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Phytogeography and Vegetation of the Soviet Arctic Region: Encyclopedia Arctica 6: Plant Sciences (Regional)
Stefansson, Vilhjalmur, 1879-1962

Phytogeography and Vegetation of the Soviet Arctic Region

EA-Plant Sciences
(Eric Hult e é n)

PHYTOGEOGRAPHY AND VEGETATION OF THE SOVIET ARCTIC REGION

CONTENTS
Page
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
12
A. Circumpolar Plants 12
B. Eurasiatic Plants 18
II. Plants occurring in middle part of the Soviet Arctic
but not at both ends
19
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
21
A. Plants reaching the western but not the eastern end of
the Eurasiatic Arctic
22
B. Plants reaching the eastern but not the western end of
the Eurasiatic Arctic
28
IV. Endemic Plants 36
V. Singular Species 39

EA-PS.
(Eric Hult e é n)

CONTENTS-continued
Page
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

EA-Plant Sciences
(Eric Hulten)

MAPS
With the manuscript of this article, the author submitted 4 map
sketches. These are being held at The Stefansson Library.

EA-Plant Sciences
(Eric Hult e é n)

PHYTOGEOGRAPHY AND VEGETATION OF THE SOVIET ARCTIC REGION
LIMITATION OF THE ARCTIC ZONE
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.

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

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,

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–
ley.

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

HISTORICAL
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;

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
tundras.
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.
GEOLOGY AND SOIL
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

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

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.
CLIMATE
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.

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

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.
THE FLORA
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
arctica
, 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.

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.

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

Distribution of Species
I. PLANTS OCCURRING THROUGHOUT THE SOVIET ARCTIC OR AT BOTH THE EASTERN
AND WESTERN ENDS
A. CIRCUMPOLAR PLANTS
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

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

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

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

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

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.

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

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)

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)

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
species.
B. EURASIATIC PLANTS
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

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)
II. PLANTS OCCURRING IN MIDDLE PART OF THE SOVIET ARCTIC BUT NOT AT BOTH ENDS
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.
A. SPECIES NOT APPARENTLY WASHED DOWN BY THE RIVERS
Bromus sibiricus (arctic Urals)
Agropyron boreale (arctic Siberia)
Carex redowskyana (lower Ob, Khatanga, and Lena rivers)

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)
B. SPECIES APPARENTLY WASHED DOWN BY THE RIVERS
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

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
III. PLANTS OCCURRING EITHER IN WESTERN OR EASTERN PART OF SOVIET ARCTIC
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
it.
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.

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

A. PLANTS REACHING THE WESTERN BUT NOT THE EASTERN END OF THE EURASTATIC 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)

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

b. Occurring in Less Arctic Eastern and Western Ends of Kola
Peninsula
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
(arctic-montane)
Silene cucubalus
Agropyron mutabile (arctic-montane) Anemone nemorosa

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

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

Other Groups
Bromus inermis (central Asiatic) Paeonia anomala intermedia (central
Asiatic)
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)

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

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
European
Phleum alpimum commutatus (circumpolar
arctic-montane
Juncus triglumis (circumpolar
arctic-montane)
Puccinellia retroflexa suecica
(north European)
Primula stricta sens. str. (amphi–
atlantic, in Siberia P. stricta
jacutensis
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)

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
arctic-montane)
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
arctic-montane)
Pedicularis hirsuta (amphiatlantic, ? in Chukotsk Peninsula)
B. PLANTS REACHING THE EASTERN BUT NOT THE WESTERN END OF THE EURASIATIC ARCTIC
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

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
glandulosum
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

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

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
Alaska)
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
Alaska)
Carex vanheurkii
Polygonum pawlovskianum Draba juvenilis

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

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
circumpolar
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,
arctic-montane)
Potentilla stipularis (Asiatic)
Carex melanocarpa (Asiatic) Geum glaciale
Tofieldia nutans (nearly circumpolar) Pyrola secunda obtusata (nearly
circumpolar)
Rumex sibiricus (also in Alaska) Pedicularis lanata (nearly
circumpolar)
Minuartia arctica (also in Alaska)
Saxifraga punctata (arctic-montane,
also in America)
Crepis chrysantha (Asiatic,
arctic-montane)

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)

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

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

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)

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

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 )

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

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 )

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

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)
V. SINGULAR SPECIES
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)

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

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

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

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
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

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.

