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    Phytogeography and Vegetation of the Soviet Arctic Region

    Encyclopedia Arctica 6: Plant Sciences (Regional)

    Phytogeography and Vegetation of the Soviet Arctic Region

    Unpaginated      |      Vol_VI-0327                                                                                                                  
    EA-Plant Sciences

    (Eric Hult e é n)




    Limitation of the Arctic Zone 1
    Historical 5
    Geology and Soil 6
    Climate 8
    The Flora 10
    Number of Species 10
    Distribution of Species 12
    I. Plants occurring throughout the Soviet Arctic or at

    both the eastern and western ends
    A. Circumpolar Plants 12
    B. Eurasiatic Plants 18
    II. Plants occurring in middle part of the Soviet Arctic

    but not at both ends
    A. Species not apparently washed down by the rivers 19
    B. Species apparently washed down by the rivers 20
    III. Plants occurring either in western or eastern part

    of Soviet Arctic
    A. Plants reaching the western but not the eastern end of

    the Eurasiatic Arctic
    B. Plants reaching the eastern but not the western end of

    the Eurasiatic Arctic
    IV. Endemic Plants 36
    V. Singular Species 39

    Unpaginated      |      Vol_VI-0328                                                                                                                  

    (Eric Hult e é n)



    Origin and Composition of Arctic Flora 40
    The Vegetation 42
    Zones of Vegetation 43
    Snow zone 43
    High-Arctic Tundra 44
    Lichen-Moss Tundra 48
    Shrub Tundra 54
    Forest Tundra 56
    Vegetation of Different Geographical Provinces 58
    Kola Peninsula 58
    Kanin Peninsula 60
    Molozemelskaya and Bolshezemelskaya Tundras 61
    Kolguev Island 62
    Polar Urals 63
    Novaya Zemlya and Vaigach 64
    Franz Josef Land 67
    Yalmal and West Siberian Lowland 67
    Gydan Tundra and Mouth of the Yenisei 68
    Taimyr Peninsula 70
    Anabar Tundra 71
    Mouths of Olenek and Lena Rivers 73
    Lena to Kolyma and Chaunskaia Bay 74
    New Siberian Islands 75
    Chukotsk Peninsula 75
    Wrangel Island 78
    Bibliography 79

    Unpaginated      |      Vol_VI-0329                                                                                                                  
    EA-Plant Sciences

    (Eric Hulten)



            With the manuscript of this article, the author submitted 4 map

    sketches. These are being held at The Stefansson Library.

    001      |      Vol_VI-0330                                                                                                                  
    EA-Plant Sciences

    (Eric Hult e é n)





            The limitation of the arctic zone to the south is variously determined

    by different authors. This is especially the case where mountains enter

    into the arctic tundra zone, as in Scandinavia, in the northern Urals, and

    in easternmost Asia. In the lowlands either the northern limit of single

    trees or the limit of real forest can be taken as the southern boundary line.

    In this article the tundras with scattered trees or very open forests are

    included in the forest zone as this seems to be most natural, and the southern

    boundary line of the Arctic is thus taken to be the polar tree line.

            From Kanin to Kolyma the timber line consists of conifers and is fairly

    well defined, but on Kola Peninsula a belt of more or less shrubby or stunted

    birches is inserted between the coniferous forest and the open tundra. This

    birch belt is a continuation eastward to the subalpine ( Betula tortuosa ) belt

    of the Scandinavian mountains. In the river valleys this birch belt reaches

    or nearly reaches the northern coast, while the headlands between the rivers

    lack this shrub vegetation. Polunin does not refer the northern coast of

    Kola Peninsula as belonging to the Arctic, probably on account of this birch

    belt or possibly because permanently frozen ground has small extension there.

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    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    As, however, the floristic composition as well as the vegetation and the soil

    of the treeless parts of the north coast of Kola Peninsula agree almost

    exactly with the corresponding features in arctic Siberia, I must include

    them in the real Arctic. The most arctic part is the middle coastal part

    of the peninsula, where solifluction is common. Eastward as well as westward

    many more southern components alien to the real arctic tundras appear step

    by step, eastward on account of the southeastern direction of the coast line,

    westward on account of the strong Atlantic influence of the Gulf Stream.

            East of Kolyma conditions change very rapidly. The last trees are not

    conifers but willows and birches forming fringing forests chiefly in the river

    valleys. The chukotsk Peninsula, the Anadyr and Penzhina basins form a large

    treeless area with larch forests only in the central western part of Anadyr

    and alluvial Salix-Betula forests in the large river valleys. This treeless

    area also continues southward along the western coast of Kamchatka to the

    southern point of that peninsula and along the Kurils to Uruppu Island on one

    hand, and on the western side of the Okhotsk Sea to about 61° N. latitude on

    the other. As on Kola Peninsula, a subalpine belt is inserted between the open

    tundras and coniferous forests in the Far East. This belt consists of shrubby

    Alnus and Pinus pumila thickets.

            The question is whether this vast area should be regarded as belonging to

    the Arctic or not. Polunin refers only the arctic drainage to the Arctic, ex–

    cluding the Anadyr Valley and everything to the south of it. It is hard to

    find a reason for this. The floristic composition in the Far East is somewhat

    different from that of other parts of the Arctic, but this holds true also

    concerning the arctic drainage of Chukotsk Peninsula. The vegetation east of

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    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    the Kolyma likewise differs somewhat from that of other parts of the Arctic,

    but there is not much difference between the vegetation on northern Chukotsk

    Peninsula and that of the more southerly treeless areas. It is therefore

    not easy to find a line of demarcation for the Arctic to the south based on

    tenable grounds.

            It has been argued that the treeless areas in the Far East are merely

    an alpine zone lowered down to sea level. Floristically this is true, as

    very many eastern Asiatic mountains species protrude much farther northward

    and eastward here than in any other place, but the same holds good of the

    flat, treeless areas of the northern Kola Peninsula. There are, however,

    several characteristics in which the treeless zone of the mountains differs

    from the tundra zone of the lowlands.

            The tundra zone is characterized by strong permanent summer light, low

    precipitation and, thus, shallow snow cover, permanently frozen soil and

    consequently bad drainage, with frequent shallow pools at melting time and

    a poor humus layer or none at all. In the southern mountains the light is

    very variable on account of the day-night variation and the direction of

    the slopes; the temperature varies a great deal likewise on account of the

    direction of the slopes; the precipitation is much hii higher and the snow

    cover is deep in places; the soil is not frozen, as the first snow falls

    early in the autumn and shelters the soil from frost. Melting starts early

    in the spring, giving a longer period of vegetation; the humus layer is

    rich, and a layer of humus and roots is formed where even geophytes can

    thrive. In the northern mountains the conditions approach those of the

    tundra. The sunlight is continuous as on the tundra; on the other hand,

    004      |      Vol_VI-0333                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    the temperatures are higher, the precipitation is higher but the draiage is

    better and the earth drier and warmer; the snow cover is in places deeper

    and the snow falls earlier; the period of vegetation is longer as the melt–

    ing of the snow starts earlier. The reason for the treelessness of the

    mountains, judging from the zone of stunted trees occurring at the timber

    line, is the strong winds, especially during the winter; while on the flat

    tundra, where the last trees are usually not stunted, but nearly branchless

    poles, the reason for the treelessness is that it is difficult for trees to

    take up enough water from the frozen soil there during the summer, when the

    evaporation is strong.

            Because of the above ecological advantages, the forests oftenetend farther

    northward in the mountains in the Urals and in Anadyr, for instance, than in

    the adjacent flat tundra.

            The questions is: Can these different ecological conditions help us to

    understand the nature of the large treeless areas of the Far East? In most

    coastal parts, at any rate, a higher soil temperature and deeper and earlier

    snow cover due to the interrupted distribution of the permafrost make the

    conditions correspond to mountain conditions, but they are hardly s sufficient

    to explain the differences, which doubtless have deep historical reasons.

            Although it is thus very difficult to find a reasonable argument for ex–

    cluding the treeless areas down to northern Kamchatka (where the permanently

    frozen ground ceases) in the Arctic, it is here taken to include the Chukotsk

    Peninsula only, that is, southward to the northern slopes of the Anadyr Val–


    005      |      Vol_VI-0334                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic



            The arctic tundra zone as defined above is a series of undulating

    plains. Mountains enter into this zone in the Urals, at Novaya Zemlya,

    in Taimyr Peninsula, and in the Far East. The tundra zone is not very

    well investigated botanically but, as large areas of it are fairly monoto–

    nous, it is possible to get a fairly accurate view of their phytogeograph–

    ical conditions.

            During the Tertiary the northern part of Eurasia was covered with a

    subtropical vegetation which, when the deterioration of the climate took

    place in the Quaternary period, was gradually replaced by a more arctic

    vegetation. The history of the tundra zone as a treeless area is thus

    not a very long one — doubtless considerably shorter than a million years.

    During the different glacial and interglacial periods great changes took

    place in the zone, changes of which we know very little at present.

            The first scientific reports on the arctic tundras were brought back

    by the Great Northern Expedition, which had worked its way along the shores

    of the Arctic Sea in central and eastern Siberia in 1734-1742. The tundras

    of arctic European Russia and western Siberia were first visited by an ex–

    pedition sponsored by the Russian Academy in 1771-1772, and of which Ozeret–

    skovski and Zuev were members. When Cook navigated Bering Strait in 1778,

    the easternmost Eurasiatic Arctic was opened up for scientific investigation.

            In 1787-1791, Billings surveyed the coast between the Kolyma and the

    Chukotsk Peninsula; in 1806, Adams investigated the mouth of the Lena River;

    in 1837, Baer visited Novaya Zemlya, and Schrenk the Bolshezemelskaya tundra;

    006      |      Vol_VI-0335                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    in 1843, Middendorff made his important journey to Taimyr; in 1850, Hoffman

    investigated the northern Urals and the Pai-khoy Range; in 1866-1880, Schmidt

    visited the Gydan tundra and lower Ob River.

            In 1878-80, Nordenskiőld and Palander made the first circumnavigation

    of Eurasia and thus sailed for the first time along the arctic coast from

    Europe to the Chukotsk Peninsula on board the Vega . The botanists on this

    famous expedition were Kjellman and Almquist. Kjellman’s reports, published

    in the Scientific Results of the Vega expedition, have up to recent times

    been the most widely quoted source of botanical information on the arctic


            In 1887, Kihlman thoroughly investigated Kola Peninsula, and, in

    1900-1902, Toll visited Taimyr and the New Siberian Islands.

            In the twentieth century, numerous botanical investigations were under–

    taken on the Russian tundras mainly by Russian botanists, among whom may be

    mentioned Tolmachev, Zhitkov, Kuznetzov, Grigoriev, Pohle, Regel, Andreev,

    Sochava, Reverdatto, Gorodkov, and Sambuk.



            The first attempt to divide the Soviet Arctic into Phytogeographic units

    was made by Trautvetter. In recent times Andreev, Gorodkov, Rikli, Sambuk,

    Sochava, and others have dealt with the zonation of this region, each one in

    a way differing somewhat from the others, so that any generally recognized

    results in regard to the zonation can hardly be said to have been achieved.

            The geological conditions in the arctic tundras are little known as yet.

    The bedrock is rarely visible, the surface being covered with gravel, sand,

    or muck. The possibility of the occurrence of calciferous rocks, which might

    007      |      Vol_VI-0336                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    be expected to exert some influence on the flora and vegetation, cannot there–

    fore be discussed here. Gorodkov has noted that on the tundras of central

    Siberia, and also on the Bolshezemelskaya tundra, crusts of lime can be

    found on the surface of the tundra polygons. They are formed by calciferous

    salts ascending with the water from the soil in the crevices of the polygons

    and drying on their surface.

            The most important ecological feature characterizing the tundras is

    doubtless the permanently frozen soil (only to a small extent occurring in

    Kola Peninsula).

            In summer the ice melts to a maximum depth of 1.5 to 2 m., usually much

    less. The temperature in the surface layer of the soil can (although rarely)

    reach 30°C. in summertime even at high latitudes but drops very quickly with

    increasing depth. Thus, for instance, the temperature 5 cm. below the sur–

    face in Franz Josef Land was 3° to 7°C.; 20 cm. below the surface in Taimyr

    (76° N.) it was 0.5°C.; while in the south of Bolshezemelskaya tundra 50 cm.

    below the surface, it was 5° to 10°C. In Penzhina it is 10° to 15°C. at a

    depth of 10 cm. in August and never above 4° or 5°C. at a depth of 20 cm.

            The permafrost is by no means restricted to the tundra belt. Districts

    with patches of frozen soil are found east of Yenisei south to about 52° N.

    latitude in central and eastern Siberia, and to about 60° N. in northern

    Kamchatka, especially in peaty soil. (See map.) South of the tundra zone

    the ice melts to a depth great enough to allow forest growth, and the layers

    of ice at considerable depths in the soil apparently have very little influ–

    ence on the vegetation.

            In wintertime the tundra soil is frozen throughout, and in spring at

    008      |      Vol_VI-0337                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    first only the top layer thaws, with the result that shallow pools are

    formed which are doubtless partially responsible for the open vegetation

    that is characteristic of the entire zone. Sometimes the tundra is covered

    at melting time by water more than a meter deep. Only such plants that can

    stand the constantly wet conditions are able to grow on the tundra. Which

    plants grow in a given place depends to a large extent on the depth to which

    the thawing proceeds in spring and early summer. Shrubs and plants with

    deep roots can grow only in places where the soil thaws deeply and quickly.

    Peat does not accumulate in the tundra zone except along the southern rim.

    In Europe and western Siberia deep treeless peat bogs are found in the

    northern parts of the forest zone, but they are lacking in the forests of

    the east. Peat bogs are not formed within the area of the permafrost, where

    they are replaced by boggy and shrubby tussock communities. In cal clayey

    or muddy soil, polygon tundras are formed by the freezing process, but not

    in sandy soil, which thaws to a greater depth than the peaty soil.

            Sochava considers that the mineralization of the plant residue does not

    fully take place; muck accumulates owing to the low temperature, and this

    soil formation differentiates the tundra from the bogs where peat is formed.

    Gorodkov opposes this view and maintains that there is no difference in

    principle between the tundra and the more southern areas in this respect.



            A characteristic of the tundra belt is the low yearly precipitation.

    Nowhere does it average more than 250 mm. except on the Kola Peninsula, where

    it reaches about 400 mm. on the north coast. It is less than 150 mm. between

    the Yenisei and the Kolyma.

