CHAPTER 3

SECTION 5. Order COLEOPTERA

Family IPIDAE (bark beetles)

The bark-beetles of the Kuril Islands have remained essentially unstudied until recently. There is only fragmentary information in the literature (Winkler, 1932; Sawamoto, 1940a, 1940b; Kurentsov, 1941, 1950; Stark, 1952; Krivolutskaya, 1956, 1958) on the finding of a few species on the Kuril Range, usually without indication of the specific places where they were found. In 1965 we published an article (Krivolutskaya, 1965a), in which 48 species of bark-beetles were cited for the Southern Kuril Islands, with data on their distribution, biotopic delimitation, trophic links, phenological periods of development, etc. S. Kuwayama (Kuwayama, 1967) did not actually enlarge our list in his work on the insects of the Southern Kurils. Of the two species additionally cited by him, Cryphalus piceae Ratz. is a European species; the specimens identified under this name should apparenly be assigned to Cryphalus piceus Egg., and under the name Ips cembrae Heer., undoubtedly to Ips subelongatus Motsch. (Stark, 1952).

In all, 53 species of bark-beetles, belonging to 20 genera, are known on the Kuril Islands at the present time: Scolytus (5 species), Hylesinus (1), Hyorrhynchus (1), Hylurgops (1), Alniphagus (1), Polygraphus (6), Crypturgus (2), Cryphalus (6), Ernoporicus (2), Eocryphalus (2), Phellodendrophagus (1), Hypothenemus (1), Dryocoetes (8), Pityophthorus (1), Trypodendron (4), Pityogenes (3), Ips (2), Orthotomicus (2), Xyleborus (2), and Scolytoplatypus (2). Of these, 3 species were found to be new to science (Krivolutskaya, 1968), 1 species was identified to the genus level and is apparently also new, but thus far cannot be described, since only one specimen has been collected; 1 genus and 5 species are cited for the first time for the fauna of the USSR.

The bark-beetle fauna of the Kuril Archipelago is appreciably impoverished as compared with adjacent islands and the continental regions of the Far East. Thus, 68 species of bark-beetles are known on Sakhalin, 95 on Hokkaido, and 128 in Primorskiy Kray. Some genera (Dendroctonus, Blastophagus, Hylastes, Ernoporus, Trypophloeus, etc.) are not observed here at all; others (Hylesinus, Hylurgops, Crypturgus, Ips, Xyleborus) are represented by only one to two species. Some representatives of the genera Scolytus, Hylastes, Hylurgops, Crypturgus, Ips, and Hyleborus, which are quite widely distributed on Sakhalin and Hokkaido, may be found in the future on the Southern Kuril Islands.

The bark-beetles are among insects which are closely linked to their food plants and, accordingly, to specific plant associations. Therefore, the character of the ipidofauna is determined, first and foremost, by the composition of the woody species forming forest stands, and their distribution on the islands. At the same time, the distribution of the bark-beetles over the territory of the archipelago is to a certain degree in accordance with general patterns which are characteristic for the Kuril entomofauna as a whole. The bark-beetle fauna is distributed in the following manner over the territory of the islands :
 
  Shikotan Kunashir Iturup Urup Paramushir Species, in all 12 40 24 9 2 % of total number of species 22.1 75.5 45.4 16.9 3.6  

Note.  Bark beetles have not been found on the islands of Simushir and Shumshu.The wealth of the ipidofauna on the southern islands is explained by the fact that dark-coniferous, broadleaved-dark coniferous, and larch forests, with which the majority of the species are associated, flourish precisely in this part of the archipelago. The taiga complex of bark-beetles, which are vitally linked with the spruces, the Yezo spruce, (Picea microsperma), the Yeddo spruce (P. ajanensis), and the Sakhalin spruce (P. glehni), and the Sakhalin fir (Abies sachalinensis); this complex, which is appreciably impoverished by comparison with Sakhalin, stands out quite distinctly. It is composed of 18 species, of which 13 live on spruce (Polygraphus sachalinensis Egg., P. poligraphus L., Crypturgus pusillus Gyll., Cryphalus piceus Egg., Dryocoetes rugicollis Egg., D. autographus Ratz., D. hectographus Reitt., Trypodendron proximum Niiji., T. lineatum Ol., Pityogenes chalcographus L., Ips typographus f. japonicus Niiji., Orthotomicus suturalis Gyll., O. laricis F.), 3 on fir (Polygraphus proximus Blandf., Dryocoetes striatus Egg., Scolytoplatypus tycon Blandf.), and 2 on both spruce and fir (Hylurgops palliatus Gyll., Cryphalus kurenzovi Stark.). The ipidofauna in the broadleaved-dark coniferous forests, especially in the southwest of Kunashir, where leaved species predominate in the stands in some places, is more diverse. Groupings of bark-beetles living on various leaved trees and shrubs predominate in such forests, and the taiga complex occupies a subordinate position.

But, the ipidofauna varies on different islands, depending on the features of the forest vegetation, and forms unique groupings. For example, on the southwest shore of Kunashir, 24 species of bark-beetles were recorded on leaved species, while on Iturup, there were only 11, although among these, southern species are encountered as well (Table 12). The comparison that has been made demonstrates that on Iturup, after many broadleaved species (elm, cork tree, magnolia, Bothrocaryum, white mulberry, and ash) have dropped out of the composition of the stands, the bark-beetles which are associated with them also disappear. On the remaining leaved species, on the other hand, which flourish on both islands, the bark-beetle groupings proved to be quite constant. The alder is an exception, on which a number of polyphagous species absent or rare on Iturup, settle in the forest of Kunashir.

The bark-beetles, which are associated with the Kuril Dahurian larch, are known only on Iturup, the only island where there are pure, large larch stands. They form a small but specific complex, which combines Polygraphus sachalinensis Egg., Dryocoetes baicalicus Reitt., Pityogenes bidentatus Herbst., and Ips subelongatus Motsch., of which the last 2 species develop en masse in windfallen and sawn larch trunks.

The groupings of bark-beetles living on the Japanese stone pine, alder, and mountain ash merit attention. On the southern islands the alder and mountain ash do not form independent stands, but rather flourish usually as an admixture in mixed forests. The Japanese stone pine here occupies elevated parts of the mountain slopes, but not infrequently descends to sea level and also grows among mixed broadleaved-dark coniferous forests. These species acquire a different position on the middle and northern islands of the range, where they form pure elfin wood stands.

Specific bark-beetle species, which develop on one or two closely-related species, are characteristic of each of those enumerated. On the alder these are Alniphagus alni Niiji., Cryphalus alni Krivol., Eocryphalus zachvatkini Krivol., and Dryocoetes ussuriensis Egg.; on the mountain ash, Cryphalus padi Krivol. and Polygraphus nigrielytris Niiji.; and on the Japanese stone pine, Pityogenes foveolatus Egg. and Dryocoetes orientalis Kurenz.

In addition, polyphagous bark-beetles which shift from other plants live on nearly each species. Their groupings vary depending upon the conditions in which the food plant flourishes. Thus, on Kunashir Polygraphus poligraphus L., Cryphalus piceus Egg., and Pityophthorus lichtensteini Ratz. occupy the Japanese stone pine; these are species which can develop on other coniferous species as well, in particular on the spruce. This complex is increased two-fold on Iturup (Polygraphus poligraphus L., P. sachalinensis Egg., Crypturgus hispidulus Thoms., C. pussilus Gyll., Dryocoetes orientalis Kurenz., Pityogenes foveolatus Egg.), on account of species which are specific for the Japanese stone pine and which shift from the larch and spruce. On Urup the grouping of "Japanese stone pine" bark-beetles is not numerically increased; its composition is simply varied somewhat by comparison with Iturup (Polygraphus sachalinensis Egg., P. gracilis Niiji., P. poligraphus L., Cryphalus kurenzovi Stark, Dryocoetes orientalis Kurenz., Pityogenes foveolatus Egg.). Four species are known as inhabitants of spruce. This suggests that dark-coniferous forests evidently flourished on Urup in the past. After their disappearance, the "spruce" bark-beetles evidently entirely shifted to the Japanese stone pine and continue to exist on it until the present day. With further movement to the north of the range (Paramushir), only Pityogenes foveolatus Egg., which is typical for it, remains on the Japanese stone pine.

A similar alternation in groupings of bark-beetles is observed on the alder and mountain ash. As was indicated above, 8 bark-beetle species live on the alder on Kunashir (3 are specific for this species and 5 are polyphagous); on Iturup, only 2 (both are typical "alder" species); and on Urup, also 2, Eocryphalus zachvatkini Krivol. and Dryocoetes ussuriensis Egg. The first of these is exceptionally rare and is still known only on Sakhalin; only D. ussuriensis Egg. which reaches the north of the range (Paramushir) is a typical inhabitant of cool moist alpine forests on the continent that also invaded Kamchatka.

Only 2 specific species are associated with the mountain ash; both are encountered on Kunashir and Iturup, while only one remains on Urup, Polygraphus nigrielytris Niiji. On the islands located to the north of Urup, it was not possible to find bark-beetles on the mountain ash, despite a careful search, although it is entirely probable that P. nigrielytris Niiji. will be found in the northern part of the range, since it is known on Kamchatka. Polyphagous bark-beetles have not been observed on the mountain ash.

Thus, the overwhelming majority of bark-beetles known on the Kuril Archipelago is concentrated on the southern islands (51 species). Only species associated with the Japanese stone pine, alder, and mountain ash penetrate the middle and northern islands. A substantial proportion of the species (29) has thus far been found only on one of the islands: there are 20 such species on Kunashir, 1 on Shikotan, 6 on Iturup, and 2 on Urup. It cannot be excluded that many of these, especially those which develop on spruce and fir, will be found also on neighboring islands within the areas that dark coniferous forests flourish.

The ipidofauna of the Kuril Islands possesses some features which distinguish it from the ipidofauna of Sakhalin. These features consist in a somewhat different species composition of the bark-beetles, in the interrelationship of their groupings, associated with coniferous and leaved species, in the population sizes of particular species, and in the phenological periods of development. Thus, bark-beetles are encountered on the Southern Kurils which are absent on Sakhalin, but which in their majority are known on Hokkaido. These are species which develop on broadleaved species: Hylesinus elatus Niiji., Hyorrhynchus lewisi Blandf., Polygraphus ssiori Niiji., Cryphalus exiguus Blandf., C. kurilensis Krivol., Ernoporicus spessivtzevi Berg., Hypothenemus insularum Krivol., Dryocoetes aceris Krivol.

At the same time, there are not many widely distributed "conifer" bark-beetles on the Kuril Archipelago. If it is the case that species associated with coniferous species (42 on coniferous, 19 on leaved) predominate on Sakhalin, in the case of the Kurils and Hokkaido, species associated with leaved species predominate (on the Kurils: 25 on coniferous, 27 on leaved; on Hokkaido: 41 on coniferous, more than 50 on leaved). To a substantial degree this brings the bark-beetle fauna of the Southern Kuril Islands closer to the ipidofauna of Hokkaido.

