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Ecologica Montenegrina 39: 121-128 (2021) This journal is available online at: www.biotaxa.org/em http://dx.doi.org/10.37828/em.2021.39.13

New occurrences, morphology, and imaginal phenology of the rarest tiger tundrana (: )

IVAN N. BOLOTOV1,2,3, VITALY M. SPITSYN1, EVGENY S. BABUSHKIN3,4, ELISAVETA A. SPITSYNA1, YULIA S. KOLOSOVA1 & NATALIA A. ZUBRII1,2

1N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Northern Dvina Emb. 23, 163000 , 2Northern Arctic Federal University, Northern Dvina Emb. 17, 163000 Arkhangelsk, Russia 3Saint-Petersburg State University, Universitetskaya Emb. 7/9, 199034 Saint Petersburg, Russia 4Surgut State University, Lenina Av. 1, 628403 Surgut, Russia *Correspondence: [email protected]

Received: 11 January 2021│ Accepted by V. Pešić: 15 February 2021 │ Published online: 17 February 2021.

The Arctic tiger moth Arctia tundrana (Tshistjakov, 1990) is one of the largest and most attractive lepidopteran species in the Eurasian Arctic, being ranged from the Kolguev Island () to the northeastern edge of the continent (Tshistjakov 1990; Bolotov et al. 2015). The type locality of this species is as follows: “Magadan Oblast, Chukotka Autonomous Okrug, Chaplino, hot springs” [64.4244°N, 172.5014°W] (Tshistjakov 1990; Bolotov et al. 2015). Previously, this Palearctic taxon was downgraded to a subspecies of the Nearctic species Arctia subnebulosa (Dyar, 1899) (Dubatolov 2010). The latter species has a vague type locality, which is based on four syntypes as follows: “Nushagak, Alaska <...>, Point Barrow <...> and Bethel, Kuskoquim River” (Dyar 1899). The range of Arctia subnebulosa covers Yukon, Alaska, the Pribiloff Islands, and the St. Matthew Island but does not cross the Bering Strait (Bolotov et al. 2015). Recently, it was shown that these taxa are distant phylogenetically based the COI gene fragment and that they most likely belong to separate species (Bolotov et al. 2015). Later, this mtDNA-based hypothesis was confirmed using several nuclear DNA markers, and Arctia tundrana was resurrected as a valid species (Rönkä et al. 2016). The ranges, ecology, and biology of Arctia tundrana and A. subnebulosa were discussed in detail using a nearly complete occurrence data set, field observations, and a thorough review of the body of available literature (Bolotov et al. 2015). Fibiger et al. (2011) transferred these taxa from Sotavalta, 1965 to Arctia Schrank, 1802 based on the male genitalia structure, and this generic placement was supported by subsequent DNA-based research (Rönkä et al. 2016). This study (1) displays markings pattern of male and female specimens of Arctia tundrana from various parts of its broad range; (2) illustrates a paratype male specimen of this species with its genitalia and aedeagus; (3) presents a few additional occurrences of A. tundrana supplementing the data set published in our earlier paper (Bolotov et al. 2015); (4) provides an updated map of the species’ occurrences; and (5) discusses its imaginal phenology based on long-term occurrence data.

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Samples were processed using standard approaches for (Bolotov et al. 2018). Museum abbreviations are as follows: RMBH – Russian Museum of Biodiversity Hotspots of the Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Arkhangelsk, Russia; ZMMU – Zoological Museum of the Moscow University, Moscow, Russia; ZISP – Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia.

Figure 1. Male specimens of Arctia tundrana. (A) [locality no. 48; see Figs. 4 and 5C], 06.viii.2019 [coll. RMBH]. (B) Tareya village, Taymyr [locality no. 12; see Fig. 4], vii.1967 [coll. ZMMU]. (C) Agapa River, Taymyr [locality no. 17; see Fig. 4], 23.vii.1973 [coll. ZMMU]. (D) Kresty village, Taymyr [locality no. 14; see Fig. 4], 12.vii.1976 [coll. ZMMU]. (E) Uelen settlement, Chukotka [locality no. 51; see Fig. 4], 19.vii.1979 [coll. ZMMU]. (F- H) Naukan settlement, Chukotka [locality no. 51; see Fig. 4], 14.vii.1979 [coll. ZMMU]. Scale bar = 10 mm. (Photos: Elisaveta A. Spitsyna and Vitaly M. Spitsyn).

