Resource partitioning of sympatric small mammals in an African forest‐grassland vegetation mosaic

It is often hypothesized that two species competing for the same resource cannot stably coexist unless they partition their resources in space and time. More recently stable isotope analyses have complemented traditional, observation‐based niche research by conceptualizing many of the characteristics of communities, for example, trophic niche width and the partitioning of resources. Here we quantify resource partitioning of sympatric small mammal species in an African ecosystem by analysing stable isotope ratios of hair collected from a South African forest‐grassland vegetation mosaic, and combine this with known spatial and temporal behavioural data to interpret community competition and resource partitioning. We observe niche separation to different degrees across the entire community, with different species displaying either unique isotopic dietary preferences, or partitioning resources in space and/or time. δ¹³C values were more enriched in species that inhabited afromontane grassland compared with those that inhabited afromontane forest, a reflection of the dominant vegetation in each habitat. Contrary to expectations, arboreal rodents occupied higher trophic positions than terrestrial rodents and approaching δ¹⁵N values similar to insectivorous shrews, suggesting that arboreal rodents feed on items such as arthropods enriched in ¹⁵N. While grassland species display phenotypic plasticity in terms of dietary preferences, small mammals that occurred in forests display narrow niche preferences, suggesting these species may be particularly sensitive to habitat modifications. Our results illustrate that the use of stable isotopes can be used in conjunction with spatial and temporal behavioural knowledge to elucidate resource partitioning in small African mammal communities.     

Interspecific competition in small rodents: from populations to individuals

The role of interspecific competition in shaping animal and plant communities has formed one of the major issues in ecology for decades. Small mammals, mainly rodents, have been among the model systems used for research on interspecific competition. Most studies within small mammal systems in the past have examined effects of competition on population attributes such as on population size, habitat use, or population dynamics. Population-level responses are the cumulative effects of individual responses, however, the influence of competition on individual life-history traits has rarely been studied. Research on life-histories may bridge gaps between population biology and effects of competition on individual behaviour. In this paper, we review recent research approaches to interspecific competition in rodents based on census data and species assemblages, that use regression analysis, time series analysis, removal and exclusion experiments, and showcase our own experimental research on the effects of interspecific competition on individual life-history traits in boreal voles.     

Habitat preferences and biomasses of small mammals in Swaziland

This paper describes the distribution and abundance of small mammals in Swaziland in relation to quantified habitat features. Eighteen habitat sites were sampled at seven different locations in all four geographical regions of Swaziland. Small mammal diversity (as described by the Shannon diversity index) was positively correlated with vegetation density. A total of twelve species of rodents and seven species of shrews were captured; however, only six species were caught at more than three sites. Small mammal biomasses and densities were highest at the highveld and middleveld sites. Tall grassland at all elevations supported a high diversity of small mammals. Highveld montane forest supported a high biomass and diversity of species, while no species were captured in moist forest in the Lubombo mountains.     

Trophic Niche Differentiation and Microhabitat Utilization in a Species‐rich Montane Forest Small Mammal Community of Eastern Madagascar

Unraveling the mechanisms facilitating species coexistence in communities is a central theme in ecology. Species‐rich tropical mammal communities provide excellent settings to explore such mechanisms as they often harbor numerous congeneric species with close phylogenetic relationships. Explicit tests for the mechanisms that allow syntopic occurrence in these assemblages, however, is often hampered because of the difficulty in obtaining detailed ecological data on the organisms making up the community. Using stable nitrogen and carbon ratios of hair samples, we examine whether trophic niche differentiation and microhabitat segregation explain the coexistence of 21 small mammal species at a montane humid forest site in eastern Madagascar. Overall, the community was trophically diverse and covered wide isotopic space. This diversity was based on: (1) a multi‐layered trophic community structure with mainly frugivorous‐granivorous rodents (subfamily Nesomyinae) as primary consumers and insectivorous tenrecs (family Tenrecidae) as secondary and tertiary consumers; (2) trophic segregation of rodents and tenrecs with the latter occupying different microhabitats; and (3) a dense and regular packing of species in the community. The 12 locally occurring Microgale shrew tenrecs (subfamily Oryzorictinae) showed high trophic redundancy, but were maximally spaced from each other within the trophic space covered by the genus. Results of stable isotope analysis suggest that in combination the differentiation of microhabitats and trophic niches explain the coexistence of small mammals in this community. Congeneric species appeared to be under more intense competition compared with non‐congeneric species and their coexistence can only partly be explained by trophic and microhabitat niche segregation.     

