Historical and contemporary factors generate unique butterfly communities on islands

Vodă, Raluca et al.The mechanisms shaping island biotas are not yet well understood mostly because of a lack of studies comparing eco-evolutionary fingerprints over entire taxonomic groups. Here, we linked community structure (richness, frequency and nestedness) and genetic differentiation (based on mitochondrial DNA) in order to compare insular butterfly communities occurring over a key intercontinental area in the Mediterranean (Italy-Sicily-Maghreb). We found that community characteristics and genetic structure were influenced by a combination of contemporary and historical factors, and among the latter, connection during the Pleistocene had an important impact. We showed that species can be divided into two groups with radically different properties: widespread taxa had high dispersal capacity, a nested pattern of occurrence, and displayed little genetic structure, while rare species were mainly characterized by low dispersal, high turnover and genetically differentiated populations. These results offer an unprecedented view of the distinctive butterfly communities and of the main processes determining them on each studied island and highlight the importance of assessing the phylogeographic value of populations for conservation.This research was supported by the Spanish Ministerio de Economía y Competitividad (Project CGL2013-48277-P) and from the European Union’s Seventh Framework programme for research and innovation under the Marie Skłodowska-Curie grant agreement No 609402 - 2020 researchers: Train to Move (T2M) postdoctoral fellowship to R. Vodă, and by the projects “Barcoding Italian Butterflies” and “Barcoding Butterflies of the Tuscan Archipelago National Park.” V. Dincă was supported by a Marie Curie International Outgoing Fellowship within the 7th European Community Framework Programme (project no. 625997). L. Dapporto was supported by European Union’s Horizon 2020 research and innovation programme under the Marie Sklodowska-Curie grant (project no. 658844 Eco-PhyloGeo).Peer Reviewe

As(V) adsorption using MFe2O4 (M=Cd2+, Ni2+) ferrite nanoparticles

The paper present the possibilities of arsenic removal from aqueous solutions using as adsorbent MFe2O4 (M=Cd2+, Ni2+) ferrite nanoparticles due to the affinity of arsenic towards iron ions. The ferrites were obtained after a heating treatment of the cadmium respectively nickel ferrioxalate coordination compounds, as precursors, at 500oC. From the two studied adsorbent material the nickel ferrite developed a higher maximum adsorption capacity (132 µg As(V)/g of ferrite) than the cadmium ferrite (109 µg As(V)/g of ferrite) in the removal process of As(V) from aqueous solutions. In both cases the equilibrium between the adsorbent and adsorbate was achieved in 60 minutes

