Predation by Tawny owls (Strix aluco) on Bank voles (Clethrionomys glareolus) and Wood mice (Apodemus sylvaticus)

During 1954–56 we made a study of the numerical relationships between a population of Tawny owls ( Strix aluco ) and populations of their main prey species, the Bank vole ( Clethrionomys glareolus ) and the Wood mouse ( Apodemus sylvaticus ), in a 48.6-ha area of mainly deciduous woodland near Oxford. The owls were censused by plotting their territorial challenges (hooting), the rodents by the capture-mark-recapture technique. Since the rodents were marked with numbered monel metal leg rings, we were able to recover a proportion of these in the pellets of undigested material cast by the owls.
The results for Bank voles, which were the more numerous of the two prey species, indicated that 20–30% of the standing crop was removed by owls in any two-month period. We also made an independent estimate of predation rate from the number of rings recovered from owl pellets in relation to the number of ringed rodents released into the population each two months. This coincided with the limits indicated by the first method for voles (20-30% removed of the standing crop each 2 months).
Wood mice were scarcer than Bank voles and were not amenable to satisfactory capturemark-recapture analysis but, when treated on the system of recovery of rings in owl pellets, we found that they were preyed upon relatively more heavily than were the voles. Of the latter 18–46% of the marked animals were recovered in the owl pellets compared with 28–70% of the marked mice. Either the mice were preferred prey or they were more vulnerable to owl predation by reason of their preference for more open habitats.     

A yardstick for measuring populations of small rodents

(1) Six-monthly trappings (summer and winter) were done between 1952 and 1969 of Wood mice (Apodemus sylvaticus) and Bank voles (Clethrionomys glareolus) in Wytham Woods, near Oxford. One hundred and fifty Longworth live traps were used on two 2-acre grids in oak-ash-sycamore woodland. (2) The traps were pre-baited for 48 hours and then set. The catch was examined on the evening of the same day and on the following morning. The total of animals caught during the 24 hours was used as an index to the actual numbers present. Animals retrapped in the morning (identified by a simple method of marking) were omitted from the total. (3) Between 1969 and 1972 the same procedure was used but trapping, evening and morning, was continued for 3-4more days until some 75–80% of the catch consisted of marked animals. (4) This enabled estimates to be made of the actual numbers of voles and mice present at each trapping and so to assess the proportion of the total present which was caught ring the first 24 hours. (5) These estimates were made by (Method la) a straight Lincoln Index, (Method lb) a Lincoln Index method as modified by Hayne (1949) and (Method 2) by Hayne's trap-out method. Over six trappings the estimates by these methods showed reasonable agreement. (6) Comparison of the index trapping (i.e. the numbers caught during the first 24 hours) with the estimates of the total numbers present showed that, on all occasions, more than half of the mice and voles present entered the traps during the first 24 hours. With Bank voles there was a tendency for about half of them to be trapped during this time during the winter and about three-quarters of them during the summer. (7) With Wood mice the figures are more variable but, again, never less than half were trapped during the first 24 hours and, on some occasions, nearly the whole of the population. (8) These results show that reasonably accurate estimates of the numbers of these rodents can be achieved within 3–4 days' trapping (estimates that can be cross-checked by different methods) by using a high density of traps (see (1) above). Furthermore, trapping for 24 hours only will always catch at least half of the population present and, on occasions, considerably more than half.     

Species and individual differences in juvenile female alloparental care are associated with oxytocin receptor density in the striatum and the lateral septum

The neuropeptide oxytocin has been implicated in the regulation of affiliative behavior and maternal responsiveness in several mammalian species. Rodent species vary considerably in the expression of juvenile alloparental behavior. For example, alloparental behavior is spontaneous in juvenile female prairie voles (approximately 20 days of age), takes 1-3 days of pup exposure to develop in juvenile rats, and is nearly absent in juvenile mice and meadow voles. Here, we tested the hypothesis that species differences in pup responsiveness in juvenile rodents are associated with oxytocin receptor (OTR) density in specific brain regions. We found that OTR density in the nucleus accumbens (NA) is highest in juvenile prairie voles, intermediate in juvenile rats, and lowest in juvenile mice and meadow voles. In the caudate putamen (CP), OTR binding was highest in prairie voles, intermediate in rats and meadow voles, and lowest in mice. In contrast, the lateral septum (LS) shows an opposite pattern, with OTR binding being high in mice and meadow voles and low in prairie voles and rats. Thus, alloparental responsiveness in juvenile rodents is positively correlated with OTR density in the NA and CP and negatively correlated with OTR density in the LS. We then investigated whether a similar receptor-behavior relationship exists among juvenile female prairie voles by correlating individual variation in alloparental behavior with variation in OTR density. The time spent adopting crouching postures, the most distinctive component of alloparental behavior in juveniles, was positively correlated with OTR density in the NA (r = 0.47) and CP (r = 0.45) and negatively correlated with OTR density in the lateral septum (r = -0.53). Thus, variation in OTR density in the NA, CP, and LS may underlie both species and individual differences in alloparental care in rodents.     

