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.     

Breeding season, litter size and nestling mortality of the Harvest mouse, Micromys minutus (Rodentia: Muridae), in Britain

Data are presented on the breeding biology of Micromys minutus in Britain. It is shown that although litters are produced over a protracted period (May-December), 74% of the litters are born in August and September. Mean litter size is 5·40±0·16( n = 62), which is a significant reduction on the mean litter size of 6·75±0·40( n = 16) calculated from pre-1917 literature. Litter size was found to be constant throughout the breeding season, and there was no difference in mean litter size between two samples collected close to the northern and southern limits of the known British range. By comparing the mean litter size of samples 1–5, 6–10 and 11+ days old, no significant loss of individual mice within litters could be detected, although 12·3% of all litters died prior to weaning. This loss was predominantly of autumnal litters, cold and wet being important climatic factors that terminate the breeding season. Comparisons are made with data from other parts of the animal's range.     

The population ecology of Rattus exulans on Tiritiri Matangi Island,and a model of comparative population dynamics in New Zealand

Abstract

On Tiritiri, a small predator-free island in northern New Zealand, kiore (Rattus exulans) were live and snap trapped in grassland and forest. In both habitats, kiore abundance peaked in late summer/autumn. The increase followed a 3 month breeding season during which females produced two to three litters, each averaging 7 young. During the population decline in autumn and winter, animals lost weight. Few bred in the breeding season of their birth and none lived to breed in a second breeding season, so the population consisted of distinct age cohorts. These patterns may relate to a highly seasonal food supply.

Kiore elsewhere in New Zealand show seasonal breeding, but the length of breeding, sexual maturation, and litter size vary. Other studies of kiore in the Pacific show less marked seasonal fluctuations, longer breeding seasons, and smaller litters. We propose a model to explain the variation in rodent demography in New Zealand. The model is based on the seasonal availability of food, along with the modifying influences of predation and dispersal.     

Reproduction of plateau pika (<Emphasis Type="BoldItalic">Ochotona curzoniae</Emphasis>) on the Qinghai–Tibetan plateau

The reproduction of the plateau pika (Ochotona curzoniae) was investigated in Guoluo District at an elevation of 4,000 m on the Qinghai–Tibetan plateau, China, from April 2007 to August 2008. Reproduction was seasonal, and the breeding season lasted from April to late June/early July. Adults produced two litters in each year, and the mean litter size, estimated by counting the number of embryos in utero, was 3.3 ± 0.1 (n = 52) in 2007 and 3.2 ± 0.1 (n = 66) in 2008. The timing of reproduction showed high inter-annual variation; lower precipitation and the concomitant delay in spring vegetation phenology may have retarded the onset of the breeding season in 2007 compared with 2008. The most frequent litter sizes were 3 and 4, which together comprised 71.2% and 83.3% of litters in May and June of 2007 and 2008, respectively. Compared with previous studies, reproduction was highly variable between geographic areas. Pikas produced between one and five litters per year in different regions of the plateau over different breeding seasons. This geographic and inter-annual variation appeared to be associated with the duration of plant growth at each site, suggesting that plateau pikas adjust the length of their breeding season to match the period when sufficient energy is available to support the high energy demands of reproduction.     

Changes in population structure and reproduction during a 3-yr population cycle of voles

Cyclic changes in population growth rate are caused by changes in survival and/or reproductive rate. To find out whether cyclic changes in reproduction are an important part of the mechanism causing cyclic fluctuations in small mammal populations, we studied changes in the population structure and reproduction of field voles ( Microtus agrestis ), sibling voles ( M. rossiaemeridionalis ), bank voles ( Clethrionomys glareolus ), and common shrews ( Sorex araneus ) in western Finland during 1984–1992, in an area with 3-yr vole cycles. We also modelled the population growth of voles using parameter values from this study. The animals studied were collected by snap trapping in April, May, June, August, September, and, during 1986–1990, also in October. We found several phase-related differences in the population structure (age structure, sex ratio, proportion of mature individuals) and reproduction (litter size, length of the breeding season) of voles. In non-cyclic common shrews, the only significant phase-related difference was a lower proportion of overwintered individuals in the increase phase. According to the analyses and the vole model, phase-related changes in litter size had only a minor impact on population growth rate. The same was true for winter breeding in the increase phase. The length and intensity of the summer breeding season had an effect on yearly population growth but this impact was relatively weak compared to the effect of cyclic changes in survival. The population increase rates of Microtus were delayed dependent on density (8–12-month time lag). Our results indicate that cyclic changes in reproduction are not an important part of the mechanism driving cyclic fluctuations in vole populations. Low survival of young individuals appeared to play an important role in the shift from the peak to the decline phase in late summer and early autumn.     