Biodiversity and comparative phylogeography of western Mediterranean butterflies

Esta tesis doctoral se basa en un enfoque macroecológico y en análisis de grandes conjuntos de datos para el estudio de la biodiversidad y filogeografía de las mariposas mediterráneas occidentales. Una de las contribuciones más importantes y originales de esta tesis doctoral fue la creación de una extensa colección de ADN y tejidos de mariposas del Mediterráneo occidental, así como la reunión de datos de presencia sobre las comunidades de las islas y del continente. Actualmente en Europa no existen conjuntos de datos similares disponibles para grupos taxonómicos enteros con tan amplia distribución y con una resolución espacial comparable. En el primer capítulo investigamos los patrones de la beta-diversidad de las mariposas del Mediterráneo occidental y proporcionamos nueva evidencia de la efectividad de diferentes medidas para la beta diversidad. Se demuestra que ningún índice por sí mismo puede recuperar amplios patrones biogeográficos simultáneamente para las islas y el continente. Por lo tanto, se propone comparar y combinar los resultados obtenidos usando los dos tipos de índices, particionados y no particionados, para obtener conclusiones válidas. En el segundo capítulo definimos objetivamente las especies crípticas y producimos una lista de grupos crípticos para todas las mariposas del Mediterráneo occidental. Se demuestra que los patrones de distribución 'chequered' dentro de los grupos crípticos son mucho más frecuentes que entre especies no crípticas congéneres y que preponderantemente se producen entre las especies no hermanas con una divergencia genética sustancial. Estos patrones de distribución son un fenómeno general de las especies crípticas de mariposas en el Mediterráneo occidental pero también podría ser el caso para otros organismos. En el tercer capítulo investigamos las causas potenciales que pueden producir patrones de distribución 'chequered', utilizando un enfoque multidisciplinario para dos pares de especies crípticas. Se demuestra que éste es un fenómeno multifacético que no puede ser explicado por hipótesis simples y establecemos algunos de los factores clave, incluyendo las interacciones entre especies, que suelen ser olvidadas en ecología dada la dificultad de evaluar su importancia. En el cuarto capítulo, proporcionamos un nuevo método para comparar directamente los diferentes tipos de marcadores, incluso si los datos disponibles para cada marcador sólo en parte se superponen, y discutimos las implicaciones biogeográficas de las concordancias y discrepancias observadas. Una de las aplicaciones más importantes de este método es que puede ser utilizado para una amplia variedad de marcadores y taxones. En el quinto y último capítulo se investigan los mecanismos que determinan y mantienen la comunidad de mariposas de las islas circum-sicilianas. Se demuestra que las poblaciones insulares tienen historias muy diferentes y presentan diferentes probabilidades de recolonización tras extinciones locales. Estos resultados tienen grandes implicaciones para centrar los esfuerzos de conservación en ciertas islas. Los resultados de esta tesis doctoral contribuyen a una mejor comprensión de la diversidad de mariposas en el Mediterráneo occidental y también proporcionan un marco para futuros estudios que investiguen no sólo las mariposas, si no también otros organismos. El trabajo realizado durante esta tesis doctoral aporta nuevos datos para la investigación (una completa colección de especímenes, datos de presencia y secuencias de ADN), herramientas originales para filogeografía comparada (algoritmos mejorados, funciones R), resultados descriptivos (mapas zoogeográficos, patrones filogeográficos) y aportaciones conceptuales (exclusión mutua, las propiedades únicas de la biodiversidad críptica, las islas como individuos, los procesos que determinan las comunidades insulares i el valor filogeográfico de las poblaciones para la priorización de la conservación)This PhD thesis consists in a comprehensive macroecological approach and in analyses of large datasets to study the biodiversity and phylogeography of western Mediterranean butterflies. One of the most important and original contributions of this PhD was creating a comprehensive butterfly DNA and tissues collection for the western Mediterranean and improving the occurrence data for island and mainland communities. Currently in Europe there are no other similar datasets available for an entire taxonomic group with such a wide distribution and with a comparable spatial resolution. In the first chapter we investigate patterns of the butterfly beta-diversity in the western Mediterranean and provide new evidence for the effectiveness of different measures of beta diversity. We demonstrate that no index by itself is able to retrieve comprehensive biogeographical patterns simultaneously for islands and mainland and the results should be compared and combined by using both unpartitioned and partitioned indices to obtain comprehensive results. In the second chapter we objectively define cryptic species and produce a list of cryptic groups for all the butterflies in the western Mediterranean. We show that chequered patterns of distribution within cryptic groups are much more frequent than among congeneric non-cryptic species and they preponderantly occur between non-sister species with substantial genetic divergence. These patterns of distribution are a general phenomenon for cryptic butterfly taxa in the western Mediterranean as could also be the case for other organisms. In the third chapter we investigate the potential causes producing such chequered distribution patterns by using a multidisciplinary approach for two pairs of cryptic species. We show that this is a multifaceted phenomenon that cannot be explained by simple hypotheses and we pinpoint some of the key players, including species interactions, which are usually forgotten in ecology given the difficulty to assess their importance. In the fourth chapter we provide a new method to directly compare different types of markers even if data available for each marker only partially overlap and discuss the biogeographic implications of the observed concordances and discrepancies. One of the most important applications of this method is that it can be used for a wide array of markers and taxa. In the fifth and last chapter we investigate the mechanisms determining and maintaining the butterfly community that occurs on the circum-Sicilian islands. We show that island populations have very different histories and are subjected to different recolonization probabilities following local extinctions. These findings have great implications for focusing conservation efforts on particular islands. The results of this PhD thesis contribute to a better understanding of the butterfly diversity in the western Mediterranean and they also provide a framework for future studies that investigate not only butterflies, but other organisms as well. The work performed during this PhD brings novel data for research (a comprehensive collection of specimens, presence data and DNA sequences), original tools for comparative phylogeography (improved algorithms, R functions), descriptive results (zoogeographic maps, phylogeographic patterns) and conceptual contributions (mutual exclusion, unique properties of cryptic biodiversity, islands as individuals, processes that determine island communities, the phylogeographic value of populations for conservation prioritization)

