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УДК 598.288.1:57.063(0470.1)
DISTRIBUTION, TAXONOMY AND ECOLOGY OF CINCLUS CINCLUS (LINNAEUS,1758) IN THE EUROPEAN NORTH OF RUSSIA
A.A. ESTAFYEV
Institute of Biology, Komi Science Centre, Ural Branch, RAS, Syktyvkar, Russia
The work is devoted to a usual for the European North of Russia nesting and wintering water bird species, the dipper C. c. cinclus L. 1758. The anthropogenic load on water ecosystems in the region largely increases negatively affecting the population density of this bird. The paper reviews the population status of dipper since the XX century using the author's data (1965-2014) and literature materials. The number of species remains high in natural habitats. Log drifting was not organized at the nesting places of dipper and so its number and nesting area remained high and spacious, correspondingly
Keywords: dipper, distribution, taxonomy, ecology, the European North of Russia
А.А. ЕСТАФЬЕВ. РАСПРОСТРАНЕНИЕ, ТАКСОНИМИЯ И ЭКОЛОГИЯ CINCLUS CINCLUS (LINNAEUS, 1758) НА ЕВРОПЕЙСКОМ СЕВЕРЕ РОССИИ
Работа посвящена обычному для Европейского Севера России гнездящемуся и зимующему виду водно-болотных птиц, оляпке C. c. cinclus (L. 1758). Антропогенная нагрузка на водные экосистемы в регионе в значительной степени увеличивается, негативно влияя на популяционную плотность этой птицы. В работе рассматривается популяционный статус оляпки с XX столетия, с использованием данных автора (1965-2014) и литературных материалов. В естественных местах обитания число видов остается высоким. Поскольку не был организован сплав древесины в местах гнездования оляпки, ее численность оставалась высокой, а территория гнездования - протяженной.
Ключевые слова: оляпка, распространение, таксономия, экология, европейский Север России
Materials and methods
The data on distribution, biology, and ecology of dipper were collected by the author for the period of 1965-2014 in the Komi Republic, the Arkhangelsk region, Finland, and Sweden. Also the appropriate published literature data were used. We conducted field routes (about 65 thousand kilometers) to count population number and number of nesting pairs. Dipper's diet was studied by the stomach's contents and winter findings on ice at feeding grounds as young fish, larvae of dipterans, beetles etc.
Study results and discussion
The dipper Cinclus cinclus L. (Linnaeus, 1758) belongs to the Cinclidae family and the Cinclidae genus. Dippers form a species nesting and wintering in the European North of Russia. The study region is inhabited by its typical subspecies C. c. cinclus (L., 1758).
Distribution
The bird can be met in Northern Europe from Finland, Sweden, and the Kola Peninsular to latitude 69° North, in the European North of Russia to the So-lovetsky Islands and the Onega River. It also inhabits the Leningrad region, Karelia, all rivers at the mentio-
Dipper Cinclus cinclus cinclus (L.,1758). Adultus male, fresh (autumn).
ned regions up to the cities of Arkhangelsk, Mezen, and Pinega. Eastwards the bird builds nests in the upper river courses of the Timan Range, northwards - up to the Tsilma and Tobysh Rivers, the Izhma River and its left-bank tributaries, the Pechora River and its right-bank tributaries, along the Ural Mountains northwards to the Western Saledy Ridge. In the Volga-Kama region its distribution area stretches to middle parts of the Southern Urals. Southwards it occurs to the western coast of the Black Sea, northern pre-mountain areas of Caucasia [1-12].
Dipper birds migrate to large distances in winter and so appear in the river valleys of the Northern Dvina, the Luza, the Mezen', the Pechora, the Izhma, the Vychegda, the Vym', in the upper Sysola course. An example was collected in the Koigorodok region of the Komi Republic at an ice-hole of the Kuim River at latitude 60° North (now it is a collection example of the Institute of Biology, Komi Sci.Centre, Ural Branch, RAS).
