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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.

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.

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.

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.

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).)

It is generally assumed that birds' choice of structurally suitable materials for nest building is genetically predetermined. Here, we tested that assumption by investigating whether experience affected male zebra finches' (Taeniopygia guttata) choice of nest material. After a short period of building with relatively flexible string, birds preferred to build with stiffer string while those that had experienced a stiffer string were indifferent to string type. After building a complete nest with either string type, however, all birds increased their preference for stiff string. The stiffer string appeared to be the more effective building material as birds required fewer pieces of stiffer than flexible string to build a roofed nest. For birds that raised chicks successfully, there was no association between the material they used to build their nest and the type they subsequently preferred. Birds' material preference reflected neither the preference of their father nor of their siblings but juvenile experience of either string type increased their preference for stiffer string. Our results represent two important advances: (i) birds choose nest material based on the structural properties of the material; (ii) nest material preference is not entirely genetically predetermined as both the type and amount of experience influences birds' choices.

While most animals live in a three-dimensional world, they move through it to different extents depending on their mode of locomotion: terrestrial animals move vertically less than do swimming and flying animals. As nearly everything we know about how animals learn and remember locations in space comes from two-dimensional experiments in the horizontal plane, here we determined whether the use of three-dimensional space by a terrestrial and a flying animal was correlated with memory for a rewarded location. In the cubic mazes in which we trained and tested rats and hummingbirds, rats moved more vertically than horizontally, whereas hummingbirds moved equally in the three dimensions. Consistent with their movement preferences, rats were more accurate in relocating the horizontal component of a rewarded location than they were in the vertical component. Hummingbirds, however, were more accurate in the vertical dimension than they were in the horizontal, a result that cannot be explained by their use of space. Either as a result of evolution or ontogeny, it appears that birds and rats prioritize horizontal versus vertical components differently when they remember three-dimensional space.

The ability of animals to remember the what, where and when of a unique past event is used as an animal equivalent to human episodic memory. We currently view episodic memory as reconstructive, with an event being remembered in the context in which it took place. Importantly, this means that the components of a what, where, when memory task should be dissociable (e.g. what would be remembered to a different degree than when). We tested this hypothesis by training hummingbirds to a memory task, where the location of a reward was specified according to colour (what), location (where), and order and time of day (when). Although hummingbirds remembered these three pieces of information together more often than expected, there was a hierarchy as to how they were remembered. When seemed to be the hardest to remember, while errors relating to what were more easily corrected. Furthermore, when appears to have been encoded as a combination of time of day and sequence information. As hummingbirds solved this task using reconstruction of different memory components (what, where and when), we suggest that similar deconstructive approaches may offer a useful way to compare episodic and episodic-like memories.

In recent years, there have been over 50 comparative analyses carried out in which social or ecological variables have been used to explain variation in whole brain size, or a part thereof, in a range of vertebrate species. Here, we review this body of work, pointing out that there are a number of substantial problems with some of the assumptions that underpin the hypotheses (e.g. what brain size means), with the data collection and with the ways in which the data are combined in the analyses. These problems are particularly apparent in those analyses in which attempts are made to correlate complex behaviour with parts of the brain that carry out multiple functions. We conclude that now is the time to substantiate these results with data from experimental manipulations.

In mammals, stress hormones have profound influences on spatial learning and memory. Here, we investigated whether glucocorticoids influence cognitive abilities in birds by testing a line of zebra finches selectively bred to respond to an acute stressor with high plasma corticosterone (CORT) levels. Cognitive performance was assessed by spatial and visual one-trial associative memory tasks. Task performance in the high CORT birds was compared with that of the random-bred birds from a control breeding line. The birds selected for high CORT in response to an acute stressor performed less well than the controls in the spatial task, but there were no significant differences between the lines in performance during the visual task. The birds from the two lines did not differ in their plasma CORT levels immediately after the performance of the memory tasks; nevertheless, there were significant differences in peak plasma CORT between the lines. The high CORT birds also had significantly lower mineralocorticoid receptor mRNA expression in the hippocampus than the control birds. There was no measurable difference between the lines in glucocorticoid receptor mRNA density in either the hippocampus or the paraventricular nucleus. Together, these findings provide evidence to suggest that stress hormones have important regulatory roles in avian spatial cognition.

Men and women differ in their ability to solve spatial problems. There are two possible proximate explanations for this: (i) men and women differ in the kind (and value) of information they use and/or (ii) their cognitive abilities differ with respect to spatial problems. Using a simple computerized task which could be solved either by choosing an object based on what it looked like, or by its location, we found that the women relied on the object's visual features to solve the task, while the men used both visual and location information. There were no differences between the sexes in memory for the visual features of the objects, but women were poorer than men at remembering the locations of objects.

A core assumption implicit in economic models of animal choice is that subjects assign absolute utilities to options that are independent of the type and number of alternatives available. Humans sometimes appear to violate this assumption and employ relative, as opposed to absolute, currencies when making choices. Recent evidence suggests that animals too might sometimes employ relative choice mechanisms. We tested this idea by measuring the foraging preferences of rufous hummingbirds (Selasphorus rufus) faced with choices analogous to those in which human use of relative currencies is evident. The birds experienced three treatments: a binary choice between two artificial flower types designated concentration (20 microl, 40% sucrose solution) and volume (40 microl, 20%), and two trinary treatments in which a third decoy option (either concentration decoy: 10 microl, 30% or volume decoy: 30 microl, 10%) was added to the set. The birds' preferences differed significantly across the three treatments. In the trinary treatments, the effect of the decoy options was to increase the preference for the option that dominated the decoy. These results are similar to those reported in the human choice literature, and are compatible with the hummingbirds using a relative evaluation mechanism in decision making.

In a number of mammalian species, males appear to have superior spatial abilities to females. The favoured explanations for this cognitive difference are hormonal, with higher testosterone levels in males than females leading to better spatial performance, and evolutionary, where sexual selection has favoured males with increased spatial abilities for either better navigational skills in hunting or to enable an increased territory size. However, an alternative explanation for this sex difference focuses on the role of varying levels of oestrogen in females in spatial cognition (the 'fertility and parental care' hypothesis). One possibility is that varying oestrogen levels result in variation in spatial learning and memory so that, when tested across the oestrous cycle, females perform as well as males on days of low oestrogen but more poorly on days of high oestrogen. If day in the oestrous cycle is not taken into account then, across an experiment, any sex differences found would always produce male superiority. We used a spatial working memory task in a Morris water maze to test the spatial learning and memory abilities of male and female rats. The rats were tested across a number of consecutive days during which the females went through four oestrous cycles. We found no overall sex differences in latencies to reach a submerged platform in a Morris water maze but, on the day of oestrus (low oestrogen), females took an extra swim to learn the platform's location (a 100% increase over the other days in the cycle). Female swim speed also varied across the oestrous cycle but females were no less active on the day of oestrus. These results oppose the predictions of the fertility and parental care hypothesis.

Song and brain structure are compared amongst 41 species of oscine birds by using the method of independent evolutionary contrasts. We find a significant correlation between the relative volume of the song control centre, the high vocal centre (HVC), and the number of song types typically found in the repertoire. Relative HVC volume is not correlated with the number of different syllable types per song bout. The relative volume of a second song nucleus, area X, is not significantly correlated with either measure. Relative HVC volume is uncorrelated with relative volume of the hippocampus, a brain area involved in other forms of memory. This is the first evidence for repeated independent evolution of an association between complexity of learned song and the relative volume of one of the song control nuclei though to be involved in song learning.