Volume 35 - No. 1 March 2014 BRILL Amphibia-Reptilia Amphibia-Reptilia Volume 35 - No. 1 - 1-148 - 2014 Publication of the Societas Europaea Herpetologica Contents Neftali Sillero, João Campos, Anna Bonardi, Claudia Corti, Raymond Creemers, Pierre- 1 Andre Crochet, Jelka Crnobrnja Isailovic,´ Mathieu Denoël, Gentile Francesco Ficetola, João Gonçalves, Sergei Kuzmin, Petros Lymberakis, Philip de Pous, Ariel Rodríguez, Roberto Sindaco, Jeroen Speybroeck, Bert Toxopeus, David R. Vieites and Miguel Vences Updated distribution and biogeography of amphibians and reptiles of Europe

Neftalí Sillero, Marco Amaro Oliveira, Pedro Sousa, Fátima Sousa and Luís Gonçalves-Seco 33 Distributed database system of the New Atlas of Amphibians and Reptiles in Europe: the NA2RE project

Toby Keswick and Margaretha Hofmeyr 41 VOLUME 35 - NO. 1 - 2014 - PP. 1-148 Refuge characteristics and preferences of Psammobates oculifer in semi-arid savanna

Neus Oromi, Fèlix Amat, Delfi Sanuy and Salvador Carranza 53 Life history trait differences between a lake and a stream-dwelling population of the Pyrenean brook newt (Calotriton asper)

Jérémie H. Cornuau, Dirk S. Schmeller, Romain Pigeault and Adeline Loyau 63 Resistance of morphological and behavioral sexual traits of the palmate newt (Lissotriton helveticus) to bacterial lipopolysaccharide treatment

Carlos Carreras, Catalina Monzón-Argüello, Luis Felipe López-Jurado, Pascual Calabuig, 73 Juan Jesús Bellido, Juan José Castillo, Pablo Sánchez, Pas Medina, Jesús Tomás, Patricia Gozalbes, Gloria Fernández, Adolfo Marco and Luis Cardona Origin and dispersal routes of foreign green and Kemp’s ridley turtles in Spanish Atlantic and Mediterranean waters

James C. Cureton II, Michael Janis, William I. Lutterschmidt, Christopher P. Randle, Donald 87 C. Ruthven III and Raelynn Deaton Effects of urbanization on genetic diversity, gene flow, and population structure in the ornate box turtle (Terrapene ornata)

Giulia Tognarelli, Marco A.L. Zuffi, Silvia Marracci and Matilde Ragghianti 99 Surveys on populations of green frogs (Pelophylax) of Western Tuscany sites with molecular and morphometric methods

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ISSN 0173-5373 E-ISSN 1568-5381 brill.com/amre Amphibia-Reptilia 35 (2014): 1-31

Updated distribution and biogeography of amphibians and reptiles of Europe

∗ Neftali Sillero1, , João Campos1, Anna Bonardi2, Claudia Corti3, Raymond Creemers4, Pierre-Andre Crochet5, Jelka Crnobrnja Isailovic´6,7, Mathieu Denoël8, Gentile Francesco Ficetola2, João Gonçalves9, Sergei Kuzmin10, Petros Lymberakis11, Philip de Pous12,13, Ariel Rodríguez14, Roberto Sindaco15, Jeroen Speybroeck16, Bert Toxopeus17, David R. Vieites18,19, Miguel Vences14

Abstract. A precise knowledge of the spatial distribution of taxa is essential for decision-making processes in land management and biodiversity conservation, both for present and under future global change scenarios. This is a key base for several scientifi disciplines (e.g. macro-ecology, biogeography, evolutionary biology, spatial planning, or environmental impact assessment) that rely on species distribution maps. An atlas summarizing the distribution of European amphibians and reptiles with 50 × 50 km resolution maps based on ca. 85 000 grid records was published by the Societas Europaea Herpetologica (SEH) in 1997. Since then, more detailed species distribution maps covering large parts of Europe became available, while taxonomic progress has led to a plethora of taxonomic changes including new species descriptions. To account for these progresses, we compiled information from different data sources: published in books and websites, ongoing national atlases, personal data kindly provided to the SEH, the 1997 European Atlas, and the Global Biodiversity Information Facility (GBIF). Databases were homogenised, deleting all information except species names and coordinates, projected to the same coordinate system (WGS84) and transformed into a 50 × 50 km grid. The newly compiled database comprises more than 384 000 grid and locality records distributed across 40 countries. We calculated species richness maps as well as maps of Corrected Weighted Endemism and define species distribution types (i.e. groups of species with similar distribution patterns) by hierarchical cluster analysis using Jaccard’s index as association measure. Our analysis serves as a preliminary step towards an interactive, dynamic and online distributed database system (NA2RE system) of the current spatial distribution of European amphibians and reptiles. The NA2RE system will serve as well to monitor potential temporal changes in their distributions. Grid maps of all species are made available along with this paper as a tool for decision-making and conservation-related studies and actions. We also identify taxonomic and geographic gaps of knowledge that need to be filled and we highlight the need to add temporal and altitudinal data for all records, to allow tracking potential species distribution changes as well as detailed modelling of the impacts of land use and climate change on European amphibians and reptiles.

Keywords: biogeography, conservation, distribution atlas, distribution types, endemism, European herpetofauna, IUCN red list, species richness.

1 - Centro de Investigação em Ciências Geo-Espaciais, 7 - Department of Evolutionary Biology, Institute for Bio- Alameda do Monte da Virgem, 4430-146 Vila Nova de logical Research “Siniša Stankovic”,´ University of Bel- Gaia, Portugal grade, Despota Stefana 142, 11000 Beograd, Serbia 2 - Department of Earth and Environmental Sciences, Uni- 8 - F.R.S. – FNRS Research Associate, Behavioural Biol- versità di Milano-Bicocca, Piazza della Scienza 1, ogy Unit, University of Liège, 22 Quai van Beneden, 20126 Milano, Italy 4020 Liege, Belgium 3 - Museo di Storia Naturale dell’Università di Firenze, 9 - CIBIO, University of Porto, R. Campo Alegre 687, Sezione di Zoologia “La Specola”, Via Romana 17, 4169-007 Porto, Portugal 50125 Firenze, Italia 10 - Institute of Ecology and Evolution, Russian Academy 4 - RAVON, Postbus 1413, 6501 BK Nijmegen, The of Sciences, Moscow 117071, Russia Netherlands 11 - Natural History Museum of Crete, University of Crete, 5 - CNRS-UMR5175 CEFE, Centre d’Ecologie Fonction- Knossou Ave., P.O. Box 2208, 71409 Heraklion Crete, nelle et Evolutive, 1919, route de Mende, 34293 Mont- Greece pellier, France 12 - Faculty of Life Sciences and Engineering, Universitat 6 - Department of Biology and Ecology, Faculty of Sci- de Lleida, Av. Rovira Roura 191, 25198 Lleida, Spain ences and Mathematics, University of Niš, Višegradska 13 - Institut de Biologia Evolutiva (CSIC-Universitat Pom- 33, 18000 Niš, Serbia peu Fabra), Animal Phylogeny and Systematics,

© Societas Europaea Herpetologica, 2014 DOI:10.1163/15685381-00002935 This is an open access article distributed under the terms of the Creative Commons Attribution-Noncommercial 3.0 Unported (CC-BY-NC 3.0) License. 2 N. Sillero et al.

Introduction In European herpetology, shortly after the Societas Europaea Herpetologica (SEH) was A good knowledge on the geographical dis- established in 1979, it became evident that a tribution of organisms is pivotal for macro- comprehensive assessment of the distribution ecological and evolutionary studies, as well as of all European amphibians and reptiles should to inform policy makers in decisions on land receive priority, as basic maps where lack- management, health, climate change and biodi- ing. A mapping committee of the SEH was versity conservation (Jetz, McPherson and Gu- established in 1983, coordinated by a team ralnick, 2011). The availability of reliable maps based at the Muséum National d’Histoire Na- that depict the historical and current distri- turelle in Paris. From the work of regional bution of species therefore constitutes an im- and national coordinators, more than 85 000 portant component in conservation-related re- grid records were collected and shown in maps search. Data on their extent of occurrence are of 50 × 50 km resolution produced by the crucial for assigning IUCN threat categories to Service du Patrimoine Naturel (Paris, France). species (IUCN, 2001). This has for instance This resulted in a distribution atlas published been a strategy in the Global Amphibian As- in 1997 (Gasc et al., 1997). This work, which sessment (Stuart et al., 2004) which provided in the following will for brevity be referred the f rst comprehensive estimate of threat cat- to as ‘the 1997 European Atlas’, has subse- egories and distribution ranges of amphibians quently provided the basis for numerous stud- worldwide, a taxon that constitutes an impor- ies, such as several conservation-oriented mod- tant model group in conservation biology (e.g. elling approaches (e.g. Araújo and Pearson, Hopkins, 2007). Furthermore, many amphibian 2005; Araújo et al., 2005; Araújo, Thuiller and species and at least some groups of reptiles are Pearson, 2006; Araújo et al., 2008). undergoing severe global declines (Wake and After the publication of the 1997 Euro- Vredenburgh, 2008; Sinervo et al., 2010; Böhm pean Atlas, there has been a high intensity of et al., 2013), making their conservation a prime mapping efforts and related research in Eu- challenge and gathering data on their current rope. Numerous regional and national soci- distribution a top research priority. eties have since then produced detailed am- phibian and reptile distributional information covering large parts of Europe, more detailed Passeig Marítim de la Barceloneta 37-49, 08003 Barcelona, Spain and reliable than the 1997 European Atlas. 14 - Technische Universität Braunschweig, Division of Many of these were published in the form of Evolutionary Biology, Zoological Institute, Mendel- regional or national atlases (e.g. Bitz et al., ssohnstr. 4, 38108 Braunschweig, Germany 1996; Günther, 1996; Pleguezuelos, 1997; Ca- 15 - c/o Museo Civico di Storia Naturale, via San Francesco bela, Grillitsch and Tiedemann, 2001; Hofer, di Sales 88, 10022 Carmagnola (TO), Italia 16 - Research Institute for Nature and Forest, Kliniekstraat Monney and Dušej, 2001; Pleguezuelos, Lizana 25, 1070 Brussels, Belgium and Márquez, 2002; Głowacinski´ and Raf nski,´ 17 - University of Twente, Faculty of Geo-Information Sci- 2003; Puky, Schad and Szövenyi, 2006; Sin- ence and Earth Observation (ITC), P.O. Box 217, 7500 daco et al., 2006; Jacob et al., 2007; Lanza et AA Enschede, The Netherlands 18 - Museo Nacional de Ciencias Naturales and Consejo al., 2007; Laufer, Klemens and Sowig, 2007; Superior de Investigaciones Científicas c/José Gutier- Proess, 2007; Creemers and van Delft, 2009; rez Abascal 2, 28006 Madrid, Spain Corti et al., 2010; Loureiro et al., 2010). Some 19 - REFER Biodiversity Chair, University of Porto, of them (e.g. UK, Netherlands, Wallonia, Flan- CIBIO, Campus Agrário de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal ders, Switzerland) were published also through ∗ Corresponding author; publicly available internet resources. Others, e-mail: [email protected] like the atlas of Sweden, were published exclu- New atlas of European amphibians and reptiles 3 sively on the internet. This wealth of novel data art, 2006), it is an urgent task to make updated claims for an update of the herpetofaunal distri- distributional information on these organisms bution data also at the European level, to quan- available. The species distribution maps of the tify Europe-wide the improvement in knowl- 1997 European Atlas (Gasc et al., 1997) have edge since the previous Atlas, as well as a f rst never been made available in GIS format. How- step towards tracking potential changes in the ever useful and original at the time, they are now distribution of the European herpetofauna in the outdated due to the considerable accumulation context of global change. of new distribution data, and especially because Novel technologies for mapping species dis- of the taxonomic progress that resulted in mul- tributions currently available, such as newly de- tiple changes of genus-level classification and a veloped Geographic Information Systems (Lon- large number of new species descriptions (Spey- gley et al., 2010) and their extensions, offer the broeck, Beukema and Crochet, 2010; Vences possibility of establishing extensive databases et al., 2013). This new taxonomy resulted in of distribution records, with associated meta- many species being split into multiple entities data such as voucher specimen lists or photos. for which the exact distribution limits are poorly Citizen-science online tools allow contributors known. entering their observations, and directly link The goal of the present study is to provoke them to analysis tools such as spatial modelling and facilitate fillin of these gaps by making or the production of customised maps. The cur- updated distribution maps for the European her- rent Mapping Committee of the SEH (estab- petofauna available. For this purpose, we have lished in 2006), together with the SEH Council compiled information from a large number of and some associated fellows, has acknowledged published and partly unpublished mapping ef- that distribution atlases should be conceived as forts at a variety of spatial scales and trans- dynamic tools, implemented in a way that al- formed those data into a 50 × 50 km UTM lows for continuous updates, extension changes, grid, similar to the one used for the 1997 Eu- and customised data extraction while respect- ropean Atlas. Based on this new compilation ing the copyright that particular organisations or of maps, all of which are made available (see individuals might hold on parts of the underly- online Supplementary Atlas S1-S5 online), we ing data. The goal is to establish a Spatial Data here (1) identify the major spatial and taxo- Infrastructure, a system of geographically dis- nomic gaps in the currently available knowledge tributed systems, where the original data remain in order to identify future research priorities, on the servers controlled by national or regional and (2) analyse patterns of species richness, en- herpetological societies, and through an online demism and main distribution types (i.e. groups network it is possible to make data queries via of species with similar distribution patterns) for the SEH portal (Sillero et al., 2014; see http:// European amphibians and reptiles. na2re.ismai.pt). For countries that do not have national databases, the SEH works on establish- Materials and methods ing a connected database linked to an internet Study area portal for data collection. A dynamic online atlas of European amphib- This compilation included almost the same area as the 1997 ians and reptiles based on an underlying dis- European Atlas (Gasc et al., 1997). We used the limits for Europe (see Supplementary fig S1 online) provided by Geo- tributed database of distribution records repre- commons (http://geocommons.com/overlays/76975). The sents a major logistic challenge and is time- geographical limits of the previous SEH 1997 European at- consuming. However, considering the current las were those define by Mertens and Wermuth (1960), covering parts or the whole of 45 countries. Partial territo- conservation crisis faced by many European ries included were: north-western tip of Turkey (European amphibians and reptiles (Cox, Chanson and Stu- Turkey), territories in the Russian Federation west of the 4 N. Sillero et al.

