Arachnologische Mitteilungen 38: 8-27 Nürnberg, Dezember 2009

Sascha BUCHHOLZ, University of Münster, Institute of Landscape Ecology, Department of Community Ecology, Robert-Koch-Str. 26, 48149 Münster, Germany E-Mail: saschabuchholz@uni-muenster.de

Martin KREUELS, AraDet, Alexander-Hammer-Weg 9, 48161 Münster, E-Mail: kreuels@aradet.de

eingereicht: 13.8.2009; akzeptiert: 25.11.2009; online verfügbar: 7.12.2009

Diversity and distribution of spiders (Arachnida: Araneae) in dry ecosystems of North Rhine-Westphalia (Germany)

Sascha Buchholz & Martin Kreuels

Abstract: The present study provides a robust data set for ecological planning and conservation of dry ecosys-tems in western Germany in general and North Rhine-Westphalia in particular. We summarised all available data from recent publications that dealt with spiders in dry ecosystems of North Rhine-Westphalia. Additionally, so far unpublished results of a detailed investigation regarding spiders in sand habitats of the Westphalian Bay that was conducted between 2006 and 2008 are presented. The analysis focussed on the habitat types according to Annex I of the EU Habitats Directive and related habitats. The investigation areas were scattered in the federal state of North Rhine-Westphalia. The data set comprised a total of 84436 individuals from 371 species and 28 families. Overall, an endangerment status is assigned to 68 species. Of these, 12 spiders are in imminent danger of becoming extinct. Two species, Erigonoplus globipes and Meioneta simplicitarsis, are believed to be extinct in North Rhine-Westphalia. Seven species (Dictyna major, Mastigusa arietina, Micaria formicaria, Styloctetor romanus, Thanatus striatus, Theridion uhligi and Xysticus ferrugineus) are new to the arachnofauna of North Rhine-Westphalia.

Keywords: biodiversity research, dry grassland, Flora-Fauna-Habitat directive, heathland, Juniperus communis heath, semi-dry grassland

In Germany, dry ecosystems, such as nutrient-poor sandy grasslands, dry heaths and semi-natural dry grasslands and scrubland facies on calcareous substra-tes are highly endangered (RIECKEN et al. 2006) and are listed in Annex I of the European Habitats Direc-tive as priority habitat types (BALZER & SSYMANK 2005). Due to increasing cultivation, especially during the past 50 years, and the lack of disturbance (drif-ting sand, grazing, fire) the area of dry ecosystems has decreased considerably in northern and western Germany (DRACHENFELS 1996, VERBÜCHELN & JÖBGES 2000, JENTSCH et al. 2002, KRATOCHWIL 2004, PARDEY 2004). Within the framework of conservation and ecological planning, updated and effective data sets concerning species inventories of endangered habitat types as well as distribution and ecology of habitat specialists are imperative. For example, stenotopic species are useful for the evaluation of the nature con-servation status of a habitat and biotic communities and may render profuse management guidelines (cf. SCHNITTER et al. 2003). Furthermore, diversity stu-dies and in particular diversity studies of arthropods

generally provide a wide spectrum of biogeographical and ecological probes for use in monitoring challen-ges (GARDNER 1991, KREMEN et al. 1993). In this context, spiders can play an important role since they are abundant, easy to record, occupy a wide array of spatial and temporal niches and respond immediately to habitat changes. Spiders provide robust data sets and statistical rigor within various kinds of ecological surveys (e.g. NEW 1999, SKERL 1999, SCHARFF et al. 2003, SCHMIDT et al. 2005, 2008, FINCH et al. 2007). Information about the diversity and distribution of spiders in dry ecosystems of western Germany is poor (KREUELS et al. 2008). Thus, the aim of this study is to present a first complete catalogue of spiders that occur in dry ecosystems of North Rhine-Westphalia. For this purpose, we have summarised all available data from the recent literature and added previously unreleased results of detailed investigations concer-ning the ecology of spiders in sand habitats of the Westphalian Bay.

Study areaThe investigation areas were scattered in the federal state of North Rhine-Westphalia that makes up part of the west and north-west of Germany (Fig. 1). The longest distance between areas was about 210 km (W-E) and 220 km (N-S). Most of the study areas were situated in the lowlands of North Rhine-West-phalia (Lower Rhine Valley, Westphalian Bay) with altitudes between 40 and 100 m a.s.l.. The climate

Spiders in dry ecosystems of North Rhine-Westphalia 9

Fig. 1: Location of the investigation areas in North Rhine-Westphalia Geographical explanations: 1 = Bockholter Berge (TK25 3912-1), 2 = Boltenmoor (3912-1), 3 = Dahlberg (4419-3), 4 = Die Spey (4606-3), 5 = Dorbaum (3912-3); 6 = Drover Heide (5205-3), 7 = Elter Sand (3711-3), 8 = Emsaue (3912-4), 9 = Heiliges Meer (3611-2), 10 = Hohenhorster Berge (4105-4), 11 = Holtwicker Wacholderheide (4208-2), 12 = Hülstener Wacholderheide (4208-2), 13 = Kaninchenberge (4306-3), 14 = Kla-tenberge (3912-4), 15 = Kooksheide (4013-2), 16 = Kregenberg (4519-3), 17 = Letter Wacholderheide (4109-1), 18 = Loosen Berge (4306-2), 19 = Moosheide (4118-1/3), 20 = Münster (4011-4), 21 = Osterklee (3712-4), 22 = Schirlheide (3913-3), 23 = Stolzenburg (5405-3), 24 = Talgraben (4014-1), 25 = Teverener Heide (5002-1/3), 26 = Venner Moor (4111-1), 27 = Vinnenberg (3913-4), 28 = Wacholderheide Hörsteloe (3907-1), 29 = Wahner Heide (5108/5109-1), 30 = Westruper Heide (4209-3), 31 = Wisseler Dünen (4203-2), 32 = Wulsenberg (4519/3).

