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.     

An estimate of the impact of predators on the British Field Vole Microtus agrestis population

The productivity of the British Field Vole population is estimated to be between 677 000 and 982 000 voles per year, depending on the precise basis of the calculation. The total consumption of Field Voles by all the major predators, mammalian and avian, is estimated to be around 980 000, if the very uncertain consumption by Feral Cats is included. Two specialist vole predators (Weasel, Kestrel) and two generalist predators (Red Fox, Feral Cat) consume 85% of the total between them, the other 10 species share the rest between them. However important voles might be for owls, owl predation is unimportant to voles. The slight evidence for cyclicity in Field Vole populations in Britain might be explained by the overall take of voles by the predators, and by the rather generalized diet of even the vole specialists.     

The diet of migrant Red Kites Milvus milvus during a Water Vole Arvicola terrestris outbreak in eastern France and the associated risk of secondary poisoning by the rodenticide bromadiolone

Poisoning by pesticides is considered one of the primary threats to the Red Kite Milvus milvus. We studied the diet of this raptor in an area of eastern France where the rodenticide bromadiolone is widely used to control Water Vole Arvicola terrestris outbreaks. A high degree of specialization for Water Voles was noted, as their remains were identified in all 119 pellets collected in autumn 2008, whereas other small rodent species and insects occurred in 27 and 9% of pellets, respectively. We estimated that Water Voles constituted 94% of the total biomass ingested by Red Kites under these conditions. Based on these data, the risk of secondary poisoning due to feeding on poisoned voles was assessed. Acute exposure on a single day was not considered a risk for Kites, but exposure to poisoned voles over 1 week represented the maximal risk for the Red Kite; the calculated dose of bromadiolone ingested by a Red Kite was 137 times higher than the toxicological benchmark for birds. A field survey in the studied area detected four dead Red Kites and one moribund bird in autumn 2008 but did not confirm that the cause of death was bromadiolone poisoning. We suggest that professional monitoring is needed to assess the impact of rodenticides on Red Kites in areas where voles are controlled.     

The prey of the Barn owl (Tyto alba alba) in East Norfolk

Some 5000 Barn owl pellets, collected from sites in East Norfolk during the past decade, have been examined. The most important prey species, by weight, were the Field vole (Microtus agrestis) 52%, the Brown rat (Rattus norvegicus) 17%, and the Common shrew (Sorex araneus) 12%. The prey varies over different habitats; Wood mice (Apodemus sylvaticus) and Bank vole (Clethrionomys glareolus) forming a higher proportion in localities with hedges, scrub and woodland than in open grasslands. These results are comparable with those of other recent work. However, when compared with studies conducted over 30 years ago, it would appear that the Field vole now constitutes a higher proportion, and the Brown rat a lower proportion, of the prey taken.     

Bird communities of highway verges: Influence of adjacent habitat and roadside management

We have investigated the effects of landscape traversed and roadside structure on the use of highway verges by birds. Three contrasted landscapes were chosen in terms of human land use and vegetation structure: an intensive farmland, a pine plantation, and a matoral. The roadside sections varied in vegetation structure, width and profile. We recorded birds present in roadsides and adjacent habitats by transect counts over all seasons. Roadside bird species appeared for a great part similar to those of adjacent habitats. However, diversity and abundance in verges did not depend on that of adjacent habitats. Woody roadsides were comparable to hedges, as trees (and shrubs) in verges enhanced species richness and abundance of birds in the farmland and woodland sites. Width and profile of verges had less influence. In all sites, typical species of the habitat traversed partly avoided roadsides. On the contrary, numerous species associated with ‘rare’ habitats in one site preferred roadsides, provided that verge vegetation contrasted with the dominant habitat. It is concluded that birds responses to highways can vary greatly with landscape traversed and verge vegetation. Highway verges could be favorable to birds, if they constitute a complementary habitat to the dominant habitat within a landscape.     

Rainfall, ambient temperature, and Clethrionomys californicus capture frequency

Sixty Western Red-backed Voles Clethrionomys californicus inhabiting an old-growth mixed conifer forest in north-western California (USA) were studied over 2 years to assess relationships among vole capture frequency, rainfall, and ambient temperature. Red-backed Voles were encountered more often and recapture rates per vole doubled with the tripling of yearly rainfall. In addition, more voles were captured during wet months than dry months. Analysed over weekly intervals, captures were positively correlated with the amount of weekly rainfall, a greater proportion of the known vole population was encountered during wet weeks than dry weeks, and increased weekly rainfall resulted in an increased proportion of the vole population captured. Significant relationships were not found between daily rainfall and vole captures, but both minimum and maximum daily ambient temperatures were lower when voles were trapped. Vole captures increased with decreases in daily and weekly minimum and maximum ambient temperatures. Weekly rainfall in combination with minimum weekly temperatures accounted for the greatest explanation of variability in vole capture frequencies. These results document that rainfall and ambient temperature have an impact on Western Red-backed Vole capture frequency; thus these weather variables should be addressed as covariates in analyses focused on capture data for this species.     

