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There is noticeable variation in male mate-seeking behavior among species of Brassolini butterflies. Males of Opsiphanes Doubleday and Caligo Hübner (Lepidoptera, Nymphalidae) species perform crepuscular displays along forest edges and in light gaps. While male Opsiphanes perform aerial displays, Caligo males perch and wait for receptive females. A comparison of five species of each genus suggests that male display behavior is associated with, and has likely influenced the evolution of their wing attributes and body design. Opsiphanes males have higher wing aspect ratio and more distal centroid position than congeneric females, suggesting that the energetic demands of aerial displays led to sexual dimorphism in wing morphology. In contrast, male and female Caligo generally showed similar wing morphology, which possibly results from the lower energy expenditure of perching behavior when compared with active flight. Likely due to a genetic correlation between sexes, female Opsiphanes and Caligo are more similar in wing morphology and body design to their congeneric males than to each other. Based on our analyses, we make predictions about reproductive behavior for four species in which male mating displays are unknown.

Based on comparative morphology of adults, a phylogeny is proposed for the butterfly tribe Amathusiini (Nymphalidae, Satyrinae). The dataset includes 92 characters scored for 45 species in 12 genera, representing the most comprehensive phylogenetic analysis for this group. Parsimony analyses produced a well-resolved strict consensus tree where genera were divided in three main groups: (clade 1) Stichophthalma; (clade 2) Aemona, Faunis, Melanocyma and Taenaris; (clade 3) Enispe, Discophora, Thaumantis, Thauria, Amathusia, Amathuxidia, and Zeuxidia. While genera in clades 1 and 2 were found to be morphologically homogeneous, clade 3 showed remarkable morphological divergence between and within genera. The monophyly of most genera was recovered with variable levels of support, but Melanocyma and Taenaris nested within Faunis. Therefore, here Melanocyma NEW SYN. is subsumed within Faunis, and Taenaris STAT. REV. is regarded as a subgenus of Faunis. Mimicry likely evolved a single time within the Faunis-Taenaris assemblage, as species of Taenaris formed a monophyletic group. Results are compared to early classifications and recent DNA-based analyses, and points of agreement and conflicts are discussed.

The global increase in species richness toward the tropics across continents and taxonomic groups, referred to as the latitudinal diversity gradient, stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluate several of these hypotheses to explain spatial diversity patterns in a butterfly family, the Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal, and also the extent to which these processes differ among regions at the same latitude. We generate a time-calibrated phylogeny containing 2,866 nymphalid species (~45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the latitudinal diversity gradient among regions because temporal diversification dynamics differ greatly across longitude. The Neotropical diversity results from low extinction rates, not high speciation rates, and biotic interchanges with other regions are rare. Southeast Asia is also characterized by a low speciation rate but, unlike the Neotropics, is the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, combined with tropical niche conservatism, played a major role in generating the modern latitudinal diversity gradient of nymphalid butterflies., F.L.C is supported by an “Investissements d’Avenir” grant managed by the Agence Nationale de la Recherche (CEBA, ref. ANR-10-LABX-25-01). D.J.L. was supported by grants DEB-1541557 from NSF and WW-227R-17 from the National Geographic Society. R.I.V.W. was supported by Leverhulme Trust emeritus programme. M.E. was supported by an ATIP grant, a grant from the Human Frontier Science Program (RGP0014/2016) and a grant from the French National Research Agency (ANR CLEARWING ANR-16-CE02-0012). E.O.A. was supported by Sigma-Xi (G20100315153261), Center for Systematic Entomology and the Council of the Linnean Society and the Systematics Association for the Systematics Research Fund. S.N. was supported by the Swedish Research Council (2015-04218 and 2019-03441). P.M.M. was supported by a Czech Science Foundation grant (Junior GAČR, 20-18566Y). R.V. was supported by projects PID2019-107078GB-I00 / AEI / 10.13039/501100011033 (Agencia Estatal de Investigación) and 2017-SGR-991 (Generalitat de Catalunya). N.W. acknowledges support by the Swedish Research Council (2015-04441) and a start-up grant from the Department of Biology, Lund University.

This study reassesses the taxonomic status of Neotropical blue and purple/rose-colored Cithaerias butterflies, thus complementing a previous study of the rose-colored species. Based on comparative study of wing coloration and genitalia morphology, I revise species definitions and the taxonomic status of: Cithaerias andromeda, C. azurina STAT. REV., C. esmeralda STAT. REV., C. bandusia STAT. REV., C. pyropina, and C. songoana STAT. REV. Photographs of adults and illustrations of male and female genitalia are provided for all species. Of particular importance are the genitalia illustrations of male and female C. azurina, presented here for the first time, as well as finding a putative first female of Ecuadorean subspecies C. pyropina julia.