Defaunation and fragmentation erode small mammal diversity dimensions in tropical forests

Forest fragmentation and defaunation are considered the main drivers of biodiversity loss, yet the synergistic effects of landscape changes and biotic interactions on assemblage structure have been poorly investigated. Here, we use an extensive dataset of 283 assemblages and 105 species of small mammals to understand how defaunation of medium and large mammals and forest fragmentation change the community composition and diversity of rodents and marsupials in tropical forests of South America. We used structured equation models to investigate the relationship between small mammal species, functional and phylogenetic diversity with forest size, forest cover and the occurrence of medium and large mammals. The best‐fit model showed that defaunation reduced functional diversity, and that species diversity of small mammals increased with forest patch size. Forest cover did not affect functional and phylogenetic diversity. Our results indicate that occurrence of medium and large sized mammals (probably acting as predators, or competitors of small mammals) and forest patch size help to retain species and functional diversity in small mammal communities. Further, the number of species in a small mammal community was critical to the maintenance of phylogenetic diversity, and may have a pronounced influence on the ecological functions played by small mammals. Identifying how phylogenetic and functional diversity change in function of human pressures allows us to better understand the contribution of extant lineages to ecosystem functioning in tropical forests.     

Small mammal assemblages in a disturbed tropical landscape at Pozuzo,Peru

Deforestation in the Peruvian pre-montane forests has increased alarmingly in recent years. Particularly, fragmentation in these forests takes place against a background of very extensive continuous forest. Until now, no studies have detailed how Peruvian small mammals responded to forest fragmentation. In this paper, we report the results of an assessment of nonvolant small mammal fauna in forest remnants (one large fragment and one small fragment) and cattle pasture in a disturbed landscape at Pozuzo, formerly a pre-montane tropical forest in central Peru. In each forest remnant, we quantify habitat parameters and identified small mammals to know if there is a relationship between forest remnant size or microhabitat structure and small mammal assemblages. In addition, we investigated the influence of habitat structure on the presence of small mammal species. We did not find any evidence that habitat structure in large fragment was greater than small fragment. Actually, both forest remnants presented habitat features typical of forests subject to higher levels of disturbance. We found that abundance of small trees (10 - 19 cm DBH) was positively associated with the presence of Marmosops noctivagus, but negatively with the presence of Oecomys bicolor. Likewise, the presence of the arboreal Rhipidomys leucodactylus was associated with the abundance of larger trees (DBH > 50 cm). We recorded 12 small mammal species in large fragment, 9 in small fragment and 3 in cattle pasture. No significant differences were found in species richness between both fragments, but marsupial abundance in the small fragment was greater than large fragment. At Pozuzo, we recorded 9 species restricted to forest habitat and 3 to forest and cattle pastures. However, it is necessary to assess more fragments to improve our knowledge about species survival in this landscape.     

Habitat patch and matrix effects on small-mammal persistence in Amazonian forest fragments

Tropical forest mammal assemblages are widely affected by the twin effects of habitat loss and habitat fragmentation. We evaluated the effects of forest patch metrics, habitat structure, age of patch isolation, and landscape metrics on the species richness, abundance and composition of small mammals at 23 forest fragments (ranging in size from 43 to 7,035 ha) in a highly deforested 3,609-km² landscape of southwestern Brazilian Amazonia. Using pitfall traps and both terrestrial and arboreal traplines of Sherman, Tomahawk and snap traps, we captured a total of 844 individuals over 34,900 trap-nights representing 26 species and 20 genera of small-mammals, including 13 rodent and 13 marsupial species. We also consider the effects of distance from forest edges on species occupancy and abundance. Overall small mammal abundance, species richness and species composition were primarily affected by the quality of the open-habitat matrix of cattle pastures, rather than by patch metrics such as fragment size. Ultimately, small mammal community structure was determined by a combination of both landscape- and patch-scale variables. Knowledge of the anthropogenic factors that govern small mammal community structure is of critical importance for managing the persistence of forest vertebrates in increasingly fragmented neotropical forest landscapes.     

