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POLISH JOURNAL OF ECOLOGY<br />

(Pol. J. Ecol.)<br />

58 4 759–768 2010<br />

Regular research paper<br />

Maria OLESZCZUK 1 *, Malwina ULIKOWSKA 2 , Krzyszt<strong>of</strong> KUJAWA 1<br />

1<br />

Institute for Agricultural and Forest Environment, <strong>Polish</strong> Academy <strong>of</strong> Sciences, Field Station,<br />

Szkolna 4, 64–000 Turew, Poland, *e-mail: oleszczukm@vp.pl (corresponding author)<br />

2<br />

Rąbiń, Gajowa 1/2, 64–010 Krzywiń<br />

EFFECT OF DISTANCE FROM FOREST EDGE<br />

ON THE DISTRIBUTION AND DIVERSITY OF SPIDER WEBS<br />

IN ADJACENT MAIZE FIELD<br />

ABSTRACT: The study was carried out on<br />

lowland agricultural lands in western Poland between<br />

July and August 2007. The influence <strong>of</strong> the<br />

proximity <strong>of</strong> forested areas on the occurrence <strong>of</strong><br />

foliage-dwelling spiders was defined by measuring<br />

the abundance <strong>of</strong> spider webs on five transects<br />

situated at distances <strong>of</strong> 0, 20, 40, 60 and 80 metres<br />

from the forest edge. Generally, the greatest diversity<br />

and the highest total density were observed<br />

close to the forest edge, but an opposite relation<br />

was found for orb webs, built by spiders from the<br />

families Araneidae and Tetragnathidae, whose<br />

abundance was the lowest in close proximity to<br />

the forest. At a distance <strong>of</strong> 80 metres from the forest<br />

a slight increase in total abundance <strong>of</strong> webs<br />

was recorded, no doubt due to the presence <strong>of</strong><br />

weeds, in which were recorded numerous occurrences<br />

<strong>of</strong> sheet and irregular webs. On all the transects<br />

studied, orb webs predominated on maize<br />

shoots, whereas only three-dimensional sheet and<br />

irregular webs occurred on weeds. Included in<br />

this discussion are some <strong>of</strong> the potential effects <strong>of</strong><br />

the proximity <strong>of</strong> forest areas on foliage-dwelling<br />

spider populations in maize fields and on the possibilities<br />

for spiders limiting this crop’s pests.<br />

KEY WORDS: abundance <strong>of</strong> spiders, agricultural<br />

landscape structure, maize, cultivated<br />

fields, ecotonal zone<br />

1. INTRODUCTION<br />

For the last two decades much attention<br />

has been paid to the issue <strong>of</strong> biodiversity. One<br />

<strong>of</strong> the reasons for the increased interest in this<br />

topic is the search for some benefits for humans<br />

in connection with ecosystem services.<br />

It is commonly accepted that high species<br />

diversity contributes to the effectiveness <strong>of</strong><br />

biocenotic regulation in an ecosystem, which<br />

among other things may serve as a mechanism<br />

for limiting the numbers <strong>of</strong> some pest<br />

species on arable lands. Many studies have<br />

shown that the general level <strong>of</strong> biodiversity<br />

in farmland greatly depends on the structure<br />

<strong>of</strong> the landscape, mainly the presence<br />

<strong>of</strong> non-farmed elements fulfilling the role <strong>of</strong><br />

