Exact reference to the published paper is: O.E. Kosterin, A.Yu. Haritonov, K.Inoue. 2001. Dragonflies of the part of Novosibirsk Province east of the Ob’ River. Sympetrum Hyogo, Vol. 7/8, pp. 24-49
(the journal of the HYOGO SOCIETY of ODONATOLOGY, Hyogo, Japan)
This paper is published as a bilingua, in Enlish and Japanese, the two text not coinciding with each other in full. Some information was doubled, some regional details were omitted in the Japanese text but some details interesting to Japanese readers were added. Besides, the paper is illustrated with 29 original photographs taken in Nature. Beow reproduced is only the English part of the text, without additional references connected to the only Japanese fragments and so far without figures. Any reference should be made to the paper in Sympetrum Hyogo.
Dragonflies of the part of Novosibirsk Province east of the Ob' River.
* Institute of Cytology & Genetics, Siberian Division of Russian Academy of Sciences, Acad. Lavrentiev ave 10, Novosibirsk 630090 Russia.
** Institute of Systematics & Ecology of Animals, Siberian Division of Russian Academy of Sciences, Frunze str. 11, Novosibirsk 630091 Russia.
In July 2001 the next XY Symposium of International Foundation S.I.O. is planned to be held in Novosibirsk, West Siberia, Russia. During the symposium, this Siberian city and its surroundings will be visited by many odonatologists, including those from Japan. Naturally, they would be interested in being introduced into odonatofauna of the latters, peovided that within the frames of the Symposium a number of odonatological excursions will be carried out. In this paper an annotated list of the dragonfly fauna is presented of the part of the Novosibirtsk Province situated east of the Ob’ River as the most interesting territory including the city, within 53o 28’ - 56 o 03’ N and 82 o 48’- 85 o 10’E, and comprising the Bolotninskii, Moshkovskii, Toguchinskii, Novosibirtskii, Iskitimskii, Cherepanovskii, Maslyaninskii, and Suzunskii Districts.
The major western part of the Province, west of theOb’ River, is situated on the great West Siberian Plain characterized by a very flat relief and only a zonal vegetation changing from taiga in the north to steppe in the south. The right bank part of the Province occupies a very gentle elevations outlined from the west by the Ob' River, the level of which at Novosibirsk is 92 m above sea level. These elevations comprise the NW extreme of the Altai-Sayan mountain country. They are composed with the Palaeozoic rocks and represented by the Herzin structures of the so-called Kolyvan'-Tomsk Priobskaya [Near-Ob'] Elevation, represented on the territory studied by the Sokur Plateau (north of the Inya River, up to 251 m above sea level) and the Bugotakskie Sopki hilly range (between the Inya and Ob' Rivers going roughly from the settlement Gornyi and village Zavyalovo, up to 381 m above sea level), and, eastwards from the formers, the Caledonian structures of a low and flat range of the Salairskii Kryazh (up to 502 m above sea level within NovosibirskProvince in its very east, in the sources of the Anfimov Mocheg Rivulet) (Chernov et al., 1988). The studied territoy resides mostly in the forest-steppe zone ("grove-belt") and its zonal vegetation is represented by birch and birch/asp groves (local 'kolki'), and also open water-shed birch forests, alternating with open meadow-steppe areas. In the south the patches of true steppes tend to play a significant role in the landscape, in the north (in Bolotninskii District) there appear birch/asp forests of the sub-taiga type with participation of spruce. Vast areas along the Ob' River, characterized by sandy soils, are occupied with pine forests (called, from south to north, Suzunskii, Karakanskii, El'tsovskii, Mochishchenskii Bors, and others). Smaller pine forests are scattered along smaller rivers (such as Inya, Berd'). With distance from rivers the pine becomes more scarce and is present as only an admixture to the birch forests and groves. The elevations also disturb the zonal disposition of vegetation. The southern slopes of major hills of the Bugotakskie Sopki are clad with true steppes while the northern slope and watershed surface of the Salairskii Kryazh are covered with asp forests of the subtaiga type and communities of the so-called chernevaya taiga (asp/fir forests), sometimes pure fir forests, in tree stands of crests and southern slopes also participates the pine and occurs the larch. Human activity lead to a substantial reduction of the area occupied by forests and groves while open areas are mostly ploughed up. Nevertheless, large areas of little disturbed natural communities persist even in the Novosibirsk close vicinities. The Ob', one of the great Siberin rivers, is partly turned into the Novosibirsk Water Reserve with scanty water vegetation and young banks still being undermined. The reserve is scarcely inhabited by dragonflies. Up- and downstream the sectionds of well-expressed floodland are retained with an intricate system of oxbow-lakes supporting flourishing populations of many dragonfly species. The Inya is rather a small lowland river with rather a weak current. The Berd' is draining the Salairskii Kryazh, twice crossing its axis and in these sections behaving as a fast mountain river. Fast and clear are also smaller rivulets of the Salairskii Kryazh, such as Chyom, Malyi Ik, Ik, Suenga etc. Within the Sokur and Bugotakskie Sopki small and middle-size rivulets are common flowing in a well developed valleys. Lakes are not common in the part of Novosibirsk Province east of the Ob' River, as different from the opposite part, theyare represented only by numerous oxbows of the Ob' and, to a less extent, Inya. But numerous are ponds on small rivulets.
It should be noted that, in spite of closeness to Novosibirsk, no regular studies of the dragonfly fauna of the territory considered were made. However, for many years a large body of data has been accumulated which allow to characterize this fauna quite completely. Most our attention was paid on the following sites: a) the Inya River left bank valley at the railway station Otgonka (Toguchinskii District); b) the Ob' River right bank vallet at villages Beloyarka, Uspenka and Dubrovino (Moshkovskii District); c) the same at village Suzun (Suzunskii District) (observations by A. H.); d) Novosibirsk surroundings, in particular, surrundings of Akademgorodok [Novosibirsk Academy Town] and of the village Nizhnyaya El'tsovka, the Inya River within few km of its mouth (Novosibirskii District) (observations by O. K.); e) since 1994, the Berd' River valley section between villages Legostaevo and Novososedovo (Iskitimskii Districrt). Weakly assessed remains Maslyaninskii District. In the below given annotated list of species information is given on distribution of dragonfly species over the territory considered and their biotopic preferences. Approximate flight periods are also outlined which, as a matter of fact, much vary anually (as it follows from the dates adduced, remarkable were an especially late spring of 1986 and 1987 and early spring of 1997).
