Dear Author,

Here are the proofs of your article.

• You can submit your corrections online, via e-mail or by fax.

• For online submission please insert your corrections in the online correction form. Always indicate the line number to which the correction refers.

• You can also insert your corrections in the proof PDF and email the annotated PDF.

• For fax submission, please ensure that your corrections are clearly legible. Use a fine black pen and write the correction in the margin, not too close to the edge of the page.

• Remember to note the journal title, article number, and your name when sending your response via e-mail or fax.

• Check the metadata sheet to make sure that the header information, especially author names and the corresponding affiliations are correctly shown.

• Check the questions that may have arisen during copy editing and insert your answers/ corrections.

• Check that the text is complete and that all figures, tables and their legends are included. Also check the accuracy of special characters, equations, and electronic supplementary material if applicable. If necessary refer to the Edited manuscript.

• The publication of inaccurate data such as dosages and units can have serious consequences. Please take particular care that all such details are correct.

• Please do not make changes that involve only matters of style. We have generally introduced forms that follow the journal’s style. Substantial changes in content, e.g., new results, corrected values, title and authorship are not allowed without the approval of the responsible editor. In such a case, please contact the Editorial Office and return his/her consent together with the proof.

• If we do not receive your corrections within 48 hours, we will send you a reminder.

• Your article will be published Online First approximately one week after receipt of your corrected proofs. This is the official first publication citable with the DOI. Further changes are, therefore, not possible.

• The printed version will follow in a forthcoming issue.

Please note After online publication, subscribers (personal/institutional) to this journal will have access to the complete article via the DOI using the URL: http://dx.doi.org/[DOI]. If you would like to know when your article has been published online, take advantage of our free alert service. For registration and further information go to: http://www.link.springer.com. Due to the electronic nature of the procedure, the manuscript and the original figures will only be returned to you on special request. When you return your corrections, please inform us if you would like to have these documents returned. Metadata of the article that will be visualized in OnlineFirst

ArticleTitle Defining the importance of landscape metrics for large branchiopod biodiversity and conservation: the case of the Iberian Peninsula and Balearic Islands Article Sub-Title Article CopyRight Springer International Publishing AG (This will be the copyright line in the final PDF) Journal Name Hydrobiologia Corresponding Author Family Name Sala Particle Given Name Jordi Suffix Division GRECO, Institute of Aquatic Ecology Organization University of Girona Address Girona, Spain Phone +34 972 418 466 Fax Email [email protected] URL ORCID http://orcid.org/0000-0002-1227-8566

Author Family Name Gascón Particle Given Name Stéphanie Suffix Division GRECO, Institute of Aquatic Ecology Organization University of Girona Address Girona, Spain Phone Fax Email URL ORCID Author Family Name Cunillera-Montcusí Particle Given Name David Suffix Division GRECO, Institute of Aquatic Ecology Organization University of Girona Address Girona, Spain Phone Fax Email URL ORCID Author Family Name Alonso Particle Given Name Miguel Suffix Division Department of Ecology Organization University of Barcelona Address Barcelona, Spain Phone Fax Email URL ORCID Author Family Name Amat Particle Given Name Francisco Suffix Division Organization Instituto de Acuicultura de Torre de la Sal, CSIC Address Castellón, Ribera de Cabanes, Spain Phone Fax Email URL ORCID Author Family Name Fonseca Particle da Given Name Luis Cancela Suffix Division Laboratório Marítimo da Guia Organization MARE– Marine and Environmental Sciences Centre Address Cascais, Portugal Division CTA - Centro de Ciências e Tecnologias da Água Organization Universidade do Algarve Address Faro, Portugal Phone Fax Email URL ORCID Author Family Name Cristo Particle Given Name Margarida Suffix Division Organization FCT, Universidade do Algarve Address Faro, Portugal Division Organization CCMar, Universidade do Algarve Address Faro, Portugal Phone Fax Email URL ORCID Author Family Name Florencio Particle Given Name Margarita Suffix Division Organization Estación Biológica de Doñana, CSIC Address Seville, Spain Division Departamento de Ecologia, Instituto de Ciências Biológicas Organization Universidade Federal de Goiás Address Goiânia, Goiás, Brazil Phone Fax Email URL ORCID Author Family Name García-de-Lomas Particle Given Name Juan Suffix Division Research Group on Ecology and Dynamics of Aquatic Ecosystems Organization University of Cádiz Address Cádiz, Spain Phone Fax Email URL ORCID Author Family Name Machado Particle Given Name Margarida Suffix Division Organization CCMar, Universidade do Algarve Address Faro, Portugal Phone Fax Email URL ORCID Author Family Name Miracle Particle Given Name Maria Rosa Suffix Division Instituto Cavanilles de Biodiversidad y Biología Evolutiva Organization University of Valencia Address Burjassot, Valencia, Spain Phone Fax Email URL ORCID Author Family Name Miró Particle Given Name Alexandre Suffix Division Integrative Freshwater Ecology Group Organization Center for Advanced Studies of Blanes, Spanish Research Council (CEAB- CSIC) Address Blanes, Girona, Catalonia, Spain Phone Fax Email URL ORCID Author Family Name Pérez-Bote Particle Given Name José Luis Suffix Division Zoology Unit Organization University of Extremadura Address Badajoz, Spain Phone Fax Email URL ORCID Author Family Name Pretus Particle Given Name Joan Lluís Suffix Division Department of Ecology Organization University of Barcelona Address Barcelona, Spain Phone Fax Email URL ORCID Author Family Name Prunier Particle Given Name Florent Suffix Division Organization Asociación de Educación Ambiental El Bosque Animado Address Córdoba, Spain Phone Fax Email URL ORCID Author Family Name Ripoll Particle Given Name Javier Suffix Division Organization Asociación de Educación Ambiental El Bosque Animado Address Córdoba, Spain Phone Fax Email URL ORCID Author Family Name Rueda Particle Given Name Juan Suffix Division Instituto Cavanilles de Biodiversidad y Biología Evolutiva Organization University of Valencia Address Burjassot, Valencia, Spain Division Department of Microbiology and Ecology Organization University of Valencia Address Burjassot, Valencia, Spain Phone Fax Email URL ORCID Author Family Name Sahuquillo Particle Given Name María Suffix Division Instituto Cavanilles de Biodiversidad y Biología Evolutiva Organization University of Valencia Address Burjassot, Valencia, Spain Division Organization Dirección General de Medio Natural, Generalitat Valenciana Address Valencia, Spain Phone Fax Email URL ORCID Author Family Name Serrano Particle Given Name Laura Suffix Division Department of Plant Biology and Ecology Organization University of Sevilla Address Seville, Spain Phone Fax Email URL ORCID Author Family Name Ventura Particle Given Name Marc Suffix Division Integrative Freshwater Ecology Group Organization Center for Advanced Studies of Blanes, Spanish Research Council (CEAB- CSIC) Address Blanes, Girona, Catalonia, Spain Phone Fax Email URL ORCID Author Family Name Verdiell-Cubedo Particle Given Name David Suffix Division Organization Asociación Columbares Address Murcia, Spain Phone Fax Email URL ORCID Author Family Name Boix Particle Given Name Dani Suffix Division GRECO, Institute of Aquatic Ecology Organization University of Girona Address Girona, Spain Phone Fax Email URL ORCID

Received 23 December 2016 Schedule Revised 20 June 2017 Accepted 30 June 2017

Abstract The deficiency in the distributional data of invertebrate taxa is one of the major impediments acting on the bias towards the low awareness of its conservation status. The present study sets a basic framework to understand the large branchiopods distribution in the Iberian Peninsula and Balearic Islands. Since the extensive surveys performed in the late 1980s, no more studies existed updating the information for the whole studied area. The present study fills the gap, gathering together all available information on large branchiopods distribution since 1995, and analysing the effect of human population density and several landscape characteristics on their distribution, taking into consideration different spatial scales (100 m, 1 km and 10 km). In overall, 28 large branchiopod taxa (17 anostracans, 7 notostracans and 4 spinicaudatans) are known to occur in the area. Approximately 30% of the sites hosted multiple species, with a maximum of 6 species. Significant positive co-occurring species pairs were found clustered together, forming 4 different associations of large branchiopod species. In general, species clustered in the same group showed similar responses to analysed landscape characteristics, usually showing a better fit at higher spatial scales. Keywords (separated by '-') Anostraca - Co-occurrences - Land cover - Notostraca - Rarity - Spinicaudata Footnote Information Electronic supplementary material The online version of this article (doi:10.1007/s10750-017-3293-1) contains supplementary material, which is available to authorized users. Guest editors: Federico Marrone, D. Christopher Rogers, Paola Zarattini & Luigi Naselli-Flores / New Challenges in Anostracan Research: a Tribute to Graziella Mura Hydrobiologia DOI 10.1007/s10750-017-3293-1

1 CHALLENGES IN ANOSTRACAN RESEARCH

2 Defining the importance of landscape metrics for large 3 branchiopod biodiversity and conservation: the case 4 of the Iberian Peninsula and Balearic Islands

5 Jordi Sala . Ste´phanie Gasco´n . David Cunillera-Montcusı´ . Miguel Alonso . 6 Francisco Amat . Luis Cancela da Fonseca . Margarida Cristo . 7 Margarita Florencio . Juan Garcı´a-de-Lomas . Margarida Machado . 8 Maria Rosa Miracle . Alexandre Miro´ . Jose´ Luis Pe´rez-Bote .

