Joah R Madden

• Senior Lecturer at University of Exeter

Wildlife–vehicle collisions (WVCs) cause millions of vertebrate mortalities globally, threatening population viability and influencing wildlife behaviour and survival. Traffic volume and speed can influence wildlife mortality on roads, but roadkill risk is species specific and depends on ecological traits. The COVID‐19 pandemic, and associated UK‐w...

Most animals confine their activities to a discrete home range, long assumed to reflect the fitness benefits of obtaining spatial knowledge about the landscape. However, few empirical studies have linked spatial memory to home range development or determined how selection operates on spatial memory via the latter’s role in mediating space use. We a...

Understanding animal movement is essential to elucidate how animals interact, survive, and thrive in a changing world. Recent technological advances in data collection and management have transformed our understanding of animal “movement ecology” (the integrated study of organismal movement), creating a big-data discipline that benefits from rapid,...

In group-living vertebrates, dominance status often covaries with physiological measurements (e.g. glucocorticoid levels), but it is unclear how dominance is linked to dynamic changes in physiological state over a shorter, behavioural timescale. In this observational study, we recorded spontaneous aggression among captive juvenile pheasants ( Phasi...

The release of gamebirds for recreational shooting exerts a series of effects on the ecosystems into which they are placed. Pheasants (Phasianus colchicus) are omnivorous and eat invertebrates, especially when young or, if females, when breeding. Consequently, the release of large numbers of pheasants into woodland release pens may affect local inv...

Cognitive biases for encoding spatial information (orientation strategies) in relation to self (egocentric) or landmarks (allocentric) differ between species or populations according to the habitats they occupy. Whether biases in orientation strategy determine early habitat selection or if individuals adapt their biases following experience is unkn...

The structure of a group is critical in determining how a socially learnt behaviour will spread. Predictions from theoretical models indicate that specific parameters of social structure differentially influence social transmission. Modularity describes how the structure of a group or network is divided into distinct subgroups or clusters. Theoreti...

Memories about the spatial environment, such as the locations of foraging patches, are expected to affect how individuals move around the landscape. However, individuals differ in the ability to remember spatial locations (spatial cognitive ability) and evidence is growing that these inter-individual differences influence a range of fitness proxies...

This review examines 128 items of primary and other literature to provide an insight into current knowledge of the effects of pheasant and red‐legged partridge releasing and associated management for shooting on habitats and wildlife in the UK. It summarizes key findings and uses them to define sub‐topic sections for which the effects are classifie...

To understand the evolution of cognitive abilities, we need to understand both how selection acts upon them and their genetic (co)variance structure. Recent work suggests that there are fitness consequences for free-living individuals with particular cognitive abilities. However, our current understanding of the heritability of these abilities is r...

Social environments influence important ecological processes and can determine how selection acts on traits. Cognitive abilities can shape these social environments and in turn, affect individuals' fitness. To understand how cognitive abilities evolve, we need to understand the complex interplay between an individual's cognitive abilities, the soci...

The differential specialization of each side of the brain facilitates the parallel processing of information and has been documented in a wide range of animals. Animals that are more lateralized as indicated by consistent preferential limb use are commonly reported to exhibit superior cognitive ability as well as other behavioural advantages. We as...

The ability to inhibit prepotent actions towards rewards that are made inaccessible by transparent barriers has been considered to reflect capacities for inhibitory control (IC). Typically, subjects initially reach directly, and incorrectly, for the reward. With experience, subjects may inhibit this action and instead detour around barriers to acce...

Inhibitory control (IC) is the ability to intentionally restrain initial, ineffective responses to a stimulus and instead exhibit an alternative behaviour that is not pre-potent but which effectively attains a reward. Individuals (both humans and non-human animals) differ in their IC, perhaps as a result of the different environmental conditions th...

The ability to inhibit prepotent actions towards rewards that are made inaccessible by transparent barriers has been considered to reflect capacities for inhibitory control (IC). Typically, subjects initially reach directly, and incorrectly, for the reward. With experience, subjects may inhibit this action and instead detour around barriers to acce...

The ability to control impulsive actions is an important executive function that is central to the self-regulation of behaviours and, in humans, can have important implications for mental and physical health. One key factor that promotes individual differences in inhibitory control (IC) is the predictability of environmental information experienced...

The recent decline in farmland songbirds in the UK has coincided with increases in the populations of many nest predators. However, studies which have removed nest predators and monitored the response of prey populations have found mixed results. One explanation for this ambiguity is that, within species, predators differ in how likely they are to...

