ISSN 1408-3671 
UDK 57(497.4) 
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Društvo biologov Slovenije 
ISSN 1408-3671 
UDK 57(497.4) 
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IJIJLJANA 
~ACTA 
~ BIOLOGICA 
SLOVE NICA 
VOL. 45 ŠT. 2 LJUBLJANA 2002 
prej/formerly BIOLOŠKI VESTNIK 
izdajate! j/pu bi isher 
Društvo biologov Slovenije 
Acta Biologica Slovenica 
ACTA BIOLOGICA SLOVENICA 
LJUBLJANA 2002 Vol. 45, Št. 2: 1- 52 
Glasilo Društva biologov Slovenije - Journal Of Biological Society of Slovenia 
Izdaja - Published by 
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ACTA BIOLOGICA SLOVENICA 
LJUBLJANA 2002 Vol. 45, Št. 2: 3 - 14 
Sprejeto (accepted): 2002-05-27 
Food choice experiments with cadmium nitrate dosed food in terrestrial 
isopod Oniscus asellus (Crustacea) 
Poskusi izbire hrane z različno vsebnostjo kadmijevega nitrata na 
kopenskem enakonožcu Oniscus asellus (Crustacea) 
Ulf Iskender KASCHL1, Primož ZIDAR2, Jasna ŠTRUS2, Volker STORCH1 
1 Institute for Zoology, University of Heidelberg, Im Neuenheimer Feld 230, 69120 Heidelberg, 
Germany 
2 Department of Biology, Biotechnical Faculty, University of Ljubljana, Večna pot 111, SI-1000 
Ljubljana, Slovenia; e-mail: primoz.zidar@uni-lj.si 
Correspondence to Primož Zidar 
Abstract. The influence of different concentrations of cadmium nitrate on food 
choice behaviour was studied in the terrestrial isopod Oniscus asellus. In paired food 
choice tests, consumption rates were compared in relation to cadmium nitrate 
concentrations and duration of feeding. The consumption of contaminated food was 
reduced at even the lowest cadmium concentration already in the first week of feeding. 
In the third week the consumption rates for uncontaminated and contaminated food 
reached a ratio of 6:4 in ali animal groups. Consumption of contaminated food during 
the experiment resulted in increased cadmium content in the animals. It is presumed 
that O. asellus cannot distinguish food according to cadmium concentration. The 
difference in consumption rates between uncontaminated and cadmium-contaminated 
food could be based on integration of feeding behaviour and the adverse metabolic 
effects of cadmium. 
Keywords: Isopods, Oniscus asellus, food-choice, cadmium nitrate, contamination, 
consumption rate, accumulation 
Izvleček. Proučevali smo vpliv različnih koncentracij hrani dodanega kadmijevega 
nitrata na izbiro hrane pri kopenskem enakonožcu Oniscus asellus. V poskusih izbire 
smo primerjali stopnjo hranjenja z neonesnaženo in onesnaženo hrano, v odvisnosti 
od koncentracije kadmijevega nitrata in trajanja hranjenja. Živali so zaužile manj 
onesnažene hrane že v prvem tednu poskusa izbire tudi pri najnižji koncentraciji 
kadmijevega nitrata dodanega hrani. V tretjem tednu poskusa je bilo pri vseh skupinah 
živali razmerje med stopnjo hranjenja z neonesnaženo in onesnaženo hrano 6:4. Pri 
živalih, ki so med poskusom jedle hrano z dodanim kadmijevim nitratom je bila telesna 
vsebnost kadmija povečana. Iz rezultatov sklepamo, da raki enakonožci verjetno ne 
razlikujejo med hrano z različno vsebnostjo kadmijevega nitrata. Razlike v stopnji 
4 
Introduction 
Acta Biologica Slovenica, 45 (2), 2002 
hranjenja z neonesnaženo in onesnaženo hrano so verjetno posledica prehranjevalnega 
vedenja živali povezanega s presnovnimi učinki kadmija. 
Ključne besede: enakonožci, Oniscus asellus, izbira hrane, kadmijev nitrat, 
onesnaževanje, stopnja hranjenja, akumulacija 
Faced with an ever growing input of waste substances to the chemical cycles of the biosphere due 
to anthropogenic activities, the importance of assessing levels of pollution by biomonitoring 
programmes in various ecosystems has gained increasing recognition over the last decades (BAYNE 
1979, HoPKJN 1989, GoLDBERG & BERTINE 2000). The success of the 'Mussel Watch Program' in 
marine ecosystems has led to considerable efforts to establish similar programmes for terrestrial 
ecosystems. Terrestrial isopods are amongst the most promising animals for globa! terrestrial 
biomonitoring, and the species Porcellio scaber has been proposed as the terrestrial biomonitoring 
equivalent of Mytilus edulis (CoRTET & al. 1999, CoUGHTREY & al. 1997, DROBNE 1997, HoPKIN & al. 
1986, HoPKIN 1990, HoPKIN & al. 1993, PAOLETTI & HASSALL 1999). However, in recent years some 
data on possible pollutant-dependent food selection behaviour have been presented (VAN CAPELLEVEEN 
& al. 1986, DALLINGER 1977, OoENDAAL & REINECKE 1999, DROBNE & al. 1995). lf terrestrial isopods 
are able to discriminate between differently contaminated food, their value as biomonitoring organisms 
would diminish. 
Terrestrial isopods play an important role in decomposition of organic material and fulfill most of 
the criteria required of a good biomonitor (HoPKIN 1989, HoPKIN & al. 1993, CouGHTREY & al. 1997, 
PAOLETTI & HASSALL 1999). Many aspects ofthe ecology of Porcellio scaber are well known , and it is 
one ofthe most studied isopods (CoRTET & al. 1999, DROBNE 1997, GuNNARSON 1987, OoENDAAL & 
REINECKE 1999, PAOLETTI & HASSALL 1999). Field studies and monitoring programmes have used the 
species successfully in assessing the bioavailability of pollutants (HAMES & HoPKIN 1989, HoPKIN 
1990, RABITSCH 1995). Terrestrial isopods accumulate the highest tissue concentrations of cadmium, 
copper, lead, and zine known for any invertebrate (HoPKIN 1989, Ho PKIN & MARTIN 1984). The 
robustness against pollution with metals in isopods apparently stems from a compartmentalization 
mechanism dependent on metal-containing granules in s mali cells of the hepatopancreas and possibly 
a detoxification mechanism with binding metallothioneins, which yet remains to be verified (CROMENTUnN 
& al. 1994, HAMES & HoPKIN 1991, HoPKIN 1989, PROSI & al. 1983). The shiny woodlouse Oniscus 
asellus has been proposed as a substitute for P scaber in areas where the latter is scarce or missing 
from the fauna (DROBNE 1997). O. asellus shows higher accumulation rates for cadmium and other 
metals than P scaber, and has likewise a wide distribution, thus constituting a suitable alternative to P 
scaber (HAMES & HoPKIN 1991, HoPKIN 1990). 
In recent years, much information has been gathered about the sensoty equipment of terrestrial 
isopods. They are able to distinguish between and show preferences for different qualities of food, e.g. 
different levels of fungal permeation of their food (ZIMMER & al 1996, GuNNARSON 1987, SzLAVECZ & 
MAIORANA 1990). In addition, some results offood choice experiments have suggested that terrestrial 
isopods may also sense different levels of contaminants in their food, e.g. P laevis has been shown to 
discriminate against cadmium sulfate added to leaves and P scaber can apparently discriminate and 
avoid lead and copper in its food (VAN CAPELLEVEEN & AL. 1986, DALLINGER 1977, ODENDAAL & 
REINECKE 1999). If terrestrial isopods have evolved mechanisms to avoid uptake of biotoxic substances 
from their food, then the concentrations of these substances in their tissues do not represent levels of 
habitat pollution, but are the result of an integrated behavioural response to pollution (DROBNE & al 
1995, DoNKER & BooERT 1991). However, one ofthe criteria for the use of an organism asa biomonitor 
U. l. Kaschl, P. Zidar, J. Štrus, V. Storch: Food choice experiments with cadmium nitrate ... 5 
is that accumulation of a pollutant should reflect exposure. Results from biomonitoring experiments 
using terrestrial isopods could therefore be unreliable and should be treated with caution (DROBNE & al. 
1995). 
In the present work we investigated the possibility of an avoidance mechanism for the consumption 
of cadmium-contaminated food by the terrestrial isopod O. asellus in paired food choice experiments. 
We aimed to show that the results from our experiments, which document preference for uncontaminated 
food, in correlation with previous publications, can be explained without assuming the existence of a 
sensory detection mechanism for cadmium in the diet, thus rendering the species an accurate biomonitor 
of bioavailable cadmium in its habitat. We assumed that the observed preference for uncontaminated 
food could be explained by the combined influence of feeding behaviour and the adverse metabolic 
effects of cadmium in the isopods. 
Materials and Methods 
Sampling of animals from the field 
About 250 specimens of Oniscus asellus were collected in October 2001 from the litter layer of a 
woodland area near a former smel ter site in the vicinity of Nussloch near Heidelberg, Germany. For 
three weeks the animals were kept in a glass container on moist plaster of Paris and fed on partly 
decomposed unpolluted leaves ofvarious tree species. The temperature in the glass container was kept 
steady at about 17°C, and exposure to direct light was prevented. To avoid desiccation, the leaves and 
the plaster of Paris were lightly sprayed with commercial bottled water every two days. After three 
weeks, ten animals were selected at random, lyophilised and weighed, and digested in a hot acid 
mixture (HN0
3
:HCl0
4 
= 7: 1; fina! temperature l 85°C) until dryness. The residue was suspended in 
1.5 ml HN0
3 
(0.2%) and analysed for cadmium content by flame atomic absorption spectrophotometry 
(AAS), using a Perkin Elmer AAAnalyst 100 atomic absorption spectrophotometer. Ali further animal 
samples were digested and analysed according to this method. 
Sampling of soil 
From the same site, five soil samples of the top 1 O cm of the soil horizon were collected, cleaned 
of ali visible organic components, and dried for two hours at 110° C, then homogenised in a mortar and 
digested in hot acid mixture (HN0
3
:HCI = 1 :3) until dryness. As the samples were not digested 
completely by this procedure, it was repeated once. The samples were then diluted with weak nitric 
acid (0.2 % ), and filtered. The filtrate was subsequently analysed for cadmium content by flame AAS 
analysis. 
Experimental set-up 
The animals were separated, sexed and weighed, and kept individually on moist filter paper in 
plastic petri dishes (0 = 9 cm). Only males and non-gravid females were selected for the experiment. 
The petri dishes were kept in a climate chamber at a relative humidity of 100% and under a 16 hours 
light and 8 hours dark regi me. Temperature was kept constant at 21 °C (±1 °C). Animals were checked 
every two days, and the filter paper was moistened with commercial bottled water if necessary. Any 
dead animals were removed immediately and their data excluded from the results. Moulting animals or 
animals that did not produce faecal pellets were marked and counted. 
The animals were fed exclusively with complex food pellets designed especially for this experiment. 
For the production offood pellets, partly decomposed hazel leaves (Corylus avellana) were collected 
in an unpolluted woodland area in the vicinity of Cerknica near Ljubljana, Slovenia. After drying the 
leaves at room temperature for severa! days, leaf stems were removed and the leaves pulverised with 
a coffee mili and subsequently sieved through a 0.25 mm mesh net. The leaf powder was dried for 2 
6 Aeta Biologiea Slovenica, 45 (2), 2002 
hours at 60°C. Other components of the complex food pellets were commercial Dr. Oetker' s gelatine, 
which was dried at 40°C, and fish food for aquarium fish (JBL Novobel; JBL GmbH& Co. KG), 
which was homogenised by hand in a glass mortar and dried for two hours at 60°C. 
Dry leaf powder, gelatine, and fish food were mixed in a 63:34:3 ratio and turned into a paste with 
demineralized water ( 15 ml per gram gelatine ). For the preparati on of contaminated food, four different 
amounts of cadmium nitrate (Cd(N0
3
)
2
) solution (2712.5 mg Cd/1) were added to the food paste to 
give nominal concentrations of O, 20, 45, 200 and 450 mg Cd kg·1 dry weight of food. In order to 
exclude food choice due to nitrate content (Hopkin 1989), corresponding amounts of potassium nitrate 
(KN0
3
) were added to the respective control food. Food pellets were formed out of the paste by 
depositing equal amounts in plastic blisters (V= 0.3 ml). The food pellets were allowed to solidify for 
24 h at 5°C, dried at room temperature for 24 h and then dried at 70°C for the next 48 h. The actual 
concentrations of cadmium in the food pellets were measured by AAS analysis and compared to the 
nominal cadmium contents (Tab. !). The amount of water-soluble cadmium in the food pellets was 
measured by soaking food pellets in demineralized water for 24 h at increasing temperatures up to 
40°C, centrifuging the solution and analysing the supernatant for Cd content by flame AAS (Tab. 1). 
The water soluble concentration of cadmium in food pellets was observed to be less than 1 O% of the 
actual cadmium concentration. 
Table 1: Concentration of cadmium in food pellets as measured by flame AAS (* = concentration 
below detection limit) 
Preglednica 1: Koncentracije kadmija v hrani izmerjene s plamensko AAS (* = koncentracija pod 
zaznavno mejo) 
Nominal Cone. (mg/kg) Measured Cone. (mg/kg) n=20 Water Soluble Cone. (mg/kg) n=IO 
Mean I SE Mean I SE 
o 0.06 0.02 * 
20 20.55 0.30 1.14 0.13 
45 45.34 O.SI 3.78 0.07 
200 194.55 2.69 14.05 0.20 
450 445.26 1.58 36.58 0.40 
Food pellets were offered in small plastic dishes (0 = 2 cm, height of rim ca. 3 mm) . Prior to food 
choice experiments, the animals were acquainted with the form and taste of the food pellets by offering 
them uncontaminated food for one week. After seven days, the animals were starved for 24 h to empty 
their guts (HAMES & HoPKIN 1991 ). Ten animals were selected randomly and analysed for tissue 
cadmium content by flame AAS. 
Food choice experiments 
The isopods were assigned to five groups of 35 individuals each, giving a total number of 175 
animals. Each animal was offered a choice between an uncontaminated and a Cd-contaminated food 
pellet; in the group "0-20" the nominal cadmium concentration in the contaminated pellet was 20 mg 
kg·1 dry food weight, in the group "0-45" 45 mg kg 1 dry food weight, in the group "0-200" 200 mg kg·1 dry 
food weight, and in the group "0-450" 450 mg kg·1 dry food weight. Animals in the control group "0-
0" were offered a choice of two uncontaminated, but differently marked food pellets. The dry weight 
of food pellets was determined before and after exposure to animals using a micro-balance. 
After seven days of the food choice experiment, the animals were starved for 24 h to empty their 
guts (HAMES & HoPKIN 1991 ). Eight animals of each group were randomly selected and analysed for 
tissue cadmium content by AAS analysis. The food pellets were collected, dried for two days at room 
U. I. Kaschl, P. Zidar, J. Štrus, V. Storch: Food choice experiments with cadmium nitrate ... 7 
temperature, cleaned from faeces and dried at 70 °C for 48 h. Consumption of food was calculated 
according to the differences in weight before and after the seven day period of exposure to the animals. 
The remaining animals were offered a new choice between new food pellets with the same Cd-
concentrations as before. The whole cycle of feeding, removal of food, starvation for 24 h, and random 
selection of eight animals for analysis of cadmium content by AAS analysis was repeated twice, 
resulting in a total exposure period of twenty-one days for the last cohort of animals. 
Analysis of data 
To compare food consumption rates (CR), the absolute consumption of food per week was di vided 
by the dry weight of animals. The <lata of the experiment were analysed using Microsoft Excel 97 
computer software. Ratios of consumption for uncontaminated food and contaminated food were 
calculated from consumption rates for each individual. The percentage of uncontaminated food consumed 
was calculated and compared to a null-hypothesis of 50 % with a two-tailed Student's t-test. Standard 
errors (SE) and 95 % confidence intervals were calculated where appropriate. Weekly consumption 
rates within groups and weekly consumption rates between groups were analysed for significant 
differences with a one-way ANOVA test and a Tukey test, using SPSS for Windows statistics software. 
Results 
Concentration of cadmium in soil samples and animals 
Analysis of soil samples by flameAtomic Absorption Spectrometry yielded a mean concentration 
of 11 .42 (SE= 1.63) mg Cd per kg dry weight of soil. Analysed animal sam ples from the field contained 
a mean of 64.54 (SE=5.26) mg Cd per kg dry weight of animals. Analysed animal samples from food 
choice experiment yielded increased levels of cadmium correlated to the duration of the experiment and 
the concentration of cadmium in the offered food (Fig. 1 ). 
:? 
180 
bO 160 ·;; 
~ 140 
>-, 
-o 120 o 
..o 
g 100 
80 
bO 
-" 60 
bO 
5 40 
-o 20 u 
o 
' 
Field '1l-O" ''0- 20" 
groups 
''0-45" "0-200" ''0-450" 
Figure 1: Concentration of cadmium in whole Oniscus asellus measured by flameAAS followed over 
three weeks ofthe food choice experiment. Concentration of cadmium in field animals is represented 
on the far left (mean ± SE). 
Slika 1: Koncentracije kadmija v telesu raka enakonožca Oniscus asellus, izmerjene s plamensko AAS 
v treh tednih poskusa izbire hrane. Povprečna koncentracija kadmija v živalih iz okolja je prikazana 
skrajno levo (povp. ± stand. napaka). 
8 Acta Biologica Slovenica, 45 (2), 2002 
Consumption rates 
During the preliminary experiment, the mean food consumption rate was 0.52 (SE=0.03) mg dry 
food weight per kg dry weight of animals. In the third week of the food choice experiment, the mean 
food consumption rate increased to 1.24 (SE=0.05). 
Combined consumption rates of contaminated and uncontaminated food increased significantly 
between the first and third week in group "0-0" (Tukey test: p < 0.05) (Fig. 2). In group "0-20", 
combined consumption rates between the first and second week are significantly different (ANOVA: 
p<0.05); however, with the Tukey test no statistically significant increase between weekly consumption 
rates could be shown. Combined consumption rates in group "0-45" were significantly higher in the 
second and third week (Tukey test: p < 0.005) than in the first week. In group "0-200", combined 
consumption rates in the third week were significantly higher than in the first week (Tukey test: p < 
0.05); between the first and second and the second and third week there was no statistically significant 
difference. In group "0-450", no statistically significant increase in weekly consumption rates could be 
shown. 
1,8 ...................... 
1,6 
1,4 
1,2 
iz u 
0,8 
0,6 
0,4 
0,2 
o 
"0-0" "O- 20" ··o-45" 
groups 
"0-2(XJ" "0-450" 
0 Weekl 
[] Week2 
m Week3 
Figure 2: Combined consumption rates (CR) of uncontaminated and contaminated food in different 
groups of animals for each of the three weeks of the food choice experiment (mean ± SE). 
Slika 2: Stopnja hranjenja (CR) z neonesnaženo in onesnaženo hrano skupaj v posameznih tednih 
poskusa izbire, pri različnih skupinah živali (povp. ± stand. napaka). 
Consumption rates of contaminated and uncontaminated food in different groups 
Consumption rates show that animals within group "0-0" did not significantly discriminate between 
food pellets during the three weeks (Fig. 3, Tab. 2). Animals within group "0-20" and "0-45" significantly 
preferred uncontaminated food to contaminated food in the first and third week but not in the second 
week. Animals of the group "0-200" consumed significantly more uncontaminated food in ali three 
weeks. In group "0-450", there was no significant preference for uncontaminated over contaminated 
food in the first week; however, in the second and third week, animals consumed significantly less 
contaminated food. 
U. l. Kaschl, P. Zidar, J. Štrus, V. Storch: Food choice experiments with cadmium nitrate ... 9 
A 
100 
90 
80 
70 
i: ••• ! ... 
60 
i 
! ... r 
% 50 l. 
40 
30 
20 
10 
o 
0-0 0-20 0-45 0-200 0-450 
B 
100 
90 
80 
70 f ... 
60 I T 
t ... 
% 50 t .... .L 
40 
30 
20 
10 
o 
0-0 0-20 0-45 0-200 0-450 
C 
100 
90 
80 
70 
f · 60 f ··· f ... 1· r 
% 50 I 
40 
30 
20 
10 
o -
0-0 0-20 0-45 0-200 0-450 
Figure 3: Consumption rate (CR) ofuncontaminated food shown as percentage values ( •) of combined 
CR of uncontaminated and Cd-contaminated food (means and 95% conf. int.): A = 1 st week, B = 2"d 
week, C = 3,d week of food choice experiments. Stars above bars represent significant differences 
between CR ofuncontaminated food and 50 % (horizontal line) (t-test: * = p < 0.05; ** = p < 0.01; *** 
= p < 0.005). 
Slika 3: Stopnja hranjenja (CR) z neonesnaženo hrano prikazana kot odstotek ( •) skupne stopnje 
hranjenja z neonesnaženo in s kadmijem onesnaženo hrano (povp. in 95% interval zaupanja): A = 1. 
teden, B = 2. teden, C = 3. teden poskusov izbire hrane. Statistična značilnost razlik med stopnjo 
hranjenja z neonesnaženo hrano in 50% (vodoravna črta) je prikazana z zvezdicami (t-test: * = p < 
0.05; ** = p <O.O!;***= p < 0.005). 
10 Acta Biologica Slovenica, 45 (2), 2002 
Differences in combined consumption rates between the groups in the first week were mainly due 
to differences in consumption of uncontaminated food (Tab. 2). In the second week, the differences 
between combined CRs of the groups were mainly connected with consumption of contaminated food, 
whereas consumption levels of uncontaminated food were roughly similar in ali groups (Tab. 2). 
During the third week, CRs between uncontaminated and contaminated food reached almost identical 
ratios of about 6:4 in ali groups (Tab. 2, Fig. 3). 
Table 2: Feeding and moulting behaviour of Oniscus asellus in different groups over three weeks of 
food choice experiment. (O = uncontaminated food, Cd = food contaminated with cadmium; CR = 
Consumption Rate) 
Preglednica 2: Prehranjevanje in levitev v treh tednih poskusa izbire pri Oniscus asellus iz različnih 
skupin. (O= neonesnažena hrana, Cd = hrana onesnažena s kadmijem; CR = stopnja hranjenja) 
''0-0" "0-20" "0-45" ''0-200" "0-450" 
Mean SE Mean SE Mean SE Mean SE Mean SE 
Week 1 CRO 0.53 0.05 0.79 0.07 0.47 0.05 0.64 0.07 0.41 0.05 
CRCd 0.50 0.06 0.31 0.03 0.29 0.02 0.29 0.03 0.36 0.05 
% of moulting animals 33.33 21.43 42.86 24.14 38.46 
Number of animals 26 27 27 28 24 
Week2 CRO 0.54 0.06 0.71 0.05 0.67 0.07 0.67 0.07 0.64 0.07 
CRCd 0.56 0.04 0.65 0.06 0.63 0.05 0.40 0.05 0.30 0.04 
% of moulting animals 5.26 10.00 10.00 33.33 41.18 
Number of animals 18 19 19 20 16 
Week3 CRO 0.66 0.04 O.SI 0.06 0.78 0.06 0.75 0.10 0.70 0.10 
CRCd 0.77 0.10 0.56 0.07 0.51 0.05 0.56 0.06 0.43 0.05 
% of moulting animals 18.18 8.33 o o 11.11 
Number of animals II 12 12 12 9 
Discussion 
Compared with <lata from HoPKIN (1989), the soil samples from Nussloch show intermediate to 
slight pollution Jevels with cadmium. Variability in body concentrations of cadmium in field animals 
was shown. It has been demonstrated that there may be variability of metal concentrations in sampled 
field populations, and that levels of metals in the hepatopancreas of individual woodlice may be more 
than twice the mean value for the whole population within a site (HoPKIN & MARTIN 1984, HoPKIN & 
al. 1986). Furthermore, the amount of pollutants in soil and litter are not always predictive of 
bioavailability and actual accumulation of pollutants, especially in environments that are polluted due 
to anthropogenic activity, such as former smelters, mines, and roadsides (HoPKIN & al. 1986). The 
variable cadmium concentrations in the sampled animals may be further explained by the variable age 
of the sample animals (reviewed in DROBNE 1997). 
Cadmium concentrations in experimental animals increased with increasing concentrations of 
cadmium in the food. This is in accordance with laboratory experiments on the accumulation of 
pollutants from food according to consumption and dry weight (DROBNE 1996, HoPKIN & MARTIN 
1984) 
Food consumption rates in ali groups increased with the course of the experiment. This may be 
explained by the great numbers of moulting animals in the first week, and the fact that animals were not 
adapted to laboratory conditions. It is recommended to accustom isopods to laboratory conditions for 
some weeks before the start of an experiment, as moulting is stimulated by changed conditions and 
U. l. Kaschl , P. Zidar, J. Štrus, V. Storch: Food choice experiments with cadmium nitrate ... 11 
thus interferes with normal behavioural responses to a stimulus (STEEL 1980; DROBNE 1996; ZIDAR & 
al. 1998). An interval of 28 days between two ecdyses has been reported for O. asellus (STEEL 1982), 
so the impact of moulting behaviour on food consumption rate cannot be excluded in long-term 
experiments. Almost 80% of animals moulted during the experiments which resulted in decreased food 
consumption rates. Another factor influencing the food consumption rate might be the phenomenon of 
hyperphagia which describes the observation that isopods in the laboratory tend to con sume more food 
than a comparable group of field animals (BECK & BRESTOWSKY 1980, HoPKIN 1990). 
