Mimetic Signals Research Articles

Propensity of Predator Mimicry in Steller’s Jays

Avian vocal mimicry has been described in a variety of contexts, suggesting its function is multifaceted within and across species; however, basic empirical data describing mimetic signal prevalence and context are lacking for numerous species. We examined the occurrence and context of mimicked Red-shouldered Hawk (Buteo lineatus) calls over a 12-month period in a population of 49 individually color-marked Steller’s Jays (Cyanocitta stelleri). We documented mimicry of Red-shouldered Hawk calls in 14 of 49 (28.6%) jays during this 12-month period. We also reviewed the occurrence of the behavior in historic observation data. Hawk mimicry occurred more often during the early breeding season when jays were within home territories, their mates were present, and aggression was absent. Younger, larger, and bolder jays were most likely to perform imitations. These results suggest jays individually vary in mimetic propensity, and individuals’ proclivity for mimicry may be influenced by social and ecological contexts, physical characteristics, and personality traits.

Spectral variation and pigmentary basis of ornamental and mimetic wing colour patches of swallowtail butterflies

Abstract Colours and colour patterns are diverse traits that are often used as visual signals. To test ecological and evolutionary drivers of these visual signals, a clear understanding of their nature and variation is necessary. Here we characterize variation in wing coloration of mormon swallowtail butterflies (Papilio, subgenus Menelaides). These species exhibit two kinds of colour patches on black wings: creamy white/yellow/green patches that are presumably used as sexual ornaments, and pure white patches that are presumably used as mimetic signals. Using reflectance spectrophotometry we quantified spectral properties of black wing background and colour patches between sexes, wing surfaces, and mimicry status. We found that brightness and saturation of the black background were less variable across sexes, wing surfaces, and mimetic/nonmimetic status. However, colour contrast and saturation were higher on dorsal surfaces than on ventral surfaces, and colour contrast between the black background and colour patches was higher in males than in females. Colour patches in nonmimetic butterflies were brighter and more saturated across the whole colour spectrum compared to mimetic butterflies. Thus, our study characterizes colour variation in an extraordinary signal radiation on the wings of swallowtail butterflies, a model clade in ecology, evolution, and genetics. Our results generate numerous hypotheses about the ecological and sexual functions of these colour patches, which may guide further behavioural experiments. Finally, using liquid chromatography-mass spectrometry we identified the pigment papiliochrome-II to be the chemical basis of the presumed sexual ornaments in mormon swallowtails.

Unexpected colour pattern variation in mimetic frogs: implication for the diversification of warning signals in the genus Ranitomeya.

Predation is expected to promote uniformity in the warning coloration of defended prey, but also mimicry convergence between aposematic species. Despite selection constraining both colour-pattern and population divergence, many aposematic animals display numerous geographically structured populations with distinct warning signal. Here, we explore the extent of phenotypic variation of sympatric species of Ranitomeya poison frogs and test for theoretical expectations on variation and convergence in mimetic signals. We demonstrate that both warning signal and mimetic convergence are highly variable and are negatively correlated: some localities display high variability and no mimicry while in others the phenotype is fixed and mimicry is perfect. Moreover, variation in warning signals is always present within localities, and in many cases this variation overlaps between populations, such that variation is continuous. Finally, we show that coloration is consistently the least variable element and is likely of greater importance for predator avoidance compared to patterning. We discuss the implications of our results in the context of warning signal diversification and suggest that, like many other locally adapted traits, a combination of standing genetic variation and founding effect might be sufficient to enable divergence in colour pattern.