Annotated list of species
Calopteryx splendens (Harris, 1782)
A rheophylic species common on all also the rivers including arms of the Ob' and to as small rivulets as the Shipunikha. As a rule imagines appear in the beginning of June but the time of appearing varies from May 28 (e.g. 1997) to June 14 (e.g. 1994 and 1996). Emerging of imagines is prolonged and may contunie up to July 20, while the flight period in general can last to the end of August.
Lestes dryas Kirby, 1890
An eurytopic and very common species. It prefers small water bodies and is very abundant on temporary ones and tussock swamps inundated in the forst half of summer, at the same time it is found on river banks as well as apart of water. Imagines fly from late June to middle September.
Lestes sponsa (Hansemann, 1823).
A mass euritopyc species which, however, avoids the smallest water bodies and is less frequently seen apart from water. The earliest imagines appears about June 10 and are observed in immense quantities in July-August.
Lestes virens Charpentier, 1825
A scarce species east of the Ob’ River, found on small water bodies, ponds. Some local populations sometimes are very abundant. The species has rather a late flight period as appearing in middle July and so becomes most noticeable in September when the two preceding species become less numerous. On 27th August 2000 on banks of a pond at Novosibirsk Academy Town O. K. observed many pairs laying eggs into dry stems of Alisma plantago aquatica L. about 0.5 m above the ground and within 0-3 m from the shoreline.
Lestes barbarus (Fabricius, 1798)
Rather a rare species east of the Ob’ River. A. H. found it seldom in the southern Cherepanovskii and Suzunskii District in ponds and small lakes on steppedied territories.
Sympecma paedisca (Brauer, 1877) ssp. braueri Bianchi, 1905
Common everywhere, permanently seen apart from water (up to 10 k). After hibernation these damseflies appear in early spring on forest edges and, in immense numbers, on the banks of all the stagnant water bodies. Their disappearance in July is not complete and some individuals can be observed throughout the warm season. The earliest recorded date of appearance is April 10, 1979, the latest date of observation is October 23, 1992.
Coenagrion armatum (Charpentier, 1840)
Prefers the smallest water bodies, little swamps and pools, as a rule covered with a carpet of Lemna minor, on large water bodies in the territory considered the relative abundence is extremely low. This is the earliest Coenagrionideae appearing in middle May (but emergence of imagines was also recorded as late as on June 24, 1987) and flying till middle July.
Coenagrion johanssoni Wallengren, 1894 (= C. concinnum (Johannson, 1859))
A rare species seldom met with in June-July in the valleys of the Izdrevaya, Inya, Verkhnii Suzun.
Coenagrion lunulatum (Charpentier, 1840) (=C. vernale (Hagen, 1839))
One of the most abundant damesflies, does not avoid the smallest water bodies and ponds. Flies from the last third of May to the end of July.
Coenagrion hastulatum (Charpentier, 1825)
Common on various stagnant and semicurrent water bodies, occurs also apart from water. Flies from the end of May to the beginning of August, but in 1987 tthe main appearance took place in 20s of June.
Coenagrion puella (Linnaeus, 1758)
Occurs quite freuently on diverse water bodies including river sections with slow current; flies from early or middle June to middle August.
Coenagrion pulchellum (Van Der Linden, 1823)
A very abundant damesfly inhabiting diverse watert bodies and flying from the end of May to middle August, often together with the previous species. Belyshev (1973) wrote that this species lies its eggs on the undersurface of flowting leaves of Potamogeton, Nymphaea, Nuphar etc. On 23th June 1996, on an Ob’ River oxbow just downstream of the dam, OK observed that oviposition of this species takesplace almost exclusively on floating leaves of Hydrocharis morsus-ranae L.: a female sits on a leaf edge and bends its abdomen under the leafwhile the male stands vertically on its abdomen on the female’s ‘neck’. Oviposition on a leaf is rather fast, within a minute, and a pair flies to another leaf.
Erythromma najas (Hansemann, 1832).
This species inhabits only suffeciently large, often semicurrent water bodies, common on oxbows and vays of river and Novosibirsk Water Reserve. Appears in early June (e.g. June 4, 1989, but in 1987 the most intense emerging took place on June 24), flies until early August (e.g. August 1, 1999). The damesflies keep to floating leaves of aquatic plants and rarely seen apart from water.
Nehalennia speciosa (Charpentier, 1840)
An extremely rare species on the territory considered. Scarce individuals were recorded by A. H. June 29, 1988 on a small swamp in the Nizhnii Suzun Rivulet valey (Suzunskii District). On 7th August 1999 a male was observed by E. A. Perepelov (pers. comm.) on a small pool on the Inya River floodland near its mouth, that is almost within the city of Novosibirsk.
Enallagma cyathigerum Charpentier, 1840 ssp. risi Schmidt, 1961
By the structure of the superior anal appendages in males our damesflies belong to the taxon risi Schmidt, 1961, which occupies arid and semiarid territories of Asia from the lowest Volga River basin through South Siberia and Central Asia to Mandzhuria. In the populations inhabiting the mountains of Middle Ural, Altai and SE Siberia exhibita great diversity in the structure of the upper anal appendages which vary between the types of risi and cyathigerum s. str. Damseflies of Europe, Anterior Asia and the taiga zone of Siberia and Far East to Korea, and also of North America, have the appendages of the cyathigerum type. This gives us a basis to consider the taxon risi as a subspecies of E. cyathigerum (Kosterin, 1999).
