CHAPTER 4

ECOLOGICAL AND GEOGRAPHICAL OUTLINE OF THE ENTOMOFAUNA OF THE KURIL ARCHIPELAGO

SECTION 3. ENTOMOLOGICAL COMPLEXES CHARACTERISTIC FOR THE PRINCIPAL BIOTOPES OF THE KURIL ISLANDS

As an invariable component of any biogeocenosis, insects populate the entire territory of the Kuril archipelago. However, as already indicated in section 2 of this chapter, entomological complexes that differ both qualitatively and quantitatively are characteristic of various biotopes. They also differ with respect to their ecological requirements, genetic associations, and origin. Of the entire diversity of biotopes existing on the Kuril Islands, we distinguish the following basic biotopes according to the character of the entomofauna: broadleaved and coniferous-broadleaved forests, dark coniferous forests, larch stands, birch stands, alder stands, floodplain stands which are represented primarily by willow stands, elfin Japanese stone pine stands, Kuril bamboo groves, herbaceous associations, and near-fumarole and supralittoral areas.

As can be seen from this list, the character of the majority of the biotopes is determined by the composition of the vegetation, on which the composition of the insects also directly depends. The more complex the composition of the vegetation, the more diverse the entomofauna associated with it. The entomofauna reaches its greatest diversity in mixed coniferous-broadleaved forests, where there are more opportunities for the multiplication of insects and other animals.
 

The entomofauna of the broadleaved and coniferous-broadleaved forests:
 

The broadleaved forests of the Southern Kuril Islands (Kunashir and Iturup) are mainly derivatives of the broadleaved-dark coniferous forests. They are exceptionally complex in composition and mosaic with respect to the distribution of the principal forest-forming species. Dark coniferous species, the degree of whose participation in the composition of the stand fluctuates markedly, also typically grow in such forests. In describing these forests, A. S. Ageenko et al. (1969) emphasize that sections of multispecies stands are frequently encountered among this forest formation so complex that it is impossible to establish the main forest-forming species in them. These forests undoubtedly are the legacy of Tertiary thermo- and hygrophilous mixed forests which were widely distributed in the past in North East Asia, and which have a relict character. Not without reason does A. I. Tolmachev (1959) assign them to the East Asian floristic region, and in a more fractional division, to the Sakhalin-Hokkaido province. It is also not accidental that the coniferous-broadleaved forests achieve special luxuriance at the present time on the Okhotsk littoral of Kunashir, in sites with the most favorable climatic conditions, in particular near Alekhino set. It is appropriate at this point to cite the remark of S. Kuwayama (Kuwayama, 1967) to the effect that the climatic conditions in the region of Alekhino set., with respect to many indicators, are more favorable than in Sapporo on Hokkaido.

In combination with the variation in the conditions of the mountain slopes of various exposures, the alternation of markedly shaded areas under the canopy of a multitiered mixed forest, where many tree trunks are densely entwined by vines, with open areas of forest clearings, occupied by vigorously growing tall herbaceous vegetation or Kuril bamboo, creates an exceptional diversity of ecological niches which support the existence of a multiplicity of diverse insects that form macro- and microcomplexes. These forests are especially interesting as preservers of an ancient moisture-loving flora and fauna of East Asia, which has preserved archaic features up until the present time. The postulate formulated by P. M. Rafes (1968), to the effect that "the formed forest, more that any other type of vegetation, and for a longer period, insulates the space delimited by it, and the forest habitat persistently preserves its specific characteristics. Therefore, forest biogeocenoses are characterized by great stability" (p. 9), is entirely applicable to the Kuril broadleaved forests. However, the character of the entomofauna of the broadleaved and coniferous-broadleaved forests of Kunashir and Iturup is not entirely identical, as is the case with the character of these forests themselves as well; this has already been mentioned in the analysis of particular groups of insects. Therefore, it is expedient to examine the corresponding entomological complexes separately.

The entomofauna of the broadleaved forests of the southern part of Kunashir, especially its southwest coast, is most distinctive and rich. The majority of southern species, which impart to the Kuril entomofauna, as it does to the entire landscape, a brilliant coloration, is concentrated precisely in this complex. Because of the multispecies stands, the combination of groupings of insects associated with many woody and shrub species is observed: oak, elms, maple, cherry, bird cherry, mountain ash, mulberry, Bothrocaryum, cork tree, sen, magnolia, ash, birch, alder, aspen, willows, trichocarpous toxicodendron, panicled hydrangea, spruces, Sakhalin fir, and Japanese stone pine. Not a few species develop on different shrubs which form the underbrush, as well as on herbaceous plants growing under the forest canopy. In connection with the fact that open sections within the woods (large clearings, logging area swaths) are occupied by Kuril bamboo, species are also present there which are associated with this specific biotope, and which supplement the diversity of the entomofauna of the broadleaved forests.

Since the presentation of complete faunistic lists would occupy too much space, let us dwell on visually-striking landscape or background species that characterize the plant formation under consideration (Figures 39, 40). These are, first and foremost, the large and brilliantly colored forms which are encountered frequently, some in large numbers. Among these are a number of species of butterflies; of the Rhopalocera, the "Maack" swallowtail (Papilio maackii kurilensis Mats.) which usually feeds on flowers and which not infrequently forms large clusters on sections of damp soil, the "Stubbendorf" swallowtail (Parnassius stubbendorfi doii Mats.), the green-veined white (Pieris napi nesis Fruhs.), several representatives of the fam. Nymphalidae (pearl butterflies Argynnis aglaja chishimensis Mats., A. laodice ferruginea Mats., A. paphia neopaphia Fruhst., the pestrokryl'nitsa bureinskaya Araschnia burejana Btl.), the "Chinese" ?? satyr (Ypthima argus jezoensis Mats.), "blues" (Cyaniris argiolus ladonides De l'Orza and Chrysophanus phlaeas daimio Seitz), the common skipper (Parnara pellucida Murr.); of the nocturnal butterflies, the "Artemis" giant silkworm (Actias artemis Brem.), a tiger moth (Parasemia plantaginis kunashirica Bryk), which is a common species on nearly the entire Kuril Range with the exception of Iturup, the "sphinx" [?? elephant] (Haemorrhagia fusciformis affinis Brem.), a geometer (Cystidia stratonice Cr.), Mimeusemia persimilis Btl. (of the fam. Agaristidae), and some others.

Representatives of other orders as well, in particular the following, have been assigned to the landscape species: some cicadas (Tibicen bihamatus Motsch., Aphrophora alpina Mel., A. intermedia Uhl.), a wingless locust (Parapodisma mikado Bol.), a earwig (Anechura harmandi Burr.), bugs (Carbula humerigera Uhl., Elasmostethus humeralis Jak., Elasmucha signoreti Scott, Palomena angulosa Motsch., Acanthosoma spinicolle Jak.), many species of beetles, including the dung beetle (Geotrupes laevistriatus Motsch.), chafers (Sericania sachalinensis Mats., Ectinohoplia rufipes Motsch., Popillia japonica Newm., Mimela flavilabris Waterh., Anomala cuprea Hope., A. lucens Ball., A. rufocuprea Motsch.), scarabaeid beetles (Gnorimus subopacus Motsch. and Cetonia roelofsi Har.), long-horned beetles (Leptura scotodes Bat., L. succedanea Dalm., Judolia cometes Bat., Strangalia arcuata Panz., S. ochraceofasciata Motsch., S. aethiops Poda, S. vicaria Bat., Leontium viride Thoms., Chlorophorus japonicus Chevrol., Allosterna tabacicolor bivittis Motsch.) which constantly cluster en masse on the flowers of various plants, especially of the carrot family, leaf-beetles (Basilepta balyi kurilensis L. Medv., Gastrolina peltoidea Gebl., Linaeidea aenea L.), and soldier beetles (Themus cyanipennis Motsch. and Athemus suturellusMotsch.); among dipterans, mainly representatives of the fam. Syrphidae (Volucella jeddona Big., V. pellucens tabanoides Motsch., Matsumyia nigrifacies Shir., Cheilosia impressa Lw., Ch. yesonica Mats., Sphegina violovitshi Stack., Chrysotoxum subbicinctum Viol.), as well as Laphria rufa R÷der and other species of the fam. Asilidae.

We shall limit the enumeration of the landscape species of the broadleaved forests of Kunashir at this point, although the list of them could be substantially augmented by particular representatives of the dipterans, hymenopterans, dragonflies, lepidopterans, and other orders. It is significant that among the species cited more than 70% are representatives of the Palaearchaearctic fauna, mainly Japanese-Manchurian and Island species, and Boreal species, widely distributed within the limits of the Palaearctic, account for less than 30%. Such a ratio is especially characteristic for the coniferous-broadleaved forests of the west coast of Kunashir, where southern forms primarily appear as background species, while the boreal elements occupy a subordinate position. This ratio changes appreciably in the direction of an increase in the role of the Boreal species in the coniferous-broadleaved forests of Iturup and even in the central part of Kunashir; we will dwell on this in greater detail below.

Southern forms, whose development is associated with the soil, also are provided an opportunity to multiply in fairly large numbers on the west coast of Kunashir, due to the favorable soil and microclimatic conditions which have developed there. These are the already-mentioned cicada, a number of species of lamellicorns (chafers, scarabaeid beetles), stag beetles (Lucanidae), some species of clickbeetles (Athous inornatus Lew., Dolopius lewisi Fleut., Ectinus candezei Lew.), many orthopterans, including a long-horned grasshoper (Metrioptera japonica Bol.), a mole cricket (Gryllotalpa africana Palis.), the "Japanese pygmy grasshopper" [??] (Tetrix japonica Bol.), the locusts (Parapodisma mikado Bol., Podismopsis genicularis Shir.), Chorthippus biguttulus maritimus Mitsch., and the solitary form of the Asiatic locust, Locusta migratoria migratoria L.

Besides the background species, which numerically predominate over other species, and which, more than anything else, determine the appearance of each entomological complex, the presence of particular representatives of more southern faunas, of Japan and China, is typical for the entomofauna of the broadleaved forests of Kunashir. These southern species are in nearly all orders of insects, but in the majority of cases they are rare, are encountered singly, and it is far from every year that it is possible to find some of them. It is entirely evident that the coniferous-broadleaved forests of Kunashir serve as the northern region of their range. Many can be cited as examples; we will limit ourselves just to some. The following belong to such southern species: leafhoppers of the tropical tribe Otiocerini fam. Derbidae, Epotiocerus flexuosus Uhler and Mysidioides sapporoensis Mats. (Anufriev, 1968a, 1968b), a number of Manchurian bug species (Urostylis annulicornis Scott, Acanthosoma expansum Horv., A. labiduroides Jak., Dinorhynchus dubowskii Jak., etc.), many species of beetles, ground-beetles (Lebidiaoctoguttata A. Mor., Parena piceola Chd.), stag beetles (Lucanus maculifemoratus Motsch., Eurytrachelus rectus Motsch. and other representatives of this Indo-Malaysian genus), Syntelia histeroides Lew., species of the tropical family Languriidae (Languriomorpha lewisii Crotch., L. kunashiriana Kno), such typically Japanese species of the fam. Melandryidae, as Phloetrya bellicosa Lew. and Ivania coccinea Lew., and a number of southern species of long-horned beetles (Eustrangalis distenoides Bat., Dihammus fraudator Bat., Pterolophia jugosa Bat., Asaperda meridiana Matsush., Eryssamena tuberculata Pic, Exocentrus testudineus Matsush., Mesosa senilis Bat., Cagosima sanguinolenta Thoms., Glenea relicta Pascoe, etc.). Even such examples are sufficient to imagine the degree of saturation of the coniferous-broadleaved forests of Kunashir with southern forms. But, a fair number of boreal Palaearctic species are encountered, along with southern elements, in the composition of the entomofauna of these forests, which, by contrast with similar forests of Sakhalin and other islands of the archipelago located further north, here prove to be sparse or rare.

Let us dwell on some features of the fauna of the forests in question which are archaic from our point of view. It was already noted in section 2 of this chapter that the rather high temperatures and humidity on the west coast of Kunashir support the existence in this region of many heat- and moisture-loving species of plants and animals. In particular, the large numbers of some moisture-loving species of animals strike the eye, for example, the woodlice (Isopoda, Oniscoidea), which are encountered en masse everywhere under the forest canopy and which openly crawl on the trunks of trees, clustering in bark fissures, on stumps, and on the ground, as well as terrestrial molluscs which are regularly encountered on the ground, trees, and bushes. Some insects also form such clusters: barklice (Copeognatha, Psocidae), large and dense groups of which can often be seen on the bark of stout leafed trees; an earwig (Anechura harmandi Burr.) which occupies the same eco-areas as the woodlouse, and which is encountered in large numbers not only under the forest canopy, but also in forest clearings, on the flowers of herbaceous plants; and spittlebugs (Aphrophoridae), whose larvae occupy all tiers of the forest vegetation, from the grassy cover to the high crowns of trees.

