Characteristics of centipede (Chilopoda) assemblies in Dinaric frost hollows 
in Velika gora (Slovenia)
Značilnosti združbe strig (Chilopoda) v dinarskih mraziščih na Veliki gori 
(Slovenija)
Branka Vode, Ivan Kos*
Biotechnical Faculty, University of Ljubljana, Večna pot 111, SI-1001 Ljubljana, Slovenia
*correspondence: ivan.kos@bf.uni-lj.si
Abstract: Current study is preliminary research of centipedes (Chilopoda) assem-
blies in Slovenian frost hollows. With two sampling methods in three series (June, 
August and November) in year 2010, 834 individuals were caught and indentified to 
30 species. Most collected species had middle Europe distribution, 6 species were 
endemic for southeast Alps and northwest Balkan region, 3 of them are probably new 
for science. Five species had paleartic distribution. Maximum number of species found 
in one location was 28. Average density of centipedes was from 218 ind./m2 to 552 
ind./m2. The alfa diversity of the centipede assemblies is medium compared to other 
locations in Dinaric part of Slovenia and evidently higher compare to other European 
forest. The same applies for average density and species richness. Overall centipede 
assemblies were more similar in one location during the season than to assemblies 
from different locations in the same month.
Keywords: Chilopoda, frost hollow, assemblies, seasonal characteristic, Dinaric 
region
Izvleček: Preučevali smo sezonsko dinamiko strig v nekaterih mraziščih na Veliki 
gori, J Slovenija. Z dvema vzorčevalnima metodama v treh vzorčenjih (junija, avgusta 
in novembra) v letu 2010 smo dobili 834 osebkov, ki smo jih uvrstili v 30 vrst. Največ 
vrst je imelo srednje evropsko razširjenost, kar 6 vrst je bilo endemnih za območje 
jugovzhodnih Alp ali severozahodnega Balkana, od tega so 3 vrste najverjetneje nove za 
znanost. Največje število dobljenih vrst v enem mrazišču je bilo 28. Povprečna gostota 
osebkov na posameznem vzorčenju je bila od 218 os./m2 do 552 os./m2. Glede na alfa 
diverziteto združbi strig v mraziščih sodita med srednje bogate na območju Slovenije 
in med zelo bogate v primerjavi z drugimi evropskimi gozdnimi združbami. Enako 
velja za vrstno bogastvo in povprečno gostoto. Združbe strig so si bile bolj podobne 
na posamezni lokaciji skozi sezono, kot združbe različnih lokacij v istem mesecu.
Ključne besede: strige, mrazišče, združba, sezonske značilnosti, dinarsko območje
 ACTA BIOLOGICA SLOVENICA 
 LJUBLJANA 2014              Vol. 57, [t. 1: 45–58
Introduction
According final revision of data (Ravnjak 
2012) there are 98 species of Chilopoda known 
in Slovenia, more than one third of them (35 spe-
cies) being endemic in southeast Alps or northwest 
Balkan (Kos 2001). But Slovenia area is not evenly 
studied. We have still very little information for 
46 Acta Biologica Slovenica, 57 (1), 2014
the southeast and northeast part of Slovenia, part 
of the southwest and surroundings of the Ljubljana 
basin (Ravnjak 2012). 
One of unique characteristic of Slovenia is also 
karst terrain and karst phenomena such as frost 
hollows. In Slovenia frost hollows are present in 
the Julian Alps, Karavanke and all Dinaric area. 
