Dinâmica espaço-temporal em uma guilda de borboletas frugívoras no limite norte da floresta atlântica nordestina

The extent of the Brazilian Atlantic rainforest, a global biodiversity hotspot, has been reduced to less than 7% of its original range. Yet, it contains one of the richest butterfly fauna in the world. Butterflies are commonly used as environmental indicators, mostly because of their strict association with host plants, microclimate and resource availability. This research describes diversity, composition and species richness of frugivorous butterflies in a forest fragment in the Brazilian Northeast. It compares communities in different physiognomies and seasons. The climate in the study area is classified as tropical rainy, with two well defined seasons. Butterfly captures were made with 60 Van Someren-Rydon traps, randomly located within six different habitat units (10 traps per unit) that varied from very open (e.g. coconut plantation) to forest interior. Sampling was made between January and December 2008, for five days each month. I captured 12090 individuals from 32 species. The most abundant species were Taygetis laches, Opsiphanes invirae and Hamadryas februa, which accounted for 70% of all captures. Similarity analysis identified two main groups, one of species associated with open or disturbed areas and a second by species associated with shaded areas. There was a strong seasonal component in species composition, with less species and lower abundance in the dry season and more species and higher abundance in the rainy season. K-means analysis indicates that choice of habitat units overestimated faunal perceptions, suggesting less distinct units. The species Taygetis virgilia, Hamadryas chloe, Callicore pygas e Morpho achilles were associated with less disturbed habitats, while Yphthimoides sp, Historis odius, H. acheronta, Hamadryas feronia e Siderone marthesia likey indicate open or disturbed habitats. This research brings important information for conservation of frugivorous butterflies, and will serve as baseline for future projects in environmental monitoring

Movimento de borboletas frugívoras e conectividade funcional em uma paisagem de Mata Atlântica

An organisms movement within and between habitats is an essential trait of life history, one that shapes population dynamics, communities and ecosystems in space and time. Since the ability to perceive and react to specific conditions varies greatly between organisms, different movement patterns are generated. These, in turn, will reflect the way species persist in the original habitat and surrounding patches. This study evaluated patterns of movement of frugivorous butterflies in order to estimate the connectivity of a landscape mosaic in an area of Atlantic Forest. For this purpose, we used the capture-mark-recapture method on butterflies trapped with fermented fruit bait in three distinct habitats. The first represents a typical Atlantic forest fragment, while the other two represent man-made matrix habitats. One contains a coconut plantation and the other a plantation of the exotic Acacia mangium species. Five traps were randomly placed in each landscape unit in areas of 40 x 40m. Using recapture data and relating it to distance between captures and habitat structure, I found that movement frequencies, both within and between landscape units were different for the analyzed species, suggesting that they do not interpret and react to the landscape in the same way. Thus this study was able to measure landscape functional connectivity. For most species, the exchange between forest and coconut plantations occurred with low frequency compared to exchanges between the forest and acacia plantations, which share more structural similarities. This seems to indicate that a matrix that is more similar to patches of native vegetation can shelter species, permit their movement and, consequently, contribute to the landscape connectivity

Borboletas no semiárido: sazonalidade e padrões de diversidade de borboletas frugívoras em um ambiente extremo

The spatial and temporal distribution of the population reflects the adjustment of their biological characteristics to environmental conditions and biotic interactions as adaptive and phylogenetic precursors elements. The habitat’s heterogeneity and alternating seasons tend to cause patterns of activity of organisms and species diversity. However, these seasonal and spatial patterns in butterfly communities in dry environments are not yet clear. We studied a community of frugivorous butterflies in ESEC Seridó, in northeastern Brazil, aiming to characterize the guild in semiarid and check the relative contribution of climate and vegetation variables on its composition, diversity and phenofaunistic. The butterflies were sampled monthly during one year, and the distribution of species was associated with structural characteristics of three vegetation types (eg. richness and abundance of tree and shrub species, canopy cover, herbaceous cover, litter) and climatological data (temperature, rainfall and humidity). We captured 9580 individuals of 16 species of butterflies belonging to four subfamilies (Biblidinae, Charaxinae, Nymphalinae and Satyrinae). The richness, abundance and diversity varied in different scales, especially in time, being higher in the rainy season, while the β-diversity and turnover was higher in the dry. The distribution of species mainly followed the changes in humidity, rainfall and vegetation phenology, with no defined boundaries between habitats. The flight period was shared within subfamilies, which should have distinct response to environmental stimuli, as well as respond to the phenology of host plants and have different reproductive strategies. There is even evidence of physiological and behavioral adaptations as seasonal reproduction and aestivation. So there was environmental control over the distribution and diversity of species, with the key role climate Association and vegetation structure in the community of differentiation in the seasons, and the availability and quality of resources on the variation of species abundance in small scales. These results may support the biomonitoring and conservation preserved areas, particularly in environments under human pressure and extreme environmental conditions such as semi-arid.

