93 Lauterbornia 62: 93-119, D-86424 Dinkelscherben, 2008-05-15
Black flies of the River Danube (Diptera, Simuliidae) Ladislav Jedlička and Gunther Seitz With 2 tables Keywords: Simuliidae, Diptera, Insecta, Danube, zonation, taxonomy, nomenclature, faunistics Schlagwörter: Simuliidae, Diptera, Insecta, Donau, Zonierung, Taxonomie, Nomenklatur, Faunistik 31 nominal black fly species have been recorded in the River Danube, including the source rivers. The most comprehensive data are from the Slovak-Hungarian section (21 species before damming), the German stretches (19 agreed taxa), the Pannonian Plain section (10 species in Vojvodina), and the Serbian-Romanian section (7 species before damming). 10 species have been recorded from the Austrian section. Very little data are available from Bulgaria, Romania (including the Ukrainian part of the delta) and the Hungarian Danube Bend. The usual zonation of black fly species assemblages along the longitudinal gradient of the river is interrupted by inversions of the rhithral and the potamal. A local inversion is found near Kelheim (Danube break through the Franconian Alb, section 2). The first major inversion is below the Inn (section 3), the second is located below Bratislava (subsection 4.2), the third one is at the Iron Gate (section 7). The latter two inversions were disturbed after the Danube was dammed. Some taxonomic and nomenclatural problems in the genus Metacnephia Crosskey 1969, Simulium maculatum (Meigen 1804), the Simulium ornatum species-group, the Simulium reptans species-group, the Simulium tuberosum species-group, and the Simulium lineatum species-group, are discussed in connection with the Danube black fly fauna.
1 Introduction With a length more than 2800 km from its source area in the Black Forest in Germany to its outlet into the Black Sea (data on its length differ according to different sources: officially it is 2826 km including river Breg and 2780 km to Donaueschingen according to ICPDR, Schmedtje 2005 and Dr. H. Weber, pers. comm. to G. Seitz), the Danube is the second longest river in Europe and is the only large European river crossing Europe in a generally eastward direction. Its watershed comprising 817 000 km² (Domokos 2001) extends from the southwestern part of the German Mittelgebirge in the west to the Wallachian Plain and Moldovan Plateau in the east, and also covers the northern and eastern part of Alps, the Vienna Basin, most of the Carpathians, the Pannonian Plain, and the Balkans. During the Pleistocene, the Danube river basin, and the river itself, was larger, as it included those parts of Alps that now form part of the Rhine basin since the Upper Rhine has eroded backward and most of the waters from this part of Alps changed their direction. This region is of zoogeo-
94 graphic interest, as it has contacts with the Quaternary refugia and is also crossed by postglacial colonization routes as the Balkans were apparently a source of colonization for all species in the east and for many species in the west for which the ice-capped Alps were an initial barrier to their northward expansion (Cooper & al. 1995, Durand & al. 1999, Hewitt 1996, 2000, 2001, Taberlet & al. 1998). At present, the Danube also remains the southern corridor of colonization routes for modern invasive species, for plants and animals, aquatic and terrestrial (e.g. Bij de Vaate & al. 2002, Jehlík & al. 2005, Paunovic & al. 2007, van der Velde & Platvoet 2007, Wittman 2007). The aquatic fauna of the Danube is generally known. Forty years ago, the first comprehensive view of the macrozoobenthos of the Danube was published (Dudich 1967). Over the following decades, knowledge of the Danube fauna has been expanded and improved through many regional studies (e.g. Kusel-Fetzmann & al. 1998, Russev & al. 1998, Schmid 1994). During the past decade, some research projects have been carried out that have resulted in an upto-date list of the taxa recorded from the Danube. In all these studies, an apparent lack of data on black flies, one of the most typical rheophilous insect groups, is evident. Although Literáthy & al. (2002) have listed 4 taxa (S. austeni, S. ornatum, S. reptans, S. reptans var. galeratum) from upper sections 1 and 2, and S. ornatum even as bioindicator (sic!), this is only a fragmentary view of the black fly fauna of the upper reaches of the Danube. No black flies were listed among the macroinvertebrate reference species for the section types and reaches of the River Danube (Sommerhäuser & al. 2003). One of the reasons could be the sometimes problematic matter of identification, as well as the special methods needed for sampling black flies which are usually underestimated in hydrobiological studies (Crosskey 1990). So, although the aquatic fauna is well known, our knowledge of the black flies remains sparse (for details see below). The aim of this study is to summarize the known data on the black flies of the Danube, to identify the gaps in our knowledge of the black fly fauna, and to stimulate further studies in this field. 2 The River Danube The River Danube originates as the confluence of two source rivers, the Breg and the Brigach, near Donaueschingen at an altitude of 678 m a. s. l. Both source rivers rise on the eastern slopes of the Black Forest mountains (Germany): the Breg river (length 46 km) at an altitude of 1078 m a. s. l. and only 100 m away from the line that forms the European Watershed, and the Brigach river (43 km) at an altitude of 925 m a. s. l. Geographically, the source of the Danube is the source of the Breg, but traditionally the beginning of the Danube is considered to be the confluence of the Breg and Brigach near Donaueschingen.
95 The Danube then drains the Black Forest Mountains, in the north the Swabian and Franconian Alb, parts of the Oberpfälzer, the Bavarian and the Bohemian Forests, the Austrian Mühl- and Waldviertel, and the Bohemian-Moravian uplands, in the south the Swabian-Bavarian-Austrian Alpine foothills as well as large parts of the Alps (Schmedtje 2005). Only 26 km after its origin at Donaueschingen, the Danube enters the Danube Sink in the Swabian Alb Mountains, where it loses about half of its discharge in the porous limestone which is then carried by subsurface flows mainly into Aachtopf north of Lake Constance and feeds the Rhine. In the Bavarian section, the Danube is a medium-sized hilly river but then changes substantially when it receives its largest upstream tributary, the Inn River, which carries more water than the Danube itself. Together with other right-side tributaries draining the Alps, the River Inn imprints on the Danube the hydrological character of an alpine river. Passing the Viennese Basin, the Danube crosses the Little Carpathians at the Devín Gate and after some 150 km reaches through the northern part of the Pannonian lowland enters the Visegrád Gorge, between the foothills of the Western Carpathian and the Hungarian Transdanubian Mountains. In this Hungarian Danube Bend, the Danube turns to the south and crosses the Pannonian Plain, confined by the Carpathians in the north and the east (including the Transsylvanian uplands), and the Karnic Alps and the Karawankas, the Julian Alps and the Dinaric Mountains in the west and south. After breaking through the Southern Carpathians at the Iron Gate Gorge (Djerdap Gorge), the Danube flows across a plain, dividing the Danubian Plain of Bulgaria and the Romanian Walachian Plain. Near Tulcea, some 80 km from the sea, the river begins to spread out into its delta, with three main channels; 80 % of delta is located in the territory of Romania and 20 % belongs to the Ukraine. More than 80 % of the length of the Danube is regulated for flood protection, and approximately 30 % of its length is additionally impounded for hydropower generation, mainly in the upper part where a total of 59 dams have been built along the river's first 1000 kilometres. The largest Water Power and Navigation System Iron Gate I was completed in 1972, and in 1985 Iron Gate II was built 80 km downstream from the first one. The backwater zone of Iron Gate I extends 250-300 km upstream in the River Danube, reaching the cities of Belgrade and Novi Sad (Serbia). The second largest dam system has been operated at Gabčíkovo, downstream of Bratislava (Slovakia), since 1992. Nowadays the former river channel receives only some 20 % of the total flow, as the other 80 % is channeled around the dam towards the power plant (Petkovic & al. 2005). Several systems to subdivide the Danube into sections have been established, based on different parameters. Based on the character of the catchment, the
96 Danube was divided into the Upper, Middle and Lower Danube (Lászlóffy 1967). The Upper Danube reach extends from the source rivers down to the Devin Gate at the Austrian-Slovakian border and the confluence with the River Morava (March); MQ Bratislava: 2044 m³/s (Belz et al. 2004). In this region the Danube is influenced mainly by tributaries from the Alps. The Middle Danube reach, crossing the Pannonian basin, extends from the Devín Gate down to the Iron Gate. (MQ Orsova: 5611 m³/s, Belz et al. 2004).The Lower Danube reach is the lowland part of the Danube from the Iron Gate to the delta (MQ 6855 m³/s, Domokos 2001); the delta was classified as special part. Taking into account the river slope, the Danube has been subdivided into six sections (Lászlóffy 1967). Compared with the previous classification, this classification differentiates within the Upper Danube a mountain section from the confluence of the source rivers Brigach and Breg down to the confluence of the River Lech (average slope 1.01 m/km), and shifts the lower boundary of the Upper Danube section from the Devín Gate down to the rejoining of the Mosoni Danube (river km 1794, average slope 0.4 ‰). In this system, the Middle Danube section reaches downstream to the beginning of the Iron Gate cataract (river km 1071, average slope 0.06 ‰), whilst the Iron Gate (Cataract) reach is considered to be a separate section (river km 1040-941, average slope 0.28 ‰). The last two sections, the Lower Danube (river km 941-80, average slope 0.039 ‰) and the Danube delta (river-km 80-0, average slope 0.001 ‰ or lower), remain unchanged. Literáthy & al. (2002) based their subdivision of the Danube on geomorphological landscape features as well as on the anthropogenic impacts. They distinguished nine distinct reaches that in general follow the sections defined by Lászlóffy (1967) but with some exceptions. First, the Reach 1 of this classification starts at Neu Ulm (river km 2581) and does not include some 200 km upstream to Donaueschingen. The Reach 3 is the 64 km long lowest part of the Upper Danube section of Lászlóffy (1967), which is impacted by the construction of the Gabčíkovo Dam. The Middle Danube section is here divided into two reaches, upstream of Budapest and downstream of Budapest respectively. The Cataract reach of Lászlóffy is classified as Reach 6, which extends from Belgrade (confluence with the River Sava) to the Iron Gate Dam (river km 943). The Lower Danube is divided into Reaches 7 and 8, with the confluence with the River Jantra as the boundary between them, and Reach 9 corresponds to the Danube delta. Due to the fact that none of these proposed subdivisions takes into account the source rivers Breg and Brigach, the real mountain headwaters of the river (the River Danube including its source rivers) are not evaluated in these classifications. The crenal, epirhithral and metarhithral are absent from the system.