Sizes. Taxonomy
Marked and complex; involves colour of head and nape and width of dark feather-fringes on remainder of upperparts (both strongly affected by bleaching and wear), colour of breast and belly (often with marked individual variation), and (slightly) size. 7 races in west Palearctic, but boundaries not clear-cut and
some variation within races.....Nominate cinclus from
northern Europe has upperparts similar to gularis, but breast and belly completely blackish-brown; some rufous brown tinge often visible along line with white chest, but not as much as in gularis. C. c. aguaticus from central Europe (south to north-west France, western Germany and Czech Republic) rather pale above and bright rufous-chestnut or deep chestnut on breast and belly; crown and nape rather pale grey-brown. C. c. caucasicus from Caucasus and Transcaucasia differs from other west Palearctic species in colour of up-perparts; dull grey-brown of crown and nape exstends to mantle and scapulars; breast and belly uniform dull grey-brown, rather like crown and nape [13].
The considerable variation within any single subspecies range merits caution in interpreting current taxonomi. The nominate C.c.cin-clus occupies the European North of Russia, the Urals, Fennoscandia, Denmark, C France, W Iberia, Corsica, Sardinia. It is the most migratory European subspecies, accounting for most wintering records outside the breeding range in the European North of Russia, Britain, France, Denmark, Finland and Baltic States. C.c.aq-uaticus occupies the remainder of the continental Europe except for the Caucasus and the Urals where caucasicus and uralensis occur. Lastly, there are two island endemics, biber-nicus in Ireland and gularis in Britain. Six, perhaps seven, subspecies occur where in the species distribution [14].
The research has shown that dippers nest in the Urals, Pre-Urals region, the Timan Ridge, in the basins of the Mezen, the Northern Dvina and further west in Finland and Sweden. Consequently, the breeding area of the C.c. cinclus in Europe is not interrupted from Scandinavia, the Kola Peninsula to the east to the mountain regions of the Urals, inclusive.
About 200 bird samples from collections of the RAS Zoological Museum, the Zoological Museum of
the Moscow University, the Institute of Biology of the Komi Science Centre, Ural Branch, RAS, the Pechora-Ilych Reserve, and the Helsinki University were analyzed by the author (Table 1).
The subspecies C.c. uralensis [15] was described by a male example with a highly light-colored head. Its habitat was reported to occupy the southern part of the Ural Mountains. The bird with a light-colored head was the only one exemplar of all birds collected in the Urals at the nesting time and had very old feathering. By Serebrovsky [15], C.c. uralensis birds in contrast to other subspecies have better expressed light-and brown (cinnamonic)-colored upper body part with their down part being more chocolate-brown than it is usually observed for C. c. cinclus. The C.c. caucasicus birds are highly variable whereby C.c. uralensis representatives are highly stable by color. Sudilovskaya [3] did not isolate the subspecies C.c. uralensis [15] and C.c. caucasicus [3]; she unified them under the subspecies C.c. cinclus L., 1758. Portenko [2] who studied 26 samples from Scandinavia, Finland, and the Kola Peninsula, 35 - from the Northern and Southern Urals, and 82 - from Caucasus confirmed the existence of C.c. uralensis. He characterized young C.c. uralensis birds at nests as being "stronger slate-colored with better expressed black edges".
Having studied 16 bird samples from the western slope of the sub-Polar Urals (the Bolshaya Synya River) and the Timan Range (the Kedva River, i.e. the Izhma River tributary) the author did not find any reliable differences by color or size of dippers between C.c. cinclus and C.c. uralensis. Dipper birds nest in the Urals, the sub-Urals, the Timan Range, in basins of
Comparative analysis of sizes of body parts (M±m and lim) and the coefficients of variation (CV) in different forms of common dipper (Cinclus Cinclus L.)
