The importance of migratory drop-off for island colonization in birds.

Seasonal migration is an underappreciated driver of animal diversification. Changes in migratory behaviour may favour the establishment of sedentary founder populations and promote speciation if there is sufficient reproductive isolation between sedentary and migratory populations. From a systematic literature review, we here quantify the role of migratory drop-off-the loss of migratory behaviour-in promoting speciation in birds on islands. We identify at least 157 independent colonization events likely initiated by migratory species that led to speciation, including 44 cases among recently extinct species. By comparing, for all islands, the proportion of island endemic species that derived from migratory drop-off with the proportion of migratory species among potential colonizers, we showed that seasonal migration has a larger effect on island endemic richness than direct dispersal. We also found that the role of migration in island colonization increases with the geographic isolation of islands. Furthermore, the success of speciation events depends in part on species biogeographic and ecological factors, here positively associated with greater range size and larger flock sizes. These results highlight the importance of shifts in migratory behaviour in the speciation process and calls for greater consideration of migratory drop-off in the biogeographic distribution of birds.

Proto-tool use for food processing in wild Arabian babblers: matching processing methods, substrates and prey types.

Cognition is a powerful adaptation, enabling animals to utilise resources that are unavailable without manipulation. Tool use and food processing are examples of using cognition to overcome the protective mechanisms of food resources. Here, we describe and examine the flexibility of proto-tool use (defined as the alteration of an object through object-substrate manipulation) for food processing in a cooperatively breeding bird, the Arabian babbler (Argya squamiceps). Field observations demonstrate that the birds transport different caterpillar species to different substrate types depending on the processing method needed to prepare the caterpillar for eating. Species with toxic setae (e.g. Casama innotata) are transported to be rubbed on rough substrates (e.g. sand) before consumption, while other species (e.g. Hyles livornica) are transported to be pounded against hard substrates until their inner organs are removed and only their external body part is consumed. These results are among the few to describe flexible proto-tool use for food processing in wild animals. They thereby contribute to the taxonomic mapping of proto-tool use and food processing in non-human species, which is a fundamental step to advance comparative studies on the evolution of these behaviours and their underlying cognitive mechanisms.

Family-living and cooperative breeding in birds are associated with the number of avian predators.

Cooperative breeding occurs when individuals contribute parental care to offspring that are not their own. Numerous intra- and inter-specific studies have aimed to explain the evolution of this behaviour. Recent comparative work suggests that family living (i.e., when offspring remain with their parents beyond independence) is a critical steppingstone in the evolution of cooperative breeding. Thus, it is key to understand the factors that facilitate the evolution of family living. Within-species studies suggest that protection from predators is a critical function of group living, through both passive benefits such as dilution effects, and active benefits such as prosocial antipredator behaviours in family groups. However, the association between predation risk and the formation and prevalence of family groups and cooperative breeding remains untested globally. Here we use phylogenetic comparative analyses including 2984 bird species to show that family living and cooperative breeding are associated with increased occurrence of avian predators. These cross-species findings lend support to previous suggestions based on intraspecific studies that social benefits of family living, such as protection against predation, could favour the evolution of delayed dispersal and cooperative breeding.

Does the expensive brain hypothesis apply to amphibians and reptiles?

Vertebrate brains show extensive variation in relative size. The expensive brain hypothesis argues that one important source of this variation is linked to a species' ability to generate the energy required to sustain the brain, especially during periods of unavoidable food scarcity. Here we ask whether this hypothesis, tested so far in endothermic vertebrates, also applies to ectotherms, where ambient temperature is an additional major aspect of energy balance. Phylogenetic comparative analyses of reptiles and amphibians support the hypothesis. First, relative brain size increases with higher body temperature in those species active during the day that can gain free energy by basking. Second, relative brain size is smaller among nocturnal species, which generally face less favorable energy budgets, especially when maintaining high body temperature. However, we do not find an effect of seasonal variation in ambient temperature or food on brain size, unlike in endotherms. We conclude that the factors affecting energy balance in ectotherms and endotherms are overlapping but not identical. We therefore discuss the idea that when body temperatures are seasonally very low, cognitive benefits may be thwarted and selection on larger brain size may be rare. Indeed, mammalian hibernators may show similarities to ectotherms.

Evidence for a reproductive sharing continuum in cooperatively breeding mammals and birds: consequences for comparative research.