Chemical aposematism: the potential for non‐host odours in avian defence

Many birds display plumage coloration and patterns that act as visual aposematic and mimetic signals for predators. Avian aposematism can also be communicated via chemical cues, where so‐called non‐host odours (NHO) emitted from the plumage, the uropygial (preen) gland or other chemical sources denote the status of birds as dangerous or unprofitable hosts for ectoparasitic and haematophagous arthropods. Avian NHO are aposematic signals that in natural concentrations repel nuisance arthropods or inhibit their attraction to hosts, as opposed to being generally unpalatable or toxic to predators. Birds that emit NHO preempt encounters with nuisance arthropods and the pathogens they transmit. Nuisance arthropods that respond to NHO avoid birds that exhibit disabling behavioural, anatomical, physiological and/or chemical defences. As with distinctive plumage patterns that evolved as visual warnings to predators, birds are selected to advertise their non‐host status to nuisance arthropods via distinctive semiochemicals. Short‐chain (C6–C10) aldehydes emitted from the plumage of some birds may act as NHO against a variety of nuisance arthropods. In some birds, the size of the uropygial gland (and presumably its secretory output) is negatively correlated with the degree of ectoparasite infestation or the incidence of malarial infection, thus implicating uropygial secretions in defence against dipteran vectors of haemosporidian pathogens. A field experiment demonstrates that volatiles emanating from the uropygial gland of the Eurasian Hoopoe Upupa epops deter some biting flies. Birds' risky habit of plucking hair from live mammals and incorporating the stolen hair tufts into their nests is hypothesized to serve as a mechanism of appropriating NHO from mammalian skin secretions for nest defence against arthropods. Feathers plucked from heterospecifics and shed snake skins that are incorporated into nests also may emit NHO. Behavioural assays measuring the repellence or attraction‐inhibition of arthropods are needed to elucidate avian NHO. Experimental efforts to elucidate the underlying defences that support avian NHO entail challenges that include the enactment of defences that are variously effective against different arthropods, and the emission of chemical signals that are variously ‘understood’, i.e. avoided, by different arthropods.

Selective alarm call mimicry in the sexual display of the male superb lyrebird (Menura novaehollandiae)

Despite much research on mimicry, little is known about the ecology of dynamic mimetic signals involving mimicry of multiple species. Some of the most conspicuous examples of phenotypically plastic mimicry are produced by oscine passerines, where vocal production learning enables some species to mimic multiple models and flexibly adjust what they mimic and when. While singing from a perch, male superb lyrebirds (Menura novaehollandiae) accurately imitate multiple songs and calls of over 20 species of bird. However, at key moments within their multimodal displays performed on display arenas on the forest floor, males mimic a small number of mobbing-alarm calls creating the acoustic illusion of a mixed-species mobbing flock (‘D-song’). Using observations from camera footage and a field-based playback experiment, we tested six hypotheses for alarm call model selection within D-song. Mimicked species were remarkably invariant, with 79% of D-song made up of imitations of just three different bird species. Males did not mimic the most common species in their general environment, but neither did they mimic rare species. Instead, males imitated the mobbing-alarm calls of heterospecific birds that foraged on or near the forest floor. Indeed, males primarily mimicked the alarm calls of heterospecific species that foraged alongside lyrebirds and were likely to appear together in experimentally-induced, terrestrial mobbing flocks. These findings support the hypothesis that males mimic a cue of a terrestrial predatory threat to lyrebirds, most likely to exploit the antipredator behaviour of female lyrebirds. Our study illustrates the importance of investigating the drivers of model selection in dynamic multi-model mimicry.

Predator-based selection and the impact of edge sympatry on components of coral snake mimicry

Mimicry is a vivid example of how predator-driven selection can impact phenotypic diversity, which itself can be influenced by the presence (sympatry) or absence (allopatry) of a dangerous model. However, the impact of sympatry and allopatry on predation on mimicry systems at fine spatial scales (e.g., edge sympatry, allopatry) is not well understood. We studied mimicry in a montane tropical site in Honduras with an abrupt elevational gradient where coral snakes and mimics exist in close sympatry and allopatry. Specifically, we used clay replicas to test the impact of edge sympatry on (1) overall attack rates, (2) the fitness benefit of mimetic coloration, and (3) predation on specific mimetic signal components. Unlike previous research, we found that mimetic phenotypes received significantly more attacks than cryptic replicas in edge sympatry, suggesting that mimetic phenotypes might not confer a fitness benefit in areas of edge sympatry. Additionally, we documented temporal variation in predator-based selection, as the impacts of allopatry on predatory attacks varied among years. Our results imply that the effect of sympatry and allopatry on predator-based selection in mimicry systems may be more complex than previously thought for species-rich assemblies of coral snakes and their mimics in the montane tropics.

