New data on the distribution and ecology of Camponotus oertzeni Forel, 1889 (Hymenoptera: Formicidae) in Eastern Europe and the Caucasus

Том 21. Вып. 2 Vol. 21. Iss. 2 CAUCASIAN ENTOMOLOGICAL BULLETIN Ростов-на-Дону 2025 РОССИЙСКАЯ АКАДЕМИЯ НАУК Южный научный центр RUSSIAN ACADEMY OF SCIENCES Southern Scientific Centre Кавказский энтомологический бюллетень 21(2): 221–226 Caucasian Entomological Bulletin 2025 New data on the distribution and ecology of Camponotus oertzeni Forel, 1889 (Hymenoptera: Formicidae) in Eastern Europe and the Caucasus © D.M. Shevchenko1, 2, Z.M. Yusupov3, C. Karaman4, K. Kiran4, D.A. Dubovikoff5 1D.I. Ivanovsky Academy of Biology and Biotechnology of Southern Federal University, Stachki Avenue, 194/1, Rostov-on-Don 344090 Russia. E-mail: [email protected] 2Federal Research Centre the Southern Scientific Centre of the Russian Academy of Sciences, Chekhov Street, 41, Rostov-on-Don 344006 Russia 3Tembotov Institute of Ecology of Mountain Territories of the Russian Academy of Sciences, Inessa Armand Street, 37a, Nalchik 360051 Russia. E-mail: yz[email protected] 4Department of Biology, Faculty of Science, Trakya University, Edirne 22030 Türkiye. E-mail: [email protected], [email protected] 5Department of Applied Ecology, Faculty of Biology, 16th line of Vasil’evskiy Island, 29, Saint Petersburg 199178 Russia. Email: [email protected]u Abstract. New data on the distribution and ecology of Camponotus oertzeni Forel, 1889 (Hymenoptera: Formicidae) are presented. The species is recorded for Russia, Ukraine, Armenia, Abkhazia and South Ossetia for the first time. In the North Caucasus (Russia), the species occupies a wide range of habitats, from humid beech-hornbeam forests in Krasnodar Region and the Republic of Adygea to moderately dry valleys in the Kabardino-Balkarian Republic. Considerable colour variation was observed among worker caste. While typical southern populations representatives of C. oertzeni exhibit yellowish-red to reddish colouration of the head, mesosoma, and legs, northern populations from the Caucasus and Ukraine are distinctly darker, with almost black heads and mesosomas, and only slightly lighter brown to yellowish legs. This trend may reflect either genetic differentiation or environmental influence. Morphological differences from the related C. aethiops (Latreille, 1798) are discussed. Key words: Camponotus oertzeni, Tanaemyrmex, Eastern Europe, colour polymorphism, population differences. Новые данные о распространении и экологии Camponotus oertzeni Forel, 1889 (Hymenoptera: Formicidae) в Восточной Европе и на Кавказе © Д.М. Шевченко1, 2, З.М. Юсупов3, К. Караман4, К. Киран4, Д.А. Дубовиков5 1Академия биологии и биотехнологии имени Д.И. Ивановского Южного федерального университета, пр. Стачки, 194/1, Ростов-на-Дону 344090 Россия. E-mail: [email protected] 2Федеральный исследовательский центр Южный научный центр Российской академии наук, пр.Чехова, 41, Ростов-на-Дону 344006 Россия 3Институт экологии горных территорий имени А.К.