HISTORY OF RESEARCH ON THE INSECTS OF THE KURIL ISLANDS
The earliest published information on the insects of the Kuril Islands appeared in the middle of the 19th century in the papers of Motschulsky (1860), Hagen (1856), and Sharp (1896). On the basis of material collected by others, they recorded the presence of particular species on the islands and described new ones: Motschulsky (1860) mentioned 13 species of beetles from the Kurils on the basis of collections made by Voznesenskiy, a laboratory assistant of the Russian Academy of Sciences and a well-known Russian traveler who visited the Kuril Islands in 1844-1845. Hagen (1856) described two species of Kuril Island dragonflies and Sharp (1896) referred to the presence of three curculio genera on the islands. The English captain H. J. Snow made a collection of insects from the islands of the central part of the Archipelago that were later studied by Sharpe and other European scientists.
Judging from the available scientific literature, research was not conducted on the Kurils for a extended period after 1875, when the Islands came under Japanese control. Particular islands were, however, occasionally visited by foreign scientists: in 1886 the Englishman D. G. Leech collected butterflies (Heterocera) on Shikotan, and in 1908 the German zoologist K. Rost collected insects on Kunashir and Iturup.
Japanese entomologists began to study the entomofauna of the Kuril Islands in detail only after a mass outbreak of the Siberian moth in the southern area of Sakhalin Island and the southern Kurils in 1920-1921. Aidzawa and Harada, the first Japanese entomologists to work in the Archipelago, devoted attention principally to the study of the Siberian moth. Matsumura (1925, 1926a-b) published a synopsis of the species of the genus Dendrolimus injurious to firs, and on their insect parasites and predators, based on material collected by Aidzawa and Harada. Doi then collected insects on Kunashir, Shikotan, Iturup, Urup, Paramushir, and Shumshu. A scientific expedition working on the northern islands of Paramushir and Shumshu collected diverse botanical, theriological, ornithological, acarological, entomological, and hydrobiological materials in 1931. Asahina, who was studying dragonflies at the time, collected insects on Paramushir in July and August 1933. Sugihara collected insects on the southern Kurils (Kunashir, Shikotan, Iturup, and Urup) over a number of years (1935, 1936, 1940): jointly with Uchida in 1935 and with Kuwayama in 1940. The majority of the specimens collected by these investigators is presently kept at Hokkaido University, Sapporo, and at the Hokkaido Scientific Research institute of Agriculture.
In 1929 and 1930, the Swedish ornithologist Bergman visited many islands in various parts of the Kuril Archipelago and assembled comprehensive zoological materials, including a collection of insects that is now held at the Museum of Natural History in Stockholm. After analyzing some of these specimens, Bryk (1942) published a large paper on the Kuril Island butterflies that has retained its scientific significance to this day. A series of articles devoted to the entomofauna of the Kurils based on materials collected by Japanese investigators was published in various Japanese journals. Typically, these papers consist of lists of species of various groups of insects, with information on the island localities where they were found and on their general distribution. In some cases, food plants and, for parasitic insects, host insects, are indicated. Data on population sizes of the various species, their confinement to particular floral assemblages, and injurious nature are generally lacking. A considerable portion of these articles were published in Japanese.
The earliest of this series of publications (Ohta, 1929; Kobayashi, 1931; Takeuchi, 1931) were devoted to sawflies and beetles of the families Coccinellidae and Chrysomelidae. A report on a comprehensive 1931 expedition, dealing with the environment of the northern Kurils, was published in 1933-1934 in the Bulletin of the Biogeographic Society of Japan. In additional to other studies, three entomological papers were included in this report: Esaki (1933-1934), Takeuchi (1933-1934), and Kano (1933-1934), in which two species of diurnal butterflies, four species of sawflies, and 62 species of beetles encountered on the Kurils are cited. The discovery there of stoneflies, mayflies, caddis-flies, and dipterans, 14 species in all, is noted in an paper by Uéno (1933-1934); but because the identifications were based on larval stages, the majority of the specimens were identified only to genus--only two were identified to species.
