Reproductive biology and ecology of female polar bears (Ursus maritimus)

Data on age-specific natality rates, litter size, interbirth interval, age of first reproduction, reproductive senescence, age of weaning and cub survival were determined for a free-ranging population of polar bears inhabiting Hudson Bay, Canada, near the southern limit of the species range. Serum progesterone levels were also determined for females at different stages of their reproductive cycle to provide corroborative support for the reproductive parameters described. Animals were live captured using immobilizing drugs and each animal uniquely marked for future identification. First parturition occurred at four or five years of age and the age-specific natality rate increased with age until approximately 20 years, after which it dropped markedly. At least 40% of adult females displayed two-year interbirth intervals and 55% of cubs in their second year were independent of their mother. Mean size of cub litters in spring was 1.9 and 13% of litters had three or more cubs. The natality rate for 5–20-year-old females was estimated as 0.9, higher than that reported for any more northerly polar bear populations where two-year interbirth intervals are rare, fewer than 5% of yearling cubs are weaned and triplet litters occur with less than 1% frequency. Cub mortality was initially high and declined with age. Although cubs in western Hudson Bay were weaned at a younger age and a lighter weight than their counterparts in more northern populations, cub mortality rates were similar. The reason for the marked differences in reproductive parameters in the western Hudson Bay population is not known. We speculate that sea-ice conditions may be sufficiently different to allow weaned bears at a lighter body weight to hunt seals more successfully there than further north.     

Population ecology of polar bears in Davis Strait,Canada and Greenland

Until recently, the sea ice habitat of polar bears was understood to be variable, but environmental variability was considered to be cyclic or random, rather than progressive. Harvested populations were believed to be at levels where density effects were considered not significant. However, because we now understand that polar bear demography can also be influenced by progressive change in the environment, and some populations have increased to greater densities than historically lower numbers, a broader suite of factors should be considered in demographic studies and management. We analyzed 35 years of capture and harvest data from the polar bear (Ursus maritimus) subpopulation in Davis Strait, including data from a new study (2005–2007), to quantify its current demography. We estimated the population size in 2007 to be 2,158 ± 180 (SE), a likely increase from the 1970s. We detected variation in survival, reproductive rates, and age-structure of polar bears from geographic sub-regions. Survival and reproduction of bears in southern Davis Strait was greater than in the north and tied to a concurrent dramatic increase in breeding harp seals (Pagophilus groenlandicus) in Labrador. The most supported survival models contained geographic and temporal variables. Harp seal abundance was significantly related to polar bear survival. Our estimates of declining harvest recovery rate, and increasing total survival, suggest that the rate of harvest declined over time. Low recruitment rates, average adult survival rates, and high population density, in an environment of high prey density, but deteriorating and variable ice conditions, currently characterize the Davis Strait polar bears. Low reproductive rates may reflect negative effects of greater densities or worsening ice conditions. © 2013 The Wildlife Society.     

Delayed female reproduction in equilibrium and chaotic populations

Behavioural and life history polymorphisms are often observed in animal populations. We analyse the timing of maturation and reproduction in risky and resource-limited environments. Field and laboratory evidence suggests that female voles and mice, for example, can adjust their breeding according to the level of risk to their own survival and to survival probabilities and recruitment of young produced under different environmental conditions. Under risky or harsh conditions breeding can be postponed until later in the current breeding season or even to the next breeding season. We develop a population dynamics and life history model for polymorphism in reproduction (co-existence of breeding and non-breeding behaviours) of females in an age-structured population, with two temporally distinct mating events within the breeding season. We assume that, after overwintering, the females can breed in spring and again in summer or they can delay breeding in spring and breed in summer only. Young females born in spring can either mature and breed in summer or stay immature and postpone breeding over the winter to the next breeding season. We show that an evolutionarily stable breeding strategy is either an age-structured combination of pure breeding behaviours (old females breed and young delay maturity) or a mixed breeding behaviour within age-classes (a fraction of females breed and the rest of the age class postpones breeding). Co-occurrence of mixed reproductive behaviour in spring and summer within a single breeding season is observed in fluctuating populations only. The reproductive patterns depend on intraspecific, possibly interspecific, and ecological factors. The density dependence (e.g. social suppression) and predation risk are shown to be possible evolutionary mechanisms in adjusting the relative proportions of the different but co-existing reproductive behaviours.     

