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1 1 Land use influences harvestman diversity in northern Spain: a case study involving 2 3 2 secondary grasslands and forest plantations (Arachnida: ) 4 5 6 3 Izaskun MERINO-SÁINZ and Araceli ANADÓN 7 8 9 10 4 Departamento de Biología de Organismos y Sistemas, Universidad de Oviedo. Catedrático 11 12 5 Valentín Andrés s/n 33006 Oviedo (Asturias, Spain) 13 14 15 6 Corresponding author: Araceli ANADÓN. E-mail: [email protected] 16 17 18 7 Telephone number 34 985 10 48 16 19 20 21 22 8 23 24 25 9 26 27 28 10 29 30 31 32 11 33 34 35 12 36 37 38 13 39 40 41 14 Acknowledgements 42 43 The Government of the Principado de Asturias provided the predoctoral grant 2008-2011 44 15 45 46 16 (research scholarship of I. Merino), reference: BP08039, obtained through the “Severo Ochoa 47 48 17 Program” of the FICYT Foundation. 49 50 51 18 We thank Carlos Prieto for help in taxonomic and identification enquiries, T.E. Díaz-González 52 53 19 and J.A. Fernández Prieto for help in habitat characterization, Mario O. Díaz Anadón for 54 55 56 20 obtaining the species indicator values and Irene Fernández-Rodríguez and Florentino Braña for 57 58 21 assistance in statistical analyses. 59 60 22 61 62 1 63 64 65 23 Abstract 1 2 24 Harvestmen have a general distribution pattern, with more species and higher abundance in 3 4 25 forests than in open habitats, previously verified in mountain Cantabrian areas of northern 5 6 26 Spain. The lower altitude areas in the same zone present a more complex mosaic landscape with 7 8 9 27 mixed natural and managed habitats, mainly secondary grasslands and forest plantations, a 10 11 28 combination of characteristics that makes a comparison of their harvestman distribution pattern 12 13 29 with that of the previously mentioned mountain areas very interesting. 14 15 30 These managed habitats, and also contiguous habitats like natural forests, non-planted young 16 17 18 31 forests, shrublands and habitat boundaries were studied. All these systems were continuously 19 20 32 sampled with 7 pitfall traps during one year at 28 sites. Their harvestman assemblages were 21 22 33 differentiated with 6 different analyses, and indicator species were identified. 23 24 34 The spatial patterns of harvestman diversity in low managed and natural areas differed from 25 26 those of mountain areas, despite their having 15 species in common. There was high average 27 35 28 29 36 harvestman species richness at each site. Shrublands were the richest habitats. The frequency 30 31 37 and abundance of harvestman species also varied between the 2 areas. 32 33 38 Grasslands had a unique harvestman composition with significant extraordinary abundances due 34 35 39 to Homalenotus quadridentatus -indicator species of this habitat- and H. laranderas. Leiobunum 36 37 38 40 rotundum was the indicator species of 2 clusters with trees. 39 40 41 H. laranderas, Paroligolophus agrestis and Ischyropsalis hispanica, which were indicator 41 42 42 species of some open habitats in the low Cantabrian area, were indicators of shady forests in 43 44 43 mountain Cantabrian territories. 45 46 47 44 None of the 16 species found was under threat. 48 49 45 50 51 52 46 Keywords: Opiliones, Iberian Peninsula, diversity patterns, agroecosystems, indicator species 53 54 55 47 56 57 48 58 59 60 61 62 2 63 64 65 49 Introduction 1 2 3 50 Harvestmen are a common component of terrestrial ecosystems and have higher species 4 5 51 diversity in tropical areas, with a decline toward the poles. In temperate areas the number of 6 7 52 harvestmen species at any given location is rarely greater than 12 (Curtis and Machado 2007). 8 9 10 53 Many harvestman species are collected with pitfall traps when wandering over the ground and 11 12 54 so they have frequently been studied together with other epigean fauna (Zingerle 1999; Ivask et 13 14 55 al 2008; Rosa García et al 2009a 2009b). 15 16 17 56 Some effort has been given to the study of harvestmen in managed habitat areas. Ivask et al. 18 19 57 (2008) found statistically significant differences between the number of Opiliones individuals 20 21 22 58 present in fields and on their edges. The influence of agricultural management type (Ivask et al. 23 24 59 2008; Marasas et al. 2001; Stašiov et al. 2011), the grazing history (Dennis et al. 2001; 25 26 60 Paschetta et al. 2013), the types of cultivated soils (Ivask et al. 2008) and types of forest 27 28 61 plantations (Hicks et al. 2003) on harvestman communities have all been studied. 29 30 31 62 Northwestern Spain sustains considerable harvestman diversity and a high number of endemic 32 33 34 63 species in the areas that have been studied (Rambla 1974, Prieto 2003, Merino Sáinz and 35 36 64 Anadón 2008, 2009). The studies on harvestman assemblages have focused mainly on areas 37 38 65 high in the mountains, far from populated nuclei and with few anthropogenic influences. 39 40 66 Though there are some taxonomic papers on the harvestmen in low Cantabrian areas (Merino- 41 42 43 67 Sáinz and Anadón 2013) the distribution of their harvestman assemblages had still not been 44 45 68 investigated until now. 46 47 48 69 Here, two types of managed habitats are studied: meadows and forest plantations. Secondary 49 50 70 grasslands were formerly natural forests and have now become grasslands due to human activity 51 52 71 and may be pastures (for grazing) or meadows (for hay-making). Secondary grasslands are an 53 54 55 72 essential part of Europe´s landscape and account for 35.3% of the utilized agricultural area 56 57 73 (Dengler el al. 2014). In Asturias, grasslands account for 25% of land use, meadows 58 59 60 61 62 3 63 64 65 74 representing 21% of the total (García Manteca et al. 2005), whilst forested areas make up 29% 1 2 75 of the land surface, 9% being forest plantations. 3 4 5 76 A comparative study of the distribution patterns of harvestmen in managed versus non-managed 6 7 77 habitats in different low Cantabrian habitats could reveal the effect of management on general 8 9 10 78 distribution patterns (Curtis and Machado 2007). Furthermore, the inclusion of managed 11 12 79 habitats might also provide some information of significance for conservation policies. Included 13 14 80 within the scope of this study are the species composition, species richness and abundance of 15 16 81 the harvestmen in the habitats of the region and the investigation of differences between the 17 18 82 harvestman assemblages. 19 20 21 22 83 The hypothesis concerning species richness was that forests and forest plantations would be 23 24 84 richer in species than the other open areas studied, whilst the hypothesis concerning abundance 25 26 85 was that forests and forest plantations would have greater abundance than open habitats. 