Life histories of Clethrionomys and Microtus (Microtinae)

Although there have been numerous life-history reviews of mammals at high taxonomic levels (e.g. among families within orders), there are far fewer studies at lower taxonomic levels (e.g. among species within genera). Data on adult weights, litter size, gestation length, neonate weight, age and weight at weaning, growth rate to weaning, maximum life span, and length of the breeding season were compiled from the literature on five species of Clethrionomys and 33 species of Microtus. Variability in litter size and male body weights was not significantly different when compared between cyclic and non-cyclic populations. Coefficients of variation were also calculated for the three species with the most data (C. gapperi, C. glareolus and M. pennsylvanicus ). These values showed that the amount of intraspecific variation differed among traits as well as among species. Gestation length was the most invariable of all traits and variation in adult weights, neonate weight, gestation length, and litter size had similar values to those reported for Peromyscus maniculatus. Five and eight traits differed among Clethrionomys and Microtus species, respectively. Differences in litter size, adult weights and length of the breeding season were common to both genera. Male weight, gestation length and neonate weight as well as length of the breeding season were different between genera. Very few traits covaried within C. gapperi, C. glareolus or M. pennsylvanicus. Similarly, few traits covaried among all Clethrionomys populations. However, among all Microtus populations and Microtus species, 11 and 12 correlations were significant. Many of the patterns found in Microtus involved positive relationships between female weight and some other trait. These patterns have also been found by broader surveys at higher taxonomic levels. Large species of Microtus had larger offspring, a greater litter size and occurred in short-season environments relative to small species of this genus.     

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.     

Dietary protein constraint on age at maturity: an experimental test with wild deer mice

1. Five populations of Peromyscus maniculatus borealis (Mearns) were live-trapped in the Kananaskis Valley, Alberta, throughout the breeding seasons of 1996 and 1997, to test the hypothesis that growth and maturation of young-of-the-year (YY) females are limited by dietary protein.
2. In 1996, two populations received a high protein (30%) food supplement, one received a high energy, but low protein (14%) food supplement and two others were unsupplemented. In 1997, three populations received the protein (30%) supplement and two served as controls.
3. In 1996, control populations had high nestling growth rates and many (43%) YY females bred. In 1997, nestling growth rates were lower in control populations and no YY females bred.
4. Supplementation of high protein food resulted in consistently high nestling growth rates in both 1996 and 1997, and a significantly higher proportion of YY females bred than controls in both years.
5. Supplementation of high energy, but low protein food had no effect on YY growth or maturation.
6. Neither protein nor energy supplementation had any effects on the number of litters conceived per season, litter success, litter size or sex ratio of litters born to over-wintered females.
7. We conclude that growth and maturation of YY female Peromyscus maniculatus in this area are limited by the availability of dietary protein.     

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.     

Comparative morphology of dark-eyed juncos Junco hyemalis breeding at two elevations: a common aviary experiment

Morphologies of bird species often vary along elevation gradients, yet causes of the variation have not been examined experimentally. We investigated variation in morphological traits of the dark‐eyed junco Junco hyemalis, breeding at 1,000 m a.s.l. (low‐elevation; i.e. low) and 2,000 m asl (high‐elevation; i.e. high) in the Rocky Mountains, Canada. Eight morphological traits were measured in free‐living birds. We found two consistent differences in populations between elevations: at high‐elevation sites, females had longer wings and males had longer tails than birds from low‐ elevation sites. Other age‐ and gender‐ specific results were observed in free‐living birds between elevations: tarsi were shorter in high‐elevation second year (SY) females and after second year (ASY) males, beak lengths were slightly longer in low‐elevation SY females, and high‐elevation ASY females tended to have lower fat than low‐elevation ASY females. Morphological differences may result from genetic differences between elevations, or phenotypic flexibility resulting from exposure to the different environmental conditions. To identify which mechanism caused the difference in morphometrics, hand‐reared birds from low‐ and high‐elevation habitats were raised in identical conditions with unlimited access to high quality food until they had replaced all feathers. The traits measured in the lab (wing and rectrix length, weight and fat score) tended to increase in magnitude compared to field values. Juncos from high‐ and low‐elevations had similar responses to the aviary environment, with one exception: males from high‐elevation sites had greater weight gain relative to free‐living juncos than males from low‐elevation sites. Thus, morphological traits in dark‐eyed juncos were phenotypically flexible, capable of growing larger in the laboratory environment. However, there were also persistent genetic or perinatal/maternal differences underlying population responses that prevented traits from converging under aviary conditions. As a result, trait size differences between high‐ and low‐elevation populations were maintained or exacerbated in the common aviary environment.     