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    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

            Nevertheless the degree of cloudiness is high, 70 to 75% of the days in

    the year being cloudy. A Only about 10% of the precipitation falls in winter–

    time. The depth of snow is accordingly small, about 20 to 30 cm. on Yalmal

    and in Taimyr, 15 to 20 cm. at the mouth of the Lena River, but much deeper

    on the forest tundra, for instance, 45 to 60 cm. on Kola Peninsula. During

    the winter the weather is of an anticyclone type, with prevailing southern

    winds in the tundra zone. In summertime the prevailing winds are northerly

    and fairly strong, for instance, at the lower Yenisei and Ob rivers, averag–

    ing 6 to 7 meters per second.

            The winter temperatures are low, though this is of little consequence

    to the plant life. Average temperatures for January are: in Kolguev, 13°C.;

    at Novaya Zemlya (Karmakuly), −16.5°; at the mouth of the Ob (Obdorsk), −25.6°;

    at the mouth of the Yenisei (Dudinka), −23.9°; at the mouth of the Lena (Bulun),

    −40°; and at the mouth of the Anadyr, −23°.

            The period of vegetation varies a good deal. In the most arctic parts it

    lasts only about 2 months, with an average temperature of 4°C., while in the

    middle tundra zone it is about 3 months with an average temperature of about

    6°, and, lastly, in the forest tundra about 4 months with 9.4° average tem–

    perature. At the mouth of the Yenisei it extends over 4 months and the average

    temperature is 8.6°; at the mouth of the Anadyr the average for July is 11°,

    and at Markova, at the eastern limit of the forest, it is 14.5°C. Many

    flowers of the Far North can apparently stand very low temperatures. For

    instance, Kihlman states that Cochlearia can freeze solid while in flower

    and continue to grow after thawing.

            The tundra zone is not — as some have called it — an arctic steppe

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    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    nor should the snow fields be called arctic deserts. The reason for the

    treelessness is quite different from that in the steppes or deserts. The

    steppes have never during their historical development had any connection

    with the tundra zone. A few steppe plants have been washed down by the

    great rivers and found themselves at home in the tundra zone, but this does

    not entitle us to regard the tundras as steppes.




    Number of Species

            In the following discussion of the flora only the vascular plants are

    taken into consideration, as the distribution of cryptogams is mostly too

    little known to allow of an analysis. However, in the part dealing with the

    vegetation, mosses and lichens characteristic of the different vegetative

    units are enumerated. It should only be mentioned that the sea algae, which

    were fairly well studied by Kjellman, are few in number, in his view owing

    to the brackishness of the water along the coast, caused by the great volume

    of fresh water emptied into the Arctic Sea by the large-rivers. The sea

    algae amount to about 35 species, of which the most important are: Polysiphonia

    , Phyllophora interrupta , Rhodomela tenuissima , and laminarias.

            The tundra zone forms a bow starting in the west on the Kola Peninsula

    at approximately 69° N. latitude, extending to about 77° N. over the Taimyr

    Peninsula, and thence to the Ch7kotsk Peninsula at about 67° N. To this must

    be added the Franz Josef Archipelago and Severnaya Zemlya, from which no

    botanical reports are known to the author; also the New Siberian Islands and

    Wrangel and Herald Islands.

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    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

            It is but natural that a wide botanical variation should be noticeable

    in a belt about 4,000 miles long and with a range in latitude of more

    than 15°.

            The exact limitation of the zone southward has much influence on its

    species content, as many species characteristic of the boreal woods accom–

    pany these woods to their northernmost boundary line. Many of these

    species protrude somewhat farther northward, thus entering into the Arctic.

    Thus along the southern rim of the tundra belt there suddenly occurs a

    great variety of forest species which are rather alien to the Arctic. It

    is hardly possible, therefore, to give a list of all species belonging to

    the arctic zone, but in the following enumeration those occurring north of

    the forest tundra are listed and classified as far as they are known to

    the author.

            The number of species occurring within a given space is somewhat

    variable. Thus, 36 species are known to occur in the Franz Josef Archi–

    pelage, 180 in Kolguev, 190 in Novaya Zemlya, 105 at Dickson Island (a

    small island near the mouth of the Yenisei), about 200 in the Taimyr Penin–

    sula, 63 in the relatively small Preobrazhenie Island, and 150 at Wrangel

    Island. The Chukotsk Peninsula is still not very well known botanically

    but it possesses more than 450 species — probably about 500.

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    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic


    Distribution of Species





            Most of the species which form the basic vegetation in the Arctic are

    circumpolar species. Other components of the flora play only a very minor

    part in the vegetation, except possibly in the Far East where plant communi–

    ties alien to other parts of the Arctic are found.

            The circumpolar plants may be divided into three groups, viz., arctic

    plants occurring around the Arctic Sea and not reaching far to the south of

    the tundra zone; arctic-montane plants occurring also in the mountains far

    south of the arctic zone; and boreal plants occurring, like the arctic ones,

    around the Arctic Sea but reaching far down into the boreal forests. These

    are natural groups, each having its special early history. Most of the

    arctic and arctic-montane species have a continuous distribution from the

    western end of the zone to the eastern, but several arctic or arctic-montane

    species are lacking in the most arctic part.

            1. With No Gap in their Eurasiatic Area

            a. Circumpolar Arctic Plants


    Seashore Plants

    Puccinellia phryganodes sens. lat. Stellaria humifusa
    Carex incurve (with several races) Cochlearia officinalis arctica
    Carex subspathacea Matricaria ambigua


    Inland Plants

    Calamagrostis neglecta borealis Minuartia rubella

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    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    Deschampsia alpine (?) Sagina intermedia
    Deschampsia caespitosa borealis Ranunculus nivalis
    Poa abbreviata (high-arctic) Papaver radicatum sens. lat.
    Poa alpigena Draba cinerea
    Poa glauca Draba lacteal
    Colpodium fulum Braya purpurascens (continuous ?)
    Eriophorum angustifolium triste Chrysosplenium tetrandum
    Carex aquatilis stans Potentilla hyparctica (= emarginata )
    Carex bigelowii Epilobium arcticum (little known)
    Carex lapponica (distribution little known) Pyrola grandiflora
    Carex rariflora Vaccinium vitis-idaea minus
    Carex ursina (distribution little known) Ameria maritima labradorica
    Luzula arctica Polemonium boreale
    Luzula frigida Erigeron eriocephalus
    Luzula wahlenbergii Taraxacum arcticum (little known)
    Rumex arcticus Taraxacum hyparcticum

            b. Circumpolar Arctic-Montane Plants Without Large Gaps in

    Arctic Eurasiatic Branch


    Absent in Mountains of Central Europe

    Eriophorum brachyantherum Arctagrostis latifolia
    Eriophorum medium Phippsia algida
    Eriophorum russeolum Carex glacialis
    Hierochloe alpina Carex rotundata
    Poa arctica sens. lat. Carex saxatilis

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    Luzula arcuata Draba hirta (= glabella )
    Luzula arcuata confusa Draba nivalis
    Tofieldia pusilla Eutrema edwardsii
    Salix arctica Cardamine bellidifolia
    Koenigia islandica Saxifraga rivularis
    Melandrium apetalum Astragalus alpinus arcticus
    Ranunculus hyperboreus Cassiope tetragona
    Ranunculus sulphureus Pedicularis lapponica
    Draba alpina Artemisia borealis

            No less than 18 of these are known from the high-arctic Wrangel Island.

            The following species have a similar distribution but are less arctic:

    Luzula parviflora Pinguicula villosa
    Epilobium davuricum


    Present in Mountains of c C entral Europe

    Woodsia glabella Salix glauca sens. lat.
    Sparganium hyperboreum Salix reticulate sens. lat.
    Trisetum spicatum Polygonum viviparum
    Eriophorum scheuchzeri Oxyria digyna
    Kobresia myosuroides Minuartia biflora
    Carex bicolor Sagina linnaei
    Carex lachenalii Thalictrum alpinum
    Carex misandra (incl. fuliginosa ) Ranunculus pygmaeus
    Carex rupestris Draba fladnizensis
    Juncus biglumis Saxifraga cernua
    Juncus castaneus Saxifraga foliclosa

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    Saxifraga groenlandica sens. lat. Epilobium anagallidifolium
    Saxifraga nivalis Arctostaphylos alpina
    Saxifraga oppositifolia Gentiana tenella
    Sedum rosea sens. lat. Pedicularis sudetica


    Antennaria [ ?] carpathica

            Seventeen of these are known from Wrangel Island.

            The following species have a similar distribution but are less arctic:

    Carex angarae Rumex acetosa lapponicus
    Carex capitata Minuartia stricta


    c. Circumpolar Boreal Plants Without Large Gaps


    Species Reaching Far to Fairly into the Arctic

    Cystopteris fragilis Betula nana sens. lat.
    Lycopodium Selago Stellaria crassifolia
    Equisetum arvense Caltha palustris sens. lat.
    Equisetum scirpoides Ranunculus lapponicus
    Equisetum variegatum Cardamine pratensis
    Festuca rubra Saxifraga hirculus
    Festuca ovina sens. lat. Rubus arcticus
    Eriophorum angustifolium Rubus chamaemorus
    Carex chordorrhiza Empetrum nigrum sens. lat.
    Carex vaginata sens. lat. Vaccinium uliginosum
    Androsace septentrionalis Senecio congestus

            Only two of these, viz., Ranunculus lapponicus and Rubus arcticus , are

    lacking in central Europe.

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    Circumpolar Boreal Species Extending to Southern Part

    of Tundra Zone Only

    Equisetum fluviatile Rosa acicularis
    Equisetum pretense Hippuris vulgaris
    Equisetum silvaticum Callitriche autumnalis
    Lycopodium annotinum Epilobium angustifolium
    Carex brunnescens Epilobium palustre
    Carex capillaries Pyrola minor
    Coeloglossum viride Andromeda polifolia
    Ranunculus reptans Menyanthes trifoliata
    Parnassia palustris Linnaea borealis
    Potentilla palustris Adoxa moschatellina

    Companula rotundifolia sens. lat.

            d. Circumpolar Boreal Plants Reaching into the Arctic at Western

    End (European Russia) and Eastern End (Chukotsk Peninsula) of Tundra Zone Only

    but lacking in Middle Part

            Equisetum palustre (Kola, Kanin, Chukotsk Peninsula)

            Botrychium lunaria (Kola, Kanin, Kolguev, Kolyma)

            Lycopodium clavatum (Kola, Kolguev, Chukotsk Peninsula)

            Montia lamprosperma (Kola, Chukotsk Peninsula, subatlantic)

            Cornus suecica (Kola, Kanin, Kolguev, subatlantic)

            Fostera marina (Kola, Chukotsk Peninsula)

            Moneses uniflora (Kola, Kolguev, Chukotsk Peninsula)

            Galium boreale (Kola, Kanin, Chukotsk Peninsula)

            Lactuca siluiica (Kola, Kolyma, Eurasiatic)

    017      |      Vol_VI-0346                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

            2. With a Large Gap in their Eurasiatic Area

            These plants (possibly with the exception of Mertensia ) probably had a

    continuous distribution there in a warmer period but have been exterminated

    in the extreme North by the deterioration of the climate.

            a. Incompletely Circumpolar Seashore Plants

            Elymus arenarius (gap between E. arenarius s. str. And subsp. mollis,

    Vaigach to Chukotsk Pennsula)

            Carex glareosa (gap Novaya Zemlya to Chukotsk Peninsula)

            Carex mackenziei (gap Kolguev to Chukotsk Peninsula)

            Honckenya peploides sens. lat. (gap Yenisei to Chukotsk Peninsula)

            Potentilla egedii groenlandica (gap Kolguev to Chukotsk Peninsula)

            Lathyrus maritimus (gap Pechora to Kolyma)

            Mertensia maritima (gap Kola to Chukotsk Peninsula)

            Hippuris tetraphylla (gap Kanin to Kolyma)

            b. Incompletely Circumpolar Arctic Inland Plants

            Calamagrostis deschampsioides (gap from European Russian to Kolyma,

    Chukotsk Peninsula, and Anadyr, but probably very incompletely known)

            Potentilla pulchella (gap from Novaya Zemlya to Wrangel Island)

            Chrysanthemum arcticum Polaris (gap from Yalmal to Chukotsk Peninsula)

            c. Arctic-Montane Circumpolar Plants Incomplete in Arctic

    Eurasiatic Branch

            Lycopodium alpinum (gap Yenisei to Chukotsk Peninsula)

            Agrostis borealis (gap the Urals to Lena River)

            Poa alpina (gap Yenisei to Chukotsk Peninsula)

            Carex atrofusca (gap Kola to Chukotsk Peninsula and Anadyr, but

    incompletely known)

    018      |      Vol_VI-0347                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

            Carex nardina (gap Franz Josef Land to Chukotsk Peninsula)

            Luzula spicata (gap the Urals to Chukotsk Peninsula)

            Silene acaulis (gap the Urals to Novaya Zemlya to Chukotsk Peninsula)

            Sibbaldia procumbens (gap Yenisei to Chukotsk Peninsula)

            Rhododendron lapponicum (gap Scandinavia to Yana River)

            Loiseleuria procumbens (gap the Urals to Chukotsk Peninsula)

            Phyllodoce coerulea (gap the Urals to Novaya Zemlya to Chukotsk Peninsula)

            Campanula uniflora (gap Novaya Zemlya to Chukotsk Peninsula)

            Erigeron humilis (= unalaschcensis ) (gap Vaigach to Chukotsk Peninsula)

            The circumpolar component in the arctic flora thus consists of about 180



            1. Eurasiatic Arctic Plants

    Phippsia concinna Salix reptans

            2. Eurasiatic Arctic-Montane Plants (Many occurring also in Alaska )

    Salix hastate Primula sibirica (in Arctic

    European Russia and Chukotsk

    Peninsula only)
    Salix lanata
    Salix nummularia Eritrichium villosum
    Salix polaris Myosotis alpestris
    Dryas octopetala punctata Pedicularis cederi
    Hedysarum obscurum sens. lat. Pedicularis verticillata
    Oxytropis campestris sordida Valeriana capitata
    Viola biflora (in the southern

    part only)
    Saussurea alpina sens. lat.
    Taraxacum glabrum

    019      |      Vol_VI-0348                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

            3. Eurasiatic Boreal Plants (Species reaching far into the Arctic )

    Calamagrostis lapponica Rumex graminifolius
    Eriophorum v aginatum (E. spissum exluded) Polygonum bistorta
    Veratrum album (subsp. lobelianum and

    subsp. oxycephalum )
    Chrysanthemum bipinnatum
    Allium schenoprasum sibiricum Petasites frigidus

            4. Eurasiatic Boreal Plants Reaching Southern Part of Tundra Zone Only

    Carex globularis Conioselinum vaginatum
    Carex heleonastes Pedicularis sceptrum carolinum
    Carex rostrata Veronica longifolia
    Salix myrtilloides Aster sibiricus
    Salix Pyrolifolia (entering the Arctic

    at Pechora and Chukotsk Peninsula only)
    Chrysanthemum vulgaris
    Thymus serpyllum sens. lat. Senecio intergrifolius
    Viola epipsila (in Arctic European

    Russia and Chukotsk Peninsula only)
    Erigeron acris asteroides (introduced)


            A number of species occur neither in the western nor in the eastern end

    of the arctic zone but only in its middle part. They are central Siberian or

    Eurasiatic continental species. In the Arctic they might be relicts from a

    warmer period when part of the continental components in the Eurasiatic flora

    were able to protrude farther northward.


