Internat. Rev. Hydrobiol.

92 2007 1 61–83

DOI: 10.1002/iroh.200510921

ANA PREVIšIĆ, MLADEN KEROVEC and MLADEN KUČINIĆ*

Department of Zoology, Faculty of Science, University of Zagreb, Rooseveltov trg 6,

10000 Zagreb, Croatia; e-mail: kucinic@zg.biol.pmf.hr

Emergence and Composition of Trichoptera from Karst Habitats,

Plitvice Lakes Region, Croatia
key words: caddisflies, emergence patterns, diversity, trophic structure, sex ratio

Abstract
Adult Trichoptera were observed at the Plitvice Lakes, Croatia during two years (2000 and 2001)
using pyramid-type emergence traps. A total of 1350 individuals and 37 species were collected.
Throughout the study, the emergence patterns showed only minor variations regarding differences in
particular habitats. During 2000 most species emerged in June, and in 2001 in May. The complete emer-
gence period is shown for all collected species and studied in detail for abundant species. Excluding
Allogamus uncatus, all abundant species emerged in summer, with most exhibiting a long emergence
period. Rhyacophila fasciata had the longest emergence period in both years of the study and also
showed winter activity. Trichopteran diversity and equitability were calculated and trophic relationships
of the collected species were identified for each sampling site in order to obtain better insight into tri-
chopteran community structure. Results indicated that the trichopteran community could be grouped
according to travertine barriers and stream habitat types.

1. Introduction
To date, there have been various studies focusing on trichopteran communities in karst
habitats, but mostly based on larval stages (e.g. BONADA et al., 2005; HABDIJA, 1988; HAB-
DIJA et al., 2002). Besides, data of adult Trichoptera ecology and faunistics are commonly
obtained by sampling with light and Malaise traps (e.g. SMITH et al., 2002; SVENSSON, 1974;
WARINGER, 1991; 2003). Thus emergence patterns and the length of the emergence periods
are still insufficiently investigated for many species inhabiting karst areas.
The emergence method has been used for various studies all over the world. At first, the
method was used for quantitative production studies (ILLIES, 1971; MALICKY, 1976), but
more recently it is mostly used for faunistic, phenological and biodiversity studies of run-
ning waters (FREITAG, 2004; FÜREDER et al., 2005; PETERSEN et al., 1999; WARINGER, 1996)
due to its advantages over some other methods, e.g. benthos sampling (MALICKY, 2002).
Hence, the emergence method was applied to investigate trichopteran community composi-
tion and emergence patterns in different karstic habitats.
The Balkan Peninsula is known to be an important evolutionary centre for a number of
trichopteran genera (e.g. Drusus, Rhyacophila, Tinodes, Potamophylax) resulting in a high
species endemism (KUMANSKI and MALICKY, 1999). The Plitvice Lakes National Park was
chosen for this study because of the presence of different habitat types typical of the karst
systems (springs, streams, lakes and travertine barriers). Moreover, some endemic species
from the mentioned genera inhabit the Plitvice Lakes area (KUČINIĆ and MALICKY, 2002;
MARINKOVIĆ-GOSPODNETIĆ, 1971) and were thus encompassed in the study.

* Corresponding author

© 2007 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1434-2944/07/102-0061

62 A. PREVIŠIĆ et al.

Figure 1. Location of emergence trap sampling sites in the Plitvice Lakes National Park, Croatia.
Abbreviations of the sampling sites: BR – spring of Bijela rijeka, CR – spring of Crna rijeka, PLJ –
Crnarijeka, lower reach, LA – Labudovac travertine barrier, KM – Kozjak-Milanovac travertine barrier.

The objectives of the study were (1) to define emergence patterns of species inhabiting
karst but also to compare these with data from different habitats and regions (2) to investi-
gate the composition, diversity and similarity of trichopteran fauna at different karst habi-
tats, (3) to determine the trophic structure of trichopteran communities at investigated habi-
tats and (4) sex ratios of abundant species.

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 Trichoptera from Karst Habitats 63

2. Study Area
The study was carried out in Plitvice Lakes National Park, located in the mountainous
region of Croatia, in the Lika region. Sixteen lakes, divided by numerous travertine barri-
ers, form the barrage system of the Plitvice Lakes. These lakes are characterised by specif-
ic hydrobiological properties such as low organic matter concentration, super saturation by
calcium salts, pH > 8.0 (SRDOČ et al., 1985) and the presence of algae and bacteria in some
mosses at the barriers (STILINOVIĆ and BOŽIČEVIĆ, 1998) that are responsible for the biody-
namic ecosystem. The process of travertine formation is responsible for the vertical growth
of the barriers, changing their geomorphology, but also for the occurrence of new and the
disappearance of old barriers and waterfalls (STILINOVIĆ and BOŽIČEVIĆ, 1998).
The rivers Crna rijeka and Bijela rijeka are the main water suppliers of the lakes (Fig. 1).
All study sites are at altitudes ranging between ca. 500 and 720 m. Physical and chemical
parameters of the water at the sampling sites are presented in Table 1.
Due to its geographical location, the Lika region is influenced by the characteristics of
both temperate and continental climates according to the KÖPPENS climate classification
(MAKJANIĆ, 1958). Total annual precipitation in the last five years (January 2000 to Decem-
ber 2004) was in the range of 1148 – 1835 mm. Most of the area is covered by forest con-
sisting of Fagus sylvatica L., Abies alba MILL. and Picea abies (L.) KARSTEN.