There were many topological and geological 
studies conducted in the hollows (Gams 1972, 
1974, Trošt 2008, Stepišnik 2006, 2010), but few 
biological studies. Authors who made studies 
of vegetation in frost hollows in Slovenia were 
Martinčič (1977), Zupančič (1980, 1999), Wraber 
M. (1969), Wraber T. (1963) and Zavadlav (1974). 
There were only few studies of the fauna in frost 
hollows, confined to the particular animal groups: 
Collembola (Červek 1967, 1968), Gastropoda (Bole 
1976) and Acarina (Tarman 1975). Chilopoda in 
frost hollows have been only partially sampled 
(Kos et al. unpublished) and collected data have 
not yet been analyzed. This survey is therefore 
first preliminary study of centipedes’ assemblies in 
frost hollows in Slovenia. Our expectations were 
as follows: (i) rare or new species for centipede 
fauna of Slovenia will be found, (ii) the majority of 
species found will be species with middle European 
and paleartic distribution, iii) species with boreal 
and alpine character will be present, iv) there will 
be season differences in densities of centipedes, 
v) individual species will have different densities 
during the season.
Materials and methods
Sampling took place in Velika gora hills in 
Dinaric region in southern of Slovenia, west 
of city Ribnica (Fig. 1). Bedrock in this area is 
mostly calcareous, dominated by limestone and 
dolomites of different ages. Area is mountainous 
but does not reach great heights, while the terrain 
is very diverse due to karst phenomena (Bole and 
Slapnik 1997). Dominant natural vegetation in 
the area is the fir – beech forest (Omphalodes - 
fagetum) (Bole and Zupančič 1992). First selected 
frost hollow was Smrekov žleb (GKX: 5061900, 
GKY: 5476404). Most numerous tree species in 
the valley are common spruce (Picea abies) and 
fir (Abies alba). The topmost layer of the soil 
in the frost hollow was acidic and covered with 
rich undergrowth. Second frost hollow was near 
Kragulovec (GKX: 5062821, GKY: 5474681). 
Common spruce was also most present tree spe-
cies there, ground were also covered with rich, 
acidophilic undergrowth. 
Sampling took place on the 1st of June, 17th of 
July and 17th of November in year 2010, between 
8 am and 15 pm. Nine (9) soil sample units (SU), 
representing one sample, on each location were 
collected with metal cylinder. The sample units 
dimensions was Ø21cm x 15cm (appr. 5.2 dm3 of 
soil). Both organic (litter) as well as the fermenta-
tion horizon were covered. Samples were taken 
randomly in single location, but minimum 10 
m apart. Samples were then placed on Tullgren 
funnels, modified so that the lower parts of fun-
Figure 1:  Left: The location of Velika gora plateau in south of Slovenia. Right: Frost hollows Kragulovec and 
Smrekov žleb.
Slika 1:  Levo: Položaj planote Velika gora v Sloveniji. Desno: Mrazišči Kragulovec in Smrekov žleb.
47Vode, Kos: Centipedes assemblies in frost hollows on Velika Gora
nels were in the dark and cooled. Animals were 
intercepted in vessels filled with ethylene glycol. 
The extraction lasted 20-30 days, depended of the 
humidity of the sample. We also collected animals 
under the dead wood plants material and stones. 
In single location we have searched for 20 – 30 
min in all area. Captured animals were directly 
put in 70% alcohol.
All statistical analysis was made following 
Krebs (1989). For estimation of species richness 
we used Jackknife for diversity Shanon-Wiener 
index and Evenness index. For calculation of 
similarity among centipede assemblies Percent-
age similarity (Renkonen index) was used (Krebs 
1989, p.304). For this we used only relative share 
of species (without juvenile lithobids and unde-
termined individuals) from each soil sampling. 
Dendrogram of similarity was made with program 
PAST (http://folk.uio.no/ohammer/past/) using 
minimum-linkage method of clustering. Centipedes 
were determined following published literature by 
Eason (1964), Koren (1986, 1992), Matic (1966, 
1972), Verhoef (1931, 1937) and other. All col-
lected material is kept in Biotehnical faculty in 
Department of biology.
Results
Review of the species
Order Geophilomorpha was recently fully 
reviewed and amended by Bonato and Minelli 
(2014). New species names are therefore used in 
the article, but to allow comparison with earlier 
data, the previous valid names and/or names that 
were used in past Slovenian literature are given 
in parentheses in list 1. For general distribution 
of species we followed Stoev (1997), Kos (2012) 
and data from Fauna europaea and Chilobase.
