CHAPTER 3

SECTION 3. Order HOMOPTERA

Suborder CICADINEA

woody species or with herbaceous vegetation that is characteristic of forest eco-areas and that grows under the forest canopy, in forest clearings, and in the flood plains of streams and rivers (Table 3). The mesohygrophilic and hygromesophilic species living in bogs and waterlogged areas and feeding on grains and sedges form a fairly small group (10 species - 15.6%). Three species are xeromesophils. They are confined to meadow and near-fumarole, relatively dry, areas. 

An abundance of spittlebugs (Aphrophoridae), which comprise about 20% of all of their fauna and not uncommonly reproduce en masse (Aphrophora alpina Mel., A. intermedia Uhl., Philaenus spumarius L.), is a characteristic feature of the Kuril cicads. At other sites, for example in the European part of the USSR, spiThe cicads of the Kuril Islands have, until recently, remained nearly uninvestigated. Only a small amount of information on the detection on the archipelago of particular species has been contained in some Japanese papers (Kato, 1933; Ishihara, 1953; Esaki, Ito, 1954). The material we collected was analyzed by G. A. Anufriev and presented by him in the special articles (Anufriev, 1968a, 1970) that have been used in the writing of the present section. 

In a recently published study, Ishihara (Ishihara, 1953) cited 36 species; according the calculations of G. A. Anufriev, only 10 of these are lacking in our collections. They have been included in the general list with the appropriate reference. A substantial proportion of the species mentioned by Ishihara were evidently misidentified by him and require refinement. At the same time, some of the identifications made by G. A. Anufriev were also imprecise, and must be corrected in the light of Ishihara's article. All of the corrections and refinements of the species names of the cicads have been made and graciously furnished to us by G. A. Anufriev; they are included in Table 3. 

The extensive group of the cicads is represented on the Kuril Islands by a highly impoverished complex, by comparison with adjacent parts of the continent and Japan. Representatives of eight families have been recorded here: Derbidae (2), Cixiiidae (2), Delphacidae (6), Cicadidae (3), Cercopidae (1), Aphrophoridae (12), Membracidae (2), and Cicadellidae (39); 67 species in all, belonging to 50 genera. Of these, 10 species and 2 genera have proven to be new to science (Emel'yanov, 1966; Anufriev, 1967, 1968c, 1970), 4 species new for the fauna of the USSR, and 4 were identified only to the generic level. It may be expected that representatives of several other families (Fulgoridae, Dictyopharidae, Achilidae, Tettigometridae) that are encountered in Primorskiy Kray and Japan, but have thus far not been noted on the Kuril Islands, possibly due to the insufficient investigation of their entomofauna, will be found on the archipelago. 

The poverty of the cicad fauna on the Kuril Islands is expressed in the limited number of families present here and of the species included in them (8 families on the Kurils, 15 in Primorskiy Kray, and 20 in Japan). According oral report of G. A. Anufriev, about 400 cicad species have been noted in Primorye, that is, six times more than in the Kurils. In Japan, 259 species have been counted in the family Cicadellidae alone (Esaki, Ito, 1954); thus far only 38 have been recorded in the Kurils. 

The cicads inhabiting the Kuril Islands are mainly represented by sylvan, mesophilic species (47 species - 73.4%), associated with ttlebugs constitute only 2.3% of the total number of cicads (Emel'yanov, 1964). Their larvae, due to the constant high humidity on the islands, live in foam exclusively on leaves, and not on their axils. They damage the most diverse woody species, shrubs, vines, and herbaceous vegetation. The majority of the species are wide polyphages; less often they are oligophages. Thus, A. intermedia Uhl., whose larvae are bred in cages directly in nature, develops, in particular, on Sakhalin fir, white-leaf Japanese magnolia, panicled hydrangea, trichocarpous toxicodendron, actinidias and tall herbaceous vegetation; A. alpina Mel., on willow, oak, and tall herbaceous vegetation; Polminellus nigripectus Mats., on Sakhalin cherry and other Rosales; Peuceptyelus coriaceus Fall., on various pines. 

The cicads are unevenly distributed on the Kuril Range. The majority of the species are associated with broadleaved-coniferous forests and are prevalent on Kunashir, Shikotan, and Iturup; their fauna is richest (43 species) on Kunashir, and in particular on the southern half of the island, especially along the western littoral. Representatives of the southern, mainly subtropical and tropical, families (Derbidae, Cicadidae, Cercopidae) have been collected principally along the southwest coast of Kunashir, in the region of the caldera of the Golovnin Volcano and in the environs of the village of Alekhino. Only the large cicada, Tibicen bihamatus Motsch., is also fairly numerous on Shikotan. The southern species no longer penetrate Iturup. Only 19 species have been recorded on this island, associated both with woody and herbaceous vegetation, or limited to moist eco-areas, not infrequently to waterlogged areas. Here some species of spittlebugs (6) and leaf hoppers (9) are quite common; Delphacidae and Cixiiidae are encountered singly. The cicad fauna is even more impoverished on Urup; 6 species have been noted:² 1 from the fam. Aphrophoridae; 2 from the fam. Cixiiidae; 3 from the fam. Cicadellidae. On islands to the north of Urup, representatives of the fam. Cicadellidae are encountered exclusively. Of these, only one species, Oncopsis sardescens An. is associated with the "Maximovich" and "Kamchatka" alders; the rest live on herbaceous plants, mainly on grains. The cicad fauna is maximally impoverished on Simushir (here only 2 species have been found), but as Kamchatka is approached, it begins to become richer once again, but now with northern species. Five species are noted on Paramushir, of which 3 are unknown on more southerly islands.

² The number of species on the islands of Urup and Simushir will undoubtedly increase in further investigations, since our visits to these islands were brief.

From the zoogeographic point of view, the cicads of the Kuril Islands are highly heterogeneous. They can be divided into four groups on the basis of the character of the ranges: 1) widely distributed species; 2) species prevalent in North East Asia and North America; 3) Manchurian species with a range encompassing the Amur basin, including Primorskiy Kray, Northern China, and the territories adjacent to it; and 4) island species, lacking on the continent.