Offspring growth, survival and reproductive success in the bank vole: a litter size manipulation experiment

To estimate the optimality of brood size, it is essential to study the effects of brood size manipulation on offspring survival and reproductive success. Moreover, testing the generality of the hypothesis of reproductive costs requires experimental data from a diversity of organisms. Here I present data on the growth, survival and reproductive success of bank vole Clethrionomys glareolus individuals from manipulated litters. Furthermore, the survival of mothers whose litter size was manipulated was studied. At weaning, the mean weight of pups from enlarged litters was lower and from reduced litters higher compared to control litters. After winter, at the start of the breeding season, individuals from enlarged litters, especially males, were still lighter than individuals from the other two treatments. Litter enlargements did not increase the number of reproducing female offspring per mother, nor did the litter sizes of female offspring differ between treatments. There were no differences between treatments in winter survival of offspring after weaning, but among female offspring, weaning weight explained the survival probabilities over winter. A higher weight of females at winter determined the probability of starting to reproduce in spring. The survival of mothers did not seem to be influenced by litter manipulation performed the previous year. According to the results, mothers nursing enlarged or reduced litters do not gain any fitness benefits in terms of number of offspring surviving to breeding. The results are consistent with the majority of experiments conducted in birds, which have found costs of enlarged brood appearing as offspring trade-offs rather than parent trade-offs. Received: 14 December 1997 / Accepted: 1 March 1998     

Two viruses isolated from rodents (Clethrionomys gapperi and Microtus pennsvlvanicus) trapped in St. Lawrence County, New York

Four strains of C. gapperi virus were isolated from 3 Clethrionomys gapperi and 47 strains of Microtus virus from 15 Microtus pennsylvanicus and 1 Mus musculus. One of the Microtus strains was isolated from a pool of 20 mites while the others were from rodent tissues. These agehts were insensitive to ether and sodium desoxycholate, withstood freezing at -70 C for 3 years and lyophilization without loss of titer, and were not killed when heated at 60 C for 1 hour. Their size as determined by filtration was less than 50 mg and greater than 20-35 mmicro. The strains within each group appear to be similar. The illness induced in suckling mice by the C. gapperi agents had a 5-day incubation period followed by prostration and death with a histologic picture of extensive encephalomalacia. The incubation period in mice for the Microtus agents was 9 to 11 days followed by convulsions and death. Histopathology showed meningeal infiltration and necrosis of the molecular layer. No antigenic similarity was detected between the C. gapperi and Microtus viruses by cross complement-fixation test.     

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.     

Evidence for plasticity in the frequency of skipped breeding opportunities in common toads

Breeding is limited by energetic or environmental constraints and long-lived species sometimes skip breeding opportunities. Environmental conditions may vary considerably across the geographic and elevational range of a species and species that can respond through variation in life history strategies are likely to maintain populations at the extremes of their ranges. The decision to skip breeding enables animals to adjust life history to circumstances, and plasticity in behavior allows implementation of adjustments. Elevational patterns suggest that breeding may be limited physiologically at high elevations (e.g., greater probability of skipped breeding; resources and environmental conditions more variable) in contrast to low elevations (probability of skipping breeding lower; resources and environmental conditions more predictable). We estimated the probabilities of survival and skipped breeding in a high-elevation population of common toads and compared estimates to existing data for common toads at low elevations, and to another toad species inhabiting a similar high elevation environment. Female common toads at high elevations tend to have high probabilities of skipping breeding and survival relative to data for common toads at low elevations, and appear to use a similar strategy of skipping breeding in response to similar environmental constraints as other toads at high elevations. We provide evidence of variability in this aspect of life history for common toads. Understanding variation in life history within widely distributed species is critical. Knowing that certain life history strategies are employed on a continuum informs conservation efforts, especially as impacts of climate change are likely to be different depending on elevation.     