The importance of considering both taxonomic and habitat guild approaches in small mammal research

Studies on the effects of habitat fragmentation on small mammals often lead to confounding results as they only consider taxonomic groups in their analysis and neglect functional diversity of the communities. Here we describe the structure and composition of small mammal communities at 22 sites, ranging from 41 to 7035 ha, in a hyper‐fragmented landscape of an Amazonia‐Cerrado ecotone. Also, in considering a taxonomic and habitat guild approach, we report the effects of habitat structures and patch spatial attributes on richness, abundance and species composition. Small mammal richness reported in southern Amazonia (N = 23 species) is greater than most previous studies in the tropics. All rare small mammals captured in this study were forest interior species. Richness of forest interior species was positively related to larger patches, as shown by the species–area relationship. However, 52% of the small mammal species were in forest fragments smaller than 50 ha, highlighting the importance of preserving both large and small forest fragments in a landscape with accelerated habitat reduction. Richness of edge‐tolerant species was not associated with the tested variables, yet edge‐tolerant species were more abundant in degraded environments. Marsupials were positively associated with vertical habitat structures, while rodents were more strongly related to a ground‐level habitat structure. The landscape studied is extremely variable and has contributed to the difficulty in detecting clear patterns, particularly when considering only one approach. Because of the complementary outputs when analysing either taxonomic groups or habitat guilds, we recommend the use of multi‐taxa studies of different guilds to assist decision makers in designing conservation strategies and appropriate management of small mammal populations.     

Niche evolution in Australian terrestrial mammals? Clarifying scale-dependencies in phylogenetic and functional drivers of co-occurrence

Interactive forces between competition and habitat filtering drive many biogeographic patterns over evolutionary time scales. However, the responsiveness of assemblages to these two forces is highly influenced by spatial scale, forming complex patterns of niche separation. We explored these spatial dependencies by quantifying the influence of phylogeny and functional traits in shaping present day native terrestrial mammal assemblages at multiple scales, principally by identifying the spatial scales at which niche evolution operates. We modelled the distribution of 53 native terrestrial mammal species across New South Wales, Australia. Using predicted distributions, we estimated the range overlap between each pair of species at increasing grain sizes (~0.8, 5.1, 20, 81, 506, 2,025, 8,100 km²). We employed a decision tree to identify how interactions among functional traits and phylogenetic relatedness translated to levels of sympatry at increasing spatial scales. We found that Australian terrestrial mammals displayed phylogenetic over-dispersion that was inversely related to spatial scale, suggesting that ecological processes were more influential than biogeographic sympatry patterns in defining assemblages of species. While the contribution of phylogenetic relatedness to patterns of co-occurrence decreased as spatial scale increased, the reverse was true for habitat preferences. At the same time, functional traits also operated at different scales, as dietary preferences dominated at local spatial scales (<10 km²) while body mass has a stronger effect at larger spatial scales. Our findings show that ecological and evolutionary processes operate at different scales and that Australian terrestrial mammals diverged slower along their micro-scale niche compared to their macro-scale niche. By combining phylogenetic and niche methods through the modelling of species distributions, we assessed whether specific traits were related to a particular niche. More importantly, conducting multi-scale spatial analysis avoids categorical assignment of traits-to-niches, providing a clearer relationship between traits and a species ecological niche and a more precise scaling for the axes of niche evolution.     

