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The best-known example of episodic memory in animals came from food-storing birds. One of the beauties of the food-storing system was that inherent in the behaviour were the elements that (at the time) made up episodic memory: what, where and when. While there were then already plenty of data on animals' ability to put together what and where, the addition of the time element in animals' memory and its testing was one that was both new and experimentally challenging. It has, however, led to an increasing variety of examples showing that animals can put together all three informational components. If episodic memories can be described as those memories that make any one of us who we are, why should non-human animals have such memories? Here, we argue that episodic memories play a significant functional role in the lives of real animals, in particular, enabling them to make decisions about how they might or should act in their future. We support our argument with data from a range of examples, focussing on data from the field.This article is part of the theme issue 'Elements of episodic memory: lessons from 40 years of research'.

Humans cooperate to build complex structures with culture-specific architectural styles. However, they are not the only animals to build complex structures nor to have culture. We show that social groups of white-browed sparrow weavers ( Plocepasser mahali ) build structures (nests for breeding and multiple single-occupant roosts for sleeping) that differ architecturally among groups. Morphological differences are consistent across years and are clear even among groups with territories a few meters apart. These repeatable differences are not explained by among-group variation in local weather conditions, bird size, tree height, or patterns of genetic relatedness. Architectural styles are also robust to the immigration of birds from other groups.

As humans increasingly modify the natural world, many animals have responded by changing their behaviour. Understanding and predicting the extent of these responses is a key step in conserving these species. For example, the tendency for some species of birds to incorporate anthropogenic items-particularly plastic material-into their nests is of increasing concern, as in some cases, this behaviour has harmful effects on adults, young and eggs. Studies of this phenomenon, however, have to date been largely limited in geographic and taxonomic scope. To investigate the global correlates of anthropogenic (including plastic) nest material use, we used Bayesian phylogenetic mixed models and a data set of recorded nest materials in 6147 species of birds. We find that, after controlling for research effort and proximity to human landscape modifications, anthropogenic nest material use is correlated with synanthropic (artificial) nesting locations, breeding environment and the number of different nest materials the species has been recorded to use. We also demonstrate that body mass, range size, conservation status and brain size do not explain variation in the recorded use of anthropogenic nest materials. These results indicate that anthropogenic materials are more likely to be included in nests when they are more readily available, as well as potentially by species that are more flexible in their nest material choice., (© 2024 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Brood (social) parasites and their hosts exhibit a wide range of adaptations and counter-adaptations as part of their ongoing coevolutionary arms races. Obligate avian brood parasites are expected to use potential host species with more easily accessible nests, while potential hosts are expected to evade parasitism by building more concealed nests that are difficult for parasites to enter and in which to lay eggs. We used phylogenetically informed comparative analyses, a global database of the world's brood parasites, their host species, and the design of avian host and non-host nests (approx. 6200 bird species) to examine first, whether parasites preferentially target host species that build open nests and, second, whether host species that build enclosed nests are more likely to be targeted by specialist parasites. We found that species building more accessible nests are more likely to serve as hosts, while host species with some of the more inaccessible nests are targeted by more specialist brood parasites. Furthermore, evolutionary-transition analyses demonstrate that host species building enclosed nests frequently evolve to become non-hosts. We conclude that nest architecture and the accessibility of nests for parasitism represent a critical stage of the ongoing coevolutionary arms race between avian brood parasites and their hosts.

The evolution of behaviour can both influence, and be influenced by, morphology. Recent advances in methods and data availability have facilitated broad-scale investigations of physical form and behavioural function in many contexts, but the relationship between animal morphology and object manipulation-particularly objects used in construction-remains largely unknown. Here, we employ a new global database of nest materials used by 5924 species of birds together with phylogenetically informed random forest models to evaluate the link between beak shape and these nest-building materials. We find that beak morphology, together with species diet and access to materials, can predict nest-material use above chance and with high accuracy (68-97%). Much of this relationship, however, is driven by phylogenetic signal and sampling biases. We therefore conclude that while variation in nest material use is linked with that of beak shape across bird species, these correlations are modulated by the ecological context and evolutionary history of these species. This article is part of the theme issue 'The evolutionary ecology of nests: a cross-taxon approach'.

Successful reproduction for most birds requires them to have built 'good' nests. The remarkable diversity of nests across approximately 10 000 species of living birds suggests that 'good' nest design depends critically on a species' microhabitat, life history and behaviour. Unravelling the key drivers of nest diversity remains a key research priority-bolstered by renewed appreciation for nest museum collections and increasing correlational field and experimental laboratory data. Phylogenetic analyses-coupled with powerful datasets of nest traits-are increasingly shedding light on the evolution of nest morphology and there are functional questions yet to be addressed. For birds, at least, developmental and mechanistic analyses of building (behaviour, hormones, neuroscience) itself, rather than measurements and analyses of nest morphology, are already becoming the next major challenge. We are moving towards a holistic picture in which Tinbergen's four levels of explanation: evolution, function, development, and mechanism, are being used to explain variation and convergence in nest design-and, in turn, could shed light on the question of how birds know how to build 'good' nests. This article is part of the theme issue 'The evolutionary ecology of nests: a cross-taxon approach'.