Reproduction in the European hare in southern Sweden

Reproduction in the European hare Lepus europaeus Pallas populations was studied in three study areas in South Sweden during 1974–1976; two on the mainland and the third on the Island of Ven in Öresund. The monthly birth rates was calculated on the basis of the age of juvenile hares shot in autumn. The first births always occurred in February, indicating that the onset of the breeding season was governed by day length rather than by climate. Later, the reproduction rate was determined by temperature, snow cover, and precipitation. Low temperatures in spring delayed the reproduction; high temperatures in autumn favoured a prolongation of the breeding season. Low precipitation in May-July caused a decline in reproduction in July in a sandy pasture area, mainly because water amount was low in the green forage. Annual fertility, estimated on the basis of placental scars, varied between 6.8 and 8.9 young per female. There was a significant negative correlation between fertility and population density. The largest variations were recorded in one-year-old females, and on average the values were significantly higher in three-year-old females. 68% of all females examined had three litters annually; four litters were found in only 13% of all females. The mean litter size for the whole period was about 2.9 in all areas. The size of the first litter was significantly higher in females producing one or two litters than in females with three or four litters. The postnatal losses were on average 30–35% lower in the island population than in the mainland populations (73–84%), probably due to lack of predators, sparse road traffic, and inhibited juvenile dispersal.     

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.     

Optimal litter size for individual growth of European rabbit pups depends on their thermal environment

In altricial mammals and birds, the presence of a large number of litter or brood mates often affects the development of individual offspring by reducing the share of resources provided by the parents. However, sibling presence can also be favourable, conferring thermoregulatory benefits when ambient temperatures are low. Consequently, shifts in the relation between costs and benefits of sibling presence can be expected as a function of the thermal environment. In a study of a European rabbit population (Oryctolagus cuniculus) living in a field enclosure, we investigated the effects of litter size and soil temperature on pup growth over 7 years. Temperatures inside the subterranean nests were positively correlated with soil temperature and with litter size. Soil temperature varied strongly across the breeding season, ranging from 3 to 21°C. Under warmer soil temperature conditions (10–15°C and >15°C), pup growth decreased with increasing litter size, where litters of two pups (smallest litter size considered) showed the highest growth rates. In contrast, under colder soil temperature conditions (<10°C), the highest growth rates were found in litters of three pups. We also asked if such temperature-dependent differences in the optimal pup growth rates might be explained by differences in maternal characteristics, which might affect lactational performance. We assessed maternal performance using females’ postpartum body mass and social rank. However, we did not find consistent differences in maternal characteristics between females giving birth to different-sized litters during different soil temperature conditions, which would have provided an alternative explanation for the observed differences in litter size-dependent pup growth. We conclude that under colder soil temperature conditions, the thermal benefits of a greater number of littermates outweigh the negative consequences of competition for milk, leading to an environment-dependent shift in the optimal litter size for individual growth in this species.     

Growth and morphology of nestling Hooded crows Corvus corone cornix, a sexually dimorphic bird species

The growth and morphology of nestling Hooded crows, Corvus corone cornix , were studied at Trondheim, Central Norway, during 1980 and 1981. The sexual size dimorphism of the nestlings developed while they were in the nest, but was less pronounced than that of the juvenile birds. Female young grew relatively faster than the males. The size variation of the morphological characters studied was greater in the males than in the females, during the entire nestling period. The order of development of the different morphological characters seemed to be adapted to the needs of the nestlings during the nestling period. Growth of the bone and bill characters was the highest priority, body weight and feather characters the lowest. In general, the sizes of the different morphological characters were highly intercorrelated, particularly those that developed at the same time during the growth period. In the favourable breeding season of 1981, the nestlings were larger than in the unfavourable season of 1980. The high priority characters of the males increased more in size than did those of the females, and the low priority characters of the females increased more in size than did those of the males. About 75% of the nestlings could be correctly sexed from their external morphological characteristics. The findings support the view that the parental cost of rearing a male offspring is greater than that of a female.     