Woodland small mammal population ecology in Britain: a preliminary review of the Mammal Society survey of Wood Mice Apodemus sylvaticus and Bank Voles Clethrionomys glareolus, 1982–87

The Mammal Society has co-ordinated a population survey of Wood Mice Apodemus sylvaticus and Bank Voles Clethrionomys glareolus in 13 0.81 -hectare sites in Britain. Numbers of mice and voles live-trapped using standard methods were collated every May/June and November/December from 1982 to 1987. The data were analysed with results from four independent studies in England and the corresponding assessments of tree seed crop size. Wood Mouse numbers are usually higher in winter than in summer but Bank Vole fluctuations are less regular. In deciduous woodland, Wood Mouse mean relative densities are significantly greater in the winter and the following summer after a good seed crop than after a poor one; rates of population change from summer to winter are significantly higher when a good seed crop falls. Bank Vole relative densities are significantly greater in the summer following a good seed crop than after a poor one, and rates of change from winter to summer are significantly higher. In Wood Mouse populations, tests for density dependence suggest that it is strong from summer to winter but absent from winter to summer; in Bank Voles weaker density dependence is present in both halves of the year. Thus, Wood Mouse numbers are regulated in autumn but are also influenced by seed crop size in winter and the following summer; Bank Vole numbers are less strongly regulated during both autumn and spring and are influenced by seed crop size in the following summer. Evidence is presented suggesting that populations of each species in deciduous woodlands are synchronized over the country in summer and that Wood Mice are also synchronized in winter; highs and lows tend to coincide between different sites. The yield of tree seed is shown to vary significantly from year to year and may be the cause of the synchrony, but weather effects may also be involved.     

Multiple captures provide evidence of small mammal social behavior

Social behavior of small mammals living under natural conditions often is inferred from live-trapping data, particularly from incidents in which two or more individuals are captured together in a trap. We examined whether multiple-capture data from a long-term study of prairie voles (Microtus ochrogaster) and meadow voles (Microtus pennsylvanicus) were consistent with well-known species differences in social behavior (whereas prairie voles are highly social and display monogamy, meadow voles are less social and promiscuous). When possible, we also examined multiple captures of two nontarget species, northern short-tailed shrews (Blarina brevicauda) and western harvest mice (Reithrodontomys megalotis). Percent of total captures that were multiple captures and percent of total adult captures that were male–female captures were highest for prairie voles and lowest for meadow voles; values for harvest mice and shrews were in between those of the vole species, but more similar to values for meadow voles. Repeat captures of the same male–female pair occurred most commonly in prairie voles, and multiple captures of this species typically involved individuals from the same social group. Multiple captures of adults and juveniles were more common in prairie voles than meadow voles, except for captures of at least one adult male and at least one juvenile, which did not differ between the two vole species. Multiple capture data for prairie voles and meadow voles were largely consistent with established species differences in social behavior, suggesting that such data can provide an accurate indication of social and mating systems of small mammals.     

The ecology of Wood mice (Apodemus sylvaticus) on arable farmland

The ecology of Wood mice ( Apodemus sylvaticus ) was studied by live trapping and by examining the stomach contents of killed animals on an arable farm in West Suffolk. The study area included several fields but only a short length of hedgerow and was distant from other hedges and woods. Wood mice were trapped in the open fields throughout the year and their burrows were also found there. Population densities were comparable with those reported from deciduous woodland and seasonal population changes, ranging behaviour and timing of the breeding season were also broadly similar. The survival rate of juvenile mice varied greatly within and between breeding seasons tending to improve when young of the year replaced overwintered animals in the adult breeding population. Wood mice showed no marked preference for fields with different crops in winter when the main foods were grain, waste sugar beet roots, weed seeds and soil invertebrates. In spring and early summer they preferred winter wheat fields where seeding weeds provided most of their food.     