Individual differences in performances on cognitive tasks have been found to differ according to social rank across multiple species. However, it is not clear whether an individual’s cognitive performance is flexible and the result of their current social rank, modulated by social interactions (social state dependent hypothesis), or if it is determ...

Dataset for Qu 2: Spatial learning performances of adult male pheasants following a rank manipulation and their most recent social rank Does variation in cognitive performances persist once social rank is standardised?

Dataset for Qu 3: Spatial learning performances of male pheasant chicks and adult social rank Do chick cognitive performances predict their future social rank?

Dataset for Qu 1: Chick and adult spatial learning performances Chick and adult spatial learning performances, measures generated from learning curves (glms) fitted on 20 trials.

Description of Supplemental Data Files

Rates of dominance behaviour by male pheasants while housed in two different social conditions Rates of dominance behaviour (crowing and lateral displays) by male pheasants while housed in two different social conditions.

Fine scale sexual segregation outside of the mating season is common in sexually dimorphic and polygamous species, particularly in ungulates. A number of hypotheses predict sexual segregation but these are often contradictory with no agreement as to a common cause, perhaps because they are species specific. We explicitly tested three of these hypot...

Individuals vary in their cognitive performance. While this variation forms the foundation of the study of human psychometrics, its broader importance is only recently being recognized. Explicitly acknowledging this individual variation found in both humans and non-human animals provides a novel opportunity to understand the mechanisms, development...

Cognitive abilities probably evolve through natural selection if they provide individuals with fitness benefits. A growing number of studies demonstrate a positive relationship between performance in psychometric tasks and (proxy) measures of fitness. We assayed the performance of 154 common pheasant (Phasianus colchicus) chicks on tests of acquisi...

Brain lateralization is considered adaptive because it leads to behavioral biases and specializations that bring fitness benefits. Across species, strongly lateralized individuals perform better in specific behaviors likely to improve survival. What constrains continued exaggerated lateralization? We measured survival of pheasants, finding that ind...

To understand how natural selection may act on cognitive processes, it is necessary to reliably determine interindividual variation in cognitive abilities. However, an individual's performance in a cognitive test may be influenced by the social environment. The social environment explains variation between species in cognitive performances, with sp...

Individuals likely vary in how quickly they learn, how many different associations they may be able to maintain, and how long they can remember previously learned associations. However, it is unclear whether capacities for these cognitive processes are consistent within individuals, or whether individual performance differs when presented with nove...

Intra-individual variation in performance within and across cognitive domains may confound interpretations of both domain-general and domain-specific abilities. Such variation is rarely considered in animal test batteries. We investigate individual consistency in performance by presenting pheasant chicks (n = 31), raised under standardized conditio...

It remains unclear whether performance of non-human animals on cognitive test batteries can be explained by domain general cognitive processes, as is found in humans. The persistence of this dispute is likely to stem from a lack of clarity of the psychological or neural processes involved. One broadly accepted cognitive process, that may predict pe...

Around 60% of pheasants released for shooting in the UK, an estimated 21 million birds, do not end up at their intended fate: being shot. This constitutes wastage, raising economic, environmental and ethical questions. We review what is known of the fates of released pheasants and consider why they do not directly contribute to the numbers harveste...

Sexual segregation is common and can occur when sexes occupy different habitats, and/or when sexes aggregate assortatively within the same habitats. However, it is rarely studied in birds, with most previous work concentrating on differential settlement by the sexes in discrete habitats, often separated by large distances. Little attention has been...

Transparent Cylinder and Barrier tasks are used to purportedly assess inhibitory control in a variety of animals. However, we suspect that performances on these detour tasks are influenced by non-cognitive traits, which may result in inaccurate assays of inhibitory control. We therefore reared pheasants under standardized conditions and presented e...

Methods, pictures of test apparatuses and flow diagram of inhibitory control performance across trials

Dominant individuals differ from subordinates in their performances on cognitive tasks across a suite of taxa. Previous studies often only consider dyadic relationships, rather than the more ecologically relevant social hierarchies or networks, hence failing to account for how dyadic relationships may be adjusted within larger social groups. We use...

Inhibitory control enables subjects to quickly react to unexpectedly changing external demands. We assessed the ability of young (8 weeks old) pheasants Phasianus colchicus to exert inhibitory control in a novel response-inhibition task that required subjects to adjust their movement in space in pursuit of a reward across changing target locations....