Diet selection and density estimates of forest buffalo in Campo‐Ma’an National Park,Cameroon

We studied diet selection and density of forest buffalo in the Campo Ma’an National Park of southern Cameroon. The buffalo’s diet in this rainforest comprised 43% grass, including 15%Leptochloa caerulecens. Other species eaten were non‐graminoid monocots (21.3%), mainly Commelinaceae (18.2%), including Palisota spp. and dicotyledons (32.7%), mainly leaves (26.5%). This diet revealed that buffalo collect their food on road verges, logging tracks and along large rivers. This agrees well with the distribution of buffalo tracks in the study area. Mean buffalo density in the forest was only 0.01 buffalo km^?2, whereas the density on the road together with its direct surroundings was 0.4 buffalo per km of road. We estimated the total number of buffalo in the 650 km² southern part of National Park Campo‐Ma’an of only twenty individuals. We suggest that the buffalo’s ancestral niche was not a primeval type of rainforest, as suggested by Kingdon, but an interface between savanna and rainforest. The Campo‐Ma’an buffalo most likely depend to a very large extent on anthropogenic vegetation types. Maintaining some grass cover by annual clear‐cutting along roads and on logging tracks might be necessary for the survival of the buffalo population within the National Park.     

Nest loss in capercaillie and black grouse in relation to the small rodent cycle in southeast Norway

Summary The relationship between nest loss in boreal forest grouse and the fluctuations in small rodents was studied at Varaldskogen in southeast Norway during 1979–1986, covering two complete rodent cycles. Nest loss in capercaillie (N=174) and black grouse (N=81) was calculated according to Mayfield (1975) based on nests from radio-equipped hens (N=77) and nests found by other methods (N=178). Small rodent density was measured by snap trapping during spring and autumn. Losses varied as predicted by the classical alternative prey hypothesis (Hagen 1952 and Lack 1954, as elaborated by Angelstam et al. 1984): high losses during rodent crash years (85.5% capercaillie, 51% black grouse), and smaller losses during peak years (54.5% capercaillie, 32.5% black grouse). Losses were inversely related to autumn abundance of rodents in capercaillie (P<0.05), but the correlation was not significant for black grouse (0.10<P<0.20). In capercaillie, the only species with an adequate sample for analysis, no relationship was detected between spring density of rodents and nest loss. Losses during the prepeak years were nearly as high as during crash years, a result inconsistent with the model. We conclude that the numerical response of predators to their cyclic main prey (i.e. small rodents) probably play a main role during the low phase and prepeak year, whereas the dietary shift is most important during the peak and crash year of the cycle.     

Effects of predator removal on vertebrate prey populations: birds of prey and small mammals

We studied the effects of removal of breeding nomadic avian predators (the kestrel, Falco tinnunculus and Tengmalm's owl, Aegolius funereus) on small mammals (voles of the genera Microtus and Clethrionomys and the common shrew, Sorex araneus) during 1989–1992 in western Finland to find out if these predators have a regulating or limiting impact on their prey populations. We removed potential breeding sites of raptors from five manipulation areas (c. 3 km² each), whereas control areas had nest-boxes in addition to natural cavities and stick-nests. Densities of small mammals were monitored by snap-trapping in April, June, and August, and densities of mammalian predators (the least weasel, Mustela nivalis nivalis, the stoat, M. erminea and the red fox, Vulpes vulpes) by snow tracking in early spring and late autumn. The yearly mean number of raptor breeding territories was 0.2–1.0 in reduction areas and 3.0–8.2 in control areas. Breeding raptors alone did not regulate prey populations in the long term, but probably caused short-term changes in the population dynamics of both the main prey, the sibling vole (Microtus rossiaemeridionalis) and an alternative prey (the common shrew). The densities of an alternative prey, the bank vole (Clethrionomys glareolus) decreased in raptor reduction areas, most likely due to increased least weasel predation pressure in the absence of breeding avian predators.     

