See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/277121114

Environmental conditions affecting spiders in grasslands at the

lower reach of the River Tisza in Hungary

Article  in  Entomologica Fennica · June 2011

DOI: 10.33338/ef.4241

CITATIONS READS

20 485

3 authors, including:

Róbert Gallé
Centre for Ecological Research
78 PUBLICATIONS   1,148 CITATIONS   

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

INFOSTER - Invertebrates of Forest Steppe research View project

East-West Project View project

All content following this page was uploaded by Róbert Gallé on 21 December 2015.

The user has requested enhancement of the downloaded file.

© Entomologica Fennica. 31 May 2011

Environmental conditions affecting spiders in grasslands

at the lower reach of the River Tisza in Hungary
Róbert Gallé, Norbert Vesztergom & Tamás Somogyi

Gallé, R., Vesztergom, N. & Somogyi, T. 2011: Environmental conditions affect-

ing spiders in grasslands at the lower reach of the River Tisza in Hungary. —
Entomol. Fennica 22: 29–38.
The aims of the present study were to reveal the differences between grassland
types, and to identify the local and landscape parameters that influence the spider
assemblages at the reach of the River Tisza. The rarefied number of spider species
was negatively correlated with the proportion of forests in a radius of 500 m
around each site. A negative correlation was found between the number of grass-
land specialist species and the proportion of the forests, but the number of forest
species increased significantly with the neighboring forest area. The relative area
of neighboring forests, the number of plant species and regular flooding played
major roles in shaping of the species composition of spiders. The results of the
present study emphasize the importance of the effect of habitat landscape proper-
ties on spider assemblages. The structural diversity of the landscape may influ-
ence species richness and composition of the habitats.
R. Gallé, N. Vesztergom & T. Somogyi, Department of Ecology, University of
Szeged, Középfasor 52, H-6726 Szeged, Hungary; Corresponding author’s e-
mail: galle.robert@gmail.com
Received 15 February 2010, accepted 3 December 2010

1. Introduction habitats (Ward et al. 2002, Lambeets et al. 2008,

2009).
In the Hungarian Great Plain, grasslands are The distribution of organisms across such a
threatened by the expansion of agricultural activ- mosaic landscape is influenced by numerous fac-
ity (Horváth et al. 2009). During the river regula- tors and presumably by the complex relationships
tions in the 19th century the historical floodplain among these factors (Turner 2005).
was divided by dikes into floodplain and non- Among the habitat features, the composition
flooded, protected areas. The remnants of the pro- and structure of vegetation (Ysnel & Canard
tected grasslands are mainly used as pastures or 2000, Heikkinen & MacMahon 2004) influence
meadows. the species composition and diversity of the in-
These small patches are enclosed between vertebrate assemblages. Plant species richness
highly modified landscape elements, i.e. arable and the height of the vegetation is supposed to
fields and forest plantations of mainly non-native have a major influence on the spider assemblages,
tree species. The natural disturbance of the regu- as higher plant species richness offers more di-
lar flooding events and the related changes in the verse habitat structure and more potential sites for
vegetation structure are expected to affect struc- web building (Jeanneret et al. 2003a, Horváth et
tural and microclimatic parameters in floodplain al. 2009).
30 Gallé et al. • ENTOMOL. FENNICA Vol. 22

