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Journal of Wildlife and Biodiversity 4(2): 73-84 (2020) (http://jwb.araku.ac.ir/) DOI: 10.22120/jwb.2020.115601.1098 Scientific Report

Birds of High-mountain Daghestan, A report based on two decades survey

that include representatives of not only High- Evgeny Ev Vilkov mountain, Intra-mountain, and Piedmont Caspian Institute of Biological Resources, Daghestan but also of the plains of the republic. Daghestan Scientific Centre RAS, M. Gadzhiev However, the formation principle and St. 45, Makhachkala, composition of these communities still have to *Email: [email protected] be revealed. Received: 11 October 2019 / Revised: 26 March 2020 / Accepted: The purpose and objectives of the research were 26 March 2020 / Published online: 15 April 2020. Ministry of Sciences, Research, and Technology, Arak University, . to describe specific features of bird for each of the 11 bird communities forming the Abstract avifauna ecological structure of High-mountain The paper is based on 19 years (1998-2017) of Daghestan. the avifauna monitoring in High-mountain Daghestan. Each of 11 bird communities of the The paper summarizes the results of 19–year study area is characterized by its whole set of (1998–2017) ornithological counts carried out in , including the rarely occurred since 13 key sites, located within 6 administrative the latter promotes the formation of a unique districts of High-mountain Daghestan. Material pattern not only of an individual faunal and methods are fully represented in Report 1. community but also the entire avifauna of the To characterize ecologically different species of region. It is assumed that a specific avifauna birds, the author's classification was used pattern of High-mountain Daghestan is (Vilkov 2010b), distinguishing the birds by their determined not only by resident communities of occurrence in preferred habitats that allowed the typical mountain birds but also by nesting in the identification of 11 bird communities (Vilkov mountains adapted populations of migratory 2020, Fig. 2). The birds, found in the air above birds of the plains. The analysis of the collected the upper boundary of the vegetation (birds of material provides a picture of the species prey, airborne birds), were subdivided into bird diversity and territorial distribution of birds and communities of hoverers and airborne birds, helps to understand the avifauna resource while the air environment, being a feeding place capacities of High-mountain Daghestan. of airborne birds and a location from where the hoverers view the surroundings in search of their Keywords: Avifauna, bird community, victims, is conventionally treated as "habitat". In ecology, High-mountain Daghestan, case of revealing other, not so significant, micropopulaton regularities, the bird communities were segregated into additional groups of birds Introduction depending on their ecological specificity. The High-mountain Province of Daghestan has the most difficult habitat conditions, Findings substantially differing from Piedmont and Proceeding from the avifauna ecological Intra-mountain provinces by the sufficient level structure of High-mountain Daghestan (Vilkov of moisture and contrasting orography 2020, Fig. 2), the species richness of forest bird intensified by intermountain trenches. It is community ranks the first, uniting 45 species occupied by quite complex bird communities

74 | Journal of Wildlife and Biodiversity 4(2): 73-83 (2020) with a total average abundance of 246 ind./km2. searching for food, move in the forest interior The highest taxonomic density and ecological as gender pairs and groups. During the differentiation of the forest bird community are daytime, differently oriented snow-covered determined by the variety of habitats, high slopes of mountains and groups of shrubs and concentration of forage and diversity of trees between them are exposed to different shelters. In High-mountain Daghestan, being a intensity of light. The slopes located in the bioclimatic province with harsh conditions, concentration areas are warmed up at a forest stands still can be found along river different time of the day. As a consequence, valleys, on mountain slopes of northern birds select their routes taking into account the expositions and mainly in the north-western and hours when it is easier to get food in this or that central parts of the highlands, whereas the part of the forest. In the afternoon, when the south-eastern part is dominated by mountain- temperature reaches its peak, birds make up meadow landscapes. The forests are mosaically mixed flocks occupying the forest canopy with fragmented, making such transitional habitats the highest concentration of . As a result, as solitary trees, shrubs and small tree-shrub an observer being in the central part of a groves critically important for forest birds. Due mountain forest in the interval between 13 and to their territorial discontinuity, these 15 hours, experiences a minimal presence of transitional formations play a role of short-term birds, since the bulk of them is concentrated at shelters by which part of the forest birds moves its upper boundary. Moreover, the birds, between distant forest areas. These mosaically forming mixed flocks, dramatically shrink their scattered "forest” habitats, located between feeding area, shifting to a strategy of detailed large forest formations, turn a sporadically "combing" of the feeding site moving at high forested area into a conventionally