Improving density estimates for elusive carnivores: Accounting for sex-specific detection and movements using spatial capture–recapture models for jaguars in central Brazil

Owing to habitat conversion and conflict with humans, many carnivores are of conservation concern. Because of their elusive nature, camera trapping is a standard tool for studying carnivores. In many vertebrates, sex-specific differences in movements – and therefore detection by cameras – are likely. We used camera trapping data and spatially explicit sex-specific capture–recapture models to estimate jaguar density in Emas National Park in the central Brazilian Cerrado grassland, an ecological hotspot of international importance. Our spatially explicit model considered differences in movements and trap encounter rate between genders and the location of camera traps (on/off road). We compared results with estimates from a sex-specific non-spatial capture–recapture model. The spatial model estimated a density of 0.29 jaguars 100 km⁻² and showed that males moved larger distances and had higher trap encounter rates than females. Encounter rates with off-road traps were one tenth of those for on-road traps. In the non-spatial model, males had a higher capture probability than females; density was estimated at 0.62 individuals 100 km⁻². The non-spatial model likely overestimated density because it did not adequately account for animal movements. The spatial model probably underestimated density because it assumed a uniform distribution of jaguars within and outside the reserve. Overall, the spatial model is preferable because it explicitly considers animal movements and allows incorporating site-specific and individual covariates. With both methods, jaguar density was lower than reported from most other study sites. For rare species such as grassland jaguars, spatially explicit capture–recapture models present an important advance for informed conservation planning.     

Integrating occupancy modeling and interview data for corridor identification: A case study for jaguars in Nicaragua

Corridors are critical elements in the long-term conservation of wide-ranging species like the jaguar (Panthera onca). Jaguar corridors across the range of the species were initially identified using a GIS-based least-cost corridor model. However, due to inherent errors in remotely sensed data and model uncertainties, these corridors warrant field verification before conservation efforts can begin. We developed a novel corridor assessment protocol based on interview data and site occupancy modeling. We divided our pilot study area, in southeastern Nicaragua, into 71, 6 × 6 km sampling units and conducted 160 structured interviews with local residents. Interviews were designed to collect data on jaguar and seven prey species so that detection/non-detection matrices could be constructed for each sampling unit. Jaguars were reportedly detected in 57% of the sampling units and had a detection probability of 28%. With the exception of white-lipped peccary, prey species were reportedly detected in 82–100% of the sampling units. Though the use of interview data may violate some assumptions of the occupancy modeling approach for determining ‘proportion of area occupied’, we countered these shortcomings through study design and interpreting the occupancy parameter, psi, as ‘probability of habitat used’. Probability of habitat use was modeled for each target species using single state or multistate models. A combination of the estimated probabilities of habitat use for jaguar and prey was selected to identify the final jaguar corridor. This protocol provides an efficient field methodology for identifying corridors for easily-identifiable species, across large study areas comprised of unprotected, private lands.     

Local perceptions of jaguars (Panthera onca) and pumas (Puma concolor) in the Iguaçu National Park area, south Brazil

Jaguars (Panthera onca) have been killed by local residents within the boundaries and lands surrounding Iguaçu National Park (INP), Brazil. Both jaguars and pumas (Puma concolor) occur in the region, however, livestock predation by pumas has rarely been reported. Our objective was to assess the local perceptions about jaguars and pumas. We identified two major factors that distinguished the perceptions towards the two species: less people feared the puma than the jaguar; and most people believed that jaguars, but not pumas, were released into INP by local authorities. Interestingly, despite those major differences in these perceptions, feelings towards the two species tended to be the same. Perceptions towards jaguars were not influenced by the predation history of the properties, suggesting that the predation impact was not remarkable enough to influence local perceptions towards carnivores. This is apparently the first study on local perceptions towards large carnivores in Brazil.     

