Blackbuck Antilope Cervicapra Research Articles

Population demography of the Blackbuck Antilope cervicapra (Cetartiodactyla: Bovidae) at Point Calimere Wildlife Sanctuary, India

Demographic research utilizing vital rates and life tables is a standard aspect of planning protection and management strategies for wildlife populations. The Blackbuck Antilope cervicapra population at Point Calimere Wildlife Sanctuary, Tamil Nadu, has undergone fluctuations in recent decades. The cause remains elusive, and conservation efforts may be hampered by a lack of population data. This study aimed to estimate demographic parameters using population count and age-sex classification data collected for the years 2017─2020. The overall mean population estimate derived from line-transect distance sampling was 719, with annual estimates of 716, 727, 711, and 722 for the years 2017–2020 respectively. In total, 64% of Blackbucks counted were adults, 19% subadults, and 17% fawns. Mortality was highest for adult and subadult classes for the composite female class, and fawn mortality was 20%. The net reproductive rate (R0) was as low as 3.28 offspring per generation contrasted with a rather longer mean generation time (G) as 4.75 years. Thus, the study observes a decrease in Blackbuck numbers postulated in earlier research to be driven principally by a conglomerate of factors, including reduction of usable space and interspecific resource competition. Our findings provide a baseline demography of the species and highlights the value of long-term demographic monitoring of age sex classes to understand the evolution of life histories.

An insight into population structure and seasonal herd pattern of blackbuck Antilope cervicapra (Linnaeus, 1758) (Mammalia: Artiodactyla: Bovidae) in semi-arid region of western Haryana, India

An insight into population structure and seasonal herd pattern of blackbuck Antilope cervicapra (Linnaeus, 1758) (Mammalia: Artiodactyla: Bovidae) in semi-arid region of western Haryana, India

Determinants of diet selection by Blackbuck Antilope cervicapra at Point Calimere, southern India: quality also matters

Unlike the wide-ranging habitat generalists that move seasonally across heterogenous habitats to optimize the energy intake, short-ranging habitat specialists fulfil the same by restricting to single habitat. Understanding how habitat-specialists do this is an interesting question and essential for their conservation. We studied the diet composition and evaluated the covariates belonging to climate, habitat and grass dynamics to assess the determinants of seasonal diet selection by Blackbuck Antilope cervicapra, an antelope endemic to the Indian subcontinent, at Point Calimere Wildlife Sanctuary, southern India. Diet composition studied following feeding trail observation (n = 102322) and the influence of covariates on the top five major diet species selected seasonally was tested using Regression with Empirical Variable Selection. The results showed that overall Blackbucks consumed 30 plant species—six browse and 27 grass species. While wet season diet was less diverse (22 species) with higher dependency on principal diet Cyperus compressus (>40%) and Aeluropus lagopoides (24%), the dry season diet was more diverse (30) species, with decreased dependency on principal diet. Among 13 covariates belonging to climate, habitat, and grass dynamics tested against selection of top five major diet plants by Blackbucks, grass dynamics covariates alone entered as the predictors both in wet and dry seasons. While cover and green leaves of the grass were the most common predictors in the top-five diets selection during wet season, in dry season besides cover and green leaves, grass texture (hard and soft), also entered as the most common predictors. The entry of grass cover, a quantitative related measure, and texture and green condition of the grass, quality related measures, as the drivers indicate that diet selection by Blackbuck is not just a matter of grass quantity, but also its quality.

Population tendencies of Blackbuck Antilope cervicapra in four consecutive years at Basur Amruth Mahal Kaval Blackbuck Conservation Reserve, Karnataka, India

This study was carried out to ascertain the status and trends of blackbuck population (Nt) in four consecutive years from 2017 to 2020 and to investigate the causative factors taking away their livelihood in a fragile and dynamic habitat of Basur Amruth Mahal Kaval Blackbuck Conservation Reserve with an area of 7.36 km2. Our results emerged with population changes (Nt) with total of 238, 262, 277 and 296 individuals in each year respectively displaying steady state increase. Adult females increased from 136 to 189 whereas adult males (59-60) showed near stable numbers prior to increased predation chances at the habitat edges between open grassland and tall bushes where fawns might be rarely detected. The population density (Dp= 32.336 to 40.217 individuals per sq. km) showed gradual increasing trend. The dwindling blackbuck population prefers open grassland habitat with less fragmentation forces and avoid thick and tall vegetations. Population estimations of blackbuck using line transect methods in these areas with mixed vegetations containing tall shrubs (Prosopis juliflora, Dodonaeaviscosa) probably lead to biased detections. Habitat fragmentation, reduced grassland area due to invasive growth of weeds including Barleriamysorensis, overgrazing and poaching activities are the timeline issues of conservation in this area.

