NORTH-WESTERN JOURNAL OF ZOOLOGY 12 (2): 255-260 ©NwjZ, Oradea, Romania, 2016 Article No.: e161301 http://biozoojournals.ro/nwjz/index.html
Influence of habitat structure on the spiders of river islands and floodplain for- ests of the lower reach of the Mures River in Western Romania
Róbert GALLÉ1,2*, Nikolett SZPISJAK1 and Attila TORMA1
1. University of Szeged, Department of Ecology, Szeged, Hungary. 2. Sapientia Hungarian University of Transylvania, Department of Environmental Sciences, Cluj-Napoca, Romania. *Corresponding author, R. Gallé, E-mail: [email protected]
Received: 21. December 2014 / Accepted: 01. February 2016 / Available online: 20. April 2016 / Printed: December 2016
Abstract. Floodplain forests are among the most threatened habitats in Europe and contain a highly diverse and characteristic fauna. We tested the effect of landscape and habitat attributes on the structure of spider assemblages of river islands and floodplain forests along the Mures River, Western Romania. The fieldwork was carried out in seven islands and seven riverbank forest sites. Spiders were collected with pitfall traps. We found significant association between habitat structure and rarefaction diversity and species composition of spider assemblages. The type of the sites (i.e. island vs. riverbank) also influenced the composition of the spider assemblages. However, there was no relationship between the landscape attributes and the spider assemblages, presumably due to the presence of large non-flooded forests in the study area, which may serve as overwintering habitat for the floodplain fauna. The present study emphasizes the importance of protecting river islands and floodplain forests for preservation of the regional species richness.
Key words: Araneae, forest, flooding, habitat structure, landscape.
Introduction ments (Ziesche & Roth 2008, Oxbrough et al. 2010). In general, forest spiders are negatively in- Natural and semi-natural floodplains have a high fluenced by the decreasing structural complexity, diversity of both plant and animal species (Roth- which possibly causes a reduction of microhabitat enbücher & Schaefer 2006). In floodplains, flood- diversity or resources (Willett 2001, Finch 2005). In ing events play a key role in shaping and main- addition to habitat structure, the diversity and taining the complex mosaic of riparian habitats composition of spider assemblages are influenced (Hughes & Rood 2003). Flooding events represent by landscape attributes and interspecific interac- a challenge for terrestrial invertebrates, which tions (e.g. Schmidt et al. 2008, Herrmann et al. therefore develop special survival strategies 2010, Wise 1999). (Rothenbücher & Schaefer 2006, Pétillon et al. Faunistical data on spiders of the Western part 2009) to cope with the long periods of submersion of Romania is relatively scarce, however data is (Pekár 1999). Floodplains are among the most di- available on the Western Romanian lowland verse and threatened ecosystems world-wide (Duma 2006a,b, 2008, 2012, Fetykó 2008, Gallé et (Lambeets et al. 2009, Gallé et al. 2011, Torma & al. 2012, Sas-Kovács et al. 2013, 2014, 2015a,b, Sas- Császár 2013). However, most European river val- Kovács & Sas-Kovács 2014a,b). Previous studies leys have been altered by river regulations during considering the spider fauna of the floodplain the last 150 years and about 90% of the former along the river Tisza (the Mures falls into the Tisza floodplains have been degraded or functionally at Szeged, Hungary) indicated that besides flood- destroyed (Tockner et al., 2009). Thus, the remain- ing regime, habitat type and landscape scale fac- ing natural and semi-natural lowland floodplain tors affect assemblage structure of spiders at the habitats are of high conservational value, includ- scales between 250 and 750 m, with a peak at 500 ing their arthropod fauna, which consists of spe- m buffer (Gallé et al. 2011, Gallé & Schwéger cialist and rare species (Rothenbücher & Schaefer, 2014). Here we test the relationship between spi- 2004, Sabo et al. 2005, Gallé et al. 2011, Torma & der assemblages and the habitat structure and Császár 2013). landscape composition in river islands and allu- Spiders are among the most diverse inverte- vial forests. brate predators in terrestrial ecosystems (Wise 1993). The distribution of forest spiders is affected Material and methods by a distinct complex of environmental factors with respect to species-specific habitat require- Spider sampling was performed on a 20 km long section 256 R. Gallé et al. of the Mures River, near Pesica, Romania (Fig. 1.). The vegetation cover at 10 cm height (R2=0.795, t=-6.840, study area is situated approximately 100 m a.s.l. and the