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.     

State‐space modeling reveals habitat perception of a small terrestrial mammal in a fragmented landscape

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Factors determining mammal species richness on habitat islands and isolates: habitat diversity, disturbance, species interactions and guild assembly rules

• 1 For over three decades the equilibrium theory of island biogeography has galvanized studies in ecological biogeography. Studies of oceanic islands and of natural habitat islands share some similarities to continental studies, particularly in developed regions where habitat fragmentation results from many land uses. Increasingly, remnant habitat is in the form of isolates created by the clearing and destruction of natural areas. Future evolution of a theory to predict patterns of species abundance may well come from the application of island biogeography to habitat fragments or isolates.
• 2 In this paper we consider four factors other than area and isolation that influence the number and type of mammal species coexisting in one place: habitat diversity, habitat disturbance, species interactions and guild assembly rules. In all examples our data derive from mainland habitat, fragmented to differing degrees, with different levels of isolation.
• 3 Habitat diversity is seen to be a good predictor of species richness. Increased levels of disturbance produce a relatively greater decrease in species richness on smaller than on larger isolates. Species interactions in the recolonization of highly disturbed sites, such as regenerating mined sites, is analogous to island colonization. Species replacement sequences in secondary successions indicate not just how many, but which species are included. Lastly, the complement of species established on islands, or in insular habitats, may be governed by guild assembly rules. These contributions may assist in taking a renewed theory into the new millennium.

     

The influence of behaviour and season on habitat selection by a small mammal

This study examined the selection of habitat by Ningaui yvonneae , a small, nocturnal insectivorous marsupial of semi-arid regions of southern Australia. In addition to idengifying habitat preferences, the study was concerned with idengifying how habitats were used, and for what purpose they were valued. Triodia irritans (hummock grass) was the most preferred habitat component overall. However, other habitat components were found to be of equal or greater preference during certain seasons and for certain behaviours. The importance of considering behaviour in studies of habitat selection was discussed. Habitat selection by N. yvonneae appeared to be influenced by predation risk and energetic reward. The selection for Triodia is thought to be due to its provision of protection and foraging opportunities. However, leaf litter was preferred over Triodia for foraging, possibly because it offered greater energetic returns. To minimise predation risk while in leaf litter, ningauis remained close to Triodia and moved larger prey items to the edge of Triodia for consumption.     

Linking small desert mammal distribution to habitat structure in a protected and grazed landscape of the Monte, Argentina

Interpretation of landscape patterns from the perspective of different species allows knowing the way in which they perceive landscape, and how their perception varies with scale. We examined distribution of four small mammal species at different scales over a landscape including protected and grazed areas, and associated species distribution with landscape structure. The study was conducted in the central Monte Desert (Reserve of Ñacuñán). Trap grids were set in both areas at two scales, varying their grain and extent. To determine whether spatial patterns are random, clumped or regular, we used a point pattern analysis. Logistic regressions were performed to relate the presence-absence of small mammals to environmental variables. Intensity of the point pattern was not constant, either in the Reserve or the grazed area. Small mammal abundance exhibited a heterogeneous distribution, and the existence of a first-order effect was detected for all species. No second-order effects were detected, the point pattern was random for all species in both areas. Both areas were differently perceived by rodent species. Habitat structure in both conditions and its variations with scale appear to be important factors affecting distribution patterns.     

Untangling the roles of fire,grazing and rainfall on small mammal communities in grassland ecosystems

In grassland systems across the globe, ecologists have been attempting to understand the complex role of fire, grazing and rainfall in creating habitat heterogeneity and the consequences of anthropogenic control of these factors on ecosystem integrity and functioning. Using a South African grassland ecosystem as a model, we investigated the impact of fire and grazing pressure on small mammal communities during three differing periods of a rainfall cycle. Over 2 years, 15,203 trap nights revealed 1598 captures of 11 species (nine rodents, one macroscelid and one insectivore). Results highlighted the importance of the interplay between factors and showed that the role of fire, grazing and rainfall in determining small mammal abundance was species-dependant. While no two species were affected by the same environmental variables, grass cover or height was important to 56% of species. Considered independently, high rainfall had a positive influence on small mammal abundance and diversity, although the lag period in population response was species-specific. High grazing negatively affected overall abundance, but specifically in Mastomys coucha; fire alone had little immediate impact on small mammal diversity. Six months after the fire, vegetation cover had recovered to similar levels as unburned areas, although small mammal diversity and richness were higher in burned areas than unburned areas. Grazing levels influenced the rate of vegetation recovery. In conclusion, low-level grazing and burning can help to maintain small mammal biodiversity, if conducted under appropriate rainfall levels. A too high grazing pressure, combined with fire, and/or fire conducted under drought conditions can have a negative impact on small mammal biodiversity. To maintain small mammal diversity in grassland ecosystems, the combined effects of the previous year’s rainfall and existing population level as well as the inhibition of vegetation recovery via grazing pressure need to be taken into consideration before fire management is applied. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Assessment of habitat quality for four small mammal species of the Monte Desert, Argentina

