Upload
others
View
0
Download
0
Embed Size (px)
Citation preview
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: [email protected]
Martin KREUELS, AraDet, Alexander-Hammer-Weg 9, 48161 Münster, E-Mail: [email protected]
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 Additionally, so far unpublished results of a detailed investigation regarding spiders in sand habitats of the Westphalian Bay that was conducted between 2006 and 2008 are presented.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
noar
eaH
abita
t ty
pes
TK
25N
EM
as
lsi
tes
trap
s pe
r site
Inve
stig
atio
n pe
riod
Pres
erva
tion
fluid
refe
renc
e
1B
ockh
olte
r Ber
geC
3912
-152
°03'
30.2
1''
07°3
9'39
.05'
'50
34
08.2
006-
07.2
008
4% fo
rmal
inco
ll. B
uchh
olz
(*3)
2B
olte
nmoo
rC
3912
-152
°01'
18.5
8''
07°4
1'10
.12'
'55
24
08.2
006-
07.2
008
4% fo
rmal
inco
ll. B
uchh
olz
(*3)
3D
ahlb
erg
F44
19-3
51°3
0'10
.76"
08°5
4'19
.24"
275
25
04.1
991-
10.1
996
4% fo
rmal
inK
reue
ls (1
998a
)4
Die
Spe
y I
4606
-351
°20'
10.0
1"06
°42'
03.4
9"25
43
06.1
992-
09.1
992
80%
eth
anol
(*1)
Gri
go (1
997)
(*3)
5D
orba
um
B, C
, H39
12-3
52°0
1'23
.06"
07°4
2'48
.17"
505
504
.200
2-04
.200
33%
form
alin
Buc
hhol
z &
Har
tman
n (2
008)
6D
rove
r Hei
de
D52
05-3
50°4
3'58
.07"
06°3
1'41
.08"
205
51
05.2
006-
05.2
007
unkn
own
coll.
Kre
uels
(*3)
7E
lter S
and
A, B
, C, E
3711
-352
°13'
27.4
5''
07°3
2'02
.04'
'55
53
05.2
005-
10.2
005
4% fo
rmal
inB
uchh
olz
(200
8)8
Em
saue
C39
12-4
52°0
10'3
.87"
07°4
6'22
.35"
451
408
.200
6-07
.200
84%
form
alin
coll.
Buc
hhol
z (*
3)9
Hei
liges
Mee
rA
, B36
11-2
52°2
1'12
.11'
'07
°38'
02.9
1''
454
408
.200
6-07
.200
84%
form
alin
coll.
Buc
hhol
z (*
3)10
Hoh
enho
rste
r Ber
geC
4105
-451
°49'
55.5
2"06
°39'
17.9
8"30
14
08.2
006-
11.2
006
4% fo
rmal
inco
ll. B
uchh
olz
(*3)
11H
oltw
icke
r Wac
hold
erhe
ide
E42
08-2
51°4
5'00
.04"
07°0
7'36
.68"
901
408
.200
6-07
.200
84%
form
alin
coll.
Buc
hhol
z (*
3)12
Hül
sten
er W
acho
lder
heid
eE
4208
-251
°47'
52.0
3''
07°0
6'30
.24"
801
408
.200
6-07
.200
84%
form
alin
coll.
Buc
hhol
z (*
3)13
Kan
inch
enbe
rge
A, B
4306
-351
°37'
39.1
3"06
°41'
46.5
2"25
43
06.1
992-
09.1
992
80%
eth
anol
(*1)
Gri
go (1
997)
(*3)
14K
late
nber
geC
3912
-452
°00'
16.7
3''
07°4
7'02
.70'
'60
14
08.2
006-
07.2
008
4% fo
rmal
inco
ll. B
uchh
olz
(*3)
15K
ooks
heid
e A
4013
-251
°58'
13.9
9"07
°59'
03.2
8"60
15
05.1
992-
08.1
992
unkn
own
Kre
uels
et a
l (20
08)
16K
rege
nber
g F
4519
-351
°26'
25.6
4"08
°51'
51.4
5"33
04
504
.199
1-10
.199
64%
form
alin
Kre
uels
(199
8a)
17L
ette
r Wac
hold
erhe
ide
B, C
4109
-151
°53'
12.0
9''
07°1
0'05
.81'
'75
14
08.2
006-
07.2
008
4% fo
rmal
inco
ll. B
uchh
olz
(*3)
18L
oose
n B
erge
A
, I43
06-2
51°4
0'07
.82"
06°4
6'40
.70"
353
306
.199
2-09
.199
280
% e
than
ol (*
1)G
rigo
(199
7) (*
3)19
Moo
shei
deA
, B, C
, H41
18-1
/351
°51'
18.7
1''
08°4
0'58
.69'
'13
05
408
.200
6-07
.200
84%
form
alin
coll.
Buc
hhol
z (*
3)20
Mün
ster
A, C
, I40
11-4
51°5
5'59
.75"
07°3
9'48
.13"
607
304
.200
0-07
.200
0br
ine
Kre
uels
et a
l (20
08)
21O
ster
klee
F37
12-4
52°1
3'17
.86"
07°4
5'38
.81"
110
14
04.2
008-
07.2
008
4% fo
rmal
inco
ll. B
uchh
olz
(*3)
22Sc
hirl
heid
eD
3913
-352
°00'
37.3
0"07
°51'
14.5
8"60
15
06.2
008-
10.2
008
Ren
ner (
*2)
coll.