continuous speed and with numerous repeats (probably due woodland massif. Besides, the diffuse to the interspecific competition). This itinerary distribution of forest birds, as a specific method of using the area allows birds to save adaptation, allows them to more effectively their time and energy expenditures when examine the entire forested territory and use exploring new feeding sites in the same area different types of forage and suitable shelters. and increases their feeding efficiency. As an According to the type of their daily movements optimal foraging strategy is formed in the and foraging the mountain forest birds can be course of natural selection and aimed at conventionally subdivided into five groups. increasing the consumption of net energy The first group consists of the species received during feeding (Cowie 1977), this characterized by the extensive range, high method of exploration of the feeding area by speed of movements and the relative this bird group can be attributed to the category permanence of their daily routes (Böhme & of specific adaptations. Banin 2001). It includes the Green Leaf The second group unites birds which diet is Warbler, Caucasian Chiffchaff, Coal Tit, Long- predominated by easily accessible vegetative tailed Tit (9.3 ind./km2), Great Tit (8.4 forage - fruits of the common sea buckthron ind./km2), Blue Tit (2.4 ind./km2), Eurasian Hippophae rhamnoides L., barberry Berberis Treecreeper (5.1 ind./km2) and some others, in vulgaris L., hawthorn Crataegus which diet a significant role is played by insects pseudoheterophylla Pojark, etc. It includes the and sporadically dispersed vegetation forage. (6.6 ind./km2), Ring Ouzel (8.4 The Green Leaf Warbler and Caucasian ind./km2), Common Blackbird (11.5 ind./km2) Chiffchaff (31 ind./km2 and 16.2 ind./km2) as and Mistle (6.4 ind./km2). Feeding of well as the Coal Tit (23 ind./km2) are absolute this bird group is characterized by a small range leaders in abundance. In the morning hours and of movements, lack of permanent routes and under uniform settled weather these species, more full consumption of revealed forage. In

75 | Journal of Wildlife and Biodiversity 4(2): 73-83 (2020) the afternoon they concentrate in areas with fidelity and aggression towards other high abundance of forage resources. When the individuals of the same species, trying to fruits are eaten to such extent that they are penetrate to their territory. found less frequently compared with other The fifth group is composed of diurnal birds of areas, the birds move to other sites (Böhme & prey and owls. It includes the Northern Banin 2001). These two groups cannot be Goshawk (1.3 ind./km2), Eurasian clearly separated since they have species with a Sparrowhawk (1.4 ind./km2), Long-legged transitional type of feeding. Depending on the Buzzard (1.6 ind./km2), Common Buzzard (1.5 environment the foraging strategy of such ind./km2), European Honey Buzzard (0.5 species (for example, Blue Tits) tends to the ind./km2), Saker Falcon (1.7 ind./km2), first or second group. Changes in the forage Eurasian Scops Owl (0.7 ind./km2), resources, depended on the productivity of trees Tengmalm’s Owl (1.2 ind./km2), Long-eared and shrubs, which are the main suppliers of Owl (1.3 ind./km2), and Eurasian Owl mass forage, have a crucial impact on the (0.4 ind./km2). A distinctive feature of this distribution of birds within their occupied area. group is an increase in the size of their feeding In this regard, none of the above-mentioned area in case of decreasing forage resources or species is a narrow foraging specialist that their availability (Galushin & Golodushko makes them more plastic and resistant to the 1963). They are also characterized by a high forage dynamics in mountains. pronounced fidelity to their nesting area and, in The third group includes birds that less particular, to forests where the same species intensively but quite widely move within (pairs) nest for many years. In fact, during long- extensive forest stands, making prolonged term counts of 2006-2010 in Tlyarosh Village stopovers in particular forest areas with the (Charodinsky District), a pair of Common highest concentration of forage. They are the Buzzards was regularly observed within the Green Woodpecker (3 ind./km2), Greater same coniferous-deciduous forest. A similar Spotted Woodpecker (2.2 ind./km2), Black territorial fidelity is observed for Northern Woodpecker (1 ind./km2), Tree Pipit (4.2 Goshawks, Eurasian Sparrowhawks and owls. ind./km2), Common Chaffinch (9.5 ind./km2), This feature of feathered raptors is traced not Common Bullfinch (4.8 ind./km2), etc. These only in the forests of High-mountain birds rarely between large forest stands. If Daghestan, but also in its Intra-mountain the ultimate goal of their flight is located at a Province (Vilkov 2010; 2013a), being considerable distance from their main habitat explained by the lack of their suitable habitats they travel (en route or by making short and, on the other hand, relative statics of stopovers) through mosaically scattered forest habitats of their prey. As for the present status groves, groups of shrubs and solitary trees. of the Caucasian Chiffchaff, as one of the three The fourth group consists of species rarely neoendemic Caucasian taxa, its abundance in leaving forest stands. It includes the Red- the High-mountain Daghestan ranges from 5 to breasted Flycatcher (3.4 ind./km2), Eurasian 54.2 ind./km2, with the total average number (8.7 ind./km2), (4.9 ind./km2), equalling to 16.8 ind./km2. Its range extends Common (5.7 ind./km2), and Eurasian from Piedmont to High-mountain Daghestan Robin (4.3 ind./km2). Sizes of their occupied (not found in lowlands), where it mosaically areas are insignificant compared with the three inhabits peripheral forest areas, forest edges above-mentioned bird groups, since these with shrub thickets and small groves species select them basing on an optimal set of surrounded by meadows with high conditions and preferred resources (including concentration of insects. This, in a grove nesting habitats). The specific features of their located at an altitude of 1850–1950 m, 5–7 km ecology are determined by a strong territorial southward of Urukh-Sota Village (Charodinsky

