Enrique García-Barros

ABSENCE OF TRADE-OFFS BETWEEN SEXUAL SIZE DIMORPHISM AND EARLY MALE EMERGENCE IN A BUTTERFLY'

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. If protandry 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 devel- opment, 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 environ- ments. Similar female-biased size dimorphism occurred in protandrous and non-protan- drous 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 phe- nomenon and may make possible independent optimization of size and development time (age at sexual maturity), and accordingly influence expected patterns of size-related trade- offs.

DNA barcode reference library for Iberian butterflies enables a continental-scale preview of potential cryptic diversity

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.

Use of space and resources in a Mediterranean population of the butterfly Euphydryas aurinia

Abstract Population parameters of a central Spanish population of the marsh fritillary butterfly, Euphydryas aurinia (Rott.), were studied between 1990 and 1991. Numbers were very similar in the two study years, but adult emergence was delayed 15 days in the second year as a consequence of variation in weather conditions. The nectar sources used by adults were different in the two years and were correlated with the most abundant plants available during the flight period. Adult abundance was correlated with the presence of the larval foodplant (Lonicera periclymenum) in open woodland and was higher where this habitat was mixed with open areas, where nectar sources abound. Estimates of male population numbers were higher than those of females, suggesting that males used wider ranges than females, or a higher residence rate for males. Nevertheless mobility did not differ significantly between the sexes. The conservation of the species in central Spain depends on maintaining patches of oak woodlands, important for the presence of larval foodplants. Large numbers of the butterfly in the Natural Park study area are favoured by traditional management including charcoal production and extensive grazing by cattle and goats.

Immature stages of Hipparchia Fabricius and the systematics of the ‘Satyrus series» (Lepidoptera: Nymphalidae: Satyrinae)

Abstract. Characters of potential use from the eggs and larvae of four species of Hipparchia (subgenera Hipparchia, Parahipparchia, Neohipparchia and Pseudotergumia) are described. These characters, together with previously published information on larval, pupal and adult morphology, are used to construct a seventy‐one character matrix for Hipparchia and four other genera of the ‘Satyrus series’ (Miller, 1968): Arethusana, Kanetisa, Chazara and Satyrus. The cladistic relationships among these genera and the relative contribution of adult and immature data sets are discussed. A list of possible synapomorphies of the immature stages of the ‘Satyrus series’ is proposed.

Morphology and chaetotaxy of the first instar larvae of six species of the Satyrus (s.l.) series (Lepidoptera: Nymphalidae)

Abstract. The morphology and chaetotaxy of the first instar larvae of six species belonging to the genera Hipparchia, Kanetisa and Chazara are described. Specific characters are stated, drawn mainly from size, setal length and morphology, and the shape of the suranal plate. Several characters, other than chaetotaxy, that are of potential use in nymphalid systematics are discussed. The larval chaetotaxy is briefly compared with that of both heliconiine and danaine first instar larvae.

Morphology and morphometry of Lycaenid eggs (Lepidoptera: Lycaenidae)

A morphological study with the use of scanning electron microscope of 67 species of Iberian Lycaenidae is presented. The study covers all the genera present in the area and shows an extraordinary variation in chorionic characters that allows egg diagnosis for most species. A morphometric study showed that the eggs from the sample have sizes that are correlated with adult size, but some species showed larger egg size than expected. Species hibernating at the egg stage proved to have on average larger sizes than those overwintering at other stages, probably because this trait might be favourable to endure the adverse conditions taking place during the winter. A cladistic analysis was performed using morphologic and morphometric characters from the egg with the result of poor discriminant power. However, some formal taxonomic groups such as the genera Lycaena and Satyrium were supported by our analysis due to specific apomorphic characters.

Ecology and recovery plans for the four Spanish endangered endemic butterfly species

Data compiled during three fieldwork seasons and habitat models provided the tools to produce Species recovery plans for the endangered and endemic butterflies in Spain: Polyommatus violetae, P. golgus, Agriades zullichi, and Euchloe bazae. The distribution of each species has been updated with new field records. For E. bazae and P. violetae the known distribution records increased 33% and 8%, respectively. Detailed habitat descriptions, based on fieldwork data, are given for the four species and models show relatively small and fragmented habitat patches for P. violetae, P. golgus, and P. zullichi. Those areas might be particularly sensitive under a climate change scenario. Habitat of the four species is very specialized and mainly consists of open areas with grassland or scrub vegetation. Population densities are provided for the first time for these species and show low numbers for E. bazae, particularly in the area around Baza in SE Spain. 25 to 54% of the populations were apparently not threatened. For the rest, apparent habitat suitability threats were abandonment for E. bazae and P. violetae, trampling for A. zullichi, and overgrazing for P. golgus. Recovery plans included aspects on legislation, habitat management, precautionary measures (limit new developments and land use changes), research, and public awareness campaigns. Suggested habitat management actions are: (1) support extensive grazing and prevent overgrazing; (2) reduce the effect of trampling by visitors in mountain areas; and (3) woodland and scrub clearing of abandoned areas and pine plantations to improve the quality of grassland habitats.

