Small mammal diversity and dynamics within Nigeria,with emphasis on reservoirs of the lassa virus

Nigeria has a rich small mammal community, with several species implicated as carriers of zoonotic microbes such as the Lassa virus (LASV). We sought to elucidate the diversity and distribution of these animals (including known LASV reservoirs) geographically, habitat-wise and seasonally. Our DNA-assisted survey detected at least 19 small mammal species amongst 790 specimens. Diversity indices were similar between ecological zones and also between endemic and non-endemic areas for Lassa fever. Mastomys natalensis, the most renowned LASV host, was present in eight out of nine localities sampled. We also described the spatial occurrence of other known LASV hosts such as M. erythroleucus and Hylomyscus pamfi, including carriers of LASV-like arenaviruses such as Mus (Nannomys) spp. The most numerous rodents (Mastomys natalensis, M. erythroleucus, and Praomys daltoni) were captured mainly inside human dwellings. Reproductive activity occurred throughout the year, but led to population peaks for M. natalensis in the dry season and for M. erythroleucus and P. daltoni in the rainy season. Extensive geographic distribution of LASV rodent reservoirs, with population peaks in different seasons, shows that the risk of rodent-to-human transmission of LASV is greater than currently realized.     

Ecology of a typical West African Sudanian savannah rodent community

The small‐mammal community of a typical Sudanian savannah area of south‐eastern Senegal was studied by way of intensive sampling at the beginning (July) and at the end (November) of the rainy season. A total of 5931 trapnights and 200 pitfallnights yielded 767 small mammals, including 757 rodents of 12 species, as well as a few shrews and hedgehogs. Accumulation curve indicated that the rodent community was well sampled. Murine rodents were far more numerous than gerbilline rodents, and among them, Mastomys erythroleucus was the dominant species, being caught in high numbers in all habitats. Arvicanthis niloticus and Praomys daltoni were also abundant, followed by Mastomys natalensis and Gerbilliscus guineae. Abundances were high at the beginning of the rainy season when reproductive activity was low in all species. At the end of the rainy season, murine rodents were actively involved in reproduction, while gerbilline rodents were at their lowest density. Significant changes in relative abundance among habitats (including human dwellings) were observed between periods in some species, suggesting seasonal trends in habitat preferences. This may help species coexistence in this species‐rich rodent community, in an area likely to be submitted to significant habitat alterations in the years to come.     

Microhabitat choice and diet of rodents in Maynugus irrigation field, northern Ethiopia

An ecological study on the choice of microhabitat and diet of small mammals was carried out in Maynugus irrigation field of the Tigray region in northern Ethiopia. Abundance of rodents was high during the reproductive phase of the agricultural plants. Mastomys erythroleucus occurred more frequently in vegetable fields, whereas Arvicanthis dembeensis was more frequent in habitats with monocot plants. Scrotal males and reproductive females consumed more food by weight than abdominal males and quiescent females. Arvicanthis dembeensis consumed more food than M. erythroleucus. Both the species consumed more food during the main rainy season. All the cultivated plants were attacked by these rodents with a degree of difference in relation to season, species, sex, age and sexual condition. As rodent attack was more intense during the fruiting phase of the plants, rodent control management should be carried out during the flowering phase.     

Host habitat patchiness and the distance decay of similarity among gastro-intestinal nematode communities in two species of <Emphasis Type="Italic">Mastomys</Emphasis> (southeastern Senegal)

Beta-diversity, or how species composition changes with geographical distance, has seldom been studied for different habitats. We present here quantitative estimates of the relationship between geographic distance and similarity of parasitic nematode communities in two closely related rodent host species that live in habitats with very different spatial configurations. In southeastern Senegal Mastomys natalensis lives exclusively inside human villages whereas M. erythroleucus is continuously distributed outside villages. Both host species and their gastro-intestinal nematodes were sampled on the same spatial scale. Beta-diversity was found to be higher in parasite communities of M. erythroleucus than in those of M. natalensis, and significantly related to geographic distance in this first species. Even on the local spatial scale studied, host dispersal limitation, and stochastic events, may affect species turnover in nematode communities of M. erythroleucus. In M. natalensis, no relationship was found between geographic distance and nematode community similarity, however, suggesting low host dispersal rates between habitat patches. Together with previous population genetic results, this study illustrates the need for different approaches with regard to dispersal in natural populations and its effect on biodiversity.     

