Responses to novelty in wild insular birds: comparing breeding populations in ecologically contrasting habitats

Islands have always provided ideal natural laboratories for assessing ecological parameters influencing behaviour. One hypothesis that lends itself well to testing in island habitats suggests that animals frequenting highly variable environments should be motivated to approach and interact with (i.e. explore) novelty. Intra-species comparisons of populations living in ecologically different island habitats may, thus, help reveal the factors that modulate animals’ responses to novelty. In this study, we presented novel objects to two geographically isolated breeding populations of the black-faced sheathbill (Chionis minor), a sedentary land-based bird that frequents remote sub-Antarctic islands. In the first population (Chionis minor ssp. crozettensis), the “Crozet group” (Baie du Marin, Ile de la Possession, Crozet Islands), breeding pairs inhabit a variable habitat close to penguin (Aptenodytes patagonicus) colonies. In the second population (Chionis minor ssp. minor), the “Kerguelen group” (île Verte, Morbihan gulf, Kerguelen Islands) breeding pairs live in penguin-free territories. In this latter population, the environment is less variable due to the presence of a broad intertidal zone which ensures year-round food availability. At both Kerguelen and Crozet, at least one breeding partner in all pairs approached at least one of the novel objects, and we found no significant differences in the latency of approach between the two populations. However, sheathbills at Crozet touched objects significantly more than birds at Kerguelen, and were also faster to touch them. We discuss how environmental variability, along with other potential influencing factors, may favour exploration of novelty in this wild insular bird

Cognition physique chez l’oiseau : général ou adapté ?

