An overview of the relationships between mimicry and crypsis

There have been many different and conflicting definitions of mimicry. Some of the definitions of mimicry include crypsis and others do not. Each definition includes different groups of phenomena and uses different criteria to distinguish mimetic from non-mimetic phenomena. The confusion is eliminated by a consideration of the criteria of all definitions. This shows that there are in fact three major criteria dividing six phenomona, rather than a single dichotomy between mimicry and crypsis (Table 2). The criteria are defined by the results of a mistake in discrimination between the model and mimìc: (a) the mistake does or does not depend upon relationship between mimic and background; (b) the mistake has or has no effect on the population dynamics or evolution of the model and (c) the mistake affects dynamics or evolution of one or of many models. The main reason for the contusion about mimicry and crypsis is that each author's definition includes differing and partially overlapping subsets of the six classes: crypsis; masquerade; Batesism; Müllerism; polymorphism and convergence.     

昆虫拟态的历史发展

昆虫的拟态理论是由英国自然学家Bates于1862年提出的,Fisher称其为"达尔文后自然选择最重要的依据之一".大量的科学研究表明,昆虫的拟态行为最晚出现在石炭纪,自那时起昆虫与捕食者、昆虫与植物之间开始出现了共同的演变和进化.拟态的模仿方式一般包括颜色、花纹以及形态,但是也可以单指行为方面,且拟态大部分情况下可能模仿的是一个动物群体或者只是另外一种动物身上的一部分.拟态包括多种定义,不同的定义之间用小同的标准来区分拟态现象和非拟态现象,如贝茨氏拟态、缪勒氏拟态、侵略性拟态和瓦曼氏拟态等.本文从其中广义拟态的角度,对当前不同类群昆虫化石中拟态现象的研究进展进行了简要总结.     

Understanding Mimicry – with Special Reference to Vocal Mimicry

The term mimicry was introduced to biology in 1862 by Henry Walter Bates in his evolutionary explanation of deceptive communication in nature, based on a three‐part interaction system of a mimicked organism or object (called model), a mimicking organism (called mimic), and one or more organisms as selecting agents. Bates gave two incongruous definitions of mimicry: one from the viewpoint of a natural agent that selects for, and in consequence is deceived by, the close resemblance of a toxic model's warning signal and the similar appearance of a palatable mimic, and another one from the viewpoint of a human taxonomist who under an evolutionary aspect focuses on convergent resemblance between model and mimic. Later definitions of Müllerian (F. Müller), arithmetic (A. Wallace) and social (M. Moynihan) mimicry abolish deception in the natural selecting agent, rely on the convergence criterion alone, fuse the roles of model and mimic but have to accept a mix of homologous and convergent resemblance amongst them for a functional explanation. The definition of vocal mimicry (E. Armstrong) refers to a learned resemblance between mimic and heterospecific model by character duplication (no convergence), so far without known (deceived or not deceived) natural selecting agents. It excludes Batesian vocal mimicry. The functional ethological understanding of mimicry as a tripartite communication system (W. Wickler) is consistent with Bates' concept and accepts deception as key element of Batesian mimicry beyond homologous and convergent resemblances. Deception is seen as caused by the divergence between a sign and its meaning for the natural selecting agent. This understanding covers mimicry in all behaviour domains, provides a generally applicable definition of mimic and model so far missing in any mimicry concept, and it distinguishes – still in line with Henry Bates – cultural from genetically determined model‐mimic‐resemblance; this applies to vocal mimicry in particular. Convergently evolved model‐mimic‐resemblance, not essential in Batesian mimicry but mandatory for its alternatives, marks a fundamental distinction between Batesian mimicry (including Mimesis) and all other conceptualized mimicries and accounts for the non‐existence of a unified meaning of the term mimicry. However, character convergence does not help to explain the mere existence of mimicry phenomena and is irrelevant for their permanence in nature. I therefore propose to remove the convergence argument from any mimicry definition.     

On the definition of mimicry

An operational distinction between crypsis and mimicry is made in terms of the cognitive and perceptual systems of signal-receivers. Cryptic organisms specialize in generating information of the type not attended to or filtered out (reference frame) by the receivers, whereas mimetic organisms specialize in producing information (signals) of the type sought out by and of interest to a receiver. Mimicry is defined in terms of a system of three living organisms, model, mimic and operator (signal-receiver), in which the mimic gains in fitness by the operator identifying it with the model. Some advantages and applications of the definition are briefly discussed.     

