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Taxa restricted to mountains may be vulnerable to global warming, unless localscale topographic variation and conservation actions can protect them against expected changes to the climate. We tested how climate change will affect the 19 mountain-restricted Erebia species of the Iberian Peninsula, of which 7 are endemic. To examine the scope for local topographic variation to protect against warming, we applied species distribution models (HadGEM2 and MPI) at two spatial scales (10 10 and 1 1 km) for two representative concentration pathways (RCP4.5 and RCP8.5) in 2050 and 2070. We also superimposed current and future ranges on the protected area (PA) network to identify priority areas for adapting Erebia conservation to climate change. 4. In 10 10 km HadGEM2 models, climatically suitable areas for all species decreased in 2050 and 2070 (average 95.7%). Modelled decreases at 1 1 km were marginally less drastic (95.3%), and 14 out of 19 species were still expected to lose their entire climatically favourable range by 2070. The PA network is well located to conserve the species that are expected to retain some climatically suitable areas in 2070. However, we identify 25 separate 10 10 km squares where new PAs would help to adapt the network to expected range shifts or contractions by Erebia. Based on our results, adapting the conservation of range-restricted mountain taxa to projected climate change will require the implementation of complementary in situ and ex situ measures alongside urgent climate change mitigation., This study was supported by the project BIOUAM07-2019 founded by Biology Department from Universidad Autonoma de Madrid, the project IBERARTRO (SBPLY/17/180501/000492) funded by European Regional Development Fund (ERDF) through the Consejería de Educacion, Ciencia y Cultura, Junta de Comunidades de Castilla-La Mancha, the project RTI2018-096739-B-C21 funded by MCIU/AEI/ FEDER, UE, and the NexTdive project (PID2021-124187NB-I00) funded by the Spanish Ministry of Science and Innovation.

Reproductive character displacement occurs when competition for successful breeding imposes a divergent selection on the interacting species, causing a divergence of reproductive traits. Here, we show that a disputed butterfly taxon is actually a case of male wing colour shift, apparently produced by reproductive character displacement. Using double digest restriction-site associated DNA sequencing and mitochondrial DNA sequencing we studied four butterfly taxa of the subgenus Cupido (Lepidoptera: Lycaenidae): Cupido minimus and the taxon carswelli, both characterized by brown males and females, plus C. lorquinii and C. osiris, both with blue males and brown females. Unexpectedly, taxa carswelli and C. lorquinii were close to indistinguishable based on our genomic and mitochondrial data, despite displaying strikingly different male coloration. In addition, we report and analysed a brown male within the C. lorquinii range, which demonstrates that the brown morph occurs at very low frequency in C. lorquinii. Such evidence strongly suggests that carswelli is conspecific with C. lorquinii and represents populations with a fixed male brown colour morph. Considering that these brown populations occur in sympatry with or very close to the blue C. osiris, and that the blue C. lorquinii populations never do, we propose that the taxon carswelli could have lost the blue colour due to reproductive character displacement with C. osiris. Since male colour is important for conspecific recognition during courtship, we hypothesize that the observed colour shift may eventually trigger incipient speciation between blue and brown populations. Male colour seems to be an evolutionarily labile character in the Polyommatinae, and the mechanism described here might be at work in the wide diversification of this subfamily of butterflies., European Regional Development Fund. Grant Numbers: CGL2016‐76322‐P, PID2019‐107078GB‐I00 Ministerio de Economía, Industria y Competitividad, Gobierno de España. Grant Number: BES‐2017‐080641 Ministerio de Ciencia, Innovación y Universidades Generalitat de Catalunya. Grant Number: 2017‐SGR‐991 Academy of Finland. Grant Number: 328895

