Who Bites the Bullet First? The Susceptibility of Leopards Panthera pardus to Trophy Hunting

Reliable data is fundamentally important for managing large carnivore populations, and vital for informing hunting quota levels if those populations are subject to trophy hunting. Camera-trapping and spoor counts can provide reliable population estimates for many carnivores, but governments typically lack the resources to implement such surveys over the spatial scales required to inform robust quota setting. It may therefore be prudent to shift focus away from estimating population size and instead focus on monitoring population trend. In this paper we assess the susceptibility of African leopards Panthera pardus to trophy hunting. This has management ramifications, particularly if the use of harvest composition is to be explored as a metric of population trend. We explore the susceptibility of different leopard age and sex cohorts to trophy hunting; first by examining their intrinsic susceptibility to encountering trophy hunters using camera-traps as surrogates, and second by assessing their extrinsic susceptibility using photographic questionnaire surveys to determine their attractiveness to hunters. We show that adult male and female leopards share similar incident rates to encountering hunters but adult males are the most susceptible to hunting due to hunter preference for large trophies. In contrast, sub-adult leopards rarely encounter hunters and are the least attractive trophies. We suggest that our findings be used as a foundation for the exploration of a harvest composition scheme in the Kwazulu-Natal and Limpopo provinces where post mortem information is collected from hunted leopards and submitted to the local provincial authorities.     

Movement,activity and hunting behaviour of leopards in the Cederberg mountains,South Africa

Few data are available on the behaviour of leopards in the absence of competing large predators and human impact, both of which are believed to influence leopard activity and movements. Remote camera traps and global positioning system (GPS) collars were used to quantify leopard activity in the Cederberg Mountains, seasonal and sexual differences in their movements, and determine whether nocturnal hunting success was related to lunar activity. Seventy‐seven per cent of camera‐trap photographs were at night, with a strong male bias (69%) in captures. Daily displacement using one location per day suggested that males moved significantly further than females. However, multiple locations (≥6 per day) showed no difference because males moved in a more linear fashion, but not further each day, than females. In the Cederberg Mountains, an open rocky habitat with low human impact and no competing predators, leopards were predominantly nocturnal, mainly hunting diurnal prey species. Hunting success was low: leopards travelled long distances between kills, with nocturnal hunting success higher on darker nights.     

Hunting behaviour in West African forest leopards

The leopard (Panthera pardus) is a major predator of mammals within the rainforest ecosystem of West Africa. Most of the available information on leopard hunting behaviour comes from studies conducted in open savannah habitats, while little is known about forest leopards. Our radio‐tracking data and scat analysis show that forest leopards differ in various ways from the savannah populations. Forest leopards are diurnal and crepuscular hunters who follow the activity pattern of their prey species. They exhibit seasonal differences in activity patterns, and they develop highly individualized prey preferences. These findings challenge the widespread notion of leopards as opportunistic nocturnal predators.     

Insights into the Management of Large Carnivores for Profitable Wildlife-Based Land Uses in African Savannas

Large African predators, especially lions (Panthera leo) and leopards (Panthera pardus), are financially valuable for ecotourism and trophy hunting operations on land also utilized for the production of other wildlife species for the same purpose. Predation of ungulates used for trophy hunting can create conflict with landholders and trade off thus exists between the value of lions and leopards and their impact on ungulate populations. Therefore productionist and conservation trade-offs are complexly graded and difficult to resolve. We investigated this with a risk-benefit analysis on a large private wildlife production area in Zimbabwe. Our model showed that lions result in substantial financial costs through predation on wild ungulates that may not be offset by profits from hunting them, whereas the returns from trophy hunting of leopards are projected to exceed the costs due to leopard predation. In the absence of additional income derived from photo-tourism the number of lions may need to be managed to minimize their impact. Lions drive important ecological processes, but there is a need to balance ecological and financial imperatives on wildlife ranches, community wildlife lands and other categories of multiple use land used for wildlife production. This will ensure the competitiveness of wildlife based land uses relative to alternatives. Our findings may thus be limited to conservancies, community land-use areas and commercial game ranches, which are expansive in Africa, and should not necessarily applied to areas where biodiversity conservation is the primary objective, even if hunting is allowed there.     

