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GROUND BEETLES (COLEOPTERA, CARABIDAE) IN FUMAROLE FIELDS OF KUNASHIR ISLAND, KURIL ARCHIPELAGO, RUSSIA

1Moscow State Pedagogical University, Institute of Biology & Chemistry Zoology & Ecology Department, Kibalchicha str. 6, build. 3, Moscow 129164, Russia E-mail: kvmac@inbox.ru, https://orcid.org/0000-0002-9184-7869 2Federal Scientific Center of East Asia Terrestrial Biodiversity, Zoology Department Laboratory of Entomology, Pr. 100-letiya Vladivostoka 159, 690022 Vladivostok, Russia E-mail: yun-sundukov@mail.ru https://orcid.org/0000-0003-3312-4029 3Pirogov Russian National Research Medical University, Biology Department Ostrovitianova Str. 1, 117997 Moscow, Russia E-mail: andrei-matalin@yandex.ru, https://orcid.org/0000-0002-7790-8709


INTRODUCTION
The Kunashir Island is the southernmost and one of the largest within the Greater Kuril Chain. It is 123 km long, 7-30 km wide, and has an area of 1490 km 2 . The island consists of three mountain blocks formed by four active volcanoes: Tyatya (1819 m) and Ruruy (1485 m) in the north, Mendeleev (889 m) in the centre, and Golovnin (541 m) in the south. These blocks are separated by the South Kuril and Sernovodsk isthmuses, composed of quaternary marine sediments and volcanic-sedimentary folded Neogene rocks.
Volcanism played and continues to play the leading role in influencing the landscapes on the island. Due to residual volcanism, the soil, water and air in the fumarole fields are enriched with sulfurous compounds; the vegetation is strongly depressed and degraded, while the temperature of the soil and above-soil air is markedly increased. As a result, the original landscapes are destroyed, while pioneer or modified habitats are formed.
Although Kunashir Island is the most biodiverse one of the Kuril Archipelago, the study of its carabid fauna started relatively recently: the first information appeared only in the second half of the 20th century (Konakov 1956, Habu 1967, Kuwayama 1967, Kryzhanovskij 1968). These publications contain data about 24 ground beetle species, 20 of which were listed by Kuwayama (1967).
Further research during the last quarter of the 20th century increased the taxonomic diversity of Carabidae of the island to 119 species (Kryvolutskaja 1973, Kryzhanovskij et al. 1975, Lafer 1989, 1992. Taking into account all recorded and still unpublished data, the carabid fauna of Kunashir Island currently amounts to at least 180 species (Sundukov, 2001, 2019a-b, Makarov & Sundukov 2011, 2016, 2019. However, there are no publications on the carabid fauna of the fumarole fields of Kunashir. Only one paper, reports three species on the fumarole fields of the volcanoes of the South Kurils Cicindela sachalinensis A. Morawitz, 1862, Cylindera elisae (Motschulsky, 1859), and Carabus blaptoides rugipennis Motschulsky, 1861 (Konakov 1956).
Thus, the adaptive responses of Carabidae to the extreme conditions of the fumarole fields on Kunashir Island remain to be described, and this is the main objective of the present paper.

Research area
According to the geological morphogenetic classification (Fedorchenko et al. 1989), active Kunashir volcanoes belong to three types: Tyatya is a single stratovolcano, Ruruy and Mendeleev are isolated "cluster" stratovolcanoes, while Golovnin is a caldera pumicepyroclastic volcano destroyed mainly by erosion and denudation. The latter three volcanoes are presently at a hydrothermal-solfataric stage of activity. Thus, denuded fumarole fields occupy the slopes or calderas of volcanic edifices, with several rivers and streams flowing from there. The temperature of these streams reaches 100 °C, sulphate and sulphate-chloride waters are acidic (minimal pH ca. 2.0), while hydro-carbonate-sulphate sodium-calcium waters are subneutral (pH up to 8.5). A detailed description of the studied solfatara fields is given in Appendix 1.

