Density‐dependent demography and movements in a cyclic brown lemming population

Theoretical modeling predicts that both direct and delayed density‐dependence are key factors to generate population cycles. Deciphering density‐dependent processes that lead to variable population growth characterizing different phases of the cycles remains challenging. This is particularly the case for the period of prolonged low densities, which is inherently data deficient. However, demographic analyses based on long‐term capture–mark–recapture datasets can help resolve this question. We relied on a 16‐year (2004–2019) live‐trapping program to analyze the summer demography and movements of a cyclic brown lemming population in the Canadian Arctic. More specifically, we examined if inversely density‐dependent processes could explain why population growth can remain low during the prolonged low phase. We found that the proportion of females in the population was inversely density‐dependent with a strong male‐biased sex ratio at low densities but not at high densities. However, survival of adult females was higher than adult males, but both had lower survival at low densities than at high ones. Distances moved by both adult males and females were density‐dependent, and proportion of females in reproductive condition was weakly density‐dependent as it tended to increase at low density. Individual body condition, measured as monthly change in body mass, was not density‐dependent. Overall, the strong male‐biased sex ratio at very low densities suggests a loss of reproductive potential due to the rarity of females and appears to be the most susceptible demographic factor that could contribute to the prolonged low phase in cyclic brown lemmings. What leads to this sex‐bias in the first place is still unclear, potentially owing to our trapping period limited to the summer, but we suggest that it could be due to high predation rate on breeding females in winter.     

Is there direct and delayed density dependent variation in population structure in a temperate European cyclic vole population?

Population structure, in terms of the body mass, condition, sex and reproductive status of individuals, has been found to vary in phase with population density in cyclic populations of microtine rodents. Because sustained population cycles involve delayed density dependent changes in the population growth rate, we would expect at least some life history traits also to depend on past densities. Detailed, long-term studies of changes in vole life history traits are however few, and are largely restricted to northern Europe. In view of the uncertainty as to whether the cyclic microtine populations of western Europe represent the same phenomenon as those of northern Europe, we studied temporal variation in the structure of a clearly cyclic population of the common vole Microtus arvalis Pallas, in the cereal plains of mid-western France. Our data set contains seasonal, individual-level data from long-term, large-scale trapping covering four entire population cycles. We found considerable cyclic variation in population structure in spring (April), but less so in summer (June). In spring of post-peak years, animals were of low body weight and body condition (particularly females), litter sizes were smaller and there was a reduction in the proportion of breeders. All of these could be proximal drivers of increased mortality rates, or decreased birth rates, contributing to the population declines. Few life history traits, however, showed direct density dependent variation, and none of the traits studied here showed delayed density dependence. We have shown declines in the fecundity and body condition of voles from a western European population that coincides with, and may be a proximal cause of, cyclic declines in population density. Closer attention to proximal causes, by which ecological processes drive cycles, could clarify the extent to which microtine cycles across Europe represent a single phenomenon.     

Are silica defences in grasses driving vole population cycles?

Understanding the factors that drive species population dynamics is fundamental to biology. Cyclic populations of microtine rodents have been the most intensively studied to date, yet there remains great uncertainty over the mechanisms determining the dynamics of most of these populations. For one such population, we present preliminary evidence for a novel mechanism by which herbivore-induced reductions in plant quality alter herbivore life-history parameters and subsequent population growth. We tested the effect of high silica levels on the population growth and individual performance of voles (Microtus agrestis) reared on their winter food plant (Deschampsia caespitosa). In sites where the vole population density was high, silica levels in D. caespitosa leaves collected several months later were also high and vole populations subsequently declined; in sites where the vole densities were low, levels of silica were low and population density increased. High silica levels in their food reduced vole body mass by 0.5% a day. We argue that silica-based defences in grasses may play a key role in driving vole population cycles.     