97 In a recent UNDP/GEF Danube Regional Project (Robert & al. 2003), a new section typology for the Danube meeting the requirements of the Water Framework Directive of the European Union was developed. This typology combines the four ecoregions according to Illies (1978), i.e. Central Highlands, Hungarian Lowlands, the Carpathians and Pontic Province, with different abiotic parameters (geomorphology, slope, lithology, water temperature etc.). The sections were proposed according to available information on the river and basin parameters (expert knowledge, geology, geomorphology, ecoregion, slope, etc) and then validated by analysis of the benthic macroinvertebrates. The Danube was subdivided into ten sections, with four of them (Nos 2, 4, 5, 6) subdivided into two and three subsection types, respectively (tab. 1). The boundaries of the sections and/or subsections differ according to different sources, but in the present study the boundaries defined by Robert & al. (2003) are adopted. Tab. 1. The section and sub-section types, compiled from Illies (1978), Moog & al. (2003) and Robert & al. (2003), with additional information. River km according to ICPDR (see Introduction). The country codes follow the internatinal codes for automobiles Eco- Section type region 9 0 Headwaters
Borders of the section types and Borders of the sub-section types and Country river km river km Brigach and Breg D
1 Upper Course of the Danube
11
10 12
Confluence of Brigach and Breg D to Neu Ulm km 2780-2581 2 Western Alpine Foot- Neu Ulm to Passau 2.1 Western Alpine Foothills D hills Danube km 2581-2225 km 2588-2376 2.2 Western Alpine Foothills/crystal- D line Basement km 2376-2225 3 Eastern Alpine Foot- Passau to Krems D, A hills Danube km 2225-2001 4 Lower Alpine Foothills Krems to Gönyü/Kližská Nemá 4.1 Krems to Devín A, SK Danube km 2001-1791/1790 (2001-1880) 4.2 Devín to Gyönyü/Kližská Nemá A, SK, H km 1880-1791/1790 5 Hungarian Danube Gönyü/ Kližská Nemá to Baja 5.1 Gyönyü/Kližská Nemá to Eszter- SK, H Bend km 1791/1790-1497 gom km 1791/1790-1719 5.2 Esztergom to Nagymaros/Visegrád SK, H km 1719-1695 5.3 Nagymaros/Visegrád to Baja H km 1695-1497 6 Pannonian Plain Dan- Baja to Bazias 6.1 Baja to tr. Drava H, HR, ube km 1497-1071 km 1497-1379 SRB 6.2 tr. Drava to tr. Sava HR, SRB km 1379-1170 6.3 tr. Sava to Bazias SRB, RO km 1170-1071 7 Iron Gate Danube Bazias to Turnu Severin SRB, RO km 1071-931 8 Western Pontic Dan- Turnu Severin to Chiciu/Silistra SRB, RO, ube km 931-378 BG 9 Eastern Wallachian Chiciu/Silistra to Isaccea RO Danube km 378-100 10 Danube Delta Isaccea to Sulina RO, UA km 100-0
Reach
Upper Danube
Middle Danube
Lower Danube
98 This typology was validated (Moog & al. 2003) using data on benthic macroinvertebrates obtained during the Joint Danube Survey (Literáthy & al. 2002). In general, the a priori typology has been validated. In the overall analysis, the three major reaches of the Danube, viz. Upper, Middle and Lower Danube, are well separated and identified, whereas the section types (e.g. section types 4 and 5) were only recognized in a detailed analysis of the separate major reaches. However, as mentioned earlier, the source rivers and Section type 1 were not included in the analysis because appropriate data were absent, and Section type 2 was also not included due to insufficient data for the section. 3 Black fly fauna The black fly fauna of the Danube is in general inadequately and unevenly known. Special attention has been paid to the black flies of the Danube in only three countries – in Germany, Serbia and Slovakia. In Germany, the study of the black flies of the Danube started with a few reports in Baden-Württemberg (Grunewald 1965) and Lower Bavaria (Seitz 1988). In Serbia, the first period of the study of black flies was mainly in connection with mass outbreaks of Simulium (Simulium) colombaschense (Scopoli 1780) and the damage to people and livestock (a series of papers published by Baranov in the years 1926-1939, see below). In the second period, the changes in black fly communities after the construction and operation of the Iron Gate I and Iron Gate II hydropower stations were documented by Živkovič (1975). In Slovakia, the study of the black flies of the Danube also started in connection with the construction of the Gabčíkovo hydropower plant and the early period of its operations in the last decade of the 20th century (Illéšová & Halgoš 1994, Illéšová & al. 1994, Illéšová & Stloukalová 1995, Jedlička & al. 1997). In the other Danube countries, the study of the black fly fauna has been less intensive or even non-existent (for details, see below). As data on the black flies of the Danube have been accumulated, recorded and published on a country-by-country basis, we have followed this scheme in the review presented below. 3.1. Germany Data on black flies from the upper reaches of the Danube, namely on those from the source rivers, are almost non-existent. Grunewald (1965), in his study on the black flies of the Black Forest, published a list of the black fly species recorded from this area, but partly without precise sites. He recorded Simulium (Nevermannia) carpathicum (Knoz, 1961) from the sources of the Breg, Simulium (Simulium) noelleri Friederichs, 1920 from the source of the Brigach, and a further 6 species [Simulium (Eusimulium) aureum Fries, 1824, Simulium
99 (Nevermannia) angustitarse (Lundström, 1911), Simulium (Wilhelmia) equinum (Linnaeus, 1758), Simulium (Boophthora) erythrocephalum (De Geer, 1776), Simulium (Simulium) ornatum Meigen, 1818, Simulium (Simulium) posticatum Meigen, 1838 (as S. venustum)] from the Danube, and of these S. aureum and S. posticatum in the Danube Sink zone only. The occurrence of another species in the Danube, Simulium (Nevermannia) vernum Macquart, 1826 (as S. latipes), is possible as Grunewald indicates its occurrence in nearly all streams. For the upper course of the Danube, Weber (1985) confirmed the occurrence of Simulium (Wilhelmia) lineatum (Meigen, 1804), S. lundstromi (as S. latigonium) and S. vernum, and Hackbarth (pers. comm.) has also collected S. noelleri there. So far as the source rivers are concerned, Hackbarth (pers. comm.) has also noted the occurrence of Prosimulium tomosvaryi (Enderlein, 1921) and Simulium (Simulium) argyreatum Meigen, 1838. More detailed data were subsequently published by Seitz (1992) and Schmid (1994). In the latter comprehensive study on the limnology and water quality of the Bavarian Danube, the German Danube stretch was divided into 6 sections delimited by the confluence of the Danube tributaries Lech, Altmühl, Regen, Isar, Inn and above Jochenstein (Austrian boundaries). From these sections, 7 black fly species have been recorded, and a further 8 black fly records which are included in the present study have not been previously published (tab. 2). 19 taxa have been found to date in the German stretch of the Danube, although the presence of S. angustitarse must be doubtful as this species prefers to live in the epirhithral of small meadow brooks (Seitz 1992). As some of the current samples from the Brigach (leg. Hackbarth), which have been made available to the junior author for identification, have to be assigned to the species S. lundstromi, it can be assumed that the black flies recorded by Grunewald (1965) as E. angustitarse were actually S. lundstromi. The Eusimulium latigonium (Rubtsov 1956) found by Weber (1985) in the Danube near Hausen (Section 1) is a synonym (cf. Bass & al. 1995) and is listed in table 2 under S. lundstromi. The Eusimulium aureum recorded by Grunewald (1965) from the Danube Sink zone and based on the determination of preimaginal stages must be listed as species-group taxon as the nominal species lives by preference in spring brooks and lake outlets in low mountain ranges. The black flies in this biotope should therefore belong to one of the sibling species. There is thus only one species in the German list of species, S. carpathicum, that lives by preference in the hypocrenal and has not been found in the Danube proper. When considering the individual sections of the Danube, it is apparent that the number of species in Section 1, Subsection 2.1 and Section 3 is in each case considerably higher than that in Subsection 2.2. This is evidently due to the fact that in this area the gradient of the Danube is relatively low and there is
Prosimulium hirtipes (Fries) Prosimulium rufipes (Meigen) Prosimulium tomosvaryi (Enderlein) Metacnephia danubica (Rubtsov) Simulium (Boophthora) erythrocephalum (De Geer) Simulium (Byssodon) maculatum (Meigen) Simulium (Eusimulium) angustipes Edwards Simulium (Eusimulium) aureum species-group [Simulium (Nevermannia) angustitarse (Lundström)]1 Simulium (Nevermannia) carpathicum (Knoz) Simulium (Nevermannia) cryophilum (Rubtsov) Simulium (Nevermannia) lundstromi (Enderlein) Simulium (Nevermannia) vernum Macquart Simulium (Obuchovia) auricoma Meigen Simulium (Simulium) argenteostriatum Strobl Simulium (Simulium) argyreatum Meigen Simulium (Simulium) colombaschense (Scopoli) Simulium (Simulium) degrangei Dorier & Grenier Simulium (Simulium) galeratum sensu Knoz2
Species
● 6
● ● ●1
●
●
●3 ●
●3,5 ● ●6 ●
1
● ●4
●3
●
●
●6 ●
●6
●6
●
9
● ●
●8 ●8 ●8 ●8 ● ● ●8
●7
●
●8 ●8 ● ●8
●
●
●
● ●
●
●
●
● ●
7
●
●
●
●
●
●8 ● ●10 ● ●
Sections/subsections (for details see Tab. 1) 0 1 2.1 2.2 3 4.1 4.2 5.1 6.1 6.2 6.3
●
8
