Observation of female-male mounting in the carrion crow.

In the biological sciences, sexual behaviours in non-human animals are traditionally investigated in the context of reproduction and direct fitness benefits. While the evolutionary functions of non-conceptive sexual behaviours ('socio-sexual behaviours') remain less well explored, these interactions and displays have been suggested to be important for shaping and maintaining social relationships. Here, we report an observation of a captive female carrion crow, Corvus corone corone, mounting her co-housed male partner. We highlight the importance of more systematic research, reporting, and discussions of rarely observed behaviours in social evolution research, including considerations for behaviours that transcend binary or heteronormative frameworks, for a more comprehensive understanding of non-conceptive socio-sexual behaviours.

Cognition: Crows are natural statisticians.

A new study shows that carrion crows use memorized reward associations in a combinatorial way to apply relative probabilistic information to optimize reward outcome. This for the first time shows that a corvid species can flexibly apply statistical inference during decision making.

Conceptualization, context, and comparison are key to understanding the evolution of fear.

The fearful ape hypothesis proposes that heightened fearfulness in humans is adaptive. However, despite its attractive anthropocentric narrative, the evidence presented for greater fearfulness in humans versus other apes is not sufficient to support this claim. Conceptualization, context, and comparison are strongly lacking in Grossmann's proposal, but are key to understanding variation in the fear response among individuals and species.

Assessing cats' (Felis catus) sensitivity to human pointing gestures.

A wide range of nonhuman animal species has been shown to be able to respond to human referential signals, such as pointing gestures. The aim of the present study was to replicate previous findings showing cats to be sensitive to human pointing cues (Miklósi et al., 2005). In our study, we presented two types of human pointing gestures-momentary ipsilateral (direct pointing) and momentary cross-body pointing. We tested nine rescue cats in a two-way object-choice task. On a group level, the success rate of cats was 74.4%. Cats performed significantly above chance level in both the ipsilateral and cross-body pointing conditions. Trial number, rewarded side, and type of gesture did not significantly affect the cats' performances in the experiment. On an individual level, five out of seven cats who completed 20 trials performed significantly above chance level. Two cats only completed 10 trials. One of them succeeded in eight and the other in six of these trials. The results of our study replicate previous findings of cats being responsive to human ipsilateral pointing cues and add additional knowledge about their ability to follow cross-body pointing cues. Our results highlight that a domestic species, socialized in a group setting, may possess heterospecific communication skills. Further research is needed to exclude alternative parsimonious explanations, such as local and stimulus enhancements. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Effects of severe anthropogenic disturbance on the heart rate and body temperature in free-living greylag geese (Anser anser).

Anthropogenic disturbances are a major concern for the welfare and conservation of wildlife. We recorded heart rate and body temperature of 20 free-living greylag geese in response to a major regularly re-occurring anthropogenic disturbance-New Year's Eve fireworks. Heart rate and body temperature were significantly higher in the first and second hour of the new year, compared with the same hour on the 31st of December, the average during December and the average during January. Heart rate and body temperature was not significantly affected by sex or age. From 0200 to 0300 onwards, 1st of January heart rates did not significantly differ from the other periods; however, body temperatures were significantly increased until 0300-0400. From 0400 to 0500, heart rate was not affected by any of the investigated factors, whereas body temperature was significantly increased on the 1st of January compared with the 31st of December and the December average but not compared with the January average. To conclude, our results show that New Year's Eve fireworks cause a substantial physiological response, indicative of a stress response in greylag geese, which is costly in terms of energy expenditure.

Learning and motor inhibitory control in crows and domestic chickens.

Cognitive abilities allow animals to navigate through complex, fluctuating environments. In the present study, we tested the performance of a captive group of eight crows, Corvus corone and 10 domestic chickens, Gallus gallus domesticus, in the cylinder task, as a test of motor inhibitory control and reversal learning as a measure of learning ability and behavioural flexibility. Four crows and nine chickens completed the cylinder task, eight crows and six chickens completed the reversal learning experiment. Crows performed better in the cylinder task compared with chickens. In the reversal learning experiment, species did not significantly differ in the number of trials until the learning criterion was reached. The performance in the reversal learning experiment did not correlate with performance in the cylinder task in chickens. Our results suggest crows to possess better motor inhibitory control compared with chickens. By contrast, learning performance in a reversal learning task did not differ between the species, indicating similar levels of behavioural flexibility. Interestingly, we describe notable individual differences in performance. We stress the importance not only to compare cognitive performance between species but also between individuals of the same species when investigating the evolution of cognitive skills.

