Food mechanical properties in three sympatric species of Hapalemur in Ranomafana National Park,Madagascar

We investigated mechanical dietary properties of sympatric bamboo lemurs, Hapalemur g. griseus, H. aureus, and H. (Prolemur) simus, in Ranomafana National Park, Madagascar. Each lemur species relies on bamboo, though previous behavioral observations found that they specialize on different parts of a common resource (Tan: Int J Primatol 20 1999 547–566; Tan: PhD dissertation 2000 State University of New York, Stony Brook). On the basis of these earlier behavioral ecology studies, we hypothesized that specialization on bamboo is related to differences in mechanical properties of specific parts. We quantified mechanical properties of individual plant parts from the diets of the bamboo lemur species using a portable tester. The diets of the Hapalemur spp. exhibited high levels of mechanical heterogeneity. The lemurs, however, could be segregated based on the most challenging (i.e., mechanically demanding) foods. Giant bamboo culm pith was the toughest and stiffest food eaten, and its sole lemur consumer, H. simus, had the most challenging diet. However, the mechanical dietary properties of H. simus and H. aureus overlapped considerably. In the cases where lemur species converged on the same bamboo part, the size of the part eaten increased with body size. Plant parts that were harvested orally but not necessarily masticated were the most demanding, indicating that food preparation may place significant loads on the masticatory apparatus. Finally, we describe how mechanical properties can influence feeding behavior. The elaborate procurement processes of H. simus feeding on culm pith and H. griseus and H. aureus feeding on young leaf bases are related to the toughnesses of protective coverings and the lemurs' exploitation of mechanical vulnerabilities in these plants. Am J Phys Anthropol, 2009. © 2008 Wiley‐Liss, Inc.     

Variability in habitat, diet, and social structure of Hapalemur griseus in Ranomafana National Park, Madagascar

This study investigated the effects of habitat on the diet, population density, and social structure of a small-bodied folivore, Hapalemur griseus. Three groups of H. griseus were followed at two study sites (Tala and Vato) within Ranomafana National Park in southeastern Madagascar. These two sites differed in degree of habitat disturbance, forest composition, and forest structure, as determined by botanical plots (50 x 10 m) which were monitored twice monthly in the home ranges of three study groups (group T, n = 4; group V1, n = 4; group V2, n = 2). Tala has experienced more habitat disturbance compared to Vato, and this appeared to influence population density and group size. The H. griseus that inhabit Tala occur at much higher densities and in larger social groups than the H. griseus at Vato. These results may be attributable to: 1) a greater number of potential food sources, 2) a more dense resource base, and/or 3) higher-quality resources at Tala for this folivore. In general, dietary diversity for all groups was low, and groups exploited similar plant species. However, within this small range of food sources, each study group specialized in food sources found in the highest abundance within their home range. This resulted in dietary differences between groups within sites, as well as differences between sites. Introduced tree species such as Chinese guava (Psidium cattleyanum) may have greatly impacted the diet and social structure of those groups at Tala, whose home ranges included this food source. In contrast, Vato group 1 ingested more new and mature leaves of Ficus spp., and Vato group 2 primarily ate spider bamboo (Nastus elongatus). In conclusion, it seems that microhabitat differences, which may be related to habitat disturbance and/or other factors such as topography, influenced the food species ingested by H. griseus. Overall habitat quality, which is likely also affected by habitat disturbance, influenced general diet, population density, and group size.     