The dynamics and regulation of small rodent populations in the woodland ecosystems of Killarney, Ireland

Populations of Wood mice Apodemus sylvaticus and Bank voles Clelhrionomys glareolus were studied by mark-recapture techniques in five different woodlands but concentrated on an oak Quercus petraea wood and a yew Taxus baccata wood, where food supply was also investigated. Much of the yew wood was on limestone pavement which was greatly creviced thus providing static and quantifiable cover. Indices of the intensity with which food was being searched for were obtained on all areas by seed-removal experiments.
Trappability varied within the populations and seasonally, being highest in summer. Unusually high densities of mice were recorded in the yew wood but not elsewhere; vole densities were always low. There was an inverse relationship between range length and density: mean range length in the yew wood was exceptionally low. Males generally ranged further than females, voles further than mice. Females outnumbered males in the oak wood, where cover was minimal, when the population was low. In the two areas with most cover, males outnumbered females. In a further area, where numbers were initially exceedingly low, the population consisted entirely of males for almost a year. Sexes differed in habitat preference but numbers, especially of males, were significantly correlated with cover. Voles usually bred for shorter periods than mice. Overwinter breeding and increased overwinter survival followed good seed years. Individuals recruited when food was plentiful lived longest. In seed-removal experiments rate of removal increased with shortage of natural seed.
Food and intraspecific aggression were the probable major factors in regulating population size, of mice at least. Cover probably modifies intrinsic mechanisms by reducing numbers of encounters between individuals.     

DISPERSAL IN THE WOOD FROG (RANA SYLVATICA): IMPLICATIONS FOR GENETIC POPULATION STRUCTURE

Recapture of marked juvenile and adult wood frogs in five Appalachian Mountain ponds showed adults to be 100% faithful to the ponds in which they first bred, but approximately 18% of the juveniles dispersed to breed in ponds other than the one of origin. Effective population sizes were generally smaller than the population censuses and genetic neighborhoods had an average radius of 1,126 meters. Values of standardized genetic variance based on effective population size and mating success were relatively small. Genetic population structure estimated from the dispersal data suggested that ponds within about a 1,000 meter radius should show little genetic differentiation; ponds separated by a distance greater than 1,000 meters should experience little gene flow and show higher genetic differentiation. Wood frogs in these ponds do not show a meta-population structure as suggested for newts.     

Maturity, castration and behaviour of male Bank voles, Clethrionomys glareolus (Mammalia)

This paper discusses the relationship between aggressive behaviour and sexual maturity, prepubertal castration and wounding in male Bank voles, Clethrionomys glareolus. In neutral arena tests in the laboratory, sexually mature males exhibited more aggressive components of behaviour than immature males or than adult males that had been castrated prepubertally. Mature males also exhibited fewer amicable components than the immature or castrated males. During several months residence in large outdoor enclosures, mature males consistently had more fresh skin wounds (inflicted by other voles) than did either immature or castrated males. The relevance of these results to the population ecology of Bank voles is discussed.     

Natal dispersal in relation to population density and sex ratio in the field vole,Microtus agrestis

Summary In a sample of 240 juvenile field voles 8% of the males and 22% of the females reached sexual maturity within their natal home range. Among individuals retrapped as adults, 58% of males and 23% of females had dispersed, i.e. had moved more than one home range diameter. The mean distance moved for males (58.5 m) exceeded that for females (28.6 m). Male movement distances were negatively associated with total density, and with density of adult females, but not with male density. Female movements were not related to population density. There were no relation between sex ratio and distance moved. The distribution of distances moved for both males and females fit a geometrical distribution, suggesting the importance of competitive processes.     

Life in varying environments: experimental evidence for delayed effects of juvenile environment on adult life history

1.?The effects of environment experienced during early development on phenotype as an adult has started to gain vast amounts of interest in various taxa. Some evidence on long-term effects of juvenile environment is available, but replicated experimental studies in wild animals are still lacking. 2.?Here we report the first replicated experiment in wild mammals which examines the long-term effects of juvenile and adult environments on individual fitness (reproduction, survival and health). The early development of bank vole (Myodes glareolus) individuals took place in either food-supplemented or un-supplemented outdoor enclosures. After the summer, adult individuals were reciprocally changed to either a similar or opposite resource environment to overwinter. 3.?Adult environment had an overriding effect on reproductive success of females so that females overwintering in food-supplemented enclosures had a higher probability of breeding and advanced the initiation of breeding. However, the characteristics of their litters were determined by juvenile environment: females initially grown in food-supplemented conditions subsequently produced larger litters with bigger pups and a male-biased sex ratio. 4.?In males, individuals growing in un-supplemented conditions had the highest survival irrespective of adult environment during winter, whereas in females, neither the juvenile nor adult environments affected their survival significantly. The physiological condition of voles in spring, as determined by haematological parameters, was also differentially affected by juvenile (plasma proteins and male testosterone) and adult (haematocrit) environments. 5.?Our results suggest that (i) life-history trajectories of voles are not strictly specialized to a certain environment and (ii) the plastic life-history responses to present conditions can actually be caused by delayed effects of the juvenile environment. More generally, the results are important for understanding the mechanisms of delayed life-history effects as well as recognizing their population dynamic consequences.     