Heart rate as a measure of emotional arousal in evolutionary biology.

How individuals interact with their environment and respond to changes is a key area of research in evolutionary biology. A physiological parameter that provides an instant proxy for the activation of the automatic nervous system, and can be measured relatively easily, is modulation of heart rate. Over the past four decades, heart rate has been used to assess emotional arousal in non-human animals in a variety of contexts, including social behaviour, animal cognition, animal welfare and animal personality. In this review, I summarize how measuring heart rate has provided new insights into how social animals cope with challenges in their environment. I assess the advantages and limitations of different technologies used to measure heart rate in this context, including wearable heart rate belts and implantable transmitters, and provide an overview of prospective research avenues using established and new technologies, with a special focus on implications for applied research on animal welfare. This article is part of the theme issue 'Measuring physiology in free-living animals (Part II)'.

Testing for Behavioral and Physiological Responses of Domestic Horses (Equus caballus) Across Different Contexts - Consistency Over Time and Effects of Context.

In a number of species, consistent behavioral differences between individuals have been described in standardized tests, e.g., novel object, open field test. Different behavioral expressions are reflective of different coping strategies of individuals in stressful situations. A causal link between behavioral responses and the activation of the physiological stress response is assumed but not thoroughly studied. Also, most standard paradigms investigating individual behavioral differences are framed in a fearful context, therefore the present study aimed to add a test in a more positive context, the feeding context. We assessed individual differences in physiological [heart rate (HR)] and behavioral responses (presence or absence of pawing, startle response, defecation, snorting) of 20 domestic horses (Equus caballus) in two behavioral experiments, a novel object presentation and a pre-feeding excitement test. Experiments were conducted twice, once between July and August, and once between September and October. Both experiments caused higher mean HR in the first 10 s after stimulus presentation compared to a control condition, but mean HR did not differ between the experimental conditions. In the novel object experiment, horses displaying stress-related behaviors during the experiments also showed a significantly higher HR increase compared to horses which did not display any stress-related behaviors, reflecting a correlation between behavioral and physiological responses to the novel object. On the contrary, in the pre-feeding experiments, horses that showed fewer behavioral responses had a greater HR increase, indicating the physiological response being due to emotional arousal and not behavioral activity. Moreover, HR response to experimental situations varied significantly between individuals. Individual average HR was significantly repeatable across both experiments, whereas HR increase was only significantly repeatable during the novel object and not the pre-feeding experiment. Conversely, behavioral response was not repeatable. In conclusion, our findings show that horses' behavioral and physiological responses differed between test situations and that emotional reactivity, shown via mean HR and HR increase, is not always displayed behaviorally, suggesting that behavioral and physiological responses may be regulated independently according to context.

Parental behaviour and family proximity as key to gosling survival in Greylag Geese (Anser anser).

Reproductive success in monogamous species is generally affected by both behavioural and hormonal fine-tuning between pair partners. Vigilance, defence and brooding of offspring are among the main parental investments, and often the sexes adopt different roles. In the present study, we investigate how sex differences in parental behaviour and family proximity in the socially monogamous Greylag Goose (Anser anser) affect gosling survival. During the reproductive season in spring 2013, we recorded the behaviour of 18 pairs with offspring and gosling survival in a semi-tame, long-term monitored, and individually marked flock of Greylag Geese in Grünau, Austria. We found that behavioural role differentiation between the parents varied with developmental phase, and thus with gosling age. Especially during the first 10 days after hatching, females were foraging more frequently than males, which were more vigilant and aggressive towards other flock members. Such differences between the sexes levelled out 20 to 30 days after hatching. In general, females stayed in closer proximity to their offspring than males. Gosling survival was high when the parents were relatively aggressive and emphasized vigilance rather than foraging behaviour. Hence, we show a direct link between pair partners' quality of parental investment and gosling survival.

Self-control in crows, parrots and nonhuman primates.

