Population growth and habitat distribution in cyclic small rodents: to expand or to change?

Habitat selection determined by intraspecific interactions (social behaviour), being either free or despotic, should result in the largest densities in the most favourable habitat at least in late increase and decline phases of cyclic populations. Habitat distribution determined by interspecific effects such as herbivore-plant or predator-prey interactions may result in higher densities in inferior habitats at late peaks and/or declines due to overgrazing of preferred habitats, or invasion of such habitats by specialist predators. An examination of the distributions of the rodent species Clethrionomys glareolus and Microtus agrestis during a population cycle on forest clearcuts at Grimsö, south-central Sweden, demonstrated clear changes towards less preferred habitats during the decline phase, particularly in M. agrestis. Intraspecific competition could be excluded as a cause since the numbers of both species declined simultaneously. There was no sign of overgrazing, while weasels, Mustela nivalis, invaded the preferred M. agrestis habitat during peak–decline. Predation thus appears more important for the local occurrence at early decline, rather than territoriality or other social behaviour. It is concluded that distributions in various habitats do not always reveal any primary habitat selection but, rather, habitat-dependent survival.     

The effect of diet quality on gut anatomy in British voles (Microtinae)

Three species of British voles, the bank vole Clethrionomys glareolus, the field vole Microtus agrestis and the water vole Arvicola terrestris were maintained on diets of seed and plant leaf material to investigate changes in gut anatomy. C. glareolus and M. agrestis showed significant changes in most regions of the gut; they developed longer and heavier tracts when on a highfibre diet. This response may be important in enabling these animals to withstand seasonal changes in diet quality.     

Spatial dynamics in fluctuating vole populations

Summary Patterns and consistency of distribution, spatial and temporal components, and the extent of spatial density-dependence were compared between semi-cyclic and cyclic populations of the vole species Clethrionomys glareolus and Microtus agrestis in south-central and north Sweden. Cyclic populations were less clumped and only C. glareolus showed a consistency in distribution between years. Spatial variation contributed little to the distributions in cyclic populations while the spatial and temporal variations were of the same magnitude in the semi-cyclic populations. The latter populations could be subdivided into areas with different spatial and temporal components. The spatial density-dependence increased from increase to decline years in C. glareolus but not in M. agrestis, which differed conspicuously between reregions in population development. The data imply that spatial dynamics should be considered as much as temporal ones for non-cyclic populations, that the same regulating or limiting factors may be at work in both spatial and temporal components and that, in addition, social behaviour may be important in explaining spatial dynamics. However, the latter effects may be fairly species-specific.     

Species diversity and biocenotic relationships of gamasid mites (Parasitiformes,Gamasoidea) in the nests of the field vole <Emphasis Type="Italic">Microtus agrestis</Emphasis> L. of the Karelian middle taiga subzone

The species diversity and habitat relations of gamasid mites from nests of the field vole (Microtus agrestis L.) were studied in the middle taiga subzone of the Republic of Karelia. The fauna of gamasid mites on the field vole, in its nests, and in the forest litter is very diverse, comprising 49 species with a complex taxonomic structure (22 genera, 12 families) and a variety of life strategies (necrophages, zoophages, and hematophages). The fauna of gamasid mites is most diverse in the nests of M. agrestis and includes both primitive free-living forms and specialized obligatory (epizoic) parasites. The group with the highest diversity of mite species is that from the nest of M. agrestis, with the diversity index amounting to 2.65 (30 species, 18 genera, 11 families).     

