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1

Hlaváček, Antonín, Klára Daňková, Daniel Benda, Petr Bogusch, and Jiří Hadrava. "Batesian-Müllerian mimicry ring around the Oriental hornet (Vespa orientalis)." Journal of Hymenoptera Research 92 (August 31, 2022): 211–28. http://dx.doi.org/10.3897/jhr.92.81380.

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Mimicry is usually understood to be an adaptive resemblance between phylogenetically distant groups of species. In this study, we focus on Batesian and Müllerian mimicry, which are often viewed as a continuum rather than distinct phenomena, forming so-called Batesian-Müllerian mimicry rings. Despite potent defence and wide environmental niche of hornets, little attention has been paid to them as potential models in mimicry research. We propose a Batesian-Müllerian mimicry ring of the Oriental hornet (Vespa orientalis, Hymenoptera: Vespidae) consisting of eight species that coexist in the Mediterranean region. To reveal general ecological patterns, we reviewed their geographical distribution, phenology, and natural history. In accordance with the ‘model-first’ theory, Batesian mimics of this ring occurred later during a season than the Müllerian mimics. In the case of Batesian mimic Volucella zonaria (Diptera: Syrphidae), we presume that temperature-driven range expansion could lead to allopatry with its model, and, potentially, less accurate resemblance to an alternative model, the European hornet (Vespa crabro: Hymenoptera: Vespidae). Colour morphs of polymorphic species Cryptocheilus alternatus (Hymenoptera: Vespidae), Delta unguiculatum (Hymenoptera: Vespidae), Rhynchium oculatum (Hymenoptera: Vespidae), and Scolia erythrocephala (Hymenoptera: Scoliidae) appear to display distinct geographical distribution patterns, and this is possibly driven by sympatry with alternative models from the European hornet (Vespa crabro) complex. General coevolution patterns of models and mimics in heterogenous and temporally dynamic environments are discussed, based on observations of the proposed Oriental hornet mimicry ring.
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2

Hlaváček, Antonín, Klára Daňková, Daniel Benda, Petr Bogusch, and Jiří Hadrava. "Batesian-Müllerian mimicry ring around the Oriental hornet (Vespa orientalis)." Journal of Hymenoptera Research 92 (August 31, 2022): 211–28. https://doi.org/10.3897/jhr.92.81380.

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Mimicry is usually understood to be an adaptive resemblance between phylogenetically distant groups of species. In this study, we focus on Batesian and Müllerian mimicry, which are often viewed as a continuum rather than distinct phenomena, forming so-called Batesian-Müllerian mimicry rings. Despite potent defence and wide environmental niche of hornets, little attention has been paid to them as potential models in mimicry research. We propose a Batesian-Müllerian mimicry ring of the Oriental hornet (Vespa orientalis, Hymenoptera: Vespidae) consisting of eight species that coexist in the Mediterranean region. To reveal general ecological patterns, we reviewed their geographical distribution, phenology, and natural history. In accordance with the 'model-first' theory, Batesian mimics of this ring occurred later during a season than the Müllerian mimics. In the case of Batesian mimic Volucella zonaria (Diptera: Syrphidae), we presume that temperature-driven range expansion could lead to allopatry with its model, and, potentially, less accurate resemblance to an alternative model, the European hornet (Vespa crabro: Hymenoptera: Vespidae). Colour morphs of polymorphic species Cryptocheilus alternatus (Hymenoptera: Vespidae), Delta unguiculatum (Hymenoptera: Vespidae), Rhynchium oculatum (Hymenoptera: Vespidae), and Scolia erythrocephala (Hymenoptera: Scoliidae) appear to display distinct geographical distribution patterns, and this is possibly driven by sympatry with alternative models from the European hornet (Vespa crabro) complex. General coevolution patterns of models and mimics in heterogenous and temporally dynamic environments are discussed, based on observations of the proposed Oriental hornet mimicry ring.
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3

Jamie, Gabriel A. "Signals, cues and the nature of mimicry." Proceedings of the Royal Society B: Biological Sciences 284, no. 1849 (2017): 20162080. http://dx.doi.org/10.1098/rspb.2016.2080.

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‘Mimicry’ is used in the evolutionary and ecological literature to describe diverse phenomena. Many are textbook examples of natural selection's power to produce stunning adaptations. However, there remains a lack of clarity over how mimetic resemblances are conceptually related to each other. The result is that categories denoting the traditional subdivisions of mimicry are applied inconsistently across studies, hindering attempts at conceptual unification. This review critically examines the logic by which mimicry can be conceptually organized and analysed. It highlights the following three evolutionarily relevant distinctions. (i) Are the model's traits being mimicked signals or cues? (ii) Does the mimic signal a fitness benefit or fitness cost in order to manipulate the receiver's behaviour? (iii) Is the mimic's signal deceptive? The first distinction divides mimicry into two broad categories: ‘signal mimicry’ and ‘cue mimicry’. ‘Signal mimicry’ occurs when mimic and model share the same receiver, and ‘cue mimicry’ when mimic and model have different receivers or when there is no receiver for the model's trait. ‘Masquerade’ fits conceptually within cue mimicry. The second and third distinctions divide both signal and cue mimicry into four types each. These are the three traditional mimicry categories (aggressive, Batesian and Müllerian) and a fourth, often overlooked category for which the term ‘rewarding mimicry’ is suggested. Rewarding mimicry occurs when the mimic's signal is non-deceptive (as in Müllerian mimicry) but where the mimic signals a fitness benefit to the receiver (as in aggressive mimicry). The existence of rewarding mimicry is a logical extension of the criteria used to differentiate the three well-recognized forms of mimicry. These four forms of mimicry are not discrete, immutable types, but rather help to define important axes along which mimicry can vary.
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4