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District), the species abundance reached 54.2 mountainous regions of Daghestan the number ind./km2. The bulk of the Caucasian Chiffchaff of grazed livestock has been decreasing that, population is also concentrated in High- presumably, may negatively affect the local mountain Daghestan. The species status is faunal communities due to the excessive currently favorable, but there are certain threats overgrowing of meadows. It can be well associated with the reduction of forest areas, illustrated by an example though not from climate mesophilization and expansion of High-mountain but Intra-mountain Daghestan. anthropogenic landscapes. Thus, the Gunibsky experimental plot of the The second place in the species diversity is Mountain Botanical Garden (MBG) (1,900– occupied by a bird community of subalpine 2,000 m, 5–7 ha), free of any anthropogenic meadows, composed of 43 species with a total impact including livestock grazing, had the average number of 423 ind./km2. The specific grass height equalling to 70–80 cm with 100% features and originality of the subalpine projective cover. As a result, no birds were community living in severe meteorological revealed there during the counts, whereas the conditions of highlands have lead to the cultivated areas (3 ha) adjacent to the formation of a peculiar set of birds, which experimental plot supported the Red-backed species are recorded nowhere else except the Shrike (11 ind./km2), Common Whitethroat (5 mountains. Among the unique species it should ind./km2), Red-fronted Serin (8.3 ind./km2), be mentioned the Caucasian Black Grouse (2.6 European Goldfinch (10 ind./km2), Whinchat ind./km2), Chukar Partridge (3.5 ind./km2), (13.3 ind./km2), and Common Rosefinch (16.6 Common (2.4 ind./km2), Blue ind./km2). This suggests that the livestock Rock Thrush (2.1 ind./km2), Water Pipit (3.6 grazing has a regulating effect on the ind./km2), Shore Lark (3.5 ind./km2), Black composition of the birds nesting and feeding on Redstart (9.4 ind./km2) and some others. In pastures since the changes in grass cover alter addition to narrow specialists, there are also the not only a pattern of the nesting habitat but also species typical for plains: Common Quail (2.3 the structure of feeding habitats as well ind./km2), Eurasian Skylark, Common (Ostashchenko 2006; Lebedev et al. 2010). Whitethroat (5.4 ind./km2), Whinchat (10.3 Describing the current status of the Caucasian ind./km2), European Stonechat (10.5 ind./km2), Black Grouse as the second neoendemic species Northern Wheatear (8.9 ind./km2), etc. The of the Caucasus, we should note that its most abundant are the Eurasian Skylark (54.9 population density in the studied highlands ind./km2), Grey Partridge (44.7 ind./km2), and varies from 0.8 to 4.5 ind./km2, and a major part Red-fronted Serin (36.5 ind./km2). It is of it is also concentrated in the High-mountain noteworthy that typical mountain bird species Daghestan. The Caucasian Black Grouse in inhabiting the subalpine belt are characterized highlands mosaically inhabits shrub thickets by sedentary or, more precisely, sedentary- and subalpine meadows along the upper forest roaming lifestyles out of the breeding season, boundary. The current number of this species in whereas adapted populations of the species of the entire Daghestan is estimated as 1,100 pairs the plains, nesting in the same belt, are (Vilkov & Pishvanov 2000). At present, the migratory. The intensity of livestock grazing species status in the ecoregion is relatively that regulates the height and density of grass stable. The main limiting factors are severe and cover in subalpine meadows can provide a snowy winters, grazing, shepherd dogs and considerable effect on the qualitative and episodic shootings of birds. quantitative composition of meadow birds, The third place is occupied by a bird including their territorial distribution in the community of agro landscapes (fields, mountainous Daghestan and in High-mountain orchards, vegetable gardens). It is composed of Province as well. In the last 2–3 decades in the 41 species with a total average abundance of

77 | Journal of Wildlife and Biodiversity 4(2): 73-83 (2020)