Tab. 2. Occurrence of the Simuliidae species in sections and subsections of the Danube; sections/subsections without records are omitted 1 = probably a misidentification - see text, 2 = see chapter 4.4 in the text, 3 = first record from the (sub)section, leg. Hackbarth, 4 = Breg only, 5 = Brigach only, 6 = first record from the (sub)section, leg. Seitz, 7 = free flowing main river only, 8 = only before damming, 9 = indirect evidence, 10 = only after damming
100
Simulium (Simulium) monticola Friederichs Simulium (Simulium) morsitans Edwards Simulium (Simulium) noelleri Friederichs Simulium (Simulium) ornatum Meigen Simulium (Simulium) posticatum Meigen Simulium (Simulium) reptans (Linnaeus)2 Simulium (Simulium) variegatum Meigen Simulium (Simulium) voilense Sherban Simulium (S.imulium) vulgare Dorogost., Rubts. & Vlasenko Simulium (Wilhelmia) balcanicum (Enderlein) Simulium (Wilhelmia) equinum (Linnaeus) Simulium (Wilhelmia) lineatum (Meigen)
Species
● 5
● ●
● ● ● ● 3
●
●
●
● ● ● ● ●6 ●8 ●8 ● ● ●6 ●6 ● ● ● 7 ● ● ● ● ● ● ● ● ●9
●
●
●
●8 ●8 ● ● ● ● ●
● ● ● ●
●
●
●
Sections/subsections (for details see Tab. 1) 0 1 2.1 2.2 3 4.1 4.2 5.1 6.1 6.2 6.3
●
●
●
7
●
8
101
102 thus a higher degree of potamalisation. Accordingly, the Simuliidae coenosis consists of only a few typical potamal species, while the rhithral parts in the other sections support a higher diversity of species. Longitudinally, S. equinum, S. erythrocephalum, S. ornatum and Simulium (Simulium) reptans (Linnaeus, 1758) show the highest constancy: they live nearly everywhere in the German Danube, starting from the confluence of the Breg and Brigach. Section 1, which is about 200 km long, is characterised by the presence of S. posticatum, the current presence of which is confirmed by Hackbarth (pers. comm.). The species may be regarded as one of the index species of the flowing water body type "Large Rivers of the Low Mountain Range" ("type 9.2" according to Pottgieser & Sommerhäuser 2004), as the Danube is here classified (StMUGV 2005). With the dealpine Iller reaching the Danube near Ulm, the discharge regime of the Danube, which is pluvial due to the low mountain range "Schwäbische Alb", changes and the average discharge is increased by more than twice the original amount. In subsection 2.1 - "Western Alpine Foothills" - which begins here, P. tomosvaryi and Simulium (Simulium) morsitans Edwards, 1915 appear for the first time, whilst S. posticatum can no longer be found. It is true that in the list of species resulting from the "Joint Danube Survey", a ship expedition that was carried out in the late summer of 2001, one finding is cited of Simulium (Simulium) austeni Edwards, 1915 (synonym of S. posticatum) from the German-Austrian Danube stretch (Literáthy & al. 2002). However, this must be considered as doubtful as at this time the species is in aestival diapause. The appearance of P. tomosvaryi upstream of Kelheim (Seitz 1992) is a special case as this species lives mainly in the rhithral of smaller flowing water bodies. The only reason for its appearance here seems to be the rhithral character of this river section; and, additionally, the fact that the shaded rocks within the Danube breakthrough along the water line are covered with moss, which seems to satisfy the specific requirements of this species for its breeding habitat (Zwick & Zwick 1990). Unfortunately, it has not yet been possible to determine to species the taxon belonging to the S. (E.) aureum group. Due to the fact that the Altmühl, which discharges from the left near Kelheim, had contained a distinct population of Simulium (Eusimulium) velutinum Santos Abreu, 1922 before its canalisation to the Main-Danube Channel (Seitz 1994), it seems possible that this species may also live in the Danube. In the river subsection 2.2 ("Western Alpine Foothills/Crystalline Basement") downstream of Regensburg, in addition to the species that were mentioned above and which are present throughout the river, Simulium (Simulium) galeratum sensu Knoz and Simulium (Wilhelmia) balcanicum (Enderlein, 1924) could be found in the area downstream of the Isar confluence.
103 The latter species also appears in the lower course of the Isar where it replaces S. lineatum. Downstream of the confluence with the Inn, which has a much higher discharge (see above), the composition of the black fly coenosis differs materially from the other research sites because of the presence of four additional species: S. argyreatum, S. colombaschense and Simulium (Simulium) variegatum Meigen, 1818, for example, also live in the dealpine Inn and this reflects the changed character in the Danube water body. Simulium (Nevermannia) cryophilum (Rubtsov, 1959), which has also been found there, is a hypocrenal and epirhithral species living in the torrents of the Danube breakthrough that flow in from the left and from the right (Seitz 1992), and its preimaginal stages also find suitable living conditions in the Danube which is now summer-cold. 3.2 Austria The black fly fauna of Austria is in general well known (e.g. Car & Moog 1993, 2003, Ofenböck & al. 2002). Although it seems to be rather well documented (on average with one investigation site per 32 km2), the analyses indicate that there are still some gaps, for example the benthic fauna of the Viennese basin, a part of Pannonian basin, is poorly documented (Ofenböck & al. 2002, for S. balcanicum see Seitz 1994). Comprehensive synopses of the black fly fauna, and not only of the black flies, of the Austrian stretches of the Danube provide an overview (Moog & al. 1994, Zauner & al. 2001), but not along a longitudinal gradient as the Danube is classified into two parts: the free-flowing main river and backwaters of impoundments, with a third class formed by the backwaters of the riverside (not included here). We are interested mainly in the free-flowing main river channel, which is the remains of the original river. According to these synopses, it is clear that the fauna of the main channel consists of species typical of hyporhithral and epipotamal assemblages, and the differences between assemblages in the free-flowing and impoundment stretches do not seem to be significant. Taking into account the fact that P. tomosvaryi and S. colombaschense were recorded near Vienna (Humpesch & Anderwald 1988), and S. colombaschense also in the rhithral of the River Inn (Supperer & Kutzer 1964), the Danube can be considered as hyporhithral, and sites with S. erythrocephalum as (epi)potamal. This could be indirectly supported by knowledge of the black fly fauna of the River Inn, which is well known on the German side of the river (Seitz 1992).
104 3.3 Slovakia The Danube enters the territory of Slovakia through the Devín Gate, crossing the Little Carpathians. With the exception of a short river stretch near Bratislava (about 30 km, eight of which form Austrian-Slovak border), the Danube forms the border between Slovakia and Hungary (down to mouth of the River Ipeľ). Downstream of Bratislava, the Danube deposited a large fan of sediments in earlier times and in this section (approximately between km 1865-1804) it forms an inland delta with three main river branches (Danube, Little Danube, and Mosoni Danube) and a network of anastomosing and meandering anabranches from the parapotamal to the oxbow type. Some of the branches are only active during floods. The slope value decreases from 0.35 ‰ at the beginning of the delta to 0.10 ‰ at its end. Downstream from the inland delta, the Danube is a lowland river (hydrological middle stream). There are no early data on the black flies of this Danube section, including the common Slovak-Hungarian stretch (for a historical review, see Jedlička 1996), with the exception of unidentified larvae (Brtek & Rothschein 1964) which were mentioned in the original manuscript version of an MSc thesis by Brtek (unpublished) as the East Siberian species Prosimulium alpestre Dorogostaisky, Rubtsov et Vlasenko, 1935. According to the first reliable data, S. ornatum and S. lineatum occurred in the main channel, and S. erythrocephalum in a transient arm of the Danube (Jedlička 1970). In the floodplain of the Danube, in a suburb of Bratislava, 4 black fly species (S. ornatum, S. lineatum, S. erythrocephalum and S. reptans) were recorded attacking horses, with their supposed breeding sites in the Danube (Halgoš & Jedlička l971). Later, Halgoš & Jedlička (1987) found S. colombaschense in the main channel of the Danube. However, these partly accidental data based on hydrobiological samples and parasitological studies do not provide evidence of the real black fly fauna of the Danube. Contrasting with the considerable knowledge of other systematic groups of the benthic fauna, the black fly fauna of the Slovak part of the Danube remained almost unknown until recently. At the same time, the black flies of Bratislava and its surroundings (Jedlička & Halgoš 1986) as well as of Žitný ostrov island (Illéšová 1992) were studied, but these papers do not contain data actually from the Danube and/or its arms. Modern studies of the black flies of this stretch of the Danube began after the start of the construction of the hydropower station in Gabčíkovo in 1991, and then continued for some years to monitor the changes following the damming of the old river and the switching of its main discharge from the old river channel to a by-pass canal. The denuded banks and groynes made access to the river line possible and thereby the collection of black flies too.
