Cent. Eur. J. Biol. • 7(5) • 2012 • 917-926 DOI: 10.2478/s11535-012-0061-0

Central European Journal of Biology

Mosquitoes (Diptera, Culicidae) of the Morava River floodplain,

Research Article

Lucia Strelková*, Jozef Halgoš

Department of Ecology, Faculty of Natural Sciences Comenius University, 842 15 Bratislava 4, Slovakia

Received 07 February 2012; Accepted 15 May 2012

Abstract: Research on the Morava River floodplain  fauna was carried out from April to October in 2009 and 2010. Altogether, 5864 adults were collected and identified from 12 selected sites. The presence of 28 mosquito species belonging to6 genera were confirmed in the study area. The floodplain of the Morava River is known for its frequent inundationevery year. The floods are quite irregular and often followed by the mass production of mosquitoes. The most abundant species were Aedes vexans (41.4%), Ae. cinereus (7.5%), Ae. rossicus (16.7%), Ochlerotatus sticticus (20.5%), and pipiens (3.1%). Ae. vexans and Oc. sticticus are typical outbreak species for the Morava River floodplain. The years 2009 and 2010 differed in the number of floods, which influenced the mosquito faunal composition and abundance during theyear. Keywords: Mosquitoes • Morava River Basin • Seasonal changes • Floods • Floodplain ©VersitaSp.zo.o.

1. Introduction There are 50 mosquito species recorded in Slovakia. More attention started to be paid to the family Culicidae Spreading of diseases across Europe in the last several at the beginning of the 20th century due to the occurrence years has led to the necessity of mosquito monitoring. of malaria in Slovakia, which was eradicated in 1963 [2]. Tropical mosquito-borne diseases like dengue, Mosquitoes transmitted or still transmit four arboviruses: chikungunya, and West Nile fever have rapidly become Ťahyňa, Čalovo, Sindbis, and West Nile [3], in the territory a threat to public health in Europe that is proving difficult of Slovakia. was reported in humans from to confront. The presence and spread of these types several countries in Central and Eastern Europe and the of diseases depend on many different and interacting Mediterranean last year, and 2010 saw the highest number factors, such as the current distribution of mosquitoes of human cases reported, suggesting it has become a re- and their population densities, climate, wind patterns, emerging pathogen in Europe [4] (http://archive.defra.gov. proximity to bodies of water, land use, and vegetation. uk/foodfarm/farmanimal/diseases/monitoring/documents/ In addition, the frequency of intercontinental travel wnv-europe.pdf). Another mosquito-borne disease in and trade and global warming allow foreign species Slovakia is dirofilariosis. The first autochthonous case in to establish themselves in new environments, where Slovakia was recorded in dogs in 2005, and first human their natural enemies are not found. In the recent past, case occurred in 2007 in Western Slovakia [5], the area considerable fluctuation in air temperature has been where our research was carried out. At present, dirofilariosis typical for spring and summer. Precipitation has been is regarded as the fastest-spreading -borne irregularly spread during the year, in contrast with disease, and represents a high infection risk for both normal conditions featuring long dry periods alternating and humans [6]. with heavy rains, which caused localised or extensive The Morava River floodplain, with various types floods with mosquito calamities [1]. of stagnant waters, offers suitable conditions for

* E-mail: [email protected] 917 Mosquitoes (Diptera, Culicidae) of the Morava River floodplain, Slovakia