27 28 29 86 The species composition of low Cantabrian areas (Merino-Sáinz and Anadón 2013) was nearly 30 31 87 the same as the species composition of Muniellos (Merino Sáinz and Anadón 2008- 2009), so 32 33 34 88 there is an opportunity to search for similarities and differences between the distribution of 35 36 89 these harvestman species in low and mountain areas in the Cantabrian region. Muniellos is a 37 38 90 “natural” (in the sense of Peterken 1993), forested Biosphere Reserve in the Cantabrian 39 40 91 mountains in Asturias. The low Cantabrian areas are in the biogeographic Cantabro-Atlantic 41 42 43 92 Province, while Muniellos is in the Orocantabrian Province. These 2 provinces are to be found 44 45 93 in Spain, on the northern fringe of the Iberian Peninsula, adjacent to the Mediterranean basin, 46 47 94 which is a biodiversity hotspot (Mittermeier et al. 2011; Myers et al. 2000). 48 49 95 Material and methods 50 51 Study area 52 96 53 54 97 The study was carried out in low Cantabrian areas in Asturias and Cantabria (Fig. 1). Over a 55 56 98 period of one year from March 2009 until April 2010, 26 plots (Table 1) were sampled in 57 58 99 Asturias and 2 plots in Cantabria. These plot areas have a temperate hyperoceanic/oceanic 59 60 61 62 4 63 64 65 100 submediterranean bioclimate and are included in the Cantabro-Atlantic Province of the 1 2 101 Eurosiberian phytogeographic Region (Rivas Martínez et al. 2004), next to the Orocantabrian 3 4 102 Province. 5 6 103 The managed habitats were 6 grasslands and 5 forest plantations. All the grassland plots were 7 8 9 104 meadows, located in 3 different municipalities: Oviedo, Muros de Nalón (both in Asturias) and 10 11 105 Piélagos (in Cantabria, locality of Vioño). Two of these meadows had fruit trees. 12 13 106 The natural habitats and forest plantations were sampled in Oviedo: 6 forests, 5 young forests, 4 14 15 107 shrublands and 2 boundaries or margins adjacent to 2 grasslands. One of the boundaries was 16 17 18 108 populated with horsetails and the other one with nettles. One plot was in the city and all the 19 20 109 other plots were in Monte Naranco, in 5 different areas or zones (Fig. 1): Ajuyán and Brañes 21 22 110 (the northern mountainside beside the river Nora), El Violeo (the western top of the mountain), 23 24 111 Ules and Naranco (the southern side). Mount Naranco has calcareous, siliceous and mixed soils. 25 26 Sampling scheme 27 112 28 29 30 113 Each plot was sampled with 7 pitfall traps which were processed as a single sample. Each pitfall 31 32 114 trap consisted of 2 plastic cups 11 cm in height with a diameter of 8 cm at the top and 5 cm at 33 34 115 the bottom. The outer cup remained in the ground and the inner cup was used to take the sample 35 36 116 and to renew the liquid each time. The pitfall traps had a solution of water and ethylene glycol 37 38 39 117 at 40% as preservative and antifreeze and 15 g/L of CALGON® sodium polymetaphosphate, as 40 41 118 emulsifier. A 10 cm long, 6 cm high roof was placed over the traps while functioning, to protect 42 43 119 them from the rain. 44 45 120 The samples were collected every 15 days. Harvestmen in the samples were sorted and 46 47 identified with the bibliography mentioned in the following preliminary studies: Merino-Sáinz 48 121 49 50 122 and Anadón (2013) and Merino-Sáinz et al. (2013). All the specimens are accessible in the 51 52 123 Harvestmen dataset of the BOS-Opi Collection of the University of Oviedo, Spain, 53 54 124 through the GBIF network. 55 56 125 Data analyses 57 58 59 60 61 62 5 63 64 65 126 Species richness, abundance, frequency and “true” diversity of harvestmen were obtained for 1 2 127 the different sites and habitats. Diversity was studied as: 2D= 1/λ (Hill 1973; Jost 2007; 3 4 128 Tuomisto 2010), called by Tuomisto “true” diversity. This diversity measure is the inverse of 5 6 129 the Simpson index of evenness; pi being the proportional abundance of the ith species. This 7 8 9 130 index increases as diversity intuitively increases. ANOVA (Analysis of Variance) was used to 10 11 131 discard the null hypothesis of no differences between the means of different habitats or clusters. 12 13 132 When the existence of differences had been proved, post hoc or “a posteriori”, multiple 14 15 133 comparisons, including the HSD test of Tukey, Scheffé and Fischer LSD (Least Significant 16 17 18 134 Difference), were used to determine between which habitats or clusters differences were found. 19 20 135 The relationship between the harvestman species richness and the harvestman abundance of the 21 22 136 sites was studied with the linear correlation coefficient r. 23 24 25 137 The smooth accumulation curves were produced to assess the quality of the inventory. The 26 27 138 sampling dates were taken as measures of sampling effort, and they were randomized 999 times. 28 29 30 139 The Simplex and Quasi-Newton method (Hortal et al. 2004) with the program Statistica V6 31 32 140 (StatSoft 2001) fitted the Clench function to the smoothed curves to estimate the asymptotes. 33 34 141 These asymptotes predicted the estimated species richness (Hortal et al. 2004) and the ratio 35 36 142 observed/estimated species richness (q) gives the proportion of the known inventories. When 37 38 39 143 the value of the final slope was lower than 0.1 and the percentage of collected species was over 40 41 144 70, the inventory was considered reliable enough and well sampled (Hortal and Lobo 2005). 42 43 145 The accumulation curves and Clench function (Table 2) confirmed that the inventories can be 44 45 146 considered reliable enough and well sampled, except for three sites that were insufficiently 46 47 sampled. The sampling efficiency percentage of these three sites was above 70% but the final 48 147 49 50 148 slope of the curves was greater than 0.1: 0.12, 0.13 and 0.16. 51 52 149 The relative position of sites and species was visualized in a correspondence analysis, run using 53 54 150 the PAST.exe statistical program (Hammer et al. 2001). 55 56 151 The hierarchical clustering (CLUSTER) was carried out with average group linkage and it used 57 58 59 152 the triangular matrices of the distances between sampling sites (according to their species 60 61 62 6 63 64 65 153 assemblages). The distance between two sites was measured with the Bray-Curtis coefficient of 1 2 154 similarity based on square root transformed abundance data of harvestmen. These matrices were 3 4 155 also used in the non-metric multidimensional scaling (MDS), which represents the distances 5 6 156 between the sites in a geometric space. 7 8 9 157 The dissimilarity between samples from different groups was obtained with the similarity 10 11 158 percentage analysis (SIMPER). The PRIMER V6 program (Clarke and Gorley 2006) was used 12 13 159 to obtain species accumulation curves, hierarchical clustering, multidimensional scaling, 14 15 160 analysis of similarity and similarity percentages. 16 17 18 161 Indicator species analyses for a cluster of sites were obtained using the package “indicspecies” 19 20 162 1.7.3 2014-07-10 (De Cáceres and Jansen 2014) in R (R Development Core Team 2012). The 21 22 163 indicator value indexes (IndVal) (De Cáceres and Legendre 2009; Dufrêne and Legendre 1997), 23 24 164 measured the association of a species for a given clustering of sites. 