Ground beetle collections
This study is based on the material taken on Kunashir Island during nine field seasons. The first and third authors, together with I. V. Melnik, A. A. Zaitsev and A. S. Prosvirov (all from Moscow), captured beetles in 2008, 2009, 2011, 2013 and 2017. The second author, together with L. A. Sundukova (Lazo, Primorsky Territory), collected insects from

Sequence alignment
The nucleotide sequences obtained were manually edited to correct possible sequencing errors and to delete low-resolution terminal segments using the Unipro UGENE v33.0 (Okonechnikov et al. 2012) and then aligned in MEGA-X v10.1.6 using the MUS-CLE algorithm (Kumar et al. 2018). Ambiguously aligned regions were excluded from the downstream analyses. The edited nucleotide sequences were deposited in GenBank under accession numbers MW240488-MW240502.

Phylogenetic analysis
To clarify the taxonomic status of the particular populations of Cylindera (Eugrapha) elisae (Motschulsky, 1859) from different Kunashir Island localities, additional C. elisae sequences from GenBank from various localities in Japan, Korea and Taiwan were used, and the sequences from GenBank corresponding to specimens of C. (E.) bonina (Nakane et Kurosawa, 1959) were also applied to the analysis as outgroup. A haplotype network to visualize the relationships among haplotypes for the C. elisae sequence dataset was calculated using PopART v.1.7 (Leigh & Bryant 2015). A Minimum spanning network was constructed with default settings for the aligned haplotype sequences (Bandelt et al. 1999).
The evolutionary analysis was inferred by using the Maximum Likelihood method and Kimura 2-parameter model (Kimura 1980, Tamura et al. 2012. Initial tree(s) for the heuristic search were automatically obtained by applying Neighbor-Join and BioNJ algorithms to a matrix of pairwise distances estimated using the Maximum Composite Likelihood (MCL) approach, and then selecting the topology with the superior log likelihood value. This analysis involved 113 nucleotide sequences. A total of 849 positions were in the final dataset. Evolutionary analyses were conducted in MEGA X (Kumar et al. 2018).

Measurements
The measurements (in mm) were made with an ocular-micrometer mounted on a Leica M165c (Carl Zeiss) stereo microscope, as follows: CI -confidence interval; ELgreatest length of elytra; EW -greatest width of elytra; Lae -length of the aedeagus; Llm2 -length of lamina 2; Lms -length of main sclerite; PLm -length of pronotum, measured along the median line; PW -greatest width of pronotum; TL -total body length without labrum (from anterior margin of clypeus to the elytral apex along the suture). The nomenclature of the sclerites of the male internal sac in Bembidion follows Belousov and Sokolov (1996), as well as Neri and Vigna Taglianti (2010).

Data analysis
Statistical analysis was performed using the PAST v4.0 software (Hammer et al. 2001). The significance was determined by Tukey's and Mann-Whitney U-test for independent variables, with a 95% confidence interval. If statistically significant differences were found, pairwise Dunn's a posteriori tests were additionally executed. Frequencies of colour variants were analysed by Mann-Whitney and χ 2 tests. The influence of temperature and pH was assessed using a multivariate multiple linear regression analysis.

RESULTS AND DISCUSSION
A total of 846 specimens belonging to 29 species of ground beetles (Appendix 4) were collected in the fumarole fields, but only five species (780 specimens) were permanent inhabitants of these unusual habitats.