Author Proof 9 Joan Lluı´s Pretus . Florent Prunier . Javier Ripoll . Juan Rueda . 10 Marı´a Sahuquillo . Laura Serrano . Marc Ventura . David Verdiell-Cubedo . 11 Dani Boix

12 Received: 23 December 2016 / Revised: 20 June 2017 / Accepted: 30 June 2017 PROOFPROOF 13 Ó Springer International Publishing AG 2017

14AQ1 Abstract The deficiency in the distributional data of study fills the gap, gathering together all available 23 15 invertebrate taxa is one of the major impediments information on large branchiopods distribution since 24 16 acting on the bias towards the low awareness of its 1995, and analysing the effect of human population 25 17 conservation status. The present study sets a basic density and several landscape characteristics on their 26 18 framework to understand the large branchiopods distribution, taking into consideration different spatial 27 19 distribution in the Iberian Peninsula and Balearic scales (100 m, 1 km and 10 km). In overall, 28 large 28 20 Islands. Since the extensive surveys performed in the branchiopod taxa (17 anostracans, 7 notostracans and 29 21 late 1980s, no more studies existed updating the 4 spinicaudatans) are known to occur in the area. 30 22 information for the whole studied area. The present Approximately 30% of the sites hosted multiple 31 species, with a maximum of 6 species. Significant 32 positive co-occurring species pairs were found clus- 33 A1 Electronic supplementary material The online version of tered together, forming 4 different associations of 34 A2 this article (doi:10.1007/s10750-017-3293-1) contains supple- A3 mentary material, which is available to authorized users. large branchiopod species. In general, species clus- 35 tered in the same group showed similar responses to 36 A4 Guest editors: Federico Marrone, D. Christopher Rogers, Paola analysed landscape characteristics, usually showing a 37 A5 Zarattini & Luigi Naselli-Flores / New Challenges in better fit at higher spatial scales. 38 A6 Anostracan Research: a Tribute to Graziella Mura

A7 J. Sala (&) Á S. Gasco´n Á D. Cunillera-Montcusı´ Á A18 L. C. da Fonseca A8 D. Boix A19 Laborato´rio Marı´timo da Guia, MARE– Marine and A9 GRECO, Institute of Aquatic Ecology, University of A20 Environmental Sciences Centre, Cascais, Portugal A10 Girona, Girona, Spain A11 e-mail: [email protected] A21 L. C. da Fonseca UNCORRECTEDA22 CTA - Centro de Cieˆncias e Tecnologias da A´ gua, A12 M. Alonso Á J. L. Pretus UNCORRECTEDA23 Universidade do Algarve, Faro, Portugal A13 Department of Ecology, University of Barcelona, A14 Barcelona, Spain A24 M. Cristo A25 FCT, Universidade do Algarve, Faro, Portugal A15 F. Amat A16 Instituto de Acuicultura de Torre de la Sal, CSIC, A26 M. Cristo Á M. Machado A17 Castello´n, Ribera de Cabanes, Spain A27 CCMar, Universidade do Algarve, Faro, Portugal

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

39 Keywords Anostraca Á Co-occurrences Á Land et al., 2016) and national level (e.g. Alonso, 1985; 63 40 cover Á Notostraca Á Rarity Á Spinicaudata Thie´ry, 1987; Defaye et al., 1998; Mura, 1999; Van 64 den Broeck et al., 2015a; Marrone et al., 2016). The 65 effects of environmental factors on the distribution of 66 large branchiopods are largely known, particularly 67 41 Introduction those related to habitat characteristics (e.g. salinity, 68 turbidity, temperature, surface, depth, altitude, vege- 69 42AQ2 Large branchiopods are a key faunal group of tation cover; Alonso, 1998; Boven et al., 2008; 70 43 crustaceans characteristic of temporary aquatic sys- Nhiwatiwa et al., 2011; Gasco´n et al., 2012; Horva´th 71 44 tems (with few exceptions) which can be found across et al., 2013; Sahuquillo & Miracle, 2013; Stoch et al., 72 45 all continents (Brendonck et al., 2008). Although 2016). Moreover, some spatial variables at regional 73 46 nowadays nearly 600 species of large branchiopods level are also known to affect species distribution, 74 47 are known worldwide (Rogers et al., 2015), a large part such as the closeness of sites (which can be related to 75 48 of them are known only from a few localities ([50% of dispersal or shared environmental characteristics; 76 Author Proof 49 the species are known from B10 localities in the case Nhiwatiwa et al., 2011; Horva´th et al., 2013), climatic 77 50 of anostracans; Rogers, 2013), indicating that the gradients (Stoch et al., 2016) or habitat fragmentation 78 51 faunal group harbours a large amount of biodiversity, (Gasco´n et al., 2012). The sensitivity of large 79 52 and that the knowledge of the group is still far from branchiopods to all this range of environmental and 80 53 being complete (Rogers et al., 2015). This is also the spatial variablesPROOFPROOF makes them especially interesting as 81 54 case in the western Mediterranean, where descriptions biological indicators of the conservation status of 82 55 of new species are still being performed (e.g. Alonso temporary wetlands (e.g. Sahuquillo & Miracle, 2015; 83 56 & Garcı´a-de-Lomas, 2009; Korn et al., 2010; Van den Broeck et al., 2015b; Lumbreras et al., 2016). 84 57 Machado & Sala, 2013; Boix et al., 2016) and some However, the effects of changes in land use and the 85 58 efforts to describe the distribution of large bran- fragmentation and loss of habitats have been less 86 59 chiopods are made at regional (e.g. Machado et al., explored on large branchiopods, although these factors 87 60 1999; Boix, 2002; Samraoui & Dumont, 2002; Culioli are recognized as some of the main threats to global 88 61 et al., 2006; Marrone & Mura, 2006; Miracle et al., biodiversity. In the case of the Mediterranean region, 89 62 2008; Prunier & Saldan˜a, 2010; Rodrı´guez-Flores larger biodiversity losses are expected in future 90

A28 M. Florencio A46 F. Prunier Á J. Ripoll A29 Estacio´n Biolo´gica de Don˜ana, CSIC, Seville, Spain A47 Asociacio´n de Educacio´n Ambiental El Bosque Animado, A48 Co´rdoba, Spain A30 M. Florencio A31 Departamento de Ecologia, Instituto de Cieˆncias A49 J. Rueda A32 Biolo´gicas, Universidade Federal de Goia´s, Goiaˆnia, A50 Department of Microbiology and Ecology, University of A33 Goia´s, Brazil A51 Valencia, Burjassot, Valencia, Spain

A34 J. Garcı´a-de-Lomas A52 M. Sahuquillo A35 Research Group on Ecology and Dynamics of Aquatic A53 Direccio´n General de Medio Natural, Generalitat A36 Ecosystems, University of Ca´diz, Ca´diz, Spain A54 Valenciana, Valencia, Spain Á Á A37 M. R. Miracle J. Rueda M. Sahuquillo A55 L. Serrano A38 Instituto Cavanilles de Biodiversidad y Biologı´a A56 Department of Plant Biology and Ecology, University of A39 Evolutiva, University of Valencia, Burjassot, Valencia, A57 Sevilla, Seville, Spain Spain UNCORRECTEDA58 D. Verdiell-Cubedo Á UNCORRECTED A40 A. Miro´ M. Ventura A59 Asociacio´n Columbares, Murcia, Spain A41 Integrative Freshwater Ecology Group, Center for A42 Advanced Studies of Blanes, Spanish Research Council A43 (CEAB-CSIC), Blanes, Girona, Catalonia, Spain

A44 J. L. Pe´rez-Bote A45 Zoology Unit, University of Extremadura, Badajoz, Spain

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

91 scenarios, due to the sensitivity of its ecosystems to branchiopod species distribution. These will set a 140 92 drivers of biodiversity change, especially those related basic framework to understand their distribution and 141 93 to land-use change and introduction of exotic species will allow us to assess whether there are particular 142 94 (Sala et al., 2000; Underwood et al., 2009). Although factors affecting this faunal group that should be taken 143 95 temporary ponds were so far only minimally impacted into consideration in the evaluation of their conserva- 144 96 by traditional agricultural uses, their disappearance tion status. 145 97 and degradation have been highly intensified during 98 the last 50 years, due to infrastructure and urban 99 development, and intensification of agricultural activ- Materials and methods 146 100 ity (Gallego-Ferna´ndez et al., 1999; Rhazi et al., 101 2012). This increasing degradation and loss of habitats Study area 147 102 (Belk, 1998; Zacharias & Zamparas, 2010) strongly 103 impact large branchiopod populations, leading to The Iberian Peninsula and the Balearic Islands are 148 104 greater distances between remaining populations, situated in the western Mediterranean, and cover an 149 Author Proof 105 eventually leading to the loss of local populations area of about 587,000 km2. The area is shared by 150 106 (Eder & Ho¨dl, 2002) and even some species on the several countries (Spain, Portugal, Andorra, France 151 107 regional scale (Martens & De Moor, 1995; Mura, and Gibraltar), although the scope of this study 152 108 1999; De Roeck et al., 2007). Therefore, the landscape focusses only on Spain and Portugal. The climate 153 109 structure surrounding the wetlands should be taken types present inPROOFPROOF the study area are mainly arid and 154 110 into account when evaluating the factors affecting temperate climates, with cold climates circumscribed 155 111 large branchiopods assemblages, since some land- to the higher peaks in the mountain ranges (Peel et al., 156 112 scape characteristics are known to have great impor- 2007; AEMET & IM, 2011). Arid climate is restricted 157 113 tance for biotic communities, such as landscape mostly to southeastern Spain, although there are some 158 114 heterogeneity or land-use types, acting at several areas in northeastern (Ebro valley), central (southern 159 115 spatial scales (Weibull et al., 2000; Hall et al., 2004; Meseta and Extremadura) and southwestern (Alen- 160 116 Hartel & von Wehrden, 2013). The evaluation of these tejo) Iberian Peninsula, and Balearic Islands (southern 161 117 landscape characteristics for the conservation of the Majorca, and the islands of Ibiza and Formentera) with 162 118 large branchiopod assemblages has been seldomly this type of climate. Temperate climates are divided 163 119 carried out, but the type of land use is known to affect into those with dry summer (which comprises the 164 120 assemblage structure of passive dispersers (Hall et al., majority of the area of the Iberian Peninsula, from the 165 121 2004), as well as some species at different spatial western Atlantic coast to the Mediterranean sea, 166 122 scales (Angeler et al., 2008), whereas for other excluding the arid southeastern Spain) or without dry 167 123 species, the land use surrounding the wetland was season (restricted to the northern Iberian Peninsula– 168 124 shown to be irrelevant (Angeler et al., 2008; Horva´th Cantabrian coast, northern Meseta and the Iberian and 169 125 et al., 2013). To this end, the integration of landscape Pyrenees mountain ranges). 170 126 characteristics will improve large branchiopod con- 127 servation programs at regional and national level, and Species distribution 171 128 they will help to better understand the drivers of 129 biodiversity change linked to future scenarios of All known bibliographical and unpublished large 172 130 global change. branchiopod records from the Iberian Peninsula and 173 131 The main goal of this article is to describe the Balearic Islands were compiled (dating back to 1916), 174 132 overall biodiversity patterns of the large branchiopods and geographically referenced to the maximum pos- 175 133 present in the Iberian PeninsulaUNCORRECTED and Balearic Islands, sible resolution whenever possible (see Online 176 134 establishing a general overviewUNCORRECTED for the conservation of Resource 1 for bibliographical references). This 177 135 this faunal group in this area. Additionally, we aim to allowed to create the checklist of the large bran- 178 136 determine if the biodiversity patterns detected are chiopod fauna, and the global distribution in the 179 137 similar across the entire study area, and to investigate Iberian Peninsula and Balearic Islands (see Online 180 138 the role of land use, landscape structure, population Resource 2). However, as it is unreliable to relate old 181 139 density and protected area networks on the large records to present-day landscape metrics, all analyses 182 123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

