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Reproduction and social behaviour of the dhole, Cuon alpinus ( Canidae).
Abstract
The home range of a pack of dholes living in the Bandipur Tiger Reserve Karnataka, India, was 40 km2, average density was 0.35-0.9 dholes/km2 in the 20 km2 core area. The hunting range of the pack during the denning period was 11 km2 and at other times >15 km2. Mean number of adults in the pack averaged 8.3 and including pups, the mean pack size was 16. Adult sex ratio consistently favoured males with a 2:1 ratio. Preexisting dens were used. Pups left the den when 70-80 days old and at 7 months of age actively took part in killing prey. Means of communication, types of play, and response to other species are described. Determinants of pack size and features of pack size regulation are discussed and data are compared with those of Canis lupus and Lycaon pictus.-from Author
... Individual dholes weigh 15-20 kg. Packs of 2-25 individuals routinely hunt wild ungulate herbivores that are 5-10 times their body size (Johnsingh, 1982;Venkataraman, Arumugam & Sukumar, 1995;. Studies examining dhole distribution and habitat associations suggest that, although generally restricted to forested Protected Areas, dholes also use unprotected secondary forests, multi-use forest fragments, and agroforests adjoining Protected Areas for movement and dispersal (Jenks et al., 2012;Gangadharan, Vaidyanathan & St. Clair, 2016;Punjabi et al., 2017;Srivathsa et al., 2019aSrivathsa et al., , 2020a. ...
... Leopards are generally nocturnal, although this can vary across locations and habitats (see Karanth et al., 2017). Their reclusive behaviour means they are reported to be socially suppressed by copredators like lions Panthera leo, spotted hyenas Crocuta crocuta, tigers and dholes (Johnsingh, 1982;Venkataraman et al., 1995;Caro & Stoner, 2003). Studies examining prey preferences of leopards across their range suggest a varied diet. ...
... Leopards, for instance, exploit arboreal habitats to escape risky encounters and also to cache their kills (Karanth & Sunquist, 2000). Dholes use dens during the breeding season; these are typically underground or in cavities in rocky habitats that are too small for the two large cats to enter (Johnsingh, 1982). Such avoidance mechanisms are important for enabling copredator coexistence, but unlikely to be captured by studies using spatially static camera-trap data. ...
Intraguild interactions among carnivores have long held the fascination of ecologists. Ranging from competition to facilitation and coexistence, these interactions and their complex interplay influence everything from species persistence to ecosystem functioning. Yet, the patterns and pathways of such interactions are far from understood in tropical forest systems , particularly across countries in the Global South. Here, we examined the determinants and consequences of competitive interactions between dholes Cuon alpinus and the two large felids (leopards Panthera pardus and tigers Panthera tigris) with which they most commonly co-occur across Asia. Using a combination of traditional and novel data sources (N = 118), we integrate information from spatial, temporal, and dietary niche dimensions. These three species have faced catastrophic declines in their extent of co-occurrence over the past century; most of their source populations are now confined to Protected Areas. Analysis of dyadic interactions between species pairs showed a clear social hierarchy. Tigers were dominant over dholes, although pack strength in dholes helped ameliorate some of these effects; leopards were subordinate to dholes. Population-level spatio-temporal interactions assessed at 25 locations across Asia did not show a clear pattern of overlap or avoidance between species pairs. Diet-profile assessments indicated that wild ungulate biomass consumption by tigers was highest, while leopards consumed more primate and livestock prey as compared to their co-predators. In terms of prey offtake (ratio of wild prey biomass consumed to biomass available), the three species together harvested 0.4-30.2% of available prey, with the highest offtake recorded from the location where the carnivores reach very high densities. When reexamined in the context of prey availability and offtake, locations with low wild prey availability showed spatial avoidance and temporal overlap among the carnivore pairs, and locations with high wild prey availability showed spatial overlap and temporal segregation. Based on these observations, we make predictions for 40 Protected Areas in India where temporally synchronous estimates of predator and prey densities are available. We expect that low prey availability will lead to higher competition, and in extreme cases, to the complete exclusion of one or more species. In Protected Areas with high prey availability, we expect intraguild coexistence and conspecific competition among carnivores, with spill-over to forest-edge habitats and subsequent prey-switching to livestock. We stress that dhole-leopard-tiger co-occurrence across their range is facilitated through an intricate yet fragile balance between prey availability, and intraguild and conspecific competition. Data gaps and limitations notwithstanding, our study shows how insights from fundamental ecology can be of immense utility for applied aspects like large predator conservation and management of human-carnivore interactions. Our findings also highlight potential avenues for future research on tropical carnivores that can broaden current understanding of intraguild competition in forest systems of Asia and beyond.
... Another carnivore species in which adult play seems to serve as regulator of social dynamics is the dhole or the Asiatic wild dog (Cuon alpinus) (Fox, 1984;Johnsingh, 1982). Dholes live in fluid packs made up of a variable number of subjects ranging from 5 to 25 individuals depending on the different sites (Durbin et al., 2004). ...
... This tolerant behavior strongly resembles that expressed by wild bonobos when subjects belonging to different communities meet each other (Behncke, 2015). In dholes as well as in African wild dogs, adult-adult play is a ritualized behavior that helps synchronizing mood around hunting activities (Fox, 1984;Johnsingh, 1982). Adult-adult play, which involves same-sex and mixed-sex chasing, ambushing, and mounting, precedes and follows both successful and unsuccessful hunting events. ...
... Adult-adult play, which involves same-sex and mixed-sex chasing, ambushing, and mounting, precedes and follows both successful and unsuccessful hunting events. Moreover, as for spotted hyenas, adults spend time playing with other adults and cubs at the dens (Johnsingh, 1982). Therefore, although few data on social behavior of this species are available, play seems to be an important part of the dholes' life tolerant style: "[…] Play is a way of coming together and staying together for a time, a means of reaffirming friendships and consolidating social bonds. ...
Play is generally considered an immature affair. However, adult play is present in several mammal species living in complex social systems. Here, I hypothesize that adult social play is favored by natural selection in those species characterized by high level of social tolerance and/or by the need of others’ cooperation to reach a goal (i.e., leverage). The integration and comparison of bio-behavioral data on non-human primates and wild social carnivores allows drawing a comprehensive picture on the importance of adult play in facing unpredictable, novel social situations and in overcoming stressful experiences. The ability to cope with potentially competitive interactions through play can favor the emergence of egalitarian societies. A further interesting and beneficial aspect of adult play is its role in synchronizing group activities and favoring collective decision making by renovating the motivation to cooperate in groupmates. As a last step, some considerations about the presence of adult play in the most egalitarian and cooperative human groups (e.g., hunter-gatherer societies) allows discussing the apparent dichotomy between cultural and biological evolution of certain behavioral traits, including social play in adulthood.
... Previous telemetry, behavioral, and genetic studies on dholes have suggested a bias toward female dispersal due to the majority of male-biased packs (Johnsingh, 1982;Venkataraman, 1998;Iyengar et al., 2005), but, to date, no extensive study has been done, comprehending the effects of competition, group size, density, and spatial heterogeneity on dhole dispersal (Miyamoto et al., 2013;Groom et al., 2017). Though the malebiased sex ratio in dhole packs possibly be a consequence of female-biased nature of dhole dispersal (Venkataraman, 1998;Modi et al., 2018), the dispersal pattern of a pack-living animal is further complicated by other variables. ...
... This can be due to the reason that tiger presence is solely correlated with prey-rich sites, while the presence probability of dhole was a trade-off between prey availability and active spatial avoidance of tigers (Steinmetz et al., 2013). The male-biased sex ratio in both populations indicates the possibility of female-biased dispersal, which is further supported by previous ecological studies (Johnsingh, 1982;Venkataraman, 1998). The probability of female-biased dispersal is suggested by the male-biased sex ratio in both populations, which is further confirmed by earlier ecological studies (Johnsingh, 1982;Venkataraman, 1998). ...
... The male-biased sex ratio in both populations indicates the possibility of female-biased dispersal, which is further supported by previous ecological studies (Johnsingh, 1982;Venkataraman, 1998). The probability of female-biased dispersal is suggested by the male-biased sex ratio in both populations, which is further confirmed by earlier ecological studies (Johnsingh, 1982;Venkataraman, 1998). According to the local resource enhancement (LRE) paradigm, research on species that breed cooperatively in hierarchies also shows that the pack tends to favor the more helpful sex (Silk and Brown, 2008). ...
Introduction
Dispersal is a multi-causal, crucial life-history event in shaping the genetic and behavioral structure of mammals. We assessed the dispersal pattern of dholes aka Asiatic wild dog (Cuon alpinus), a social monogamous mammal at two tiger reserves of Maharashtra with different degrees of pack size and competition with tigers i.e. Tadoba-Andhari (TATR, smaller pack size, higher tiger density) and Nawegaon-Nagzira (NNTR, larger pack size, lower tiger density).
Methods
We used the microsatellite data of 174 individual genotypes (98 males and 67 females) to assess the dispersal pattern of dholes from two populations with varying pack size, tiger density, and landscape connectivity using gene flow as a proxy. We compared the population structure, pairwise F statistics, assignment index, and relatedness across a spatial scale.
Results and discussion
Overall, the results suggested a difference in sex-bias dispersal pattern for the two sub-populations, exhibiting significant results for female-biased dispersal in the TATR population with a smaller pack size and higher tiger density. Our study highlights the variability in sex-biased dispersal patterns in two different populations which could be the consequence of different variables such as pack size, tiger density, and geographical scale. The study warrants further quantitative investigation including several factors such as individual behavior, pack composition, pack size, tiger density, etc. In the present Anthropocene era, determining the sex bias in dispersal patterns for a short-range, pack-living carnivore will help in devising an effective conservation management plan for their long-term survival.
... Dholes inhabiting various habitat types in their geographical ranges [64]. In India, dholes use dry deciduous forest and savannah woodlands which interspersed with meadows [48,[67][68][69][70][71], and Dipterocarp Forest [72]. In Thailand, dholes use lowland rainforest [73], mixed evergreen and deciduous forest [74], mountainous semi-evergreen forest [75], and grassland [74] as their habitat. ...
... Cohen et al. (1978) observed that one solitary fawn was killed by a pack of 8 dholes and solitary dholes should be most successful at hunting small prey such as rodents and lagomorphs. A chital fawn can be killed by two dholes, while three dholes were successfully killing a sambar fawn and chital doe [69]. ...
... Female dhole begins to reproduce at the age of three years [89]. Only one individually alpha female received reproductive behaviors from males, despite many females being present in the pack during her tenure [49,69,89]. The mean tenure for females was six years, while the tenure for males was two years [89]. ...
The success of conservation and management of endangered species is highly dependent on the manager's understanding on the biology and ecology of the target species. However, most wildlife management and conservation activities in developing countries are not based on scientific data, but more on the political agenda and intuition of managers. As a result, the impact of conservation and management efforts was far from what was expected and sometimes even caused new problems to arise (e.g., the rapid population decline of certain species, introduction of alien invasive species etc.). Wildlife legal status is also important to protect a certain species from exploitation and to ensure the fairness by regulating social conduct and enforcing policy associated with wildlife management and conservation, and, therefore, it has play an important role in achieving management goal. We attempt to review the management and conservation of dholes (Cuon alpinus) at the human-dominated landscape through ecological and legal approach. Our finding showed that an approach which was mostly used along the history of dhole’s management was lethal control due to livestock depredation. This approach has resulted dramatic decline of dhole’s population and explosion of wild ungulates population. In Indonesia, the government regulation to conserve this species is already exists, but lack of implementation due to management limitations. The absence and lack of baseline data for dhole’s conservation are common phenomenon since this species has not listed as a priority species. To ensure the long-term survival of this species, suitable conservation strategy supported by good scientific data and regulations related to dhole’s conservation is absolutely needed. Gathering scientific data related to the dhole’s conservation by the multidisciplinary team will benefit to the management in resolving the problem arises during the management process. The availability of adequate ecological data will also be useful for increasing public understanding in the importance of dhole conservation and attracting public sympathy, thus they become more tolerant to the existence of this species.