105 In the old channel, the old Danube, 21 black fly species were recorded before and shortly after the damming (Illéšová 1992, Illéšová & al. 1994, Illéšová & Stloukalová 1995, Tab. 2) against 11 species known from watercourses in the adjacent lowlands (including Veľký Žitný ostrov, Great Danube Island); 5 species known from the lowland flows [S. angustipes, S. angustitarse, Simulium (Simulium) galeratum Edwards, 1920 sensu Knoz 1965, S. lundstromi, Simulium (Simulium) trifasciatum Curtis, 1839] have not been found in the Danube. Only 6 species occur in both the Danube and the adjacent lowlands (S. erythrocephalum, S. morsitans, S. noelleri, S. ornatum, S. equinum, and S. lineatum) whilst another 15 species have been found only in the Danube. The occurrence of some rhithral species was unexpected (e.g. Prosimulium hirtipes (Fries, 1824), Prosimulium rufipes (Meigen, 1830), P. tomosvaryi, S. vernum, Simulium (Obuchovia) auricoma Meigen, 1818, Simulium (Simulium) argenteostriatum Strobl, 1898, S. argyreatum, Simulium (Simulium) degrangei Dorier & Grenier, 1960, Simulium (Simulium) monticola Friederichs, 1920, S. reptans, S. variegatum, Simulium (Simulium) voilense Sherban, 1960, S. balcanicum). They are known from the rhithral of submontane flows and their presence indicates that this part of the Danube has the character of the rhithral zone. After the Gabčíkovo power plant was put into operation, the majority of the discharge was directed into the by-pass canal. Only 20 % of the original discharge remained in the old channel, and so the width decreased from 200-300 to 100 m and the flow rate from 1.5-2 m/s to 0.5-1 m/s or less (Kľúčovská & Topoľská 1995). As result of these changes, the species number decreased to 9 (S. erythrocephalum, S. colombaschense, S. noelleri, S. ornatum, S. reptans, Simulium (Simulium) vulgare Dorogostaisky, Rubtsov et Vlasenko, 1935, S. balcanicum, S. equinum, S. lineatum). The first to disappear were all the rhithral species. The species composition shifted towards the fauna of the adjacent lowland flows (Soerensen’s index of similarity rose from 0.42 before damming to 0.57 after damming). The species richness decreased much more in the upstream part of the old Danube, just below the dam, than it did in the downstream sites (Jedlička & al. 1997). Not only did the number of species change, but substantial changes were also recorded in community structure. The relative abundance of the pre-damming dominant species decreased significantly and rapidly: the highest relative abundance was attained by S. erythrocephalum (up to 85 %) while the cumulative relative abundance of all the other species together (S. colombaschense, S. noelleri, S. ornatum, S. reptans, S. vulgare, S. balcanicum, S. equinum, S. lineatum) decreased to 15 %. The community structure changed from the earlier rhithral to the potamal. There is no direct data on the black flies from the Danube subsection 5.1 downstream of Kližská Nemá. The females of S. erythrocephalum and S. linea-
106 tum (as salopiense) collected near the river bank for trials of arbovirus isolation (Danielová & al. 1978) may be considered to be indirect evidence of the possible breeding of these two species in the Danube. However, both species are known to breed in adjacent lowland water flows. 3.4 Hungary There are no reliable contemporary data except from the common SlovakHungarian part. Old records in Fauna Regni Hungariae (Thalhammer 1890) which are over a century old mentioned Simulium (Simulium) maculatum Meigen, 1804 and S. noelleri (as argyreata) from the Danube surroundings, but these data might not have been connected with the Danube itself as at that time the aquatic stages were not studied and all records were based on adults only. Furthermore, the data are not considered to be reliable, and revision of the material is no longer possible because the collection was completely destroyed (Papp 2001). More recent studies (Rubtsov 1967, Szabó 1971) did not deal with the black flies of the Danube itself. The only reliable data on the black flies of this stretch of the Danube were published recently by Kúdela (2003), who recorded 3 Wilhelmia species from the Danube: S. balcanicum, S. lineatum, S. equinum. According to the lowland landscape and potamal river character, we could infer that the same species occur there as in the Serbian part of the Danube and, as mass black fly attacks (Szabó 1971) have been recorded, the presence of S. erythrocephalum seems to be definite. 3.5 Serbia and Croatia The black flies of the Serbian part of the Danube, including the Serbian-Croatian border which is repeatedly crossed by the Danube, are among the best known thanks to the studies by Baranov and mainly by Živkovič. Baranov (1926, 1937, 1938a,b, 1939) recorded 4 species from the Danube at the Iron Gate (Djerdap Gorge): S. maculatum, S. reptans, S. colombaschense, S. equinum. From the adjacent regions (e.g. Golubac, Tuman), he named and described several specific and infrasubspecific taxa, now mostly considered to be synonyms (for details, see Crosskey & Peterson 1972). Most of them were described from the adult stage, with possible breeding sites in the Danube as well as in smaller tributaries, which were explicitly mentioned for some of them. As these taxa were not confirmed in the Danube by Živkovič (1955, 1967, 1969, 1971, 1975), we do not include them in the list of Danubian black flies for Serbia. Živkovič (1967, 1971, 1975) studied the entire river stretch, almost 600 km in length, which could be divided, based on the black fly fauna, into two parts the Pannonian part and the Iron Gate. The black fly fauna of each part is quite different. The presence of the most abundant species in Vojvodina (Pannonian
107 part) was recently confirmed in studies on black fly attacks (Ignjatović-Ćupina & al. 2003, 2006a,b). Živkovič (op. cit.) found 12 species in the Danube. Six of them (S. balcanicum, S. maculatum, S. reptans [Simulium (Simulium) pictum Meigen, 1818 with some doubt], S. galeratum, S. voilense, S. colombaschense) occur throughout the entire Serbian stretch, five only in the Pannonian stretch (S. equinum, S. lineatum, S. erythrocephalum, S. sp. ex gr. ornatum and S. sp. ex gr. vernum), and only one species, Metacnephia danubica (Rubtsov, 1956), restricted to the Danube at the Iron Gate. She thus recorded 11 species in the Pannonian part, the most abundant of which were S. erythrocephalum, S. maculatum, and species of the S. reptans group. At the Iron Gate, seven species were recorded, and those that also occurred in the Pannonian part were found at the Iron Gate mainly or exclusively in basins with a slower current, whereas M. danubica, S. colombaschense and S. balcanicum were found in rapids. After the damming of the Danube at the Iron Gate, M. danubica was no longer found, the population of S. colombaschense, the dominant species before damming, decreased substantially, and S. galeratum became the most abundant species, appearing in masses. S. erythrocephalum, which previously occurred in the Pannonian part, has only colonized breeding sites in the Iron Gate stretch, with a relative abundance up to 9 %. In general, the relative abundance of lowland species increased (Živkovič 1975) in the same way that we have found at Gabčíkovo more recently. 3.6 Romania, Bulgaria and Ukraine The Romanian part of the Danube includes the Iron Gate stretch (Portile de Fier) and all the lower reaches including the delta. As the entire Bulgarian stretch forms the common Bulgarian-Romanian border, and only about 20 % of the Danube delta belongs to the Ukraine, all the lower reaches including the delta are discussed here. In spite of the fact that a comprehensive book on the Romanian black flies was published by Dinulescu (1966), original primary data on Danubian black flies are scarce. The older data are connected with massive outbreaks of S. colombaschense (e.g. Ciurea & Dinulescu 1924). Dinulescu (1966) himself took over data partly from Živkovič (see also Živkovič 1971) and partly from Rubtsov, without original new records. For the Danube at the Iron Gate he accepted Rubtsov’s data on M. danubica, S. maculatum maculatum (Meigen, 1804) and S. maculatum danubense (Rubtsov, 1956), Živkovič’s (op. cit.) data on S. erythrocephalum, and Metacnephia blanci (Grenier & Theodorides, 1953) (without source) which was said to occur at the Iron Gate together with S. colombaschense, S. reptans and S. lineatum. He also listed some of Baranov’s names
