Publicaciones EA

This paper studies the effect of wastewater discharges on benthic macroinvertebrates in the Furnia River (Pontevedra, NW Spain). Semiquantitative surveys were carried out in spring 2008 and 2017 in three different locations,  upstream and downstream of a sewage treatment plant built in 2013. Different indexes were calculated based on benthic macroinvertebrate assemblages: abundance, richness, Shannon-Wiener, EPT, IASPT, IBMWP and several physicochemical variables were measured concurrently. Although the indexes values decreased slightly along the water course, the results indicate an optimal water quality of the Furnia River, supporting a very diverse community of aquatic macroinvertebrates.

We present the first records of Coquillettidia (Coquillettidia) buxtoni (Edwards, 1923) (n=15), Culex (Neoculex) impudicus Ficalbi, 1890 (n=12), Culex (Neoculex) territans Walker, 1856 (n=52), Culiseta (Culicella) fumipennis (Stephens, 1825) (n=535), and Culiseta (Culicella) morsitans (Theobald, 1901) (n=54) in the Autonomous Community of Galicia (NW Spain), including the first evidence of the genus Coquillettidia Dyar, 1904 in the northwest of Spain. The field research was carried out in different water ecosystems (ponds, streams, rivers, lakes, and lagoons) belonging to 39 zones throughout the entire Galician territory. The samples were collected intermittently and  seasonally through different projects of water quality monitoring between 1998 and 2018. A total of 1,614 mosquito larvae have been analyzed, and 668 of them were identified as species not previously recorded in Galicia. Between 1999 and 2003, the first catches of these species were obtained and since then, more specimens have been detected in different wetlands of Galicia, mainly in spring and summer.

We present the first records of Anopheles (Anopheles) plumbeus Stephens, 1828 and Culex (Culex) torrentium Martini, 1925 in the autonomous region of Galicia (NW Spain), obtained through the Rede Galega de Vixilancia de Vectores (ReGaViVec). This entomological surveillance network, which was initiated in 2017 by the Xunta de Galicia in collaboration with the University of Vigo and the University of Santiago de Compostela, aims to detect the  arrival of invasive vectors as well as to improve the knowledge about mosquito populations (Diptera: Culicidae) in the Galician territory. This study shows the first findings of these species in Galicia, which have been reported in six different locations throughout the region: five in the province of Pontevedra and one in the province of Lugo. The 51 captured specimens were collected at different stages of development between July and September, 2018 and 2019 by using specialized traps arranged in favorable regions for the settlement and development of culicids.

Despite the fundamental importance of hydroelectric power for socioeconomic development, the presence of hydropower plants cause large-scale alterations to the natural flow regime of rivers and profoundly influences aquatic processes and biodiversity. This study evaluates the seasonal impact of small hydropower stations by analysing and measuring macroinvertebrate community composition. Our objectives were to (1) examine whether the abundance and richness of benthic macroinvertebrates vary according to alterations to the river, (2) identify the families of macroinvertebrates most sensitive to depletion as a consequence of the impact of a hydropower station, and (3)  determine whether there is a seasonal component in river regulation that impacts on macroinvertebrate communities. A 167 848 individuals were sampled from 6 different phyla and 10 different orders of Arthropoda representing 116 different macroinvertebrate families, whose distribution and abundance depended on the river, sampling time and sampling site. Ephemeroptera, Diptera, Trichoptera, Coleoptera and Plecoptera were the most representative orders in all seasons of the year, and also had the most abundant families (Baetidae, Caenidae, Chironomidae and Simuliidae). There was no great variation in autumn and winter for the most abundant taxa but it was important variation in spring and summer. Our findings point to differences in macroinvertebrate abundance and richness in aquatic systems impacted by hydropower stations and to a recovery process downstream, where the water level and habitats are not negatively affected by these stations.

The feeding habits of 17 species of Nemouroidea stoneflies (belonging to the families Nemouridae, Leuctridae and Capniidae) from the north-western Iberian Peninsula are described and quantified by means of gut content analyses. As a whole, fungi, followed by pollen (both from angiosperm and, in a lesser degree, Pinaceae), leaf fragments (CPOM), detritus and, more punctually, lichens and animalmatter are themost common items found in their guts. Nevertheless, variations among species were observed. For a species with a flight period that extends throughout the year, Protonemura globosa, variations in the diet were noticed among seasons, with pollen being more important during the spring and summer period. Together with a few previous studies, the present one reaffirms that feeding is an important aspect of the adult life in males and females of Nemouroidea. © 2017 The Entomological Society of Japan.

We analysed the microhabitat preferences of caddisfly species in four rivers in north-western Spain. In each river, we sampled five sites with different types of substrate. These sites were characterised in situ according to the predominant material type (macrophytes, moss, pebbles and sand). A one-way ANOVA was used to test for significant differences in species abundance among substrates, and a nested ANOVA was used to test for significant differences in physical and chemical variables among rivers and sampling sites. A similarity percentage-species contribution analysis was performed to estimate the contribution of each species to the characterisation of each substrate. We observed significant differences in the abundance of seven species (Drusus bolivari, Glossosoma privatum, Larcasia partita, Micrasema longulum, M. servatum, M. gr. moestum and Sericostoma sp.) among substrates, confirming that these species have substrate preferences. According to similarity percentage-species contribution, the species that contributed most to differences were as follows: for sand, Sericostoma sp.; for macrophytes, Rhyacophila adjuncta, Hydropsyche ambigua, Sericostoma sp.; for moss, Micrasema gr. moestum, M. servatum, M. longulum, Hydropsyche tibialis and R. adjuncta; and for pebbles, Sericostoma sp., Larcasia partita and H. tibialis. These preferences can be explained by the fact that different species use different materials from the substrate to build their cases and also as a source of stability, protection and food. © CSIRO 2017.

