Protandry, here defined as the earlier emergence of males, is a common feature in life histories and could be the result of sexual selection on males to maximize matings, or alternatively an incidental by-product of other selection pressures on the sexes. lfprotandry is selected for per se, theory predicts that it should be associated with seasonal environments where there is little overlap between generations. The degree of protandry should be insensitive to environmental conditions. Moreover, on the assumption that males and females grow at the same rate as larvae, a trade-off between development time and size is expected to result in a strong association between protandry and female-biased sexual size dimorphism. These predictions were tested by a combination of comparative and experimental studies on five populations of the speckled wood butterfly, Pararge aegeria, from central and south Sweden, England, Spain, and the island of Madeira. Protandry was associated with seasonal environments, as it was only exhibited in the three northernmost populations. Protandry in these populations remained largely constant in a variety of temperatures, both under direct development, when protandry results from a sex difference in development time through the egg, larval, and pupal stages, and under diapause development, when it results from a sex difference in pupal development time only. These results indicate that protandry is selected for per se through sexual selection in seasonal environments. Similar female-biased size dimorphism occurred in protandrous and non-protandrous populations alike, and hence sexual size dimorphism in P. aegeria is not a result of selection for protandry, nor the causal factor behind protandry. Protandry and sexual size dimorphism appear to be largely decoupled traits in the life history evolution of P. aegeria. This is achieved by means of variation in pupal developmental time and variation in the relative growth rates of the sexes. Variation in growth rates is likely to be a general phenomenon and may make possible independent optimization of size and development time (age at sexual maturity), and accordingly influence expected patterns of size-related tradeoff's.
How common are cryptic species - those overlooked because of their morphological similarity? Despite its wide-ranging implications for biology and conservation, the answer remains open to debate. Butterflies constitute the best-studied invertebrates, playing a similar role as birds do in providing models for vertebrate biology. An accurate assessment of cryptic diversity in this emblematic group requires meticulous case-by-case assessments, but a preview to highlight cases of particular interest will help to direct future studies. We present a survey of mitochondrial genetic diversity for the butterfly fauna of the Iberian Peninsula with unprecedented resolution (3502 DNA barcodes for all 228 species), creating a reliable system for DNA-based identification and for the detection of overlooked diversity. After compiling available data for European butterflies (5782 sequences, 299 species), we applied the Generalized Mixed Yule-Coalescent model to explore potential cryptic diversity at a continental scale. The results indicate that 27.7% of these species include from two to four evolutionary significant units (ESUs), suggesting that cryptic biodiversity may be higher than expected for one of the best-studied invertebrate groups and regions. The ESUs represent important units for conservation, models for studies of evolutionary and speciation processes, and sentinels for future research to unveil hidden diversity.
Aim The geographical distributions of animal and plant species endemic to the Iberian Peninsula and Balearic Islands were analysed to locate and designate areas of endemicity.Location The Iberian Peninsula and the three largest Balearic Islands (Mallorca, Menorca and Ibiza) in the western Mediterranean, West Palaearctic region. MethodsThe information analysed consisted of presence/absence data of animal and plant species, recorded on a 100´100 km grid based on the UTM projection system. From a larger initial data set, a simpli®ed matrix of 480 species present in at least two quadrats was obtained, and processed to estimate the overall similarity patterns across land squares, and the areas of endemism. Two methods were employed to detect areas of endemism: Wagner Parsimony (PAE, or parsimony analysis of endemicity) and compatibility. A modi®cation of PAE, PAE±PCE (Parsimony analysis of endemicity with progressive character elimination) was applied to overcome some of the potential shortcomings of the method. ResultsThe results represent the ®rst attempt for a combined analysis of animal and plant distributions in the western Mediterranean. The proposed PAE±PCE procedure proved useful to identify areas of endemism that would have been otherwise overlooked. Up to thirty-six different areas of endemisms were identi®ed. Some of these represent concentric (hierarchically nested) structures, while other are partly overlapping sectors. The endemism areas, as derived from parsimony and compatibility analyses, generally ®t within the frame of the overall similarity approach.Main conclusions The areas of endemicity identi®ed often coincide with mountain sectors, and this may be of incidental interest for conservation policies as most natural preserves in the study area are located in mountain ranges. The conclusions are of interest for large scale approaches to the biogeography of the Mediterranean Basin, facilitating the selection of endemism areas for operative purposes. However, most of the best supported areas of endemism detected are relatively small, or overlap with neighbouring endemism areas. Hence, adopting large area units such as`Iberia' for historical analysis at a wider geographical scale may be risky, because such units may actually represent composite sectors of an heterogeneous nature. The distribution of the areas of endemism, as well as the results of the overall similarity classi®cation, share a number of features with previous sectorizations from independent, mostly phytogeographical, approaches. Parsimony analysis of endemicity is a potentially useful tool for identifying areas designated by species with congruent distributions, but (1) the results Correspondence: Enrique Garcõ Âa-Barros, have no direct historical implications (for phylogenetic information is not incorporated), and (2) unless modi®cations such as the PAE±PCE procedure are applied, the number of potential areas of endemism (in the sense stated above) will often be underestimated. It is also shown that, in a PAE, a`total evidence' approac...
The interspecific relationships between egg size and body size in buttedies (Papilionoidea and Hesperiidae), and between size and egg and larval development time, larval trophic specificity, foodplant structure, climate, and phenology were investigated based on a sample of more than 1180 species. The independent contrasts method was used to avoid taxonomydependent results. Egg size is allometrically related to adult wing length by a slope of 0.43. Based on a subset of species, fecundity is correlated to adult body size, and there is evidence for a compromise between egg number and egg size (relative to adult size) across species. Butterfly size increases in correlation to the mean annual temperature of the species geographic range, but decreases in relation to increased aridity (or the length of the dry season). Larger butterflies tend to have longer larval development times, use large or structurally complex host plants, and are more likely to lay their eggs in batches, irrespective of climate. Larger eggs tend to develop more slowly, and give rise to larvae with longer developmental periods that will result in larger adults. No evidence was found to support a relationship between butterfly body size and polyphagy. A complex pattern of interrelationships links body size (and egg size) to other traits, although correlations other than that between egg size and body size are generally low. The results suggest the necessity of separating climate and seasonality into components that are relevant to insect life histories in comparative studies.
A three‐step protocol described elsewhere is used to obtain a map of butterfly species density in Portugal on a 50×50 km grid. First, all available faunistic information was compiled and analysed to explore the historic patterns of butterfly sampling in Portugal, and to determine which grid cells are sufficiently prospected to produce reliable estimates of species richness. Then, we relate the estimated species richness scores from these areas to a set of environmental and spatial variables by means of General Linear Models, obtaining a function to extrapolate of species density scores to the rest of Portugal. Finally, the model is validated, results explored and outliers identified and deleted. Any spatial autocorrelation remaining in the residuals is examined. Lastly, model parameters are recalculated in absence of deleted outliers, and the resulting function is used to predict species richness scores throughout mainland Portugal. A highly‐predictive function based on some variables previously related to butterfly composition at macro‐scale, such as number of sunny days per year, temperature or environmental heterogeneity, was obtained. However, in Portugal those variables are highly spatially structured along a steep latitudinal gradient, leading to difficulty in ascertaining if the latitudinal gradient detected by our analysis is due to macroecological or historic effects. Information on European and Iberian butterfly assemblages and causal processes are discussed in the light of the patterns observed. Then, previous information obtained on Portuguese scarabs is added to identify conservation areas, biogeographically important for both insect groups. Finally, the main drawbacks and advantages of this approach to mapping biodiversity for conservation are discussed briefly.
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