Structures génétiques comparées de trois espèces de rongeurs africains du genre Mastomys au Sénégal

Résumé Une analyse par électrophorèse des protéines à 20 locus a été réalisée sur trois espèces du genre Mastomys au Sénégal. Malgré l'absence de locus diagnostique, une approche multivariée par analyses factorielles (AFC et AFD) permet néanmoins de reconnaître de façon sûre une des espèces en présence (M. cf. natalensis, 100% d'individus bien classés par l'AFD) et d'attribuer correctement les individus des deux autres espèces (M. erythroleucus et M. huberti) dans 92% et 77% des cas respectivement. Bien qu'écologiquement nettement différenciées, ces deux dernières espèces apparaissent très proches génétiquement (D_Nei=0,12) et montrent par ailleurs de hétérozygoties très elevées. Les faibles distances génétiques entre les trois espèces contrastent avec la forte différenciation chromosomique observée par ailleurs. Chez M. erythroleucus et M. huberti, les distances génétiques observées entre populations continentales ne sont pas corrélées avec les distances géographiques alors que les populations insulaires présentent une nette baisse de variabilité, en relation probable avec leur isolement géographique important. Par ailleurs, chez M. huberti, plusieurs populations s'écartent fortement de la panmixie. Ces tensions génétiques pourraient résulter de phénomènes d'introgression avec M. erythroleucus.

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Compared genetic structures of three species of African rodents of the genus Mastomys from Senegal. A protein electrophoresis analysis at 20 loci was performed on three species of the genus Mastomys from Senegal. Although no diagnostic locus was found between the three species, they can be readily recognized by multivariate analysis (100% of the M. cf. natalensis, 92% of the M. erythroleucus and 77% of the M. huberti individuals could be correctly classified by discriminant analysis). Although ecologically quite clearly differentiated, M. erythroleucus and M. huberti were found to be genetically very close (D_Nei=0.12) and display very high heterozygosities. On the whole, the small genetic distances between the three species contrast with the high chromosomal differentiation that has been reported in another study. In M. erythroleucus and M. huberti, genetic distances between mainland populations are not correlated with geographic distances, whereas insular populations show an important decrease of variability probably linked with their geographic isolation. In M. huberti, some populations show a strong departure from panmictic equilibrium: these genetical tensions could result from introgression phenomena with M. erythroleucus.

     

Diversity,dynamics and reproduction in a community of small mammals in Upper Guinea,with emphasis on pygmy mice ecology

As part of a large survey on reservoirs of Lassa fever in Guinea, three villages were investigated in high endemic zone, close to Sierra Leone border. Biodiversity of the small mammal community is presented in this study through a standardized trapping in houses, cultivations and forest. Identification of the small mammals was based on morphology and by molecular technique for sibling species. Of the 1123 specimens collected in 2003–2005, we identified seventeen species (thirteen Muridae, four Soricidae), leading to high diversity (Shannon index = 1.6–1.8) and high equitability (evenness index = 0.7–0.8) in cultivations and forest. In houses conversely, the rodent community was dominated by Mastomys natalensis (95–98%), leading to low diversity and equitability. Dynamics and reproduction were investigated in two species of pygmy mice, Mus mattheyi and Mus minutoides, two species of Praomys, P. daltoni and P. rostratus, and in Mastomys erythroleucus. The pygmy mice were abundant in cultivations in early rainy season, and reproduced from rainy to dry season. Praomys daltoni was also found more abundant in cultivations and seemed to reproduce between rainy and dry season, whereas P. rostratus preferred forest and cultivations in late rainy season, and reproduced throughout the year. Finally, M. erythroleucus was more abundant in forest in dry season, and seemed to reproduce from late rainy to dry season. This species had a low occurrence (6.5%) in the Faranah’s zone, and probably lived at its southern limit in Guinea. The presence of other Murinae, such as M. natalensis, Praomys spp as possible competitors in the same habitats, is discussed. For the first time, this study relates population biology of pygmy mice with molecular identification.     