Physical cognition is defined as the knowledge that we possess about the inanimate objects surrounding us, and their relation with the environment. According to the general social intelligence hypothesis, physical cognition would have developed as a continuum (like all the other cognitive domains), in species living in complex social groups. By contrast, the adapted social intelligence hypothesis assumes that interacting with conspecifics has allowed development of superior, but specific, cognitive capacities related to the social realm. Although research on the evolution of cognition first focused on primates, we now know that some avian species are capable of interacting with their physical world in a complex way by using and manufacturing tools (e.g., Article 1). However, to date, various bird families are still unstudied, leaving open this debate. The general goal of this work is to contribute to a better understanding of the factors acting on the evolution of cognition, thanks to the theoretical input of the social intelligence hypothesis according to its general and adapted aspect. Four species that had never been studied in cognitive studies before were selected according to their degree of sociality but also according to the specificity of their ecology. This latter factor has been largely ignored in behavioural biology, although it is crucial for a more holistic comprehension of the behaviour. This work aimed to assess the ability of two avian species, great white pelicans Pelecanus onocrotalus (social birds) and yellow-crowned bishops Euplectes afer afer (gregarious birds), to imitate the behaviour of a trained conspecific (social domain) with a social learning task (Articles 2 part I & 3, respectively). Subsequently, we have studied heir ability to use an object in order to get an out-of-reach food reward (physical domain) with a tool use task (Articles 2 part II & 4). Although pelicans were capable of rapidly solving the social learning task, they did not succeed in using tools spontaneously (cf. discussion Article 2 part II). The bishops were able to imitate the behaviour of a conspecific, however they were not capable of using tools in the foraging context, although these birds are well known to use and manufacture tools in quite a complex way in order to build their nests. We have also administrated to two other avian social species, ground-hornbills Bucorvus and toucans Ramphastidae, an experimental paradigm to assess cognition in the physical domain: the string-pulling test (Articles 5 & 6, respectively). The task involved pulling on a string in order to obtain a food reward attached to its extremity. Ground-hornbills failed to pull on the string in the vertical configuration, but rapidly solved the task in various conditions within the horizontal configuration. In toucans, however, only one subject succeeded in the vertical configuration. These results allow us to refute the social intelligence hypothesis in its general aspect. Indeed, no link seems to be drawn between the social and the physical domains in the four species studied. Although the social intelligence hypothesis is supported since decades, a unitary theory is required. A new model of cognitive evolution, that allows assessing the importance of general intelligence in species, may be particularly promising.La cognition physique correspond à l’ensemble des connaissances que nous possédons sur les objets inanimés qui nous entourent, et à leurs relations avec l’environnement. Selon l’hypothèse de l’intelligence sociale générale, la cognition physique se serait développée tel un continuum (à l’instar de tous les autres domaines cognitifs), chez les espèces vivant au sein de groupes sociaux complexes. A l’inverse, l’hypothèse de l’intelligence sociale adaptée suppose que le fait d’interagir avec des congénères a permis de développer des capacités cognitives supérieures, mais spécifiques, du domaine social. Bien que les recherches relatives à l’évolution de la cognition physique se soient d’abord focalisées sur les primates, nous savons aujourd’hui que certains oiseaux sont capables d’interagir de manière complexe avec leur monde physique, en utilisant et en fabriquant des outils (p. ex., voir Article 1). Néanmoins, de nombreuses familles aviaires restent à ce jour non étudiées, laissant ce débat en suspens. L’objectif général de ce travail de recherche est de contribuer à une meilleure compréhension des facteurs responsables de l’évolution de la cognition, grâce à l’apport théorique de l’hypothèse de l’intelligence sociale suivant son aspect général et adapté. Quatre espèces, jusqu’alors inconnues sur le plan cognitif, ont été sélectionnées suivant leur degré de socialité mais également la spécificité de leur écologie. Ce dernier facteur, trop souvent ignoré en biologie du comportement, est crucial pour apprécier le comportement dans son ensemble. Nous avons ainsi estimé la faculté des sujets, à savoir des pélicans blancs Pelecanus onocrotalus (sociaux) et des euplectes vorabés Euplectes afer afer (grégaires), à reproduire le comportement d’un congénère (domaine social) grâce à un test d’apprentissage social (Articles 2 part I & 3). Nous avons ensuite évalué leur capacité à se servir d’un objet pour obtenir une récompense alimentaire hors de portée (domaine physique), grâce à un test d’utilisation d’outils (Articles 2 part II & 4). Bien que les pélicans fussent capables de résoudre rapidement la tâche d’apprentissage social, ils ne réussirent pas à utiliser spontanément des outils (cf. discussion Article 2 part II). A l’instar du pélican, l’euplecte imita le comportement d’un congénère. Néanmoins, il échoua à utiliser des outils dans le contexte du fourragement, malgré le fait que cet oiseau utilise et fabrique des outils de manière complexe pour construire son nid.A défaut d’avoir pu étudier l’apprentissage social et l’utilisation d’outils chez deux autres espèces sociales, les calaos terrestres Bucorvus et les toucans Ramphastidae, cette recherche consistait également à administrer un paradigme permettant d’apprécier le domaine physique : le test de la ficelle (Articles 5 & 6, respectivement). La tâche impliquait de tirer sur une ficelle afin d’obtenir une récompense alimentaire accrochée à son extrémité. Les calaos terrestres échouèrent à tirer sur la ficelle dans la configuration verticale, mais réussirent rapidement la tâche dans diverses conditions de la configuration horizontale. Chez les toucans, cependant, un seul sujet réussit le test dans sa configuration verticale. Les résultats obtenus nous permettent de réfuter l’hypothèse de l’intelligence sociale dans son aspect général. En effet, aucun lien ne semble se dessiner entre le domaine social et physique chez les quatre espèces aviaires étudiées. Bien que l’hypothèse de l’intelligence sociale soit soutenue depuis plus de quatre décennies, une théorie unitaire est requise. Dans ce cadre, un nouveau modèle d’évolution cognitive, permettant d’évaluer l’importance de l’intelligence générale chez une espèce donnée, pourrait s’avérer particulièrement prometteur

Physical cognition in birds : general or adapted?