The first group (24 species) combines species prevalent throughout the world: cosmopolites (4.7%), holarctic (4.7%), and transpalaearctic (26.6%). They are distributed more or less evenly throughout the territory of the archipelago (see Table 3), but they predominate numerically on the southern islands.

Only 2 species are included in the second, smallest group; one is Beringian (Diplocolensis evansi Aschm.), prevalent in Alaska, Canada, on the Chukot Peninsula, and on the Northern Kuril Islands; the other is Okhotsk (Oncopsis sardescens An.), known in Magadanskaya Oblast (based on the collections of D. G. Kononov), and found throughout the entire Kuril Range.

The Manchurian group is somewhat smaller than the first (18 species - 26.6%), but in essence makes up, together with the Island species, the nucleus of the fauna on the southern islands. Representatives of this group do not penetrate north of Urup.

The fourth group (19 species - 28.2%) consists of Island, namely Kuril-Japanese, Sakhalin-Kuril-Japanese, Sakhalin-Kuril, and Kuril proper, species, also concentrated on the southern islands (Kunashir and Shikotan). Only isolated representatives of it reach Iturup and Urup, and they are absent on the central and northern islands.

The zoogeographical position of the four (6.2%) species that have been identified only to the generic level is thus far unclear.

For purposes of economy of space, the list of the cicad species observed on the Kuril Range is presented in Table 3, in which a list of synonyms based on the articles of G. A. Anufriev (1968a) and T. Ishihara (Ishihara, 1966) is also included.

Three species mentioned by Ishihara are not included in our list: Oncopsis alni Schrk., Macrosteles sexnotatus Fall., and M. quadrimaculatus Mats. In the opinion of G. A. Anufriev, the identification of M. quadrimaculatus Mats. and M. sexnotatus Fall. needs to be checked, and the finding of these species on the Kuril archipelago is in need of further confirmation; Oncopsis alni Schrk. evidently belongs to one of the species of the genus Oncopsis described by Anufriev.
 

Suborder APHIDINEA - APHIDS

Information on the aphid fauna of the Kuril Islands has until recently been entirely lacking in the Soviet literature. Only brief reports of the finding of individual species of aphids on the Kurils were published by Japanese investigators, starting in 1932, but unfortunately all of these publications were in the Japanese language and have been practically inaccessible for Soviet specialists. An article was published (Krivolutskaya, Ivanovskaya-Shubina, 1966) in 1966 on the basis of our collections and observations, in which 48 species of aphids, belonging to 7 families, were cited. Later S. Kuwayama (Kuwayama, 1967), in a synoptic article on the entomofauna of the Southern Kuril Islands, added another 18 species to our list, mainly associated with herbaceous plants, both cultivated and wild (of these, 4 were identified only to the generic level). But this list also cannot be regarded as exhaustive, since aphids have been gathered both by us and apparently by Japanese entomologists from the above-ground parts of the plants, while those living on the roots have to a substantial degree escaped the notice of investigators. Species associated with herbaceous plants are still insufficiently explored. Therefore, later, when special collections are made, the number of aphid species on the Kuril Islands, especially the southern islands, will undoubtedly increase.

Nevertheless, the material we have at our disposal makes it possible even now to make judgments regarding the composition of the aphid fauna populating the islands, its disposition on their territory, the trophic and genetic linkages, as well as the practical significance of some mass species in the archipelago's conditions. The aphids we have collected have been analyzed by O. I. Ivanovskaya. She has described two new species and one genus from Kunashir (Ivanovskaya-Shubina, 1966a, 1966b).

In all, 66 species of aphids, belonging to 7 families and 33 genera, are known on the Kuril Islands at the present time: Adelgidae (3), Pemphigidae (5), Lachnidae (8), Thelaxidae (8), Callaphididae (12), Chaitophoridae (4), and Aphididae (33). Of these, 56 have been identified to the species and 10 to the generic level.

The majority of the species (41) are associated with woody and scrub plants; the minority (20), with herbaceous plants; in five species trees are the main food plant and herbaceous plants supplemental. Aphids living on coniferous species are represented by nine species: 3 on fir, 3 on Japanese stone pine, 1 on fir and spruce, 1 on Japanese stone pine, and 1 on larch. A substantially larger number of them (38) are associated with leaved varieties: 4 species on willow, 6 on birch, 6 on alder, 3 on oak, 3 on elm, 4 on mountain ash, 4 on cherry, 2 on bird cherry, 4 on maple, 1 on udo, 2 on elder, 1 on honeysuckle, and 1 on panicled hydrangea. Of the aphids observed on herbaceous plants, the majority of the species live on Compositae (10) and grains (5); the "bamboo" aphid (Takecallis bambusae Mats.), which lives on the Kuril bamboo (Sasa kurilensis) and which is penetrating the Central Kuril Islands in its wake, is of substantial interest.

The aphid fauna is distributed on the territory of the islands in the following manner:
 
 

The figures given demonstrate the clear impoverishment of the aphid fauna in the central and northern parts of the Kuril Range. A definite specificity in the composition of the fauna is characteristic for individual islands. This is expressed in the fact that there are only 14 species that are prevalent on two or several islands. The rest are found on only one of the islands and are not encountered at other sites. There are 30 such species on Kunashir, 13 on Iturup, 6 on Shikotan, 2 on Urup, and 1 on Paramushir.

At the present time, due to the insufficient exploration of the aphid fauna in the eastern part of the country and the lack of regional compendia on this group, it is difficult to judge the degree of impoverishment of the Kuril aphid fauna as compared with nearby regions of the continent. But on neighboring islands, Sakhalin and Hokkaido, where the aphids have been studied in greater detail by Japanese investigators (Inouye, 1936, 1938, 1953, 1956; Hori, 1938), their species composition is substantially richer than on the Kuril Range, and the numbers of the individual species are greater. For example, the fam. Adelgidae is represented on Hokkaido by 11 species, on Sakhalin by five, and only three on the Kurils. Moreover, species of the genus Adelges have not been observed at all on the Kuril Range, while Adelges japonicus Monz. is encountered en masse on Sakhalin and Hokkaido, and is a serious pest of the Yeddo spruce (Novokhatka, Kovalenko, 1968).