Community assembly is determined by a combination of historical events and contemporary processes that are difficult to disentangle, but eco-evolutionary mechanisms may be uncovered by the joint analysis of species and genetic diversity across multiple sites. Mountain streams across Europe harbour highly diverse macroinvertebrate communities whose composition and turnover (replacement of taxa) among sites and regions remain poorly known. We studied whole-community biodiversity within and among six mountain regions along a latitudinal transect from Morocco to Scandinavia at three levels of taxonomic hierarchy: genus, species and haplotypes. Using DNA barcoding of four insect families (>3100 individuals, 118 species) across 62 streams, we found that measures of local and regional diversity and intraregional turnover generally declined slightly towards northern latitudes. However, at all hierarchical levels we found complete (haplotype) or high (species, genus) turnover among regions (and even among sites within regions), which counters the expectations of Pleistocene postglacial northward expansion from southern refugia. Species distributions were mostly correlated with environmental conditions, suggesting a strong role of lineage- or species-specific traits in determining local and latitudinal community composition, lineage diversification and phylogenetic community structure (e.g., loss of Coleoptera, but not Ephemeroptera, at northern sites). High intraspecific genetic structure within regions, even in northernmost sites, reflects species-specific dispersal and demographic histories and indicates postglacial migration from geographically scattered refugia, rather than from only southern areas. Overall, patterns were not strongly concordant across hierarchical levels, but consistent with the overriding influence of environmental factors determining community composition at the species and genus levels. © 2017 John Wiley & Sons Ltd

Aim: To reconstruct the evolutionary history of a genus of freshwater beetle with a pan-Mediterranean distribution, to test classic hypotheses which proposed a Miocene origin for groups with high biodiversity in the Iberian and Anatolian peninsulas. Location: Mediterranean basin. Methods: We sequenced four mitochondrial and one nuclear gene from 51 specimens of 30 of the c. 60 extant species of Deronectes (Dytiscidae), all typical of mid-mountain streams from North Africa and Iberia over most of Europe to the Middle East. We used maximum likelihood, Bayesian probabilities with an a priori evolutionary rate and a dispersal–extinction–cladogenesis model to reconstruct their biogeographical history. Results: Deronectes has two major lineages which originated in the mid Miocene; one including mostly eastern and another mainly western and central Mediterranean species. From these two areas, range expansions, mainly at the end of the Miocene and beginning of the Pliocene, resulted in the many species groups and some of the extant species of the genus. Most of the current diversity and distributions are, however, of Plio-Pleistocene origin, particularly in widespread European species. Main conclusions: In line with traditional hypotheses, we found an ancient division between eastern and western Mediterranean lineages of Deronectes, likely resulting from the isolation of Europe west of the Alps from the Balkans and Anatolia during the early-middle Miocene. The history of the genus was strongly influenced by major geological and climatic events, with successive cycles of fragmentation and subsequent eastward and westward range expansions, resulting in a steady accumulation of species across the basin. Most of these range movements took place through the north side of the Mediterranean, with only local displacements in the south during the Messinian salinity crisis and a recent (Pleistocene) colonization of the Italian Peninsula, which remained largely submerged through most of the genus’ evolutionary history. © 2016 John Wiley & Sons Ltd

During composting, facilities usually exert greater control over the bio-oxidative phase of the process, which uses a specific technology and generally has a fixed duration. After this phase, the material is deposited to mature, with less monitoring during the maturation phase. While there has been considerable study of biological parameters during the thermophilic phase, there is less research on the stabilization and maturation phase. This study evaluates the effects of the type of starting material on the evolution of microbial dynamics during the maturation phase of composting. Three waste types were used: sludge from the fish processing industry, municipal sewage sludge and pig manure, each independently mixed with shredded pine wood as bulking agent. The composting system for each waste type comprised a static reactor with capacity of 600 L for the bio-oxidative phase followed by stabilization and maturation phase in triplicate 200 L boxes for 112 days. Phospholipid fatty acids, enzyme activities and physico-chemical parameters were measured throughout the maturation phase. The evolution of the total microbial biomass, Gram + bacteria, Gram – bacteria, fungi and enzymatic activities (β-glucosidase, cellulase, protease, acid and alkaline phosphatase) depended significantly on the waste type (p < 0.001). The predominant microbial community for each waste type remained present throughout the maturation process, indicating that the waste type determines the microorganisms that are able to develop at this stage. While fungi predominated during fish sludge maturation, manure and municipal sludge were characterized by a greater proportion of bacteria. Both the structure of the microbial community and enzymatic activities provided important information for monitoring the composting process. More attention should be paid to the maturation phase in order to optimize composting. © 2016 The Authors.

Temporary ponds are habitats that undergo periods of drying and flooding. They have been neglected for many years and changes produced by climatic change will greatly affect them. Thus, nowadays they constitute an endangered ecosystem due to their characteristics and to human pressures. These habitats support a high biological richness with species adapted to extreme conditions. Assuming that hydroperiod is the main factor structuring aquatic assemblages in this type of ecosystem, the aim of this study was to assess the effect of the dry period on the faunal composition and the natural succession process of macroinvertebrate assemblages in two temporary ponds and to analyze the differences between two periods, before and after the dry period. A total of 7225 individuals belonging to 93 macroinvertebrate taxa (Nematoda, Hirudinea, Oligochaeta, Mollusca, Crustacea, Acari, Insecta) were collected. The most abundant and richest group were insects. Cluster and Non-Metric Multi-Dimensional Scaling (NMDS) analyses showed the clustering of the samples in two groups, before and after the dry period, as we had expected. Thus, there was a change in faunal composition in both ponds, corresponding to a successional process. According to the SIMPER analysis, the most contributive taxa in both ponds were mostly insects and crustaceans. Regarding feeding traits, predators and shredders were the dominant groups. However, there was a change in the trophic structure of the assemblages between the two periods. Most taxa resist the drying season with resting eggs, cocoons or simply by flying to more permanent freshwater bodies. Although the two studied ponds are temporary habitats, they support a different faunal composition hosting species that are endemic or rare at regional or national level. © 2015, Page Press Publications. All rights reserved.

In this work, we studied Trichoptera assemblages in different rivers in NW Spain affected by hydroelectric power stations, and assessed the influence of environmental variables on the distribution of species. Twenty sites in four rivers were sampled during eight sampling campaigns (2001-2002). The fauna was collected with a quantitative Surber sampler. In addition, several physical, chemical and habitat variables were measured at each site. A distance-based redundancy analysis (dbRDA) was done to investigate the relationship between the assemblages and the environmental variables. Assemblage composition was analyzed by non-metric multi-dimensional scaling (NMDS) and differences between groups were tested using the analysis of similarity (ANOSIM) considering two grouping variables, the river basin and the position of the site (location). The SIMPER routine was used to verify species contribution to dissimilarity. A total of 53 taxa of Trichoptera belonging to 16 families were found, 52% of which were Iberian endemics. The dbRDA revealed that altitude, conductivity, total suspended solids, temperature and location were the variables that most influenced the studied fauna. According to the NMDS analysis, significant differences in faunal composition were recorded between up and downstream sites and between river basins. The fauna seems to respond to a longitudinal gradient, but also to the impact of hydropower stations. The main effects we observed were variations in water temperature and changes in fauna composition, which may be due to the presence of hydropower stations. © 2015 Elsevier GmbH.