La cognition physique correspond à l’ensemble des connaissances que nous possédons sur les objets inanimés qui nous entourent, et à leurs relations avec l’environnement. Selon l’hypothèse de l’intelligence sociale générale, la cognition physique se serait développée tel un continuum (à l’instar de tous les autres domaines cognitifs), chez les espèces vivant au sein de groupes sociaux complexes. A l’inverse, l’hypothèse de l’intelligence sociale adaptée suppose que le fait d’interagir avec des congénères a permis de développer des capacités cognitives supérieures, mais spécifiques, du domaine social. Bien que les recherches relatives à l’évolution de la cognition physique se soient d’abord focalisées sur les primates, nous savons aujourd’hui que certains oiseaux sont capables d’interagir de manière complexe avec leur monde physique, en utilisant et en fabriquant des outils (p. ex., voir Article 1). Néanmoins, de nombreuses familles aviaires restent à ce jour non étudiées, laissant ce débat en suspens. L’objectif général de ce travail de recherche est de contribuer à une meilleure compréhension des facteurs responsables de l’évolution de la cognition, grâce à l’apport théorique de l’hypothèse de l’intelligence sociale suivant son aspect général et adapté. Quatre espèces, jusqu’alors inconnues sur le plan cognitif, ont été sélectionnées suivant leur degré de socialité mais également la spécificité de leur écologie. Ce dernier facteur, trop souvent ignoré en biologie du comportement, est crucial pour apprécier le comportement dans son ensemble. Nous avons ainsi estimé la faculté des sujets, à savoir des pélicans blancs Pelecanus onocrotalus (sociaux) et des euplectes vorabés Euplectes afer afer (grégaires), à reproduire le comportement d’un congénère (domaine social) grâce à un test d’apprentissage social (Articles 2 part I & 3). Nous avons ensuite évalué leur capacité à se servir d’un objet pour obtenir une récompense alimentaire hors de portée (domaine physique), grâce à un test d’utilisation d’outils (Articles 2 part II & 4). Bien que les pélicans fussent capables de résoudre rapidement la tâche d’apprentissage social, ils ne réussirent pas à utiliser spontanément des outils (cf. discussion Article 2 part II). A l’instar du pélican, l’euplecte imita le comportement d’un congénère. Néanmoins, il échoua à utiliser des outils dans le contexte du fourragement, malgré le fait que cet oiseau utilise et fabrique des outils de manière complexe pour construire son nid.A défaut d’avoir pu étudier l’apprentissage social et l’utilisation d’outils chez deux autres espèces sociales, les calaos terrestres Bucorvus et les toucans Ramphastidae, cette recherche consistait également à administrer un paradigme permettant d’apprécier le domaine physique : le test de la ficelle (Articles 5 & 6, respectivement). La tâche impliquait de tirer sur une ficelle afin d’obtenir une récompense alimentaire accrochée à son extrémité. Les calaos terrestres échouèrent à tirer sur la ficelle dans la configuration verticale, mais réussirent rapidement la tâche dans diverses conditions de la configuration horizontale. Chez les toucans, cependant, un seul sujet réussit le test dans sa configuration verticale. Les résultats obtenus nous permettent de réfuter l’hypothèse de l’intelligence sociale dans son aspect général. En effet, aucun lien ne semble se dessiner entre le domaine social et physique chez les quatre espèces aviaires étudiées. Bien que l’hypothèse de l’intelligence sociale soit soutenue depuis plus de quatre décennies, une théorie unitaire est requise. Dans ce cadre, un nouveau modèle d’évolution cognitive, permettant d’évaluer l’importance de l’intelligence générale chez une espèce donnée, pourrait s’avérer particulièrement prometteur.Physical cognition is defined as the knowledge that we possess about the inanimate objects surrounding us, and their relation with the environment. According to the general social intelligence hypothesis, physical cognition would have developed as a continuum (like all the other cognitive domains), in species living in complex social groups. By contrast, the adapted social intelligence hypothesis assumes that interacting with conspecifics has allowed development of superior, but specific, cognitive capacities related to the social realm. Although research on the evolution of cognition first focused on primates, we now know that some avian species are capable of interacting with their physical world in a complex way by using and manufacturing tools (e.g., Article 1). However, to date, various bird families are still unstudied, leaving open this debate. The general goal of this work is to contribute to a better understanding of the factors acting on the evolution of cognition, thanks to the theoretical input of the social intelligence hypothesis according to its general and adapted aspect. Four species that had never been studied in cognitive studies before were selected according to their degree of sociality but also according to the specificity of their ecology. This latter factor has been largely ignored in behavioural biology, although it is crucial for a more holistic comprehension of the behaviour. This work aimed to assess the ability of two avian species, great white pelicans Pelecanus onocrotalus (social birds) and yellow-crowned bishops Euplectes afer afer (gregarious birds), to imitate the behaviour of a trained conspecific (social domain) with a social learning task (Articles 2 part I & 3, respectively). Subsequently, we have studied heir ability to use an object in order to get an out-of-reach food reward (physical domain) with a tool use task (Articles 2 part II & 4). Although pelicans were capable of rapidly solving the social learning task, they did not succeed in using tools spontaneously (cf. discussion Article 2 part II). The bishops were able to imitate the behaviour of a conspecific, however they were not capable of using tools in the foraging context, although these birds are well known to use and manufacture tools in quite a complex way in order to build their nests. We have also administrated to two other avian social species, ground-hornbills Bucorvus and toucans Ramphastidae, an experimental paradigm to assess cognition in the physical domain: the string-pulling test (Articles 5 & 6, respectively). The task involved pulling on a string in order to obtain a food reward attached to its extremity. Ground-hornbills failed to pull on the string in the vertical configuration, but rapidly solved the task in various conditions within the horizontal configuration. In toucans, however, only one subject succeeded in the vertical configuration. These results allow us to refute the social intelligence hypothesis in its general aspect. Indeed, no link seems to be drawn between the social and the physical domains in the four species studied. Although the social intelligence hypothesis is supported since decades, a unitary theory is required. A new model of cognitive evolution, that allows assessing the importance of general intelligence in species, may be particularly promising

Exclusion by donkey’s ears: Donkeys (Equus asinus) use acoustic information to find hidden food in a two-way object-choice task.