Post‐breeding movements of Wood Storks in Brazil and Argentina

Seasonal migration and the dispersal of juvenile and adult Wood Storks (Mycteria americana) after breeding have been documented in the United States, but little is known about the post‐breeding movements of Wood Storks in South America. Our objective was to identify the locations of post‐breeding areas used by Wood Storks banded as nestlings in breeding colonies in Brazil by analyzing banding data. During the period from 1984 to 2007, 2543 nestlings were banded at breeding colonies in three regions of Brazil, with most (94%) banded in the Pantanal wetland in west‐central Brazil. Seventeen bands were subsequently recovered, with most (14) recovered in southern Brazil and northern Argentina. The mean distance between banding and recovery sites was 1265 km. Our results suggest that Wood Stork movements from breeding areas in Brazil are, as also reported in the United States, in response to changing water levels. The rainy season begins at the end of the breeding season and, in apparent response to rising water levels, Wood Storks in our study moved to drier areas further south with shallower water where they can forage more efficiently. Because only a small percentage of the area where Wood Stork bands were recovered in our study is currently protected, measures are needed to prevent habitat destruction and preserve wetland habitats used by Wood Storks during the post‐breeding period in southern Brazil and Argentina.     

布氏田鼠种群生理年龄结构的研究

本文根据胴体重分布将布氏田鼠划分为越冬鼠与当年鼠两组,然后再根据生理学指标将当年鼠划分为性成熟鼠(成年)和性未成熟鼠(幼年)。由于生理年龄与时间年龄不一致,故本文并不强调各种形态指标来划分时间年龄组。5月下半月以前种群主体为越冬鼠,但在当年鼠出现地表面后,当年鼠取代越冬鼠成为种群主体,到7月下半月越冬鼠已降到10％以下。当年生的幼鼠春季生长发育旺盛,很快成为当年成鼠。晚夏和秋季出生的个体当年性不发育成熟。秋季种群主体为当年幼鼠。越冬鼠和大部分当年成鼠从种群中消失。比较种群数量上升与下降年份,春季越冬鼠胴体重的轻重与第一批当年鼠(K_1)出现规模可能是种群数量升降的重要指标。     

The suprachiasmatic nucleus in organotypic slice cultures of the common vole (Microtus arvalis): comparison of development with rat and hamster and the effect of age

The intrinsic properties of the suprachiasmatic nucleus (SCN), the site of the main circadian pacemaker in mammals, have recently been studied in vitro by means of organotypic slice culturing. So far, only neonatal rats and mice have been used for such developmental and functional analyses of the isolated pacemaker. Here, the authors present a comparative developmental study of the SCN of voles, rats, and hamsters in organotypic slice cultures. In contrast to strictly circadian organization of behavior in rats and hamsters, common voles (Microtus arvalis) are characterized by large variability in the strength of circadian organization of behavior. It is not known to what extent this variability is reflected in the intrinsic features of the SCN. Cultures were prepared from rat, hamster, and vole pups (6 to 9 days old) for the purpose of species comparison. In addition, the authors studied the relation between age and development in cultures from pup (7 to 10 days old), juvenile (15 to 16 days old), and young adult (1 to 2 months old) voles. In contrast to the situation in rat and hamster, the most striking feature in neonatal voles is the variability in shape of the final, fully developed culture and its poor resemblance with the in vivo SCN. The SCN of adult voles, however, could be cultured successfully while retaining its morphological organization seen in situ. Phase-contrast microscopy and immunocytochemical staining for vasopressin and glial fibrillary acidic protein revealed that cultures of pup and juvenile voles still have potential for neurogenesis and morphological reorganization. Young voles, therefore, can serve as a model to study the developmental establishment of a functional circadian pacemaker, while adult voles allow the study of intrinsic pacemaker properties in relation to previously recorded behavior of the donor and aging-related pacemaker dysfunction.     

