Multi-trait mimicry and the relative salience of individual traits

Mimicry occurs when one species gains protection from predators by resembling an unprofitable model species. The degree of mimic–model similarity is variable in nature and is closely related to the number of traits that the mimic shares with its model. Here, we experimentally test the hypothesis that the relative salience of traits, as perceived by a predator, is an important determinant of the degree of mimic–model similarity required for successful mimicry. We manipulated the relative salience of the traits of a two-trait artificial model prey, and subsequently tested the survival of mimics of the different traits. The unrewarded model prey had two colour traits, black and blue, and the rewarded prey had two combinations of green, brown and grey shades. Blue tits were used as predators. We found that the birds perceived the black and blue traits to be similarly salient in one treatment, and mimic–model similarity in both traits was then required for high mimic success. In a second treatment, the blue trait was the most salient trait, and mimic–model similarity in this trait alone achieved high success. Our results thus support the idea that similar salience of model traits can explain the occurrence of multi-trait mimicry.     

Numbers,neighbors, and hungry predators: What makes chemically defended aposematic prey susceptible to predation?

Many chemically defended aposematic species are characterized by relatively low toxin levels, which enables predators to include them in their diets under certain circumstances. Knowledge of the conditions governing the survival of such prey animals—especially in the context of the co‐occurrence of similar but undefended prey, which may result in mimicry‐like interactions—is crucial for understanding the initial evolution of aposematism. In a one‐month outdoor experiment using fish (the common carp Cyprinus carpio) as predators, we examined the survival of moderately defended aposematic tadpole prey (the European common toad Bufo bufo) with varying absolute densities in single‐species prey systems or varying relative densities in two‐species prey systems containing morphologically similar but undefended prey (the European common frog Rana temporaria). The density effects were investigated in conjunction with the hunger levels of the predator, which were manipulated by means of the addition of alternative (nontadpole) food. The survival of the B. bufo tadpoles was promoted by increasing their absolute density in the single‐species prey systems, increasing their relative density in the two‐species prey systems, and providing ample alternative food for the predator. Hungry predators eliminated all R. temporaria individuals regardless of their proportion in the prey community; in treatments with ample alternative food, high relative B. bufo density supported R. temporaria survival. The results demonstrated that moderately defended prey did benefit from high population densities (both absolute and relative), even under long‐term predation pressure. However, the physiological state of the predator was a crucial factor in the survival of moderately defended prey. While the availability of alternative prey in general should promote the spread and maintenance of aposematism, the results indicated that the resemblance between the co‐occurring defended and undefended prey may impose mortality costs on the defended model species, even in the absence of actual mimicry.     

The normalized segment classification model: A new tool to compare spectral reflectance curves

1. Color patterns are complex traits under selective pressures from conspecifics, mutualists, and antagonists. To evaluate the salience of a pattern or the similarity between colors, several visual models are available. Color discrimination models estimate the perceptual difference between any two colors. Their application to a diversity of taxonomic groups has become common in the literature to answer behavioral, ecological, and evolutionary questions. To use these models, we need information about the visual system of our beholder species. However, many color patterns are simultaneously subject to selective pressures from different species, often from different taxonomic groups, with different visual systems. Furthermore, we lack information about the visual system of many species, leading ecologists to use surrogate values or theoretical estimates for model parameters.
2. Here, we present a modification of the segment classification method proposed by Endler (Biological Journal of the Linnean Society, 1990 41, 315–352): the normalized segment classification model (NSC). We explain its logic and use, exploring how NSC differs from other visual models. We also compare its predictions with available experimental data.
3. Even though the NSC model includes no information about the visual system of the receiver species, it performed better than traditional color discrimination models when predicting the output of some behavioral tasks. Although vision scientists define color as independent of stimulus brightness, a likely explanation for the goodness of fit of the NSC model is that its distance measure depends on brightness differences, and achromatic information can influence the decision‐making process of animals when chromatic information is missing.
4. Species‐specific models may be insufficient for the study of color patterns in a community context. The NSC model offers a species‐independent solution for color analyses, allowing us to calculate color differences when we ignore the intended viewer of a signal or when different species impose selective pressures on the signal.