During this same period, the well-known Japanese entomologist Kôno was analyzing materials obtained from the islands and concentrating specifically on the beetles of the Kuril Archipelago and Sakhalin Island. He was responsible for a large number of papers relating to various coleopteran families. Ten articles (Kôno, 1935a-e, 1936a-c, 1937a-b), under the general title "Beitrag zur Kenntnis der Käferfauna der Kurilen," were devoted especially to the Coleoptera of the Kuril Islands. Numerous families of beetles are considered: Lycidae, Cantharidae, Lampyridae, Dasytidae, Lucanidae, Scarabaeidae, Cleridae, Byturidae, Nitidulidae, Endomychidae, Lathridiidae, Coccinellidae, Oedemeridae, Pyrochroidae, Mordellidae, Meloidae, Serropalpidae, Othniidae, Cucurlionidae, Cerambycidae, Lagriidae, and Languriidae. Collections made by the investigators mentioned above served as the basis for these publications: Doi, Uchida, Sukihara, and in particular, Kuwayama. In addition, data on the discovery of individual species of beetles in the Kurils are cited by Kôno in other papers on the fauna of Sakhalin and Japan (Kôno, 1929a-b, 1932, 1934, 1936d, 1937c, 1938a-b, 1939, 1940). In all, he cited about 150 beetle species for the Kuril Islands. However, many large coleopteran families were not considered in Kôno's studies: among these are the Carabidae, Histeridae, Silphidae, Staphylinidae, Elateridae, Buprestidae, Chrysomelidae, Ipidae, and others. Kôno evidently did not analyze these families because mention is made of finding them on the Kuril Islands in the work of Kano (1933-1934) and other investigators (Heikertinger, 1949; Kimoto, 1965).
A relatively small amount of information on the Coleoptera of the Kuril Islands is also found in the studies of Miwa (1934), who cited 13 species of clickbeetles from this region; Matsushita (1933b), who described Strangalia doii, a new species of long-horned beetle from Iturup; Tamanuki (1933), who cited seven species of long-horned beetles from the Kurils; and Sawamoto (1940a-b), who cited two species of Kuril Island bark-beetles, Drycoetes rugicollis Egg and Ips cembrae Heer (the latter in error).
Studies of a similar character, touching upon other orders of insects, relate to the same period. Uchida published a series of articles on the ichneumonids (Uchida, 1936c, 1940b) and lepidopterans (Uchida, 1935, 1936a-b) of the Kuril Islands. In these papers, he cited 125 species of ichneumonid flies and 46 species of butterflies (Rhopalocera).
Kuwayama (1936a-b) observed eight species of neuropterans (Neuroptera), one species of sialid (Megaloptera), one species of scorpion fly (Mecoptera), and eight species of caddis-flies (Trichoptera). Okumura (1941, 1942) cited 11 species of dragonflies for the Kurils, two of which, according to Asahina (1958), are doubtful. Sugihara (1939) published a short note on the wasps (Vespidae) of the Kuril Islands. Okada (1936, 1937), in two brief reports on dipterans of the genus Nematocera, cited six species for the southern islands and two for Urup. Additional information on the incidence of particular species of insects on the Kuril Islands can be found in general articles on the entomofauna of Japan published during this period (e.g., Matsumura, 1933; Takeuchi, 1934; Alexander, 1935).
Field studies on the Kuril Islands by Japanese investigators were suspended after 1941, and the number of Japanese publications devoted to the entomofauna of the Kuril Archipelago appreciably declined in the postwar years. Nevertheless, a few additional publications dealing with the insects of the archipelago, based on old material collected as far back as the 1930s, appeared in Japan after the end of the 1940s; but these papers do not differ in character from the prewar articles, containing not much more than lists of species, although often more complete ones. Exceptions among these are two review articles by Takeuchi (1955, 1956) on the sawflies of the Archipelago that mention one species of horntail and 86 species of sawflies belonging to six families (Cephidae, Pamphiliidae, Blasticotomidae, Argididae, Cimbicidae, and Tenthredinidae) of which eight species and four subspecies are described as new to science.