A common damesfly, especially on relatively large stagnant and current water bodies, the necessary condition is an open water surface and more or less open and solid banks. Tends to keep to water surface. As different from Erythromma najas, which flies relatively a little and keeps to areas with dense underwater vegetation, this species is very mobile and easy to fly and likes surface of free water, with scarce emerging stems used as perches. Flies from early June to middle August (recorded on August 21, 1999).
Ischnura elegans (Van Der Llinden, 1820)
These damesflies are rather abundant on the Inya River banks and also occur on smaller rivulets such as Koinikha (Iskitim District). They keep to thickets of rush emerging from water and fly from middle June to early September. [Note: the underlined text is absent from the version published in the jounral and is added here, O. K.]
Platycnemis pennipes (Pallas, 1771)
A rheophylic species common on the banks of larger rivers, such as the Berd’ and Inya, as well as rivulets intermediate in size, such as Shipunikha and Koinikha. In general appears in early June, but in some years (say, 1990) the beginnig of emerging can happen as late as in the end of this month. The flight period ends in middle August.
Aeshna mixta Latreille, 1805 (= Ae. coluberculus (Harris, 1782))
Not rarely occurs since middle July to middle September, in some years were met with until middle October (for instance, recorded by A. H. on the Inya River on October 16, 1992). Appearing of some imagines can be prolonged until late August (e.g., a teneral make was collected at a pond on the Zyryanka Rivulet, Novosibirsk Province, on August 23, 1987). This species is more frequently observed far from water.
Aeshna affinis Van Der Linden, 1825
In the southern part of the territoey considered this species is not rare from middle July to middle September. Thus, on August 5-6, 1996 it was quite abundant on the Ob’ right bank floodland at the village Verkhnii Suzun (Suzunskii District) where an intense crepuscular swarming trofical flight was observed. On August 8, 1998 on a large pomd on the Koinikha River at the settlement Linyovo (Iskitimskii District) rather numerous males were observed which examined riparian reed thickets with a slow flight interrupted with frequent stops in the air , they as well often sat on the reed. It was obviously a reproductive behaviour but no individual areas seemed to exist. This species occurs as well at Novosibirsk: in early August 1986 a swarming trophical flight of several individuals was observed above one of the streets of Alademgorodok; a male was collected at the Inya River mouth on July 25, 1992.
Aeshna crenata Hagen, 1856
This species is common on forest lakes and ponds, it appears in the beginning of July and flies until the end of September, but in the autumn it does not look abundant. At water the males are patrolling their well expressed individual areas by evenly flying to and back along a bank section on a height of 1-2 m. Besides, his species is often observed on forest glades and cuttings far from water.
Aeshna serrata Hagen, 1856
This species is rather frequently met with but mostly far from water. Morphologically it is closely related to the previous one but they have differences in their ecological preferences: Ae. crenata is mostly a forestspecies while Ae. serrata is a steppen species. In the forest/steppe zone these two species occur together but in the pine forests along the Ob’ River and more woody areas at the Salairskii Kryazh Ae.crenata predominates substantially while in the more southern mostly steppen regions in Iskitimskii and Cherepanovskii Districts Ae. serrata is more common.
Aeshna juncea (Linnaeus, 1758)
Not rare, appears in early July amd flies to the beginning of October (the latest record dating October 5, 1997). This species attracts attention on those water bodies where Ae. crenata is scarce which excludes this species (Belyshev, 1973) and also in the end of summer and in autumn when the abundance of Ae. crenata decreases more rapidly. It is common on the Ob’ valley oxbows asn found on small forest pools and ponds as well. As different from Ae. crenata, the males fly low above the water manoevring among emerging vegetation and frequently stopping in the air. No individual areas can be observed but two males, having met each other, rise high and try tochase each other out.
Aeshna subarctica Walker, 1908
Known by a single finding in the Bolotninskii District in the Lebyazhya River headwaters (B. F. Belyshev, personal communication).
Aeshna grandis (Linnaeus, 1758)
Common, prevails of other Aeshna species on small forest lakes and ponds, inhabits also rivers of intermediate size, such as Berd’ and Inya. Flies from the end of June to the end of September. Imagines have in general a diurnal activity but participate as well in a crepuscular trophical flying together with Ae. viridis.
Aeshna viridis Eversmann, 1835
This species on the larval stage is tightly connected with thickets of Stratiotes aloides, .which are very common in the Ob’ River oxbows where it is also very abundant. However thanks to the fact that this dragonfly in West Siberia has mostly a crepuscular activity (Kosterin, 1996a) it is hardly seen in the dayime. Yet it must be very numerous, as its crepuscular flying is quite noticeable in July even on those steets of Akademgorodok which are close to forest, notwithstanding they are situated 6 km from the closest breeding sites. Here in Akademgorodok, on 3rd August 2000 a female was observed flying in dusk among a number of individuals of Ae. grandis above the surface of a small pond on the Zyryanka Rivulet lacking Stratiotes. The species appears in late June (say, recorded on June 28, 1997) and flies to the end of September. On the Ob’ River right bank valley oxbows at the village Nizhnyaya El’tsovka on July 16, 1994 and August 3, 1998 in an overcast weather an intense oviposition by numerous females into Stratiotes rosettes was observed, while in the closest sunny days no female was seen.
Ophiogomphus cecilia (Fourcroy, 1785)
Numerous in the Bred’ River valley, breeds also in the Inya River and smaller rivers such as Koyon, Izdrevaya. In the Inya River valley these dragonflies are masked with much more numerous G. epophthalmus, in late August/early September this species still flies and becomes noticeable while G. epophthalmus mostly disappears. An intense emergence was observed at village Legostaevo on June 13, 1994. In the Bolotninskii District adjacent to the Tomsk Province finding is possible of the Siberian-Far-Eastern species Ophiogomphus obscurus Bartenef, 1930 (Haritonov, Borisov, 1990).