is sub-Atlantic with a mean annual temperature of 9.5 to 10 °C and mean annual precipitation of 700 to 750 mm. Further study areas were located in the geographic region of the Eifel (Stolzenburg) and the Süder mountains (Hochsauerland: Dahlberg, Kre-genberg, Wulsenberg) at an elevation of 450 m a.s.l.

and about 345 m a.s.l., respectively. Both regions are characterised by mean annual temperatures below 7 °C and more than 1000 mm of annual precipitation. For further detailed information on the landscape and natural regions of North Rhine-Westphalia see DINTER (1999) and LÖBF (2005).

10 S. Buchholz & M. Kreuels

MethodsAll available data from recent publications that dealt with spiders in dry ecosystems of North Rhine-Westfalia were condensed into this study. Here, we analyse mainly the investigation sites representing the habitat types according to Annex I of the EU Habitats Directive (2310 - Dry sand heaths with Calluna and Genista; 2330 - Inland dunes with open Corynephorus and Agrostis grasslands; 4030 - European dry heaths; 5130 - Juniperus communis formations on heaths or calcareous grasslands; 6210 - Semi-natural dry grass-lands and scrubland facies on calcareous substrates) (cf. BALZER & SSYMANK 2005) and related habitats like semi-dry grasslands, dry Avenella-grasslands as well as ruderalised dry grasslands and heathlands. Investigation sites that could not be clearly assigned to one of the above listed habitat types were excluded from the analysis. Furthermore, so far unpublished results of a detailed investigation of spiders in sand habitats of the Westphalian Bay that was conducted between 2006 and 2008 are presented. A detailed overview of the investigated sites, study periods and methods is given in Tab. 1. Due to the differences in methodology, sampling intensity and investigation period, this study should only provide a qualitative description of the species inventories. All information on distribution and status of endangerment spiders of North Rhine-Westphalia were taken from KREUELS & BUCHHOLZ (2006) and KREUELS et al. (2008).

ResultsA total of 84436 individuals from 371 species and 28 families were summarised (Table 2). Altogether, for 67 species a status of endangerment is given. Apart from 22 species of category V (endangerment may be assumed), 22 endangered (category 3) and 9 highly endangered (category 2) species, 12 spiders are in imminent danger of becoming extinct (category 1). Furthermore, the linyphiid spiders Erigonoplus globipes and Meioneta simplicitarsis are now considered extinct in North Rhine-Westphalia (category 0). The records of seven species (Dictyna major, Mastigusa arietina, Micaria formicaria, Styloctetor romanus, Thanatus stri-atus, Theridion uhligi and Xysticus ferrugineus) new to the arachnofauna of North Rhine-Westphalia are noteworthy. Further faunistically interesting spe-cies discovered during this study were Agnyphantes expunctus, Diplocephalus connatus, Halorates reprosus, Hypselites jacksoni and Linyphia tenuipalpis, all of which are extremely rare in North Rhine-Westphalia and Germany.

DiscussionSeven species have been recorded for the first time for the arachnofauna of North Rhine-Westphalia: According to STAUDT (2009), the rare Dictyna major is mainly distributed in northern Germany, for exam-ple in dune habitats (HEYDEMANN 1964, SCHULTZ & PLAISIER 1995) and dry grasslands (BOCHMANN 1941, MERKENS 2002). This species also seems to inhabit dry Pinus forests (SCHAEFER 1980, SIMON 1995). Mastigusa arientina has so far only been recorded in southern and eastern Germany (STAUDT 2009). The biology and ecology of this rare species is rather unclear. According to MARTIN (1983) Mastigusa ari-entina is a myrmecophil spider that occurs exclusively in nests of Formica rufa ants. However, SIMON (1995) and KIELHORN & BLICK (2007) found this species at trees and on treetops, respectively. These records may explain the scarcity of this spider since pitfall traps seem to be an inappropriate method to study these strata. The gnaphosid Micaria formicaria is classified as a xerophilous species that inhabits mainly dry and semi-dry grasslands and Juniperus heaths (HAUK 1996, PLATEN et al. 1999) but also occurs in open pas-tures (HÄNGGI & BAUR 1998) and dry forest edges (BAUCHHENSS 2002). According to BAUCHHENSS (1995) and BAUR et al. (1996), Micaria formicaria inhabits sandy substrates as well as calcareous soils. This species is distributed mainly in the south-western regions of Germany (LEIST 1978, BAEHR & BAEHR 1984, HAUK 1996) but is also found in eastern Ger-many (PLATEN et al. 1999, STAUDT 2009). The linyphiid Styloctetor romanus was recorded mainly in southern and eastern Germany (STAUDT 2009) but also seems to be distributed in northern Germany (MERKENS 2002). According to MAR-TIN & UHLIG (1986), SACHER & BREINL (1999), PLATEN et al. (1999), RATSCHKER (2001), SACHER (2001) and SCHNITTER et al. (2003), this species is a typical inhabitant of dry grasslands or dry ecosystems in general (for example dry fallow land), respectively. In contrast, GÖTZE (1992) found a single individual in a salt marsh of northern Germany. On the other hand, up to now, Thanatus striatus has been found in a variety of different habitat types, such as dry and semi-dry grassland (HÖREGOTT 1958, HEYDEMANN et al. 1994, KUSCHKA 2004, AL HUSSEIN & LÜBKE-AL HUSSEIN 2007), heathlands (SCHMIDT & MELBER 2004), humid habitats (NENT-WIG 1983, PLATEN et al. 1999, STAUDT 2000), salt