Calibrating a camera trap–based biased mark–recapture sampling design to survey the leopard population in the N'wanetsi concession,Kruger National Park,South Africa

Estimating large carnivore abundance can be challenging. A biased leopard (Panthera pardus) population survey was conducted in the N'wanetsi concession in the Kruger National Park (KNP), South Africa, using motion‐sensitive camera traps from April to August 2008. Survey effort included 88 trapping occasions and 586 trap days. The survey yielded 24 leopard photographs, comprising fourteen adults of eleven males and three females. The capture rate was determined to be 24.4 trap days per leopard. Estimates of population abundance stabilized at approximately 500 trap days. Precision of population estimates began to stabilize after 378 trap days. We estimated that there were nineteen leopards in an area of 150 km². Leopard density was estimated at 12.7 leopards per 100 km². We explore the possibility of employing the methods used in this study to survey the leopard population in the KNP and surrounding areas.     

Intraspecific interactions in a high‐density leopard population

Although less studied than interspecific interactions, interactions among members of the same species can influence space use and temporal activity. Using techniques commonly applied to the analysis of interspecific interactions—multispecies occupancy modeling and the analysis of temporal activity patterns—we studied intraspecific interactions within a high‐density population of Persian leopards (Panthera pardus saxicolor) in Tandoureh National Park, northeastern Iran. Using camera‐trap data, we investigated spatiotemporal interactions between male leopards, lone female leopards, and families (cubs/females with cubs). While we hypothesized that male and female leopards would display different temporal activity patterns, we did not predict spatial avoidance between these groups. We also predicted that leopard families would exhibit spatiotemporal avoidance from male leopards due to the risk of infanticide. Contrary to our expectations, we did not find any evidence for spatial or temporal avoidance between leopard families and adult male leopards. Male and lone female leopards exhibited positive pairwise co‐occurrence, consistent with reports of high overlap between male and female leopard home ranges. While a high level of overlap in temporal activity patterns was found between males/lone females and males/families, there was evidence for variation in the proportion of time each leopard group was active in particular periods of the diel cycle. Male leopards showed cathemeral activity, while lone females and families were more active during daylight hours. The application of these techniques to interactions within a species has improved understanding of the ecology and behavior of this endangered solitary carnivore.     

Selectivity of harvesting differs between local and foreign roe deer hunters: trophy stalkers have the first shot at the right place

Harvesting represents a major source of mortality in many deer populations. The extent to which harvesting is selective for specific traits is important in order to understand contemporary evolutionary processes. In addition, since such data are frequently used in life-history studies, it is important to know the pattern of selectivity as a source of bias. Recently, it was demonstrated that different hunting methods were selected for different weights in red deer (Cervus elaphus), but little insight was offered into why this occurs. In this study, we show that foreign trophy stalkers select for larger antlers when hunting roe deer (Capreolus capreolus) than local hunters, but that close to half of the difference in selectivity was due to foreigners hunting earlier in the season and in locations with larger males. The relationship between antler size and age was nevertheless fairly similar based on whether deer was shot by foreign or local hunters.     

Leopard diets and landowner perceptions of human wildlife conflict in the Soutpansberg Mountains,South Africa

Human-wildlife conflict between carnivores and livestock and game owners is an issue of high conservation concern and has led to the global decline of many large carnivore species. Research has shown that carnivores are often blamed for higher levels of predation of livestock and game than actually occurs and this often leads to retaliatory killing. The aim of this study was to obtain information via scat analysis on the range of prey species taken by leopards in the Soutpansberg Mountains, South Africa, and combine these data with self-reported accounts of livestock predation from local landowners to examine differences between real and perceived leopard predation. Results showed that despite landowners reporting frequent events of leopard predation of livestock and introduced farmed game across the Soutpansberg farming community, no evidence of these species were found in leopard diets. The most frequently eaten species by relative biomass were bushbuck, hyrax and vervet monkeys; in contrast, the farmers reported cattle and impala as often being taken by leopards. Despite sharing the landscape with domestic cattle and introduced game, leopards in the Soutpansberg do not frequently utilise these species as prey and instead focus their diets on wild species. Human-carnivore conflict can be reduced by overcoming the mismatch between actual and perceived levels of predation via landowner education, effective anti-predation measures, an improved government response to reports of livestock predation and potentially giving economic value to problem animals via trophy hunting.     