Population ecology of polar bears at Svalbard, Norway

The population ecology of polar bears at Svalbard, Norway, was examined from 1988 to 2002 using live-captured animals. The mean age of both females and males increased over the study, litter production rate and natality declined and body length of adults decreased. Dynamics of body mass were suggestive of cyclical changes over time and variation in body mass of both adult females and adult males was related to the Arctic Oscillation index. Similarly, litter production rate and natality correlated with the Arctic Oscillation index. The changes in age-structure, reproductive rates and body length suggest that recovery from over-harvest continued for almost 30 years after harvest ended in 1973 and that density-dependent changes are perhaps being expressed in the population. However, the variation in reproduction and body mass in the population show a relationship between large-scale climatic variation and the upper trophic level in an Arctic marine ecosystem. Similar change in other polar bear populations has been attributed to climate change, and further research is needed to establish linkages between climate and the population ecology of polar bears.     

Sex-specific costs of reproduction on survival in a long-lived seabird

Costs of reproduction on survival have captured the attention of researchers since life history theory was formulated. Adults of long-lived species may increase survival by reducing their breeding effort or even skipping reproduction. In this study, we aimed to evaluate the costs of current reproduction on survival and whether skipping reproduction increases adult survival in a long-lived seabird. We used capture–mark–recapture data (1450 encounters) from two populations of Bulwer''s petrel (Bulweria bulwerii), breeding in the Azores and Canary Islands, North Atlantic Ocean. Using a multi-event model with two different breeding statuses (breeders versus non-breeders), we calculated probabilities of survival and of transitions between breeding statuses, evaluating potential differences between sexes. Females had lower survival probabilities than males, independent of their breeding status. When considering breeding status, breeding females had lower survival probabilities than non-breeding females, suggesting costs of reproduction on survival. Breeding males had higher survival probabilities than non-breeding males, suggesting that males do not incur costs of reproduction on survival and that only the highest quality males have access to breeding. The highest and the lowest probabilities of skipping reproduction were found in breeding males from the Azores and in breeding males from the Canary Islands, respectively. Intermediate values were observed in the females from both populations. This result is probably due to differences in the external factors affecting both populations, essentially predation pressure and competition. The existence of sex-specific costs of reproduction on survival in several populations of this long-lived species may have important implications for species population dynamics.     

Overestimates of maternity and population growth rates in multi-annual breeders

There has been limited attention to estimating maternity rate because it appears to be relatively simple. However, when used for multi-annual breeder species, such as the largest carnivores, the most common estimators introduce an upward bias by excluding unproductive females. Using a simulated dataset based on published data, we compare the accuracy of maternity estimates derived from standard methods against estimates derived from an alternative method. We show that standard methods overestimate maternity rates in the presence of unsuccessful pregnancies. Importantly, population growth rates derived from a matrix model parameterized with the biased estimates may indicate increasing populations although the populations are stable or even declining. We recommend the abandonment of the biased standard methods and to instead use the unbiased alternative method for population projections and assessments of population viability.     

Variation in the response of an Arctic top predator experiencing habitat loss: feeding and reproductive ecology of two polar bear populations

Polar bears (Ursus maritimus) have experienced substantial changes in the seasonal availability of sea ice habitat in parts of their range, including the Beaufort, Chukchi, and Bering Seas. In this study, we compared the body size, condition, and recruitment of polar bears captured in the Chukchi and Bering Seas (CS) between two periods (1986–1994 and 2008–2011) when declines in sea ice habitat occurred. In addition, we compared metrics for the CS population 2008–2011 with those of the adjacent southern Beaufort Sea (SB) population where loss in sea ice habitat has been associated with declines in body condition, size, recruitment, and survival. We evaluated how variation in body condition and recruitment were related to feeding ecology. Comparing habitat conditions between populations, there were twice as many reduced ice days over continental shelf waters per year during 2008–2011 in the SB than in the CS. CS polar bears were larger and in better condition, and appeared to have higher reproduction than SB bears. Although SB and CS bears had similar diets, twice as many bears were fasting in spring in the SB than in the CS. Between 1986–1994 and 2008–2011, body size, condition, and recruitment indices in the CS were not reduced despite a 44‐day increase in the number of reduced ice days. Bears in the CS exhibited large body size, good body condition, and high indices of recruitment compared to most other populations measured to date. Higher biological productivity and prey availability in the CS relative to the SB, and a shorter recent history of reduced sea ice habitat, may explain the maintenance of condition and recruitment of CS bears. Geographic differences in the response of polar bears to climate change are relevant to range‐wide forecasts for this and other ice‐dependent species.     

