Reproduction strategy in an island population of yellow-necked mice

A population of yellow-necked mice, Apodemus flavicollis, inhabiting Crabapple Island (NE Poland) was sampled from 1994–2002. The catch-mark-release method was applied. Sexually mature males and females (including pregnant ones) as well as immatures of each sex were identified. During the study period 3,036 mice were caught 10,672 times. The number of mice caught varied from 6 individuals (April 1999) to 272 (July 1996). The number of both mature females and mature males increased linearly with population size. Differences in the reproductive strategies of yellow-necked mouse and bank voles, Clethrionomys glareolus, are discussed. The respective strategies led to high variability in the population size of mice and much less variability in the case of the voles during the breeding season.     

Hypothalamo-pituitary neurosecretory system of the Northern redbacked vole Clethrionomys rutilus in the course of population cycle

Hypothalamo-pituitary neurosecretory system (HPNS) of the Northern redbacked vole, Clethrionomys rutilus was studied at different stages of the population cycle using paraldehyde-fuchsin staining and immunohistochemical revealing of vasopressin and oxytocin. We found at the stages of high voles number (peak and recession), an increase of vasopressin synthesis in the neurosecretory cells (NSC) of paraventricular (PVN) and supraoptic (SON) nuclei, as well as its active transport and release to the portal capillaries of the outer zone of median eminence (ME). At the stages of low voles number (depression and growth) was demonstrated that level of oxytocin synthesis in the NSC of SON was high and moderate in the NSC of PVN, which was accompanied by an extensive release of oxytocin to capillaries of the posterior pituitary (PP). Increased supply of vasopressin to portal blood flow of the vole pituitary in conditions of overpopulation is suggested to have highly stimulating influence on adrenocorticotropic function of pituitary, which negatively affects the reproductive function of the voles and leads to a decrease of their number. At the stages of low number of population in conditions favoring the life of the voles, the increased supply of oxytocin to the systemic blood flow stimulates the reproductive behavior of the voles, which results in rise of their population during this period.     

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.     

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.     

HETEROZYGOSITY,AGGRESSION, AND POPULATION FLUCTUATIONS IN MEADOW VOLES (MICROTUS PENNSYLVANICUS)

We tested whether variation in heterozygosity could produce cyclic changes in population size in meadow voles (Microtus pennsylvanicus). For this to occur, three conditions must be met: (1) populations are more outbred (heterozygotic) at high than low population density, (2) heterozygotic voles are more aggressive than relatively inbred individuals, and (3) heterozygotic voles have lower reproductive fitness, though being superior at defending resources. We found no evidence that heterozygosity varied with population size or that reproductive success varied with heterozygosity. However, the former test was indirect and relatively weak. We directly measured aggression and heterozygosity of individual voles. Aggression was significantly related to heterozygosity: higher heterozygosity correlated with more aggression in males and less aggression in females. The proportion of variance in aggression that could be explained by heterozygosity was small. These results suggest that changes in population size of meadow voles could not be driven by systematic changes in heterozygosity with population size.     

Is space management of female meadow voles (Microtus pennsylvanicus) related to nutritive quality of plants?

Summary It is thought by many (see Ims 1987 for review; Desy and Batzli 1989) that high quality food regulate population processes, territoriality and mating systems among small herbivores like meadow voles (Microtus pennsylvanicus). We thought that comparisons of nutritive components from selected plants eaten by sexually active and inactive voles, as well as between territorial and non territorial sexually active females would bring some light into these theoretical considerations. Sexually active females did have a higher diet quality over inactive ones and over active and inactive males. Nutritive components of selected species from territorial reproductive females did not vary significantly from those of the non territorial females the year of higher crowding conditions but they varied significantly the following year when population density of voles was much lower. This decline in food quality coincided with a switch in food selection. Since there were only eight plant species involved in such processes, we think that crowding condition and availability of high quality food are two factors involved concurrently in space management and territoriality among voles.     