Keywords: Cuon alpinus, Conservation, Ecology, Java, Legal status, Management.
... Prater (1965) stated that a number of female wild dogs (Cuon alpinus) may select a den site and form a breeding colony. A group of new young ones may have several different aged that create confusion to determine which one is more younger or offspring of which female wild dog (Cuon alpinus) then, Johnsingh (1982) clears that confusion and he stated that breeding is restricted to only a single female wild dog (Cuon alpinus) within a pack, as is also the case for the highly social, pack living African wild dogs (Lycaon pictus) and for wolves (Canis lupus). In a pack of Wild dogs (Cuon alpinus) every adult ones help to young ones to provide food at their den and protect all descendants from the predators like Tiger (Panthera tigris) and other big carnivores who can eat their descendants along with the lactating females. ...
... In a pack of Wild dogs (Cuon alpinus) every adult ones help to young ones to provide food at their den and protect all descendants from the predators like Tiger (Panthera tigris) and other big carnivores who can eat their descendants along with the lactating females. "Guards"-Wild dog (Cuon alpinus) that remain behind at the den site with the young ones while the rest are out for the hunting-are also fed (Johnsingh 1982;Fox 1984). Many times, in a group, members of a group of Wild dogs (Cuon alpinus) after hunting, it is not mandatory to all members of group to assemble again. ...
... Belligerent interactions between the group members will seen extremely rare and group member showing willingness to each other for the killing purpose and in a group generally competition for the hunting at very low level. The relating places where wild dogs (Cuon alpinus) can live and their range of that kind of places in India is generally in a group is around 35-40 km 2 , this home range is only when they are in no breeding season but when we talk about during their breeding season then their suitable habitat range will rapidly decrease from 35-40 km 2 to 10-12 km 2 (Johnsingh 1982). ...
Habitat suitability is an illustrious mission to conserve the ecosystems and its components. Human wildlife conflicts have increased tremendously due to habitat fragmentation; reason being human encroachment for their own use or for commercial purpose like forest products and unsustainable development programs leading to major decline in Biodiversity. Pench Tiger Reserve (PTR), Madhya Pradesh has great capacity to nurture young Wild dogs (Cuon alpinus) because it supports lots of biodiversity especially herbivores which will be great availability of food and have great tree cover on the landscape. The motive of this study is to find out the landscape which will suitable for Wild dog (Cuon alpinus) at PTR, MP using presence SDM (Species Distribution Model) at 900 m grain Size using Maxent, Ecological Niche Factor analysis and Bioclim and compare with AHP (Analytical hierarchical process). Maxent gives contributions of every factor in percentage also gives responses curve of every factor separately and also give permutation importance of every factor. Results show that the model with accuracy more than 0.90 AUC value. Bioclimatic variable 17 and LULC contributes the moderately high in the model. For AHP only need experts, who fill the Saaty table on the basis of environmental variables importance to wild dog (Cuon alpinus). Calculate or predict the habitat suitability of wild dog (Cuon alpinus) by experts rating to every environmental variable (comparison of every variables with respect to each other) from 1 to 9 or inverse like 1/9. Maxent shows that out of the about 750 km2 areas, more than 150 km2 found to be highly suitable at 900 m grain size SDM. This shows PTR, MP has the great potential to nurture or sustain Wild dog (Cuon alpinus) because wild dog (Cuon alpinus) need larger area to walk, they walk daily around 20–25 km to find their food and PTR, MP is immensely rich in large size herbivore animals like Chital (Axis axis). Strict implementation of laws is required this can be achieved by government agencies like forest department and others along with that awareness can be spread among the local residents of the PTR, MP so the wildlife and their habitat can be conserve by the conservationists and wildlife experts.
... They are one of the few carnivorous mammals in the canid group that hunt as a group (Marshall-Pescini and Kaminski 2014). Normally, there are 5-12 members in a pack (Johnsingh 1982;Durbin et al. 2004) to increase the efficiency of hunting large prey such as banteng, gaur, and sambar. The number of pups per litter is from 2 to 8 (Cohen et al. 1978). ...
... The total distance of the walking paths is about 100 km From February 2017 until September 2018, when scat is found, it is classified according to characteristics, and the size and footprints are measured in that area (Cohen et al. 1978;Johnsingh 1992;Karanth and Sunquist 1995;Selvan et al. 2013). Normally dhole packs hunt, feed, and often defecate together (Johnsingh 1982;Karanth and Sunquist 1995;Durbin et al. 2004;Thinley et al. 2011). Therefore, we sampled only 1 scat from each dropping to help ensure that the scats were from independent feeding events. ...
... That confirms the activity period of dholes in Thailand mostly occur during day time. This is consistent with Johnsingh (1982); Karanth and Sunquist (2000); Acharya (2007); Kawanishi and Sunquist (2008) and Majumder (2011) who found that dholes mostly hunt prey during the day time especially in the morning and afternoon, which is different from tigers and leopards that hunt during the night time until the morning. However, Ghaskadbi et al. (2016) reported that dholes at the Tadoba-Andhari Tiger Reserve, India, have the activity pattern as a crepuscular activity pattern as in the report by Nurvianto et al. (2016) which studied dholes on Java Island, Indonesia, which shows that dholes have the ability to adapt their hunting times. ...
Sukmasuang R, Charaspet K, Reontik J, Pla-ard M. 2020. Temporal overlap of carnivorous mammal community and their prey in Khao Ang Rue Nai Wildlife sanctuary, Chachoengsao Province, Thailand. Biodiversitas 21: 922-932. This study on the temporal overlap of the carnivorous community and their prey in Khao Ang Rue Nai Wildlife Sanctuary was conducted from March 2017 to February 2018. Camera traps were deployed systematically with a total of 4,463 trap nights. Fourteen carnivorous mammals were recorded, which were mainly present at night, with the exception of the dhole, small Indian mongoose, crab-eating mongoose, and yellow-throated marten. The clouded leopard's presence overlapped between day and night. Using the average coefficient overlap(Δ) between a carnivore, and the other carnivorous species, the leopard cat was found to have the highest Δ value, followed by the hog badger, Asiatic jackal, small Indian civet, Asian palm civet, large Indian civet, large spotted civet, Asiatic black bear, dhole, Malayan sun bear, yellow-throated marten, small Indian mongoose, crab-eating mongoose, and clouded leopard. The potential prey species that had the highest Δwith the carnivorous species, was the Siamese hare. This study shows the importance of preserving the carnivorous community within the area. An important threat is a likelihood that carnivorous species in the area may be exposed to external diseases from infected domestic animals when coming out to hunt in the communities surrounding the protected area.
... Зграю формують або члени однієї сім'ї, або декількох споріднених родин. Щенята покидають лігво у віці від 70 до 80 днів, а у семимісячному віці активно беруть участь у полюванні (Johnsingh 1982; Venkataraman et al. 1995). ...
... Саме це призвело до негативного ставлення місцевого населення до питання збереження дхолів (Arya et al. 2015). Ареал полювання зграї під час денного періоду становить близько 11 км 2 , а в інший час -понад 15 км 2 (Johnsingh 1982). ...
У навчальному посібнику детально розглянуто представників чоти-рьох родин хижаків, зокрема ведмедевих, псових (вовчих), котових (котя-чих) і мустелових (куницевих). Подано загальні відомості про тварин, ареа-ли їх поширення, спосіб життя, розмноження, харчування. Висвітлено про-блеми збереження і приналежність до видових категорій згідно з Червоним списком Міжнародного союзу охорони природи.
Для студентів і викладачів географічних факультетів вищих закладів освіти, майбутніх фахівців у галузі географії і природничих наук, усіх, хто цікавиться життям тварин.
... Despite the knowledge of these ongoing situations of dhole biology and conservation, appropriate studies to verify genetic diversity and differentiation patterns are limited 14 . To date, most of the studies have focused on their behaviour [20][21][22] , occupancy 23 , population pattern 24,25 , genetics [26][27][28] at local/regional scales, still an in-depth understanding of population/demographic patterns are lacking. ...
... Generating detailed information on dhole population parameters at the landscape scale is highly challenging due to their obligate forest dependency, elusive nature and sensitivity towards anthropogenic activities. Therefore, dhole research has generally been focused on local habitat scales (for example, behavioural studies by 20,22,32 , population patterns by 28 ) or at their distribution and occupancy using standard approaches 33 . The present study is perhaps the most exhaustive primary information on dhole genetics and population patterns across the species distribution to the best of our knowledge. ...
Deforestation and agricultural intensification have resulted in an alarming change in the global land cover over the past 300 years, posing a threat to species conservation. Dhole is a monophyletic, social canid and, being an endangered and highly forest-dependent species, is more prone to the loss of favorable habitat in the Anthropocene. We determined the genetic differentiation and demographic history of dhole across the tiger reserves of Maharashtra using the microsatellite data of 305 individuals. Simulation-based analyses revealed a 77–85% decline in the major dhole sub-populations. Protected areas have provided refuge to the historically declining dhole population resulting in clustering with strong genetic structure in the remnant dhole population. The historical population decline coincides with the extreme events in the landscape over the past 300 years. The study highlights the pattern of genetic differentiation and diversity of a highly forest-dependent species which can be associated with the loss of forest cover outside tiger reserves. It also warrants attention to develop conservation plans for the remnant surviving population of dholes in India.
... Additionally, dholes are only 1 of 3 canid species with specialized morphological and behavioral adaptations for an exclusively carnivorous diet (i.e., hypercarnivory; Van Valkenburgh 1991). Behavioral traits associated with hypercarnivory include forming exceptionally large packs (often >10 members) to more efficiently hunt and consume large numbers of prey (Johnsingh 1982). Therefore, maintaining relatively high densities of preferred ungulate prey species might be especially important for conserving a hypercarnivorous canid species. ...
... For each scat, we recorded the diameter, date, and global positioning system (GPS) location. Packs of dholes hunt, feed, and defecate together (Johnsingh 1982, Karanth and Sunquist 1995, Durbin et al. 2004, Thinley et al. 2011. Specifically, while traveling through their territory, the alpha pair often defecate together along their travel path, then beta pack members will defecate in the immediate vicinity (A. ...
Endangered dholes ( Cuon alpinus ) are restricted to small and declining populations in Southeast Asia, and little is known about how their ecology differs within the region. We used DNA‐confirmed scats and prey surveys to determine the seasonal diet and prey selection of dholes in 2 different landscapes that dominate Southeast Asia: closed evergreen forests in hilly terrain in northern Laos, and open deciduous forests in relatively flat terrain in eastern Cambodia. On both sites, muntjac ( Muntiacus spp.; 20–28 kg) was the dominant prey item and was selectively consumed over other ungulates in all seasons. Our findings differ from previous conclusions, based largely on studies from India, that the preferred prey weight range of dholes was either 40–60 kg or 130–190 kg. Other important prey were sambar ( Rusa unicolor ) in Laos, and wild pig ( Sus scrofa ) and banteng ( Bos javanicus ) in Cambodia. Seasonal differences in overall diet occurred in Laos, but not Cambodia, primarily because of an increase in livestock consumption. The mean number of dhole scats in group defecation sites was higher in Cambodia (5.9 ± 0.5 [SE]) than Laos (2.4 ± 0.2), suggesting pack sizes were larger in Cambodia. Our results suggest that regardless of land cover type, prey diversity, or pack size, the management of muntjac will be important for conserving dhole populations in Southeast Asia. In Laos, we recommend that local villagers remove livestock from the protected area during the hot‐dry season to reduce livestock predation by dholes. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.