Assessing trophy hunting in South Africa by comparing hunting and exporting databases

Trophy hunting constitutes a major part of the global wildlife tourism industry and is connected through the export of kills to international wildlife trade. Inconsistencies between kills and exports can contribute to identifying illegal trophy hunting that constitutes a major threat to biodiversity conservation. This paper quantitatively analyses to what extent the data of trophy hunting kills and of trophy exports are consistent using the example of South Africa. Data was extracted from two different sources for the year 2018. These sources were trade data reported under the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) and its Trade Database on the export of trophy items from South Africa, which is compared with the South African Professional Hunting statistics (SAPHs) containing trophy kills information for each species and the country of origin of the hunter. The data of trophy hunting kills and data of trophy hunting export was found to differ to varying degrees across the countries contributing to trophy hunting and CITES-listed trophy-hunted species. We found that both databases report hunting of the same 28 threatened taxa. On the other hand, the same data reports that hunters of 64 nationalities participated in a total of 4,726 trophy kills, while only 3,131 hunting trophy items were exported from South Africa, to 37 countries as final destinations. Among the possible reasons for the discrepancies found, we suggest that the time required to taxidermize trophy specimens may delay the items being addressed to their final destination, in addition to the dual citizenship of some hunters that should also have some influence on our results, as well as the important commercial destination of Brussels Airport. The USA show the highest absolute number of trophy hunters, followed by Spain, but Denmark is the highest driver per capita. Therefore, greater participation of these countries in conservation policies for endangered species is necessary. Additionally, a more detailed differentiation of the term “trophy” to more specific terms such as claws, skins, skulls, etc. may improve reporting systems to easier identify illegal activities related to hunting.     

Does selection on horn length of males and females differ in protected and hunted populations of a weakly dimorphic ungulate?

Weaponry in ungulates may be costly to grow and maintain, and different selective pressures in males and females may lead to sex‐biased natural survival. Sexual differences in the relationship between weapon growth and survival may increase under anthropogenic selection through culling, for example because of trophy hunting. Selection on weaponry growth under different scenarios has been largely investigated in males of highly dimorphic ungulates, for which survival costs (either natural or hunting related) are thought to be greatest. Little is known, however, about the survival costs of weaponry in males and females of weakly dimorphic species. We collected information on horn length and age at death/shooting of 407 chamois Rupicapra rupicapra in a protected population and in two hunted populations with different hunting regimes, to explore sexual differences in the selection on early horn growth under contrasting selective pressures. We also investigated the variation of horn growth and body mass in yearling males (n = 688) and females (n = 539) culled in one of the hunted populations over 14 years. The relationship between horn growth and survival showed remarkable sexual differences under different evolutionary scenarios. Within the protected population, under natural selection, we found no significant trade‐off in either males or females. Under anthropogenic pressure, selection on early horn growth of culled individuals showed diametrically opposed sex‐biased patterns, depending on the culling regime and hunters’ preferences. Despite the selective bias between males and females in one of the hunted populations, we did not detect significant sex‐specific differences in the long‐term pattern of early growth. The relationship between early horn growth and natural survival in either sex might suggest stabilizing selection on horn size in chamois. Selection through culling can be strongly sex‐biased also in weakly dimorphic species, depending on hunters’ preferences and hunting regulations, and long‐term data are needed to reveal potential undesirable evolutionary consequences.     