Pheasants (Phasianus colchicus) are commonly killed on UK roads, presenting a threat to motorists and a loss to the game shooting industry. Pheasants may be inherently susceptible, or the recent increase in their artificial rearing and release may have exacerbated the situation, either through population increases or because artificial rearing has...

Hypotheses for why animals sexually segregate typically rely on adult traits, such as differences in sexual roles causing differential habitat preferences, or size dimorphism inducing differences in diet or behaviour. However, segregation can occur in juveniles before such roles or size dimorphism is well established. In young humans, leading hypot...

In many areas of animal behaviour research, improvements in our ability to collect large and detailed data sets are outstripping our ability to analyse them. These diverse, complex and often high-dimensional data sets exhibit nonlinear dependencies and unknown interactions across multiple variables, and may fail to conform to the assumptions of man...

Simple Summary In this placebo controlled study, we exposed 28 dogs to Pet Remedy (a natural stress relief product) to investigate whether Pet Remedy lowered stress-affected behaviour. No statistically significant differences were found when dogs were exposed to Pet Remedy or the placebo condition. We suggest that Pet Remedy, in this particular stu...

A strong motivation for one individual to aggregate with others is to reduce their vigilance because other group members provide coverage and warning of approaching predators. This collective vigilance means that a focal individual is usually less susceptible to predation than when alone. However, individuals differ in their vigilance levels depend...

Poster Presentation at ISAE Congress 2016

Stress affecting behaviour in companion animals can have an adverse effect on animal health and welfare and their relationships with humans. This stress can be addressed using chemical treatments, often in conjunction with behavioural therapies. Pet Remedy, a natural valerian based stress relief product for mammals is commercially available on th...

Poster Presentation

Rates of innovative foraging behaviours and success on problem-solving tasks are often used to assay differences in cognition, both within and across species. Yet the cognitive features of some problem-solving tasks can be unclear. As such, explanations that attribute cognitive mechanisms to individual variation in problem-solving performance have...

Subtle variations in early rearing environment influence morphological, cognitive and behavioural processes that together impact on adult fitness. We manipulated habitat complexity experienced by young pheasants (Phasianus colchicus) in their first seven weeks, adding a third accessible dimension by placing elevated perches in their rearing pens mi...

Behavioural and physiological deficiencies are major reasons why reintroduction programmes suffer from high mortality when captive animals are used. Mitigation of these deficiencies is essential for successful reintroduction programmes. Our study manipulated early developmental diet to better replicate foraging behaviour in the wild. Over 2 years,...

Our understanding of the processes underlying animal cognition has improved dramatically in recent years, but we still know little about how cognitive traits evolve. Following Darwinian logic, to understand how selection acts on such traits we must determine whether they vary between individuals, influence fitness, and are heritable. A handful of r...

Third party punishment can be evolutionarily stable if there is heterogeneity in the cost of punishment or if punishers receive a reputational benefit from their actions. A dominant position might allow some individuals to punish at a lower cost than others and by doing so access these reputational benefits. Three vignette-based studies measured pa...

Social network analysis has become an increasingly popular method to link individual behaviour to population level patterns (and vice versa). Technological advances of recent years, such as the development of spatial proximity loggers, have enhanced our abilities to record contact patterns between animals. However, loggers are often deployed withou...

Individuals exhibiting a high level of cognitive ability may also exhibit more elaborate traits and so gain higher levels of mating success. This suggests that selection may act on cognitive performance through mate choice. Studies investigating this relationship have tended to focus on single cognitive tasks, or tasks that are closely related to e...

Females vary in their mate choice and consequent fitness outcomes. Individual differences may be explained by conditions experienced early in life. We tested whether the sex ratio at which young pheasants, Phasianus colchicus, were reared affected their adult sexual behaviour. Females reared in equal sex ratios discriminated strongly between males...

Recreational hunting can disrupt the population structure or alter the morphology of target populations. More subtly, such hunting may alter the behaviour of individuals in the target population, especially if individuals are culled nonrandomly. We assayed the behavioural temperaments of a sample of hand-reared and released pheasants, Phasianus col...

Cultivation may be described as a process of co-evolution and niche construction, with two species developing a mutualistic relationship through association, leading to coordinated change [1 • Rindos D. The Origins of Agriculture: An Evolutionary Perspective. Academic Press, New York1984 • Google Scholar ]. Cultivation is rare but taxonomically w...