Antelope space-use and behavior indicate multilevel responses to varying anthropogenic influences in a highly human-dominated landscape.

A primary means of conserving a species or a habitat in a human‐dominated landscape is through promoting coexistence with humans while minimizing conflict. For this, we should understand how wildlife is impacted by direct and indirect human activities. Such information is rare in areas with high human densities. To investigate how animals respond to altered ecological conditions in human‐dominated landscapes, we focused on a wild herbivore of conservation concern in the Krishnasaar Conservation Area (KrCA) in Nepal. Here, blackbuck Antilope cervicapra, a generalist grazer, lives in refugia located with a growing human population. We studied the impacts of humans on habitat use and behavior of blackbuck. We laid 250 × 250 m grid cells in the entire KrCA and carried out indirect sign surveys with three replications for habitat use assessment. We observed herds of blackbuck for 89 h in different habitat types using scan sampling methods. Our habitat‐use survey showed that habitats under intensive human use were hardly used by blackbuck, even when high‐quality forage was available. Habitat openness was the major predictor of habitat use inside the core area, where levels of human activities were low. We also found a positive correlation between habitat use by blackbuck and livestock. Blackbuck were substantially more vigilant when they were in forest than in grassland, again indicating an influence of risk. Overall, blackbuck appear to be sensitive to the risk associated with both natural and anthropogenic factors. Our findings have direct implications for managing human–wildlife interactions in this landscape, specifically regarding strategies for livestock grazing in habitats highly used by blackbuck and concerning predictions of how changing land use will impact the long‐term persistence of blackbuck. Our work suggests that wild herbivores may be able to persist in landscapes with high human densities so long as there are refuges where human activities are relatively low.

Ecology of Indian wolf Canis lupus pallipes in the Great Indian Bustard Sanctuary, Nannaj, Solapur, Maharashtra, India

The Indian wolf, Canis lupus pallipes is an endangered carnivore of India listed in Schedule-1 of the Indian Wildlife (Protection) Act, 1972. There is not much information available on the species and with this perspective, the present study was initiated in the Great Indian Bustard Sanctuary in Maharashtra. Direct and indirect evidences were used to assess its population status and distribution, habitat use, feeding ecology, denning behaviour and wolf-human conflict emanating due to livestock depredation. Intensive ecological and behavioural studies were conducted at Nannaj on one pack between 1991 and 1994. The wolf survives all over the Sanctuary, which is densely populated by humans as elsewhere in its distribution range in India where wolf-human conflicts are common arising either due to livestock depredation or child-lifting. Usually, large packs were present in Blackbuck Antilope cervicapra areas whereas smaller packs inhabited areas not supporting healthy populations of Blackbuck. In Blackbuck poor areas, the wolf survived mainly on livestock. The Sanctuary supports 40-58 wolves. The wolves had a seasonal pattern of habitat use related to movement of prey and livestock. Wolves selected those areas as rendezvous sites which had higher prey density, vegetation cover, minimum human disturbance and were located close to water source. On an average wolves consumed 3.68 (S.E.=0.17) kg/wolf/kill and kill periodicity was 3.65 days (S.E.=0.58). Wolves preyed largely on blackbuck during the non-breeding period and livestock during denning period. Predation was higher on male blackbuck than females despite female biased sex ratio. The average litter size of three packs was recorded to be 5 (range 4-6). The dominant male and female wolves guarded the dens more often than the helpers. They used multiple dens for raising pups that were excavated and/or renewed during denning period. Important conservation problems of wolf have also been discussed in the paper.