Urals, north-eastern tip of Azerbaijan, north-western tip of Database compilation Kazakhstan, Greece minus the Sporades Islands. However, the Geocommons limits do not include parts of Azerbaijan Our goal in compiling updated distribution maps for the Eu- and Kazakhstan, while the Ural limits are define more pre- ropean fauna was to cover as many European countries as possible with national atlas data or new personal records. cisely. These limits for Europe are widely accepted by many The species data included in these updated maps were ob- geographical atlases (e.g. Cheers, 2005). tained from different data sources, namely (1) published (in books or websites) or on-going national atlases, (2) per- Taxa sonal data kindly provided to the SEH, (3) the 1997 Euro- pean Atlas, and (4) the Global Information Facility (GBIF: For historical consistency and to facilitate reading, in this www.gbif.org). Because the GBIF data originate from many paper we use the traditional term ‘reptiles’ for the para- different data sources and contain numerous errors and phyletic group including the vertebrate orders Squamata, discrepancies, we tried to minimise their use as explained Testudines, Crocodylia, and Rhynchocephalia, i.e. Saurop- below. However, a few of the national atlas data were di- sida excluding birds (of which only Squamata and Tes- rectly available only from GBIF (e.g. Denmark and Nor- tudines are represented in Europe’s extant fauna). The way) and in these cases, the data were labelled as National species-level taxa considered in this compilation were de- Atlas Data rather than as GBIF data. Some countries pro- termined by the SEH, using Speybroeck, Beukema and Cro- vided databases used in already published atlases (whole chet (2010) as starting point (see Supplementary Text S1 on- database with temporal data series: e.g. Spain and Portu- line). In numerous cases, although the species status of two gal; simplifie database: e.g. The Netherlands) or before or more related taxa is undisputed, we were unable to assign publishing as an atlas (e.g. Slovenia and France). For other all available records to a species. This was either because the countries, we digitised the data from published books (e.g. original databases had been compiled following an outdated Hungary). We also included large unpublished databases for taxonomy, or because many records could not be identifie several countries compiled by some co-authors of this study up to species level in the fiel (such as for instance, Tritu- (e.g. S.L. Kuzmin, P. de Pous). In the case of territories of rus marmoratus and T. pygmaeus in the Iberian Peninsula). former Yugoslavia, J. Crnobrnja Isailovic´ and collaborators In these cases, we merged the respective species into a sin- provided some of the original data used in the 1997 Eu- gle entry in our database, which therefore in several cases ropean Atlas. National atlases and personal databases were represents a simplificatio of current taxonomy. subsequently merged in one database, which in the follow- The sampling effort was obviously not homogeneous ing will be referred to as COUNTRIES. A second database, across the whole study area. Some countries have a very hereafter named SEH/GBIF database, contained the data of good knowledge on the ranges of their species while others the 1997 European Atlas and GBIF, but only for those coun- have large gaps of chorological information. Although the tries for which no national atlas data were available. For present compilation is represented at a rather coarse scale the f nal compilation, the same exclusion strategy was also (50 × 50 km grid), gaps in the species distributions are still employed at the level of single UTM squares. Whenever a observable. Similarly, not all national and regional data sets record from the COUNTRIES database was available for a are fully consistent in their treatment of marine and intro- UTM grid (only in personal databases: e.g. S.L. Kuzmin’s personal database) we used that one rather than the dupli- duced species. Where available, our compilation includes cate record from the SEH/GBIF database. This process was terrestrial as well as marine taxa (i.e. marine turtles). Be- performed using spatial queries in ArcGIS 9.3. sides native species and populations, a number of national Many original databases contained erroneous records. data sets also included introductions, i.e. introduced species The databases were therefore reviewed and validated by from outside Europe as well as introduced populations of members of the SEH Council and its Mapping Committee European species occurring outside their natural range. In in various rounds. Erroneous records were excluded from this case our compilation is not fully consistent. For ma- the two main databases (COUNTRIES and SEH/GBIF) and rine turtles, some countries included records on sightings stored in a different file During this revision of the point (on coast and ocean) and reproduction places (i.e. Portu- locality data in the COUNTRIES and SEH/GBIF database, gal and Spain), while other countries only included repro- we furthermore f agged introduced species and species loca- duction places (i.e. Italy and Balkan countries). In general, tions, and these were transferred to a third database hereafter we did not include single records of escaped exotic species called INTRODUCED. As such, we never deleted a record: where there was no indication of naturalised populations. keeping all erroneous records rather than simply deleting For non exotics, we considered as introduced those cases them allowed tracking validation errors and makes our de- where the origin of the introduction is well known and can cisions verifiable Introduction records were define using be traced back into recent history, such as the populations of our current knowledge, which is not homogeneous, thus bias Discoglossus pictus in southern France and in Spain (Cat- may be present for some species and regions. alonia), but not those cases where ancient introductions are The three databases were composed by point records. suspected (e.g. various species on Mediterranean islands). The numerous data (table 1; 30 databases) have been re- In this sense, much of the actual herpetofaunal composition ceived in multiple digital formats, with disparate informa- in the Mediterranean is probably related to or at least influ tion and in different spatial resolutions (ranging from point enced by human activities (Corti et al., 1999). centroids of 50 × 50 km UTM grid cells to very precise New atlas of European amphibians and reptiles 5

Table 1. List of databases used in this atlas compilation. Resolution, records, and sources refer to data obtained and used for the compilation of the European atlas. References to published atlases are mentioned. Some of these databases included more than one country (e.g. S.L. Kuzmin). See table 2 for number of records per country.

Resolution Records Sources Published atlases

NATIONAL DATABASES Austria 5 × 5 km 14 136 digitised from Atlas Cabela, Grillitsch and Tiedemann, 2001 Bosnia and Herzegovina 10 × 10 km 152 provided by D. Dobrnjicand´ E. Tanovic´ Brussels 10 × 10 km 59 provided by Natagora Weiserbs and Jacob, 2005 Bulgaria 10 × 10 km 3170 digitised from website http://www.oocities.org/ herpetology_bg/ Estonia 10 × 10 km 2872 provided by Riinu Rannap Flanders 5 × 5 km 38 945 provided by Natuurpunt-Hyla Bauwens and Claus, 1996 France 50 × 50 km 11 071 provided by Service du Lescure and De Massary, Patrimoine Naturel (Muséum 2012 National d’Histoire Naturelle) Germany 10 × 10 km 31 065 digitised from Atlas Günther, 1996 Greece exact coordinates 9893 provided by P. Lymberakis Valakos et al., 2008 Hungary 10 × 10 km 13 582 digitised from Atlas Puky, Schad and Szövenyi, 2006 Italy 50 × 50 km 4292 provided by SHI (Societas Sindaco et al., 2006 Herpetologica Italica) data through R. Sindaco Luxembourg exact coordinates 10 642 provided by Musée National Proess, 2003, 2007 d’Histoire Naturelle du Luxembourg Malta 50 × 50 km 37 compiled by Claudia Corti Poland 10 × 10 km 15 502 digitised from Atlas Głowacinski´ and Raf nski,´ 2003 Portugal 10 × 10 km 17 431 provided by A. Loureiro Loureiro et al., 2010 Romania exact coordinates 5454 provided by D. Cogalniceanu˘ Cogalniceanu˘ et al., 2013a, 2013b Slovenia 10 × 10 km 3414 provided by Societas Slovenica Herpetologica Spain 10 × 10 km 68 618 provided by Sociedad Pleguezuelos, Lizana and Herpetológica Española Márquez, 2002, updated until 2005 Sweden exact coordinates 30 778 obtained from GBIF Switzerland 10 × 10 km 5705 provided by Koordinationsstelle Meyer et al., 2009 für Amphibien- und Reptilienschutz in der Schweiz (KARCH) The Netherlands 10 × 10 km 8061 provided by RAVON Creemers and van Delft, 2009 UK + Ireland 10 × 10 km 20 289 digitised from Atlas Arnold, 2005 Ukraine 10 × 10 km 1162 digitised from Atlas Kypnjehko and Bepbec, 1999 Wallonia 4 × 4 km 7269 provided by Raînne-Natagora Jacob et al., 2007

PERSONAL DATABASES J. Crnobrnja-Isailovic,´ 50 × 50 km 1128 D. Dobrnjic,´ E. Tanovic,´ Idriz Haxhiu P. de Pous Several 10 405 D. Jablonski 50 × 50 km 685  S.L. Kuzmin 1 17 865 Kuzmin, 2013 6 N. Sillero et al.

Table 1. (Continued.)

Resolution Records Sources Published atlases

CONTINENTAL DATABASES Europe 50 × 50 km 12 155 SEH Gasc et al., 1997 GBIF Several 18 772 GBIF

TOTAL 384 609

GPS point locality records). Therefore, the databases were Biogeographical analyses homogenised, deleting all other information except species × names, coordinates, and data source, and projected to the The coarse 50 50 km occurrence data were not suitable same coordinate system (WGS84). for sophisticated analyses (e.g. calculation of ecological niche models; Sillero, 2011), and these were not the main Map production goal of this compilation. We therefore did not apply any methods based on environmental niche modelling which at As an atlas is usually the representation of the species’ this level had already been carried out by Araújo, Thuiller distributions by uniform units (Sillero, Celaya and Martín- and Pearson (2006) and Araújo et al. (2008). Instead, we Alfageme, 2005; Loureiro and Sillero, 2010), record points used a number of descriptive statistics to visualise general were transformed into a grid. We used the officia UTM biogeographic patterns. Besides calculating species rich- × grid of 50 50 km, that it is freely available from the Eu- ness, we also used clustering analysis to defin chorotypes ropean Environment Agency (http://www.eea.europa.eu/). and applied a measure of regional endemism. Chorotypes This grid is based on the one used for the European Atlas of were define by Baroni-Urbani, Ruffo and Vigna Taglianti Flora, the f rst biological distribution atlas for Europe (Jalas (1978) as clusters of species with statistically similar distri- and Suonuinen, 1972). It includes 4524 land squares. There- butions for a specifi area. However, Vigna-Taglianti et al. fore, each point database (COUNTRIES, SEH/GBIF, and (1999) stated that to defin chorotypes the whole species’ INTRODUCED) was transformed to a grid file by spatially distribution should be used. In fact, Vigna-Taglianti et al. overlapping with the 50 × 50 km UTM grid. This transfor- (1999) proposed a standard classificatio of chorotypes us- mation from the point databases (e.g. GPS points, as well as centroids of grids of 1 × 1km,4× 4km,5× 5km, ing several groups of animals (e.g. beetles, amphibians, and 10 × 10 km, and 50 × 50 km squares) to a grid database reptiles). Nevertheless, the term chorotypes has been widely was performed by a set of GIS scripts for ArcGIS 9.3 (see used when applied to the herpetofauna of certain regions Supplementary table S1 online) in which for each species, (e.g. Corti et al., 1991, 1997; Olivero, Real and Márquez, each grid was assigned 0 for absence or 1 for presence. 2011; Sillero et al., 2009, and reference therein). Our in- The species maps (see example in f g. 1; all maps are tention here was not to establish a standard classificatio of provided online in Supplementary Atlases S1 and S2, and biogeographical regions for the European amphibians and the corresponding GIS f les in Supplementary Atlases S3 reptiles, but to classify species by their distribution simi- and S4; species codes are provided in Supplementary At- larity using the current available knowledge. Notwithstand- las S5) were created automatically by overlapping the three ing this, and for avoiding misunderstandings, we will use grid file (COUNTRIES, SEH/GBIF, and INTRODUCED), the term distribution type instead of chorotype, proposed by using a script written in the R language (R 2.15, R Develop- Baroni-Urbani and Collinwood (1976) and Baroni-Urbani ment Core Team, 2012). The script (included online in Sup- and Collinwood (1977). In these two works, distribution plementary Text S2) looked sequentially for each species types were calculated using incomplete species’ distribu- in the three grids, representing them with different colours. tions. The resulting maps were exported to images in .jpg format. Identificatio of the main distribution types of amphib- Species richness maps for amphibians and reptiles were cal- ians and reptiles in Europe was carried out following Sillero culated by the sum of all species present in each grid cell. et al. (2009). The merged species distribution f les (COUN- We then compared species richness maps with those based TRIES and SEH/GBIF) were transformed into two separate entirely on the original data of the 1997 European Atlas, and data matrices for amphibians and reptiles, respectively (.csv for each grid cell we subtracted the old from the new number of species occurring therein. The resulting value was subse- format) and analysed using the R 2.15 software (R Devel- quently represented on the same grid to indicate areas of opment Core Team, 2012). Distribution types were deter- increased vs. decreased quantity of recorded species. For a mined by a Hierarchical Cluster Analysis using Jaccard’s bi- better cartographical representation, all maps are shown in nary index and UPGMA as clustering method (Sillero et al., the Albers Conical projection for Europe. This projection 2009), which is a measure of similarities among species dis- (EPSG code: 9822; http://spatialreference.org/ref/sr-org/44/ tributions. This analysis was performed using the function html/) reduces cartographical distortions of Europe, by a “vegdist” of the R package “vegan” (Oksanen et al., 2012), better adjustment to the central meridian (Greenwich) and which computes the Jaccard’s index as 2B = (1 + B), both standard parallels. where B represents Bray-Curtis dissimilarity. The Bray- New atlas of European amphibians and reptiles 7

