Habitat structure and disturbance affect small mammal populations in Mediterranean forests

Habitat structure, the physical arrangement of objects in space, affects animal populations, ecological interactions, species diversity, and ecosystem functioning. Conventional forest management practices that reduce habitat structural complexity are thus under reappraisal, and there are increasing efforts to understand how habitat structure affects functionally important species. We investigate here the effect of habitat structures on small mammal dens and their interactions with widespread antagonistic species (wild boars). Among habitat structures we considered tree dimension (diameter), shrub cover, rock cover, dead wood volume, and stump area. Ground cover with rooting signs measured wild boar disturbance. The number of small mammal dens was related positively to rock cover and tree diameter, and negatively to wild boar disturbance. Additionally, a positive interaction between wild boar disturbance and tree diameter emerged, showing that the effect of big trees on small mammal nesting was more evident with higher levels of disturbance. This study suggests that habitat structures that can be affected by forest management provide effective refuges for functionally important species. The effect of habitat structure on species survival and interactions thus needs to be addressed to understand ecosystem management and functioning.     

Energy flow through small mammal populations

In order to compile an annual or seasonal energy budget for a small mammal population, it is necessary to derive estimates of standing crop, consumption, egestion, digestion, excretion, assimilation, respiration, growth and reproduction. This review gives an account of the methods available for compiling such budgets, and uses results obtained from a study of Microtus agrestis in south west Britain as a worked example. This population was live–trapped on twenty occasions during the course of 2 years. Total energy flow was 309 kJ/m² in the first year and 173 kJ/m² in the second. The average coefficient of digestibility of the natural diet was 54–3%, the average net production efficiency was 1 –0% and the average gross ecological efficiency was 1 –2%. Peak energy flow occurred in spring and late summer each year, and coincided with peaks in the abundance and growth rate of the grasses which formed the main source of food. Previously published estimates show that the annual flow of energy is higher amongst small mammal populations inhabiting open habitats, such as grasslands, than it is in successionally more mature habitats, such as woodlands.     

A mathematical analysis of small mammal populations

Populations of Microtus montanus, the montane vole, have been extensively studied. It is known that their reproductive activity is closely linked to the availability of the chemicals in growing plants. We use a mathematical model here to study how the length of the vegetative season and the natural reproduction rhythm of voles are involved in the long term dynamics of the population numbers. In particular, we use data obtained from Timpie Springs, Utah, and from Jackson Hole, Wyoming, to formulate a model. The novelty of this model is its use of littering curves that highlight the temporally discrete nature of vole reproduction. The model shows how the timing of the vegetative season can influence vole population sizes.This work was supported in part by NSF Grant No. MCS-82-08986 and DMS85-14000 (FCH) and by the University of Utah where Dr. Murphy was a visitor     

Demographic strategies in fluctuating populations of small rodents

Summary A graphic model for individual selection determined by the logarithmic growth rates, dN _i /N _i ·dt, is developed for studying optimal demographic strategies at different phases of microtine cycles. In a density-independent situation (no crowding) selection leads to maximization of total life-time reproductive output (or equivalently, the Malthusian parameter, m) at the expense of competitive (contest type) abilities. In a density-dependent situation (crowding), selection leads to maximization of contest type competitive abilities at the expense of reproductive output. These two modes of selective pressure are called r- and -selection. r-selection is presumed to occur during the increase phase of a cycle. As the habitat becomes crowded, -selection takes over and is assumed to be extreme at high densities.The characteristics of r- and -strategists are found to be similar to those of the docile and aggressive forms presumed in Chitty's theory for fluctuating populations. Literature supports the attributes predicted by the theory. I argue that sensitivity to density-independent factors is higher in the -strategy. On the basis of a graphic model, I show that the -strategists' high sensitivity to extrinsic factors will account for the crash in microtine cycles.On the basis of these theoretical considerations, Chitty's theory for fluctuating populations is interpreted to imply that interactions between intrinsic and extrinsic (random) factors will result in cycles. A graphic model for this interpretation of Chitty's theory is discussed. The heterogeneity of the habitat is an important aspect.According to theory, dispersal of pregnant females is explained as an adaptation leading to increased current reproductive output. This behaviour is presumed to dominate numerically during the increase phase of a cycle, a prediction supported in literature.     