On the landscape scale, heterogeneity of the grasslands around the study sites, as the the-
surrounding habitat patches may influence the di- ory of island biogeography states that more
versity and the number of species of the assem- isolated habitats retain less habitat specialist
blages at a given patch (Duelli 1997). Although species than less isolated ones (e.g. Lövei et
numerous authors have demonstrated the signifi- al. 2006).
cance of the spatial structure of the landscape on (4) According to the above-mentioned hypothe-
invertebrate assemblages (e.g. Diekötter 2008, ses we expected both the proportion of forests
Ricketts 2001), it is difficult to generalize the ef- and grasslands to influence the species com-
fect of the landscape heterogeneity, as various position of spider assemblages.
taxa react differently to the habitat and landscape
features, since species ecology and dispersal abil-
ities are different for every organism (Burel & 2. Materials and methods
Baudry 1995, Jeanneret et al. 2003a).
The present study aimed at testing the effect 2.1. Study sites and sampling
of habitat parameters (soil humidity, regular
flooding, structure and cover of the vegetation) Two habitat complexes of the lower Tisza-valley
and landscape (proportion of grasslands, forests were selected for sampling spider assemblages.
and arable fields) on the spider assemblages, and The southern habitat complex (Vesszõs) is situ-
to reveal the differences between assemblages of ated near Szeged (46°17’30”N; 20°14’20”E),
the floodplain and on protected grasslands. Spi- where the landscape mainly consists of arable
ders were chosen for the study as they are the fields with small patches of grasslands, and oak
most diverse and abundant predatory invertebrate and poplar forest plantations are embedded in the
group of grasslands (Wise 1993). matrix of arable fields. The Dóc habitat complex
lies 40 km north of Szeged (46°26’20”N; 20°10’
The following hypotheses were tested: 40”E), in a structurally more complex landscape.
Higher proportion of semi-natural grasslands and
(1) In terms of the habitat features we expected forests occur here.
the number of species to increase with more The spider assemblages were studied at six
structured vegetation, as it is well-known that grasslands (sampling sites) in the southern habitat
vegetation height and density is important for complex and four in the northern habitat com-
spiders (e.g. Greenstone 1984). We did not plex, respectively. In each habitat complex, hay-
expect the number of species to change with meadow, saline grassland, floodplain meadow,
regular flooding event as floodplain habitats degraded grassland, and at the southern habitat
do not necessarily harbor more species but complex a pair of dike-side meadows (i.e. flood-
rather exhibit different species compositions plain and protected) were studied.
and different diversity patterns (Lambeets et To sample the fauna pitfall traps were applied
al. 2009). (diameter 65 mm, filled with ethylene glycol as
(2) According to Entling et al. (2007) and Lam- preservative, Koivula 2003, Schmidt et al. 2006).
beets et al. (2008, 2009) we also expected the At each site three lines of traps (sampling plots)
vegetation structure, soil moisture and regular were placed, each line consisted of five traps
flooding to alter the species composition of keeping a distance of 4 m between them. The av-
spiders. erage distance between the lines was 150 m. Prior
(3) As regards the effect of the landscape compo- to the data analysis, the data of the five traps were
sition, we expected the number of species to pooled. We expected an underestimation of the
increase with increasing proportion of forests abundance of web-building species, as pitfall
around the study sites, as a consequence of the traps measure the activity-density of species at
occurrence of forest specialist and generalist the ground level, thus they capture the spiders
species in the grasslands (Usher et al. 1993). with an active hunter lifestyle on the soil surface
We also expected the number of species to in- more efficiently (Topping & Sunderland 1992,
crease with the increasing proportion of Urák & Samu 2008). However, pitfall traps offer
ENTOMOL. FENNICA Vol. 22 • Spiders of river valley grasslands 31

a relatively good alternative for comparing the To assess the significance of their effect, the
activity-density of epigaeic spider fauna of differ- explanatory variables suggested by the URT me-
ent habitats (Scmidt et al. 2005, Öberg et al. thod were subjected to a linear mixed-effect
2007). The traps were kept open for four 2-week model with nested random effects using the li-
sampling periods (27 May–13 June, 30 June–15 brary nlme in R (Pinheiro et al. 2007). The effect
July, 16–30 Aug., 18 Sept.–01 Oct.) in 2007 to of the different habitat complexes and sampling
cover the main activity period of the spider spe- sites were used as random effects and the explan-
cies. The data of the four sampling periods were atory variables suggested by the regression tree
pooled. method were used as fixed effects in the mixed ef-
To characterize the structure of the vegeta- fect linear model (Crawley 2007). The collected
tion, the number of plant species and the total species were categorized according to their habi-
cover of the vegetation at the ground level, at 10 tat requirements (Buchar & Ruzicka 2002, Szita
and 40 cm above the ground, were measured et al. 2006, Schmidt & Hanggi 2007, Batáry et al.
within 1 × 1 meter quadrates near the traps, as 2008). The significant landscape scale parame-
structurally complex habitats may provide more ters were included in two subsequent linear mod-
potential sites for web building, and because spi- els to assess their effects on the number of forest
ders are strongly influenced by the habitat struc- and grassland specialist species.
ture (Schwab et al. 2002). Rényi’s diversity ordering was used to com-
To assess the features of the landscape, the pare the diversity of the floodplain and protected
proportion of land-use types was measured in a habitats (hypothesis 1). Diversity ordering pro-
radius of 500 m around each site, based on aerial vides the diversity profiles of the assemblages.
photographs (i.e. grasslands, forests and arable These curves consist of a series of diversity indi-
fields), as several studies suggest that landscape ces, as the scale parameter is changed, including
composition at scales of 500 m radius can be rele- indices sensitive to both rare and abundant spe-
vant for spiders (Clough et al. 2005, Schmidt et cies (Carranza et al. 2007). An assemblage is con-
al. 2005). The remaining landscape elements sidered to be more diverse than another if its pro-
(e.g. surface of the river) were not entered into the file lies above and they do not intersect (Tóth-
analysis in order to decrease the number of fac- mérész 1995). R software (R Development Core
tors which were tested. Team 2007) with the BiodiversityR Package
(Kindt 2008) was used for the calculations.
2.2. Data analysis The influence of the measured explanatory
variables was tested on spider assemblages (hy-
Habitat structure affects the trappability of spider pothesis 2 and 4) by using the multivariate regres-
individuals, which may result in biased data for sion tree (MRT) method. This method is an exten-
studies comparing species richness of habitats sion of univariate regression trees having multi-
with different structures (Melburne 1999). Due to variate response (De’ath 2002). MRT does not as-
the different habitat properties of the sampling sume particular form of relationship between spe-
plots and the large variation in the number of indi- cies abundances and explanatory variables. This
viduals recorded over the trapping period, rar- method can be used to identify interactions be-
efaction was used to standardise the number of tween explanatory variables (De’ath 2002,
species recorded within each of the sampling 2010).
plots (Heck et al. 1975, Gotelli & Colwell 2001). The size of the tree was selected based on the
As regards hypothesis 1 and 3, the univariate minimum tree size that falls below 1 SE of the
regression tree method (URT) was used to gain minimum cross-validation estimate. The final
insight into the structure of the data and identify tree size was selected based on the frequently oc-
the interactions between the variables (Tree curring number of leaves from 100 individual
Package, Ripley 2009). The method performs a trees (Work et al. 2004).
binary recursive partitioning of the dataset. It also The species that characterize each leaf and
offers the opportunity to identify the influential node of the MRT were identified using the indica-
explanatory variables (Crawley 2007). tor species index (IndVal) based on the relative
32 Gallé et al. • ENTOMOL. FENNICA Vol. 22