List 1:  A list of all taxa collected with hand sampling (*) and with soil sampling method (+) in both locations. 
Currently known distribution (C.D.): pa – paleartic, il – Illyric, en – endemic, Ev – european, miE - mid-
dle european, sE -  south european, seE- southeast european, eE – east european, Me –Mediterranean
Seznam 1: Seznam vseh taksonov. Vrste, ki smo jih dobili pri ročnem pobiranju * in vrste, ki smo jih dobili pri 
talnem vzorčenju +. Splošna razširjenost (C.D.): pa – palearktična, il – ilirska, en – endemit, Ev – 
evropska, miE – srednjeevropska, sE – južnoevropska, seE -  jugovzhodnoevropska, eE – vzhodno 
evropska, Me – mediteranska
C.D o. GEOPHILOMORPHA
f. Geophilidae Leach, 1815
ili Clinopodes carinthiacus (Latzel, 1880) + (Clinopodes trebevicensis (Verfoeff, 1898) )
En Eurygeophilus pinguis (Brölemann, 1898) + (Chalandea scherpeltzi Attems, 1952 )
miE Geophilus alpinus Meinert, 1870 + (Geophilus (Orinophilus) insculptus Attems, 1895)
En Geophilus n.sp. + (Geophilus (Orinophilus) n.sp. )
sE Stenotenia sorrentina (Attems, 1903) +
f.Linotaeniidae Cook, 1899
Ev Strigamia acuminata (Leach, 1815) +
miE Strigamia transsilvanica (Verhoeff, 1928) + *
f. Mecistocephalidae Bollman, 1893
ili Dicellophilus carniolensis (C.L.Koch), 1847 + *
f. Schendylidae Cook, 1896
Al-ka Schendyla tyrolensis (Meinert, 1870) + * (Schendyla montana Attems, 1895)
Ili Schendyla carniolensis (Verhoef, 1902) +
Pa Schendyla nemorensis (C. L. Koch, 1837) +
48 Acta Biologica Slovenica, 57 (1), 2014
o. LITHOBIOMORPHA
f. Lithobiidae
sE Eupolybothrus (Leptopolybothrus) tridentinus (Fanzago, 1874) *
En Harpolithobius cf. gottscheensis (Verhoeff, 1937) +
Harpolithobius sp. + *
Lithobius juv + *
Lithobius sp. +
Pa Lithobus borealis Meinert, 1868 *
Me Lithobius castaneus Newport, 1844 +
Pa Lithobius forficatus (Linnaeus, 1758) *
miE Lithobius latro Meinert, 1872 +
miE Lithobius nodulipes Latzel, 1880 + *
miE Lithobius pelidnus Haase, 1880 +
miE Lithobius pygmaeus Latzel, 1880 +
Ev Lithobius tenebrosus Meinert, 1872 + *
miE Lithobius punctulatus C.L. Koch, 1847 + * ( Lithobius validus Meinert, 1872)  
miE L. (Monotarsobius) aeruginosus C. L. Koch, 1862 +
En ? L. (M.) n.sp. + *
En L. (Sigibius) burzenlandicus carinthiacus (Koren, 1992) + *
En ? L. (Sigibius) n. sp. «anici« +
o. SCOLOPENDROMORPHA  
f. Cryptopidae
Pa Cryptops anomalans Newport, 1844 *
Pa Cryptos hortensis Leach, 1815 +
Ev Cryptos parisi Brolemann, 1920 + *
seE Cryptos rucneri Matic & Teodoreanu, 1966 +
All species that were found in this study have 
been already found in area of Slovenia. Four spe-
cies were found only by hand collecting method, 
17 species only with soil sampling method. Nine 
species were found with both sampling methods 
(List 1). Most collected species had middle Euro-
pean distribution (8), followed by endemic species 
(6) and paleartic species (5). We found 3 Illiric 
species and 3 species with european distribution 
(List 1). One Mediterranean (Lithobius castaneus), 
one alpine-karpatian (Schendyla tyrolensis) and 
one species with southeast distribution (Cryptops 
rucneri) were also found.