Dinâmica espaço-temporal em uma guilda de borboletas frugívoras no limite norte da Floresta Atlântica Nordestina

The extent of the Brazilian Atlantic rainforest, a global biodiversity hotspot, has been reduced to less than 7% of its original range. Yet, it contains one of the richest butterfly fauna in the world. Butterflies are commonly used as environmental indicators, mostly because of their strict association with host plants, microclimate and resource availability. This research describes diversity, composition and species richness of frugivorous butterflies in a forest fragment in the Brazilian Northeast. It compares communities in different physiognomies and seasons. The climate in the study area is classified as tropical rainy, with two well defined seasons. Butterfly captures were made with 60 Van Someren-Rydon traps, randomly located within six different habitat units (10 traps per unit) that varied from very open (e.g. coconut plantation) to forest interior. Sampling was made between January and December 2008, for five days each month. I captured 12090 individuals from 32 species. The most abundant species were Taygetis laches, Opsiphanes invirae and Hamadryas februa, which accounted for 70% of all captures. Similarity analysis identified two main groups, one of species associated with open or disturbed areas and a second by species associated with shaded areas. There was a strong seasonal component in species composition, with less species and lower abundance in the dry season and more species and higher abundance in the rainy season. K-means analysis indicates that choice of habitat units overestimated faunal perceptions, suggesting less distinct units. The species Taygetis virgilia, Hamadryas chloe, Callicore pygas e Morpho achilles were associated with less disturbed habitats, while Yphthimoides sp, Historis odius, H. acheronta, Hamadryas feronia e Siderone marthesia likey indicate open or disturbed habitats. This research brings important information for conservation of frugivorous butterflies, and will serve as baseline for future projects in environmental monitoring

Gene duplication is an evolutionary mechanism for expanding spectral diversity in the long-wavelength photopigments of butterflies

Butterfly long-wavelength (L) photopigments are interesting for comparative studies of adaptive evolution because of the tremendous phenotypic variation that exists in their wavelength of peak absorbance (lambda(max) value). Here we present a comprehensive survey of L photopigment variation by measuring lambda(max) in 12 nymphalid and 1 riodinid species using epi-microspectrophotometry. Together with previous data, we find that L photopigment lambda(max) varies from 510-565 nm in 22 nymphalids, with an even broader 505- to 600-nm range in riodinids. We then surveyed the L opsin genes for which lambda(max) values are available as well as from related taxa and found 2 instances of L opsin gene duplication within nymphalids, in Hermeuptychia hermes and Amathusia phidippus, and 1 instance within riodinids, in the metalmark butterfly Apodemia mormo. Using maximum parsimony and maximum likelihood ancestral state reconstructions to map the evolution of spectral shifts within the L photopigments of nymphalids, we estimate the ancestral pigment had a lambda(max) = 540 nm +/- 10 nm standard error and that blueshifts in wavelength have occurred at least 4 times within the family. We used ancestral state reconstructions to investigate the importance of several amino acid substitutions (Ile17Met, Ala64Ser, Asn70Ser, and Ser137Ala) previously shown to have evolved under positive selection that are correlated with blue spectral shifts. These reconstructions suggest that the Ala64Ser substitution has indeed occurred along the newly identified blueshifted L photopigment lineages. Substitutions at the other 3 sites may also be involved in the functional diversification of L photopigments. Our data strongly suggest that there are limits to the evolution of L photopigment spectral shifts among species with only one L opsin gene and that opsin gene duplication broadens the potential range of lambda(max) values.

Adaptive evolution of color vision as seen through the eyes of butterflies

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

Opsin clines in butterflies suggest novel roles for insect photopigments

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

Beyond e-commerce: when caterpillars know what butterflies understand

Many small firms increasingly operate in markets under siege from new entrants who exploit the technologies associated with the Internet's World Wide Web (the web). In these circumstances, interpreting the operating environment is like a vu jade, the opposite of deja vu, a time in space where they have never been, have no idea what they are doing and who it is that could help them. Through the use of the story of the Caterpillar and the Butterfly, this paper considers the inherent difficulties faced by small firms considering the prospect of becoming an e-firm. When considered from an evolutionary perspective, the journey from small firm to small e-firm is not seen as one of choice, but rather one of necessity. In such markets, a race currently appears to exist between entrepreneurs exploiting the web's technologies, and the process of natural selection acting upon firms whose routines have lost favour.

A generic individual-based spatially explicit model as a novel tool for investigating insect-plant interactions: A case study of the behavioural ecology of frugivorous Tephritidae

Computational modelling of mechanisms underlying processes in the real world can be of great value in understanding complex biological behaviours. Uptake in general biology and ecology has been rapid. However, it often requires specific data sets that are overly costly in time and resources to collect. The aim of the current study was to test whether a generic behavioural ecology model constructed using published data could give realistic outputs for individual species. An individual-based model was developed using the Pattern-Oriented Modelling (POM) strategy and protocol, based on behavioural rules associated with insect movement choices. Frugivorous Tephritidae (fruit flies) were chosen because of economic significance in global agriculture and the multiple published data sets available for a range of species. The Queensland fruit fly (Qfly), Bactrocera tryoni, was identified as a suitable individual species for testing. Plant canopies with modified architecture were used to run predictive simulations. A field study was then conducted to validate our model predictions on how plant architecture affects fruit flies’ behaviours. Characteristics of plant architecture such as different shapes, e.g., closed-canopy and vase-shaped, affected fly movement patterns and time spent on host fruit. The number of visits to host fruit also differed between the edge and centre in closed-canopy plants. Compared to plant architecture, host fruit has less contribution to effects on flies’ movement patterns. The results from this model, combined with our field study and published empirical data suggest that placing fly traps in the upper canopy at the edge should work best. Such a modelling approach allows rapid testing of ideas about organismal interactions with environmental substrates in silico rather than in vivo, to generate new perspectives. Using published data provides a saving in time and resources. Adjustments for specific questions can be achieved by refinement of parameters based on targeted experiments.