Figure 1. Example of species distribution map (Ichthyosaura alpestris) showing, in different colours, records corresponding to the COUNTRIES (red), SEH/GBIF (green) and INTRODUCED (purple) databases used in this study. Brown colours represent higher elevations. We used the officia UTM grid of 50 × 50 km from the European Environmental Agency (www.eea.europa.eu/). COUNTRIES database included data from published or on-going national atlases, and from personal data kindly provided to the SEH. SEH/GBIF included data from the 1997 European Atlas and the Global Information Facility (GBIF: www.gbif.org). We only included data from SEH/GBIF when data from COUNTRIES database were not available. Datasets for introduced species were not available in all countries.

Curtis dissimilarity is calculated as (a + b − 2j)/(a + b), (Laffan and Crisp, 2003). The sum of the weighted values where a and b are the numbers of species on compared for a given cell (weighted endemism) is then divided by squares, and j is the number of species in both squares com- the number of species occurring in the cell. This correction pared. The Jaccard’s index is 1 when species composition is for the cell species richness ensures that CWE values high- identical between squares and 0 when two squares have no light areas with a high proportion of endemic species but not species in common. According to the values of Jaccard’s in- necessarily high in richness (Crisp et al., 2001; Laffan and dex, the species were clustered into a dissimilarity tree, and Crisp, 2003; Laffan, Ramp and Roger, 2012). We calculated the branches with a minimum of at least three species and CWE using the “endemicity tools” extension for ArcView splitting off the basal polytomy of this tree were define as 3.2 (Danho, 2003), and performed computations at the cell the main distribution types. level (radius = 1), excluding empty grid cells from analysis. Using occurrence data of amphibians and reptiles, we Single cell calculations provide the maximum resolution for separately calculated for the two groups the Corrected the analysis at the expense of artefacts occurring in poorly Weighted Endemism index (CWE) (Crisp et al., 2001). For sampled cells (Laffan and Crisp, 2003). We assumed that calculating this index, the species are weighted by the in- herpetological explorations in Europe have been intensive verse of their cell ranges so that species with narrow ranges enough to allow calculations at single-cell level (see below are assigned relatively high weights, while species with for a discussion of this assumption; see also Ficetola et al., broader ranges are assigned progressively lower weights 2013). 8 N. Sillero et al.