Self-control is critical for both humans and nonhuman animals because it underlies complex cognitive abilities, such as decision-making and future planning, enabling goal-directed behavior. For instance, it is positively associated with social competence and life success measures in humans. We present the first review of delay of gratification as a measure of self-control in nonhuman primates, corvids (crow family) and psittacines (parrot order): disparate groups that show comparable advanced cognitive abilities and similar socio-ecological factors. We compare delay of gratification performance and identify key issues and outstanding areas for future research, including finding the best measures and drivers of delayed gratification. Our review therefore contributes to our understanding of both delayed gratification as a measure of self-control and of complex cognition in animals. This article is categorized under: Cognitive Biology > Evolutionary Roots of Cognition Psychology > Comparative Psychology.

Monkeys (Sapajus apella and Macaca tonkeana) and great apes (Gorilla gorilla, Pongo abelii, Pan paniscus, and Pan troglodytes) play for the highest bid.

Many studies investigate the decisions made by animals by focusing on their attitudes toward risk, that is, risk-seeking, risk neutrality, or risk aversion. However, little attention has been paid to the extent to which individuals understand the different odds of outcomes. In a previous gambling task involving 18 different lotteries (Pelé, Broihanne, Thierry, Call, & Dufour, 2014), nonhuman primates used probabilities of gains and losses to make their decision. Although the use of complex mathematical calculation for decision-making seemed unlikely, we applied a gradual decrease in the chances to win throughout the experiment. This probably facilitated the extraction of information about odds. Here, we investigated whether individuals would still make efficient decisions if this facilitating factor was removed. To do so, we randomized the order of presentation of the 18 lotteries. Individuals from 4 ape and 2 monkey species were tested. Only capuchin monkeys differed from others, gambling even when there was nothing to win. Randomizing the lottery presentation order leads all species to predominantly use a maximax heuristic. Individuals gamble as soon as there is at least one chance to win more than they already possess, whatever the risk. Most species also gambled more as the frequency of larger rewards increased. These results suggest optimistic behavior. The maximax heuristic is sometimes observed in human managerial and financial decision-making, where risk is ignored for potential gains, however low they may be. This suggests a shared and strong propensity in primates to rely on heuristics whenever complexity in evaluation of outcome odds arises. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

How does cognition shape social relationships?

The requirements of living in social groups, and forming and maintaining social relationships are hypothesized to be one of the major drivers behind the evolution of cognitive abilities. Most empirical studies investigating the relationships between sociality and cognition compare cognitive performance between species living in systems that differ in social complexity. In this review, we ask whether and how individuals benefit from cognitive skills in their social interactions. Cognitive abilities, such as perception, attention, learning, memory, and inhibitory control, aid in forming and maintaining social relationships. We investigate whether there is evidence that individual variation in these abilities influences individual variation in social relationships. We then consider the evolutionary consequences of the interaction between sociality and cognitive ability to address whether bi-directional relationships exist between the two, such that cognition can both shape and be shaped by social interactions and the social environment. In doing so, we suggest that social network analysis is emerging as a powerful tool that can be used to test for directional causal relationships between sociality and cognition. Overall, our review highlights the importance of investigating individual variation in cognition to understand how it shapes the patterns of social relationships.This article is part of the theme issue 'Causes and consequences of individual differences in cognitive abilities'.

Temporal modification of social interactions in response to changing group demographics and offspring maturation in African lions (Panthera leo).

In group living animals, affiliative social interactions maintain cohesion between individuals. Involvement in these interactions is likely to differ between individuals, depending on their sex, age and life history stages. Here we investigated different social network measures to describe greeting interactions within two prides of captive-origin African lions (Panthera leo). We aimed to determine if the introduction of cubs to these prides altered the strength of greeting networks among female lions. We also tested if the strength of greeting interactions changed between the age classes as younger lions matured. We found that interactions amongst female lions decreased from the period before cubs were born (least square means [95% CIs] 15.3 [7.67-22.93]) compared to after their integration into a pride (5.63 [-1.99-13.26] χ21 = 210.03, p < 0.001). In contrast, greeting network strength increased as offspring matured, with adults directing more interactions towards younger lions from 30 months of age (12.89 [1.96-23.82]) compared to when offspring were cubs (-0.69 [-11.63--10.24] χ210 = 156.03, p < 0.001). We suggest that social interactions between age classes may mediate recruitment and dispersal in lion prides.

Free-living greylag geese adjust their heart rates and body core temperatures to season and reproductive context.

