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Cent. Eur. J. Biol. • 6(1) • 2011 • 118–129 DOI: 10.2478/s11535-010-0098-x

Central European Journal of Biology

Ecology of (Diptera) in a wetland habitat and their potential role as bioindicators

Research Article Ivan Gelbič*, Jiří Olejníček

Biological Centre of Czech Academy of Sciences, Institute of Entomology, CZ 370 05 České Budějovice,

Received 28 April 2010; Accepted 06 September 2010

Abstract: Ecologicalinvestigationsoflong-legged(Dolichopodidae)werecarriedoutinwetmeadowwetlandsnearČeskéBudějovice, Czech Republic. Sampling was performed during the adult flies’ seasonal activity (March-October) in 2002, 2003 and 2004 using yellow pan traps, Malaise traps, emergence traps, and by sweeping. Altogether 5,697 specimens of 78  of Dolichopodidae were collected, identified and analysed. The study examined community structure, species abundance, and diversity(Shannon-Weaver’sindex-H’;Sheldon’sequitabilityindex-E). cilipes,C. gramineus and ungulatus were the most abundant species in all three years. Species richness and diversity seem strongly affected by soil moisture.

Keywords: Long-legged Flies • Ecology • Conservation • Bioindication ©VersitaSp.zo.o.

usually do not too far from their breeding places, 1. Introduction which is convenient for their use as bioindicators. The Dolichopodid flies represent a good model for the study aims of this paper are (i) to extend our knowledge on of bioindication because they meet all necessary criteria the community structure of dolichopodid fauna and for this role [1]. Pollet has indicated four such criteria: (ii) to explore possibilities for the use of these as 1) easy determination of species, 2) a taxonomic bio-indicators. group comprised of a sufficient number of species, 3) satisfactory knowledge about ecology/biology of the species, and 4) species should reveal specific ecological 2. Experimental Procedures requirements [2]. North Central , roughly delimited by 5–20° E 2.1 Study area and 45–55° N, has been the subject of numerous The study was carried out during 2002–2004, inclusive, at studies that have provided basic information about Švábův Hrádek near České Budějovice, Czech Republic habitat evaluation for Dolichopodidae [3-22]. Most of (Figure 1) (48°58’26.84” N, 14°25’55.93” E). The sampling these studies relate to , Czech Republic and site represents a mosaic of different landscape elements, . Other studies, however, have also reported on with a wetland of less than 0.1 km2 with wet meadows, two the dipteran fauna (: Limoniidae; Tipulidae; small permanent pools of about 25 and 300 m2 (the latter ; : ; ; one is actually a chain of several connected smaller pools Dolichopodidae) in other parts of the region [23-29] and with alders, birches and oaks on their banks), and several are important in this respect. old fruit trees adjacent to the wet meadows. The sampling Larvae and adults of the dolichopodid flies are site is immediately surrounded by intensively farmed land predators [29,30] that can be found in all wet biotopes. bordering coniferous forest (mixed spruce and pine). The Larvae are mostly aquatic or semi-aquatic. Adults wetland is situated in the lower part of a former open-cast