Ecology of Indian wolf Canis lupus pallipes in the Great Indian Bustard Sanctuary, Nannaj, Solapur, Maharashtra, India

The Indian wolf, Canis lupus pallipes is an endangered carnivore of India listed in Schedule-1 of the Indian Wildlife (Protection) Act, 1972. There is not much information available on the species and with this perspective, the present study was initiated in the Great Indian Bustard Sanctuary in Maharashtra. Direct and indirect evidences were used to assess its population status and distribution, habitat use, feeding ecology, denning behaviour and wolf-human conflict emanating due to livestock depredation. Intensive ecological and behavioural studies were conducted at Nannaj on one pack between 1991 and 1994. The wolf survives all over the Sanctuary, which is densely populated by humans as elsewhere in its distribution range in India where wolf-human conflicts are common arising either due to livestock depredation or child-lifting. Usually, large packs were present in Blackbuck Antilope cervicapra areas whereas smaller packs inhabited areas not supporting healthy populations of Blackbuck. In Blackbuck poor areas, the wolf survived mainly on livestock. The Sanctuary supports 40-58 wolves. The wolves had a seasonal pattern of habitat use related to movement of prey and livestock. Wolves selected those areas as rendezvous sites which had higher prey density, vegetation cover, minimum human disturbance and were located close to water source. On an average wolves consumed 3.68 (S.E.=0.17) kg/wolf/kill and kill periodicity was 3.65 days (S.E.=0.58). Wolves preyed largely on blackbuck during the non-breeding period and livestock during denning period. Predation was higher on male blackbuck than females despite female biased sex ratio. The average litter size of three packs was recorded to be 5 (range 4-6). The dominant male and female wolves guarded the dens more often than the helpers. They used multiple dens for raising pups that were excavated and/or renewed during denning period. Important conservation problems of wolf have also been discussed in the paper.

Plant diversity of Point Calimere Wildlife Sanctuary and fodder species grazed by the Blackbuck Antilope cervicapra L.

A rapid but intense survey was conducted using visual landmarks in the Point Calimere Wildlife Sanctuary to enumerate the flora and foraging habits of the Blackbuck Antelope cervicapra. The area was divided into various segments such as the sanctuary entrance, Maattu muni kovil, Savukku plot or Casuarina plantation, S-Bend road and the old light house for precise enumeration. A total of 111 plant species that include 50 herbs, 16 climbers/lianas, 30 shrubs and sub-shrubs, and 15 trees belonging to 39 plant families were recorded in this study. Visual observations showed that Blackbucks grazed on grasses such as the Mangrove Grass Aeluropus lagopoides (L.) Thwaites, Dog’s Tooth Grass Cynodon barberi Rang. & Tadul., Indian Durva Grass Cynodon dactylon (L.) Pers., Feather Finger Grass Chloris virgata Sw., and a sedge, the pointed fimbristylis Fimbristylis acuminata Vahl during the day time. They were also observed browsing on the leaves and pods of Algaroba Prosopis juliflora (Sw.) DC. in the evenings. Our observation on the presence of feral horses and stray cattle in the Point Calimere Wildlife Sanctuary shows that they compete for food and water with the Blackbuck. The spread of invasive alien plant species competes with and reduces the space for native species.

Seasonal food preferences and group activity pattern of Blackbuck Antilope cervicapra (L., 1758) (Mammalia: Cetartiodactyla: Bovidae) in a semi-arid region of western Haryana, India

To evaluate food preferences and group activity patterns, a fragmented population of Blackbuck Antilope cervicapra was selected for observation in a semi-arid ecosystem of western Haryana. A field survey was conducted fortnightly, from dawn to dusk, between September 2019 and August 2020, covering every season. Scan sampling and quadrat methods were used to record data on group size and vegetation. Group sizes ranged from 3 to 72 individuals. Based on visual observation, blackbuck seasonally consumed 26 species belonging to 25 families with varied preferences, out of a total of 53 plant species documented from the study site. Some plant species with high medicinal and therapeutic values were preferred, including Artemisia scoparia, Cucumis callous, Ziziphus jujuba, and Ziziphus nummularia. Unlike most herbivores, Blackbuck also consumed the toxic and medicinally rich Calotropis procera. We suggest that zoos which house blackbuck include these preferred wild plant species in their diet. Observations on group activity were analyzed on hourly, monthly and seasonal bases, and converted into time percentages. Group foraging activity was at a maximum in the monsoon (62%) and minimum in winter (50%), followed by resting: maximum in winter (21%) and minimum (12%) in monsoon, largely influenced by food availability. Foraging/walking ratio was at a maximum (5.2) in monsoon and minimum (3.1) in winter, and was correlated with the number of group sightings (maximum in winter and minimum in monsoon) in nearby farmland, when the animals faced food scarcity in their natural habitat and fed on crops.