Evaluating data quality and inventory completeness must be a preliminary step inany biodiversity research, particularly in the case of insects and high biodiversityareas. Yet, this step is often neglected or, at best, assessed only for one insectgroup, and the degree of congruence of sampling effort ffor different insect groupsremains unexplored. We assess the congruence in the spatial distribution of sampling effort for fiveinsect groups (butterflies, caddisflies, dung beetles, moths, and aquatic beetles) inthe Iberian Peninsula. We identify well-surveyed areas for each taxonomic groupand examine the degree to which the patterns of sampling effort can be explainedby a set of variables related to environmental conditions and accessibility. Irrespective of the general lack of reliable inventories, we found a general but lowcongruence in the completeness patterns of the different taxa. This suggests thatthere is not a common geographical pattern in survey effort and that idiosyncraticand contingent factors (mainly the proximity to the workplaces of entomologists)are differentially affecting each group. After many decades of taxonomic and faunistic work, distributional databases ofIberian insects are still in a very preliminary stage, thus limiting our capacity toobtain reliable answers to basic and applied questions. We recommend carrying out long-term, standardised and well-designed entomolog-ical surveys able to generate a reliable image of the distribution of different insect groups. This will allow us to estimate accurately insect trends and better under-stand the full extent of global biodiversity loss., This study has been supported by the projects BioWeb (MINECO:CGL2011-15622-E BOS), BANDENCO (JCMM: POII11-0277-5747)and IBERARTRO (SBPLY/17/180501/000492) founded by European Regional Development Fund (ERDF) through the Consejería de Educación, Ciencia y Cultura, Junta de Comunidades de Castilla-La Mancha. David Sánchez-Fernández is funded by a postdoctoral contract fromthe Spanish Ministry of Science and Innovation (Ramón y Cajal program; RYC2019-027446-I

We estimated the potential impact of Global Warming on the species richness of Iberian butterflies. First, we determined the grid size that maximized the balance between geographic resolution, area coverage and environmental representativeness. Contemporary richness was modelled in several alternative ways that differed in how sampling effort was controlled for, and in whether the non-climatic variables (physiography, lithology, position) were incorporated. The results were extrapolated to four WorldClim scenarios. Richness loss is to be expected for at least 70% of the area, with forecasts from the combined models being only slightly more optimistic than those from the purely climatic ones. Overall, the most intense losses are predicted for areas of highest contemporary species richness, while the potential slightly positive or nearly neutral changes would most often concentrate in cells of low to moderate present richness. The environmental determinants of richness might not be uniform across the geographical range of sampling effort, suggesting the need of additional data from the least intensively surveyed areas., Open Access funding provided thanks to the CRUE-CSIC agreement with Springer Nature. This study was partly funded by JJ CC Castilla—La Mancha (Spain), Project SBPLY/17/180501/000492 and MINECO, Project CGL2017-86926-P

ContextUrbanisation is an environmental filter for many species that leads to community homogenisation, with a few species inhabiting isolated patches (e.g. public and private gardens and parks) embedded within the urban landscape. Promoting biodiversity in urban areas requires understanding which species traits allow species to survive the urban landscape.ObjectivesThe objective of this study was to assess how species traits and landscape factors combine to allow species functional groups to live in the city.MethodsWe used butterfly count data collected by volunteers in 24 gardens of Barcelona city, during 2018 and 2019. Species were clustered in functional groups according to their traits. We applied a multinomial choice model to test for the effect of the landscape on the different functional groups.ResultsThree functional groups became prevalent in the city while a fourth, containing most sedentary specialist species, was filtered out. Although the observed groups had similar species richness, abundances varied depending on urban landscape characteristics. Specialist sedentary specialists and medium mobile species were all favoured by patch connectivity; while the presence of mobile generalist species was only enhanced by habitat quality. Our results indicate that butterfly communities are more diverse in highly connected gardens.ConclusionsOur study highlights the need of contextualised management with actions accounting for the species functional groups, rather than a management focused on general species richness. It demonstrates that urban landscape planning must focus on improving connectivity inside the city in order to diversify the community composition.