* E-mail: [email protected] 118 I. Gelbič, J. Olejníček

pit for extracting clay used in bricks manufacturing, but has 2. Emergence traps (ET): Six ETs (metal construction not been in production for about the past 50 years. The of conical shape covered by black waterproof serge) upper part of the pit (outside the study location) was used were placed at various locations in the wet meadows as dump for communal refuse. Our monitoring was initiated and at the pool edges, two of them standing just in 10 years after this activity ceased. The dump was thinly shallow water (littoral) near the pool bank [10]. The covered with soil and overgrown by self-seeding plants. traps covered 0.25 m2 and the collected insects were In 2003, the entire dump was “reclaimed” by elimination trapped in a bottle containing 75% ethyl alcohol which of herbage and adding a 50 cm layer of clay followed by was emptied every two weeks. a layer of soil of the same thickness and then sown with 3. Sweeping: For an additional sampling method, we grass. This decreased the level of pool water by about used sweep nets once each month (50 sweeps linearly 50 cm and caused partial desiccation of the wet meadows for 20 min of continuously sweeping). in 2003 and 2004. Climatic data for all dates 2002 to 2004 4. Malaise traps (MT): Two MTs, each controlled were obtained from the hydro-meteorological station every two weeks, were used and the material was in České Budějovice, which is part of a meteorological collected in 75% alcohol. monitoring network. 2.3 Material processing, identification and 2.2 Sampling statistical analysis During 2002 to 2004, from March to October, Preservation and mounting: individual specimens were Dolichopodid flies were collected in the above site using dried prior to identification. Flies preserved in alcohol the following sampling strategy: were dried by: 1) transferring from alcohol to an alcohol- 1. Yellow pan traps (YT): Thirty plastic rectangular formaldehyde mixture; 2) transferring to alcohol-diethyl dishes (size 10x7x4cm), each containing 1% of ether mixture; then 3) transferring to diethyl ether, the formaldehyde solution with a few drops of the detergent evaporation of which left the material dried. Dried flies Lena natur®, were distributed linearly (with 5 m distances were mounted on triangular paper labels. between two neighbouring dishes) at the investigated Several monographs were used for identification locality. YTs were in operation for one day per week and [34-47]. The relative abundance (%) of each species yields were transferred to and stored in 75% ethanol. was calculated separately for each year of the three-year According to Pollet and Grootaert [32] and Barták [33], study. The following scale of dominance was used: more yellow pan traps are the most effective collecting method than 10% of total number of Dolichopodidae caught per for the majority of species of the Dolichopodidae studied year was regarded as eudominant (ED), 5–10% although the traps are less suitable for the primarily dominant (D), 2–5% subdominant (SD), 1–2% recedent arboreal . (R) and less than 1% subrecedent (SR). The Shannon-

Figure 1. Picture of the studied locality (part of a wet meadow with pools).

119 Ecology of Dolichopodidae (Diptera) in a wetland habitat and their potential role as bioindicators

Weaver diversity index (H’) and equitability index On the other hand, analysis of the sex ratio shows the (E) were monitored for each year. A linear regression greatest disproportion in the genera Chrysotus and was fitted to a linear model to determine relationships Sybistroma. In species of the genera Chrysotus and between the size of the studied area and biodiversity. Sybistroma males predominated over females. Just the reverse was true in P. nobilitatus. Moreover, the degree of sexual disproportion rose with increasing level of 3. Results and Discussion dominancy (compare Figure 2 with Table 1). 3.1 General results and sex ratio 3.2 Community structure The total number of dolichopodid specimens collected Semi aquatic localities are typical for most species of during 2002–2004 was 5,697 (3,700, 1,659 and 338 the Dolichopodidae, and specimens of this family often in 2002, 2003 and 2004, respectively). The taxonomic represent the most frequent captures in samples from representation encompassed 18 genera and 78 species such biotopes. (57 species in 2002, 48 species in 2003 and 37 species Table 1 shows the total number of individuals of in 2004; see Table 1). Most of this material was obtained each species captured, their relative abundance (%), using the yellow water trap method. Only three species and the classification of dominancy. Only three species ( cinereus, apicalis, and – Chrysotus cilipes, Ch. gramineus and Dolichopus bicolorellum) were collected by a different method – the ungulatus – were consistently dominant or eudominant emergence traps. Yellow water traps were most effective in all three years, but none of these species was itself in catching males (Figure 2). This is consistent with the eudominant for all three years. Dolichopus latilimbatus, observations of Pollet and Grootaert [10] who noted that D. plumipes, Poecilobothrus nobilitatus and Poecilobothrus in-water traps located on the soil surface, the frequency chrysozygos were dominant or eudominant for one year of males is higher than in traps at 25 cm height. Figure 2 only. An unexpectedly large number of species for such presents the percentages for species that were at least a small locality was detected during the first year of subdominant in at least one of the monitored years. Sex monitoring, although there was a decrease in individual ratio is generally close to 1:1, and this was shown in and species abundance after drainage of the locality the majority of investigated species of Dolichopodidae. caused partial drying of the wetland.

