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SUMMARY: The aim of this study was to investigate responses of ant and
carabid assemblages and their morpho-ecological traits to habitat differences
within natural temperate forests in Medvednica Nature Park. Toquantify habitat
differences in examined areas, both structural heterogeneity of the vegetation
and taxonomic diversity of plants were measured on six plots.

Habitat complexity was quantified using four habitat characteristics within
the site: tree canopy cover; shrub canopy cover; ground herbs and leaf litter
cover. Ants and carabids were sampled using pitfall traps.

Ant species richness and abundance, unlike carabid species richness were
positively correlated with habitat complexity, especially with leaf litter cover on
plots. The responses of insects morpho-ecological traits to habitat were recorded,
with more large bodied carabids present in more complex site and higher
abundance of opportunist ant species in more open sites with low complexity of
vegetation. Higher dominance of certain carabid species at the lower plots then
those on the top of the mountain, suggest competitive exclusion, confirming
lower areas as more stable. Species adapted to colder climate, that inhabit higher
elevations such as flightless forest specialist Cychrus caraboidesandCara-
bus irregularis, and boreo-montane ant species Camponotus herculeanus, are
less competent to colonize lower areas. Furthermore, they may not survive severe
instability of their habitats, especially in a changing climate. Overall results
suggest that conservation issues need to be focused on preserving stability
and structural complexity of forest habitat in summit areas of the mountain.

Key words: biodiversity, vegetation structure, litter, altitude, nature
conservation, forest habitat


Mountain landscapes usually offer steep gradients perature, precipitation, etc.), what is reflected in the
for a wide range of environmental parameters (i.e. tem-adaptation and distribution of different types of organisms
and ecological communities that occupy different


Dr. sc.Lucija Šerić Jelaska, dipl. prof. Ana Ješovnik,
prof. dr. sc. Mladen Kučinić, prof. dr. sc. Paula Durbešić,

positions along these gradients (review inHodkinson
Division of Biology (*Group for Systematic Zoology & Entomo

2005). Many of these environmental factors are multifa

logy), Faculty of Science, University of Zagreb, Rooseveltov trg 6,

ceted and interlinked in defining overall structural com

10000 Zagreb, Croatia, Phone: +385 1 4877711;
Fax: +385 1 4826260, e-mail:

plexity of insect habitats, which Hodkinson (2005)


Doc. dr. sc. Sven D. Jelaska, Division of Biology (*Group

has found to decrease with increasing altitude.The latter

forTerrestrial biodiversity), Faculty of Science, University

affects not only the species richness but also the species

of Zagreb, Marulićev trg 20, 10000 Zagreb, Croatia


composition of insect communities (Whittaker 1952

Mr. sc. Aljoša Pirnat, Groharjeva 18, SI-1241 Kamnik, Slovenia

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in Hodkinson 2005). The physical structure of the

environment, mediated by plant communities, can inf

luence the distribution and interactions of species (review
in Lawton 1983). “Structural heterogeneity
hypothesis” assumes that structurally complex habitats
may provide more niches and environmental resources
for exploitation and thus increase species diversity, although
empirical support for this relationship is biased
towards studies of vertebrates and habitats under anthropogenic
influence (Tews et al. 2004). Lassau and
Hochuli (2004) and Lassau et al. (2005) showed
that habitat complexity in forests may affect the composition
of ant and beetles assemblages.Brose (2003b)
showed that effects of habitat heterogeneity on ground
beetle assemblages were positive on the micro- and


meso-scale (0.25 and 500–1000 m, respectively), and


were not significant on a macro-scale (10 km).
In “taxonomic diversity hypothesis” plant taxonomic
diversity is positively correlated with the diversity of
herbivore insects (Murdoch etal. 1972,Root1973),
and thus with predator diversity (Hunter and Price
1992). The importance of floristic richness for the
ground beetle abundance was also reported by Baguette(
Ants and ground beetles have been widely recommended
as environmental, ecological and biodiversity
indicators (e.g. Altegrim et al. 1997, Andersen
1997, Niemelä 2000, Szyszko et al. 2000, Andersen
et al. 2002,Antonova and Penev 2006,
Pearce and Venier 2006, Šerić Jelaska et al.
2007,Šerić Jelaska and Durbešić 2009). They
respond well to natural and anthropogenic disturbances.
Microhabitat characteristics, i.e. litter type, organic
matter content, insolation, temperature fluctuation,
are important for providing hunting, foraging niches
and for protection of beetles from predators, ovipositioning,
larval development, over wintering etc.

(Thiele 1977,Pearce and Venier 2006). Theuse
of carabids morpho-ecological traits like wing morphology,
body size or diet are recommended in the evaluation
of habitat quality (Blake etal. 1994,Gutiérrez
et al. 2004,Gobbi andFontaneto 2008).Brose
(2003a) found that large carabids, as more preferable
prey because of their higher nutritive value and reduced
foraging time required, prefer dense vegetation
plots as enemy-free spaces.According toAndersen
et al. (2002), ant functional groups can be used as
indicators of habitat quality and forest health under different
management (Stephens and Wagner2006).

Lassau andHochuli (2004) suggested measurement
of habitat complexity as a surrogate for the diversity
of a range of arthropods including ants.Very often,
different taxonomic groups indicate certain areas as important
for protection or nature conservation, showing
mutual positive correlations. Based on the existence of
these correlations, the use of “surrogate species groups”
for estimation of overall biodiversity has been reported
(Garson etal. 2002;Satersdal etal. 2003); being
particularly useful where limited biodiversity data exist.
Vascular plants have been reported bySatersdal et
al. (2003) as a good surrogate group, which also works
for estimation of ground beetle diversity. Although,
some habitat preferences are known for many species of
ants and ground beetles, it is not always evident which
environmental factors are the most responsible for species
assemblage and distribution.