The ecological connections of dispersal in small mammal populations]

The dispersal of 27 Holarctic species of rodents was quantitatively described on the base of long-term catch method. The relative proportion of non-territorial individuals is an important parameter that reflects the intensity of dispersal. This parameter is statistically related to population (abundance and population type) and species characters (the size of species range and the level of intraspecific differentiation). The contribution of species characters to the variance of migrant portion is lower than contribution of population ones. Isolation and population specialisation are accompanied by the decrease in the portion of disperser, while in pessimal environment the increase in population fluidity is observed. Intensity of dispersal is positively correlated with the size of species range and negatively--with the level of intraspecific differentiation. Thus the dispersal in populations of small mammals serves for colonisation and reparation, providing the stable existence of given group in variable environment.     

The structure and stability of small mammal faunas

Summary I examined questions about the local and global stability and resilience characteristics of six small mammal faunas and their relationship to connectance, average interaction strength, community covariance, and indirect competitive pathways. Community matrices estimated elsewhere were used in the analyses. The faunas had from three to nine species and represented several habitat types. The following properties were observed: (1) all community matrices were globally stable, (2) return rates to equilibrium were generally maximized, and (3) competitive interactions were strictly hierarchical. Global stability resulted from a reduction in connectance with increasing species and not from structural characteristics (i.e., community covariance). Average interaction strength did not change with the number of species in these faunas. Increased resilience was due largely to reduced community covariance. Two of the six faunas showed some evidence for indirect pathways, but this appeared to be artifactual. Randomizations of the original community matrices showed that indirect pathways have a high probability of occurrence. Monte Carlo simulations indicated that the probability of indirect pathways increases with increasing number of species or magnitude of competition. These results bring into question previous observational studies invoking indirect competitive pathways. Communities without indirect pathways have a special linear hierarchical structure. This structure has greater resilience than when indirect pathways are present. The observed patterns in community structure are discussed with regard to current theories of habitat selection.     

Road effects on demographic traits of small mammal populations

Recent studies have highlighted the positive effects of road verges on the abundance of small mammals. However, most of these studies occurred in intensively grazed or cultivated areas, where verges were the last remnants of suitable habitats, which could mask the true effects of roads on population traits. We analysed the effects of roads on small mammal populations living in a well-preserved Mediterranean forest. We used the wood mouse (Apodemus sylvaticus) as a model of forest-dwelling small mammals that probably are among the species most affected by road clearings. Our study compared populations in similar habitat areas with and without road influence. We assessed abundance, survival and temporary emigration using extended Pollock’s robust design capture-recapture models. Moreover, we analysed population turnover, sex ratio, age structure and body condition. We found that wood mouse abundance and body condition were lower at the road bisected area, whereas the remaining population traits were similar. This suggests that the reduced habitat availability and quality due to the physical presence of the road and verge vegetation clearing are the main drivers of demographic differences in wood mouse populations between areas. Nevertheless, our results also suggest that in high-quality habitats surrounding national roads, wood mouse populations present similar dynamics to others living in undisturbed areas, despite the decrease in abundance and body condition. Overall, the often-reported increased small mammal abundance in road surroundings should not be generalized independently of habitat quality or to other population traits.     

On the problem of trap-response types of small mammal populations

1. So far as the population data thus far amassed go, commensal rats in residential habitats are of type II; house mice may be of type I or III in fields but might show type II in residences, and fieral species or subspecies may range from type I to III. Altogether alloresponsive populations seem to be much commoner than the isoresponsive.
2. It is as yet little disclosed to what degree the differentiation of the type is correlated with speciation or subspeciation, even if only a proof that a subspecific population would hold type III as its fixed feature is given. The response types are supposed to be associated with the levels of population shyness. Hence some characters reflected in those may possibly be connected with sexual isolation provoking speciation.
3. Some awkward respects due to the heterogeneous trappability, depending upon the trap-response type, among a mixed population of marked and unmarked animals have been confirmed regarding the methodology for estimating parameters.

     

Synchrony in small mammal populations of montane forest patches in southern India

1. Small mammal populations were studied in montane evergreen forests in the Nilgiris, southern India, using live-trapping from January 1994 to September 1996. Two sites were selected, one with a single large forest patch and adjoining plantations, and the other with several small forest patches separated by grassland.
2. Nine species were recorded, of which eight were trapped in the forest patches, two in the grasslands and five in plantations. Rattus rattus was the most abundant species (2–36 individuals ha⁻¹) followed by Suncus montanus (0–11 individuals ha⁻¹). Densities of seven other species recorded were very low.
3. Synchrony in various population variables – density, biomass, mean weight, density of adults and adult females, and proportion of adults, adult females and sub-adults – was examined for Rattus rattus populations in the forest patches and plantations. Density and biomass were studied in seven other species recorded at these sites. Spearman's rank order correlation was used as a measure of synchrony between the population variables.
4. Within-site synchrony was higher than between-site synchrony in population characters. Synchrony was also higher between plots within the unfragmented site than they were between plots in the fragmented site. Relatively high synchrony in proportions of adults, adult females and juveniles in the forest patches implied that breeding is probably influenced by climate and food availability, which are seasonal in this habitat.
5. Given the small patch sizes (≈1–10 ha) and low population sizes, asynchrony is likely to be an outcome of demographic and environmental stochasticity, and low dispersal rates may impede establishment of synchrony.     