            Bromus sibiricus (arctic Urals)

            Agropyron boreale (arctic Siberia)

            Carex redowskyana (lower Ob, Khatanga, and Lena rivers)

    020      |      Vol_VI-0349                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

            Carex sabynensis (arctic Urals, mouth of Yenisei)

            Salix dasyclados (distribution unclear)

            Delphinium cheilantham (arctic Urals, mouth of Kolyma)

            Delphinium elatum (Kanin to Kolyma, only in southern parts)

            Trollius asiaticus (Vaigach, lower Yenisei and Lena rivers)

            Sorbus sibirica (arctic Europe and Siberia)

            Astragalus australis (= A. uralensis ) (arctic Siberia)

            Geranium albiflorum (arctic Eruopean Russia, arctic Urals)

            Rhododendron anthopogon (arctic Urals, mouth of Lena River)

            Pedicularis compacta (lower Pechora and Yenisei rivers)

            Chrysanthemum sibiricum (Taimyr)


            A number of central Asiatic species or species with wide southern dis–

    tribution have apparently been washed down from central Asia along the great

    rivers, especially along the Yenisei, since they occur along rivers far north

    of their compact area of distribution. Probably several others just reach

    the southern limit of the Arctic at the mouths of the Yenisei and Lena rivers.


    Found at Mouth of Yenisei River

    Polygonum tomentosum Angelica decurrens
    Stellaria bungeana Limosella aquatic
    Cardamine macrophylla Pedicularis uncinata
    Viola uniflora Campanula glomerata
    Henacleum dissectum Achillaea impatiens

    021      |      Vol_VI-0350                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

            Close to Mouth of Yenisei (Doubtful Whether They Reach the Arctic)

    Potamogeton pectinatus Potamogeton pusillus (also in

    Kola Peninsula)
    Potamogeton perfoliatus Phalaris arundinacea
    Potamogeton praelongus Carex rostrata


    Found at Mouth of Lena River

    Braya aenea Caragana jubata (entering

    into the Arctic ?)


    Swertia perennis


    Found at Mouth of Kolyma River


    Beckmannia zizygachne


            In the foregoing pages have been enumerated those plants which occur all

    along the arctic belt, at both ends of it, or [ ?] else along the middle part of


            Many plants, however, occur within the arctic belt in its western or in

    its eastern part only. The reason for this is either that they belong to

    western (European, amphiatlantic) or to eastern (Eastern Asiatic, American)

    groups, or that they are more southern species of circumpolar or Eurasiatic

    groups which can enter into the tundra zone at its least arctic parts only,

    viz., in the European Arctic or in the Arctic of the Far East. As it is of

    great phytogeographical interest to know the order in which these species dis–

    appear as one moves from Kola Peninsula eastward or from Chukotsk Peninsula

    westward, and as no such enumeration has previously existed, lists of them

    are given here, starting with the European series and concluding with the

    Eastern Asiatic.

    022      |      Vol_VI-0351                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic


            1. In the Arctic Only in Kola Peninsula

            a. Occurring in More Arctic Parts


    Circumpolar Species

    Cystopteris montana (arctic-montane) Carex salina
    Dryopteris austriaca Carex vesicaria
    Dryopteris linnaeana Juncus filiformis
    Dryopteris Phaegopteris Stellaria calycantha (arctic-montane)
    Potamogeton filiformis Subularia aquatic
    Triglochin palustre Trifolium pratense (introduced)
    Milium effusum Ligusticum scoticum
    Poa annua (introduced) Rhinanthus minor groenlandicus
    Poa nemoralis Utricularia minor


    European or Eurasiatic Plants

    Pinus silvestris Rubus saxatilis
    Calamagrostis purpurea Vicia cracca
    Carex juncella Callitriche polymorpha
    Orchis maculata Viola montana
    Gymandenia canopsea Viola palustris
    Rumex pseudonatronatus Anthriscus silvestris
    Stellaria nemorum Melampyrum pratense


    Hieracium to the group tridentate

    Other groups

            Deschampsia atropurpurea

            Alchemilla glomerulans (amphiatlantic)

    023      |      Vol_VI-0352                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

            b. Occurring in Less Arctic Eastern and Western Ends of Kola



    Circumpolar Species

    Equiaetum silvaticum Carallorhiza trifida
    Juncus gerardi Populus tremula
    Lazula multiflora Pyrola secunda


    European or Eurasiatic Species

    Luzula pilosa Vicia sepium
    Salix caprea Melampyrum silvaticum
    Melandrium rubrum Plantago media
    Ranunculus auricomus Matricaria inodora
    Potentilla erecta Tussilago farfara

            c. Occurring at Ponoi or Ponoi-Yokanga at Southern Edge of

    Arctic Zone


    Circumpolar Species

    Lycopodium complanatum Carex disperma
    Botrychium boreale Carex livida
    Athyrium filix-femina Carex limosa
    Dryopteris filix-mas Juncus alpinus alpestris (arctic-montane)
    Polypodium vulgare Alnus incana
    Potamogeton pusillus Chenopodium album (Introduced)
    Phalaris arundinacea Stellaria longifolia
    Nardus stricta Sagina procumbens
    Agropyron labiglume eurasiaticum

    Silene cucubalus
    Agropyron mutabile (arctic-montane) Anemone nemorosa

    024      |      Vol_VI-0353                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    Capsella bursa - pastoris (intoduced) Galeopsis bifida (introduced)
    Rorippa islandica Veronica scutellata (introduced)
    Trifolium repens (introduced) Veronica serpyllifolia (introduced)
    Callitriche verna Plantago major (introduced)
    Drosera rotundifolia Galium palustre
    Epilobium lactiflorum (arctic-montane) Galium trifidum
    Arctostaphylos uva - ursi Sonchus arvensis


    European or Eurasiatic Species

    Athyrium crenatum Ranunculus sceleratus reptabundus
    Calmagrostis epigeios Erysimum hieracifolium
    Melica nutans Fumaria officianalis
    Molinia coerulea Prunus padus
    Poa remota Cotoneaster melanocarpus
    Poa trivialis Vicia silvatica
    Agropyron caninum Geranium pratense
    Carex heleonastes Daphne mezereum
    Carex loliacea Epilobium alsinifolium (arctic-montane)
    Carex caespitosa Angelica silvestris
    Carex gracilis Calluna vulgaris
    Carex globularis Myosotis silvatica gfrigida (arctic-montane)
    Gegea lutea Myosotis sparsiflora
    Paris quadrifolia Euphrasia brevipila
    Nuphar pumilum [ ?]Calium uliginosum
    Thalictrum simplex boreale Senecio nemorensis jacquinianus
    Actaea erythrocarpa Crepis paludosa

    Crepis tectorum

    025      |      Vol_VI-0354                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic


    Other Groups

    Bromus inermis (central Asiatic) Paeonia anomala intermedia (central

    Urtica dioeca gracilis (American) Ranunculus peltatus
    Aconitum septentrionale (central Asiatic) Rosa cinnamomea (central Asiatic)

    Gentiana nivalis (amphiatlantic)

            d. Occurring in Western Part of Kola Peninsula

    Alopecurus aequalis (circumpolar) Carex flava (circumpolar)
    Carex goodenowii (circumpolar) Luzula pallescens (circumpolar)
    Carex arctogena (amphiatlantic) Betula callosa
    Carex atrata (circumpolar, arctic-montane) Polygonum raji norvegicum (European)

    Alchemilla alpina (amphiatlantic)

            2. In the Arctic from Kola Peninsula to Arctic Russian Mainland

    Puccinellia distans pulvinata Carex microglochin
    Puccinellia maritima Sorbus aucuparia glabrata
    Carex magellanica

            3. In the Arctic from Kola Peninsula to Kanin Peninsula


    Boreal Circumpolar Species

    Triglochin meritimum Lemna trisulca
    Hierochloe ordrata Salicornia europaea
    Carex canescens Sagina nodosa
    Carex pauciflora Callitriche autumnalis
    Lemna minor Hieracium to the group vulgtiformia


    Circumpolar Arctic-Montane Species

    Athyrium alpestre Selaginella selaginoides

    Epilobium hornemannii (incompletely known)

    026      |      Vol_VI-0355                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic


    Other Groups

    Carex halleri (amphiatlantic) Gnaphalium norvegicum (amphi–

    atlantic, also in middle Urals)
    Viscaria a lpina (amphiatlantic)
    Barbarea stricta (Eurasiatic, from Kolyma

    to Anaydr Bay orthoceras
    Cirsium heterophylllum (north

    Cenolophium fischeri (Eurasiatic) Ligularia sibirica (Eurasiatic)

            4. In the Arctic from Kola Peninsula to Kanin and Kolguev

    Agrostis stolonifera (boreal circumpolar) Angelica archangelica (north European)
    Juncus arcticus (circumpolar arctic-montane) Gentiana aurea (amphiatlantic)
    Salix lapponum (north European, also on

    the mainland)
    Lomatogonium rotatum (circumpolar

    arctic-montane, also on the mainland)
    Salix phylicifolia (north European, also

    on the mainland)
    Euphrasia frigida (amphiatlantic)
    Rumex acetosa (circumpolar arctic-montane) Antennaria dioica (Eurasiatic)
    Aconitum septentrionale (central Asiatic) Gnaphalium supinum (amphiatlantic)
    Geum rivale (boreal circumpolar) Fetasites laevigatus (central Asiatic)
    Alchemilla glemerulans (amphiatlantic) Solidago virgaurea (Eurasiatic)

            5. In the Arctic from Kola Peninsula to Urals

    Deschampsia fletuosa (several separate

    Trollius europaeus (north European)
    Carex adelestoma (area little known) Fillipendula ulmaria (north European)
    Dianthus superbus (Eurasiatic) Veronica alpina (circumpolar arctic–


    Pinguicula vulgaris (circumpolar)

    027      |      Vol_VI-0356                                                                                                                  
    E A-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

            6. In the Arctic from Kola Peninsula to Novaya Zemlya or Vaigach


    Amphiatlantic Plants

    Carex paralella Draba norvegica (= rupestris)
    Stellaria crassipes Arabis alpina (also in the

    Kara tundra)
    Cerastium alpinus sens. str.
    Cerastium trigynum Saxifraga stellaris
    Arenaria pseudofrigida Bartia alpina
    Ranunculus glacialis sens. str. Taraxacum brachyceras (= simulum)


    Other Groups

    Alopecurus ventricosus (group doubtful) Elymus arenarius sens, str. (north

    Phleum alpimum commutatus (circumpolar

    Juncus triglumis (circumpolar

    Puccinellia retroflexa suecica

    (north European)
    Primula stricta sens. str. (amphi–

    atlantic, in Siberia P. stricta

    Puccinellia distans (Eurasiatic)

            7. In the Arctic from Kola to Lower Ob River

    Juncus trifidus (amphiatlantic) Geranium silvaticum (north European)
    Salix herbacea (amphiatlantic) Alchemilla murbeckiana (north European)
    Melandrium furcatum angustiflorum

    (arctic European)
    Vaccinium myrtillus (north European)
    Ribes rubrum (Eurasiatic) Castilleja pallida arctica (endemic)

    Plantago maritima sens. lat. (several separate areas, goes to Yalmal)

            8. In the Arctic from Kola to Yenisei

    Pices abies obovata (central Asiatic) Larix sibirica (central Asiatic)
    Juniperus communis (boreal circumpolar) Scripus caespitosus austriacus

    (boreal circumpolar)

    028      |      Vol_VI-0357                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    Carex dioeca (Eurasiatic) Myosotis palustris (Eurasiatic)
    Salix arbuscula (Eurasiatic arctic-montane) Ledum palustre sens. str. (north

    European, east of Yenisei

    subsp. decumbens)
    Salix myrsinites (Eurasiatic arctic-montane)
    Salix nigricans (North European, only along

    southern rim of the Arctic)
    Cassiope hypnoides (amphiatlantic)
    Betula tortuosa (Eurasiatic arctic-montane) Diapensia lapponica sens. str.

    (amphiatlantic, in the east

    subsp. obovata)
    Stellaria graminea (Eurasiatic) Euphrasia frigida (amphiatlantic)
    Stellaria palustris (central Asiatic) Pinguiculs alpina (Eurasiatic


            9. In the Arctic from Kola to Taimyr

            Potentilla crantzii (amphiatlantic)

            10. In the Arctic from Kola to Lena

    Creastium regelii (arctic Eurasiatic) Astragalus norvegicus (Eurasiatic


    Pedicularis hirsuta (amphiatlantic, ? in Chukotsk Peninsula)


            1. In the Arctic in Chukotsk Peninsula only


    Boreal Circumpolar Plants

    Woodsia ilvensis Potentilla fruticosa


    Circumpolar Arctic-Alpine Plant

            Gentiana Prostrata


    Eastern Asiatic Inland Plants

    Larix dahurica Salix krylovii
    Carex novograblenovii Claytonia eschscholtzii
    Salix berberifolia Stellaria dehurica

    029      |      Vol_VI-0358                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    Saxifraga dahurica Oxytropis adamsiana
    Saxifraga nudicaulis Pinguicula glandulosa

    Saussurea tilesii


    Eastern Asiatic Coastal Plants

    Carex scita karaginensis Potentilla fragiformis
    Trollius membranestylic Cnidium ajanense
    Draba eschscholtzii Rhodedendron kamtschaticum

    Ermania parryeides
    Dicentra peregrinus Aster consanguineus
    Oxytropis revoluta Artemisia glomerata


    Plants Confined to Bering Strait Area

    Colpodium vrightii Saxifraga dahurica grandipetala
    Stellaria dicrancides Chrysosplenium wrightii
    Minuartia elegans Eritrichium chamissonis
    Smelovskia calycina integrifolia Artemisia senjavinensis


    Pacific Plants

            Several of the species belonging to this group are apparently rare on

    the Chukotsk Peninsula. They are probably relicts from a warmer period, during

    which they had a more northerly area on the shores of the Bering Sea.

