Aposematic Warning Research Articles

Selfish herd effects in aggregated caterpillars and their interaction with warning signals.

Larval Lepidoptera gain survival advantages by aggregating, especially when combined with aposematic warning signals, yet reductions in predation risk may not be experienced equally across all group members. Hamilton's selfish herd theory predicts that larvae that surround themselves with their group mates should be at lower risk of predation, and those on the periphery of aggregations experience the greatest risk, yet this has rarely been tested. Here, we expose aggregations of artificial 'caterpillar' targets to predation from free-flying, wild birds to test for marginal predation when all prey are equally accessible and for an interaction between warning coloration and marginal predation. We find that targets nearer the centre of the aggregation survived better than peripheral targets and nearby targets isolated from the group. However, there was no difference in survival between peripheral and isolated targets. We also find that grouped targets survived better than isolated targets when both are aposematic, but not when they are non-signalling. To our knowledge, our data provide the first evidence to suggest that avian predators preferentially target peripheral larvae from aggregations and that prey warning signals enhance predator avoidance of groups.

Colour polymorphism associated with a gene duplication in male wood tiger moths.

Colour is often used as an aposematic warning signal, with predator learning expected to lead to a single colour pattern within a population. However, there are many puzzling cases where aposematic signals are also polymorphic. The wood tiger moth, Arctia plantaginis, displays bright hindwing colours associated with unpalatability, and males have discrete colour morphs which vary in frequency between localities. In Finland, both white and yellow morphs can be found, and these colour morphs also differ in behavioural and life-history traits. Here, we show that male colour is linked to an extra copy of a yellow family gene that is only present in the white morphs. This white-specific duplication, which we name valkea, is highly upregulated during wing development. CRISPR targeting valkea resulted in editing of both valkea and its paralog, yellow-e, and led to the production of yellow wings. We also characterise the pigments responsible for yellow, white, and black colouration, showing that yellow is partly produced by pheomelanins, while black is dopamine-derived eumelanin. Our results add to a growing number of studies on the genetic architecture of complex and seemingly paradoxical polymorphisms, and the role of gene duplications and structural variation in adaptive evolution.

Hidden black and yellow thigh color acts as an aposematic signal in the Eastern Gray Treefrog (Hyla versicolor)

AbstractRecognizing form and function of animal defenses is paramount to understanding the ecological and evolutionary forces behind predator and prey dynamics. Color patterns are strongly related to defensive strategies in animals. Some rely on camouflage to avoid detection, while others are brightly colored and conspicuously signal their noxiousness to potential predators. Still others combine cryptic dorsal coloration with colorful patches that are concealed in resting position but are facultatively unveiled by special behavior or simply during activity. Such hidden conspicuous color patches may be an intermediate stage in the evolution from camouflage to aposematism. Many species, especially treefrogs of the family Hylidae, are generally considered to be cryptic, yet show colorful patches on their flanks or thighs and have defensive skin secretions that may have unpalatable properties. We investigated whether the conspicuous black and yellow spotted pattern on the thighs of Eastern Gray Treefrogs (Hyla versicolor) serve as an aposematic signal. We conducted a palatability study to test whether Gray Treefrog skin secretions are unpalatable, and a clay model field study to test whether the conspicuous black and yellow thigh pattern is an aposematic warning coloration that reduces predation. Frog secretions were discriminated against during palatability assays, and clay models painted with a spotted black and yellow pattern suffered lower predation rates. Our findings support the hypothesis that hidden color patches act as aposematic signals in Gray Treefrogs and suggest that more species benefit from aposematic coloration than currently appreciated.