Hunters and hunting patterns in part of the Kaimanawa Recreational Hunting Area

Abstract

A postal survey of 156 hunters who used the three southeastern-most hunting blocks in the Kaimanawa Recreational Hunting Area (RHA) in 1986 and 1987 achieved a 74% response rate. Comparison of age distributions from this and earlier hunter surveys confirmed an apparent decline in recruitment of young hunters entering the sport during the 1980s. Most hunters had considerable experience (>10 years) and were motivated not only by hunting success but also by the aesthetic value of the outdoor experience. The prime motivation for hunting in the Kaimanawa RHA was the presence of sika deer, and more than two-thirds of the hunters came from outside the Tongariro/Taupo Conservancy. Hunting pressure over current sika deer range in the central North Island may decline if this species continues to disperse naturally or through illegal liberations to other areas. Most (80%) hunters felt that deer densities were acceptable and were satisfied with the present unrestricted hunting system. Fewer than half (41%) the respondents were in favour of active management in the Kaimanawa RHA. Suggestions for active management focused on increasing hunter safety and increasing hunting success rates. Deer densities, as indicated by faecal pellet counts, were inversely related to hunting pressure. Variation in hunting pressure between areas was caused mainly by differences in accessibility. This indicates some potential for manipulation of deer density through options such as increasing access to remote areas or restrictions on hunting pressure.     

泰国Khao Yai国家公园豹猫和云豹的生态和保护

于1997年10月到1999年10月在泰国KhaoYai国家公园对两种同域分布的物种（即豹猫和云豹）进行了研究。使用带诱饵的陷阱捕捉了这些动物,进行麻醉,确定了性别、年龄,并戴上了无线电项圈,对6只雌性豹猫和4只雄性豹猫进行了1—18个月的无线电追踪。豹猫在干旱季节的利用区域大于雨季,雄性利用的区域大于雌性。在夜间和晨昏时节,豹猫活动增加,但是并没有节律性活动。所有豹猫个体在旱季和雨季的活动都相似,但雄性个体的白天活动多于雌性。豹猫的行走距离有性别差异但是没有季节差异;其食物以鼠类为主。分别对1只雌性云豹和1只雄性云豹进行了17个月和7个月的无线电追踪,雌性个体的活动面积为39．4km^2,而雄性个体活动面积为42．2km^2,核心区都是2．9km2。云豹对半绿林的使用大于其它类型的植被,在晨昏和夜间的活动增加,其活动无节律。     

Density-Dependent Natal Dispersal Patterns in a Leopard Population Recovering from Over-Harvest

Natal dispersal enables population connectivity, gene flow and metapopulation dynamics. In polygynous mammals, dispersal is typically male-biased. Classically, the ‘mate competition’, ‘resource competition’ and ‘resident fitness’ hypotheses predict density-dependent dispersal patterns, while the ‘inbreeding avoidance’ hypothesis posits density-independent dispersal. In a leopard (Panthera pardus) population recovering from over-harvest, we investigated the effect of sex, population density and prey biomass, on age of natal dispersal, distance dispersed, probability of emigration and dispersal success. Over an 11-year period, we tracked 35 subadult leopards using VHF and GPS telemetry. Subadult leopards initiated dispersal at 13.6 ± 0.4 months. Age at commencement of dispersal was positively density-dependent. Although males (11.0 ± 2.5 km) generally dispersed further than females (2.7 ± 0.4 km), some males exhibited opportunistic philopatry when the population was below capacity. All 13 females were philopatric, while 12 of 22 males emigrated. Male dispersal distance and emigration probability followed a quadratic relationship with population density, whereas female dispersal distance was inversely density-dependent. Eight of 12 known-fate females and 5 of 12 known-fate male leopards were successful in settling. Dispersal success did not vary with population density, prey biomass, and for males, neither between dispersal strategies (philopatry vs. emigration). Females formed matrilineal kin clusters, supporting the resident fitness hypothesis. Conversely, mate competition appeared the main driver for male leopard dispersal. We demonstrate that dispersal patterns changed over time, i.e. as the leopard population density increased. We conclude that conservation interventions that facilitated local demographic recovery in the study area also restored dispersal patterns disrupted by unsustainable harvesting, and that this indirectly improved connectivity among leopard populations over a larger landscape.     