108 which are now considered to be synonyms. Only the records published originally by Živkovič are accepted here. No species records were mentioned from the Walachian part. In a study on the black flies of the Danube tributaries, Kovatschev (1979) mentioned two species from the Romanian-Bulgarian Danube stretch, from the Danube itself, i.e. S. colombaschense and S. equinum, and these are the only records from the section 8. These records were quoted in a study on the diversity of zooplankton and zoobenthos in the Danube, its tributaries, and adjacent waterbodies (Russev & al. 1998); 60 black fly species were mentioned in this study, among them only two unnamed species from the Danube (river km 845-375). The citation in some bibliographical sources of a study by Zhdanova (1984) as being a study on black flies of the Danube delta is a mistake in the English translation. The paper actually deals with the biting midges (Ceratopogonidae) of the Dunayskie plavni reserve in the Ukrainian part of the Danube delta (Dr. A. V. Yankovski, pers. comm.). 3.7 Black fly fauna of the River Danube In summary, 31 named black fly species have been recorded in the Danube, but for two reasons this number may not be final: insufficient investigations and taxonomic problems. In spite of this, however, some conclusions are possible. Very little data are available from the lower reaches in Bulgaria and Romania as well from the Hungarian Danube Bend. 10 species have been recorded from the Austrian section, in both the main river channel and its backwaters. The most comprehensive data are from the German section (19 agreed taxa), the Slovak-Hungarian section (21 species before damming), the Pannonian Plain Danube section (10 species in Vojvodina), and the Serbian-Romanian sections (7 species before damming). There are no reliable records of black flies from 3 of the 10 proposed river sections. The highest species richness was found in subsection 4.2 (Devín Gyönyü/Kližská Nemá) with 21 recorded species, but this was the situation before the Danube was dammed at Gabčíkovo; after damming the number of species dropped to 9. The black fly fauna in sections 1 to 3 (German stretch) is reliably known, with 19 species pooled but 10 (11) species in each section; the real species number may be higher, particularly in rhithral sites. Reliable data on the black flies were compiled by Živkovič (1967, 1969, 1971, 1975) for sections 6 (Pannonian Plain Danube, 11 species) and 7 (Iron Gate Danube, 8 species pooled but only 7 after damming). However, the number of black fly species is not definitive. A higher number of species may be expected in the source rivers Breg and Brigach, where the oc-
109 currence of montane rhithral species can be anticipated by analogy with the rhithral stretches of other mountain water bodies and with subsection 4.2 before damming. From the ecological point of view, both source rivers are an integral part of the river regardless of the fact that geographically the Danube begins only at their confluence. A smaller increase in the species number is to be expected in the lower reaches of the Danube. Only a few species typical for large lowland rivers may occur there, and have not yet been recorded. Paradoxically, the increase in our knowledge of the black fly fauna of two stretches, i.e. 4.2 and 7, was a consequence of the damming of the river and the construction of two major hydropower plants (Iron Gate I, II and Gabčíkovo), which have substantially changed the nature of the river and the composition of its fauna and have probably caused the global (M. danubica) or regional extinction of some black fly species. 3.8 Inversions The available data on the black flies and their longitudinal distribution in the Danube do not enable us to assess the longitudinal and altitudinal zonation of black fly communities or to analyse zoogeographic aspects and the formation of the fauna in terms of the Quaternary climatic processes. In spite of the fact that many sections of the River Danube are in a far from natural condition, it seems to be possible to infer the original species assemblages. One phenomenon which seems to be clear is that the usual typical longitudinal zonation of the river along the longitudinal gradient sensu Vannote & al. (1980) is not continuous but is interrupted by inversions of the rhithral and the potamal. This means that the rhithral and potamal stretches with their characteristic species assemblages alternate between rapids and limnetic stretches, not as transition assemblages but as quite well differentiated units. A locally inferred inversion is found in the Bavarian stretch near Kelheim, where the Danube breaks through the Franconian Alb and where P. tomosvaryi occurs. The first great inversion is below the Inn, resulting in a hyporhithral character with S. argyreatum, S. colombaschense, S. cryophilum and S. variegatum as typical species. This inversion is caused by confluence with the summer-cold River Inn, a large alpine river that strongly affects the character of the Danube. In the upper reaches above the Inn and especially above the Isar, the Danube is a lowland river, but below the Inn (and other alpine rivers) the Danube becomes distinctly alpine in character, also hydrologically. The following inversions are connected with the Danube break through the Carpathians. The second inversion was located below Bratislava, where the Danube breaks through promontories of the West Carpathians and enters the Pannonian lowland. Even though the benthic fauna in the Viennese basin was
110 poorly known, we can suppose that potamal assemblages are still there or had originally developed there (occurrence of S. erythrocephalum). This inversion is a thing of the past since the Gabčíkovo hydropower station began operating. Then, after more than 600 km, the Danube breaks through the Carpathians for a second time, this time through the South Carpathians at the Iron Gate. The flow of the Danube along this gorge, with a length of more than 100 km, was never uniform, and consisted of four rapids and three basins, with different species assemblages as found by Živkovič. The situation here changed to a lesser extent than it did at Gabčíkovo. At Gabčíkovo, about 80 % of the discharge is diverted into the canal and the old channel receives some 20 %, up to 400 m3/s, following which the old Danube changed into a lowland river. At the Iron Gate the water flow was not diverted, and the stream velocity did indeed decrease, but it remains up to 1 m/s, which is suitable for rheophile species. 4 Taxonomy and nomenclature: unresolved questions around the Danube Some taxonomic questions concerning the black flies of the Danube remain unresolved, and this is why the species list of Danube black flies is not completely reliable. The most problematic areas are the Simulium reptans species-group, Wilhelmia species, Metacnephia species, S. maculatum, and the S. ornatum species-group. Another problem is posed by the groups of sibling species (e.g. the cryophilum or vernum species-groups) each of which is listed in the present paper as a single taxon on the morphospecies level. The clarification of these taxonomic questions is essential for resolving problems in faunistics, zoogeography and ecology - and not only in the Danube region. 4.1 Metacnephia species The description of M. danubica, which according to original description was similar to Cnephia pallipes (Fries, 1824), was based on larvae and on pupal gills with 30-32 filaments dissected from a mature larva, "breeding in the swiftest part of a large river (Danube) at Iron Gate (Bosman), with pupation time by end of April" (Rubtsov 1956). According to Živkovič (1975), the species has not been found there since the damming of the Danube. In an earlier paper, Živkovič (1955) mentioned M. blanci from Serbia (Danube, in the Djerdap Gorge) and Macedonia, but in her later papers she listed only M. danubica for the Danube. These earlier records of M. blanci (and that by Dinulescu 1966 for Romania) are probably the basis for the Serbian, Macedonian and Romanian entries in Fauna Europaea and in the inventory of world black flies (Crosskey 2005, Crosskey & Howard 2004). The question to be resolved is whether M. danubica and M. blanci are two separate species and, if that is indeed the case,
111 whether both are distributed in the lower reaches of the Danube and/or other rivers in the region. Moreover, Metacnephia lesnei (Séguy 1925), a questionable synonym of M. blanci described from the Eastern Pyrenees, may also be distributed in southern Europe and the Balkans according to Rubtsov (1956). Another species, Metacnephia uzunovi Kovachev, 1965 was described and is only known from Bulgaria (Kovachev 1985). It should be closely related to (or a synonym of?) Metacnephia nigra (Rubtsov, 1940), with a Caucasus-Turanian distribution (and reported from Romania by Dinulescu 1966), and to Metacnephia subalpina (Rubtsov, 1956) from the Caucasus. In the pupal stage, all three species differ in the shape of the pupal cocoon and in the number of gill filaments (36–40 in M. nigra, 47 in M. uzunovi, 50–56 in M. subalpina), and in the shape of the ventral plate. The status of these species remains unclear (cf. Crosskey & Zwick 2007). To make the matter even more obscure, Metacnephia fuscipes (Fries, 1824), distributed in northern Europe, was also listed from Hungary (Crosskey & Howard 2004, Papp 2001). 4.2 Simulium maculatum (Meigen, 1804) Simulium maculatum danubense Rubtsov, 1956 was described from larvae and pupae "breeding in the most rapid part of Danube river, where it crosses through the Carpathian range" (Rubtsov 1956) and is found only there. Rubtsov (1956) has described a number of infrasubspecific forms within S. maculatum and although these forms have no status in nomenclature the question arises: is this taxon with such a wide Holarctic distribution homogenous across the whole area? 4.3 Simulium ornatum species-group S. ornatum and/or the ornatum species-group/complex has been recorded in all sections of the Danube. S. baracorne Smart, 1944 was described from Golubac, Serbia (as ruficorne Baranov, 1926, preocc.). With regard to the present state of the ornatum-group taxonomy, it remains an open question as to which species, one or more, are breeding in the Danube. All the ornatum data listed in the present study should be therefore understood as an umbrella name for unseparated taxa within the ornatum species-group. 4.4 Simulium reptans species-group The entire reptans species-group, especially from central and southern Europe at least, is in need of a general revision. Two main questions are of major importance in connection with the Danube black flies.
