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The zoogeography of theYamal‐Gydan area, Western SiberiaG. L. Rutilevskiy

Version of record first published: 23 Dec 2008.

To cite this article: G. L. Rutilevskiy (1979): The zoogeography of the Yamal‐Gydan area,Western Siberia, Polar Geography, 3:2, 88-117

To link to this article: http://dx.doi.org/10.1080/10889377909377107

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THE ZOOGEOGRAPHY OF THE YAMAL-GYDAN AREA,WESTERN SIBERIA

G. L. Rutilevskiy

From: Yamalo-Gydanskaya oblast': fiziko-geograficheskaya kharakteristika[The Yamal-Gydan area: physical geographical characterization].

R. K. Sisko (ed.).Leningrad: Gidrometeoizdat, 1977, pp. 226-260.

Abstract: The author has compiled a detailed survey of the fauna of the Yamal-Gydan area, the major stress being on the ranges and movements of all the species withinthis tundra environment. Utilization of these wildlife resources is discussed whererelevant. The author has relied both on the available published literature, and particularlyon field research during the Arctic-Antarctic Institute's expedition to the area in1971-1972. (The translation is by William Barr, University of Saskatchewan, Saskatoon.)

In examining the fauna of the Yamal-Gydan tundra (Fig. 1) it is necessary totouch briefly on the composition of the fauna of the Arctic as a whole; this allowsone to sketch in the specific peculiarities of this area against the general back-ground. In so doing one must bear in mind that the paleontological data for theselatitudes, relevant to the vertebrate fauna, are so meager that they cannot provide afull picture of that fauna. If one excludes the fauna of the mammoth complex,containing remains of mammals, there are practically no paleontological materialsrelating to other classes of vertebrates within the limits of the present Arctic. Apartfrom the present Arctic areas, representatives of the mammoth complex are alsofound far to the south of their main habitats.

Thus in any appraisal of the mammalian fauna one must rely mainly onpaleontological data derived from only a very brief time period. As is well known, areassessment is now under way of many theories regarding the dating of variousdeposits, the tendency being to move the dates closer to the present. Questions arebeing clarified with regard to ice cover in general, and in particular the existence ofan ice cover on the low-lying plains of many areas of Siberia, including theYamal-Gydan area, is being called in question. Ideas as to the dominant role ofmollusks are changing. Current methods of such investigations are still not per-fected.

As is well known, during the Karginian Period, many species of mammals died out;these included mammoths, oxen, bison, horses, cave lions, woolly rhinoceroses, etc.;and also cave bears, fossil elk and some species of deer which have retained theirsystematic classes.

Within the Eurasian Arctic the muskox has died out, but it has survived inGreenland and on the islands of the Canadian Arctic Archipelago down to thepresent. Finally, the mammoth complex of the Karginian Period also includespresent-day mammals: deer of the genus Corvus, the saiga and the present reindeer,the fox, wolverine, wolf, Arctic hare, Arctic and Ob' lemming, and some others.

While the fossil mammalian fauna permits one, to some extent, to fix the dateof its extinction, the emergence of present-day species, unfortunately, remains

88ISSN 0148-7671/79/0020-0088$07.50/0

© 1979 Scripta Publishing Co.

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LBELYY is.x^£?*v:::::;::::::x :7DIKSON \

SIBIRYAKOV IS. \

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Fig. 1. Location map of Yamal and Gydan Peninsulas.

unclear. One can assume that it occurred considerably earlier than the disappearanceof the main body of the mammoth complex.

During the flourishing of that complex, woodland forms also lived side-by-sidewith the animals of the open country. Since it is known that during the KarginianPeriod land extended considerably farther north, one must assume that many of theoffshore islands did not exist as such, and that part of the present shelf comprisedpart of the continent. Since the climate was somewhat milder than at present, the

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natural zones were displaced north, and the forest grew farther north than itspresent limits. This provides evidence that even during the short period of Karginiantime natural conditions underwent significant modifications. Hence it is notsurprising to find remains of ecologically differing animals within the present Arctic.One must also take into consideration the fact that the fossil finds, as a rule, aremainly redeposited materials, even in the case of assemblages of massive fossils.

It follows from what has been said that in analyzing the present fauna it ismore expedient to consider the distribution of species and their ecological peculi-arities. If one examines the vertebrate fauna of the Arctic as a whole, one has torecognize its uniformity and homogeneity with regard to speciation. The presentfauna of the Arctic areas of North America and Eurasia are almost indistinguishablein terms of species variation. In both cases the main taxonornic differences scarcelygo beyond the level of species. Endemic species are encountered only in isolatedcases. Naturally, in the present context, one has in mind the strict Linnaeanapproach to species determination.

Throughout the Arctic the distribution of ranges obeys the same laws: viz. first,as one moves north the quantitative composition of species decreases (the phe-nomenon of latitudinal zonality); second, as one moves from west to east, or in thereverse direction, there usually occur taxonomic differences generally of specific orsubspecific significance (the phenomenon of provincial differentiation); third, themajor systematic differences are more markedly noticeable between species of theAtlantic vs. the Pacific sectors of the Arctic, than between animals occupying thecorresponding areas of North America and Asia. These circumstances emphasize thatthe formation of the Arctic fauna occurred synchronously.

Thus one may assume that the establishment of the fauna in these land areasand bodies of water was subject to the same or similar influences which affected itsdevelopment during more remote epochs. Even during the Quaternary, inasmuch asthe paleontological materials are clearly not unequivocal, it is best to resolve thesequestions by means of a comparison of the present ranges and the influence of theinorganic environment on the fauna as a whole [18].

A fleeting glance at the map of Siberia is all that is required to assess theexceptional role of the marine and fresh bodies of water bordering the coasts andalso the interior areas of the Yamal-Gydan tundra, as compared to other areas ofthe Arctic. These factors are of considerable significance in the formation anddevelopment of the fauna in general, and of the aquatic fauna in particular. Theinfluence, on the one hand, of the discharge of such large rivers as the Ob' and theYenisey into Ob' Bay and Yenisey Gulf, whose regimes are influenced by the KaraSea, and, on the other hand, of the penetration of fresh water northwards beyondthese gulfs, inlets and offshore islands, make a marked impression on the distri-bution of aquatic fauna of both marine and continental origins.

The exchange of aquatic fauna between the sea and the gulfs, and also therivers, occurred more intensively at an earlier date. It is sufficient to point out thatduring the Yamal transgression, such Arctic representatives as the omul and the sealpenetrated up the Yenisey as far as Lake Baykal, and have since become totallyfreshwater species. Apart from this, the mingling of the habitats of the freshwaterand marine plankton and benthonic fauna is extremely typical of the Yamal-Gydanarea; freshwater forms persist in the inshore coastal areas of the Kara Sea and

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marine forms penetrate far to the south. In this respect some similarities can beseen between the Yamal-Gydan area and the White Sea. Thus the White Sea has servedin its time as a water route for the colonization by seals of several freshwater lakes.

The abundance of plankton and benthos in the bodies of water has had afavorable impact on the distribution and colonization by fish of both Ob' Bay andYenisey Gulf, these being excellent fishing areas. It is not by accident that theichthyofauna of the Yamal-Gydan area occupies first place in the Arctic in terms offisheries resources.

Fish

The Yamal and Gydan peninsulas projecting far out to sea, the markedlydissected coastline, the gulfs and inlets running deep into the continental landmass,the large number of rivers and lakes concentrated in the relatively restricted areabetween the Urals and the Taymyr Peninsula, and also the influence of thedischarge of such large rivers as the Ob' and the Yenisey, all place this region in anexceptional position in terms of both marine and freshwater habitats, conditionswhich are extremely favorable to colonization by fish. These circumstances have anequally positive impact on the development of aquatic invertebrates, including bothbenthos and plankton, which serve as food for many fish.

Apart from normal fish migrations provoked by ecological peculiarities andassociated with definite seasons (spawning runs, feeding migrations and shoaling runsin Ob', Taz and Gyda bays, and also in the Yuribey, Messoyakha and many otherrivers) also characteristic are regular migrations resulting as a response to dangers ofwinterkill. Winterkill is controlled directly by the marshiness of the area along thelower parts of the rivers, and by the general abundance of marshes, which provokesan oxygen deficiency. This phenomenon is particularly severe in winter since the iceprevents exchange of gases between the atmosphere and the water surfaces.

Various species of fish react differently to insufficient oxygen. Thus at watertemperatures near 0°C the normal oxygen content is about 14 mg/1. A decrease inoxygen levels to 7—7.5 mg/1 provokes depressed respiration among some fish, whileto survive they require at least 3.5 mg/1. I have cited some examples (Table 1).

Naturally an oxygen deficiency provokes in the fish an attempt to leave thezones of winterkill hazard. Hence muksun, Cisco, peled, young sturgeon, nelma(inconnu) and some other species commonly move out from the rivers into the baysand gulfs. The degree of impact of oxygen deficiency makes itself felt more stronglywith decreasing depth and rising water temperatures. In particular, in summer, witha rise in water temperature fish begin to move out of the floodplain lakes, i.e., fromlocations with an abundance of food, into areas less well provided with food.

It is also essential to note that although Baydaratskaya, Ob', Gyda and Tazbays and Yenisey Gulf are areas of intensive fishing, the questions of fish speciescomposition, seasonal migration and their ecological peculiarities are little studied.This also applies in like degree to the fish occupying the rivers and lakes and theinshore waters of the Kara Sea. The characteristics of the fish of the Yamal-Gydanarea are listed below.

Of the lampreys (Lampetra), the arctic lamprey {Lampetra faponica) and the

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TABLE 1

Reaction of Various Species of Fish to Varying OxygenContents of Water (mg/1) at a Water Temperature

of 0.2°-0.3°C [9]

Fish species

SterletNelmaMuksunPeledPerchIdePikeCrucian carp

Beginning of depressedrespiration

7.5-77-6

4.5-44-3.54-34-33-22-1

Survivallimit

3.54 . 5 ^

2-1.51.5-11.1-0.6

0.50.6-0.3

0.1

arctic brook lamprey (L. Kessleri) occur. According to L. S. Berg [2] the latter is asmall riverine form of the arctic lamprey, with a distribution from the Pechora tothe Anadyr'. Ye. V. Burmakin [5] reports that the only specimen of a lampreycollected at Cape Drovyanoy was a young individual 171 mm long and weighing 5g. N. A. Valikov [23] reported both subspecies from the southern part of Ob' Bayand also from Taz Bay and Yenisey Gulf. P. A. Dryagin [7] points out that adiadromous form, 43-48 cm long and weighing 100—200 g is envountered as far asNovyy Port, the mouths of the Yada, Napalkov, Yaptiksale and Tambey rivers andKotel'nikovo. It has been reported from Taz Bay by A. A. Pnev [14]. Apart fromthe arctic lamprey many authors also refer to a small riverine form. The questionsof the systematic position of the lampreys in the Yamal-Gydan area, especially ofthe diadromous form, require special investigation. Lamprey larvae (of whichspecies?) are extremely numerous in the northern parts of the Ob' and Gyda baysand in Yenisey Gulf.