... Provisioning both the pups and lactating mother is widely documented paternal care amongst canids (Asa and Valdespino, 1998). As Macdonald (1992) noted, regurgitation of partially digested food is widely described in the lupine canid lineage (present in all species in the genera Canis, Lycaon, Cuon, Chrysocyon and Speothos; Biben, 1982;Johnsingh, 1982;Rasmussen and Tilson, 1984;Asa and Valdespino, 1998;Lord et al., 2013) but absent in the vulpine lineage (though see Poessel and Gese, 2013). In these more carnivorous canids, this economical means of transporting prey to the den without the risk of kleptoparasitism is clearly advantageous (e.g., van Lawick-Goodall and Lawick-Goodall, 1970). ...
... An innovation in canids is the ability to directly feed both pups and mother by regurgitation of partially digested food. This ability is found in all wolf-like canids (Canis, Cuon, and Lycaon genera, Johnsingh, 1982;Lord et al., 2013) as well as maned wolves (Rasmussen and Tilson, 1984) and bush dogs (Biben, 1982). Regurgitation is generally absent from vulpine canids, although it was recently reported in the swift fox (Poessel and Gese, 2013). ...
The Canidae are successful, being a widespread, abundant, speciose, and adaptable family. Several canids in particular have recently experienced rapid expansions in range and abundance, with similar situations mirrored on several continents by different species. Despite extreme behavioral diversity between and within species, monogamy is a common denominator in canid societies. In this review, we ask why canids are monogamous and how monogamy is related to their success. We begin with an overview of canid social monogamy, describing the pair bonding, paternal care, and often alloparental care that is characteristic of the family, and discuss theories on the evolution of mammalian social monogamy. We discuss why and how monogamy is maintained in canids, either voluntarily or enforced, and how ecological conditions influence either the functional advantages of monogamy or ability for enforcement and thus whether social monogamy is maintained. Social monogamy does not necessitate exclusive mating and many canids exhibit extra-pair paternity. We consider the costs and benefits of extra-pair mating for male and female canids and how ecological conditions can shift this cost/benefit balance and thus affect its prevalence. Monogamy may be responsible for many of the unusual canid reproductive characteristics through facilitating alloparental care and monogamy enforcement, and the domestic dogs' departure from monogamy supports our interpretation that it is an adaptation to resource availability. In asking whether monogamy is responsible, at least in part, for their success, we propose the monogamy as pro-cooperative hypothesis, suggesting four characteristics have contributed to canid success: (1) ecological flexibility, (2) high mobility, (3) high reproductive rates, and (4) sociality/cooperation, with the latter two being consequences of monogamy. These four interconnected traits enhance one another and it is their combination, with monogamy at its foundation enabling cooperative sociality and thereby enhanced reproduction and survival, that together comprise the formula of canid success.
... The Asiatic wild dog or dhole (Cuon alpinus) is a highly elusive, endangered, social canid distributed in south and southeast Asia (Johnsingh, 1982;Durbin et al., 2004) occupying a range of habitat types including alpine, temperate, sub-tropical and tropical forests (Durbin et al., 2004). Driven by habitat loss, prey depletion, disease transmission from domestic dog, human persecution and interspecific competition (Hayward, Lyngdoh & Habib, 2014;Kamler et al., 2015), dholes are currently found in about 75% of their historical global range (Durbin et al., 2004;Kamler et al., 2015). ...
... Dholes prefer dense forested habitats (Johnsingh, 1985) where the social groups defaecate in communal latrine sites mostly found on the junctions of roads/trails (Johnsingh, 1982). Their elusive nature and highly social behaviour present unique challenges in scat sampling for individual identification. ...
Background:
The Asiatic wild dog or dhole (Cuon alpinus) is a highly elusive, monophyletic, forest dwelling, social canid distributed across south and Southeast Asia. Severe pressures from habitat loss, prey depletion, disease, human persecution and interspecific competition resulted in global population decline in dholes. Despite a declining population trend, detailed information on population size, ecology, demography and genetics is lacking. Generating reliable information at landscape level for dholes is challenging due to their secretive behaviour and monomorphic physical features. Recent advances in non-invasive DNA-based tools can be used to monitor populations and individuals across large landscapes. In this paper, we describe standardization and validation of faecal DNA-based methods for individual identification of dholes. We tested this method on 249 field-collected dhole faeces from five protected areas of the central Indian landscape in the state of Maharashtra, India.
Results:
We tested a total of 18 cross-species markers and developed a panel of 12 markers for unambiguous individual identification of dholes. This marker panel identified 101 unique individuals from faecal samples collected across our pilot field study area. These loci showed varied level of amplification success (57-88%), polymorphism (3-9 alleles), heterozygosity (0.23-0.63) and produced a cumulative misidentification rate or PID(unbiased) and PID(sibs) value of 4.7 × 10-10 and 1.5 × 10-4, respectively, indicating a high statistical power in individual discrimination from poor quality samples.
Conclusion:
Our results demonstrated that the selected panel of 12 microsatellite loci can conclusively identify dholes from poor quality, non-invasive biological samples and help in exploring various population parameters. This genetic approach would be useful in dhole population estimation across its range and will help in assessing population trends and other genetic parameters for this elusive, social carnivore.
... For animals, the shelter is an indispensable resource for their survival and propagation (McComb and Noble 1981;Ruggiero et al. 1998). Animals use shelters predominantly as sleeping or resting sites (Miles et al. 1981;Carter and Encarnação 1983;Beck-King et al. 1999;Pardini and Trajano 1999), temporary resting sites (Prestrud 1992;Larivière and Messier 1998), food caches (Post et al. 1993), and natal burrows (Johnsingh 1982). A suitable natal burrow site is a key requirement for many small to medium mammals which provides a safe dwelling for reproduction and hence these structures become significant in population growth function (Kinlaw 1999), by supporting the breeding success of many burrow-dependent species (Alt and Gruttadauria 1984;Ruggiero et al. 1998). ...
... Most of the canids are seasonal breeders (Holt et al. 2014), albeit canids such as bush dog exhibiting no seasonality. The breeding timings of different (Rodden et al. 1996(Rodden et al. , 2004, the Indian fox (Acharjyo and Misra 1976;Johnsingh 1978;Johnsingh and Jhala 2004), and the dholes (Cuon alpinus) (Davidar 1973;Johnsingh 1982;Venkataraman 1998) in India are all breeding during the dry spell. In India, the dry spell is followed by the monsoon and several of these abiotic factors affect the breeding onset and subsequent birth of young ones (Lehman et al. 1997;Prendergast 2005;Holt et al. 2014), thus optimizing (Moehlman 1979(Moehlman , 1983(Moehlman , 1987. ...
Availability and use of a natal burrow site is a prerequisite for survival, propagation, and breeding success for many burrow-dependent species. Among mammals, canids typically use existing burrows of other animals during the breeding season. The study describes the factors influencing the golden jackal (Canis aureus) in selecting appropriate burrows in Keoladeo National Park (KNP), India, including a description of site-specific habitat characteristics, microhabitat conditions, and the burrow-specific activities. Jackals had occupied 11 of the 47 recorded burrows and had chosen the area with more wood cover. The occupied burrows were found active throughout the day. The mean detections per day were significantly highest in the month of May for “rearing pups.” The nocturnal activity was associated with the lunar phase, with a significant peak between the “third quarter” to “new moon” phase. The intensity of the activity of adults and pups were analyzed across 6 weeks (April–May), wherein feeding, grooming, playing, and guarding significantly occurred between fourth and sixth weeks. The pups suckled every 8 h, and at about 3 weeks old, regurgitated food also became part of their diet. Jackal pups and adults spent an average of 103.60 and 36.13 min per day playing and guarding the burrows, respectively. Increased temperature with the day’s progression significantly reduced jackal’s activities and they retreated around 1200 h. Guarding of the burrows by adults significantly increased during night hours (0030–0500 h), when the pups are more susceptible to predation. Both the sexes participated in parental care activities, taking turns to guard the burrow, grooming, and playing with the pups. The study thus provides insights on the golden jackal’s natal site selection, observances, and use of sub-surface earthen refuges in semi-arid condition.
... entrance, complex den with several entrances, simple den in a cave, under a rock or between two rocks, and finally a complex den in a cave with several connected entrances. Inside these dens faeces, hair and remains of regurgitated food from adults to feed the offspring can be found (Johnsingh, 1982). Some authors have claimed that bones found in dens were due to the presence of hyenas or porcupines (Johnsingh, 1982), but others, Fox among them, have pointed out that the bones could be there because of the offspring. ...
... Inside these dens faeces, hair and remains of regurgitated food from adults to feed the offspring can be found (Johnsingh, 1982). Some authors have claimed that bones found in dens were due to the presence of hyenas or porcupines (Johnsingh, 1982), but others, Fox among them, have pointed out that the bones could be there because of the offspring. Dholes, according to the characteristics of the pack, can occasionally transport part of the prey to their den. ...
... The fieldwork was conducted by two teams, each consisting of three surveyors experienced in recognizing dhole scats and mammal signs in general. Dhole scats were easily identified because these animals tend to defecate together in latrines (Johnsingh, 1982;Karanth & Sunquist, 1995;Durbin et al., 2004;Thinley et al., 2011;Kamler et al., 2012), a phenomenon not reported for sympatric domestic dogs or Asiatic jackals (Canis aureus) (Cohen et al., 1978), and the scat identification was confirmed by their distinctive odor, appearance and presence of dhole tracks near the collection site (Cohen et al., 1978;Johnsingh, 1992;Karanth & Sunquist, 1995;Selvan et al., 2013b). Once scats were found, they were collected and stored in an airtight plastic bag (Selvan et al., 2013a) and the locations of the scats were recorded using GPS Garmin 76 CSX. ...
... Only scats which were clearly identifiable were collected. As stated above, dholes hunt, feed, and defecate together in latrines (Johnsingh, 1982;Karanth & Sunquist, 1995;Durbin et al., 2004;Thinley et al., 2011;Kamler et al., 2012), so that individual latrines contained scats that nearly always have the same content, suggesting they came from the same feeding event (Thinley et al., 2011). Hence, as a general rule, only one scat from each latrine was collected to help ensure that scats were from independent feeding events (Kamler et al., 2012). ...
We conducted a study on the feeding habits of dholes in the Baluran National Park, Indonesia. Scat analysis was employed to identify the prey consumed. In total, 54 scats were collected across the park during the dry season of 2013 and analyzed to identify the prey of this species; at least 20 prey species were identified. Ungulates are the most important prey, estimated to contribute more than 95 % of the biomass consumed by dholes in BNP. Efforts to ensure availability of ungulates and to secure habitat will be the key to the dhole's conservation in Java.
... To optimise capture probability; we maintained a mean distance of 672 m (SE ± 15.73) between neighbouring camera traps. Previous studies indicated home ranges of 23-199 km 2 for dholes (Johnsingh 1982;Venkataraman et al. 1995;Karanth and Sunquist 2000;Acharya 2007); hence, we believe this inter-trap distance was suitable for our target species. The average temperature and humidity during the entire sampling period were 24 °C (± 0.29 SE) and 79.4% (± 0.54 SE), respectively (Time and Date 2021). ...