The size of a bird's nest can play a key role in ensuring reproductive success and is determined by a variety of factors. The primary function of the nest is to protect offspring from the environment and predators. Field studies in a number of passerine species have indicated that higher-latitude populations in colder habitats build larger nests with thicker walls compared to lower-latitude populations, but that these larger nests are more vulnerable to predation. Increases in nest size can also be driven by sexual selection, as nest size can act as a signal of parental quality and prompt differential investment in other aspects of care. It is unknown, however, how these microevolutionary patterns translate to a macroevolutionary scale. Here, we investigate potential drivers of variation in the outer and inner volume of open cup nests using a large dataset of nest measurements from 1117 species of passerines breeding in a diverse range of environments. Our dataset is sourced primarily from the nest specimens at the Natural History Museum (UK), complemented with information from ornithological handbooks and online databases. We use phylogenetic comparative methods to test long-standing hypotheses about potential macroevolutionary correlates of nest size, namely nest location, clutch size and variables relating to parental care, together with environmental and geographical factors such as temperature, rainfall, latitude and insularity. After controlling for phylogeny and parental body size, we demonstrate that the outer volume of the nest is greater in colder climates, in island-dwelling species and in species that nest on cliffs or rocks. By contrast, the inner cup volume is associated solely with average clutch size, increasing with the number of chicks raised in the nest. We do not find evidence that nest size is related to the length of parental care for nestlings. Our study reveals that the average temperature in the breeding range, along with several key life-history traits and proxies of predation threat, shapes the global interspecific variation in passerine cup nest size. We also showcase the utility of museum nest collections-a historically underused resource-for large-scale studies of trait evolution., (© 2022 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Typically, investigations of animal cognition couple careful experimental manipulations with examination of the animal's behavioural responses. Sometimes those questions have included attempts to describe the neural underpinnings of the behavioural outputs. Over the past 25 years, behaviours that involve spatial learning and memory (such as navigation and food storing) has been one context in which such dual or correlated investigations have been both accessible and productive. Here I review some of that work and where it has led. Because of the wealth of data and insights gained from that work and song learning before it, it seems that it might also be useful to try to add some neurobiology to other systems in animal cognition. I finish then, with a description of recent work on the cognition and neurobiology of avian nest building. It is still relatively early days but asking questions about the cognition of nest building has already shown both neural correlates of nest building and that learning and memory play a much greater role in this behaviour than previously considered. While it is not yet clear how putting these components together will be synergistic, the examples of song learning and food storing provide encouragement. Perhaps this might be true for other behaviours too?, (© 2022. The Author(s).)

Bird nests have long attracted human interest, both as beautiful creations and as important constructions. They are important constructions because avian parents use them to protect their developing eggs and young from the dangers presented by the elements and predators. Despite sharing these protective functions, however, bird nests can be remarkably structurally variable, being built from a wide diversity of materials, and placed in all manner of locations. For example, male Australian brush turkeys scrape material into large nest mounds, woodpeckers excavate holes, storks build stacks, sparrows and swallows build cups (but of different materials), while weavers weave their famous hanging baskets. Nests might be built in bushes or holes in trees, dangle from a branch, or sit on the ground. They might be solitary and hidden, or conspicuous, together housing hundreds of families (Figure 1)., Competing Interests: Declaration of interests The author declares no competing interests., (Copyright © 2022 Elsevier Inc. All rights reserved.)

Rufous hummingbirds ( Selasphorus rufus ) will readily learn the location and the colour of rewarded flowers within their territory. But if these birds could apply a relational concept such as 'the larger flowers have more nectar', they could forego learning the locations of hundreds of individual flowers. Here, we investigated whether wild male territorial rufous hummingbirds might use 'larger than' and 'smaller than' relational rules and apply them to flowers of different sizes. Subjects were trained to feed consistently from one of two flowers. Although the flowers differed only in size, the reward was always contained in the same-size flower. The birds were then tested on a choice of two empty flowers: one of the familiar size and the other a novel size. Hummingbirds applied relational rules by choosing the flower that was of the correct relational size rather than visiting the flower of the size rewarded during training. The choices made by the hummingbirds were not consistent with alternative mechanisms such as peak shift or associative learning. We suggest that while hummingbirds are very good at remembering the spatial locations of rewarding flowers, they would be able to use relative rules when foraging in new and changing environments.

Traplining, when animals repeat the order in which they visit a number of locations, is taxonomically widespread, but little is known about which factors influence the routes that animals follow. For example, as the quality of rewarding locations changes over time, foragers are expected to update their traplines, either to prioritize locations where the reward increases or to avoid locations that have ceased to be profitable. Here, we tested how traplining wild hummingbirds responded to increases or to decreases in the sucrose concentration of one of the flowers on their trapline. Hummingbirds did not change their trapline to visit the flower with the increased reward first, but by changing the order in which they visited flowers, they avoided a flower that contained a decreased reward. Depending on where along the trapline the reduced-content flower occurred, hummingbirds either changed the origin of their trapline or changed the direction in which they flew around their trapline. It may be that this asymmetric modification of foraging traplines is especially noticeable in risk-averse foragers, such as these territorial hummingbirds., (© 2021. The Author(s).)