Two species of sturgeon (Acipenseridae) have been recorded: the sterlet(Acipenser ruthenus) and the Siberian sturgeon (A. baeri). In practice the sterletmust be considered as a representative of more southerly latitudes, and the rareappearances of isolated individuals in Ob' Bay (Tambey River) and in Yenisey Gulf(at the latitude of the mouths of the Dudinka and Igarka), as accidental visits.

The Siberian sturgeon is a common fish in all the bays and inlets, and isencountered as far as the shores of the Kara Sea, but it is mainly immatureindividuals which frequent the most northerly areas. Sturgeon are also found asvisitors in rivers and lakes. As one moves north the numbers of sturgeon decreasemarkedly. Large sturgeon are caught mainly in the southern parts of Ob' Bay andYenisey Gulf. The once rich stocks of sturgeon are almost completely exhausted atthe present time. The sturgeon fishery suffered particularly badly during the War.At the present time small, dead sturgeon, killed by water pollution, are frequentlyfound in the vicinity of Cape Kamennyy on the shores of Ob' Bay.

Of the herring family (Clupeidae) two subspecies are found in the Yamal-

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Gydan area. In 1936 and 1937 Pacific herring (Clupea harengus pallasi) were foundin the vicinity of Cape Drovyanoy in Ob' Bay [8]. The systematics of minortaxonomic divisions at the level of the subspecies are far from clear and requirestudy, the more so since the Pacific herring is not found east of the Ob' as far asthe Lena. The stocks of this herring in Ob' Bay have not been determined, and thetotal number of individuals caught so far does not exceed a few dozen. Sincethere was a female with ripe eggs taken among the herring caught, the hypothesishas been proposed that this herring spawns nearby, and its appearance in Ob' Bay islooked upon as a consequence of the warming of the Arctic [2].

The second subspecies, the Atlantic herring (Clupea harengus harengus) is not atall typical of the Kara Sea. Nevertheless odd specimens of the Atlantic herring areencountered among catches of cisco in Omulevaya Bay (Yenisey Gulf)-

Of the salmon family (Salmonidae), five genera are recorded from here: salmon(Salmo), char (Salvelinus), taimen (Hucho), nelma (inconnu) (Stenodus), andwhitefish (Coregonus). They are all of great value in terms of fisheries, but thesalmon and taimen are not of commerical significance due to their limited numbers.The char plays a relatively minor role in terms of the fisheries. The nelma andparticularly the whitefish species comprise the bulk of the catch.

An Atlantic salmon (Salmo salar) was found in 1932 by A. N. Probatov [16],west of the Kara River. According to questionnaire data, odd specimens visit therivers and lakes of Western Yamal from Baydaratskaya Bay [29]. This matter needsto be verified. The same may be said about the taimen (Hucho taimen), which isoccasionally encountered in the southern part of Yamal, especially in the Sh-chuch'ya and in some lakes. One specimen was acquired in a salted condition by theAANII expedition in 1971 at Neyto Lake. Taimen are occasionally reported in thesouthern part of Yenisey Gulf.

Char and palii (Salvelinus alpinus and subspecies) represent a very complextaxonomic group and their systematics have not been worked out at all. Apart fromthe anadromous char, which annually run down to the sea, there exist lake-and-riverforms and even purely lake forms (in landlocked bodies of water), which are calledpalii. Both are found in the basins of the large Siberian rivers. Anadromous char andpalii may be either large or relatively small. Anadromous char make runs down tothe sea, where they disappear beneath the ice, but every year they return to therivers before freeze-up. Mature fish run upstream to the spawning grounds. Havingspawned they either remain for the winter in the river, where they select deep holes,or else move out to deep lakes. As a rule the largest forms occupy the large rivers.Mature individuals do not spawn every year. On their return to the rivers young andimmature fish often remain in the lowest sections of the rivers near their mouths.

Large palii, which spawn in the rivers, occupy deep lakes. Finally palii fromsmall lakes without any outlet spend their entire lives in the same body of waterand are of small dimensions. Palii in breeding coloration are usually more brightlycolored than the anadromous char. Anadromous char are known in Baydaratskayaand Gyda bays and the northern part of Ob' Bay, in Munga Bay and Yenisey Gulf,and also in the Seyakha, Nurmayakha, Yuribey and several other rivers [2, 5, 24,27]; the same applies to the landlocked variant. On the Yamal Peninsula the fisheryis for local consumption, but on the Gydan Peninsula and in Yenisey Gulf it is ofminor commerical significance.

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The nelma (Stenodus leuckhthus nelma) is a very valuable and fairly commonfish in Ob', Taz and Gyda bays and in Yenisey Gulf. Its main range coincides withthe Ob'-Irtysh basin (from 48°N to Cape Drovyanoy and Shokal'skiy Island).Farther north it is encountered in Yenisey Gulf. The nelma is widely fished in Ob'Bay at Novyy Port, Cape Munga, Cape Kamennyy, Cape Taran, Cape Trekh-bugornyy, Cape Drovyanoy, and in the Seyakha and Tambey rivers. It runs up therivers on the west coast of the Yamal Peninsual from Baydaratskaya Bay reachingsome lakes.

At the present time the fishery in Ob' Bay has declined drastically, and ittypically is based upon immature individuals. In Gyda Bay the nelma are somewhatlarger, but here too it is mainly young fish which are being caught. It is alsoencountered in the rivers of the Gydan Peninsula, the Yuribey, the Ayvayakha, theGnezu, the Gyda, the Nyauyakha, and the Nadayakha, and also in lakes Khaseyntoand Yambuto. The nelma run up the rivers in the spring, and leave them again priorto freeze-up. A small number of fish stay all winter in the southern part of YeniseyGulf. The largest individual was caught in 1936 and weighed 16.2 kg, for a length of103 cm [4]. Nelma run up several rivers from Taz Bay, including the Taz and the Pur.

The Siberian cisco (Coregonus sardinella) belongs to a group of widelydistributed whitefish which inhabit the rivers and lakes of the area. In the northernparts of its range the cisco does not shun saline inshore marine waters, frequentinginlets and bays. These fish, the systematics of which have not yet been properlyworked out, should correctly be divided into separate populations. As a rule thecisco of the northern populations are larger than the southern ones. In the case ofthe northern populations the feeding migrations are made in salt water, maturityoccurs later, the fish live longer, and they winter in rivers (in deep holes and pits)with depths of 5—6 m or more, close to their spawning grounds.

The tugun (Soregonus tuguri) is not typical of the Ob' and Gyda bays. Isolatedindividuals have been taken in the southern part of Yenisey Gulf [3]. Occasionallytugun are encountered in the Shchuch'ya, Taz and Pur rivers [7].

The omul {Coregonus autumnalis) is one of the main objects of the fishery inthe northern parts of Ob' and Gyda bays and also in Yenisey Gulf. The omul runsup many of the rivers of northern Yamal: the Seyakha, Tambey, Yagodyyakha,Khobeyyakha, etc. It is common in the inshore waters around Shokal'skiy andOlen'iy islands, in Baydaratskaya Bay and in Malygin Strait. On the GydanPeninsula the omul has been recorded in the lowest reaches of the Nyauyakha,Yuribey and Gyda rivers, while along the northwest shore of Yenisey Gulf it isparticularly numerous in the mouths and lower reaches of the Mongocheyakha(Sosnovaya), Omulevaya and Nyarmkhoyyakha rivers. According to the findings ofthe AANII expedition in 1972, the omul run begins here on July 20 and isparticularly intense in the second half of August. According to the fishermen, theomul stays here throughout the winter. In former years ice fishing for omul wascarried out at the mouths of the Omulevaya and Mongocheyakha rivers. In thesouthern part of Ob' Bay isolated individuals range as far south as Novyy Port.There are no omul in the Ob', but in the Yenisey they penetrate far upstream tothe South. They apparently spawn in the Khobeyyakha and Tambey rivers, sincethey run upstream for 40—50 km in these rivers [15]. According to V. I. Dmitriyev[6], the spawning run in Yenisey Gulf occurs particularly intensively from September5 to 15.

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The peled {Coregonus peled) is basically widely distributed in the rivers andlakes of the Yamal, Taz and Gydan peninsulas. Apart from the lake-and-river andlake populations, there also exist populations which run down into the bays andinlets, including saltwater areas. In the spring many peled move from Ob' Bay intothe flood-plain lakes of the Ob' to feed.

The chir or broad whitefish {Coregonus nasus) is a valuable food fish whichoccupies most of the rivers and bodies of water of the Yamal-Gydan tundra andalso the Ob', Taz and Gyda bays and Yenisey Gulf. It concentrates in the largestnumbers in the southern and central parts of Ob' and Taz bays [7]. In Ob' Bay itgoes as far north as Cape Drovyanoy. On the Gydan Peninsula the chir is to befound in the following rivers: the Yuribey, Aybayakha, Gnezyu, Nyaruyakha, Gyda,Gadasyo, and in lakes Khaseynto, Yambuto and some other bodies of water [5]. In1972 members of the AANII expedition caught chir in the Yambuto-Neyto groupof lakes and in the mouth of the Nurmayakha River. The largest male, at a lengthof 570 mm, weighed 3,212 g. Ye. V. Burmakin [5] quotes the maximum weight fora chir from the Gydan Peninsula as 3,060 g. A. I. Dmitriyev [6] notes that on theYenisey at the latitude of the Brekhovskiye Islands the chir is a common commercialfish. In the case of the Ust'-Port fisheries chir comprised 9.1% of the catch (in the1938 summer fishery).

The Siberian whitefish or pizhyan {Coregonus lavaretus pidschiari) is a commonfish in the Yamal-Gydan area. In the case of the rivers and lakes of the YamalPeninsula it is cited by Zhitkov [29] as being one of the main objects of localfishing by the reindeer herders. P. A. Dryagin [7], in indicating the centers ofdistribution of this whitefish, includes the southern and northern parts of Ob' Bayand also Taz and Gyda bays. The local population in the southern part of Ob' Bayis particularly large; from here the whitefish make a feeding migration up the Ob',and in part, they spawn in tributaries of the Ob'. The local population from thecentral and northern parts of Ob' Bay (estuarine whitefish) fatten and spawn ininflowing rivers and also in the bays and inlets. The whitefish of Taz Bay run upthe Taz, Pur, Messoyakha, Payutayakha and Poylovayakha to spawn. According toYe. V. Burmakin [4] the whitefish of Gyda Bay are encountered in the Yuribey,Gnezyu, Nyauyakha, Gadasyo and Nadayakha rivers and in lakes Khaseynto andYambuto.