Spiders in dry ecosystems of North Rhine-Westphalia 11

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84%

form

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4% fo

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3)27

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502

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100

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, E, G

4209

-351

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07.0

3''

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4'16

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74

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4% fo

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inco

ll. B

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ler D

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6'03

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06°1

7'57

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153

306

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2-09

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280

% e

than

ol (*

1)G

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(199

7) (*

3)

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g F

4519

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3"08

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59.7

2"34

54

504

.199

1-10

.199

64%

form

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uels

(199

8a)

Tab

. 1: O

verv

iew

of i

nve

stig

ated

sit

es, s

tud

y p

erio

ds

and

stu

dy

des

ign

s. (*

1) =

20

% g

lyce

rin

an

d 1

% t

hym

ol a

dd

ed. (

*2) =

acc

ord

ing

to

REN

NER

(198

2): e

than

ol +

ace

tic

acid

+ g

lyce

rin

+

wat

er. (

*3) =

hit

her

to u

np

ub

lish

ed d

ata.

Hab

itat

typ

es: A

= b

are

san

d, B

= d

ry s

and

hea

ths

wit

h C

allu

na a

nd

Gen

ista

(FFH

-co

de

2310

), C

= in

lan

d d

un

es w

ith

op

en C

oryn

epho

rus

and

Ag

rost

is g

rass

lan

ds

(FFH

-co

de

2330

), D

= E

uro

pea

n d

ry h

eath

s (F

FH-c

od

e 40

30),

E =

Juni

peru

s com

mun

is fo

rmat

ion

s o

n h

eath

s o

r ca

lcar

eou

s g

rass

lan

ds

(FFH

-co

de

5130

), F

= s

emi-

na-

tura

l dry

gra

ssla

nd

s an

d s

cru

bla

nd

faci

es o

n c

alca

reo

us

sub

stra

tes

(FFH

-co

de

6210

), G

= A

vene

lla d

om

inat

ed d

ry g

rass

lan

d, H

= s

emi-

dry

gra

ssla

nd

, I =

ru

der

alis

ed s

emi-

dry

an

d d

ry

gra

ssla

nd

.

12 S. Buchholz & M. Kreuels

meadows (SPARMBERG & SACHER 1997, BARNDT 2007, FINCH et al. 2007) and dunes (HEYDEMANN 1964, SCHULTZ 1992). T. striatus is distributed all over Germany (STAUDT 2009). MARTIN (1973a) described the theridiid spider Theridion uhligi. He found specimens during an investigation of the nature reserve Rietzer See (cf. MARTIN 1973b). Since then, this species has only been recorded in very few studies in dry grasslands of eastern Germany (PLATEN et al. 1999, JAKOBITZ 2003, STAUDT 2009). Conversely, T. uhligi was recorded in dry grasslands and heathlands in the Netherlands and Belgium ( JOCQUÉ 1977, KEER & VANUYTVEN 1993, PRINSEN 1996, HELSDINGEN 1999) so that the distribution gap is closed now. One further record refers to DUMA (2008) who found T. uhligi in a dry and sandy place of south-eastern Ro-mania. According to HERZOG (1968), BAUCHHENSS (1995), PERNER (1997), PLATEN et al. (1999) and SACHER (2002), the thomisid Xysticus ferrugineus is stenotopic of dry and calcareous grasslands. This species is known from only few locations in central and eastern Germany (STAUDT 2009).

The linyphiid spiders Erigonoplus globipes and Meio-neta simplicitarsis are now considered to be extinct in North Rhine-Westphalia: E. globipes was last recorded by KREUELS (1998b). Due to destruction of the former locations caused by land-use, this population has disappeared. Since then, this species has been considered to be extinct in North Rhine-Westphalia (KREUELS & BUCHHOLZ 2006). E. globipes is distributed mainly in higher altitudes of central and southern Germany (BRAUN 1960, BAEHR & BAEHR 1984, BAUCHHENSS & SCHOLL 1985, JOGER 1997) and seems to be absent in the lowlands (STAUDT 2009). According to all the previous records, this species is stenotopic for dry calcareous grasslands (BAEHR 1988, KÖHLER et al. 1989, PERNER 1997). Meioneta simplicitarsis is a rare species that has until now been found in eastern Germany (e.g. SACHER & BREINL 1999), Rhineland-Palatinate (WEBER 1999) and North Rhine-Westphalia (CASEMIR 1982). It seems to prefer dry grasslands (BRAUN 1969, BUCHAR & RŮŽIČKA 2002) but also occurs in wet meadows and pastures (HEIMER & NENTWIG 1991, KREUELS & BUCHHOLZ 2006). Agnyphantes expunctus, Diplocephalus connatus, Halorates reprosus, Hypselites jacksoni and Linyphia tenuipalpis are rarely distributed to North Rhine-Westphalia in particular and Germany in general