In the conditions of the Kuril Islands, the most active reproduction of aphids is observed in the first half of the summer, approximately up to the middle of July. Winged individuals which fly to other food plants appear en masse at that time, and therefore are conspicuous more often.

The aphids of the Kuril Islands are represented mainly by moisture-loving forms. Only several species are relative xerophils to one degree or another; these are Aphis jacobae Schr., A. spiraephilae Patch., Myzus cerasi F., and species of the genus Dactynotus, the overwhelming majority of which are associated with herbaceous vegetation, and settle on plants of open, stony sites or on those springing up along forest margins and clearings.

The aphids do not inflict substantial damage on the islands, with the exception of particular species that reproduce en masse. Among these are the common "bird cherry" aphid (Rhopalosiphum padi L.),which severely damages a bird cherry (Padus ssiori) on Iturup, Periphyllus aceris L., which is often encountered on the painted maple (Acer pictum) in the leaved forests of Kunashir and Iturup, the "cherry" aphid (Myzus cerasi Fabr.), which is widely distributed on Iturup, and which causes curling of "Sakhalin" cherry (Cerasus kurilensis) leaves. The remaining species are encountered sporadically, sometimes forming colonies of varying size on individual trees or groups of trees. Thus, large colonies of Symydobius oblongus Heyd. can be found on birch shoots at some sites on Kunashir. Among the aphids living on herbaceous plants, the "bamboo" aphid (Takecallis bambusae Mats.) reproduces en masse. Bamboo stands were solidly infested with this aphid on the central part of Iturup in 1963. Large black colonies of Dactynotus facae L. can often be encountered on stems of the hair-vein agrimony (Agrimonia pilosa).

The aphids are a rather conservative group. They feed on strictly determined plants, and migrating species require a particular spectrum of plants for their development. If one of the host-plants for dioecius aphids disappears at a particular site, the aphids living on those plants cannot complete the full cycle of development, and become incomplete-cycle forms, multiplying parthenogenetically on one plant. At the same time, aphids can follow their host-plant and spread quite far from the initial habitat. Therefore, the ranges of many species of aphids extend beyond the boundaries of one faunistic region.

The Kuril aphid fauna is represented in the main by widely distributed boreal species (31 species - 46.9%), the majority of which (37.9%) are indigenous to the Palaearctic, and 9% to the Holarctic. A group of species which are also prevalent in other faunistic regions, the Indo-Malaysian, Ethiopian, Neotropical, and Australian, occupies a substantial place (13 species - 19.8%).

By contrast with other insect orders inhabiting the Kuril Range, the island species of aphids are relatively few in number - the Kuril-Japanese (Byrsocrypta ulmi jezoensis Mats., Cinarra longipennis Mats., C. laricicola Mats., Betacallis alnicollens Mats., Aulacorthium magnoliae Ess. et Kuw., Dactynotus giganteus Mats., Macrosiphoniella yomogicola Mats.) and Kuril proper (Sorbaphis kurilensis Iv., Heterogenaphis kunashyri Iv.). Manchurian elements make up quite a small group (Tuberculatus favus Mordv., Aphis araliae Mats., Megoura viciae japonica Mats.). It is true that the zoogeographical affiliation has not yet been established for 10 species (15.2%) thus far identified only to the generic level. It must be assumed that after it has been determined, the percentage ratios of the zoogeographical groupings among the aphids will shift in the direction of an increase in the Kuril-Japanese and Manchurian groupings, since almost all the unidentified species are found on Kunashir. The overwhelming majority of the Manchurian, Kuril-Japanese, and Kuril proper species are also concentrated on Kunashir. Southern elements essentially do not penetrate the remaining, especially the northern, islands of the archipelago. On those islands the aphid fauna consists mainly of boreal, i.e., broadly palaearctic and holarctic, species.

The ranges of many palaearctic species remain insufficiently explored; in particular, there is little information on the distribution of the aphids over the enormous territory of Siberia. Therefore, at the present time the ranges of some species (Periphyllus aceris L., P. acericola Walk., P. lyropictus Kessler., and others) may only provisionally be considered amphipalaearctic.

A more complete characterization of all the species of aphids observed on the Kuril Islands is given below. Information on the general distribution is presented on the basis of the data of G. Kh. Shaposhnikov (1955, 1964), A. A. Popova (1967), C. Börner (Börner, 1952), M. Inouye (1953), and S. Kuwayama (Kuwayama, 1967), as well as on the basis of the collections of O. I. Ivanovskaya in Siberia.

SYSTEMATIC LIST OF SPECIES

Family ADELGIDAE

1. Pineus cembrae Chol. (=Chermes sibiricus Chol.) - the "Siberian" chermes. Shu - July 29, 1964, isolated apterous individuals, on Japanese stone pine (Pinus pumila); P - env. of Severo-Kuril'sk, July 27-28, 1964, isolated apterous individuals and colonies on Japanese stone pine; Southern Kurils (without indication of island, Kuwayama, 1967). Palaearctic species, widespread in the European part of the USSR; known in Kazakhstan, Western and Eastern Siberia, the Baikal region, Primorskiy Kray, Sakhalin, the Kuril Islands, and outside the USSR, in Western Europe and Japan (Hokkaido). Usually associated with spruce, cedar, and pines. Lives on Japanese stone pine under Kuril Islands conditions. According to our observations, may induce yellowing of the needles and drying of isolated shoots.

2. Pineus pini L. - the "pine" chermes. K - env. of the village of Alekhino, July 4, 1962, winged form, dark coniferous-broadleaved forest, on Sakhalin fir (Abies sakalinensis). The species range goes beyond the limits of the Palaearctic. Prevalent on the entire territory of Europe, in Siberia, the Southern Kuril Islands, in North Africa, Western Asia, Japan (Hokkaido), New Zealand, and Australia. Usually develops on pines. Rare on the Kuril Islands; evidently associated with dark coniferous varieties.