Some species which are encountered in broadleaved-dark coniferous forests in the extreme southwest of Sakhalin and which are rare there, are pests under the conditions of the Kuril Islands. The following can be classified in this group: Polygraphus nigrielytris Niiji., Cryphalus alni Krivol., Eocryphalus semenovi Kurenz., Phellodendrophagus elegans Krivol., Scolytoplatypus daimio Blandf. Apparently, at the present time the central region of the range of the enumerated species is on the territory of Kunashir and Hokkaido; this is governed by the optimal life conditions. Isolated finds of such species on Sakhalin and in individual cases in Primorskiy Kray, on the other hand, point only to the boundaries of their ranges and suggest a past wider distribution of the fauna and flora existing in these regions. The finding of Ernoporicus spessivtzevi Berg. in large numbers on Kunashir, which is previously known only in Primorskiy Kray, but which is scanty there, and which has not been noted either on Sakhalin, or in Japan, is highly significant in this connection.

On the Kuril Islands, thanks to the absence of recent burns and to limited logging operations, favorable conditions have not been created for the occurrence of foci of mass reproduction of bark-beetles on large areas. In nearly all types of forests, especially in stands which include coniferous species, the preconditions exist for the appearance of bark-beetle colonies. At the present time, diffuse small foci of the reproduction of bark-beetles appear at sites of windfalls and storm damage, in areas with overmature and weakened, as well as dead standing trees, close to solfatars on trees which have been depressed by the constant discharge of hydrogene sulphide and other noxious gases, on coastal slopes in Japanese stone pine elfin stands, depressed by the prolonged action of the winds blowing from the sea, and at the sites of logging operations, where unbarked wood is left in the forest in the summer. Of the bark-beetles which are capable of multiplying en masse under the conditions of the Kuril Islands, the following can be observed on coniferous species: Ips typographus f. japonicus Niiji., I. subelongatus Motsch., Pityogenes chalcographus L., P. bidentatus Herbst., Polygraphus proximus Blandf., P. sachalinensis Egg., Cryphalus piceus Egg., Crypturgus pusillus Gyll., Trypodendron proximum Niiji., Orthotomicus laricis F.; on leaved species: Polygraphus nigrielytris Niiji., Hylesinus elatus Niiji., Cryphalus exiguus Blandf., C. padi Krivol., C. alni Krivol., Ernoporicus spessivtzevi Berg., Eocryphalus semenovi Kurenz., Phellodendrophagus elegans Krivol., Scolytoplatypus daimio Blandf.

The phenological periods of development of the bark-beetles on the Kuril Islands coincide with those on Sakhalin, and are somewhat delayed by comparison with the continental regions of the Far East. Habitation in a marine climate environment which is characterized by a late spring, a moist and relatively warm summer, and a prolonged warm autumn is the reason for this.

In terms of its origin, the bark-beetle fauna of the Kuril archipelago can be divided into two principal complexes, the Boreal (23 species) and the Palaearchaearctic (30 species). They in their turn are divided into a number of groups and subgroups, depending on the character of the ranges of the species included in these complexes (Table 13).

The Boreal complex combines primarily widely distributed representatives of the taiga fauna which have a holarctic and transpalaearctic distribution, and which also inhabit territories surrounding the aquatorium of the Sea of Okhotsk. This complex almost entirely consists of bark-beetles associated with coniferous species (21 species on coniferous, 2 on leaved species).

The Holarctic group includes Hylurgops palliatus Gyll., Dryocoetes autographus Ratz., Trypodendron lineatum Ol., and Xyleborus saxeseni Ratz.; the Transpalaearctic includes Polygraphus poligraphus L., Crypturgus pusillus Gyll., C. hispidulus Thoms., Dryocoetes baicalicus Reitt., D. hectographus Reitt., Pityophthorus lichtensteini Ratz., Pityogenes chalcographus L., P. bidentatus Herbst., Ips typographus f. japonicus Niiji., I. subelongatus Motsch., Orthotomicus suturalis Gyll., and O. laricis F. These species are widely distributed on the Southern Kuril Islands within the areas where coniferous forests flourish. At some sites they represent the principal composition of the ipidofauna and predominate over the remaining zoogeographical groupings.

The Okhotsk species, which are apparently a derivative of a Beringian fauna, also live almost exclusively on coniferous trees. To these we assign Polygraphus sachalinensis Egg., P. gracilis Niiji., P. proximus Blandf., Cryphalus kurenzovi Stark, C. piceus Egg., Dryocoetes ussuriensis Egg., Pityogenes foveolatus Egg., and provisionally Dryocoetes orientalis Kurenz. They are quite evenly distrubuted over the archipelago, and Pityogenes foveolatus Egg. and Dryocoetes ussuriensis Egg., which are confined to Japanese stone pine forests, reach the northern islands.

Bark-beetles with a less broad range, which encompasses the basins of the Ussuri and Amur Rivers, partially the territory of North East China, Sakhalin, the Kuril Islands, and Japan, belong to the Palearchearctic complex. In accordance with the distribution over the territory mentioned, they are combined in three groups, the Manchurian, Ussuri-Island, and Island groups. The close association of the overwhelming majority of the species with leaved species (26 species on leaved, 4 on coniferous) and overall with moist broadleaved forests is a characteristic feature of the Palaearchaearctic complex. The entire complex undoubtedly has its common origin in the time of the existence of lush broadleaved forests on the islands. Remainders of the past fauna have survived to the present day (even if in altered form) in the relict coniferous-broadleaved forests of Hokkaido, Kunashir, and partially Iturup (Tolmachev, 1959).

Species which are fairly widely distributed on the continent and in Japan are included in the Manchurian group: Scolytus shikisanii Niiji., S. trispinosus Strohm., S. aratus Blandf., Alniphagus alni Niiji., Dryocoetes rugicollis Egg., Trypodendron proximum Niiji., T. niponicum Blandf., Scolytoplatypus tycon Blandf. Scolytus dahuricus Chap., which is common in the Primor'ye, the Priamur'ye, and on Sakhalin, but which does not invade Japan, constitutes the only exception.

Bark-beetles which we have set apart in the Ussuri-Island group are distinguished by a unique range: Scolytus ventrosus Schev., Ernoporicus spessivtzevi Berg., Eocryphalus semenovi Kurenz., Phellodendrophagus elegans Krivol., Dryocoetes striatus Egg. They are encoutered in the basin of the Ussuri (usually in the south of the Sikhoté-Alin'), on Sakhalin, and the Kuril Islands, but they are thus far unknown in Japan, except for D. striatus Egg. Such a distribution points to the existence in the geological past of direct connections between the continent and the modern territory of the Kuril archipelago. The discovery of some of them on Hokkaido, which is closer in its natural conditions to Sakhalin and the Southern Kurils than to the remaining Japanese islands, is entirely probable. This is confirmed by the discovery on Hokkaido of D. striatus Egg.

Bark-beetles which have an island origin are set apart in the Island group.⁵ They are combined into four subgroups depending on distribution on the islands: 1) the Kuril-Japanese group which includes the species encountered in Japan and on the Southern Kuril Islands (Hylesinus elatus Niiji., Hyorrhynchus lewisi Blandf., Polygraphus ssiori Niiji., Cryphalus exiguus Blandf., Dendrotrypum pubipenne Blandf.); 2) the Sakhalin-Kuril-Japanese group, which combines species distributed on Sakhalin, in Japan, on the Kuril Archipelago, and partially on Kamchatka and the Korean Peninsula (Polygraphus nigrielytris Niiji., Xyleborus germanus Blandf., Scolytoplatypus daimio Blandf.); 3) the Sakhalin-Kuril group, to which bark-beetles known only on Sakhalin and the Kuril Range belong (Cryphalus padi Krivol., C. alni Krivol., Eocryphalus zachvatkini Krivol.); and 4) the Kuril proper group, which is represented by new species found thus far only on Iturup and Kunashir (Cryphalus kurilensis Krivol., Ernoporicus sp., Hypothenemus insularum Krivol., Dryocoetes aceris Krivol.). The representatives of the Kuril proper subgroup cannot be regarded as indisputable Kuril endemics, since during future investigations they may be found on Sakhalin, Hokkaido, and even on the continent, as was the case with Phellodendrophagus elegans Krivol.

⁵ Some of these species, which definitely have an island origin, are encountered also on Kamchatka and the Korean Peninsula, i.e., on those territories which closely adjoin the modern islands, with which in relatively recent geological time dry-land connections existed (Minato, 1966).

The zoogeographical analysis of the Kuril ipidofauna that has been presented offers a fairly clear notion of the elements composing it and reflects general patterns which are characteristic for the Kuril entomofauna as a whole. Information on the general distribution of the bark-beetles that is cited in the systematic list is based on our own collections and observations, as well as on that borrowed from published studies (Kurentsov, 1941, 1950, 1951; Stark, 1952; Kuwayama, 1967).

  SYSTEMATIC LIST OF SPECIES

  1. Scolytus shikisanii Niiji. - the "Japanese elm" engraver. K - env. of Alekhino set., August 7, 1962, 14 spec., coniferous-broadleaved forest, from trunk of drying Manchurian elm. Not encountered in other sites, apparently because of the absence of the food plant. According to our observations, the beetles begin to build nests in weakened and drying elm trees at the end of July-beginning of August. Development continues up until autumn. The larvae and pupae overwinter. WhenS. shikisanii Niiji. maintained in the laboratory were extracted from cut-sections, older instar larvae and pupae were found under the bark in the first ten-day period of April, and the mass flight of the beetles began on April 22 and continued for 4 days. The emerged beetles mated and again crept under the bark, where they underwent additional feeding, gnawing out wide galleries in the surface of the sapwood to the depth of 2-3 mm, or burrowing into the wood to the depth of 3-4 mm. Judging from the literature (Kurentsov, 1941; Stark, 1952; Krivolutskaya, 1965a), the pattern of the galleries of this engraver is quite variable. By observing the behavior of the bark beetles in containers, we established that the flight of the beetles began at 10 a.m. At that time they exited the galleries, flew and actively crawled along the surface of trunks, but again hid in the galleries by 1600 hrs. A Manchurian species. Distributed in Primorskiy Kray, on Sakhalin, K, and in Japan (Hokkaido, Honshu).

2. Scolytus ventrosus Schev. - the "eastern Siberian" engraver. K - env. of Alekhino set., August 1, 1961, 1 male, 1 female, broadleaved forest; the beetles were establishing a nest under the bark of a Manchurian elm near notching. Just as with the preceding species, it occupies overmature and weakened Manchurian elm trees, primarily the lower portions of thick trunks. The flight of the beetles takes place at the end of July-beginning of August. During this time they gnaw out mother galleries and deposit eggs. The larvae overwinter and complete development in the spring of the following year. Thus, we were able to observe the nests of S. ventrosus Schev. with older instar larvae and the pupae at the end of June, 1962. Belongs to the Ussuri-Island group. Known in Primorskiy Kray, on Sakhalin, and K. Everywhere quite rare.

3. Scolytus trispinosus Strohm. - the "three-toothed" engraver. K - env. of Alekhino set., June 27-August 7, 1961, 1962, 17 spec. in all, coniferous-broadleaved forest. One of the most numerous species among the engravers observed on the Kuril Islands. Lives on the Manchurian elm. Settles on thick trunks and small branches of living and dying trees, more often near flawing birch-bark defects, mechanical injuries to the bark, and notchings. May cause noticeable physiological damage to weakened trees. The developmental periods of this species are similar to those of S. ventrosus Schev.: the flight of the beetles and oviposition take place at the beginning of August, and the development of the larvae from August through June of the following year. Older instar larvae, pupae, and young beetles are encountered at the end of June in the nests of S. trispinosus Strohm. During flight and oviposition, some of the beetles undergo additional feeding, gnawing out irregular crook-shaped galleries under the bark (Figure 13, 1). A typical representative of the Manchurian zoogeographical grouping. Distributed on K, Sakhalin, in Primorskiy Kray, Eastern Siberia, in North East China, and in Japan (Hokkaido, Honshu).

