Testing the predatory behaviour of Podarcis sicula (Reptilia: Lacertidae) towards aposematic and non-aposematic preys

Bonacci, Teresa, Gaetano Aloise, Pietro Brandmayr, Tullia Z. Brandmayr and Massimo Capula. 2008. Testing the predatory behaviour of Podarcis sicula (Reptilia: Lacertidae) towards aposematic and non-aposematic preys. Amphibia- Reptilia. 29 (3): 449-453. [t.bonacci@unical.it]

Abstract: Food preferences and the effects of prey chemical repellents in the dietary behaviour of Podarcis sicula were tested using four species of Carabid beetles as prey models. The goal of the study was to assess (i) the ability of P. sicula to recognize insect preys provided with chemical repellents and aposematic colorations under laboratory conditions, and (ii) the importance of chemical signals used by the prey model as antipredatory strategy. Preys used in this study were Brachinus sclopeta and Anchomenus dorsalis (aposematic species) and Amara anthobia and A. aenea (non-aposematic species). Aposematic species are characterized by warning color pattern and by production of chemical repellents, while non-aposematic ones do not. Amara anthobia and A. aenea were attacked with high frequency by P. sicula, Brachinus sclopeta and Anchomenus dorsalis with low frequency. Non-aposematic species were preyed more often than the aposematic ones. Brachinus sclopeta was preyed after low latency, while Amara anthobia and A. aenea after long latency. Non-aposematic species were captured and eaten without difficulty, while when B. sclopeta or A. dorsalis were captured, lizards always tossed their head and then rub the snout on the soil, probably because of the unpalatability of aposematic preys.

Optimal-Foraging Predator Favors Commensalistic Batesian Mimicry

Optimal-Foraging Predator Favors Commensalistic Batesian Mimicry

Atsushi Honma1¤*, Koh-ichi Takakura2, Takayoshi Nishida1

1 Laboratory of Insect Ecology, Graduate School of Agriculture, Kyoto University, Kyoto, Japan, 2 Osaka City Institute of Public Health and Environmental Science, Osaka, Japan

Background

Mimicry, in which one prey species (the Mimic) imitates the aposematic signals of another prey (the Model) to deceive their predators, has attracted the general interest of evolutionary biologists. Predator psychology, especially how the predator learns and forgets, has recently been recognized as an important factor in a predator–prey system. This idea is supported by both theoretical and experimental evidence, but is also the source of a good deal of controversy because of its novel prediction that in a Model/Mimic relationship even a moderately unpalatable Mimic increases the risk of the Model (quasi-Batesian mimicry).

Methodology/Principal Findings

We developed a psychology-based Monte Carlo model simulation of mimicry that incorporates a “Pavlovian” predator that practices an optimal foraging strategy, and examined how various ecological and psychological factors affect the relationships between a Model prey species and its Mimic. The behavior of the predator in our model is consistent with that reported by experimental studies, but our simulation's predictions differed markedly from those of previous models of mimicry because a more abundant Mimic did not increase the predation risk of the Model when alternative prey were abundant. Moreover, a quasi-Batesian relationship emerges only when no or very few alternative prey items were available. Therefore, the availability of alternative prey rather than the precise method of predator learning critically determines the relationship between Model and Mimic. Moreover, the predation risk to the Model and Mimic is determined by the absolute density of the Model rather than by its density relative to that of the Mimic.

Conclusions/Significance

Although these predictions are counterintuitive, they can explain various kinds of data that have been offered in support of competitive theories. Our model results suggest that to understand mimicry in nature it is important to consider the likely presence of alternative prey and the possibility that predation pressure is not constant.