34, thus the species composition differed consid-

erably between the studied grasslands.

3.1. Hypothesis 1: The influence of habitat

features on the rarefied number of the
species

According to the univariate regression tree me-

thod the total vegetation cover is the only habitat
parameter that had a possible influence on the rar-
efied number of species in case of higher propor-
tion of forests (Fig. 1). However, the vegetation
cover had no significant effect on the rarefied
Fig. 1. Univariate regression tree for the rarefied spe-
cies richness. Node 1 is based on Forest (%), the pro-
number of the species according to the results of
portion of forests around the sampling plots; node 2 the linear mixed-effect model (t17 = –0.361; p =
on Veg. cov. (%), the total vegetation cover; node 3 on 0.722).
Arable field (%), the proportion of the arable field. The The diversity profiles of the spider assem-
different lengths of the lines following each split are blages inhabiting the floodplain and protected
proportional to the variance explained by the split. The habitats are given in Fig 2. The curve for the pro-
numbers below each leaf show the number of samp- tected grasslands runs above that for the
les falling into that leaf. floodplain grasslands and the two curves do not
intersect. Thus, the assemblages of the protected
abundance and relative frequency of occurrence grasslands were more diverse for the whole range
(Dufrene & Legendre 1997, De’ath 2002). of the scale parameter.

3.2. Hypothesis 2. The influence of habitat

3. Results features on the species composition
A total number of 3547 spider specimens were The cross validation of 100 MRT trees yielded
collected, representing 95 species in 17 families trees of two, four and five leaves, 11, 46 and 40
(Appendix, downloadable from http://ojs.tsv.fi/ times, respectively, with three trees resulting in
index.php/entomolfennica/). The species rich- 6–8 leaves. The four-leaf tree is presented in Fig
ness of grassland patches ranged between 23 and 3. The MRT, indicating the influence of the num-

Fig. 2. The results of

Rényi’s diverisity order-
ing. The diversity pro-
files do not intersect,
thus the assemblages
of the protected grass-
lands are more diverse.
ENTOMOL. FENNICA Vol. 22 • Spiders of river valley grasslands 33

Fig. 3. Multivariate regression tree for spiders. Bray-Curtis dissimilarity was used for splitting. The final tree size
was selected based on the most frequently occurring number of leaves from 100 individual trees. The numbers
below each leaf show the number of samples falling into that leaf. The abbreviated names of species with signifi-
cant indicator value are given at each leaf, the indicator values are given in parenthesis (*** p<0.001, ** p<0.01, *
p<0.05). Abbreviations: T are: Thanatus arenarius, H rad: Hogna radiata, M ross: Micaria rossica, G luc: Gna-
phosa lucifuga, Z seg: Zelotes segrex, Z lon: Z. longipes, D con: Diplostyla concolor, P ala: Pardosa alacris, Z lat:
Z. latreillei, O sim: Ozyptila simplex, O api: Oedothorax apicatus, A ele: Antistea elegans, A leo: Arctosa leopar-
dus, E den: Erigone dentipalpis, P lat: Pirata latitans, A. hum: Araeoncus humilis, D lut: Drassyllus lutetianus,
P ame: P. amentata, P cri: P. cribrata, P deg: Pachygnatha degeeri, P pro: Pardosa proxima, T str: T. striatus,
Z mun: Z. mundus.