     

Balanced polymorphisms and their divergence in a Heliconius butterfly

The evolution of mimicry in similarly defended prey is well described by the Müllerian mimicry theory, which predicts the convergence of warning patterns in order to gain the most protection from predators. However, despite this prediction, we can find great diversity of color patterns among Müllerian mimics such as Heliconius butterflies in the neotropics. Furthermore, some species have evolved the ability to maintain multiple distinct warning patterns in single populations, a phenomenon known as polymorphic mimicry. The adaptive benefit of these polymorphisms is questionable since variation from the most common warning patterns is expected to be disadvantageous as novel signals are punished by predators naive to them. In this study, we use artificial butterfly models throughout Central and South America to characterize the selective pressures maintaining polymorphic mimicry in Heliconius doris. Our results highlight the complexity of positive frequency‐dependent selection, the principal selective pressure driving convergence among Müllerian mimics, and its impacts on interspecific variation of mimetic warning coloration. We further show how this selection regime can both limit and facilitate the diversification of mimetic traits.     

A complex mode of aggressive mimicry in a scale-eating cichlid fish

Aggressive mimicry is an adaptive tactic of parasitic or predatory species that closely resemble inoffensive models in order to increase fitness via predatory gains. Although similarity of distantly related species is often intuitively implicated with mimicry, the exact mechanisms and evolutionary causes remain elusive in many cases. Here, we report a complex aggressive mimicry strategy in Plecodus straeleni, a scale-eating cichlid fish from Lake Tanganyika, which imitates two other cichlid species. Employing targeted sequencing on ingested scales, we show that P. straeleni does not preferentially parasitize its models but—contrary to prevailing assumptions—targets a variety of co-occurring dissimilar looking fish species. Combined with tests for visual resemblance and visual modelling from a prey perspective, our results suggest that complex interactions among different cichlid species are involved in this mimicry system.     

Masquerading predators deceive prey by aggressively mimicking bird droppings in a crab spider

In aggressive mimicry, a predator accesses prey by mimicking the appearance and/or behavior of a harmless or beneficial model in order to avoid being correctly identified by its prey. The crab spider genus Phrynarachne is often cited as a textbook example of masquerading as bird droppings (BDs) in order to avoid predation. However, Phrynarachne spiders may also aggressively mimic BDs in order to deceive potential prey. To date, there is no experimental evidence to support aggressive mimicry in masquerading crab spiders; therefore, we performed a field survey, a manipulative field experiment, and visual modeling to test this hypothesis using Phrynarachne ceylonica. We compared prey-attraction rates among BDs, spiders, and control empty leaves in the field. We found that although all prey combined and agromyzid dipterans, in particular, were attracted to BDs at a higher rate than to spiders, other dipterans and hymenopterans were attracted to BDs at a similar rate as to spiders. Both spiders and BDs attracted insects at a significantly higher rate than did control leaves. As predicted, prey was attracted to experimentally blackened or whitened spiders significantly less frequently than to unmanipulated spiders. Finally, visual modeling suggested that spiders and BDs can be detected by dipterans and hymenopterans against background leaves, but they are indistinguishable from each other. Taken together, our results suggest that insects lured by spiders may misidentify them as BDs, and bird-dropping masquerading may serve as aggressive mimicry in addition to predator avoidance in P. ceylonica.     

Juvenile Snooks (Centropomidae) as Mimics of Mojarras (Gerreidae), with a Review of Aggressive Mimicry in Fishes

Aggressive mimicry has been proposed for several unrelated fish species both in freshwater and marine environments. I describe herein a few additional examples, including the first ones from brackish water. In one well documented case, juvenile snooks, Centropomus mexicanus (Centropomidae) join bottom-foraging groups of the superficially similar mojarras, Eucinostomus melanopterus (Gerreidae) and prey on small fishes and crustaceans under such disguise. Two other snook species and two species of groupers (Serranidae), are here suggested as additional instances of aggressive mimicry. Furthermore, I review published examples of aggressive mimicry in fishes and indicate trends in the relationships between the mimics, their feeding tactics, and their putative models. Three large families, Serranidae, Cichlidae, and Blenniidae display most of the examples of aggressive mimicry, serranids being largely represented by the genus Hypoplectrus and blenniids by the tribe Nemophini only. Three major trends are here indicated for aggressive mimics: (1) fish species that feed on prey smaller than themselves tend to mimic and join fish species harmless to their prospective prey; (2) fish species that feed on prey larger than themselves tend to mimic mostly beneficial fish species (cleaners) or, less frequently, join species harmless to their prospective prey; (3) fish species that feed on prey about their own size tend to mimic their prospective prey species, the perfect wolf in a sheep's clothes disguise type. The latter deceit is recorded mostly for scale and fin-feeding freshwater fishes.     