This study assessed riparian vegetation as a source of energy in different ponds. The trophic importance of food sources from the riparian zones for invertebrate consumers was analysed using the dual stable isotope technique. We measured stable carbon (δ13C) and nitrogen (δ15N) isotope values for three energy sources and primary consumers in four ponds in NWSpain to determine food sources that support primary consumers. Samples of littoral macroinvertebrates, macrophytes, epiphyton, and riparian vegetation were collected during two periods (summer 2010 and summer 2011) at each site.We used the MixSIR Bayesian mixing model to model the contribution of potential food sources of organic carbon supporting primary consumers at the study sites. The δ13C signatures of food sources varied between -35.5 ‰ and -20.76 ‰ , whereas the δ15N signatures ranged from -6.2 h to 11.27 h. Primary consumer isotope signatures corrected for the discrimination factor were within the range of the isotope values of the potential food sources in three of the ponds. Significant differences were observed in δ13C and δ15N signatures among sources at each site and overall ( p < 0.05). Significant differences were also found among sampling sites for each source separately and for overall food sources ( p < 0.05). TheMixSIR Bayesian stable isotope mixing model indicated that riparian vegetation and epiphyton were the most important production sources supporting the biomass of invertebrates overall. In addition, riparian vegetation and epiphyton were, most likely, a food source in the three ponds. Macrophytes were an unlikely food source for invertebrates in all ponds, although they could have some importance as a source in the Bodeira pond. © Asociación Ibérica de Limnología, Madrid. Spain.

The diet of Perla madritensis, endemic species of the northern half of the Iberian Peninsula, is described for the first time by means of a study carried out in north-western Spain. As other species of the genus Perla, this taxon behaves mainly as predator, with Chironomidae, followed by Baetidae, being the most abundant prey in its gut. Shifts in diet composition were detected in relation to size in this species, with smaller prey being replaced by larger ones such as Simuliidae and Leptophlebiidae when the nymphs become larger. © 2014 Société entomologique de France.

Wetlands are diverse and productive ecosystems endangered by human pressure, which degradation implies a biodiversity loss worldwide. Among the biological assemblages of these habitats, aquatic Coleoptera is one of the most diverse and useful groups when assessing the ecological conditions of the ecosystems they inhabit. The aims of the present study were to analyze the diversity and composition of aquatic Coleoptera assemblages in 24 wetlands protected by the Natura 2000 network of North-western Spain and the influence of environmental variables on the distribution of species, in order to detect differences between the different types of standing water habitats. A total of 11,136 individuals of 105 species belonging to 12 families of aquatic Coleoptera (Gyrinidae, Haliplidae, Noteridae, Paelobiidae, Dytiscidae, Helophoridae, Hydrochidae, Hydrophilidae, Hydraenidae, Scirtidae, Elmidae and Dryopidae) were collected. In general, wetlands presented high richness and diversity values, Dytiscidae and Hydrophilidae having the highest species richness. Most of recorded species have a wide biogeographical distribution and only 12 endemic ones were captured. Cluster and Non-Metric Multi-Dimensional Scaling (NMDS) analyses showed the clustering of the studied ponds and lagoons in four groups based on biological data. In general, the wetlands of each group seem to have distinct aquatic Coleoptera faunas, as showed by the most representative species. A combination of altitude, SST and hydroperiod was the best explaining factor of the distribution of the species throughout the study area. This study shows the high biodiversity of standing water habitats in North-western Spain and the usefulness of water beetles in establishing habitat typologies.

Coastal lagoons are high productivity ecosystems with high biodiversity, but very unstable systems. When assessing the ecological status of these ecosystems it is necessary to consider the influence of natural variations. In order to understand the community variation of coastal lagoons we studied spatial (among-lagoons) and temporal (among-years) changes in aquatic insect assemblages and variations in 23 measures potentially useful for bioindication purposes. Two series of data were examined: (a) aquatic insect families and (b) Coleoptera and Hemiptera species. Coefficients of variation were used to quantify the effect of among-lagoon and interannual variability on the selected metrics. Data analyses showed that within-lagoon variation (temporal) was higher than among-lagoon variation (spatial). Both aquatic insect families and Coleoptera and Hemiptera species provided similar information about among-lagoon similarity. According to the results, taxa richness, rarefied richness (ES200), species richness of Coleoptera and Hemiptera (%S CH), the Margalef diversity index (d), and richness and abundance of predators could be the most useful metrics when assessing ecological status in these coastal lagoons. Diversity metrics and fauna composition may be used together for a better understanding of the diversity patterns in these ecosystems.

Environmental drivers shaping beetle assemblages in four rivers of NW Spain affected by hydroelectric power stations. In this work, we studied the environmental drivers that shape the water beetle assemblages in four rivers from NW Spain. We also tested whether hydroelectric power stations have a negative impact on these assemblages. A total of 24931 specimens were collected at 20 sites in eight sampling fields (during years 2001-2002). The fauna was collected with a quantitative Surber net. In addition to habitat and spatial parameters, several physical and chemical variables were measured at each site. The environmental variables were then condensed by means of Principal Component Analysis (PCA). The relationship between species and environmental variables was explored by Canonical Correspondence Analysis (CCA). The assemblage composition was analyzed by a non-metric multidimensional scaling (NMDS), and the differences between groups were tested using the analysis of similarity (ANOSIM) considering two factors, the NMDS grouping and the power station impact. The SIMPER routine was used to verify the species contribution to the dissimilarity. We detected a total of 47 species of Coleoptera belonging to 11 families, 43 % of which were Iberian endemics. A Canonical Correspondence Analysis revealed that altitude, distance from source and conductivity were the variables that most influence the studied fauna. Significant differences in the faunal composition were registered in NMDS groups 1 and 2, which were made up of sites belonging to different river basin and lithology. Moreover, differences related to the hydroelectric power stations impact could not be detected due to the experimental design. The SIMPER routine highlighted that Elmis aenea (Müller, 1806), Dupophilus brevis Mulsant & Rey, 1872 and E. rioloides (Kuert, 1890) were the species that most contributed to explain the differences between groups.

