Population dynamics of the Smith's red-backed vole in highlands of Shikoku

Population dynamics of theSmith 's red-backed vole predominantly common through uplands of Shikoku have been in some degree disclosed by the use of my own and Government Forest Station's samples collected since 1955 onward. It has proved that the upper-range population reaches its density peak possibly in late summer or early fall, but the lower-range one does probably in October–November as the seasonal trend in usual years. The upper one produced a peak three times at 3–4 year intervals, the first peak being an outbreak followed by a crash, during the last decade. It seems likely that all the populations through the range have, in the gross, fluctuated in phase after 1959 at least. The cyclic fluctuation may readily be explained by the theory of intrinsic mechanism, because no external factors are considered to have exerted a conclusive effect. Except what was concerned in the outbreak, the role of the social stress could be set at naught. The regulation of fecundiy by density was expressed more markedly in litter size and less in active reproductivity rate and incidence of pregnancy. The mean litter size at term as small as 2.00 is contrary to our expectation in view of the supposed prolific potential, nevertheless the observed frequency of litter poduction and intra-uterine survival rate has proved not to be so high as to make up for the small litter size. The problem in the postnatal mortality has remained to be solved.     

Investigation into the edge effect by use of capture-recapture data in a vole population

A substantial explication of the edge effect has been attempted by use of capture-recapture data for a vole population (Microtus montebelli), gathered intwo plots of 100×100 m or less during 12 days, cheked twice daily, in August 1970; the sample was quite sufficient for the aim. The edge effect as guessed by increased catch per trap is usually suspected to ensue from range-settlers in the outside boundary strip of a plot and immigrants. But by a theoretical analysis I could attain a tentative conclusion that no increased catch per trap will occur unless any invasion takes place. Then it follows that, apart from the effect of invasion, the role of the adjoining outside settlers in the edge effect is essentially required to be studied in the light of knowledge on the truth of size and shift in home range. The variation in range behavior for 183 adult voles, captured 6 times or more, could be grouped into eight types, of which the range-conservative type possessed 52% of the sample and the group of the type was justly utilized for giving averages of range size. Besides, it was seen from the observed frequency of types that a considerable number of immigrants onto the census plot were induced perhaps being allured by trap baits, but the majority of them proved to be assigned to the voles that have their ranges inside the assessment line ofDice ; the rest referable to effective immigrants was only a few (7%). I could perceive no reason such as disproves the idea ofDice 's additional boundary strip. Viewed from maps of ingress shift of ranges, the effect of ingress must have been greater in the outer trap rows than in the inner within the plot, so that it might well be called edge effect in general; such effect, however, is seen gradually diminishing toward the center, and hence it is almost unlikely that one should find any clear-cut intra-plot assessment lines demarcating such an inner square as quite free from edge effects. Averages of observed range length and width (ORL and ORW), as reliable measures for the true range size, were determined from the above group of specimens; as a result, the remarkable concept of elliptic range shape was established by regarding ORL as long axis and ORW as short one, and, directly from these averages, the mean range sizes worked out at 0.04 for females and 0.09 for males in acreage which proved to be surprisingly well agreeable with those of isotope-revealed ranges for voles given byGodfrey (1954) andAmbrose (1969). The catchability for marked voles () was estimated by the maximum likelihood method by use ofJolly 's formulae (1965), but that for unmarked ones () was made by the regression census formula; as a result it was shown that the population was clearly of π>p type and that the trap-experience that voles underwent one month or more ago can make them retain as high catchability as π.     