49Vode, Kos: Centipedes assemblies in frost hollows on Velika Gora
Table 1:   N- number of centipedes collected with hand collecting method.
Tabela 1:  N- število ulovljenih osebkov pri ročnem nabiranju. Vrste označene “odebeljeno”, so bile najdene le s  to 
 metodo 
Sampling date 1. 6. 2010 17. 8. 2010 17. 11. 2010
o. Species/Vrste Kragulovec Smrekov žleb Kragulovec
Smrekov 
žleb Kragulovec
Smrekov 
žleb
geo D. carniolensis 3   5  
geo S. tyrolensis 1        
geo S. transsilvanica   1   1  
lit E. tridentinus 1     5  
lit L. borealis       4  
lit L. burzenlandicus carinthiacus     1  
lit L. castaneus   1 1  
lit L. forficatus 1      
lit L. nodulipes   1  
lit L. pelidnus       1  
lit L. punctulatus   1 1 1
lit L. tenebrosus 1 2 1 1 1 17
lit L. (M.) n.sp.     1  
sco C. anomalans   3 1  
sco C. parisi   6 1 2  
lit Harpolithobius.sp.   1  
lit Lithobius juv   1    
N 3 6 14 19 10 17
Average density and species richness 
Juvenile Lithobiidae had the highest density 
in location Kragulovec in summer and autumn 
followed by species Lithobius pygmaeus, Clino-
podes carinthiacus and Cryptops parisi. The last 
two had highest densities in the spring and then 
progressively lower. Schendyla carniolensis, S. 
tyrolensis and Stenotaenia sorerentina on the other 
hand had the lowest densities in the summer and 
the highest in the autumn. 
In Smrekov žleb species with very high 
density in all three samplings were Lithobius 
(Sigibius) n.sp. »anici«, followed by L. pygmaeus 
and again C. parisi. Juvenile Lithobiidae also had 
very high density, but lower than in Kragulovec. 
In this location L. pygmaeus, Cryptops hortensis 
and C. parisi had highest densities in the summer. 
Opposite had Eurygeophilus pinguis, Lithobius 
(Monotarsobius) n. sp. and L. (S.) n. sp. ”anici” 
lowest densities in the summer. 
Average density of centipedes was from 218 
ind./m2 to 552 ind./m2 (Tab. 2,Tab. 3). In location 
Smrekov žleb the density was higher in all three 
samplings comparing to samplings in location 
Kragulovec. The number of species caught with 
square soil sampling method at one sampling was 
between 12 in 20 (Tab. 2,Tab. 3). Total number of 
species (soil sampling and hand collecting method 
in all three samplings) was 22 in location Kragu-
lovec and 28 in location Smrekov žleb.
Diversity index, evenness and Similarity of the 
assemblies
Shannon – Wiener diversity index was between 
1.9 and 2.47 and the Evenness index was between 
0.77 and 0.86 (Tab. 2, Tab. 3). In both locations the 
highest S-W index and Evenness was in August. 
Calculation of percentage similarity (Renkonen 
index) showed the greatest similarity between the 
centipedes assemblies in June and August on loca-
50 Acta Biologica Slovenica, 57 (1), 2014
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– 
W
2.
14
2.
15
1.
89
E
0.
77
0.
86
0.
76
51Vode, Kos: Centipedes assemblies in frost hollows on Velika Gora
Ta
bl
e 
3:
  
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ca
tio
n 
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re
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v 
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52 Acta Biologica Slovenica, 57 (1), 2014
tion Kragulovec (68.2%), followed by assemblies 
in Smrekov žleb in June and November (64.7%) 
(Tab.4). Overall centipede assemblies were more 
similar in one location during the season than 
assemblies from different locations in the same 
month (Fig. 2).
