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Mutualistic interactions between butterflies and ants can evolve into complex social parasitism. `Cuckoo¿ caterpillars, known only in the Lycaenidae, use multimodal mimetic traits to achieve social integration into ant societies. Here, we present the first known `cuckoo¿ butterfly in the family Riodinidae. Aricoris arenarum remained in taxonomic limbo for > 80 years, relegated to nomen dubium and misidentified as Aricoris gauchoana. We located lost type material, designated lectotypes and documented the morphology and natural history of the immature stages. The multifaceted life cycle of A. arenarum can be summarized in three phases: (1) females lay eggs close to honeydew-producing hemipterans tended by specific Camponotus ants; (2) free-living caterpillars feed on liquids (honeydew and ant regurgitations); and (3) from the third instar onward, the caterpillars are fed and tended by ants as `cuckoos¿ inside the ant nest. This life cycle is remarkably similar to that of the Asian lycaenid Niphanda fusca, despite divergence 90 Mya. Comparable eco-evolutionary pathways resulted in a suite of ecomorphological homoplasies through the ontogeny. This study shows that convergent interactions can be more important than phylogenetic proximity in shaping functional traits of social parasites., Financial support for this research was provided by National Geographic Society (WW-224R-17) and a Programa Nacional de Pos-Doutorado (PNPD)-Coordenação de Aperfeicoamento de Pessoal de Nivel Superior (CAPES) fellowship to L.A.K., and by the Spanish Ministerio de Ciencia, Innovación y Universidades (PRX19/00067 and PID2019-107078GB-I00 / AEI / 10.13039/501100011033) to R.V. This paper is dedicated to Ivone Erichsen (1937–2019) and Beatriz D. Ponce (1933–2016) for fundamental symbiotic support to L.A.K. and L.V., respectively.

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...

Tropical fruit-feeding nymphalid butterflies generally restrict adult foraging exclusively to either the canopy or understory strata. We compared canopy and understory butterfly communities using data from four long-term studies in Central and South America. At all study sites we found little similarity in species composition between canopy and understory, with most species showing a strong affinity for one of the two habitats. There was a consistent phylogenetic signal for canopy and understory association, suggesting a substantial evolutionary history with these habitats. In addition to compositional differences, we found different patterns of beta diversity between canopy and understory communities. Across all study sites, the canopy had greater temporal and spatial beta diversity compared to the understory. Although these two communities are composed of the same feeding guild and separated only by a relatively small vertical space, each has its own stratum-specific species composition and community dynamics.

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.

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.

Tropical forest insects are vertically stratified between the canopy and understorey. Using 60 traps set at two heights above the forest floor (30 at 15 cm and 30 at 1 m) we compared abundances in capture height, persistence in traps, and sex of three co-occurring understorey butterflies (Cithaerias pireta,Dulcedo politaandPierella helvina) in Costa Rica. We captured, marked and released 283 individual butterflies (65C. pireta, 79D. polita, 139P. helvina) and showed all three species were captured more often in low traps, andP. helvinawas captured only in low traps. The probability of remaining in traps for 24 h did not differ significantly forD. politaandP. helvina, but was significantly lower forC. pireta. The odds of trapping either sex did not differ significantly forP. helvinaandC. pireta, but they were significantly lower forD. politamales. We experimentally demonstrate that these co-occurring species fly and feed just above the forest floor, but differ with respect to their persistence in traps and attraction to traps by sex. Our study implies that closely related species can exhibit behavioural differences that may influence population abundance estimates in multi-species studies.

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.