25 26 27 165 Results 28 29 166 The total number of epigean harvestmen studied were 12,208 specimens, of the following 16 30 31 167 species. Their distribution was Ho, holarctic, Eu, European or IE, Iberian endemic. 32 33 34 168 Suborder Hansen & Sørensen 1904 35 36 169 Superfamily Latreille 1802 37 38 39 170 Family Phalangiidae Latreille, 1802 40 41 171 Subfamily Oligolophinae Banks, 1893 42 43 172 1.- Paroligolophus agrestis (Meade, 1855). Ho 44 45 173 2.- Odiellus simplicipes (Simon, 1879). IE 46 47 3.- Odiellus seoanei (Simon, 1878). IE 48 174 49 50 175 Subfamily Phalangiinae Latreille, 1802 51 52 176 4.- Phalangium opilio Linnaeus, 1761. Ho 53 54 177 Family Simon, 1879 55 56 178 Subfamily Gyinae Šilhavý, 1946 57 58 59 179 5.- Gyas titanus Simon, 1879. Eu 60 61 62 7 63 64 65 180 Subfamily Leiobuninae Banks, 1893 1 2 181 6.- Leiobunum rotundum (Latreille, 1798). Eu 3 4 182 7.- Leiobunum blackwalli Meade, 1861. Eu 5 6 183 Subfamily Sclerosomatinae Simon, 1879 7 8 9 184 8.- Homalenotus quadridentatus (Cuvier, 1795). Eu 10 11 185 9.- Homalenotus laranderas Grasshoff, 1959. IE 12 13 186 Suborder Dyspnoi Hansen & Sørensen 1904 14 15 187 Superfamily Ischyropsalidoidea Simon 1879 16 17 18 188 Family Ischyropsalididae Simon 1879 19 20 189 10.- Ischyropsalis hispanica Roewer, 1953. IE 21 22 190 Family Sabaconidae Dresco, 1970 23 24 191 11.- Sabacon franzi Roewer, 1953. IE 25 26 Superfamily Troguloidea Sundevall 1833 27 192 28 29 193 Family Nemastomatidae Simon, 1872 30 31 194 Subfamily Nemastomatinae Simon, 1872 32 33 195 12.- Nemastomella dentipatellae (Dresco, 1967). IE 34 35 196 13.- Nemastoma hankiewiczii (Kulczynski, 1909). IE 36 37 38 197 Family: Trogulidae Sundevall, 1833 39 40 198 14- Anelasmocephalus cambridgei (Westwood, 1874). Eu 41 42 199 15.- Trogulus nepaeformis s.l. Eu? 43 44 200 Suborder Laniatores Thorell, 1876 45 46 47 201 Superfamily Travunioidea Absolon & Kratochvil 1932 48 49 202 Family Travuniidae Absolon & Kratochvil, 1932 50 51 203 16.- Hadziana clavigera (Simon, 1879). IE 52 53 54 55 204 This check-list is very similar to the list of the Opiliones fauna of Muniellos, since there are 15 56 57 205 shared species. H. quadridentatus is the sole species absent from Muniellos, which had 4 58 59 206 species not present in this catalogue. The Analysis of Variance of the distribution of the relative 60 61 62 8 63 64 65 207 abundance and the relative frequency of occupancy of sites of these 16 harvestman species in 1 2 208 low Cantabrian territories and Muniellos (Table 3), gave significant differences (p-value 0.015) 3 4 209 only for the frequencies. The relative frequency of many species was high in low Cantabrian 5 6 210 areas (Table 4). Half of the 16 harvestman species studied were found in many habitats and 7 8 9 211 were eurychorous, with a wider ecological niche: L. blackwalli, T. nepaeformis s.l., N. 10 11 212 dentipatellae, O. simplicipes and N. hankiewiczii were very abundant, while A. cambridgei, P. 12 13 213 opilio and L. rotundum were not abundant. The first 2 species were collected in all the sites. The 14 15 214 species with a smaller range, stenochorous, were G. titanus and H. clavigera, rare; O. seoanei, 16 17 18 215 P. agrestis, I. hispanica and S. franzi, not abundant species and H. quadridentatus and H. 19 20 216 laranderas, very abundant. 21 22 23 217 European species represent more than half the total harvestman abundance in low Cantabrian 24 25 areas, followed by endemic Iberian species (see last lines of Table 3). 26 218 27 28 219 Harvestman species richness and abundance will be treated separately here, since the linear 29 30 220 correlation coefficient was near to zero (r = 0.007), indicating that their values were 31 32 221 independent. 33 34 222 Species richness.- Each sampled site had between 6 and 12 harvestman species (Table 4) and 35 36 37 223 each species was found in a minimum of 2 to a maximum of 28 sites (average 16.4 sites). The 38 39 224 average harvestman species richness/site in these low Cantabrian areas, 9.6 ± 1.7 species/site, 40 41 225 was higher than in Muniellos, which had 6.2 ± 3.5 species/site. The specific inventories for the 42 43 226 different zones include 14 species in Violeo, 13 in Brañes, 13 in Ules, 12 in Ajuyán, 11 in 44 45 46 227 Muros and 9 species in Vioño. 47 48 228 The average harvestman species richness in the different habitats was always above 8 species 49 50 229 (Table 5). The managed habitats had the lowest average values: 8.4 for forest plantations and 51 52 230 8.8 for grasslands. The highest number of species was found in boundaries and shrublands. 53 54 55 231 In the low Cantabrian area, the average species richness was 1.09 times greater in habitats 56 57 58 232 which were not forests: forests had 9.2 ± 1.9 species/site. In Muniellos, however, the non-forest 59 60 233 habitats had only 0.48 times the number of species/site that the forest environments had; 61 62 9 63 64 65 234 Muniellos forests had 8.3 ± 3.1 harvestman species. The forest plantations with chestnuts and 1 2 235 pedunculate oaks had 6 to 9 harvestman species, while the eucalyptus plantation had 11 species. 3 4 5 236 There were no great differences in species richness between either sites or clusters. Analysis of 6 7 237 Variance showed differences in richness which were close to being significant (p = 0.065, 8 9 10 238 between habitats and p = 0.067 between clusters). 11 12 13 239 Abundance.- Each sampled site had between 41 and 1817 specimens (Table 4). The average 14 15 240 abundance value was close to the standard deviation: 436 ± 470.3 specimens/site. The mean 16 17 241 harvestman abundance was 2.6 times greater in the 5 habitats which were not forests (Table 5). 18 19 242 Shrublands and young forests followed grasslands in abundance. The Analysis of Variance 20 21 22 243 showed there are significant differences in harvestman abundance comparing habitats (p-value 23 24 244 0.000) and comparing clusters (p-value 0.000). Multiple comparisons “a posteriori” with HSD 25 26 245 Tukey and Scheffé of the abundance between all the habitats and all the clusters showed the 27 28 246 grasslands, which constitute one of the clusters, had significant differences, p-value of 0.000, 29 30 with respect to all the remaining habitats and clusters. The differences in abundance between 31 247 32 33 248 any of the other habitats or clusters were not significant. 34 35 249 Grasslands housed well-defined harvestman assemblages which had 3.73 times greater average 36 37 250 abundance than shrublands. There was a gradual decrease in the average number of specimens 38 39 251 from shrublands to young forests, boundaries, forests and forest plantations. Forest populations 40 41 42 252 were slightly more abundant and diverse than those of forest plantations (Table 4). 43 44 253 Diversity.- Harvestman diversity 1/λ values/site ranged between 1.38 and 7.67 (Table 4). The 45 46 254 highest mean values (Table 5) were obtained in cluster C2.1, which includes shrublands plus 47 48 255 adjacent young forest at the top of and on the southern side of Monte Naranco. The lowest mean 49 50 51 256 value was for grasslands. The Analysis of Variance showed there are significant differences in 52 53 257 the diversity values between clusters (p-value 0.018) and between habitats (p-value 0.037). 