Female zebra finches Taeniopygia guttata will copy the novel foraging choice of males. The degree to which they do so, however, can vary considerably. Among-individual differences in social learning and their underlying neural pathways have received relatively little attention and remain poorly understood. Here, then, we allowed female zebra finches to observe live-streamed male demonstrators feeding from one of two novel-coloured feeders (social information acquisition phase). After this social information acquisition phase, we tested from which feeder the females preferred to feed to determine whether they copied the feeder choice of the male demonstrator (social learning test phase). We then examined the brains of these females for immediate early gene activity (c-fos) in the neural social behaviour network for the time during which they were observing the male feeding. Of the 12 regions and sub-regions in the brain examined we found only one weak correlation: greater copying was associated with lower activity in the bed nucleus of the stria terminalis, BSTmv. Future work should perhaps focus on neural activity that occurs during the stage in which there is evidence that animals have copied a demonstrator (i.e., social learning test phase in the current experiment) rather than during the period in which those animals observe that demonstrator (i.e., social information acquisition phase in the current experiment). What is clear is that the considerable emphasis on examining the behavioural component of social learning has not yet been accompanied by neural analyses., (© 2022. The Psychonomic Society, Inc.)

But fish cognitive ecology did not begin in rivers and streams. Rather, one of the starting points for work on fish cognitive ecology was work done on the use of visual cues by homing pigeons. Prior to working with fish, Victoria Braithwaite helped to establish that homing pigeons rely not just on magnetic and olfactory cues but also on visual cues for successful return to their home loft. Simple, elegant experiments on homing established Victoria's ability to develop experimental manipulations to examine the role of visual cues in navigation by fish in familiar areas. This work formed the basis of a rich seam of work whereby a fish's ecology was used to propose hypotheses and predictions as to preferred cue use, and then cognitive abilities in a variety of fish species, from model systems (Atlantic salmon and sticklebacks) to the Panamanian Brachyraphis episcopi . Cognitive ecology in fish led to substantial work on fish pain and welfare, but was never left behind, with some of Victoria's last work addressed to determining the neural instantiation of cognitive variation., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Healy and Patton.)

Birds' nests represent a rich behavioural 'fingerprint', comprising several important decisions-not the least of which is the selection of appropriate material. Material selection in nest-building birds is thought to reflect, in part, builder-birds' use of the 'best' material-in terms of physical properties (e.g., rigidity)-refined across generations. There is, however, little experimental evidence to link the physical properties of nest material to both birds' nest-building and breeding performance. We examined individual-level material-use consequences for breeding zebra finches by manipulating the kind of material available to laboratory-housed pairs: stiff or flexible same-length string. We show that higher fledgling numbers were related to: (i) fewer pieces used in nest construction by stiff-string builders; and conversely, (ii) more pieces used in nest construction by flexible-string builders. Together, these data suggest that physical differences in nest material can affect avian reproduction (here, the trade-off between nest-construction investment and fledgling success), highlighting the adaptive significance of nest-building birds' material selectivity., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Innovation-the combination of invention and social learning-can empower species to invade new niches via cultural adaptation. Social learning has typically been regarded as the fundamental driver for the emergence of traditions and thus culture. Consequently, invention has been relatively understudied outside the human lineage-despite being the source of new traditions. This neglect leaves basic questions unanswered: what factors promote the creation of new ideas and practices? What affects their spread or loss? We critically review the existing literature, focusing on four levels of investigation: traits (what sorts of behaviours are easiest to invent?), individuals (what factors make some individuals more likely to be inventors?), ecological contexts (what aspects of the environment make invention or transmission more likely?), and populations (what features of relationships and societies promote the rise and spread of new inventions?). We aim to inspire new research by highlighting theoretical and empirical gaps in the study of innovation, focusing primarily on inventions in non-humans. Understanding the role of invention and innovation in the history of life requires a well-developed theoretical framework (which embraces cognitive processes) and a taxonomically broad, cross-species dataset that explicitly investigates inventions and their transmission. We outline such an agenda here. This article is part of the theme issue 'Foundations of cultural evolution'.

Our current understanding of manipulation is based on primate hands, resulting in a detailed but narrow perspective of ways to handle objects. Although most other animals lack hands, they are still capable of flexible manipulation of diverse objects, including food and nest materials, and depend on dexterity in object handling to survive and reproduce. Birds, for instance, use their bills and feet to forage and build nests, while insects carry food and construct nests with their mandibles and legs. Bird bills and insect mandibles are much simpler than a primate hand, resembling simple robotic grippers. A better understanding of manipulation in these and other species would provide a broader comparative perspective on the origins of dexterity. Here we contrast data from primates, birds and insects, describing how they sense and grasp objects, and the neural architectures that control manipulation. Finally, we outline techniques for collecting comparable manipulation data from animals with diverse morphologies and describe the practical applications of studying manipulation in a wide range of species, including providing inspiration for novel designs of robotic manipulators.

