Millipede and isopod auna o Swiss ci ies 199
Occu ence and assemblage composi ion o millipedes
(My iapoda, Diplopoda) and e es ial isopods
(C us acea, Isopoda, Oniscidea) in u ban a eas
o Swi ze land
Fe enc Vilisics1, Dá id Bogyó2, Thomas Sa le 3, Ma co Mo e i3
1 Uni e si y o Helsinki, Facul y o Biological and En i onmen al Science, Depa men o En i onmen al
Science, 00014 Helsinki, Viikinkaa i 2, Finland 2 Depa men o Ecology, Uni e si y o Deb ecen, H-4010
Deb ecen, PO Box 71, Hunga y 3 Swiss Fede al Resea ch Ins i u e WSL, Communi y Ecology Resea ch Uni ,
Via Belsoggio no 22, 6500 Bellinzona, Swi ze land
Co esponding au ho : Fe enc Vilisics ([email p o ec ed])
Academic edi o : S. S en hou akis|Recei ed 29 Sep embe 2011|Accep ed 17 Feb ua y 2012|Published 20 Ma ch 2012
Ci a ion: Vilisics F, Bogyó D, Sa le T, Mo e i M (2012) Occu ence and assemblage composi ion o millipedes
(My iapoda, Diplopoda) and e es ial isopods (C us acea, Isopoda, Oniscidea) in u ban a eas o Swi ze land. In: Š us J, Tai-
i S, S en hou akis S (Eds) Ad ances in Te es ial Isopod Biology. ZooKeys 176: 199–214. doi: 10.3897/zookeys.176.2153
Abs ac
Te es ial isopods and millipedes, membe s o he in e eb a e mac o-decompose guild, we e collec ed
h ough pi all aps in h ee Swiss ci ies (Zu ich, Luce ne, Lugano). A o al o 7,198 indi iduals o 17
isopod species (7093 ind.), and 10 millipede species (105 ind.) we e cap u ed. Besides he Alpine en-
demic isopod (T ichoniscus alemannicus) and millipede (Cylind oiulus e hoe i), u ban assemblages we e
mainly composed o widesp ead, na i e Eu opean and e en cosmopoli an species, which a e equen in
an h opogenic a eas. O e all species ichness (isopods and millipedes combined) was simila in Zu ich
(17 species) and Luce ne (16), while only 13 species we e sampled in Lugano. Acco ding o he Sø ensen
index o simila i y, species composi ion o Zu ich and Luce ne we e mo e alike, while he one o Lugano
was mo e dis inc om he o he wo ci ies. This esul can be explained by he spa ial p oximi y o Zu ich
and Luce ne in he no h o he Alps compa ed o Lugano, which is loca ed mo e dis an ly and in he
sou h o he Alps. Dominan isopods and millipedes in Zu ich and Luce ne we e ound o be widesp ead
synan h opic species in empe a e Eu ope (Po cellio scabe , T achelipus a hkii and Ophyiulus pilosus) while
he dominan isopod in Lugano (T achelipus azzau ii) is a species wi h a no h-eas e n Medi e anean
dis ibu ion. Ou s udy e eals ha he u ban millipede and isopod auna in Swiss ci ies mainly consis s
o widesp ead species, bu species o na owe dis ibu ion (e.g. T. alemannicus, C. e hoe i ) may also ind
sui able habi a s in ci ies. Despi e some signs o bio ic homogeniza ion, ou s udy also ound composi-
ional di e ences o millipede and isopod assemblages be ween no he n and sou he n ci ies ha sugges
geog aphical e ec s o he egional species pool.