    Deschampsia beringensis Draba borealis
    Poa komarowii Potentilla villosa
    Poa arctica williamsii Saxifraga bracteata
    Carex anthoxanthea Geranium erianthum
    Carex macrochaeta Angelica lucida
    Carex lyngbyaei cryptocarpa Phyllodoce aleutica

    030      |      Vol_VI-0359                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    Primula cureifolia saxifragifolia Trientalis europaea arctica
    Primula tschuktschorum Artmisia globularia

    Taraxacum kamtschaticum


    Eastern Asiatic-Western American Plants

    Selaginella schmidtii Saxifraga eschscholtzii
    Selaginella sibirica Saxifraga unalaschcensis
    Carex montanensis Spiraea beauverdiana
    Salix alaxensis Potentilla biflora
    Salix phlebophylla Geum rossii
    Salix reticulata orbiculata Rubus stellatus
    Claytonia sarmentosa Campanula laciocarpa
    Anemone narcissiflora sibirica Antennaria monocephala
    Draba densifolia Arnica lessingii
    Alyssum americanum Saussurea angustifolia
    Saxifraga bronchialis funstonii Saussurea nuda

    Taraxacum lateritium


    American Plants

    Elymus arenarius mollis Stellaria monantha
    Poa leptocoma Silene stenophylla
    Bromus pumpellianus arcticus Thalictrum sparsiflorum
    Carex circinata Cardamine purpurea
    Carex gynocrates Arabis lyrata kamtschatica
    Carex membrancea Saxifraga exilis
    Carex scripoides Parnassia kotzebuei

    031      |      Vol_VI-0360                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    Potentilla vahliana Dodecatheon frigidum
    Dryas integrifolia Gentiana propinqua
    Oxytropis bellii Solidago multiradiata
    Oxytropis maydeliana Chrysanthemum integrifolium
    Conioselinum benthamii Antennaria ekmanniana

    Scencio pseudo-arnica

            2. In the Arctic from Chukotsk Peninsula to Kolyma


    Northeastern Asiatic Plants

    Calamagrostis kolymensis Rhododendron chrysanthemum
    Arenaria tschuktchorum Dracocephalum palmatum


    Eastern Asiatic-Western American Plants

    Selaginella sibirica Androsace ochotensis
    Aconitum delphinifolium Arnica louiseana frigida


    American Plants

    Carex stylosa Pyrola asarifolia incarnata

            3. In the Arctic from Chukotsk Peninsula to Lena River


    Arctic-Montana Plants

    Kobresia schoenoides (Eurasiatic) Potentilla elegans (Asia and

    Salix torulosa
    Oxytropis glacialis Aster alpinus (Eurasia, America)
    Corydalis pauciflora (Asia and Alaska) Crepis nana (Asia, America)


    Eastern Asiatic Plants

    Carex lenaensis Claytonia acutifolia (also in

    Carex vanheurkii
    Polygonum pawlovskianum Draba juvenilis

    032      |      Vol_VI-0361                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    Draba pseudopilosa (also in Alaska) Saxifraga redowskiana

    Primula stricta jakutensis (= egalikensis ?)


    Eastern Asiatic-Western American Plants

    Calamagrostis purpurascens arctica Cardamine richardsonii
    Puccinellia hauptiana Draba barbata
    Carex lugens (= soczaweana ) Draba caesia (= paladeriana )
    Salix arbutifolia (= fuscescens ) Potentilla uniflora
    Salix chamissonis Primula borealis
    Merckia physodes Gentiana acuta
    Delphinium brachycentrum Gentiana glauca
    Cardamine microphylla Taraxacum sibiricum


    American Plants

    Anemone richardsonii Hedysarum mackenziei


    4. In the Arctic from Chukotsk Peninsula to Taimyr


    Arctic-Montane plants

    Saxifraga serpyllifolia Artemisia trifurcata


    North Asiatic Plants

    Elymus interior ( villosissimus of

    Russian authors)
    Oxytropis middendorffii
    Cal Claytonia tuberosa (also in Alaska) Pedicularis adamsii
    Oxytropis mertensiana (also in Alaska) Petasites gmelinii (also in the

    central Urals)


    Eastern Asiatic-American Plants

    Eriophorum callitrix (to Khatanga River) Diapensia lapponica obovata
    Lesquerella arctica Pedicularis langsdorffii (incl.

    P. arctica )
    Ribes triste

    Artemisia arctica

    033      |      Vol_VI-0362                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic


    5. In the Arctic from Chukotsk Peninsula to Yeninsei River


    Arctic Asiatic-American Plants

    Festuca altaica (arctic-montane) Alyssum americanum (Alaska only

    in America)
    Betula nana exilis
    Polygonum laxmannii (arctic-montane) Oxytropis leucantha
    Claytonia arctica (to Norilsk Mts.) Oxytropis nigrescens
    Melandrium furcatum sens. str. Oxytropis roaldi (little known)
    Melandrium taylorae Ladum palustre decumbens
    Ranunculus pygmaeus sabinei Eritrichium aretioides (in America

    west of Mackenzie River)
    Draba pilosa (nearly circumpolar) Pedicularis capitata (nearly

    Castilleja pallida elegans Boschniskia rossica


    Eurasiatic Plant

            Androsace filiformis (southern, farther westward)


    6. In the Arctic from Chukotsk Peninsula to Ob River


    Asiatic-American Plants

    Carex bonanzensis Descurainia sophioides

    Pedicularis labradorica


    7. In the Arctic from Chukotsk Peninsula to the Urals


    Arctic or Arctic-Montane Plants

    Dryopteris fragrans (circumpolar,

    Potentilla stipularis (Asiatic)
    Carex melanocarpa (Asiatic) Geum glaciale
    Tofieldia nutans (nearly circumpolar) Pyrola secunda obtusata (nearly

    Rumex sibiricus (also in Alaska) Pedicularis lanata (nearly

    Minuartia arctica (also in Alaska)
    Saxifraga punctata (arctic-montane,

    also in America)
    Crepis chrysantha (Asiatic,


    034      |      Vol_VI-0363                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic


    8. In the Arctic from Chukotsk Peninsula to Novaya Zemlya


    Arctic Plants

    Calamagrostis holmii (Asiatic, also in Alaska) Draba macrocarpa (nearly circumpolar)
    Koeleria asiatica (Asiatic) Draba micropetala (nearly circumpolar)
    Puccinellia langeana asiatica (Asiatic) Draba oblongata (nearly circumpolar)
    Salix pulchra (also in Alaska) Draba pauciflora (nearly circumpolar)
    Salix rotundifolia (also in Alaska) Draba subcapitata (nearly circumpolar)
    Minuartia macrocarpa (also in Alaska) Hesperis pallasii (nearly circumpolar)
    Ranunculus affinis (nearly circumpolar) Androsace triflora (Asiatic)


    Arctic-Montane Plants

    Festuca brevifolia Saxifraga flagellaris
    Festuca supine Androsace chamae jasme lehmanniana
    Lloydia serotina Senecio atropurpureus (“frigidus”)
    Stellaria longipes Senecio resedifolius

    Cerastium maximum

            9. In the Arctic from Chukotsk Peninsula to Arctic Russian Mainland

            Hierochloe pauciflora (nearly circumpolar, arctic)

            Trisetum sibiricum litorale (also in Alaska)

            Alnus crispa (to Mezen, nearly circumpolar)

            Dianthus repens (to Pechora, also in Alaska)

            Draba sibirica (to Pechora, Asiatic)

            Epilobium latifolium (circumpolar arctic-montane)

            Cortusa mathioli (Eurasiatic)

            Pedicularis amoena (to Bolshezemelskaia tundra, Asiatic)

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            10. In the Arctic from Chukotsk Peninsula to Kolguev

            Festuca brachyphlla (nearly circumpolar)

            Calamagrostis canadensis langsdorffii (also in Alaska)

            Stellaria ciliatosepala (nearly circumpolar)

            11. In the Arctic from Chukotsk Peninsula to Kanin


    Arcctic Plants

            Alopecurus alpinus

            Dupontia fischeri (nearly circumpolar)

            Deschampsia arctica (nearly circumpolar)

            Cerastium berringianum (also in Alaska)

            Astragalus umbellatus (also in Alaska)

            Lagotis stelleri (Eurasiatic)

            Artemisia tilesii (also in Alaska)


    Arctic-Montane Plants

            Parrya nudicaulis (also in western America)

            Ligusticum nutellinoides alpinum (also in Alaska)

            12. In the Arctic from Chukotsk Peninsula to Kola Peninsula (but

    lacking in Scandinavia )


    Arctic-Montane Plants

    Salix arctica Myosotis alpestris
    Eutrema edwardsii Pedicularis sudetica
    Hedysarum obscurum sens . lat. Pedicularis verticillata
    Lomatogonium rotatum Valeriana capitata

    036      |      Vol_VI-0365                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic


    Arctic Circumpolar Plants

    Rumex arcticus Vaccinium vitis - idaea minus
    Ranunculus pallasii


    Northern Eurasiatic Plants

    Salix reptans Chrysanthum bipinnatum

            IV. ENDEMIC PLANTS

            Numerous plants are endemic in the Soviet Arctic, that is, they occur

    nowhere else. However, the endemism is not very strongly marked, as most of

    them differ only slightly from close relatives in the arctic or boreal belt.

    These plants in many cases should rather be regarded as arctic races of more

    widespread species, but the present tendency to recognize very small systematical

    units as species causes most authors to quote them as distinct species.

            An enumeration of the endemics of the Soviet Arctic is given below.

    They are roughly divided into four geographical groups.

            1. Endemics with Wide Distribution Within the Soviet Arctic or Even

    Occurring Somewhat Outside It

            Trisetum sibiricum litorale (arctic European Russia to Chukotsk Peninsula,

    also in Alaska)

            Calamagrostis holmii (Novaya Zemlya to Alaska, related to C . d eschampsioides )

            Puccinellia sibirica (lower Yenisei and 0b Rivers)

            Salix reptans (Kola peninsula to Anadyr and Kamchatka)

            Cerastium regelii (Kola Peninsula to Lena River; also on Spitsbergen, related

    to C. alpinum )

            Silene pauciflora (arctic Europe and Siberia, related to S. graminifolia)

            Ranunculus hyperboreus samojedorum (Kolguev to Chukotks Peninsula; also

    in Alaska)

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            Arabis septentrionalis (Novaya Zemlya to Kolyma, related to A. petraea )

            Draba glacialis (arctic Europe and Siberia, related to D. alpina )

            Oxytropis campestris sordida (Kola to Chukotsk Peninsula)

            Hedysarum arcticum (race of H. obscurum )

            Arnica alpina iljini (Novaya Zemlya to Chukotsk Peninsula)

            2. Endemics Occurring in Most Arctic Central Parts Only

            Poa taimyrensis (Taimyr, related to P. lanatiflora )

            Poa tolmatchevii (Taimyr, related to P. arctica )

            Lynchnis villosula (Taimyr to Lena River, related to L. sibirica )

            Melandrium taimyrense (Taimyr, related to M. furcatum )

            Caltha caespitosa (Novaya Zemlya to Taimyr, related to C. palustris )

            Papaver pulvinatum (Taimyr, related to P. radicatum )

            Draba parvisiliquosa (Taimyr, related to D. cineres )

            Draba pohlei (Taimyr, related to D. alpina )

            Oxytropis sverdrupi (Taimyr, little known)

            Senecio taimyrensis (Gorodkov, unpublished)

            3. Endemics Occurring in Western Parts of Soviet Arctic

            Koehleria gorodkovi (Gydan tundra)

            Kochleria pohleana (Kolguev, related to K. glauca )

            Calamagrostis steinbergii (Northern Island of Novaya Zemlya)

            Puccinellia jenisejensis (= Atropis jenisejensis Roshew ) (lower Yenisei River)

            Poa petschorica (Pechora, related to P. arctica )

            Carex bucculenta (Kanin, Novaya Zemlya, race or form of C. incurva )

            Carex selina (western Siberia, race or form of C. incurva )

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            Lychnis samojedorum (Pechora, arctic Siberia, related to L. sibirica )

            Melandrium furcatum angustifolium (Scandinavia to Ob River)

            Ranunculus glabriusculus (race of P. acris )

            Papaver angustifolium (Gydan tundra, lower Yenisei, related to P. radicatum )

            Draba kjellmanii (Novaya Zemlya and Vaigach; also in Spitsbergen, related to

    D. alpina )

            Rhodiola arctica (Kola Peninsula to Novaya Zemlya, in [se ?] parable from R. rosea ?)

            Sorbus polaris (lower Ob River, variety of S. sibirica )

            Astragalus grigorievii (Kanin Peninsula, related to A. umbellatus )

            Epilobium tundrarum (Vaigach to Taimyr, related to E. palustre )

            Castilleja pallida arctica (Kola Peninsula to Yenisei)

            Taraxacum brevicorne (Novaya Zemlya to Yenisei)

            Taraxacum hjeltii (Scandinavia to Kanin Peninsula)

            Taraxacum nova zemliae (Novaya Zemlya (= T. brachyceras ?))

            Taraxacum platyphyllum (Novaya Zemlya)

            4. Endemics Occurring in Eastern Part of Soviet Arctic

            Calamagrostis kolymaensis (Kolyma to Chukotsk Peninsula, related to C. neglecta )

            Calamagrostis bungeana (Yana River, related to C. neglecta )

            Koehleria seminude (lower Lena River, related to K. cristata )

            Puccinellia laeviuscula (Chukotsk Peninsula; also in Alaska)

            Carex arakamensis (Chukotsk Peninsula, related to C. holostoma )

            Monolepis asiatica (Yenisei to Kolyma, related to M. nuttalliana )

            Cerastium bialynichii Cerastium bialynichii (Yenisei to Chukotsk Peninsula; also in Alaska?, related

    to C. alpinum )

            Arenaria tschuktchorum (Kolyma to Chukotsk Peninsula, related to A. capillaris )

    039      |      Vol_VI-0368                                                                                                                  
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            Trollius chartosepalus (Chukotsk Peninsula)

            Draba chamissonis (Chukotsk Peninsula, related to D. nivalis )

            Saxifraga redowskii (Lena to Kolyma, related to S. stellaris )

            Rhodiola borealis (Henisei to Chukotsk Peninsula, doubtfully distinct from

    R. rosea

            Oxytropis middendorffii (Taimyr to Kolyma, related to O. schmidtii and

    O. leucantha )

            Oxytropis schmidtii (Yenisei to Kolyma, related to O. middendorffii and

    O. leucantha

            Artemisia laciniatiformis (Chukotsk Peninsula, related to A. laciniata )

            Taraxacum macilentum (Yenisei to Lena River)

            Taraxacum longicorne (lower Lena River)

            Taraxacum macrodens (lower Lena River)


            A few species have such a singular distribution that for the present

    at least, they cannot be grouped with other plants. They are enumerated below.