The atlas of mitochondrial genetic diversity for Western Palaearctic butterflies

Motivation Butterflies represent a model in biology and a flagship group for invertebrate conservation. We provide four new resources for the Western Palaearctic butterflies: (1) an updated checklist comprising 552 species; (2) a curated dataset of 32,126 mitochondrial cytochrome c oxidase subunit I (COI) sequences for 532 species, including a de novo reference library for the Maghreb (Morocco, northern Algeria and Tunisia) and Macaronesia (Azores, Madeira and Canary Islands); (3) seven indexes of intraspecific genetic variation (IGV): observed and expected number of haplotypes, haplotype and nucleotide diversity, two fixation indexes and maximum p-distance; and (4) species-level maps illustrating the distribution of COI variability and haplotype networks. The updated checklist will be fundamental for any application dealing with butterfly diversity in the Western Palaearctic. The IGV indexes provide measures for genetic polymorphism and spatial structure and represent proxies for dispersal capacity. These resources will facilitate comparative studies of macrogenetics, foster integrative taxonomy and aid conservation strategies. Main types of variables contained A complete species checklist in table format, 32,126 mitochondrial DNA barcodes provided with metadata (species membership, WGS84 coordinates and sequence length) and a book in PDF format, including the IGV atlas and indexes, are provided. Spatial location and grain The checklist encompasses Europe up to the Urals in the east, north Macaronesia (the Azores, Madeira and the Canary Islands) and the Maghreb (Morocco, northern Algeria and Tunisia). COI sequences have been retained in the geographical interval of -31.3 to 67.5° of longitude and 27.5 - 71.2° of latitude. Time period and grain COI sequences originate from studies published between 1998 and 2022 and from de novo sequencing of 2541 specimens done between 2007 and 2022. Major taxa and level of measurement Butterflies (Lepidoptera: Papilionoidea), analysed from individual to species level. Software format Data and functions to manage the dataset are provided in the iodatabase R package (https://github.com/leondap/iodatabase) and in Dryad (https://doi.org/10.5061/dryad.9w0vt4bjj).Support for this research was provided by the Academy of Finland (Academy Research Fellow, decision no. 328895) and by a Marie Curie International Outgoing Fellowship within the 7th European Community Framework Programme (project no. 625997) to V.D., by “la Caixa” Foundation (ID 100010434) to M.M. (grant LCF/BQ/DR20/11790020), and by projects CGL2010-21226/BOS and CGL2013-48277-P (Spanish Ministerio de Economía y Competitividad), CGL2016-76322 (AEI/FEDER, UE), PID2019-107078GB-I00 (MCIN/AEI/10.13039/501100011033) and 2017-SGR-991 (Generalitat de Catalunya) to R.Vi., the grant BES-2017-080641 funded by MCIN/AEI/10.13039/501100011033 and by “ESF Investing in your future” to J.C.H., the project PID2020-117739GA-I00 (MCIN/AEI/10.13039/501100011033) to G.T., and by the Direttiva Biodiversità 2019 and 2020 projects (Ministero della Transizione Ecologica) to L.D.1 INTRODUCTION 2 METHODS 2.1 Checklists and geographical ranges 2.2 Data acquisition, curation and quality control 2.3 Indexes of genetic variation 2.4 Maps of genetic variation 2.5 Haplotype networks 2.6 Script availability 3 RESULTS AND DISCUSSION ACKNOWLEDGMENTS FUNDING INFORMATION CONFLICT OF INTEREST BIOSKETC

Integrating three comprehensive datasets shows that mitochondrial DNA variation is linked to species traits and paleogeographic events in European butterflies