Blackbuck Antilope cervicapra (Mammalia: Cetartiodactyla: Bovidae) estimates in human-dominated landscape in Aligarh, Uttar Pradesh, India

Information on the status of the Indian Blackbuck Antilope cervicapra is mostly available from protected areas (PA), although 80% of its population occurs outside PAs. We conducted surveys to assess the status, age structure, sex ratio, and conservation issues of Blackbuck in and around Aligarh between February and June 2014. A median of 672 individuals at 18 separate locations were recorded with a group size ranging 4–216 except for solitary individuals. The abundance of Blackbuck was maximum in Sikandra Rao (range: 154–216) followed by Andla (range: 47–65), and Pala-Sallu (range: 53–62). Sex ratio was skewed towards females (1:4.5) with yearling to female and fawn to female ratio of 7.8:100 and 6.7:100, respectively. The percentage of adult males of Blackbuck (12.8% adult males, 8.4% sub-adult males) as well as adult females (56.4% adult females, 11.9% sub-adult females) was higher than other age classes or groups in the population. The preliminary observations indicate that the increasing population of free-ranging feral dogs, degradation of forest patches, social forestry plantations, competition with livestock, and poaching pressure are the major conservation issues of Blackbuck in the area. The current information is expected to serve as baseline in assessing the population of Blackbuck in the future.

From extinction to recovery: the case of blackbuck Antilope cervicapra from Nepal

From extinction to recovery: the case of blackbuck Antilope cervicapra from Nepal

Influence of invasive Prosopis juliflora on the distribution and ecology of native blackbuck in protected areas of Tamil Nadu, India

Baseline data on intrinsic–extrinsic determinants of spatial distribution of flagship species helps wildlife managers develop their management strategies. Globally, large herbivores are in range contraction due to the expansion of human habitation. Of late, alien invasion is raising management concerns due to its detrimental effect on native biota. This study evaluating the determinants of spatial detectability and density distribution of blackbuck Antilope cervicapra, an endemic species of the Indian subcontinent, demonstrates the negative influence of Prosopis juliflora, an alien invasive, on the native blackbuck at three reserves—Point Calimere Wildlife Sanctuary, Guindy National Park and Sathyamangalam Tiger Reserve, Tamil Nadu, India. Using distance sampling technique, the study has surveyed blackbuck from 26 grid-based, spatially replicated line-transects between January 2018 and July 2019. Furthermore, 27 covariates related to climate, habitat and anthropogenic pressure were evaluated to determine their effect on spatial detectability and density distribution. Regression by Empirical Variable Selection showed that blackbuck spatial detectability and density distribution increase significantly with the extent of grassland, habitat openness and grass biomass, but decreases with Prosopis cover, shrub cover, percentage of woodland and distance to water, thus revealing their negative effect on the blackbuck. Besides, the Prosopis cover also decreases significantly the key positive predictors of the density distribution of blackbuck, namely the extent of grassland, habitat openness and grass productivity. Thus, the study concludes that the alien invasive species has a negative effect on the native blackbuck population. This raises the need for controlling or eliminating the invasive species in order to conserve, in the long run, the endemic blackbuck at the three reserves.

Dry season resource selection among sympatric ungulates in a tropical coastal landscape: implications for conservation and management

Differential resource use by sympatric species mediates coexistence by specialising in individual ecological niches. We investigated the population, habitat use and dietary overlaps between wild ungulates, i.e., blackbuck Antilope cervicapra, chital Axis axis, domestic livestock and feral horse at Point Calimere Wildlife Sanctuary, India. The food habits and diet overlap of ungulates were estimated using the feeding site method and the density were estimated using distance sampling-based line transect approach. The estimated overall density (number of individuals per km2 ± SE) of blackbuck was (37.1 ± 5.2), chital (4.5 ± 0.9), feral horse (7.6 ± 1.2) and cattle (21.9 ± 3.3). The target species fed on 43 plant species, of which, plant use was high in feral horse (35), followed by chital (33), cattle (29) and blackbuck (23). Dung/pellet analysis revealed that all the target species used monocots (graminoids) in high proportion. Feral horse had the highest habitat niche breadth (0.89) followed by cattle (0.72), blackbuck (0.48) and chital (0.26). Diet overlap was maximum between blackbuck and cattle (95%) followed by feral horse (81%), and chital (38%) with respect to blackbuck. The observed high degree of diet overlaps between wild ungulates with domestic livestock and feral horses is a matter of concern for the long-term survival of the wild ungulates.