Темботова Российской академии наук, ул. Инессы Арманд, 37a, Нальчик 360051 Россия. E-mail: [email protected] 4Кафедра биологии, факультет естественных наук, Фракийский университет, Эдирне 22030 Турция. E-mail: [email protected], [email protected] 5Кафедра прикладной экологии, биологический факультет, Санкт-Петербургский государственный университет, 16-я линия Васильевского острова, 29, Санкт-Петербург 199178 Россия. E-mail: [email protected] Резюме. Приведены новые данные о распространении и экологии Camponotus oertzeni Forel, 1889 (Hymenoptera: Formicidae). Вид впервые указан для России, Украины, Армении, Абхазии и Южной Осетии. На Северном Кавказе (Россия) вид населяет широкий спектр местообитаний– от влажных буково-грабовых лесов на территории Краснодарского края и Республики Адыгея до сухих котловин в Кабардино-Балкарской Республике. Среди рабочих особей наблюдались значительные различия в окраске. В то время как типичные представители южных популяций C.oertzeni имеют от желтовато-красной до красной окраску головы, мезосомы и конечностей, северные популяции, обнаруженные на Кавказе и на Украине, явно темнее, с почти черной окраской головы и мезосомы и лишь слегка более светлыми, от коричневого до желтоватого цвета, ногами. Эта тенденция может отражать либо генетическую дифференциацию, либо влияние факторов среды. Обсуждаются морфологические различия с родственным видом C.aethiops (Latreille, 1798). Ключевые слова: Camponotus oertzeni, Tanaemyrmex, Восточная Европа, цветовой полиморфизм, популяционные отличия. Introduction The ant fauna (Hymenoptera: Formicidae) is relatively well studied in Europe. However, detailed studies have led to the discovery of previously unknown species in Eastern Europe and the North Caucasus [Yusupov, 2009, 2014; Radchenko, Yusupov, 2012, 2017; Radchenko et al., 2016; Shevchenko, 2024, etc.]. Camponotus oertzeni Forel, 1889 is a member of the subgenus Tanaemyrmex Ashmead, 1905. According to Borowiec and Salata [2022], C. oertzeni belongs to the complex of large species with setose gena, pubescence of DOI: https://doi.org/10.5281/zenodo.17465240 Research Article / Научная статья antennal scapi and posterior tibiae lacking suberect hairs. This complex also includes Camponotus aethiops (Latreille, 1798), C.jaliensis Dalla Torre, 1893 and C.sannini Tohmé et Tohmé, 2000. Camponotus oertzeni is most similar to C. aethiops, which is widespread in Europe and Central Asia. Previously, C. oertzeni was known from Slovenia, Croatia, Serbia, Montenegro, Northern Macedonia, Bulgaria, mainland Greece, the Aegean Islands, Crete, the Cyclades, the Dodecanese, the Ionian Islands, Türkiye, Azerbaijan and Iran [Forel, 1889; Emery, 1908, 1915; Finzi, 1928, 1939; Menozzi, 1936; Agosti, Collingwood, 1987; Radchenko, 1997; Borowiec, 2014; Borowiec, Salata, 2012, 2017a, b, 2018a, b, 2022; Bračko et al., 2016; Karaman, Kiran, 2022; Lapeva-Gjonova, Antonova, 2022]. Through the examination of the collections of the Zoological Museum of Moscow State University (ZMMU, Moscow, Russia) and the Zoological Institute of the Russian Academy of Sciences (ZISP, Saint Petersburg, Russia), as well as our own collected material, we record C.oertzeni for Russia (Krasnodar Region, the Republic of Adygea, Stavropol Region, the Kabardino-Balkarian Republic, the Republic of Dagestan), Ukraine (Kiev and Vinnitsa regions), Armenia (Gegharkunik Region), Abkhazia and South Ossetia for the first time. Ecological preferences in the North Caucasus