Cicindela (Cicindela) sachalinensis sachalinensis
Cicindela sachalinensis occurs in eastern and central China, Korea, eastern Mongolia, the Russian Far East, and Japan (Wiesner 1992, Puchkov & Matalin 2003. Among the five subspecies, the nominal one is known exclusively from insular habitats of Sakhalin, Moneron, Iturup, Kunashir, Shikotan (Kryzhanovskij et al. 1975, Lafer 1978, Makarov et al. 2020, Hokkaido and Honshu (Wiesner 1992, Puchkov & Matalin 2003. This species is widely distributed over the Kunashir Island (Appendices: Fig. A13) and is mostly observed in habitats without or only with scant plant cover (Kryzhanovskij et al. 1975, Lafer 1978, Makarov et al. 2019b. It was present in almost all fumarola fields studied (Appendices: Fig. A8) but recorded in high numbers only in Golovnin Volcano on the fumaroles of the central domes. The specimens inhabiting the fumarole fields were similar in size to specimens from other habitats, except for the significantly smaller females (Figs 9A, B). On the fumaroles, significantly more (p << 0.01) dark-pigmented adults were collected (  Cylindera elisae is an Asiatic tiger beetle occurring from the Amur River valley through Mongolia, Korea and China (Puchkov & Matalin 2003 to northern Vietnam (Wiesner et al. 2017), and also in Sakhalin, Kunashir, Taiwan and various Japanese islands. On Kunashir Island, this species is found only on fumarole fields of two volcanoes (Appendices: Figs A9, A12). One form, C. e. kunashirensis Pütz et Wiesner, 1994, inhabits the valley of Kislaya River on the Mendeleev Volcano (Kryzhanovskij et al. 1975, Lafer 1978, Pütz & Wiesner 1994, Sabirov et al. 2014, Makarov et al. 2019b, Sundukov & Makarov 2019, while a different form populates the Neskuchenskiye Streams on the Ruruy Volcano (Makarov et al. 2019b, Sundukov & Makarov 2019. The specimens from these localities are well distinguishable by size, the proportions of the labrum, and elytral coloration. Thus, C. e. kunashirensis is smaller: in males, TL = 8.74 mm [CI = 7.8-9.2 mm] vs. 9.38 mm [CI = 8.55-10.05 mm] in the specimens from the Ruruy Volcano (p << 0.01), in females, TL = 9.81 mm [CI = 8.6-10.6 mm] vs. 10.19  in specimens from the Ruruy Volcano (p < 0.05). Besides, C. e. kunashirensis shows a significantly (p << 0.001) narrower labrum, darker elytra (Figs 1A-F), and darker elytral punctation. The barcoding technique also verified the distinction of C. e. kunashirensis. According to the data obtained, specimens from the Kislaya River formed a compact cluster clearly separated from most other local forms of C. elisae (Fig. 2), their haplotypes being separated from the basic one by 206 mutational steps. By the white pattern on the elytra (Figs 1B-F), as well as the shape of the aedeagus (Figs 1G-L), specimens from the Ruruy Volcano are more similar to those from Sakhalin and the continental Russian Far East than to C. e. novitia, the latter inhabiting northern Hokkaido. All these observations allow us to conclude that the Ruruy specimens belong to the nominative subspecies. Interestingly, in the Japanese sub-prefecture of Nemuro, where the eastern coast of the Shiretoko Peninsula is 24-43 km from the Kunashir Island, C. elisae has not yet been recorded (Kimoto & Yasuda 1995).