4. Scolytus aratus Blandf. - the "Korean" engraver. K - env. of Alekhino set., August 3, 1961, 1 female, on Manchurian elm in coniferous-broadleaved forest. According to our observations, very rare on the Kurils. A characteristic Manchurian species. Ditsributed on K, Sakhalin, in the basins of the Ussuri River and middle reach of the Amur River, in Primorskiy Kray, on the Korean Peninsula, and in Japan (Hokkaido, Honshu, Sikoku).

5. Scolytus dahuricus Chap. - the "Dahurian" engraver. K - valley of Alekhin Stream, July 13, 1962, 1 female; I - valley of the Kurilka River, July 5, 1963, 6 males, 9 females, mixed forest, in a birch pile. Encountered rarely in the Kuril Range. It has not been possible to trace the developmental periods under the conditions of the islands. On I, the beetles were undergoing additional feeding at the beginning of July. One beetle each was found in galleries 1-4 cm in length. A Manchurian species whose range encompasses the basins of the Ussuri and Amur rivers, Dauriya, Sakhalin, K, and I. It is unknown in Japan, but might be found on Hokkaido.

6. Hylesinus (Longulus) elatus Niiji.⁶ - "elm" bark-beetle. K - env. of Alekhino set., June 27-August 7, 1961, 1962, a series of 385 spec., on Manchurian elm in coniferous-broadleaved forest. Described on Hokkaido (Niijima, 1915). We had set it apart in the subgenus Longulus along with the close Far Eastern species H. costatus Blandf. Has been described again in detail, since the initial description had been done schematically (Krivolutskaya, 1968).

⁶ We had previously cited it as Ulmiphagus ulmi (Krivolutskaya, 1965a); later this name was reduced to a synonym of H. elatus Niiji. (Krivolutskaya, 1968).

  Lives in dark coniferous-broadleaved forests of the southwest part of K on the Manchurian elm, Ulmus laciniata, and on Hokkaido on U. campestris. Damages weakened, drying, and windthrown trees with fairly fresh phloem. Substantially damages young saplings with a trunk diameter at the brest height of 8-15 cm. Occupies primarily thick small branches with a diameter of 12-15 cm on stout trees. It settles in the lower parts of trunks in close proximity to species of the genus Scolytus. The flight of the beetles lasts about a month and a half, from the end of June to the beginning of August. The development of beetles of the younger generation has sufficient time to be completed in nearly all nests before autumn. Larvae and pupae overwinter only in some nests. The young beetles remain to overwinter in the mother nests (hatch sites). They undergo additional feeding in the spring before emerging. Overwintering larvae turn into beetles only by the middle of July. The structure of the galleries of the "elm" bark-beetle is similar to that of the galleries of many species of the genus Hylesinus (Figure 13, 2). Two transverse, relatively short mother galleries, weakly imprinted on the sapwood and gnawed out by different females, take off from the nuptial chamber, which looks like a small triangle. Larval galleries are encountered frequently and diverge fanwise in opposite directions from the mother galleries. They are gnawed through in the phloem or the bark and also are not imprinted on the sapwood. The nests in occupied trees are closely adjacent to one another, such that nearly no intact phloem remains between them. H. elatus Niiji. is a common and even numerous species on K at sites where the Manchurian elm flourishes. The parasitic wasp Dinotiscus aponius Wlk. parasitizes in the galleries of this bark-beetle. Based on the character of the range it is assigned to the Island group, and has a local distribution within the limits of K and Hokkaido, but will possibly be found in the southern region of Sakhalin.

7. Hyorrhynchus lewisi Blandf. - "Lewis's ambrosia beetle" (Figure 14,1-3). K - env. of Alekhino set., July 3, 1962, August 4, 1961, 16 spec., coniferous-broadleaved forest, on the "controversial" Bothrocaryum. Encountered only in the southwest of K, where the "controversial" Bothrocaryum flourishes. According to the data of Japanese investigators (Niijima, 1909), it develops on elm (Ulmus campestris) in Japan. It is associated exclusively with the Bothrocaryum on the Kuril Islands, despite the presence of an elm (Ulmus laciniata) in broadleaved forests. According to its biology, this bark beetle belongs to the "ambrosia beetles" and lives in close proximity to Scolytoplatypus daimio Blandf., selecting the same trees to colonize. The beetles gnaw out round galleries, which enter the wood deeply and which are disposed perpendicularly to the longitudinal axis of the trunk. A bluntly terminating lateral canal branches off the main galleries in the same plane (Figure 13, 3). The larvae do not gnaw out independent galleries and apparently feed on the hyphae of fungi on the walls of the mother gallery. The colonization of trees by the beetles continues throughout July and August; the young beetles appear in September, but the majority of them evidently remain to overwinter in the hatch sites, i.e., in the wood. The beetles attack standing, markedly weakened and recently dried out trees with wood which is still moist. They usually settle in the trunk, more rarely in thick small branches. They are commercial pests. Fungi which destroy the wood penetrate the trunk along old galleries of H. lewisi Blandf. Damage by this bark beetle is encountered quite frequently in the broadleaved forests of K. A Kuril-Japanese species. Distributed on K, Hokkaido, and Honshu.

8. Hylurgops palliatus Gyll. - "small spruce ambrosia beetle". K - valley of the Lesnaya River, July 25, 1962, 8 spec., felling site in spruce-fir forest, from a Sakhalin fir stump. The only representative of the genus Hylurgops found on the Kuril Range. It is apparently encountered sporadically. Pupae and young beetles undergoing additional feeding were found on July 25 in nests of H. palliatus Gyll. A Holarctic species, distributed in the forest zone from Europe to North America.

9. Alniphagus alni Niiji. - "Ussuri alder ambrosia beetle". K - shore of Lake Goryacheye, env. of Alekhino set., region of Cape Petrov, outside find dates June 29-August 26, 34 spec. in all; I - env. of Lesozavodsk set., August 28, 1961, 8 spec. Lives on the Kuril Islands in dark coniferous-broadleaved and floodplain forests which include alder. Occupies weakened but still growing, drying old trees and the small trunks of the understory. The nests of this "ambrosia beetle" are located in the bark and phloem, and do not go down as far as the sapwood. The developmental periods are shifted even by comparison with Sakhalin. Flight begins in the first ten-day period of July and continues to the beginning of August. The development of the larvae is completed by the end of August-beginning of September in the majority of cases; older instar larvae, pupae, and young beetles are found in the nests. The last of these overwinter in the hatch sites and undergo additional feeding in the mother nests in the spring; they leave the mother nests only at the end of June-beginning of July. Thus, we were able to observe nests of A. alni Niiji. with actively feeding young beetles on K on June 29, 1962. The parasite Dinotiscus aponius Wlk. is occasionally encountered in the nests. On the Kuril Islands this species is less numerous than in the forests of Sakhalin, and the damage which it inflicts is insignificant. Belongs to the Manchurian group. Distributed in Primorskiy Kray where various species of alder flourish, on Sakhalin, I, K, and in Japan (Hokkaido, Honshu).

10. Polygraphus gracilis Niiji. - the "elegant bark-beetle". U - Okhotsk littoral near Cape Zhdi, August 14, 1963, 1 spec., in Japanese stone pine growths. Very rare in the Kuril archipelago. Evidently should be encountered on other islands as well, especially where dark coniferous forests flourish, since the spruce and fir are the principal food plants. Based on the character of the range it is included in the Okhotsk group. Known from the northern region of Primorskiy Kray (basin of the Khor River), on Sakhalin, U, and Hokkaido.

11. Polygraphus sachalinensis Egg. - the "Sakhalin bark-beetle". K - env. of Lake Peschanoye, July 10, 1962, 3 spec.; Sh - env. of Malokuril'sk set., August 12-14, 1961, 2 spec.; I - env. of Kuybyshevo and Burevestnik settlements, env. of Kuril'sk, Listvennichnoye Plateau, outside collection dates August 19-25, 18 spec. in all; U - env. of Lake Tokotan, August 12, 1963, 1 spec. One of the representatives of the genus Polygraphus which is widely distributed on the Kuril Islands. It lives in the dark coniferous forests of K and Sh on the Yezo spruce and the Japanese stone pine, on I in Japanese stone pine forests of Pinus pumila, as well as on the larch, and on the Japanese stone pine on U. The beetles gnaw out mother galleries and deposit eggs in the first half of July. The development of the larvae continues throughout July and August. New generation beetles appear in the nests of P. sachalinensis Egg. at the end of August. Evidently the majority of these remain for the winter in the mother nests, where they undergo additional feeding in the autumn and the following spring. Despite the relatively frequent incidence, this bark beetle is distinguished by its small numbers and does not inflict substantial damage. The parasitic chalcid Dinotiscus eupterus Wlk. can be encountered fairly frequently in its galleries; it evidently substantially reduces the number of the larvae. Belongs to the group of Okhotsk species. Distributed in the south of Primorskiy Kray, on Sakhalin, Kamchatka, Kuril Islands (K, Sh, I, U), and in Japan (Hokkaido).

12. Polygraphus poligraphus L. - the common double-eyed spruce bark-beetle. K - env. of Sernovodsk and Alekhino settlements, valley of the Lesnaya River, July 7-25, 1962, 14 spec. in all; Sh - env. of Malokuril'sk set., August 13-27, 1961, 4 spec.; I - env. of Peschanyy set., August 23, 1961, 1 spec.; U - Okhotsk littoral near Cape Zhdi, August 14, 1963, 2 spec. Quite common in dark coniferous forests, where it lives on the Yezo spruce, and in Japanese stone pine growths. Frequently encountered together with P. sachalinensis Egg. Occupies drying trees and lumber; it builds its nests in trunks and small branches. The beetles gnaw out mother galleries and lay eggs in the first half of July. The development of the larvae continues until September; the larvae, pupae, and new generation beetles overwinter. The parasitic wasp Dinotiscus eupterus Wlk. parasitizes in the nests. Just as in Sakhalin, despite the relatively frequent incidence, P. poligraphus L. colonies are not numerous in the forests of the Kuril Islands. A Transpalaearctic species, widely distributed in the forest zone of European USSR, throughout Siberia, in Khabarovskiy and Primorskiy Krays, on Sakhalin, Kamchatka, K, Sh, I, and U; known in Western Europe, Western Turkey, and Northern Mongolia.

