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Understanding the origin and maintenance of adaptive phenotypic novelty is a central goal of evolutionary biology. However, both hybridization and incomplete lineage sorting can lead to genealogical discordance between the regions of the genome underlying adaptive traits and the remainder of the genome, decoupling inferences about character evolution from population history. Here, to disentangle these effects, we investigated the evolutionary origins and maintenance of Batesian mimicry between North American admiral butterflies (Limenitis arthemis) and their chemically defended model (Battus philenor) using a combination of de novo genome sequencing, whole-genome resequencing, and statistical introgression mapping. Our results suggest that balancing selection, arising from geographic variation in the presence or absence of the unpalatable model, has maintained two deeply divergent color patterning haplotypes that have been repeatedly sieved among distinct mimetic and nonmimetic lineages of Limenitis via introgressive hybridization.

Biological mimicry is regarded by many as a textbook illustration of Darwin’s idea of evolution by random mutation followed by differential selection of reproductively fit specimens, resulting in gradual phenotypic change in a population. In this paper, I argue that some cases of so-called mimicry are probably merely look-a-likes and do not gain an advantage due to their similarity in appearance to something else. In cases where a similar appearance does provide a benefit, I argue that it is possible that these forms of mimicry were created in a single generation. An interpretive response to an appearance as a sign can make a new structure perform drastically differently in an environment. In such cases, Darwin’s natural selection mechanism only helps to explain gradual the spread of these new forms, not the creation of them. I argue that biosemiosis should be regarded as a much more powerful mechanism for affecting evolutionary trajectories than the gradualist view allows. I focus on two cases of butterfly mimicry: the Viceroy (Nymphalidae: Limenitis archippus) and Monarch (Nymphalidae: Danaus plexippus) butterflies, supposed Mullerian mimics, and deadleaf mimic butterflies (Kallima).

The phylogenetic relationships of the nymphalid butterfly tribe Limenitidini are best known for the genera Limenitis and Adelpha, model taxa for evolutionary processes such as Batesian mimicry and rapid adaptive radiations. Whereas these American limenitidines have received the most attention, phylogenetic relationships of their Asian relatives are still controversial and largely unexplored. Even one of the largest genera in Asia, Athyma, is polyphyletic. To clarify the phylogenetic relationships of these Asian Limenitidini, a total of 53 representatives were sampled; 37 have their mitogenomes sequenced for the first time. Our phylogenetic results confirm that mitogenomic data provides well-resolved relationships at most major levels of the phylogeny, even using different partition schemes or different inference methods. Interestingly, our results show that some Athyma taxa are embedded within the genus Limenitis, whereas the genus Tacola, previously considered to be a synonym of Athyma, needs to be recognized as a valid clade. Additionally, the other Limenitidini genera in Asia (namely Tarattia, Litinga, Sumalia, Pandita and Patsuia) are now grouped either within Athyma or Limenitis, so these genera need to be sunk. Importantly, we also show that the mainly Old World Limenitis and entirely New World Adelpha are sister groups, confirming the relevance of Asian lineages to global studies of Limenitis evolution.

Emmanouil Limenitis XV. yüzyılda Rodos adasında yaşamış bir şairdir. Toplamda ona atfedilen üç şiir vardır. Tezimin çeviri ve inceleme konusu olan “Rodos’ta Veba” isimli son eseri, 1500 yılında, Osmanlı fethi öncesi Rodos’ta yazılan en önemli eserlerden biri sayılmaktadır. Eserin adından da anlaşılacağı üzere, yazar, Rodos adasını iki yıl boyunca etkisi altına alan veba salgını hakkında bilgiler sunmaktadır. 1498-1500 yıllarında Rodos’ta yaşanan veba salgını yazarın ailesinden de olmak üzere adada yaşayan pek çok kişinin hayatını kaybetmesine sebep olmuştur. Şüphesiz ki bu trajik olay yazarı bu şiiri yazmaya teşvik etmiştir, ancak eseri bu kadar mühim kılan vebaya ek olarak adanın tarihi ve geleneksel yaşantısı hakkında bilgiler içermesidir. Yunanların, Katolik Latin Saint Jean Şövalyeleri yönetimi altında Rodos adasında hangi şartlar altında yaşadıklarını, yazarın Latinlere ve Türklere bakış açısını, Ortaçağ Hıristiyan dünyasının dini ve sosyal yaşantısını, vebanın Ortaçağ edebiyatı ve sanatına yansımalarını ve yazarın kullandığı dil özellikleri gibi birçok konu çalışmamızda ele alınmıştır. Eser içerdiği bu bilgilerle XV. yüzyılda On İki Adalar kültürüne ışık tutmuştur. Emmanouil Limenitis was a poet who lived in the 15th century on the island of Rhodes. Three poems are attributed to him. The title of my dissertation is the translation and analysis of his last poem titled “The Plague of Rhodes”. The work was written in 1500 and is one of the most important works written on Rhodes before the Ottomans captured the island. As can be seen from the name of the poem, the author gave information about the plague epidemic that struck for two years the island of Rhodes. This epidemic took place in 1498-1500 and caused death to the inhabitants of the island even from the family of the poet. Undoubtedly this tragic event became the reason for writing this poem but the thing that makes the work so remarkable is the information the poet gave us about the history and the traditional life of the island. The work includes topics such as the life of the Greeks under the domination of the Catholic Knights Hospitaller, the poet’s view of the Latins and the Turks, the religious and social life of the Christian inhabitants of the island in the Middle Ages, the influence of plague on literature and the art in the Middle Ages and properties of the medieval Greek language. The information contained in the work illuminates the 15th century Dodecanese culture.

