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Research Article
New faunistic records of chironomids and phantom midges (Diptera, Chironomidae and Chaoboridae) from Ukraine indicate recent climatic refugia in the Eastern Carpathians
expand article infoPeter Bitušík‡, Milan Novikmec§, Marek Svitok§|, Ladislav Hamerlík‡
‡ Matej Bel University, Banská Bystrica, Slovakia
§ Technical University in Zvolen, Zvolen, Slovakia
| Czech University of Life Sciences Prague, Prague, Czech Republic
¶ Institute of Zoology, Slovak Academy of Sciences, Bratislava, Slovakia

Corresponding author: Peter Bitušík ( peter.bitusik@umb.sk )

Academic editor: Viktor Baranov
© 2024 Peter Bitušík, Milan Novikmec, Marek Svitok, Ladislav Hamerlík.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Bitušík P, Novikmec M, Svitok M, Hamerlík L (2024) New faunistic records of chironomids and phantom midges (Diptera, Chironomidae and Chaoboridae) from Ukraine indicate recent climatic refugia in the Eastern Carpathians. ZooKeys 1211: 349-367. https://doi.org/10.3897/zookeys.1211.125436
ZooBank: urn:lsid:zoobank.org:pub:E6BB88C5-5795-4E80-9888-82146DD4CE9C
Open Access

Abstract

The aquatic insect fauna of the Eastern Carpathians is poorly known, especially in Ukraine. To address this knowledge gap, we conducted faunistic surveys of Chironomidae and Chaoboridae in 2018 and 2021. The study involved sampling of 11 watercourses and 10 mountain lakes situated in the Ukrainian part of the Eastern Carpathians. A total of 101 taxa were identified, including 40 chironomid species and one genus that have been recorded for the first time from Ukraine. The occurrence of one species previously considered as “doubtfully present” in Ukraine was confirmed by this study. One of the two identified phantom midge species, Chaoborus (s. str.) obscuripes (van der Wulp, 1859), is recorded for the first time from Ukraine. The most intriguing records are chironomid species Cricotopus (s. str.) beckeri Hirvenoja, 1973, Eukiefferiella bedmari Vilchez-Quero & Laville, 1987, and Pseudorthocladius (s. str.) berthelemyi Moubayed, 1990. These species have Mediterranean distribution and their occurrence in the Eastern Carpathians could be remains of once-widespread populations that currently survive in the Carpathian refugia due to adverse climatic conditions in the former distribution area. The high number of first records from a relatively small number of sites indicates a great gap in the knowledge of the Ukrainian chironomid fauna.

Key words

climatic relicts, mountain lakes, pupal exuvia, submontane rivers

Introduction

Chironomids are the most ubiquitous free-living holometabolous insects known from all zoogeographic regions, and all climatic zones from the tropics to the polar regions, including Antarctica (Ashe and O’Connor 2009). Recently, 7290 species belonging to nearly 440 genera and 11 subfamilies have been described worldwide (Ferrington 2007; Pape et al. 2011). In immature stages, most species inhabit various types of freshwaters although some species thrive in brackish water and intertidal pools, and few are truly marine. Finally, semi-terrestrial and fully terrestrial species are also known (Sæther et al. 2000). Among aquatic insects, Chironomidae is the most species-rich insect family found in freshwater ecosystems (Cranston 1995; Ferrington 2007). The species richness of the family recorded from only one stream locality is often astonishing. In some cases, the number of chironomid species recorded is higher than the diversity of all the other benthic macroinvertebrates (own observations).

Chironomidae can withstand an extremely wide range of environmental conditions in terms of water column depth, temperature, pH, dissolved oxygen, habitat drying and, finally, the gradient of human impacts such as pollution, habitat modification, and changes in watersheds (Ferrington 2007 and references therein). Consequently, chironomids have attracted the attention of researchers around the world as biological indicators for environmental impact assessments, ecosystem health, palaeolimnological reconstructions, and climate change (Resh and Rosenberg 2008; Eggermont and Heiri 2012; Nicacio and Juen 2015 and references therein). Compared to chironomids, the global diversity of phantom midges is low. The Chaoboridae family consists of about 51 extant species in six genera and two subfamilies (Borkent 2014). The immature stages usually live in standing waters, in some cases in small, temporary ponds. Larvae are predators and mostly planktonic; they are often considered keystone species that can eliminate or strongly suppress other invertebrates in the community (MacKay et al. 1990). Subfossil remains of the Chaoborus genus have been used in palaeoenvironmental research but also in contemporary ecological studies (e.g., Luoto and Nevalainen 2009; Tolonen et al. 2012). Despite the indisputable importance of both the chaoborids and chironomids and the rapid progress in the knowledge of their distribution, there are still areas that are “terra incognita”. The Eastern Carpathians are undoubtedly one of them.