Phylogenetic relationships among Lemuridae (Primates): evidence from mtDNA

The family Lemuridae includes four genera: Eulemur, Hapalemur, Lemur,Varecia. Taxonomy and phylogenetic relationships between L. catta, Eulemur and Hapalemur, and of Varecia to these other lemurids, continue to be hotly debated. Nodal relationships among the five Eulemur species also remain contentious. A mitochondrial DNA sequence dataset from the ND 3, ND 4 L, ND 4 genes and five tRNAs (Gly, Arg, His, Ser, Leu) was generated to try to clarify phylogenetic relationships w ithin the Lemuridae. Samples (n=39) from all ten lemurid species were collected and analysed. Three Daubentonia madagascariensis were included as outgroup taxa. The approximately 2400 bp sequences were analysed using maximum parsimony, neighbor-joining and maximum likelihood methods. The results support monophyly of Eulemur, a basal divergence of Varecia, and a sister-group relationship for Lemur/Hapalemur. Based on tree topology, bootstrap values, and pairwise distance comparisons, we conclude thatVarecia and Eulemur both represent distinct genera separate from L. catta. H. griseus andH. aureus form a clade with strong support, but the sequence data do not permit robust resolution of the trichotomy involving H. simus, H. aureus/H. griseus and L. catta. Within Eulemur there is strong support for a clade containing E. fulvus, E. mongoz and E. rubriventer. However, analyses failed to clearly resolve relationships among those three species or with the more distantly related E. coronatus and E. macaco. Our sequencing data support the current subspecific status of E.m. macaco and E.m. flavifrons, and that of V.v. variegata and V.v. rubra. However, tree topology and relatively large genetic distances among individual V.v. variegata indicate that there may be more phylogenetic structure within this taxon than is indicated by current taxonomy.     

Chromosomal evolution in Malagasy lemurs. III. Chromosome banding studies in the genus Hapalemur and the species Lemur catta.

The karyotypes of five Malagasy lemurs are described and compared after the successive use of various banding methods. Phylogenic relationships appear to exist between the respective ancestors of Lemur catta, Hapalemur simus, H. griseus occidentalis, H. griseus ssp, and H. griseus griseus. Their karyotypes can be derived from that of L. fulvus fulvus by chromosomal rearrangements affecting mainly chromosomes 1 and X. Intense staining of juxtacentromeric heterochromatin was detected among the different subspecies of H. griseus.     

Effects of Transect Selection and Seasonality on Lemur Density Estimates in Southeastern Madagascar

I investigated how transect type (trails vs. cut transects) and seasonality influenced density estimates for 5 lemur taxa (Avahi laniger, Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus) in the Vohibola III Classified Forest in SE Madagascar. I surveyed tree height and diameter and lemur populations from June 1 to December 28, 2004 along 2 1250-m trails local people used and 2 1250-m transects cut parallel to the trails in primary rain forest. Despite dendrometric variations within and between trails and transects, only density estimates of Hapalemur griseus griseus differed significantly by transect type. The spatial variation may be a result of removal by local people of giant bamboo, which is the main food for Hapalemur griseus griseus, along trails. Conversely, seasonality influenced density estimates for Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus. The temporal variations may be related to seasonal torpor for Cheirogaleus major and increased detection probabilities during periods of fruit exploitation for Eulemur rubriventer, Microcebus rufus, and Hapalemur griseus griseus. Transect type and seasonality did not affect density estimates for Avahi laniger, which may be related to the highly folivorous and low-energy diet of the nocturnal lemur. Researchers surveying lemurs along line transects should be aware that transect selection may influence density estimates for Hapalemur griseus griseus and that seasonality may influence density estimates for Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus.     

Seasonal variation in activity and diet in a small-bodied folivorous primate,Hapalemur griseus,in southeastern Madagascar

How small-bodied (500–1,200 g) folivorous prosimian primates cope with large amounts of foliage in their diet seasonally has yet to be determined for many species such as Hapalemur griseus, which specializes on bamboo. To address this issue, we present results on seasonal variation in activity and diet from a wild group of H. griseus in southeastern Madagascar. Throughout the study (which was conducted from July–November 1994 and July 1995–February 1996), H. griseus primarily fed on new growth from three species of bamboo: two species of liana bamboo and Cephalostachyum perrieri. Bamboo species were used in different ways seasonally; liana bamboo was consumed more during the dry, cool season, and C. perrieri was eaten more often during the wet, warm season. H. griseus also spent more of their time feeding and traveling than nocturnal folivores of similar body size during the dry season. During the warm wet season, H. griseus decreased the amount of time spent feeding and traveling and rested more often. We hypothesize that seasonal changes in activity may be primarily related to the distribution and availability of food sources and/or reproductive cycles. Am. J. Primatol. 43:211–223, 1997. © 1997 Wiley-Liss, Inc.     