(KREUELS et al. 2008, STAUDT 2009). Agnyphantes expunctus seems to be restricted to humid habitats in low mountain ranges and mountains, e.g. in the Eifel (CASEMIR 1976, 1982), Harz (HEIMER 1980, SACHER 1997, 1998) and the Alps (KREUELS & LÜCKMANN 1998, MUSTER 2001). Diplocephalus connatus apparently prefers humid habitats as well (HÄNGGI et al. 1995, KREUELS & BUCHHOLZ 2006) and was hitherto sporadical recorded in western and central Germany (MORITZ 1973, ALBRECHT et al. 1994, ESSER 1997). Furthermore, Halorates reprosus was rarely found in the northern part of North Rhine-Westphalia (BUCHHOLZ & KREUELS 2005) and sin-gle locations along the Rhine (BRAUN 1960) and the North Sea coast (HELSDINGEN 2003, STAUDT 2009). Apart from southern Germany, Hypselites jacksoni was sampled in northern (SCHAEFER 1970, 1972), west-ern (CASEMIR 1960, 1976, RASKIN 2000) and eastern (HERZOG 1974, HIEBSCH 1985, OTTO et al. 2001) parts of the country. According to previous records, both, Halorates reprosus and Hypselites jacksoni appar-ently prefer humid habitats (cf. HÄNGGI et al. 1995). Finally, Linyphia tenuipalpis is mainly distributed to the lowlands (e.g. NW-Germany: MERKENS 2002, SCHMIDT & MELBER 2004; NE-Germany: PLATEN et al. 1999, SCHNITTER et al. 2003, BARNDT 2005; Netherlands: TUTELAERS 2000, 2001) with south-ernmost records in Thuringia (MALT & PERNER 2002). When working with the present data one has to consider several taxonomical problems. For example, Alopecosa accentuata and Alopecosa barbipes are closely related species that were once considered to be syno-nyms. However, several studies have confirmed the separation of both species (DAHLEM et al. 1987, CORDES & HELVERSEN 1990, CORDES 1995, VINK & MITCHELL 2002). On the other hand, at least the identification of female specimens is very difficult, while males can be clearly distinguished by the hair coat on Tibia I. Furthermore, both species show a different phenology and distribution (CORDES & HELVERSEN 1990, STAUDT 2009). Nevertheless, SCHMITT (2008) stated that several authors might have ignored the differences between both species found during previous studies, which makes the cur-rent status of A. accentuata and A. barbipes question-able. Consequently, we checked specimens from most of the lowland sites (2, 5, 7, 9, 10, 19, 21, 30) and one highland site (16). As a result of this, we state that all lowland records belong to Alopecosa barbipes while individuals from the low mountain ranges belong

Spiders in dry ecosystems of North Rhine-Westphalia 13

to A. accentuata. Thus, former data on A. accentuata that were published by BUCHHOLZ & HARTMANN (2008) and BUCHHOLZ (2008) have to be transferred to A. barbipes. Further taxonomic questions arise concerning Dicymbium nigrum brevisetosum which was first de-scribed by LOCKET (1962) and WIEHLE (1965) as a form and recognised as a species by LOCKET et al. (1974). Later, THALER (1986) discussed the existence of further forms of Dicymbium nigrum s. str. in the southern Alps. Finally, ROBERTS (1987) downgraded D. n. brevisetosum back to a form. This taxonomical problem is not completely solved yet, but differences in the hair coat of Tibia I which is considerably shorter in D. n. brevisetosum might justify the separation of both as valid species. Both have been recorded from Germany (WIEHLE 1965, HARMS 1987) but it is as-sumed that numerous records of Dicymbium nigrum s. str. might belong in fact to D. n. brevisetosum since, for example, drawings of the first one given in the refer-ence guide of NENTWIG et al. (2003) in truth refer to D. n. brevisetosum (cf. WIEHLE 1960, 1965). Hence, we checked all available material and exclusively found specimens with short hair coats on Tibia I which according to ROBERTS (1987) indicated the occur-rence of only Dicymium nigrum brevisetosum within the study area. Consequently, records for Dicymbium nigrum published by BUCHHOLZ & HARTMANN (2008) and BUCHHOLZ (2008) have to be adjusted to D. n. brevisetosum. Finally, we have to keep in mind the fact that several parts of North Rhine-Westphalia are hitherto poorly investigated. Especially for the eastern und southern parts of the federal state (e.g. Sauerland, Weserbergland), as well as for the mountains of the Eifel, faunistic records are almost entirely missing (KREUELS et al. 2008). This is a drawback for this study since large parts of the semi-natural dry grass-lands and scrubland facies on calcereous substrates are situated in these regions and thus remained under-sampled yet. As opposed to this, the coverage level for dry and sandy heathlands and grasslands has been thoroughly improved within detailed studies during the last years.

AcknowledgementsWe thank the district administration (Untere Land-schaftsbehörde) of Arnsberg, Borken, Coesfeld, Gütersloh, Paderborn, Recklinghausen, Steinfurt and Warendorf for enabling field work between 2006 and 2008. We also wish to express our gratitude to Andreas Beulting, Karsten Hannig, Kristian Mantel, Mathias Olthoff, Niels Ribbrock, Michael

Schwartze, Heinrich Terlutter and Christian Venne for information on study areas and Witold Arndt, Mareike Breuer, Nele Kloster and Tim-Martin Wertebach for their assistance during field work. Furthermore, we are very thankful to Theo Blick, Oliver-D. Finch, Volker Hartmann and two anonymous reviewers for valuable comments on an earlier draft of the manuscript, and to Robert Baumgartner for linguistic revision of the text. Sascha Buchholz received funding through a scholarship from the Friedrich-Ebert-Foundation (FES).