Mode of action, chemistry and defensive efficacy of the osmeterium in the caterpillar Battus polydamas archidamas

Chemical secretions are one of the main defensive mechanisms in insects. The osmeterium is a unique organ in larvae of Papilionidae (Lepidoptera), which is everted upon disturbance, secreting odoriferous volatiles. Here, using larvae of the specialized butterfly Battus polydamas archidamas (Papilionidae: Troidini), we aimed to understand the mode of action of the osmeterium, the chemical composition and origin of the secretion, as well as its defensive efficiency against a natural predator. We described osmeterium’s morphology, ultramorphology, structure, ultrastructure, and chemistry. Additionally, behavioral assays of the osmeterial secretion against a predator were developed. We showed that the osmeterium is composed of tubular arms (made up by epidermal cells) and of two ellipsoid glands, which possess a secretory function. The eversion and retraction of the osmeterium are dependent on the internal pressure generated by the hemolymph, and by longitudinal muscles that connect the abdomen with the apex of the osmeterium. Germacrene A was the main compound present in the secretion. Minor monoterpenes (sabinene and ß-pinene) and sesquiterpenes ((E)-β-caryophyllene, selina-3,7(11)-diene, and other some unidentified compounds) were also detected. Only sesquiterpenes (with the exception of (E)-β-caryophyllene) are likely to be synthesized in the osmeterium-associated glands. Furthermore, the osmeterial secretion proved to deter predatory ants. Our results suggest that the osmeterium, besides serving as an aposematic warning for enemies, is an efficient chemical defense, with its own synthesis of irritant volatiles.

Social information use about novel aposematic prey depends on the intensity of the observed cue

Abstract Animals gather social information by observing the behavior of others, but how the intensity of observed cues influences decision-making is rarely investigated. This is crucial for understanding how social information influences ecological and evolutionary dynamics. For example, observing a predator’s distaste of unpalatable prey can reduce predation by naïve birds, and help explain the evolution and maintenance of aposematic warning signals. However, previous studies have only used demonstrators that responded vigorously, showing intense beak-wiping after tasting prey. Therefore, here we conducted an experiment with blue tits (Cyanistes caeruleus) informed by variation in predator responses. First, we found that the response to unpalatable food varies greatly, with only few individuals performing intensive beak-wiping. We then tested how the intensity of beak-wiping influences observers’ foraging choices using video-playback of a conspecific tasting a novel conspicuous prey item. Observers were provided social information from 1) no distaste response, 2) a weak distaste response, or 3) a strong distaste response, and were then allowed to forage on evolutionarily novel (artificial) prey. Consistent with previous studies, we found that birds consumed fewer aposematic prey after seeing a strong distaste response, however, a weak response did not influence foraging choices. Our results suggest that while beak-wiping is a salient cue, its information content may vary with cue intensity. Furthermore, the number of potential demonstrators in the predator population might be lower than previously thought, although determining how this influences social transmission of avoidance in the wild will require uncovering the effects of intermediate cue salience.

Seasonal variation of mortality, detectability, and body condition in a population of the adder (Vipera berus).

We analyzed seasonal variation in mortality rates in adult males and females of the European adder (Vipera berus), using data collected during a 13‐year capture–recapture study (2005–2017) in a large population. We concurrently obtained quantitative information on the seasonal variation in the detectability and body condition of adders. Our results show strong seasonality in body condition, encounter, and capture rates of adult adders, and these patterns differ markedly between sexes and between breeding and nonbreeding females. Seasonal variation in mortality rates was however virtually nonexistent in males and moderately low in both breeding and nonbreeding females. In addition, we found no evidence for among‐year differences in the seasonal mortality schedules of males and females. During periods of intensive basking, both males and pregnant females are highly visible for humans, but are not subject to strong natural mortality. This low susceptibility to predation is presumably induced by various factors, including the limitation of overt exposure to short periods of time and specific microhabitats, the dorsal coloration pattern that provides cryptic protection and possibly also an aposematic warning signal, and presumed seasonal differences in the foraging behavior and food requirements of natural predators. Our data provide some evidence that female adders, but not males, are relatively vulnerable to predation during the seasonal migrations between the hibernation and feeding habitats. Mortality in the females was not much elevated during their breeding years, but was notably highest in the spring of the ensuing nonbreeding year. After giving birth, reproductive females are extremely emaciated and have a weakened general condition. They then run the risk of dying from starvation either before, during, or after hibernation. The higher mortality after giving birth, that is sustained over a period of ca. 9 months, should be considered as an indirect and delayed survival cost of reproduction.