Extreme reproductive skew occurs when a dominant female/male almost monopolizes reproduction within a group of multiple sexually mature females/males, respectively. It is sometimes considered an additional, restrictive criterion to define cooperative breeding. However, datasets that use this restrictive definition to classify species as cooperative breeders systematically overestimate reproductive skew by including groups in which reproduction cannot be shared by definition (e.g. groups with a single female/male). Here, we review the extent of reproductive sharing in 41 mammal and 37 bird species previously classified as exhibiting alloparental care and extreme reproductive skew, while only considering multi-female or multi-male groups. We demonstrate that in groups where unequal reproduction sharing is possible, extreme reproductive skew occurs in a few species only (11/41 mammal species and 12/37 bird species). These results call for significant changes in datasets that classify species' caring and mating system. To facilitate these changes, we provide an updated dataset on reproductive sharing in 63 cooperatively breeding species. At the conceptual level, our findings suggest that reproductive skew should not be a defining criterion of cooperative breeding and support the definition of cooperative breeding as a care system in which alloparents provide systematic care to other group members' offspring.

What is cooperative breeding in mammals and birds? Removing definitional barriers for comparative research.

Cooperative breeding (i.e. when alloparents care for the offspring of other group members) has been studied for nearly a century. Yet, inconsistent definitions of this breeding system still hamper comparative research. Here, we identify two major inconsistencies, discuss their consequences and propose a way forward. First, some researchers restrict the term 'cooperative breeding' to species with non-breeding alloparents. We show that such restrictive definitions lack distinct quantitative criteria to define non-breeding alloparents. This ambiguity, we argue, reflects the reproductive-sharing continuum among cooperatively breeding species. We therefore suggest that cooperative breeding should not be restricted to the few species with extreme reproductive skew and should be defined independent of the reproductive status of alloparents. Second, definitions rarely specify the type, extent and prevalence of alloparental care required to classify species as cooperative breeders. We thus analysed published data to propose qualitative and quantitative criteria for alloparental care. We conclude by proposing the following operational definition: cooperative breeding is a reproductive system where >5% of broods/litters in at least one population receive species-typical parental care and conspecifics provide proactive alloparental care that fulfils >5% of at least one type of the offspring's needs. This operational definition is designed to increase comparability across species and disciplines while allowing to study the intriguing phenomenon of cooperative breeding as a behaviour with multiple dimensions.

The effect of climate change on avian offspring production: A global meta-analysis.

Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young.

Extended parental provisioning and variation in vertebrate brain sizes.

Large brains provide adaptive cognitive benefits but require unusually high, near-constant energy inputs and become fully functional well after their growth is completed. Consequently, young of most larger-brained endotherms should not be able to independently support the growth and development of their own brains. This paradox is solved if the evolution of extended parental provisioning facilitated brain size evolution. Comparative studies indeed show that extended parental provisioning coevolved with brain size and that it may improve immature survival. The major role of extended parental provisioning supports the idea that the ability to sustain the costs of brains limited brain size evolution.

Parental provisioning drives brain size in birds.

Large brains support numerous cognitive adaptations and therefore may appear to be highly beneficial. Nonetheless, the high energetic costs of brain tissue may have prevented the evolution of large brains in many species. This problem may also have a developmental dimension: juveniles, with their immature and therefore poorly performing brains, would face a major energetic hurdle if they were to pay for the construction of their own brain, especially in larger-brained species. Here, we explore the possible role of parental provisioning for the development and evolution of adult brain size in birds. A comparative analysis of 1,176 bird species shows that various measures of parental provisioning (precocial vs. altricial state at hatching, relative egg mass, time spent provisioning the young) strongly predict relative brain size across species. The parental provisioning hypothesis also provides an explanation for the well-documented but so far unexplained pattern that altricial birds have larger brains than precocial ones. We therefore conclude that the evolution of parental provisioning allowed species to overcome the seemingly insurmountable energetic constraint on growing large brains, which in turn enabled bird species to increase survival and population stability. Because including adult eco- and socio-cognitive predictors only marginally improved the explanatory value of our models, these findings also suggest that the traditionally assessed cognitive abilities largely support successful parental provisioning. Our results therefore indicate that the cognitive adaptations underlying successful parental provisioning also provide the behavioral flexibility facilitating reproductive success and survival.

Contrasting coloured ventral wings are a visual collision avoidance signal in birds.