Does maternal stress influence winter survival of offspring in root voles Microtus oeconomus? A field experiment

Maternal stress can have long‐term adverse consequences on immunocompetence and disease risk of offspring, and winter survival is a crucial demographic parameter in the life‐history of an individual that can substantially affect northern rodent population dynamics. An understanding of the effects of maternal stress on winter survival of offspring may help identify mechanisms driving population fluctuations of northern small mammals. Thus, we assessed the effects of maternal stress, resulting from high population densities, on winter survival of first generation (F₁) and second generation (F₂) in root voles Microtus oeconomus. Replicate high‐ and low‐density enclosed parental populations were established, from which we obtained F₁ generation that were used to establish new enclosed, equal‐density populations. The adults of the high‐density parental populations had higher corticosterone levels, an indication of physiological stress, than did those of the low‐density parental populations. Over‐winter survival of the F₁ generation voles from the low‐density parental populations was greater than that of those from the high‐density parental populations. Over‐winter survival of F₂ generation voles did not differ between the two treatments. Our results suggest that maternal stress affected over‐winter survival of first generations but not second generations. Reduced immunocompetence, resulting from high population density stresses, transferred to offspring may be a factor in annual (winter) population declines. Because the effect is transitory, i.e. immunocompetence of F₂ voles is not affected, reduced immunocompetence resulting from high density stresses would not contribute to lengthy periods of low population densities that are characteristic of multi‐annual population fluctuations.     

MICROGEOGRAPHIC VARIATION IN MITOCHONDRIAL DNA OF MEADOW VOLES (MICROTUS PENNSYLVANICUS) IN RELATION TO POPULATION DENSITY

We examined mitochondrial-DNA (mtDNA) sequence heterogeneity on four adjacent trapping grids in an island population of meadow voles (Microtus pennsylvanicus) at two different population densities. Four restriction endonucleases revealed 20 different mtDNA composite phenotypes in samples totaling 198 meadow voles. There were significant heterogeneities in the distribution of four common mtDNA composite phenotypes among the four trapping grids, suggesting that there is population subdivision on a fine scale. Genetic distances between grids, mtDNA diversity within grids, and G_ST also varied during the study period. We found a decrease in genetic distance and an increase in diversity when the population density was high and vice versa when the population density was low. When population density was high, the coefficient of gene differentiation was smaller than the same coefficient observed when the population density was low. These changes in population subdivision and diversity are consistent with theoretical expectations of population structure in which effective female population size and dispersal are the critical variables. The data also support the hypothesis of maintenance of mtDNA diversity by population subdivision, rapid population growth rate, and dispersal.     

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.     

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.     

The cost of habitat selection in prairie voles: an empirical assessment using isodar analysis

The ideal free distribution assumes that habitat selection is without cost and predicts that fitness should be equal in different habitats. If habitat selection has a cost, then individuals should only move to another habitat when potential fitness in the new habitat exceeds that in the source habitat by an amount greater than the cost of habitat selection. We used isodar techniques to assess the cost of habitat selection. In an experimental landscape, we monitored density, movement, and reproductive success of adult female prairie voles, Microtus ochrogaster, in adjacent paired habitats with low and high cover. We tested the following hypotheses: (1) adult female prairie voles exhibited density-dependent habitat selection; (2) the cost of habitat selection was density-independent. Habitat quality based on population density and fitness of adult females was higher in high cover habitats. Net movement was from low cover to high cover habitats. The results indicated that adult female prairie voles exhibited density-dependent habitat selection. Furthermore, there was a significant cost of habitat selection, and the cost was density-independent.     