The fitness effects of a pale mutant in the aposematic seed bug Lygaeus simulans indicate pleiotropy between warning coloration and life history.

Conspicuous warning colors that signal chemical or other defenses are common in the natural world. For instance, such aposematic warning patterns of red‐and‐black or yellow‐and‐black are common among insect taxa, particularly in the order Hemiptera, often forming the basis of Batesian and/or Müllerian mimicry rings. In addition, it has been repeatedly noted that color polymorphisms or mutants that influence pigmentation can show pleiotropy with other behavioral, physiological, or life‐history traits. Here, we describe a pale mutant of the seed bug Lygaeus simulans that appeared in our laboratory population in 2012, which differs in color to the wild‐type bugs. Through multigenerational experimental crosses between wild‐type and pale mutant L. simulans, we first show that the pale phenotype segregates as a single Mendelian locus, with the pale allele being recessive to the wild type. Next, we show (a) that there is a large heterozygous advantage in terms of fecundity, (b) that pale females suffer reduced longevity, and (c) that pale males have increased body length compared to wild‐type homozygotes. Our data therefore suggest that the color locus is pleiotropic with a number of life‐history traits, opening the door for a more complete genetic analysis of aposematic coloration in this species. In addition, this phenotype will be useful as a visible genetic marker, providing a tool for investigating sperm competition and other post‐copulatory drivers of sexual selection in this species.

Diversity in warning coloration: selective paradox or the norm?

ABSTRACTAposematic theory has historically predicted that predators should select for warning signals to converge on a single form, as a result of frequency‐dependent learning. However, widespread variation in warning signals is observed across closely related species, populations and, most problematically for evolutionary biologists, among individuals in the same population. Recent research has yielded an increased awareness of this diversity, challenging the paradigm of signal monomorphy in aposematic animals. Here we provide a comprehensive synthesis of these disparate lines of investigation, identifying within them three broad classes of explanation for variation in aposematic warning signals: genetic mechanisms, differences among predators and predator behaviour, and alternative selection pressures upon the signal. The mechanisms producing warning coloration are also important. Detailed studies of the genetic basis of warning signals in some species, most notably Heliconius butterflies, are beginning to shed light on the genetic architecture facilitating or limiting key processes such as the evolution and maintenance of polymorphisms, hybridisation, and speciation. Work on predator behaviour is changing our perception of the predator community as a single homogenous selective agent, emphasising the dynamic nature of predator–prey interactions. Predator variability in a range of factors (e.g. perceptual abilities, tolerance to chemical defences, and individual motivation), suggests that the role of predators is more complicated than previously appreciated. With complex selection regimes at work, polytypisms and polymorphisms may even occur in Müllerian mimicry systems. Meanwhile, phenotypes are often multifunctional, and thus subject to additional biotic and abiotic selection pressures. Some of these selective pressures, primarily sexual selection and thermoregulation, have received considerable attention, while others, such as disease risk and parental effects, offer promising avenues to explore. As well as reviewing the existing evidence from both empirical studies and theoretical modelling, we highlight hypotheses that could benefit from further investigation in aposematic species. Finally by collating known instances of variation in warning signals, we provide a valuable resource for understanding the taxonomic spread of diversity in aposematic signalling and with which to direct future research. A greater appreciation of the extent of variation in aposematic species, and of the selective pressures and constraints which contribute to this once‐paradoxical phenomenon, yields a new perspective for the field of aposematic signalling.