The combined consumption rates for the whole duration of the food choice experiment show that 
low levels of cadmium (20 and 45 mg kg·1 dry food weight) seem to have only little orno adverse 
effects on consumption rate. This is in accordance with dala from the literature, which reports the low 
observed effective concentration (LOEC) for Cd in P. scaber at about 20 mg Cd kg·1 (DoNKER & 
BoGERT 1991 ). For intermediate and high levels of Cd-pollution in the food a decrease of food 
consumption was observed. 
After three weeks of the food choice experiment, the amounts of uncontaminated food and 
contaminated food consumed reached an apparently stable ratio of 6:4 in ali groups, thus suggesting 
some kind of avoidance reaction to cadmium independent of cadmium concentration. If O. asellus was 
indeed able to taste and discriminate against cadmium contaminated food, we would expect a reaction 
in relation to cadmium concentration. 
However, some interesting pattems of discrimination against contaminated food related to the 
concentration of cadmium and the duration of exposure can be observed. In groups "0-20" and "0-45" 
we found a similar pattem of rejection over three weeks. In the first week, there was a strong 
preference for uncontaminated food, whereas in the second week, the avoidance reaction against 
cadmium contaminated food was barely observable. In the third week, animals again discriminated 
significantly against contaminated food. In group "0-200", preference for uncontaminated food dropped 
from a ratio of 7:3 in the first week to afina! 6:4 in the third week. In group "0-450", discrimination 
against contaminated food in the first week was not very pronounced, whereas in the second week, 
animals strongly discriminated against contaminated food, reaching a consumption ratio of about 6:4 
in the third week. 
The differences and time-dependent pattem of the avoidance reactions to cadmium contaminated 
food observed in the food-choice experiment might be explained by the assumption oftwo combined 
physiological effects: Firstly, an animal that starts to feed on cadmium contaminated food will suffer 
adverse me tabo lic effects from cadmium which will cause a premature cessation of feeding . Secondly, 
the intake of cadmium initiates a detoxification mechanism. 
As described in the literature, isopods show a regular cycle of feeding behaviour (HAMES & HoPKIN 
1990). After a phase of resting, animals void their guts, search for food, and, upon finding suitable 
food, start feeding. It may be assumed that once an animal starts feeding on suitable food, it will 
continue to feed on the same food if not disturbed. If animals start to feed on contaminated food, they 
will ingest this food until the ingested cadmium produces the assumed adverse metabolic effect, which 
will res uit in cessation of feeding. The higher the concentrations of ingested cadmium, however, the 
more pronounced the toxic reaction will be, thus reducing uptake even of uncontaminated food. The 
effect observed in group "0-450", where consumption levels of contaminated and uncontaminated 
food are equally low, might therefore be explained by assuming that the ingested levels of Cd are 
already so toxic that it also reduces the consumption of uncontaminated food . 
Comparing the course of food consumption in the respective groups over tirne supports the second 
assumption. If animals that are exposed to small concentrations of cadmium acclimatise to these levels 
of contaminants in their food, after some tirne they will not be subject to the negative metabolic effects 
of low doses of cadmium in their food. Consumption rates of uncontaminated and contaminated food 
will be more equal, as was observed in the second week for groups exposed to low cadmium 
concentrations (groups "0-20" and "0-45"), and to a lesser degree in the group exposed to intermediate 
12 Acta Biologica Slovenica, 45 (2), 2002 
cadmium concentrations (group "0-200"). In the group exposed to high cadmium concentrations 
(group "0-450"), this acclimatisation process will take longer. In the third week, the acclimatisation to 
cadmium in the food reached a steady s tate, which allows consumption of contaminated food, but stili 
favours consumption of uncontaminated food. This s tate is represented by similar consumption ratios 
for uncontaminated and contaminated food in ali groups. 
Conclusions 
In paired food choice tests, the consumption of cadmium contaminated food by terestrial isopods 
is reduced. This has formerly been interpreted as the ability of isopods to detect and discriminate 
against pollutants in their food, rendering results from biomonitoring experiments less significant. In 
our experiment, we could show that O. ase/lus does eat less contaminated food than uncontaminated 
if offered a choice. However, this effect is not related to the concentrations of the pollutant or the 
duration of exposure. This leads to the presumption that O. asellus (and perhaps other terrestrial 
isopods) cannot distinguish food according to cadmium concentration. The difference in consumption 
rates between uncontaminated and contaminated food is based on integration offeeding behaviour and 
the adverse metabolic effects of cadmium. 
Ifthis presumption is proven to be correct, results from experiments where O. asellus is employed 
as biomonitoring organism are accurate. The findings of this paper strongly support the generally held 
concept that the maximum amount of knowledge has to be accumulated about a species prior to its 
potential use as a biomonitor. 
Acknowledgements 
We are grateful to Doc. Dr. A. Blejec for his critical comments on the statistical evaluations 
performed. This work was financially supported by the Slovenian Ministry ofEducation, Science and 
Sport (projects No. Zl-3189 and PO-0525) and bilateral cooperation within the Socrates programme. 
Povzetek 
Kopenski raki enakonožci v telesu kopičijo kovine, kar jih uvršča med možne pokazatelje dostopnosti 
kovin v okolju. Nekatere raziskave so pokazale, da enakonožci izbirajo hrano z nižjo vsebnostjo kovin. 
Izbira hrane z manj kovin lahko pomeni, da vsebnost kovin v enakonožcih ne kaže količine dostopnih 
kovin v okolju. V tem primeru bi bila vprašljiva uporaba enakonožcev kot pokazateljev obremenjenosti 
okolja s kovinami. 
V predstavljenem delu smo proučevali vpliv različnih koncentracij hrani dodanega kadmijevega 
nitrata na izbiro hrane pri kopenskem enakonožcu Oniscus ase/lus. Živalim smo hkrati ponudili hrano 
brez dodanega kadmija in hrano z dodanim kadmijevim nitratom (20, 45, 200 in 400 mg Cd kg· 1 suhe 
teže hrane). Hrano smo pripravili kot mešanico mletih listov leske, ribje hrane in želatine v razmerju 
63:3:34. Količino nitratnih ionov v ponujeni hrani smo uravnotežili z dodajanjem raztopine kalijevega 
nitrata (KNO
3
). Primerjali smo stopnjo hranjenja z neonesnaženo in onesnaženo hrano, v odvisnosti 
od koncentracije kadmijevega nitrata v hrani in trajanja hranjenja. 
Živali so zaužile manj s kadmijem onesnažene hrane že v prvem tednu poskusa izbire tudi pri 
najnižji koncentraciji kadmijevega nitrata dodanega hrani. V tretjem tednu poskusa je bilo pri vseh 
skupinah živali razmerje med stopnjo hranjenja z neonesnaženo in onesnaženo hrano 6:4. Pri živalih, 
U. I. Kaschl, P. Zidar, J. Štrus, V. Storch: Food choice experiments with cadmium nitrate ... 13 
ki so med poskusom jedle hrano z dodanim kadmijevim nitratom je telesna vseb:10st kadmija naraščala 
s koncentracijo kadmija v hrani in časom hranjenja. 
Rezultati poskusov izbire hrane so pokazali, da O. asellus zaužije manj s kadmijem onesnažene 
hrane, če ima na voljo neonesnaženo. Vendar razmerje med zaužitjem neonesnažene in onesnažene 
hrane ne narašča s koncentracijo kadmija v hrani kakor tudi ne s časom hranjenja. Tako sklepamo, da 
O. asellus verjetno ne razlikujejo med hrano z različno vsebnostjo kadmijevega nitrata. Razlike v 
stopnji hranjenja z neonesnaženo in onesnaženo hrano so verjetno le posledica prehranjevalnega 
vedenja živali povezanega s presnovnimi učinki kadmija. Če je naša domneva pravilna, uporaba O. 
asellus kot pokazatelja dostopnosti kovin v okolju ni vprašljiva. 
Literature: 
BAYNE B.L. 1979: Assessing effects of marine pollution. Nature 280: 14-15. 
BECK L. & E. BRESTOWSKY 1980: Auswahl und Verwertung verschiedener Fallaubarten durch Oniscus 
asellus (Isopoda). Pedobiologia 20: 428-441. 
CoRTET J., A. GoMoT-DE V AUFLERY, N. Po1NsoT-BALAGUER, L. GoMoT, C. TEXIER & D. CLUZEAU 1999: 
The use ofinvertebrate soil fauna in monitoring pollutant effects. Eur. J. Soil. Biol. 35 (3): 
115-134. 
CouoHTREY P. J., M. H. MARTIN & E. W. YouNo 1997: The woodlouse, Oniscus asellus, asa monitor 
of environmental cadmium levels. Chemosphere 12: 827-832. 
CROMMENTUIJN T., J. A. M. DooDEMAN & A. DooRNEKAMP 1994: Lethal body concentrations and 
accumulation patterns determine time-dependent toxicity of Cadmium in soil arthropods. 
Environmental Toxicology and Chemistry 13 (11): 1781-1789. 
DoNKER M.H. & C.G. BooERT 1991: Adaptation to cadmium in three populations of the isopod 
Porcellio scaber. Comp. Biochem. Physiol. 12: 143-146. 
DALLINGER R. 1977: The flow of copper through a terrestrial food chain: III. Selection of an optimum 
copper diet by isopods. Oecologia 30: 273-276. 
DROBNE D., P. ZIDAR & P. BrnRREGAARD 1995: Could proposed bio-monitoring organisms seleet 
differently contaminated food? Abstracts, Sixth International Symposium, Metal Compounds 
in Environment and Life, Jiilich, Germany, May 9-12, p. 35. 
DROBNE D. 1996: Zine toxieity to terrestrial isopods (Isopoda, Crustaeea). Dissertation Thesis. Univerza 
v Ljubljani, Biotehniška fakulteta, Oddelek za biologijo, Ljubljana, Slovenia. 
DROBNE D. 1997: Terrestrial isopods- a good choice for toxicity testing ofpollutants in the terrestrial 
environment. Environmental Toxicology and Chemistry 16: 1159-1164. 
GoLDBERG E. D. & K. K. BERTINE 2000: Beyond the mussel wateh - new direetions for monitoring 
marine pollution. The Scienee of the Tota) Environment 247: 165-174. 
GuNNARSSON T. 1987: Soil arthropods and their food: choiee, use and consequences. Doctoral thesis. 
Department of Ecology, University of Lund, Sweden. 
HAMEs C. A. C & S. P. HoPKIN 1989: The structure and function of the digestive system of terrestrial 
isopods. J. Zool. 217: 599-627. 
HAMES C. A. C. & S. P. HoPKIN 1990: A daily eycle of apoerine secretion by the B eells in the 
hepatopancreas ofterrestrial isopods. Can. J. Zool. 69: 1931-1937. 
HAMES C. A. C. & S. P. HoPKIN 1991: Assimilation and loss of Hl9Cd and 65Zn by the terrestrial isopods 
Oniscus asel/us and Porcel/io scaber. Buli. Environ. Contam. Toxicol. 47: 440-447. 
HoPKIN S. P. & M. H. MARTIN 1984: Heavy metals in woodliee. Symp. Zool. Soe. (London) 53: 143-
166. 
HoPKIN S. P, G. N. HARDISTY & M. H. MARTIN 1986: The woodlouse Porcellio scaber asa 'biologieal 
Indicator' of zine, cadmium, lead and copper pollution. Environmental Poli uti on 11: 271-290. 
14 Acta Biologica Slovenica, 45 (2) , 2002 
HoPKIN S. P. 1989: Ecophysiology ofmetals in terrestrial invertebrates. Elsevier Applied Sciences, 
London and New York. 
HorKIN S. P. 1990: Species-specific differences in the net assimilation of zine, cadmium, lead, copper 
and iron by the terrestrial isopod Oniscus asellus and Porcellio scaber. Joumal of Applied 
Ecology 27: 460-474. 
HOPKIN S . P. , DT. JoNES & D. DIETRICH 1993: The isopod Porcellio scaber as a monitor of the 
bioavailability ofmetals in terrestriaJ ecosystems: towards aglobal ' woodlouse watch ' scheme. 
The Science of the Tota! Environment, Supplement J 993 , Elsevier Science Publishers B.V., 
Amsterdam, 357-365. 
OoENDAAL J. P. & A . J. REJNECKE 1999: The sublethal effects and accumulation of cadmium in the 
terrestrial isopod Porcellio laevis Latr. (Crustacea, Isopoda). Arch. Environ. Contam. Toxicol. 
36: 64-69. 
PAOLETTI M . G. & M . HASSALL J 999: Woodlice (lsopoda: Oniscidea): their potential for assessing 
sustainability and use as bioindicators. Agriculture, Ecosystems and Environment 74: 157-
165. 
PROSI F., V. STORCH & H. H. JANSSEN (1983) : Small cells in the midgut glands of terrestrial Isopoda: 
Sites of heavy metal accumulation. Zoomorphology 102: 53-64. 
RABITSCH W. B. 1995: Metal accumulation in arthropods near a lead/zinc smelter in Arnoldstein, 
Austria. I. Environmental Pollution 90, No. 2: 221-237. 
STEEL C. G. H. 1980: Mechanisms of coordination between moulting and reproduction in terrestrial 
isopod (Crustacea). Biol. Buli. 159, 206-218. 
STEEL C. G. H. 1982: Stages of the intermoult cycle in the terrestrial isopod Oniscus asellus and their 
relation to biphasic cuticle secretion. Can. J. Zool. 60: 429-437. 
SzLAVEcz K. & V. C. MAIORANA 1990: Food selection by isopods in pared choice tests. In: Biology of 
Terrestrial Isopods. Third Intemational Symposium. Poitiers, July 10-12, 1990. 
V AN CAPELLEVEEN H. E. , N. M. VAN STRAALEN, M. VAN DEN BERG & E. VAN W ACHEM 1986: Avoidance as 
a mechanism of tolerance for lead in terrestrial arthropods . In: Proc 3rd Eur. Congress. 
Entom. Part 2 (H.H.W. Veldhuis, Ed.), 251-255 . N. Ev., Amsterdam. 
ZIDAR P., D. DROBNE & J. ŠTRus 1998: Determination of moult stages of Porcellio scaber (Isopoda) for 
routine use. Crustaceana 71, 6: 646-654. 
ZJMMER M., G. KAuTZ & W. ToPP 1996: Olfaction in terrestrial isopods (Crustacea: Oniscidea): responses 
of Porcellio scaber to the odour oflitter. Eur. J . Soil. Biol. 32 (3) : 141-147. 
ACTA BIOLOGICA SLOVENICA 
LJUBLJANA 2002 Vol. 45, Št. 2: 15 - 24 
Sprejeto (accepted): 2002-05-27 
Adaptive mutation: shall we survive bacterial genetic skills? 
Adaptivna mutacija: bomo preživeli genetske veščine bakterij? 
Rok KRAŠOVEC, Igor JERMAN 
Institute BION, Stegne 21, 1000 Ljubljana, Slovenia 
Corespondent author: 
Rok Krašovec, Institute Bion, Stegne 21, SI-1000 Ljubljana; Slovenia 
tel. 386 1 513 11 47; fax. 386 1 513 11 46; e-mail: rok.krasovec@volja.net 
Abstract. The origin and dynamics of genetic variations is one of the key questions 
in the modem science that has stili not come out with afina! answer. Emerging concepts 
regarding genetic variation have always produced a great controversy because they 
hold a key to unlock a great mystery of evolution. With such a powerful motivation 
scientist working in the molecular biology, genetics and biochemistry gathered a vast 
amount of experimental data showing us that a genome is a dynamic, hierarchically 
organized and complex integrated system for storing and processing information. 
Dynamic balance between stability and mutability of DNA nucleotide sequences is 
essential for a proper functioning of the organism. Besi de many DNA repairing proteins 
and DNA protective mechanisms organisms possess also biochemical systems capable 
of changing DNA information. One of the most controversial and at the same tirne the 
most informative one is a phenomenon called adaptive mutation . We shall review 
findings conceming the phenomenon of adaptive mutation in prokaryotes and point 
out an urgent need for the upgrade of the awkward neo-darvinistic view on the origin 
.of the genetic variation. 
Keywords: adaptive mutation, inducible mutagenesis, transposable elements, 
signal transduction network, neo-darvinism 
Izvleček. Izvor in dinamika genetskih variacij je eden od ključnih vprašanj modeme 
znanosti, ki še vedno čaka na dokončen odgovor. Koncepti v zvezi z genetskimi 
variacijami povzročajo veliko polemik, saj nosijo ključ do skrivnosti evolucije. 
Molekularni biologi, genetiki in biokemiki, so zavoljo tako močnega motiva zbrali 
ogromno količino eksperimentalnih podatkov, ki kažejo, da je genom dinamičen, 
hiearhično organiziran in kompleksno integriran sistem za shranjevanje in obdelovanje 
informacij. Dinamično ravnotežje med stabilnostjo in spremenljivostjo DNK zapisa je 
nujno za pravilno funkcioniranje organizma. Poleg številnih DNK popravljalnih 
proteinov in DNK varovalnih mehanizmov, imajo organizmi tudi biokemične sisteme, 
sposobne spreminjanja DNK informacije. Eden najbolj polemičnih in hkrati najbolj 
poučnih je fenomen imenovan adaptivna mutacija. Najin namen je pregledati dognanja 
16 
Introduction 
Acta Biologica Slovenica, 45 (2), 2002 
o adaptivni mutaciji pri prokariontih in pokazati, da je nadgradnja togega 
neodarvinističnega pogleda na izvor genetskih variacij, nujno potrebna. 
Ključne besede: adaptivna mutacija, inducibilna mutageneza, transponibilni 
elementi, signal prevajalna mreža, neodarvinizem 
In its essence genetic variation stirred up scientists as far back as the Darwin's era in the 19th 
century. Despite the fact that Darwin possessed no knowledge about genetics, he suspected that 
variations between individuals or mutations did not originale only from random events. He thought 
that there were also adaptive mutations induced by the environment. However, he commented that it 
was reasonable to treat them as random, as long as we do not know their origin (Darwin 1859). 
At the start of the antibiotic era in 1940' the emergence of antibiotic resistant mutants stimulated 
Luria & DelbrUck ( 1943) to perform a classical experiment, where they exposed bacterial population 
to a lethal selective pressure of a bacteriophage T 1. Phage immediately killed non-resistant cells and 
only cells with a pre-existing specific mutation could survive the exposure to the phage. A thorough 
analysis of a number of surviving colonies and their distribution in independent cultures proved the 
existence of random mutations that arose during the growth with no relation to the selective pressure. 
Together with the persuasive results from Lederberg & Lederberg (1952) and Cavalli-Sforza & 
Lederberg (1956) researchers concluded that all mutations arose randomly, prior to or in the absence 
of the selective pressure. They considered mutations as only a consequence of a non-perfection DNA 
replication machinery. Genetic variations thus appeared to be totally independent from the needs of the 
organisms in the environment with natura! selection asa fina! statistical filter to decide which organism 
will survive. This notion became the central idea of the neo-darvinistic evolutionary biology. 
First results that contradicted this well established dogma came already in the 50' when Ryan 
demonstrated genomic changes without the DNA replication (Ryan & Wainwright 1954, Ryan 1955), 
but he failed to show that changes follow the applied selective pressure. First indication that the 
environment can influence the mutation process came much later when Shapiro ( 1984) used genetically 
engineered bacterial cells with a mutation that prevented them to use a specific carbon source. By using 
a non-lethal selective pressure, he observed accumulation of mutants on the plates during the selection. 
The tuming point in the research on the origin and dynamics of the genetic variation came four 
years later when Cairns and co-workers (1988) challenged the established biological dogma with an 
argument that mutations in the direction of phage or antibiotic resistance are not expressed until after 
the period of growth. Since lethal conditions used killed bacterial cells orat least completely inhibited 
their growth, classical experiments could not detect and did not exclude the existence of mutations that 
could arise after the selective pressure was applied. The tem pora] and numerical distribution of surviving 
mutant colonies clearly demonstrated that during non-lethal and non-mutagenic selective pressure 
non-growing or slowly growing bacterial cells experience a specific mutation, named adaptive mutation 
that relieves the selective pressure. 
The notion that mutations arise in non-dividing stationary cells (Caims & Foster 1991) was 
strongly reminiscent of Lamarck's ideas. This, of course, provoked a great upheaval in the scientific 
community (Lenski & al. 1989, Lenski & Mittler 1993), and it was mainly due to the fact that adaptive 
mutations ari se only after the selective pressure was applied and in the presence of the selective agent. 
So it seemed that in some way the applied stress directs mutations in a useful way on appropriate sites 
(Cairns & al. 1988, Foster & Caims 1992). 
After the presen tati on of such challenging resul ts and thinking, researchers from various countries 
and different backgrounds performed many new studies and a great deal of experimental data were 
collected. Improvements in experimental techniques enabled researchers to geta deeper understanding 
R. Krašovec, l. Jerman: Adaptive mutation: shall we survive bacterial genetic skills 17 
of the complexity of molecular events taking place during the processes of mutation, which inevitably 
lead to a general agreement that an awkward classical neo-darvinistic doctrine, claiming that there is no 
relation between the needs of the organism and its mutability, must be upgraded. 
Currentfindings in adaptive mutation research 
Great efforts from many researchers in the last 14 years gave rise to numerous mutational systems, 
from which some of them became representative and often more explored and understandable. Our 
intention is to arrange a review of current findings in adaptive mutation research according to such 
systems and at the end present a discussion on the origin and dynamics of the bacterial genetic variation 
with pointing out some possible effects of such bacterial genetic variability on the species Homa 
sapiens sapiens. 
Cairns system 
Escherichia coli cells strain FC40 carry an a+ 1 frameshift mutation in an F' -located lacI-lacZ 
fusion gene (Caims & Foster 1991, Foster 1999) and therefore cannot metabolise lactose. After plating 
ona minimal medium with lactose as the only carbon and energy source, they readily revert to lactose 
utilization. After many studies and many proposals it tumed out that this most popular adaptive 
mutation assay system, also called Caims system, is explainable by a standard Darwinian process 
(Andersson & al. 1998, Hendrickson & al. 2002). The explanation of this phenomenon, called the 
amplification-mutagenesis model, presupposes that a non-selective growth before plating enables few 
cells to acquire a simple duplication of the leaky lac mutant allele. After plating, such cells initiate 
slowly growing clones and with further amplification soon dominate the colony. Eventually a celi of 
the clone experiences an adaptive mutation that enables it to utilize lactose asa sole carbon and energy 
source. Selection then favours only stable mutants carrying only the revertant allele; these cells overgrow 
the clone and begin strongly to predominate in the mature revertant colony. 
A recombination between repeated sequences of amplificated alleles releases DNA fragments, 
which are degraded to single DNA strands that induce the SOS system. Induction of the SOS system 
was confirmed in the E. coli strain FC40 (McKenzie & al. 2000). The SOS response induces an error-
prone DNA polymerase DinB (Wagner & al 1999, Wagner & Nohmi 2000, Tang & al. 2000, McKenzie 
& al. 2001) which leads to extra mismatches that exceed the capacity of the mismatch repair system and 
therefore a celi experiences a genome-wide mutagenesis known as hypermutation (Hall 1990, Foster 
1997, Torkelson & al. 1997, Rosenberg & al. 1998, Rosche & al. 1999, Lombardo & al. 1999, Buli & 
al. 2000a,b, Caims 2000, Godoy & Fox 2000a, Godoy & al. 2000b, Foster 2000, Rosenberg 2001, 
Bridges 2001). Temporary hypermutability is therefore a side effect of the lac operon amplification 
(Hendrickson & al. 2002). 
The amplification-mutagenesis model is totally in tune with the neo-darvinistic view ofthe origin 
and dynamics of the genetic variation because here the environment does not direct mutation in specific 
regions, does not have direct effects on the general mutation rale and mutations do not occur in non-
growing stationary cells but in growing subpopulations. But as it will be seen, this model cannot 
explain other cases of the adaptive mutation in cells under a specific selection pressure. 
Adaptive mutation mediated by Mu prophage 
Probably the clearest example showing us the incompleteness of the classical neo-darvinistic 
paradigm is a Mu-mediated adaptive mutation. Bacteriophage Mu is a very notorious mutator phage 
18 Acta Biologica Slovenica, 45 (2), 2002 
that induces mutations in a host genome. Mu element is integrated into the bacterial genome and exists 
inside the celi in a latent prophage state, as long as the genes controlling its lytic pathway are not 
expressed. A phage-encoded repressor maintains the control of the expression. In the case of E. coli the 
repressor of the strain used by Shapiro (1984) was temperature sensitive and the Mu prophage was 
inserted into the araB gene. The prophage represents both a translational and a transcriptional block 
preventing the expression of downstream Jocated genes lacZ and lac Y. When the prophage excises, 
genes lacZY can be fused to an araB gene and the celi encodes a hybrid AraB-LacZ protein. 
Shapiro ( 1984) reported that the Mu-mediated formation of araB-lacZ fusions occurred when cells 
were plated on the selective lactose minimal medium with the arabinose as an inducer. Mutants 
accumulate on the selective media more frequently than during the normal growth. Later it was 
demonstrated that fusions occurred only in the presence of the lactose (Caims & al. 1988), but this was 
subsequently denied. Today it is known that an aerobic starvation could induce the formation of araB-
lacZ fusion s also in the absence oflactose (Mittler & Lenski 1990, Maenhaut-Michel & Shapiro 1994, 
Foster & Caims 1994, Sniegowski 1995). 
But it tumed out that the most staggering point regarding Mu-mediated adaptive mutations was a 
demonstration that the structures of araB-lacZ fusions occurring during aerobic starvation on the 
glucose or on the lactose-arabinose medium differed from each other (Maenhaut-Michel & Shapiro 
1994, Maenhaut-Michel & al. 1997). This means that selective conditions have some influence on the 
mechanism by which the fusions occur. Through the known complexity of the fusion formation 
(Shapiro & Leach 1990, Gomez-Gomez & al. 1997, Lamrani & al. 1999) it becomes clear that we are 
witnessing a multi-step process and not some stochastic individual mutational event. This process 
enables bacterial cells to control their genetic variation according to the environmental conditions. 
