Road verges as habitat for small mammals in Britain

There are 370 000 kilometres of roads in Great Britain, mostly bordered by a verge that is potential habitat for small mammals. The present study assessed the importance of road verges as small mammal habitat and investigated the influence of some key features on rodent abundance. Five rodent and three shrew species were live‐trapped on 14 road verges in late summer 1994 and nine verges in autumn 1996 in north Cambridgeshire, UK. On average, between three and four species were captured per verge. Bank Voles Clethrionomys glareolus, Wood Mice Apodemus sylvaticus and Field Voles Microtus agrestis were the most abundant species, with mean densities of 45.5, 40.2 and 29.5 animals km^–1 in summer and 52.8, 181.9 and 47.2 animals km^–1 in autumn. Numbers varied between verges and this was significantly correlated with particular features on the verge. Bank Vole and Field Vole numbers showed a significant positive correlation with the dimensions of hedges and the width of the tall grass area, respectively. Wood Mice were also more numerous on verges with big hedges but the relationship between mouse abundance and verge structure was complex. The number of mice in 1994 was positively and significantly correlated with hedge features and with the width of the short grass sightline, whereas, in autumn 1996, they were only significantly correlated with total verge width (positive association) and ditch width (negative association).     

Controlled experiments of habitat fragmentation: a simple computer simulation and a test using small mammals

Habitat fragmentation involves a reduction in the effective area available to a population and the imposition of hard patch edges. Studies seeking to measure effects of habitat fragmentation have compared populations in fragments of different size to estimate and area effect but few have examined the effect of converting open populations to closed ones (an effect of edges). To do so requires a shift in spatial scope-from comparison of individual fragments to that of fragmented versus unfragmented landscapes. Here we note that large-scale, controlled studies of habitat fragmentation have rarely been performed and are needed. In making our case we develop a simple computer simulation model based on how individual animals with home ranges are affected by the imposition of habitat edges, and use it to predict population-level responses to habitat fragmentation. We then compare predictions of the model with results from a field experiment on Peromyscus and Microtus. Our model treats the case where home ranges/territories fall entirely within or partially overlap with that of sample areas in continuous landscapes, but are restricted to areas within habitat fragments in impacted landscapes. Results of the simulations demonstrate that the imposition of hard edges can produce different population abundances for similar-sized areas in continuous and fragmented landscapes. This edge effect is disproportionately greater in small than large fragments and for species with larger than smaller home ranges. These predictions were generally supported by our field experiment. We argue that large-scale studies of habitat fragmentation are sorely needed, and that control-experiment contrasts of fragmented and unfragmented microlandscapes provide a logical starting point.     

Highway verges as habitat providers for small mammals in agrosilvopastoral environments

The Mediterranean Basin has an important conservation value given its high biodiversity and high number of endemic species, which have co-existed with human traditional practices for centuries. However, northern areas as the Iberian Peninsula have experienced intensification in livestock production in recent past, with consequent reduction in habitat quality. In this study we assessed the importance of fenced highway verges as habitat for small mammals in Mediterranean agrosilvopastoral landscapes. More specifically, we compared small mammal abundance between highway verges and the adjacent two main land uses (“montado” and open areas); compared the vegetative structure among these land uses; and addressed how vegetation structure influences species occupancy. Thirty-six sites were sampled in agrosilvopastoral system areas in southern Portugal (sampling effort 8,840 trap-nights). A total of 351 individuals from target species were captured: 157 wood mice (Apodemus sylvaticus), 95 western Mediterranean mice (Mus spretus) and 99 greater white-toothed shrews (Crocidura russula). Capture–mark–recapture analyses were performed to estimate population size. Our data suggests that fenced highway verges promote better vegetative structure conditions which in turn favor a higher animal abundance therein. We suggest the adoption of management practices to increase the height and cover of herbaceous and shrub layers in road verges, together with creating grazing controlled areas in highway vicinity, particularly in “montado” patches, linked by vegetated linear features. This would increase habitat and refuge for a large numbers of species, including small mammals, and thus benefiting the trophic chain and the whole agrosilvopastoral system.     

Motor vehicles as vectors of plant species from road verges in a suburban environment

The objective of this study was to assess the potential contribution of car-borne flora to species dispersal in a suburban area in Flanders (Belgium). Therefore, motor vehicle-borne flora was evaluated monthly for a random sample of 20 cars between October 1988 and September 1989. At the end of the experiment 33 car-borne plant species, with a total of 690 seedlings were identified from mud, attached to these vehicles. There was considerable temporal variation in the mud mass attached to the vehicles and therefore also the size and composition of the vehicle-borne flora varied during the year. Because mud retention on cars is a crucial factor in seed dispersal, the number of emerging seedlings is negatively correlated with precipitation in the week preceding the sampling. The flora dispersed by vehicles consists mostly of species with small, light seeds and a persistent seed bank. Usually, they are pioneer species of nitrogen-rich habitats. The nature of the vehicle dispersed flora was compared with the local flora of road verges and differed significantly from this in, e.g., seed length, seed weight, ecological groups, Ellenberg indicator values for nitrogen supply, potential dispersal strategies and rarity. A larger resemblance with the flora of the zone immediately adjacent to the carriageway was found. We only found a low similarity to other motor vehicle-borne floras.     