Population parameters and harvest risks for polar bears (<Emphasis Type="Italic">Ursus maritimus</Emphasis>) of Kane Basin,Canada and Greenland

We estimated demographic parameters and current harvest risks for a population of polar bears (Ursus maritimus Phipps) inhabiting northern Smith Sound and Kane Basin, Canada and Greenland. Our demographic analysis included a detailed assessment of age- and sex-specific survival and recruitment from 141 marked polar bears, using information contained within the standing age distribution of captures and mark-recapture analysis. Total survival rates for females were: 0.374 ± 0.180 (cubs), 0.686 ± 0.157 (ages 1–4), and 0.967 ± 0.043 (ages 5+). Mean litter size was 1.67 ± 0.08 cubs. Females did not reproduce until at least age 6, which is late compared to other populations of polar bears. The model-averaged, mark–recapture estimate of mean abundance (±1 SE) for years 1994–1997 was 164 ± 35 bears. We incorporated demographic parameters and their variances into a harvest risk analysis (i.e., a stochastic, harvested population viability analysis, PVA). Results suggest that polar bears in the region were severely over-harvested during the mark–recapture interval (1992–1997). The current status of the population is unknown.     

Life history characteristics of brown trout in lakes at different stages of acidification

In 66 Norwegian lakes with allopatric populations of brown trout Salmo trutta that ranged from highly acidified to non-acid (pH 4·7–6·6) there was a significant inverse correlation between mean age and relative abundance (CPUE). As mean annual survival rates were not significantly related to CPUE, this population response may indicate that recruitment was lower in low-density lakes. Age-specific masses were significantly higher in populations that had declined in abundance than in unaffected populations. Mean body mass at sexual maturity in both males and females was inversely correlated with CPUE. Between populations, high age at maturity was associated with high survival rates in both sexes. Age at maturity correlated positively with specific growth rate between the ages of 1 and 2 years in females only.     

Effects of female kin groups on reproduction and demography in the gray-tailed vole, Microtus canicaudus

I conducted an experiment with gray-tailed voles, Microtus canicaudus , to test the hypothesis proposed by Charnov and Finerty that populations of voles comprised of female kin groups would grow more rapidly and reach higher densities more quickly than populations in which female kin groups were disrupted. The experiment was conducted in 0.2-ha semi-natural enclosures planted with mixed grasses. In four enclosures, females were unmanipulated (control) and in four enclosures all newly caught females were removed from their natal enclosures and replaced with females of comparable age from another enclosure, such that juvenile females did not settle near their siblings or parents (treatment). I found no significant differences in survival, reproduction, juvenile recruitment, population growth rates, or population size between control and treatment populations. The only difference was the time to sexual maturation for young females, which was 3.1 weeks for control enclosures compared with 4.2 weeks for treatment enclosures. I could not measure reproductive success for individual females, but my results did not support the hypothesis that the presence or absence of kin groups resulted in any biologically meaningful population-level effects. Female voles that have nesting territories adjacent to relatives may accrue some individual benefits, but these benefits are unlikely to contribute to population regulation in gray-tailed voles.     

Life on the edge: the demography of short-season populations of deer mice

We documented populations of deer mice in the Kananaskis Valley, southwestern Alberta, Canada from 1979 to 1997 to determine whether these short-season populations were more, or less, variable than populations in more temperate environments. We then examined patterns of reproduction, age-specific survival, and immigration to explain variation in summer population growth. Population densities showed no multi-annual periodicity and were generally low. At maximum, numbers doubled over the breeding season, but declined over the breeding season in 4 of 16 yr. Variability in population density was low, and similar to that of Peromyscus populations in more temperate environments. No demographic parameters were related to spring population densities, and immigration rates were low when conditions for survival of nestlings and adults were favorable. Variation in summer population growth was attributed primarily to variation in nestling survival among years.     