The aim of this study was to analyze ants and carabid
species richness and abundance, carabids body size
and ants functional groups, in different habitats within
natural temperate mature forests, and to test whether
higher habitat complexity and plant species richness
support higher insect diversity along vertical gradient
in Mt. Medvednica.

MATERIALAND METHODS – Materijal i metode

Study Area – Područje istraživanja

Mount Medvednica is part of the Croatian continental
karst, located north of Zagreb, the capital of Croatia.
The great floristic diversity (Dobrović et al. 2006),
due to the mosaic structure of forest communities, geographical
position and geological structure make this
area very interesting for biodiversity investigations.
Dobrović etal. (2006) reported that the floristic richness
and diversity of the mountain was higher than similar
regions in Croatia and several European countries
(Italy,Austria, Slovakia, Poland and Serbia). For protection
of its valuable forest areas, the western part of Mt.


Medvednica (228 km) was proclaimed a nature park by
the Nature ProtectionAct in1981. Thehighestpeak of

the mountain, named Sljeme, is placed 1035 m above sea

level. Despite protection, anthropogenic pressures on

Med vednica are constantly increasing (road construction,
recreation, ski trails, logging, etc.), mainly at its
upper elevations, changing the habitat quality.This can
change structural complexity of forests, especially where
trees and shrubs form part of the natural landscape.

Surveyed forests are natural temperate ones on
brown acid soil with silicate parent rocks (Basch
1995). For this investigation, six plots (50 x 50 m) were
selected along the central profile of Mt. Medvednica
across a vertical gradient on both slopes of the mountain
(Figure 1).They were placed in mature stands of
five different forest communities, that account for 78%
of the total forest cover of the nature park:Querco–Castaneetum
sativaeHt. 1938, (plot 1);Luzulo–Fagetum
sylvaticaeMausel 1937, (plot 2);Lamio orvale–Fage

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Figure 1 Position of the study area - Medvednica Nature Park (black polygon) and position of investigated plots in the Park

Slika 1. Područje istraživanja i položaj istraživanih ploha na području Parka prirode Medvednica

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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486

tum sylvaticae Ht. 1938, (plot 3), Festuco drymeiae-sured. Because of its circular nature, the terrain aspect
Abietetumass. nova,Vukelić &Baričević2007, was transformed into a continuous north-south gradient
(plots 4 and 6);Chrysanthemo macrophylli–Aceretum (northness) by calculating the cosine of aspect values
pseudoplatani (Ht. 1938) Borh. 1962 (plot 5), (Table (Guisan etal. 1999,Jelaskaetal. 2003).
1). For each plot, altitude, slope and aspect were mea-

Table 1 Ecological variables and their values for six plots

Tablica 1.Vrijednosti ekoloških varijabli na šest istraživanih ploha

Variable/plot (50x50 m) 1 2 3 4 5 6
community/zajednica* qc lf lo af cm af
Organic matter content/udio org. tvari (%) 8.99 16.36 15.96 21.47 18.82 13.28
Number of plant species/br. biljnih vrsta 62 19 41 43 36 39
Altitude/visina (m) 400 550 660 970 810 660
Slope/nagib (o) 21 18 17 28 21 7
Aspect/aspekt (o) 145 272 102 325 354 335
Northness/sjevernost -0.819 0.035 -0.208 0.819 0.995 0.906
Canopy openness/otvorenost sklopa (%) 4.04 10.62 4.95 12.69 4.98 4.94
Leaf area index/indeks lisne površine 3.91 2.59 4.13 2.34 3.81 3.41
Soil type/tip tla Brown acid/kiselo smeđe

* qc- Querco- Castaneetumsativae, lf- Luzulo- Fagetumsylvaticae, lo- Lamioorvale- Fagetumsylvaticae,
af – Festuco drymeiae - Abietetum, cm- Chrysanthemomacrophylli–Aceretumpseudoplatani

Habitat complexity was quantified using four habitat used them in our analyses.Three characters (shrubs cavariables
within the site: tree canopy cover; shrubs ca-nopy cover; ground herb cover; amount of leaf litter)
nopy cover; ground herb cover and amount of leaf litter.

were visually quantified into four categories (from score
Each variable was scored with 0, 1, 2 and 3 using an

0 to score 3: low contribution to site complexity (score
ordinal scale, where increasing scores indicate greater

0), the highest contribution to site complexity (score 3).
habitat complexity.This is modified version of the tech-

Tree canopy cover was measured as Leaf Area Index
niques used byLassau andHochuli (2004, 2008).

(LAI) and classified into four categories with scores
Compared to theLassau andHochuli (2004) sco

from 0 to 3. Summing up the scores for all quantified
ring method, there were no differences in two additional

characters, habitat complexity range from 5 (lowest ha-
variables i.e. soil moisture and portion of logs, rocks and bitat complexity) to 10 (maximum habitat complexity)
debris between investigated plots, hence we haven’t per plot (Table 2).