The dhole or Asiatic wild dog Cuon alpinus is the only endangered social canid in the tropical Indian forests and is considered at high risk of extinction in the wild. Despite this, knowledge about its ecology is still scarce. We investigated relative abundance, habitat utilisation and spatial and temporal ecology of dholes in Manas National Park, North-East India. Camera traps (n = 473) were deployed across forested habitats of Manas for 11,388 trap nights. Independent records and relative abundance index were 163 and 1.431 ± 0.387 SE per 100 trap nights, respectively. Dhole habitat use exhibited a non-significant relationship with habitat and anthropological variables. The availability of small prey, i.e., rhesus macaque, showed a tendency of a positive association with dhole habitat use and, therefore, intrinsically highlights the potential conservation importance of small prey species. Low temporal overlaps (< 0.50) were observed between dhole and other sympatric carnivores. Dholes showed a positive co-occurrence pattern with tigers and leopards. To coexist, dholes segregate themselves from other sympatric carnivores, primarily by utilising different temporal niches and adjusting their activity peaks when encountering them. This spatio-temporal segregation suggests a partial avoidance of dholes from dominant predators that may decrease competition and the risk of intraguild predation.
... In a typical defecation posture, hoary foxes lower the hindquarters and bend their tails, moving their tails up and down when defecation is complete. Unlike truly social canids (Johnsingh 1982), hoary foxes defecated a regular amount each time in a certain place, instead of small portions in several places. Individual or communal latrines were not observed, although the finding of a greater number of scats in certain sites of the reproductive areas could suggest this possibility. ...
The hoary fox, Lycalopex vetulus, is a small insectivorous canid endemic to grasslands of Central Brazil. Herein we characterize hoary fox social behavior, based on qualitative and quantitative differences in intraspecific communication among males, females, and subadults. A family group, two reproductive pairs and three other adult individuals of unidentified social status were directly observed from 1995 to 2001, for the continuous recording of all occurrences of behavioral events. Hoary fox exhibited behaviors related to communication and food consumption. A total of 4,240 behavioral events were recorded, most in the categories ‘Consumption’ (58%) and ‘Olfactory’ (23%). The frequencies of signaling types varied among individuals according to sex, age, and socioecological phase. Olfactory signalings were more frequent in adult males, and during the Mating and Dispersal phases. Acoustic signalings were more frequent in adult females and individuals rearing the young. ‘Consumption’ and ‘Tactile’ behaviors were more frequent in subadults. The hoary fox intraspecific communication is characteristic of solitary canids, but during mating season and rearing period, individuals become more ‘socialized.’ Data on courtship and mating, partnership dissolution phase, and the breakup of the family group until young dispersal, are necessary for a better understanding of intraspecific communication of the hoary fox.
... Among communal breeders, such as the African lion Panthera leo, the spotted and the brown hyaenas, and the banded mongoose, most females breed during each reproductive cycle and participate in some degree of alloparental care although temporarily nonbreeding females and males may also contribute to the care of young born in the group (Mills 1990;Lewis and Pusey 1997). In facultative cooperative breeders such as the black-backed jackal Canis mesomelas and Arctic fox, the parents and nonbreeding helpers alike care for the young (Johnsingh 1982) although the number of helpers is small, and parents can successfully raise their young without helper assistance (Clutton-Brock 2006). Obligate cooperative breeders, such as the African wild dog and the meerkat Suricata suricatta, require assistance from nonbreeding helpers to successfully raise their offspring. ...
The mammalian order Carnivora is generally defined as species that feed exclusively or to some degree by eating other animals. The Carnivora comprise around 280 species, divided into 16 families, 13 of which are terrestrial and 3 aquatic. Carnivores are spread across the entire planet, including the two polar regions and on land and sea. Consistent with such diverse ecologies, there is no typical pattern of parental care distinguishing carnivores from other mammals. Using examples from different taxonomic families, our aim is to illustrate the diversity of parental care in Carnivora. Major topics include parental care before and after birth of the young, paternal, and alloparental care and the process of weaning. Given the position of many carnivores at the apex of food chains, a greater understanding of their patterns of parental care as a vital part of reproductive biology is essential to conservation programs.
... Among communal breeders, such as the African lion Panthera leo, the spotted and the brown hyenas, and the banded mongoose, most females breed during each reproductive cycle and participate in some degree of alloparental care although temporarily nonbreeding females and males may also contribute to the care of young born in the group (Mills 1990;Lewis and Pusey 1997). In facultative cooperative breeders such as the black-backed jackal Canis mesomelas and Arctic fox, the parents and nonbreeding helpers alike care for the young (Johnsingh 1982) although the number of helpers is small, and parents can successfully raise their young without helper assistance (Clutton-Brock 2006). Obligate cooperative breeders, such as the African wild dog and the meerkat Suricata suricatta, require assistance from nonbreeding helpers to successfully raise their offspring. ...
Direct care of offspring by the father (sire) is relatively rare in primates. Besides humans, there are a number of species where the male is essential for the survival of offspring: marmosets, tamarins, titis and owl monkeys, some lemurs, and siamangs. All these species show reduced sexual dimorphism, territoriality, and biparental care. However, timing and levels of direct care may vary among these species. Here, relying on both lab and field data, we address the variability found in father's involvement with his infants, the behavioral, neuroendocrine and sensory systems that are a cause and consequence of paternal care, and social bonds between the breeding pair. We integrate studies of laboratory animals (where detailed observations and experimentation are possible) with field studies (which illuminate the ecological and evolutionary functions of paternal care) and discuss the future directions for examining the proximate and ultimate mechanisms of paternal care in nonhuman primates.
... Scent-marking in latrines is widespread among the Canidae. Kit foxes (Vulpes macrotis) (Ralls and Smith 2004), coyotes (Canis latrans) (Bowen and McTaggart-Cowan 1980;Ralls and Smith 2004), Ethiopian wolves (Canis simensis) (Sillero-Zubiri and Macdonald 1998), wolves (Canis lupus) (Barja et al. 2004), golden jackals (Canis aureus) (Macdonald 1979), racoon dogs (Nyctereutes procyonoides) (Ikeda 1984), dholes (Cuon alpinus) (Johnsingh 1982;Thinley et al. 2011), maned wolves (Chrysocyon brachyurus) (Dietz 1984) and swift foxes (Vulpes velox) (Darden et al. 2008) all defecate in latrines, and on this basis, African wild dogs can be expected to use latrines for scent communication (expectation 1). ...
Understanding the use of space by endangered African wild dogs (Lycaon pictus) can contribute to their conservation because persecution by livestock farmers is a major cause of mortality when wild dogs range outside protected wildlife areas. Scent-marking is likely to be a key mechanism by which African wild dog packs organise their mutual use of space, and here, we report the discovery that African wild dog packs repeatedly shared a scent-marking latrine. We present the first systematic records of scent communication between packs of African wild dogs. Camera trapping for 13 months recorded four wild dog packs in northern Botswana repeatedly scent-marking with urine and faeces at a latrine where two dirt roads crossed a large game trail, in the overlap zone of two of the packs’ home ranges. This site was visited by wild dogs at a mean interval of 12.2 days, with a mean of 10.3 days between consecutive visits by different packs, and a mean of 28.7 days between consecutive visits by the same pack. Dominant dogs scent-marked every time they visited this site for a minute or longer. Sniffing and countermarking of scent-marks from previous visits showed that this shared marking site acted as a bulletin board, with 12 exchanges of information between packs over a period of 13 months.
Significance statement
African wild dogs are endangered large carnivores, whose populations suffer continual attrition from lethal conflict with livestock owners. Deterring wild dogs from leaving protected wildlife areas will reduce this conflict-related mortality. All terrestrial mammals mark their home ranges with scent, and using artificial scent-marks to simulate African wild dog home range boundaries along protected area borders has the potential to reduce conflict killings by deterring the dogs from leaving the protected areas. Our discovery of multiple African wild dog packs scent-marking at a persistent, shared latrine, where scent-marking depended on sex and social status, is a breakthrough in our understanding of inter-pack communication in this endangered species and a key step towards developing artificial home range boundary markers that will deter them from leaving protected wildlife areas.
... Species specific studies: Unlike most carnivores, the wild dogs have a unique social structure. They are cooperative breeders (Johnsingh, 1982;Venkataraman, 1998) and packs can sometimes be of up to 25 adults (Kamler et al., 2015). This in turn can determine the prey species, home range as well as heterogeneity of the population, and it is thus important to focus on species specific studies in order to understand their current population, ecology as well as natural history within the reserve. ...
A short-term field study on the status of Endangered and Threatened species of Mammals (Medium and large sized) and Herpetofauna in the Sahyadri Tiger Reserve, located in the northern Western Ghats, in Maharashtra, was conducted between November 2016 and June 2017. This intensive field study was aimed in providing robust scientific information for the Sahyadri Tiger Reserve, on the mammals and herpetofauna found in this landscape.
... The alpha pair are the only breeding pair in the pack while the other members support the pair in raising and protecting the pups. Females give birth to up to 7 pups per litter after a gestation period o f 60-62 days (Johnsingh, 1982;Paulraj et al., 1992). In order to feed such large litters, the females have eight pairs of mammae. ...
The book "Wild Mammals of the Shuklaphanta National Park" describes a total of 56 mammalian species. We have described 56 mammals in the book; however additional species "Himalayan Goral" was spotted in the park when this book was in the press. The new update for the park is 57 mammalian species. The documentation of 56 species were based on camera-trap photographs (37), other photographic evidences (4) published and unpublished records with locality details (5) and published reports without locality details (10). Out of 56 mammal species, 17 are globally threatened, 3 are near threatened in IUCN Red List and 34 species are listed in CITES Appendices (10 in Appendix I; 8 in Appendix II; and 16 in Appendix III) In National Red List of Nepal, 23 species are nationally threatened, 1 near threatened and 5 are data deficient. Ten species are protected by the National Parks and Wildlife Conservation Act 1973 as Schedule I species.
... Dholes are also estimated to require five times more land area than large-bodied carnivores such as tigers (Kamler et al., 2012), mainly because of the social structure of populations living in exclusive territories (Johnsingh, 1982), unlike solitary tigers with overlapping territories (Carter et al., 2015). Large space requirements in conjunction with a hypercarnivorous diet (Van Valkenburgh, 1991) make dholes more vulnerable to extirpation, as evidenced by their disappearance from more reserves than tigers (Woodroffe and Ginsberg, 1998). ...
Most canids face population declines and range contractions worldwide. Although the dhole (Cuon alpinus) is widely distributed across 10 countries in South and Southeast Asia, limited studies exist on this species. Despite its globally “Endangered” status and ecological role as an apex predator, assessments on its distribution are limited to a few landscapes and countries. This explains the lack of a dhole-specific species conservation plan in most range countries, including Bhutan where no current population estimate exists. The species has also recovered from a country-wide poisoning campaign in the 1970s and 80s. In this study, we determine the dhole's distribution pattern and assess the protection and connectivity of dhole habitat in Bhutan. We anticipated dholes to be extant within their habitat well-represented in protected areas (PAs) and biological corridors (BCs). We used 721 georeferenced dhole occurrence records and eight environmental variables in MaxEnt software to model potential dhole distribution and habitat suitability. The model output was overlaid on the spatial layers of PAs and BCs to assess habitat protection and connectivity. As anticipated, we found the dhole widely distributed in all districts, PAs, and BCs in Bhutan. Dholes were recorded at the highest elevation range limit of 4,980 m above sea level, which overlapped with the “Vulnerable” snow leopard (Panthera uncia). Our model identified 72% (27,634 km2) of the country as suitable areas for dholes, of which, 31% (11,899 km2) was highly suitable and 41% (15,735 km2) was moderately suitable. Contrary to our expectation, PAs and BCs encompassed only 29% (8,046 km2) and 12% (3,185 km2) of suitable areas for dholes, respectively. A vast majority of the areas we deemed suitable for dholes currently remain unprotected, thus making dholes more vulnerable to human persecution and local extermination. We recommend adjusting PA boundaries to fully encompass suitable dhole habitat, and also advocate improved livestock husbandry to reduce dhole related livestock predation and minimize conflict, thereby ensuring its long-term survival in Bhutan.
