The demography of the lion-tailed macaque (Macaca silenus) in rain forest fragments in the Anamalai Hills, South India

We examined demographic consequences of habitat fragmentation in the lion-tailed macaque in the Anamalai Hills in southern Western Ghats. The parameters examined were group size, age/sex composition, and birth rate, in relation to various habitat parameters. Demographic parameters were estimated for 11 groups in 8 rain forest fragments, during January to May 1996. Area, tree density, canopy cover, canopy height, and tree basal area were estimated for these fragments. As fragment area decreased, there was a decline in birth rate and proportion of immatures in the group, and an increase in the number of adult males, and the variability in group size and adult sex ratios. A similar pattern was also observed with decreasing values of other habitat variables. Differences in the founder population size and age structure, demographic randomness, and history of poaching have caused greater variability in group size and adult sex ratios in the small fragments. The lack of dispersal in the small fragments is another reason for the high variability.     

Densities of Two Frugivorous Primates with Respect to Forest and Fragment Tree Species Composition and Fruit Availability

Conservation of wildlife populations requires extensive knowledge of their habitat requirements, efficient methods to evaluate habitat quality, and an understanding of the value of fragments and edges. Kibale National Park, Uganda has areas that differ in the densities of 2 species of frugivorous monkeys—Cercopithecus mitis and Lophocebus albigena—including one on an edge and forest fragments outside the park that lack both species. We compared the basal area densities of important food trees with primate densities. The density of Cercopithecus mitis correlates most strongly with the basal area density of all types of food trees combined. The density of Lophocebus albigena does not correlate with the basal area densities of any category of food trees or with fruit availability. An index of their density—number of groups seen per km walked—correlates to fruit availability but with marginal significance. Lack of a relationship between the basal area densities of food trees and density of Lophocebus albigena may be the result of a mismatch in scale between the forest area measured and their large home ranges. We compared the unused area of forest to the other areas of the forest and the fragments and found it had higher basal area densities in all food tree categories for both species than the fragments and lower basal area densities of most categories than the other parts of the forest, indicating that the fragments are poor quality and would probably be unused even if dispersal were likely.     

Forest Fragmentation Modifies Habitat Quality for Alouatta palliata

Fragmentation reduces habitat area, increases the number of habitat patches, decreases their size, and increases patch isolation. For arboreal mammals such as howlers (Alouatta palliata), canopy modifications from fragmentation processes could also negatively affect habitat quality. We analyzed changes in the composition and plant structure of 15 fragments (1–76 ha) and compared them with vegetation from a continuous tropical rain forest reserve (700 ha) in Los Tuxtlas, Mexico. At each site, we sampled 1000 m² of all trees, shrubs, and lianas with a diameter at breast height (DBH) ≥10 cm. We obtained estimates of species richness, density, and basal area for different ecological groups, DBH ranges, and top food resources for howlers. We used a stepwise multiple regression analysis to determine relationships between fragment characteristics (size, shape index, and isolation) and plant variables. Compared to continuous forest, fragments have altered composition and plant structure, with large trees absent from the canopy. The basal area of top food resources is higher in continuous forest. Fragment size is the best explanation for the differences in composition and plant structure. The largest fragments had greater basal area of top food resources and more large primary trees in the canopy. Overall, our results suggest that fragmentation altered the habitat quality for howlers.     