Bembidion (Ocydromus) dolorosum
Bembidion dolorosum is widespread in Japan (Hokkaido, Honshu), also inhabiting southern Sakhalin (north to Dolinsk, 47°19') and Moneron Island, as well as the southern Kuriles (Urup, Iturup, Kunashir, Shikotan, and islands of the Lesser Kuril Chain) (Nakane 1963, Inouye 1971, Kryzhanovskij et al. 1975, Watanabe 1989, Lafer 1998, 2002a, 2006, Yoshitake et al. 2011, Sundukov 2017, Sundukov & Makarov 2013, 2016, Makarov et al. 2019b). On Kunashir Island, it occurs on the banks of almost all rocky or sandy rivers and streams (Appendices: Fig. A14), in all fumarola fields studied (Appendices: Figs A2, A4, A6), sometimes in very high numbers. In fumarolas, we observed it feeding on small dead insects, mostly flies (Appendices: Figs A10, A11). In the northern part of the distribution area (Sakhalin and Kuril Islands), B. dolorosum is characterized by moderate variability and a clear-cut sexual dimorphism (Figs 10A,B). In males, TL = 5.19 mm [CI = 5.14-5.23 mm], in females, TL = 5.35 mm . The proportions of the head and pronotum are slightly variable. The differences between the populations from different islands are usually not significant, only the males from Yuriy Island and the females from Sakhalin Island in most cases differ significantly. These demonstrate clinal variability, because the body size, coloration and main body proportions gradually changed under an environmental acid- ity gradient. Along transects with a moderate gradient, these changes are relatively small (Fig. 4A), but is more pronounced under greater range (Fig. 4B).
The COI sequences of the specimens inhabiting the fumarola fields and other habitats were identical or differed only by 1−2 mutation steps. All specimens formed a compact cluster on the Ocydromus phylogenetic tree (Appen-  Fig. A95). D. Maddison (pers. comm.) has sequenced six genes (COI, 28S, wingless, CAD, topoisomerase, and Muscle-Specific Protein 300 (MSP)) for one specimen of the B. dolorosum form (from the Shiretoko Peninsula, Hokkaido Isl.) and one specimen we provided of the fumarole from the caldera of Golovnin Volcano, Kunashir Isl. These two specimens had identical sequences in COI, 28S, and differed by a single base (among 487 bases) in wingless, one base (among 767 total bases) in CAD, four bases (among 876 bases) in topoisomerase, and two bases (among 765 bases) in MSP.
Note. Lafer (2002b) described Bembidion (Peryphus) kuznetsovi from the bank of Lake Kipyashcheye and compared his new species to Bembidion (Pery-

Bembidion (Peryphanes) sanatum
This species is widely distributed in different habitats over a wide altitudinal range on the Japanese islands of Hokkaido and Honshu (Bates 1883, Jedlička 1965, Watanabe 1989, Kimoto & Yasuda 1995, Yasui & Shiyake 2008, Yoshitake et al. 2011, Mori 2016, Yoshimatsu et al. 2018), but in the Kunashir Island, it was found only on the slopes of the Mendeleev Volcano (Appendices: Fig. A15), in the valley of Kislaya River (Kryzhanovskij et al. 1975, Makarov et al. 2019b. The beetles inhabit the bank of an acidic mineralized stream flowing down from fumarola fields (Appendices: Fig. A4). It noteworthy that these fumarola fields and streams are also the sole habitat for the endemic Kunashir tiger beetle, Cylindera elisae kunashirensis. Notes. In the original description, Bates (1883) considered B. sanatum as closer to the European B. lunatum (Duftschmid, 1812). Later authors placed this species to the subgenus Peryphus Dejean, 1821 (Netolitzky 1943, Jedlička 1965, Kirschenhofer 1984, while others to Bembidion incertae sedis (Marggi et al. 2017). Based on the structure of the male aedeagus and female spermatheca, B. sanatum belongs to Peryphanes . Most species of this subgenus are restricted to the western Palaearctic. In eastern Asia, only five species are known to occur (Jedlička 1933, Habu & Uéno 1955, Habu 1973, Nakane 1979, Kirschenhofer 1984, which are recorded from Japan (B. dostali Kirschenhofer, 1984;B. hayachinense Nakane, 1979;B. hikosanum Habu et Uéno, 1955), China (B. parepum Jedlička, 1933) or Taiwan (B. lulinense Habu, 1973). Originally, all those species were described in different subgenera, and only recently were they all classified as Peryphanes (Toledano 2009, Marggi et al. 2017. Based on the conformation of the spermatheca (a large number of whorls of a sclerotized vas deferens, the proportions and shape of the chambers), as well as structure of the aedeagus (large lateral scle- rites, large basal sclerites protruding far from the base, the absence of a ventral tubercle), B. sanatum is very similar to the western Palaearctic Bembidion (Peryphanes) stephensii Crotch, 1866 (Fig. 7).

