Ecologica Montenegrina 31: 46-49 (2020) This journal is available online at: www.biotaxa.org/em http://dx.doi.org/10.37828/em.2020.31.10 Hot spring puddling by butterflies YULIA S. KOLOSOVA*, OLGA V. AKSENOVA, ILYA V. VIKHREV & IVAN N. BOLOTOV N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Northern Dvina Emb. 23, 163000, Arkhangelsk, Russia *Corresponding author: [email protected] Received: 2 May 2020│ Accepted by V. Pešić: 20 May 2020 │ Published online: 23 May 2020. Puddling behavior of butterflies and moths is a well-known phenomenon driven by a deficit of several minerals and nutrients in larval and imago diet, especially sodium and proteins (Arms et al. 1974; Adler 1982; Boggs and Jackson 1991; Beck et al. 1999; John & Tennent 2012; Inoue et al. 2012). In particular, sodium and albumin were found to be the most attractive puddling resources for tropical butterflies on Borneo based on the results of a long-term experimental study (Beck et al. 1999). This kind of behavior is more characteristic for males, while female butterfly puddling occurs only occasionally (Beck et al. 1999; Adler & Pearson 1982; Scriber 1987, 2002; John & Tennent 2012; John & Dennis 2019). Male puddling could increase reproductive success in butterflies because minerals and nutrients are transferred through the spermatophore at mating (Boggs & Gilbert 1979; Pivnick & McNeil 1987; Smedley & Eisner 1996; Dennis et al. 2014; Mitra et al. 2016). It was shown that another purpose of puddling by males of swallowtail butterflies is to excrete excessive potassium (Inoue et al. 2015). A large body of literature describes the utilization by butterflies of various puddling sources such as mud puddles, edges of streams, aphid honeydew, tree sap, carrion, perspiration, saliva, urine, and excreta (e.g. Scriber 1987; Scriber & Ayres 1988; Boggs & Jackson 1991; Otis et al. 2006). A few unusual examples of moths feeding on bird tears were recorded in South America and Madagascar (Hilgartner et al. 2007; Sazima 2015; De Lima Moraes 2019). Different lepidopteran species appear to prefer different puddling substrates, probably looking for species-specific sets of nutrients (Boggs & Dau 2004). It was shown that carnivore urine was more preferable for puddling butterflies in North America compared with herbivore and omnivore urine that could be influenced by a specific sodium or amino acid/volatile content (Bodri 2018). Marine (seawater) puddling is known to occur in at least 21 species of butterflies, including papilionids, nymphalids, pierids, lycaenids, and hespereiids (Pola & García-París 2005; John & Tennent 2012; John & Dennis 2019). If marine puddling by butterflies was considered a rare (or rarely seen) kind of behavior (John & Tennent 2012), puddling on geothermal sources was virtually unknown until recently, when large aggregations of Trogonoptera brookiana (Wallace, 1855) males were discovered puddling on a hot spring in Malaysia (Phon et al. 2017). To the best of our knowledge, this observation is the only example of hot spring (geothermal) puddling by butterflies published to date. In this correspondence, we report on the field observations of puddling by butterflies on warm mineral water from hot springs in Northern Asia. Water temperatures were measured using a digital thermometer (TK-5.05, TekhnoAS, Russia). Water samples were collected from each puddling site using a Ecologica Montenegrina, 31, 2020, 46-49 KOLOSOVA ET AL. 0.5 L polyethylene bottle and kept cool and dark until transfer to the laboratory. Hydrochemical analyses were performed in the core facility center of the Northern Arctic Federal University, Arkhangelsk, Russia. Photographs were taken using a Canon EOS 60D camera (Canon Inc., Tokyo, Japan). First, we recorded hot spring puddling by two males of the Chinese peacock butterfly Papilio bianor Cramer, [1777] at the travertine field of the Neskuchensky hot springs [44.4853°N, 146.0978°E, Kunashir Island, Kurile Archipelago, eastern Russia, 