Aggressive use of Batesian mimicry by an ant-like jumping spider

Batesian and aggressive mimicry are united by deceit: Batesian mimics deceive predators and aggressive mimics deceive prey. This distinction is blurred by Myrmarachne melanotarsa, an ant-like jumping spider (Salticidae). Besides often preying on salticids, ants are well defended against most salticids that might target them as potential prey. Earlier studies have shown that salticids identify ants by their distinctive appearance and avoid them. They also avoid ant-like salticids from the genus Myrmarachne. Myrmarachne melanotarsa is an unusual species from this genus because it typically preys on the eggs and juveniles of ant-averse salticid species. The hypothesis considered here is that, for M. melanotarsa, the distinction between Batesian and aggressive mimicry is blurred. We tested this by placing female Menemerus sp. and their associated hatchling within visual range of M. melanotarsa, its model, and various non-ant-like arthropods. Menemerus is an ant-averse salticid species. When seeing ants or ant mimics, Menemerus females abandoned their broods more frequently than when seeing non-ant-like arthropods or in control tests (no arthropods visible), as predicted by our hypothesis that resembling ants functions as a predatory ploy.     

Fenestration: a window of opportunity for carnivorous plants

A long-standing but controversial hypothesis assumes that carnivorous plants employ aggressive mimicry to increase their prey capture success. A possible mechanism is that pitcher plants use aggressive mimicry to deceive prey about the location of the pitcher''s exit. Specifically, species from unrelated families sport fenestration, i.e. transparent windows on the upper surfaces of pitchers which might function to mimic the exit of the pitcher. This hypothesis has not been evaluated against alternative hypotheses predicting that fenestration functions to attract insects from afar. By manipulating fenestration, we show that it does not increase the number of Drosophila flies or of two ant species entering pitchers in Sarracenia minor nor their retention time or a pitcher''s capture success. However, fenestration increased the number of Drosophila flies alighting on the pitcher compared with pitchers of the same plant without fenestration. We thus suggest that fenestration in S. minor is not an example of aggressive mimicry but rather functions in long-range attraction of prey. We highlight the need to evaluate aggressive mimicry relative to alternative concepts of plant–animal communication.     

Who Resembles Whom? Mimetic and Coincidental Look-Alikes among Tropical Reef Fishes

Studies of mimicry among tropical reef-fishes usually give little or no consideration to alternative explanations for behavioral associations between unrelated, look-alike species that benefit the supposed mimic. I propose and assess such an alternative explanation. With mimicry the mimic resembles its model, evolved to do so in response to selection by the mimicry target, and gains evolved benefits from that resemblance. In the alternative, the social-trap hypothesis, a coincidental resemblance of the model to the “mimic” inadvertently attracts the latter to it, and reinforcement of this social trapping by learned benefits leads to the “mimic” regularly associating with the model. I examine three well known cases of supposed aggressive mimicry among reef-fishes in relation to nine predictions from these hypotheses, and assess which hypothesis offers a better explanation for each. One case, involving precise and complex morphological and behavioral resemblance, is strongly consistent with mimicry, one is inconclusive, and one is more consistent with a social-trap based on coincidental, imprecise resemblance. Few cases of supposed interspecific mimicry among tropical reef fishes have been examined in depth, and many such associations may involve social traps arising from generalized, coincidental resemblance. Mimicry may be much less common among these fishes than is generally thought.     

Temporal changes in hamlet communities (Hypoplectrus spp., Serranidae) over 17 years

Transect surveys of hamlet communities (Hypoplectrus spp., Serranidae) covering 14 000 m² across 16 reefs off La Parguera, Puerto Rico, are presented and compared with a previous survey conducted in the year 2000. The hamlet community has noticeably changed over 17 years, with a > 30% increase in relative abundance of the yellowtail hamlet Hypoplectrus chlorurus on the inner reefs at the expense of the other hamlet species. The data also suggest that the density of H. chlorurus has declined and that its distribution has shifted towards shallower depths. Considering that H. chlorurus has been previously identified as one of the few fish showing a positive association with seawater turbidity on the inner reefs of La Parguera and that sedimentation of terrestrial origin has increased over recent decades on these reefs, it is proposed that turbidity may constitute an important but so far overlooked ecological driver of hamlet communities.     

