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Research Article

Water bugs (Heteroptera: Gerromorpha, Nepomorpha) of the Caucasus ecoregion

G. M. Berchi1 Institute for Advanced Environmental Research, West University of Timișoara, Timișoara, RomaniaCorrespondence[email protected]
ORCID Iconhttps://orcid.org/0000-0002-9873-0647
ORCID Icon,
D. Copilaș-Ciocianu2 Laboratory of Evolutionary Ecology of Hydrobionts, Nature Research Centre, Vilnius, LithuaniaORCID Iconhttps://orcid.org/0000-0002-6374-2365
ORCID Icon,
P. Kment3 Department of Entomology, National Museum, Prague, Czech RepublicORCID Iconhttps://orcid.org/0000-0002-7026-5691
ORCID Icon &
L. Mumladze4 Institute of Zoology, Ilia State University, Tbilisi, GeorgiaORCID Iconhttps://orcid.org/0000-0002-2172-6973
ORCID Icon
Pages 167-192 | Received 28 Sep 2022, Accepted 24 Jan 2023, Published online: 26 Feb 2023

Abstract

The Caucasian fauna of water bugs (Heteroptera: Gerromorpha, Nepomorpha) is reviewed based on data originating from literature survey, museum and private collections, and extensive field sampling. The diversity of Caucasus ecoregion is quite significant with 77 species and subspecies framed within 25 genera (17 in Nepomorpha, 8 in Gerromorpha), and 13 families (8 in Nepomorpha, 5 in Gerromorpha). Micronecta anatolica anatolica Lindberg, 1922, Sigara iranica Lindberg, 1964, S. samani tigranes Jansson, 1986, Velia kiritshenkoi Tamanini, 1958, and Gerris asper (Fieber, 1860) are reported for the first time from Georgia, M. anatolica anatolica, Notonecta maculata Fabricius, 1794, Mesovelia thermalis Horváth, 1915, and Gerris lacustris (Linnaeus, 1758) represent new records for Armenia, and Microvelia macani Brown, 1953 is new for Azerbaijan. A chorological and similarity analysis highlighted the connections with and between adjacent countries, with Caucasus ecoregion having the highest similarity of the water bug fauna with Turkey, whereas Iran and Russia (European territory) have the lowest. The largest number of shared species/subspecies was found between the Caucasus and Turkey (64 taxa), while the lowest was between Iran and Russia (40 taxa). Our results indicate that the Caucasus represents a composite of various faunal elements of different origin, ranging from Caucasian endemics to Afrotropico-Indo-Mediterranean and Holarctic.

Introduction

The water bugs (Hemiptera: Heteroptera: Gerromorpha, Nepomorpha) are worldwide spread, excepting polar regions, and occur in a remarkable broad range of aquatic ecosystems (Polhemus & Polhemus Citation2008; Schuh & Weirauch Citation2020). They comprise two relatively small, but very diverse groups of insects. The Gerromorpha, also known as semiaquatic bugs, are mostly living on the water surface of freshwater habitats, but some are adapted to moist terrestrial habitats or even marine environments (Andersen Citation1982; Spence & Andersen Citation1994). The Nepomorpha, or aquatic bugs, occur particularly in freshwater habitats, but some species are known to live in dry terrestrial or marine environments as well (Schuh & Slater Citation1995; Schuh & Weirauch Citation2020).

The world’s gerromorphans are currently framed in eight, whereas nepomorphans in 13 families (Nieser Citation1999; Damgaard Citation2008b; Damgaard & Zettel Citation2014; Wang et al. Citation2021). The highest species richness is found in the Neotropical, Oriental, and Afrotropical regions, whereas in the Palaearctic the diversity is significantly lower. Overall, 17 families, 85 genera, and about 525 species and subspecies are known from the latter (Aukema & Riger Citation1995; Polhemus & Polhemus Citation2008; Kment et al. Citation2016). Although the Palaearctic water bug fauna is considered to be well documented in comparison with that of other regions, there are still large areas which lack adequate surveys (i.e., southeastern Europe, the Caucasus, and a large portion of the Asian territories) (see e.g., Aukema & Rieger Citation1995; Fent et al. Citation2011; Linnavuori et al. Citation2011; Ghahari et al. Citation2013; Kment et al. Citation2016; Csabai et al. Citation2017; Berchi et al. Citation2018a, Citation2018b; Cianferoni Citation2019).