The underlying kin structure of groups of animals may be glimpsed from patterns of spatial position or temporal association between individuals, and is presumed to facilitate inclusive fitness benefits. Such structure may be evident at a finer, behavioural, scale with individuals preferentially interacting with kin. We tested whether kin structure...

The Montserrat Oriole (Icterus oberi) is a critically endangered species, confined to a small range in the hill forests of the volcanic island of Montserrat in the eastern Caribbean. From 1998 to 2005 we studied its breeding biology and survival of adults, finding that the Montserrat Oriole has a smaller clutch, more extended parental care, and hig...

Individuals in social groups interact with numerous other group members in a polyadic network. Interactions can depend on the individual's own attributes (age, sex, status etc.), on their partner's attributes, and the group's network of social interactions. Previous studies tend to look at a subset of dyadic interactions, focusing on particular cla...

Behavioural ecologists are increasingly using social network analysis to describe the social organisation of animal populations and to test hypotheses. However, the statistical analysis of network data presents a number of challenges. In particular the non-independent nature of the data violates the assumptions of many common statistical approaches...

The evolution and expression of different forms of cooperative behaviour (e.g. feeding, guarding, sentinel duties, etc.) are usually studied independently, with few studies considering them as a single syndrome. However, studies investigating individuals' investment across a suite of different behaviours reveal that they are correlated, suggesting...

Knowledge of the structure of networks of social interactions is important for understanding the evolution of cooperation, transmission of disease, and patterns of social learning, yet little is known of how environmental, ecological, or behavioural factors relate to such structures within groups. We observed grooming, dominance, and foraging compe...

Begging by young provokes adults to provide food for them. However, eventually begging by young and provisioning by adults cease and young become nutritionally independent. Why do young cease begging and so forgo food brought to them by adults? Three explanations have been proposed: (1) adults may not respond to begging anymore and cease feeding be...

Adults vary in their generosity in provisioning the young and their sensitivity to the need of the young. Do the young modulate their behaviour so as to specifically target more high-provisioning adults? This is especially likely in situations with mobile, nutritionally dependent young. We studied cooperatively breeding meerkats Suricata suricatta,...

Groups of individuals frequently interact with each other, but typically analysis of such interactions is restricted to isolated dyads. Social network analysis (SNA) provides a method of analysing polyadic interactions and is used to analyse interactions between individuals. We use a population of 12 groups (ca. 250 animals) of wild meerkats (Suric...

It is thought that allogrooming is practised strategically in order to establish, maintain and reinforce social bonds between group members, exchanging one altruistic behaviour for a different form of reciprocated benefit at a later date. Correlational evidence supports this, but evidence of causality is lacking. We reduced parasite loads in eight...

Offspring are frequently raised alongside their siblings and are provisioned early in life by adults. Adult provisioning is stimulated by offspring begging, but it is unclear how each offspring should beg, given the begging behaviour of their siblings. It has previously been suggested that siblings may compete directly through begging for a fixed l...

In many bird species with biparental care for young in the nest, hungry chicks beg repeatedly, and parents adjust their feeding rate to the call rate of young. Repetitive calling also occurs in fledglings and in some mammals where offspring follow provisioners. It is not yet clear whether, in mobile systems with dispersed young where adults cannot...

Baylis (1982, Acoustic Communication in Birds, Academic Press) decried the serious lack of experimental verification for the various hypotheses proposed to explain vocal mimicry in songbirds. With few exceptions, our understanding of the function and acquisition of this fascinating behaviour seems to have scarcely progressed. We examine the propose...

In species where young are provisioned by both parents, males commonly contribute less to parental care than females and are less responsive to variation in begging rates. Similar differences in the care of young occur among adults in cooperative breeders but fewer studies have investigated whether these are associated with differences in responsiv...

Definitions of what features constitute cultural behaviour, and hence define cultures are numerous. Many seem designed to describe those aspects of human behaviour which set us apart from other animals. A broad definition prescribing that the behaviour is: learned; learned socially; normative and collective is considered to apply to several species...

In some species, dependent offspring join foraging providers and beg for food. Mobile offspring might benefit from evolving begging signals adapted to the different situations they are exposed to, but this possibility has been ignored. In cooperatively breeding meerkats (Suricata suricatta), dependent offspring use a repertoire of several begging c...

Much attention has been paid to innovative foraging methods, but little to innovative sexual displays. Innovations may be common in behavioural display, such as song or object use, and could occur in both male display form and female preferences. Similar evidence exists for innovation in display as in foraging methods, but in smaller quantities. Ra...