The visual ecology of selective predation: Are unhealthy hosts less stealthy hosts?

Predators can strongly influence disease transmission and evolution, particularly when they prey selectively on infected hosts. Although selective predation has been observed in numerous systems, why predators select infected prey remains poorly understood. Here, we use a mathematical model of predator vision to test a long‐standing hypothesis about the mechanistic basis of selective predation in a Daphnia–microparasite system, which serves as a model for the ecology and evolution of infectious diseases. Bluegill sunfish feed selectively on Daphnia infected by a variety of parasites, particularly in water uncolored by dissolved organic carbon. The leading hypothesis for selective predation in this system is that infection‐induced changes in the transparency of Daphnia render them more visible to bluegill. Rigorously evaluating this hypothesis requires that we quantify the effect of infection on the visibility of prey from the predator''s perspective, rather than our own. Using a model of the bluegill visual system, we show that three common parasites, Metschnikowia bicuspidata, Pasteuria ramosa, and Spirobacillus cienkowskii, decrease the transparency of Daphnia, rendering infected Daphnia darker against a background of bright downwelling light. As a result of this increased brightness contrast, bluegill can see infected Daphnia at greater distances than uninfected Daphnia—between 19% and 33% further, depending on the parasite. Pasteuria and Spirobacillus also increase the chromatic contrast of Daphnia. These findings lend support to the hypothesis that selective predation by fish on infected Daphnia could result from the effects of infection on Daphnia''s visibility. However, contrary to expectations, the visibility of Daphnia was not strongly impacted by water color in our model. Our work demonstrates that models of animal visual systems can be useful in understanding ecological interactions that impact disease transmission.     

Population genetic structure and evolution of Batesian mimicry in Papilio polytes from the Ryukyu Islands,Japan, analyzed by genotyping‐by‐sequencing

Batesian mimicry is a striking example of Darwinian evolution, in which a mimetic species resembles toxic or unpalatable model species, thereby receiving protection from predators. In some species exhibiting Batesian mimicry, nonmimetic individuals coexist as polymorphism in the same population despite the benefits of mimicry. In a previous study, we proposed that the abundance of mimics is limited by that of the models, leading to polymorphic Batesian mimicry in the swallowtail butterfly, Papilio polytes, on the Ryukyu Islands in Japan. We found that their mimic ratios (MRs), which varied among the Islands, were explained by the model abundance of each habitat, rather than isolation by distance or phylogenetic constraint based on the mitochondrial DNA (mtDNA) analysis. In the present study, this possibility was reexamined based on hundreds of nuclear single nucleotide polymorphisms (SNPs) of 93 P. polytes individuals from five Islands of the Ryukyus. We found that the population genetic and phylogenetic structures of P. polytes largely corresponded to the geographic arrangement of the habitat Islands, and the genetic distances among island populations show significant correlation with the geographic distances, which was not evident by the mtDNA‐based analysis. A partial Mantel test controlling for the present SNP‐based genetic distances revealed that the MRs of P. polytes were strongly correlated with the model abundance of each island, implying that negative frequency‐dependent selection interacting with model species shaped and maintained the mimetic polymorphism. Taken together, our results support the possibility that predation pressure, not isolation by distance or other neutral factors, is a major driving force of evolution of the Batesian mimicry in P. polytes from the Ryukyus.     

Comparison of native and non‐native predator consumption rates and prey avoidance behavior in North America and Europe