Inquiry into effects of prebaiting on removal census in a vole population

A field work with two plots of grid, a snap trap being set on each station spaced 5 m apart, was executed in the summer of 1968 to evaluate prebaiting in census trapping by comparing the result in one plot, prebaited for three days, with that in the other not prebaited. Since the population was as high as some 230 per acre on the average in density and formed of the vole, Microtus montebelli, alone, sufficient samples were gathered irrespective of the plot size as small as 50×50 m. Owning to the circumstances, multiple collisions inflicted so intense influence on sampling especially in the prebaited plot that z-equation for census adjusted to the effect was well applicable to the data in either plot. In sampling, the fact that small voles are apt to be caught later than large voles was statistically evidenced in either sex, and yet any proof that males tend to be caught prior to females was not offered. It was ascertained in either plot that the daily catch was realized according to the same rule through the whole period of trapping in both external belt and internal square within the plot; hence it follows that no considerable immigration occurred. One of the beneficial effects of prebaiting is sure to be that the probability of capture was markedly enhanced in the prebaited plot, and a second is supposed, though inconclusively, to be that a good sampling could be executed consistently through the census period giving rise to no inordinate catches perhaps due to heterogeneous sampling as was seen in earlier days in the not prebaited plot. The supposition has derived from the condition that most of the whole population is trappable, which is established by interrelation among population density, size of home range and trap spacing. It was suggested that the effect of prebaiting should be evaluated from the view-point of the interrelation, because the basic utility of prebaiting consists in that it may help to our utlimate purpose to estimate the whole population.     

Adrenal analysis for critique of the social stress theory in natural populations of a montane vole

Summary The vole,Eothenomys smithi, in highlands of Shikoku was studied to disclose ups and downs of adrenal activities under the influence of the breeding level in natural populations. It has proved that the vole's adrenal in both sexes may be much more strongly by rise and fall of the fertility as well as by shift of the age structure in populations than the rat's. Therefore, as for perhaps all kinds of small mammals, we consider necessary to follow a pertinent procedure to avoid thoroughly these sources of error in studies to draw any conclusion for or against theChristian's theory. The present data analysed by such a method failed to afford any confirmative proofs; the fact that the pregnancy rate was increased in parallell to the density is against his view. However, never the whole content of the theory has been invalidated by these results.     

A field study of the effect of prebaiting on censusing by the capture-recapture method in a vole population

Populations of the vole, Clethrionomys rufocanus, in a lowland woodlot of Hokkaido were studied for the presence of effects of prebaiting on censusing by the capture-recapture method. A grid of 121 live-traps, spaced 5 m apart, was laid out on each of two plots, one of which alone was prebaited three days long. Owing to very high densities and great trap-efficiency, sufficient and favourable samples could be available for statistical analysis, except the trend of delayed catch for young in sampling. The population on either plot, however, proved to be markedly variable in catchability of unmarked animals in the course of trapping; while the probability of recapture was counted as invariable on the average from day to day, the recapture frequency was different between juveniles, subadults and adults. Needless to say, the catchability was distinctly greater for marked voles than for unmarked ones, whether prebaited or not, through the trapping period, except that the first-day catchability for unmarked ones on the prebaited plot seems not to be significantly lower than that for marked ones. Consequently, it turns out that the prebaiting has almost never helped to eliminate the important bias induced by differential trap-response of marked and unmarked animals; its contribution is only that the catchability for unmarked ones is slightly higher on the part of the prebaited plot on earlier days of the period. In accordance with the heterogeneous catchability, the Γ-form distribution analogous to the geometric could be applied with thorough fitness to the capture frequency in order to estimate the whole populations. The fact that the estimates are reliable, being not at any rate underestimates, was further confirmed by the result of a follow-up work conducted by means of the removal method with wider trap-spacing which brought forth distinct underestimation chiefly referable to unexposure to traps of the partial populations. The subject of unexposure was discussed by laying stress on the relation between minimum range length and trap-spacing.     