89% 90 84% 90 83% ♂ 78% 80% ♂ ♂ ♂ ♂ 70 66% 67% 70 61% 63% ♂ 58% ♂ ♂ ♂ 52% ♂ 47% 50 45% ♂ 50 ♂ ♂ 39% ♂ 30 30

10 10

Ch. angulicornis Ch. cilipes Ch. cupreus Ch. gramineus Ch. neglectus festivus D. D. latilimbatus D. plumipes D. trivialis D. ungulatus H. chrysozygos N. nodicornis P. nobilitatus zetterstedti R. ♂ ♀ 1 2 3 4 5 6 7 8 9 10 11 12 13 14 -10 ♀ -10 ♀ ♀ 11% ♀ ♀ 17% 16% 22% 20% -30 ♀ ♀ -30 ♀ ♀ 34% ♀ 33% 39% 37% ♀ 42% -50 ♀ -50 ♀ 48% 55% ♀ 53% 61% -70 -70

Figure 2. Sex ratios of selected species from yellow water trap.

120 I. Gelbič, J. Olejníček

2002 2003 2004 2002–2004 Species No % CD No % CD No % CD No % CD Achalcus cinereus 1 0.03 SR 1 0.02 SR argentina 1 0.30 SR 1 0.02 SR 6 0.36 SR 6 0.11 SR 2 0.05 SR 3 0.18 SR 15 4.44 SD 20 0.35 SR 2 0.05 SR 15 SR 1 0.30 SR 18 0.32 SR 1 0.03 SR 4 0.24 SR 5 0.09 SR curvipes 1 0.24 SR 4 0.24 SR 5 1.48 R 18 0.32 SR Campsicnemus magius 1 0.06 SR 1 0.02 SR Campsicnemus marginatus 1 0.30 SR 1 0.02 SR Campsicnemus scambus 1 0.30 SR 1 0.02 SR Chrysotus cilipes 286 7.73 D 777 46.84 ED 68 20.12 ED 1131 19.85 ED Chrysotus cupreus 265 7.16 D 20 1.21 R 285 5.00 D Chrysotus femoratus 19 0.51 SR 3 0.18 SR 22 0.39 SR Chrysotus gramineus 1804 48.76 ED 169 10.19 ED 30 8.88 D 2003 35.16 ED Chrysotus laesus 1 0.06 SR 1 0.02 SR Chrysotus monochaetus 7 0.19 SR 7 0.12 SR 76 4.58 SD 11 3.25 SD 87 1.53 R Chrysotus pulchellus 22 0.59 SR 17 1.02 R 5 1.48 R 44 0.77 SR Chrysotus suavis 31 1.87 R 6 1.78 R 37 0.65 SR Dolichopus agilis 1 0.06 SR 1 0.02 SR Dolichopus arbustorum 12 0.32 SR 12 0.21 SR Dolichopus brevipennis 9 0.24 SR 7 0.42 SR 5 1.48 R 21 0.37 SR Dolichopus campestris 6 0.16 SR 1 0.06 SR 2 0.59 SR 9 0.16 SR Dolichopus cilifemoratus 1 0.03 SR 1 0.02 SR Dolichopus claviger 13 0.35 SR 5 0.30 SR 2 0.59 SR 20 0.35 SR Dolichopus clavipes 2 0.05 SR 2 0.04 SR Dolichopus cruralis 2 0.59 SR 2 0.04 SR Dolichopus festivus 91 2.46 SD 20 1.21 R 9 2.66 SD 120 2.11 SD Dolichopus flavipes 1 0.03 SR 1 0.02 SR Dolichopus griseipennis 5 0.30 SR 5 0.09 SR Dolichopus latilimbatus 13 0.35 SR 89 5.36 D 16 4.73 SD 118 2.07 SD Dolichopus longicornis 17 0.46 SR 3 0.18 SR 20 0.35 SR Dolichopus nigricornis 1 0.03 SR 3 0.18 SR 4 0.00 SR