mosquito hatching. Variation in climatic conditions can Devín Castle near the town of Bratislava to the village of lead to noticeable changes taking place in freshwater Moravský Svätý Ján. The study sites represent different ecosystems, which are particularly evident in the types of stagnant waters (including plesiopotamal, species composition and ecology of the examined palaeopotamal, breeding sites in forests and on groups [7]. The key factors that influence meadows, and temporary ponds created and filled by composition, abundance, and ecology of mosquito rainfall or groundwater). Selected characteristics of the species are changes in the hydrological regime and 12 study sites are summarized in Table 1. water and air temperatures. The Morava River floodplain Material was collected at least once a month from is characterized by its frequent and often irregular floods April to October, depending on floods. Some of the every year. These floods create ideal breeding habitats study sites were not always accessible due to flooding. for floodwater species of mosquitoes of the genera Adults were collected by sweeping the vegetation with Aedes and Ochlerotatus. The years 2009 and 2010 entomological nets (diameter 25 cm) for 10 minutes at differed in the number and frequency of flood events. each breeding site. Collection began in the early morning Changes in climatic conditions lead to changes in the at site 1 and continued over all sites in one day. Water phenology of mosquitoes [7]. Earlier snow melt and temperature was also measured at breeding sites. The spring precipitation could result in a shift in the larval collected mosquitoes were killed by ether, transported development of the spring monovoltine species [7]. The to the laboratory, and stored in the freezer. Sex was mosquito production can be influenced by the extent determined using a stereomicroscope. All males were of flooding and previous precipitation, and the final dissected and external genitalia were mounted in mosquito assemblage depends on the local hydrological Canada balsam. Morphological identification keys were conditions [8]. used for species identification [10,11]. We used PAST software version 1.95 (http://folk.uio.no/ ohammer/past/) to evaluate the data for site similarity based 2. Experimental Procedures on the presence/absence of species (Jaccard index).

The Morava River floodplain is one of the last well- preserved wetlands in Central Europe [9]. Collections 3. Results of mosquitoes were made at 12 selected sites in the Morava River floodplain (Figure 1) from April to October Years 2009 and 2010 were very similar when considering in 2009 and 2010. The study area extends from the the mean monthly air temperature (Figure 2). The

Figure 1. Locations of collection sites in the Morava River floodplain, Slovakia (source: www.maps.google.com).

918 L. Strelková, J. Halgoš

Altitude (m Depth Shading Sediment Site Latitude (N) Longitude (E) Biotope Ambience above sea level) (m) (%) type 1. Moravský Sv. Ján - before 48°36’01,4’’ 16°56’39,9’’ 83.3 SP 0.4 W, M 20 earth embankment 2. Moravský Sv. Ján - behind 48°36’01,4’’ 16°56’39,9’’ 83.3 D 0.4 W 100 mud embankment 3. Lantov - tributary 48°34’10,5’’ 16°57’16,0’’ 150.6 Pal 1 M 40 mud

4. Lantov - meadow 48°34’10,5’’ 16°57’16,0’’ 150.6 Moor 0.7 M 0 mud

5. Malé Leváre a 48°33’15,2’’ 16°57’38,0’’ 140 D 0.5 W 100 mud

6. Malé Leváre b 48°30’50,6’’ 16°56’00,4’’ 112 D 0.4 W 100 mud

7. Vysoká pri Morave 48°19’14,8’’ 16°54’29,8’’ 137 Ple 0.5 S,M 75 mud

8. Devínske jazero - meadow 48°16’23,4’’ 16°55’46,1’’ 136.1 Ple 0.2 M 10 mud

9. Devínske jazero - forest 48°16’18,9’’ 16°55’52,0’’ 131 D 0.6 W 100 earth

10. Devínske jazero - tributary 48°15’42,2’’ 16°57’30,9’’ 123.6 Ple 0.5 M 10 mud 11. Devínska Nová Ves - 48°19’49,0’’ 16°97’40,4’’ 124 D 0.2 W 100 earth alluvial forest 12. Devín - near the castle 48°10’33,4’’ 16°58’40,1’’ 124 SD 0.8 W, M 50 mud

Table 1. Characteristics of the study sites. Abbreviations: D = depression filled by groundwater, SP = pool filled with water in spring only, Ple = plesiopotamal, Pal = palaeopotamal, W = wood, M = meadow, S = suburban area.