It is precisely on the west coast of Kunashir that leeches which are apparently capable of surviving for a prolonged period of time without water, using only atmospheric moisture, have been found on the ground. This fact points to a [temporally] remote similarity of the ecological conditions of the Kunashir broadleaved forests and tropical forests in which leeches typically live on trees (Panfilov, 1961). A tree frog (Hyla japonica Gunther.), a representative of amphibians, mainly distributed in southern subtropical and tropical forests is known only in those forests (Terent'ev, Chernov, 1949). Three species of rat snakes of Japanese origin, live in the broadleaved forests of Kunashir: the "island rat snake" (Elaphe dimacophora Boie.), the "small-scaled rat snake" (E. quadrivirgata Boie.), and the "Maki Japanese climbing rat snake" (E. japonica Maki.). These are most often encountered in the region of the caldera of Golovnin Volcano and in the environs of Alekhino set., and on the east coast stay mainly near hot springs (env. of Goryachiy Plyazh set.), from whence the majority of specimens kept in the collections of the Zoological Institute of the USSR Academy of Sciences were collected (Pereleshin, Terent'ev, 1963). The "small-scaled rat snake" is the more common of these; the other two species are encountered more rarely. A brightly colored lizard, the Far-Eastern skink (Eumeces latiscutatus Hallow.), which belongs to a tropical genus distributed in North and Central America, Africa, and South East Asia, also lives in the environs of Alekhino set. With regard to birds, V. A. Nechaev (1969) characterizes the ornithological complex confined to the broadleaved and mixed forests of Kunashir as rich in species and having mainly a Sino-Japanese appearance.

The concentration of a large number of southern species of plants and animals is apparently determined not only by the high temperature and humidity, but by an entire complex of specific ecological conditions existing in these forests, which have been preserved in relatively limited areas in the southern part of Kunashir. For it is difficult to assume that all southern forms migrated there from Japan and then were confined in their distribution only to a small territory occupied by broadleaved forests. It is more probable that an indigenous flora and fauna are existing here, which have come down to our times in markedly altered and impoverished form, but which have preserved until the present time particular features of ancient biogeocenoses of thermophilous broadleaved forests that predominated on this segment of the dry land in the Tertiary. The facts under consideration attest to the relict character of the biogeocenosis of the Kunashir coniferous-broadleaved forests.

The entomofauna of the broadleaved and coniferous-broadleaved forests of Iturup contrasts appreciably with that of Kunashir, as does the character of the forests themselves as well. On Iturup, the broadleaved forests do not have so complex a composition as on Kunashir, and do not achieve such luxuriance (Figure 41). They are encountered in the central portion of the island and along its west coast up to the Chirip Peninsula; they are represented on the coastal slopes mainly by elfin woods.

The appearance of the entomofauna of this natural complex is quite different; it is not distinguished by the diversity which was observed in the Kunashir forests. Although the number of southern species here is still fairly high, they lose their leading role and yield it to representatives of the boreal fauna. The overwhelming majority of background species on Iturup are different from those on Kunashir, and their number is markedly diminished, since the southern forms that are present usually do not achieve high numbers. Complexes of species associated with birch, mountain ash, bird cherry, and Sakhalin cherry, some of which were not observed on Kunashir or were encountered there in small numbers (see Table 19), here are more distinctly prominent. Such southern species of butterflies as Actias artemis Brem., Cystidia stratonice Cr., Mimeusemia persimilis Btl., Ypthima argus jezoensis Mats., Araschnia burejana Btl., and many others which on Kunashir are landscape species are absent on Iturup. The extreme paucity and impoverishment of both diurnal and nocturnal butterflies, especially by comparison with the coniferous-broadleaved forests of Kunashir, is striking overall. The blues (Lycaenidae) and tiger moths (Arctiidae) are exceptionally rare; Maack, Bianor, and common swallowtails are encountered sporadically and in small numbers, just as are representatives of the Nymphalidae, Satyridae, Sphingidae families and some others. In essence not a single species of the butterflies on Iturup can be regarded as a landscape species, with the exception, perhaps, of individual leaf-rollers, geometers, and the Gaschkewich's satyr (Neope gaschkewischi [sic] Mn.) which is associated with the Kuril bamboo, and for that reason penetrates nearly all forest biotopes. Two species of white butterflies (Pieris napi nesis Fruhs. and Colias erata poliographus Motsch.) and the pearl butterfly (Argynnis aglaja chischimensis Mats.), as well as butterflies associated with the Kuril bamboo (Neope gaschkewitschi [sic] Mn., Lethe diana Btl., Halpe varia Murr.) are encountered relatively frequently.

The crickets (Grylloidea), southern species of cicadas of the subfamilies Cicadidae, Derbidae, and Cercopidae, the scorpion fly (Panorpa pryeri Mac Lach.), which is common on Kunashir, some families of bugs (Tingidae, Reduviidae, Rhopalidae, etc.), beetles (Cleridae, Erotylidae, Languriidae), and other insects are absent here. The long-horned grasshoppers (Tettigonioidea) and the locusts (Acridoidea) are extremely poorly represented; the same is true, of the bugs - the shield bugs (Acanthosomatidae and Pentatomidae), of the beetles - the stag beetles (Lucanidae), chafers (subfamilies Rutelinae, Trichiinae, Cetoniinae), the lycids (Lycidae), false darkling beetles (Melandryidae), etc., many species of which are common and even background on Kunashir.

The following have been distinguished as landscape insects in the broadleaved forests of Iturup: beetles - the leaf-beetle (Clitena fuscipennis Jac.), a soldier beetle (Podabrus marginellus Motsch.), long-horned beetles (Allosterna tabacicolor bivittis Motsch., Strangalia aethiops Poda, Paraclytus excultus Bat.), a borer (Agrilus ignoratus Obenb.), the seven-spotted ladybeetle (Coccinella septempunctata bruckii Mulsant); dipterans - a horsefly (Hybomitra montana Meigen), flower-flies (Syrphus ribesii L., S. balteatus Deg., Eristalis cerealis F., Helophilus sapporensis Mats., etc.); some species of aphids (the "bird cherry-oat aphid" Rhopalosiphum padi L., the "black cherry aphid" Myzus cerasi F. and Dactynotus jaceae L.); leafhoppers (Aphrophora alni Fall. and Philaenus spumarius L.); and bumblebees (Bombus japonicus Fr. et Wag. and B. konakovi Panf.), of which the former is widely distributed throughout the entire Kuril Range. Among the landscape species on Iturup, southern Palaearchaearctic elements now constitute only 40%, while boreal Palaearctic elements constitute 60%.

It is characteristic that the mass clustering of moisture-loving species of insects and crustaceans which we saw on Kunashir is not observed in the sparser coniferous-broadleaved forests of Iturup. Reptiles and amphibians are absent here. According to V. A. Nechaev (1969), the ornithofauna is also represented by an impoverished forest complex, from which a number of southern species of birds have fallen out.

Groupings of insects associated with particular broadleaved species on the Kuril Islands continue to remain far from adequately studied; this is the subject of special investigations. However, the materials we have collected may provide a general notion of such entomological complexes, and also may underscore the difference and similarity of these complexes on Kunashir and Iturup. Therefore, we will present brief lists of them (Table 19), although many species of butterflies and sawflies whose caterpillars and larvae were encountered on leaves have not been included in them, since it was quite often impossible to establish the species affiliation on the basis of the larval phase. Despite this, the complexes presented reflect connections with the corresponding groupings of insects on the continent, attest to substantial impoverishment and at the same time the originality of the Kuril complexes as compared with the continental, and also point to the presence in their composition of ancient species associated with relict plants. Let us dwell in somewhat greater detail on these groupings.

The entomofauna of the oak (Quercus). First and foremost, by comparison with the diverse entomofauna of the oak stands of Primorskiy Kray, the marked poverty of the Kuril complex of insects associated with the oak is striking. The majority of butterflies which develop on oak in the Primor'ye and Priamur'ye (Neptis thisbe Mn., Nyphanda fusca Brem., Zephyrus brilliantina Stg., Z. saphirina Stg., Dendrolimus undans fasciatella Mn., Rhodinia fugax diana Obert., Antheraea pernyi Quer., Ephesia dissimilis Brem.), and many others (Kurentsov, 1939, 1965a) are absent on the islands. Those of the polyphages which would be able to develop on oak (Actias artemis Brem. and Dictyoploca japonica Btl.) prefer other plants under Kuril Islands conditions: the former prefers birch and the latter, alder (Konovalova, 1968). There is not a single one of the 12 species of long-horned beetles which are among the entomofauna of the Mongolian oak on the continent (Shabliovskiy, 1968) on the Kuril Islands. And long-horned beetles have not been found here on the oak in general. The same can be said of the bark-beetles and other insects. At the same time, species which suggest the existence of associations between the entomofauna of oak stands of the islands and of the continent are encountered in the broadleaved forests of Kunashir and Iturup. These are a bug, Urostylis annulicornis Scott, of a primitive family of shield bugs (Urostylidae), a metallic wood-boring beetle (Agrilus ignoratus Obenb.), a leaf-beetle (Aphthona perminuta Baly), and a blue (Zephyrus lutea Yew.), which are typical representatives of the Manchurian fauna, trophically associated with the Far-Eastern oaks.

The existing differences of the island entomofauna of the oak from the continental fauna are explained by a number of factors, evidently including, in particular, the different species composition of the oak stands themselves. It is worth remembering that the oak stands in the continental Far East are mainly formed of the Mongolian oak (Quercus mongolica), while on the Kurils they are formed of the shallow-cup Mongolian oak (Q. crispula), which is morphologically more ancient. The daimyo oak (Q. dentata) is encountered rarely in the Primor'ye and on the Kurils and is not significant as a forest-forming species.

The group of amphipalaearctic species which has a range that is substantially interlinked with the range of the genus Quercus is of great interest. To these have been assigned a butterfly, Stauropus fagi persimilis Btl., which is distributed in the Far East (the south of Primorskiy Kray, Iturup, Japan), in the Transcaucasus, and in broadleaved forests of European USSR, but which is absent in Siberia, and 2 species of aphids, Lachnus pallipes Hart. and Tuberculoides annulatus Hart., of which the former is common in European USSR and was recently found by us in Primorskiy Kray and on Kunashir, while the latter is known in Europe (Western and Eastern), West Kazakhstan, Central Asia, Kunashir, North America, and Australia. Both species reveal a discontinuity of range in Siberia; T. annulatus Hart. has evidently shifted to feeding of other plants in West Kazakhstan and Central Asia, where the oak is absent. E. M. Danzig (1968) has observed a very similar distribution interlinked with the oak in some genera of scale insects [Trans. note: this term in Russian may also be translated as "mealybugs", and therefore it is probably necessary to consult the original reference to determine which term is correct in this context], and V. I. Kuznetsov has observed the same (1968a, 1968b, 1969) in leaf-rollers which have corresponding vicariants in European oak forests which are associated in this case with a different species of oak (Quercus robur). This discontinuity of range in Siberia attests to its relict character, both for the oak itself (Sokolov, Svyazeva, 1965) as well as for the insects which accompany it, and which are evidently derivatives of a Tertiary fauna.

The entomofauna of the maple (Acer). The island complex of insects which live on maple is also somewhat impoverished as compared with the continent, mainly due to the absence of a number of species of butterflies and other leaf-gnawing insects. Among the latter only 2 species are distinguished by their numbers: a leaf-beetle (Clitena fuscipennis Jac.) on Iturup, and a butterfly (species not established), whose caterpillars have been encountered en masse on maple leaves on Kunashir.

Aphids which are associated with maple comprise a highly interesting group. All 3 species of aphids which we collected on the islands are distinguished by a discontinuous amphiboreal range, which repeats the modern range of Palaearctic species of the genus Acer to a substantial degree (Sokolov, Svyazeva, 1965):

Judging by their range, within the limits of which various species of maples also flourish, all of these aphids are oligophages. In the past (apparently as far back as the beginning of the Pleistocene), when the genus Acer had an unbroken distribution in the moderate zone of Eurasia (Poyarkova, 1933), the aphids associated with it also evidently had an unbroken range, later disrupted in connection with the disappearance from Siberia of their food plants that had retreated southward under the influence of the cold episode which had set in. In that case, just as with the "oak" aphids, which occupy an analogous range, we are apparently dealing with ancient forms linked by their roots with the Tertiary fauna.