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

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,

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

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

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

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

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
globosum
, 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

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
fischeri
, Hierochloe pauciflora , Salix reptans , Caltha palustris , Comarum
palustre
, 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

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
crispa
, C. cucullata , Cladonia gracilis , Dicranum congestum , Dufouraea
arctica
, Hylocomium proliferum ( splendens ), Polytrichum hyperboreum , Ptilidium
ciliare
, 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

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
sylvatica
, 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
cucullata
, C. crispa , Cladonia pleurota , C. gracilis , Dicranum elongatum ,
Dufourea arctica , Polytrichum hyperboreum , P. piliferum , and Rhacomitrium
hypnoides
( lanuginosum ), the last-mentioned being especially common in the
west. The vascular plants consist of scattered specimens of Hierochloe
alpina
, Arctagrostis latifolia , Luzula confusa , Pedicularis hirsuta , Parrya
nudicaulis
, 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
alpina
. 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

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
aphtosa
. 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

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
vaginatum
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

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

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
vitis
- 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

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

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
aquatilis
, 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
vulgaris
. 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.

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
strictum
, 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
aphtosa
, 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

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
nana
, 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
sibirica
, 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

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
graminifolia
, 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.

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.

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
rariflora
community covers large areas. On the slopes shrubs such as Salix
glauca
, S. lanata , and S. myrsinites with Calamagrostis are common. In Andreev
(1931) a fairly detailed account is given of the different small sociological
units.
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
longifolia
, Pedicularis sceptrum - carelinum , Achillaea millefolium , and
Solidago virgaurea , some species even reaching northern Kanin, e.g., Parnassia
palustris
, 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

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
nana
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.

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
frigidus
is common on all plains. In wet places Carex aquatilis , Colpodium
fulum
, 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
ambigua
, and in the salt marshes Puccinellia vilfoidea , Carex subspathacea ,
and Stellaria humifusa .

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 .

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.

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
hieracifolia
, 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
pallasii
, 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

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
nivalis
, Oxytropis campestris sordida , Myosotis alpestris , Eritrichium
villosum
, 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
acaulis
, 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
subcapitata
, Saxifraga oppositifolia , and Potentilla hyparctica predominate.
Po a , Papaver , and Potentilla form a community alternating with Saxifraga
oppositifolia
.

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
asiaticus
, Parnassia palustris , Rubus arcticus , Viola biflora , Epilobium
dahuricum
, 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
rivularis
, 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

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
turgidum
, 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
pubescens
, 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

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
furcatum
, Androsace septentrionalis , Castilleja pallida elegans , Erigeron
eriocephalus
, 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

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.

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
sibirica
, 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
sibirica
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. Tolmachev in 1905, and by Sochava and Efimov in 1932.
The Anabar tundra is a mesozoic plain covered with innumerable small
lakes, mostly less than 800 meters across.
Sochava has given a very detailed review of its phytogeographcal con–
ditions, with descriptions of all plant communities. Anabar, which until
recently was one of the phytogeographically least known parts of the Arctic,
is now particularly well known. Sochava described 76 different associations,
of which 37 are from the high-arctic tundra and the lichen tundra.

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

In the high-arctic zone along the arctic shore (north of the 7° isotherm
for July) the soil thaws down only to 25 to 35 cm. The precipitation is lower
and the winds less strong than in the same zone farther west. Dryas octo–
petala punctata
is the most characteristic plant in the entire zone, having
the widest ecological amplitude, from rocks to swamps and from [: ] muck to sandy
soil.
Sphagnum ( S. girgensohnii , S. squarrosum , S. teres ) is a floristic
rarity, A Dryas tundra containing Cetraria cucullata , Hierochloe alpina,
Carex rupestris
, Diapensia , and Cassiope occupies the largest areas. Lichen
tundra with Alectoria ocroleuca and Cornioularia divergens is found on hilltops
only. Swamps with Carex aquatilis stans and C. retundata are very commn.
Shrubs such as Betula nana exilis , Rubus chamaemorus , Empetrum , Vaccinium
vitis
- idaea minus , V. uliginosum , and Arctostaphylus alpina are lacking within
the high-arctic zone. The shrubs consists of Salix arctica , S. polaris, S. pul
chra , and S. rotundifolia . Moss tundra is rare. Its most common species are
Rhytidium rugosum , Aulacomnium turgidum , and Hylocomium proliferum ( splendens ).
The swamps have a network of 70 to 100-cm.-high peat walls, or tussocks of
that height, covered with Carex peat.
In the lichen-moss tundra zone, which occupies a belt about 100 km. broad
south of the high-arctic zone, the Alectoria tundra occupies the largest areas
on hilltops and on sandy soil. Within it are found Luzula confusa , Polygonum
pawlovskianum
, and Cassiope tetragona. Cetraria tundra occurs on muddy or
clayey soil. Cladonia societies are poorly represented. Eriophorum vaginatum ,
Carex aquatilis stans , and Betula nana exilis are prevalent inthem. Moss
tundra with Ptilidium ciliare and Dryas tundra occurs here and there. Betula