Indications of higher diversity and abundance of small rodents in human‐influenced Sudanian savannah than in the Niokolo Koba National Park (Senegal)

Expansion of human activities in the Sudanian savannah is changing natural habitats together with their animal communities. The abundance and diversity of small mammal communities were investigated in human‐influenced savannahs in close proximity to Niokolo Koba National Park (NKNP) in south‐eastern Senegal and compared with those from NKNP. Two localities were sampled in natural and two in human‐influenced areas during two dry and two rainy seasons. Total sampling effort was 5400 trap‐nights. Ninety‐eight individuals of eleven species were captured outside NKNP, while only 28 individuals of five rodent species were caught inside the park (all of which were found also outside NKNP). The most abundant species outside NKNP was Mastomys erythroleucus, while inside NKNP it was Praomys daltoni. Both relative abundance and diversity were significantly higher in human‐influenced landscape than in natural savannahs of NKNP. The difference in species richness may be linked with lower population densities of certain species in NKNP and the effect of traditional agriculture, which may support the presence of species typical for deforested landscape. Furthermore, food availability, and predator abundances are discussed as possible reasons for the higher rodent abundances outside NKNP.     

Intraspecific functional diversity of common species enhances community stability

Common species are fundamental to the structure and function of their communities and may enhance community stability through intraspecific functional diversity (iFD). We measured among‐habitat and within‐habitat iFD (i.e., among‐ and within‐plant community types) of two common small mammal species using stable isotopes and functional trait dendrograms, determined whether iFD was related to short‐term population stability and small mammal community stability, and tested whether spatially explicit trait filters helped explain observed patterns of iFD. Southern red‐backed voles (Myodes gapperi) had greater iFD than deer mice (Peromyscus maniculatus), both among habitats, and within the plant community in which they were most abundant (their “primary habitat”). Peromyscus maniculatus populations across habitats differed significantly between years and declined 78% in deciduous forests, their primary habitat, as did the overall deciduous forest small mammal community. Myodes gapperi populations were stable across habitats and within coniferous forest, their primary habitat, as was the coniferous forest small mammal community. Generalized linear models representing internal trait filters (e.g., competition), which increase within‐habitat type iFD, best explained variation in M. gapperi diet, while models representing internal filters and external filters (e.g., climate), which suppress within‐habitat iFD, best explained P. maniculatus diet. This supports the finding that M. gapperi had higher iFD than P. maniculatus and is consistent with the theory that internal trait filters are associated with higher iFD than external filters. Common species with high iFD can impart a stabilizing influence on their communities, information that can be important for conserving biodiversity under environmental change.     

Living on the verge: are roads a more suitable refuge for small mammals than streams in Mediterranean pastureland?

The retention of natural habitat corridors is a useful and practical conservation tool that can attenuate the effects of habitat loss and fragmentation on wildlife. Linear structures may contribute to the conservation of biodiversity by providing additional habitats for small fauna living in highly modified environments. We assessed the importance of road verges as refuge areas for small mammals, in highly intensified grazed pastures, within a Mediterranean landscape and compared the role of road verges as refuges with that of riparian galleries, which have been described as important shelter locations for small fauna. For this purpose, a small mammal trapping study was undertaken on two road verges and beside two small streams in southern Portugal. We captured 457 individuals of five different species, with Mus spretus the most common species captured, followed by Crocidura russula. Captures were 4.6-fold higher immediately beside both roads and streams than 12 m away in the surrounding matrix. Individuals captured in the matrix presented a smaller body size and lower body condition, suggesting that this suboptimal habitat is occupied mainly by subadults. M. spretus was 46% more abundant by roads than by streams, while C. russula was present in similar numbers in both habitats. M. spretus individuals were larger near streams but exhibited no difference in body condition between habitats. C. russula had a better body condition and slightly higher body lengths at roadsides. Our results show that roadside verges in intensively grazed Mediterranean landscapes act as important refuges and constitute equally vital habitats for small mammals as do riparian vegetation strips in landscapes where other suitable habitats are scarce.     