The evolution of Batesian mimicry within the North American Asidini (Coleoptera: Tenebrionidae)

The asidine darkling beetles (Coleoptera: Tenebrionidae: Asidini) are a diverse tribe of flightless tenebrionids found in many arid and sub‐arid habitats around the world. The 263 currently described North American species are contained in ten genera, all of which are restricted to the western half of the continent. The Asidini, like all members of the subfamily Pimeliinae, lack defensive glands. Instead, several phenotypic traits occur within the tribe that may help limit predation. These include the contrasting defensive strategies of crypsis, through either background matching or pattern disruption, and Batesian mimicry of the chemically defended genus Eleodes. Dorsal elytral morphology was assessed between 53 North American asidine species and 13 common Eleodes model species using multiple methodologies to assess similarities between species in the two groups that might indicate mimetic relationships. A phylogeny of the North American asidines is used to map the occurrence of differing defensive strategies within the tribe. Crypsis is reconstructed as the ancestral state, with two origins for Batesian mimicry and multiple reversals. The combination of strongly to weakly cryptic species and varying levels of mimetic fidelity to Eleodes model species make the asidines a promising lineage upon which to further explore the evolution of defensive phenotypes.     

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.     

Cryptic cuckoo eggs hide from competing cuckoos

Interspecific arms races between cuckoos and their hosts have produced remarkable examples of mimicry, with parasite eggs evolving to match host egg appearance and so evade removal by hosts. Certain bronze-cuckoo species, however, lay eggs that are cryptic rather than mimetic. These eggs are coated in a low luminance pigment that camouflages them within the dark interiors of hosts'' nests. We investigated whether cuckoo egg crypsis is likely to have arisen from the same coevolutionary processes known to favour egg mimicry. We added high and low luminance-painted eggs to the nests of large-billed gerygones (Gerygone magnirostris), a host of the little bronze-cuckoo (Chalcites minutillus). Gerygones rarely rejected either egg type, and did not reject natural cuckoo eggs. Cuckoos, by contrast, regularly removed an egg from clutches before laying their own and were five times more likely to remove a high luminance model than its low luminance counterpart. Given that we found one-third of all parasitized nests were exploited by multiple cuckoos, our results suggest that competition between cuckoos has been the key selective agent for egg crypsis. In such intraspecific arms races, crypsis may be favoured over mimicry because it can reduce the risk of egg removal to levels below chance.     

A relatively cheap photographic film eye-piece grid

A ‘signaller’ (for example, a prey) can avoid detection by a ‘receiver’ (for example, a predator) if its visual signals are difficult to separate from the background. There are two ways by which signallers match the coloration of their background— ‘crypsis’ and ‘masquerade’. We draw attention to the relationships between these two phenomena and how they differ from Batesian mimicry, another adaptation against detection.     

Model toxin level does not directly influence the evolution of mimicry in the salamander <Emphasis Type="Italic">Plethodon cinereus</Emphasis>

The resemblance between palatable mimics and unpalatable models in Batesian mimicry systems is tempered by many factors, including the toxicity of the model species. Model toxicity is thought to influence both the occurrence of mimicry and the evolution of mimetic phenotypes, such that mimicry is most likely to persist when models are particularly toxic. Additionally, model toxicity may influence the evolution of mimetic phenotype by allowing inaccurate mimicry to evolve through a mechanism termed ‘relaxed selection’. We tested these hypotheses in a salamander mimicry system between the model Notophthalmus viridescens and the mimic Plethodon cinereus, in which N. viridescens toxicity takes the form of tetrodotoxin. Surprisingly, though we discovered geographic variation in model toxin level, we found no support for the hypotheses that model toxicity directly influences either the occurrence of mimicry or the evolution of mimic phenotype. Instead, a link between N. viridescens size and toxicity may indirectly lead to relaxed selection in this mimicry system. Additionally, limitations of predator perception or variation in the rate of phenotypic evolution of models and mimics may account for the evolution of imperfect mimicry in this salamander species. Finally, variation in predator communities among localities or modern changes in environmental conditions may contribute to the patchy occurrence of mimicry in P. cinereus.     