and variability in worker caste colouration are discussed. Material and methods Most of the specimens studied were found in the collections of ZMMU and ZISP. The majority of specimens were labeled as C.aethiops (C.maculatus aethiops) or had only a geographic label. Material was also collected by the first and second authors in the period from 2005 to 2024, on the territory of Russia (Republic of Adygea, Republic of Dagestan and Kabardino-Balkarian Republic, Krasnodar and Stavropol regions), Armenia and Abkhazia. The material from South Ossetia was collected by Yu.Е.Komarov. Hand collection and nest excavation were the main methods used. Ants were also collected using standard pitfall traps. The material was preserved in 70%ethanol. The photos were taken using a Leica M205C microscope at the Research Park of StPetersburg State University (“Resource Centre Figs 1–4. Camponotus oertzeni worker, habitus, details. 1, 3 – worker: 1 – lateral view, 3 – dorsal view; 2 – head, frontal view; 4 – the transverse row of short setae on the posterior margin of the first gastral tergite (white line indicates the end of the first tergite, the arrows point to the ends of the setae). Рис. 1–4. Camponotus oertzeni, рабочий, габитус, детали строения. 1, 3 – рабочий: 1 – латерально, 3 – дорсально; 2 – голова, фронтально; 4 – поперечный ряд прилежащих волосков на заднем крае первого тергита брюшка (белая линия обозначает заднюю границу первого тергита, стрелки указывают на вершины прилежащих волосков). 222 D.M. Shevchenko, Z.M. Yusupov, C. Karaman, K. Kiran, D.A. Dubovikoff for Microscopy and Microanalysis”). Specimens are temporarily kept in the author’s collections (PCSH – D.M. Shevchenko’s collection, PCZY – Z.M. Yusupov’s collection) and will be deposited in ZISP, ZMMU and the Tembotov Institute of Ecology of Mountain Territories of the Russian Academy of Sciences (Nalchik, Russia). The following abbreviations are given in the “Material” section for ant castes: “w”for worker, “g”for gyne and “m”for male. Camponotus oertzeni Forel, 1889 (Figs 1–10) Material. Russia. Krasnodar Region: Sochi District: 1 w (ZMMU), “326. Abrau, N-W Caucas. 26.07.[19]21 Arnoldi”; 3 w (ZMMU), “Sochi, 20.04.1916, №660 N Koltzov”; 1 w (ZMMU), “642. Bez. Sotschi 08.1926 W. Alpatow”; 2 w (ZMMU), “Сочи (Кавказ), 14.07.1927, И. Гудим” [Sochi (Caucasus), I. Gudim]; 2 w (ZMMU), “5256 Sotschi 07.[19]31, V. Lutschnik”; 3 w (ZMMU), “5258 Sotschi 08.[19]31, V. Lutschnik”; 1 w (ZMMU), Matsesta, 6.05.1931 (Y.I. Kostylev); Tuapse District: 2 w (PCZY), Gorskoevill.env., edge of a broad-leaved forest, forb-couch grass community, near a stream, 44°21′36.0′′N/ 38°43′12.0′′E, 108m, 14.06.2008 (Z.M.Khanov). Republic of Adygea, Maikop District: 4w (ZMMU), “6097 Шунтук, Майкоп, Сев. Кавказ, Арнольди” [Shuntuk, Maykop, North Caucasus, Arnoldi]; 12w (PCZY), Kamennomostskiy settlementenv., oak forest with rhododendron, nest in soil, workers collected in the leaf litter and in the thicket of ferns, 44°17′00.0′′N/ 40°09′00.0′′E, 652m, 29.07.2011 (Z.M. Yusupov); 5 w (PCSH), Nickel settlement env., Granitnoe Gorge, on a rocky slope in the floodplain of the Belaya River, 44°09′48.23′′N / 40°08′28.64′′E, 520 m, 19.06.2024 (D.M. Shevchenko); 1 w (PCSH), Ust’-Sakhray settlementenv., Sakhray waterfall, beech and hornbeam forest, on rocky granite outcrops, 44°09′04.67′′N/ 40°17′43.31′′E, 600m, 24.06.2024 (D.M. Shevchenko); 2 g, 3 w (PCSH), Dakhovskaya Cossack vill. env., oak forest on the stony slope of Dakhovskaya kuesta, 44°14′31.96′′N / 40°10′23.02′′E, 740 m, 22.06.2023 (D.M. Shevchenko). Stavropol Region: 25w (PCZY), Pyatigorsk, MashukMt., mixed-grass meadow on the edge Figs 5–10. Camponotus oertzeni, habitus, details. 