Influence of within‐patch habitat quality on high‐Andean Polylepis bird abundance

Habitat restoration strategies for fragmented high Andean forest landscapes must consider the influence of within‐patch habitat quality on bird abundance. I examined vegetation and bird abundance at three locations within a highly fragmented Polylepis forest landscape in the Cordillera Vilcanota, southern Peru. Across the landscape, there was significant variation in the vegetation structure of Polylepis forest patches of different size categories, especially in terms of tree girth, tree height, tree density, and canopy vegetation structure. Principal Component Analysis extracted five factors of habitat quality, which together accounted for 74.2% of the variability within 15 habitat variables. Polylepis bird species differed in their responses to habitat quality but, overall, variation in Polylepis bird abundance was not fully captured by the range of habitat quality variables. Tall, dense vegetation cover was clearly important for 11 conservation‐important species, a high density of large trees was important for 10 species and primary forest ground cover was important for eight species. Habitat quality exhibited no significant influence on the abundance of only one species –Asthenes urubambensis. The abundance of seven species was associated with lower elevation forest, but only one species was associated with higher elevation forest. Management of habitat quality in large and medium remnant forest patches throughout the Cordillera Vilcanota, particularly in the 3800–4200 m elevation range, will be a cornerstone in ensuring the persistence of the majority of conservation‐important bird species populations.     

Small mammal responses to moose supplementary winter feeding

Supplementary feeding of wild large herbivores is a widespread practice in North America and Europe. The presence of feeding stations may have ecological consequences through changes to animal distributions, patterns of herbivory and a net nutrient input into the ecosystem. In Fennoscandia, supplementary feeding of moose in winter (Alces alces) is increasing. Although it has been shown to affect bird communities, its effects on small mammal communities were unknown. Here, we studied the effects of moose supplementary feeding stations on plants and on abundance, reproduction, and biomass of small mammals in years with low and high vole abundance. We sampled small mammals with snap traps and conducted surveys of the field layer vegetation, at varying distances from moose supplemental feeding stations. Due to the vegetation changes induced by feeding stations, abundance of common shrews (Sorex araneus) and Microtus voles were positively affected by long-term moose winter feeding, while bank voles (Myodes glareolus) were not affected. Moose feeding stations did not affect reproduction, individual body mass, or the total biomass of small mammals. Moose winter-feeding stations have impacts on nontarget species, providing islands of preferred grass and forb habitat for Microtus spp. and common shrews, allowing them to penetrate into a matrix of less preferred forest habitat.     

Individual flexibility in habitat selection in the ortolan bunting Emberiza hortulana

In order to understand habitat selection in birds, it is important to know how individuals respond when encountering a variety of habitats during dispersal and must choose between them. However, very few field studies have addressed this question. We compared habitat selection of ortolan buntings Emberiza hortulana before and after dispersal events. In Norway, this species has a patchy distribution and breeds in different open habitats of which the two major ones are botanically distinct: raised peat bogs and forest clear‐cuts on dry, sandy soil. There was no evidence that habitat selection of males after natal dispersal was influenced by natal habitat, and 83% changed vegetation type. Habitat selection was random when taking the availability of each vegetation type into account, both at the landscape level and among habitats encountered along likely dispersal routes. Habitat selection after breeding dispersal was not influenced by the habitat of the patch of origin, and 62–71% changed habitat during successive dispersal events. Changing habitat did not seem to affect pairing or breeding success, but decreased with age. Lifetime patterns of habitat selection indicated that the majority (86%) changed habitat one or more times. However, the proportion never changing habitat (14%) was significantly higher than expected if settlement after dispersal was random (5%), suggesting that the majority of males were flexible whereas a minority was habitat conservative. These results provide some of the first systematic evidence based on settlement decisions of individuals that habitat selection of birds can be flexible with regard to vegetation type.     

Habitat preferences of the bank voleMyodes glareolus in a Mediterranean mountain range

