THE CAUCASIAN GROUSE LYRURUS MLOKOSIEWICZI TACZANOWSKI, 1875: HISTORY OF STUDIES AND MODERN PROBLEMS Potapov R.L., Pavlova E.A.

Zoological Institute, Russian Academy of Sciences, Universitetskaya Enb.,1 Saint-Petersburg, Russia. E-mail:[email protected] ABSTRACT The paper is devoted to the history of studies of a relic species and Caucasian endemic, the Caucasian Grouse Lyrurus mlokosiewiczi, starting from the appearance of the first zoologists in the Caucasus (1770-1773), discovery and description of the species, and its subsequent studies till the present time. The paper comprises the data on 128 works containing data on this species. Many of these publications are hardly available not only to foreign, but even to Russian specialists. In the conclusion, the authors determine the most important tasks in future studies of this species. They include clarification of the range borders, the detailed study of the biology of southern populations of the grouse, the further study of the structure of local populations and the degree of bird’s mobility with the use of modern technique (radio tracking, ringing), elaboration of methods for the cultivation of this species in capture in order to obtain hybrids between the Caucasian and the Black Grouses, for further expose of ancestral features, appeared usually in the hybrid offspring, and also for the study of the degree of stability of species characters, first of all, morphological and behavioral (voice) ones. And, at last, the further activation of paleontological investigations that can result in new findings of the fossils of this species, which now are very scarce. Key words: Caucasian Black Grouse, Lyrurus mlokosiewiczi, history of studies, bibliography.

INTRODUCTION At present, bibliographic, archive, and historic studies attract more and more attention because of a constantly extending time period separating us from the first original stages of certain investigations, appearance of new generations of scientists, disappearance of old scientific schools, unavailability of old literature, language problems, and some other reasons. Unavailability of original publications, in its turn, gives birth to new problems. On the one hand, they include problems of conscientious ornithologists, naturally obliged to know all the previous results obtained in their field of study. On the other hand, deliberate plagiarism becomes possible. The present communication is devoted to the history of the study of a relic tetraonid bird, the Caucasian grouse, one of the last birds discovered and described in the family 1

Tetraonidae in 1875. After the description of this species, only two new species of this Family were described, the Severtsov’s hazel grouse (1876} and Gunnison sage grouse (2001). According to these reasons, the Caucasian grouse remains poorly investigated even nowadays. The necessity of the present review is also explained by an interest of German ornithologists to the Caucasian grouse, developing since 1980-ies. According to philological problems or by some other reasons, they ignore all the works published by Russian, Georgian, Armenian, and Azerbaijan ornithologists. Even Russian ornithologists are poorly acquainted with the original studies. All this was a stimulating motive for the compiling of the present review. THE DISCOVERY OF THE SPECIES AND THE FIRST DATA The first naturalists that had visited the Caucasus as members of the famous expeditions organized by the Russian Academy of Sciences, namely, J.Guldenstadt who worked in upper Terek and Kuma Rivers in 1770, Ossetia, and Kabarda; P. S. Pallas who studied the nature of Kabarda in 1773; and E. Menetries who in 1829-1830 collected the material for St. Petersburg Kunstkammer in the Caucasus together with his assistant I. Voznesenskii, did not mention any grouses from Caucasian mountains. Caucasian Black Grouse was also not mentioned in famous Pallas’s voluminous review “Zoographia Rosso-Asiatica”. For the first time, grouses dwelling in the Caucasus were mentioned by A. Nordmann (1840). In 1834, he traveled along the Black Sea coast from Gelendzhik to the Turkish border and, supposedly, heard a grouse voice in the Gurian Mountains (the present Abkhazia). According to a fragment from his work, where the author points to the fact that the grouse (he talks about the black grouse) also dwells in the Gurian Mountains, because he heard its specific voice, which is usually produced during courtship. Nordmann did not see the bird directly, but in any case it was impossible to hear the loud song of the Black Grouse (Lyrurus tetrix) in those places and what sounds were heard by this scientist, is a great puzzle yet. Soon after Nordmann, the Austrian naturalist F. Kolenati traveled from upper Kuban River to Tbilisi in 1843. In August he was near Mt Kazbek, where the Caucasian grouse is frequently found even now, but did not find this bird. Following Nordmann’s example, many naturalists believed the Caucasus is populated by the black grouse, inhabited plains near foothills of Northern Caucasus. M. Bogdanov, the next ornithologist who visited the Caucasus in 18711872, searched for grouses in foothills, usual habitats of the black grouse, and did not examine the subalpine mountain belt (Bogdanov, 1879). He wrote about this afterwards, when the Caucasian grouse was described for the first time as a new species by Taczanowski (1875). The first ornithologist who managed to obtain the Caucasian grouse was the famous Gustav Radde, who studied the bird fauna of the Caucasus for 20 years. Radde appeared in Tiflis (the present Tbilisi) in 1863. Already in the next year, during his first voyage to mountains of Swanetia in 2

1864, he saw a grouse (he thought it was the Black grouse) in Mingrel Alps, at an altitude of about 2400 m. Later, in late June, he got a nestling and saved it in alcohol. Moreover, he mentioned that such grouses were found, although not frequently, everywhere in Mingrel mountains, and also near the source of the Tskhenis-Tskhali River. Already at that time he understood that this is a new species, different from Lyrurus tetrix; he even selected the name (Tetrao acatoptricus), underlying, in his opinion, the main difference between the new species and the black grouse, namely, the absence of the white mirror on wings. However, Radde did not hurry with publication of his results, continuing to collect the material. In 1884, when his review “The ornithological fauna of the Caucasus” was published, he possessed a collection of 8 specimens, including a skeleton (Radde, 1884). Some of these specimens were exhibited in the Caucasus Museum, created by Radde and opened in 1871. Later, this museum was visited by M. Bogdanov, who saw stuffed skins of grouses in a closed cabinet in a rather dark room; he did not, however, discerned these birds and their labels, because Gustav Radde was absent in Tiflis at that time. Later, we shall return to the contribution to the study of the Caucasian grouse made by this remarkable naturalist, Gustav Radde. The tireless investigator and collector, Radde was very slow in his publications: he published his main work on Caucasian birds only in 1884. In this work, he confesses that the description of a new species published by Taczanowski in 1875 came as a surprise for him. In 1874, L. Mlokosiewicz, a retired officer who worked as a forester in Lagodekhi in Central Caucasus? Not far from Tbilisi, shot two grouses in order to reveal possible differences between these birds and black grouses. This inquisitive observant was doubtful about the belonging of Caucasian birds to the species Lyrurus tetrix for a long time. In order to determine the birds more precisely, he sent them to L. Taczanowski, an well known ornithologist and the head of the Warsaw University museum. On the basis of these specimens, the latter immediately described a new species, naming it after his friend Tetrao mlokosiewiczi (at that time, and even now, many ornithologists included grouses and capercaillies into a single genus Tetrao). This description was very brief, as it is typical of the diagnosis. It mentioned that the Caucasian grouse strongly differs from the “European” grouse in the size, shorter wing proportions, and different shape, length, and width of the male’s retrices. The tail is forked less deeply owing to longer middle retrices. All the retrices are narrower; edges of ultimate feathers are slightly bent laterally, but are distinctly directed downwards. Then he gave a description of a female; unfortunately, this description was erroneous, because both Mlokosiewicz and Taczanowski took a one-year-old male for the female, and all the description of the female belongs to a young male. Lorentz (1884) was the first who noticed this mistake; he found that one-year-old males of the Caucasian grouse strongly differ from adult males in coloration and resemble females. In other words, the black color is entirely absent in these birds. Taczanowski 3

briefly described the coloration of the new species and main differences between this bird and the black grouse. He, however, did not describe young birds (the young male was described as the adult female), nestlings, eggs, the distribution of the species, etc. Nevertheless, the Caucasian grouse got his “civil rights” in taxonomy as the independent species, occupying the corresponding position in the structure of the family Tetraonidae and, which is especially important, attracted attention of zoologists (and ornithologists in particular). Just from this time, the study of the species began. Later, describing the history of the discovery of this species, Mlokosiewicz (1879) reports that he would get this bird and send it for determination long time ago, but all the specialists convinced him that this is a usual black grouse of no particular interest. More than that, he mentioned that he saw stuffed skins of this grouse in the “Museum” in Tiflis, labeled as Lyrurus tetrix. The “Museum” in Tiflis (Tbilisi) was the Caucasian Museum founded by Radde (at present, the State Museum of Georgia). Mlokosiewicz acquainted Dresser with his observations; the latter was preparing a voluminous review on European birds, published in 1876. Observations of Mlokosiewicz were published there virtually without any comments (Dresser, 1876). Dresser mentions Mlokosiewicz’s data on the finding of the grouse in southern slopes of the Main Caucasian Range in Georgia, in the Minor Caucasus in Armenian mountains, and in territories at present belonging to Turkey (near the Turkish border); on altitudes of their habitats; on some peculiarities of their behavior and feeding; and on the first nest with 10 eggs found. Mlokosiewicz, in particular, pointed that this grouse can be found at various circumstances and at different altitudes: he mentioned altitudes of 2610 m (8700 ft), 1896 m (6320 ft), and even 3300 m (in the latter case, “in a snowy land”). At the same time, Mlokosiewicz erroneously believed that this species also dwells in Mt Ararat and. probably, in Kurdistan. Already at that time, he wrote about the negative influence of cattle grazing on the population density of grouses, but published these data later (Mlokosiewicz, 1879). The description of the new species became known to all the naturalists only later and some of them continued writing about this bird as about the black grouse. Two communications more appeared in the same 1875. Shcherbakov (1875) found grouses in the southern Daghestan, in forests near Rugul Village (Samur district), mentioning them as black grouses (the Russian term “chernysh” used by him was commonly used in the game literature as the synonym of the black grouse). He also mentions that he saw “many grouses”, but reports nothing about their mode of life. In his communication, devoted to birds of Daghestan, Samsonov (1875) also mentions the Black grouse; from his text it is clear that he talks about the Caucasian black grouse. A year later, O. Markgraf (1876), one of enlightened hunters of that time, in his description of the Caucasian game described habitats of the Caucasian grouse for the first time. These habitats included a transitional zone between forests and alpine meadows. At the same time, he calls this grouse 4