Prosimian hemoglobins--V. The primary structures of the alpha-I, alpha-II and beta-hemoglobin chains of Hapalemur griseus, with a note on the classification of Microcebus

1. The duplicated adult hemoglobins were isolated from a mature Hapalemur griseus and the constituent chains prepared. Sequence analysis of the isolated alpha-globins showed that the alpha-I and alpha-II chains differed by a glycine for lysine substitution at position 15. 2. The complete amino acid sequence of the single adult beta-globin of Hapalemur griseus was determined. The beta-globin sequence of Hapalemur griseus clusters with those of other authentic lemurs, and is clearly separated from the sequences characteristic of lorisiform primates. 3. Partial sequence analysis of the beta-globin of Microcebus murinus showed only a single amino acid difference when compared to the Hapalemur globin. 4. Partial sequence analysis of the alpha-globin of Microcebus murinus showed only three amino acid residues that are not found in other lemuriform alpha-globins; two of these are unique to Microcebus among all known prosimian alpha-globin sequences. 5. The Microcebus alpha- and beta-globins are more similar to the homologous lemuriform sequences than they are to lorisiform sequences.     

Description of the gastrointestinal tract of five lemur species: Propithecus tattersalli, Propithecus verreauxi coquereli, Varecia variegata, Hapalemur griseus, and Lemur catta

The objective of this project was to better define the similarities and differences in gastrointestinal morphology present in lemur species. Measurements of the gastrointestinal tract of lemurs were obtained at necropsy from the captive population at Duke University Primate Center. Measurements of body length and weight, as well as gastrointestinal length, were recorded from five prosimian species: Propithecus tattersalli, Propithecus verreauxi, Varecia variegata, Hapalemur griseus, and Lemur catta. Photographs and measurements were used to obtain illustrations. Preliminary results suggest differences in gastrointestinal morphology among lemur species that coincide with differences in diet. Distinct sacculations in either the cecum or the colon were present for H. griseus, L. catta, P. verreauxi, and P. tattersalli, but not for V. variegata. The Propithecus specimens possessed a much greater ratio of gastrointestinal length to body length than the other three species. A short, blunt cecum and a shortened and sacculated colon were unique characteristics of the H. griseus specimens. These differences correlate well with a dietary shift from consumption of large amounts of structural plant cell wall (Propithecus sp.) to consumption of variable or moderate amounts (H. griseus, L. catta, and V. variegata). They also suggest that captive groups would benefit from further diet refinement in captivity.     

Confirmation of the specific status ofHapalemur aureus (Primates,Strepsirhini) by restriction genomic DNA banding patterns

This study confirms, on the basis of our molecular biology results and in accordance with cytogenetic, morphological and ethological data, the specific status ofHapalemur aureus. Furthermore, it appears clearly thatHapalemr simus began its differentiation fromHapalemur griseus griseus andHapalemur aureus (wich have a common branch) shortly after the separation ofLemur catta from the phylogenetic tree of theL. catta/ Hapalemur group.     

Diet,Activity Patterns,and Ranging Ecology of the Bale Monkey (<Emphasis Type="BoldItalic">Chlorocebus djamdjamensis</Emphasis>) in Odobullu Forest,Ethiopia

Bale monkeys (Chlorocebus djamdjamensis) are little-known primates endemic to the forests of the Bale Massif and Hagere Selam regions of Ethiopia. From August 2007 to May 2008, we conducted the first ever study of the species’ behavior and ecology, focusing in particular on its diet, activity patterns, and ranging ecology in the Odobullu Forest. We studied 2 neighboring groups (group A: 55–60 members; group B: 46–50 members) and conducted behavioral scan samples on the first 2–5 individuals sighted at 15-min intervals. Feeding accounted for 65.7% of the activity budget, followed by moving (14.4%), resting (10.7%), social (7.1%), and other behaviors (2.4%). Overall diet during the study was dominated by young leaves (80.2%), though subjects also ate fruits (9.6%), flowers (3.1%), animal prey (2.3%), shoots (1.5%), stems (1.4%), mature leaves (1.1%), and roots (0.9%). Bale monkeys consumed only 11 plant species; of these, the top 5 species accounted for 94.3% of their diet. The top food item, bamboo (Arundinaria alpina), was responsible for a remarkable 76.7% of their diet, with most (95.2%) of the bamboo consumption consisting of young leaves. Mean daily path length for the study groups was 928 m and mean (100% minimum convex polygon) home range size was 15.2 ha. Though we are cautious in drawing conclusions from only 2 groups, the larger group traveled further per day and occupied a larger home range, patterns suggesting scramble competition may be occurring in Bale monkey groups at Odobullu. The dietary specialization of Bale monkeys on bamboo makes them unique among Chlorocebus spp. and suggests an intriguing ecological convergence with the golden monkeys (Cercopithecus mitis kandti) of Uganda and bamboo lemurs (Hapalemur spp.) of Madagascar. Their narrow ecological niche, limited geographic distribution, and bamboo harvesting by local people for commercial purposes place Bale monkeys at risk of extinction. To ensure the long-term survival of Bale monkeys, appropriate management action should be taken to conserve the species and the bamboo forests upon which it depends.     