Diversity and assemblage structure of water beetles sampled from twenty eight streams covering northern Tunisia have been analyzed. In addition to habitat and spatial variables, several physical and chemical variables were measured at each site. Samples from mountainous streams were dominated by rheophilous species of the family Hydaenidae and the subfamily Hydroporina while lowland streams are colonized by limnophilous species belonging to the families Hydrophilidae, Haliplidae and Dytiscidae. The most abundant taxa were Berosus affinis, Haliplus lineaticollis, Laccobius atratus, Hydroporus feryi, Laccophilus minutus, L. hyalinus and Hydraena leprieuri. Species richness revealed seasonal and spatial variation. Non-parametric multidimensional scaling (NMDS) ordination highlighted the faunistic affinities between sites revealing five main groups. Species-environmental variables relationships were examined using principal component analysis (PCA) showing two main species groups. Conductivity, water temperature, pH, dissolved oxygen, water permanence, current velocity, substrate type, altitude and abundance of aquatic vegetation were identified to be the major drivers structuring beetle assemblages and influencing species abundance and richness. These results suggest that habitat heterogeneity was the predictor of beetle assemblages, while species richness could be expected at a landscape scale using physicochemical features.

The understanding of community structuring factors is a fundamental issue in community ecology. We investigate the biodiversity of the water beetle in the Ichkeul Lake stream system located in Northern Tunisia, with emphasis on the effect of several environmental variables on their assemblage structure. This area is of a great importance since it was included in the list of preserved waterbodies by RAMSAR convention on wetlands, MAB (Man And Biosphere), UNESCO (United Nations Educational, Scientific and Cultural Organization). Consequently, the study of its biodiversity constitutes a required way to better conduct this protection aim. Special attention must be paid to species endemic to North Africa, as all of them require conservation measures. Sampling was carried out over the course of a year. Dissolved oxygen, conductivity, pH, temperature and turbidity were measured for each study site. Water permanence, velocity, aquatic vegetation and substrate type were considered. The investigated streams showed a spatial heterogeneity in water depth, water velocity, turbidity, mineralization and substrate type, but homogeneity in temperature and dissolved oxygen. A total of 68 coleopteran species were identified from the investigated area, belonging to different chorotypes and phenological categories. The coleopteran assemblage was assessed in terms of four community parameters: abundance, richness (S), diversity (Shannon-Wiener index: H′), and evenness (Pielou index: J′). Species composition was analyzed by multivariate analysis. Sites were arranged into two groups based on their species affinity by non-parametric multidimensional scaling (MDS) ordination on the basis of Bray-Curtis similarity at a similarity of 50%. Principal component analysis (PCA) highlighted the species response to the environmental variables. It revealed the distribution profile of sampled species; on the first axis, F1, a maximum number of variables are projected. Its positive pole is defined by the majority of species, with tolerance of high salinity. On its negative pole only three species are present. Its positive pole is defined by lentic phytophagous species, showing a preference for well vegetated waters. Its negative pole is marked by the species L. orientalis, associated with the majority of water parameters. © 2013 Société entomologique de France.

We analysed the microhabitat preferences of water beetle species in four rivers in Northwest Spain. In each river, we sampled 5 sites with different types of substrate. These sites were characterised in situ according to the predominant material type (macrophytes, moss, pebbles and sand). The occurrence of a substrate preference was verified from a comparative study of species richness and abundance among different microhabitats. The differences in abundance and richness between substrates and in the abundance of each species were tested with an ANOVA. The similarity between microhabitats was tested with non-metric multidimensional scaling (NMDS), and the correlation between fauna and substrates was verified with a correspondence analysis (CA). We observed different species distribution patterns, and these patterns reflected the microhabitat preference of each species. Both the ecological parameters and the correspondence analysis indicated that the preferred substrate for most of the species was moss, followed by pebbles. © Asociación Ibérica de Limnología, Madrid. Spain.

Recent molecular analyses revealed unexpected genetic variability within Hydraena gracilis, one of the most common and widespread European moss beetles, belonging to the West Palaearctic endemic «Haenydra» lineage. For this taxon, molecular data univocally evidenced two distinct clades, the first one including populations from the Iberian Peninsula, and the second including populations from central and eastern Europe. Aedeagal morphology of H. gracilis was further investigated, finally demonstrating the actual existence of a new cryptic species of the H. gracilis complex in Portugal, Spain and the French Pyrenees. The geographical range of the true H. gracilis was consequently revised. Finally, dichotomous keys for males of all species belonging to the H. gracilis complex, and for males of all the Iberian «Haenydra» species, are also supplied. © 2012 Taylor and Francis Group, LLC.

We review the distribution of the water scavenger beetles (Hydrophilidae) in the northern part of Tunisia, based on data from 54 sites collected over the course of a year. In total, 21 hydrophilid species were found. The most interesting ones from the zoogeographical point of view are: Enochrus nigritus Sharp, 1872, Enochrus affinis Thunberg, 1794, Laccobius revelierei Perris, 1864 (all newly recorded for Tunisia), and Laccobius orientalis Knisch, 1924 and Berosus spinosus Steven, 1808 (newly recorded for northern Africa). An updated checklist of hydrophilid species occurring in the northern Tunisia is provided. The composition of the hydrophilid fauna in the area is assessed using the indexes of richness, abundance and diversity. We discuss possible causes of this phenomenon; identify the most valuable water beetle habitats within the investigated area that should be protected by law. © Pacific Coast Entomological Society.