With regard to xylophages associated with the maple, they form a characteristic group on the Kuril Islands, which consists primarily of Island, Ussuri-Island, and partially, typically Manchurian species. The bark-beetles (Trypodendron niponicum Blandf. and Scolytoplatypus daimio Blandf.) are distinguished by a broad polytrophism. Within the limits of their range, they have adapted to life on plants which belong to various families, but which are endemic for Far Eastern and sometimes only for Island flora. In particular, Trypodendron niponicum Blandf. lives on the Kurils on Alnus maximowiczii and Betula ermanii, in addition to the maple (Krivolutskaya, 1965a); on Sakhalin on Ulmus propinqua as well (Krivolutskaya, 1958); and in the Southern Primor'ye, on Tilia amurensis and Betula costata (Kurentsov, 1941). The Island Scolytoplatypus daimio Blandf. has a no less broad circle of food plants, including Acer pictum, Cerasus sachalinensis, Alnus maximowiczii, Cornus controversa, and Fraxinus mandshurica on the Kurils, and Juglans sieboldiana on Sakhalin. The trophic associations enumerated and the limited range of these bark-beetles suggest their probable ancient confinement to distinctive, multi-species mixed forests of the Manchurian type.

By contrast with the bark-beetles, the horntails which live on maple (Euxiphydria potanini A. Jac. and Xiphydria ogasawarai Mats.) reveal a narrower food specialization. The former of these may develop on Erman's birch as well, and the latter only on maples. It is significant that both species are associated on the islands with Acer pictum, while on the continent they are associated with A. mono, i.e., with the most primitive representatives of the genus Acer, close to its prototype (Poyarkova, 1933). Xiphydria ogasawarai Mats., according to the observations of V. K. Stroganova (1968), is adapted to habitation in high levels of humidity; as a result it keeps to shaded sites in Primorskiy Kray, typically under the forest canopy, and avoids open spaces. This ecological characteristic is, evidently, testimony to the adaptation of the species to habitation in damp forests, which was elaborated in the process of its evolution. Taking all of the above into account, it can be concluded that both horntails are distinguished by their archaic character and belong to the group of Ussuri-Island autochthons.

Of the three species of long-horned beetles which develop on maples in the continental region of the Far East (Shabliovskiy, 1968), there is not a single one in the broadleaved forests of Kunashir and Iturup. Evidently, these long-horned beetles are associated on the continent with other species of maples and, it must be assumed, this is the cause of their absence on the islands.

The entomofauna of the elm (Ulmus). The family Ulmaceae, as is the case for the beeches (Fagaceae) and the maples (Aceraceae), is characterized by a discontinuous range in the moderate zone of Eurasia, and is regarded as a relict family (Sokolov, Svyazeva, 1965). It is represented in the Far East by four species, of which only two (Ulmus laciniata and U. propinqua) flourish on Kunashir, and singly on Shikotan. However, the elm stands occupy only limited sections in the southwest of Kunashir, where U. laciniata predominates. Therefore, all insects associated with the elm (mainly with U. laciniata) are also localized in the broadleaved stands of Kunashir and are absent on Iturup, with the exception of particular polyphages.

The Kuril entomofauna of the elm has features similar to those of the entomofaunas of the oak and maple just considered. It is appreciably impoverished by comparison with the continental and even Sakhalin elm entomofauna, consists in the overwhelming majority of Manchurian and Island species, and the more widely distributed Palaearcts included in it often have disjunctive ranges. For example, the aphid Byrsocrypta hirsuta Baker., which clearly has an East Asian origin and is widely distributed in the southern regions of East Asia, Japan, on Kunashir, and in the south of Primorskiy Kray, is absent in Siberia, and appears again in Central Asia (Uzbekistan and Kirghizia), but does not penetrate further to the west. Evidently in the past, when the elm was unbrokenly distributed in Eurasia, the range of this aphid was limited to the boundaries of Siberia, and after the displacement of the boundaries of the range of Ulmus to the south, this aphid was preserved only in Central Asia and in the Far East (Shaposhnikov, 1955; Krivolutskaya, Ivanovskaya-Shubina, 1966). The range of another aphid, B. ulmi L., entirely coincides with the contemporary distribution of the elm. It is known in the Far East (Kunashir, Japan, the south of Primorskiy Kray, China), in Central Asia, and Europe, but is absent in Siberia; it forms a special subspecies, B. ulmi jezoensis Mats., on the islands.

The leafhopper Jassus lanio L., which in various regions of the range has adapted to feeding on various broadleaved species which have a disjunctive range, is also distinguished by a discontinuous distribution. In the Far East it lives on the elm (Anufriev, 1970), in European USSR on the oak (Emel'yanov, 1964), and in Siberia, apparently on the linden, since it is known that other representatives of the genus Jassus are associated with the linden, the only one of the broadleaved species which is preserved on limited areas in the south of Siberia (Tolmachev, 1962). It must be assumed that these species, just as is the case with their food plants, are Tertiary or early Quaternary relicts.

Of the xylophages recorded on the elm (see Table 19), only a polyphage long-horned beetle (Allosterna tabacicolor bivittis Motsch.) is widely distributed in the Palaearctic; the remaining species are Manchurian or Island autochthons. Among these, a group of Island species is quite conspicuous; this group includes long-horned beetles Mesosa japonica Bat., Pterolophia jugosa Bat., and Glenea relicta Pascoe, and a bark-beetle Hylesinus elatus Niiji.

It is curious that a bark-beetle (Ernoporicus spessivtzevi Berg.), which in Primorskiy Kray lives on the ash, lives on the elm under the conditions of the Kurils (Kurentsov, 1941). A second bark-beetle, Hylesinus elatus Niiji., which multiplies en masse on the elm on Kunashir and Hokkaido, is supplanted in the Primor'ye by a close species, H. costatus Blandf. (Krivolutskaya, 1968), which lives on ash. Overall, the bark-beetles of the genus Hylesinus are associated almost exclusively with the ash, and their migration to other plants is an extremely rare phenomenon. The causes which elicit this in this case become clear following an analysis of the Kuril ash entomofauna.

The entomofauna of the ash (Fraxinus). The ash grows only in broadleaved forests of Kunashir, but here it is sparse and does not play an appreciable role in the composition of the stands. On the basis of the character of the range, it, like many other broadleaved species, is regarded by botanists (Sokolov, Svyazeva, 1965, etc.) as a relict plant. The specific entomofauna of the Manchurian ash (F. manschurica [sic]), which is so richly represented in the Primor'ye and the Priamur'ye (Kurentsov, 1939, 1956; Lyubarskiy, et al., 1961), as well as in Japan, and which contains a large number of species of various groups of insects, is extremely impoverished on Kunashir. Here the ash, as it were, loses all of the insects which are characteristic of it, including the bark-beetles, as was referred to above. It is characteristic that on Sakhalin as well not one bark-beetle species is found on the ash, although in general its entomofauna is much richer there. A number of species of leaf-beetles (Basilepta fulvipes Motsch., Gastrolina peltoidea Gebl., Clitena fuscipennis Jac.), which usually damage the ash on the continent, also prefers to live on other plants, the alder and the maple, in the Kuril conditions. A fairly small group of insects which develop on the ash in the forests of Kunashir consists mainly of polyphagous species which have migrated to it from other woody plants. The conclusion that the ash disappeared from the composition of the broadleaved forests of the Kuril Islands for a prolonged period at a certain stage of geological history and appeared here again relatively recently, after insects characteristic of it had adapted to life on other plants, suggests itself from an analysis of all of the facts presented. It is entirely probable that it was secondarily introduced here by man. This conclusion also relates to a certain degree to the "ash" entomofauna of Sakhalin.

The entomofauna of the cork tree (Phellodendron). The entomofauna of the Sakhalin cork tree is represented by an impoverished complex of Manchurian species, by comparison with the Amur cork tree, although it does have certain specific characteristics. If on the continent the principal pests of the cork tree are leaf-gnawing (the butterflies Papilio bianor Cr., P. xuthus L., the leaf-beetles Basilepta orientale Jac., Chrysomela guttata Gebl., Melasoma cupreum Fabr.) and sucking insects (a plant louse Calophya nigra Kuw.), and a long-horned beetle Mesosa myops Dalm. is of secondary significance (Lyubarskiy, 1952; Kurentsov, 1956; Konovalova, 1966a), only Papilio bianor Cr. and P. maackii kurilensis Mats. remain of this complex on the Kurils, while a bark-beetle (Phellodendrophagus elegans Krivol.) which has been found on the continent only recently on the Manchurian aralia (Krivolutskaya, Kupyanskaya, 1970) is becoming the main pest of the cork tree.

The entomofauna of the sen (Kalopanax). The complex of species associated with the sen is in general highly limited and is still inadequately studied. It consists mainly of polyphages both on the islands and on the continent. Only one species, a bark-beetle Eocryphalus semenovi Kurenz., which is encountered in the Primor'ye, on Sakhalin, and in the Southern Kurils is specific for this species.

The entomofauna of the magnolia (Magnolia). The white-leaf Japanese magnolia is one of the most ancient broadleaved species present in the broadleaved forests of Kunashir. By contrast with the evergreen representatives of the genus, this form is deciduous and has a limited range within the limits of Kunashir and the northern region of Japan. The insects which are found on magnolia form a fairly small and at the same time distinctive grouping (see Table 19), consisting almost entirely of Island, primarily Japanese, species. All of them are polyphages which have trophic associations of various latitudes, but which are at the same time restricted to life on relict or tropical plants. It is significant that not one monophage has been observed on the magnolia. V. I. Kuznetsov (1969), analyzing the alimentary associations of the leaf-rollers, reaches the conclusion that caterpillar polyphagy is characteristic of archaic tribes ofTortricidae, developed in them as an adaptation to life in multi-species forests, and is at the present time characteristic of relict biogeocenoses. D. V. Panfilov (1961), in his turn, observes polyphagy as a characteristic of the entomofauna of tropical forests. Evidently an analogous phenomenon occurs in this case, and the insects associated with the magnolia must be regarded as a relict complex, formed under the conditions of moist coniferous-broadleaved forests, complex in composition, which predominated on the eastern edge of Eurasia in the Tertiary. To some degree this entomological complex has been preserved until the present time.

The entomofauna of the Rosaceae (Rosaceae). The rose family is represented on the Kurils by a large number of herbaceous, bushy, and woody species. Let us dwell on the better studied groupings of insects which are associated with the woody Rosaceae, the cherry (Cerasus), the bird cherry (Padus), and the mountain ash (Sorbus), which are widely distributed in coniferous-broadleaved forests. These species belong to a group of ancient Rosaceae, fossils of which are known from the Paleogene (Vasil'yev, 1958). Each of these species is represented on the archipelago by two species with fairly narrow ranges, an Island species (Cerasus kurilensis, Padus ssiori) or one which is limited to the islands and the coastal region of the continent (Cerasus sachalinensis, Sorbus sambucifolia, S. commixta, Padus maximowiczii).

The largest number of insects in the coniferous-broadleaved forests of Kunashir and Iturup live on the cherry, and the smallest number on the mountain ash. Insects living on leaves (aphids, leafhoppers, bugs singly, leaf-beetles, butterflies singly) constitute a large group. Some of these, for example the aphids and leafhoppers, are the oligophages, adapted to life on various species of one genus of plants, on cherries, mountain ashes, or bird cherries. These are mainly Palaearcts or still more widely distributed species which may be regarded as cosmopolites. Two new species and 1 genus of aphids described from the Southern Kurils on the basis of our collections from the cherry constitute an exception, but it is difficult thus far to make a judgment regarding the latitude of their range and the specificity of their alimentary associations. With regard to the beetles which are included in this group (leaf-beetles, leafrollers, etc.), these are distinguished by broad polyphagy, and migrate to the Rosaceae from other plants, often remote from the systematic point of view. Thus, specialized Island forms are not found among the phyllophages of the Rosaceae.

The bark-beetles are of interest in this connection. They form a fairly distinct group of intergenus oligophages, whose range is limited to the islands. Polygraphus nigrielytris Niiji., which is known on Sakhalin, the southern part of the Kuril Range (all the way to Urup), Kamchatka, in Japan, and on the Korean Peninsula, turns out to be the most widely distributed of these, capable of developing with the equal success on all of the enumerated species. The second species, P. ssiori Niiji. has been observed only on Iturup, where it is quite common on the Kuril cherry; in Japan (Hokkaido, Honshu, Sikoku), it damages fruit trees (Kuwayama, 1967). It is characteristic that among the bark-beetles of the genus Polygraphus, which are mainly associated with the coniferous species, only 3 species develop on deciduous Rosaceae, and in particular on the woody Rosaceae mentioned. In the Far East these are the Island P. nigrielytris Niiji. and P. ssiori Niiji.; in Europe, P. grandiclava Thoms. The latter must evidently be regarded as a vicariant of P. nigrielytris Niiji. (Krivolutskaya, 1958). These species are phylogenetically younger than the other representatives of the genus Polygraphus, which was originally associated with coniferous species, since they undoubtedly migrated to the Rosaceae from more ancient coniferous species. At the same time, a number of facts suggests the Tertiary age of these beetles: the Island range of P. nigrielytris Niiji. and P. ssiori Niiji., the presence of a vicariant in Europe, the intergenus oligophagy, and the adaptive adjustments to island conditions of existence in P. ssiori Niiji. (see section 2 of the present chapter).

