We present a model of cultural evolution in which an individual's propensity to engage in social learning is affected by social learning itself. We assume that individuals observe cultural traits displayed by others and decide whether to copy them based on their overall preference for the displayed traits. Preferences, too, can be transmitted between individuals. Our results show that such cultural dynamics tends to produce conservative individuals, i.e., individuals who are reluctant to copy new traits. Openness to new information, however, can be maintained when individuals need significant time to acquire the cultural traits that make them effective cultural models. We show that a gradual enculturation of young individuals by many models and a larger cultural repertoire to be acquired are favorable circumstances for the long-term maintenance of openness in individuals and groups. Our results agree with data about lifetime personality change, showing that openness to new information decreases with age. Our results show that cultural remodeling of cultural transmission is a powerful force in cultural evolution, i.e., that cultural evolution can change its own dynamics

We call "regulatory traits" those cultural traits that are transmitted through cultural interactions and, at the same time, change individual behaviors directly influencing the outcome of future cultural interactions. The cultural dynamics of some of those traits are studied through simple simulations. In particular, we consider the cultural evolution of traits determining the propensity to copy, the number of potential demonstrators from whom one individual may copy, and conformist versus anti conformist attitudes. Our results show that regulatory traits generate peculiar dynamics that may explain complex human cultural phenomena. We discuss how the existence and importance of regulatory traits in cultural evolution impact on the analogy between genetic and cultural evolution and therefore on the possibility of using evolutionary biology inspired models to study human cultural dynamics.

Flight ability is generally expected to increase with relative flight muscle mass. Changes in weight can therefore be expected to influence the capacity to rapidly take-off, which can determine mating success and predator avoidance. This study examined the influence of relative flight muscle mass, sex, and season on free take-off flight ability in a butterfly model (Aglais urticae) that undergoes adult winter hibernation. Mass change and take-off flight ability (velocity and take-off angle), was predicted to fluctuate with season (before, during and after hibernation) and sex (due to reproductive investment). Our results indeed showed changes in take-off ability in relation to both parameters. Females maintained velocity across seasons but reduced take-off angles during and after hibernation. Male flight speed increased during and after hibernation, whereas take-off angles were significantly reduced during hibernation. Finally, we showed that investment in relative flight muscle mass increased velocity in female, but not in male butterflies.

Knowing the abundance of a population is a crucial component to assess its conservationstatus and develop effective conservation plans. For most cetaceans, abundanceestimation is difficult given their cryptic and mobile nature, especially when thepopulation is small and has a transnational distribution. In the Baltic Sea, the numberof harbour porpoises (Phocoena phocoena) has collapsed since the mid-20thcenturyand the Baltic Proper harbour porpoise is listed as Critically Endangered by the IUCNand HELCOM; however, its abundance remains unknown. Here, one of the largestever passive acoustic monitoring studies was carried out by eight Baltic Sea nationsto estimate the abundance of the Baltic Proper harbour porpoise for the first time. Bylogging porpoise echolocation signals at 298 stations during May 2011–April2013,calibrating the loggers’ spatial detection performance at sea, and measuring the clickrate of tagged individuals, we estimated an abundance of 71–1105individuals (95% CI,point estimate 491) during May–Octoberwithin the population's proposed managementborder. The small abundance estimate strongly supports that the Baltic Properharbour porpoise is facing an extremely high risk of extinction, and highlights theneed for immediate and efficient conservation actions through international cooperation.It also provides a starting point in monitoring the trend of the populationabundance to evaluate the effectiveness of management measures and determine itsinteractions with the larger neighboring Belt Sea population. Further, we offer evidencethat design-basedpassive acoustic monitoring can generate reliable estimatesof the abundance of rare and cryptic animal populations across large spatial scales.

Aposematic color patterns that signal prey unprofitability are suggested to work best when there is high contrast within the animal color pattern or between the animal and its background. Studies show that prey contrast against the background increases the signal efficiency. This has occasionally been extended to also explain the presence of internal patterns. We used domestic chicks, Gallus gallus domesticus, to investigate the relative importance for avoidance learning of within-prey pattern contrast and prey contrast against the background. In a series of trials, birds were first trained to avoid artificially made aposematic mealworms that were plain red or red with black stripes, and to discriminate them from palatable brown mealworms, on either a red or a brown background. Second, we investigated how the birds generalized between striped and nonstriped prey. The chicks showed faster avoidance learning when the basic color of the aposematic prey (red) contrasted with the background color (brown). However, there was no similar effect of internal pattern contrast. The generalization test showed a complete generalization between the nonstriped and the striped prey. We conclude that contrasting internal patterns do not necessarily affect predator avoidance learning the same way as shown for prey-to-background contrast in aposematic prey.