Summary 1. We develop a Bayesian method for analysing mark–recapture data in continuous habitat using a model in which individuals movement paths are Brownian motions, life spans are exponentially distributed and capture events occur at given instants in time if individuals are within a certain attractive distance of the traps. 2. The joint posterior distribution of the dispersal rate, longevity, trap attraction distances and a number of latent variables representing the unobserved movement paths and time of death of all individuals is computed using Gibbs sampling. 3. An estimate of absolute local population density is obtained simply by dividing the Poisson counts of individuals captured at given points in time by the estimated total attraction area of all traps. Our approach for estimating population density in continuous habitat avoids the need to define an arbitrary effective trapping area that characterized previous mark–recapture methods in continuous habitat. 4. We applied our method to estimate spatial demography parameters in nine species of neotropical butterflies. Path analysis of interspecific variation in demographic parameters and mean wing length revealed a simple network of strong causation. Larger wing length increases dispersal rate, which in turn increases trap attraction distance. However, higher dispersal rate also decreases longevity, thus explaining the surprising observation of a negative correlation between wing length and longevity.

We present the first dated higher-level phylogenetic and biogeographic analysis of the butterfly family Riodinidae. This family is distributed worldwide, but more than 90% of the c. 1500 species are found in the Neotropics, while the c. 120 Old World species are concentrated in the Southeast Asian tropics, with minor Afrotropical and Australasian tropical radiations, and few temperate species. Morphologically based higher classification is partly unresolved, with genera incompletely assigned to tribes. Using 3666 bp from one mitochondrial and four nuclear markers for each of 23 outgroups and 178 riodinid taxa representing all subfamilies, tribes and subtribes, and 98 out of 145 described genera of riodinids, we estimate that Riodinidae split from Lycaenidae about 96 Mya in the mid-Cretaceous and started to diversify about 81 Mya. The Riodinidae are monophyletic and originated in the Neotropics, most likely in lowland proto-Amazonia. Neither the subfamily Euselasiinae nor the Nemeobiinae are monophyletic as currently constituted. The enigmatic, monotypic Neotropical genera Styx and Corrachia (most recently treated in Euselasiinae: Corrachiini) are highly supported as derived taxa in the Old World Nemeobiinae, with dispersal most likely occurring across the Beringia land bridge during the Oligocene. Styx and Corrachia, together with all other nemeobiines, are the only exclusively Primulaceae-feeding riodinids. The steadily increasing proliferation of the Neotropical Riodininae subfamily contrasts with the decrease in diversification in the Old World, and may provide insights into factors influencing the diversification rate of this relatively ancient clade of Neotropical insects., The study was funded by grant #204308 from the Research Council of Norway to M.E., and grants from the Baker Foundation, the Putnam Expeditionary Fund of the Museum of Comparative Zoology, and NSFDEB-0447242 to N.E.P. D.C.L. was funded by INRA and by ERC EMARES #250325 during writing. L.-W.W. and Y-F.H. were funded by NSC97-2621-B-003-003-MY3 and COA101-CF-02.1-C12.

1. Documenting species abundance distributions in natural environments is critical to ecology and conservation biology. Tropical forest insect faunas vary in space and time, and these partitions can differ in their contribution to overall species diversity. 2. In the Neotropics, the Central American butterfly fauna is best known in terms of general natural history, but butterfly community diversity is best documented by studies on South American fruit-feeding butterflies. Here, we present the first long-term study of fruit-feeding nymphalid species diversity from Central America and provide a unique comparison between Central and South American butterfly communities. 3. This study used 60 months of sampling among multiple spatial and temporal partitions to assess species diversity in a Costa Rican rainforest butterfly community. Abundance distributions varied significantly at the species and higher taxonomic group levels, and canopy and understorey samples were found to be composed of distinct species assemblages. 4. Strong similarities in patterns of species diversity were found between this study and one from Ecuador; yet, there was an important difference in how species richness was distributed in vertical space. In contrast to the Ecuadorian site, Costa Rica had significantly higher canopy richness and lower understorey richness. 5. This study affirms that long-term sampling is vital to understanding tropical insect species abundance distributions and points to potential differences in vertical structure among Central and South American forest insect communities that need to be explored.