Microhabitat use, giving-up densities and temporal activity as short- and long-term anti-predator behaviors in common voles

We used depletable food patches to determine the effect of microhabitat (mowed versus unmowed adjacent grasslands) and time (day versus night) on the foraging behavior of common voles (Microtus arvalis). The food remaining after 12‐h periods (giving‐up density, GUD) measured the vole's habitat selection under predation risk. In accord with several other rodent species and the effects of avian predators, voles had significantly lower GUDs in the unmowed than mowed portion of the grassland. GUDs in patches along the border between adjacent habitats were more similar to the risky mowed grassland than the safe unmowed grass. Time interacted strongly with microhabitat. In the mowed grass, voles had significantly higher GUDs at night than day. Whereas in the unmowed grass, GUDs were significantly higher during the day than night. Vole GUDs did not vary with time along the boundary. This suggests that predators are more abundant or effective in the mowed grass at night (owls?), and in the unmowed grass during the day (weasels?). In terms of predation risk, the voles perceived the mowed grass at night as the riskiest and the unmowed grass at night as the safest. Voles may have difficulties assessing resources under high predation risk: GUDs among patches were well equalized in the unmowed microhabitat whereas in the mowed grass only day GUDs did not vary significantly among patches. We linked these results to the vole's day‐night‐activity and life span. For the 533 voles live‐trapped at the study area, the ratio of day versus night captures for each individual served as an activity index and the span between first and last capture measured minimum life span. In accord with higher GUDs at night, very few individuals behaved selectively towards the night, but individual life expectancy increased with temporal opportunism. Microhabitat differences in GUDs reflect short‐term strategies of predator avoidance and the trapping data reflect long‐term patterns of anti‐predator behavior.     

Effects of narrow roads on the movement of carabid beetles (Coleoptera,Carabidae) in Nopporo Forest Park,Hokkaido

Forest-dwelling carabid beetles that have no flight ability were studied using mark-recapture methods in late-June to mid-October 2007. This study was done to determine the effects of narrow roads in Nopporo Forest Park, Hokkaido on carabid beetle movement and habitat use. The investigation was conducted at four sites: one site was an abandoned grassy road with a width of 3.5 m, two sites were gravel roads with widths of 3.5 and 4.5 m, and another site was an asphalt-paved road with a width of 4.5 m. A total of 3,580 individuals from six species of carabid beetles were collected using dry pit-fall traps, and recapture rates ranged from 6.1 to 36.2%. All examined roads acted as barriers against the movement of Leptocarabus arboreus ishikarinus. All roads, except the abandoned grassy road, acted as a barrier against Carabus granulatus yezoensis movement. Forest–roadside verge comparisons demonstrated that some carabid beetles avoid even narrow roadside verges. Harmful effects increase with increasing road width and both paved roads and narrow roads negatively affect the movement of carabid beetles inhabiting the bordering forest. Therefore, forest specialist beetles are influenced by a barrier effect that starts at the forest road verge, and this barrier effect may be exacerbated by vehicular traffic. Therefore, these barrier effects on carabid beetles should be considered when planning and implementing road construction and maintenance in forests.     

Infrastructure features outperform environmental variables explaining rabbit abundance around motorways

Human disturbance is widespread across landscapes in the form of roads that alter wildlife populations. Knowing which road features are responsible for the species response and their relevance in comparison with environmental variables will provide useful information for effective conservation measures. We sampled relative abundance of European rabbits, a very widespread species, in motorway verges at regional scale, in an area with large variability in environmental and infrastructure conditions. Environmental variables included vegetation structure, plant productivity, distance to water sources, and altitude. Infrastructure characteristics were the type of vegetation in verges, verge width, traffic volume, and the presence of embankments. We performed a variance partitioning analysis to determine the relative importance of two sets of variables on rabbit abundance. Additionally, we identified the most important variables and their effects model averaging after model selection by AICc on hypothesis‐based models. As a group, infrastructure features explained four times more variability in rabbit abundance than environmental variables, being the effects of the former critical in motorway stretches located in altered landscapes with no available habitat for rabbits, such as agricultural fields. Model selection and Akaike weights showed that verge width and traffic volume are the most important variables explaining rabbit abundance index, with positive and negative effects, respectively. In the light of these results, the response of species to the infrastructure can be modulated through the modification of motorway features, being some of them manageable in the design phase. The identification of such features leads to suggestions for improvement through low‐cost corrective measures and conservation plans. As a general indication, keeping motorway verges less than 10 m wide will prevent high densities of rabbits and avoid the unwanted effects that rabbit populations can generate in some areas.     