Neotropical butterflies in the nymphalid genera Pierella Herrich-Schäffer, 1865 (Lepidoptera, Nymphalidae) and Cithaerias Hübner, 1819 (Lepidoptera, Nymphalidae) are known to glide near ground level, and have a high forewing aspect ratio. The hind wings of Pierella are proportionately larger than those of Cithaerias, suggesting a potential role in gliding flight. We show that Pierella helvina (Hewitson, 1860) (Lepidoptera, Nymphalidae) is capable of gliding for longer periods of time than Cithaerias pireta (Stoll, 1780) (Lepidoptera, Nymphalidae) both in the field and a laboratory setting. Experimental reduction of hind wing area in P. helvina led to a decrease in male gliding performance, and an increase in female wing-beat frequency. We also found that escape velocity decreased in both sexes after hind wing area reduction, but the effect was strongest in females. Although the increased drag associated with large hind wings could be detrimental to gliding flight, the large hind wing area of P. helvina effectively enhances gliding performance in ground effect. This study is the first to investigate the role of hind wings on butterfly gliding performance.

Species delimitation is at the core of biological sciences. During the last decade, molecular-based approaches have advanced the field by providing additional sources of evidence to classical, morphology-based taxonomy. However, taxonomy has not yet fully embraced molecular species delimitation beyond threshold-based, single-gene approaches, and taxonomic knowledge is not commonly integrated to multi-locus species delimitation models. Here we aim to bridge empirical data (taxonomic and genetic) with recently developed coalescent-based species delimitation approaches. We use the multispecies coalescent model as implemented in two Bayesian methods (DISSECT/STACEY and BP&P) to infer species hypotheses. In both cases, we account for phylogenetic uncertainty (by not using any guide tree) and taxonomic uncertainty (by measuring the impact of using or not a priori taxonomic assignment to specimens). We focus on an entire Neotropical tribe of butterflies, the Haeterini (Nymphalidae: Satyrinae). We contrast divergent taxonomic opinion—splitting, lumping and misclassifying species—in the light of different phenotypic classifications proposed to date. Our results provide a solid background for the recognition of 22 species. The synergistic approach presented here overcomes limitations in both traditional taxonomy (e.g. by recognizing cryptic species) and molecular-based methods (e.g. by recognizing structured populations, and not raise them to species). Our framework provides a step forward towards standardization and increasing reproducibility of species delimitations.

Throughout its range in Africa, Papilio dardanus Brown, 1776 (Lepidoptera: Papilionidae) displays femalelimited mimicry of multiple model species, and the absence of hind wing tails is an important component of their mimetic convergence. Nonmimetic P. dardanus females have a narrow, disjunct distribution (Ethiopia, Madagascar, Comoros), and resemble males in color and by possessing hind wing tails. We used elliptical Fourier analysis to investigate whether mimetic P. dardanus female forms converged on the wing shape of their unpalatable models in the tribes Acraeini and Danaini (Lepidoptera: Nymphalidae). Models varied in forewing and hind wing shapes, and separated in shape space according to phylogenetic affinities. Forewing and hind wing shapes of mimics did not closely match those of models. Nonetheless, we found that mimetic P. dardanus female hind wings differed from conspecific males that had their tails photographically removed to allow standardized comparisons. Four nonmimetic Papilio Linnaeus, 1758 (Lepidoptera: Papilionidae) species did not show significant wing shape dimorphism between sexes, supporting the idea that in P. dardanus females, the evolution of mimicry led to changes in hind wing shape beyond the loss of tails.

The latitudinal diversity gradient (LDG) is arguably one of the most striking patterns in nature. The global increase in species richness toward the tropics across continents and taxonomic groups stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluated several of these hypotheses to explain spatial diversity patterns in the butterfly family, Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal to the LDG, and also the extent to which these processes differ among regions at the same latitude. We generated a new, time-calibrated phylogeny of Nymphalidae based on 10 gene fragments and containing ca. 2,800 species (∼45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the LDG among regions because temporal diversification dynamics differ greatly across longitude. For example, we found that Neotropical nymphalid diversity results from low extinction rates, not high speciation rates, and that biotic interchanges with other regions were rare. Southeast Asia was also characterized by a low speciation rate but, unlike the Neotropics, was the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, particularly during the Eocene-Oligocene transition, combined with the conserved ancestral tropical niches, played a major role in generating the modern LDG of butterflies.

Papilio hectorides larvae reared on Piper xylosteoides mostly followed a usual five-instar larval stage, but one third of the individuals underwent a supernumerary molt — the majority of which were male. The smaller size of the second instar indicated that a larva would experience a supernumerary molt. Along their development, larvae that underwent six instars showed smaller increments in head capsule size and body mass at each molt than those that experienced five instars. Nonetheless, six-instar larvae were larger at the onset of the penultimate instar, and also produced larger pupae than five-instar larvae. Adding a sixth instar predictably increased the time spent in the larval stage. A previous study showed that P. xylosteoides is a low quality larval host for P. hectorides compared to three other native plants. As larvae that underwent supernumerary molts produced large pupae, plasticity in instar number seems to allow them to compensate for lower quality food.

Cunizza hirlanda tirimbina Chacon, DeVries & Penz, new subspecies is described from Costa Rica. The male and female genitalia of this species are illustrated here for the first time. The DNA barcode sequence is provided for one of the paratypes, and it differs by 2.58% from C. hirlanda niguida from Peru.