... Recent studies of the mitochondrial DNA of the African wild dog showed a close relationship with canines (Canidae) of the Canis, Cuon, Cerdocyon, Atelocynus, Atelocynus, Dusicyon and Lycalopex genus [37]. According to Johnsing [38] and Venkataraman [39], the African wild dogs and dholes (Cuon alpinus) have a similar morphology, behavior and ecology. Radinsky [40] writes in his study that the South African Painted Dogs has the same number of chromosomes and similar neuroanatomy as the domestic dog (Canis lupus familiaris). ...
In this study, we present the first data concerning the anatomical, morphometrical, histological and histochemical study of the orbit, eye tunics, eyelids and orbital glands in South African Painted Dogs ( Lycaon pictus pictus ). The study was performed using eyeball morphometry, analysis of the bony orbit including its morphometry, macroscopic study, morphometry, histological examination of the eye tunics and chosen accessory organs of the eye and histochemical analysis. The orbit was funnel shaped and was open-type. There was a single ethmoid opening for the ethmoid nerve on the orbital lamina. The pupil was round, while the ciliary body occupied a relatively wide zone. The iris was brown and retina had a pigmented area. The cellular tapetum lucidum was semi-circular and milky and was composed of 14–17 layers of tapetal cells arranged in a bricklike structure. In the lower eyelid, there was a single conjunctival lymph nodule aggregate. One or two additional large conjunctval folds were observed within the posterior surface of the upper eyelids. The superficial gland of the third eyelid had a serous nature. The third eyelid was T-shaped and was composed of hyaline tissue. Two to three conjunctival lymph nodul aggregates were present within the bulbar conjunctiva of the third eyelid. The lacrimal gland produced a sero-mucous secretion. A detailed anatomic analysis of the eye area in the captive South African Painted Dogs females showed the similarities (especially in the histological examination of the eyetunics and orbital glands) as well as the differences between the Painted dog and the other representatives of Canidae . The differences included the shape and size od the orbita with comparison to the domestic dog. Such differences in the orbit measurements are most likely associated with the skull type, which are defined in relation to domestic dogs. The presented results significantly expand the existing knowledge on comparative anatomy in the orbit, eye and chosen accessory organs in wild Canidae .
... Only a few wild dog species in this world hunt in packs (Marshall-Pescini and Kaminski 2014). Normally, there are 5-12 members in a Dhole pack (Johnsingh 1982;Durbin et al. 2004) to increase the efficiency in hunting large prey such as Banteng (Bos javanicus d'Alton, 1823), Gaur (Bos gaurus Smith, 1827), Sambar deer (Rusa unicolor Kerr, 1792), etc. These pack hunters have a great number of members and will need spacious areas for storing the prey, which is mainly ungulates, to appropriately maintain the Dhole populations in the ecosystem. ...
Charaspet K, Sukmasuang R, Khoewsree N, Pla-ard M, Chanachai Y. 2020. Prey species and prey selection of dholes at three different sites in Thailand. Biodiversitas 21: 5248-5262. The study of prey species and prey selection of Dholes at 3 different sites was conducted at Khao Yai National Park, Salak Pra, and Huai Kha Khaeng Wildlife Sanctuaries from 2013 to 2020. Information on Dhole prey at the sites was collected from the residues of dhole scats, from which the selection index, the relative biomass of the prey, and the relative amounts of the consumed prey were calculated. The data were collected simultaneously with the use of camera traps at each site. The study revealed that there were 13 species of Dhole prey with body weight over 5 kg. The result indicated that there were 7 species of even-toed ungulates. The relative biomass of even-toed ungulates ranged between 76.78 - 90.50% of the total biomass of all the Dholes’ consumed prey for all study sites. The dietary diversity index unveiled a similar index in all areas, which proved the adequacy of the analyzed scats. However, the Niche breadth index, which indicates the relevance of prey selection and prey species to the appearances of the prey at each site, was found to be high at Huai Kha Khaeng Wildlife Sanctuary, Khao Yai National Park, while the index was found to be low at Salak Pra Wildlife Sanctuary. The results revealed that Dholes consumed viverrid species and Malayan porcupine more often at the site where there were large carnivores. The recommendation from this study was the conservation and restoration of the ungulate populations, the main prey, as it greatly affects the conservation of the Dhole populations in Thailand. Grassland and salt lick sites, water sources improvements are also important to promote prey population. The conservation of wildlife prey by releasing them to nature, as currently conducted, has an effect on the increase of Dholes’ prey species.
... Among the sub-populations of Dhole in southern Asia, Johnsingh (1985) reported that it is frequently seen in many of the protected areas south of the Ganga River, with the central Indian highland forests having the largest population of Dhole, followed by the Western Ghats of southern India. In Western Ghats, Bandipur National Park was presumed to have had the largest subpopulations of Dhole four decades back, a total population of 207-304 individuals and estimated 44-64 mature individuals (20-29 alpha males and 20-29 alpha females with 4-6 sub-dominant breeders (Johnsingh 1982). Dhole density in southern India over the last four decades vary between 14-100 /km 2 ; e.g., 31/100 km 2 (Venkataraman et. ...
The Asiatic Wild Dog has a wide global range covering Central, South and South-East Asia. It has been recorded over most of the Indian subcontinent except for the deserts of western India and Eastern Ghats of Tamil Nadu. We hereby provide photographic evidence of the Asiatic Wild Dog Cuon alpinus from Khandige Estate in Sirumalai.
... The Red List assessment by the IUCN suggests that India harbors the largest population of dholes; Thailand and Myanmar support medium populations, whereas Bhutan, Cambodia, China, Indonesia, Laos, Malaysia and Nepal support small populations (Kamler et al., 2015). Over the past five decades, most ecological studies of dhole have dealt with diet analyses and behavioral observations (Johnsingh, 1982;Venkataraman et al., 1995;Karanth and Sunquist, 2000;Kumara et al., 2004;Ghaskadbi et al., 2016; see Supplementary information 1 for a full list of diet studies). More recently, select studies have examined distribution and habitat associations at various spatial scales (Karanth et al., 2009;Jenks, 2012;Srivathsa et al., 2014;Punjabi et al., 2017). ...
Livestock depredation is the most ubiquitous type of negative interaction between humans and carnivores. We conducted a range-wide assessment linking diet patterns of the endangered dhole Cuon alpinus, with livestock consumption and human-dhole interactions. We first performed a reanalysis of dhole diet data from all published studies (1973-2013) incorporating a recently-developed non-linear correction factor for quantifying prey biomass consumed. We then determined the relative livestock numbers consumed by dholes over time across its range, compared these with earlier estimates, and investigated the relative importance of wild vs. non-wild prey in dhole diet. Using information from >70 studies, we explored links between livestock consumption by dholes, availability of wild versus non-wild prey, sympatric depredation-prone carnivores, and people's perception of dholes as livestock predators. We found that (a) dhole diet profiles varied regionally, (b) dholes consumed fewer livestock compared to estimates generated using other, widely used methods, (c) livestock consumption by dholes was associated with wild and non-wild prey densities, and number of co-predator species, and (d) people's negative perception of dholes was associated with pack sizes, levels of livestock depredation and number of sympatric carnivore species. Global efforts for dhole conservation should involve different strategies based on region-specific realities that account for ecological context as well as human perceptions, which would require well-designed studies of dhole social and population dynamics, and human-dhole interactions. We also call for more such range-wide assessments of livestock depredation by wild canids, complemented with direct investigations of human-canid interactions.
... For faecal sampling, the entire study area was intensively surveyed using a vehicle as well as on foot to look for dhole latrine sites. Dholes have a communal latrine system where they defecate in groups generally at the junction of roads (Johnsingh 1982). All samples were collected fresh from latrine sites during our surveys. ...
Asiatic wild dog (Cuon alpinus) or dhole is an endangered canid with fragmented distribution in South, East and Southeast Asia. The remaining populations of this species face severe conservation challenges from anthropogenic interventions, but only limited information is available at population and demography levels. Here, we describe the novel molecular approaches for unambiguous species and sex identification from noninvasively collected dhole samples. We successfully tested these assays on 130 field-collected dhole faecal samples from the Vidarbha part of central Indian tiger landscape that resulted in 97 and 77% success rates in species and sex identification, respectively. These accurate, fast and cheap molecular approaches prove the efficacy of such methods in gathering ecological data from this elusive, endangered canid and show their application in generating population level information from noninvasive samples.
... The presented data on cues to orientation in the Dhole calls highlight the probable role of contact biphonic calls in communication of this species. The Dhole is pack-living canid, communally hunting on large prey and inhabiting areas with complex relief in mountains and in locations with dense vegetation; with primary breeding by a dominant pair and with other group members functioning as helpers, and with very low intrapack aggression (COHEN 1977, JOHNSINGH 1982, KARANTH & SUNQUIST 1995, VENKATARAMAN et al. 1995, VENKATARAMAN 1998, LUDWIG & LUDWIG 2000. In the previous study, we showed that the biphonic yap-squeaks provide significantly better individual discrimination in comparison with non-biphonic yaps and squeaks occurring as separate vocalisations. ...
Cues to orientation of a caller to a listener in biphonic and non-biphonic close range contact calls in the Dhole (Cuon alpinus) Dholes produce two call types with wide frequency ranges, potentially providing cues to orientation of a caller to a listener because of different distribution for the higher and lower frequencies. One call type is biphonic (yap-squeak), the second one (yap) includes only one fundamental with its harmonics. Here we compare the abilities the biphonic and non-biphonic, but rich in harmonics call types to encode orientation of a caller to a listener. We recorded calls and movements from three male Dholes, running singly forward and back in their identical enclosures and subdivided the recorded calls into two groups: produced toward a microphone in sector ±45°, and produced away from a microphone in sector 135-225°. For each call, within 20.3 ms time segment taken in a call centre, we calculated the amplitude ratio of sum of amplitudes higher 5 kHz to sum of amplitudes lower 5 kHz. For the pooled sample of yaps and yap-squeaks, the amplitude ratio was significantly higher for "toward" than for "from" call group. For yaps and yap-squeaks separately the results were similar. Also, Dhole showed interindividual differences both in preference of a particular call type and in reliability of cues to orientation. Overall, both biphonic yap-squeaks and non-biphonic yaps bored cues to orientation. We discuss, that the increased occurrence of biphonic yap-squeaks in the Dhole may be conditioned by their additional function as individual markers, lacking in non-biphonic yaps.
... comm.) due to: 1) their requirement for larger group sizes to kill larger prey compared to jackals which hunt smaller prey (Johnsingh, 1982;Moehlman, 1983;Mukherjee et al., 2004;Jaeger et al., 2007); 2) amicable behaviour between within-group individuals (Fox, 1984). Further, when local people enter Chhep Wildlife Sanctuary they often bring dogs and dholes are often killed in snares in Cambodia (J. ...