In spring the pizhyan enter the rivers; in summer some of them remain in thelakes. Fish running downstream are often encountered in the lower reaches of therivers in the second half of October and in November. Some individuals spend theentire winter in the bays. Members of the AANII expedition in 1972 caughtwhitefish in the mouth of the Nyarmkhoyyakha and in Lake Yaruto. On theYenisey the best feeding areas are the left bank tributaries and the channelsbetween the low, alluvial islands. The shallow depths, slow current velocities and therapid warming of the water have a favorable influence on the development ofbenthos and plankton. Hence it is no accident that the waters surrounding theBrekhovskiye Islands are the best for the fattening migrations of fish of the genusCoregonus. The catching of whitefish is mainly limited to the summer fishery, andin different years may account for 20 to 37% of the total catch.

The muksun (Coregonus muksuri) is a common commercial fish in Ob', Taz andGyda bays and in Yenisey Gulf. In Ob' Bay the major concentration of muksunoccurs in the southern part, whereas to the north, in the area of Cape Drovyanoy,

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concentrations drop appreciably. The muksun occupies many lakes and rivers onthe Yamal Peninsula, including those debouching into Baydaratskaya Bay. On theGydan Peninsula it is encountered in the Yuribey, Gyda, Ayvayakha, Nyarmkh-oyyakha and other rivers, and also in lakes Khazeynto, Yambuto and Yaruto. FromTaz Bay the muksun run up the Taz, Pur and Aderpayuta rivers. In Yenisey Gulfthe main population of muksun sticks to the area of the Brekhovskiye Islands andfarther north.

The Salmonidae represent the main species fished in the Yamal-Gydan area.

The grayling (Thymallus arcticus) occupies the rivers and lakes of the Yamaland Gydan peninsulas and is also encountered in Baydaratskaya, Ob', Taz and Gydabays and in Yenisey Gulf. On the east coast of the Yamal Peninsula the graylingextends north to the Tambey River and on the west coast to north of theMordyyakha. It was once caught in a bay at Cape Drovyanoy [29]. On the GydanPeninsula it is encountered in many of the rivers in the system of tributaries andlakes involving the Yuribey, Gyda and Aybayakha. It spawns in Lake Khaseyntoand is common in Lake Yambuto. In 1972 it was caught in the mouth of theNyarmkhoyyakha River and in Lake Yarunto. The absolute dimensions of maleswere from 348 to 427 mm, with weights from 406 to 554 g, while females were338 to 403 mm in length and weighed 360 to 614 g. Grayling are encountered insmall numbers in the rivers of the Taz Bay and Yenisey Gulf catchment basins.

The Asiatic (or arctic) smelt (Osmerus eperlanus) inhabits Ob', Taz and Gydabays and Yenisey Gulf. It enters many of the rivers, apparently to spawn, althoughit also lays its eggs in the bays and inlets. The stocks of smelts are fairlyencouraging, but the fishery has begun to develop only relatively recently. Accord-ing to P. A. Dryagin [7], 10,500 centners of smelt were caught from Novyy Port in1938-1943. In general no more than 15-20% of the stocks are utilized.

The pike (Esox lucius) is one of the most numerous fish of both large andsmall bodies of fresh water in the Yamal-Gydan area. It is encountered in smallnumbers in the bays and gulfs, mainly in the southern parts. From the data of theAANII expeditions of 1971 and 1972 pike live in all the rivers and lakes of thesouthern tundra subzones, whereas there are few of them in the northern subzones,and it is totally absent from bodies of water on the islands.

The Siberian roach (Rutilus rutilus) and the Siberian dace (Leuciscus leuciscus)are found in small numbers in the bodies of water of the region, and areencountered, although only as solitary individuals, in the southern parts of Ob' andTaz bays and in Yenisey Gulf [2, 7 ] .

The ide {Leuciscus idus) is rare. It is distributed only in the southern andcentral parts of Ob' Bay from Novyy Port and Cape Munga to the Seyakha River.Isolated individuals are encountered in Taz and Gyda bays, Yenisey Gulf and in thechannels at the latitude of the Brekhovskiye Islands [5, 7] . As a rule the ide in thebodies of water at these northern latitudes are small and are of no significance interms of fishing.

The lake minnow (Phoxinum percnurus) and Chekanov's minnow (Phoxinusczekanowskii) penetrate into the bodies of water of the southern margins of thearea, particularly in the basins of the Shchuch'ye, Pur and Taz rivers, and

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apparently also the Yenisey [2, 7] . The former prefers small lakes, the latter is alsoencountered in river channels.

Of the family Cobitidae the Siberian stone loach (Nemachilis borbatulus)inhabits the southern parts of Ob' Bay and Yenisey Gulf. Apparently it alsooccupies the lowest reaches of the Taz and the Pur.

The perch (Perca fluviatilis) is occasionally encountered in the southern parts ofOb' and Taz bays and in Yenisey Gulf [5, 7] . The perch is of no significance interms of fishing.

The ruffe (Acerina cernua) is common in the fresh-water areas of the baysalthough it is not completely absent from the saltier areas. Its numbers decreaseconsiderably toward the north. The ruffe is rare in Gyda Bay but is caught in theYuribey River and in the Yuntose channel of the Gyda [4]. It is typical of manyrivers on the Yamal Peninsula and is encountered in Ob' Bay. It is particularlynumerous in the Taz.

In 1935 approximately 4,000 centners of dead fish were found on a beach atCape Parusnyy in a strip about 10 km long and 10 m wide; in some places the layerof fish reached 20 cm in depth [7]. In Yenisey Gulf the ruffe is common at thelatitude of the Brekhovskiye Islands. It is of no significance in terms of fisheries inthe area.

The burbot (Lota lota) is widely distributed in the rivers and lakes of theYamal and Gydan peninsulas, and is also encountered in the many bays and gulfs.In Ob' Bay the burbot is typical of the southern and central parts, and isencountered in rivers and lakes as far as the Tambey River. The burbot is equallycommon in Taz and Gyda bays. On the Gydan Peninsula it is caught in particular inthe Yuribey, Aybayyakha, Gnezyu, Nuyakha and Gyda rivers and in Lake Yambutoand others [4]. The burbot is taken in the southern parts of Yenisey Gulf andcomprises up to 15% of the total catch [6].

The navaga (Eleginus navaga) is typical of the waters around the northerncoasts of the Yamal and Gydan peninsulas. In Baydaratskaya Bay the navaga hasbeen recorded south of the mouth of the Yuribey, where particularly largespecimens have been caught. Thus in 1946 two females were caught there, 441 and419 mm in length. They weighed 703 and 713 g [2]. The navaga is fairly commonin Gyda Bay and is also encountered around the coasts of Shokal'skiy, Oleniy andBelyy Islands. In the late fall it commonly enters the mouths of rivers on the floodtide, then runs back out again on the ebb. The navaga at present is of nocommercial significance.

The Arctic cod (Boreogadus saida) is a fish which concentrates in large schools,occurring in the waters of the Kara Sea and in the northern parts of the bays andgulfs adjacent to the Yamal and Gydan peninsulas. The Arctic cod prefers salt waterareas. During their mass runs catching them is quite easy. The question oforganizing a fishery depends upon the possibilities of using the fish. In terms ofquality it is a low-grade fish, but quite edible, and with appropriate processing andpreserving techniques it could serve as a cheap food for animal farms, zoos,dog-rearing farms, poultry farms and pig farms. The Arctic cod is a food source formany fish, mammals and birds, which not only catch it in the water, but also whereit has accumulated on shore, thrown up in large numbers during storms.

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The Arctic flounder (Liopsetta glacialis) is encountered along the coasts of thebays and gulfs. In the summer of 1942, 59 centners were caught in the northernpart of Ob' Bay, incidental to other species, and 39 centners in 1943 [7]. It iscommon around Cape Drovyanoy, the mouth of the Tambey, the shores ofShokal'skiy Island, and in Yenisey Gulf [5]. The flounder is of no commercialsignificance here.

The nine-spined stickleback (Pungitius pungitius) is among those fish which,apart from fresh bodies of water including small lakes with no discharge, can easilylive in bodies of water with a fairly high salinity. In practice it may be encounteredanywhere.

One specimen of a fish which is quite rare in the Yamal-Gydan area, thelumpsucker (Cyclopterus lumpus), was caught at Cape Drovyanoy [5].

More common along the coasts of the Yamal and Gydan peninsulas isDeryugin's lumpsucker (Eumicrotremus derjugini), which is related to the Atlanticspiny lumpsucker [1].

Several species of sculpins inhabit the waters of the Kara Sea adjacent to theislands and to the coasts of Yamal and Gydan peninsulas; they are encountered onlyin small numbers or singly. An exception is the Arctic fourhorn sculpin (Myoxo-cephalus quadricornis) which is resident in large numbers in the salt water of thebays and gulfs. In the southern part of Ob' Bay the sculpin penetrates as far asNapalkova. In the northern areas it causes great trouble to fishermen in clearingtheir nets, especially small-mesh nets. The sculpin often runs into the lower reachesof rivers on the flood tide.

Apart from this the sharp-nosed sculpin (Triclops pingeli) is caught off BelyyIsland. According to Yesipov the spatulate sculpin (Icelus spatula) and the twohornsculpin (/. bicornis) are encountered fairly commonly in the Kara Sea. The Arcticstaghorn sculpin (Gymnocampus tricuspis) has been observed in Ob' Bay and inYenisey Gulf. Possibly several other species may be found in inshore areas, which sofar have been found only in the northern parts of the Kara Sea at relatively greatdepths. In completing this survey of the fishes of the bodies of water of theYamal-Gydan area one should mention two more freshwater sculpins: the Europeansculpin (Cottus gobio), which is encountered in the south of the Yamal Peninsula,and the Siberian sculpin (C. sibiricus) from Taz Bay and the southern part ofYenisey Gulf. Both species are also found in salt water to some extent, but do notgo very far north.

In conclusion one should note that more than 40 species of fish have beenrecorded in the waters of the Yamal and Gydan peninsulas and in the adjacentwaters of the Kara Sea. In this connection one should realize that due to theincomplete nature of the systematics and to differences in interpretation oftaxonomic classes, an accurate count of species is very difficult. Thus, one mayassume that at the present time in the area under study the total number of speciesof fish is 41—43, belonging to 15 families and 29 genera. Apart from this, oneshould also take into consideration that in the future this number may be increasedby the addition of rare or accidental visitors, mainly from the north, since theoffshore waters are inadequately investigated.

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Birds

Birds represent the largest group of vertebrates. Of the 26 orders of birdsrecorded in the USSR, 12 occur in the Yamal-Gydan tundra. Of these, repre-sentatives of four orders do not nest within the area: alcids, woodpeckers, cranesand procellariiformes. As regards the procellariiformes and the alcids, this is to beexplained by the lack of suitable nesting biotopes appropriate to the birds of theseorders. As a result of the total lack of rock outcrops, birds which nest in cliffcolonies do not nest here at all. This applies not only to the alcids andprocellariiformes but also to some of the gulls, and especially the kittiwake.Glaucous gulls and herring gulls, which live parasitically on cliff colonies, are alsolimited in their nesting numbers. The limited amount of rock-and-gravel tundraexplains the sporadic distribution of dotterels. Cranes are only visitors to the tundrazone, nesting much farther south. Visits by woodpeckers are explained by the factthat they may be able to find food here while in transit.