Collisions between fast-moving objects often cause severe damage, but collision avoidance mechanisms of fast-moving animals remain understudied. Particularly, birds can fly fast and often in large groups, raising the question of how individuals avoid in-flight collisions that are potentially lethal. We tested the collision-avoidance hypothesis, which proposes that conspicuously contrasting ventral wings are visual signals that help birds to avoid collisions. We scored the ventral wing contrasts for a global dataset of 1780 bird species. Phylogenetic comparative analyses showed that larger species had more contrasting ventral wings than smaller species, and that in larger species, colonial breeders had more contrasting ventral wings than non-colonial breeders. Evidently, larger species have lower manoeuvrability than smaller species, and colonial-breeding species frequently encounter con- and heterospecifics, increasing their risk of in-flight collisions. Thus, more contrasting ventral wing patterns in these species are a sensory mechanism that facilitates collision avoidance.

Forest management affects seasonal source-sink dynamics in a territorial, group-living bird.

The persistence of wildlife populations is under threat as a consequence of human activities, which are degrading natural ecosystems. Commercial forestry is the greatest threat to biodiversity in boreal forests. Forestry practices have degraded most available habitat, threatening the persistence of natural populations. Understanding population responses is, therefore, critical for their conservation. Population viability analyses are effective tools to predict population persistence under forestry management. However, quantifying the mechanisms driving population responses is complex as population dynamics vary temporally and spatially. Metapopulation dynamics are governed by local dynamics and spatial factors, potentially mediating the impacts of forestry e.g., through dispersal. Here, we performed a seasonal, spatially explicit population viability analysis, using long-term data from a group-living territorial bird (Siberian jay, Perisoreus infaustus). We quantified the effects of forest management on metapopulation dynamics, via forest type-specific demography and spatially explicit dispersal, and how forestry impacted the stability of metapopulation dynamics. Forestry reduced metapopulation growth and stability, through negative effects on reproduction and survival. Territories in higher quality natural forest contributed more to metapopulation dynamics than managed forests, largely through demographic processes rather than dispersal. Metapopulation dynamics in managed forest were also less resilient to disturbances and consequently, may be more vulnerable to environmental change. Seasonal differences in source-sink dynamics observed in managed forest, but not natural forests, were caused by associated seasonal differences in dispersal. As shown here, capturing seasonal source-sink dynamics allows us to predict population persistence under human disturbance and to provide targeted conservation recommendations.

Who do you trust? Wild birds use social knowledge to avoid being deceived.

Many species give deceptive warning calls, enabled by the high risk of ignoring them. In Siberian jays, a territorial, group-living bird, individuals give warning calls toward perched predators and mob them. However, intruding neighbors can emit these warning calls in the absence of predators to access food, but breeders often ignore these calls. Playback field experiments show that breeders flee sooner and return later after warning calls of former group members than those of neighbors or unknown individuals. Thus, breeders respond appropriately only to warning calls of previous cooperation partners. This mechanism facilitates the evolution and maintenance of communication vulnerable to deceptive signaling. This conclusion also applies to human language because of its cooperative nature and thus, its vulnerability to deception.

Sex-specific effects of cooperative breeding and colonial nesting on prosociality in corvids.

The investigation of prosocial behavior is of particular interest from an evolutionary perspective. Comparisons of prosociality across non-human animal species have, however, so far largely focused on primates, and their interpretation is hampered by the diversity of paradigms and procedures used. Here, we present the first systematic comparison of prosocial behavior across multiple species in a taxonomic group outside the primate order, namely the bird family Corvidae. We measured prosociality in eight corvid species, which vary in the expression of cooperative breeding and colonial nesting. We show that cooperative breeding is positively associated with prosocial behavior across species. Also, colonial nesting is associated with a stronger propensity for prosocial behavior, but only in males. The combined results of our study strongly suggest that both cooperative breeding and colonial nesting, which may both rely on heightened social tolerance at the nest, are likely evolutionary pathways to prosocial behavior in corvids.

Introduction of a New In-Situ Measurement System for the Study of Touch-Feel Relevant Surface Properties.