3. Aphrastasia pectinatae Chol. - the "brown spruce-pine" chermes. I - env. of the village of Lesozavodsk, July 26, 1963, apterous virgins, larvae, in mixed broadleaved-dark coniferous forests, on Sakhalin fir (Abies sakalinensis). Characteristic for the forest zone of the Palaearctic, prevalent in the European part of the USSR, Western and Eastern Siberia, the Baikal region, Primorskiy Kray, Sakhalin, and Hokkaido. M. Inouye (Inouye, 1953) distinguishes a special subspecies of this chermes on Sakhalin and Hokkaido - A. pectinata var. ischiharai Inou. Spruces (on the islands, Picea Glehni and P. microsperma), on which the aphids form galls, is the main food plant of the species; the winged individuals migrate to the Sakhalin fir, and the following year again return to the spruce. This species is rarely encountered under the conditions of the Kuril Islands.

Family PEMPHIGIDAE

4. Byrsocrypta hirsuta Baker. K - env. of the village of Alekhino, July 4, 1962, larvae and nymphs in galls on Manchurian elm (Ulmus laciniata) leaves. Within the USSR, observed in Central Asia, Uzbekistan, Kirgiziya (Nevskiy, 1929), in the south of Primorskiy Kray and on K, apparently along the northern border of the range, which encompasses Japan (the islands of Hokkaido and Honshu), China (including Taiwan), the Philippines, India, South Africa. Lives on leaves of various elms in sac-like short-stemmed galls, and migrants shift to the roots of grains, where they can exist as an incomplete-cycle form. Encountered on K at sites where elm springs up, and severely damages leaves on isolated trees.

5. Byrsocrypta ulmi jezoensis Mats. (=B. gallarum Gmel., Tetraneura ulmi Deg.) - "elm-grain" aphid. K - close to the village of Alekhino, August 1, 1962, alate virgins and nymphs on Manchurian elm leaves in a broadleaved forest. A Kuril-Japanese subspecies, according to the data of Kuwayama (Kuwayama, 1967), has been known on the islands of Hokkaido and Honshu. The principal form is widespread in the Palaearctic - Europe, the Caucasus, Central Asia, South Primorye, and China. As in the case of the preceding species, lives in galls on elm leaves and migrates to the roots of grains (Shaposhnikov, 1955).

6. Byrsocrypta sp. K - env. of the village of Alekhino, June 27, 1962, apterous individuals and larvae on Manchurian elm.

7. Paraprociphilus baicalensis Chol. (=Pemphigus baicalensis Chol.). Southwest slope of Mendeleev Volcano, July 26, 1962, apterous individuals on "Maksimovich" alder (Alnus maximoviczii). A transsiberian species, prevalent in the taiga zone from the Urals to Kamchatka. The aphids are in long white tomentum, suck on the inferior surface of leaves; on the continent they are also associated with various alder species. It is known (Mordvilko, 1929) that larvae and virgins overwinter by penetrating the understory around the trunks of trees.

8. Prociphilus xylostei Deg. "honeysuckle-spruce" aphid. I - env. of the village of Lesozavodsk, July 19, 1963, nymphs on edible honeysuckle (Lonicera edulis). A widespread Holarctic species, known in the European part of the USSR, the Caucasus, Central Asia, Primorskiy Kray, Western Europe, and North America. Lives on young shoots and honeysuckle leaves rolled into a small tube. Colonists migrate to the fine roots of firs and spruces (Shaposhnikov, 1955).

Family LACHNIDAE

9. Cinara brauni C. B. K - env. of the village of Alekhino, July 15, 1962, alate, 1 spec., in flight. Species range insufficiently investigated, but may be amphipalaearctic, since C. brauni C. B. is widespread in Western Europe, is known in the Near East and has been found on K. According to C. Börner (Börner, 1952), lives between cones on young shoots of coniferous trees.

10. Cinara cembrae Chol. (=C. pinea Mordv.) "cedar" aphid. I - west coast in the region of the Listvennichnoye Plateau, July 11, 1963, alate forms on Japanese stone pine (Pinus pumila). Boreal palaearct. Known in Western Europe (Alps) and the northern Soviet Union (Shaposhnikov, 1964), in Eastern Siberia, and the Baikal region (Popova, 1967), in Japan on Hokkaido (Kuwayama, 1967). It is associated with cedar on the continent (P. cembrae, P. sibirica). On the islands, with Japanese stone pine. Does not inflict mass damage on I, but small damaged branches are encountered here and there, which are readily noticed due to the white tomentum secreted by the aphids.

11. Cinara piceae Panz. (=C. abieticola Chol.). "large fir" aphid. K - east coast close to Petrov Cape, August 23, 1964, colonies of apterous virgins on Sakhalin fir (Abies sakalinensis). Prevalent in Western Europe, in the northwest of the European part of the USSR; mentioned by A. A. Popova (1967) for the Baikal region. Large aphids, usually settles on the bark of thick branches, trunks, and thick roots of various fir species.

12. Cinara pinihabitans Mordv. "pine" aphid. K - shore of Lake Goryachee, July 18, 1962, apterous forms on Japanese stone pine. Palaearctic species. Previously known in the European part of the USSR, Western Siberia, Western Europe. The Scotch pine (Pinus sylvestris) and presumptively spruces and larches are mentioned as its food plants (Shaposhnikov, 1955).

13. Cinara longipennis Mats. Not found by us on the Kuril Islands. According to the data of Kuwayama (Kuwayama, 1967), was observed on Sh on the Sakhalin fir. A Kuril-Japanese species. prevalent on the islands of Sakhalin, Sh, in Japan (Hokkaido, Honshu), and in Korea.

14. Cinara laricicola Mats. (=C. taeniata Koch.). We did not find it. Cited on the basis of S. Kuwayama's study (Kuwayama, 1967). An island species, prevalent in Japan (Hokkaido, Honshu) and on I, where it lives on the larch (Larix kurilensis). In an earlier work, M. Inouye (Inouye, 1956) cites this species under the name Cinarataeniata Koch. and mentions its distribution in Europe and Japan. Judging by S. Kuwayama's study (Kuwayama, 1967), the material for which was identified by M. Inouye, the earlier designation by this author must be considered to be incorrect.