Microevolutionary change in mimicry? Potential erosion of rattling behaviour among nonvenomous snakes on islands lacking rattlesnakes

Batesian mimics – harmless species that converge on the warning signals of a dangerous species – are spectacular examples of adaptation, but few documented cases involve acoustic signals. Even fewer studies have documented microevolutionary change in mimicry of any kind. Here, we describe a potential evolutionary change in acoustic mimicry. Many nonvenomous snakes vibrate their tail tip when threatened, making a sound resembling venomous rattlesnakes, which can vibrate their tail rapidly and for a long duration. When we compared this behavior between Gophersnakes from mainland California (where rattlesnakes are present) versus nearby derived island populations (where rattlesnakes are absent), we found that mainland snakes vibrated their tail for a longer duration, thereby producing a signal more similar to rattlesnakes. At the same time, we did not find evidence that island snakes differ from mainland snakes in the rate at which they vibrate their tails, or their propensity to vibrate their tails. Taken together, these data lend some support to the longstanding hypothesis that defensive tail vibration by nonvenomous snakes represents acoustic mimicry of rattlesnakes. These data further suggest that this mimetic signal may be undergoing erosion in populations where rattlesnakes are absent.

Sensory bias and signal detection trade-offs maintain intersexual floral mimicry

Mimicry is common in interspecies interactions, yet conditions maintaining Batesian mimicry have been primarily tested in predator-prey interactions. In pollination mutualisms, floral mimetic signals thought to dupe animals into pollinating unrewarding flowers are widespread (greater than 32 plant families). Yet whether animals learn to both correctly identify floral models and reject floral mimics and whether these responses are frequency-dependent is not well understood. We tested how learning affected the effectiveness and frequency-dependence of imperfect Batesian mimicry among flowers using the generalist bumblebee, Bombus impatiens, visiting Begonia odorata, a plant species exhibiting intersexual floral mimicry. Unrewarding female flowers are mimics of pollen-rewarding male flowers (models), though mimicry to the human eye is imperfect. Flower-naive bees exhibited a perceptual bias for mimics over models, but rapidly learned to avoid mimics. Surprisingly, altering the frequency of models and mimics only marginally shaped responses by naive bees and by bees experienced with the distribution and frequency of models and mimics. Our results provide evidence both of exploitation by the plant of signal detection trade-offs in bees and of resistance by the bees, via learning, to this exploitation. Critically, we provide experimental evidence that imperfect Batesian mimicry can be adaptive and, in contrast with expectations of signal detection theory, functions largely independently of the model and mimic frequency. This article is part of the theme issue 'Signal detection theory in recognition systems: from evolving models to experimental tests'.

A pheromone antagonist liberates female sea lamprey from a sensory trap to enable reliable communication

The evolution of male signals and female preferences remains a central question in the study of animal communication. The sensory trap model suggests males evolve signals that mimic cues used in nonsexual contexts and thus manipulate female behavior to generate mating opportunities. Much evidence supports the sensory trap model, but how females glean reliable information from both mimetic signals and their model cues remains unknown. We discovered a mechanism whereby a manipulative male signal guides reliable communication in sea lamprey (Petromyzon marinus). Migratory sea lamprey follow a larval cue into spawning streams; once sexually mature, males release a pheromone that mimics the larval cue and attracts females. Females conceivably benefit from the mimetic pheromone during mate search but must discriminate against the model cue to avoid orienting toward larvae in nearby nursery habitats. We tested the hypothesis that spawning females respond to petromyzonol sulfate (PZS) as a behavioral antagonist to avoid attraction to the larval cue while tracking the male pheromone despite each containing attractive 3-keto petromyzonol sulfate (3kPZS). We found 1) PZS inhibited electrophysiological responses to 3kPZS and abated preferences for 3kPZS when mixed at the same or greater concentrations, 2) larvae released more PZS than 3kPZS whereas males released more 3kPZS than PZS, and 3) mixtures of 3kPZS and PZS applied at ratios measured in larval and male odorants resulted in the discrimination observed between the natural odors. Our study elucidates how communication systems that arise via deception can facilitate reliable communication.