1. Indices of environmental associations such as the Community Temperature Index (CTI) and Community Precipitation Index (CPI) can be derived from occurrence data to extend the geographic scope or time frame of evidence for responses of insect diversity to global change., 2. We tested whether occurrence records from 1901 to 2016 from the Iberian Peninsula could shed light on butterfly community responses to changes over space and time in the climate; and whether local climatic variation caused by topographic heterogeneity could buffer communities against the effects of climate change., 3. CTI and CPI were closely related to variation in temperature and precipitation across 115 well-sampled 10 km grid squares. However, whereas temperature and precipitation changed systematically from 1901–1979 to 1980–2016, and these changes were positively related to changes in CTI and CPI, community climatic associations did not change significantly over time. Butterfly communities became more associated with closed vegetation, suggesting that land cover changes overshadowed the effects of climate change., 4. Local (1 km) climatic variation generally exceeded change over time at 10 km resolution, and heterogeneity in elevation slowed rates of warming. In turn, spatial variation in climatic conditions dampened butterfly community responses to heating and drying., 5. Occurrence data are limited by their spatial resolution but can inform understanding of insect community responses to global change for regions lacking long-term monitoring data. Our results suggest that local climatic variation accompanying topographic heterogeneity can shield regional butterfly faunas from the impacts of climate change., Analyses were funded by project RTI2018-096739-B-C21 (MCIU/AEI/FEDER,UE) to R​.W. Atlas data were developed by M.L.M., E.G.B. and H.R. using funding from I + D + I National Plan project DynaGen (CGL2013-48277-P) and regional projects BANDENCO (POII-2014-018-P) and IBERARTHRO (SBPLY/17/180501/000492). Atlas data verification and project development were supported by Comunidad de Madrid funding to M.L.M. (PEJD-2017/PRE/AMB-4719) and R.W. (2016-T3/AMB-1073).

Trait-based analyses explaining the different responses of species and communities to environmental changes are increasing in frequency. European butterflies are an indicator group that responds rapidly to environmental changes with extensive citizen science contributions to documenting changes of abundance and distribution. Species traits have been used to explain long- and short-term responses to climate, land-use and vegetation changes. Studies are often characterised by limited trait sets being used, with risks that the relative roles of different traits are not fully explored. Butterfly trait information is dispersed amongst various sources and descriptions sometimes differ between sources. We have therefore drawn together multiple information sets to provide a comprehensive trait database covering 542 taxa and 25 traits described by 217 variables and sub-states of the butterflies of Europe and Maghreb (northwest Africa) which should serve for improved trait-based ecological, conservation-related, phylogeographic and evolutionary studies of this group of insects. We provide this data in two forms; the basic data and as processed continuous and multinomial data, to enhance its potential usage., Measurement(s) resources • Egg Laying • larval environment • pupal environment • geographic location • behavior • size • voltinism • phenology • host plant Technology Type(s) digital curation Factor Type(s) species Sample Characteristic - Organism Papilionoidea Sample Characteristic - Location Europe • Northwest Africa Machine-accessible metadata file describing the reported data: 10.6084/m9.figshare.12998828

"This is the peer reviewed version of the following article: Pulido-Pastor, Antonio et al. "Identification of potential source and sink areas for butterflies on the Iberian Peninsula", which has been published in final form at https://doi.org/10.1111/icad.12297. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions.", 1. 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. 2. 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. 3. 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. 4. 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). 5. 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, Parts of this work were supported by projects POII11-0277-5747/BANDENCO, CGL2017-86926-P and sDIV (iDIV)WG 4.28 sECURE

The climatic preferences of the species determine to a large extent their response to climate change. Temperature preferences have been shown to play a key role in driving trends in animal populations. However, the relative importance of temperature and precipitation preferences is still poorly understood, particularly in systems where ecological processes are strongly constrained by the amount and timing of rainfall. In this study, we estimated the role played by temperature and precipitation preferences in determining population trends for birds and butterflies in a Mediterranean area. Trends were derived from long-term biodiversity monitoring data and temperature and precipitation preferences were estimated from species distribution data at three different geographical scales. We show that population trends were first and foremost related to precipitation preferences both in birds and in butterflies. Temperature preferences had a weaker effect on population trends, and were significant only in birds. The effect of precipitation on population trends operated in opposite directions in the two groups of species: butterfly species from arid environments and bird species from humid habitats are decreasing most. Our results indicate that, although commonly neglected, water availability is likely an important driver of animal population change in the Mediterranean region, with highly contrasting impacts among taxonomical groups.

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., We thank MAVA Fondation pour la Nature for financial support (Project No. 11/198).