Adaptive mutation mediated by insertion sequences 
Another example of a mutational event affected by the selective environment represents adaptive 
mutations in the ebg operon mediated by insertion sequences (IS). IS are less than 2,5kb long segments 
of the DNA that code only elements responsible for their mobility (Mahillon & al. 1998). They are 
known because of their ability to affect different parts of the genome (Naas 1994 & al. , Fedoroff 1999, 
Shneider & al. 2000) and together with transposons and viruses like Mu constitute a storehouse of 
mobile DNA elements. IS not only inactivating genes butare also capable of activating the so-called 
silent or cryptical genes (Reynolds & al. 1981, Hall 1998a, Hall 1999a,b). 
E. coli possesses at least four silent systems for the uptake and metabolism of the beta-glucoside 
sugars (Hall 1998a and references therein). One of them is called the ebg o peron and it is organized in 
the same way as the lac operon. Ebg operon encodes a repressor EbgR and a beta-galactosidase 
EbgAC (Hall 1989). If a strain is deleted for the JacZ, the ebgAC represents the only beta-galactosidase 
in the celi. The expression of the ebgAC is under the control of the repressor ebgR and mutations in the 
ebgR gene allow cells to grow on the lactose or related sugars such as lactu!ose, as a sole source of 
energy and carbon. 
It was shown that during a prolonged starvation on the lactulose 61 % of the growth dependent and 
80% of adaptive mutations in the ebgR gene are mediated by IS (Hall 1999a). 6% of the growth 
dependent and 39% of adapti ve mutations were due to insertions of IS30 in the ebgR, so it appears that 
IS30 transposition may be at least partially directed by some adequate environmental condition (Foster 
1999). It was also demonstrated that only adaptive mutations in the ebgR gene and not growth 
dependent mutations are positively regulated by a two-component regulatory system PhoPQ (Hall 
1998b). 
PhoPQ is a typical two component regulatory system with a regulatory protein in the cytoplasm 
and a sensory kinase located in the membrane (Stock & al. 2000). Primary signal for the PhoQ sensory 
R. Krašovec, l. Jerman: Adaptive mutation: shall we survive bacterial genetic skills 19 
kinase is an extracellular Mg2+ (Sancini & al. 1996, Vescovi & al. 1997). After an extracellular signal 
is sensed by the sensory kinase, an autophosphorylation takes place at the histidine residue of the 
sensory kinase, thus creating a high-energy phosphoryl group. This group is then transferred to an 
aspartate residue in the regulatory protein, which induces a conformational change in the regulatory 
domain. The result is an activation of at least 50 genes in the E. coli (Kasahara & al. 1992, Groisman 
2001). PhoPQ regulated proteins are therefore directly or indirectly responsible for the adaptive 
mutation at the ebgR repressor gene. 
Promoter-creating mutations 
An incredible adaptation to a starving condition stili unexplainable by the classical neo-darvinistic 
approach was shown with Pseudomonas putida cells carrying a plasmid with a promotorless pheAB 
operon (Kasak & al. 1997) that encodes for the enzyme that decompose phenol. Phenol utilising 
mutants that accumulated in a starving cul ture experience base substitutions, deletions and insertions 
of Tn4652 that created active promoters permitting starving P.putida cells to use phenol as a sole 
source of carbon. Mutants accumulate only in the presence of the phenol and plating stationary phase 
cells showed higher rate of mutants' accumulation than plating exponential cells. 
Another exiting and stili unexplainable case of the adaptive mutation is the accumulation of 
resistant mutants during a non-lethal selective pressure from the antibiotic chloramfenicol (Lioy & al. 
2001). During stressful selective conditions sensitive E. coli cells carrying a plasmid with a cam 
promotorless gene, encoding for chloramfenicol acetyl transferase enzyme, experience the insertion of 
the IS 1 0R mobile element into the upstream of the cam gene. Mobile element is transposed from the 
bacterial genome and allowed the bacterial RNA polymerase to efficiently transcript cam gene located 
on the plasmid, which in turn enable celi to survive a non-lethal selective pressure from the antibiotic. 
Discussion 
Ever since the discovery of the transposable elements in 1950 (McC!intock 1984) genome has 
been viewed as a complex, dynamical and highly organised system for storing and processing 
information (Berg & Howe 1989, Shapiro 1991, Shapiro 1992, Shapiro 1999a,b, Fedoroff 1999). 
This means that a bacterial genome is not only a conservative library of tri plet codes for cell's constituent 
elements but also represents a dynamical storage system subject to constant and at least partially 
regulated changes. After this short but thorough review of the current status in the adaptive mutation 
research it is obvious that bacterial genetic change can originate from growth dependent random events 
independently from the environment or from some processes that show sensitivity to specific 
environmental conditions. The big question is which mode of the mutation generation is more 
biologically relevant. 
To answer that we have to bear in mind the fact that within bacterial genomes coexist highly 
mutable 'contingency' genes and so called 'housekeeping' genes with low mutation rates (Moxon & al. 
1994). This means that not ali parts of the bacterial genome are equally mutable and bacterial cells 
obviously have capacities to control and maintain a balance between stability and mutability of DNA 
nucleotide sequences. With this in mind it is not surprising that bacterial cells are known to possess 
many DNA repairing proteins and DNA protecting mechanisms which preserve nucleotide sequences 
(Radman & al. 1999) and protect organism' s needing triplet codes from unpredictable ran dom mutations. 
However, on the other hand, many times bacterial cells exit stressful conditions only if they acquire 
specific genetic information. Celi s that gain such information sometimes in the past through a random 
mutational event are not sensing any stress at ali. But other sensitive cells that are threatened can 
20 Acta Biologica Slovenica, 45 (2), 2002 
accomplish such a task with de novo mutation or with a reorganization of the already existing genetic 
information that may include a horizontal gene transfer or insertions of transposable elements. 
Knowledge gained from the adaptive mutation research demonstrates that during non-lethal stress 
bacterial cells can experience controlled DNA changes influenced by selective environmental conditions. 
Bacterial cells must therefore possess some biochemical systems, active only under stress, capable of 
inducing changes or the reorganisation of genetic information (Shapiro 1991, Shapiro 1992, Shapiro 
1997, Radman & al. 1999, Shapiro 1999a,b, Capy 2000). Biochemical systems such as ROSE 
mutagenesis (Taddei & al. 1995, Taddei & al. 1997), SOS mutagenesis (Fijalkowska & al. 1997) and 
the adaptive mutation process constitute strategic mechanisms for the genetic variation available to 
cells during stressful situations. Which cell's response or which inducible biochemical system will 
predominate depends on many external and interna! signals sensed by the cellular signal transduction 
network. Or in another words, these inducible biochemical systems are regulated by specific controlling 
mechanisms and show sensitivity to environmental conditions sensed through a complex bacterial 
signal transduction network (Shapiro 1999a, Massey & al. 1999, Poster 1999). 
Conclusions 
It may be concluded that both random mutations and controlled genetic changes are present as a 
constituent elements of the bacterial life cycle. Growth dependent random mutations are unpredictable 
and bacterial cells are trying to suppress them by numerous repairing strategies. On the other hand, 
controlled inducible DNA changes are needed during stressful conditions and help many bacterial 
cells to conquer stress and stay alive. Therefore, we must acknowledge that bacterial cells possess 
genetic skills that enable a bacterial celi to gain the needed DNA information to exit stress s tate. We can 
say that the bacterial genome is evolved to cope with the predictable and also unpredictable challenges 
that come out from the environment (Caporale 1999). 
Last but not least, let us discuss some of the consequences of such bacterial genetic skills on our 
human lives. Adaptive mutations were demonstrated to be important in a development of antibiotic 
resistance mutations (Riesenfeld & al 1997, Alonso & al. 1999, Martinez & Baquero 2000, Karunakaran 
& Davies 2000) and may also provide models for human cancer (Strauss 1992, Hall 1995 Cairns 
1998). Needless to say, these two phenomena represent a gigantic problem for modem human society. 
Butat the same tirne they present a high motivation for researchers to try to understand more deeply 
and thoroughly the nature of bacterial genetic variation. Therefore it rests on us, researchers, to stay 
open minded and admit that bacterial cells possess genetic skills that go beyond the ideas of the 
ordinary neo-darwinian evolutionary doctrine and should not be underestimated. 
Acknowledgements 
This work was supported by grant L3-24 l 6-0487-00 from the Slovenian Ministry of Education, 
Science and Sport. 
R. Krašovec, I. Jerman: Adaptive mutation: shall we survive bacterial genetic skills 21 
References 
ALONSO A., E. CAMPANARIO & J.L. MARTINEZ 1999: Emergence of multidrug-resistant mutants is 
increased under antibiotic selective pressure in Pseudomonas aeruginosa. Microbiology-
(UK) 145: 2875-2862. 
ANDERSSON D. I., E. S. SLECHTA & J. R. Rom 1998: Evidence that gene amplification underlies adaptive 
mutability ofthe bacterial operon. Science 282: 1133-1135. 
BERG D. E. & M. M. HoWE (ed.) 1989: Mobile DNA. ASM Press, Washington D.C. 
BRIDGES B. A. 2001: Hypermutation in bacteria and other cellular systems. Philos. Trans. R. Soc. Lond. 
B 356: 29-39. 
BuLL H. J., G. J. McKENZIE, P. J. HASTINGS & S. M. RosENBERG 2000a: Response to John Cairns: 
Contribution of transiently hypermutable cells to mutation in stationary phase. Genetics 156: 
925-926. 
BuLL H. J., G. J. McKENZIE, P. J. HASTINGS & S. M. RosENBERG 2000b: Evidence that stationary-phase 
hypermutation in the E.coli chromosome is promoted by recombination. Genetics 154: 
1427-1437. 
CAIRNS J. 2000: The Contribution ofbacterial hypermutators to mutation in stationary phase. Genetics 
156: 923. 
CAIRNS J. & P.L. FosTER 1991: Adaptive reversions of a frameshift mutation in Escherichia coli. 
Genetics 128: 695-701. 
CAIRNS J. 1998: Mutation and cancer: The antecedents to our studies of adaptive mutation. Genetics 
148: 1433-1440. 
CAIRNS J., J. OvERBAUGH & S. MILLER 1988: The origin of mutants. Nature 335: 142-145. 
CAPORALE L.H. 1999: Chance favors the prepared genome. Ann. N.Y. Acad. Sci. 870: 1-21. 
CAPY P., G. GASPERI, C. BIEMONT & C. BAZIN 2000: Stress and transposable elements: co-evolution or 
useful parasites? Heredity 85(2): 101-106. 
CAVALLI-SFORZA L. L. & J. LEDERBERG 1956: Isolation of pre-adaptive mutants in bacteria by sib 
selection. Genetics 41: 367-381. 
DARWIN C. 1859: The origin of species., J. Murray, Albemarle Street, London 
FEDOROFF N. V. 1999: Transposable elements asa molecular evolutionary force. Ann. N.Y. Acad. Sci. 
870: 251-264. 
FIJALKOWSKA I. J ., R. L. DuNN & R.M. SettAAPER 1997: Genetic requirements and mutational specificity 
of the E.coli SOS mutator activity. J. Bacteriol. 179: 7435-7445. 
FosTER P. L. & J. CAIRNS 1992: Mechanisms of directed mutation. Genetics 131: 783-789. 
FosTER P. L. & J. CAIRNS 1994: The occurence of heritable Mu excisions in starving cells of E.coli. 
EMBO J. 13: 5240-44. 
FosTER P. L. 1997: Non-adaptive mutations occur on the F' episome during adaptive mutation conditions 
in Escherichia coli. J. Bacteriol. 179: 1550-1554. 
FosrnR P. L. 1999: Mechanisms of stationary phase mutation: A decade of adaptive mutation. Annu. 
Rev. Genet. 33: 57-88. 
FosrnR P. L. 2000: Adaptive mutation: implications for evolution. BioEssays 22: 1067-1074. 
GALITSKI T. & J. R. Rom 1996: A search for a general phenomenon of adaptive mutability. Genetics 
143: 645-659. 
GoooY V. G. & M. S. Fox 2000a: Transposon stability and a role for conjugational transfer in adaptive 
mutability. Proc. Natl. Acad. Sci. U. S. A. 97: 7393-7398. 
Goooy V. G., F. S. GIZATULLIN & M.S. Fox 2000b: Some features of the mutability of bacteria during 
nonlethal selection. Genetics 154: 49-59. 
GoMEz-GoMEZ J. M., J. BLAZQUEZ, F. BAQUERO & J.L. MARTINEZ 1997: H-NS and RpoS regulate 
emergence of LacAra+ mutants of E.coli MCS2. J. Bacteriol. 179: 4620-4622. 
22 Acta Biologica Slovenica, 45 (2), 2002 
GROISMAN E. A. 2001: The pleiotropic two-component regulatory system PhoP-PhoQ. J. Bacteriol. 
183: 1835-1842. 
HALL B. G. 1988: Adaptive evolution that requires multiple spontaneus mutations. I. Mutations 
involving an insertion sequence. Genetics 120: 887-897. 
HALL B. G. 1990: Spontaneous point mutations that occur more often when advantageous than when 
neutral. Genetics 126: 5-16. 
HALL B. G. 1995: Adaptive mutations in E.coli as a model for the multiple mutational origins of 
tumors. Proc. Natl. Acad. Sci U.S.A. 92: 5669-5673. 
HALL B. G. 1998a: Activation of the bgl operon by adaptive mutation. Mol. Biol. Evo!. 15(1): 1-5. 
HALL B. G. 1998b: Adaptive mutagenesis at ebgR is regulated by PhoPQ. J. Bacteriol. 180: 2862-
2865. 
HALL B. G. 1998c: Adaptive mutagenesis: a process that generates almost exclusively beneficial 
mutations. Genetica 102/103: 109-125. 
HALL B. G. 1999a: Spectra of spontaneous growth-dependent and adaptive mutations at ebgR. J. 
Bacteriol.181: 1149-1155. 
HALL B. G. 1999b: Transposable elements as activators of cryptic genes in E.coli. Genetica 107: 181-
187. 
HALL B. G., P.W. BETTS & J.C WoorroN 1989: DNA sequence analysis of artificially evolved ebg 
enzyme and ebg repressor genes. Genetics 123: 635-648. 
HARRIS S. B., G. FENG, K. J. Ross, R. SmHu, C. THULIN, S. LoNGERICH, S. K. SzIGETY, M. E. WINKLER 
& S.M. RosENBERG 1997: Mismatch repair protein MutL becomes limiting during stationary-
phase mutation. Genes Dev. 11: 2426-2437. 
HENDRICKSON H., E. S. SLECHTA, U. BERGTHORSSON, D. I. ANDERSSON & J. R. Rorn 2002: Amplification-
mutagenesis: Evidence that "directed" adaptive mutation and general hypermutability result 
from growth with a selected gene amplification. Proc. Natl. Acad. Sci. U. S. A. 99: 2164-
2169. 
JAYARAMAN R. 2000: Modulation of allele leakiness and adaptive mutability in E.coli. J. Genetics 79(2): 
55-60. 
JAYARAMAN R. 1995: Leakiness of genetic markers and susceptibility to post-plating mutagenesis in 
E.coli. J. Genetics 74(3): 85-79. 
KARUNAKARAN P. & J. DAvIEs 2000: Genetic antagonism and hypermutability in Mycobacterium 
smegmatis. J. Bacteriol. 182: 3331-3335. 
KASAHARA M., A. NAKATA & H. SHINAGAWA 1992: Molecular analysis of the Escherichia coli phoP-
phoQ operon. J. Bacteriol. 174: 492-498. 
KASAK L., R. HoRAK & M. K1v1sAAr 1997: Promoter-creating mutations in Pseudomonas putida: A 
model system for the study of mutation in starving bacteria. Proc. Natl. Acad. Sci. U.S.A. 
94: 3134-3139. 
LAMRANI S., C. RANQUET, M-J. GAMA, H. NAKAI, J. A. SHAPIRO, A. ToussAINT & G. MAENHAUT-MICHEL 
1999: Starvation-induced Mucts62-mediated coding sequence fusion: a role for ClpXP, 
Lon, RpoS and Crp. Mol. Microbiol. 32: 327-343. 
LEDERBERG J. & E. M. LEDERBERG 1952: Replica plating and indirect selection ofbacterial mutants. J. 
Bacteriol. 63: 399-406. 
LENSKI R. E. & J. E. MITTLER 1993: The directed mu tati on controversy and neo-darwinism. Science 
259: 188-194. 
LENSKI R. E., M. SLATKIN & F. J. A YALA 1989: Mutation and selection in bacterial populations: Altematives 
to the hypothesis of directed mutation. Proc. Natl. Acad. Sci. U. S. A. 86: 2775-2778. 
LIOY M., S. DABIZZI, S. AMMANATO, A. CACIOTTI, L. CioNI & R. FANI 2001: Activation of cam 
promotorless gene by ISRIO transposition in an Echerichia coli population under stress 
conditions. Ann. Microbiol. 51: 225-233. 
R. Krašovec, I. Jerman : Adaptive mutation: shall we survive bacterial genetic skills 23 
LOMBARDO M . J., J. ToRKELSON, H. J. Bu:..L, G. J. McKENZIE & S. M . RosENBERG 1999: Mechanisms of 
genome-wide hypermutation in stationary phase. Ann. N.Y. Acad. Sci. 870: 275-289. 
LuRIA S. E. & M. DELBRDcK 1943: Mutations of bacteria from virus sensitivity to virus resistance. 
Genetics 28: 491-511. 
MADIGAN M. T., J. M. MARTINKO & J. PARKER (ed.) 2000: Brock Biology of microorganisms. Ninth 
edition. Prentice Hall, NY. 
MAENHAUT-MICHEL G. & J. A. SHAPIRO 1994: The roles of starvation and selective substrates in the 
emergence of araB-lacZ fusion clones. EMBO J. 13: 5229-5239. 
MAENHAUT-MICHEL G., C. E. BLAKE, D. R. F. LEACH & J. A. SHAPJRO 1997: Different structures of 
selected and unselected araB-lacZ fusions. Mol. Microbiol. 23: 1133-1145. 
MAHILLON J. & M. CHANDLER 1998: Insertion sequences. Microbiol. Mol. Biol. Rev. 62(3): 725-774. 
MARTINEZ J. L. & F. BAQUERO 2000: Mutation frequencies and antibiotic resistance. Antimicrob. 
Agents. Chemother. 44: 1771-1777. 
MASSEY R. C., P. B. RAINEY, B. J. SHEEHAN, KEANE O. M. & C. J. DoRMAN 1999: Environmentally 
constrained mutation and adaptive evolution in Salmonella. Curr. Biol. 9: 1477-1480. 
McCLINTOCK B. 1984: The significanse of responses of the genome to challenge. Science 226: 792-80 l. 
McKENZIE G. J., P. L. LEE, M. J. LOMBARDO, P. J. HASTINGS & S. M. RosENBERG 2001: SOS mutator 
DNA polymerase IV functions in adaptive mutation and not adaptive amplification. Mol. 
Cell. 7: 571-579. 
McKENZIE G. J. , R. S. HARRIS, P. L. LEE & S. M. RosENBERG 2000: The SOS response regulates 
adaptive mutation. Proc. Natl . Acad. Sci. U. S. A. 97: 6646-6651. 
MITTLER J. E. & R. E. LENSKI 1990: New data on excisions of Mu from E.coli MCS2 cast doubt on 
directed mutation hypotheses. Nature 344: 173-175. 
MoxoN E. R. , P. B. RAINEY, A. M. NowAK & R. E. LENSKI 1994: Adaptive evolution of highly mu table 
loci in pathogenic bacteria. Curr. Biol. 4: 24-33. 
NAAS T., M. BLOT, W. M. FITCH & W. ARBER 1994: Insertion sequence-related genetic variation in 
resting E.coli K-12. Genetics 136: 721-730. 
RADMAN M, I. MATIC & F. TADDEI 1999: Evolution of evolvability.Ann. N.Y.Acad. Sci. 870: 146-155. 
REYNOLDSA. E., J. FELTON & A. WRIGHT 1981: Insertion of DNA activates the cryptic bgl operon in 
E.coli Kl2. Nature 293: 625-629. 
R1ESENFELD C., M. EvERETT, L.J.V. PmDocK & B.G. HALL 1997: Adaptive mutations produce resistance 
to ciprofloxacin. Antimicrob. Agents Chemoter. 41: 2059-2060. 
RoscHE W. A. & P. L. FosTER 1999: The role of transient hypermutators in adaptive mutation in 
Escherichia coli. Proc. Natl. Acad. Sci. U.S.A. 96: 6862-6867. 
RosENBERG S. M. 2001: Evolving responsively: adaptive mutation. Nat. Rev. Genet. 2: 504-515. 
RosENBERG S. M., C. THULIN & R. S. HARRIS 1998: Transient and heritable mutators in adaptive 
evolution in the lab and in nature. Genetics 148: 1559-1566. 
RYAN F. J. & L. K. W AINWRIGHT 1954: Nuclear segregation and the growth of the clones of spontaneous 
mutants of bacteria. J. Gen. Microbiol. 11: 364-379. 
RYAN F. J. 1955: Spontaneus mutations in non-dividing bacteria. Genetics 40: 726-738. 
ScHNEIDER D., E. DuPERCHY, E. CouRsANGE, R. E. LENSKI & M. BLOT 2000: Long-term experimental 
evolution in Escherichia coli. IX. Characterization of insertion sequence-mediated mutations 
and rearrangements. Genetics 156: 477-488. 
SHAPIRO J. A. & D. LEACH 1990: Action of a transposable element in coding sequence fusions . Genetics 
126: 293-299. 
SHAPIRO J. A. 1984: Observations on the formati on of clones containing araB-lacZ cistron fusions. 
Mol. Gen. Genet. 194: 79-90. 
SHAPIRO J. A. 1991: Genomes as smart systems. Genetica 84: 3-4. 
SHAPIRO J. A. 1992: Natura) genetic engineering in evolution. Genetica 86: 99-111. 
24 Acta Biologica Slovenica, 45 (2), 2002 
SHAPIRO J. A . 1995: Adaptive mutation: Who's really in the garden? Science 268: 373-374. 
SHAPIRO J. A. 1997: Genome organization, natura! genetic engineering and adaptive mutation. Trend 
Genet. 13(3): 98-104. 
SHAPIRO J. A. 1999a: Genome system architecture and natura! genetic engineering in evolution. Ann. 
N .Y.Acad. Sci. 870: 23-35. 
SHAPIRO J. A. 1999b: Transposable elements as the key to a 21 st century view of evolution. Genetica 
107: 171-179 
SNIEGOWSKI P. D. 1995 : A test of the directed mutation hypothesis in E.coli MCS2 using replica 
plating. J. Bacteriol. 177: 1119-1120. 
SoNCINI F. C. & E. A. GROISMAN 1996: Two component regulatory systems can interact to process 
multiple environmental signals. J. Bacteriol. 178: 6796-6801. 
SrncK A. M ., V. L. ROBINSON, P. N . GouDREAU 2000: Two-component signal transduction. Annu. Rev. 
Biochem. 69: 183-215 . 
STRAUSS B. S. 1992: The origin of point mutations in human tumor cells. Cancer Res. 52: 249-253. 
TADDEI F., l. MATIC & M. RADMAN 1995: cAMP-dependent SOS induction and mutagenesis in resting 
bacterial populations. Proc. Natl. Acad. Sci. U. S. A. 92: 11736-11740. 
TADDEI F., J.A. HALLIDAY, I. MATIC & M. RADMAN 1997: Genetic analysis of mutagenesis in aging 
E.coli colonies. Mol. Gen. Genet. 256: 277-281. 
TANC M ., P. PHAM, X. SttEN, J .S . TAYLOR, M . O ' DoNNEL, R. WooDGATE & M .F. GOODMAN 2000: Roles 
of E .coli DNA polymerases IV and V in lesion-targeted and untargeted SOS mutagenesis. 
N ature 404: 1014-1018. 
ToRKELSON J., R . S. HARRIS, M.J. LOMBARDO, J. NAGENDRAN, C. THULIN & S.M. RosENBERG 1997: 
Genome-wide hypermutation in a sub-population of stationary-phase cells underlies 
recombination-dependent adaptive mutation. EMBO J. 16: 3303-3311 . 
Vescov1 E. G ., Y. M. AYALA, E. D1 CERA & E .A. GROISMAN 1997: Characterization of the bacterial 
sensor protein PhoQ. Evidence for distinct binding sites for Mg2+ and Ca2+. J. Biol. Chem. 
272: 1440-1443. 
WAGNER J. & T. NoHMI 2000: Escherichia coli DNA polymerase IV mutator activity: genetic requirements 
and mutational specificity. J. Bacteriol. 182: 4587-4595 . 
WAGNER J ., P. GRuz , S . R . K1M, M. YAMADA, K. MATSUI, R.P. FuCHs . & T. NoHMI 1999: The dinB gene 
encodes a novel E.coli DNA polymerase, DNA pol IV, involved in mutagenesis. Mol. Celi. 
4: 281-286. 
ACTA BIOLOGICA SLOVENICA 
LJUBLJANA 2002 Vol. 45, Št. 2: 25 - 34 
Sprejeto (accepted): 2002-05-27 
Aquatic macrophytes of the mountain lake Krnsko jezero, Slovenia 
Vodni makrofiti Krnskega jezera, Slovenija 
Olga URBANC-BERČIČ, Alenka GABERŠČIK, Milijan ŠIŠKO, Anton BRANCELJ 
National Institute of Biology, Večna pot 111, I 000 Ljubljana, Slovenia; 
Tel.+386 I 4233388; E-mail : olga.urbanc@nib.si 
Abstract. The macrophyte vegetation in the mountain lake Krnsko jezero has 
been monitored since 1991. Five species of aquatic macrophytes, occupying different 
depths, were present in the lake. Changes in depth distribution were detected due to 
accelerated eutrophication caused by inputs of nutrients from the watershed. In the 
period between 1997 and 1998 repeated earthquakes additionally influenced the 
processes in the lake Krnsko jezero, by increased input of matter. The occasional 
phytoplankton blooms reduced water transparency and consequently disturbance to 
growth, development and distribution of anchored macrophytes occurred. In 1998 
macrophytes spread down to 7.0 m, reaching the maximum at 5.8 kg DW m·2 at 2 m 
depth. The total primary production in the lake was estimated at 4991 kg organic matter 
per year out of which Chara and Potamogeton species presented 95.5 % and 0.6 %, 
respectively. 