How cognitive biases select for imperfect mimicry: a study of asymmetry in learning with bumblebees

Imperfect mimicry presents a paradox of incomplete adaptation - intuitively, closer resemblance should improve performance. Receiver psychology can often explain why mimetic signals do not always evolve to match those of their models. Here, we explored the influence of a pervasive and powerful cognitive bias where associative learning depends upon an asymmetric interaction between the cue (stimulus) and consequence (reinforcer), such as in rats, which will associate light and tone with shock, and taste with nausea, but not the converse. Can such biases alter selection for mimicry? We designed an artificial mimicry system where bees foraged on artificial flowers, so that colours could be switched between rewarding or aversive. We found that when the colour blue was paired with a sucrose reward, other cues were ignored, but not when blue was paired with aversive compounds. We also tested the hypothesis that costs of errors affect how receivers sample imperfect mimics. However, costs of errors did not affect bee visits to imperfect mimics in our study. We propose a novel hypothesis for imperfect mimicry, in which the pairing between specific cues and reinforcers allows an imperfect mimic to resemble multiple models simultaneously. Generally, our results emphasize the importance of receiver psychology for the evolution of signal complexity and specificity.

Supramolecular Nanostructure Activates TrkB Receptor Signaling of Neuronal Cells by Mimicking Brain-Derived Neurotrophic Factor.

Brain-derived neurotrophic factor (BDNF), a neurotrophin that binds specifically to the tyrosine kinase B (TrkB) receptor, has been shown to promote neuronal differentiation, maturation, and synaptic plasticity in the central nervous system (CNS) during development or after injury and onset of disease. Unfortunately, native BDNF protein-based therapies have had little clinical success due to their suboptimal pharmacological properties. In the past 20 years, BDNF mimetic peptides have been designed with the purpose of activating certain cell pathways that mimic the functional activity of native BDNF, but the interaction of mimetic peptides with cells can be limited due to the conformational specificity required for receptor activation. We report here on the incorporation of a BDNF mimetic sequence into a supramolecular peptide amphiphile filamentous nanostructure capable of activating the BDNF receptor TrkB and downstream signaling in primary cortical neurons in vitro. Interestingly, we found that this BDNF mimetic peptide is only active when displayed on a peptide amphiphile supramolecular nanostructure. We confirmed that increased neuronal maturation is linked to TrkB signaling pathways by analyzing the phosphorylation of downstream signaling effectors and tracking electrical activity over time. Furthermore, three-dimensional gels containing the BDNF peptide amphiphile (PA) nanostructures encourage cell infiltration while increasing functional maturation. Our findings suggest that the BDNF mimetic PA nanostructure creates a highly bioactive matrix that could serve as a biomaterial therapy in injured regions of the CNS. This new strategy has the potential to induce endogenous cell infiltration and promote functional neuronal maturation through the presentation of the BDNF mimetic signal.

Frequency dependence shapes the adaptive landscape of imperfect Batesian mimicry.

Despite more than a century of biological research on the evolution and maintenance of mimetic signals, the relative frequencies of models and mimics necessary to establish and maintain Batesian mimicry in natural populations remain understudied. Here we investigate the frequency-dependent dynamics of imperfect Batesian mimicry, using predation experiments involving artificial butterfly models. We use two geographically distinct populations of Adelpha butterflies that vary in their relative frequencies of a putatively defended model (Adelpha iphiclus) and Batesian mimic (Adelpha serpa). We found that in Costa Rica, where both species share similar abundances, Batesian mimicry breaks down, and predators more readily attack artificial butterfly models of the presumed mimic, A. serpa By contrast, in Ecuador, where A. iphiclus (model) is significantly more abundant than A. serpa (mimic), both species are equally protected from predation. Our results provide compelling experimental evidence that imperfect Batesian mimicry is frequency-dependent on the relative abundance of models and mimics in natural populations, and contribute to the growing body of evidence that complex dynamics, such as seasonality or the availability of alternative prey, influence the evolution of mimetic traits.