Kuwayama (1956, 1961a-b, 1962) refined the species composition of the Neuroptera of the Kurils, expanding the list to 13 species, providing descriptions of two new species, and reducing some previously recognized forms to synonyms. Asahina (1958) revised the Kuril Island dragonflies (recognizing 19 species), defined their distribution on the islands more precisely, advanced the hypothesis that all the species, except Somatochlora graeseri, can be found on Hokkaido, noted the absence on the Kurils of endemic species and subspecies, and provided a comparison with the dragonfly fauna of Kamchatka. Gilmour (1960) published a short article on the long-horned beetles of the Kuril Islands, based on Bergman's collections in the Museum of Natural History in Stockholm, in which he reported on 13 species of long-horned beetles from Urup and Kunashir. Kimoto (1965) analyzed a collection of leaf-beetles collected on the Kurils by Kuwayama and Sugihara as far back as 1940, and published for the first time a list of leaf-beetles of the Kuril Islands that includes 36 species in nine subfamilies: Donaciinae, Criocerinae, Clytrinae, Cryptocephalinae, Eumolpinae, Chrysomelinae, Synetinae, Galerucinae, and Alticinae; another five species were cited on the basis of published data. A small amount of information on the occurrence of particular leaf-beetles on the Kuril Islands is also contained in the earlier papers of other authors (Heikertinger, 1948; Chujo and Kimoto, 1961; Gressitt and Kimoto, 1961).
In a short note on the Collembola of the southern Kurils, Uchida (1965) cited five species distinguished by having a wide Holarctic distribution. A short article by Oku (1965) contains descriptions of two species of leaf rollers (Tortricidae) from the islands of Kunashir and Khoromushir. In a paper devoted to the stoneflies (Plecoptera) of the southern Kurils, Kawai (1966) cited seven species, one of which, Nemoura kuwayamai, was described as new to science; distributions of stoneflies on the southern islands of Kunashir, Shikotan, and Iturup are also given. Scattered mention of the distribution of various species of insects (borrowed mainly from the above-mentioned studies) can be found in monographs and review papers of a number of authors (e.g., Gressitt, 1951; Konishi, 1955; Plavil'shchikov, 1958; Uchida, 1958).
Finally, Kuwayama's (1967) large synoptic monograph on the insects of the southern Kuril Islands summarizes all known information, including published materials of Soviet investigators. It consists of seven chapters, in addition to a large bibliography. An introductory section provides general information on the geography and natural conditions of the southern Kurils as well as a review of Japanese industry that existed there prior to the second World War. This is followed by a brief history of entomological research in the region, general information on the entomofauna, and a list of the 20 orders of insects recorded from the islands. Various hypotheses proposed to explain the regionalization of the insects of the archipelago are considered; the insect of pests of agricultural plants, forests, fish, and fish products are cited; and, in conclusion, a list of the insects found in the southern Kuril Islands is given, which accounts for 1,917 species and takes up three-quarters of the book.
Despite an attempt to provide general information, Kuwayama's (1967) book cannot pretend to be give a complete analysis. As pointed out by the author himself, a substantial amount of material available even in Japanese collections is not included and the investigations of Russian entomologists of recent years (e.g., G. O. Krivolutskiy, V. M. Ermolenko, E. N. Savchenko, V. I. Kuznetsov, E. M. Danzig, and G. S. Zolotarenko) have shown that the diversity of some groups of insects on the Kurils is actually much greater than that cited by Kuwayama: for example, the number of known species of leaf rollers (Lepidoptera, Tortricidae) is not five but 160, of coccids (Homoptera, Coccoidea) not one but 60, of bugs (Heteroptera) not 49 but 130, of ground beetles (Coleoptera, Carabidae) not 31 but 100, and of worm moths (Lepidoptera, Noctuidae) not 92 but 165, and so on.
Kuwayama devoted considerable attention to the distribution of insects throughout the Kuril Archipelago and to the regionalization of the entomofauna within its limits. He presented interesting data on the commonality of the fauna of the Kurils and of Kamchatka, Sakhalin, Hokkaido, and to some extent other Japanese islands, but did not provide an extensive zoogeographic analysis of the entomofauna. Ecological information is almost non-existent.
Japanese entomologists have made a substantial contribution to the study of the entomofauna of the Kuril Islands. Nevertheless, various groups of insects here have been far from uniformly studied, and some of them still await investigation. In addition, many general questions relating to the ecology, morphogenesis, and origin of the island entomofauna, remain unanswered. It is also worth noting here that the Japanese literature has been difficult to use; most of it is published in Japanese and in little-known journals.