Figure 2. Mean monthly air temperature and precipitation in Moravský Svätý Ján (data from Slovak Hydrometeorological Institute in Bratislava). greatest differences occurred in April (14.4°C in 2009 lasted from May to September, with the highest values and 10.4°C in 2010) and September (16.5°C in 2009 in May and August. and 13.9°C in 2010). The other months differed by no A total of 5864 adult mosquitoes belonging to 28 more than 2°C. The years 2009 and 2010 differed in the species were collected (Table 2). A total of 3205 adults water level and precipitation values throughout the year were collected in 2009. The months with the highest (Figures 2 and 3). While the floods in 2009 occurred in abundance were July and August. The most abundant the spring and summer months only, the year 2010 was species were Ae. vexans, Oc. sticticus, Ae. rossicus, atypical with respect to the frequency and duration of and Ae. cinereus (Table 3), together representing over floods. The water level of the Morava River reached its 85% of all specimens collected. There was only one highest values in March, May and June, but Figure 3 peak of abundance of all species sampled in 2009, shows that raised water levels occurred many times although there were several generations of some throughout the year. In 2009, higher rates of precipitation species (Figure 4). A total of 2659 adult mosquitoes were lasted from February until March, from May until August, collected in 2010. The most abundant species were Ae. and in November. In 2010, high mean precipitation vexans, Oc. sticticus, Ae. rossicus, and Cx. pipiens

919 Mosquitoes (Diptera, Culicidae) of the Morava River floodplain, Slovakia

Figure 3. Daily water level values of the Morava River in Moravský Svätý Ján in 2009 and 2010 (data from the Slovak Hydrometeorological Institute, Bratislava).

1 2 3 4 5 6 7 8 9 10 11 12

An. claviger ○

An. hyrcanus ○ ● ○

An. maculipennis s.l. ● ● ● ● ●

An. plumbeus ●

Ae. cinereus ○ ● ○ ● ○ ○ ● ○ ● ○ ○ ● ○ ○ ● ○ ○ ●

Ae. geminus ●

Ae. rossicus ○ ● ○ ● ○ ○ ● ○ ● ○ ● ○ ● ○ ● ○ ● ○ ● ○ ●

Ae. vexans ○ ● ○ ● ○ ● ○ ● ○ ● ○ ● ○ ● ○ ● ○ ● ○ ● ○ ● ○ ●

Oc. annulipes ● ○ ● ○ ● ○ ○ ○ ○ ○

Oc. cantans ○ ● ○ ● ○ ● ○ ● ○ ○ ● ○ ○ ○ ○

Oc. caspius ● ● ●

Oc. cataphyla ● ● ○ ○ ○ ○ ○ ● ○ ●

Oc. communis ○ ● ○ ○ ○

Oc. excrucians ● ○ ● ○ ● ○ ● ○ ● ○ ●

Oc. flavescens ● ○ ○ ○ ○ ○ ○

Oc. genniculatus ○

Oc. intrudens ● ● ● ○ ● ● ● ●

Oc. leucomelas ○ ○ ○

Oc. punctor ○ ○

Oc. rusticus ● ●

Oc. sticticus ○ ● ○ ● ○ ○ ● ○ ● ○ ● ○ ○ ● ○ ○ ○ ● ○ ●

Cx. hortensis ● ● ○ ● ● ●

Cx. modestus ● ○ ● ○ ● ● ● ● ○ ● ○ ● ○ ● ● ●

Cx. pipiens ● ○ ● ○ ● ○ ● ○ ● ○ ● ○ ● ○ ● ● ● ○ ●

Cx. territans ○ ● ● ● ○ ● ○ ● ○ ● ○ ● ●

Cs. annulata ● ●

Cs. morsitans ● ○

Ur. unguiculata ●

Table 2. List of species collected at selected study sites (○ – 2009, ● – 2010).