183 were limited to records from 1995 onwards. Moreover, of sites that showed co-occurring species. Seventy- 230 184 some records were not considered for the study of the seven UTM 100 km grid squares had presence of large 231 185 biodiversity patterns due to particularities in the branchiopods in the Iberian Peninsula and Balearic 232 186 biology or in the taxonomic resolution of different Islands, but only those with more than 5 sites were 233 187 taxa. Thus, the exotic invasive Artemia franciscana included in the analyses (39 out of 77). 234 188 Kellogg, 1906 was not considered as this species 189 competitively displaces the autochthonous species of Landscape metrics 235 190 Artemia. Records of parthenogenetic strains of 191 Artemia with undefined ploidy were also not consid- Landscape metrics were calculated considering 236 192 ered, because parthenogenetic strains with different human population density (DP, based on local admin- 237 193 ploidies (i.e. diploid and tetraploid) were considered istrative units), protection status (N2000, Natura 2000 238 194 as two different taxa, as recent studies suggest that network areas), land cover (CORINE land cover) and 239 195 they could be related to different bisexual Artemia landscape structure. We assigned to each site the 240 196 species (Mun˜oz et al., 2010; Asem et al., 2016). In the corresponding population density of the local admin- 241 Author Proof 197 case of genus Triops, we considered all species of the istrative units (LAUs) where it was located. LAUs, as 242 198 genus as one taxon, due to the difficulties to allocate well as their corresponding population, were obtained 243 199 the bibliographical records of this genus to the 6 from 2011 population census (European Statistical 244 200 existing species present in the Iberian Peninsula (Korn System, 2011). Natura 2000 network areas (EEA, 245 201 et al., 2006, 2010); at this moment, syntopic occur- 2015) were usedPROOFPROOF as the protection category for the 246 202 rences of different species of Triops have not been studied region instead of other national protected areas 247 203 recorded in the Iberian Peninsula or the Balearic due to its high representativeness of biodiversity 248 204 Islands. (Rosati et al., 2008; Abella´n&Sa´nchez-Ferna´ndez, 249 205 For each species, the number of occurrences in the 2015). Sites that were in a Natura 2000 network area 250 206 study area allowed to determine their rarity with rank– were classified as Protected. Land covers were clas- 251 207 frequency curves following Siqueira et al. (2012). The sified in 6 categories based on CORINE Land Cover 252 208 inflection point of the curve classified the species as inventories (EEA, 2006): artificial surfaces (AS), 253 209 common (those at the left side of the curve) or rare forests and semi-natural areas (FS), irrigated agricul- 254 210 (those at the right side of the curve; in this case those tural areas (IA), non-irrigated agricultural areas (nIA), 255 211 that were present in less than 50 sites; see Online wetland areas (W) and water bodies, although this last 256 212 Resource 3). Moreover, those species that contributed category was not used to test its effects on the biotic 257 213 with less than 1% to the total occurrences were assemblages. Then, for each site, we calculated the 258 214 considered very rare (in this case, those that were relative percentage of surface occupied by each 259 215 present in less than 15 sites). category using 3 different spatial scales (circular areas 260 216 Biodiversity metrics were calculated using two were constructed using 100 m, 1 km and 10 km radius 261 217 different approaches. In one hand, for each site species distances to the central point of each site as centroid). 262 218 richness and composition was recorded in order to We used ArcMap GIS 10.0 to process all datasets and 263 219 describe biodiversity patterns across the whole study calculate landscape metrics (ESRI, 1999). Finally, 264 220 area. We considered co-occurring species as those following Weibull et al. (2000), landscape diversity 265 221 species that were recorded in the same site, but not (Ldiv) was calculated as a measure of landscape 266 222 necessarily at the same time (i.e. syntopic species). structure using the Shannon diversity index, 267 223 However, data on synchronic species (i.e. those Ldiv =-pi ln(pi), where pi is the proportion of 268 224 species that were recorded at the same time in the habitat i in the circular areas used for calculating the 269 225 same site) appear in OnlineUNCORRECTED Resource 4. On the other land covers. In the case of the UTM grid square 270 226 hand, for each UTM 100UNCORRECTED km grid square we calculated approach, we calculated the mean values of the 271 227 the number of sites with presence of large bran- relative percentage of surface occupied by each 272 228 chiopods, the mean species richness per site, the total category of land covers and landscape diversity for 273 229 species richness in each grid square and the percentage all the sites present at each grid square. 274

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

275 Data analysis (Logan, 2010). In order to evaluate whether the 319 magnitude of the contribution of a variable is signif- 320 276 Species associations icant (Z C 1.65 at the 95% level), a randomization 321 procedure was used in which the independent contri- 322 277 Species associations were explored using two different butions of each predictor variable were compared to 323 278 approaches: (1) co-occurrence analysis and (2) clas- distributions of such contributions generated by 324 279 sification analysis. The co-occurrence analysis repeated randomizations of the data matrix (number 325 280 allowed us to identify significant co-occurring species, of randomizations = 100). While a binomial family 326 281 either with positive or negative co-occurrences. error was specified for the species hierarchical parti- 327 282 Moreover, the effect size (i.e. the difference between tioning and randomization procedures, a Poisson 328 283 expected and observed frequency of co-occurrence) family error was used for species richness analyses. 329 284 was also obtained in order to quantify the strength of When over- or under-dispersion was detected, P values 330 285 the pattern. In the second approach, a non-hierarchical were corrected using quasi-likelihood model (Logan, 331 286 cluster approach was used to identify species assem- 2010). The sign of the relationship between species 332 Author Proof 287 blages. Partitioning around medoids (PAM) was the presence–absence or richness data and landscape 333 288 classification technique chosen to cluster species into metrics was assessed with generalized linear models 334 289 groups, since it has been pointed out as more robust (GLM) using all environmental variables as predic- 335 290 than more classical approaches like k-means cluster- tors. To assess the model goodness-of-fit, we 336 291 ing (Borcard et al., 2011). Sørensen similarity was employed the squaredPROOFPROOF Pearson correlation coefficient 337 292 used as the similarity measure. The number of groups between observed and predicted values. Hierarchical 338 293 tested representing existing species assemblages was partitioning and randomizations were carried out 339 294 from 2 to 16 (number of large branchiopod species using package ‘‘hier.part’’ (Walsh & Mac Nally, 340 295 minus 1, only considering the species with more than 5 2013). 341 296 occurrences). Silhouette widths of the resulting groups 297 were obtained to examine group membership’s appro- Biodiversity and landscape metrics 342 298 priateness, as well as the overall classification result 299 using the Rousseeuw quality index. The silhouette Conditional inference tree (CIT) models were used to 343 300 width ranges between -1 and 1, and it is a measure of assess the relationship between biodiversity (response 344 301 the degree of membership of an object to its cluster, variables: mean species richness per site, the total 345 302 negative values indicating misclassified objects. Co- species richness in each UTM 100 km grid square and 346 303 occurrence analysis was performed using package the percentage of sites with co-occurrences) and 347 304 ‘‘cooccur’’ (Griffith et al., 2016), and PAM analysis landscape metrics (explanatory variables: DP, 348 305 was performed using package ‘‘cluster’’ (Maechler N2000, AS, FS, IA, nIA, W, Ldiv) in the 3 tested 349 306 et al., 2016). spatial scales (100 m, 1 km and 10 km radius). This 350 type of regression displays a binary tree, built by a 351 307 Species and landscape metrics process known as binary recursive partitioning, which 352 gives a very clear picture of the structure of the data, 353 308 Hierarchical partitioning was used to identify the and provides a highly intuitive insight into the kinds of 354 309 relationship’s sign (positive or negative) and the interactions between variables (Crawley, 2002). CIT 355 310 relative magnitude of the effects of the landscape are not affected by over-fitting and are unbiased with 356 311 metrics tested at different scales (areas of 100 m, 1 km regard to the types of explanatory variables used 357 312 and 10 km radius). This method assesses the indepen- (Hothorn et al., 2006; Strobl et al., 2007). CIT models 358 313 dent, joint and total contributionUNCORRECTED (relative influence) of were developed using the ‘‘party’’ package for R 359 314 each predictor variableUNCORRECTED by averaging a measure of (Hothorn et al., 2006). Three models, one for each 360 315 goodness-of-fit over all possible models that include biodiversity metric, were run with the different spatial 361 316 that predictor variable. Therefore, it is less susceptible scales. 362 317 to multicollinearity problems than are the single- All statistical analyses were run using R 3.0.1 (R 363 318 model approaches when looking for the ‘‘best’’ model Core Team, 2013). 364

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

365 Results 1), corresponding to 1167 different sites (i.e. water 369 bodies). One hundred seventy sites had only records 370 366 In total, 1808 bibliographical and unpublished records before 1995 (Fig. 1), and were used only for the 371 367 of large branchiopods in the Iberian Peninsula and the overall distribution of species (see Online Resource 2 372 368 Balearic Islands were obtained (see Online Resource for distribution maps with all records), leaving 997 373

Records before 1995 Records from 1995 onwards Artemia salina part. Artemia (diploid) Author Proof

PROOFPROOF part. Artemia (tetraploid) Artemia franciscana Branchinecta ferox Branchinecta orientalis

Branchinectella media Branchipus cortesi Branchipus schaefferi Chirocephalus diaphanus

Cyzicus grubei Lepidurus apus Leptestheria mayeti Linderiella baetica

Linderiella sp. Maghrebestheria maroccana Phallocryptus spinosus Streptocephalus torvicornis UNCORRECTEDUNCORRECTED

Tanymastix stagnalis Tanymastix sp. Tanymastigites lusitanica Triops spp.