A comprehensive and well-designed study by Soviet investigators of the environment of this interesting and in many respects unique territory began soon after the Kuril Islands were returned to the Soviet Union after the war. Beginning in 1946, the Archipelago was visited annually by participants of numerous expeditions designed to address diverse scientific questions. The following account describes only the entomological investigations.
The entomologist N. N. Konakov worked on the Kuril Islands in 1946 as a member of the comprehensive Kurils Expedition organized by the Far Eastern research base of the USSR Academy of Sciences and the Primorye affiliate of the All-Union Geographic Society. He collected not only insects, but also birds and mammals. Although he published two papers on the basis of this material (Konakov, 1947, 1956), a complete analysis of the Kuril collection was circumvented by his untimely death in 1947. Bei-Bienko (1949, 1953), however, published two papers on the basis of Konakov's collections, which were transferred to the Zoological Institute of the USSR Academy of Sciences. In these papers, Bei-Bienko provided a list of species, but also a zoogeographic analysis of the orthopteroid insect fauna of the archipelago and adjacent territory. Medvedev (1952, 1960) also used Konakov's material when he described the lamellicorn beetles (Scarabaeidae) of the USSR.
The zoologists O. A. Skarlato and A. A. Strelkov made small collections of insects incidentally in 1947-1949 on Shikotan Island. These collections are housed at the Zoological Institute of the USSR Academy of Sciences. Later, in 1950-1957, N. A. Violovich examined collections of some families of dipterans on the Kuril Islands and published several articles on the flower flies (Syrphidae) and horse flies (Tabanidae). Violovich (1955, 1956b-c, 1957, 1960a-b), in his studies of flower flies, described a number of new species, increased the list of this group for Sakhalinskaya Oblast from 36 to 185 species (of these, he cited 103 for the southern Kurils), and provided a detailed zoogeographic analysis. His papers on the horseflies (Violovich, 1954, 1956a, 1968) contain descriptions of new species and a review of the Tabanidae of Sakhalinskaya Oblast; he cites 10 species of horseflies for the Kuril Islands.
Shtakel'berg (1950, 1952, 1953a-b, 1955a-b, 1956, 1958), using the collections of Konakov and Violovich, described a number of new dipteran species for the Kurils, and presented new data on their distribution in the Archipelago. Savchenko (1956) published an paper, based on materials collected by Violovich, on the crane flies (Tipulidae) of the Kuril Islands, which contains much new and interesting information on the species composition inhabiting the islands, as well as an analysis of the origin of this taxon on the Archipelago.
The entomologist N. N. Filippov visited the southern Kuril Islands (Yuriy, Zelionyi, and Kunashir) in 1954-1955 and made a large collection of insects, primarily beetles, studies of which have never been published. The entomologist R. A. Vasil'ev worked on the Kurils (Kunashir and Iturup) in September-October 1956 as a member of an expedition of the Far Eastern affiliate of the USSR Academy of Sciences; the material he collected also remain unpublished. The geologist O. N. Kabakov made a small collection of beetles on Kunashir in 1960 in which new species of leaf-beetles were later found and described by Medvedev (1966). Belyshev (1962, 1963) examined the pathways of dispersal and the history of formation of the island and peninsular faunas of northeast Asia, including the Kuril Islands, as exemplified by the dragonflies (but based on the collections of Asahina, 1958).
Kurentsov (1959a, 1960a-b, 1963a-b, 1965a, 1967, 1968a) touched on the insects of the Kuril Archipelago in a series of general papers on the zoogeography of Kamchatka and the entire Far East. He cited a number of examples from insect taxa (primarily butterflies and beetles) when reviewing specific ecological complexes and zoogeographic groupings of animals. In particular, in speaking of the problems of Beringia in the zoogeography of Kamchatka, he listed species of insects that are common to Kamchatka and the Kurils, and also cited three species of butterflies associated with Kuril bamboo: Neope gaschkewischi Mèn., Lethe diana Btl., and L. callipteris But.