Results and discussion Table 2. Point records per country from the three main databases (COUNTRIES, SEH/GBIF, and INTRODUCED) Database compilation of this compilation, for amphibians and reptiles, and for both groups together. See table 1 for number of records per The COUNTRIES database includes a total of national and personal databases. 364 814 records; the SEH/GBIF database in- Country Amphibians Reptiles Amphibians cludes 15 485 records; and the INTRODUCED and reptiles database includes 4310 records. Our compila- Albania 163 852 1015 tion thus totals 384 609 entries from 28 national Andorra 12 23 35 Austria 8365 5872 14 237 ∗ and personal databases, plus the original SEH Belgium 40 413 4251 44 664 and GBIF databases (table 1). The Spanish Her- Bosnia and 177 312 489 petological Society provided the largest amount Herzegovina Bulgaria 1108 2565 3673 of records (68 618; updated until 2005; table 2). Belarus 1258 195 1453 Other countries, like Portugal and Luxembourg Croatia 471 1924 2395 also provided their entire database, with data Czech Republic 648 436 1084 Denmark 3695 1452 5147 about locality, author, and date. Records with a Estonia 2525 480 3005 high spatial resolution (table 1) were also avail- Finland 1845 2264 4109 able for instance in Flanders (5 × 5km),Wal- F.Y.R. of Macedonia 74 201 275 lonia (4 × 4 km), and Portugal (GPS points). France 6865 5881 12 746 Georgia 742 18 760 Table 1 details the characteristics of the dif- Germany 24 380 11 116 35 496 ferent databases that were used in this study. Greece 1430 11 367 12 797 The fina number of records per species repre- Hungary 8227 3738 11 965 × Ireland 459 530 989 sented in the 50 50 km grids (total: 48 440 Italy 1583 2736 4319 occurrence records at the 50 × 50 km grid Latvia 368 63 431 level) is lower than in the sum of the three Liechtenstein 8 5 13 Lithuania 432 90 522 databases (COUNTRIES, SEH/GBIF, INTRO- Luxembourg 9539 1054 10 593 DUCED) due to record duplications caused by Malta 83240 the reduction in the spatial resolution of the Moldova 356 72 428 UTM squares (e.g. from GPS points in the Por- Montenegro 94 228 322 × Netherlands 6249 2012 8261 tuguese database to the fina 50 50 km UTM Norway 6958 3359 10 317 square). Poland 11 264 4127 15 391 Overall, 218 taxa were mapped (73 species Portugal 8054 9101 17 155 Romania 3084 4470 7554 of amphibians and 145 of reptiles; table 3), in- Russia 14 315 2695 17 010 cluding 13 amphibian and 18 reptile species that Serbia 493 721 1214 were not represented in the 1997 European At- Slovakia 1694 641 2335 Slovenia 1522 1489 3011 las (Gasc et al., 1997). However, as the study Spain 27 797 41 059 68 856 area is slightly different, 18 species from the Sweden 26 562 4253 30 815 eastern edges of the area covered by the 1997 Switzerland 3015 2464 5479 European Atlas were not mapped in our compi- Ukraine 4031 881 4912 United Kingdom 10 880 8417 19 297 lation (see Study Area section). Therefore, and considering also taxonomical changes, our com- TOTAL 241 163 143 446 384 609 pilation includes 31 newly mapped species (ta- ∗ Belgium data was composed by three different databases: ble 3). We merged 46 taxa with others in the Flanders, Wallonia, and Brussels. same species-level map (usually not more than 2-3 species per map) when their taxonomic sta- tus and/or their precise distribution boundaries them on separate maps (see section on taxo- were insufficientl known to warrant plotting nomic gaps of knowledge below and table 3). New atlas of European amphibians and reptiles 9 9 2 4 3 1 5 20 18 28 − − 134 52 63 12 76 17 38 2130 12 25 13 137 64 110 782 214 157 53 701 237 556 45 570 51 1997 Atlas Difference 2059 554 1266 296 1213 27 esponds to the 1997 European yet been evaluated by IUCN at a lassification separately for Amphibia, (partim) (partim) 448 29 (partim) (partim) (partim) (partim) (partim) (partim) (partim) Hyla meridionalis Bufo viridis Hyla arborea Triturus boscai Alytes cisternasii Alytes obstetricans Bufo calamita Euproctus asper Chioglossa lusitanica Discoglossus pictus Discoglossus sardus Hyla arborea Discoglossus galganoi Bufo bufo Bufo viridis Bombina bombina Alytes obstetricans Triturus alpestris Alytes muletensis Euproctus asper Discoglossus montalentii Euproctus montanus Euproctus platycephalus Hyla arborea Bombina variegata 3 1 7 9 8 33 93 20 28 40 26 28 18 201 134 119 210 938 996 621 477 601 data as in Atlas 1997 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 28 95 75 230 1562 359 2613 155 123 1240 ). Numbers in parentheses refer to species complexes which subsume species either not considered as ∗ 1 4 0 0 0 0 3 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 14 11 13 2 1 7 9 8 19 93 20 28 40 26 28 18 200 210 525 783 963 134 119 466 513 1332 2254 1117 50 km UTM squares) per species for this compilation and the 1997 European atlas. COUNTRIES, INTRODUCED and SEH/GBIF corresponds × LC LC LC LC NT VU VU LC LC CR NT VU NT LC LC EN LC LC LC LC IUCN COUNTRIES INTRODUCED SEH/GBIF All Species name status LC ∗ ∗ (5) LC ∗ ∗ ∗ (14) LC ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ complex (2) LC complex (11) LC ∗ Total number of records (50 Hyla sarda Hyla meridionalis Bombina bombina Alytes dickhilleni Alytes muletensis Discoglossus galganoi Discoglossus sardus Euproctus montanus Ichthyosaura alpestris Calotriton arnoldi Bufo viridis Chioglossa lusitanica Discoglossus pictus Euproctus platycephalus Discoglossus montalentii Alytes cisternasii Bufo calamita Bufo bufo Bombina variegata Calotriton asper Hyla arborea Lissotriton boscai Alytes obstetricans Bufo boulengeri Hyla intermedia MPHIBIANS A Table 3. Species valid (Speybroeck, Beukema and Crochet,The 2010) second or column for summarises whichDeficient the distribution records global LC, cannot extinction Least be risk Concern; unambiguouslyglobal NT, status Near assigned level). of Status Threatened; in each in VU, the parentheses Vulnerable; species available EN, refers databases according Endangered; to (see to cases CR, footnotes where Critically the for confirmatio Endangered IUCN detailed (dashes red explanations). indicate is list species necessary (IUCN, that due have 2012), to not taxonomic according uncertainties. to IUCNes categori (IUCN, 2001): DD, Data to the record numbersatlas. per Difference species is of the eachTestudines, subtraction database and between Squamata. included this Species in endemic compilation this to and compilation, Europe the are and 1997 marked ‘All with European data’ an atlas. summarises asterisk Species ( the are total listed number alphabetically of according records. to Atlas current 1997 c corr 10 N. Sillero et al. 4 3 1 5 7 8 12 82 35 17 − − 463 − − − − 267 030 6 31 027 18 29 23 31 8 79 11 0 82 635125 15 16 4 63 6 56 4 22 44 132 34 685 35 852 486 60 197 56 295 11 270 113 854 59 1997 Atlas Difference 1169 436 1147 417 1874 1460 520 1782 563 Rana balcanica (partim) (partim) (partim) Rana lessonae + (partim) (partim) (partim) and Rana ridibunda Rana arvalis Rana macrocnemis Rana temporaria Pleurodeles waltl Rana catesbeiana Triturus vulgaris Mertensiella luschani Triturus vittatus Rana shqiperica Rana dalmatina Rana temporaria Salamandra salamandra Salamandra lanzai Proteus anguinus Rana kl. esculenta Pelobates fuscus Pelobates syriacus Pelodytes caucasicus Triturus helveticus Triturus italicus Pelobates cultripes Rana epeirotica Rana perezi Rana graeca Rana iberica Rana italica Rana latastei Salamandra atra Triturus montandoni Salamandra salamandra Salamandrella keyserlingii Pelodytes punctatus data as in Atlas 1997 0 166 2 69 00 3 30 0 2 00 5 0 7 4 0 47 90 44 27 10 546 39 0 253 00 11 605 0000 65 78 67 29 7 69 0 383 00 8 (not included) 12 16 60 12 22 80 913 21 66 157 310 1564 366 2345 289 1980 106 720 104 1411 388 770 0 0 0 0 0 0 0 1 0 0 0 3 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 43 47 3 2 5 7 4 0 8 8 30 35 27 39 44 11 11 65 78 67 29 62 55 LC 613 NT 166 LC LC 432 LC 1254 (LC)LC 67 1979 (LC) 383 IUCN COUNTRIES INTRODUCED SEH/GBIF All Species name status LC LC NT LCLC LC 545 VU NT 253 EN VU NT LCEN 306 VU LC NT LC VU EN LC LC VU LC 833 LC 1691 LC (1) LC 1304 ∗ (12) LC 1405 ∗ / lessonae ∗ ∗ ∗ ∗ / bedriagae ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ esculentus grafi (10) ∗ ∗ ∗ ∗ ∗ kl. kl. sp. (Continued.) Salamandra atra Rana dalmatina Rana pyrenaica Pleurodeles waltl Proteus anguinus Rana iberica Lissotriton italicus Lissotriton montandoni Lyciasalamandra helverseni Ommatotriton vittatus Pelobates cultripes Pelobates fuscus Pelophylax epeiroticus Pelophylax Pelophylax ridibundus Pelophylax shqipericus Rana arvalis Rana graeca Rana italica Rana latastei Rana macrocnemis Rana temporaria Salamandra corsica Salamandra lanzai Salamandra salamandra Lissotriton helveticus Pelophylax cretensis Pelophylax Lithobates catesbeianus Pelobates syriacus Pelodytes caucasicus Pelodytes Pelophylax perezi Lissotriton vulgaris Salamandrella keyserlingii Table 3. Species New atlas of European amphibians and reptiles 11 2 1 1 7 3 15 26 14 88 91 − − − 396 − − − 2 040 88 20 20 31 40 6 0297 012 043 00 41 17 27 208 62 22 5 15 96 110 46 109 27 284 48 223 33 786 309 134 66 127 1997 Atlas Difference (partim) (partim) (partim) (partim) superspecies (partim) 1209 341 Acanthodactylus erythrurus Salamandrina terdigitata Ablepharus kitaibelii Agkistrodon halys Triturus marmoratus Caretta caretta Chelonia mydas Speleomantes italicus Speleomantes ambrosii Testudo hermanni Testudo marginata Algyroides fitzingeri Speleomantes ambrosii Speleomantes flavus Speleomantes genei Speleomantes imperialis Speleomantes supramontis Algyroides marchi Triturus cristatus Dermochelys coriacea Emys orbicularis Eretmochelys imbricata Lepidochelys kempii Lepidochelys olivacea Mauremys caspica Mauremys caspica Testudo graeca Mauremys leprosa 1 data as in Atlas 1997 0 156 0 396 0 235 0000 1 2 2 4 0 4 0 5 0 58 0 332 0 40 0 297 000 16 1 15 (not included) 0 36 0 27 0 7 (not included) 0 12 0 43 0 200 15 136 20 256 31 124 182 1550 318 1095 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 6 0 6 7 0 0 1 0 2 2 396 0 1 2 2 4 4 5 1 0 50 69 58 40 12 16 13 36 27 43 91 156 297 235 230 121 776 198 1368 LC LC LC LC status IUCN COUNTRIES INTRODUCED SEH/GBIF All Species name EN EN CR NT VU VU NT NT VU NT EN NT LC LC EN LC CR NT CR –0088 – –8 VU (4) LC/NT 332 ∗ (20) LC / terdigitata ∗ ∗ ∗ ∗ ) ∗ ∗ ∗ ) ∗ ∗ ∗ (13) ∗ ∗ complex ∗ (17) ESTUDINES QUAMATA (T (S (Continued.) Acanthodactylus erythrurus (not included) Xenopus laevis Caretta caretta Chelonia mydas Speleomantes sarrabusesis Testudo marginata Trachemys scripta Speleomantes genei Speleomantes imperialis Speleomantes italicus Speleomantes strinatii Speleomantes supramontis Triturus cristatus Algyroides marchi Salamandrina perspicillata Speleomantes ambrosii Ablepharus kitaibelii Algyroides fitzingeri Triturus/ pygmaeus marmoratus Dermochelys coriacea Speleomantes flavus Eretmochelys imbricata Emys orbicularis Lepidochelys kempii (not included) Mauremys caspica Mauremys leprosa Mauremys rivulata Testudo graeca Testudo hermanni EPTILES EPTILES R R Table 3. Species 12 N. Sillero et al. 2 6 1 1 3 1 4 6 1 1 3 50 61 61 − − − − − − − − − − 511 10 10 9 8 50 06 1 5 57 28 36 10 1914 1 12 2 22 0 36 42 14 24 16 57 14 91 30 118 56 120 123201 45 45 246 51 276 92 1997 Atlas Difference 1701 119 1042 181 (partim) (partim) 220 (partim) (partim) 11 2 Anguis fragilis Lacerta saxicola Blanus cinereus Lacerta caucasica Lacerta saxicola Elaphe dione Elaphe quatuorlineata Eremias arguta Lacerta praticola Algyroides moreoticus Algyroides nigropunctatus Alsophylax pipiens Anguis cephallonicus Lacerta bedriagae Chalcides striatus Lacerta oxycephala Lacerta armenaica Lacerta rudis Lacerta mosorensis Eirenis collaris Chalcides bedriagae Lacerta mosorensis Elaphe quatuorlineata Coluber caspius Coluber schmidti Lacerta derjugini Chalcides ocellatus Eirenis modestus Chalcides chalcides Chamaeleo chamaeleon Cyrtodactylus russowi Coronella austriaca Coronella girondica Cyrtodactylus caspius 3 7 6 6 0 1 1 0 6 1 8 data as in Atlas 1997 001 0 20 55 0 11 14 0 246 0 22 0 168 1 3 159 0 174 6 8 0 121 0 40 72 61 44 0 368 01 46 78 13 13 101 133 120 297 463 1820 219 1223 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 6 0 0 1 0 0 0 0 0 05 0 0 01 01 0 1 07 32 32 20 55 11 14 22 34 168 156 177 246 121 368 1003 (LC) 174 status IUCN COUNTRIES INTRODUCED SEH/GBIF All Species name LC (LC) 1355 –0011 NT NT LC NT NT NT LC LC –6071 LC VU NT –0022 –0055 –002 –001 – LC –204 –5 – LC LC – –001 –7 – LC –0022 ∗ ∗ ∗ ∗ ∗ (6) (not included) – ∗ ∗ ∗ ∗ ∗ ∗ ∗ (not included) sp. (7) (Continued.) Dinarolacerta montenegrina Archaeolacerta bedriagae Alsophylax pipiens Darevskia praticola Algyroides moreoticus Chalcides bedriagai Darevskia derjugini Darevskia lindholmi Anguis cephallonica Darevskia caucasica Darevskia saxicola Elaphe quatuorlineata Eremias arguta Algyroides nigropunctatus Anguis Blanus/ mariae cinereus Darevskia armenaica Darevskia rudis Dinarolacerta mosorensis Eirenis modestus Elaphe dione Elaphe sauromates Chalcides chalcides Chamaeleo africanus Dolichophis caspius Eirenis collaris Dolichophis schmidti Chalcides striatus Chalcides ocellatus Chamaeleo chamaeleon Coronella austriaca Dalmatolacerta oxycephala Coronella girondica Mediodactylus russowi Cyrtopodion caspium Table 3. Species New atlas of European amphibians and reptiles 13 3 1 1 2 2 2 5 2 7 2 1 4 2 81 81 10 13 − − − − − − − − 159 113 415 375 − − − − − 7 10 11 7 9 5 6 11 2 53 25 18 23 80 14 18 18 32 26 17 56 3 29 15 18 112 0 237 121 311 64 142 25 107 28 746 1997 Atlas Difference 1172 (partim) 361 (partim) (partim) 90 (partim) (partim) (partim) (partim) (partim) (partim) (partim) (partim) (partim) (partim) Cyrtodactylus kotschyi Malpolon monspessulanus Macrovipera lebetina Macroprotodon cucullatus Lacerta bonnali Malpolon monspessulanus Macroprotodon cucullatus Hemidactylus turcicus Lacerta graeca Coluber algirus Hierophis gemonensis Coluber viridiflavus Lacerta bonnali Lacerta bonnali Lacerta bonnali Lacerta bonnali Lacerta bonnali Lacerta horvathi Lacerta monticola Lacerta viridis Lacerta schreiberi Macrovipera schweizeri Coluber hippocrepis Coluber ravergeri Laudakia stellio Lacerta trilineata Lacerta agilis Lacerta strigata Eremias velox Lacerta viridis Eryx jaculus Laudakia caucasia Phyllodactylus europaeus Eumeces schneiderii Eryx miliaris 9 1 2 2 6 5 2 1 8 13 78 11 21 94 50 data as in Atlas 1997 2 112 0 280 0 0 0 0 0 0 0 0 0 167 0 113 0 358 0 0 0 375 0 0 00 415 0 9 135 56 159 68 371 50 375 1185 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 5 0 6 2 0 12 9 2 2 6 5 2 1 1 3 0 13 78 11 21 94 106 103 280 167 113 374 415 346 126 297 808 LC –0044 LC – LC –0202 LC –0077 NT NT LC LC EN EN NT EN NT NT CR VU LC NT EN LC LC –004343 status IUCN COUNTRIES INTRODUCED SEH/GBIF All Species name LC LC – 46 0 13 59 LC –0044 NT 44 0 0 44 –0022 –001616 (15) (LC) ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ (Continued.) Macrovipera schweizeri Mediodactylus kotschyi Macrovipera lebetina Malpolon monspessulanus Iberolacerta aranica Iberolacerta monticola Macroprotodon cucullatus Malpolon insignitus Iberolacerta cyreni Hemidactylus turcicus Hemorrhois algirus Hemorrhois hippocrepis Hierophis gemonensis Hierophis viridiflavus Iberolacerta aurelioi Iberolacerta bonnali Iberolacerta galani Iberolacerta horvathi Iberolacerta martinezricai Lacerta agilis Lacerta schreiberi Lacerta strigata Hemorrhois ravergieri Eremias velox Macroprotodon brevis Lacerta bilineata Lacerta trilineata Lacerta viridis Eryx jaculus Laudakia stellio Laudakia caucasia Euleptes europaea Eryx miliaris Hellenolacerta graeca Eumeces schneiderii Table 3. Species 14 N. Sillero et al. 1 5 3 6 0 1 6 0 2 3 7 3 6 3 1 18 42 − − − − − − − 20 8 4 77 12 14 35 40 45 24 18 22 107 76 30 32 78 36 14 23 219 36 163 452 68 181 55 244 38 158 33 236 43 413 49 665 79 1997 Atlas Difference 1967 159 (partim) 18 (partim) (partim) 62 (partim) (partim) (partim) (partim) (partim) Psammodromus algirus Podarcis taurica Pseudopus apodus Podarcis perspicillata Coluber rubriceps Coluber najadum Phrynocephalus helioscopus Phrynocephalus mystaceus Ophisops elegans Phrynocephalus guttatus Natrix tessellata Ophiomorus punctatissimus Psammodromus hispanicus Podarcis bocagei Podarcis bocagei Podarcis erhardii Podarcis hispanica Podarcis melisellensis Podarcis peloponnesiaca Podarcis tiliguerta Podarcis wagleriana Podarcis sicula Elaphe scalaris Vipera xanthina Natrix maura Natrix natrix Podarcis erhardii Podarcis filfolensis Podarcis erhardii Podarcis erhardii Podarcis lilfordi Podarcis milensis Podarcis pityusensis Podarcis wagleriana Podarcis muralis 2 6 5 3 1 8 4 9 3 36 18 65 36 16 30 18 data as in Atlas 1997 0 255 0 0 236 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 191 0 279 0 282 22 162 35 129 29 744 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 6 0 0 0 9 0 13 0 6 5 3 1 8 4 3 3 94 36 18 57 36 16 30 18 255 140 236 182 279 702 282 LC LC LC –0033 LC 92 0 14 106 LC–001919 0 0 5 5 10314 –110 –002626 LCLC 347 12 1 0 172 0 520 12 – LCLCLC 3 461 1613 0 1 4 0 509 0 2126 462 3 IUCNstatus COUNTRIES INTRODUCED SEH/GBIF All Species name LC LC EN EN LC LC VU VU EN LC VU LC NT CR LC LC LC LC (19) LC ∗ ∗ ∗ ∗ complex (3) (LC) ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ ∗ (Continued.) Psammodromus hispanicus Podarcis tiliguerta Rhinechis scalaris Psammodromus algirus Podarcis tauricus Scelarcis perspicillata Podarcis bocagei Platyceps collaris Platyceps najadum Phrynocephalus helioscopus Phrynocephalus mystaceus Ophisops elegans Phrynocephalus guttatus Natrix tessellata Ophiomorus punctatissimus Pseudopus apodus Podarcis carbonelli Podarcis cretensis Podarcis levendis Podarcis milensis Podarcis pityusensis Podarcis siculus Podarcis waglerianus Podarcis filfolensis Montivipera xanthina Natrix maura Natrix natrix Podarcis gaigeae Podarcis melisellensis Podarcis muralis Podarcis raffonei Podarcis erhardii Podarcis hispanicus Podarcis lilfordi Podarcis peloponnesiacus Table 3. Species New atlas of European amphibians and reptiles 15 9 0 9 − − 123 − 05 50 50 9 40 10 82 12 270 163 172 224311 29 48 104 7 487 102 284 41 142 40 100 28 1997 Atlas Difference 1325 81 1403 158 41 540 6900 hispanicus . , and the hemiclone kl. Tarentola mauritanica Vipera seoanei Vipera ursinii Vipera ammodytes Vipera aspis Vipera berus Typhlops vermicularis Vipera dinniki Vipera kaznakovi Telescopus fallax Elaphe longissima Lacerta vivipara Elaphe hohenackeri Lacerta lepida Trapelus sanguinolentus Vipera latasti Vipera nikolskii Elaphe situla P. lessonae 3 2 50 49 data as in Atlas 1997 as subspecies of 0 0 0 359 21 0 325 0 182 0 128 28 253 76 589 462 1406 470 1561 9463 48 440 bergeri and several yet undescribed candidate species; the precise distribution areas of these taxa remain due to uncertain identificatio to species level of numerous records especially from Portugal which , which are currently not accepted at species level by the SEH. ,aswellas 0 0 0 0 0 0 0 0 0 0 123 P. vaucheri , esculentus T. pygmaeus H. orientalis which due to their morphological similarity are not distinguished in the available databases. Z. lineatus ; records of these species are not unambiguously distinguished in the databases available to us. .kl. P and which is currently not accepted at species level by the SEH. , and P. liolepis 4 0 97 B. mariae 49 50 325 359 225 944 182 513 128 1091 V. renardi and H. molleri ,and 41 465 P. lessonae ; the distinction of these three taxa at the species level as well as their precise distribution areas require confirmatio by additional , sensu lato, T. marmoratus and Z. longissimus P. punctatus , and two undescribed candidate species of the Iberian Peninsula; the distribution area and taxonomy of these taxa require which is currently not accepted at species level by the SEH. B. cinereus A. fragililis H. arborea V. ursinii , LC 401 27 0 433 VU NT LC LC LC LC –0055 –001 VU IUCN COUNTRIES INTRODUCED SEH/GBIF All Species name status LCLC 0 98 1 0 0 13 111 1 (not included) 1 –0055 –0055 –6 LC LC P. hispanicus P. ibericus , includes records of D. jeanneae A. graeca includes records of , includes records of (8) (LC) / lessonae includes records of contains includes records of (9) complex includes A. colchica esculentus complex includes records of ∗ / renardi ∗ / renardi V. berus kl. sp. includes records of ∗ sp. includes (Continued.) Pelodytes Podarcis hispanicus Blanus/ mariae cinereus Vipera ursinii Hyla arborea Triturus/ pygmaeus marmoratus Discoglossus galganoi Zamenis longissimus / lineatus Pelophylax Anguis Tarentola mauritanica Vipera ursinii Vipera ammodytes Vipera aspis Vipera berus Vipera dinniki Vipera kaznakovi Vipera latastei Teira dugesii Telescopus fallax Timon lepidus Zamenis hohenackeri Trapelus agilis Typhlops vermicularis merged with Zamenis longissimus / lineatus Zamenis situla Zootoca vivipara Vipera seoanei OTAL T (10) more study. (6) (7) (9) (2) (3) to be elucidated. (4) are based on larvae. (5) study. (8) Table 3. Species (1) 16 N. Sillero et al.