Since the phenomenon of mimicry was first described by Bates in 1862 it has become one of the foundational examples of adaptive evolution. Numerous subcategories of mimicry and dozens of hypotheses pertaining to its evolution and maintenance have been proposed. Many of these hypotheses, however, are difficult to test in experimental settings, and data from natural observations are often inadequate. Here we use data from a long-term survey of butterfly presence and abundance to test several hypotheses pertaining to Batesian and female-limited polymorphic mimicry (FPM; a special case of Batesian mimicry). We found strong evidence that models outnumber mimics in both mimicry systems, but no evidence for an increase in relative abundance of FPM mimics to their Batesian counterparts. Tests of the early-emergence/model first hypothesis showed strong evidence that the Batesian mimic routinely emerges after the model, while emergence timing in the FPM system was site specific, suggesting that other ecological factors are at play. These results demonstrate the importance of long-term field observations for testing evolutionary and ecological hypotheses.

Currently, DNA barcodes are often required to be analyzed using old museum specimens when they are the only available specimens for rare or endangered species, or even type series. In this study, using eight universal primers and newly designed 315 species-specific primers, we tried to recover full-length barcode sequences from 45 dried specimens of 36 butterfly species collected between 1959 and 1980 in Korea. The eight universal primers failed entirely in the PCR amplification and sequencing of all the specimens. On the other hand, 284 primer pairs consisting of the 315 primers, targeting fragments of 71–417 bp, amplified various lengths of barcode sequences from all specimens. The fragments were successfully combined to generate the barcode sequences ranging from 444 bp to 658 bp. Notably, of the 284 primer pairs, 26 primer pairs designed for Limenitis camilla , Argynnis niobe , and Brenthis daphne successfully amplified the barcode sequences of congeneric species, Limenitis doerriesi , Argynnis nerippe , and Brenthis ino , suggesting that the species-specific primers can be available for analyzing barcode sequences of closely related species. Our study reveals that the newly designed species-specific primers will be effective in acquiring COI sequences from old butterfly specimens.

Butterflies and primates are interesting for comparative color vision studies, because both have evolved middle- (M) and long-wavelength- (L) sensitive photopigments with overlapping absorbance spectrum maxima (λ max values). Although positive selection is important for the maintenance of spectral variation within the primate pigments, it remains an open question whether it contributes similarly to the diversification of butterfly pigments. To examine this issue, we performed epimicrospectrophotometry on the eyes of five Limenitis butterfly species and found a 31-nm range of variation in the λ max values of the L-sensitive photopigments (514–545 nm). We cloned partial Limenitis L opsin gene sequences and found a significant excess of replacement substitutions relative to polymorphisms among species. Mapping of these L photopigment λ max values onto a phylogeny revealed two instances within Lepidoptera of convergently evolved L photopigment lineages whose λ max values were blue-shifted. A codon-based maximum-likelihood analysis indicated that, associated with the two blue spectral shifts, four amino acid sites (Ile17Met, Ala64Ser, Asn70Ser, and Ser137Ala) have evolved substitutions in parallel and exhibit significant d N /d S >1. Homology modeling of the full-length Limenitis arthemis astyanax L opsin placed all four substitutions within the chromophore-binding pocket. Strikingly, the Ser137Ala substitution is in the same position as a site that in primates is responsible for a 5- to 7-nm blue spectral shift. Our data show that some of the same amino acid sites are under positive selection in the photopigments of both butterflies and primates, spanning an evolutionary distance >500 million years.