The difference between generalist and specialist: the effects of wide fluctuations in main food abundance on numbers and reproduction of two co-existing predators

Specialist individuals within animal populations have shown to be more efficient foragers and/or to have higher reproductive success than generalist individuals, but interspecific reproductive consequences of the degree of diet specialisation in vertebrate predators have remained unstudied. Eurasian pygmy owls (hereafter POs) have less vole-specialised diets than Tengmalm's owls (TOs), both of which mainly subsist on temporally fluctuating food resources (voles). To test whether the specialist TO is more limited by the main prey abundance than the generalist PO, we studied breeding densities and reproductive traits of co-existing POs and TOs in central-western Finland during 2002–2019. Breeding densities of POs increased with augmenting densities of voles in the previous autumn, whereas breeding densities of TOs increased with higher vole densities in both the previous autumn and the current spring. In years of vole scarcity, PO females started egg-laying earlier than TOs, whereas in years of vole abundance TO females laid eggs substantially earlier than PO females. The yearly mean clutch size and number of fledglings produced of both POs and TOs increased with abundance of voles in the current spring. POs laid large clutches and produced large broods in years of both high and low vole abundance, whereas TOs were able to do so only in years of high vole abundance. POs were able to raise on average 73% of the eggs to fledglings whereas TOs only 44%. The generalist foraging strategy of POs including flexible switching from main prey to alternative prey (small birds) appeared to be more productive than the strictly vole-specialized foraging strategy of TOs. In contrast to earlier studies at the individual-level, specialist predators at the species level (in this case TOs) appear to be less effective than generalists (POs), but diet specialisation was particularly costly under conditions when scarcity of main foods limited offspring production.     

Dampening prey cycle overrides the impact of climate change on predator population dynamics: a long‐term demographic study on tawny owls

Predicting the dynamics of animal populations with different life histories requires careful understanding of demographic responses to multifaceted aspects of global changes, such as climate and trophic interactions. Continent‐scale dampening of vole population cycles, keystone herbivores in many ecosystems, has been recently documented across Europe. However, its impact on guilds of vole‐eating predators remains unknown. To quantify this impact, we used a 27‐year study of an avian predator (tawny owl) and its main prey (field vole) collected in Kielder Forest (UK) where vole dynamics shifted from a high‐ to a low‐amplitude fluctuation regime in the mid‐1990s. We measured the functional responses of four demographic rates to changes in prey dynamics and winter climate, characterized by wintertime North Atlantic Oscillation (wNAO). First‐year and adult survival were positively affected by vole density in autumn but relatively insensitive to wNAO. The probability of breeding and number of fledglings were higher in years with high spring vole densities and negative wNAO (i.e. colder and drier winters). These functional responses were incorporated into a stochastic population model. The size of the predator population was projected under scenarios combining prey dynamics and winter climate to test whether climate buffers or alternatively magnifies the impact of changes in prey dynamics. We found the observed dampening vole cycles, characterized by low spring densities, drastically reduced the breeding probability of predators. Our results illustrate that (i) change in trophic interactions can override direct climate change effect; and (ii) the demographic resilience entailed by longevity and the occurrence of a floater stage may be insufficient to buffer hypothesized environmental changes. Ultimately, dampened prey cycles would drive our owl local population towards extinction, with winter climate regimes only altering persistence time. These results suggest that other vole‐eating predators are likely to be threatened by dampening vole cycles throughout Europe.     

Reproductive investment under fluctuating predation risk: Microtine rodents and small mustelids