Species diversity and abundance of small mammals in Nechisar National Park,Ethiopia

Species diversity and abundance of small mammals were studied in Nechisar National Park, Ethiopia, during August 2008 – March 2009. Twenty species of rodents and four species of insectivores were recorded from the study area. Mastomys natalensis (17.37%), Arvicanthis dembeensis (17.09%), Mastomys erythroleucus (8.90%), Stenocephalemys albipes (8.76%), Arvicanthis niloticus (8.19%), Acomys cahirinus (7.34%), Lemniscomys striatus (6.92%), Gerbilliscus nigricauda (6.21%), Grammomys dolichurus (3.67%), Gerbilliscus robusta (2.12%), Mus proconodon (1.98%), Mus mahomet (1.41), Dendromus melanotis (1.27%), Arvicanthis abyssinicus (1.13%), Mus musculus (0.99%), Praomys fumatus (0.85%), Xerus erythropus (0.85%), Lemniscomys barbarus (0.71%), Mus tenellus (0.71%) and Otomys typus (0.28%) were the rodents and their respective relative abundance in the study area. Crocidura olivieri (1.55%), Crocidura fumosa (0.85%), Crocidura bicolor (0.57%) and Elephantulus rufescens (0.28%) were the insectivores recorded with their respective relative abundance. Mastomys natalensis was the most abundant and O. typus and E. rufescens were the least (two each). Diversity of small mammals ranged from 2.299 to 2.625 with an average of 2.412. The highest small mammal diversity was in grasslands and the lowest was in Lake Chamo shore. Small mammal density varied from 5 to 43 ha^?1 and biomass varied from 244 to 2559 g ha^?1 with significant changes in relation to seasons and habitats.     

Habitat preferences,seasonal abundance and diets of rodents in Alage,Southern Ethiopia

To understand habitat preferences, seasonal abundance and diets of rodents in wet and dry season surveys were conducted in Alage, Southern Ethiopia. Sherman and snap traps were used to capture rodents from the four habitats: bushland, Acacia woodland, maize and wheat farmlands. A total of 3312 trap nights, from the four trapping habitats, yielded 776 individuals that represented 11 species of rodents. The distribution of rodents varied between habitats and seasons. Wet season rodent abundance was 52.3% while in the dry season it was 47.7%. Seasonal differences in species abundance were insignificant. Bushland habitat had high wet and dry season abundances with 137 and 211 individuals, respectively. Abundance was low in maize farm (57 individuals) in the wet season and wheat farm (10 individuals) in the dry season. Stomach content composition analysis of snap‐trapped rodents from different habitats showed differences between species and across seasons. Six rodent species were recorded as pests on the farmlands in this study area. In conclusion, variation in habitat preferences and diet of rodents in different habitats and across seasons might be due to the role of ground cover and food sources.     

Natural levels of fungal infections in grasshoppers in Northern Benin

The infection of grasshoppers by naturally occurring, entomopathogenic fungi was monitored at two sites in Malanville, northern Benin, Africa. Grasshoppers were collected and recorded from the sites between June and December 1992 and all of them, barring the first instars, were incubated in ventilated cages. At the first site, 1343 individuals of 35 grasshopper species were incubated, and at the second site, 857 individuals of 36 grasshopper species were incubated. Three hyphomycete fungi (Deuteromycotina: Hyphomycetes), Metarhizium flavoviride Gams and Rozsypal, Beauveria bassiana (Bals.) Vuillemin and Sorosporella sp. were found infecting grasshoppers. The average incidence of M. flavoviride infection was 2.9% and 1.8% at the two sites for all host species. M. flavoviride sporulated on most grasshopper cadavers within 10 days of collection. B. bassiana and Sorosporella sp. were only collected from one and five grasshopper individuals respectively. A significant difference was noted in the time to death of small grasshopper species infected with M. flavoviride compared to larger species. At one site, M. flavoviride infection was positively correlated with rainfall during the 10‐day period in which samples were taken.     