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.     

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

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Non-native plantation forests as alternative habitat for native forest beetles in a heavily modified landscape

The once extensive native forests of New Zealand’s central North Island are heavily fragmented, and the scattered remnants are now surrounded by a matrix of exotic pastoral grasslands and Pinus radiata plantation forests. The importance of these exotic habitats for native biodiversity is poorly understood. This study examines the utilisation of exotic plantation forests by native beetles in a heavily modified landscape. The diversity of selected beetle taxa was compared at multiple distances across edge gradients between each of the six possible combinations of adjacent pastoral, plantation, clearfell and native forest land-use types. Estimated species richness (Michaelis–Menten) was greater in production habitats than native forest; however this was largely due to the absence of exotic species in native forest. Beetle relative abundance was highest in clearfell-harvested areas, mainly due to colonisation by open-habitat, disturbance-adapted species. More importantly, though, of all the non-native habitats sampled, beetle species composition in mature P. radiata was most similar to native forest. Understanding the influence of key environmental factors and stand level management is important for enhancing biodiversity values within the landscape. Native habitat proximity was the most significant environmental correlate of beetle community composition, highlighting the importance of retaining native remnants within plantation landscapes. The proportion of exotic beetles was consistently low in mature plantation stands, however it increased in pasture sites at increasing distances from native forest. These results suggest that exotic plantation forests may provide important alternative habitat for native forest beetles in landscapes with a low proportion of native forest cover.     

Effects of forest fragmentation on small mammals in an Atlantic Forest landscape

Recent studies on the effects of tropical forest fragmentation indicate that fragmented landscapes are complex and heterogeneous systems influenced by factors other than the size or degree of isolation of forest remnants: of particular importance are the quality of the matrix and the edge-induced habitat changes. In order to investigate the influence of these factors, small mammals were surveyed in 36 sites in the landscape of Una, a region that encompasses some of the last and largest Atlantic Forest remnants in northeastern Brazil. Six sites were distributed on each of six landscape components – the interiors and edges of small remnants, the interiors and edges of large remnants, and the most common forested habitats found in the matrix. The survey comprised 46,656 trap-nights and yielded 1725 individuals of 20 species of rodents and marsupials. Results revealed: an increase in beta-diversity caused by fragmentation; the contrasting effects of the altered forested habitats of the matrix, which harbor both forest and disturbance-adapted species; a greater importance of edge effect than of patch size to the observed changes in small mammal community in remnants; an association among terrestrial forest species and among arboreal forest species in terms of the distribution and abundance in the Una mosaic; and a distinctive vulnerability of these two groups of species to fragmentation. Results emphasize the biological importance and conservation value of both fragmented landscapes and small remnants in the Atlantic Forest, as well as the importance of management techniques to control and attenuate edge effects.     

Metacommunities of spiders in grassland habitat fragments of an agricultural landscape

Arthropod communities in fragmented agricultural landscapes depend on local processes and the interactions between communities in the habitat islands. We aimed to study metacommunity structure of spiders, a group that is known for high dispersal power, local niche partitioning and for engaging in species interactions. While living in fragmented habitats could lead to nestedness, other ecological traits of spiders might equally lead to patterns dominated either by species interactions or habitat filtering. We asked, which community pattern will prevail in a typical agricultural landscape with isolated patches of semi-natural habitats. Such a situation was studied by sampling spiders in 28 grassland locations in a Hungarian agricultural landscape. We used the elements of metacommunity structure (EMS) framework to distinguish between alternative patterns that reveal community organization. The EMS analysis indicated coherent species ranges, high turnover and boundary clumping, suggesting Clementsian community organization. The greatest variation in species composition was explained by local habitat characteristics, indicating habitat filtering. The influence of dispersal could be detected by the significant effect of landscape composition, which was strongest at 500 m. We conclude that dispersal allows spiders to respond coherently to the environment, creating similar communities in similar habitats. Consistent habitat differences, such as species rich versus species poor vegetation, lead to recognisably different, recurrent communities. These characteristics make spiders a predictable and diverse source of natural enemies in agricultural landscapes. Sensitivity to habitat composition at medium distances warns us that landscape homogenization may alter these metacommunity processes.     