15. Tuberolachnus salignus Gmel. K - env. of the village of Alekhino, July 15, 1962, apterous virgins on willow (Salix sp.); Sh - env. of the village of Krabovoye, August 13, 1961, apterous virgins, larvae, nymphs on willow. Forms smallish colonies on small branches. Prevalent almost worldwide: the European part of the USSR, the Caucasus, Central Asia, Northern India, China (including Taiwan), the Korean Peninsula, Japan (Hokkaido, Honshu, Kyushu), the Kuril Islands (K, Sh), North and South America.

16. Lachnus pallipes Hart. K - env. of the village of Golovnino, August 8, 1961, large apterous individuals and larvae in smallish colonies on leaf stalks, in groves of shallow-cup Mongolian oak (Quercus crispula). A palaearctic species. Common in the European part of the USSR; we encountered it in Primorskiy Kray (the Suputinskiy Reserve) on maple.

Family THELAXIDAE

17. Glyphina betulae Kalt. - "birch" Glyphina. K - shore of Lake Goryachee, July 18, 1962, apterous virgins and larvae on "Tausch" birch (Betula tauschii). Aphids form smallish colonies on shoots and the inferior aspect of leaves. Palaearctic species. Prevalent in the European part of the USSR, Western Kazakhstan, Central Asia, Western Siberia, Primorye, Western Europe.

Family CALLAPHIDIDAE

18. Symydobius oblongus Heyd. (=S. alter Mordv.). K - shore of Lake Goryachee, July 17, 1962, adult apterous virgins and larvae. Often encountered on "Tausch" birch; form very large colonies, frequented by ants. Palaearctic species, prevalent in the European part of the USSR, Western Kazakhstan, Central Asia, in Altai, Western Siberia, Western Europe.

19. Betacallis comes Walk. (=Clethrobius giganteus Chol., C. betulae Mordv.). I - env. of the village of Lesozavodsk, July 19, 1963, adult alate virgins and nymphs in smallish colonies on birch shoot. Palaearctic species, known in the European part of the USSR, in the Caucasus, Central Asia, Primorskiy Kray, Siberia, and Western Europe.

20. Betacallis alnicolens Mats. We did not find it on the Kuril Range. Cited by S. Kuwayama (Kuwayama, 1967) for K (village of Grigor'evo) on alder (Alnus sp.). Kuril-Japanese species, prevalent within the limits of Honshu, Hokkaido, K.

21. Betacallis sp. K - env. of the village of Alekhino, June 27, 1962, apterous individuals, and apparently, stem-mothers on "Maksimovich" alder (Alnus maximoviczii). Possibly Kuril-Japanese species.

22. Euceraphis punctipennis Zett. (=Eu. nigritarsis Heyd.). I - env. of the village of Kuril'sk, June 30, 1963, alate virgins and nymphs; August 5, 1963, mainly alate virgins. Imagos - only alate. Live separately on the inferior aspect of birch leaves. Species widespread in the Palaearctic and beyond its borders, ubiquitously, in the European part of the USSR, in the Caucasus, Primorskiy Kray, Western Europe, and Japan (Hokkaido, Honshu), North America, and Australia.

23. Euceraphis sp. Sh - env. of the village of Malokuril'sk, August 22, 1963, alate virgins and nymphs on alder; I - env. of the village of Lesozavodsk, August 28, 1961, adult virgins and nymphs on "Maksimovich" alder.

24. Tuberculoides annulatus Hart. K - env. of the village of Golovnino, August 8, 1961, alate individuals and nymphs on shallow-cup Mongolian oak, which forms groves on ridges amidst bogs. The aphids suck singly on the inferior aspect of leaves without forming colonies. The species is prevalent in the European part of the USSR, in the Western Caucasus, Primorskiy Kray, and Central Asia; it is known in Western Europe, North America, and Australia.

25. Tuberculoides flavus Mordv. K - groves of shallow-cup Mongolian oak near the village of Golovnino, August 8, 1961, alate virgins and nymphs. The aphids live on the inferior, more rarely the superior, aspect of leaves, forming smallish colonies. Apparently a Palaearctic species. Known only in southern Primorye (Mordvilko, 1929) and on K.

26. Myzocallis sp. K - east coast close to Petrov Cape, according to the data of S. Kuwayama (Kuwayama, 1967).

27. Takecallis bambusae Mats. (=T. arundicolens Clark.).³ I - along the valley of the Kurilka River, July 5, 1963, dense colonies of apterous and alate virgins, larvae, and nymphs on Kuril bamboo (Sasa kurilensis); U - region of Lake Tokotan, August 11, 1963, apterous and alate virgins, larvae, and nymphs on Kuril bamboo, en masse. Form large dense colonies, completely covering all the leaves of the bamboo from below. This aphid evidently originates in East Asia and is associated with various species of bamboo. Following its food plant, it spread far from the original habitat. It is known at the present time in England, the Mediterranean, East Asia, the Korean Peninsula, Taiwan, Japan (Hokkaido, Honshu), and North America.

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³ S. Kuwayama (Kuwayama, 1967) cites this species under the name T. arundicolens Clark. In view of the fact that the question of the synonymy of this species remains not entirely clear (Hille Ris Lambers, 1965), we retain the old name.

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28. Calaphis betulicola Kalt. Sh - env. of the village of Malokuril'sk, August 22, 1963, apterous virgins, larvae, and nymphs on birch; U - near Lake Tokotan, August 11, 1963, apterous virgins and larvae on Erman's birch (Betula ermani). The aphids suck on the inferior aspect of leaves, forming smallish colonies. Information on distribution is clearly incomplete: the European part of the USSR, Western Siberia, and Western Europe.

