+ individuals, 327 on one plot (grid 1), 751 on the other (grid 2). The mean (±SE) number of frogs found per night was 11.3 (±0.6) on grid 1 and 25.6 (±1.4) on grid 2. We used capture‐recapture models to estimate population size, proportion of animals remaining beneath the surface and survival rate. The mean (±SE) population estimate was 131 (±14.7) frogs on grid 1 and 367 (±38.7) on grid 2. Over 25 years the estimated population increased on grid 1 and fluctuated on grid 2. Some frogs were captured on most sampling visits, others less often, evidently failing to emerge from cover each visit. Using a combination of open and closed population models, we estimated the mean (±SE) proportion remaining underground was 0.63 (±0.12) on grid 1 and 0.53 (±0.07) on grid 2. Our research represents one of the longest‐run population studies of any frog, and we recorded significant longevity, two males reaching 35+ and 37+ years, a female 34+ years. No significant differences occurred between mean annual survival rates of apparent females and males, or between the two sites. The number of toes clipped for individual identification had little influence on the return rate, once the effect of time of first capture was removed.     

Age differences in blue tit Parus caeruleus plumage colour: within‐individual changes or colour‐biased survival?

In many species of passerine birds yearlings display a less elaborate version of the adult secondary sexual traits, but the causes of such differences in ornamentation are not always well understood. We studied age-related changes in blue tit Parus caeruleus UV/blue structural crown coloration, a sexually selected trait. In our Austrian study population, older blue tits, irrespective of sex, displayed on average a more ultraviolet (lower hue, higher UV chroma), more chromatic and brighter crown coloration than yearlings. This age dichromatism was caused by within-individual changes in the expression of crown coloration between years since males and females became more UV, more chromatic and brighter as they aged. Colour biased survival did not contribute to the observed pattern of age dichromatism since crown coloration was largely unrelated to overwinter survival. Between-year repeatability of crown colour was significant for most colour variables but low in general, and lower for females than for males. In the blue tit, yearling males might benefit from being less ornamented by avoiding adult aggression but at the expense of sexual attractiveness. Adaptive explanations of blue tit age dichromatism should however take into account that age effects were of similar magnitude in males and females. This suggests that both male and female yearlings could benefit from being less ornamented and hence that sexual selection might be acting on both sexes simultaneously in this species.     

Age-specific reproductive performance of Black Grouse Tetrao tetrix females

As part of a demographic study of Black Grouse in the southern French Alps, we estimated variations in reproductive parameters in relation to age of females. Both yearlings and adults initiated a clutch, but adults raised about eight times more young (yearlings 0.25 chicks/females, adults 1.85 chicks/female). Yearlings laid significantly fewer eggs and initiated incubation later than older females. Success of nests did not differ between the two age classes, but yearlings tended to re-nest less frequently and exhibited higher chick loss than older females. The lower reproductive success of yearlings was primarily due to their lower ability to raise young. At nest sites, nest concealment, as well as vertical and horizontal structure and density of the vegetation, did not differ between yearling and older females, which is consistent with the similarity in nesting success in the two age classes. Results support the hypothesis that the extent of the difference in number of young raised by yearling and older females increases with predation pressure.     

Body size and condition of coyotes in southern Texas

Body size and condition of coyotes (Canis latrans) from a high-density population in Webb County, Texas (USA) were analyzed for age, sex and seasonal differences during 1980 to 1986. Mean body mass was progressively greater for juvenile, yearling and adult coyotes. Males were heavier and longer than females in each age class. Indices of intraperitoneal fat deposits were similar between sexes. Juveniles continued growth from fall to spring. Adults and yearlings both lost intraperitoneal fat overwinter. Mean body mass of adults decreased overwinter but mass of yearlings did not differ significantly between fall and spring. Territorial and transient female coyotes did not differ in mean body mass, length or indices of subcutaneous fat deposits.     

Population size and survival of the Brazilian Torrent Frog Hylodes heyeri (Anura,Hylodidae)

