Dogs are one of the most common species to be found as pets and have been subjects of human curiosity, leading to extensive research on their socialization with humans. One of the dominant themes in dog cognition pertains to their capacity of understanding and responding to human referential gestures. The remarkable socio-cognitive skills of pet dogs, while interacting with humans, is quite well established. However, studies regarding the free-ranging subpopulations are greatly lacking. The interactions of these dogs with humans are quite complex and multidimensional. For the first time, we tested 160 adult free-ranging dogs to understand their ability to follow relatively complex human referential gestures using dynamic and momentary distal pointing cues. We found that these dogs are capable of following distal pointing cues from humans to locate hidden food rewards. However, approximately half of the population tested showed a lack of tendency to participate even after successful familiarization with the experimental set-up. A closer inspection revealed anxious behavioural states of the individuals were responsible for such an outcome. Finally, we compared the results using data from an earlier study with dynamic proximal cues. We found that free-ranging dogs follow distal cues more accurately compared to proximal cue. We assume that life experiences with humans probably shape personalities of free-ranging dogs, which in turn influence their responsiveness to human communicative gestures.

Cooperative breeding is an excellent example of altruistic cooperation in social groups. Domestic dogs have evolved from cooperatively hunting and breeding ancestors, but have adapted to a facultatively social scavenging lifestyle on streets, and solitary living in human homes. Pets typically breed and reproduce under human supervision, but free-ranging dogs can provide insights into the natural breeding ecology of dogs. We conducted a five year-long field based behavioural study on parental care of free-ranging dogs in India. 23 mother-litter units, belonging to 15 groups were observed, which revealed the presence of widespread allo-parenting by both adult males and females. While all the females were known to be related to the pups receiving care, the relatedness with the males could not be determined. Hence, we coined the term "putative father" for caregiving males. Allomothers provided significantly less care than the mothers, but the putative fathers showed comparable levels of care with the mothers. Mothers invested more effort in nursing and allogrooming, while the putative fathers played and protected more. Our observations provide support for both the "benefit-of-philopatry" and "assured fitness returns" hypotheses. Free-ranging dogs are not cooperative breeders like wolves but are rather communal breeders; their breeding biology bearing interesting similarities with the human joint family system. This breeding strategy is likely to have played an important role in increasing pup survival in a stochastic environment and helping to adapt to living among humans during the domestication of dogs.

Differences in pet dogs’ and captive wolves’ ability to follow human communicative intents have led to the proposition of several hypotheses regarding the possession and development of social cognitive skills in dogs. It is possible that the social cognitive abilities of pet dogs are induced by indirect conditioning through living with humans, and studying free-ranging dogs can provide deeper insights into differentiating between innate abilities and conditioning in dogs. Free-ranging dogs are mostly scavengers, indirectly depending on humans for their sustenance. Humans can act both as food providers and as threats to these dogs, and thus understanding human gestures can be a survival need for the free-ranging dogs. We tested the responsiveness of such dogs in urban areas toward simple human pointing cues using dynamic proximal points. Our experiment showed that pups readily follow proximal pointing and exhibit weaker avoidance to humans, but stop doing so at the later stages of development. While juveniles showed frequent and prolonged gaze alternations, only adults adjusted their behaviour based on the reliability of the human experimenter after being rewarded. Thus free-ranging dogs show a tendency to respond to human pointing gestures, with a certain level of behavioural plasticity that allows learning from ontogenic experience.

Parent-offspring conflict theory predicts the emergence of weaning conflict between a mother and her offspring arising from skewed relatedness benefits. Empirical observations of weaning conflict have not been carried out in canids. In a field-based study on free-ranging dogs we observed that nursing/suckling bout durations decrease, proportion of mother-initiated nursing bouts decrease and mother-initiated nursing/suckling terminations increase with pup age. We identified the 7th - 13th week period of pup age as the zone of conflict between the mother and her pups, beyond which suckling solicitations cease, and before which suckling refusals are few. We also report for the first time milk theft by pups who take advantage of the presence of multiple lactating females, due to the promiscuous mating system of the dogs. This behaviour, though apparently disadvantageous for the mothers, is perhaps adaptive for the dogs in the face of high mortality and competition for resources.