Srygley, Robert B. "Locomotor mimicry in Heliconius butterflies: contrast analyses of flight morphology and kinematics." Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences 354, no. 1380 (1999): 203–14. http://dx.doi.org/10.1098/rstb.1999.0372.

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Müllerian mimicry is a mutualism involving the evolutionary convergence of colour patterns of prey on a warning signal to predators. Behavioural mimicry presumably adds complexity to the signal and makes it more difficult for Batesian mimics to parasitize it. To date, no one has quantified behavioural mimicry in Müllerian mimicry groups. However, morphological similarities among members of mimicry groups suggested that pitching oscillations of the body and wing–beat frequency (WBF) might converge with colour pattern. I compared the morphology and kinematics of four Heliconius species, which comprised two mimicry pairs. Because the mimics arose from two distinct lineages, the relative contributions of mimicry and phylogeny to variation in the species' morphologies and kinematics were examined. The positions of the centre of body mass and centre of wing mass and wing shape diverged among species within lineages, and converged among species within mimicry groups. WBF converged within mimicry groups, and it was coupled with body pitching frequency. However, body–pitching frequency was too variable to distinguish mimicry groups. Convergence in WBF may be due, at least in part, to biomechanical consequences of similarities in wing length, wing shape or the centre of wing mass among co–mimics. Nevertheless, convergence in WBF among passion–vine butterflies serves as the first evidence of behavioural mimicry in a mutualistic context.
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5

Sherratt, Thomas N. "The evolution of Müllerian mimicry." Naturwissenschaften 95, no. 8 (2008): 681–95. http://dx.doi.org/10.1007/s00114-008-0403-y.

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6

Stoddard, Mary Caswell. "Mimicry and masquerade from the avian visual perspective." Current Zoology 58, no. 4 (2012): 630–48. http://dx.doi.org/10.1093/czoolo/58.4.630.

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Abstract Several of the most celebrated examples of visual mimicry, like mimetic eggs laid by avian brood parasites and palatable insects mimicking distasteful ones, involve signals directed at the eyes of birds. Despite this, studies of mimicry from the avian visual perspective have been rare, particularly with regard to defensive mimicry and masquerade. Defensive visual mimicry, which includes Batesian and Müllerian mimicry, occurs when organisms share a visual signal that functions to deter predators. Masquerade occurs when an organism mimics an inedible or uninteresting object, such as a leaf, stick, or pebble. In this paper, I present five case studies covering diverse examples of defensive mimicry and masquerade as seen by birds. The best-known cases of defensive visual mimicry typically come from insect prey, but birds themselves can exhibit defensive visual mimicry in an attempt to escape mobbing or dissuade avian predators. Using examples of defensive visual mimicry by both insects and birds, I show how quantitative models of avian color, luminance, and pattern vision can be used to enhance our understanding of mimicry in many systems and produce new hypotheses about the evolution and diversity of signals. Overall, I investigate examples of Batesian mimicry (1 and 2), Müllerian mimicry (3 and 4), and masquerade (5) as follows: 1) Polymorphic mimicry in African mocker swallowtail butterflies; 2) Cuckoos mimicking sparrowhawks; 3) Mimicry rings in Neotropical butterflies; 4) Plumage mimicry in toxic pitohuis; and 5) Dead leaf-mimicking butterflies and mantids.
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7

Hoyal Cuthill, Jennifer F., Nicholas Guttenberg, Sophie Ledger, Robyn Crowther, and Blanca Huertas. "Deep learning on butterfly phenotypes tests evolution’s oldest mathematical model." Science Advances 5, no. 8 (2019): eaaw4967. http://dx.doi.org/10.1126/sciadv.aaw4967.

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Traditional anatomical analyses captured only a fraction of real phenomic information. Here, we apply deep learning to quantify total phenotypic similarity across 2468 butterfly photographs, covering 38 subspecies from the polymorphic mimicry complex of Heliconius erato and Heliconius melpomene. Euclidean phenotypic distances, calculated using a deep convolutional triplet network, demonstrate significant convergence between interspecies co-mimics. This quantitatively validates a key prediction of Müllerian mimicry theory, evolutionary biology’s oldest mathematical model. Phenotypic neighbor-joining trees are significantly correlated with wing pattern gene phylogenies, demonstrating objective, phylogenetically informative phenome capture. Comparative analyses indicate frequency-dependent mutual convergence with coevolutionary exchange of wing pattern features. Therefore, phenotypic analysis supports reciprocal coevolution, predicted by classical mimicry theory but since disputed, and reveals mutual convergence as an intrinsic generator for the unexpected diversity of Müllerian mimicry. This demonstrates that deep learning can generate phenomic spatial embeddings, which enable quantitative tests of evolutionary hypotheses previously only testable subjectively.
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8

Ihalainen, Eira, Hannah M. Rowland, Michael P. Speed, Graeme D. Ruxton, and Johanna Mappes. "Prey community structure affects how predators select for Müllerian mimicry." Proceedings of the Royal Society B: Biological Sciences 279, no. 1736 (2012): 2099–105. http://dx.doi.org/10.1098/rspb.2011.2360.