            Pleuropogon sabinei : Novaya Zemlya, mouth of Lena River, Wrangel Island,

    Altai, Greenland, northeastern America (arctic-montane species with several

    large gaps in area)

            Colpodium vahlianum (= Puccinellia kjellmanii): Novaya Zemlya, Wrangel

    Island, Spitsbergen, Greenland (circumpolar?)

            Agropyron angustiglume : Arctic Urals, northern Europe, central Asiatic

    mountains, Kamchatka

            Torularia humilis : Mouth of Lena and Yana Rivers, the Himalayas, moun–

    tains of China and Korea, Altai, Alaska, Newfoundland, Greenland (arctic–

    montane with large gaps)

    040      |      Vol_VI-0369                                                                                                                  
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            Braya linearis : Scandinavia, Pechora, Vaigach, mouth of Lena River,

    eastern America (area very little known)

            Centiana verna : Kanin, Kolguev, Samoyed tundra in the Arctic, then in

    central and southern Europe, in England, Oreland, Caucasus and central

    Asiatic mountains (Eurasiatic arctic-montane)

            Artemisia richardsoniana : Taimyr, arctic America (distribution but

    little known)

            Antennaria alpina : Circumpolar type divided into spomictic types -–

    A. alpina s. str., Scandinavia, Kola, Kanin, Greenland, Iceland


    Origin and Composition of Arctic Flora

            The circumpolar arctic and boreal species form the core of the arctic

    flora. They constitute the chief components in most plant communities. Their

    origin is obscure, as they are plants with a marked ability to spread and

    often with large ecological amplitude. Gaps in their area caused by climatic

    or other changes during their earlier history are rapidly filled up when bet–

    ter conditions permit, and they then penetrate to their climatic and ecological

    limits. They must be plants that occurred in the Tertiary period in the

    northern zone of the earth.

            The arctic-montane group must have had a different history. They are

    Tertiary mountains plants which survived the glacial periods on plains or on

    mountains in unglaciated areas, chiefly in easternmost Asia and in unglaciated

    Alaska, and spread eastward and westward from that center during warmer periods.

    Most of them are more or less circumpolar, while a few are Eurasiatic. Many

    of them show large gaps in their area, proving that the drought during warm

    periods or other changes in the climatic or ecological conditions exterminated

    041      |      Vol_VI-0370                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    them from part of their area and that they have not been able subsequently to

    fill the gaps. A detailed study of their interesting and diverse areas will

    give the key to an interpretation of their history.

            Numerous species bordering the shores of both the Atlantic and the Pacific

    occur. In Europe they form the rich Atlantic and sub-Atlantic components in

    the flora; in eastern America it is for the most part other species that form

    corresponding groups. In the Pacific the flora is to a large extent the same

    on the borders of the Asiatic and on American shores. In no case do the

    European Atlantic and sub-Atlantic plants reach the Arctic (possibly with

    the exception of Cornus suecica and Montia lamprosperma ). In the Pacific the

    conditions are quite the opposite. Numerous plants bordering the American

    and Asiatic shores occur in the Chukotsk Peninsula, and even farther north

    in Wrangel Island. Many of them are found far to the west in the Arctic,

    as will be seen from the lists of distribution. The reason for this is doubt–

    less that the Pacific was very little glaciated during the glacial periods,

    and plants which have been exterminated elsewhere were thus able to survive

    these glaciations and spread when the amelioration of the climate set in.

            The plants occurring only in the western part of the Eurasiatic Arctic

    are mostly boreal circumpolar or boreal Eurasiatic species which barely reach

    into the arctic zone in its least arctic parts. A few amphiatlantic plants

    also belong here. They would seem to be remnants of formerly circumpolar

    species which died out in the Pacific. This view, however, is opposed by some

    other phytogeographers, who consider them to be species of a sunken North

    Atlantic land bridge. Most amphiatlantic species extend to Novaya Zemlya.

            A singular position in the Arctic is occupied by Cornus suecica , and

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    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    Montia lamprosperma which occur both in the European part and in the Chukotck

    Peninsula. They may be said to be circumpolar subocean species, as they occur

    on all the four ocean shores of the boreal belt.

            Central Asiatic or Eurasiatic plants occur for the most part on the

    southern border of the Arctic. They play no part in the vegetation. They

    are either relicts from a warmer period or plants whose present ecological

    amplitude allows them to penetrate northward into the tundra zone.

            The endemics have already been dealt with in the above pages.

            Finally it may be emphasized that ferns are very rare. Cystopteris

    fragilis, Woodsia ilvensis, W. glabella, Athyrium alpestre, Cryptogramma

    stelleri, Dryopteris fragrans
    , and D. robertiana are present, in Kola

    Peninsula still others occur, but some in single localities only. Annuals

    are lacking, except for Koenigia , which, however, might be expected to be

    biennial in the Arctic. Geophytes are extremely rare as the bulbs cannot

    stand the frozen soil. Lloydia serotina is the only real arctic geophyte.

    Allium schoenoprasum sibiricum and Coeloglosum viride occur, however, on the

    southern rim of the tundra zone.

            The largest genus in the Arctic is Carex , with about 60 species, next

    comes Salix with 35, Draba with 31, Saxifraga with 22, and Poa , Potentilla ,

    and Pedicularis with 16 each.



            The vegetation of the Arctic is fairly uniform as most of the species

    that play a large part in the prevailing plant communities belong to the

    circumpolar group of distribution. The vegetation of the Chukotsk Peninsula

    is the most prominent exception to this rule. As only very limited parts of

    043      |      Vol_VI-0372                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    the Arctic have been well investigated phytogeogrphically, only a somewhat

    schematic review of the conditions is possible. A general characterization

    of the main vegetational zones will first be given, mainly according to the

    system of Gorodkov. This will be followed by a short description of the

    different geographical provinces of the Arctic, starting in the west with

    the Kola Peninsula and ending in the east with the Chukotsk Peninsula.


    Zones of Vegetation

            The snow zone comprises the Franz Josef Land Archipelago, the northern

    island of Novaya Zemlya except the Matochkin Shar district, Severnaya

    Zemlya north of Cape Chelyuskin, the northernmost part of that Cape and

    the northern New Siberian Islands. Nothing is known of the flora and

    vegetation of the small islands between Yalmal and Severnaya Zemlya, of

    which Lonely Island has long been known, or of the two islands between Franz

    Josef Land and Severnaya Zemlya, Ushakov Island (covered with ice) and Vize

    Island, but they naturally belong to the snow zone.

            In the snow zone vascular plants are found only in favored places well

    sheltered by snow in wintertime and warmed by insolation in summertime. In

    the snow fields red snow ( Chlamydomonas nivalis ) is common, and brown patches

    of diatoms also occur on the glaciers and snow fields.

            The vascular plants grow mostly in clumps of thawing earth found in the

    pits of frost cracks. Only the most hardy species, such as Alopecurus alpinus ,

    Poa alpigena , Silene acaulis , Draba alpina , Saxifraga groenlandica , S. oppo

    sitifolia , Papaver radicatum , Siversia glacialis , and Myosotis alpestris , are

    found. At Cape Chelyuskin, for instance, Kjellman found very few lichens and

    mosses but mostly Deschampsia arctica and Catabrosa algida in the cracks of

    the earth.

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            High-Arctic Tundra Zone . South of the snow zone follows the high-arctic

    tundra. It comprises the southern island of Novaya Zemlya, the northern

    shore of Matochkin Shar, Vaigach, the northern half of Yalmal Peninsula, the

    northern part of Taimyr Peninsula, and a narrow coastal strip from the mouth

    of Khatanga River to somewhat east of Chaun Bay, including the Lena Delta.

    Wrangel Island must also be regarded as belonging to this zone as it has a

    flora of 150 species, several of them not high-arctic, although Soviet

    botanists refer it to the snow zone.

            The snow cover is usually thin in the high-arctic zone and the ground

    ice thaws to depths of only 20 to 30 cm.

            Along the seashore of the high-arctic zone are found salt or brackish

    meadows. Their vegetation consists of Dupontia fischeri, Festuca rubra ,

    Calamagrostis deschampsioides , Carex rariflora and C. subspathacea. East

    of the Lena River the saline meadows are of a different character, Carex

    aquatilis stans
    and Arctophila fulva alternating with mosses. Along the

    seashore are found Puccinellia phryganode s, Elymus , Honckenya peploides ,

    Stellaria humifusa , Cochlearia , and Magricaria ambigua .

            South of the shore line, more or less wet polygon-tundra, Dryas tundra,

    and polygon swamps cover most of the ground. They are all characterized by

    their open vegetation. Lichen tundra, moss tundra, and meadows occupy small

    areas. In the inundation zone along the rivers occur Salix reptans and, to

    a lesser extent, S. lanata and S. pulchra .

            The polygon tundra is formed by the action of strong wind and heavy

    frost. In wintertime the wind blows away the snow from the summits of small

    elevations of the ground. Any existing vegetation is also blown away there,

    045      |      Vol_VI-0374                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

    and only bare earth remains throughout the winter. The earth freezes, and

    cracks forming polygons around the bare patches are formed by the expanding

    ice. In these craks the melting water accumulates in the spring and re–

    peatedly freezes and thaws. This widens the cracks. A solid lump of ice

    usually remains in the center of the polygons until the middle of the summer.

    The vegetation finds a foothold only in the cracks, which form a sluggish

    drainage system, so that water remains in them during most of the summer.

    Such polygons are formed chiefly on clayey or muddy ground, but not in sandy

    soil. The scraping of the hoofs of the reindeer may initiate the formation

    of polygon tundra in moss or lichen tundra.

            Lichens and mosses are poorly developed in the polygon tundra. Among

    the lichens occur Alectoria ochroleuca and Cornicularia divergens (predom–

    inating) and, further, Cetraria crispa , C. cucullata , Stereocaulon alpinum ,

    and Thamnolia vermicularis. Cladonia species are lacking.

            The mosses are Drepanoclades uncinatus , Polytrichum alpinum , Rhacmitrium

    canescens, R. hypnoides ( Lanuginosum ), and Webera cruda .

            The most characteristic vascular plants are:

    Hierochloe alpina Betula nana (or nana exilis )
    Poa arctica Polygonum viviparum
    Festuca brachyphylla Minuartia macrocarpa
    Alopecurus alpinus Draba fladnizensis
    Carex bigelowii Parrya nudicaulis
    Luzula confusa Papaver radicatum
    Salix rotundifolia Dryas octopetala punctata
    Salix polaris Pedicularis oederi

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            In addition to the species enumerated above, the wet polygon tundra is

    characterized by Eriophorum scheuchzeri , Calamagrostis neglecta borealis ,

    Arctagrostis latifolia , Pedicularis sudetica , and others.

            In the southernmost part of the high-arctic zone the polygon tundras

    show transgressions to patchy tundra where low grasses and even low shrubs

    occur among rich patches of lichens and mosses. This patchy tundra, how–

    ever, mostly belongs to the lichen-moss tundra zone.

            Ranunculus hyperboreus and R. pallasii are found in very wet places.

            The Dryas tundra dominates in many places in the southern part of the

    high-arctic zone, as in southern Taimyr and at the mouth of the Yenisei.

            Besides Dryas octopetala punctata , the following are characteristic:

    Hierochloe alpina Astragalus umbellatus
    Carex rupestris Oxytropis nigrescens (in

    the east)
    Lloydia serotina Saxifraga oppositifolia
    Salix arctica Diapensia lapponica
    Parrya nudicaulis

            In the thin and sparse lichen-moss layer are found Ditrichum flexicaule ,

    Distichium capillarum , and others.

            Dryas tundra occurs southward, mostly as patches in the polygon tundra

    southward to the forests.

            Where sandy soil predominates over large areas, lichen tundra occurs.

    Its vegetation may be said to be an extension northward of that of the northern–

    most sandy larch forests. In the high-arctic lichen tundra, patches of lichens

    alternate with patches of bare soil, which sometimes occupy large areas. The

    cryptogams are represented by Cetraria cucullata , C. islandica , Alectoria

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    nigrescens , A. chalybeiformis , Cornicularia aculeata , and Polytrichum hyper

    boreum . Middendorff found such lichen tundra in the Taimyr only in the

    Byrranga Mountains, the vegetation of which is still very little known. In

    the northern part of the high-arctic zone the lichen tundra depauperates

    and transgressions occur to polygon tundra.

            In the southern part of the zone, where domesticated reindeer are fre–

    quent, the lichen tundra is badly damaged by them. The composition of the

    lichen flora is influenced by their grazing. Cetaria nivalis , Sphaerophorus

    , and Stereocaulon paschale are less eaten and are thus favored by

    grazing. Lichens grow slowly in northern countries, and the lichen tundra

    recovers slowly from the grazing.

            The moss tundra is poorly developed in the high-arctic zone and is found

    chiefly along the shore of the Arctic Sea and in the southern central part

    eastward to the Lena River. Farther eastward, in spite of the lower precipita–

    tion, the vegetation is so boggy that moss tundra is not formed. Rather wet

    moss tundras occur with Eriophorum angustifolium , Carex aquatilis stans ,

    C. misandra , Salix pulchra , S. reticulata , Polygonum viviparum , and Dryas .

            Peat bogs or mires with a continuous peat cover are not present in the

    high-arctic zone, where Sphagnum is mostly lacking or occurs merely as a

    floristic rarity. They are replaced by polygonal sedge swa m ps, which often

    cover large areas. These swamps are divided into polygons by cracks with

    somewhat raised margins. The ice persists in the cracks up to late summer.

    These polygonal swamps correspond to the drier polygon tundra. The humus

    layer is usually shallow, 25 to 30 cm., and is formed by mosses such as

    Calliergon richardsonii , C. sarmentosum , C. stramineum , Drepanocladum exanula

    tus , D. revolvens , D. sendtneri , and D. vernicosus . If Sphagnum is present

    048      |      Vol_VI-0377                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    at all, S. obtusum and S. squarrosum are the most common species. Of higher

    plants Carex predominates, with C. aquatilis stans , C. rariflora , and

    C. rotundata ; and, further, Eriophorum angustifolium , E. russeolum , Dupontia

    , Hierochloe pauciflora , Salix reptans , Caltha palustris , Comarum

    , and Patesites frigidus are commoncomponents in these high-arctic

    polygon swamps.

            Tundra meadows are poorly developed especially in the central Siberian

    parts. They are formed in places where deep and long-remaining snow cover

    kills most of the lichens and mosses. The grasses and sedges in these

    meadows are the same as in the polygon swamps. They are low but grow thickly.

    Salix pulchra , Ranunculus acris borealis , Lagotis stelleri , and others are

    found in these meadows. The bottom layer is formed by a few scattered lichens

    and mosses that can stand the long snow cover, such as Dicranum congestum ,

    Drepanocladus uncinatus , Polytrichum hyperboreum , Cetraria delisei .