A third bark-beetle, Cryphalus padi Krivol., is associated on Sakhalin with the bird cherry, and on the Kurils with the mountain ash. Finding it on the cherry tree as well cannot be excluded. In Primorskiy Kray it has been displaced by a very close species C. pruni Egg., which lives on a bird cherry (Padus asiatica), the Manchurian apricot (Armeniaca manshurica), the Ussurian plum (Prunus ussuriensis), and the mulberry (Morus alba). These species have become separated relatively recently, after the separation of the islands from the continent had already occurred, which is indicated by their large morphological and ecological similarity, and are the result of a later process of species formation.

The lists of insects associated with particular broadleaved species that we have presented (Table 19) hardly exhaust the entire diversity of the entomofauna of the broadleaved forests; we cannot attempt to do this here. An analysis of the entomofauna of the principal broadleaved species shows an appreciable impoverishment of all of the island groups of insects considered, demonstrates ancient associations between corresponding ecological groupings on the islands and the continent, and also identifies the presence in nearly all of these of ancient autochthonous elements.

The entomofauna of the dark coniferous forests. The dark coniferous forests, as has already been indicated, grow in the southern part of the Kuril archipelago, on the islands of Kunashir, Shikotan, and Iturup (Figures 42, 43). They are formed in the main by the Sakhalin fir and the Sakhalin spruce; here and there the Sakhalin spruce and the Japanese yew appear in their composition, but the significance of the latter [two] as forest-forming species is fairly small. The dark coniferous forests essentially do not form pure stands. The birch and broadleaved species, the proportions of which is variable at different sites, as are those of the coniferous species, are commonly encountered in the composition of the dark coniferous forests. In the past, the dark coniferous forests were more widely distributed on the islands, but their areas have been substantially reduced, and their composition altered by both clear-cutting and selective felling.

The entomofauna of the dark coniferous forests is not so diverse as that of the broadleaved forests, although these two entomological complexes coexist in close proximity to one another as a result of the growth jointly of the dark coniferous and broadleaved species. In essence, they comprise only parts of a unitary entomofauna of the biogeocenosis of the dark coniferous-broadleaved or broadleaved-dark coniferous forests. But, depending on the predominance in the stands of various species, the basic background of the entomofauna varies. Insects associated with broadleaved species predominate in the dark coniferous-broadleaved forests, while in the broadleaved-dark coniferous forests, those associated with the dark coniferous species predominate. The latter predominate on the territory of Kunashir (apart from its southern and southwestern regions), and in the south of Iturup and on Shikotan. The number of landscape species of insects in the dark coniferous forests is not so great (Figure 44). On different islands and in particular areas within the limits of the same island, different species appear as landscape species. For example, the "white-striped silkworm" (Dendrolimus superans albolineatus Mats.), some geometers (Boarmia ribeata Cl., etc.), and long-horned beetles (Leptura succedanea Lew., L. scotodes Bat., Strangalia arcuata Panz., S. vicaria Bat., Megasemum quadricostulatum Kr.) may be classified as landscape species of the dark coniferous forests on the western shore of Kunashir; a leafroller (Eurydoxa advena Fil.) and particular species of long-horned beetles and horntails may be classified as landscape species on the east coast of the same island; this is true of some long-horned beetles (Leptura succedanea Lew., L. scotodes Bat., to some extent Strangalia arcuata Panz.) on Iturup; on Shikotan, a geometer (Arichana melanaria L.) and long-horned beetles (Leptura succedanea Lew., to some extent Strangalia arcuata Panz., S. vicaria Bat., Megasemum quadricostulatum Kr., Monochamus grandis Waterh.).

The reasons for the limited number of landscape species in the dark coniferous forests as compared with the broadleaved forests reside in the following: first, the overwhelming majority of insects associated with the dark coniferous species are xylophages, and lead a concealed mode of life, and second, representatives of a Boreal fauna predominate in the complex under consideration, many of which are rare on the Kurils or do not multiply in large numbers. Apparently, it is for this reason that Manchurian and island species also are landscape species in dark coniferous forests, although they occupy a subordinate position in the overall composition of this grouping.

The complexes of the entomofauna of the dark coniferous forests on different islands are not equivalent. The maximal number of species have been recorded on Kunashir (54), and approximately in equal numbers on Shikotan (22) and Iturup (23). But on Iturup the entomofauna of the dark coniferous forests has a more northern cast. Such typically taiga elements as Nivellia sanguinosa Gyllh., Judolia sexmaculata L., Oedecnema dubia Motsch., and Rhagium inquisitor rugipenne Reitt. of the long-horned beetles and Polygraphus poligraphus L., Crypturgus pusillus Gyll., and C. hispidulus Thoms. of the bark beetles are present in its composition, and a number of representatives of the Palaearchearctic fauna (Strangalia vicaria Bat., Megasemum quadricostulatum Kr., Monochamus grandis Waterh., Dryocoetes striatus Egg., and some others) which are characteristic for Kunashir and Shikotan are absent. At the same time, the principal background is created by Manchurian species (Leptura succedanea Lew., L. scotodes Bat., Asemum amurense Kr., Polygraphus proximus Blandf.), which multiply in larger numbers than the Boreal Palaearctic species on Iturup as well, just as on Kunashir and Shikotan.

The groupings of insects associated with dark coniferous species (aphids, long-horned beetles, bark beetles, and horntails) have been elucidated in considerable detail in special sections (see Chapter 3); therefore there is no sense in repeat those listings. Here let us only summarize briefly these data and dwell on the specific characteristics of the entomofauna of the dark coniferous forests as a whole. The Japanese yew (Taxus cuspidatus), by contrast with other dark coniferous species, is almost never colonized by insects. Not a single species of the xylophages is found on it, and single butterfly caterpillars are very rarely encountered.

A fairly large group of insects, numbering more than 50 species (4 species of aphids, 17 of long-horned beetles, 18 of bark beetles, 6 of horntails, and 4 of butterflies²) live on spruce under the conditions of the Kurils. Some of these, the black arches moth [??] (Ocneria monacha L.), the "large conifer" long-horned beetle (Monochamus urussovi Fisch.), and a timber beetle (Scolytoplatypus tycon Blandf.) are broad polyphages which migrate to leafed species. But, the overwhelming majority (47 species) are oligophages, capable of developing on a number of coniferous species, although the spruce nevertheless remains the favorite for many species. This suggests a primordial association of such species with the spruce. Their migration to other coniferous species was probably induced by a change in the structure of the biocenoses in the past, and could have been accomplished given the flourishing jointly of various coniferous species on the same territory, and given the high degree of ecological plasticity of the insects. There are no true monophages on the spruce, if one does not count species with insufficiently studied biology that may also turn out to be oligophages (the long-horned beetles Asemum punctulatum Bless. and Monochamus nitens Bat., and the bark beetle Trypodendron proximum Niiji.).
 

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² The number of species of butterflies associated with dark coniferous forests ought to increase somewhat, since we have not investigated the inhabitants of cones, and the sparse and rare species have not been studied sufficiently.

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Species possessing a wide, namely latitudinally-extended holarctic, transpalaearctic, or transsiberian range, are in the main characteristic for the entomofauna of the spruce. Species with a Manchurian and Island type of range, that have a meridional directionality, constitute less than 30%. Their distribution is linked to a substantial degree with the range of the spruces of the Omorica section (VasilÆyev, 1958). To these we assign long-horned beetles (Distenia gracilis Bless., Leptura succedanea Lew., Megasemum quadricostulatum Kr., Asemum punctulatum Bless., A. amurense Kr., Monochamus nitens Bat.), bark beetles (Polygraphus sachalinensis Egg., Cryphalus kurenzovi Stark, C. piceus Egg., Dryocoetes rugicollis Egg., Trypodendron proximum Niiji., Scolytoplatypus tycon Blandf.), and horntails (Xoanon mysta Sem., Urocerus antennatus Marl.). In the complex of the spruce entomofauna this group should evidently be regarded as phylogenetically more ancient, analogously to the entomofauna of the Schrenk spruce, which also has its original appearance and evidently a more ancient age (Kostin, 1964).

The complex of insects that live on fir, by contrast with the other coniferous species, has a specific character. Apart from the broad oligophages, it has within it a group of species associated nearly exclusively with the fir. This group includes aphids (Cinara piceae Panz., C. longipennis Mats.), long-horned beetles (Leptura scotodes Bat., Strangalia vicaria Bat., Monochamus grandis Waterh., Acanthocinus griseus F.), a borer (Niphades variegatus Roel.), bark beetles (Polygraphus proximus Bladf., Dryocoetes striatus Egg.), and, possibly, a horntail (Urocerus umbra Sem.). They constitute the principal nucleus of this faunistic complex and are distinguished by a range confined to the limits of the Palaearctic, extreme northeast Asia (the Okhotsk type of range), or just the islands. As a rule, insects which have adapted to life on fir do not migrate to other coniferous species. At the same time, the migration of insects usually associated with other species to the fir is limited to a certain degree. I. A. Kostin (1967) notes this characteristic for the bark beetles; it is also characteristic of some long-horned beetles, weevils and other trunk insects. The distinctive anatomical structure of the fir bark and wood, which is different from that of the spruce and cedar, the system of resin canals, and the different chemical composition of the resin and other substances (Ivanov, 1961; Treinis, 1961; Krivolutskaya, 1965b) are evidently a significant impediment to such migration. Of the oligophages, especially those which also frequent the spruce, the following are quite common on the fir under Kuril Islands conditions: long-horned beetles (Leptura succedanea Lew., Strangalia arcuata Panz., Megasemum quadricostulatum Kr.), bark beetles (Cryphalus kurenzovi Stark, Scolytoplatypus tycon Blandf.), the "white-striped silkworm/moth" (Dendrolimus superans albolineatus Mats.), a geometer (Boarmia angulifera Btl.), and horntails (Xeris spectrum L., Urocerus antennatus Marl.), and on the east coast of Kunashir, a leafroller (Eurydoxa advena Fil.).

The presence of exclusively eastern species associated with local dark coniferous species, which in fact determine its appearance, is characteristic for the entomofauna of the spruce-fir forests of the Kuril Islands overall, although widely distributed Boreal and taiga species proper participate significantly in this faunistic grouping. Such a composition of entomofauna reflects ancient connections of the Kuril dark coniferous forests with the corresponding forest formations of the continental Far East (the moist southern spruce stands and fir-broadleaved forests of Sikhot-AlinÆ), with the Siberian dark coniferous taiga, as well as with the mountain dark coniferous forests of Japan, Kamchatka, and North America.

In summarizing the above, we may briefly dwell on the distinctiveness of the entomofauna of the coniferous species in general. An appreciable disproportion in the numerical relationship of its components is a general feature of the entomofauna of the coniferous species. It is expressed in a clear predominance of xylophages over the phyllophages. The xylophages are represented by a large number of forms which are adapted to habitation in the bark, phloem, and wood, which are usually combined on the basis of this character into a unitary entomological complex which has received the name "trunk" insects. The emergence of such a complex was evidently brought about in various insects by the necessity of affording their progeny an abundance of food and simultaneously protecting them from unfavorable ecological conditions of the milieu and from enemies; this was achieved by the concealed mode of life of the larvae in the trunks and branches of trees. A convergent similarity in the general body structure, in the morphological characteristics of the structure of the internal organs, and in anabolism gradually developed in the larvae of systematically remote xylophage insects (Mamaev, 1966). It must be assumed that the evolution of this complex of insects took place in parallel with the evolution of the plants themselves; the strict confinement of a number of species to habitation on specific woody species points to this. But, due to the narrow specialization which reached a high degree in a number of instances, this complex may be regarded overall as a progressively developing group. The shift of many trunk insects from monophagy to oligophagy and, to some extent, to polyphagy is undoubtedly a later phenomenon which developed in the insects in the last stages of their evolution. The wide distribution which the broad oligophages and polyphages in particular have attained on the territory of the forest zone of the Palaearctic in the modern epoch attests to this.

Insects which feed on needles (phyllophages) do not form so extensive a group as the xylophages and do not exhibit such specialization, although particular representatives of the order of lepidopterans (leafrollers, moths, etc.) also shift to a concealed mode of life within needles, and then within shoots, while aphids (Chermes) do so within galls.
 

The Entomofauna of the Larch Stands
 

The Kuril Dahurian larch (Larix kurilensis) is encountered on Iturup and Shikotan. On Iturup it grows in the central part of the island, for the most part in mixed open stands of ErmanÆs birch and broadleaved species with a cover of Kuril bamboo (Figure 45), and on necks with less favorable climatic conditions, in combination with ErmanÆs birch, Japanese stone pine, and bamboo. The single area of nearly pure larch forests occupies a limited area on the Listvennichnoye Plateau south of the Kuibyshevskiy Neck, but this forest tract is drying out quite intensively. On Shikotan the larch is encountered singly and does not form forests.