“kosach” (Russian game term for the black grouse) again. In December 1869, Radde (1876) got two adult birds near the source of the Khrama River. He reports about male and female (at that time he did not distinguish young males from females), deposited at the Caucasian Museum created by him. He also points to the fact that grouses dwell southwards to Akhaltsykh “near the Turkish border.” In 1878 году, Transcaucasia (the source of the Kura River in the western edge of the Dzhavakhet Plateau) was visited by I. Mikhailovskii, in order to get a specimen of the new grouse species for the Zoological Museum in St. Petersburg. Although he did not manage to shot any bird, he assumed that this grouse “must be usual in mountain meadows, possessing a vast horizontal distribution in Caucasian mountains.” He was the first who reported about dwelling of the Caucasian grouse in this part of the Minor Caucasus. K. Kessler, the head of the Zoology Chair in St. Petersburg University, believed that the Caucasian grouse dwells only in the Caspian area of Transcaucasia (Kessler, 1878). His paper describes the journey of the author to Transcaucasia in 1875, when he observed no grouses in general. When he returned to St. Petersburg, he got information on the discovery of a new grouse species from Caucasian mountains and received two well-done stuffed skins (male and female) of the Caucasian grouse, sent by his friend N. Zeidlits from Tiflis. Surprisingly and fortunately, one of these exhibits, the male stuffed skin is still deposited at a small museum of the Chair of Vertebrate Zoology in St. Petersburg State University. M. Bogdanov (1879) was the first who summarized the results of the study of the new species. He analyzed the reasons of his own failures in search for the Caucasian grouse and also summarized all the known data on this species. In particular, he reported for the first time about wide distribution of the Caucasian grouse in Caucasian mountains “from Daghestan to the Black Sea and from the Main Range to Armenia. This bird dwells exclusively in the upper border of coniferous forests in birch groves and in the rhododendron belt” (Bogdanov, 1879). L. Mlokosiewicz (1879) gives mainly the data sent by him to Dresser earlier and published by the latter in English. In this publication, he excludes Mt Ararat, Elbur Mountain Range, and Lenkoran from the range of the species. The year 1884 was very rich in publications; each of them significantly widened our knowledge on this species. First of all, sketches on the Caucasian grouse were published by a naturalist N. Dinnik (1884, 1884a, 1884b, 1884c), who reported about his observations of this species. He found these birds quite common in northern slopes of the Main Caucasian Range, near the source of the Kuban River, and in slopes of Mt Oshten (the north-westernmost locality of the species). His communications also contain brief descriptions of habitats, meetings with broods, behavior of females with broods, and relationships with humans. T. Lorenz (1884) published his very important observations in the same year. He devoted two spring seasons (1883 and 1884) to the study of lekking of the Caucasian grouse, until that time completely unknown and even mysterious. He 5

founds leks of this species in Mt Bergamyt, 40 km south of Kislovodsk, on the northernmost border of the range, at comparatively low altitudes (1250 - 1500 m). Later, nobody saw leks of these birds at such low altitudes. As a result, he finally revealed that the absence of vocalization was the brightest feature in lekking of this species. In other words, the lek of the Caucasian grouse is silent. Just by this reason naturalists could not find leks of this species for a long time. Lorenz described, although rather briefly, characteristic poses and jumps and main difference in lekking behavior between the Caucasian and the black grouse. He was also the first who established strong differences in coloration of one-year-old and older males and pointed to a mistake in the original description of Taczanowski, which is especially important. Later (Lorenz, 1887), he repeated the description of lekking in German and gave a detailed description of the specimens obtained, pointing again to the mistake of Taczanowski. In the same work, he gave the comparative description of the syrinx of the Caucasian and the black grouse with schematic pictures in the natural dimension. These descriptions and pictures demonstrate the strong reduction of the vocal apparatus in the Caucasian species. Lorenz was also the first who attracted attention to the structure of grouse “brows” and its definite similarity to that in the Ptarmigans.The detailed description of the “brow” was given only a century later (Potapov, 1978). A review on birds of the Caucasus with a large section devoted to the Caucasian grouse was published in the same year by Radde (1884). There, the author summarized the results of all his long-term observations: he described male and female dimensions, coloration of the plumage (incorrectly), pointing to some age peculiarities. He also described general distribution of the Caucasian grouse and explained its absence in the Crimea, Mt Ararat, Elburs Mts, and mountains of Asia Minor and Afghanistan. Not arguing about the priority of Taczanowski in the description of the species, Radde explains the reasons of his tardiness in the description of the Caucasian grouse. At the same time, Radde did not notice Taczanowski’s mistake (the latter described a one-year-old male as a female), because he also paid no attention to strong differences in coloration of young and mature males. A map demonstrating the distribution of the Caucasian grouse with all collecting sites is the most important part of the publication. It is the first sketch of the species range; the distribution of some other bird species spread in the Caucasus is also shown. Interesting and first observations of the behavior of this species in Daghestan were published by K. Rossikov (1884), who visited upper flow of the Andiskii Koisu River in 1883. He was the first who described large aggregations of the Caucasian grouse formed in autumn (October-November) in feeding sites and described their food on the basis of the content of grouse crops. The role of young shoots and leaves of the shrub birch and aspen and cowberries in their ration was underlined. Two specimens of adult males from these places were granted by Rossikov to the Zoological Museum in St. Petersburg, where they are deposited now (collection 6

of the Zoological Institute, Russian Academy of Sciences, nos. 23932 and 23933). Together with Dinnik, he was the first who found the Caucasian grouse to the north of the watershed of the Main Caucasian Mountain Range and described the autumn life of the species. Apparently, he performed his journey in 1883, visiting the mountain Lake Retlo (Azen-Am) in Chechnya; he mentioned “a grove with grouses” above this lake. Rossikov visited this grove together with native hunters, but found no birds there; in his opinion, because of too many people and cattle. Later, he found grouses in mountains near Ingordakh Village and described snowy depressions with shrubs, small meadows, and barrows on slopes with numerous grouses that formed groups containing up to 6 birds, and also pairs of birds and solitary grouses that fed on leaved and buds of shrubs and grasses and with red cowberries. Females (he treated all gray birds as females) “fed and rested carefree”, and adult males frequently stretched their necks and looked around. After 2 gunshots, “flock after flock left their places; a low noise, something like ss-ss-rr-rr-rr, lasted for several seconds. It was striking that upper flocks as if notified in advance lower ones; the latter followed them immediately. After crossing of two thirds of the depression southwards, they changed the horizontal line of the flight altogether and, making a parabola, landed in a hundred meters from the starting place”. But not all the birds flew away, solitary individuals stayed between stones and shrubs. “In the daytime, grouses appeared in the same depression with the first sunrays. This time, they formed denser groups of 10 and more birds. Northern slopes were covered with dry and coarse grass and the eastern slope, with luxurious, although frostbitten green alpine grass; southern and western slopes were framed by sparse low shrubs of birch and aspen, and alder and heather with a brightly green curly carpet of cowberry. I found there the male grouse killed yesterday and dissected the crop; two thirds of the latter were filled with young shoots and leaves of birch and alder; the rest content was represented mainly by cowberries and a small amount of clover, buttercups, and flax. Noteworthy, that around me I could see bilberries and strawberries, low rhododendrons on rocks, and rare junipers. Lower part of the slope was covered with oaks, ashes, beeches, dog-roses, etc. Large number of limestone fragments scattered along slopes in grass and shrub thickets give birds numerous shelters, where they can spend nights, have rest, and build nests” (Rossikov, 1884, pp. 262 - 267). Together with Ingordakh, the Caucasian grouse was found in southern slopes of the Incharo Mountain Range, on its southern slopes and spurs running along the Zonokal Canyon. Two years later, Dinnik (1886) summarizes his observations on this species during numerous journeys to the Main Caucasian Mountain Range and uses new data published (Lorenz, 1884). This work contains no new data, except for the mentioning of the lowest altitude where the Caucasian Grouses were obtained (5500 ft, or 1650 m). In summer 1888, Rossikov (1890) examined the zoogeography of the northwestern Caucasus (the Kuban River basin); he reports, 7