CONCLUSIONS
In most cases, the study of the influence of volcanic activity is limited to direct, often catastrophic effects on animals, including Coleoptera (see review by Elizalde 2014). Fumarola fields are one of the manifestations of volcanism under constant change. The micro-relief of the fumarola fields is formed or destroyed by thermal springs, mud pools and steam-gas emissions, also because hydrothermally altered rocks are subjected to denudation. According to lichen The presence of such environmental gradients creates the prerequisites for gradual development of the adaptations in different species during their occupation of fumarola fields. Thus, the species inhabiting the fumarolas could be considered as model objects for the study of microevolutionary processes. On Kunashir Island, only a few species of ground beetles have been able to colonize such fumarole fields. It is hardly surprising that the inhabitants of open spaces or the rocky banks of streams have become successful colonists. However, only C. sachalinensis and B. dolorosum were recorded from all volcanoes studied (Appendices: Figs A13, A14), while the remaining few species were narrowly localized. Cylindera elisae was found only on the Mendeleev and Ruruy volcanoes (Appendices: Fig. A12), B. sanatum only on the Mendeleev Volcano (Appendices: Fig. A15), while P. samurai, widespread across the island (Appendices: Fig. A16), only on the fumaroles of the Dokuchaev Mountain Ridge. Obviously, this unevenness and patchiness could have hardly been caused by environmental conditions, which were generally similar in all fumarole fields studied, and requires additional explanation.
The reaction of carabids to the special conditions of fumarole fields was species-specific. On the one hand, fumarola-inhabiting specimens of C. sachalinensis slightly differed from those in other habitats (Figs 8A, 9A, B); likewise, B. sanatum from the Kunashir, hardly differed from specimens from Honshu. On the other hand, specimens of P. samurai populating the fumarole fields differed noticeably in size and melanization (Figs 8B, 9D; Appendices: Figs A96-A97 vs. A98-A99), while C. elisae kunashirensis, in addition to the smaller size (Fig. 9C) and melanization (Fig. 1A), showed a relatively narrow labrum and was noticeably separated genetically (Fig. 2). The most interesting pattern of variability in all studied fumarole fields is demonstrated by B. dolorosum: as both temperature and acidity increase, specimens of this species become relatively small, elongated, and partially depigmented.
These differences could be associated both with certain features of different species and with the time of their adaptations to the conditions of fumarola fields. According to the data obtained, morphological changes do not depend on the taxonomic similarity of a species. Thus, B. dolorosum was extremely variable, while B. sanatum showed almost no particular change on the fumarola fields. Similar differences can also be seen between C. sachalinensis and C. elisae.
It is more difficult to assess the influence of time in the adaptation process. We can reasonably assume that widespread, flying species found in high numbers near the borders of fumarole fields (C. sachalinensis, B. dolorosum) have constantly inhabited these particular habitats. Specimens taken within or beyond fumarola fields were not distinguishable by their barcodes. Yet, GROUND BEETLES IN FUMAROLE FIELDS OF KUNASHIR ISLAND, RUSSIA they showed differences in morphological adaptations. Cicindela sachalinensis changed only slightly, while B. dolorosum, under increasing temperatures and acidity, was capable of developing forms that could erroneously be determined as different species.
The denuded landscapes of the fumarole fields and the headwaters of the streams on the Mendeleev Volcano, the single place where both Kunashir endemic carabids, C. elisae kunashirensis and B. sanatum, have been found, are formed at the site of explosion craters about 2,100-1,500 yBP (Abdurakhmanov et al. 2004). It is evident that such landscapes could have existed on the cone of Mendeleev Volcano earlier. The oldest volcanic layers not covered by marine sediments date back to about 39,000 yBP (Lebedev et al. 1980). We believe that the conditions of thermal (including fumarole) fields could have ensured the survival of some species under the colder conditions during the Pleistocene and Holocene, as in Bembidion ruruy Makarov et Sundukov, 2014 and several rove beetles (Shavrin & Makarov 2019). Therefore, we believe that C. elisae kunashirensis might have colonized the Kunashir Island much earlier, and the time of its isolation could probably be associated with the Last Glacial Maximum, ca. 20,000 yBP (Clark et al. 2009).
Earlier DNA studies support our hypothesis. Based on the sequences of the 28S rDNA and COI genes (Sota et al. 2011), C. elisae from various localities in Korea, Japan and Taiwan shows minimal divergence, especially in the COI sequences. At all localities studied, only one haplotype is widely distributed, the one that seems to have given rise to the remaining 17 haplotypes. Thereby most of those numerous haplotypes are separated from the basal one by no more than one to three mutational steps. Only the Taiwanese C. e. reductelineata is distinct from the basal haplotype by 18-26 mutational steps (Sota et al. 2011: fig. 5A). The divergence time of this subspecies is ca. 0.6 Mya, this being comparable with the divergence time of ca .0.9 Mya for the more strongly related Cylindera bonina (Nakane et Kurosawa, 1959) which is endemic to the Bonin Islands (Sota et al. 2011: Fig. 4). According to these data, most of the insular and mainland subspecies of C. elisae (at least C. e. koreanica, C. e. novitia, C. e. mikurana and C. e. formosana) could be considered varieties. The presence of two subspecies of C. elisae on the Kunashir Island is not unique, because two subspecies of this species co-occur also in Taiwan. According to Sota et al. (2011), C. e. reductelineata could have populated Taiwan much earlier than C. e. formosana did. Based on the results of DNA analyses performed by us and considering the results by Sota et al. (2011), we can assume that C. e. kunashirensis could have colonized the Kunashir Island much earlier than the nominative subspecies. In our opinion, the evolution under the particular conditions of fumarola fields and the long-term isolation might have caused a significant genetic divergence of this subspecies.
Estimating the divergence time of B. sanatum is more difficult because the species, both morphologically and genetically, is very close to the European B. stephensii, due to possible convergence (Netolitzky 1943: 37). Such patterns/disjunctions (amphi-Palaearctic or Euro-Manchurian) are also found among plants (Nakamura 2008, Denk & Grimm 2009, vertebrates and various insect orders, such as Lepidoptera (Dubatolov & Kosterin 2000), Odo natoptera (Kosterin 2002), Diptera (Oosterbroek et al. 2001), and Coleoptera (Semenov-Tian-Shansky 1911). Usually, the time of the origin of such disjunctions is attributed to the Pliocene or Miocene (Mikkola 1987, Nakamura 2008. However, is likely that the later, repeated periods of warming allowed repeated recolonisation of former distribution areas, and that an estimate of 110,000-100,000 yBP is more realistic (Belova 1985, Dubatolov & Kosterin 2000, Kosterin 2002). Thus, we can assume that the ancestors of B. sanatum could have entered this region no later than 100,000 yBP, and the isolation of the B. sanatum was associated with the Last Glacial Maximum under the conditioning influence of fumarola fields.
Although the isolation of the endemic forms of C. elisae and B. sanatum on the Mendeleev Volcano might have occurred at about the same time, the level of genetic divergence differs by an order of magnitude. The variance in the COI sequence of C. elisae elisae and C. elisae kunashirensis amounts to 206 mutation steps, while the same parameter for B. sanatum and B. stephensii is only 66.
We conclude that the degree of the divergence of ground beetles, both morphological and genetic, during the adaptation to the specific conditions of fumarole fields varies very considerably and does not depend on taxonomic similarity or the time of colonization of fumarolas. This forces us to be careful in generalizations such as "the level of morphological (genetic) differences is enough to distinguish a species or subspecies".
Taking the ground beetles living in the specific and extreme conditions of fumarola fields as examples, we can see that the combination of flexible and conservative traits is one of the important reasons for the exceptionally high biological diversity of this family Appendix 1. Detailed characteristics of the studied fumarola fields.