Summary We studied the reproductive investment of microtine rodents (bank vole (Clethrionomys glareolus),Microtus epiroticus andMicrotus agrestis) in western Finland under predation risk from small mustelids. During 1984–1992, the yearly mean litter size of overwintered bank voles was smaller at high least weasel and stoat densities than at low densities (close to 3 versus 4–5). In addition, the annual mean litter size of young bank voles was negatively correlated to the least weasel density. In youngM. agrestis voles, the yearly late summer litter size was negatively associated with the autumn density of small mustelids. In the crash phase of the vole cycle (1989 and 1992), we removed small mustelids (mainly least weasels) from four unfenced areas in late April to late May and studied the reproduction of voles in four removal and comparable control areas (each 2–4 km²). Reduction of small mustelids significantly increased the proportion of pregnant bank vole females, but not that of pregnantMicrotus vole females. We conclude that predation risk apparently reduced reproductive investment of free-living bank vole females; these voles appear to trade their current parental investment against future survival and reproductive prospects. Accordingly, the presence of small mustelids (or their scent) may slow down the reproductive rate of voles. As antipredatory behaviours occurred on a large scale, our results add evidence to the hypothesis that crashes in multiannual vole cycles are driven by small mustelid predators.     

Fading out of vole and predator cycles?

Northern voles and lemmings are famous for their spectacular multiannual population cycles with high amplitudes. Such cyclic vole populations in Scandinavia have shown an unexpected and marked long-term decline in density since the early 1970s, particularly with a marked shift to lower spring densities in the early 1980s. The vole decline, mainly characterized by a strongly decreased rate of change in numbers over winter, is associated with an increased occurrence of mild and wet winters brought about by a recent change in the North Atlantic Oscillation. This has led to a decrease in winter stability and has shortened the period with protective snow cover, the latter considered as an important prerequisite for the occurrence of multiannual, high-amplitude cycles in vole populations. Although the vole decline is predicted to be negative for predators' reproduction and abundance, empirical data showing this are rare. Here we show that the dynamics of a predator-prey system (Tengmalm's owl, Aegolius funereus, and voles), have in recent years gradually changed from 3-4 yr, high-amplitude cycles towards more or less annual fluctuations only.     

Factors affecting fecal glucocorticoid levels in semi-free-ranging female mandrills (Mandrillus sphinx)

Subordinate female cercopithecine primates often experience decreased reproductive success in comparison with high-ranking females, with a later age at sexual maturity and first reproduction and/or longer interbirth intervals. One explanation that has traditionally been advanced to explain this is high levels of chronic social stress in subordinates, resulting from agonistic and aggressive interactions and leading to higher basal levels of glucocorticoids. We assessed the relationships among fecal cortisol levels and reproductive condition, dominance rank, degree of social support, and fertility in female mandrills (Mandrillus sphinx) living in a semi-free-ranging colony in Franceville, Gabon. Lower-ranking females in this colony have a reproductive disadvantage relative to higher-ranking females, and we were interested in determining whether this relationship between dominance rank and reproductive success is mediated through stress hormones. We analyzed 340 fecal samples from 19 females, collected over a 14-month period. We found that pregnant females experienced higher fecal cortisol levels than cycling or lactating females. This is similar to results for other primate species and is likely owing to increased metabolic demands and interactions between the hypothalamus-pituitary-adrenal axis, estrogen, and placental production of corticotrophin releasing hormones during pregnancy. There was no influence of dominance rank on fecal cortisol levels, suggesting that subordinate females do not suffer chronic stress. This may be because female mandrills have a stable social hierarchy, with low levels of aggression and high social support. However, we found no relationship between matriline size, as a measure of social support, and fecal cortisol levels. Subordinates may be able to avoid aggression from dominants in the large enclosure or may react only transiently to specific aggressive events, rather than continuously expecting them. Finally, we found no relationship between fecal cortisol levels and fertility. There was no difference in fecal cortisol levels between conceptive and nonconceptive cycles, and no significant relationship between fecal cortisol level and either the length of postpartum amenorrhea or the number of cycles before conception. This suggests that the influence of dominance rank on female reproductive success in this population is not mediated through chronic stress in subordinate females, and that alternative explanations of the relationship between social rank and reproduction should be sought.     