Key words: aquatic macrophytes, chlorophyll a, lake Krnsko jezero, mountain 
lake, nutrients, organic matter, primary production 
Izvleček. Makrofite v Krnskem jezeru spremljamo od 199 l leta. V jezeru je 
prisotnih pet vrst podvodnih rastlin, ki uspevajo na različnih globinah. Zaradi pospešene 
evtrofikacije, kije odraz povečanega vnosa nutrientov iz zaledja, se globina uspevanja 
spreminja. Med letoma 1997 in 1998 je bila serija potresov, zaradi česar se je vnos 
snovi iz zaledja povečal in dodatno vplival na procese v jezeru. Občasno cvetenje 
fitoplanktona je zmanj šalo prosojnost vode, kar se je odrazilo na rasti , razvoju in 
razporeditvi submerznih makrofitov. Tako so v letu 1998 makrofiti segali do globine 7 
m, njihova največja biomasa je bila 5,8 kg suhe teže m·2 na globini 2 m. Celotna 
primarna produkcija organske snovi v jezeru je bila ocenjena na 4991 kg leto-1, pri 
čemer je bil delež parožnic 95,5 %, delež dristavcev pa le 0,6 %. 
Ključne besede: vodni makrofiti , klorofil a, Krnsko jezero, gorsko jezero, 
nutrienti, organska snov, primarna produkcija 
26 Acta Biologica Slovenica, 45 (2), 2002 
Introduction 
Mountain lakes, usually located in remote natura! environment, are becoming interesting sites of 
ecological research for two reasons inter alta: they are relatively small and they are sensitive to 
different influences from the environment. The alpine lake Krnsko jezero is an eutrophic lake (GABERŠČIK 
& URBANC-BERČIČ 1996). In the last century some events had put an evident mark on this ecosystem. 
During the First World War the wider area was the centre of severe battles that had resulted in increased 
input of matter from the watershed. Latter on two species of fish were introduced in the lake; in late 
twenties Salvelinus alpinus (Linnaeus, 1758) and two decades latter Phoxinus phoxinus (Linnaeus, 
1758) (BRANCEU & al.1997). Other human activities, pasturing and mountaineering at most, presented 
an influential activity, the latter being intensified in the last decade. The influence of these activities had 
been closely studied in the nearby lake (BRANCEU & al. 2000). 
The first investigation ofbasic geographic and hydrobiologic characteristics was carried out in the 
late fifties (GAMS 1962), but no record on macrophytes was made until 1988 (BLAŽENČIČ & al. 1990). 
In 1996 the systematic research of 14 alpine lakes started as an addition to the previous 5-year 
monitoring. The study had revealed that the most abundant aquatic vegetation was in the lake Krnsko 
jezero (GABERŠČIK & URBANC-BERČIČ, 1996). Comparable studies to ours on primary production in 
mountain lakes was not found but in our case it was evident that macrophytes contributed a great deal 
to the production ofthe lake. Researches on lowland lakes, rich with aquatic vegetation showed, that 
macrophytes presen! an important factor maintaining the stability of the ecosystem (Oz1MEK & al. 1990, 
R0RSLETT 1991, RASPOPOV & al. 1996). 
The present study was carried out in order to evaluate the abundance and production of 
anchored macrophytes, related to the main trophic parameters and their contribution to the primary 
production of the lake. 
Material and methods 
Site description 
The lake Krnsko jezero is located well bellow the timberline at the altitude of 1383 m. It is of 
glacial origin. Lake is the largest and the deepest in the Triglav National Park, covering the area of 
49600 m2• The lake is 17 .6 m deep (Fig. 1 ). At the highest water leve! the to tal vol ume of 446700 
m3 of water accumulates in the lake. At heavy rains fluctuations of water leve! could be as high as 4 
m, reaching the normal leve! within two days. The lake is ice covered usually from November to 
May. During the summer it is stratified, with late spring (after ice-cover melting) and late autumn 
homothermy. The nearby watershed is diverse, formed of stones, gravel, rocks, meadows and 
pastures. The wider area is geologically unstable and some severe earthquakes with soil avalanches 
have caused strong disturbances in watershed and increased the input of matter. 
Water chemistry 
The sampling of water for chemical analyses and chlorophyll a was carried out with Van Dom 
sampler. Samples were collected on the vertical profile every 2.5 m (O m, 2.5 m, 5 m, 7.5 m, JO.O m, 
12.5 m and above bottom) in monthly intervals in the ice-free period from May 1996 to the end of 
1999. Electric conductivity was measured in laboratory at 25 °C with conductivity meter (MA 5950, 
Iskra, Slovenia) and concentration of oxygen was measured in situ with an oxygenmeter (WTW Oxi 
320, Germany). To evaluate chemical variables analytical methods according to APHA (1992) were 
applied. Ali spectrophotometric measurements were performed with spectrophotometer Perkin Elmer 
(UV NIS, Lambda 12). The temperature was measured with a thermistor (Iskra, Slovenia). Chlorophyll 
O. Urbanc-Berčič, A. Gaberščik, M. Šiško, A. Brancelj: Aquatic macrophytes of the mountain ... 27 
a concentration was determined by photometry (GOLTERMAN & al. 1978, WETZEL & LIKENs 1995). 
Biomass of phytoplankton was calculated on the basis of chlorophyll a concentrations (T ALLING 
1975). Transparency was estimated by means of Secchi disc. The physical and chemical data are 
presented as median, minimum and maximum of total measured values. 
,__ ______ \00 m 
Figure 1: Bathimetric map of the lake Krnsko jezero 
Slika 1: Batimetrična karta Krnskega jezera 
Macrophyte survey 
The survey of submersed vegetation was carried out from 1991 to 1999 along 20 permanent 
transects. The shore line was divided into the sections of different lengths, based on the uniformity 
of morphological characteristics (bathimetric map), water depth and substrate types. For graphic 
presentation the homogenous sections were additionally divided into the fina! 31 sections, with the 
length of 35±2 m (Fig. 2). Abundance of the observed macrophytes was estimated semi-
quantitatively using a 5-level scale (!- present, 2 - rare, 3 - common, 4 - abundant, 5 - predominant) 
(MELZER 1992, PALL & JANAUER 1995). The survey was made from the boat using depthmeter, view 
box and sampling rake. In September 1998, macrophyte community was surveyed by scuba-diving 
on six representative transects. The total biomass of macrophytes was collected in the area of 0.1 m2 
on the depths of 1, 2, 3, 4 and 7 m, washed thoroughly to remove periphyton, oven dried at 105 °C 
and weighted. The ratio of organic matter was determined by ignition at 550 °C following the 
method from APHA (1992). 
Results and Discussion 
As the majority oflakes world-wide the lake Krnsko jezero is subjected to accelerated eutrophication 
due to diverse reasons. The disturbances to the lake were caused by the First World War, introduction 
of fish, traditional pasturing, earthquakes and mountaineering (GABERŠČIK & al. 1997). In the period 
between 1997 and 1998 after repeated earthquakes the sedimentation rate in the lake increased. The 
analyses of the material in the sediment traps revealed that the annual amount of gathered matter was 
increasing from 0.84 g DW m 2 in 1997, 1.01 g DW m·2 in 1998 to 3.68 g DW m 2 in 1999 (MURI&. 
al. 2002). 
28 
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O) 
Cl 
10 
12,5 t'Q\,'.#Ml\ 
15 JOJ 
2,5 
5 
7,5 
10 
12,5 
15 
O 100 200 300 
Tota! P mg m ·3 
o 10 20 
Water temperature °C 
Acta Biologica Slovenica, 45 (2), 2002 
7,5 
10 
12,5 
15 
o 1000 2000 3000 
Tota! N mg m·3 
12,5 <:-::"/~!/"< 
o 5 10 15 
Oxygen gm·3 
7 ,5 ~:fq:it':i~~.-:•.,-. .. , ..... , .. -.-.. ... t ... w.·.·;-'j 
10 
o 
2,5 
5 
7,5 
10 
12,5 
15 
o 6 12 18 
Chlorophylla mg m·3 
"._·ft> --------' 
··--·~x=-:::i 
O 100 200 300 400 
Conductivity µS cm·1 
Figure 2: Main physical and chemical parameters measured in the depth profile of the lake Krnsko 
jezero in the period from 1996 to 1999- Median values, minima and maxima are presented_ 
Slika 2: Glavni fizikalni in kemični parametri merjeni na globinskem profilu v Krnskem jezeru, v 
obdobju od 1996 do 1999 leta_ Pedstavljene so mediane, minimalne in maximalne vrednosti_ 
o. Urbanc-Berčič, A. Gaberščik, M. Šiško, A. Brancelj: Aquatic macrophytes of the mountain... 29 
Concentrations of nutrients in water column measured in a period from 1996 to 1999 varied to a great 
extent (Fig. 2). The median values of nutrients increased with depth, the total phosphorus ranging from 
21 to 44 mg m·3 and the total nitrogen from 915 to 1463 mg m·3. The concentrations of total phosphorus 
in the water column differed a lot, exhibiting the lowest variability in the bottom sam ples, due to releasing 
of phosphorus from the sediment under anaerobic conditions. The concentrations of total nitrogen and 
dissolved solids rneasured as electric conductivity showed less variability than total phosphorus. 
The introduction of fish influenced the food web in the lake, and consequently only one zooplankton 
species, Cyclops vicinus (Uljanin 1875) was present. In the nearby lakes zooplankton was stili diverse 
and up to 6 species could be found (BRANCEU & al. 1997). The reduction of zooplankton diversity 
resulted further in the increase of chlorophyll a concentration and in lower water transparency (Tab. 1 ). 
During the monitoring Secchi depth varied in the range from 2.5 m to 1 1 m, median value being 4 m. 
The concentration of chlorophyll a asa measure of phytoplankton production was rather constant in 
the upper layers, reaching median value about 3 mg m·3• The highest concentration was measured at the 
bottom layer (median value 13 mg m-3), likely due to alga] sedimentation, while the variability from 1 
to 17 mg m·3 was deterrnined at the depth of 7.5 m, where thermocline usually occurred. 
Table 1: Data on minimal water tranparencies and maximal depth distribution of Characeae in the lake 
Krnsko jezero from 1994 to 1999; n = 3 - 6, * BLAŽENčrč & al. (1990). 
Tabela 1: Najnižje vrednosti prosojnosti vode in največja globina uspevanja parožnic (Characeae) v 
Krnskem jezeru v obdobju od 1994 do 1999 (n=3-6). Podatek* BLAžENČIČ & al. (1990). 
Year 
Secchi depth (m) 
Maximal depth 
distribution (m) 
1988 1991 1992 1993 1994 1995 1996 1997 1998 1999 
10.0* 8.0 8.0 8.0 8.0 
4.2 
7.0 
3.5 
7.0 
3.0 
7.5 
2.5 
7.0 
2.5 
7.5 
By preventing the penetration of light, planktonic algae posed depth limits for anchored macrophytes. 
In spite of this phenomenon the littoral area of the lake Krnsko jezero is large. We defined it by the 
extension of macrophyte community (WETZEL 1990) on the basis of bathimetric map. In 1998 it 
presented about 19600 m2, that is nearly 40 % of the projected lake area. Five species of submersed 
macrophytes i.e. Chara delicatula Ag., Chara contraria f capillacea Mig., Potamogeton alpinus 
Balbis, Potamogeton pusillus L. and Batrachium trichophyllum Chaix. colonising different depths, 
were determined in the lake (Fig. 3). The community is considered as flora! rich in comparison to other 
13 Slovene alpine lakes (GABERŠČIK & URBANC-BERČIČ 1996). This coincides with the data ofnorthern 
European lakes elaborated by R0RSLETI (1991) which revealed that upland lakes exhibited proportionally 
fewer species than lowland sites. 
During the monitoring the maximal depths colonised by macrophytes oscillated (Tab. 1) showing 
trend of gradual reduction. In 1988 Chara delicatula spread down to l O m. The maximum depth of this 
species was 8 m from 1991 to 1994 and 7 - 7.5 m since then (BLAŽENČic & al. 1990, GABERŠČIK & URBANC-
BERČIČ 1996). The competition for light and nutrients with filamentous and planktonic algae was estimated 
to be the main reason for this feature. BuNoow ( 1992a,b) found out that charophytes were sensitive to the 
reduction of light more than to the increase of total phosphorus, as it was indicated in the previous studies 
(FoRSBERG 1964). LEHMANN & LACHAVANNE (1999) compared the relevance of two indices to retine 
changes in macrophyte community in Lake Geneva. They demonstrated that saprobic index based on 
organic matter inputs, was more closely related to subtle vegetation changes than index, based on 
nutrients load. Our survey indicated that increased inputs detected in sediment traps, originating from the 
disturbed watershed supposed to be a trigger for changing of submersed vegetation. Survey of 116 high 
mountain Pyrenean lakes, led by GACIA & al. ( 1994) pointed out two environmental variables, vegetation 
coverage of the catchment and altitude as the main factors that influence the distribution and composition 
30 
s o,~~----
..5 , ✓ 
& ~ Q -2 " C> 
5 
5 
Acta Biologica Slovenica, 45 (2), 2002 
Potamogeton alpinus 
~~) { o 1 D--<J ( : c~J 
10 
10 
10 
15 20 25 30 
Potamogeton pusillus 
)tli(~~ 
15 20 25 30 
Chara contraria 
15 
Chara delicatula 
Filamentous algae 
15 
Locations 
20 25 30 
20 25 30 
Figure 3: Depth distribution and abundance (rated from I to 5) of macrophytes in 31 sections (length 
35±2 m) in the littoral of the lake Krnsko jezero in September 1998. Section 1 isto the right of the lake 
outflow (counter clockwise). 
Slika 3: Globinska razporeditev in številčnost makrofitov, ocenjena od 1 do 5, na 31 odsekih litorala v 
Krnskem jezeru, september 1998. Odseki ( 1 je desno od iztoka) potekajo v obratni smeri urinega kazalca. 
o. Urbanc-Berčič, A. Gaberščik, M. Šiško, A. Brancelj: Aquatic macrophytes of the mountain... 31 
of macrophytes. These two determinants reflect potential inputs in whole while a nutrie:1t availability in 
the water as a relevant variable for the primary producers, depends on the processes in lake, as well. 
In 1998, when a detailed survey was made by scuba-di vers, the maximum biomass of macrophytes 
was determined between the depths of2 and 3 m (the estimation of abundance was 5) even lhough the 
continuous stands of stoneworts spread to the depth of7.5 m (Fig. 3). Plants were extremely elongated 
reaching the length up to 1 m at the depth of 3 m. The prevailing species in the lake were Chara 
delicatula and Chara contrariaf capillacea. In 1998 their share per lake was 2700 and 2068 kg of 
organic matter, respectively. The form er reached up to 3 kg DW m·2 on lhe depth of 3 m. The highest 
dry weight 5.8 kg m·2 of Chara contrariaf capillacea was determined on the depth of 2 m (Fig. 4). 
o 2 4 6 8 
Biomass kg DW 1112 
Figure 4: Depth distribution of macrophytes biomass in September 1998 (white column - average, 
grey column - maximum value). Figures present the values calculated from the macrophytes' samples 
obtained by scuba diving on 6 transects. 
Slika 4 : Globinska razporeditev biomase makrofitov v septembru 1998 (bel stolpec so povprečne, siv 
stolpec maximalne vrednosti). Številke predstavljajo biomaso, preračunano iz vzorcev makrofitov, 
zbranih s pomočjo potapljačev na 6 transektih. 
Both species contributed a lot to the total dry weight which was determined at 17537 kg. The value 
corresponds to 4991 kg of organic matter. Converted to energy content it means that about 108 KJ were 
integrated in the ecosystem from solar energy. With such a production the lake reaches the leve! of 
fertile ecosystems (OZIMEK & al. 1990). The predominance of stonewort species which function asa 
stabiliser of the sediment is beneficial for the system resilience conceming inputs from the watershed 
i.e. nutrients, as well as particulate matter. The other two submersed macrophytes Potamogeton alpinus 
and Potamogeton pusillus were not abundant and their biomass presented a negligible share of 21 and 
33 kg DW per lake in 1998 (Fig.S) but having the highest organic content (Fig. 6). In spite of low 
production pondweeds presen! a factor stabilising the shallow, sensitive parts of the littoral and 
promoting sedimentation of fine particles. The macrophytic biomass distribution in the lake Krnsko 
jezero reflects the characteristics of the lake, i.e. basin configuration, high availability of nutrients, 
waler leve! fluctuations and modest light conditions . Stoneworts that have lower light demands for 
photosynthesis (BLINDOW l 992a,b) express a competitive advantage over higher submersed plants. 
Occasionally, when the runoff of nutrients from the watershed was increased, the biomass of filamentous 
and planktonic algae increased. Even though planktonic algae presented a minor share in the primary 
production of the lake, they affected macrophytes (LACHAVANNE 1985, HouGH & al. 1989). Calculated 
32 Acta Biologica Slovenica, 45 (2), 2002 
to the whole lake volume their average concent~ation would be 4 mg m·3, which is equivalent to the 
production of 92 kg organic matter per lake, presenting 1.8 % of the lake primary production (Fig. 5). 
In 1998, nearly the same share of primary production was contributed from filamentous algae. They 
Cha del 
-H,4% 
Pot alp 
0,1% 
Cha con 
54,1% 
Figure 5. Partition oftotal organic matter among different primary producers in the lake Krnsko jezero 
in September 1998 when the total primary production was estimated to 4991 kg DW of organic matter. 
Periphyton biomass is excluded. 
Slika 5: Delež organske snovi med različnimi primarnimi producenti v Krnskem jezeru, september 1998. 
Celotna primarna produkcija je bila ocenjena na 4991 kg suhe organske snovi brez biomase perifitona. 
H 
~ ro s 
(.) 
·a 
ro 
e!l o 
~ 
100 
80 
60 
40 
20 
o 
'°"'•.· 0,.s 
¾ 
~ 
Figure 6: The ratio of organic matter in the biomass of different primary producers from the lake 
Krnsko jezero. 
Slika 6: Razmerje organske snovi v biomasi različnih primarnih producentov v Krnskem jezeru. 
appeared mainly in the shallow water competing with pondweeds. On the sites with steep slopes where 
the conditions for anchoring were not favourable, higher plants and stoneworts were scarce. On these 
locations filamentous algae were not in a position of direct competitors (locations from 18 to 30 on Fig. 3). 
Conclusions 
In the Jake Krnsko jezero accelerated eutrophication resulted in high primary production that 
presented very high absorbance of solar energy into the lake. In 1998, littoral presented 40 % of a 
O. Urbanc-Berčič, A. Gaberščik, M. Šiško, A. Brancelj: Aquatic macrophytes of the mountain... 33 
projected Jake area and was densely colonised with macrophytes. The majo:- share of biomass belonged 
to charophyte species (95 .5%) while planktonic and filamentous algae contributed 3.5%, only. In spite 
of this minority they presented an influential competitor to stonewort for light. Reduction of light 
penetration was a feature which have been measured in the last years. Successful competition of 
filamentous algae for nutrients was evident in the shallowest parts, with exception ofthe Jocations with 
steep slopes and unsuitable substrates for stoneworts and higher plants to anchor. The colonisation of 
littoral with anchored macrophytes increases the ecosystem resilience, by stabilising the erosion zone 
which overtakes function of a sedimentation zone. 
In spite of different influences from the watershed, the direct impact of a single disturbance was 
difficult to define on a tem pora! scale, but the consequences of integral impact resulted in reduction of 
water transparency, enhanced appearance of filamentous algae and shrinking of littoral zone. Those 
features indicated the eutrophication in lake Krnsko jezero and showed trend of accelerated succession 
in a remote mountain lake ecosystem. 
Povzetek 
Pospešena evtrofikacija Krnskega jezera se odraža v visoki primarni produkciji , kar pomeni zelo 
visoko vezavo sončne energije v jezeru . Litoralje v letu 1998 zavzemal kar40 % jezerske površine, ki 
je bila gosto poseljena z makrofiti. Večinski delež biomase je pripadel parožnicam (95,5% ), medtem ko 
je bil delež planktonskih in nitastih alg samo 3,5%. Kljub podrejenemu deležu so bile alge vpliven 
kompetitor za svetlobo, katere intenziteta se je v jezeru zmanjševala. Uspešnost nitastih alg v tekmovanju 
za hranila je bila očitna predvsem v plitvejših predelih, z izjemo odsekov na strmih brežinah, kjer je bil 
substrat neprimeren za zakoreninjenje parožnic in višjih rastlin. Poselitev litorala z makrofitskimi 
vrstami, ki se zakoreninijo, povečuje elastičnost ekosistema, saj te vrste s koreninskim sistemom 
stabilizirajo potencialne erozijske cone, ki tako postanejo sedimentacijske cone. 
Kljub očitnim vplivom iz zaledja je neposreden vpliv posamezne motnje tako časovno kot prostorsko 
težko razločiti. Vendar pa so posledice vseh motenj določljive in prepoznane kot znižanje prosojnosti 
vode, kot povečano pojavljanje nitastih alg in kot krčenje litoralne cone. Vsi ti pojavi, ki so prepoznani 
kot odraz pospešene evtrofikacije, so v Krnskem jezeru časovno zgoščeni in kot taki dober prikaz 
sukcesijskih procesov v oddaljenem gorskem jezerskem ekosistemu. 
Acknowledgement 
Project was supported by Ministry of Science and Technology Republic of Slovenia (SLO-ALPE 
project Paleoecology and ecology in mountain lakes, JI 7 414) and partly by European Commission as 
M O LAR project (Measuring and modelling the dynamic response of remote mountain lake ecosystems 
to environmental change, IC20-CT96-02 J ). 
References 
APHA 1992: Standard Methods for the examination of Water a Wastewater, J S•h edition. Washington 
DC, USA. 
BLAŽENČIČ, J., O. URBANC- BERČIČ & D. VRHOVŠEK 1990: Makrofiti v jezerih Triglavskega narodnega 
parka. Biološki vestnik 38: 1-14. 
BuNnow, 1,1992a: Decline of charophytes during eutrophication; a comparison to angiosperms. 
Freshwater Biology, 28: 9 -14. 
34 Acta Biologica Slovenica, 45 (2), 2002 
BLINDOW, 1.1992b: Long and short term dynamics of submerged macrophytes in two shallow eutrophic 
lakes. Freshwater Biology, 28: 15-27. 
BRANCELJ, A., G. Kosi & M. ŠIŠKO 1997: Distribution of algae and crustacea in mountain lakes in 
Slovenia with different trophic levels. Periodicum Biologorum 99: 87-96. 
BRANCELJ, A., M. Š1šKo, l. REJEC BRANCELJ, Z. JERAN & R. JAč1Mov1c 2000: Effects ofland use and fish 
stocking on a mountain lake - evidence from the sediment. Periodicum Biologorum 102: 
259-268. 
FoRSBERG, C.1964: Phosphorus, a maximum factor in the growth ofCharaceae, Nature, 201: 517 - 518. 
GABERŠČIK, A. & O. URBANC-BERČIČ 1996: Lakes of the Triglav National park: water chemistry and 
macrophytes. In: GABERŠČIKA., O. URBANC-BERČIČ, G.A. JANAUER (eds.): Proceedings of the 
Intemational Workshop and 8th Macrophyte Group Meeting IAD-SIL, Bohinj 1996, pp. 23-29. 
GABERŠČIK, A., O. URBANC-BERČIČ, A. BRANCELJ & M. Š1šKo 1997: Mountain lakes - remote but 
endangered. In: ROŠ, M. ( ed. ): Proceedings l st Internati ona! Conference on environmental 
restoration, IAWQ, SDZV, Ljubljana 1997, pp. 452-457. 
GACIA, E., E. BALLESTEROS, L. CAMARERO, O. DELGADO, A. PALAU, J.L. RIERA & J. CATALAN 1994: 
Macrophytes from lakes in the eastem Pyrenees: community composition and ordination in 
relation to environmental factors. Freshwater Biology, 32: 73 -81. 
GAMS, l. 1962: Visokogorska jezera. Poročilo, pp. 197-261. 
GoLTERMAN, H. L., R.S. CLLYMO, & M. A. M OHNSTAD 1978: Methods for Physical and Chemical 
Analysis of Freshwaters. Billing and Sons Limited, pp. 162. 
HouoH, R. A., M. D FoRNWALL, B. J. NEGELE, R. L. THOMPSON & D. A. Purr 1989: Plant community 
dynamics in a chain of lakes: principal factors in the decline of rooted macrophytes with 
eutrophication. Hydrobiologia, 173: 199 - 217. 
LACHAVANNE, J. - B.1985: The influence of accelerated eutrophication on the macrophytes of Swiss 
lakes: Abundance and distribution. Verh. Int. Ver. Limnol. 22: 2950 -2955. 
LEHMANN, A. & J. - B. LACHAVANNE 1999: Changes in the water quality ofLake Geneva indicated by 
submersed macrophytes. Freshwater Biology, 42: 457-466. 
MELZER A.1992: Submersed macrophytes. In: Scharf, B.W. & S. Bjorks (eds.): Limnology of Eifel 
maar lakes, Ergebnisse der Limnologie 38: pp. 223-237. 