Kre
uels
(*3)
23St
olze
nbur
g F
5405
-350
°30'
55.8
2"06
°34'
00.5
7"45
01
1004
.197
1-09
.197
14%
form
alin
Bec
ker (
1977
)24
Tal
grab
en
A40
14-1
51°5
7'15
.53"
08°0
1'36
.96"
501
505
.199
2-08
.199
2un
know
nK
reue
ls e
t al (
2008
)25
Tev
eren
er H
eide
A
, C, D
5002
-1/3
50°5
7'35
.05"
06°0
4'08
.22"
905
520
045%
ace
tic a
cid
Kre
uels
(200
6)26
Ven
ner M
oor
D41
11-1
51°5
1'51
.16"
07°3
2'14
.25"
651
405
.200
7-10
.200
74%
form
alin
coll.
Buc
hhol
z (*
3)27
Vin
nenb
erg
I39
13-4
52°0
1'33
.96"
07°5
8'01
.35"
551
505
.199
2-08
.199
2un
know
nK
reue
ls e
t al (
2008
)28
Wac
hold
erhe
ide
Hör
stel
oeC
, E39
07-1
52°0
5'45
.50'
'06
°54'
43.2
0''
502
408
.200
6-07
.200
84%
form
alin
coll.
Buc
hhol
z (*
3)29
Wah
ner H
eide
A
, C, D
5108
/510
9-1
50°5
2'19
.48"
07°1
0'06
.46"
100
45
01.1
994-
10.1
994
70%
eth
anol
Jäge
r (19
96)
30W
estr
uper
Hei
deB
, E, G
4209
-351
°44'
07.0
3''
07°1
4'16
.47'
'45
74
08.2
006-
07.2
008
4% fo
rmal
inco
ll. B
uchh
olz
(*3)
31W
isse
ler D
ünen
A
, C42
03-2
51°4
6'03
.44"
06°1
7'57
.39"
153
306
.199
2-09
.199
280
% e
than
ol (*
1)G
rigo
(199
7) (*
3)
32W
ulse
nber
g F
4519
-351
°26'
44.1
3"08
°51'
59.7
2"34
54
504
.199
1-10
.199
64%
form
alin
Kre
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).
ReferencesALBRECHT C., T. ESSER & J. WEGLAU (1994): Un-
tersuchungen zur Wiederbesiedlung unterschiedlich strukturierter Feldraine durch ausgewählte Arthropo-dengruppen (Araneae, Isopoda, Carabidae, Heteroptera, Lepidoptera und Saltatoria) im landwirtschaftlichen Rekultivierungsgebiet des Braunkohletagebaus `Zu-kunft-West` bei Jülich. – Entomologische Mitteilungen Löbbecke-Museum Aquazoo Düsseldorf 7: 1-222
AL HUSSEIN I.A. & M. LÜBKE-AL HUSSEIN (2007): Faunistisch-Ökologische Erhebungen zu Webspinnen (Arachnida: Araneae) und Laufkäfern (Coleoptera: Carabidae) im ehemaligen Braunkohlentagebau Nach-terstedt. – Entomologische Mitteilungen Sachsen-An-halt 15: 43-45
BAEHR B. (1988): Die Bedeutung der Araneae für die Na-turschutzpraxis, dargestellt am Beispiel von Erhebungen im Landkreis Weißenburg-Gunzenhausen (Mittelfran-ken). – Schriftenreihe des Bayerischen Landesamt für Umweltschutz 83: 43-59
BAEHR B. & M. BAEHR (1984): Die Spinnen des Lau-tertales bei Münsingen (Arachnida, Araneae). – Veröf-fentlichungen für Naturschutz und Landschaftspflege in Baden-Württemberg 57/58: 375-406
BALZER S. & A. SSYMANK (2005): Natura 2000 in Deutschland. Naturschutz und biologische Vielfalt 14 [CD Rom]
BARNDT D. (2005): Beitrag zur Arthropodenfauna des Na-turparks Schlaubetal und Umgebung. - Faunenanalyse und Bewertung (Coleoptera, Heteroptera, Saltatoria, Araneae, Opiliones u.a.). – Märkische Entomologische Nachrichten 7 (2): 45-102
BARNDT D. (2007): Beitrag zur Arthropodenfauna der Binnensalzwiesen von Storkow und Philadelphia (Brandenburg/Landkreis Oder-Spree) - Faunenanalyse und Bewertung - (Coleoptera, Heteroptera, Auchenor-rhyncha, Saltatoria, Araneae, Isopoda u. a.). – Märkische Entomologische Nachrichten 9 (1): 1-54
BAUCHHENSS E. (1995): Die epigäische Spinnenfauna auf Sandflächen Nordbayerns (Arachnida: Araneae). – Zoologische Beiträge N.F. 36: 221-250
BAUCHHENSS E. (2002): Die Spinnenfauna eines thermo-philen Waldrandes in Mittelfranken (Bayern). – Arach-nologische Mitteillungen 23: 1-21
14 S. Buchholz & M. Kreuels
BAUCHHENSS E. & G. SCHOLL (1985): Bodenspinnen einer Weinbergsbrache im Maintal (Steinbach, Lkr. Haßberge). Ein Beitrag zur Spinnenfaunistik Unter-frankens. – Abhandlungen des Naturwissenschaftlichen Vereins Würzburg 23/24 (1982/83): 3-23
BAUR B., J. JOSHI, B. SCHMID, A. HÄNGGI, D. BORCARD, J. STARY, A. PEDROLI-CHRISTEN, G.H. THOMMEN, H. LUKA, H.-P. RUSTERHOLZ, P. OGGIER, S. LEDER-GERBER & A. ERHARDT (1996): Variation in species richness of plants and diverse groups of invertebrates in three calcareous grasslands of the Swiss Jura mountains. – Revue suisse de zoologie 103: 801-833
BECKER J. (1977): Die Trockenrasenfauna des Natur-schutzgebietes Stolzenburg (Nordeifel). – Decheniana 130: 101-113
BOCHMANN G. VON (1941): Die Spinnenfauna der Strandhaferdünen an den deutschen Küsten. – Kieler Meeresforschung 4: 38-69
BRAUN R. (1960): Neues zur Spinnenfauna des Rhein-Main-Gebietes und der Rheinpfalz. – Jahrbücher des Nassauischen Vereins für Naturkunde 95: 28-89
BRAUN R. (1969): Zur Autökologie und Phänologie der Spinnen (Araneida) des Naturschutzgebietes „Mainzer Sand“. – Mainzer naturwissenschaftliches Archiv 8: 193-288
BUCHAR J. & V. RŮŽIČKA (2002): Catalogue of spiders of the Czech Republic. Peres Publisher, Praha. 349 pp.