The complex of insects associated with the larch is not numerous and is markedly impoverished as compared with the continent and even with Sakhalin. On Iturup, it is represented by several species characteristic for the larch - an aphid (Cinara laricicola Mats.), bark beetles (Dryocoetes baicalicus Reitt., Pityogenes bidentatus Herbst., Ips subelongatus Motsch.), a sawfly (Pristiphora erichsonii Htg.), and according to the data of K. Takeuchi (Takeuchi, 1955), by a webworm (Cephalcia alpina Kl.), as well as by oligophages of coniferous species which have migrated to the larch from spruce (Rhagium inquisitor rugipenne Reitt., Leptura succedanea Lew., Polygraphus sachalinensis Egg.), fir (Leptura scotodes Bat.), and from Japanese stone pine (Dryocoetes orientalis Kurenz.). In the region of the drying larch tract, the majority of the trunk insects enumerated multiply en masse, accelerating the destruction of the weakened trees. Small local foci of mass reproduction in individual wind-thrown or weakened larch trees are formed by some bark beetles (Pityogenes bidentatus Herbst. and Ips subelongatus Motsch.) in mixed larch-broadleaved forests. On Shikotan we found only isolated species, usually associated with the larch, in particular the "slender-antennaed" long-horned beetle (Tetropium gracilicorne Reitt.), which migrates here to spruce.

The impoverishment of the "larch" complex of entomofauna is obvious. Not one butterfly species which is specific for this species, the "Dahurian larch casebearer moth" (Coleophora dahurica Flkv.), which multiplies en masse on Sakhalin, the larch bud moth (Zeiraphera diniana Gn.), the "larch cone-roller" (Semasia perangustana Snell.), and others, which are common in Eastern Siberia (Raigorodskaya, 1966), has been found on the Kurils. According to L. A. IvlievÆs report, he observed a similar phenomenon on the Kamchatka Peninsula as well, where butterflies characteristic for the larch were also not found. The sole larch monophage among the bark beetles, the "Morawitz bark beetle" (Scolytus morawitzi Sem.), is also absent on the Kurils and Kamchatka. Judging by the character of the entomofauna, the larch forests tracts of Iturup and Kamchatka which are formed by the Kuril Dahurian larch have long been isolated from the principal, and essentially continuous, range of this species on the continent.
 

The Japanese Stone Pine Entomofauna
 

The Japanese stone pine (Pinus pumila) is one of the most widely distributed woody plants of the Kuril archipelago. It grows on the entire Greater Kuril Range, with the exception of the islands of Matua and Alaid. It is absent on the Lesser Kuril Range. On the southern islands it forms a fairly prominent altitude zone on the slopes of mountains, above the areas on which the dark coniferous and ErmanÆs birch forests flourish. It is also encountered under the canopy of various forests (dark coniferous, dark coniferous-broadleaved, ErmanÆs birch stands, and larch stands), in undergrowth, and on seashores. It frequently grows in groups among the Kuril bamboo. Groves of crooked Japanese stone pine are the predominant formation of woody vegetation, on a par with alder stands, on the central and northern islands (Figure 46).

The groupings of insects which live on the Japanese stone pine vary in different parts of the archipelago. Only two species (an aphid Pineus cembrae Chol. and the "alpine tundra bark beetle" Pityogenes foveolatus Egg.) accompany it throughout the entire extent of the range. On the southern islands, all the way to Iturup, species which have migrated to it from dark coniferous species, mainly from spruce, predominate on the Japanese stone pine. These are long-horned beetles (Judolia sexmaculata L., Strangalia arcuata Panz., Asemum amurense Kr.) and bark beetles (Polygraphus poligraphus L., P. sachalinensis Egg., P. gracilis Niiji., Crypturgus pusillus Gyll., C. hispidulus Thoms., Cryphalus piceus Egg., C. kurenzovi Stark, Pityophthorus lichtensteini Ratz., Dryocoetes orientalis Kurenz.). Of the insects biologically associated with the Japanese stone pine and which are specific for this plant in particular, only one species can be cited on the Kurils, namely, Pityogenes foveolatus Egg. With regard to inhabitants of the needles, this grouping is represented even more scantily. It includes 4 species of aphids widely distributed in the Palaearctic, oligophages of the genus Pinus (Pineus cembrae Chol., Cinara brauni CB., C. cembrae Chol., C. pinihabitans Mordv.), and a webworm of the Pamphiliidae family, larval nests of which are occasionally encountered on Iturup. We did not happen to observe butterfly caterpillars feeding on Japanese stone pine needles. It cannot be excluded that some butterflies develop in the cones and will still be found, since pests of the Japanese stone pine cones and seeds have not been studied thus far in the conditions of the Kurils.

In the Central Kurils, and particularly on Simushir, not a single insect species has been found on the Japanese stone pine. This agrees entirely with the opinion of B. A. Tikhomirov (1946b) on the later (after the last marine transgression) penetration of Pinus pumila into the central part of the Kuril Range, where its entomofauna is extremely impoverished. In general, the entomofauna of the Japanese stone pine over its entire range, which mainly encompasses the northeast part of Asia (Figure 47), has been insufficiently studied. At the present time, there are only two papers devoted especially to insect pests of this bush on Kamchatka (Kurentsov, Ivliev, 1960) and in Magadanskaya Oblast (Ivliev, Kononov, 1966a). Judging by these studies, as well as by our investigations on Sakhalin and the Kuril Islands, it can be concluded that an independent complex of insects biologically associated with the Japanese stone pine has not yet formed. This is evidently explained by historical factors, namely the "ancient genetic connections of associations of Pinus pumila with the forests" (Tikhomirov, 1946b, page 534) and by the specific characteristics of the growth of the Japanese stone pine in combination with various coniferous species in the modern epoch.

Depending upon growth conditions, under the forest canopy, in open areas of mountain slopes at the upper edge of the forest vegetation or on alpine tundra slopes [goltsy], the Japanese stone pine produces various ecological forms, from fairly small, nearly straight-trunked trees to prostrate bushes pressed to the ground. Its entomofauna is also directly dependent on the same conditions; its appearance differs in different regions. If the Japanese stone pine grows among dark coniferous forests or in immediate proximity to them, oligophages migrate to it from spruce and fir, as occurs on the Kurils, Sakhalin, in Sikhot-AlinÆ, on Kamchatka, and at some other sites; if, on the other hand, it grows among larch stands, insects associated with the larch are, as a rule, encountered on it; this is manifested especially clearly in Magadanskaya Oblast, and to some extent on Kamchatka and Sakhalin. Therefore, one can judge on the basis of the character of the entomofauna the species in combination with which the Japanese stone pine is growing, or has previously grown in one locality or another.

Only in the alpine belt, where other coniferous trees are absent and the climate is more severe, do monophages which are adapted to life exclusively on this shrub remain on it. There are a few, only 4, species of Japanese stone pine monophages. Of these, only the "alpine tundra bark beetle" is widely distributed, and evidently has a more ancient link with its food plant; it is known in Sikhot-AlinÆ, on Sakhalin, Iturup, Urup, Kamchatka, in Magadanskaya Oblast, and in North and Central Japan. The remaining monophages are encountered in various parts of the range of Pinus pumila, and the boundaries of their distribution are not contiguous; this points to the autochthonous origin of such species under conditions of territorial separation. These are the "Kamchatka" geometer (Cidaria kamtschatica D.), whose range is confined to the limits of the Kamchatka Peninsula proper south of ParapolÆskiy Dol (Kurentsov, 1963b), the "Japanese stone pine" long-horned beetle (Cornumutila semenovi Pl.), which is distributed north of ParapolÆskiy Dol in the basin of the Penzhina River, in Magadanskaya Oblast and Yakutiya, and the "Japanese stone pine" geometer (Bupalus cembraria Motsch.), which is known only in the alpine regions of Sikhot-AlinÆ. Since the Japanese stone pine emerged from beneath the coniferous forests and has formed independent plant associations, a phenomenon which is evidently secondary and historically relatively recent (Tikhomirov, 1946b), the 3 last-named species of insects, which have adapted to life only on this plant primarily in the subalpine belt, should probably be regarded as phylogenetically young. It is otherwise difficult to explain their absence on adjacent territories.
 

The Entomofauna of the Alder Stands
 

The alder is one of the most widely distributed woody plants of the Kuril archipelago. It grows on all the more or less large islands, but is represented by different species in different parts of the Range. Three alder species are encountered on the southern islands: the Japanese (Alnus japonica), the "hirsute" (A. hirsuta), and the "Maksimovich" (A. maximoviczii) alders. Of these, the first is known only on Kunashir, where it is infrequently encountered by chance; the second is present on Kunashir, Shikotan, and Iturup, where it occupies fairly small areas; its role as a forest-forming species is minor. The "Maksimovich" alder is predominant on the southern islands all the way to Urup. It is this species in particular which forms the majority of alder stands there. It grows at various altitudes, from the seashore to the mountain peaks, and depending on growth conditions produces various ecological forms, from small trees, 2-7 m in height (in the floodplains of rivers and among mixed forests), to small prostrate shrubs, 10-15 cm in height (on the stony peaks of mountain ridges). Elfin stands of the "Kamchatka" alder, A. kamtschatica, is the principal vegetation formation, on a par with Japanese stone pine groves, in the central and northern islands of the archipelago (Figure 48).

The entomofauna of the alder on the Kuril Islands is highly numerous and diverse (Figure 49). According to our data, it numbers more than 80 species belonging to various groups of insects; this substantially exceeds the volume of the entomofauna of any woody species on the archipelago. Among the insects associated with the alder we did not observe selectivity in relation to one or another of its species, even among the monophages. Although botanists (VasilÆyev, 1958, and others) note the great antiquity of the Japanese alder and the relative phylogenetic youth of the "hirsute" alder, D. P. VorobÆyev et al. (1966) point to the fact that both species frequently form hybrids. The phylogenetic age of the "Maksimovich" alder and the "Kamchatka" alder has not thus far been established. Apparently, the entire complex of the entomofauna of the alder stands is historically linked with the genus Alnus, the representatives of which are known on Sakhalin and Hokkaido as far back as the end of Mesozoic era (Krishtofovich, 1937b). They have thrived on islands, and evidently on the Kurils as well, throughout the entire Tertiary and Quaternary periods (Krishtofovich, 1936a; Neishtadt, 1955; Borsuk, 1956; Tolmachev, 1959; Tanai, Huzioka, 1967). Therefore, the association of a particular group of insects with the alder undoubtedly has ancient roots.

The alder stands within the limits of the Kuril Range are evidently a primordial and constant type of vegetation, which especially took root here in the Quaternary period, after the cooling off which had set in and the decline in the wide distribution here of the thermophilous broadleaved forests. The migration to alder of some insects from broadleaved species (oak, ash, and mulberry), referred to above, points to this. It can be seen from table 20 that polyphages which are trophically associated also with other plants constitute a substantial proportion (49 species) among the insects living on alder under the conditions of the Kuril Islands. For some of these an association with alder on the continent has not been established at all. It cannot be excluded that these species (Gastrolina peltoidea Gebl., Eugnamptus amurensis Fst., Depasophilus paeatus Fst., Deporaus mannerheimi Humm., Dictyoploca japonica Btl., Papilio machaon chishinana Mats., and others) adapted to habitation on alder secondarily, due to the altered ecological situation and the dropping out of initial food plants from the composition of the stands.

But, in addition to the polyphages, a group of monophages exists in this complex which are associated constantly with the genus Alnus and which exceptionally rarely migrate to other plants. This group includes all of the aphids observed on alder, a leafhopper Oncopsis sardescens An., long-horned beetles Plectrura metallica Bat., Eryssamena tuberculata Pic and Cagosima sanguinolenta Thoms., leaf beetles Basilepta balyi kurilensis L. Medv., Linaeidea aenae L., Luperus viridipennis laricis Motsch., and Stenoluperus cyaneus Baly, bark beetles Alniphagus alni Niiji., Cryphalus alni Krivol., Eocryphalus zachvatkini Krivol., and Dryocoetes ussuriensis Egg., a leafroller Epinotia rubricana Vl. Kuzn., and apparently some other species, whose biology remains insufficiently studied, in particular, a horntail Xiphydria jakovlevi Sem. et Guss. It is characteristic that the majority of monophages possess a narrow range which encompasses only the islands or a fairly small territory of the continent adjacent to them as well. There are species among the monophages which have vicariants in Eurasia and North America. For example, a bark beetle Dryocoetes ussuriensis Egg. has been supplanted in the western part of the Palaearctic by the close species D. alni Georg., which lives on the European alder (Alnus glutinosa). A bark beetle ["bast-eating"], Alniphagus alni Niiji., has two vicariants in North America, A. aspericollis Le Conte. and A. hirsutus Schedl, of which the former develops on the red and Sierra alders (Alnus rubra and A. rhombifolia), and is widely distributed in North America; the latter lives on the Sitka alder (A. sitchensis), and is distinguished by a narrower range, confined to the limits of British Columbia and the Copper Mountains (Swaine, 1918; Chamberlin, 1958). There are vicariants of the long-horned beetle, Plectrura metallica Bat., on the Korean Peninsula and in North America, and of the aphid Boernerina sp. in Europe, Japan, and North America. The presence of vicariants points to the ancient associations of these species with alder. All of the aphids observed on alder belong to ancient families (Shaposhnikov, 1951). These facts make it possible to regard the group of monophages as an indigenous complex of alder entomofauna, although the species included in it evidently are of varied phylogenetic age.