5–7 – gyne: 5 – lateral view, 6 – dorsal view, 7 – head, frontal view; 8–10 – male: 8 – head, frontal view, 9 – habitus, lateral view, 10– habitus, dorsal view. Рис. 5–10. Camponotus oertzeni, габитус, детали строения. 5–7 – самка: 5 – габитус, вид сбоку, 6 – габитус, вид сверху, 7 – голова, вид спереди; 8–10 – самец: 8 – голова, вид спереди, 9 – габитус, вид сбоку, 10– габитус, вид сверху. New data on the distribution and ecology 223 of broad-leaved forest, nest in soil, 44°02′58.0′′N / 43°04′45.8′′E, 658 m, 1.06.2013 (Z.M. Yusupov). Kabardino-Balkarian Republic, Cherekskiy District: 4 w (PCZY), Verkhnyaya Balkariya vill. env., semiarid valley, SE slope, upland xerophytic vegetation, nest in soil, 43°08′29.7′′N / 43°27′34.7′′E, 1232m, 13.08.2005 (Z.M.Yusupov); 5w (PCZY), the same place, nest in soil, 18.08.2005 (Z.M.Yusupov); 9w (PCZY), the same place, nest in soil, 17.06.2007 (Z.M. Yusupov). Republic of Dagestan, Buynaksk District: 3w (PCZY), 1.5km SE of Dubki vill., oak forest, nest in soil, under the stone, 43°00′18.0′′N/ 46°51′10.8′′E, 610m, 3.05.2019 (Z.M.Yusupov). Armenia. Gegharkunik Region: 4w (PCZY), Sevan Lakeenv., rock outcrops, xerophytic vegetation, on stones, 40°35′43.8′′N / 44°57′55.6′′E, 1920 m, 27.05.2021 (Z.M. Yusupov); Lori Region: 15 w (PCZY), 2 km N Pambak vill., oak forest with large stones, nest in soil, 40°50′29.8′′N / 44°33′48.6′′E, 1319m, 29.05.2021 (Z.M.Yusupov). Abkhazia. Gagra District: 1 w (ZMMU), “Ermolovskoe pr. Gagry 08.08.1933, G. Kostylev” (= Tsandripsh village); 3 w (ZISP), “Гагры, зап.Кавказ, Арнольди 18.06.1933” [Gagra, Western Caucasus, Arnoldi]; 1w (ZMMU), “Абхазия, Гагра, 20.09.1963, В.Ковалёв” [Abkhazia, Gagra, V. Kovalyov]; Gulripsh District: 4 m, 1 w (ZISP), “Ажары у Чхатлы на Кодоре, Сухум. Калишевс. 14.07.05” [Azhara vill. near Chkhatla River on Kodor, Sukhum. Kalishevskiy. 14.07.1905]; 3w (ZISP), same data, but 17.07.1905; 2g, 1w (ZISP), same data, but 18.07.1905; 1g, 2w (ZISP), same data, but 21.07.1905; 6g, 1w (ZISP), “р.Чхалта, увпад. в Кодор, Сухум. о. Калишевc 01.08.05” (Chkhatla River at the confluence with the Kodor River, Sukhumenv. Kalishevskiy 1.08.1905]; 2g, 5w (ZISP), “Лата на Кодоре, Сухум. окр. Калишевс 19.08.05” [Latavill. on Kodor River, Sukhumenv. Kalishevskiy 19.08.1905]; Sukhum District: 3 w (PCZY), Sukhum, agrocenosis, nest in soil, on the area with sparse herbaceous vegetation, 42°58′25.0′′N/ 41°04′02.7′′E, 10m, 12.07.2006 (Z.M.Yusupov); 6w (PCZY), in the same place, in the grass, 12.07.2006 (Z.M.Yusupov). South Ossetia. Tskhinval District: 4w (PCZY), 1km NW Prisvill., oak forest, 42°13′53.9′′N/ 43°58′57.3′′E, 850m, 10.05.2013 (Yu.E.Komarov); 1w(PCZY), in the same place, pine forest, 13.07.2013 (Yu.E.Komarov);35w (PCZY), in the same place, oak forest, 13.07.2013 (Yu.E. Komarov); 10w(PCZY), oak