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At first glance, there are some longitudinal shifts in the markings pattern of Arctia tundrana imagoes (Figs. 1-2). In particular, male samples from Yamal and Taymyr are slightly darker compared with those from Chukotka (Fig. 1), while a female specimen from Taymyr shared a more developed yellow markings pattern compared with those from Yamal and Kolguev (Fig. 2). However, a much larger series is needed to estimate the geographic variability of this rare species in detail. Additionally, a paratype male specimen of Arctia tundrana from Chukotka is illustrated here for the first time, including the genitalia and aedeagus (Fig. 3).

Figure 2. Female specimens of Arctia tundrana. (A) Near Bugrino village, Kolguev Island [locality no. 1; see Fig. 4], 26.vii.2009 [coll. RMBH]. (B) Near Seyakha village, Yamal Peninsula [locality no. 49; see Fig. 4 and 5D], 31.vii.2014 [coll. RMBH]. (C) Agapa River, Taymyr [locality no. 17; see Fig. 4], 12.vii.1976 [coll. ZMMU]. Scale bar = 10 mm. (Photos: Elisaveta A. Spitsyna and Vitaly M. Spitsyn).

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Figure 3. Male paratype of Arctia tundrana [coll. ZMMU]. (A) Dorsal view of the specimen with corresponding labels. The geographic label [in Russian] reads as follows: “Chukotka, Koluchinskaya Bay , Belyak’s Sandspit, 17.vii.1988, P. Tomkovitch leg.” [locality no. 24; see Fig. 4] (B) Genitalia and aedeagus. (Photos: Vitaly M. Spitsyn).

Table 1. List of additional occurrences of Arctia tundrana. This data supplements Table S2 of Bolotov et al. (2015). The numbers of localities in the table correspond to those in the map (Fig. 4).

Coordinates** No Country Region Locality and sample data* Altitude, m Reference Latitude Longitude 48 Russia Yamal-Nenets Shokalsky Island at the eastern 1 72.8597 74.6200 This study Autonomous side of the mouth of the District River, , 06.viii.2019, 1♂, Lapsui leg. [coll. RMBH; see Figs. 1A and 5C] 49 Russia Yamal-Nenets Yamal Peninsula, grass-herb 3 70.1525 72.4361 This study Autonomous nival meadow on a southern District slope of a hill near Seyakha village, 31.vii.2014, 1♀ (it was jumping through plants over distances <1 m), Zubrii leg. [coll. RMBH; see Figs. 1B and 5D] 50 Russia Arkhangelsk , Yuzhny 7 71.5590 52.3240 Kullberg et Region (Southern) Island, Belushja al. (2019) Guba, 14.vii.1907, 1 larva [coll. ZISP] 51 Russia Chukotka Naukan settlement, 140 66.0272 -169.7022 This study Autonomous 14.vii.1979, 3♂, Tomkovich District leg. [coll. ZMMU; see Fig. 1F-H] 51 Russia Chukotka Uelen settlement, 19.vii.1979, 10 66.1578 -169.8009 This study Autonomous 1♂, Tomkovich leg. [coll. District ZMMU; see Fig. 1E]

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Altogether five occurrences of Arctia tundrana were added to our earlier data set (Bolotov et al. 2015), i.e. those from Novaya Zemlya, Shokalsky Island, Yamal Peninsula, and two new records from Chukotka (Table 1; Fig. 4). The two first insular occurrences are somewhat remarkable, as the brachypterous females of this species can fly (or rather jump) over very short distances only, i.e. less than 1 m (see Table 1: locality no. 49). The Shokalsky Island is located at the eastern side of the mouth of the Ob River, and is separated from the mainland by a narrow and shallow strait. It is likely that this island was isolated from the continent in the Early Holocene, as was Kolguev (Bolotov et al. 2015; Artamonova et al. 2020).

Figure 4. Updated map of Arctia tundrana (Palearctic species) and A. subnebulosa (Nearctic species) occurrences (earlier version of this map is presented in Bolotov et al. 2015: Fig. 1A). The red circles indicate new occurrences of A. tundrana (Table 1). The yellow circles indicate A. tundrana occurrences published by Bolotov et al. (2015). The violet circles indicate occurrences of A. subnebulosa, a Nearctic vicariate of A. tundrana (after Bolotov et al. 2015). The numbers of localities correspond to those in Table 1 (this study) and Table S2 (Bolotov et al. 2015: Online Resource 1).