International audienceOnce believed uniquely human, the capacity to reason is now investigated in a wide range of species. One component of this ability, inference by exclusion, has been traditionally explored through the cups task, where 2 containers are presented but only 1 covers a food reward (if Cup A is empty, then choose Cup B). Often based on low-level cognitive mechanisms (learning), performance on this task can also reflect reasoning processes in some individuals. Limited taxonomic investigation, however, has hindered our understanding of the factors that contribute to the evolution of reasoning in animals. Using the cups task, we first investigated the ability of 14 semiwild donkeys to locate a food reward by using visual or acoustic cues alone (pretest phase). Although all subjects failed the task when provided with visual cues, 7 donkeys succeeded in the acoustic modality. We thus tested the capacity of donkeys to make choices by exclusion in the acoustic modality (test phase). Three conditions were presented, where subjects received information about either both containers (full information condition), solely the empty container (exclusion condition), or no information (control condition). Most subjects chose the correct container in the full information condition, whereas they chose randomly in the control condition. Except for 1 individual, donkeys improved their performance across trials in the exclusion condition. Our study contributes to the very small body of literature on exclusion based on acoustic cues in animals and paves the way to further experiments on the cognitive processes underlying exclusion performance in donkeys. (PsycInfo Database Record (c) 2022 APA, all rights reserved)

Wild skuas can follow human-given behavioural cues when objects resemble natural food

International audienceThe capacity to follow human cues provides animals with information about the environment and can hence offer obvious adaptive benefits. Most studies carried out so far, however, have been on captive animals with previous experience with humans. Further comparative investigation is needed in order to properly assess the factors driving the emergence of this capacity under natural conditions, especially in species that do not have longstanding interactions with humans. Wild brown skuas (Catharacta antarctica ssp. lonnbergi) are non-neophobic seabirds that live in human-free habitats. In test 1: food objects, we assessed this species’ capacity to use human behavioural cues (i.e., pecking at the same object previously picked up and lifted by a human experimenter) when the items presented were food objects: anthropogenic objects (wrapped muffins) and resembling-natural food objects (plaster eggs). In test 2: non-food objects, we tested the response of another skua population towards non-food objects (sponges). Although all skuas in test 1: food objects pecked at an object, they pecked significantly more at the same previously handled items when they resembled natural food (plaster eggs). Most skuas in test 2: non-food objects, however, did not approach or peck the non-food objects presented. Our results lead us to suggest that the use of human-behavioural cues may be influenced by skuas’ foraging ecology, which paves the way to further field studies assessing whether this capacity develops specifically towards food objects or after previous interaction with human

Wild skuas can use acoustic cues to locate hidden food

International audienceAmong animals, the visual acuity of several predatory bird species is probably the most outstanding. This, and the ease with which visually based tasks are administered, has led researchers to predominantly use the visual modality when studying avian cognition. Some wild skua populations routinely use acoustic cues emitted by their prey during foraging. In this study, we thus assessed whether this species was able to locate hidden food using acoustic cues alone (training phase). During the subsequent test phase, we investigated the capacity of successful individuals to choose the correct baited container in four conditions: (i) baited (shaking the baited container), (ii) full information (shaking both containers), (iii) exclusion (shaking the empty container), and (iv) control (shaking neither container). Four out of ten subjects succeeded in locating the baited container in the training phase. During the test phase, most subjects chose the baited container significantly more than the empty container in the baited and full information condition, while their performance was at chance level in the control condition. When no sound emanated from the empty container in the exclusion condition, one out of four skuas chose the baited container with more accuracy than predicted by chance. As this bird chose correctly on the first trial and during the first five trials, its performance is unlikely due to learning processes (learning to exclude the empty container). Although further tests are necessary to draw firm conclusions, our results open the way for assessing further this species’ reasoning abilities in the wild

Exclusion in the field: wild brown skuas find hidden food in the absence of visual information