With regard to species composition the passerines predominate in terms ofnumbers. But they occupy mainly the most southerly margins of the area. Next interms of numbers of species come the Charadriiformes (shore birds), Anseriformes(geese), gulls and raptors. The remaining orders are limited to only a few species.

The order Columbidae is represented by three species in the Yamal-Gydantundra: the red-throated diver (Gavia stellata), the black-throated diver (G. arctica)and the white-billed diver (G. adamsii). The best-adapted of these is the red-throateddiver, which occupies the entire tundra area, including the offshore islands. Theblack-throated diver nests up to the Kara Sea coast but is not numerous here. Thenesting range of the white-billed diver coincides with the forest-tundra and thesouthern subzone of the tundra. All these divers nest close to water and feed mainlyon mollusks and fish. They arrive late, after the rivers have broken up and after thelake ice has melted away from the shores. The red-throated diver makes use of thesea more frequently than the other species in procuring food. Indeed, in areas witha limited food supply, e.g., on islands, the red-throated diver derives its food mainlyfrom the sea. The white-billed diver is generally rare in the northern tundrasubzone, preferring to occupy large lakes.

Procellariiformes (Fulmarus). The only representative of this order, the fulmar(Fulmarus glacialis) is encountered in the northern parts of the bays and in YeniseyGulf while on passage. The lighter form predominates.

Alcids (Alcae). There are no nesting species among the alcids, although twospecies have been recorded as transients in the northern parts of Ob' Bay andYenisey Gulf. These are the black guillemot (Cepphus grylle) and Briinnich'sguillemot (Una lomvia). They are observed more frequently out over the Kara Sea,beyond the offshore islands.

Gulls (Lari). This order includes three families: jaegers (Stercoraridae), gulls(Laridae) and terns (Sternidae), although some authors combine all three into onefamily, Laridae. Within the first of the three families, three species are found in theYamal-Gydan tundra: the pomarine jaeger (Stercorarius pomarinus), the parasiticjaeger (5. parasiticus), and the long-tailed jaeger (S. longicaudus). The ranges of all

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three species practically coincide within the area. They nest both on the mainlandtundra and on the offshore islands, but numerically the pomarine jaeger pre-dominates on the islands. It also frequents the coast more commonly. Thelong-tailed jaeger penetrates somewhat farther south than the others. When rodentnumbers peak on the tundra the number of nesting pairs rises sharply. With lowrodent numbers the number of surviving jaeger nestlings drops sharply, or else theadults simply do not begin nesting at all. Jaegers do not catch fish for themselves,but steal them from other birds. They frequently eat carrion, including dead fish.Apart from rodents, jaegers also catch small birds for food, usually passerines andshorebirds, destroying their nests and eating nestlings and eggs. Jaegers take theplace of the raptors on the tundra, the latter being quite limited in numbers; jaegersbehave extremely aggressively and provoke retaliatory action even from smallnesting birds.

Of the true gull family (Laridae), six species have been recorded. Three of themhave been noted in passage, or on migration, one only sporadically, and threenesting regularly; these last occur in quite small numbers. The ivory gull (Pagophiluseburnea) visits the coasts only very rarely in spring, while solitary Sabine's gulls(Xema sabini) may be encountered on the tundra in the interior. The situation withregard to the kittiwake (Rissa tridactyla) is somewhat different. It does not nest inthis area, either, but is fairly common on migration in the northern parts of Ob'and Gyda bays, in Yenisey Gulf and also in the adjacent parts of the Kara Sea. It•frequently accompanies ships, picking up galley refuse; otherwise they catch arcticcod for food. Glaucous gulls (Larus hyperboreus) nest mainly in the northerntundra subzone along the sea coast; they do not form large colonies. Isolated pairsnest on headlands, on island beaches, and on lakes. Exactly the same applies to theherring gull (L. argentatus). They also nest on islands, but they are also found inthe southern tundra subzone. Neither species of gull is particularly numerous.

The common gull (L. canus) is the third species of gull which nests sporadicallyin this area, although it nests in mass colonies farther south. However, isolated pairsnest and raise their young on the southern part of Yenisey Gulf, in the area of theBrekhovskiye Islands, and have been recorded as visitors on Taz and Ob' bays.Apart from these, Ross's gull (Rhodostethia rosed) is encountered on passage in thenorthern parts of the Kara Sea in the fall.

The representative of the third family in the Yamal-Gydan tundra—the Arctictern {Sterna paradisea)—nests throughout the area on coastal beaches and along theshores of rivers and lakes. Isolated pairs also nest on the islands.

After the passerines the shorebirds (Limicolae) represent one of the mostnumerous orders of birds inhabiting the Yamal-Gydan tundra in terms of number ofspecies. They occupy varied biotopes: marshes, river banks, lakes and the sea coast.Their nesting ranges encompass both open areas and shrub-grown terrain. Theyrepresent the most widely distributed and most varied of the arctic fauna, enliveningthe landscape of this severe environment, especially in spring. As distinct from thepasserines, the shorebirds also occupy the northern subzone; many of them nest onthe islands.

The plover family (Charadriidae) occupies an important place among theshorebirds, amounting to two genera and five species. The largest within this family(by size), the black-bellied plover (Squatarola squatarola), is allocated a separate

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genus of its own; it occupies the entire area of the mainland tundra and also nestsin small numbers on the islands. The European golden plover (Chaadrius apricarius)and the American golden plover (Ch. dominicus) are widely distributed throughoutthe tundra. These are closely related species, but while the former is numericallypredominant on the Yamal Peninsula and farther east (although not beyond theKhatanga), the latter is restricted to Yamal as the western limit of its range. TheEuropean golden plover prefers drier habitats than the American golden plover andis not found in the northern tundra subzone of the Yamal Peninsula, although onthe Gydan Peninsula it nests right to the sea coast.

Another two shore birds which nest on the tundra of the Yamal and Gydanpeninsulas also belong to this same genus. These are the dotterel (Ch. morinellus)and the ringed plover (Ch. hiaticula). The former is encountered only sporadicallyon the tundra of the Yamal-Gydan area, as in many other areas; the northern limitof its range is controlled not only by high latitude but also by the lack of suitablenesting habitat. Observations by B. M. Zhitkov [29], who encountered dotterelsgiving their mating call on the Kara Sea coast in the vicinity of the Mordyyakha onMay 31, and also by V. A. Gayev, who recorded a pair of these birds at LakeTaureto on June, 1972 (on the AANII expedition), leave no doubt that the dottereloccasionally nests on the Yamal Peninsula. In 1972 downy nestlings were also foundby V. A. Gayev on the shore of the Nyarmkhoyyakha River on Gydan Peninsula.Particularly large numbers of dotterels were observed here from August 24 toSeptember 10. Ringed plovers are not very numerous but nest regularly in the area,right to the northern coasts, and have been recorded on Belyy Island on passage.

The genus Arenaria is represented by a species which is extremely characteristicof the Yamal and Gydan tundras, the turnstone (Arenaria interpres): it nests rightto the Kara Sea and is also found on Belyy Island. The turnstone nests relativelyrarely on dry, lightly vegetated tundra; it much prefers sand and gravel beachesalong the shores of rivers and lakes, and also spits along the sea coast. This noisy,energetic little bird plays a major role in the life of its ecosystem, actively involvingitself in every event relating to the defense of nests, and warning as to the approachof enemies, including man.

Of the sandpiper genus (Calidris), eight species have been recorded. The curlewsandpiper (C. testacea), evidently a rare nesting species, was encountered by S. P.Naumov [10] in the eastern part of Gydan Peninsula. The curlew sandpiper wasfound on Taz Peninsula at Cape Kruglyy. This is the western limit of sightings. Thedunlin (C. alpina) is an extremely common bird on the Yamal-Gydan tundra,nesting as far as the Kara Sea littoral, but it is not very common in the northernsubzone, although it has been reported from the islands during the nesting season.The little stint (C. minuta) is an extremely characteristic and numerous species,nesting throughout the entire region, including the islands. Temminck's stint (C.temminckii) is a close relative of the little stint, but nests in more varied habitatsand prefers the southern tundra subzone. It was encountered repeatedly in the areaof Cape Leskin but no nesting has been confirmed as far north as this. The knot (C.canutus) was observed on Belyy Island during fall migrations. Three specimens werecollected but no nesting was confirmed. Two pectoral sandpipers (C. melanotos)were collected at Cape Leskin by the AANII expedition in 1972, evidently inpassage. The spotted redshank (Tringa erythropus) is also encountered in passage.Two sanderlings (Crocethia alba) were collected on Belyy Island on fall migration inSeptember 1936.

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The ruff (Philomachus pugnax) is a common nesting species on the Yamal-Gydan tundra, and has been recorded by all investigators, although it is lessnumerous in the north. Ruffs have not been observed on Belyy Island. Extremelycharacteristic of the Yamal-Gydan tundra are the two phalaropes; the red-necked(Phalaropus lobatus) and the grey phalarope (Ph. fulicarius). The former is moretypical of the southern subzone, although it does also nest on the islands. Inparticular, a brood of four downy young was found, along with a male adult, onJuly 31, 1936 [22]. The red-necked phalarope also nests right to the Kara Sea onthe Gydan Peninsula. As a rule the grey phalarope is not encountered on the YamalPeninsula during the nesting season, and only occasionally it is observed along theKara Sea coast; but on the Gydan Peninsula it nests regularly in small numbers inthe northeast. Typical of the grey phalarope is the fact that after the nesting periodthey fly out to the sea coast and roam along the coast in large flocks, consisting ofmales and young; the females migrate south earlier.

The pintail snipe (Gallinago stenura) is a bird of the forest tundra, and it nestssporadically on the Yamal and Gydan peninsulas only in marshes in the southerntundra subzone. The great snipe (G. media) may be observed even more rarely onthe left bank of the Yenisey in the southern tundra zone; it is encountered here inisolated pairs and may possibly nest. The bar-tailed godwit (Limosa lapponicd) nestsoccasionally in the southern tundra subzone in isolated pairs.

Anseres. This order consists of one family (Anatidae) which is divided into twosubfamilies: the geese (Anserinae), which includes all the swans and geese, and theducks (Anatinae), which includes all. the remaining birds of this order.

Only two species of the swan genus (Cygnus) have been recorded: the whooperswan (C. cygnus) which visits the southern part of Ob' Bay, and Bewick's swan (C.bewickii), which nests regularly throughout the entire area, with the exception ofthe northern offshore islands. The latter is a fairly common species, preferring largelakes. Of the geese the white-fronted goose (Anser albifrons) and the bean goose (Afabalis) occur in small numbers both on the nesting grounds and while moulting.The former exceeds the bean goose in numbers in the northern region, including theislands. The latter sticks to the vicinity of large lakes and ranges farther south thanthe white-fronted. The lesser white-fronted goose (A. crythropus) is generally notnumerous on the Yamal-Gydan tundra; it does not occur along the northern KaraSea coast, nor on the islands.