When presented with resources that differ in quantity, many animals use a numerosity system to discriminate between them. One taxonomically widespread system is the approximate number system. This is a numerosity system that allows the rapid evaluation of the number of objects in a group and which is regulated by Weber's Law. Here we investigated whether wild, free-living rufous hummingbirds (Selasphorus rufus) possess an approximate number system. The hummingbirds were presented with two experiments. In the first we investigated whether hummingbirds spontaneously chose an array containing more flowers than an alternate array. In the second we asked whether the hummingbirds could learn to use numerosity as a cue to which of two arrays contained the better reward. The birds did not spontaneously prefer an array containing more flowers. After minimal training, however, they learned to choose the more numerous array and could differentiate between arrays of five and seven flowers. These data support the presence of an approximate number system in the rufous hummingbird. It seems plausible that having such a system would enable much more efficient foraging in this species.

Husbandry procedures and facility settings, such as low-frequency fire alarms, can produce noises in a laboratory environment that cause stress to animals used in research. However, most of the data demonstrating harmful effects that have, consequently, led to adaptations to management, have largely come from laboratory rodents with little known of the impacts on avian behavior and physiology. Here we examined whether exposure to a routine laboratory noise, a low-frequency fire alarm test, induced behavioral changes in laboratory zebra finches ( Taeniopygia guttata ). Twenty-four breeding pairs of zebra finches were randomly selected and exposed to the low-frequency fire alarm (sounding for 10-20 s) or no noise (control) on separate test days. All birds were filmed before and after the alarm sounded and on a control day (without the alarm). The zebra finches decreased their general activity and increased stationary and social behaviors after exposure to the alarm. Brief exposure to a low-frequency alarm disrupted the birds' behavior for at least 15 min. The induction of this behavioral stress response suggests that low-frequency sound alarms in laboratory facilities have the potential to compromise the welfare of laboratory birds., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Corbani, Martin and Healy.)

Decision makers can be described as economically rational (making consistent choices), or economically irrational (making choices that vary with the options available). As the extent to which animals can and do make rational versus irrational decisions remains unclear, we tested the decision-making strategies of female Nasonia vitripennis parasitic wasps in two behavioural contexts: oviposition and foraging. In our first experiment, to determine whether oviposition preferences changed depending on the options available, we presented females with a high and a medium-quality blow fly host to parasitize, and gave some females an additional low or very low quality 'decoy' host. Presence of decoy options did not affect females' oviposition choices, either in willingness to parasitize a host or the number of offspring laid. In our second experiment, we tested the effects of a low-quality decoy option on foraging preference for a high and a medium-quality sucrose concentration option. Here, presence of the low-quality decoy enhanced female preference for the high-quality option. Females therefore made economically rational decisions when ovipositing and economically irrational decisions when foraging. This difference in decision outcomes suggests that the cost/benefit ratio of making one type of decision over another may differ with the behavioural task., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Nest building consists of a series of motor actions, which are concomitant with activity in regions of the anterior motor pathway, the social behavior network, and the reward circuity in nest building adult male zebra finches (Taeniopygia guttata). It is not clear, however, whether this activity is due to nest building, collection, and/or manipulation of nest material. To identify which areas of the brain are specifically involved, we used immunohistochemistry to quantify the immediate early gene c-Fos in male zebra finches that were nest building (Building), birds given a nest box but could interact only with tied down nest material (Fixed), and birds that were not given a nest box or nest material (Control). We investigated the following brain regions: the anterior motor pathway (anterior ventral mesopallium [AMV], AN, anterior striatum [ASt]), areas of the social behavior network (bed nucleus of the stria terminalis, dorsomedial subdivision [BSTmd], lateral septum [LS]), the dopaminergic reward circuitry (ventral tegmental area), and the cerebellum. We found that there was greater Fos immunoreactivity expression in the BSTmd, LS, and AMV with increased material deposition; in LS, AMV ASt, and Folium VI with increased material carrying; in LS, AMV, and ASt with increased nest material tucking; and in LS and all folia (except Folium VIII) with increased tugging at tied down material. These data confirm a functional role for areas of the anterior motor pathway, social behavior network, and the cerebellum in nest material collection and manipulation by birds., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Ordinality is a numerical property that nectarivores may use to remember the specific order in which to visit a sequence of flowers, a foraging strategy also known as traplining. In this experiment, we tested whether wild, free-living rufous hummingbirds ( Selasphorus rufus ) could use ordinality to visit a rewarded flower. Birds were presented with a series of linear arrays of 10 artificial flowers; only one flower in each array was rewarded with sucrose solution. During training, birds learned to locate the correct flower independent of absolute spatial location. The birds' accuracy was independent of the rewarded ordinal position (1st, 2nd, 3rd or 4th), which suggests that they used an object-indexing mechanism of numerical processing, rather than a magnitude-based system. When distance cues between flowers were made irrelevant during test trials, birds could still locate the correct flower. The distribution of errors during both training and testing indicates that the birds may have used a so-called working up strategy to locate the correct ordinal position. These results provide the first demonstration of numerical ordinal abilities in a wild vertebrate and suggest that such abilities could be used during foraging in the wild.

The duration of the developmental period represents a fundamental axis of life-history variation, yet broad insights regarding the drivers of this diversity are currently lacking. Here, we test mechanistic and ecological explanations for the evolution of developmental duration using embryological data and information on incubation and fledging for 3096 avian species. Developmental phases associated primarily with growth are the longest and most variable, consistent with a role for allometric constraint in determining the duration of development. In addition, developmental durations retain a strong imprint of deep evolutionary history and body size differences among species explain less variation than previously thought. Finally, we reveal ecological correlates of developmental durations, including variables associated with the relative safety of the developmental environment and pressures of breeding phenology. Overall, our results provide broad-scale insight into the relative importance of mechanistic, ecological and evolutionary constraints in shaping the diversification of this key life-history trait.