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18 S. Buchholz & M. KreuelsTa

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Ner

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mon

tana

*

1

23

Ner

iene

pel

tata

*

13

1216

Nus

oncu

s nas

utus

*

1

1O

edot

hora

x ag

rest

is 3

1

1O

edot

hora

x ap

icat

us*

1

13

43

331

319

27

61

31

428

442

1O

edot

hora

x fu

scus

*

213

172

11

21

11

50O

edot

hora

x gi

bbos

us*

2

2O

edot

hora

x re

tusu

s*

17

511

412

103

203

115

129

01

1599

Ost

eari

us m

elan

opyg

ius

*

11

2Pa

llidu

phan

tes e

rica

eus

*

12

101

115

Palli

duph

ante

s pal

lidus

*

12

11

21

112

122

112

3895

Pana

mom

ops i

ncon

spicu

us2

C2

373

192

567

Para

pele

cops

is ne

mor

alis

R

22

Pele

cops

is el

onga

ta

R

16

310

Pele

cops

is m

enge

iR

1

11

3Pe

leco

psis

para

llela

*

334

1055

134

8322

15

11

729

7282

313

181

7Pe

pono

cran

ium

ludi

crum

*C

11

Pepo

nocr

aniu

m o

rbicu

latu

m*

C9

3847

Poca

dicn

emis

junc

eaV

1

1Po

cadi

cnem

is pu

mila

*

21

52

1537

2564

911

123

21

11

151

17

203

7938

9Po

rrho

mm

a ca

mpb

elli

*

32

5Po

rrho

mm

a m

icroc

aven

seR

1

1Po

rrho

mm

a m

icrop

htha

lmum

*

36

21

19

152

443

Pri

neri

gone

vag

ans

*

22

4Sa

arist

oa a

bnor

mis

*

31

31

19

Saar

istoa

firm

aR

1

1Si

lom

etop

us b

ones

si1

12

2684

122

Silo

met

opus

eleg

ans

*C

11

Silo

met

opus

reus

si*

C27

27Si

ntul

a co

rnig

er*

C2

2St

emon

ypha

ntes

line

atus

*

25

711

15

16

17

42

67

65St

yloc

teto

r rom

anus

nr

3

3St

yloc

teto

r sta

tivu

s *

21

21Sy

edra

gra

cilis

R

15

6Ta

llusia

expe

rta

*

52

11

14

14

22 S. Buchholz & M. Kreuels

fam

iliy

[no

spec

ies]

/sp

ecie

s

End

Dis

stud

y ar

ea [n

o of

site

s]

su

m

1

23

45

67

89

1011

1213

1415

1617

1819

2021

2223

2425

2627

2829

3031

32

[3]

[2]

[2]

[4]

[5]

[5]

[5]

[1]

[4]

[1]

[1]

[1]

[4]

[1]

[1]

[4]

[1]

[3]

[5]

[7]

[1]

[1]

[1]

[1]

[5]

[1]

[1]

[2]

[4]

[7]

[3]

[4]

Ta

pino

cyba

inse

cta

*

123

327

Tapi

nocy

ba p

raec

oxV

9

6828

16

31

4810

575

145

300

Tapi

nocy

boid

es p

ygm

aeus

2

51

232

322

353

Tapi

nopa

long

iden

s*

1

22

5T

enui

phan

tes c

rist

atus

*

11

Ten

uiph

ante

s flav

ipes

*

24

85

21

12

73

742

Ten

uiph

ante

s men

gei

*

11

23

1910

12

11

81

26

159

Ten

uiph

ante

s ten

ebri

cola

*

12

11

5T

enui

phan

tes t

enui

s*

74

3013

420

343

1315

412

121

101

395

74

63

12

11

929

4614

912

05T

enui

phan

tes z

imm

erm

anni

*

33

11

11

10T

hyre

osth

eniu

s par

asit

icus

*C

11

Tiso

vag

ans

*

82

125

123

126

12

11

1532

21

132

53

381

Tri

chon

cus a

ffini

s1

C1

23

Tri

chon

cus s

axico

la1

C4

4T

rich

opte

rna

cito

2

102

896

112

313

253

Tri

chop

tern

a th

orel

li *

1

12

Tro

xoch

rus s

cabr

iculu

s*

2

881

11

1737

147

Typ

hoch

rest

us d

igita

tus

2

22

631

832

8W

alck

enae

ria

acum

inat

a*

7

11

526

12

522

31

11

82

3514

8W

alck

enae

ria

alti

ceps

*

21

23

16

116

106

48W

alck

enae

ria

anti

ca*

4

512

36

463

523

1053

170

Wal

cken

aeri

a at

roti

bial

is*

18

16

373

6751

13

630

22

61

2513

22

276

Wal

cken

aeri

a ca

pito

*

1

1W

alck

enae

ria

corn

icula

ns

*

11

163

221

89W

alck

enae

ria

cucu

llata

*

57

52

101

13

34W

alck

enae

ria

cusp

idat

a*

2

18

213

Wal

cken

aeri

a dy

sder

oide

s*

1

112

15

62

11

11

21

1921

75W

alck

enae

ria

furc

illat

a*

13

28

77

31

56

13

153

3510

9W

alck

enae

ria

mit

rata

*

1

15

7W

alck

enae

ria

mon

ocer

os*

1

244

24

1140

86W

alck

enae

ria

nodo

sa

*

11

Wal

cken

aeri

a nu

dipa

lpis

*

12

31

13

112

Wal

cken

aeri

a ob

tusa

*

32

163

33

12

336

Wal

cken

aeri

a un

icorn

is*

2

13

Wal

cken

aeri

a vi

gila

x*

1

11

25

Tet

ragn

athi

dae

[7]

M

etel

lina

men

gei

*

21

3M

etel

lina

segm

enta

ta*

1

12

Pach

ygna

tha

clerc

ki*

1

398

32

11

13

11

81

124

Pach

ygna

tha

dege

eri

*

6333

161

131

3818

1553

2816

47

921

594

45

132

192

879

4Pa

chyg

nath

a lis

teri

*

21

12

3137

Spiders in dry ecosystems of North Rhine-Westphalia 23

fam

iliy

[no

spec

ies]