Frequency and density-dependent selection on life-history strategies--a field experiment

Negative frequency-dependence, which favors rare genotypes, promotes the maintenance of genetic variability and is of interest as a potential explanation for genetic differentiation. Density-dependent selection may also promote cyclic changes in frequencies of genotypes. Here we show evidence for both density-dependent and negative frequency-dependent selection on opposite life-history tactics (low or high reproductive effort, RE) in the bank vole (Myodes glareolus). Density-dependent selection was evident among the females with low RE, which were especially favored in low densities. Instead, both negative frequency-dependent and density-dependent selection were shown in females with high RE, which were most successful when they were rare in high densities. Furthermore, selection at the individual level affected the frequencies of tactics at the population level, so that the frequency of the rare high RE tactic increased significantly at high densities. We hypothesize that these two selection mechanisms (density- and negative frequency-dependent selection) may promote genetic variability in cyclic mammal populations. Nevertheless, it remains to be determined whether the origin of genetic variance in life-history traits is causally related to density variation (e.g. population cycles).     

Effects of competition and season on survival and maturation of young bank vole females

In territorial microtines intra-specific density dependent processes can limit the maturation of individuals during the summer of their birth. This may have demographic consequences by affecting the number and the age distribution of breeding individuals in the population. Little is known about this process on a community level, though populations of many northern microtine species fluctuate in synchrony and are known to interfere socially with each other. We experimentally studied the influence of the field vole Microtus agrestis on maturation, breeding, space use and survival of weanling bank voles, Clethrionomys glareolus. Two additive competition experiments on bank vole populations were conducted in large outdoor enclosures, half of them additionally housing a field vole population. In a mid-summer experiment low population density and absence of older breeding females minimised intra-specific competition. Survival was not affected by the presence of field voles. Season had a significant effect on both the probability of maturation and breeding of the weanlings. Competition with field voles significantly delayed breeding, and coupled with seasonal effects decreased the probability of breeding. In a late-summer experiment breeding and survival of bank vole weanlings were studied for three weeks as part of a high density breeding bank vole population. Weanlings did not mature at all nor were their space use and survival affected by the presence of field voles. Our results show that competition with other species can also have an impact on breeding of immatures. In an extreme seasonal environment, even a short delay of breeding may decrease survival chances of offspring. Seasonal and competition effects together may thus limit the contribution of year born females to reproductive output of the population. Other studies have shown that adult breeding bank voles suffer lower survival in the presence of field voles, but this study showed no survival effects on the weanlings. Thus it might be beneficial for weanlings to stay immature especially in the end of the breeding season and postpone reproduction to the next breeding season if densities of competing species are high.     

Dynamic effects of predators on cyclic voles: field experimentation and model extrapolation

Mechanisms generating the well-known 3-5 year cyclic fluctuations in densities of northern small rodents (voles and lemmings) have remained an ecological puzzle for decades. The hypothesis that these fluctuations are caused by delayed density-dependent impacts of predators was tested by replicated field experimentation in western Finland. We reduced densities of all main mammalian and avian predators through a 3 year vole cycle and compared vole abundances between four reduction and four control areas (each 2.5-3 km(2)). The reduction of predator densities increased the autumn density of voles fourfold in the low phase, accelerated the increase twofold, increased the autumn density of voles twofold in the peak phase, and retarded the initiation of decline of the vole cycle. Extrapolating these experimental results to their expected long-term dynamic effects through a demographic model produces changes from regular multiannual cycles to annual fluctuations with declining densities of specialist predators. This supports the findings of the field experiment and is in agreement with the predation hypothesis. We conclude that predators may indeed generate the cyclic population fluctuations of voles observed in northern Europe.     