Urinary Progestogen and Estrogen Excretion During Pregnancy in Eulemur macaco flavifrons,E. rubriventer,and Hapalemur griseus occidentalis

Via a non-invasive approach, we aimed to provide comparative data on the presence and relative abundance of progestogen and estrogen metabolites excreted into the urine of Eulemur rubriventer, E. macaco flavifrons and Hapalemur griseus occidentalis and to characterize the patterns of progestogen and estrogen excretion during pregnancy. We found that estrone is the major urinary estrogen in Eulemur macaco flavifrons and Hapalemur griseus occidentalis, while 16-hydroxyestrone appeared to be the predominant estrogen in the urine of E. rubriventer. Estradiol-17ß was either absent (Hapalemur) or present in very low amounts. HPLC of progestogens had high levels of immunoreactivity in an assay against 5-pregnane-3-ol-20-one (5-P-3OH) in each species, though the nature of the progestogens measured by the 5-P-3OH assay differed among species. During pregnancy, 5-P-3OH levels remained relatively low in Eulemur rubriventer, whereas in E. macaco flavifrons and Hapalemur griseus occidentalis, a sustained elevation in levels occurred from mid-pregnancy until the final weeks before parturition. Estrogen excretion differed depending on the sex of the fetus in each species, with only females carrying male infants showing clearly elevated estrogen levels during the last 6–8 weeks of gestation. In conclusion, we demonstrate both species similarities and differences in the metabolism and urinary excretion patterns of reproductive hormones throughout gestation in the Lemuridae. The data not only extend our knowledge on the reproductive physiology of lemurs but also show that more studies on other lemur taxa are needed to provide a broader basis for interspecific comparison.     

Linking Laboratory and Field Approaches in Studying the Evolutionary Physiology of Biting in Bamboo Lemurs

A realistic understanding of primate morphological adaptations requires a multidisciplinary approach including experimental studies of physiological performance and field studies documenting natural behaviors and reproductive success. For primate feeding, integrative efforts combining experimental and ecological approaches are rare. We discuss methods for collecting maximum bite forces in the field as part of an integrated ecomorphological research design. Specifically, we compare maximum biting ability in 3 sympatric bamboo lemurs (Hapalemur simus, H. aureus, and H. griseus) at Ranomafana National Park, Madagascar to determine if biting performance contributes to the observed partitioning of a shared bamboo diet. We assessed performance by recording maximum bite forces via jaw-muscle stimulations in anesthetized subjects from each species. Behavioral observations and food properties testing show that the largest species, Hapalemur simus, consumes the largest and most mechanically challenging foods. Our results suggest that Hapalemur simus can generate larger bite forces on average than those of the 2 smaller species. However, the overlap in maximum biting ability between Hapalemur simus and H. aureus indicates that biting performance cannot be the sole factor driving dietary segregation. Though maximum bite force does not fully explain dietary segregation, we hypothesize that size-related increases in both maximum bite force and jaw robusticity provide Hapalemur simus with an improved ability to process routinely its more obdurate diet. We demonstrate the feasibility of collecting physiological, ecological, and morphological data on the same free-ranging primates in their natural habitats. Integrating traditionally laboratory-based approaches with field studies broadens the range of potential primate species for physiological research and fosters improved tests of hypothesized feeding adaptations.     