Population dynamics are influenced by environmental variability and understanding the abundance and persistence of individuals and populations is a fundamental goal of population ecology. Thus, estimating demographic parameters to identify the factors important for population variability is required to understand temporal and spatial dynamics. The stream-living diurnal frog Hylodes heyeri is endemic to the Atlantic Forest of Brazil in the states of Paraná, São Paulo and Santa Catarina. Here we use capture-mark-recapture methods to estimate survival rates and population size of this Brazilian Torrent Frog in Pico do Marumbi State Park, Paraná. We used CJS models for an open population to estimate apparent survival, capturability and population size in two streams. The number of captures during each session was positively correlated with the minimum weekly temperature. Despite that correlation, the most parsimonious model of survival and capturability was the constant model for both parameters, resulting in a monthly survival rate of 0.38 (95% CI = 0.30–0.46). Thus, only the abundance of the frog differed in the two streams (79 vs. 36), with the population size estimate of 187 individuals. Reproduction is seasonal in the Brazilian Torrent Frog and so the low monthly survival rate suggests that animals move over time rather than die, because 38% month⁻¹ survival should result in <1% of the population remaining after 5 months. Thus, researchers must recognize that populations are labile and individuals often move or are washed downstream during heavy rainfall, generating apparently rapid local turnover that is unlikely to reflect true mortality.     

Annual survival and breeding dispersal of a migratory passerine,the Scissor‐tailed Flycatcher

Knowledge of survival rates is critical for advancing our understanding of the dynamics of populations and here we report apparent annual survival and breeding dispersal of Scissor‐tailed Flycatchers (Tyrannus forficatus) breeding at two sites in southwest Oklahoma (Ft. Sill and Wichita Mountain Wildlife Refuge [WMWR]). Our Cormack‐Jolly‐Seber estimate of apparent adult survival for the period from 2008 to 2105 was relatively low (0.514) compared to estimates for 36 other migratory and socially monogamous passerines breeding in North America, and was independent of sex (males: N = 151; females: N = 119), breeding status (territory holder or floater), body mass, site, year, and precipitation during the year prior to breeding. Although apparent survival did not differ between sites, dispersal (N = 66 individuals) was more common and breeding dispersal distance (BDD) was greater for Scissor‐tailed Flycatchers at Ft. Sill where anthropogenic disturbance was more frequent. BDD also increased with body mass at Ft. Sill (but not at WMWR) and, after accounting for it, BDD at Ft. Sill tended to be greater for birds that failed to breed successfully in the past year. Older birds and males had the longest BDDs at WMWR, and males exhibited a similar tendency at Ft. Sill. We contend that our estimate of apparent survival is low, not because of inherently low survivorship, but, instead, as a consequence of frequent permanent emigration from our population. We also suggest that the greater BDD of older birds (WMWR) and males (both sites) reflects a history of selection for dispersal in response to frequent habitat disturbance. Frequent habitat disturbance, in addition to the opportunity to prospect for territories both before and after breeding, probably enable the earliest spring arrivals (typically older birds and males) to often relocate between years.     

Variation in selection pressure acting on body size by age and sex in a reverse sexual size dimorphic raptor

Body size strongly influences fitness, with larger individuals benefiting in terms of both greater productivity and survivorship; for reverse sexual size dimorphic (RSD) species, this relationship may be more complex. We examined the selection pressures acting on body size in male and female Merlins Falco columbarius to assess whether larger or smaller individuals of this RSD species were favoured in terms of survival and breeding performance. For males and females there were clear links between body size and survival but the exact relationship varied by sex. Among males, birds that survived each year class were larger than those that died and yearlings were on average smaller than older birds, but there were no measurable differences among adult males (age 2+). Among females, larger individuals aged 1 and 2 years were more likely to survive, but this size‐based pattern was not apparent in older age classes. Size early in life predicted the lifespan in male Merlins but not as strongly as for females and not for the largest individuals. Reproductive performance based on brood size was not associated with body size in either males or females, but there was a weak positive relationship between female body size and lifetime reproductive success. Selection appears to favour larger males and females but there is no indication that the population is evolving towards bigger individuals, perhaps in part due to selection against the largest birds. Increased survival may allow larger and higher quality individuals to occupy higher quality territories as they age and thereby to accrue greater lifetime reproductive success in the process.     

Identification and breeding of yearling Piping Plovers

ABSTRACT Age of first breeding is an important life history trait. Many Piping Plovers (Charadrius melodus) do not breed as yearlings, but little information is available concerning the age of first breeding. From 2000 to 2006, we marked 991 chicks in three areas in Saskatchewan, Canada, and subsequently determined when 102 (49 females and 53 males) first bred. Females bred significantly earlier, on average, than males. More females (68%) bred as yearlings than did males (41%; P= 0.04), with most others first nesting when 2‐yr old (29% of females and 50% of males). As expected from differences between the sexes in age of first breeding, younger females were more likely to pair with older males than were younger males with older females. Chicks that hatched early in the breeding season did not breed at an earlier age than those that hatched late in the year. Unlike older birds, juvenile Piping Plovers do not replace flight feathers during their first winter. As a result, 18 of 27 yearlings (67%) had worn outer primaries, whereas only one of 123 (1%) older birds had worn primaries. In addition, whereas 20 of 24 yearlings (83%) retained a few buff‐tipped median coverts, none of 119 known older birds had such coverts. As a result, we were able to identify all yearlings by their worn primaries, buff‐tipped median wing coverts, or both. Wing lengths of yearling Piping Plovers were 3% shorter than those of older birds, presumably due to wear. Because there is no evidence of differences in adult survival rates between the sexes and breeding habitat is available, we speculate that fewer yearling males than females breed because primary wear may reduce the ability of yearling males to perform aerial breeding displays.     