Fig. 1 Distribution maps of all records of large branchiopods before 1995, of all records from 1995 onwards, and of the different large branchiopod taxa present in Iberian Peninsula and Balearic Islands from 1995 onwards 123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

A B

no. sites < 11 mean S < 1.15

31

Author Proof CD

PROOFPROOF

total S < 3 % sites < 15

Fig. 2 Geographical patterns of large branchiopod assem- in each UTM 100 km grid square. D Percentage of sites with blages. A Number of sites within each UTM 100 km grid more than 1 species in each UTM 100 km grid square. In all square. B Mean species richness (S) per site recorded in each panels, only UTM grid squares with more than five sites are UTM 100 km grid square. C Total species richness (S) recorded shown

374 sites to use for analyses. The records were scattered all with two known sites; Korn et al., 2010). Fifteen taxa 390 375 across the Iberian Peninsula and the Balearic Islands, can be considered rare (\50 sites); within this group, 391 376 but they were more frequent in the southern regions 11 taxa are extremely rare, being in less than 15 sites 392 377 (Andalusia, Alentejo and Algarve) than in the central (Table 1). To our knowledge, Cyzicus tetracerus 393 378 and northern ones, with the exception of Catalonia (Krynicki, 1830) is the only taxon that has not been 394 379 (Fig. 2A). There are still some regions (Navarra, captured again since 1983 (Alonso, 1996), so it was 395 380 Cantabria and several northern Portuguese districts) not present in further analyses. The majority of the 396 381 without records of large branchiopods. taxa are considered as Trans-Iberian species with wide 397 382 Twenty-eight taxa have been recorded to date in the distributions across the western Palaearctic, although 398 383 Iberian Peninsula and Balearic Islands: 17 anostracans there are also a large number of Iberian endemic 399 384 (one of them the invasiveUNCORRECTEDUNCORRECTED exotic Artemia franciscana), species (10 species, of which 4 can be considered as 400 385 7 notostracans and 4 spinicaudatans. Seven taxa can be extremely rare). Spinicaudatans, notostracans and 401 386 considered common ([50 sites) (Table 1), although extremely rare anostracans tended to co-occur fre- 402 387 we considered all Triops spp. together, and within this quently with other species, with the exception of 403 388 group there are some species with restricted distribu- Tanymastigites lusitanica Machado & Sala, 2013, 404 389 tions (e.g. T. emeritensis Korn & Pe´rez-Bote, 2010, usually found alone (Table 1). Approximately 30% of 405

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

Table 1 Taxa of large branchiopods found in the Iberian Peninsula and Balearic Islands from 1995 onwards in decreasing order of occurrence Taxon Abbreviation No. sites Rarity General Co-occurrences distribution with other spp. (%)

Chirocephalus diaphanus Pre´vost, 1803 CHR_DIA 377 C T 38.2 Branchipus schaefferi Fischer, 1834 BCH_SCH 293 C T 22.9 Triops spp. TRP_SPP 241 C – 79.7 Tanymastix stagnalis (L., 1758) TAN_STA 169 C N 64.5 Branchipus cortesi Alonso & Jaume, 1991 BCH_COR 111 C EN 77.5 Streptocephalus torvicornis (Waga, 1842) STR_TOR 62 C T 71.0 Cyzicus grubei (Simon, 1886) CYZ_GRU 60 C EN 95.0 Maghrebestheria maroccana Thie´ry, 1988 MAG_MAR 23 R S 100.0

Author Proof Artemia salina (L., 1758) ART_SAL 20 R T 25.0 Branchinectella media (Schmankewitsch, 1873) BRT_MED 18 R T 22.2 Artemia franciscana Kellogg, 1906 ART_FRA 16 – EX – Tanymastix sp. TAN_SP 15 R EN 46.7 Phallocryptus spinosus (Milne-Edwards, 1840) PHC_SPI 9PROOFPROOF RR T 44.4 Tanymastigites lusitanica Machado & Sala, 2013 TAG_LUS 9 RR EN 11.1 parthenogenetic Artemia (tetraploid) ART_PAT 8 RR T 12.5 Branchinecta orientalis Sars, 1901 BRN_ORI 8 RR T 50.0 parthenogenetic Artemia (diploid) ART_PAD 7 RR T 57.1 Lepidurus apus (L., 1758) LEP_APU 7 RR T 14.3 Leptestheria mayeti (Simon, 1885) LPT_MAY 4 RR M 100.0 Branchinecta ferox (Milne-Edwards, 1840) BRN_FER 3 RR T 66.7 Linderiella sp. LIN_SP 3 RR EN 100.0 Linderiella baetica Alonso & Garcı´a-de-Lomas, 2009 LIN_BAE 1 RR EN 100.0 Cyzicus tetracerus (Krynicki, 1830) CYZ_TET 0 RR T – For each taxon, number of sites, rarity class, general distribution in the study area (adapted from Abella´n et al., 2005) and percentage of sites where each taxon co-occurred with any other taxa. Note that Triops spp. is a heterogeneous group that includes 6 species, 4 of them endemic from the Iberian Peninsula (1 of them extremely rare) C common, EN endemic species, EX exotic species, M Mediterranean species, N northern species (present in Europe and the Iberian Peninsula, but not in north Africa), R rare, RR extremely rare, S southern species (present in north Africa and the Iberian Peninsula, but not in Europe north of the Pyrenees), T Trans-Iberian species (present in Europe, the Iberian Peninsula and north Africa)

406 sites presented co-occurrence of large branchiopod Number of sites with large branchiopods 418 407 species, with decreasing site frequencies when number (mean = 23.3; range 6–125; Fig. 2A) had high values 419 408 of co-occurring species increased (see Online in SW and NE Iberian Peninsula, similarly to the 420 409 Resource 4). A maximum of 6 species were found pattern found for the total species richness per UTM 421 410 co-occurring together in 2 different sites in SW Iberian grid square (mean = 4.6; range 1–11; Fig. 2C). In the 422 411 Peninsula (in Andalusia and Algarve), followed by 5 case of the mean species richness per site 423 412 species found in 7 sitesUNCORRECTEDUNCORRECTED (Andalusia, Extremadura and (mean = 1.4; range 1.0–2.2; Fig. 2B), high values 424 413 Alentejo) and 4 species found in 34 sites (Andalusia, were also more common in SW Iberian Peninsula, but 425 414 Extremadura, Algarve, Alentejo and Valencia). there were also some UTM grid squares with high 426 415 The UTM 100 km grid square approach showed values in the eastern part of the Peninsula and in the 427 416 that the general biodiversity patterns were geograph- Balearic Islands. The percentage of sites with co- 428 417 ically heterogeneous in the study area (Fig. 2). occurrences within a UTM grid square (mean = 26.8; 429

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

Fig. 3 Pairwise combinations of large branchiopod species in the Iberian Peninsula and Balearic Islands determined by probabilistic co- occurrence analysis. Circles indicate significant co- occurrences, and the size of the circles represents the effect sizes of the association. Only significant co-occurrences are shown. White circles in white background positive co- occurrences; black circles in grey background negative

Author Proof co-occurrences. Abbreviations of species are the same as in Table 1 PROOFPROOF

430 range 0.0–67.4; Fig. 2D) was especially high in SW, and the diploid parthenogenetic Artemia), group 3 451 431 whereas lower values were observed in the NE and (including B. cortesi, T. stagnalis and Maghrebesthe- 452 432 some areas in the SE Iberian Peninsula. ria maroccana Thie´ry, 1988), group 4 (including 453 433 The probabilistic co-occurrences analysis showed Streptocephalus torvicornis (Waga, 1842), Triops 454 434 some species pairs with significant positive associa- spp., C. grubei, C. diaphanus and B. schaefferi) and 455 435 tions (e.g. Branchipus cortesi Alonso & Jaume, 1991 group 6 [including the halophilous Branchinectella 456 436 and Tanymastix stagnalis (L., 1758); Cyzicus grubei media (Schmankewitsch, 1873) and Phallocryptus 457 437 (Simon, 1886) and Triops spp.; Fig. 3), whereas other spinosus (Milne-Edwards, 1840)]. Group 1 had the 458 438 pairs had significant negative co-occurrences (e.g. highest ASW, whereas group 4 had the lowest ASW, 459 439 Branchipus schaefferi Fischer, 1834 and Chiro- pointing that the species in this cluster were not tightly 460 440 cephalus diaphanus Pre´vost, 1803; T. stagnalis and grouped. 461 441 B. schaefferi; C. diaphanus and T. stagnalis; Fig. 3). Overall, hierarchical variation partitioning showed 462 442 The partitioning around medoids (PAM) analysis significant effects of landscape metrics on large 463 443 obtained similar results, clustering together the species branchiopods at several spatial scales. In general, the 464 444 that showed positive co-occurrences (Fig. 4). How- best fit was observed at the largest spatial scale 465 445 ever, the global averageUNCORRECTEDUNCORRECTED silhouette width (ASW; a (10 km). In the case of species richness, although the 466 446 measure of how well the species are clustered) was proportion of the variation explained was low, the 467 447 quite low, indicating that in general the affinity among presence of non-irrigated agriculture (nIA) revealed 468 448 the species in each cluster was not very strong. PAM positive relationships at lower spatial scales (100 m 469 449 analysis classified the species in four significant and 1 km), whereas forests and semi-natural vegeta- 470 450 groups: group 1 (including Artemia salina (L., 1758) tion (FS) showed significant positive relationship at 471

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

472 the lowest spatial scale (100 m) but negative relation- Fig. 5 Plots showing species richness (S) and species groups ofc 473 ships at larger scales (Fig. 5). At the largest spatial PAM analysis (groups 1, 3, 4, 6 and the rest of species not clustered in any group of PAM analysis) in the three spatial 474 scale, presence of wetlands and landscape diversity scales (100 m, 1 km and 10 km). The first column shows the fit 475 (Ldiv) showed a positive relationship, and population of the binomial models calculated as the Pearson’s correlation 476 density showed a negative relationship. between observed and expected values (white bars), whereas the 477 In the case of group 1 of PAM analysis, the rest of columns show the independent contribution of the landscape metrics obtained from the hierarchical partitioning 478 proportion of the variation explained was relatively (grey bars indicate positive effects while black bars indicate 479 high, especially for the diploid parthenogenetic negative effects). Only significant relationships detected after 480 Artemia. Wetlands (W) and population density (DP) the randomization process are shown (see ‘‘Materials and 481 had significant positive relationships on the distribu- methods’’ section for more details). Abbreviations of species are the same as in Table 1, and those of landscape metrics appear in 482 tion of both taxa at different scales, whereas the rest of Materials and methods 483 metrics had low relation (Fig. 5). For group 3 of PAM 484 analysis, composed by an assemblage characteristic of 485 the SW Iberian Peninsula, protected areas (N2000) some species was defined by being outside of 496 Author Proof 486 and Ldiv (especially at large spatial scales) showed protected areas (C. diaphanus, S. torvicornis and 497 487 significant positive relationships on the distribution of Triops spp.). Interestingly, B. schaefferi was positively 498 488 B. cortesi and T. stagnalis. FS and nIA showed related to population density (DP), artificial surfaces 499 489 different effects depending of the spatial scale anal- (AS), irrigated agriculture (IA), FS and nIA, whereas 500 490 ysed, being negative at larger scales and positive at ChirocephalusPROOFPROOF diaphanus was negatively related to 501 491 smaller ones (Fig. 5). In contrast, the group 4 of PAM DP, IA and Ldiv. The group 6 of PAM analysis was 502 492 analysis (Fig. 5) was characterized by significant composed of Branchinectella media and Phallocryp- 503 493 positive relationships of nIA, especially at small tus spinosus, two characteristic species of saline 504 494 spatial scales, although the proportion of the variation endorrheic wetlands, and their distribution were 505 495 explained was relatively low. Also, the distribution of explained by being positively related to N2000, Ldiv 506