Cycles in voles and small game in relation to variations in plant production indices in Northern Sweden

Population dynamics for voles (Cricetidae), Tengmalm's owl (Aegolius funereus (L.)), red fox (Vulpes vulpes (L.)) willow grouse (Lagopus lagopus (L.)), black grouse (Lyrurus tetrix (L.)), capercaillie (Tetrao urogallus L.), hazel hen (Tetrastes bonasia (L.)), mountain hare (Lepus timidus L.) and tularemia (Francisella tularensis (McCoy & Chapin)) and game bird recruitment were studied by index methods in northern Sweden. In addition contemporary temperature records and spruce (Picea abies (L.) Karst.) and pine (Pinus silvestris L.) cone crops (as indices for plant production) and the occurrence of forest damage, caused by voles eating bark, were studied.During 1970–80 two synchronous 4-year cycles were observed for voles, predators (Tengmalm's owl and red fox) and their alternative prey species (grouse and mountain hare). In grouse the change of numbers was correlated with that of recruitment. Autumn vole numbers peaked about a year before the other species and extensive forest damage occurred at winter peak densities of voles. These population fluctuations are consistent with a predator-prey model for their regulation. In short the model suggests that vole-food plant interactions trigger the cycle of voles, that voles generate the cycle of predators and that these in turn synchronize alternative prey populations to the others at vole declines.For voles, grouse and red fox the amplitude was higher in the first cycle compared to the second one whilst the opposite was true for the mountain hare. Although temperature and cone crops showed large interannual variations they still implied that herbivore food conditions were better during the former cycle. Hence, the reduction of the amplitude of the vole cycle may be explained by inter-cyclic differences in plant food conditions, implying food shortage (as indicated by bark-eating) at different population levels. The similar decrease of grouse and red fox populations may also be explained by deteriorated food conditions and/or for the fox by an outbreak of sarcoptic mange (Sarcoptes scabiae var. vulpes). The increased amplitude of the mountain hare cycle was part of a long-term rise in numbers after a tularemia epidemic in 1967. This is interpreted as a recovery, probably towards the generally higher pre-epidemic population level.     

东方田鼠种群密度制约的迟滞效应

本研究中用成年东方田鼠不同时间不同密度笼养后（笼内雌雄各半）,低密度配对饲养,观察各阶段东方田鼠繁殖指标的差异。试鼠144只,处理时分为LL（低密度长时间）（2只/笼,共20笼,90d）、HL（高密度长时间）（8只/笼,共5笼,90d）、HM（高密度中等时间）（8只/笼,共4笼,20d）与HS（高密度短时间）（8只/笼,共4笼,10d）4组。处理后在低密度条件下观察繁殖情况（胎仔数、怀孕率、分娩频率以及产仔间隔等）,直至180 d。实验数据按时间划分为3个部分统计：0~90 d为第一阶段(不同密度处理期),90~109 d为过渡阶段（处理后的过渡期）,109 d以后为第二阶段（正常低密度配对繁殖期）。结果表明：经过长时间高密度处理后,平均产仔数显著减少,组间平均分娩频率无显著差异。怀孕率由HL组到HM组,再到LL和HS组显著增加。在第一阶段,除了LL组之外,其余各组都未发现繁殖。过渡阶段的怀孕率组间差异显著,LL组以及HM组显著高于HL组。第二阶段的怀孕率以及胎仔数的组间差异显著,都为LL、HS两组较高,HL、HM两组较低。平均分娩频率、平均胎仔数以及产仔间隔各组间无显著差异。结论：不同持续时间的密度效应有较大差别。对于东方田鼠,20d的高密度处理相比于10d更能对其繁殖起到抑制作用。恢复低密度后,存在种群的迟滞性密度制约。     

Alternative male reproductive tactics in a natural population of prairie voles <Emphasis Type="Italic">Microtus ochrogaster</Emphasis>

Alternative reproductive tactics have been described in male mammals, but little information exists regarding fitness benefits and whether males change tactics. Adult male prairie voles Microtus ochrogaster (Wagner, 1842) display alternative tactics described as resident and wanderer. Enclosure studies provide conflicting data concerning the relative success of each tactic and whether males display one tactic throughout adulthood. To characterize further residents and wanderers in this species, we examined data collected during 5 years of monitoring a natural population in Illinois, USA. We found that during the breeding period, wandering males survived longer, moved longer distances, and were more likely than residents to have scrotal testes. During the nonbreeding period, wandering and resident males differed only in whether or not they established residency. Data on sources and fates of resident and wandering males revealed that a substantial proportion of males switched tactics. Our estimate of the reproductive contribution of wandering males to the population, which is based on the premise that wandering males typically mate with single females, suggests that wanderers contribute 34–38% of young recruited during March through October and 4–12% in November, when single females are less common. Parentage studies in natural populations are necessary to test our estimates.     