Spatial organization and food habits of jaguars (Panthera onca) in a floodplain forest

For most carnivore populations, territoriality is the regulating social system ultimately determined by food abundance and/or strife. However, in some food-based territorial felid species such as the jaguar (Panthera onca), the influence of food availability on territoriality remains unclear and may be lessened because of a tendency for individuals to occur at high densities across the landscape. We examined spatial organization and use of food in a population of jaguars in the southern region of the Pantanal, Brazil (2003-04). We predicted that if territoriality plays an important role in determining jaguar population dynamics, exclusive use of territories should be observed and thereby influence prey selection patterns. We determined that home range sizes were comparable between sexes and overlapped little at the core area level. Line transect surveys revealed that large mammals comprised the bulk of available wild prey for jaguars, and scat analysis indicated that jaguars relied mostly on large mammalian prey. The most common wild species killed by jaguars were capybara (Hydrochaeris hydrochaeris) and caiman (Caiman yacare). We estimated that the wild prey base was adequate to support the jaguar population. Larger wild prey species were considerably more likely to die from predation than from other causes, and predation was more likely to occur in jaguar core areas than in areas of home range overlap. Modest cattle depredation rates had little demographic importance to the local jaguar population. We conclude that spacing patterns in the local jaguar population were likely based on exclusion through territoriality rather than food limitation.     

Protecting more than the wetland: The importance of biased sex ratios and habitat segregation for conservation of the Hine’s emerald dragonfly, Somatochlora hineana Williamson

Within species habitat use may depend on age, season or sex of an individual. The distribution of males and females may vary both temporally and spatially due to differences in the costs of reproduction and the distribution of critical resources. Conservation of a species requires knowledge of the habitat use of both sexes in order to predict the population size and protect all habitats that a species requires. Adult dragonfly populations often have highly male-biased sex ratios at the breeding habitat. This bias has been attributed to females using alternative habitats to avoid male harassment, or to high female mortality. We monitored adult Hine’s emerald dragonfly (Somatochlora hineana Williamson) populations, in breeding and non-breeding habitats in Door County, Wisconsin and found significant differences in habitat use between males and females. Males primarily used wetland habitats, while females primarily used dry meadows and marginal breeding habitats, only coming into wetlands to lay-eggs or find mates. We assessed food resources in the different habitats and found that high quality insect prey (primarily adult Diptera) were more available in the wetland habitat, indicating that these areas were likely a more productive foraging area for adult dragonflies. The fact that females appear to avoid the wetland habitat is consistent with the hypothesis that male harassment alters female distribution patterns. Consideration of the patterns of habitat use by S. hineana indicates the need to develop a broader understanding of the importance of non-wetland areas in the conservation of wetland species.     

Estimating the density of a jaguar population in the Brazilian Pantanal using camera-traps and capture-recapture sampling in combination with GPS radio-telemetry

The jaguar (Panthera onca) is the largest feline in the Americas and third largest world-wide, smaller in size only to the tiger (P. tigris) and lion (P. leo). Yet, in comparison, relatively few studies on jaguar population densities have been conducted and baseline data for management purposes are needed. Camera trapping and capture-recapture sampling methods were used to estimate the size of a jaguar population in the Pantanal’s open wet grassland habitat, an important area for the long-term survival of the species. This study is the first jaguar population estimate conducted in co-operation with a GPS-telemetry study providing an important opportunity for comparing different methods of density estimation. An accessible area within a 460 km² privately-owned ranch was sampled with equal effort during the dry seasons of 2003 and 2004. Thirty-one and twenty-five individual jaguars were identified in 2003 and 2004, respectively. Estimates of jaguar abundance were generated by program CAPTURE. Density estimates were produced according to different methods used to calculate the effectively sampled areas which ranged from 274 to 568 km². For 2003, the currently-used mean maximum distance moved (MMDM) method produced a density of 10.3 jaguars/100 km², while GPS-telemetry-based calculations produced a mean density of 6.6 jaguars/100 km². For 2004, the MMDM method produced an estimate of 11.7 jaguars/100 km² while GPS-telemetry calculations produced a density of 6.7 jaguars/100 km². Our results suggest that the widely-used MMDM method used to calculate effectively sampled areas is significantly under-reflecting maximum distances moved by jaguars and their range-use and, thereby, considerably inflating cat density estimates. This overestimation could place a population in a difficult situation by lengthening the time taken to initiate protection measures because of underestimating the risk to that population.     