Foraging in space and time structure an African small mammal community

We used live-trapping and foraging to test for the effect of habitat selection and diet on structuring a community of six small mammals and one bird within the Soutpansberg, South Africa. We established grids that straddled adjacent habitats: woodland, rocky hillside, and grassland. Trapping and foraging were used to estimate abundance, habitat use, and species-specific foraging costs. The species with the highest abundance and foraging activity in a habitat, activity time, or food was considered the most efficient and presumed to have a competitive advantage. All species exhibited distinct patterns of spatial and temporal habitat preference which provided the main mechanism of coexistence, followed by diet selection. The study species were organized into three assemblages (α diversity): grassland, Rhabdomys pumilio, Dendromus melanotis, and Mus minutoides.; woodland, Aethomys ineptus and Micaelamys namaquensis; and rock-dwelling, M. namaquensis and Elephantulus myurus. Francolinus natalensis foraged in open rocky areas and under wooded islands within the grassland. Species organization across the habitats suggested that feeding opportunities are available within all habitats; however, distinct habitat preferences resulted from differing foraging aptitudes and efficiencies of the competing species. At Lajuma, species distribution and coexistence are promoted through distinct habitat preferences that were shaped by competition and species-specific foraging costs. The combination of trapping and foraging provided a mechanistic approach that integrates behavior into community ecology by ‘asking’ the animal to reveal its perspective of the environment. Using spatial and temporal foraging decisions—as behavioral indicators—enables us to guide our understanding for across-taxa species coexistence.     

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.     

Genetic structuring in fluctuating populations

The effect of genetic drift in spatially distributed dispersal-linked and density-regulated populations is studied in a classical one-locus two-allele system. We analyse emergence of genetic differentiation assuming random drift only, where the noise-like variability is due to demographic stochasticity. We find emergence of clusters of sub-units with local allele fixation and persistence of both alleles in lengthy simulations. We demonstrate that local allele fixation (extending over a number of adjoining spatial sub-units) – without global loss of alleles – may occur when the carrying capacities of local patches are small, under a full range population dynamic regimes, when dispersal rate is small, and when redistribution (through dispersal) does not act as global mixer. These results are novel. The key to the observations is that drift is simultaneously influenced by distance-dependent dispersal, demographic stochasticity and autocorrelated population fluctuations due to delayed-density dependence. These are standard elements of contemporary population models in spatially structured context. With stable large populations, no stochasticity and dispersal limited to neighbours only, our model collapses to the stepping-stone model, while with dispersal being random and global, the model collapses to Wright's island model.     

A long term study of small mammal populations in a Brazilian agricultural landscape

Long-term monitoring of small mammal populations is very important to understand the variations in temporal abundance on a large time scale, which are related to ecological, economic and epidemiological phenomena. The aim of this study is to monitor the populations of the marsupials Didelphis aurita and Philander frenatus and the rodents Nectomys squamipes, Akodon cursor and Oligorysomys nigripes in a locality of typical Brazilian rural landscape, Sumidouro Municipality, Rio de Janeiro State, Brazil. A mark-recapture study was conducted during five years. We analyzed the population dynamics, the reproduction and age structure of these species. Both marsupials presented higher population sizes in the end of wet period and beginning of the dry period, which can be explained by the seasonal reproduction which begins in the middle of the dry period and ends in the last months of the wet period. N. squamipes reproduced throughout the year but mostly during rainy periods, due to the close association of this rodent to resources found in the water. Higher survivorship and recruitment rates were in the end of the wet season. The rodent A. cursor had an opportunist reproduction, resulting in high turnover rates. Survivorship increased with the effects of the dry periods. O. nigripes showed a clear annual pattern of population cycle with peaks during the dry season. The rodents did not show potential to present outbreaks and become agricultural pests. The annual population cycles of O. nigripes and the unique peak of A. cursor population during five years highlight attention to their importance as wild reservoirs of the hantavirus disease. Their ecological characteristics associated to their opportunistic behavior make these species prone to be good reservoirs of zoonoses.     