Non-linear relationship between body size of terrestrial carnivores and their trophic niche breadth and overlap

Studying food partitioning of mammalian predators is important for understanding trophic structures and interactions between coexisting carnivore species. This is particularly pertinent in the light of expanding ranges of populations of generalist species whose habitat and diet overlap with more specialized species. Here, we tested the resource partitioning hypothesis in terrestrial carnivores, predicting that trophic niche breadth and overlap relate positively to body mass. We used dietary data from 18 terrestrial carnivore taxa in four families (Canidae, Mustelidae, Felidae and Ursidae; body mass 0.1–173.6 kg) in three regions in Central and Eastern Europe, i.e. deciduous forest and forest-steppe region (DFR), temperate deciduous and mixed forest region (MFR) and transitory mixed forest regions (TFR). We ranked carnivores along an axis of trophic niche (breadth and overlap), and analysed the relationship between trophic niche and body mass (or pair-wise difference in body mass). A hierarchical cluster analysis of diet composition divided carnivores into four ecological groups: wild ungulate predators; small-mammal predators; amphibians and small mammal predators and omnivores. The relationship between body mass of predators and both trophic niche breadth and trophic niche overlap were hump-shaped. The trophic niche breadth to body mass ratio was significantly lower in DFR than in TFR and trophic niche overlap was significantly higher in DFR than in MFR and TFR. The predominant food resource is small mammals whose abundance is related to local agricultural and forestry management practices. Modifications of management techniques can affect population dynamics and community composition of carnivore species, especially in the case of small-mammal predators.     

Mammal assemblages in forest fragments and landscapes occupied by black howler monkeys

Species assemblages in disturbed habitats vary as a function of the interaction between species requirements and the spatial configuration of the habitat. There are many reports accounting for the presence of howler monkeys in fragments where other mammals are absent, suggesting that they are more resilient. In the present study we explored this idea and predicted that if howler monkeys were more resilient to habitat loss and fragmentation than other mammals, mammal assemblages in fragments occupied by howler monkeys should include fewer species with decreasing amount of habitat (smaller fragment size and less habitat in the landscape) and increasing number of forest fragments. We explored these relationships by additionally considering the feeding and life habits of mammal species, as well as the isolation and proximity of each fragment to human settlements and roads. We sampled the presence of mammals in five fragments occupied by black howler monkeys (Alouatta pigra) in the Mexican state of Campeche. Through direct sights performed during 240 h in each fragment, we observed 23 species. At the landscape scale, higher fragmentation was associated with a decrease in herbivores, omnivores and total number of species. At the fragment scale semiarboreal, omnivore, and total number of species increased with increasing fragment size. This study supports the idea that howler monkeys are more resilient to forest loss and fragmentation than other native mammals, and our exploratory analyses suggest that the specific mammal assemblages that are found in fragments are related to both landscape and fragment scale spatial attributes, as well as with species-specific characteristics.     

Shifts from native to invasive small mammals across gradients from tropical forest to urban habitat in Borneo

Urbanization has paved the way for the spread of commensal rodents at global scale. However, it is largely unknown how these species use tropical anthropogenic landscapes originally covered with forests and inhabited by diverse small mammal assemblages. We surveyed non-flying small mammals in various urban and suburban habitat types and adjacent forest in the tropical town of Kota Kinabalu in Borneo. We used occupancy and polynomial regression models to determine variation in species occurrences along gradients of land-use intensity. Müller’s sundamys (Sundamys muelleri) was the only native small mammal species found in urban and suburban landscapes with a continuous decrease in occurrence probability from forests to urban habitats. The invasive Asian black rat (Rattus rattus species complex) and the invasive Asian house shrew (Suncus murinus) had the highest occurrence probabilities in habitats of intermediate land-use intensity, but Asian black rats are also likely to occasionally invade forested habitats and occupied urban habitats in sympatry with the Norway rat (Rattus norvegicus). In urban and suburban habitats, fallow land possibly favoured the occurrence of S. muelleri and S. murinus. Other native small mammal species (Muridae, Sciuridae, Tupaiidae) were found only in forested areas. Our study shows that native small mammals found in forest are largely replaced by invasive species in urban and suburban habitats. Due to their occurrence in habitats of various land use intensities, S. muelleri and R. rattus comprise central links between forest wildlife and urban species, an association that is important to consider in studies of parasite and disease transmission dynamics.     