Nine species (six amphibians and three rep- tiles) represented more than 10 000 records in the whole compiled point databases, cor- responding in almost all cases to the most widespread species in Europe. From lesser (11 696) to larger (31 638), these were: Zootoca vivipara, Anguis sp., Ichthyosaura alpestris, ). Natrix natrix, Triturus cristatus complex, Pelo- phylax kl. esculentus/lessonae, Lissotriton vul- garis, Rana temporaria, and Bufo bufo.Inthe opposite extreme, there were 41 species (13 Chrysemys picta amphibians and 28 reptiles) with less than 10 records. These species corresponded to en- demisms of mainland Europe (e.g. Iberolacerta aranica) and of the Mediterranean islands (e.g. Podarcis filfolensis). However, and particularly

or related genera (e.g. for the most widespread taxa, the higher num- ber of records also correspond to species present in distribution atlases with a high resolution, T. macedonicus ; records of these species are not unambiguously distinguished

Trachemys i.e. a high number of records. In relation with the whole database in grid format (table 3), 16 species included more than 1000 records (i.e.

. present in more than 1000 grid cells), three

T.,and dobrogicus of them with more than 2000 (i.e., Natrix na- trix, Rana temporaria, Bufo bufo). All these,

B. variegata again, were species widespread in Europe. On

T. cristatus , the other hand, 59 species were present in less P. occidentalis , which are currently not accepted at species level by the SEH. , than 10 cells, many of them endemisms (e.g. Podarcis levendis), but others were marginal

T. carnifex species with their main distribution range out- Eirenis modestus

, which are currently not accepted at species level by the SEH. side the study area (e.g. ). P. hispanicus ,and

, The increment in distribution knowledge was Rana balcanica ) which is not accepted at species level by the SEH. T. arntzeni , considerable (4224 new grid records, 19.6%). variabilis = ( which is not recognized as a species by the SEH. Although the taxa entities are not completely and which is treated as a subspecies of . The latter is currently not accepted at species level by the SEH. congruent, 44 (8.3%) taxa presented less records P. ewardsianus

T., karelinii than in the 1997 European Atlas; 17 (7.8%) the kurtmuelleri

P. cerigensis same number; and 152 (69.7%) more records balearicus might also include records of introduced specimens of other species of B. pachypus

includes Pelophylax escu-

includes (table 3). The extremes are kl. Emys trinacris refers to the Balearic populations. siculus . The latter is currently not accepted at species level by the SEH. lentus lessonae includes / with a loss of 463 records, and includes

includes Rana temporaria with a gain of 563 records. includes includes The reasons for the changes in the number of complex includes Trachemys scripta grid cells per species are manifold. Increases are usually due to an improved mapping intensity and coverage, whereas decreases are often ex- Triturus cristatus complex Bufo viridis Pelophylax ridibundus Bombina variegata Macropotodon cucullatus Pelophylax bedriagae Emys orbicularis Bufo boulengeri Psammodromus hispanicus plained by changes in taxonomy such as split- (11) (12) (13) Records of (14) (15) (16) (17) (18) in the databases available to us. (20) (19) ting of previously widespread species into dif- New atlas of European amphibians and reptiles 17 ferent species, or redefinition of taxa with cor- very patchy distribution (fig 4): north-western responding reduction of their actual ranges, but Iberian Peninsula, Po lowland, Sardinia, and also because of the low number of recent data western Greek coast were the areas with a for some countries devoid of distribution atlas higher number of threatened amphibians. On programs (see table 1). the other hand, threatened reptiles were Patterns of species richness were different widespread, especially in the Iberian and Italian in amphibians and reptiles as we will fur- peninsulas as well as in Central Europe. These ther explore in the biogeography section be- different patterns are due to the species com- low. Species richness of amphibians was high- position: threatened amphibians were mostly est in Western-Central Europe, while for rep- composed by localised endemics (e.g. Alytes tiles the southern peninsulas had the highest muletensis) while threatened reptiles included concentration of species, in particular Greece some widespread species (e.g. Emys orbic- (fig 2), which is in general agreement with anal- ularis). However, the European herpetofauna yses based on the 1997 European Atlas (Araújo, might have a higher level of conservation threat Thuiller and Pearson, 2006; Araújo et al., 2008) than currently recognised (Denoël, 2012). Fu- and the Global Amphibian Assessment (e.g. ture evaluations such as those provided through Anthony et al., 2008; Baha el Din et al., 2008). herpetological atlases could thus shed light on Several countries such as Albania, Bosnia and wider patterns of vulnerability (see e.g. Denoël, Herzegovina, Latvia, Lithuania, Ireland, F.Y.R. 2012). of Macedonia, Moldova, Montenegro and Ser- bia presented low levels of species richness, Biogeographical analysis mainly due to insufficien coverage, impossibil- The analysis of corrected weighted endemism ity of digitising chorological information pub- (CWE) highlighted the importance of Mediter- lished in journals, or because database chairs ranean islands as centres of endemism for both decided not to collaborate in our compilation. amphibians and reptiles (fig 5). For amphib- No atlases or articles with chorological data are ians, highest CWE values were found in Sar- currently available for some of these countries, dinia and Corsica, Mallorca, Sicily, and south- as far as we know. Calculating species richness ern Aegean islands. In addition, some grid cells for endemic European species only (i.e. exclud- on the Balkans and the Western Caucasus stand ing all species which have ranges extending out- out with high local endemism values. Reptiles side the study area) leads to a strong shift of showed an overall similar pattern, but some ar- species richness towards Western Europe, re- eas such as Corsica, Sicily and the southern flectin that the Balkan Peninsula holds many Aegean presented lower CWE values while ad- species with ranges extending into the Middle ditional areas of endemism were identifie on East and Caucasus, and Central Europe holds smaller Mediterranean islands such as Malta, as many widespread species with ranges extend- well as certain areas in Spain (corresponding to ing east of the Ural Mountains (fig 3). Simi- the microendemic Iberolacerta species) and the larly, the Caucasus region was not identifie as Balkans. an area of endemism because most of the nu- However, these CWE calculations were some- merous species endemic to the Caucasus Moun- what biased due to our definitio of the study tains are distributed on the southern slopes as area. Because the CWE calculation took the full well, i.e. outside Europe as we define it. range size of a species into account, and the The species richness of European threat- full range sizes of some species (104) were not ened amphibians, following the IUCN cat- included in the study area (and thus not com- egories Vulnerable (VU), Endangered (EN), plete in the compilation database used for anal- and Critically Endangered (CR), presented a ysis, especially regarding species distributed in 18 N. Sillero et al.

Figure 2. Maps of Europe showing species richness separately for amphibians and reptiles, based on species distribution maps of all non-introduced species occurring in the study area. New atlas of European amphibians and reptiles 19

Figure 3. Maps of Europe showing species richness based on species distribution maps of European endemic amphibians and reptiles (i.e. including only species whose range does not extend beyond the study area). 20 N. Sillero et al.

Figure 4. Maps of Europe showing species richness based on species distribution maps of European threatened amphibians and reptiles, including the IUCN categories Vulnerable, Endangered, and Critically Endangered. New atlas of European amphibians and reptiles 21

Figure 5. Maps of Europe showing Corrected Weighted Endemism (CWE) based on species distribution maps of European endemic amphibians and reptiles. 22 N. Sillero et al. the Balkan Peninsula or widespread in Central Nine and 13 main distribution types were Europe), it is possible that CWE values in the identifie for amphibians (named CA1-CA9; eastern part of Europe were slightly inflated table 4 and online Supplementary fig S2) In other words, many species occur in just a and reptiles respectively (named CR1-CR13; small part of the study area and thus appear table 5 and online Supplementary fig S3). to be range-restricted and micro-endemic, while Many species that formed clusters of only one in fact they have wide ranges extending further member, corresponding mainly to regional en- east outside our study area. This phenomenon is demics, were not assigned to a distribution inversely analogous to the species richness pat- type number and are not further considered here. In the following, we will briefl charac- terns of endemic European amphibians, where terise distribution types and mention one rep- the same artefacts lead to inflate species rich- resentative species for each. Amphibian distri- ness values in Western Europe. These prob- bution types can be characterised as follows lems call for caution in interpreting the biogeo- (table 4 and online Supplementary fig S2): graphic analysis of our data, but do not invali- CA1, species distributed in the Iberian Penin- date the observed general patterns. Furthermore, sula and western France (e.g. Hyla meridio- from the point of view of conservation priori- nalis); CA2, western European species (e.g. ties, the observed patterns of Europe-endemic Alytes obstetricans); CA3, species widespread species richness are highly relevant since they in Europe (e.g. Bufo bufo); CA4, Pyrenaean highlight the importance of specifi areas (es- species (e.g. Calotriton asper); CA5 and CA6, pecially the Mediterranean islands), where the species distributed in Corsica, Sardinia, and the survival of a large number of European en- southern Mediterranean coast (e.g. Euproctus demics fully depends on European conservation platycephalus, CA5; and Discoglossus sardus, efforts. CA6); CA7, species from the Italian Penin-

Table 4. Amphibian species grouped by main distribution types. See the dendrogram in online Supplementary fig S1. Distribution types were named with codes following Baroni-Urbani and Collingwood (1976) and Baroni-Urbani and Collingwood (1977).

Distribution type CA1 Distribution type CA3 Distribution type CA6 Alytes cisternasii Bombina bombina Euproctus platycephalus Chioglossa lusitanica Bufo bufo Speleomantes flavus Discoglossus galganoi Bufo viridis Speleomantes supramontis Hyla meridionalis Lissotriton montandoni Distribution type CA7 Lissotriton boscai Lissotriton vulgaris Pelobates cultripes Pelobates fuscus Hyla intermedia Pelodytes sp. Pelophylax kl. esculentus/lessonae Rana italica Pelophylax perezi Pelophylax ridibundus Salamandrina Pleurodeles waltl Rana arvalis perspicillata/terdigitata Rana iberica Rana temporaria Speleomantes italicus Triturus marmoratus/pygmaeus Triturus cristatus complex Lissotriton italicus Distribution type CA2 Distribution type CA4 Distribution type CA8 Alytes obstetricans Calotriton asper Ommatotriton vittatus Bombina variegata Pelophylax kl. grafi Pelodytes caucasicus Bufo calamita Rana pyrenaica Rana camerani Hyla arborea complex Distribution type CA5 Distribution type CA9 Ichthyosaura alpestris Lissotriton helveticus Discoglossus sardus Proteus anguinus Salamandra salamandra Euproctus montanus Rana latastei Rana dalmatina Hyla sarda Salamandra atra Discoglossus montalenti Salamandra corsica New atlas of European amphibians and reptiles 23

Table 5. Reptile species grouped by main distribution types. See dendrogram in online Supplementary fig S2. Distribution types were named with codes following Baroni-Urbani and Collingwood (1976) and Baroni-Urbani and Collingwood (1977).