Here, we present results from an ongoing faunistic inventory of chironomids and chaoborids from Ukrainian Carpathian Mountain lakes, which are supplemented by results from earlier investigations of the flowing waters in this territory.

Materials and methods

Study area

This study was conducted in the Ukrainian Carpathians located in the northern part of the Eastern Carpathians, extending through the western part of Ukraine (Fig. 1). The total length of this mountain range is approximately 240 km with total area of ca 24,000 km2. The Ukrainian Carpathians are medium-high mountains with the highest elevation slightly exceeding 2000 m a.s.l. (Novikoff and Hurdu 2015; Vyshnevskyi and Donich 2021). The studied area is characterised by complex geology consisting mostly of flysch with different constituents. Only small areas are formed by limestone, shales, and volcanic rocks, predominantly andesites and gneisses (Ivanik et al. 2019). We sampled 11 streams and 10 mountain lakes (Table 1). Except for lakes Sinevir and Dragobratske located in deciduous and coniferous forests, respectively, all studied lakes are situated above 1500 m a.s.l., i.e., above formerly accepted climatic tree line (Kobiv 2017). The semi-natural meadows and pastures (so-called polonynas in the vernacular) dominate the catchment vegetation. The proportion of dwarf pine (Pinus mugo), dwarf juniper (Juniperus communis subsp. nana), green alder (Alnus viridis), and scattered Norway spruce (Picea abies) is different in individual catchments.

Figure 1. 

Map showing of sampling localities of Chironomidae and Chaoboridae in the Ukrainian part of the Eastern Carpathians. The site abbreviations correspond to the Table 1. Inset shows location of the investigated territory within the Carpathian Mountains (grey area).

Table 1.
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Basic characteristics of the sampling sites. Strahler stream order was estimated from Google Earth Pro. Stream width, depth and lake depth were estimated in the field. Lake area was derived from Google Earth Pro using Polygon tool.

Stream/Lake name Code Latitude, Longitude Elevation (m) Stream order Av. width (m) Av. depth (m)
Zhdenivka River S1 48°46.29'N, 22°58.68'E 409 IV. 6 0.30
Nameless headwater stream S2 48°49.19'N, 22°58.58'E 777 II. 2 0.15
Tributary of the Zhdenivka River S3 48°47.98'N, 22°57.41'E 454 III. 3 0.20
Latoricja River S4 48°45.00'N, 23°00.75'E 360 IV. 20 0.30
Gluhana peat bog channel S5 48°27.94'N, 23°37.94'E 596 I. 0.4 0.10
Rika River (village Soimy) S6 48°34.00'N, 23°28.53'E 449 IV. 35 0.15
Rypenka River (village Soimy) S7 48°33.92'N, 23°28.29'E 448 IV. 25 0.30
Rika River (above Zaperedillia village) S8 48°29.08'N, 23°30.24'E 382 V. 30 0.50
Nameless tributary of the Volovets River S9 48°33.14'N, 23°35.51'E 725 II. 2 0.15
Tereblia River (village Sinevyr) S10 48°28.25'N, 23°38.16'E 573 IV. 18 0.30
Sukhar brook (village Kolochava) S11 48°25.40'N, 23°42.58'E 580 III. 10 0.30
Tisa River (Vinohradiv) S12 48°08.06'N, 23°05.15'E 124 VII. 120 –
Area (ha) Max. depth (m)
Sinevir Lake L1 48°37.01'N, 23°41.04'E 989 4.20 22
Brespo Lake L2 48°08.62'N, 24°30.94'E 1627 0.03 0.4
Bolotnoe ozerce L3 48°08.55'N, 24°31.25'E 1695 0.01 0.9
Brebeneskul Lake L4 48°06.10'N, 24°33.74'E 1793 0.60 3.2
Dragobratske Lake L5 48°14.45'N, 24°14.45'E 1382 0.07 1.5
Apshynets Lake L6 48°16.91'N, 24°09.53'E 1491 2.97 1.8
Geryshaska (Dohiaska) Lake L7 48°16.25'N, 24°09.93'E 1585 2.58 2
Kosivske Lake L8 48°15.78'N, 24°11.96'E 1614 0.13 1
Ivor Lake L9 48°13.70'N, 24°14.10'E 1606 0.04 2
Small Ivor Lake L10 48°13.69'N, 24°14.07'E 1602 0.04 0.6

All the studied flowing water sites except the Tisa River were located in the Carpathians. Most of the sampling sites were situated in valleys of the submontane belt up to 550–600 m a.s.l. (established according to climate and vegetation characteristics; Golubets 1977), only two sites (S2, S9) were situated in the lower montane belt above 700 m a.s.l. Stream bottom substrates were mostly dominated by cobbles with an admixture of finer components such as gravel and sand. The bottom habitats of both mountain brooks were more complex due to the presence of boulders and a considerable proportion of woody debris. Because of the bed surface roughness and shallow depth, turbulent flow is generally prevailing. Even though most sites were located near roads and often in settlements, the channels, banks, and riparian vegetation were visibly well preserved, without the signs of artificial modifications.