ZooKeys 176: 199–214 (2012)
doi: 10.3897/zookeys.176.2153
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Fe enc Vilisics e al. / ZooKeys 176: 199–214 (2012)
200
Keywo ds
Decompose s, u baniza ion, woodlice, u ban biodi e si y, a h opods
In oduc ion
As one o he majo ac o s o global change, u baniza ion and i s e ec s on biodi e -
si y ha e a ac ed g ea scien i ic a en ion in he pas decade (e.g. McDonnell and
Hahs 2008, Niemelä e al. 2000). Nume ous s udies ha e added an inc easing knowl-
edge o he unde s anding o he ecology o many axa in u ban en i onmen s, such
as plan s (e.g. Walke e al. 2009, Gulezian and Nybe g 2010), insec s (e.g. Magu a
e al. 2004, Sa le e al. 2010a, 2010b, 2011), spide s (e.g. Magu a e al. 2010), and
e eb a es (e.g. Melles e al. 2003, Tó h e al. 2009, Fon ana e al. 2011).
U ban soil meso- and mac o-a h opods ha e ecei ed less a en ion (bu see Ko -
sós e al. 2002, Vilisics e al. 2007), despi e hei impo ance in ecosys em p ocesses
such as decomposi ion o o ganic ma e (Hassall e al. 1987). Decay o dead plan
ma e esul s in ions eadily a ailable o up ake o plan s. The majo i y o allen lea es
and woody deb is a e b oken down by mic obes, ungi and in e eb a es (Be g and
McClaughe y 2003).
Soil mac o- meso- and mic o-in e eb a es con ibu e in he decomposi ion cas-
cade by ei he agmen ing, o u he mine alizing dead plan ma e (Ve hoe and
B ussaa d 1990). Key o ganisms in ini ial b eakdown and comminu ion o dead ma -
e a e isopods, millipedes, e mi es, an s, and membe s o o he in e eb a e g oups
(Coleman and Hend ix 2000), while nema odes and annelids a e essen ial o mine -
aliza ion (Ve hoe and B ussaa d 1990, Flegel and Sch ade 2000). Thus, millipedes
(My iapoda: Diplopoda) and isopods (C us acea, Isopoda: Oniscidea) belong o he
same unc ional guild, e en hough hey a e axonomically qui e dis an (Scheu and
Falca 2000). Based on his common ecosys em unc ion, we p opose o pool he wo
axa in he same analyses which, o he bes o ou knowledge, is a a he no el ap-
p oach in soil zoology and ecology.
Millipedes and isopods a e known o inhabi Eu opean u ban habi a s, mainly by
cosmopoli an and Hola c ic species (Smi h e al. 2006, Riedel e al. 2009, Vilisics and
Ho nung 2009). Human ac i i ies, such as ga dening, anspo a ion o soil, and cul i-
a ion o o namen al plan s, a e suspec ed o be he mos impo an ac o s o species
exchanges o less mobile o ganisms such as millipedes and isopods be ween dis an
loca ions. A he same ime o he s udies (Bogyó and Ko sós 2009, Riedel e al. 2009)
ha e shown ha also he na i e species pool has an e ec on u ban soil auna, p o id-
ing subs a e o na i e species ypical in na u al and semi-na u al habi a s in ci ies.
Pa icula u ban habi a s, such as bo anical and p i a e ga dens and pa ks, howe e ,
ha bou es ablished popula ions o alien soil a h opods oo (Vilisics and Ho nung
2009, Cocha d e al. 2010).
U ban assemblages o soil in e eb a es show con o e sial pa e ns o species com-
posi ion: some s udies sugges ha species composi ions di e along u baniza ion g a-
Millipede and isopod auna o Swiss ci ies 201
dien s (Ko sós 1992, Ko sós e al. 2002, Bogyó and Ko sós 2009, Riedel e al. 2009),
while o he s ha e shown no di e ences along simila g adien s (Ho nung e al. 2007,
Vilisics e al. 2007). In his s udy we analyse he occu ence and assemblage composi-
ion o u ban millipede and isopod aunas in h ee u ban a eas o Swi ze land. We
discuss he mos in e es ing indings wi h espec o Eu opean soil auna. This con i-
bu ion is pa o he Biodi e Ci y p ojec (www.biodi e ci y.ch) ha aims o assess bio-
di e si y in u ban en i onmen s and i s accep ance by ci izens in he amewo k o he
Swiss na ional esea ch p og am ‘Sus ainable de elopmen o he buil en i onmen ’.