One source of public information may be the enduring products of others' behaviour, such as discarded tools or vacated nests. Here, we examined whether observation of a nest affects the material captive zebra finch males prefer when they construct their first nest. It does: for first-time nest construction, males that viewed only an empty cage preferred the colour of material each initially favoured but those males that had observed a pre-built nest of material of their non-preferred colour lost their material-colour preference altogether. Additionally, half of the males that viewed a nest were tested in an environment (the laboratory) different to that in which they were reared (an outdoor aviary). We had expected the aviary-reared (versus laboratory-reared) males would be more uncertain, and thus more likely to select material for their first nest that matched in colour to the colour of the 'demonstrated' nest-but this was not the case. The aviary-reared males did, however, tend to touch first the demonstrated colour of material more than did the laboratory-reared males. Together these results show that both observation of a nest and a change in environment can influence the material choices of novice builders. For naïve animal builders, then, construction artefacts can be information resources for learning about potential construction material.

Stress exposure during prenatal and postnatal development can have persistent and often dysfunctional effects on several physiological systems, including immune function, affecting the ability to combat infection. The neuroimmune response is inextricably linked to the action of the hypothalamic-pituitary-adrenal (HPA) axis. Cytokines released from neuroimmune cells, including microglia, activate the HPA axis, while glucocorticoids in turn regulate cytokine release from microglia. Because of the close links between these two physiological systems, coupled with potential for persistent changes to HPA axis activity following developmental stress, components of the neuroimmune system could be targets for developmental programming. However, little is known of any programming effects of developmental stress on neuroimmune function. We investigated whether developmental stress exposure via elevated prenatal corticosterone (CORT) or postnatal unpredictable food availability had long-term effects on pro- ( IL-1β ) and anti-inflammatory ( IL-10 ) cytokine and microglia-dependent gene ( CSF1R ) expression within HPA axis tissues in a precocial bird, the Japanese quail ( Coturnix japonica ). Following postnatal stress, we observed increased IL-1β expression in the pituitary gland, reduced IL-10 expression in the amygdala and hypothalamus, and reduced CSF1R expression within the hypothalamus and pituitary gland. Postnatal stress disrupted the ratio of IL-1β : IL-10 expression within the hippocampus and hypothalamus. Prenatal stress only increased IL-1β expression in the pituitary gland. We found no evidence for interactive or cumulative effects across life stages on basal cytokine and glia expression in adulthood. We show that postnatal stress may have a larger impact than elevated prenatal CORT on basal immunity in HPA-axis-specific brain regions, with changes in cytokine homeostasis and microglia abundance. These results provide evidence for postnatal programming of a pro-inflammatory neuroimmune phenotype at the expense of reduced microglia, which could have implications for central nervous system health and subsequent neuroimmune responses., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Animals that feed from resources that are constant in space and that refill may benefit from repeating the order in which they visit locations. This is a behavior known as traplining, a spatial phenomenon. Hummingbirds, like other central-place foragers, use short traplines when moving between several rewarding sites. Here we investigated whether traplining hummingbirds also use relevant temporal information when choosing which flowers to visit. Wild rufous hummingbirds that were allowed to visit 3 artificial flower patches in which flowers were refilled 20 min after they had been depleted repeated the order in which they visited the 3 patches. Although they tended to visit the first 2 patches sooner than 20 min, they visited the third patch at approximately 20-min intervals. The time between visits to the patches increased across the experiment, suggesting that the birds learned to wait longer before visiting a patch. The birds appeared to couple the sequential pattern of a trapline with temporal regularity, to some degree. This suggests that there is a temporal component to the repeated spatial movements flown by foraging wild hummingbirds., (© 2018 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

As an increasing number of researchers investigate the cognitive abilities of an ever-wider range of animals, animal cognition is currently among the most exciting fields within animal behavior. Tinbergen would be proud: all four of his approaches are being pursued and we are learning much about how animals collect information and how they use that information to make decisions for their current and future states as well as what animals do not perceive or choose to ignore. Here I provide an overview of this productivity, alighting only briefly on any single example, to showcase the diversity of species, of approaches and the sheer mass of research effort currently under way. We are getting closer to understanding the minds of other animals and the evolution of cognition at an increasingly rapid rate., (© 2018 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

Determining the role that social learning plays in construction behaviors, such as nest building or tool manufacture, could be improved if more experimental control could be gained over the exact public information that is provided by the demonstrator, to the observing individual. Using video playback allows the experimenter to choose what information is provided but will only be useful in determining the role of social learning if observers attend to, and learn from, videos in a manner that is similar to live demonstration. The goal of the current experiment was to test whether live-streamed video presentations of nest building by zebra finches Taeniopygia guttata would lead observers to copy the material choice demonstrated to them. Here, males that had not previously built a nest were given an initial preference test between materials of 2 colors. Those observers then watched live-stream footage of a familiar demonstrator building a nest with material of the color that the observer did not prefer. After this experience, observers were given the chance to build a nest with materials of the 2 colors. Although two-thirds of the observer males preferred material of the demonstrated color after viewing the demonstrator build a nest with material of that color more than they had previously, their preference for the demonstrated material was not as strong as that of observers that had viewed live demonstrator builders in a previous experiment. Our results suggest researchers should proceed with caution before using video demonstration in tests of social learning., (© 2018 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