p<0.001), vegetation cover at 40 cm height (R2=0.656, t=- mean annual precipitation is 550–600 mm (Andó 1995, 4.789, p<0.001), vegetation height (R2=0.302, t=-2.282, Erdős et al. 2012). The soil in the region is grey soil with p=0.041), and positively correlated with the cover of leaf high clay content (Jakab 1995). The river runs out its bed litter (R2=0.851, t=8.337, p<0.001), canopy closure and invades the floodplain for 6 days/year in average (R2=0.850, t=8.285, p<0.001). Thus, lower values represent (Sipos et al. 2011). The river regulations in the 19th century sites with highly structured and dense understory vegeta- significantly decreased the area of the floodplain and the tion, whereas higher values represent closed and shaded length of the river, resulting in a higher and steeper slope forests with high amount of leaf litter. The flooding re- of the riverbed and decreased number of meanders. The gime was similar at all sites, thus the effect of flooding modified river dynamics and flooding regimes are in was not analysed. close relation with the decreasing number of the river is- To assess the composition of the landscape, the pro- lands (Sipos et al. 2011). However, numerous islands are portion of forests was measured in a radius of 500 meters present in the study area. around each site, using QGIS software and satellite im- ages, as several studies suggest that landscape composi- tion at scales of 500 m radius can be relevant for spiders (Schmidt et al. 2005, Gallé & Schwéger 2014). The link between habitat structure (PC1), landscape composition (proportion of forests) and location (island vs. riverbank) and species richness was tested with gen- eral linear models (GLM) with log-link function. The negative binomial error term was used because of the overdispersion of the data. Large between-site variation was found in the num- ber of collected individuals (63.2 ± 40.6, mean ± SD), thus rarefaction diversity was used to standardize the number
Figure 1. Map of the study region. The dotted line repre- of species recorded within each of the sampling sites sents the border between Hungary and Romania, for- (Gotelli & Colwell, 2001).The effects of habitat structure ests are marked with grey and the dashed line repre- (PC1), landscape composition (proportion of forests) and sents the study site. The island-bank sampling pairs are location (island vs. river bank) on the rarefaction diversity marked with open circles. of spider assemblages were tested with linear regression. The influential points were identified with the Cook’s dis- tance plot and were excluded from further analysis. A total of 14 study sites were established, comprising Stepwise model selection on the basis of Akaike Informa- seven islands and seven riverbank sites near the islands tion Criterion was used to identify the best predictive with similar flooding regime. Spiders were sampled with model. pitfall traps. Pitfall traps measure the activity-density of To explore which of the environmental variables epigeic species, thus they capture ground-dwelling spi- mentioned above were most important in shaping the ders more efficiently (Topping & Sunderland 1992, Urák species composition of ground-dwelling spider assem- & Samu 2008). Three traps were installed at each of the blages Multivariate Regression Tree (MRT) analysis sites with 5 meters distance between them (Ward et al. (De’ath, 2002) was used. This method can be used to iden- 2001). The traps consisted of plastic jars (diameter of the tify relationship between species abundances and ex- opening 9 cm) filled up to one third with a 70% ethylene planatory variables and also explore interactions between glycol solution. Trapping was done continuously from explanatory variables (De’ath, 2002, 2010). To visualize 15th June to 6th July, 2012, (22 days). To characterize the the results of the MRT analysis, a Principal Component structure of the habitat, the percentage cover of the litter, analysis (PCA) was performed where the site were used bare ground, herbaceous vegetation at the ground level, as objects and species abundances as variables. The char- at 10 and at 40 cm height and the average height of the acteristic species for each MRT group was explored by the herbaceous vegetation were measured within four 1 × 1 IndVal (Indicator Value) procedure (Dufrêne and Legen- meter quadrates at each site. We did not measure litter dre, 1997). The statistical significance was calculated by depth as the regular flooding events redistribute leaf litter 1000 random Monte Carlo permutations. (Xiong & Nilsson 1997). We measured the closure of the All statistical analyses were performed using the R canopy using digital photographs of the canopy taken software (R