Metabolic rate associates with, but does not generate covariation between, behaviours in western stutter-trilling crickets, Gryllus integer.

The causes and consequences of among-individual variation and covariation in behaviours are of substantial interest to behavioural ecology, but the proximate mechanisms underpinning this (co)variation are still unclear. Previous research suggests metabolic rate as a potential proximate mechanism to explain behavioural covariation. We measured the resting metabolic rate (RMR), boldness and exploration in western stutter-trilling crickets, Gryllus integer, selected differentially for short and fast development over two generations. After applying mixed-effects models to reveal the sign of the covariation, we applied structural equation models to an individual-level covariance matrix to examine whether the RMR generates covariation between the measured behaviours. All traits showed among-individual variation and covariation: RMR and boldness were positively correlated, RMR and exploration were negatively correlated, and boldness and exploration were negatively correlated. However, the RMR was not a causal factor generating covariation between boldness and exploration. Instead, the covariation between all three traits was explained by another, unmeasured mechanism. The selection lines differed from each other in all measured traits and significantly affected the covariance matrix structure between the traits, suggesting that there is a genetic component in the trait integration. Our results emphasize that interpretations made solely from the correlation matrix might be misleading.

A dark cuticle allows higher investment in immunity, longevity and fecundity in a beetle upon a simulated parasite attack.

Cuticle melanism in insects is linked to a number of life history traits: a positive relationship is hypothesized between melanism, immune function, fecundity and lifespan. However, it is not clear how activation of the immune system affects trade-offs between life history traits in female mealworm beetles (Tenebrio molitor) differing in cuticle melanization. The females with tan, brown and black cuticles examined in the present study did not differ in the intensity of encapsulation response, fecundity and longevity when their immune system was not activated. However, we found that immune activation and cuticle melanization have a significant effect on life history traits. Offspring number and lifespan decreased in females with tan and brown cuticles, while the fecundity and lifespan of black females were not affected. Importantly, we inserted the implants again and found a significant decrease in the strength of encapsulation response in females with tan and brown cuticles. In contrast, black females increased melanotic reactions against the nylon implant, suggesting immunological priming. The results show that cuticle melanization plays an important adaptive role under the risk of being infected, while the lack of these benefits before the insertion of nylon monofilaments suggests that there are costs associated with an activated immunity system.

Why can't we all just get along? Integration needs more than stories.

As astutely noted by the authors of this provocative article, it is time for evolutionary psychology (EP) to be incorporated into clinical and educational interventions. However, two issues from this article are raised in the current commentary: some historical misconceptions of the evolutionary label and a lack of clear and specific guidelines for developing or improving interventions based on EP.

Animal play and evolution: Seven timely research issues about enigmatic phenomena.

The nature of play in animals has been long debated, but progress is being made in characterizing play and its variants, documenting its distribution across vertebrate and invertebrate taxa, describing its mechanisms and development, and proposing testable theories about its origins, evolution, and adaptive functions. To achieve a deeper understanding of the functions and evolution of play, integrative and conceptual advances are needed in neuroscience, computer modeling, phylogenetics, experimental techniques, behavior development, and inter- and intra-specific variation. The special issue contains papers documenting many of these advances. Here, we describe seven timely areas where further research is needed to understand this still enigmatic class of phenomena more fully. Growing empirical and theoretical evidence reveals that play has been crucial in the evolution of behavior and psychology but has been underestimated, if not ignored, in both empirical and theoretical areas of evolutionary biology and neuroscience. Play research has important ramifications for understanding the evolution of cognition, emotion, and culture, and research on animals can be both informative and transformative.

Social learning in Cartilaginous fish (stingrays Potamotrygon falkneri).