Nihonogomphus ruptus (Selys, 1857)
B. F. Belyshev (1973) supposed that the larva collected at Novosibirsk by S. G. Lepneva (1930) and identfied as Anisogomphus forcipatus belongs to this species (their larvae are indeed similar). Undoubtedly this species was recorded from Novosibirsk Province thrice:
by an exuvium collected by O. K. on June 13, 1994 at the village Legostaevo among numerous exuvia of the previous species and determined by Richard Seidenbusch, Sulzbarch-Rosenberg, Germany;
by a teneral male collected by O. G. Berezina on June 16, 1996 on the Berd’ River left bank at thye village Starososedovo (Iskitimskii Distict) among much more numerous teneral individuals of Gomphus vulgatissimus and less frequent G. epophthalmus, Ophiogomphus cecilia and Stylurus flavipes;
by a female collected by A. Perepelov (pers. comm.) on 18th June 2000 at a pond on the Zyryanka Rivulet in Akademgorodok.
Most probably these are the westernmost findings of this species, which is common more easterly, in Central and East Siberia, that is the western border of its range coincides with the Ob’ River. Already in the piedmont hills of Gornaya Shoria Mts. (S Kemerovo Province) this species becomes abundant (E. I. Malikova, personal communication; observations by O. Kosterin).
Stylurus flavipes (Charpentier, 1825)
Occurs in the valleys of the Ob’, Berd’, and Inya Rivers. On the Berd’ emergence of imagines was observed on June 15-16, 1996. On the other hand, a very teneral male and female were met with on a sandy beach of the Novosibirsk Water Reserve at Akademgorodok on August 3, 1987 at 20-21 o’clock. Thus, this is one of the few dragonflies able to breed along open banks of the Reserve, their development being delayed.
Gomphus vulgatissimus (Linnaeus, 1758)
For a long time this species was considered as absent from Siberia (Belyshev, 1973). There was a report by Prof. S. D. Lavroff (1927) for Omsk (considered by B. F. Belyshev as erroneous) where it was not found later (Kosterin, 1996). However, it turned out that it attain a great abundance in the valleys of the Berd’ and Ik Rivers on the terrotiry considered, where it prevails overwhelmingly over all other Gomphids. An intence emergence was observed by O. K. at the village Novososedovo and Starososedovo on the Berd’ and Ik Rivers on June 15-16, 1996, May 31, 1997 and June 7, 1998. In the latter case the dragonflies emerged in especially enormous quantities, so that each step along the bank frightened several teneral individual from bordering vegetation! The dragonfly hatch mostly on this vegetation and also just on the ground up to 1 m off the water, both at the river sections with slow and fast current. Besides, an intense emergence was observed by O. K. on June 5, 1994 also on a much smaller River Bugotak not far from its fall into the Inya River (Toguchinskii District); scarce individuals of this species were met with as well on the Koyon River of about the same size (Iskitimskii District). At the same time, according to observations by A. H., in the Inya River valley only scarce individuals of this species occurs, for instance, at ailway stations Otgonka and Izynskii (Toguchinskii Dostrict).
The males of this species occupy perches at river bank being bunches of rush and other reparian vegetation and chase out passing by Anisoptera, including such large ones as Macromia amphigena.
Thus, in Siberia the species is known only from Novosibirsk and, with a doubt, Omsk Provinces.
Gomphus epophthalmus Selys, 1872
This species, in general rare in Siberia, for some reason is the mass one at the Inya River throughout its section on the territory considered, where it predominates overwhelmingly over all other Gomphids. Its life history was studied in 1979 and 1980 at the Otgonka railway station by A. H. and I. N. Haritonova (Haritonov, 1981; Haritonov, Haritonov, 1982). The larvae develop in river sections with slow current and silty bottom (which prevail in the Inya). Emergence of imagines has, as a rule, a mass nature and takes place on average in10s of June (not far from Inya River mouth was observed by O. K. also on June 22, 1986 and June 7, 1994). The imagines hatch both on riparian grass vegetation and on solid ground (see photo). Just ahter harding the tenerals leave breeding places and disperse off the river up to 1 km to concentrate on forest glades and along roads. 4-6 days later the reproductive period starts which lasts to middle August. The males migrate back to the river where they occupy small individual areas from where they chase out all large dragonflies but females of the same species. Most of the time the males sit on branches of riparian bushes sometimes undertaking short flight around their alotments. Females staydispersed for a much greater territory as they continue active feeding. Their main pray are butterflies and also other dragonfles, up to the size of Sympetrum flaveolum. Individuals of G. epophthalmus wait for their prays a’sitting on their perches and get to the air only with a pray having approached closely. The females come to water only for a short time for mating and oviposition. They lay up to 120 eggs at once. Imagines are active on average between 8 and 20 o’clock.
Some individuals can be observed at the banks of the Novosibirsk Water Reserve, on forest edges at Akademgorodok etc. But there is no direct observations of ability of the species to breed in the Reserve; most probably those mentioned were migrants. On June 16, 1996, rather few teneral individuals were met with by O. K. also on the Berd’ River at the village Starososedovo where they were much less abundant than G. vulgatissimus.
Beyond the Novosibirsk Province G. epophthalmus is reliably known from a number of pints throughout the Kemerovo Province: from the same Inya River (at the village Orlovka) (Belyshev, 1962) and its tributary the Kos’ma River (Gagina, 1997); from the Zolotoi Kitat River (collected by O. K. on 30th June 2000); from the Novokuznetsk surroundings (Dronzikova, 1998), from village Mundybash surroundings (A. Perepelov, pers. comm.); and from the Stolby Nature Reserve at Krasnoyarsk, Krasnoyarsk Province (Belyshev, 1971). The taxon was described from «Irkutsk» (Selys Longshamps et MacLachlan, 1872), however there are doubts that the labes expressed an actual place of collection rather than the administrative centre of the province. In the same work, for instance, «Gomphus» maacki was described which is absent from the Irkutsk Province and appears only in E Transbaikalia (Kosterin, 1999) and eastwards. No recent reports of G. epophthalmus exist from East Siberia. In East Russia this species appears in the Amurskaya Provin ce (Malikova, 1997), while the report by Belyshev (1974) for Primorye in fact was based on a female of Aisagomphus melaenopsoides (Doi, 1943) (Malikova, 1995). Beyond Russia G. epophthalmus is known for NE China (?Belyshev, 1973) and Korea (Asahina, 1989a).