BUCHHOLZ S. (2008): Spider assemblages in an inland dune complex of Northwest Germany. – Drosera 2008: 63-76
BUCHHOLZ S. & V. HARTMANN (2008): Spider fauna of semi-dry grasslands on a military training base in Northwest Germany (Münster). – Arachnologische Mitteilungen 35: 51-60
BUCHHOLZ S. & M. KREUELS (2005): Die Spinnen (Arachnida: Araneae) im Naturschutzgebiet "Heiliges Meer" - eine vorläufige Artenliste. – Natur und Heimat 65 (4): 97-112
CASEMIR H. (1960): Beitrag zur Kenntnis der Niederrhei-nischen Spinnenfauna. – Decheniana 113: 239-264
CASEMIR H. (1976): Beitrag zur Hochmoor-Spinnenfauna des Hohen Venns (Hautes Fagnes) zwischen Nordeifel und Ardennen. – Decheniana 129: 38-72
CASEMIR H. (1982): Zweiter Beitrag zur Spinnenfauna des Bausenberges (Brohltal, östl. Vulkaneifel). – Decheniana 27: 47-55
CORDES D. (1995): Alopecosa accentuata and A. barbipes (Araneae, Lycosidae) in Central and Northern Europe. – Proceedings of the 15th European Colloquium of Arachnology: 213
CORDES D. & O. V. HELVERSEN (1990): Indications for the existence of Alopecosa barbipes (Sundevall 1832) as a sibling species to Alopecosa accentuata (Latreille 1817). Results of morphological, ethological and biogeogra-
phical studies. – Bulletin de la Société Européenne d’Arachnologie (H.S.) 1: 70-74
DAHLEM B., C. GACK & J. MARTENS (1987): Balzverhal-ten von Wolfspinnen der Gattung Alopecosa (Arachnida: Lycosidae). – Zoologische Beiträge N. F. 31: 151-164
DINTER W. (1999): Naturräumliche Gliederung. In: Lan-desanstalt für Ökologie, Bodenordnung und Forsten NRW (ed.): Rote Liste der gefährdeten Pflanzen und Tiere in Nordrhein-Westfalen, 3. Fassung. – LÖBF-Schriftenreihe 17: 29-36
DRACHENFELS O. V. (1996): Rote Liste der gefährdeten Biotoptypen in Niedersachsen – Bestandsentwicklung und Gefährdungsursachen der Biotop- und Ökosys-temtypen sowie ihrer Komplexe – Stand Januar 1996. – Naturschutz und Landschaftspflege in Niedersachsen 34: 1-148
DUMA I. (2008): Theridion uhligi Martin, 1974 (Araneae: Theridiidae) new to Romania. – Entomologica Roma-nica 13: 297-299
ESSER T. (1997): Artenvielfalt in der modernen Agrar-landschaft: Der Feldrain rekultivierter Anbauflächen als Lebensraum für Spinnen (Arachnida, Araneae) und Asseln (Isopoda, Oniscoiidea). – Acta Biologica Benrodis Supplement 6: 1-131
FINCH O.-D., H. KRUMMEN, F. PLAISIER & W. SCHULTZ (2007): Zonation of spiders (Araneae) and carabid bee-tles (Coleoptera: Carabidae) in island salt marshes at the North Sea coast. – Wetlands Ecology and Management 15: 207-228
GARDNER S. M. (1991): Ground beetle (Coleoptera: Carabidae) communities on upland heath and their as-sociation with heathland flora. – Journal of Biogeography 18: 281-289
GÖTZE W. (1992): Beweidung und Vertritt als Belas-tungsfaktoren der Spinnenfauna in Sandsalzwiese und Graue-Dünen-Formation. – Supplement zu Faunis-tisch-Ökologische Mitteilungen 13: 45-67
GRIGO M. (1997): Vergleichende Untersuchungen zur Spinnenfauna (Araneae) verschiedener Sandbiotope am Niederrhein. Diploma thesis University of Cologne. 134 pp.
HÄNGGI A. & B. BAUR (1998): The effect of forest edge on ground-living arthropods in a remnant of unferti-lized calcareous grassland in the Swiss Jura mountains. – Mitteilungen der Schweizerischen Entomologischen Gesellschaft 71: 343-354
HÄNGGI A., E. STÖCKLI & W. NENTWIG (1995): Le-bensräume mitteleuropäischer Spinnen. – Miscellanea Faunistica Helvetiae 4: 1-460
HARMS K.H. (1987): Spinnen und Weberknechte aus Grünlandbrachen des südlichen Pfälzerwaldes. In: ROWECK H. (ed.): Beiträge zur Biologie der Grünland-brachen im Südlichen Pfälzerwald. – Pollichia-Buch 12: 169-205
Spiders in dry ecosystems of North Rhine-Westphalia 15
HAUK B. (1996): Die Spinnenfauna ausgewählter Wachol-derheiden im Landkreis Calw. – Veröffentlichungen für Naturschutz und Landschaftspflege in Baden-Württem-berg / Beiheft 88: 259-288
HEIMER S. (1980): Eine bemerkenswerte Kugelspinne aus dem Harz. – Faunistische Abhandlungen Museum für Tierkunde Dresden 7: 179-181
HEIMER S. & W. NENTWIG (1991): Spinnen Mitteleuro-pas: ein Bestimmungsbuch. Parey, Berlin. 543 pp.