Social learning is considered one of the hallmarks of cognition. Observers learn from demonstrators that a particular behavior pattern leads to a specific consequence or outcome, which may be either positive or negative. In the last few years, social learning has been studied in a variety of taxa including birds and bony fish. To date, there are few studies demonstrating learning processes in cartilaginous fish. Our study shows that the cartilaginous fish freshwater stingrays (Potamotrygon falkneri) are capable of social learning and isolates the processes involved. Using a task that required animals to learn to remove a food reward from a tube, we found that observers needed significantly (P < 0.01) fewer trials to learn to extract the reward than demonstrators. Furthermore, observers immediately showed a significantly (P < 0.05) higher frequency of the most efficient "suck and undulation" strategy exhibited by the experienced demonstrators, suggesting imitation. Shedding light on social learning processes in cartilaginous fish advances the systematic comparison of cognition between aquatic and terrestrial vertebrates and helps unravel the evolutionary origins of social cognition.

Play, animals, resources: the need for a rich (and challenging) comparative environment.

Van de Vliert proposes a comprehensive explanation for differences in "freedoms" in diverse human populations based on climate and monetary resources. This intriguing approach, though derived from an evolutionary view covering all species, is based exclusively on human populations. This anthropocentric lens is challenged by ways of testing Van de Vliert's thesis more generally using playfulness as a surrogate for freedom.

A review of interspecific social play among nonhuman animals.

Few species play socially with another species, hereafter called interspecific social play (ISP). ISP involves reading and responding appropriately to social cues of other species, often taxonomically remote, and has implications for perception, communication, and cognition. We reviewed information on non-human ISP from both print media and videos from YouTube and Reddit. We found over 200 instances of ISP. The literature predominantly featured wild primates, carnivores, and marine mammals. Carnivores and terrestrial ungulates were common in videos. ISP in avian and reptile species were found in both sources, including instances of playing with mammals. Animals may engage in ISP because it is risky and stimulating, they lack age-appropriate conspecifics, the play motivation is high, or to maintain social bonds in mixed-species groups. Cataloguing ISP uncovers which species are interacting and how. Systematic studies of ISP are difficult and many reports are brief and anecdotal. Minimally, future research should record information about each observation, including the age, sex, and history of participants.

Comparing the ontogeny, neurobiology, and function of social play in hamsters and rats.

Syrian hamsters show complex social play behavior and provide a valuable animal model for delineating the neurobiological mechanisms and functions of social play. In this review, we compare social play behavior of hamsters and rats and underlying neurobiological mechanisms. Juvenile rats play by competing for opportunities to pin one another and attack their partner's neck. A broad set of cortical, limbic, and striatal regions regulate the display of social play in rats. In hamsters, social play is characterized by attacks to the head in early puberty, which gradually transitions to the flanks in late puberty. The transition from juvenile social play to adult hamster aggression corresponds with engagement of neural ensembles controlling aggression. Play deprivation in rats and hamsters alters dendritic morphology in mPFC neurons and impairs flexible, context-dependent behavior in adulthood, which suggests these animals may have converged on a similar function for social play. Overall, dissecting the neurobiology of social play in hamsters and rats can provide a valuable comparative approach for evaluating the function of social play.

A Descriptive Methodology for Studying the Ontogeny of Object Play and Breed Differences in Dogs (Canis lupus familiaris).

Play behavior is a prominent aspect of juvenile behavior for many animals, yet early development, especially play with objects, has received little attention. Our previous study on object play introduced our general methods, focusing on litter differences in the developmental trajectory of object play and toy preferences. Here, we present a detailed ethogram of more than 30 observed object play behaviors. We focus on breed differences in the development of play in the three following breeds: Welsh Terriers, Vizslas, and standard Poodles. Puppies were video recorded from 3 to 7 weeks of age at half-week intervals upon the introduction of a standard set of five toys into their home environments. Ten minutes of video from each session for each puppy were analyzed using the Noldus Observer XT program. Aside from analyzing individual behaviors, they were also grouped into three behavioral categories. These were behaviors that occurred only in a solitary context, only in a social context, or in both contexts. Solitary object play developed first, and social object play developed later across breeds. There was a significant three-way interaction between breed, developmental age, and the context in which play occurred. Pairwise comparisons within each breed, age, and context are discussed, but a prominent result is that the onset of many behaviors occurred later in Welsh Terriers compared to the other breeds.