Lion-tailed macaques (Macaca silenus) in a disturbed forest fragment: Activity patterns and time budget

We describe the activity patterns and time budget of a feral group of lion-tailed macaques that were confined to a disturbed forest fragment of 65 ha and compare the results with those obtained for groups in protected forests. The degraded nature of the study site was reflected in low tree densities, low specific diversity, gaps in the girth distribution of trees, and frequent disturbance by humans. The study group of 43 subjects was twice as large as lion-tailed macaque groups in protected habitats. They spent the most time ranging (34.0%), followed by foraging (23.7%), feeding (17.9%), resting (16.0%), and other activities such as social interactions (8.4%). Monthly variations are significant for all activity categories except ranging. Times spent resting and foraging are negatively correlated (r = −0.5) and show significant seasonal differences. Foraging time was highest from September to November, when key food sources such asCullenia andArtocarpus were absent or marginally available. The study group spent most time (40.4%) at canopy levels between 21 and 30 m. They spent more time each day ranging than resting or feeding and more time terrestrially compared with groups in protected forests. Large group size, poor habitat quality, and seasonal variation in food availability were the major variables affecting their time budget, and these variables accounted for differences from the time budgets of groups in protected forests.     

Influence of the landscape matrix on the abundance of arboreal primates in fragmented landscapes

Composition of the landscape matrix of surrounding forest fragments is thought to be critically important to the survival of arboreal primates because it offers structures that help the animals move between fragments and other foraging sites. However, little is known about the composition of the matrix used by these animals. The aim of this study was to quantitatively assess the importance of the landscape matrix and its effects on primate abundance, using black howler monkeys (Alouatta pigra) living in a landscape fragmented by the expansion of agriculture and pastures for livestock in southeastern Mexico. In 2008, a complete census of the monkeys was carried out across the 2000-ha landscape matrix, and for every site where we observed monkeys, we recorded canopy height, tree basal area, food-source abundance, and distance to the nearest fragment. A total of 244 howler monkeys, distributed among 48 groups (including six solitary males) were counted in the matrix. Mean troop size was 5.6 ± 2.8 individuals, and the mode was three individuals. The highest number of troops and greatest howler monkey abundance were recorded in the isolated trees, the eucalyptus plantation, and orchards. A generalized linear model revealed that monkey abundance tended to be higher in matrix elements with higher canopy height, greater food availability, and closest to rainforest fragments. These results suggest that it is necessary to take into account the many elements of the landscape when drawing up conservation and habitat management plans, particularly in order to establish connectivity among the fragments and elements of the matrix with native trees.     

Dispersal Patterns in <Emphasis Type="Italic">Tarsius spectrum</Emphasis>

One of the most widely accepted explanations for the difference in the sex bias between mammals and birds is that male-biased dispersal in mammals is due to the preponderance of polygynous mating systems exhibited by this class, whereas birds are predominantly monogamous. Spectral tarsiers (Tarsius spectrum) are unusual in that they exhibit variation in its mating system. Although the majority of spectral tarsier groups are monogamous, ca. 15% are polygynous. If mating system influences dispersal, then I predicted that the polygynous groups would exhibit male biased dispersal whereas I predicted that the dispersal patterns of the monogamous groups would be analogous to that exhibited by birds, specifically female biased. Alternatively, I hypothesized that ecological variation may influence dispersal habits in this species. Specifically, I predicted that polygynous groups would exhibit greater habitat quality than monogamous groups. The 2 hypotheses are not mutually exclusive. On the basis of 14 individuals birdbanded between 1994 and 1999, I determined that individuals of both sexes were equally likely to disperse (males, n = 5; females, n = 9). Males dispersed twice as far as females did. The mean dispersal distance for males was 660 m, and for females it was 266 m. Females (77%) were more likely to form a territory adjacent to the parental territory than were males (20%). Individuals exhibited relatively high amounts of site fidelity (86%) that were related to physical characteristics of the sleeping site. Adults that dispersed a second time (n = 4) initially resided in trees that were shorter and had a smaller diameter-at-breast height than the trees of individuals that exhibited site fidelity. The results of my study partly support the parental mating system hypothesis and also support the habitat quality hypothesis.     