It is worthwhile to distinguish polyphages associated, in addition to the alder, with the birch, willow, and herbaceous plants, into a special group. This group most likely was formed in the Holocene, when microphyllous forests of various species of birch and alder predominated on the territory of Eastern Siberia and the Far East (south of the 60th parallel). The same vegetation occurred on the coasts of Kamchatka and evidently on many Kuril Islands, where it has persisted for a long period of time, whereas on the continent more rapid successions of vegetative zones took place in the middle and late Holocene (Neishtadt, 1955). It is significant that the species which are capable of developing on alder and birch (Linaeidea aenea L., Cimbex femorata L., Xiphydria camelus L.), as well as on alder and some herbaceous plants (Aphrophora similis Leth., Selatosomus affinis Pk., Dolerus yukonensis Nort.), are distinguished by a wide Transpalaearctic distribution. Insects associated with alder, willow, and tall herbaceous vegetation (Sinophora submacula Metc. et Hort., Cryptocephalus pumilo Sffr., Luperus flaviventris Motsch., Stenoluperus nipponensis Lab., Coenorrhinus interruptus Voss., Parapodisma mikado Bol., Ophthalmoserica karafutoensis Niiji. et Kinosh., Sericania sachalinensis Mats., Gonioctena japonica Chujo et Kimoto, Galeruca weisei Rtt.), by contrast, have a narrower range of the Manchurian and Island type.

The distribution across the archipelago of insects associated with alder conforms to a general regularity which is characteristic of the entomofauna of the Kuril Islands as a whole: the overwhelming majority of the species (73) is concentrated on the southern islands all the way to Urup. Of these, only 4 reach the north of the range, and only 7 species are known on the northern islands but are unknown on the southern islands.

According to our observations, only 11 species of insects live on the "Kamchatka" alder within the limits of the Kurils (see Table 20); this is explained by its territorial separation from the remaining species of the genus Alnus and by its growth in more severe climatic conditions. It is curious that of this group, only Dryocoetes ussuriensis Egg. is present on Kamchatka, while the remaining insects are represented by other species and are encountered rarely (Ivliev, Kononov, 1966a). On the Kurils, on the other hand, all species associated with Alnus kamtschatica are very common, and some (a leafhopper Oncopsis sardescens An., a leafroller Epiblema solandriana sinuana Hb., and the pyralid moth, Pyralidae sp.) multiply in great numbers.

As can be seen from the above, the alder entomofauna of the Kuril Islands is heterogeneous by origin, breadth of trophic associations, and phylogenetic age. But it was formed mainly under the influence of local fauna; the obvious predominance in its composition of Island, Ussuri-Island, Manchurian, and Okhotsk species, making up more than 75% of its total number, attests to this.
 

The Entomofauna of the Birch Stands
 

In the main, two birch species, the ErmanÆs birch (Betula ermanii) and the Asian white birch (B. platyphylla = B. tauschii), of which the former, which forms the majority of independent stands and is common in the composition of other forest formations, predominates, grow on the Kuril Range (VorobÆyev, 1963; Voroshilov, 1966). The former is encountered at various altitudes, from the seashore to a height of 600-700 m. The cover in the ErmanÆs birch stands most often consists of Kuril bamboo, and less frequently, of herbaceous vegetation. The birches are distributed within the limits of the archipelago only on the southern islands (Shikotan, Kunashir, Iturup) and on Urup, where apparently, as in the case of the alder stands, they are an ancient component of the forest vegetation.

The entomofauna of the Kuril birch stands is highly diverse. In addition to species associated directly with birch, a large number of insects which develop on other woody, scrub, and herbaceous plants and the Kuril bamboo growing among the birch stands, are present in it. However, the complex of "birch" insects proper is not that large; it numbers, according to our data, about 50 species. True, this number cannot be considered exhaustive, since it must increase somewhat when special investigations are carried out. Nevertheless, as compared with the alder, the birch entomofauna is appreciably scantier. In addition to the monophages which comprise approximately half of the complex (more than 20 species), a number of oligophages and polyphages which are also associated with alder, willow, oak, and other plants are included in it. A characteristic group of cockchafers (Ectinohoplia rufipes Motsch., Anomala lucens Ball., A. rufocuprea Motsch.) which feed on leaves may also be assigned to their number. During flight, the beetles of these species are found constantly in the birch stands, and Ectinohoplia rufipes Motsch. sometimes forms large clusters and severely damages leaves, especially in the understorey. The entomofauna of ErmanÆs birch stands of the Kuril archipelago differs from that of Sakhalin, the continental regions of the Far East, and Kamchatka. Its distinctiveness resides in a certain impoverishment of the species composition due to the absence of a number of widely distributed Palaearctic species, and in the presence of a substantial number of island forms which constitute almost the quarter of the entire faunistic complex.

This fauna is heterogeneous in origin. It consists of an ancient nucleus, initially apparently associated with primitive birches of the section Costatae Rge., to which Betula ermanii belongs (VasilÆev, 1942), and of a group of species of later genesis which have a broad range and which are distinguished by greater ecological plasticity. Species which have not exceeded the limits of the range of ErmanÆs birch in their distribution comprise the more ancient portion of the complex. These are mainly representatives of the Palaearchearctic fauna which have an Island, Ussuri-Island, Manchurian, and to some extent, Eastern Siberian type of range (the bugs Physatocheila orientis Drake and Elasmucha amurensis Kerzh., the already mentioned species of cockchafers, the long-horned beetles Paraclytus excultus Bat., Cyrtoclytus caproides Bat., Chlorophorus japonicus Chevrol., Exocentrus testudineus Matsush., and Pogonocherus dimidiatus Bless., the leaf beetle Phratora inhonesta Weise., the bark beetles Scolytus dahuricus Chap. and Trypodendron niponicum Bladf., the cutworm moths Acronycta incretata Btl. and Amphipyra schrenkii Mn., the geometer Cabera exanthemata Scap., and some others). Their trophic associations are typically limited to the birch alone, or include also the alder and some far eastern broadleaved species (oak, mulberry, etc.). Particular Island species have vicariants on the continent beyond the borders of the Far East, in particular the long-horned beetle Paraclytus excultus Bat., in Tibet and in the Caucasus; this provides grounds for regarding it as a Tertiary relict.

Insects which are characteristic of the birch stands of the Boreal zone (the broad Palaearctic and Holarctic species) are apparently phylogenetically younger. They are characterized, in addition to a wide distribution, by a different type of trophic associations. Various birch species which flourish in the northern part of Eurasia, willows, and herbaceous plants are included among their food plants. The following belong to this group: all of the aphids which we have found on birch (Glyphina betulae Kalt., Symydobius oblongus Heyd., Betacallis comes Walk., Euceraphis punctipennis Zett., Calaphis betulicola Kalt., Aphis galiae Iv.), a bug (Elasmostethus interstinctus L.), long-horned beetles (Strangalia aethiops Poda, S. quadrifasciata L., Xylotrechus ibex Gebl.), a leaf beetle (Cryptocephalus parvulus Mll.), a number of cutworm moths (Acronycta leporina L., Eurois exusta Btl., Mamestra thalassina contrastata Bryk, Enargia paleacea Esp.), geometers (Geometra papilionaria subrigua Prout., Boarmia bistortata Goeze), sawflies (Arge ustulata fuscipes Fall., Cimbex femorata L.), and a horntail (Xiphydria camelus L.). This portion of the fauna evidently formed in more severe, cryoxerophytic conditions, mainly as long ago as the Quaternary. On Urup, where the climatic conditions are less favorable than on the southern islands, species exclusively of this group form the birch entomofauna. On Iturup, they make up only half, while as one moves further southward, they occupy an obviously subordinate position as compared with the southern grouping of the ErmanÆs birch entomofauna.

Thus far the leafhoppers of the genus Oncopsis (O. adusta An., O. caliginosa An., O. sulphurea An., O. sepulcralis An.), described recently by G. A. Anufriev (1967) on the Kuril Islands, which live there on the ErmanÆs birch and are unknown at other sites, occupy an uncertain position. They are possibly the result of a later process of species formation and must be regarded as the youngest elements of the fauna.

The territorial separation of the Kuril birch stands from the Kamchatka birch stands also affects the character of their entomofauna, which contains relatively few common ("birch" proper) species, only several Palaearctic long-horned beetles and sawflies (Strangalia aethiops Poda, S. quadrifasciata L., Xylotrechus ibex Gebl., Cimbex femorata L., and some others) which have penetrated the Kurils and Kamchatka, evidently by different routes. Rather it is the insects associated with tall herbaceous vegetation that accompanies the ErmanÆs birch stands on Kamchatka which are common for these regions (Kurentsov, 1963b). Under the conditions of the Kuril Islands, where in the majority of cases tall herbaceous vegetation has been crowded out of the birch stands by the Kuril bamboo, the grouping of insects which live in the tall herbaceous vegetation cannot be regarded as an integral part of the entomofauna of the birch stands. The presence of a large number of endemics which are characteristic for the birch on Kamchatka, and their absence on the islands, suggest a rather prolonged separation of these biocenotic complexes overall. The varied zoogeographic associations also point to this: the substantial commonality of the Kamchatka entomofauna with the Nearctic, and the Kuril with the Japanese-Manchurian.
 

The Entomofauna of the Floodplain Forests
 

The floodplain forests on the Kurils form fairly narrow strips along the shores of rivers, streams, and lakes. They consist primarily of willows (Salix sachalinensis, S. urbaniana, and others on the southern islands, and S. chamissonis, S. cuneata, S. polaris on the northern), and the Manchurian and "Kamchatka" alders; on Iturup, they consist in places of the Japanese poplar (Populus maximoviczii), the Japanese bird cherry, the Korean mountain ash, and some other trees and shrubs. The grassy cover in the floodplain forests is usually thick, and consists of the many species of large-stemmed grasses (Filipendula kamtschatica, Senecio palmatus, Petasites amplus, Calamagrostis langsdorffii, Polygonum thunbergii, etc.) which form the groves of tall herbaceous vegetation that are characteristic for the islands (Figure 50).

In terms of species diversity and size of the populations of individual species, the entomofauna of the floodplain stands yields only to the entomofauna of the broadleaved forests. Such abundance is a result of the concentration on comparatively small areas of species associated with willow, alder, and in places poplar, and number of other woody plants and tall herbaceous vegetation. Broadleaved forests frequently abut floodplain forests on the Southern Kurils; these broadleaved forests occupy more elevated terraces of river valleys and lakes (for example, in the valley of the Alekhin Stream, around Goryacheye and Peschanoye Lakes and another sites); or they are abutted by dark coniferous forests growing higher than the zone of the yearly inundation with flood waters. A fairly close contact takes place under such conditions between the faunas of the floodplain, broadleaved, and dark coniferous forests, as a result of which an impression is created of a great diversity of the insects in the floodplain stands, into which many species of butterflies, beetles, flies, and hymenopterans fly in from neighboring biotopes to feed on the flowers of herbaceous plants, especially the large members of the carrot family. In addition, insects whose larvae develop in the water or on aquatic plants - dragonflies, caddis flies, Mayflies, stoneflies, some beetles, flies, and bugs, typically keep to the flood plains.

We have already touched on the entomofauna of the floodplain stands above to some extent when we considered the ecological features of the Golovnin Volcano caldera, and when we analyzed the entomofauna of the alder stands. Here let us dwell briefly only on the characterization of the complex of insects associated with the willow, one of the principal forest-forming species in the narrow and relatively short flood plains of the Kuril rivers and streams. The entomofauna of the poplar and aspen has been studied insufficiently because of the scarcity of these species on the Kurils, and it will not be examined separately, although there exist many oligophages among the insects associated with the Salicacea family (Salicacea) that are capable of developing on various species of willows and poplars with equal success. In general, the number of insects vitally associated with the Salicacea within the limits of the extensive range of these species is very great, but does have its specificity in various regions. Suffice is to say that more than 700 species of insects live on the poplars alone in the moderate zone of the Palaearctic (Grechkin, Vorontsov, 1962). This is explained, on the one hand, by the great antiquity of the genera Salix and Populus, which are known as far back as the Cretaceous (Krishtofovich, 1933), and, on the other hand, evidently, by the features of the biochemical makeup of these plants (Blagoveshchenskiy, 1966) and by their flourishing in more moist ecological conditions.