... Studies from southeast Asia suggest that smaller packs in tropical forests of Thailand have home-range sizes between 50 and 100km 2 (Grassman et al. 2005, Jenks et al. 2012). In the central part of Western Ghats (our study area), dhole packs appear to have much smaller home-range sizes (seasonally 20-60km 2 ; Johnsingh 1982, Venkataraman et al. 1995, Karanth and Sunquist 2000, a likely consequence of high densities of ungulate prey in these well-protected deciduous forest reserves (Karanth et al. 2004). Our estimate presented here is much larger compared to that reported by Karanth and Sunquist (2000), who estimated home-range size of ~27km 2 for a pack of dholes from an adjacent area in Nagarahole between 1988-1991. ...
The dhole Cuon alpinus is one of the least studied large carnivores in the world. Unlike many other social canids, dholes occur at low densities in tropical forests. Furthermore, they are wary, and difficult to capture and radio-tag, thereby posing challenges in the field for tracking their movements or making behavioural observations. We conducted intensive camera-trap surveys in Nagarahole and Wayanad wildlife reserves in the Western Ghats of southern India, as part of a long-term study of tiger population dynamics. The survey duration was kept short, from 28 November 2014 to 12 January 2015 (45 days), to ensure demographic closure. Besides generating data on tigers, the surveys also yielded incidental photographic captures of dholes. In general, individual dholes cannot be uniquely identified from camera-trap photographs. But we were able to identify two individuals in a pack based on distinct markings on their pelage, enabling us to map locations of the pack during the survey period. We present here, an estimate of home-range size (~85km 2) for this dhole pack from non-invasive camera-trap surveys.
... Dholes hunt in a group and prefer medium-sized ungulates, although they also prey on smaller sized animals and scavenge (Fox, 1984). Ethological studies show that dholes consume their prey very quickly at the place of the kill (Domínguez Rodrigo, 1993), although they may occasionally transport part of the carcasses to their dens, where they may also defecate (Johnsingh, 1982;Fox, 1984). In the Iberian Peninsula the dhole has been found in several Middle Pleistocene sites, although they were more highly distributed during the Late Pleistocene, even conserving some populations into the early Holocene (Pérez Ripoll et al., 2010). ...
Bone accumulations created by carnivores during the Pleistocene have been largely associated with the action of cave hyenas (Crocuta spelaea), with breeding dens identified all over Europe. Yet in recent years it has been shown that leopards (Panthera pardus) played a role in the creation of sites in the Iberian Peninsula. In this study we present the taphonomic study of the bone accumulation of the Pre-Solutrean level at the Racó del Duc Cave (south of the province of Valencia), where the dominant species (Capra pyrenaica) has been accumulated by leopards. Together with new data and other references taken from the bibliography, we have summarised the main taphonomic characteristics of the sites accumulated by leopards. The predator-prey relationship has also been analysed and it has shown that during the Pleistocene, leopards in the Iberian Peninsula were specialised in catching goats, showing similar behaviour to that of snow leopards. Said specialisation may have been due to the occupation of caves in steep areas where rupicolous fauna predominated.
... Dholes are considered to be primarily diurnal in habit (Johnsingh 1980;Acharya 2007;Kawanishi and Sunquist 2008;Majumder 2011); however, we observed a crepuscular activity pattern similar to that reported by Nurvianto et al. (2015) in Java, Indonesia. The dhole's vocal repertoire was similar to that reported by Johnsingh (1982), with the exception of "Chuckle" (Table 3). Additional research focused on vocalizations will be needed to compile a comprehensive vocal repertoire for the dhole. ...
The dhole (Cuon alpinus) is an endangered social canid that inhabits the forests of southern and southeastern Asia. A scarcity of field studies and inconsistent findings have led to a poor understanding of their ecology and conservation status. We compiled an ethogram of dhole behavior based on analysis of 395.35min of video recordings. We recorded 3,394 behavioral events in 1,654 video clips lasting 10s each. We classified behavioral events into 6 categories: Locomotion, Resting, Social Behavior, Feeding, Scent Marking, and Miscellaneous. Behavioral events associated with Locomotion were most frequent (40.95%), as was the proportion of time spent on such behaviors (41.89%). Dholes spent the least time exhibiting Miscellaneous and Scent Marking behaviors (1.45% and 2.64%, respectively), as well as the lowest frequency (0.74% and 4.01%, respectively). Although scent marking was relatively rare, we observed unique scent-marking behaviors in this study, including “hind bounce” and “hind scrub.” The time spent on different categories of behaviors differed significantly among males, females, and subadults. We also used camera traps and opportunistic observations to investigate the activity patterns of dholes in dry deciduous forest of Tadoba-Andhari Tiger Reserve in central India, where they are sympatric with other large predators including tigers (Panthera tigris) and leopards (P. pardus). Our findings suggested that dholes were primarily crepuscular. Fundamental knowledge about behavioral ecology is crucial for the conservation of any species, and our findings provide a new foundation for future behavioral research on this endangered social canid.
... This method has been criticized however, because a home range can include large areas of land which are never visited, and in some cased even impossible to access because of their geographical constraints [26]. However, because previous studies have used this method to estimate the home range of dholes [20,[29][30][31], we used the 95% MCP method in order to be able to compare our results to previous research findings. The KUD method was used to identify areas of high use, and is considered to be a method which provides a very good means of highlighting areas of concentrated activity [26]. ...
We conducted the first research on the activity patterns and movement of denning dhole in the Baluran National Park, Indonesia. Camera traps and radio telemetry surveys were employed to observe dhole activity patterns and movement over the denning period. The dholes showed crepuscular and diurnal activity patterns with most activities intensifying at dawn and dusk, and becoming less intense in the middle of the day. The dhole’s home range comprised of hunting grounds, water resources, and a den in the centre. The home range size was estimated at 744.86 ha (using the 95% Minimum Convex Polygon/MCP method), at 1418.28 ha (using the 80% Kernel Utilization Distribution/KUD analysis), and at 479.59 ha (using 90% Local Convex Hull/LoCoH), whereas the core area was estimated to be 636.36 ha (50% KUD) and 67.37 ha (50% LoCoH), and the size of the most greatly used area at 231.57 ha (25% KUD) comprising of the den sites and the hunting grounds. The dhole’s den ecology strategy was to use other animal’s burrows on steep slopes with dense vegetation cover and located on the opposite side of the hill to where human activity occurred. Den switching occurred every 2 weeks. These results indicate that dholes selected a den site that fulfilled their needs for food, water, cover, and predator evasion.
Keywords: Cuon alpinus, den ecology, radio telemetry, home range, predator.
Cooperative breeding (i.e. when alloparents care for the offspring of other group members) has been studied for nearly a century. Yet, inconsistent definitions of this breeding system still hamper comparative research. Here, we identify two major inconsistencies, discuss their consequences and propose a way forward. First, some researchers restrict the term 'cooperative breeding' to species with non-breeding alloparents. We show that such restrictive definitions lack distinct quantitative criteria to define non-breeding alloparents. This ambiguity, we argue, reflects the reproductive-sharing continuum among cooperatively breeding species. We therefore suggest that cooperative breeding should not be restricted to the few species with extreme reproductive skew and should be defined independent of the reproductive status of alloparents. Second, definitions rarely specify the type, extent and prevalence of alloparental care required to classify species as cooperative breeders. We thus analysed published data to propose qualitative and quantitative criteria for alloparental care. We conclude by proposing the following operational definition: cooperative breeding is a reproductive system where >5% of broods/litters in at least one population receive species-typical parental care and conspecifics provide proactive alloparental care that fulfils >5% of at least one type of the offspring's needs. This operational definition is designed to increase comparability across species and disciplines while allowing to study the intriguing phenomenon of cooperative breeding as a behaviour with multiple dimensions.
Das Bedürfnis nach sozialer Anerkennung ist in unserem Alltag allgegenwärtig. Wir machen Überstunden, um Lob vom Chef zu bekommen, strengen uns beim Sport an, um Pokale zu gewinnen, und kaufen Statussymbole, um von anderen bewundert zu werden. Warum wir diese und viele weitere Bräuche durchführen, welche Rituale es früher gab und welche psychologischen Mechanismen hinter der Suche nach Anerkennung stecken, erfahren Sie in diesem Kapitel.
Scent marks deposited as semiochemical signals are a primary mode of communication for a broad range of mammal species. Such scent signals are often deposited at specific, frequently visited marking sites called latrines. Despite descriptions of widespread latrine use by numerous mammal species, detailed understanding of site visit rates and latrine function is lacking. Here we report for the first time a quantitative assessment of scent-marking behaviours that represent interpack olfactory communication by African wild dogs, Lycaon pictus, at latrines visited by multiple resident neighbouring packs, hereafter called a ‘shared marking site’ (SMS). We show that multiple packs visited specific SMSs frequently and regularly throughout the year, with a notable decrease in visits during the 3-month denning season coinciding with a contraction in range size. In addition to resident neighbouring packs, dispersing individuals visited and scent-marked at SMSs, suggesting that latrines function at least in part as sites communicating information about residence and possibly reproductive status. Further detailed investigation of the relevance of latrine use to territorial behaviour, ranging, habitat use and dispersal in this species is required, particularly as it may have direct applied conservation implications for this wide-ranging but territorial endangered species.
Latrines are accumulations of two to several hundred faeces resulting from the repeated use of the same defecation sites by the same or several individuals. Many carnivores deposit their faeces in such dedicated latrine sites, which are often shared by several animals either from the same social group or from neighbouring territories. Although latrines are assumed to play an important role in olfactory communication, detailed knowledge of specific information exchange is still lacking. Four different categories of data are important in trying to understand the function of latrines in animal societies: (i) spatial distribution patterns; (ii) temporal usage patterns; (iii) individual visit and contribution patterns; and (iv) information content of the signal. While the spatial distribution of latrines in relation to territory boundaries, landmarks and resources has been studied in a variety of species, only a few studies concentrated on temporal variation in latrine usage. Even fewer studies provide insights into inter‐individual differences in visit and contribution patterns or the olfactory information content of latrines. In this review, we outline potential functional hypotheses for latrine use and develop a research framework for the study of latrine function. We then present three model species – European badger, Meles meles , meerkat, Suricata suricatta , and banded mongoose, Mungos mungo – for which we have detailed data for at least three of the four above‐mentioned categories, which we will use to test these hypotheses. Throughout the chapter, we review the different techniques used to collect these data in different species, discuss the limitations of using spatial data alone to test functional hypotheses, and highlight the value of a combined approach.
The function of holding territories is primarily to have access to resources like food and mates. However, it is costly in terms of energy and time investment. Solitary-living, territorial species are known to reduce these costs by being more aggressive towards unfamiliar strangers and less aggressive towards neighbors. However, in social, territorial species, neighbors can impose a greater threat than strangers. We tested whether the highly social Asiatic wild dogs/dholes (Cuon alpinus) exhibit the “nasty neighbor” or the “dear enemy” phenomena in Tadoba Andhari Tiger Reserve (TATR), Maharashtra, India. We conducted scat translocation experiments where we presented fresh scats collected from unique donor groups to a resident dhole group and tested the type and the intensity of behavioral response (duration) to the stimulus. Dholes responded differentially to the two treatments suggesting they exhibit neighbor-stranger discrimination. Overall, strangers elicited a stronger response with longer duration and larger packs were less likely to respond to the stimulus than smaller packs. Differences found between categories of dhole scent marks establish the importance of olfactory communication, especially “counter-marking” in the species. Within recipient packs, individual status affected the response to trials wherein the alpha pair reacted more intensively to strangers than others. Our study provides experimental evidence to demonstrate that dholes exhibit the “dear enemy” phenomenon.