Dispersal mode,shade tolerance,and phytogeographical affinity of tree species during secondary succession in tropical montane cloud forest

Secondary succession following land abandonment, represented by a chronosequence of 15 old fields (0–80 years old) and two old-growth forests, was studied in the tropical montane cloud forest region of Veracruz, Mexico. The objective was to determine successional trajectories in forest structure and species richness of trees ≥5 cm DBH, in terms of differences in seed dispersal mode, shade tolerance, and phytogeographical affinity. Data were analyzed using AIC model selection and logistic regressions. Mean and maximum canopy height reached values similar to old-growth forest at 35 and 80 years, respectively. Species richness and diversity values were reached earlier (15 and 25 years, respectively) while basal area and stem density tended to reach old-growth forest values within 80 years. Along the chronosequence, the proportion of species and individuals of wind-dispersed trees declined, that of bird dispersed small seeded trees remained constant, while that of gravity and animal dispersed large seeded trees increased; shade-intolerant species and individuals declined, while intermediate and shade-tolerant trees increased. Shade-tolerant canopy trees were rare during succession, even in the old-growth forest. Tropical tree species were more frequent than temperate ones throughout the chronosequence, but temperate tree individuals became canopy dominants at intermediate and old-growth forest stages.     

Abnormal Pelage Color in an Isolated Population of <Emphasis Type="Italic">Alouatta guariba clamitans</Emphasis> Cabrera, 1940 in South Brazil

We located 4 brown howlers (1 adult male, 2 adult females, and 1 juvenile male) showing abnormally lighter pelage in 3 social groups comprising 5, 6, and 9 individuals in a 20 ha-forest fragment in the State of Rio Grande do Sul, Brazil. Two additional groups composed only of normally colored individuals also live in the fragment, which is isolated from nearby fragments by 267–1009 m. They were the only brown howlers with abnormal pelage color out of a total of 386 individuals belonging to 67 groups in 21 fragments in the 5876-ha study area. The isolation of the forest fragment, its high howler density (2.2 individuals⁄ha), and large group size (8.8 ± 2.4 individuals) may decrease the likelihood of successful immigration into the population, leading to an increased probability of inbreeding that may facilitate the expression of rare alleles.     

Foraging in a pathogen reservoir can lead to local host population extinction: a case study of a Lepidoptera-virus interaction

In 1990, natural infestations of the polyphagous vapourer moth, Orgyia antiqua (Lepidoptera: Lymantriidae) in lodgepole pine plantations in northern Scotland, were studied to ascertain the role of host foraging behaviour on the prevalence of nucleopolyhedrovirus (NPV; Baculoviridae) infection in the population. Aerial dispersal of early instar larvae (L1–L3) from the tree canopy onto heather foliage at the forest understorey, with subsequent relocation back onto the tree as late-instar larvae (L4–L6) appeared to play a significant role in the development of a widespread virus epizootic in which approximately 80% of L4–L6 individuals succumbed to disease. Bioassays of foliage 1 year later showed that the distribution of NPV followed a pronounced vertical gradient through the forest canopy culminating in high concentrations of virus in the forest understorey. Experimental systems comprising potted pine trees positioned above heather bases showed that NPV infections could be acquired by early stage larvae following dispersal from the tree and feeding on the undercanopy vegetation, then translocated to the tree component for secondary transmission to susceptible tree-feeding individuals. Behavioural studies indicated that the tendency for first-, second- and third-instar larvae to disperse to the understorey was probably not influenced by larval density on the tree but was strongly dependent on larval instar. In contrast, the tendency for larvae to relocate from the understorey heather to the tree was affected by both larval density and larval instar, suggesting that both these factors may significantly affect virus acquisition, translocation and transmission in the host population. In the present study, the heather understorey appeared to act as a pathogen reservoir in which virus could persist between host generations. Spatial heterogeneity in virus distribution combined with host foraging behaviour (dispersal and feeding) resulted in the pathogen playing a major role in host population dynamics over an extended time period (3 years). The reservoir theory is supported by the observation that similar dynamics were not observed in O. antiqua populations at neighbouring sites which lacked understorey food plants. Received: 8 June 1998 / Accepted: 5 October 1998     