The Caucasus ecoregion is a biodiversity crossroads which completely or partially extends over the territories of Georgia, Azerbaijan, Armenia, Russia, Turkey, and Iran (). While being acknowledged as one of the richest biodiversity hotspots on Earth (Myers et al. Citation2000), and one of the Pleistocene refugia and post-glacial colonization source for northern Europe (Hewitt Citation2000), the Caucasus has supposedly received comparatively less scientific attention, mainly due to the political context of the last decades (Mumladze et al. Citation2020).

Figure 1. Map of the Caucasus ecoregion and distribution of the sampling sites.

Figure 1. Map of the Caucasus ecoregion and distribution of the sampling sites.

The most significant work on the water bugs of Caucasus is that of Kiritshenko (Citation1918), which covers Armenia, Azerbaijan, Georgia, and areas of the south European Russia and northeastern Turkey. Additional studies for some smaller areas are listed by Aukema and Rieger (Citation1995). More recently, Linnavuori & Hosseini (Citation2000), Kanyukova et al. (Citation2002), Prokin et al. (Citation2008, Citation2009), Neimorovets (Citation2010), Shapovalov et al. (Citation2012, Citation2014, Citation2018, Citation2019) improved the knowledge in some areas, whereas the works of Kanyukova (Citation2006), Fent et al. (Citation2011), and Ghahari et al. (Citation2013) provide comprehensive lists and distributions by provinces/republics for the faunas of Russia, Turkey, and Iran respectively. Although, no comprehensive survey has been carried out on the Caucasian ecoregion as a whole.

The main aim of this paper is to improve the knowledge of the Caucasus ecoregion’s water bug taxonomic diversity. Therefore, we provide an annotated catalogue of the gerromorphans and nepomorphans from this region based on a review of the published records, data from museum and private collections, and extensive sampling. A chorological analysis is performed in order to evaluate the Caucasus’ composition of faunal elements. The faunal connections with and between adjacent countries are compared and discussed.

Materials and methods

The study is based on data originating from literature survey, museum and private collections, and our own extensive field sampling. Most of the newly obtained material was collected in the period 2013–2020, and is preserved in GMB’s private collection (see below), unless otherwise stated in the material examined; the collectors, which were not repeated in the material examined, are Gavril Marius Berchi, Denis Copilaș-Ciocianu, and Levan Mumladze. Specimens were collected from 116 localities in Armenia, Azerbaijan, and Georgia (), using standard water nets (0.5 mm mesh size) and stored in 96% ethanol. Another portion of material comes from the Hungarian Natural History Museum, Budapest, Hungary, National Museum, Prague, Czech Republic (including the collection of the late Miroslav Papáček), and Michal Straka’s personal collection, Hustopeče, Czech Republic, and was identified by Petr Kment unless stated otherwise.

Table I. List of sampling sites with their habitats, geographic coordinates, elevation and collecting dates.

Abbreviations of the depositories:

GBTR = Gavril Marius Berchi personal collection, Timișoara, Romania

HNHM = Hungarian Natural History Museum, Budapest, Hungary

LMTG = Levan Mumladze personal collection, Tbilisi, Georgia

MSHC = Michal Straka personal collection, Hustopeče, Czech Republic

NMPC = National Museum, Prague, Czech Republic

Abbreviations of the wing morphs: ap = apterous, br = brachypterous, ma = macropterous, mi = micropterous.

The system and nomenclature mostly follows Aukema & Rieger (Citation1995) and Aukema et al. (Citation2013). Nieser (Citation1999, Citation2002) raised Micronectinae that belonged to Corixidae to family rank, and therefore, we adopt this change which has been recently confirmed through a phylogenomic analysis by Wang et al. (Citation2021). We used the following identification keys: Tamanini (Citation1947, Citation1958, Citation1979), Poisson (Citation1957), Jansson (Citation1986b), Andersen (Citation1990, Citation1993), Kanyukova (Citation1973a, Citation1973b, Citation1974, Citation1982), and Berchi et al. (Citation2016).