HELSDINGEN P. J. VAN (1999): Catalogus van de Neder-landse Spinnen (Araneae). – Nederlandse faunistische mededelingen 10: 1-191
HELSDINGEN P.J. VAN (2003): Halorates reprobus (O.P.-Cambridge, 1879) Nieuw voor Nederland. – Nieuwsbrief Spined 18: 10-11
HERZOG G. (1968): Beiträge zur Kenntnis der Spinnen-fauna der südlichen Mark. – Veröffentlichungen des Bezirksheimatmuseums Potsdam 16: 5-10
HERZOG G. (1974): Zur Spinnenfauna der westlichen Niederlausitz und benachbarter Gebiete. – Biologische Studien Luckau 3: 20-27
HEYDEMANN B. (1964): Die Spinnenfauna des Natur-schutzgebietes `Bottsand`, der Kohlberger Heide und des Schönberger Strandes (Araneae). – Faunistische Mitteilungen aus Norddeutschland 2: 133-141
HEYDEMANN B., W. GÖTZE & U. RIECKEN (1994): Öko-logische Analyse der Fauna des NSG `Barker Heide`. In: VOIGT, N. (ed.): Bedeutung von Heideökosystemen für die Wirbellosenfauna. – Supplement zu Faunistisch-Ökologische Mitteilungen 16: 13-47
HIEBSCH H. (1985): Beitrag zur Spinnenfauna der Moore im NSG `Serrahn`. – Zoologischer Rundbrief für den Bezirk Neubrandenburg 4: 15-33
HÖREGOTT H. (1958): Arachnologische Studien auf den Sandfluren bei Kleinsaubernitz/Oberlausitz. – Natura lusatica 4: 20-35
JÄGER P. (1996): Spinnen (Araneae) der Wahner Heide bei Köln. – Decheniana 35: 531-572
JAKOBITZ J. (2003): Neue und besonders gefährdete Spinnenarten (Araneae) für Brandenburg im NSG Pimpinellenberg. – Naturschutz und Landschaftspflege in Brandenburg 12(2): 51-53
JENTSCH A., W. BEYSCHLAG, W. NEZADAL, T. STEIN-LEIN & W. WELSS (2002): Bodenstörung – treibende Kraft für die Vegetationsdynamik in Sandlebensräumen - Konsequenzen für Pflegemaßnahmen im Naturschutz. – Naturschutz und Landschaftsplege 34: 37-44
JOCQUÉ R. (1977): Contribution à la connaissance des araignées de Belgique, V. Description de Theridion hublei n. sp. – Revue Arachnologique 1: 59-63.
JOGER H.G. (1997): Untersuchungen zur epigäischen Fauna von Halbtrockenrasen: Anpassungen von Spinnen und Insekten an einen Extrem-Lebensraum. Cuvelier, Göttingen. 210 pp.
KEER J. VAN & H. VANUYTVEN (1993): Catalogus van de Spinnen van Belgie. Del XI. Theridiidae, Anapidae en Theridiosomatidae. – Studiedocumenten van het Ko-ninklijk Belgisch Instituut voor Natuurwetenschappen 71: 7-44
KIELHORN K.-H. & T. BLICK (2007): Erstfund von Hahnia picta (Araneae, Hahniidae) in Deutschland – mit Angaben zu Habitatpräferenz und Verbreitung. – Arachnologische Mitteilungen 33: 7-10
KÖHLER G., V. VOPEL & R. BALLMANN (1989): Un-tersuchungen zum Einfluß der Verbuschung auf die Vegetations- und Faunenstruktur von Muschelkalk-steilhängen - ein Beitrag zur Sukzessionsforschung. – Archiv für Naturschutz und Landschaftsforschung 29 (3): 129-142
KRATOCHWIL A. (2004): Sand-Ökosysteme im Binnen-land: Dynamik, Restitution und Beweidungsmanage-ment – das Beispiel: Emsland. In: Westfälischer Na-turwissenschaftlicher Verein (ed.): Dünen und trockene Sandlandschaften – Gefährdung und Schutz. Verlag Wolf & Kreuels, Havixbeck-Hohenholte. pp. 13-21
KREMEN C., R.K. COLWELL, T.L. ERWIN, D.D. MUR-PHY, R.F. NOSS & M.A. SANJAYAN (1993): Terrestrial arthropod assemblages: their use in conservation plan-ning. – Conservation Biology 7: 796-808
KREUELS M. (1998a): Zur Frage strukturbezogener und phänologischer Anpassungen epigäischer Spinnen (Araneae) auf Kalkmagerrasen im Raum Marsberg. PhD thesis University of Münster. 108 pp.
KREUELS M. (1998b): Erstnachweis von Erigonoplus globipes (L. Koch, 1872) (Araneae: Linyphiidae) in Nordrhein-Westfalen und eine Verhaltensbeobachtung. – Arachnologische Mitteilungen 15: 77-80
KREUELS M. (2006): Die Webspinnen (Arachnida: Araneae) aus Beifängen des NSG Teverener Heide (NRW, Kreis Heinsberg). – Acta Biologica Benrodis 13: 185-193
KREUELS M. & S. BUCHHOLZ (2006): Ökologie, Ver-breitung und Gefährdungsstatus der Webspinnen Nordrhein-Westfalens. Verlag Wolf & Kreuels, Havix-beck-Hohenholte. 116 pp.
KREUELS M., S. BUCHHOLZ & V. HARTMANN (2008): Atlas der Webspinnen Nordrhein-Westfalens. Verlag Wolf & Kreuels, Bösensell. 135 pp.