Table 1. Characteristics of the five sampling sites in the Plitvice Lakes National Park. BR –
spring of Bijela rijeka, CR – spring of Crna rijeka, PLJ – Crna rijeka, lower reach, LA –
Labudovac travertine barrier, KM – Kozjak-Milanovac travertine barrier.

Site BR CR PLJ LA KM

Elevation (m) 720 677 670 630 546

Latitude N 44°50′005″ N 44°50′143″ N 44°50′10,5″ N 44°52′17,5″ N 44°53′39,9″
Longitude E 15°33′436″ E 15°36′282″ E 15°36′30,7″ E 15°35′59,8″ E 15°36′32,8″
Substrate stones, gravel, stones, gravel stones, gravel, travertine travertine
composition sand sand
Surrounding meadow, beech beech and beech forest beech and fir willow tree,
vegetation and spruce spruce forest forest bush
forest
Riparian canopy open closed closed closed closed
Water
temperature (°C) 7.40/7.70 7.70/8.00 6.20*/8.70 2.60/20.40** 1.60/22.10**
min/max
Oxygen (mg/l) 10.30/14.18 9.80/17.65 10.66/16.95 9.33/13.09** 9.32/13.19**
min/max
Oxygen saturation 92.90/128.90 87.00/166.50 98.1/150.00 83.75/124.37** 82.59/118.43**
(%) min/max
Conductivity 466/516 408/437 420/437 400 368
(µS/cm)
min/max
pH min/max 7.30/7.54 7.30/7.54 7.76/7.78 8.03/8.30** 8.10/8.50**
Carbonate 12.99/15.08 11.59/13.97 10.8/13.98 11.06/12.57** 10.34/11.54**
hardness (d °H)
min/max

(Physico-chemical data unpublished except *MATONIĆKIN and PAVLETIĆ, 1967 and **SRDOĆ et al., 1985)

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64 A. PREVIŠIĆ et al.

The study was carried out at four sampling sites in 2000: two headwater springs of the
Bijela rijeka (BR) and Crna rijeka (CR) rivers; lower reach of the river Crna rijeka (PLJ);
and a travertine barrier Labudovac (LA). An additional travertine barrier, Kozjak-Milanovac
(KM) was sampled in 2001 (Fig. 1).

3. Methods
Four pyramid-type emergence traps were installed during 2000 at each site. Unfortunately, in Feb-
ruary 2001, one trap at the headwater springs of Crna rijeka and Bijela rijeka, respectively, were
destroyed, there was therefore one trap less at each sampling site in the second year of the study. The
traps were placed at various positions across the stream section to cover different microhabitats present
at each site. Each trap was a 50 cm high, four-sided pyramid with a base of 45 × 45 cm, and fastened
to the stream bed in a way that allowed the free movement of larvae in and out of the sampling area.
The side frames of each trap were covered by net 1 mm mesh-size, and at the top of each trap was a col-
lecting container filled with preservative (2% formaldehyde with detergent). The containers were checked
monthly from January 2000 until December 2001, and the samples pooled for each site.
Identification of the collected material was based on MALICKY (1983; 2004) and the systematic
review on BOTOSANEANU and MALICKY (1978). As it was not possible to identify Glossosoma and
Hydropsyche females to species level, they were only included in data analysis examining the number
of emerging species, diversity and similarity of trichopteran fauna if no males were caught. In the analy-
sis of sex ratios, the data of the above mentioned genera were pooled and analysed at the genus level.
Although one female belonging to the family Rhyacophilidae could not be identified to the species level
it was nevertheless included in the complete data analysis, because it could not be associated with any
of the known species caught. Emergence patterns were analysed in detail for species constituting more
than 3% (11 species) of the total catch in at least one year of the study, solely at the sampling sites
where they accounted for at least 7% of the catch in at least one year of the study.
The determination of the trophic relationships of the collected species was based on GRAF et al.
(1995) and WIGGINS (1996). Drusus croaticus MARINKOVIĆ -GOSPODNETIĆ , 1971 was classified as
scraper according to HABDIJA et al. (2002) and personal observation. The abundance of feeding types
for each sampling site was expressed in percentages of total number of individuals.
In order to find out more about the faunistic diversity of Trichoptera, the number of individuals, the
number of species, the Shannon diversity index (SHANNON, 1948), the Simpson’s diversity index (SIMP-
SON, 1949) and the equitability (PIELOU, 1969) were calculated for each site. Cluster analysis using pres-
ence/absence data with the Bray-Curtis coefficient and the group average (UPGMA) clustering method
were used to detect faunistic similarity of Trichoptera between sampling sites. For this analysis, sam-
ples were combined to give one pooled sample for each year of the study. The analysis was preformed
with PC-ORD Version 4 for Windows.