Fig. 4 Silhouette plots showing the partition around medoids (PAM) analysis results. Different colours indicate different groups. Average silhouette width of the overall analysis is indicated at the bottom of the plot. j stands for the number of the cluster; nj indicates the number of species within each cluster j; aveieCj si corresponds to the average silhouette width for each cluster. Abbreviations of species are the same as in Table 1

UNCORRECTEDUNCORRECTED

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia GROUP 1 GROUP 3 Author Proof

PROOFPROOF GROUP 4 GROUP 6

UNCORRECTEDUNCORRECTED

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

507 (mainly at small spatial scales), nIA and W (both at exception of the tetraploid parthenogenetic Artemia 518 508 large spatial scales), while FS had negative significant (Fig. 5). 519 509 relationship on them (Fig. 5). The rest of the species Wetland and non-irrigated agriculture land covers 520 510 formed a heterogeneous group of extremely rare were the only factors that were related to the 521 511 species that did not cluster significantly in PAM biodiversity metrics at the UTM 100 km grid square 522 512 analysis. Therefore, there was not a common pattern in approach. Significantly higher values of mean species 523 513 the landscape metrics affecting the species distribu- richness per site at a spatial scale of 1 km radius 524 514 tions, although DP had negative significant relation- (Fig. 6A) were found in UTM grid squares with high 525 515 ships on the distribution of all species except values of wetland land cover (more than 6.9 ha), and 526 516 Lepidurus apus (L., 1758), whereas nIA had positive also in non-irrigated agriculture land cover (more than 527 517 significant relationships at larger scales with the 177.4 ha). Similar results were obtained for total 528 Author Proof

PROOFPROOF

UNCORRECTEDUNCORRECTED

Fig. 6 Significant results of the conditional inference trees C Total species richness per UTM grid square at a spatial scale between biodiversity and landscape metrics for each UTM of 10 km radius. D Percentage of sites with co-occurrences at a 100 km grid square at each spatial scale. A Mean species spatial scale of 1 km radius. Abbreviations of landscape metrics richness per site at a spatial scale of 1 km radius. B Total species appear in ‘‘Materials and methods’’ section richness per UTM grid square at a spatial scale of 1 km radius. 123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

529 species richness per UTM grid square at a spatial scale Triops had high occurrences, but paradoxically it is 575 530 of 10 km radius (Fig. 6C), and for the percentage of one of the taxa that gathers more uncertainties about 576 531 sites with co-occurrences at a spatial scale of 1 km its distribution due to recent taxonomical modifica- 577 532 radius (Fig. 6D). In the case of total species richness tions (until 2006 it was considered as only one species, 578 533 per UTM grid square at a spatial scale of 1 km radius T. cancriformis (Bosc, 1801); Korn et al., 2006, 2010), 579 534 (Fig. 6B), significantly higher values were found only that difficult the allocation of all previous biblio- 580 535 in UTM grid squares with high values of wetland land graphical records to the 6 species present in the Iberian 581 536 cover (more than 2.4 ha). Peninsula. A detailed study of the identities, distribu- 582 tions and conservation status of the genus Triops in the 583 Iberian Peninsula (especially in the northern and 584 537 Discussion eastern regions) and Balearic Islands is therefore 585 strongly recommended. For the extremely rare 586 538 This work represents an exhaustive update of the species, monitoring and conservation programs should 587 539 distribution and the biodiversity patterns of the large be mandatory, due to their low occurrences, in some 588 Author Proof 540 branchiopod fauna of the Iberian Peninsula and cases with the major part of their localities outside 589 541 Balearic Islands, similarly to other studies in neigh- protected areas. A paradigmatic case is the anostracan 590 542 bouring countries (e.g. Defaye et al., 1998; Mura, Linderiella baetica Alonso & Garcı´a-de-Lomas, 2009, 591 543 1999; Van den Broeck et al., 2015a; Marrone et al., of which the only known remaining population is 592 544 2016). Since the major works by Alonso critically endangeredPROOFPROOF by urbanization (Garcı´ade 593 545 (1985, 1996, 1998) on Iberian and Balearic large Lomas et al., 2016), and that urgent measures are 594 546 branchiopods, six new species were described (L. needed to avoid extinction. However, surveillance and 595 547 baetica, Tanymastigites lusitanica, Triops baeticus monitoring programs should be also regarded for not 596 548 Korn, 2010, T. gadensis Korn & Garcı´a-de-Lomas, so rare species, because their populations could be 597 549 2010, T. vicentinus Korn, Machado, Cristo & Cancela declining rapidly, even at supranational scales. This is 598 550 da Fonseca, 2010 and T. emeritensis; Alonso & the case of the autochthonous species of genus 599 551 Garcı´a-de-Lomas, 2009; Korn et al., 2010; Machado Artemia, which are also of great concern. The 600 552 & Sala, 2013), 2 more are currently being described abandonment of the salterns, together with the expan- 601 553 (Linderiella sp. and Tanymastix sp.), 1 is new for the sion of the invasive A. franciscana, are leading to the 602 554 Iberian Peninsula (Triops simplex Ghigi, 1921; Korn decline and loss of populations of native species 603 555 et al., 2010) and an exotic species was recorded all through competition and displacement (Amat et al., 604 556 along the Peninsula (Artemia franciscana; Amat et al., 2007; Redo´n et al., 2015), and there is an urgent need 605 557 2005). To date, the number of species found in the to update the distributional data and to assess the 606 558 study area, 28 species, is similar to neighbouring conservation status of the Mediterranean brine 607 559 countries (e.g. 22 species in Morocco (Thie´ry, 1986; shrimps at national level or higher in order to evaluate 608 560 Amat et al., 2005; Boix et al., 2016; Korn & the extent of damage to their populations (Mura et al., 609 561 Hundsdoerfer, 2016), and 25 in Italy (Cottarelli & 2006). 610 562 Mura, 1983; Mura, 1999; Marrone & Mura, 2006; Four large branchiopod associations were detected 611 563 Mura et al., 2006; Scanabissi et al., 2006; Alfonso, which corresponded in part to those already described 612 564 2017)) and slightly higher than others [e.g. 19 species by Alonso (1998). Two associations were character- 613 565 both in Tunisia (Ben Naceur et al., 2013; Marrone istic of saline habitats: group 1 is a characteristic 614 566 et al., 2016) and Algeria (Samraoui & Dumont, 2002; association with Artemia present mainly in salterns, 615 567 Samraoui et al., 2006; Ghomari et al., 2011), and 18 whereas group 6 contained the halophile species 616 568 species in France (NourissonUNCORRECTED & Thie´ry, 1988; Defaye Phallocryptus spinosus and Branchinectella media, 617 569 et al., 1998; Amat et al.,UNCORRECTED2005; Rabet et al., 2005)]. The which is also present in other areas with steppic 618 570 taxa with high occurrences were species common in habitats (e.g. Samraoui et al., 2006; Marrone et al., 619 571 the western Palaearctic (C. diaphanus, B. schaefferi, T, 2016). The other two associations were characteristic 620 572 stagnalis, S. torvicornis), together with two Iberian of freshwater temporary waterbodies. Group 3 is a 621 573 endemics (B. cortesi, C. grubei) widely distributed characteristic association from the SW Iberian Penin- 622 574 mainly in the western Peninsula. Also, the genus sula with B. cortesi, T. stagnalis and sometimes M. 623 123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

624 maroccana, partly described in Cancela da Fonseca that the extent of wet areas or the pond density also 673 625 et al. (2008). This association is partly similar to the influences invertebrate and plant assemblages (e.g. 674 626 one observed in Morocco, where M. maroccana Boix et al., 2008; Bagella et al., 2010). It is interesting 675 627 usually also appears with Tanymastix affinis Daday, to note that the majority of the relationships had a 676 628 1910 and sometimes with B. schaefferi (Thie´ry, 1991; better fit at larger spatial scales, in accordance with the 677 629 Van den Broeck et al., 2015a). Group 4 corresponds to results found by Angeler et al. (2008), suggesting that 678 630 an association that coincides partly with the one management at larger scales have also to be taken into 679 631 described by Alonso (1998), but probably corresponds account for the conservation of large branchiopods. 680 632 to two different groups due to the fact that several Moreover, biodiversity metrics were influenced by 681 633 species of genus Triops are included in this group. In the presence of wetlands and/or land use linked to non- 682 634 this sense, an association with B. schaefferi, Triops irrigated agricultural practices, especially at larger 683 635 spp. and sometimes S. torvicornis, is present in the arid scales. Our results showed that wetlands, floodplains 684 636 climates of the north and the east of the Peninsula, and and surrounding areas in the Iberian Peninsula (e.g. 685 637 in the Balearic Islands, whereas an association with C. Guadiana, Don˜ana, Ca´diz and Emporda` wetlands) 686 Author Proof 638 diaphanus, C. grubei and Triops spp. is characteristic presented high values of large branchiopod species 687 639 of the south of the Iberian Peninsula. richness at different spatial scales, probably due to the 688 640 The necessity of increasing our knowledge on large heterogeneity of the water body types that could host 689 641 branchiopod ecology should not be restricted only to different communities. Moreover, these areas were 690 642 species or population level, but they should also also importantPROOF forPROOF the presence of rare and/or Iberian 691 643 encompass higher levels of organization. The func- endemic species. The importance of this kind of 692 644 tioning of biodiversity patterns at different spatial habitats for the biodiversity of large branchiopods has 693 645 scales must be taken into account for improving also been observed in other regions (e.g. Eder et al., 694 646 conservation actions, because disturbances are scale 1997; Timms & Sanders, 2002; Waterkeyn et al., 695 647 dependent (Caro, 2010). In general, our study detected 2009; Nhiwatiwa et al., 2014; Stoch et al., 2016). 696 648 that land use had an effect on the species richness and However, land-use analyses also revealed that tempo- 697 649 distribution of large branchiopods. However, not all rary water bodies in non-irrigated agricultural land- 698 650 the species were affected by the same landscape scapes not linked directly to large wetland areas also 699 651 metrics. These results are in concordance with those hosted a large amount of biodiversity. Traditional, 700 652 observed by Angeler et al. (2008), with T. cancri- non-intensive agricultural practices are compatible 701 653 formis being affected by irrigated agriculture at large with the existence of highly biodiverse temporary 702 654 spatial scale, whereas Branchinecta orientalis was not. ponds (e.g. Grillas et al., 2004; Robson & Clay, 2005), 703 655 In contrast, other studies found no effects of land use even in areas where temporary ponds are human-made 704 656 on large branchiopod communities (e.g. Horva´th et al., and the presence of natural temporary water bodies is 705 657 2013; Mabidi et al., 2016). We found that species very rare (e.g. Verdiell-Cubedo & Boix, 2014). The 706 658 clustered together were similarly affected by the same large branchiopod fauna associated to these habitats is 707 659 landscape metrics, but that different clusters were usually very rich, and they host multiple species (e.g. 708 660 affected by different landscape metrics, which could Boven et al., 2008; Cancela da Fonseca et al., 2008; 709 661 partially explain those contrasting results. Agricultural Prunier & Saldan˜a, 2010; Marrone et al., 2016; 710 662 (irrigated and non-irrigated) and wetland land covers Alfonso, 2017). However, these habitats are rapidly 711 663 were the metrics that affected to more species (positive declining due to intensification of agricultural prac- 712 664 or negative, and with different magnitudes). It is tices, changes in land use and population growth (e.g. 713 665 known that agricultural land use around temporary Underwood et al., 2009; Rhazi et al., 2012), affecting 714 666 ponds can have importantUNCORRECTED effects on their biota (Rhazi the biotic populations linked to these habitats (e.g. 715 667 et al., 2001; O’Neill etUNCORRECTED al., 2016), although traditional Euliss Jr. & Mushet, 1999; Beja & Alcazar, 2003; Van 716 668 agricultural practices are a key factor for temporary den Broeck et al., 2015a). 717 669 pond conservation (Bagella et al., 2016). This is in Since the extensive surveys performed by Alonso 718 670 concordance with the positive relationships that (1985, 1996, 1998) in the Iberian Peninsula and 719 671 showed the non-irrigated agriculture on the majority Balearic Islands during the last quarter of the twentieth 720 672 of large branchiopod species. Similarly, it is known century, no more studies exist updating the 721 123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