Foraging habitats of bats in southern Finland

We determined the foraging habitats of the northern batEptesicus nilssonii (Keyserling et Blasius, 1839), Brandt’s batMyotis brandtii (Eversmann, 1845), whiskered batMyotis mystacinus (Kuhl, 1819), Daubenton’s batMyotis daubentonii (Kuhl, 1819) and brown long-eared batPlecotus auritus (Linnaeus, 1758) in southern Finland. Among these species, we compared the diversities of foraging habitats, linear feature preference and the bats’ tendencies to forage simultaneously.Eptesicus nilssonii was the most opportunistic, foraging in a wide range of habitats.Myotis daubentonii (94%) foraged mainly on water habitats, whileM. brandtii/mystacinus (89%) andP. auritus (66%) foraged mainly in forest habitats. The diversities of foraging habitats used byE. nilssonii andP. auritus were higher than those ofM. brandtii/mystacinus andM. daubentonii. Both E.nilssonii andP. auritus foraged mostly alone or in small numbers, whileM. brandtii/mystacinus tended to gather in large numbers to forage in the same habitat. Half ofE. nilssonii and 46% ofM. daubentonii foraged over linear features, while other species did not use linear features to such an extent. Management and conservation of foraging habitats are needed especially forM. brandtii/mystacinus andM. daubentonii, which are more specialized thanE. nilssonii and P. auritus.     

Species composition and community structure of small pest rodents (Muridae) in cultivated and fallow fields in maize‐growing areas in Mayuge district,Eastern Uganda

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Resource partitioning confirmed by isotopic signatures allows small mammals to share seasonally flooded meadows

Meadows in river deltas are characterized by a high diversity and abundance of small mammals. However, neither their spatial arrangement nor differences in their use of microhabitat can necessarily explain the dense co‐occurrence of sympatric species. We investigated how several small mammal species share a seasonally flooded meadow of limited size, testing predictions (P1) that herbivore, granivore, insectivore, and omnivore species are separated in time (dominant in different years), (P2) that sympatric species undergo isotopic partitioning, and (P3) that there are intraspecific differences in diet. Stable carbon and nitrogen isotope signatures in the hair of seven synantropic shrew, vole, and mice species were used as a proxy for their diet. We found that the three most abundant species in eight of the nine years were from different diet groups. However, based on the number of species in the functional groups, the state of small mammal community was considered unfavored in five out of the nine investigation years. In years with the greatest dominance of Apodemus agrarius, the small mammal community was characterized by decreased diversity and Micromys minutus was either in low abundance or absent. In 2014 and 2016, years of low abundance or absence of M. oeconomus, M. agrestis, and M. glareolus were both recorded in high numbers. Differences in the isotopic signatures of the three most abundant small mammal species in the community were clearly expressed and core areas in the isotopic space were separated, showing their dependence on different dietary resources. Intraspecific dietary separation between young and adult animals was observed only in M. oeconomus. Thus, the high species diversity of small mammals and the formation of their community in this investigated flooded meadow are maintained by isotopic partitioning (segregation in dietary space) and by changes in their number over time (shifting dominance).     

Vulnerability of small and medium-sized prey mammals in relation to their habitat preferences,age classes and locomotion types in the temperate Monte Desert,Argentina

In this study we examined predation by small felids upon small and medium-sized prey mammals in the central region of the Monte Desert. We analyzed the degree of vulnerability of prey mammals in relation to their habitat use patterns, modes of locomotion and age classes. Medium and small rodents were the predominant prey in the diet of small cats. The most consumed small mammal was Akodon molinae, which inhabits areas of dense cover, while species occurring in open habitats such as Eligmodontia typus were less consumed. Among medium-sized mammals, Galea leucoblephara that depends on patches with high plant cover for refuge was more consumed than Microcavia australis that lives in colonies and displays a complex set of antipredator strategies. Prey selectivity on small rodents showed that E. typus, which uses bipedal locomotion and an erratic escape behaviour, was consumed in greater proportion than its availability. We suggest that although its specialized locomotory mode would diminish predator attacks, its preference for open habitats would increase the probability of predator–prey encounter. Consumption of small mammals focuses mostly on adult individuals. However, we found a significant consumption of senile individuals of A. molinae, which points out that vulnerability tends to be high towards the age of maturity in this species.     