MURI, G., Z. JERAN, R. JAČIMOVIc & O. URBANC-BERČIČ 2002: Fizikalne lastnosti sedimentov in 
onesnaževalci v sedimentih. V: (ur. A. Brancelj)Visokogorskajezera vzhodnem delu Julijskih 
Alp. Založba ZRC, ZRC - SAZU.159 - 177. 
OZIMEK, T., R.D. GULATI & E.vAN DoNK1990: Can macrophytes be useful in biomanipulation oflakes? 
The Lake Zwemlust example. Hydrobiologia 200/201: 39-407. 
PALL, K. & G.A. JANAUER 1995: Die Makrophytenvegetation von Flul3stauen am Beispiel der Donau 
zwischen Flul3-km 2552,0 und 2511,8 in der Bundesrepublik Deutchland. Arch. Hydrobiol. 
Suppl. 101, Large Rivers 9, 2: 91-109. 
RAsPoPov I. M., I. N. ANDRONIKOVA, O.N. DorsENKO, G.I. KuRASHov, G. I LETANSKAYA,. V.E. PANov, 
M.A. RYcHKOVA, LV. TELESH, O.A. TcHERNYKH & F.F. VoRoNrsov 1996: Littoral zone of 
Lake Lado ga: ecological state evaluation, Hydrobiologia 322: 39 - 47. 
R0RSLETT, B. 1991: Principal determinants of aquatic macrophytes richness in Northem European 
lakes. Aquatic Botany 39: 173 - 193. 
TALLING, J. F. 1975: Primary production of aquatic plants - conclusions. In: Cooper J. P. (ed.): 
Photosynthesis and productivity in different environments. Cambridge Univ. Press, 
Cambridge, pp 281-294. 
WETZEL, G. R. & G. E. LIKENS 1995: Limnological analyses. Second Edition. Springer -Verlag, pp. 
156-161. 
WETZEL, G. R. 1990: Land- water interfaces: Metabolic and limnological regulators. In Edgardo Baldi 
Memorial Lecture. Verh. Int. Verein. Limnol. 24: 6 -24. 
ACTA BIOLOGICA SLOVENICA 
LJUBLJANA 2002 Vol. 45, Šf. 2 : 35 - 52 
Sprejeto (accepted): 2002-05-27 
Goli polži (Gastropoda: Pulmonata: Milacidae, Limacidae, 
Boettgerillidae, Agriolimacidae, Arionidae) Slovenije 
Slugs (Gastropoda: Pulmonata: Milacidae, Limacidae, Boettgerillidae, 
Agriolimacidae, Arionidae) of Slovenia 
Marjan VAUPOTIČ*, France VELKOVRH** 
*Generala Maistra 5, SI-9000 Murska Sobota, Slovenia; e-mail: marjan.vaupotic@uni-lj.si 
**Lackova ulica 52, SI-2230 Lenart v Slovenskih goricah, Slovenia 
Uvod 
Izvleček. Podan je prvi obsežnejši pregled favne golih polžev Slovenije. Z obdelavo 
približno 600 vzorcev z golimi polži (Milacidae, Limacidae, Boettgerillidae, 
Agriolimacidae in Arionidae) iz treh zbirk in z analizo podatkov iz literature je bilo na 
območju Slovenije ugotovljenih 27 vrst. Med njimi je šest vrst, za katere so to prvi 
podatki o prisotnosti na območju Slovenije. Največ vrst golih polžev v Sloveniji je iz 
družine Milacidae (8). Seznam vrst je dopolnjen s kartami njihove razširjenosti po 
kvadratih UTM (10 x 10 km). 
Ključne besede: polži (Gastropoda), goli polži, favna, razširjenost, Slovenija. 
Abstract. Slugs are among the less investigated gastropods in Slovenia. The 
study of approximatelly 600 samples of slugs (Milacidae, Limacidae, Boettgerillidae, 
Agriolimacidae and Arionidae) from three collections and the analysis of data from the 
literature gave a total of 27 species for the territory of Slovenia. For six species this is 
the first record for Siovenia. Most slugs in Slovenia belong to the family Milacidae 
(8). A list of species with distribution maps in UTM grids (10 x 10 km) is given. 
Key words: snails (Gastropoda), slugs, fauna, distribution, Slovenia. 
Na območju Slovenije živijo predstavniki petih družin golih polžev: Milacidae - grebenarji, Limacidae -
slinarji, Boettgerillidae - črvarji, Agriolimacidae - poljski slinarji, Arionidae - lazarji. Po ocenah 
WIKTORJA ( 1997) obsegajo te družine vsega približno 300 vrst. V Sloveniji so goli polži med slabše 
raziskanimi skupinami polžev. 
V starejši literaturi že zasledimo nekaj podatkov o golih polžih iz Slovenije ali njene bližnje 
soseščine. SCHMIDT (1847) v prvem favnističnem popisu mehkužcev na slovenskih tleh tedanje 
36 Acta Biologica Slovenica, 45 (2), 2002 
avstro-ogrske dežele Kranjske še ne navaja golih polžev. Šele ERJAVEC (1877) omenja deset vrst iz 
Posočja. SIMROTHje opisal nove vrste golih polžev tudi iz Slovenije, leta 1885 vrstoAmalia robici 
(danes jo imenujemo Tandonia robici) iz Suhega dola nad Zgornjo Kokro v Savinjskih Alpah in nato 
leta 19IO vrsto Amalia kobelti z Mangarta (danes je to Tandonia simrothi). BOLE (1969) navaja 
štirinajst vrst v določevalnem ključu za mehkužce Slovenije. Tudi v nekaterih drugih delih navaja 
podatke o golih polžih Slovenije (BOLE 1962, 1966, 1976a, 1976b, 1977, 1979a, 1979b, 1981, 
1992a, 1992b, 1994). V delu o golih polžih nekdanje Jugoslavije so bili prvič (WIKTOR 1996) zbrani 
natančnejši podatki o vrstah iz Slovenije. VAUPOTIČ & VELKOVRH (1997) sta z intenzivnim 
vzorčenjem favne mehkužcev v Pomurju ugotovila 11 vrst golih polžev, med njimi je bila prvič odkrita 
Boettgerilla pallens. Gole polže z ozemlja Slovenije navajajo v svojih delih še STOSSICH (1899), 
SAJOVIC (1908), WAGNER (1931), KOS (1933), JAECKEL et al. (1958), ALTENA (1973, 1977), 
REISCHUTZ (1978), WIKTOR (1982), WOLF & RAHLE (1987), SLAPNIK (1998). 
Predloženo delo je prikaz trenutnega poznavanja favne golih polžev Slovenije (Sl. 1) in temelji na 
analizi podatkov iz literature ter taksonomski obdelavi skoraj 700 vzorcev, ki jih je prvi avtor članka 
pregledal v okviru svojega magistrskega dela. Vsi podatki so iz 112 kvadratov UTM (10 x JO km). 
~ v 
/ 
'j' i'-.. 1 1 \ r'f 1 Hu~gar' __,, 
" 
8ustrla 
l 21 ' - 1 I'-... '- ,.i-"-- .......... ......... -., ~ ~3 2 "- 12 ~ - ,........,~ --, ./ 1 3 5 1 ~ :.-r ["\1 11 3 6' i.-~ 
u 1z .... 9 -z;-~ '" i.L - 1 5 4, 5 2 .., j'\_ ' --- '\. ~II" .J2 5 5 j O 2. 4\J / 7 rJ-... 1 JI... 3 ~ rr ._ "'\ !--' -
~ <;, \.3 6 ~ 
_, 
1 l'\1. 1 9 5 2 - '\ '2 1""" ./ lr ,,_,, 
M ... ~ 3 '\1 { 1 -... ) 4 1 3 1 3 ::. ' "' ~/r 1 8' , 6 3 L. "6' ~ ' .,-,, l_ " 2 2 3 1 1 3 " -1.. [ J ,,,.JJ1 6 5 7 1 1 2 1> 2 ~ l { 
...... 2 3 4 3 7 7 1 4 iJ'-- j,,, J~ "'K 
~~' 4 2 1 4 1 2 
~ o 
T 
,, 4 6 .f 1 5 1 J ....__ 
l!. 1 6 r 1l - 1 1 \ .. 5 ) / 
'2 ~ '-' -" ..... , .... i""\ ,,~ T ~ 
~ _/ ' Cr.oati · / ,, 
Figure 1: Number of slug species known in individual UTM squares in Slovenia. 
Slika 1: Število ugotovljenih vrst golih polžev v posameznih kvadratih UTM v Sloveniji. 
Material in metode dela 
Pregledala sva vzorce iz zbirke mehkužcev J. Boleta na Biološkem inštitutu ZRC SAZU (BI), in 
lastnih zbirk (FV in MV). Vseh vzorcev v zbirkah je bilo 686, od tega največ v zbirki FV (562 
vzorcev), manj v zbirki MV (117 vzorcev) in najmanj v zbirki Biološkega inštituta (7 vzorcev). V 38 
vzorcih so bili le rudimenti hišic golih polžev, ki zaenkrat niso uporabni za določitev vrste (SOUTH 
1992 cit. REUSE 1983). Pregledani material je bil nabran v obdobju od leta 1957 do 1998. Živali so 
shranjene v 70 % etanolu. Vseh živali v vzorcih je bilo približno 4000. Nabirali smo na dva različna 
M. Vaupotič, F. Velkovrh: Slugs (Gastropoda: Pulmonata: Milacidae, Limacidae, Boettgerillidae ... 37 
načina: naključno in intenzivno. Pogostejši je bil naključni način vzorčenja, ko število primerkov v 
vzorcih ni presegalo števila 10. V nekaterih vzorcih, ki so rezultat intenzivnega vzorčenja, je bilo tudi 
več sto živali, zaradi zelo uspešne metode nabiranja, ki jo je drugi avtor članka izvajal le na eni lokaliteti 
(Lenart v Slovenskih Goricah, WM65). Zvečer je na namočena tla nastavljal vlažne, 4-5 cm debele 
deske. Čez noč so se pod te deske zatekli mnogi polži, ki jih je naslednji dan pobral, še preden je 
dopoldansko sonce segrelo deske. 
Taksonomsko pomembni znaki za določitev vrste so na spolnem aparatu, zato je bilo večinoma 
treba opraviti sekcijo živali pod stereo-mikroskopom. Za določanje vrst so bila uporabljena dela več 
avtorjev (KERNEY et al. 1983; LIKHAREV & WIKTOR 1980; QUICK 1960; WIKTOR 1973, 
1982. 1983. 1987, 1989, 1996; BACKELJAU & VAN BEECK 1986; BACKELJAU & DE BRUYN 
1990; GARRIDO et al. 1995; LUPU 1977; VON PROSCHWITZ 1989; DE WINTER 1984,1989; 
WIKTOR & MILANI 1995). 
Ves pregledani material je shranjen v zbirki Franceta Velkovrha, Lenart v Slovenskih goricah, 
Lackova ulica 52. 
Rezultati 
Ugotovljene vrste so naštete v seznamu. Pri vsaki vrsti je navedena njena razširjenost v Sloveniji. 
Prvi podatki so iz literature, večinoma po WIKTORJU (1996), ki navaja tako rezultate lastnih raziskav 
favne tedanje Jugoslavije, kot tudi podatke drugih avtorjev (ALTENA l 973, 1977; BOLE 1962, 1966, 
1976a, 1976b, 1977, 1979a; GROSSU 1972; JAECKEL & MEISE 1956; SIMROTH 1885, 1910; 
WAGNER 1931; WOLF & RAHLE 1987). Nekatera dela pa v prej navedeni monografiji niso 
upoštevana (BOLE 1969, 1981, 1992b, 1994; REISCHUTZ 1978; SLAPNIK 1998; VAUPOTIČ & 
VELKOVRH 1997). Sledijo lastni podatki po zgoraj omenjenih zbirkah. Štirideset vzorcev iz zbirke 
MV je bilo že uporabljenih in objavljenih (VAUPOTIČ & VELKOVRH 1997). V karte razširjenosti so 
iz literature vneseni podatki, ki so geografsko dovolj natančni, vrste pa identificirane po anatomskih 
znakih. Upoštevane so bile tudi navedbe o vrstah, ki se jih da zanesljivo določiti že po zunanjih znakih 
(Limax cinereoniger). V literaturi navedene vrste, ki jih nisva odkrila v vzorcih iz zbirk, so v pregledu 
vrst označene s črtico(-). 
Najdišča vrst so prikazana v mreži UTM. Pri navajanju podatkov iz literature je za imenom avtorja 
navedena samo koda UTM, medtem ko je pri podatkih iz sedaj obdelanih zbirk za kodo kvadrata UTM 
zapisana lokali teta, datum nabiranja in kratica zbirke s kataloško številko. Število osebkov v vzorcu je 
navedeno le v primeru, ko gre za eno samo lokaliteto oziroma lokalitete iz enega samega kvadrata 
UTM. Če je bilo več vzorcev z istega nahajališča, je pri ponovitvah naveden samo datum vzorčenja na 
najdišču in kataloška številka vzorca; če pa je bil drugi vzorec nabran tudi isti dan, je navedena le 
kataloška številka. 
Legenda: L - podatki iz literatur 
ZT - podatki, dobljeni z obdelavo vzorcev treh zbirk 
FV - zbirka Franceta Velkovrha 
BI - zbirka Biološkega inštituta SAZU 
MV - zbirka Marjana Vaupotiča 
? - podatki, ki niso vneseni v zemljevide, ker geografsko niso dovolj natančni ali pa 
je taksonomska določitev nezanesljiva 
38 Acta Biolo11ica Slovenica, 45 (2), 2002 
Table 1: Sistematic check-list of the slugs of Slovenia. 
Tabela I: Sistematski pregled vrst golih polžev Slovenije. 
DRUZINA VRSTA 
Milacidae l. Milax nigricans 
2. Tandonia budapestensis 
3. Tandonia rara 
4. Tandonia reu/eauxi 
5. Tandonia robici 
6. Tandonia rustica 
7. Tandonia simrothi 
8. Tandonia sowerbvi 
Limacidae 9. Malacolimax mrazeki 
JO. Lehmannia marginata 
11. Limax cinereoniger 
12. limax maximus 
13. Limax fJ.avus 
Boettgerillidae 14. BoettHerilla eallens 
Agriolimacidae 15. Deroceras laeve 
16. Deroceras lothari 
17. Deroceras reticulatum 
18. Deroceras rodnae 
19. Deroceras sturanyi 
20. Deroceras turcicum 
Arionidae 21. Arion lusitanicus 
22. A rion subfuscus 
23. A rio,1 circumscriprus 
24. Arion fasciatus 
25. Ario11 silvaticus 
26. Arion a/pinus 
27. A rio11 distinctus 
DRUŽINA: Milacidae - grebenarji (ali vrtni lazarji) 
- Milax gagates (Draparnaud, 1801) - temni grebenar 
L:? BOLE (1969),? BOLE (1981). 
l. Milax nigricans (Schulz, 1836) - črni grebenar (Sl. 2 A) 
Z: VL04: Pridvor = Sv. Anton, Dekani, Koper, 8.1995, FV48653. V vzorcu je 5 osebkov. 
2. Tandonia budapestensis (Hazay, 1881) - madžarska grebenarka (Sl. 2 A) 
Z: VL08: Dornberk, 11.10.1998, MV697; VL18: Vipavski Križ, l l.J0.1998, MV705; WM65: 
Lenart v Slovenskih Goricah, 8. 1986, FV45623; 1995, FV49989; 7.-10.1996, FV50877; 7.-
10.1996, FV51065; 7.-10.1997, FV51442; 5. - 7.1994, MV607; WM96: Generala Maistra 5, 
Murska Sobota, 29.9.1996, MV616; WM96: žive meje, vrtovi, Murska Sobota, 8.-9.1993, 
MV674. 
-Tandoniafejervaryi (Wagner, 1929) - Fejervaryjeva grebenarka 
Milax ( Milax) Adensameri Wagner 1931. 
L: ALTENA ( 1977)/ VM63. 
3. Tandonia rara Wiktor, 1996 - redka grebenarka (Sl. 2 A) 
Tandonia schleschi partim in Wiktor 1982 
Z: VM62: Planina Koren, Krvavec, 6.1974, FV26345. V vzorcu so 3 osebki. 
4. Tandonia reuleauxi (Clessin, 1887) - Reuleauxjeva grebenarka (Sl. 2 A) 
Aspidoporus limax Fitzinger 1883 
L: WAGNER (1931) - ? Krain (as Aspidoporus limax) (Kranjska (kot Aspidoporus limax)). 
Z:VL06: Jama v Doktorjevi ogradi, Pliskovica, Tomaj , 1.6.1996, MV654. V vzorcu je en sam 
osebek. 
5. Tandonia robici (Simroth, 1885) - Robičeva grebenarka (Sl. 2 B) 
Amalia Robici Simroth 1885, Milax robici (Simroth 1885) 
M. Vaupotič, F. Velkovrh: Slugs (Gastropoda: Pulmonata: Milacidae, Limacidae, Boettgerillidae... 39 
L: SIMROTH (1885)/ VM62; WAGNER (1931)/ UM92;? JAECKEL ET AL. (1958);? 
BOLE ( 1969); WIKTOR ( 1982)/ VM23; WOLF & RAHLE ( 1987)/ VM04; WIKTOR ( 1996)/ 
VL27; VLS4. 
Z: VL45: melišče, pod steno Pešišče, Koritnice, 10.11.1994, MV656; VL47: Rakov Škocjan, 
10.1969, FV10123; pred vhodom: Planinska jama, Planina, 6.1976, FV32608; pred jamo: 
Mačkovica, Planina, 4.1977, FV36015; Pod stenami, Planinsko polje, 5.1982, FV44616; 
Laška kukava, Logaška planota, Laze, Planina, 6.1995, FV48795; vhodna udornica, Vranja 
jama, Planinsko polje, N rob, 6.1995, FV48998; Pod stenami, Planinsko polje, Planina, 
6.1982, FV51328; 7.1982, FV50876; VL48: N Ljubljanski vrh, Vrhnika, 6.1977, FV36046; 
Planinski polje, N rob, Planina, 6.1985, FV50865; VL57: Slivnica, Cerknica, 6.1971, FV50870; 
6.1971, FV51307; FV51323; VL95: Stojna, Dolga vas, Kočevje, 9.1996, FV51451; VM04: 
Prisojnik, W pobočje, Vršič, 6.1981, FV46026; VM43: vas Dolina, Tržič, 5. 1974, FV25433 ; 
VMS0: Tume, Grmada, Tacen, Ljubljana, 6.1997, FV50615; Podutik, nad kamnolomom, 
Ljubljana, 6.-7.1997, FV50982; VM61: stena nad vasjo Rašica, Ljubljana, 7.1994, FV47407, 
FV 47408; Rašica SW pobočje, Ljubljana, 6.1995, FV 48883; VM62: Planina Koren, Krvavec, 
6.1974, FV26348; Dolge njive, Krvavec, 6.1978, FV38691 ; Dolina Korošice, Krvavec, 6.1978, 
FV38775; desno pobočje: Dolina Korošice, Stahovica, 6.1998, FV51058; Korošica- Krvavec, 
4.1970, FV51319; Davovec, Cerklje, 6.1998, FV51622; VM95: pred Jamo v Votli peči, 
Ravne DRJS-3263, 5.1975, FV50875; WL0S: Podstene, Kočevski Rog, 6.1980, FV45783; 
6.1986, FV 45662; WLlS: Semič , Bela Krajina, 14.4.1990, FV 47476; WLl 7: gozd, Otočec, 
26.9.1998, MV701. 
6. Tandonia rustica (Millet, 1843) - podeželska grebenarka (Sl. 2 B) 
Milax rusticus (Millet 1843) 
L: ? BOLE ( 1969); ? BOLE ( 1979a); WOLF & RAHLE ( 1987)/ UM83. 
Z: VL08: Dornberk, 11.10.1998, MV696; VL16: W Dane, Sežana, 9.1997, FV50636; VL18: 
Vipavski Križ, 11.10.1998, MV698. 
7. Tandonia simrothi (Hesse, 1923) - Simrothova grebenarka (Sl. 2 A) 
Amalia Kobe/ti Simroth 1910, Milax Simrothi Hesse 1923, Milax (Milax) Sch/eschi Wagner 1930, 
Tandonia schleschi (Wagner 1930) 
L: SIMROTH(l910)/ UM94; BOLE (1969)/ UM94; WOLF & RAHLE (1987)/ VM04, 
UM92; WIKTOR (1996)/ VM63. 
Z: UM92: Planina Duplje, Julijske Alpe, 7. 1981, FV41412; pri Dupeljskem jezeru, Planina 
Duplje, Julijske Alpe, 7.1982, FV43645; UM94: Mangart, pod Rdečo skalo, 9.1970, FV 12602; 
VM02: Komna, 8.1958, FVl 1387; 8.1958, FVI 1388; Planina Govnač, Komna, 5.1972, 
FV20525; Planina Govnač, Komna, 7.1982, FV43614; Komna - Bogatinsko sedlo, 7.1985, 
FV45008; 7.1985, FV45891/51311; VM04: Erika (koča), Kranjska Gora - Vršič, 6.1975, 
FV32163; Vršič - Trenta, 6.1975, FV32178; 6.1975, FV32179; Vršič, 7.1975, FV35407; 
Prisojnik, S pobočje: pod oknom, Kranjska Gora, 6.1977 , FV36616; Mihov dom, Vršič -
Kranjska Gora, 7.1981, FV42013; Prisojnik, W pobočje, Vršič, 7.1980, FV45416; 7.1984, 
FV45773; 6.1984, FV45826; 7.1984, FV45867; Kranjska Gora - Vršič, 6.1981, FV45944; 
VM73, Kamniško sedlo, Kamniška Bistrica (Sl640a), 21.8.1994, B1003; Kamniško sedlo, 
8.9.1987, BI004. 
8. Tandonia sowerbyi (Ferussac, 1823) - Sowerbyjeva grebenarka (Sl. 2 C) 
Z: UL94: Portorož - Beli Križ - Fiesa, 24.10.1998, MV703; Piran, 24.10.1998, MV704; 
VL04: Pridvor = Sv. Anton , Dekani, Koper, 8.1995, FV50860; VL18: Vipavski Križ, 
11.10.1998, MV699; VM60: Ljubljana, 10.1976, FV33182; Ljubljana, Kolodvor, 10.1976, 
FV34231. 
DRUŽINA: Limacidae - slinarji 
9. Malacolimax mrazeki (Simroth, 1904) - Mrazekov mali slinar (Sl. 2 D) 
40 Acta Biologica Slovenica, 45 (2), 2002 
Malacolimax kostali Wiktor 1982, nec Babor 1900 
L: WIKTOR (1982)/ VLSS; WIKTOR (1996)/ VM14, VL54. 
Z: VL47: gozd, pasti, Laze, Planina, 3.1995, FV 49911; VM04: pod slapom, Gozd Martuljek, 
6.1974, FV26395; Erjavčeva koča, Vršič, 7.1981, FV41422; Prisojnik, W pobočje, Vršič, 
6.1981, FV51441; VM72: Velika planina, Savinjske Alpe, 6.1976, FV38659; WL13: hrib 
Sebetih, Sinji vrh, Črnomelj, 8.1974, FV28582; WL17: Otočec, gozd, 26.9.1998, MV694; 
WM0l: Marija Reka, Prebold , 9.1996, MV659; WM0S: Jesenkov vrh, Slovenj Gradec, 
9.1996, FV51303; 9.1996, MV661; WM33: Oplotnica, Pohorje, 8.1967, FVl 1474; WM34: 
Šumnik, Pohorje, 8.1967, FV11486; Ruško Pohorje, 8.1995, FV48915; WM35: kmetija 
Strnad, Smolnik, Ruše, 9.1996, MV668; WM65: Lenart v Slovenskih Goricah, 8.1986, 
FV45726; WM66: Zgornja Ščavnica, Slovenske Gorice, 10.1987, FV46544; WM97: 
Sebeborci, gozd, Goričko, Prekmurje, 9.9.1995, MV617. 
- Malacolimax tenellus (0. F. Milller, 1774) - gobji mali slinar 
Limax tenellus Nillson 1822 
L: ? BOLE ( 1969) - samo v Karavankah, ? BOLE ( 1979a) - v gozdovih na Javornikih, ? 
WOLF & RAHLE (1987)- N ofVrišič pass (severno od prelaza Vršič/VM04). 
10. Lehmannia marginata (O. F. Milller, 1774)- obrobljena slinarka (Sl. 2 E) 
Limax marginatus O.F. Miiller 1774 
L: BOLE (1962)/ VM03; BOLE (1969)/ VL19, VL54; BOLE (1977)/ VMS0; WOLF & 
RAHLE(1987)/VM04, UM83;WIKTOR(1996)/VM13, VM33, VL27, WMOO;VAUPOTIČ 
& VELKOVRH (1997)/ WM95, WM96; SLAPNIK (1998)/ WM00. 