431 ind./km2. The distinctive feature of these 2010). birds is mosaicity and local character of their The fourth place is occupied a community of distribution, determined by the boundaries of birds of cliffs and rock outcrops, uniting 31 agricultural landscapes. The most abundant are species with a total average abundance of 206 the Eurasian Skylark (50.3 ind./km2), Eurasian ind./km2. This type of landscapes is typical for Tree Sparrow (41.1 ind./km2) and Grey High-mountain Daghestan (mainly for its Partridge (39.9 ind./km2). The population north-western part). A high bird species density of individual species in agricultural diversity of these areas is because despite the landscapes reaches their peak values, since the harsh habitat conditions with a strict specificity area has both easily accessible forage for birds- of the species arrangement, these landscapes gatherers (Rock Dove (20.1 ind./km2), House support not only stenotopic taxa such as the (4.6 ind./km2) and Tree Sparrows, Red-fronted Water Pipit (3.6 ind./km2), Red-billed Chough, Serin (16.6 ind./km2), European Goldfinch (9.8 Blue Rock Thrush (3.3 ind./km2), Wall Creeper ind./km2), etc. and various shelters, represented (5.3 ind./km2), Chukar Partridge (10.1 by shrub thickets and high grass. Besides, these ind./km2), Güldenstädt’s Redstart (8.3 territories propose quite suitable nesting ind./km2) and some others, but also adapted habitats for birds of open landscapes thus populations of birds of the plains – the Common supporting annual breeding of the Grey (3.6 ind./km2), Northern Wheatear (8.9 Partridge, Common Quail (2.3 ind./km2), ind./km2), etc. The most abundant are the Rock Eurasian Skylark, Whinchat and European Bunting (24.4 ind./km2), Red-fronted Serin Stonechat (11 and 8.9 ind./km2), Red-backed (16.5 ind./km2), (11.8 Shrike (16.4 ind./km2), Common Rosefinch ind./km2) and Red-billed Chough (10.8 (10.2 ind./km2), etc. However, the structure of ind./km2). A characteristic feature of the breeding birds in agro landscapes is highly ecology of these birds is their sporadic unstable since the ecological niches presented distribution and static habitat borders, there can satisfy the needs of only a limited associated with the concentration of preferred group of birds. For the same reason the bulk of resources (nesting habitats, shelters, etc.) in birds only feed in agro landscapes, and for local areas of the mountains. Thus, in some shelter in bad weather and for breeding they fly regions of High-mountain Daghestan, there can to other, more suitable habitats. In fact, the last be found "nesting cliffs" with a negative slope 2–3 decades in the High-mountain Daghestan, (45–50 degrees) arranged as a niche, where, similarly to other mountain regions of the according to local residents, colonies of the republic, has shown a steady trend towards the Common House Martin, numbering up to 5–15 lesser exploitation of agricultural lands that, nesting pairs, have been nesting for many years. according to the information from local It is curious that such kind of colonies of the residents, has led to a dramatic decrease in the species, built in grotto-like covers, are number of Rock Doves, Grey Partridges, widespread at different altitudes and southern Common Quails and Eurasian Tree Sparrows. slopes of mountains of southern expositions At the same time, the remained agrolandscapes, from Piedmont to High-mountain Daghestan. additionally to breeding species, regularly Similar breeding habitats are also typical for the continue attracting the migratory birds using Eurasian (average total abundance agricultural lands as key ecological channels, – 7.5 ind./km2) but, unlike the Common House by which the migrants step-by-step move along Martin, it prefers breeding in separate pairs or the Transcaucasian Flyway. Regular migrants in a small colony of 3 to 7 pairs. include the Eurasian Bee-eater, Common In the fifth place in terms of the species Hoopoe, Eurasian Skylark, Red-backed Shrike, variation there is a community of synanthropic Whinchat and European Stonechat, etc. (Vilkov and conventionally synanthropic birds, uniting

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28 species with a total average abundance of Falcon, , Common Hoopoe, 232 ind./km2. Typical representatives of the Barn , Common House Martin, synanthropic group of birds include the Rock wagtails, wheatears, and Dove, Common Magpie (5.1 ind./km2), Hooded Eurasian Robin migrate to their wintering Crow (3.1 ind./km2), House (4.6 ind./km2) and grounds. However, part of the Eurasian Tree Sparrows. It is noteworthy that population (1.4 ind./km2) still remain in villages the Common Magpie, Hooded Crow and House for winter. During the winter and early spring Sparrow are not found in all settlements of such birds as the Red-billed Chough, Common High-mountain Daghestan, whereas the Raven and some others, usually not showing and the Rock Dove synanthropic, penetrate to the outskirts of inhabit almost all residential landscapes of the villages, where sufficient food and suitable highlands. All the above-mentioned shelters can be found. It should be emphasized synanthropic species have penetrated into that the number of the Hooded Crow in the mountainous areas since ancient times studied anthropogenic landscapes is extremely following human beings and since then always limited, and it is very important since its low prefer urbanized landscapes, where different numbers ensure the safety of a vulnerable types of anthropogenic forage and suitable community of mountain birds from this shelters can be found. Many rodents, available dangerous nest-destroyer. In the absence of there, also promote the breeding of the Little suitable nesting trees in some settlements, the Owl (2.1 ind./km2), which can be considered as Hooded Crow fly to nest in adjacent forests, a conventional synanthropic, because inhabits located as far as 500 and more meters away, practically all settlements of mountainous where it nests on tall trees along the forest Daghestan (Vilkov, 2007). The most abundant periphery. However, the nesting birds regularly