Understanding the dynamics of biodiversity, including the spatial distribution of genetic diversity, is critical for predicting responses to environmental changes, as well as for effective conservation measures. This task requires tracking changes in biodiversity at large spatial scales and correlating with species functional traits. We provide three comprehensive resources to understand the determinants for mitochondrial DNA differentiation represented by i) 15,609 COI sequences and ii) 14 traits belonging to 307 butterfly species occurring in Western‐Central Europe and iii) the first multi‐locus phylogenetic tree of all European butterfly species. By applying phylogenetic regressions we show that mitochondrial DNA spatial differentiation (as measured with Gst, G'st, D and Dst) is negatively correlated with species traits determining dispersal capability and colonization ability. Thanks to the high spatial resolution of the COI data, we also provide the first zoogeographic regionalization maps based on intraspecific genetic variation. The overall pattern obtained by averaging the spatial differentiation of all Western‐Central European butterflies shows that the paradigm of long‐term glacial isolation followed by rapid pulses of post‐glacial expansion has been a pervasive phenomenon in European butterflies. The results and the extensive datasets we provide here constitute the basis for genetically‐informed conservation plans for a charismatic group in a continent where flying insects are under alarming decline

Butterfly-parasitoid-hostplant interactions in Western Palaearctic Hesperiidae: a DNA barcoding reference library

The study of ecological interactions between plants, phytophagous insects and their natural enemies is an essential but challenging component for understanding ecosystem dynamics. Molecular methods such as DNA barcoding can help elucidate these interactions. In this study, we employed DNA barcoding to establish hostplant and parasitoid interactions with hesperiid butterflies, using a complete reference library for Hesperiidae of continental Europe and north-western Africa (53 species, 100% of those recorded) based on 2934 sequences from 38 countries. A total of 233 hostplant and parasitoid interactions are presented, some recovered by DNA barcoding larval remains or parasitoid cocoons. Combining DNA barcode results with other lines of evidence allowed 94% species-level identification for Hesperiidae, but success was lower for parasitoids, in part due to unresolved taxonomy. Potential cases of cryptic diversity, both in Hesperiidae and Microgastrinae, are discussed. We briefly analyse the resulting interaction networks. Future DNA barcoding initiatives in this region should focus attention on north-western Africa and on parasitoids, because in these cases barcode reference libraries and taxonomy are less well developed.Support for this research was provided by the Spanish National Research Council (CSIC) with a JAE-Intro fellowship for the introduction to research to ETD (reference numbers JAEINT_20_00248 and JAEINT20_EX_0638) and by projects PID2019-107078GB-I00/MCIN/AEI/10.13039/501100011033 and 2017-SGR-991 (Generalitat de Catalunya) to RV, and PID2020-117739GA-I00/MCIN/AEI/10.13039/501100011033 to GT. We thank the Rachadaphiseksomphot Fund, Graduate School, Chulalongkorn University, for the award of a Senior Postdoctoral Fellowship to DLJQ. Further support for this research was provided by the Academy of Finland (Academy Research Fellow, decision no. 328895) to VD. PDNH acknowledges support from Genome Canada through Ontario Genomics. BV has been funded by the CERCA Programme of the Generalitat de Catalunya and by the Grant RYC-22243-2017, whose PI is Josep Sardanyés. SV was supported by the Spanish Ministry of Economy and Competitiveness, grant PID2020-117822GB-I00 MINEICO/AEI/ FEDER and the European Union.INTRODUCTION MATERIAL AND METHODS RESULTS DISCUSSION CONCLUSION SUPPORTING INFORMATION ACKNOWLEDGEMENTS DATA AVAILABILITY REFERENCES Supplementary dat