By comparison with Sakhalin and various regions of the continent, the entomofauna of the willow and the poplar on the archipelago is appreciably impoverished. On the Kurils this complex comprises only about 60 species, and the phyllophages markedly predominate in it, while the xylophages are represented by an extremely limited number of polyphagous species which migrate to the willow from other plants. Of the long-horned beetles, Plectrura metallica Bat. and Rhopaloscelis unifasciatus Bless. have been observed on the willow; on Iturup, Lamia textor L. (according to Japanese data); of the weevils, only Cryptorrhynchidius lapathi L. has been observed. Of these, the first lives mainly on alder, and settles on willow relatively rarely; the second, being a polyphage, also migrates to the willow from other plants; the third is known from a single find on Iturup, and only the last is actually associated with the willow and forms small foci on the southern islands. The long-horned beetles of the genera Saperda and Obrium, borers, bark beetles, and goat moths which are characteristic of the willow and the poplar are entirely absent on the Kuril Range; damage caused by clear-winged moths are not encountered.

Leaf beetles (Syneta adamsi Baly, Smaragdina aurita nigrocyanea Motsch., Cryptocephalus approximatus Baly, C. parvulus Mll., C. pumilo Sffr., Chrysolina surichalcea Mann., Plagiodera versicolora Laich., Chrysomela tremulae F., Gonioctena chujoi L. Medv., Pyrrhalta lineola Fabr., Clitena fuscipennis Jac., Luperus flaviventris Motsch., L. viridipennis laricis Motsch., Stenoluperus nipponensis Lab., Crepidodera japonica Baly) are the most numerous group among the phyllophages. The overwhelming majority of them are common, but not populous species, with the exception of Plagiodera versicolora Laich., which is constantly present in the floodplain stands in the south of the Range (all the way to Urup), and which is distinguished by large numbers. The butterfly fauna, which is still insufficiently studied, is quite diverse in the willow stands. Of these, only the leaf-roller Epinotia salicicolama Vl. Kuzn. multiplies in appreciable numbers (Kuznetsov, 1968a); the others are encountered sporadically and in small numbers. These include the tussock moths Subacronicta megacephala Schiff. and Acronycta [sic - spelling??]leporina L., the cutworm moths Eurois exusta Btl., Scoliopteryx libatrix L., and Colobochyla salicalis Schiff., a geometer Abraxas orientalis Sby., and some others. Ussuri puss moth (Dicranura vinula felina Btl.) caterpillars are encountered here and there on aspen.

Diverse sawflies are constant inhabitants of the floodplain stands, but a fairly small group (Arge enodis L., A. ustulata fuscipes Fall., Cimbex lutea L., Rhogogaster viridis L., several gall-forming species) lives on willow; the rest of the sawflies for the most part are associated with herbaceous or scrub vegetation. Only Rhogogaster viridis L. is distinguished by high population size and wide distribution throughout the entire archipelago. Aphids (Tuberolachnus salignus Gmel., Pterocomma salicis L., Aphis furcula Zett., A. galiae Iv.) and leafhoppers (Sinophora submacula Metc. et Hort., Aphrophora alpina Mel., A. alni Fall., Macropsis sp., Idiocerus ikumai Mats., Linnavuoriana sexpunctata Fall.), of the other components of the willow entomofauna, may also be noted. In terms of the character of their alimentary associations, oligophages and polyphages which are capable of developing, besides the willow, on alder, birch, and herbaceous plants, predominate in the complex under consideration. Monophages, on the other hand, occupy a subordinate position: they account for no more than one third of the species.

Despite the wide distribution of willow over the entire Kuril Range, the majority of the insects confined to it are concentrated on the southern islands; this is in agreement with the distribution over the archipelago of the alder entomofauna and other entomological complexes.

In the opinion of A. I. Kurentsov (1963a, 1963b), V. N. VasilÆyev (1958), and other investigators, the biocenoses of the floodplain forests of poplar and willow in Eastern Siberia and on Kamchatka comprise the legacy of ancient Tertiary forests that have been markedly impoverished and altered in the course of the Quaternary. This is applicable also to some degree to the Kuril floodplain stands, in which a number of relict species of insects have been preserved on alder, tall herbaceous vegetation, and some other plants. With regard to willow in particular, on the other hand, its entomofauna overall is rather of Quaternary age, with the exception of a small group of oligophages and polyphages which are also associated with alder and with individual broadleaved species. The following can be assigned to this group: the long-horned beetles Plectrura metallica Bat. and Rhopaloscelis unifasciatus Bless., some leaf beetles, butterflies, and other insects which exhibit limited ranges of the Island, Ussuri-Island, and Manchurian type and stenomorphism. At the same time, young evolving species also have a place here, for example the leaf beetle Gonioctena chujoi L. Medv., which has formed several subspecies on the relatively small territory of its range, evidently the leafhopper Idiocerus ikumai Mats., which has a narrow Kuril-Japanese range, and some others.

It is known that many endemic species of willows, which probably arose as a result of an active process of recent species formation that is taking place under island conditions, exist on Sakhalin and the Kurils (Sokolov, Svyazeva, 1965, etc.). It cannot be excluded that this process is also reflected in insects which live on rapidly evolving plants, and in its turn induces adaptive transformations in them.
 

The Entomofauna of the Herbaceous Vegetation
 

The herbaceous vegetation on the Kuril Islands is encountered in various forest formations, under the forest canopy and in forest clearings, on seashores and marine terraces, around rivers and lakes, and in waterlogged areas. The species composition of the grasses is highly diverse. Growths of tall herbaceous vegetation and forb meadows are the most characteristic for the Kurils. Tall herbaceous vegetation proliferates luxuriantly in the moist flood plains of rivers and streams, here and there along seashores, among cliffs, and in humid sites on mountain glades (Figure 51). It consists of robust tall grasses that reach a height of 2-3 m and form a number of associations with predominance of various plants: Sakhalin knotweed (Polygonum sachalinense) or Weyrich knotweed (P. weyrichii), meadowsweet (Filipendula kamtschatica), "hastate" Cacalia (Cacalia hastata), groundsel (Senecio palmatus), common cow parsnip (Heracleum lanatum), "Japanese" coltsfoot (Petasites japonicus), etc. The forb meadows are located mainly on marine terraces and upper sections of mountain slopes, where the snow is blown off in the winter and therefore the Kuril bamboo does not grow. Meadow areas alternate not infrequently with groves of shrubs and groups of dwarf trees.

The entomofauna of the herbaceous vegetation (hortophils) has not been studied in detail; this extensive topic constitutes the subject of special research. However, investigations have shown that an extensive complex of insects, encompassing the majority of the large orders, develops on various groupings of herbaceous plants. A certain differentiation of the entomofauna with respect to groups of plant associations is observed (species associated with tall herbaceous vegetation, various meadows, bogs, and coastal vegetation). In addition, many insects which fly in from neighboring biotopes feed on flowering grasses during the flight period. Thus, various hymenopterans (sawflies, ichneumon flies, bumblebees, wasps, and solitary bees), flies (mainly flourflies), diverse butterflies, beetles (cockchafers, scarabaeid beetles, long-horned beetles, weevils, blossom beetles, soldier beetles, click- beetles, etc.), and some groups of bugs typically concentrate en masse on the flowers of the "bear" angelica, other large members of the carrot family, and meadowsweet. Lists of the most common species frequenting flowers have already been cited in the description of the entomofauna of the coniferous-broadleaved forests, and to some extent, in the consideration of the individual groups of insects; therefore, we will not repeat them. Such a clustering of insects on the flowers of herbaceous plants is possible because of the fact that various plant communities on the Kuril Islands are in close proximity and frequently alternate with one another; the permanently elevated humidity moderates the ecological barriers between eco-areas; therefore, forest species regularly fly to meadows, while meadow species regularly fly to thinned-out stands or to forest clearings.

The composition of the insects living on tall herbaceous vegetation and meadow vegetation differs in different parts of the archipelago; this is also characteristic of other entomological complexes and is determined, apart from climatic conditions, by the boundaries of the ranges of the particular species. For example, the grouping of insects associated with tall herbaceous vegetation is much richer on Kunashir than on Iturup and on the islands located further to the north. Almost all species of long-horned grasshoppers and locusts which are known on the archipelago are included in it; of these, a wingless locust (Parapodisma mikado Bol.) vigorously eats round the leaves of many large herbaceous plants; the Japanese beetle (Popillia japonica Newm.), the "28-spotted" ladybird (Epilachna vigintioctomaculata Motsch.), a leafroller (Euops polita Roel.), and others feed on the leafs of Sakhalin knotweed, while the "urticarial" coccid scale (Orthezia urticae L.) feeds on its stems; evidently, the beetles of the tropical family Languriidae and another series of species of insects encountered only on Kunashir are associated with tall herbaceous vegetation. Several species of weevils (Hylobius gebleri Boh., H. perforatus Roel., Larinus formosus Petri., Trichalophusalbonotatus Motsch., etc.), striking because of their fairly massive body, are regularly encountered on the large leaves of various large grasses under the canopy of the broadleaved and floodplain forests, primarily in shaded areas.

The complex of hortophils which inhabit meadows differs appreciably from that living on tall herbaceous vegetation, although they do not yield to the latter in numbers and in species diversity. It consists of small cicadas belonging mainly to the fam. Cicadellidae, aphids of the fam. Aphididae, bugs, primarily of the fam. Miridae, leaf beetles, of which Atrachya menethriesi Fald. is distinguished by higher numbers, small weevils, click-beetles, small flies (from various families), numerous butterflies (especially moths, leafrollers, and cutworm moths), sawflies, ichneumon flies, and other insects, the detailed enumeration of which would occupy too much space.

Various background species are characteristic in the majority of cases for the meadow eco-areas of various islands. On Kunashir these are blues (Lycaena euphemus doii Mats. and Chrysophanus phlaeas daimio Seitz), cutworm moths (Hermonassa arenosa Btl., Axylia putris L., Crino adusta Esp., Trachea auriplena Wlk., Plusia nadeja Oberth.), and bumblebees (Bombus sp.); on Kunashir and Iturup, a cutworm moth (Parastichtis rurea F.), bumblebees (Bombus japonicus Fr. et Wag., which is characteristic also for the meadows of the central islands, and B. hypocrita Perez), a number of species of flourflies, sawflies, and other insects. On the north of the archipelago, the entomofauna of the meadow vegetation is distinguished mainly by a different set of species, including background species, including the "common fritillary" (Brenthis selene Sch.), the "strong-headed palaemon/leaf-rolling sawfly" [??] (Pamphila palaemon murasei Mats.), a green-veined white butterfly (Pieris napi bryoniae O.), a tiger moth (Parasemia plantaginis kunashirica Bryk), a bug (Irbisia sericans Stl), a leaf beetle (Chrysolina staphylea L.), a sawfly (Byrrhus fasciatus L.), and bumblebees (Bombus albocinctus Smith. and B. tichenkoi Skor.).

By comparison with the meadows, the entomofauna of waterlogged areas is somewhat impoverished, but in general it is nevertheless diverse. In bog vegetation we collected several species of cicadas (Elymodelphax excisa Mel., Aphrophora similis Leth., Neophilaenus albipennis F., Notus flavipennis Zett., Macrosteles striifrons An., Limotettix kuwayamai Ish., Sorhoanus tritici Mats.), psyllids, bugs of the fam. Saldidae, flies of the fam. Dolichopodidae, a number of species of cutworm moths, and several other insects.

A cursory review of the entomofauna of the herbaceous plant formations, without detailed analysis of the trophic associations of the insects with specific taxonomic groups of plants, does not make it possible to draw specific conclusions regarding its phylogenetic age. So far only some general considerations, which come down to the following, can be advanced in this connection on the basis of the character of the ranges of particular species. The Kuril hortophils are heterogeneous in origin. They have ranges which differ in latitude, and which gravitate to different zoogeographical regions. Among the species vitally associated with tall herbaceous vegetation flourishing on the southern and central islands, the overwhelming majority have ranges limited to the boundaries of the Palaearctic. These are mainly Manchurian and Island species, apparently comprising the most ancient, autochthonous nucleus of the fauna of the Kuril hortophils. The small number of subspecies forms also points to a certain conservatism of the group.

Part of the entomofauna that is limited to meadow vegetation must evidently be regarded as younger, formed on the islands after the appearance there of independent herbaceous associations. It is not accidental that widely distributed Holarctic and Transpalaearctic species predominate among the insects inhabiting meadow and bog vegetation. They are widely distributed over the archipelago; many are represented by Island or Eastern Siberian subspecies. A substantial proportion of these species penetrated the islands by different routes, probably as far back as the Quaternary. Some of these display a tendency under the conditions of insular isolation to develop local forms. A substantial number of North Palaearctic, Arctic, and Beringian species are present in the north of the Range in the structure of the meadow entomofauna.
 