Me hods
S udy si es
The s udy ook place in h ee Swiss ci ies, namely Zu ich (371,000 inhabi an s /92 km2),
Luce ne (59,000 /24 km2) and Lugano (49,000/ 26 km2), which ep esen small o me-
dium sized ci ies in cen al Eu ope. The ci ies s udied lay on a no h o sou h g adien
(app ox. 200 km, wi h Lugano sou h o he Alps) and all a e bo de ed by a lake and moun-
ains > 800 m. O iginally, 36 sampling si es we e selec ed in each ci y bu a he end only
106 could be used o he analyses: 36 in Zu ich and Lugano, 34 in Luce ne.
The h ee ci ies sha e common ea u es such as his o ical cen es, esiden ial a eas,
business qua e s, public g een a eas, pa ks and ceme e ies, and o me indus ial a eas.
The ci ies a e cha ac e ized by mode a e empe a u e (No h: a e age Janua y em-
pe a u e 1°C, July 17°C; Sou h: Janua y 3°C, July 20°C) wi h an annual p ecipi a ion
o 1000 mm o Zu ich, 1150 mm o Luce ne and 1600 mm o Lugano.
Wi hin each o he h ee ci ies sampling poin s we e selec ed along a con inuous
u baniza ion g adien , which was measu ed as he ac ion o sealed and buil a ea in
he 50 m adius a ound he sampling poin s. The selec ion o he indi idual sampling
poin s ollowed a easoned choice sampling s a egy o co e he en i e u banisa ion
g adien (3% o 92% sealed and buil a ea). We included a wide ange o u ban habi-
a ypes (p i a e ga dens, semi-public spaces o apa men buildings, public pa ks
and cou ya ds o indus ial buildings) in o he s udy. Mean dis ance be ween s udy
si es was 388 m (± 21 m SE). A minimal dis ance o 250 me es was kep be ween
sampling si es and he ci y inge. P ecise loca ions o he s udy si es a e gi en in
Ge mann e al. (2008).
Da a collec ion
Isopods and millipedes we e sampled h ough pi all aps, consis ing o 3 plas ic cups
(opening diame e 75 mm) pe ap si e ecessed in o he soil and a anged in an isos-
celes iangle wi h a dis ance o one me e . T anspa en oo s ins alled app oxima ely 8
cm abo e he cups p o ided p o ec ion om ain. T aps we e emp ied weekly du ing
Fe enc Vilisics e al. / ZooKeys 176: 199–214 (2012)
202
7 weeks om June 13 h o Augus 3 d 2006 (Sa le e al. 2010a), which co esponds o
he pe iod wi h highes a h opod ac i i y in Swi ze land (Duelli e al. 1999, Ob is
and Duelli 2010).
Iden i ica ion o millipedes was based on Schuba (1934) and Blowe (1985). Fo
isopod iden i ica ion we used he keys o Schmölze (1965) and G une (1966). Valid
nomencla u e was applied acco ding o Schmal uss (2003) and Engho (2010). The e -
e ence collec ion is deposi ed a he Na u al His o y Museum in Lugano (Swi ze land).
Da a analyses
We used species ichness (numbe o species) as he mos common measu e o quan i y
biodi e si y (Magu an 2004). Incidence is he equency wi h which he species oc-
cu s a all in he s udy si es o a ci y. This alue was used o assess s eadiness o a spe-
cies in he h ee ci ies. This alue is indica i e o he egula i y o a species’ occu ence
which does no necessa ily co ela e wi h abundance. We ega ded incidence a e ‘high’
when i was o e 50%, and ‘low’ when i did no each 10%.