© 2015 The Linnean Society of London. Since the phenomenon of mimicry was first described by Bates in 1862 it has become one of the foundational examples of adaptive evolution. Numerous subcategories of mimicry and dozens of hypotheses pertaining to its evolution and maintenance have been proposed. Many of these hypotheses, however, are difficult to test in experimental settings, and data from natural observations are often inadequate. Here we use data from a long-term survey of butterfly presence and abundance to test several hypotheses pertaining to Batesian and female-limited polymorphic mimicry (FPM; a special case of Batesian mimicry). We found strong evidence that models outnumber mimics in both mimicry systems, but no evidence for an increase in relative abundance of FPM mimics to their Batesian counterparts. Tests of the early-emergence/model first hypothesis showed strong evidence that the Batesian mimic routinely emerges after the model, while emergence timing in the FPM system was site specific, suggesting that other ecological factors are at play. These results demonstrate the importance of long-term field observations for testing evolutionary and ecological hypotheses.

Limenitis camilla (Linnaeus, 1764) Papilio camilla Linnaeus, 1764: 304. FIRST RECORD FOR THE VALDIERI AREA. — Cassulo L. 2001 pers. coll. (Andonno, 900 m). CHOROTYPE. — European; NE China and Amur to Japan (disjunct). ECOLOGY. — Nemoral, lowland-lower montane, mesophilous, mesophilous. EDIT PROJECT SAMPLING. — Not found., Published as part of Bonelli, Simona, Barbero, Francesca, Casacci, Luca Pietro, Cerrato, Cristiana & Balletto, Emilio, 2015, The butterfly fauna of the Italian Maritime Alps: results of the EDIT project, pp. 139-167 in Zoosystema 37 (1) on page 158, DOI: 10.5252/z2015n1a6, http://zenodo.org/record/5154896, {"references":["LINNAEUS C. 1764. - Museum S: ae R: ae M: tis Ludovicae Ulricae Reginar Svecorum. Laur. Salvii: [i-vi], 1 - 713 [+ index]."]}

The evolution of wing pattern diversity in butterflies has emerged as a model system for understanding the origins and maintenance of adaptive phenotypic novelty. Admiral butterflies (genus Limenitis) are an attractive system for studying wing pattern diversity because mimicry is common among the North American species and hybrid zones occur wherever mimetic and non-mimetic wing pattern races meet. However, the utility of this system has been limited because the evolutionary relationships among these butterflies remain unclear. Here I present a robust species-level phylogeny of Limenitis based on 1911 bp of two mitochondrial genes (COI and COII) and 904 bp of EF1-alpha for all five of the Nearctic species/wing pattern races, the majority of the Palearctic species, and three outgroup genera; Athyma, Moduza (Limenitidini), and Neptis (Limenitidinae: Neptini). Maximum-likelihood and Bayesian analyses indicate that the North American species are a well-supported, monophyletic lineage that is most closely related to the widespread, Palearctic, Poplar admiral (L. populi). Within North America, the Viceroy (L. archippus) is the basal lineage while the relationships among the remaining species are not well resolved. A combined maximum-likelihood analysis, however, indicates that the two western North America species (L. lorquini and L. weidemeyerii) are sister taxa and closely related to the wing pattern subspecies of the polytypic Limenitis arthemis species complex. These results are consistent with (1) an ancestral host-shift to Salicaceae by the common ancestor of the Poplar admiral and the Nearctic admiral lineage, (2) a single colonization of the Nearctic, and (3) a subsequent radiation of the North American forms leading to at least three independent origins of mimicry.

Rapid diversification is often associated with morphological or ecological adaptations that allow organisms to radiate into novel niches. Neotropical Adelpha butterflies, which comprise over 200 species and subspecies, are characterized by extraordinary breadth in host plant use and wing colour patterns compared to their closest relatives. To examine the relationship between phenotypic and species diversification, we reconstructed the phylogenetic history of Adelpha and its temperate sister genus Limenitis using genomewide restriction-site-associated DNA (RAD) sequencing. Despite a declining fraction of shared markers with increasing evolutionary distance, the RAD-Seq data consistently generated well-supported trees using a variety of phylogenetic methods. These well-resolved phylogenies allow the identification of an ecologically important relationship with a toxic host plant family, as well as the confirmation of widespread, convergent wing pattern mimicry throughout the genus. Taken together, our results support the hypothesis that evolutionary innovations in both larvae and adults have permitted the colonization of novel host plants and fuelled adaptive diversification within this large butterfly radiation.

