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.     

Senescence and costs of reproduction in the life history of a small precocial species

Species following a fast life history are expected to express fitness costs mainly as increased mortality, while slow‐lived species should suffer fertility costs. Because observational studies have limited power to disentangle intrinsic and extrinsic factors influencing senescence, we manipulated reproductive effort experimentally in the cavy (Cavia aperea) which produces extremely precocial young. We created two experimental groups: One was allowed continuous reproduction (CR) and the other intermittent reproduction (IR) by removing males at regular intervals. We predicted that the CR females should senesce (and die) earlier and produce either fewer and/or smaller, slower growing offspring per litter than those of the IR group. CR females had 16% more litters during three years than IR females. CR females increased mass and body condition more steeply and both remained higher until the experiment ended. Female survival showed no group difference. Reproductive senescence in litter size, litter mass, and reproductive effort (litter mass/maternal mass) began after about 600 days and was slightly stronger in CR than IR females. Litter size, litter mass, and offspring survival declined with maternal age and were influenced by seasonality. IR females decreased reproductive effort less during cold seasons and only at higher age than CR females. Nevertheless, offspring winter mortality was higher in IR females. Our results show small costs of reproduction despite high reproductive effort, suggesting that under ad libitum food conditions costs depend largely on internal regulation of allocation decisions.     

On the virtue of being the first born: the influence of date of birth on fitness in the mosquitofish, Gambusia affinis

We found in an earlier study that mosquitofish (Gambusia affinis and G. holbrooki) ceased reproduction in the late summer, long before the end of warm weather, stored fat, then utilized reserves to survive the winter and initiate reproduction the following spring. We hypothesized that this pattern of fat utilization was a life history adaptation that enabled the fish to acquire food resources in the autumn then allocate them to reproduction the following spring when the fitness of the young would be greater. Here we evaluate one aspect of this hypothesis by evaluating the probability of survival to maturity and fecundity of young as a function of date of birth. We placed cohorts comprising eight to ten litters of young born early‐, mid‐ or late in the reproductive season in replicate field enclosures. The entire experiment was repeated in two different years. Early‐born young had a significantly higher probability of survival to maturity but did not differ in fecundity relative to the last cohort of the season. Early‐born young also attained maturity early enough to reproduce in their year of birth while late‐born young had to overwinter before reproduction. The fitness consequences to the mother of either producing one more litter of young at the end of the season, versus instead storing fat and reproducing the following spring are not as determinate as are the effects of date of birth on offspring fitness. Females most often gain fitness by not producing one last litter and instead over‐wintering. If, however, the overwinter survival of offspring is not influenced by their size at the end of the season, then a female's fitness could be enhanced by producing one more litter late in the season. If instead the probability of overwinter survival is strongly influenced by the size of offspring at the end of the season, then our results suggest that a female gains more by deferring reproduction and storing for overwinter survival and reproduction the following spring.     

Variations in litter size and reproductive effort within and between some populations of Lacerta vivipara

We studied the relationships between litter size, litter weight, newborn weight, relative clutch mass and the female snout-vent length in some Lacerta vivipara populations over a period of three years.
Litter size and litter weight were positively correlated with female snout-vent length in all the populations for all the years, as in most other lizard species. Relative clutch mass generally increased with female size, though correlations appear not to be very tight.
Considering the two best studied populations suggests that montane females invest less in reproduction than lowland ones.
The main reproductive traits of the species appeared highly variable between as well as within the different populations hitherto studied.
We argue that current theory about lizard reproductive strategy requires, first to work out a good estimate of reproductive effort, and second to get more information about the relations between the species and their environmental, biotic and abiotic conditions.     