Consecutive virgin births in the new world boid snake, the Colombian rainbow Boa, Epicrates maurus.

Until recently, facultative automictic parthenogenesis within the squamate reptiles exhibiting ZZ:ZW genetic sex determination has resulted in single reproductive events producing male (ZZ) or female (ZW) offspring. With the recent discovery of viable parthenogenetically produced female (WW) Boa constrictors, the existence of further parthenogenetic events resulting in WW females was questioned. Here, we provide genetic evidence for consecutive virgin births by a female Colombian rainbow boa (Epicrates maurus), resulting in the production of WW females likely through terminal fusion automixis. Samples were screened at 22 microsatellite loci with 12 amplifying unambiguous products. Of these, maternal heterozygosity was observed in 4, with the offspring differentially homozygous at each locus. This study documents the first record of parthenogenesis within the genus Epicrates, a second within the serpent lineage Boidae, and the third genetically confirmed case of consecutive virgin births of viable offspring within any vertebrate lineage. Unlike the recent record in Boa constrictors, the female described here was isolated from conspecifics from birth, demonstrating that males are not required to stimulate parthenogenetic reproduction in this species and possibly other Boas.

How comparative was (is) the Journal of Comparative Psychology? A reptilian perspective.

[Correction Notice: An Erratum for this article was reported in Vol 135(3) of Journal of Comparative Psychology (see record 2021-87304-004). In the article "How Comparative Was (Is) the Journal of Comparative Psychology? A Reptilian Perspective" by Gordon M. Burghardt (Journal of Comparative Psychology. Advance online publication. August 5, 2021. http://doi.org/10.1037/com0000290), the phrase in the introduction that includes the Dewbury (1998) citation also includes an extra word. The phrase should appear as Dewbury (1998) noted that the focus. The year of publication for the Journal of Animal Behavior that appears in the third line of the Method section should appear as (1911-1917). The last sentence in the first paragraph of the Method section should appear as The 8,911 entries over this 110-year period constituted the data analyzed here in detail. The first sentence of the Results section should appear as The Journal of Animal Behavior published 238 articles in its 7-year run. The last phrase of the first paragraph of the Results section should appear as and 8,635 published items of the JCP and JCPP....] Comparative psychology, and particularly the Journal of Comparative Psychology, has been criticized for a lack of taxon diversity. The nature and consequences of the critiques are discussed and assessed by analyzing the representation of nonavian reptiles in the journal over its 100-year existence. Although reptiles are indeed rare in the journal, their representation has greatly increased in recent decades, and especially since about 1980. More interestingly, the mix among the major reptilian groups: turtles, lizards, snakes, and crocodylians, has shifted. First turtles predominated in studies, but in recent decades, snakes were far more prominent. In the last 50 years of the journal, there were 10 times the number of articles on snakes than in the first 50 years, turtles declined, and lizards increased greatly, although their totals remained less than half the number of snake articles. Crocodylians only appeared in the first several volumes in the 1920s and never again. The predominance of snakes, not known for their cognitive prowess, in a journal viewed increasingly as an outlet for work on comparative cognition, is discussed. Finally, it appears that the low representation of reptile behavioral research is not peculiar to the Journal of Comparative Psychology, but animal behavior journals more generally. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

A new method for studying problem solving and tool use in stingrays (Potamotrygon castexi).