Carlson and colleagues introduce mobbing an anti-predator behaviour found in many animals., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Navigation is an essential skill for many animals, and understanding how animal use environmental information, particularly visual information, to navigate has a long history in both ethology and psychology. In birds, the dominant approach for investigating navigation at small-scales comes from comparative psychology, which emphasizes the cognitive representations underpinning spatial memory. The majority of this work is based in the laboratory and it is unclear whether this context itself affects the information that birds learn and use when they search for a location. Data from hummingbirds suggests that birds in the wild might use visual information in quite a different manner. To reconcile these differences, here we propose a new approach to avian navigation, inspired by the sensory-driven study of navigation in insects. Using methods devised for studying the navigation of insects, it is possible to quantify the visual information available to navigating birds, and then to determine how this information influences those birds' navigation decisions. Focusing on four areas that we consider characteristic of the insect navigation perspective, we discuss how this approach has shone light on the information insects use to navigate, and assess the prospects of taking a similar approach with birds. Although birds and insects differ in many ways, there is nothing in the insect-inspired approach of the kind we describe that means these methods need be restricted to insects. On the contrary, adopting such an approach could provide a fresh perspective on the well-studied question of how birds navigate through a variety of environments.

Hummingbirds feed from hundreds of flowers every day. The properties of these flowers provide these birds with a wealth of information about colour, space and time to guide how they forage. To understand how hummingbirds might use this information, researchers have adapted established laboratory paradigms for use in the field. In recent years, however, experimental inspiration has come less from other birds, and more from looking at other nectar-feeders, particularly honeybees and bumblebees, which have been models for foraging behaviour and cognition for over a century. In a world in which the cognitive abilities of bees regularly make the news, research on the influence of ecology and sensory systems on bee behaviour is leading to novel insights in hummingbird cognition. As methods designed to study insects in the laboratory are being applied to hummingbirds in the field, converging methods can help us identify and understand convergence in cognition, behaviour and ecology., (© 2017 The Author(s).)

The transmission of information from an experienced demonstrator to a naïve observer often depends on characteristics of the demonstrator, such as familiarity, success or dominance status. Whether or not the demonstrator pays attention to and/or interacts with the observer may also affect social information acquisition or use by the observer. Here we used a video-demonstrator paradigm first to test whether video demonstrators have the same effect as using live demonstrators in zebra finches, and second, to test the importance of visual and vocal interactions between the demonstrator and observer on social information use by the observer. We found that female zebra finches copied novel food choices of male demonstrators they saw via live-streaming video while they did not consistently copy from the demonstrators when they were seen in playbacks of the same videos. Although naive observers copied in the absence of vocalizations by the demonstrator, as they copied from playback of videos with the sound off, females did not copy where there was a mis-match between the visual information provided by the video and vocal information from a live male that was out of sight. Taken together these results suggest that video demonstration is a useful methodology for testing social information transfer, at least in a foraging context, but more importantly, that social information use varies according to the vocal interactions, or lack thereof, between the observer and the demonstrator., (Copyright © 2017 The Authors. Published by Elsevier B.V. All rights reserved.)

Animals use cues from their environment to orient in space and to navigate their surroundings. Geometry is a cue whose informational content may originate from the metric properties of a given environment, and its use has been demonstrated in the laboratory in nearly every species of animal tested. However, it is not clear whether geometric information, used by animals typically tested in small, rectangular boxes, is directly relevant to animals in their natural environment. Here we present the first data that confirm the use of geometric cues by a free-living animal in the wild. We trained rufous hummingbirds to visit a rectangular array of four artificial flowers, one of which was rewarded. In some trials a conspicuous landmark cued the reward. Following array translocation and rotation, we presented hummingbirds with three tests. When trained and tested with the landmark, or when trained and tested without it, hummingbirds failed to show geometric learning. However, when trained with a landmark but tested without it, hummingbirds produced the classic geometric response, showing that they had learned the geometric relationships (distance and direction) of several non-reward visual elements of the environment. While it remains that the use of geometry to relocate a reward may be an experimental artefact, its use is not confined to the laboratory., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Establishment in urbanized environments is associated with changes in physiology, behaviour, and problem-solving. We compared the speed of learning in urban and rural female common mynas, Acridotheres tristis, using a standard visual discrimination task followed by a reversal learning phase. We also examined how quickly each bird progressed through different stages of learning, including sampling and acquisition within both initial and reversal learning, and persistence following reversal. Based on their reliance on very different food resources, we expected urban mynas to learn and reversal learn more quickly but to sample new contingencies for proportionately longer before learning them. When quantified from first presentation to criterion achievement, urban mynas took more 20-trial blocks to learn the initial discrimination, as well as the reversed contingency, than rural mynas. More detailed analyses at the level of stage revealed that this was because urban mynas explored the novel cue-outcome contingencies for longer, and despite transitioning faster through subsequent acquisition, remained overall slower than rural females. Our findings draw attention to fine adjustments in learning strategies in response to urbanization and caution against interpreting the speed to learn a task as a reflection of cognitive ability.