Recent studies have examined the ecological and evolutionary bases for variation in animal personality. However, only a few such studies have examined how foraging parameters are influenced by different personality domains. In wild ungulates, the trade-off between the time spent on food intake and antipredator behaviour differs between individuals, but the underlying reason for this is not yet well understood. One possibility is that this trade-off reflects personality dimensions such as boldness. To relate foraging decisions to personality we measured personality and performed feeding experiments with familiar and novel food in familiar and novel situations. We measured personality traits in 15 tame fallow deer, using novel object tests (NO), behavioural observations (BO) and personality ratings (PR). Boldness dimensions were found using PR and NO, dominance dimensions were found using BO and PR, and a flexibility dimension was found using BO. Multitrait–multimethod analysis showed that similar dimensions were significantly correlated across different methods and that different dimensions were not significantly correlated, even if measured using the same method. We also found that novel food eaten in familiar situations and familiar food eaten in novel situations were strongly related to boldness but not dominance, flexibility or age. Thus the trade-off between the benefits of gaining more food and the costs of reduced vigilance or increased toxin ingestion reflect boldness. These findings highlight the nature of personality dimensions in ungulates and how boldness impacts foraging behaviour.

In a previous study, we found that juvenile northern wheatears (Oenanthe oenanthe) exposed to a magnetic displacement to the west of their natural migration route increased their body mass. The total intensity and inclination used for the western displacement may also have been interpreted as northern compared to the experimental site (stronger total field intensity and steeper inclination angle). In order to investigate whether the fuelling response was a response to an unexpected magnetic field or specific to the northern magnetic field, we conducted a new experiment. Juvenile wheatears from the same study population were magnetically displaced to southwestern magnetic fields, exposing the birds to unexpected magnetic combinations, but eliminating the possible effect of a northern magnetic field. A control group was kept in the local geomagnetic field in Sweden for comparison. There was no difference in body mass increase between treatments, suggesting that the fuelling response previously found was not a simple response to an unexpected magnetic field, but rather a specific response to the northern magnetic field. Juvenile wheatears may have developed a fuelling response to northern magnetic fields in order to enable a successful flight towards the migration goal.

Recent experiments exposing migratory birds to altered magnetic fields simulating geographical displacements have shown that the geomagnetic field acts as an external cue affecting migratory fuelling behaviour. This is the first study investigating fuel deposition in relation to geomagnetic cues in long-distance migrants using the western passage of the Mediterranean region. Juvenile wheatears (Oenanthe oenanthe) were exposed to a magnetically simulated autumn migration from southern Sweden to West Africa. Birds displaced parallel to the west of their natural migration route, simulating an unnatural flight over the Atlantic Ocean, increased their fuel deposition compared to birds experiencing a simulated migration along the natural route. These birds, on the other hand, showed relatively low fuel loads in agreement with earlier data on wheatears trapped during stopover. The experimental displacement to the west, corresponding to novel sites in the Atlantic Ocean, led to a simulated longer distance to the wintering area, probably explaining the observed larger fuel loads. Our data verify previous results suggesting that migratory birds use geomagnetic cues for fuelling decisions and, for the first time, show that birds, on their first migration, can use geomagnetic cues to compensate for a displacement outside their normal migratory route, by adjusting fuel deposition.

Over recent decades, substantial research has focused on fish cognitive evolution to increase our understanding of the evolution of the enormous diversity of cognitive abilities that exists in fishes. One important but understudied aspect of cognitive evolution is sexual dimorphism in cognitive abilities. Sex-specific variation in brain region morphology has been proposed to be an important mechanism in this context. However, it is also common to find sex-specific variation in behavior and cognition without associated differences in brain morphology among the sexes. The telencephalon is the major cognitive center in the vertebrate brain and variation in telencephalon size has been associated with variation in cognition. Here, we utilize recently developed guppy artificial selection lines with ca. 10% differences in relative telencephalon size to investigate whether similar responses to selection of the size of this region may affect cognitive abilities differently in males and females. To that end, we compared two ecologically relevant aspects of cognition, detour learning and binary spatial discrimination. We tested the significance of the interaction between telencephalon size and sex, and we found no sex-specific effects of evolutionary increases in telencephalon size in the cognitive abilities tested. This study indicates that no clear cognitive sex-specific effects occur in response to rapid selection of telencephalon size. We suggest that future research on sexual dimorphism in cognitive abilities in fish could use various cognitive tests and examine telencephalic sub-regions to gain a more comprehensive understanding of their evolution.