Macromia amphigena Selys, 1871 ssp. fraenata Martin, 1906 (= M. sibirica Djakonov, 1926)
Until recently this dragonfly was known in Siberia and Mongolia under the name Macromia sibirica Djakonov, 1926. S. Asahina (1964) has studied the type specimens of Macromia fraenata and shown that this is a continental subspecies of Macromia amphigena and that M. sibirica equals to it. Later this point of view was supported by E. I. Malikova (1995) who studied specimens of «M sibirica» from Novosibirsk to Primorye.
As well as Gomphus epophthalmus, this very rare in Siberia dragonfly is common in the region considered where it inhabits the Inya and Berd’ Rivers (Haritonov, Haritonova, 1989). It should be noted that Macromia sibirica was described just from the Inya River at Novosibirsk (Dyakonov, 1926).
During the flight period of this species, lasting from early or middle June to middle August an observer happened to be on the bank of one of the mentioned rivers would observe one or several individuals ranging along the bank just above the bank. The bank sections convenient for meeting of sexes and oviposition are patrolled by several males moving one after another according to the infinit ribbon principle (Haritonov, Haritonova, 1989). Mass emergence of this species can be observed on some sections of the Inya and Berd’ River in early or middle June. Such an emergence was observed by O. K. and O. Berezina on June 16, 1996 5 km NE of the village Starorososedovo. Imagines emerged on a floodland damp meadow on the Berd’ River left bank (see photo) where the river slightly bends right and has a stony bed and fast current as just having broken through the axis of the Salairskii Kryazh range and passed large rocky cliffs. On this meadow, about 200 m long, 48 exuviae were collected, sometimes several emerging dragonflies could be observed from the same point of view. The imagines emerged at 10-12 o’clock at an overcast weather within 5-15 m (not closer) from the water. The larvae had to pass the riparian margin of Scirpus lacustris and rather a dense grass and, in some places, Salix cinerea bushes. The dragonflies emerged on grasses or, more frequently, low juvenile willow bushes on the height of about 0.5 m. In other downstream sites of the bank, rather similar to this one, only rare exuviae vere seen. Upstreams the bank terrace come close to the water and the bank becomes higher, there we found some exuviae only 1-2 m apart from the water and 1-2 m above it. It can be supposed that the larva tends to climb up for a certain height above water level and if the bank is flat this aspiration makes the larva to move for a long distance. Perhaps, this trait evidences that the species was formed in a mountainous conditions, as in the mountains, where the rivers have an unstable diurnal level, it can be of an adaptive value. This interpretation is supported by an observation by O. K. and O. Berezina in the Gornaya Shoria Mts. in the Kemerovo Province, where 5 exuviae of M. fraenata were found on the Kondoma River right bank just downstream of the village Kuzedeevo on June 25, 1996. They were disposed on almost a vertical rock wall at about 1 m above its base. In their way from the river the larvae had to pass not less than 20 m through willow bushes, a bank covered with concrete, a railroad on a road-metal embankment, and another bush stripe!
On the Berd’ river an entire hatching process of a dragonfly was observed (see photos.). For about an hour the larva rested immovable on an ear of Dactylis glomerata before the hatching started. The hatching as such happened rather quickly but then the teneral imago for about 15 min stayed on its exuvium with the wings not elongating, only sometimes it moved with its legs. Then the teneral moved from the exuvuim to the stem and started wing elongation. The abdomen started to elongate after wing elongation finished.
Many teneral individuals became the preys of spiders.
Distribution of this species in Siberia much resembles that of Gomphus epophthalmus. Beyond the territory considered in the paper it is hitherto known from the Inya River valley in the Kemerovo Province (the village Orlovka (Belyshev, 1973), the Zolotoi Kitat River (observed by O.K. in 2000)), from North Altai (the Isha River at the Lebed’ River mouth and the Lebed’ River lower reaches (Belyshev, 1973)), from the Kondoma River in the Gornaya Shoria Mts. in the Kemerovo Province (Dronzikova, 1998; unpublished data by O. Kosterin and E. Malikova) and the Krasnoyarsk environs (Belyshev, 1971; 1973). Besides, the species was reported from the Bulgan-Gol River in the very south-western Mongolia, in a desert habitat of the Dzhungarian Gobi so strikingly differing from the Siberian ones (Peters, 1995). Further east this species is not known from Siberia; it appears again in Amurland (Malikova, 1997) and inhabits Primorye (Belyshev, 1973, Malikova 1995), Korea (Asahina, 1989b) and NE China (??Belyshev, 1973) .
Epitheca bimaculata (Charpentier, 1825)
This species is in general rare on the territory considered but sporadically some populations in the Ob’ River oxbows attain high abundance. It is not excluded that this species is able to breed in the Novosibirsk Waree Reserve, as a female was recorede in down on June 28, 1986 flying above its bluffy bank at Akademgorodok.
On June 13, 1994 a about noon on the left bank oxbow lake of the Berd’ River 2 km NWW of the village Legostaevo O. K. happened to observe and photograph (see photo) an intergeneric copulation of a male Cordulia aenea and female Epitheca bimaculata. The copulation sat on a dry willow branch about 1.5 m above the ground and several m from the water. Behaviour of the dragonflies seemed quite normal, their abdomina made slow pulsating movements usual for copulations. Copulating lasted for a time sufficient to make about a dozen of photographic shots, about 0.5-1 min, then the pair was frightened by the observer. It should be noted that B. F. Belyshev (1973) repoprted that he several times observed copulations of E. bimaculata with females of Libellula quadrimaculata.
Cordulia aenea (Linnaeus, 1758)
Quite common on various stagnant and weakly currrent water bodies, mostly forest ones. The species appears in the end of May (in 1997 was recorded as early as May 21), flies to early August. In late May/early June these dragonflies are abundant on forest edges and glades far from the water.