Testing the cognitive abilities of cartilaginous fishes is important in understanding the evolutionary origins of cognitive functions in higher vertebrates. We used five South American fresh water stingrays (Potamotrygon castexi) in a learning and problem-solving task. A tube test apparatus was developed to provide a simple but sophisticated procedure for testing cognitive abilities of aquatic animals. All five subjects quickly learned to use water as a tool to extract food from the testing apparatus. The experimental protocol, which gave the animals the opportunity of correcting a wrong visual cue decision, resulted in four out of five subjects correcting an error rather than making an initial right choice. One of five subjects reached 100% correct trials in the visual discrimination task. The ability to use water as an agent to extract food from the testing apparatus is a first indication of tool use in batoid fishes. Performance in the instrumental task of retrieving food from a novel testing apparatus and the rapid learning in the subsequent discrimination/error correction task shows that cartilaginous fish can be used to study the origins of cognitive functions in the vertebrate lineage.

Behavior in Free-Living American Black Bear Dens: Parturition, Maternal Care, and Cub Behavior.

Denning behavior has long remained the least observed aspect of bear behavior. During 2010-2013, we used webcams, microphones, the internet, and 14,602 h of archived video to document the denning behaviors of two adult wild black bears (Ursus americanus) as they gave birth and cared for four litters through six winters in northeastern Minnesota. Observations included types of dens, labor, pre-parturient genital swelling, birthing positions, post-partum vocalizations, mothers removing amniotic tissues and warming newborn cubs in sub-freezing temperatures, frequency of nursing, cubs establishing nipple order, yearlings suckling, the ingestion of snow and icicles, the ingestion of foot pads, urination and defecation in latrine areas, toilet-licking, eye opening, reciprocal tongue-licking, play, rapid eye movement (REM) sleep and possible dreaming, and reactions to wildlife intruders. The use of this new method for observing natural bear dens allowed the identification of many behaviors undescribed for any species of wild bear in dens. We also discuss the need for future studies and how the depth and duration of black bear hibernation varies with body condition and geographic region.

Darwin's legacy to comparative psychology and ethology.

Charles Darwin made numerous seminal contributions to the study of animal behavior over his long career. This essay places these contributions in the context of Darwin's life, showing his long-standing interest in psychological and behavioral issues encompassing all species, including humans. Ten areas are highlighted: natural history; communication; sexual selection and courtship; comparative cognition; emotion; instinct and behavioral development; inheritance of behavior; phylogeny of behavior; sociobiology and behavioral ecology; and applied animal behavior, animal welfare, and conservation. Several newer emphases that Darwin anticipated are briefly discussed. Darwin, while not always correct by current standards, crucially aided the process of firmly embedding psychological phenomena in a naturalistic scientific ethos.

A place for emotions in behavior systems research.

Although issues of motivation, including appetitive searching behavior, have been crucial aspects of behavior systems approaches since their inception, as well as in the ethological research and models that inspired them, emotions and affect have been noticeably absent in such analyses. Emotions and affect may have been lying below the surface all the time, however, as motivation, emotion, and cognitive processing are embedded in all aspects of behavior, including conditioning and learning. Here a brief case is made that emotions and related hedonic processes, can and should be explicitly incorporated into behavior systems approaches. Evidence from recent behaviorist, neuroscience, and animal behavior (including human) studies suggest that emotions may, just as motivational drives, lead to appetitive searching and avoidance, both general and focal, as well as consummatory acts with emotional consequences and satiation. Research testing these claims, as well as theoretical formulations and evaluations, would be timely and extend the reach of behavior systems approaches to a far wider swath of psychological research than it has engaged with hitherto.

Development of social play in hamsters: Sex differences and their possible functions.