Life history characteristics of the Western Viceroy (Limenitis archippus obsoleta), an obligate riparian nymphalid butterfly in the desert southwestern United States, are described and related to Colorado River riparian restoration efforts. Riverine disturbance regimes and associated fluvial and hydrological dynamics may provide resources critical to this butterfly. Puddling by adult butterflies may require flood-cleared surfaces and an obligate riparian plant, Gooddings willow, was a larval host plant. This butterfly needs a variety of resources that are only found in close proximity in naturally functioning riparian ecosystems. Habitat heterogeneity required for colony persistence depends largely upon the natural dynamic character of flowing water systems. Because of the links between this butterfly and riparian structure and function it may be a useful indicator for monitoring riparian ecosystem restoration in the area.

Limenitis rileyi is recorded for the first time in India from Upper Dibang Valley District, Arunachal Pradesh.

© The Author 2014. Published by Oxford University Press on behalf of the society for Molecular Biology and Evolution.. Opsins are ancient molecules that enable animal vision by coupling to a vitamin-derived chromophore to form light-sensitive photopigments. The primary drivers of evolutionary diversification in opsins are thought to be visual tasks related to spectral sensitivity and color vision. Typically, only a few opsin amino acid sites affect photopigment spectral sensitivity. We show that opsin genes of the North American butterfly Limenitis arthemis have diversified along a latitudinal cline, consistent with natural selection due to environmental factors. We sequenced single nucleotide (SNP) polymorphisms in the coding regions of the ultraviolet (UVRh), blue (BRh), and long-wavelength (LWRh) opsin genes from ten butterfly populations along the eastern United States and found that a majority of opsin SNPs showed significant clinal variation. Outlier detection and analysis of molecular variance indicated that many SNPs are under balancing selection and show significant population structure. This contrasts with what we found by analysing SNPs in the wingless and EF-1 alpha loci, and from neutral amplified fragment length polymorphisms, which show no evidence of significant locus-specific or genome-wide structure among populations. Using a combination of functional genetic and physiological approaches, including expression in cell culture, transgenic Drosophila, UV-visible spectroscopy, and optophysiology, we show that key BRh opsin SNPs that vary clinally have almost no effect on spectral sensitivity. Our results suggest that opsin diversification in this butterfly is more consistent with natural selection unrelated to spectral tuning. Some of the clinally varying SNPs may instead play a role in regulating opsin gene expression levels or the thermostability of the opsin protein. Lastly, we discuss the possibility that insect opsins might have important, yet-to-be elucidated, adaptive functions in mediating animal responses to abiotic factors, such as temperature or photoperiod.

Reciprocal double mating experiments were done between the northern banded (disruptively colored) form [Limenitis arthemis arthemis (Drury)] and the southern unbanded mimetic form [L. arthemis astyanax (F.)] of the banded purple butterfly. The results showed that following a second mating (and the successful transfer of a second spermatophore into the bursa copulatrix of each female), the first male continued to sire the subsequent progeny in five of 17 (29%) cases. The sequential double matings of these 17 different females with 34 different males resulted in full, partial, and unbanded wing phenotypes which were used as paternal genetic markers in this intergrading complex of butterflies. The containment of the spermatozoa within separate spermatophores at the time of mating normally resulted in only one effective sire. Among three broods (all one of four cross types) some sperm mixing may have occurred following the second matings, as evidenced by the occurrence of a few unbanded insects among...

In the eastern United States, the mimetic viceroy butterfly (Limenitis archippus) exhibits clinal variation in wing color, ranging from a tawny orange phenotype (L. a. archippus) in the N to a dark mahogany phenotype (L. a. floridensis) in Florida. Geographic distributions of these two subspecies are roughly coincident with the ranges of the viceroy's two eastern mimicry models: the monarch (Danaus plexippus) in the N, and the queen (D. gilippus) in the S. This coincidence has historically been attributed to “model-switching”: presumably, southern viceroys have switched from mimicking the monarch to mimicking the locally predominant queen, due to selective pressure exerted by visually foraging predators. As an initial test of this hypothesis, I sought evidence of selective predation on light and dark viceroys by captive red-winged blackbirds previously exposed to either monarchs or queens. Results were consistent with the model-switching hypothesis: queen-conditioned birds preferentially avoided ...