Effects of ectoparasite infestation during pregnancy on physiological stress and reproductive output in a rodent-flea system

Biotic and abiotic stressors impose various fitness costs on individuals across a variety of taxa. In vertebrates, these stressors typically trigger complex neuroendocrine responses that stimulate glucocorticoid (GC) secretion from the adrenal cortex. Short-term elevation of GCs can be adaptive as it shifts energy toward physiological processes that cope with acute stressors; however, chronic increases in GC levels could have detrimental effects on fitness. Parasitism can be considered an important biotic stressor in nature and a possible cause of reproductive failure that could substantially affect an individual’s fitness. Thus, we aimed to test the effects of parasitism and maternal stress, as measured by GCs, during pregnancy and the relationship between these variables and measures of reproductive output using a rodent-flea system. Female Egyptian spiny mice (Acomys cahirinus) were randomly assigned to flea (Parapulex chephrenis) infested or uninfested treatments before and during pregnancy. The offspring of these females were flea-free. Feces were collected at five time points during the experiment to determine maternal fecal glucocorticoid metabolite (FGCM) concentrations. Overall, infested females had lower FGCM levels during gestation but higher FGCM levels post-parturition and larger mass changes than uninfested females. Additionally, models related to pup quality and quantity often included some measure of maternal investment or body condition moderating relationships between infestation and stress. This suggests that flea parasitism or high GC levels alone might not significantly impact host reproduction but rather females can experience different effects depending on their level of investment, which could be limited by body condition and/or the number of pups present in a litter.     

Resource levels,reproduction and resistance to haematozoan infections

Compromise of immune function during reproduction may form a link between parental effort and the cost of reproduction, but the role of environmental variation in structuring intra-individual life-history trade-offs has been poorly investigated. We manipulated the need for parental effort in Eurasian kestrels, Falco tinnunculus, by food-supplementing broods for three nestling periods, during which the natural main food supply (voles) varied, and found that parental parasitaemia was inversely related to yearly vole densities. The level of parasitaemia in females was, however, reduced by food supplements. No effect on males was expected, as earlier work has shown that only females responded to the supplements by changing their behaviour. We show directly that the likelihood of female parasitaemia was diminished by spending less time in flight-hunting, which was related to reproduction during a good vole year, to our supplementary feeding, and to being mated to a male with high parental effort. Our results represent a novel direct benefit for females in resource - providing species, linked to female, as well as offspring, well-being, and they provide insight into why the appearance of reproductive costs may be linked to gender or environmental conditions.     

Competition, predation and interspecific synchrony in cyclic small mammal communities

Although competition and predation are considered to be among the most important biotic processes influencing the distribution and abundance of species in space and time, the relative and interactive roles of these processes in communities comprised of cyclically fluctuating populations of small mammals are not well known. We examined these processes in and among populations of field voles, sibling voles, bank voles and common shrews in western Finland, using spatially replicated trapping data collected four times a year during two vole cycles (1987–1990 and 1997–1999). Populations of the four species exhibited relatively strong interspecific temporal synchrony in their multiannual fluctuations. During peak phases, we observed slight deviations from close temporal synchrony: field vole densities peaked at least two months earlier than those of either sibling voles or bank voles, while densities of common shrews peaked even earlier. The growth rates of all four coexisting small mammal species were best explained by their own current densities. The growth rate of bank vole populations was negatively related to increasing densities of field voles in the increase phase of the vole cycle. Apart from this, no negative effects of interspecific density, direct or delayed, were observed among the vole species. The growth rates of common shrew populations were negatively related to increasing total rodent (including water voles and harvest mice) densities in the peak phase of the vole cycle. Sibling voles appeared not to be competitively superior to field voles on a population level, as neither of these Microtus voles increased disproportionately in abundance as total rodent density increased. We suggest that interspecific competition among the vole species may occur, but only briefly, during the autumn of peak years, when the total available amount of rodent habitat becomes markedly reduced following agricultural practices. Our results nonetheless indicate that interspecific competition is not a strong determinant of the structure of communities comprised of species exhibiting cyclic dynamics. We suggest that external factors, namely predation and shortage of food, limit densities of vole populations below levels where interspecific competition occurs. Common shrews, however, appear to suffer from asymmetric space competition with rodents at peak densities of voles; this may be viewed as a synchronizing effect.     