The above characteristics mostly do not apply to the Upper Tisa River. The river stretch close to the town of Vinohradiv flows in a lowland landscape (the Hungarian lowland ecoregion; Afanasyev et al. 2020). The floodplain is covered with alluvial forests containing willow and poplar. In the studied section, the low slope and high discharge altered the river channel morphology significantly, forming a high degree of channel sinuosity. The basic characteristics of the sampling sites are summarised in Table 1.

Sampling and identification

Chironomids and chaoborids were collected during the sampling campaigns in May 2018 (flowing waters and Sinevir Lake) and in August 2021 (mountain lakes in the Chornohora and Svidovets Massifs). A hand net attached to a telescopic handle (mesh size 250 μm, frame diameter 25 cm) was used to skim the water surface and collect floating material along the shores of streams and lakes. In lakes, the material was collected at the leeward shore; in flowing waters, the floating material was collected along an ~ 100-meter-long stretch while moving upstream.

On the shore, the netted sample was placed in a labelled 100-ml plastic bottle and preserved with 75% ethanol. In the laboratory, the samples were placed in a Petri dish and all chironomid material was picked up under a stereomicroscope (7.5–50×). Sorted exuvia, pupae, and adults were mounted on microscopic slides and identified following the keys of Langton and Visser (2003), Ekrem (2004), Stur and Ekrem (2006), and Langton et al. (2013) for pupal exuvia and Langton and Pinder (2007a, 2007b) for adults under high magnification (400×) using phase contrast. Chaoborid larvae, which were collected accidentally during the chironomid sampling of lakes, were identified using Parma (1969) and Sæther (1997).

The nomenclature and distribution of species are consistent with Fauna Europaea (de Jong 2016), and Ashe and O’Connor (2009, 2012). Voucher specimens are deposited in the collections of the Dept. of Biology and Environmental Studies, Faculty of Natural Sciences, Matej Bel University in Banská Bystrica.

Results and discussion

A total of 2088 specimens were collected and identified as 99 chironomid species/taxa (belonging to 43 genera from 5 subfamilies) and two chaoborid species of the same subfamily and genus. Altogether, 40 species and one genus of Chironomidae, and one species of Chaoboridae were recorded for the first time in Ukraine. The occurrence of one chironomid species, Nilotanypus dubius (Meigen, 1804), previously considered as “doubtfully present” in Ukraine (Spies and Sæther 2013) was finally confirmed. A list of all species/ taxa recorded is given below (sampling site codes refer to Table 1; Pe after the genus name refers to a morphotype not associated with an adult by Langton (1991); # – previously doubtfully present in Ukraine, * – first record of species/ genus from Ukraine). For detailed information on the abundance and life stages of collected specimens see Supplementary file 1.