Dolichopus nitidus 3 0.08 SR 1 0.06 SR 4 0.07 SR Dolichopus nubilus 55 1.49 R 67 4.04 SD 3 0.89 SR 125 2.19 SD Dolichopus pennatus 1 0.03 SR 1 0.02 SR Dolichopus plumipes 76 2.05 SD 57 3.44 SD 21 6.21 D 154 2.70 SD Dolichopus popularis 2 0.05 SR 11 0.66 SR 13 0.23 SR Dolichopus signatus 2 0.05 SR 2 0.04 SR Dolichopus signifer 3 0.08 SR 3 0.05 SR Dolichopus simplex 7 0.19 SR 1 0.30 SR 8 0.14 SR Dolichopus subpennatus 1 0.03 SR 1 0.30 SR 2 0.04 SR Dolichopus trivialis 76 2.05 SD 1 0.06 SR 9 2.66 SD 86 1.51 R 486 13.14 ED 134 8.08 D 23 6.80 D 643 11.29 ED Dolichopus wahlbergi 1 0.03 SR 5 0.30 SR 6 1.78 R 12 0.21 SR Ethiromyia chalybea 3 0.08 SR 4 0.24 SR 7 0.12 SR aerosus 18 0.49 SR 6 0.36 SR 2 0.59 SR 26 0.46 SR Gymnopternus angustifrons 1 0.03 SR 1 0.02 SR Hercostomus nanus 1 0.06 SR 1 0.02 SR Hercostomus parvilamellatus 1 0.03 SR 1 0.02 SR Lamprochromus bifasciatus 1 0.06 SR 1 0.02 SR Medetera apicalis 1 0.03 SR 1 0.02 SR Medetera bisecta 6 1.78 R 6 0.11 SR Medetera mixta 1 0.03 SR 1 0.02 SR Medetera pallipes 2 0.59 SR 2 0.04 SR 3 0.08 SR 4 1.18 R 7 0.12 SR Nematoproctus distendens 1 0.06 SR 1 0.02 SR quadrifasciata 21 0.57 SR 14 0.84 SR 35 0.61 SR Poecilobothrus chrysozygos 268 7.24 D 23 1.39 R 1 0.30 SR 292 5.13 D Poecilobothrus nobilitatus 37 1.00 R 6 0.36 SR 53 15.68 ED 96 1.69 R albifrons 3 0.18 SR 3 0.05 SR Rhaphium appendiculatum 2 0.12 SR 2 0.04 SR Rhaphium auctum 2 0.05 SR 2 0.04 SR Rhaphium caliginosum 12 0.32 SR 36 2.17 SD 10 2.96 SD 58 1.02 R Rhaphium commune 1 0.03 SR 1 0.02 SR Rhaphium laticorne 1 0.30 SR 1 0.02 SR Rhaphium micans 6 0.16 SR 4 0.24 SR 10 0.18 SR Rhaphium nigribarbatum 1 0.06 SR 1 0.02 SR platypterus 4 0.11 SR 9 0.54 SR 2 0.59 SR 15 0.26 SR Sybistroma crinipes 1 0.06 SR 1 0.02 SR Sybistroma nodicornis 12 0.32 SR 6 0.36 SR 7 2.07 SD 25 0.44 SR pulicarius 5 0.14 SR 1 0.30 SR 6 0.11 SR Syntormon bicolorellum 1 0.03 SR 1 0.02 SR Syntormon pumilum 2 0.05 SR 2 0.04 SR Syntormon rufipes 1 0.03 SR 1 0.30 SR 2 0.04 SR Teuchophorus spinigerellus 2 0.05 SR 1 0.06 SR 4 1.18 R 7 0.12 SR tenellus 1 0.03 SR 3 0.18 SR 4 0.07 SR Total specimens 3700 1659 338 5697 Total species 57 48 37 77 H’ 2,96 3.17 4.12 3.47 E 0,51 0.57 0.79 0.55