n Leng th, mm
wing CV tail CV hypotar-sus CV beak (from feathering) CV
M a l e s
Cinlus Cinclus Cinlus 46 90.1±1.12 81.5-96.2 4.77 54.8±85 51.8-61.0 5.58 27.7±0.18 27.3-27.4 2.28 15.5±0.30 14.3-17.0 6.91
C.c.uralen-sis 32 89.3±1.47 83.0-93.9 5.44 54.4±1.00 52.0-60.7 5.82 3 6 <° CO 02 +i и .6.0 22 10.00 15.5±1.82 13.5-10.0 5.72
C.c.cau-casicus 67 88.7±0.87 81.4-97.6 4.68 50.9±0.84 46.3-56.0 7.88 28.1±0.28 27.0-29.3 3.59 15.5±0.25 14.1-17.0 5.66
F e m a l e s
C.c.cinclus 28 86.3±1.33 82.0-93.0 4.08 52.2±1.46 48.3-58.3 7.39 26.6±0.26 26.5-27.3 2.55 15.0±0.60 14.4-15.4 10.00
C.c.uralen-sis 24 88.2±0.91 85.0-93.4 3.27 53.0±1.37 46.4-60.3 8.16 27.9±0.39 26.4-23.3 4.46 15.6±0.28 14.9-17.5 5.94
C.c.cau-casicus 31 83.3±1.12 78.6-93.2 4.46 49.5±0.79 46.0-56.3 5.31 27.6±0.19 27.3-28.5 2.15 14.9±0.18 14.1-16.0 3.35
else-
the Mezen' and the Northern Dvina rivers and westwards in Finland and Sweden. Consequently, distribution area of dippers does not stop by Scandinavia and the Kola Peninsula but stretches eastwards to the Ural Mountains. Southwards, C.c. uralensis dippers gradually become light-colored in head and belly, and reduce in size. These signs are better pronounced for C.c. caucasicus. Comparing body parts of subspecies by size (Table 1), we did not find any reliable differences between C.c. cinclus and C.c. uralensis. But they both do differ from C.c. caucasicus: males - by tail's length,
C.c. uralensis and C.c. caucasicus females - by wing's and tail's length. In all cases, C.c. caucasicus birds were smaller [6].
Among 21 nestlings obtained at different time of the nesting period, the author did not find any reliable difference between the subspecies. Only C.c. cinclus and C.c. caucasicus nestlings have a darker edge of back feathers. The author relates this divergence to the different age of collection material. The ecology of birds from the Timan Range, the Ural Mountains, and Karelia also does not much differ excluded for nesting time start [5-8].
The collection material confirms the subspecies C.c. cinclus and C.c. caucasicus met in the Southern Urals and produced hybrids. Collected C.c. uralensis birds have some examples with transitional signs which can be easily attributed to any of the studied subspecies [6]. By Portenko [2] "some C.c. caucasicus individuals collected at nesting time do not differ from C.c. uralensis by color, other C.c. caucasicus birds have the upper part of head and the fore-belly even darker than those of C.c. uralensis".
But these collection samples were normally old with worn feathers. Consequently, the whole area of the European North and the Ural Mountains is inhabited by Cinclus cinclus cinclus L. [6] which agrees with the data of Sudilovskaya [3].
Molting
Adult birds molt in the middle of July (females) and in the third decade of July (males) when they change small feathers. From early August till middle September they change contour feathers, quill feathers, and tail feathers. In the Pechora-Ilych Reserve, they found semi-fledged young birds in nests on June 19 and fledged ones on June 26 but with short quill and tail feathers [4,17]. Northwards in the upper Shchugor stream, nestlings were semi-fledged on July 11 and they left nests on July 27 with underdeveloped feathers [2]. Furthermore in the upper course of the Bolshaya Synya River, they found 4 semi-fledged nestlings. A young male from the upper course of the Bolshaya Synya River did not stop moulting towards October 6. Its head, neck, back, uropygium feathers and feathers on the lower part of belly were partly tubed (three quarters of the adult feathers length). The other feathers were new [7]. Young birds change their nestling dress for the first autumn dress from middle July till early October (small feathers, covering wing feathers; the other feathers are replaced in the next year).
Habitat
The species prefers taiga streams with clear water and does not inhabit mountain tundra regions. In the Ural Mountains and the Timan Range it occurs in rivers with rocky bottom with a stream velocity of 0.71.0 m/s. It was not met above the forest belt. In the Northern Urals it reaches areas with heights of up to 800 m a.s.l. Dipper birds make nests in niches of riverside cheer cliffs and in roots of fallen trees along river banks. The height of cheer cliffs with cracks and small niches covered by mosses and lichens sometimes reaches 10 m above river surface. These niches are
normally fed on ground waters. In pre-mountain and plain areas, the species inhabit sand-pebble tongues with outside fallen trees, stumps, and roots. River bottom on plain is normally sand with some rocks [7].