Conserving tigers Panthera tigris in selectively logged Sumatran forests

The response of most large carnivores to selective logging is poorly understood. On the one hand, selective logging may represent loss of important habitat, yet, on the other hand, selective logging may increase browse availability for a terrestrial ungulate prey base, thereby indirectly benefiting large carnivores. Using a camera trap-based sampling method, we estimate tiger density in two primary-selectively logged forest areas that straddle Kerinci Seblat National Park, Sumatra. We then investigate potential differences between the habitat use of tigers: within these study areas and forest types; and, within the finer-scale landscape features associated with these covariates. Across the mixed forest study areas, tiger density estimates (adult individuals/100 km² ± S.E.) of 2.95 ± 0.56 and 1.55 ± 0.34 were produced. However, within these areas, tigers showed a preference for primary over degraded forest, and this was related to the greater accessibility of degraded forest sites to people, e.g., through their proximity to roads. Presently, the majority of Sumatran tigers occur within large tracts of primary forest, but these extend outside of the island’s protected area borders, and these unprotected forests are especially at risk from the high levels of deforestation in Sumatra. As forest is cleared, previously remote, and therefore safer, tracts of primary forest become accessible and, eventually, degraded. Yet, from our study, degraded forest in combination with primary forest supported sufficiently high tiger densities and can, therefore, make an important contribution to tiger conservation. It is therefore essential to lessen the detrimental effects of accessibility through increasing law enforcement and destroying ex-logging roads.     

Grizzly bear response to human development and activities in the Bow River Watershed, Alberta, Canada

Few studies have reported the effects of multiple human activities on grizzly bears, Ursus arctos. We document the degree of grizzly bear response to various human developments as a function of multiple interacting variables based on observed median distances to roads, trails and development features in a landscape where human presence is widespread. Female grizzly bears remained further than males from paved roads regardless of habitat quality or time of day. Males were found closer to paved roads when within or adjacent to high quality habitat and during the period of least human activity. The combination of traffic volume and highway configuration, however, overrides a bear’s attraction to high quality habitats for high-speed, high-volume, highways. Avoidance of busy transportation corridors was strongest in the adult segment of the population. Bears were found closer to trails during the human inactive period when within high quality habitat and further from trails when distant to high quality habitat. Our data indicated an inverse relationship between the sexes in response to vehicles and traffic noise compared to the response to human settlement and encountering people. Female bears were found further away than males in relation to vehicles and traffic noise, yet found closer than males to human settlement and places where people may be encountered. Those males that were more willing to exploit high quality habitat near roads, did so at night and where hiding cover was present. Adult females were the most risk-averse cohort, choosing to avoid humans instead of seeking out high quality habitats. Adult female grizzly bears were influenced most by human activities and development. Management agencies must maintain access to high quality habitat, especially for adult females, and create new opportunities to support the reproductive potential of the population.     

Linking resource use with demography in a synanthropic population of common ravens

A thorough understanding of a population’s ecology requires knowledge of the relationship between habitat use, resource use and demographic parameters. We undertook an empirical investigation of habitat use, resource use and demography in a population of common ravens (Corvus corax), a species widely distributed throughout the Northern Hemisphere. The abundance of ravens is increasing in many parts of western North America, which represents a conservation concern since predation by ravens is thought to contribute to the decline of several sensitive species. We defined resources as the suite of physical and biological components in the environment that led to occupancy of a particular place by ravens. The home ranges of breeding and nonbreeding ravens contained similar proportions of resources, but breeding ravens used more edges, roads, forest, clearcuts, and towns than nonbreeders. We detected no differences in survival between the sexes, but breeding ravens survived at higher rates than nonbreeders, due to exclusion by breeding ravens from those resources positively associated with survival. Raven use of mature forests and anthropogenic land use types was positively associated with survival. Breeding raven use of clearcuts and patchy areas contributed to increased reproduction, but the use of clearcuts along with the use of roads was negatively associated with survival due to illegal shooting. Greater insight into the demography of synanthropic species such as the common raven will enable managers to make informed decisions for protecting biodiversity. This study is the first to consider the demographic consequences of habitat use and resource use for both nonbreeding and breeding common ravens.     