Effects of Forest Fragmentation on the Abundance of <Emphasis Type="Italic">Colobus angolensis palliatus</Emphasis> in Kenya’s Coastal Forests

We documented the occurrence and abundance patterns of Angola black-and-white colobus (Colobus angolensis palliatus) in 46 coastal forest fragments ranging from 1 ha to >1400 ha in the Kwale District, Kenya. In field surveys conducted in 2001, we also recorded forest spatial, structural, resource, and disturbance characteristics to determine the effects of habitat quality and fragmentation and the factors most critical to the continued survival of the little-known species. We tested 13 hypotheses to explain variation in patch occupancy and abundance patterns of Colobus angolensis palliatus in relation to habitat attributes. Minimal adequate models indicated that the occurrence of colobus in forest fragments is positively associated with fragment area and canopy cover, whereas the density of colobus in occupied fragments is attributable to forest area, the proportion of forest change over the previous 12 yr, and the basal area of 14 major food trees. Large-scale illegal extraction of major colobus food trees in the District for human resource use, in both protected and unprotected forests, together with ongoing forest clearance and modification, are the major threats to Colobus angolensis palliatus in Kenya.     

Population Declines of Colobus in Western Uganda and Conservation Value of Forest Fragments

The processes of habitat loss and fragmentation are probably the most important threats to biodiversity. It is critical that we understand the conservation value of fragments, because they may represent opportunities to make important conservation gains, particularly for species whose ranges are not in a protected area. However, our ability to understand the value of fragments for primates is limited by the fact that researchers have conducted many studies in protected areas, which do not represent most fragments, and studies are typically short term. Here we determine the long-term survival probability of red (Procolobus pennantii) and black-and-white colobus (Colobus guereza) inhabiting forest fragments outside of Kibale National Park, Uganda. Local communities use the fragments primarily for subsistence agriculture and fuelwood. We surveyed primate populations 3 times over 8 yr, made a total inventory of all trees 2 times, contrasted behavior of groups inhabiting 1 fragment with groups in the continuous forest, and judged the conservation value of the fragments by quantifying patterns of forest use by local people. Of the 20 fragments surveyed, 16 supported resident populations of colobus in 1995, 2 were cleared in 2000, and an additional 2 fragments were cleared by 2003. In 1995 we counted 165 black-and-white colobus, whereas in 2000 and 2003, we counted 119 and 75 individuals, respectively. Seven fragments supported red colobus in 1995, 11 in 2000, and 9 in 2003. In 2000 we counted 159 red colobus, while in 2003, we saw 145 individuals. For both species, activity patterns in continuous forest were similar to those in a fragment, with the exception that individuals in the fragment rested more. Colobus in the fragment ate more mature leaves than colobus in the continuous forest did. Fragments supported all the fuelwood needs of an average of 32 people who lived immediately adjacent to them, and partially supported families up to 3 farms away (ca. 400 m), representing 576 people. Intensive harvesting for fuelwood occurred when neighboring households engaged in beer brewing (an average of 9.6% of the households), gin distilling (8.8%), or charcoal production (14.5%). Overall, between 2000 and 2003, the average density of trees declined by 14 trees/ha (range = 0–60 trees/ha). If current rates of clearing continue, the probability that the fragments will continue to support colobus populations is low.     

Behavioural responses of lion-tailed macaques (Macaca silenus) to a changing habitat in a tropical rain forest fragment in the Western Ghats, India.

This study reports critical changes in the behaviour patterns of lion-tailed macaques (Macaca silenus) inhabiting a continuously changing and deteriorating rain forest fragment in the Western Ghats, India. The study area, a privately owned rain forest patch in a tea/coffee garden called Puthuthotam, has suffered two massive selective logging episodes. Over the years, the native rain forest trees have been largely replaced by non-native/pioneer species resulting in loss of canopy contiguity and significant changes in other vegetation parameters. The almost wholly arboreal lion-tailed macaque now spends a considerable amount of time on the ground in this area. The species has also experienced a major shift in its diet, ranging patterns and other activities.     