Number
In rivers of the Northern and sub-Polar Urals it is a usual nesting species. In rivers of the Timan Range it is seldom. Within 14 km along the upper course of the Bolshaya Synya River (sub-Polar Urals, at latitude 65° North) we found seven pairs in 1968, four pairs of birds each year for the period 1970-1992. Every 10 km approximately counted 3.6 pairs. The middle part of the Timan Range (a 45-km piece along the Chernaya Kedva River) was identified for three nesting pairs. In the Belaya Kedva River and in the upper course of the Mezen' River, this species is seldom [16].
In autumn and winter, dippers concentrate at rifts of non-frozen mountain streams and rivers. A 15-km-long piece in the upper course of the Bolshaya Synya River was identified for 14 dippers (October 13, 1968). They fed at light and twilight time, spent nights in roots of trees fallen near river. The northern slopes of the Sablya Ridge saw a decrease in number of wintering dippers (when comparing between 1968-1970 and 1992). A 14 km piece of the Voi-Vozh River mountain floodplain (the sub-Polar Urals) was identified for only eight birds. This resulted from a strong river overfreez-ing [7]. In the Pechora-Ilych Reserve it goes down to pre-mountain areas forming flocks of 20-30 individuals at non-frozen grounds in winter. At moderately cold winters it can be met on plain rivers [17].
Norway. 5000-25 000 pairs 1970-90. Sweden. 5000-50 000 pairs in late 1980s. Finland. 250-300 pairs in late 1980s (of which southern population 20-30 pairs). Estonia. Up to 5 pairs 1991. Latvia. 1-5 pairs. Russia. 1000-10 000 pairs [13].
Reproduction
The mating season starts with end April. In upper courses of the Pechora River and in the Timan tributaries of the Izhma River (the sub-Polar Urals) it starts in end May. From this very time birds form pairs and gradually, as rivers get free of ice, occupy a nesting area (usually the same as in the former year). The nesting area is 2-3 km long in the Urals and 5-6 km long in the Timan Range. The pairs build their nests for 1-2 weeks at a height of 1.5-2.5 m above water surface; they usually do not use the old one. The interior nest part is composed of moss and grass, is dry. The exterior part is composed of green moss and dry plant stems together with plant roots; it is moist as it touches soil or rock. The laying counts 4-5, sometimes 6 eggs. The brooding starts in end May and in the western slope of the sub-Polar Urals - in end June. In the Pechora-Ilych Reserve, there were 4 semi-fledged nestlings (49 g each) in a nest on June 19. The other nest included 5 nestlings with underdeveloped quill and tail feathers being ready to fly out of nest on June 26. In the upper course of the Shchugor River, there were half-dressed nestlings on July, 11. Three of them flew away on July, 27 [2]. The nest in the upper course of the Bolshaya Synya River included 4 semi-fledged nes-
tlings on July 6, 1972. In 1968, young birds flew away from nest on July, 25. In 1970, on a 14-km-piece in the upper stream of the Bolshaya Synya River we saw 4 nesting pairs of dippers. On July 16-20, each nest was left by 2-3 nestlings. Once left of nest, nestlings are still fed by adult birds. The number of nesting pairs at this river piece remained stable for 1968-1972. The number of birds in broods was 4 for 1968, 2 for 1969, 2-3 for 1970, 4-5 for 1972-1992 [7].
In the sub-Polar Urals along mountain streams, it seldom builds nests and occurs at a height of 800 m a.s.l. On July 13, 1968 we met a dipper bird with feed in the upper course of the Bolshaya Synya River. On July 25 we observed how adult birds fed their nestlings. They carried feed from stream every 3-4 minutes. In 1970, there were four nesting pairs within a 14-km-long piece along the Bolshaya Synya upper course. On July, 16 we recognized a family of 2 well flying nestlings. Female bird sometimes fed up both of them. Broods normally counted 2-3 nestlings. From mid July adult birds left nesting areas. In the upper Shchugor River, we found a nest on July, 11. It was built in tree roots and presented a rough sphere-shaped foundation. Three nestlings left on July, 27. In early July 1972, a nest built in a niche of an abrupt river bank at a height of 1.5 m hosted four semi-fledged nestlings [7]. Dmokhovsky [1] observed a brood of four young birds and one adult on August 19.