Table 4:  Matrix of similarity coefficients (Renkonen 
index) for all samples. K1 - Kragulovec 
1.6.2010, K2 - Kragulovec 17.8.2010, K3 – 
Kragulovec 17.11.2010; SmZ1 – Smrekov žleb 
1.6.2010, SmZ2 – Smrekov žleb 17.8.2010, 
SmZ3 – Smrekov žleb 17.11.2010
Tabela 4: Matrica podobnosti (Rankonenov indeks) med 
posameznimi vzorčenji. K1 – Kragulovec 
1.6.2010, K2 - Kragulovec 17.8.2010, K3 – 
Kragulovec 17.11.2010; SmZ1 – Smrekov žleb 
1.6.2010, SmZ2 – Smrekov žleb 17.8.2010, 
SmZ3 – Smrekov žleb 17.11.2010
 K1 K2 K3 SmZ1 SmZ2 SmZ3
K1      
K2 0,682      
K3 0,449 0,499     
SmZ1 0,427 0,361 0,248    
SmZ2 0,547 0,564 0,331 0,569   
SmZ3 0,487 0,423 0,407 0,647 0,582  
Discussion
Presentation of species
Many studies was made stating that for most 
representative faunistic studies combination of 
quantitative and qualitative sampling methods 
should be used (Kos, 1988a, Fründ et al. 1997, 
Leśniewska 2000, Grgič 2005). Much smaller 
number of individuals obtained with hand collect-
ing method is the result of lower effort compared 
with soil sampling method. Nevertheless accord-
ing to the given estimation of species richness 
repeated sampling would result in 2 (in location 
Kragulovec) to 6 (location Smrekov žleb) addi-
tional species at most. We can therefore conclude 
that our sampling was sufficiently intense and that 
most of the species present in frost hollows were 
captured. Below are in more detail presented 4 
species which are endemic to the southeast Alps 
or/and the northwest Balkan region, one species 
with alpine character and two species with boreal 
character (in alphabetical order).
Eurygeophilus pinguis Brolemann, 1898
In Slovenia there were 36 individuals found 
in the Dinaric and alpine region (Kos 1988a, Kos 
unpublished) and 28 more individuals in frost 
hollows (Vode & Kos this research). They were 
first determined as Chalandea scheerpeltzi Attems, 
1952, following Attems (1952) and Koren (1986) 
with reference to their physical characteristics 
and number of legs. All male individuals had 33 
and all female indivividuals had 35 pairs of leg 
namely. After the revision (Bonato et al. 2006) 
C. scheerpeltzi was put in the synonimics with 
E. pinguis.
Individuals of Geophilus (“Orinophilus”) 
n. sp. were found in Kočevska reka (Kos and 
Praprotnik 2000), Mala gora near Kočevje (Grgič 
2002), near Ribnica (Kos 1988a) in the valley Iška 
(Grgič 2005), in Cerkno (Pagon 2006) and in the 
0,96
0,88
0,8
0,72
0,64
0,56
0,48
0,4
K
3
K
2
K
1
S
m
Z2
S
m
Z1
S
m
Z3
S
im
ila
rit
y
Figure 2:  Dendrogram of similarity using Renkonen index 
of similarity) between locations (soil sampling 
method). Key: K1 - Kragulovec 1.6.2010, 
K2 - Kragulovec 17.8.2010, K3 – Kragulovec 
17.11.2010; SmZ1 – Smrekov žleb 1.6.2010, 
SmZ2 – Smrekov žleb 17.8.2010, SmZ3 – 
Smrekov žleb 17.11.2010
Slika 2:  Dendrogram podobnosti med posameznimi 
lokacijami (talni vzorci) po Renkonenovemu 
indeksu podobnosti. Legenda: K1 – Kragulovec 
1.6.2010, K2 - Kragulovec 17.8.2010, K3 – 
Kragulovec 17.11.2010; SmZ1 – Smrekov žleb 
1.6.2010, SmZ2 – Smrekov žleb 17.8.2010, 
SmZ3 – Smrekov žleb 17.11.2010
53Vode, Kos: Centipedes assemblies in frost hollows on Velika Gora
frost hollows near Velika gora (Vode & Kos, this 
research). Altogether 60 individuals were found 
and identification key for thee potentially new 
species is in preparation.