4. Results
4.1. Composition and Diversity of Trichopteran Fauna

A total of 1350 individuals and 37 species were collected comprising 779 individuals
belonging to 34 species, and 571 individuals belonging to 32 species during 2000 and the
2001, respectively. A complete list of the collected species at each sampling site during both
years is presented in Table 2.
Species were generally not equally abundant in the two years of the study. In 2000 43.5%
of the total catch consisted of two species, Synagapetus krawanyi ULMER, 1938 and D. croat-
icus (25.5% and 18%, respectively). In 2001 these two species accounted for 33.6% of the
total catch, of which only D. croaticus accounted for 28.2% (Table 2). Six species, Rhya-
cophila sp., Chaetopteryx gonospina MARINKOVIĆ, 1966, Micropterna lateralis (STEPHENS,
1834), Potamophylax latipennis (CURTIS, 1834), Halesus tessellatus (RAMBUR, 1842) and
Oecetis testacea (CURTIS, 1834), were represented by single individuals (Table 2).

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 Trichoptera from Karst Habitats 65

Table 2. Trichoptera species caught by emergence traps at five sites in the Plitvice Lakes
National Park during 2000 and 2001. Glossosoma and Hydropsyche females could not be
identified to species level with certainty and therefore shown as Glossosoma sp. and
Hydropsyche sp.
Species Sampling sites Total % of total
emergence

BR CR PLJ LA KM

2000 2001 2000 2001 2000 2001 2000 2001 2001 2000 2001 2000 2001

RHYACOPHILIDAE
Rhyacophila aurata 1 4 1 1 5 0.1 0.9
BRAUER, 1857
Rh. d. plitvicensis 13 26 3 13 29 1.7 5.1
KUČINIĆ and MALICKY,
2002
Rhyacophila fasciata 10 18 13 10 8 2 31 30 4.0 5.3
HAGEN, 1859
Rhyacophila tristis 4 8 2 12 2 1.5 0.4
PICTET, 1834
Rhyacophila sp. 1 0 1 0.0 0.2

GLOSSOSOMATIDAE
Glossosoma bifidum 1 1 1 1 0.1 0.2
MCLACHLAN, 1879
Glossosoma discopho- 10 5 1 5 11 0.6 1.9
rum KLAPALEK, 1902
Glossosoma sp.

3 42 7 1 3 2 13 45 1.7 7.9
Synagapetus krawanyi 1 3 91 22 107 6 199 31 25.5 5.4
ULMER, 1938

PHILOPOTAMIDAE
Wormaldia subnigra 1 16 16 3 17 19 2.2 3.3
MCLACHLAN, 1865

HYDROPSYCHIDAE
Hydropsyche instabilis 13 3 1 13 4 1.7 0.7
(CURTIS, 1834)
Hydropsyche saxonica 43 17 3 43 20 5.5 3.5
MCLACHLAN, 1884
Hydropsyche sp.

73 37 7 73 44 9.4 7.7

POLYCENTROPODIDAE
Neureclipsis bimaculata 1 3 5 1 8 0.1 1.4
(LINNAEUS, 1758)
Plectrocnemia brevis 1 3 1 3 0.1 0.5
MCLACHLAN, 1871
Polycentropus flavoma- 2 5 2 5 0.3 0.9
culatus (PICTET, 1834)

PSYCHOMYIDAE
Lype reducta 6 4 2 6 6 0.8 1.1
(HAGEN, 1868)
Tinodes dives 2 6 49 4 18 1 1 2 70 13 9.0 2.3
(PICTET, 1834)
Tinodes unicolor 3 1 3 1 0.4 0.2
(PICTET, 1834)

LIMNEPHILIDAE
Drusus croaticus 129 156 10 5 1 140 161 18.0 28.2
MARINKOVIĆ-GOS-
PODNETIĆ, 1971

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66 A. PREVIŠIĆ et al.

Table 2. (continued)

Species Sampling sites Total % of total

emergence

BR CR PLJ LA KM

2000 2001 2000 2001 2000 2001 2000 2001 2001 2000 2001 2000 2001

Limnephilus lunatus 1 11 22 11 23 1.4 4.0

CURTIS, 1834
Limnephilus rhombicus 4 4 0 0.5 0.0
(LINNAEUS, 1758)
Grammotaulius nigro- 1 1 2 0.3 0.0
punctatus (RETZIUS,
1783)
Potamophylax latipen- 1 0 1 0.0 0.2
nis (CURTIS, 1834)
Potamophylax nigricor- 5 1 1 5 0.1 0.9
nis (PICTET, 1834)
Potamophylax pallidus
(KLAPALEK, 1899) 1 1 1 14 3 16 4 2.1 0.7
Halesus digitatus 6 2 1 8 1 1.0 0.2
(SCHRANK, 1781)
Halesus tessellatus 1 1 0 0.1 0.0
(RAMBUR, 1842)
Micropterna lateralis 1 0 1 0.0 0.2
(STEPHENS, 1834)
Allogamus uncatus 1 16 42 31 8 1 51 48 6.55 8.41
(BRAUER, 1857)
Chaetopteryx fusca 5 1 5 1 0.6 0.2
BRAUER, 1857
Chaetopteryx gono- 1 1 0 0.1 0.0
spina MARINKOVIĆ,
1966

GOERIDAE
Lithax niger (HAGEN, 1 1 6 2 6 0.3 1.1
1859)
Silo pallipes 1 2 1 3 1 0.4 0.2
(FABRICIUS, 1781)