KREUELS M. & J. LÜCKMANN (1998): Arachnologische und koleopterol. Ergebnisse der zoologischen Alpen-exkursionen der Westfälischen Wilhelms-Universität Münster nach Österreich in das Kl. Walsertal und in die Silvretta 1993-1997. – Jahrbuch Vorarlberger Lan-desmuseumsverein 1998: 9-17
KUSCHKA V. (2004): Ackerbrachen als Chance für den Na-turschutz? Struktur und Dynamik von Spinnenzönosen der Bodenoberfläche als Indikatoren der Habitatqualität. ökom Verlag, München. 200 pp.
16 S. Buchholz & M. Kreuels
LEIST N. (1978): Die Spinnen des Rußheimer Altrheins. In: Der Rußheimer Altrhein, eine nordbadische Auen-landschaft. – Die Natur- und Landschaftsschutzgebiete Baden-Württembergs 10: 365-398
LOCKET G.H. (1962): Miscellaneous notes on linyphiid spiders. – Annals and Magazine of Natural History (13) 5: 7-15
LOCKET G.H., A.F. MILLIDGE & P. MERRETT (1974): British Spiders 3. Ray Society, London. 315 pp.
LÖBF (Landesanstalt für Ökologie, Bodenordnung und Forsten NRW) (2005): Natur und Landschaft in Nord-rhein-Westfalen 2005. LÖBF-Mitteilungen 4/2005: 1-282
MALT S. & J. PERNER (2002): Zur epigäischen Arthro-podenfauna von landwirtschaftlichen Nutzflächen der Unstrutaue im Thüringer Becken. Teil 1: Webspinnen und Weberknechte (Arachnida: Araneae et Opiliones). – Faunistische Abhandlungen Museum für Tierkunde Dresden 22: 207-228
MARTIN D. (1973a): Theridion uhligi nov. spec., eine bis-her unbekannte Kugelspinne (Araneae: Theridiidae). – Deutsche Entomologische Zeitschrift (N.F.) 22: 113-115
MARTIN D. (1973b): Zur Kenntnis der Spinnenfauna des Naturschutzgebietes Rietzer See. Brandenburgische Naturschutzgebiete 16. – Naturschutzarbeit in Berlin und Brandenburg 9: 1-4
MARTIN D. (1983): Die Spinnenfauna des Naturschutzge-bietes 'Ostufer der Müritz'. – Zoologischer Rundbrief für den Bezirk Neubrandenburg 3: 1-40
MARTIN D. & M. UHLIG (1986): Die Spinnen- und Kurz-flüglerfauna (Araneae et Staphylinidae) der Silbergras-rasen (Corynephoreten) des Buhnenwerder, Stadtkreis Brandenburg, Bezirk Potsdam (Arachnida; Insecta, Coleoptera). – Faunistische Abhandlungen Museum für Tierkunde Dresden 14: 31-35
MERKENS S. (2002): Epigeic spider communities in inland dunes in the lowlands of Northern Germany. – Proceed-ings of the 19th European Colloquium of Arachnology: 215-222
MORITZ M. (1973): Neue und seltene Spinnen (Araneae) und Weberknechte (Opiliones) aus der DDR. – Deut-sche Entomologische Zeitschrift (N.F.) 20: 173-210
MUSTER C. (2001):Biogeographie von Spinnentieren der mittleren Nordalpen (Arachnida: Araneae, Opiliones, Pseudoscorpiones). – Verhandlungen des Naturwissen-schaftlichen Vereins in Hamburg (N.F.) 39: 5-196
NENTWIG W. (1983): Die Spinnenfauna (Araneae) eines Niedermoores (Schweinsberger Moor bei Marburg). – Decheniana 136: 43-51
NENTWIG W, A. HÄNGGI, C. KROPF & T. BLICK (2003): Spinnen Mitteleuropas / Central European Spiders. An internet identification key. Internet: http://www.araneae.unibe.ch, version 8.12.2003
NEW T.R. (1999): Untangling the web: spiders and the challenges of invertebrate conservation. – Journal of Insect Conservation 3: 251-256
OTTO B., T. SÜSSMUTH & F. MEYER (2001): Zur Schutz-würdigkeit und -bedürftigkeit von Verlandungsmooren in der Mittleren Mark - dargestellt am Naturschutzge-biet "Rauhes Luch" bei Luckenwalde. – Naturschutz und Landschaftspflege in Brandenburg 10 (2): 62-70
PARDEY A. (2004): Dünen und Sandlandschaften in Nord-rhein-Westfalen unter besonderer Berücksichtigung der Situation in Westfalen. In: WESTFÄLISCHER NATURWIS-SENSCHAFTLICHER VEREIN (ed.): Dünen und trockene Sandlandschaften – Gefährdung und Schutz. Verlag Wolf & Kreuels, Havixbeck-Hohenholte. pp. 3-11
PERNER J. (1997): Zur Arthropodenfauna der Kalktrocken-rasen im Mittleren Saaletal (Ostthüringen). Teil 1: Co-leoptera, Diptera, Auchenorrhyncha, Saltatoria, Araneae (Insecta et Arachnida). – Faunistische Abhandlungen Museum für Tierkunde Dresden 21: 53-90
PLATEN R. , B. V. BROEN, A. HERRMAN, U.M. RATSCH-KER & P. SACHER (1999): Gesamtartenliste und Rote Liste der Webspinnen, Weberknechte und Pseudoskor-pione des Landes Brandenburg (Arachnida: Araneae, Opiliones, Pseudoscorpiones) mit Angaben zur Häufig-keit und Ökologie. – Naturschutz und Landschaftspflege in Brandenburg 8 (2): 1-79
PLATNICK N.I. (2009): The world spider catalog, version 10.0. – American Museum of Natural History, Internet: http:// research.amnh.org/entomology/spiders/catalog/index.html (16.11.2009)
PRINSEN J.D. (1996): Theridion uhligi Martin, 1974 (Araneae: Theridiidae), een zeldzame kogelspin van heideterreinen. – Nieuwsbrief Spined 10: 4-7
RATSCHKER U.M. (2001): Die Zönose der Spinnen und Weberknechte in der Agrarlandschaft des Biosphä-renreservates Schorfheide-Chorin. PhD thesis TU Dresden. 218 pp.