54 55 56 258 The analysis “a posteriori” found significant differences between the diversity of the cluster of 57 58 259 grasslands (A) and cluster C2.1. Regarding the differences in diversity between the habitats, 59 60 61 62 10 63 64 65 260 only the test for multiple comparisons “a posteriori” Fischer LSD (Least Significant Difference) 1 2 261 gave some difference: between grasslands and boundaries (p-value 0.015), grasslands and 3 4 262 shrublands (p-value 0.011) and grasslands and young forests (p-value 0.010). 5 6 7 263 Differences between harvestman assemblages.- The harvestman assemblages of grasslands were 8 9 10 264 clearly differentiated from the rest of habitats by all the analyses carried out in order to 11 12 265 investigate the differences: correspondence analysis, cluster, MDS, ANOSIM and SIMPER. 13 14 15 266 In the correspondence analysis (CA) (Fig. 2) all the grasslands, plus 3 harvestman species, 16 17 267 Homalenotus quadridentatus, H. laranderas and Odiellus seoanei, were isolated to the left, on 18 19 268 the first axis. The two Homalenotus spp. were the most abundant of the sampled species; O. 20 21 22 269 seoanei, however, was not abundant and it was present in only 2 sampling sites, one of which 23 24 270 was grassland; nonetheless, this species was present in 47% of sites in Muniellos. 25 26 271 The Pyrenean oak young forest of Naranco, a very small number of trees on the Naranco 27 28 272 mountain, was also separated to the upper right of the CA with Hadziana clavigera and 29 30 Paroligolophus agrestis. Gyas titanus was also isolated in the lower right. The rest of the places 31 273 32 33 274 had a more central position in the CA, together with 10 frequent species (see Table 4). L. 34 35 275 blackwalli, present in 100% of the sites and Nemastoma hankiewiczii, present in 72% of the 36 37 276 sites, had an intermediate position in CA, between grasslands and the central sites: they were 38 39 277 quite abundant in grasslands, more abundant than in other clusters (Table 4). 40 41 42 278 The cluster analyses of sites with harvestman abundance data discriminate between the 43 44 279 assemblages better, giving 5 sets of sites (Fig. 3, Table 4). Only one cluster included sites with 45 46 280 just one habitat type: cluster A included all the grasslands but no other habitat. The other 47 48 281 clusters had a mixture of habitats, and 4 habitat types were scattered in different clusters. 49 50 51 282 Cluster C1 included most of the forests and 2 young forests (one bay young forest and one 52 53 283 hazelnut young forest, shaded, humid with calcareous soil and a northern orientation). They 54 55 284 were all in Ajuyán and Brañes, on the northern side of the mountain, except for one 56 57 285 oligotrophous forest in Violeo, with northwestern orientation. 58 59 60 61 62 11 63 64 65 286 Cluster C2.1 included all the shrublands and 2 young forests of Pyrenean oaks and one young 1 2 287 forest of willow trees, all in Naranco and Violeo, at over 400 m altitude and in sunny 3 4 288 orientations. 5 6 289 Cluster C2.2 included 2 boundaries with 3 forest plantations, one of eucalyptus and 2 7 8 9 290 plantations of pedunculate oaks and chestnuts, in a lower, southern position on the mountain, all 10 11 291 in Ules, except one of the boundaries, which was in Oviedo. The area was a characteristic site 12 13 292 for Pyrenean oak trees. 14 15 293 Cluster B had the places that were poorest in abundance, quite near each other, one forest 16 17 18 294 plantation and one eutrophous gallery forest at the head of a stream, high up in Violeo. 19 20 295 In the MDS (Fig. 4) all the grasslands were grouped to the left, separated from the other 21 22 296 habitats. The sites with the same habitat type were near to each other, except for forest 23 24 297 plantations and young forests. Harvestman assemblages of forest plantations were scattered in 25 26 the MDS (Fig. 4). 27 298 28 29 299 The ANOSIM tests showed that the harvestman assemblages of grasslands and forests were the 30 31 300 most clearly differentiated from those of the other habitats (Table 6). Boundaries had the least 32 33 301 distinct harvestman assemblages since they only differed from grasslands. 34 35 36 302 The highest dissimilitude percentages between harvestman assemblages were found between the 37 38 39 303 grasslands and the other habitats (Table 6) with the SIMPER analysis of similitude. 40 41 42 304 Forest harvestman assemblages were different from the harvestman assemblages of all the other 43 44 305 types of habitats, excluding boundaries populated with horsetails or nettles. There was a gradual 45 46 306 increase in differences between forests and boundaries, young forests, forest plantations, 47 48 307 shrublands and finally grasslands (Table 6). 49 50 51 Indicator species.- Seven indicator species for certain habitats, for certain clusters or a 52 308 53 54 309 combination of habitats or clusters have been identified. H. quadridentatus was an indicator 55 56 310 species of cluster A, grasslands. This species was also indicator species of grasslands + 57 58 311 herbaceous boundaries. H. laranderas was in turn an indicator species of the sum of 2 clusters 59 60 61 62 12 63 64 65 312 and indicator of the combination of grasslands + shrublands + herbaceous boundaries (Table 7). 1 2 313 H. laranderas was present in surprisingly high numbers in grasslands. 3 4 5 314 Leiobunum rotundum was an indicator species of the sum of clusters C1 + C2.2, 2 clusters 6 7 315 which include forests, most forest plantations and boundaries (Table 6). 8 9 10 316 Ischyropsalis hispanica was an indicator species of the sum of open habitats: boundaries plus 11 12 317 shrublands and young forests, excluding grasslands. Sabacon franzi was indicator species only 13 14 318 of shrublands plus some young forests. Paroligolophus agrestis was an indicator species of the 15 16 319 cluster C2.1, which includes all the shrublands and 3 young forests. Odiellus simplicipes was an 17 18 320 indicator species of the sum of clusters with all the shrublands, boundaries, grasslands and 3 19 20 21 321 young forests (Table 4). 22 23 24 322 Discussion 25 26 323 The independence of the species richness and abundance of harvestmen seen in this study has 27 28 324 already been described in a National Park in the Czech Republic (Klimeš 1999) and the Pre- 29 30 Pyrenees with the linear correlation coefficient (r= 0.039) near to zero calculated from the data 31 325 32 33 326 of Rambla (1985). The abundance/site and diversity 1/λ in the habitats studied in the low 34 35 327 Cantabrian area differed between sites, as in the Pre-Pyrenees, where 12 different habitats 36 37 328 studied with a similar sampling device to ours showed a high standard deviation in average 38 39 329 harvestman abundance, 441 ± 352.6 (obtained from Rambla 1985 data). 