Chronic stress in pregnant guinea pigs (<Emphasis Type="Italic">Cavia aperea f. porcellus</Emphasis>) attenuates long-term stress hormone levels and body weight gain,but not reproductive output

Stress, when extreme or chronic, can have a negative impact on health and survival of mammals. This is especially true for females during reproduction when self-maintenance and investment in offspring simultaneously challenge energy turnover. Therefore, we investigated the effects of repeated stress during early- and mid-gestation on the maternal stress axis, body weight gain and reproductive output. Female guinea pigs (Cavia aperea f. porcellus, n = 14) were either stressed (treatment: exposure to strobe light in an unfamiliar environment on gestational day -7, 0, 7, 14, 21, 28, 35, 42) or left completely undisturbed (control) throughout pregnancy. Females of both groups received the same respective diets, and reproductive parameters were evaluated upon parturition. Additionally, hormonal data were obtained from blood and feces. The stress exposure induced a significant increase in plasma cortisol concentrations during the afternoon. In contrast to this short-term response in plasma cortisol concentrations, we found no significant differences in the levels of cortisol metabolites in feces collected after stress exposure between groups and even significantly decreased levels of fecal cortisol metabolites on non-stress days over time in treatment females. Among treatment females, gain in body weight was attenuated over gestation and body weight was lower compared to control females during lactation, especially in cases of large litter sizes. No differences could be seen in the reproductive parameters. We conclude that repeated stress exposure with strobe light during early- and mid-gestation results in a down-regulation of the hypothalamic–pituitary–adrenal axis and lower weight gain in treatment females, but has no effect on reproductive output.     

Is negative density‐dependent reproduction regulated by density‐induced stress in root voles? Two field experiments

Density dependence in reproduction plays an important role in stabilizing population dynamics via immediate negative feedback from population density to reproductive output. Although previous studies have shown that negative density‐dependent reproduction is associated with strong spacing behavior and social interaction between individuals, the proximal mechanism for generating negative density‐dependent reproduction remains unclear. In this study, we investigated the effects of density‐induced stress on reproduction in root voles. Enclosed founder populations were established by introducing 6 (low density) and 30 (high density) adults per sex into per enclosure (four enclosures per density in total) during the breeding season from April to July 2012 and from May to August 2015. Fecal corticosterone metabolite (FCM) levels, reproductive traits (recruitment rate and the proportion of reproductively active individuals), and founder population numbers were measured following repeated live trapping in both years. The number of founders was negatively associated with recruitment rates and the proportion of reproductively active individuals, displaying a negative density‐dependent reproduction. FCM level was positively associated with the number of founders. The number of founder females directly affected the proportion of reproductive females, and directly and indirectly through their FCM levels affected the recruitment rate; the effect of the number of male founders on the proportion of reproductive males was mediated by their FCM level. Our results showed that density‐induced stress negatively affected reproductive traits and that density‐induced stress is one ecological factor generating negative density‐dependent reproduction.     

Seasonality, body condition, and timing of reproduction in Propithecus verreauxi verreauxi in the Kirindy Forest

Mammals that live in seasonal environments may adjust their reproductive cycles to cope with fluctuations in food availability. Because lemurs in Madagascar experience highly seasonal variation in food availability, we examined the effects of fluctuating food availability on body condition and reproduction in one of the larger living species, Verreaux's sifaka (Propithecus verreauxi verreauxi), in the Kirindy Forest of western Madagascar. Seven years of demographic data were combined with an intensive study of 25 individuals over the course of 18 months. In contrast to other populations of Verreaux's sifaka, females were found to have greater body mass than males. Both male and female sifaka exhibited significant losses of body mass and fat during the dry season. Females were more likely to give birth and successfully wean an infant when they had higher body mass during the mating season. They mated during the periods of high and declining food availability, gave birth during the lean season, and then timed mid/late lactation with the period of increasing food availability. Thus, we conclude that sifaka follow the "classic" reproductive strategy (sensu van Schaik and van Noordwijk [Journal of Zoology (London) 206:533-549, 1985]).     