Butterfly life-history features are expected to co-vary along environmental gradients related to changes in the vegetation structure or composition; however the direction and intensity should vary across regions at the large scale. This study focuses on the butterfly communities of Portuguese ‘montados’. Sixteen sites (mostly cork oak fields) were selected, reflecting a succession gradient in the vegetation of the understorey after human intervention. While controlling for vegetation and broader geographical and climate effects, we looked for trends in butterfly species richness and abundance (using generalised linear models) and for trends in species composition (using redundancy analyses). Moreover, we tried to uncover the co-variation between the butterfly life-history characteristics and succession. The results revealed that butterfly species richness was not significantly influenced by any of the considered variables. In contrast, abundance depended on geographic and oceanity–continentality gradients as it increased towards the East and with more marked temperature annual ranges and less dry summer conditions. Species composition was influenced by temperature ranges and by shrub coverage. There was no strong evidence in favour of fast–slow or generalist–specialist syndromes co-varying along human imposed environmental gradients. However, after controlling for the broad scale variables (geography and climate) shrub cover emerged as a relevant factor. This reinforces the idea that late successional stages are not optimal for butterfly communities. It implies the importance of the extensive methods of traditional management and the negative effects of long-term abandonment.

The Iberian Peninsula is a major European region of biodiversity, as it harbours more than 30% of European endemic species. Despite a number of studies having evaluated the ability of nature reserves to protect certain taxa, there is still a lack of knowledge on how Iberian endemic fauna are represented in these reserves. We detected biodiversity hotspots of Iberian endemicity and evaluated the effectiveness of the Natura 2000 network (N2000) in representing 249 endemic species from eight animal taxonomic groups (amphibians, mammals, freshwater fishes, reptiles, water beetles, butterflies, lacewings and dung beetles). We found that only the 10% of these Iberian endemic species are considered species of community interest (i.e. species included in the Annexes of the Habitats Directive). We conducted gap analyses and null models of representativeness in N2000. Generally, N2000 is effective in its representation of Iberian endemic fauna, although we detected species and few hotspots of endemism that were still not represented. It is necessary to declare a few new protected areas, thus enhancing N2000's effectiveness in the conservation of the Iberian endemic fauna. Although the aim of N2000 is to protect species listed in the Birds and Habitats Directives, the conservation status of endemic species from one of the most important areas of Europe in terms of biodiversity, could be also a concern for the European Union. Our results are useful in the context of the recent European Commission mandate calling for a ‘fitness check’ of the Birds and Habitats Directives. This approach could be also applicable to other regions with high value of endemicity., DS‐F was supported by a post‐doctoral contract funded by Universidad de Castilla‐La Mancha and the European Social Fund (ESF). PA was supported by a ‘Ramón y Cajal’ contract (RYC‐2011‐07670, MINECO). This research was partially funded by project POII11‐0277‐5747 (Junta de Castilla‐La Mancha).

SummaryWe test the extent to which the combination of 69 morphological characters from the early stages (18 from the egg, six larval and eight pupal) with 37 adult morphological characters of the lycaenids (Lepidoptera: Lycaenidae) improves the results of a parsimony analysis to reconstruct the evolutionary history of the group, using 51 sample species from the West Palaearctic. The results show three main clades, although with low support from bootstrap and jackknife resamplings; they represent the four tribes present in our sample, i.e. Lycaenini, Theclini, Eumaeini and Polyommatini. The tribe Lycaenini (the one that has the highest node support) is in the base of the tree. Theclini + Eumaeini appear as sister clades with the Polyommatini as the sister taxon to both of them. At the lowest taxonomic levels our results are often in agreement with those from recent studies done on a molecular basis. Adding morphological information from the immature stages to that of the adult contributed significantly to ...

Measuring threat and selecting ‘flagship’ species for conservation planning should ideally rely on ‘dynamic’ criteria (i.e.: decreasing range and population sizes) which may not be available for most taxa and geographic regions. We address the question of what features of the organisms or of their geographic distributions lie behind selections of potentially threatened species when there is insufficient information on temporal trends. Focusing on Iberian butterflies, we evaluate the support for past and present lists of target species from the most easily quantifiable features of the species’ ranges and conspicuousness (size and visual apparency). Among the features tested, geographic rarity has the highest weight in all the species lists. However, the explanation is rather modest statistically, especially when the species selection is of a supra-regional nature (in this case European). Among the rarest species, those with geographic ranges concentrated in the study area (a) are comparatively few and (b) on average, have less restricted ranges than those whose geographic rarity results from their marginal occurrence in the study area. Finally, the total geographic coverage of the EU-threatened butterflies is relatively large in Iberia. As a consequence species-oriented conservation planning in this region may be rather unrealistic if only geographic rarity is emphasized, or if the levels of threat determined at the supra-regional (European) level are directly translated to the regional legislations. Thus we support the regionally hierarchical approach to conservation proposed by former authors.