Two species of the genus Branta have been recorded from the Yamal-Gydantundra: the brant (Branta bernicla) and the red-breasted goose (B. ruficollis). Thefirst of these is common along the Kara Sea coast and in the northern parts of thebays and gulfs, and also on the offshore islands. They nest in isolated pairs on riversand lakes. They are also encountered during the moult. The red-breasted gooseinhabits mainly the Gydan tundra, while on the Yamal tundra it is restricted to theshores of Ob' Bay where it nests in the valleys of small streams, preferring driersites. The red-breasted goose is endemic. Its nesting range is sporadic and although itextends east to the western shores of Lake Taymyr, this goose has been severelyreduced in numbers. The killing of red-breasted geese is forbidden, and they arenow protected. This is a very placid goose and not at all timid; it always remains indensely-packed flocks and hence suffers particularly heavily from man's activities; itis also poorly adapted to combat predation by either birds or mammals.

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The genus of river ducks (Anas) is represented by four species. Most typical ofthese is the pintail (A. acutd). It nests regularly in the southern tundra subzone, andoccasionally in the northern subzone. It is not found on the islands, but wasencountered repeatedly in the northern parts of the bays and gulfs. The teal (A.crecca) and the widgeon (A. penelope) nest in small numbers in the southernsubzone; isolated nests or broods were found as far north as 71° [29]. Baikal teal(Anser formosum) have been observed only in passage. The red-breasted merganser(Mergus senator), which belongs to the diving ducks, nests in limited numbers onthe Yamal-Gydan tundra. A nest with a fertilized clutch was found at LakeTaureto. It was observed frequently on the Nurmayakha River, at Cape Leskin onthe Gydan Peninsula, and at the mouth of the Salyyadayakha (AANII expeditionsof 1971 and 1972).

In the southern tundra of both the Yamal and Gydan peninsulas there nestisolated pairs of ducks whose main ranges are located in the forest tundra and thenorthern tayga. These include the tufted duck (Ayihya fuligula), the greater scaup(Aythya marila), the common scoter (Melanitta nigra) and the velvet scoter (M.fused). They are atypical of the tundra zone, but are recorded regularly as visitorsalong its southern borders, and in isolated cases occur as nesting species within theregion [17].

The long-tailed duck (Clangula hyemalis) occupies a somewhat different posi-tion. Its normal nesting range is located in the southern tundra and forest tundra,but it also extends into the northern (tundra) subzone, including the offshoreislands [19]. After the breeding season the main mass of males fly out to the shoresof the Kara Sea where they moult in the northern parts of the bays and gulfs andalong the sea coast, then head west along the coast.

One visit by a harlequin duck (C. histrionicus) has been recorded in thenorthern part of Yenisey Gulf [13].

Among the eiders the king eider (Somateria spectabilis) represents a nestingseries which is even more numerous than the long-tailed duck. Large flocks ofnonbreeding and moulting birds permanently frequent the northern coasts of theYamal and Gydan peninsulas. The main nesting range of the king eider is located onislands along the northern coasts of these peninsulas; it nests less frequently alongthe shores of rivers and lakes. The remaining two species of eiders have beenrecorded only as visitors. These include the chance appearance of solitary commoneiders (S. mollissimd) on the Yamal Peninsula and in Yenisey Gulf; this species doesnot usually range as far east as the peninsula. A sighting on Belyy Island must beconsidered as erroneous, since A. N. Tyulin [22] evidently confused common eiderswith king eiders. The Steller's eider (S. stelleri) is a common visitor to the Yamaland Gydan peninsulas.

The order Galliformes is limited to two species of ptarmigan: the willowptarmigan {Lagopus lagopus), which occupies the mainland tundra in its entirety,but does not nest on the offshore islands, and the rock ptarmigan (L. mutus), whichis encountered farther north and greatly exceeds the former numerically. As a rule,the willow ptarmigan is encountered on the offshore islands only as a visitor.Although in the southern tundra subzone of the Yamal and Gydan peninsulas thenumerical superiority of the rock over the willow ptarmigan still obtains, it hasleveled off significantly. In winter both species of ptarmigan migrate south into the

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forest tundra and the northern taiga, where they are of great significance to hunters.Isolated individuals remain for the winter in the southern tundra of the Yamal andGydan peninsulas.

The cranes (Grues) are represented by only two visiting species. Neither thecommon crane (G. grus) nor the Siberian crane (G. leucorgeranus) nests on theYamal or Gydan tundra. Visits to the southern tundra by the Siberian crane areaccidental, evidently from the isolated colonies on the lower reaches of the Ob',whereas the common crane is recorded more frequently, inasmuch as its nestingrange extends down the Ob' as far as the Arctic Circle.

Diurnal raptors (Accipitres) are not numerous on the tundra of the Yamal andGydan peninsulas, either in terms of species or individuals. There are three speciesof the falcon family (Falconidae). The peregrine falcon (Falco peregrinus) occupiesthe entire area right out to the offshore islands. This falcon has been recorded as avisitor on Belyy Island. According to the literature, the gyrfalcon (F. gyrfalco) is aresident of the mainland tundra, the islands and the coast, but unfortunately therehas been no factual data on this bird for several decades now, although the mapscontinue to delimit its range just as before. The third representative of the family,the pigeon-hawk (F. columbaris) nests in the southern tundra subzone. Theremaining three species of raptors belong to the hawk family (Accipitridae): thegoshawk (Accipiter gentilis) has been recorded in passage on the tundra of theGydan Peninsula. The gray sea-eagle (Haliacetus albiciUa) occurs in very smallnumbers, but is encountered throughout the tundra, preferring large lakes and thesea coast. It has been observed several times on Belyy Island. In 1971 these eaglesappeared frequently at Lake Taureto, circling over the lake and were also twicesighted on the cliffed banks of the Nurmayakha River. The eagles often frequentwaterfowl moulting areas, including those of ducks, white-fronted geese and beangeese. The best-adapted species of raptor in the tundra zone is the rough-leggedhawk (Buteo lagopus although its nesting on the latter has not been confirmed. Onthe Yamal Peninusla the rough-legged hawk nests as far as Malygin Strait.

Owls (Striges). This order is represented on the Yamal-Gydan tundra by twospecies: the snowy owl (Nyctea scandiaca) and the short-eared owl (Asio flammeus).The former is a native of the tundra zone and only in years of rodent scarcity doesit also nest in the forest tundra. Its nesting range coincides very closely with theranges of the arctic and Ob' lemmings. The snowy owl is a large and powerful bird.It defends its nest aggressively, putting even foxes to flight. The owls usually do notwinter on the tundra, but migrate south; however, isolated individuals may beencountered here in any month of the year. If there is a food shortage, apart fromlemmings they catch hares and also ptarmigan, ducks, geese and many other birds.The short-eared owl is regularly encountered as a nesting species in the region, butnever in large numbers, and occurs particularly rarely in the northern subzone.

Woodpeckers (Picariae). Representatives of this order have been recorded onlyas visitors. A greater spotted woodpecker (Dendrocopos major) was collected onAugust 9, 1971, among willow bushes by V. A. Gayev (of the AANII expedition)on the Yamal Peninsula, 4 km north of the Nurmayakha River. On August 6 oneindividual was observed at the mouth of that river. From the literature sources thenorthern three-toed woodpecker (Picoides tridactylm) has been encountered on

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Yamal in the region of the Nurmayakha River. Woodpeckers are atypical of thetundra zone but they are able to feed there in summer.

Passerines (Passeriformes). This order includes both birds recorded as nesting onthe Yamal-Gydan tundra and those encountered on migration or as visitors. Themajority of these nest in the southern tundra subzone and are tied to thedistribution of bushy vegetation. Some of the species gravitate to the Yenisey valleyand, as a rule, are encountered on its left bank considerably less frequently than onthe Taymyr coast, but sometimes they penetrate considerably farther north than onthe Yamal Peninsula, indeed right into the tundra zone.

The largest birds of this order, the raven (C. corax) and the hooded crow (C.comix), are encountered here only as visitors. The raven has been observed on thelower Ob' and in southern Yamal. The crow is recorded frequently and farthernorth. On the Yamal Peninsula it was recorded in the area of Lake Taureto and onthe Gydan Peninsula at Lake Yaruto (AANII expedition of 1971 and 1972). Thefield sparrow (Passer montanus) nests as far as the mouth of the Yenisey and insome years even as far as 72°39'N.

The redpoll (Acanthus flammea, A. hornemannii). Both of these variants (thedivision of the redpolls into two separate species is not adequately substantiated)nest on the tundra of the Yamal and Gydan peninsulas, in the very same locations,differing not at all in their ecological peculiarities. They form mixed nesting pairsand during migration form common flocks. On the Yamal Peninsula a nest wasfound in the Yeryakha-Yuribey interfluve, and on Gydan Peninsula, on the shoresof the gulf of the same name. On the Yenisey the redpoll nests to 72°.N, but is notfound on the islands. The least bunting (Emberisa pusilla) nests in the southernsubzone on the Yamal at Cape Yamsale, and as far as 72°N on the Gydan Peninsula.The reed bunting (E. schoeniclus) nests in the southern tundra subzone on the YamalPeninsula and on the lower reaches of the Taz.

Lapland longspur (Calcarius lapponica) and the snowbunting (Plectrophenaznivalis) represent two of the most typical birds of the Yamal-Gydan tundra andnest right to the northern coasts, including the islands. The former prefershummocky, well vegetated tundra, and the latter the sea coast, strewn withdriftwood, or gullied relief with steep slopes. Sometimes they select gaps betweenrocks or abandoned lemming burrows as their nesting sites. Snowbuntings also buildnests in dwellings and other buildings, or in stacks of firewood, and like sparrowsbecome quite accustomed to people, even feeding on garbage dumps. Anotherindigenous species of the tundra zone is the horned lark (Eremophila alpestris)which nests both on the Yamal and on Gydan Peninsula, including the offshoreislands.

The wagtail family (Motacillidae), which includes both pipits and wagtails, isrepresented by six species. Of these, only the red-throated pipit (Anthus cervinus)and the white wagtail (Motacilla alba) penetrate into the northern tundra subzone,but even then only in small numbers. The former has been reported from theGydan Peninsula right to the sea coast, and has been collected on Shokal'skiyIsland, evidently just as a visitor. On the Yamal Peninsula it was observed at CapeKamennyy in considerable numbers and also along the shores of the middle courseof the Shchuch'ye River. According to the data from the AANIPs 1971 expedition,the red-throated pipit is very common at Lake Taureto, where both nests andunfledged nestlings were found. The white wagtail was also recorded from Cape

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Kamennyy on the Yamal Peninsula, where its nests were frequently found; it rangesas far as the Mordyyakha River. On the Gydan Peninusla it nests to the shores ofthe Yuribey and on the Yenisey it is encountered at 73°N.