13. Polygraphus proximus Blandf. - "white-fir bark-beetle". K - valley of the Lesnaya River, env. of Lake Lagunnoye, Sernovodsk set., env. of Alekhino set., Pacific Ocean coast near Cape Petrov, outside find dates July 25-August 27, 46 spec. in all; Sh - in the region of Cape Kray Sveta, August 27, 1963, 8 spec.; I - Lesozavodskiy Neck, env. of Lesozavodsk set., outside collection dates July 24-August 28, 22 spec. in all. It is encountered everywhere on the Kuril Range where dark coniferous forests flourish. It is the principal pest of the Sakhalin fir. Attacks weakened, drying, windthrown trees and unbarked lumber. Forms local foci of mass reproduction at the sites of windfalls and in logging regions. The gnawing out of nests and oviposition by the beetles take place in the second half of June. The development of the larvae is completed by the end of July. Older instar larvae and pupae are found in the nests in the last ten-day period of July, and sexually immature beetles of the new generation appear. Young beetles undergo additional feeding in the mother nests throughout all of August, and the overwhelming majority of them evidently remain here to overwinter. They fly out in the spring, with the commencement of the warm days when the daytime air temperature rises above +15° . This is one of the early-flying species. The range of P. proximus Blandf. encompasses the region of the distribution of far eastern firs, including Primorskiy and Khabarovskiy Krays, Sakhalin, K, Sh, I, the Korean Peninsula, and Japan (Hokkaido, Honshu, Sikoku, Kyushu).

14. Polygraphus nigrielytris Niiji. - the "black", or "mountain ash bark-beetle". K - env. of Sernovodsk set., western slopes of Mendeleev Volcano, July 27-30, 1961, 1962, 31 spec. in all; I - env. of Kuril'sk, env. of Lesozavodsk and Kuybyshevo settlements, outside collection dates July 4-September 7, 72 spec. in all; U - shore of Lake Tokotan, Cape Zhdi, August 10-15, 1963, 25 spec.

A common species of dark coniferous-broadleaved forests of K and the leaved elfin wood stands of I and U. Attacks weakened but still viable mountain ash, cherry, and bird cherry trees en masse. Occupies trunks from base to top and small branches. The flight of the beetles is markedly protracted, but is approximately a month behind by comparison with Sakhalin. The first beetles of the new generation appear in the nests at the end of July; however their emergence continues throughout all of August and the first half of September. Thus,, we observed a mass flight and oviposition of P. nigrielytris Niiji. in 1961 on I. On August 19 and on September 7, we found larvae, pupae, and young beetles in the nests of this bark beetle. Beetles emerged in 1962 from August 4 to the beginning of October from bird cherry sections collected on July 27 on K, but their greatest mass emergence was observed at the end of August. Soon after flying out (2-3 days later), the beetles begin to gnaw out mother galleries and to deposit eggs. P. nigrielytris Niiji. families are polygamous; this is reflected in the structure of the galleries in the nest. The male (he is usually smaller than the female) gnaws out the nuptial chamber; two to three females fly to him; after mating these gnaw independent mother galleries through, which take off from the nuptial chamber in different directions (Figure 13, 4). The beetles emplace the mother galleries in the phloem and in the bark layer, not infrequently at its very surface, such that the bark along the gallery frequently bursts and narrow clefts are formed which are reminiscent of knife slashes. The nests are located in the trunks in the bark and phloem layer, and do not impinge upon the sapwood. Typically the entire pattern of the galleries is imprinted on the sapwood surface in small branches where the bark is thin. The parasite Rhoptrocerus xylophagorum Ratz. has been observed on I in the galleries of the "black bark-beetle", but it apparently has little significance in reducing the numbers of this bark beetle.

Mountain ash and bird cherry trees occupied by this bark beetle are readily distinguished by the abundance of rust-red bore meal spilling from the galleries and covering the trunk with a continuous rusty bloom, and sometimes accumulating at the base. The secretion of gum, which has the appearance of transparent gelatinous excrescences, takes place on damaged small cherry trunks. The injuries caused by P. nigrielytris Niiji. which we observed on Sakhalin were not so massive in character. On Kamchatka, according to the data of A. I. Kurentsov and D. G. Kononov (1961), it is infrequently encountered and does not inflicts substantial damage. Apparently the central portion of the range of this species (the zone of its injuriousness) is in the region of the Kuril Islands and Northern Japan. A typical Island species. Distributed on Sakhalin, Kamchatka, U, I, K, Hokkaido, and Honshu.

15. Polygraphus ssiori Niiji. - "Japanese cherry bark-beetle". I - env. of Kuril'sk, July 4-August 20, 1961, 1963, 35 spec. in all. Found in elfin leaved stands which grow on coastal marine terraces and slopes. Lives on weakened and drying saplings of the Sakhalin cherry, in close proximity to P. nigrielytris Niiji. It occupies the lower portions of the trunks and thick small branches around twigs [knots?]. The phenology of both species is evidently similar, since adult and young beetles, eggs, and developing larvae were found on August 20, 1961 in the galleries of P. ssiori Niiji. The galleries of this bark beetle have a characteristic pattern: a short (1.5-2.0 cm in length) and fairly wide mother gallery takes off from the nuptial chamber, on both sides of which closely spaced, slightly diverging larval galleries are disposed (Figure 13, 5). It is distinguished from other species of the genus Polygraphus by markedly shiny, glinting integuments and by the absence of hair and scales on the pronotum and elytra. A fairly small number of fine bright scales are preserved only on the very slope of the elytra. A Kuril-Japanese species, which evidently has an island origin. Known on I, Hokkaido, Honshu, and Sikoku.

16. Crypturgus pusillus Gyll. - the "spruce bark-beetle-chafer". K - valley of the Lesnaya River, July 27, 1962, 19 spec.; I - Okhotsk littoral near Kuybyshevo set., August 22, 1961, 1 spec. Under Kuril Islands conditions it is associated mainly with the Yezo spruce, as well as with the Japanese stone pine. It accompanies Ips typographus and some other larger bark beetle species. We observed the mass flight of C. pusillus Gyll. at an active felling site on July 27, 1962. It was possible to see hundreds of "spruce bark-beetle-chafers" crawling along the logs and flying on piles of unbarked lumber. A Transpalaearctic species, a typical inhabitant of coniferous forests whose range occupies all of European USSR including the Crimea and the Caucasus, all of Siberia to the shores of the Pacific Ocean, Primorskiy Kray, Sakhalin, I, and K; it is distributed beyond the boundaries of the USSR in Western Europe, Algeria, Western Turkey, on the Korean Peninsula, in China (Taiwan), and in Japan (Hokkaido, Honshu).

17. Crypturgus hispidulus Thoms. - the "conifer bark-beetle-chafer". I - Okhotsk littoral near Kuybyshevo set., Pacific Ocean littoral near Burevestnik set., August 22-25, 1961, 22 spec. It is found on the Kuril Range only on Japanese stone pine, where it accompanies Pityogenes foveolatus Egg. We occasionally found adult beetles which were apparently undergoing additional feeding in C. hispidulus Thoms. nests at the end of August. The phenology of this bark beetle in island conditions was not traced in greater detail. A Transpalaearctic species, distributed in the taiga zone of Eurasia, Primorskiy Kray, on Sakhalin, Kamchatka and I.

18. Cryphalus kurenzovi Stark. - the "Kurenzov" Cryphalus. K - env. of Alekhino set., valley of the Lesnaya River, July 6-27, 1962, 17 spec.; I - env. of Lesozavodsk set., July 24, 1963, 20 spec.; U - env. of Lake Tokotan, August 12, 1963, 2 spec. Quite common on the southern islands of the Kuril archipelago, but is encountered sporadically. Develops here on all dark-coniferous species. More numerous in lumbering regions where it inhabits the small branches of sawn spruce. Older instar larvae, pupae, and young and adult beetles are found in the nests of this species at the end of July. At the beginning of August the young beetles emerge from the breeding sites, but begin to reproduce after overwintering. An Okhotsk species, known in the central and southern Sikhoté-Alin', on K, Sakhalin, I, and U. Finding this Cryphalus also in the mountainous coniferous forests of Hokkaido cannot be excluded.

19. Cryphalus piceus Egg. - the "Japanese" Cryphalus. K - env. of Sernovodsk and Alekhino settlements, shore of Lake Goryacheye, Okhotsk coast near Lake Peschanoye, valley of the Lesnaya River, July 7-August 4, 1962, 35 spec. in all; Sh - env. of Malokuril'sk set., August 12-14, 1961, 36 spec.; I - env. of Lesozavodsk set., August 29, 1961, 8 spec. Common in dark-coniferous forests of the Southern Kuril Islands. Lives on the spruce, occupying the small branches and trunks of weakened, drying, and windfallen trees, as well as unbarked lumber. In certain instances may also colonize the Japanese stone pine.

The flight of the beetles, their gnowing out of nests, and deposition of eggs begin in the last days of June and continue to the middle-end of July. The development of the larvae proceeds throughout July and August, but concludes at various times, depending on the time the eggs were deposited. In some nests the beetles of the new generation appear as early as the second half of July, and at the end of July leave the breeding sites. For example, in 1962 on K , we observed mass flight of C. piceus Egg. on July 27. In other nests, larvae, pupae, and young beetles feeding additionally can still be encountered at the end of August-beginning of September, as occured on August 29, 1961 on I. Not infrequently, when the eggs are deposited late, the larvae do not have the time to develop before the beginning of the cool weather; they overwinter, and pupate only in the spring of the following year. The "Japanese" Cryphalus may form local foci in the forests of the Kuril Islands, reproducing en masse in clutter areas of the forest and in lumbering regions. An Okhotsk species. Distributed in coniferous forests of Primorskiy and Khabarovskiy Krays, on Sakhalin, I, K, Sh, in Japan (Hokkaido, Honshu, Sikoku), and in North East China.

20. Cryphalus exiguus Blandf. - the "mulberry" Cryphalus. K - Okhotsk littoral near Lake Peschanoye, env. of Alekhino set., July 8-August 11, 1962, 46 spec. in all.; Sh - env. of Malokuril'sk set., August 24-28, 1963, 25 spec. The white mulberry (Morus alba), which it attacks en masse, is the principal food plant. Ocassionally and in small numbers it also colonizes the "Maksimovich" alder. The flight of the beetles and deposition of eggs by them take place in the first half of July. It occupies slightly broken small branches and weakened trunks of trees, preferentially the lower parts of the crown and trunks. The beetles do not lay mother galleries, but gnaw nuptial chambers of irregular form which have somewhat bluntly terminating depressions in the form of pockets (Figure 13, 6).

The families are monogamous: in all of the nuptial chambers which we opened only two beetles each, a male and a female, were found. The female lays eggs in the depressions of the nuptial chamber in a tight row or in clumps and does not cover them with bore meal as do many other species of bark-beetles. The larvae initially gnaw a common family gallery in the phloem, but before pupation form branches from it, not infrequently burrowing into the wood to a depth of 1.5-2.0 mm, where they gnaw pupal cells. The development of the larvae continues 35-40 days, and as early as the end of the first ten days of August the beetles of the new generation appear in the nests. They feed additionally for several days under the bark, entirely eating out the interlayer of phloem, and partially the sapwood, after which they fly out and again embark on the building of nests and deposition of eggs.

Under the conditions of the warm and prolonged Kuril autumn, the larvae continue to feed to the end of September; by this time they achieve the older instars, and some of them possibly have sufficient time to pupate. In the spring development terminates, and at the end of June-beginning of July the beetles of the second generation emerge. Thus, C. exiguus Blandf., by our observations, puts forth two generations a year on the Southern Kurils. But, apparently, it is not in the entire population that two generations develop. In connection with the great protraction of the flight (some of the beetles continue to emerge and deposit eggs to the end of July), the development in nests that are founded late is markedly delayed, in this case the young beetles which appear later remain to overwinter in the maternal nests or emerge, but do not have time to lay eggs. The nuptial chambers, larval galleries, and pupal cells in this species are imprinted on the sapwood, especially in nests in thin branches. The young beetles when feeding additionally gnaw more narrow intertwined galleries which are also deeply incised in the sapwood.