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Müllerian mimicry describes the close resemblance between aposematic prey species; it is thought to be beneficial because sharing a warning signal decreases the mortality caused by sampling by inexperienced predators learning to avoid the signal. It has been hypothesized that selection for mimicry is strongest in multi-species prey communities where predators are more prone to misidentify the prey than in simple communities. In this study, wild great tits ( Parus major ) foraged from either simple (few prey appearances) or complex (several prey appearances) artificial prey communities where a specific model prey was always present. Owing to slower learning, the model did suffer higher mortality in complex communities when the birds were inexperienced. However, in a subsequent generalization test to potential mimics of the model prey (a continuum of signal accuracy), only birds that had foraged from simple communities selected against inaccurate mimics. Therefore, accurate mimicry is more likely to evolve in simple communities even though predator avoidance learning is slower in complex communities. For mimicry to evolve, prey species must have a common predator; the effective community consists of the predator's diet. In diverse environments, the limited diets of specialist predators could create ‘simple community pockets’ where accurate mimicry is selected for.
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9

Lev-Yadun, Simcha. "Müllerian mimicry in aposematic spiny plants." Plant Signaling & Behavior 4, no. 6 (2009): 482–83. http://dx.doi.org/10.4161/psb.4.6.8848.

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10

Speed, Michael P. "Mistakes not necessary for Müllerian mimicry." Nature 396, no. 6709 (1998): 323. http://dx.doi.org/10.1038/24519.

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11

Bosque, Renan Janke, Chaz Hyseni, Maria Luiza Gonçalves Santos, et al. "Müllerian mimicry and the coloration patterns of sympatric coral snakes." Biological Journal of the Linnean Society 135, no. 4 (2022): 645–51. http://dx.doi.org/10.1093/biolinnean/blab155.

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Abstract Coral snakes in the genus Micrurus are venomous, aposematic organisms that signal danger to predators through vivid coloration. Previous studies found that they serve as models to several harmless species of Batesian mimics. However, the extent to which Micrurus species engage in Müllerian mimicry remains poorly understood. We integrate detailed morphological and geographical distribution data to investigate if coral snakes are Müllerian mimics. We found that coloration is spatially structured and that Micrurus species tend to be more similar where they co-occur. Though long supposed, we demonstrate for the first time that coral snakes might indeed be Müllerian mimics as they show some convergence in coloration patterns. Additionally, we found that the length of red-coloured rings in Micrurus is conserved, even at large geographic scales. This finding suggests that bright red rings may be under more substantial stabilizing selection than other aspects of coloration and probably function as a generalized signal for deterring predators.
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12

Ezray, Briana D., Drew C. Wham, Carrie E. Hill, and Heather M. Hines. "Unsupervised machine learning reveals mimicry complexes in bumblebees occur along a perceptual continuum." Proceedings of the Royal Society B: Biological Sciences 286, no. 1910 (2019): 20191501. http://dx.doi.org/10.1098/rspb.2019.1501.

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Müllerian mimicry theory states that frequency-dependent selection should favour geographical convergence of harmful species onto a shared colour pattern. As such, mimetic patterns are commonly circumscribed into discrete mimicry complexes, each containing a predominant phenotype. Outside a few examples in butterflies, the location of transition zones between mimicry complexes and the factors driving mimicry zones has rarely been examined. To infer the patterns and processes of Müllerian mimicry, we integrate large-scale data on the geographical distribution of colour patterns of social bumblebees across the contiguous United States and use these to quantify colour pattern mimicry using an innovative, unsupervised machine-learning approach based on computer vision. Our data suggest that bumblebees exhibit geographically clustered, but sometimes imperfect colour patterns, and that mimicry patterns gradually transition spatially rather than exhibit discrete boundaries. Additionally, examination of colour pattern transition zones of three comimicking, polymorphic species, where active selection is driving phenotype frequencies, revealed that their transition zones differ in location within a broad region of poor mimicry. Potential factors influencing mimicry transition zone dynamics are discussed.
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13

Pfennig, David W., and David W. Kikuchi. "Competition and the evolution of imperfect mimicry." Current Zoology 58, no. 4 (2012): 608–19. http://dx.doi.org/10.1093/czoolo/58.4.608.