Ongoing ultradian activity rhythms in the common vole,Microtus arvalis,during deprivations of food,water and rest

Summary The timing mechanism underlying ultradian (2–3 h) activity patterns in the common vole, Microtus arvalis, was studied using behavioural deprivation experiments. These were aimed at distinguishing between a homeostatic control mechanism, in which the rhythmic behaviour itself is part of the causal loop, and a clock mechanism, independent of the behaviour.In 175 experiments, deprivation of food during 3 ultradian cycles in (subjective) daytime did not result in significant changes in the ultradian periodicity of attempts to obtain the food, compared with ad lib. access to food and water. A minor, but significant increase in ultradian activity time () occurred in the course of the deprivation, but this was compensated by a shorter ultradian rest (). These results were obtained both in intact animals (n = 24), which showed ultradian and circadian rhythmicity in behaviour, and in animals (n = 21) with electrolytic lesions aimed at the suprachiasmatic nuclei (SCN), which lacked the circadian modulation of behaviour. Simultaneous deprivation of water and food in 8 voles without circadian rhythmicity during 40 experiments also did not lead to any change in the ultradian periodicity of feeding attempts.Rest deprivation was studied in 5 SCN lesioned voles, by forcing running wheel activity to continue following spontaneous running. Thus, the experimental activity bout was artificially lengthened to 2–9 h in 67 experiments. The onset of the subsequent rest episodes occurred independent of the duration of the preceding . The duration of was dependent on the preceding, experimental in a periodic fashion. The interval experimental (=lengthened +following ) was equal to one, two or three times the control (obtained on nonexperimental days). This result fits the prediction of a clock model and is in conflict with a monotonicincrease of with , as expected in a homeostatic, restorative process.It is concluded that the ultradian timing of activity in the common vole can be explained neither by homeostatic hunger or thirst mechanisms nor by homeostatic rest/activity regulation. The results strongly suggest an independent clock system generating ultradian feeding rhythms in the common vole.Abbreviations DD continuous darkness - LD light-dark regime - LL continuous light - RCA retrochiasmatic area - ARC arcuate nucleus - SCN suprachiasmatic nuclei - ultradian period - ultradian activity time - ultradian rest time     

A possible discrepancy between the exposed and the whole population depending on range-size and trap-spacing in vole populations

Population Ecology - From a field study for the vole population (Clethrionomys rufocanus) in Hokkaido in the late summer of 1965, it has been proved that the range length may decrease from 25 to 18...     

Climate change in Norway: warm summers limit grouse reproduction

Grouse and vole numbers may peak after peaks in the seed crop of bilberry (Vaccinium myrtillus) because of reduced levels of feeding deterrents in bilberry plants. We predicted that grouse reproduction depends also on summer (June–September) temperatures in the 2 previous years, because bilberry plants will be less exhausted after a high seed crop in or after warm summers, and thus rebuild their chemical defence more quickly. After berry peak years, population indices of capercaillie (Tetrao urogallus) and bank vole (Myodes glareolus) in southern Norway were negatively related to summer temperatures in the previous year or previous 2 years. Willow grouse (Lagopus lagopus) chick production in five areas in Norway was negatively related to summer temperatures in the 2 previous years when controlling for vole density. A similar pattern was found for the bilberry-feeding moth (Eulithis populata), an important prey for grouse chicks. In eastern Norway, autumn densities of capercaillie and black grouse (Tetrao tetrix) were more likely to peak in vole peak years at high altitudes, where summer temperatures are low. We conclude that high summer temperatures may limit grouse reproduction through the effect on bilberry plants and that a warm climate thus adversely affects population levels of grouse.     

Does time after pair bond disruption affect subsequent reproduction in the socially monogamous woodland vole (Microtus pinetorum)?

Disruption of the pair bond between socially monogamous animals leads to changes in behavior, which may have reproductive consequences. There are two alternative hypotheses to explain the effect of the length of time since pair bond disruption on subsequent reproduction. One hypothesis predicts that voles housed immediately with a new opposite-sex conspecific will be as likely to produce litters and will produce them as quickly as voles separated from their initial mate for longer. Alternatively, if attachment between mates is enduring, we expect that more voles separated longer from their previous mates will produce litters and produce them sooner than voles re-paired immediately after separation from their initial mates. Woodland voles, paired with opposite-sex conspecifics, remained together until parturition. Mates were then separated for 0, 7, or 14 days until re-pairing with an opposite-sex conspecific. Pair bond disruption did not prevent males and females from mating subsequently, which was consistent with data from our breeding colony. In addition, the length of time an individual remained alone after pair bond disruption did not affect the latency to produce a litter. Our results show that having been paired previously does not affect subsequent reproduction in this socially monogamous vole.     