40 41 42 330 Managed habitats had their own harvestman peculiarities, and both managed and natural 43 44 331 habitats in the low Cantabrian area of the biogeographic Cantabro-Atlantic Province had 45 46 332 different harvestman distribution patterns to the habitats of the mountain Cantabrian area of the 47 48 333 Orocantabrian Province. Among managed habitats, grasslands were clearly differentiated from 49 50 51 334 the remaining habitats, while forest plantations did not have characteristic harvestman 52 53 335 assemblages. 54 55 336 Managed habitats 56 57 58 59 60 61 62 13 63 64 65 337 The species richness and abundance of harvestman species in managed habitats were quite 1 2 338 unexpected. They had greater average harvestman species richness than natural habitats in the 3 4 339 Orocantabrian Province. 5 6 7 8 340 Grasslands were expected to be the poorest in both harvestman species and abundance and 9 10 341 forest plantations were expected to have quite high harvestman species richness and abundance. 11 12 342 However, the managed habitats were the two poorest in harvestmen species, and grasslands, 13 14 343 among all the habitats studied, were the most abundant in harvestmen. 15 16 17 344 Forest plantations had the lowest average harvestman species richness of all the habitats (8.4 18 19 345 species, 70% of that of the boundaries, with the highest harvestman richness), and they also had 20 21 346 the lowest average harvestman abundance of all the habitats (10% of that of the grasslands). 22 23 24 347 Since forest plantations are wooded or forested habitats, they were expected to have, in some 25 26 348 degree, greater harvestman species richness and abundance than open habitats in low Cantabrian 27 28 349 areas, considering the ratio found in mountain Cantabrian areas (Merino-Sáinz and Anadón 29 30 350 2015) and the generally observed patterns. Muniellos in the Cantabrian mountains had a ratio of 31 32 33 351 harvestman species richness of forests/open habitats of 2.06 and a ratio of harvestman 34 35 352 abundance forests/open habitats of 2.79. Curtis and Machado (2007) provided a general pattern 36 37 353 of harvestman distribution: they compiled the local richness of harvestman species in 89 38 39 354 forested and 70 open habitats from the data of many authors and found that the average 40 41 harvestman species richness in forested habitats was 2.8 times higher than in open habitats. 42 355 43 44 45 356 Forest plantation harvestman assemblages were different from the harvestman assemblages of 46 47 357 grasslands, forests and shrublands but presented no differences with harvestman assemblages of 48 49 358 young forests or boundaries. These forest plantations did not have a characteristic harvestman 50 51 52 359 assemblage but their populations seemed to be dependent on the harvestmen of neighbouring 53 54 360 habitats. 55 56 361 Grasslands in low Cantabrian areas had the next poorest average harvestman species richness, 57 58 362 8.8 species. However, this value was higher than the average harvestman species richness of the 59 60 61 62 14 63 64 65 363 forests in Muniellos, the richest habitat in the mountains, and was also higher than the 1 2 364 harvestman species richness of grasslands in mountain sites (Fig.1) of similar latitude such as 3 4 365 Muniellos (Merino-Sáinz and Anadón 2015), Illano (Rosa García et al. 2010a), open areas of 5 6 366 the Pre-Pyrenees (Rambla 1985) and Eastern Pyrenees (Ledoux and Emerit, 2006). In 7 8 9 367 mountains, the low number of harvestmen in the grasslands could be explained because 10 11 368 grasslands are more exposed than forests to changes in climatic factors (Curtis and Machado 12 13 369 2007). 14 15 370 The poorer harvestman species richness of grasslands was expected but, on the contrary, the 16 17 18 371 surprising abundance of harvestmen that was observed was most unexpected. The harvestman 19 20 372 abundance of grasslands significantly exceeded the abundance recorded in any other of the 21 22 373 habitats studied and this high abundance was due to the 2 Homalenotus species. There is some 23 24 374 previous knowledge of harvestman abundance in grazing grasslands in Alpine pastures and 25 26 from grazing experiments in the Cantabrian mountains, where the dominance of certain species 27 375 28 29 376 has been shown, but together with poorer overall harvestmen abundance. In the Northwest of 30 31 377 Italy Mitopus morio dominated all pastoral types in an Alpine environment, together with 32 33 378 Dasylobus ligusticus. In the Cantabrian mountains H. laranderas dominated the grazing 34 35 379 heathlands of Illano, whether the predominant vegetation was heather and heaths, gorse or grass, 36 37 38 380 though it was less abundant in grass (Rosa García et al 2010b). In Illano, shrublands had 97% of 39 40 381 the opilionid abundance and 10 species, while grasslands had only 3% of abundance and 7 41 42 382 species in experimental plots grazed by sheep or cattle (Rosa García et al 2010a). In Muniellos, 43 44 383 grasslands had only 4 species and very few specimens (Merino-Sáinz and Anadón 2015). 45 46 47 384 Grasslands in low Cantabrian areas had rich, unique and exclusive harvestman assemblages 48 49 385 very different to those of all the other habitats and they had the lowest diversity 1/λ. Their high 50 51 386 species richness could be related to the biodiversity of secondary grasslands which house many 52 53 387 plant species in Europe (Dengler et al. 2014) and to the structure of these grasslands (Morris 54 55 56 388 2000). Asturias and Cantabria are well known for their rich meadows within the association 57 58 389 Lino biennis - Cynosuretum cristatus Tüxen & Oberdorfer 1958 (Díaz González and Fernández 59 60 390 Prieto 1994). These are permanent grasslands which are not ploughed, two conditions which 61 62 15 63 64 65 391 favour higher species richness in harvestman communities (Ivask et al. 2008; Stašiov et al. 1 2 392 2011). 3 4 393 Natural areas.- 5 6 394 The natural habitats studied in the low Cantabrian area also had unexpected harvestman 7 8 9 395 assemblages with higher species richness and abundance than the habitats in the Orocantabrian 10 11 396 Province. Semi-natural areas tend to have much greater arthropod abundance than adjacent 12 13 397 arable fields (Pfiffner and Luka 2000). 14 15 398 Forests in low Cantabrian areas were not the habitats richest in harvestman species, as had been 16 17 18 399 expected, and neither were they the most abundantly populated habitats. However, they had 19 20 400 higher average harvestman species richness than mountain forests in Muniellos and the Pre- 21 22 401 Pyrenees. Mountain forests were the most abundantly populated habitats, though not all the 23 24 402 forests had the same abundance: those forests with a sunny orientation in Muniellos (Merino- 25 26 Sáinz and Anadón 2015) or with Mediterranean characteristics in the Pre-Pyrenees (Rambla 27 403 28 29 404 1985) were less abundant in harvestmen than the remainder of the forests. The riverside forests 30 31 405 of Muniellos had the most abundant harvestman assemblages of all the habitats studied there, 32 33 406 the opposite case to low Cantabrian gallery forests, which had low harvestman abundance when 34 35 407 compared to the other habitats. 