Table 1. ListofspeciescollectedduringthemonitoredperiodatŠvábůvHrádek,includingnumberofindividuals(No),relativeabundance(%) andclassificationofdominance(CD)(eudominant–ED,dominant–D,subdominant–SD,recedent–R,subrecedent–SR).EDandD are accentuated by bold face.

121 Ecology of Dolichopodidae (Diptera) in a wetland habitat and their potential role as bioindicators

Chrysotus gramineus (48.76%, 10.19% and locality, changes of vegetation) and climate (changes 8.88%, respectively) and C. cilipes (7.73%, 46.84% of temperature and precipitation). The impact of and 20.12%, respectively) were the most abundant environmental pollution on the diversity of dolichopodid species. C. gramineus was found to be dominant in flies could not be ascertained, but analysis of water and 2002 (before amelioration) and C. cilipes in the second soil samples showed only slightly increased amounts of year. This shows that the drier environment might be mercury above standard level in one of the soil samples. preferred by the latter species. Even though the same Slightly increased amounts of mercury had also been species of Dolichopodiade were studied, our result observed before amelioration. has been contrasted with Pollet [48], who found that While 2002 was marked by much Dolichopodidae can be used for quality assessment of higher precipitation than in 2003 and 2004 reed marshes and grass lands in Belgium. That may (Figure 3). This fact, together with catastrophic be related to location differences between Czech and destruction due to amelioration, was possibly Belgium. responsible for the decrease in species richness Another dominant species during the first year of and specimen abundance. The decrease continued monitoring was C. cupreus. This species decreased until 2004. While species richness was highest in in the second year, coincident with amelioration, and the first year of monitoring and decreased in each disappeared completely in the third year, most likely as subsequent year, the index of diversity (H’) and a result of altered drainage. Dolichopus ungulatus was index of equitability (E) rose with decreasing the only large species that was dominant at all times richness (Table 1). The species spectrum was (see Table 1). All the above mentioned species are larger in comparison with results reported for other common throughout the Czech Republic [3,4,20,49,50]. localities (including larger and smaller study sites) Poecilobothrus nobilitatus is a frequent inhabitant of [3-7,10,13,15,16,24,32,48,51]. In spite of the partial small periodical pools west of the Bohemian-Moravian drying of the wetland and decreasing dolichopodid Uplands, but it is very rare east of these Uplands numbers, certain species were recorded only during (J. Olejníček, unpublished data). the third year of our investigations. Most species were collected throughout the entire Temperatures were warmer than the long-term study period (2002–2004). Only a few species occurred averages during our study (Figure 4). There was also in just one or two years (e.g. Argyra argyria, Argyra much higher fluctuation in the amount of precipitation. vestita, Campsicnemus magius, Medetera mixta, The very low humidity in February and March together Lamprochromus bifasciatus), and then mostly just at with extremely dry August in 2003 evidently reduced the one time per year (Table 2). The data in Tables 1 and number of species for this month. Only about half the 2 reveal important changes in the diversity of some number of species was recorded in this year compared species affected by amelioration (partial drying of the with the two years monitored earlier (Table 2).

450

400

350

300

250

200

Precipitation [mm] 150

100

50

0 I II III IV V VI VIIVIII IX X XI XII I II III IV V VI VIIVIII IX X XI XII I II III IV V VI VIIVIII IX X XI XII 2002 2003 2004

Figure 3. Monthlyprecipitation(thickline,blacksquares)andlong-termaverageprecipitation(thinline,opencircles)atthestudiedlocality.