The Impact of Roads on the Demography of Grizzly Bears in Alberta

One of the principal factors that have reduced grizzly bear populations has been the creation of human access into grizzly bear habitat by roads built for resource extraction. Past studies have documented mortality and distributional changes of bears relative to roads but none have attempted to estimate the direct demographic impact of roads in terms of both survival rates, reproductive rates, and the interaction of reproductive state of female bears with survival rate. We applied a combination of survival and reproductive models to estimate demographic parameters for threatened grizzly bear populations in Alberta. Instead of attempting to estimate mean trend we explored factors which caused biological and spatial variation in population trend. We found that sex and age class survival was related to road density with subadult bears being most vulnerable to road-based mortality. A multi-state reproduction model found that females accompanied by cubs of the year and/or yearling cubs had lower survival rates compared to females with two year olds or no cubs. A demographic model found strong spatial gradients in population trend based upon road density. Threshold road densities needed to ensure population stability were estimated to further refine targets for population recovery of grizzly bears in Alberta. Models that considered lowered survival of females with dependant offspring resulted in lower road density thresholds to ensure stable bear populations. Our results demonstrate likely spatial variation in population trend and provide an example how demographic analysis can be used to refine and direct conservation measures for threatened species.     

Frequency and inheritance of non‐male producing clones in Daphnia magna: evolution towards sex specialization in a cyclical parthenogen?

In Daphnia (Cladocera, Crustacea), parthenogenetic reproduction alternates with sexual reproduction. Individuals of both sexes that belong to the same parthenogenetic line are genetically identical, and their sex is determined by the environment. Previously, non-male producing (NMP) genotypes have been described in species of the Daphnia pulex group. Such genotypes can only persist through phases of sexual reproduction if they co-occur with normal (MP) genotypes that produce both males and females, and thus the breeding system polymorphism is similar to gynodioecy (coexistence of females with hermaphrodites), which is well known in plants. Here we show that the same breeding system polymorphism also occurs in Daphnia magna, a species that has diverged from D. pulex more than 100 MY ago. Depending on the population, between 0% and 40% of D. magna females do not produce males when experimentally exposed to a concentration of the putative sex hormone methyl farnesoate that normally leads to male-only clutches. Natural broods of these NMP females never contained males, contrasting with high proportions of male offspring in MP females from the same populations. The results from a series of crossing experiments suggest that NMP is determined by a dominant allele at a single nuclear locus (or a several closely linked loci): NMP × MP crosses always yielded 50% NMP and 50% MP offspring, whereas MP × MP crosses always yielded 100% MP offspring. Based on cytochrome c oxidase subunit I-sequences, we found that NMP genotypes from different populations belong to three highly divergent mitochondrial lineages, potentially representing three independent evolutionary origins of NMP in D. magna. Thus, the evolution of NMP genotypes in cyclical parthenogens may be more common than previously thought. Moreover, MP genotypes that coexist with NMP genotypes may have responded to the presence of the latter by partially specializing on male production. Hence, these populations of D. magna may be a model for an evolutionary transition from a purely environmental to a partially genetic sex determination system.     

Reproductive Ecology and Cub Survival of Florida Black Bears

Abstract: We investigated reproductive ecology and cub survival of Florida black bears (Ursus americanus floridanus) in Ocala National Forest and the adjacent residential area of Lynne, Florida, USA, 1999-2003. We documented production of 81 cubs from 39 litters. Average litter size was 2.08 ± 0.11 (SE) cubs. The mean age of first reproduction was 3.25 ± 0.27 years. Excluding females that reproduced in consecutive years due to litter loss, interlitter interval was 2.11 ± 0.11 years. The mean annual fecundity rate was 0.57 ± 0.06. We used expandable radiocollars to monitor the fate of 41 bear cubs. The probability of cubs surviving to 9 months of age was 0.46 ± 0.09 and did not differ between cohorts or study locations. The most important causes of cub mortality included infanticide and mortality caused directly or indirectly by collisions with vehicles. Our results indicate that reproductive rates of female black bears in the Ocala study area are comparable to those reported for other black bear populations from eastern United States, but cub survival rates are lower than those reported for most black bear populations. Management of Florida black bears should emphasize strategies to reduce the mortality of cubs.     