Viral tRNA Mimicry from a Biocommunicative Perspective.

RNA viruses have very small genomes which limits the functions they can encode. One of the strategies employed by these viruses is to mimic key factors of the host cell so they can take advantage of the interactions and activities these factors typically participate in. The viral RNA genome itself was first observed to mimic cellular tRNA over 40 years ago. Since then researchers have confirmed that distinct families of RNA viruses are accessible to a battery of cellular factors involved in tRNA-related activities. Recently, potential tRNA-like structures have been detected within the sequences of a 100 mRNAs taken from human cells, one of these being the host defense interferon-alpha mRNA; these are then additional to the examples found in bacterial and yeast mRNAs. The mimetic relationship between tRNA, cellular mRNA, and viral RNA is the central focus of two considerations described below. These are subsequently used as a preface for a final hypothesis drawing on concepts relating to mimicry from the social sciences and humanities, such as power relations and creativity. Firstly, the presence of tRNA-like structures in mRNAs indicates that the viral tRNA-like signal could be mimicking tRNA-like elements that are contextualized by the specific carrier mRNAs, rather than, or in addition to, the tRNA itself, which would significantly increase the number of potential semiotic relations mediated by the viral signals. Secondly, and in particular, mimicking a host defense mRNA could be considered a potential new viral strategy for survival. Finally, we propose that mRNA’s mimicry of tRNA could be indicative of an ancestral intracellular conflict in which species of mRNAs invaded the cell, but from within. As the meaning of the mimetic signal depends on the context, in this case, the conflict that arises when the viral signal enters the cell can change the meaning of the mRNAs’ internal tRNA-like signals, from their current significance to that they had in the distant past.

Polymorphism at a mimicry supergene maintained by opposing frequency-dependent selection pressures

Explaining the maintenance of adaptive diversity within populations is a long-standing goal in evolutionary biology, with important implications for conservation, medicine, and agriculture. Adaptation often leads to the fixation of beneficial alleles, and therefore it erodes local diversity so that understanding the coexistence of multiple adaptive phenotypes requires deciphering the ecological mechanisms that determine their respective benefits. Here, we show how antagonistic frequency-dependent selection (FDS), generated by natural and sexual selection acting on the same trait, maintains mimicry polymorphism in the toxic butterfly Heliconius numata Positive FDS imposed by predators on mimetic signals favors the fixation of the most abundant and best-protected wing-pattern morph, thereby limiting polymorphism. However, by using mate-choice experiments, we reveal disassortative mate preferences of the different wing-pattern morphs. The resulting negative FDS on wing-pattern alleles is consistent with the excess of heterozygote genotypes at the supergene locus controlling wing-pattern variation in natural populations of H. numata The combined effect of positive and negative FDS on visual signals is sufficient to maintain a diversity of morphs displaying accurate mimicry with other local prey, although some of the forms only provide moderate protection against predators. Our findings help understand how alternative adaptive phenotypes can be maintained within populations and emphasize the need to investigate interactions between selective pressures in other cases of puzzling adaptive polymorphism.