Different colour morphs of the poison frog Andinobates bombetes (Dendrobatidae) are similarly effective visual predator deterrents

AbstractAposematism is the use of warning signals to advertise unpleasant or dangerous defences to potential predators. As the effectiveness of this strategy depends on predator learning, little variation is expected in aposematic warning signals, as similar signals facilitate predator learning. However, warning signals are frequently variable in aposematic species. Such variability could arise as a result of geographic variation in the interpretation that local predators give warning signals. We tested this divergent learning hypothesis in the polytypic poison frog Andinobates bombetes (Anura: Dendrobatidae), focusing on visual predators. Our study was conducted in two populations of this species located in the Western Andes of Colombia, where individuals at some localities exhibit red dorsolateral stripes, while those in others exhibit yellow dorsolateral stripes. We deployed paraffin models imitating both forms of A. bombetes in size and colouration, as well as dull‐coloured controls, at sites inhabited by either red‐striped or yellow‐striped frogs. Red and yellow models were attacked at similar rates at both sites, and brown models were attacked more frequently at one of the sites. These results suggest that red and yellow colourations function as similarly effective aposematic signals for primarily visual predators, regardless of the form previously experienced by these predators. Therefore, our results do not support the hypothesis of divergent predator learning as a driver of the polytypism present in this species. Finally, we discuss other mechanisms that may be involved in the evolution and maintenance of this polytypism.

Bioluminescence in Dinoflagellates: Evidence that the Adaptive Value of Bioluminescence in Dinoflagellates is Concentration Dependent.

Three major hypotheses have been proposed to explain why dinoflagellate bioluminescence deters copepod grazing: startle response, aposematic warning, and burglar alarm. These hypotheses propose dinoflagellate bioluminescence (A) startles predatory copepods, (B) warns potential predators of toxicity, and (C) draws the attention of higher order visual predators to the copepod's location. While the burglar alarm is the most commonly accepted hypothesis, it requires a high concentration of bioluminescent dinoflagellates to be effective, meaning the bioluminescence selective advantage at lower, more commonly observed, dinoflagellate concentrations may result from another function (e.g. startle response or aposematic warning). Therefore, a series of experiments was conducted to evaluate copepod grazing (Acartia tonsa) on bioluminescent dinoflagellates (during bioluminescent and nonbioluminescent phases, corresponding to night and day, respectively) at different concentrations (10, 1000, and 3000 cells mL-1 ), on toxic (Pyrodinium bahamense var. bahamense) and nontoxic (Lingulodinium polyedrum) bioluminescent dinoflagellates, and in the presence of nonluminescent diatoms (Thalassiosira eccentrica). Changes in copepod ingestion rates, clearance rates, and feeding preferences as a result of these experimental factors, particularly during the mixed trails with nonluminescent diatoms, indicate there is a concentration threshold at which the burglar alarm becomes effective and below which dinoflagellate bioluminescence functions as an aposematic warning.

Aposematic coloration does not deter corallivory by fish on the coral Montastraea cavernosa

Predation on corals by visual predators is a significant source of partial or total mortality on coral reefs, and corals have evolved strategies, including chemical defenses, to deter predation. One mechanism that organisms use to communicate the presence of chemical defenses is aposematic coloration, or the display of bright coloration as a warning to visual predators such as fish. Corals exhibit multiple colors, and it has been hypothesized that one role for this variability in coloration is as an aposematic warning of adverse palatability. Here, we test green and orange color morphs of the Caribbean coral Montastraea cavernosa for the presence of chemical defenses and whether their differences in coloration elicited different feeding responses. While M. cavernosa is chemically defended, there is no difference in feeding deterrence between color morphs; thus, the different color morphs of this coral species do not appear to represent an example of aposematic coloration.

Two Defensive Lines in Juvenile Leaf Beetles; Esters of 3-nitropropionic Acid in the Hemolymph and Aposematic Warning.