Elevational gradient in clutch size of Red‐faced Warblers

Our understanding of life history evolution has benefited from debates regarding the underlying causes, and geographic ubiquity, of spatial patterns in avian clutch sizes. Past studies have revealed that birds lay smaller clutch sizes at higher elevation. However, in most previous studies, investigators have failed to adequately control for elevational differences in breeding phenology. To better understand the elevational gradient in avian clutch size, we need to know how clutch size changes across the entire elevational breeding range of a species (i.e., the shape of the relationship between elevation and clutch size), and whether the elevational gradient in clutch size is merely an artifact of elevational gradients in breeding phenology or breeding season length. We examined the relationship between breeding elevation and clutch size of Red‐faced Warblers (Cardellina rubrifrons) along a 1000‐m elevational gradient in Arizona. Our objectives were to determine how clutch size changed with elevation, and if the relationship between clutch size and elevation merely reflected elevational changes in breeding season length or phenology. The proportion of 5‐egg clutches decreased and the proportion of 3‐ and 4‐egg clutches increased non‐linearly with increasing elevation, even after controlling for the elevational gradient in nest initiation date. Thus, average clutch size declined across the elevational breeding range of Red‐faced Warblers, but this decline was not due to elevational variation in breeding phenology. Timing of breeding changed, but the duration of the breeding season did not change appreciably across the elevational gradient. Hence, elevational differences in breeding season length or breeding phenology cannot explain why Red‐faced Warblers (and perhaps other birds) breeding at higher elevations have smaller clutches.     

Diet quality differentially affects breeding effort of Mastomys coucha and M. natalensis: Implications for rodent pests

In this study we compare the reproductive ability of Mastomys natalensis, an agricultural rodent pest and that of the closely related species M. coucha, which has not been ascribed as a pest. We suggest that the ability of M. natalensis to plague, as well as dominate, resource-limited habitats, may in part be related to its differential breeding response to variation in diet quality. Thus, we examined the reproductive response of M. coucha and M. natalensis mothers to variation in dietary protein intake. Our results demonstrated that typical levels of reproduction extended over a narrower range of dietary protein content for M. coucha than M. natalensis females. Only M. natalensis females bred on 6% protein diets, while on 20% protein diets the reproductive output of M. coucha was lower than on diets containing 10- and 15% protein. M. natalensis responded to low protein diets by reducing litter size and litter mass but not individual pup mass. Thus, providing diet quality improves before parturition, conception under sub-optimal conditions may not compromise individual pup growth. Furthermore, the ability of M. natalensis to breed under conditions of low diet quality may advance their breeding season compared to species that cannot breed until diet quality improves, allowing M. natalensis to produce additional litters through the season, while more daughters could reproduce during the season of their birth, both factors that are known to contribute to the plaguing tendency of M. natalensis.     

Carbon and nutrient mineralization and fungal spore composition of fecal pellets from voles in Minnesota

Numerous investigators have suggested that small mammals may regulate ecosystem processes because they couple producers and decomposers through deposition of fecal matenals and by dispersing spores from mycorrhizal symbiotic fungi We investigated carbon nitrogen, and phosphorus mineralization rates from feces of two arvicoline rodents, Clethrionomn gapperi and Microtus pennsylianicus common in the forests of the Lake Superior region We also examined fungal spore composition from these feces Mineralizable pools of C and N and their decay rates were higher in feces from C gapperi than from M pennsvhamcus , but there were no differences in sizes of mineralizable pools of phosphorus Feces of C gapperi contained four times more fungal spores than those of M pennsylvanicus Twenty-three fungal genera were represented in these feces, and all but one genus have predominantly ectomycorrhizal forms Fungal species composition also differed significantly between the two arvicoline species Although the amount of N and P mineralized annually from populations of these two species is small compared with nutrient budgets for the forests as a whole, these species may be important in dispersing spores and labile nutrient pools to microsites of seedling establishment     