Ruruy Volcano
Despite the high density and variety of volcanic edifices in the northern part of the Kunashir Island, two areas of thermal-solfatara activity: "Neskuchenskie springs" and "Dal'nie springs" are known only on the western seashore.
The "Neskuchenskie Springs" is stretched along the seacoast for ca. 1.2 km and covers an area of about 1.5 km 2 . The fumarola fields a total area ca. 3000 m 2 (Zharkov 2014). The "First Upper" solfatara field is located at an altitude of 70 m a.s.l. and covers an area of about 1700 m 2 . The solfatara activity is observed in its apical part, where the gas temperature reaches 100 °C. The "Second Upper" solfatara field with an area of about 800 m 2 , is located slightly to the north. Its fumarola outcrops also have a temperature of 100.0-100.9°C, and the waters of the streams flowing from there are subneutral (pH = 6.9-7.3). The "Third Upper" solfatara field is located at an altitude of 210 m a.s.l. Its sulphate calcium-sodium waters are acidic (pH = 2.2), with the temperatures of up to 91.7 °C. Besides, there are two "extinct" fumarola sites and numerous acidic hydrothermal and solitary fumarola outcrops.
The "Dal'nie Springs" site was discovered only in 2017 (Sundukov & Kozlovski 2017) and volcanologists have never studied it. The site with hydrothermal-solfataric activity was located in a bowl-shaped depression at an altitude of 390-560 m a.s.l. and covered an area of about 1 km 2 . It includes two solfatara fields, "Dlinnoe" and "Bolshoye". The "Bolshoye" field is 3500 m 2 located at 390-400 m a.s.l. The "Dlinnoe" field is 100 m to the southwest and has an area of about 1500 m 2 . In both these fields, only a weak solfataric activity is presently observed. The gases escaping from the fumarolas have a temperature of 18.0-22.3 °C, the temperature of the streams flowing through the fields ranges from 20.4 °C to 22.5 °C, their water being weakly acidic (pH = 4.0-4.2).