Reproduction of the bank vole, Clethrionomys glareolus, in northern and southern Sweden during several seasons and in different phases of the vole population cycle

The breeding biology of the bank vole was studied in northern and southern populations in Sweden in the years 1973–76. Trapped specimens supplied organ weights and histological data, which were utilized to describe the breeding season, number and size of litters, reproductive losses, spermatogenesis, and sexual maturation. In the north the populations are cyclic. During the increase phase of the cycle the breeding season was long, litter sizes and gonad weights were large, and there was maturation of yearlings in the summer. During the peak phase reproduction was impaired in all these respects. In southern Sweden there was no cyclicity, and the reproductive events varied less, but in 1976 the litters were larger than usual and the season was much extended.
In the south the voles born during the early part of the season regularly became sexually mature during their first summer. The differences in reproductive biology are related to population dynamics and discussed in terms of different hypotheses concerning vole population cyclicity.     

Increasing returns in the life history of Columbian ground squirrels

1. We examined positive associations and trade-offs of maternal and reproductive traits in a population of Columbian ground squirrels, Spermophilus columbianus .
2. Structural size, body condition, mother's personal allocation to body mass during reproduction, and timing of littering were estimated for live-trapped reproductive females that were observed during an 8-year period, and were compared to litter mass, litter size, and average pup mass using path analyses.
3. Mothers exhibited age-structured traits that influenced reproductive patterns. Yearling mothers were significantly smaller, bred later, and had smaller litters than older females. Mothers that gained more body mass during reproduction and older mothers in good body condition that were structurally large had larger litters.
4. Early seasonal timing of littering was an important positive influence on successful reproduction by older mothers only in early breeding seasons and in years when conditions for reproduction were good for all females.
5. The number of offspring that survived to 1 year of age was most strongly associated with litter mass and litter size; date of breeding was of secondary influence, with earlier litters exhibiting greater success.
6. In general, mothers that gained the most in body mass during reproduction were concurrently more successful in weaning large litters (perhaps due to better quality of foraging habitat).
7. In addition to expected reproductive trade-offs, reproduction by Columbian ground squirrels exhibited positive associations of life-history traits that may reflect evolutionary increasing returns.     

Variation in reproductive investment among and within populations of the scorpion Centruroides vittatus

Although information concerning variation among and within populations is essential to understanding an organism's life history, little is known of such variation in any species of scorpion. We show that reproductive investment by the scorpion Centruroides vittatus varied among three Texas populations during one reproductive season. Females from the Kickapoo population produced smaller offspring and larger litters than females from the Independence Creek or Decatur populations; this pattern remained when adjusting for among population variation in either female mass or total litter mass. Relative clutch mass (RCM) and within-litter variability in offspring mass (V*) did not differ among populations. Among-population variation may result from genetic differences or from phenotypically plastic responses to differing environments. Within populations, the interrelationships among reproductive variables were similar for Decatur and Independence Creek: females investing more in reproduction (measured by total litter mass, TLM) produced larger litters and larger offspring, and V* decreased with increased mean offspring mass (and with decreased litter size at Decatur). At Kickapoo, larger females produced larger litters and had larger TLM; females investing more in reproduction produced larger litters but not larger offspring. Within litter variability in offspring mass was not correlated with any reproductive variables in this latter population. These patterns may be explained by the fractional clutch hypothesis, the inability of females precisely to control investment among offspring or morphological constraints on reproduction.     