RASKIN R. (2000): Renaturierung eines Heidemoores im Hohen Venn. Ergebnisse einer fünfjährigen ökologi-schen Effizienzkontrolle. – Naturschutz und Land-schaftsplanung 32: 212-221
RENNER K. (1982): Coleopterenfänge mit Bodenfallen am Sandstrand der Ostseeküste - ein Beitrag zum Problem der Lockwirkung von Konservierungsmitteln. – Faunis-tisch-Ökologische Mitteilungen 5: 127-146.
RIECKEN U., P. FINCK, U. RATHS, E. SCHRÖDER & A. SSYMANK (2006): Rote Liste der gefährdeten Biotop-typen Deutschlands. Zweite fortgeschrieben Fassung 2006. – Naturschutz und Biologische Vielfalt 34: 1-318
ROBERTS M.J. (1987): The spiders of Great Britain and Ireland. Volume 2: Linyphiidae and checklist. Harley Books, Colchester. 204 pp.
SACHER P. (1997): Webspinnen (Arachnida: Araneae) im Nationalpark Hochharz. – Berichte der Naturhistori-schen Gesellschaft Hannover 139: 259-276
Spiders in dry ecosystems of North Rhine-Westphalia 17
SACHER P. (1998): Spinnen (Araneae) an Fichten im Moorbereich - Ergebnisse von Klopffängen im Hoch-harz. – Abhandlungen und Berichte aus dem Museum Heineanum Halberstadt 4: 87-97
SACHER P. (2001): Beiträge zur Wirbellosenfauna des NSG "Harslebener Berge und Steinholz" im Nordharzvor-land/Sachsen-Anhalt. Teil 1: Webspinnen (Arachnida: Araneae). – Abhandlungen und Berichte aus dem Mu-seum Heineanum 5: 105-124
SACHER P. (2002): Webspinnen (Arachnida: Araneae). In: LANDESAMT FÜR UMWELTSCHUTZ SACHSEN-AN-HALT (ed.): Management von FFH-Lebensraumtypen: Untersuchungen zu den Auswirkungen von Maßnahmen zur Heide-Pflege (Flämmen, Mahd) auf Gliederfüßer (Arthropoda). – Berichte des Landesamtes für Umwelt-schutz Sachsen-Anhalt Sonderheft 3/2002: 15-17
SACHER P. & K. BREINL (1999): Neue Spinnenarten für Thüringen aus dem Kyffhäuser. (Arachnida: Araneae). – Thüringer Faunistische Abhandlungen 6: 51-60
SCHAEFER M. (1970): Einfluß der Raumstruktur in Land-schaften der Meeresküste auf des Verteilungsmuster der Tierwelt. – Zoologische Jahrbücher Systematik 97: 55-124
SCHAEFER M. (1972): Beitrag zur Kenntnis der Spinnen-fauna Schleswig-Holsteins (Araneae: Linyphiidae und Micryphantidae). – Schriften des Naturwissenschaftli-chen Vereins für Schleswig-Holstein 42: 94-103
SCHAEFER M. (1980): Sukzession von Arthropoden in ver-brannten Kiefernforsten. II. Spinnen und Weberknechte. – Forstwissenschaftliches Centralblatt 99: 341-356
SCHARFF N., J. A. CODDINGTON, C. E. GRISWORLD, G. HORMIGA & P. DE PLACE BJORN (2003): When to quit? Estimating spider species richness in a northern european deciduous forest. – Journal of Arachnology 31: 246-273
SCHMIDT L. & A. MELBER (2004): Einfluss des Heide-managements auf die Wirbellosenfauna in Sand- und Moorheiden Nordwestdeutschlands. – NNA-Berichte 17: 145-164
SCHMIDT M. H., I. ROSCHEWITZ, C. THIES & T. TSCHARNTKE (2005): Differential effects of landscape and management on diversity and density of ground-dwelling farmland spiders. – Journal of Applied Ecology 42: 281-287
SCHMIDT M.H., S. ROCKER, J. HANAFI & A. GIGON (2008): Rotational fallows as overwintering habitat for grassland arthropods: the case of spiders in fen meadows. – Biodiversity and Conservation 17: 3003-3012
SCHMITT M. (2008): Erstnachweis der Wolfspinne Alope-cosa barbipes (Araneae: Lycosidae) in Nordrhein-Westfa-len. – Arachnologische Mitteilungen 36: 1-3
SCHNITTER P.H., M. TROST & M. WALLASCHEK (eds.) (2003): Tierökologische Untersuchungen in gefähr-
deten Biotoptypen des Landes Sachsen-Anhalt. I. Zwergstrauchheiden, Trocken- und Halbtrockenrasen. – Entomologische Mitteilungen Sachsen-Anhalt, Son-derheft 2003: 1-216
SCHULTZ W. (1992): Beitrag zur Spinnenfauna (Arachnida, Araneida) der Tertiärdünen der ostfriesischen Insel Nor-derney. – Verhandlungen des naturwissenschaftlichen Vereins Hamburg (N.F.) 33: 239-245
SCHULTZ W. & F. PLAISIER (1995): Zum gegenwärtigen Besiedlungsstand der Strandinsel Minsener Oog durch Spinnen (Arachnida, Araneida) und Laufkäfer (Coleo-ptera, Carabidae). – Drosera 95: 85-100
SIMON U. (1995): Untersuchung der Stratozönosen von Spinnen und Weberknechten an der Waldkiefer. Wis-senschaft und Technik Verlag, Berlin. 142 pp.