Juveniles of the leaf beetles in subtribe Chrysomelina have efficient defense strategies against predators. When disturbed, they transiently expose volatile deterrents in large droplets from nine pairs of defensive glands on their back. Here, we report on an additional line of defense consisting of the non-volatile isoxazolin-5-one glucoside and its 3-nitropropanoyl ester in the larval hemolymph. Because isoxazolin-5-one derivatives were not detectable in related leaf beetle taxa, they serve as a diagnostic marker for the Chrysomelina subtribe. Conjugation of isotopically labelled 3-nitropropionic acid to isoxazolin-5-one glucoside in vivo demonstrates its function as a carrier for the 3-nitropropanoyl esters. The previous identification of characteristic glucosides as precursors of the volatile deterrents underlines the general importance of glucosides for sequestration from food plants, and the subsequent transport in the hemolymph to the defense system. The combination of repellent volatiles with non-volatile toxic compounds in the hemolymph has the potential to create synergistic effects since the odorant stimulus may help predators learn to avoid some foods. The combination of the two defense lines has the advantage, that the hemolymph toxins provide reliable and durable protection, while the repellents may vary after a host plant change.Electronic supplementary materialThe online version of this article (doi:10.1007/s10886-016-0684-0) contains supplementary material, which is available to authorized users.

Evolutionary losses of facial stripes in New World pitvipers

Associations between the evolutionary loss or gain of colours and habitat can be informative about how environment drives colour evolution, and provide insight into their functions. Despite a wealth of studies on colours, the function of many colour patches are not well understood. This is especially the case for dark facial markings. Dark facial stripes are common in snakes, including many pitvipers, which possess facial stripes that extend from the eye towards the corner of the mouth. We investigated whether the evolutionary loss or gain of facial stripes in New World pitvipers is associated with activity period and habitat, hoping to shed light on their function. First, we examined whether evolutionary loss or gain of facial stripes is associated with nocturnal or diurnal activity. It has been suggested that facial stripes may protect the venom from ultraviolet radiation, which would be higher in diurnal species. Similarly, snakes in open habitats may experience higher ultraviolet radiation than those in closed habitats. Alternatively, we examined whether evolutionary loss or gain of facial stripes is associated with terrestrial or non-terrestrial habitat use. Studies on similar facial markings in other vertebrates indicated that they have a signalling function. If facial stripes in pitvipers have a signalling function, arboreal or saxicolous habitat use may limit signal effectiveness. Hence, we hypothesized that there may be an association between facial stripe evolution and habitat use. Using combined phylogenies that included almost every New World pitviper species, we tested correlations between facial stripes and ultraviolet exposure (activity period and habitat openness) and habitat use (terrestrial or non-terrestrial) using concentrated-changes tests and Pagel’s correlation tests. Our data did not support the hypothesis that ultraviolet exposure influenced facial stripe evolution. Instead, the evolutionary loss of facial stripes was associated with non-terrestrial habitat use, such as arboreal and saxicolous habitats. In these habitats, we suggest the effectiveness of facial stripes as a signal would be limited, leading to their evolutionary loss. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104, 923–933. ADDITIONAL KEYWORDS: aposematic warning – rattlesnakes – signals – ultraviolet radiation.

Bioluminescent aposematism in millipedes

Summary Bioluminescence — the ability of organisms to emit light — has evolved about 40–50 times independently across the tree of life [1]. Many different functions for bioluminescence have been proposed, for example, mate recognition, prey attraction, camouflage, and warning coloration. Millipedes in the genus Motyxia produce a greenish-blue light at a wavelength of 495 nm that can be seen in darkness [2]. These detritivores defend themselves with cyanide, which they generate internally and discharge through lateral ozopores [3]. Motyxia are an ideal model system to investigate the ecological role of bioluminescence because they are blind, thus limiting their visual signalling to other organisms, for example predators. While the biochemical mechanisms underlying Motyxia bioluminescence have been studied in detail [2,4], its adaptive significance remained unknown [5,6]. We here show that bioluminescence has a single evolutionary origin in millipedes and it serves as an aposematic warning signal to deter nocturnal mammalian predators. Among the numerous examples of bioluminescence, this is the first field experiment in any organism to demonstrate that bioluminescence functions as a warning signal. Video Abstract The editors of Current Biology welcome correspondence on any article in the journal, but reserve the right to reduce the length of any letter to be published. All Correspondence containing data or scientific argument will be refereed. Queries about articles for consideration in this format should be sent by e-mail to cbiol@current-biology.com

The role of the proteinase inhibitor ovorubin in apple snail eggs resembles plant embryo defense against predation.