Density-dependent reproduction causes winter crashes in a common vole population

Common voles in western France exhibit three-year population cycles with winter crashes after large outbreaks. During the winter of 2011–2012, we monitored survival, reproduction, recruitment and population growth rate of common voles at different densities (from low to outbreak densities) in natura to better understand density dependence of demographic parameters. Between October and April, the number of animals decreased irrespective of initial density. However, the decline was more pronounced when October density was higher (loss of ≈54 % of individuals at low density and 95 % at high density). Using capture-mark-recapture models with Pradel's temporal symmetry approach, we found a negative effect of density on recruitment and reproduction. In contrast, density had a slightly positive effect on survival indicating that mortality did not drive the steeper declines in animal numbers at high density. We discuss these results in a population cycle framework, and suggest that crashes after outbreaks could reflect negative effects of density dependence on reproduction rather than changes in mortality rates.     

Limits to food intake and fiber utilization in the prairie vole,Microtus ochrogaster: effects of food quality and energy need

We fed prairie voles (Microtus ochrogaster) rat chow diluted with variable amounts of -cellulose to determine 1) how much fiber the voles could tolerate in their diet; 2) changes in food intake and digestibility of dry matter and of fiber; 3) the extent to which voles utilized fiber as an energy source; and 4) whether any of these variables differed between groups of animals maintained at 5 or 22°C. Fiber content of the diets ranged from 20 to 84%. Animals held at 5°C maintained body mass through a diet containing 69% fiber, while animals held at 22°C maintained body mass through the 84% fiber diet. Dry matter intake increased with fiber level from 9.3 to 15.0 g·day^-1 for animals at 5°C and from 5.6 to 14.0 g·day^-1 for animals at 22°C; intake on the highest fiber diet eaten by either group was not different. Dry matter digestibility decreased significantly as the fiber in the diets increased, but was not affected by temperature treatments. Digestible dry matter intake for each group remained constant regardless of diet quality, but on each diet digestible dry matter intake for animals at 5°C was significantly higher than that of the animals held at 22°C. Digestibility of the fiber portion of the experimental diets remained constant as food quality decreased, so the percent of daily energy need met by fiber utilization increased with higher food intake. On the lowest quality diet each group tolerated, fiber digestion provided approximately 42 and 68% of the energy needs of voles at 5 and 22°C, respectively.Abbreviations BM body mass - BMR basal metabolic rate - DE digestible energy - DM dry matter - DMD dry matter digestibility - DDMI digestible dry matter intake - MR metabolic rate - NDF neutral detergent fiber (=cell walls) - NDS neutral detergent solubles (=cell solubles) - SEM standard error of mean - T _a ambient temperature     

Early-summer inhibition of reproduction in wild white-footed mice (Peromyscus leucopus noveboracensis): influence of supplemental food

Adults of wild white-footed mouse (Peromyscus leucopus noveboracensis) populations in southeastern Virginia have been shown to be reproductively suppressed during May–July compared with other months of the reproductive season. The gonads and sex accessory glands are also significantly smaller during May–July than at other times of the year. The causes of this early summer reproductive suppression are unknown. In this study, food (Agway 3000) that could not be hoarded was supplied in feeding stations on one-half the 11-ha wooded study area during 1990 and 1991. Food consumption (g) was positively but not significantly correlated with population density (adults/ha) during both years. Population density was higher during 1991 on the food-supplemented area than on the nonsupplemented area, but the statistical significance was marginal. Mice of both sexes living on both areas during 1990 and 1991 exhibited significant reproductive suppression during May–June. Further, mice of both sexes living on both areas during both years had significantly smaller gonads and sex accessory glands during May–June compared with other months of the year. Thus, during both years, surplus food did not prevent suppression of reproductive function in this wild population, and this phenomenon remains unexplained. Received: November 4, 1998 / Accepted: October 5, 1999     