29. Boernerina sp. (pr. depressa Bramst.). U - on the Okhotsk shore near Lake Tokotan, August 10, 1963, apterous virgins and their larvae, nymphs singly in colonies on the inferior aspect of "Maksimovich" alder leaves; S - env. of the village of Skalistyy, September 8, 1964, apterous sexuparae, apterous normal females and their larvae on "Kamchatka" alder (Alnus kamtschatica) leaves; P - env. of Severo-Kuril'sk, July 19, 1964, apterous virgins and their larvae form colonies along the veins on the inferior aspect of "Kamchatka" alder leaves; July 27, 1964, apterous and alate (en masse) virgins, larvae, and nymphs on alder. Only 4 species are known in the genus Boernerina: one each in Europe and Japan (B. depressa Bramst. and B. alni Takah.), 2 in North America (B. occidentalis H. R. L. in Alaska and B. variabilis Rich. in North America). The Boernerina sp. on the Kuril Islands is distinguished from all these species by a number of characters (Krivolutskaya, Ivanovskaya-Shubina, 1966), but it reveals the greatest similarity to B. depressa Bramst. In this case we are evidently dealing with vicarious species.

Family CHAITOPHORIDAE

30. Periphyllus aceris L. - "maple leaf" aphid. K - env. of the village of Alekhino, July 27, 1962, alate virgins; Cape Zyryanskiy, July 4, 1962, alate virgins; shore of Lake Goryachee, alate virgins and nymphs, everywhere on maple; I - env. of the village of Kitovyy, June 29, 1963, apterous and alate virgins; env. of Kuril'sk, July 1, 1963, apterous virgins, larvae, and nymphs, everywhere on maple. The large bright yellow aphids of this species do not form pronounced colonies, but rest in groups of several individuals each on the inferior aspect of leaves and on leaf stalks. They are encountered in relatively large numbers here and there on the western coast of I, and noticeably damage the painted maple (Acer pictum). A widespread holarctic species: ubiquitously in the European part of the USSR, in the Caucasus, the Transcaucasus, the Far East, Western Europe, and North America. Lives on various maples.

32. Periphyllus lyropictus Kessler (=P. fuscus Börn.). K - env. of the village of Alekhino, July 3, 1962, apterous and alate virgins, larvae, and nymphs, on the inferior surface of painted maple leaves; I - env. of Kuril'sk, August 19, 1961, apterous virgins, larvae, and nymphs. As in the case of the preceding species, evidently has an amphiboreal distribution, since it is known in the European part of the USSR, the Kuril Islands, Western Europe, and North America.

33. Periphyllus viridulus Mamont. (?). I - western shore close to the village of Kuybyshevo, September 7, 1961; older instar larvae on shoots of the "Sakhalin" cherry (Cerasus sachalinensis). Described by V. A. Mamontova (1955) in the Ukraine, where it lives on young shoots and leaf stalks of maple. Finding the larvae we collected on the cherry may be accidental. They could have been hatched by alate sexuparae of this species which had migrated from the maple. Distribution inadequately investigated; we assign it to the palaearctic group.

Family APHIDIDAE

34. Pterocomma salicis L. P - env. of Severo-Kuril'sk, August 5, 1964, apterous virgins, larvae on willow. Usually lives on large willow shoots and small branches. Widespread in the European part of the USSR; has been observed in Central Asia; previously unknown in the eastern part of the country.

35. Rhopalosiphum padi L. - common "bird cherry" aphid. I - env. of Kuril'sk, June 30-July 2, 1963, apterous and alate virgins, larvae, and nymphs, on the "Ssiori" bird cherry (Padus ssiori); region of the Listvennichnoye Plateau, July 2, 1963. This species has not been observed on the other islands. It severely damages the bird cherry on I. In the last ten days of June and the first half of July, dense colonies of this aphid solidly cover the young branches, leaves, and flowers. As a result of their sucking, the leaves curl, turn pale, and then yellow, and later turn brown and dry. By the end of June, the majority of the bird cherry leaves in the region of the focus have a pallid, yellow-orange, or reddish color and are curled. In the middle of July the alate virgins have left the bird cherry and have migrated to grains; by the beginning of August there were no aphids on the bird cherry, and all the damaged leaves had completely turned dry. There was no fruiting on the damaged trees. Distributed almost worldwide: in the European part of the USSR (ubiquitously), in the Transcaucasus, Kazakhstan, Central Asia, Western and Eastern Siberia, the Baikal region, Primorskiy Kray, Western Europe (everywhere), Africa, India, China (including Taiwan), the Korean Peninsula, Japan (Hokkaido, Honshu, Sikoku, Kyushu), North America, and Australia.

36. Rhopalosiphum maidis Fitch. We have not found it on the Kuril Islands. K - east coast (the villages of Tyatino and Grigor'evo); I - the villages of Kuybyshevo and Slavnoye (Kuwayama, 1967). S. Kuwayama mentions barley (Hordeum vulgare var. nudum var. hexastichon) and broomcorn millet (Panicum miliaceum) as food plants. Has not been listed previously for USSR fauna. The range of the species is very wide, encompasses Sakhalin, the Southern Kurils, Japan (Hokkaido, Honshu, Sikoku, Kyushu), the Korean Peninsula, Taiwan, New Zealand, Java, the Near East, Africa, and North America.

37. Aphis sambucia L. - the "elder" aphid. As in the case of the preceding species, cited according to the data of Kuwayama (Kuwayama, 1967). Sh - Tserkovnaya Bay, on "Siebold" elder (Sambucus sieboldiana). Evidently rare on the Kuril Islands. Distribution holarctic: the European part of the USSR (ubiquitously), in the Caucasus, the Transcaucasus, Central Asia, the Baikal region, Southern Kurils, Japan (Hokkaido, Honshu, Sikoku, Kyushu), the Korean Peninsula, North America, and Western Europe.

38. Aphis rumicis L. - the "bean" aphid. K - eastern shore (the village of Grigor'evo); Sh - the village of Malokuril'sk, on the "Japanese sorrel/dock", Rumex japonicus (Kuwayama, 1967). Distributed nearly worldwide: the European part of the USSR (ubiquitously), the Transcaucasus, Kazakhstan, Central Asia, Primorskiy Kray, Southern Kurils, Western Europe, Africa, Taiwan, India, Japan (Hokkaido, Honshu, Sikoku, Kyushu), the Hawaiian Islands, North America.