that the Caucasian grouse was common in Gerpegem heights, in particular, near Storozhevaya Stanitsa, at very low altitudes of about only 1200 – 1300 m, in meadows among birch groves. He mentioned, however, that those birds were not breeding; nevertheless, the mentioned altitudes are the lowest where the Caucasian grouse was observed. Nobody found this bird at such low altitudes, although later, Averin and Nasimovich (1938) reported that “in western-northernmost localities of its range, it stays at altitudes of about 1000 all year round”. It should also be noted that Storozhevaya Station is situated on northern horns of the mountain massif reaching an altitude of 1800 m. Bilkevich (1893) found this species in all the examined mountains above the snow line, mainly in the western half of Daghestan, in sources of Andiiskii and Avarskii Koisu. His statement that males of the Caucasian grouse keep closely to females with the offspring was never confirmed by anyone, except for a single case (Orlovskii, 1905). In Daghestan (July 24), the author observed an offspring with a female and a male and noticed that young grouses were larger than a quail. Kratkii (1894), who participated in the game of Prince Sergei Mikhailovich in its game area (the present Caucasian State Nature Reserve) in September 1894 г., points to the abundance of Caucasian grouses near the source of the Urushten River, the upper left tributary of the Malaya Laba River in September 1-5 (Julian calendar). The author got 2 two-year-old males with still many-colored head and neck. The author described plumage of these birds and also assumed that the completely black plumage appears in males only in the third year of their life. It was a mistake, because the specimens examined by Kratkii were simply molted incompletely. Vilkonskii (1897) studied the fauna of the western part of the Minor Caucasus (Adzharia, Guria, Lazistan) under the supervision of M. Menzbier in 1891-1894. He revealed that the Caucasian grouse nests in small number over the entire area of mountain meadows of Adzharia and Guria at altitudes of 2100 – 2400 m, in low thickets of the rhododendron (it was evidently an erroneous observation). In June 8 1893, he got a female (near the apex of Mt Samiminomsta) with a large nesting spot. Its crop contained rhododendron buds, bilberry leaves, “blue flowers”, and a grasshopper. By words of natives, grouses are frequently found in winter in spruce forests near the Khanli Pass, Arsianskii Mountain range, 41º40́΄N. In 1895, a very informative and important (first of all, for ornithologists who did not know Russian) paper by Noska (1895) on the Caucasian grouse was published. Working as an administrator in the vast territory of the Emperor’s game reserve, coinciding in general with modern borders of the Caucasian State Nature Reserve, he published all the data known, with German punctuality mentioning all the details from publications known at that time. Nothing new was added to observations of preceding authors (especially, to the Lorenz’ data), but the data on the size, weight, and collecting dates of the large number of grouse specimens are still interesting. Vasilyev (1896) did not perform any field investigations of the Caucasian grouse. Some data on the vertical 8

distribution of this species are briefly mentioned in a section of his work devoted to the Caucasian grouse. In particular, it is mentioned that in Kuban area (the basin of the Kuban River), the Caucasian grouse is found starting from altitudes of about 5 000 ft (1500 m), and in the Black Sea area, i.e., in the southwestern side of the Main Caucasian Range, starting from 6000 ft (1800 m). It is interesting that Deryugin (1899) never mentioned the Caucasian grouse in his descriptions of the birds of Chorokh territory. STUDIES OF THE CAUCASIAN GROUSE IN THE FIRST HALF OF THE 20TH CENTURY Kalinovskii (1901) mentioned the high population density of grouses observed in AkhaltsikhImereti, Tatuzol, and Latpar Passes; in Dilizhan, Argun, and Tars canyons; and in the Dadian Svanetia (i.e., in regions where this species was found by Radde for the first time), and also in mountains around Borzhomi. It was the first report on successful attempts of domesticating of the Caucasian grouse. Captured birds rapidly get accustomed to new food (millet, buckwheat, rice) and to hens. Being really free, these birds frequently flew into the nearest forest, but always returned back. These striking data were never confirmed by anyone later. Dinnik (1902) reported about high population density of these birds in the upper Malaya Laba River: “In spring, they gather in leks as large groups”. Orlovskii (1905) described the distribution of the Caucasian grouse in Ossetia, where this bird was rather abundant (Lars, basins of Ardon and Urup Rivers, western slope of the Fiterta range, Chirkh, and Saturindzar) and also gave the data on lekking and the number of birds in leks (no more than 17 males; usually, 6-10 males), on dates of hatching, on their food and behavioral activity; it was a rather complete review of the ecology of the species. Observation of adult males near broods (very rare) attract special attention (this phenomenon was observed by Bilkevich earlier, see above), as well as the first description of the phenomenon of digging of these birds under the snow in winter. The author mentioned a case, when he “raised masses of birds” from under the snow, when he was going down from rocks above the border of the forest. K.A. Satunin, a well-known investigator of the Caucasian fauna, especially of the Minor Caucasus, did not study the Caucasian grouse especially, mentioning only solitary meetings with this bird during his excursions. All these meetings occurred in territories for which this grouse was already known (Satunin, 1904, 1907, 1911). In particular, he reported about comparative abundance of grouses in the Abkhazo-Imeretinskii Mountain Range. Many grouses, mainly young molting males, were collected there in 31.VII – 1.VIII in SaryChair-Kuga in subalpine meadows with thickets of juniper, rowan, birch, blueberry and some rhododendrons (Satunin, 1911). His report (from words of local hunters) on dwelling of the Caucasian grouse in the Okhochinskoe Mountain Range along the road to Nakhichevan 9

remained unverified. Satunin (1907) also mentioned separate places where these birds can be found: “I can add to what is already known on the distribution of these birds that they are found in mountains near Dilizhan, in Aratvinskii district near the Turkish border and in the Adzharskii Mountain Range.” In 1911, Satunin reported about the distribution of the Caucasian grouse in the alpine belt beginning from the border of the forest over the entire territory, excluding southern Transcaucasia «we got many grouses, mainly young molting males, 31.VII-1.VIII in Sair-Chary-Kurga in subalpine meadows with thickets of juniper, rowan, blueberry, and some rhododendrons.” Nesterov (1911), collector of birds from the southwestern Transcaucasia and adjacent regions of Turkey, worked in Transcaucasia simultaneously with Satunin. Both ornithologists exchanged with critical notes in scientific magazines, blaming each other for some erroneous statements. However, it was P.V. Nesterov who had an honor to discover the Caucasian grouse in the territory of modern Turkey, near the source of the Oltu River, in mountains of northeastern Turkey connecting with the Arsian Mountain Range (40º43’N – 44º34’EE, the former Russian-Turkish border, Barduz guard-post) in 20-ies of July 1910. At that tine, it was a solitary documented finding of the Caucasian grouse to the south of the Pontiiskii Mountain Range. Finally, in 1911, N.Ya. Dinnik found the Caucasian grouse in Zakatal district of the Main Caucasian Range, the easternmost point of the range at that time. The author does not mention this species from Daghestan. B.A. Dombrovskii (1913) analyses literary data on this species in Adzharia..E. Kudashov (1916) reported that “in the stomach of a nestling, collected in June 26 in Mt Anikho, I found grasshoppers, beetles, and flower buds of the Caucasian rhododendron”. The beginning of the First World War stopped publication of the data on the Caucasian grouse; new works appeared only in 1925. During this decade, the situation was unfavorable for ornithologists, but favorable for birds, owing to the decreased economical activity of humans in mountains; this phenomenon was reflected in new publications. Three of the latter were published in the Georgian magazine “Nadiroba”, at present hardly available. G. Mosienko (1925) mentioned the presence of the Caucasian grouse in Adzharia and described hunting for grouses with dogs near Goderskii Pass, which from Akhaltsikh was seen. “In the first day, about 40 birds were disturbed by dogs in the territory of half a square mile; we got 12 males, 3 females, and 2 youngsters.” Mlokosiewicz (1925) reported about attempts of keeping Caucasian grouses in captivity, about hatching of nestlings from eggs incubated by hens, and about feeding and behavior of these nestlings. A year later (Mlokosiewicz, 1926), he published another work on maintenance of the Caucasian grouse in captivity, in general repeating his work of 1925. E. Markov (1925) hunted for grouses in a watershed mountain range (altitude 7 000 ft) near Tbilisi, between Ksanskoe Canyon and the Aragva River valley. He noticed accumulations of birds in places with abundant cowberries, bilberries, blackberries, currant, and rowan. 10

Apparently, he was the first ornithologist who occasionally saw the autumn lek of grouses, but did not understand the scene. The author described as closer to the evening 7 males landed steeply on a clearing in a steep slope and after that set motionless in this clearing. The author did not understand what kind of a “club” it was and decided that they appeared in this clearing for watering, because a small brook was situated nearby. An unknown author under initials N.K. (1925) gave a list of food of the Caucasian grouse on the basis of the study of crops of two birds obtained in August near Bakuriani. P.V. Serebrovskii (1925) mentioned that this species was common in 1916 in Zakatalskii district of Transcaucasia, in places were later the Zakatalskii Nature Reserve was established. He gave the data on its population density obtained from local hunters; reported about flocks comprising 30-50 birds (in seasons when birds do not breed); and also on the unsuspecting behavior of these birds, that can be “killed by a stick” (probably, he talked about the period of lek activity). L.B. Beme (1925) published the results of his observations on 3 grouse broods performed during a month, describing the behavior of males and females, of young and old birds. He described places where broods feed and spend nights; noted that they were associated with certain territories and that their disintegrations began in the end of August. He mentions extreme altitudes of meetings of old males and single females in summer in alpine meadows near Devdorakskii glacier at an altitude of 2800 m, and on meeting with male flocks comprising 1520 birds in autumn near Kazbek. A new description of the nest (the first one was made by Mlokosiewicz in 1879) is given together with some data on molting. In particular, the author described the development of pale feathers on the body of adult males (especially on the neck and on the head) in the end of summer-beginning of autumn and mentioned that such plumage is characteristic of males in autumn. He clearly talked about the summer plumage of adult males, described later (Potapov, 1985) and similar to that in the black grouse. Mlokosiewicz (1926) gave a general description of the mode of life of the Caucasian grouse. He noted that usual habitats of these birds are situated at altitudes of 6000 – 8000 feet (1800 – 2400 m), less frequently, up to 9000 ft (2700 m). Winter habitats are situated at altitudes of about 6000 – 7000 ft (1800 – 2100 m); very rarely from 4 000 ft (1200 м). His author had also described the lek (by memory): «It stretches its tail like a fan; during jumps, occasionally produces a kind of a prolonged whistle; it is possible to approach this bird very closely; it is not afraid of gunshots; once, I shot 11 times into a single bird”. He realized that as many as 6 eggs can be found in a single nest and that birds nest in rhododendron thickets and, rarely, in thickets of birch and rowan, but only with an admixture of rhododendron. Birds fly up trees mainly in autumn and winter. Once, the author observed 10 frightened males flowing from under the snow; he had never heard about such mode of spending night. S.S. Turov (1928) noticed the negative influence 11