The effects of size, age and a cost of early breeding on reproduction in female mountain hares

Both age and size may influence female reproductive performance in mammals, and successful early reproduction may lead to reduced success at later attempts. The effects of age, size and early reproduction on distribution of reproductive effort throughout a single breeding season was examined in female mountains hares Lepus timidus L. Hind foot length was used as an index of body size, because, unlike body weight, it did not fluctuate with reproductive status. Fifty-six female carcasses were collected from March to October 1984, and their litters were assigned to one of three chronologically equal'litter periods'(1–3) of equal length. Whereas number of ova shed was always independent of age, large females shed more ova than did smaller females in litter periods 1 and 2. Prenatal mortality of ova and embryos was highest during litter period 1, when it was independent of age and size. Although prenatal mortality remained high in first year females in litter period 2, there was an overall decline through to the final litter period when it was negligible. Total number of young produced through the season increased with skeletal size in old females (age > 1), but not significantly in first year females. It is concluded that large size, rather than age, favours early reproduction in mountain hares. Every additional offspring produced in litter periods 1 and 2 reduced that female's production in period 3. After correcting for this cost of early reproduction the number of young produced in the final litter period also increased with maternal size.     

Short-term versus long-term effects of food intake on reproductive output in a viviparous snake, Vipera aspis

Feeding rates influence reproductive output in many kinds of animals, but we need to understand the timescale of this influence before we can compare reproductive energy allocation to energy intake. A central issue is the extent to which reproduction is fuelled by long-term energy stores ("capital" breeding) versus recently-acquired resources ("income" breeding). Our data on free-living aspic vipers show that there is no simple answer to this question: reproductive frequency is determined by long-term energy stores, offspring size is influenced by maternal food intake immediately before ovulation, and litter size is influenced by both long-term stores and short-term energy acquisition. Thus, offspring size in free-living vipers reflects the mother's energy balance over the preceding year (via a trade-off between litter size and offspring size) as well as her energy balance in the current breeding season. Hence, different components of a given reproductive output (litter) are not only functionally linked, but also respond to different temporal scales of prey availability. A female's body size has little effect on her reproductive output. Attempts to quantify reproductive energy allocation must take into account the fact that different reproductive traits (such as offspring size versus number) may respond to energy availability over different timespans. Thus, although the aspic viper is a typical "capital breeder" in terms of its reliance on stored reserves for maternal "decisions" concerning reproductive frequency, it is to some degree a facultative "income breeder" with respect to the determination of offspring size and litter size.     

Correlates of seasonal change in the body condition of an Arctic top predator

Climate‐driven sea ice loss has led to changes in the timing of key biological events in the Arctic, however, the consequences and rate of these changes remain largely unknown. Polar bears (Ursus maritimus) undergo seasonal changes in energy stores in relation to foraging opportunities and habitat conditions. Declining sea ice has been linked to reduced body condition in some subpopulations, however, the specific timing and duration of the feeding period when bears acquire most of their energy stores and its relationship to the timing of ice break‐up is poorly understood. We used community‐based sampling to investigate seasonality in body condition (energy stores) of polar bears in Nunavut, Canada, and examined the influence of sea ice variables. We used adipose tissue lipid content as an index of body condition for 1,206 polar bears harvested from 2010–2017 across five subpopulations with varying seasonal ice conditions: Baffin Bay (October–August), Davis Strait and Foxe Basin (year‐round), Gulf of Boothia and Lancaster Sound (August–May). Similar seasonal patterns were found in body condition across subpopulations with bears at their nadir of condition in the spring, followed by fat accumulation past break‐up date and subsequent peak body condition in autumn, indicating that bears are actively foraging in late spring and early summer. Late season feeding implies that even minor advances in the timing of break‐up may have detrimental effects on foraging opportunities, body condition, and subsequent reproduction and survival. The magnitude of seasonal changes in body condition varied across the study area, presumably driven by local environmental conditions. Our results demonstrate how community‐based monitoring of polar bears can reveal population‐level responses to climate warming in advance of detectable demographic change. Our data on the seasonal timing of polar bear foraging and energy storage should inform predictive models of the effects of climate‐mediated sea ice loss.     