Cognitive biases, which are defined as distortions in cognitive processes that are influenced by a background emotional state, can provide information about an individual's affective state. For instance, negative cognitive biases, where individuals assess ambiguous situations as unrewarding, are commonly found in humans suffering from anxiety disorders. Cognitive biases are also increasingly used as indicators of affective state in animals. As it is not clear whether female and male animals differ in performance on cognitive bias tasks, we used a spatial location task to examine cognitive bias in female and male adult Norway rats (Rattus norvegicus). We trained the rats to distinguish between reward and unrewarded locations, and then provided food pots at ambiguous, intermediate positions. We found that, during testing, females were slowest to approach the unrewarded location, while they approached ambiguous and rewarded locations similarly quickly. In contrast, the males approached all locations quickly. This sex difference is consistent with previous evidence that male rats are quicker than females to extinguish previously learned associations. Cognitive bias tasks could therefore be used to examine sex differences in learning strategies, as well as providing opportunities to test predictions about sex differences in welfare requirements., (Copyright © 2016 Elsevier B.V. All rights reserved.)

When making decisions between options, humans are expected to choose the option that returns the highest benefit. In practice, however, adding inferior alternatives to the choice set can alter these decisions. Here we investigated whether decisions over the facial features that people find healthy looking can also be affected by the context in which they see those faces. To do this we examined the effect of choice set on the perception of health of images of faces of light-skinned Caucasian females. We manipulated apparent facial health by changing yellowness of the skin: the healthy faces were moderately yellow and the less healthy faces were either much more yellow or much less yellow. In each experiment, two healthy faces were presented along with a third, less healthy face. When the third face was much more yellow, participants chose the more yellow of the two healthy faces more often as the most healthy. However, when the third face was the least yellow, participants chose the less yellow of the two healthy faces more often. A further experiment confirmed that this result is not due to a generalised preference for an intermediate option. These results extend our understanding of context-dependent decision-making in humans, and suggest that comparative evaluation may be a common feature across many different kinds of choices that humans have to make., (Copyright © 2016. Published by Elsevier B.V.)

It is becoming apparent that birds learn from their own experiences of nest building. What is not clear is whether birds can learn from watching conspecifics build. As social learning allows an animal to gain information without engaging in costly trial-and-error learning, first-time builders should exploit the successful habits of experienced builders. We presented first-time nest-building male zebra finches with either a familiar or an unfamiliar conspecific male building with material of a colour the observer did not like. When given the opportunity to build, males that had watched a familiar male build switched their material preference to that used by the familiar male. Males that observed unfamiliar birds did not. Thus, first-time nest builders use social information and copy the nest material choices when demonstrators are familiar but not when they are strangers. The relationships between individuals therefore influence how nest-building expertise is socially transmitted in zebra finches., (© 2016 The Author(s).)

An animal's behavior is affected by its cognitive abilities, which are, in turn, a consequence of the environment in which an animal has evolved and developed. Although behavioral ecologists have been studying animals in their natural environment for several decades, over much the same period animal cognition has been studied almost exclusively in the laboratory. Traditionally, the study of animal cognition has been based on well-established paradigms used to investigate well-defined cognitive processes. This allows identification of what animals can do, but may not, however, always reflect what animals actually do in the wild. As both ecologists and some psychologists increasingly try to explain behaviors observable only in wild animals, we review the different motivations and methodologies used to study cognition in the wild and identify some of the challenges that accompany the combination of a naturalistic approach together with typical psychological testing paradigms. We think that studying animal cognition in the wild is likely to be most productive when the questions addressed correspond to the species' ecology and when laboratory cognitive tests are appropriately adapted for use in the field. Furthermore, recent methodological and technological advances will likely allow significant expansion of the species and questions that can be addressed in the wild., (© 2016 Society for the Experimental Analysis of Behavior.)

Animals may remember an important location with reference to one or more visual landmarks. In the laboratory, birds and mammals often preferentially use landmarks near a goal ("local landmarks") to return to that location at a later date. Although we know very little about how animals in the wild use landmarks to remember locations, mammals in the wild appear to prefer to use distant landmarks to return to rewarded locations. To examine what cues wild birds use when returning to a goal, we trained free-living hummingbirds to search for a reward at a location that was specified by three nearby visual landmarks. Following training we expanded the landmark array to test the extent that the birds relied on the local landmarks to return to the reward. During the test the hummingbirds' search was best explained by the birds having used the experimental landmarks to remember the reward location. How the birds used the landmarks was not clear and seemed to change over the course of each test. These wild hummingbirds, then, can learn locations in reference to nearby visual landmarks., (Copyright © 2015 Elsevier B.V. All rights reserved.)