Table 2 Ecological variables used to derive measures of habitat complexityand their values for six plots

Tablica 2.Ekološke varijable korištene za određivanje kompleksnosti structure staništa na šest istraživanih ploha

Variable/plot (50x50 m) 1 2 3 4 5 6
Tree canopy cover (LAI)/sklop krošnji (LAI) 3 (3.91) 1 (2.59) 3 (4.13) 1 (2.34) 3 (3.81) 2 (3.41)
Shrub canopy cover/pokrovnost grmlja 2 0 1 0 0 2
Ground flora cover/pokrovnost prizemnog bilja 2 3 2 3 3 1
Leaf litter cover/količina listinca 3 3 3 1 2 1
Habitat complexity scores/vrijednosti kompleksnosti staništa 10 7 9 5 8 6

In each of the six plots, the number of vascular
plants species was recorded on successive visits during
the season (March – October). Data on canopy openness
and LAI, as indirect estimators of available under
story light, were measured using hemispherical photographs
(as described inJelaska 2004 andJelaska
et al. 2006) and analyzed with Gap Light Analyzer
(GLA) software (Frazer etal. 1999).

Soil samples were collected twice during this investigation,
at the beginning of the season in March 2001
and in May 2002. Four samples of the top 10 cm of soil
were collected from each plot, after removal of the litter.
Samples were diagonally placed across the plot, at
14 m intervals following general scheme in Scholz et
al. 1994.To determine the percentage of organic matter,
10 g of soil was ashed at 400oC. All environmental
variables are displayed inTable 1.

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Carabid Beetles andAnts Sampling – Uzorkovanje trčaka i mrava

Carabid beetles and ants were collected in 16 pitfall
traps on each plot, placed in a regular rectangular net
with cell size of 10 x 10 meters placed five meters from
the edges of the plot.Altogether, 96 pitfall traps were exposed
through the investigation period.Traps were filled
with ethanol (96%), acetic acid (9%) and water in equal
proportions and emptied every two to three weeks from
March 2001 toApril 2002. Carabids and ants were identified
using specialized keys (Collingwood 1979,
Agosti and Collingwood 1987, Trautner and

Data Analyses –

The total abundance and species richness of ants and
carabids was determined for each habitat (i.e. plot).The
Shannon-Wiener and Sorensen indices (Krebs 1989)
were calculated for all plots to assess ground beetles and
ants’diversity and similarity using Programs for Ecological
methodologyVer.5.2 (15-III. 2000). Sorensen similarity
values between plots were used in cluster
analyses with the Euclidean distance as distance measure
and single linkage as linkage rule for constructing

Atotal of 9288 beetles and 2958 ants belonging to 43

and 20 species, respectively, were trapped (Table 3). The
number of captured species per plot varied from 17 to 27
for carabids, and from 5 to 14 for ants.Abax parallelepipedus,
Abax paralellus, Aptinus bombarda,Carabus
violaceusandCychrus attenuatuswere carabids found at
all surveyed plots (Table 3). Only one ant species Myr-
Geigenmüller 1987,Hůrka 1996,Freude et al.
2004 andSeifert 2007). AccordingtoAndersen et
al. (2002) ants were classified in four functional groups.
Based on personal measurements of carabid specimens
and data from Hůrka (1996) and Freude et al.
(2004), average body size for analyzed species were
classified into three categories: (1) small (4–8 mm); (2)
medium (8–21 mm); and (3) large (22–40 mm) using a
size distribution graph.

Analiza podataka

dendrograms using STATISTICA 6.1 (StatSoft Inc.
2003).Wecompare habitat complexity scores, plant species
richness and measured environmental variables
(soil organic matter, slope, aspect and elevation) with
data about richness and abundance of ants and carabids,
ant functional groups and carabids body size distribution
patches using Pearson Product Moment correlations in
STATISTICA6.1 software.

– Rezultati
mica ruginodiswas found at all plots, unlikeCrematogaster
schmidti,Camponotus herculeanus,Formica gagatesandFormica
rufarecorded at single plot (Table 3).

Ant species richness was negatively associated with carabid
richness per plot (r = -0.86;p= 0.029), opposite to
the trend for their abundance (r = 0.85; p=0.031).

Table 3 Carabid and ant species with number of specimens per plot. Plot number corresponds to those in Figure 1

Tablica 3.Vrste i brojnost jedinki trčaka (Carabid) i mrava (Ant) na istraživanim plohama.
Broj plohe odgovara onome na slici 1.

Carabid species/Plots 1 2 3 4 5 6
Abax carinatus(Duftschmid 1812) 156 23 2 1 26
Abax ovalis(Duftschmid 1812) 1 3 37 1
Abax parallelepipedus(Piller & Mitterpacher 1783) 654 89 328 5 61 50
Abax parallelus(Duftschmid 1812) 498 2 283 1 145 136
Agonum gracilipes(Duftschmid 1812 1
Platynus scrobiculatus(Fabricius 1801) 1 19
Amara aeneaDe Geer 1774 3 2 1 1
Aptinus bombarda (Illiger 1800) 1108 4 11 3 213 45
Bembidion lampros (Herbst 1784) 1
Bembidion deletumAudinet-Serville 1821 1
Carabus convexusFabricius 1775 445 343 362 1
Carabus coriaceusL. 1758 34 23 82 2 10
Carabus gigasCreutzer 1799 16 5 1
Carabus intricatusL. 1761 132 49 78 1 3
Carabus irregularis Fabricius 1792 72 85 12
Carabus nemoralisO. F.Müller 1764 636 379 52 15 6
Carabus praecellensPalliardi 1825 18 3 2 24 35
Carabus ulrichiiGermar 1824 548 29 100 52 8
Carabus violaceusL. 1758 23 168 43 29 35 63

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Cychrus attenuatus(Fabricius 1792) 43 221 70 44 99 43
Cychrus caraboides(Linnaeus 1758) 5 1
Dromius fenestratus (Fabricius 1794) 3
Harpalus atratusLatreille 1804 1
Harpalus marginellus Dejean 1829 2
Harpalus laevipesZetterstedt 1828 1
Harpalus dimidiatus(P. Rossi 1790) 1
Leistus piceusFrölich 1799 5 3 1
Leistus rufomarginatus (Duftschmid 1812) 3 1 1 62 67
Licinus hoffmannseggii(Panzer 1803) 2 23 83 3
Molops elatus(Fabricius 1801) 9 27 4
Molops piceus(Panzer 1793) 20 27 5 1 37
Nebria brevicollis (Fabricius 1792) 1