Seasonal changes in density, demography and body composition of small mammals in a southern temperate forest

Eleven small mammal species (nine rodents and two shrews) were recorded in 16 months of trapping in the Karkloof Forest, Natal. Total densities varied seasonally, ranging from 60.7 animals/ha in the dry autumn and winter to 29.1 animals/ha at the onset of the rains in spring and early summer. Peaks in breeding coincided with high body fat and water content in the five common species during summer: in winter, when body fat and water levels were low, there was no breeding. This may be related to poor food quality at this time. Annual changes in density, breeding season and juvenile recruitment of Rhabdomys pumilio, Mastomys natalensis and Myosorex varius are the same as recorded in conspecific populations from other habitat types and do not seem modified by the habitat stability of the forest environment. Of the five common species, only Grammomys dolichurus , one of the few forest-associated small mammals in southern Africa, and Graphiurus murinus demonstrated stable, K-selectéd life-history styles.     

Interspecific relations in forest floor coleopteron assemblages: Niche overlap and guild structure

In six different environments of the Towada-Hachimantai National Park, the forest floor coleopterons were sampled by means of pitfall traps with five different baits. Their niche breadths and niche overlaps were assessed for the electivity of habitat, bait, and habitat × bait, and compared with those expected from neutral models constructed randomly. The niche breadths in the observed assemblages were smaller than those expected from the neutral model, while the niche overlaps were more widely distributed than those expected from the neutral model. Mean overlaps between species neighbors in niche space, cluster analyses, and ordinations of species revealed a guild for habitat, bait and habitat × bait. Lawlor's (1979) gamma matrices concerning the habitat electivity show that the rates of mutualism resulted from indirect interspecific competitions are significantly smaller in the guild than in the whole assemblage. This suggests that the interspecific competition among the guild members is mitigated or even converted into mutualism by joining of the species which do not belong to the guild. The rates of mutualism were also significantly larger in the resource state of two dimensions (habitat × bait) than in that of either one. Therefore, if niche overlaps are measured on the basis of resource state of multi-dimensions, the rates of mutualism may increase further.     

Diversity of rodent and shrew assemblages in different vegetation types of the savannah biome in South Africa: no evidence for nested subsets or competition

Identifying nonrandom species composition patterns predicted by assembly rules has been a central theme in community ecology. Few studies have investigated the prevalence of multiple drivers on species composition patterns in small mammal assemblages in the Old World. This study investigated seasonal changes in rodent and shrew diversity in eleven savannah vegetation types in South Africa. We tested whether species composition patterns are nonrandom with respect to predictions from Diamond's assembly rules, niche limitation hypothesis and nestedness hypothesis. Species richness estimators indicated that inventories for the rodents (80%) and shrews (100%) were relatively complete. Rodent (n = 11 species) diversity and shrew (n = 5 species) diversity were highest in summer and lowest in autumn. Rodent richness was highest in the Terminalia sericea bushveld and woodlands and lowest in the Drypetes arguta sand forest, whilst shrew richness was highest in the T. sericea bushveld and woodlands and lowest in the Acacia nilotica/Dichrostachys cinerea open shrub savannah. We found no support for the predictions of competition and nestedness hypotheses and suggest that this was probably due to the high seasonal and annual variability in rodent and shrew diversity.     