Z: VL09: Trnovski gozd, 1969, FV09352; VL47: Pod stenami , Planina, 6.1982, FV43483; 
7.1982, FV43659; Vranja jama, vhodna udornica, Planinsko polje, N rob, 6.1995, FV5l315; 
Rakov Škocjan, 10.1969, FV51429; VL48: S Ljubljanski vrh, Vrhnika, 6.1977, FV36047 ; 
Planinsko polje, N rob, Planina, 6.1985, FV46530; VL48: Bistra, izvir v Bistri, Vrhnika, 
25 .6.1990, FV47623; VLSS: grad Snežnik, Cerknica, 9.1977, FV36518; FV36520; VL56: 
pri Mrzli jami, Bločice, 29.7.1994, MV655; VL57: Slivnica, 6.1971, FV15846; FV19776; 
VL78: Vodice na Dolenjskem, Dobrepolje, 6.1958, FVl 1374; FV 11375; VMOl: dolina pri: 
Doblarec, Doblar, 6.1998, FV51632; VM02: Savica - Komna, Bohinj, 7.1985, FV45249; 
7.1986, FV45794; FV45997; 7.1982, FV46439; 7.1981, FV47719; 6.1981, FV51542; VM04: 
Vršič, 7.1975, FV35401; Prisojnik, W pobočje, Vršič, 7.1980, FV45412; 6.1981, FV51320; 
6.1984, FV45817; 7.1984, FV45866; Mihov dom, Kranjska Gora - Vršič, 6.1980, FV45805; 
6.1981 , FV50866; 7.1981, FV50873; FV51324; Kranjska gora - Vršič , 6.1981, FV45935; 
FV45937; Vršič - Trenta, 6.1975 , FV51312; Vršič, 7.1975, FV5l325; VM24: Mežakla, 
6. 1969, FV11144; FV50867 ; FV51435 ; 6.1971 , FV18795 ; Srednja Radovna, 6. 1970, 
FVl4657; VM33: gozd, Mošnje, Dobruša, levi breg, Brezje, 6.1995, FV48710; FV487l1; 
FV48712; VM41: Moškrin, Škofja Loka, 6.1997, FV50750; Grenc, Škofja Loka, 6.1997, 
FV50850; VMS0: Turne, Grmada, Tacen, Ljubljana, 6.1997 , FV50612; VM62: Dolina 
Korošice, Kamniška Bistrica, 6.1974, FV26313; Dolge njive, Krvavec, 6.1978, FV38692; 
Planina Koren, Krvavec, 6.1978, FV38751; VM63: Rokovnjaške luknje, Kamniška Bistrica, 
7.1979, FV41243; VM72: Kamniška Bistrica, 7.1979, FV4!699; VM82: Javoršek, Tuhinj, 
5.1970, FV11780; FV11785; VM82: Mačkin kot, Gornji grad, 5.1976, FV25529; VM95: 
pred Jamo pri Votli peči , Ravne DRJS-3263, 5.1975, FV32534; VM95: Jama v Votli peči, 
Ravne, Koroška, 3.1978, FV 42066; 1.1978, FV 45394; S pobočje, Straži ščec (karbonati), NE 
Prevalje, 9.1994, FV47342; WL0S: Podstene, Kočevski Rog, 7 .1980, FV42778; FV51431 ; 
6. 1980, FV45784; FV5l423; WL26: Gospodična, Gorjanci, 5. 1974, FV26109; WL48: 
Vitovec, E Malence, Brežice, 7.1994, FV47326; WM0S: Jesenkov vrh, Slovenj Gradec, 
9 .1996, MV 660; WM16: Kremenice, Šmartno Pohorje, 7 .1986, FV 46080; WM24: Pohorje, 
8.1967, FV50880; WM33: Oplotnica, Pohorje, 8.1967, FV50883; WM34: Pragozd, Lobnica, 
M. Vaupotič, F. Velkovrh: Slugs (Gastropoda: Pulmonata: Milacidae, Limacidae, Boettgerillidae.. . 41 
Pohorje, 6.197~, FV30199; FV30200; Šumnik, Pohorje, 8.1967, FV 50886; WM42: pragozd, 
Boč, Poljčane, Stajerska, 8.1997, FV50555; WM43: Bistriški vindgar, N Slovenska Bistrica, 
6.1997, FV50564; WM44: Areh, Pohorje, 8.1967, FV46172; WM46: Brestemica, 9.1996, 
MV665; WM65: Lenart v Slovenskih Goricah, 8.1985, FV44326; 9.1986, FV45705; WMSS: 
gozd, Desnjak, SW Žerovinci, Ljutomer, Prlekija, 7.8.1994, MV646; WM95: vrt, Veržej, 
Prlekija, 6.11 .1993, MV 641; WM96: vrtovi , žive meje, Murska Sobota, Prekmurje, 8.+9 .1993, 
MV626; vrtovi, žive meje, Murska Sobota, Prekmurje, 9.9.1993, MV627; Murski logi, SW 
Dokležovje, Prekmurje, 11.8.1993, MV630; dvorišča, vrtovi, Dokležovje, Prekmurje, 9.9.1993, 
MV638. 
11. Limax cinereoniger Wolf, 1803 - pepelnočrni slinar (Sl. 2 F) 
Limax cinereo-niger Wolf 1803, Limax maximus Linnaeus 1758 (partim), Limax lineo/atus Dumont 
et Mortillet 1852. 
L: BOLE ( 1962)/ VL47; BOLE (1969); BOLE (l 976a)/ VL19, VLOS, VL47 (3), VL49, 
VL47, VL54, VL95; BOLE (1976b)/ VL54; BOLE (1977)/ VMSO, BOLE (1978)/ VL46; 
WIKTOR (1982)/ VL55; WOLF & RA.HLE (1987)/ VM04, UM83 (3); WIKTOR (1996)/ 
WMlO, VL37, VM14, VM33, VM13, VL27; VAUPOTIČ & VELKOVRH (1997)/ WM96. 
Z: UM94: Mangart, pod: Rdeča skala, 9.1970, FVl2603; VL16: Laze, Novo mesto, 6.1997, 
FV50350; FV50351; VL17: Mlake pri Vipavi , l l. l0.1998, MV682; VL18: Vipavski Križ, 
11. l0.1998, MV683; VL29: Godovič. Idrija, 6.1997, FV5055 I; VL45: Koritnice, Boršt -
hrastov gozd, Knežak, JO. ! 0. 1988, MV708; MV709; Jurišče, Gradec, Pivka, l0.10.1988, 
MV7IO; Koritnice, pod lesom, Knežak, 15.I0.1998, MV717; VL47: Rakov Škocjan, 10.1969, 
FVI0!22; FV51317; 1968, FVI 1550; pred vhodom: Planinska jama, Planina, 6.1976, 
FV32605; Pod stenami, Planinsko polje, Planina, 6.1982, FV 43481; 6. I 992, FV 47890; Vranja 
jama, vhodna udornica, Planinsko polje, N rob, 6.1995, FV48987; VL48: Bistra, Vrhnika, 
4.1971, FVI4913; 5.1977, FV36762; 6.1980, FV42056; S Ljubljanski vrh, Vrhnika, 6.1977, 
FV51422; FV36022; FV36032; Laška kukava, Logaška planota, Laze, Planina, 6.1995, 
FV48790; VL54: Sviščaki , Snežnik, 6.1978, FV38469; FV38471 ; VL55: grad Snežnik, 
Cerknica, 9.1977, FV365 l 7; Leskova dolina, Snežnik, 6.1978, FV387 l 7; Mašun, Vidovi lazi, 
I0.10.1988, MV7I 3; VL56: Gorica, Cerkniško jezero, Cerknica, 6.1980, FV42053; VL57: 
Slivnica, 6.1971, FVI5843; Slivnica, gozd pod vrhom, 6.1971, FV19773; FVI9774; FV51322; 
FV50869; VLSS: Pekel, Borovnica, 6.1976, FV32653; VL67: Turjaški grad, Turjak, Velike 
Lašče, 6.1997, FV50786; VL68: Draga, Ig, 5.1967, FVl9778; VL78: Vodice, Dolenjsko, 
8.6.1958, FVI 1373; Limberk, Predole, 6.1998, FV51727; VLSS: Vir pri Stični, 6.1976, 
FV32652; VL95: Stojna, Dolga vas, Kočevje, 9.96,6.97, FV50357; FV50358; FV50359; 
VL97: Dvor pri Žužemberku, 6. 1970, FVI8796; FVl8797; VMOl: Tolmin, ob Soči , 4.1977, 
FV36804; VM02: Komna, 8.1958, FVI 1390; Savica- Komna, Julijske Alpe, 7.1982, FV46435 ; 
7. 1983, FV 43502; 7.1982, FV 43630; 7 .1985, FV 45250; 6.1986, FV 45584; Komna - Bogatinsko 
sedlo, Julijske Alpe, 7.1985, FV45891?; VM04: Gozd Martuljek pod slapom, 6.1974, 
FV26416; Mihov dom, Kranjska Gora- Vršič, 6.1980, FV50871 ; FV45798; 6.1981, FV45966; 
7 .1981, FV 42009; FV5 l 309; Kranjska Gora - Vršič, 6.1981, FV 45931; FV50872; Prisojnik, 
W pobočje, Vršič, 6.1981, FV51440; VM23: Kranjska dolina, Pokljuka, 9.1967, FV 11553; 
FV24762; VM24: Mežakla, Ravne, 6.1969, FV09901(4); FV50868; 6.1971, FVl8794; 
Srednja Radovna, 6.1970, FV14658; FV 17213; Črni vrh, Jesenice, 7.1957, FV24777; VM33: 
Mošnje, gozd, Dobruša: levi breg, Brezje, 6. I 995, FV48725; VM34: Završnica, Jesenice, 
6.1974, FV26136; VM41: Grenc, Škofja Loka, 6.1997, FV50851; VMSO: Tume, Grmada, 
Tacen, Ljubljana, 6.1997, FV50622; Podutik, nad kamnolomom, 6.-7.1997, FV50779; 
FV50972; VM61: Rašica SW pobočje, Ljubljana, 6.1995, FV48882; VM62: Korošica -
Krvavec, travniki , 9 .1970, FV I 2787; Dolina Korošice, S Kamniška Bistrica, 6.1974, FV263 l 8; 
Planina Koren, Krvavec, 6.1974, FV26339; 6.1978, FV38750; FV38753; Kopišče, Kamniška 
42 Acta Biologica Slovenica, 45 (2), 2002 
Bistrica, 6.1977, FV36733; 6.1976, FV36737; desno pobočje, Dolina Korošice pod Krvavcem, 
6.1988, FV51052; FV51053; Davovec, Cerklje, 6.1998, FV51623; VM71: Boštanovajama, 
Krtina, Moravče, 1.1980, FV 40366; VM72: Velika planina, Savinjske Alpe, 6.1976, FV38657; 
VM74: Olševa, gora, 7.1974, FV26591; VM80: Kresnice, Litija, 6.1974, FV26192; VM82: 
Javoršek, Tuhinj, 5.1970, FVl 1775; FVI 1776; FVl 1782(3?), FV51438; Mačkin kot, Gornji 
grad, 5.1976, FV51447; VM94: Skobirjeva votlica, Zg. Razbor, Slovenj Gradec, 10.1978, 
FV38 l 54; VM95: Ošven, Podgora, Ravne na Koroškem, 6.1988, FV51670; WL05: Podstene, 
Kočevski Rog, 7. l 980, FV 41124; WLl 7: gozdovi, Otočec, 1998, MV689; WL29: Studenec, 
Sevnica, 8.1972, FV20516; WL39: Bučerca, Krško, 6.1970, FV24902; Anže, Brestanica, 
5.1976, FV32568; WM00: Škofja Riža, 5.6.1991, B1001; Kum, zgornji del ob pretvorniku, 
6.5.1988, B1005; WM0l: Marija Reka, Prebold, 9.1996, MV658; WM05: Gradišče (silikati), 
N Slovenj Gradec, Pohorje, 9.1994, FV47243; FV47244; Jesenkov vrh, Slovenj Gradec, 
9.1996, MV657; WM13: Huda luknja, okolica, Velenje, 6.1994, FV50859; FV50895; WM14: 
Špehovka, jama, vhod, Mislinja, 9.1970, FV13652; WM16: Suhi Vrh nad: Radlje ob Dravi, 
7.1986, FV45629; WM24: Pohorje, 8.1967, FVI 1547; WM34: Pragozd, Lobnica, Pohorje, 
6.1975, FV30197; WM35: Ruško Pohorje, 8.1995, FV48910; kmetija Strnad, Smolnik, 
Ruše, 9 .1996, MV667; WM42: pragozd, Boč, Poljčane, Štajerska, 8.1997, FV50554; 8.1997, 
FV50556; okolica vasi, Hrastje, Poljčane, 7.1997, FV50572; FV50576; WM43: Slovenska 
Bistrica, 6.1988, FV5 l 798; FV51802; WM45: dolina: Razvanjski potok, Razvanje, Pohorje, 
Maribor, 6.1997 , FV50756; WM46: Bresternica, 9 .1996, MV663; WM65: Lenart v 
Slovenskih Goricah, 1995, FV 49990; 7 .-10.1997, FV5 l 445; WM66: Zgornja Ščavnica, Lenart 
v Slovenskih Goricah, 6.1985, FV44657; FV44658; Krivi Vrh, Sv. Ana, Lenart v Slovenskih 
Goricah, 8.1986, FV45622; WM96: Murski logi, W Stara Mura, S Dokležovje, Prekmurje, 
12.8.1993, MV622. 
12. Limax maximus Linnaeus, 1758 - veliki slinar (Sl. 2 G) 
Limax cinereus Lister 1678. 
L:? BOLE (1969); VAUPOTIČ & VELKOVRH (1997)/ WM95, WM96. 
Z: rastlinjak: B. Sket,Aškerčeva 12, Ljubljana, FV42237; UL93: Sečovlje, 9.1971, FV19646; 
VL04: Šalara, Koper, 6.1974, FV26337; Pridvor = Sv. Anton, Dekani , Koper, 8.1995, 
FV 48649; VL48: Bistra, Vrhnika, 6.1989, FV46953; VL55: Mašun, Vidovi, Lazi, 10.10.1988, 
MV707; VL59: Vrhovci, Ljubljana, 1958, FV 11348; WM14: Pečenikov vrh, Mislinja, 6. I 998, 
FV51764; WM65: Lenart v Slovenskih Goricah, 8.1987, FV46433; 5. - 7.1994, MV608; 
1995, FV49992; 7.-10.1996, FV51064; 7.-10.1997, FV50862; WM95: vrt, Veržej, Prlekija, 
6.11.1993, MV643; MV612; WM95: Noršinci, Ljutomer, Prlekija, 3.5.1995, MV670; Noršinci, 
Ljutomer, 3.5.1995, MV618; WM96: dvorišča, žive meje, Dokležovje, 9.9.1993, MV610; 
vrtovi, žive meje, Beltinci, Prekmurje, 9.9.1993, MV61 l ; vrtovi, žive meje, Murska Sobota, 
Prekmurje, 8. + 9.1993, MV621; Borovnjakova ulica, Murska Sobota, Prekmurje, 11 .6.1994, 
MV651. 
13. Limaxflavus Linnaeus, 1758 - rumenkasti slinar (Sl. 2 H) 
Limax variegatus Drapamaud 1801. 
L: ? BOLE (1969); WOLF & RAHLE ( 1987)/ UM83. 
Z: VL04: Pridvor = Sv. Anton, Dekani , Koper, 8. 1995, FV50861 ; VL45: Koritnice, pod 
lesom, 15.10.1998, MV711. 
DRUŽINA: Boettgrillidae - črvarji 
14. Boettgerilla pallens Simroth, 1910 - Boettgerjev črvar (Sl. 2 H) 
Boettgerilla vermiformis Wiktor 1959 
L: VAUPOTIČ & VELKOVRH (1997)/ WM96. 
Z: WM96: Generala Maistra 5, Murska Sobota, 29.9.1996, MV673; dvorišča, vrtovi , 
M. Vaupotič, F. Velkovrh: Slugs (Gastropoda: Pulmonata: Milacidae, Limacidae, Boettgerillidae... 43 
Dokležovje, Prekmurje, 9.9.1993, MV637; vrtovi, žive meje, Murska Sobota, Prekmurje, 
8.+9.1993, MV623. 
DRUŽINA: Agriolimacidae - poljski slinarji 
- Deroceras agreste (Linnaeus 1758) - enobarvni poljski slinar 
Limax agrestis Linnaeus 1774, Agriolimax agrestis var. Fedschenkoi Koch et Heynemann 1874. 
L:? BOLE (1969);? BOLE (1962);? BOLE (1976b);? BOLE (1977);? BOLE (1992). 
15. Deroceras laeve (0. F. Miiller, 1774) - obvodni poljski slinar (Sl. 2 H) 
Limax laevis O.F. Miiller 1774. 
L:? BOLE (1981);? BOLE (1994); WIKTOR (1996)/ VM33. 
Z: rastlinjak v Biološkem središču, Ljubljana, 7 .1994, MV620; VL14: Osp, Koper, 11.1984, 
FV4630I.V vzorcu je en osebek. 
16. Deroceras lothari Giusti, 1971 - Lotharjev poljski slinar (Sl. 3 I) 
Deroceras klemmi Grossu 1972. 
L: REISCHUTZ ( 1978)/ VL15, VM52, WM21; WOLF & RAHLE ( 1987)/VM04; WIKTOR 
(1996)/ VM60, VM52, VM33, VM14, VM13, WM00; ALTENA ( 1973)/ VM0l, VM62 (2); 
VAUPOTIČ & VELKOVRH (1997)/ WM95, WM96. 
Z: UL94: Portorož- Fiesa, 24.10.1998, MV687; Piran, 24.10.1998, MV688; VL04: Pridvor 
= Sv. Anton, Dekani, Koper, 8.1995, FV48655; VL0S: Dornberk, 11.10.1998, MV692; 
VL29: Idrija, 13.9.1998, MV706; VL45: Koritnice, na cesti, v dežju, 1.11.1998, MV714; 
Koritnice, v jami nad Koritnicami, 2.4.1995, MV715; pod lesom, Koritnice, 15 .10.1998, 
MV716; VL54: gozd Sviščaki, Ilirska Bistrica, 8.-10.9.1991, FV51437; VL69: Ljubljansko 
barje, ob Ižanski cesti, 10.1976, FV33410; VL77: Velike Lašče, 9.1984, FV43827; 11.1984, 
FV43950; FV43951; FV51302; VM02: Komna, 8.1958, FVI 1389; VM04: Mihov dom, 
Kranjska Gora - Vršič, 7 .1984, FV 42022; VM23: Kranjska dolina, Pokljuka, 9.1967, FV24763; 
VM50: Rožnik, Ljubljana, 6.1972, FV22805; Koseze, Ljubljana, 12.10.1994, MV614; WM24: 
Pohorje, 8.1967, FVI 1548; WM43: Kajuhova cesta, Slovenska Bistrica, 9.1996, MV662; 
WM65: Lenart v Slovenskih Goricah, 11.1984, FV 43939; 1995, FV 49985; FV 49986; 
FV50879; 7.-10.1996, FV51068; FV51436; 7.-10.1997, FV51443; WM95: vrt, Veržej, 
Prlekija, 6. 11.1993, MV644; Noršinci, Ljutomer, 10.9.1994, MV672; WM96: vrtovi, žive 
meje, Beltinci, Prekmurje, 9.9.1993, MV628; poplavni travnik, S Dokležovje, Prekmu1je, 
12.8.1993, MV635. 
17. Deroceras reticulatum (O. F. Miiller, 1774)- mrežasti poljski slinar (Sl. 3 J) 
L: WIKTOR (1996)/ VM33),? BOLE (1994); VAUPOTIČ & VELKOVRH (1997)/ WM95, 
WM96; SLAPNIK (1998)/ WM00. 
Z: VL07: Komen, 11.10.1998, MV702; VLOS: Dornberk, 11 .10.1998, MV691; VL16: Orlek, 
Sežana, 1.1977, FV35309; VL17: Mlake pri Vipavi, 11.10.1998, MV681; VL18: Vipavski 
Križ, 11.10.1998, MV700; VL29: Idrija, 13.9.1998, MV690; VL57: Slivnica, Cerknica, 
6.1971, FV51304; VL77: Velike Lašče, Ljubljana, 4.1984, FV43472; 9.1984, FV43826; 
11.1984, FV43949; FV43952; FV50887; VM60: Ljubljana, kolodvor, 10.1976, FV34232; 
stena nad vasjo, Rašica, Ljubljana, 7 .1994, FV47405; Ljubljana, 10.1976, FV51450; VM62: 
Dolge njive, Kalški greben, Savinjske Alpe, 6.1967, FVI 1508; Planina Koren, Krvavec, 
6.1974, FV26347; WL05: Podstene, Kočevski Rog, 6.1986, FV45661; WL18: Mokronog, 
9.1998, MV693; 10.1998, MV712; WM46: Bresternica, 9.1996, MV664; WM65: Lenart v 
Slovenskih Goricah, 1995, FV49987; FV49988; 7.-10.1996, FV51066; 7.-10.1997, FV51078; 
5.- 7.1994, MV606; WM95: Noršinci, Ljutomer, 10.9.1994, MV613; vrt, Veržej, Prlekija, 
11.6.1993, MV645A-D; vrtovi, žive meje, Murska Sobota, Prekmurje, 8.+9.1993, MV625; 
WM96: vrtovi, žive meje, Beltinci, Prekmurje, 9.9.1993, MV629; dvorišča, vrtovi, Dokležovje, 
Prekmurje, 9.9.1993, MV639; Bratonci, Prekmurje, 7.1994, MV648; Borovnjakova ulica, 
Murska Sobota, Prekmurje, 11.6.1994, MV649; MV675. 
44 Acta Bioloeica Slovenica, 45 (2), 2002 
18. Deroceras rodnae Grossu et Lupu, 1965 - svetli poljski slinar (Sl. 3 K) 
Z: VL57: Slivnica, Cerknica, 6.1971, FV51305; FV51306; Slivnica, gozd pod vrhom, Cerknica, 
6.1971, FV51449; VM62: Korošica- Krvavec, 9.1970, FVl2789; Dolina Korošice, Kamniška 
Bistrica, 6.1974, FV26304. 
19. Deroceras sturanyi (Simroth, 1894) - Sturanyjev poljski slinar (Sl. 3 K) 
L: WIKTOR (1996)/ VM33), VAUPOTIČ & VELKOVRH (1997)/ WM95, WM96. 
Z: VM50: Koseze, okolica ribnika, Ljubljana, 6.1997, FV5 l l l 9; VM60: Ljubljana, kolodvor, 
10.1976, FV5 l 308; WL18: Mokronog, 1998, MV680; WM65: Lenart v Slovenskih Goricah, 
7.-10.1996, FV50878; FV51069; WM95: vrt, Veržej, Prlekija, 6.11.1993, MV671; WM96: 
vrtovi, žive meje, Murska Sobota, Prekmurje, 8.+9.1993, MV624; dvorišča, vrtovi, Dokležovje, 
Prekmurje, 9.9.1993, MV640. 
20. Deroceras turcicum (Simroth, 1894) - turški poljski slinar (Sl. 3 K) 
?Deroceras forcarfi Grossu et Lupu 1961. 
L: ALTENA (1973)/ VM72, VL56; WIKTOR (1996)/ VL26. 
Z: VM0l: Tolmin, ob Soči, 4.1977, FV36805. V vzorcu je en sam osebek. 
DRUŽINA: Arionidae - lazarji 
21. Arion lusitanicus J. Mabille, 1868 - španski lazar (Sl. 3 L) 
L: WIKTOR (1996)/ VL37; VAUPOTIČ & VELKOVRH (1997)/ WM96. 
Z: UL98: ob potokuLijak,Ajševica, 27.7.1998, MV679; VL07: Komen, 11.10.1998, MV684; 
VL08: Dornberk, 11.10.1998, MV686; VL17: Mlake pri Vipavi, 11.I0.1998,MV685; VM33: 
gozd, Mošnje, levi breg Dobruše, 6.1995, FV48707; VM50: Biološko središče, Ljubljana, 
7.7.1994, MV602; VM60: Tomačevo, Ljubljana, 6.1973, FV25637; VM70: bregovi, reka 
Sava, Dolsko, 6.1970, FV12083; WL48: Brežice, 15.9.1991, FV47255; WM0l: Marija 
Reka, Prebold, 9.1996, MV666; WM65: Lenart v Slovenskih Goricah, 9.1985; 5.-7.1994, 
MV605; FV44603; 1995, FV49982; 7.-10.1996, FV51063; WM85: vrt, Ključarovci, Ljutomer, 
Prlekija, 7.8.1994, MV601; WM96: Murski logi, 0,5km W Babičov mlin, Veržej, Prlekija, 
8.7.1995, MV603; Borovnjakova ulica, Murska Sobota, Prekmurje, 11.6.1994, MV609. 
- Arion rufus (Linnaeus 1758) - veliki lazar 
Arion afer Linnaeus 1758 (partim, Arion afer rufus Linnaeus 1758), Arion empiricorum Ferussac 
1819 (partim), Arion afer rufus (Linnaeus 1758). 
L:? BOLE (1969),? BOLE (1976b),? BOLE (1977),? BOLE (1978),? BOLE (1994). 
22. Arion subfuscus (Draparnaud, 1805) - rjavkasti lazar (Sl. 3 M) 
L:? BOLE (1962),? BOLE (1966),? BOLE (1969),? BOLE (1976a),? BOLE (1976b),? 
BOLE (1979),? BOLE (1994); WIKTOR (1982)/ VM23, VLSS), WIKTOR (1996)/ VM23, 
WM00, VM03, VM13, VM33, VL27, VL26; VAUPOTIČ & VELKOVRH (1997)/ WM96; 
SLAPNIK (1998)/ WM00. 