of pasturing on the size of the range of the Caucasian grouse; as an example, he mentioned the total disappearance of this species in Mts Fisht and Oshten. It should be noted however, that the Caucasian grouse never disappeared totally in these mountains and is rather usual there even now. I.D. Chkhikvishvili (1930) found that the Caucasian grouse was rather common species in subalpine and alpine meadows of southern slopes of the Main Range and reported that in winter this species inhabits the upper part of the forest zone, feeding there on birch buds. L.B. Krasovskii (1932) studied terrestrial vertebrates in southeastern Daghestan in 1929. Only three lines are devoted to the Caucasian grouse in this publication, where the existence of the grouse (”occasionally found”) in upper border of the forest, in particular, near Rutul and Gelmets Villages is confirmed (see Shcherbakov, 1875). In this paper, the author also gave the general characteristics of natural conditions of the region and mentioned that the upper part of the forest belt over the entire northern slope is represented by pure pine forests. During the following study of the fauna of Zakatalskii Reserve, Markov and Mlokosiewicz (1935) revealed the negative influence of the eagle-owl on the bird fauna; the Caucasian grouse was found among victims of the eagle-owl. The next stage in the study of the Caucasian grouse is associated with Yu. V. Averin, the future known Russian ornithologist and a student at that time, who started investigation of just this species in one of the most favorable places for this work: in the territory of the Caucasian State Reserve, established in the territory of the former “Royal Game Territory”. After 4-year field studies performed in 1933 – 1936, Averin (1938) published his first monograph on the ecology of the Caucasian grouse; this aspect of the grouse life stayed unexamined in spite of the numerous published notes and sketches. First of all, Averin studied in detail the northwestern part of the range and demonstrated that the range is represented by a series of small isolated islands. He was the first who underlined the phenomenon that these islands are associated with mountain apices above the upper forest border; in such cases, grouses dwell not in slopes, but only at the very apices of mountains. In such conditions, Averin found grouse habitats at altitudes of about 1000 m, i.e., lower than it was known. He demonstrated that vertical distribution of the grouse, fluctuating near the upper border of the forest, can strongly vary in dependence of the total height of mountains, being determined by the distance between the altitude of this border and apices of a certain mountain range or the border of the snow. In the mentioned review, he paid attention to the seasonality of the species and to seasonal changes in habitats for the first time. Averin also described for the first time in detail the life of the Caucasian grouse in winter. He showed that the winter mode of life of this species is quite typical of any other tetraonid bird, namely:

12

1. Winter feeding of the Caucasian grouse appeared to be similar to that of the black grouse, being based on the same birch buds and catkins and also on juniper needles and berries. Episodic feeding on fir’s needles in crowns of these trees was also noted for the first time. 2. It was found that birds regularly spend nights in under-snow chambers, choosing places with friable snow. Moreover, even the selection of winter dwellings depends mainly on the presence of the suitable snow cover. At the same time, strong snowfalls concealing low birch trees and juniper shrubs force birds to migrate to lower altitudes in the forest belt. The separate section of this work is devoted to feeding of the Caucasian grouse; in this section, its seasonal peculiarities were characterized and a list of plant and animal trophic objects consumed by adult birds and nestlings was given. The latter was obtained on the basis of the analysis of the content of bird stomachs. Sex ratio (1:1) in local populations was established for the first time and it was also demonstrated that rather low percentage of these birds live in flocks in this part of the range, by contrast to more southern populations. Flock formation reaches its maximum in autumn-winter, although the number of birds in each flock does not exceed 10. The author explained this phenomenon by the abundance of suitable habitats (including feeding conditions) and by the absence of other factors determining concentration of birds in limited territories. Averin also firstly described lek and breeding of the Caucasian grouse and clarified the description of lek made by Lorenz, pointing that birds make a 180° turn during a jump. He found that the specific noise, produced by grouses and believed to be produced by vocal organs (Lorenz, 1884), in reality is produced by wings of a flying-up bird. Among vocal sounds, he described only rather rare clicks and an original rattle resembling the voice of the corncrake. The author also described firstly male’s nuptial fights, their poses and behavior during such confrontations, and the daily rhythm of lek, with maximal activity in nearly complete darkness of dusk and down, distribution and types of leks; he also revealed the territorial character of behavior of males, possessing their own constant plots in the lek. The data are also given on nesting (9 nests found), the largest number of eggs in an egg-laying (9), duration of brooding (24 - 25 days), and the size of eggs. In the beginning of 1920-es, the significant decrease in the population density of the Caucasian grouse near the northwestern border of its range was noted together with the complete disappearance of this species in Chernye Mts, the northern advanced post of the Main Caucasian Range (in this region, grouses were common in summer in Gerpegem Heights near Storozhevaya Stanitsa already 30 years ago, Rossikov, 1890). This is the first evidence of the decrease of the range in the 20th century. In the end of the publication, the species is estimated as an archaic life form adopted to the Caucasian highlands; the further prospects of its existence are estimated as unfavorable because of the distinctly negative reaction of the species to human agricultural activity (first of all, cattle grazing). Some data on the mode of life of the Caucasian grouse in 13

the Lagodekhi Nature Reserve (typical habitat of the species) is found in the description of game animals of the reserve published by E.L. Markov (1938); this publication contains the solitary mentioning in the literature on the presence of about 20-25 birds in a single lek, with prevalence of females. The habit to dig themselves in the snow not only at night, but also at the daytime, was also mentioned (this habit is typical for grouse birds in the middle of winter). A case was mentioned when about ten birds flew away from under the snow being disturbed by the author. I.D. Chkhikvishvili (1939) gives the data on the distribution and ranging of grouses in the Kodora Canyon in Abkhazia, mentioning that they are numerous in mountains near the upper border of the arboreal vegetation in thickets of juniper, rowan, and shrubby birch. E.P. Spangenberg (1941) wrote about very easy grouse game in Armenia and on the harm to local grouse population as a result of hunting for broods with beagle dogs. Lyaister and Sosnin (1942) gave the detailed data on the distribution of the Caucasian grouse in Armenia, on its habitats, a propos, differing from those in the Main Caucasus, on dates of breeding, and on the mode of life. The authors again paid attention to the original whistling sound produced by males during flight. They mentioned that molting males did not produce this sound during the flight. STUDIES OF THE CAUCASIAN GROUSE IN THE SECOND HALF OF THE 20TH CENTURY The first volume of the 4-volumed review “Birds of the USSR” was published in 1951. In a sketch devoted to the Caucasian grouse, the data of Nesterov (1911) were used for the first time; it was also mentioned that this species is also spread in mountains Turkey adjoining Armenia. In a multivolume publication “The Life of Animals” (1952 and the following editions), A.V. Mikheev gave a brief review on the biology and ecology of the Caucasian grouse. K.M. Gambarov (1954) wrote about the distribution of the Caucasian grouse in the southern slope of the Main Range within the borders of Azerbaijan and about food of grouses (on the basis of the study of content of crops of 3 birds collected in upper Kusarchai River). The lek behavior was described by the author on the basis of the observations made by N.K. Vereshchagin and his personal communication; probably, that was the reason of some temporal mess: Vereshchagin reported that he observed lek (aggregations of 5 – 10 males and 6 females) in June 12 and 13. These dates are extremely late; even in more northern territories lek’s activity is finished completely to this time. In this area, female with the offspring was observed in June 26. The easternmost limit of the distribution of the Caucasian grouse in the Main Caucasian Range was pointed by the author as the upper Girdymanchai River in the souther slope and as the upper Kusarchai River, in the northeastern slope (47°50΄E). V.F. Larionov and A. Cheltsov (1958) published a brief review on the biology of the species based on the literary data, mentioning L.F. Mlokosiewicz as the first person who noticed the difference between the black and Caucasian 14

grouses. Actually, these differences were noticed by Radde, as we mentioned above; unfortunately, he published the results of his observations too late. Molamusov (1959) very briefly described the distribution of the Caucasian grouse in Kabardino-Balkaria. Later (Molamusov, 1961) the author analyzed this species in more detail. In this publication, he mentioned the main reason of the decreased population density of the grouse in KabardinoBalkaria, namely, pasturing of cattle in nesting territories of the grouse. Besides, the author gave some data on molting and reproduction of the grouse, in particular, he described 2 nests with 5 and 6 eggs, found by him in small plots among rhododendron thickets. The altitude where these nests were found (3500 and 3000 m above sea level) seems erroneous, making its identification doubtful. It is well known that the upper border of the rhododendron in Kabardino-Balkaria runs at altitudes below 2900 m above sea level. The earliest of 7 broods observed by the author was met in July 4. The author underlined the differences in feather coloration between one-year-old males and females and mentioned that the complicated adult plumage appears in males only after the third autumn molting; this is erroneous, because, as was described already by Lorenz (1884), male grouses become completely black already after the second autumn molting. The author also reported that he never met flocks exceeding 6 birds, and that in January 1960 he saw the same birds in the same places as in summer. M.E. Kutubidze (1961) performed the most detailed study of the ecology of the Caucasian grouse in the Greater and Lesser Caucasus within the borders of the Georgian SSR. He described the distribution of the grouse, its distribution in biotopes of Georgia, daily activity, molting, feeding, reproduction, and natural enemies of these birds. The most recent data on the distribution and population density of the Caucasian grouse in separate ranges of the Lesser Caucasus and confirmation of the existence of these birds in the Suramskii Mountain Range, connecting subalpine belts of the Greater and lesser Caucasus are the most valuable. The work contains the large volume of the original data devoted to different aspects of the biology of the Caucasian grouse in different localities of its range. The work is very valuable as the original source of new data; however, its accessibility is strongly limited by the fact of its publication in Georgian. The work was cited by R.L. Potapov in his monograph on tertraonid birds (Potapov, 1985), who received a Russian translation of this work from A.I. Ivanov only in 1980. I.B. Volchanevskii, I.I. Puzanov, and V.S. Petrov (1962) reported on the distribution and biotopes of the Caucasian grouse in the northwestern Caucasus, also giving the ecological, faunistic, and zoogeographical characteristics of the region. In June 15, 1935, they found 4 weakly hatched eggs in the fine beech forest of the subalpine belt; in July 7, 1935, a brood of young nestlings that could not fly in the Pseushkho Pass: and in July 11, nestlings able to fly. His report on the presence of the Caucasian grouse in forestless passes Nauzhi, Bekshei, and Tubinskii, and also in Mt Pseushkho, at altitudes of about 1000 – 1200 m above sea level in 15