On Sh, the nests of C. exiguus Blandf. are in close proximity to colonies of Phellodendrophagus elegans Krivol. Multiplying en masse, the "mulberry" Cryphalus inflicts substantial damage on the mulberry stands, all the more so since cases of its attacking not only weakened but healthy trees as well were observed. The chalcid of the fam. Pteromalidae (the genus and species of the parasite have thus far not been identified) that is frequently encountered in the nests is evidently not capable of markedly decreasing the population size of the host. A Kurilo-Japanese species, distributed on K, Sh, in Japan (Hokkaido, Honshu, Sikoku, Kyushu), and on the Korean Peninsula. The reference by S. Kuwayama (Kuwayama, 1967) for China requires more precise determination.

21. Cryphalus padi Krivol. - the "Sakhalin bird cherry" Cryphalus. K - env. of Alekhino set., August 2, 1962, 186 spec.; I - env. of Kuril'sk, August 19, 1961, 41 spec. We have described it on Sakhalin (Krivolutskaya, 1958) where it is associated with the "downy bird cherry" (Padus pubescens). It is observed on the Kuril Islands only on the mountain ash (Sorbus commixta). It occupies drying, thin, small branches. Usually reproduces in large numbers.

K. E. Schedl (Schedl, 1962a) reduced it to a synonym of C. scopiger Berg., on the basis of review of one specimen; from our point of view, this is unjustified. At the present time we have at our disposal large series of C. scopiger Berg. from Primorskiy Kray, and C. padi Krivol. from the Kuril Islands. Comparison of the serial materials reveals quite clear distinctions between these species:

   
   C. padi Krivol. C. scopiger Berg. The frons is shiny, delicately stippled, with a prominent fine keel, which reaches the vertex; there is no rugose sculpturation around the mouth Frons matte, coarsely stippled, with a shorter keel; rugose around the mouth The rim of denticles around the anterior margin of the pronotum consists of 5-7 tubercles, of which the 4 middle ones are larger The rim of denticles on the anterior margin of the pronotum consists of 4-8 denticles, of which the 2 middle ones are larger Small punctate furrows of the elytra are slightly depressed; the intervals are slightly terete or entirely flat; they sometimes appear slightly convex only at the very base of the elytra and are stippled with delicate fine dots Small punctate furrows of the elytra are distinct, deep; the intervals are markedly convex, terete all over the entire extent of the elytra, especially at the base; covered, in addition to dots, by transverse oblique rugae, which are more coarse in the anterior half     The humeral tubercles at the base of the elytra are large, shiny, and protrude prominently The humeral tubercles of the elytra do not protrude, are matte, and poorly discernible Elytra shiny Elytra matte Lives on the mountain ash and bird cherry Strictly associated with the Manchurian walnut  

This analysis gives grounds for regarding C. padi Krivol. as an independent species; the features of its biology point to this as well.

Young beetles, and pupae and larvae singly, are found en masse in the nests of C. padi Krivol. under Kuril Islands conditions in August (2-19). The emergence of the adult beetles from butt-ends in the laboratory was observed during the second half of August and in September. The phenological periods of development were not followed in greater detail. The chalcid Dinotiscus sp. parasitizes in the galleries of this Cryphalus. Belongs to the Island species of the Sakhalin-Kuril subgroup. Known on Sakhalin, I, and K.

22. Cryphalus alni Krivol. - the "alder" Cryphalus. K - env. of Alekhino set., July 8, 1962, 5 spec.; shore of Lake Goryacheye, July 19, 1962, 8 spec. As was the case with the preceding species, has been described on Sakhalin (Krivolutskaya, 1958). Here and there on the Kuril Islands its habitats are floodplain mixed stands of leaved species with the predominance of "Maksimovich" alder, its principal food species. It often colonizes trees growing at the very edge of the water. Damage inflicted by this bark-beetle was found in several trees. It colonizes trunks 15-20 cm in diameter, small sapling trunks and small branches. The galleries are laid in the phloem layer and do not touch the sapwood. Females which have deposited eggs and first instar larvae were found in the nests on July 19. The parasite Rhopalicus sp. was removed from the galleries. Belongs to the Sakhalin-Kuril endemics. Known thus far only on Sakhalin and K. Rare everywhere.

23. Cryphalus kurilensis Krivol. - the "Kuril" Cryphalus. K - env. of Alekhino set, July 22, 1962, 44 spec.; I - basin of the Kurilka River, July 5, 1963, 79 spec. We have described it on the Kuril Islands (Krivolutskaya, 1968). Lives on trichocarpous toxicodendron (Toxicodendron trichocarpum). Inhabits weakened and drying small trunks, measuring 3-10 cm in diameter. The flight of the beetles, their founding of nests and depositing eggs take place at the beginning of July (according to observations on I). The development of the larvae takes place throughout July and August. The emergence of the beetles (judging by observations in the laboratory on colonized butt-ends) takes place from the end of August to the end of September. The flight of the beetles is evidently markedly protracted; therefore the precise alternation of generations has not been traced. It is possible that the secondary deposition of eggs and sister generations occur. The overwintering of various phases of development, larvae, pupae, and young beetles, points to this. It cannot be excluded that two generations of beetles a year develop on K . Thus, on July 22, 1962 in the env. of Alekhino set., pupae and adult beetles which had laid eggs were found simultaneously in the galleries of the "Kuril" Cryphalus.

Families in this species are monogamous; only two beetles each are found in the nuptial chambers, a male and a female. The pattern of the galleries is reminiscent of the galleries of C. exiguus Blandf.: the beetles gnaw a nuptial chamber and a small depression taking off from it in the form of a pocket, in which the eggs are deposited in a clump. The larvae gnaw a family gallery in one direction. In some cases, the galleries are laid in the phloem layer, in others they are imprinted on the sapwood. The larvae pupate under the bark without burrowing into the wood. The beetles develop in the colonized trees en masse, although parasites and predators do destroy the larvae: the parasitic wasp, Cosmophorus cembrae Rusch., and the flat bark-beetle, Prostominia lewisi Rtt. A Kuril endemic, thus far found only on K and I.

24. Ernoporicus spessivtzevi Berg. - the "Spessivtsev" Cryphalus. K - env. of Alekhino set., July 13-August 7, 1961, 1962, 44 spec. in all. Previously known only in Primorskiy Kray where it is associated with the ash. It lives in the forests of K on the Manchurian elm. Inhabits weakened, thin, fine sapling trunks and drying small branches; on the thicker small branches it colonizes in close proximity to Hylesinus elatus Niiji., through whose galleries it evidently penetrates beneath the bark. The galleries are laid in the bark layer and in the superficial layers of the phloem, without touching the sapwood. It forms extensive colonies at the sites of reproduction. The flight is apparently markedly protracted, since fully developed nests with larvae of various instars, pupae, and young beetles have been simultaneously observed from the middle of July to the middle of August. By multiplying in large numbers, it promotes the drying of the elm sapling, by which means it undoubtedly inflicts substantial damage. It is common in the broadleaved forests of K where the elm flourishes. We have assigned it to the Ussuri-Island subgroup, since at the present time it is known in Primorskiy Kray and on K. The discontinuity of its range is explained most likely by the fact that it has been poorly studied.

25. Ernoporicus sp. K - env. of Alekhino set., August 3, 1961, 1 spec., extracted from magnolia butt-end. Close to E. spessivtzevi Berg., but is distinguished from it by larger dimensions, by the elongate clava of the antennae, by the dense covering of small scales on the elytra, and by the number of denticles on the anterior margin of the pronotum. Since the genus Ernoporicus is monotypic, the bark-beetle we found is a new species, but at the present time we are not in a position to describe it, due to the discovery of only one specimen.

26. Eocryphalus semenovi Kurenz. - the "Semenov" Cryphalus. K - env. of Alekhino set., June 28-August 31, 1962, 10 spec.; I - env. of Lesozavodsk set., Okhotsk coast near Kuybyshevo set., August 28-September 7, 1961, 23 spec. in all. Encountered on the Kuril Islands in broadleaved and dark coniferous-broadleaved forests, where the sen flourishes. Colonizes weakened and drying small sen trunks, 2.5-3.5 cm in diameter, and live small branches of trees, more often overmature or weakened, but may also attack healthy trees. The beetles penetrate the bark at the base of buds and young shoots. They gnaw through longitudinal mother galleries in the bark layer or on the surface of the phloem. The phenological periods of development are somewhat intertwined; this evidently takes place because of the prolonged flight. It also cannot be excluded that two generations of beetles develop over the course of the year. Thus, we have observed the flight of the beetles and oviposition in the middle of July and at the very end of August; we have observed larvae and pupae in the nests at the end of June and in the beginning of September, and young beetles in the first half of September. The larvae feed on the phloem and on the superficial layers of the sapwood; before pupation they burrow 1.0-1.5 mm into the wood. The beetles feed additionally under the bark and convert it into rotten wood, leaving untouched only the thin superficial bark. The damage inflicted by this bark-beetle is insignificant; in particular instances it may cause drying of small branches. An Ussuri-Island species, distributed in southern Primorskiy Kray, on southern Sakhalin, and on the Okhotsk littoral of I and K.

27. Eurocryphalus zachvatkini Krivol. - the "Zakhvatkin" Cryphalus. U - the Okhotsk coast near Cape Zhdi, August 15, 1963, 16 spec., procumbent growth of "Maksimovich" alder. Described on Sakhalin, where was also collected from "Maksimovich" alder (Krivolutskaya, 1958). Judging by the solitary finds, the species is rare, apparently strictly confined to sea shores. Inhabits small branches of varied thickness and small trunks. The pattern of the galleries is reminiscent of galleries of representatives of the genus Cryphalus, but by contrast with the typical cryphalids, a short mother gallery takes off from the nuptial chamber; the eggs are laid like "stitching" on both sides of it (Figure 13, 7). The larvae gnaw individual galleries in the phloem, which are not imprinted on the sapwood. Typical cryphalids lay eggs in a clump, and their larvae gnaw a common, family, gallery. The families are evidently monogamous, since in the majority of cases two beetles are found in the nuptial chambers, rarely three, and one mother gallery takes off from the nuptial chamber in each nest. In view of the rare incidence, the phenological periods of development of the "Zakhvatkin" Cryphalus were not traced. At the time of collection (August 15), larvae, adult beetles, and eggs were found in the nest. The settlements do not form dense colonies; the nests are located at a considerable distance from one another. An Island species, belonging to the Sakhalin-Kuril subgroup. Known on Sakhalin and on U.