36 37 38 408 The harvestman assemblages of low Cantabrian forests were different to harvestman 39 40 409 assemblages of all the other habitats except boundaries. The diversity of forests was similar to 41 42 410 the harvestman diversity in forest plantations and it was intermediate among all the habitats. 43 44 411 Shrublands, young forests and boundaries had the highest harvestman richness and an 45 46 47 412 intermediate abundance between forests and forest plantations on one end and grasslands on the 48 49 413 other end, probably related to a more complex structure and greater capacity for shelter than 50 51 414 grasslands. As a consequence, these 3 intermediate open habitats had the highest harvestman 52 53 415 diversity values (1/λ). Really, they constituted a transition between the most differentiated 54 55 56 416 grasslands and forests, as seen with ANOSIM and SIMPER analyses. In Illano a higher 57 58 417 abundance and richness of harvestmen in shrubland experimental plots compared to grassland 59 60 418 plots was found (Rosa García et al. 2010a). The discovery of the highest species richness in 61 62 16 63 64 65 419 herbaceous boundaries resembles the greater number of harvestmen found in Estonia along field 1 2 420 edges than in the centre of the fields with three different types of soils, (Ivask et al. 2008) and 3 4 421 on the field margins of Northern Europe (Marshall and Moonen 2002). 5 6 422 General considerations. 7 8 9 423 The number of 16 harvestman species found in low Cantabrian areas is quite high for temperate 10 11 424 areas (see Curtis and Machado 2007). The harvestman species richness found in mountain areas 12 13 425 of approximately the same latitude (Fig. 1) were: 11 in Pre-Pyrenees (Rambla 1985), 12 in 14 15 426 Pyrenees (Rambla and Perera 1989), 16 in Montseny mountain in Catalonia (Rambla and Perera 16 17 18 427 1995) and 14 in France (Ledoux and Emerit 2006). In western Asturias Illano had 14 species 19 20 428 and Muniellos had 19 species. 21 22 429 The most abundant harvestman species, the indicator species of some habitats, and the 23 24 430 frequencies of the species were different in the 2 Cantabrian territories. H. quadridentatus was 25 26 the most abundant species in the low Cantabrian areas (Table 3), while P. agrestis was the most 27 431 28 29 432 abundant species in Muniellos. P. agrestis, H. laranderas and I. hispanica, which in low 30 31 433 Cantabrian areas were indicators of open habitats, including young forests, were, in Muniellos, 32 33 434 indicators of the lower forest sites in shady habitats (Merino-Sáinz and Anadón 2015). 34 35 36 37 435 The mosaic landscape in low Cantabrian territories may have facilitated the presence of many 38 39 436 species in different habitats, which, in turn, could be related to the high relative frequency of 40 41 437 many species. The species may find shelter in adjoining areas, and may move easily between 42 43 438 adjacent, small-sized patches of different vegetation structure. In cultivated areas of 44 45 46 439 Switzerland, a mosaic landscape of small-sized crop fields and semi-natural habitats maximizes 47 48 440 arthropod diversity and may decrease the probability of overall extinction even of rare species 49 50 441 (Duelli 1990). 51 52 53 54 442 The pool of harvestman species studied in mixed managed and natural habitats is not in danger 55 56 443 in the low Cantabrian area. Furthermore, these species have also been found in mountain 57 58 444 Orocantabrian areas, which ensures their persistence in the region during future years, bearing 59 60 61 62 17 63 64 65 445 in mind that they have been shown to live at different altitudes and many of them in a variety of 1 2 446 habitats. 3 4 5 447 The harvestmen assemblages in low Cantabrian areas, -of different habitats and different 6 7 8 448 clusters-, were closer to each other than in Muniellos, where some groups of habitats were 9 10 449 clearly separated from each other in the analyses such as CA and MDS. The absolute exception 11 12 450 in low Cantabrian areas was the distinguished position of grassland assemblages, widely 13 14 451 separated from all the other assemblages in all the analyses performed, despite the fact that the 15 16 17 452 grasslands studied were spread across 3 different municipalities. 18 19 20 453 The different distribution of harvestman species in areas that are not widely separated, like 21 22 454 mountain Cantabrian and low Cantabrian territories warns against making easy generalizations 23 24 and shows that regional or even local features must be considered in conservation policies. 25 455 26 27 28 456 References 29 30 457 Clarke KR, Gorley RN (2006) PRIMER v6: User Manual/Tutorial. PRIMER-E Ltd. Plymouth 31 32 33 458 Curtis DJ, Machado G (2007) Ecology. In: Pinto-da-Rocha R, Machado G, Giribet G. (eds) 34 35 36 459 Harvestmen. The biology of Opiliones. 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49 582 50 51 583 52 53 584 54 55 585 56 57 586 58 59 60 587 61 62 23 63 64 65 588 Figure captions 1 2 589 Fig. 1 3 4 590 Location of sampling places and localities. Centre: 22 sampling points (black triangles) around 5 6 591 the city of Oviedo and their local zones, numbered according to Table 1. Lower left corner, 7 8 9 592 Iberian Peninsula with the communities of Asturias and Cantabria delimited. Upper right corner 10 11 593 Asturias and its municipalities. Black crosses: six localities whose harvestmen were compared: 12 13 594 PI Parque Integral Natural; PNO, Parque Nacional de Ordesa y Monte Perdido; SJP, Macizo de 14 15 595 San Juan de la Peña, PNM, Parque Natural del Montseny and RPM, Rèserve Naturelle de Prats- 16 17 18 596 de-Mollo. 19 20 597 Fig. 2 21 22 598 Correspondence analysis of the sampling sites and their harvestman species. Grasslands are 23 24 599 within the left ellipse. Most sites are within the right ellipse. Abbreviated and complete name of 25 26 the sites and their characteristics are in Table 1. Harvestman species complete names are in the 27 600 28 29 601 text. 30 31 602 Fig. 3 32 33 603 Cluster analysis of harvestman assemblages of the sites, obtained with abundance data. 34 35 604 Abbreviated and complete name of the sites and their characteristics are in Table 1. The names 36 37 38 605 of each cluster are below. 39 40 606 Fig. 4 41 42 607 Multidimensional scaling (MDS) of the harvestman assemblages of the sites studied. Each type 43 44 608 of habitat has a different tag and the clusters of Figure 3 are marked. 45 46 47 609 48 49 610 50 51 611 52 53 612 54 55 56 613 57 58 614 59 60 615 61 62 24 63 64 65 616 1 2 617 3 4 618 5 6 619 Table 1. Position and characteristics of the sites studied. Ab., abbreviation; (Nar), Naranco; 7 (Ul), Ules; (Vle), Violeo; (Bra), Brañes; (Aju), Ajuyán; (Ovi), Oviedo; (Mur), Muros de Nalón; 8 620 9 621 (Vio), Vioño; DD Coo., Decimal Degrees coordinates; m: altitude in meters. Sites beginning 10 622 with E, boundary; F, forest; G, grassland; P, forest plantation; S, shrubland; Y, Young forest. 11 623 12 Site Ab. DD Coo. Habitat Phytosociological association m 13 1.