SKERL K.L. (1999): Spiders in conservation planning: a survey of US natural heritage programms. – Journal of Insect Conservation 3: 341-347
SPARMBERG H. & P. SACHER (1997): Webspinnen an Binnensalzstellen Thüringens. – Thüringer Faunistische Abhandlungen 4: 44-55
STAUDT A. (2000): Neue und bemerkenswerte Spinnen-funde im Saarland und angrenzenden Gebieten in den Jahren 1996-99. – Abhandlungen der Delattinia 26: 5-22
STAUDT A. (2009): Nachweiskarten der Spinnentiere Deutschlands. – Internet: http://www.spiderling.de/arages
THALER K. (1986) Über wenig bekannte Zwergspinnen aus den Alpen, VII. – Mitteilungen der Schweizerischen Entomologischen Gesellschaft 59: 487-498
TUTELAERS P. (2000): Spinnen uit Cranendonck. – Ni-euwsbrief Spined 15: 5-21
TUTELAERS P. (2001): Herfstspinnen bij onze Hunebed-den. – Nieuwsbrief Spined 16: 7-9
VERBÜCHELN G. & M. JÖBGES (2000): Verbreitung und aktueller Zustand der Heide, Sandtrockenrasen und Borstgrasrasen in Nordrhein-Westfalen. – NUA-Hefte 6: 6-23
VINK C.J. & A. D. MITCHELL (2002): 12S DNA sequence data confirms the separation of Alopecosa barbipes and Alopecosa accentuata (Araneae: Lycosidae). – Bulletin of the British Arachnological Society 12: 242-244
WEBER M. (1999): Artenliste der Spinnen (Araneae) aus der Stadtbiotopkartierung Mainz. – Arachnologische Mitteilungen 17: 51-71
WIEHLE H. (1960): Spinnentiere oder Arachnoidea, XI. Micryphantidae - Zwergspinnen. Tierwelt Deutschlands 47. Fischer, Jena. 615 pp.
WIEHLE H. (1965): Beiträge zur Kenntnis der deutschen Spinnenfauna IV. – Mitteilungen aus dem Zoologischen Museum in Berlin 41: 11-57
18 S. Buchholz & M. KreuelsTa
b. 2
: Sp
ecie
s lis
t (n
om
encl
atu
re fo
llow
s Pl
atn
ick
2009
). A
bb
revi
atio
ns:
En
d =
sta
tus
of e
nd
ang
erm
ent
in N
ort
h R
hin
e-W
estp
hal
ia (a
cco
rdin
g t
o K
REU
ELS
& B
UC
HH
OLZ
200
6): *
= n
ot
end
ang
ered
, V =
en
dan
ger
men
t m
ay b
e as
sum
ed, R
= e
xtre
mel
y ra
re, n
ot
dec
linin
g s
pec
ies,
3 =
en
dan
ger
ed, 2
= h
igh
ly e
nd
ang
ered
, 1 =
in im
min
ent
dan
ger
of b
eco
min
g e
xtin
ct,
0 =
ext
inct
, nr
= n
ew r
eco
rd fo
r N
ort
h R
hin
e-W
estp
hal
ia, ?
= h
ith
erto
no
t as
sess
ed; D
is =
Dis
trib
uti
on
: C =
sp
ecie
s th
at e
xclu
sive
ly o
ccu
rred
on
cal
care
ou
s su
bst
rate
s.
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]
A
typi
dae
[2]
A
typu
s affi
nis
*
322
34A
typu
s pice
us
1
44
Sege
stri
idae
[1]
Se
gest
ria
seno
cula
ta*
1
21
14
211
Dys
deri
dae
[3]
D
ysde
ra er
ythr
ina
*
126
146
12
8925
6H
arpa
ctea
hom
berg
i*
4
1115
Har
pact
ea r
ubicu
nda
*
11
Mim
etid
ae [1
]
Ero
furc
ata
*
11
11
12
310
Nes
tici
dae
[1]
N
esti
cus c
ellu
lanu
s*
C2
2T
heri
diid
ae [2
0]
Asa
gena
pha
lera
ta*
19
3928
3051
91
555
226
423
599
1542
9C
rust
ulin
a gu
ttat
aV
1
46
11D
ipoe
na co
racin
a2
6
463
7713
2E
nopl
ogna
tha
lati
man
a3
1
12
Eno
plog
nath
a ov
ata
*
12
14
Eno
plog
nath
a th
orac
ica
*
407
411
113
2113
174
217
615
49
19
119
313
Epi
sinus
ang
ulat
us*
C1
1E
pisin
us tr
unca
tus
*
12
3E
uryo
pis fl
avom
acul
ata
*
41
413
15
333
291
11
298
Neo
ttiu
ra b
imac
ulat
a*
6
6Pa
idisc
ura
palle
ns*
C7
1724
Phol
com
ma
gibb
um*
1
11
14
Phyl
lone
ta im
pres
sa*
2
2Ph
yllo
neta
sisy
phia
*C
11
Rob
ertu
s aru
ndin
eti
*
19
111
Rob
ertu
s liv
idus
*
26
1527
61
71
17
17
283
Rob
ertu
s neg
lect
us*
C1
12
4Se
ycel
loce
sa v
itta
tus
*
11
2St
eato
da a
lbom
acul
ata
3
483
455
The
ridi
on u
hlig
i nr
1
12
Lin
yphi
idae
[160
]
Aba
copr
oece
s sal
tuum
R
33
Aca
rtau
chen
ius s
curr
ilis
2
11
Spiders in dry ecosystems of North Rhine-Westphalia 19
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]
A
gnyp
hant
es ex
punc
tus
*
11
Agy
neta
caut
a 3
1
12
Agy
neta