The structure of common cuckoo nestling begging calls differs between the two host-races parasitizing reed warblers (reed warbler-cuckoos) and dunnocks (dunnock-cuckoos; longer syllable duration, lower peak and maximum frequency, narrower bandwidth). Cross-fostering experiments demonstrated that this difference is not genetically fixed but develops...

Male Spotted Bowerbirds Chlamydera maculata build and decorate structures, bowers, that act as targets for female choice. Bower construction, orientation and distribution all differed between two populations, as did rates of intrasexual aggression. Although there was a broad similarity in classes of decorations used on bowers, the exact types and p...

The common cuckoo has several host-specific races, each with a distinctive egg that tends to match its host's eggs. Here, we show that the host-race specializing on reed warblers also has a host-specific nestling adaptation. In playback experiments, the nestling cuckoos responded specifically to the reed warbler's distinctive 'churr' alarm (given w...

We tested whether nestlings of a generalist brood parasite, the brown-headed cowbird, Molothrus ater, had special adaptations for tuning into host parent food and alarm calls. We compared the responses of 7-day-old cowbird nestlings reared by eastern phoebes, Sayornis phoebe, with those of a related, nonparasitic icterid, the red-winged blackbird,...

Begging by nestlings can prove costly, either through energy expenditure if food-bearing parents are not present, or through increased predation risk. Therefore, parents may provide cues to modulate begging. We investigated responses of 7-day old nestlings of eastern phoebes, Sayornis phoebe, and red-winged blackbirds, Agelaius phoeniceus, to vario...

Animal color pattern phenotypes evolve rapidly. What influences their evolution? Because color patterns are used in communication, selection for signal efficacy, relative to the intended receiver's visual system, may explain and predict the direction of evolution. We investigated this in bowerbirds, whose color patterns consist of plumage, bower st...

Animal color pattern phenotypes evolve rapidly. What influences their evolution? Because color patterns are used in communication, selection for signal efficacy, relative to the intended receiver's visual system, may explain and predict the direction of evolution. We investigated this in bowerbirds, whose color patterns consist of plumage, bower st...

Parent birds often give alarm calls when a predator approaches their nest. However, it is not clear whether these alarms function to warn nestlings, nor is it known whether nestling responses are species-specific. The parental alarms of reed warblers, Acrocephalus scirpaceus ("churr"), dunnocks, Prunella modularis ("tseep"), and robins, Erithacus r...

Males with elaborate secondary sexual traits can enhance their mating success both by choosing to display at a site that increases the efficacy of their signal and by displaying at a site where the population is structured to provide inclusive fitness benefits via kin selection. Theoretical models and field studies reveal that, across a range of sp...

Recent work has shown that elaborate secondary sexual traits and the corresponding preferences for them may be transmitted culturally rather than by genetic inheritance. Evidence for such cultural transmission commonly invokes spatial patterns of local similarity, with neighbouring individuals or populations appearing similar to each other. Alterna...

Young brood parasites that tolerate the company of host offspring challenge the existing evolutionary view of family life. In theory, all parasitic nestlings should be ruthlessly self-interested and should kill host offspring soon after hatching. Yet many species allow host young to live, even though they are rivals for host resources. Here we show...

The Spotted Bowerbird, Chlamydera maculata, appears to be sexually monomorphic. We caught and marked 118 birds in central Queensland, and sexed 88 using molecular methods. We found that our catch was strongly male-biased, both at bower sites and at non-bower feeding sites. We continued to observe the bird's behaviour after their release and so sub-...

Females target elaborate secondary sexual traits to acquire either direct benefits, represented or provided by the trait, or indirect benefits usually in the form of genetic components. The link between a males trait and his potential genetic contribution is often unclear. Bowers are extreme examples of secondary sexual traits, built and decorated...

The sensory bias model of sexual selection suggests that elaborate male secondary sexual traits evolved to exploit biases in the female's sensory system. Such biases may have evolved in a nonsexual context. Male bowerbirds build and decorate elaborate structures, bowers, which function as targets of female choice. The colour of decorations used on...

Variation in the mating success of males can often be predicted by considering measures of their exaggerated sexual traits. Male spotted bowerbirds Chlamydera maculata build and decorate elaborate structures--bowers--that function in mate choice. I show that numbers of certain decorations correlate with the mating success of the bower owner. Specif...