Modelling the effect of covariates on the detectability and density of native blackbucks and invasive feral-horse using Multiple Covariate Distance Sampling at Point Calimere Wildlife Sanctuary, Southern India

Reliable estimates of population size and a knowledge of determinants of detectability and density estimates are crucial for effective conservation of species. Using Multiple Covariate Distance Sampling (MCDS), we modelled the influence of covariates on detection probability and density estimates of native blackbuck Antilope cervicapra and the invasive feral-horse Equus caballus at Point Calimere Wildlife Sanctuary. Grids of 1 × 1 km size were overlaid on the study area with a 1-km line transect in alternate grid cells. Sixteen transects were walked four times each, which detected 199 blackbuck and 152 feral-horse clusters. On each sighting, the climatic, habitat and anthropogenic covariates were recorded, which are likely to affect detection probability. At first, exploratory analyses were made using Conventional Distance Sampling (CDS) to arrive at estimates. Later, in Multiple Covariate Distance Sampling (MCDS), key models were fit into the dataset after selecting the potential covariates that had a significant effect on detection distances obtained from FAMD/PCA, and the best one was selected based on AIC. The MCDS analysis in blackbuck included covariates, viz., distance to water in the best model (29 individuals/km2; CI 22–36; detection probability = 0.58), followed by distance to feral-horse, sampling time and principal diet. The covariate distance to water emerged as the best model for feral-horse as well (13 individuals/km2; CI 9–18; detection probability = 0.36), followed by distance to cattle and principal diet. While the MCDS approach outperformed the CDS global and survey-strata estimates in blackbuck, both the approaches had a marginal difference in feral-horse. Post-stratification analysis showed that blackbuck density increased significantly with distance to water and feral-horse, but insignificantly with the absence of principal diet; whereas, the density of the feral-horse increased significantly with distance to water and cattle in the presence of principal diet. These findings suggest the ability of feral-horse to keep the blackbuck away from optimal areas, where the principal diet is abundant. Our study, thus, illustrates the need for the use of MCDS approach that ascertained (i) reliable population density estimates, (ii) spatiotemporal constraint on large herbivores caused by overabundance of water during the wet season, and (iii) competitive interaction of the invasive feral-horse with the native blackbuck and its likely effect on the latter keeping away from the former and the principal diet to overcome competition. Thus, the study highlights the impact of feral-horse on the native species and suggests measures for the long-term conservation of the blackbuck.

<b>Population size, herd structure and sex ratio of the Blackbuck <I>Antilope Cervicapra</I> (Mammalia: Cetartiodactyla: Bovidae) in a human dominated area in Odisha, India</b>

Blackbuck Antilope cervicapra in human dominated landscapes are vulnerable to poaching, habitat loss and competition with livestock for forage. I estimated population sizes and age structure of Blackbucks in an unprotected site of 61.21km² in Odisha over a period of one year (October 2012 to October 2013). A total of 7,134 individuals in 366 herds were documented ranging from a single individual to the largest herd of 51 animals. Average herd size was 19.49±0.03 (SE) and ranged from 13.34±0.06 in summer to 31.86±0.07 during the monsoon. Sex ratio was skewed towards females by 3:1. The young constituted 16% of the population. This indicates that a healthy population of blackbuck is surviving in this area; therefore measures need to be taken to conserve this site and manage the area as a Blackbuck reserve.

<b>Measuring Indian Blackbuck <I>Antilope cervicapra</I> (Mammalia: Cetartiodactyla: Bovidae) abundance at Basur Amruth Mahal Kaval Conservation Reserve, Chikkamagaluru, southern India</b>