Novel predator–prey interactions can contribute to the invasion success of non‐native predators. For example, native prey can fail to recognize and avoid non‐native predators due to a lack of co‐evolutionary history and cue dissimilarity with native predators. This might result in a competitive advantage for non‐native predators. Numerous lady beetle species were globally redistributed as biological control agents against aphids, resulting in novel predator–prey interactions. Here, we investigated the strength of avoidance behavior of the pea aphid (Acyrthosiphon pisum) toward chemical cues of native lady beetles and non‐native Asian Harmonia axyridis and European Coccinella septempunctata and Hippodamia variegata in North America, hypothesizing that cues of non‐native lady beetles induce weaker avoidance behavior than cues of co‐evolved native lady beetles. Additionally, we compared aphid consumption of lady beetles, examining potential predation advantages of non‐native lady beetles. Finally, we compared cue avoidance behavior between North American and European pea aphid populations and aphid consumption of native and non‐native lady beetles in North America and Europe. In North America, pea aphids avoided chemical cues of all ladybeetle species tested, regardless of their origin. In contrast to pea aphids in North America, European pea aphids did not avoid cues of the non‐native H. axyridis. The non‐native H. axyridis and C. septempunctata were among the largest and most voracious lady beetle species tested, on both continents. Consequently, in North America non‐native lady beetle species might have a competitive advantage on shared food resources due to their relatively large body size, compared to several native American lady beetle species. In Europe, however, non‐native H. axyridis might benefit from missing aphid cue avoidance as well as a large body size. The co‐evolutionary time gap between the European and North American invasion of H. axyridis likely explains the intercontinental differences in cue avoidance behavior and might indicate evolution in aphids toward non‐native predators.     

Mito‐nuclear discordance at a mimicry color transition zone in bumble bee Bombus melanopygus

As hybrid zones exhibit selective patterns of gene flow between otherwise distinct lineages, they can be especially valuable for informing processes of microevolution and speciation. The bumble bee, Bombus melanopygus, displays two distinct color forms generated by Müllerian mimicry: a northern “Rocky Mountain''’ color form with ferruginous mid‐abdominal segments (B. m. melanopygus) and a southern “Pacific''’ form with black mid‐abdominal segments (B. m. edwardsii). These morphs meet in a mimetic transition zone in northern California and southern Oregon that is more narrow and transitions further west than comimetic bumble bee species. To understand the historical formation of this mimicry zone, we assessed color distribution data for B. melanopygus from the last 100 years. We then examined gene flow among the color forms in the transition zone by comparing sequences from mitochondrial COI barcode sequences, color‐controlling loci, and the rest of the nuclear genome. These data support two geographically distinct mitochondrial haplogroups aligned to the ancestrally ferruginous and black forms that meet within the color transition zone. This clustering is also supported by the nuclear genome, which, while showing strong admixture across individuals, distinguishes individuals most by their mitochondrial haplotype, followed by geography. These data suggest the two lineages most likely were historically isolated, acquired fixed color differences, and then came into secondary contact with ongoing gene flow. The transition zone, however, exhibits asymmetries: mitochondrial haplotypes transition further south than color pattern, and both transition over shorter distances in the south. This system thus demonstrates alternative patterns of gene flow that occur in contact zones, presenting another example of mito‐nuclear discordance. Discordant gene flow is inferred to most likely be driven by a combination of mimetic selection, dominance effects, and assortative mating.     

High-model abundance may permit the gradual evolution of Batesian mimicry: an experimental test

In Batesian mimicry, a harmless species (the ‘mimic’) resembles a dangerous species (the ‘model’) and is thus protected from predators. It is often assumed that the mimetic phenotype evolves from a cryptic phenotype, but it is unclear how a population can transition through intermediate phenotypes; such intermediates may receive neither the benefits of crypsis nor mimicry. Here, we ask if selection against intermediates weakens with increasing model abundance. We also ask if mimicry has evolved from cryptic phenotypes in a mimetic clade. We first present an ancestral character-state reconstruction showing that mimicry of a coral snake (Micrurus fulvius) by the scarlet kingsnake (Lampropeltis elapsoides) evolved from a cryptic phenotype. We then evaluate predation rates on intermediate phenotypes relative to cryptic and mimetic phenotypes under conditions of both high- and low-model abundances. Our results indicate that where coral snakes are rare, intermediate phenotypes are attacked more often than cryptic and mimetic phenotypes, indicating the presence of an adaptive valley. However, where coral snakes are abundant, intermediate phenotypes are not attacked more frequently, resulting in an adaptive landscape without a valley. Thus, high-model abundance may facilitate the evolution of Batesian mimicry.     