forest, on the trail, 42°13′34.9′′N/ 44°00′42.0′′E, 870m, 25.05.2013 (Yu.E.Komarov); 19w (PCZY), in the same place, 16.06.2013 (Yu.E. Komarov); 1 w (PCZY), 500 m NW of Tskhinval, 42°13′58.0′′N / 43°57′15.6′′E, 810m, oak forest, 15.06.2013 (Yu.E.Komarov); 9w (PCZY),in the same place, pine forest, 13.07.2013 (Yu.E. Komarov); 1 w (PCZY), inthe same place, oak forest, 13.07.2013 (Yu.E.Komarov); 1w (PCZY), in thesame place, pine forest, 30.08.2013 (Yu.E.Komarov); 12w (PCZY), in the sameplace, oak forest, 30.08.2013 (Yu.E.Komarov); 1w (PCZY), Tskhinval, wasteland behind the former railway station, 42°14ʹ36.2ʺN/ 43°58ʹ02.2ʺE, 800 m, 22.06.2014 (Yu.E. Komarov); 1 w (PCZY), Atsriskhev vill. env., South Ossetian State Reserve, in the grass, 42°17′32.7′′N / 44°13′23.1′′E, 1220m, 20.06.2014 (Yu.E.Komarov); 52w (PCZY), 1km SE Atsriskhevvill., South Ossetian State Reserve, hornbeam forest, under the stone, 42°17′45.4′′N / 44°13′44.8′′E, 1280 m, 21.06.2014 (Yu.E. Komarov); 78 w (PCZY), in the same place, under the stone, 21.06.2014 (Yu.E.Komarov); 27 w (PCZY), Zonkar vill. env., Malo-Liakhvi gorge, S slope, hornbeam forest, 42°17′27.2′′N/ 44°09′31.0′′E, 1372m, 21.06.2014 (Yu.E.Komarov); 2 w (PCZY), Atsriskhevvill.env., Malo-Liakhvi gorge, Sslope, hornbeam forest, pitfall traps, 42°18′10.6′′N / 44°14′32.6′′E, 1260m, 2.10.2014 (Yu.E.Komarov); 1w (PCZY), 2.5km S Belotvill., SEslope, mixed-grass meadow among hornbeam forest, pitfall traps, 42°17′35.9′′N/ 44°07′45.1′′E, 1250m, 27.08.2015 (Yu.E.Komarov); 27w (PCZY), 2km E Atsriskhevvill., Malo-Liakhvi gorge, South Ossetian State Reserve, Sslope, glade among hornbeam forest, 42°18′19.9′′N / 44°15′00.3′′E, 1393 m, 18.05.2016 (Yu.E.Komarov); Znaur District: 1w (PCZY), Verkhniy Bekmarvill.env., clearing among deciduous forest, on plants, 42°14′59.5′′N / 43°49′40.0′′E, 950m, 30.08.2013 (Yu.E.Komarov). Ukraine. Vinnitsa Region: 2w (ZISP), “Цыбулевский лес Подольской губ. 03.04.1911 И. Якубовский №6/77” [Tsibulevskiy forest, Podolsk Province, I.Yakubovskiy]; Kiev Region: 1w (ZMMU), “Канев. Киев. Обл. 17.06.1952г. Кришталь №124” [Kanev, Kiev Region, Kryshtal]. Notes. Camponotus oertzeni is morphologically similar to C.aethiops, which has probably led to misidentifications in the past, as the main distinguishing feature between these two species was thought to be the colouration of the workers. Radchenko [1997] stated that C.oertzeni differs from C.aethiops by the orange-red colouration of the head and mesosoma. More recent studies [Salata, Borowiec, 2018] have reported a wide range of colour variation in both species, indicating that colouration by itself is not a reliable diagnostic character. Instead, the most consistent morphological difference is the length of the transverse row of short setae on the posterior margin of the first abdominal tergite: in C.aethiops, the setae of this row are long, usually reaching or exceeding the posterior margin of the tergite, whereas in C.oertzeni it is short, typically reaching only 1/2 to 2/3 of the tergite margin length (Fig. 4). Nonetheless, Salata and Borowiec [2018] noted that C.aethiops is generally darker, with a predominantly dark brown to almost black body and usually brown to black legs (although, in rare cases, individuals