Investigating the Function of Play Bows in Dog and Wolf Puppies (Canis lupus familiaris, Canis lupus occidentalis).

Animals utilize behavioral signals across a range of different contexts in order to communicate with others and produce probable behavioral outcomes. During play animals frequently adopt action patterns used in other contexts. Researchers have therefore hypothesized that play signals have evolved to clarify communicative intent. One highly stereotyped play signal is the canid play bow, but its function remains contested. In order to clarify how canid puppies use play bows, we used data on play bows in immature wolves (ages 2.7-7.8 months) and dogs (ages 2 to 5 months) to test hypotheses evaluated in a previous study of adult dogs. We found that young dogs used play bows similarly to adult dogs; play bows most often occurred after a brief pause in play followed by complementary highly active play states. However, while the relative number of play bows and total observation time was similar between dog and wolf puppies, wolves did not follow this behavioral pattern, as play bows were unsuccessful in eliciting further play activity by the partner. While some similarities for the function of play bows in dog and wolf puppies were documented, it appears that play bows may function differently in wolf puppies in regards to re-initiating play.

Investigating the function of play bows in adult pet dogs (Canis lupus familiaris).

Play bows are a common, highly stereotyped canine behavior widely considered to be a 'play signal,' but only one study has researched their function. Bekoff (1995) found that play bows function as behavioral modifiers to help clarify playful intent before or after easily misinterpretable behaviors, such as bite-shakes. To further examine the function of play bows, the current study analyzed five types of behaviors displayed by the bower and the partner immediately before and after a play bow during dyadic play. We found that play bows most often occurred after a brief pause in play. Synchronous behaviors by the bower and the partner, or vulnerable/escape behaviors by the bower (such as running away) and complementary offensive behaviors by the partner (such as chasing) occurred most often after the play bow. These results indicate that during adult dog dyadic play, play bows function to reinitiate play after a pause rather than to mediate offensive or ambiguous actions.

Rough-and-tumble play as a window on animal communication.

Rough-and-tumble play (RT) is a widespread phenomenon in mammals. Since it involves competition, whereby one animal attempts to gain advantage over another, RT runs the risk of escalation to serious fighting. Competition is typically curtailed by some degree of cooperation and different signals help negotiate potential mishaps during RT. This review provides a framework for such signals, showing that they range along two dimensions: one from signals borrowed from other functional contexts to those that are unique to play, and the other from purely emotional expressions to highly cognitive (intentional) constructions. Some animal taxa have exaggerated the emotional and cognitive interplay aspects of play signals, yielding admixtures of communication that have led to complex forms of RT. This complexity has been further exaggerated in some lineages by the development of specific novel gestures that can be used to negotiate playful mood and entice reluctant partners. Play-derived gestures may provide new mechanisms by which more sophisticated communication forms can evolve. Therefore, RT and playful communication provide a window into the study of social cognition, emotional regulation and the evolution of communication systems.

Rollovers during play: Complementary perspectives.

In this commentary, we compare and contrast Norman et al.s' findings on rollovers during dog play (Norman et al., 2015; the "target article") with our work on dog play fighting (Bauer and Smuts, 2007; Ward et al., 2008). We first review our major findings and then correct some errors in the target article's descriptions of our work. We then further explore the concept of "defensive" rollovers proposed in the target article. We conclude that a combination of the target article's approach and ours should inform future investigations of dog rollovers.

Quantity-based judgments in the domestic dog (Canis lupus familiaris).

We examined the ability of domestic dogs to choose the larger versus smaller quantity of food in two experiments. In experiment 1, we investigated the ability of 29 dogs (results from 18 dogs were used in the data analysis) to discriminate between two quantities of food presented in eight different combinations. Choices were simultaneously presented and visually available at the time of choice. Overall, subjects chose the larger quantity more often than the smaller quantity, but they found numerically close comparisons more difficult. In experiment 2, we tested two dogs from experiment 1 under three conditions. In condition 1, we used similar methods from experiment 1 and tested the dogs multiple times on the eight combinations from experiment 1 plus one additional combination. In conditions 2 and 3, the food was visually unavailable to the subjects at the time of choice, but in condition 2, food choices were viewed simultaneously before being made visually unavailable, and in condition 3, they were viewed successively. In these last two conditions, and especially in condition 3, the dogs had to keep track of quantities mentally in order to choose optimally. Subjects still chose the larger quantity more often than the smaller quantity when the food was not simultaneously visible at the time of choice. Olfactory cues and inadvertent cuing by the experimenter were excluded as mechanisms for choosing larger quantities. The results suggest that, like apes tested on similar tasks, some dogs can form internal representations and make mental comparisons of quantity.

A mechanism for the evolution of altruism among nonkin: positive assortment through environmental feedback.

The evolution of altruism often requires genetic similarity among interactors. For structured populations in which a social trait affects all group members, this entails positive assortment, meaning that cooperators and noncooperators tend to be segregated into different groups. Several authors have claimed that mechanisms other than common descent can produce positive assortment, but this claim has not been generally accepted. Here, we describe one such mechanism. The process of "environmental feedback" requires only that the cooperative trait affects the quality of the local environment and that individuals are more likely to leave low-quality than high-quality environments. We illustrate this dynamic using an agent-based spatial model of feeding restraint. Depending on parameter settings, results included both positive assortment (required for the evolution of altruism) and negative assortment (required for the evolution of spite). The mechanism of environmental feedback appears to be a general one that could play a role in the evolution of many forms of cooperation.

Reproduction in wild female olive baboons.

The purpose of this paper is to evaluate several factors that influence female reproduction in a large troop of wild olive baboons (Papio cynocephalus anubis) based on 4 consecutive years of demographic data. Interbirth intervals were significantly shorter for females whose infants died before their next conception than for females whose infants survived. High-ranking mothers of surviving infants had significantly shorter birth intervals than comparable low-ranking mothers, independent of maternal age. This occurred mainly because the interval from resumption of cycling to conception was significantly shorter for high-vs. low-ranking females. Dominance rank did not influence sex ratio at birth, infant survival in the first 2 years, or adult female mortality. Age was also significantly related to interbirth intervals, with older females having shorter intervals. Primiparous females had consistently longer reproductive intervals than did multiparous females, but this difference reached statistical significance only for females whose infants died before the next conception. Primiparous females also experienced significantly higher infant mortality. Data on body size and estrous cycle length indicated no differences between high- and low-ranking females. Nutritional and stress-related mechanisms that may underlie the reproductive advantages of high rank are discussed.