[Article]Being conspicuous and defended: selective benefits for the individual

Being conspicuous and defended: selective benefits for the individual

Christina G. Halpin, John Skelhorn and Candy Rowe

Centre for Behaviour and Evolution, Institute of Neuroscience, Newcastle University, Henry Wellcome Building, Framlington Place, Newcastle upon Tyne, NE2 4HH, UK

Aposematic insects conspicuously advertise their unprofitability to potential predators. However, when these prey initially evolved, they were likely to have been rare and presumably at a greater risk of being detected and killed by naive predators. Both kin and individual selection theories have been used in attempts to explain this apparent paradox, with much of the empirical research supporting kin selection–based theories. Here, we experimentally test how chemical defence levels in prey and avian color biases influence the probability of a rare conspicuous morph having an initial survival advantage. We used newly hatched domestic chicks (Gallus gallus domesticus) foraging on green and purple prey, on a green or purple background, to model the evolutionary scenario of a rare conspicuous morph arising in a population of already defended cryptic prey. Defended prey were produced by spraying them with quinine solution, which the birds readily detect and can learn to avoid. Although attack rates were initially similar for both defended prey types, the chicks only learned to avoid defended prey when they were conspicuous, not when they were cryptic. In addition, defended conspicuous prey were more likely to be rejected on attack than defended cryptic prey, even when first encountered by a predator. These data suggest that there could be a selective advantage for a rare conspicuous morph to arise in a population of cryptic defended prey due to increased avoidance learning and taste-rejection in naive predators. Our findings also suggest that being a non-preferred color and/or highly defended will increase the probability of this evolutionary scenario.

keywords: aposematism, avoidance learning, color bias, predation, receiver psychology, taste-rejection, warning signal.

[Article]Dazzle coloration and prey movement

Dazzle coloration and prey movement

Martin Stevens1, Daniella H. Yule1, Graeme D. Ruxton2

1Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK
2Faculty of Biomedical and Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK

Many traits in animals reduce the rate of attack from visually hunting predators, including camouflage, warning signals and mimicry. In addition, some animal markings may reduce the likelihood that an attack ends in successful capture. These might include dazzle markings, high-contrast patterns that make the estimation of speed and trajectory difficult. However, until now, no study has experimentally tested whether some markings may achieve such an effect. We developed a computer ‘game’ where human ‘predators’ have to capture computer-generated prey moving across a background. In two experiments, we find that although uniform camouflaged targets were among the hardest to capture, so were a range of high-contrast conspicuous patterns, such as bands and zigzags. Prey were also more difficult to capture against more heterogeneous than uniform backgrounds, and at faster speeds of movement. As such, we find the first experimental evidence that conspicuous patterns, similar to those found in a wide range of real animals, make the capture of moving prey more challenging. Various anti-predator markings may work prey during motion, and some animals may combine such dazzle patterns with other functions, such as camouflage, thermoregulation, sexual and warning signals.

keywords: protective coloration, motion, conspicuousness, vision, predation, dazzle

[Artical]Can't tell the caterpillars from the trees: countershading enhances survival in a woodland

Can't tell the caterpillars from the trees: countershading enhances survival in a woodland

Hannah M. Rowland1, Innes C. Cuthill2, Ian F. Harvey1, Michael P. Speed1, Graeme D. Ruxton3

1 School of Biological Sciences, University of Liverpool, Biosciences Building, Crown Street, Liverpool L69 7ZB, UK
2 School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 1UG, UK
3Division of Environmental and Evolutionary Biology, Institute of Biomedical and Life Sciences, University of Glasgow, Graham Kerr Building, Glasgow G12 8QQ, UK

Perception of the body's outline and three-dimensional shape arises from visual cues such as shading, contour, perspective and texture. When a uniformly coloured prey animal is illuminated from above by sunlight, a shadow may be cast on the body, generating a brightness contrast between the dorsal and ventral surfaces. For animals such as caterpillars, which live among flat leaves, a difference in reflectance over the body surface may degrade the degree of background matching and provide cues to shape from shading. This may make otherwise cryptic prey more conspicuous to visually hunting predators. Cryptically coloured prey are expected to match their substrate in colour, pattern and texture (though disruptive patterning is an exception), but they may also abolish self-shadowing and therefore either reduce shape cues or maintain their degree of background matching through countershading: a gradation of pigment on the body of an animal so that the surface closest to illumination is darker. In this study, we report the results from a series of field experiments where artificial prey resembling lepidopteran larvae were presented on the upper surfaces of beech tree branches so that they could be viewed by free-living birds. We demonstrate that countershading is superior to uniform coloration in terms of reducing attack by free-living predators. This result persisted even when we fixed prey to the underside of branches, simulating the resting position of many tree-living caterpillars. Our experiments provide the first demonstration, in an ecologically valid visual context, that shadowing on bodies (such as lepidopteran larvae) provides cues that visually hunting predators use to detect potential prey species, and that countershading counterbalances shadowing to enhance cryptic protection.