Aim Differentiation of sites or communities is often measured by partitioning regional or gamma diversity into additive or multiplicative alpha and beta components. The beta component and the ratio of within-group to total diversity (alpha/gamma) are then used to infer the compositional differentiation or similarity of the sites. There is debate about the appropriate measures and partitioning formulas for this purpose. We test the main partitioning methods, using empirical and simulated data, to see if some of these methods lead to false conclusions, and we show how to resolve the problems that we uncover. Location South America, Ecuador, Orellana province, Rio Shiripuno. Methods We construct sets of real and simulated tropical butterfly communities that can be unambiguously ranked according to their degree of differentiation. We then test whether beta and similarity measures from the different partitioning approaches rank these datasets correctly. Results The ratio of within-group diversity to total diversity does not reflect compositional similarity, when the Gini–Simpson index or Shannon entropy are used to measure diversity. Additive beta diversity based on the Gini–Simpson index does not reflect the degree of differentiation between N sites or communities. Main conclusions The ratio of within-group to total diversity (alpha/gamma) should not be used to measure the compositional similarity of groups, if diversity is equated with Shannon entropy or the Gini–Simpson index. Conversion of these measures to effective number of species solves these problems. Additive Gini–Simpson beta diversity does not directly reflect the differentiation of N samples or communities. However, when properly transformed onto the unit interval so as to remove the dependence on alpha and N, additive and multiplicative beta measures yield identical normalized measures of relative similarity and differentiation.

Aim We explore the potential role of the ‘tropical conservatism hypothesis’in explaining the butterfly species richness gradient in North America. Itsapplicability can be derived from the tropical origin of butterflies and thepresumed difficulties in evolving the cold tolerance required to permit thecolonization and permanent occupation of the temperate zone.Location North America.Methods Digitized range maps for butterfly species north of Mexico were usedto map richness for all species, species with distributions north of the Tropic ofCapricorn (Extratropicals), and species that also occupy the tropics (Tropicals).A phylogeny resolved to subfamily was used to map the geographical pattern ofmean root distance, a metric of the evolutionary development of assemblages.Regression models and general linear models examined environmental correlatesof overall richness and for Extratropicals vs. Tropicals, patterns in summer vs.winter, and patterns in northern vs. southern North America.Results Species in more basal subfamilies dominate the south, whereas morederived clades occupy the north. There is also a ‘latitudinal’ richness gradient inCanada/Alaska, whereas in the conterminous USA richness primarily varieslongitudinally. Overall richness is associated with broad- and mesoscaletemperature gradients. The richness of Tropicals is strongly associated withtemperature and distance from winter population sources. The richness ofExtratropicals in the north is most strongly correlated with the pattern of glacialretreat since the more recent Ice Age, whereas in the south, richness is positivelyassociated with the range of temperatures in mountains and the presence offorests but is negatively correlated with the broad-scale temperature gradient.Main conclusions The tropical conservatism hypothesis provides a possibleexplanation for the complex structure of the species richness gradient. TheCanada/Alaska fauna comprises temperate, boreal and tundra species that arenevertheless constrained by cold climates and limited vegetation, coupled withpossible post-Pleistocene recolonization lags. In the USA tropical species areconstrained by temperature in winter as well as recolonization distancesin summer, whereas temperate-zone groups are richer in cooler climates inmountains and forests, where winter conditions are more suitable for diapause.The evolution of cold tolerance is key to both the evolutionary and ecologicalpatterns.KeywordsButterflies, cold tolerance, diapause, diversity gradients, latitudinal gradients,niche conservatism, North America, Rhopalocera, species richness gradients,tropical conservatism hypothesis.

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.

We describe the immature stages Pedaliodes poesia Hewitson, 1862 from northeastern Ecuador. Chusquea scandens (Poaceae, Bambusoidea) is the larval food plant. Eggs are laid singly or in pairs on the bottom side of host plant leaves. The duration of the egg, larval, and pupal stages, combined, is 99-107 days.