species, in addition to typical synanthropic such return to feed in “their” settlement. as the Eurasian Tree Sparrow (52.2 ind./km2) The sixth place is occupied by a tree-shrub bird and Rock Dove (23.3 ind./km2), also include community, uniting 28 taxa with a total average conventional synanthropic such as the Rock abundance of 215 ind./km2. The most abundant Bunting (22.5 ind./km2) and Common species are the Rock Bunting (29 ind./km2), Blackbird (21.6 ind./km2), typical for High- Caucasian Chiffchaff (26 ind./km2) and Red- mountain Daghestan. The most typical fronted Serin (14.4 ind./km2). Such habitats as conventional synanthropic include the Eurasian shrubs and solitary trees in the north-western Jay (6.6 ind./km2), Great Tit (8.4 ind./km2), and central parts of High-mountain Province Black Redstart (9.4 ind./km2), Gray and White are usually used by non-breeding birds as Wagtails (7.7 and 8.1 ind./km2) and some temporary shelters, as we have discussed above. others. According to the original data and In conditions of the deforested south-eastern inquires, a major part of the synanthropic bird highlands dominated by mosaic shrub thickets population is sedentary or semi-sedentary as in subalpine meadows and sibljak-covered arid they occur in residential landscapes all year slopes with frequent winds and the deficit of round, including the autumn-winter season. The forage, shelters and nesting habitats, the role of only exception is their short-term departures to such refuges dramatically increases since birds feeding areas located on the outskirts of not only regularly feed and hide in them but villages. Most of the conventional also nest annually. The breeding group of these synanthropes, occurring in settlements, habitats consists of the Chukar Partridge (13.3 periodically move to adjacent territories in ind./km2), Common Blackbird (3 ind./km2), search of more suitable forage or breeding Red-backed Shrike (2.5 ind./km2), European habitats. In the autumn-winter season, such Goldfinch (14.3 ind./km2), Red-fronted Serin conventional synanthropic as the Hobby 6.7 ind./km2), Common Whitethroat (0.8

79 | Journal of Wildlife and Biodiversity 4(2): 73-83 (2020) ind./km2), Rock Bunting (36 ind./km2), etc. The were, "squeezing" him from their occupied species variation in breeding birds proves that territories. A similar reaction to the visitor was the harsh bioclimatic conditions of the High- demonstrated 2 hours later in the same site by mountain Province can support not only highly the same birds. The above suggests that, despite specialized mountain taxa, but also adapted the harsh conditions of the alpine zone, birds populations of ecologically plastic bird species gave a clear preference to the microhabitats that of the plains, which have formed stable long- were suitable for them, though located at term relationships with local habitats. To be untypical altitude. In fact, the main part of the more persuasive, we will illustrate this. Thus, local breeding population of the Eurasian Wren during the annual counts (2006–2010) in the (2.5 ind./km2) and Eurasian Robin (3.2 vicinity of Tlyarosh Village (Charodinsky ind./km2) are found in their traditional habitats District) (Vilkov 2013a), on the same bushed located 250–300 metres down the precipitous subalpine meadow (1,550–1,600 m) the same slope in a mature mixed-deciduous forest. male of the Red-Backed Shrike with a very The seventh place is occupied by the bird noticeable defect of his right wing was recorded community of alpine meadows and upper forest every year. The presence of this distinctive boundaries, uniting 17 species with a total character in a particular bird can be regarded as average abundance of 98 ind./km2. In High- equal to the reliable ringing data. Indeed, this Mountain Daghestan the alpine zone starts from bird from year to year bred in the same site an altitude of 2,500 m, and its fragments are within a radius of 100–150 m from the initial mosaically scattered. The severe point of its nesting record. In fact, the given meteorological conditions of the alpine zone, example suggests a strong long-term deficit of forage and suitable shelters determine relationship with the nesting site not only of a the presence of mainly stenotopic bird groups particular bird but also, probably, of the entire including Caucasian Snowcock (3.4 ind./km2), micropopulation adapted to the conditions of Caucasian Black Grouse (4.5 ind./km2), Alpine the mountains. And, as observations in other Chough (5.4 ind./km2), Red-billed Chough, regions of High-mountain and Intra-mountain Great Rosefinch (3.7 ind./km2), Water Pipit (3.6 Daghestan have shown (Vilkov 2010; 2011a, b; ind./km2), Snowfinch (6 ind./km2), 2013a), such habitat fidelity of Güldenstädt’s Redstart (8.3 ind./km2), micropopulations is recorded for many species Common Rock Thrush (2.4 ind./km2), Alpine of migratory birds of the plains preferring the Accentor (8.3 ind./km2), etc. The most same areas for many years. As for the fidelity to abundant are the Common Rosefinch (11.2 inverted habitats in the deforested south-eastern ind./km2), Red-billed Chough (10.8 ind./km2) part of High-mountain Province, there, in the and Black Redstart (9.4 ind./km2). Most absence of woodland areas, suitable breeding stenotopic taxa are disjunctively distributed, in habitats for some forest bird species are accordance with the location of alpine ridges. replaced by outcrops and piles of stones with They are also characterized by diurnal and mosaic shrub thickets. Thus, on 