Notiophilus biguttatus(Fabricius 1779) 1 5 1
Notiophilus rufipesCurtis 1829 23 3 1
Platyderus rufus(Duftschmid 1812) 4 24
Pseudoophonus rufipes(De Geer 1774) 1 1
Pterostichus anthracinus (Illiger 1798) 1
Pterostichus fasciatopunctatus (Creutzer 1799) 2 11
Pterostichus oblongopunctatus (Fabricius 1787) 121 120
Pterostichus transversalis (Duftschmid 1812) 5 7 79
Stomis rostratus (Sturm in Duftschmid 1812) 1 1
Synuchus vivalis(Illiger 1798) 1 4 1
Trechus cardioderus Putzeys 1870 1

Number of species/broj vrsta 18 1721 2726

Number of specimens/broj jedinki
Ant species/Plots
4350 1346 1449 374 1117 631
Aphaenogaster subterranea(Latreille 1798)
2 3
4 5 6
Camponotus fallax(Nylander 1856) 2
Camponotus herculeanus(Linnaeus 1758) 2
Camponotus ligniperda(Latreille 1802) 11 144 4 7
Crematogaster schmidti (Mayr 1853) 26
Dolichoderus quadripunctatus(Linnaeus 1771) 8 1 1 1
Formica fuscaLinnaeus 1758 14 18 1
Formica gagatesLatreille 1798 22
Formica polyctenaFörster 1850 154 1
Formica rufaLinnaeus 1758 2
Lasius brunneus(Latreille 1798) 14 65 11 2 91
Lasius citrinusEmery 1922 1 1
Lasius emarginatus(Olivier 1792) 629 180 4 1 3
Lasius fuliginosus(Latreille 1798) 1 3 1
Myrmecina graminicola(Latreille 1802) 28 16
Myrmica ruginodisNylander 1846 50 244 127 199 54 156
Myrmica sabuletiMeinert 1861 4 2
Prenolepis nitens(Mayr 1853) 135 11
Stenamma debile(Förster 1850) 6 10 17 35
Temnothorax crassispinus(Karavajev 1926) 134 44 165 2 20
Number of species/broj vrsta 14 12 11 5 5 10

Number of specimens/broj jedinki 1126

No significant correlations were found between
plant richness and arthropod diversity. Habitat complexity
scores were positively correlated with carabid and
ant abundances and ant species richness, but negatively
correlated with carabid species richness (Table 4). Of

761 478 209 66 318

the four environmental variables assessed by scoring
method, shrub and leaf litter cover were positively correlated
with ant species richness and carabid abundance.
Large negative correlations were found between
carabid richness and litter cover (-0.90,p= 0.015).

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Figure 2
Tree diagram of cluster analyses using Sorensen indices based on carabid species composition in investigated
plots as similarity measure. Numbers on X axes denote investigated plots

Slika 2. Dendrogram sličnosti istraživanih ploha koristeći Sorensen indeks ovisno o prisutnosti vrsta trčaka tijekom
razdoblja uzorkovanja. Broj na osi X označava plohe

Tree diagram of cluster analyses using Sorensen indices based on ant species composition as similarity measure.
Numbers on X axes denote investigated plots

Slika 3. Dendrogram sličnosti istraživanih ploha koristeći Sorensen indeks ovisno o prisutnosti vrsta mrava tijekom
razdoblja uzorkovanja. Broj na osi X označava plohe

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Table 4 Correlation coefficients between ant and carabid assemblages and habitat variables

Tablica 4.Koeficijenti korelacije između sastava mrava i trčaka sa stanišnim čimbenicima

Habitat variables /stanišne varijable
Community features /osobine zoocenoza
score /
richness /
Altitude /
matter /
Carabid richness /broj vrsta trčaka
Carabid abundance /abundancija trčaka
Carabid Shannon indeks /Shannon indeks trčaka

There was no correlation between the heterogeneity
of vegetation structure and plant species richness. Habitat
complexity was negatively correlated with altitude
(r = -0.70;p=0.12) and with aspect (r = -0.77;p= 0.074).
The same trend was found for ant richness, abundance
and diversity indices for ants’ communities (Table 4).
Carabid species richness was positively correlated with
altitude and aspect, opposite to carabids abundance and
ant assemblages (Table 4). Large and small bodied species
showed opposite trends in their distribution across
investigated sites. Large carabids were positively cor related
with leaf litter cover (r= 0.81, p=0.05). Small
carabids positively correspond with higher altitude
(r= 0.89, p=0.018).

The most abundant carabids wereA. parallelepipedus,
A. paralellus, A. bombarda,C. nemoralisandC.
ullrichi(Table 3) respectively, accounting for 37.16%
of the total catch. The highest number of specimens,
comprising 47.11% of all specimens, was captured in
the most complex site (plot 1), though this plot recorded
almost the lowest number of species (i.e. 18, as
compared to the minimum of 17 on plot 2). On the contrary,
the highest number of carabid species (27) was
recorded on plot 4 (with the lowest habitat complexity
score) but with the lowest abundance making only 4%
of captured specimens.