1930ies (based on hunters reports), attracts attention. R.G. Zhordania (1962) gives literary data on the distribution of the Caucasian grouse in the Lesser Caucasus within the borders of the Georgian SSR, together with generalized data of breeding dates and molting of these birds. S.P. Chunikhin (1964) briefly describes the results of his observations on the Caucasus grouse in Kabardino-Balkaria, but does not separate his own results from the results obtained by other authors. In particular, he mentions the weight of 3-day-old and month-old nestlings (23 - 30 and 150 -200 g, respectively); analyzes the behavior of growing nestlings and females; reports on the significant degree of mortality in young nestlings during first weeks of life. According to the author, male molting begins in the second ten days of June, and remiges are exchanged slowly and gradually, do not influencing flight abilities of birds. It is also reported that molting males form temporary flocks and are very careful; that the species is rare outside nature reserves; he also gives some recommendations to hunters (in some regions, hunting for the Caucasian grouse was not prohibited at that time). N.M. Drozdov (1965) gives some data on the population density of the species in the western part of the Main Caucasian Range in Azerbaijan (Zakatalskii district), mentioning that the average population density of the Caucasian grouse in subalpine meadows in August 1961 constituted 2 specimens per 1 km2. A.I. Khanmamedov (1965) gives some data on the dates of reproduction and population density of the Caucasian grouse and also shows collecting sites as dots in the map eastwards to the upper Samur and Kusarchai Rivers; the easternmost point of distribution of the grouse in the Main Caucasian Range is pointed as the source of the Velvichai River, Mt Karlika-Dag (40º56”N, 48º32”E; 2631 m above sea level). The author does not mention the presence of this species near Mt Dyubrar. A.I. Khanmamedov and F.A. Aslanbekova (1965) observed the Caucasian grouse during the reproductive period in northeastern Azerbaijan. A total of 37 males of different age were obtained; the data on the weight of their testicles at different dates of the reproductive period are given. The authors give the data on the dates of lek activity and egg-laying; the arrangement and structure of 9 nests was described together with the mentioning of the number of eggs in each egg-laying (a single nest contained as much as 12 eggs, the record number), its weight and size; the duration of hatching (23-25 days) was established for the first time. The authors observed mass hatching of nestlings in the first weak of June (the earliest dates known for the Caucasian grouse). V.I. Tkachenko (1966), in his voluminous and detailed review of galliform birds of the northern Caucasus highlands (made mainly on the basis of the study of these birds in the Teberdinskii State Nature Reserve), described the ecology and biology of the Caucasian grouse. The author describes in detail the distribution and habitats of the grouse, seasonal changes of stations and associated migrations of birds. He analyzed the content of 30 crops and stomachs of grouses obtained in different seasons; as a result, he gives a detailed list of animal and plant food consumed by birds 16

in different seasons of the year; he distinguishes the main, secondary, and occasional food; calculates nutrition needs in dependence of the bird weight; and also estimates approximately food resources in different grouse habitats. The reproduction of the Caucasian grouse is described in detail, together with dates of lek activity, nest stations, and the structure of three nests. The author analyzes natural enemies of grouses and their endoparasites in detail. Besides, the work contains the data on the morphology of the species, its first plumage, and molting. Special attention is devoted to the population density of birds and probable reasons of its changes. The actual population density of birds in the Teberdinskii Reserve was estimated for the first time, constituting 1.1 specimens per 1 km2 (average data for 4 springs). The method of the registration of the population density of birds, associated with spring lek sites, suggested by the author, remains the most fruitful. On the basis of the material collected in the Caucasian Nature Reserve in 1950-1966 (the number of broods observed and the number of nestlings per brood), V.A. Kotov (1968) demonstrated the direct dependence of the breeding success of the Caucasian grouse on the climatic conditions during brooding and first days of life of nestlings; mentioning in particular that strong decrease of temperature and snowfalls decrease the average volume of broods by 2 and more times. When the number of nestlings per brood in different favorable years in June constituted 4.1 – 4.7, in unfavorable years, it fluctuated from 1 to 2.2. Strong cooling also resulted in freezing of females in nests. E.V. Chernyavskaya (1971) described the lek behavior of the Caucasian grouse. Unfortunately, her paper does not contain any new data, but, by contrast, is full of erroneous impressions, testifying to extremely inaccurate observations in the leks. A.A. Kuznetsov (1972) describes in detail his single observation of the morning lek in southeastern slope of Mt Ushba at an altitude of 2800 m above sea level. His observation of males that appeared in the lek site from neighboring slopes (they did not spend night in the lek) is very interesting. In 1974, the Ministry of Agriculture of the USSR established the Red Data Book of the USSR (a list of rare and endangered plant and animal species). The Caucasian grouse was included into this and all the subsequent issues of the Red Book. Airumyan and Margaryan (1974) determined the total number of Caucasian grouses in Armenia as 500 specimens. By the opinion of R.L. Potapov (1984), this number is lower than real one. N.I. Burchak-Abramovich (1974) published the first data on fossils of the Caucasian grouses from upper Acheulle-Mouster (about 300 000 years ago) in layers of a Paleolithic site in the Kudaro-1 cave (1700 m above sea level), in Mouster layers of the cave Kapshinskaya (250 m above sea level), in Mouster layers of a cave near Erevan, and also in upper Paleolithic layers (Madlen, about 80 000 years ago) of the cave Dvardzhis-Glde in Imeretia. Z.D. Sikharulidze (1974) published a very informative paper based on his all-year-round field observations of birds in Pshavo-Khevsuretia (northwestern Georgia). The author characterized the feeding of the Caucasian grouse in different seasons, 17

including winter. The author also determined the average population density of the species in summer; the latter varied from 2.9 to 4.8 specimens per square km (3.5 specimens on average) in hayfields and from 0.3 to 0.7 specimens (0.5 specimens on average) in pastures. These recording confirmed repeatedly published data on extremely negative influence of cattle pasturing on the population density of grouses. He also mentioned late lek of adult males (up to June 20). D. Scott (1975) and his Iranian colleagues revealed the Caucasian grouse in the northern slope of the Karadag Mountain Range, in a small area between Mt Kamki and Mt Suradzhartepe. This mountain territory adjoins the southern part of the Zangezur Mountain Range in Armenia; the Iranian population is separated from the Armenian one only by the deep valley of the Araks River. A year later, this author (Scott, 1976) described the state of a population of the Caucasian grouse in Kalibar Mts in the north of the Iranian Azerbaijan, where 200 – 300 birds were found in forest clearings and alpine forests at altitudes of 1800-2100 m above sea level. This species is protected in Iran since 1971, when the grouse game was prohibited; however, the species still suffers from human activity, ruining natural habitats, including protected territories of Arasbaran. M.N. Zhuravlev (1977) characterized the modern distribution and population density of the Caucasian grouse in the Caucasus. In particular, he pointed to the significantly higher population density of the species in the Main Range in comparison with the Lesser Caucasus; population density of grouses is higher in the western part of the Main Range (on average, 2.5 specimens per 1 km2, occasionally, up to 10 specimens per 1 km2). The lowest position of the border of the vertical distribution of the species in the northwestern Caucasus was also confirmed; in some places of these area, the grouse dwells at an altitude of only 1000 m above sea level and even somewhat lower. It was reported on cases of aggregation of these birds in groups of 20 and more birds in the end of summer-autumn, after the end of cattle pasturing. He also mentioned the disappearance of this grouse in the Lagonak Plateau because of over-grazing; later, this observation was disproved (Vitovich, 1986). The author mentions Mt Shessi as the westernmost point of the species range in the Main Range, and the line from middle Sulak River to Kuba (46°50́΄- 48º30΄ E) as the easternmost point. The author did not know about dwelling of the bird in Turkey and Iran. The eastern border of the species range in the Lesser Caucasus is described very uncertainly (Zangezur Mountain Range). R.L. Potapov and E.A. Pavlova (1977) performed a special study of the reproductive behavior of the Caucasian grouse (the work was conducted in the territory of the Teberdinskii Nature Reserve in 1973) in comparison with similar behavior of other tetraonid birds. For the first time they described in detail and illustrated the nuptial flight, the main element of reproductive behavior of theses birds, previously named the lek jump. The 18