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.     

Crypsis hypothesis as an explanation for evolution of impermeable coats in seeds is anecdotal

Impermeable seed/fruit coat, i.e. physical dormancy (PY) occurring in seeds of many genera of 19 angiosperm plant families has been traditionally viewed as a form of dormancy that regulates germination timing. However, this view was recently challenged by an alternative explanation claiming that the impermeable seed coat evolved as a coping mechanism to escape predators, i.e. crypsis hypothesis. Here, I wish to call for more careful attention on crypsis as an evolutionary factor because (1) information of volatile compounds is not known in PY families except Fabaceace; (2) impermeability is not induced until the moisture content of the seeds drops below species-specific threshold suggesting that drying determines development of impermeable seed coats; (3) the crypsis hypothesis does not explain the year-to-year or between site variations in proportions of impermeable seeds produced by plants; and (4) dry seeds of species from non-PY families also do not emit volatile compounds, and do not develop impermeable seed coats. For these reasons, it appears crypsis might be an exaptation, i.e., a trait that performs a function for which it was not originally evolved. Based on the available evidence, it is suggested that climate drying might have resulted in evolution of impermeable seed coats.     

Batesian insect-insect mimicry-related explosive radiation of ancient alienopterid cockroaches

Batesian mimicry is a relationship in which a harmful organism (the model) is mimicked by a harmless organism (the mimic), which gains protection because predators mistake it for the model. It is the most widely studied of mimicry complexes and has undoubtedly played an important role in the speciation of various animals especially insects. However, little is known about the early evolution of this important behavior and its evolutionary significance owing to a dearth of paleontological records. Here we report several specialized representatives of the family Alienopteridae from the Early Cretaceous of Brazil, mid-Cretaceous Burmite, and the Eocene of the USA. They exhibit unique morphological adaptations for wasp and ant mimicry and represent one of the oldest evidence of Batesian mimicry in the insect fossil record. Our findings reveal at least 65-million-year coevolution between extinct alienopterids and aculeates. Phylogenetic Bayesian network analysis houses Alienopteridae within Umenocoleidae explosively radiating ~127 Ma. Alienopteridae is the only Mesozoic-type cockroach family which passed K/Pg.     

Microbial Maintenance: A Critical Review on Its Quantification

Microbial maintenance is an important concept in microbiology. Its quantification, however, is a subject of continuous debate, which seems to be caused by (1) its definition, which includes nongrowth components other than maintenance; (2) the existence of partly overlapping concepts; (3) the evolution of variables as constants; and (4) the neglect of cell death in microbial dynamics. The two historically most important parameters describing maintenance, the specific maintenance rate and the maintenance coefficient, are based on partly different nongrowth components. There is thus no constant relation between these parameters and previous equations on this subject are wrong. In addition, the partial overlap between these parameters does not allow the use of a simple combination of these parameters. This also applies for combinations of a threshold concentration with one of the other estimates of maintenance. Maintenance estimates should ideally explicitly describe each nongrowth component. A conceptual model is introduced that describes their relative importance and reconciles the various concepts and definitions. The sensitivity of maintenance on underlying components was analyzed and indicated that overall maintenance depends nonlinearly on relative death rates, relative growth rates, growth yield, and endogenous metabolism. This quantitative sensitivity analysis explains the felt need to develop growth-dependent adaptations of existing maintenance parameters, and indicates the importance of distinguishing the various nongrowth components. Future experiments should verify the sensitivity of maintenance components under cellular and environmental conditions.     

Evolution of color variation in dragon lizards: quantitative tests of the role of crypsis and local adaptation