Somatochlora metallica (Van Der Linden, 1825)
Prefers current and semicurrent water bodies, common on semicurrent Ob’ River oxbows and on small forest rivulets and springs numerous in the area considered (thus, in the Zyryanka strean on May 31, 1987 quite numorous larvae were found). Appears in early June (the earliest records date to June 7-10, but on the Berd’ River at the village Starososedovo emergence was recorede on June 16, 1996), flies until middle August. The dragonflies as a rule range low over the water along banks or small stream courses.
Somatochlora flavomaculata (Van Der Linden, 1825)
Occurs much more rarely than the previous species, recorded on the Ob’ oxbows at the village Nizhnyaya El’tsovka neighbouring to a pine forest (July 31, 1999) and far from the water in Akademgorodok (July 6, 1993). B. F. Belyshev (1973) notes that in Siberia this species occurs exclusively on lakes situated within or at pine forests.
Somatochlora arctica (Zettlestedt, 1840)
A species very rare on the territory considered; recorded by A. H. on small swampy forest lakes in Moshkovskii and Bolotninskii Districts in late Jule/early August.
Leucorrhinia caudalis (Charpentier, 1840)
A rare species known from the territory considered by scarce findings by A. H. on the Inya River oxbows at the station Otgonka (Toguchinskii District).
Leucorrhinia rubicunda (Linnaeus, 1758)
One of the most common dragonflies preferring very small stagnant forest lakes and pools. Appears about May 20 and flies up to middle July.
Leucorrhinia pectoralis (Charpentier, 1825)
Occurs together with the previous species but is less abundant. Imhabits mostly small stagnant water bodies surrounded with arboreal vegetation. Appears in the end of May and flies to the middle July.
Leucorrhinia albifrons (Burmeister, 1839)
A scarce dragonfly. It was observed on semicurrent oxbows of the Ob’ River right floodland downstream of the Reserve. In a good weather these dragonflies keep to open water surface apart from the banks, resting on emerging vegetation slightly appearing from water, such as Scirpus lacustris L., or on floating leaves of Nymphaea tetragona Georgi (a habit rarely seen in Anisoptera). When the sun disappears for a long time, they moved to riparian trees where sat 2-6 m above the ground.
Leucorrhinia dubia (Van Der Linden, 1825)
A scarce species recorded on mortlaes of the Inya and Ob’.
Libellula quadrimaculata Linnaeus, 1758.
A aommon and numerous species inghabiting diverse stagnant water bodies, including very small. As a rule it appears around May 20 and flies up to late July (recorded on July 31, 1999). In some years in the end of May in Akademgorodok O. K. observed the following phenomenon: for a day or several days numerous dragonflies fly in the same direction 3-10 m above the ground. At the same time several dragnflies can be observed from the same point of view, but their movement does not live an impression of a coherent mass migration which are often exhibited by this species in the steppen zone (Belyshev, 1973): the dragonfly looked perfectly autonomous from each other and following the same direction independently from each other.
Sympetrum croceolum (Selys, 1883).
This miracable dragonfly, ranging in Japan, NE China, Korea, S Primorye (Needham, 1930; Belyshev, 1973; Asahina, 1990) and later found in the Amurskaya Province (Malikova, 1997) was unexpectedly discovered in West Siberia by O. K. in 1982 on Lake Manzherok in NE Altai Mts. (Kosterin, 1987a, b; 1996b) and by A. H. in 1987 on the very south of the eastern part of the Novosibirsk Province (this finding was only shortly mentioned in press in Belyshev et al., 1989). At last, on 25th August 2000 three specimens were unexpectedly observed and one captured by A. Yu’s student E. Shtrekker on a small lake on the Ob’ River right bank floodland WITHIN the city of Novosibirsk, at the Communal Bridge! No other records are hitherto known from Siberia.
Taking into account a great interest of the western range of this species just for a Japanese reader we give below a detailed information on both localities.
Lake Manzherok (Lake Doingol) is situated o the very NW edge of the Altai Mts. in the Katun’ River valley at 423 m above sea level W of a woody mountain Sinyukha (1218 m above sea level), 2.5 km of the Katun’ bank and 88 m above it. Its size is about 1100 x 400 m and area about 37.6 ha; the depth does not exceeds 3 m. It is an eutrophyc lake with a sapropel layer on the bottom more than 5 m thick. The bottom temperatures are 5-6o lower than the surface ones due to cold ground sources. The lake feeds by these surces and springs and had an outflux before it was crossed with a dam. The eastern, north eastern and southern banks are covered with a depressed peat-moss pine/birch forest, at present partly cut. The lake bears a flourishing populations of many aquatic plants. There is a floating swing mat up to 45 m wide developed along these banks, then there goes a carpet of Nymphaea candida and other floating vegetation 5-50 m wide changing with even more wide carpet of water chestnut Trapa pectinata rosettes. Only the central part of the lake has an open surface. The NW bank is sandy, hard and open. Remarkable is that the water chestnut growing in this lake (not present elsewhere in the Altai Mts.) is represented by a form which is regarded as either a species Trapa pectinata V. Vassil. endemic for this very lake or as ranging in NE China, Japan and the Russian Far East species Trapa manshurica Fler. (Tsvelev, 1995), at least it is very close to the latter. Thus, an isolate of the East Asian dragonfly has a parallel in an isolate of an East Asian plant.