Individual heterogeneity in life-history trade-offs with age at first reproduction in capital breeding elephant seals

Recruitment age plays a key role in life-history evolution. Because individuals allocate limited resources among competing life-history functions, theory predicts trade-offs between current reproduction and future growth, survival and/or reproduction. Reproductive costs tend to vary with recruitment age, but may also be overridden by fixed individual differences leading to persistent demographic heterogeneity and positive covariation among demographic traits at the population level. We tested for evidence of intra- and inter-generational trade-offs and individual heterogeneity relating to age at first reproduction using three decades of detailed individual life-history data of 6,439 capital breeding female southern elephant seals. Contrary to the predictions from trade-off hypotheses, we found that recruitment at an early age was associated with higher population level survival and subsequent breeding probabilities. Nonetheless, a survival cost of first reproduction was evident at the population level, as first-time breeders always had lower survival probabilities than prebreeders and experienced breeders of the same age. However, models accounting for hidden persistent demographic heterogeneity revealed that the trade-off between first reproduction and survival was only expressed in “low quality” individuals, comprising 35% of the population. The short-term somatic costs associated with breeding at an early age had no effect on the ability of females to allocate resources to offspring in the next breeding season. Our results provide strong evidence for individual heterogeneity in the life-history trajectories of female elephant seals. By explicitly modeling hidden persistent demographic heterogeneity we show that individual heterogeneity governs the expression of trade-offs with first reproduction in elephant seals.     

Age and size-dependent variation in the seasonal timing and probability of reproduction among mature female pumpkinseed,Lepomis gibbosus

Synopsis In this study, we used pumpkinseed,Lepomis gibbosus, populations in two east-central Ontario lakes to test for age and size-dependent effects on the probability and timing of reproduction of mature females. Pumpkinseed body size characteristics differed in the two lakes; Little Round Lake harbours a stunted population and Beloporine Lake does not. Age-class distribution of mature females was determined by a maturity assessment on fish collected just prior to spawning and at the mid-spawning period, combined with an early-season capture-mark-recapture survey. Spawning females were collected throughout the breeding season to assess age and size-related temporal trends, and to compare their age and size distribution with that of mature females at large. The proportion of age 2 females in Little Round Lake that actually spawned was considerably lower than the proportion of mature age 2 females at large. Furthermore, age 2 females that spawned in this lake did not do so until late in the breeding season. In contrast, the proportion of young/small females spawning in Beloporine Lake was comparable to the proportion of young/small mature females at large, and both small and large females spawned throughout the breeding season. Small mature females in Little Round Lake may have had no other option but to spawn late in the season because of their poor body condition. In Beloporine Lake, condition factor early in the breeding season in age 2–4 females was higher than that of Little Round Lake females, suggesting that limited energy reserves in the spring may have prevented young Little Round Lake females from early spawning. Our results show that the likelihood and timing of reproduction are both age and size-dependent in some populations. Small individuals that delay seasonal maturation and spawn late in the summer probably contribute little to the population due to the restricted growth and reduced overwinter survival of their progeny.To whom correspondence should be addressed     

Determinants of local recruitment in a growing colony of Audouin's gull

1. Local recruitment of Audouin's gull ( Larus audouinii Payraudeau) was studied between 1988 and 1997 at the Ebro Delta colony (north-western Mediterranean). Since its establishment in 1981, the colony has dramatically grown to include, in 1997, 65% of the total world population. Several hypotheses were tested, involving the effects of a badger predatory event in 1994, and sex, age and cohort (year of birth) on recruitment.
2. Results supported the prediction that colony size influenced recruitment: the probability for any individual to have previously bred increased throughout the study, together with colony size. At the end of the study, 90% of breeders aged 4 years had already been recruited at age 3, the age of first reproduction by Audouin's gulls. As expected by the dramatic increase of breeding numbers, most local recruitment occurred at very young ages, especially when compared with other Laridae.
3. Neither food availability nor reproductive success affected recruitment. Recruitment was not affected by high nest predation by the badger, although after the event, the proportion of Ebro Delta birds nesting on the nearby Columbretes Islands tripled.
4. Probability of first reproduction depended on age: it was the highest at ages 3 and 4, and then decreased sharply with age to stabilize beyond age 6 to a value depending on the year and cohort but always very low (< 5%). Cohort and sex did not influence local recruitment.
5. Annual resighting rates ranged between 35% and 82%, and were higher for females. This may represent a sex-dependent suspension of breeding, probably as a trade-off between early recruitment and future survival.     