Evaluation of the pituitary-gonadal response to GnRH, and adrenal status, in the leopard (Panthera pardus japonensis) and tiger (Panthera tigris)

Frequent blood samples were collected to study hormonal responses to GnRH in male and female leopards and tigers. Animals were anaesthetized with ketamine-HCl and blood samples were collected every 5 min for 15 min before and 160 min after i.v. administration of GnRH (1 micrograms/kg body weight) or saline. No differences in serum cortisol concentrations were observed between sexes within species, but mean cortisol was 2-fold greater in leopards than tigers. GnRH induced a rapid rise in LH in all animals (18.3 +/- 0.9 min to peak). Net LH peak height above pretreatment levels was 3-fold greater in males than conspecific females and was also greater in tigers than leopards. Serum FSH increased after GnRH, although the magnitude of response was less than that observed for LH. Basal LH and FSH and GnRH-stimulated FSH concentrations were not influenced by sex or species. Serum testosterone increased within 30-40 min after GnRH in 3/3 leopard and 1/3 tiger males. Basal testosterone was 3-fold greater in tiger than leopard males. LH pulses (1-2 pulses/3 h) were detected in 60% of saline-treated animals, suggesting pulsatile gonadotrophin secretion; however, in males concomitant testosterone pulses were not observed. These results indicate that there are marked sex and species differences in basal and GnRH-stimulated hormonal responses between felids of the genus Panthera which may be related to differences in adrenal activity.     

Patterns of hunting mortality in Norwegian moose (Alces alces) populations

We used a simple life table approach to examine the age-specific patterns of harvest mortality in eight Norwegian moose populations during the last 15 years and tried to determine if the observed patterns were caused by hunter selectivity. The general opinion among local managers is that hunters prefer to shoot female moose not in company with calves to keep a high number of reproductive females in the population (and because of the emotional stress involved in leaving the calf/calves without a mother), and relatively large males because of the higher return with respect to meat and trophy. In support of the former view, we found the harvest mortality of adult females to be higher among pre-prime (1–3 years old) than prime-aged age classes (4–7 years old). This is probably because prime-aged females are more fecund and, therefore, more likely to be in company with one or two calves during the hunting season. As the season progressed, however, the selection pressure on barren females decreased, probably due to more productive females becoming ‘legal’ prey as their calf/calves were harvested. In males, we did not find any evidence of strong age-specific hunter selectivity, despite strong age-dependent variation in body mass and antler size. We suggest that this was due to the current strongly female-biased sex ratio in most Norwegian moose populations, which leaves the hunters with few opportunities to be selective within a relatively short and intensive hunting season. The management implications of these findings and to what extent the results are likely to affect the future evolution of life histories in Norwegian moose populations are discussed.Electronic Supplementary Material Supplementary material is available for this article at and accessible for authorised users.     

Leopard predation and primate evolution

Although predation is an important driving force of natural selection its effects on primate evolution are still not well understood, mainly because little is known about the hunting behaviour of the primates' various predators. Here, we present data on the hunting behaviour of the leopard (Panthera pardus), a major primate predator in the Tai; forest of Ivory Coast and elsewhere. Radio-tracking data showed that forest leopards primarily hunt for monkeys on the ground during the day. Faecal analyses confirmed that primates accounted for a large proportion of the leopards' diet and revealed in detail the predation pressure exerted on the eight different monkey and one chimpanzee species. We related the species-specific predation rates to various morphological, behavioural and demographic traits that are usually considered adaptations to predation (body size, group size, group composition, reproductive behaviour, and use of forest strata). Leopard predation was most reliably associated with density, suggesting that leopards hunt primates according to abundance. Contrary to predictions, leopard predation rates were not negatively, but positively, related to body size, group size and the number of males per group, suggesting that predation by leopards did not drive the evolution of these traits in the predicted way. We discuss these findings in light of some recent experimental data and suggest that the principal effect of leopard predation has been on primates' cognitive evolution.     

Field age determination of leopards by tooth wear

Age determination is an important tool in wildlife studies. Estimating the age of animals in the field using tooth wear criteria may be subject to error as a result of variations between individuals, habitats and populations. Data on age estimation of leopards and tooth wear characteristics are lacking. Nineteen leopards in Namibia were assessed for tooth eruption and wear. Between 1991 and 1995 leopards (including 13 individuals of known age) were monitored at one year intervals ('28 leopard years') to record age and tooth wear. At the age of two years leopards had fully developed dentition. Wear started with the incisors and canines, and spread to the premolars and molars. A chronology of tooth eruption and wear in relation to age is presented. Above the age of three years, male leopards showed higher frequencies of enamel flaking and canine fractures than females.     