We found one individual of the species Har-
polithobius gottscheensis Verhoef, 1937. Till 
now several individuals were found in the Karst 
Edge (Kos 1990), Kočevski Rog (Kos & Grgič 
2001), valley Iška (Grgič 2005) and Boč plateau 
(Ravnjak 2006). According to current knowledge 
this species is rare in Slovenia and it is considered 
endemic in northern Dinarides and southeast Alps 
(Kos, 2012). 
Lithobius borealis Meinert, 1868 is paleartic 
species, common in boreal forests in northern 
Eurasian area. It was found sparsely in Slovenian 
Alps (Kos et al. 2000), valley Iška (Grgič 2005) 
and Dinaric region (Kos 1988a, 1995a, b; Vode 
& Kos this research). Altogether there have been 
only 17 individuals found in Slovenia so far.
Lithobius tenebrosus Meinert, 1872 is 
widespread in boreal forests in Northern and 
Central Europe and the Carpathian Mountains 
(Chilobase, Kos 1995b). With its unique metabo-
lism it is adapted to the environments with low 
temperature (Kos unpublished). It was found in 
alpine, sub alpine and Dinaric region of Slovenia 
(Kos 1988b, Kos et al. 2000, Pagon 2006, Grgič 
2002, 2005, respectively) almost exclusively with 
hand collecting method. Its colder preferences 
were confirmed with this study as it was most 
abundant species found in frost hollows with hand 
collecting method.
There were 38 individuals of the species Litho-
bius (Monotarsobius) n. sp. found in Kočevski 
Rog (Kos 1995b), Kočevje area (Kos and Praprot-
nik 2000), the valley Iška (Grgič 2005), Cerkno 
(Pagon 2006) and Kras (Kos et al. unpublished). 
In this study 23 more individuals were found, 
which is noticeable contribution to facilitate the 
description of this potentially new species.
Lithobius (Sigibius) n. sp. “anici” is new 
species for science and was fist found in Kočevje 
area, where it was one of the most abundant species 
(Kos 1988a). Later it was also found in Bosnia 
(Kos 1992), Slovenian Karst (Kos 1995a), in 
the valley Iška (Grgič 2005), Kočevje area (Kos 
in Praprotnik 2000, Vode & Kos this research). 
Altogether there were almost 800 individuals 
found which makes this species one of the most 
numerous species of Dinaric region in Slovenia.
Centipede assemblies’ characteristics
Comparing to other locations in Dinaric region 
in Slovenia (where densities and species number 
were as high as 44 species and 892 ind./m2 (Grgič 
2005)) average density of centipedes from 218 ind./
m2 to 552 ind./m2 and number of species 22 to 28 
defines centipede assemblies in frost hollows as 
medium rich. But when comparing to forest as-
semblies in Europe can be defined as (very) rich. 
In Poland for instance Leśniewska (2000) found 
up to 19 species with average density 181 ind./m2 
in Querco-Carpinetum forest with two sampling 
methods. Tuf (2000) found 2 - 9 species (max. aver. 
density 250 ind./m2 with soil sampling method) in 
different successional stages of Querco-Ulmetum 
association and Wytwer (2000) found 6 – 9 species 
(max. aver. density 40 ind/m2 with Barber’pitfall 
traps) in 5 different forests types. Studies made 
in Germany by Albert (1982) and Fründ (1987) 
also reviled smaller numbers of species (4 – 9) 
and lower densities (up to 171 ind/m2) as have 
studies in England (Roberts 1957: 7 species, 
150 ind./m2) and in Hungary (Loksa, 1979: 7-9 
species, up to 234 ind./m2). Jet we must point out 
that all densities made by Slovenian authors were 
determined on the basis of sampling the unoc-
cupied (“free”) surface. The data are therefore 
not fully compatible with other authors who used 
different combination of sampling methods and 
statistical analysis.