BackgroundFieldwork has thoroughly established that most eggs are intensely predated. Among the few exceptions are the aerial egg clutches from the aquatic snail Pomacea canaliculata which have virtually no predators. Its defenses are advertised by the pigmented ovorubin perivitellin providing a conspicuous reddish coloration. The nature of the defense however, was not clear, except for a screening for defenses that identified a neurotoxic perivitellin with lethal effect on rodents.Ovorubin is a proteinase inhibitor (PI) whose role to protect against pathogens was taken for granted, according to the prevailing assumption. Through biochemical, biophysical and feeding experiments we studied the proteinase inhibitor function of ovorubin in egg defenses.Methodology/Principal FindingsMass spectrometry sequencing indicated ovorubin belongs to the Kunitz-type serine proteinase inhibitor family. It specifically binds trypsin as determined by small angle X-ray scattering (SAXS) and cross-linking studies but, in contrast to the classical assumption, it does not prevent bacterial growth. Ovorubin was found extremely resistant to in vitro gastrointestinal proteolysis. Moreover feeding studies showed that ovorubin ingestion diminishes growth rate in rats indicating that this highly stable PI is capable of surviving passage through the gastrointestinal tract in a biologically active form.ConclusionsTo our knowledge, this is the first direct evidence of the interaction of an egg PI with a digestive protease of potential predators, limiting predator's ability to digest egg nutrients. This role has not been reported in the animal kingdom but it is similar to plant defenses against herbivory. Further, this would be the only defense model with no trade-offs between conspicuousness and noxiousness by encoding into the same molecule both the aposematic warning signal and an antinutritive/antidigestive defense. These defenses, combined with a neurotoxin and probably unpalatable factors would explain the near absence of predators, opening new perspectives in the study of the evolution and ecology of egg defensive strategies.

Familiarity breeds contempt: effects of striped skunk color, shape, and abundance on wild carnivore behavior

Multicomponent aposematic warning signals are generally accepted to function as a deterrent to predatory attacks; however, the relative importance of specific visual cues used by wild predators to recognize defended animals is poorly understood. Here, I use naturally and reciprocally colored taxidermy models of striped skunks (Mephitis mephitis) and gray foxes (Urocyon cinereoargenteus) to explore the impact of aposematic coloration and body shape on the behavior of wild mammalian predators. Models were baited, and all visiting species were recorded with remote video cameras at 10 wilderness sites in California, USA. I found that mammalian predators not only approached black-and-white models more hesitantly than gray-colored models but also reacted negatively to skunk-shaped models, suggesting that predators may generalize noxious qualities to both skunk coloration and skunk shape. This study also found an effect of skunk abundance on animal behavior with predators less likely to visit black-and-white models and more hesitant in their approach to skunk-shaped models at sites with greater skunk abundance. These findings suggest that prior experience and the frequency with which aposematic prey are encountered in a landscape are important predictors of predator avoidance of defended prey.

What is going on between aposematic carabid beetles? The case of Anchomenus dorsalis (Pontoppidan 1763) and Brachinus sclopeta (Fabricius 1792) (Coleoptera Carabidae)