CHIRONOMIDAE

Tanypodinae

Procladius (Holotanypus) choreus (Meigen, 1804): L5, L6, L9

Nilotanypus dubius (Meigen, 1804)#: S1, S6, S7, S8

Thienemannimyia carnea (Fabricius, 1805)*: S7

Zavrelimyia barbatipes (Kieffer, 1911): L6

Diamesinae

Diamesa (Diamesa) cinerella Meigen, 1835*: S8

Diamesa (Diamesa) cf. tonsa (Haliday, 1856): S6

Diamesa (Diamesa) vaillanti Serra-Tosio, 1972*: S7

Potthastia Pe1 Langton, 1991: S4, S7, S8

Sympotthastia macrocera Serra-Tosio, 1973*: S6

Prodiamesinae

Prodiamesa olivacea (Meigen, 1818): L1

Orthocladiinae

Brillia bifida (Kieffer, 1909): S3

Brillia flavifrons (Johannsen, 1905)*: S7

Corynoneura celtica Edwards, 1924*: S1, S3, S10

Corynoneura cf. scutellata Winnertz, 1846: L1, L6

Corynoneura Pe2a Langton, 1991: S1, S4, S5, S6, S7, S8

Corynoneura Pe4 Langton, 1991: S1

Cricotopus (Cricotopus) annulator Goetghebuer, 1927: S1, S4, S5, S6, S7

Cricotopus (Cricotopus) beckeri Hirvenoja, 1973*: S4, S6, S8

Cricotopus (Cricotopus) curtus Hirvenoja, 1973*: S6

Cricotopus (Cricotopus) fuscus (Kieffer, 1909): S1, S11

Cricotopus (Cricotopus) pallidipes Edwards, 1929*: S8

Cricotopus (Cricotopus) similis Goetghebuer, 1921*: S1, S7, S8

Cricotopus (Cricotopus) tremulus (Linnaeus, 1758)*: S1, S3, S6, S8

Cricotopus (Cricotopus) trifascia Edwards, 1929: S4, S11

Cricotopus (Cricotopus) vierriensis Goetghebuer, 1935: S1, S4, S6, S7, S8

Cricotopus Pe 17 Langton, 1991: S4, S8

Cricotopus (Isocladius) reversus Hirvenoja, 1973*: L1

Cricotopus (Isocladius) sylvestris (Fabricius, 1794): L6, L7

Cricotopus (Isocladius) Pe 5 Langton, 1991: L6, L7

Cricotopus (Paratrichocladius) rufiventris (Meigen, 1830): S4, S7, S10

Eukiefferiella bedmari Vilchez-Quero & Laville, 1987*: S4

Eukiefferiella brevicalcar (Kieffer, 1911): S2, S3

Eukiefferiella clypeata (Thienemann, 1919)*: S6, S7, S11

Eukiefferiella coerulescens (Kieffer, 1926): S3, S8

Eukiefferiella devonica (Edwards, 1929): S9

Eukiefferiella fuldensis Lehmann, 1972: S10

Eukiefferiella ilkleyensis (Edwards, 1929): S4, S6, S7, S8, S10

Euryhapsis Pe1 Langton, 1991: S11

Heleniella serratosioi Ringe, 1976*: S1, S4, S6, S7, S8, S9, S10, S11

Krenosmittia boreoalpina (Goetghebuer, 1944)*: S1, S3, S6, S7, S8, S10

Krenosmittia camptophleps (Edwards, 1929)*: S10

Limnophyes cf. asquamatus Andersen, 1937: L3

Nanocladius (Nanocladius) parvulus (Kieffer, 1909): S3, S4, S6, S8

Nanocladius (Nanocladius) rectinervis (Kieffer, 1911)*: S6, S7, S8, S10, S11

Orthocladius (Orthocladius) dentifer Brundin, 1947: L7

Orthocladius (Orthocladius) excavatus Brundin, 1947*: S7, S8, S11

Orthocladius (Orthocladius) oblidens (Walker, 1856)*: S11

Orthocladius (Orthocladius) pedestris Kieffer, 1909*: S1, S3, S4, S7, S8, S11

Orthocladius (Orthocladius) rivinus Potthast, 1914*: S3

Orthocladius (Orthocladius) rubicundus (Meigen, 1818): S1, S3, S4, S7, S8, S10

Orthocladius (Euorthocladius) ashei Soponis, 1990*: S4, S7, S8, S11

Orthocladius (Euorthocladius) rivicola Kieffer, 1911: S3, S6, S8, S10, S11

Orthocladius (Euorthocladius) rivulorum Kieffer, 1909: S4, S6, S7, S8

Paracricotopus niger (Kieffer, 1913)*: S4, S6, S7, S8, S11

Parakiefferiella bathophila (Kieffer, 1912)*: S4, S7, S8

Parametriocnemus stylatus (Spaerck, 1923): S1, S3, S4, S6, S7, S8, S9, S10, S11

Psectrocladius (Psectrocladius) limbatellus (Holmgren, 1869): S11

Psectrocladius (Psectrocladius) oligosetus Wuelker, 1956: L2, L3, L9, L10

Psectrocladius (Psectrocladius) schlienzi Wuelker, 1956*: L1

Pseudorthocladius (Pseudorthocladius) berthelemyi Moubayed, 1990*: S6

Rheocricotopus (Psilocricotopus) chalybeatus (Edwards, 1929): S1, S4, S6, S7, S8

Rheocricotopus (Rheocricotopus) fuscipes (Kieffer, 1909): S1, S3, S7, S8

Rheosmittia spinicornis (Brundin, 1956)*: S2, S3, S10

Symbiocladius rhithrogenae (Zavrel, 1924): S11