The touch-feel sensation of product surfaces arouses growing interest in various industry branches. To entangle the underlying physical and material parameters responsible for a specific touch-feel sensation, a new measurement system has been developed. This system aims to record the prime physical interaction parameters at a time, which is considered a necessary prerequisite for a successful physical description of the haptic sensation. The measurement setup enables one to measure the dynamic coefficient of friction, the macroscopic contact area of smooth and rough surfaces, the angle enclosed between the human finger and the soft-touch surfaces and the vibrations induced in the human finger during relative motion at a time. To validate the measurement stand, a test series has been conducted on two soft-touch surfaces of different roughness. While the individual results agree well with the literature, their combination revealed new insights. Finally, the investigation of the haptics of polymer coatings with the presented measuring system should facilitate the design of surfaces with tailor-made touch-feel properties.

Extended parenting and the evolution of cognition.

Traditional attempts to understand the evolution of human cognition compare humans with other primates. This research showed that relative brain size covaries with cognitive skills, while adaptations that buffer the developmental and energetic costs of large brains (e.g. allomaternal care), and ecological or social benefits of cognitive abilities, are critical for their evolution. To understand the drivers of cognitive adaptations, it is profitable to consider distant lineages with convergently evolved cognitions. Here, we examine the facilitators of cognitive evolution in corvid birds, where some species display cultural learning, with an emphasis on family life. We propose that extended parenting (protracted parent-offspring association) is pivotal in the evolution of cognition: it combines critical life-history, social and ecological conditions allowing for the development and maintenance of cognitive skillsets that confer fitness benefits to individuals. This novel hypothesis complements the extended childhood idea by considering the parents' role in juvenile development. Using phylogenetic comparative analyses, we show that corvids have larger body sizes, longer development times, extended parenting and larger relative brain sizes than other passerines. Case studies from two corvid species with different ecologies and social systems highlight the critical role of life-history features on juveniles' cognitive development: extended parenting provides a safe haven, access to tolerant role models, reliable learning opportunities and food, resulting in higher survival. The benefits of extended juvenile learning periods, over evolutionary time, lead to selection for expanded cognitive skillsets. Similarly, in our ancestors, cooperative breeding and increased group sizes facilitated learning and teaching. Our analyses highlight the critical role of life-history, ecological and social factors that underlie both extended parenting and expanded cognitive skillsets. This article is part of the theme issue 'Life history and learning: how childhood, caregiving and old age shape cognition and culture in humans and other animals'.

Remotely sensed forest understory density and nest predator occurrence interact to predict suitable breeding habitat and the occurrence of a resident boreal bird species.

Habitat suitability models (HSM) based on remotely sensed data are useful tools in conservation work. However, they typically use species occurrence data rather than robust demographic variables, and their predictive power is rarely evaluated. These shortcomings can result in misleading guidance for conservation. Here, we develop and evaluate a HSM based on correlates of long-term breeding success of an open nest building boreal forest bird, the Siberian jay. In our study site in northern Sweden, nest failure of this permanent resident species is driven mainly by visually hunting corvids that are associated with human settlements. Parents rely on understory nesting cover as protection against these predators. Accordingly, our HSM includes a light detection and ranging (LiDAR) based metric of understory density around the nest and the distance of the nest to the closest human settlement to predict breeding success. It reveals that a high understory density 15-80 m around nests is associated with increased breeding success in territories close to settlements (<1.5 km). Farther away from human settlements breeding success is highest at nest sites with a more open understory providing a favorable warmer microclimate. We validated this HSM by comparing the predicted breeding success with landscape-wide census data on Siberian jay occurrence. The correlation between breeding success and occurrence was strong up to 40 km around the study site. However, the HSM appears to overestimate breeding success in regions with a milder climate and therefore higher corvid numbers. Our findings suggest that maintaining patches of small diameter trees may provide a cost-effective way to restore the breeding habitat for Siberian jays up to 1.5 km from human settlements. This distance is expected to increase in the warmer, southern, and coastal range of the Siberian jay where the presence of other corvids is to a lesser extent restricted to settlements.

The syntax-semantics interface in animal vocal communication.