Tropical dry forests are among the most diverse and threatened habitats in the world, yet they are rarely protected and remain poorly studied. In Brazil, dry forests are naturally fragmented and embedded within various biomes, thus making it important to assess biotic homogeneity among geographically separated forest fragments. We sampled 7732 individuals belonging to 48 species to quantify the diversity of fruit-feeding butterfly communities at four Brazilian dry forest sites, and found differences in community structure between northern and central sites. Species richness per plot was the same in both areas, but abundance per plot was higher in northern sites. Species composition differed between sites mostly due to species of Satyrinae. Additive partitioning showed that beta diversity corresponded to 70.1% of all diversity. Rather than species loss, beta diversity primarily represented species turnover that was potentially driven by differences in the surrounding habitats. Butterfly community composition and abundance were influenced by vegetation where abundance increased with tree density and basal area, and decreased with the average tree height. Butterfly species richness and abundance were higher in the wet season than in the dry season, and all species sampled in the dry season were present in the wet season. Differences in community structure across relatively short geographic distances in the same type of habitat highlight the importance of conserving tropical dry forest fragments to ensure the maintenance of butterfly diversity and, presumably, other insects.

We assessed responses in grooming behavior to ectoparasite densities in naturally occurring white-tailed deer (Odocoileus virginianus) in southwest Alabama, and tested predictions of the stimulus-driven and programmed grooming models with respect to intraspecific body size and vigilance. Stimulus-driven grooming predicts greater tick densities would lead to an increase in grooming, whereas the programmed model predicts a higher rate of grooming would decrease tick densities. Within the programmed model, smaller individuals are predicted to groom more and host fewer ticks, and, due to increased vigilance, breeding males will groom less than females and bachelor males, and thus host more ticks during the rut. We used generalized linear models to determine males had a higher average tick density than females and exhibited complete separation of tick parasitism between non-rutting and rutting periods. Our results support the stimulus-driven grooming model as both fawns and yearlings had significantly higher d...

When considering Neotropical Lepidoptera for conservation purposes moths are usually neglected because little is known about their biology, ecology and taxonomy. Using light-traps, we sampled moths in the subfamily Arctiinae (Noctuoidea: Erebidae) along an elevational transect (7–927 m asl) on a steep slope of Atlantic Forest in the southern region of Serra do Mar, Brazil. Serra do Mar assemblages clustered according to three elevational zones. We recorded 294 species of Arctiinae in Serra do Mar, and an approach using rarefaction and extrapolation indicated that species richness decreases with increasing elevation, similarly to patterns found in southern Ecuador and Costa Rica, although diversity in Serra do Mar is lower than in southern Ecuador. The elevational transect samples from Serra do Mar were compared with available data for two other biomes. For such comparative analyses, we focused on the tribe Arctiini, for which assemblages were partitioned into Serra do Mar, Interior Atlantic forest and Pampa. The Brazilian Pampa was expected to be less diverse due to low habitat complexity, but it was not possible to explain the higher diversity of Arctiines in Serra do Mar compared to Interior Atlantic Forest. This discrepancy and the probable reasons behind it are discussed, and directions for future research are proposed. Undoubtedly, there is a great need of extensive taxonomical revisions and basic biology research on Arctiines in the Atlantic Forest, Pampa and probably in other tropical and subtropical habitats. These data should be used to improve future biodiversity research and to produce high quality information as a foundation for effective conservation measures.

Flight is a key innovation in the evolutionary success of insects and essential to dispersal, territoriality, courtship and oviposition. Wing shape influences flight performance and selection likely acts to maximize performance for conducting essential behaviours that in turn results in the evolution of wing shape. As wing shape also contributes to fitness, optimal shapes for particular flight behaviours can be assessed with aerodynamic predictions and placed in an ecomorphological context. Butterflies in the tribe Haeterini (Nymphalidae) are conspicuous members of understorey faunas in lowland Neotropical forests. Field observations indicate that the five genera in this clade differ in flight height and behaviour: four use gliding flight at the forest floor level, and one utilizes flapping flight above the forest floor. Nonetheless, the association of ground level gliding flight behaviour and wing shape has never been investigated in this or any other butterfly group. We used landmark-based geometric morphometrics to test whether wing shapes in Haeterini and their close relatives reflected observed flight behaviours. Four genera of Haeterini and some distantly related Satyrinae showed significant correspondence between wing shape and theoretical expectations in performance trade-offs that we attribute to selection for gliding in ground effect. Forewing shape differed between sexes for all taxa, and male wing shapes were aerodynamically more efficient for gliding flight than corresponding females. This suggests selection acts differentially on male and female wing shapes, reinforcing the idea that sex-specific flight behaviours contribute to the evolution of sexual dimorphism. Our study indicates that wing shapes in Haeterini butterflies evolved in response to habitat-specific flight behaviours, namely gliding in ground effect along the forest floor, resulting in ecomorphological partitions of taxa in morphospace. The convergent flight behaviour and wing morphology between tribes of Satyrinae suggest that the flight environment may offset phylogenetic constraints. Overall, this study provides a basis for exploring similar patterns of wing shape evolution in other taxa that glide in ground effect.