Jaguars, pumas, their prey base, and cattle ranching: ecological interpretations of a management problem

Jaguar and puma depredation on livestock may be influenced by (1) innate and learned behavior; (2) health and status of individual cats; (3) division of space and resources among jaguar and puma; (4) cattle husbandry practices; and (5) abundance and distribution of natural prey. Our study in Los Llanos of Venezuela aimed to establish how all these elements related to cattle being lost to cat depredation. Prey distribution was influenced by forest composition, topographical characteristics, and degree of habitat interspersion. The biomass of natural prey in the study area was adequate to support the resident large cats without a subsidy of livestock. Selective rather than opportunistic hunting by the cats reinforced that conclusion. Puma were responsible for more attacks on livestock than jaguar, frequently in maternity pastures in upland areas of relatively low prey availability. Management recommendations are discussed that may be relevant to other savanna/forest mosaics of South America.     

Eucalypt plantations as habitat for birds on previously cleared farmland in south-eastern Australia

In parts of Australia, extensive areas of cleared land are now being planted with commercial plantations of native eucalypts. Questions arise about the extent to which such plantations can rectify previous loss of habitat and contribute to biodiversity conservation. This study assessed abundance of bird species (as one aspect of biodiversity) on 105 sites (25 cleared land, 58 plantations and 22 native forest) in two regions of rural Victoria, south-eastern Australia. Generalised linear modelling was used to assess some of the landscape and habitat variables that contributed to the value of plantation sites for particular groups of bird species. These models demonstrated the importance of on-site habitat variables in explaining the abundance of groups of bird species, with landscape context making small additional contributions.Mean abundance of forest and woodland birds was higher in eucalypt plantations than cleared farmland, and marginally lower than in native forest. Patterns differed between bird guilds. For example, insectivores that forage in the canopy and tall shrub layers were at least as common in plantations as in native forest, with birds in the latter group using young eucalypts as if they were tall shrubs. Birds that forage from open ground among trees were more common in plantations than native forest, and may benefit substantially from the new habitat fortuitously provided for them. This group includes several species that have declined in natural woodland habitats. Nectarivores, carnivores and birds that forage among low shrubs were less common in plantations than in native forest. Insectivores that forage from eucalypt bark made little use of plantations. Different approaches to plantation design and management would be needed to cater for groups such as these. Specific measures include planting of rough-barked eucalypts in addition to smooth-barked species, and provision of artificial hollows. Retention of existing remnants of native forest (e.g. old trees and forest patches) is a priority, to supply habitat elements that would otherwise be missing for long periods.     

Using occupancy models of forest breeding birds to prioritize conservation planning

As urban development continues to encroach on the natural and rural landscape, land-use planners struggle to identify high priority conservation areas for protection. Although knowing where urban-sensitive species may be occurring on the landscape would facilitate conservation planning, research efforts are often not sufficiently designed to make quality predictions at unknown locations. Recent advances in occupancy modeling allow for more precise estimates of occupancy by accounting for differences in detectability. We applied these techniques to produce robust estimates of habitat occupancy for a subset of forest breeding birds, a group that has been shown to be sensitive to urbanization, in a rapidly urbanizing yet biological diverse region of New York State. We found that detection probability ranged widely across species, from 0.05 to 0.8. Our models suggest that detection probability declined with increasing forest fragmentation. We also found that the probability of occupancy of forest breeding birds is negatively influenced by increasing perimeter-area ratio of forest fragments and urbanization in the surrounding habitat matrix. We capitalized on our random sampling design to produce spatially explicit models that predict high priority conservation areas across the entire region, where interior-species were most likely to occur. Finally, we use our predictive maps to demonstrate how a strict sampling design coupled with occupancy modeling can be a valuable tool for prioritizing biodiversity conservation in land-use planning.     