Distribution type CR1 Distribution type CR4 Distribution type CR8 Ablepharus kitaibelii Algyroides moreoticus Darevskia caucasica Darevskia praticola Anguis cephalonica Eirenis collaris Dolichophis caspius Eryx jaculus Eirenis modestus Elaphe sauromates Hellenolacerta graeca Hemorrhois ravergieri Lacerta viridis Hierophis gemonensis Laudakia caucasia Montivipera xanthina Lacerta trilineata Distribution type CR9 Natrix tessellata Malpolon insignitus Ophisops elegans Mauremys rivulata Darevskia derjurgini Podarcis tauricus Mediodactylus kotschyi Darevskia saxicola Testudo graeca Ophiomorus punctatissimus Vipera kaznakovi Vipera ammodytes Platyceps najadum Podarcis erhardii Distribution type CR10 Distribution type CR2 Podarcis peloponnesiacus Dolichophis schmidtii Acanthodactylus erythrurus Pseudopus apodus Eumeces schneiderii Blanus sp. Telescopus fallax Macrovipera lebetina Chalcides bedriagae Testudo marginata Mauremys caspica Chalcides striatus Typhlops vermicularis Chamaeleo chamaeleon Zamenis situla Distribution type CR11 Coronella girondica Distribution type CR5 Elaphe dione Hemidactylus turcicus Eremias arguta Hemorrhois hippocrepis Algyroides nigropunctatus Eremias velox Macroprotodon brevis Dalmatolacerta oxycephalus Eryx miliaris Malpolon monspessulanus Dinarolacerta mosorensis Lacerta strigata Mauremys leprosa Podarcis melisellensis Phrynocephalus guttatus Natrix maura Distribution type CR6 Phrynocephalus mystaceus Podarcis hispanicus complex Trapelus agilis Psammodromus algirus Anguis sp. Vipera ursinii/renardi Psammodromus hispanicus complex Coronella austriaca Rhinechis scalaris Emys orbicularis Distribution type CR12 Lacerta agilis Tarentola mauritanica Iberolacerta aranica Natrix natrix Timon lepidus Iberolacerta aurelioi Vipera latastei Vipera berus Iberolacerta bonnali Zootoca vivipara Distribution type CR3 Distribution type CR13 Distribution type CR7 Algyroides fitzingeri Iberolacerta galani Chalcides chalcides Archaeolacerta bedriagae Iberolacerta martinezricai Elaphe quatorlineata Euleptes europea Iberolacerta monticola Podarcis tiliguerta Hierophis viridiflavus Lacerta schreiberi Lacerta bilineata Podarcis bocagei Podarcis muralis Podarcis carbonelli Podarcis sicula Vipera seoanei Testudo hermanni Vipera aspis Zamenis longissimus/lineatus sula (e.g. Hyla intermedia); CA8, Caucasian ranean countries (e.g. Malpolon monspessu- species (e.g. Pelodytes caucasicus); and CA9, lanus); CR3, Corsican and Sardinian species alpine and dinaric species (e.g. Salamandra (e.g. Archaeolacerta bedriagae); CR4, species atra). In the case of reptiles (table 5 and on- from the Balkan Peninsula and Eastern Eu- line Supplementary fig S3): CR1, species dis- rope (e.g. Malpolon insignitus); CR5, species tributed along the Italian and Balkan Penin- from the eastern Adriatic coast (e.g. Podarcis sulas as well as south-eastern Europe (e.g. melisellensis); CR6, widespread in all of Eu- Natrix tessellata); CR2 grouped species dis- rope (e.g. Anguis sp.); CR7, western-central Eu- tributed along the western-southern Mediter- ropean species (e.g. Vipera aspis); CR8, CR9, 24 N. Sillero et al. and CR10, Caucasian species (e.g. Darevskia including species for the three Mediterranean caucasica, CR8; Vipera kaznakovi, CR9; Mau- peninsulas (CR4 and CR7) are also considered remys caspica, CR10); CR11, South-eastern by Rueda, Rodríguez and Hawkins (2010). No European species (e.g. Eremias velox); CR12, widespread species (e.g. Bufo bufo or Vipera species from the Central Pyrenees (e.g. Ibero- berus) f t in any of the regions identifie by lacerta bonnali); and CR13, species occurring Bunce et al. (2002) or Rueda, Rodríguez and in the north-western Iberian Peninsula (e.g. La- Hawkins (2010). certa schreiberi). Taxonomic and mapping gaps of knowledge These distribution types were partly but not fully congruent with those published for a more About ten species-level units in our analysis limited study area (i.e. the Iberian Peninsula; are characterised by taxonomic uncertainty or Sillero et al., 2009). The discordances can be by difficultie in species identification some explained by a higher number of species in- of these are (or might be) composed of dif- cluded in the present analysis, a larger size of ferent taxa (see footnotes in table 3). Particu- the study area, and a different spatial resolution lar taxonomic efforts are needed to clarify both the status and the precise distribution limits of the grid. As Europe holds more species and is of the Bufo viridis complex (balearicus, vari- considerably larger than the Iberian Peninsula, abilis, viridis), the Hyla arborea complex (H. the resulting main distribution types at least arborea, H. molleri, and H. orientalis), Iberian partially included the Iberian distribution types. Pelodytes (P. ibericus, P. punctatus, and two The definitio and interpretation of distribution yet undescribed candidate species), the Anguis types is always relative and strongly depends on fragilis complex (A. colchica, A. graeca, A. the study area. fragilis), and the Podarcis hispanicus complex The distribution types define in this work for (P. hispanicus sensu lato, P. liolepis, P. vaucheri European amphibians and reptiles are not in full and several undescribed candidate species). Fur- agreement with previous biogeographical clas- thermore, in the following species complexes, sifications because our classificatio was based the precise distribution ranges of each species on the distribution of species (always incom- need to be determined (preferably using ge- plete) and not on environmental data (Bunce et netic methods; Joger et al., 2007) and the avail- al., 2002) or distribution data from herpetolog- able records (and new future records) need ical guide books (e.g. range polygons on con- to be refine to distinguish between the dif- tinental maps; Rueda, Rodríguez and Hawkins, ferent species: Triturus marmoratus/pygmaeus, Triturus carnifex cristatus dobrogicus karelinii 2010). Bunce et al. (2002) define 59 environ- / / / / macedonicus Blanus cinereus mariae Psam- mental classes based in a grid square of 0.5 min , / , modromus hispanicus complex (P. edward- (i.e. ca. 55 km). As Bunce et al. (2002) did sianus, P. hispanicus, and P. occidentalis), not provide a hierarchical tree of environmental Vipera ursinii/renardi, and Zamenis longis- classes, only some of these classes had corre- simus/lineatus. This list of taxa in need of tax- spondence with our distribution types (e.g. CR6 onomic and distributional revision is clearly and CR13). Rueda, Rodríguez and Hawkins not exhaustive and was driven by the particu- (2010) identifie respectively seven and eight lar problems that we have identifie while as- biogeographical regions for amphibians and sembling the distributional data sets. It is clear reptiles in Europe. In the case of amphibians, that taxonomic revision is also needed in other Rueda, Rodríguez and Hawkins (2010) clus- species of European amphibians and reptiles, tered the distribution types CA8 and CA9 in one especially those in south-eastern Europe. In- single region. In reptiles, the distribution types deed, even for the most studied complexes, such New atlas of European amphibians and reptiles 25 as the crested newts (Wielstra and Arntzen, Conclusions and future tasks: the 2011), reliably attributing all grid cells to ei- distributed database network system ther of the newly recognized species within Distribution maps are ephemeral products in the limits of distribution of the entire complex constant need for updating. Therefore, the most turned out to be impossible. The issues identi- important part of a chorological atlas is its fie here are particularly pressing, because they database, which should be operative for a long often concern widespread species where clari- time. For this reason, the SEH Mapping Com- ficatio of the exact distribution boundaries re- mittee decided to implement a system of dis- quires intensive sampling. tributed online databases, as this is the only so- Subtracting the number of species (amphib- lution to avoid problems of data duplication and ians and reptiles merged) for each grid cell in actualisation, and to ensure that the owners of the 1997 European Atlas from the respective each sub-database maintain the control over its value in our compilation yields a pattern reflect administration. The f rst prototype of this sys- ing the overall increased coverage and mapping tem is ready (see Sillero et al., 2014). In the near intensity, especially in Western and Central Eu- future, we hope the system might connect the rope (fig 6). However, in the new compilation a databases of each European country. lower overall number of species per grid cell is An important future aspect will be to stan- present in some countries. This counter-intuitive dardise the date of each record in each of pattern is partly explained by the fact that for the national databases as well as in the SEH some areas the 1997 European Atlas was based database (see also Denoël, 2012). At present, on expert opinion about the occurrence of a precise dates of observation are provided for species in a grid cell (Gasc et al., 1997), and each record in some of the databases, but com- underlying records were not available any more pletely lacking in others. Furthermore, histori- for the new national atlases. Contemporary re- cal records often lack any precise date. A sys- gional and national atlases, on the contrary, typ- tem of minimum date (at least year) for each ically only take fully documented records into record needs to be implemented to allow query- ing the databases for possible changes in species account. Moreover, we mainly compiled pub- range, e.g. in the context of both climate change lished data. Therefore, our compilation lacks all and land cover use, and accurate dates would chorological information in personal databases even allow evaluating phenological changes. or journals not available to us. For those coun- This would imply to have multiple records for tries where new national mapping data exist we each cell grids when data are available for sev- excluded the SEH/GBIF database records from eral years for instance. our compilation, therefore for countries such One major problem cannot be solved by the as Greece and Ukraine the current compilation distributed database system, namely the lack contains fewer grid cell records which however of funding and personnel in many countries are better documented than those in the 1997 to set up a national database, collect mapping European Atlas. In general, south-eastern Eu- data, validate each record, and feed them into rope concentrates a high species richness espe- the system. It therefore will be important to cially of reptiles, but many countries in this area activate also other sources from which these lack national atlases. Future efforts should be data could be obtained. Distribution mapping targeted to encourage and support national map- and species monitoring are research field with ping efforts in this region. In addition, a Euro- a well-developed tradition of citizen science pean initiative might be useful to set up a map- contribution. In many countries of central and ping campaign to fil in these crucial distribu- northern Europe, the bulk of amphibian and tional gaps. reptile distribution data are collected by volun- 26 N. Sillero et al.