Mendeleev Volcano
The outputs of the solfatara gases are concentrated on the Mendeleev Volcano in four fields: southeastern, eastern, northeastern, and northwestern. In the valleys of the rivers and streams originating from these fields, there are groups of thermal springs with varying temperatures and chemical composition (Tables 1 & 2).
The southeastern field is 400-575 m a.s.l. and has an area of about 75,000 m 2 , with numerous outlets of small gas jets, as well as thermal and cold mineral springs in its upper part. Thermophilic algae grow in places where the thermal waters come out, and there is a strong odour of hydrogen sulfide. The water is acidic (pH = 2.5), siliceous, sulphatechloride, with a complex cationic composition and a mineralization level of about 1 g/l (Markhinin & Stratula 1977).

GROUND BEETLES IN FUMAROLE FIELDS OF KUNASHIR ISLAND, RUSSIA
The eastern field is stretched out in a narrow strip along the upper reaches of Lechebnyi Stream at 650-450 m a.s.l. It includes a small thermal lake (T = 60-85 °C, pH = 3.3) and numerous springs, mud griffins and funnels with the temperatures of 53-101 °C and pH = 2.7-3.6. The thermal waters are sulphate calcium-magnesium with increased content of Al 3+ , Fe 2+ , H + and a mineralization level of about 1 g/l. According to the composition of gases, they are carbonic: CO 2 -43.6%, N 2 -39.1%, CH 4 -14.2%, with no H 2 S detected (Markhinin & Stratula 1977).
The northeastern field is the most active area. In 1880, a weak phreatic eruption occurred there (Milne 1896(Milne ), and in 1901(Milne , 1946(Milne , 1977(Milne and 1987 emissions of the steam-gas jets up to a height of 150-200 m and tangible earthquakes were observed. This fumarola field is at 250-350 m a.s.l. and an area ca. 2 km 2 . There are three eroded funnels of phreatic explosions, which are the centres of discharging acid sulphate-chloride fluids . There are numerous fumarolas with the temperatures of up to 99.6 °С, and in the valley of Kislaya River, flowing from this field, there are dozens of springs with the temperatures of 90.0-98.5 °С and pH = 2.4-2.5. The waters of these springs are acidic, carbonic (CO 2 -about 89%, H 2 S -up to 4.7%, O 2 -0.5%) sulphate sodium-calcium-magnesium hydrothermal fluids (Markhinin & Stratula 1977). The northwest field is at a 400 m a.s.l. and is about 50,000 m 2 in area. Its solfataric activity is relatively high: in 1978 and 1984, the temperature increased to 111-113 °C, accompanied by the appearance of molten sulfur (Zharkov 2014). This field also supports a "boiling" spring with a temperature of > 90 °C. Its water is highly acidic (pH = 2.5-3.0), sulphate-chloridic, with a total mineralization level of up to 4.4 g/l. Carbon dioxide and hydrogen sulphide (CO 2 -88.1%, H 2 S -9.8%, N 2 -2.1%, the CH 4 content being negligible) prevail in the gas composition (Markhinin & Stratula 1977).