Dominance relations in captive groups of adult and juvenile arctic blue foxes (Alopex lagopus)

Dominance relationships were studied in captive Arctic blue fox (Alopex lagopus) groups comprising adults (four males, five females) and juveniles (four males, five females). The results showed that Arctic blue foxes easily formed a social organization with an observable hierarchy, in which adults typically dominated over juveniles. Within the same age group, males usually dominated over females. Dominance correlated most significantly with body weight in autumn, but later that correlation decreased. Urine marking activity was very low during autumn and early winter, but increased significantly prior to and during the breeding season when aggressive encounters were also most pronounced. In addition to several adults, the social status of some juveniles was high at breeding time. Altogether 7 out of 11 females (63.6%) whelped, but the survival rate of litters was low and kits of only two adult females survived (18.2%). It can be concluded that hierarchical development and reproduction in Arctic blue fox groups are markedly influenced by dominance relationships.     

Effect of indomethacin on reproduction under laboratory and field conditions in deermice (Peromyscus maniculatus)

The purpose of this investigation was to determine the long-term effect of enterically administered indomethacin (a prostaglandin synthetase inhibitor) on reproduction in Peromyscus maniculatus (deermice) under both laboratory and field conditions. Indomethacin-coated wheat (1 mg/kg wheat) was provided to the experimental animals in the laboratory and under field conditions ad libitum. Rat chow was also provided to the animals in the laboratory. The laboratory data indicate that the number and size of the litters from the control breeding pairs were larger (number of litters=22; average number of pups/litter=4.9) than the experimental breeding pairs (number of litters=2; average number of pups/litter=2.5) over a 100-day experimental period. Field data indicate that the number of pregnant animals, the total number of animals, and mean juvenile males and females live-trapped decreased in the experimental group with respect to the control group during the experiment. The data suggest that deermice will consume indomethacin under both laboratory and field conditions, and that it effectively inhibits reproduction on a long-term basis. It is, therefore, possible that prostaglandin synthetase inhibitors may be used to control populations of rodent pests. Additionally, the data raise questions about the effect of long-term use of nonsteroidal analgesic compounds similar to indomethacin on human reproduction.     

Costs of reproduction and terminal investment by females in a semelparous marsupial

Evolutionary explanations for life history diversity are based on the idea of costs of reproduction, particularly on the concept of a trade-off between age-specific reproduction and parental survival, and between expenditure on current and future offspring. Such trade-offs are often difficult to detect in population studies of wild mammals. Terminal investment theory predicts that reproductive effort by older parents should increase, because individual offspring become more valuable to parents as the conflict between current versus potential future offspring declines with age. In order to demonstrate this phenomenon in females, there must be an increase in maternal expenditure on offspring with age, imposing a fitness cost on the mother. Clear evidence of both the expenditure and fitness cost components has rarely been found. In this study, we quantify costs of reproduction throughout the lifespan of female antechinuses. Antechinuses are nocturnal, insectivorous, forest-dwelling small (20-40 g) marsupials, which nest in tree hollows. They have a single synchronized mating season of around three weeks, which occurs on predictable dates each year in a population. Females produce only one litter per year. Unlike almost all other mammals, all males, and in the smaller species, most females are semelparous. We show that increased allocation to current reproduction reduces maternal survival, and that offspring growth and survival in the first breeding season is traded-off with performance of the second litter in iteroparous females. In iteroparous females, increased allocation to second litters is associated with severe weight loss in late lactation and post-lactation death of mothers, but increased offspring growth in late lactation and survival to weaning. These findings are consistent with terminal investment. Iteroparity did not increase lifetime reproductive success, indicating that terminal investment in the first breeding season at the expense of maternal survival (i.e. semelparity) is likely to be advantageous for females.     

Seasonal variation in fertility, fecundity and litter sex ratio in laboratory and wild stocks of house mice (Mus domesticus)

Data compilations were made for three parameters pertaining to reproduction in a domestic strain (14 years) and a wild stock (4 years) of commensal house mice: (a) the percentage of females mated that produced litters; (b) the average number of pups per litter; and (c) the sex ratio of the pups in the litters. Fecundity and fertility varied seasonally in both domestic and wild mice. More females become pregnant and litter sizes were larger in the spring, summer, and fall months than during the winter season. Sex ratios also varied seasonally with more male biased litters produced during the 4 winter months. There appear to be seasonal shifts in productivity for both stocks of mice and these seasonal trends have not been altered by domestication under laboratory conditions. In spite of the fact that house mice are generally opportunistic, it is possible that there has been selection in the mice for shifting rates of production in relation to the best seasons of the year in terms of climate and resource availability. The higher production of males during summer months may be geared toward greater success in dispersal at that season and a higher probability that the males can find a territory and mate successfully in summer rather than winter. These results have potential implications for animal breeders and for those who maintain mouse colonies to produce animals for scientific investigations.     