Synorthocladius semivirens (Kieffer, 1909): S3, S10

Thienemanniella majuscula (Edwards, 1924): S1

Thienemanniella Pe 1b Langton, 1991: S3

Thienemanniella Pe 2 Langton, 1991: S10

Tvetenia verralli (Edwards, 1929)*: S6

Chironominae

Benthalia sp.: L7

Demicryptochironomus Pe1 Langton, 1991: S11

Microtendipes chloris (Meigen, 1818): S5, S7

Microtendipes pedellus (De Geer, 1776): L6

Paracladopelma mikianum (Goetghebuer, 1937)*: S6, S11

Phaenopsectra flavipes (Meigen, 1818): S5, L1, L2, L6, L7

Polypedilum (Polypedilum) albicorne (Meigen, 1838)*: S4, S6, S7, S11

Polypedilum (Polypedilum) laetum (Meigen, 1818): S11

Polypedilum (Polypedilum) nubeculosum (Meigen, 1804): S11

Polypedilum (Pentapedilum) sordens (van der Wulp, 1875): S7

Polypedilum (Pentapedilum) cf. uncinatum (Goetghebuer, 1921): L2, L5, L7

Polypedilum (Tripodura) cf. apfelbecki (Strobl, 1900): S6

Polypedilum (Uresipedilum) convictum (Walker, 1856): S4, S11

Cladotanytarsus (Cladotanytarsus) atridorsum Kieffer, 1924: L6, L7, L8

Cladotanytarsus (Cladotanytarsus) vanderwulpi (Edwards, 1929): S6, S7, S8

Micropsectra atrofasciata (Kieffer, 1911): S6, S7

Micropsectra lindrothi Goetghebuer, 1931*: L7

Micropsectra lindebergi Saewedal, 1976/ insignilobus Kieffer, 1924: S5

Neozavrelia Pe1 Langton, 1991*: S1, S4, S5, S6, S7, S8

Paratanytarsus austriacus (Kieffer, 1924): L6

Paratanytarsus dissimilis (Johannsen, 1905)*: S5

Paratanytarsus laccophilus (Edwards, 1929): L2, L5, L7, L8, L9, L10

Rheotanytarsus pentapoda (Kieffer, 1909)*: S6, S7

Rheotanytarsus rhenanus Klink, 1983*: S1, S6, S7

Stempellinella flavidula (Edwards, 1929)*: S8

Tanytarsus aberrans Lindeberg, 1970*: L6

Tanytarsus debilis (Meigen, 1830)*: L5, L7

Tanytarsus gregarius Kieffer, 1909: L2, L4

Tanytarsus heusdensis Goetghebuer, 1923*: S6, S7

Virgatanytarsus Pe1 Langton, 1991: S4

CHAOBORIDAE

Chaoborinae

Chaoborus (Chaoborus) crystallinus (De Geer, 1776): L3

Chaoborus (Chaoborus) obscuripes (van der Wulp, 1859)*: L7

Of the 99 recorded chironomid taxa, 22 were found exclusively in lakes, and 20 of them only in alpine lakes. Fourteen lacustrine species/ taxa (i.e., 70%) were also found in lakes during our previous research (Bitušík et al. 2020) indicating a similar species composition of all the studied lakes. Phantom midges were not deliberately targeted since the larvae recorded were caught by chance while collecting pupal exuvia. However, one of the species recorded, Chaoborus (s. str.) obscuripes (van der Wulp, 1859), appears to be the first record in the Ukrainian Carpathian alpine lakes. Most chironomid species have been found in flowing waters. Overall, they are considered typical for streams and rivers of Western and Central Europe (e.g., Caspers 1991; Laville and Vinçon 1991; Ruse 1995; Orendt 2002a, 2002b; Schöll and Haybach 2004; Bitušík et al. 2006; Calle-Martínez and Casas 2006; Prat et al. 2016 and citation therein). Because most of the newly recorded species are widespread in stagnant and flowing waters in Europe and the Palaearctic Region, and do not increase our knowledge on their ecology, only species with restricted distributions and rarely collected elsewhere are discussed further in more detail. Regarding the common species not discussed further in the text, their geographical distribution is documented in Ashe and O’Connor (2009, 2012) and de Jong (2016), while their ecology is summarised in Moller Pillot (2009, 2013) and Vallenduuk and Moller Pillot (2007).

Family Chironomidae

Subfamily Diamesinae

Tribe Diamesini

Diamesa (Diamesa) vaillanti Serra-Tosio, 1972

Material examined

1 pupal exuvium, Rypenka river (S7), 7 May 2018.

Distribution

Palaearctic: Germany, Switzerland, France, Italy, Austria, Slovakia, Poland, Russia, Spain, Turkey, Morocco (Ashe and O’Connor 2009), and Azerbaijan (Kownacki 1985).