A rapid approach to modeling species-habitat relationships

A basic element in the success of managing species of conservation concern is knowledge of the species’ habitat occupancy. Often, predictive species-habitat models are developed from GIS data sources that were intended for purposes other than predicting species habitat occupancy and are of inappropriate scale. In addition, the techniques used to quantify predictor variables from such data sources are often time consuming and cannot be repeated efficiently to reflect changing conditions. We used digital orthophotos and a grid cell classification scheme to develop an efficient technique to quantify predictor variables to model Florida scrub-jay habitat occupancy. We combined our classification scheme with a priori hypothesis development using expert knowledge and a previously published habitat suitability index and used an objective model selection procedure to choose candidate models. We classified a large area (43,000 ha) in a fraction of the time that would be required to map vegetation classes and were able to test models at varying scales using a grid-cell windowing process. Interpretation of the selected models confirmed existing knowledge of factors important to Florida scrub-jay habitat occupancy. The potential uses and advantages of using a grid cell classification scheme in conjunction with expert knowledge and an objective model selection procedure are discussed.     

A range-wide model of landscape connectivity and conservation for the jaguar, Panthera onca

Large, wide-ranging carnivores face greater threats and more persistent declines than most other mammal species. An important conservation tool for these carnivores has been range-wide priority-setting exercises that have helped identify critical threats and key populations. However, such exercises often fail to identify functional movement corridors or account for genetic connectivity. We present a new model for jaguar (Panthera onca) conservation that uses a geographic information system (GIS) and expert input to create a dispersal cost surface and identify least-cost corridors connecting the 90 known populations across the jaguar’s range. Results indicate 78% of historic jaguar range, an area of approximately 14.9 million km², still holds potential for jaguar movement and dispersal. We identified 182 potential corridors between populations, ranging from 3 to 1607 km in length; 44 of these corridors are characterized as being of immediate concern due to their limited width, and thus their high potential for being severed. Resultant maps, displaying priority populations and corridors, are used to direct field-based research and conservation efforts. Field assessment and refinement of the corridors is ongoing. This is the first attempt to create and implement such a holistic model of range-wide conservation for a large carnivore species.     

Cattle depredation by puma (Puma concolor) and jaguar (Panthera onca) in central-western Brazil

In this study, data on cattle depredation by puma (Puma concolor) and jaguar (Panthera onca) were recorded for six years (1998-2003) in a cattle ranch in central-western Brazil. Depredation represented 18.9% of the overall cattle mortality, being predominant on calves. In biomass, kills represented 0.4% (63.8 kg/km²) of the ranch’s annual stock. In economic loss, kills represented 0.3% of the cattle stock value. Depredation was mainly associated with cattle’s age class and location along with the time of birth of calves. The proportion of pastures next to forest with depredation (n = 33, 48.5%) was not distinguished to the proportion of pastures not bordering forest with depredation (n = 35, 51.5%). However, the proportion of pastures next to forest with depredation represented 54% (n = 33) of the 61 total pastures that were at least partially surrounded by forest patches or riparian forests that comprised eight continuum blocks of forest fragments of different sizes in the ranch and adjacent areas. No kills occurred in the central portion (main house) of the farm, close to the headquarters where the pastures not bordering forest. The distances of the kills in relation to areas of native forest was 1317.48 ± 941.03 m. In order to reduce depredation, calves should be kept as far as possible from forest areas and concentrated cattle breeding and calving seasons should be encouraged.     

Population ecology of the Chinese grouse, Bonasa sewerzowi, in a fragmented landscape

We report the first population ecology study of the endangered Chinese grouse (Bonasa sewerzowi). The study population was stable during 1995-2000. Annual rates of survival were 0.64 for adults and only 0.17 for chicks to 13 weeks of age. Reproductive parameters were 6.1 eggs/clutch, 63% nest success, and overall 3.6 chicks hatched/female/year. The spring sex ratio was 64% males, with 44% of the males unmated in spring. Local people found an estimated 10-29% of the nests and took the eggs for food. Only about 9% of the reserve containing the study area is mature mixed forest, the preferred habitat. Inadequate brood habitat may have influenced chick survival and the skewed adult sex ratio may be a result of habitat fragmentation. We recommend protecting the remaining mixed forest, creating more brood habitat, and stopping the taking of eggs.     