The simila i y be ween he millipede and isopod species assemblages combined
sampled in he h ee ci ies was assessed using he Sø ensen index (Sø ensen 1948),
which is a widely used index in ecology and hus sui able o compa a i e pu poses.
Fo hese analyses we used da a o specimens iden i ied o species le el only.
Resul s
Species ichness and composi ion
O e all, 17 species o isopods (7015 indi iduals) and 8 species millipedes (98 ind.)
we e iden i ied in he h ee s udied Swiss ci ies; one isopod could only be iden i ied a
genus le el, while wo millipedes only a amily le el.
Isopod species ichness was highes in Luce ne (14 species) and lowes in Lugano
(10 species), wi h 11 species in Zu ich (Table 1). One hi d (6 species) o all Isopoda
species occu ed in all h ee ci ies. Fi e addi ional species we e cap u ed in bo h Lu-
ce ne and Zu ich bu no in Lugano (Table 1). Th ee isopod species we e dominan in
he h ee ci ies (Fig. 1), i.e. he cosmopoli an Po cellio scabe La eille, 1804 in Zu ich
(1216 indi iduals, 32% ela i e abundance); he widesp ead Eu opean T achelipus
a hkii (B and , 1833) in Zu ich (1934, 33%) and Luce ne (1234, 72%), and he
Medi e anean T achelipus azzau ii (A cangeli, 1913) in Lugano (307, 52.3%).
We ound se en millipede species in Zu ich, ou in Lugano, and h ee in Luce ne.
Ophyiulus pilosus (Newpo , 1842) occu ed in all h ee ci ies, bu was dominan in Zu-
ich (62.5%) and Lugano (42.5%), while Polydesmus angus us La zel, 1884 was dominan
in Luce ne (85%) (Fig. 1). Two o he species [B achydesmus supe us La zel, 1884 and Ox-
ydus g acilis (C. L. Koch, 1847)] we e exclusi ely ound in Zu ich and Luce ne (Table 1).
Millipede and isopod auna o Swiss ci ies 203
able 1. Incidence o isopod and millipede species in sampling si es in he ci ies o Zu ich, Luce ne,
and Lugano.
Species
Species occu ences in aps
Zu ich
(n=36)
Luce ne
(n=34)
Lugano
(n=36)
And oniscus den ige Ve hoe , 1908 + + 0
A madillidium nasa um Budde-Lund, 1885 + + +
A madillidium ulga e (La eille, 1804) + + +
Cylis icus con exus (De Gee , 1778) + + 0
Haploph halmus danicus Budde-Lund, 1880 0 0 +
Hyloniscus ipa ius (C. Koch, 1838) +! +! +
Ligidium hypno um (Cu ie , 1792) 0 + 0
Oniscus asellus Linnaeus, 1758 +! +! 0
O home opon planum (Budde-Lund, 1885) 0 0 +
Philoscia musco um (Scopoli, 1763) + +! 0
Pla ya h us ho mannseggii (Budde-Lund, 1893)* + + +
Po cellio scabe La eille, 1704 +! +! +
Po cellionides p uinosus (B and , 1833) + 0 0
T achelipus a hkii (B and , 1833) +! +! +
T achelipus azzau ii (A cangeli, 1913) 0 0 +!
T ichoniscus alemannicus Ve hoe , 1917 + + 0
T ichoniscus pusillus B and , 1833 0 + 0
T ichoniscus sp.0++
Isopod species 11 13 9
Isopod specimens 3738 2690 587
B achydesmus supe us La zel, 1884 + 0 +
Cylind oiulus cae uleocinc us (Wood, 1864) + 0 0
Cylind oiulus e hoe i (B olemann, 1896) 0 0 +
Nemasoma a ico ne C. L. Koch, 1847 + 0 0
Ophyiulus pilosus (Newpo , 1842) + + +!