In contrast, a deep and wide marine barrier separates Novaya Zemlya from the , which cannot easily be crossed not only by flightless but also by some bumblebees (Potapov et al. 2019). A growing body of phylogenetic and paleogeographic research revealed that Novaya Zemlya was a cryptic refugium for some terrestrial and freshwater organisms during the entire Pleistocene epoch, e.g. a variety of plants (Serebryanny and Malyasova 1998), the Glacier bumblebee Bombus glacialis Friese, 1902 (Potapov et al. 2017, 2019), and the Arctic charr Salvelinus alpinus (Linnaeus, 1758) (Makhrov et al. 2019). Based on these data, we could assume that Arctia tundrana immigrated into Novaya Zemlya well before the Last Glacial Maximum but this preliminary hypothesis needs to be checked using a phylogenetic approach. Another explanation is that wind-borne dispersal of early instar larvae might support Arctia tundrana expansion throughout shelf islands of the , as it was shown for other arctiine species (review: Bolotov et al. 2015). In total, this species was collected from 35 localities throughout Northern Eurasia during the period of 1904-2019 (Fig. 4). The map of European distribution of Arctia tundrana published in “Noctuidae Europaeae, vol. 13” (Fibiger et al. 2011: 122) reveals that this species is ranged in northern Scandinavia. However, it seems to be a technical failure, as it is unknown west of the Kolguev Island, and there is no occurrences from Fennoscandia (Bolotov et al. 2015).

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Figure 5. Habitats of Arctia tundrana in Northeastern (A-B) and Western Siberia (C-D). (A) Bog with lichens, mosses, and cottongrasses near Bugrino village, Kolguev Island [locality no. 1]. (B) Mountain dry willow-grass tundra near Amderma village, Yugor Peninsula [locality no. 2]. (C) Wet tundra, Shokalsky Island [locality no. 48]. (D) Nival grass-herb meadow on a southern slope of a hill near Seyakha village, Yamal [locality no. 49]. The numbers of localities correspond to those in the map (Fig. 4). (Photos: Boris Y. Filippov [A-B], Evgeny S. Babushkin [C], and Natalia A. Zubrii [D]).

Based on the number of available occurrences, Arctia tundrana could be considered one of the rarest Palearctic tiger moths, and is comparable with A. menetriesii (Eversmann, 1846), a widespread but extremely rare boreal species (Bolotov et al. 2013; Berlov and Bolotov 2015). Although larvae of Arctia tundrana could be numerous locally, the abundance of imago is always low due to the high levels of parasitoid infestation (Bolotov et al. 2015). It was shown that the high parasitoid pressure could significantly reduce the imago abundance of several other tiger moth species (Shilenkov and Richter 1998). This species inhabits a broad range of habitats, which vary from mountain dry tundra and grass-herb meadows to lowland wet tundra and bogs (Fig. 5). Our phenological analysis reveals that Arctia tundrana flights from mid-June to early August (Fig. 6). However, most specimens were collected in July, with the peak value between 11 and 20 July. It was shown that the emergence of its imago is clearly linked to the warmest days of the summer season (Bolotov et al. 2015). It is likely that the development of this species lasts two years, although this hypothesis is yet to be confirmed experimentally (Bolotov et al. 2015). As a conclusion, the range of this species is rather well known but its life history and geographical patterns of morphological variability need future research efforts. Although Arctia tundrana could be a polyphagous species like the majority of Palearctic (e.g. Berlov and Bolotov 2015; Koshkin 2019), its native host plants are still unknown (Bolotov et al. 2015). Earlier, we described the morphology of the last instar larva, pupa, and cocoon (Bolotov et al. 2015) but external patterns of early instar larvae as well as the entire life history of this species are yet to be characterized. Finally, widespread tiger moth taxa such as

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Arctia tundrana, A. menetriesii, and A. lapponica (Thunberg, 1791) can be used as appropriate models for continent-scale phylogeographic and genomic research combined with morphological analyses (e.g. Hegna et al. 2015).

Figure 6. Imaginal phenology of Arctia tundrana based on the long-term collection data of 1904-2019 (Bolotov et al. 2015: Table S2; this study: Table 1). Specimens with uncertain collecting dates were excluded from the data set.

Acknowledgements

We are grateful to an anonymous reviewer who helped us to improve an earlier version of this correspondence. This study was partly supported by the Ministry of Science and Higher Education of the Russian Federation (projects 0409-2019-0042 to V.M.S. and 0793-2020-0005 to I.N.B.), Russian Science Foundation (project 19-14-00066 to E.S.B. and I.N.B.), and Russian Foundation for Basic Research (project 19-34-90012 to V.M.S. and I.N.B.), and the Department of Education and Youth Policy of the Khanty- Mansiysk Autonomous Okrug – Ugra (to E.S.B.). Special thanks goes to Dr. Andrey V. Sviridov (ZMMU, Moscow, Russia) for his help during this survey.

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