Lemur responses to edge effects in the Vohibola III classified forest, Madagascar

Forest edges are dynamic zones characterized by the penetration (to varying depths and intensities) of conditions from the surrounding environment (matrix) into the forest interior. Although edge effects influence many tropical organisms, they have not been studied directly in primates. Edge effects are particularly relevant to lemurs because of the highly fragmented forest landscapes found in Madagascar. In this study, data are presented regarding how the densities of six lemur species (Avahi laniger, Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, Microcebus rufus, and Propithecus diadema edwardsi) varied between six 500-m interior transects and six 500-m edge transects in the Vohibola III Classified Forest in SE Madagascar. Diurnal (n = 433) and nocturnal (n = 128) lemur surveys were conducted during June-October 2003 and May-November 2004. A. laniger, E. rubriventer, and H. g. griseus exhibited a neutral edge response (no differences in densities between habitats). M. rufus and P. d. edwardsi had a positive edge response (higher densities in edge habitats), which may be related to edge-related variations in food abundance and quality. Positive edge responses by M. rufus and P. d. edwardsi may ultimately be detrimental due to edge-related anthropogenic factors (e.g., hunting by local people). The negative edge response exhibited by C. major (lower densities in edge habitats) may result from heightened ambient temperatures that inhibit torpor in edge habitats.     

High flexibility in diet and ranging patterns in two golden monkey (Cercopithecus mitis kandti) populations in Rwanda

Many primates exhibit behavioral flexibility which allows them to adapt to environmental change and different habitat types. The golden monkey (Cercopithecus mitis kandti) is a little-studied endangered primate subspecies endemic to the Virunga massif and the Gishwati forest in central Africa. In the Virunga massif, golden monkeys are mainly found in the bamboo forest, while in the Gishwati forest they live in mixed tropical montane forest. Here we describe and compare the diet of golden monkeys in both fragments. Over 24 consecutive months from January 2017 we used scan sampling to record feeding and ranging behavior of two Virunga groups and one Gishwati group totaling ca. 240 individuals. We also examined the phenology of bamboo and fruit trees, key seasonal food plant species for the monkeys. Golden monkeys fed on more than 100 plant species. The Virunga groups were mostly folivorous (between 72.8% and 87.16% of the diet) and fed mostly on young bamboo leaves and bamboo shoots, while 48.69% of the diet of the Gishwati group consisted of fruit from 22 different tree and shrub species. Bamboo shoots and fruit are seasonally available foods and were consumed regularly throughout the period when they were available. Despite being the smallest of the three study groups, the Gishwati group had a larger home range area (150.07 ha) compared to both Virunga groups (25.24 and 91.3 ha), likely driven by the differences in availability and distribution of fruit and bamboo in the habitats. Like other blue monkey subspecies, golden monkeys appear to have a flexible dietary strategy enabling them to adjust diet and ranging behavior to local habitats and available food resources. Additional studies and continuing conservation efforts are needed to better understand how variation in feeding and ranging ecology affects reproduction, population growth, and carrying capacity.     

Characterization of 14 microsatellite marker loci in the grey bamboo lemur (Hapalemur griseus)

Fourteen nuclear microsatellite loci were isolated from a genomic DNA library derived from a free‐ranging bamboo lemur (Hapalemur griseus griseus) from Ranomafana National Park, Madagascar. Expected and observed heterozygosity levels were determined for populations of bamboo lemurs from Ranomafana National Park and a nonprotected area near Tsinjoarivo, Madagascar. Although genotype frequencies for all but one marker were within Hardy–Weinberg expectations for the Tsinjoarivo population, this was not the case for the Ranomafana samples. It is probable that the deviation of genotype frequencies from Hardy–Weinberg expectations in Ranomafana was precipitated by the presence of two taxa within the sample population.     

Comparative use of color vision for frugivory by sympatric species of platyrrhines

Ateles spp. and Alouatta spp. are often sympatric, and although they are mainly frugivorous and folivorous, respectively, they consume some of the same fruit species. However, they differ in terms of color vision, which is thought to be important for fruit detection. Alouatta spp. have routine trichromatic color vision, while Ateles spp. presents the classic polymorphism of platyrrhines: heterozygous females have trichromatic color vision, and males and homozygous females have dichromatic vision. Given these perceptual differences, one might expect Alouatta spp. to consume more reddish fruits than Ateles spp., since trichromats have an advantage for detecting fruits of that hue. Furthermore, since Ateles spp. have up to six different color vision phenotypes, as do most other platyrrhines, they might be expected to include fruits with a wider variety of hues in their diet than Alouatta spp. To test these hypotheses we studied the fruit foraging behavior of sympatric Alouatta palliata and Ateles geoffroyi in Costa Rica, and modeled the detectability of fruit via the various color vision phenotypes in these primates. We found little similarity in fruit diet between these two species (Morisita = 0.086). Furthermore, despite its polymorphism, A. geoffroyi consumed more reddish fruits than A. palliata, which consumed more greenish fruits. Our modeling results suggest that most fruit species included in the diet of A. geoffroyi can be discriminated by most color vision phenotypes present in the population. These findings show that the effect of polymorphism in platyrrhines on fruit detection may not be a disadvantage for frugivory. We suggest that routine trichromacy may be advantageous for other foraging tasks, such as feeding on young leaves.     