coni
gera
*
11
Agy
neta
dec
ora
*
22
Agy
neta
subt
ilis
*
14
5A
llom
enge
a vi
dua
*
1414
Ang
ulip
hant
es a
ngul
ipal
pis
RC
11
Ara
eonc
us cr
assic
eps
*
11
Ara
eonc
us h
umili
s*
36
9811
2058
1317
144
1012
374
317
206
459
1A
sthe
narg
us p
agan
us*
1
54
10B
aryp
hym
a pr
aten
se
*
11
Bat
hyph
ante
s app
roxi
mat
us*
10
15
31
21
23B
athy
phan
tes g
racil
is*
13
942
277
17
210
21
162
310
11
22
213
273
293
Bat
hyph
ante
s nig
rinu
s*
7
65
422
Bat
hyph
ante
s par
vulu
s*
13
91
101
236
Bol
ypha
ntes
alti
ceps
*C
11
Cen
trom
erita
bico
lor
*
3127
524
1078
11
111
3491
503
210
961
3C
entr
omer
ita co
ncin
na*
8
1658
920
23
126
223
196
38
7019
11
142
314
066
530
81C
entr
omer
us b
revi
vulv
atus
*
1
11
3C
entr
omer
us ca
vern
arum
RC
11
Cen
trom
erus
dilu
tus
*
33
21
41
11
117
Cen
trom
erus
incil
ium
*
1217
42
2267
1121
581
421
042
9C
entr
omer
us le
vita
rsis
R
31
4C
entr
omer
us p
abul
ator
*
401
56
71
446
55
103
52
238
792
Cen
trom
erus
pru
dens
*
1
12
Cen
trom
erus
sylv
aticu
s*
65
1911
216
1549
482
385
168
69
1262
383
Cer
atin
ella
bre
vipe
s*
1
11
11
5C
erat
inel
la b
revi
s*
1
11
11
38
Cer
atin
ella
scab
rosa
*
61
18
Cne
phal
ocot
es o
bscu
rus
*
443
189
22
9713
195
212
Col
linsia
dist
inct
a*
15
215
2C
ollin
sia in
erra
ns*
2
31
12
31
114
Dic
ymbi
um n
igru
m b
revi
seto
sum
V
11
101
31
28
81
310
62
2178
Dic
ymbi
um ti
bial
e*
2
427
79
150
Dip
loce
phal
us co
nnat
usR
1
1D
iplo
ceph
alus
crist
atus
*
62
615
216
6D
iplo
ceph
alus
lati
fron
s*
9
132
143
Dip
loce
phal
us p
icinu
s*
2
16
110
Dip
lost
yla
conc
olor
*
12
177
24
11
52
21
61
205
Dism
odicu
s bifr
ons
*
13
32
9E
ntel
ecar
a co
ngen
era
*C
33
20 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]
E
ntel
ecar
a er
ythr
opus
*
11
2E
rigo
ne a
rcti
ca m
arit
ima
3
221
462
71E
rigo
ne a
tra
*
155
6866
7581
312
230
659
21
111
129
1069
22
19
232
89
347
311
1872
Eri
gone
den
tipa
lpis
*
973
236
641
110
437
602
125
23
531
84
6510
647
31
138
642
03
452
90E
rigo
ne lo
ngip
alpi
s*
1
119
41
127
Eri
gone
lla h
iem
alis
*
11
12
623
34E
rigo
nopl
us g
lobi
pes
0C
9696
Flo
roni
a bu
ccul
enta
*
11
Gna
thon
ariu
m d
enta
tum
*
33
Gon
atiu
m p
arad
oxum
*C
11
Gon
atiu
m r
ubel
lum
*
31
4G
onat
ium
rub
ens
*
13
164
325
138
268
143
Gon
gylid
iellu
m la
tebr
icola
*
13
21
83
93
13
21
11
140
Gon
gylid
iellu
m v
ivum
*
154
41
11
11
24
373
Gon
gylid
ium
rufi
pes
*
71
8H
alor
ates
repr
osus
R
2
2H
ypom
ma
bitu
berc
ulat
um*
19
19H
ypse
liste
s jac
kson
iR
1
1Ja
ckso
nella
falco
neri
*
16
2027
Lep
thyp
hant
es m
inut
us*
1
12
4L
epto
rhop
trum
robu
stum
*
5252
Les
sert
ia d
enti
chel
isR
1
1L
inyp
hia
hort
ensis
*
11
Lin
yphi
a te
nuip
alpi
s R
1
1L
inyp
hia
tria
ngul
aris
*
22
15
Lop
hom
ma
punc
tatu
m
*
22
Mac
rarg
us r
ufus
*
2338
18
82
15
33
12
95M
aso
galli
cus
R
11
Mas
o su
ndev
alli
*
12
22
18
Mei
onet
a af
finis
3
12
5255
Mei
onet
a in
nota
bilis
*
1
1M
eion
eta
mol
lis*
2
2M
eion
eta
rure
stri
s*
50
4894
1652
720
11
3523
116
68
1734
12
917
192
710
78
111
928
Mei
onet
a sa
xati
lis*
2
21
32
10M
eion
eta
simpl
icita
rsis
0
77
Mer
mes
sus t
rilo
batu
s*
3
12
15
11
210
26M
etop
obac
trus
pro
min
ulus
*
13
1014
Micr
argu
s ape
rtus
*
11
Micr
argu
s her
bigr
adus
*
73
211
24
71
53
91
32
11
63
3110
2M
icrar
gus s
ubae
qual
is*
16
47
14
234
Micr
olin
yphi
a pu
silla
*
11
817
11
12
12
11
37
Spiders in dry ecosystems of North Rhine-Westphalia 21
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]
M
icron
eta
viar
ia*
2
54
11
21
39
28M
inyr
iolu
s pus
illus
*
12
123
91
11
30M
ioxe
na b
land
aV
4
21
148
56M
onoc
epha
lus f
uscip
es*
2
2N
erie
ne cl
athr
ata
*
11
18
11N
erie
ne fu
rtiv
aR
1
12
Ner
iene
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