Interspecific aggregations among insects are quite common, but not among carabid beetles. Moreover, aggregations of aposematic species, and morphologically similar beetles such as Anchomenus dorsalis and Brachinus sclopeta are even rarer. Most interestingly, in these aggregations, we observed a peculiar rubbing behaviour of A. dorsalis towards B. sclopeta. In this study we aimed (i) to verify if A. dorsalis displays the rubbing behaviour towards non-aposematic species, (ii) to describe in detail and to quantify this behaviour between A. dorsalis and B. sclopeta, and (iii) to verify if previous experience of inter-specific gregariousness with the same species may exert some effects on the frequency of the rubbing display. We have investigated the qualitative and quantitative characteristics of rubbing behaviour towards aposematic and non-aposematic carabide beetle species, and then we used A. dorsalis and B. sclopeta as model species. We tested the rubbing of A. dorsalis towards B. sclopeta with either solitary A. dorsalis or in A. dorsalis coming from monospecific (A. dorsalis) or from heterospecific (A. dorsalis and B. sclopeta) aggregations. We finally tested the effectiveness of the B. sclopeta cuticular odour in eliciting rubbing from A. dorsalis by presenting olfactory dummies (special paper) either soaked with water or after it had previously been placed in a monospecific group of B. sclopeta for 15 days. First of all, we found important differences in the occurrence of rubbing behaviour towards aposematic and non-aposematic species, as A. dorsalis displayed the rubbing only towards the former. Secondly, rubbing frequency depended on the pre-test conditions (isolated, mono- or heterospecific aggregations) of the tested A. dorsalis individuals, as no rubbing occurred in individuals coming from heterospecific aggregations. The paper presentation tests demonstrated that the odour of B. sclopeta was sufficient to provoke rubbing in A. dorsalis. We conclude that aggregation in these species of carabid beetles may influence their anti-predatory defences possibly increasing the effectiveness of the aposematic (warning) signals. More work is needed to delineate in detail the evolutionary significance of this behaviour, focusing on the possibility that interspecific aggregations may not be so rare in ground beetles, and that this positive interaction could involve a larger number of species and genera, at least in wet, sun-exposed and open land habitat types.

Signals of profitability? Food colour preferences in migrating juvenile blackcaps differ for fruits and insects

Red is a common colour signal in both aposematic warning displays, and in fruit displays. One common feature is that red is conspicuous against the natural background of the prey and fruits. However, there is a potential conflict between fruits and aposematic prey in how a bird predator should react to red colours, where fruits aim to attract birds and aposematic insects aim to ward off, often the same bird individuals. Here we investigate possible differences in red/green colour preferences of frugivorous, wild-caught, young blackcaps (Sylvia atricapilla), when food is either a fruit or an insect. Birds in two groups were presented with a series of pairs of food items that had been artificially painted red and green, in the order of (I) fruits, crickets and maggots, or (II) crickets, fruits, and maggots. Birds first presented with crickets or fruits differed in first attacks directed at the two colours: They showed no colour preference between fruits, but showed a clear preference for green over red crickets. Also, birds in both experimental groups clearly preferred green to red maggots. These results provide evidence that wild, frugivorous birds are able to differentiate between prey types, and show different colour preferences depending on whether food is insect or fruit. We conclude that blackcaps show an attack bias against red insects, and that one important function of the signal in insects, is to inhibit attack after discovery. However, the lack of preference for red fruits suggests other functions to red fruit displays, such as facilitating discovery per se, rather than directly stimulating attack after discovery.

The function of facial masks in "midguild" carnivores

A group of medium sized carnivores possesses conspicuously colored facial markings or masks. This facial coloration is most compatible with the aposematic warning hypothesis and functions to deter predation by larger carnivores. Other hypotheses addressing ecological or social functions are not supported by the data. Facial masks may be a recent character derived from an ancestral contrast in body coloration of carnivores that functioned to warn potential predators of a disproportionate danger in attacking the smaller guild member. This danger may be predicated either on the relative ferocity of the smaller carnivore (wolverine/badger‐type) or that the smaller carnivore is equipped with a defensive nauseous discharge that is directly harmful to larger would‐be predators. Facial masks in themselves appear to function as an aposematic warning of aggressive defensive behavior that could be harmful to predators.

Hissing in bumblebees: an interspecific defence signal

Bumblebees (Bombus terrestris) react to disturbances within the nest by a conspicuous hissing sound. The sound is characterised by a high intensity in the ultrasonic frequency range. It is elicited by vibrations of the nest and by mammalian breath and artificial air currents containing CO2. Domestic mice entering a bumblebee nest elicit these sounds and retreat immediately in response to the bumblebees' signal. It is concluded that the hissing sounds serve as aposematic warning signals aimed at predators entering the nest.