The highest number of ant species (14) and abundance
(1126 specimens) was recorded on plot 1. The
lowest number of ant species and their abundance was
recorded on plots 4 and 5. 45% of them are woodland
species and only 10% of sampled species prefer open
habitats.The rest of them occur in both woodland and
open habitats.Analyzing the functional groups, according
toAndersen (1997), 65% of recorded species (13
species) are cold climate specialists (CCS), 3 species
are opportunists (O), 3 of them are subordinate camponotini
(SC) and 1 species belong to generalized Myrmecinae
(GM). Presence of species representatives of
all four groups were recorded on plot 1, only.M. ruginodis(
opportunist) that was found on all plots had the
highest abundance in the low complexity areas (plots 2
and 5).The abundance of opportunist ants was negatively
correlated with canopy cover (r= -0.91, p=0.05).
High correlations were found between ant abundance
and richness with the amount of organic matter in the
soil (Table 4). Organic matter content in the soil rise
with higher altitude (r= 0.9, p=0.012).

Cluster analyses of species composition on plots revealed
differences in carabid assemblages between the
southern (plots 1, 2, 3) and northern slopes (plots 4, 5, 6)
of Mt. Medvednica (Figure 2). Some species such asCarabus
convexus,Notiophilus rufipesandCarabus intricatuswere
found exclusively on the southern slope of
mountain, unlike Carabus irregularis,Notiophilus biguttatus,
Molops elatus,Platyderus rufusand all of the
collected species of the genusPterostichus, which were
recorded exclusively on northern slope of the mountain.
There was also a significant difference in number of collected
ants on southern and northern slopes (2365 on
south, 593 on north). Cluster analyses using Sorensen
indices based on ants’species composition, separate the
two highest plots, 4 and 5, from others (Figure 3).An ant
speciesC. herculeanusis recorded for plot 4, which is
970 m high and has northern exposure.C. herculeanus
has altitudinal range from 1000–1700 m, and boreal
mountain distribution, with a very restricted area. It prefers
coniferous mountain forests and can withstand very
low temperatures, 38.5 oC below zero (Seifert2007).

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The relationship between habitat features and diversity
of ants and carabid beetles was analyzed.We recorded
great differences in carabid species composition and
abundance among plots, where carabid abundance, but
not the richness, was significantly higher in habitats
with higher complexity scores.Brose (2003a) showed
that an experimental reduction of vegetation complexity
reduced the activity-abundance of large carabid species.
Our results supports “enemy-free space hypothesis”
(Lawton,1983) that prey species have more chances
of escaping from natural enemies in dense vegetations.
We found that large carabids prefer dense vegetation
plots covered with leaf litter as enemy-free space.

Furthermore, carabid species richness was negatively
correlated with ant species richness.The results of
ant richness analyses differ from those of Lassau and
Hochuli (2004), but not those analyzing functional
groups where they also found the largest abundance of
opportunist species in low complexity sites. Stephens
and Wagner (2006) have found that species
richness, diversity, and dominance were a less satisfactory
measure of various forest management impacts on
ants than functional group analysis.

The highest number of ant species (14) and abundance
(1126 specimens) was recorded on plot 1, and
the lowest number of the species and their abundance
on plots 4 and 5. This result was in accordance with
their habitat preferences.Ants in general, with excep

tion of a few cold-temperate species, are thermophilic

animals, and function poorly below 20 oC and not at all

below 10 oC (Hölldobler and Wilson, 1994).
Greater abundance and species number in lower altitudes
can be explained, in addition to temperature, with
precipitations, thickness and volume of leaf litter and
available food resources (Brühl etal. 1999).

The highest Shannon-Wiener index for carabid diversity
(Table 5) was recorded on plot 5, situated in the
Chrysanthemo macrophylli–Aceretum pseudoplatani
forest, with high concentration of soil organic matter as
a consequence of the longer persistence of snow-cover
and hence higher soil humidity and a shorter micro-organism
activity period.The herbaceous layer of this forest
is characterized by nitrophilous plant species e.g.
Lunaria rediviva L., Urtica dioicaL., Corydalis solida
(L.) Swartz, etc.The impact of leaf litter origin on carabids
has also been reported byNiemeläetal. (1992),
Koivula etal. (1999) andMagura etal. (2005), in
which the latter two authors observed significant impact
of leaf litter on certain species e.g. Pterostichus
oblongopunctatus and C. caraboides. In this study,
both above mentioned species were found only on plots
4 and 5, which had the highest soil organic matter content.
Soils in cooler climates commonly have more organic
matter because of slower decomposition rate

Table 5
Shannon-Wiener (H’) indices, numbers of equally common species (N) and Smith & Wilson evenness
(S&W) for investigated plots

Tablica 5.Shannon-Wiener (H’) indeksi raznolikosti, broj zajedničkih vrsta (N) te Smith & Wilson jednolikost
(S&W) trčaka (carab.) i mrava (ants) na istraživanim plohama

Plots /
carab. ants
carab. ants
carab. ants
carab. ants
carab. ants
carab. ants
H’ 3.049 2.262 2.724 2.585 2.951 2.074 3.201 0.366 3.711 0.977 3.625 1.936
N 8.27 4.80 6.61 6 7.73 4.21 9.19 1.29 13.10 1.97 12.34 3.83
S&W 0.138 0.26 0.153 0.39 0.128 0.322 0.291 0.22 0.182 0.293 0.218 0.295

On plots 1, 2 and 3, smaller number of species was
recorded, while plots 4, 5 and 6 had a higher number of
species with a low abundance, indicating less structured
arthropods communities without dominant species.
Also, cluster analyses of species composition on six
plots divided carabids based on aspect and ants species
based on elevation of investigated plots.