Oxidus g acilis (C. L. Koch, 1847) 0 + +
Polydesmus angus us La zel, 1884 0 + 0
P opolydesmus es aceus (C. L. Koch, 1847) + 0 0
Cho deuma idae sp.0+0
C aspedosoma idae sp.00+
inde + +00
Millipede species 5 3 4
Millipede specimens 32 27 46
Legend: 0: species absen ; +: species p esen ; +!: species p esen a > 25% o si es pe ci y; * axa iden i ied
as a my mecophilous species (ubiqui ous in an nes s); numbe o specimens we e no coun ed, see ex
o explana ion
Millipedes show a di e en occu ence pa e n han isopods, o he la e he
di e ences among he h ee ci ies seem o be g ea e . O e all, 50% o all millipede
and isopod species we e obse ed in only one ci y. The mean numbe o indi iduals

Fe enc Vilisics e al. / ZooKeys 176: 199–214 (2012)
204
Figu e 1. Abundances o isopods and millipedes in Zu ich, Luce ne and Lugano in loga i hmic scale.
Dashed line sepa a es isopods (on he le ) om millipedes. Black ba s ep esen abundances, whi e ci cles
show he species’ incidences pe all si es pe ci y.
Millipede and isopod auna o Swiss ci ies 205
o isopods and millipedes pe si e a ied subs an ially in he h ee ci ies, i.e. Luce ne
[mean 159.7 (SE: 75.3)], Lugano [mean 45 (SE: 12.56)], and Zu ich [mean 221.7
(SE: 77.22)]. Sø ensen simila i y index o species composi ions (isopods and millipedes
combined) was highes be ween Zu ich and Luce ne (0.67) and lowes be ween Zu ich
and Lugano (0.40), wi h an in e media e alue o 0.58 be ween Luce ne and Lugano.
Species incidence
Incidences o isopod and millipede species pe ci y we e gene ally low, i.e. below 10% o
he o al numbe o aps (Figu e 1). Fo isopods, he mos abundan species we e also
he mos equen ones. The mos widesp ead isopod in Zu ich was P. scabe (55.6% o
36 aps), while in Luce ne i was T. a hkii (58.8% o 34 aps), and in Lugano T. az-
zau ii (44.4% o 36 aps). Luce ne and Zu ich, ci ies loca ed no h o he Alps, sha ed
ou isopod species [Hyloniscus ipa ius (C. Koch, 1838), P. scabe , Oniscus asellus Lin-
naeus, 1758, and T. a hkii] ou o he i e mos equen ly sampled (> 25%). The only
millipede wi h an incidence o e 25% (ou o 36 aps) was O. pilosus in Lugano, while
he es o millipede species o ou s udy occu ed wi h incidences < 10% (Figu e 1).
The e we e also examples whe e highe ela i e abundances pai ed wi h ela i ely
low incidences (<10% occu ence pe all si es o pe all ci y) which sugges an agg e-
ga ed dis ibu ion o some species. This was he case o T. a hkii ( ela i e abundance
20%; incidence 6%) and A madillidium ulga e (La eille, 1804) in Lugano (14%;
6%), and A madillidium nasa um Budde-Lund, 1885 in Zu ich (30%; 6%).
Discussion
Faunis ic esul s
Nex o many axonomic and aunis ic s udies on Eu opean isopods (Schmal uss and
Wol -Schwenninge 2002), o he bes o ou knowledge, he e we e nei he e iews
no aunis ic pape s published on Swiss Isopoda auna in he las 100 yea s since he
las o e iew by Ca l (1911). The ecen millipede auna has been summa ized by
Ped oli-Ch is en (1993), desc ibing 127 species om he coun y. The e o e ou e-
sul s a e somewha challenging o in e p e a ion and need o be pu in con ex by
conside ing o he s udies on he Eu opean le el.