In the temperate desert of Argentina, the combined action of climatic and anthropogenic factors has contributed to the formation of a highly heterogeneous landscape. In the central region of the Monte desert, four small mammal species (Eligmodontia typus, Calomys musculinus, Akodon molinae and Graomys griseoflavus) coexist and show different habitat uses in response to spatial variability. Three main habitat types are present in the region: mesquite forest, the creosotebush community and sand dunes. These habitat types are present also in the surrounding grazing area.The objective of this study was to determine habitat quality for these species in a protected area (Reserve MaB Ñacuñán) and in the adjacent grazed area. For each species we estimated demographic parameters that are highly correlated to fitness in each habitat, and for both treatments (protected and grazed).We found that the protected area offered a higher quality habitat than the grazed area for all species, but principally A. molinae and G. griseoflavus. At a local scale, we found that A. molinae and C. musculinus clearly showed higher fitness in the more complex habitats as the creosotebush community and the mesquite forest. In contrast, for E. typus, open and simplest patches, such as sand dunes, were optimal for its survival and reproduction.     

Fire,landscape change and models of small mammal habitat suitability at multiple spatial scales

Fire is an important process in many ecosystems, but inappropriate fire regimes can adversely affect biodiversity. We identified a naturally flammable heathy woodland ecosystem where the use of planned fire had increased the extent of older vegetation, and quantified the abundance of two small native mammals in this landscape (silky mouse Pseudomys apodemoides and heath rat P. shortridgei). We defined four time‐since‐fire (TSF) categories representing a 2‐ to 55‐year post‐fire sequence and, on the basis of a habitat accommodation model, predicted that both species would select younger age‐classes over older ones. We also predicted that (i) much of the variance in vegetation structure would remain unexplained by TSF and (ii) statistical models of mammal abundance and occupancy including structural variables as predictors would be better than models including TSF. Pseudomys apodemoides selected 17‐ to 23‐year‐old sites, while there was no evidence that P. shortridgei selected a particular TSF category, findings that were inconsistent with our predictions. In line with our predictions, relatively large portions of the variance in vegetation structure remained unexplained by TSF (adjustedr² for four structural variables: 0.24, 0.29, 0.35 and 0.57), and in three of four cases there was strong evidence that statistical models of mammal abundance and occupancy including structural variables were better than those including TSF. At the site scale (hectares), P. shortridgei abundance was positively related to the cover of dead material at the base of Xanthorrhoea plants and at the trap scale (metres), the trapability of both species was significantly related to vegetation volume at 0–20 cm. Our findings suggest that TSF may not be a good proxy for either vegetation structure or species abundance/occupancy.     

Marten space use and habitat selection in managed coniferous boreal forests of eastern Canada

Effects of habitat loss and fragmentation on the behavior of individual organisms may have direct consequences on population viability in altered forest ecosystems. The American marten (Martes americana) is a forest specialist considered as one of the most sensitive species to human-induced disturbances. As some studies have shown that martens cannot tolerate >30–40% clear-cuts within their home range, we investigated marten space use (home range size and overlap) and habitat selection in landscapes fragmented by 2 different patterns of timber harvesting in the black spruce boreal forest: dispersed-cut landscapes (10–80 ha cut-blocks) and clustered-cut landscapes (50–200 ha cut-blocks). We installed radio-collars on female martens and determined 20 winter home ranges (100% minimum convex polygons and 60–90% kernels) in dispersed-cut (n = 8) and clustered-cut (n = 12) landscapes. Home range size was not related to the proportion of clear-cuts (i.e., habitat loss), but rather to the proportion of mixedwood stands 70–120 years old. However, female body condition was correlated to habitat condition inside their home ranges (i.e., amount of residual forest and recent clear-cuts). At the home range scale, we determined that mixedwood forests were also among the most used forest stands and the least used were recent clear-cuts and forested bogs, using resource selection functions. At the landscape scale, home ranges included more mixedwood forests than random polygons and marten high activity zones were composed of more residual forest and less human-induced disturbances (clear-cuts, edges, and roads). These results suggest that mixedwood forests, which occupy approximately 10% of the study area, play a critical role for martens in this conifer-dominated boreal landscape. We recommend permanent retention or special management considerations for these isolated stands, as harvesting mixedwood often leads to forest composition conversion that would reduce the availability of this highly used habitat. © The Wildlife Society, 2013     