Aim We test whether species of western Mediterranean aquatic Coleoptera of the ‘Haenydra’ lineage (Hydraenidae, Hydraena) originated through: (1) successive periods of dispersal and speciation, (2) range fragmentation by random vicariance, or (3) range fragmentation by geographic isolation owing to a general reduction of population density. Location Europe. Methods To discriminate between scenarios we use contrasting predictions of the relationship between phylogenetic and geographic distance. The phylogeny was based on 3kb of four mitochondrial and two nuclear gene fragments of about half of the known species of ‘Haenydra’, including most western Mediterranean taxa. Divergences were estimated using a molecular clock. The relationship between phylogenetic and geographic distance was tested using bivariate plots, Mantel tests and comparison of the observed phylogeny with the one minimizing geographic distances between species, as measured using Euclidean minimum spanning trees (EMSTs). Results The monophyly of ‘Haenydra’ was strongly supported, although its phylogenetic placement was not resolved. ‘Haenydra’ was estimated to be of late Miocene age, with most species originating during the Pleistocene. In two clades (Hydraena tatii and Hydraena emarginata clades) there was a significant association between geographic and phylogenetic distance, and the reconstructed phylogeny was identical to that obtained through the EMST, demonstrating a strong non-randomness of the geographic distribution of the species. In two other clades (Hydraena iberica and Hydraena bitruncata clades) there was no association between geographic and phylogenetic distance, and the observed phylogeny was not the one minimizing geographic distances. In one of the clades this seems to be due to a secondary, recent range expansion of one species (H. iberica), which erased the geographic signal of their distributions. Main conclusions We show that it is possible to obtain strong evidence of stasis of the geographic ranges of narrow-range endemic species through the study of their phylogenetic relationships and current distributions. In at least two of the studied clades, current species seem to have originated through the fragmentation of a more widely distributed species, without further range movements. A process of range expansion and fragmentation may have occurred repeatedly within the ‘Haenydra’ lineage, contributing to the accumulation of narrow-range endemics in Mediterranean Pleistocene refugia. © 2010 Blackwell Publishing Ltd.

An ecological study of the geographical distribution of aquatic Adephaga and Polyphaga (Coleoptera) in the mountains of northern Tunisia was carried out with an analysis of the effect of altitude on the distribution of 123 species collected from 64 sampling sites. Species richness was analyzed at different altitudinal levels and the indicator species were determined by establishing their altitude profile in terms of reciprocal species-factor information. Species richness decreases with increasing altitude. Thirty representative species are placed into three groups: species present along the entire altitudinal gradient; species present at high altitude; and species occurring at low altitude.

The nymphal feeding of Siphonoperla torrentium (Pictet, 1842) in Galicia (NW Iberian Peninsula) is studied. The main diet component is CPOM (approximately 33.5 % of the gut content), followed by animal prey, detritus, fungi and pollen. Chironomidae was the main ingested prey. Particularly outstanding is the importance of pollen in the feeding of these nymphs. Ontogenetic shift in the diet is detected only for prey, with animal matter being more important in the largest nymphs. Thus, the nymph of 5. torrentium in the study area can be catalogued as mainly shredder and, especially when older, as predator. Obtained data are compared with those previously known for this species in other areas, showing that different populations of the same species can present different diets and they can even be catalogued in different functional feeding groups.

Chorological and phenological analyses of the aquatic beetle fauna (Gyrinidae, Haliplidae, Hygrobiidae, Noteridae, Dytiscidae, Hydrophilidae, Hydraenidae, Dryopidae, Elmidae, and Scirtidae) of northern Tunisia are presented. Water beetles were sampled from freshwater ecosystems at 64 sites distributed throughout northern Tunisia. In total, 123 species were identified and their distribution in the study area is discussed. Chorological categories were determined according to the geographical distribution of sampled species. Three groups of species were established based on their phenology: permanent, frequent, and seasonal species.

Native and exotic seaweeds frequently lie on the beach and sustain part of the benthic food web. However, the role of exotic seaweeds as food sources for beach consumers has been poorly studied. We studied the temporal and spatial variability in the trophic significance of the invasive brown seaweed Sargassum muticum on sandy beaches. We measured the stable isotopes (δ13C and δ15N) in the tissues of S. muticum and of invertebrate consumers and estimated the dietary biomass proportion of S. muticum during four sampling dates at two beaches and heights on the shore. Samples were collected from eight pitfall traps placed at a distance of 2 m from each other. Detrital macroalgae and seagrasses were also collected by hand within an area of 30 cm around each pitfall trap. We measured the spatial and temporal variability in the isotope composition of the beach consumers and of S. muticum using different models of analyses of variance. We then calculated the biomass proportion of S. muticum to the animal diet with a two-isotopic mixing model. The invasive alga S. muticum seemed to be one of the main food sources for the amphipod Talitrus saltator and, to a less extent, for the isopod Tylos europaeus. The importance of S. muticum was however temporally variable and decreased during spring (in March and May), probably due to the availability of native macrophytes. The supply of invasive wrack to beach food webs thus deserves more attention if we want to understand their role in influencing food web dynamics. © 2009 Elsevier B.V. All rights reserved.

The accumulation patterns of different trace elements (As, Cd, Cr, Cu, Fe, Mn, Ni, Pb and Zn) were studied in three species of dragonfly larvae (Cordulegaster boltoni, Boyeria irene and Onychogomphus uncatus). Additionally these species were assessed as potential biomonitors in a lotic ecosystem (Louro River, Spain). Element concentrations were determined by inductively coupled plasma optical emission spectrometry (ICP-OES) in both dragonfly larvae and river sediments. The surface of the larvae was observed and analyzed with scanning electron microscopy (SEM). A desorption test was used to establish the percentage of element adsorbed into the exoskeleton. Field biomonitoring studies were made considering the first part of the river as a control area in relation to the second part, which was severely polluted by industrial and domestic activities. Upon application of principal component analysis (PCA), two different element groups were found in relation to element bioaccumulation. Cu, Cd and Zn were mainly associated with the inner part of the larvae. As, Cr, Fe, Mn, Ni and Zn were found on the outer body parts, related with deposition of oxyhydroxides in the hydrocycle. SEM revealed a layer of inorganic particles, similar in composition to fine bed sediments. Significant relations between the element content of this group and that of sediments at the sampling site were found. Differences in bioaccumulation for each of the three species, except for As, were observed. Dragonfly larvae revealed their ability to reflect environmental concentrations of As in freshwater media. © The Royal Society of Chemistry 2010.

The cycling of organic matter through food webs is a fundamental process that may be altered by the invasion of non-indigenous plants. We explored consequences of the invasion of non-indigenous Spartina patens to the composition of soil organic matter (SOM) and to detritivore and herbivore diets in the upper salt marsh within Corrubedo National Park, northwest Spain. We tested for the incorporation of S. patens carbon (C) into SOM and by detritivores and herbivores using stable isotope analysis, focusing primarily on detritivorous enchytraeid oligochaetes and herbivorous insects. Stable isotope results indicated that C derived from S. patens has been incorporated into SOM. Elevated densities of enchytraeids in stands of S. patens, and their incorporation of C derived from this plant, suggested that dense patches of S. patens may facilitate detritivore populations. In contrast, although insect herbivores used S. patens as habitat, there was little isotopic evidence for the widespread incorporation of S. patens-derived C by these consumers. The population and dietary response of enchytraeids to S. patens suggests that S. patens invasion could indirectly influence soil processes and pathways mediated by detritivore activity (e. g., soil respiration rates, nutrient retention and transformation, energy flow). The loss of food resources to insect herbivores alters local food webs. However, insect herbivores may move and feed on native plants elsewhere. As a result, insect populations may be less immediately impacted than soil detritivore populations by S. patens. Our study suggests that the influences of S. patens invasion extend beyond the more obvious changes in native plant abundance, to include differing responses in the cycling of organic matter between detritivore and insect herbivore food web pathways. © 2010 The Author(s).