ber of plant species (node 2, Fig. 3), accounted for For the fourth leaf we identified 14 indicator
12.9% of the total variance and the regular flood- species. Out of these spiders the generalist Oedo-
ing events (node 3, Fig. 3) accounted for 8.9% of thorax apicatus (Blackwall, 1850), Erigone den-
the total variance. tipalpis (Wider, 1834) and Pachygnatha degeeri
The IndVal analysis identified one indicator Sundevall, 1830 occur under the regularly dis-
species of the first leaf (Thanatus arenarius Tho- turbed conditions of agricultural fields and 11
rell, 1872). This species lives in open, dry habi- species are associated with wet meadows and
tats. pond margins (Buchar & Ruzicka 2002, Schmidt
For the second leaf the IndVal analysis identi- & Hanggi 2007).
fied 6 species. These spiders occur in the natural
alkali grasslands of the Great Hungarian Plain
(Szita et al. 2006, Batáry et al. 2008). 3.3. Hypothesis 3. The effect of landscape
For third leaf the IndVal analysis identified scale parameters on the rarefied number
two forest specialist species (Pardosa alacris of the species
(C.L. Koch, 1833) and Diplostyla concolor (Wi-
der, 1834)), and the generalist Zelotes latreillei According to the URT the two most influential
(Simon, 1878) and Ozyptila simplex (O.P.-Cam- parameters were the proportion of forests and ara-
bridge, 1862) (Buchar & Ruzicka 2002). ble fields (Fig. 1). As predicted, the proportion of
34 Gallé et al. • ENTOMOL. FENNICA Vol. 22

forests had a significant effect according to the water regime play an important role in the shap-
subsequent GLMM (t17 = –2.228, p = 0.039). ing of the diversity of spider assemblages at rive-
However, no significant relationship was found rine landscapes. Numerous studies have demon-
between the proportion of arable fields and the strated that spiders are particularly sensitive to
rarefied number of the species (t17 = –1.801, p = flooding. The spider diversity is not only influ-
0.089). The subsequent linear models revealed enced by the occurrence of flooding events but
the significant negative effect of forests on the also by the duration and frequency of these events
number of grassland specialist species (t19 = – (Uetz 1976, Paetzold et al. 2008).
2.166, p = 0.043) and the marginally significant
positive effect on the forest specialist species (t19
= 2.024, p = 0.057). 4.2. The number of plant species
and the regular flooding events influenced
the species composition of spiders
3.4. Hypothesis 4. The influence of landscape
scale parameters on the composition The vegetation and flooding events are con-
of species straints for spider occurrence, indicating the im-
portance of the quality of local habitats. In low-
The MRT analysis identified only one influential land floodplains, flooding dynamics determine
landscape scale parameter. The proportion of fo- the composition of the species and the structure of
rests in a radius of 500 m around each site (Fig. 3, the vegetation which in turn influence the compo-
node 1) accounted for 26.2% of the total variance. sition of spider fauna (Ballinger et al. 2005). As in
most habitat types, vegetation plays a major role
in determining the physical parameters of the
4. Discussion habitat, and therefore, has a significant influence
on the distributions of invertebrate species (Bonte
4.1. No influential habitat parameter on the 2002, Tews et al. 2004). Gravesen (2000) and
rarefied number of the species, but there is Lambeets et al. (2006) also support the hypothe-
a higher diversity at the protected grasslands sis that a combination of flooding intensity and
vegetation structure is important factor for the
Although numerous studies have demonstrated species composition of spiders in wet grassland
the effect of plant species richness and habitat areas.
structure on the number of species and the diver- Sabo et al. (2005) reviewed the literature on
sity of spiders (e.g. Jeanneret et al. 2003b, Tews et the species richness and assemblage composition
al. 2004), we found no significant relationship of flooded and non-flooded habitats. In accor-
between the vegetational parameters and the rar- dance with the results of the present study they
efied number of the species. Gallé & Torma found that the species composition differs be-
(2009) and Gallé et al. (2010) demonstrated that tween the regularly flooded and non-flooded hab-
different habitat types may exhibit spider assem- itats. This can increase the regional species rich-
blages with similar number of species but differ- ness. Several studies have confirmed the impor-
ent species compositions. Thus, for a detailed tance of fluvial dynamics affecting spider assem-
study into the relationship between the spiders blages by altering the habitat conditions (Bonn et
and the habitat parameters the examination of the al. 2002, Paetzold et al. 2008).
species abundance data is required in order to as- At regularly flooded habitats Lambeets et al.
sess the impact of these parameters on the spider (2009) found that the dispersal ability of species
assemblages without the loss of information plays a prominent role in structuring carabid and
when summarizing the data on the assemblages in spider assemblages, thus life-history traits affect
a single value such as species richness (Jeanneret species composition of spiders. Rothenbücher
et al. 2003b, c). and Schaefer (2006) studied the impact of flood-
In accordance with Bell et al. (1999) and ing on invertebrate assemblages. They distin-
Bonn et al. (2002), our results also indicate that guished two types of adaptation in invertebrates:
ENTOMOL. FENNICA Vol. 22 • Spiders of river valley grasslands 35