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.

Este manual es el resultado de la revision de los guiones que a lo largo de ya numerosos cursos hemos usado como apoyo en las practicas de las asignaturas de Zoologia General a lo largo de cuatro planes de estudio (Licenciatura primero, luego Grado) en la Universidad Autonoma de Madrid. Algunos de ellos tienen su origen en versiones procedentes de otras universidades, adaptados para su uso en el, entonces, tercer curso de Ciencias Biologicas de nuestro centro, hace ya treinta anos. Otros han surgido de esquemas mas personales. En uno y otro caso, han sufrido sucesivas transformaciones a lo largo del tiempo. El resultado cubre las necesidades del curso de manera mas especifica que otros manuales disponibles (HICKMAN & HICKMAN, 1991; MUNILLA LEON, 1992; o MILLER, 1995) al apoyarse en el aprovechamiento de materiales asequibles en nuestro entorno geografico, el area ibero-balear. DOI: http://dx.doi.org/10.15366/pract.zoologia2016

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., Support for this research was provided by the Spanish Ministerio de Economía y Competitividad (project CGL2010-21226/BOS and CGL2013- 48277-P), by Marie Curie International Outgoing Fellowships within the 7th European Community Framework Programme to V. Dincă (project no. 625997) and G. Talavera (project no. 622716), by grant 1.50.1617.2013 at Saint Petersburg State University for G. Talavera, and by the government of Canada through Genome Canada and the Ontario Genomics Institute in support of the International Barcode of Life project.

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.

A database with comprehensive butterfly faunistic information from the Iberian Peninsula and the Balearic Islands was used to estimate inventory completeness as well as the environmental, spatial. and land-use effects on sampling intensities, on a 50 x 50 km UTM grid. The degree of sampling effort was assessed by means of accumulation curves based on the Clench function. Using the General Linear Model regression procedure, the effects of 22 variables on the estimated sampling efforts were assessed. This combination of methods is proposed as a preliminary step in biodiversity studies, in order to evaluate not only the degree of geographic coverage of existing faunistic data. but also the amount and nature of the bias on the faunistic work done throughout the last two centuries. The degree of spatial effects on the data was greater than the effects of environmental or land-use variables, although the latter two proved to be locally relevant. The results confirm previous findings that collecting is often skewed by relatively simple factors that affect collector activity, such as accessibility and attractiveness of sampling sites. With regard to Iberian and Balearic butterflies. adequate inventories on the scale investigated may probably suffice for further studies of the diversity of this insect group. Additionally, the results enabled us to develop general guide lines for the design of further faunistic work in the area.

A new species of the genus Stachorutes, Stachorutes cabagnerensis n. sp., from central Spain is described. It is characterized by the presence of 6+6 eyes in the head, retinaculum 2+2 teeth, dentes with 5 hairs, and the absence of mucron. A phylogenetic analysis of this genus was attempted. Potential synapomorphies supporting the monophyly of Stachorutes are presented. One member of the genus (the Nearctic S. navajellus) appears as a basal form, phylogenetically distant from the remaining (Old World) species. There is evidence for a monophyletic infrageneric clade with the species S. dematteisi, S. jizuensis and S. sphagnophilus. However, more information is required for further phylogenetic resolution., Se describe una nueva especie del genero Stachorutes de la region central de España. Stachorutes cabagnerensis nov. sp. se caracteriza por la presencia de 6+6 ojos en la cabeza, retinaculum con 2+2 dientes y 5 sedas en cada rama del dentes; la furca carece de mucrón. Se ha efectuado un análisis filogenético. Las sinapomorfias potenciales establecen la monofilia del género. Una especie del mismo, S. navajellus, aparece como forma basal, filogenéticamente distante del resto de especies (Viejo Mundo). Se podría establecer un clado infragenérico con las especies S. dematteisi, S. jizuensis y S. sphagnophilus. Sin embargo, se precisa de mayor información para poder confirmarlo.

A three-step protocol described elsewhere is used to obtain a map of butterfly species density in Portugal on a 50 x 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.