By the beginning of our investigations, Soviet entomological studies and expeditions devoted to the Kuril entomofauna were limited to the above, but in the 1960s study of the insects of this region became more active. Concurrently with us and afterwards, a large number of researchers made entomological collections, primarily working on the southern islands of the Archipelago: V. A. Nechaev in 1962-1963 and N. G. Vasil'ev in 1968, Institute of Biology and Soil Sciences, Far Eastern affiliate of the Siberian Division of the USSR Academy of Sciences; A. K. Tsimbalyuk, 1964, Far Eastern State University; V. M. Ermolenko, 1965, and E. N. Savchenko and V. I. Monchenko, 1968, Zoological Institute of the Ukrainian SSR Academy of Sciences; E. M. Danzig and V. I. Kuznetsov, 1967, and K. B. Gorodkov, É. P. Narchuk, V. A. Rikhter, and M. N. Meyer, 1968, Zoological Institute of the USSR Academy of Sciences; and G. S. Zolotarenko and V. V. Nikolaev, 1968, Biological Institute of the Siberian Division of the USSR Academy of Sciences. The entomologist G. S. Lafer made large collections of insects, primarily beetles, on Kunashir, Shikotan, and Iturup. The collections made by all of these individuals have been used in the present work.
Our investigations have provided additional data on the species composition of the insects of the Kurils, defined more precisely their distribution within the limits of the Archipelago, added new ecological and phenological information, identified harmful species, provided a zoogeographic analysis of the insect fauna of the Archipelago, and demonstrated the pathways of its formation and origin. A number of papers on particular insect groups inhabiting the Kurils have been published in recent years on the basis of materials we have collected: Bei-Bienko (1966), orthopteroid insects; Krivolutskaya and Ivanovskaya-Shubina (1966), aphids; Anufriev (1968a, 1970), cicadans; Medvedev (1966), net-winged beetles; Gur'eva and Krivolutskaya (1968), clickbeetles; Krivolutskaya (1962b, 1966), long-horned beetles; Krivolutskaya and Medvedev (1966), leaf-beetles; Krivolutskaya (1963, 1965a), bark-beetles; Krivolutskaya and Stroganova (1966), horntails; Savchenko and Krivolutskaya (1966), crane flies; Konovalova (1966b-c, 1967, 1968), butterflies.
Prior to these studies, information on the distribution of some of these groups on the islands (aphids, cicadans, bark-beetles, horntails) had been nearly absent in the literature. In other groups (orthopteroids, clickbeetles, leaf-beetles, long-horned beetles, crane flies), the number of known species has been increased two-fold and, in some cases, three-fold. A considerable number of new species have been described (e.g., Emel'yanov, 1966; Krivolutskaya and Ivanovskaya-Shubina, 1966; Medvedev, 1966; Savchenko and Krivolutskaya, 1966; Kerzhner, 1968; Krivolutskaya, 1968). Species lists published in these articles up to 1967 were included by Kuwayama (1967) in his book on the entomofauna of the southern Kuril Islands.
General problems relating to patterns of distribution of insects in the Archipelago, their trophic linkages, ecological and geographic peculiarities, and the morphogenesis and origin of the entomofauna of the Kuril Islands, were addressed by Krivolutskaya (1962a, 1964, 1966, 1967, 1969), Krivolutskaya and Medvedev (1966), Konovalova (1967), Kurentsov and Krivolutskaya (1967). More detailed commentary on these works is provided in the chapters below.
New information on the Kuril entomofauna continues to be received. In particular, Kuznetsov (1986b) and Danzig (1968) presented interesting reports on the Kuril Island leaf-rollers (Lepidoptera, Tortricidae) and coccids (Homoptera, Coccoidea) at the XIII International Entomological Congress in Moscow. Medvedev and Ermolenko (1969) published a paper on the lamellicorn beetles (superfamily Lamellicornia) of the Kurils in which several species are described as new for the islands. Savchenko (1970) published additions to the crane fly fauna (Diptera, Tipulidae) of the southern Kurils, describing yet another new species and increasing the list of Kuril tipulids by 16 species.
In concluding this review of investigations devoted to the entomofauna of the Kuril Archipelago, it must be emphasized that despite the large number of publications, the insects of the Kurils have been far from fully studied. Considerably greater efforts are required to resolve the wide range of questions relating to the entomofauna of this region.
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