Habitat

Rheophilic species inhabiting high-altitude springs, streams (including glacier-fed), and rivers with rocky bottoms, but also alpine lakes (Bitušík 2004; Lencioni et al. 2011; Rossaro and Lencioni 2015). The species is cold-stenothermal (e.g., Rossaro 1991) although the findings from streams in the High Atlas indicate that it can tolerate relatively high temperatures of up to 22 °C (Azzouzi et al. 1992).

Remarks

The occurrence of the species is limited to waters at high altitudes. Since our record is from 448 m a.s.l., our finding is exceptional.

Sympotthastia macrocera Serra-Tosio, 1973

Material examined

2 pupal exuvia, Rika River (S6) in Soimy village, 7 May 2018.

Distribution

Palaearctic. For a long time, known only from Western Europe (France, Germany; Ashe and O’Connor 2009). More recent data comes from the Drava River in Croatia (Kresonja 2018). Our record is evidence for the current easternmost occurrence in Europe, but there are indications that the distribution of this species may extend as far as the Ural Mountains (Krasheninnikov 2012).

Habitat

Generally, larvae of the genus Sympotthastia inhabit cold running waters and springs (Sæther and Andersen 2013). A few data indicate that S. macrocera is probably a rheophilic species.

Remarks

Sympotthastia macrocera appears to be a relatively rare species with little known ecology.

Subfamily Orthocladiinae

Cricotopus (Cricotopus) beckeri Hirvenoja, 1973

Material examined

1 pupal exuvium, Latoricja River (S4), 5 May 2018; 1 pupal exuvium, Rika River (S6); 2 pupal exuvia, Rika River (S8) above Zaperedillia village, 7 May 2018.

Distribution

Palaearctic. France, Spain, Greece, Madeira, Corsica, Turkey, Algeria, Morocco, Slovakia. Its questionable occurrence in Finland (Ashe and O’Connor 2012) was not accepted later Paasivirta (2014).

Habitat

Principally inhabits the rhithral zone of streams at lower altitudes (Hirvenoja and Moubayed 1989; Kettani and Langton 2012; Moubayed-Breil and Ashe 2016).

Remarks

Cricotopus beckeri has been considered an exclusively Mediterranean species (Laville and Reiss 1992). Our record is the second reliable finding in the Carpathians (Bitušík and Langton 1994) far from its continuous distribution. We assume that the isolated populations in the Carpathians could be remnants of once widespread populations, which currently survive in refugia due to adverse climatic conditions. Thus, they could be considered a climatic relict, as customary for some plant species (Molnár et al. 2017). Interestingly, Reiss (1986) already hypothesised that the extra-Mediterranean occurrence of another Mediterranean chironomid, Paratanytarsus mediterraneus Reiss, 1981, could have a relict character in the Middle Rhine.

Cricotopus (Cricotopus) pallidipes Edwards, 1929

Material examined

1 pupal exuvium, Rika River (S8), 7 May 2018.

Distribution

Palaearctic. Finland, Norway, France, Portugal, Spain, Germany, Great Britain, Ireland, Romania, Hungary, Russia, Lebanon, and Morocco (Cobo et al. 2002; Soriano and Cobo 2006; Móra et al. 2006; Ashe and O’Connor 2012).

Habitat

The ecological requirements of this species are still unclear. It has been found in flowing and stagnant waters in cold climatic zones (Aagaard et al. 1997; Pozdeev 2012) to warm rivers, canals, lakes, and marshes in central and southern Europe, and North Africa (e.g., Laville and Tourenq 1968; Móra et al. 2006; Abbou and Fahde 2017; Moubayed et al. 2019). French authors (Tourenq 1976; Moubayed et al. 2019) consider it a lacustrine species, tolerant of low oxygen content.

Remarks

Currently known only from few European countries. It does not seem to be abundant anywhere. In Bavaria and the Sauerland Mountains (Germany), it is listed among possibly endangered species; however, its status is unknown (Orendt and Reiff 2003; Dittmar 2012).

Eukiefferiella bedmari Vilchez-Quero & Laville, 1987

Material examined

3 pupal exuvia, Latoricja River (S4), 5 May 2018.

Distribution

Palaearctic. France, Spain, Greece, Corsica, Turkey, Lebanon, Algeria, and Morocco (Ashe and O’Connor 2012).

Habitat

Streams and rivers (Laville and Langton 2002; Chaib et al. 2013; Moubayed-Breil and Ashe 2016).