The geographical coordinates of the newly collected material are given in the World Geodetic System (WGS84) (see ). The Caucasus ecoregion is defined as proposed by Olson et al. (Citation2001). The map () was created with ArcGIS 10 (ESRI Citation2011). When referring to the published records from Iran, Russia, and Turkey, we only considered those administrative units (provinces, republics) completely or partially overlapping the ecoregion.

The chorotypes are assigned according to Vigna Taglianti et al. (Citation1993, Citation1999) and they mostly derive from the distributional data provided by Aukema & Rieger (Citation1995) and Aukema et al. (Citation2013). The Turano-Caucasian (TUC), Balkano-Pontic (BAP), Turano-Arabian (TUA), S-Asiatic (SOA), and S-Palaearctic (SPA) chorotypes were newly created.

The material examined is arranged alphabetically by country and locality; each location of the newly collected material has a code which corresponds to the specific additional information (habitat, geographical coordinates, elevation, and the date of sampling) presented in . Information about the material in HNHM, MSHC, and NMPC is presented as almost as written on the labels and is not included in .

In order to analyse community similarity among neighboring countries we used the Sørensen index (Magurran Citation2004).

Results

In total, 1448 newly collected specimens were identified at species/subspecies level, and 196 specimens from HNHM, MSHC and NMPC were identified or revised. All in all, they belong to 43 taxa. Five of them [Micronecta anatolica anatolica Lindberg, 1922, Sigara iranica Lindberg, 1964, S. samani tigranes Jansson, 1986, Velia kiritshenkoi Tamanini, 1958, and Gerris asper (Fieber, 1860)] are reported for the first time from Georgia, four taxa [M. anatolica anatolica, Notonecta maculata Fabricius, 1794, Mesovelia thermalis Horváth, 1915, and Gerris lacustris (Linnaeus, 1758)] represent new elements for the Armenian fauna, whereas one species [Microvelia (Picaultia) macani Brown, 1953] is new for Azerbaijan.

Altogether with the literature review, we find that up to 77 species/subspecies (51 in Nepomorpha, 26 in Gerromorpha) occur in the Caucasus ecoregion. They are framed within 25 genera (17 in Nepomorpha, 8 in Gerromorpha), and 13 families (8 in Nepomorpha, 5 in Gerromorpha) ().

Table II. Diversity of the aquatic and semiaquatic Heteroptera in the Caucasus ecoregion.

Some possible taxonomic issues were identified and briefly discussed [see below the Remarks sections for Aphelocheirus (Aphelocheirus) kolenatii Kiritshenko, 1925, and Notonecta (Notonecta) glauca poissoni Hungerford, 1934].

The annotated list of taxa is presented below.

Taxonomic accounts

Order Hemiptera Linnaeus, 1758

Suborder Heteroptera Latreille, 1810

Infraorder Nepomorpha Popov, 1968

Family Nepidae Latreille, 1802

Subfamily Nepinae Latreille, 1802

Genus Nepa Linnaeus, 1758

Nepa cinerea Linnaeus, 1758

Discussion

From a Palaearctic perspective, the Caucasus ecoregion comprises 8 of 11 nepomorphan, and 5 of 6 gerromorphan families. The nepomorphan Belostomatidae, Gelastocoridae, Helotrephidae and gerromorphan marine Hermatobatidae are not reported from the Caucasian fauna, and except the first family, the occurrence of all the others can be excluded due to ecological and geographical reasons. The largest two families of water bugs in the Palaearctic region, Corixidae and Gerridae with 14 and 20 genera, and more than 120 and 100 species and subspecies, respectively (Aukema & Rieger Citation1995; Aukema et al. Citation2013), have also the highest richness in the Caucasus, both in terms of genera (Corixidae: 8; Gerridae: 3) and species and subspecies (Corixidae: 29; Gerridae: 12).

The chorological analysis () revealed 24 chorotypes, underlining the diversity and various faunal influences in the Caucasus. Four species (i.e., Callicorixa raddei, Gerris kiritshenkoi, Hebrus liliimacula, and Monticorixa armeniaca) are Caucasian endemics. Other seven taxa are endemic to the Ponto-Caucasian area (i.e., Aphelocheirus kolenatii, Gerris caucasicus, Notonecta glauca poissoni, Sigara daghestanica, S. iranica, S. stagnalis pontica and Velia kiritshenkoi), whereas three (i.e., Corixa jakowleffi, Hebrus pilipes, and Micronecta mesmini) have a Turano-Caucasian distribution. A few species are Mediterranean extending their distribution to the Caucasus, whereas another small portion are restricted to southwestern Asia.