LEPIDOSTOMATIDAE
Lepidostoma hirtum 11 20 11 20 1.4 3.5
(FABRICIUS, 1775)

LEPTOCERIDAE
Mystacides azurea 7 13 6 7 19 0.9 3.3
(LINNAEUS, 1761)
Oecetis testacea 1 1 0 0.1 0.0
(CURTIS, 1834)

SERICOSTOMATIDAE
Notidobia ciliaris 5 1 5 1 0.6 0.2
(LINNAEUS, 1761)
Sericostoma flavicorne 2 2 1 2 6 1 0.8 0.2
SCHNEIDER, 1845

Total 148 251 224 88 169 20 238 180 32 779 571

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 Trichoptera from Karst Habitats 67

Table 3. Diversity of Trichoptera caught by emergence traps at five sites; shown as num-
ber of individuals (N), number of species (S), equitability (PIELOU, 1969) (E), Simpson’s
diversity index (SIMPSON, 1949) (D) and Shannon diversity index (SHANNON, 1948) (H’) per
site per year of the study. ND = no data. For the abbreviations of the localities see legend
of Figure 1.

Site No. species Equitability Shannon Simpson’s

diversity index diversity index

Year 2000 2001 2000 2001 2000 2001 2000 2001

BR 8 6 0.251 0.522 0.522 0.934 0.2142 0.4263

CR 13 9 0.622 0.813 1.594 1.786 0.7294 0.7811
PLJ 14 12 0.533 0.892 1.408 2.216 0.5645 0.8457
LA 21 17 0.832 0.838 2.533 2.373 0.8862 0.8842
KM ND 9 ND 0.918 ND 2.016 ND 0.8480

Species richness was highest at the Labudovac barrier and lowest at the Bijela rijeka
spring in both years of the study (Table 3). In 2000 the lowest number of individuals were
collected at Bijela rijeka and the highest at Labudovac (Table 3). In contrast, the highest
number of individuals collected in 2001 were at Bijela rijeka, and the lowest at the lower
reach of Crna rijeka (Table 3).
The highest number of Trichoptera species was collected at Labudovac and the lowest at
Bijela rijeka during both years (Table 3). Both Shannon’s and Simpson’s diversity index
were highest at Labudovac during both years, whereas equitability was highest at Labudovac
in 2000 and Kozjak-Milanovac in 2001 (Table 3).
Cluster analysis shows that trichopteran community similarities were highest between
sites in Crna rijeka and between travertine barriers, in 2000 and 2001, respectively (Fig. 2).
Tinodes dives (PICTET, 1834), Potamophylax pallidus (KLAPALEK, 1899) and Sericostoma
flavicorne SCHNEIDER, 1845 were the only species recorded at all habitat types (springs,
lower reach and travertine barriers), whereas there were 12 species recorded in two differ-
ent habitat types (Table 2). Furthermore, there were three species collected solely at the
springs, and four species solely at the lower reach of the river Crna rijeka. Fifteen species
were only ever caught at the travertine barriers (Table 2).

4.2. Emergence Patterns

The complete period of emergence is presented for all collected species in Table 4. For
about half of the species collected in the two years emergence started 1 to 3 months earlier
in 2001 and the other half emerged either at the same time in both years or 1 month earlier
in 2000 (Table 4). The maximum number of emerging species were collected in June and
May, in 2000 and 2001, respectively (Table 4).
Regarding the number of collected individuals and recorded species at a particular sam-
pling site, considerable regularity in emergence patterns in the two years of the study was
found (Fig. 3). Moreover, there was a coinciding trend in the number of individuals collect-
ed and the number of species recorded in the two years (Fig. 3). At all sites emergence peaks
were recorded in the summer and autumn, mainly in June and September or October, respec-
tively (Fig. 3). The Kozjak-Milanovac barrier showed the highest variation, though emer-
gence patterns were generally in accordance with those observed at other sampling sites
(Fig. 3b, d).
All abundant species emerged during summer except Allogamus uncatus (BRAUER, 1857)
which was caught only in autumn (Fig. 4i, Table 4). Short emergence periods of 3 months

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68 A. PREVIŠIĆ et al.

2000
Distance (Objective Function)
5E-02 2,7E-01 4,8E-01 7E-01 9,1E-01

Information Remaining (%)

100 75 50 25 0

BR

CR

PLJ

LA

a)

2001
Distance (Objective Function)
2,7E-01 5,6E-01 8,4E-01 1,1E+00 1,4E+00

Information Remaining (%)

100 75 50 25 0

BR

CR

PLJ

LA

KM

b)
Figure 2. Cluster analysis based on the Bray-Curtis coefficient showing similarities of Trichoptera
community at sampling sites in 2000 (a) and 2001 (b).

were recorded for Lepidostoma hirtum (FABRICIUS, 1775), A. uncatus and Mystacides azurea
(LINNAEUS, 1761), whereas the remaining species had emergence periods of 4 months or
more in at least one year of the study (Fig. 4, Table 4).
In the two years of the study, most of the analysed species exhibited unimodal emergence
patterns regardless of the numbers caught although the emergence patterns were found to be
erratic for Limnephilus lunatus CURTIS, 1834, M. azurea and Rhyacophila fasciata HAGEN,
1859 (Fig. 4). R. fasciata had the longest emergence period (8 – 9 months) and was active all
year (although there was only one individual caught each month in the period from April to
July 2001) (Fig. 4b, Table 4).