large number of photographs allowed compilation of the complete scheme of bird movements. Noteworthy, video recording made later (Klaus et al., 1990) confirmed the accuracy of that description. Some elements of the lek’s behavior were described for the first time, including the “duel in parallel”, known for other tetraonid species, etc. Territorial and age structure of the lek was analyzed in detail; its three-step hierarchy was established; the size of dominating male’s plots in the lek’s arena were determined; and daily activity of birds in reproductive period was described. Comparison of peculiarities of sexual behavior of the Caucasian grouse and other tetraonid birds allowed revealing these peculiarities; their functional analysis was given for the first time and possible trends in their development were assumed. In particular, the absence of noticeable role of the male tail in the breeding ritual (by contrast to other species of the family) was explained and its significance as a signal element important for the location of the male in dense grass was shown; advantages of a 180° turn during lek’s jumping flight were analyzed; it was assumed that the “silence” of the breeding ritual is the explainable result of the reduction of vocal elements of the reproductive behavior in highlands, etc. In other words, the adaptive character of these peculiarities stipulated by specificity of the highlands was demonstrated. Moreover, the conclusion, that the rate of evolutionary changes of the breeding behavior in the black and the Caucasian grouse differs, was made for the first time. This rate is pronouncedly less in the Caucasian species owing to its rather small range of island character in relatively stable highland environment. The success of field studies was to a greater extent stipulated by the help of O.V. Vitovich, a zoologist of the Teberdinskii Nature Reserve, who knew in detail the arrangement of leks in the territory of the reserve. During discussions, the authors interested O.V. Vitovich in further investigations of the biology of this poorly examined species. During next 10 years, O.V. Vitovich fulfilled nearly all the tasks that were envisaged during discussions and that appeared during his studies; his investigations covered mountains of the Lesser Caucasus. In his first work (Vitovich, 1977а), he gave the material on the reestablishment of the population density of the Caucasian grouse in the territory of the Teberda Nature Reserve after finishing of cattle pasturing in 1943. Partly, these data were published earlier (Tkachenko, 1966), but Vitovich continued these studies and found that the population density of the species, which increased to 1964 from 1.4 to 3.8 specimens per 1 km2, later stabilized and only small fluctuations were observed; these fluctuations are stipulated only by natural factors, the most important of which included the weather during brooding. In another work (Vitovich, 1977b), the first data on the embryonic mortality of the Caucasian grouse were given; these data were obtained as a result of observation of 13 nests with 80 eggs. Mortality differed in different years (fluctuations from 0 to 43%) and depended, first of all, on the weather conditions during brooding; embryos froze to death during strong cold snaps with snowfalls. In his third work, 19

Vitovich (1977c), besides of these data gave another ones concerning the biology of breeding and analyzed also conditions necessary for reestablishment of the population density of the grouse in territories used as pastures. Field material obtained during his work in the Teberdinskii Nature Reserve and also examination of the voluminous material deposited at the Zoological Museum (Moscow State University) and especially at the Zoological Institute RAS in St. Petersburg, allowed R.L. Potapov (1978) publishing one more work, in which characters common for the species of this group were revealed together with the highland specificity. Before this investigation, R.L. Potapov had studied tetraonid birds already for 10 years. The mentioned publication contained the first detailed map of the species range, where reliable collecting sites were designated by dots. The analysis of archaic features that retained in this species allowed the author talking about close relations between the ancestral forms of grouses and willow grouses. The structure of brows in the Caucasian grouse, similar to that in the willow grouse Lagopus lagopus is very demonstrative in this respect (The sketch of brow transsection was given for the first time). The original shape of the tail in male grouses (and, to a lesser extent, in females) is also analogous to that in willow grouses (not so strong but, nevertheless, quite evident). The delay in the growth of central and, occasionally, also of external tail feathers in the latter species during termination of molting results (in some cases) in the formation of the fork-shaped tail. It was also revealed that the Caucasian grouse possesses the longest legs among other species of the family (distinct adaptation to walking along steep slopes); its short and wide wing is adapted to the sliding-diving flight, advantageous in mountain landscapes and similar in structure to the wing of representatives of the genus Tetraogallus. The mechanics of the flight was analyzed and the source of whistling sounds accompanying the flight were revealed: the sounds are produced by primary remiges; the latter demonstrated the strong deterioration of inner margins of the vexilla in one-year-old and older birds (the photo of them and, for comparison, the same ones in Lyrurus tetrix were done). Finally, the first hypothesis on the origin of the Caucasian grouse was assumed in this publication. According to this hypothesis, the ancestral form had subdivided into two species in the Middle Pleistocene, and the ancestor of the Caucasian grouse reached Caucasus being already the mountain bird. This species was formed in Balkan mountains, where a series of early tetraonid birds were found; further on, the species spread eastwards along mountains of the Asia Minor Peninsula as far as the Caucasus during one of cold epochs of the Middle Pleistocene. Later, in the “Priroda” popular science magazine, Potapov (1978a) characterized the biology of the Caucasian grouse; gave a map of its range; and offered some methods of its protection. In the Red Data Book of the USSR (1984), Potapov estimated (for the first time) the total number of grouses as about 70000 specimens. A.D. Lipkovich (1985) reported on comparatively well situation with the population density of the 20

Caucasian grouse in the North Ossetian State Nature Reserve (he gave no certain numbers), and also gave some data on the phenology of the species. In his monograph devoted to tetraonid birds, Potapov (1985), together with the data on all the taxa of the family Tetraonidae from genera to subspecies, summarized all the data on the Caucasian grouse, including his own data. This is the most complete description of the Caucasian grouse performed in the 20th century; it includes the data on the coloration of birds of all the ages, their size and weight, distribution, population density, mode of life in different seasons, feeding, breeding, data on paleontology, and all the other data allowing understanding of the origin and evolutionary history of the species. This study was based on the own data of the author and also on examination of the voluminous collections from museums and institutions of the USSR and all the available literary data. The author demonstrated the presence of all the typical tetraonid features in the Caucasian grouse, and also the development of specific features associated with the adaptation to highland dwelling. The work was supplied with the most accurate at that period map; in subsequent publications of different authors it was rather copied or copied with mistakes. The existence of two groups of populations in the species range: in the northern part of the range (the Greater Caucasian Range) and in the southern one (ranges of the Lesser Caucasus and adjoining ranges of the Turkey and Iran), connected by a fine thread passing through the Suram Mountain Range, were shown for the first time.. In the following 1986, a very important work of Vitovich (1986) to the ecology of the Caucasian grouse was published. This publication, the most complete of all the ecological studies, was based mainly on the personal experience of the author, who collected the material not only in the Greater, but, which is more important, in the Lesser Caucasus, wonderfully supplemented the monograph by Potapov (1985). As a result, the Caucasian grouse turned from the most poorly studied species of the family Tetraonidae into one of the most completely studied species. The ecology of this species is better studied in the Main Caucasian Range. Vitovich (1986) demonstrated that these parts of the range strongly differ in the climate and vegetation, testifying to a rather high degree of ecological plasticity typical of the Caucasian grouse just in the subalpine habitats. In spite of differences in northern and southern habitats, association with the same natural complex (the border between the forest and adjoining subalpine meadows and shrubs, their characteristic in all the parts of the range. The mentioned work also describes in detail the habitat’s preferences and mode of life of birds in different seasons; flock behavior and sex ratio; population density and its dynamics; breeding period and lek characters; nesting period; growth, development, and molting of nestlings; feeding (including the data on daily nutrition needs); natural enemies, etc. The publication also included a large number of new facts, in particular, the unknown previously winter lek activity, some courtship poses and rituals (lek jump without turning), etc. Tilba and Cheprakov (1986) reported on the 21

strong (nearly by 2 times) decrease in the population density of the Caucasian grouse in summer of 1983, after the bacterial “attack” of an unknown origin that had destroyed shrubs and arboreal vegetation that usually stays green in winter (coniferous plants, rhododendrons, berry shrubs, etc.) in the early spring of that year. Authors concluded that this event strongly decreased the volume of the nutritive supply of birds, feeding just on these plants. They also noted that the strong cooling with snowfalls was observed in June 7-8 of the same year, i.e., during the breeding period; this phenomenon usually also strongly decreases the population density (Vitovich, 1976). In the voluminous book “Birds of the USSR. Gruiformes and Galliformes”, published in 1987, R.L. Potapov gave a brief description of the all species of grouse Family, including Caucasian Black grouse. Klaus et al. (1988) described the breeding behavior of the Caucasian grouse observed by the authors in the Teberdinskii Reserve in spring of 1987 (May 12 - 22) after a snowy winter, when the first lek stages were performed on snow cover. The paper was illustrated by drawings made from photographs and 16-mm film. The authors assumed that the reduction of all the white feathers in males, typical of the black grouse, became unnecessary in the high grass. According to the same reason, males do not spread wings during courtship and also during the copulation. Prilutskaya and Pishvanov (1989) confirmed the presence of the Caucasian grouse in Tlyarotinskii, Tsunkinskii, Tsummadinskii, Sovetskii, Rutulskii, and Akhvakhskii districts of Daghestan, in all the places with coniferous forests with admixture of birches. The report on the presence of the Caucasian grouse in foothill districts (Buinakskii and Kaitagskii districts) seems doubtful. The total number of grouses dwelling in Daghestan was estimated by the authors as 1100-1200 birds. A.D. Lipkovich (1989) pointed to the negative influence of forest felling and cattle’s grazing on the population density of the Caucasian grouse. In 1990, a group of authors (Klaus, Bergmann, Marti, Muller, Vitovich, and Wiesner) published a voluminous review devoted to Black grouses. This publication acquainted readers with different aspects of bird biology, including paleontological data, modern distribution, geographical variability, peculiarities of feeding and reproduction, etc.; hybrids formed with participation of Caucasian grouses were also described. This review also included a section devoted to the Caucasian grouse (pp. 198 - 266), compiling the data published earlier by Potapov (1978, 1985, 1987, 1989) and Vitovich (1986). These data were supplemented by the results of own observations of the authors performed during several trips to the Teberdinskii Nature Reserve, where Vitovich, who worked at that time in the Reserve, acquainted his co-authors with the biology of the species. Vitovich is also the author of the most of photographs and of several schemes illustrating the review. The authors had also filmed lek jumps and recorded sonograms of sounds produced by jumping and flying males. In 1991, H.H. Bergmann, S. Klaus, J. Wiesner, and O.A. Vitovich (Bergman et al., 1991) reported again that males of the Caucasian grouse do 22

