Do dogs live in joint families? Understanding allo-parental care in free-ranging dogs

📝 Abstract

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 biology of dogs. We conducted a five year long study on parental care of free-ranging dogs in India. We observed widespread alloparenting by both adult males and females. Allomothers provided significantly less care that the mothers, but the putative fathers showed comparable levels of care with the mothers. However, the nature of care varied; mothers invested more effort in feeding and allogrooming, while the putative fathers played and protected more. We were unsure of the relatedness of the pups with the putative fathers, but all the allomothers were maternal relatives of the pups, which provides support for both the benefit-of-philopatry and assured fitness returns hypotheses. Free-ranging dogs are not cooperative breeders like wolves, but are more similar to communal breeders. Their breeding biology bears interesting similarities with the human joint family system.

💡 Analysis

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 biology of dogs. We conducted a five year long study on parental care of free-ranging dogs in India. We observed widespread alloparenting by both adult males and females. Allomothers provided significantly less care that the mothers, but the putative fathers showed comparable levels of care with the mothers. However, the nature of care varied; mothers invested more effort in feeding and allogrooming, while the putative fathers played and protected more. We were unsure of the relatedness of the pups with the putative fathers, but all the allomothers were maternal relatives of the pups, which provides support for both the benefit-of-philopatry and assured fitness returns hypotheses. Free-ranging dogs are not cooperative breeders like wolves, but are more similar to communal breeders. Their breeding biology bears interesting similarities with the human joint family system.

📄 Content

Do dogs live in joint families? Understanding allo-parental care in free-ranging dogs

Manabi Paul1 and Anindita Bhadra1,*

1Behaviour and Ecology Lab, Department of Biological Sciences, Indian Institute of Science Education and Research Kolkata, India

*Behaviour and Ecology Lab, Department of Biological Sciences,
Indian Institute of Science Education and Research Kolkata Mohanpur Campus, Mohanpur,
PIN 741246, West Bengal, INDIA
tel. 91-33-66340000-1223 fax +91-33-25873020 e-mail: abhadra@iiserkol.ac.in

Abstract 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 biology of dogs. We conducted a five year-long study on parental care of free-ranging dogs in India. We observed widespread allo-parenting by both adult males and females. Allomothers provided significantly less care that the mothers, but the putative fathers showed comparable levels of care with the mothers. However, the nature of care varied; mothers invested more effort in feeding and allogrooming, while the putative fathers played and protected more. We were unsure of the relatedness of the pups with the putative fathers, but all the allomothers were maternal relatives of the pups, which provides support for both the “benefit-of-philopatry” and “assured fitness returns” hypotheses. Free-ranging dogs are not cooperative breeders like wolves, but are more similar to communal breeders. Their breeding biology bears interesting similarities with the human joint family system.

Keywords: free-ranging dogs; maternal care; allomother; putative father; kin selection; philopatry

Introduction Cooperation and conflict drive the dynamics of social groups in species as diverse as insects to humans. While selfishness is easily explained by the theory of natural selection (1,2), cooperation and altruism are more difficult to understand as behavioural traits (3). The most extreme form of cooperation in animal societies is manifested by cooperative breeding, where only a few individuals in a social group reproduce and the others help to rear their offspring while forfeiting reproduction themselves (4,5). Cooperative breeding is observed in social insects like ants, bees, wasps and termites, but is relatively less common in mammals. Among mammals, naked mole rats, lemurs and some species of canids like wolves and coyotes are well known examples of cooperative breeders (5–9).Cooperative breeding is sometimes confused with communal breeding, though these are quite distinct breeding systems. In communally breeding species, multiple females share dens or birthing sites and help each other to rear offspring (6,8). Thus the reproductive hierarchy evident in cooperative breeders is absent in communal breeders.

For the last few decades, cooperative breeding has been mostly explained by kin selection theory (10,11). It has been argued that the benefits gained by the alloparent through inclusive fitness are not enough to compensate for their own reproduction (12). Therefore kin selection is not a sufficient condition for the evolution of cooperative breeding. However, kin selection, in conjugation with philopatry, i.e., the tendency of an individual to stay in or return to a particular area, leading to an increased probability of kin living in close proximity, could explain the evolution of altruism in animal groups (13). Females of most group-living mammals are reported to be philopatric (14) and the average kinship between females is highest for smaller groups (15,16). Hence cooperative breeding could be a consequence of kin selection and philopatry in such groups; for example, subordinates of a wolf pack are usually philopatric offspring of the dominant breeding pair (17,18) that share a common territory and exhibit cooperative breeding (5).

Domestic dogs (Canis lupus familiaris) share a common ancestry with modern day gray wolves (Canis lupus lupus) (19), but show much variation in their social organization. They are capable of living solitarily as pets, in artificially formed groups as pack dogs, and as natural social groups in free-ranging populations (20–22). They have a promiscuous mating system that lacks reproductive hierarchy (23,24). Their group dynamics greatly depend on their mating and denning seasons (22), and often multiple females of a group give birth in neighbouring dens. Maternal care is the predominant form of care received by the pups (25) where mothers adjust their care with pup age and litter sizes (Paul et al. submitted). They are predominantly scavengers, surviving on human-generated wastes, but a

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