Ou esul s e eal ha he obse ed ci ies ha bou mos ly species widesp ead in
Eu ope. Rega ding isopods, six species a e known as widesp ead in empe a e and
no he n Eu ope, occupying bo h u banized and u al a eas: A. ulga e, O. asellus,
Philoscia musco um (Scopoli, 1763), P. scabe , T. a hkii, T ichoniscus pusillus B and ,
1833 (e.g. Ho nung e al. 2007, Vilisics e al. 2007). O home opon planum (Budde-
Lund, 1885), a Cen al Eu opean species, is known o dwell in b oadlea o es s
Fe enc Vilisics e al. / ZooKeys 176: 199–214 (2012)
206
(Vilisics e al. 2008), bu has also been ound in he u ban inge o Budapes (Vilisics
and Ho nung 2009). The only equen isopod species in Lugano, T. azzau ii, seems,
ins ead, o be mainly es ic ed o no heas e n Medi e anean (Schmal uss 2003).
Among millipedes, se en species a e widely dis ibu ed ac oss Eu ope (Engho
2010), six o which a e known o occu in a eas unde human in luence: B. supe us,
Cylind oiulus cae uleocinc us (Wood, 1864), O. pilosus, O. g acilis, P. angus us, and
P opolydesmus es aceus (C. L. Koch, 1847) (Blowe 1985, Kime 1990, Ped oli-Ch is en
1993, Tadle and Thale 1993).
One isopod (T ichoniscus alemannicus Ve hoe , 1917) and one millipede (C.
e hoe i) species a e known o be es ic ed o he Alps (A ems 1949, Ped oli-Ch is en
1993, Schmal uss, 2003). These cases show ha u ban a eas may suppo he su i al
also o na i e species wi h a mo e es ic ed Eu opean dis ibu ion.
The isopod Pla ya h us ho mannseggii (Budde-Lund, 1893), a depigmen ed and
blind my mecophilous species, ubiqui ous in an nes s, was equen ly sampled in Lu-
ce ne. The dis ibu ion pa e n o such isopods may ollow he dis ibu ion o hei
an hos s.
The in a-Eu opean alien (Cocha d e al. 2011) A. nasa um is commonly
in oduced o g eenhouses ac oss Eu ope (Schmal uss 2003). Repo s om a ious
ci ies show ha he species can su i e in ou doo habi a s, oo (e.g. Ams e dam: Be g
e al. 2008; Budapes : Vilisics and Ho nung 2010). The o he non-na i e species was
he millipede O. g acilis, a ue alien o opical eas e n Asian o igin. I has been in-
oduced by human ac i i ies in o Eu opean g eenhouses (Blowe 1985) and seldom
su i es ou doo s in Eu ope (Ped oli-Ch is en 1993).
Species composi ions, abundances and incidence
Ou su ey in h ee Swiss ci ies esul ed in a ela i ely high species ichness and abun-
dance o isopods, while millipedes we e cap u ed in lowe numbe o species and in-
di iduals.
The o e all numbe o millipede species e ealed in he h ee ci ies is ela i ely
low (8.7%) compa ed wi h he known millipede auna o Swi ze land (127 species;
Ped oli-Ch is en 1993). In empe a e Eu ope he a e age u ban species numbe is
be ween 14 and 26 (Engho 1973, Tischle 1980, Ko sós 1992, Ko sós e al. 2002,
S oe 2004, Bogyó and Ko sós 2009, Riedel e al. 2009), despi e he ac ha only 9
species we e eco ded in London by Smi h e al. (2006).
Repo s on u ban isopod and millipede auna show ela i ely high species ichness
in empe a e ci ies as compa ed o known na i e, local aunas. Such epo s a e, how-
e e , ha dly compa able due o di e ences in sampled habi a s, sampling e o and
me hodology. Pi all apping in pa ks o he ci y Deb ecen (Hunga y) esul ed o a
19% (14 species) o he known millipede auna o Hunga y (Bogyó and Ko sós 2009).
