Anthropogenic nest material use in a global sample of birds.

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.

Nest architecture influences host use by avian brood parasites and is shaped by coevolutionary dynamics.

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.

Bird nest building: visions for the future.

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'.

Beak shape and nest material use in birds.

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'.

Adding the neuro to cognition: from food storing to nest building.

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?

Global drivers of variation in cup nest size in passerine birds.

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.

Nests and nest building in birds.

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).

Size is relative: use of relational concepts by wild hummingbirds.

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.

It Began in Ponds and Rivers: Charting the Beginnings of the Ecology of Fish Cognition.

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.

Involvement of the neural social behaviour network during social information acquisition in zebra finches (Taeniopygia guttata).

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.

Hummingbirds modify their routes to avoid a poor location.

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.

Reproductive consequences of material use in avian nest construction.

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.

Not by transmission alone: the role of invention in cultural evolution.

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'.

Object manipulation without hands.

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.

Estimating on the fly: The approximate number system in rufous hummingbirds (Selasphorus rufus).

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.

The rationality of decisions depends on behavioural context.

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.

Neural Circuits Underlying Nest Building in Male Zebra Finches.

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.

Numerical ordinality in a wild nectarivore.

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.

Ecology and allometry predict the evolution of avian developmental durations.

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.

The Impact of Acute Loud Noise on the Behavior of Laboratory Birds.

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.