28. Phellodendrophagus elegans Krivol. - the "glossy" bark-beetle. K - env. of Alekhino set., July 1-August 4, 1961, 1962, 107 spec. in all; Sh - env. of Malokuril'sk set., August 21, 1963, 11 spec.; I - env. of Lesozavodsk set., August 30, 1961, 97 spec. Described on Sakhalin (Krivolutskaya, 1958). Subsequently the genus Phellodendrophagus Krivol. was assigned by K. E. Schedl (Schedl, 1962b) to the tribe Pityophthorini and was reduced to a synonym of the Japanese genus Eidophelus Eichh. Examination of large series of this species collected in recent years on the Kuril Islands and on the continent has shown that by general habitus the "glossy" bark-beetle can actually be assigned to the tribe Pityophthorini; however, the structure of the slope of the elytra and the clava of the antennae markedly distinguishes it from the representatives of this tribe. With regard to the genus Eidophelus Eicch., judging by the description of Eichhoff (Chapuis et Eichhoff, 1875), it is not identical to the genus Phellodendrophagus Krivol. (we, unfortunately, do not have specimens of the genus Eidophelus Eichh.). The differences between the two genera in question are the following:

 
   Phellodendrophagus Krivol. Eidophelus Eichh.     The clavae of the antennae are sutureless, with the external aspect having a noticeably bare spot, it remaining surface being covered with rather dense fine white hairs Clava has barely noticeable sutures, without a bare spot The anterior margin of the pronotum is even, without tubercles, only sometimes appears slightly undulating; its bases and sides are clearly marginated Anterior margin of the pronotum has tubercles; the pronotum is not marginated The base of the elytra is straight, and not interrupted at the scutellum; the ends of the elytra are smoothly incised, the sutural [??] angle does not stand out; the slope of the elytra is markedly convex, without any signs of tubercles or other sculpturated formations The base of the elytra at the scutellum is rounded or obliquely incised; the sutural [??] angle is somewhat elongated; the slope is flattened, covered with fine tubercles  

Taking all the above into account, we cannot consider the reduction of the genus Phellodendrophagus to a synonym of the genus Eidophelus to be substantiated. The position of this genus in the system was recently determined by A. Nobuchi (Nobuchi, 1969) in a comparative study of the masticatory stomachs of adult beetles of the superfam. Scolytoidea. On the basis of comparative morphological data, he assigns the genus Phellodendrophagus to the tribe Cryphalini and to the type Ernoporus, bringing it in proximity to the genera Ernoporus, Cryphalus, Ernoporicus, and Eocryphalus.

On Sakhalin it lives in the extreme southwest on the Sakhalin cork tree (Krivolutskaya, 1058). On the Kuril Islands (K, Sh, I), is distributed substantially more widely and is a common species of dark coniferous-broadleaved forests. Here it is already associated with several woody species: the Sakhalin cork tree, the silver magnolia, the white mulberry, the trichocarpous toxicodendron, and the spindle tree. Does the greatest damage to the cork tree, attacking young weakened saplings en masse. The beetles prefer to settle on young small trunks and small branches with thin bark. Their settlements were not observed on trunks with a thick cork layer. The pattern of the galleries is very reminiscent of the galleries of bark-beetles of the genus Eocryphalus: one to two short S-shaped curved mother galleries take off from a triangular nuptial chamber. The character of the larval galleries is difficult to establish, since they are gnawed in the phloem, are not imprinted on the sapwood, and are completely deformed during additional feeding of the young beetles.

The flight of the beetles takes place in the second half of July. The larvae develop throughout July and August. They feed on the phloem and the inner layers of bark, and burrow 1-2 cm into the wood only for pupation. In such shrubs as the trichocarpous toxicodendron and the spindle tree, which have loose pith in small trunks, the larvae pupate in the pith. For example, we were able to observe a large cluster of young beetles and pupae in the pith of trichocarpous toxicodendron in the breeding period (at the end of August- beginning of September). The beetles feed additionally under the bark, consuming the cork layer. The overwhelming majority of them remain in the maternal nests to overwinter. At sites of mass settlements of the "glossy" bark-beetle, an entire complex of entomophages develop in its nests: parasitic wasps (Dinotiscus eupterus Wlk. and Rhopalicus sp.), the larvae of gnats and other dipterans, the larvae and imagines of the flat bark-beetle (Prostominia lewisi Rtt.), and the larvae and imagines of a large thrips.

Until recently, P. elegans Krivol. was considered an Island species. The region of the Southern Kuril Islands is evidently the central part of its range; this is suggested by the large number of food plants with which it is associated here. The majority of them are lacking in the forests of Sakhalin. Recently this species was found on the littoral of the Sea of Japan on the continent (the City of Nakhodka), where it had colonized weakened saplings of the Manchurian aralia en masse (Krivolutskaya, Kupyanskaya, 1979). It can be expected to be found on Hokkaido as well. In light of the new data we have assigned it to the subgroup of Ussuri-Island species.

29. Hypothenemus insularum Krivol. - the "island" Cryphalus (Krivolutskaya, 1968). K - env. of Alekhino set., July 3, 1962, 17 spec. The third species of the genus Hypothenemus found on the territory of the USSR. Very close to Hypothenemus corni Kurenz. from Primorskiy Kray. It lives on the "controversial" Bothrocaryum (Bothrocaryum controversum) of the dogwood family (Cornaceae). Colonizes the tops of drying small trunks and small branches along with Scolytoplatypus daimio Blandf. The nests are encountered singly and are disposed separately. The mother and larval galleries are gnawed in the phloem layer; only for pupation do the larvae burrow into the wood to a distance of 1.5-2.0 mm, and stop up the entry with a small plug of bore meal. The phenological periods of development have not been followed fully. On July 3, 1962, beetles could be seen running along the bark and gnawing nuptial chambers. At the same time, pupae and young beetles of the new generation, singly, could be found in the old nests. The emergence of the beetles from butt-ends continued throughout July in the laboratory. Belongs to the subgroup of Kuril endemics.

30. Dryocoetes striatus Egg. - the "fir" Dryocoetes-bark-beetle. Sh - valley of the Otradnyy Stream, August 27, 1963, 20 spec., dark coniferous forest, from freshly-sawed logs of Sakhalin fir. It is evidently rare on the Kuril Islands or is encountered sporadically, as in other parts of the Range. The possibility of finding it on K and I cannot be excluded. It lives on various species of Far Eastern firs. The periods of development on the archipelago are evidently delayed as compared with Sakhalin. Eggs, larvae of various instars, pupae, young beetles of the new generation, and old darkened beetles which had in fact laid eggs were found simultaneously by us in the nests of D. striatus Egg. at the end of August. Colonizes trunks with fairly fresh phloem and still liquid resin; can attack weakened growing trees. The nests are located in close proximity to settlements of Polygraphus proximus Blandf., but are distinguished by a highly characteristic pattern of the galleries: three to four mother galleries take off in the longitudinal direction from the nuptial chamber; along their sides the eggs are laid in dense "stitching", slightly covered by bore meal. The mother and larval galleries are quite distinctly imprinted on the sapwood. The entrance and flight openings are larger than in P. proximus Blandf. The range of the "fir" Dryocoetes-bark-beetle is typical for the Ussuri-Island species; it encompasses the southern spurs of Sikhoté-Alin', Sakhalin, Sh, and Hokkaido.

31. Dryocoetes baicalicus Reitt. - the "Baikal" Dryocoetes-bark-beetle. I - env. of Kuril'sk, August 5, 1963, 12 spec.; Listvennichnoye Plateau, August 23, 1961, 11 spec., N. A. Popov. We had previously cited it as D. kurilensis sp. n. in litt. (Krivolutskaya, 1965a). This name was later reduced to a synonym of D. baicalicus Reitt. (Krivolutskaya, 1968). Is encountered sporadically and in small numbers. Occupies thick branches of windthrown Kuril Dahurian larch trees. The phenological periods of development under the conditions of the Kuril Islands have not been tracked. According to our observations, development is completed by the end of July, since at the beginning of August mainly young, not yet darkened, beetles are found in the nests. A Transpalaearctic species, distributed in the taiga zone from European USSR to the Pacific Ocean; along the Sikhoté-Alin', reaches the southern spurs, and is known on Sakhalin and I.

32. Dryocoetes orientalis Kurenz. - the "Japanese stone pine" Dryocoetes-bark-beetle. I - Listvennichnoye Plateau, env. of Kuril'sk, Okhotsk littoral near Kuybyshevo set., Pacific Ocean littoral near Kasatka set., outside find dates July 11-August 23, 27 spec. in all; U - Okhotsk coast near Cape Zhdi, August 15, 1963, 7 spec. The biology of this bark-beetle, like its range, is still insufficiently studied, evidently due to its rare occurence and small numbers, as well as its morphological closeness to D. baicalicus Reitt., with which it may be confused. The Japanese stone pine is the main food plant on the Kuril Archipelago; on occassion it settles on the Kuril Dahurian larch. In Primorskiy Kray it lives mainly on the Korean pine, but was found once on the Manchurian fir (Kurentsov, 1941). On Sakhalin K. Tamanuki collected it from the Yeddo spruce (Krivolutskaya, 1958). One specimen of D. orientalis Kurenz. was found in the Transbaikalia on the Siberian larch (Lur'e, Lindeman, 1961). However, the latter report requires revision and checking. The phenological periods of development under Kuril Islands conditions have not been tracked. It is only known that the development of the larvae is completed at the end of August, and that young beetles which overwinter in the breeding sites appear in the nests.

Undoubtedly a Far Eastern species by origin. We have conditionally included it in the Okhotsk group, since the reference for the Transbaikalia raises doubts. Known on the southern spurs of Sikhoté-Alin', on Sakhalin, I, U, and in the Transbaikalia. S. Kuwayama (Kuwayama, 1967) cites the name D. orientalis Kurenz. as a synonym of D. infuscatus Mur., a species which is unknown to us.

33. Dryocoetes ussuriensis Egg. - the "Ussuri" Dryocoetes-bark-beetle. K - western slope of the Mendeleev Volcano, July 26, 1962, 7 spec.; I - southern slope of the Atsonupuri Volcano, August 31, 1961, 29 spec.; U - Okhotsk littoral close to Cape Zhdi, August14-15, 1963, 31 spec.; P - env. of Severo-Kuril'sk, July 19, 1964, 19 spec. On the Southern Kuril Islands it lives in mountainous mixed forests on "Maksimovich" alder; in the central and northern parts of the range, also on "Kamachatka" alder in floodplain and coastal elfin stands, nearly at sea level. Inhabits trunks and small branches of trees that are weakened and undercut by water; the galleries are laid in the phloem. The larvae develop over the course of July and August. At the end of August-beginning of September, the beetles of the new generation are found in the nests; they evidently remain here to overwinter. An Okhotsk species, distributed in Primorskiy and Khabarovskiy Krays, in Magadanskaya Oblast, on Kamchatka, Sakhalin, K, I, U, and P.

34. Dryocoetes aceris Krivol. - the "maple" Dryocoetes-bark-beetle. I - Okhotsk littoral in the region of Listvennichnoye Plateau, July 11, 1963, 1 spec.; Okhotsk littoral near Kuybyshevo set., September 7, 1961, 22 spec. Very close to D. ussuriensis Egg. and, apparently, genetically closely associated with it. Attacks painted maple trees which are suppressed by the constant action of the wind but which are still entirely viable, and occupies the lower parts of the trunks (from the middle to the root collar). The beetles gnaw small nuptial chambers and short mother galleries (1.5-2.0 cm in length) which are imprinted on the sapwood (Figure 13, 8). The larvae gnaw their own galleries in the phloem, not touching the sapwood. Their development is completed at the beginning of September; older instar larvae, pupae, young and adult beetles are found simultaneously in the nests. The majority of the young beetles feed additionally under the bark and evidently remain there to overwinter. The rest leave the maternal nests and found new families, gnawing out mother galleries and laying eggs in the very same trees. Thus far known only on I, but its discovery at other sites is not excluded. In particular, the specimens collected by A. I. Kurentsov from a maple in Primorksiy Kray and assigned by him to D. ussuriensis Egg. (Kurentsov, 1941) need to be checked.