Significance statement
Animals defend territories from other members of their own species, but intrusions are commonplace in the wild. Different intruders may pose different levels of threats, and hence, intruders are treated differentially to minimize the energetic costs of territorial defense. In some animals, neighbors with well-established territories may become less aggressive towards each other. This is known as the dear enemy effect. By contrast, at times neighbors may represent a greater threat than strangers which is known as the “nasty neighbor” effect. We experimentally show that dholes exhibit the dear enemy phenomenon by responding more intensively to strangers than familiar neighbors. We show how response varied based on hierarchy in a pack as well as the pack sizes. Furthermore, we found that, both in core as well as buffer areas of their own territory, this relationship was consistent.
Robust population estimation is indispensable for well-informed management and conservation of species. Capture-recapture models provide a robust framework for density estimation of animals with unique identifying features. The challenge is amplified for group-living species lacking identifiable marks where both the number of groups and number of individuals needs to be estimated simultaneously. We developed a model to estimate the population of group-living animals using a hierarchical clustering approach. We used weighted geographical distance and probability of missing an individual from photo-capture sequence to estimate number of groups and individuals in a group. We simulated scenarios with a varying number of capture sequence to test the effect of number of sequences on a detection probability and group-size. We applied the proposed method on a group living, unidentifiable canid, dhole (Cuon alpinus) following a multidisciplinary approach. We examined the average radius estimated from clustering method against home-range radius obtained from GPS-collared individuals. Additionally, we compared population estimates with individuals identified through molecular tools. Simulations showed there was no substantial change in the number of individuals estimated with the number of photo-capture sequences. The efficacy of our model was more than 70% until the ratio of the radii of the pack operating area and pack range centres were less than 0.8. The spatial extent and population estimate derived from clustering model was comparable with GPS telemetry data and individuals identified using molecular methods. Our model would broaden the scope of the applicability of camera trap studies for social species with uniquely unidentifiable individuals. With assumptions like territoriality, independent captures and cohesion of the study species, this model can aid in the monitoring and conservation of rare, endangered species like dholes, which otherwise pose a challenge to managers and conservation practitioners.
The Dhole Cuon alpinus used to be the meso carnivore of the forests throughout Indian subcontinent; however, habitat loss, low prey biomass, and human disturbance exterminated the species from India’s 60% historic range, and the numbers are less than 1,500 individuals in wild. Following the same shrinking trend, Dholes were extirpated from Gujarat. A few doubtful sightings and inevident reportings generated ambiguity of Dhole presence in Gujarat. We conducted a study in Vansda National Park with 15,660 trap nights at 30 trap locations, and have confirmed the rediscovery of Dholes in Gujarat after 70 years. We estimated the Dhole’s minimum home range as 13.7km2 and also analyzed relative abundance index of other mammals. The future retention of Dholes requires detailed range, diet, and adaption studies along with conservational efforts to reduce re-extinction probabilities.
Synopsis
Many large predators are also facultative scavengers that may compete with and depredate other species at carcasses. Yet, the ecological impacts of facultative scavenging by large predators, or their “scavenging effects,” still receive relatively little attention in comparison to their predation effects. To address this knowledge gap, we comprehensively examine the roles played by, and impacts of, facultative scavengers, with a focus on large canids: the African wild dog (Lycaon pictus), dhole (Cuon alpinus), dingo (Canis dingo), Ethiopian wolf (Canis simensis), gray wolf (Canis lupus), maned wolf (Chrysocyon brachyurus), and red wolf (Canis rufus). Specifically, after defining facultative scavenging as use or usurpation of a carcass that a consumer has not killed, we (1) provide a conceptual overview of the community interactions around carcasses that can be initiated by facultative scavengers, (2) review the extent of scavenging by and the evidence for scavenging effects of large canids, (3) discuss external factors that may diminish or enhance the effects of large canids as scavengers, and (4) identify aspects of this phenomenon that require additional research attention as a guide for future work.
Resource depletion exerts opposing pressures on co‐occurring consumers to expand diets while limiting overlap with competitors. Using foraging theory as a framework, we tested the effects of prey availability on diet specialization and overlap among competing Asian predators: dhole, leopard, tiger. We used scat analysis from a prey‐poor site, combined with a quantitative synthesis of 40 other diet studies, to determine biomass of different prey types consumed by each predator. We then assessed diet composition in relation to prey density, and compared diet breadth and overlap between prey‐poor and prey‐rich sites. In prey rich areas, all three predators specialized on energetically profitable medium and large ungulates (> 30 kg), resulting in narrow, overlapping niches. Each predator shifted towards less profitable small‐bodied prey (≤ 30 kg) as preferred ungulates declined, whereas consumption of preferred ungulates was unrelated to small prey abundance, as predicted by foraging theory. Diet breadths doubled under prey depletion (except leopard), but overlap declined as diets diverged via species‐specific traits that facilitated capture of different types of alternative prey. Asia's apex predators adapt similarly to depletion of mutually preferred ungulates by switching to more numerous but less profitable small prey. Yet they can also partition a depleted prey base through intrinsic niche differences, thereby avoiding competitive exclusion. Our findings illuminate the stabilizing properties of adaptive foraging and niche differences in ecological communities, and provide insights into the behavior and resilience of Asia's endangered apex predators in response to prey depletion in the heavily‐poached forests of this region.
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In animal societies, individuals can cooperate in a variety of tasks, including rearing young. Such cooperation is observed in complex social systems, including communal and cooperative breeding. In mammals, both these social systems are characterized by delayed dispersal and alloparenting, whereas only cooperative breeding involves reproductive suppression. While the evolution of communal breeding has been linked to direct fitness benefits of alloparenting, the direct fitness cost of reproductive suppression has led to the hypothesis that the evolution of cooperative breeding is driven by indirect fitness benefits accrued through raising the offspring of related individuals. To decipher between the evolutionary scenarios leading to communal and cooperative breeding in carnivores, we investigated the coevolution among delayed dispersal, reproductive suppression, and alloparenting. We reconstructed ancestral states and transition rates between these traits. We found that cooperative breeding and communal breeding evolved along separate pathways, with delayed dispersal as the first step for both. The three traits coevolved, enhancing and stabilizing one another, which resulted in cooperative social systems as opposed to intermediate configurations being stable. These findings promote the key role of coevolution among traits to stabilize cooperative social systems and highlight the specificities of evolutionary patterns of sociality in carnivores.
In most mammalian species, females regularly interact with kin, which is expected to reduce aggressive competitive behaviour among females. It may thus be difficult to understand why infanticide by females has been reported in numerous species and is sometimes perpetrated by groupmates. Here, we investigate the evolutionary determinants of infanticide by females by combining a quantitative analysis of the taxonomic distribution of infanticide with a qualitative synthesis of the circumstances of infanticidal attacks in published reports. Our results show that female infanticide is widespread across mammals and varies in relation to social organization and life history, being more frequent where females breed in groups and have intense bouts of high reproductive output. Specifically, female infanticide occurs where the proximity of conspecific offspring directly threatens the killer's reproductive success by limiting access to critical resources for her dependent progeny, including food, shelters, care or a social position. By contrast, infanticide is not immediately modulated by the degree of kinship among females, and females occasionally sacrifice closely related infants. Our findings suggest that the potential direct fitness rewards of gaining access to reproductive resources have a stronger influence on the expression of female aggression than the indirect fitness costs of competing against kin.
This article is part of the theme issue ‘The evolution of female-biased kinship in humans and other mammals’.
The food habits of dhole were evaluated in the tropical forests of Silent Valley National Park (SVNP) from December 2011 to May 2012 by analysing their scats. Eleven prey species were identified. Sambar was found to be the principal prey species for dhole as inferred from the relative biomass consumption of prey remains in dhole scats. Regarding prey biomass contribution, sambar was highest (66.74%) while grey jungle fowl was the lowest (0.32%). The aim of this study was to assess the food habits of dhole co-existing with large predators, tiger and leopard in the tropical forests of SVNP.
The impact of environment, interspecific competition, and, to a lesser extent history, on the structure of the guild of large predatory mammals is explored in one fossil and four Recent communities. Two aspects are emphasized: (1) the number of species within each guild and (2) the extent of locomotor convergence as inferred from morphology among the constituent species. Locomotor behavior reflects habitat choice, hunting mode, and escape strategy, all of which appear to be important avenues of adaptive divergence among coexisting predators.
Locomotor behavior in extinct and extant predators is determined from body weight and five measured characteristics of the postcranial skeleton, including ungual shape, elbow shape, and limb proportions. Results indicate that levels of morphologic and inferred ecologic similarity between large predators are higher in the tropical grassland guild of East Africa than in the equivalent guilds of either tropical or temperate forest. This may be due to the great density and diversity of terrestrial herbivores in the productive grasslands. The fossil guild, from the Late Chadron–Orellan (Oligocene) of North America, appears most similar to the tropical forest guilds, but the predators seem to have been slower and more robust than their modern counterparts. Since the Orellan represents an early stage in the evolution of large, fissiped carnivores, both ancestry and time could have influenced Orellan guild structure.
The African wild dog Lycaon pictus was originally described by C.J. Temminck in 1820 from a specimen collected on the Mozambique coast. He called it a form of hyaena Hyaena picta, but it was soon recognised as one of the canids, and was first renamed Lycaon tricolor by Brookes in 1827. The name Lycaon is derived from the Greek word lykaios which means wolfish. It was then given the specific epithet pictus,which is derived from the original picta and which is the Latin word for spotted, as directed by the International Rules on Taxonomic Nomenclature. The African wild dog is today also sometimes called the Cape hunting dog, or the painted dog. The latter name refers to the wild dog’s varied coat colour with its blotched patches of white, black and yellow, which led to the use of the name tricolor by Brookes. Although all wild dogs have this patchy coat colour, those of the southern savannas have a bit more white in their coat than their northern counterparts.
The domestic dog (Canis lupus f. familiaris) and cat (Felis silvestris f. catus), which are quite vocal by mammalian standards, are not good representatives of the acoustic activities of fissiped carnivores. Fissipeds are generally thought of as mammals that communicate with smell rather than with vocalizations (Gorman and Trowbridge, this volume). Nevertheless, several carnivore acoustic signals like the howling of gray wolves (Canis lupus), the whooping of spotted hyenas (Crocuta crocuta), and the roaring of African lions (Panthera leo) capture the human imagination as few other animal sounds do. It is probably no accident that wolf howling—unlike other acoustic signals of carnivores—is one of the best-studied mammalian vocalizations (Theberge and Falls 1967; Cohen and Fox 1976; Tembrock 1976a, 1976b; Fox and Cohen 1977; Shalter et al. 1977; Field 1978, 1979; Fox 1978; Harrington and Mech 1978a, 1978b, 1979, 1982, 1983; Schassburger 1978; Klinghammer and Laidlaw 1979; Filibeck et al. 1982; Harrington 1986, 1987; Nikolskii and Frommolt 1986).
This paper presents an overview of large carnivore research and a perspective on the deficiencies in our knowledge about this group. Since the landmark research done in the 1960’s, our knowledge of carnivore ecology and the amount of research has been unbalanced. In some areas of carnivore ecology, our knowledge has remained practically unchanged since the early 1970’s. Notable exceptions have been work on the Hyaenidae and Felidae in open habitats of Africa, work with the felids on the Indian subcontinent, and work on the Ursidae and Canidae in North America. Over this same time period (and starting much earlier), the knowledge base for game species around the world has increased several-fold. We discuss the special research needs of carnivore ecology and conservation, especially in the areas of human/carnivore conflicts, population viability, and experimental approaches. We stress the urgent need for work on large carnivores in developing countries, particularly in those regions of the world where little or no modern ecological work has been performed.