24 September seasonal movements to lower parts of the 2015 we encountered a Eurasian Wren and a mountains. However, despite the harsh Eurasian Robin on a juniper-covered rocky conditions of their habitats the alpine meadows slope surrounded by alpine meadows. The site are also visited by birds from other bird was located at an altitude of 1,950 m on the communities. Among them there are the right slope of the side gorge 6.5 km southward , Eurasian Skylark, Rock of Rutul Village (Rutulsky District). Both birds Bunting and some others. Observations show showed clear signs of aggressive territorial that these species are found within the alpine fidelity, manifested in loud cries of the birds belt in summer time or in the period of seasonal repeatedly approaching the visitor and, as it migrations. The species composition and

80 | Journal of Wildlife and Biodiversity 4(2): 73-83 (2020) population density of the alpine bird group are chicks from nests to sell them to falconers, relatively stable, though in recent years, due to photographers, etc.) also has substantially the growing hunting pressure, the numbers of increased in the recent decade (Vilkov 2014). In the Caucasian Black Grouse and Caucasian addition to the above-mentioned species the Snowcock might decrease. The Caucasian hoverers also include the Saker and small Snowcock, being the third neoendemic taxa, falcons – Merlin, Hobby Falcon and Common has a very sporadic range in the alpine belt of Kestrel. The Saker, as a nesting migrant, was Daghestan highlands, varying from 1 to 7.3 repeatedly recorded in different regions of ind./km2. A major part of its population is mountainous Daghestan. A particular case of its concentrated in High-mountain Province. breeding was recorded on 19 August 2003 at the During winter in highlands the Caucasian edge of the pine-birch forest in the vicinity of Snowcock usually prefers southern mountain the alpine village of Verkhnee Gakvari slopes, where snow patches is melted under the (Tsumadinsky District) (Vilkov 2014). A sun or quicker blown by wind. Seasonal vertical migrating Merlin (3.3 ind./km2) was twice movements are also observed: in the second recorded (20 July 2009 and 23 August 2017) in half of summer the species ascends to summits a subalpine meadow with inclusions of shrub and crests of mountains where it hides among thickets and large orchards in the vicinity of stones avoiding predators (Golden Eagle), and Tsurib Village (Charodinsky District). in winter, with the appearance of snow, it According to inquires, this falcon was descends to lower mountain belts. In winter the frequently recorded in this site. In contrast to all Snowcock prefers the lower half of the alpine feathered raptors, the Hobby Falcon in High- belt where it searches for snowless areas mountain Daghestan is directly associated with finding the forage; in early spring it descents up anthropogenic landscapes because it breeds in to the subalpine zone. The factors, limiting the nests of the Hooded Crows, and the latter, in species number, include cold winters with a lot turn, are typical synanthropes making their of snow and occasional shooting. nests in the human neighbourhood or at the The eight place is occupied with a bird periphery of nearby forests. As for the Common community of hoverers, uniting 13 species with Kestrel, it is quite widespread in different a total average abundance of 19 ind./km2. This habitats of High-mountain Daghestan breeding group includes large and middle-sized birds of in cliff niches surrounded by diverse landscapes prey – the Golden Eagle, Common Buzzard, where it hunts. In addition to typical hoverers Long-legged Buzzard, European Honey the discussed bird group also includes Buzzard, etc. and necrophages – the Griffon conventional hoverers, namely two species of Vulture, Lammergeier and Black Vulture. The hawks – the and highest density of hoverers are recorded in the . They are regarded as areas of rodents concentration in the subalpine “conventional” since they rarely leave forest zone, burial grounds for refuses, stands though sometimes move to subalpine landfills, and mature forests. Human activity and alpine meadows, less frequently - to the has an essential impact on the number dynamics nival belt and are often seen in settlements and distribution of necrophagous raptors in where they catch poultry on farmsteads. This Daghestan mountains. It can be negative or causes their periodical shooting by local positive, depending on the intensity of livestock residents (limiting factor). A quite significant grazing which number has noticeably reduced role of the Northern Goshawk as a key regulator (5–7 and more times) in the last 2–3 decades, of the number of some bird species should be partly due to the emigration of people from noted (Belik 2013). Thus, a pair of Northern mountains to plain regions. A direct persecution Goshawks, nested in 2017 and raised two of birds of prey by humans (hunting and taking chicks in the regularly surveyed forest of the