Figure 2. Chorotypes of the Caucasian fauna. Abbreviations: AIM = Afrotropico-Indo-Mediterranean; ASI = Asiatic; ASE = Asiatic-European; BAP = Balkano-Pontic; CAUC = Caucasian endemic; CAS = Centralasiatic; CAE = Centralasiatic-European; CEM = Centralasiatic-Europeo-Mediterranean; CAM = Centralasiatic-Mediterranean; EME = East-Mediterranean; EUR = European; OLA = Holarctic; MED = Mediterranean; PAL = Palaearctic; POCA = Ponto-Caucasian; SOA = South-Asiatic; SIE = Sibero-European; SWA = Southwest-Asiatic; TUA = Turano-Arabian; TUC = Turano-Caucasian; TUE = Turano-European; TEM = Turano-Europeo-Mediterranean; TUM = Turano-Mediterranean; WPA = West-Palaearctic.

Figure 2. Chorotypes of the Caucasian fauna. Abbreviations: AIM = Afrotropico-Indo-Mediterranean; ASI = Asiatic; ASE = Asiatic-European; BAP = Balkano-Pontic; CAUC = Caucasian endemic; CAS = Centralasiatic; CAE = Centralasiatic-European; CEM = Centralasiatic-Europeo-Mediterranean; CAM = Centralasiatic-Mediterranean; EME = East-Mediterranean; EUR = European; OLA = Holarctic; MED = Mediterranean; PAL = Palaearctic; POCA = Ponto-Caucasian; SOA = South-Asiatic; SIE = Sibero-European; SWA = Southwest-Asiatic; TUA = Turano-Arabian; TUC = Turano-Caucasian; TUE = Turano-European; TEM = Turano-Europeo-Mediterranean; TUM = Turano-Mediterranean; WPA = West-Palaearctic.

The Palaearctic, West-Palaearctic, and Sibero-European chorotypes comprise the highest number of taxa (9). Eleven chorotypes are represented by only one taxon each. The majority of taxa are widely spread throughout parts of Asia, Europe, northern Africa or even the entire Palaearctic. One taxon (i.e., Anisops sardeus sardeus) has an Afrotropico-Indo-Mediterranean distribution, whereas another one (i.e., Limnoporus rufoscutellatus) is Holarctic.

Territories of the neighboring countries (Turkey: ~785,000 km2, Iran: ~1,745,000 km2, Russia (European part): ~3,969,000 km2) are significantly larger than the Caucasus ecoregion (~580,000 km2). Therefore, we analyzed the total and shared number of species/subspecies, and compared the similarity of the water bug taxa with and between the ecoregion and these countries.

Our results indicate an overall number of 77 species and subspecies in the Caucasus ecoregion. Eighty three taxa are listed from Turkey by Fent et al. (Citation2011); however, Velia rhadamantha rhadamantha Hoberlandt, 1941 is endemic to Crete (see Csabai et al. Citation2017; Berchi et al. Citation2018b), Hebrus pusillus pusillus has been recently excluded by Kment et al. (Citation2016), whereas two more species have been recorded afterwards: Hydrometra gracilenta (Banbal & Fent Citation2016; Çerçi & Koçak Citation2016), and V. mancinii mancinii (Berchi et al. Citation2018b). Yet, the presence of other seven species is considered doubtful. From Iran, 82 species have been reported by Ghahari et al. (Citation2013), but the occurrence of some is questionable (see e.g., Kment et al. Citation2016), whereas some certainly does not occur in this country i.e., Velia caprai caprai Tamanini, 1947, Gerris gibbifer Schummel, 1832, which for e.g., in eastern Europe are restricted to north of the Mediterranean peninsulas and Caucasus (see e.g., Fent et al. Citation2011; Küçükbasmacı & Kıyak Citation2015; Berchi et al. Citation2018a, Citation2018b), G. lateralis Schummel, 1832, which is also a Nordic species, and H. pusillus pusillus, which has been recently excluded by Kment et al. (Citation2016). Two other taxa, Ranatra parvipes vicina Signoret, 1880 and Sigara marginata (Fieber, 1851) have been listed by Andersen (Citation1995e) and omitted by Ghahari et al. (Citation2013), whereas the occurrence of H. pusillus arabicus Linnavuori, 1971 has been recently confirmed by Kment et al. (Citation2016). According to Aukema & Rieger (Citation1995), Kanyukova et al. (Citation2002), Kanyukova (Citation2006), and Shapovalov et al. (Citation2018), 83 species are known from the European territory of Russia.