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Abstract Mimicry is widely used to exemplify natural selection’s power in promoting adaptation. Nonetheless, it has become increasingly clear that mimicry is frequently imprecise. Indeed, the phenotypic match is often poor between mimics and models in many Batesian mimicry complexes and among co-mimics in many Müllerian mimicry complexes. Here, we consider whether such imperfect mimicry represents an evolutionary compromise between predator-mediated selection favoring mimetic convergence on the one hand and competitively mediated selection favoring divergence on the other hand. Specifically, for mimicry to be effective, mimics and their models/co-mimics should occur together. Yet, co-occurring species that are phenotypically similar often compete for resources, successful reproduction, or both. As an adaptive response to minimize such costly interactions, interacting species may diverge phenotypically through an evolutionary process known as character displacement. Such divergence between mimics and their models/co-mimics may thereby result in imperfect mimicry. We review the various ways in which character displacement could promote imprecise mimicry, describe the conditions under which this process may be especially likely to produce imperfect mimicry, examine a possible case study, and discuss avenues for future research. Generally, character displacement may play an underappreciated role in fostering inexact mimicry.
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14

Sadahisa, Yagi, Hirowatari Toshiya, and Arita Yutaka. "A remarkable new species of the genus Teinotarsina (Lepidoptera, Sesiidae) from Okinawa-jima, Japan." ZooKeys 571 (March 7, 2016): 143–52. https://doi.org/10.3897/zookeys.571.7780.

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A new species of long-legged clearwing moth Teinotarsina aurantiaca Yagi, Hirowatari & Arita, sp. n. is described from Okinawa-jima, the Ryukyus, Japan. The species is distinguishable at a glance from other related congeners by the remarkable orange scales ornamenting many parts of the body, such as antennae, palpi, legs, and wings. We hypothesize that the species acquired these differences as a result of convergence with toxic species (Batesian mimicry) or other mimics (Müllerian mimicry).
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15

Ihalainen, Eira, Leena Lindström, Johanna Mappes, and Sari Puolakkainen. "Can experienced birds select for Müllerian mimicry?" Behavioral Ecology 19, no. 2 (2008): 362–68. http://dx.doi.org/10.1093/beheco/arm151.

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16

LANE, CHARLES, and MIRIAM ROTHSCHILD. "A case of Müllerian mimicry of sound." Proceedings of the Royal Entomological Society of London. Series A, General Entomology 40, no. 10-12 (2009): 156–58. http://dx.doi.org/10.1111/j.1365-3032.1965.tb00305.x.

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17

Speed, M. P. "Batesian, quasi-Batesian or Müllerian mimicry? Theory and data in mimicry Research." Evolutionary Ecology 13, no. 7-8 (1999): 755–76. http://dx.doi.org/10.1023/a:1010871106763.

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18

Muñoz-Ramírez, Carlos P., Pierre-Paul Bitton, Stéphanie M. Doucet, and Lacey L. Knowles. "Mimics here and there, but not everywhere: Müllerian mimicry in Ceroglossus ground beetles?" Biology Letters 12, no. 9 (2016): 20160429. http://dx.doi.org/10.1098/rsbl.2016.0429.

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The ground beetle genus Ceroglossus contains co-distributed species that show pronounced intraspecific diversity in the form of geographical colour morphs. While colour morphs among different species appear to match in some geographical regions, in others, there is little apparent colour matching. Mimicry is a potential explanation for covariation in colour patterns, but it is not clear whether the degree of sympatric colour matching is higher than expected by chance given the obvious mismatches among morphs in some regions. Here, we used reflectance spectrometry to quantify elytral coloration from the perspective of an avian predator to test whether colour similarity between species is, indeed, higher in sympatry. After finding no significant phylogenetic signal in the colour data, analyses showed strong statistical support for sympatric colour similarity between species despite the apparent lack of colour matching in some areas. We hypothesize Müllerian mimicry as the responsible mechanism for sympatric colour similarity in Ceroglossus and discuss potential explanations and future directions to elucidate why mimicry has not developed similar levels of interspecific colour resemblance across space.
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19

Llaurens, V., M. Joron, and S. Billiard. "Molecular mechanisms of dominance evolution in Müllerian mimicry." Evolution 69, no. 12 (2015): 3097–108. http://dx.doi.org/10.1111/evo.12810.

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20

Beatty, Christopher D., Kirsten Beirinckx, and Thomas N. Sherratt. "The evolution of müllerian mimicry in multispecies communities." Nature 431, no. 7004 (2004): 63–66. http://dx.doi.org/10.1038/nature02818.

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21

Speed, Michael P., Nichola J. Alderson, Christine Hardman, and Graeme D. Ruxton. "Testing Müllerian mimicry: an experiment with wild birds." Proceedings of the Royal Society of London. Series B: Biological Sciences 267, no. 1444 (2000): 725–31. http://dx.doi.org/10.1098/rspb.2000.1063.

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22

Kumazawa, Fuga, Takahiro Asami, Nariyuki Nakagiri, et al. "Population dynamics of Müllerian mimicry under interspecific competition." Ecological Modelling 220, no. 3 (2009): 424–29. http://dx.doi.org/10.1016/j.ecolmodel.2008.11.007.

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23

Merrill, Richard M., and Chris D. Jiggins. "Müllerian Mimicry: Sharing the Load Reduces the Legwork." Current Biology 19, no. 16 (2009): R687—R689. http://dx.doi.org/10.1016/j.cub.2009.07.008.