Literature regarding the biology of the dhole (Cuon alpinus) is reviewed, summarized, and re-interpreted, providing a framework for further research on this endangered species. The basis for the taxonomic classification of the dhole (subfamily Simocyoninae) is reviewed and questioned. Distribution, habitat, and physical characteristics are noted and compared between populations in India and the U.S.S.R. Dholes are typically seen in packs of 5-12 individuals and more than one female in the pack may breed simultaneously. Dens may be earthen or cavernous and may connect to others. Pups often engage in agonistic encounters until 7-8 months of age, when the dominance hierarchy may be established. Co-operative hunting strategies may operate to kill large ungulates, but lagomorphs and rodents may be taken by individuals or pairs. Domestic livestock species are rarely taken. Several myths concerning hunting behavior are discussed and (hopefully) laid to rest. The known communicative signals are reviewed and suggestions made to orient future research. Relations with man reveal a history of unmerited persecution. Indian dhole populations are considered to have declined due to indiscriminate slaughter, habitat destruction, and reduced prey aváilability.
African wild dog (Lycaon pictus) predation was observed in Ngorongoro Crater, Tanzania, between September, 1964, and July, 1965, when packs were in residence. The original pack of 21 dogs remained only 4 months, but 7 and then 6 members of the group reappeared in the Crater at irregular intervals. The ratio of males:females was disproportionately high, and the single bitch in the small pack had a litter of 9 in which there was only one female. The pack functions primarily as a hunting unit, cooperating closely in killing and mutual defense, subordinating individual to group activity, with strong discipline during the chase and unusually amicable relations between members. A regular leader selected and ran down the prey, but there was no other sign of a rank hierarchy. Fights are very rare. A Greeting ceremony based on infantile begging functions to promote pack harmony, and appeasement behavior substitutes for aggression when dogs are competing over meat. Wild dogs hunt primarily by sight and by daylight. The pack often approaches herds of prey within several hundred yards, but the particular quarry is selected only after the chase begins. They do not run in relays as commonly supposed. The leader can overtake the fleetest game usually within 2 miles. While the others lag behind, one or two dogs maintain intervals of 100 yards or more behind the leader, in positions to intercept the quarry if it circles or begins to dodge. As soon as small prey is caught, the pack pulls it apart; large game is worried from the rear until it falls from exhaustion and shock. Of 50 kills observed, Thomson's gazelles (Gazella thomsonii) made up 54 percent, newborn and juvenile wildebeest (Connochaetes taurinus) 36 percent, Grant's gazelles (Gazella granti) 8 percent, and kongoni (Alcelaphus buselaphus cokei) 2 percent. The dogs hunted regularly in early morning and late afternoon, with a success rate per chase of over 85 percent and a mean time of only 25 minutes between starting an activity cycle to capturing prey. Both large and small packs generally killed in each hunting cycle, so large packs make more efficient use of their prey resource. Reactions of prey species depend on the behavior of the wild dogs, and disturbance to game was far less than has been represented. Adult wildebeest and zebra (Equus burchelli) showed little fear of the dogs. Territorial male Thomson's gazelles, which made up 67 percent of the kills of this species, and females with concealed fawns, were most vulnerable. The spotted hyena (Crocuta crocuta) is a serious competitor capable of driving small packs from their kills. A minimum of 4-6 dogs is needed to function effectively as a pack. It is concluded that the wild dog is not the most wantonly destructive and disruptive African predator, that it is an interesting, valuable species now possibly endangered, and should be strictly protected, particularly where the small and medium-sized antelopes have increased at an alarming rate.
The eleven different functions for which mammals use urine marking are reviewed in this paper, and the urine marking behavior of the red fox (Vulpes vulpes) is described in detail. A new hypothesis is advanced that urine marking may serve as a "book keeping system" in the red fox's scavenging behavior. Foxes consistently investigate and urine mark inedible food remnants (e.g., bones, bird wings, and dried out pieces of hide). When a fox re-investigates a marked remnant, the urine mark signals "no food present," and the fox investigates this object for only a brief period of time. This use of urine marking may increase the efficiency of its scavenging behavior, i.e. more food-items found per hour of scavenging. This efficiency may be particularly important during periods of food shortage. The hypothesis is tested in three different experiments, using free-ranging red foxes as subjects. Experiment I establishes that fox do urine mark food remnants. Experiment II shows that foxes investigate for a significantly shorter period of time (P<0.001) food remnants exhibiting both the odor of food and the odor of urine as compared to remnants exhibiting just the odor of food. Experiment III suggests that there a hierarchy of stimuli which determines different responses in the fox's scavenging behavior. The experiments also suggest that there is a degree of social behavior in the scavenging activities of red foxes. Foxes appear to use each other's urine marks to increase the efficiency of their scavenging behavior. Thus this study definitely support LEYHAUSEN'S (1965) statement that the social life of solitary animals is frequently more complex than we realize. Solitary species probably show many ingeniously adapted mechanisms for occupying niches where highly social species could not be maintained. The social evolution and ecological advantages of solitary species deserve to be the focus of future research.
Observations of wolves on Isle Royale are reported for 1961–66, with interpretations including the earlier 3-year period described by Mech (1966). On this 210-square-mile island the fully protected wolf population varied from approximately 22 to 28 in midwinter. The major and minor foods were moose and beaver, respectively.
The main pack varied in number between 11 and 22 with about three breeding pairs believed present. The population remained relatively stable; mating occurred every winter; and adult mortality appeared to be low. High mortality among pups seemed to be the point of population control. Socio-economic factors may have controlled the size of the large pack. Availability of food during the period of parturition and rearing probably was critical to survival of young.
Recruitment of young appeared to take place in years of high production of moose calves. Numbers in the large pack probably were curtailed through the progressive exclusion of aged and socially subordinate individuals. Under harassment these animals separated and became pack-following scavengers, then probably true loners ranging outside the area used by the pack. Smaller aggregations of two or three non-breeders were seen each winter, as were the loners, some of which appeared thin and weak.
The only known breeding outside the big pack was in a group of five present in the winter of 1965. This group was probably a family unit which separated from the main pack. A year later the male had disappeared, and remains of a pup, probably theirs, were found. In the winter of 1966 the alpha male of the large pack became lame and apparently was killed. This background appears favorable for further changes in social organization.
Information on Alaskan wolf populations was obtained from examination of bounty records, 4,150 wolf radii and ulnae, 1,262 wolf carcasses, and from observations of wolves inhabiting an area of 20,000 square miles where wolves were protected.
Pregnant adult female wolves averaged 6.5 fetuses; two-year-old females averaged 5.3 fetuses;female pups were not sexually mature.
In Alaska, wolves conceive from late February through early April but most females breed in March. Multiparous females breed earlier than first breeders. Multiparous females produce an average of 7.3 ova and 6.5 implanted fetuses. The loss of ova from ovulation to implantation is significant. Multiparous females produce more ova than first breeders; the difference is highly significant.
Mortality of pups rather than the lack of initial production of pups is believed to be the reason for the observed variations in the proportion of pups in wolf populations. Wolf packs include members in all categories of sex and age during the breeding season. Size of the pack is an indicator of abundance.
Wolves in an area where they were protected increased at an average rate of 20–30% per year during an 11-year period.
Miniature collar-type transmitters originally designed by W. W. Cochran, Illinois, were adapted for use on timber wolves (Canis lupus sp.) in east-central Ontario. Wild timber wolves were captured in steel traps, restrained with a forked stick, fitted with radio-collars and released at point of capture. Receivers were adapted for use in trucks, airplanes, and for walking in rough bush country. Maximum ranges were 3.2 km with ground and 9.6 km with aircraft receivers. A preadult female tagged in July, 1964, and a lactating adult female tagged in June, 1965, were tracked intermittently for 5.5 and 2.5 months, respectively. Tracking periods for six other animals of both sexes, ranging in age from pups to adults, varied from 2 days to 4 months. The lactating female and her associated pack regularly returned to three preferred “resting sites” for various periods during July and August. Preferred areas were well drained, semi-open, mixed conifer-hardwood stands in close proximity to swamps or beaver ponds. The preadult female ranged in an area frequented by a pack, but frequently remained independent of it. A preadult male, tagged in the same region, wandered over a slightly larger area than the female. Tagged animals were active throughout all periods of the day or night. Activity increased slightly during the early evening hours. There was a slight correlation between weather conditions and patterns of behavior and activity. Apparently, tagged individuals were quickly accepted by other members of the pack.
Studies of the interaction of wolves and their prey during the past twenty years have demonstrated that ungulates are the primary prey of wolves both in winter and summer and that predation in summer tends to be concentrated on the young of the year while in winter it is concentrated on animals in older-age classes. There appears to be intrinsic control of wolf numbers and there is evidence which suggests that a wolf per 10 square miles may approach the maximum density that is attainable in most ranges. The adaptations between ungulates and their predators may have evolved in relatively stable forest environments which could not support high-density prey populations. This could explain why wolves do not appear to be capable of controlling moose and deer populations in environments that have been drastically altered by man.
The population composition and density of wild and domestic ungulates in selected areas of Chitawan Valley, Nepal, are estimated and compared with other regions. Species considered include Axis axis, Axis porcinus, Cervus unicolor, Muntiacus muntjak, Sus scrofa, Rhinoceros unicornis, and domestic cattle, water buffalo, sheep, and goats. Rhinoceros constituted the bulk of the wild ungulate biomass in the riverine forest and grassland within the Royal Chitawan National Park. Species differed in their occurrence in various vegetation types and these differences afforded a degree of ecological separation. Counts indicated that juvenile mortality for most wild ungulates was high and seemed related to both nutritional and predatory factors. Suggestions for increasing the precision of counts of wild ungulates in monsoonal forest areas are included and conservation implications are discussed.
African wild dogs were studied from 1967 to 1978 on the Serengeti Plains in northern Tanzania. The main objectives were to determine the status of the sub-population and to elucidate the ecology and behavior of this social carnivore. This paper describes the decline of the sub-population, the dynamics of pack composition, and patterns of dispersal.
Wolves live in territorial packs of up to 20 animals. To gain insight into the regulation of wolf populations it is necessary to understand the regulation of pack size. Based on a long-time study of the social behaviour of a wolf pack in a large enclosure, a model of the regulation of pack size is constructed. Ecological factors such as prey size and prey biomass have an influence on wolf mortality, as well as on aggressive, sexual and spatial behaviour within the pack.
Wild dogs attacking a tiger
- Connell W.
Dhole or Indian wild dog (Cuon alpinus) mating
- Davidar E. R. C.
A wild boar (Sus scrofa) sharing wild dogs (Cuon alpinus) kill
- Johnsingh A. J. T.
An instance of wild dogs scavenging on a tiger's kill
- Johnsingh A. J. T.
Census of wild animals in Bandipur Tiger Reserve
- Wesley D. G.
Jungle memories: Part IV
- Adams E. G. P.
Observations at the dens of the Dhole or Indian wild dog (Cuon alpinus)
- Davidar E. R. C.
Panther and wild dogs
- Burton R. W.
Hunting and feeding in wild dog
- Fox M. W.
Einige Beobachtungen zum verhalten des Dekkan-Rothundes (Cuon alpinus dukhunensis Sykes) in Kanha-National Park
- Keller R.
Radio tracking wolves in Algonguin park
- Kolenosky