submersion tolerance and regular migration be- thus the isolation of the habitats. Higher propor-
fore and after the flooding. Most spiders over- tion of forests may serve as dispersal barriers for
winter at juvenile or adult stages (Pekár 1999). grassland specialist species.
These stages of terrestrial arthropods are less
flood-resistant than are their eggs, thus they are
not able to tolerate long winter submersions. 4.4. The fauna of the surrounding forests
Rothenbücher and Schaefer (2006) demonstrated influence the species composition of spiders
that the floodplain spider fauna consists of spe-
cies immigrating with receding water levels. The According to the MRT landscape composition is
abundance of cursorial spiders depends on the important in determining the species composition
proximity and connectivity of natural habitats, in grasslands. Similarly to our results, numerous
possibly for seasonal migration towards hiberna- studies have shown that landscape variables are
tion sites (Bonte et al. 2003, Lambeets et al. important determinants of species composition of
2008), while good ballooners are able to colonize spiders (e.g. Öberg et al. 2007, Batáry et al. 2008,
form greater distances (Bonte et al. 2002). Schmidt et al. 2008). The IndVal analyses identi-
Although the IndVal analysis identified 14 fied two forest specialist species (P. alacris and
species associated with floodplain grasslands, D. concolor) at non-flooded grasslands with a
many species of the floodplain habitats are oppor- high proportion of surrounding forests, and ac-
tunistic species from the surrounding non- cording to the linear models the number of forest
flooded habitats that can colonize after the flood- specialist species increased and the number of
ing events (Adis & Junk 2002). grassland specialist species decreased with the
proportion of surrounding forests. Although
Schmidt et al. (2005) found no significant corre-
4.3. The proportion of forests lation between the percentage of forests and spi-
had a significant negative effect der assemblages, Kajak (2007) demonstrated that
on the rarefied number of the species forests influence the species composition of ce-
real fields by enhancing the number and abun-
In accordance with the results of the present dance of forest specialist species. She also found
study, Clough et al. (2005) reported that the rar- significant effect of forest age and naturalness.
efied number of species is influenced by the land- Landscape scale forestry operations may af-
scape parameters. Öberg et al. (2007) and Drape- fect the diversity and species composition of the
la et al. (2008) found that a higher proportion of surrounding grassland habitats by altering the
forests in the surrounding landscape were associ- proportion or the habitat quality of forests. This
ated with a higher number of observed spider spe- emphasizes the importance of the landscape-level
cies, as forests may serve as source habitat for spi- approach to nature conservation.
ders. Our results also support this hypothesis as
we found increasing number of forest specialist
species with the proportion of surrounding fo- 5. Conclusions
rests. Species arriving in grasslands from forests
are likely to add new species to the spider assem- In the present study we have assessed the compo-
blage, as forests and open habitats are home to sitional differences of spider assemblages of vari-
contrasting spider assemblages (Entling et al. ous grassland types and the effects of habitat and
2007, Drapela et al. 2008). landscape properties on the rarefied species rich-
The effect of forests on the grassland special- ness and composition of spiders. Lowland flood-
ist species is possibly due to the fact that smaller, plain habitats are of high conservational value, as
more isolated fragments retain fewer habitat spe- their associated arthropod fauna consists of spe-
cies than larger less isolated ones (e.g. Lövei cialist and rare species (Rothenbücher & Schae-
2006). Several papers emphasize the effects of fer 2004). Even small fragments play an impor-
surrounding habitats on the dispersal of species tant role in maintaining the regional diversity of
(Bonte et al. 2004, Thorbek & Topping 2005) and spiders, as they are not entirely isolated, because
36 Gallé et al. • ENTOMOL. FENNICA Vol. 22