The effect of varying calcium intake on calcium metabolism in wild rodent species

Calcium is essential for mammals but distributed heterogeneously in the environment and so the calcium content of the diet of wild small mammals can vary with habitat geology. The effect of varying the calcium intake of two wild rodent species, the granivorous wood mouse Apodemus sylvaticus and the herbivorous bank vole Clethrionomys glareolus , was investigated. Wood mice were similar to laboratory rodents in their calcium metabolism. Net calcium gut absorption efficiency was 11–15% in wood mice fed either high (0˙87%) or low (0˙30%) calcium diet. No more than the equivalent of 6% of the ingested calcium load was excreted in urine but this was sufficient to eliminate the equivalent of all the extra calcium absorbed across the gut when mice were fed the high calcium diet. In contrast, mean net calcium gut absorption efficiency in bank voles was high and increased from 28% to 42% when dietary calcium content was tripled. Urine was important for excreting calcium, the equivalent of up to 23% of intake being eliminated by this route. Despite this, bank voles fed the high calcium diet assimilated three times more calcium than voles fed the low calcium diet; this was in direct proportion to the three-fold difference in dietary calcium concentration. Possible implications for wild rodents in calcium-rich and calcium-poor habitats are discussed.     

Small mammal populations in relation to hedgerow structure in an arable landscape

The population ecology of small mammals in hedgerows in arable farmland in eastern England is described. Features of hedgerows of importance to individual species are examined. Some 97% of the total 3042 mammals captured were wood mouse Apodemus sylvaticus , yellow-necked mouse Apodemus flavicollis , bank vole Clethrionomys glareolus and common shrew Sorex araneus . Small numbers of harvest mice Micromys minutus , field voles Microtus agrestis , pygmy shrews Sorex minutus and water shrews Neomys fodiens were also caught. Wood mouse, the most numerous species, showed a typical pattern of large numbers in autumn and winter, followed by a simultaneous decline over all hedges in early spring. Population changes were less clear in yellow-necked mouse and bank vole but the yellow-necked mouse was more scarce in the second year of study. Common shrews were most numerous in summer and declined rapidly in autumn. Hedgerow coppicing had a marked effect on yellow-necked mouse numbers but not on wood mouse. In an extensive survey of mammal numbers in relation to hedgerow features, ground cover was found to be the single largest factor influencing size of bank vole populations. Hedgerow condition (lack of gaps) was important to yellow-necked mice, which thrived only in well-established hedgerows. Wood mice appeared little influenced by the characteristics of the hedge. Common shrews were more abundant in hedgerows with adjacent permanent water.     

Effect of conspecifics on sexual maturation in female European pine voles (Pitymys subterraneus)

Using the number of large ovarian follicles (Type 8) as an indicator of sexual maturation we found that urinary compounds released by adult males accelerated puberty while urine from females suppressed hormonal activity in juvenile female European pine voles. The release of chemosignals that delayed puberty of juvenile females was not influenced by ovarian hormones; urine from ovariectomized females was as effective as urine from unoperated animals.     

Recognition of interspecific familiar versus unfamiliar odours among bank voles and yellow-necked mice

Two experiments (laboratory and field-laboratory) were designed to determine whether individual bank volesClethrionomys glareolus (Schreber, 1780) and yellow--necked miceApodemus flavicollis (Melchior, 1834) could distinguish heterospecific odour cues from familiar and unfamiliar individuals. In the laboratory experiment each male bank vole was familiarized for 24 h with odour (cotton wool impregnated with urine and faeces) of male yellow-necked mice and yellow-necked mice were familiarized with odour of male bank voles. In the field-laboratory experiment the individual bank voles and yellow-necked mice captured at the same point were considered familiar and transfered to the laboratory. In laboratory, these individuals were tested in a box (for 5 h) affording them the choice between the odours of familiar and unfamiliar heterospecific males. Bank voles discriminated between familiar and unfamiliar yellow--necked mouse odours. Male yellow-necked mice seemed to have a similar ability to recognise odours of familiar bank voles. It is proposed that interactions between these two species occur not only on the species level, but also on the level of individual. This phenomenon (probably asymmetric) can play an important role in spatial orientation, and influence direct contacts between individuals of these species.     

Parentage assignment detects frequent and large-scale dispersal in water voles

Estimating the rate and scale of dispersal is essential for predicting the dynamics of fragmented populations, yet empirical estimates are typically imprecise and often negatively biased. We maximized detection of dispersal events between small, subdivided populations of water voles (Arvicola terrestris) using a novel method that combined direct capture-mark-recapture with microsatellite genotyping to identify parents and offspring in different populations and hence infer dispersal. We validated the method using individuals known from trapping data to have dispersed between populations. Local populations were linked by high rates of juvenile dispersal but much lower levels of adult dispersal. In the spring breeding population, 19% of females and 33% of males had left their natal population of the previous year. The average interpopulation dispersal distance was 1.8 km (range 0.3-5.2 km). Overall, patterns of dispersal fitted a negative exponential function. Information from genotyping increased the estimated rate and scale of dispersal by three- and twofold, respectively, and hence represents a powerful tool to provide more realistic estimates of dispersal parameters.