Syntax (rules for combining words or elements) and semantics (meaning of expressions) are two pivotal features of human language, and interaction between them allows us to generate a limitless number of meaningful expressions. While both features were traditionally thought to be unique to human language, research over the past four decades has revealed intriguing parallels in animal communication systems. Many birds and mammals produce specific calls with distinct meanings, and some species combine multiple meaningful calls into syntactically ordered sequences. However, it remains largely unclear whether, like phrases or sentences in human language, the meaning of these call sequences depends on both the meanings of the component calls and their syntactic order. Here, leveraging recently demonstrated examples of meaningful call combinations, we introduce a framework for exploring the interaction between syntax and semantics (i.e. the syntax-semantic interface) in animal vocal sequences. We outline methods to test the cognitive mechanisms underlying the production and perception of animal vocal sequences and suggest potential evolutionary scenarios for syntactic communication. We hope that this review will stimulate phenomenological studies on animal vocal sequences as well as experimental studies on the cognitive processes, which promise to provide further insights into the evolution of language. This article is part of the theme issue 'What can animal communication teach us about human language?'

Food sharing patterns in three species of callitrichid monkeys (Callithrix jacchus, Leontopithecus chrysomelas, Saguinus midas): Individual and species differences.

Food sharing (FS) in cooperatively breeding callitrichids is unusual among nonhuman primates because they regularly share significant amounts of preferred food with immatures and engage in proactive FS. However, it is still unclear which classes of individuals (males or females, breeders or helpers) engage most in FS, and whether differences exist among callitrichid species. In the first part of this study, we characterized general FS patterns in common marmosets (Callithrix jacchus). We found substantial adult-immature FS, and female breeders shared the least with immatures. This conflicts with previously published studies, where data were collected with the prevailing standard method of providing a food bowl to the entire group. In the second part, a comparison of our access-bias-free method and the standard method suggested that previous findings are likely the result of access bias. In the third part, we investigated species differences in adult-adult FS among common marmosets, golden-headed lion tamarins (Leontopithecus chrysomelas), and red-handed tamarins (Saguinus midas). As common marmosets show lower levels of interdependence within groups, we expected more adult-adult FS in tamarins compared with marmosets. Adult-adult FS was indeed more prevalent in tamarins and not exclusively directed from male breeders to female breeders. Therefore, our results suggest that adult-adult FS in marmosets mostly reflects the high energetic demands of female breeders, whereas in the more interdependent tamarins, it may be used to reinforce cooperative bonds between adult group members. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Call combinations in birds and the evolution of compositional syntax.

Syntax is the set of rules for combining words into phrases, providing the basis for the generative power of linguistic expressions. In human language, the principle of compositionality governs how words are combined into a larger unit, the meaning of which depends on both the meanings of the words and the way in which they are combined. This linguistic capability, i.e., compositional syntax, has long been considered a trait unique to human language. Here, we review recent studies on call combinations in a passerine bird, the Japanese tit (Parus minor), that provide the first firm evidence for compositional syntax in a nonhuman animal. While it has been suggested that the findings of these studies fail to provide evidence for compositionality in Japanese tits, this criticism is based on misunderstanding of experimental design, misrepresentation of the importance of word order in human syntax, and necessitating linguistic capabilities beyond those given by the standard definition of compositionality. We argue that research on avian call combinations has provided the first steps in elucidating how compositional expressions could have emerged in animal communication systems.

The interacting effects of forestry and climate change on the demography of a group-living bird population.

Anthropogenic degradation of natural habitats is a global driver of wildlife population declines. Local population responses to such environmental perturbations are generally well understood, but in socially structured populations, interactions between environmental and social factors may influence population responses. Thus, understanding how habitat degradation affects the dynamics of these populations requires simultaneous consideration of social and environmental mechanisms underlying demographic responses. Here we investigated the effect of habitat degradation through commercial forestry on spatiotemporal dynamics of a group-living bird, the Siberian jay, Perisoreus infaustus, in boreal forests of northern Sweden. We assessed the interacting effects of forestry, climate and population density on stage-specific, seasonal life-history rates and population dynamics, using long-term, individual-based demographic data from 70 territories in natural and managed forests. Stage-specific survival and reproductive rates, and consequently population growth, were lower in managed forests than in natural forests. Population growth was most sensitive to breeder survival and was more sensitive to early dispersing juveniles than those delaying dispersal. Forestry decreased population growth in managed forests by reducing reproductive success and breeder survival. Increased snow depth improved winter survival, and warmer spring temperatures enhanced reproductive success, particularly in natural forests. Population growth was stable in natural forests but it was declining in managed forests, and this difference accelerated under forecasted climate scenarios. Thus, climatic change could exacerbate the rate of forestry-induced population decline through reduced snow cover in our study species, and in other species with similar life-history characteristics and habitat requirements.