Abstract Many animal species display striking color differences with respect to geographic location, sex, and body region. Traditional adaptive explanations for such complex patterns invoke an interaction between selection for conspicuous signals and natural selection for crypsis. Although there is now a substantial body of evidence supporting the role of sexual selection for signaling functions, quantitative studies of crypsis remain comparatively rare. Here, we combine objective measures of coloration with information on predator visual sensitivities to study the role of crypsis in the evolution of color variation in an Australian lizard species complex (Ctenophorus decresii). We apply a model that allows us to quantify crypsis in terms of the visual contrast of the lizards against their natural backgrounds, as perceived by potential avian predators. We then use these quantitative estimates of crypsis to answer the following questions. Are there significant differences in crypsis conspicuousness among populations? Are there significant differences in crypsis conspicuousness between the sexes? Are body regions “exposed” to visual predators more cryptic than “hidden” body regions? Is there evidence for local adaptation with respect to crypsis against different substrates? In general, our results confirmed that there are real differences in crypsis conspicuousness both between populations and between sexes; that exposed body regions were significantly more cryptic than hidden ones, particularly in females; and that females, but not males, are more cryptic against their own local background than against the background of other populations. Body regions that varied most in contrast between the sexes and between populations were also most conspicuous and are emphasized by males during social and sexual signaling. However, results varied with respect to the aspect of coloration studied. Results based on chromatic contrast (“hue’ of color) provided better support for the crypsis hypothesis than did results based on achromatic contrast (“brightness’ of color). Taken together, these results support the view that crypsis plays a substantial role in the evolution of color variation and that color patterns represent a balance between the need for conspicuousness for signaling and the need for crypsis to avoid predation.     

Optimization of cryptic coloration in heterogeneous habitats

We present a theoretical approach to the optimization of crypsis in heterogeneous habitats. Our model habitat consists of two different microhabitats, and the optimal combination of crypsis in the microhabitats is supposed to maximize the probability of escaping detection by a predator. The probability of escaping detection for a prey is a function of: (i)degree of crypsis, (ii) probability of occurrence in the microhabitats and (iii) probability of encountering a predator in the microhabitats. Because crypsis is background-specific there is a trade-off between crypsis in two visually different microhabitats. Depending on the nature of the trade-off, the optimal coloration is either a compromise between the requirements of the differing microhabitats or entirely adapted to only one of them. An increased risk of predation in one of the microhabitats favours increased crypsis in that microhabitat. Because the trade-off constrains possible optimal solutions, it is not possible to predict the optimal coloration only from factors (i)-(iii). However, habitat choice may fundamentally change the situation. If minimizing predation risk does not incur any costs, the prey should exclusively prefer the microhabitat where it has a lower probability of encountering a predator and better crypsis. The implications of these results for variation in cryptic coloration and polymorphism are discussed.     

Perspectives and Debates: Mimicry,Signalling and Co‐Evolution (Commentary on Wolfgang Wickler – Understanding Mimicry – With special reference to vocal mimicry)

In his stimulating discussion, Wolfgang Wickler criticizes fuzzy usage of term mimicry by drawing attention to its original definition by H. Bates. Mimicry refers to functional ‘model–mimic–selecting agent’ trinity (with varying number of species involved) when the selecting agent (i.e. signal receiver) responds similarly to mimic and model to the advantage of the mimic. Concurring with Wickler I argue that convergence is neither necessary nor sufficient to support similarity as evidence for mimicry and that it is artificial and unproductive to classify mimicry with respect to ontogeny (innate vs. learned similarity) or model species identity (learning from conspecifics vs. heterospecifics). Using butterfly ‘eye’‐spots, I argue that just identifying each of the supposed model, the mimic and the selective agent, and even demonstrating that mimic‐model similarity affects the agent's behaviour, provides no conclusive evidence for mimicry. Even a demonstration that the mimic benefits from receiver response may not provide conclusive evidence for mimicry. Using avian brood parasite–host egg and nestling mimicry, I emphasize that without experimental manipulation of the hypothesized mimetic traits, it is impossible to test the mimicry hypothesis robustly. Due to fundamental constraints on human perception, some cases of mimicry may in fact be just a by‐product of human inability to perceive relevant differences between animal phenotypes (what is similar for human eye, nose or ear may not be viewed, smelled or heard as similar for relevant animal observers), whereas many cases of real mimicry may escape our attention from the same reason (‘hidden’ mimicry). Surprisingly, the same mimetic phenotype may show completely different effects on selective agents under different ecological circumstances. Finally, relatively dissimilar species may be more mimetic than highly similar model–mimic pairs because mimicry may be more fruitfully understood as a co‐evolutionary process rather than a similarity.     

What kind of signals do mimetic tiger moths send? A phylogenetic test of wasp mimicry systems (Lepidoptera: Arctiidae: Euchromiini).