Brain size varies substantially across the animal kingdom and is often associated with cognitive ability; however, the genetic architecture underpinning natural variation in these key traits is virtually unknown. In order to identify the genetic architecture and loci underlying variation in brain size, we analysed both coding sequence and expression for all the loci expressed in the telencephalon in replicate populations of guppies (Poecilia reticulata) artificially selected for large and small relative brain size. A single gene, Angiopoietin-1 (Ang-1), a regulator of angiogenesis and suspected driver of neural development, was differentially expressed between large-and small-brain populations. Zebra fish (Danio rerio) morphants showed that mild knock down of Ang-1 produces a small-brained phenotype that could be rescued with Ang-1 mRNA. Translation inhibition of Ang-1 resulted in smaller brains in larvae and increased expression of Notch-1, which regulates differentiation of neural stem cells. In situ analysis of newborn large-and small-brained guppies revealed matching expression patterns of Ang-1 and Notch-1 to those observed in zebrafish larvae. Taken together, our results suggest that the genetic architecture affecting brain size in our population may be surprisingly simple, and Ang-1 may be a potentially important locus in the evolution of vertebrate brain size and cognitive ability.

One of the most spectacular displays of social behavior is the synchronized movements that many animal groups perform to travel, forage and escape from predators. However, elucidating the neural mechanisms underlying the evolution of collective behaviors, as well as their fitness effects, remains challenging. Here, we study collective motion patterns with and without predation threat and predator inspection behavior in guppies experimentally selected for divergence in polarization, an important ecological driver of coordinated movement in fish. We find that groups from artificially selected lines remain more polarized than control groups in the presence of a threat. Neuroanatomical measurements of polarization-selected individuals indicate changes in brain regions previously suggested to be important regulators of perception, fear and attention, and motor response. Additional visual acuity and temporal resolution tests performed in polarization-selected and control individuals indicate that observed differences in predator inspection and schooling behavior should not be attributable to changes in visual perception, but rather are more likely the result of the more efficient relay of sensory input in the brain of polarization-selected fish. Our findings highlight that brain morphology may play a fundamental role in the evolution of coordinated movement and anti-predator behavior.

The organization and coordination of fish schools provide a valuable model to investigate the genetic architecture of affiliative behaviours and dissect the mechanisms underlying social behaviours and personalities. Here we used replicate guppy selection lines that vary in schooling propensity and combine quantitative genetics with genomic and transcriptomic analyses to investigate the genetic basis of sociability phenotypes. We show that consistent with findings in collective motion patterns, experimental evolution of schooling propensity increased the sociability of female, but not male, guppies when swimming with unfamiliar conspecifics. This finding highlights a relevant link between coordinated motion and sociability for species forming fission–fusion societies in which both group size and the type of social interactions are dynamic across space and time. We further show that alignment and attraction, the two major traits forming the sociability personality axis in this species, showed heritability estimates at the upper end of the range previously described for social behaviours, with important variation across sexes. The results from both Pool-seq and RNA-seq data indicated that genes involved in neuron migration and synaptic function were instrumental in the evolution of sociability, highlighting a crucial role of glutamatergic synaptic function and calcium-dependent signalling processes in the evolution of schooling.

The Human Evolutionary Transition offers a unified view of the evolution of intelligence, presenting a bold and provocative new account of how animals and humans have followed two powerful yet very different evolutionary paths to intelligence. This incisive book shows how animals rely on robust associative mechanisms that are guided by genetic information, which enable animals to sidestep complex problems in learning and decision making but ultimately limit what they can learn. Humans embody an evolutionary transition to a different kind of intelligence, one that relies on behavioral and mental flexibility. The book argues that flexibility is useless to most animals because they lack sufficient opportunities to learn new behavioral and mental skills. Humans find these opportunities in lengthy childhoods and through culture. Blending the latest findings in fields ranging from psychology to evolutionary anthropology, The Human Evolutionary Transition draws on computational analyses of the problems organisms face, extensive overviews of empirical data on animal and human learning, and mathematical modeling and computer simulations of hypotheses about intelligence. This compelling book demonstrates that animal and human intelligence evolved from similar selection pressures while identifying bottlenecks in evolution that may explain why human-like intelligence is so rare.

The ability to acquire knowledge and skills from others is widespread in animals and is commonly thought to be responsible for the behavioural traditions observed in many species. However, in spite of the extensive literature on theoretical analyses and empirical studies of social learning, little attention has been given to whether individuals acquire knowledge from a single individual or multiple models. Researchers commonly refer to instances of sons learning from fathers, or daughters from mothers, while theoreticians have constructed models of uniparental transmission, with little consideration of whether such restricted modes of transmission are actually feasible. We used mathematical models to demonstrate that the conditions under which learning from a single cultural parent can lead to stable culture are surprisingly restricted (the same reasoning applies to a single social-learning event). Conversely, we demonstrate how learning from more than one cultural parent can establish culture, and find that cultural traits will reach a nonzero equilibrium in the population provided the product of the fidelity of social learning and the number of cultural parents exceeds 1. We discuss the implications of the analysis for interpreting various findings in the animal social-learning literature, as well as the unique features of human culture.