/sp

ecie

s

End

Dis

stud

y ar

ea [n

o of

site

s]

su

m

1

23

45

67

89

1011

1213

1415

1617

1819

2021

2223

2425

2627

2829

3031

32

[3]

[2]

[2]

[4]

[5]

[5]

[5]

[1]

[4]

[1]

[1]

[1]

[4]

[1]

[1]

[4]

[1]

[3]

[5]

[7]

[1]

[1]

[1]

[1]

[5]

[1]

[1]

[2]

[4]

[7]

[3]

[4]

T

etra

gnat

ha m

onta

na*

1

1T

etra

gnat

ha p

inico

la*

1

1A

rane

idae

[12]

A

gale

nate

a re

dii

3

22

15

Ara

neus

dia

dem

atus

*

15

11

21

112

Ara

neus

qua

drat

us

*

11

2A

rani

ella

opi

stho

grap

ha*

1

12

Arg

iope

bru

enni

chi

*

11

2C

ercid

ia p

rom

inen

s*

1

223

61

41

38H

ypso

singa

alb

ovit

tata

3

138

22

1112

66H

ypso

singa

pyg

mae

a*

17

219

Hyp

sosin

ga sa

ngui

nea

*

16

281

541

Lar

inio

ides

corn

utus

*

11

2M

ango

ra a

caly

pha

*

112

11

15N

eosc

ona

adia

nta

*

11

Lyc

osid

ae [3

6]

Alo

peco

sa a

ccen

tuat

aV

C59

4626

131

Alo

peco

sa a

cule

ata

*

2323

Alo

peco

sa b

arbi

pes

?

4616

1027

72

23

16

129

81

663

Alo

peco

sa cu

neat

a*

5

253

798

442

121

121

260

413

011

273

252

11

181

7022

21A

lope

cosa

fabr

ilis

*

154

19A

lope

cosa

inqu

ilina

1C

22

Alo

peco

sa p

ulve

rule

nta

*

6429

175

308

1328

52

272

5058

811

76

1121

388

136

126

1351

103

3623

423

98A

lope

cosa

stri

atip

es3

C19

6584

Alo

peco

sa tr

abal

is3

1

146

531

149

9A

rcto

sa fi

gura

ta1

5

52

12A

rcto

sa le

opar

dus

*

113

245

43A

rcto

sa lu

teti

ana

*

301

221

162

112

920

2A

rcto

sa p

erita

*

731

5721

256

241

11

222

170

42

166

212

273

6A

ulon

ia a

lbim

ana

*

133

116

355

135

1670

3480

Hyg

roly

cosa

rub

rofa

scia

ta*

13

2336

Pard

osa

agre

stis

*

117

44

62

174

421

2Pa

rdos

a am

enta

ta*

81

91

715

212

21

32

113

6Pa

rdos

a ho

rten

sis3

28

36

289

Pard

osa

lugu

bris

s. st

r.*

39

417

410

149

117

125

3919

912

83

24

272

214

750

1010

1627

178

360

1876

Pard

osa

mon

tico

la*

1

333

4118

12

937

133

5168

287

69Pa

rdos

a ni

grice

psV

3

659

582

193

51

630

538

195

610

Pard

osa

palu

stri

s*

10

941

716

7522

4919

13

2350

221

173

8757

911

211

345

22

1738

36Pa

rdos

a pr

ativ

aga

*

86

1669

69

215

215

3419

11

203

Pard

osa

prox

ima

R

1818

Pard

osa

pulla

ta*

2

169

31

172

62

133

538

119

52

252

186

3517

597

233

127

085

1211

27

653

4080

24 S. Buchholz & M. Kreuels

fam

iliy

[no

spec

ies]

/sp

ecie

s

End

Dis

stud

y ar

ea [n

o of

site

s]

su

m

1

23

45

67

89

1011

1213

1415

1617

1819

2021

2223

2425

2627

2829

3031

32

[3]

[2]

[2]

[4]

[5]

[5]

[5]

[1]

[4]

[1]

[1]

[1]

[4]

[1]

[1]

[4]

[1]

[3]

[5]

[7]

[1]

[1]

[1]

[1]

[5]

[1]

[1]

[2]

[4]

[7]

[3]

[4]

Pa

rdos

a sa

ltans

*

33

Pira

ta h

ygro

philu

s*

34

161

121

2486

762

22

11

11

361

306

Pira

ta la

tita

ns*

1

21

11

6Pi

rata

pir

aticu

s*

1

11

12

21

9Pi

rata

tenu

itars

isR

2

2Pi

rata

ulig

inos

us*

3

39

3854

51

113

Tro

chos

a ru

rico

la*

2

2019

62

103

11

13

231

143

58

22

297

Tro

chos

a sp

inip

alpi

s *

1

1T

roch

osa

terr

icola

*

246

162

400

263

5917

19

453

116

199

196

2221

035

2051

213

915

3414

3012

9725

524

763

244

49X

erol

ycos

a m

inia

taV

23

633

821

912

54

1659

13

2063

311

312

185

319

67X

erol

ycos

a ne

mor

alis

*

320

72

33

135

116

6011

45

2633

2755

4P

isau

rida

e [2

]

Dol

omed

es fi

mbr

iatu

s*

1

1Pi

saur

a m

irab

ilis

*

74

66

208

11

11

16

11

11

28

101

87A

gele

nida

e [6

]