Core Team 2014) with MVPART (De’ath from the ground level. The photographs were analysed in 2010), labdsv (Roberts 2013) and Vegan (Oksanen et al. the laboratory with visual assessment of the proportion of 2013) packages. leaves. As the habitat parameters were highly correlated, a PCA analysis was performed and the PC1 axis was used as a novel variable to characterize habitat structure. The Results PC1 axis explained 77.4% of the total variance and signifi- cant negative correlation was found with vegetation cover at the ground level (R2=0.968, t=-19.186, p<0.001), A total of 851 adult spiders belonging to 56 species Spiders fauna of Mures floodplain 257 were collected during the study (Table 1). The Discussion most abundant species were Ozyptila praticola (C.L. Koch, 1837), Pardosa lugubris (Walckenaer, 1802) In the present study, we have assessed the compo- and Phrurolithus festivus (C.L. Koch, 1835). sitional differences of spider assemblages of is- We did not find significant association be- lands and riverbank and the effects of habitat tween the predictor variables and the species rich- structure and landscape properties on the rarefac- ness of spiders according to the general linear tion diversity and composition of spider assem- model. Habitat structure and proportion of forests blages. were included in the best predictor model for the Floodplain forests provide great habitat diver- rarefaction diversity of spiders. Significant effect sity and in turn have high species diversity of habitat structure was found (β=0.0276, t=3.167, (Buchholz 2009), in the present study 58 species P=0.013). However we did not confirm the effect were collected. Gallé & Schwéger (2014) collected of proportion of forests (β=-1.383, t=-1.615, NS.). 42 species in floodplain forests at the lower reach The species composition was affected by the of river Tisza and also found that P. alacris and location of sampling sites and, in the case of is- O.praticola are among the most abundant spider lands, the habitat structure had a major effect t ac- species, emphasizing the relatively similar species cording to the Multivariate Regression Tree analy- composition of dominant spider species of flood- sis (Fig. 2). The subsequent PCA showed the sepa- plain forests. In the case of regularly flooded habi- ration of leaf 2, islands with relatively simple tats, only the specialized species may be able to vegetation structure and closed canopy. survive short-term disturbances (Rothenbücher & Schaefer 2006), and when the magnitude of dis- turbance is higher, only highly dispersive species occur due to repeated colonization (Lambeets et al. 2008). Out of the differential species, P. festivus was a significant indicator of islands with rela- tively complex vegetation structure and open can- opy, this generalist species is known to occur in different habitat types, e.g. in sunny forests near water banks (Nentwig et al. 2015). N. reticulatus indicated islands with relatively simple vegetation structure and closed canopy, this species being abundant on the ground layer of different forest types (Buchar & Ruzicka 2002). The present study demonstrated, that the floodplain spider fauna of the Mures River is in- fluenced by habitat structure rather than land- scape composition, however the location of the study site had an effect on the species composition of the assemblages. Species composition of the vegetation and habitat structure are determined
by flooding (Ballinger et al. 2005, Plum, 2005). The Figure 2. Multivariate Regression Tree analysis (a) and habitat structure and flooding events are con- the subsequent Principal Component Analysis (b). Black straints for spider diversity, indicating the impor- circles represent the study sites of leaf 1, open circles tance of the local habitat attributes (Lambeets et al. leaf 2 and grey circles leaf 3, respectively. 2006, Gallé et al. 2011, Gallé & Schwéger 2014). Flooding disturbance appears to be especially Significant indicator values were found for P. relevant in shaping the invertebrate fauna of river- festivus on islands with relatively complex vegeta- ine landscapes. This localized disturbance is im- tion structure and open canopy (third MRT leaf, portant to facilitate the occurrence of specialized IndVal=0.803, P=0.042) and Neon reticulatus species and thus increases the regional diversity (Blackwall, 1853) only found on islands with rela- (Robinson et al. 2002, Lambeets et al. 2008, Gallé et tively simple vegetation structure and closed can- al. 2011, Gallé & Schwéger 2014). Numerous stud- opy (second MRT leaf, IndVal=1, P=0.003.) ies have demonstrated that anthropogenic altera-
Table 1. Habitat parameters and list of species, see text for details.