Ant and carabid diversity and species body size distribution
highly correlate with altitude and aspect (expressed
as northness). On warmer, more south exposed
plots (plots 1, 2 and 3) smaller number of carabid species
was recorded. On the contrary, plots on northern
slopes (plots 4, 5 and 6) had a higher number of species
with a smaller number of specimens. Salgado et al.
(1997) reported similar findings in their research of deciduous
oak forests, where 10 of 42 species represented
90% of the total catch on five plots. They recorded a
smaller number of species on plots with higher abundance
of specimens and those with smaller abundance
and larger number of species, the latter having unstable
climatic conditions. In this study, plot 1, with recorded
capture of 47.11% of the total catch, is situated at the lowest
altitude with southern exposure, thereby ensuring
more stable and warmer climate conditions that enable
formation of ground beetle communities with the dominant
and co-dominant species present (e.g. A. bombarda,
A. parallelus, A. parallelepipedus, C. ullrichi, C.
nemoralis, C. convexusandC. intricatus; Table 3). The
highest number of carabid species, but the lowest abundance
was recorded at the highest altitude (plot 4) with
the lowest habitat complexity score. Flightless forest
specialist such as C. caraboides that preferred higher

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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486

elevations and lower temperatures andC. irregularisare
distributed on top of the mountain with a small number
of individuals on the northern plots. Geomorphologic
and climatic conditions, large habitat complexity and
very low disturbances in plot 1, contribute in forming
more stable carabid and ant communities. Community
composition can be used to indicate broader aspects of
habitat quality and more general changes (i.e. degradation
and recovery following stress or disturbances),
(Hodkinson andJackson 2005). High abundance
of carabids but low richness at the warmer plots could
suggest competitive exclusion confirming that habitat
stability may have unimodal effect on richness. The intermediate
disturbance hypothesis predicts that diversity
will be greatest when physical disturbances prevent
competitively dominant species from excluding other
species from the community.At the low level of disturbances,
diversity is low because only the best competitors
persist and competitive exclusion leads to species
loss at either end of the disturbance continuum (Death
andWinterbourn 1995).Stephens andWagner
(2006) found that different ant functional groups were
dominant under different levels of disturbance intensity.

Data about species richness and habitat complexity
within undisturbed forest systems with low direct anthropogenic
impacts can be used as reference data for environmental
monitoring of changes in temperate forests.
Knowledge on how management of forests relates to forest
carabids and ants diversity is poorly documented in
Croatia.Wefound that ant functional groups and carabids
body size analyses respond well to differences in
habitat complexity.These results confirm the need for
sustainable forest management that will preserve higher
level of habitat complexity that provide more niches and
environmental resources for exploitation and thus support
dominance of larger carabids and high animal biomass.
Furthermore, carabids and ants may serve as
target groups in climate change risks assesment in
mountain ecosystems.

Grateful thanks to Gregor Bračko for helping us in tian Ministry of Science, Education and Sports (Grants
species determination.This study was financed by Croa-0119-123, 119-1193080-1206 and 119-0000000-3169).

REFERENCES – Literatura

Agosti,D., and C.A.Collingwood,1987: Aprovisional
list of the Balkan ants with a list to the
worker caste. II. Key to the worker caste, including
the European species without the Iberian.
Mitteilungen der Schweizerischen Entomologischen
Gesellschaft 60:193–261.

Altegrim,O., K.Sjöberg,and J. P.Ball,1997:
Forestry effects on a boreal ground beetle community
in spring: Selective logging and clear-
cutting compared. Entomol. Fennica 8: 19–26.

Andersen, A.N., 1997: Using ants as bioindicators:
Multiscale issues in ant community ecology.
Conserv. Ecol. (online) 1 (1): 8


Andersen, A. N., B. D. Hoffmann, J. Müller,
and A. D. Griffiths, 2002: Using ants as
bioindicators in land management: simplifying
assessment of ant community responses. J.Appl.
Ecol. 39: 8–17.

Antonova,V.,and Ly.Penev,2006: Change in the
zoogeographical structure of ants (Hymenoptera:
Formicidae) caused by urban pressure in
the Sofia region (Bulgaria). Myrmecologische
Nachrichten 8: 271–276.

Basch,O., 1995: Geološka karta Medvednice (Geological
map of Medvednica), 1:62500. Institut za
geološka istraživanja (In: Geološki vodić Medvednice,
Ed. Šikić K.). Institut za geološka
istraživanja, Zagreb, INANaftaplin, Zagreb.

Baguette, M., 1993: Habitat selection of carabid
beetles in deciduous woodlands of southern Belgium.
Pedobiologia 37: 365–387.

Blake,S., G. N. Foster, M. D. Eyre,and M. L.
Luff, 1994: Effects of habitat type and grassland
management practices on the body size
distribution of carabid beetles. Pedobiologia 38
(6): 502–512.

Bot,A., and J. Benites, 2005: The importance of
soil organic matter: Key to drought-resistant soil
and sustained food production. FAO Soils Bulletin
80 - URL: http://www.

Brose, U., 2003a: Bottom-up control of carabid
beetle communities in early successional wetlands:
mediated by vegetation structure or plant
diversity? Oecologia 135: 407–413.
Brose, U., 2003b: Regional diversity of temporary
wetland Carabid beetle communities: a matter of
landscape features or cultivation intensity?Agr.
Ecosyst. Environ. 98: 163–167.
Brühl, C.A., M. Mohamed,and K. E. Linsenmair,
1999:Altitudinal distribution of leaf litter
ants along a transect in primary forest on Mount
Kinabalu, Sabah, Malaysia. J. Trop. Ecol. 15:
Collingwood, C.A., 1979:The Formicidae (Hymenoptera)
of Fennoscandia and Denmark. Fauna
Entomologica Scandinavica 8: 9–175.