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

nana exilis nana exilis is common, but does not predominate. Eriophorum vaginatum
tussock swamps occur, but is more common in the shrub zone. In grass swamps
with stagnant water Carex rotundata is prevalent (lacking farther north) and
in flowing water Carex aquatilis stans . There is no Polytrichum tundra in
Anabar.
In the shrub zone, which is somewhat narrow, Salix s hrubs, chiefly
Salix pulchra , predominate, together with Betula nana exilis. Calamagrostis
canadensis langsdorffii
, Salix alaxensis , Aconitum , and Rosa acicularis are
characteristic plants in this zone. Eriophorum vaginatum tussock tundra is
common. Sphagnum plays a larger part that it does farther north.
The most northerly trees, which are invariably larch, almost entirely
lack branches, and look like striaght poles. In the northern part of the
forest tundra the trees stand 30 to 40 per hectare and have very poorly devel–
ped crowns. The northern limit of Alnus usually runs along the northern timber
line, though low suppressed Alnus shrubs can be found even in the lichen tundra.
It is lacking along the rivers where Salix alaxensis and S. lanata occur.
Mouths of Olenek and Lena Rivers . The mouth of the Olenek was investigated
by Chekanovsky and Müller in 1874 and by Chekanovsky in 1875. The lower Lena
was first visited for botanical purposes by Adams in 1806, later by Bunge in
1883, Nilsson-Ehlein 1889, Cajander and Poppius in 1901, and by several others.
Nevertheless the Arctic at the Olenek and Lena is but little known. The high–
arctic tundra occupies only a very narrow strip about 100 km. broad and 250 km.
long along the Arctic Sea at the Olenek and the delta of the Lena River. The
lichen-moss tundra zone extends eastwards to the Olenek River, but the tundra
between the Olenek and Lena is replaced by a mountainous country with Carex

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

mires on the divides and meadows in the valleys, of which very little is
known. The shrub tundra likewise stops at the Olenek, where larches grow
down to about 100 km. from the mouth and ascend the eastern tributaries of
the river.
The Lena Delta has a very poor vegetation, although about 200 flowering
plants are known from there, most of them from the southern and eastern parts.
In the lower Lena Valley, which is surrounded by mountains, the last
larch is found at Tit-ary on 72° N., i.e., at the beginning of the delta.
No shrub zone is found there. The trees are crippled, being 3 to 5 meters
high and covered with lichen. Salix , Betula , Ribes triste , Rosa acicularis ,
Vaccinium uliginosum , and Cassiope tegragona are found in their undergrowth.
Lena to Kolyma and Chaunskaia Bay . The first botanical investigations
were made in 1784 by Merck, who visited Nizhne Kolymsk. In 1865-66, Maydel
explored the shore between the Yana and Indigirka, and in 1875 Augustinovich
visited the lower Kolyma River.
The high-arctic tundra belt is about 100 km. broad; lichen-moss tundra
is lacking or plays a small part. The shrub tundra is also about 100 km.
broad.
The vegetation is very little known. Large areas are occupied by
Eriophorum vaginatum tussock-swamp tundra with bare earth between the tussocks.
Large areas are distitute of vegetation. At least to the west, green mosses
but few Sphagna occur in those swamps and the peat layer is not over 10 cm.
deep. The tundra thaws to a depth of only about 25 cm. Large areas are
covered with Cladonia tundra, C. rangiferina predominating. Typical polygon
tundra seems to be lacking or rare. The vegetation of the mountains, which
enter the Arctic east of the Yana and on both sides of Chaunskaia Bay, is