Age

lena

laby

rint

hica

*

72

113

4922

12

32

352

157

Hist

opon

a to

rpid

a*

1

412

62

21

2351

Mal

thon

ica

pict

a*

10

515

Mal

thon

ica

silve

stri

s*

3

13

12

199

110

Teg

enar

ia a

gres

tis

*

27

112

91

33

261

34

58

85T

egen

aria

atr

ica

*

13

81

13H

ahni

idae

[6]

A

ntist

ea el

egan

s*

2

19

25

19H

ahni

a he

lveo

la*

15

75

31

818

11

211

1610

711

4H

ahni

a m

onta

na*

9

57

134

41

52

14

174

Hah

nia

nava

*

154

378

41

126

21

69

269

986

Hah

nia

onon

idum

*

492

758

Hah

nia

pusil

la*

3

24

164

817

93

3014

223

8D

icty

nida

e [7

]

Alte

lla b

iunc

ata

1C

11

Arg

enna

subn

igra

1

53

8C

icuri

na ci

cur

*

721

1610

21

214

92

22

16

410

5128

6D

ictyn

a m

ajor

nr

1

1D

ictyn

a pu

silla

*

11

2L

athy

s hum

ilis

*

12

14

Mas

tigu

sa a

rien

tina

nr

11

Am

auro

biid

ae [3

]

Am

auro

bius

fene

stra

lis*

1

23

Coe

lote

s ter

rest

ris

*

135

626

16

148

106

Eur

ocoe

lote

s ine

rmis

*

5170

43

713

5

Spiders in dry ecosystems of North Rhine-Westphalia 25

fam

iliy

[no

spec

ies]

/sp

ecie

s

End

Dis

stud

y ar

ea [n

o of

site

s]

su

m

1

23

45

67

89

1011

1213

1415

1617

1819

2021

2223

2425

2627

2829

3031

32

[3]

[2]

[2]

[4]

[5]

[5]

[5]

[1]

[4]

[1]

[1]

[1]

[4]

[1]

[1]

[4]

[1]

[3]

[5]

[7]

[1]

[1]

[1]

[1]

[5]

[1]

[1]

[2]

[4]

[7]

[3]

[4]

T

itan

oeci

dae

[1]

T

itano

eca

quad

rigu

ttat

a3

C4

2428

Mit

urgi

dae

[1]

C

heir

acan

thiu

m er

rati

cum

*

21

21

17

Che

irac

anth

ium

onc

ogna

thum

R

11

Che

irac

anth

ium

vir

esce

ns3

3

42

61

35

61

21

12

37A

nyph

aeni

dae

[1]

A

nyph

aena

acc

entu

ata

*

31

4L

iocr

anid

ae [6

]

Agr

oeca

bru

nnea

*

4812

229

2220

118

81

24

51

21

929

61

221

Agr

oeca

cupr

ea3

1

541

1685

157

Agr

oeca

lusa

tica

1

22

22A

groe

ca p

roxi

ma

*

453

239

1315

32

162

61

226

316

2622

0A

post

enus

fusc

us*

C3

771

800

881

Scot

ina

cela

ns*

33

7594

202

Clu

bion

idae

[14]

0C

lubi

ona

brev

ipes

*

11

2C

lubi

ona

com

ta*

5

13

22

11

116

Clu

bion

a di

vers

a*

3

75

62

13

14

32C

lubi

ona

frut

etor

um3

1

23

Clu

bion

a lu

tesc

ens

*

31

4C

lubi

ona

negl

ecta

*

66

13

11

16

126

Clu

bion

a pa

llidu

la*

1

1C

lubi

ona

phra

gmit

is*

5

5C

lubi

ona

pseu

done

glec

ta

R

11

Clu

bion

a re

clusa

*

23

22

12

12C

lubi

ona

subs

ulta

ns

*

12

3C

lubi

ona

subt

ilis

*

12

3C

lubi

ona

terr

estr

is*

1

12

26

Clu

bion

a tr

ivia

lis*

3

11

5C

orin

nida

e [2

]

Phru

rolit

hus f

esti

vus

*

102

521

233

22

183

23

41

52

53

311

1016

430

2Ph

ruro

lithu

s min

imus

*

13

393

749

209

338

Zod

ariid

ae [1

]

Zod

ario

n ita

licum

1

99

Gna

phos

idae

[26]

C

allil

epis

noct

urna

*

35

17

16D

rasso

des c

upre

us*

1

837

910

91

11

135

347

Dra

ssode

s lap

idos

us*

3

13

255

41

39

315

99D

rasso

des p

ubes

cens

*

1260

391

36

33

114

32

32

141

9335

9D

rass

yllu

s lut

etia

nus

*

172

120

26 S. Buchholz & M. Kreuels

fam

iliy

[no

spec

ies]

/sp

ecie

s

End

Dis

stud

y ar

ea [n

o of

site

s]

su

m

1

23

45

67

89

1011

1213

1415

1617

1819

2021

2223

2425

2627

2829

3031

32

[3]

[2]

[2]

[4]

[5]

[5]

[5]

[1]

[4]

[1]

[1]

[1]

[4]

[1]

[1]

[4]

[1]

[3]

[5]

[7]

[1]

[1]

[1]

[1]

[5]

[1]

[1]

[2]

[4]

[7]

[3]

[4]