Seasonal depended centipede assemblies 
changes
According to the temperature of the soil and 
air in the frost hollows made in previous studies 
(see “Introduction”) we presume that evaporation 
in the frost hollows is smaller and soil retains 
more moisture than in the surroundings. Knowing 
that centipedes actively choose the most optimal 
locations available (Grgič and Kos 2001) and that 
they are capable of relatively long distance migra-
54 Acta Biologica Slovenica, 57 (1), 2014
tion (referring to larger epidaphic species) (Kos 
1995a, Grgič 2005) we were anticipating there 
will be seasonal migration to frost hollows from 
the surrounding area in the summer, meaning that 
centipede densities in the frost hollows would be 
higher in the summer months. Our results reject 
this theory as the average centipede density was in 
the summer in both locations lower than in spring 
or in autumn. Our presumptions are therefore that 
in summer months when the temperatures were 
higher, larger more mobile species migrated out 
of frost hollows, while smaller less mobile species 
moved deeper in the soil. The disclosure of reasons 
for this migration as well as the confirmation of 
our migrational theory still have to be done. 
Diversity, evenness and similarity 
Although the average densities were in the 
summer the lowest, the S – W diversity index 
and the evenness index were in the summer the 
highest in both frost hollows. Interpretation of the 
dominance tells us why. In every sampling 2 - 3 
species and juvenile Lithobiidae had very high 
dominance. In location Kragulovec they were 
presenting 57 – 77% of all captured individuals. 
In spring species Clinopodes carinthiacus had 
24.8% dominance. Increased evenness rate in 
summer in Kragulovec can be attributed to more 
balanced proportions of species present while C. 
carinthiacus dominance declined to 10.3%. Again 
lower evenness rate and S – W index in autumn 
are due to the increased dominance of Schendyla 
carniolensis, Stenotaenia sorrentina and Lithobius 
pygmaeus. In location Smrekov žleb S – W diver-
sity index and evenness rate were higher compar-
ing to Kragulovec as three most dominant taxa 
represent “only” 48 – 49% of all species found. In 
this location Lithobius (Sigibius) n.sp. “anici” was 
most dominant species in all samplings followed 
by one more species (C. parisi in the spring, L. 
pygmaeus in the summer and S. tyrolensis in the 
autumn) and juvenile Lithobiidae. Whereas the 
total dominance of the 3 most dominant species 
did not change during the season so did not the 
S – W and evenness index. 
Dendrogram of similarity show distinct dif-
ference between frost hollows. The assemblies 
were more similar in the same location during the 
season, than to assemblies in the same month in 
other location. We assume that species composition 
of centipede assembly therefore depends more 
with respect to the surroundings rather than by 
environmental conditions in the frost hollows.
Conclusions
According to results we can confirm all hy-
potheses. We found 6 species that are endemic 
for northeren Dinarides or southeasteren Alps, 
3 of them are probably new to science. Spe-
cies with middle European distribution were 
far most numerous frost hollows, followed by 
species with palearctic distribution. We found 
Eurygeophilus pinguis which is alpine species 
and two species with boreal character: Lithobius 
borealis and Lithobius tenebrosus, last being the 
most numerous species obtained with hand col-
lecting method. There were statistical differences 
in species composition and densities during the 
season. From the results we predict that there is 
an active season migration between frost hollows 
and surroundings. We assume that frost hollows 
in Dinaric region are “refugia” of local popula-
tions of certain species thus having an important 
role when concerning maintaining population 
viability and biodiversity of the area. Due to this 
features we believe that frost hollows are suitable 
sampling sites for monitoring of impact of climatic 
changes on centipedes. To confirm our theories 
more comprehensive seasonal sampling with ad-
ditional sampling methods should be carried out. 