1 Butterflies are frequently used to investigate Neotropical diversity, but the family Hesperiidae is almost never employed as a focal group. Sampling Hesperiidae with artificial lures has been used to assist in species richness estimates of males that use bird droppings as a resource, but its effectiveness for estimating total hesperiid site diversity is unknown. 2 This study characterises species richness and abundance in a diverse assemblage of Hesperiidae sampled at artificial lures. Sampled and estimated richness among taxonomic subsets were compared to the known site richness. These comparisons are used to assess artificial lures as a method for sampling total richness of Hesperiidae. We tested for potential differences in attraction in lures placed with and without the presence of army ant swarms. 3 Five years of intermittent sampling with lures recovered 65% of the known Hesperiidae species richness at the study site. We found the subfamilies were differentially attracted to lures, where 86% of Eudaminae, 56% of Hesperiinae and 55% of Pyrginae species known from the site were recovered by lures. Species accumulation and taxonomic diversity differed among lures placed alone, and those placed in the presence of army ants. 4 Our results suggest that sampling with artificial lures can be used to estimate diversity of Neotropical Eudaminae and a few tribes of Hesperiinae at some sites. Available evidence indicates that Hesperiidae feeding at bird droppings is predominately a Neotropical phenomenon. Finally, we conclude that estimating diversity in Neotropical Hesperiidae will require long-term studies that use multiple sampling techniques.

A sample of over 400 co-occurring species of Neotropical Hesperiidae attracted to artificial lures was used to test for temporal structure among species and explore potential consequences of temporal partitioning within assemblages. Five years of intermittent transect samples at the same site revealed that diel activity was significantly partitioned among subfamilies, genera and species within this species assemblage. Calculations of reproductive isolation indicate that partitions in activity times are sufficient to provide significant prezygotic reproductive isolation among some congeneric species pairs. Temporal patterns in this assemblage are considered in light of field observations and temporal partitions are examined as a potential isolating barrier among insects. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society 2008, 94, 723–736.

Adult butterflies feed from a variety of substrates and have appropriate adaptations. We examined proboscis morphology in a community of fruit-feeding butterflies (Nymphalidae) in a tropical forest in Uganda. These data were supplemented with behavioural observations and measurements of intake rate on natural and artificial substrates. We found no sexual dimorphism in proboscis morphology even though puddling behaviour is usually performed by males. Two main feeding techniques could be distinguished on the basis of behaviour and morphology: the piercing technique, typically found in Charaxinae, and the sweeping technique employed by both Nymphalinae and Satyrinae. These techniques, distinguished in previous studies, are described in more detail and their relative efficiencies are discussed in the context of sexual dimorphism, food-choice and life history evolution. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86, 333–343.

We present a novel analytical method for assessing spatial and temporal structure in community samples that is useful for comparing large data-sets that include species abundance data. The model assumes that species numbers in two samples are drawn from a bi-variate Poisson log-normal species abundance distribution and parameters from the fitted distribution are estimated to assess community structure. We assessed three tropical butterfly data-sets for spatial structure in the vertical dimension, and tested for changes in structure as a result of temporal variance, disturbance regimes, and geographic location. Our results indicate that the vertical dimension is a major structural component in tropical forest butterfly communities that varies little through time and is not measurably affected by small-scale disturbances. However, there is evidence that the degree of vertical structure may vary among geographic regions. These results are discussed in terms of the mechanisms maintaining vertical structure, and the implications of changes in forest architecture on butterfly communities.

Aerodynamic theory predicts that migrant fliers should reduce their speed of flight as endogenous energy reserves are gradually consumed. This prediction was tested for Urania fulgens (Walker) moths migrating through central Panama in 1987 and again in 1998. Direct airspeed measurements together with chloroform:methanol extractions of abdominal lipids were used to determine that, contrary to theoretical predictions, the speed of flight among individuals was independent of both body mass and abdominal lipid mass.