The diversity of true water beetles (Coleoptera: Gyrinidae, Haliplidae, Dytiscidae, Helophoridae, Hydrochidae, Hydrophilidae, Hydraenidae, Elmidae, and Dryopidae) in Picos de Europa National Park (Cantabrian Mountains, Spain) was examined. Taking into account historic long-term sampling (all collections from 1882 to the present), a total of 117 species are recorded. Species accumulation models and non-parametric estimators were used to estimate the actual species richness of aquatic Coleoptera occurring in Picos de Europa National Park. Estimates were generated by analyzing both the collector’s curve from the long-term sampling and the historic cumulative curve of species recorded from the park since 1882. Values of species richness estimated by different methods range from 127 to 170 species (mean = 148 ± 15 SD). Therefore, it seems that the current inventory has reached a reasonably good level of completeness as estimates indicate that about 80% of the water beetle fauna has already been recorded. The inventory is used to analyze the biological uniqueness of the park and its outstanding level of species richness and endemism (33 Iberian endemic species). Richness and endemism of water beetles in Picos de Europa National Park is compared to those of other national parks in Spain. Finally, the conservation status and threat level is assessed for two Cantabrian endemic species, Deronectes costipennis gignouxi Fery and Brancucci and Ochthebius (Asiobates) cantabricus Balfour-Browne.

The objective of this study was to identify the response of aquatic Coleoptera to water quality and habitat variables in two rivers in northwest Spain with different degrees of contamination. The water beetle assemblages of the Lagares and Minor Rivers were studied during an annual cycle (May 2001-January 2002). Several physical and chemical variables, in addition to habitat and spatial factors were measured at each site. The environmental variables were then subjected to PCA to facilitate the interpretation of the principal components. The structure of the assemblage was evaluated for different richness measures: Species richness rarefied (ES); Abundance (N) AND Shannon-Wiener Diversity Index (H’). Regression analysis was used to examine the relationships between the richness measures and the water quality variables and the relationship between species and environmental, variables was explored by Canonical Correspondence Analysis (CCA). Differences in assemblage structure were examined using analysis of similarities with Bray-Curtis index. We found a total of 35 species of Coleoptera belonging to 10 families, 37 % of which are endemic to the Iberian Peninsula. All richness measures decrease in degraded sites, with a negative correlation between these factors and sites with high levels of contamination. Data analyses also indicated significant differences among the assemblages of non-polluted and polluted waters. CCA showed that most of the species seemed to be sensitive to pollution, only Limnius volckmari (Panzer) and Elmis aenea (Müller) might be considered tolerant species.

The aim of this study was to gather new data about the life cycle phenologies of several species of aquatic insects in the «Gándaras de Budiño» (Galicia, NW Spain) protected wetland, included in the Natura 2000 network. During an annual cycle (2004-2005), three shallow lakes and four streams were sampled monthly using a semi-quantitative sampling method. The body lengths of the larvae and nymphs of thirteen species were measured, and their life cycles were analysed. All species had univoltine or semivoltine cycles. Additionally, a possible correlation between larval and nymphal lengths and water temperature was examined. We found a significant correlation for two species: the water beetle (Noterus laevis) and the dragonfly (Boyeria Irene). It is important to highlight the presence in the studied area of a species included in the Habitats Directive: the damselfly (Coenagrion mercuriale).

Wrack (dead, washed-up seaweed and seagrass) buried in soft substrata may increase the organic content and alter the physical structure of sediments. These effects may influence the composition and structure of macrofaunal assemblages in the sediment. Such influences can be expected to vary according to the type and amount of wrack as well as the presence of invasive seaweeds in the wrack. In this study, we deliberately buried different amounts of the invasive species Sargassum muticum in isolation or mixed to the native species Ulva sp. and Fucus vesiculosus, in two intertidal sandflats to test some hypotheses about the response of macrofaunal assemblages. We tested whether (1) diversity of detritus (i.e. different mixtures), and (2) the amount of detritus of S. muticum influenced the composition and the relative abundance of macrofaunal assemblages. We also assessed whether the sediment organic carbon and the biomass of benthic microalgae varied depending on the diversity of detritus and the amount of detritus of S. muticum. Finally, we tested if these effects of wrack were consistent across sites. Results indicated that buried wrack affected the composition and structure of macrofaunal assemblages in short-term (i.e. 4 weeks), but there were no differences depending on detritus diversity or the amount of S. muticum. In addition, sediment organic matter and microalgal biomass were not affected by the addition of wrack. They instead varied greatly among small spatial scales (i.e. plots). Wrack composition or abundance of the invasive species S. muticum played thus a small role in shaping the structure of macrofaunal assemblages or the biomass of benthic microalgae in these intertidal sediments, probably because these sediments are frequently affected by various inputs of organic matter and benthic assemblages are already adapted to organically enriched sediments. © Elsevier B.V.

In this work we present the data of the study of the aquatic coleopteran community of the Gandaras of Budiño wetland (LIC area) (Pontevedra, NW of Spain). Semi quantitative samplings were conducted monthly during a year (february 2004-february 2005) in 8 points of two different types of habitat (4 in ponds and 4 in streams). A total of 2610 individuals belonging to 40 species of the families Gyrinidae, Haliplidae, Noteridae, Dytiscidae, Helophoridae, Hydrochidae, Hydrophiiidae, Hydraenidae, Scirtidae, Elmidae and Dryopidae were collected. Different community parameters (species richness, abundance and diversity) were analysed, as well as the influence of different environmental variables (water temperature, pH, conductivity, % O2, and TDS) on the community. The results suggest a marked deterioration of the stagnant waters of the wetland due to a possible pollution. © Asociación Ibérica de Limnología, Madrid, Spain.