A context analysis of bobbing and fin‐flicking in a small marine benthic fish

Most antipredator strategies increase survival of individuals by signaling to predators, by reducing the chances of being recognized as prey, or by bewildering a predator''s perception. In fish, bobbing and fin‐flicking are commonly considered as pursuit‐deterrent behaviors that signal a predator that it has been detected and thus lost its surprise‐attack advantage. Yet, very few studies assessed whether such behavioral traits are restricted to the visual presence of a predator. In this study, we used the yellow black‐headed triplefin Tripterygion delaisi to investigate the association between these behaviors and the visual exposure to (a) a black scorpionfish predator (Scorpaena porcus), (b) a stone of a size similar to that of S. porcus, (c) a conspecific, and (d) a harmless heterospecific combtooth blenny (Parablennius sanguinolentus). We used a laboratory‐controlled experiment with freshly caught fish designed to test for differences in visual cues only. Distance kept by the focal fish to each stimulus and frequency of bobbing and fin‐flicking were recorded. Triplefins kept greater distance from the stimulus compartment when a scorpionfish predator was visible. Bobbing was more frequent in the visual presence of a scorpionfish, but also shown toward the other stimuli. However, fin flicks were equally abundant across all stimuli. Both behaviors decreased in frequency over time suggesting that triplefin become gradually comfortable in a nonchanging new environment. We discuss why bobbing and fin‐flicking are not exclusive pursuit‐deterrent behaviors in this species, and propose additional nonexclusive functions such as enhancing depth perception by parallax motion (bobbing) or signaling vigilance (fin‐flicking).     

Tail autotomy works as a pre‐capture defense by deflecting attacks

Caudal autotomy is a dramatic antipredator adaptation where prey shed their tail in order to escape capture by a predator. The mechanism underlying the effectiveness of caudal autotomy as a pre‐capture defense has not been thoroughly investigated. We tested two nonexclusive hypotheses, that caudal autotomy works by providing the predator with a “consolation prize” that makes it break off the hunt to consume the shed tail, and the deflection hypothesis, where the autotomy event directs predator attacks to the autotomized tail enabling prey escape. Our experiment utilized domestic dogs Canis familiaris as model predator engaged to chase a snake‐like stimulus with a detachable tail. The tail was manipulated to vary in length (long versus short) and conspicuousness (green versus blue), with the prediction that dog attacks on the tail should increase with length under the consolation‐prize hypothesis and conspicuous color under the deflection hypothesis. The tail was attacked on 35% of trials, supporting the potential for pre‐capture autotomy to offer antipredator benefits. Dogs were attracted to the tail when it was conspicuously colored, but not when it was longer. This supports the idea that deflection of predator attacks through visual effects is the prime antipredator mechanism underlying the effectiveness of caudal autotomy as opposed to provision of a consolation prize meal.     

Similar yet different: differential response of a praying mantis to ant‐mimicking spiders

Batesian mimics typically dupe visual predators by resembling noxious or deadly model species. Ants are unpalatable and dangerous to many arthropod taxa, and are popular invertebrate models in mimicry studies. Ant mimicry by spiders, especially jumping spiders, has been studied and researchers have examined whether visual predators can distinguish between the ant model, spider mimic and spider non‐mimics. Tropical habitats harbour a diverse community of ants, their mimics and predators. In one such tripartite mimicry system, we investigated the response of an invertebrate visual predator, the ant‐mimicking praying mantis (Euantissa pulchra), to two related ant‐mimicking spider prey of the genus Myrmarachne, each closely mimicking its model ant species. We found that weaver ants (Oecophylla smaragdina) were much more aggressive than carpenter ants (Camponotus sericeus) towards the mantis. Additionally, mantids exhibited the same aversive response towards ants and their mimics. More importantly, mantids approached carpenter ant‐mimicking spiders significantly more than often that they approached weaver ant‐mimicking spiders. Thus, in this study, we show that an invertebrate predator, the praying mantis, can indeed discriminate between two closely related mimetic prey. The exact mechanism of the discrimination remains to be tested, but it is likely to depend on the level of mimetic accuracy by the spiders and on the aggressiveness of the ant model organism.     

Mimics without models: causes and consequences of allopatry in Batesian mimicry complexes

Batesian mimicry evolves when a palatable species (the ‘mimic’) co-opts a warning signal from a dangerous species (the ‘model’) and thereby deceives its potential predators. Longstanding theory predicts that this protection from predation should break down where the model is absent. Thus, mimics are expected to only co-occur with their model. Yet, many mimics violate this prediction and occur in areas where their model is absent. Here, we discuss the causes and consequences of such allopatric mimics. We also describe how these ‘rule-bending’ mimics provide critical insights into diverse topics ranging from how Batesian mimicry evolves to its possible role in speciation.