At the crossroads of physiology and ecology: Food supply and the timing of avian reproduction

The decision of when to breed is crucial to the reproductive success and fitness of seasonally breeding birds. The availability of food for adults prior to breeding has long been thought to play a critical role in timing the initiation of seasonal reproductive events, in particular laying. However, unequivocal evidence for such a role remains limited and the physiological mechanisms by which an increase in food availability results in seasonal activation of the reproductive system are largely speculative. This lack of mechanistic information partly reflects a lack of integration of ecological and physiological approaches to study seasonal reproduction. Indeed, most work pertaining to the role of food availability for adults on the timing of avian reproduction has been ecological and has focused almost exclusively on female traits associated with reproductive timing (e.g., lay date and clutch size). By contrast, most work on the physiological bases of the relationship between food availability and the timing of reproduction has investigated male traits associated with reproductive development (e.g., reproductive hormones and gonadal development). To advance our understanding of these topics, we review the role of proximate factors including food availability, social factors, and ambient temperature in the control of breeding decisions, and discuss the role of three potential candidates (leptin, glucocorticoids, and GnIH–neuropeptide Y) that may mediate the effects of food availability on these decisions. We emphasize that future progress in this area is heavily contingent upon the use of physiology-based approaches and their integration into current ecological frameworks.     

Experimental manipulation of breeding density and litter size: effects on reproductive success in the bank vole

1. Reproductive success of individual females may be determined by density-dependent effects, especially in species where territory provides the resources for a reproducing female and territory size is inversely density-dependent.
2. We manipulated simultaneously the reproductive effort (litter size manipulation: ± 0 and + 2 pups) and breeding density (low and high) of nursing female bank voles Clethrionomys glareolus in outdoor enclosures. We studied whether the reproductive success (number and quality of offspring) of individual females is density-dependent, and whether females can compensate for increased reproductive effort when not limited by saturated breeding density.
3. The females nursing their young in the low density weaned significantly more offspring than females in the high density, independent of litter manipulation.
4. Litter enlargements did not increase the number of weanlings per female, but offspring from enlarged litters had lower weight than control litters.
5. In the reduced density females increased the size of their home range, but litter manipulation had no significant effect on spacing behaviour of females. Increased home range size did not result in heavier weanlings.
6. Mother's failure to successfully wean any offspring was more common in the high density treatment, whereas litter manipulation or mother's weight did not affect weaning success.
7. We conclude that reproductive success of bank vole females is negatively density-dependent in terms of number, but not in the quality of weanlings.
8. The nursing effort of females (i.e. the ability to provide enough food for pups) seems not to be limited by density-dependent factors.     

Day length and estradiol affect same-sex affiliative behavior in the female meadow vole

Non-sexual social bonding between adult mammals is poorly understood, despite its importance in many species. Female meadow voles are territorial and nest alone in long summer day lengths when circulating estradiol concentrations are high, but cohabit in groups in short winter photoperiods when estradiol secretion is low. The influence of day length and estradiol on same-sex huddling behavior was assessed in adult female pairs housed together in long day lengths (LDs) or short day lengths (SDs) from weaning. The behavior of intact, ovariectomized, and estradiol-treated ovariectomized females from each photoperiod was assessed during 3 h partner-preference tests. Intact SD voles, unlike intact LD voles, spent the majority of the test in proximity to their cage-mates. Estradiol treatment of SD voles significantly reduced time spent huddling with the partner. Neither ovariectomy nor estradiol treatment significantly affected the amount of time LD females spent in contact with their partners. Low estradiol availability is therefore a necessary but not sufficient condition for maintenance of high levels of huddling. These results establish that ovarian hormones interact with photoperiod to affect same-sex social behavior.     