Although small-scale navigation is well studied in a wide range of species, much of what is known about landmark use by vertebrates is based on laboratory experiments. To investigate how vertebrates in the wild use landmarks, we trained wild male rufous hummingbirds to feed from a flower that was placed in a constant spatial relationship with two artificial landmarks. In the first experiment, the landmarks and flower were 0.25, 0.5 or 1 m apart and we always moved them 3-4 m after each visit by the bird. In the second experiment, the landmarks and flower were always 0.25 m apart and we moved them either 1 or 0.25 m between trials. In tests, in which we removed the flower, the hummingbirds stopped closer to the predicted flower location when the landmarks had been closer to the flower during training. However, while the distance that the birds stopped from the landmarks and predicted flower location was unaffected by the distance that the landmarks moved between trials, the birds directed their search nearer to the predicted direction of the flower, relative to the landmarks, when the landmarks and flower were more stable in the environment. In the field, then, landmarks alone were sufficient for the birds to determine the distance of a reward but not its direction.

In nature, many animals build structures that can be readily measured at the scale of their gross morphology (e.g. length, volume and weight). Capturing individuality as can be done with the structures designed and built by human architects or artists, however, is more challenging. Here, we tested whether computer-aided image texture classification approaches can be used to describe textural variation in the nests of weaverbirds (Ploceus species) in order to attribute nests to the individual weaverbird that built them. We found that a computer-aided texture analysis approach does allow the assignment of a signature to weaverbirds' nests. We suggest that this approach will be a useful tool with which to examine individual variation across a range of animal constructions, not just for nests.

It is now well established that individuals can differ consistently in their average levels of behaviour across different contexts. There have recently been calls to apply the same adaptive framework to interindividual differences in cognition. These calls have culminated in the suggestion that variation in personality and cognition should correlate. We suggest that both these appealing notions are conceptually and logistically problematic. We identify the first crucial step for establishing any cognition-personality relationship. This is to determine the degree to which cognitive abilities yield consistent task performance. We then suggest how to establish whether such consistency exists. Finally, we discuss why formulating predictions about how cognition might be related to personality is much more difficult than is currently realised., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

During nest building in zebra finches (Taeniopygia guttata), several regions in the social behaviour network and the dopaminergic reward system, which are two neural circuits involved in social behaviour, appear to be active in male and female nest-building finches. Because the nonapeptides, mesotocin and vasotocin and the neurotransmitter, dopamine, play important roles in avian social behaviour, we tested the hypothesis that mesotocinergic-vasotocinergic and dopaminergic neuronal populations in the social behaviour network and dopaminergic reward system, respectively, are active during nest building. We combined immunohistochemistry for Fos (an indirect marker of neuronal activity) and vasotocin, mesotocin or tyrosine hydroxylase on brain tissue from nest-building and non-nest-building male and female zebra finches and compared Fos immunoreactivity in these neuronal populations with the variation in nest-building behaviour. Fos immunoreactivity in all three types of neuronal populations increased with some aspect of nest building: (i) higher immunoreactivity in a mesotocinergic neuronal population of nest-building finches compared to controls; (ii) increased immunoreactivity in the vasotocinergic neuronal populations in relation to the amount of material picked up by nest-building males and the length of time that a male spent in the nest with his mate; and (iii) increased immunoreactivity in a dopaminergic neuronal population in relation to the length of time that a male nest-building finch spent in the nest with his mate. Taken together, these findings provide evidence for a role of the mesotocinergic-vasotocinergic and dopaminergic systems in avian nest building., (© 2014 The Authors. Journal of Neuroendocrinology published by John Wiley & Sons Ltd on behalf of British Society for Neuroendocrinology.)

Despite centuries of observing the nest building of most extant bird species, we know surprisingly little about how birds build nests and, specifically, how the avian brain controls nest building. Here, we argue that nest building in birds may be a useful model behaviour in which to study how the brain controls behaviour. Specifically, we argue that nest building as a behavioural model provides a unique opportunity to study not only the mechanisms through which the brain controls behaviour within individuals of a single species but also how evolution may have shaped the brain to produce interspecific variation in nest-building behaviour. In this review, we outline the questions in both behavioural and comparative neuroscience that nest building could be used to address, summarize recent findings regarding the neurobiology of nest building in lab-reared zebra finches and across species building different nest structures, and suggest some future directions for the neurobiology of nest building.

Free-living hummingbirds can learn the refill schedules of individual experimental flowers but little is known about what information they use to do this. Colour cues, in particular, may be important to hummingbirds when learning about rewarded flower properties. We investigated, therefore, whether colour cues facilitated the learning of flower refill schedules in wild, free-living rufous hummingbirds (Selasphorus rufus). In the Cued condition, we presented birds with an array of six flowers, three of one colour, each of which were refilled 10min after being emptied by the bird and three of a different colour, which were refilled 20min after being emptied. In the Uncued condition we presented birds with six flowers of the same colour, three of which were refilled after 10min and three of which were refilled after 20min as for the birds in the Cued condition. In the second part of the experiment, we moved the array 2m and changed the shape of the array. Across both phases, birds in the Cued condition learned to discriminate between 10 and 20-min flowers more quickly than did the birds in the Uncued condition. The Cued birds were also better at discriminating between the two distinct refill intervals. Colour cues can, therefore, facilitate learning the refill schedules of experimental flowers in these birds. This article is part of a Special Issue entitled: Cognition in the wild., (Copyright © 2014 Elsevier B.V. All rights reserved.)