Does novelty influence the foraging decisions of a scavenger?

Acquiring knowledge about the environment is crucial for survival. Animals, often driven by their exploratory tendencies, gather valuable information regarding food resources, shelter, mating partners, etc. However, neophobia, or avoiding novel environmental stimuli, can constrain their exploratory behaviour. While neophobia can reduce potential predation risks, decreased exploratory behaviour resulting from it may limit the ability to discover highly rewarding resources. Dogs (Canis familiaris) living in semi-urban and urban environments as free-ranging populations, although subject to various selection forces, typically have negligible predation pressure. These dogs are scavengers in human-dominated environments; thus, selection against object-neophobia can provide benefits when searching for novel food resources. Although captive pack-living dogs are known to be less neophobic than their closest living ancestors, wolves (Canis lupus), little is known about free-ranging dogs' behavioural responses to novel objects, particularly in foraging contexts. Using an object choice experiment, we tested 259 free-ranging dogs from two age classes, adult and juvenile, to investigate their object-neophobia in a scavenging context. We employed a between-subject study design, providing dogs with a familiar and a potentially novel object, both baited with equal, hidden food items. Adult and juvenile dogs significantly inspected the novel object first compared to the familiar one, even when the hidden food item was partially visible. To validate these findings, we compared novel objects with different strengths of olfactory cues (baited vs. false-baited) and found that they were inspected comparably by adults and juveniles. No significant differences were found in the latencies to inspect the objects, suggesting that free-ranging dogs may still be cautious when exploring their environments. These results indicate that free-ranging dogs, evidently from an early ontogenetic phase, do not show object-neophobia, as demonstrated by their preference for novel over familiar food sources. We conclude that little to no constraint of neophobia on exploratory behaviour in semi-urban and urban-dwelling animals can guide foraging decision-making processes, providing adaptive benefits.

Adjustment in the point-following behaviour of free-ranging dogs - roles of social petting and informative-deceptive nature of cues.

Animals of different taxa can read and respond to various human communicative signals. Such a mechanism facilitates animals to acquire social information and helps them react in a context-dependent manner. Dogs have garnered extensive attention owing to their socio-cognitive skills and remarkable sensitivity to human social cues. For example, dogs readily respond to different human pointing gestures to locate hidden food rewards. However, a general inclination towards testing highly socialized pet dogs has resulted in a dearth of information on other sub-populations of dogs. Free-ranging dogs are one of the least socialized dog populations yet exhibit point-following behaviour flexibly. As a consequence of frequent negative interspecific interactions, they are typically wary of unfamiliar humans; thus, contextual recognition of human actions is paramount for these dogs to avoid potential conflict. However, the mechanisms influencing their point-following behaviour remain unidentified. We asked to what extent the informative-deceptive nature of cues and positive human interactions influence the interspecific communicative behaviour of these minimally socialized dogs. Using a point-following experiment with a 2 × 2 design, we focused on adult free-ranging dogs' behavioural adjustments. Dogs were randomly divided into two groups, with only one receiving brief social petting. Further, informative and deceptive cues were given to separate subsets within each group. Our findings suggest that brief social petting strongly affects the likelihood of free-ranging dogs' point-following tendencies. Dogs who received petting followed the pointing cues regardless of their informative or deceptive nature, whereas dogs who did not receive petting discriminated between informative and deceptive pointing. This study highlights the contribution of positive human interaction and informative-deceptive quality of cues in modulating the behavioural responses of free-ranging dogs in an interspecific communicative context.

Power-laws in dog behavior may pave the way to predictive models: A pattern analysis study.

Apparently random events in nature often reveal hidden patterns when analyzed using diverse and robust statistical tools. Power law distributions, for example, project diverse natural phenomenon, ranging from earthquakes to heartbeat dynamics into a common platform of self-similarity. Animal behavior in specific contexts has been shown to follow power law distributions. However, the behavioral repertoire of a species in its entirety has never been analyzed for the existence of such underlying patterns. Here we show that the frequency-rank data of randomly sighted behaviors at the population level of free-ranging dogs follow a scale-invariant power law behavior. It suggests that irrespective of changes in location of sightings, seasonal variations and observer bias, datasets exhibit a conserved trend of scale invariance. The data also exhibits robust self-similarity patterns at different scales which we extract using multifractal detrended fluctuation analysis. We observe that the probability of consecutive occurrence of behaviors of adjacent ranks is much higher than behaviors widely separated in rank. The findings open up the possibility of designing predictive models of behavior from correlations existing in true time series of behavioral data and exploring the general behavioral repertoire of a species for the presence of syntax.