According to Teplova [4], dippers build nests in pre-mountain and mountain parts of the Pechora-Ilych Reserve and are abundant in upper tributaries of the Pechora River. In summer they live on cliffs. Nests were found on cliffs of the Bolshoi Shizhem, Shai-tanovka, Pechora Rivers (Verkhnie Klyuchi region). Normally they cannot be surveyed without special equipment as they are attached to an abrupt cliff above river. In 1944, they surveyed two nests. The first was in the Verkhnie Klyuchi region; it was organized in a niche of a cliff at a height of 1.5 m above river, made of moss and looked like a round mossy tussock. The nest included four semi-fledged nestlings on June 19. One nestling was taken away for investigation. It weighed 49 g; its stomach had only water or ground insects (larvae and imago of caddis-flies, saw-flies). The other nest in the upper course of the Bolshoi Shizhem River was surveyed on June 26. It was also made of moss and attached to a cliff 2.5 m above river. The nest held 5 big non-flying nestlings. They were smoky-grey-colored, body fully covered by feathers, but quill and tail feathers were very short; they jumped off and dove. In the upper course of the Bolshoi Shizhem River they met two hardly flying nestlings with an adult female on May 30, 1947.
In winter, dippers concentrate on unfrozen river parts, plain areas. Already in October they can be observed on rifts of the Pechora River and its Ural tributaries as the Unya, the Ilych, the Shchugor, Pod-cherem, Bolshaya Synya etc. [5-8,16].
Nutrition
Throughout a breeding range encompassing boreal, temperate, steppe and mediterranean climatic zones, the Dipper breeds alongside and feeds almost
exclusively in well-oxigenated waters of fast-flowing streams and rivers, especially where a stony bed provides preferred invertebrates prey such as mayfly (Ephemeroptera) adults and nymphs, caddis (Trichop-tera), larvae and stonefly (Plecoptera) nymphs. True fly (Diptera) larvae may replace mayflyes in the diet at high altitudes [14].
Birds enter water for feeding 15-20 times during an hour (usually without diving) in November. They stay under water surface 3-5 seconds with an interval of 3-4 minutes. Consequently, they were about a minute under water surface during an hour. Dippers started feeding at 9-10 a.m. and ended at twilight time. Birds passed nights on willow branches at river banks [7]. Without nestlings birds look for feed under river surface (larvae of caddis-flies, water coleopterans, other insects and mollusks).
Vladimirskaya [4] called dipper an enemy of young Atlantic salmon fish Salmo salar L. in the upper course of the Pechora River: "Small fish samples of minnow, bullhead, and taimen were found on ice at icehole taken from water by dippers in winter 1952-1953. From November 24 to December 8 we collected 13 taimen samples being 65-81 mm long. There were no smaller fish samples because possibly they were swallowed by birds. After December 8 we did not saw any fish samples on ice though dipper stayed here till spring.
Dipper obviously feeds on small fish in autumn-summer but seldom and does not insert any negative effect for fishery industry in the Pechora River basin. Having nestlings, birds usually feed when fly or take it from ground surface near streams. A dipper sample collected in February had fragments of water insects in stomach and in August - only ground insects as dip-terans, ants, and small beetles. Adult male weighs 71 g and young male - 65 g. Bird stomach contains gastric concretions: October 11 - 1.1 g, October 31 - 0.8 g, November 1 - 0.3 g, November 4 - 0.7 g. Stomach without gastric concretions weighed 0.1-0.5 g at day time.
Conclusion
The nesting area of Cinclus cinclus (L. 1758) in the European North of Russia is still safe of any anthropogenic load which provides conditions for successful reproduction and high population size. Stop of loose wood floating also has positive after-effects for nesting and feeding of dipper birds. The work is devoted to a usual nesting and wintering for the European North of Russia water bird species, the dipper C. c. cinclus L. 1758.
References
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