Talitrid amphipods are the most abundant herbivores on exposed sandy beaches. Despite their important role as trophic intermediates between macrophytes and higher levels (i.e. insect and bird) of beach food webs, very little information is available on their feeding patterns. The main aim of this study was to investigate intraspecific differences in the feeding behaviour of Talitrus saltator. We tested the hypotheses that: (1) adult females and males showed different isotope signatures and therefore relied on different sources of food; and (2) patterns of variation of isotope signatures of juveniles differed from those of adult specimens, evidencing a diet shift during the development. We used stable isotope signatures and tested for differences upon the level on the shore, times of the year and beaches experiencing similar morpho-dynamic and environmental conditions. Finally, we investigated the trophic significance of macrophyte detritus in the diet of males, females and juveniles. Results showed that adult males had a more variable diet than females and juveniles (inferred from δ13C and δ15N values). Dual-isotope graphs suggested that Sargassum muticum and Cystoseira baccata wrack could be among the main food sources for both juvenile and adult stage. © 2009 Elsevier Ltd. All rights reserved.

The paper deals with the composition of the arthropod fauna in nineteen shag (Phalacrocorax aristoteli Linnaeus, 1761) nests collected in two sampling sites from the National Park of the Atlantic Islands: in the Cíes and Ons Islands located in the NW of the Iberian Peninsula during 1997. The occurrence and abundance of arthropods in the nests have been studied in three different breeding phases: nests with eggs, nests with broods and empty nests. In these nineteen nests, the authors found 25 families of acari (92.1 % of the total fauna), 14 families of Diptera (98 % were in a saprophagous larval stage), 8 families of Hymenoptera, 16 families of Coleoptera and 5 families of spiders. The ecological and the trophic relations of these families have been studied. A large proportion of arthropod families found in the nests occur there occasionally. Furthermore, the nests provided few groups with the right conditions to develop and complete their biological cycle. Only saprophagous mites, fly larvae and predator beetles found suitable conditions to survive in the nests. Saprophagous arthropods were the most abundant, independently of the nest occupation phase. The difficult access to the cliffs along with the necessity to transport the nests into the laboratory hampered the sampling process and consequently made this study the first about entomological fauna in shag nests.

1. The objectives of the present study were to describe and analyse the composition and structure of aquatic Coleoptera and Hemiptera assemblages in three coastal lagoons of north-western Spain during a one year cycle, in order to evaluate their relative adequacy to provide information about the conservation value of lagoonal habitats and sites. The lagoons are designated as Special Areas of Conservation (SAC) under the European Union Habitats Directive and two of them are also protected by the Ramsar Agreement. Several abiotic variables, including salinity, were recorded at the time of sampling. 2. In total, 67 species (52 Coleoptera and 15 Hemiptera) and 6568 adult individuals (2664 Coleoptera and 3904 Hemiptera) were collected. In all pair-wise comparisons Kendall’s coefficients of concordance between lagoons were higher for Hemiptera than for Coleoptera, indicating that Hemiptera assemblages were more similar across sites. 3. Most species recorded had a wide Palearctic distribution and only three species of water beetles could be considered endemic to Iberia: Hydroporus vagepictus, Hydroporus vespertinus and Hydrochus angusi. Similarly, the species collected had a widespread distribution in the Iberian Peninsula as the only species considered to be rare taxa at this scale were Hydrochus angusi and Cymbiodyta marginella among the Coleoptera and Sigara scotti and Notonecta glauca glauca among the Hemiptera. 4. The results do not support the idea of distinct aquatic insect assemblages for coastal lagoons in the Iberian Peninsula. Comparisons of the numbers of interior (non-coastal) and coastal provinces of the Iberian Peninsula where the species had been recorded showed they had been recorded in a larger proportion of interior provinces. 5. Salinity seemed to reduce species richness in both groups while the results of a canonical correspondence analysis (CCA) showed that the majority of species responded negatively to salinity. 6. This study suggests that species richness and rarity of aquatic insect assemblages may underestimate the conservation value of lagoonal habitats because their net contribution to catchment biodiversity is likely to be low. Copyright © 2008 John Wiley & Sons, Ltd.

In this work the influence of the concentration of dissolved iron and nitrate in the water on the structure of the community of aquatic Coleoptera of a river in northwest Spain is analyzed. A total of 45 species and subspecies of aquatic Coleoptera (Adephaga and Polyphaga) were collected between March and November 1999 from 10 sites distributed along the Avia river (Ourense, N.W. Spain). Also, 24 physical and chemical parameters of water were measured and a CCA was made to evaluate which of these factors had the greatest influence on the studied fauna. At the same time an ecological analysis was made using the distribution of species and subspecies in relation to two water parameters (concentration of nitrate and dissolved iron). The study of distribution in relation to these factors by means of elaborating their ecological profiles suggested indicator species for each of these parameters. Seventeen species were found to be indicators for at least one of the parameters analyzed. The species Deronectes costipennis gignouxi, Enichocerus legionensis, Hydraena testacea, Megasternum obscurum, Nebrioporus depressus elegans and Stenelmis canaliculata are the best indicators of these parameters, if we consider only taxa collected in class 4, that is, with high values of iron and nitrate. © Asociación Ibérica de Limnología, Madrid.

This study deals with the analysis of the coleopteran assemblages in a beachdune system located on the Galician coast. In particular, we used experimental manipulation of algal wrack, i.e. artificial patches of Saccorhiza polyschides, to test hypotheses about influences on coleopteran assemblages inhabiting the upper shore level along an exposed sandy beach. Specifically, we tested that (1) abundance of colonizing individuals and species vary with the size of wrack patch and time, and (2) as a result, coleopteran assemblages also vary with patch size and time. Furthermore, responses could differ among sites because of their different environmental conditions. A total of 3 980 individuals belonging to 8 families and 18 species were collected. Tenebrionidae was the most abundant family followed by Staphylinidae and Hydrophilidae families, respectively. The tenebrionid Phaleria cadaverina (Fabricius, 1792) was the most abundant species within this family, with a total of 1 064 adults and 1 124 larvae. Phytosus spinifer Curtis, 1838 was the most abundant staphylinid species represented by 736 adults and 3 larvae. This study showed that coleopterans are very common colonizers in wrack patches on sandy beaches. Patterns of succession differed among species and colonization of patches was rapid, i.e. most species colonised patches on day 3. Coleopteran assemblages changed depending on patch size and time. Wrack may act as islands of food and/or habitat for several coleopteran species on sandy beaches. Cleaning activities on beaches used for recreational purposes might alter the diversity of coleopteran assemblages associated to wrack patches. Thus, for managerial and conservational purposes cleaning activities on beaches should be regulated.