722 information for the whole study area. In this sense, the Branchiopoda, Anostraca, Chirocephalidae), a new species 774 723 present study represents a new step on the knowledge from southern Spain. Zoosystema 31: 807–827. 775 Amat, F., F. Hontoria, O. Ruiz, A. J. Green, M. I. Sa´nchez, J. 776 724 of large branchiopod distribution in this area, not only Figuerola & F. Hortas, 2005. The American brine shrimp as 777 725 because it gathers all available information on the an exotic invasive species in the western Mediterranean. 778 726 distribution of this taxonomic group (historical records Biological Invasions 7: 37–47. 779 727 and new observations), but also because it analyses the Amat, F., F. Hontoria, J. C. Navarro, N. Vieira & G. Mura, 2007. 780 Biodiversity loss in the genus Artemia in the Western 781 728 effect of several landscape characteristics on its Mediterranean Region. Limnetica 26: 387–404. 782 729 distribution, detailing the spatial scale at which its Angeler, D. G., O. Viedma, S. Sa´nchez-Carrillo & M. A´ lvarez- 783 730 influence is noticeable for large branchiopods pres- Cobelas, 2008. Conservation issues of temporary wetland 784 731 ence. All this information would be highly valuable Branchiopoda (Anostraca, Notostraca: crustacea) in a 785 semiarid agricultural landscape: what spatial scales are 786 732 for land stakeholders under future conservation relevant? Biological Conservation 141: 1224–1234. 787 733AQ3 scenarios. Asem, A., A. Eimanifar & S.-C. Sun, 2016. Genetic variation 788 and evolutionary origins of parthenogenetic Artemia 789 734 Acknowledgements We dedicate this paper to the memory of (Crustacea: Anostraca) with different ploidies. Zoologica 790

Author Proof 735 Prof. Graziella Mura that inspired many of us with her Scripta 45: 421–436. 791 736 passionate approach to these interesting organisms. We also Bagella, S., S. Gasco´n, M. C. Caria, J. Sala, M. A. Mariani & D. 792 737 dedicate this study to the memory of our co-author Prof. Maria Boix, 2010. Identifying key environmental factors related 793 738 Rosa Miracle, who was an admired and stimulating researcher in to plant and crustacean assemblages in Mediterranean 794 739 the Iberian limnology. We want to sincerely thank all people that temporary ponds. Biodiversity and Conservation 19: 795 740 selflessly supplied us with large branchiopod records. We also 1749–1768. PROOF 796 741 thank two anonymous reviewers for providing valuable Bagella, S., S. GascoPROOF´n, R. Filigheddu, A. Cogoni & D. Boix, 797 742 comments to an earlier version of the manuscript. MF is 2016. Mediterranean temporary ponds: new challenges 798 743 supported by a grant from the Brazilian Conselho Nacional de from a neglected habitat. Hydrobiologia 782: 1–10. 799 744 Desenvolvimento Cientı´fico e Tecnolo´gico—CNPq (401045/ Beja, P. & R. Alcazar, 2003. Conservation of Mediterranean 800 745 2014-5), program Cieˆncia sem Fronteiras. DC-M held a grant temporary ponds under agricultural intensification: an 801 746 from the Spanish Ministry of Education, Culture and Sport evaluation using amphibians. Biological Conservation 802 747 (FPU014/06783). 114: 317–326. 803 Belk, D., 1998. Global status and trends in ephemeral pool 804 invertebrate conservation: implications for Californian 805 fairy shrimp. In Witham, C. W., et al. (eds.), Ecology, 806 Conservation, and Management of Vernal Pool Ecosys- 807 748 References tems – Proceedings from a 1996 Conference. California 808 Native Plant Society, Sacramento: 147–150. 809 749 Abella´n, P. & D. Sa´nchez-Ferna´ndez, 2015. A gap analysis Ben Naceur, H., A. Ben Rejeb Jenhani & M. S. Romdhane, 810 750 comparing the effectiveness of Natura 2000 and national 2013. Morphometric characterization of adult Artemia 811 751 protected area networks in representing European (Crustacea: Branchiopoda) populations from costal and 812 752 amphibians and reptiles. Biodiversity and Conservation 24: inland Tunisian salt lakes. African Invertebrates 54: 813 753 1377–1390. 543–555. 814 754 Abella´n, P., D. Sa´nchez-Ferna´ndez, J. Velasco & A. Milla´n, Boix, D., 2002. Aportacio´ al coneixement de la distribucio´ 815 755 2005. Assessing conservation priorities for insects: status d’anostracis i notostracis (Crustacea: Branchiopoda) als 816 756 of water beetles in southeast Spain. Biological Conserva- Paı¨sos Catalans. Butlletı´ de la Institucio´ Catalana d’His- 817 757 tion 121: 79–90. to`ria Natural 70: 55–71. 818 758 AEMET & IM, 2011. Iberian climate atlas. Air temperature and Boix, D., S. Gasco´n, J. Sala, A. Badosa, S. Brucet, R. Lo´pez- 819 759 precipitation (1971-2000). Agencia Estatal de Meteo- Flores, M. Martinoy, J. Gifre & X. D. Quintana, 2008. 820 760 rologı´a & Instituto de Meteorologia de Portugal, Madrid. Patterns of composition and species richness of crustaceans 821 761 Alfonso, G., 2017. Diversity and distribution of large bran- and aquatic insects along environmental gradients in 822 762 chiopods (Branchiopoda: Anostraca, Notostraca, Spini- Mediterranean water bodies. Hydrobiologia 597: 53–69. 823 763 caudata) in Apulian ponds (SE Italy). The European Boix, D., J. Sala, D. Escoriza & M. Alonso, 2016. Linderiella 824 764 Zoological Journal 84: 172–185. jebalae sp. nov. (Crustacea: Branchiopoda: Anostraca), a 825 765 Alonso, M., 1985. A survey of the Spanish Euphyllopoda. new species from the Rif mountains (northern Morocco). 826 766 Miscel la`nia Zoolo`gicaUNCORRECTEDUNCORRECTED 9: 179–208. Zootaxa 4138: 491–512. 827 767 Alonso, M., 1996. Crustacea. Branchiopoda. In Ramos, M. A., Borcard, D., F. Gillet & P. Legendre, 2011. Numerical Ecology 828 768 et al. (eds), Fauna Ibe´rica, Vol. 7. Museo Nacional de with R. Springer, New York. 829 769 Ciencias Naturales, CSIC, Madrid. Boven, L., B. Vanschoenwinkel, E. R. De Roeck, A. Hulsmans 830 770 Alonso, M., 1998. Las lagunas de la Espan˜a peninsular. Lim- & L. Brendonck, 2008. Diversity and distribution of large 831 771 netica 15: 1–176. branchiopods in Kiskunsa´g (Hungary) in relation to local 832 772 Alonso, M. & J. Garcı´a-de-Lomas, 2009. Systematics and habitat and spatial factors: implications for their conser- 833 773 ecology of Linderiella baetica n. sp. (Crustacea, vation. Marine and Freshwater Research 59: 940–950. 834

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

835 Brendonck, L., D. C. Rogers, J. Olesen, S. Weeks & W. R. Hoeh, Garcı´a de Lomas, J., C. M. Garcı´a, F. Hortas, F. Prunier, D. 896 836 2008. Global diversity of large branchiopods (Crustacea: Boix, J. Sala, D. Leo´n, L. Serrano, J. Prenda, J. D. Gilbert, 897 837 Branchiopoda) in freshwater. Hydrobiologia 595: 167–176. F. J. Guerrero, F. Marrone, M. Sahuquillo, A. Camacho, C. 898 838 Cancela da Fonseca, L., M. Cristo, M. Machado, J. Sala, J. Reis, Olmo, M. R. Miracle, C. Zamora-Mun˜oz, G. Mura, M. 899 839 R. Alcazar & P. Beja, 2008. Mediterranean temporary Machado,´ I.Sa´nchez, J. A´ .Ga´lvez, M. Florencio, J. 900 840 ponds in Southern Portugal: key faunal groups as man- L. Pe´rez-Bote & M. Alonso, 2016. Linderiella baetica 901 841 agement tools? Pan-American Journal of Aquatic Sciences Alonso & Garcı´a-de-Lomas 2009 (Crustacea, Bran- 902 842 3: 304–320. chiopoda, Anostraca): >al borde de la extincio´n? Revista de 903 843 Caro, T., 2010. Conservation by Proxy. Indicator, Umbrella, la Sociedad Gaditana de Historia Natural 10: 15–26. 904 844 Keystone, Flagship, and Other Surrogate Species. Island Gasco´n, S., M. Machado, J. Sala, L. Cancela da Fonseca, M. 905 845 Press, Washington. Cristo & D. Boix, 2012. Spatial characteristics and species 906 846 Cottarelli, V. & G. Mura, 1983. Anostraci, Notostraci, Con- niche attributes modulate the response by aquatic passive 907 847 costraci (Crustacea: Anostraca, Notostraca, Conchostraca). dispersers to habitat degradation. Marine and Freshwater 908 848 In Ruffo, S. (ed.), Guide per il riconoscimento delle specie Research 63: 232–245. 909 849 animali delle acque interne italiane, 18. Consiglio Nazio- Ghomari, M. S., G. S. Selselet, F. Hontoria & F. Amat, 2011. 910 850 nale delle Ricerche, Verona. Artemia biodiversity in Algerian sebkhas. Crustaceana 84: 911 851 Crawley, M. J., 2002. Statistical Computing. An Introduction to 1025–1039. 912