Estimating fluctuating selection in age-structured populations

In age-structured populations, viability and fecundity selection of varying strength may occur in different age classes. On the basis of an original idea by Fisher of weighting individuals by their reproductive value, we show that the combined effect of selection on traits at different ages acts through the individual reproductive value defined as the stochastic contribution of an individual to the total reproductive value of the population the following year. The selection differential is a weighted sum of age-specific differentials that are the covariances between the phenotype and the age-specific relative fitness defined by the individual reproductive value. This enables estimation of weak selection on a multivariate quantitative character in populations with no density regulation by combinations of age-specific linear regressions of individual reproductive values on the traits. Demographic stochasticity produces random variation in fitness components in finite samples of individuals and affects the statistical inference of the temporal average directional selection as well as the magnitude of fluctuating selection. Uncertainties in parameter estimates and test power depend strongly on the demographic stochasticity. Large demographic variance results in large uncertainties in yearly estimates of selection that complicates detection of significant fluctuating selection. The method is illustrated by an analysis of age-specific selection in house sparrows on a fitness-related two-dimensional morphological trait, tarsus length and body mass of fledglings.     

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.     

Patterns in the structure of Asian and North American desert small mammal communities

Aim We compared assemblages of small mammal communities from three major desert regions on two continents in the northern hemisphere. Our objective was to compare these with respect to three characteristics: (1) species richness and representation of trophic groups; (2) the degree to which these assemblages exhibit nested community structure; and (3) the extent to which competitive interactions appear to influence local community assembly. Location We studied small mammal communities from the deserts of North America (N=201 sites) and two regions in Central Asia (the Gobi Desert (N=97 sites) and the Turan Desert Region (N=36 sites), including the Kara-Kum, Kyzyl-Kum, NE Daghestan, and extreme western Kazakhstan Deserts). Method To provide baseline data we characterized each desert region in terms of alpha, beta, and gamma diversity, and in terms of the distribution of taxa across trophic and locomotory groups. We evaluated nestedness of these communities using the Nestedness Temperature Calculator developed by Atmar & Patterson (1993, 1995) , and we evaluated the role of competitive interactions in community assembly and applied a null model of local assembly under varying degrees of competitive interaction ( Kelt et al., 1995, 1996 ). Results All three desert regions have low alpha diversity and high beta diversity. The total number of species in each region varied, being highest in North America, and lowest in the Turan Desert Region. The deserts studied all present evidence of significant nestedness, but the mechanism underlying this structure appears different in North American and Asia. In North America, simulations strongly implicate interspecific competition as a dominant mechanism influencing community and assemblage structure. In contrast, data from Asian desert rodent communities suggest that these are not strongly influenced by competition; in fact, they have greater numbers of ecologically and morphologically similar species than expected. These results appear to reflect strong habitat selection, with positive associations among species that share similar habitat requirements in these communities. Our analyses support earlier reports suggesting that predation and abiotic forces may have greater influences on the assembly and organization of Asian desert rodent communities, whereas interspecific competition dominates assembly processes in North America. Additionally, we suggest that structuring mechanisms may be very different among the two Asian deserts studied. Gobi assemblages appear structured by trophic and locomotory strategies. In contrast, Turan Desert Region assemblages appear to be randomly structured with respect to locomotory strategies. When trophic and locomotory categories are combined, however, Turan species are positively and nonrandomly associated. Main conclusions Very different ecological dynamics evidently exist not only between these continents, but within them as well. These small mammal faunas differ greatly in terms of community structure, but also appear to differ in the underlying mechanisms by which communities are assembled. The underlying role of history and geography are strongly implicated as central features in understanding the evolution of mammalian faunas in different deserts of the world.     

Temporally fluctuating prey and interfering predators: a positive feedback

Prey densities often show fluctuating patterns over various timescales. Focusing on short-term, within-generation fluctuating patterns of local prey availability, we suggest that prey that show synchronized and high-amplitude fluctuations in availability experience decreased risks of predation, but also enhance the maintenance of predator interference hierarchies by affecting the relative foraging success of unequal conspecific interferers. When predators interfere with each other, they forage less intensely on prey, which benefits prey in terms of decreased predation risk. The system hence involves a positive feedback. We thus argue that short-term temporal fluctuations in local prey availability could be an important mechanism behind how interference-structured social predator systems are developed and sustained. The temporal fluctuations also have implications for the phenotypic diversity of predators, and may be involved in speciation processes.     

British mammal populations: fifty years of change

• 1 Agricultural intensification, urbanization, overpopulation by both native and invasive species and diseases have all had marked effects on populations of British mammals over the last 50 years.
• 2 Here, accounts are presented of the effects of these processes with a prospective view of what the future holds for the mammals of Britain.