Elevation induced variation in the breeding traits of a nectar-feeding non-flying mammal

Factors that influence reproduction in nectar-feeding non-flying mammals are poorly described. We investigated factors that may influence the breeding traits of the eastern pygmy-possum (Cercartetus nanus), a small (25 g) nectar-feeding marsupial from eastern Australia. Females at our coastal site produced 1–3 litters (frequently of 5–6 young) over an 8-month period within the flowering period of the dominant food plant (Banksia ericifolia). The number of lactating females over time was highly correlated (R?=?0.9) with the abundance of flowers on B. ericifolia, suggesting that flower availability has a substantial influence on breeding. To assess the generality of these findings and investigate whether elevation or latitude influence breeding in this species, we examined the breeding traits previously described at three other locations. Females on the tableland 70 km away produced one litter of 3–4 young over a 4-month period within the flowering period of B. ericifolia, the dominant food plant. At other coastal and tableland locations 500 km away, coastal females produced larger numbered litters more often than tableland females. A hypothesis relating to minimum temperature appears the most plausible explanation for this pattern that reflects elevation but not latitude. The mean minimum temperature drops below 5 °C for 2–5 months each year at the tableland locations but not at all at the coastal locations. Low temperatures are known to reduce nectar secretion in the dominant food plants. Thus, although the breeding traits of the eastern pygmy-possum are influenced by flowering in their dominant food plants, low temperature appears to impose a constraint on reproductive output.     

Numbers of litters, litter size and survival in two species of microtines at two elevations

The number of litters, litter size and survival of Clethrionomys gapperi and Microtus pennsylvanicus were documented at two elevations (1450 m; 2240 m) in the Canadian Rocky Mountains to determine if those life history traits changed with the length of the breeding season. The high elevation was colder, had fewer frost-free days, and more precipitation than the low elevation. The length of the breeding season of C. gapperi and M. pennsylvanicus averaged 8 d and 34 d longer, respectively, at the low elevation than at the high elevation. In both species and at both elevations, most females survived only long enough to produce one or two litters. Litter size was significantly greater at the high elevation than at the low elevation in M. pennsylvanicus , but not different in C. gapperi. Overall, nestling survival was greater at the low elevation than at the high elevation in M. pennsylvanicus , but not different in C. gapperi. Summer and winter survival rates were not different between elevations in either species. Differences in litter size and nestling survival between elevations in M. pennsylvanicus may be related to differences in habitat quality.     

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.     

Testing the reproductive and somatic trade‐off in female Columbian ground squirrels

Energetic trade‐offs in resource allocation form the basis of life‐history theory, which predicts that reproductive allocation in a given season should negatively affect future reproduction or individual survival. We examined how allocation of resources differed between successful and unsuccessful breeding female Columbian ground squirrels to discern any effects of resource allocation on reproductive and somatic efforts. We compared the survival rates, subsequent reprodction, and mass gain of successful breeders (females that successfully weaned young) and unsuccessful breeders (females that failed to give birth or wean young) and investigated “carryover” effects to the next year. Starting capital was an important factor influencing whether successful reproduction was initiated or not, as females with the lowest spring emergence masses did not give birth to a litter in that year. Females that were successful and unsuccessful at breeding in one year, however, were equally likely to be successful breeders in the next year and at very similar litter sizes. Although successful and unsuccessful breeding females showed no difference in over winter survival, females that failed to wean a litter gained additional mass during the season when they failed. The next year, those females had increased energy “capital” in the spring, leading to larger litter sizes. Columbian ground squirrels appear to act as income breeders that also rely on stored capital to increase their propensity for future reproduction. Failed breeders in one year “prepare” for future reproduction by accumulating additional mass, which is “carried over” to the subsequent reproductive season.     