Sociability of Indian free-ranging dogs (Canis lupus familiaris) varies with human movement in urban areas.

Cohabiting with humans in the same ecological space requires significant variation in the behavioral repertoire of animals. Behavioral variation can potentially improve the chances of survival of an individual. The influence of humans can be measured by quantifying specific behavioral parameters of the interacting individuals. Sociability or the tendency to be friendly toward others is one of many personality traits in animals that can provide us with insights regarding their relationship with humans. Free-ranging dogs are one of the successful urban-adapted species that interact with humans regularly, which, in turn, influences their behavioral properties. In this study, we tested 600 adult dogs from 60 sites across India, categorized as high, low, and intermediate human flux zones, to understand their degree of sociability toward an unfamiliar human. Initially, a "positive vocalization phase" was carried out. Unresponsive dogs were further tested in a "stimulus phase." The first phase was characterized by positive vocal sounds, while the second one included both food and positive vocalization. In addition, we surveyed a total of 1,200 people from the 60 sites to understand their perception of free-ranging dogs. Dogs in the IF zones were highly sociable compared to the other zones. High human flux zone dogs were reluctant to approach initially but showed an increased approach when food was provided. Low human flux zone dogs were the least sociable, and even the food reward had minimal impact on them. Our study provides the first evidence of behavioral variation in the degree of sociability of free-ranging dogs in urban environments. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Humans Dominate the Social Interaction Networks of Urban Free-Ranging Dogs in India.

Research on human-animal interaction has skyrocketed in the last decade. Rapid urbanization has led scientists to investigate its impact on several species living in the vicinity of humans. Domesticated dogs (Canis lupus familiaris) are one such species that interact with humans and are also called man's best friend. However, when it comes to the free-ranging population of dogs, interactions become quite complicated. Unfortunately, studies regarding free-ranging dog-human interactions are limited even though the majority of the world's dog population is free-ranging. In this study, we observed twelve groups of free-ranging dogs in their natural habitat, the streets. We quantified their interactions at the intra (dog-dog) and interspecific (dog-human) levels. The study areas were divided into two zones, namely - intermediate and high flux, based on human activity or movement. Social network analysis revealed higher instances of interspecific than intraspecific interactions, irrespective of the human flux zones. Humans, in significantly higher occasions, initiated both positive and negative behaviors in comparison to dogs. Our findings conclude that humans are a crucial part of the interaction network of Indian free-ranging dogs.

The effects of human attentional state on canine gazing behaviour: a comparison of free-ranging, shelter, and pet dogs.

The ability of animals to communicate using gaze is a rich area of research. How domestic dogs (Canis lupus familiaris) use and respond to the gaze of humans is an area of particular interest. This study examined how three groups of domestic dogs from different populations (free-ranging dogs, pet dogs, and shelter dogs) responded to a human during three attentional state conditions: when the human was making eye contact (attentive), when the human was turned away (inattentive), and when the human exited the testing area. We found that dogs from different populations differed in their gazing behaviour. Free-ranging dogs responded to the human's change in attentional state by looking significantly less at the human in the inattentive condition compared to the attentive condition. Pet and shelter dogs did not differ in their gazing behaviour between these conditions. However, they gazed significantly more at the human in both the inattentive and attentive conditions compared to the free-ranging dogs and also spent more time in the proximity of the experimenter. This study suggests that life experience plays an important role in how dogs respond to the attentional state of a human.

Free-Ranging Dogs Are Capable of Utilizing Complex Human Pointing Cues.

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 for understanding and responding to human referential gestures. The remarkable sociocognitive 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 setup. A closer inspection revealed that anxious behavioral 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.

The great Indian joint families of free-ranging dogs.

Cooperative breeding is an excellent example of 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.

Free-ranging dogs prefer petting over food in repeated interactions with unfamiliar humans.

Dogs (Canis lupus familiaris) are the first species to have been domesticated and, unlike other domesticated species, they have developed a special bond 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 dogs to investigate their response to immediate social and long-term food and social rewards. The dogs were provided a choice of obtaining food either from an 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 social contact. Our results revealed that these dogs tend to build trust based on affection, not food. This study provides significant insights into the dynamics of dog-human interactions on the streets and subsequent changes in behaviour of dogs through the process of learning.

Free-ranging dogs show age related plasticity in their ability to follow human pointing.

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.

Practice makes perfect: familiarity of task determines success in solvable tasks for free-ranging dogs (Canis lupus familiaris).