A bee or not a bee: an experimental test of acoustic mimicry by hoverflies

The degree of similarity between Batesian mimics and their models varies widely and occurs across a range of sensory modalities. We use three complementary experimental paradigms to investigate acoustic mimicry in hoverflies (Diptera: Syrphidae) which mimic stinging Hymenoptera. First, we analyse sounds made by 13 hoverfly species and 9 Hymenoptera species with and without simulated predation (“alarm” and “flight” sounds, respectively). We demonstrate that the bumblebees Bombus terrestris, B. hortorum, and B. lucorum, and the hoverfly Cheilosia illustrata exhibit alarm sounds that are significantly different to their respective flight sounds, and indistinguishable between species. We then demonstrate that the B. terrestris alarm sound reduces predation on artificial prey by wild birds, but that the hoverfly mimic alarm sound does not. Finally, we trained chicks to avoid distasteful food in the presence of different acoustic stimuli. Overall the chicks showed no difference in response to bee and hoverfly stimuli, demonstrating no innate aversion to the Bombus alarm sound. We therefore conclude that (i) similarity of acoustic signals exists among Hymenoptera and hoverflies, (ii) acoustic aposematic signals (but not the almost identical mimetic signals) are effective at reducing predation, and (iii) wild birds exhibit learned rather than innate aversion to certain acoustic stimuli.

Unlinked Mendelian inheritance of red and black pigmentation in snakes: Implications for Batesian mimicry

Identifying the genetic basis of mimetic signals is critical to understanding both the origin and dynamics of mimicry over time. For species not amenable to large laboratory breeding studies, widespread color polymorphism across natural populations offers a powerful way to assess the relative likelihood of different genetic systems given observed phenotypic frequencies. We classified color phenotype for 2175 ground snakes (Sonora semiannulata) across the continental United States to analyze morph ratios and test among competing hypotheses about the genetic architecture underlying red and black coloration in coral snake mimics. We found strong support for a two-locus model under simple Mendelian inheritance, with red and black pigmentation being controlled by separate loci. We found no evidence of either linkage disequilibrium between loci or sex linkage. In contrast to Batesian mimicry systems such as butterflies in which all color signal components are linked into a single "supergene," our results suggest that the mimetic signal in colubrid snakes can be disrupted through simple recombination and that color evolution is likely to involve discrete gains and losses of each signal component. Both outcomes are likely to contribute to the exponential increase in rates of color evolution seen in snake mimicry systems over insect systems.

Locomotory mimicry in ant-like spiders

The jumping spider genus Myrmarachne (Salticidae) consists of over 200 species of morphologically accurate ant mimics, enabling Myrmarachne to evade ant-averse predators that confuse the spiders with ants (Batesian mimicry). A conspicuous but untested trait of Myrmarachne, which is frequently mentioned in the literature, is locomotory mimicry. For these spiders, locomotion that is more ant like than salticid like may be an integral part of the suite of mimetic signals. We quantified the locomotory pattern of several species of non-ant-like salticids, Myrmarachne, and ants from Australia. We found that the locomotion of the mimics generally resembles that of ants, but not of other salticids. To tease out the effects of locomotion from those based on morphology, ant-eating salticids as visual predators were presented with 3D animations of ant-like salticids walking either in ant or in salticid fashion. Ant-eating salticids spent significantly longer looking at the stimulus with ant-like movement and were significantly more likely to visually track and stalk the stimulus with ant-like movement. Overall, our results suggest that there is selection pressure on Myrmarachne for increased resemblance to a model by locomotor mimicry, despite the associated costs when faced with ant-eating predators.

Costs of Learning and the Evolution of Mimetic Signals.

Predators must use the appearance of their prey to decide whether it is likely to be defended. Most theory assumes that predators have completed learning about prey appearance, yet we do not understand how predators learn which aspects of appearance to use for classifying prey. If sampling prey can be risky, predators might forgo opportunities to learn about the relationship between prey appearance and defense. Using Bayesian inference and dynamic programming, we modeled how the immediate risks and future rewards of learning about prey appearance influence how predators learn. In addition, we explored how variation in predator learning affects the evolution of mimicry, which occurs when two prey evolve to share a common signal to predators. We found that when learning about prey with distinct appearances was expensive, optimal predators tended to lump them into the same category or exhibit an unwillingness to sample at all (neophobia). This resulted in a reduction in selection for defensive mimicry. However, the same predator behavior favored the evolution of aggressive mimicry, because in that case, mimics benefited from being sampled. When prey were very rare and costs of sampling them were high, predators exhibited neophobia, refusing to attack. This behavior could forestall the evolution of mimicry and instead select for polymorphism.