Diversity and structure of ant communities associated with oak,pine, eucalyptus and arable habitats in Portugal

Summary In central Portugal, 28 species of ants were found in oak, 25 in pine, 35 in eucalyptus and 12 in arable habitats, a total of 43 species. They include five numerically dominant species, the exotic Argentine ant.Linepithema (Iridomyrmex) humile and four native species,Crematogaster scutellaris, Pheidole pallidula, Tapinoma nigerrimum andTetramorium hispanicum. L. humile occurred in 34% of sampled sites. When present, it was abundant in all three arboreal habitats, where it was associated with strikingly decreased ant species richness and equitability. Such diversity was greater inL. humile-colonised arable habitats, where the ant was much less abundant than in the arboreal habitats.WhereL. humile was absent,C. scutellaris andP. pallidula were usually co-dominants in oak and pine, andT. nigerrimum andT. hispanicum in the arable habitat, but all were absent or rare in eucalyptus. Ants were numerically least abundant in eucalyptus but, in the absence ofL. humile, species richness was greater and the community more equitable than in the other habitats.L. humile-absent arable habitats supported the poorest and least equitable ant community.     

Guinean biodiversity at the edge: Rodents in forest patches of southern Mali

Southern Mali mainly belongs to the Sudanian savanna bioclimatic zone, but forest patches showing botanical affinities with Guinean humid forest remain as gallery forests or ravine forests. To characterize the rodent diversity of this area and check for the presence of some species of Guinean affinities in this group, rodent assemblages were sampled in four regions of southern Mali, using trapping and observational data in forest and surrounding habitats. Twenty-four species were recorded, comprising a representative sample of the expected overall diversity in this group according to rarefaction curves. Praomys rostratus was the dominant species in the most humid, closed lowland forest. Praomys daltoni was also present in this habitat type, being all the more abundant as habitat degradation was apparent. It became the dominant species in ravine forest on rocky substrate where P. rostratus was completely absent. In Sudanian savanna habitats and in herbaceous and cultivated areas, Mastomys erythroleucus dominated a diverse rodent community. A few species were found that testified for Guinean affinities of the most humid forest patches, especially in the extreme southeast of the country (region of Sikasso). Rodent assemblages of the Bafing and Mts Mandingues areas, in the western part of the study area, showed the highest similarity, in relation with environmental characteristics of this region representing an extension of the Fouta Djallon plateau in Guinea. The results obtained highlight the high biodiversity value of this forest-savanna mosaic, and provide new arguments in favour of the preservation of West African forest patches and their surrounding habitats.     

Distribution of fungi on two mite species and their habitats in Egypt

Fungi associated with two mite species,Acotyledon krameri andTyrophagus putrescentiœ, and their habitats were studied using 1% glucose-Czapek's and potato-dextrose agar media at 28±2°C. A total of 54 species and one variety belonging to 25 genera were recovered from different habitats of mite species; 36 species and one variety belonging to 18 genera were associated with mite bodies. Nearly most fungal species isolated from mite bodies were also encountered in their habitats. The most common species wereAspergillus flavus, A. fumigatus, A. niger, Mucor racemosus, Nectria hœmatococca. A. terreus, Scopulariopsis brevicaulis andTrichurus spiralis were isolated only from mite bodies whereasFusarium oxysporum, Penicillium chrysogenum, P. corylophilum, P. funiculosum andRhizopus stolonifer were recovered from the mite habitats.A. krameri individuals survived well onCunninghamella elegans, F. oxysporum andM. racemosus cultured on both type of media, whereasCylindrocarpon destructans was best forT. putrescentiœ survival.     

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.