Impacts of habitat on butterfly dispersal in tropical forests,parks and grassland patches embedded in an urban landscape

Dispersal distances of 17 species of butterflies in tropical Singapore were significantly greater in forest than in urban habitat. Butterflies in urban plots frequently moved within suitable habitat (park/grassland) patches but rarely crossed non-habitat patches suggesting potential isolation and a need for urban corridors.     

Cape ground squirrels as ecosystem engineers: modifying habitat for plants,small mammals and beetles in Namib Desert grasslands

Burrowing and foraging of semi‐fossorial rodents can affect species distribution and composition. Ground squirrels dig large burrow systems for refuge from predators and temperature extremes. Burrowing and foraging around burrows by squirrels may affect habitat and resource distributions for other organisms. We examined the impact of Cape ground squirrels (Xerus inauris) on vegetation, small mammals and beetles during winter and summer in grasslands on the edge of the Namib Desert. At each burrow system and paired control site without burrows, we estimated plant cover and height using quadrats (N = 8 paired sites), small mammal abundance and species richness using mark‐recapture techniques (N = 8 paired sites) and beetle abundance and species richness using pitfall traps (N = 6 paired sites, winter only). Squirrel burrowing and foraging activities resulted in lower plant cover and height, higher small mammal abundance and lower beetle abundance and species richness. Squirrels also reduced more plant cover in winter compared to summer, but had no effect on small mammal species richness. Furthermore, plant cover and height were higher in summer, whereas small mammal abundance and species richness were higher in winter. Our results suggest that Cape ground squirrels are important ecosystem engineers that influence plant and animal communities in the Namib Desert grasslands.     

The role of small secondary biotopes in a highly fragmented landscape as habitat and connectivity providers for dragonflies (Insecta: Odonata)

Habitat loss and degradation are considered major threats to freshwater biodiversity and to invertebrates in particular. These often irreversible processes may lead to local and regional extinctions of species, most notably of stenotopic taxa. In spite of this, a number of studies have shown that small habitat patches can sustain rich and abundant communities. The present work assesses the relevance of a group of four small man-made (secondary) wildlife ponds to Odonata species diversity and abundance. Results obtained on pond recruiting capacity, species richness, abundance and habitat use by means of exuviae collection and monitoring of adults using a capture–mark–recapture (CMR) method indicate the potential suitability of these small aquatic biotopes and the surrounding landscape as habitat providers and stepping stone connectors in the Vallès lowlands (Catalonia, Spain). This region, close to the metropolitan area of Barcelona, has severely degraded natural habitats and high landscape fragmentation due to infrastructure, urban and industrial expansion. A comparison among a greater number of sites (ponds and sections of streams and rivers) distributed across the region showed that adequately managed small waterbodies harbour richer Odonata communities than others that are unmanaged or managed specifically for other types of fauna or uses. Appropriate care of these small biotopes avoids disturbance and keeps them free from vertebrates like fish and waterfowl which, under certain conditions, may have a strong influence on the invertebrate communities because, apart from feeding on larvae, they may have a negative impact on macrophyte development and water quality. While rivers and streams, the only natural aquatic habitats in the area, may be both expensive and technically challenging to restore and manage successfully, the creation and/or restoration of small ponds and short river sections in suitable locations can be a cost-effective method for enhancing freshwater vegetation and invertebrate diversity in this impacted landscape.     

A first assessment of the land management effect on the ecological role of large trees as habitat refuges for desert small mammals

Large old trees are keystone organisms that generate a highly connected network of interactions because they provide refuge and feeding sites to mammals with different habitat requirements through their under canopy structure and deadwood. In dry woodlands, these keystone trees are found within agricultural landscapes, where grazing and deadwood removal are the main subsistence activities carried out by local people. These activities can modify the structure of trees and, in turn, affect small mammal communities. Our objective was to assess how different land management types modify the structure of P. flexuosa trees, and to determine the effects of modified tree structure on the abundance and composition of small mammal communities. The study was conducted in P. flexuosa forests within a protected area and in grazing fields. We found that the trees and the vegetation structure beneath their canopies reflect the management history of areas. Trees in the protected area and in an abandoned field were structurally more similar to one another than were those in grazing fields with deadwood removal and, in turn, presented greater total abundances of small mammals. Under tree canopy, the amount of deadwood and grass cover favor the presence of species that need closed and more complex habitats. Also, protection provided by the trees was differentially perceived by species, according to their ecological requirements. We can conclude that different land management scenarios allow for the conservation of the whole rodent assemblage and that, the trees determine the presence of species, particularly of those needing more complex habitats.     