Diabetes and myocarditis in voles and lemmings at cyclic peak densities—induced by Ljungan virus?

Although it is well-documented from theoretical studies that pathogens have the capacity to generate cycles, the occurrence and role of pathogens and disease have been poorly empirically studied in cyclic voles and lemmings. In screening for the occurrence of disease in cyclic vole and lemming populations, we found that a high proportion of live-trapped Clethrionomys glareolus, C. rufocanus, Microtus agrestis and Lemmus lemmus at high collective peak density, shortly before the decline, suffered from diabetes or myocarditis in northern Scandinavia. A high frequency of animals had abnormal blood glucose (BG) levels at the time of trapping (5–33%). In contrast, C. rufocanus individuals tested at a much lower overall density, and at an earlier stage relative to the decline in the following cycle, showed normal BG concentrations. However, a high proportion (43%) of a sample of these individuals kept in captivity developed clinical diabetes within five weeks, as determined by BG levels and a glucose tolerance test performed at that later time. A new picornavirus isolated from the rodents, Ljungan virus (LV), was assumed to cause the diseases, as LV-induced diabetes and myocarditis, as well as encephalitis and fetal deaths, were observed in laboratory mice. We hypothesize that LV infection significantly affects morbidity and mortality rates in the wild, either directly or indirectly, by predisposing the rodents to predation, and is at least involved in causing the regular, rapid population declines of these cyclic voles and lemmings. Increased stress at peak densities is thought to be an important trigger for the development of disease, as the occurrence of disease in laboratory mice has been found to be triggered by introducing stress to LV-infected animals.     

To breed, or not to breed? Predation risk induces breeding suppression in common voles

Breeding suppression hypothesis (BSH) predicts that, in several vole species, females will suppress breeding in response to high risk of mustelid predation; compared to breeding females, suppressing females would gain higher chances of survival. Seminal evidence for BSH was obtained in the laboratory, but attempts to replicate breeding suppression under field conditions were less conclusive. We tested whether breeding suppression occurs in common voles (Microtus arvalis), and how population density and predation risk combined affect voles’ reproductive activity. We found that, in contrast to males, female common voles suppress reproductive activity when faced with high predation risk. Population size was not reduced despite breeding suppression. A model of the interaction between predation risk and population density revealed that predator-induced breeding suppression depends on the density of conspecifics. We concluded that breeding suppression is a viable adaptation only at low vole densities, when per capita predation risk is high. Finally, we identified the key issues of experimental design required for the consistency of future studies on breeding suppression.     

Tawny owl reproduction and offspring sex ratios under variable food conditions

Tawny owl reproduction and offspring sex ratios have been considered to depend on the abundance of small voles. We studied reproductive performance (laying date, clutch and brood size) during 1995–2003 and offspring sex ratios from 1999 to 2003 in relation to the abundance of small voles and food delivered to the nest in a tawny owl population in southern Finland. Abundance of small voles (field and bank voles) was based on trappings in the field, and estimates of food delivery was based on diet analysis of food remains in the nest boxes. In this population, reproductive output was not related to the abundance of small voles. Analysis of food delivered to the nest showed that the prey weight per offspring varied more than twofold between years and revealed that this difference was mainly related to the proportion of water voles in the diet. Only the number of water voles correlated with laying dates. Offspring sex ratios were weakly male biased (55%) but did not differ from parity. Sex ratios were not related to the abundance of small voles, and we found no evidence that parents delivered more food to nests with proportionally more offspring of the larger (female) sex. Our results underline the notion that populations may differ in their sex allocation pattern, and suggest such differences may be due to diet.     

Does avian predation risk depress reproduction of voles?