            In the extreme north the twigs of the shrubs shade the earth and the

    ground ice persists longer under them than in their surroundings. Farther

    south the conditions are just the opposite. The twigs are warmed by the sun

    and the ice melts more quickly beneath them than on the open tundra. Shrubs

    are therefore suppressed in the north but favored in the south.

            Lichen-Moss Tundra Zone . South of the high-arctic tundra zone stretches

    the lichen-moss tundra zone in the west, replaced east of the Lena River by

    the Eriophorum -tussock tundra. The lichen-moss tundra occupies northern Kanin

    Peninsula, Kolguev, a narrow strip of the mainland, the northernmost Urals,

    southern Yalmal Peninsula, and from northern Taz Bay to lower Khatanga and

    Olenek rivers. The Eriophorum -tussock tundra stretches from west of the Yana

    049      |      Vol_VI-0378                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    River to the mountains east of the lower Kolyma River. It also occurs in

    the shrub zone south of Channkaia Bay and in Chukotsk Peninsula.

            It is characteristic of the lichen-moss tundra zone that dwarf shrubs

    are common and that the plant communities are more numerous and sometimes

    more or less closed.

            The chief vegetational types are polygon tundra, Dryas tundra, lichen

    tundra, moss tundra, mucky bogs, and tundra meadows.

            The polygon tundra described above is distributed throughout the zone,

    but its floristic composition is poorer and the dwarf shrubs play a greater

    part here than in the high-arctic zone. Betula nana, Empetrum, Arctostaphylos

    alpina, Ledum palustre decumbens, Vaccinium vitis-idaea minus
    , and V. uligi

    nosum are frequent and lichens are more abundant. Grass patches occur with

    such plants are Carex melanocrapa, Festuca brachyphylla, Juncus biglumis,

    J. castaneus, Papaver radicatum, Petasites gmelini

            The Dryas tundra is of a different composition, especially in the western

    Siberian lowlands, with Alopecurus alpinus, Arctagrostis latifolia, Carex

    bigelowii, Luzula arctica
    , and Salix lanata in the upper layer and Cetraria

    , C. cucullata , Cladonia gracilis , Dicranum congestum , Dufouraea

    , Hylocomium proliferum ( splendens ), Polytrichum hyperboreum , Ptilidium

    , and Salix polaris in the lower.

            The lichen tundra occurs preferably on sandy soil. It is usually poor

    in species as well as in individuals. Their poorness is in part the result

    of reindeer grazing.

            The Alectoria tundra is most hardy against frost and wind and therefore

    predominates in the northern and in the more continental parts, while the

    050      |      Vol_VI-0379                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    Cladonia tundra, which suffers badly from heavy frosts, is found chiefly in

    the southern part of the zone, especially from Kola to the Yenisei and in the

    Far East, where they are particularly common. In the loose lichen-moss mat

    of the Alectoria tundra, Alectoria nigricans predominates, and Cetraria

    species, especially C. cucullata and C. nivalis , are much less prominent.

    Vascular plants are Carex bigelowii , Luzula confusa , Ledum palustre decum

    bens , Vaccinium vitis - idaea minus , Cassiope tegragona , Diapensia lapponica ,

    and, in the east, Polygonum pawlovskianum . On the Cladonia tundra, Cladonia

    , C. rangiferina , C. uncialis and C. amaurocraea predominate, and

    in Kola Peninsula also C. alpestris . Other characteristic species of the

    ground layer are Cornicularia divergens , Aulacomnium turgidum , Cetraria

    , C. crispa , Cladonia pleurota , C. gracilis , Dicranum elongatum ,

    Dufourea arctica , Polytrichum hyperboreum , P. piliferum , and Rhacomitrium

    ( lanuginosum ), the last-mentioned being especially common in the

    west. The vascular plants consist of scattered specimens of Hierochloe

    , Arctagrostis latifolia , Luzula confusa , Pedicularis hirsuta , Parrya

    , and others. Dwarf shrubs are rare and low-growing, as the snow

    cover is shallow. Among them predominate Betula nana , Salix rotundifolia ,

    Empetrum , Vaccinium vitis - idaea minus , V. uliginosum , and Arctostaphylus

    . In the west, Cladonia -moss tundra occurs intermingled with the

    Cladonia tundra, and also a hillock tundra with Alectoria societies on the

    almost barren tops of the hillocks and Cladonia societies at the base.

            The moss tundra has a wide distribution on clayish or peaty soil and

    is especially common in the southern part of the lichen-moss tundra zone.

    The bottom layer consists of mosses mixed with some lichens. Representative

    species are Aulacomnium palustre , A. turgidum , Dicranum angustum , Hylocomium

    051      |      Vol_VI-0380                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    proliferum ( splendens ), Polytrichum stricutum , Ptilidium ciliare , and the

    lichens Cetraria cucullata , Cladonia gracilis , C. sylvatica , and Peltigera

    . Vascular plants are Calamagrostis neglecta borealis , Festuca supina ,

    Empetrum , Vaccinium vitis - idaea minus , Petasites frigidus , and others, the

    height of which depends on the depth of the snow cover.

            Peat bogs or mires with a continuous peat cover are lacking or very poorly

    developed within the area of the permanently frozen soil. They occur, however,

    in the southern part of the lichen-moss tundra zone, where they are shallow,

    at most 4 6 meters deep. The peat is perpetually frozen and does not

    increase. In Europe, peat bogs reach the coast of the Arctic Sea and they

    occur in the southern part of the zone to the Yenisei. Farther east they do

    not enter into the tundra zone. Thus in the Arctic they are confined to the

    border of the permanently frozen ground. Those found farther to the north

    are probably relicts from a warmer period. While the peat bogs in the wood

    tundra and in the shrub zone are characterized by large hillocks 4 to 8 meters

    high, the more northern ones in districts where the soil thaws only superfi–

    cially have very low hillocks only 0.5 to 0.7 meter high. The northern mires

    also have a poorer vegetation and are very shallow, only 0.5 to 1 meter deep.

    Sphagnum plays a less prominent part, and the peat consists of other mosses.

    Lichens of a more arctic type such as Cetraria cucullata and C. nivalis pre–

    dominate. The dwarf shrubs are few and chiefly comprise Betula nana , Vac

    cinium , and Ledum .

            Between the hillocks occur Sphagnum balticum , S. fimbriatum , S. lindbergii ,

    Calliergon cordifolium , C. sarmentosum , Drepanocladus exannulatus , D. vernico

    sus , Polytrichum jensenii , and others. Scattered specimens of Eriophorum

    052      |      Vol_VI-0381                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    vaginatum , E. russeolum , Carex aquatilis stans , C. rariflora , and C. rotundata

    are found in this moss mat.

            East of the Lena River, in spite of the lower precipitation, a wet muddy

    Eriophorum vaginatum tussock tundra with Sphagnum is formed and replaces the

    moss tundra and the mires of the west. The reason for this is unclear. This

    tussock tundra thaws to a depth of 0.5 to 1 meter in summertime. Circular

    spots lacking vegetation and covered with ground-water form around the tus–

    socks — a very characteristic feature of the tussock tundra. In Europe and

    western Siberia such tussock tundras are lacking in the north; some occur in

    the southern part of the lichen-moss tundra zone, but they are more common

    still farther south. East of the Lena and especially in the Chukotsk Penin–

    sula, the Anadyr and Penzhina regions they form a special tussock tundra zone.

    The tussocks cover 30 to 50% of the surface. They consist of Eriophorum

    30 to 50 cm. high.

            East of the Indigirka, Carex lugens takes part in the tussock formation,

    especially in the forest tundra. The peat, which is shallow — usually 15 to

    20 cm. deep only — is mostly composed of Eriophorum vaginatum and Carex lugens .

    The mosses and lichens sometimes vary widely, as Sphagnum lenense , S. balticum ,

    and S. Warnstorffianum ( Warnstorfii ), occur in Europe and western Siberia, but

    in other places Aulocomnium turgidum , Dicranum elongatum , Polytrichum jensenii ,

    P. strictum predominate, and still elsewhere lichens such as Cetmaria cucullata ,

    C. crispa , Cladonia rangiferina , and others are found in the ground layer. Vas–

    cular plants are Betula nana (or nana exilis ) Ledum , Vaccinium vitis - idaea minus ,

    and a few others. The snow cover reaches to about the top of the tussocks.

            Where the snowdrifts are deep and remain long, grass meadows are formed with

    053      |      Vol_VI-0382                                                                                                                  
    EA- [ ?] PS . Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    a close mat of vegetation. Their species content is very variable in the

    different parts of the tundra zone, as many plants with a restricted dis–

    tribution occur among them. The following species are often found in the

    tundra meadows.

    Poa alpigena Draba glacialis
    Carex lachenalii Draba hirta
    Oxyria digyna Saxifraga cernua
    Polygonum bistorta Pachypleurum alpinum
    Ranunculus nivalis Valeriana capitata

            The dwarf shrubs are suppressed by the long-lasting snow cover, but

    some of them occur also in the meadows, mostly along their margins, such as

    Salix herbacea , S. lanata , S. pulchra , S. reptans in the west, and Salix

    chamissonis , Alnus crispa , Spiraea beauverdiana , and Rhododendron chrysanthum

    in the east.

            While meadows prevail at the base of slopes, another plant community with

    continuous plant mats occurs at the upper, less snow-covered, parts of dry

    slopes. It is related to the meadows and transgresses into them. These xero–

    phytic flower mats consist of Dryas, Empetrum, Equisetum arvense, Lloydia

    serotina, Salix reticulata, Polygonum viviparum Trollius europeus
    ( asiaticus

    to the east), Thalictrum alpinum, Sedum rhodiola, Hedysarum obscurum, Oxytropis

    campestris sordida, Geranium silvaticum, Pedicularis oederi, Taraxacum

    and other. In the bottom layer Hylocomium proliferum ( splendens ), Rytidium

    , Thuidium abietinum , and also Cetraria islandica , and Cladonia occur.

            Ponds and lakes thaw late in the lichen-moss tundra zone and the water

    remains cold throughout the summer. They are therefore practically destitute

    of vascular plants.

    054      |      Vol_VI-0383                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

            On the sea beach in the Chukotsk Peninsula, Elymus europeus mollis is

    present in sandy places. Senecio pseudo-arnica, Honckenya peploides, and Merckia physodes are also found there.

            Shrub Tundra Zone . South of the lichen-moss tundra zone comes the shrub

    tundra, the southern limit of which is the arctic timber line. It is char–

    acterized by deeply thawing soil and by an abundance of shrubs in the moss

    tundra, mires, bogs, and meadows. It stretches from northern Kola Peninsula

    over southern Kanin Peninsula, lower Pechora River, the northern Urals, middle

    Taz Bay, lower Yenisei, and Khatanga rivers to lower Olenek River. There

    it is interrupted by mountains and continues from lower Yana River to lower

    Kolyma River. Most of the area between Anadyr River and the Gizhiga and northern

    Kamchatka also belongs to this zone, but it is not reckoned here as belong–

    ing to the Arctic.

            Polygon tundra, moss tundra, mires, Eriophorum vaginatum tussock tundra,

    shrub thickets, and meadows constitute the main vegetational types.

            The polygon tundra stretches throughout the zone but has poorer species

    content and more dwarf shrubs than in the more northerly tundra zones. Patches

    of Dryas tundra occur intermingled with it.

            The shrub-covered moss tundras with Betula nana and Ledum and wet

    plant communities occupy by far the largest area of the shrub tundra zone.

    Besides Betula nana ( exilis ) and Ledum , Salix glauca , S. pulchra , Vaccinium

    - idaea minus , and V. uliginosum occur in the moss tundra. The shrubs

    cover 50 to 60% of the area but are more scattered eastward. In the grass

    layer are frequently found Carex bigelowii , Festuca supina , Empetrum , and

    Petasites frigidus . In the southernmost part of the zone, forest plants that are

    055      |      Vol_VI-0384                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    otherwise alien to the Arctic, such as Lycopodium annotinum , Carex globu

    lari s, Delphinium elatum , Geum rivale , Angelica decurrens , Myosotis al

    pestris , and Circium heterophyllum , occur in the moss tundra. In the bottom

    layer mosses and lichens occur, mostly in patches with wet open soil between

    them, viz., Dicranum elongatum , Drepanocladus uncinatus , Polytrichum strictum ,

    Sphagnum compactum , Gymnocolea inflata , Cetraria islandica , C. nivalis ,

    Cladonia gracilis , C. silvatica , and, especially in the southernmost part,

    Nephroma arctica . The shrubs increase in size southward. At the arctic

    tree line Betula nana reaches a height of about 50 cm. No such shrubby moss

    tundra occurs east of the Lena River.

            The mires are shallow and are composed of Sphagnum lenense , S. squar

    rosum , and S. subsecundum and other mosses, but not Sphagnum fuscum or

    S. medium ( magellanicum ) as in more southerly bogs. Bogs with hillocks more

    than 2 meter high are common in the southern tundra forests (except in

    central Siberia), where permanently frozen soil occurs only in patches, but

    they are rarely found in the shrub tundra zone except west of Pechora. The

    peat bogs with hillocks in arctic European Russia occur preferably in closed

    basins. They are covered with deep snow in wintertime and with melting water

    in springtime. The hillocks are dome-shaped or else elongated, with a center

    of mineral earth. The peat on the top of the hillocks consists of Sphagna ,

    Dicranum congestum , D. elongatum , and Polytrichum strictum , which in a later

    stage of development are replaced by Ochrolechia tartares , Pertusariae and

    stunted individuals of Cladonia deformis and Cetraria nivalis .

            On the slopes of the hillocks are found Sphagnum angustifolium , S. medium

    ( megellanicum ), S. fuscum , green mosses, Cetraria cucullata , C. islandica ,

    C. nivalis , Cladonia rangiferna , Eriphorum vaginatum , Betula nana , Rubus

    055a      |      Vol_VI-0385                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    chamaemorus , Empetrum , Vaccinium vitis - idaea minus , V. uliginosum , and

    others. In the wet places between the hillocks a society containing

    Sphagnum balticum , S. lindbergii , Calliergon cordofolium , C. stramineum ,

    Drepanoclodus exannulatus , and Paludella squarrosa occurs, with Carex

    , C. chordorrhiza , C. limosa , C. rariflora , C. rotundata , Erio

    phorum angustifolium , and Andromeda polifolia in the grass layer.

            In ponds and lakes, especially in the southern part of the zone, are

    found Sparganium hyperboreum , Potamogeton alpinus , P. natans , and Hippuris

    . The ponds develop into mires chiefly owing to the growth of

    Arctophila fulva , Carex aquatilis stans , Eriophorum augustifolium , Hippuris ,

    Ranunculus pallasii , Drepanocladus , and Calliergon , and not, as in the case

    of the forest tundra, through the growth of Sphagnum .