Povzetek
Namen raziskave je bil ugotoviti vrstno sestavo 
in gostoto strig v mraziščih, ter kako se tekom 
vegetacijske sezone spreminjata. Predvidevali smo, 
da bomo našli redke ali nove vrste za slovensko 
favno strig, da bomo poleg vrst s srednjeevrop-
skim in palearktičnim arealom razširjenosti našli 
vrste z alpskim in borealnim značajem, kar smo 
potrdili. Našli smo 6 endemnih vrst, od katerega 
so 3 najverjetneje nove za znanost, vrsto Euryge-
ophilus pinguis,ki ima alpski značaj in dve vrsti 
z borealnim značajem, Lithobius tenebrosus in 
Lithobius borealis. Skupno smo dobili 22 in 28 
vrst v posameznem mrazišču, povprečna gostota 
55Vode, Kos: Centipedes assemblies in frost hollows on Velika Gora
je bila od 218 os./m2 do 552 os./m2, kar uvršča 
združbo strig v mraziščih med srednje bogate 
združbe v primerjavi z drugimi lokacijami v 
slovenski dinarski regiji oziroma med zelo bogate 
v primerjavi z drugimi evropskimi gozdnimi 
lokacijami. Povprečna gostota je bila poleti na 
obeh lokacijah najmanjša, kar je v nasprotju z 
našimi predvidevanji, da strige poleti migrirajo v 
mrazišča zaradi nižjih temperatur in posledično 
večje vlažnosti. Zato predvidevamo, da se strige 
poleti ali umaknejo ven iz mrazišč ali pa se po-
maknejo globlje v tla, kar pa s to študijo nismo 
uspeli potrditi. Prav tako ostaja nepojasnjeno, kateri 
parametri so glavni razlog za sezonske spremembe 
v združbi strig v mraziščih, saj temperatura sama 
očitno nima glavnega vpliva. Vrstna pestrost v 
izbranih mraziščih je primerljiva z vrstno pestrostjo 
dinarskih bukovo-jelovih gozdov in je višja od 
mezofilnih gozdov srednje in severne Evrope. V 
obeh mraziščih sta bila vrednost Shannon – Wie-
nerjevega indeksa ter indeksa stalnosti najvišja v 
avgustovskem vzorčenju. To lahko razložimo s 
stopnjo dominance treh najštevilčnejših taksonov. 
Poleti so bile vrste bolj enakomerno zastopane 
kot spomladi ali jeseni, ko so dominirali juve-
nilni litobidi in vrste Clinopodes carinthiacus, 
Lithobius pygmeus, Schendyla carniolensis in 
Stenoteania sorrentina v lokaciji Kragulovec, 
oziroma juvenilni litobidi in vrste Schendyla 
tyrolensis, Lithobius (Sigibius) n.sp. »anici« in 
Cryptops parisi v lokaciji Smrekov žleb. Ocena 
podobnosti po Renkonenu je pokazala, da so bile 
združbe strig v posameznem mrazišču tekom 
sezone bolj podobne med sabo, kot z združbo v 
drugem mrazišču vzorčeno v istem obdobju, kar 
pomeni, da je vrstna sestava združbe strig bolj 
odvisna od vrstne sestave okolice, kot od okoljskih 
značilnosti mrazišča. Ta preliminarna raziskava 
združb strig v mraziščih utemeljuje potrebo po 
sezonskem vzorčenju z dodatnimi metodami, da 
bomo lahko potrdili naše domneve glede sezonske 
migracije strig med mraziščem in okolico in o 
refugialnem pomenu mrazišč za določene vrste. 
Že sedaj pa lahko predvidevamo, da so mrazišča 
primerna vzorčna mesta za spremljanje vplivov 
klimatskih sprememb na združbe strig. 
Acknowledgements
This work was supported by Slovenian Re-
search Agency by different projects or program 
such as “CRP project V4-0497 (C): Spatial and 
population dynamics of wildlife in Slovenian 
forests as a consequence of climate changes”, or 
program “P1-0184 (D): Investigations in zoology 
and speleobiology«. Authors would like to thank 
Franci Kljun for irreplaceable help with the field 
and laboratory work and Blanka Ravnjak for 
guidance with statistical analysis of data. We are 
also in debt to Ivan H. Tuf for his many construc-
tive remarks and improvements of our original 
manuscript.
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