Smirnov_06_2022.indd UDC 597.6(477.85) THE CASES OF ABNORMAL AMPLEXUS IN ANURA ON THE TERRITORY OF THE CHERNIVTSI REGION, UKRAINE N. A. Smirnov Chernivtsi Regional Museum, vul. Olha Kobylianska, 28, Chernivtsi, 58002 Ukraine E-mail: nazarsm@ukr.net https://orcid.org/0000-0001-9731-6404 urn:lsid:zoobank.org:pub:0D20DC00-A170-4D75-B030-75B487743B1B Th e Cases of Abnormal Amplexus in Anura on the Territory of the Chernivtsi Region, Ukraine. Smirnov, N. A. — Th e paper contains information about 21 cases of abnormal amplexus among anurans (Amphibia) detected on the territory of the Chernivtsi Region, Ukraine. Multiple amplexus, consisting of one female and two or more males, was registered in three species: Bombina variegata, Bufo bufo, Rana temporaria. Th e author described interspecifi c amplexus in six cases. Five of them were among anurans (Bufo bufo male × Pelophylax lessonae male, Rana dalmatina males × Rana temporaria female or males, Rana temporaria male × Pelophylax ridibundus female) and one case — among Anura and Caudata (Bom- bina variegata male × Lissotriton montandoni female). K e y w o r d s : Amphibia, abnormal spawning, mating behaviour, Lissotriton montandoni . Introduction Amplexus is a form of mating behaviour known in most species of anurans (Duellman & Trueb, 1994). Normally amplexus is a part of mating behaviour when a male wraps its forelegs around a female and fertilizes the eggs that are being laid. Such behaviour enables partners to synchronize allocating reproductive products and increase the chances of successful external fertilization due to the closest approaches of male’s and female’s cloacas. Currently, seven types of amplexus are known among anurans, the most common of which are axillary and inguinal types (Duellman & Trueb, 1994; Willaert et al., 2016). As a rule, amplexus is an interaction of one male with one female of the same species, however diff erent deviations also can take place. For example, there have been cases of multiple amplexus with interaction among one female and several males, among males of the same species or with amphibians of diff erent species, with non-living individuals of its or other species or inanimate objects of the surrounding nature (Mollov et al., 2010; Kuzmin, 2012). Zoodiversity, 56(6): 489–494, 2022 DOI 10.15407/zoo2022.06.489 490 N. A. Smirnov Abnormal amplexus is a result of a breakdown in mating behaviour. Such a phenomenon is relatively rare because it leads to unreasonable spending of resources – energy and time (Simović et al., 2014). As an exception, there is a multiple amplexus common to species with “explosive” reproduction and which were documented, for example, for Bombina variegata (Linnaeus, 1758), Bufo bufo (Linnaeus, 1758), Rana temporaria Linnaeus, 1758, Rana iberica Boulenger, 1878 and others (Hartel et al., 2007; Ayres, 2008; Kuzmin, 2012; Đorđević & Simović, 2014). Fewer interspecifi c amplexus was found (among beings of diff erent species) (Gherghel et al., 2008 a; Strugariu & Gherghel, 2008; Simović et al., 2014; Rocha et al., 2015; Gül et al., 2018; Mačát & Jablon- ski, 2018; Mačát et al., 2019; Vásquez-Cruz et al., 2019; Decemson et al., 2020; Amin et al., 2021; Koynova & Natchev, 2021; Pupins et al., 2022), between males and dead females (Pintanel et al., 2021; Pupins et al., 2022), with other males (Mollov et al., 2010) or with inanimate objects (Mollov et al., 2010; Đorđević & Simović, 2014; Pupins et al., 2022) and even the cases with females act like a male (Malashichev, 1999). Th e information about abnormal amplexus for the territory of Ukraine is rather scarce. Th ere are re- ports on copulation of Pelophylax ridibundus (Pallas, 1771) with Bufotes viridis (Laurenti, 1768) in Crimea (Shcherbak, 1966), Pelobates fuscus (Laurenti, 1768) with Epidalea calamita (Laurenti, 1768) in Volyn (Zabroda and Ilyenko, 1981), Rana dalmatina Fitzinger in Bonaparte, 1839 with R. temporaria in Chernivtsi Region (Smirnov, 2013), multiple amplexus of B. bufo and attempt of R. temporaria to interact with other species (Pysanets, 2014). In the Carpathians (Shcherbak & Shcherban, 1980) abnormal amplexus was registered for B. bufo (multiple amplexus, couples with other species, interacting with dead females or a piece of wood), B. viridis (multiple and interspecifi c amplexus), Pelophylax lessonae (Camerano, 1882) (interspecifi c amplexus), R. temporaria (multiple amplexus). Diff erent types of abnormal amplexus were notifi ed in Kharkiv Region (multiple amplexus in