Seasonal and sex differences in the structure and chemical composition of adipose tissue in wild polar bears (Ursus maritimus)

Biopsy samples of adipose tissue from the upper thigh were collected in spring and in late summer/autumn from 370 wild polar bears ( Ursus maritimus ), including adult females with and without cubs, adult males and juveniles. Mean adipocyte volume was measured from all samples and chemical assays of the lipid, total protein and collagen were also performed on samples from 53 bears. Mean adipocyte volume was smaller in all specimens in spring than in late summer/autumn, but the differences were greatest for solitary adult females. The range of adipocyte volumes was much greater for adult females than for adult males, and in females only, mean adipocyte volume correlated significantly with total body mass. Therefore, adipocyte volume measurements from biopsy samples provide some information about fatness in adult females, but are worthless as an indicator of body composition in males and juveniles. In juveniles and females, but not adult males, the lipid content of the adipose tissue was up to 18% lower in autumn than in spring. The collagen content was significantly higher in autumn than in spring in all bears except females with cubs. We suggest that these differences in chemical composition arise from accumulation of water within and between the adipocytes, which would minimize tissue shrinkage, and from changes in the vascularization of the adipose tissue. These properties may be adaptations to rapid fattening and prolonged fasting and the sex differences may reflect the contrasting reproductive strategy of female and male polar bears.     

Reproductive effort during gestation and lactation by Richardson's ground squirrels

Summary Reproductive effort by yearling and older female Richardson's ground squirrels was studied over a 4-year period in southern Alberta by obtaining serial weight records from marked individuals to compare the mother's mass at critical points in the annual cycle (emergence from hibernation, estrus, parturition, and litter emergence) with her litter's mass at birth and weaning. Yearlings weighed only 80% of older adults at emergence from hibernation, but they mated at the same time as older females, attained adult mass coincident with pregnancy, and weaned litters that were not significantly smaller in size or mass than those of older females. Age and maternal mass were weak predictors of litter size and litter mass. Of the net increase in mass of the combined mother-litter unit during gestation, over half (60% of 139 g for yearlings; 52% of 127 g for older females) was attributable to an increase in the mother's own mass, whereas during lactation almost all of the net increase (93% of 545 g for yearlings; 96% of 567 g for older females) was attributable to an increase in the litter's mass. On a daily basis, deposition of mass in the litter was 6 times greater during lactation than gestation. On average, neonates weighed 2.3% (6.5 g) of maternal mass at birth and 23.1% (81 g) at emergence from the natal burrow; offspring masses at birth and at emergence were significantly negatively correlated with litter size. On average, litters weighed 16.3% (48 g) of maternal mass at birth and 157.5% (578 g) at emergence from the natal burrow. Compared with other hibernating sciurids, Richardson's ground squirrels have a similar offspring mass relative to maternal mass both at birth and at emergence from the natal burrow. However, because of the large litter size (typically 6–8), absolute reproductive effort, measured either as litter mass at birth or at natal emergence, is large for the body size of the species.     

Reproductive biology of Philodryas patagoniensis (Snakes: Dipsadidae) in south Brazil: Female reproductive cycle

In this study, we describe the female reproductive cycle of Philodryas patagoniensis in south Brazil, which was described through morpho‐anatomical and histological analyses. The peak of secondary vitellogenesis occurred during winter–spring (July–December), ovulation in spring (October–December), mating and fertilization in spring–summer (October–February), oviposition in spring–autumn (October–May) and births from late spring to autumn (December–July). The diameter of vitellogenic follicles/eggs was larger in winter–spring than in other seasons. The diameter of the shell glands was also larger in winter–spring. In spite of the clear reproductive peak, gonads only showed reduced activity in the autumn. Therefore, at the individual level, females have a discontinuous cyclical reproduction; in the populational level, the reproductive cycle is seasonal semisynchronous. We support the hypothesis that P. patagoniensis have the ability to produce multiple clutches with long‐term stored sperm. Sexual dimorphism in body size was evident, and females are significantly larger and heavier than males. Larger females were able to produce follicles and eggs in larger amount and size. The maternal body size was positively related to the reproductive effort and fecundity. To conclude, we deliberated about the proximal and distal causes that influence the reproductive traits and patterns of P. patagoniensis.     