may have a reddish mesosoma and yellowish legs), while C. oertzeni populations commonly exhibit partly pale, yellowish-red to reddish heads, mesosomas, and legs. However, in the North Caucasus and Ukraine, the species shows darker colouration: head, mesosoma and abdomen are almost black, the femora and tibiae are light brown to yellow (Figs1–3). Camponotus oertzeni can also be distinguished from C.aethiops by its ecological preferences. While the latter species prefers dry open habitats, C. oertzeni inhabits wetter habitats like deciduous forests and open, yet moist rocky outcrops in floodplains. In the Republic of Adygea of Russia the species inhabits oak and beech-hornbeam forests, in which it settles in open areas such as clearings, edges and open rocky outcrops (Figs11,12). In the second half of June 2024 gynes of C. oertzeni were often found nesting in limestone crevices and under rocks in moist beech and hornbeam forests after the nuptial flight. In the Kabardino-Balkarian Republic of Russia the species also inhabits moderately dry valleys above the belt of deciduous forests (where it is suddenly not represented), in the belt of mountain xerophytes at an altitude of about 1200m (Fig.13). Discussion A wide colour polymorphism is a well-known phenomenon among various ant genera, including Camponotus Mayr, 1861 [Seifert, 2019; Csösz et al., 2022]. In the paper on the Formica cinerea species-group Seifert [2002] reported that the colour polymorphism in the workers of Formica cinerea Mayr, 1853 has more likely genetic basis than the ecological one, as he found three sites with syntopic occurrence of reddish and dark morphs. The difference in mesosoma colouration in northern populations of C. oertzeni may also be both genetic differences and environmental impact. Interestingly, C. aethiops, which is more typical for semiarid valleys, in the Kabardino-Balkarian Republic of Russia is absent in these semiarid basins. On the territory of the region, it is represented only in the plain part, in the steppe zone. Probably, C.aethiops could not penetrate through the belt of moist broad-leaved forests, while C.oertzeni could and therefore occupy a habitat untypical for the species. Records in southwestern Ukraine and in the North Caucasus significantly extend the northern boundary of the species distribution (Fig.14). The presence of the species in Kiev Region of Ukraine is doubtful and needs to be verified. Since morphological characteristics allow us to clearly classify the specimens as C.oertzeni, an error in the labeling of the material seems possible. 224 D.M. Shevchenko, Z.M. Yusupov, C. Karaman, K. Kiran, D.A. Dubovikoff Acknowledgements The authors are grateful to Bernhard Seifert (Senckenberg Museum of Natural History, Görlitz, Germany) and the second anonymous reviewer for valuable comments and corrections. The study was supported by the state project 125011200139-7 for D.M.Shevchenko. References Agosti D., Collingwood C.A. 1987. 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DOI: 10.1134/S0013873814020195 226 D.M. Shevchenko, Z.M. Yusupov, C. Karaman, K. Kiran, D.A. Dubovikoff Received / Поступила: 24.03.2025 Accepted / Принята: 26.05.2025 Published online / Опубликована онлайн: 28.10.2025