Remarks

E. bedmari is a circum-mediterranean faunistic element (Laville and Reiss 1992; Moubayed-Breil 2008). Our unexpected extra-Mediterranean finding suggests the relict character of its population in the Carpathians (see comments to Cricotopus beckeri).

Orthocladius (Orthocladius) rivinus Potthast, 1914

Material examined

1 pupal exuvium, left-hand tributary of Zhdenivka River (S3), 7 May 2018.

Distribution

Palaearctic. Norway, Great Britain, Ireland, Austria, Slovakia, Hungary, Poland, Belarus, Germany, Switzerland, France, Italy, Spain, Canary Islands, and Portugal (Cobo et al. 2002; Ashe and O’Connor 2012; Moller Pillot 2013; Móra et al. 2013; Sołtys-Lelek et al. 2014).

Habitat

Rheophilic species inhabiting springs and flowing waters from small streams to large rivers, although it has been reported also from lakes (Langton and Visser 2003). Rossaro et al. (2003) underline its preference for cold waters, but some findings question this (e.g., Langton and Orendt 1996; Móra et al. 2013).

Remarks

The species is known from a few European countries and is generally considered rare. Like the ambiguous data on its ecology, this may also be the result of misidentification.

Psectrocladius (Psectrocladius) schlienzi Wuelker, 1956

Material examined

1 pupal exuvium, Lake Sinevir (L1), 7 May 2018.

Distribution

Palaearctic? In addition to some European countries (Austria, Denmark, Finland, Germany, Great Britain, Italy, Moldova, Netherlands, Norway, Portugal, Slovakia, Czech Republic, Spain, Sweden, Switzerland; Ashe and O’Connor 2012; Syrovátka and Langton 2015), it was also recorded in Mongolia (Hayford 2005). However, the species may have a Holarctic distribution, provided that its presence in North America can be confirmed (Sealock and Ferrington 2008). Baranov et al. (2024) found a species resembling P. schlienzi in Namibia, but it is possible that the specimen belongs to a yet undescribed species of Psectrocladius related to P. schlienzi.

Habitat

Different types of stagnant waters from lakes to pools. For example, the only record from the Carpathians comes from a shallow pond in an exploited part of an alkaline fen (Bitušík and Illéšová 1998). Its occurrence in slowly flowing waters is exceptional (de Beauvesère-Storm and Tempelman 2009).

Remarks

The records are scattered across Europe, and it seems that the species is not abundant anywhere (Moller Pillot 2013).

Pseudorthocladius (Pseudorthocladius) berthelemyi Moubayed, 1990

Material examined

11 pupal exuvia, Rika River (S6), 7 May 2018.

Distribution

Palaearctic. Austria, Bulgaria, Corsica, France, Germany, Portugal, Slovakia, Spain, Turkey, and Morocco (Ashe and O’Connor 2012).

Habitat

Mountain streams and rivers with stony bottoms. The species is rheophilic, cold-stenothermal with high demand for dissolved oxygen (Martínez et al 1995; Moubayed-Breil et al. 2012). It can also inhabit hygropetric sites (Moubayed-Breil 2008).

Remarks

The species is considered a Mediterranean element (Moubayed-Breil 2008) with an originally circum-mediterranean distribution (Laville and Langton 2002). The extra-Mediterranean occurrence in more northerly countries indicates its relict character (see comments to Cricotopus beckeri).

Subfamily Chironominae

Tribe Chironomini

Paracladopelma mikianum (Goetghebuer, 1937)

Material examined

2 pupal exuvia, Rika River (S6), 7 May 2018; 1 pupal exuvium, Tisa River (S12), 8 May 2018.

Distribution

Palaearctic. The species was recorded only from a few countries in Europe (e.g., Spain, Hungary, Slovakia, Portugal, France, Germany, Romania) and North Africa (Morocco, Lebanon).

Habitat

It is a rheophilic species inhabiting fast-flowing streams and rivers (Moller Pillot 2009). Although Calle-Martínez and Casas (2006) listed the species in a chironomid community usually associated with low-temperature or torrential mountain streams, our finding from Tisa River and other records from large lowland rivers (Gandouin et al. 2006; Klink 2010) indicate that the species does not have as strict cold temperature preferences as though previously (Ringe 1974).

Remarks

Laville and Vinçon (1986) considered the species to be a Mediterranean-Palaearctic element whose northern limit is located in the Pyrenees, the Alps, and the Carpathians.

Tribe Tanytarsini

Neozavrelia Goetghebuer, 1941

Material examined

11 pupal exuvia, Zhdenivka River (S1), 5 May 2018; 17 pupal exuvia, Latoricja River (S4), 5 May 2018; 1 pupal exuvium, Rika River (S6), 7 May 2018; 18 pupal exuvia Rika River (S8), 7 May 2018; 19 pupal exuvium, Rypenka River (S7), 7 May 2018; 1 pupal exuvium, channel at Gluhana peat bog (S5), 7 May 2018.