Predicting the Competitive Regime of Female <Emphasis Type="Italic">Colobus vellerosus</Emphasis> from the Distribution of Food Resources

We examined the spatial and temporal distribution of the foods of ursine colobus (Colobus vellerosus) at Boabeng-Fiema, Ghana as a means to predict the monopolizablity and usurpability of their food resources. Recent evidence suggests that food may not be limiting for folivorous primates, and that male sexual coercion may be a more important influence on folivore social organization. To address the question, we collected focal data on the feeding behavior of adult females and males over 11 mo (September 2000-August 2001) on 2 groups: WW (n = 31–33 individuals) and B (n = 8–16 individuals). We also conducted phenological monitoring and a tree survey of the two-group home ranges to establish food availability and distribution. We used 2 behavioral or organism-defined indicators of feeding behavior to assess potential resource contestability: food site residence time and distance moved between food sites. The colobus fed on a high diversity of species, most of their food trees were not clumped in distribution, within-tree interfood distances were short, and food trees were large. The only condition associated with the potential for monopolization was low food tree density. However, low food tree density may be offset by the colobus’ use of large trees. Taken together, the ecological and behavioral indicators suggest the food resources of Colobus vellerosus had a low potential for monopolization. Our results also indicate mature leaves had the longest food site residence time, which may suggest they should be the most usurpable plant part, though their presumed low quality and high abundance probably counteracted the effect. The pattern implied the potential for direct feeding competition among Colobus vellerosus at Boabeng-Fiema was low and agonistic interactions over food are not expected. Instead, a group size effect on feeding efficiency should be a more predominant influence on feeding efficiency, if food is limiting for the species.     

Genetic Consequences of Habitat Fragmentation in Black-and-Gold Howler (<Emphasis Type="BoldItalic">Alouatta caraya</Emphasis>) Populations from Northern Argentina

Human-induced habitat fragmentation might seriously affect behavioural patterns and the survival of species whose ecological requirements strongly depend on specific environmental conditions. We compared the genetic structure and dispersal patterns of 2 populations of Alouatta caraya (Plathyrrhini, Atelidae) to understand how habitat reduction and fragmentation affect gene flow in this species. We sampled individuals from 7 groups living in continuous forest (CF, n = 46, 22 males and 24 females), and 11 groups that inhabit a fragmented forest (FF, n = 50, 24 males and 26 females). FST values based on 11 microsatellite loci showed a recent genetic differentiation among groups in the FF. In contrast, the CF showed no differentiation among groups. Further, FST values between sexes, as well as kinship relationships, also exhibited differences between habitats. In the CF, both males and females disperse, leading to nondifferentiated groups composed of adults that are not close relatives. Conversely, in the FF, some groups are differentiated, males disperse more than females, and groups are composed of closely related adult females. Our results suggest that habitat fragmentation modifies the dispersal patterns of black-and-gold howlers. These differences between habitats may reflect a reduced gene flow, providing genetic evidence that suggests that habitat fragmentation severely limits the howler’s ability to disperse. An increasing level of isolation due to uncontrolled deforestation may cause similar loss of genetic diversity on other arboreal primates, and nonprimates that depend on forest continuity to disperse, reducing their abilities to cope with environmental changes.     