spiders are especially successful in dispersal fects of aerial dispersal, habitat specialisation, and
(Horváth et al. 2009). landscape structure on spider distribution across frag-
mented grey dunes. — Ecography 27: 343–349.
A total number of 95 species were collected, Buchar, J. & Ruzicka, V. 2002: Catalogue of Spiders of the
and the species richness of grassland patches Czech Republic. — Peres Publishers: Praha, Czech
ranged between 23 and 34, thus the species com- Republic. pp: 1–349.
position differed considerably between the stud- Burel, F. & Baudry, J. 1995: Species biodiversity in chang-
ied grasslands. Our results suggest that beside the ing agriculture landscapes: A case study in the Pays
d’Auge, France. — Agriculture Ecosystems and Envi-
conservation of floodplain grasslands, the main- ronment 55: 193–200.
tenance of diversity of grassland habitats is also Carranza, M. L., Acosta, A. & Ricotta, C. 2007: Analyzing
essential for preserving the regional species pool landscape diversity in time: The use of Rènyi’s genera-
of invertebrates. lized entropy function. — Ecological Indicators 7:
505–510.
Acknowledgements. We would like to thank László Clough, Y., Kruess, A. & Tscharntke, T. 2005: Spider di-
Körmöczi and Márta Zalatnai for their help with the botan- versity in cereal fields: comparing local, landscape and
ical surveys, Gábor Laskay for the linguistic correction of regional scales. — Journal of Biogeography 32: 2007–
the manuscript. We would also acknowledge Katja 2014.
Matveinen-Huju and an anonymous Referee for the con- Crawley, M. J. 2007: The R Book. — John Wiley: New
siderable efforts in suggesting improvements in the manu- York pp: 627–685.
script. The present study was supported by NKFP De’ath, G. 2002: Multivariate regression trees: a new tech-
6/013/2005 grant and the Regional University Knowledge nique for modeling species-environment relations-
Centre for Environmental and Nanotechnology, Univer- hips. — Ecology 83: 1105–1117.
sity of Szeged. De’ath, G. 2010: mvpart: Multivariate partitioning. R
package version 1.3-1. — [www document] URL
http://CRAN.R-project.org/package=mvpart (Site vi-
sited on 14 August 2010.)
References Diekötter, T., Billeterand, R. & Crist, T. O. 2008: Effects of
landscape connectivity on the spatial distribution of
Adis J. & Junk, W. J. 2002: Terrestrial invertebrates inhabi- insect diversity in agricultural mosaic landscapes. —
ting lowland river floodplains of Central Amazonia Basic and Applied Ecology 9: 298–307.
and Central Europe: a review. — Freshwater Biology Drapela, T., Moser, D., Zaller, J. G. & Frank, T. 2008: Spi-
47: 711–731. der assemblages in winter oilseed rape affected by
Ballinger, A., Mac Nelly, R. & Lake, P. S. 2005: Immediate landscape and site factors. — Ecography 31: 254–262.
and longer-term effects of managed flooding on flood- Duelli, P. 1997: Biodiversity evaluation in agricultural
plain invertebrate assemblages in south-eastern Au- landscapes: An approach at two different scales. —
stralia: generation and maintenance of a mosaic land- Agriculture Ecosystems and Environment 62: 81–91.
scape. — Freshwater Biology 50: 1190–1205. Dufrene, M. & Legendre, P. 1997: Species assemblages
Batáry, P., Kovács, A. & Báldi, A. 2008: Management ef- and indicator species: the need for a flexible asymme-
fects on carabid beetles and spiders in Central Hunga- trical approach. — Ecological Monographs 67: 345–
rian grasslands and cereal fields. — Community Eco- 366.
logy 9: 247–254. Entling, W., Schmidt, M. H., Bacher, S., Brandl, R. &
Bell, D., Petts, D. E. & Sadler, J. P. 1999: The distribution Nentwig, W. 2007: Niche properties of Central Euro-
of spiders in the wooded riparian zone of three rivers in pean spiders: shading, moisture and the evolution of
Western Europe. — Regulated Rivers: Research & the habitat niche. — Global Ecology and Biogeo-
Management 15: 140–158. graphy 16: 440–448.
Bonn, A., Hagen, K. & Reiche, D. W. 2002: The signifi- Gallé, R. & Torma, A. 2009: Epigeic spider (Araneae)
cance of flood regimes for carabid beetle and spider assemblages of natural forest edges in the Kiskunság
communities in riparian habitats – a comparison of (Hungary). — Community Ecology 10: 146–151.
three major rivers in Germany. — River Research and Gallé, R., Torma, A. & Körmöczi, L. 2010: Small-scale ef-
Applications 18: 43–64. fect of habitat heterogeneity on invertebrate assembla-
Bonte, D., Baert, L. & Maelfait, J.-P. 2002: Spider assem- ges in Hungarian sandy grasslands. — Polish Journal
blage structure and stability in a heterogeneous coastal of Ecology 58: 333–346.
dune system (Belgium). — Journal of Arachnology Gotelli, N. J. & Colwell, R. K. 2001: Quantifying biodiver-
30: 331–343. sity: procedures and pitfalls in the measurement and
Bonte, D., Lens, L., Maelfait, J.-P., Hoffmann, M. & Kuij- comparison of species richness. — Ecology Letters 4:
ken, E. 2003: Patch quality and connectivity influence 379–391.
spatial dynamics in a dune wolfspider. — Oecologia Gravesen, E. 2000: Spiders (Araneae) and other inverteb-
135:227–233. rate groups as ecological indicators in wetland area. —
Bonte, D., Baert, L., Lens, L. & Maelfait, J.-P. 2004: Ef- Ekologika (Bratislava) 19: 39–42.
ENTOMOL. FENNICA Vol. 22 • Spiders of river valley grasslands 37