Spatial movement is fundamental to ecological and evolutionary processes because of its role in habitat colonization and diversification. This study examines the dispersal potential of two Neotropical butterflies, Morpho achilles and Bia actorion, to estimate the time necessary to expand into their current geographical distribution in the Amazon Forest. To do so, we use life history data, dispersal rates estimated from mark-release-recapture experiments, and reaction-diffusion or integro- difference equations to calculate an annual speed for the wave of dispersal across suitable unoccupied habitat. We show that times required for these butterflies to disperse across Amazonia are orders of magnitude smaller than phylogenetic estimates of the species ages. We compare dispersal rates of Morpho and Bia to other Neotropical butterflies, including taxa that could have moved relatively rapidly through Central America and Mexico. Our findings demonstrate that butterflies (and possibly many other organisms) have tremendous potential for rapid colonization of suitable habitats. Our results also suggest that dispersal can easily increase local diversity in areas where speciation has been attributed mostly to vicariance (such as the eastern slope of the Andes).

Based on the most recent nymphalid phylogeny, the Satyrinae can be tentatively organized into the species-rich tribe Satyrini plus a clade that includes the Morphini, Brassolini, Haeterini, Elymniini, Melanitini, Dirini, Zetherini, and Amathusiini. Members of the latter eight tribes have the largest body sizes within Satyrinae and also show extraordinary wing pattern variation. Representatives of these tribes are illustrated herein, and pattern elements of the nymphalid ground plan are identified. Five themes are briefly discussed in light of their pattern diversification: (1) central symmetry system dislocations, (2) variation in ventral hind wing ocelli, (3) the color band between elements f and g, (4) sexual dimorphism and mimicry, and (5) transparency. Within an ecological and evolutionary standpoint, selected genera are provided as examples to explore wing patterns involved in male mating displays, camouflage, and mimicry.

This study describes and compares the diverse dorsal and ventral wing color patterns of Brassolini butterflies. Thirty-three species are illustrated, where pattern elements of the nymphalid ground plan are labeled in color. In general, a larger number of pattern elements can be identified on the ventral than on the dorsal surface of both wings, and the forewing has a larger number of discernible pattern elements than the hind wing. The dorsal elements are broad, diffuse, and more difficult to identify against the typically brown brassoline wing background color. Species with a light colored dorsal background served as a guide for our proposal that fewer pattern elements are present dorsally, particularly on the hind wing. Colorful bands or markings generally present on the dorsal surface seem to be associated with specific pattern elements and have correspondence to the ventral pattern. We refer to these as trailing bands, and they constitute a predominant feature of the brassoline dorsal coloration. We propose a subordinate groundplan for brassolines and interpret some of the ventral pattern variation in light of their phylogeny. Dorsal color pattern variation that leads to sexual dimorphism and mimetic resemblance are also discussed.

A new phylogenetic hypothesis for the Neotropical butterfly genus Hamadryas, based on the combination of a morphological matrix, one mitochondrial (COI) and four nuclear markers (CAD, RpS5, EF1a, and Wingless), is presented. Results from analyses of the molecular evidence are compared with a previously published morphological phylogeny. Molecular data and the analysis of the complete dataset support the monophyly of Hamadryas and most sister groups suggested by morphological data alone. The addition of DNA sequences to the morphological matrix helped define species groups for which no morphological synapomorphies were found. Partitioned Bremer support indicates that COI, CAD, and morphology were consistently in agreement with the combined evidence tree. In contrast, signal from the nuclear markers Rps5, EF1a, and Wingless showed indifference at most levels of the tree, and minor conflict at nodes solving the relationships between species groups. Though resolved, the combined evidence tree shows low resample values, particularly among species groups whose relationships were characterized by short internodes. A reassessment about the pattern of character change for sound production is presented and discussed.

This study examines the contribution of early-stages and adult characters to the reconstruction of the phylogeny of Brassolini butterflies. Parsimony analyses used both equal weights and implied weights, and a series of analyses were performed. First, we analysed adult and early-stages partitions independently and in combination for a subset of 27 species; in these cases the matrices were mostly complete. Whereas the adult partition alone produced a topology that was well resolved and congruent with previous studies, the early-stages partition produced a poorly resolved tree under equal weights. Furthermore, implied weights produced a well-resolved early-stages topology that differed significantly from the adult topology. When both partitions were combined for 27 species, implied weights yielded a topology that resembled the adult tree except for the positions of Bia and Penetes, but statistical node support was generally lower. This suggests that stochastic noise increased when early-stage characters were added to the adult partition, but the combined partitions topology was not statistically different from that based on adult characters alone. Second, given that preserved early stages are not as readily available as adults, we analysed a matrix including 45 species in which early-stage data were missing for 18 species, and compared the topology to that produced by the adult partition alone. Results were similar to the analyses including fewer species; the combined partitions tree was similar to that from the adult partition except for the position of Bia and Penetes. We compare our findings to other genus-level phylogenetic studies within Lepidoptera that have also used early-stages and adult characters.