The effect of experimental removal of red foxesVulpes vulpes on spring density of brown haresLepus europaeus in western Poland

Red foxesVulpes vulpes (Linnaeus, 1758) were experimentally removed in two nearby areas located in western Poland to verify the hypothesis about the limiting impact of their predation on the low-density population of brown haresLepus europaeus (Pallas, 1778) (4.4-10.6 ind./km² in late autumn). In 1996/1997–2001/2002 foxes were culled (mainly in autumn and winter) in the reduction area (32 km²), whereas in the control area (34 km²) intensive culling was carried out only in 2000/2001–2001/2002. Indices of fox and hare spring densities were estimated using spotlight counts, as mean numbers of individuals observed per 10 km of the counting route. Annual changes in the fox density indices were negatively correlated with the bag of foxes, and annual changes in the hare density indices were negatively related to the annual changes in fox density indices. The fox density indices were significantly lower in the reduction area than in the control one only in 2000–2001 (2.8 times, on average), and in the same years, the hare population responded with higher density (1.7 times, on average). The hare responses took place without time delay, which suggests that the changes in fox abundance affected the situation of hares primarily in the autumn-winter season.     

The role of food supply in the dispersal behaviour of juvenile Tawny Owls Strix aluco

We investigated the effects of food supply on decisions made by dispersing juvenile Tawny Owls Strix aluco in Kielder Forest, Northumberland, in 1996 and 1997. Field Voles Microtus agrestis were the main food of the owls and clear-cuts the main habitat for voles. A vole sign index was used to estimate vole abundance. In areas near to roosting owls, mean vole densities were 83 and 115 ha⁻¹ in 1996 and 1997, respectively. The prediction that birds would perform area-restricted searches when prey was more abundant was not confirmed. Moreover, we found no evidence that juveniles avoided conspecifics. Owls appeared to have an imperfect knowledge of the environment as they responded to variability in Field Vole densities by altering the time spent in different areas rather than by moving to areas with successively greater vole densities. Vole abundance explained 25.7% of the variation in the time spent in different areas. Movements did not decrease with time after dispersal, although the detection of such movements was prone to error. This study supports recent work suggesting that although dispersal may be initiated by a variety of proximate and ultimate factors, individual decisions made during dispersal may depend partly on environmental conditions encountered during the process itself.     

The spread of the Bank vole Clethrionomys glareolus in Ireland

The discovery of the Bank vole Clethrionomys glareolus in Ireland and the published information on its spread are reviewed, and the results of a complete resurvey of its distribution, in 1 982 , presented. The rate of spread is extremely variable and somewhat unpredictable. Over suitable habitat—agricultural land with hedges and other cover (and sometimes mountain valleys)—spread may average 2–4.5 kmyr^-1 but can sometimes be very much slower. This might be influenced by the type and extent of hedges and of permanent pasture. High ground and moorland usually prevent spread but there have been dramatic exceptions. In 1982 Bank voles occupied probably the whole of Co. Limerick; most of Co. Kerry, with the exception of substantial parts of the mountainous peninsulas in the southwest and contiguous interior; a large part of Co. Cork, though not the south-east and again excepting portions of the mountainous south-west, together with a substantial strip of east Co. Tipperary, and the south-east quarter of Co. Clare.     

Responses of stoats and least weasels to fluctuating food abundances: is the low phase of the vole cycle due to mustelid predation?

Summary We studied responses of stoats and least weasels to fluctuating vole abundances during seven winters in western Finland. Density indices of mustelids were derived from snow-tracking, diet composition from scat samples, and vole abundances from snap-trapping. Predation rate was estimated by the ratio of voles to mustelids and by the vole kill rate by predators (density of predator x percentage of voles in the diet). We tested the following four predictions of the hypothesis that small mustelids cause the low phase of the microtine cycle. (1) The densities of predators should lag well behind the prey abundances, as time lags tend to have destabilizing effects. The densities of stoats fluctuated in accordance with the vole abundances, whereas the spring densities of least weasels tracked the vole abundances with a half-year lag and the autumn densities with a 1-year lag. (2) Predators should not shift to alternative prey with declining vole densities. The yearly proportion of Microtus voles (the staple prey) in the diet of stoats varied widely (range 16–82%) and was positively correlated with the winter abundance of these voles. In contrast, the same proportion in the food of least weasels was independent of the vole abundance. (3) The ratio of voles to small mustelids should be smallest in poor vole years and largest in good ones. This was also observed. (4) Vole densities from autumn to spring should decrease more in those winters when vole kill rates are high than when they are low. The data on least weasels agreed with this prediction. Our results from least weasels were consistent with the predictions of the hypothesis, but stoats behaved like semi-generalist predators. Accordingly, declines and lows in the microtine cycle may be due to least weasel predation, but other extrinsic factors may also contribute to crashes.     