112 S. colombaschense was described from the middle Danube region (the authorship and spelling were discussed by Thompson 2001) and was originally considered to be endemic to the Danube at the Iron Gate. We now know that it is distributed not only at the Iron Gate and not only in the Danube - its distribution area reaches as far as the Alps, Harz Mountains (Germany) and Italy (Halgoš & Jedlička 1987, Kowald 1971, Rivosecchi 1978, Seitz 1992, Supperer & Kutzer 1964, Werner & Adler 2004). However, its abundance at the terra typica (the Danube at the Iron Gate) decreased substantially after the river was dammed (Živkovič 1975), and one could now consider it as a threatened species in terms of an IUCN assessment. Serban (1960) has described S. voilense, which is similar to S. colombaschense but with minor differences. The latter species was said to be restricted to the Danube and its affluents whilst S. voilense was said to occur in submontane rivers and streams, but doubt has been cast on this, as both species have been found together in the Danube (Živkovič 1971, 1975, Jedlička & al. 1997). Although both species are accepted as valid, their validity needs to be verified. One problem is that the type of S. colombaschense probably no longer exists, even if it ever existed, and the search for the type of S. voilense has remained unsuccessful until now. Another problem is posed by S. reptans and S. galeratum (or other species). Culex reptans was described by Linnaeus from Sweden as the most annoying black fly there. Edwards (1920) considered S. reptans to consist of one main form, S. reptans, and a variety, galeratum, which Rubtsov (1956) raised to species rank but later considered to be a synonym of reptans (pers. comm. from Rubtsov to Carlsson, see Carlsson 1962), and this is how these forms have been treated by recent authors (e.g. Crosskey 2005, Crosskey & Howard 2004). The two forms are sometimes sympatric (Carlsson 1962) but morphologically they differ especially by the head spots in the larvae and the gills in the pupae; Rubtsov (1956) also found small differences in the genitalia. All the specimens from Scandinavia examined by Carlsson (1962) have been S. galeratum sensu Edwards, and Carlsson considers it to be most probable that all pupae and adults from Scandinavia are S. galeratum auct. as used in the modern literature. Consequently, the true S. reptans sensu Linnaeus is identical with S. galeratum auct. (or S. reptans var. galeratum) as used in the modern literature, and all specimens previously considered to be galeratum are in reality S. reptans sensu Linnaeus. The taxon named reptans by Edwards (1920), Grenier (1953) and Rubtsov (1956) would then be a distinct species, for which Carlsson proposed the name S. reptantoides Carlsson, 1962. The existence of two species was supported by Knoz (1965), who gave a description of galeratum based on material from the former Czechoslovakia, as the species named by Edwards as var. galeratum and by Carlsson as reptans (partim.). If we accept Carlsson’s
113 view, then we have in Central Europe S. reptans sensu Linnaeus (galeratum auct.) and S. reptantoides (reptans auct.), but paradoxically reptantoides (recorded here as reptans) is found in cold submontane streams and the true northern reptans (recorded as galeratum sensu Knoz) in southern and warmer lowland flows. If galeratum sensu Knoz is another species, what is its identity? In the present study we retain the name galeratum sensu Knoz for the taxon known from lowland rivers in Central and South Europe, including the Danube, but we are conscious that this conclusion is only tentative. Another question is the identity of S. pictum sensu Rivosecchi (1978) (a synonym of reptans – Crosskey & Howard 2004). Živkovič (1969, 1971, 1975) mentioned all three names, reptans, galeratum and pictum, but pictum with a question mark as a possible synonym of reptans. 4.5 Simulium tuberosum species-group Three nominal species of the tuberosum species-group have been recorded from Central Europe (chronologically): Simulium janzeni Enderlein, 1922, Simulium tuberosum (Lundström, 1911), and S. vulgare. The latter species was first recorded in Central Europe by Novák (1956) and Knoz (1965), partly as S. tuberosum, and it was only distinguished later (Knoz & Šašinková 1969, Knoz 1980). This is still the current viewpoint, although there are some doubts as to whether S. vulgare from Central Europe is the same species that was described from East Siberia and is known from North America, China, Mongolia, etc. S. tuberosum itself is a complex of karyologically defined forms. More than ten years ago, Adler & Kuusela (1994) found larvae in material from Baden-Württemberg (Germany) which, after karyotaxonomic analysis, appeared to be a species close to Simulium (Simulium) subtile Rubtsov, 1956, described from the Caucasus and recorded in Italy and Romania. This conclusion was recently confirmed by Adler (pers. comm. to the junior author) for the entire material from Bavaria identified by the junior author as S. vulgare sensu Knoz. Could this be identical with the forgotten species S. janzeni, described by Enderlein (1922) from Styria? 4.6 Simulium lineatum species-group Although both S. lineatum and S. balcanicum have repeatedly been recorded in Central Europe including the River Danube, the specific status and validity of the latter is doubtful. The authors dealing with Danube black flies have based their identification on pupae, using the gill filaments which are forked in balcanicum but separated in lineatum. However, Zwick could not find any differences between the adults (Crosskey 1998, Crosskey & Zwick 2007, Zwick 1995). Both forms can be found together as well as separately, but it seems that S. bal-
114 canicum has only recently invaded the German stretch of the Danube (cf. Seitz 1994). Acknowledgements The authors would like to thank Dr. A. V. Yankovsky, Saint Petersburg, and Dr. A. IgnjatevićĆupina, Novi Sad, for help with providing otherwise inaccessible literature resources, W. Hackbarth for confirmations of faunistic data, and Dr. M. Kúdela for a thorough reading of an earlier version of this manuscript and for his valuable comments. We gratefully acknowledge the help of Adrian C. Pont, Goring-on-Thames, for linguistic revisions of the manuscript. The study was partly supported by grant number 1/2349/05 of VEGA Grant Agency of the Ministry of Education of Slovak Republic and Slovak Academy of Sciences. This publication is part of the main research topics of the International Association for Danube Research (IAD). References Adler, P. H. & K. Kuusela (1994): Cytological identities of Simulium tuberosum and S. vulgare (Diptera: Simuliidae) with notes on other Palaearctic members of the S. tuberosum speciesgroup.- Entomologica Scandinavica 25: 439-446, Lund Baranov, N. (1926): Über die serbischen Simuliiden.- Neue Beiträge zur systematischen Insektenkunde 3: 193-194, Berlin Baranov, N. (1937): K poznavanju golubačke mušice V. (Studij epidemiologije golubačke mušice na invaziji g. 1936).- Veterinarski Arhiv 7: 229-276, Beograd Baranov, N. (1938a): Die Kolumbatscher Mücke in Jugoslawien im Jahre 1937.- Archiv für wissenschaftliche und praktische Tierheilkunde 72: 158-164, Berlin Baranov, N. (1938b): K poznavanju golubačke mušice VI. (Studij golubačke mušice I njeznih sinbiocenota).- Veterinarski Arhiv 8: 313-328, Beograd Baranov, N. (1939): La mouche de Golubatz.- Bulletin - Office International des epizooties 18: 311-322, Paris Bass, J. A. B., R. W. Crosskey & D. Werner (1995): On the European blackfly Simulium lundstromi and inclusion within this species of S. latigonium as a new synonym.- The British Simulium Group Bulletin 5: 7-19, Liverpool Belz, J. U., L. Goda, Z. Buzas, M. Domokos, H. Engel & J. Weber (2004): Das Abflussregime der Donau und Ihres Einzugsgebietes. Die Donau und ihr Einzugsgebiet - eine hydrologische Monographie, Folgeband VIII/2.- Deutsches Nationalkommitee für das International Hydrological Programme (IHP) der Unesco und das Hydrology and Water Resources Programme (HWRP) der WMO, 152 pp + 4 maps, Koblenz Bij de Vaate, A., K. Jazdzewski, H. A. M. Ketelaars, S. Gollasch & G. Van der Velde (2002): Geographical patterns in range extension of Ponto-Caspian macroinvertebrate species in Europe.Canadian Journal of Fishery and Aquatic Sciences 59: 1159-1174, Toronto Brtek, J. & J. Rothschein (1964): Ein Beitrag zur Kenntnis der Hydrofauna und des Reinheitszustandes des tschechoslowakischen Abschnittes der Donau.- Biologické práce 10(5): 1-62, Bratislava Car, M. & O. Moog (1993): Höhen- und längenzonale Verteilung der Simuliidenfauna (Diptera) Österreichs und Liechtensteins.- Essener Oekologische Schriften 2: 63-79, Essen Car, M. & O. Moog (2003): Simuliidae. In Moog, O. (ed.): Fauna Aquatica Austriaca. Ministry of Agriculture, Forestry, Environment and Water Management, Vienna Carlsson, G. (1962): Studies on Scandinavain Black Flies (fam. Simuliidae).- Opuscula Entomologica supplements 21: 1-280, Lund