The bank voleMyodes glareolus Schreber, 1780 is a widely distributed rodent in Europe, being numerically dominant in small mammal communities living in temperate woodlands. However, it becomes scarce in southern Europe (Mediterranean area) where it reaches the southernmost limit of its distribution range. We studied the habitat preferences of bank voles in 9 plots in a transitional area between Mediterranean and Eurosiberian regions within a Mediterranean mountain. During the study period we captured 1919 small mammals of 9 species, including 287 bank voles (14.95%). Mean density ranged from zero individuals per plot (1.1 ha) at the boreo-subalpine scrubland to 10.27 ± 2.84 (SE) at a Mediterranean river woodland. Statistical path analysis was used to investigate relationships between mean bank vole density and climate and vegetation structure measured within plots. The variables selected by the structural equation model were those related to forest structure, like tree cover and height, dead vegetation, moss, and rock cover. Habitat moisture was also important (microclimatic conditions). Mean climate conditions (and elevation) did not have any significant effect on mean bank vole density, and no significant association with understorey vegetation (eg shrub and herbaceous cover) was observed. Our results pointed out that bank voles were habitat specialists in our study area, being more abundant and frequent in moist woodlands, and rare or absent in shrublands and grasslands.     

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.     

A Hierarchical Analysis of Habitat Selection by Raccoons in Northern Indiana

Abstract: Although numerous studies have examined habitat use by raccoons (Procyon lotor), information regarding seasonal habitat selection related to resource availability in agricultural landscapes is lacking for this species. Additionally, few studies using radiotelemetry have investigated habitat selection at multiple spatial scales or core-use areas by raccoons. We examined seasonal habitat selection of 55 (31 M, 24 F) adult raccoons at 3 hierarchical orders defined by the movement behavior of this species (second-order home range, second-order core-use area, and third-order home range) in northern Indiana, USA, from May 2003 to June 2005. Using compositional analysis, we assessed whether habitat selection differed from random and ranked habitat types in order of selection during the crop growing period (season 1) and corn maturation period (season 2), which represented substantial shifts in resource availability to raccoons. Habitat rankings differed across hierarchical orders, between seasons within hierarchical orders, and between sexes within seasons; however, seasonal and intersexual patterns of habitat selection were not consistent across hierarchical orders of spatial scale. When nonrandom utilization was detected, both sexes consistently selected forest cover over other available habitats. Seasonal differences in habitat selection were most evident at the core-area scale, where raccoon selection of agricultural lands was highest during the maturation season when corn was available as a direct food source. Habitat use did not differ from availability for either sex in either season at the third-order scale. The selection of forest cover across both seasons and all spatial orders suggested that raccoon distribution and abundance in fragmented landscapes is likely dependent on the availability and distribution of forest cover, or habitats associated with forest (i.e., water), within the landscape. The lack of consistency in habitat selection across hierarchical scales further exemplifies the need to examine multiple biological scales in habitat-selection studies.     

Local coexistence and niche differences between the Lusitanian and Mediterranean pine voles (<Emphasis Type="Italic">Microtus lusitanicus</Emphasis> and <Emphasis Type="Italic">M. duodecimcostatus</Emphasis>)

In the present study, we analyzed the coexistence pattern of the Lusitanian pine vole (Microtus lusitanicus) and the Mediterranean pine vole (Microtus duodecimcostatus) in a potential area of sympatry in a Mediterranean landscape (Portugal). We also determined the relative contribution of local, landscape, and spatial factors explaining the differences in the distribution patterns of the two species in the region. Using a kriging interpolation method, we obtained a map of sympatric and allopatric areas of species occurrence. The estimated sympatry area corresponded to a northwest–southeast belt representing 11.3% of the study area. Habitat niche differences were assessed with binomial GLMs followed by a variance partitioning. At a local scale, higher altitude, higher cover of shrubs, lower clay content in the soil, and lower cover of tree canopy were the most important factors distinguishing M. lusitanicus presence sites from those with M. duodecimcostatus. At a larger scale, the presence of forest landscape units and the low abundance of “montado” units were the most influencing landscape factors in the identification of M. lusitanicus occurrence sites when compared to M. duodecimcostatus. Our results suggested that local coexistence of M. lusitanicus and M. duodecimcostatus in the field is a rare event. The differences in distribution patterns of the two pine vole species were mostly explained by fine-scale environmental factors and by shared spatial effects.     

Will Environmental Stewardship enhance small mammal abundance on intensively managed farmland?