On Lake Manzherok in 1982 teneral individuals started to emerge on August 5 and continue emerging until the end of observations on August 15. For 8 days of regular dragonfly accounts undertaken 2-3 times a day along the entire bank 15 tenerals were observed in total (Kosterin, 1987b). When the lake was visited again on August 20-21, 1994 (Kosterin, 1996b) both teneal and mature individuals were as well observed. The imagines hatched on emerging vegetation along the floating mat edge from where tenerals flied to riparian trees and bushes, often becoming preys of Aeshna crenata. , Mature dragonflies almost exclusively keeped to swamped banks and did not visited the hard bank. Ther activity took place mostly above water surface within 0-40 m off the bank, i. e. within the zone of the aquatic plant carpet with dominance of Nymphaea candida. They used as perches dead willow branches, tips of Scirpus lacustris sprouts, or floating leaves of Nymphaea candida - the latter trait was never seen in other our Anisoptera. The dragonflies either had a steady perch from which they flew no more than for 5 m (up to 20 m when frightened) or flew slong the banks often stopping in the air and making fast rushes to the sides. This species has a strong and impetous flight mostly 0.2-0.5 m above the water surface (Kosterin, 1987b)
In the Novosibirsk Province a small isolated population of S. crcceolum was discovered by A. H. on August 26-28, 1987 on a small floodland lake without an oficial name several km of village Meret’ in Suzunskii District, just at the border with the Altaiskii Krai Province. The lake is situated at the outer edge of the floodland at the Ob’ valley board, on a border of the Suzunskii Bor pine forest. It is roundish, about 200 m in diameter, shallow, densely grown up with water vegetation. The banks are mostly bogged and covered with dense bush thickets. The dragonflies were observed above the water surface, flying and resting on emerging vegetation, and along the bank and exhibited no trend to disperse from the water, as other species of the genus. The size of the population is hard to estimate, in imaginal phase it does not exceed several hundred individuals. A population within Novsibirsk is soon to be examined. Both populations in the Novosibirsk Province are situated on the Ob’ River floodland plains, that is in relatively unstable habitats.
The West Siberian specimens do not differ from the Primorian ones available to O. K. and A. H. for comparison, but the mature specimens have brown darkenings of the distal parts of the wings expressed into a different extent (Kosterin, 1987b), which were not recorded in the Primorian ones.
Sympetrum flaveolum (Linnaeus, 1758)
The most numerous and omnipresent Anisoptera in the territory considered. It often breeds in temporal water bodies and is very common for grassy tussock swamps inundated in spring and early summer. It appears in middle or late June, dependent on the year, and flies to the middle October. In the Ob’ floodland at the village Dubrovino (Moshkovskii District) rather numerous copulating individuals were recorded by A. haritonov on October 14, 1998.
Sympetrum pedemontanum (Mueller, 1766)
Hemireophilous species, in West Siberia inhabiting only semicurrent (such as the Ob’ oxbows with the water filtrating from the river, ponds on rivulets and bays of the Novosibirsk Water Reserve) and current (the Berd’ and Inya Rivers) water bodies. A common species which, however, is never very abundant. It a appears in middle July and flies to m\middle September.
Sympetrum danae (Sulzer, 1776) (= S. scoticum Donovan, 1811)
A numerous species developing in diverse stagnant water bodies, including inundating grassy swamps. It appears in early July and flies to middle October, in general this is the latest dragonfly in the region, as remaining very abundant in autumn.
Sympetrum vulgatum (Linnaeus, 1758)
A common and very numerous species developing in diverse water bodies but, as compared with two rpevious species, prefers larger and avoids temporal ones. It appears somewhat later than S. sanguineum and S. flaveolum, in middle July, and flies to early October. But on August 1-2, 1998 the emergence of this species in the Ob’ right floodland boggy oxbows at village Nizhnyaya El’tsovka was still intensive.
Sympetrum sanguineum (Muller, 1764).
A common species emerging in late June, substantially earlier than the previous species. Flies to middle September. It is more numerous in woody territories of the region considered.
Besides, on the territory considered findings of some other species known from the adjacent regions cannot be excluded. They are: Calopteryx japonica Selys, 1869 (occurs in the northern Altai Mts. (Malikova, 1995), common in the Gornaya Shoriya Mts. in the Kemerovo Province (data by O. K.)), Coenagrion ecornutum (Selys, 1872) (quite numerous somewhat more southly, in the Altaiskii Krai Province (Belyshev, 1973)), Ischnura pumilio (Charpentier, 1825) (known from two points in the North Altai Mts. (Belyshev, 1973; Kosterin, 1987a), Aeshna caerulea (Stroemberg, 1783) (a species inhabiting the taiga zone (Belyshev, 1973), Anax parthenope Selys, 1839 (found out in the Novokuznetsk environs and the Kondoma River valley in the Kemerovo Province (Dronzikova, 1998), Orthetrum cancellatum (Linnaeus, 1758) (common in the forest-steppe and steppe regions to the south, in the Altaiskii Krai Province (Belyshev, 1973), Sympetrum meridionale (Selys, 1841) (reported from the Kemerovo and Tomsk Provinces, by one locality each (Belyshev, 1973).
In 1981 B. F. Belyshev and A. Y. Haritonov (Belyshev, Haritonov, 1981) wrote that the Upper Ob’ Basin seems to be in odonatological aspect one of the best studied territories of the world. However it turned out that even on this territory striking findings are still possible, such as the discovery of Sympetrum croceolum. Besides, only recently in the Upper Ob’ Basin there was found Gomphus vulgatissimus, and in the Novosibirsk Province - Nihonogomphus ruptus.