115 Ciurea, I. & G. Dinulescu (1924): Ravages causés par la mouche de Golubatz en Roumanie, ses ataques contre les animaux et contre l’homme.- Annals of Tropical Medicine and Parasitology 18: 323-337, Liverpool Cooper S. J. B., K. M. Ibrahim & G. M. Hewitt (1995): Postglacial expansion and genome subdivision in the European grasshopper Chorthippus parallelus.- Molecular Ecology 4: 49-60, Oxford Crosskey R. W. (1990): The natural history of blackflies.- 711 pp., (John Willey & Sons Ltd.) Chichester Crosskey, R. W. (1998). Records of blackflies from mainland Greece (Diptera: Simuliidae).Entomologist's Gazette 49: 277-283, Brightwood Crosskey R. W. (2005): Fauna Europaea: Simuliidae. In Jong H. de (ed.): Fauna Europaea: Diptera, Nematocera. Fauna Europaea, version 1.2, http://www.faunaeur.org, Amsterdam Crosskey R. W, & T. M. Howard (2004): A Revised Taxonomic and Geographical Inventory of World Blackflies (Diptera: Simuliidae) http://www.nhm.ac.uk/research-curation/projects/blackflies/index.html Crosskey, R. W. & B. V. Peterson (1972): The Simuliidae described by N. Baranov and their types (Diptera). Bulletin of British Museum (Natural History) 27: 189-214, London Crosskey, R. W. & Zwick, H., (2007): New faunal records, with taxonomic annotations, for the blackflies of Turkey (Diptera, Simuliidae).- Aquatic Insects 29: 21-48, London Danielová, V., D. Málková, J. Minář, B. Rehse-Küpper, Z. Hájková, J. Halgoš & L. Jedlička (1978): Arbovirus isolations from mosquitoes in south Slovakia.- Folia parasitologica 25: 187-191, Praha Dinulescu, G. (1966): Diptera. Fam. Simuliidae (Muştele columbace). Fauna Republicii socialiste România, Insecta, XI, 8.- 600 pp., (Editura Acad. Rep. Socialiste Romane) Bucuresti Domokos, M. (2001): Drei Jahrzehnte hydrologische Zusammenarbeit der Staaten des Donaueinzugsgebietes (1971-2000).- Hydrologie und Wasserbewirtschaftung 45, H. 4: 144-154, Koblenz Dudich, E. (1967): Systematisches Verzeichnis der Tierwelt der Donau mit einer zusammenfassenden Erläuterung.- In: Liepolt, R. (ed.): Limnologie der Donau - Eine monographische Darstellung. Kapitel V: 4-69 (Schweizerbart) Stuttgart Durand, J. D., H. Persat & Y. Bouvet (1999): Phylogeography and postglacial dispersion of the chub (Leuciscus cephalus) in Europe.- Molecular Ecology 8: 989–997, Oxford Edwards F.W. (1920): On the British species of Simulium. II. The early stages; with corrections and additions to part I.- Bulletin of Entomological Research 11: 211-246, Cambridge Grenier, P. (1953): Simuliidae de France et d'Afrique du nord, systématique, biologie, importance médicale.- Encyclopédie Entomologique Paris, sér. A, 29: 1-170, Paris Grunewald, J. (1965): Zur Kenntnis der Simuliidenfauna (Diptera) des Süd-Schwarzwaldes und seiner Randgebiete.- Beiträge zur Naturkundlichen Forschung in Südwest-Deutschland 24: 143-152, Karlsruhe Halgoš, J. & L. Jedlička (1971): The black-flies (Diptera, Simuliidae) attacking horses in the environs of Bratislava.- Biológia (Bratislava) 26: 431-433, Bratislava Halgoš, J. & L. Jedlička (1987): Faunistical notes: Diptera, Simuliidae: Simulium colombaschense.Biológia (Bratislava) 42: 1026, Bratislava Hewitt, G. M. (1996): Some genetic consequences of ice ages, and their role in divergence and speciation.- Biological Journal of the Linnean Society 58: 247-276, London Hewitt, G. M. (2000): The genetic legacy of the Quaternary ice ages.- Nature 405: 907-913, London Hewitt, G. M., (2001): Speciation, hybrid zones and phylogeography - or seeing genes in space and time.- Molecular Ecology 10: 537–549, Oxford Humpesch, U. H. & P. Anderwald (1988): Beitrag zur Faunistik der österreichischen Donau - das Makrozoobenthos bei Stromkilometer 2005.- Wasser und Abwasser 32: 41-55, Wien
116 Ignjatović-Ćupina, A., D. Petrić, M. Zgomba, A. Konjević & S. Grabovac (2003): Distribution of some blackfly species (Diptera: Simuliidae) in the region of Novi Sad.- The British Simuliid Group Bulletin 21: 6-7, London Ignjatović-Ćupina, A., D. Petrić, M. Zgomba, A. Konjević, S. Grabovac & D. Marinković (2006a): Estimating the biting risk to humans by the black fly species that are most abundant in the region of Novi Sad (Vojvodina Province, Serbia and Montenegro).- Proceedings of the International Symposium on Simuliidae.- Studia dipterologica Supplement 14: 31-44, Halle (Saale) Ignjatović-Ćupina, A., M. Zgomba, L. Vujanović, A. Konjević, D. Marinković & D. Petrić (2006b): An outbreak of Simulium erythrocephalum (Dre Geer, 1776) in the region of Novi Sad (Serbia) in 2006.- Acta entomologica Serbica, Suplement: 97-114, Beograd Illéšová, D. (1992): Black-flies (Diptera, Simuliidae) of Žitný ostrov on Podunajská nížina Lowland.- Biológia (Bratislava) 47: 107-122, Bratislava Illéšová, D. & J. Halgoš (1994): Faunistic notes. New findings of black flies (Diptera, Simuliidae) from the River Danube on the territory of Slovakia.- Biológia, Bratislava, 49: 228, Bratislava Illéšová, D., L. Jedlička & J. Halgoš (1994): Black flies of the Danube inland delta (Diptera, Simuliidae).- Biológia, Bratislava, 49: 223-227, Bratislava Illéšová, D. & V. Stloukalová (1995): Changes in the black fly communities (Diptera, Simuliidae) of the Danubian inland delta.- In: Mucha I. (ed.): Gabčíkovo part of the Hydroelectric Power Project - Environmental impact review: 307-310 (Faculty of Natural Sciences, Comenius University) Bratislava Illies, J. (1978): Limnofauna Europaea - Eine Zusammenstellung aller die europäischen Binnengewässer bewohnenden mehrzelligen Tierarten mit Angaben über ihre Verbreitung und Ökologie. 2. Auflage.- 532 pp., (G. Fischer) Stuttgart Jedlička, L. (1970): Die Verbreitung der Kriebelmücken (Diptera, Simuliidae) in der Südslowakei.Acta Rerum Naturalium Musei Nationalis Slovaciae 16: 45-56, Bratislava Jedlička, L. (1996): A synopsis of blackfly fauna of Slovakia (Diptera, Simuliidae).- Acta Zoologica Universitatis Comenianae 40: 49-67, Bratislava Jedlička, L., J. Halgoš (1986): Poznámky o muškovitých v mestskom prostredí Bratislavy (Diptera, Simuliidae).- Dipterologica bohemoslovaca 4: 105-107, Česká Budějovice Jedlička, L., D., Illéšová & V. Stloukalová (1997): Die Kriebelmückenfauna (Diptera: Simuliidae) der Donau im Bereich des Inlanddeltas bei Gabčíkovo (Slowakei).- Schriftenreihe des Instituts für Parasitologie und Zoologie der Veterinärmedizinschen Universität Wien 1: 25-29, Wien Jehlík, V., J. Dostálek & M. Zaliberová (2005): Spreading of adventive plants on river banks of the Elbe River in the Czech Republic and the Danube River in Slovakia outside of harbours.Thaiszia.- Journal of Botany 15: 35-42, Košice Kľúčovská J. & J. Topoľská (1995): Water level regime in the Danube river and its river branches.In: Mucha, I. (ed.): Gabčíkovo part of the Hydroelectric Power Project - Environmental impact review: 33-42 (Faculty of Natural Sciences, Comenius University) Bratislava Knoz, J. (1965): To Identification of Czechoslovakian Black-Flies (Diptera, Simuliidae).- Folia Facultatis Scientiarum Naturalium Universitatis Purkynianae Brunensis, Biologia 2 (6/5): 1-54 + 425 figs, Brno Knoz, J. (1980): Simuliidae.- In: Chvála, M. (ed.): Fauna ČSSR vol. 22, Krevsající mouchy a střečci: 144-281, Praha Knoz, J. & V. Šašinková (1969): Zur Kenntnis der Kriebelmücken (Simuliidae, Diptera) im DyjeGebiet in Morava.- Folia Facultatis Scientiarum Naturalium Universitatis Masarykinae Brunensis, Biologia 10(25): 13-44, Brno Kovatschev, S. (1979): Bemerkungen über die Zusammensetzung, Verbreitung und Ökologie der Simuliden (Diptera, Simulidae) in der Bulgarischen Donauzuflüssen.- XXI. Arbeitstagung IAD, 392-396, Wien Kovachev, S. G. (1985): Metacnephia uzunovi sp. n. - a new black fly (Diptera, Simuliidae) from Bulgaria, with notes on its bionomy.- Acta zoologica bulgarica 28: 63-65, Sofia