Phylogeny of the genus Palmanura (Collembola: Neanuridae)

In order to assess the phylogenetic structure of the springtail genus Palmanura, as well as to test the monophyly of the tribe Sensillanurini (Neanuridae: Neanurinae), a data matrix of morphological (chaetotactic and other) characters of members of this group was assembled and analysed in the light of Wagner parsimony. The data matrix included all the known members of the Neotropical genus Palmanura, plus representatives of Sensillanura and Americanura. Although not all the clades obtained were highly supported by bootstrap resampling, some structures were relatively constant under different approaches. Alternative analyses (unordered and ordered character states, rescaled weighting procedure) were applied. While alternative solutions were obtained, a number of structures were shared by the results irrespective of the method used. On this basis, the results suggest that some further reassessment is required to confirm formally the monophyly of the tribe Sensillanurini. The genera Palmanura and Americanura are mutually poly/paraphyletic; we thus suggest that Palmanura should be considered as a synonym of Americanura, although some character reassessment and more varied outgroup species may be necessary before a formal generic redefinition can be proposed. Finally, a comparison of the performance of the characters under Wagner parsimony analysis indicated that differences in the characters’ retention indexes are due not to the topological (tagmal) position of the traits involved, but to character coding: the characters describing quantitative features (generally numbers of setae) generally performed worse than other types of characters under parsimony. An updated list of the known members of the Sensillanurini (Collembola: Neanuridae: Neanurinae) is presented.
© The Willi Hennig Society 2009.

Identification of potential source and sink areas for butterflies on the Iberian Peninsula

The dynamic analysis of geographical distribution is relevant to identify the processes that underlie any changes of geographical ranges. This is an essential element of both biogeography and conservation biology. Fuzzy logic, in particular the fuzzy concept of favourability for species occurrence, helps to perform a dynamic interpretation of the internal complexity of species ranges. We modelled the distributions of the 222 Iberian butterflies using favourability functions and 92 environmental variables (spatial, climatic, topographic, geological and indicators of human activity). We obtained a significant environmental favourability model for each butterfly. We identified the potential sources and sinks in the distribution area of each butterfly species using their respective favourability and presence–absence maps, considering as sources only those areas with high favourability where the butterfly is present, and sinks only those areas with low favourability where the butterfly is present too. The source areas for the Iberian butterflies are concentrated in the north of the peninsula, mainly in the mountain ranges (Cantabrian Range, Pyrenees and Central Range). Sink areas are more dispersed all around the peninsula. We found a concentration of sink locations in eastern Iberia (Alicante and Murcia provinces) and in the southwestern area (Doñana National Park). This may be helpful when implementing conservation measures, by providing a biogeographical dynamic interpretation of the roles of different parts of the distribution range of the species. Preserving the processes that link sources with sinks is critical for maintaining or improving the sustainability of populations.

Corrigendum: DNA barcodes highlight unique research models in European butterflies

V. Dincă and P.D.N. Hebert. Biodiversity Institute of Ontario, University of Guelph, 50 Stone Road East, ON N1G 2W1, Canada. N. Backstrom. Department of Evolutionary Biology, Uppsala University, Norbyvagen 18D, 75236 Uppsala, Sweden. L. Dapporto. Department of Biological and Medical Sciences, Oxford Brookes University, Headington, Oxford, OX3 0BP, UK. M. Friberg. Department of Ecology and Genetics, Plant Ecology and Evolution, Uppsala University, Norbyvagen 18D, 75236 Uppsala, Sweden. E. Garcia-Barros, J. Hernandez-Roldan, and M.L. Munguira. Department of Biology, Universidad Autonoma de Madrid, Campus Cantoblanco 28049, Madrid, Spain. E. Hornett. Department of Zoology, University of Cambridge, Cambridge, CB2 3EJ, UK. V. Lukhtanov. Department of Karyosystematics, Zoological Institute of Russian Academy of Sciences, Universitetskaya nab. 1, 199034 St. Petersburg, Russia. F. Marec and J. Sichova Faculty of Science, University of South Bohemia, Ceske Budějovice, 370 05, Czech Republic. S. Montagud. Institut Cavanilles de Biodiversitat i Biologia Evolutiva (ICBiBE) Universitat de Valencia, Carrer Catedratic Jose Beltran 2, 46980, Paterna, Spain. M. Olofsson and C. Wiklund. Department of Zoology, Stockholm University, Svante Arrhenius vag 18B, S-106 91, Stockholm, Sweden. G. Talavera. Department of Organismic and Evolutionary Biology, Museum of Comparative Zoology, Harvard University, 26 Oxford St., 02138 Cambridge, MA, USA. J.C. Vicente-Arranz. Asociacion Espanola para la Proteccion de las Mariposas y su Medio (Zerynthia), Logrono (La Rioja), Spain. R. Vila. Institut de Biologia Evolutiva (CSIC-Universitat Pompeu Fabra), Passeig Maritim de la Barceloneta 37, 08003, Barcelona, Spain. Corresponding author: Vlad Dincă (e-mail: vdinca@uoguelph.ca). 391