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 Trichoptera from Karst Habitats 69

Table 4. The period of emergence shown for all species collected during the two years of
the study; ● represents 2000 and ▲ 2001. Glossosoma and Hydropsyche females could not
be identified to species level with certainty and therefore are shown as Glossosoma sp.
and Hydropsyche sp.
Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Rhyacophila aurata ●▲ ▲
Rh. dorsalis plitvicensis* ▲ ▲ ▲ ●▲ ●▲ ●▲ ●▲ ●
Rhyacophila fasciata* ▲ ●▲ ▲ ●▲ ●▲ ●▲ ●▲ ●▲ ●▲ ●
Rhyacophila tristis ▲ ● ● ●
Rhyacophila sp.
▲
Glossosoma bifidum ● ▲
Glossosoma discophorum ▲ ●▲ ●▲ ●▲ ▲
Glossosoma sp.

●▲ ●▲ ●▲ ●▲ ● ●
Synagapetus krawanyi* ● ●▲ ●▲ ●▲ ▲
Wormaldia subnigra* ●▲ ●▲ ●▲ ▲
Hydropsyche instabilis ●▲ ▲
Hydropsyche saxonica* ▲ ●▲ ●▲ ●▲ ●▲ ●▲
Hydropsyche sp.

▲ ●▲ ●▲ ●▲ ●▲ ●▲
Neureclipsis bimaculata ●▲ ▲ ▲
Plectrocnemia brevis ▲ ●
Polyce. flavomaculatus ▲ ▲ ● ▲
Lype reducta ▲ ●▲ ●▲ ● ▲
Tinodes dives* ▲ ●▲ ●▲ ●▲ ●▲ ▲
Tinodes unicolor ●▲
Drusus croaticus* ●▲ ●▲ ●▲ ●▲ ●▲ ●▲ ●
Limnephilus lunatus* ▲ ●▲ ▲ ● ▲
Limnephilus rhombicus ● ●
G. nigropunctatus ● ●
Potamophylax. latipennis ▲
Potamoph. nigricornis ▲ ●▲
Potamoph. pallidus ● ●▲
Halesus digitatus ●▲
Halesus tessellatus ●
Micropterna lateralis ▲
Allogamus uncatus * ●▲ ●▲ ●
Chaetopteryx fusca ● ●▲
Chaetopteryx gonospina ●
Lithax niger ▲ ● ●
Silo pallipes ▲ ●
Lepidostoma hirtum* ●▲ ●▲ ●▲
Mystacides azurea* ▲ ●▲ ●▲ ●
Oecetis testacea ●
Notidobia ciliaris ●▲
Sericostoma flavicorne ▲● ●

Number of 2000 0 0 1 0 11 19 16 13 14 6 4 0
species 2001 1 0 1 6 17 16 14 10 16 8 0 0

*abundant species (≥ 3% of the total catch in at least one year of the study)

For most species, emergence peaks were recorded in the summer, mainly in June (Table 4).
Two emergence peaks were recorded for D. croaticus; one in June (both years) and a later
peak in September 2000 and August 2001 (Fig. 4g).

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70 A. PREVIŠIĆ et al.

12

10
Number of species

8
BR
CR
6
PLJ
LA
4

0 a)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
2000

10

7
Number of species

BR
6
CR
5 PLJ
LA
4
KM
3

0 b)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
2001

Figure 3a, b

4.3. Trophic Structure of Trichopteran Community

Considering the trophic structure of the trichopteran community (presented in Fig. 5) at
sampled habitats, marked differences were observed.
At the Bijela rijeka spring the community was dominated by scrapers, represented by
D. croaticus, accounting for a very large portion of the collection in both years (87% and

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 Trichoptera from Karst Habitats 71

100

90

80

70
Number of individuals

60 BR
CR
50
PLJ
40 LA

30

20

10

0 c)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
2000

80

70

60
Number of individuals

50 BR
CR
40 PLJ
LA
30 KM

20

10

0 d)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
2001

Figure 3. Number of species recorded in a) 2000 and b) 2001 and number of individuals caught in
c) 2000 and d) 2001 at all sampling sites.

62%, respectively). Scrapers– collectors gatherers were well represented in 2001 (22%) by
Glossosomatidae. Predators accounted for about 7% in both years and were represented by
R. fasciata.
In 2000, the community at the Crna rijeka spring was dominated by scrapers– collectors
gatherers (45%) represented mainly by S. krawanyi. Scrapers– collectors filterers– collectors
gatherers represented by T. dives and shredders– scrapers– predators, represented by Alloga-

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72 A. PREVIŠIĆ et al.