The joint spatial and temporal fluctuations in the community structure of tropical butterflies are analyzed by fitting the bivariate Poisson lognormal distribution to a large number of observations in space and time. By applying multivariate dependent diffusions for describing the fluctuations in the abundances, the environmental variance is estimated to be very large and so is the strength of local density regulation. The variance in the lognormal species abundance distribution is partitioned into components expressing the heterogeneity between the species, independent noise components for the different species, a demographic stochastic component, and a component due to overdispersion in the sampling. In disagreement with the neutral theory, the estimates show that the heterogeneity component is the dominating one, representing 81% of the total variance in the lognormal model. Different spatial components of diversity, the alpha, beta, and gamma diversity, are also estimated. The spatial scale of the autocorrelation function for the community is of order 1 km, while sampling of a quadrat would need to be 10 km on a side to yield the total diversity for the community.

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.

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.

To test the veracity of previous studies and illuminate major community patterns from an intact community, a guild of nymphalid butterflies was sampled at monthly intervals for five consecutive years by trapping in the canopy and understorey of five contiguous forest plots in the same rainforest. Significant numbers of species belonged to either the canopy or understorey fauna, confirming fundamental vertical stratification, and showing that sampling in one vertical position is a poor estimator of diversity. Significant monthly variation showed that intermittent or short-term sampling would underestimate diversity, and significant variation among years and areas showed that diversity was strongly influenced by sampling year. Even when the underlying communities were the same, temporal interactions strongly affected species diversity in both horizontal and vertical dimensions. An unprecedented seasonal inversion of species richness and abundance was detected between the canopy and understorey that occurred at the onset of all rainy seasons. This investigation suggests that long-term studies evaluating spatial and temporal patterns of species diversity among many sites may be required for a better understanding of tropical communities and how best to conserve them.

Diversity in biological communities frequently is compared using species accumulation curves, plotting observed species richness versus sample size. When species accumulation curves intersect, the ranking of communities by observed species richness depends on sample size, creating inconsistency in comparisons of diversity. We show that species accumulation curves for two communities are expected to intersect when the community with lower actual species richness has higher Simpson diversity (probability that two random individuals belong to different species). This may often occur when comparing communities that differ in habitat heterogeneity or disturbance, as we illustrate using data from neotropical butterflies. In contrast to observed species richness, estimated Simpson diversity always produces a consistent expected ranking among communities across sample sizes, with the statistical accuracy to confidently rank communities using small samples. Simpson diversity should therefore be particularly useful in rapid assessments to prioritize areas for conservation.

To test the hypotheses that butterflies in an intact lowland rainforest are randomly distributed in space and time, a guild of nymphalid butterflies was sampled at monthly intervals for one year by trapping 883 individuals of 91 species in the canopy and understory of four contiguous, intact forest plots and one naturally occurring lake edge. The overall species abundance distribution was well described by a log-normal distribution. Total species diversity (γ-diversity) was partitioned into additive components within and among community subdivisions (α-diversity and β-diversity) in vertical, horizontal and temporal dimensions. Although community subdivisions showed high similarity (1-β-diversity/γ-diversity), significant β-diversity existed in each dimension. Individual abundance and observed species richness were lower in the canopy than in the understory, but rarefaction analysis suggested that the underlying species richness was similar in both canopy and understory. Observed species richness varied among four contiguous forest plots, and was lowest in the lake edge plot. Rarefaction and species accumulation curves showed that one forest plot and the lake edge had significantly lower species richness than other forest plots. Within any given month, only a small fraction of total sample species richness was represented by a single plot and height (canopy or understory). Comparison of this study to a similar one done in disturbed forest showed that butterfly diversity at a naturally occurring lake edge differed strongly from a pasture-forest edge. Further comparison showed that species abundance distributions from intact and disturbed forest areas had variances that differed significantly, suggesting that in addition to extrapolation, rarefaction and species accumulation techniques, the shapes of species abundance distributions are fundamental to assessing diversity among sites. This study shows the necessity for long-term sampling of diverse communities in space and time to assess tropical insect diversity among different areas, and the need of such studies is discussed in relation to tropical ecology and quick surveys in conservation biology.