Animals adaptively regulate their metabolic rate and hence energy expenditure over the annual cycle to cope with energetic challenges. We studied energy management in greylag geese. In all geese, profound seasonal changes of heart rate (fH) and body temperature (Tb) showed peaks in summer and troughs during winter, and also daily modulation of fH and Tb. Daily mean fH was on average 22% lower at the winter trough than at the summer peak, whereas daily mean Tb at the winter trough was only about 1 °C below the summer peak. Daily means of Tb together with those of air temperature and day length were the most important predictors of daily mean fH, which was further modulated by precipitation, reproductive state, and, to a minor degree, social rank. Peaks of fH and Tb occurred earlier in incubating females compared to males. Leading goslings increased daily mean fH. Our results suggest that in greylag geese, pronounced changes of fH over the year are caused by photoperiod-induced changes of endogenous heat production. Similar to large non-hibernating mammals, tolerance of lower Tb during winter seems the major factor permitting this. On top of these major seasonal changes, fH and Tb are elevated in incubating females.

Seasonal differences of corticosterone metabolite concentrations and parasite burden in northern bald ibis (Geronticus eremita): The role of affiliative interactions.

The reproductive season is energetically costly as revealed by elevated glucocorticoid concentrations, constrained immune functions and an increased risk of infections. Social allies and affiliative interactions may buffer physiological stress responses and thereby alleviate associated effects. In the present study, we investigated the seasonal differences of immune reactive corticosterone metabolite concentrations, endoparasite burden (nematode eggs and coccidian oocysts) and affiliative interactions in northern bald ibis (Geronticus eremita), a critically endangered bird. In total, 43 individually marked focal animals from a free-ranging colony were investigated. The analyses included a description of initiated and received affiliative interactions, pair bond status as well as seasonal patterns of hormone and endoparasite levels. During the reproductive season, droppings contained parasite eggs more often and corticosterone metabolite levels were higher as compared to the period after reproduction. The excretion rate of endoparasite products was lower in paired individuals than in unpaired ones, but paired animals exhibited higher corticosterone metabolite concentrations than unpaired individuals. Furthermore, paired individuals initiated affiliative behaviour more frequently than unpaired ones. This suggests that the reproductive season influences the excretion patterns of endoparasite products and corticosterone metabolites and that affiliative interactions between pair partners may positively affect endoparasite burden during periods of elevated glucocorticoid levels. Being embedded in a pair bond may have a positive impact on individual immune system and parasite resistance.

Effects of mate separation in female and social isolation in male free-living Greylag geese on behavioural and physiological measures.

The presence of a social partner may significantly contribute to coping with stressful events, whereas dyadic separation generally increases glucocorticoid levels and, thereby, might also affect immune function and health. To study the covariation between social factors, immuno-reactive corticosterone metabolites, haematology and parasite product excretion patterns in a free-living, long-term monogamous bird, we separated pair mates in Greylag geese (Anser anser). We isolated the males of eight pairs for 48h to examine behavioural, adrenocortical, haematological and parasitological responses to mate removal in the female partners, and to social isolation in the males. Females showed no elevated levels of corticosterone metabolites in their droppings, but their haematocrit decreased during mate removal, whereas leucocyte number and heterophil/lymphocyte (H/L) ratio remained unchanged. In contrast, the socially isolated males excreted significantly elevated levels of corticosterone metabolites compared to baseline and showed a decrease in haematocrit as well as elevated leucocyte number and H/L ratio. In both sexes, the excretion of coccidian oocysts increased within 48h of the start of the separation, remained high one week after separation, and returned to baseline four weeks later. Described effects were generally more pronounced in males than in females. Our results suggest relatively swift potential health effects of mate loss and social isolation in an unfamiliar confinement in free-living geese.

Social and environmental factors modulate leucocyte profiles in free-living Greylag geese (Anser anser).