Greenstone, M. H. 1984: Determinants of web spider spe- re on pitfall traps: an experimental evaluation. — Au-
cies diversity: vegetation structural diversity vs. prey stralian Journal of Ecology 24: 228–239.
availability. — Oecologia 62: 299–304. Öberg, S., Ekbom, B. & Bommarco, R. 2007: Influence of
Heck, K. L, van Belle, G. & Simberloff, D. 1975: Explicit habitat type and surrounding landscape on spider di-
calculation of the rarefaction diversity measurement versity in Swedish agroecosystems. — Agriculture,
and the determination of sufficient sample size. — Ecosystems and Environment 122: 211–219.
Ecology 52: 1459–1461. Paetzold, A., Yoshimura, C. & Tockner, K. 2008: Riparian
Heikkinen, M. W. & MacMahon, J. A. 2004: Assemblages arthropod responses to flow regulation and river chan-
of spiders on models of semi-arid shrubs. — The Jour- nelization. — Journal of Applied Ecology 45: 894–
nal of Arachnology 32: 313–323. 903.
Horváth, R., Magura, T., Szinetár, Cs. & Tóthmérész, B. Pearce, J. L. & Venier, L. A. 2006: The use of ground beet-
2009: Spiders are not less diverse in small and isolated les (Coleoptera: Carabidae) and spiders (Araneae) as
grasslands, but less diverse in overgrazed grasslands: bioindicators of sustainable forest management: A re-
A field study (East Hungary, Nyírség). — Agriculture, view. — Ecological Indicators 6: 780–793.
Ecosystems and Environment 130: 16–22. Pekár, S. 1999: Some observations on overwintering of
Jeanneret, Ph., Schüpbach, B., Pfiffner, L., Herzog, F. & spiders (Araneae) in two contrasting orchards in the
Walter, Th. 2003a: The Swiss agri-environmental pro- Czech Republic. — Agriculture, Ecosystems and En-
gramme and its effects on selected biodiversity indica- vironment 73: 205–210.
tors. — Journal for Nature Conservation 11: 213–220. Pinheiro, J., Bates, D., DebRoy, S. & Sarkar, D. 2007:
Jeanneret, Ph., Schüpbach, B., Pfiffner, L. & Walter, Th. nlme: Linear and nonlinear mixed effects models. R
2003b: Arthropod reaction to landscape and habitat fe- package version 3, 1–86. — [www document] URL
atures in agricultural landscapes. — Landscape Eco- http://ftp.uni-bayreuth.de/math/statlib/R/CRAN/doc/
logy 18: 253–263. packages/nlme.pdf (Site visited on 14 February 2010).
Jeanneret, Ph., Schüpbach, B. & Luka, H. 2003c: Quanti-
R Development Core Team. 2007: R: A Language and En-
fying the impact of landscape and habitat features on
vironment for Statistical Computing. R Foundation for
biodiversity in cultivated landscapes. — Agriculture,
Statistical Computing, Vienna. — [www document]
Ecosystems and Environment 98: 311–320.
URL http://www.R-project.org. (Site visited on 14
Kajak, A. 2007: Effects of forested strips on spider assem-
August 2010).
blages in adjacent cereal fields: dispersal activity of
Ricketts, T. H. 2001: The matrix matters: effective isola-
spiders. — Polish Journal of Ecology 55: 691–704.
tion in fragmented landscapes. — The American Na-
Kindt, R. 2008: The BiodiversityR Package. R package
turalist 158: 87–99.
ver. 1.2. — [www document] URL http://cran.r-pro-
ject.org/. (Site visited on 14 August 2010). Ripley, B. 2009: Tree: Classification and regression trees.
Koivula, M., Kotze, D. J., Hiisivuori, L. & Rita, H. 2003: R package version 1.0-27. — [www document] URL
Pitfall trap efficiency: do trap size, collecting fluid and http://CRAN.R-project.org/package=tree (Site visited
vegetation structure matter? — Entomologica Fennica on 14 August 2010).
14: 1–14. Rothenbücher, J. & Schaefer, M. 2006: Submersion tole-
Lambeets, K., Bonte, D., Van Looy, K., Hendrickx, F. & rance in floodplain arthropod communities. — Basic
Maelfait, J.-P. 2006: Synecology of spiders (Araneae) and Applied Ecology 7: 398–408.
of gravel banks and environmental constraints along a Sabo, J. L., Sponseller, R., Dixon, M., Gade, K., Harms, T.,
lowland river system, the Common Meuse (Belgium, Heffernan, J., Jani, A., Katz, G., Soykan, C., Watts, J.
the Netherlands) — In: Deltshev C & Stoev P (eds.), & Welter, J. 2005: Riparian zones increase regional
European Arachnology 2005 Acta Zoologica Bulgari- species richness by harboring different, not more, spe-
ca Suppl.1, pp: 137–149. cies. — Ecology 86: 56–62.
Lambeets, K., Hendrickx, F., Vanacker, S., Van Looy, K., Samu, F., Sunderland, K. D. & Szinetár, Cs. 1999: Scale-
Maelfait, J.-P. & Bonte, D. 2008: Assemblage structu- dependent dispersal and distribution patterns of spi-
re and conservation value of spiders and carabid beet- ders in agricultural systems: a review. — Journal of
les from restored lowland river banks. — Biodiversity Arachnology 27: 325–332.
and Conservation 17: 3133–3148. Schmidt, M. H. & Hänggi, A. 2007: Zelotes mundus (Ara-
Lambeets, K., Vandegehuchte, M. L., Maelfait, J.-P. & neae: Gnaphosidae) in the Camargue: a continental
Bonte, D. 2009: Integrating environmental conditions species reaches the western Mediterranean coast. —
and functional life history traits for riparian arthropod Bulletin of the British Arachnological Society 14: 27–
conservation planning. — Biological Conservation 29.
146: 625–637. Schmidt, M. H., Roschewitz, I., Thies, C. & Tscharntke, T.
Lövei, G. L., Magura, T., Tóthmérész, B., Ködölöcz, V. 2005: Differential effects of landscape and manage-
2006: The influence of matrix and edges on species ment on diversity and density of ground-dwelling
richness patterns of ground beetles (Coleoptera: Cara- farmland spiders. — Journal of Applied Ecology 42:
bidae) in habitat islands. — Global Ecology and. Bio- 281–287.
geography 15: 283–289. Schmidt, M. H., Clough, Y., Schulz, W., Westphalen, A.,
Melbourne, B. A. 1999: Bias in the effect of habitat structu- Tscharntke, T. 2006: Capture efficiency and preserva-
38 Gallé et al. • ENTOMOL. FENNICA Vol. 22