Author Proof 852 Data Analysis Using S-Plus. John Wiley & Sons, Griffith, D. M., J. A. Veech & C. J. Marsh, 2016. Cooccur: 913 853 Chichester. probabilistic species co-occurrence analysis in R. Journal 914 854 Culioli, J.-L., C. Mori, A. Orsini & B. Marchand, 2006. Distri- of Statistical Software 69: 1–17. 915 855 bution and status of the large Branchiopoda (Crustacea) in Grillas, P., P. Gauthier, N. Yavercovski & C. Perennou (eds), 916 856 Corsica, France. First International Symposium on Envi- 2004. Mediterranean Temporary Pools. Volume 1 – Issues 917 857 ronment Identities and Mediterranean Area (ISEIMA). relating to conservation,PROOF functioning and management. 918 858 Corte-Ajaccio, Corsica, France, 9-12 July 2006: 271–273. Station BiologiquePROOF de la Tour du Valat, Le Sambuc. 919 859 De Roeck, E. R., B. J. Vanschoenwinkel, J. A. Day, Y. Xu, L. Hall, D. L., M. R. Willig, D. L. Moorhead, R. W. Sites, E. 920 860 Raitt & L. Brendonck, 2007. Conservation status of large B. Fish & T. R. Mollhagen, 2004. Aquatic macroinverte- 921 861 branchiopods in the western Cape, South Africa. Wetlands brate diversity of playa wetlands: the role of landscape and 922 862 27: 162–173. island biogeographic characteristics. Wetlands 24: 77–91. 923 863 Defaye, D., N. Rabet & A. Thie´ry, 1998. Atlas et bibliographie Hartel, T. & H. von Wehrden, 2013. Farmed areas predict the 924 864 des crustace´s branchiopodes (Anostraca, Notostraca, distribution of amphibian ponds in a traditional rural 925 865 Spinicaudata) de France me´tropolitaine. In Maurin, H. landscape. PLoS ONE 8: e63649. 926 866 (ed), Coll. Patrimoines Naturels, 32. Service du Patrimoine Horva´th, Z., C. F. Vad, L. Vo¨ro¨s & E. Boros, 2013. Distribution 927 867 Naturel/IEGB/MNHN, Paris. and conservation status of fairy shrimps (Crustacea: 928 868 Eder, E. & W. Ho¨dl, 2002. Large freshwater branchiopods in Anostraca) in the astatic soda pans of the Carpathian basin: 929 869 Austria: diversity, threats and conservational status. In the role of local and spatial factors. Journal of Limnology 930 870 Escobar-Briones, E. & F. A´ lvarez (eds.), Modern 72: 103–116. 931 871 Approaches to the Study of Crustacea. Kluwer Academic/ Hothorn, T., K. Hornik & A. Zeileis, 2006. Unbiased recursive 932 872 Plenum Publishers, New York: 281–289. partitioning: A conditional inference framework. Journal 933 873 Eder, E., W. Ho¨dl & R. Gottwald, 1997. Distribution and phe- of Computational and Graphical Statistics 15: 651–674. 934 874 nology of large branchiopods in Austria. Hydrobiologia Korn, M. & A. K. Hundsdoerfer, 2016. Molecular phylogeny, 935 875 359: 13–22. morphology and taxonomy of Moroccan Triops granarius 936 876 EEA, 2006. CORINE land cover 2006 project. European (Lucas, 1864) (Crustacea: Notostraca), with the description 937 877 Environment Agency [Accessed 1 Dec 2016]. of two new species. Zootaxa 4178: 328–346. 938 878 EEA, 2015. Natura 2000 data – the European network of pro- Korn, M., F. Marrone, J. L. Pe´rez-Bote, M. Machado, M. Cristo, 939 879 tected sites. European Environment Agency. http://www. L. Cancela da Fonseca & A. K. Hundsdoerfer, 2006. Sister 940 880 eea.europa.eu/data-and-maps/data/natura-5. [Accessed 1 species within the Triops cancriformis lineage (Crustacea, 941 881 Dec 2016]. Notostraca). Zoologica Scripta 35: 301–322. 942 882 ESRI, 1999. ArcMap 10.0 GIS software. Redlands, California Korn, M., A. J. Green, M. Machado, J. Garcı´a-de-Lomas, M. 943 883 USA. Cristo, L. Cancela da Fonseca, D. Frisch, J. L. Pe´rez-Bote 944 884 Euliss Jr., N. H. & D. M. Mushet, 1999. Influence of agriculture & A. K. Hundsdoerfer, 2010. Phylogeny, molecular ecol- 945 885 on aquatic invertebrate communities of temporary wet- ogy and taxonomy of southern Iberian lineages of Triops 946 886 lands in the Prairie Pothole Region of North Dakota, USA. mauritanicus (Crustacea: Notostraca). Organisms, Diver- 947 887 Wetlands 19: 578–583. sity & Evolution 10: 409–440. 948 888 European Statistical System,UNCORRECTEDUNCORRECTED 2011. Population and Housing Logan, M., 2010. Biostatistical Design and Analysis Using R. 949 889 Census database. Eurostat. http://ec.europa.eu/eurostat/ A Practical Guide. Wiley, Oxford. 950 890 web/population-and-housing-census/census-data/2011- Lumbreras, A., J. T. Marques, A. F. Belo, M. Cristo, M. Fer- 951 891 census [Accessed 1 Dec 2016]. nandes, D. Galioto, M. Machado, A. Mira, P. Sa´-Sousa, R. 952 892 Gallego-Ferna´ndez, J. B., M. R. Garcı´a-Mora & F. Garcı´a- Silva, L. G. Sousa & C. Pinto-Cruz, 2016. Assessing the 953 893 Novo, 1999. Small wetlands lost: a biological conservation conservation status of Mediterranean temporary ponds 954 894 hazard in Mediterranean landscapes. Environmental Con- using biodiversity: a new tool for practitioners. Hydrobi- 955 895 servation 26: 190–199. ologia 782: 187–199. 956

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

957 Mabidi, A., M. S. Bird, R. Perissinotto & D. C. Rogers, 2016. Peel, M. C., B. L. Finlayson & T. A. McMahon, 2007. Updated 1018 958 Ecology and distribution of large branchiopods (Crustacea, world map of the Ko¨ppen-Geiger climate classification. 1019 959 Branchiopoda, Anostraca, Notostraca, Laevicaudata, Hydrology and Earth System Sciences 11: 1633–1644. 1020 960 Spinicaudata) of the Eastern Cape Karoo, South Africa. Prunier, F. & S. Saldan˜a, 2010. Grandes branquio´podos (Crus- 1021 961 ZooKeys 618: 15–38. tacea: Branchiopoda: Anostraca, Spinicaudata, Notostraca) 1022 962 Machado, M. & J. Sala, 2013. Tanymastigites lusitanica sp. nov. en la provincia de Co´rdoba (Espan˜a) (an˜o hidrolo´gico 1023 963 (Crustacea: Branchiopoda: Anostraca) from Portugal, first 2009/2010). Boletı´n de la Sociedad Entomolo´gica Arago- 1024 964 representative of the genus in Europe. Zootaxa 3681: nesa 47: 349–355. 1025 965 501–523. R Core Team, 2013. R: A language and environment for sta- 1026 966 Machado, M., M. Cristo & L. Cancela da Fonseca, 1999. Non- tistical computing. R Foundation for Statistical Comput- 1027 967 cladoceran branchiopod crustaceans from southwest Por- ing, Vienna. http://www.R-project.org. 1028 968 tugal. I. Occurrence notes. Crustaceana 72: 591–602. Rabet, N., J.-F. Cart, D. Montero & H. Boulekbache, 2005. First 1029 969 Maechler, M., P. Rousseeuw, A. Struyf, M. Hubert & K. Hornik, record of Lynceus brachyurus Mu¨ller, 1776 (Bran- 1030 970 2016. Cluster: Cluster Analysis Basics and Extensions. chiopoda, Laevicaudata, Lynceidae) in France. Crus- 1031 971 Foundation for Statistical Computing, Vienna, Austria. taceana 78: 931–940. 1032 972 Marrone, F. & G. Mura, 2006. Updated status of Anostraca, Redo´n, S., F. Amat, M. I. Sanchez & A. J. Green, 2015. Com- 1033 973 Notostraca and Spinicaudata (Crustacea Branchiopoda) in paring cestode infections and their consequences for host 1034