Domestic dogs' (Canis lupus familiaris) socio-cognitive faculties have made them highly sensitive to human social cues. While dogs often excel at understanding human communicative gestures, they perform comparatively poorly in problem-solving and physical reasoning tasks. This difference in their behaviour could be due to the lifestyle and intense socialization, where problem solving and physical cognition are less important than social cognition. Free-ranging dogs live in human-dominated environments, not under human supervision and are less socialized. Being scavengers, they often encounter challenges where problem solving is required in order to get access to food. We tested Indian street dogs in familiar and unfamiliar independent solvable tasks and quantified their persistence and dependence on a novel human experimenter, in addition to their success in solving a task. Our results indicate that free-ranging dogs succeeded and persisted more in the familiar task as compared to the unfamiliar one. They showed negligible amount of human dependence in the familiar task, but showed prolonged gazing and considerable begging behaviour to the human experimenter in the context of the unfamiliar task. Cognitive abilities of free-ranging dogs thus play a pivotal role in determining task-associated behaviours based on familiarity. In addition to that, these dogs inherently tend to socialize with and depend on humans, even if they are strangers. Our results also illustrate free-ranging dogs' low competence at physical cognitive tasks.

Differences in problem-solving between canid populations: Do domestication and lifetime experience affect persistence?

Past research has suggested that a variety of factors, phylogenetic and ontogenetic, play a role in how canines behave during problem-solving tasks and the degree to which the presence of a human influences their problem-solving behaviour. While comparisons between socialized wolves and domestic dogs have commonly been used to tease apart these predictive factors, in many cases a single dog population, often pets, have been used for these comparisons. Less is understood about how different populations of dogs may behave when compared with wolves, or with each other, during an independent problem-solving task. This experiment compared the independent persistence of four populations of canines (two groups of pet domestic dogs, a group of free-ranging domestic dogs, and human-socialized wolves) on an independent problem-solving task in the presence of an on looking human. Results showed that wolves persisted the most at the task while free-ranging dogs persisted the least. Free-ranging dogs gazed at the human experimenter for the longest durations during the task. While further research is needed to understand why these differences exist, this study demonstrates that dogs, even those living outside human homes as scavengers, show comparatively low levels of persistence when confronted with a solvable task in the presence of a human as well as significantly greater duration of human-directed gaze when compared with wolves.

Clever mothers balance time and effort in parental care: a study on free-ranging dogs.

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.

Selfish Pups: Weaning Conflict and Milk Theft in Free-Ranging Dogs.

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.

Denning habits of free-ranging dogs reveal preference for human proximity.

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.

High early life mortality in free-ranging dogs is largely influenced by humans.

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.

When Love Is in the Air: Understanding Why Dogs Tend to Mate when It Rains.

Seasonality of reproduction is observed in many species of organisms, across taxa, and is influenced by both biotic and abiotic factors. While such seasonality is easy to understand in temperate species exposed to extreme climates, it is more difficult to explain in the tropics. In many tropical species offspring are born during the season of high precipitation, which also coincides with high resource availability. Interestingly, in India, free-ranging dogs seem to mate, and not whelp, when it rains--an observation that cannot be explained by the resource abundance hypothesis. We carried out an extensive study to identify the mating seasons of free-ranging dogs, and observed a strong correlation between both the incidence and frequency of mating related behaviours of dogs, and precipitation levels. There are two clear mating seasons, of which the primary mating season coincides with the monsoon (rainy season) and the secondary mating season coincides with the nor'westerlies in this part of India. We speculate that this strong correlation is an effect of chemistry, rather than biology. While male dogs can mate round the year, females come into estrous seasonally. In the urban environment, dogs are exposed to a lot of olfactory noise, which can dilute the signal present in sex pheromones of the females in heat. A shower leads to increased humidity and reduced temperature of the air, leading to intensification of pheromone signals that trigger a sexual response in the dogs.

Selfish mothers indeed! Resource-dependent conflict over extended parental care in free-ranging dogs.

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.

Selfish mothers? An empirical test of parent-offspring conflict over extended parental care.

Parent-offspring conflict (POC) theory is an interesting conceptual framework for understanding the dynamics of parental care. However, this theory is not easy to test empirically, as exact measures of parental investment in an experimental set-up are difficult to obtain. We have used free-ranging dogs Canis familiaris in India, to study POC in the context of extended parental care. We observed females and their pups in their natural habitat for the mother's tendency to share food given by humans with her pups in the weaning and post-weaning stages. Since these dogs are scavengers, and depend largely on human provided food for their sustenance, voluntary sharing of food by the mother with her pups is a good surrogate for extended parental care. Our behavioural observations convincingly demonstrate an increase of conflict and decrease of cooperation by the mother with her offspring over given food within a span of 4-6 weeks. We also demonstrate that the competition among the pups in a litter scales with litter size, an indicator of sib-sib competition.