122 I. Gelbič, J. Olejníček

Three species groups, with respect to ecological because of different species composition. For example, plasticity, could be distinguished. The first group is the Belgian wetlands contain the genera Achalcus and represented by species with a great ecological plasticity. Lamprochromus as the most abundant dolichopodids These species are usually present in most biotopes and but these are quite rare in the Czech Republic. often create dense populations (Chrysotus gramineus, Dolichopus plumipes, D. ungulatus). Species of this 3.3 The relationship between species richness group have a more limited usefulness. The second group and study area comprises highly specialised species--the occurrence There are several places in the Czech Republic where of which is limited by specific biotopes (like peat-bogs, dolichopodid fauna have been previously studied. salt marshes) or species that are endemic to limited Detailed studies were made using identical sampling territories (usually mountains or hot springs). These methods in Pálava Biospherical Reserve [49] and in the species are usually dominant in their typical biotopes Bílina and Duchcov reclaimed area at a former open- and absent outside of these localities. However, their cast brown-coal mine [50]. The Švábův Hrádek locality occasional occurrence in other localities can indicate used in the present study has a more limited range of that these species can be infrequent migrants or that biotopes compared with the other studied areas. Švábův their presence depends on the higher concentration of Hrádek represents only single types of biotopes – trees, salt in the soil,resulting from drought. A typical example lowland wet meadows and very small pools – while is the halophilous Campsicnemus magius, which is Pálava and Bílina or Duchcov contain large varieties of rare in central Europe [48] but which appeared at the lowland or gradient and submountain biotopes. These studied locality after the extremely dry August in 2003 include, for example, stream or river banks, flooded (Figure 3). forest and fishpond (or permanent pools) banks, as well The third species group includes the remaining as peat-bogs with mountain faunal biotopes. Combined, dolichopodid species, which occur in relatively large these biotopes encompass practically all types that can areas and exhibit rather large ecological plasticity, be found in central Europe. although they prefer one or only a limited number of The species abundance (77 species) on our different biotopes. These species form the majority relatively small locality is rather high. The larger study of the investigated species, and knowledge of their sites mentioned above revealed 152 and 128 species. biology, occurrence, and dominance rates can provide A regression line fitted to a plot of number of species useful information on the particular locality. These against geographical area is shown in Figure 5. species can be very abundant in favourable biotopes It suggests a positive linear relationship between these while being scare in other localities. It is sometimes variables. The relationship is obviously valid only for problematic to compare localities from different regions the mild-climate areas of similar geographical latitude.

20

15

10

5 Temperature [°C]

0

-5 I II III IV V VI VII VIII IX X XI XII I II III IV V VI VII VIII IX X XI XII I II III IV V VI VII VIII IX X XI XII 2002 2003 2004

Figure 4. Comparisonofmonthlytemperaturesinmonitoredyears(thickline,blacksquares)withlong-termaverage(thinline,opencircles) obtainedfromČeskéBudějovicemeteorologicalstation.

123 Ecology of Dolichopodidae (Diptera) in a wetland habitat and their potential role as bioindicators

350 CZ&SK

300 CZ

250 y = 18,51⋅ln(x) + 89,35 200 R2 = 0,95

150 Pa

No. of of No. species B&D 100 SH 50

0 0,01 0,1 1 10 100 1000 10000 1000001000000 Area [km2]

Figure 5. RelationbetweenmonitoredareaandsupposednumberofspeciesofDolichopodidae.SH,ŠvábůvHrádek–77species,0.10km2; B&D,BílinaandDuchcov–128species,48km2;Pa,Pálava–152species,83km2;CZ,CzechRepublic–305species,78,866km2; CZ &SK,CzechandSlovakrepublics–324species,127881km2.