            The Eriphorum tussock tundra described above frequently occurs in the

    shrub tundra zone. To the south, especially in the forest tundra, it has

    a Sphagnum facies.

            The shrubs in the tundra shrub zone differ from those in the lichen–

    moss tundra zone in the stronger development of their branches. They occur

    especially on muddy soil and on the lower parts of the slopes where the snow

    cover is deep. Their roots do not go deep into the cold soil and they are

    therefore easy to root up. In the neighborhood of human habitation, there–

    fore, they are often lacking as they have been used for fuel. The domesticated

    reindeer, which are especially common on the Molozemelskaya and Bolshezemelskaya

    tundras, feed on the shrubs and destroy them to a large extent. The shrub

    vegetation is then replaced by grassland. Salix myrsinites is considered to

    be especially favored by reindeer grazing, as it is avoided by the animals.

    056      |      Vol_VI-0386                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

            The shrubs are 1/2 to 1 meter high in the northern part of the shrub

    tundra zone, but 1 1/2 to 3 meters in its southern part. The ground ice

    thaws to a depth of 1 to 2 meters beneath the shrubs but usually to a depth

    of only 40 to 50 cm. in the surrounding moss tundra. In the western part

    of the shrub tundra zone the following species predominate in muddy soil:

    Betula nana , Salix glauca , S. lanata , S. lapponum , S. phylicifolia , S. pulchra ,

    and S. reptans . In sandy soil Betula nana , with Cladonia sylvatica and

    C. amauracea in the bottom layer, predominates on Kola Peninsula and in

    European Russia, but is less common on the continental tundras of Siberia.

    In Anadyr and Penzhina, Betula nana exilis replaces B. nana and Betula midden

    dorffii and Alnus crispa the Salix shrubs. Alnus crispa enters into the shrub

    tundra zone on Bolshezemelskay tundra and eastward, and occurs in narrow

    strips along the river valleys. The ground layer in the tundra shrub forma–

    tion is formed by mosses such as Aulacomnium palustre , A. turgidum , Drepano

    cladus uncinatus , Hylocomium proliferum ( splendens ), Mnium rugicum , Polytrichum

    , and, in the southernmost part, Polytrichum commune , Sphagnum girgen

    sohnii , and S. squarrosum . Lichens are poorly developed on account of the

    deep shade. Cetraria islandica , Cladonia gracilis , Nephroma arctica , Peltigera

    , and others are, however, found.

            Forest Tundra . The tundras south of the northernmost trees, the forest

    tundra, are here not included in the Arctic. They form a rather broad belt

    where the tundra formations predominate with single, often palelike, nearly

    branchless trees or groups of trees scattered here and there. The forest

    tundra transgresses almost continuously into open woods, and the arctic forest

    line is often somewhat arbitrary. In European Russia the forest tundra zone

    057      |      Vol_VI-0387                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    is about 100 km. wide, but narrows to the eastward. The plant communities

    of the woods are found there mixed with those of the shrub tundra. Betula

    , Empetrum , Ledum , Vaccinium , and Salix dominate in the shrub layer.

            The last trees are in Kola Peninsula: Scotch pine ( Pinus silvestris )

    and in some places spruce ( Picea abies obovata ). North of the limit of the

    conifers is a brushwood belt of Betula tortuosa . East of Kola Peninsula the

    pine retreats southward when it meets the perpetually frozen soil. Its long

    central polelike root makes it very poorly adaptable to the shallow earth

    of the permafrost districts. On Bolshezemelskaya tundra, spruce, which has

    a flat root system, forms the timber line. From the Urals to the upper

    Anadyr River (Markovo) the last trees are larch, west of the Yenisei Larix

    , and east of that river L. dahurica .

            The larch is especially well adapted to frozen soil as it has a falt

    root system. In the Urals the buds of the larch burst while there is still

    a thick layer of snow remaining under the trees. In this [way ?] the period of

    vegetation is prolonged.

            The timber line has not always been the same. Finds of dead trunks far

    north of the present timber line show that in a warmer period trees grew

    farther northward than they do now. In Kanin Peninsula Ramsey found remains

    of spruce in more northerly situations than where it is found now. In the

    Molozemelskaya and Bolshezemelskaya tundras, logs of larch, birch, and spruce

    are found up to the very shore of the Arctic Sea. In the Gydan tundra,

    Gorodkov found birch remains 300 km. north of the present forest line; at

    the lower Yenisei fossil larch occurs up to 70° N. In northern Taimyr,

    Urvantsev found fossil larch, but it is possible that it had been carried

    058      |      Vol_VI-0388                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    to that place by a river. In the Anabar tundra are fossil larches,

    doubtless in situ . As the patches of wood in the forest tundra often de–

    pauperate, it seems probable that the timber line is still receding. In

    a warmer period southern plants may have been distributed farther north than

    they occur now. Such “steppe plants,” probably relicts from a warmer period

    are still found in the tundra woods, plants such as Carex supina , Arenaria

    , Cerastium maximum , Lychnis sibirica , Sisymbriu m , junceum ,

    Erysimum hieracifolium , Galium verum . Many of the mosses and lichens of

    the tundra zone are characteristic forest species and may be expected to

    be relicts from a period when the forests reached farther northward than

    they do now.


    Vegetation of Different Geographical Provinces

            Kola Peninsula was investigated by several Russian, Finnish, and Scan–

    dinavian botanists, among them Kihlman in 1887, Regel 1913, and Zinserling

    in 1931. The author visited the north coast of Kola Peninsula in 1926.

            The vegetation in Kola Peninsula is to a certain extent analogous to

    that of the Anadyr-Penzhina district. In both places the alpine zone of

    the nearby mountains goes down to sea level and a subalpine shrub-brushwood

    zone is inserted between this alpine zone and the conifer woods. In other

    parts of the Arctic such a brushwood zone is lacking (with the possible

    exception of the Urals) and the shrub tundra borders the woods. Kola is

    a hilly tableland and the zones are therefore more schematical there, while

    in the Far East a more complicated topography, with mountain chains and

    valleys, makes the zonation of the vegetation more complicated.

    059      |      Vol_VI-0389                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

            The phytogeographical interpretation of this alpine zone at sea level

    is not quite clear, but it seems difficult to find any essential difference

    between it and the flat tundras of the Arctic. In any case a narrow strip

    of the Kola Peninsula north of the brushwood of Betula tortuosa must be re–

    garded as belonging to the Arctic. Permanently frozen soil is very rare

    in the forest zone of the peninsula but occurs, although not continuously,

    over large areas, especially in peatbogs, which do not thaw deeper than 40

    to 50 cm. Bogs with large hillocks characteristic of the border line of

    frozen soil occur frequently in the birch belt. At Cape Orlov the soil was

    frozen to a depth of 10 cm. in the middle of June and to about 20 cm. in the

    middle of July, when the author visited that place.

            North of the timber line, which consists of pine, in some places replaced

    by spruce, a 60 to 100-k. -broad belt of Betula tortuosa is found, which in

    the river valleys reaches nearly to the Arctic Sea. The birches are 2 to 3

    meters high at the shore, but 3 to 5 meters farther inland. Single spruces

    occur here and there in the birch belt. The lower layer within the birch

    belt is very variable; Polytrichum , Sphagnum , or Cornus suecica undergrowth

    occur. On the divides and between the river valleys a Cladonia tundra pre–

    dominates. In the tundra meadows grow Nardus stricta , Salix herbacea , Sib

    baldia procumbens , Viola biflora , and Gnaphalium supinum. Juniperus communis

    is common north to the Arctic coast. Kihlman counted the annual rings of a

    trunk of Juniperus 8.3 cm. thick and found the tree to be 544 years old.

    coast of Kola Peninsula.

    060      |      Vol_VI-0390                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

            Kanin Peninsula was investigated botanically by Ruprecht in 1841,

    by Pohle (1898-99) and by Andreev (1930, 1931). The mountains of the penin [ ?]

    sula do not exceed 200 meters. The northern part belongs to the lichen-moss

    tundra zone. Real moss tundra is lacking and is replaced by a mixed lichen–

    moss tundra with low hillocks, covered with Deschampsia alpina , Carex bigelowii ,

    Betula nana , Empetrum , Salix glauca , S. herbacea , Polygonum bistorta , Arcto

    staphylus alpina , and Vaccinium vitis - idaea . The hillocks are about 75 cm.

    high and about 2 meters in diameter and cover about 15% of the surface. Mires

    containing Eriophorum angustifolium and E. scheuchzeri are common, and a Carex

    community covers large areas. On the slopes shrubs such as Salix

    , S. lanata , and S. myrsinites with Calamagrostis are common. In Andreev

    (1931) a fairly detailed account is given of the different small sociological


            The southern part of the Kanin Peninsula belongs to the shrub tundra zone.

    Picea abies obovata , Alnus crispa , and Betula tortuosa occur on the eastern

    coast in patches to the Pesh River, close to its mouth, and to within about

    25 km. of the mouth of the Vizhas River. In southwestern Kanin patches of

    Picea abies obovata about 15 cm. high and 50 cm. in diameter occur at Cape

    Konushin. There are a few forest species in Kanin, as Veratrum album lobeli

    anum , Rumex aquaticus , Dianthus superbus , Polemonium acutiflorum , Veronica

    , Pedicularis sceptrum - carelinum , Achillaea millefolium , and

    Solidago virgaurea , some species even reaching northern Kanin, e.g., Parnassia

    , Filipendula ulmaria , and Geum rivale . They are probably relicts

    from a warmer period, when the woods extended farther northward.

            Dunes with Elymus and Matricaria ambigua are found at the mouths of the

    061      |      Vol_VI-0391                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    rivers. The mead ows along the seashore are often 7 to 8 km. broad. They

    are marshy, with abundant Carex subspathacea . Their peat is used as fuel.

            Molozemelskaya and Bolshezemelskaya Tundras . These districts were

    investigated in 1837 [ ?]by Schrenk, by Sambuk and Dedov in 1926-29, and by

    Andreev in 1930.

            Whereas on the Kola Peninsula the pine and the spruce grow equally

    far to the north, east of the White Sea, the fir goes at least 1/2° farther

    northward than the pine. The Timan Mountains do not enter into the Arctic.

            Only the eastern part of the Timan coast from about 50° E. and a nar–

    row strip along the coast of Bolshevemelskaya tundra belongs to the lichen–

    moss tundra belt. South of this strip is found the shrub tundra, with Betula

    prevailing. This shrub belt is about 50 km. broad in the west but widens

    eastward to about 150 km. close to the Urals. At the mouth of the Pechora

    dense thickets of Salix pyrolifolia 1.5 meters high are found in the river

    valley. Alluvial meadows and sand dunes are also found there. On the Timan

    tundra and in the southern part of Molozemelskaya tundra many peat bogs with

    large hillocks are found. On Bolshezemelskaya tundra deep peat bogs, un–

    doubtedly relicts from a warmer period, occur. The Cladonia on the lichen

    tundra is badly damaged by the numerous reindeer, whose grazing favor Stereo

    caulon , Sphaerophorus globosus , and Festuca supina . On Molozemelskaya tundra

    southern species such as Anthoxantum odoratum , Solidago virgaurea , and Hiera

    cium of the alpinum group occur and on Bolshezemelskaya tundra grow Poa alpina ,

    Polygonum viviparum , Veratrum album lobeliamu m, Ranunculus acris borealis ,

    Geranium silvaticum , and Achillaea millefolium. Alnus crispa , which forms a

    subalpine belt in the Urals, occurs here in the lowlands along the river

    valley westward to the base of Kanin Peninsula. The northernmost trees are

    spruce trees.

    062      |      Vol_VI-0392                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

            Andreev (1932) describes a large number of plant communities from the

    Bolshezemelskaya tundra.

            Kolguev Island was investigated botanically by Ruprecht in 1841, by

    Pohle, and by Tolmachev (1929).

            The entire island belongs to the lichen-moss tundra zone. It is a

    round moraine island about 60 km. in diameter, with steep shores 40 to 50

    meters high. The flora comprises about 200 species. Several of them do

    not occur on the opposite mainland or farther to the east. This is doubt–

    less due to the influence of the warm water of the Gulf Stream. Mean tempera–

    ture in July is 8°C.; fogs are frequent in summer. Some of the southern

    species, such as Geranium silvaticum , Delphinium elatum , and Salix arbuscula ,

    occur only in the southern part of the island. Otherwise the northern part

    of the island has a richer vegetation, with meadows containing T r ollius europeus,

    Veronica longifolia , and Geum rivale. Salix nummularia and Silene acaulis are,

    besides Dryas , the most common plants in dry places in the northern part. A

    wet tundra containing Eriophorum angustifolium , E. scheuchzeri , Carex big

    elowii , C. rariflora , Dupontia fischeri , Arctagrostis latifolia , and Petasites

    is common on all plains. In wet places Carex aquatilis , Colpodium

    , and Caltha palustris occur. Eriophorum vaginatum , Rubus chamaemorus ,

    Betula nana tundra is common. Salix glauca shrubs occur in abundance. In

    sheltered places are found Salix hastata , S. herbacea , S. lanata (predominating),

    Ranunculus borealis , Alchemilla glomerulans , A. murbeckiana , Oxytropis cam

    pestris sordida , G entiana verna , Myosotis alpestris , Valeriana capitata , and

    Senecio campestris . Along the coasts are found Cochlearia and Matricaria

    , and in the salt marshes Puccinellia vilfoidea , Carex subspathacea ,

    and Stellaria humifusa .

    063      |      Vol_VI-0393                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

            Polar Urals . In the Polar Urals, the Pai-khoy Mountains are not very

    well known botanically. The first information was brought back by Ruprecht

    in 1848. Gorodkov investigated the Sob River (1924), Sochava the Liapina

    River (1926-27) and Andreev, Igoshina, and Keskov the eastern slopes from

    Obdorsk to Baidaratskaya (Kara) Bay (1935). These investigations mostly

    concern the southern part of the area.

            The p P ai-khoy Mountains fall within the lichen-moss tundra zone, only the

    shores of Yugor Shar can be regarded as belonging to the high-arctic tundra.