Relative efficiency of preferred and nonpreferred patterns of lateralized foraging in the gentle lemur (Hapalemur griseus)

Lateralized patterns of hand use in species-typical bamboo shoot foraging were evaluated for efficiency in five female and six male gentle lemurs (Hapalemur griseus sp.). Efficiency was defined as amount of time required to complete a foraging response sequence. The foraging pattern consisted of four component movements: PULL-IN, COUNTERFORCE, TURN, and FEED-IN. These component movements had been shown in a previous study to incorporate lateral hand biases that formed subsets of patterns that were characteristic for each gentle lemur. The duration of each foraging sequence was measured from the beginning of the first component to the initiation of the terminal component. Frequency of use scores were employed to divide the pattern sequences of each subject into preferred and nonpreferred categories. A within-subjects comparison of the mean durations of preferred patterns (M = 2.56 sec) with those of non- preferred patterns (M = 3.02 sec) found that preferred patterns were executed more rapidly, t(10) = 3.36, P = .007. A multiple regression showed that order of pattern preference accounted for 89% of the variance in mean duration of response time (R² = .89, P = .056). Thus, the use of preferred lateralized hand patterns resulted in more rapid bamboo shoot harvesting. Speed in foraging may be regarded as an adaptive strategy for a species that subsists on a high bulk, low nutrient density food such as bamboo. © 1995 Wiley-Liss, Inc.     

Resource use by howler monkeys (Alouatta palliata) in the rain forest of Los Tuxtlas,Veracruz, Mexico

The feeding preferences of howler monkeys at their northernmost distribution in the Neotropics are reported for an annual cycle. A remarkable selectivity for 27 species representing 15 families was observed. The Moraceae and Lauraceae plant families were the most important in the diet. The howlers spent an almost equal proportion of their feeding time eating leaves and fruit, and displayed a marked preference for young leaves and mature fruit. The consumption of different plant parts was markedly seasonal and the howlers’ ranging behavior was closely associated with the availability of young leaves and mature fruit. Their home range was unusually large (ca. 60 ha) for howlers and the food species exploited occur at very low densities (93%, ≤ 4 ind/ha). They chose food items richer in protein and energy. Alkaloid compounds, present in some of the leaves, play a secondary role in their dietary selectivity.     

Food habits and habitat selection of suburban badgers (Meles meles) in Japan

A study of the Japanese badger Meles meles anakuma was undertaken in Hinode, a suburb of Tokyo, between 1992 and 1998. Faecal analysis, based on 82 samples, revealed that during spring and summer, earthworms ( Megaseolocidae spp.) occurred at high frequency in the diet, with berries ( Rubus spp.), beetles and persimmon Dymopyrus kaki also eaten during summer months. Scavenged food was eaten in early spring when earthworm availability was low, and badgers switched from worms when persimmon became abundant in autumn. Twenty-one Japanese badgers (14 males and seven females) were radio-tracked. Adult badger home ranges were stable, and those of males [40±19 ( sd ) ha, n =7] were larger than those of females [11±6 ( sd ) ha, n =4]. Badger resting sites in each home range were located within 630 m of each other and categorized as setts or couches. Setts were sited within core areas (30% adaptive kernel method) of home ranges. Most setts were on a sub-ridge and avoided west-facing slopes. Couches, mainly in deciduous forest and forest edge, were generally sited towards the periphery of home ranges. Most badger foods were distributed along ecotones between forestry plantations and farmland; earthworms, their main food from late spring to summer, and berry thickets were both concentrated at the edge of conifer plantations. Persimmon trees, the main food source for badgers in autumn, were also found in agricultural land bordering forest edge. Badger home range size was related to forest edge density.