• 1 The creation of grassy field margins, as part of the UK Government's Environmental Stewardship (ES) scheme, is one of a number of measures proposed to mitigate the adverse effects of arable intensification on wildlife. Widespread development of these margins will potentially increase the amount of habitat available to small mammals in arable landscapes, as many species do not inhabit the cropped area.
• 2 The aim of this study was to determine what impact ES-type margins might have on small mammals. We compared small mammal abundance and biomass in spring and autumn on 3-m-wide and 6-m-wide grassy margins with that on conventionally managed field edges that have no margin (0 m wide) and are intensively cultivated to the field edge.
• 3 Bank voles Clethrionomys glareolus, wood mice Apodemus sylvaticus and common shrews Sorex araneus were the most abundant species; few field voles Microtus agrestis were captured on any margin. In autumn, bank vole and common shrew numbers were higher on the grassy margins than on conventional field edges, and margin width per se was positively associated with bank vole abundance. The number of wood mice captured did not differ among the different margin types.
• 4 Total small mammal biomass increased between spring and autumn on the 3-m- and 6-m-wide grassy margins, but decreased on the 0-m-wide margins. Total biomass in autumn was three times higher on 6-m-wide margins compared with the conventional arable field edges.

     

An appraisal of the fitness consequences of forest disturbance for wildlife using habitat selection theory

Isodar theory can help to unveil the fitness consequences of habitat disturbance for wildlife through an evaluation of adaptive habitat selection using patterns of animal abundance in adjacent habitats. By incorporating measures of disturbance intensity or variations in resource availability into fitness-density functions, we can evaluate the functional form of isodars expected under different disturbance-fitness relationships. Using this framework, we investigated how a gradient of forest harvesting disturbance and differences in resource availability influenced habitat quality for snowshoe hares (Lepus americanus) and red-backed voles (Myodes gapperi) using pairs of logged and uncut boreal forest. Isodars for both species had positive intercepts, indicating reductions to maximum potential fitness in logged stands. Habitat selection by hares depended on both conspecific density and differences in canopy cover between harvested and uncut stands. Fitness-density curves for hares in logged stands were predicted to shift from diverging to converging with those in uncut forest across a gradient of high to low disturbance intensity. Selection for uncut forests thus became less pronounced with increasing population size at low levels of logging disturbance. Voles responded to differences in moss cover between habitats which reflected moisture availability. Lower moss cover in harvested stands either reduced maximum potential fitness or increased the relative rate of decline in fitness with density. Differences in vole densities between harvested and uncut stands were predicted, however, to diminish as populations increased. Our findings underscore the importance of accounting for density-dependent behaviors when evaluating how changing habitat conditions influence animal distribution.     

Parental investment related to social systems in microtines

Summary Laboratory offspring of wild-caught voles Clethrionomys glareolus and Microtus agrestis, bred for 1 year under constant conditions, were examined with regard to sex ratios and weights at weaning and at 2 months of age. C. glareolus exhibits female territoriality and M. agrestis male territoriality in summer. The adults die away in late summer-autumn. Early-summer young mature in the year of birth but late-summer young do not reach maturity until the following year. C. glareolus young showed a male bias in early summer and a female bias in late summer. Conditions were the opposite but less clear in M. agrestis. C. glareolus males grew comparatively faster than M. agrestis males and showed a markedly higher early summer male: female weight ratio at 2 months of age. Maternal investment thus appears clearly related to the social system; the sex with the largest number and highest quality of young was that which was not limited in number by territoriality. However, the investment depended also on the time until maturation of the young.     

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.     

Geographic differences in the sociability of voles in relation to cyclicity

Bank voles, Clethrionomys glareolus, and field voles, Microtus agrestis, redistribute food hoards and alter the location of their activity upon the introduction of a foreign conspecific in laboratory experiments. This behaviour showed geographic variation in C. glareolus but no clear geographic variation in M. agrestis; north Swedish cyclic C. glareolus were repelled by the new animals while southern non-cyclic C. glareolus were attracted. There were no differences between animals from various population phases at the same locality. The geographic differences were retained in laboratory-born animals kept in an abnormal social environment. Other studies have demonstrated similar differences in M. agrestis. Cyclic voles may be negatively affected by density-related stimuli while non-cyclic ones may instead gather in environments that are fairly safe from predators.     