An extensive faunistic database has been set up gathering bibliographic information, data from scientific collections and other relevant unpublished data from the 143 species of continental Portuguese butterflies and burnets (Superfamily Papilionoidea, Families Hesperiidae and Zygaenidae). This database was used to define distribution patterns through the application of factorial correspondence analysis (FCA) to the matrices of frequency and presence-absence data in a 50 x 50 km UTM grid. The identification of faunistic elements and regions was obtained through an automatic application of the classification analysis to the FCA results. In a second instance, we used the same methodology to establish the relationship between previously obtained patterns and a set of environmental variables, in order to determine the factors related to the observed classes. The annual average temperature proved to be the best factor to explain the observed latitudinal gradient in the butterfly distribution. The results show a well-defined northern interior region of high average altitudes and limited in the South by the Serra da Estrela. This area has the largest species richness and hosts the Montesinho Natural Park, which is the most diverse in the country. In the southern half of Portugal, the Algarve deserves a special mention for its group of species with typical Mediterranean requirements.La información corológica (datos de bibliografía, colecciones e inéditos) sobre las 143 especies de mariposas (Superfamilia Papilionoidea, y Familias Hesperiidae y Zygaenidae) de Portugal continental se ha reunido en una base de datos. Ésta se emplea para definir patrones de distribución, aplicando el análisis factorial de correspondencias (AFC) a las matrices de frecuencias y de presencia-ausencia, basadas en una rejilla UTM de 50 x 50 km. En primer lugar, se identifican los elementos y regiones faunísticas mediante la aplicación automática del análisis de clasificación a los resultados del AFC. En una segunda fase, se establece la relación entre los patrones obtenidos y un conjunto de variables ambientales, para determinar los posibles factores asociados a las regularidades detectadas. La temperatura media anual es el principal factor explicativo del gradiente latitudinal observado. Queda bien definida una región norte interior, de altitudes medias elevadas, y limitada hacia el sur por la Serra da Estrela. Esta área presenta mayor riqueza de especies que el resto del territorio, e incluye el Parque Natural de Montesinho, donde se localiza la máxima diversidad de mariposas del país. En la mitad meridional del país, destaca la región del Algarve, con un conjunto de especies de requisitos biológicos típicamente mediterráneos.

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. Methods The 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 simplified 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 modification of PAE, PAE–PCE (Parsimony analysis of endemicity with progressive character elimination) was applied to overcome some of the potential shortcomings of the method. Results The results represent the first 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 identified. 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 fit within the frame of the overall similarity approach. Main conclusions The areas of endemicity identified 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 classification, 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 have no direct historical implications (for phylogenetic information is not incorporated), and (2) unless modifications 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' approach is to be preferred to a consensus of partial (taxon-specific) results.

Global climatic change exerts a generalised impact on species ranges, which could be problematic if ecologically-related species differ in their geographical response to climate change. Therefore, a combination of distribution modelling protocols is required to predict the future impacts of global warming on sets of different ecologically-dependent species. We test the extent to which the predicted future distributions of Spanish monophagous butterfly species, as estimated on the basis of climatic and physiographic variables, would differ depending on whether the geographic distribution of their larval food plants was taken into account or not. Using the favourability function as the modelling tool and fuzzy logic to combine butterfly and plant models, we extrapolated climatic favourability models for the butterfly, the plant and the butterfly–plant combination to the periods 2011–2040, 2041–2070, 2071–2100. All the models obtained were significant and the predicted butterfly–plant interactions indicated that larval food plants will represent in the future a greater constraint on butterfly species distributions. Climatic favourability for butterflies was expected to increase in the future more than the climatic favourability for the food plants. The plant data had relevant effects on the predicted future ranges of butterflies, which were generally expected to contract due to the effect of climate change on the plant. This highlights the view that opposite to recent results stating that climate is the primary driving force of butterfly distributions in Europe, variables other than those directly related to climate may exert a leading role in the near future. These include direct biotic interactions such as dependence on host plants, at least for highly specialised (monophagous) species of phytophagous insects.

The interspecific relationships between egg size and body size in butterflies (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 taxonomy-dependent 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.

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.