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Gunib Mountain Botanical Garden of Russian Daghestan, are located at different altitudes Academy of Sciences (Intra-mountain and, as a consequence, their inhabitants are Daghestan, 1,800– 1,900 m), dramatically represented by various hydrophilic birds with a reduced the number of average-sized birds, wide range of environmental preferences. which had been constantly seen in this site since These species have insignificant abundance and 2014–2016, namely the Eurasian Jay – from 10 low population density as the rapid water flow, to 2 ind./km2 and Common Blackbird – from rocky bottom and rocky-pebble banks make 53.2 to 11.7 ind./km2. Also, the Common Wood mountain watercourses unsuitable as a habitat Pigeon (2 ind./km2), Green Woodpecker (6 for many plain hydrophilic species. This has ind./km2), Common Buzzard (2.9 ind./km2) and lead to the formation of a small group of birds, Long-legged Buzzard (2 ind./km2) completely which distribution is directly associated with disappeared from the study area. Quite possible the mountain hydrological network. It is that having depleted their current forage composed of the species which populations are resources, the Northern Goshawks will further most adapted to the water bodies and streams in have to hunt on small birds, not only in the harsh mountain conditions thereby turning given forest but also in adjacent open them into typical mountain bird species. They landscapes that may decrease the number of are the Common Sandpiper, Gray Wagtail, some small bird species (Belik 2013). White Wagtail, and White-throated . The ninth place is occupied by a community of These species use sharp forms of coastal relief subnival-nival birds, consisting of 7 species for nesting, and watercourses – for feeding. It is with a total average abundance of 194 ind./km2. interesting that despite the mudflow of 2009 In High-mountain Daghestan the subnival-nival that had radically changed the outlines of banks zone starts from 3,000 m. For most bird species and pebble beaches of the Karala-Zurger River the habitat conditions of this zone are extreme, (near Tlyarosh Village, Charodinsky District, and in this context, similar to the community of 1,530 m), a micropopulation of the Common alpine birds, the taxonomic structure of this Sandpiper, regularly nested there before the avian group unites narrow specialists well mudflow, not only returned in subsequent years adapted to dwelling in snow-covered areas of to their former breeding area, but significantly mountains and glaciers. The species diversity increased from 8.5 to 33.3 ind./km2. In addition and ecology of subnival-nival birds have much to a stable breeding group of mountain in common with the alpine species but there are hydrophiles, distributed within a branched no such taxa as the Caucasian Black Grouse, hydrological network of the highlands, there is Wallcreeper and Common Rock Thrush. also found the Green Sandpiper – a scanty However, other stenotopic forms are always summering species, occurring occasionally present (except for the winter season): along the banks of rivers and shores of water Caucasian Snowcock (1–5 ind./km2), Red- bodies. Thus, two Green Sandpipers were billed Chough, Alpine Chough, Alpine recorded on 10 September, 2000 at the shore of Accentor, Güldenstädt's Redstart (2.2 an alpine lake (2,800 m) in the vicinity of ind./km2), Snowfinch (6 ind./km2), and Great Chirag (Agulsky District) where the birds were Rosefinch (3.7 ind./km2). The most abundant feeding on Gammarus sp., living in shallows of species are the Red-billed Chough (10 the water body. Another species of the group of ind./km2), Alpine Accentor (2.5–14 ind./km2), mountain hydrophiles is the Marsh Warbler, and Alpine Chough (5.4 ind./km2). inhabiting shores of small lakes and spills of The tenth place is occupied by waterbirds, mountain watercourses overgrown with consisting of 6 species with a total average macrophytes. This species is widely distributed abundance of 209 ind./km2. The water objects, from Piedmont to High-mountain Daghestan as intrazonal inclusions of High-mountain (Vilkov 2010; 2011b). Thus, a solitary

82 | Journal of Wildlife and Biodiversity 4(2): 73-83 (2020) individual of the Marsh Warbler was recorded noticeable decrease in the number of Barn on 22.08.2011 on a marshy alpine meadow at that has been observed at the an altitude of 2,300 m in the spurs of the Nukatl macropopulation level over the last 10–15 years Ridge (Charodinsky District). (Vilkov 2013b). The decline in abundance of And, finally, the 11th bird group is represented the alpine population of this species is by airborne birds, consisting of 6 species of presumably associated with an integrated effect open air environment with a total average of 3 limiting factors: 1 – the climate abundance of 213 ind./km2. Despite the single mesophilization of mountain ecosystems and (air) habitat, the population diversity of these reduction in the wetland areas used by the Barn birds varies in different parts of High-mountain Swallow in the nest-building period (clay Daghestan. This is due to the fact that their intake); 2 – the reduction in the number of qualitative and quantitative composition is livestock and farms where it previously nested; determined not only by the orography of 3 - the application of new building materials suitable rocky habitats located at different (brick instead of natural stone, plastic windows, altitudes, but also by the habitat availability of cermets, etc.), worsening the quality of terrestrial ecosystems. In this regard, breeding habitats (farms, houses, etc.). geographically distant natural complexes The specific pattern of the avifauna of High- supply birds with different amount of forage, mountain Daghestan is provided not only by suitable shelters or nesting areas. When resident communities of typical mountain birds, feeding, the airborne birds actively use forage but also by nesting in the mountains adapted resources of their