The Siberian pipit (Anthus gustavi) normally nests in the southern tundrasubzone, but is nowhere numerous, while the meadow pipit (A. pratensis) isrecorded only from the Yamal Peninsula. Its nesting in the tundra zone has notbeen confirmed, although apparently isolated pairs raise their young in the south ofYamal. The yellow-headed wagtail (Motacilla citroela) nests on the Gydan Peninsulaand on the Yamal Peninsula to 70°N, while on the Yenisey it is numerous on theBrekhovskiye Islands. The yellow wagtail (M. flava) nests along the shores of thesoutheastern part of Taz Bay, and along the lower reaches of the Taz. It penetratesfar to the north along the Yenisey, whereas on the Ob' it only reaches Ob' Bay.

Of the tits (chickadees), the Siberian tit (gray-headed chickadee) (Pants cintus)is fairly common in the fall in the southern tundra subzone, and on migration alongthe lower reaches of the Yenisey. The northern shrike (Lanius excubitor) has beenrecorded from the lower reaches of the Yenisey in the southern tundra subzone,evidently just as a visitor.

Of the warblers (Sylviidae) five species have been recorded from the tundra ofthe Yamal and Gydan peninsulas. All of these birds are encountered only veryrarely, preferring the forest-tundra environment, or in the southern tundra subzoneareas with shrub vegetation. The willow warbler (Phylloscopus trochilus) and thechiff-chaff (Ph. collybitd) penetrate into the southern tundra of Yamal as far as 69°30'N, to 70° on the Gydan Peninsula, and to 71° on the Yenisey where they nestin limited numbers. The arctic warbler (Ph. borealis) extends somewhat farthernorth, especially on the Yenisey. Finally the yellow-browed warbler (Ph. inornatus)does not penetrate beyond the forest tundra zone as a nesting species, but inJanuary 1933 was collected as a visitor at Gyda-Yamo. The sedge warbler(Acrecephalus schoenobaenus) extends east as far as the Yenisey and nests on theYamal Peninsual as far as 70°N, and on the Gydan Peninsula as far as 71°N.

Of the passerines, apart from the warblers, birds of the flycatcher and thrushfamilies are encountered in the southern tundra, and individual species, some ofthem nesting, penetrate north to 71°. Two species of martins also raise their youngin the southern tundra subzone: the sand martin (Riparia riparis) and the housemartin (Delichon urbica). The latter nests along the Yenisey valley right to itsmouth, and the former along the left bank of Ob' Bay, at the mouth of the Nyda,and in the northern part of the Taz Peninsula.

With regard to hunting, the Anseriformes are of the greatest significance: theseinclude the bean goose and the white-fronted goose, and also some ducks: thelong-tailed duck, the king eider, and the pintail to some extent. The red-breastedgoose, which is endemic to the USSR is of exceptional value as an exchange reservefor zoos. Apart from these, both species of ptarmigan are of great importance forhunting by the local population.

Mammals

The insectivores (Insectivora) represent one of the oldest orders of mammals.Only moles (Talpidae) and shrews (Soricidae) sometimes occur in the Arctic. The

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fullest data on shrews appear in the works of B. S. Vinogradov [26], S. I. Ognev[12] and V. A. Yudin [28]. Moles are atypical of the tundra zone. The Siberianmole (Asioscalops altaica) is the only representative of the Talpidae family topenetrate into the tundra and the estuarine areas of the Yenisey as far as 70°N. Insize it is larger than the common mole, has visible eyes and movable eyelids, ashortened tail, and relatively small, narrow teeth. Six subspecies of Siberian moleshave been described; of these only Asioscalops altaica sibirica ranges far north downthe Yenisey valley. The northern forms of moles are colored a lighter greyish tone.Due to their limited numbers and limited range on the tundra, moles are of nopractical significance.

Seven species of shrews have been recorded on the Yamal-Gydan tundra,belonging to two genera: Neomys with one species, and Sorex with six species.

The water shrew (Neomys fodiens), a species adapted to an aquatic lifestyle,occupies the banks of rivers, streams and lakes, old river channels and swampylowland areas. The distribution of the water shrew has not been studied at thenorthern limits of its range. Its systematics are incomplete. On the Yamal Peninsulaindividual specimens penetrate into the southern tundra along streams which flowdown from the eastern slopes of the Urals. It is also found on the lower reaches ofthe Pur; here hill ridges do not present a barrier to its spread. At the northern limitsof its range the numbers of shrews are very low. This explains the paucity of dataon shrews from the tundra and forest-tundra zones.

Sorex isodon resembles members of the muridae in general appearance. It is notfound on the Yamal Peninsula, but is encountered on the right bank of the Ob', inthe basins of the Nyda, Pur and Taz, and also farther north between Taz Bay andYenisey Gulf, although it does not reach the latter. The distribution of this specieshas been very little studied. No subspecies have been described.

S. Boboratus. Although this shrew is somewhat smaller than the previousspecies, it belongs among the larger shrews. Four subspecies have been describedfrom Siberia. It penetrates into the tundra zone only between Taz Bay and YeniseyGulf. It is abundant on the left bank of the Ob' only south of the delta, and isabsent from Yamal.

Laxmann's shrew (S. caecutiens). This medium-sized shrew occupies a variety ofbiotopes: tundra, forest tundra, taiga, larch stands, open steppe areas and hills. Fivesubspecies have been described from Siberia and the Far East. The northernsubspecies (S. c. koreni) lives in the tundra zone from the Pechora to the ChukchiPeninsula inclusive. This shrew is not found on the Yamal Peninsula but is commonalong river valleys emptying into Taz Bay from the east. The numbers of Laxmann'sshrews on the Gydan tundra are small, comprising 2.7% of the total of all shrews.This percentage increases to 9.6% in the forest tundra of the Yamal-NenetsAutonomous Okrug.

The Arctic shrew (S. arcticus) is the only small insectivore which penetratesfarther north than the others and occupies some territory in the northern tundrasubzone. The Arctic shrew is common in Yamal north of 71° and on the right bankof the Yenisey as far as 73° 40'N. Six subspecies have been described from Siberiaand the Far East. The Pechora subspecies (S. ar. petschorae) occupies theYamal-Gydan tundra and extends east as far as the Yenisey. This shrew prefers

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habitats near water, either overgrown with shrubs and bushes, or well wooded andwith good grass cover, but it does not completely avoid mossy hummocks as long asthey are not too marshy. The Arctic shrew is close to the Laxmann's shrew in size.

Lesser shrew (S. minutus). This shrew is among the smaller members of itsgenus. It is only slightly larger than the least shrew. The lesser shrew is common inthe forest tundra, but is rare in the southern tundra and occurs only sporadically asfar as 69° 40'N on the Yamal Peninsula, and as far as the Arctic Circle on the TazPeninsula. In these northern ranges this shrew is not found east of the Taz basin.

Least shrew (S. minutissimus) is the smallest shrew of the genus Sorex. Itpenetrates into the southern tundra zone in extremely limited numbers, mainly inthe areas of the Pur and the Taz.

Rodents (Rodentia). The total number of species of rodents recorded in theYamal-Gydan tundra is 12. These comprise three families: squirrels (Sciuridae) withone genus and one species; mice, with seven genera and 10 species; and hares(Leptoridae) with one genus and one species. The squirrel (Sciurus vulgaris) appearsin the southern tundra in years when there is a surge in the population, during thesnowfree season, and just before winter when the animals are on the move. In suchyears it is taken in limited quantities on the Yamal-Gydan tundra. The farthestpenetrations by squirrels onto the tundra are recorded along the left bank of theYenisey.

By species composition the most numerous rodents are the mice, and parti-cularly the voles. Of these the only true tundra residents are the Ob' and Arcticlemmings, although these do penetrate south into the forest tundra. They form themajor food source for predatory mammals, raptors, and also for jaegers and somegulls, especially in years of lemming population explosion. The remainder of thevoles penetrate onto the tundra only in small numbers from the forest andforest-tundra zones. The Middendorf s vole and the Northern red-backed vole gofarther north than the other species. The two species of muridae, the house mouseand the Norway rat are parasitic companions of man and in his wake they penetratefarther north than the other species of voles with the exception of the lemmings.

The house mouse (Mus domesticus) gradually colonizes newly developed areas,as a common occupant of man's dwellings. Penetration by mice into the Arcticbegan relatively recently. At the present time mice are resident at several polarstations, settlements and ports. In particular they have become established at CapeKamennyy, Novyy Port, Ust'-Port and other settlements. The usual route ofpenetration by mice as far north as this is in association with the delivery of foodsupplies by ship.

The Norway rat (Rattus norvegicus). Rats have reached the North by exactlythe same route as mice. According to V. N. Skalon's [20] information, cases of thetransport of rats into the settlement of Khal'mersede at the mouth of the Taz havebeen recorded, but they have not survived there. Rats are now resident in NovyyPort, Cape Kamennyy and Ust'-Port.

The Ob' lemming (Lemmus obensis) occupies the Yamal-Gydan tundra as far asthe sea coast, and also the offshore islands: Belyy, Shokal'skiy, Vil'kitskiy,Sibiryakov, Oleniy and a few others. This is the most abundant species of rodent in

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the tundra zone. Numbers of lemmings are particularly large in years when apopulation explosion occurs, when they become the main food source for severalspecies of carniverous mammals (fox, ermine and weasel) as well as for many birds(white-tailed eagles, buzzards, glaucous gulls, herring gulls, jaegers, snowy owls andboreal owls). The migration of foxes, the nesting of gulls and owls, the numbers ofyoung in litters, and of nestlings in owl broods all depend to a great degree on thenumbers of lemmings. At high densities lemming migrations usually occur, whenthey abandon their home territories. Under these circumstances they swim acrossrivers (although they dislike getting wet), travel across ice, and cross areas of thetundra devoid of vegetation. Massive fatalities of these little animals are observedduring these migrations. Normal seasonal migrations consist of the movement oflemmings into low areas of tundra with a rich vegetation and deeper snow cover forthe winter. When snowmelt begins in spring the lemmings move out to the slopes ofdrainage divides. In summer they occupy practically the entire landscape avoidingonly excessively marshy areas and areas of bare ground totally devoid of vegetation.

The Arctic lemming (Dicrostonyx torquatus). This little animal is very closelyrelated to the Ob' lemming, but despite close similarities, it is distinguished by anumber of ecological peculiarities. Essentially it occupies (in the widest sense) thesame sites at the Ob' lemming, but it prefers elevated, dry areas of gravel-and-rocktundra and bedrock outcrops. The limited development of this tundra type and theabsence of outcrops in the Yamal-Gydan area markedly control the numbers ofArctic lemmings. There are no Arctic lemmings on the offshore islands but this doesnot signify the general northern limits of its range. The Arctic lemming is betteradapted to the severe conditions of the Arctic than the Ob' lemming in terms ofcoloration, pelage, the development of specialized winter claws, right down to suchaspects as diet. Apart from vascular plants the Arctic lemming eats several lichenswith are avoided by the Ob' lemming.