not use vocal signals during courtship displays, but can produce sounds by their wings. These sounds are produced during jumps and flight and are represented by two types: metallic whistle during the gliding stage of the flight and wing flaps, that was established before (Potapov, Pavlova, 1977; Potapov, 1978). Flying females occasionally produce similar whistling sounds. Two sound sources on wings can work independently. Besides, fighting or copulating males can produce low caws by the larynx, that was mentioned before by several authors. In his work devoted to monitoring of birds dwelling in the territory of the North Ossetian Nature Reserve, Komarov (1991) gave some data on the population density of the Caucasian grouse. In subalpine meadows in summer, population density of the species constituted 7 and 9 specimens per 1 km2 in August and September, respectively. Temple-Lang and Cocker (1991) mentioned finding of the Caucasian grouse in rhododendron thickets in mountains of the Pont Mountain Range in Turkey. Tkachenko (1992) reported on meeting with a brood of the Caucasian grouse, with seven 2-3-day-old nestlings in June 7, 1991. By phenology dates, spring of that year was quite usual and other broods were met in usual dates, i.e., in the end of June. The author notes that such early breeding dates were recorded only in the southern Transcaucasia (Khanmamedov and Aslanbekova, 1965). In the book “Birds in Europe”, devoted to preservation of endangered European bird species, Potapov (1994) gave a brief review of the distribution, ecology, and general abundance of the Caucasian grouse. The author estimated the number of grouse specimens approximately constituting 70000 in the Greater Caucasus (including Russia, Georgia, and Azerbaijan) and 500 in the Lesser Caucasus (Armenia). Tilba (1994) describes findings of the Caucasian grouse in different parts of the Caucasian Nature Reserve and adjacent territories, including the number of birds, collecting dates, and sources of information. He concluded that the population density of the species did not decrease in the northwestern part of the range, and that a report on finding of the Caucasian grouse in the Gerpegem Heights (Rossikov, 1890) belongs to summer birds that allegedly cannot breed in that kind of area, because only dry steppes, but not subalpine landscapes are found there. At the same time, the author missed the fact that Lorentz (see above), who had described the lek of the Caucasian grouse for the first time, was basing on observations performed in the Mt Bermamyt, situated even northerner and at the same altitude of 1250 m above sea level. In the same Caucasian Nature Reserve, Averin (1938) pointed to several mountain peaks lower than 1000 m above sea level, where, nevertheless, the Caucasian grouse was successfully dwelling. Atkinson et al. (1995) gave the results of the study of distribution of the Caucasian grouse in the Pontiiskii Mountain Range (northeastern Turkey). The northernmost point of the range was Mt Ablaris (40º N). In a 150-km part of the range, they found 33 leks with 138 males. Unfortunately, the authors did not mention, whether only black mature males were recorded, or gray one-year-old males were recorded also. 23

In any case, the authors demonstrated that these birds are not rare in the area; nevertheless, the species remains endangered one. In his short thesis, Potapov (1995) described adaptive morphological peculiarities of the species, promoting its highland dwelling; these features were described earlier in Russian (Potapov, 1978). In an Atlas of European birds, Tilba and Potapov (1997) gave the general data on the species and its distribution, and the latest data on its abundance. According to these data, the Greater Caucasus is populated by 70000 specimens (Potapov, 1984, 1985); the Lesser Caucasus, by approximately 8400, and Armenia, by 500 birds (Airumyan and Margaryan, 1974). On the basis of unpublished data of Potapov, 2500, 400, and 75000 birds were mentioned for Turkey, Azerbaijan, and Georgia, respectively. Solokha (1997) published the results of his field observations on the Caucasian grouse in four sites in Armenia: in the center of the Zakhungans Mountain Range near Takiyarlu (Artavaz) and Megradzor, 23002800 m above sea level, 40˚32’N, 44˚30’ - 44˚37’E, where he saw 10-11 and 5-7 birds in May 17-23, 1995 and June 4-21, 1996, respectively. The second site included the eastern part of the Pambak Mountain Range near Dilizhan, 2050-2550 m above sea level, 40˚41’N, 44˚48’E, where he saw no birds in June 24 -27, 1996, although other authors reported that this species is common in this area. The third site included northeastern parts of the Zangezur Mountain Range near Kadgaran, 3000 m above sea level, 39˚08’N, 46˚05’E, where only 2 or 3 birds were recorded in July 4-6, 1996. The fourth site included the central part of the Takhniyants Mountain Range near Agveran, 2300-2500 m above sea level, 40˚30’N, 44˚33’E, where 9-10 birds were recorded in July 10-14, 1996. Extrapolating these data on all the suitable places, the author assumed that the total number of birds in Armenia constitutes about 100 -120 specimens. Lipkovich (1999) hypothesized on the origin of the Caucasian grouse from plain populations of the ancestor form dwelling in the Russian Plain to the north of Caucasian Mountains. However, no paleontological data on findings of tetraonid birds in the southern Russian Plain in the first half of the Pleistocene are mentioned by the author. The reason is simple: these data are absent. One of the main arguments of the author in favor of a hypothesis on penetration of the ancestor of the Caucasian grouse from the north includes correlations between modern distribution of the grouse and the presence of such trophic plants as cowberries and bilberries, which penetrated to the Caucasus from the north; the range of these plants supposedly coincides with the range of the grouse. In reality, the degree of coincidence is only 60%. In summer of 1998, Drovetski and Rohwer (2000) performed observations and collected the material in the northwestern border of the grouse range in the Lagonakskoe Plateau; in summer of 1999, they examined the Magisho Mountain Range. They observed a total of 27 broods, mainly in meadows, but also in gullies and in pine forests. One-year-old males were also noted in those places. Adult males (7 – 8) preferred other habitats, poorer in food but safer from predators, namely, high grass plots in gullies. A nest 24

with 5 eggs was also found. Broods observed differed in age by 2-2.5 weeks. The presence of “late” broods point to later breeding dates of young females rather than to secondary egg-laying. The authors also determined population density of adult birds in summer; this density constituted 2.3 specimens per km2, corresponding to the population density of the Caucasian grouse over the entire range. Kaphengst (2003) saw the reasons of the decrease in population density of the Caucasian grouse in over-pasturing of cattle in alpine meadows, forest cutting, disturbance caused by cattle and humans, poaching, and dogs (cit, from Klaus et al., 2003). Klaus and Storch (2003) described their observations performed in September 24 – 26 in slopes of Mt Kazbek on a small (no more than 18 specimens) flock of Caucasian grouses comprising young and mature males and females. Birds were feeding, but in the evening of September 25, some adult males demonstrated several elements of the courtship rituall, including jumps, although these males were not very active. Klaus et al. (2003) summarized the literary data and their own material on the ecology and behavior of the Caucasian grouse obtained during 7 field seasons in different places of the Caucasus, mainly, in the Teberdinskii Reserve. Territorial and lek behavior was analyzed once more, and early data on sounds produced by lek birds were confirmed. Baskaya (2003) examined the eastern part of the Pont Mts. The authorstudied the distribution of the grouse in the Karadeniz Mts and determined its population density as 3.9 birds per square km on average (0.4 - 7.75); additionally, he pointed to the factors negatively affecting the population density of these birds, including game activity (both legal and illegal ones) and also egg collecting, predation by foxes and shepherd dogs, disturbance from berry collectors, photographers, alpinists, and other humans appearing in nesting sites of the Caucasian grouse. This negative influence is supplemented by forest cutting, active pasturing of cattle, construction of roads in mountains, etc. The author confirmed the distribution of the grouse along these mountains as far as 40˚E, observed earlier (Atkinson et al., 1995). Potapov (2004) supplemented his talk on the International Symposium on Tetraonid Birds, made in 1993, by new data on the absence of geographical variability and on the reasons of the presence of such an unusual plumage in one-year-old males, strongly differing from that in mature males. This plumage plays an important cryptic role, especially in winter. The first winter is spent by males in flocks, together with females, dwelling among forest vegetation, by contrast to mature males, living separately in more exposed stations near the upper border of the forest. The author gave the data on perfect flying abilities of males that can widely migrate within the range, providing genetic exchange between populations. The Suram Mountain Range, connecting the Greater and the Lesser Caucasus and still populated by the Caucasian grouse, also promotes this exchange. Segelbacher and Storch (2004) analyzed the genome DNA from molted feathers by PCR with the following sequencing in the polyacrylamid gel. This method was already used by the authors 25