Density,diet and productivity of Long-eared Owls Asio otus in the Italian Alps: the importance of Microtus voles

Capsule Relatively large populations, feeding predominantly upon voles, were present at higher elevations.

Aims To determine the density, productivity and diet composition of Long-eared Owls breeding at higher elevations.

Methods Population census and breeding biology were investigated, and dietary analysis performed for 32 Long-eared Owl territories that were occupied over a six-year period (2000–05), in a 155 km² study area located in the Noce Valley of the central-eastern Italian Alps.

Results Territories were mainly situated at the edge of large pine forests, near large patches of grassland and/or open-structured apple orchards, at elevations ranging between 540 and 1210 m. Density varied between 10 and 15 pairs/100 km². Mean intraspecific nest spacing averaged 1727 m and territories were either solitary or clumped in loose aggregations of one to five pairs. Mean laying date was 27 March and the mean number of fledged young was 0.95 and 2.13 per territorial and successful pair, respectively. Adults and nestlings were preyed upon by Eagle Owls Bubo bubo and Common Buzzards Buteo buteo, both of them abundant in the study area. Diet was dominated by Common Voles Microtus arvalis, complemented by Apodemus mice and thrushes. Annual variations in owl density, productivity and diet breadth varied in parallel with the occurrence of Common Voles in the diet, suggesting that the owls responded numerically to the availability of their main prey. This confirms earlier analyses on other European populations, but without the marked three- to four-year cycles observed in such areas.

Conclusion The conservation status of the species in these Alpine habitats seemed currently satisfactory. However, the unregulated use of rodenticides in apple orchards and the loss of open habitats associated with land abandonment may represent long-term threats for the species in these habitats.     

内蒙古草原布氏田鼠和长爪沙鼠体重及身体成分的季节变化

研究了内蒙古草原布氏田鼠(Lasiopodomys brandtii)和长爪沙鼠(Meriones unguiculatus)的体重、身体脂肪、水分含量及身体热值等指标的季节变化.2种动物的体重都是在春季最高,布氏田鼠的体重具显著的季节变化,长爪沙鼠的鲜重具季节变化,但干重则基本维持恒定.2种动物的脂肪含量和身体热值都...     

Changes in the food of British Barn Owls (Tyto alba) between 1974 and 1997

Comparison of the results of a 1993–97 Barn Owl Tyto alba pellet survey with those of a similar survey from 1956–74 showed that Barn Owl diet had changed significantly. The primary differences were a widespread decrease in the percentage of Common Shrew Sorex araneus, combined with an increase in Pygmy Shrew Sorex minutus. The percentage of Wood and Yellow‐necked mice Apodemus sylvaticus and A. flavicollis and Bank Vole Clethrionomys glareolus in the diet also increased. Changes in Barn Owl diet since 1974 were independent of land‐class group, but were dependent upon region. This was due primarily to a large increase in the percentage of Apodemus spp. in Eastern England. Whilst the percentage of Pygmy Shrew in Barn Owl diet showed significant regional variation, there was no significant variation between land‐class groups. The diversity of Barn Owl diet increased between 1974 and 1997, although it was still lower in 1997 than earlier in the century. This increase was dependent upon region, but independent of land‐class group. The combined results of both surveys showed significant interland‐class group variation in dietary diversity. Changes in diet are discussed in relation to the intensification of agriculture and other changes in land management since the 1970s. The effects on Barn Owls of these changes in prey abundance are discussed, particularly in relation to the decline in Barn Owl numbers during the twentieth century.     

The habitat use and diet of Black Grouse Tetrao tetrix in the Pennine hills of northern England

The habitat and dietary preferences of Black Grouse were studied from 1989 to 1991 in a largely treeless habitat within a 17 km² area of the Pennine hills of northern England using radiotelemetry. Grassland habitats were preferred by 84% of 19 birds for much of the year and heather moorland was selected in winter; the other birds, all females, remained in heather habitats all year. Most females reared broods in damp, rushy grass moorlands, but one of six broods used hay fields. Diet closely reflected seasonal plant availability in selected habitats and the leaves, flowers, fruits and seeds of a wide variety of plants were eaten. Cotton-grass Eriophorum vaginatum was important in spring, grassland herbs, grasses, sedges and rushes in summer and autumn and Ling Heather Calluna vulgaris in late autumn and winter. Males fed substantially more on heather in winter than females which partially replaced heather in their diet with the leaves of grasses and herbs. Young chicks showed considerable preference for Sawfly (Symphyta) larvae. Suggested management recommendations include the regulation of sheep-grazing, encouragement of wet flush areas and the late cutting of hay fields.