This study compiles previously published morphological, colour and behavioural characters and includes new DNA sequence data for eight markers (one mitochondrial and seven nuclear) to re-evaluate phylogenetic relationships and estimate times of divergence for Morpho butterflies using parsimony and Bayesian methods. We note an effect of missing data on phylogenetic inference and calculations of Partitioned Bremer Support. Morphology and DNA trees were moderately congruent, and the combined analyses of all data included elements of both sources. Both morphology and DNA support the monophyly of Morpho and the early separation of the sister pair M. marcus plus M. eugenia, but trees from different data sources are congruent mostly at derived nodes, and differ at several internal nodes. The analyses of combined data indicate that Morpho is composed of four clades each of which include one or more previously proposed subgenera. The subgenera Pessonia and Morpho were not monophyletic, and to address this issue we propose that Pessonia, syn.nov. be subsumed within Morpho. The ancestor of Morpho probably arose during the Oligocene, and most diversification seems to have occurred during the late Miocene. S-DIVA analysis suggests eastern Andean region as the ancestral area for Morpho, and that the South American Atlantic Forest was colonized multiple times.

This study encompasses three trophic levels, including description and observations of a butterfly caterpillar, its host plant, and two parasitoids, one of which is described here as a new species. We describe the immature stages and larval and adult behaviors of the butterfly Eryphanis greeneyi Penz & DeVries, 2008 (Lepidoptera: Nymphalidae) from the east Andean slopes of Ecuador. We identify Chusquea cf. scandens (Poaceae: Bambusoideae) as the larval food plant and review host associations in the genus Eryphanis and related taxa. We compare the morphology of E. greeneyi caterpillars to related taxa feeding on Chusquea and discuss the function of anal combs and associated frass throwing behavior. We describe a new species of braconid wasp Protapanteles eryphanidis sp. n. Whitfield and a tachinid fly, Winthemia sp. nr. analis, reared from field-collected larvae of E. greeneyi and evaluate their taxonomic position and association with E. greeneyi.

Here we describe the immature stages and ecological associations of Antirrhea adoptiva porphyrosticta Watkins, 1928 (Lepidoptera:Nymphalidae:Morphinae). The cloud forest bamboo, Chusquea scandens Kunth (Bambusoidea: Poaceae), serves as the larval food plant for this butterfly in eastern Ecuador, the first hostplant record for Antirrhea outside the family Arecaceae. The larvae of A. adoptiva porphyrosticta are superficially similar to those of other Antirrhea species. We also provide observations on adult and larval behavior. Caterpillars of this butterfly species are parasitized by tachinid flies, as well as by Ichneumonidae and a newly described braconid wasp.

This study provides a species-level phylogeny for the Neotropical brassoline genus Opoptera Aurivillius based on 37 morphological characters. A revised generic definition is given, and two species groups are supported. The syme-group includes three species from the Brazilian Atlantic forest. The aorsa-group includes O. staudingeri (Godman & Salvin) from Central America, three species from western and northern South America, and one species from the Atlantic forest. Two subspecies are elevated to species status: O. hilaris Stichel, NEW STATUS and O. bracteolata Stichel, NEW STA- TUS. Two new combinations are proposed: O. hilaris fuscata Stichel, NEW COMBINATION and O. hilaris colombiana Rothshild, NEW COMBINATION. Diagnoses and illustrations of habitus and genitalia are provided for the eight recognized Opoptera species. Wing color, male scent organs, and male and female genitalic morphology are described and discussed.

This study uses 80 morphological characters and cladistic analysis to evaluate the monophyly and phylogenetic relationships of 18 genera that constitute the butterfly tribe Brassolini. Most characters derive from genitalia, confirming previous generic definitions based mainly on wing characters, and showing that 16 of 18 genera are monophyletic. Mimoblepia Casagrande, syn.n. was subsumed within Opoptera Aurivillius to address the paraphyly of the latter, but resolution of the status of Aponarope Casagrande requires further study. The results suggest that the Brassolini includes six suprageneric groups/clades. Although this study verifies some genus-level relationships put forward over 100 years ago, some new hypotheses of relationships are proposed. Tracing larval host plant use onto the Brassolini phylogeny indicates that species in this tribe retain the use of Arecaceae and Poaceae from their ‘satyroid’ ancestors.

Birds are able to recognize and learn to avoid attacking unpalatable, chemically defended butterflies after unpleasant experiences with them. It has also been suggested that birds learn to avoid prey that are efficient at escaping. This, however, remains poorly documented. Here, we argue that butterflies may utilize a variety of escape tactics against insectivorous birds and review evidence that birds avoid attacking butterflies that are hard to catch. We suggest that signaling difficulty of capture to predators is a widespread phenomenon in butterflies, and this ability may not be limited to palatable butterflies. The possibility that both palatable and unpalatable species signal difficulty of capture has not been fully explored, but helps explain the existence of aposematic coloration and escape mimicry in butterflies lacking defensive chemicals. This possibility may also change the role that putative Mullerian and Batesian mimics play in a variety of classical mimicry rings, thus opening new perspectives in the evolution of mimicry in butterflies.