In several rodent species social play appears to be necessary for proper deployment of species-specific patterns of aggressive and reproductive behavior. Specifically, in male Syrian hamsters (Mesocricetus auratus), play has been linked to the development of adult aggression. We quantified several types of social play behavior in same-sex peer groups of Syrian hamsters three times per week for three consecutive weeks after weaning, which included postnatal days 22-42 (PD22 to PD42). Male hamsters increased playful contact during PD36-PD42, whereas females showed peak playful contact during PD29-PD35. These findings suggest that the motivation for social play increases during mid-adolescence in males, but dissipates in females. To investigate the effects of social play deprivation, one hamster per litter remained pair-housed with its mother forthree weeks after weaning its littermates. In adulthood, both play-deprived and play-exposed animals received acute social defeat stress followed by social interaction testing. Play deprivation led to increased defeat-induced social avoidance in both males and females. In males, play deprivation increased fighting back during social defeat stress, whereas in females it reduced aggressive behavior during conditioned defeat testing. We suggest that social play deprivation disrupts neural circuits regulating aggression in a sex-specific manner, perhaps related to sex differences in territorial defense, but has similar effects on neural circuits regulating stress responsivity. Overall, these findings suggest that juvenile social play functions to promote coping with stress and appropriate social behavior in adulthood.

Updating von Uexküll: new directions in communication research.

The study of communication in all its forms has come a long way conceptually and methodologically in the last decade, and this is more true of auditory communication than of the other sensory modalities. In considering the issues involved in communication, it is clear that simple dyadic considerations leave out too many important issues. Background noise, eavesdropping, and deception represent important neglected factors driving the evolution of the mechanisms, development, and ecological adaptiveness of communicatory processes in natural environments.

Wallace Craig's Appetites and aversions as constituents of instincts: A centennial appreciation.

Wallace Craig's "Appetites and Aversions as Constituents of Instincts," one of the seminal articles in animal behavior, comparative psychology, and ethology, appeared 100 years ago this year. The influence of this classic article is continuing and perhaps even expanding. Here we review the major ideas Craig offered in the article, provide a brief sketch of his scientific career and historical context, and examine the impact of the article on ethological and instinct theory. We also review some current applications of his work in diverse areas and discuss why it is cited more now than in earlier decades, especially in articles on causal mechanisms. (PsycINFO Database Record (c) 2018 APA, all rights reserved).

Toward a Theory of the Evolution of Fair Play.

Juvenile animals of many species engage in social play, but its functional significance is not well understood. This is especially true for a type of social play called fair play (Fp). Social play often involves behavioral patterns similar to adult behaviors (e.g., fighting, mating, and predatory activities), but young animals often engage in Fp behaviors such as role-reversals and self-handicapping, which raises the evolutionary problem of why Fp exists. A long-held working hypothesis, tracing back to the 19th century, is that social play provides contexts in which adult social skills needed for adulthood can be learned or, at least, refined. On this hypothesis, Fp may have evolved for adults to acquire skills for behaving fairly in the sense of equitable distribution of resources or treatment of others. We investigated the evolution of Fp using an evolutionary agent-based model of populations of social agents that learn adult fair behavior (Fb) by engaging in Fp as juveniles. In our model, adults produce offspring by accumulating resources over time through foraging. Adults can either behave selfishly by keeping the resources they forage or they can pool them, subsequently dividing the pooled resources after each round of foraging. We found that fairness as equitability was beneficial especially when resources were large but difficult to obtain and led to the evolution of Fp. We conclude by discussing the implications of this model, for developing more rigorous theory on the evolution of social play, and future directions for theory development by modeling the evolution of play.

Training and long-term memory of a novel food acquisition task in a turtle (Pseudemys nelsoni).

We developed a shaping procedure for training Florida red-bellied cooters, Pseudemys nelsoni, to dislodge clear plastic bottles to obtain food pellets. The animals were then trained in a 2-choice problem to choose only the bottle containing pellets. All nine turtles learned the task of knocking over bottles for food. For the discrimination task, turtles chose the correct bottle 71% on average. After 2 months (82-84 days), and again after another 7.5 months (228 days) of no interaction with the bottles, turtles were retested and many retained both the response and the discrimination (mean success rates 77-81%), with significant savings in retraining all turtles. The turtles showed two basic response strategies, which changed across time for some individuals. This study demonstrates that turtles can learn and retain a novel skill in a laboratory context.