Fasting physiology of polar bears in relation to environmental change and breeding behavior in the Beaufort Sea

We examined the use of the ratio of serum urea to serum creatinine as a physiological biomarker of fasting to monitor temporal patterns in the feeding ecology of polar bears (Ursus maritimus). Blood was collected from 436 polar bears in the eastern Beaufort Sea during April and May of 1985–1986 and 2005–2006. The proportions of polar bears fasting were 9.6% in 1985, 10.5% in 1986, 21.4% in 2005, and 29.3% in 2006. We used stepwise logistic regression analysis to evaluate factors that could influence the binary response variable of fasting or not fasting. Significant predictor variables of fasting were: the 2005 and 2006 capture years, solitary adult male bears, and adult male bears that were accompanying an estrous female. The increased number of polar bears in a physiological fasting state from all sex, age, and reproductive classes in 2005 and 2006 corresponded with broad scale changes in Arctic sea ice composition, which may have affected prey availability. The higher proportion of adult males fasting from all years was attributed to spring breeding behavior.     

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.     

Assessing stress in Western Hudson Bay polar bears using hair cortisol concentration as a biomarker

The development of novel biomarkers to help assess whether polar bear (Ursus maritimus) health is impacted by long-term physiological stress associated with climate change represents an emerging area of research. Reductions in sea ice cover and food availability are potentially stressful, and chronic stress can have deleterious effects that may impair individual and population level health. Cortisol is the principal effector hormone of the stress response and has previously been linked to aspects of polar bear life history (e.g., reproduction and growth) known to be negatively influenced by environmental change. Understanding stress is important for polar bears at the southern limit of their range, such as those in Western Hudson Bay (WH), where rapidly changing sea ice phenology threatens population viability. We examined the relationships between age, reproductive status, and body condition (fatness) and hair cortisol concentration (HCC) in 729 polar bear hair samples collected in WH from 2004–2013. Overall, there was a negative relationship between fatness and HCC, suggesting that bears in poorer body condition experienced higher levels of stress. However, when reproductive status was included in our analysis, this relationship only held for male and lone female bears. Females with dependent offspring had consistently low fatness and elevated HCC, likely because of the high cost of maternal care. We also found a positive correlation between HCC and age for: (1) bears in poor body condition, possibly due to nutritional stress compounding effects of aging; and (2) male bears, potentially due to stress and injury associated with intrasexual mate competition. These findings support the use of HCC as a biomarker for polar bear health. Furthermore, we have established a HCC benchmark against which future population-level effects of climate change in WH polar bears can be compared.     

The stress of Arctic warming on polar bears

Arctic ecosystems are changing rapidly in response to climate warming. While Arctic mammals are highly evolved to these extreme environments, particularly with respect to their stress axis, some species may have limited capacity to adapt to this change. We examined changes in key components of the stress axis (cortisol and its carrier protein—corticosteroid binding globulin [CBG]) in polar bears (Ursus maritimus) from western Hudson Bay (N = 300) over a 33 year period (1983–2015) during which time the ice‐free period was increasing. Changing sea ice phenology limits spring hunting opportunities and extends the period of onshore fasting. We assessed the response of polar bears to a standardized stressor (helicopter pursuit, darting, and immobilization) during their onshore fasting period (late summer–autumn) and quantified the serum levels of the maximum corticosteroid binding capacity (MCBC) of CBG, the serum protein that binds cortisol strongly, and free cortisol (FC). We quantified bear condition (age, sex, female with cubs or not, fat condition), sea ice (breakup in spring–summer, 1 year lagged freeze‐up in autumn), and duration of fasting until sample collection as well as cumulative impacts of the latter environmental traits from the previous year. Data were separated into “good” years (1983–1990) when conditions were thought to be optimal and “poor” years (1991–2015) when sea ice conditions deteriorated and fasting on land was extended. MCBC explained 39.4% of the variation in the good years, but only 28.1% in the poor ones, using both biological and environmental variables. MCBC levels decreased with age. Changes in FC were complex, but more poorly explained. Counterintuitively, MCBC levels increased with increased time onshore, 1 year lag effects, and in poor ice years. We conclude that MCBC is a biomarker of stress in polar bears and that the changes we document are a consequence of climate warming.     