The colony structure and reproductive biology of the afrotropical Mashona mole-rat, Cryptomys darlingi

Cryptomys darlingi occurs in the mesic Miombo woodland of north-eastem Zimbabwe. It occurs in colonies of up to nine individuals, in which reproduction is limited to one of the largest males and the largest female in the colony.
Reproduction and details of colony size and number of breeding animals in a colony are described for five complete field-captured colonies.
In captivity, mating is not confined to a particular season, and up to three litters of pups are orn per annum. The reproductive female initiates the pre-copulatory behaviour. The gestation lengti is 56–61 days ( n = 2 ). The new-born pups are altricial and litter size is small = 1.7 ± 0.5 ( n = 6). In this case, the pups first left the nest 10 days after birth, began to eat solids when 14 days old, and were fully weaned at five weeks. They began to spar with each other when 36–40 days old, but did not disperse and were incorporated into the colony. This suggests that the Mashona mole-rat colonies are composed of a founding pair and at least three successive litters of pups.     

高原鼠兔(Ochotona curzoniae)冬季自然死亡率

以耳标观察法。1985年4-7月共捕标了样地内高原鼠兔(Ochotona curzoniae)幼鼠114只。到入冬前,死亡率为50.88%。第2胎出生的鼠死亡率明显地高于第1胎的死亡率。冬季死亡率呈现波动,入冬及开春时死亡率较高。鼠兔种群进入繁殖前期死亡率趋于零。初春样区存活标志鼠6只,经夏秋冬3季总计死亡率达94.74%。如按入冬时实有标志鼠计算,冬季死亡率为91.04%。     

Breeding and juvenile survival among slender-tailed meerkats (Suricatu suricatta) in the south-western Kalahari: ecological and social influences

Reproductive success of co-operatively-breeding slender-tailed meerkats ( Suricata suricatta ) in the Kalahari was monitored over four breeding seasons and 26 band years, concentrating on three focal bands. Breeding was strongly seasonal, with peak numbers of births coinciding with maximum rainfall between January and March. Breeding seasons were extended, and more litters were produced, during the years with above average rainfall. For dominant females, the mean annual rate of litter production was 1.9 ± 0.8 litters per year and short interbirth intervals (mean = 90 %pL 18 days) indicated that females came into oestrus within three weeks of giving birth. Births were synchronous within but not between bands. Major known causes of kitten mortality were cold weather and predation, and losses mainly occurred between three and five weeks of age. Sixty-seven percent of juveniles survived between emergence from the den at three or four weeks old and attainment of effective foraging independence at 12 weeks. There were no significant effects of birth date within the season upon litter size or survival. Several incidents of apparent infanticide were recorded. These meerkats were highly co-operative and adult band members assiduously guarded and provisioned the kittens. Nevertheless, regression analysis showed that neither rainfall (an index of prey availability) nor band size variables accounted for much variance in juvenile survival to 12 weeks, which appeared to be heavily influenced by chance events such as flash floods. In contrast, rainfall between January and March had a significantly positive effect on the total number of juveniles produced during the breeding year.     

Body size,litter size,timing of reproduction,and juvenile survival in the Unita ground squirrel,Spermophilus armatus

The timing of reproduction affected litter size, offspring mass, and offspring survival in the Uinta ground squirrel, Spermophilus armatus, in Grand Teton National Park, Wyoming. Survival of juvenile females to yearling age varied negatively with date of weaning and positively with individual offspring mass. At the same time, juveniles weaned early in the season were lighter, and juveniles weaned later in the season were heavier. The coefficient of variation for juvenile body mass, originally measured at weaning, significantly decreased by the time juveniles entered hibernation, indicating that individuals weaned early and light caught up in body mass to individuals weaned later and heavier. From the perspective of the mother's investment in the litter, litter size (corrected for mother's mass) decreased with later wcaning dates, while the relationship of weaning date to litter mass (corrected for mother's mass) was significant in only one year. Maternal allocation of resources in litters changed over the season so that mothers produced many, small offspring early in the season, and fewer, large offspring late in the season.