Figure 6. Differences in species richness of all non-introduced species of European amphibians and reptiles between this compilation and the 1997 European Atlas (Gasc et al., 1997). New atlas of European amphibians and reptiles 27 teers, many of which do not hold biology de- ing ways to gather all species occurrence data grees and are not professionally working as her- available in Europe is a major challenge for petologists. We feel that activating such vol- the future. Integrating the temporal dimension unteering work especially in southern coun- and measures of spatial uncertainty to all point tries, and among tourists visiting these coun- records in the original databases is another nec- tries, should be an important resource to fil essary improvement to allow detailed modelling mapping gaps (Bonardi et al., 2011). Provid- of the impacts of land use and climate change, ing a common platform to enter such observa- and we call for concerted and varied efforts to tions, accompanied by photographic documen- fil the geographic and taxonomic gaps identi- tation, will be a step to achieve this goal, if fied coupled with a functional and robust validation procedure. Such an online platform for enter- ing data will be provided by the SEH on- Acknowledgements. We are grateful to a large number of persons who have contributed or facilitated data, ideas in- line database system. However, major chal- formation, and discussions to this project: Albert Montori, lenges remain, such as integration with other Ali Salamun, Armando Loureiro, Anamarija Zagar, Annie systems like iNaturalist (www.inaturalist.org), Zuiderwijk, Benedikt Schmidt, Burkhard Thiesmeier, Cá- tia Matos, Claes Andrén, Ciprian Samoila, Edmée Engel, Observado.org (www.observado.org), or Tel- Thierry Kinet, Dan Cogalniceanu,˘ Daniel Jablonski, Darvin mee (www.telmee.nl), the feedback of the cen- Dobrnjic,´ David Tarknishvilli, Elena Argaña, Elvir Tanovic,´ trally collected data into the national databases, Idriz Haxhiu, Jean-Christophe de Massary, Jean-Philippe Siblet, Horace Da Costa, Patrick Haffner, Benoit Lefeuvre, and especially, the review and scientifi valida- Jiri Moravec, Katja Poboljšaj, Marco Zuff , Michael Veith, tion process before the contributed data are in- Miguel A. Carretero, Monika Hachtel, Nikolay Tzankov, cluded in these databases (Boakes et al., 2010; Oldrich Kopecky, Oleksandr Zinenko, Rainer Günther, Robert Jooris, Sandra Nieto-Roman, Sergé Bogaerts, Silvia Bonter and Cooper, 2012; Ficetola et al., 2013). Zumbach, Wouter Beukema and Susanne Hauswaldt. Our An important point is also that each national or project was based on a varied set of national and regional local database should use the same taxonomic atlases. We are therefore very grateful to the different so- cieties and the atlas coordinators: A. Loureiro (Portugal), list. Finally, there is a large variation of resolu- Asociación Herpetológica Española (Spain), RAVON (The tion between distribution atlases: although some Netherlands), Service du Patrimoine Naturel (Muséum Na- use point coordinates, others provide only large tional d’Histoire Naturelle, France), Koordinationsstelle für Amphibien- und Reptilienschutz in der Schweiz (KARCH, areas. At the current stage, the grid size resolu- Switzerland), Societas Herpetologica Italica (Italy), Natu- tion of 50 × 50 km reduces this problem, but in urpunt – Hyla (Flanders), Raînne-Natagora (Wallonia and the long term, the realisation of more detailed Brussels), Musée National d’Histoire Naturelle (Luxem- maps would require the centralization of highly bourg), Societas Herpetologica Slovenica and Center za kar- tografij favne in flor (Slovenia). We deeply thank the detailed data from each database. thousands of volunteers that contributed to the atlases and In summary, the data presented here pro- distribution databases. List of names can be found through vide a f rst, tentative step towards an interac- their relevant societies and local atlases. We also thank the group of specialists of the SEH than define the taxonomic tive, dynamic and distributed database of the reference list. This research project was funded by Soci- spatial distribution of European amphibians and etas Europaea Herpetologica. Authors were funded by the reptiles. The grid maps of all species made following institutions: the Fundação para a Ciência e Tec- nologia (Portugal) grant SFRH/BPD/26666/2006 to NS, the available along with this paper will facilitate Spanish Ministry of Environment 206/2010 grant and the conservation-related studies and actions, and BiodivERsA: EC21C to DRV, the Ministry of Education, will inform and guide further activities to im- Science and Technological Development of Republic of Ser- bia grant 173025 to JCI, Fonds Spéciaux de la Recherche prove and complete the database. However, it grant C11/23 and Fonds de la Recherche Scientifique-FNR should be kept in mind that they are currently Crédit aux Chercheurs grant 1.5.040.10.F to MD (Fonds de dependent on availability of digital databases, la Recherches Scientifique-FNR Research associate, Bel- gium), the Generalitat de Catalunya FI-DGR grant, Spain to and not only on species presence or even on PdD, and a Georg Forster Fellowship of the Alexander von current knowledge on species distribution. Find- Humboldt Foundation to AR. 28 N. Sillero et al.

References A., García, A., Aguilar, C., Ajtic, R., Akarsu, F., Alen- car, L.R.V., Allison, A., Ananjeva, N., Anderson, S., An- Anthony, B., Arntzen, J.W., Baha el Din, S., Böhme, drén, C., Ariano-Sánchez, D., Arredondo, J.C., Auliya, W., Cogalniceanu,˘ D., Crnobrnja-Isailovic,´ J., Crochet, M., Austin, C.C., Avci, A., Baker, P.J., Barreto-Lima, P.-A., Corti, C., Griff ths, R., Kaneko, Y., Kuzmin, S., A.F., Barrio-Amorós, C.L., Basu, D., Bates, M.F., Batis- Lau, M.W.N., Li, P., Lymberakis, P., Marquez, R., Pa- tella, A., Bauer, A., Bennett, D., Böhme, W., Broadley, penfuss, T., Pleguezuelos, J.M., Rastegar, N., Schmidt, D., Brown, R., Burgess, J., Captain, A., Carreira, S., B., Slimani, T., Sparreboom, M., Ugurtas, I., Werner, Castañeda, M.D.R., Castro, F., Catenazzi, A., Cedeño- Y., Xie, F. (2008): Amphibians of the Palearctic realm. Vázquez, J.R., Chapple, D.G., Cheylan, M., Cisneros- In: Threatened Amphibians of the World, p. 106-111. Heredia, D.F., Cogalniceanu, D., Cogger, H., Corti, Stuart, S.N., Hoffmann, M., Chanson, J.S., Cox, N.A., C., Costa, G.C., Couper, P.J., Courtney, T., Crnobrnja- Berridge, R.J., Ramani, P., Young, B.E., Eds, Lynx, Isailovic, J., Crochet, P.-A., Crother, B., Cruz, F., Dal- Barcelona. try, J.C., Daniels, R.J.R., Das, I., de Silva, A., Dies- Araújo, M.B., Pearson, R.G. (2005): Equilibrium of species’ mos, A.C., Dirksen, L., Doan, T.M., Dodd, C.K., Doody, distributions with climate. Ecography 5: 693-695. J.S., Dorcas, M.E., Duarte de Barros Filho, J., Egan, Araújo, M.B., Thuiller, W., Williams, P.H., Reginster, I. V.T., El Mouden, E.H., Embert, D., Espinoza, R.E., Fal- (2005): Downscaling European species atlas distribu- labrino, A., Feng, X., Feng, Z.-J., Fitzgerald, L., Flores- tions to a f ner resolution: implications for conservation Villela, O., França, F.G.R., Frost, D., Gadsden, H., Gam- planning. Glob. Ecol. Biogeogr. 1: 17-30. ble, T., Ganesh, S.R., Garcia, M.A., García-Pérez, J.E., Araújo, M.B., Thuiller, W., Pearson, R.G. (2006): Climate Gatus, J., Gaulke, M., Geniez, P., Georges, A., Ger- warming and the decline of amphibians and reptiles in lach, J., Goldberg, S., Gonzalez, J.-C.T., Gower, D.J., Europe. J. Biogeogr. 33: 1712-1728. Grant, T., Greenbaum, E., Grieco, C., Guo, P., Hamilton, Araújo, M.B., Nogués-Bravo, D., Valdes, P.J., Rahbek, C. A.M., Hare, K., Hedges, S.B., Heideman, N., Hilton- (2008): Quaternary climate changes explain diversity Taylor, C., Hitchmough, R., Hollingsworth, B., Hutchin- among reptiles and amphibians. Ecography 31: 8-15. son, M., Ineich, I., Iverson, J., Jaksic, F.M., Jenkins, Arnold, H.R. (1995): Atlas of amphibians and reptiles R., Joger, U., Jose, R., Kaska, Y., Kaya, U., Keogh, in Britain. ITE research publication no. 10, London, J.S., Köhler, G., Kuchling, G., Kumluta¸s, Y., Kwet, A., HMSO. La Marca, E., Lamar, W., Lane, A., Lardner, B., Latta, Baha el Din, S., Böhme, W., Corti, C., Crnobrnja-Isailovic,´ C., Latta, G., Lau, M., Lavin, P., Lawson, D., LeBre- J., Lymberakis, P., Marquez, M., Miaud, C., Slimani, ton, M., Lehr, E., Limpus, D., Lipczynski, N., Lobo, T., Ugurtas, I., Werner, Y. (2008): The status and dis- A.S., López-Luna, M.A., Luiselli, L., Lukoschek, V., tribution of amphibians in the Mediterranean Basin. In: Lundberg, M., Lymberakis, P., Macey, R., Magnusson, Threatened Amphibians of the World, p. 113. Stuart, W.E., Mahler, D.L., Malhotra, A., Mariaux, J., Maritz, S.N., Hoffmann, M., Chanson, J.S., Cox, N.A., Berridge, B., Marques, O.A.V., Márquez, R., Martins, M., Master- R.J., Ramani, P., Young, B.E., Eds, Lynx, Barcelona. son, G., Mateo, J.A., Mathew, R., Mathews, N., Mayer, Baroni-Urbani, C., Collingwood, C.A. (1976): A numerical G., McCranie, J.R., Measey, G.J., Mendoza-Quijano, analysis of the distribution of British Formicidae (Hy- F., Menegon, M., Métrailler, S., Milton, D.A., Mont- menoptera, Aculeata). Verhandl. Naturf. Ges. Basel 85: gomery, C., Morato, S.A.A., Mott, T., Muñoz-Alonso, 51-91. A., Murphy, J., Nguyen, T.Q., Nilson, G., Nogueira, Baroni-Urbani, C., Collingwood, C.A. (1977): The zoo- C., Núñez, H., Orlov, N., Ota, H., Ottenwalder, J., Pa- geography of ants (Hymenoptera, Formicidae) in North- penfuss, T., Pasachnik, S., Passos, P., Pauwels, O.S.G., ern Europe. Acta Zool. Fennica 152: 2-34. Pérez-Buitrago, N., Pérez-Mellado, V., Pianka, E.R., Baroni-Urbani, C., Ruffo, S., Vigna Taglianti, A. (1978): Pleguezuelos, J., Pollock, C., Ponce-Campos, P., Powell, Materiali per uma biogeografi italiana fondata se alcuni R., Pupin, F., Quintero Díaz, G.E., Radder, R., Ramer, J., generi di coleotteri cicindelidi, carabidi, e crisomelidi. Rasmussen, A.R., Raxworthy, C., Reynolds, R., Rich- Estr. Mem. Soc. Ent. Ital. 56: 35-92. man, N., Rico, E.L., Riservato, E., Rivas, G., da Rocha, Bauwens, D., Claus, K. (1996): Verspreiding van Amfi P.L.B., Rödel, M.-O., Rodríguez Schettino, L., Roosen- bieën en Reptielen in Vlaanderen. De Wielewaal Natu- burg, W.M., Ross, J.P., Sadek, R., Sanders, K., Santos- urvereniging, Turnhout, Belgium. Barrera, G., Schleich, H.H., Schmidt, B.R., Schmitz, A., Bitz, A., Fischer, K., Simon, L., Thiele, R., Veith, M., Sharifi M., Shea, G., Shi, H.-T., Shine, R., Sindaco, Eds (1996): Die Amphibien und Reptilien in Rheinland- R., Slimani, T., Somaweera, R., Spawls, S., Stafford, P., Pfalz. GNOR, Nassau, Germany. Stuebing, R., Sweet, S., Sy, E., Temple, H.J., Tognelli, Boakes, E.H., McGowan, P.J.K., Fuller, R.A., Ding, C.Q., M.F., Tolley, K., Tolson, P.J., Tuniyev, B., Tuniyev, S., Clark, N.E., O’Connor, K., Mace, G.M. (2010): Dis- Üzüm, N., van Buurt, G., Van Sluys, M., Velasco, A., torted views of biodiversity: spatial and temporal bias Vences, M., Veselý, M., Vinke, S., Vinke, T., Vogel, in species occurrence data. PLoS Biology 8: e1000385. G., Vogrin, M., Vogt, R.C., Wearn, O.R., Werner, Y.L., Böhm, M., Collen, B., Baillie, J.E.M., Bowles, P., Chan- Whiting, M.J., Wiewandt, T., Wilkinson, J., Wilson, B., son, J., Cox, N., Hammerson, G., Hoffmann, M., Liv- Wren, S., Zamin, T., Zhou, K., Zug, G. (2013): The con- ingstone, S.R., Ram, M., Rhodin, A.G.J., Stuart, S.N., servation status of the world’s reptiles. Biol. Conserv. van Dijk, P.P., Young, B.E., Afuang, L.E., Aghasyan, 157: 372-385. New atlas of European amphibians and reptiles 29