Vigorous Dynamics Underlie a Stable Population of the Endangered Snow Leopard Panthera uncia in Tost Mountains,South Gobi,Mongolia

Population monitoring programmes and estimation of vital rates are key to understanding the mechanisms of population growth, decline or stability, and are important for effective conservation action. We report, for the first time, the population trends and vital rates of the endangered snow leopard based on camera trapping over four years in the Tost Mountains, South Gobi, Mongolia. We used robust design multi-season mark-recapture analysis to estimate the trends in abundance, sex ratio, survival probability and the probability of temporary emigration and immigration for adult and young snow leopards. The snow leopard population remained constant over most of the study period, with no apparent growth (λ = 1.08+−0.25). Comparison of model results with the “known population” of radio-collared snow leopards suggested high accuracy in our estimates. Although seemingly stable, vigorous underlying dynamics were evident in this population, with the adult sex ratio shifting from being male-biased to female-biased (1.67 to 0.38 males per female) during the study. Adult survival probability was 0.82 (SE+−0.08) and that of young was 0.83 (SE+−0.15) and 0.77 (SE +−0.2) respectively, before and after the age of 2 years. Young snow leopards showed a high probability of temporary emigration and immigration (0.6, SE +−0.19 and 0.68, SE +−0.32 before and after the age of 2 years) though not the adults (0.02 SE+−0.07). While the current female-bias in the population and the number of cubs born each year seemingly render the study population safe, the vigorous dynamics suggests that the situation can change quickly. The reduction in the proportion of male snow leopards may be indicative of continuing anthropogenic pressures. Our work reiterates the importance of monitoring both the abundance and population dynamics of species for effective conservation.     

Sex ratio variation in harvested moose (<Emphasis Type="Italic">Alces alces</Emphasis>) calves: does it reflect population calf sex ratio or selective hunting?

During the last 30 years, the proportion of males in the calf harvest of moose (Alces alces) in Norway has decreased, indicating a decline in proportions of males recruited to the autumn populations. At the same time, the percentages of exclusive calf hunting permits and of calves shot have increased. The change in calf sex ratio may thus simply be the result of hunter preferences for slightly larger (6.2% higher body mass) male calves combined with fewer opportunities for selective hunting due to increasing hunting quotas of calves. We examined this hypothesis by analyzing the variation in sex, number of siblings, carcass mass, date, and location of kill of 16,330 moose calves harvested during 1970–2004. In the presence of hunting selection for larger calves, we predicted larger proportions of male calves to be harvested in populations with large sexual size dimorphism among calves. Similarly, we expected more males to be harvested from twin than single litters because hunters then can more easily compare twins and select the larger calf, which is more often a male. Increasing proportions of single female calves were also expected to occur in the daily harvest as the accumulated number of harvested calves increased and the proportion of calves left in the population decreased. We found no positive relationship between the proportion of male calves and the level of sexual size dimorphism, no clear difference in sex ratio between harvested single and twin calves, and no increase in the proportion of single female calves as the accumulated number of calves in the harvest increased. This suggests that the spatiotemporal variation in the harvest calf sex ratio in Norway most likely reflects differences in population calf sex ratios prior to the hunting season and not varying degrees of hunting selectivity.     

Hunting activity and aging among the gidra papuans: A biobehavioral analysis

From the individual-based records of hunting practice for 1,633 hr in total and from the hunters' visual acuity and grip strength observed among the Gidra Papuans in lowland Papua New Guinea, this paper aims to analyze the relationship between the hunting activity and aging. The sensorimotor functions determine the range of age in which the individuals act as active or productive hunters: determined to be from the late teen-age years to about 45 years among the present subjects. In this age range, hunting efficiency increases with age. In terms of weight of animals killed per hunting time, the efficiency of the elder married (aged about 35 to 45 years) was almost four times higher than that of the unmarried (16–17 to late 20s). This aging effect is judged to depend on behavioral abilities that increase in accordance with experience and cumulative knowledge. Simultaneously, the comparison of individual hunters' records in 1971–72 and in 1981 reveals that hunting efficiency is associated with the individualities of the hunters.