35. Dryocoetes rugicolis Egg. - the "large spruce" Dryocoetes-bark-beetle. K - env. of Lake Goryacheye, valley of the Lesnaya River, slope of the Golovnin Volcano, from the stomach of a white-backed woodpecker (Dryobatos leucotos), V. A. Nechaev; Pacific Ocean littoral near Cape Petrov, outside collection dates July 26-November 8, 37 spec. in all; Sh - env. of Malokuril'sk set., August 13, 1961, 6 spec. Has already been cited by a number of authors for the Kuril Range (Kôno, 1938a; Sawamoto, 1940a, 1940b; Stark, 1952), without indication of the specific places found on the islands. A common inhabitant of Far Eastern dark coniferous forests. Settles on thick trunks of windfallen, windbroken spruce trees and on unbarked lumber. Prefers shaded, damp sites, encountered more often under the forest canopy. A Manchurian species, distributed in southern Primorskiy Kray, on Sakhalin, K, Sh, in North East China, and in Japan (Hokkaido, Honshu, Sikoku).

36. Dryocoetes autographus Ratz. - the "conifer" Dryocoetes-bark-beetle. K - env. of Lake Goryacheye, August 2, 1961, 6 spec. Rare on the Kuril Islands. The beetles were collected along with D. hectographus Reitt. and D. rugicollis Egg. from the trunks of thrown Sakhalin spruce trees. A Holarctic species. Widely distributed in European USSR, including in the Crimea and the Caucasus, in Siberia, the Transbaikalia, on Sakhalin, K, and outside of the USSR, in Western Europe, China (including Taiwan), on the Korean Peninsula, in Japan (Hokkaido, Honshu), and in North America.

37. Dryocoetes hectographus Reitt. - the "taiga" Dryocoetes-bark-beetle. K - env. of Lake Goryacheye, August 2, 1961, 12 spec.; Sh - env. of Malokuril'sk set., August 14, 1961, 1 spec. It is encountered in dark coniferous forests of the Kuril Islands on spruce, more frequently than the preceding species, but is also sparce. A Transpalaearctic species, whose range encompasses the taiga and forest zones of European USSR, all of Siberia to the shores of the Pacific Ocean, Primorskiy Kray, Sakhalin, Sh, K, North East China, and Japan (Hokkaido, Honshu).

38. Pityophthorus lichtensteini Ratz. - the "Lichtenstein" Pityophthorus. K - shores of Lake Goryacheye, July 17, 1962, 22 spec. Had not been observed previously east of the Southern Pribaikal'ye. Known as a pest of various coniferous species - pine, cedar, spruce, and fir. On K the beetles occupy small trunks and small branches of the Japanese stone pine which grows near solfatars and which has been weakened by the action of toxic sulfurous gases. They possibly also attack still viable plants, since we found colonies of this Pityophthorus on recently dried small branches with yellowed, but not yet fallen needles. In the middle of July larvae of various instars and adult beetles were found in the nests; at the same time the beetles continued to gnaw new mother galleries and lay eggs. Observations on butt-ends showed that the development of the larvae continues until autumn (September-October), but it is only completed the following spring in nests where eggs have been deposited late. Young beetles which have appeared in the autumn season remain to overwinter in the maternal nests, where they undergo supplementary feeding. Thus, the older instar larvae, pupae, and young beetles overwinter. In isolated areas P. lichtensteini Ratz. may do appreciable damage to Japanese stone pine stands, promoting their necrosis. A Transpalaearctic species, widely distributed in the forest zone of the Palaearctic.

39. Trypodendron proximum Niiji. - the "Ussuri" timber beetle. K - valley of the Lesnaya River, July 25-27, 1962, 13 spec., felling site in dark coniferous forest, from logs and stumps of the Yezo spruce. A typical inhabitant of Far Eastern dark coniferous forests. Multiplying in large numbers, it is one of the principal pests of lumber. On K the flight of the beetles, their founding of nests, and laying of eggs take place in the second half of July. Logs and stumps damaged by the "Ussuri" timber beetle are readily distinguished by the white bore meal which abundantly spills from its galleries. It is included in the Manchurian group. Distributed in Primorskiy and Khabarovskiy Krays, on Sakhalin, K, in Japan (Hokkaido, Honshu, Sikoku, Kyushu), and on the Korean Peninsula.

40. Trypodendron niponicum Blandf. - the "Japanese" timber beetle. K - env. of Alekhino set., valley of the Lesnaya River, slope of the Mendeleev Volcano, outside collection dates July 24-August 4, 12 spec. in all. Quite rare on the Kuril Islands. Lives on weakened "Maksimovich" alder trees, the painted maple, and Erman's birch; sometimes colonizes stumps. The phenological periods of development have not been established. Adult beetles are encountered in the galleries at the end of July-beginning of August. A Manchurian species, known in the Southern Pribaikal'ye, the Priamur'ye and Primor'ye, on Kamchatka, Sakhalin, K, the Korean Peninsula, and in Japan (Hokkaido, Honshu, Sikoku, Kyushu).

41. Trypodendron lineatum Ol. - the spruce ambrosia beetle. K - valley of the Lesnaya River, July 25, 1962, 1 spec., in stacks of unbarked lumber. Lives on the Yezo spruce in dark coniferous forests. Apparently more rare than on Sakhalin. Has a Holarctic distribution. The species range encompasses European USSR, the Caucasus, Kirghizia, all of Siberia to the Pacific Ocean littoral, Primorskiy Kray, Kamchatka, Sakhalin, K, Europe, Mongolia, Japan (Hokkaido, Honshu), and North America.

42. Dendrotrypum pubipenne Blandf. (=Trypodendron pubipenne Blandf.) - the "Korean" timber beetle. Described by M. A. Blandford (Blandford, 1894) on Hokkaido under the name Trypodendron pubipenne. Later, K. E. Schedl (Schedl, 1951), on the basis of a number of morphological and biological characteristics, separated out a new genus, Dendrotrypum, from the genus Trypodendron, including in it several close Japanese species, in particular D. pubipenne Blandf. We have not found it on the islands; evidently it is very rare here. Cited by V. N. Stark (1952) for the Kuril Islands on the basis of the collections of N. N. Konakov without indication of a specific find site. K. E. Schedl notes the fig tree (Ficus caria), the leaf flower (Phyllanthus flexuosus), Machilis thunbergii, and Benzoin thunbergii as food plants of this bark-beetle. The food plants on the Kurils have not been elucidated. Known on Sakhalin, the Kuril Islands, Honshu, Hokkaido, Sikoku, Kyushu, Tsushima, Quelpart Island [Cheju Island - Trans.], and the Korean Peninsula. Judging by its distribution, we have assigned it to the group of Island (Kuril-Japanese) species.

43. Pityogenes chalcographus L. - the six-toothed spruce bark-beetle. K - env. of Sernovodsk set., July 7, 1962, 6 spec., from small branches of windthrown spruce; valley of the Lesnaya River, July 25-27, 1962, 32 spec., felling site in dark coniferous forest, from trunks and small branches of thrown spruce trees. Under Kuril Islands conditions the flight of the beetles is markedly protracted, from the end of June up until August. In connection with this, nests with larvae of various instars and mother galleries with just-deposited eggs are encountered simultaneously. Beetles of the new generation appear at the end of August-in September; the overwhelming majority overwinter in the breeding sites. P. chalcographus L. families are polygamous; typically three to five mother galleries take off from the nuptial chamber; they are clearly imprinted on the sapwood. Along with other bark-beetles they may inflict a certain amount of damage in logging regions. A Transpalaearctic species whose range encompasses all of Europe, all of Siberia (to the shores of the Pacific Ocean), North Mongolia, Primorskiy Kray, Kamchatka, Sakhalin, K, Japan (Hokkaido, Honshu), and the Korean Peninsula.

44. Pityogenes foveolatus Egg. - the "alpine tundra" bark-beetle. I - env. of Kuybyshevo set., August 22-23, 1961, 41 spec.; U - Okhotsk littoral near Cape Zhdi, the shores of Lake Tokotan, August 12-15, 1963, 39 spec.; P - env. of Severo-Kuril'sk, July 28, 1964, 3 spec. One of the principal pests of the Japanese stone pine on I, and the central and the northern islands of the Kuril Range. Attacks weakened and drying small trees. Occupies trunks and small branches along with Dryocoetes orientalis Kurenz. and Polygraphus sachalinensis Egg. We have at our disposal only fragmentary information regarding the developmental periods of this species under Kuril Islands conditions. At the end of August the development of the larvae is mainly completed; solitary older instar larvae, pupae, and young beetles are found in the nests; the latter evidently remain to overwinter under the bark, where they undergo supplementary feeding. A typical Okhotsk species, known from the high mountains of Sikhoté-Alin', in Magadanskaya Oblast, on Kamchatka (Kurentsov, Ivliev, 1969), Sakhalin, I, U, P, and from the mountains of Japan (Hokkaido, Honshu).

45. Pityogenes bidentatus Herbst. - the two-toothed pine wood engraver. I - env. of Kuril'sk, July 3-August 20, 1961, 1963, 321 species in all. Lives on the Kuril Dahurian larch. Specimens of the two-toothed pine wood engraver which we collected on I have a distinguishing feature: an additional pair of small sharp denticles is situated on the slope of the elytra of the males, above large hook-shaped denticles (Figure 14, 4). In the typical form these denticles are usually absent, or if they are present, then only in isolated individuals as an exception. In the Kuril population this character proves to be very stable, since additional denticles are present in all males and vary only somewhat in size. It is possible that P. bidentalis Herbst. living on the Kuril Islands forms a geographical race, all the more so since the body lengths in the Kuril specimens also are somewhat larger; they range from 2.5-3.5 mm, whereas typically (Stark, 1952) the length of the body does not exceed 2.8 mm.

Under islands conditions the flight of the beetles takes place at the end of June. At the beginning of July they gnaw through mother galleries and deposit eggs. The families are polygamous, i.e., three to four mother galleries typically take off from the nuptial chamber. By the beginning of August the development of the larvae is completed and the young beetles appear; they undergo supplementary feeding in the maternal nests for about a month; some of them remain in the breeding sites for overwintering. It multiplies en masse, forming local bark-beetle colonies, on weakened or windthrown Kuril larch trees. We have observed such a bark-beetle colony, for example, at the end of August, 1961 near Kuril'sk: the trunk and small branches of a thrown larch were solidly covered with nests of the two-toothed pine wood engraver. Thousands of young beetles which had turned the phloem and inner bark layers into rotten wood and had completely destroyed the initial mother and larval galleries were found under the bark. Some of the beetles had gnawed out flight openings and had emerged. Many beetles, running and flyng back and forth, could be seen on the bark. Characteristic for the zone of coniferous forests of Eurasia; in the Far East it is known in Primorskiy and Khabarovskiy Krays, on Sakhalin, I, and in Japan (Hokkaido).