The Entomofauna of the Kuril Bamboo
 

Groves of Kuril bamboo that occupies extensive areas and acts as one of the basic landscape plants (Figure 52) form a distinctive vegetative formation in the southern half of the archipelago. An original grouping of insects, consisting entirely of southern species adapted to habitation in the conditions of moderately cool climate of the Kuril Islands, is associated with this "specific type of vegetation, characteristic mainly of the typical tropics and subtropics" (Yaroshenko, 1960b, p. 157) (Figure 53).

Butterflies belonging to the Satyridae (Lethe diana Btl., L. callipteris Btl., Neope gaschkewischi Mn.) and Hesperiidae (Parnara pellucidae sachalinensis Mats., Halpe varia Murr.) families comprise half the group. All of them, with the exception of Lethe callipteris Btl., are highly numerous on Kunashir, where, just as the bamboo itself, they are background. Only the diana [??] satyr butterfly is encountered en masse on Shikotan. Only the diana [??] satyr butterfly and Gashkevich satyr penetrate Iturup and Urup; but there they are sparse and even rare.

Included in the other half of this complex are the bamboo aphid (Takecallis bambusae Mats.), shield bugs (Nikkoaspis shiranensis Kuw., etc.), bugs (Paraplesius unicolor Scott of the Coreidae fam., and, according to the observations of . P. Narchuk, Erimiris tenuicornis Miy. et Has., and Dolichomiris kuwayamai Miy. of the fam. Miridae), as well as an entomophage which accompanies them, the beetle Trypherus niponicus Lew. of the Cantharididae fam., which has protective coloration against the color of the bamboo stems. But this predator is not found to the north of Kunashir; this is possibly one of the reasons for the mass multiplication of the bamboo aphid on Iturup and Urup.

The cockchafer, Ectinohoplia rufipes Motsch., feeds facultatively on Kuril bamboo leaves on Kunashir, while the "leaf long-horned grasshopper" Kuwayamaea sapporensis Mats. et Shir. does so on Shikotan.

The entomofauna of the Kuril bamboo comprises a markedly impoverished variant of the richer fauna of bamboo stands of the subtropics and tropics. The majority of the species included in it are trophically associated only with bamboo. The region of their distribution encompasses Japan, the Korean Peninsula, and China, and more rarely only the islands. Some species live on other graminaceous plants on the continent outside the limits of the range of the bamboo. For example, the skipper Parnara pellucida Murr. damages rice in Primorskiy Kray. It may have adapted to life on the bamboo secondarily.
 

The Entomofauna of the Near-Fumarole Areas
 

As we know, the Kuril Islands are in a region of active volcanism which exerts a substantial influence on nature on the archipelago. Even in periods between eruptions of volcanoes, the volcanic activity is felt here quite strongly due to the abundance of hot springs, fumaroles, and solfataras situated at the foot and on the slopes of active volcanoes, and sometimes at the seashore itself, in the intertidal zone (Figure 54). Special microclimatic conditions are created at such sites, and thermophilous species of animals which form distinctive microbiocenoses are frequently concentrated there.

We have observed original groupings of insects and other animals at the sites of the outflow of hot springs on the shore of the Sea of Okhotsk to the southeast of Alekhino set., along the shores of Goryacheye and Kipyashcheye Lakes, in the environs of Reidovoye set. on Iturup, and in the fumarole fields of Mendeleev Volcano. The faunistic microcomplexes populating near-fumarole areas around Goryacheye and Kipyashcheye Lakes were described in considerable detail when the microbiocenoses of the caldera of Golovnin Volcano were considered. Here we will dwell on the characterization of other faunistic groupings.

Hot sulfurous springs vent in the environs of Alekhino set. in small bights along the Okhotsk coast. They are found at some distance from the shore or nearly at the water's edge among large stones. The soil, stones, and even sea water along the shore are substantially heated by the hot water of the springs. The small bights themselves are rimmed by fairly steep cliffs which protect a relatively narrow shore strip from the direct action of the easterly winds in the summer period. These areas are settled by original animals which are not encountered in like combination in other sites. A brightly colored lizard, the Far-Eastern skink (Eumeces latiscutatus Hallow.), which belongs to a tropical genus distributed in South East Asia, Africa, and North and Central America; a wingless tropical long-horned grasshopper Diestrammena japonica Karny (Figure 55, 4); a large wingless earwig (Anisolabis maritima Gn), an inhabitant of the shores of seas and oceans, primarily in more southerly latitudes; and the "Japanese great dragonfly" Polycanthagyna melanictera Selys. live there. Snow does not stick at such sites and is quickly melted by the heated ground and the warm evaporations; this allows the tropical long-horned grasshopper to remain in an active state year-round.

There are also similar vents of hot springs on the ocean shore of Kunashir, in the environs of Goryachiy Plyazh set. But here the insect complex under consideration is absent; it is evidently represented by a different grouping which is still insufficiently investigated. In particular, only the locust Podisma kurilensis B.-Bien., which is endemic for Kunashir, is known here. It cannot be excluded that the daily powerful ocean tides prevent the formation of a stable entomocenosis on this area; during those tides the area occupied by the springs is entirely covered by sea water.

We have observed an interesting grouping of insects around a hot sulfurous spring several kilometers away from the sea in the region of Reidovoye set. on Iturup. There the hot water does not flow to the surface, since the spring itself is enclosed in a well, is covered and is used for medicinal purposes. But soil surrounding it is waterlogged and permeated by hot sulfurous evaporations. Mass breeding of the small fly Caenia fumosa Sth. of the Ephydridae fam. took place during our visit on August 3, 1963 (Figure 55, 3); individual representatives of this family live in hot sulfurous springs (Narchuk, 1970). The flies surrounded the well, soil, grass, and in general everything around the spring en masse (tens of thousands). According to the testimony of local inhabitants, such a massive appearance of these flies is observed periodically. Their larvae evidently develop in the warm waterlogged soil, permeated by sulfurous water and fumes. Very small bogs with sulfurous but already cold water, located at a short distance from the spring, are populated by different faunistic communities. Water skaters/striders (Limnoporus rufoscutellatus Latr.), small beetles, water scavenger beetle (Enochrus sp.), and the larvae of small dragonflies (Sympetrum maculatum Oguma) live in them, while a small predatory bug (Salda littoralis L.) lives along the shores.

N. N. Konakov (1956) described specific complexes of insects in the fumarole fields on the slopes of Mendeleev Volcano. He distinguished typhobiontic insects constantly living in immediate proximity to the fumaroles, solfataras, or in the bare "waste grounds" adjacent to them and participating in the formation of very simple biocenoses, as a special ecological group. N. N. Konakov assigned the following to these complexes: small flies from the genus Discocerina Mcq. (Ephydridae), which cluster en masse at the very openings of fumaroles and on their walls, which have a temperature of +47 to +61; tiger beetles Cicindela elisae Motsch. and C. sachalinensis A. Mor., which are encountered on the territory of fumarole fields, but which do not fly closer than 1-2 m to the fumaroles themselves; single sweat bees (Halictus rufitarsis Zett., H. albipes Fabr., Halictus sp.), which dig burrows right there in the pumice-like rock; and inhabitants of the Kislaya River (flowing from the described fumarole field, and having a water temperature from +40 to +43C and a sharply acidic taste) - a skater/strider (Aquarius paludum F.) and larvae of dragonflies close to the genus Orthetron. The small flies were absent during our visit to these fumarole fields at the end of July of 1962; this was apparently associated with the periodicity of their appearance.

N. N. Konakov called the temporary frequenters of the near-fumarole areas, which fly there from other biotopes, typhoxenes. They are common also on other fumarole fields, for example in the caldera of Golovnin Volcano, in the crater of Trezubets Volcano on Urup, and at other sites. These accidental immigrants, attracted by the heat or borne by the wind to an ecological niche which is not characteristic of them, cannot be regarded as constituents of the near-fumarole entomocenoses; this is confirmed by the mass destruction of such insects which fly close to the fumaroles and which do not tolerate a prolonged stay in the atmosphere of poisonous sulfurous fumes. N. N. Konakov describes a graveyard of various insects near a fumarole in the crater of the Trezubets Volcano in detail.

The faunistic complexes in question attest to the unusual diversity of the entomofauna of the near-fumarole areas, which is far from exhausted by the examples cited. In each individual case they are represented by original and unique variants of faunistic groupings, which remain insufficiently investigated. The fauna of the near-fumarole areas is undoubtedly richer than appears at first glance. It evidently consists of many stenobiontic species, for the identification of which special, more prolonged, and more detailed investigations are required. Some of these are primordial and already specialized inhabitants of these specific landscapes. Among these are flies which are adapted to residence in hot sulfurous fumes and water, and the larvae of some dragonflies and other insects. Other, mainly thermophilous, species (the wingless long-horned grasshopper, tiger beetles, skaters/striders, etc.) find favorable temperature conditions near the springs and are gradually adapted to life in an atmosphere of poisonous gases, but can also live among other landscapes given the presence there of the ecological optimum that they require.

Evidently, local microclimatic conditions existing around thermal springs and among fumaroles have served as refuges for a thermophilous fauna in the past, and permitted it to survive a general cooling off of the climate and to be preserved at such sites in the form of fragments of relict biocenoses. The presence on the Kuril Islands of special ecological complexes, confined to near-fumarole areas, heated as the result of volcanic activity, must be regarded as a distinctive feature of the entomofauna of the archipelago.
 

The Entomofauna of the Seashores
 

The seashores are one of the most typical island biotopes, thickly populated by various animals, among which the insects also find a place for themselves (Figure 56). The entomofauna of the seashores, especially of the supralittoral, is highly distinctive and consists to a substantial degree of specialized species, leading for the most part a concealed mode of life. The ecological conditions on the shores are more severe than on the marine terraces, and, all the more so, at sites remote from the shore by a substantial distance. Here the action of the constantly blowing winds is felt more strongly, while in the intertidal zone the breakers and the regular flood tides acquire great significance; this compels insects to seek reliable shelters or to develop special adaptations for life in this specific biotope.

The absence of phyllophages, with the exception of species living at a remote distance from the water, at the boundary of marine terraces where herbaceous vegetation appears, is a characteristic feature of shore entomofauna. Predatory insects or those feeding on various decomposing vegetable or animal remains cast up by the sea as well as dead crustaceans - beach fleas (Orchestia ochotensis) - which populate sandy shores at the water's edge en masse, are concentrated on sea shores. Several species of ground beetles, distinguished by various behaviors and occupying different ecological niches, are encountered here. Some of them lead nocturnal and others diurnal lives. A large East Asian ground beetle (Craspedonotus tibialis Schaum), a typical inhabitant of river and marine sandy shores (see Figure 2), begins to hunt at twilight. It constructs long burrows in the sand that exit at the surface in the form of horseshoe-shaped funnel traps, into which various prey (insects and crustaceans) fall accidentally. The beetle sits in the burrow, at the very entrance, and seizes prey which has fallen into the trap. By day it is difficult to see this beetle; it is usually deep in the burrow or in other shelters, and appears rarely at the surface. It does not move rapidly across the sand and does not have great agility; but on the other hand, it does dig rapidly into the sand. A second large ground beetle - Carabus (Damaster) rugipennis Motsch. - may be encountered fairly frequently on the sea shores of Kunashir, less often on Iturup, in the daytime (see Figure 34). These beetles roam quite clumsily on their long legs along the ebb tide in search of molluscs and other small animals cast up by the sea. Other ground beetles (Carabus maeander Fisch.-W., Nebria ochotica Sahlb., Pterostichus oblongopunctatus Fabr., Pterostichus sp.) hunt openly in the near-littoral zone as darkness sets in; this is suggested by their adroitness and capacity to run quickly. By day they hide under stones and various objects on the shore.

Algae, cast up on the shore and rotting, that form thick layers in places, comprise an ecological niche populated by numerous insects which are, however, relatively similar faunistically. Thus, flies of the specialized family Coelopidae, adapted to life in algae that have been cast up on the shore (see Figure 36), and of two other families (Rhagionidae and Dryomizidae), develop on decomposing sea kale (Laminaria). Larvae of the last two evidently live not only on algae but in the surface layers of sand as well, since their empty puparia collect in large numbers in small pits on the shore. Besides flies, several species of water scavenger beetles of the genus Cercyon (fam. Hydrophilidae), predatory beetles (Staphylinidae), and darkling beetles (Tenebrionidae) are encountered in the layers of algae. The predatory beetles and their larvae are common in general on the sandy shores, where they feed on small crustacea, the larvae of flies, and various animal remains cast up by the sea. They have the capacity to dig completely into the sand, are not discovered immediately, and probably tolerate a brief time in sea water, since they live in the tidal zone. Of the staphylinids, the most common are Cafius nudus Scharf., Cafius sp., Liusius hilleri Wse.; of the darkling beetles, the most common are Phaleromela subhumeralis Mars. and Emypsara riederi Fald. (Figure 57). The majority of the shore species have ranges limited to the boundaries of northeast Asia (the Okhotsk and the Ussuri-Island); only some are distributed more widely, throughout the entire northern Palaearctic or in Japan and China.