substitute habitats for many other species, including<br />

predators and parasites. Due to the<br />

presence <strong>of</strong> these habitats, species diversity<br />

in arable fields is much higher than on more<br />

uniform landscapes, where non-farmed habitats<br />

are very sparse. The mosaic-like structure<br />

<strong>of</strong> most farmland provides favourable<br />

conditions for many species <strong>of</strong> predatory or<br />

parasitic species, which can effectively reduce<br />

the populations <strong>of</strong> some pest species (Karg<br />

and Ryszkowski 1996, Karg 1999, 2004,<br />

Kajak 1990, 1998, Kajak and Oleszczuk<br />

journal 24b.indb 759 2011-01-14 21:37:24


760<br />

Maria Oleszczuk et al.<br />

2004, Thomas et al. 1991, Sunderland<br />

and Samu 2000, S chmidt et al. 2003, 2004,<br />

Thies et al. 2005, Flückiger and S chmidt<br />

2006). In consequence, recent studies on biodiversity<br />

have dealt with functional aspects,<br />

taking into account the role <strong>of</strong> a given species<br />

in the biocenosis, its trophic relations with<br />

other species and how these relations depend<br />

on habitat and landscape structure (Jeanneret<br />

et al. 2003, Birkh<strong>of</strong>er et al. 2007,<br />

Har wood and Obr ycki 2007, Öberg et al.<br />

2007, 2008, S chmidt et al. 2008, Richardson<br />

and Hanks 2009).<br />

One <strong>of</strong> the group <strong>of</strong> invertebrates that is<br />

<strong>of</strong>ten regarded as playing an important role<br />

in controlling the abundance <strong>of</strong> pest species<br />

in crop fields are spiders (Nyffeler 1982,<br />

1999, Rypstra 1995, Marc et al. 1999, Sunderland<br />

1999, Symondson et al. 2002,<br />

Maloney et al. 2003). Spiders catch their<br />

prey in two ways: by means <strong>of</strong> a web or by<br />

active hunting. Spider webs are very diverse<br />

in construction and size. The features <strong>of</strong> webs<br />

are strictly related to the taxonomy <strong>of</strong> spiders<br />

(Roberts 1995):<br />

• Linyphiidae build sheet-like webs<br />

without a retreat and the spider hangs<br />

beneath the web. The spider inserts<br />

a scaffolding <strong>of</strong> threads above the<br />

sheet<br />

• Araneidae and Tetragnathidae – spin<br />

orb webs, mostly almost vertical.<br />

These have developed as an efficient<br />

means <strong>of</strong> capturing flying insects.<br />

Their structure provides a unique<br />

combination <strong>of</strong> large capture area<br />

with close almost invisibility, making<br />

detection and avoidance difficult, especially<br />

at night.<br />

• Theridiidae – spin a tangled threedimensional<br />

web.<br />

The type <strong>of</strong> web construction is an evolutionarily<br />

adaptation towards the most effective<br />

way <strong>of</strong> catching a given set <strong>of</strong> insect<br />

species with given habitat preferences and<br />

type <strong>of</strong> flight (Nyffeler 1982, 1999, Sunderland<br />

et al. 1986, Har wood and Obr y-<br />

cki 2007, Ludy 2007). In consequence, the<br />

impact <strong>of</strong> spiders on a prey species depends<br />

on spider density and the web types present<br />

in a given habitat.<br />

The aim <strong>of</strong> our study was to determine<br />

the effect <strong>of</strong> the proximity <strong>of</strong> forest areas<br />

adjacent to fields planted with maize on the<br />

abundance <strong>of</strong> spider webs, the composition <strong>of</strong><br />

web construction types and the final effect on<br />

the diversity <strong>of</strong> prey species.<br />

2. STUDY AREA AND METHODS<br />

The study was carried out near the village<br />

<strong>of</strong> Turew, located in the central part <strong>of</strong><br />

the Dezydery Chłapowski Landscape Park<br />

(80 m a.s.l.), 30 kilometres south <strong>of</strong> Poznań<br />

(western Poland). The area is predominantly<br />

flat agricultural landscape, with gradients<br />

<strong>of</strong> less than 5%. The density <strong>of</strong> small rivers<br />