To test whether ithomiine butterfly species within Mullerian mimetic classes are associated in space and time, we sampled a community of ithomiine butterflies at monthly intervals with traps in the canopy and the understory of four forest habitats: primary, higrade, secondary and edge. A species accumulation curve reached an asymptote at 22 species, suggesting that these species have a greater preference for feeding on fruit juices than other ithomiines known to occur at the study site. Species richness and individual abundance showed marked temporal variation, and there were slight differences in the distribution of species richness and individual abundance among the four habitats. The 22 species sampled in this study were not stratified vertically. The five mimetic colour classes of these butterflies were unequally distributed among the four habitats and over the course of the twelve months. There is suggestive evidence that co-mimic species occurred in the same habitats, and strong evidence that they occurred at the same times. Habitat and temporal effects each contributed approximately 10% to the total mimetic class diversity, with the temporal effect being slightly larger than that of habitat. This study demonstrates that Mullerian co-mimic associations can be measured on a much smaller scale than has been done previously.

To test the hypotheses that fruit-feeding nymphalid butterflies are randomly distributed in space and time, a community of fruit-feeding nymphalid butterflies was sampled at monthly intervals for one year by trapping 6690 individuals of 130 species in the canopy and understory of four forest habitats: primary, higraded, secondary, and edge. The overall species abundance distribution was well described by a lognormal distribution. Total species diversity (γ-diversity) was partitioned into additive components within and among community subdivisions (α-diversity andβ-diversity) in vertical, horizontal and temporal dimensions. Although community subdivisions showed high similarity (1−β-diversity/γ-diversity), significantβ-diversity existed in each dimension. Individual abundance and observed species richness was lower in the canopy than in the understory. However, rarefaction analysis and species accumulation curves revealed that canopy had higher species richness than understory. Observed species richness was roughly equal in all habitats, but individual abundance was much greater in edge, largely due to a single, specialist species. Rarefaction analysis and species accumulation curves showed that edge had significantly lower species richness than all other habitats. Samples from a single habitat, height and time contained only a small fraction of the total community species richness. This study demonstrates the feasibility, and necessity, of large-scale, long-term sampling in multiple dimensions for accurately measuring species richness and diversity in tropical forest communities. We discuss the importance of such studies in conservation biology.

1. We develop a Bayesian method for analysing mark-recapture data in continuous habitat using a model in which individuals movement paths are Brownian motions, life spans are exponentially distributed and capture events occur at given instants in time if individuals are within a certain attractive distance of the traps. 2. The joint posterior distribution of the dispersal rate, longevity, trap attraction distances and a number of latent variables representing the unobserved movement paths and time of death of all individuals is computed using Gibbs sampling. 3. An estimate of absolute local population density is obtained simply by dividing the Poisson counts of individuals captured at given points in time by the estimated total attraction area of all traps. Our approach for estimating population density in continuous habitat avoids the need to define an arbitrary effective trapping area that characterized previous mark-recapture methods in continuous habitat. 4. We applied our method to estimate spatial demography parameters in nine species of neotropical butterflies. Path analysis of interspecific variation in demographic parameters and mean wing length revealed a simple network of strong causation. Larger wing length increases dispersal rate, which in turn increases trap attraction distance. However, higher dispersal rate also decreases longevity, thus explaining the surprising observation of a negative correlation between wing length and longevity.