tion attributes of different fluids in pitfall traps. — of the science? — Annual Review of Ecology, Evolu-
Journal of Arachnology 34: 159–162. tion, and Systematics 36: 319–44.
Schmidt, M. H., Thies, C., Nentwig, W. & Tscharntke, T. Uetz, G. W. 1976: Gradient analysis of spider communities
2008: Contrasting responses of arable spiders to the in a streamside forest. — Oecologia 22: 373–385.
landscape matrix at different spatial scales. — Journal Urák, I. & Samu, F. 2008: Contribution to the spider fauna
of Biogeography 35: 157–166. of the Mohoº peat bog from Transylvania, with some
Schwab, A., Dubois, D., Fried, P. M & Edwards, P. J. 2002: new data for Romania. — North-Western Journal of
Estimating the biodiversity of hay meadows in north- Zoology 4: 50–60.
eastern Switzerland on the basis of vegetation structu- Usher, M., Field, J. P., Bedford, S. E., 1993: Biogeography
re. — Agriculture, Ecosystems and Environment 93: and diversity of ground-dwelling arthropods in farm
197–209. woodlands. — Biodiversity Letters 1: 54–62.
Szita, É., Samu. F., Szinetár, Cs., Dudás, Gy., Botos, E., Vasques, G. M., Grunwald, S. & Sickman, J. O. 2008:
Horváth, R. & Szalkovszki, O. 2006: New data on the Comparison of multivariate methods for inferential
occurence of Gnaphosa rufula (L. Koch, 1866) and modeling of soil carbon using visible/near-infrared
Gnaphosa mongolica Simon, 1895 in Hungary (Ara- spectra. — Geoderma: 146: 14–25.
neae: Gnaphosidae) — In: Deltschev, C. & Stoev, P. Ward, J. V., Tockner, K., Arscott, D. B. & Claret, C. 2002:
(eds.), European Arachnology 2005 pp. 329–334. Riverine landscape diversity. — Freshwater Biology
Tews, J., Brose, U., Grimm, V., Tielbörger, K., Wichmann, 47: 517–539.
M. C., Schwager, M. & Jeltsch, F. 2004: Animal spe- Wise, D. H. 1993: Spiders in ecological webs. — Cam-
cies diversity driven by habitat heterogeneity/diversi- bridge University Press. pp: 1–328.
ty: the importance of keystone structures. — Journal of With K. A, Pavuk, D. M., Worchuck, J. L, Oates, R. K. &
Biogeography 31: 79–92. Fisher, J. L. 2002: Threshold effects of landscape
Thorbek, P. & Topping, C. J. 2005: The inuence of landsca- structure on biological control in agroecosystems. —
pe diversity and heterogeneity on spacial dynamics of Ecological Applications 12: 52–65.
agrobiont linyphiid spiders: An individual based mo- Work, T. T., Shorthouse, D. P., Spence, J. R., Volney, W. J.
del. — BioControl, 50: 1–33. A. & Langor, D. 2004: Stand composition and structu-
Topping, C. J. & Sunderland, K. D. 1992: Limitations to re of the boreal mixedwood and epigaeic arthropods of
the use of pitfall traps in ecological studies exemplifi- the Ecosystem Management Emulating Natural Dis-
ed by a study of spiders in a field of winter wheat. — turbance (EMEND) landbase in northwestern Alberta.
Journal of Applied Ecology 29: 485–491. — Canadian Journal of Forest Research 34: 417–430.
Tóthmérész, B. 1995: Comparsion of different methods for Ysnel, F. & Canard, A. 2000: Spider biodiversity in con-
diversity ordering. — Journal of Vegetation Science 6: nection with the vegetation structure and the foliage
283–290. orientation of hedges. — Journal of Arachnology 28:
Turner, M. G. 2005: Landscape ecology: What is the state 107–114.

View publication stats