39. Aphis araliae Mats. (=Arimakia araliae Mats.). - the "aralia" aphid. K - eastern shore of Lake Goryachee, August 16, 1962, larvae and nymphs (rarely) on udo (Aralia cordata). The aphids damage young shoots, leaf stalks, and the leaves themselves. An Ussuri-Japanese species. Prevalent in the south of Primorskiy Kray, on the Southern Kuril Islands and in Japan (Hokkaido, Honshu). On the continent, lives on Manchurian aralia (A. mandshurica).

40. Aphis furcula Zett. - the "willow" aphid. K - shore of Lake Goryachee, July 18, 1962, apterous virgins and larvae in colonies on willow (Salix sp.). The aphids of this species live on young shoots and on the inferior aspect of leaves. They harm various willow species, especially young trees. Known in the Palaearctic: the European part of the USSR, Siberia, K, Western Europe. This species is clearly insufficiently studied.

41. Aphis galiae Iv. U - Okhotsk littoral near the village of Podgornyy, August 10, 1963; apterous virgins and larvae in colonies on small branches of Erman's birch. Described by O. I. Ivanovskaya (Ivanovskaya, 1958, 1959) in Novosibirsk, where it was collected from the bark of trunks and thick small branches of poplar; later found by her in the Altai Mountains on leaf stalks. Not known elsewhere. Evidently has a transsiberian distribution.

42. Aphis jacobeae Schr. Sh - env. of the village of Krabovoye; apterous virgins, larvae, and nymphs, in colonies on Siberian golden ray (Ligularia sibirica). Palaearctic species. prevalent in the European part of the USSR (ubiquitously), Siberia, and Western Europe.

43. Aphis spiraephilae Patch. K - env. of the village of Alekhino; apterous and alate virgins, larvae, and nymphs, on leaves of the "Kamchatka" meadowsweet (Filipendula kamtschatica). Sometimes form large colonies. The species has a circumboreal distribution from Western Europe to North America. Not noted in Japan.

44. Lipaphis pseudobrassicae Davis. K - the village of Alekhino; I - Kuril'sk, the village of Reydovoye, Dobroye, according to the data of Kuwayama (Kuwayama, 1967). Cucumis sativus, Brassica campestris, B. oleracea var. capitata are mentioned as food plants. Is evidently a pest of garden crops. Judging by distribution, the species obviously originates in East Asia. The range encompasses Sakhalin, Japan (Hokkaido, Honshu, Sikoku, Kyushu), China (including Taiwan), the Korean Peninsula, India, Africa, the Near East, Java, the Hawaiian Islands, and North America.

45. Sorbaphis chaetosiphon Shap. K - env. of the village of Alekhino; apterous virgins and nymphs on Sakhalin knotweed (Polygonum sachalinense). Described by G. Kh. Shaposhnikov (1950) on mountain ash (Sorbus aucuparia), on which it causes curling of leaves. Where migrants move to has not been establish. The species range has been insufficiently explored. Known in the Southern Urals, Novosibirsk, and the lower reaches of the Amur.

46. Sorbaphis kurilensis Iv. K - env. of the village of Alekhino, July 14, 1963, apterous virgins, larvae, and nymphs on "Sakhalin" cherry; I - env. of the village of Goryachiye Klyuchi, July 14, 1963, apterous virgins and nymphs on "Sakhalin" cherry (Cerasus C. sachalinensis). The aphids form colonies on the inferior surface of leaves, which as a result roll up slightly and yellow, and sometimes take on a red color. The species was described by O. I. Ivanovskaya-Shubina (1966b) on the basis of our material, and thus far it has not been found elsewhere. It may be an endemic of the Kuril Islands.

47. Heterogenaphis kunashyri Iv. K - Cape Zyryanskiy, July 13, 1962, apterous virgins, larvae, and nymphs on "Sakhalin" cherry; I - env. of the village of Alekhino, June 29-July 15, 1962, apterous and alate virgins and nymphs on "Sakhalin" cherry. The aphids live in colonies on the inferior surface of leaves. Damaged leaves thicken from above; their borders curve downward and are externally reminiscent of galls, acquiring a pink or bright red color. The genus and species were described by O. I. Ivanovskaya-Shubina (1966b) on K on the basis of our collections. Until it is found elsewhere it may be considered an endemic of K.

48. Cavariella caprae Fabr. I - flood pain of the Kurilka River, July 1, 1963, apterous and alate virgins, larvae, and nymphs on the "Kamchatka" meadowsweet (Filipendula kamtschatica). Constantly associated with willows, and facultatively with the carrot family. Distributed nearly worldwide: the European part of the USSR, Western Siberia, I, Western Europe, Africa, India, China (including Taiwan), the Korean Peninsula, Japan (Honshu), North and South America, and Australia. G. Kh. Shaposhnikov (1955) notes that various species of the genus are cited in the literature under the name of C. caprae Fabr.; it does not seem possible to identify these. It is evidently for this reason that so wide a distribution is noted for this species.

49. Myzus cerasi F. - a cherry aphid. I - env. of the village of Yasnoye and the city of Kuril'sk, June 29, 1963, larvae on "Kuril" cherry; env. of the village of Kuybyshevo, September 7, 1961, larvae on "Kuril" cherry growing along coastal slopes. The aphids form smallish colonies on the inferior aspect of leaves. It is considered a dangerous pest on the continent, especially in the south (Shaposhnikov, 1955). A Palaearctic species, distributed with man nearly worldwide. Is known at the present time in the European part of the USSR, Western Siberia, Central Asia, the Kuril Islands, Western Europe, Africa, the Near East, India, Japan (Hokkaido), North America, New Zealand, and Australia.

50. Myzus padellus H. R. L. et Roger. I - env. of Kuril'sk, June 30, 1963, apterous virgins, larvae, and nymphs on "Ssiori" bird cherry. The aphids live on leaves in open galls of a yellowish and reddish color. They migrate to the mint family. A Palaearctic species Prevalent in the European part of the USSR, in Siberia, in the Far East only on the Kuril Islands, and Western Europe.