Microhabitat use by the larvae of cryptic lamprey species in <Emphasis Type="Italic">Lethenteron reissneri</Emphasis> in a sympatric area

Microhabitat use by the larvae of two cryptic lamprey species in Lethenteron reissneri, Lethenteron sp. N (northern form) and Lethenteron sp. S (southern form) larvae, as well as intraspecific variations on a size-related basis, were examined in a study area (500 × 5 m) located on a stream where they showed sympatric distribution. The total number of individuals collected from each of 31 quadrats (60 × 60 cm) ranged from 0 (0 n/m²) to 13 (36.1 n/m²). The distinct difference in habitat use was not observed between the two species in the present study area. Lamprey density was not significantly independent among the size groups (small and large) for each of Lethenteron sp. N and Lethenteron sp. S. The density of each size group of the two species was significantly correlated with some habitat variables, and stepwise multiple regression analysis revealed a positive association for a substrate with a smaller particle size and deeper depth for large individuals of Lethenteron sp. N and Lethenteron sp. S, respectively, suggesting that these conditions would be suitable for burrowing. On the other hand, small individuals of both species tended to occur in areas where there was a high content of medium sand and a faster current velocity, probably reflecting a preference for respiratory efficiency. Various condition of habitat should be maintained to conserve the cryptic lampreys.     

The Influence of Large Tree Density on Howler Monkey (Alouatta palliata mexicana) Presence in Very Small Rain Forest Fragments

The populations of the Mexican mantled howler monkey ( Alouatta palliata mexicana ) in the Los Tuxtlas region, Mexico, have declined drastically due to habitat loss and fragmentation. Nevertheless, several troops still inhabit very small and isolated rain forest fragments. We identified the main vegetation attributes that can favor the presence of howlers within 18 small (< 10-ha) fragments that did not differ significantly in size, shape, and isolation (nine occupied and nine unoccupied by howlers). We found that habitat quality ( i.e. , food resources and vegetation structure) affected howler incidence in small fragments. Particularly, the occupied fragments showed greater density of big trees (dbh > 60 cm), greater total basal area, greater basal area of persistent tree species, and greater basal area of top food species than the unoccupied fragments; suggesting that even for small fragments the loss of big trees and particularly the decrease in size class of the top food species can negatively affect howler distribution in highly fragmented landscapes. These findings could be used to establish foreground conservation areas for this critically endangered subspecies in fragmented landscapes of Los Tuxtlas.     

Low genetic diversity and fragmentation effects in a wind-pollinated tree, Polylepis multijuga Plige (Rosaceae) in the high Andes

Fragmentation is predicted to increase inbreeding depression and lower the evolutionary potential of organisms by disrupting dispersal. Trees may be more resilient to fragmentation effects due to potential long-distance dispersal mechanisms that genetically connect fragments. Polylepis woodlands in the high Andes are highly fragmented and are currently the focus of reforestation and conservation efforts. Polylepis multijuga Plige (Rosaceae) is a threatened, endemic tree species in the northern Andes of Peru. Samples were collected from 371 adult trees in nine forest fragments separated by 0.5–80 km and genotyped at amplified fragment length polymorphism loci (AFLP) and chloroplast intergenic regions to determine the connectedness of fragments and their suitability for collecting seed for restoration efforts. P. multijuga is wind-pollinated and dispersed; however, genetic diversity in P. multijuga was about half that reported for other wind-pollinated species. Genetic spatial autocorrelation and patterns of chloroplast and AFLP diversity suggest seed dispersal is very limited and that wind dispersed pollen does not effectively connect all fragments. Conservation of this species will require reforestation efforts and possibly augmentation of some fragments to increase their genetic diversity. Collecting seed from multiple large fragments and from individuals separated by at least 25 m within fragments would maximize the genetic diversity of seed collections for reforestation or augmentation. Future studies of this and other Polylepis species should determine how complex topography may affect wind mediated dispersal between fragments and patterns of genetic diversity.     