- SbN (Nar) Shrub: furze Ulici europaei-Genistetum occidentalis 460 14 43.3815,-5.8573 15 2.- SgN (Nar) Shrub: heather-gorse Ulici europaei-Ericetum vagantis 547 16 43.3836,-5.8535 17 3.-YpN (Nar) Young forest: Pyrenean oaks Previous to Blechno spicanti-Quercetum roboris facies Q. pyrenaica 540 43.3835,-5.8536 18 4.-PoU (Ul) For. Plantation: oaks & chestnuts Corresponding to Polysticho setiferi-Fraxinetum excelsioris 350 19 43.3785,-5.8869 20 5.PooU (Ul) For. Plantation old: oaks & chestnuts Corresponding to Polysticho setiferi-Fraxinetum excelsioris 379 21 43.3793,-5.8880 22 6.-EhU (Ul) Herb. border: horsetail Picridio hieracioides-Eupatorietum cannabini subasociation equisetosum telmateia 355 43.3786,-5.8874 23 7.- PecU (Ul) For. Plantation: eucalyptus Eucalyptus globulus 363 24 43.3788,-5.8893 25 8.- ShV (Vle) Shrub: heather Ulici europaei-Ericetum vagantis 428 26 43.3922,-5.9087 27 9.-YwV (Vle) Young forest: willow trees Betula-Salicetum atrocinerea 423 43.3922,-5.9092 28 10.- SgV (Vle) Shrub: gorse edge Ulici europaei-Ericetum vagantis 421 29 43.3929,-5.9092 30 11.- FeV (Vle) Forest: eutrophous forest Polysticho setiferi-Fraxinetum excelsioris 418 31 43.3927,-5.9082 32 12.-PchV (Vle) For. plantation: chestnut trees Derived from Blechno spicanti-Quercetum roboris facies Q. pyrenaica 411 43.3939,-5.9087 33 13.-YpV (Vle) Young forest: Pyrenean oaks Previous to Blechno spicanti-Quercetum roboris facies Q. pyrenaica 339 34 43.3948,-5.9133 35 14.- FolV (Vle) Forest: oligotrophous forest Blechno spicanti-Quercetum roboris facies Q. pyrenaica 354 36 43.3981,-5.9131 15.-FeB (Bra) Forest: gallery eutrophous forest Polysticho setiferi-Fraxinetum excelsioris 126 37 43.4117,-5.9156 38 16.- FmB (Bra) Forest: mixed forest transition Polysticho setiferi-Fraxinetum excelsioris to Hyperico androsaeni- Alnetum glutinosae 126 39 43.4112,-5.9163 40 17.- FaB (Bra) Forest: gallery alder tree forest Hyperico androsaeni- Alnetum glutinosae 125 41 43.4113,-5.9164 18.- YhA (Aju) Young forest: hazel-nut tree forest Rubu ulmifoli- Tametum communis 242 42 43.4083,-5.8987 43 19.-YlA (Aju) Young forest: bays Hedero helicis-Lauretum nobilis 231 44 43.4098,-5.8960 45 20.-GcA (Aju) Grassland with cherry tree Lino biennis-Cynosuretum cristati 226 46 43.4098,-5.8941 21.- FeA (Aju) Forest: eutrophous forest Polysticho setiferi-Fraxinetum excelsioris 199 47 43.4125,-5.8925 48 22.-PoA (Aju) For. plantation: oaks & chestnuts Derived from Polysticho setiferi-Fraxinetum excelsioris 244 49 43.4055,-5.8923 50 23.-EnO (Ovi) Herb. border: nettle-elder tree Urtico dioicae-Sambucetum ebuli 224 51 43.3525,-5.8551 24.-GO (Ovi) Grassland: meadow Lino biennis-Cynosuretum cristati Subassociation brometosum erecti 314 52 43.3556,-5.8744 53 25.-GaM (Mur) Grassland with apple trees Lino biennis-Cynosuretum cristati 115 54 43.5545,-6.0916 55 26.- GM Mur) Grassland: meadow Lino biennis-Cynosuretum cristati 115 56 43.5542,-6.0920 27.-G1Vi (Vio) Grassland: meadow Lino biennis-Cynosuretum cristati 43 57 43.3616,-3.9780 58 28.-G2Vi (Vio) Grassland: meadow Linno biennis-Cynosuretum cristati 46 59 43.3613,-3.9773 60 624 61 62 25 63 64 65 625 1 2 626 3 4 627 5 6 628 Table 2. Accumulation curves and Clench equation parameters. Sampling sites as in Table 1 7 8 9 629 (number of samples with harvestman specimens in brackets); S harvestman species richness; 10 2 11 630 SEXP (a/b), expected richness according to Clench equation; r , determinant coefficient; q 12 13 631 proportion of the inventory; % sampling efficiency percentage; f.s. final slope of the curve; E.V. 14 15 632 explained variance. 16 17 2 18 Sampling sites S SEXP (a/b) r q % f.s E.V. 19 20 SbN (25) 11 11.73 0.99 0.94 93.78 0.04 98.98 21 SgN (24) 12 13.57 0.99 0.9 90.36 0.05 99.91 22 YpN (25) 10 11.2 0.99 0.89 89.2 0.04 99.8 23 24 PoU (23) 8 8.57 0.99 0.93 93.35 0.03 99.27 25 PooU (24) 9 10.23 0.99 0.88 87.98 0.05 99.8 26 EhU (24) 12 13.77 0.99 0.9 90.7 0.05 99.9 27 28 PecU (24) 11 12.28 0.99 0.89 89.57 0.04 99.73 29 ShV (25) 9 9.64 0.99 0.93 93.36 0.02 98.77 30 31 YwV (25) 11 11.8 0.96 0.93 93.22 0.02 96.08 32 SgV (24) 11 11.8 0.99 0.93 93.22 0.04 99.99 33 FeV (16) 9 12.43 0.99 0.72 72.4 0.16 99.5 34 35 PchV (16) 6 6.7 0.99 0.89 89.55 0.04 99.7 36 FolV (25) 8 8.32 0.99 0.96 96.15 0.02 99.46 37 38 YpV (25) 11 11.72 0.99 0.94 93.86 0.03 99.58 39 FeB (19) 8 9.07 0.99 0.88 88.23 0.05 99.82 40 FmB (22) 11 14.48 0.99 0.76 75.97 0.13 98.78 41 42 FaB (24) 12 14.53 0.99 0.83 82.59 0.09 99.47 43 YhA (23) 8 9.05 0.99 0.88 88.4 0.03 99.54 44 45 YlA (21) 11 13.29 0.99 0.83 82.77 0.09 99.98 46 GcA (25) 9 9.45 0.99 0.95 95.24 0.01 99.49 47 FeA (23) 7 7.69 0.99 0.9 91.03 0.03 99.65 48 49 PoA (19) 8 10.88 0.98 0.73 73.53 0.12 98.25 50 EnO (24) 12 13.59 0.99 0.88 88.3 0.07 98.56 51 GO (24) 8 8.36 0.99 0.96 95.69 0.02 98.66 52 53 GaM (25) 10 10.01 0.96 0.99 99.9 0.01 95.57 54 GM (25) 11 11.74 0.99 0.94 93.7 0.03 98.98 55 56 G1Vi (23) 7 7.28 0.99 0.96 96.15 0.02 98.93 57 G2Vi (23) 8 8.76 0.97 0.91 91.32 0.04 97.43 58 633 59 60 61 62 26 63 64 65 634 1 2 635 3 4 636 5 6 637 Table 3. Relative abundance percentage (Abu) and relative frequency percentage (Fr) of the 7 8 9 638 harvestman species found in low Cantabrian areas (LC) and Muniellos Reserve (Mu). European 10 11 639 (Eu), Holarctic (Ho) or Iberian endemic (IE) species distribution. 12 13 640 14 15 16 Species Abu LC Abu Mu Fr LC Fr Mu 17 1 Anelasmocephalus cambridgei Eu 1.4 0.5 79 11 18 2 Gyas titanus Eu 0.05 0.4 7 5 19 20 3 Hadziana clavigera EI 0.06 0.3 11 5 21 4 Homalenotus laranderas EI. 21.6 7.3 64 47 22 5 Homalenotus quadridentatus Eu 27.7 0 47 0 23 24 6 Ischyropsalis hispanica EI 2.27 3.5 61 53 25 7 Leiobunum blackwalli Eu 11.8 11.8 100 58 26 8 Leiobunum rotundum Eu 1.2 10 61 47 27 28 9 Nemastoma hankiewiczii EI 4.9 2.6 79 21 29 10 Nemastomella dentipatellae EI 5.8 0.5 93 5 30 11 Odiellus seoanei EI 0.6 3.3 7 47 31 32 12 Odiellus simplicipes EI 9.2 6.7 58 53 33 13 Paroligolophus agrestis Ho. 2 23.3 32 53 34 14 Phalangium opilio Ho. 2 14.3 82 74 35 36 15 Sabacon franzi EI 1 0.9 61 16 37 16 Trogulus nepaeformis s.l. Eu 8.5 6.7 100 42 38 17 Dicranopalpus sp. 0 0.3 0 11 39 40 18 Oligolophus hansenii (Kraepelin, 1896) EU 0 7.3 0 58 41 19 Paramiopsalis ramblae Benavides & Giribet, 2017 IE 0 0.1 0 5 42 20 Megabunus diadema (Fabricius, 1779) Eu 0 0.1 0 5 43 44 European species combined 50.6 36.6 45 Iberian Endemic species combined 45.5 25.1 46 Holarctic species combined 3.9 37.5 47 48 641 49 50 642 51 52 643 53 54 644 55 56 57 645 58 59 646 60 61 62 27 63 64 65 647 1 2 648 3 4 649 5 6 650 7 8 9 651 10 11 652 12 13 653 14 15 654 16 17 18 655 19 20 656 21 22 657 23 24 658 25 26 Table 4. Number of harvestmen at each site and global diversity values. Lower lines: 27 659 28 660 abundance and number of sites for each species. Sites described by their abbreviations in Table 29 661 1 and ordered according to the cluster analysis in Figure 3. Sites beginning with E boundary; F, 30 662 forest; G, grassland; P, forest plantation; S, shrubland; Y Young forest. 31 32 663 33 34 664 35 36 665 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 28 63 64 65 666 1 2 667 3 4