A comparison of life‐history and parental care in temperate and tropical wrens

Comparing closely related species that live in different environments is a powerful way to understand selective pressures that influence life‐history evolution. We examined a suite of life‐history traits and parental care in neotropical buff‐breasted wrens Cantorchilus leucotis and north‐temperate Carolina wrens Thryothorus ludovicianus (Family Troglodytidae), to test hypotheses about life‐history evolution. As expected, buff‐breasted wrens exhibited smaller clutch sizes and higher annual adult survival than Carolina wrens. We found minimal support for the nest predation hypothesis, as nest survival and age‐corrected provisioning rates to whole broods were similar between species, and number of breeding attempts and breeding season length were greater in temperate wrens. Critical predictions of the food limitation hypothesis were not supported; in particular age‐corrected provisioning rates per nestling were higher in the tropical than temperate species. The adult survival and offspring quality hypothesis garnered the most support, as buff‐breasted wrens exhibited greater age‐corrected provisioning rates per nestling, a longer nestling period, longer re‐nesting intervals following nest success, and lower annual fecundity than Carolina wrens. Despite similarly prolonged breeding seasons, reproductive strategies differ between species with buff‐breasted wrens investing considerably in single broods to optimize first‐year survival and Carolina wrens investing in multiple small broods to optimize annual fecundity.     

Flowering phenology and reproduction of the Solidago virgaurea L. complex along an elevational gradient on Mt Norikura,central Japan

This study examined the flowering phenology and reproductive traits of the Solidago virgaurea complex at four elevations in the subalpine zone in Japan using a bagging experiment. Flowering started earlier at higher elevations. Syrphid flies mainly visited flowers of the S. virgaurea complex, and the number of visits was considerably lower at the lowest elevation than at the three other elevations. Although the number of seeds per individual did not differ among the four elevations, total achene weight per individual was lower at the lowest elevation than at the three other elevations. The weight of an achene and seed germination rate of the control were much greater at higher elevations than at the lowest elevation. The weight of an achene and germination rate for the control were as low as the bagging treatment at the lowest elevation with infrequent flower visitors, which indicates that the S. virgaurea complex is a facultative outcrosser. The S. virgaurea complex is thought to produce seeds during a short growing season at high elevations by starting to flower earlier, and the large seed size is advantageous for seedling establishment at high elevations. Therefore, elevational changes in flowering phenology and reproductive traits are thought to be an adaptation to the short growing season at high elevations.     

High and low, fast or slow: the complementary contributions of altitude and latitude to understand life-history variation

Dark-eyed juncos ( Junco hyemalis ) from two elevations in the Rocky Mountains of Canada display different life histories. Birds breeding at high elevation face a later and compressed reproductive season during which they raise on average half the number of broods (young) than their low-elevation conspecifics. Fledglings at high elevation are heavier, fatter, and have an increased chance of surviving to 25 days of age. Likewise, survival of adults (males) increases at higher elevation. Put into a broader perspective, high-elevation juncos show the life-history strategy characteristic for low-latitude birds. This raises questions about the mechanisms influencing life-history evolution.     

Selection for synchronous breeding in the European starling

Colonial birds often demonstrate considerable breeding synchrony. In southern Sweden the semi-colonial European starling initiated the vast majority of clutches within one week. Laying dates were positively skewed so that many birds initiated clutches at similar dates early in the season. Breeding was further synchronised by a particularly strong clutch-size reduction equivalent to one third of an egg per day during the first part of the breeding season. The decline in clutch size with season also held true for separate age-classes of females, for individual females laying at different times at different years and for individual females laying at different times the same year. Trends in breeding success during nestling rearing were unlikely to explain the high degree of breeding synchrony or the seasonal decline in clutch size; nestling survival and growth were weakly related or unrelated to reproductive timing. In contrast recruitment success of fledged offspring declined sharply with season. Even within the synchronous laying period, defined as clutches initiated during the first week each year, local recruitment success declined. It is suggested that the early seasonal decline is caused by selection for synchronous fledging permitting the immediate formation of flocks after fledging, whereas the late seasonal trends may be caused by either population differences in female quality or deteriorating conditions for raising young.