Phenotypic Plasticity Confers Multiple Fitness Benefits to a Mimic

Animal communication is often deceptive; however, such dishonesty can become ineffective if it is used too often, is used out of context, or is too easy to detect [1-3]. Mimicry is a common form of deception, and most mimics gain the greatest fitness benefits when they are rare compared to their models [3, 4]. If mimics are encountered too frequently or if their model is absent, avoidance learning of noxious models is disrupted (Batesian mimicry [3]), or receivers become more vigilant and learn to avoid perilous mimics (aggressive mimicry [4]). Mimics can moderate this selective constraint by imperfectly resembling multiple models [5], through polymorphisms [6], or by opportunistically deploying mimetic signals [1, 7]. Here we uncover a novel mechanism to escape the constraints of deceptive signaling: phenotypic plasticity allows mimics to deceive targets using multiple guises. Using a combination of behavioral, cell histological, and molecular methods, we show that a coral reef fish, the dusky dottyback (Pseudochromis fuscus), flexibly adapts its body coloration to mimic differently colored reef fishes and in doing so gains multiple fitness benefits. We find that by matching the color of other reef fish, dottybacks increase their success of predation upon juvenile fish prey and are therefore able to deceive their victims by resembling multiple models. Furthermore, we demonstrate that changing color also increases habitat-associated crypsis that decreases the risk of being detected by predators. Hence, when mimics and models share common selective pressures, flexible imitation of models might inherently confer secondary benefits to mimics. Our results show that phenotypic plasticity can act as a mechanism to ease constraints that are typically associated with deception. VIDEO ABSTRACT.

Avian vocal mimicry: a unified conceptual framework.

Mimicry is a classical example of adaptive signal design. Here, we review the current state of research into vocal mimicry in birds. Avian vocal mimicry is a conspicuous and often spectacular form of animal communication, occurring in many distantly related species. However, the proximate and ultimate causes of vocal mimicry are poorly understood. In the first part of this review, we argue that progress has been impeded by conceptual confusion over what constitutes vocal mimicry. We propose a modified version of Vane-Wright's (1980) widely used definition of mimicry. According to our definition, a vocalisation is mimetic if the behaviour of the receiver changes after perceiving the acoustic resemblance between the mimic and the model, and the behavioural change confers a selective advantage on the mimic. Mimicry is therefore specifically a functional concept where the resemblance between heterospecific sounds is a target of selection. It is distinct from other forms of vocal resemblance including those that are the result of chance or common ancestry, and those that have emerged as a by-product of other processes such as ecological convergence and selection for large song-type repertoires. Thus, our definition provides a general and functionally coherent framework for determining what constitutes vocal mimicry, and takes account of the diversity of vocalisations that incorporate heterospecific sounds. In the second part we assess and revise hypotheses for the evolution of avian vocal mimicry in the light of our new definition. Most of the current evidence is anecdotal, but the diverse contexts and acoustic structures of putative vocal mimicry suggest that mimicry has multiple functions across and within species. There is strong experimental evidence that vocal mimicry can be deceptive, and can facilitate parasitic interactions. There is also increasing support for the use of vocal mimicry in predator defence, although the mechanisms are unclear. Less progress has been made in explaining why many birds incorporate heterospecific sounds into their sexual displays, and in determining whether these vocalisations are functionally mimetic or by-products of sexual selection for other traits such as repertoire size. Overall, this discussion reveals a more central role for vocal mimicry in the behavioural ecology of birds than has previously been appreciated. The final part of this review identifies important areas for future research. Detailed empirical data are needed on individual species, including on the structure of mimetic signals, the contexts in which mimicry is produced, how mimicry is acquired, and the ecological relationships between mimic, model and receiver. At present, there is little information and no consensus about the various costs of vocal mimicry for the protagonists in the mimicry complex. The diversity and complexity of vocal mimicry in birds raises important questions for the study of animal communication and challenges our view of the nature of mimicry itself. Therefore, a better understanding of avian vocal mimicry is essential if we are to account fully for the diversity of animal signals.