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24

Loeffler-Henry, Karl, and Thomas N. Sherratt. "A case for mutualistic deceptive mimicry." Biological Journal of the Linnean Society 133, no. 3 (2021): 853–62. http://dx.doi.org/10.1093/biolinnean/blaa219.

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Abstract It has long been understood that species that are profitable for predators to attack can gain protection if they resemble unprofitable species (Batesian mimicry), and that unprofitable species may face selection to evolve a common warning signal (Müllerian mimicry). Here we suggest that there may be widespread selection for another form of protective mimicry, so far unrecognized, that can arise even among profitable prey. Specifically, when predators adopt species-specific attack strategies, then co-occurring prey species that are caught in different ways may be selected to resemble one another. This is because the mimicry may increase the chance that the predator deploys an inappropriate attack strategy, thereby increasing the probability the prey will escape. We refer to this phenomenon as “mutualistic deceptive mimicry”, since the mimicry misleads the predator yet potentially benefits all co-mimics. We show that this hypothesis is quantitatively plausible. We then provide an empirical ‘proof of concept’ demonstrating that predators can learn to attack distinct prey types in specific ways and that this behaviour readily generates selection for mimicry. Finally, we discuss how this unrecognized form of mimicry fits into an earlier classification of protective mimicry and suggest a number of potential examples in the natural world.
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Chouteau, Mathieu, Kyle Summers, Victor Morales, and Bernard Angers. "Advergence in Müllerian mimicry: the case of the poison dart frogs of Northern Peru revisited." Biology Letters 7, no. 5 (2011): 796–800. http://dx.doi.org/10.1098/rsbl.2011.0039.

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Whether the evolution of similar aposematic signals in different unpalatable species (i.e. Müllerian mimicry) is because of phenotypic convergence or advergence continues to puzzle scientists. The poison dart frog Ranitomeya imitator provides a rare example in support of the hypothesis of advergence: this species was believed to mimic numerous distinct model species because of high phenotypic variability and low genetic divergence among populations. In this study, we test the evidence in support of advergence using a population genetic framework in two localities where R. imitator is sympatric with different model species, Ranitomeya ventrimaculata and Ranitomeya variabilis . Genetic analyses revealed incomplete sorting of mitochondrial haplotypes between the two model species. These two species are also less genetically differentiated than R. imitator populations on the basis of both mitochondrial and nuclear DNA comparisons. The genetic similarity between the model species suggests that they have either diverged more recently than R. imitator populations or that they are still connected by gene flow and were misidentified as different species. An analysis of phenotypic variability indicates that the model species are as variable as R. imitator . These results do not support the hypothesis of advergence by R. imitator . Although we cannot rule out phenotypic advergence in the evolution of Müllerian mimicry, this study reopens the discussion regarding the direction of the evolution of mimicry in the R. imitator system.
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Lev-Yadun, Simcha. "Müllerian and Batesian mimicry out, Darwinian and Wallacian mimicry in, for rewarding/rewardless flowers." Plant Signaling & Behavior 13, no. 6 (2018): e1480846. http://dx.doi.org/10.1080/15592324.2018.1480846.

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Llaurens, V., S. Billiard, and M. Joron. "The effect of dominance on polymorphism in Müllerian mimicry." Journal of Theoretical Biology 337 (November 2013): 101–10. http://dx.doi.org/10.1016/j.jtbi.2013.08.006.

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28

MACDOUGALL, ANGUS, and MARIAN STAMP DAWKINS. "Predator discrimination error and the benefits of Müllerian mimicry." Animal Behaviour 55, no. 5 (1998): 1281–88. http://dx.doi.org/10.1006/anbe.1997.0702.

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Raška, Jan, Jan Krajíček, Zuzana Bosáková, Pavel Štys, and Alice Exnerová. "Larvae of pyrrhocorid true bugs are not to spiders' taste: putative Müllerian mimicry." Biological Journal of the Linnean Society 129 (October 21, 2019): 199–212. https://doi.org/10.1093/biolinnean/blz174.

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Raška, Jan, Krajíček, Jan, Bosáková, Zuzana, Štys, Pavel, Exnerová, Alice (2020): Larvae of pyrrhocorid true bugs are not to spiders' taste: putative Müllerian mimicry. Biological Journal of the Linnean Society 129: 199-212, DOI: 10.1093/biolinnean/blz174
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Willmott, Keith R., Julia C. Robinson Willmott, Marianne Elias, and Chris D. Jiggins. "Maintaining mimicry diversity: optimal warning colour patterns differ among microhabitats in Amazonian clearwing butterflies." Proceedings of the Royal Society B: Biological Sciences 284, no. 1855 (2017): 20170744. http://dx.doi.org/10.1098/rspb.2017.0744.