and canals amounts to 0.4 km km -2 and small<br />

bodies <strong>of</strong> water – 1.3 km -2 . Large, intensively<br />

farmed crop fields dominate the study area,<br />

but the landscape <strong>of</strong> the Park as a whole is<br />

characterized by a highly diversified structure,<br />

due to the presence <strong>of</strong> various nonfarmed<br />

habitats: woodland, copses, bodies <strong>of</strong><br />

water, wetlands, as well as a mosaic <strong>of</strong> smallsized<br />

crop fields. Crop fields and grasslands<br />

cover about 74% <strong>of</strong> the Park area, with woodland<br />

constituting about 15%. Woods are numerous,<br />

but highly fragmented. In most cases,<br />

small woods are composed <strong>of</strong> a mosaic <strong>of</strong><br />

small forest stands dominated by either Pinus<br />

sylvestris L. or Quercus robur L., sometimes<br />

with an admixture <strong>of</strong> Robinia pseudoacacia<br />

L., Picea abies L., Larix decidua Mill., Alnus<br />

glutinosa Gaertn., Betula pendula Roth and<br />

Fraxinus excelsior L. The undergrowth layer<br />

consists mainly <strong>of</strong> Padus serotina (Ehrh.)<br />

Borkh. The unique, mosaic structure <strong>of</strong> the<br />

farmland, rich in diverse non-farmed habitats,<br />

creates a favourable habitat for speciesrich<br />

communities <strong>of</strong> different organisms<br />

(Ryszkowski and Karg 1997).<br />

In the studied field, apart from maize<br />

plants, the following weeds were present:<br />

Echinochloa crus-galli (L.) P. Beauv., Chenopodium<br />

album L., Elymus (Agropyron) repens<br />

(L.) Gould, Stellaria media (L.) Vill. and Poa<br />

annua L. The two first mentioned plant species<br />

were the most numerous.<br />

The abundance <strong>of</strong> spider webs was determined<br />

twice: the first counting was performed<br />

from 18 th to 20 th <strong>of</strong> July and the second<br />

one – from 6 th to 9 th <strong>of</strong> August. The webs were<br />

counted on five transects <strong>of</strong> a maize field that<br />

were parallel to each other and to the edge <strong>of</strong><br />

a forest adjacent to this field. The tree stand at<br />

journal 24b.indb 760 2011-01-14 21:37:24


Effect <strong>of</strong> forest on crop field spider community 761<br />

Fig. 1. Scheme <strong>of</strong> sampling plots distribution.<br />

Fig. 2. Scheme <strong>of</strong> sampling plot. Grey color indicates the area where webs were counted.<br />

the edge <strong>of</strong> this forest consisted <strong>of</strong> ca 60-year<br />

old Robinia pseudoacacia with common elder<br />

and young robinia in the undergrowth layer.<br />

The distance between each <strong>of</strong> the transects<br />

amounted to 20 metres. Each transect consisted<br />

<strong>of</strong> 10 three metre long sampling plots<br />

(sections) distributed evenly, every 10 meters.<br />

The first transect was located in the row <strong>of</strong><br />

maize planted right at the forest edge and the<br />

others were located at distances <strong>of</strong> 20, 40, 60<br />

and 80 metres from the forest (Fig. 1).<br />

Each sampling plot consisted <strong>of</strong> a single<br />

row <strong>of</strong> maize (with stems and foliage up to 1.5<br />

metres in height) as well as a 0.7 metre wide<br />

strip <strong>of</strong> ground (with thinly growing weeds)<br />

between the two neighbouring rows <strong>of</strong> maize<br />

(Fig. 2).<br />

Spider webs were counted after spraying<br />

with water the whole <strong>of</strong> a sampling plot. This<br />

measure made webs visible even when they<br />

were <strong>of</strong> a very small size. The type <strong>of</strong> construction<br />

and location <strong>of</strong> the webs were used<br />

as features enabling their classification to one<br />

<strong>of</strong> the four spider families: Linyphiidae (sheet<br />

webs), Theridiidae (irregular webs) or Araneidae<br />

and Tetragnathidae (orb webs) (Roberts<br />

1995).<br />

3. RESULTS<br />

The abundance <strong>of</strong> spider webs was negatively<br />

correlated with distance from the forest<br />

(Fig. 3). The gamma correlation coefficient<br />

amounted to −0.67 (P< 0.001), but this<br />

journal 24b.indb 761 2011-01-14 21:37:24


762<br />

Maria Oleszczuk et al.<br />

Fig. 3. Total density <strong>of</strong> spider webs (arithmetical means and SD) in relation to distance from the forest<br />