51. Aulacorthum magnoliae Ess. et Kuw. K - the village of Yuzhno-Kuril'sk, July 25, 1961, apterous and alate virgins, larvae, and nymphs on the leaves of "Siebold" elder (Sambucus sieboldiana). Forms large colonies. A Kuril-Japanese species. Prevalent on the islands of Hokkaido, Honshu, and Taiwan, and on the Korean Peninsula. Lives on various tropical and subtropical plants, including citruses and magnolia; in a moderate climate, on elder (Shaposhnikov, 1955).

52. Aulacorthum solani Kalt. (=Au. pelargonii Mordv.). - "common potato aphid". K - villages of Alekhino, Golovnino; I - the village of Kuybyshevo. We have not collected it on the islands. Cited according to the data of Kuwayama (Kuwayama, 1967), who mentioned the potato (Cucumis sativus) and soya (Glycine Max.) as food plants. It is not mentioned as a potato pest on the Kuril Islands. Spread by man throughout the world.

53. Rhopalosiphoninus sp. K - valley of the Zyryanka River, July 2, 1962, apterous virgins, larvae, and nymphs on panicled hydrangea (Hydrangea paniculata). Species very close to Rh. ribesinus Goot., which lives on mulberry.

54. Dactynotus jacae L. K - env. of Lake Peschanoye, July 10, 1962, apterous virgins, larvae, and nymphs; env. of Lake Lagunnoye, July 27, 1962, apterous virgins, larvae, and nymphs, mixed forest; valley of the Lesnaya River, July 27, 1962, apterous and alate virgins, larvae, and nymphs; env. of the village of Golovnino, August 8, 1961, apterous and alate virgins, larvae, and nymphs, shallow-cup Mongolian oak stand; U - Okhotsk shore close to the village of Podgornoye, August 10, 1963, apterous virgins, larvae, and nymphs. In all cases the aphids were collected from "Japanese" agrimony (Agrimonia japonica); they typically live on various Compositae. They form colonies on the stems and the bases of leaves. They are oriented head downward on the stems, and when approached or touched they drop to the ground and run off. A Palaearctic species. Prevalent in the south of the European part of the USSR, in the Caucasus, Central Asia, Western Siberia, Western Europe, Western Asia and Asia Minor; has not previously been listed for the east.

55. Dactynotus mulgedus Nevs. Sh - env. of the village of Malokuril'sk, August 13, 1961, apterous virgins and larvae on "Glehn" aster (Aster glehnii) in mixed forest. It is known that this species forms huge colonies on the stems and on the base of leaves on plants of the genus Aster, and in Central Asia, on Mulgedium tataricum. A Palaearctic species, known in the south of the European part of the USSR, Central Asia, the Altai, and Kulunda (based on the collections of O. I. Ivanovskaya-Shubina); has not previously been listed for the east of the country.

56. Dactynotus picridis Fabr. I - env. of the village of Burevestnik, August 25, 1961, apterous and alate virgins, larvae, and nymphs on "Japanese oxtongue" (Picris japonica), in mixed forest with Kuril bamboo cover. The aphids form smallish colonies on stems that are not frequented by ants. Is quite widespread in the Palaearctic: the European part of the USSR, in the Caucasus, Western Siberia, Western Europe, Western Asia, and Japan (Hokkaido, Honshu).

57. Dactynotus similis H. R. L. Sh - Cape Nepokorennyy, according to the data of Kuwayama (Kuwayama, 1967). Lives on Canadian fleabane (Erigeron canadensis). We have not found it. Apparently a Palaearctic species, known in Europe.

58. Dactynotus giganteus Mats. K - Tyatya Volcano, according to the data of Kuwayama (Kuwayama, 1967). Food plant - "Kamchatka" thistle (Cirsium kamtschaticum). A Kuril-Japanese species, prevalent only on the islands of K, Hokkaido, Honshu, and Kyushu.

59. Dactynotus gobonis Mats. K - according to the data of Kuwayama (Kuwayama, 1967), without indication of the specific site of the finding. We have not found it. Lives on great burdock (Arctia lappa). Type of range Sino-Japanese; K, Hokkaido, Honshu, Sakhalin, Korean Peninsula, and China (including Taiwan).

60. Dactynotus sp. Sh - env. of the village of Malokuril'sk, August 21, 1963, apterous virgins, larvae, and nymphs; I - env. of the village of Lesozavodsk, July 19, 1963, apterous virgins, larvae, and nymphs on golden-rod (Solidago virga-aurea).

61. Dactynotus sp. K - Tyatya Volcano, according to the data of Kuwayama (Kuwayama, 1967). Aphids collected from Cirsium kamtschaticum ("Kamchatka" thistle). Not encountered at other sites.

62. Macrosiphoniella yomogicola Mats. Sh - the village of Malokuril'sk (Kuwayama, 1967). Lives on wormwood (Artemisia sp.). We have not found it. A Kuril-Japanese species, prevalent within the limits of the islands of Sh, Hokkaido, Honshu, Kyushu.

63. Macrosiphum (Sitobion) avenae Fabr. K - the village of Tyatino (Kuwayama, 1967), on common oat (Avena sativa). We have not found it. A widespread species, apparently imported into various countries by man along with the food plant. Known in the European part of the USSR (ubiquitously), in the Transcaucasus, Central Asia, Primorskiy Kray, Western Europe, East Africa, the islands of Java, Taiwan, Japan (Hokkaido, Honshu, Sikoku, Kyushu), and the Korean Peninsula.

64. Macrosiphum sp. K - the village of Tyatino (Kuwayama, 1967). Aphids collected from broomcorn millet (Panicum miliaceum).

65. Macrosiphum sp. K - the village of Alekhino (Kuwayama, 1967), on Cucumis sativus.

66. Megoura viciae japonica Mats. K - the village of Alekhino, according to the data of Kuwayama (Kuwayama, 1967), on vetch (Vicia Fabr.). We have not encountered it. A Manchurian species, prevalent on K, in Japan (Hokkaido), on the Korean Peninsula, and in China.