The northern red-backed vole (Evotomys rutilus). The major habitats of thisvole are in the forest zone. According to B. S. Vinogradov [26] it is a visitor to thetundra zone, but S. P. Naumov [10] considers it to be a common species in theforest-tundra of the Gydan Peninsula. Inasmuch as there is a specimen in Yakovlev'scollection [10] from Yenisey Gulf (from Tol'styy Nos at 70°N), without any doubtthis little animal also occupies the southern tundra [10]. Apart from this, one mayassume that the northern red-backed vole will also be found in the southwesternpart of the Yamal tundra, adjacent to the Northern Urals.

The large-toothed red-backed vole (E. rufocanus). This vole is typical of theswampy areas of the taiga and the forest-tundra, but it is not completely alien tothe southern tundra, as witnessed by the find of a lower jaw of this species infaeces at 69°N. One can assume that with a more detailed study of the Yamalrodents, it will be found in the southern subzone of that peninsula.

European water vole (Arvicola terrestris). V. N. Skalon has recorded findingwater voles in the valley of the Taz, quite close to its mouth, i.e., within the tundrazone [20]. It is also resident on the shores of the Ob' delta and, evidently,penetrates into the southern tundra on Yamal. On the lower reaches of the Ob' thenumbers of these voles become very considerable in peak years.

The tundra vole (Microtus oeconomus). From S. P. Naumov's data [10] itwould appear that this animal penetrates in small numbers into the southern tundra

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on the Gydan Peninsula, and is common in the forest tundra. In particular, theremains of the skull of a tundra vole were found in a faeces on the banks of theLutsempyayakha, within the tundra zone (69° 30'N). This is confirmed by B. S.Vinogradov [26] who notes that the vole is edging its way into the tundra zone inlow marshy areas.

Middendorf s vole (M. middendorffi). This vole is a visitor to the southerntundra of the Yamal and Gydan peninsulas and the lower reaches of the Taz andYenisey, preferring hummocky, marshy areas, where it builds a system of open andclosed runways and nesting chambers. These little animals also commonly buildwinter houses woven out of grass.

The large narrow-skulled vole (M. major). This species occupies the YamalPeninsula and the lower reaches of the Ob'; it is distinguished by its large size andby its light coloring. It is possible that this form and the European narrow-skulledvole (M. gregalis) are one and the same species. The ecology and taxonomic statusof these voles require further study. The large narrow-skulled vole is not found onthe Gydan Peninsula, hence it is quite possible that a break occurs in its dis-tribution between the Yamal and the Taymyr populations. The voles found onthe Yamal Peninsula lead a communal lifestyle; up to 20 voles of varying ages maylive in a single burrow, occupying up to 10 m2 in area. Up to 30—40 side burrowsleading to the outside branch off in various directions.

The snow hare (Lepus timidus). This species occupies the Yamal-Gydan tundraright to the shores of the Kara Sea. In the northern parts of these areas, however, itis encountered in appreciable numbers only in peak reproduction years. As a rule,previous authors [12, 20, 29] have placed the northern limit of its range much toofar south. It occupies the tundra everywhere, right to the Arctic Ocean coast,including Cape Chelyuskin; it is the only rodent of any significance in terms ofhunting, and even then only on a minor scale and not every year.

Carnivores (Carnivora). Almost all carnivorous mammals belong to fur-bearingspecies and are widely distributed throughout a great variety of natural zones. Themajority of them live in the forest zone but at the same time may include withintheir ranges several zones which they either occupy year-round, or on an alternatingbasis, depending on the season. They are all the object of hunting or trapping. Thewolverine prefers the forest-tundra and northern taiga zones although it does notentirely shun the tundra. The wolf, weasel and ermine are common on the tundra,the forest-tundra, the forest and the steppe. The red fox is encountered in thesezones also, but only rarely visits the tundra. One may encounter the brown bear inthe southern tundra, and even more rarely, the otter. Thus nine species ofcarnivores have been recorded from the Yamal-Gydan tundra, belonging to threefamilies. The bear family (Ursidae) includes two genera and two species; the dogfamily (Canidea), three genera and three species, and the weasel family (Mustelidae),three genera and four species.

The polar bear (Thalarctos maritimus) occupies three zoogeographical zones:the Arctic Basin, the arctic deserts and northern shelves of the marginal seas, andalso the tundra and southern shelves of the marginal seas [18]. In the latter zone itis encountered along the coasts, on islands and in inshore waters. There are fewbears in the Arctic Basin, and they normally only remain there in summer. Isolatedindividuals, mainly males, do reach the North Pole. The majority of bears spend the

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bulk of the year in the arctic-desert-and-northern-shelf zone. In winter, and thisapplies particularly to pregnant females who den up for lengthy periods, they leavethe pack ice, come ashore on island coasts, and den up in snowdrifts. Males den upfor shorter periods in the very darkest period of the year. Bears very rarely excavatedens on the mainland coast. Particularly in connection with the general decline inthe number of bears, for years now no dens have been found on the offshoreislands or on the mainland coasts of the Yamal-Gydan area. Earlier there was comehunting on the Yamal and Gydan peninsulas, yielding up to 20—30 hear per year.Bear hunting has been prohibited since 1956.

The grizzly bear (Ursus arctos) is not at all typical of the Yamal-Gydan tundra,but nonetheless lone individuals are sometimes observed along the lower reaches ofthe Taz and Pur rivers. They are also rare in the forest-tundra and have beenreported only during the summer.

Arctic fox (Alopex lagopus). The total range of this fox is very extensive, andvaries greatly from year to year. On their wanderings during winter foxes may beencountered out on the ice of the Arctic Basin and south into the forest zone. Thebreeding range is also variable depending on peaks in the reproductive cycle of thelemmings. With minor exceptions, foxes do not breed in the arctic desert zone. Thelitter size in the tundra is also small when there is a poor lemming "crop" prior tothe birth of the pups. After the cubs have been weaned and until they start toemerge from the den, the main diet of the cubs consists of small rodents providedby the parents. The absence of owls, jaegers and some other birds, or their failureto nest on a large scale, are good indicators of a poor breeding year for the localfoxes. The most productive area for raising cubs is the southern tundra subzone andthe forest tundra where, when there are few lemmings, they may be replaced byother species of rodents.

Fox trapping in the Yamal-Gydan tundra is effected by the use of steel trapsand deadfall traps. In that the main nomadic population has left the northernregions by the beginning of the trapping season, there are few well equippedtraplines there. Since they migrate with their reindeer, the Nentsy are unable todevote much attention to fox trapping and limit themselves to setting traps on atemporary basis. At the present time trapping is carried out by the residents of thesettlements. In good years totals of up to 25,000 foxes are harvested here. In the1974—75 trapping season the harvest of foxes was very poor.

Wolf (Canis lupus). Wolves are encountered only sporadically on the Yamal-Gydan tundra, and despite the intensity of domestic reindeer herding, they do notinflict heavy losses on the herds in summer. Researchers of the AANII's physicalgeographical expedition working here in 1971—73 saw wolf tracks only twice, andobserved a solitary individual only once. A. N. Tyulin, who wintered on BelyyIsland in 1931, noted the sighting of a wolf in February and saw a solitaryindividual on May 30 [22]. In general, wolves occupy the forest tundra in winter.In the tundra zone they are most frequently encountered in the northern areas inwinter, where the wild reindeer are concentrated. According to S. P. Naumov [10]wolves remain for the whole winter on the Gydan Peninsula in particular, andespecially on the Yavay Peninsula.

Red fox (Vulpes vulpes). At present the red fox rarely penetrates into the

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tundra zone, but individuals have been killed there on the Yenisey, on the lowerreaches of the Taz and the Pur and in other areas.

Wolverine (Gulo giild). The main habitat of the wolverine is confined to theforest tundra and the: northern taiga subzones; nevertheless they visit the tundraevery year, travelling north as far as the sea coast. They have been recorded on theYamal Peninsula by S. P. Naumov [10]. They are fairly common on the southerntundra where they inflict losses on the trappers by eating foxes caught in the trapsand caches of fish and meat left on the tundra. Questionnaires revealed that in thewinter of 1973—74 individual trappers killed 6—7 wolverines each during thetrapping season on the tundra of the Gydan Peninsula.

Otter (Lutra lutrd). Along the Ob' the otter ranges north right to the delta, butit has not been observed in the rivers of Yamal or in Ob' Bay, although incursionsinto the southern tundra are quite feasible. This statement is based on the fact thatin some years otters have been killed on the lower reaches of the Pur and Taz, andon the very lowest reaches of the Yenisey, well within the southern tundra.

The ermine (Mustek erminea) is a common resident of the Yamal and Gydantundras. It is encountered everywhere, right to the coast and incontestably occurson the offshore islands. In particular, we have frequently recorded it on DiksonIsland. In the southern tundra it is encountered in commercial quantities, but as aresult of the nature of the evolution of employment among the local people,nobody specifically concentrates on trapping ermine. In lemming peak years thenumber of ermine also increases. With regard to the systematic classification ofsubspecies of ermine, this needs to be thoroughly examined since the data from thenorthern areas are extremely meagre and are not mutually comparable.

Weasel (Mustek nivalis). The range of the weasel differs little from that of theermine. As a rule the weasel does not range so far north, and is everywhere lessnumerous than the ermine. In the Gydan Peninsula S. P. Naumov collected a weaselwhich had been killed in the middle of the Yuribey (at 70° 47'N). The AANIIexpedition to Yamal recorded a weasel in the vicinity of Cape Kamennyy.

Ungulates (Ungalata). These belong to two different genera: elk (Alces) andreindeer (Rangifer). The former is not typical of the tundra and only isolatedindividuals (mainly bulls) wander out onto the tundra, reaching 71°N on the YamalPeninsula. They may also be encountered as far as approximately the same latitudeon the Gydan Peninsula. The European elk (Alces alces), which occupies WesternSiberia, does not extend farther east than the Yenisey; here its place is taken byanother elk, the American elk [moose] [30]. Evidently it is more appropriate toconsider the elk as a single species, particularly since the differences can be reducedto color variations, size indices, and a few skull variations. This being the case, onlyextreme forms, widely separated from each other can be easily distinguished, whilein contiguous ranges the differences can only be detected with difficulty.

Thus the geographical variability of the elk (e.g., the general increase in sizetoward the east) must be seen as a subspecific index, and the less significant changesas a racial index. The northern forms of the elk are larger than the southern onesand also lighter in color. Penetration by elk into the tundra and forest tundra (theyare rare in the latter zone too) only occurs in summer. This may be explained bythe fact that elk, as the largest representative of the deer family, feeding in winter

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on coarse woody and shrubby vegetation, including conifers, simply are unable tosupport themselves on the tundra except during the vegetative period, when theycan compensate for the absence of their normal foods with marsh grasses. Forestecotones are the major habitat of the elk. Thus the appearance of elk on theYamal-Gydan tundras must be seen as an accidental phenomenon.