Epizootiology of blood parasites in an Australian lizard: a mark-recapture study of a natural population

The dynamics of a naturally endemic blood parasite (Hepatozoon hinuliae) were studied in a lizard (Eulamprus quoyii) host population, using 2 years of longitudinal data. We investigated how parasite abundance in the population varied over time, examined whether certain host sub-populations were more prone to infection, and compared parasite loads in relation to host reproductive behaviour. We recorded blood parasite infections of 331 individuals, obtained in 593 captures. Prevalence (the proportion of the host population infected) of blood parasites was high; approximately 66% of the lizard population was infected. Probability of infection increased with host age and size, but did not differ between the sexes. Within individuals, parasite load (the intensity of infection within individuals) did not vary over time, and was independent of host reproductive behaviour. Parasite load was significantly higher in males compared to females.     

Survival rates and causes of mortality of roe deer <Emphasis Type="Italic">Capreolus capreolus</Emphasis> in a rural landscape,eastern Poland

Game species in the cultural landscapes of Europe are exposed to several sources of anthropogenic pressure. Whilst hunting is part of coordinated wildlife management, poaching is difficult to quantify, and hence, relatively little is known about its impact on survival patterns of game species. We estimated survival and cause-specific mortality rates of 31 roe deer Capreolus capreolus in the surroundings of the Bia?owie?a Primeval Forest, eastern Poland, from 2010 to 2013. The study area consisted of large, extensively used agricultural fields and a mosaic of pine plantations, which were interspersed with natural forest stands and small agricultural fields and meadows. Roe deer were regularly hunted and the forested part of the study area was traversed by large carnivores. Annual survival rate of all deer was 0.69 and did not differ between seasons. Annual survival rates of females and males in each age class (young and adult) were similar. Poaching was a major source of mortality (6 out of 13 roe deer deaths), and poachers did not prefer any sex or age. Adult survival (0.77) was higher than survival of young individuals (0.53); this trend was stronger when poaching was excluded from the analysis. All cases of poaching were recorded between mid-September and February. Although the two parts of the study area were exposed to a similar degree of anthropogenic pressure, all poached individuals occupied home ranges in the mosaic, where the forest cover in home ranges of roe deer was significantly higher than in the field.     

Male quality and territory quality in the great tit parus major

In this study there was no positive significant relationship between the quality of male great tits, as measured by average strophe length (a strophe is a short burst of song), and territory quality. (1) Males singing longer strophes did not defend territories in areas of higher recruitment, and (2) naturally settling males were better singers than replacement birds were. The second result was probably caused by a difference in age composition of the two groups, because naturally settling yearlings and replacement yearlings did not differ significantly in average strophe length. These observations on a non-migratory species are in contrast to those of several other studies on passerine birds, which were mostly concerned with migratory species. The relationship between male quality and territory quality is predicted to be closer in species that migrate than in non-migratory species.     

Age-Dependent Expression of Song in the Collared Flycatcher, Ficedula albicollis

Theory suggests that male ornaments should be reliable signals of age, with more elaborated ornaments reflecting superior quality in terms of experience and/or viability. Bird song is immensely involved in sexual selection, thus not‐surprisingly, it usually shows age‐dependent variation. Although the collared flycatcher Ficedula albicollis has been intensively studied for its sexual traits, and female preference for male age is expected to be strong, there is no quantitative information on age‐dependent expression of song in this species. Here, we fill this gap and, based on phenotypic correlations, we report the relationship between age and several song features. Repertoire size was consistently smaller in yearlings than in older males, but it also tended to increase after the second year of breeding. In a meta‐analysis of effect sizes using data from the literature, we found that the strength of the relationship between age and repertoire size in the collared flycatcher is similar to patterns observed in other passerines. Song rate was inversely related to age, as yearlings sang at higher rates than older males. Generally, effect sizes for the relationship between age and other song traits were of medium level on average, and had broad confidence intervals. Song traits covaried with survival in a direction suggesting that differences in song between age categories are unlikely to result from phenotype dependent survival. Our results generally support the hypothesis that song holds the potential to function as a reliable indicator of male age in the collared flycatcher. However, it seems that not all song traits are unambiguously more expressed in older males than in yearlings.     