The butterfly genus Opoptera includes eight species, three of which have diurnal habits while the others are crepuscular (the usual activity period for members of the tribe Brassolini). Although never measured in the field, it is presumed that diurnal Opoptera species potentially spend more time flying than their crepuscular relatives. If a shift to diurnal habits potentially leads to a higher level of activity and energy expenditure during flight, then selection should operate on increased aerodynamic and energetic efficiency, leading to changes in wing shape. Accordingly, we ask whether diurnal habits have influenced the evolution of wing morphology in Opoptera. Using phylogenetically independent contrasts and Wilcoxon rank sum tests, we confirmed our expectation that the wings of diurnal species have higher aspect ratios (ARs) and lower wing centroids (WCs) than crepuscular congeners. These wing shape characteristics are known to promote energy efficiency during flight. Three Opoptera wing morphotypes established a priori significantly differed in AR and WC values. The crepuscular, cloud forest dweller Opoptera staudingeri (Godman & Salvin) was exceptional in having an extended forewing tip and the highest AR and lowest WC within Opoptera, possibly to facilitate flight in a cooler environment. Our study is the first to investigate how butterfly wing morphology might evolve as a response to a behavioral shift in adult time of activity.

Porto Alegre was already a big city when I was born in 1961. Some time before my parents’ wedding, my grandmother Morena (nee Adelina) had her house remodeled into an upstairs–downstairs duplex, and the upper floor was designed for the newlyweds. She also had the entire backyard paved with a mosaic of uneven pieces of tricolored ceramic tiles. According to my mother Isolde, grandma was tired of the “dirt” that was dragged into the house under everybody’s shoes. A few planters were left along the walls to house Sansevieria, Asparagus, and a Rhapis clump. Introduced European house sparrows roosted on a tall Dracaena that was fragrant at night when in bloom. Grandma also had a few large concrete planters cast to resemble tree trunks where Anthurium plants grew. From our house, the distant noise of a sawmill could be heard all day, and there was also the hustle and bustle of a Volkswagen car dealership located across the street. How could a biologist emerge from such an urban place?

This study provides updated species definitions for five rose-colored Cithaerias butterflies, starting with a historical overview of their taxonomy. Given their mostly transparent wings, genitalia morphology yielded the most reliable characters for species definition and identification. Genitalic divergence is more pronounced when multiple species occur in sympatry than between parapatric taxa. Cithaerias aurorina is granted full species status, C. cliftoni is reinstated as a full species, and one new combination is proposed, i.e. C. aurora tambopata. Two new synonyms are proposed, Callitaera phantoma and Callitaera aura = Cithaerias aurora.

Phylogenetic relationships among Heliconiinae genera are proposed based on early-stage and adult morphology. Parsimony analyses of forty-nine species in twenty-nine genera indicate that Heliconiinae can be divided into four main groups: (1) Pardopsis, Acraea and Actinote; (2) Cethosia plus Neotropical genera; (3) Oriental genera and (4) fritillaries. Analyses of adult characters suggested that Oriental genera form a monophyletic group, whereas those of adult plus early-stage characters artificially split this group into three separate lineages. Our character set does not contain enough phylogenetic information to resolve relationships among fritillary genera with confidence, and further studies of this group are needed. The classification of the Heliconiinae is revised based on our results.

The classification of butterflies in the widely recognized genus Morpho previously used subgenera that were assumed to constitute natural species groups. Cladistic analysis of 120 characters provided a well-resolved tree showing that some subgenera do not constitute monophyletic groups. This study supported some traditional taxonomic species groupings, but rejected the concept of subgenera for Morpho. Therefore, we formally redefined the genus to be consonant with the assumptions of phylogenetic classification. Predictions about Morpho life histories, the correlation of color pattern and flight behavior with vertical flight height, and the evolution of sexual dimorphism are discussed in light of our phylogeny.

A comparative study of over 1,000 specimens allowed us to revise the taxonomy of the Amazonian butterfly genus Bia. We redescribed the genus, and used a selected set of characters to define and describe new species and subspecific taxa. We found that male genitalia showed little variation among taxa, and that wing and genitalia characters varied independently across the range of the genus. We also noted that species diversification seems to follow a north-south Amazonian divide, with Bia actorion and decaerulea occupying the northern portion of the genus range while the remaining four species are found in southern Amazonia. As defined here Bia includes six species: B. actorion (Linnaeus), B. actorion ecuatoria DeVries & Penz, NEW SSP., B. actorion occulta Casagrande & Penz, NEW SSP., B. decaerulea Weymer STAT. NOV., B. decaerulea cayana Simonsen & Penz, NEW SSP., B. decaerulea pallida Casagrande & Penz, NEW SSP., B. rebeli Bryk STAT. NOV., B. rebeli aegina Penz & Simonsen, NEW SSP., B. rebeli acreana Casagrande & Penz, NEW SSP., B. rebeli arikeme Penz & Casagrande, NEW SSP., B. rebeli pareci Penz & DeVries, NEW SSP., B. rebeli cuprea Penz & Casagrande, NEW SSP., B. rebeli tapajos Penz & Simonsen, NEW SSP., B. caelestis Penz & DeVries, NEW SP., B. pucallpa Casagrande & Penz, NEW SP., and B. peruana Röber.