Bonardi, A., Manenti, R., Corbetta, A., Ferri, V., Fiacchini, Ficetola, G.F., Bonardi, A., Sindaco, R., Padoa-Schioppa, D., Giovine, G., Macchi, S., Romanazzi, E., Soccini, C., E. (2013): Estimating patterns of reptile biodiversity in Bottoni, L., Padoa Schioppa, E., Ficetola, G.F. (2011): remote regions. J. Biogeogr. 40: 1202-1211. Usefulness of volunteer data to measure the large scale Gasc, J.-P., Cabela, A., Crnobrnja-Isailovic,´ J., Dolmen, D., decline of “common” toad populations. Biol. Conserv. Grossenbacher, K., Haffner, P., Lescure, J., Martens, H., 144: 2328-2334. Martínez-Rica, J.P., Maurin, H., Oliveira, M.E., Sofi Bonter, D.N., Cooper, C.B. (2012): Data validation in cit- anidou, T.S., Veith, M., Zuiderwijk, A. (1997): Atlas izen science: a case study from Project FeederWatch. of Amphibians and Reptiles in Europe. Societas Eu- Front Ecol. Environ. 10: 305-309. ropaea Herpetologica and Muséum national d’Histoire Bunce, R.H., Carey, P., Elena-Rossello, R., Orr, J., Watkins, naturelle, Paris. J., Fuller, R. (2002): A comparison of different biogeo- Głowacinski,´ Z., Raf nski,´ J., Eds (2003): Atlas płazów i graphical classification of Europe, Great Britain and gadów Polski. Status – Rozmieszczenie – Ochrona. In- Spain. J. Environ. Managem. 2: 121-134. spekcja Ochrony Srodowiska,´ Instytut Ochrony Przy- Cabela, A., Grillitsch, H., Tiedemann, F. (2001): Atlas zur rody PAN, Warszawa-Kraków, Poland. Verbreitung und Ökologie der Amphibien und Rep- Günther, R., Ed. (1996): Die Amphibien und Reptilien tilien in Österreich: Auswertung der herpetofaunistis- Deutschlands. Gustav Fischer Verlag, Jena. chen Datenbank der herpetologischen Sammlung des Hofer, U., Monney, J.-C., Dušej, G. (2001): Die Reptilien naturhistorischen Museums in Wien. Umweltbunde- der Schweiz: Verbreitung, Lebensräume, Schutz / Les samt, Vienna, Austria. Reptiles de Suisse: Répartition, Habitats, Protection / Cheers, C. (2005): Geographica – The Complete Illustrated I Rettili della Svizzera: Distribuzione, Habitat, Pro- Atlas of the World. Random House, Australia. tezione. Birkhäuser, Basel. Cogalniceanu,˘ D., Szekely, P., Samoila,˘ C., Iosif, R., Tudor, Hopkins, W.A. (2007): Amphibians as models for studying 48 M., Plaia¸˘ su, R., Stanescu,˘ F., Rozylowicz, L. (2013a): environmental change. Ilar. J. : 270-277. IUCN (International World Conservation Union) (2001): Diversity and distribution of amphibians in Romania. IUCN red list categories. Version 3.1. Species Survival ZooKeys 296: 35-57. Commission, IUCN, Gland, Switzerland, and Cam- Cogalniceanu,˘ D., Rozylowicz, L., Székely, P., Samoila,˘ bridge, United Kingdom. C., Stanescu,˘ F., Tudor, M., Székely, D., Iosif, R. IUCN (International World Conservation Union) (2012): (2013b): Diversity and distribution of reptiles in Roma- The IUCN Red List of Threatened Species. Ver- nia. ZooKeys 341: 49-76. sion 2012.2. http://www.iucnredlist.org. Downloaded on Corti, C., Nistri, A., Poggesi, M., Vanni, S. (1991): Biogeo- 17 October 2012. graphical analysis of the Tuscan herpetofauna (Central Jacob, J.-P., Percsy, C., de Wavrin, H., Graitson, E., Kinet, Italy).Rev.Esp.Herp.5 [1990]: 51-75. T., Denoël, M., Paquay, M., Percsy, N., Remacle, A. Corti, C., Lo Cascio, P., Vanni, S., Turrisi, G.F., Vaccaro, A. (2007): Amphibiens et Reptiles de Wallonie. Série (1997): Amphibians and Reptiles of the circumsicilian Faune – Flore – Habitats no 2. Aves – Raînne et islands: new data and some considerations. Boll. Mus. Direction Générale des Ressources naturelles et de 15 Reg. Sci. Nat. Torino (1): 179-211. l’Environnement, Ministère de la Région wallonne, Na- Corti, C., Masseti, M., Delfino M., Perez-Mellado, V. mur, Belgium. (1999): Man and herpetofauna of the Mediterranean is- Jalas, J., Suonuinen, J. (1972): Atlas Florae Europaea. Dis- 13 lands. Rev. Esp. Herp. : 83-100. tribution of Vascular Plants in Europe, Vol. 1: Pterido- Corti, C., Capula, M., Luiselli, L., Sindaco, R., Razzetti, phyta. (T. C. for Mapping the Flora of Europe and Soci- E., Eds (2010): Fauna d’Italia, Vol. XLV, Reptilia. etas Biologica Fennica Vanamo, Ed.). Helsinki, Finland. Calderini, Bologna, Italy. Jetz, W., McPherson, J.M., Guralnick, R.P. (2011): Integrat- Cox, N., Chanson, J., Stuart, S. (2006): The Status and Dis- ing biodiversity distribution knowledge: toward a global tribution of Reptiles and Amphibians of the Mediter- map of life. Trends. Ecol. Evol. 27: 151-159. ranean Basin. Gland, Switzerland and IUCN, Cam- Joger, U., Fritz, U., Guicking, D., Kalyabina-Hauf, S., bridge, UK. Nagy, Z.T., Wink, M. (2007): Phylogeography of west- Creemers, R.C.M., van Delft, J.J.C.W., Eds (2009): De ern Palaearctic reptiles – Spatial and temporal speciation Amfibieë en Reptielen van Nederland. – Nederlandse patterns. Zoolr. Anz. 246: 293-313. Fauna. Nationaal Natuurhistorisch Museum Naturalis, Kuzmin, S.L. (2013): The Amphibians of the Former Soviet European Invertebrate Survey – Leiden, Netherlands. Union. Pensoft Publisher, Sofi and Moscow. Crisp, M.D., Laffan, S., Linder, H.Ü., Monro, A. (2001): Kypnjehko, B.E., Bepbec, H.T. (1999): Amphibians and Endemism in the Australian f ora. J. Biogeogr. 28: 183- Reptiles of Ukraine. Genesa, Kiev. 198. Laffan, S.W., Crisp, M.D. (2003): Assessing endemism Danho, N. (2003): Endemicity tools. ESRI Support. http:// at multiple spatial scales, with an example from the arcscripts.esri.com/details.asp?dbid=14537 (accessed: Australian vascular f ora. J. Biogeogr. 30: 511-520. November 2012). Laffan, S.W., Ramp, D., Roger, E. (2012): Using endemism Denoël, M. (2012): Newt decline in Western Europe: high- to assess representation of protected areas – the family lights from relative distribution changes within guilds. Myrtaceae in the Greater Blue Mountains World Her- Biodivers. Conserv. 21: 2887-2898. itage Area. J. Biogeogr. 40: 570-578. 30 N. Sillero et al.

Lanza, B., Andreone, F., Bologna, M.A., Corti, C., Razzetti, Sillero, N. (2011): What does ecological modelling model? E., Eds (2007): Fauna d’Italia, Vol. XLII, Amphibia. A proposed classificatio of ecological niche models Calderini, Bologna, Italy. based on their underlying methods. Ecol. Model. 222: Laufer, H., Klemens, F., Sowig, P., Eds (2007): Die Amphi- 1343-1346. bien und Reptilien Baden-Württembergs. Ulmer Verlag, Sillero, N., Celaya, L., Martín-Alfageme, S. (2005): Using Stuttgart. GIS to make an atlas: A proposal to collect, store, map Lescure, J., de Massary, J.C., Eds (2013): Atlas des Amphi- and analyse chorological data for herpetofauna. Rev. biens et Reptiles de France. Muséum National d’Histoire Esp. Herpetol. 19: 87-101. Naturelle. Biotope Eds, Paris. Sillero, N., Brito, J.C., Toxopeus, B., Skidmore, A.K. Longley, P.A., Goodchild, M.F., Maguire, D.J., Rhind, D.W. (2009): Biogeographical patterns derived from remote (2010): Geographical Information Systems and Science, sensing variables: the amphibians and reptiles of the 3rd Edition. John Wiley, New York. Iberian Peninsula. Amphibia-Reptilia 30: 185-206. Loureiro, A., Ferrand, N., Carretero, M.A., Paulo, O. Sillero, N., Amaro, M., Sousa, F., Sousa, P., Gonçalves- (2010): Atlas dos Anfíbios e Répteis de Portugal. Sfera Seco, L. (2014): Distributed database system of the new do Caos, Lisboa. atlas of amphibians and reptiles in Europe: the NA2RE Loureiro, A., Sillero, N. (2010): Metodologia. In: Atlas dos project. Amphibia-Reptilia 35: 33-39. anfíbios e répteis de Portugal, p. 66-74. Loureiro, A., Sindaco, R., Doria, G., Razzetti, E., Bernini, F., Eds (2006): Ferrand, N., Carretero, M.A., Paulo, O., Eds, Esfera do Atlante degli Anfib e dei Rettili d’Italia/Atlas of Italian Caos, Lisboa. Mertens, R., Wermuth, H. (1960): Die Amphibien und Amphibians and Reptiles. Societas Herpetologica Ital- Reptilien Europas. (Dritte Liste, nach dem Stand vom 1. ica. Edizioni Polistampa, Firenze, Italy. Januar 1960). Frankfurt am Main, Verlag Waldemar Sinervo, B., Méndez-de-la-Cruz, F., Miles, D.B., Heulin, B., Kramer. Bastiaans, E., Villagrán-Santa Cruz, M., Lara-Resendiz, Meyer, A., Zumbach, S., Schmidt, B., Monney, J.-C. (2009): R., Martínez-Méndez, N., Calderón-Espinosa, M.L., Les Amphibiens et les Reptiles de Suisse Bern, Switzer- Meza-Lázaro, R.N., Gadsden, H., Avila, L.J., Morando, land. Haupt Verlag. M., De la Riva, I.J., Victoriano Sepulveda, P., Rocha, Oksanen, J., Blanchet, F.G., Kindt, R., Legendre, P., C.F., Ibargüengoytía, N., Aguilar Puntriano, C., Massot, Minchin, P.R., O’Hara, R.B., Simpson, G.L., Solymos, M., Lepetz, V., Oksanen, T.A., Chapple, D.G., Bauer, P., Stevens, M.H.H., Wagner, H. (2012): vegan: Commu- A.M., Branch, W.R., Clobert, J., Sites, J.W. Jr. (2010): nity Ecology Package. R package version 2.0-5. http:// Erosion of lizard diversity by climate change and altered CRAN.R-project.org/package=vegan thermal niches. Science 328: 894-899. Olivero, J., Real, R., Márquez, A.L. (2011): Fuzzy Speybroeck, J., Beukema, W., Crochet, P.A. (2010): A tenta- chorotypes as a conceptual tool to improve insight into tive species list of the European herpetofauna (Amphibia biogeographic patterns. Systc. Biol. 60: 645-660. and Reptilia): an update. Zootaxa 2492: 1-27. Pleguezuelos, J.M., Ed. (1997): Distribución y Biogeografía Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Ro- de los Anfibio y Reptiles en España y Portugal. drigues, A.S.L., Fischman, D.L., Waller, R.W. (2004): Asociación Herpetológica Española – Universidad de Status and trends of amphibian declines and extinctions Granada, Spain. worldwide. Science 306: 1783-1786. Pleguezuelos, J.M., Márquez, R., Lizana, M., Eds (2002): Valakos, E., Pafilis P., Sotiropoulos, K., Lymberakis, P., Atlas y Libro Rojo de los Anfibio y Reptiles de Maragou, P., Foufopoulos, J. (2008): The Amphibians España. Dirección General de la Conservación de la a and Reptiles of Greece. Edition Chimaira, Frankfurt am naturaleza-Asociación Herpetológica Española (2 im- Main. presión). Madrid, Spain. Vences, M., Guayasamin, J.M., Miralles, A., de la Riva, I. Proess, R., Ed. (2003): Verbreitungsatlas der Amphibien (2013): To name or not to name: Criteria to promote des Grossherzogtums Luxemburg. Ferantia 37, Travaux economy of change in Linnaean classificatio schemes. Scientifique du Musée national d’Histoire naturelle, Zootaxa 3636: 201-244. Luxembourg. Vigna Taglianti, A., Audisio, P., Biondi, M., Bologna, M.A., Proess, R., Ed. (2007): Verbreitungsatlas der Reptilien des Carpaneto, G.M., De Biase, A., Fattorini, S., Piattella, Großherzogtums Luxemburg. Ferantia 37, Travaux Sci- entifique du Musée national d’histoire naturelle, Lux- E., Sindaco, R., Venchi, A., Zapparoli, M. (1999): A pro- embourg. posal for a chorotype classificatio of the Near East Puky, M., Schad, P., Szövenyi, G. (2006): Herpetological fauna, in the framework of the Western Palearctic region. atlas of Hungary/Magyarorszag herpetologiai atlasza. Biogeographia 20: 31-59. IUCN SSC Hungary. Wake, D.B., Vredenburg, V.T. (2008): Are we in the midst R Development Core Team (2012): R: a Language and of the sixth mass extinction? A view from the world Environment for Statistical Computing. R Foundation of amphibians. Proc. Natl. Acad. Sci. USA 105: 11466- for Statistical Computing, Vienna, Austria. ISBN 3- 11473. 900051-07-0, URL http://www.R-project.org/. Weiserbs, A., Jacob, J.P. (2005): Amphibiens et Reptiles de Rueda, M., Rodríguez, M.A., Hawkins, B.A. (2010): To- la Région de Bruxelles-Capitale. Aves & Institut Bruxel- wards a biogeographic regionalization of the European lois pour la Gestion de l’Environnement, Bruxelles, Bel- biota. J. Biogeogr. 11: 2067-2076. gium. New atlas of European amphibians and reptiles 31

Wielstra, B., Arntzen, J.W. (2011): Unraveling the rapid ra- Submitted: September 2, 2013. Final revision received: diation of crested newts (Triturus cristatus superspecies) December 23, 2013. Accepted: January 14, 2014. using complete mitogenomic sequences. BMC Evol. Associated Editors: Sylvain Ursenbacher and Sebastian Biol. 11: 162. Steinfartz.