Viceroy butterflies (Limenitis archippus), long considered palatable mimics of distasteful danaine butterflies, have been shown in studies involving laboratoryreared specimens to be moderately unpalatable to avian predators. This implies that some viceroys are Müllerian co-mimics, rather than defenseless Batesian mimics, of danaines. Here, I further test this hypothesis by assessing the palatability of wild-caught viceroys from four genetically and ecologically diverse populations in the southeastern United States. Bioassays revealed that viceroys sampled from three sites in Florida and one in South Carolina were all moderately unpalatable to captive redwinged blackbird predators, which ate fewer than half of the viceroy abdomens presented. Red-wings commonly exhibited long manipulation times and considerable distress behavior when attempting to eat a viceroy abdomen, and they taste-rejected over one-third of viceroys after a single peck. These findings, the first based on wild-caught butterflies, support the hypothesis that the viceroy-danaine relationship in some areas represents Müllerian mimicry, prompting a reassessment of selective forces shaping the interaction. Moreover, considerable variation in palatability of individual viceroys, and in behavior of individual birds, contributes to the complexity of chemical defense and mimicry in this system.

This study documents the existence of a food plant related palatability spectrum in Florida queen butterflies (Danaus gilippus). Abdomens of butterflies reared on four asclepiad food plants differed significantly in cardenolide content and palatability to a generalist avian predator (Red—winged Blackbird). However, unpalatability did not directly parallel food plant or butterfly cardenolide content. Queens reared on Sarcostemma clausum were cardenolide free and essentially palatable (85% eaten). Butterflies reared on Asclepias tuberosa and A. incarnata were also virtually cardenolide free, but were weakly and moderately unpalatable, respectively (62 and 46% eaten). Queens reared on the high cardenolide Asclepias curassavica contained 101—753 mg cardenolide and were very unpalatable (8% eaten). This results suggest that in addition to cardenolides, other asclepiad phytochemicals are directly or indirectly utilized by queens as components of their systemic chemical defense. The demonstrated variation in queen palatability is predicted to affect the aversive conditioning of predators and thus the outcome of queen—predator encounters. Queen populations feeding on more than one asclepiad species must exhibit automimicry, with attack rates on individual butterflies dependent upon both the proportion of palatable automimics and the noxiousness of unpalatable automodels in the population. The verification of a queen palatability spectrum also contributes to understanding the dynamics of mimicry between queens and viceroy butterflies (Limenitis archippus). This study predicts that, as a consequence of regional differences in asclepiad floras, the viceroy queen mimicry relationship in Florida switches between the Batesian and Mullerian mimicry modes. In the extreme, queens feeding on certain food plants may be Batesian mimics of sympatric, moderately unpalatable viceroys, reversing the traditionally cited mimicry roles of these butterflies.

As butterflies, monarchs (Danaus plexippus) (Fig. 7.1) do not fight predators or act passively if an attack by a predator is imminent; instead, the monarch uses warning coloration and chemical defenses against predation. As adults, monarchs are brightly colored in orange, black, and white; they, therefore, stand out in the environment to a predator, much like the pattern of a poisonous coral snake (Micrurus fulvius) (Zug et al. 2001). What does the coloration of a monarch signal to a potential predator? That is unpalatable, so stay away. Monarch becomes unpalatable because of toxins that larvae feed upon in milkweed (Asclepias syriaca). These toxins are cardio glycosides, which elicits vomiting in birds that may feed upon them. Only two bird species, the black-headed oriole (Icterus abeillei) and the black-headed grosbeak (Pheucticus melanocephalus) are known to feed on monarch with no ill effect. In fact, the viceroy butterfly (Limenitis archippus) mimics the coloration of the monarch to offset predation on it, which is a form of mimicry known as Batesian.

Butterflies have long inspired evolutionary biologists with the striking diversity of their wing colors. A more recent surprising finding has been the remarkable diversity of their color vision systems. Butterfly photoreceptors, unlike those of bees and moths, have diversified extensively through processes such as opsin gene duplications, positive selection on single opsin loci, and heterogeneous expression of filtering pigments. Further, lineage-specific opsin duplications and expression of filtering pigments have facilitated the evolution of sexually dimorphic butterfly eyes. Although the molecular basis of visual system diversification is relatively well understood, its ecological and evolutionary significance remains to be elucidated.

