The impact of detoxification costs and predation risk on foraging : implications for mimicry dynamics

This work was supported by the European Research Council (Advanced Grant 250209 to Alasdair Houston), a Natural Environment Research Council Independent Research Fellowship (NE/L011921/1) awarded to A.D.H., a BBSRC-NERC project grant (BB/G00188X/1) awarded to J.S., C.R. and G.D.R. and a faculty fellowship awarded to C.G.H. (Medical Sciences, Newcastle University) with strategic support funding from the Wellcome Trust.Prey often evolve defences to deter predators, such as noxious chemicals including toxins. Toxic species often advertise their defence to potential predators by distinctive sensory signals. Predators learn to associate toxicity with the signals of these so-called aposematic prey, and may avoid them in future. In turn, this selects for mildly toxic prey to mimic the appearance of more toxic prey. Empirical evidence shows that mimicry could be either beneficial (‘Mullerian’) or detrimental (‘quasi-Batesian’) to the highly toxic prey, but the factors determining which are unknown. Here, we use state-dependent models to explore how tri-trophic interactions could influence the evolution of prey defences. We consider how predation risk affects predators’ optimal foraging strategies on aposematic prey, and explore the resultant impact this has on mimicry dynamics between unequally defended species. In addition, we also investigate how the potential energetic cost of metabolising a toxin can alter the benefits to eating toxic prey and thus impact on predators’ foraging decisions. Our model predicts that both how predators perceive their own predation risk, and the cost of detoxification, can have significant, sometimes counterintuitive, effects on the foraging decisions of predators. For example, in some conditions predators should: (i) avoid prey they know to be undefended, (ii) eat more mildly toxic prey as detoxification costs increase, (iii) increase their intake of highly toxic prey as the abundance of undefended prey increases. These effects mean that the relationship between a mimic and its model can qualitatively depend on the density of alternative prey and the cost of metabolising toxins. In addition, these effects are mediated by the predators’ own predation risk, which demonstrates that, higher trophic levels than previously considered can have fundamental impacts on interactions among aposematic prey species.Publisher PDFPeer reviewe

Pattern contrast influences wariness in naïve predators towards aposematic patterns

This work was funded by BBSRC grants awarded to J.M.H. and O.P. (BB/N006569/1), C.R. and J.S (BB/N00602X/1), P.G.L (BB/N005945/1), and I.C.C. (BB/N007239/1).An apparent and common feature of aposematic patterns is that they contain a high level of achromatic (luminance) contrast, for example, many warning signals combine black spots and stripes with a lighter colour such as yellow. However, the potential importance of achromatic contrast, as distinct from colour contrast, in reducing predation has been largely overlooked. Here, using domestic chicks as a model predator, we manipulated the degree of achromatic contrast in warning patterns to test if high luminance contrast in aposematic signals is important for deterring naïve predators. We found that the chicks were less likely to approach and eat prey with high contrast compared to low contrast patterns. These findings suggest that aposematic prey patterns with a high luminance contrast can benefit from increased survival through eliciting unlearned biases in naïve avian predators. Our work also highlights the importance of considering luminance contrast in future work investigating why aposematic patterns take the particular forms that they do.Publisher PDFPeer reviewe

Using preferred fluids and different reward schedules to motivate rhesus macaques (Macaca mulatta) in cognitive tasks

Rhesus macaques (Macaca mulatta) used in behavioural neuroscience are often required to complete cognitively complex tasks, for which a high level of motivation is essential. To induce motivation, researchers may implement fluid-restriction protocols, whereby freely available water is limited, such that fluid can be used as a reward in the laboratory. A variety of different rewards and schedules are used, but there exists a lack of data assessing their effectiveness. In this study, we aimed to quantify fluid preference in rhesus macaques and to use these preferences to compare the motivational quality of different reward schedules: the monkey's previous reward (i.e. the fluid used to reward them in past studies), their new preferred reward, a variable schedule of previous and preferred reward, and a choice between the previous and preferred rewards. We found that it may be possible to reduce the level of restriction if an adequately motivating preferred reward is identified, but that this is dependent on the animal. Each monkey responded differently to both the fluid-preference assessments and to the different reward schedules. As such, monkeys should not be subject to ‘blanket’ protocols but should be assessed individually to maintain adequate scientific data collection at the least severe level of fluid restriction

Negative mood affects the expression of negative but not positive emotions in mice