Sexual ornaments used during mate choice can be costly to produce and maintain. The expression of such costly sexual traits is expected be dependent on the acquisition and allocation of resources, such as food, and therefore offer an honest signal of individual quality during mate choice. While cost of male ornaments, and how these ornaments impact female choice, is well studied, far less attention is directed towards resource-dependent expression of female ornaments and their role in shaping male mate choice decisions. Here, we examine how experimental resource restriction influences the expression of female sexual ornaments in the pygmy halfbeak (Dermogenys collettei). Specifically, we focus examine how dietary restriction influences the size and coloration of the female’s gravid spot, an orange ornament that is expressed on females’ abdomen that is used by males during mate choice. We find that the gravid spot reduced in size in females maintained on a restricted quantity diet, while it was maintained in females fed a high quantity diet. Females in the restricted diet treatment also had reduced red and green reflectance of the gravid spot. We then investigated if this resource-dependent expression a female sexual ornament influences male mate choice. As predicted, male halfbeaks show a preference for females with larger sexual ornaments. This suggests that resource quantity can explain variation in female ornament expression in halfbeaks and that female ornaments are costly. Moreover, female ornaments can serve as honest signals of quality and be used by males during mate choice, opening the door to indirect benefits of male mate choice. 

Social animals can greatly benefit from well-developed social skills. Because the frequency and diversity of social interactions often increase with the size of social groups, the benefits of advanced social skills can be expected to increase with group size. Variation in social skills often arises during ontogeny, depending on early social experience. Whether variation of social-group sizes affects development of social skills and related changes in brain structures remains unexplored. We investigated whether, in a cooperatively breeding cichlid, early group size (1) shapes social behavior and social skills and (2) induces lasting plastic changes in gross brain structures and (3) whether the development of social skills is confined to a sensitive ontogenetic period. Rearing-group size and the time juveniles spent in these groups interactively influenced the development of social skills and the relative sizes of four main brain regions. We did not detect a sensitive developmental period for the shaping of social behavior within the 2-month experience phase. Instead, our results suggest continuous plastic behavioral changes over time. We discuss how developmental effects on social behavior and brain architecture may adaptively tune phenotypes to their current or future environments.

Migratory birds wintering in Africa face the challenge of passing the Sahara desert with little opportunities to forage. During spring migration birds thus arrive in the Mediterranean area after crossing the desert with very low energy reserves. Since early arrival to the breeding grounds often is of importance to maximize reproductive success, finding stopover sites with good refuelling possibilities after the Saharan passage is of utmost importance. Here we report on extensive fuelling in the great reed warbler, Acrocephalus arundinaceus, on the south coast of Crete in spring, the first land that they encounter after crossing the Sahara desert and the Mediterranean Sea in this area. Birds were trapped with mist nets at a river mouth, individually ringed and information about body mass, wing length, muscle score and fat score were recorded. Due to an exceptional high recapture rate at the trapping site (45%), we were able to calculate minimum stopover time and fuel deposition rates in 25 individual great reed warblers during one spring season. The large proportion of trapped great reed warbler compared to other species and the large number of recaptures suggests that great reed warbler actively choose this area for stopover. The relatively long stopover period at the site, the high fuel deposition rate (1 g day-1) and the large body mass increase show that great reed warblers at this site regularly deposit a much larger fuel load than needed for one continued flight stage to the north. It was also shown that birds with lower body mass at first capture had a higher fuel deposition rate than birds with higher body mass. This indicates that individuals are able to adjust their food intake in relation to energy reserves.

Aposematic animals are often conspicuous. It has been hypothesized that one function of conspicuousness in such prey is to be detected from afar by potential predators: the ‘detection distance hypothesis’. The hypothesis states that predators are less prone to attack at long detection range because more time is allowed for making the ‘correct’ decision not to attack the unprofitable prey. The detection distance hypothesis has gained some experimental support in that time-limited predators make more mistakes. To investigate effects of prey presentation distance we performed two experiments. First, in experiment 1, we investigated at what distance chicks, Gallus gallus domesticus, could see the difference in colour between aposematic and plain mealworms. Birds chose the correct track in a two-way choice when prey were at 20, 40 and 60 cm distance but not at 80 cm. Second, in experiment 2, fifth-instar larvae of the aposematic bug Lygaeus equestris were presented to experienced chicks at 2, 20 or 60 cm distance. We found no difference in attack probability between distances. However, prey mortality was significantly lower for the shortest presentation distance. In conclusion, we found no support for the hypothesis that aposematic prey benefit from long-range detection; in fact they benefit from shortdistance detection. This result, and others, suggests that the conspicuousness of aposematic prey at a distance may simply be a by-product of an efficient signalling function after detection.

 Summary: In many anuran species, larvae modify their developmental trajectories and behaviour in response to chemical cues that predict predator risk. Recent reviews highlight a dearth of studies on delayed (post-metamorphic) consequences of larval experience.