D

rass

yllu

s pra

eficu

sV

25

838

782

727

Dra

ssyl

lus p

usill

us*

12

5184

296

421

223

24

119

79

113

32

66

5914

29

127

6611

31D

rass

yllu

s vill

icus

1C

11

Hap

lodr

assu

s dal

mat

ensis

R

1313

Hap

lodr

assu

s sig

nife

r*

38

3783

149

422

281

203

12

164

84

1420

37

110

7012

381

2H

aplo

dras

sus s

ilves

tris

*

12

14

Hap

lodr

assu

s soe

rens

eni

R

41

5H

aplo

dras

sus u

mbr

atili

s*

3

212

51

163

13

26

6527

2M

icar

ia fo

rmic

aria

nr

2

2M

icar

ia fu

lgen

sV

25

141

122

108

24

817

6M

icar

ia g

uttu

lata

RC

77

Mic

aria

pul

icar

iaV

7

331

117

101

11

1410

21

99M

icar

ia si

lesia

ca3

1

109

20T

rach

yzel

otes

ped

estr

is3

4

11

6Z

elot

es a

eneu

sV

4

312

81

136

Zel

otes

apr

icoru

m*

1

11

3Z

elot

es el

ectu

sV

5

2541

1229

35

12

528

119

81

2999

842

0Z

elot

es la

trei

llei

*

2437

687

2812

3454

23

125

212

1439

173

526

Zel

otes

long

ipes

V

213

13

1921

793

13

3529

7Z

elot

es p

etre

nsis

*

2473

352

8581

211

622

120

55

2818

362

5910

445

1148

319

1321

Zel

otes

subt

erra

neus

*

422

14

317

111

16

32

12

11

44

103

209

Zor

idae

[3]

Z

ora

nem

oral

is*

C2

2Z

ora

silve

stri

s*

1

31

128

151

2676

Zor

a sp

inim

ana

*

131

21

2615

1316

702

114

22

25

113

32

147

232

286

Spar

assi

dae

[1]

M

icrom

mat

a vi

resc

ens

3

11

2P

hilo

drom

idae

[8]

Ph

ilodr

omus

aur

eolu

s*

2

2Ph

ilodr

omus

cesp

itum

*

21

21

11

227

210

01

383

Philo

drom

us co

llinu

s*

1

11

11

5Ph

ilodr

omus

disp

ar*

1

12

Philo

drom

us h

istri

o *

5

415

226

Philo

drom

us m

arga

rita

tus

R

11

Tha

natu

s for

mici

nus

1C

77

Tha

natu

s str

iatu

s nr

2

2T

hom

isid

ae [1

8]

Cor

iara

chne

dep

ressa

R

22

Dia

ea d

orsa

ta*

1

1O

zypt

ila a

tom

aria

*

2436

140

13

2823

2O

zypt

ila cl

avea

taV

C3

1022

202

1047

Spiders in dry ecosystems of North Rhine-Westphalia 27

fam

iliy

[no

spec

ies]

/sp

ecie

s

End

Dis

stud

y ar

ea [n

o of

site

s]

su

m

1

23

45

67

89

1011

1213

1415

1617

1819

2021

2223

2425

2627

2829

3031

32

[3]

[2]

[2]

[4]

[5]

[5]

[5]

[1]

[4]

[1]

[1]

[1]

[4]

[1]

[1]

[4]

[1]

[3]

[5]

[7]

[1]

[1]

[1]

[1]

[5]

[1]

[1]

[2]

[4]

[7]

[3]

[4]

O

zypt

ila p

rati

cola

*

25

42

21

117

Ozy

ptila

pul

lata

V

428

271

61

110

816

Ozy

ptila

scab

rico

la2

C14

317

Ozy

ptila

trux

V

3

25

Xys

ticu

s ace

rbus

3

13

151

20X

ysti

cus a

udax

*

32

12

8X

ysti

cus b

ifasc

iatu

s*

12

574

4111

4X

ysti

cus c

rist

atus

*

81

1867

336

191

310

26

120

3141

3616

12

220

443

2258

4X

ysti

cus e

rrat

icus

*

951

214

122

311

146

493

Xys

ticu

s fer

rugi

neus

nr

275

32X

ysti

cus k

empe

leni

*

2

2X

ysti

cus k

ochi

*

7440

3251

1735

110

326

5789

11

23

310

964

4X

ysti

cus r

obus

tus

2C

3634

70X

ysti

cus u

lmi

*

11

2Sa

ltic

idae

[20]

A

elur

illus

v-i

nsig

nitu

sV

12

8216

141

92

131

2117

1B

allu

s cha

lybe

ius

*

31

4E

uoph

rys f

ront

alis

*

135

52

518

65

101

11

804

58

21

11

26

513

231

9E

varc

ha a

rcua

ta3

1

23

Eva

rcha

falca

ta*

1

12

21

21

53

31

11

16

233

Eva

rcha

laet

abun

daR

4

72

13H

elio

phan

us a

urat

us3

1

1H

elio

phan

us cu

preu

s*

3

11

89

22H

elio

phan

us fl

avip

es*

7

11

11

15

13

122

Myr

mar

achn

e for

mic

aria

3

11

Neo

n re

ticu

latu

s*

2

11

11

14

127

39Pe

llene

s tri

punc

tatu

sV

16

251

1413

518

76

210

716

0Ph

legr

a fa

scia

taV

3

94

122

25

139

74

11

61

281

Pseu

deuo

phry

s lan

iger

a*

C1

1Sa

lticu

s sce

nicu

s*

C2

13

Sibi

anor

aur

ocin

ctus

s. s

tr.*

1

21

4Si

tticu

s pub

esce

ns*

1

34

Sitt

icus s

alta

tor

2

22

13

8Ta

lave

ra a

equi

pes

V

422

43

114

85

69

15

112

435

1Ta

lave

ra p

etre

nsis

V

11

14

11

143

35

61