    The shrub tundra zone is replaced by the subalpine belt. Above the forests,

    which consist of larch and to a certain extent of Picea abies obovata with

    the mosses Betula nana or Salix shrubs as undergrowth, stretches the subal–

    pine belt. It consists on the Asiatic side of Alnus crispa and Betula tor

    tuosa with grass undergrowth. In the alpine zone barren screes predominate

    and snow fields occur on the summits. At Sob River the forests reach up to

    200 meters, and single larch trees extend to 300 meters within the subalpine

    Alnus belt. The Alnus thickets gradually diminish in height the farther up

    they grow. Their upper limit is about 350 meters; farther northward and west–

    ward the limit is lower. Single crippled low shrubs of Alnus grow at an

    elevation considerably above 350 meters. The Alnus belt reaches close to the

    southwestern shore of Baidaratskaya (Kara) Bay, where it borders the arctic

    lichen-moss tundra. Alnus does not thrive on olivine, a common mineral in the

    northeastern Urals. There it is replaced by Betula tortuosa and Juniperus

    sibirica with single larch trees.

            In places where snow persists long in the Alnus belt, and in gullies

    along the brooks, there are meadows with high thickets of Salix lanata , S. has

    tata , and S. phylicifolia .

    064      |      Vol_VI-0394                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

            In the lower alpine zone moss societies predominate. A Rhacomitrium

    tundra with R. hypnoides ( lanuginosum ) is especially common on olivine.

    Lichens are rare in the Rhacomitrium tundra, but Salix shrubs occur. The

    rocks are often red with Trentepholia jolithus . Above 500 meters in the

    mountains the moss associations are much torn up by water erosion, and

    Alectoria tundra with Cornicularia divergens predominates on the poorly

    snow-covered summits, as in the high-arctic flat tundra. At elevations

    over 500 meters only small groups of lichen and moss tundra occur between

    the rocks, even on flat tops.

            The vegetation of the Polar Urals is much destroyed by the grazing of

    the reindeer. Only steep places are unaffected by the grazing.

            Novaya Zemlya and Vaigach . Novaya Zemlya has been more thoroughly in–

    vestigated botanically than any other part of the Russian arctic, possibly

    with the exception of Kola Peninsula. The first botanical investigations

    were made by von Baer and Ziwolka in 1837 and by Middendorff in 1870, both

    published by Trautvetter. Later many botanists collected on Novaya Zemlya,

    among whom may be mentioned the Nordenskiőld expedition in 1875 and in 1878

    with Kjellman, Lundstrőm, and Almquist as botanists, the British Markham

    expedition in 1879, Ekstam in 1891 and 1895, the British Pearson-Feilden

    expedition in 1895, the Russian Ermak expedition in 1901 published by Palibin

    (1903-06), the French officer B e é nard in 1908 and 1914, the Norwegian expedi–

    tion with Lynge as botanist (1929), A. Tolmachev in 1923-27, and lastly

    Zubkov (1930). Although members of the B e é nard expedition crossed the

    island, which is about 100 km. broad, its interior is still unknown botan–

    ically. The mountains certainly to a great extent lack vegetation even in

    the south.

    065      |      Vol_VI-0395                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

            The mountainous and glacier-filled north island of Novaya Zemlya belongs

    to the snow zone with the exception [ ?] of the southernmost part along Matochkin

    Shar. All the rest of it belongs to the high-arctic tundra zone.

            At Matochkin Shar the first signs of spring can be noted in the middle

    of April, according to Tolmachev, but it is not until the end of May that

    the shoots of the earlier species begin to grow. These are Poa abbreviata ,

    Minuartia rubella , Papaver radicatum , Saxifraga groenlandica , S. nivalis ,

    and Potentilla hyparctica . On the 8th of June was seen the first flower of

    the earliest plant, Saxifraga oppositifolia , and by the end of June many

    plants were flowering.

            On the south island polygon tundra, with cracks due to the coastal climate,

    is rare. Large areas with muddy and stony soil have a relief recalling that

    of the northern peat bogs, with flat hillocks about 40 cm. high and about 10

    meters in diameter, but Sphagnum is lacking. They are covered with Cladonia

    moss- Salix polaris communities containing such herbs as Arctagrostis latifolia ,

    Carex bigelowii , Luzula confusa , Cerastium regelii , Silene acaulis , Saxifraga

    , and Dryas .

            The vegetation of the western coast is richer than that of the eastern.

    On the eastern (Kara) coast only about 130 vascular plants are known, which

    is about 60 less than on the western. The plants also extend farther north–

    ward on the western coast than on the eastern. A few Siberian species, how–

    ever, occur only on the eastern coast, such as Minuartia macrocarpa , Hesperis

    , and Lagotis .

            Tolmachev differentiates one coastal and one inner zone of vegetation at

    the eastern entrance to Matchkin Shar. The coastal zone is 5 to 6 km. broad

    066      |      Vol_VI-0396                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    and is characterized by abundant Luzula confusa and Hesperis pallasii .

    Luzula diminishes in the interior and is replaced by Dryas , and Hesparis

    is absent there.

            Together with Salix arctica and S. polaris , Dryas plays a large part

    in the vegetation in places with an even snow cover. There are also found

    Poa arctica , Carex rupestris , Silene acaulis , Draba nivalis , Saxifraga

    , Oxytropis campestris sordida , Myosotis alpestris , Eritrichium

    , Campanula uniflora , Taraxacum platylepium , and others.

            The mires occupy smaller areas than the society described above. They

    contain Eriophorum angustifolium , E. scheuchzeri , Arctagrostis latifolia ,

    Dupontia fischeri , Pleuropogon sabinei , Deschampsia alpina , Carex b [ ?] elowii ,

    C. misandra , Juncus biglumis , Polygonum viviparum , and Caltha palustris .

    Mosses play a minor part in them.

            In the large river valley the vegetation is more closed than on the

    divides. On well-drained slopes arctic flower mats are found containing

    such species as Poa alpigena , Trisetum spicatum , Polygonum viviparum , Silene

    , Astragalus alpinus arcticus , Thalictrum alpinum , Ranunculus acris

    borealis, Saxifraga hieracifolia , Potentilla emarginata , Polemonius boreale ,

    and Artemisia tilesii .

            Both shallow snow cover and very deep snow cover result in areas of open

    earth, and close to the snow fields all vegetation is lacking. In places

    that become free from snow early Poa abbreviate , Papaver radicatum Draba

    , Saxifraga oppositifolia , and Potentilla hyparctica predominate.

    Po a , Papaver , and Potentilla form a community alternating with Saxifraga


    067      |      Vol_VI-0397                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

            The water plants comprise Puccinellia phryganodes , Pleuropogon sabinei ,

    Phippsia algida , Caltha palustris , and Ranunculus hyperboreus .

            A fairly detailed description of the vegetation of the south island

    of Novaya Zemlya is given by Zubkov (1932).

            Vaigach has a similar flora and vegetation to those of Novaya Zemlya.

    A few plants reach Vaigach but not Novaya Zemlya, such as Elymus arenarium ,

    Veratrum album lobelianum , Allium schoenorasum , Polygonum bistorta , Trollius

    , Parnassia palustris , Rubus arcticus , Viola biflora , Epilobium

    , Cortusa matthioli , Achillaea millefolium , and Chrysanthemum bi

    pinnatum .

            Franz Josef Land . The flora of Franz Josef Land belonging to the snow

    zone is known from brief notes in the reports of several expeditions visiting

    this archipelago. The first records come from Payer, a member of the Austro–

    Hungarian expedition of 1873-74. More information was supplied by Fischer,

    a member of the Jackson-Harnsworth expedition 1894-97, by the Yermak expedi–

    tion in 1901, and by the Norwegian expedition in 1930. Thirty-six vascular

    plants are known from the archipelago. The vegetation is extremely scanty

    with only a few specimens represented by a few individuals in each small

    place free of ice. No seashore plants occur. The only place in which the

    vegetation is somewhat richer is at the base of the bird cliffs. The follow–

    ing plants reach their northern limit in Franz Josef Land, which extends to

    82° N.: Phippsia concinna , Cerastium hyperboreum , C. regelii , Saxifraga

    , S. foliolosa , S. nivalis tanuis .

            Ya l mal and the West Siberian Lowlands . This district is very little

    known botanically. Northern Ya l mal was visited by Kjellman during the Vega

    068      |      Vol_VI-0398                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    expedition (1878). Andreev (1933) gives a review of the vegetation of Yalmal,

    dealing chiefly with its value as reindeer food.

            The northern half of Ya l mal belongs to the high-arctic tundra zone; the

    lichen-moss tundra stretches nearly to the base of the peninsula, and the

    shrub tundra nearly to the mouth of the Ob River, where the northernmost

    trees occur. The peninsula between Ob Bay and Taz Bay belongs to the shrub

    tundra, with the exception of the northernmost rim. The trees meet at about

    the Arctic Circle. In the high-arctic zone polygon tundra and Carex bogs are

    common. In the lichen-moss tundra zone moss tundras containing Aulacomnium

    , Dicranum elongatum , Polytrichum strictum , Ptilium ciliare , though

    few Sphagna , S. acutifolium , ( S. nemoreum ), S. compactum , and lichens, occupy

    large areas. Low Salix shrubs occur in the river valleys. In swampy places a

    wet tundra with low hillocks and Betula nana is common.

            Gydan Tundra and Mouth of the Yenisei . The Gydan tundra was investigated

    by A. Tolmachev (1926) and by Gorodkov (1927), the lower Yenisei was visited by

    several botanists, among whom may be mentioned Schmidt in 1866, Lundstrőm (1875),

    Arnell (1876), Brenner (1876), Kjellman (1878), Reverdatto (1912), and A. Tol–

    machev (1926-30). Approximately north of 72° the high-arctic tundra prevails,

    and the lichen-moss tundra extends from there to about 70°.

            Along the Yenisei the forest stops at Dudinka, 69° 23′ N. latitude. It

    consists of Picea abies obovata. Pinus cembra sibirica , Abies sibirica , Betula

    , B. tortuosa , and, in places, Larix sibirica. Abies sibirica forms

    thickets rarely exceeding 2 meters in height, with Ledum and Vaccinium in the

    bottom layer. Alnus thickets are also common along rivers.

            North of Dudinka, Alnus thickets and Salix shrubs prevail, with single

    069      |      Vol_VI-0399                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    larch trees among them. North of Ust Yeniseisk, at 69° 39′, only a few

    larch trees are found in sheltered ravines. The Salix lanata shrubs about

    2 meters high, sometimes mixed with Alnus , form almost impenetrable thickets,

    often bordered by lower Salix glauca shrubs. Betula nana abounds on the

    wet tundra. Eriophorum angustifolium , Rubus chamaemorus , Ledum , and Vac

    cinium uliginosum are also common components. Here is also a form of Dryas

    tundra with Salix rotundifolia , Minuartia arctica , M. macrocarpa , and Pedi

    cularis capitata. Close to the shore of the Yenisei a drier type of polygon

    tundra and Dryas tundra occurs.

            On the slopes down to the great river grow Equisetum arvense , Melandrium

    , Androsace septentrionalis , Castilleja pallida elegans , Erigeron

    , Artemisia tilesii , and others.

            On the sand banks at the mouth of the Yenisei are found Salix reptans ,

    Polygonum laxmannii , P. viviparum , Astragalus alpinus , Matricaria ambigua ,

    and other species. The last Alnus thickets are found a little north of 70°,

    but Salix thickets extend considerably farther north.

            The islands in Yenisei Bay, Minin (Olenii), Sibiriakov, and Dickson,

    were investigated by Kjellman and Tolmachev. Sibiriakov Island is almost

    in its entirety a monotonous swamp tundra, with Eriophorum angustifolium ,

    E. scheuchzeri , Dupontia fischeri , Arctagrostis latifolia , Luzula confusa ,

    Poa arctica , and Calamagrostis neglecta borealis , and, in wettish places,

    Ranunculus pallasii and R. hyperboreus . On the seashore Puccinellia phry

    ganodes , Carex subspathacea , and Stellaria humifusa occur.

            Dickson Island and N M inin Island are covered with wet polygon tundra.

    On the west coast of Dickson Island there is a drier rock tundra with

    070      |      Vol_VI-0400                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of the Soviet Arctic

    Alopecurus alpinus , Deschampsia alpina , Luzula confusa , Salix polaris ,

    Draba , Saxifraga oppositifolia , and Sieversia glacialis .

            The Gydan tundra is covered with more or less wet polygon tundra,

    Dryas tundra, moss and lichen tundra.

            Taimyr Peninsula was first investigated botanically by Middendorff in

    1843. He crossed the central part and proceeded to the Kara Sea. During

    the Vega expedition in 1878, Kjellman botanized at Cape Minin, Cape Chelyus–

    kin, and at Preobrazhenie Island. In 1918-25 the Norwegian North Polar

    expedition with the Maud visited several places on the coast. In 1928

    Tolmachev investigated the area south of Lake Taimyr, the largest Lake in

    the Eurasiatic Arctic.

            The part north of a line from about 71° N. at the Yenisei, south of

    Lake Taimyr, to the mouth of Khatanga River belongs to the high-arctic zone,

    through which run the 800 to 900-meter high, botanically almost unknown,

    Byrranga Mountains. Probably their slopes are for the most part without

    vegetation. [ ?]Polygon tundra and Dryas tundra are the most conspicuous

    vegetational types. Dryas tundra is especially common in the interior.

            South of the high-arctic tundras there is a belt of lichen-moss tundras

    100 to 200 km. broad, and still farther south a shrub tundra belt 50 to

    150 km. broad is found, with a narrow (less than 50 km. borad) strip of

    forest tundra south of it.

            A larch with a trunk 120 to 230 cm. high was found by Tolmachev at

    Novoi River, a tributary of the Khatanga River, 72° 40′ N. latitude. At

    72° 12′ N., on the Khatanga the larches are 6 to 8 meters high and often

    30 cm. thick.

    071      |      Vol_VI-0401                                                                                                                  
    EA-PS. Hult e é n: Phytogeography and Vegetation of Soviet Arctic

            Probably single larches grow a few minutes farther north to the

    east of the Khatanga River. These trees and forests are the most northerly

    on earth.

            The timber line to the west of Piasina River is composed of Larix

    , and to the east of that river of L. dahurica . See map fig.

    Between the Piasina and the Khatanga the boundary of the forest is well

    marked. Low espalier larches penetrate 10 to 12 km. north of it. Larix

    seems to be receding while L. dahurica seems to move forward.

    Betula tortuosa and Picea abies obovata extend eastward to Norilsk but

    they are lacking in the Khatanga basin. At Norilsk and east of Khatanga

    the shrub tundra borders the mountains. The n N orilsk Mountains have a vege–

    tation similar to that of the northern Urals, with single larch trees on

    the plain and in the lower valleys, a subalpina belt of Alnus crispa and

    Salix shrubs above this lichen-moss tundras, and lastly rock screes with

    lichens on the summits. The vegetation of the mountains south of the

    Khatanga mouth is unknown. There is, no doubt, a belt of Alnus there too.

            Anabar Tundra . The Anabar region was investigated by Toll in 1893, by

    I. P.