Benefits of habitat restoration to small mammal diversity and abundance in a pastoral agricultural landscape in mid-Wales

Changes in agricultural practice are predicted across the UK following agricultural reform driven by government policy. The suitability of agri-environment schemes for many species is currently debated because limited quantitative data are collected. In order to understand the changes to biodiversity due to agri-environment schemes, there is a need for studies to not just compare biodiversity and species composition in and out of agri-environment areas, but to factor in the influence of temporal habitat changes. In this study, we investigate the suitability of an agri-environment initiative to support and enhance a small mammal fauna among pastoral hill farms in mid-Wales. Grazed and ungrazed woodlands, riparian habitats, and broadleaf plantations, were compared for small mammal abundance and diversity following a trapping study. Mammal diversity was similar across habitats, though abundance varied significantly. A principle component analysis identified that mammal abundance clustered into three main habitat groups separated by seral stage (early, mid, late). No relationship between mammal abundance and stock grazing was found. A canonical correspondence analysis confirmed that vegetation structure was important in explaining the distribution of captures of mammal species across the landscape. The results for habitat type, and habitat context, suggest that a mix of vegetation seral stages, reflecting a varied vegetation structure, is important to maintain small mammal diversity and abundance across the study area. Heterogeneity in structural diversity at the landscape scale is important to maintain a variety of ground-dwelling mammal species, and particularly because trends in countryside surveys show that woodlands are skewed towards late seral stages. Habitat heterogeneity can be maintained because the hill farms neighbour each other, and the farmers co-operate as a group to manage the landscape. Habitat diversity is therefore possible. These results help us to advocate, and anticipate, the benefits of groups of farms within a landscape.     

Results of a food addition experiment in a north-central Chile small mammal assemblage: evidence for the role of "bottom-up" factors

Since 1989, effects of biotic interactions including predation and herbivory have been examined in a replicated experimental study in a north‐central Chilean semiarid thorn scrub community. Strong responses of small mammals and plants to El Niño Southern Oscillations (ENSO) have also been documented suggesting that “bottom‐up” factors related to high rainfall are important. To simulate increased primary productivity effects on small mammals, ad lib rabbit pellet additions were initiated in mid‐1997 on unfenced grids near the experimental complex. Following the 1997 El Niño event with three times normal precipitation, numbers of small mammals during pre‐addition months and the first treatment year were similar on control and food addition grids. During the second year (1998–1999), a period of severe drought, food additions had significant positive effects on numbers of two predominantly herbivorous “core” (resident) species, Octodon degus and Phyllotis darwini, and an omnivorous “quasi‐core” (resident but highly fluctuating) species, Akodon olivaceus; however, all three species declined towards the end of the second treatment year. Two “opportunistic” (temporarily resident) species, Abrothrix longipilis (an insectivore) and Oligoryzomys longicaudatus (a granivore), showed no responses to food additions. An insectivorous marsupial, Thylamys elegans (also a “core species”), had significantly lower numbers on food addition grids. Changes in body weight distributions and proportions of reproductive individuals particularly in O. degus indicate in situ responses. Whereas no differences in residency, numbers of stations visited, and trappability were observed, energy compensation ratios greater than one suggest significant immigration in the second year. Thus, food additions elicited strong responses by herbivorous/omnivorous “core” and “quasi‐core species” whereas they had no effects on “opportunistic species”. These results reinforce the view that “bottom‐up” factors influencing food availability exert prevailing control on numerically important small mammal species by temporarily increasing carrying capacity, and that “top‐down” factors (i.e., biotic interactions) become important when small mammal numbers are at or near their carrying capacity. Spatial dynamics may be important in explaining declines of species populations exhibiting initially positive responses to food additions.     