A simila me hod cap u ed ca. 14% (11 species) o he known millipede auna o he
Millipede and isopod auna o Swiss ci ies 207
Czech Republic om pa ks o he ci y Olomouc (Riedel e al. 2009). Pi all apping
om pa ks and nea by u al o es ed a eas in Deb ecen and So ø (Denma k) esul ed
in Isopoda species (Ho nung e al. 2007, Vilisics e al. 2007) comp ising 11% o Hun-
ga ian and ca. 27% o known Danish Isopoda auna (Meine z 1964).
Pooled abundances o isopods and millipedes we e well o e 1000 indi iduals in
each o he abo e men ioned s udies (90 o 120 ope a ing aps o 6 o 9 mon hs). As
he numbe o millipedes in ou s udy was less han 100 indi iduals, he ques ion a ises
why abundances we e so low in he h ee Swiss ci ies? In his e iew Da id (2009) has
pinpoin ed he nega i e e ec s o habi a loss and low ood quali y on millipede assem-
blages, and demog aphy. Fu he mo e, managemen p ac ices al e ing mic ohabi a s
(like coa se woody deb is and bo h li e quan i y and quali y) has g ea e ec on soil
mac o-in e eb a es, including isopods and millipedes. (e.g. Topp e al. 2006, Kappes
e al. 2009). The low ac i i y densi y alues in ou s udied ci ies may hus be a esul o
he in ensi e managemen such as mowing and emo al o plan li e .
The only alien isopod cap u ed, A. nasa um, also known as he “g eenhouse pill-
bug”, was among he domina ing species in Zu ich, bu i only occu ed in 5.7% o he
sampling si es (ou o he o al 36), u ning o be one o he a es species in he ci y.
Simila ly, he alien millipede O. g acilis (S oe e al. 2010) showed low incidences, as i
appea ed in only 3% o he si es in Luce ne, and 12% o Lugano. We assume ha hese
species we e ei he in oduced ecen ly in hese places o su i e in si es which p o ide
special en i onmen al condi ions (such as highe annual a e age empe a u e), so hey
may agg ega e in high numbe s a ce ain spo s o a ci y, while ne e es ablish in o he s.
The h ee millipedes (O. pilosus, B. supe us, O. g acilis) occu ing in wo o mo e
ci ies a e widesp ead in Eu ope and occupy u al as well as u ban se lemen s (Blowe
1985, Ped oli-Ch is en 1993, Engho 2010). Ophyiulius pilosus in Lugano showed he
highes incidence o occu ences among he millipedes. We ound a somewha simila
end in isopods, as widely dis ibu ed species showed he g ea es incidences. Mo e-
o e , his esul is consis en wi h Kime (1990) and Voig lände (2011), who sugges ed
ha his species migh p o i om human ac i i ies and dis u bances.
Pi all apping, as he sampling me hod employed in his s udy, is a passi e sam-
pling me hod ha has been de eloped o ca ch specimens ac i e on he soil and li e
su ace. I has p o en o e icien ly ep esen species ichness and ac i i y densi y da a
o se e al a h opod g oups (e.g. A aneae, Coleop e a) (S anden 2000). As only a ac-
ion o isopod and millipede species (e.g. isopods: amily T ichoniscidae, millipedes:
Geoglome is sp., Polyxenus sp.) ac i ely mo e in soil/li e and unde ba k, pi all ap-
ping is expec ed o miss ssome less mobile isopod species ha a e unde ep esen ed in
he p esen samples. Tu (2003) has cap u ed he isopod T. pusillus wi h pi all aps
in good numbe s, while Vilisics e al. (2007) epo ed only a ew cap u ed specimens
in he mids o a dense T. pusillus popula ion wi h he same me hod. We he e o e
suppose he pi all apping can be use ul o assess species di e si y and occu ence,
while cau ion emains o abundance da a o some species. The da a p esen ed he e
a e compa able wi h o he s udies ha collec ed isopods and millipedes wi h he same
Fe enc Vilisics e al. / ZooKeys 176: 199–214 (2012)
214
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