Sex differences in population genetics, home range size and habitat use of the parti-colored bat (Vespertilio murinus, Linnaeus 1758) in Switzerland and their consequences for conservation

When habitats are declining, niche segregation by demographic groups, such as the two sexes, can have a profound impact on the extinction risk of a species as a whole. Thus, differences in the requirements of demographic groups are of importance in conservation. We combined behavioural and genetic data to investigate whether the sexually segregated parti-colored bat (Vespertilio murinus) exhibits sex-specific niche partitioning. We use our data to evaluate implications for conservation of this potentially vulnerable species in Switzerland, the western boundary of its range. Using radio-telemetry, we found sex-specific differences in habitat use. Foraging females strongly relied on lakes while foraging males displayed more flexibility in their habitat use. Moreover, males covered significantly larger foraging areas than females. Sequencing 341 base pairs of the mitochondrial D-loop of 247 individuals revealed sex-specific differences in the genetic structure of colonies, but no such difference was observed for three nuclear micro-satellite markers. We found high mtDNA diversity in two Swiss male colonies and one German female colony, but low mtDNA diversity in two Swiss female colonies. Our genetic data suggest that considerable gene flow occurs via male dispersal and mating. At the same time immigration of females into the existing female colonies in Switzerland is rare compared to the immigration of new males into male colonies. Since we found the sexes in Vespertilio murinus to differ markedly in their ecology, population genetics, and behaviour, we conclude that sex-specific conservation plans are required to protect this species efficiently.     

Sex-specific roost movements and population dynamics of the vulnerable long-fingered bat, Myotis capaccinii

The roosts of many IUCN-listed cave-roosting bat species are under threat from tourist development in SE Europe and other regions of the world. Much-needed conservation strategies require, among other information, an understanding of their roost movements and population dynamics, which can now be obtained relatively quickly using advanced models. We have studied the long-fingered bat, Myotis capaccinii, an obligate cave-dweller, in Dadia National Park, Greece. The species formed colonies of up to a few thousand individuals and was highly mobile, frequently switching summer roosts up to 39 km apart, even during late pregnancy. The bats migrated to distant hibernacula including a cave in Bulgaria 140 km NW of the Park. Adult recapture probabilities varied with season and sex: low female recapture rates in autumn, relative to spring and summer, indicated non-random temporary emigration following nursery colony dispersal. The opposite pattern was seen in males: increasing recapture rates in the autumn suggest that males gather in these roosts to mate with females in transit. Adult survival (0.86-0.94) was similar in females and males, similar in winter and summer, and comparable to recent estimates for other bats based on similar modelling techniques. Sex-based differences in juvenile recapture suggest female philopatry and male-biased dispersal. Our work shows that protection of M. capaccinii roosts must extend beyond the Park’s and indeed the country’s boundaries: its conservation requires large-scale, trans-national integrated conservation plans. Our results will apply to many other warm-temperate species with similar life history cycles.     

Habitat use of ponies and cattle foraging together in a coastal dune area

Grazing by large ungulates has been chosen as a management tool in scrub-dominated dune reserves at the Belgian coast. Due to morphological and physiological differences between cattle and ponies, differences in foraging behaviour and habitat use are expected, and these may result in a different impact on the spatially heterogeneous and nutrient-limited ecosystem. Grazing behaviour and habitat use of Shetland ponies and Highland cattle, grazing together in a coastal dune area (60 ha) were investigated at various levels of the foraging hierarchy (habitat, vegetation type, sward height and diet). Habitat use overlap was high in all seasons; both cattle and pony spent most of their grazing time in the grass-dominated habitat. However, Shetland ponies concentrated their grazing activity more on the grass-dominated habitat than did cattle. Cattle spent a greater proportion of their grazing activity in woodland and scrub, compared to the ponies. Foraging activity in woodland and scrub is strongly influenced by season. Within the grass-dominated habitat both species preferred foraging in the grasslands and avoided open vegetation and moss dunes. Within the grasslands, cattle grazed less on the short swards than did ponies. Both cattle and ponies predominantly foraged on graminoids, though there are minor differences between both species and among seasons. Browsing of woody plants occurred only by cattle. Where grazing management has been implemented to maintain dune grasslands and to avoid further invasion by scrub, a combination of cattle and ponies appears to be adequate. Ponies are suitable for maintaining grasslands, but they have no impact on invading scrub. Cattle have an impact on scrub development, both by direct consumption of various shrub species and by opening initially closed scrub.