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Mimicry is one of the best-studied examples of adaptation, and recent studies have provided new insights into the role of mimicry in speciation and diversification. Classical Müllerian mimicry theory predicts convergence in warning signal among protected species, yet tropical butterflies are exuberantly diverse in warning colour patterns, even within communities. We tested the hypothesis that microhabitat partitioning in aposematic butterflies and insectivorous birds can lead to selection for different colour patterns in different microhabitats and thus help maintain mimicry diversity. We measured distribution across flight height and topography for 64 species of clearwing butterflies (Ithomiini) and their co-mimics, and 127 species of insectivorous birds, in an Amazon rainforest community. For the majority of bird species, estimated encounter rates were non-random for the two most abundant mimicry rings. Furthermore, most butterfly species in these two mimicry rings displayed the warning colour pattern predicted to be optimal for anti-predator defence in their preferred microhabitats. These conclusions were supported by a field trial using butterfly specimens, which showed significantly different predation rates on colour patterns in two microhabitats. We therefore provide the first direct evidence to support the hypothesis that different mimicry patterns can represent stable, community-level adaptations to differing biotic environments.
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MAREK, PAUL E., JACKSON C. MEANS, and DEREK A. HENNEN. "Apheloria polychroma, a new species of millipede from the Cumberland Mountains (Polydesmida: Xystodesmidae)." Zootaxa 4375, no. 3 (2018): 409. http://dx.doi.org/10.11646/zootaxa.4375.3.7.

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Millipedes of the genus Apheloria Chamberlin, 1921 occur in temperate broadleaf forests throughout eastern North America and west of the Mississippi River in the Ozark and Ouachita Mountains. Chemically defended with toxins made up of cyanide and benzaldehyde, the genus is part of a community of xystodesmid millipedes that compose several Müllerian mimicry rings in the Appalachian Mountains. We describe a model species of these mimicry rings, Apheloria polychroma n. sp., one of the most variable in coloration of all species of Diplopoda with more than six color morphs, each associated with a separate mimicry ring.
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Halbritter, Dale, Johnalyn Gordon, Kandy Keacher, Michael Avery, and Jaret Daniels. "Evaluating an Alleged Mimic of the Monarch Butterfly: Neophasia (Lepidoptera: Pieridae) Butterflies are Palatable to Avian Predators." Insects 9, no. 4 (2018): 150. http://dx.doi.org/10.3390/insects9040150.

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Some taxa have adopted the strategy of mimicry to protect themselves from predation. Butterflies are some of the best representatives used to study mimicry, with the monarch butterfly, Danaus plexippus (Lepidoptera: Nymphalidae) a well-known model. We are the first to empirically investigate a proposed mimic of the monarch butterfly: Neophasia terlooii, the Mexican pine white butterfly (Lepidoptera: Pieridae). We used captive birds to assess the palatability of N. terlooii and its sister species, N. menapia, to determine the mimicry category that would best fit this system. The birds readily consumed both species of Neophasia and a palatable control species but refused to eat unpalatable butterflies such as D. plexippus and Heliconius charithonia (Lepidoptera: Nymphalidae). Given some evidence for mild unpalatability of Neophasia, we discuss the results considering modifications to classic mimicry theory, i.e., a palatability-based continuum between Batesian and Müllerian mimicry, with a quasi-Batesian intermediate. Understanding the ecology of Neophasia in light of contemporary and historical sympatry with D. plexippus could shed light on the biogeography of, evolution of, and predation pressure on the monarch butterfly, whose migration event has become a conservation priority.
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33

Nedvěd, Oldřich, Amir Biranvand, Jahanshir Shakarami, and Derya Şenal. "Potential Müllerian Mimicry between Adalia bipunctata (Linnaeus) and Oenopia conglobata (Linnaeus) (Coleoptera: Coccinellidae) in Iran." Coleopterists Bulletin 74, no. 1 (2020): 161–67. https://doi.org/10.1649/0010-065X-74.1.161.

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Nedvěd, Oldřich, Biranvand, Amir, Shakarami, Jahanshir, Şenal, Derya (2020): Potential Müllerian Mimicry between Adalia bipunctata (Linnaeus) and Oenopia conglobata (Linnaeus) (Coleoptera: Coccinellidae) in Iran. The Coleopterists Bulletin 74 (1): 161-167, DOI: 10.1649/0010-065X-74.1.161, URL: http://dx.doi.org/10.1649/0010-065x-74.1.161
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34

Perger, Robert. "Are goldish spiders able to teach naïve predators to avoid bullet ants? A possible case of Müllerian mimicry in spiders and ants." Journal of Natural History 55, no. 9-10 (2021): 625–27. https://doi.org/10.1080/00222933.2021.1914450.

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Perger, Robert (2021): Are goldish spiders able to teach naïve predators to avoid bullet ants? A possible case of Müllerian mimicry in spiders and ants. Journal of Natural History 55 (9-10): 625-627, DOI: 10.1080/00222933.2021.1914450, URL: http://dx.doi.org/10.1080/00222933.2021.1914450
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35

Kapan, Durrell D. "Three-butterfly system provides a field test of müllerian mimicry." Nature 409, no. 6818 (2001): 338–40. http://dx.doi.org/10.1038/35053066.

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36

IHALAINEN, E., L. LINDSTRÖM, and J. MAPPES. "Investigating Müllerian mimicry: predator learning and variation in prey defences." Journal of Evolutionary Biology 20, no. 2 (2006): 780–91. http://dx.doi.org/10.1111/j.1420-9101.2006.01234.x.