1. Documenting species abundance distributions in natural environments is critical to ecology and conservation biology. Tropical forest insect faunas vary in space and time, and these partitions can differ in their contribution to overall species diversity. 2. In the Neotropics, the Central American butterfly fauna is best known in terms of general natural history, but butterfly community diversity is best documented by studies on South American fruit-feeding butterflies. Here, we present the first long-term study of fruit-feeding nymphalid species diversity from Central America and provide a unique comparison between Central and South American butterfly communities. 3. This study used 60 months of sampling among multiple spatial and temporal partitions to assess species diversity in a Costa Rican rainforest butterfly community. Abundance distributions varied significantly at the species and higher taxonomic group levels, and canopy and understorey samples were found to be composed of distinct species assemblages. 4. Strong similarities in patterns of species diversity were found between this study and one from Ecuador; yet, there was an important difference in how species richness was distributed in vertical space. In contrast to the Ecuadorian site, Costa Rica had significantly higher canopy richness and lower understorey richness. 5. This study affirms that long-term sampling is vital to understanding tropical insect species abundance distributions and points to potential differences in vertical structure among Central and South American forest insect communities that need to be explored.

An acoustical comparison between calls of parasitic butterfly caterpillars and their host ants is presented for the first time. Overall, caterpillar calls were found to be similar to ant calls, even though these organisms produce them by different means. However, a comparison of Maculinea caterpillars with those of Myrmica ants produced no evidence suggesting fine level convergence of caterpillar calls upon those of their species specific host ants. Factors mediating the species specific nature of the Maculinea-Myrmica system are discussed, and it is suggested that phylogenetic analysis is needed for future work.

A facultative mutualism between the riodinid butterfly Thisbe ircnca and the ponerine ant Ectafomma ruidurn is described from Panama. Ants protect larvae against attacks of predatory wasps, but not against tachinid parasitoids. Several potential sources of ecological variation affecting the larval survival of Thisbe irenea are noted. A preliminary means of testing the ability of larvae to appease ants is described that may be applied to all butterfly-ant systems. Observations and literature records indicate that ant taxa which tend butterfly larvae are the same taxa that tend extrafloral nectaries and Homoptera. A general hypothesis for the evolution of myrmecophily among butterflies suggests that ant taxa which utilize secretions in their diet are major selective agents for the evolution of the larval ant-organs, and hence, ant-larval mutualisms. This idea is extended to suggest how appeasement of predaceous ant taxa through the use of larval ant-organs can influence an ant-larval relationship, eventually leading to mutualism.

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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 .

Entomophagy in the riodinid butterfly tribe Eurybiini is demonstrated for the first time. Alesa amesis caterpillars and adults possess behavioral and morphological traits for feeding on Homoptera nymphs, and oviposition by A. amesis females is mediated by the combined presence of Camponotus femoratus ants and homopteran nymphs. Caterpillars are entirely entomophagous, and do not eat plant tissues. Alesa amesis caterpillars have distinct behaviors for feeding on their prey, and for soliciting and drinking honeydew secretions from homopteran nymphs. The leg lengths of entomophagous Alesa caterpillars are shown to be longer than phytophagous relatives. The legs of Alesa are used for prey handling and soliciting honeydew secretions. We suggest that elongation of the thoracic legs has been a general consequence of entomophagy in butterfly caterpillars. This study clarifies our understanding of A. amesis and its interactions with multiple species, and points to behavioral and morphological traits impor...

Gregarious roosting behaviour and nocturnal activity is virtually unknown within the family Hesperiidae. Observations on Celaenorrhinus fritzgaertneri (Bailey) from Guanacaste Province, Costa Rica, are presented which demonstrate that, during the dry season, individuals roost gregariously in cave-like situations during the day and are nocturnally active. Circadian behaviour is highly synchronized with regard to timing of departures and arrivals at the roosting site, and appears to be regulated by light levels. Individuals on the roost are in reproductive diapause and show marked roost-site fidelity and longevity. The break-up of roosts appears to be mediated by the onset of the first rains of the season. These observations are compared and discussed in relation to gregarious roosting in other Lepidoptera. Two possible life history strategies to explain roosting in C. fritzgaertneri, with appropriate experiments, are suggested for future studies.