46. Ips typographus f. japonicus Niiji. - the "oriental" spruce bark-beetle. K - env. of Sernovodsk set., region of the caldera of Golovnin Volcano, including from the stomach of the white-backed woodpecker, V. A. Nechaev; valley of the Lesnaya River, env. of Mendeleevo set., Sernovodsk set., the Pacific Ocean coast near Cape Petrov, outside collection dates July 7-November 8, 1962, 1964, 58 spec. in all; Sh - env. of Krabovoye and Malokuril'sk settlements, August 13-25, 1961, 1963, 31 spec. in all; I - env. of Lesozavodsk set., August 29, 1961, 8 spec. One of the common species in the dark coniferous forests of the Southern Kuril Islands; damages various spruce species. Multiplies en masse at the sites of wind-falls, windslashes, and logging, and forms bark-beetle colonies. The flight of the beetles extends from the middle of June to the middle of July. The formation of nests and oviposition take place in that time period, following which the beetles leave the nests they have established and undergo supplementary feeding until the maturation of new sexual products. At the end of July-beginning of August they apparently again begin to multiply and produce a second, sister generation of bark-beetles. Thus, on July 25, 1962, when larvae of various instars, pupae, and in individual cases not yet darkened, new generation beetles were found in the nests of the "oriental" spruce bark-beetle, the old beetles were gnawing out supplementary feeding galleries in fresh stumps. These galleries were irregular in form, comparatively short, and differed appreciable from the usual mother galleries (Figure 13, 9); there were no eggs in them.

By the end of August, young beetles are encountered en masse in families of the first round of reproduction. Some of these gnaw exit holes and emerge; the rest remain under the bark for the winter. Ant beetles (Thanasimus formicarius L.), rove beetles of the subfam. Aleocharinae, flies, and robber flies are encountered at the sites of settlements of the "oriental" spruce bark-beetle on the trunks of thrown trees in large numbers; the larvae of the above-mentioned predatory beetles and the parasitic wasp, Rhoptrocerus xylophagorum Ratz., are found in its galleries.

An oriental spruce bark-beetle, which comprises a geographical race of the Transpalaearctic species I. typographus L., distributed in regions with a moist marine climate which is characteristic for seashores and islands. It is known in the Tugurskiy Rayon of Nizhneamurskaya Oblast (Kurentsov, 1951), on Sakhalin, I, K, Sh, and in Japan - Hokkaido and Honshu (Kôno, Tamanuki, 1938). The reference of B. V. Sokanovskiy (1954) for the Aldan is highly doubtful.

47. Ips subelongatus Motsch. - the "greater larch" bark-beetle. I - env. of Kuril'sk, July 3, 1963, 8 spec., mixed forest, from the trunk of a sawn larch; August 18, 1961, 16 spec.; Listvennichnoye Plateau, July 11, 1963, 20 spec., from the trunks and small branches of drying larches; Kuybyshevo set., outside find dates July 3-August 24, 1961, 1963, 44 spec. in all, from larch logs. Characteristic for the larch forests of I, where it is a principal pest of the Kuril Dahurian larch. Attacking thrown trees and unbarked lumber, it forms dense colonies. It occupies the trunk and thick small branches. The mass emergence of the beetles takes place in the last ten-day period of August; they either remain until the spring in the maternal nests, or emerge and gnaw supplemental feeding galleries in other trees, sometimes gathering in one site in clusters of several (4-7) beetles. A Transpalaearctic species, widely distributed in the north of European USSR, in Siberia, including the Transbaikalia and Yakutiya, Mongolia, in the mountain forests of Khabarovskiy and Primorskiy Krays, in Magadanskaya Oblast, in the north of Kamchatskaya Oblast, the headwaters and middle region of the basin of the Penzhina River (Kurentsov, Kononov, 1961), on Sakhalin, I, and in Northern Japan (Hokkaido).

48. Orthotomicus suturalis Gyll. - the "burn" bark-beetle. K - env. of Lake Goryacheye, August 2, 1961, 1 spec.; Sh - env. of Malokuril'sk set., August 13, 1961, 1 spec. Very rare on the Kuril Islands. Lives on various spruce species. The phenological periods of development under island conditions have not been elucidated. Distributed in coniferous forests of Europe, Siberia, Mongolia, Khabarovskiy and Primorskiy Krays, on Sakhalin, K, Sh, and has been observed in Japan (Honshu).

49. Orthotomicus laricis F. - the "lesser larch" bark-beetle. K - env. of Lake Goryacheye, valley of the Lesnaya River, outside find dates July 16-August 2, 1961, 1962, 17 spec. in all. Dark coniferous forests of varied density and felling sites are this bark-beetle's habitats. Lives on spruce deadwood and unbarked lumber. The beetles begin to multiply apparently at the end of June-beginning of July, since nests with developed larvae are already found in the middle of July. The mother galleries are short and wide, take off from the nuptial chamber in one direction, and bluntly terminate in a small expansion, in which the eggs are deposited in a clump. The larvae gnaw a wide family gallery which forms several ramifications at the end. Before pupation, the larvae burrow into the bark layer and gnaw pupal cells in its superficial layers. The young beetles are also found in the outer layers of the bark during supplementary feeding. At the beginning of August some of them emerge from the maternal nests and continue to feed in stumps and fresher neighboring logs. The colonies of this bark-beetle are very dense, solidly covering the trunks of fallen trees. The development of O. laricis F. proceeds on dead trees, and its galleries do not affect the wood; therefore it does not have practical significance as a forest pest. A Transpalaearctic species with a range encompassing the territory of all of Europe, Siberia to the Pacific Ocean, Primorskiy Kray, Sakhalin, and K; it is observed on Kamchatka as an ecdemic; known in Japan (Honshu) and on the Korean Peninsula.

50. Xyleborus saxeseni Ratz. - the "polyphagous unpaired" bark-beetle. K - shore of Lake Goryacheye, env. of Alekhino set., July 17-August 2, 1962, 2 spec. in all; Sh - env. of Malokuril'sk set., August 28, 1963, 3 spec. Rare and scanty on the Kuril Islands. Inhabits drying thick small branches and trunks of alder and Manchurian ash. Adult beetles have been found in the middle of July and the beginning of August in galleries deep in the wood. It was not possible to track the phenological periods of development under island conditions due to the rare incidence of the species. Its galleries in small ash branches alternate with the galleries of X. germanus Blandf. The "polyphagous unpaired" bark-beetle develops in various woody species and is characterized by a circumboreal range. Distributed throughout Europe, including the Crimea and Caucasus, in Central Asia, Siberia, including the Transbaikalia and Southern Yakutiya, Primorskiy Kray, on Kamchatka, Sakhalin, K, Sh, in Iran, Mongolia, North China, Japan (Hokkaido, Honshu, Sikoku, Kyushu), and North America.

51. Xyleborus germanus Blandf. - the "Japanese unpaired" bark-beetle. K - env. of Alekhino set., July 3-August 2, 1962, 7 spec. Had not been listed previously for the fauna of the USSR. Its distribution on the Kuril Islands is apparently limited to the Okhotsk littoral of K, within areas where broadleaved species flourish in regions with the most favorable microclimatic conditions. Has been observed on the Manchurian ash and the silver magnolia. The beetles occupy drying small branches of varied thickness and thin small trunks on standing trees; they have not been observed on deadwood. The beetles burrow long, spiral-curved galleries in the wood and not infrequently reach the pith. Scolytoplatypus daimio Blandf. settles in close proximity to X. germanus Blandf. The nests of these two species are easily distinguished on the basis of the size of the entrances into the wood: for X. germanus Blandf. the diameter of the entrance is 1.0-1.2 mm, for Scolytoplatypus daimio Blandf. it is 1.6-2.0 mm. Adult beetles were observed at the beginning of August in the galleries. It has not been possible to establish the phenological periods of development in greater detail in connection with its rare incidence. An Island (Sakhalin-Kuril-Japanese) species, known on Sakhalin (?) (Kuwayama, 1967), K, in Japan (Hokkaido, Honshu, Sikoku, Kyushu), and on the Korean Peninsula.

52. Scolytoplatypus daimio Blandf. - the "Sakhalin xylophagous" bark-beetle. K - env. of Alekhino set., valley of the Lesnaya River, July 3-August 4, 1961, 70 spec. in all; Sh - env. of Malokuril'sk set. (valley of the Otradnyy Stream), August 27, 1963, 9 spec.; I - Okhotsk littoral in the region of Listvennichnoye Plateau, July 11, 1963, 2 spec. It is a serious pest of many broadleaved species in the dark coniferous-broadleaved forests of K. Attacks weakened and drying "controversial" Bothrocaryum, Sakhalin cherry, painted and "yellow" maples [Acer ukurunduense], "Maksimovich" and Manchurian alder, and Manchurian ash trees. On Sh it has been observed only on alder, and on I, on maple. Occupies the entire trunk from the base to the top, as well as thick small branches. Forms fairly dense colonies, frequently in combination with other bark-beetles: with Hyorrhynchus lewisi Blandf. on Bothrocaryum, with Xyleborus germanus Blandf. on ash, and with Xyleborus saxeseni Ratz. on alder.

The flight begins at the end of June-beginning of July and reaches maximal intensity in the middle of July, when a large number of beetles may be seen running along the trunks of trees and boring into the wood. The mother galleries are bored deeply into the wood; they are covered by a black fungal fur as is the case with the common timber beetles. Larval galleries and larvae themselves were not found in a single nest; the larvae probably confine themselves to feeding on the hyphae of fungi on the walls of the mother galleries. Adult beetles remain for a long time in the galleries and constantly eject white bore meal from them, by which means they evidently ensure ventilation of the nest. They stay close to the entrance in such a way that the slope of their elytra or the entire abdomen, protruding from the gallery, are visible. It is easy to distinguish "live" nests from old nests or those abandoned by the beetles by this feature. The parasitic wasps, Dinotiscus aponius Wlk. and Pteromalidae sp., have been extracted from the galleries of this bark-beetle; they evidently parasitize its larvae.

In terms of its origin, S. daimio Blandf. belongs to the Island (Sakhalin-Kuril-Japanese) species. Distributed on Sakhalin, K, Sh, I, in Japan (Hokkaido, Honshu, Sikoku), and on the Korean Peninsula, but the area of the damage it does is on the territory of K, Sh, and Japan; it is encountered rarely in the other sites and has no practical significance.

53. Scolytoplatypus tycon Blandf. - the "Ussurit xylophagous" bark-beetle. K - env. of Alekhino set., env. of Lake Lagunnoye, July 3-27, 1961, 1962, 14 spec. in all. Develops on many woody species within the limits of its range. On K it has been found only on three species: "Maksimovich" alder, "controversial" Bothrocaryum, and Sakhalin fir. It settles both on growing and on windthrown trees. Isolated adult beetles have been observed in the galleries during July. It was not possible to elucidate the developmental periods in greater detail. A Manchurian species, known in Primorskiy and Khabarovskiy Krays, on Sakhalin and K, and outside the USSR, in Japan (Hokkaido, Honshu, Sikoku, Kyushu), on the Korean Peninsula, and in North East China.