920 L. Strelková, J. Halgoš

16/4 7/5 11/6 15/7 13/8 24/8 9/9 21/9 9/10

An. claviger 0.2

An. hyrcanus 1.5

Ae. cinereus 1.1 1.9 4.8 8.5 25.4 1.5 12.5 5.2 5.3

Ae. rossicus 6.9 52.3 21.9 14.9 18.9 16.3 18.8 8.6

Ae. vexans 3.4 5.7 45.7 27.4 4.3 44.2 33.3 19.8 31.6

Oc. annulipes 4.6 1.6 2.9 0.7 0.3 4.7 2.8

Oc. cantans 29.9 5.2 4.8 0.1 2.0 7.0 8.3

Oc. cataphyla 26.4 9.3 2.5 1.7 0.3

Oc. communis 8.5 1.4 0.8 0.4

Oc. excrucians 1.6 0.5 0.4 0.2

Oc. flavescens 2.7 0.8 0.4 0.5 4.2 5.3

Oc. genniculatus 1.2

Oc. intrudens 0.5 0.8 0.1

Oc. leucomelas 6.9 0.5 0.4

Oc. punctor 0.5

Oc. sticticus 12.6 9.3 15.8 43.5 7.2 14.0 18.8 12.9 1.5

Cx. hortensis 0.7 0.2 0.9

Cx. modestus 0.2 11.3 5.3

Cx. pipiens 0.8 0.7 3.7 2.3 2.8 12.7 26.3

Cx. territans 0.8 1.8 28.4 5.3

Cs. morsitans 0.9

100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0

Total number of individuals 87 193 392 1353 911 86 48 116 19

Table 3. Species composition of mosquito fauna with relative proportions (%) of samples on collection dates in 2009.

(Table 4). The first peak of abundance occurred in An. plumbeus, Ae. geminus, Oc. caspius, Oc. July, and another, larger peak occurred at the end of geniculatus, Oc. punctor, Oc. rusticus, Cs. annulata, August after another long period of high precipitation Cs. morsitans, and Ur. unguiculata. Four species of the (Figure 5). The composition of the mosquito fauna at genus Culex were recorded: Cx. pipiens, Cx. modestus, these two abundance peaks was quite similar (12 to Cx. territans, and Cx. hortensis. Culex species occurred 10 species). The greatest contrast between the two from June to October during both years of research. The studied years was in the abundance of Oc. intrudens most abundant species was Cx. pipiens, followed by Cx. (1 specimen in 2009 to 31 specimens at 8 different sites territans, Cx. modestus, and Cx. hortensis in 2009, and in 2010). Cx. modestus, Cx. territans, and Cx. hortensis in 2010. In 2009, the first adults of early spring mosquitoes Cx. modestus was the fifth most abundant species in (Oc. communis, Oc. cataphyla, Oc. leucomelas) and 2010, with its highest numbers occurring in July. late spring mosquitoes (Oc. annulipes, Oc. cantans, The 12 selected sites were distributed into 4 Oc. sticticus, Ae. cinereus, Ae. rossicus and Ae. groups in the value 0.56 by cluster analysis based on vexans) appeared in mid-April. The number of species the Jaccard index (Figure 6). Two groups consisted of and their abundances rose in May. Most of the early single sites. Site 4 (Lantov - meadow) was separated spring species occurred until July, with Oc. leucomelas because of the presence of species that were not persisting until mid-August. In 2010, only adults of two recorded at other sites (An. plumbeus, Culiseta species, Oc. cataphyla and Oc. intrudens, occurred in annulata, and Uranotaenia unguiculata). Site 3 (Lantov April. The number of species rose to 10 in May. - tributary) was separated because of the presence Species with total relative frequency equal to of An. hyrcanus and An. macculipenis s.l., and the or less than 0.1% were An. claviger, An. hyrcanus, absence of Cx. hortensis. Third group was made of

921 Mosquitoes (Diptera, Culicidae) of the Morava River floodplain, Slovakia

Figure 4. Number of mosquitoes collected per date in the Morava River Basin in 2009.