Differentiated macrophyte types as a habitat for rotifers in small mid-forest water bodies

The rotifer community structure may be shaped by a variety of environmental factors, including biological parameters, such as predation or competition, as well as by physical-chemical factors, among which the kind of macrophyte substratum and parameters relating to the trophic state and to the catchment area conditions are of great importance. Another impact on rotifer composition, abundance and frequency may be expected when considering differentiated macrophyte types (including helophytes, nymphaeids and elodeids) within a group of eight ponds located within a mid-forest catchment area, which was the main aim of the present study. Detailed analysis on the participation of indicator eutrophic rotifer species provided an additional goal of this investigation in order to qualify the trophic state of this kind of water body located within the Wielkopolska region, in the central western part of Poland. The presence of 117 taxa, including 7 species that are rare or infrequently distributed in the Polish fauna was recorded. The number of taxa and rotifer abundance differed greatly in respect to the specific water body and to the type of substratum. The vegetated zones dominated by elodeids were characterised by the most various and most abundant rotifer communities as well as by the highest values of the Shannon-Weaver index. This reflects a positive relationship between the heterogeneity of habitat and the structure of rotifer communities. It was also found that helophytes and the open water zone possessed the highest, while elodeids the lowest percentage of eutrophic species.     

The eco-field: A new paradigm for landscape ecology

In the spirit of the theory of biocomplexity and of the non-linear emergent characters of ecological systems, the eco-field is a new paradigm that integrates the vision of the landscape as a neutral matrix (like a habitat) in which organisms are living, and contemporarily as a product of the human mind. Eco-field is defined a cognitive field created by the interference between functional traits and the real world. Species-specific environmental suitability is the result of the quality of the different eco-fields and the landscape becomes a cognitive entity. The eco-field paradigm can be extended to the emergent properties of the systems. The eco-field of emergences is the geographic space in which the emergent properties appear. The eco-field of organisms and the eco-field of emergences, like results of aggregated entities, have in common the multidimensionality of landscapes, refusing the vision of landscape like a neutral geographic matrix for organisms and processes.     

Mediterranean Europe as an area of endemism for small mammals rather than a source for northwards postglacial colonization.

There is a general perception that central and northern Europe were colonized by range expansion from Mediterranean refugia at the end of the last glaciation. Data from various species support this scenario, but we question its universality. Our mitochondrial DNA studies on three widespread species of small mammal suggest that colonization may have occurred from glacial refugia in central Europe-western Asia. The haplotypes on the Mediterranean peninsulae are distinctive from those found elsewhere. Rather than contributing to the postglacial colonization of Europe, Mediterranean populations of widespread small mammals may represent long-term isolates undergoing allopatric speciation. This could explain the high endemism of small mammals associated with the Mediterranean peninsulae.     

Role of small mammals in the ecology of Borrelia burgdorferi in a peri-urban park in north coastal California

The role of small mammals other than woodrats in the enzootiology of the Lyme disease spirochete, Borrelia burgorferi, was assessed in a peri-urban park. Mammals were collected monthly from September through to April. Following tick removal, the animals were tested for B. burgdorferi by culture of ear-punch biopsies. Larvae and nymphs that were intermediate in morphology between Ixodes spinipalpis and Ixodes neotomae occurred on several species of rodents (Peromyscus truei, Peromyscus californicus, Microtus californicus, Rattus rattus and Reithrodontomys megalotis) and the brush rabbit (Sylvilagus bachmani). Morphometric analyses of these I. spinipalpis-like ticks and the offspring from two I. neotomae females from this site suggest that I. neotomae may be conspecific with I. spinipalpis. Borrelia burgdorferi was isolated from eight out of 109 (7.3%), three out of 16 (18.8%), two out of 38 (5.3%) and two out of six (33.3%) P. truei, P. maniculatus, M. californicus and R. rattus, respectively. One brush rabbit yielded the first isolate of B. burgdorferi from a lagomorph in western North America. This isolate and three others derived from unfed I. spinipalpis-like nymphs failed to produce infection when inoculated intradermally into 11–12 P. maniculatus each. Likewise, no spirochetes were detected in 420 Ixodes pacificus nymphs derived from larvae fed on animals inoculated with these isolates. An additional isolate, derived from an I. spinipalpis-like nymph, was recovered by ear-punch biopsies from five out of 12 (42%) needle-inoculated P. maniculatus. However, spirochetes were not detected in 20 I. pacificus nymphs fed as larvae on each of five mice (two infected and three uninfected) inoculated with this isolate. We conclude that brush rabbits and several species of rodents besides woodrats may contribute to the maintenance of B. burgdorferi because they harbour the spirochete and are fed upon by competent enzootic and bridge vectors.