Variation in the diet composition of a generalist predator, the red fox, in relation to season and density of main prey

Diet composition of a generalist predator, the red fox (Vulpes vulpes) in relation to season (winter or summer) and abundance of multi-annually cyclic voles was studied in western Finland from 1983 to 1995. The proportion of scats (PS; a total of 58 scats) including each food category was calculated for each prey group. Microtus voles (the field vole M. agrestis and the sibling vole M. rossiaemeridionalis) were the main prey group of foxes (PS = 0.55) and they frequently occurred in the scats both in the winter and summer (PSs 0.50 and 0.62, respectively). There was a positive correlation between the PSs of Microtus voles in the winter diet of foxes and the density indices of these voles in the previous autumn. Other microtine rodents (the bank vole Clethrionomys glareolus, the water vole Arvicola terrestris and the muskrat Ondatra zibethicus) were consumed more in winter than in summer. The unusually high small mustelid predation by red foxes (PS = approx. 0.10) in our study area gives qualitative support for the hypothesis on the limiting impact of mammalian predators on least weasel and stoat populations. None of the important prey groups was preyed upon more at low than at high densities of main prey (Microtus voles). This is consistent with the notion that red foxes are generalist predators that tend to opportunistically subsist on many prey groups. Among these prey groups, particularly hares and birds (including grouse), were frequently used as food by foxes.     

A preliminary assessment of the habitat selection of two Palaearctic migrant passerine species in West Africa

We investigated the habitat selected by two Palaearctic migrants (Pied Flycatcher, Ficedula hypoleuca, Willow Warbler, Phylloscopus trochilus) in a patchy landscape in Ivory Coast and compared it with the habitat selection of Afrotropical species in the same foraging guilds. Transect counts were used to test the hypothesis that migrants use more open and more seasonal habitats and have a broader use of habitats compared with resident species. Habitats compared were, in order of decreasing tree density, gallery forest, an isolated forest and bush/tree savanna. The isolated forest had the most pronounced seasonal changes (deciduous trees) and was the one with the most diverse vegetation structure. The habitat where both migrants were most frequent was the isolated forest, and thus occurred in the habitat with the most pronounced seasonal change. Diversity of habitats selected was highest in migrants but in the Pied Flycatcher this was possibly an artefact due to subdominant individuals being excluded from the preferred habitat by territorial birds. Potential competition for habitat with Afrotropical species was found to be low.     

Landscape and local correlates with two green tree-frogs,Rhacophorus (Amphibia: Rhacophoridae) in different habitats,central Japan

Changes in land-use and local factors have strong influences on amphibian distributions. However, the effects of both combined factors on assemblages are still unclear. The green tree-frogs, Rhacophorus, are sensitive to differences in landscape fragmentation. Herein, two species of Rhacophorus schlegelii and Rhacophorus arboreus have been surveyed, along the ecological gradient ranging from paddy-dominated plains to forest-dominated mountains (located in Toyota, Okazaki, and Shinshiro, central Japan). The effects of landscape and local factors on the existence of the above frogs were investigated using generalized linear models (GLMs); the Akaike Information Criteria (AIC) were used for model selection. The lowest AIC for R. schlegelii and R. arboreus were obtained at a 250-m-radius and 500-m-radius buffers, respectively. Habitat selections of each species showed distinct results in the paddy-field vs. ponds group. Nevertheless, both species exhibited a high abundance with increasing elevation and forest cover. At the local level, R. schlegelii displayed positive relationships with the presence of trees and the proportion of embankment vegetation. Our findings demonstrate that each species responded to various landscape scale and spatial composition, habitat, and species-dependent achievements of appropriate management actions to improve restoration practices are required.

     

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

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