Environmental determinants of the small mammal assemblage in an agroecosystem of central Argentina: The role of Calomys musculinus

Agricultural intensification in the central region of Argentina has been accompanied by rodent assemblage changes, involving an increase in Calomys species densities. In particular, Calomys musculinus, the main reservoir of Junín virus (etiological agent of the Argentine Haemorrhagic Fever, AHF) selects field borders over crop-fields. Borders represent relatively-stable habitats within agroecosystems, otherwise highly human modified landscapes. In this article, we assessed the effect of environmental variables on rodent species occurrence and assemblage in the field borders of a pampean agroecosystem. Small rodent occurrence was examined for variation according to vegetation species composition, structure and productivity, using canonical ordination techniques. In a total of 72 crop-field borders of Córdoba province, in spring, summer and autumn (2005-2006), we captured 1041 rodents of 8 different species. Akodon azarae, Calomys venustus and C. musculinus were the dominant species of the assemblage, with the first two associated one another, while C. musculinus tended to be related to different environmental variables. We showed that rodent species associations were mediated by resources that changed seasonally, such as vegetation cover. Also, the strength and “sign” of the association between species seemed to be a consequence of different habitat preferences and activity patterns. Finally, the intra-annual population cycle seemed to influence the relation between species and habitat structure and, possibly, inter-specific competition for resources. Therefore, we suggest that less generalist species of the rodent assemblage, well adapted to linear habitats, may limit C. musculinus occurrence throughout the agroecosystem by habitat and spatial competition. Any action aimed to control AHF, and therefore C. musculinus population density, should exclude negative effects on coexisting species.     

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.     

Use of arboreal and terrestrial space by a small mammal community in a tropical rain forest in Borneo, Malaysia

Aim Small mammals were live‐trapped in a primary rain forest to evaluate the relative distribution of species to each other and to microhabitat properties on the ground and in the canopy. Location Kinabalu National Park in Borneo, Sabah, Malaysia. Methods Seven trapping sessions were conducted along two grids with 31 trap points at distances of 20 m on the ground and in the lower canopy at an average height of 13.5 m. Results Species diversity and abundance of small mammals proved to be high: 20 species of the families Muridae, Sciuridae, Tupaiidae, Hystricidae, Viverridae and Lorisidae were trapped, with murids being dominant in both habitat layers. The terrestrial community was significantly more diverse with 16 captured species (Shannon–Wiener's diversity index = 2.47), while 11 species were trapped in the canopy ( = 1.59). The Whitehead's rat, Maxomys whiteheadi, and the red spiny rat, Maxomys surifer, dominated the terrestrial community whereas the large pencil‐tailed tree mouse, Chiropodomys major, was by far the most abundant species in the canopy. Other abundant species of the canopy community, the dark‐tailed tree rat, Niviventer cremoriventer, and the lesser treeshrew, Tupaia minor, were also abundant on the ground, and there was no clear boundary between arboreal and terrestrial species occurrences. Main conclusions As most species were not confined to specific microhabitats or habitat layers, species seemed to rely on resources not necessarily restricted to certain microhabitats or habitat layers, and separation of species probably resulted mainly from a species’ concentrated activity in a preferred microhabitat rather than from principal adaptations to certain habitats. Ecological segregation was stronger in the more diverse terrestrial community, though microhabitat selection was generally not sufficient to explain the co‐occurrences of species and the variability between local species assemblages. Constraints on small mammal foraging efficiency in the three‐dimensional more complex canopy may be responsible for the similarity of microhabitat use of all common arboreal species. Community composition was characterized by mobile species with low persistence rates, resulting in a high degree of variability in local species assemblages with similar turnover rates in both habitats.     

Behavioral states in space and time: understanding landscape use by an invasive mammal

Animal movement models can be used to understand species behavior and assist with implementation of management activities. We explored behavioral states of an invasive wild pig (Sus scrofa) population that recently colonized central Michigan, USA, 2014–2018. To quantify environmental factors related to wild pig movement ecology and spatio-temporal landscape use, we predicted wild pig behavioral states relative to land cover type, landscape structure (i.e., edge and patch cohesion), and weather conditions. We used global positioning system (GPS)-collars and monitored 8 wild pigs from 2014–2018. We fit local convex hulls and calculated movement metrics revealing 3 wild pig behavioral states (resting, exploratory, and relocating) and constructed a 3-level model to predict behavioral state probabilities relative to biotic and abiotic conditions. Probabilities of exploratory and resting behaviors were higher nearer to riparian and open herbaceous cover types (oftentimes emergent marsh), indicating that these cover types provided security cover during activity and bedding. Hard mast cover types had a strong positive association with relocating behaviors. More cohesive patches of agriculture and shrub cover types were associated with higher probabilities of exploratory behaviors, while resting was more likely in continuous patches of agriculture (mostly mid-summer corn). The probability of exploratory behaviors increased exponentially with warming ambient temperature. Our results may be used by managers to develop control strategies conducive to landscape and environmental conditions where the likelihood of encountering wild pigs is highest or targeting wild pigs when in a behavioral state most vulnerable to a particular removal technique.     