Z: UM92: pri Dupeljskem jezeru, Planina Duplje, Julijske Alpe, 7 .1982, FV 43464; 7 .1982, 
FV43616; VL09: Trnovski gozd, 1969, FV50882; VL47: Rakov Škocjan, 4.1968, FVl 1301; 
10.1969, FV51316; Pod stenami, Laze, Planina, 6.1982, FV43485; 6.1980, FV51427; Laze, 
gozd, pasti, Planina, 9.1994, FV49947; Vranja jama, vhodna udornica, Planinsko polje, N 
rob, 6.1995, FV51434; VL48: Bistra, Vrhnika,4.1971, FV14912; 5.1977, FV36764; FV36765; 
12.6.1980, FV5 l 329; S Ljubljanski vrh, Vrhnika, 6.1977, FV36030; VL54: Sviščaki, Snežnik, 
6.1978, FV384 70; VL55: grad Snežnik, Cerknica, 9 .1977, FV365 l 9; VLSS: Leskova dolina, 
Snežnik, 6.1978, FV38718; VL56: Gorica, Cerkniško jezero, 6.1980, FV42052; VL57: 
Slivnica, 6.1971, FV15844; Slivnica, gozd pod vrhom, 6.1971, FV19775; VL69: Mestni log, 
Ljubljana, 1967, FV50888; VM02: Komna, 8.1958, FVl 1386; 7.1971, FV16806; Planina 
Govnač, Komna, 5.1972, FV20524; Bogatinsko sedlo, 5.1972, FV20527; Komna - Sedmera 
jezera, 7.1973, FV25615; Savica - Komna, Bohinj, 7.1983, FV43503; 7.1985, FV51425; 
Planina Govnač, Komna, 7 .1982, FV 43613; Komna - Bogatinsko sedlo, Julijske Alpe, 7 .1985, 
M . Vaupotič, F. Velkovrh: Slugs (Gastropoda: Pulmonata: Milacidae, Limacidae, Boettgerillidae... 45 
FV45007; 7.1985, FV45892; Savica - Komna, Julijske Alpe, 7 .1986, FV45949; Planina Na 
kraju, Komna, Julijske Alpe, 7.1985, FV46430; VM03: Ute, Dolina Triglavskihjezer, 9.1958, 
FVI 1370; VM04: Vršič, 6.1975, FV32177; 7.1975, FV35402; Prisojnik, W pobočje, Vršič, 
7.1980, FV45413; FV50863; 6.1981, FV46025; 6.1984, FV45818; 7 .1984, FV45766; 
FV45869; Prisojnik, S pobočje, 6.1977, FV46180; Kranjska Gora - Vršič, 6.1981, FV45936; 
Vršič - Trenta, 6.1975, FV51310; FV51313; Mihov dom, Vršič - Kranjska Gora, 6.1980, 
FV51318; 7.1981, FV51448; Erika (koča), Kranjska Gora - Vršič, 6.1975, FV51426; VM23: 
Kranjska dolina, Pokljuka, 9.1967, FVl 1554; VM24: Mežakla, Ravne, 6.1969, FV09902; 
FV09903; FVl 1145; 6.1971 , FV51327; Srednja Radovna, 6.1970, FV17214; VM31: nad 
vasjo Prtovč, Ratitovec, 7.1960, FVl 1480; VM34: Završnica, Jesenice, 6.1974, FV26144; 
VMS0: Tume, Šmarna gora, 6.1971, FV 19245; potok: Trnovec, dolina Ločnice, Sora, 6.1973, 
FV23526; Biološko središče, Rožnik, Ljubljana, 18.7. 1994, FV49952; VM61: Rašica, SW 
pobočje, Ljubljana, 6. 1995, FV48884; VM62: Dolge njive -Kalški greben, Savinjske planine, 
6.1967, FV 11473; Korošica - Krvavec, 4.1970, FV 12790; Dolina Korošice, Kamniška Bistrica, 
6.1974, FV26306; FV26314; Planina Koren, Krvavec, 6.1974, FV26344; 6.1978, FV38763; 
Dolge njive, Krvavec, 6.1978, FV51430; VM63: Kokrško sedlo, Savinjske planine, 1957, 
FVl 1364; VM70: breg, reka Sava, Dolsko, 6.1970, FV12089; FV51418; VM72: Velika 
planina, Savinjske Alpe, 6.1976, FV38658; VM74: Olševa, gora, 7.1974, FV26590; WL04: 
Medvedak, plato, 25.9.1998, MV695; WL0S: Podstene, Kočevski Rog, 6.1980, FV45780; 
WM00: Kum, 6.5 .1988, B1006; WM0S: Jesenkov vrh, Slovenj Gradec, 9.1996, MV677; 
WM14: Pečenikov vrh, Mislinja, 6.1998, FV511765; WM24: Pohorje, 8.1967, FV50881 ; 
WM33: Pesek - Oplotnica, Pohorje, 8. l 975 , FV3448 l; Oplotnica, Pohorje, 8.1967, FV50884; 
WM34: Pragozd, Lobnica, Pohorje, 6.1975, FV30l98; Ruško Pohorje, 8.1995, FV48911; 
Šumnik, Pohorje, 8.1967, FV50885; WM44: Areh, Pohorje, 7 .1986, FV 46087; Areh, Pohorje, 
8.1967, FV46171 ; WMSS: Zgornji Duplek, Maribor, 6.1991, FV47473; WM77: Cankova, 
Prekmurje, 1994, MV678; WM85: Stara Cesta, Ljutomer, Prlekija, 11.6.1994, MV604; WM96: 
Murski logi, W Stara Mura, S Dokležovje, Prekmurje, 12.8.1993, MV63 l. 
23. Arion circumscriptus Johnston, 1828 - očrtani lazar (Sl. 3 O) 
L: WIKTOR (1996)/ VM33. 
Z: WM77: Cankova, Prekmurje, 1994, MV653. V vzorcu je 14 osebkov. 
24. Arionfasciatus (Nilsson, 1823) - vrtičkarski lazar (Sl. 3 N) 
L: VAUPOTIČ & VELKOVRH (1997)/ WM95, WM96. 
Z: WM14: Pečenikov vrh, Mislinja, 6.1998, FV5 l 766; WM65: Lenart v Slovenskih Goricah, 
1995, FV49991; 7.-10.1996, FV51321; WM95: vrt, Veržej, Prlekija, 6.11.1993, MV642. 
WM96: vrtovi, žive meje, Beltinci, Prekmurje, 9.9.1993, MV632; Murski logi, SW Dokležovje, 
Prekmurje, 11.8.1993, MV633; vrtovi, žive meje, Murska Sobota, Prekmurje, 8.+9.1993, 
MV634; poplavni travnik, S Dokležovje, Prekmurje, 12.8.1993, MV636; Bratonci, Prekmurje, 
7 .1994, MV647; Borovnjakova ulica, Murska Sobota, Prekmurje, 11.6.1994, MV650; MV676. 
25. Arion silvaticus Lohmander, 1937 - gozdni Lazar (Sl. 3 O) 
L: WIKTOR (1996)/ VL37, VM13; VAUPOTIČ & VELKOVRH (1997)/ WM96. 
Z: VL48: Bistra, Vrhnika, 12.6.1980, FV47282; VL57: Slivnica, Cerknica, 6.1971, FV15845; 
VL 77: Velike Lašče, 4.1984, FV 43471 ; VM70: bregovi, reka Sava, Dolsko, 6.1970, FV 12090; 
WL14: pred jamo Gadina, Črnomelj , 6.1975, FV342 l 2; WL39: Anže, v vasi Brestanica, 
5.1976, FV32551 ; WM34: Pragozd, Lobnica, Pohorje, 6.1975, FV51330; WM96: gozd N 
Ižakovci, Prekmurje, 6.11.1993, MV619. 
26. Arion alpinus Pollonera, 1887 - gorski lazar (Sl. 3 P) 
L: WIKTOR (1996)/ VM33, VM14. 
Z: VL47: Pod stenami, Laze, Planina, 6.1980, FV43485; 7.1982, FV43658; Vranja jama, 
vhodna udornica, Planinsko polje, N rob, 6.1995, FV48990; VL48: S Ljubljanski vrh, Vrhnika, 
46 Acta Biologica Slovenica, 45 (2) , 2002 
6.1977, FV36031; FV51433; Planinsko polje, N breg, Planina, 6.1985, FV46537; VL69: 
Rudnik pri Ljubljani, 4. 1969, FV09906; VL 76: Travna gora, 6.1972, FV23744; VL 77: Podgora, 
Dobrepolje, 1.1976, FV32038; VM02: Planina Govnač , Komna, 5.1972, FV20526; Savica -
Komna, Julijske Alpe, 7.1982, FV43629; 7.1983, FV51420; 7.1985, FV45253; 7.1986, 
FV45787; FV46002;FV51424; VM04: Erika (koča), Kranjska Gora- Vršič, 6.1975, FV32157; 
Mojstrovka, E pobočje, 6.1974, FV34234; Mihov dom, Kranjska Gora - Vršič, 6.1980, 
FV45804; 7.1981, FV42021 ; Vršič, 7.1975, FV51326; VM23: Bohinjska Bela, Bled, 6.1995, 
FV 48951; VM32: Brezovica, Kropa, 6.1998, FV5 l 722; VM33: Mošnje, gozd, Dobruša, levi 
breg, Brezje, 6. 1995, FV5 l 428; VM34: ? levi breg od mosta do jezu, Breg, Žirovnica (S 1893a), 
23 .5.1996, BI007; VM41: Moškrin, Škofja Loka, 6.1998, FV50746; VM42: Amševajama, 
okolica (Udin boršt), Sp. Duplje, Kranj, 6. 1995, FV48682; 8.1995, FV 48673; VM43: Jelendol, 
Tržič, 6.1974, FV28238; VMS0: Tume, Grmada, Tacen, Ljubljana, 6.1997, FV50614; Podutik, 
nad kamnolomom, Ljubljana, 6.-7.1997, FV50984; VM51: pobočje: reka Sava, Mavčiče, 
Smlednik, 6.1998, FV51781; VM60: stena nad vasjo, Rašica, Ljubljana, 7.1994, FV47404; 
VM62: Dolge njive, Krvavec, 6.1978, FV38690; desno pobočje: Dolina Korošice, Stahovica, 
6.1998, FV51056; Stari grad, Kamnik, 6.1998, FV51478; Davovec, Cerklje, 6.1998, FV51619; 
VM63: Rokovnjaške luknje, Kamniška Bistrica, 7.1979, FV50864; VM83: Zamernikova 
jama, Konjski vrh, Luče v Savinjski dolini, 4.1978, FV45373; WL48: Vitovec, E Malence, 
Brežice, 7 .1994, FV 47332; WMOO: Škofja Riža, Dobovec, Kum, 5.6.1991, BI002; WM05?15: 
Jesenkov vrh, Slovenj Gradec, 7.1994, FV47366; 9.1996, MV669; WM14: Pečenikov vrh, 
Mislinja, 6.1998, FV51452. 
27. Arion distinctus J. Mabille, 1868 - navadni vrtni lazar (Sl. 3 O) 
Z: WM65: Lenart v Slovenskih Goricah, 1995, FV49993; 7.-10.1996, FV51067; FV51079. 
Skupaj je v vseh treh vzorcih 43 osebkov. 
- Arion hortensis Ferussac 1819 - vrtni lazar 
L:? BOLE (1962),? BOLE (1969),? BOLE (1979a) . 
. 1 1 1 
' 1 -f-- ltrJ.'rh ->- 1 \ h :JMr ' 1 1 ~ ) ~ ~ - - () - .... --- -
~ " .,. II l ~ .. - -+i:r -~ ' C 
C 1 1 \ ' '" ' 
utrla 
' - ' 
,-, . - ~ - • -.. , _ - -'"" "' " ·'-· -'"'-~ -i-. • 1  • ~ \ ,,_-r • ~- ' • 'l. -~- '\ t) ,_ Mi/as; 11igrica1i, ~ ... ,. • Tundonitl robid 
1 ,, ~ ( () 
1 , () () " 1 Tundoriia budo;wsrmsis 
" ~ 
o ,,. :., Tunc;b,iwrora 
v e 1 I - ( ~ Tcmdonia ,nl,mai l -· .., TandonJa sl/N'Olhl .., "" 1/1\. 1 v - A Cru\ 
·~ ( o 
,, 00 • .., { ~ Tcmdmria rlllli,·a 
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o ,-
• J ••• -• • l 1 "" 
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,, • ' ~ - • • -~ - -
~ "'-
\ 
_.., 
1,... - 1• - - :-.._ ;t .... ~ "\, lll 'l. ,~ 
f - I"\ • - ~ v Tandonla SO>t-tf'byf f ,-. I"\ • -f-- - i - • - Mufaco!itm1X1t1ruttA:i 
1 ,, ,~ 1 
1 ,, • , , 1 
1 " 
...._ 
C', ; 1 1 ' I 
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..... v ,,, 
r l - 1 
l 1 
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er.ut 1--1 C I" 
• ..... , . v '"' 1 1 
re r ...... 
• l II .., ' -- 1-
c,on 1,- D 
M. Vaupotič, F. Velkovrh: Slugs (Gastropoda : Pulmonata: Milacidae, Limacidae, Boettgerillidae ... 47 
T 1 1 
1 ' 1 ~il.!!L-..... """ .~ r-; ~L.... " I -.. ,. .... " l \. '-. • 1\. • - ' - •• ., fl..~ • -... • • • ,. ·- II . ~ ' - - • • • R '-"f;l • i le .- "":;: ••• ,_ • l e '- T '- 1-,-l -~ ~ i'" ••• I'\. :11 ,"[ ,_ ,i '-'"' e l'i" • -~- ''- ••• ,.. _.., L_,:_ ... e l'I •• • • UhtlfLlnn/a moryinato . ·- limax dnerMniger r, •• ·, - 1 1 •• • ,,. ,,. r II • • ., ---, ,  .. •• ••1• '- 1 • •• ..... ,_ r:, R • • -~ ., • L..'-• 1 ' , 1· ' w ,~ L ~i,. " l,- E - ·--··· . 
,. J 
. , • • v 
l • ': { , • -· -It=-, 
'J 
1 M ... 
1 crn 1,- 1 F 
T r 1 1 1 
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~ ~ f\_ ' - l,e,_ 
: l'i" H~ -. 
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~ '-
t::;:~ I\._ -K' 1 
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1 1) 
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II 
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• 1 '-
i'i rr ,L.>- ,_, 
'- ,_ G 1:1.uth II 
II 
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::,: - ], __ ... H CrHU 
Figure 2: Distribution of slugs species in Slovenia; A - Milax nigricans, Tandonia budapestensis, 
Tandonia rara, Tandonia reuleauxi, Tandonia simrothi, B - Tandonia robici, Tandonia rustica, C -
Tandonia sowerbyi, D - Malacolimax mrazeki, E - Lehmannia marginata, F - Limax cinereoniger, G 
- Limax maximus, H - Limaxflavus, Boettgerilla pallens, Deroceras laeve. 
Slika 2: Razširjenost vrst v Sloveniji; A - Milax nigricans, Tandonia budapestensis, Tandonia rara, 
Tandonia reuleauxi, Tandonia simrothi, B - Tandonia robici, Tandonia rustica, C - Tandonia sowerbyi, 
D - Malacolimax mrazeki, E - Lehmannia marginata, F - Limax cinereoniger, G - Limax maximus, H 
- Limaxflavus, Boettgerilla pallens, Deroceras laeve. 
1. . 7 
1 
1) ~ il!1-
'-'-"' 
1trh ( 1\: 
"' • - ' ~ ;-., . iliiL J .._ -, _ • ,_ •• 1 ' , . ........ h.,_ 
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-
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Duocera.r rt:tk:ulatum 
" • • ,.. \. • -~·. ; C\. 1 11 ee • "- 1 
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llr 
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' • it' ~ 11 ; 
L> ..... ,-,-
tr,Htl 
1...-1 J 
48 Acta Biologica Slovenica, 45 (2), 2002 
1 1 • i 
< 
t.•u1rt1 i-.!.!!l!!rF--r-, ,_ 
1 1 •-L+ , ..... 1 1 h a 1r 
I'-. lllUl'l.11 1 
I' "-. , .... "- . - ,- ~ • ~,.-- '' 
1, I I ~ 1 ~:r 
~ ~ • e h ":c:e, -= C-- - i1 ... 1- >1-b.. fr!<c 1,· - - ,_ ,,._ , ..... 1 ~ -- l'I. IC () ,_ i v 1 -- ''- ,_ - . L-1...:::L-
" ' o Deroceras rmbwe E 
,-; !\ • • ·-·- •- J .Arionlusircmicttr 1, r ·~ "- ~ • • M DtNwu·as sh1nmyi - '-' - o () !J Du ocerasturcicum -,, () () v 
' I -
~ f\" 1;,al.L ~- :., " ..... - ~ 
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1 ,,, ~ 1 
1 • ~ 1 1 1: • .... IJ v -~ l \ -"- ~ 
J 
r,oatl 
IJ L 
1 -
' Ln1r11 I-- J j!!fl. "-f"'\ - ~ ,. 1- ;~ 1- C-' 'J~:-, 1urla 
1 
, 
h•1r 
' 
-
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~ 
,-
~ • ,. • ~ e1• •• "' 1-
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' 
,1. 
~~ '- •• ,,. ! I.A ~~ I'\.. ' - 1 -
" • '. • f ,. Aritm suhfascu.1· " '' 1 -· :;; - "' hionfasdaJus 1/ • • '- t 1 ., ~ 1 ,, • '- I 1 1 /J 1 I :;_ •• ' IJ J • .1-" ~ .... ,--• • l l • \ 'iP.L 1 
M 1 Cr..11t1 ,,. 
1 ' 1, 1, 
II' 
,_ 
1 ~ 
' "' ..; ' ..... - N Ct'.flat '-
1 1 
,-. 
nu!rh 
\ 1.. . .!I.• "' -r-, ~ 
1 1 1 ~ 1-
" 1 1 1 ' ' 
hstl'h 1 
• llutU 1-
I' r-,,. ~ -- ri.- Q t.. 's 1--,_ ·• 1-1-,. 
<i:_; 
,_ ,_ • 1 ... \ ""' rr '"h .._ _, "" 'l. - _..,.. J ' • f ,,frj,msilvaJi~·ILv 
1 ,,. ---~ ,.,_ J .. 
1~ • 
1 Ariondistlndus 
• • • r-- o ,, ! J .Arion cinwn:,ai~· 
I ~ -
~ •1 ,i,,"' __, -- ,- ~ 1-,.. o Crut! 
-- ' ,_ __ --, •• - ), ..... ~ 
---.--s 
~ 
- l cc'-
~ ••• • .. - ·'---t7 • '-' •• • 'l. -!':" - ~ ..... 
~ •• • ,, A.rionolpinus 
1 ,, • ,._ 1 1 IJ • 1 • • ..... -: J • v 1: I - 1 1l \ 1 \ -
/' era.ti 
,,, 1 p 
Figure 3: Distribution of slugs species in Slovenia; I - Deroceras lothari, J - Deroceras reticulatum, 
K - Deroceras rodnae, Deroceras sturanyi, Deroceras turcicum, L - Arion lusitanicus, M - Arion 
subfuscus, N -Arionfasciatus, O -Arion silvaticus, Arion distinctus, Arion circumscriptus, P-Arion 
alpinus. 
Slika 3: Razširjenost vrst v Sloveniji; I - Deroceras lothari, J - Deroceras reticulatum, K - Deroceras 
rodnae, Deroceras sturanyi, Deroceras turcicum, L - Arion lusitanicus, M - Arion subfuscus, N -
Arionfasciatus, O -Arion silvaticus, Arion distinctus, Arion circumscriptus, P -Arion alpinus. 
M. Vaupotič, F. Velkovrh: Slugs (Gastropoda: Pulmonata: Milacidae, Limacidae, Boettgerillidae ... 49 
Razprava 
Po dosedanjih izsledkih živi na območju Slovenije sedemindvajset vrst golih polžev. Za enaindvajset 
od teh vrst lahko zasledimo podatke v literaturi, za šest vrst (Milax nigricans, Tandonia budapestensis, 
Tandonia rara, Tandonia sowerbyi, Deroceras rodnae, Arion distinctus) pa so to prvi podatki za 
Slovenijo. Od teh imajo tri vrste (Milax nigricans, Tandonia rara, Arion distinctus) samo po eno 
najdišče v Sloveniji. Milax nigricans in Arion distinctus sta zunaj Slovenije precej razširjeni, zato 
lahko pričakujemo, da sta tudi v Sloveniji bolj razširjeni. WIKTOR (1996) piše za vrsto Tandonia 
rara, da je razširjena na Velebitu in v Dalmaciji. !zgleda, da poteka severna meja njenega areala v 
Sloveniji. Podobno velja najverjetneje tudi za vrsto Tandonia reuleauxi. V Sloveniji je znana samo iz 
ene lokalitete. Vrsta je sicer razširjena v južnih delih nekdanje Jugoslavije (WIKTOR 1996), pod 
imenom Aspidoporus limax jo je z območja Kranjske navedel že WAGNER (1931 ). WIKTOR ( 1996) 
opozarja, daje starejše podatke v zvezi s to vrsto treba revidirati. Vseh vrst, za katere so znane lokali tete 
le iz enega ali dveh kvadratov UTM, je sedem (Milax nigricans, Tandonia rara, Tandonia reuleauxi, 
Boettgerilla pallens, Deroceras laeve, Arion distinctus, Arion circumscriptus). 
WIKTOR (1996) navaja tudi šest vrst (Milax gagates, Tandoniafejervaryi, Malalcolimax tenellus, 
Deroceras agreste, Ar ion rufus, Arion hortensis), za katere je našel podatke v literaturi, vendar jih sam 
na območju Slovenije ni ugotovil. Tudi pri obdelavi vzorcev iz naših zbirk nisva ugotovila nobene od 
navedenih vrst. Sinantropna vrsta Milax gagates je znana iz neke jame blizu Trsta. Za vrste Malacolimax 
tenellus, Deroceras agreste in Arion rufus nimamo zanesljivih podatkov; so pa znana najdišča v 
Avstriji (REISCHUTZ 1986), ki so blizu avstrijsko-slovenske meje in zato je mogoče, da te vrste 
živijo tudi v Sloveniji. WIKTOR (1996) piše, da bi bilo treba razširjenost vrste Malacolimax tenellus 
južno od Alp podrobneje raziskati, zaradi možne zamenjave z vrsto Malacolimax mrazeki. Sodeč po 
lokalitetah blizu slovensko-avstrijske meje in po podatku, ki ga navajata WOLF & RAHLE ( 1987), 
najverjetneje poteka južna meja areala vrste Malacolimax tenellus v Sloveniji. Vrsto Arion rufus je 
marsikje v Evropi izpodrinila vrstaArion lusitanicus (FECHTER & FALKNER 1990). To najverjetneje 
velja tudi za Slovenijo, kjer so vrsto Ar ion lusitanicus ugotovili WIKTOR ( 1996) ter VAUPOTIČ & 
VELKOVRH (1997). Najstarejši vzorec s to vrsto v naših zbirkah je iz leta 1970, medtem ko vrste 
Arion rufus sploh nismo zanesljivo ugotovili. 
Zaradi medsebojne zunanje podobnosti vrst rodu Lehmannia bo v bodoče treba pregledati prav 
vse primerke v razpoložljivih vzorcih, saj bi bilo mogoče, daje tudi v Sloveniji razširjena katera izmed 
drugih vrst tega rodu. Vrsta Lehmannia nyctelia je na primer znana iz Madžarske (PINTER et al. 1979, 
WIKTOR & SZIGETHY 1983) in Avstrije (REISCHUTZ 1986), vrsta Lehmannia brunneri pa iz 
Hrvaške (WIKTOR 1996). 
V Avstriji (REISCHUTZ 1986) in na Hrvaškem (WIKTOR 1996) je po 33 vrst golih polžev. Od 
sosednjih držav je manj vrst kot v Sloveniji le na Madžarskem, kjer je znanih 22 (WIKTOR & 
SZIGETHY 1983). Slovenija je na stičišču štirih naravnogeografskih makroregij (Alpe, Dinarsko 
gorstvo, Sredozemlje, Panonska kotlina) in s tem pod vplivom različnih dejavnikov, ki vplivajo na 
razširjenost vrst, zato bi tukaj pričakovali več kot 30 vrst golih polžev. 
Pregled nahajališč golih polžev v Sloveniji (Sl. 1) kaže, da podatki prihajajo iz približno polovice 
kvadratov UTM. Nekatere podatke iz literature bi bilo treba preveriti s ponovnim pregledom navedenih 
lokalitet. Število ugotovljenih vrst v posameznem kvadratu UTM je največje tam, kjer je potekalo 
intenzivno nabiranje živali. Takšna sta kvadrata WM65 in WM96 z enajstimi oziroma dvanajstimi 
ugotovljenimi vrstami, kar predstavlja skoraj polovico vseh vrst, znanih iz Slovenije. Med ugotovljenimi 
vrstami iz teh kvadratov sta tudi dve v Sloveniji redki vrsti (Arion distinctus poznana iz kvadrata 
WM65 ter Boettgerilla pallens iz kvadrata WM96), kar poleg velikega števila ugotovljenih vrst kaže 
na uspešnost intenzivne metode vzorčenja. Slike razširjenosti vrst kažejo, da je poznavanje arealov 
mnogih vrst v Sloveniji še zelo nepopolno, iz nekaterih območij nimamo sploh nobenih podatkov in 
zato bi v bodoče bilo smiselno intenzivno vzorčiti predvsem na teh območjih . 
50 Acta Biologica Slovenica, 45 (2), 2002 
Zahvala 
Za vse nasvete se zahvaljujeva prof. dr. Borisu Sketu. Prav tako se zahvaljujeva prof. dr. Andrzeju 
Wiktorju za prijazno sodelovanje pri determinaciji nekaterih vzorcev. 
Summary 
Slugs are among the least investigated snails in Slovenia. SIMROTH described two new slug 
species from this area (1885, 1910). BOLE mentioned 14 species in his identification key (1969). 
WIKTOR (1996) presented the first accurate data about slugs in Slovenia in his work on slugs from 
the form er Yugoslavia. VAUPOTIČ & VELKOVRH ( 1997) found out 11 species in the north-eastem 
Slovenia. That was the first recording of Boettgerilla pallens in Slovenia. 
With the analysis of data from the literature and the study of 686 samples of slugs from three 
collections (J. Bole collection ofBiological Institute ZRC SAZU, our own collections), collected in 
years from 1957 to 1998, a list of slug species and their distribution in Slovenia is given. Data of 
species distribution are presented in the UTM squares (10 x 10 km). 