(see Fig.1). Solid line illustrates relationship between distance (x) and density (y): y=8.87−1.09log(x)<br />

Table 1. Statistical significance <strong>of</strong> differences in total spider webs density between studied distances<br />

from the forest ( see Fig.1).<br />

Distance 20 40 60 80<br />

0 ns ** *** **<br />

20 ns ** ns<br />

40 ns ns<br />

60 ns<br />

**–P


Effect <strong>of</strong> forest on crop field spider community 763<br />

4. DISCUSSION<br />

4.1. Web abundance and the contribution<br />

<strong>of</strong> particular spider families<br />

The diversity and abundance <strong>of</strong> web-spinning<br />

spiders occurring in crop fields depends<br />

considerably on the type <strong>of</strong> crop, and thus on<br />

the structure (construction) <strong>of</strong> the cultivated<br />

plant species and the accompanying weeds.<br />

Web abundance in the maize field in our study<br />

(taking into account only those transects separated<br />

by a distance <strong>of</strong> over 20 metres from<br />

the forest in order to eliminate the influence<br />

<strong>of</strong> the surrounding environment) amounted<br />

to 5 webs per m 2 . Assuming that web abundance<br />

is closely related to total abundance <strong>of</strong><br />

web-spinning spiders, the figures recorded<br />

are similar to those obtained in the study carried<br />

out in Germany, where spider abundance<br />

in the maize field studied amounted to 5.9<br />

ind. m -2 and the majority <strong>of</strong> these were webspinning<br />

spiders (Ludy and L ang 2004).<br />

Nyffeler and Benz (1979) found a markedly<br />

lower abundance <strong>of</strong> foliage-dwelling spiders<br />

in similar crop: 0.1 ind. m -2 . In a soybean<br />

field over a similar period as that <strong>of</strong> the present<br />

study, i.e. in the middle <strong>of</strong> the vegetative<br />

season, spider web abundance amounted to 2<br />

webs per m 2 (Balfour and Rypstra 1998).<br />

In the same area where our study was conducted,<br />

foliage-dwelling spider abundance<br />

was previously studied in arable fields in July<br />

and August, and it was found that the majority<br />

<strong>of</strong> these were web-spinning species. The<br />

recorded abundance <strong>of</strong> web-spinning spiders<br />

ranged between 0.6 and 14.0, in alfalfa – 3.6<br />

and 3.0, in potato – 0.7 and 14.1 and in rye<br />

– 0.6 and 1.0 per m 2 (Łuczak 1975). The<br />

higher spider abundance (with exception <strong>of</strong><br />

potato) observed in the maize crop is probably<br />

most commonly caused by the lack <strong>of</strong><br />

other cereal crops in the fields in the period<br />

<strong>of</strong> the studies (July/August), because this is<br />

after the harvest time for these plants. Many<br />

<strong>of</strong> the spiders usually occurring on cereals<br />

may move to adjacent fields with crop plants,<br />

for example maize yet to be harvested, and for<br />

this reason spider abundance in these crops<br />

can be relatively high during this period.<br />

Fig 4. Density <strong>of</strong> Linyphiidae (A) and Theridiidae (B) spider webs (arithmetical means and 95% confidence<br />

level) in relation to distance from the forest (see Fig.1).<br />

Fig. 5. Relationship between web density (per m 2 ) <strong>of</strong> spiders from families Linyphiidae and Theridiidae.<br />

journal 24b.indb 763 2011-01-14 21:37:24


764<br />

Maria Oleszczuk et al.<br />

Fig 6. Combined density <strong>of</strong> Araneidae and Tetragnathidae spider webs (arithmetical means and 95%<br />

confidence level) in relation to distance from the forest (see Fig.1).<br />

Fig. 7. Density <strong>of</strong> spider’s web in relation to distance from forest (Fig.1) and place <strong>of</strong> their building.<br />

