Domestication is a selection process where animals become more docile and easier to handle than their wild counterparts. This selection for ‘tameness’ has been suggested to lead to several behavioral modifications in domesticated animals, such as becoming more social and more playful towards humans, and also less aggressive and less fearful than their wild counterparts. The first animal to be domesticated was the dog, which is well-known for their pro-social relationship with humans. In an attempt to understand which behavioral changes occurred when the dog was domesticated, we used wolf-dog hybrids as a proxy for wolves and compared their behavior with the behavior of German shepherds, Siberian huskies, and Alaskan malamutes, which were the main dog breeds they were mixed with. We found that wolf-dog hybrids were as social as the dogs and that they did not show any signs of aggression. While the wolf-dog hybrids were less playful than German shepherds, they had the same levels of human-directed playfulness as Siberian huskies and Alaskan malamutes. The long-lasting fear reactions of the wolf-dog hybrids in this study, while almost absent in the three dog breeds, suggest that this fearfulness is a heritage from wolves, while dogs have adapted during domestication to have less fearful reactions in human environments.

An overrepresentation of stepchildren as victims of filicide has been explained as a consequence of ‘discriminative parental solicitude’. The idea being that Darwinian selection has favoured parental love and concern only for biological children, and when such parental feelings are absent, as in stepparents, conflicts with a child could easier escalate to lethal violence. An alternative explanation for this overrepresentation of stepchildren is that risk factors for filicide, such as criminal behaviour and mental health problems, are more prevalent in stepparents. This study focused on paternal filicide in Sweden and investigated (i) if stepchildren are overrepresented as victims of filicide compared with biological children, (ii) if filicides are committed in a context that implies a ‘conflict with the child victim’ and (iii) if stepfathers and biological fathers differ in characteristics associated with filicide risk. The analyses showed that stepchildren were overrepresented as victims compared with children of fathers in families with two biological parents and this overrepresentation was even higher in young children. Children of single biological fathers and children of non-residential biological fathers were also overrepresented as victims of filicide. Less than 20 percent of the filicides were committed in the context of a ‘conflict with the child’ and in these cases only stepchildren were overrepresented as victims. In the population at large, both stepfathers and single biological fathers had higher rates of mental health problems, violent criminality and illegal possession of drugs compared with fathers in families with two biological parents.

Domesticated animals are generally assumed to display increased sociability toward humans compared to their wild ancestors. Dogs (Canis familiaris) have a remarkable ability to form social relationships with humans, including lasting attachment, a bond based on emotional dependency. Since it has been specifically suggested that the ability to form attachment with humans evolved post-domestication in dogs, attempts to quantify attachment behavior in wolves (Canis lupus) have subsequently been performed. However, while these rare wolf studies do highlight the potential for wolves to express human-directed attachment, the varied methods used and the contrasting results emphasize the need for further, standardized testing of wolves. Here, we used the standardized Strange Situation Test to investigate attachment behavior expressed in wolves and dogs hand-raised and socialized under standardized and identical conditions up until the age of testing. We found that 23-week-old wolves and dogs equally discriminated between a stranger and a familiar person, and expressed similar attachment behaviors toward a familiar person. Additionally, wolves, but not dogs, expressed significantly elevated stress-related behavior during the test, but this stress response was buffered by the presence of a familiar person. Together, our results suggest that wolves can show attachment behaviors toward humans comparable to those of dogs. Importantly, our findings demonstrate that the ability to form attachment with humans exists in relatives of the wild ancestor of dogs, thereby refuting claims that this phenotype evolved after dog domestication was initiated.

Domestication dramatically alters phenotypes across animal species. Standing variation among ancestral populations often drives phenotypic change during domestication, but some changes are caused by novel mutations. In dogs (Canis familiaris) specifically, it has been suggested that the ability to interpret social-communicative behavior expressed by humans originated post-domestication and this behavior is thus not expected to occur in wolves (Canis lupus). Here we report the observation of three 8-week-old wolf puppies spontaneously responding to social-communicative behaviors from an unfamiliar person by retrieving a ball. This behavioral expression in wolves has significant implications for our understanding and expectations of the genetic foundations of dog behavior. Importantly, our observations indicate that behavioral responses to human social-communicative cues are not unique to dogs. This suggests that, although probably rare, standing variation in the expression of human-directed behavior in ancestral populations could have been an important target for early selective pressures exerted during dog domestication.

Human-directed play behaviour is a distinct behavioural feature of domestic dogs. But the role that artificial selection for contemporary dog breeds has played for human-directed play behaviour remains elusive. Here, we investigate how human-directed play behaviour has evolved in relation to the selection for different functions, considering processes of shared ancestry and gene flow among the different breeds. We use the American Kennel Club (AKC) breed group categorization to reflect the major functional differences and combine this with observational data on human-directed play behaviour for over 132 breeds across 89 352 individuals from the Swedish Dog Mentality Assessment project. Our analyses demonstrate that ancestor dogs already showed intermediate levels of human-directed play behaviour, levels that are shared with several modern breed types. Herding and Sporting breeds display higher levels of human-directed play behaviour, statistically distinguishable from Non-sporting and Toy breeds. Our results suggest that human-directed play behaviour played a role in the early domestication of dogs and that subsequent artificial selection for function has been important for contemporary variation in a behavioural phenotype mediating the social bond with humans.