The number of identified slug species in Slovenia is 27. For six species (Milax nigricans, Tandonia 
budapestensis, Tandonia rara, Tandonia sowerbyi, Deroceras rodnae, Arion distinctus) this is the 
first data for Slovenia. In Slovenia is the northern border of the distribution area for the species 
Tandonia rara and Tandonia reuleauxi. We suppose that the real number of the species is o ver thirty; 
we expect some species known from neighbouring countries (for example Milax gagates, Lehmannia 
nyctelia, lehmannia brunneri, Deroceras agreste, Arion rufus) to be presen! in Slovenia. The species 
Malacolimax tenellus is probably presen! in Slovenia due to the localities near the Slovene-Austrian 
border and according to the data by WOLF andRA.HLE (1987). WIKTOR (1996) writes that its wide 
range south of the Alps should be investigated in detail, because of the possibility to confound it with 
Malacolimax mrazeki. 
The data on slugs localities in Slovenia is known from only one half of the UTM squares. The 
largest number of species was found in the UTM squares, where the sampling was very intensive; e. 
g., in squares WM65 with 11 species and in WM96 withl2 species. In square WM65, a very succesful 
sampling was done in one locality, where 5 cm thick wet boards were placed during the night out on the 
ground, and under those boards the slugs were collected in the morning. 
In the future, more intensive research will be needed to geta more accurate number of the slugs 
species and their distribution in Slovenia. Our data enable us to direct future investigations. Some 
locations, which are mentioned in the literature, will have to be checked up and also the areas, for which 
there are no data for slugs, will have to be researched. 
Literatura 
ALTENA C. O. VAN REGTEREN 1973: On two species of Deroceras from Jugoslaviaand Austria. 
Basteria 37: 85-92. 
ALTENA C. O. VAN REGTEREN 1977: Notes on land slugs. 25. On little known species of the 
genus Milax. Basteria 41: 65-70. 
BACKELJAU T & M. VAN BEECK 1986: Epiphallus anatomy in the Arion hortensis species 
agregate (Mollusca, Pulmonata). Zool. ser. 15(1): 61-68. 
BACKELJAU T & L. DE BRUYN 1990: On the infrageneric systematics of the genus Arion Ferussac, 
1819 (Mollusca, Pulmonata). Buli. - Inst. r. sci. nat. Belg. 60: 35-68. 
M. Vaupotič , F. Velkovrh: Slugs (Gastropoda: Pulmonata: Milacidae, Limacidae, Boettgerillidae ... 51 
BOLE J. 1962: Mehkužci Triglavskega narodnega parka in okolice (Mollusca. Gastropoda, Bivalvia). 
Varst. narave 1: 57-85. 
BOLE J. 1966: Mehkužci in zoogeografski položaj Rakovega Škocjana. Varst. narave 5: 129-137. 
BOLE J. 1969: Ključi za določevanje živali - Mehkužci. Mollusca. Inštitut za biologijo Univerze 
v Ljubljani in Društvo biologov Slovenije, Ljubljana, 115 pp. 
BOLE J. 1976a: Mehkužci Notranjskega Snežnika in okolice. Varst. narave 9: 55-63. 
BOLE J. 1976b: Malakološke razmere v mraziščih. Razprave IV razreda SAZU 19(5): 151-183. 
BOLE J. 1977: Mehkužci Šmarne gore. Varst. narave 10: 57-62. 
BOLE J. 1979a: Mehkužci Cerkniškega jezera in okolice. Acta carsologica 8(3): 204-236. 
BOLE J. 1979b: Malakološke raziskave v nekaterih fitocenozah Slovenije. Savez društava 
ekologa Jugoslavije (II. Kongres ekologa Jugoslavije), Zagreb: 183-190. 
BOLE J. 1981: Zoogeographische Analyse der Landschnecken des dinarischen Gebietes 
Sloweniens. Razprave IV razreda SAZU 23(4): 125-146. 
BOLE J. 1992a: Rdeči seznam ogroženih kopenskih in sladkovodnih mehkužcev (Mollusca) v Sloveniji. 
Varst. narave 17: 183-189. 
BOLE J. 1992b: Mehkužci Kraškega roba. Proteus 54(6-7): 234-235 . 
BOLE J. 1994: Polži tal. Prirodoslovno društvo, Ljubljana, 38 pp. 
DE WINTER A.J. 1984: The Arion hortensis complex (Pulmonata, Arionidae): designation of types, 
descriptions, and distributional patterns, with special reference to the Netherlands. Zool. 
Med. Leiden 59: 1-17. 
DE WINTER A. J. 1989: Arion lusitanicus Mabille in Nederland (Gastropoda, Pulmonata, Arionidae). 
Basteria 53: 49-51. 
ERJAVEC F. 1877: Die malakologischen Verhaltnisse GEFURSTETEN GRAFSCHAFfGORZ im 
osterreichischen Kiistenlande, Gorz, Mailing, 82 pp. 
FECHTER R. & G. FALKNER 1990: Steinbachs Naturfiihrer: Weichtiere. Mosaikverlag, Miinchen, 
287 pp. 
GARRIDO C., J. CASTILLEJO & J. IGLESIAS 1995: The Arion subfuscus complex in the eastern 
part of the Iberian Peninsula, with redescription of Arion subfuscus (Draparnaud 1805). 
Arch. Moll., Frankfurta. M. 124 (1/2): 103-118. 
GROSSU A. V. 1972: Fiinf neue Arten der Gattung Deroceras von der Balkanhalbinsel in der Sammlung 
des Naturhistorischen Museums in Wien (Gastropoda, Limacidae). Ann. Naturhist. Mus. 
Wien 76: 639-648 . 
JAECKEL S. sen.& W. MEISE 1956: Ober Land- und Siisswasserschnecken Jugoslawiens und 
Albaniens. Mitt. Hamburg. Zool. Mus Inst. 54: 21-32. 
JAECKEL S.G., W. KLEMM & W. MEISE 1958: Die Land- und Siisswasser-Mollusken der 
nordlichen Balkanhalbinsel. Dresden, Abh. Ber. Mus. Tierk. 23 (2): 141-205. 
KERNEY M.P., R.A.D. CAMERON & J.H. JUNGBLUTH 1983: Die Landschnecken Nord- und 
Mitteleuropas. Verlag Paul Parey, Hamburg und Berlin, 384 pp. 
KOS F. 1933. Vodnik po zbirkah Narodnega muzeja v Ljubljani, Prirodopisni del. Narodni muzej v 
Ljubljani, Ljubljana, 219 pp. 
LIKHAREV I.M. & A. WIKTOR 1980: The fauna of slugs of the USSR and adjancent countries 
(Gastropoda terrestria nuda). Fauna SSSR (NS 122) Molluski III, 3 (5) . Leningrad, 434 pp. 
LUPU D. 1977: Le polymorphisme chez quelveques especes appartenant aux familles Limacidae et 
Arionidae de Roumanie. Malacologia 16( 1 ): 21-33. 
PINTER L., A. RINCHNOVSKY & A.S. SZIGETHY 1979: A magyarorszagi recens puhatestiiek 
elterjedese. Budapest, A tudomanoyos ismeretterjszto tarsulat, Bacskiskun megyei szervezete, 
Biologiai szakosztalyanak, Idoszakos malakologiai kozlemenyei, Soosiana, 351 pp. 
QUICK H. E. 1960: British slugs (Pulmonata: Testacellidae, Arionidae, Limacidae), Bulletin of the 
British Museum (Natura! History) Zoology 6: 103-226. 
52 Acta Biologica Slovenica, 45 (2), 2002 
REISCHUTZ P. L. 1978: Bemerkungen zu Deroceras klemmi Grossu, 1972. Mitt. Abt.Zool. 
Landesmus. Joanneum, Graz 7(1): 39-44. 
REISCHUTZ P. L. 1986: Die Verbreitung der Nacktschnecken 6sterreichs (Arionidae, Milacidae, 
Limacidae, Agriolimacidae, Boettgerillidae ). Supplement 2 des Catalogus Faunae Austriae, 
6sterreichische Akademie der Wissenschaften, Wien, 190 pp. 
SAJOVIC G. 1908: Kranjski mehkužci (Mollusca camiolica). Izvestja Muz. društva za Kranjsko, 21 pp. 
SCHMIDT F. J. 1847: Systematisches Verzeichniss der in der Provinz Krain vorkommenden Land-
und Suesswasser-Conchylien (mit Angabe der Fund-Orte). Blasnik, Ljubljana, 27 pp. 
SIMROTH H. 1885: Versuch einer Naturgescichte der deutschen Nacktschnecken und ihrereuropaischen 
Verwandten. Zeitschrift fiir Wissenschaftliche Zoologie 42: 203-366. 
SIMROTH H. 1910: Nacktschneckenstudien in den Siidalpen. Abh. Senckenb. Natforsch. Ges. 32: 
275-348. 
SLAPNIK R. 1998: Polži Kuma (Gastropoda: Prosobranchia, Pulmonata), Posavsko hribovje 
(Slovenija). Acta biol. slov. 42(1): 57-67. 
SOUTH A. 1992: Terrestrial slugs. Chapman & Hall, London, 428 pp. 
STOSSICH A. 1899: Contibuzione alla Fauna malacologica terrestre e Fluviatile del territorio di 
Trieste. 35 pp. 
VAUPOTIČ M . & F. VELKOVRH 1997: Prispevek k poznavanju favne mehkužcev (Mollusca) v 
severovzhodni Sloveniji. Acta biol. slov. 41 (1): 37-45. 
VAUPOTIČ M . 2002: Taksonomska in biogeografska analiza kopenskih golih polžev (Gastropoda: 
Pulmonata p.p.) v Sloveniji. Magistrska naloga. Biotehniška fakulteta, Oddelek za biologijo, 
Ljubljana, 107 pp. 
VON PROSCHWITZ T. 1989: Arion lusitanicus Mabille - en for Sverige ny snigelart. Goteborgs 
Naturhistoriska museum: 43-53 
WAGNER H. I 93 I: Diagnosen neuer Limaciden aus dem Naturhistorischen Museum in Wien. 
Zoologischer Anzeiger 95: 194-202. 
WIKTORA. 1973: Die Nacktschnecken Polens. Monografie fauny palski 1, Warszawa, Krakow, 279 pp. 
WIKTOR A. 1982: Contribution to the knowledge of the slugs ofYugoslavia (Arionidae, Milacidae, 
Limacidae, Agriolimacidae - Gastropoda, Pulmonata). Ann . Zool. , 36(24): 465-489. 
WIKTOR A. 1983: The Slugs of Bulgaria (Arionidae, Milacidae, Limacidae, Agriolimacidae -
Gastropoda, Stylommatophora). Ann. Zool. 37(3): 71-206. 
WIKTOR A. 1987: Milacidae (Gastropoda, Pulmonata)- systematic monograph. Ann. Zool. 41(3): 
154-313. 
WIKTOR A. 1989: Limacoidea et Zonitoidea nuda. Slimaki pomrowioksztaltne (Gastropoda, 
Stylommatophora), Fauna Poloniae 12, Warszawa, 208 pp. 
WIKTOR A. 1996: The Slugs of the Former Yugoslavia (Gastropoda terrestria nuda - Arionidae, 
Milacidae, Limacidae, Agriolimacidae). Ann. Zool. 46( 1-2): 1-110. 
WIKTORA. 1997. Endemism of slugs within the Balkan Peninsula and adjancent islands (Gastropoda: 
Pulmonata: Arionidae, Milacidae, Limacidae, Agriolimacidae). Genus 8(1): 205-22 I. 
WIKTOR A. & N. MILANI 1995: Contribution to the knowledge of two scarcely known Alpine 
slugs, Tandonia simrothi (HESSE, I 923) and Deroceras planarioides (SIMROTH, 1910) 
(Gastropoda: Pulmonata: Milacidae et Agriolimacidae). Malakol. Abh. 17( 12): 151-160. 
WIKTORA. & A.S. SZIGETHY 1983: The distribution of slugs in Hungary (Gastropoda: Pulmonata), 
Soosiana 10() !): 87-111. 
WOLF M., RAHLE W. 1987. Ergebnisse einer Molluskenexcursion in die westlichen Julischen 
Alpen . Mitt. Dtsch . Malakozool. Ges. 41: 31- 41. 
NAVODILA AVTORJEM 
l. Vrste prispevkov 
a) ZNANSTVENI ČLANEK je celovit opis originalne raziskave in vključuje teoretični pregled 
tematike, podrobno predstavljene rezultate z diskusijo in sklepe ter literatumi pregled: shema IMRAD 
(Introduction, Methods, ResultsAnd Discussion). Dolžina članka, vključno s tabelami, grafi in slikami, 
ne sme presegati 15 strani; razmak med vrsticami je dvojen. Recenzirata ga dva recenzenta. 
b) PREGLEDNI ČLANEK objavi revija po posvetu uredniškega odbora z avtorjem. Število strani 
je lahko večje od 15. 
c) KRATKA NOTICA je originalni prispevek z različnih bioloških področij (sistematike, bioke-
mije, genetike, mikrobiologije, ekologije itd.), ki ne vsebuje podrobnega teoretičnega pregleda. Njen 
namen je seznaniti bralca s preliminarnimi ali delnimi rezultati raziskave. Dolžina na sme presegati 5 
strani. Recenzira ga en recenzent. 
d) KONGRESNA VEST seznanja bralce z vsebinami in sklepi pomembnih kongresov in posveto-
vanj doma in v tujini. 
e) DRUŠTVENA VEST poroča o delovanju slovenskih bioloških društev. 
2. Originalnost prispevka 
Članek, objavljen v reviji Acta Biologica Slovenica, ne sme biti predhodno objavljen v drugih revijah 
ali kongresnih knjigah. 
3.Jezik 
Teksti naj bodo pisani v angleškem jeziku, izjemoma v slovenskem, če je tematika zelo lokalna. 
Kongresne in društvene vesti so praviloma v slovenskem jeziku. 
4. Naslov prispevka 
Naslov (v slovenskem in angleškem jeziku) mora biti kratek, informativen in razumljiv. Za naslovom 
sledijo imena avtorjev in njihovi polni naslovi (če je mogoče, tudi štev. faxa in e-mail). 
5. Izvleček - Abstract 
Podati mora jedrnato informacijo o namenu, uporabljenih metodah, dobljenih rezultatih in zaključkih. 
Primerna dolžina za znanstveni članek naj bo približno 250 besed, za kratko notico pa l 00 besed. 
6. Ključne besede - Keywords 
Število naj ne presega 1 O besed; predstavljati morajo področje raziskave, obravnavane v članku . 
Člankom v slovenskem jeziku morajo avtorji dodati ključne besede v angleškemjeziku. 
7. Uvod 
Nanašati se mora le na tematiko, ki je predstavljena v članku ali kratki notici. 
8. Slike in tabele 
Tabele in slike (grafi, dendrogrami, risbe, fotografije idr.) naj v članku ne presegajo števila 10, v 
članku naj bo njihovo mesto nedvoumno označeno . Ves slikovni material naj bo oddan kot fizični 
original (fotografija ali slika). Tabele in legende naj bodo tipkane na posebnih listih (v tabelah naj bodo 
le vodoravne črte) . Naslove tabel pišemo nad njimi, naslove slik in fotografij pod njimi. Naslovi tabel in 
slik ter legenda so v slovenskem in angleškem jeziku. Pri citiranju tabel in slik v besedilu uporabljamo 
okrajšave (npr. Tab. 1 ali Tabs. 1-2, Fig. l ali Figs. 1-2; Tab. 1 in Sl. 1). 
9. Zaključki 
Članek končamo s povzetkom glavnih ugotovitev, ki jih lahko zapišemo tudi po točkah . 
10. Povzetek - Summary 
Članek, ki je pisan v slovenskem jeziku, mora vsebovati še obširnejši angleški povzetek. Velja tudi 
obratno. 
11. Literatura 
Uporabljene Iiteratume vire citiramo med tekstom. Če citiramo enega avtorja, pišemo ALLAN (1995) ali 
(ALLAN 1995), če sta dva avtorja (TRINAJSTIC & FRANJIC 1994), če je več avtorjev (PULLIN & al. I 995). 
Kadar navajamo citat iz večih del hkrati, pišemo (HoNSIG-ERLENBURG & al. 1992, W ARD 1994a, ALLAN 
1995, PULLIN & al. 1995). V primeru, če citiramo več del istega avtorja, objavljenih v enem letu, 
posamezno delo označimo s črkami a, b, c itd. (W ARD 1994a,b ). Če navajamo dobesedni citat, označimo 
dodatno še strani: TOMAN (1992: 5) ali (TOMAN 1992: 5-6). Literaturo uredimo po abecednem redu, 
začnemo s priimkom prvega avtorja, sledi leto izdaje in naslov članka, mednarodna kratica za revijo 
(časopis), volumen poudarjeno, številka v oklepaju in strani. Npr.: 
HoNSIG-ERLENBURG W., K. KRAINER, P. MILDNER & C. WmsERR 1992: Zur Flora und Fauna des 
Webersees. Carinthiall 182/102 (1) : 159-173. 
TRINAJSTIC I. & J. FRANJJc 1994: Ass. Salicetum elaeagno-daphnoides (BR.-BL. et VOLK, 1940) M. 
MOOR I 958 (Salicion elaeagni) in the Vegetation in Croatia. Nat. Croat. 3 (2): 253-256. 
W ARD J. V. I 994a: Ecology of Alpine Streams. Freshwater Biology 32 (1): 10-15. 
W ARD J. V. 1994b: Ecology of Prealpine Streams. Freshwater Biology 32 (2): 10-15. 
Knjige, poglavja iz knjig, poročila, kongresne povzetke citiramo sledeče: 
ALLAN J. D. 1995: Stream Ecology. Structure and Function ofRunning Waters, 1 st ed. Chapman & Hall, 
London, 388 pp. 
PULLIN A. S., I. F. G. McLEAN & M. R. WEBB 1995: Ecology and Conservation of Lycaena dispar: 
British andEuropean Perspectives. In: PuLLINA. S. (ed.): Ecology and Conservation ofButterflies, 
1 st ed. Chapman & Hall, London, pp. 150-164. 
TOMAN M. J. 1992: Mikrobiološke značilnosti bioloških čistilnih naprav. Zbornik referatov s posveto-
vanja DZVS, GozdMartuljek, pp. 17. 
12. Format in oblika članka 
Članek naj bo napisan v programu Wordfor Windows ali WordPerfect, v pisavi "Times New Roman CE 
12" z dvojnim medvrstnim razmakom in levo poravnavo ter s 3 cm robovi naA4 formatu . Odstavki naj 
bodo med seboj ločeni s prazno vrstico. Naslov članka in poglavij naj bodo pisani krepko in v velikosti 
pisave 14. Vsa latinska imena morajo biti napisana ležeče . V besedilu navedemo uporabljene nomenklatur-
ne vire. Tabele in slike so posebej priložene tekstu. Glavnemu uredniku je potrebno oddati original, dve 
kopiji in disketni zapis na disketi 3,5" ( odda avtor po opravljenih strokovnih in jezikovnih popravkih). 
13. Recenzije 
Vsak znanstveni članek bosta recenzirala dva recenzenta ( en domači in en tuji), kratko notico pa domači 
recenzent. Avtor lahko v spremnem dopisu predlaga tuje recenzente. Recenziran članek, ki bo sprejet v 
objavo, popravi avtor. Po objavi prejme 50 brezplačnih izvodov. V primeru zavrnitve se originalne 
materiale vrne avtorju skupaj z negativno odločitvijo glavnega urednika. 
INSTRUCTIONS FORAUTHORS 
l. Types of Articles 
a) SCIENTIFIC ARTICLES are comprehensive descriptions of original research and include a 
theoretical survey of the topic, a detailed presentation of results with discussion and conclusion, and a 
bibliography according to the IMRAD outline (lntroduction, Methods, Results, and Discussion). The 
length of an article including tables, graphs, and illustrations may not exceed fifteen (15) pages; lines 
must be double-spaced. Scientific articles shall be subject to peer review by two experts in the field. 
b) REVIEW ARTICLES will be published in the journal after consultation between the editorial 
board and the author. Review articles may be longerthan fifteen (15) pages. 
c) BRIEFNOTES are original articles from various biological fields (systematics, biochemistry, 
genetics, microbiology, ecology, etc.) that do not include a detailed theoretical discussion. Their aim isto 
acquaint readers with preliminary or partial results of research. They should not be longer than five (5) 
pages. Brief note articles shall be subject to peer review by one expert in the field. 
d) CONGRESS NEWS acquaints readers with the content and conclusions of important congresses 
and seminars at home and abroad. 
e) ASSOCIATION NEWS reports on the work of Slovene biology associations. 
2. Originality of Articles 
Manuscripts submitted for publication in Acta Biologica Slovenica should not contain previously 
published material and should not be under consideration for publication elsewhere. 
3.Language 
Articles and notes should be submitted in English, or as an exception in Slovene if the topic is very 
Iocal. Asa rule, congress and association news will appear in Slovene. 
4. Titles of Articles 
Titles (in Slovene and English) must be short, informative, and understandable. The title should be 
followed by the name and full address of the author (and if possible, fax number and e-mail address). 
S.Abstract 
The abstract must gi ve concise information about the objective, the methods used, the results obtai-
ned, and the conclusions. The suitable length for scientific articles is approximately 250 words, and for 
briefnote articles, 100 words. 
6.Keywords 
There should be no more than ten (10) keywords; they must reflect the field ofresearch covered in 
the article. Authors must add keywords in English to articles written in Slovene. 
7. Introduction 
The introduction must refer only to topics presented in the article or brief note. 
8. Illustrations and Tables 
Articles should not contain more than ten ( 1 O) illustrations (graphs, dendrograms, pictures, photos 
etc.) and tables, and their positions in the article should be clearly indicated. Ali illustrative material 
should be provided as physical originals (photographs or illustrations). Tables with their Iegends should 
be submitted on separate pages (only horizontal lines should be used in tables). Titles oftables should 
appear above the tables, and titles of photographs and illustrations below. Titles of tables and illustrations 
and their legends should be in both Slovene and English. Tables and illustrations should be cited shortly 
in the text (Tab. 1 or Tabs. 1-2, Fig. 1 or Figs. 1-2; Tab. 1 and Sl. 1). 
9. Conclusions 
Articles shall end with a summary of the main findings which may be written in point form. 
10.Summary 
Articles written in Slovene must contain a more extensive English summary. The reverse also 
applies. 
11. Literature 
References shall be cited in the text. If a reference work by one author is cited, we write ALLAN ( 1995) 
or (ALLAN 1995); if a work by two authors is cited, (TRINAJSTIC & FRANJJC 1994); if a work by three or 
more authors is cited, (PuLLIN & al. 1995); and if the reference appears in severa! works, (HoNsJG-
ERLENBURG & al. 1992, W ARD 1994a, ALLAN 1995, PuLLIN & al. 1995). If severa! works by the same 
author published in the same year are cited, the individual works are indicated with the added letters a, b, 
c, etc.: (WARD l 994a,b). If direct quotations are used, the page numbers should be included: TOMAN 
( 1992: 5) or (TOMAN 1992: 5-6). 
The bibliography shall be arranged in alphabetical order beginning with the surname of the first 
author followed by the year of publication, the title of the article, the internati ona! abbreviation for the 
joumal (periodical), the vol ume (in bold print), the number in parenthesis, and the pages. Examples: 
H0Ns10-ERLENBURG W., K. KRAINER, P. M1LDNER & C. WIESER 1992: Zur Flora und Fauna des Weber-
sees. Carinthia II 182/102 (!): 159-173. 
TRINAJSTIC I. & J. FRANJic 1994: Ass. Salicetum elaeagno-daphnoides (BR.-BL. et VOLK, 1940) M. 
MOOR 1958 (Salicion elaeagni) in the Vegetation in Croatia. Nat. Croat. 3 (2): 253-256. 
WARDJ. V. 1994a: Ecology of Alpine Streams. FreshwaterBiology 32 (!): 10-15. 
W ARD J. V. 1994b: Ecology of Prealpine Streams. Freshwater Biology 32 (2): 10-15. 
Books, chapters from books, reports, and congress anthologies use the following forms: 
ALLAN J. D. 1995: Stream Ecology. Structure and Function ofRunning Waters, 1 st ed. Chapman & Hall, 
London, 388 pp. 
PuLLIN A. S., I. F. G. McLEAN & M. R. WEBB 1995: Ecology and Conservation of Lycaena dispar: 
British and European Perspectives. In: PULLIN A. S. (ed.): Ecology and Conservation of Butterflies, 
1 st ed. Chapman & Hall, London, pp. 150-164. 
TOMAN M. J. 1992: Mikrobiološke značilnosti bioloških čistilnih naprav. Zbornik referatov s posveto-
vanja DZVS, Gozd Martuljek, pp. 17. 
12. Format and Form of Articles 
Articles should be written with Wordfor Windows or WordPetfect using "Times New Roman CE 
12" font with double spacing, align left and margins of 3 cm on A4 pages. Paragraphs should be 
separated with an empty line. The title and chapters should be written bold in font size 14. Ali scientific 
names must be properly italicized. Used nomenclature source should be cited. Tables and illustrations 
shall accompany the texts separately. The original manuscript, two copies, and a copy ona 3.5" computer 
diskette must be given to the editor-in-chief. Ali articles must be proofread for professional and language 
errors before submission. 
13. Peer Review 
Ali Scientific Articles shall be subject to peer review by two experts in the field ( one Slovene and one 
foreign) and BriefNote articles by one Slovene expert in the field. Authors may nominate a foreign 
reviewer in an accompanying !etter. Reviewed articles accepted for publication shall be corrected by the 
author. Authors shall receive fifty (50) free copies of the journal upon publication. In the event an article 
is rejected, the original material shall be retumed to the author together with the negative detennination of