Dogs (Canis lupus familiaris) are the first species to have been domesticated, and unlike other domesticated species, they have developed a special bonding with their owners. The ability to respond to human gestures and language, and the hypersocial behaviours of dogs are considered key factors that have led them to become man's best friend. Free-ranging dogs provide an excellent model system for understanding the dog-human relationship in various social contexts. In India, free-ranging dogs occur in all possible human habitations. They scavenge among garbage, beg for food from humans, give birth in dens close to human habitations, and establish social bonds with people. However, there is ample dog-human conflict on the streets, leading to morbidity and mortality of dogs. Hence the ability to assess an unfamiliar human before establishing physical contact could be adaptive for dogs especially in the urban environment. We tested a total of 103 adult free-ranging dogs to investigate their response to immediate social and long-term food and social rewards. The dogs were provided a choice of obtaining a food reward either from the experimenter's hand or the ground. The dogs avoided making physical contact with the unfamiliar human. While immediate social reward was not effective in changing this response, the long-term test showed a strong effect of the social contact on the dogs' response. Our results revealed that these dogs tend to build trust based on affection, and not food rewards. The study provides significant insights into the dynamics of dog-human interactions on the streets and subsequent changes in behaviours of dogs through the process of learning.

Dens are crucial in the early development of many mammals, making den site selection an important component of parental care in such species. Resource availability and shelter from predators primarily govern den selection. Species inhabiting human-dominated landscapes typically den away from human disturbance, often shifting dens to avoid humans during the early life of their young. Domesticated dogs have evolved in human proximity over centuries, being bred and reared in human homes for generations. While pets rely on their owners for shelter and care, free-ranging dogs roam uncared, and typically whelp in dens. We conducted a study on 148 free-ranging dog dens in India to understand their denning habits. Distance from resources influenced den choice, but anthropogenic disturbance did not. Dens were found in areas of high human activity, and begging from humans was preferred over scavenging. A study on 15 pregnant females revealed that females actively searched for denning sites, rejecting several intermediate ones before selecting the final den. We propose that the obvious preference of dogs for denning close to humans is a behavioural adaptation that helps them to survive in the urban landscape, in spite of the high human induced mortality during the early life of pups.

Mammalian offspring require parental care, at least in the form of nursing during their early development. While mothers need to invest considerable time and energy in ensuring the survival of their current offspring, they also need to optimize their investment in one batch of offspring in order to ensure future reproduction and hence lifetime reproductive success. Free-ranging dogs live in small social groups, mate promiscuously and lack the cooperative breeding biology of other group-living canids. They face high early-life mortality, which in turn reduces fitness benefits of the mother from a batch of pups. We carried out a field-based study on free-ranging dogs in India to understand the nature of maternal care. Our analysis reveals that mothers reduce investment in energy-intensive active care and increase passive care as the pups grow older, thereby keeping overall levels of care more or less constant over pup age. Using the patterns of mother–pup interactions, we define the different phases of maternal care behaviour.

Free-ranging dogs are a ubiquitous part of human habitations in many developing countries, leading a life of scavengers dependent on human wastes for survival. The effective management of free-ranging dogs calls for understanding of their population dynamics. Life expectancy at birth and early life mortality are important factors that shape life-histories of mammals. We carried out a five year-long census based study in seven locations of West Bengal, India, to understand the pattern of population growth and factors affecting early life mortality in free-ranging dogs. We observed high rates of mortality, with only ~19% of the 364 pups from 95 observed litters surviving till the reproductive age; 63% of total mortality being human influenced. While living near people increases resource availability for dogs, it also has deep adverse impacts on their population growth, making the dog-human relationship on streets highly complex.

Parent–offspring conflict (POC) theory provides an interesting premise for understanding social dynamics in facultatively social species. In free-ranging dogs, mothers increase conflict over extended parental care with their pups beyond the weaning stage. In this study, we investigated whether resource quality affects POC in the dogs that typically live in a highly competitive environment as scavengers. We built a theoretical model to predict the alternative options available to the mother in the context of food sharing with her pups when protein-rich food (meat) is provided, as compared to carbohydrate-rich food (biscuits). We fit the mothers’ response from experimental data to the model and show that the mothers choose a selfish strategy, which can in turn ensure higher lifetime reproductive success, while depriving the current litter access to better resources. These results have interesting implications for understanding the social dynamics of the dogs, and the emergence of facultative sociality in a species that evolved from strongly social ancestors. We speculate that the tendency of increased conflict in resource-rich conditions might have driven the process of domestication in the ancestors of dogs which defected from their groups in favour of richer resources around human settlements.