occupied territory, where they populations of migratory birds of the plains. are constantly moving following the The relationships of birds with their mountain successively warming by the sun landscape habitats (including the species of the plains), surface since the increased temperature in the having established during a long historical "sunspot" epicentre substantially activates the period, lead to the appearance of highly insect activity. In addition, the flying insects specialized and, as a rule, territorially isolated form so-called forage "aeroplankton", which micropopulations supposedly inhabiting the components attract airborne birds. In different same habitats for many years. highland areas the structure and density of Conclusively, the avifauna of High-mountain aeroplankton prey species greatly vary, directly Daghestan is characterized by the cyclic reflecting in the quantitative structure of the changes in periods of seasonal migrations and airborne birds. Thus, the presence of the the long-term changes under the impact of a set Eurasian Bee-eater, a migrating and summering of limiting factors: global warming, reduction species of the Daghestan highlands, is primarily in cultivation areas, decreasing livestock determined by the presence of apiaries around population, increasing anthropogenic which these birds regularly concentrate and transformation of the environment, hunting feed, irritating beekeepers who regularly shoot pressure and birds of prey. them (limiting factor) (Vilkov 2016). For The analysis of the collected material provides example, only on 7 September 2017 the apiaries not only a picture of the species diversity and located near Tlyarosh Village (Charodinsky territorial distribution of birds but also gives a District) concentrated, after the rainy weather, representation of the avifauna abundance of the over 150–180 Eurasian Bee-eaters, causing High-mountain Daghestan. panic among local beekeepers as the birds pecked bees directly at bee-entrances of the hives, preventing them from flying out to References collect nectar. Also, a long-term trend of all Belik V.P. 2013. [Current changes in the mountain ecosystems in Daghestan is a avifauna of the North-Western Caucasus

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and their causes] Transactions of Menzbir Daghestan. In: Zalibekov Z.G. (ed.). Arid Ornithological Society (in the memory of ecosystems]. Vol. 17, Part 2 (47): 55–62. E.N. Kurochkin) 2: 208–230. (in Russian). KMK Press. (in Russian) Böhme R.L., Banin D.A. 2001. [Mountain Vilkov E.V. 2011b. [Structure and ecological avifauna of the southern Palearctic. diversity of birds in Intra-Mountain Ecological and geographical analysis]. Daghestan (by the example of Gotsatl Moscow University Press, Moscow, 256 p. Village). In: Birds of the Caucasus: history (in Russian). of study, life in urbanized environment. Cowie R.I. 1977. Optimal foraging in Great Tits Proceedings of the scientific conference]. (Parus major). Nature 286 (5616): 137–39. North-Caucasian State Technical Galushin V.M., Golodushko B.Z. 1963. University Press, Stavropol, pp. 25–34. (in [Variability and factors determining sizes of Russian) hunting areas of birds of prey. In: Vilkov E.V. 2013a. [History of study and Proceedings of the 5th Baltic Ornithological structural-territorial relationships of birds of Conference] Tartu, pp. 36–40. (in Russian) High-mountain Daghestan (by the example Lebedeva N.V., Ponomarev A.V., Savitsky of Charodinsky and Tlyaratinsky Districts). R.M., Arzanov Yu.G., Ilyina L.P. 2010. In: Birds of the Caucasus: history of study, [Terrestrial fauna as an indicator of grazing life in urbanized environment. Proceedings pressure] Bulletin of Southern Scientific of the scientific conference]. Alpha Print, Centre of Russian Academy of Sciences 6 Stavropol, pp. 25–52. (in Russian). (4): 84–95. (in Russian) Vilkov E.V. 2013b. Population trends in regular Ostashchenko A.N. 2006. [The effect of migrants as the basis for a prediction model decreasing anthropogenic pressure on birds for conservation of the birds of Eurasia. of Kyrgyzstan. In: Ornithological Studies in Russian Journal of Ecology 44 (2): 142– Northern Eurasia. Proceedings of the 12th 157. International Ornithological Conference of Vilkov E.V. 2014. [Retrospective analysis and Northern Eurasia] Stavropol State the current state of the Saker Falcon (Falco University Press, Stavropol, pp. 406–407. cherrug) in Daghestan]. Bulletin of St. (in Russian) Petersburg State University. Series Biology Vilkov E.V., Pishvanov Yu.V. 2000. [Rare and 4: 38 - 48. (in Russian). scanty bird species of Daghestan. In: Vilkov E.V. 2016. [Features of ecology of bee- Priklonsky S.G. and Zubakin V.A.(eds.). eaters (Merops apiaster, M. superciliosus) Rare, threatened and poor studied birds of in Daghestan as a region of intensive Russia]. RBCU Press, Moscow, pp. 13–29. migrations] South of Russia: Ecology, (in Russian) Development 11 (3): 90 – 105. DOI: Vilkov E.V. 2007. [Owls of Daghestan] Berkut 10.18470/1992-1098-2016-3-90-105 (in 16 (12): 78–86. (in Russian) Russian). Vilkov E.V. 2010. [Structure and ecological Vilkov, E. 2020. The structure, territorial diversity of birds of the High-mountain relationships, and ecology of birds in high- Daghestan] Bulletin of the Southern mountain Daghestan', Journal of Wildlife Scientific Centre of the Russian Academy and Biodiversity, 4(1): 8-17. of Sciences 6 (2): 52–59. (in Russian) Vilkov E.V. 2011a. [Materials on the winter bird population of Intra-Mountain