BACKGROUND: Blood parameters such as haematocrit or leucocyte counts are indicators of immune status and health, which can be affected, in a complex way, by exogenous as well as endogenous factors. Additionally, social context is known to be among the most potent stressors in group living individuals, therefore potentially influencing haematological parameters. However, with few exceptions, this potential causal relationship received only moderate scientific attention. METHODS: In a free-living and individually marked population of the highly social and long-lived Greylag goose, Anser anser, we relate variation in haematocrit (HCT), heterophils to lymphocytes ratio (H/L) and blood leucocyte counts to the following factors: intrinsic (sex, age, raising condition, i.e. goose- or hand-raised), social (pair-bond status, pair-bond duration and parental experience) and environmental (biologically relevant periods, ambient temperature) factors. Blood samples were collected repeatedly from a total of 105 focal birds during three biologically relevant seasons (winter flock, mating season, summer). RESULTS: We found significant relationships between haematological parameters and social as well as environmental factors. During the mating season, unpaired individuals had higher HCT compared to paired and family individuals and this pattern reversed in fall. Similarly, H/L ratio was positively related to pair-bond status in a seasonally dependent way, with highest values during mating and successful pairs had higher H/L ratio than unsuccessful ones. Also, absolute number of leucocytes tended to vary depending on raising condition in a seasonally dependent way. DISCUSSION: Haematology bears a great potential in ecological and behavioural studies on wild vertebrates. In sum, we found that HTC, H/L ratio and absolute number of leucocytes are modulated by social factors and conclude that they may be considered valid indicators of individual stress load.

Excretion patterns of coccidian oocysts and nematode eggs during the reproductive season in Northern Bald Ibis (Geronticus eremita).

Individual reproductive success largely depends on the ability to optimize behaviour, immune function and the physiological stress response. We have investigated correlations between behaviour, faecal steroid metabolites, immune parameters, parasite excretion patterns and reproductive output in a critically endangered avian species, the Northern Bald Ibis (Geronticus eremita). In particular, we related haematocrit, heterophil/lymphocyte ratio, excreted immune-reactive corticosterone metabolites and social behaviour with parasite excretion and two individual fitness parameters, namely, number of eggs laid and number of fledglings. We found that the frequency of excretion of parasites' oocysts and eggs tended to increase with ambient temperature. Paired individuals excreted significantly more samples containing nematode eggs than unpaired ones. The excretion of nematode eggs was also significantly more frequent in females than in males. Individuals with a high proportion of droppings containing coccidian oocysts were more often preened by their partners than individuals with lower excretion rates. We observed that the more eggs an individual incubated and the fewer offspring fledged, the higher the rates of excreted samples containing coccidian oocysts. Our results confirm that social behaviour, physiology and parasite burden are linked in a complex and context-dependent manner. They also contribute background information supporting future conservation programmes dealing with this critically endangered species.

Carrion crows learn to discriminate between calls of reliable and unreliable conspecifics.

Partner choice on the basis of an individual's reliability is expected to stabilize social interactions. In this experiment, we tested whether carrion crows (Corvus corone corone) learn to differentiate between calls of reliable or unreliable individuals. Crows were kept in an aviary that comprised four visually but not acoustically isolated compartments, separated by a central room. In an association phase, a dead crow placed in the central compartment was visible only to one of the four crow groups, whilst alert calls of a conspecific were played back. Therefore, these calls were reliable for that group, but unreliable for the three other groups. The procedure was repeated, using a different reliable caller for each group. In two test sessions, 1 month apart, reliable and unreliable model individuals were played back, but no dead crow was presented. We quantified birds' attention behaviour and the number of vocalisations emitted. In the association phase, crows were more attentive towards the reliable compared with the unreliable stimuli and called more in response to reliable compared to unreliable individuals. In the test and repeat phase, attention behaviour did not differ between reliability conditions, but the pattern of vocal behaviour reversed, with crows calling less frequent when listening to reliable compared with unreliable calls. Vocal responses of crows suggest that they can discriminate between reliable and unreliable callers.

With whom to dine? Ravens' responses to food-associated calls depend on individual characteristics of the caller.

Upon discovering food, common ravens, Corvus corax, produce far-reaching 'haa' calls or yells, which are individually distinct and signal food availability to conspecifics. Here, we investigated whether ravens respond differently to 'haa' calls of known and unknown individuals. In a paired playback design, we tested responses to 'haa' call sequences in a group containing individually marked free-ranging ravens. We simultaneously played call sequences of a male and a female raven in two different locations and varied familiarity (known or unknown to the local group). Ravens responded strongest to dyads containing familiar females, performing more scan flights above and by perching in trees near the respective speaker. Acoustic analysis of the calls used as stimuli showed no sex-, age- or familiarity-specific acoustic cues, but highly significant classification results at the individual level. Taken together, our findings indicate that ravens respond to individual characteristics in 'haa' calls, and choose whom to approach for feeding, i.e. join social allies and avoid dominant conspecifics. This is the first study to investigate responses to 'haa' calls under natural conditions in a wild population containing individually marked ravens.