Summary The first case is reported of a non-lethal interaction between a butterfly and tachinid which had little effect on host or parasite, and is discussed in its implications in relation to tropical tachinids and their hosts.

The surface ultrastructure of the setae and epidermal organs of the larva of Curetis regula is described. Their possible functions are discussed and comparisons made with similar structures found i...

Based on a Wagner tree analysis of ninety-two characters (eighty-five larval, one egg, six adult), the nymphalid butterfly genera Antirrhea and Caerois are demonstrated to be the closest relatives of the genus Morpho. Accordingly, Antirrhea and Caerois are formally transferred from the Satyrinae to the Morphinae. Without these two genera, the Morphinae (sensu Ehrlich) is at best a paraphyletic group. During the study, fourteen nymphalid genera were treated as potential outgroups. The analysis suggests that many currently accepted higher taxa within the Nymphalidae are probably untenable: the Satyrinae and Nymphalinae (sensu Ehrlich) are both probably polyphyletic; the Biinae (sensu Miller) must be abandoned, being polyphyletic; the Charaxidae (sensu Rydon), although probably monophyletic, appear to form a group subordinate to part of the ‘Satyrinae’; and Apatura does not cluster with the ‘Nymphalinae’, but appears to form the sister-group of the ‘Satyrinae’ (less Antirrhea and Caerois) plus the Charaxinae. Re-analyses of reduced data sets, in which potentially homoplasious larval head-horn and adult wing venational characters were eliminated, leaves these conclusions essentially unaltered. The authors suggest that a solution to the seemingly intractable problem posed by nymphalid higher classification can be sought by the application of cladistic analysis to a large data set gathered from all developmental stages, with special emphasis on detailed comparative larval morphology.

The neotropical riodinid butterfly Sarota gyas is shown to utilize non-nitrogen fixing epiphylls as larval hostplants. The host substrate where the epiphylls grow appears unimportant to oviposition by adults or feeding by larvae. The larva of S. gyas is described and the use of long setae as a defense by amymecophilous riodinids is discussed.

The lar\ae or thc lycaenid subfamily Curetinae have never been reported to be associated with ants. Observations on Cureti3 regulu Evans from Brunei are presented which show that this species may be tended by ants both as larvae and adults. The observations are discussed in relation to a rccent review on lycaenid/ant associations. IL is suggested that the Curetinae will be found to be associated with ants when more species have been reared, on evidence of the larval tentacle organs and apparent ‘pore cupolas’, both of which are ant adaptations. More studies are needed on (hrurptz~ biology and larval morphology to resolve the relationships of this enigmatic genus within the Lycaenidae.

The generic names Euptychia, Cissia and Argyreuplychia are reviewed in relation to the Cissia confusa species-group in Costa Rica and Trinidad, for which the generic name Cissia Doubleday is formally resurrected. Three new species of Cissia are described, C. confusa itself is resurrected from synonymy, three neotypes and three lectotypes are designated, and six new combinations established. On the basis of early instar morphology three subgroups are recognized and the general biology of the confusa species-group is summarized. A key to species in Costa Rica is provided and the ecology of the adults is reviewed.

Observations and experiments are presented on the use of larval ant-organs by the riodinid butterfly Thisbe irenea with emphasis on their function in myrmecophily. The results indicate that each ant organ plays a distinct role in larval-ant association and that all influence the behaviour of attending ants. Interpretations of the roles that lycaenid larval organs play in ant associations are evaluated and discussed in light of myrmecophilous riodinids. Finally, an ‘enticement and binding’ process, involving the concerted use of the larval organs, is proposed as the behavioural mechanism that 7. irenea and other riodinid larvae use to secure the attentions of individual ants for extended periods of time.