Orb webs, characteristic for Araneidae<br />

and Tetragnathidae, prevailed among the spider<br />

webs recorded in our study. Among the<br />

various types <strong>of</strong> webs recorded in maize fields<br />

in Switzerland (Nyffeler 1982) and the USA<br />

(Uetz et al. 1999), the highest contribution<br />

was also that <strong>of</strong> orb weaving spiders. Similar<br />

results were obtained in research carried out<br />

in a maize crop in an area located close to<br />

Turew village in an earlier period, when again<br />

spiders from the Araneidae and Tetragnathidae<br />

families were the most abundant. Three<br />

dominant foliage-dwelling species were recorded:<br />

Mangora acalypha Walck., Aculepeira<br />

ceropegia Walck. and Araniella cucurbitina<br />

Cl. The first two species mentioned are photophilous<br />

spiders (Nentwig et al. 2003), preferring<br />

areas with low plant cover and usually<br />

occurring in meadows but also in cultivated<br />

fields (Barthel and Platcher 1995, 1996;<br />

Wolak 2002, 2004, Ludy and L ang 2004,<br />

2006). Łuczak (1975) observed that in the<br />

fields located in the mosaic <strong>of</strong> habitats around<br />

the village <strong>of</strong> Turew, the species M. acalypha<br />

was 2.4 times more numerous than in<br />

landscape lacking <strong>of</strong> shelterbelts and forests.<br />

Thus, it seems that the presence in the agricultural<br />

landscape <strong>of</strong> semi natural habitats,<br />

such as belts <strong>of</strong> trees or shrubs, favours a high<br />

number <strong>of</strong> some orb web spider species.<br />

The significance <strong>of</strong> the type <strong>of</strong> vegetation<br />