Diversity of rodents and shrews along an elevational gradient in Bwindi Impenetrable National Park, south-western Uganda

Small mammal species diversity in the major vegetation zones of Bwindi Impenetrable National Park is discussed in relation to altitude. Species richness of the small mammals was found to decrease with an increase in altitude. The main factors accounting for the observed diversity are the wide altitudinal variation and a complex array of vegetation types. Sixty‐seven species of rodents and shrews were found to exist in the Park; 47 of which were rodents and 20 shrews. Of these, 26 species are new to the Bwindi Park list. Three species have probably not been described before. The study found 10 species of small mammals to be Albertine Rift endemics. Three genera are recorded in Uganda for the first time: Rwenzorisorex, Suncus and Paracrocidura. Five species are new records for East Africa. These are Crocidura stenocephala, Lophuromys rahmi, L. medicaudatus, Paracrocidura maxima and Hylomyscus aeta. Because of the high endemism of plants, butterflies, birds and now of small mammal species, Bwindi forest is a unique biodiversity hotspot and is among the highest conservation priorities in the Albertine Rift.     

Termites, Large Herbivores, and Herbaceous Plant Dominance Structure Small Mammal Communities in Savannahs

Large herbivores and termites are important functional groups in African savannahs. Both groups affect small mammals, which are also important determinants for savannah structure and function. Because vegetation on Macrotermes mounds are preferentially grazed by large herbivores, and mounds represent resource-rich distinct habitat patches for small mammals in relatively resource-poor savannahs, termite mounds are ideal sites for studies of how grazing by large mammals and productivity affect communities of small mammals. We conducted an experiment in Lake Mburo National Park, Uganda, with four treatments: large vegetated Macrotermes mounds (with and without large herbivores) and adjacent savannah areas (with and without large herbivores). We replicated the treatment blocks nine times and trapped small mammals regularly over a period of almost 2 years. Small mammal species assemblages differed considerably between mounds and savannah areas. Grazing had a substantial effect on small mammal species assemblages in the resource-poor savannah, but not in the relatively resource-rich termitaria. Small mammal species abundance, biomass, and richness were higher on termite mounds than adjacent savannah areas. Excluding large herbivores caused a major increase in species abundance, biomass, and richness both on savannah and termitaria. Herbaceous plant species evenness was an important determinant of the small mammal community. Small mammal biomass increased with high plant dominance, indicating that a few dominant plant species are important for biomass production of small mammals. Small mammal diversity was not related to any of the treatments, but increased with plant species evenness as well as richness. Fencing increased species dominance in the small mammal community on both savannah and termitaria, probably because competitive patterns shift from inter-guild (that is, between large and small mammals) to intra-guild (that is, between small mammals) when large mammals are excluded. The study highlights the complex interactions among large herbivores, termites, herbaceous plants, and small mammals in African savannahs. When studying the structure and function of small mammal communities it is therefore important to consider several coexisting functional groups.     

Habitat preferences of medium/large mammals in human disturbed forests in Central Japan

It is a great concern whether human disturbed forest, i.e., secondary forest or monoculture forest, can be a habitat for the wildlife. We examined habitat preference of medium/large terrestrial mammal species in a human-disturbed mountainous settlement area of Akiyama Region, Central Japan, by using camera-traps, which were placed at conifer plantation forest, high/low-disturbed deciduous broad-leaved forest, and broad-leaved/conifer mixed forest in 2008 and 2009. Camera-traps were operational for 4568.6 trap-nights producing 740 photo-captures of 13 medium/large terrestrial mammal species. Japanese serow and Japanese hare dominated 54 % of all photo-captures. Low-disturbed deciduous broad-leaved forest showed the richest mammal fauna (12 spp.). We suggested that fruit trees and understory vegetation provided suitable habitat for frugivorous as well as herbivorous mammals. The mixed forest showed the lowest mammal fauna (6 spp.), which located higher elevation. High-disturbed deciduous broad-leaved forest (9 spp.) was also supposed to be a better habitat for some frugivorous mammals and herbivorous mammals, but tended to be avoided by Japanese marten and Japanese macaque. Conifer plantation forest (7 spp.) with understory vegetation was supposed to be a better habitat for Japanese serow and Japanese hare, but not for the other species. Without fruit trees, conifer plantation forest was supposed to be non-attractive habitat for frugivorous mammals. We demonstrated that, if a forest had been disturbed by human, the forest can be a habitat for terrestrial herbivores, but this was not always true for frugivores.