Grasslands are among the most critically endangered ecosystems in the tropics, but they are often treated as wastelands and conservation efforts are seldom directed towards these landscapes. The Blackbuck Antilope cervicapra is a large wild herbivore found in most grassland ecosystems across India. Despite their critical role in their trophic web, there are no reliable estimates of Blackbuck populations from their geographic range that takes detection probability into consideration. In this study, we conducted field surveys to estimate Blackbuck density in Basur Amruth Mahal Kaval Conservation (BAMKCR) with an area of 7.36km2 in southern India. We surveyed Blackbucks for a week in July 2014 along straight line transects between 09:00–12:00 hr and used the distance sampling approach to address the imperfect detection. A total of three transect lines of lengths 3.01km, 2.4km and 1.2km were sampled for seven temporal replicates. With an effort of 46.27km, 56 sightings of Blackbucks were recorded that was analyzed using the program DISTANCE. With a detection probability of 0.58 (0.053 SD) the estimated density of Blackbuck was 26.23 (6 SD) individuals/km2. The derived abundance estimate was 193 (c. 148–238) individuals in the study area. Our results show implications of a statistically robust design that accounts for imperfect detection. It provides an insight into a resident population of Blackbucks in a dynamic and fragile habitat. Blackbuck density estimate from this study sets the background for periodic monitoring of their populations, examination of the impacts of habitat modifications and gauge long-term viability of the grassland habitat in BAMKCR.

Crop Damage by the Blackbuck Antilope Cervicapra in and around Chitta Reserve Forest of Bidar, Karnataka

Crop Damage by the Blackbuck Antilope Cervicapra in and around Chitta Reserve Forest of Bidar, Karnataka

Intraspecific variation in group size in the blackbuck antelope: the roles of habitat structure and forage at different spatial scales

The main ecological factors that are hypothesized to explain the striking variation in the size of social groups among large herbivores are habitat structure, predation, and forage abundance and distribution; however, their relative roles in wild populations are not well understood. I combined analyses of ecological correlates of spatial variation in group size with analyses of individual behaviour in groups of different sizes to investigate factors maintaining variation in group size in an Indian antelope, the blackbuck Antilope cervicapra. I measured group size, habitat structure, forage, and the occurrence of predators in ten blackbuck populations, and, at a smaller spatial scale, within an intensively studied population. To examine the processes by which these ecological factors influence group size, I used behavioural observations and an experiment to estimate the shape of the relationship between group size and potential costs and benefits to individuals. Group size varied extensively both among and within populations. Analyses of spatial variation in group size suggested that both forage and habitat structure influence group size: large-scale, among-population variation in group size was primarily related to habitat structure, while small-scale, within-population variation was most closely related to forage abundance. Analyses of individual behaviour suggested that larger groups incur greater travel costs while foraging. However, individuals in larger groups appeared to experience greater benefits, namely the earlier detection of a "predator", a reduction in vigilance, and an increase in the time spent feeding. Overall, these findings suggest that individuals in groups experience a trade-off between predation-related benefits and costs arising from feeding competition. Habitat structure and forage likely influence the nature of this trade-off; thus, variation in these ecological factors may maintain variation in group size. The role of predation pressure and other factors in explaining the remaining variation needs further exploration.

VARIATION IN LEKKING COSTS IN BLACKBUCK (ANTILOPE CERVICAPRA): RELATIONSHIP TO LEK-TERRITORY LOCATION AND FEMALE MATING PATTERNS

AbstractWe studied variation in lekking costs in an Indian antelope, the blackbuck Antilope cervicapra, in relation to female mating patterns. We tested the hypothesis that central males had a higher mating success and faced higher costs than peripheral males. We used continuous focal animal sampling to estimate time-activity budgets of individually identified central and peripheral lekking males and bachelor males. Scan sampling was used to estimate the time spent on the lek by central and peripheral males and to monitor female visits to the lek. We mapped lek-territories and monitored territory additions, territory turnover, and the location of matings. We found that central males faced higher costs than peripheral males. Central males foraged less, spent more time on the lek and tended to have higher fighting rates than peripheral males. Corresponding to this difference in costs, mating benefits were also greater for central males. Our results suggest that lek-territory location may be an important cue in female choice. Male mating success was skewed and 90% of matings observed occurred in the lek-centre. Furthermore, three of the five most successful males previously held territories in the lek-periphery where they were not observed to mate. Territory turnover was higher in the centre than in the periphery and males tended to move towards the centre while shifting territories. Based on these findings, we suggest that central and peripheral males follow two different strategies in response to a female preference for central territories: while central males may try to maximise encounters with oestrous females by investing heavily in lekking, peripheral males seem to attempt to maximise their chances of gaining central territories by not compromising on foraging time and investing less in lekking. Since we did not directly measure male phenotype, its role in explaining the patterns we found remains to be studied.