Rhyacophila dorsalis plitvicensis

12

10
Number of individuals

2000 LA
6
2001 LA

0 a)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Date

Rhyacophila fasciata

5
Number of individuals

4 2000 BR
2001 BR
2000 CR
3
2001 CR
2000 PLJ
2 2001 PLJ

0 b)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Date

Figure 4a, b

mus uncatus and Potamophylax species, also accounted for a relatively large proportion of
the collection (22% and 19%, respectively). In 2001, the community was dominated by
shredders–scrapers– predators (41%), followed by scrapers– collectors gatherers (34%).
Predators represented by Rhyacophila species composed 8% and 11% in 2000 and 2001,
respectively.
Similarly, the trichopteran community in the lower reach of the river Crna rijeka (PLJ)
was dominated by scrapers– collectors gatherers represented mainly by S. krawanyi, account-

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 Trichoptera from Karst Habitats 73

Synagapetus krawanyi

90

80

70
Number of individuals

60
2000 CR
50
2000 PLJ
40 2001 CR
2001 PLJ
30

20

10

0 c)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Date

Wormaldia subnigra

10

7
Number of individuals

6
2000 LA
5
2001 LA
4

1
d)
0
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Date

Figure 4c, d

ing for a large proportion of the collection in both years (69% and 50%, respectively). Shred-
ders–predators–scrapers, represented by seven Limnephilidae species, accounted for 9%
and 25%, respectively. Predators, represented by R. fasciata accounted for 5% and 10%,
respectively. Scrapers– collectors filterers– collectors gatherers represented by T. dives
accounted for 11% in 2000.
Compared to previous sites, the travertine barriers showed a completely different trophic
structure of the trichopteran community. At LA the community was dominated by collectors
filterers–predators–scrapers represented by two Hydropsyche species (54% and 32%, in

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74 A. PREVIŠIĆ et al.

Hydropsyche saxonica

25

20
Number of individuals

15
2000 LA
2001 LA
10

0 e)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Date

Tinodes dives

20

18

16
Number of individuals

14

12 2000 CR
2000 PLJ
10
2001 CR
8 2001 PLJ

0 f)
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Date

Figure 4e, f

2000 and 2001, respectively). Predators, represented by O. testacea and two Rhyacophila
species, accounted for 10% and 18%, followed by shredders– predators– scrapers,
represented by four Limnephilidae species, composing 8% and 13% of collected individu-
als.
At KM the community was also dominated by collectors filterers– predators– scrapers,
represented by two Hydropsyche species (34%), followed by collectors gatherers– shred-

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 Trichoptera from Karst Habitats 75

Drusus croaticus

60

50
Number of individuals

40

2000 BR
30
2001 BR

20

10

g)
0
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Date

Limnephilus lunatus

14

12

10
Number of individuals

8
2000 LA
2001 LA
6

0 h)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Date

Figure 4g, h

ders–scrapers–predators, represented by M. azurea (19%). Predators– collectors filterers

and predators, represented by N. bimaculata and three Rhyacophila species, accounted for
about 16% each. Collectors filterers represented by W. subnigra accounted for 9% and scrap-
ers–xylophagous represented by L. reducta for 6%. The remaining feeding types present at
the investigated sampling sites were generally poorly represented.

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76 A. PREVIŠIĆ et al.

Allogamus uncatus

40

35

30
Number of individuals

25

2000 CR
20
2001 CR

15

10

0
i)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Date

Lepidostoma hirtum

16

14

12
Number of individuals

10

2000 LA
8
2001 LA

j)
0
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Date

Figure 4i, j

4.4. Sex Ratio

Due to the relatively few individuals caught, the sex ratio is only shown (as the percent-
age of males) for species that constituted more than 3% of the catch in at least one year of
study. The sex ratios are presented in Table 5. In 2000 there were five species and Hydropsy-
che spp. trapped in sufficient numbers to test the significance of deviations from 1 :1, and
in 2001, there were nine species, Glossosoma and Hydropsyche spp. Rhyacophila fasciata
was the only species with a sex ratio not significantly different from 1 :1 in either year of

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 Trichoptera from Karst Habitats 77

Mystacides azurea

6
Number of individuals

4
2000 LA
2001 LA
3

0 k)
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Date

Figure 4. Number of the individuals of the eleven most abundant species recorded at selected sam-
pling sites in the two years of the study.

predators
100%
predators - collector filterers

collectors gatherers - shredders -

scrapers - predators
80%
collectors filterers

collectors filterers - predators - s crapers

60% scrapers

scrapers - x ylophagus

scrapers - x ylophagous - shredders

40%
scrapers - c ollectors gatherers

scrapers - c ollector filterers - collector

gatherers
20%
shredders

shredders - predators

0% shredders - predators - scrapers

BR BR CR CR PLJ PLJ LA LA KM
shredders - s crapers - predators
2000 2001 2000 2001 2000 2001 2000 2001 2001

Figure 5. The trophic structure of trichopteran community at sampled habitats in the Plitvice Lakes
National Park.

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78 A. PREVIŠIĆ et al.

Table 5. Sex ratios (as percentage of males) presented for abundant species (≥3% of the
total catch in at least one year of the study). Glossosoma and Hydropsyche species were
pooled as females could not be reliable discriminated. χ2 tests were used to asses the sig-
nificance of deviations from 1 :1 (*P < 0.05; **P < 0.01; ***P < 0.001).

Species 2000 2001

% of males

Rhyacophila dorsalis plitvicensis 13.79***

Rhyacophila fasciata 51.61 56.67
Glossosoma sp. 21.05***
Synagapetus krawanyi 32.16*** 41.94
Wormaldia subnigra 47.37
Hydropsyche sp. 43.41 35.29*
Tinodes dives 50.00
Drusus croaticus 40.71* 32.91***
Limnephilus lunatus 47.83
Allogamus uncatus 47.06 33.33*
Lepidostoma hirtum 45.00
Mystacides azurea 47.37

the study. In species with ratios significantly different from 1 :1, females predominated (by
about 1.5:1 to 6 :1).