Among-population variance of phenotypic traits is of high relevance for understanding evolutionary mechanisms that operate in relatively short timescales, but various sources of nonindependence, such as common ancestry and gene flow, can hamper the interpretations. In this comparative analysis of 138 dog breeds, we demonstrate how such confounders can independently shape the evolution of a behavioural trait (human-directed play behaviour from the Dog Mentality Assessment project). We combined information on genetic relatedness and haplotype sharing to reflect common ancestry and gene flow, respectively, and entered these into a phylogenetic mixed model to partition the among-breed variance of human-directed play behaviour while also accounting for within-breed variance. We found that 75% of the among-breed variance was explained by overall genetic relatedness among breeds, whereas 15% could be attributed to haplotype sharing that arises from gene flow. Therefore, most of the differences in human-directed play behaviour among breeds have likely been caused by constraints of common ancestry as a likely consequence of past selection regimes. On the other hand, gene flow caused by crosses among breeds has played a minor, but not negligible role. Our study serves as an example of an analytical approach that can be applied to comparative situations where the effects of shared origin and gene flow require quantification and appropriate statistical control in a within-species/among-population framework. Altogether, our results suggest that the evolutionary history of dog breeds has left remarkable signatures on the among-breed variation of a behavioural phenotype.

Domestication is hypothesized to drive correlated responses in animal morphology, physiology and behaviour, a phenomenon known as the domestication syndrome. However, we currently lack quantitative confirmation that suites of behaviours are correlated during domestication. Here we evaluate the strength and direction of behavioural correlations among key prosocial (sociability, playfulness) and reactive (fearfulness, aggression) behaviours implicated in the domestication syndrome in 76,158 dogs representing 78 registered breeds. Consistent with the domestication syndrome hypothesis, behavioural correlations within prosocial and reactive categories demonstrated the expected direction-specificity across dogs. However, correlational strength varied between dog breeds representing early (ancient) and late (modern) stages of domestication, with ancient breeds exhibiting exaggerated correlations compared to modern breeds across prosocial and reactive behaviours. Our results suggest that suites of correlated behaviours have been temporally decoupled during dog domestication and that recent shifts in selection pressures in modern dog breeds affect the expression of domestication-related behaviours independently.

Selection of behavioral traits holds a prominent role in the domestication of animals, and domesticated species are generally assumed to express reduced fear and reactivity toward novel stimuli compared to their ancestral species. However, very few studies have explicitly tested this proposed link between domestication and reduced fear responses. Of the limited number of studies experimentally addressing the alterations of fear during domestication, the majority has been done on canids. These studies on foxes, wolves, and dogs suggest that decreased expression of fear in domesticated animals is linked to a domestication-driven delay in the first onset of fearful behavior during early ontogeny. Thus, wolves are expected to express exaggerated fearfulness earlier during ontogeny compared to dogs. However, while adult dogs are less fearful toward novelty than adult wolves and wolf-dog hybrids, consensus is lacking on when differences in fear expression arise in wolves and dogs. Here we present the first extended examination of fear development in hand-raised dogs and European gray wolves, using repeated novel object tests from 6 to 26 weeks of age. Contrary to expectations, we found no evidence in support of an increase in fearfulness in wolves with age or a delayed onset of fear response in dogs compared to wolves. Instead, we found that dogs strongly reduced their fear response in the period between 6 and 26 weeks of age, resulting in a significant species difference in fear expression toward novelty from the age of 18 weeks. Critically, as wolves did not differ in their fear response toward novelty over time, the detected species difference was caused solely by a progressive reduced fear response in dogs. Our results thereby suggest that species differences in fear of novelty between wolves and dogs are not caused by a domestication-driven shift in the first onset of fear response. Instead, we suggest that a loss of sensitivity toward novelty with age in dogs causes the difference in fear expression toward novelty in wolves and dogs.

The domestication of animals and plants offers an exceptional opportunity to study evolutionary adaptations. In particular, domesticated animals display several behavioral alterations, including increased sociability and decreased fearfulness and aggression, when compared with their wild ancestors. However, studies quantifying simultaneous changes in multiple behaviors during domestication are lacking. Moreover, the role of human-directed play behavior has been largely neglected when studying the domestication process. Here we address these issues by examining behavioral changes during the domestication of the dog (Canis familiaris) from the gray wolf (Canis lupus) using a standardized behavioral test applied to wolf hybrids and several dog breeds. Contrary to expectations, our study provides little support for collective behavioral alterations. Specifically, although we found that wolf hybrids were less playful and overall more fearful than dogs, we did not detect any differences in sociability or aggression between wolf hybrids and dog breeds. Instead, our results suggest that behavioral alterations during domestication do not necessarily occur in concert and point to an important, but previously overlooked, role of selection on play behavior directed at humans during the domestication of dogs.