Effects of Habitat Structure and Adjacent Habitats on Birds in Tropical Rainforest Fragments and Shaded Plantations in the Western Ghats, India

As large nature reserves occupy only a fraction of the earth’s land surface, conservation biologists are critically examining the role of private lands, habitat fragments, and plantations for conservation. This study in a biodiversity hotspot and endemic bird area, the Western Ghats mountains of India, examined the effects of habitat structure, floristics, and adjacent habitats on bird communities in shade-coffee and cardamom plantations and tropical rainforest fragments. Habitat and birds were sampled in 13 sites: six fragments (three relatively isolated and three with canopy connectivity with adjoining shade-coffee plantations and forests), six plantations differing in canopy tree species composition (five coffee and one cardamom), and one undisturbed primary rainforest control site in the Anamalai hills. Around 3300 detections of 6000 individual birds belonging to 106 species were obtained. The coffee plantations were poorer than rainforest in rainforest bird species, particularly endemic species, but the rustic cardamom plantation with diverse, native rainforest shade trees, had bird species richness and abundance comparable to primary rainforest. Plantations and fragments that adjoined habitats providing greater tree canopy connectivity supported more rainforest and fewer open-forest bird species and individuals than sites that lacked such connectivity. These effects were mediated by strong positive effects of vegetation structure, particularly woody plant variables, cane, and bamboo, on bird community structure. Bird community composition was however positively correlated only to floristic (tree species) composition of sites. The maintenance or restoration of habitat structure and (shade) tree species composition in shade-coffee and cardamom plantations and rainforest fragments can aid in rainforest bird conservation in the regional landscape.     

Status and Distribution of Macaca fascicularis umbrosa in the Nicobar Islands, India

We surveyed the distribution and status of the long-tailed macaque (Macaca fascicularis umbrosa) in the Nicobar Islands of India. Long-tailed macaques live on 3 islands: Great Nicobar, Little Nicobar and Katchal. There are a total 788 groups. Group size ranged from 25 to 56 individuals with a mean size of 36. Long-tailed macaques are a multimale-multifemale society. Adult males, adult females and immatures constituted 9.7 per cent, 43.2 per cent and 47.1 per cent of the population, respectively. A fairly high ratio of immatures to adult females indicates a healthy population turnover. Vegetation parameters including tree density, canopy cover, canopy height and basal area indicated a still healthful habitats on all the 3 islands. We observed a few threats to the macaques. Measures should be undertaken to ensure survival of the Nicobar subspecies.     

Foraging behaviour of the root vole Microtus oeconomus in fragmented habitats

The effect of habitat fragmentation on spatial foraging behaviour in the root vole Microtus oeconomus was investigated in seven experimental populations. Four of the populations were established in large, continuous blocks (30 × 95 m) of meadow habitat (treatment plots), whereas the three remaining populations had six small rectangular habitat fragments (30 × 7.5 m) with variable inter-fragment distances (control plots). Both the small habitat fragments and the large continuous habitat were embedded in a non-habitat matrix area which was regularly mowed. Half-way through the study period, the continuous habitat in treatment plots was destroyed by mowing to give a configuration identical to the control plots. Dyed bait placed at the edges and in the interior of habitat fragments as well as in the matrix area was used to reveal differential use of these areas for foraging. Animals in the small-fragment plots fed more than expected along the edges, while edges were used according to availability in the large blocks of continuous habitat. In the fragmented plots, the frequency of foraging in the matrix decreased with increasing distance to the fragment border and with increasing inter-fragment distances. Furthermore, the frequency of use of more than one habitat fragment in individual foraging ranges decreased with increasing inter-fragment distances. Reproductively inactive animals of both sexes fed more often along habitat edges than reproductively active animals. Reproductively active females fed exclusively in one habitat fragment, whereas inactive animals and especially reproductively active males frequently included more than one fragment in their foraging ranges. The only effect of habitat destruction was less foraging in the matrix habitat in the post-destruction treatment plots compared to the permanently fragmented control plots. This was probably an effect of different matrix quality. Root voles in these experimental populations forage in edge and matrix habitat with great risk of becoming victims to predation, and the results are interpreted in this context. Received: 19 August 1998 / Accepted: 30 June 1999