shows the Sørensen’s similarity indices with and between the ecoregion and the neighboring countries. The Caucasus ecoregion and Turkey had the highest similarity index of 0.800, followed by Caucasus ecoregion and Russia, and Turkey and Iran, which had 0.763 and 0.695 respectively. Russia and Iran had the least similarity index of 0.488. The largest number of shared species/subspecies is between the Caucasus and Turkey (64 taxa) and Caucasus and Russia (61 taxa), whereas the lowest was between Iran and Russia (40 taxa) ().

Table III. Sørensen similarity index of the water bug species of Caucasus ecoregion and neighboring countries.

Figure 3. Diagram of the total and shared number of taxa in and between the Caucasus ecoregion and neighboring countries. The total number is presented under the country/region name, and numbers within overlapping areas represent the shared number of taxa.

Figure 3. Diagram of the total and shared number of taxa in and between the Caucasus ecoregion and neighboring countries. The total number is presented under the country/region name, and numbers within overlapping areas represent the shared number of taxa.

Although the overall number of taxa occurring in the ecoregion is significant, some additional species may be expected as well i.e., Lethocerus patruelis (Stål, 1854) (Belostomatidae), Micronecta wui alkani Hoberlandt, 1952 (Micronectidae), Sigara albiventris (Horváth, 1911), S. kervillei (Poisson, 1927), Sigara mayri (Fieber, 1860) (all Corixidae), Naucoris maculatus maculatus Fabricius, 1798, (Naucoridae), Anisops debilis perplexus Poisson, 1929 (Notonectidae), Microvelia hozari Hoberlandt, 1952, Velia mariae Tamanini, 1971, V. mancinii lyciae Tamanini, 1955 (all Veliidae), or Gerris kabaishanus Linnavuori, 1998 (Gerridae). Many of them are only known from a handful of localities from Iran, Iraq or Turkey.

Conclusion

The significant number of taxa concentrated on a relatively small area acknowledge the expected richness, whereas the relatively similar number of shared species with the adjacent countries emphasize that the Caucasus ecoregion represents a water bug fauna crossroad. This is also highlighted by the variety of chorotypes which represents a composite of various faunal elements of diverse biogeographic origin. The overall number of species reported in the Caucasus ecoregion is large, but definitely incomplete, and notwithstanding this reason, it is likely that several additional taxa will be discovered. Some possible taxonomic issues were identified and are awaiting to be solved. We hope that our work will encourage further studies in the area, in order to create a solid base for other research.

Acknowledgements

We thank Meri Arzumanyan (Yerevan State University, Armenia) for her help in the field work;Maka Bitsadze and Giorgi Beruchashvili (both WWF Caucasus Programme Office, Georgia) for sharing with us the shapefile of the Caucasus ecoregion; Janeta Shubitidze, Ani Bikashvili, Bella Japoshvili, Ekaterine Pankvelashvili, Giorgi Nozadze for helping in collecting and sorting materials at the Institute of Zoology of Ilia State University. We are also obliged to Peter Manko and Jozef Oboňa (both University of Prešov, Slovakia) for providing their samples collected in Georgia and Azerbaijan and permission to use the data in this paper.

LM’s work was partially supported by the Shota Rustaveli National Science Foundation grant (FR217-086) and CaBOL (Caucasus Barcode of Life) Project funded by Federal Ministry of Education and Research of Georgia under the grant number 01DK20014A. The contribution of PK received financial support from the Ministry of Culture of the Czech Republic (DKRVO 2019–2023/5.I.d, National Museum, 00023272).

Disclosure statement

No potential conflict of interest was reported by the author(s).

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