adjoining crop fields has been confirmed by<br />

research carried out in Germany (Bavaria). In<br />

maize fields surrounded by margins covered<br />

in nettles, the species from the families spinning<br />

space webs (Theridiidae and Linyphiidae)<br />

predominated among the spiders spinning<br />

webs on plants. The spiders represented<br />

in lesser numbers were those catching their<br />

prey in orb webs – Tetragnathidae and Araneidae.<br />

However, the contribution <strong>of</strong> the<br />

family Araneidae was considerably higher on<br />

field margins (almost 20%, compared to 10%<br />

in maize fields), this being, in the authors’<br />

opinion, connected with the dense vegetation<br />

cover (Ludy and L ang 2004). In Hungary’s<br />

maize crops on the other hand, the predominant<br />

species among foliage-dwelling spiders<br />

was Phylloneta impressa L. K. – one <strong>of</strong> the<br />

most common species in Europe, spinning<br />

journal 24b.indb 764 2011-01-14 21:37:24


Effect <strong>of</strong> forest on crop field spider community 765<br />

irregular, three-dimensional webs. This is<br />

a photophilous species, which in Poland can<br />

be found in cereal fields, their abundance being<br />

higher on plants with greater branching<br />

<strong>of</strong> shoots, such as potatoes. The above mentioned<br />

studies as well as our own results reveal<br />

that maize fields are inhabited by fairly variable<br />

assemblages <strong>of</strong> spider species, in which<br />

different families may dominate, leading<br />

to different web types predominating from<br />

one site to another. The results <strong>of</strong> the present<br />

study also suggest that the type <strong>of</strong> vegetation<br />

covering the margins <strong>of</strong> these fields has<br />

an important influence on the composition <strong>of</strong><br />

the spider species occurring.<br />

It is worth adding that although this study<br />

was conducted over just a short period <strong>of</strong><br />

time, the results concerning the contribution<br />

<strong>of</strong> particular families can be treated as representative,<br />

because the peak figures for the<br />

spider families discussed occur in the same<br />

periods (Ludy and L ang 2004).<br />

4.2. The influence <strong>of</strong> distance from<br />

a forest on spider webs distribution<br />

The present study demonstrates a distinct<br />

connection between spider web types and<br />

abundance in maize fields and the proximity<br />

<strong>of</strong> forest. As the distance from the forest<br />

increased, total web abundance decreased,<br />

and among the three spider families studied,<br />

only spiders building orb webs demonstrated<br />

an opposite tendency, i.e. abundance <strong>of</strong> webs<br />

was the lowest right next to the forest edge<br />

(an average <strong>of</strong> less than one web per m 2 ), but<br />

already from a distance <strong>of</strong> 20 metres it was as<br />

high as at any greater distance. This is probably<br />

caused by the preference <strong>of</strong> that family’s<br />

species to well insulated biotopes. The only<br />

exception is Araniella cucurbitina, which is<br />

commonly observed in forests, so it is quite<br />

likely to occur in the ecotonal zone between<br />

forest and crop field. The webs distribution<br />

pattern in the Linyphiidae and Theridiidae<br />

families confirms the thesis that spiders<br />

do not spin webs on random sites, preferring<br />

areas where the abundance and diversity<br />

<strong>of</strong> potential prey species is high (Har wood<br />

and Obr ycki 2007). Ecotonal zones abound<br />

in diverse species <strong>of</strong> those insects that constitute<br />

the staple diet <strong>of</strong> spiders (Karg and<br />

Ryszkowski 1996, Karg 2004), and this<br />

partially explains the highest web abundance<br />

at the border <strong>of</strong> maize fields adjoining a forest<br />

and the reduced abundance inside the field,<br />

where insect numbers are generally lower<br />

(Karg 2007).<br />

The data collected on the transect situated<br />

at a distance <strong>of</strong> 80 meters from the forest<br />

indicate that local differentiation <strong>of</strong> vegetation<br />

cover also has an important influence on<br />

web abundance. On this transect, web abundance<br />

was higher than expected according to<br />

the observed relationship between total web<br />

abundance and distance from the forest. This<br />

inconsistency is due to the slightly higher<br />

than expected abundance <strong>of</strong> the sheet webs<br />

<strong>of</strong> Linyphiidae, this pattern being illustrated<br />

by the curve in Fig. 3. This can be explained<br />

by the occurrence <strong>of</strong> numerous weeds on<br />

some <strong>of</strong> the sections <strong>of</strong> this transect, covering<br />

a considerable part <strong>of</strong> the area and thus making<br />

suitable conditions for sheet webs. This<br />

observation also indicates a mechanism by<br />

which web abundance increases nearer forest.<br />

A belt several metres in width adjoining<br />

the forest is usually characterized by a greater<br />

amount <strong>of</strong> weeds, as was indeed observed in<br />

the study area. As a result, most <strong>of</strong> the sheet<br />

webs <strong>of</strong> linyphiid spiders were found just adjacent<br />

to the forest. So, for the species from<br />

this family (or at least some <strong>of</strong> them), proximity<br />

<strong>of</strong> a forest or the presence <strong>of</strong> the habitat<br />