International audienceInferential reasoning by exclusion allows responding adaptively to various environmental stimuli when confronted with inconsistent or partial information. In the experimental context, this mechanism allows selecting correctly between an empty option and a potentially rewarded one. Recently, the increasing reports of this capacity in phylogenetically distant species have led to the assumption that reasoning by exclusion is the result of convergent evolution. Within one largely unstudied avian order, i.e. the Charadriiformes, brown skuas (Catharacta antarctica ssp lonnbergi) are highly flexible and opportunistic predators. Behavioural flexibility, along with specific aspects of skuas' feeding ecology, may act as influencing factors in their ability to show exclusion performance. Our study aims to test whether skuas are able to make choice by exclusion in a visual two-way object-choice task. Twenty-six wild birds were presented with two opaque cups, one covering a food reward. Three conditions were used: 'full information' (showing the content of both cups), 'exclusion' (showing the content of the empty cup), and 'control' (not showing any content). Skuas preferentially selected the rewarded cup in the full information and exclusion condition. The use of olfactory cues was excluded by results in the control condition. Our study highlights the cognitive potential of this predatory seabird and opens new investigations for testing further its cognition in the wild

Exclusion in the field: wild brown skuas find hidden food in the absence of visual information

International audienceInferential reasoning by exclusion allows responding adaptively to various environmental stimuli when confronted with inconsistent or partial information. In the experimental context, this mechanism allows selecting correctly between an empty option and a potentially rewarded one. Recently, the increasing reports of this capacity in phylogenetically distant species have led to the assumption that reasoning by exclusion is the result of convergent evolution. Within one largely unstudied avian order, i.e. the Charadriiformes, brown skuas (Catharacta antarctica ssp lonnbergi) are highly flexible and opportunistic predators. Behavioural flexibility, along with specific aspects of skuas' feeding ecology, may act as influencing factors in their ability to show exclusion performance. Our study aims to test whether skuas are able to make choice by exclusion in a visual two-way object-choice task. Twenty-six wild birds were presented with two opaque cups, one covering a food reward. Three conditions were used: 'full information' (showing the content of both cups), 'exclusion' (showing the content of the empty cup), and 'control' (not showing any content). Skuas preferentially selected the rewarded cup in the full information and exclusion condition. The use of olfactory cues was excluded by results in the control condition. Our study highlights the cognitive potential of this predatory seabird and opens new investigations for testing further its cognition in the wild

First description of nest-decoration behaviour in a wild sub-Antarctic shorebird

International audienceA wide range of animal species accumulate objects in, on, and/or around structures they build. Sometimes, these accumulations serve specific functions (e.g. structural or isolating features) or are purely incidental, while in other cases the materials are deliberately displayed to serve signalling purposes (extended phenotype signals). In this pilot study, we employed systematic in situ observations and camera trapping to describe for the first time that both partners of a territorial shorebird, the black-faced sheathbill (Chionis minor ssp minor) collect, carry, and arrange colourful marine shells and dry twigs within and around their nest cavity. Our observations expand the taxonomic breadth of avian extended phenotype signals, by showing that at least one species within a largely understudied group i.e., Charadriiformes, exhibits nest-decoration behaviour. Multiple manipulative experiments are needed to explore further the signalling function of these decorations, which opens new exciting avenues for animal communication and cognition research

Responses of wild skuas (Catharacta antarctica ssp. lonnbergi) to human cues in cooperative and competitive social contexts.

Many animals respond to and use social cues emitted by other species (e.g., head direction). In the context of human–animal communication, these capacities have been attributed to regular and longstanding exposure to humans. We presented wild brown skuas (Catharacta antarctica ssp. lonnbergi) with two versions of an object-choice paradigm. In the cooperative version (Experiment 1), one human experimenter provided a simple and salient cue indicating which of two containers covered a food reward. The cues administered consisted of touching, looking at, pointing at, or pointing and looking at the container hiding food. In Experiment 1, skuas could thus cooperate with an experimenter by using the cues provided to locate the rewarded container. In the competitive version (Experiment 2), two human experimenters presented a platform with a visible food reward. In six experimental conditions, we varied experimenters’ body orientation, head orientation, eye-gaze direction, face occlusion, and mouth occlusion, as well as the platform’s location, ensuring that in each case only one experimenter had visual access to the rewarded platform. Here, birds could compete with the experimenters by robbing the human who does not see the food. Skuas failed to use human-given cues spontaneously in Experiment 1, and took the reward regardless of whether the experimenters could see in Experiment 2. Our results contrast with those obtained on other wild birds with pre-experience with humans. Hopefully, our findings will stimulate further research in order to illuminate the potential role of such experience in the capacity to respond to and use human-given cues