Testing generalizations about latitudinal variation in reproduction and recruitment patterns with sicyoniid and caridean shrimp species

Summary

Study of latitudinal variation in seasonality of reproduction and recruitment of benthic marine invertebrates is useful in generating and testing hypotheses about causal factors acting on reproduction such as temperature and larval food supply that might be altered by changes in world climate. Analysis of latitudinal variation in reproductive patterns might be made with comparisons (a) among species with a common phylogenetic history from different latitudes and habitats and (b) among phylogenetically different taxa from the same location. Hypotheses on variation of reproductive seasonality with latitude are tested here with results of a study on nine species of caridean and two species of sicyoniid shrimp sampled from a tropical seagrass meadow in Puerto Rico. Breeding condition was determined by the presence or absence of incubated embryos (carideans) and the state of ovarian development in both carideans and sicyoniids. Recruitment was estimated from the percentage of individuals of monthly population samples in the juvenile size classes. Comparison of reproductive patterns among tropical, subtropical, and cool temperate Sicyonia spp. supports the paradigm of continuous reproduction in the tropics with increased restriction of breeding season with an increase in latitude. A greater intensity of breeding effort appears to accompany the shorter breeding period associated with an increase in latitude. At the tropical site most females of all caridean species carried embryos during all months of the year. With the onset of sexual maturity, caridean females produced consecutive broods for the rest of their relatively short (< 6 month) life span. In both sicyoniid and caridean species, recruitment occurred throughout the year but was highly variable, i.e., episodic rather than truly continuous or seasonal. Patterns of recruitment were highly concordant among but not between sicyoniid and caridean species, indicating that different sets of environmental factors controlled recruitment in the two groups. It is suggested that simultaneous study of adult reproduction and larval ecology is necessary to understand patterns of reproduction and recruitment. Coordinated effort on a global scale in studying latitudinal variation in reproduction and recruitment is suggested in order to predict the consequences of climate change on commercially and ecologically important marine invertebrate species.     

Parental age and lifespan influence offspring recruitment: a long-term study in a seabird

Recent studies of wild populations provide compelling evidence that survival and reproduction decrease with age because of senescence, a decline in functional capacities at old ages. However, in the wild, little is known about effects of parental senescence on offspring quality. We used data from a 21-year study to examine the role of parental age on offspring probability of recruitment in a long-lived bird, the blue-footed booby (Sula nebouxii). Offspring probability of recruiting into the breeding population varied over the life of parents and effects age were similar in mothers and fathers. Offspring recruitment was high when parents were roughly 6-12 years old and low before and after then. Effects of parental age on offspring recruitment varied with lifespan (parental age at last reproduction) and previous breeding experience. Offspring recruitment from young and old parents with long reproductive lifespans was greater than that of offspring from parents with short lifespans at young and old ages. For parents with little previous breeding experience recruitment of offspring decreased with their hatch date, but experienced parents were no similarly affected. We found evidence of terminal effects on offspring recruitment in young parents but not in older parents, suggesting that senescence is more likely a gradual process of deterioration than a process of terminal illness. Failure to recruit probably reflects mortality during the first years after independence but also during the fledgling transition to full independence. Our results show effects of parental age and quality on offspring viability in a long-lived wild vertebrate and support the idea that wild populations are composed of individuals of different quality, and that this individual heterogeneity can influence the dynamics of age-structured populations.     

Annual changes in serum sex steroids in male and female black (Ursus americanus) and polar (Ursus maritimus) bears

The adaptation of black and polar bears to their environments is proportional to the severity of climate and food restriction. Both black and polar bears mate during the spring, despite differences in their recent metabolic state. Reproductive activity in black bears follows 4 mo of torpor, whereas reproduction in polar bears occurs prior to torpor. The goals of this study were to measure the annual changes in serum sex steroids in male and female black and polar bears, and to determine if changes in serum levels of these steroids were associated with metabolic condition or photoperiod. Serum testosterone (T) concentrations were elevated during spring in black and polar bears. Moreover, this increase in serum T in polar bears during spring was correlated with age and testis size. Serum progesterone (P4) concentrations increased in pregnant polar bears in fall coincident with the time of expected implantation. No increases in serum P4 were observed in nonpregnant black and polar bears. Serum estradiol (E2) was elevated in nonpregnant and pregnant polar bears 2 mo prior to the time of expected implantation. We found that serum sex steroids measured in black and polar bears change independent of torpor. Therefore, our results suggest that photoperiod may be a more important regulator of serum steroid levels and reproduction than metabolic condition.