Thus, for the territory considered 48 species are recorded. The fauna has in general a western appearance. 5 species (Lestes dryas, Aeshna subarctica, Ae. juncea, Sympetrum danae, Libellula quadrimaculata) have Holarctic ranges, 21 species (Lestes sponsa, Sympecma paedisca, Coenagrion armatum, C. johannsoni, C. lunulatum, Erythromma najas, Enallagma cyathigerum, Ishnura elegans, Nechalennia speciosa, Aeshna mixta, Ae. crenata, Ae. serrata, Ae. viridis, Stylurus flavipes, Epitheca bimaculata, Somatochlora metallica, Somatochlora arctica, Cordulia aenea, Sympetrum flaveolum, S. pedemontanum, S. vulgatum) have Transpalaearctic or nearly Transpalaearctoc ranges, 18 species (Calopteryx splendens, Lestes barbarus, L. virens, Coenagrion hastulatum, C. puella, C. pulchellum, Platycnemis pennipes, Aeshna affinis, Ae. grandis, Ophiogomphus cecilia, Gomphus vulgatissimus, Somatochlora flavomaculata, Leucorrhinia caudalis, L. pectoralis, L. albifrons, L. dubia, L. rubicunda, Sympetrum sanguineum) have West Palaearctic ranges and only 4 species (Nihonogomphus ruptus, Gomphus epophthalmus, Macromia amphigena, Sympetrum croceolum) have distinctly East Palaearctic ranges being distributed from the Pacific to the Ob’ River, i. e. find in the territory considrede the western borders of their ranges. Distribution of three species of the latter group is most interesting, namely, Gomphus epophthalmus, Macromia amphigena and Sympetrum croceolum. The two former are East-Asiatic species absent (or at least extremely local) from the vast territory of East Siberia (if not to take into account a very doubtful original report of G. epophthalmus for Irkutsk) but appear in the piedmonts of the western South Siberian mountains: in the environs of Krasnoyarsk, in North Altai, Gornaya Shoria and Salairskii Kryazh Mts. For S. croceolum only three localities are hitherto known in its western range: on NW Altai and the Novosibirsk Province. Such a disjunctive Altai-Far Eastern range type was recently isolated for some Lepidoptera connected with nemoral forest formations (Dubatolov, Zolotarenko, 1995; Dubatolov, Kosterin, 1998; 2000). This type is supposed to be exhibited by some species which acquired a continuous Siberian-Far Eastern distribution during the Climatic Optimum of the Holocene due to westward migrations from the Pacific regions (since the Optimum in the West started and finished later than in the East) but later retrived due to cooling and aridisation of the climate, leaving isolates in a sufficiently moist NW part of the Altai-Sayan Mountain System (Dubatolov, Kosterin, 1998; 2000).
An interesting and so far poorly understood phenomenon is the Gomphid populations on the rivers of the territory considered: dominance of an in general rare Gomphus epophthalmus over other species on the Inya River and prevailing of G. vulgatissimus (so far not found in other regions in Siberia) on all other rivers and rivulets. Either these two species compete with each other and which one would dominate on a river is rather a matter of chance or G. epophthalmus prefers larger rivers with slow current and silty bed while G. vulgatissimus prefers smaller and faster rivers with patches of stony bottom. Most probably, the eastern Novosibirsk Provinceis the only place where these two species are sympatric and contacting with each other.
An annotated list of the dragonfly fauna is presented of the part of the Novosibirtsk Province situated east of the Ob’ River as the most interesting territory including the city, within 53o 28’ - 56 o 03’ N and 82 o 48’- 85 o 10’E, and comprising the Bolotninskii, Moshkovskii, Toguchinskii, Novosibirtskii, Iskitimskii, Cherepanovskii, Maslyaninskii, and Suzunskii Districts. A short sketch of natural conditions of this region is presented. 48 species are recorded. Information is given on distribution of dragonfly species over the territory considered, their biotopic preferences and approximate flight periods. The fauna has in general a western appearance. 5 species (Lestes dryas, Aeshna subarctica, Ae. juncea, Sympetrum danae, Libellula quadrimaculata) have Holarctic ranges, 21 species (Lestes sponsa, Sympecma paedisca, Coenagrion armatum, C. johannsoni, C. lunulatum, Erythromma najas, Enallagma cyathigerum, Ishnura elegans, Nechalennia speciosa, Aeshna mixta, Ae. crenata, Ae. serrata, Ae. viridis, Stylurus flavipes, Epitheca bimaculata, Somatochlora metallica, Somatochlora arctica, Cordulia aenea, Sympetrum flaveolum, S. pedemontanum, S. vulgatum) have Transpalaearctic or nearly Transpalaearctoc ranges, 18 species (Calopteryx splendens, Lestes barbarus, L. virens, Coenagrion hastulatum, C. puella, C. pulchellum, Platycnemis pennipes, Aeshna affinis, Ae. grandis, Ophiogomphus cecilia, Gomphus vulgatissimus, Somatochlora flavomaculata, Leucorrhinia caudalis, L. pectoralis, L. albifrons, L. dubia, L. rubicunda, Sympetrum sanguineum) have West Palaearctic ranges, 4 species (Nihonogomphus ruptus, Gomphus epophthalmus, Macromia amphigena, Sympetrum croceolum) have East Palaearctic ranges being distributed from the Pacific to the Ob’ River and find in the territory considrede the western borders of their ranges. Distribution in Siberia of Gomphus epophthalmus, Macromia amphigena and Sympetrum croceolum is given in detail as most interesting. The two former are East-Asiatic species absent (or at least extremely local) from the vast territory of East Siberia (if not to take into account a very doubtful original report of G. epophthalmus for Irkutsk) but appear in the piedmonts of the western South Siberian mountains: in the environs of Krasnoyarsk, in North Altai, Kuznetskii Alatau, Gornaya Shoria and Salairskii Kryazh Mts. For S. croceolum only two localities are hitherto known in its western range: on NW Altai (Kosterin, 1987a, b, 1997b) and the SE extreme of the Novosibirsk Province (Belyshev et al., 1989) (detailed information on these two localities is given in this paper). Such a disjunctive Altai-Far Eastern range type was recently isolated for some Lepidoptera connected with nemoral forest formations. It is supposed to be exhibited by some species which acquired a continuous Siberian-Far Eastern distribution during the Climatic Optimum of the Holocene due to westward migrations but later retrived due to cooling and aridisation of the climate, leaving isolates in sufficiently moist NW part of the Altai-Sayan Mountain System (Dubatolov, Kosterin, 1998; 2000). Interesting is the fact that an in general rare species Gomphus epophthalmus predominates over other Gomphid species on the Inya River while G. vulgatissimus (so far not found in other regions in Siberia) prevails on all other rivers and rivulets. An intergeneric copulation was observed and photograhed between a male Cordulia aenea and a female Epitheca bimaculata.
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