Author Proof 974 Sicily (Italy): review and new records. Naturalista Siciliano fitness in two sexual branchiopods: alien Artemia francis- 1035 975 30: 3–19. cana and native A. salina from syntopic-populations. PeerJ 1036 976 Marrone, F., M. Korn, F. Stoch, L. Naselli-Flores & S. Turki, 3: e1073. 1037 977 2016. Updated checklist and distribution of large bran- Rhazi, L., P. Grillas, A. Mounirou Toure & L. Tan Ham, 2001. 1038 978 chiopods (Branchiopoda: Anostraca, Notostraca, Spini- Impact of land use in catchment and human activities on 1039 979 caudata) in Tunisia. Biogeographia 31: 27–53. water, sedimentPROOF and vegetation of Mediterranean tempo- 1040 980 Martens, K. & F. De Moor, 1995. The fate of Rhino Ridge pool rary pools. ComptesPROOF Rendus de l’Academie des Sciences 1041 981 at Thomas Baines Nature Reserve: a cautionary tale for Serie III, Sciences de la vie 324: 165–177. 1042 982 nature conservationists. South African Journal of Science Rhazi, L., P. Grillas, E.-R. Saber, M. Rhazi, L. Brendonck & A. 1043 983 91: 385–387. Waterkeyn, 2012. Vegetation of Mediterranean temporary 1044 984 Miracle, M. R., M. Sahuquillo & E. Vicente, 2008. Large pools: a fading jewel? Hydrobiologia 689: 23–36. 1045 985 branchiopods from freshwater temporary ponds of Eastern Robson, B. J. & C. J. Clay, 2005. Local and regional macroin- 1046 986 Spain. Verhandlungen der Internationalen Vereinigung fu¨r vertebrate diversity in the wetlands of a cleared agricultural 1047 987 Limnologie 30: 501–505. landscape in south-western Victoria, Australia. Aquatic 1048 988 Mun˜oz, J., A´ .Go´mez, A. J. Green, J. Figuerola, F. Amat & C. Conservation 15: 403–414. 1049 989 Rico, 2010. Evolutionary origin and phylogeography of the Rodrı´guez-Flores, P. C., A. Sa´nchez-Vialas & M. Garcı´a-Parı´s, 1050 990 diploid obligate parthenogen Artemia parthenogenetica 2016. Muestreos taxono´micos en charcos estacionales: una 1051 991 (Branchiopoda: Anostraca). PLoS ONE 5: e11932. herramienta imprescindible para el conocimiento de la 1052 992 Mura, G., 1999. Current status of the Anostraca of Italy. distribucio´n geogra´fica de Anostraca (Crustacea: Bran- 1053 993 Hydrobiologia 405: 57–65. chiopoda) en el centro de la Penı´nsula Ibe´rica. Hetero- 1054 994 Mura, G., I. Kappas, A. D. Baxevanis, S. Moscatello, Q. pterus Revista de Entomologı´a 16: 29–52. 1055 995 D’Amico, G. M. Lo´pez, F. Hontoria, F. Amat & T. Rogers, D. C., 2013. Anostraca catalogus (Crustacea: Bran- 1056 996 J. Abatzopoulos, 2006. Morphological and molecular data chiopoda). The Raffles Bulletin of Zoology 61: 525–546. 1057 997 reveal the presence of the invasive Artemia franciscana in Rogers, D. C., M. Schwentner, J. Olesen & S. Richter, 2015. 1058 998 Margherita di Savoia salterns (Italy). International Review Evolution, classification, and global diversity of large 1059 999 of Hydrobiology 91: 539–554. Branchiopoda. Journal of Crustacean Biology 35: 297–300. 1060 1000 Nhiwatiwa, T., L. Brendonck, A. Waterkeyn & B. Vanschoen- Rosati, L., M. Marignani & C. Blasi, 2008. A gap analysis 1061 1001 winkel, 2011. The importance of landscape and habitat comparing Natura 2000 vs National Protected Area net- 1062 1002 properties in explaining instantaneous and long-term dis- work with potential natural vegetation. Community Ecol- 1063 1003 tributions of large branchiopods in subtropical temporary ogy 9: 147–154. 1064 1004 pans. Freshwater Biology 56: 1992–2008. Sahuquillo, M. & M. R. Miracle, 2013. The role of historic and 1065 1005 Nhiwatiwa, T., A. Waterkeyn, B. J. Riddoch & L. Brendonck, climatic factors in the distribution of crustacean commu- 1066 1006 2014. A hotspot of large branchiopod diversity in south- nities in Iberian Mediterranean ponds. Freshwater Biology 1067 1007 eastern Zimbabwe. African Journal of Aquatic Science 39: 58: 1251–1266. 1068 1008 57–65. Sahuquillo, M. & M. R. Miracle, 2015. Crustacean diversity and 1069 1009 Nourisson, M. & A. Thie´ry, 1988. Introduction practique a` la conservation value indexes in pond assessment: implica- 1070 1010 syste´matique des organismesUNCORRECTEDUNCORRECTED des eaux continentales fran- tions for rare and relict species. Limnetica 34: 333–348. 1071 1011 c¸aises. -9. Crustace´s Branchiopodes (Anostrace´s, Noto- Sala, O. E., F. Stuart Chapin III, J. J. Armesto, E. Berlow, J. 1072 1012 strace´s, Conchostrace´s) (suite et fin). Bulletin mensuel de Bloomfield, R. Dirzo, E. Huber-Sanwald, L. F. Huenneke, 1073 1013 la Socie´te´ linne´enne de Lyon 57: 104–135. R. B. Jackson, A. Kinzig, R. Leemans, D. M. Lodge, H. 1074 1014 O’Neill, B. J., D. C. Rogers & J. H. Thorp, 2016. Flexibility of A. Mooney, M. Oesterheld, N. L. Poff, M. T. Sykes, B. 1075 1015 ephemeral wetland crustaceans: environmental constraints H. Walker, M. Walker & D. H. Wall, 2000. Global biodi- 1076 1016 and anthropogenic impacts. Wetlands Ecology and Man- versity scenarios for the year 2100. Science 287: 1077 1017 agement 24: 2279–2291. 1770–1774. 1078

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Hydrobiologia

1079 Samraoui, B. & H. J. Dumont, 2002. The large branchiopods Timms, B. V. & P. R. Sanders, 2002. Biogeography and ecology 1114 1080 (Anostraca, Notostraca and Spinicaudata) of Numidia of Anostraca (Crustacea) in middle Paroo catchment of the 1115 1081 (Algeria). Hydrobiologia 486: 119–123. Australian arid-zone. Hydrobiologia 486: 225–238. 1116 1082 Samraoui, B., K. Chakri & F. Samraoui, 2006. Large bran- Underwood, E. C., J. H. Viers, K. R. Klausmeyer, R. L. Cox & 1117 1083 chiopods (Branchiopoda: Anostraca, Notostraca and M. R. Shaw, 2009. Threats and biodiversity in the 1118 1084 Spinicaudata) from the salt lakes of Algeria. Journal of mediterranean biome. Diversity and Distributions 15: 1119 1085 Limnology 65: 83–88. 188–197. 1120 1086 Scanabissi, F., G. Alfonso, S. Bergamaschi & B. Mantovani, Van den Broeck, M., A. Waterkeyn, L. Brendonck & L. Rhazi, 1121 1087 2006. Primo ritrovamento di Lepidurus couesii Packard, 2015a. Distribution, coexistence, and decline of Moroccan 1122 1088 1875 in Italia. Thalassia Salentina 29: 111–122. large branchiopods. Journal of Crustacean Biology 35: 1123 1089 Siqueira, T., L. M. Bini, F. O. Roque, S. R. Marques Couceiro, S. 355–365. 1124 1090 Trivinho-Strixino & K. Cottenie, 2012. Common and rare Van den Broeck, M., A. Waterkeyn, L. Rhazi, P. Grillas & L. 1125 1091 species respond to similar niche processes in macroinver- Brendonck, 2015b. Assessing the ecological integrity of 1126 1092 tebrate metacommunities. Ecography 35: 183–192. endorheic wetlands, with focus on Mediterranean tempo- 1127 1093 Stoch, F., M. Korn, S. Turki, L. Naselli-Flores & F. Marrone, rary ponds. Ecological Indicators 54: 1–11. 1128 1094 2016. The role of spatial environmental factors as deter- Verdiell-Cubedo, D. & D. Boix, 2014. Primeros datos sobre la 1129 1095 minants of large branchiopod distribution in Tunisian distribucio´n de grandes branquio´podos (Crustacea: Bran- 1130

Author Proof 1096 temporary ponds. Hydrobiologia 782: 37–51. chiopoda) en la Regio´n de Murcia (SE Espan˜a). Anales de 1131 1097 Strobl, C., A.-L. Boulesteix, A. Zeileis & T. Hothorn, 2007. Bias Biologı´a 36: 65–69. 1132 1098 in random forest variable importance measures: Illustra- Walsh, C. & R. Mac Nally, 2013. Hier.part: Hierarchical Par- 1133 1099 tions, sources and a solution. BMC Bioinformatics 8: 25. titioning. https://cran.r-project.org/web/packages/hier. 1134 1100 Thie´ry, A., 1986. Les crustace´s branchiopodes (Anostraca, part/index.html. 1135 1101 Notostraca et Conchostraca) du Maroc occidental. I. In- Waterkeyn, A., P.PROOF Grillas, E. R. De Roeck, L. Boven & L. 1136 1102 ventaire et re´partition. Bulletin de la Socie´te´ d’Histoire Brendonck, 2009.PROOF Assemblage structure and dynamics of 1137 1103 Naturelle de Toulouse 122: 145–155. large branchiopods in Mediterranean temporary wetlands: 1138 1104 Thie´ry, A., 1987. Les Crustace´s Branchiopodes Anostraca, patterns and processes. Freshwater Biology 54: 1139 1105 Notostraca et Conchostraca des milieux limniques tempo- 1256–1270. 1140 1106 raires (dayas) au Maroc. Taxonomie, bioge´ographie, Weibull, A.-C., J. Bengtsson & E. Nohlgren, 2000. Diversity of 1141 1107 ecologie. Ph.D. Thesis. Universite´ d’Aix-Marseille III, butterflies in the agricultural landscape: the role of farming 1142 1108 Marseille. system and landscape heterogeneity. Ecography 23: 1143 1109 Thie´ry, A., 1991. Multispecies coexistence of branchiopods 743–750. 1144 1110 (Anostraca, Notostraca & Spinicaudata) in temporary Zacharias, I. & M. Zamparas, 2010. Mediterranean temporary 1145 1111 ponds of Chaouia plain (western Morocco): sympatry or ponds. A disappearing ecosystem. Biodiversity and Con- 1146 1112 syntopy between usually allopatric species. Hydrobiologia servation 19: 3827–3834. 1147 1113 212: 117–136.

UNCORRECTEDUNCORRECTED

123

Journal : Medium 10750 Dispatch : 5-7-2017 Pages : 18 Article No. : 3293 h LE h TYPESET MS Code : HYDR-D-16-01181 h44CP h DISK Journal : 10750 Article : 3293 123 the language of science Author Query Form

Please ensure you fill out your response to the queries raised below and return this form along with your corrections

Dear Author

Author Proof During the process of typesetting your article, the following queries have arisen. Please check your typeset proof carefully against the queries listed below and mark the necessary changes either directly on the proof/online grid or in the ‘Author’s response’ area provided below

Query Details Required Author’s Response AQ1 Kindly check the edits made in the Affiliations 2, 9, 15, 17 and amend if necessary. AQ2 Please confirm the section headings are correctly identified. AQ3 Kindly check and confirm the inserted details in the Reference Maechler (2016) and amend if necessary.