Distribution

Species-rich genus (38 valid species, de Jong 2016) with a worldwide distribution except for Africa and Neotropics (Epler et al. 2013). Five species have been recorded in Europe, three of which are reliably confirmed in the Carpathians: N. improvisa Fittkau, 1954, N. luteola Goetghebuer, 1941 (Giłka 2007), and N. cuneipennis (Edwards, 1929) (Tatole 2023).

Habitat

Larvae of Neozavrelia inhabit streams, rivers, lakes, and ponds in peat bogs; they are also known from hygropetric sites, and one species lives in a hot spring (Epler et al. 2013).

Remarks

Except for N. cuneipennis (= N. longappendiculata Albu, 1980), the morphological characteristics of the pupae do not yet allow for distinguishing the European species (Langton and Visser 2003). The morphotype Neozavrelia Pe1 Langton, 1991 includes four species: N. bernensis Reiss, 1968, N. fuldensis Fittkau, 1954, N. improvisa, and N. luteola.

Family Chaoboridae

Chaoborus (Chaoborus) obscuripes (van der Wulp, 1859)

Material examined

1 larva, Lake Geryshaska (L7), 15 September 2021.

Distribution

Palaearctic. The species is widespread mainly in Northern and Western Europe, but also in Poland and the European part of Russia (Borkent 1981; de Jong 2016).

Habitat

Small, shallow nutrient-poor, meso- and polyhumic ponds with pH 4.5–5.5 (Nilssen 1974; Joniak and Domek 2006; Kuper and Verberk 2011), often fishless. Larger larvae with darker pigmentation are more sensitive to visually dependent predators (Stenson 1981).

Remarks

The species seems to occur sporadically and mostly in small numbers (Borkent 1981), which is probably related to its ecological requirements for water chemistry and the absence of fish.

The first annotated checklist of Ukrainian Chironomidae consists of 302 species (Baranov 2011a). However, this list requires revision because it contains invalid species identified solely on the basis of larvae using outdated identification keys. In recent decades, the study of taxonomy, ecology, and biogeography of chironomids in Ukraine has intensified (Baranov 2011b, 2013, 2014; Baranov and Przhiboro 2014; Baranov and Ferrington 2013; Moubayed-Breil and Baranov 2018; Didenko et al. 2021). Our survey revealed a significant gap in the taxonomic knowledge of Ukrainian chironomids. The high number of new records suggests that the chironomid fauna, especially from flowing waters is far from being fully discovered. Undoubtedly, it is necessary to continue the study of the chironomid fauna of the Eastern Carpathians. Particularly, the collection of the pupal exuvia could be a very useful tool in studying species richness, ecology, and distribution, but also for water quality assessment and monitoring purposes.

Acknowledgements

We gratefully acknowledge the support of the Carpathian Biosphere Reserve Administration. We are indebted to Mykola M. Voloshchuk, Peter Turis, and Milan Valachovič for providing field collection assistance. We are grateful to the reviewer and subject editor for their valuable comments to the previous version of the manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The study of mountain lakes in the Carpathians was supported by the Slovak Research and De-velopment Agency (project No. APVV-20-0358) and Grant Agency VEGA (project No. 1/0400/21).

Author contributions

PB identified Chironomidae pupal exuviae and adults and wrote the text, MN and MS collected the data, prepared the map, tables and wrote part of the text, LH wrote part of the text.

Author ORCIDs

Milan Novikmec https://orcid.org/0000-0002-5192-4575

Marek Svitok https://orcid.org/0000-0003-2710-8102

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

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  • Tolonen KT, Brodersen KP, Kleisborg TA, Holmgren K, Dahlberg M, Hamerlik L, Hämäläinen H (2012) Phantom midge-based models for inferring past fish abundances. Journal of Paleolimnology 47(4): 531–547. https://doi.org/10.1007/s10933-012-9579-4
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  • Vallenduuk HJ, Moller Pillot HKM (2007) Chironomidae larvae of the Netherlands and adjacent lowlands. General ecology and Tanypodinae. KNNV Publishing, Zeist, 144 pp.

Supplementary material

Supplementary material 1 

Checklist of chironomid and chaoborid taxa recorded in studied streams and lakes of the Ukrainian part of the Eastern Carpathians

Peter Bitušík, Milan Novikmec, Marek Svitok, Ladislav Hamerlík

Data type: xlsx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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