5

6

7 s.l.

8

9

10

titanus

11

Sites

12 Cluster

oma hankiewiczii

t

sp. richness

Gyas

Diversity 1/λ

13 tot abundance

Sabacon franzi

Odiellus seoanei

Phalangium opilio

Hadziana clavigera

14 Odiellus simplicipes

Leiobunum rotundum

Leiobunum blackwalli

Ischyropsalis hispanica

Paroligolophus agrestis

Nemas Homalenotus laranderas

15 Trogulus nepaeformis

Nemastomella dentipatellae

Homalenotus quadridentatus

16 Anelasmocephalus cambridgei 17 18 G1Vi 46 9 8 2 6 944 101 1116 7 1.38 19 G2Vi 1 44 7 4 5 3 720 235 1019 8 1.8 20

21 GM 6 4 77 23 4 125 5 17 1 551 613 1426 11 2.9 22 GcA 11 44 184 32 52 12 7 866 609 1817 9 2.84 23 GaM 3 1 367 230 7 129 58 30 192 240 1257 10 5.21 24

A25 GO 19 115 2 19 1 54 643 54 907 8 1.9 26 ShV 50 151 17 105 39 7 5 97 6 477 9 4.77 27

28 SbN 4 13 54 12 48 94 7 33 2 2 82 351 11 5.54 29 SgN 4 8 5 61 119 27 15 1 12 2 1 12 267 12 3.71 30 31 YpN 183 5 22 68 45 3 20 3 9 5 363 10 3.2 32 YwV 15 6 17 13 47 50 49 12 23 11 20 263 11 7.67 33 SgV 11 8 9 30 72 56 8 40 2 3 17 256 11 5.68 34 C2.135 YpV 15 9 18 8 60 109 30 53 1 36 339 11 5.58 36 EnO 2 1 12 72 27 70 1 12 5 4 33 7 246 12 4.92 37 38 EhU 1 2 1 24 39 56 10 16 7 16 7 8 187 12 5.81 39 PecU 1 3 1 16 115 27 2 12 9 6 8 200 11 2.74 40 PooU 14 5 34 52 11 4 1 1 9 131 9 4.31 41

C2.242 PoU 2 57 31 48 7 19 1 4 169 8 4.11 43 PoA 1 2 37 2 4 1 17 1 65 8 2.5 44

45 FolV 1 3 14 13 160 145 28 15 379 8 2.99 46 YlA 5 1 6 4 13 12 14 55 4 11 1 126 11 4.23 47 48 FeA 7 3 12 12 29 13 5 81 7 4.75 49 FmB 1 13 1 101 1 38 6 6 2 2 171 11 2.51 50 FaB 1 1 1 8 75 2 43 12 8 3 3 4 161 12 3.32 51 52 FeB 11 20 42 39 13 45 1 171 8 5.03

C153 YhA 4 3 5 24 85 22 9 18 170 8 3.3 54 55 FeV 1 3 1 2 6 5 22 1 41 9 3.13 56 B57 PchV 1 11 20 10 10 52 6 4.01

58 Abun. 130 238 277 238 1126 1434 1034 594 705 170 151 3382 2641 8 6 74 12208 59 60 Sites 17 9 17 23 16 28 28 22 26 22 17 13 18 3 2 2 61 62 29 63 64 65 668 1 2 669 3 4 670 5 6 671 7 8 9 672 10 11 673 12 13 674 Table 5. Harvestman average abundance, average species richness and average true diversity 14 675 (1/λ) of the clusters and the habitats. N, number of sites studied. 15 16 17 Clusters N Abundance Richness 1/λ 18 Cluster A 6 1257 8.8 2.7 19 Cluster B 2 46.5 7.5 3.6 20 21 Cluster C1 8 165.5 9.1 3.6 22 Cluster C2.1 7 330.9 10.8 5.2 23 24 Cluster C2.2 5 186.6 10.4 4.4 25 Habitats 26 Grasslands 6 1257 8.8 2.7 27 28 Shrublands 4 337.8 10.8 4.9 29 Young forests 5 252.2 10.2 4.8 30 31 Forest plantations 5 123.4 8.4 3.5 32 Forests 5 167.2 9.2 3.6 33 Boundaries 2 216.5 12 5.4 34 35 676 36 37 677 38 39 678 Table 6. Analyses of similarities ANOSIM with Bray-Curtis index of dissimilarity between the 40 41 679 harvestman assemblages of the aggregate sites for each habitat. S significance level, *, 42 43 44 680 differences error ≤ 0.05); **, differences error ≤ 0.01). % dissimilarity percentages between the 45 46 681 harvestman assemblages based on SIMPER analyses. 47 48 Grasslands Shrublands Y. forests Bound. Forests 49 50 S % S % S % S % S % 51 Shrublands 0.005 ** 62.1 52 53 Y. forests 0.002 ** 64.4 0.452 41.0 54 Boundaries 0.036 * 57.1 0.133 37.1 0.714 33.8 55 56 Forests 0.002 ** 73.8 0.005 ** 58.4 0.048 * 46.4 0.25 44.2 57 For. Plant. 0.002 ** 70.5 0.032 * 53.1 0.246 47.7 0.81 41.6 0.013 * 50.8 58 682 59 60 61 62 30 63 64 65 683 1 2 684 3 4 685 5 6 686 7 8 9 687 10 11 688 Table 7. Indicator species of the clusters of sites and the habitats. B, boundaries populated with 12 689 horsetail or nettle; GRASS, grasslands, SR, shrubland, YF, young forest. Ind. v., indicator 13 690 value; p, probability; s.l., significance level. 14 15 Indicator species Clusters Ind. v. p s.l. 16 17 Homalenotus quadridentatus A 0.916 0.002 ** 18 Homalenotus laranderas A + C2.1 0.993 0.001 *** 19 Paroligolophus agrestis C2.1 0.866 0.028 * 20 21 Leiobunum rotundum C1 + C2.2 0.882 0.012 * 22 Odiellus simplicipes A + C2.1 + C2.2 0.876 0.013 * 23 24 Habitats 25 Homalenotus quadridentatus B + GRASS 0.998 0.001 *** 26 Homalenotus laranderas B + GRASS + SR 0.985 0.002 ** 27 28 Ischyropsalis hispanica B + SR + YF 0.983 0.001 *** 29 Sabacon franzi SR + YF 0.874 0.03 * 30 31 691 32 33 692 34 35 693 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 31 63 64 65 Figure 1 pdf Click here to download Figure Fig 1 Asturias.pdf

Muros del Nalón + Illano Oviedo + PIMuniellos Ajuyán 21 ASTURIAS Brañes 16 20 19 15 18 N 17 22

200 14 13 12 400 11 1 km El Violeo 10 9 8 500 Naranco 2 3 7 6 5 1 Ules 4

FRANCE PNO Vioño + + + RPM SJP + PNM SPAIN OVIEDO 24 El Cristo 23 Parque de Invierno Figure 2 EPS Click here to download Figure Merino Anadon Fig 2.eps Figure 3 EPS Click here to download Figure Merino Anadon Fig 3.eps Figure 4 EPS Click here to download Figure Merino Anadon Fig 4.eps