Habitat type (island:1, floodplain:0) 0 1 1 0 1 0 1 0 1 0 1 0 1 0 PC1 -44 42 -10 32 -6 -93 3.7 0.6 -19 6.5 29 11 17 30 Proportion of forests 0.8 0.8 0.5 0.6 0.6 0.6 0.8 0.8 0.3 0.3 0.4 0.4 0.5 0.6 Spider species richness 18 13 9 12 17 6 12 7 10 14 9 13 18 6 Spider abundance 117 50 49 50 151 21 112 38 52 64 23 91 58 9 Rarefaction diversity 5.8 6.1 3.7 4.6 4.5 4.1 4.2 4.1 4.3 5.4 5.2 4.7 5 6 Dysdera hungarica Kulczynski, 1897 3 0 1 0 0 0 0 0 0 0 0 0 0 0 Ceratinella brevis (Wider, 1834) 0 1 0 0 0 0 0 0 0 0 0 0 0 0 Diplocephalus cristatus (Blackwall, 1833) 1 0 0 0 0 0 0 0 0 0 0 0 0 0 Diplocephalus picinus (Blackwall, 1841) 0 1 0 0 0 0 0 0 1 1 0 0 0 0 Diplostyla concolor (Wider, 1834) 2 0 0 0 1 2 2 2 0 0 0 1 1 1 Maso sundevalli (Westring, 1851) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Meioneta rurestris (C.L. Koch, 1836) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Neriene clathrata (Sundevall, 1830) 0 0 0 0 0 0 0 0 0 1 0 0 0 0 Tenuiphantes flavipes (Blackwall, 1854) 0 0 0 1 0 0 0 0 0 0 1 0 1 1 Pelecopsis radicicola (L. Koch, 1872) 0 4 0 1 0 0 0 0 0 0 0 0 0 0 Walckenaeria alticeps (Denis, 1952) 1 0 0 0 4 2 0 0 0 1 1 0 1 0 Walckenaeria cucullata (C.L. Koch, 1836) 0 0 0 0 0 0 0 0 0 0 0 0 1 0 Pachygnatha degeeri Sundevall, 1830 0 0 0 0 1 0 0 0 0 0 0 0 0 0 Pachygnatha listeri Sundevall, 1830 1 0 0 0 1 0 0 0 0 0 0 0 0 0 Episinus angulatus (Blackwall, 1836) 0 0 1 0 0 0 0 0 0 0 0 0 0 0 Robertus lividus (Blackwall, 1836) 1 0 0 0 0 0 0 0 0 0 0 0 0 0 Arctosa leopardus (Sundevall, 1833) 0 0 0 0 0 0 0 0 0 0 0 0 1 0 Arctosa lutetiana (Simon, 1876) 0 4 0 0 0 0 0 0 0 3 0 0 1 0 Pardosa agrestis (Westring, 1862) 0 0 1 1 0 0 1 0 2 0 0 1 0 0 Pardosa agricola (Thorell, 1856) 0 0 0 0 0 0 2 0 0 0 0 0 0 0 Pardosa amentata (Clerck, 1757) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Pardosa lugubris (Walckenaer, 1802) 23 13 0 23 11 1 10 15 1 9 1 39 1 2 Pardosa prativaga (L. Koch, 1870) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Pirata hygrophilus Thorell, 1872 7 0 0 1 2 0 0 2 0 0 0 0 0 0 Pirata latitans (Blackwall, 1841) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Trochosa ruricola (De Geer, 1778) 5 0 0 1 43 0 1 0 1 0 0 1 1 0 Trochosa terricola Thorell, 1856 2 3 1 1 4 0 0 0 0 0 1 3 4 1 Xerolycosa miniata (C.L. Koch, 1834) 0 0 0 0 0 0 2 0 0 0 0 0 0 0 Pisaura mirabilis (Clerck, 1757) 0 0 0 0 0 0 0 0 0 0 0 0 1 0 Agelena labyrinthica (Clerck, 1757) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Phrurolithus festivus (C.L. Koch, 1835) 11 0 2 1 37 3 40 0 26 1 7 16 0 0 Urocoras longispinus (Kulczyński, 1897) 0 0 0 0 0 0 0 0 0 0 1 0 0 0 Agroeca brunnea (Blackwall, 1833) 0 2 0 0 0 0 0 0 0 0 0 0 0 0 Agroeca cuprea Menge, 1873 0 0 0 0 0 0 0 0 0 0 0 0 1 0 Liocranoeca striata (Kulczyński, 1882) 0 0 0 0 1 0 0 0 0 0 0 0 0 0 Scotina celans (Blackwall, 1841) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Clubiona lutescens Westring, 1851 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Clubiona pallidula (Clerck, 1757) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Zodarion germanicum (C.L. Koch, 1837) 12 0 18 3 1 1 0 0 1 6 0 7 0 0 Drassyllus pusillus (C.L. Koch, 1833) 1 0 0 0 1 0 2 0 0 1 0 1 1 0 Drassyllus villicus (Thorell, 1875) 1 0 0 0 1 0 0 0 1 2 0 1 2 0 Haplodrassus minor (O.P.