keywords: countershading, crypsis, predation, animal coloration, defensive coloration

[Article]Learning and the mimicry spectrum: from quasi-Bates to super-Müller

Learning and the mimicry spectrum: from quasi-Bates to super-Müller

Alexandra C.V. Balogh^{, a,} , Gabriella Gamberale-Stille^a and Olof Leimar^a

^aDepartment of Zoology, Stockholm University, Sweden

Müllerian mimicry is the mutualistic resemblance between two defended species, while Batesian mimicry is the parasitic resemblance between a palatable species (the mimic) and an unpalatable one (the model). These two kinds of mimicry are traditionally seen as extreme ends of a mimicry spectrum. For the range in between, it has been suggested that mimetic relations between unequally defended species could be parasitic, and this phenomenon has been referred to as quasi-Batesian mimicry. Where a mimetic relation is placed along the mimicry spectrum depends on the assumptions made about predator learning. We used a variant of the Rescorla–Wagner learning model for virtual predators to analyse the different possible components of the mimicry spectrum. Our model entails that the rate of associative learning is influenced by variation in the stimuli to be learned. Variable stimuli, that is, unequal defences, can increase the predator learning rate and thus lead to an increased level of mutualism in a mimetic relation. In our analysis, we made use of the concepts of super-Müllerian mimicry, where the benefit of mimicry is even greater than in traditional Müllerian mimicry, and quasi-Müllerian mimicry, where mimicry by a palatable mimic is mutualistic. We suggest that these types of mimicry should be included in the mimicry spectrum along with Müllerian, Batesian and quasi-Batesian mimicry.

Keywords: associative learning; mutualism; Müllerian mimicry; quasi-Batesian mimicry; Rescorla–Wagner model

[Article]Multimodal warning signals for a multiple predator world

Nature 455, 96-99 (4 September 2008) | doi:10.1038/nature07087; Received 10 February 2008; Accepted 14 May 2008

Multimodal warning signals for a multiple predator world

John M. Ratcliffe^1,3 & Marie L. Nydam^2,3

1. Center for Sound Communication, Institute of Biology, University of Southern Denmark, DK-5230 Odense M, Denmark
2. Department of Ecology and Evolutionary Biology, Cornell University, Ithaca, New York 14853, USA
3. These authors contributed equally to this work.

Correspondence to: John M. Ratcliffe^1,3Marie L. Nydam^2,3 Correspondence and requests for materials should be addressed to M.L.N. (Email: mln32@cornell.edu) or J.M.R. (Email: jmr@biology.sdu.dk).

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Abstract

Aposematism is an anti-predator defence, dependent on a predator's ability to associate unprofitable prey with a prey-borne signal¹. Multimodal signals should vary in efficacy according to the sensory systems of different predators; however, until now, the impact of multiple predator classes on the evolution of these signals had not been investigated^{2, 3}. Here, using a community-level molecular phylogeny to generate phylogenetically independent contrasts, we show that warning signals of tiger moths vary according to the seasonal and daily activity patterns of birds and bats—predators with divergent sensory capacities. Many tiger moths advertise chemical defence^{4, 5} using conspicuous colouration and/or ultrasonic clicks^{3, 6}. During spring, when birds are active and bats less so, we found that tiger moths did not produce ultrasonic clicks. Throughout both spring and summer, tiger moths most active during the day were visually conspicuous. Those species emerging later in the season produced ultrasonic clicks; those that were most nocturnal were visually cryptic. Our results indicate that selective pressures from multiple predator classes have distinct roles in the evolution of multimodal warning displays now effective against a single predator class. We also suggest that the evolution of acoustic warning signals may lack the theoretical difficulties associated with the origination of conspicuous colouration.