itself is probably not as important as the fact<br />

that in the belt <strong>of</strong> the field adjoining the forest<br />

the abundance <strong>of</strong> weeds is usually higher than<br />

at any greater distance from the forest. It can<br />

therefore be concluded that for spider species<br />

from this family observed in the maize field,<br />

the limiting factor is a shortage <strong>of</strong> weeds as<br />

elements <strong>of</strong> suitable scaffolding.<br />

However, among the spider species<br />

studied, one group can be distinguished,<br />

for which distance from the forest is probably<br />

a key factor. These are species building<br />

their webs next to the ground – in hollows or<br />

near the surface. These are tiny spiders from<br />

the subfamily Erigonine (Linyphiidae). The<br />

abundance <strong>of</strong> these webs was the highest next<br />

to the forest, whereas in the transect located<br />

the farthest from the forest – despite the large<br />

numbers <strong>of</strong> weeds growing – web abundance<br />

was not different from that at a distances <strong>of</strong><br />

60 or 40 metres from the forest.<br />

journal 24b.indb 765 2011-01-14 21:37:25


766<br />

Maria Oleszczuk et al.<br />

4.3. The influence <strong>of</strong> forest proximity<br />

on the potential impact <strong>of</strong> spiders on<br />

insect abundance in cultivated fields<br />

Among foliage-dwelling spiders, each<br />

family and in some cases particular species<br />

build characteristic types <strong>of</strong> web. Moreover,<br />

even within the same family, webs may differ<br />

in vertical distribution. Particular web types<br />

are adapted to catch defined types <strong>of</strong> prey, for<br />

example orb webs mainly filter from the air<br />

flying insects such as Diptera and Homoptera<br />

(Nyffeler 1999, Ludy 2007). These two orders<br />

<strong>of</strong> insect include among others important<br />

maize pest species (Oscinella frit L., Elachiptera<br />

cornuta Fall., Delia platura Mig. and<br />

Aphidae), which can be successfully eliminated<br />

by being caught in orb webs and falling<br />

prey to spiders. Some spider species hunt<br />

much more diverse prey in terms <strong>of</strong> both their<br />

taxonomical and ecological features. For instance,<br />

the spider species Phylloneta impressa<br />

(Theridiidae) catches Aphidae and Diptera in<br />

its webs (Nyffeler 1982, Pekár 2000), but<br />

at the same time these webs also catch insects<br />

that jump and run among plants (Nyffeler<br />

1982). Insects from the Heteroptera order are<br />

<strong>of</strong>ten found in the irregular webs <strong>of</strong> theridiid<br />

spiders too (Árpás et al. 2005). So it may be<br />

assumed that more taxonomically diverse the<br />

spider commmunity caused the greater web<br />

diversity and this increases the possibilities<br />

for spiders contributing to the effective control<br />

<strong>of</strong> insect pest numbers. Since the diversity<br />

and abundance <strong>of</strong> webs are the highest<br />

just adjacent to a forest adjoining a field, the<br />

influence <strong>of</strong> the presence <strong>of</strong> forest on the potential<br />

reduction <strong>of</strong> insect numbers by spiders<br />

may be considerable.<br />

5. CONCLUSIONS<br />

1. Proximity <strong>of</strong> a forest positively affects<br />

spider web abundance and diversity<br />

in an adjoining crop field, which <strong>of</strong>fers<br />

possibilities for natural control<br />

<strong>of</strong> pest numbers by foliage-dwelling<br />

spiders.<br />

2. The highest predation potential <strong>of</strong><br />

spiders is localized in the crop fieldforest<br />

ecotonal zone due to the high<br />

abundance <strong>of</strong> webs and their diversity.<br />

3. The presence <strong>of</strong> weeds in a crop favours<br />

a high abundance <strong>of</strong> three-dimensional<br />

spider webs.<br />

4. The predominance <strong>of</strong> orb web numbers<br />

on maize shoots favours the<br />

capture <strong>of</strong> this crop’s pests, because<br />

a considerable proportion <strong>of</strong> these<br />

are precisely the tiny insects filtered<br />

by this type <strong>of</strong> web.<br />

ACKNOWLEDGMENTS: We acknowledge<br />

the help <strong>of</strong> Richard Bialy who corrected our<br />

English.<br />

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Received after revision February 2010<br />

journal 24b.indb 768 2011-01-14 21:37:25

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