ŠUMARSKI LIST 9-10/2010 str. 47     <-- 47 -->        PDF

L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486

Death, R.G., and M. J. Winterbourn,1995: Diversity
patterns in stream benthic invertebrate
communities: the influence of habitat stability.
Ecology 76 (5): 1446–1460.

Dobrović,I., T.Nikolić, S. D.Jelaska, M.Plazibat,
V.Hršak,and R.Šoštarić,2006: An
evaluation of floristic diversity in Medvednica
Nature Park (northwestern Croatia). Plant Biosystems.
140 (3): 234–244.

Frazer, G. W., C. D. Canham, and K. P. Lertzman,
1999: Gap LightAnalyzer (GLA),Version
2.0. Simon Fraser University, Burnaby,
British Columbia, and the Institute of Ecosystem
Studies, Millbrook, NewYork, 36 pp.

Freude, H., K. W. Harde, G. A. Lohse, and B.
Klausnitzer,2004: Die Käfer Mitteleuropas,
Band 2.Elsevier, München, 521 pp.

Garson,J., A.Aggarwal,and S.Sarkar,2002:
Birds as surrogates for biodiversity: an analysis
of a data set from southern Quebec. J. Biosci. 27

(4)Suppl. 2: 347–360.
Gobbi,M., and D.Fontaneto,2008: Biodiversity
of ground beetles (Coleoptera: Carabidae) in different
habitats of the Italian Po lowland. Agr.
Ecosyst. Environ. 127: 273–276.
Guisan,A., S. B. Weiss,andA. D. Weiss,1999:
GLM versus CCA spatial modelling of plant
species distribution. Plant Ecol. 143: 107–122.
Gutiérrez,D., R.Menéndez,and M.Méndez,
2004: Habitat-based conservation priorities for
carabid beetles within the Picos de Europa National
Park, northern Spain. Biol. Conserv. 115:
Hodkinson, I. D., 2005: Terrestrial insects along
elevation gradients: species and community responses
to altitude. Biol. Rev. 80: 489–513.
Hodkinson, I.D., and J. K.Jackson,2005: Terrestrial
and aquatic invertebrates as bioindicators
for environmental monitoring, with particular
reference to mountain ecosystems. Environ.
Manag. 35 (5): 649–666.
Hölldobler,B., and E. O. Wilson,1994: Journey
to the ants.The Belknap Press of Harvard University
Press. Cambridge, Massachusetts, and
London, England, pp 191–204.
Hunter, M.D., and P.W.Price,1992: Playing chutes
and ladders: heterogeneity and the relative
roles of bottom-up and top-down forces in natural
communities. Ecology73: 724–732.
Hůrka,K., 1996: Carabidae of the Czech and Slovak
Republics. Kabourek, Zlin, 565 pp.
Jelaska, S.D., 2004:Analysis of canopy closure in the

Omphalodo-Fagetumforests in Croatia using hemispherical
photography. Hacquetia3 (2): 43–49.

Jelaska, S. D., O. Antonić, T. Nikolić, V.
Hršak, M.Plazibat,and J.Križan,2003:
Estimating plant species occurrence in MTB/64
quadrants as a function of DEM-based variables

– a case study for Medvednica Nature Park,
Croatia. Ecol. Model. 170 (2–3): 333–343.

Jelaska, S. D., O. Antonić, M. Božić, J. Križan,
and V.Kušan,2006: Responses of forest
herbs to available understory light measured
with hemispherical photographs in silver fir –
beech forest in Croatia. Ecol. Model. 194 (1–3):

Koivula,M., P.Punttila,Y.Haila,and J.Niemelä,
1999: Leaf litter and the small-scale distribution
of carabid beetles (Coleoptera,
Carabidae) in the boreal forest. Ecography 22:

Krebs, C.J., 1989: Ecological methodology. Harper
and Row, New York, 654 pp.

Lassau, S.A., and D. F.Hochuli,2004: Effects of
habitat complexity on ant assemblages. Ecography
27: 157–164.

Lassau, S.A., and D. F. Hochuli, 2008: Testing
predictions of beetle community patterns derived
empirically using remote sensing. Diversity
and Distribution. 14: 138–147.

Lassau, S. A., D. F. Hochuli, G. Cassis, and

C.A.M.Reid,2005: Effects of habitat complexity
on forest beetles diversity: do functional
groups respond consistently? Diversity and Distribution.
11: 73–82.

Lawton, J.H., 1983: Plant architecture and the diversity
of phytophagous insects. Ann. Rev. Entomol.
28: 23–39.

Magura,T., B.Tóthmérész,and Z.Elek,2005:
Impacts of leaf-litter addition on carabids in a conifer
plantation. Biodivers. Conserv. 14: 475–491.

Murdoch,W.W., F. C. Evans,and C. H. Peterson,
1972: Diversity and pattern in plants and
insects. Ecology 53 (5): 819–829.

Niemelä,J., 2000: Biodiversity monitoring for decision-
making.Ann. Zool. Fenn. 37: 307–317.