Behavioral Ecology of Coyotes: Social Organization,Rearing Patterns,Space Use,and Resource Defense

Two groups of coyotes in which genealogical relationships were known were studied in the Grand Teton National Park, outside of Jackson, Wyoming, U.S.A., from 1977–1982. One group, a pack consisting of parents and some non-dispersing and non-breeding offspring, defended a territory and the food (mainly elk carrion) contained within it, especially during winter, and also had helpers at den sites (5 of 6 were males). The other group, a mated resident pair, all of whose young dispersed during the first year of life, did not defend a territory and never had helpers at dens. Delayed dispersal and retention of some offspring as helpers was related to the presence of an abundant, clumped, and defendable winter food resource. Dispersing yearlings suffered higher mortality than did non-dispersing individuals. Litter size was the same for the pack and resident pair; litter size was not significantly correlated with number of adults in the group or with the number or percentage of pups that survived to 5–6 months of age. The presence of pack helpers was not significantly correlated with pup survival, although there was a positive correlation (r^s = +0.37) between the number of adults attending a den(s) and pup survival. Helpers rarely fed pups and their presence had no appreciable effect on juvenile weight. Helpers partook in den-sitting (pup-guarding), but they did not reduce the amount of time that parents spent at den sites. Helpers also actively initiated and took part in territorial and food defense. The proportion of times that pack members initiated defense was inversely related to intruder density (r = –0.94).     

Progeny sex ratios in a short-lived lizard: seasonal invariance despite sex-specific effects of hatching date on fitness

When fitness returns are sex-specific, selection should favor the facultative adjustment of offspring sex ratios. Seasonal shifts in offspring sex ratios are predicted to be particularly beneficial in short-lived, sexually dimorphic species in which hatching date is linked to adult size, which is related to fitness in a sex-specific fashion. We used four time series of hatching dates and progeny sex ratios in the brown anole (Anolis sagrei), a short-lived lizard with male-biased sexual size dimorphism, to test for such a seasonal shift in progeny sex ratio. In 2 of the 4 years, we also released hatchlings to their natural environment to test for sex-specific effects of hatching date on juvenile survival and adult size. We found that the relationship between hatching date and size the following year was significantly steeper in males than in females, and previous work has shown that adult size is more strongly tied to fitness in males than in females. Based on those results and on further evidence linking hatching date and body size to sex-specific survival and reproductive success, we predicted that sex ratios should shift from male- to female-biased as the breeding season progressed. Contrary to our prediction, we detected no clear seasonal shift in progeny sex ratio. Furthermore, although juvenile survival was correlated with hatching date, this relationship did not consistently differ between the sexes. The observation that progeny sex ratios are seasonally invariant despite several apparent links to adult fitness suggests that the evolution of a seasonal sex-ratio bias is either inherently constrained or requires a stronger selective advantage with respect to juvenile survival.     

Sexually mature and immature yearling male European ground squirrels: a comparison of behavioral and physiological parameters

Early puberty may increase lifetime reproductive success, but may also entail diverse costs. In male European ground squirrels (Spermophilus citellus), age at sexual maturity varies among individuals. We compared sexually active and inactive yearling males under standardized conditions in outdoor enclosures. Non-reproductive yearling males hibernated significantly longer than mature males and emerged when the mating period had already started. Testosterone concentrations peaked in the precocious males during premating and mating, and were depressed in the non-reproductive individuals throughout the study period. Emergence body mass was similar in both groups, although sexually immature males were significantly heavier at the end of the active season. Additionally, non-reproductive yearlings grew faster than precocious individuals. Male–male aggression peaked during mating and the following 4 weeks. Intense aggressive interactions like chases and fights were mostly initiated by reproductive males and were directed towards mature competitors. Aggressive behavior in immature males was mainly expressed as spatial displacements. Non-agonistic interactions with females were only observed in sexually active males. Concentrations of fecal cortisol metabolites did not differ between the two male groups and were elevated in the second half of the active season. Leucocyte numbers were reduced during mating in mature males, whereas in immature animals, cell numbers did not change significantly. Our results indicate that early puberty in European ground squirrels involves lower growth rates, high intrasexual aggression, lower prehibernatory body mass, and probably worse immunological condition during mating.