The Na+, K+-ATPase of the Monarch butterfly (Danaus plexippus) is insensitive to the inhibition by cardiac glycosides due to an amino acid replacement: histidine instead of asparagine at position 122 of the α-subunit representing the ouabain binding site. By PCR amplification of the DNA sequence of this site, a PCR product of 270 bp was obtained from DNA extracted from Danainae species (Danaus plexippus, D. chrysippus, D. gillipus, D. philene, D. genutia, Tirumala hamata, Euploea spp., Parantica weiskei, P. melusine), Sphingidae (Daphnis nerii) and mimics of milkweed butterflies (Hypolimnas missipus, Limenitis archippus and L. arthemis, Nymphalidae). Analysis of the nucleotide sequences revealed that the single point mutation in the ouabain binding domain (AAC-Asn for CAC-His) was present only in Danaus plexippus, but not in the other species investigated. Since these milkweed butterflies also store cardenolides, other structural modifications of the Na+, K+-ATPase may have occurred or other strategies of cardenolide tolerance have been developed.

Hybrid zones present opportunities to study the effects of gene flow, selection, and recombination in natural populations and, thus, provide insights into the genetic and phenotypic changes that occur early in speciation. Here we investigate a hybrid zone between mimetic (Limenitis arthemis astyanax) and nonmimetic (Limenitis arthemis arthemis) populations of admiral butterflies using DNA sequence variation from mtDNA and seven nuclear gene loci. We find three distinct mitochondrial clades within this complex, and observe a strong overall concordance between wing-pattern phenotypes and mitochondrial variation. Nuclear gene genealogies, in contrast, revealed no evidence of exclusivity for either wing-pattern phenotype, suggesting incomplete barriers to gene exchange and/or insufficient time for lineage sorting. Coalescent simulations indicate that gene flow between these two subspecies is highly asymmetric, with the majority of migration occurring from mimetic into nonmimetic populations. Selective sweeps of alleles responsible for mimetic phenotypes may have occurred more than once when mimetic and nonmimetic Limenitis occurred together in the presence of the model (Battus philenor).

Batesian mimics gain protection from predation through the evolution of physical similarities to a model species that possesses anti-predator defences. This protection should not be effective in the absence of the model since the predator does not identify the mimic as potentially dangerous and both the model and the mimic are highly conspicuous. Thus, Batesian mimics should probably encounter strong predation pressure outside the geographical range of the model species. There are several documented examples of Batesian mimics occurring in locations without their models, but the evolutionary responses remain largely unidentified. A mimetic species has four alternative evolutionary responses to the loss of model presence. If predation is weak, it could maintain its mimetic signal. If predation is intense, it is widely presumed the mimic will go extinct. However, the mimic could also evolve a new colour pattern to mimic another model species or it could revert back to its ancestral, less conspicuous phenotype. We used molecular phylogenetic approaches to reconstruct and test the evolution of mimicry in the North American admiral butterflies (Limenitis: Nymphalidae). We confirmed that the more cryptic white-banded form is the ancestral phenotype of North American admiral butterflies. However, one species,Limenitis arthemis, evolved the black pipevine swallowtail mimetic form but later reverted to the white-banded more cryptic ancestral form. This character reversion is strongly correlated with the geographical absence of the model species and its host plant, but not the host plant distribution ofL. arthemis. Our results support the prediction that a Batesian mimic does not persist in locations without its model, but it does not go extinct either. The mimic can revert back to its ancestral, less conspicuous form and persist.

Understanding the dynamics of defensive mimicry requires accurately characterizing the comparative palatability of putative models and mimics. The Florida viceroy butterfly (Limenitis archippus floridensis) is traditionally considered a palatable Batesian mimic of the purportedly distasteful Florida queen (Danaus gilippus berenice). I re-evaluated this established hypothesis by directly assessing palatability of viceroys and queens to red-winged blackbirds in a laboratory experiment. Representative Florida viceroys were surprisingly unpalatable to red-wings; only 40% of viceroy abdomens were entirely eaten (compared to 98% of control butterfly abdomens), and nearly one-third were immediately tasterejected after a single peck. In fact, the viceroys were significantly more unpalatable than representative Florida queens, of which 65% were eaten and 14% taste-rejected. Thus, viceroys and queens from the sampled populations exemplify Mullerian rather than Batesian mimicry, and the viceroy appears to be the stronger model. These findings prompt a reassessment of the ecological and evolutionary dynamics of this classic mimicry relationship.