Biodiversity and comparative phylogeography of western Mediterranean butterflies

Esta tesis doctoral se basa en un enfoque macroecológico y en análisis de grandes conjuntos de datos para el estudio de la biodiversidad y filogeografía de las mariposas mediterráneas occidentales. Una de las contribuciones más importantes y originales de esta tesis doctoral fue la creación de una extensa colección de ADN y tejidos de mariposas del Mediterráneo occidental, así como la reunión de datos de presencia sobre las comunidades de las islas y del continente. Actualmente en Europa no existen conjuntos de datos similares disponibles para grupos taxonómicos enteros con tan amplia distribución y con una resolución espacial comparable. En el primer capítulo investigamos los patrones de la beta-diversidad de las mariposas del Mediterráneo occidental y proporcionamos nueva evidencia de la efectividad de diferentes medidas para la beta diversidad. Se demuestra que ningún índice por sí mismo puede recuperar amplios patrones biogeográficos simultáneamente para las islas y el continente. Por lo tanto, se propone comparar y combinar los resultados obtenidos usando los dos tipos de índices, particionados y no particionados, para obtener conclusiones válidas. En el segundo capítulo definimos objetivamente las especies crípticas y producimos una lista de grupos crípticos para todas las mariposas del Mediterráneo occidental. Se demuestra que los patrones de distribución ‘chequered’ dentro de los grupos crípticos son mucho más frecuentes que entre especies no crípticas congéneres y que preponderantemente se producen entre las especies no hermanas con una divergencia genética sustancial. Estos patrones de distribución son un fenómeno general de las especies crípticas de mariposas en el Mediterráneo occidental pero también podría ser el caso para otros organismos. En el tercer capítulo investigamos las causas potenciales que pueden producir patrones de distribución ‘chequered’, utilizando un enfoque multidisciplinario para dos pares de especies crípticas. Se demuestra que éste es un fenómeno multifacético que no puede ser explicado por hipótesis simples y establecemos algunos de los factores clave, incluyendo las interacciones entre especies, que suelen ser olvidadas en ecología dada la dificultad de evaluar su importancia. En el cuarto capítulo, proporcionamos un nuevo método para comparar directamente los diferentes tipos de marcadores, incluso si los datos disponibles para cada marcador sólo en parte se superponen, y discutimos las implicaciones biogeográficas de las concordancias y discrepancias observadas. Una de las aplicaciones más importantes de este método es que puede ser utilizado para una amplia variedad de marcadores y taxones. En el quinto y último capítulo se investigan los mecanismos que determinan y mantienen la comunidad de mariposas de las islas circum-sicilianas. Se demuestra que las poblaciones insulares tienen historias muy diferentes y presentan diferentes probabilidades de recolonización tras extinciones locales. Estos resultados tienen grandes implicaciones para centrar los esfuerzos de conservación en ciertas islas. Los resultados de esta tesis doctoral contribuyen a una mejor comprensión de la diversidad de mariposas en el Mediterráneo occidental y también proporcionan un marco para futuros estudios que investiguen no sólo las mariposas, si no también otros organismos. El trabajo realizado durante esta tesis doctoral aporta nuevos datos para la investigación (una completa colección de especímenes, datos de presencia y secuencias de ADN), herramientas originales para filogeografía comparada (algoritmos mejorados, funciones R), resultados descriptivos (mapas zoogeográficos, patrones filogeográficos) y aportaciones conceptuales (exclusión mutua, las propiedades únicas de la biodiversidad críptica, las islas como individuos, los procesos que determinan las comunidades insulares i el valor filogeográfico de las poblaciones para la priorización de la conservación).This PhD thesis consists in a comprehensive macroecological approach and in analyses of large datasets to study the biodiversity and phylogeography of western Mediterranean butterflies. One of the most important and original contributions of this PhD was creating a comprehensive butterfly DNA and tissues collection for the western Mediterranean and improving the occurrence data for island and mainland communities. Currently in Europe there are no other similar datasets available for an entire taxonomic group with such a wide distribution and with a comparable spatial resolution. In the first chapter we investigate patterns of the butterfly beta-diversity in the western Mediterranean and provide new evidence for the effectiveness of different measures of beta diversity. We demonstrate that no index by itself is able to retrieve comprehensive biogeographical patterns simultaneously for islands and mainland and the results should be compared and combined by using both unpartitioned and partitioned indices to obtain comprehensive results. In the second chapter we objectively define cryptic species and produce a list of cryptic groups for all the butterflies in the western Mediterranean. We show that chequered patterns of distribution within cryptic groups are much more frequent than among congeneric non-cryptic species and they preponderantly occur between non-sister species with substantial genetic divergence. These patterns of distribution are a general phenomenon for cryptic butterfly taxa in the western Mediterranean as could also be the case for other organisms. In the third chapter we investigate the potential causes producing such chequered distribution patterns by using a multidisciplinary approach for two pairs of cryptic species. We show that this is a multifaceted phenomenon that cannot be explained by simple hypotheses and we pinpoint some of the key players, including species interactions, which are usually forgotten in ecology given the difficulty to assess their importance. In the fourth chapter we provide a new method to directly compare different types of markers even if data available for each marker only partially overlap and discuss the biogeographic implications of the observed concordances and discrepancies. One of the most important applications of this method is that it can be used for a wide array of markers and taxa. In the fifth and last chapter we investigate the mechanisms determining and maintaining the butterfly community that occurs on the circum-Sicilian islands. We show that island populations have very different histories and are subjected to different recolonization probabilities following local extinctions. These findings have great implications for focusing conservation efforts on particular islands. The results of this PhD thesis contribute to a better understanding of the butterfly diversity in the western Mediterranean and they also provide a framework for future studies that investigate not only butterflies, but other organisms as well. The work performed during this PhD brings novel data for research (a comprehensive collection of specimens, presence data and DNA sequences), original tools for comparative phylogeography (improved algorithms, R functions), descriptive results (zoogeographic maps, phylogeographic patterns) and conceptual contributions (mutual exclusion, unique properties of cryptic biodiversity, islands as individuals, processes that determine island communities, the phylogeographic value of populations for conservation prioritization)