-Cambridge, 1879) 0 0 0 0 0 0 1 0 0 0 0 0 0 0 Micaria pulicaria (Sundevall, 1832) 1 0 0 0 0 0 0 0 0 0 0 0 0 0 Trachyzelotes pedestris (C.L. Koch, 1837) 24 5 3 5 0 0 7 12 0 13 5 6 10 1 Zelotes apricorum (L. Koch, 1876) 2 1 1 2 0 0 0 1 9 2 0 4 3 0 Zelotes gracilis Canestrini, 1868 0 0 0 0 0 0 2 0 0 0 0 0 0 0 Zelotes longipes (L. Koch, 1866) 0 0 0 0 0 0 0 0 0 1 0 0 1 0 Zora spinimana (Sundevall, 1833) 0 0 0 0 0 0 0 1 6 0 0 0 0 0 Ozyptila praticola (C.L. Koch, 1837) 19 8 21 10 29 12 40 5 4 20 5 10 26 3 Xysticus luctator L. Koch, 1870 0 3 0 0 0 0 0 0 0 3 0 1 0 0 Euophrys frontalis (Walckenaer, 1802) 0 0 0 0 1 0 0 0 0 0 0 0 0 0 Euophrys obsoleta (Simon, 1868) 0 0 0 0 1 0 0 0 0 0 0 0 0 0 Myrmarachne formicaria (De Geer, 1778) 0 1 0 0 0 0 0 0 0 0 0 0 0 0 Neon reticulatus (Blackwall, 1853) 0 4 0 0 0 0 0 0 0 0 1 0 1 0
Spiders fauna of Mures floodplain 259 tions of flooding regime and river channelizations References result in a decrease in arthropod abundance and Adis, J., Marques, M.I.M., Wantzen, K.M. (2001): First observations diversity (Laeser et al. 2005, Lambeets et al. 2008, on the survival strategies of terricolous arthropods in the Paetzold et al. 2008). The increased and irregular northern Pantanal wetland of Brazil. Andrias 15: 127-128. Andó, M. (1995): The geography of the River Maros (Mureş) and its flooding induced by the regulation of Central river system. pp. 7-24. In: Hamar, J., Sárkány-Kiss, A. (eds): The European rivers alters the hydrological processes Maros/Mureş River Valley. A study of the geography, and enhances the colonization of generalist spe- hydrobiology and ecology of the river and its environment. Szolnok - Szeged –Tîrgu Mureş. cies, while specialists tend to disappear. The proc- Ballinger, A., MacNelly, R., Lake, P.S. (2005): Immediate and ess results in changes in assemblage composition, longer-term effects of managed flooding on floodplain leading to a more uniform and less specialized invertebrate assemblages in south-eastern Australia: generation and maintenance of a mosaic landscape. Freshwater Biology 50: species composition, and shifts towards higher 1190–1205. dispersal ability (Lambeets et al. 2008, 2009). Buchholz, S. (2009): Community structure of spiders in coastal Landscape attributes influence spider species habitats of a Mediterranean delta region (Nestos Delta, NE Greece). Animal Biodiversity and Conservation 32: 101–115. composition (e.g., Öberg et al. 2007, Batáry et al. Buchar, J, Ruzicka, V (2002): Catalogue of Spiders of the Czech 2008, Schmidt et al. 2008). In the case of lowland Republic Peres Publishers: Praha, Czech Republic. river valleys, Gallé et al. (2011) and Gallé & De’ath, G. (2002): Multivariate regression trees: a new technique for modeling species-environment relationships. Ecology 83: 1105– Schwéger (2014) found significant effect of land- 1117. scape composition within 500 m buffer around the De’ath, G. (2010): mvpart: Multivariate partitioning. R package sampling sites on the diversity assemblage com- version 1.3-1.