In this study, we used experimental manipulation of algal wrack to test hypotheses about influences on macrofaunal assemblages inhabiting the upper shore level of different sites along an exposed sandy beach. First, we hypothesized that decomposition of algal wrack depends on wrack patch size and site. With respect to macrofauna, we tested the hypotheses that (1) abundance of colonising individuals and species vary with wrack patch size, (2) succession (i.e. sequence of colonisation and species replacement) depends on time, and (3) as a result, macrofaunal assemblages associated with wrack patches vary with the patch size and time. We also predicted that responses could be different across sites because of their slightly different environmental conditions. The decomposition of wrack patches was similar in all sites and was dependent on wrack patch size. It was strongly influenced by time-specific environmental and/or biological factors. The pattern of colonisation, i.e. total number of species and individuals, varied among wrack patch sizes. Small patches had fewer species and individuals than medium and large patches. Nevertheless, pattern of colonisation varied among species, across sites and through time. Colonisation of wrack patches was rapid (i.e. within 3 days) for most species. There was some evidence to support the hypothesis that macrofaunal assemblages change in response to patch size and time. © 2006 Elsevier Ltd. All rights reserved.

In this work we studied the fauna of aquatic Coleoptera (Adephaga and Polyphaga) of the hydrographic river basins of the Lagares and Miñor Rivers (Pontevedra, NW of Spain). Twenty sites were sampled in four seasonal campaigns between May of 2001 and January of 2002. A total of 1,597 specimens belonging to 35 species of the families Haliplidae, Gyrinidae, Dytiscidae, Helophoridae, Hydrochidae, Hydrophilidae, Hydraenidae, Elmidae, Dryopidae and Scirtidae were collected. The family Hydrochidae (Hydrochus angustatus), the genus Esolus (Esolus parallelepipedus), and Megasternum (Megasternum concinnum), and the species Helophorus flavipes, Elmis aenea, and Elmis maugetii maugetii are all new records for the province. Spatial and seasonal variations in richness, abundance and diversity were analysed, in addition the similarity between sites was also analysed based on the total fauna composition of each. The results indicate that the typical communities of non-polluted waters are separated from those of polluted waters, indicating the negative influence of the contamination on the fauna. © Asociación Española de Limnología.

Extant populations of Orculella bulgarica (Hesse, 1915) are described from Iberia for the first time. We present the first description of the reproductive system anatomy of the species, as well as the structure of the periostracum of young shells. Assignment to the genus Orculella Steenberg 1925 is confirmed. The populations in Iberia are found in hygrophilous habitats such as small marshes. It is recommended that the species be classified as critically endangered based on the small size of the four known populations, three of which have disappeared in the last decade.

New data concerning the morphology of males and females of Hydraena bisulcata Rey, 1884, and H. barrosi d’Orchymont, 1934, are given, including ilustrations. The taxonomic position and Iberian distribution of these species are discussed. Two previously proposed synonymies are validated: Hydraena bisulcata Rey, 1884 (= H. berthelemyi Lagar et Fresneda, 1990) and H. barrosi d’Orchymont, 1934 (= H. valladaresi Lagar et Fresneda, 1990).

Aquatic Coleoptera in shallow lakes associated with the Canal de Castilla (Palencia Province, Spain) in the northern Iberian Meseta were sampled over the course of a year (spring 1998-winter 1999). These waterbodies are typical plateau wetlands with dense vegetation and vary in permanence and area (from 3.3 ha to 29.35 ha). Oxygen concentration, conductivity and pH were recorded at the time of sampling. Lake area, depth, water permanence and type of vegetation were also taken into account. Ninety two species were collected. Species richness was high in comparison with other wetlands in Spain. The assemblage structure was assessed in terms of three community parameters: richness, abundance and diversity (Shannon index). Their relationships with environmental variables were explored using correlation coefficients. The assemblage composition was analysed by multivariate techniques. First, the sites were classified by means of TWINSPAN. The presence of each species in the different TWINSPAN groups was used to assess their habitat preferences. Second, the sites and species were ordinated by Detrended Correspondence Analysis (DCA) using the CANOCO statistical package. Richness was significantly correlated with water permanence, conductivity and aquatic macrophyte cover. The first DCA axis was significantly correlated with water permanence and conductivity, but not with any of the other parameters. Conductivity was significantly inter-correlated with permanence. Therefore, water permanence and aquatic vegetation cover seem to be the main factors influencing richness, but only water permanence appears to determine species composition.

The study of the chorology and ecology of the andalusian terrestrial malacofauna (South of the Iberian Peninsula), has showed the presence of a group of climatic relicts characteristics from the Eurosiberian region, in the Eastern Betics Mountains Chains. The ecology and distribution of two of these species (Vitrina pellucida O. F. Müller, 1774 and Cepaea nemoralis Linnaeus, 1758) is exposed, being the main limiting factor the typical mediterranean summer drought. The adaptative response of these species against the scarcity of water depends on its different ecophysiologies. Vitrina pellucida (eurithermic) lives in low-lands where the availability of water along the year is constant, as well as in dry soils of high mountains because of the cold weather. Cepaea nemoralis (stenothermic) lives only in habitats with wet soils and heavy vegetal cover placed below 1500 m. The absence of similar vegetation over this altitude and the extreme climate conditions of the High Mediterranean Mountains prevents its appearance, contrarily to the High Eurosiberian Mountains.

We have studied 415 specimens of coleoptera belonging to the genus Hydraena Kugelann, 1794, collected during random sampling carried out at different sites in the Iberian Peninsula and the Balearic Islands. New localities are given for the 24 species identified, 14 of which are recorded for the first time in seven provinces. We also provide additional comments concerning their geographical distribution.

Aquatic beetles were sampled on a monthly basis between July 1991 and June 1992 in the wetland pond of La Nava de Fuentes (Palencia), a rehabilitated wetland intended to restore just part of the ancient Laguna de La Nava, one of the biggest wet areas in continental Spain until its virtual disappearance in the 1950s. The ecology and the phenology of 50 species are described, and discussed to determine the degree of maturity reached by this newly created wetland. © Gauthier-Villars, 1994.