5. Discussion
5.1. Emergence Patterns

Differences in the length of recorded emergence periods and flight periods reported by
many authors (HICKIN, 1967; SVENSSON, 1972; WARINGER, 1991; 2003) may be the conse-
quence of the sampling method, but also of number of factors influencing the life cycle of
caddisflies, such as variability in habitat conditions affecting the larval stage, mainly food
resources and water temperature (e.g. TRACHET, 1967; NOVAK and SEHNAL, 1962; SWEENEY,
1984; WAGNER, 2002) or differences in the adult longevity (MORSE, 2003).
The life-cycles of most species in northern temperate climates involve emergence peaks
in the summer (HICKIN, 1967; ILLIES, 1971) what was also recorded in the present study.
Furthermore, there are only five species of the tribes Chaetopterygini and Stenophylacini,
which are typical autumnal species (genera Allogamus, Chaetopteryx, Halesus) in the whole
species inventory (according to CRICHTON in HICKIN, 1967; MALICKY, 1973).
Detailed analysis of abundant species in particular habitats suggests that there is specific
variability in emergence patterns caused by differences in the life cycles of each species.
Observed emergence patterns were in accordance with previously published data (e.g.
HICKIN, 1967; SVENSSON, 1972; WARINGER, 1996) for some species but for the others some
discrepancies were recorded. For S. krawanyi the emergence period was more extended than
that reported by WARINGER (1996). The emergence pattern for W. subnigra was found to be
unimodal over the two years and its emergence period was in accordance with the flight
period reported by HICKIN (1967). On the other hand, for L. hirtum and M. azurea, the flight
periods reported by HICKIN (1967) and WARINGER (1991; 2003) were more extended than
the emergence periods recorded in our study. The flight period for R. dorsalis dorsalis

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 Trichoptera from Karst Habitats 79

reported by HICKIN (1967) and for R. dorsalis persimilis by WARINGER (1991) is consistent
with the emergence period recorded for R. dorsalis plitvicensis in our study. Similarly, the
recorded emergence peak was in accordance with light-trapping results of KUČINIĆ (2002) at
the same site. According to SVENSSON (1972), R. fasciata, emerges and mates over a long
time, and consequently has a very long flight period without distinct peaks including win-
ter-flight activity. Winter activity of R. fasciata was previously recorded in Germany (one
individual caught during December 2000) by P. NEU (personal communication). Adults of
R. fasciata have also been collected during December 2004, January and February 2005 at
some sites in the Mediterranean parts of Croatia (unpublished data). These findings are in
accordance with OTTO’s (1981) hypothesis that caddisflies with predatory larvae, due to con-
tinuous availability of food resources during the year, exhibit nearly no synchronisation and
long emergence period. For L. lunatus the emergence period was recorded to be short, aes-
tival and variable in the two years, whereas the flight period reported by HICKIN (1967) and
WARINGER (1996) was more prolonged, during summer and autumn. The erratic emergence
of L. lunatus and M. azurea may be explicable if one assumes that their main populations
live in the lake above, hence they are species typical of stagnant waters (MALICKY, 1973;
GRAF et al., 1995). The total numbers are quite low and they may have intruded into the
traps, since intruding species were sometimes observed in the emergence traps (e.g. ILLIES,
1971; MALICKY, 2002). Furthermore, L. lunatus is a species that aestivates, with the main
part of the population emerging in spring/early summer and returning to the aquatic habitats
for ovoposition in autumn (NOVAK and SEHNAL, 1962). To explain the latter emergence pat-
terns, the emerging rates at the lakes should also be investigated.

5.2. Composition and Diversity of Trichopteran Fauna

Differences in numbers of individuals and species collected during the two years are prob-
ably partly attributable to one fewer trap installed at each site in 2001, and also to the col-
lecting method properties and variability of environmental conditions. Some long-term stud-
ies of the effects of environmental variables on the macroinvertebrate community (e.g.
WOOD et al., 2000; WAGNER and SCHMIDT, 2004) showed that discharge variations had the
highest influence on community structure. In the present study, lower numbers of individu-
als collected at each site were in accordance with the higher discharge recorded. MALICKY
(2002) recorded more variable results for the pyramid type emergence traps than for the larg-
er traps and suggested that this was due to their small bottom surfaces. Thus, a longer col-
lecting period and an enlarged bottom surface covered by emergence traps (e.g. increased
number of traps) probably could improve our results.
Our results suggest that some species show habitat specific distributions. For instance,
Tinodes unicolor (PICTET, 1834), recorded solely at the travertine barriers, is a well known
specialist of travertine habitats (EDINGTON and HILDREW, 1995). Furthermore, Glossosoma-
tidae were restricted to spring and stream habitats, corresponding to observations of
light trapping (KUČINIĆ, 2002), whereas Hydropsychidae were only recorded from travertine
barriers, which does not correspond with data provided by light traps (KUČINIĆ, 2002), sug-
gesting that the passive pyramid trapping method is very important factor in obtaining infor-
mation on the breeding habitat of collected species rather than attractant traps such as light
traps.
Considering the diversity of aquatic insects in the longitudinal profile of a stream, it gen-
erally increases downstream (WILLIAMS and FELTMATE, 1992). In the current study, both
diversity and equitability of Trichoptera were generally lower at springs than at downstream
sites (an exception was CR in 2000, Table 3). All values were by far the lowest at the Bijela
rijeka spring due to the high proportion of D. croaticus in the catch. Habitat characteristics
at this site are obviously highly suitable for the latter species, regarding its distribution

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80 A. PREVIŠIĆ et al.