Long-term decline in numbers of cyclic voles in boreal Sweden: analysis and presentation of hypotheses

Cyclic vole populations, defined as showing fairly regular 3–4 yr density fluctuations but with variable amplitudes, were monitored in boreal Sweden in spring and fall 1971–2002, starting in fall 1971. Voles were snap-trapped on permanent sampling plots at the landscape level within a 100 by 100 km study area north of Umeå. The predominating species trapped were Clethrionomys glareolus, C. rufocanus and Microtus agrestis. In addition to the 3–4 yr cycles, there was a long-term decrease in numbers and amplitude of the fluctuations, which was especially conspicuous in C. rufocanus. In this latter species there was a persistent decline of both spring and fall densities, apparently bringing the population close to extinction in the area. However, the decline of spring densities from the 1970s to the 1980s and onwards was also evident in C. glareolous and M. agrestis. The declines in numbers and amplitude were largely linked to an increased frequency and/or accentuation of winter declines, which more or less neutralized or even overrode the density increase during the reproductive season in the previous summer, especially so in the second year of the cycles. Thereby the gradual two-large-step build-up of high spring densities, very much founding the base for the very large peak densities and amplitudes in the 1970s, was successively replaced by a one-smaller-step build-up of more modest spring densities, leading to lower peak densities and amplitudes in the 1980s, 1990s and early 2000s. Understanding the causes of the increased frequency and/or severeness of winter declines appears critical to understanding the observed long-term changes in numbers. However, the underlying causes of the increase of winter declines and the decrease of densities and amplitudes are unknown, but some hypotheses are presented and discussed here. Also, some implications from the decreased vole abundance for reproduction and densities of predators on the voles, and on predators’ alternative prey species, are briefly discussed.     

Effects of forage availability on growth and maturation rates in water voles

1. In populations of small mammals, food supplementation typically results in higher population densities, body weights, growth rates and reproductive rates. However, few studies have demonstrated a relationship between forage levels and demographic rates in wild populations in the absence of supplementation. 2. We examined the association of levels of available forage with individual growth rates and time to sexual maturity in eight re-introduced and three naturally occurring populations of water voles (Arvicola terrestris). 3. Range sizes were smaller at sites with higher population densities. Mean forage availability and individual growth rates covaried with range size at each site. 4. The weight at which water voles became sexually mature was 112 g for females and 115 g for males and did not vary between study sites. Differences in growth rates therefore translated into differences in the time taken to reach maturity between sites. 5. In the re-introduced populations, mean days to maturity varied inversely with mean range length. Females took 7 days (18%, range 40-47 days) longer and males 5 days (13%, range 40-45 days) longer to reach breeding condition at the sites with the shortest mean range lengths. 6. Evidence from this study suggests a possible mechanism by which increased population densities may reduce maturation rates in water voles through a reduction in mean range size, thereby limiting the availability of forage to each individual.     

Food limitation and social regulation in a red fox population

This study evaluates a conceptual model on functional and numerical response to short-term fluctuating vole populations of a red fox ( Vulpes vulpes L.) population in south-central Sweden. The model assumes that this particular population is located in between socially regulated stable populations to the south and direct food-limited populations to the north. The model predicts: (1) food availability as the primary factor for limiting fox numbers, causing reduced rates of reproduction and survival during years of low vole densities, and (2) density-dependent regulation during years of increasing and high vole densities resulting in increased group sizes within territories of fixed dimensions. During 1973–1980 data were obtained from 1216 fox scats, 874 fox carcasses, 63 tagged foxes, nine radio-collared females and from yearly den counts in an area of 130 km², Eight predictions of the model were tested. These concerned the occurrence of small rodents in fox diet, fluctuations in the density of foxes, variations in the number of fox litters, the effect on reproduction of providing supplemental food during January–May, the proportion of vixens bearing a litter different years, dispersal of young males relative to that of young females throughout the vole cycle, and variations in mortality rates of young males and females. All tests were in favour of the conceptual model, and contradictory to alternative models.