Mutual exclusion between cryptic species generates most butterfly beta-diversity

Trabajo presentado en el XVIII European Congress of Lepidopterology, celebrado en Blagoevgrad (Bulgaria) del 29 de julio al 4 de agosto de 2013.The cryptic fraction of biodiversity is composed by morphologically similar spe - cies that are or have been overlooked by scientists. Most research is now directed towards documenting such cases, and estimating what will be the impact of this hid- den fraction on the monumental task of describing Earth’s biodiversity. We address for the first time the question whether the cryptic fraction of biodiversity displays particular properties beyond the fact that it is composed of morphologically similar species. By using the butterfly fauna of the Western Mediterranean we demonstrate that the cryptic fraction of diversity is overwhelmingly composed by groups of spe - cies that are not sympatric. Using two model cryptic species pairs we show that nei- ther dispersal capacity, nor climatic factors seem to explain the observed chequered pattern of distributions, and that the existence of species interactions resulting in mutual exclusion is the most likely hypothesis. Finally, we demonstrate that mutual exclusion between cryptic species on islands accounts for most beta-diversity turn- over. Thus, we show that the inclusion of cryptic diversity, frequently excluded in large-scale surveys and studies because of intrinsic identification difficulties, is of capital importance.N

Integrating distributions, community structure and genetic data to understand the biogeography of west-Mediterranean butterflies

Dapporto, Leonardo et al.-- Trabajo presentado en el XXV Congresso Nazionale Italiano de Entomologia, celebrado en Padua del 20 al 24 de junio de 2016.It is widely recognized that community characteristics and genetic structure of island biotas are determined by a interplay between contemporary and historical physi cal factors and by differences in ecological traits among species. Nevertheless, the mechanisms shaping island biotas are not yet well understood mostly, because of a lack of studies comparing eco-evolutionary fingerprints over entire taxonomic groups. In the last years, our research group integrated data about community structure (richness, frequency and nestedness), genetic differentiation (based on mitochondrial DNA) and species traits (degree of generalism, dispersal ability and climatic preferences) of butterfly communities occurring over the western Mediterranean region, a key biodiversity hot-spot located at the interface between two continents. An outstanding result emerging from these studies is the evidence that biogeographic dynamics can change butterfly communities in short time frames (thousands or hundreds of years), generating unexpected contrasts between insular and mainland populations. Understanding which ecological factors promote and reduce species colonization and survival is thus crucial in order to preserve such a patterned and peculiar insular biodiversity. Our integrated approach recently allowed us to tie apart and compare the relative relevance of several species traits in explaining species occurrence and genetic diversification in two butterfly-rich and highly diverse areas (Tuscan Archipelago and South Italy). We here show that climatic preferences play a dominant influence, with species living in drier climate showing high genetic structure and those preferring cooler and wetter climate facing the highest extinction risk. This suggests that environmental changes might selectively erase different fractions of the existing island diversity. Our approach provides an example of how series of comprehensive analyses on a wide area and la rge taxonomic groups can test rarely assessed biogeographic principles like the links of genetic structure with dispersal tendency and frequency on islands, or the relative effects of contemporary and historical determinants on island populations. The challenge of integrating community ecology and phylogeographic approaches can also provide the baseline information for developing conservation strategies that maximize biodiversity at both the species and intraspecific genetic levels.N