Explaining the seasonal decline in litter size in European ground squirrels

In European ground squirrels Spermophilus citellus as in many ground squirrel species. late born litters are composed of fewer young than early born litters. Two alternative though not mutually exclusive hypotheses may explain this seasonal pattern of change in litter size. On the one hand. the production of few large young late in the season may be an adaptation to time limitations on the offspring. that have to complete growth and fattening prior to hibernation. Then one would expect a trade-off between offspring number and size as the breeding season progresses. At its extreme. this hypothesis would predict that total maternal effort should be equal independent of litter size. Alternatively. litter size may be determined by physiological limitations on the mother. in that highly constrained mothers breed later and produce smaller litters. Then one would expect reduced overall maternal effort in highly constrained mothers of smaller litters. In this case. a trade-off between litter size and offspring size would not be expected. We found that total maternal effort in terms of gestation length and the duration of lactation increased with increasing litter size. thus supporting the second hypothesis. Lactation was not terminated at natal emergence. It extended a relatively long period of time beyond the time of first litter emergence depending on litter size. During prolonged lactation. individual young of large litters made up body mass to young of small litters. As a consequence. juvenile weaning body mass was unaffected by litter size although offspring body mass at natal emergence was inversely related to litter size. This additional weight gain in young of large litters compensated for initial survival disadvantages and presumably affected fecundity at yearling age.     

Selection of maternity dens by female polar bears in western Hudson Bay,Canada and the effects of human disturbance

We deployed 19 satellite radio collars on adult female polar bears (Ursus maritimus) in western Hudson Bay to study how pregnant females select maternity-den sites and the degree to which human disturbance might influence this process. To eliminate the possibility that being handled while pregnant might influence the final choice of a suitable site, the collars were attached 1 year before the females were expected to select dens. Our results indicated that pregnant females moved inland and selected an area to den shortly after coming ashore in the summer and remained there if not disturbed. We analyzed 24 years of consecutive data (1979–2002) to examine whether disturbance of pregnant females in the denning area in the autumn affected litter size or mass of cubs the following spring. We defined three levels of disturbance (none—bear not seen; low—bear seen but not handled; high—bear seen and handled), and found that disturbance had no effect on either litter size or the mass of male cubs. However, we did find that females handled in the autumn had significantly lighter female cubs than females that were not seen. Our disturbance of pregnant females occurred in August and September, which is only a few months prior to parturition; it is not known what the effects of disturbance might be in late October or early November, closer to the time when cubs are born.     

Seasonal allocation of energy to respiration, growth and reproduction in the freshwater clams, Pisidium variabile and P. compressum (Bivalvia: Pisidiidae)

SUMMARY. 1. The ecological energetics of the pisidiid clams, Pisidium variabile and P. compressum , from permanent ponds in southwestern Ohio were studied from July 1978 to July 1980. Productivity (P; as g C m⁻² yr ⁻¹) was 1.82 (0.80 and 1.02 for the spring and autumn generations) and 4.22 (2.69 and 1.53 for the spring and autumn generations) for P. variabile and P. compressum , respectively.
2. The average biomass (B; as g C m⁻²) of P. variabile was 0.54 (0.30 and 0.24 for the spring and autumn generations) and of P. compressum 1.09 (0.69 and 0.40 for the spring and autumn generations). P:B ratios for the spring and autumn generations of P. variabile were 2.7 and 4.2 and of P. compressum 3.9 and 3.8, which are below those reported for other iteroparous invertebrates.
3. Reproductive effort (Re:Px100) was 5.1 and 5.7 for the generations of P. variabile and 7.7 and 8.8 for the generations of P. compressum. These low values relative to other molluses can be attributed to a viviparous mode of reproduction and exposure to local environmental variability. In addition, the differences between the reproductive efforts of the spring and autumn generations of both species can be ascribed to season of birth and altometric constraints on reproductive capacity.
4. The data from this and related studies on pisidiid clams are used to generate several hypotheses concerning the evolution of life cycles in the family: (1) reproductive effort is limited by allometric constraints on clam size; (2) individuals of different ages can have different demographies and physiological rate functions; and (3) current one-dimensional life history theories are largely inappropriate when dealing with an organism that has a wide range of phenotypic responses to environmental variability as well as complex age structure.     