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37

Sasaki, Akira, Isao Kawaguchi, and Akira Yoshimori. "Spatial Mosaic and Interfacial Dynamics in a Müllerian Mimicry System." Theoretical Population Biology 61, no. 1 (2002): 49–71. http://dx.doi.org/10.1006/tpbi.2001.1552.

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38

Barber, Jesse R., and William E. Conner. "Acoustic mimicry in a predator–prey interaction." Proceedings of the National Academy of Sciences 104, no. 22 (2007): 9331–34. https://doi.org/10.5281/zenodo.14820946.

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(Uploaded by Plazi for the Bat Literature Project) Mimicry of visual warning signals is one of the keystone concepts in evolutionary biology and has received substantial research attention. By comparison, acoustic mimicry has never been rigorously tested. Visualizing bat–moth interactions with high-speed, infrared videography, we provide empirical evidence for acoustic mimicry in the ultrasonic warning sounds that tiger moths produce in response to echolocating bats. Two species of sound-producing tiger moths were offered successively to naïve, free-flying red and big brown bats. Noctuid and pyralid moth controls were also offered each night. All bats quickly learned to avoid the noxious tiger moths first offered to them, associating the warning sounds with bad taste. They then avoided the second sound-producing species regardless of whether it was chemically protected or not, verifying both Müllerian and Batesian mimicry in the acoustic modality. A subset of the red bats subsequently discovered the palatability of the Batesian mimic, demonstrating the powerful selective force these predators exert on mimetic resemblance. Given these results and the widespread presence of tiger moth species and other sound-producing insects that respond with ultrasonic clicks to bat attack, acoustic mimicry complexes are likely common components of the acoustic landscape.
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39

Garg, Kritika M., Katerina Sam, Balaji Chattopadhyay, et al. "Gene Flow in the Müllerian Mimicry Ring of a Poisonous Papuan Songbird Clade (Pitohui; Aves)." Genome Biology and Evolution 11, no. 8 (2019): 2332–43. http://dx.doi.org/10.1093/gbe/evz168.

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Abstract Müllerian mimicry rings are remarkable symbiotic species assemblages in which multiple members share a similar phenotype. However, their evolutionary origin remains poorly understood. Although gene flow among species has been shown to generate mimetic patterns in some Heliconius butterflies, mimicry is believed to be due to true convergence without gene flow in many other cases. We investigated the evolutionary history of multiple members of a passerine mimicry ring in the poisonous Papuan pitohuis. Previous phylogenetic evidence indicates that the aposematic coloration shared by many, but not all, members of this genus is ancestral and has only been retained by members of the mimicry ring. Using a newly assembled genome and thousands of genomic DNA markers, we demonstrate gene flow from the hooded pitohui (Pitohui dichrous) into the southern variable pitohui (Pitohui uropygialis), consistent with shared patterns of aposematic coloration. The vicinity of putatively introgressed loci is significantly enriched for genes that are important in melanin pigment expression and toxin resistance, suggesting that gene flow may have been instrumental in the sharing of plumage patterns and toxicity. These results indicate that interspecies gene flow may be a more general mechanism in generating mimicry rings than hitherto appreciated.
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40

Winters, Anne E., Nerida G. Wilson, Cedric P. van den Berg, et al. "Toxicity and taste: unequal chemical defences in a mimicry ring." Proceedings of the Royal Society B: Biological Sciences 285, no. 1880 (2018): 20180457. http://dx.doi.org/10.1098/rspb.2018.0457.

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Mimicry of warning signals is common, and can be mutualistic when mimetic species harbour equal levels of defence (Müllerian), or parasitic when mimics are undefended but still gain protection from their resemblance to the model (Batesian). However, whether chemically defended mimics should be similar in terms of toxicity (i.e. causing damage to the consumer) and/or unpalatability (i.e. distasteful to consumer) is unclear and in many studies remains undifferentiated. In this study, we investigated the evolution of visual signals and chemical defences in a putative mimicry ring of nudibranch molluscs. First, we demonstrated that the appearance of a group of red spotted nudibranchs molluscs was similar from the perspective of potential fish predators using visual modelling and pattern analysis. Second, using phylogenetic reconstruction, we demonstrated that this colour pattern has evolved multiple times in distantly related individuals. Third, we showed that these nudibranchs contained different chemical profiles used for defensive purposes. Finally, we demonstrated that although levels of distastefulness towards Palaemon shrimp remained relatively constant between species, toxicity levels towards brine shrimp varied significantly. We highlight the need to disentangle toxicity and taste when considering chemical defences in aposematic and mimetic species, and discuss the implications for aposematic and mimicry signal evolution.
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41

Skelhorn, John, and Candy Rowe. "Tasting the difference: do multiple defence chemicals interact in Müllerian mimicry?" Proceedings of the Royal Society B: Biological Sciences 272, no. 1560 (2005): 339–45. http://dx.doi.org/10.1098/rspb.2004.2953.