Phylogenetics and biogeography of a spectacular Old World radiation of butterflies: the subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrini)

Background: Butterflies of the subtribe Mycalesina (Nymphalidae: Satyrinae) are important model organisms in ecology and evolution. This group has radiated spectacularly in the Old World tropics and presents an exciting opportunity to better understand processes of invertebrate rapid radiations. However, the generic-level taxonomy of the subtribe has been in a constant state of flux, and relationships among genera are unknown. There are six currently recognized genera in the group. Mycalesis, Lohora and Nirvanopsis are found in the Oriental region, the first of which is the most speciose genus among mycalesines, and extends into the Australasian region. Hallelesis and Bicyclus are found in mainland Africa, while Heteropsis is primarily Madagascan, with a few species in Africa. We infer the phylogeny of the group with data from three genes (total of 3139 bp) and use these data to reconstruct events in the biogeographic history of the group.,Results: The results indicate that the group Mycalesina radiated rapidly around the Oligocene-Miocene boundary. Basal relationships are unresolved, but we recover six well-supported clades. Some species of Mycalesis are nested within a primarily Madagascan clade of Heteropsis,while Nirvanopsis is nested within Lohora. The phylogeny suggests that the group had its origin either in Asia or Africa, and diversified through dispersals between the two regions, during the late Oligocene and early Miocene. The current dataset tentatively suggests that the Madagascan fauna comprises two independent radiations. The Australasian radiation shares a common ancestor derived from Asia. We discuss factors that are likely to have played a key role in the diversification of the group. Conclusions: We propose a significantly revised classification scheme for Mycalesina. We conclude that the group originated and radiated from an ancestor that was found either in Asia or Africa, with dispersals between the two regions and to Australasia. Our phylogeny paves the way for further comparative studies on this group that will help us understand the processes underlying diversification in rapid radiations of invertebrates.

A meta-analysis of the traits affecting dispersal ability in butterflies: can wingspan be used as a proxy?

1. Dispersal ability of a species is a key ecological characteristic, affecting a range of processes from adaptation, community dynamics and genetic structure, to distribution and range size. It is determined by both intrinsic species traits and extrinsic landscape-related properties. 2. Using butterflies as a model system, the following questions were addressed: (i) given similar extrinsic factors, which intrinsic species trait(s) explain dispersal ability? (ii) can one of these traits be used as a proxy for dispersal ability? (iii) the effect of interactions between the traits, and phylogenetic relatedness, on dispersal ability. 3. Four data sets, using different measures of dispersal, were compiled from published literature. The first data set uses mean dispersal distances from capture-mark-recapture studies, and the other three use mobility indices. Data for six traits that can potentially affect dispersal ability were collected: wingspan, larval host plant specificity, adult habitat specificity, mate location strategy, voltinism and flight period duration. Each data set was subjected to both unifactorial, and multifactorial, phylogenetically controlled analyses. 4. Among the factors considered, wingspan was the most important determinant of dispersal ability, although the predictive powers of regression models were low. Voltinism and flight period duration also affect dispersal ability, especially in case of temperate species. Interactions between the factors did not affect dispersal ability, and phylogenetic relatedness was significant in one data set. 5. While using wingspan as the only proxy for dispersal ability maybe problematic, it is usually the only easily accessible species-specific trait for a large number of species. It can thus be a satisfactory proxy when carefully interpreted, especially for analyses involving many species from all across the world.

Does size matter? Comparative population genetics of two butterflies with different wingspans

The dispersal ability of a species is central to its biology, affecting other processes like local adaptation, population and community dynamics, and genetic structure. Among the intrinsic, species-specific factors that affect dispersal ability in butterflies, wingspan was recently shown to explain a high amount of variance in dispersal ability. In this study, a comparative approach was adopted to test whether a difference in wingspan translates into a difference in population genetic structure. Two closely related butterfly species from subfamily Satyrinae, family Nymphalidae, which are similar with respect to all traits that affect dispersal ability except for wingspan, were studied. Melanitis leda (wingspan 60-80 mm) and Ypthima baldus (wingspan 30-40 mm) were collected from the same areas along the Western Ghats of southern India. Amplified fragment length polymorphisms were used to test whether the species with a higher wingspan (M. leda) exhibited a more homogenous population genetic structure, as compared to a species with a shorter wingspan (Y. baldus). In all analyses, Y. baldus exhibited greater degree of population genetic structuring. This study is one of the few adopting a comparative approach to establish the relationship between traits that affect dispersal ability and population genetic structure.