Figure 5. Number of mosquitoes collected per date in the Morava River Basin in 2010.

Figure 6. Cluster analysis of the similarity of sites based on presence/absence of species (complete linkage, Jaccard index of similarity).

922 L. Strelková, J. Halgoš

16/4 18/5 8/6 23/6 13/7 4/8 25/8 20/9 13/10

An. hyrcanus 1.2

An. maculipennis s.l. 0.4 0.6 1.4 3.6

An. plumbeus 2.5

Ae. cinereus 1.4 1.9 1.9 0.4 1.7 1.8 1.4 1.2

Ae. geminus 0.1

Ae. rossicus 1.,6 2.5 9.3 1.2 14.4 18.4 25.2 1.2

Ae. vexans 1.,6 48.8 34.3 53.8 62.8 65.3 51.6 27.6

Oc. annulipes 1.2 0.5 0.7

Oc. cantans 4.5 0.7 0.5 1.7 0.1

Oc. caspius 0.4 0.1

Oc. cataphyla 5.6 6.8

Oc. communis 1.4 1.2

Oc. excrucians 4.5 0.6 0.7 0.5

Oc. flavescens 0.2

Oc. intrudens 94.4 18.9 0.6

Oc. rusticus 1.4 0.4

Oc. sticticus 27.3 42.8 5.2 15.6 1.3 9.5 1.8 1.2

Cx. hortensis 1.3 0.4 6.1

Cx. modestus 0.4 12.9 2.1 0.3 1.6 2.5

Cx. pipiens 2.5 4.1 8.5 1.5 7.8 21.4

Cx. territans 0.6 1.7 7.4 2.3 9.6 15.4

Cs. annulata 2.5

Cs. morsitans 0.4

Ur. unguiculata 7.1

100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0

Total number of individuals 18 74 166 277 653 298 793 282 98

Table 4. Species composition of mosquito fauna with relative proportions (%) of samples on collection dates in 2010. sites 9, 10, and 12, which are all located in meadows. Slovakia. Michalková [14] found 22 species in the They are filled with water for short periods only, and the floodplain of the Morava River. Our researched area mean water temperature is higher than at the sites in neighbors with the South Moravia region in the Czech the following group. The fourth group was consisted of Republic, where many mosquito researches were carried sites 1, 2, 5, 6, 7, 8, and 11, which are all in or in close out. Olejníček et al. [15] found 19 species during research proximity to forests. The rare species Oc. caspius was in 1997–2002 in the Morava and Dyje river basins. Rettich only recorded at meadow sites (sites 3, 8, and 10). et al. [16] recorded larvae of 17 mosquito species during Another rare species, An. hyrcanus, was only recorded floods in 2005 and 2006 in south and central Moravia and at undisturbed sites (sites 3, 4, and 6). central Bohemia. In total, 39 species of mosquitoes have been confirmed in Moravia [17]. We did not confirm the presence of An. atroparvus, 4. Discussion Cq. richiardii, Oc. pulchritarsis, Oc. refiki, and Cx. theileri in the Morava floodplain. Cx. theileri was The first extensive research on mosquitoes in western recorded for the first time only recently in the territory Slovakia was carried out 40 years ago. Labuda [12] of Slovakia by Halgoš and Petrus [18], where it was recorded 27 species in lowland Záhorská nížina. Later, found in Devínske jazero, an area where three of our Jalili [13] collected 29 species of mosquitoes in Western sites are situated.