In other climatic regions, the line might not only have a Acknowledgements different slope but could be of different form. The results indicate that, like many other insects, dolichopodid flies This paper was supported by Grant No. S6022201 of are suitable species as bioindicators of human influence the Academy of Sciences of the Czech Republic and by on ecosystems. For example, using dolichopodid flies grants Z60220518 and Z 50070508. We wish to thank as bioindicators of climatic changes would require more Prof. Dr. Rudolf Rozkošný, DrSc. and Dr. Karel Sláma, detailed information, including the study of weather and CSc. for reviewing a draft of this paper. biodiversity on a much larger territory.

124 I. Gelbič, J. Olejníček X X XX X X X XX X X X X X X X X XX X XXX 2004 XXX XXX XXX XXX XXX X XXXX XXXX X X X X X X X X X 2003 XXXX X XX XX X XX X XXXXX XX XXX XX XX XX XXX XXXX XXX X X X XXXXX XXXXX XXX XXXX XXXXX XX X X X X X X X X 2002 XX X XXX X X XX X XXX X XXX X X XX XX XXXX XX X XXX XXX XXXXX XXXX XX X XXX XXXXX XXX XXXXX XXX XXXX I II III IV V VI VII VIII IX X XI XII I II III IV V VI VII VIII IX X XI XII I II III IV V VI VII VIII IX X XI XII Occurrence of monitored species by individual years at the experimental area. Species Achalcus cinereus Argyra argyria Argyra diaphana Argyra leucocephala Argyra vestita Campsicnemus curvipes Campsicnemus magius Campsicnemus marginatus Campsicnemus scambus Chrysotus cupreus Chrysotus femoratus Chrysotus laesus Chrysotus monochaetus Chrysotus neglectus Chrysotus pulchellus Chrysotus suavis Dolichopus agilis Dolichopus arbustorum Dolichopus brevipennis Dolichopus campestris Dolichopus cilifemoratus Dolichopus claviger Dolichopus clavipes Dolichopus cruralis Dolichopus festivus Dolichopus flavipes Dolichopus griseipennis Dolichopus latilimbatus Dolichopus longicornis Dolichopus nigricornis Dolichopus nitidus Dolichopus nubilus Dolichopus pennatus Dolichopus plumipes Dolichopus popularis Dolichopus signatus Dolichopus signifer Dolichopus simplex Dolichopus subpennatus Dolichopus trivialis Dolichopus wahlbergi Ethiromyia chalybeus Gymnopternus aerosus Chrysotus cilipes Chrysotus gramineus Dolichopus ungulatus Gymnopternus angustifrons Table 2. Table

125 Ecology of Dolichopodidae (Diptera) in a wetland habitat and their potential role as bioindicators X X XXX XX X XX 2004 XX X X X XXX XXX X X X X X X X 2003 X XXX X XX X X X XX XX X XXXXX X X X X XX 2002 X X XX XXX XX XXX XX XXXX X XX XX X XX X I II III IV V VI VII VIII IX X XI XII I II III IV V VI VII VIII IX X XI XII I II III IV V VI VII VIII IX X XI XII Occurrence of monitored species by individual years at the experimental area. Table 2. Table Species Hercostomus chrysozygos Hercostomus nanus Hercostomus parvilamellatus Lamprochromus bifasciatus Medetera apicalis Medetera bisecta Medetera mixta Medetera pallipes Medetera truncorum Nematoproctus distendens Neurigona quadrifasciata Poecilobothrus nobilitatus Rhaphium albifrons Rhaphium appendiculatum Rhaphium auctum Rhaphium caliginosum Rhaphium commune Rhaphium laticorne Rhaphium micans Rhaphium nigribarbauma Sciapus platypterus Sybistroma crinipes Sybistroma nodicornis Sympycnus pulicarius Syntormon bicolorellum Syntormon pumilum Syntormon rufipes Teuchophorus spinigerellus Teuchophorus Xanthochlorus tenellus Continued

126 I. Gelbič, J. Olejníček

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