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42

Balogh, Alexandra C. V., and Olof Leimar. "Müllerian mimicry: an examination of Fisher's theory of gradual evolutionary change." Proceedings of the Royal Society B: Biological Sciences 272, no. 1578 (2005): 2269–75. http://dx.doi.org/10.1098/rspb.2005.3227.

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43

Mallet, James. "Causes and Consequences of a Lack of Coevolution in Müllerian mimicry." Evolutionary Ecology 13, no. 7-8 (1999): 777–806. http://dx.doi.org/10.1023/a:1011060330515.

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44

Balogh, Alexandra Catherine Victoria, Gabriella Gamberale-Stille, Birgitta Sillén Tullberg, and Olof Leimar. "FEATURE THEORY AND THE TWO-STEP HYPOTHESIS OF MÜLLERIAN MIMICRY EVOLUTION." Evolution 64, no. 3 (2010): 810–22. http://dx.doi.org/10.1111/j.1558-5646.2009.00852.x.

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45

Sherratt, Thomas N. "Spatial mosaic formation through frequency-dependent selection in Müllerian mimicry complexes." Journal of Theoretical Biology 240, no. 2 (2006): 165–74. http://dx.doi.org/10.1016/j.jtbi.2005.09.017.

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46

Páez, Erika, Janne K. Valkonen, Keith R. Willmott, Pável Matos-Maraví, Marianne Elias, and Johanna Mappes. "Hard to catch: experimental evidence supports evasive mimicry." Proceedings of the Royal Society B: Biological Sciences 288, no. 1946 (2021): 20203052. http://dx.doi.org/10.1098/rspb.2020.3052.

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Most research on aposematism has focused on chemically defended prey, but the signalling difficulty of capture remains poorly explored. Similar to classical Batesian and Müllerian mimicry related to distastefulness, such ‘evasive aposematism' may also lead to convergence in warning colours, known as evasive mimicry. A prime candidate group for evasive mimicry areAdelphabutterflies, which are agile insects and show remarkable colour pattern convergence. We tested the ability of naive blue tits to learn to avoid and generalizeAdelphawing patterns associated with the difficulty of capture and compared their response to that of birds that learned to associate the same wing patterns with distastefulness. Birds learned to avoid all wing patterns tested and generalized their aversion to other prey to some extent, but learning was faster with evasive prey compared to distasteful prey. Our results on generalization agree with longstanding observations of striking convergence in wing colour patterns amongAdelphaspecies, since, in our experiments, perfect mimics of evasive and distasteful models were always protected during generalization and suffered the lowest attack rate. Moreover, generalization on evasive prey was broader compared to that on distasteful prey. Our results suggest that being hard to catch may deter predators at least as effectively as distastefulness. This study provides empirical evidence for evasive mimicry, a potentially widespread but poorly understood form of morphological convergence driven by predator selection.
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47

Ihalainen, Eira, Leena Lindström, Johanna Mappes, and Sari Puolakkainen. "Butterfly effects in mimicry? Combining signal and taste can twist the relationship of Müllerian co-mimics." Behavioral Ecology and Sociobiology 62, no. 8 (2008): 1267–76. http://dx.doi.org/10.1007/s00265-008-0555-y.

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48

Van Belleghem, Steven M., Paola A. Alicea Roman, Heriberto Carbia Gutierrez, Brian A. Counterman, and Riccardo Papa. "Perfect mimicry between Heliconius butterflies is constrained by genetics and development." Proceedings of the Royal Society B: Biological Sciences 287, no. 1931 (2020): 20201267. http://dx.doi.org/10.1098/rspb.2020.1267.

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Müllerian mimicry strongly exemplifies the power of natural selection. However, the exact measure of such adaptive phenotypic convergence and the possible causes of its imperfection often remain unidentified. Here, we first quantify wing colour pattern differences in the forewing region of 14 co-mimetic colour pattern morphs of the butterfly species Heliconius erato and Heliconius melpomene and measure the extent to which mimicking colour pattern morphs are not perfectly identical. Next, using gene-editing CRISPR/Cas9 KO experiments of the gene WntA , which has been mapped to colour pattern diversity in these butterflies, we explore the exact areas of the wings in which WntA affects colour pattern formation differently in H. erato and H. melpomene. We find that, while the relative size of the forewing pattern is generally nearly identical between co-mimics, the CRISPR/Cas9 KO results highlight divergent boundaries in the wing that prevent the co-mimics from achieving perfect mimicry. We suggest that this mismatch may be explained by divergence in the gene regulatory network that defines wing colour patterning in both species, thus constraining morphological evolution even between closely related species.
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49

Baxter, Simon W., Riccardo Papa, Nicola Chamberlain, et al. "Convergent Evolution in the Genetic Basis of Müllerian Mimicry in Heliconius Butterflies." Genetics 180, no. 3 (2008): 1567–77. http://dx.doi.org/10.1534/genetics.107.082982.

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50

Wright, Jeremy J. "CONSERVATIVE COEVOLUTION OF MÜLLERIAN MIMICRY IN A GROUP OF RIFT LAKE CATFISH." Evolution 65, no. 2 (2010): 395–407. http://dx.doi.org/10.1111/j.1558-5646.2010.01149.x.

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