The Parque Regional de la Cuenca Alta del Manzanares (PRCAM) is a protected area at the North of Madrid. Eight sampling sites were selected representing different ecosystems and land uses. In these sites we studied the communities of four insect groups: Collembolla, Chrysomelida, Curculionoidea and Papilionoidea and Hesperioidea. The objetive of the study was to assess the conservation status of the Park ecosystems and the paralelism of the results in these four insect groups. The results show that the Park ecosystems are heavily disturbed. The lowland areas being damaged to a greater extent. On the other hand the four groups have a different behaviour in similar conditions. This shows the importance of gathering information from different groups in order to assess or manage the biological diversity of an area.El Parque Regional de la Cuenca Alta del Manzanares (PRCAM) es un área protegida del norte de Madrid. En el se han elegido ocho puntos representativos de sus ecosistemas y del uso del territorio. En estos puntos se han estudiado las comunidades de cuatro grupos de insectos: Colémbolos, Crisomélidos, Curculiónidos y Mariposas. El objetivo del estudio es valorar el estado de conservación de los ecosistemas del parque, y el grado de paralelismo de la información suministrada por los cuatro grupos de insectos sobre unas mismas áreas. Los resultados indican que los ecosistemas del Parque están muy alterados. El grado de deterioro es mayor en las zonas de menor altitud. Por otro lado las comunidades de estos cuatro grupos de insectos responden de formas distintas ante situaciones idénticas. De ello se deduce la importancia de tener en cuenta varios grupos de organismos a la hora de manejar o valorar la diversidad biológica de un zona.

The geographic range of a species is influenced by past phylogenetic and biogeographic patterns. However, other historical interactions, including the interplay between life history and geography, are also likely involved. Therefore, the range size of a species can be explained on the basis of niche-breadth or dispersal related hypotheses, and previous work on European butterflies suggests that both, under the respective guise of ecological specialisation and colonising ability may apply. In the present study, data from 205 species of butterflies from the Iberian peninsula were processed through multiple regression analyses to test for correlations between geographic range size, life history traits and geographic features of the species distribution types. In addition, the percentage of variance explained by the subsets of variables analyzed in the study, with and without control for phylogenetic effects was tested. Despite a complex pattern of bivariate correlations, we found that larval polyphagy was the single best correlate of range size, followed by dispersal. Models that combined both life history traits and geographic characteristics performed better than models generated independently. The combined variables explained at least 39% of the variance. Bivariate correlations between range size and body size, migratory habits or egg size primarily reflected taxonomic patterning and reciprocal correlations with larval diet breadth and adult phenology. Therefore, aspects of niche breadth i.e. potential larval diet breadth emerged as the most influential determinants of range size. However, the relationships between these types of ecological traits and biogeographic history must still be considered when associations between life history and range size are of interest.

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.

V. Dinca 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 Dinca (e-mail: vdinca@uoguelph.ca). 391

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.

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.

Aim: Species distribution models built with geographically restricted data often fail to capture the full range of conditions experienced by species across their entire distribution area. Using such models to predict distribution shifts under future environmental change may, therefore, produce biased projections. However, restricted-scale models have the potential to include a larger sample of taxa for which distribution data are available and to provide finer-resolution projections that are better applied to conservation planning than the forecasts of broad-scale models. We examine the circumstances under which the projected shifts in species richness patterns derived from restricted-scale and broad-scale models are most likely to be similar. Location: Europe. Methods: The distribution of butterflies in Finland, Belgium/Netherlands and Spain was modelled based on restricted-scale (local) and broad-scale (continental) distribution and climate data. Both types of models were projected under future climate change scenarios to assess potential changes in species richness. Results: In Finland, species richness was projected to increase strongly based on restricted-scale models and to decrease slightly with broad-scale models. In Belgium/Netherlands, restricted-scale models projected a larger decrease in richness than broad-scale models. In Spain, both models projected a slight decrease in richness. We obtained similar projections based on restricted-scale and broad-scale models only in Spain because the climatic conditions available here covered the warm part of the distributions of butterflies better than in Finland and Belgium/Netherlands. Main conclusions: Restricted-scale models that fail to capture the warm part of species distributions produce biased estimates of future changes in species richness when projected under climatic conditions with no modern analogue in the study area. We recommend the use of distribution data beyond the boundaries of the study area to capture the part of the species response curves reflecting the climatic conditions that will prevail within that area in the future.