A higher level phylogeny for the passion-vine butterflies (Nymphalidae, Heliconiinae) was generated by cladistic analysis of 146 morphological characters from all life stages. The 24 species studied were selected representatives of the ten currently accepted genera of the sub-tribe Heliconiiti. Analyses of only characters from larvae and pupae did not produce well resolved trees. However, some characters of the immature stages provided critical support for the monophyly of two clades. Analysis of only adult characters yielded a tree that closely resembled that obtained from all data combined. The phylogeny here derived from the combined analysis of early stage and adult characters differed in topology from all previously proposed hypotheses, and supported the monophyly of all currently recognized genera. Characters supporting each clade are described and illustrated, and various hypotheses of phylogenetic relatedness of passion-vine butterfly taxa are discussed.

We demonstrate that the mating patterns of owl butterflies Caligo illioneus (Cramer)and C. oileus (Felder) are leks. During 1993–1994, we recorded distributions of male and female butterflies and larval hostplants in a lowland Neotropical rain forest in Panama. Caligo illioneus males aggregated along forest edges and defended territories against both conspecifics and males of the related species C. oileus, which exhibited similar behaviors. Male perch sites were not associated with hostplant dispersion or the local abundance of females. However, unmated female C. illioneus were observed to arrive and copulate with males on territories that were located near where streams intersected the roadway. We found some evidence that these leks overlap to form multiple-species aggregations. Caligo illioneus and C. oileus used the same sites at similar frequencies during 1993, a pattern that was repeated during 1994. We could not detect if members of different species were being attracted by similar environmental features or if they were effectively attracting one another to the display sites. Independent of population growth, the abundance of males at a particular site was correlated with the abundance of heterospecific males during 1993, but this pattern was not confirmed in 1994. Overlap in the leks serves as evidence against a resource-based “hot-spot” hypothesis of lek formation.

Proboscis length, the length of the tip, the number and length of the various sensilla throughout the proboscis, and the size and shape of the labial palpi were compared in 25 species of pollen-feeding and non-pollen-feeding Heliconiinae (Lepidoptera, Nymphalidae). The mouthparts of pollen-feeding species (all belonging to the genera Heliconius and Laparus ) do not have structures exclusive to them. However, in comparison with non-pollen-feeding Heliconiiti, the pollen-feeding species have a significantly longer proboscis without elongation of the tip-region ; the bristle-shaped sensilla trichodea were found to be significantly more numerous and longer on the proximal and mid-region of the proboscis, while the sensilla of the tip-region are significantly shorter. In addition to these proboscis features, the labial palpi were shorter in the pollen-feeding species, which is likewise possibly associated with pollen-feeding behavior. The biological role of these features is discussed and the evolution of this unique feeding behavior among Lepidoptera is considered in the context of the phylogenetic relationships among genera of Heliconiini.

1. Flight is a key innovation in the evolution of insects that is crucial to their dispersal, migration, territoriality, courtship and predator avoidance. Male butterflies have characteristic territoriality and courtship flight behaviours, and females use a characteristic flight behaviour when searching for host plants. This implies that selection acts on wing morphology to maximize flight performance for conducting important behaviours among sexes. 2. Butterflies in the genus Morpho are obvious components of neotropical forests, and many observations indicate that they show two broad categories of flight behaviour and flight height. Although species can be categorized as using gliding or flapping flight, and flying at either canopy or understorey height, the association of flight behaviour and flight height with wing shape evolution has never been explored. 3. Two clades within Morpho differ in flight behaviour and height. Males and females of one clade inhabit the forest understorey and use flapping flight, whereas in the other clade, males use gliding flight at canopy level and females use flapping flight in both canopy and understorey. 4. We used independent contrasts to answer whether wing shape is associated with flight behaviour and height. Given a single switch to canopy habitation and gliding flight, we compared contrasts for the node at which the switch to canopy flight occurred with the distribution of values in the two focal clades. We found significant changes in wing shape at the transition to canopy flight only in males, and no change in size for either sex. A second node within the canopy clade suggests that other factors may also be involved in wing shape evolution. Our results reinforce the hypothesis that natural selection acts differently on male and female butterfly wing shape and indicate that the transition to canopy flight cannot explain all wing shape diversity in Morpho. 5. This study provides a starting point for characterizing evolution of wing morphology in forest butterflies in the contexts of habitat selection and flight behaviour. Further, these observations suggest that exploring wing shape evolution for canopy and understorey species in other insects may help understand the effects of habitat destruction on biological diversity.

This study examines the phylogenetic relationships among species of the butterfly genera Dynastor and Brassolis using 57 characters from adult morphology and wing coloration. We provide evidence for the monophyly of both genera. The relationships among Dynastor species are well resolved, but we were unable to find informative characters that could resolve the relationships inside Brassolis. We provide diagnoses for Dynastor and Brassolis and all species included in these genera, including illustrations that show geographical variation in wing color. The status of one subspecies is changed to species; Brassolis dinizi d’Almeida, 1956, NEW STATUS.