Reproductive output and the growth of captive voles were quantified under high and low avian predation risk in a semi-natural experiment. Voles were exposed to Eurasian kestrels (Falco tinnunculus), the main avian predator of vole species studied (Clethrionomys glareolus, Microtus agrestis and M. rossiaemeridionalis). Vole pairs were housed in cages settled under nest-boxes occupied by breeding kestrels or in control cages settled under empty nest-boxes for 2 weeks. The experiment was conducted in mid-summer when kestrels had half-grown nestlings, because in that time hunting adults and begging nestlings produce noise and scats which may indicate significant predation threat to voles housed underneath the nest-boxes. The risk of kestrel predation did not have any obvious impact on pregnancy rates, mean litter sizes, or growth rates of kestrel-exposed voles compared with control voles studied. These results indicate that the risk of avian predation does not depress the reproductive investment of voles. Received: 3 November 1997 / Accepted: 16 February 1998     

Induction of mictic females in the rotifer Brachionus: oocytes of amictic females respond individually to population-density signal only during oogenesis shortly before oviposition

1. One at a time during the reproductive period of amictic females, oocytes fill with yolk and undergo a mitotic maturation division (oogenesis), are oviposited as single cells, and then develop parthenogenetically into females. Sexual reproduction in Brachionus and several other genera is initiated when amictic females are crowded and oviposit some eggs induced to differentiate into mictic females. Mictic females produce haploid eggs that can develop parthenogentically into males or be fertilised and develop into diapausing embryos called resting eggs. 2. This study examines the time when oocytes in amictic females respond to maternal population density. Is the fate of all oocytes in the germarium irreversibly determined during the early postnatal life of the mother, or is each oocyte labile until just before oviposition? In the former case, the probability of an amictic female producing a mictic daughter at any time throughout her reproductive period would reflect the population density she experienced while young and not that at the time she oviposited an egg. 3. Amictic females of two clones of a Florida strain of B. calyciflorus were cultured singly from birth at a low or high density (in a large or small volume) until about halfway through their reproductive period and then switched (experimental treatment), or not (control treatment), to the other density condition. The results indicate that the female fate of an oocyte is determined by maternal population density during oogenesis. Eggs oviposited soon after transfer from low to high density had the same, or a higher, probability of becoming mictic females compared with those produced by control females kept at the high density; eggs oviposited after transfer from the high to the low density had the same low probability of becoming mictic females as those produced by control females kept at the low density. 4. Control females kept at the high density were less likely to produce mictic daughters as they aged. This decline is not because of a decreased propensity of older females to respond to crowding, as older females responded maximally when transferred from a low to a high population density. 5. As oocytes in amictic females respond to maternal population density only during oogenesis, there is a negligible lag between the population‐density signal in the environment and the commitment to sexual reproduction. This minimises the obligatory two‐generation lag between this signal and production of resting eggs, and thus reduces the possibility that crowding will lead to food limitation before production of these eggs.     

Characteristics of erythrocyte lipids in blood of tundra voles (Microtus oeconomus Pall.) inhabiting areas with increased natural radioactivity

Interrelations between the lipid characteristics of the blood erythrocytes and 226Ra accumulation in the body of tundra voles (Microtus oeconomus Pall.) inhabiting areas with different levels of the radiation background were investigated. It is shown that the ratio of the phospholipid (PL) fractions which cause the blood erythrocyte structure depends on the phase of the population cycle, as well as on the sex and age of tundra voles. The statistically significant interrelation between lysoforms and the sphingomielin content has been revealed in the blood erythrocyte PL of tundra voles; its scale somewhat differs for the animals from the reference and Ra areas. The peroxide concentration in the blood erythrocyte lipids of tundra voles from the Ra area exceeded the control values in all mature groups of the animals trapped at the depression phase of the population density. The 226Ra content in the bodies of the tundra voles which were trapped in the Ra area at the increased phase of the population cycle is for certain higher than that for the animals from the reference area. Interrelations between the lipid peroxidation parameters in the blood erythrocytes and the body 226Ra content for separate sex-age groups of tundra voles have been revealed.