117 Kúdela, M. (2003): First Hungarian records of three blackfly species representing the subgenus Wilhelmia (Diptera: Simuliidae, Simulium).- Folia Entomologica Hungarica 63: 363-365, Budapest Kusel-Fetzmann, E., W. Naidenow & B. Russev (1998): Plankton und Benthos der Donau.- Internationale Arbeitsgemeinschaft Donauforschung (ed.): Ergebnisse der Donauforschung Band 4, 376 pp. Wien Lászlóffy, W. (1965): Die Hydrographie der Donau. Der Fluss als Lebensraum.- In: Liepolt, R. (ed.): Limnologie der Donau - Eine monographische Darstellung. Kapitel II: 16-57. (Schweizerbart) Stuttgart Literáthy, P., V. Koller-Kreimel & I. Liška (2002) (eds.): Final Report of the Joint Danube Survey.261 pp., (International Commission for the Protection of the Danube River) Wien Moog, O., M. Konar & U. H. Humpesch (1994): The macrozoobenthos of the River Danube in Austria.- Lauterbornia 15: 25-51, Dinkelscherben Moog, O., T. Ofenböck & T. Battisti (2003): Typology of the Danube River - part 2: bottom-up validation.- In: Sommerhäuser, M., S. Robert, S. Birk, D. Hering, O. Moog, I. Stubauer, T. Ofenböck (eds): GEF Danube regional project "Strengthening the implementation capacities for nutrient reduction and transboundary cooperation in the Danube River basin, Activity 1.1.6 "Developing the typology of surface waters and defining the relevant reference conditions". Final report: 15-24, (University of Duisburg-Essen) Essen Novák, V. (1956): Příspěvek k poznání muchniček (Simuliidae, Diptera) ČSR.- Věstník Československé zoologické společnosti 20: 224-248, Praha Ofenböck, T., O. Moog & M. Car (2002): Do the Austrian blackfly fauna (Diptera: Simuliidae) support the typological approach of the EU water framework directive?- Limnologica 32: 255-272, Amsterdam Papp, L. (2001): Simuliidae.- In: Papp, L. (ed.): Checklist of the Diptera of Hungary: 87-89 (Hungarian Natural History Museum) Budapest Paunovic, M. M., D. G. Jakovcev-Todorovic, V. M. Simic, B. D. Stojanovic & P. D. Cakic (2007): Macroinvertebrates along the Serbian section of the Danube River (stream km 1429–925).- Biologia (Bratislava) 62: 214-221, Bratislava Petkovic, S., M. Babic-Mladenovic & M. Damnjanovic (2005): 4.9 Technical, hydrological and environmental aspects of the Iron Gate I and II Dams.- In: Bloesch, J., D. Gutknecht, & V. Iordache, (eds): Hydrology and Limnology –Another Boundary in the Danube River Basin. A contribution to the IHP Ecohydrology Project IHP. UNESCO IHP-VI Technical Document in Hydrology 75: 46-48, (UNESCO) Paris Pottgiesser, T. & M. Sommerhäuser (2004): Fließgewässertypologie Deutschlands.- In: Steinberg, C., W. Calmano, W. Klapper & R.-D. Wilken: Handbuch Angewandte Limnologie 19. Ergänzungslieferung. VIII-2.1: 1-16 + Anh., (Ecomed) Landsberg Rivosecchi, L. (1978): Simuliidae. Diptera Nematocera.- Fauna d'Italia, 533 pp. (Edizioni Calderini) Bologna Robert, S., S. Birk & M. Sommerhäuser (2003): Typology of the Danube River - part 1: top-down approach.- In: Sommerhäuser, M., S. Robert, S. Birk, D. Hering, O. Moog, I. Stubauer, T. Ofenböck (eds): GEF Danube regional project "Strengthening the implementation capacities for nutrient reduction and transboundary cooperation in the Danube River basin, Activity 1.1.6 "Developing the typology of surface waters and defining the relevant reference conditions". Final report: 6-12, (University of Duisburg-Essen) Essen Rubtsov, I. A. (1965): Moshki (Simuliidae). Fauna SSSR.- 860 pp. (Izdateľstvo Akademii Nauk SSSR) Moskva Rubzow, I. A. 1967: Über die Kriebelmücken Ungarns.- Annales Historico-Naturales Musei Nationalis Hungarici. Pars Zoologica 59: 303-318, Budapest Russev, B., A. Petrova, I. Janeva & S. Andreev (1998): Diversity of Zooplankton and Zoobenthos in the Danube River, its tributaries, and adjacent water bodies.- In: Meine, C. (ed.): Bulgaria's
118 Biological Diversity: Conservation Status and Needs Assessment. Biodiversity Support Program. Volumes I and II. http://www.worldwildlife.org/bsp/publications/europe/bulgabulgaria/bulgaria.html#toc Schmedtje, U. (2005) (ed.): The Danube River Basin District.- ICPDR Document IC/084, 184 pp. (ICPDR) Wien Schmid, R. (1994) (ed.): Limnologie und Gewässergütezustand der bayerischen Donau und ihrer wichtigsten Zubringer. Donaubasisuntersuchung 1985-1992. Ergebnisse der Donauforschung.291 pp., (Internationale Arbeitsgemeinschaft Donauforschung) Wien Seitz, G. (1988): Beitrag zur Kriebelmückenfauna Niederbayerns (Insecta, Diptera, Simuliidae).Nachrichtenblatt der Bayerischen Entomologen 37: 89-94, München Seitz, G. (1992): Verbreitung und Ökologie der Kriebelmücken (Diptera: Simuliidae) in Niederbayern.- Lauterbornia 11: 1–230, Dinkelscherben Seitz, G. (1994): Neue und bemerkenswerte Kriebelmückenfunde (Diptera: Simuliidae) für die deutsche Fauna.- Lauterbornia 15: 101-109, Dinkelscherben Sommerhäuser, M., S. Robert, S. Birk, D. Hering, O. Moog, I. Stubauer & T. Ofenböck (2003) (eds): GEF Danube regional project "Strengthening the implementation capacities for nutrient reduction and transboundary cooperation in the Danube River basin, Activity 1.1.6 "Developing the typology of surface waters and defining the relevant reference conditions". Final report.- 97 pp. (University of Duisburg-Essen) Essen Sommerhäuser, M., S. Robert & S. Birk (2003): Proposal of the reference communities of macroinvertebrates of the Danube River.- In: Sommerhäuser, M., S. Robert, S. Birk, D. Hering, O. Moog, I. Stubauer & T. Ofenböck (eds): GEF Danube regional project "Strengthening the implementation capacities for nutrient reduction and transboundary cooperation in the Danube River basin, Activity 1.1.6 "Developing the typology of surface waters and defining the relevant reference conditions". Final report: 53-58, (University of Duisburg-Essen) Essen Stančik, A., S. Jovanovič, A. Sikora, L. Ürge & D. Miklós (1988): Hydrology of the River Danube.271 pp., (Publishing House Príroda) Bratislava StMUGV (2005): Umsetzung der Europäischen Wasserrahmenrichtlinie 2000/60/EG (WRRL). Bericht zur Bestandsaufnahme für das Deutsche Donaugebiet.- 133 pp., (Bayerisches Staatsministerium für Umwelt, Gesundheit und Verbraucherschutz, Ministerium für Umwelt und Verkehr Baden-Württemberg) München Supperer, R. & E. Kutzer (1964): Beitrag zur Kriebelmücken-Fauna (Diptera, Simuliidae) Österreichs. II.- Zeitschrift für Parasitenkunde 25: 49-51, Wien Szabó, J. B. (1971): Black fly swarms and the possibilities of eradication at Tata, Hungary.- Parasitologica Hungarica 4: 169-179, Budapest Taberlet, P., L. Fumagalli, A. G. Wust-Saucy & J.-F. Cossons (1998): Comparative phylogeography and postglacial colonization routes in Europe.- Molecular Ecology 7: 453–464, Oxford Thalhammer, J. (1890): Ordo Diptera.- In: A Magyar Birodalom Állatvilága. A Magyar Birodalomból eddig ismert állatok renderszeres lajstroma. Fauna Regni Hungariae. Animalium Hungariae hucusque cognitorum enumeratio systematica. (A. K. M. Természettudományi társulat) 76 pp. + 1 map, Budapest Thompson, F. C. (2001): The name of the type species of Simulium (Diptera: Simuliidae): a historical footnote.- Entomological News 112: 125-129, Philadelphia van der Velde, G. & D. Platvoet (2007): Quagga mussels Dreissena rostriformis bugensis (Andrusov, 1897) in the Main River (Germany).- Aquatic Invasions 2: 261-264, St. Petersburg Vannote, R. L., G. W. Minshall, K. W. Cummins, J. R. Sedel & C. E. Cushing (1980): The river continuum concept.- Canadian Journal of Fishery and Aquatic Science 37: 130-137, Toronto Weber, A. (1985): Cytotaxonomische und ökologische Untersuchungen an Larven von Wilhelmia equina (Linne, 1747) und Wilhelmia lineata (Meigen, 1804) (Simuliidae, Diptera).- Diplomarbeit der Fakultät für Biologie der Universität Tübingen
119 Werner, D. & P. H. Adler (2004): A faunistic review of the black flies (Simuliidae, Diptera) of the federal state of Sachsen-Anhalt, Germany.- Abhandlungen und Berichte für Naturkunde 27: 205-245, Magdeburg Wittman, K. J. (2007): Continued massive invasion of Mysidae in the Rhine and Danube river systems, with first records of the order Mysidacea (Crustacea: Malacostraca: Peracarida) for Switzerland.- Revue Suisse de Zoologie 114: 65-86, Genève Zauner, G., P. Pinka & O. Moog (2001): Pilotstudie Oberes Donautal.- Wasserstraßendirektion Wien Zhdanova, T. G. (1984): Ekologo-faunisticheskiye issledovaniya krovososushchikh mokretsov v rayone Dunayskikh plavney.- In Assanovich, T. A. (ed.): Dvukrylye fauny SSSR i ikh rol' v ekosistemakh: 36-38 (Akademiya Nauk SSSR, Zoologicheskiy Institut) Leningrad Zwick, H. (1995): Contribution to the European blackfly taxa (Diptera: Simuliidae) described by Enderlein.- Aquatic Insects 17: 129-173, Lisse Zwick, H. & P. Zwick (1990): Terrestrial Mass-Oviposition of Prosimulium-Species (Diptera: Simuliidae).- Aquatic Insects 12: 33-46, Lisse Živkovič, V. (1955): Morfološka i ekološka ispitivan’a simulida Dunava s naročnym osvrtom na golubačku mušicu.- Srpska Akademia Nauka Monographies 245(4): 1-95, Beograd Živkovič, V. (1967): Simulide (Diptera, Simuliidae) Dunava na području Vojvodine.- Acta veterinaria (Beograd) 17: 433-438, Beograd Živkovič, V. (1969): Simuliide (Diptera, Simuliidae) Vojvodine. II deo.- Acta veterinaria Beograd 19: 397-306, Beograd Živkovič, V. (1971): Black flies (Diptera, Simuliidae) of the Danube in Yugoslavia.- Acta veterinaria (Beograd) 21: 225-236, Beograd Živkovič, V., (1975): Present state of black flies (Diptera, Simuliidae) in the Djerdap gorge (Iron gate) of the Danube in Yugoslavia.- Acta veterinaria (Beograd) 25: 279-285, Beograd Addresses of the authors: Prof. Dr. Ladislav Jedlička, Department of Zoology, Comenius University, Mlynská dolina B-1, SK-84215 Bratislava, Slovakia;
[email protected] Dr. Gunther Seitz, District Government of Lower Bavaria, Regierungsplatz 540, D-84028 Landshut, Germany;
[email protected] Received: 2008-01-24
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