Penz, Carla M., Devries, Philip J. (2006): Systematic position of Apodemia paucipuncta (Riodinidae), and a critical evaluation of the nymphidiine transtilla. Zootaxa 1190: 1-50, DOI: 10.5281/zenodo.2646145, {"references":["Bremer, K. (1994) Branch support and tree stability. Cladistics, 10, 295-304.","Callaghan, C.J. (1983) A new genus of riodinid butterflies. Bulletin of the Allyn Museum, 78, 1-7.","Callaghan, C.J. & Lamas, G. (2004) Riodinidae. In: Lamas, G. (Ed.) Atlas of Neotropical Lepidoptera. Checklist: Part 4A Hesperioidea-Papilionoidea. Scientific Publishers, Gainesville. p 141-170.","Carpenter, J.M. (1988) Choosing among multiple equally parsimonious cladograms. Cladistics, 7, 351-366.","D'Abrera. B. (1994) Butterflies of the Neotropical region. Part VI, Riodinidae. Hill House, Australia.","DePinna, M.C.C. (1991) Concepts and tests of homology in the cladistic paradigm. Cladistics, 7, 367-394.","DeVries, P. J. 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In certain times of the year, Actinote butterflies can be vastly abundant in Southeastern Brazil, thus representing conspicuous denizens of our fauna. Their wing coloration allows for the recognition of five major patterns, which probably resulted from mimicry. Furthermore, examination of series of field-collected and reared specimens clearly demonstrates a high intra-specific variation in wing color pattern. In concert, inter-specific resemblance (mimicry) and intra-specific variation make it difficult to discriminate Actinote species from each other. This key aims at providing a tool for identification of male Actinote. To that effect, we provide diagnostic characters and illustrations of wings and male genitalia for 22 species. We hope that the clear identification of these species will lead to the discovery of species yet unrecognized in Southeastern Brazil. Em certas épocas do ano, as borboletas do gênero Actinote podem ser muito abundantes no sudeste do Brasil, constituindo, então, elementos conspícuous da nossa fauna. Sua coloração alar permite a separação em cinco padrões básicos, provavelmente relacionados com o mimetismo. Além disso, séries de exemplares coletados no campo ou criados em laboratório demonstram claramente que existe uma grande variabilidade intra-específica na coloração alar destas borboletas. Em conjunto, a semelhança entre as espécies (mimetismo) e variação intra-específica dificultam o reconhecimento das espécies de Actinote. Esta chave tem como objetivo auxiliar a identificação dos machos destas borboletas e, para tanto, são apresentados caracteres diagnósticos, ilustrações das asas e genitália de 22 espécies. Esperamos que a clara identificação destas espécies possa revelar a presença de espécies ainda não reconhecidas no sudeste do Brasil.

This study provides updated species definitions for five rose-colored Cithaerias butterflies, starting with a historical overview of their taxonomy. Given their mostly transparent wings, genitalia morphology yielded the most reliable characters for species definition and identification. Genitalic divergence is more pronounced when multiple species occur in sympatry than between parapatric taxa. Cithaerias aurorina is granted full species status, C. cliftoni is reinstated as a full species, and one new combination is proposed, i.e. C. aurora tambopata . Two new synonyms are proposed, Callitaera phantoma and Callitaera aura = Cithaerias aurora .

This study re-describes and provides a phylogeny for the Brassolini genus Blepolenis, which includes three species from the Brazilian Atlantic forest. A diagnosis and illustrations of habitus and genitalia are given for each species. We compare variation in wing color pattern among the genera Blepolenis, Opsiphanes, Orobrassolis and Mielkella, and discuss the repeated loss of male wing androconial organs within Blepolenis. DNA sequence (nuclear and mitochondrial genes) provided the strongest signal for phylogeny reconstruction, given that Blepolenis species are morphologically homogeneous. Estimated time of divergence between Blepolenis and Opsiphanes dates from the Mid Miocene (ca. 15 million years ago), and was followed by an apparent period of stasis. Extant Blepolenis species seem to have diverged in the Pleistocene (ca. 2.5 mya).

A new species of the brassoline genus Orobrassolis (Lepidoptera, Nymphalidae, Satyrinae) is described based on specimens collected in the early 1900’s from the highland grasslands of Paraná, Brazil. The geological history of these highland grasslands suggests that they underwent climatic fluctuations, with warm climate periods leading to contraction and fragmentation. This fragmentation may have led to the isolation and divergence of southern Orobrassolis populations. Human impact to these areas may have led to the extinction of the species described here.

Two new species of Actinote (Nymphalidae, Heliconiinae, Acraeini) are described from southeastern Brazil. Actinote eberti sp. nov. occurs in the Serra da Mantiqueira region and resembles members of the Actinote black-yellow mimicry group. Actinote pratensis sp. nov. is found in widely scattered points in the rich-soil central S o Paulo transition between montane and semideciduous atlantic forest, and belongs to the orange-yellow mimicry group.

This study describes proboscis morphology and identifies morphometric differences among five species of noctuid moths with different feeding habits (fruit versus nectar-feeding). Morphological and morphometric parameters were analyzed using scanning electron microscopy and light microscopy. Measurements included: galea height in ten sites from base to tip, total proboscis length, and length of the distal region that contains large sensilla styloconica and / or tearing hooks and erectible barbs. Both morphometric and morphological differences were identified among species within and between feeding guilds, and these results are discussed in light of the feeding habits of each species.