Reproductive output, maternal age, and survivorship in captive common marmoset females (Callithrix jacchus)

Common marmosets (Callithrix jacchus) demonstrate significant variation in reproductive output on both a yearly and lifetime basis in comparison to other anthropoid primates. We explore the factors that may be most important in determining reproductive variation in captive common marmosets. Studies have found that maternal age, maternal condition, and dam survivorship are related to reproductive output; however, these reports are not in agreement and are far from conclusive. With the use of a large, multicolony, demographic database pooling data across five marmoset colonies, we examined litter information for 1,649 litters, and reproductive summaries for 400 dams to assess 1) how reproductive output variation (total production, total weaned production) is determined by litter size, interbirth interval (IBI), age at first birth, and dam survival age; 2) the relationship between maternal age and reproductive output variables; and 3) relationship between the reproductive output variables and survival. We used stepwise regression procedures to describe the amount of variation in lifetime reproductive output among dams, and found that mean litter size accounted for 18% of the variance in total production, survival age accounted for 10.6%, age at first birth accounted for 8.8%, and mean IBI accounted for 5%. For total (nonzero) weaned production, survival age accounted for 7.6% of variance, age at first birth accounted for 7.2%, mean IBI accounted for 2%, and mean litter size accounted for 1.6%. We identified significant effects (P<0.05) of maternal age on litter size and IBI length, but no effect of dam age on weaned litter size. Cox proportional hazards regression analyses revealed significant effects (P<0.01) of number of litters, age at first birth, and site on dam survivorship. Dams that produced more litters showed higher survivorship. Age at first birth showed a positive relationship with dam survivorship, i.e., dams that delayed first reproduction had higher survival. Our findings about reproductive variation in marmosets may have practical applications for the management of marmoset breeding colonies.     

Age specific fecundity,litter size,and sex ratio in the chacoan peccary (Catagonus wagneri)

Ten years of data on the reproductive biology of the Chacoan peccary (Catagonus wagneri) were analyzed to determine average litter size, sex ratio, timing of births, and individual and age-specific fecundity. Data were obtained from a captive herd of chacoan peccaries located in the western Paraguayan Chaco. Births peak in the austral spring months of September, October, and November, with fewer litters born during the dry season months of June, July, and August. The average litter size was 2.4 with a sex ratio of 56:44 (M:F). There was no significant difference in litter size among individual females or among females of different ages. Finally, there was some individual variation in the age at which sows produced their first litter, but no discernible variation among sows in the average time between litters. Zoo Biol 16:301–307, 1997. © 1997 Wiley-Liss, Inc.     

Reproductive changes in fluctuating house mouse populations in southeastern Australia

House mice (Mus domesticus) in the Victorian mallee region of southeastern Australia show irregular outbreaks. Changes in reproductive output that could potentially drive changes in mouse numbers were assessed from 1982 to 2000. Litter size in females is positively correlated with body size. When standardized to an average size female, litter size changes seasonally from highest in spring to lowest in autumn and winter. Litter size is depressed throughout breeding seasons that begin when the abundance of mice is high, but is similar in breeding seasons over which the abundance of mice increases rapidly or remains low. Breeding begins early and is extended on average by about five weeks during seasons when mouse abundance increases rapidly. The size at which females begin to reproduce is larger during breeding seasons that begin when mouse abundance is high. An extended breeding season that begins early in spring is necessary for the generation of a house mouse plague, but it is not in itself sufficient. Reproductive changes in outbreaks of house mice in Australia are similar but not identical to reproductive changes that accompany rodent population increases in the Northern Hemisphere. We conclude that food quality, particularly protein, is a probable mechanism driving these reproductive changes, but experimental evidence for field populations is conflicting.