for the analysis of the Alpine population of the Black grouse. As a result, genetic differences between three main populations of the Caucasian grouse (Armenian, Azerbaijan, and Georgian ones) were demonstrated. At the same time, these data did not answer on the question on the degree of geographical isolation of these populations. Sultanov et al. (2004) examined the state of population of the Caucasian grouse in the Azerbaijan part of the Lesser Caucasus and confirmed that the main habitats of birds are represented by subalpine meadows and marging of birch forests, forming these the upper border of the forest zone. The diapason of altitudes of grouse habitats varied from 1800 - 2000 to 2500 m above sea level. Although some rhododendrons (along northern slopes) and juniper (along southern slopes) are found in grouse habitats, the bird is associated mainly with the birch, especially in winter, when birch buds form its main food. The total number of these birds in the Azerbaijan part of the Lesser Caucasus is estimated as 1000 – 1500 specimens. The authors confirm the fact that the population density of the species in the Lesser Caucasus is significantly (by 3-4 times) lower that in the Greater Caucasus. G. Dzhamirsoev (2004) published the short paper, in which offered the hypothesis about the penetration of the ancestor of Caucasian grouse to the Caucasian Mountains through the South Russian Planes in the end of Pliocene (i.e. repeated the hypothesis of Lipkovich, 1999) without any real paleontological evidences and without any knowledge about the environmental situation in these plains at that times. The English variant of this paper was published later (Sultanov, 2006). It included only the data on the state of Caucasian grouse populations in the Nakhichevan Autonomous Republic and did not mention the Karabakh Highland, where both distribution and population density of the grouse stay unexamined. In the end of the 20th century, the Caucasian grouse was included into the list of rare and disappearing species; it was included into the first edition of the Red Data Book of the USSR (1984); attention of ornithologists was attracted to the study of its population density and the range. With the help of international organizations (including financial support), and first of all, with the help of the BirdLife International, the activity of ornithologists of Azerbaijan, Armenia, Georgia, Iran, and Turkey was activated. This continuing activity is manifested in organizing of symposia and in performing of different scientific projects, e.g., the project “Caucasian Black Grouse Research, Monitoring and Conservation Management in Georgia” (2004). The data on finding of this species in some points in Georgia, mainly based on the material of different authors, including hardly available papers in Georgian, are the most interesting in the publication on the project mentioned. Unfortunately, only a single finding of Radde was mentioned (he found 7 birds); his map on the distribution of the grouse in the Greater and Lesser Caucasus was also forgotten. Potapov (2005) published the results of his studies on the biology and phylogeny of the genus Lyrurus, 26

explaining the impossibility of its uniting with the genus Tetrao, and mentioning arguments in favor of close phylogenetic relations between the genera Lyrurus and Lagopus. The author pointed to the structure of the “brow” in the Caucasian grouse, treated as a relic inherited from the ancestor common with representatives of the genus Lagopus. Masoud and Fanid (2006), citing their previous publications (1993а, 1993b, 1995, 2004а, 2004b), mentioned that the number of Caucasian grouses in eastern Azerbaijan (10 000 ha of the Azerbaijan Biosphere Reserve and 20 000 ha to the west of the reserve) constituted no more than 250 specimens and, apparently, it decreased since 1971, when, according to Scott (1975), 200 – 250 birds were dwelling only in the protected territory of the reserve. The authors explain this phenomenon by high degree of nestling mortality during spring rains and cold and also by human agricultural activity. Finally, R.L. Potapov (2008) once more documented his version on the penetration of the Caucasian Grouse ancestor into the Caucasus from mountain regions of the Balkan Peninsula along mountain systems of the Anatolian Peninsula, using the most recent data on new early findings of the tetraonid birds in Balkans and on the character of environmental conditions, mainly of the vegetation, existing in these mountain systems in the late Pliocene – early Pleistocene. In this work, the impossibility of accepting a hypothesis on the penetration of the ancestor form into the Caucasus via southern Russian plains is well reasoned. THE FURTHER GOALS IN STUDIES OF THE CAUCASIAN GROUSE As seen from our review, no monographs devoted to the Caucasian grouse were published during the last 20 years. After Vitovich (1986), only some reviews were published, only supplementing the existing data with some new features of confirming the old data. This phenomenon creates an impression that the species is studied sufficiently, although this picture is not true. 1. The complete scheme of the species range is still absent. “White spots” exist in Azerbaijan (the Karabakh Highland), Daghestan, and Turkey. In the latter country, the distribution of the grouse is studied only in the Northern Pont Mountain Range. Both size of the range and its borders remain unknown. It is especially true for those parts of the range that are situated in Daghestan and eastern Turkey, and also in the eastern part of the Lesser Caucasus. It is necessary to perform field investigations in these regions in order to reveal range borders, both geographical and topographical (altitude above sea level). This will allow estimating of the stability of the range during last 50 years. 2. It is essentially necessary to investigate in detail the mode of life of the grouse in the southern part of its range, including mountains of Turkey and Iran, in the way it was done for the northern part of the range by Tkachenko, Potapov, and Vitovich. Only these data will allow estimating 27

adaptive peculiarities of the species, populating areas with strongly different environmental conditions without forming separate geographical forms. 3. The structure of the most of local populations of this species remains poorly examined. Its study is very important although hardly performed in highlands. The ringing and especially the method of radio-tracing, never used for the study of this species, seems most prospective in relation to the revealing of mobility of separate individuals in dependence of their sex, age, and season. Territoriality of males and females (in particular, movements of males between different leks during the reproductive period), dispersion of young birds can be studied with the use of local or satellite microtransmitters. These data were very important for organizing the effective measures for protection of the species protection in different parts of the range. 4. It is necessary to elaborate methods of maintenance of birds in captivity, including their reproduction, and also to obtain hybrids between Caucasian and Black grouses for further studies of ancestral characters, probably appearing in the offspring, and also for the study of the degree of stability of species characters, first of all, morphological and behavioral (voice) ones. Such studies can be very useful for the study of the origin and evolutionary history of both species. 5. It is necessary to take part in any archeological and paleontological works performed in caves of the Anatolian Peninsula, in order to obtain new paleontological material important for the study of the genesis and evolution of the species. REFERENCES Airumyan K.A. and Margaryan P.A. 1974. Resources of Game Birds in Armenia. (Tezisy VI Vsesoyuznoi Ornitologicheskoi konferentsii, Mocsow), pp. 239 – 241. [In Russian] Atkinson W., Humpage W.A., Jowitt A.D.J., Ogurlu I. and Marcus J. 1995. Distribution and status of Caucasian black grouse in northeast Turkey. Proceedings of the VI International Grouse Symposium: 131 – 133. World Pheasant Association, Raeding, Institutio Nationale per la Fauna Selvatica, Ozzano dell’Emilia. Averin Yu. V. and Nasimovich А.А. 1938. Birds of the mountain part of the northwestern Caucasus. Trudy Kavkazskogo Gosudarstvennogo Zapovednika, 1: 5 – 56. [In Russian] Averin Yu. V. 1938. The Caucasian Grouse. Trudy Kavkazskogo Gosudarstvennogo Zapovednika, 1: 56 – 86. [In Rusian] Baskaya S. 2003. Distribution and principal threats to Caucasian black grouse Tetrao mlokosiewiczi in the Eastern Karadeniz Mountain in Turkey. Wildlife Biology, 9: 377 – 383. Beme L.B. 1925. Contribution to the biology of the Caucasian grouse (Lyrurus mlokosiewiscii Tacz.). Ukrainskii Rybolov I Okhotnik, 4: 18-19. [In Russian] 28

Beme L.B. 1926. Birds of northern Ossetia and Ingushetia. Uchenye Zapiski SeveroKavkazskogo Instituta Kraevedenia, 1: 18 – 19. [In Russian] Bergmann H.H., Klaus S., Wiesner J. and Vitovich O.A. 1991. Die “stumme Balz”: Lautaußerungen mannlicher Kaukasusbirkhuhner (Tetrao mlokosiewiczi) auf dem balzplatz. Journal für Ornithologie, 132: 267 - 278. Bilkevich S.I. 1893. Materials to the study of the ornithological fauna of Daghestan. Supplement to Protocols of the Society of Naturalists at the Kazan University, 24 (135): 1 – 24. [In Russian] Bogdanov M.N. 1879. Birds of the Caucasus. Trudy Obshchestva Estestvoispytatelei pri Kazanskom Universitete 8 (4): 1 – 188. [In Russian] Burchak-Abramovich N.I. 1974. Fossil birds from Palaeolitic sites of the Caucasus. Ornitologiya, 11: 329 – 333. [In Russian] Caucasian black grouse research, monitoring and conservation management in Georgia. 2004. Quarterly Progress Report N 1. GCCW, WPA, U of I and BirdLife International. 16 pp. Chernyavskaya E.V. 1971. Peculiarities of lekking behavior of the Caucasian mountain grouse (Caucasian Reserve). Trudy Voronezhskogo Gosudarstvennogo Universiteta, 93: 88 – 90. [In Russian] Chkhikvishvili I.D. 1930. Materials on the bird fauna of Kakhetia (Telavi district). Zakavkazskii Kraevedcheskii Sbornik, 1: 1 – 24. [In Russian] Chkhikvishvili I.D. 1939. To the fauna of mammals and birds of Abkhazia. Pp. 1-44 in Materialy k faune Abkhazii [Materials to the fauna of Abkhazia], Tbilisi. [In Russian] Chunikhin S.P. 1964. Caucasian grouse. Okhota i Okhotnich’e Khozyastvo, 8: 22 – 23. [In Russian] Deryugin K.M. 1899. A report on journeys and zoological observations in the Choroikhskii Territory (western Transcaucasia) and environs of Trapezund. Trudy Sankt-Peterburgskogo Obshchestva Estestvoispytatelei, 30 (2): 49 – 115. [In Russian] Dinnik N.Ya. 1884. The Caucasian grouse (Tetrao mlokosiewiczi Tacz.). Priroda I Okhota, 3: 15 – 20. [In Russian] Dinnik N.Ya. 1884а. Oshten and neighboring parts of Kuban Province. Zapiski Kavkazskogo Otdela IRGO, 16: 357 – 421. [In Russian] Dinnik N.Ya. 1884b. Mountains and canyons of the Kuban area. Zapiski Kavkazskogo Otdela IRGO, 13 (1): 307 – 363. [In Russian] Dinnik N.Ya. 1884c. Mountains and canyons of the Tersk province. Zapiski Kavkazskogo Otdela IRGO, 13 (1): 1 – 48. [In Russian]

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