Coverage provided by the network of protected areas in the Iberian Peninsula and Balearic Islands was tested by measuring the coincidence between the squares protected by the network and the butterfly species recorded for such UTM grid squares. Five species were found to be absent in the network. The protected areas with the highest numbers of butterfly species were Ordesa National Park and Monte Perdido and the Posets–Maladeta Natural Park. Priority areas were selected using WORLDMAP software and showed that the all species of butterflies in the Iberian Peninsula and Balearic Islands can be found within 16 squares of 10 x 10 km (nine of them not within the network of protected areas). More specific area selections were also carried out: eight squares supported the total number of threatened species, five hosted all the Iberian endemisms and 13 harboured the rare butterfly species. This study detected 16 squares that are not currently protected but are important for butterfly conservation in the Iberian Peninsula and Balearic Islands. Key words: Conservation, Butterflies, Gap analysis, Protected areas, Iberian Peninsula, Balearic Islands., Se ha analizado el nivel de cobertura que proporciona la red de espacios protegidos en la Península Ibérica e Islas Baleares comprobando la coincidencia entre éstos y el número de especies de mariposas registrado. Cinco especies quedan excluidas de la red. Los espacios protegidos con mayor número de especies fueron el Parque Nacional de Ordesa y Monte Perdido y el Parque Natural de Posets–Maladeta. Se realizó también una selección de áreas utilizando el programa WORLDMAP en la que 16 cuadrículas de 10 km de lado albergan a la totalidad de mariposas de la Península Ibérica y Baleares (nueve de ellas no se encuentran dentro de la red de espacios protegidos). También se realizaron selecciones de áreas más específicas: ocho cuadrículas fueron suficientes para albergar la totalidad de especies amenazadas, cinco para los endemismos ibéricos y 13 para las especies de mariposas raras. Basándonos en estos resultados se seleccionaron 16 cuadrículas en el área de estudio que son importantes para la conservación de mariposas y que actualmente no están protegidas. Palabras clave: Conservación, Mariposas diurnas, Análisis de huecos, Espacios protegidos, Península Ibérica, Islas Baleares.

1 Several references indicate that the period of flight of the European satyrine butterfly Hipparchia semele (L.) (Nymphalidae, Satyrinae) starts earlier in southern latitudes, where summers are longer and drier than in the north. However, summer drought has an adverse effect on the growth of grasses on which larval feeding depends. Growth of the grasses is delayed as long as the drought lasts. 2 From laboratory and field observations in a mid altitude area near the centre of the Iberian Peninsula, a mechanism that can be interpreted as an adjustment of this insect's life cycle to the host plant's phenology has been observed, i.e. delayed gonadal maturation of adult females. This delay is not associated with female diapause. Although the mean delay in oviposition after copulation was 43 days some captive females were able to oviposit much earlier, and this suggests variability in oviposition dates which might have an environmental or a genetic basis. 3 A mechanism of delayed ovarian maturation similar to that of H.semele is also known to occur in the satyrine Maniola jurtina (L.); it is suggested that this adaptation enables these species to occupy wider geographical ranges than other univoltine satyrines in Europe.

The fresh weight, dry weight, and C and N content of the eggs, egg shells and neonate larvae of several satyrines were measured. This was done in order to assess the specificity of the composition of the egg and larvae, the phylogenetic or ecological nature of the variation and the existence of structural constraints on the composition of the offspring. All the traits investigated were found to be highly species-specific. The nature of the variation was not primarily phylogenetic, suggesting that the composition of the offspring has an ecological meaning. However, only a slight association was detected between three life history traits or habitat features and the compositions of the eggs or larvae, namely: female egg dropping was associated with a high C content of the eggs, xerophily with a high C : N ratio, and a high content of N in the larvae with egg diapause. The evidence for intra-specific allometry between the traits investigated and egg weight varied among the species, suggesting that the slope of such relationship may be a spe- cific feature. There was a close to isometric relationship between C and N contents in every species. Therefore simple C : N ratios are independent of egg size, hence they can be used directly in comparative studies. Across species analyses indicated that small off- spring contained a proportionally low amount of carbon and had a high dry matter content, suggesting that selection for small eggs was accompanied by selection for an enhanced proportion of nitrogen per egg. Finally, the species with large adult females invested comparatively more nitrogen per egg, which indicates a potential, constraint-based advantage of large adult size.

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.