923 Mosquitoes (Diptera, Culicidae) of the Morava River floodplain, Slovakia

Another Mediterranean species recently reported so the water level was very high during most of the in Slovakia is An. hyrcanus [19], and as a potential months. These conditions led to massive amounts of malaria vector it should not be ignored [10]. This mosquitoes in the summer months after the water level study found a few additional specimens in October of fell. Due to favorable hydrologic conditions for mosquito 2009 and 2010. The only other record of this species breeding until autumn in 2010, Culex species composed in Slovakia is by Škultétyová Bolgárová [20] from the a large part of the mosquito outbreaks. Labuda [12] floodplain of the Ipeľ River. In the same month, October, also observed that the four abovementioned floodwater we also found the thermophilic species Ur. unguiculata. species are typical and predominant in the Morava River It was first recorded from Slovakia over 20 years ago floodplain, especially in areas very close to the main [21]. We found one male specimen of Ae. geminus in channel. He also found that in cases of dry summers August 2010 at site 9 (Devinske jazero – forest). The with no floods in the River Morava floodplain, there were first record of this species was reported by Labuda [22], no outbreaks of these species. Michalková [14] obtained who collected a few specimens in April and May of 1973 similar results when the floodwater species Ae. vexans, from four sites in the Borská nížina Lowland. No other Oc. sticticus, Ae. rossicus, and Ae. cinereus dominated specimens have been recorded until this study, which the mosquito populations in the Morava River floodplain. we have identified with certainty through their diagnostic Ae. vexans, Oc. sticticus, Ae. cinereus, and Cx. pipiens hypopygial characteristics [10]. The species with the also dominated the mosquito populations of the Morava lowest relative frequencies found in this study are all floodplain during floods in 1997 [25]. rare in Western Slovakia. Cx. modestus is an important West Nile virus vector. Hydrological conditions varied between the years of Labuda [12], Jalili [26], and Michalková [14] have our study. We observed the first adults of snow melting previously found this species only very sporadically in mosquitoes in the middle of April in both studied years. the Morava River floodplain. We collected this species Because of high precipitation in February and March in both years of our research, and it was even the fifth and relatively high temperatures in April in 2009, we most abundant species in 2010. Though previously found higher numbers of spring species in 2009 than rare in Bohemia [27], Votýpka et al. [28] confirmed in 2010. In 2010, the precipitation was low from the that thermophilic Cx. modestus appeared to be a more beginning of the year until April, and the mean monthly appropriate emerging vector for West Nile virus than the temperature in April was 4°C lower than in 2009. Due abundant species Cx. pipiens in the , to these conditions, only two species occurred in April as a consequence of its recent spread within the Czech of 2010. Minář et al. [7] also observed emergence of Republic and its willingness to feed on both avian and the first adults of the early spring species in the second mammalian hosts [28]. week of April, which was caused by favorable conditions Oc. caspius was an important species in the cluster for hatching in 2005 (earlier snow melt and early spring analysis for site similarity. It was collected solely at sites precipitation), while the year with weaker and late spring in meadows. This species is West Nile and Tahyna floods led to the emergence at the beginning of May. virus vector. Tahyna virus was originally isolated from Michalková [14] also recorded the first occurrence of Ae. vexans and Oc. caspius in Slovakia [29]. Labuda adults of early spring mosquitoes in May during years [12] also collected this species only sporadically, finding 2001-2003 in the Morava River floodplain. only a few specimens of larvae and adults, and it has Ae. vexans and Oc. sticticus are the predominant not been recorded in Morava River Basin since until this species in frequently inundated areas, such as study, although there have been several occurrences floodplains of rivers and lakes with fluctuating water level of this species in some areas in Western Slovakia. [10,23,24]. In Central Europe, they are associated with Oc. caspius has not been detected in the Morava and the hatching of the larvae of Ae. rossicus and Ae. cinereus Dyje basins in the Czech Republic in the last decade [10]. There were only two periods of flooding in 2009. [16], though it was previously found in high numbers in Only the second was followed by mosquito outbreak, flooded meadows there [30]. because the first was too early in the spring, so the air temperature was too low for the floodwater species (Ae. vexans, Oc. sticticus, Ae. rossicus, and Ae. cinereus). Acknowledgements The opposite hydrologic conditions were observed in 2010, which featured extensive and long lasting floods, This research was supported by Grant 1/0176/12.

924 L. Strelková, J. Halgoš

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