24 July 2011, observations by Ilya V. Vikhrev] during at least 30 min (Fig. 1). The water was slightly brackish with high mineral [Na+: 51.8; Ca2+: 43.1; Mg2+: 17.0; K+: 15.9; 2– – -1 + – -1 SO4 : 97.5; and Cl : 16.1 mg×L ] and nitrogen content [NH4 : 4.6; and NO3 : 0.2 mg×L ] (sample no. 3- 2011-Ku), and a temperature of 40°C. Figure 1. Hot spring puddling by the Chinese peacock butterfly Papilio bianor Cramer, [1777] at the travertine field of the Neskuchensky hot springs, Kunashir Island, Russian Far East, 24 July 2011. (A) Observation site on the shore of the island. (B-C) Puddling by a male butterfly on warm mineral water. A dead larva of the carrion beetle species Silpha perforata Gebler, 1832 can be seen to the right of the butterfly (bottom photo). (Photos: Yulia Kolosova [A] and Ilya V. Vikhrev [B-C]). Ecologica Montenegrina, 31, 2020, 46-49 47 HOT SPRING PUDDLING BY BUTTERFLIES Second, males of three butterfly species were observed puddling on a geothermal swamp with algal mats near the Kotel hot springs [53.5139°N, 158.7543°E, Nalychevo Natural Park, Kamchatka, eastern Russia, 01 July 2012, observations by Ivan N. Bolotov]. These species were as follows: the Old World swallowtail Papilio machaon Linnaeus, 1758 [N = 2], the green-veined white Pieris napi (Linnaeus, 1758) [N = 3], and the northern chequered skipper Carterocephalus silvicola (Meigen, 1829) [N = 1]. The water + 2+ 2+ + 2– was also slightly brackish with high mineral [Na : 160.0; Ca : 45.6; Mg : 5.9; K : 23.0; SO4 : 59.3; and – -1 + – -1 Cl : 189.0 mg×L ] and nitrogen content [NH4 : 3.9; and NO3 : 2.3 mg×L ] (sample no. 2210-Wn), and a temperature of 25–30°C. At first glance, puddling on geothermal sources can be considered similar to the utilization of seawater, although hot springs usually share a much lower concentration of total dissolved solids compared with that in marine waters. Coastlines of tropical and subtropical seas with warm saline or brackish water and algal mats are rather similar environmentally to hot springs. For example, seawater puddling by a group of Papilio polytes males was recorded at low tide on algal mats and exposed corals of a reef shelf with a water temperature range of 36–42ºC on Guam Island, Micronesia (Pola & García-París 2005). The antennae of swallowtail butterflies bear specific sensillae responding to ammonia and humidity that could be useful in puddling behavior because NH3-emitting water bodies may represent sources rich in sodium and other nutrients (Inoue et al. 2019). This finding indicates that geothermal sources can attract butterflies by way of higher levels of ammonia emission and ammonium concentration compared with those in surrounding water bodies. In summary, five butterfly species were found puddling on geothermal water so far (Table 1). Warm mineral water from hot springs seems to be an appropriate but rarely exploited resource of sodium and other nutrients for male butterflies. We would hope that our brief report could motivate further observations on hot spring puddling behavior of butterflies worldwide. Table 1. Occurrences of hot spring puddling by butterflies. Family Species Sex Region Reference Papilionidae Papilio bianor Cramer, [1777] Male Kunashir Island [44.4853°N, 146.0978°E], This study eastern Russia Papilionidae Papilio machaon Linnaeus, Male Nalychevo Natural Park, Kamchatka This study 1758 [53.5139°N, 158.7543°E], eastern Russia Papilionidae Trogonoptera brookiana Male Ulu Geroh [4.4402°N, 101.2505°E], Phon et al. (Wallace, 1855) Perak, Malaysia (2017) Pieridae Pieris napi (Linnaeus, 1758) Male Nalychevo Natural Park, Kamchatka This study [53.5139°N, 158.7543°E], eastern Russia Hesperiidae Carterocephalus silvicola Male Nalychevo Natural Park, Kamchatka This study (Meigen, 1829) [53.5139°N, 158.7543°E], eastern Russia Acknowledgements We are grateful to the anonymous reviewer who helped us to improve an earlier version of this paper. 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