B. bufo, interspecifi c amplexus between Rana arvalis Nilsson, 1842 and P. fuscus, R. ar- valis and B. bufo, B. bufo and P. ridibundus, P. fuscus and Triturus cristatus (Laurenti, 1768)) (Shabanov, 2011). Th e purpose of this work is to characterize the cases of abnormal amplexus detected in the Chernivtsi Region of Ukraine. Material and methods Th e paper is based on the results of fi eld studies conducted by the author during March–June 2008– 2021. We studied the reproductive biology of amphibians during surveys of various types of water bodies in Chernivtsi Region in the breeding season. Th e cases of abnormal amplexus were registered by photos or videos where possible. Coordinates were determined by using the package of Google Earth or GPS navigators Garmin eTrex 10 and eTrex 30. We measured the temperature of air (Ta) or water (Tw) by a mercury-in-glass thermometer (resolution 0.1 °С). Results and discussion During the research we collected information about 21 cases of abnormal amplexus (table 1) from the following types: multiple amplexus (fi g. 1, a, b), interspecifi c amplexus (fi g. 1, c–e), amplexus with a dead specimen, incorrect embrace, amplexus between a male and inanimate object (fi g. 1, f). In some cases, we observed a combination of several types of amplexus, e. g. amplexus among two males of R. dalmatina and dead male R. temporaria, incorrect embrace (from the abdominal side) of a dead R. temporaria female by a male of the same species, multiple amplexus and embrace of R. temporaria female from the abdominal side (fi g. 1, b). More than half of the noted cases (11 or 52.3 %) were multiple amplexus among one female and two or more males. Such behaviour was observed in three species of amphibi- ans — B. variegata, B. bufo and R. temporaria. Moreover, in most cases registered in B. variegata and R. temporaria (3 and 6 accordingly), it was a combination of multiple amplexus and incorrect embrace (a part of males grasped female from the abdominal side). Th e cases of multiple amplexus for this species were known from a number of publications (Shcherbak and Shcherban, 1980; Hartel et al., 2007; Gherghel et al., 2008 b; Mollov et al., 2010; Kuzmin, 2012; Đorđević and Simović, 2014; Pysanets, 2014; Pupins et al., 2022). Th e disparity in the ratio of sex during reproduction may be a possible explanation of this phenomenon. Males of these species outnumber females in a breeding site, thus, females are a valuable resource for males (Hartel et al., 2007; Mollov et al., 2010). Interspecifi c amplexus is the second type by the frequency of detection (6 cases or 28.6  %). Males of R. dalmatina tended to make mistakes, trying to make amplexus with other species, more oft en (three cases). One mistake registered in B. bufo, B. variegata and 491Th e cases of abnormal amplexus in Anura on the territory of the Chernivtsi Region… T a b l e 1 . Cases of abnormal amplexus in anurans in the Chernivtsi Region Type of aberration Species Date, Locality, Coordinates Notes Multiple amplexus Bombina varie- gata 13.06.2010, Stebnyk tract, Bere- homet urban vill., Vyzhnytsia Distr. (48.145 N, 25.278 E) Two cases: 1 }, 5 {; 1 }, 2 { Bombina varie- gata 2.06.2011, Krasnoilsk urban vill., Vyzhnytsia Distr. (48.021 N, 25.506 E) 1 }, 5 { Rana temporaria 27.03.2012, Stebnyk tract, Bere- homet urban vill., Vyzhnytsia Distr. (48.145 N, 25.266 E) 1 }, 3 { (Tw 3.5 °C) Bombina varie- gata 22.05.2012, Stebnyk tract, Bere- homet urban vill., Vyzhnytsia Distr. (48.143 N, 25.259 E) 1 }, 7 { Rana temporaria 5–6.04.2016, Perkalab forest plot, Vyzhnytsia distr. (47.80 N, 24.95 E) Five cases: 1 }, 2 { Bufo bufo 22.03.2017, “Hariachyi Urban” forest park, Chernivtsi (48.296 N, 25.988 E) 1 }, 2 { (Ta 16.0 °C; Tw 10.5 °C) Interspecifi c amplexus Bufo bufo × Pelo- phylax lessonae 31.03.2009, Shypyntsi vill., Chernivt- si Distr. (48.369 N, 25.734 E) A { B. bufo has embraced a { P. lessonae (dead animals on the road) Bombina varie- gata × Lissotriton montandoni 13.05.2010, Stebnyk tract, Bere- homet urban vill., Vyzhnytsia Distr. (48.145 N, 25.280 E) A { B. variegata has em- braced a } L. montandoni (Tw 18.9 °C) Rana dalmatina × Rana tempo- raria 20.03.2012, “Hariachyi Urban” forest park, Chernivtsi (48.293 N, 25.990 E) A { R. dalmatina has embraced a } R. temporaria (on the bank of a pond; Ta 11.0 °C) Rana dalmatina × Rana tempo- raria 21.03.2012, “Tsetsyno” reserve, Chernivtsi (48.305 N, 25.837 E) Two { of R. dalmatina has embraced a dead { R. tem- poraria (Tw 9.0 °C) Rana dalmatina × Rana tempo- raria 2.04.2013, “Hariachyi Urban” forest park, Chernivtsi (48.293 N, 25.990 E) A { R. dalmatina has embraced a { R. temporaria (Ta 10.0 °C) Rana temporaria × Pelophylax ridibundus 29.03.2016, Kitsman` town, Cher- nivtsi Distr. (48.454 N, 25.738 E) A { R. temporaria has em- braced a } P. ridibundus (on the bank of a pond; Ta 14.5 °C) Incorrect embrace and amplexus between alive male and dead female Rana temporaria 25.03.2010, “Hariachyi Urban” forest park, Chernivtsi (48.293 N, 25.989 E) A { R. temporaria has embraced a dead } R. temporaria from the abdominal side (Tw 12.2 °C) Incorrect embrace Rana dalmatina 12.03.2014, “Hariachyi Urban” forest park, Chernivtsi (48.293 N, 25.990 E) A { R. dalmatina has embraced a } R. dalmatina from the abdominal side (Tw 7.5°C) Rana dalmatina 26.03.2021, near Valia Kuzmina vill., Chernivtsi Distr. (48.164 N, 26.047 E) A { R. dalmatina has em- braced a } upside down in the axillar area (Tw 7.5 °C) Amplexus between the male and inanimate object Bufo bufo 3.04.2016, “Krugle Boloto” Lake, Shypyntsi vill., Chernivtsi Distr. (48.367 N, 25.732 E) A { B. bufo has embraced a piece of candock’s root (Nuphar lutea (L.) Smith 1809), fl oating on the sur- face of the water R. temporaria. Victims of this type of amplexus were R. temporaria (three cases), P. lessonae and P. ridibundus. Amplexus between individuals from diff erent orders, not only between diff erent species, genera or families, is especially interesting. Th at was observed in interact- ing between a B. variegata male and a Lissotriton montandoni (Boulenger, 1880) female (fi g. 1, e). Th e cases of amplexus between anurans and urodelans were described in scien- 492 N. A. Smirnov Fig. 1. Th e examples of abnormal amplexus: a — multiple amplexus in Bombina variegata (one female, seven males), May 22, 2012, Stebnyk tract, Berehomet, Vyzhnytsia District; b — multiple amplexus in Rana temporaria (one female, two males), April 6, 2016, Perkalab, Vyzhnytsia District; c — interspecifi c amplexus ({ R. temporaria × } P. ridibundus), March 29, 2016, Kitsman, Chernivtsi District; d — interspecifi c amplexus ({ R. dalmatina × } R. temporaria), March 20, 2012, “Hariachyi Urban” Forest Park, Chernivtsi; e — interspecifi c amplexus ({ B. variegata × } L. montandoni), May 13, 2010, Stebnyk tract, Berehomet, Vyzhnytsia District; f — amplexus between a male and inanimate object ({ Bufo bufo and the root of Nuphar lutea (L.) Smith 1809), April 3, 2016, Lake “Krugle boloto”, Shypyntsi, Chernivtsi District. f d b e c a 493Th e cases of abnormal amplexus in Anura on the territory of the Chernivtsi Region… tifi c literature, though they are extremely rare (Shabanov, 2011; Simović et al., 2014; Mačát et al., 2019; Koynova and Natchev, 2021). Moreover, no information on similar cases with the participation of L. montandoni in published sources has been found. In three cases there were incorrect grasping of females by the males. In one of them, the R. temporaria male captured a dead female of the same species from the abdominal side. Th at is a combination of some types of abnormality — incorrect embrace and amplexus with a dead female. In another case, a R. dalmatina male interacted by inguinal amplexus with a female of the same species but mates’ heads were turned in diametrically opposite directions. One case of a male R. dalmatina interaction with female of the same species from the abdominal side was observed. Also, we observed amplexus between the male B. bufo and the piece of candock’s root (N. lutea) (fi g. 1, f). Interacting of males of this species with inanimate objects are also known. For example, some investigators described amplexus of B. bufo with a piece of wood (Shcherbak and Shcherban, 1980), a lump of silt (Đorđević and Simović, 2014) and, even, plastic cup (Mollov et al., 2010) and plastic bottle (Pupins et al., 2022). Seemingly, that is the result of a lack of females in breeding sites. In general, our research confi rms that such abnormal behaviour is rather rare. Th us, throughout the studies during 14 fi eld seasons (2008–2021) we observed thousands of cases of amplexus in many species of anurans. But given data represent only a small percentage of the total observations. 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Peculiarities of reproduction and larval development of common spade- foot — Pelobates fuscus (Laur.) from the Ukrainian Polesie. Vestnik zoologii, 4, 66–71 [In Russian]. Received 21 September 2022 Accepted 24 November 2022