Determinism in a transient assemblage: the roles of dispersal and local competition

Both dispersal and local competitive ability may determine the outcome of competition among species that cannot coexist locally. I develop a spatially implicit model of two-species competition at a small spatial scale. The model predicts the relative fitness of two competitors based on local reproductive rates and regional dispersal rates in the context of the number, size, and extinction probability of habitat patches in the landscape. I test the predictions of this model experimentally using two genotypes of the bacteriophagous soil nematode Caenorhabditis elegans in patchy microcosms. One genotype has higher fecundity while the other is a better disperser. With such a fecundity-dispersal trade-off between competitors, the model predicts that relative fitness will be affected most by local population size when patches do not go extinct and by the number of patches when there is a high probability of patch extinction. The microcosm experiments support the model predictions. Both approaches suggest that competitive dominance in a patchily distributed transient assemblage will depend upon the architecture and predictability of the environment. These mechanisms, operating at a small scale with high spatial admixture, may be embedded in a larger metacommunity process.     

Effects of land use,habitat characteristics,and small mammal community composition on Leptospira prevalence in northeast Madagascar

Human activities can increase or decrease risks of acquiring a zoonotic disease, notably by affecting the composition and abundance of hosts. This study investigated the links between land use and infectious disease risk in northeast Madagascar, where human subsistence activities and population growth are encroaching on native habitats and the associated biota. We collected new data on pathogenic Leptospira, which are bacteria maintained in small mammal reservoirs. Transmission can occur through close contact, but most frequently through indirect contact with water contaminated by the urine of infected hosts. The probability of infection and prevalence was compared across a gradient of natural moist evergreen forest, nearby forest fragments, flooded rice and other types of agricultural fields, and in homes in a rural village. Using these data, we tested specific hypotheses for how land use alters ecological communities and influences disease transmission. The relative abundance and proportion of exotic species was highest in the anthropogenic habitats, while the relative abundance of native species was highest in the forested habitats. Prevalence of Leptospira was significantly higher in introduced compared to endemic species. Lastly, the probability of infection with Leptospira was highest in introduced small mammal species, and lower in forest fragments compared to other habitat types. Our results highlight how human land use affects the small mammal community composition and in turn disease dynamics. Introduced species likely transmit Leptospira to native species where they co-occur, and may displace the Leptospira species naturally occurring in Madagascar. The frequent spatial overlap of people and introduced species likely also has consequences for public health.     

Predator effects on aquatic community assembly: disentangling the roles of habitat selection and post-colonization processes

Top predators are known to play an important role in the assembly of communities via two mechanisms: (1) by altering the colonization (or emigration) patterns of prey through behavioral habitat selection, and (2) by altering vital rates (e.g. mortality, birth) of prey after colonization. While both these mechanisms act to determine assembly, research has focused on either their combined overall effects (confounding them), or examined them singly. As a result, it remains unclear how these mechanisms act to sequentially shape community structure. In this study, we experimentally disentangle habitat selection and post‐colonization effects of predaceous fish to test their independent and combined influence on the assembly of insect and larval amphibian communities in experimental freshwater habitats. Specifically, we ask, ‘do the behavioral choices of colonists continue to structure aquatic communities even after post‐colonization processes have occurred?’ Like previous studies, we found that colonization was strongly reduced by the presence of fish cues. More importantly, these effects of fish on prey colonization behavior combined independently with post‐colonization processes to determine the overall effect of predators on community assembly. Although habitat selection and predation both reduced abundance and biomass of most taxa in the post‐colonization communities, these factors had qualitatively different effects on aspects of trophic structure. Habitat selection altered the ratio of secondary to primary consumer abundance and biomass, while post‐colonization predation drove strong trophic cascades not observed in response to habitat selection. Our results suggest that behavioral choices regarding habitat selection can have lasting and unique effects on the structure of aquatic communities.