Niemelä,J., Y.Haila, E. Halme,T. Pajunen,
and P.Punttila,1992: Small-scale heterogeneity
in the spatial distribution of carabid beetles
in the southern Finnish taiga. J. Biogeogr. 19:

Pearce, J.L., and L.A. Venier,2006:The use of
ground beetles (Coleoptera: Carabidae) and spiders
(Araneae) as bioindicators of sustainable
forest management:A review. Ecol. Indicat. 6:

Root, R.B., 1973: Organization of a plant-arthropod
association in simple and diverse habitats. The

ŠUMARSKI LIST 9-10/2010 str. 48     <-- 48 -->        PDF

L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486

Fauna of Collards (Brassica oleracea). Ecol. Szyszko, J., H. J. W. Vermuelen, M. Klimas-

Monogr. 43: 95–124. zewski,and A. Schwerk,2000: Mean Indi -
Salgado, J.M., J. F.Gallardo, I. S.Regina, and
vidual Biomass (MIB) ground

of beetles

(Carabidae) as an indicator of the state of the ention
of ground beetle communities in several oak
M. E.Rodriguez,1997: Ecosociological rela

vironment, pp. 289-294. In P. Brandmayr, G.
forests of western Spain (Coleoptera: Carabi-

Lövei,T.Zetto Brandmayr,A. Casale,A.Vigna
dae). Entomol. Gen. 22(1): 029–043.

Taglianti (eds.), Natural history and applied ecology
of carabid beetles. Pensoft publishers, Sofia,

Satersdal M., I. Gjerde, H. H. Blom, P. G.


Ihlen, E.W.Myrseth, R.Pommeresche,
ŠerićJelaska,L., P.Durbešić,2009: Comparison

J.Skartveit,T.Solhoy,and O.Aaas,2003.
of the body size and wing form of carabid species

Vascular plants as a surrogate species group in
(Coleoptera: Carabidae) between isolated and

complementary site selection for bryophytes, ma-
continuous forest habitats.Annales de la Société

crolichens, spiders, carabids, staphylinids, snails,
Entomologique de France 45 (3): 327–338.

and wood living polypore fungi in a northern forest.
Biol. Conserv. 115: 21–31.

ŠerićJelaska,L., M.Blanuša,P.Durbešić, S.

D.Jelaska,2007: Heavy metal concentrations

Scholz, R. W., N. Nothbaum, and T. W. May,
in ground beetles, leaf litter and soil of a forest

1994: Fixed and hypothesis-guided soil samecosystem.
Ecotoxicology and Environmental

pling methods – Principles, strategies and exam-
Safety 66: 74–81.

ples (In: Environmental Sampling for Trace
Analysis, Ed. Market B.). VCH Verlagsgesell

Tews, J., U. Brose,V. Grimm, K. Tielbörger,

M. C. Wichmann, M. Schwager, and F.
Jeltsch,2004:Animal species diversity driven

schaft GmbH,Weinheim, pp. 335–345.
Seifert,B., 2007: DieAmeisen Mittel- und Nordeuby
habitat heterogeneity/diversity: the importance

ropas. LutraVerlags undVertriebsgesellschaft

of keystone structure. J. Biogeogr. 31: 79–92.

Görlitz/Tauer, pp. 368.
Thiele, H.U., 1977: Carabid beetles in their environ-

Stephens, S.S., and M. R. Wagner,2006: Using

ments. Zoophysiology and ecology 10. Springer

Ground Foraging Ant (Hymenoptera: Formici-

Verlag, Berlin, 369 pp.

dae) Functional Groups as Bioindicators of FoTrautner,
J., and K.Geigenmüller,1987: o San-

rest Health in NorthernArizona Ponderosa Pine
dlaufkäfer Laufkäfer. J. Margraf Publisher,

Forests. Environ. Entomol. 35: 937–949.
Aichtal, 488 pp.

SAŽETAK: Ovim istraživanjem željeli smo utvrditi razlike u sastavu te morfološkim i ekološkim
značajkama trčaka i mrava u različlitim šumskim staništima na području Parka prirode Medvednica.
Kako bi kvantificirali razlike između istraživanih područja, odredili smo raznolikost strukture
prisutne vegetacije te raznolikost biljnih vrsta na šest ploha. Značajke kao što su pokrovnost drveća,
grmlja, prizemnog bilja te listinca na svakoj plohi, koristili smo u mjerenju kompleksnosti strukture
staništa. Strukturna kompleksnost staništa opadala je s nadmorskom visinom. Mravi i trčci uzorkovani
su metodom lovnih posuda.

Bogatstvo vrsta mrava pozitivno je korelirano s kompleksnošću staništa, posebice s količinom
listinca, za razliku od trčaka. Veći udio mrava koji su ekološki oportunisti zabilježen je u otvorenijim
staništima sa slabije izraženom heterogenom strukturom staništa, za razliku od trčaka kod
kojih veličina tijela korelira s kompleksnošću staništa. Stabilnija struktura zajednica trčaka s
većom brojnosti dominantnih vrsta na plohama smještenim na nižoj nadmorskoj visini može se
objasniti kompetitivnim isključivanjem, kao posljedicom stabilnijih ekoloških uvjeta u odnosu na
plohe smještene na vrhu planine. Vrste koje su prilagođene hladnijim klimatskim uvjetima te nastanjuju
zasjenjena područja na višim nadmorskim visinama, kao što su beskrilne vrste trčaka,
specijalistiCychrus caraboidesiCarabus irregularis, te boreo-montana vrsta mrava Camponotus
herculeanus, zabilježene su na najvišim nadmorskim visinama i sjevernim ekspozicijama Medvednice.
Neke od ovih vrsta ne nastanjuju niža područja, te u slučaju uništavanja povoljnih staništa
njihov opstanak na Medvednici može postati ugrožen. Jedan od razloga može biti i uslijed klimatskih
promjena koje utječu na visokoplaninske vrste hladnijih područja. Sveukupni rezultati ovog
istraživanja ukazuju na potrebu očuvanja složene strukture šumskih staništa, posebice na vršnim
dijelovima Medvednice.

Ključne riječi: biološka raznolikost, struktura vegetacije, listinac, nadmorska visina,
očuvanje prirode, šumsko stanište