Golovnin Volcano
A large eruption that formed the modern caldera about 4.7 km in diameter, according to various estimates, occurred about 52,000-30,000 yBP (Melekestsev et al. 1988, Bulgakov 1994. At present, the caldera contains five terrestrial and one underwater fumarola fields and two lakes, with thermal waters. Its chemical composition is given in Tables 1 and 2. The Central Eastern solfatara field is located at the foot of the southern slope of the Central Eastern dome. All gas outlets and thermal springs are situated along the shores or under the water of Lake Kipyashcheye. Both solfataras (T = 90-100 °C), as well as the boiling cauldrons of various sizes, consistencies and colors (T = 60-95 °C), are located there. Their waters range from subneutral hydrocarbonate-sulphate sodium-calcium (pH = 6.0-8.5) to acidic sodium sulphate (pH = 2.0-2.5) (Zharkov 2014).
Lake Kipyashcheye is a phreatic explosion funnel filled with water, with an area of 66,000 m 2 and a depth of 16 m (Kozlov & Zharkov 2010). Its acidic (pH = 2.5), chloridesulphate sodium waters are heated up to 90 °С in the places where gas-hydrothermal vents emerge. In the central part of the lake, the chemical composition of the water is predominantly sodium chloride, pH = 3.7, and the surface temperature is about 30 °C (Markhinin & Stratula 1977).
Lake Goryacheye occupies the northern part of the caldera with a 3.1 km 2 area, 62.3 m maximum depth, and a surface water temperature of 17-18 °C. The water is acidic (pH = 2.5-3.0) and contains sulphate-chloride sodium-calcium (Zotov et al. 1988). There are four fumarola fields at the lake shores.
The Central Western solfatara field is at the southern shore of the lake. Its formation seems to have been associated with a phreatic eruption on the northern slope of the Central West extrusive dome. The solfataras and thermal boilers are the hottest ones on the volcano -from 90 °С to 102.5 °С (Zharkov 2014). In terms of gas composition, they are similar to the solfatars of the Central Eastern field, but carbon dioxide and hydrogen sulphide are contained in equal proportions (48% each), and the condensate of the gases has a sulphate calcium chemical composition and pH = 2.8 (Markhinin & Stratula 1977). The lower part of this field shows a temperature of 98 °C and has highly acidic (pH = 2.0), sulphate-chloride calcium-sodium waters.
At the northern bank of Lake Goryacheye, three small solfatara fields are located: Cherepakhovoye, Nabokovskoye, and Bezymyannoye. The solfatars and hydrosulfatars of the Cherepakhovoye field are quite intense. The temperature of the solfataras reaches 98 °C, while the temperature of the springs is 80 °C; the solfataras are carbonic (CO 2 -94%) with acidic spring (pH = 2.3), while sulphate calcium-sodium with medium mineralized, waters (Mar khinin & Stratula 1977). The Nabokovskoye field has four low-rate extinct sources with a temperature of 38-52 °C and pH = 6.0 (Zharkov 2014). Within the Bezy myannoye field, the solfataras are concentrated to two areas: at and a few dozen meters off the bank of Lake Goryacheye. This field is characterised by the development of mud pots with temperatures up to 89 °C and pH = 3.4. In the channel of the stream and the upland areas, there are Aggressive gases and hightemperature solutions continually affected the lithogenic base of all fumarola fields for an extended period. These processes have completely changed the composition of loose sediments and bedrocks, being accompanied by exogenous processes and the transformation of landforms (Razjigaeva 2005). These resulted in denuded volcanic landscapes with destroyed or sparse vegetation (Figs A2-A7) and a buried soil cover (Ganzey 2004).