Gallé, R., Schwéger, Sz. (2014): Habitat and landscape attributes Sabo, J.L, Sponseller, R., Dixon, M., Gade, K., Harms, T., Heffernan, influencing spider assemblages at lowland forest river valley J., Jani, A., Katz, G., Soykan, C., Watts, J., Welter, J. (2005): (Hungary).North-Western Journal of Zoology 10: 36–41. Riparian zones increase regional species richness by harboring Gotelli, N.J., Colwell, R.K. (2001): Quantifying biodiversity: different, not more, species. Ecology 86: 56-62. procedures and pitfalls in the measurement and comparison of Sas-Kovács, É.H., Urák, I., Sas-Kovács, I. (2013): First record of the species richness. Ecology Letters 4: 379-391. rare species Pardosa maisa Hippa & Mannila, 1982 (Araneae: Herrmann, J.D., Bailey, D., Hofer, G., Herzog, F., Schmidt-Entling, Lycosidae) in Romania. Archives of Biological Science Belgrade M.H. (2010): Spiders associated with the meadow and tree 65: 1605–1608. canopies of orchards respond differently to habitat Sas-Kovács, É.H., Sas-Kovács I. (2014a): Lycosidae (Arachnida: fragmentation. Landscape Ecology 25: 1375-1384. Araneae) in “Câmpia Careiului” (north-western Romania): Hughes, F.M.R., Rood, S.B. (2003): Allocation of river flows for preliminary assessment of composition, distribution, habitat restoration of floodplain forest ecosystems: A review of preference and conservation. North-Western Journal of Zoology approaches and their applicability in Europe. Journal of 10: 102-114. Environmental Management 32: 12–33. Sas-Kovács, É. H., Sas-Kovács, I. (2014b): Note on the distribution Jakab, S. (1995): Soils of the flood plain of the Mureş (Maros) River. of Geolycosa vultuosa (Araneae: Lycosidae) in the “Câmpia pp. 25-47. In: Hamar, J. and Sárkány-Kiss, A. (eds): The Careiului” Natura 2000 site, north-western Romania. Biharean Maros/Mureş River Valley. A study of the geography, Biologist 8: 117-119 hydrobiology and ecology of the river and its environment. Sas-Kovács, É.H., Urák, I., Sas-Kovács, I., Covaciu-Marcov, S.D., Szolnok - Szeged -TîrguMureş. Rákosy, L. (2015a): Winter-active wolf spiders (Araneae: Laeser, S.R., Baxter, C.V., Fausch, K.D. (2005): Riparian vegetation Lycosidae) in thermal habitats from western Romania. Journal loss, stream channelization, and web-weaving spiders in of Natural History 49: 675-683. northern Japan. Ecological Research 20: 646–651. Sas-Kovács, E. H., Sas-Kovács, I., Urák, I. (2015b). Alopecosa Lambeets, K., Bonte, D., Van Looy, K., Hendrickx, F.,Maelfait, J.-P. psammophila Buchar, 2001 (Araneae: Lycosidae): morphometric (2006): Synecology of spiders (Araneae) of gravel banks and data and first record for Romania. Turkish Journal of Zoology environmental constraints along a lowland river system, the 39: 353-358. Common Meuse (Belgium, the Netherlands). In: Deltshev, C., Sipos, Gy., Kiss, T., Oroszi, V. (2011): Geomorphological processes Stoev, P. (eds.): European Arachnology 2005. Acta Zoologica along the lowland sections of the Maros/Mures and Körös/Criş Bulgarica, Suppl.1 : 137-149. Rivers. pp. 97–110.In: Körmöczi L. (ed.): Ecological and socio- Lambeets, K., Vandegehuchte, M.L., Maelfait, J.-P., Bonte, D. (2008): economic relations in the valleys of river Körös/Cris and river Understanding the impact of flooding on trait‐displacements Maros/Mures. Tiscia monograph series, Szeged-Arad. and shifts in assemblage structure of predatory arthropods on Schmidt, M.H., Roschewitz, I., Thies, C. Tscharntke, T. (2005): riverbanks. Journal of Animal Ecoogy 77: 1162–1174. Differential effects of landscape and management on diversity Lambeets, K., Vandegehuchte, M.L., Maelfait, J.P., Bonte, D. (2009): and density of ground-dwelling farmland spiders. Journal of Integrating environmental conditions and functional life history Applied Ecology 42: 281–287. traits for riparian arthropod conservation planning. Biological Schmidt, M.H., Thies, C., Nentwig, W., Tscharntke, T. (2008): Conservation 146: 625–637. Contrasting responses of arable spiders to the landscape matrix Larrivee, M., Buddle, C.M. (2011): Ballooning propensity of canopy at different spatial scales. Journal of Biogeography 35: 157–166. and understorey spiders in a mature temperate hardwood Rothenbücher, J., Schaefer, M. (2006): Submersion tolerance in forest. Ecological Entomology 36:144–151. floodplain arthropod communities. Basic and Applied Ecology Nentwig, W., Blick, T., Gloor, D., Hänggi, A., Kropf, C. (2015): 7: 398–408. Spiders of Europe.