Whether and to what extent animals experience emotions is crucial for understanding their decisions and behaviour, and underpins a range of scientific fields, including animal behaviour, neuroscience, evolutionary biology and animal welfare science. However, research has predominantly focused on alleviating negative emotions in animals, with the expression of positive emotions left largely unexplored. Therefore, little is known about positive emotions in animals and how their expression is mediated. We used tail handling to induce a negative mood in laboratory mice and found that while being more anxious and depressed increased their expression of a discrete negative emotion (disappointment), meaning that they were less resilient to negative events, their capacity to express a discrete positive emotion (elation) was unaffected relative to control mice. Therefore, we show not only that mice have discrete positive emotions, but that they do so regardless of their current mood state. Our findings are the first to suggest that the expression of discrete positive and negative emotions in animals is not equally affected by long-term mood state. Our results also demonstrate that repeated negative events can have a cumulative effect to reduce resilience in laboratory animals, which has significant implications for animal welfare

The environmental setting of Epipalaeolithic aggregation site Kharaneh IV

The archaeological site of Kharaneh IV in Jordan's Azraq Basin, and its relatively near neighbour Jilat 6 show evidence of sustained occupation of substantial size through the Early to Middle Epipalaeolithic (c. 24,000 - 15,000 cal BP). Here we review the geomorphological evidence for the environmental setting in which Kharaneh IV was established. The on-site stratigraphy is clearly differentiated from surrounding sediments, marked visually as well as by higher magnetic susceptibility values. Dating and analysis of off-site sediments show that a significant wetland existed at the site prior to and during early site occupation (~ 23,000 - 19,000 BP). This may explain why such a substantial site existed at this location. This wetland dating to the Last Glacial Maximum also provides important information on the palaeoenvironments and potential palaeoclimatic scenarios for today's eastern Jordanian desert, from where such evidence is scarce

Considerations on the use of video playbacks as visual stimuli: The Lisbon workshop consensus

This paper is the consensus of a workshop that critically evaluated the utility and problems of video playbacks as stimuli in studies of visual behavior. We suggest that video playback is probably suitable for studying motion, shape, texture, size, and brightness. Studying color is problematic because video systems are specifically designed for humans. Any difference in color perception must lead to a different color sensation in most animals. Another potentially problematic limitation of video images is that they lack depth cues derived from stereopsis, accommodation, and motion parallax. Nonetheless, when used appropriately, video playback allows an unprecedented range of questions in visual communication to be addressed. It is important to note that most of the potential limitations of video playback are not unique to this technique but are relevant to all studies of visual signaling in animals

Predators' Toxin Burdens Influence Their Strategic Decisions to Eat Toxic Prey

SummaryToxic prey advertise their unprofitability to predators via conspicuous aposematic coloration [1]. It is widely accepted that avoidance learning by naive predators is fundamental in generating selection for aposematism [2, 3] and mimicry [4, 5] (where species share the same aposematic coloration), and consequently this cognitive process underpins current evolutionary theory [5, 6]. However, this is an oversimplistic view of predator cognition and decision making. We show that predators that have learned to avoid chemically defended prey continue to attack defended individuals at levels determined by their current toxin burden. European starlings learned to discriminate between sequentially presented defended and undefended mealworms with different color signals. Once birds had learned to avoid the defended prey at a stable asymptotic level, we experimentally increased their toxin burdens, which reduced the number of defended prey that they ingested in the subsequent trial. This was due to the birds making strategic decisions to ingest defended prey on the basis of their visual signals. Birds are clearly able to learn about the nutritional benefits and defensive costs of eating defended prey, and they regulate their intake according to their current physiological state. This raises new perspectives on the evolution of aposematism, mimicry, and defense chemistry

Frequency-dependent taste-rejection by avian predation may select for defence chemical polymorphisms in aposematic prey

Chemically defended insects advertise their unpalatability to avian predators using conspicuous aposematic coloration that predators learn to avoid. Insects utilize a wide variety of different compounds in their defences, and intraspecific variation in defence chemistry is common. We propose that polymorphisms in insect defence chemicals may be beneficial to insects by increasing survival from avian predators. Birds learn to avoid a colour signal faster when individual prey possesses one of two unpalatable chemicals rather than all prey having the same defence chemical. However, for chemical polymorphisms to evolve within a species, there must be benefits that allow rare chemical morphs to increase in frequency. Using domestic chicks as predators and coloured crumbs for prey, we provide evidence that birds taste and reject proportionally more of the individuals with rare defence chemicals than those with common defence chemicals. This indicates that the way in which birds attack and reject prey could enhance the survival of rare chemical morphs and select for chemical polymorphism in aposematic species. This is the first experiment to demonstrate that predators can directly influence the form taken by prey's chemical defences