Mimicry has been examined in field and laboratory studies of butterflies and its evolutionary dynamics have been explored in computer simulations. Phylogenetic studies examining the evolution of mimicry, however, are rare. Here, the phylogeny of wasp-mimicking tiger moths, the Sphecosoma group, was used to test evolutionary predictions of computer simulations of conventional Müllerian mimicry and quasi-Batesian mimicry dynamics. We examined whether mimetic traits evolved individually, or as suites of characters, using concentrated change tests. The phylogeny of these moth mimics revealed that individual mimetic characters were conserved, as are the three mimetic wasp forms: yellow Polybia, black Polybia and Parachartergus mimetic types. This finding was consistent with a 'supergene' control of linked loci and the Nicholson two-step model of mimicry evolution. We also used a modified permutation-tail probability approach to examine the rate of mimetic-type evolution. The observed topology, hypothetical Müllerian and Batesian scenarios, and 1000 random trees were compared using Kishino-Hasegawa tests. The observed phylogeny was more consistent with the predicted Müllerian distribution of mimetic traits than with that of a quasi-Batesian scenario. We suggest that the range of discriminatory abilities of the predator community plays a key role in shaping mimicry dynamics.     

Lattice models in ecology and social sciences

Random matching between individuals, or the complete-mixing model, is often assumed in analyzing evolutionary or population dynamics in ecology and game theory or other models in social sciences. Making and analyzing a model is not difficult under this simple assumption. However spatial- or network-structured populations, including the lattice model and the power-law network, are more realistic for many ecological and social phenomena than the complete-mixing model. In this review, I will show first that a lattice model can be useful in investigating the effect of neighborhood interactions on the dynamics, not only of plants and forests, but also of animal and human societies. Second, the lattice model promotes the evolution of spiteful behavior, even though it is well-known that the lattice model promotes the evolution of cooperation. Finally, different social networks result in traits, such as social norms, spreading at different speeds.     

OPTIMAL DEFENSIVE COLORATION STRATEGIES DURING THE GROWTH PERIOD OF PREY

Defensive coloration that reduces the risk of predation is considered to be widespread in animals. Many closely related species adopt differing coloration strategies during the life cycle, including crypsis, conspicuousness, and ontogenic change between the two coloration types. Here, we use a dynamic state-dependent approach to use ecological and intrinsic factors to predict the proportion of the developmental period of immature animals that should be spent as cryptic or conspicuous, and when conspicuous coloration should be reliably associated with investment in defenses. The model predicts that animals should change color more than once during development only in specific circumstances. In contrast, change from crypsis to conspicuous can occur over a range of conditions related to the frequency of detection by predators, but may also depend on the opportunity costs of crypsis and the effect of size on the deterrent effect of conspicuous coloration. We also report the results of a survey of coloration strategies in lepidopteron larvae, and note a qualitative agreement with the predictions of our model in the relationship between body size and coloration strategy. Our results provide explanations for several widespread antipredator coloration phenomena in prey animals, and provide a comprehensive predictive framework for the types of coloration strategies that are employed in nature.     

Ultraviolet and green parts of the colour spectrum affect egg rejection in the song thrush (Turdus philomelos)

Much attention has been devoted to understanding the evolution of egg mimicry in avian brood parasites. The majority of studies have been based on human perception when scoring the mimicry of the parasitic egg. Surprisingly, there has been no detailed study on the recognition and sensitivity towards differently coloured parasitic eggs. We investigated effect of different colours of the experimental eggs measured by ultraviolet (UV)-visible reflectance spectrophotometry on rejection behaviour in the song thrush ( Turdus philomelos ). We carried out a set of experiments with four blue model eggs representing mimetic eggs, whereas six other colours represented nonmimetic eggs. Our results revealed that two colours originally designed as a mimetic were rejected at a high rate, whereas one group of the nonmimetic was accepted. A multiple regression model of absolute differences between song thrush and experimental eggs on rejection rate showed that the level of mimicry in the UV and green parts of the colour spectrum significantly influenced egg rejection in the song thrush. To our knowledge, this is the first detailed study showing that different colour perception by the birds can affect their responses towards the parasitic egg. These findings suggest that the combination of UV and visible ranges of the spectra plays a major role in the evolution of discrimination processes, as well as in the evolution of the mimicry of the parasitic egg.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 269–276.