Reindeer (Rangifer tarandus). As distinct from the elk the reindeer is a trulyindigenous species of the tundra zone, where it spends the major part of the year.Admittedly there are isolated populations located far to the south in the forest-tundra zone and also to the north in the arctic desert zone. In the Yamal-Gydantundra the reindeer occupies not only mainland areas but also offshore islands,including Belyy, Shokal'skiy, Vil'kitskiy, Oleniy, etc. Nonetheless, as compared toother areas of the Arctic, the numbers of wild reindeer are small. This may beexplained by the fact that in the tundra areas located between Baydaratskaya Bayand Yenisey Gulf, intensive domestic reindeer herding constricts the range of thewild animals. In the Yamal-Gydan tundra the migrations northwards in spring andsouthwards in fall, so typical of the reindeer in the tundra zones of the Taymyr,Khatanga, Anabar, Lena, Yana and Indigirka, are not observed at all. The majorityof the reindeer, which formerly migrated in this pattern, have changed their routes,while some have also reduced their range.

At present few reindeer are encountered in the central parts of the Yamal andGydan peninsulas, whereas they are common farther south in the forest tundra andfarther north along the Kara Sea littoral, and also on the islands. At present thehighest densities of reindeer are recorded on the islands. The reindeer occupying thesouthern forest-tundra zone do not move en masse to the tundra zone in spring,nor do those occupying the islands reach the forest zone in winter, but winter onthe tundra of the Yamal and Gydan peninsulas; to some degree they also winter onthe islands. In general the number of reindeer in the Yamal-Gydan area hasdecreased to 50%, 33% or even less, as compared to the 1920s to 1940s. One canjudge as to this from the observations by the AANII expedition of 1970-73, ascompared with the data presented by B. M. Zhitkov [29], S. P. Naumov [10] andA. N. Tyulin [22]. As regards reindeer systematics, they are so incomplete that abasic reorganization is required, which should take into consideration reindeerranges abroad.

Pinnipeds (Pinnipedia). Only four species of pinnipeds have been recorded asvisiting the waters of the Yamal-Gydan area, comprising two families and fourgenera. Of these the harp seal must be considered as a very occasional visitor, whilethe walrus has lost its once considerable significance as a representative of the localfauna.

Walrus {Odobaenus rosmarus). This, the sole representative of the familyOdobaenidae, forms an independent genus Odobaenus with only one species,Odobaenus rosmarus. In the seventeenth and eighteenth centuries walrus werepresent in large numbers in the waters adjacent to the coasts of Yamal and Gydan,both in the northern part of Yenisey Gulf and in Ob' and Gyda bays; howeverthey were particularly numerous in Baydaratskaya Bay.

In September 1690 Rodion Ivanov, who was shipwrecked with some com-panions on the Sharapovyye Koshki, reported an abundance of walrus and sealsthere. They amassed 40 poods of walrus tusks [a pood is a Russian unit of weight

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equal to about 36 pounds] [25]. Right down to the end of the 1920s walrus werehunted in Baydaratskaya Bay, in Malygin Strait and in the Kara Sea by Russian andNorwegian hunters.

According to the Nentsy, solitary walrus are occasionally encountered even nowon the west coast of the Yamal Peninsula as far as the Mordiyakha River. They havebeen observed in Ob' Bay at Cape Drovyanoy. According to S. P. Naumov [10],who got his information from the Nentsy, walrus are sometimes killed in Gyda Bay.He also quotes P. Yu. Shmidt's rather old data concerning walrus going up theYenisey as far as Dudinka. The decline in the numbers of walrus is general, andtheir almost complete disappearance from the waters in question is to be explainedby excessive hunting during the nineteenth century and subsequently. At thepresent time walrus of this subspecies are encountered only singly or in smallgroups, and nowhere do they haul out east of Novaya Zemlya. In the 1940s and1950s the author repeatedly saw walrus, singly and in groups of up to four on theice of the Kara Sea, well out from the mainland coast.

Bearded seal (Erignathus barbatus). An alleged subspecies of this seal (E. bar.barbatus) occupies the waters adjacent to the coasts of the Yamal-Gydan area.Bearded seals are encountered more often in the northern areas, but even there theyare not numerous. They prefer to lead a solitary type of life but sometimes one mayobserve several animals at once on the ice. During the open water period beardedseals enter the rivers, sometimes running upstream for 20—30 km. In the souththese seals are occasionally reported as far as the mouths of the Ob' and theYenisey and they have also been killed in Taz Bay. No special hunt for beardedseals occurs here, but while hunting ringed seals the local residents and the thehunting artels also shoot these seals, since their hides are highly valued, being usedfor the manufacture of lines, harness and other equipment for reindeer herding.Bearded seals are most frequently killed off the Yavay coast, off the west coast ofYamal and in the Dikson Island area.

The Greenland seal (Histriophoca groenlandica) or harp seal, is included in thelist of local fauna solely on the basis of a report by M. Dukhovnyy, which wasquoted by S. I. Ognev [12]. Judging from this information, the most easterlyoccurrence of this seal is Gyda Bay, where it is hunted occasionally. Evidently thisstatement refers to inshore waters since out in the Kara Sea, well away from themainland coast, this seal is observed much farther east during migrations. There isno other reliable material to substantiate the occurrence of harp seals in thesewaters.

Ringed seal (Phoca hispida). This species may be subdivided into severalsubspecies. The "pomor" ringed seal (Phoca hispida pomowrum) occupies thewestern part of the Kara Sea. It is found in all the bays and gulfs of theYamal-Gydan area and remains there throughout the year. It is frequentlyencountered in the mouths of the rivers, up which it travels, penetrating into lakes.Cases have been recorded from several lakes in both Yamal and Gydan of ringedseals having wintered. In winter the ringed seal leads a hidden, subnival sort of life,excavating dens in the fast ice. Inside the den, apart from an area for the animal tohaul out, is located a "breathing hole" through which the ringed seal dives down tocatch fish. By swimming back to its breathing hole it can always refill its lungs withair. The den is kept perfectly clean, and there are never any food remnants orfaeces in it. Pupping also occurs in the den. The water in the breathing hole does

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not freeze since apparently a sufficiently high temperature is maintained inside theden from the seal's breathing.

The hunting of ringed seals is of great significance to the local population; theirmeat and blubber are used for food and for bait, while the skins are used formaking clothing and for other domestic purposes. The best hunting period isJune—July when the seals haul out on the ice. Hunting artels in Ob', Gyda and Tazbays and in Yenisey Gulf are all involved in hunting ringed seals.

Whales (Cetacea). Two representatives of the cetaceans have been recorded fromthe waters of the Yamal-Gydan area, belonging to two different suborders. One ofthese, the beluga, belongs to the toothed whales (Odontoceti) and to the in-dependent genus Delphinapterus. The second, the fin whale (a rorqual) belongs tothe suborder Mystacoceti.

The beluga {Delphinapterus leucas) is a long-established resident of the arcticseas. The nominal form occurs in the waters of the Yamal-Gydan area, but morespecifically it is the Kara beluga (Del. leuc. leucas) which mainly stays along thecoast, readily visiting bays and inlets, entering rivers and even penetrating up themfar into the interior; it even appears in some lakes. On the Yamal Peninsula theoccurrence of beluga has been recorded from Neyto Lake; to get there they have toswim up the Mordyyakha and its tributary the Seyakha, where they have frequentlybeen killed. On the Gydan Peninsula beluga have been observed to penetrate up theYuribey as far as 200 km from its mouth, right up to Lake Yaruto.

Beluga are encountered regularly throughout the entire extent of Ob' Bay inboth small and large schools. The farthest observed penetrations by beluga up theOb' have been as far as Berezovo and even farther, a distance of up to 1,500 kmfrom the mouth, and up the Irtysh as far as Khanty-Mansiysk and the Tobol. InGyda Bay and neighboring inlets beluga are encountered constantly, as is also truefor Yenisey Gulf. Penetrations of beluga up the Yenisey are reported fairlyfrequently. These mammals may be encountered throughout the whole of Taz Bayand individual schools run up the Pur and Taz as far as 67°. Beluga are just ascommon in the waters of the Kara Sea around the more northerly coasts of theYamal and Gydan peninsulas as in the bays and gulfs.

Beluga do not winter in the waters of the Ob' or Yenisey or even in the KaraSea, unless one includes isolated, rare, accidental cases; however they do winter inthe Barents Sea. The spring reappearance of the beluga occurs in late May and June,and in some places even later, depending on hydrologic and ice conditions. Theseanimals are better adapted than other species of whales to heavy ice conditions.They avoid only major fields of pack ice, or fast ice occupying extensive areas.

Many authors have generally linked the appearance of large schools of belugawith massive runs of Arctic cod. In the case of the Kara, Barents and White seas,where the Arctic cod does not represent the main item in diet of the beluga, thisfactor is not of such critical significance. Here, apart from Arctic cod the belugafeed on many species of shoaling fish. The beluga penetrates very far to the northand is even encountered north of Franz Josef Land and Severnaya Zemlya. As onemoves out toward the open sea, away from land, the role of the Arctic cod as afood source for the beluga, increases.

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As a rule, the beluga gradually move out from Ob' Bay and Yenisey Gulf intothe Barents Sea via the Novaya Zemlya straits, or by rounding Novaya Zemlya.Although it is possible to observe large schools of beluga of up to several hundredanimals in late September in the Kara Sea, their numbers by then are noticeablydiminishing. The last small schools are recorded as late as mid-October.

Beluga occur in commerical quantities in the Yamal-Gydan area and hence theyare caught by hunting artels at many points in the bays and gulfs.

Since 1953 beluga hunting has been carried out by the Tyumen' Oblastauthorities using hunting vessels. Nevertheless the stocks of this species are notbeing utilized to the optimum. At the present, hunting of beluga from ships is alsopracticed to the east of Yenisey Gulf.

Fin whale (Balaenoptera physalus). On September 14, 1950, a rorqual wasobserved in the Yenisey, swimming upstream, and on the 18th it was found dead onthe left bank of the river, 15 km from Dudinka. It turned out to be a young finwhale 14.7 m long, which had only recently been weaned [21]. This is the firstknown occurrence of a rorqual in the Kara Sea.

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2. Berg, L. S. Ryby presnykh vod SSSR i sopredel'nykh stran [Freshwater Fish ofthe USSR and Neighboring Countries]. Moscow-Leningrad: Izdatel'stvo ANSSSR, Pt. 1, 1948; Pt. 2, 1949; Pt. 3, 1949.

3. Berezovskiy, A. N. "The whitefish (Coregonus) of the Yenisey River," TrudySib. ikhtiol. lab., Vol. 2, No. 1, 1924.

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6. Dmitriyev, V. I. "Fish and fisheries in the lower reaches of the Yenisey River,"Trudy Nauch.-issled. in-ta polyarn. zemled., zhivotnovod. i promysl. khoz-va.Ser. "Promysl. khoz-va,"No. 16, 1941.

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26. Vinogradov, B. S. Mlekopitayushchiye SSSR. Gryzuny [Mammals of the USSR.Rodents]. Leningrad: Izdatel'stvo AN SSSR, 1933.

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