Tawny owl prey remains indicate differences in the dynamics of coastal and inland vole populations in southern Finland

Some studies suggest that mild winters decrease overwinter survival of small mammals or coincide with decreased cyclicity in vole numbers, whereas other studies suggest non-significant or positive relationships between mild winter conditions and vole population dynamics. We expect for the number of voles to be higher in the rich and low-lying habitats of the coastal areas than in the less fertile areas inland. We assume that this geographical difference in vole abundances is diminished by mild winters especially in low-lying habitats. We examine these relationships by generalized linear mixed models using prey remains of breeding tawny owls Strix aluco as a proxy for the abundance of voles. The higher number of small voles in the coastal area than in the inland area suggest that vole populations were denser in the coastal area. Vole populations of both areas were affected by winter weather conditions particularly in March, but these relationships differed between areas. The mild ends of winter with frequent fluctuations of the ambient temperature around the freezing point (“frost seesaw”) constrained significantly the coastal vole populations, while deep snow cover, in general after hard winters, was followed by significantly lowered number of voles only in the inland populations. Our results suggest that coastal vole populations are more vulnerable to mild winters than inland ones. We also show that tawny owl prey remains can be used in a meaningful way to study vole population dynamics.     

Density-dependent growth and reproduction of the apple snail, Pomacea canaliculata: a density manipulation experiment in a paddy field

To examine density dependence in the survival, growth, and reproduction of Pomacea canaliculata, we conducted an experiment in which snail densities were manipulated in a paddy field. We released paint-marked snails of 15–20 mm shell height into 12 enclosures (pens) of 16 m² at one of five densities – 8, 16, 32, 64, or 128 snails per pen. The survival rate of released snails was 95% and was independent of snail density. The snail density had a significant effect on the growth and egg production of individual snails. This density dependence may have been caused by reduced food availability. The females at high density deposited fewer and smaller egg masses than those at low density, and consequently produced fewer eggs. The females at densities 8 and 16 deposited more than 3000 eggs per female, while the females at density 128 oviposited only 414 eggs. The total egg production per pen was, however, higher at higher snail density. The survival rates of juvenile snails were 21%–37% and were independent of adult density. The juvenile density was positively correlated with the total egg production per pen and hence was higher at higher adult density. However, the density of juveniles larger than 5 mm in shell height, i.e., juveniles that can survive an overwintering period, was not significantly different among density treatments. These results suggest that snail density after the overwintering period is independent of the density in the previous year. Thus, density dependence in growth and reproduction might regulate the population of P. canaliculata in paddies. Received: October 23, 1998 / Accepted: July 16, 1999     

An approach to the edge effect in proof of the validity ofDice's assessment lines in small-mammal censusing

A second fieldwork for the sake of solving the edge-effect subject was carried out in almost the same ways and plans in summer of 1971 in succession to that of 1970. It was found out that the population density was reduced to two-thirds, but the data were sufficient to form the subsequent conclusions:

1. The initial population estimated by the removal census method turned out to be well compatible with the numbers of survivals and emigrants of marked voles calculated byJolly 's method.
2. The natural, entire home ranges of both sexes reached by determining observed range length and width with 24 voles, selected on rigid conditions, agreed with those of the range-conservative type in 1970 in both acreage and elliptic shape.
3. In the majority of the select specimens, considered to reveal the natural ranges, the random distribution in number of captures per station within range was proved by means of I_σ so far as such capture density as treated here is concerned. On the ground of the empirical evidence, the above range size approached by the routine observational method could be made surprisingly accordant with the range size in terms of r-values calculated byWierzbowska 's method.
4. From these proofs, the validity ofDice 's assessment lines makes evident, and besides, it could be further substantiated by use ofMarten 's notion of mouse-equivalents through the mediation ofWierzbowska 's method.
5. Consequently, I have largely come to the conclusion thatDice 's assessment lines stand on justifiable basis from theoretical and empirical angles.