(springs and upper reaches), feeding (scraper; HABDIJA et al. 2002; personal observation) and
ovoposition behaviour (personal observation). High community diversity and species rich-
ness usually reflect high habitat diversity (e.g. WARINGER, 1996; WIBERG-LARSEN et al.,
2000), thus the highest diversity and species richness at Labudovac could be the conse-
quence of the variety of microhabitats present at the barrier and the availability of various
food resources (HABDIJA et al., 2004; MILIšA et al., 2006).
Cluster analysis results grouped the trichopteran fauna into two main groups; at travertine
barriers and stream habitats. The similarity of trichopteran communities at sites at the Crna
rijeka is greater than at the two springs, and most probably due to significant differences in
morphological and hydrogeological features (MATONIČKIN and PAVLETIĆ, 1967; SRDOČ et al.,
1985) e.g. water velocity, substrate composition, shading by riparian vegetation etc. The
influence of various environmental variables (e.g. substrate composition, stability and het-
erogeneity, organic matter, water temperature, riparian vegetation etc.) on aquatic insect
communities was outlined by many authors, for instance MINSHALL (1984) and WILLIAMS
and FELTMATE (1992) and references therein, SMITH et al. (2003) etc.

5.3. Trophic Structure of Trichopteran Community

Differences in the trophic structure of trichopteran community at the investigated habitats

reflect differences in the availability of various food resources (VANNOTE et al., 1980).
According to the River Continuum Concept (VANNOTE et al., 1980) the upper reaches of
headwaters are under strong influence from the surrounding vegetation, and the high input
of coarse particulate organic matter (CPOM) in the shaded headwaters results in a benthic
community dominated by shredders. The Crna rijeka spring is shaded by surrounding veg-
etation, with markedly higher input of CPOM (such as leaf litter) than the unshaded Bijela
rijeka spring, hence the differences in trophic structure of trichopteran communities. Simi-
lar habitat characteristics at the spring (CR) and lower reach of the river Crna rijeka (PLJ)
most likely resulted in similarities in composition and trophic structure of trichopteran com-
munities at these two sampling sites.
Sites at travertine barriers showed a completely different trophic structure compared to
the stream habitats (Fig. 5), which is attributable to the character of these sites, for they are
lake outlets and therefore contain more seston (e.g. more biomass of plankton) (ŠPOLJAR,
2003; HABDIJA et al., 2004). Hence these communities are dominated by collectors such as
Hydropsyche species.

5.4. Sex Ratio

Uneven sex ratios were recorded in studies based on the use of emergence traps
(WARINGER, 1996; MALICKY, 2002) and some other methods as well (e.g. light traps and
malaise traps) (WARINGER, 2003; SMITH et al., 2002; SVENSSON, 1974). It is thought that
some factors may influence the sex ratios, like emergence and oviposition behaviour of
females and emergence trap design (MALICKY, 2002) were determined. Hence, in the cur-
rent study only few species were trapped in sufficient numbers to allow statistical testing
and comparison of the sex ratios.

6. Conclusions
In the present study, specific differences in trichopteran community structure in different
karstic habitat types were recorded (springs, lower reach and travertine barriers). Emergence

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 Trichoptera from Karst Habitats 81

patterns and emergence periods and also sex ratios were analysed for the abundant species.
For some species emergence patterns were in accordance with previously published data but
for the others some discrepancies, attributable to various factors, were recorded. The current
study represents the first study which investigates the life cycle of D. croaticus and gives
the exact larval and pupal habitat preferences for some insufficiently known south-eastern
European species. Results of similarity and trophic structure analysis indicated that the tri-
chopteran community could be grouped according to travertine barriers and stream habitat
types. Such distribution and community structure of Trichoptera reflect specific habitat con-
ditions and life history traits. Altogether, our results suggest that emergence studies are
important for obtaining data on the emergence patterns and faunistics of caddisflies from
differing habitats, but also to provide information on species life history traits, as well as tri-
chopteran species diversity and ecology of the investigated area.

7. Acknowledgements
The authors would like to thank the many colleagues from Croatian Natural History Museum and
Department of Zoology who helped with this study, assisting with emergence trap construction and field
work. We are very grateful to the Plitvice Lakes National Park authorities, especially Dr. NATALIJA
PAVLUS, for their support and permission in field sampling. We thank the National Meteorological and
Hydrological Service for providing us data about precipitation and discharge. We also thank two anony-
mous reviewers who provided very useful comments and suggestions that improved the manuscript.
This research was supported by Croatian Ministry of Science, Education and Sports as a part of the Pro-
ject number 0119-121.

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Manuscript received November 3rd, 2005; revised October 6th, 2006; accepted November 7th, 2006

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