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

unknown. In this Verkhoyansk Mountains open larch woods with single pines and
a lower layer of Alnus , Betula , and Salix predominate.
The New Siberian Islands were investigated botanically by Bunge and Toll
in 1885-86. They consist of four large and several small islands, all within
the high-arctic zone. The largest, Kotelnyi Island, is about 150 km. across.
The vegetation is very little known. Snow field and open soil are com–
mon, and no drawf shrubs are present nor vegetation mats. Patches of vegeta–
tion and single Eriophorum tussocks occur, also fragments of lichen tundra.
In some places Ranunculus nivalis ( sulphureus ?) predominates. Saxifraga [: ]
groenlandica , S. hirculus , Papaver radicatum , Oxyria luzula , and Sieversia
glacialis
occur. Cochlearia is found on the shore.
Bennett, Zhokhov, and Henrietta Islands, north of the New Siberian
Islands, are entirely covered with ice.
Chukotsk Peninsula . The first botanical observations on the Chukotsk
Peninsula were made by Chamisson in 1816 at St. Lawrence Bay. In 1827 Mertens
visited the Bering Strait coast. During the Expeditions of Commander Ringgold
and Rogers in 1853-56, with C. Wright as botanical collector, Arakamchechen ?
Island was visited.
In 1879 Kjellman wintered with the Vega Expedition at Pitlekai. He also
collected at Cape Yakan, Irkaipij (Cape Schmidt), Konyam Bay and at St. Law–
rence Bay. In 1881 the brothers Krause investigated the Bering Strait coast,
and in 1934 Gorodkov worked at Cape Schmidt and Cape Vankarem on the north
coast, making short stops at St. Lawrence Bay and Port Providence.
The high-arctic tundra stops at about 175° E. The entire peninsula
west of the Stanovoi Mountains and southward to the mouth of the Anadyr River

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

belongs to the lichen-moss tundra zone. The peninsula is mountainous toward
its eastern end
It Real polygon tundra is lacking in Chukotsk Peninsula. Cladonia tundra
and Sphagnum play a large part in the vegetation, as in Kola Peninsula, contrary
to what is the case in the central parts of the Arctic zone. On the north
coast of the Chukotsk Peninsula special tundras occur where patches of bare
earth alternate with drier or wetter patches of vegetation containing Alectoria
nigricans
, Ochrolechia tartares , Rhacomitrium , Thamnolia vermicularis , and the
mosses Aulocomnium turgidum , Camptothesium trichoides ( Tomentophytum nitens ,)
Dicranum elongatum , Drepanoclades uncinatus, and Hylocomium proliferum ( splen
dens ). In these patches occur also vascular plants, such as Carex lugens ,
C. misandra , Salix pulchra , S. reptans , Ranunculus nivalis , Parrya nudicaulus ,
Dryas punctata , Sieversia glacialis , Oxytropis nigrescens , Pedicularis lanata ,
and Artemisia trifurcata .
The moss tundra is common on slopes and between the tussock swamps. Prom–
inent mosses are Camptothecium trichoides ( [: ] Tomenthypnum nitens ), Dicranum
elongatum
, Hylocomium proliferum , and Ptilidium ciliare . Lichens are of very
little significance. Among the herbs, Eriophorum angustifolium , Carex bigelowii ,
C. aquatilis stans , Polygonum viviparum , Salix pulchra , Dryas punctata , and
others occur. Dry moss tundra, which is so common west of the Lena River, is
lacking in the eastern part of the peninsula but occurs, at least to some ex–
tent, in the western parts. In the lichen tundra Cassiope tetragona , Rhodo
dendron kamtschaticum glandulosum , R. lapponicum , and Saxifraga bronchialis
funstonii
occur with Cetraria cucullata , which predominates especially in the
mountains. A characteristic feature of northeastern Asia is the vast areas of

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

Eriophorum vaginatum tussock tundra. They reach as far as to the northern
coast of Chukotsk Peninsula. The tussocks. 20 to 30 cm. high and 30 to 40
cm. broad, cover about 40% of the tussock tundra. Between them Calliergon
sarmentosum
and Polytrichum jensenii are common. Several mosses and lichens
are found on the top of the tussocks. The following herbs are found there:
Salix phlebophylla , S. pulchra , Vaccinium vitis - idaea minus , Carex lugens ,
Juncus biglumis , Pedicularis langsdorffii .
In the meadows many plants not otherwise occurring in the Arctic are
found, such as Carex scirpoidea , Salix chamissonis , Anemone richardsonii ,
Trollius membranostylis , Parnassia kotzebuei , and Gentiana glauca .
In the mountains there is a subalpine belt of Alnus crispa and Pinu s
pumil a. Pinus is more frequent in the upper part of the zone. It is lacking
in the extreme northeast, where it is replaced by Salix shrubs. Along the
seashores the subalpine belt is lacking, being replaced by Salix alaxensis
and S. pulchra shrubs about 1 meter high. The undergrowth in the Pinus
thickets is very varied; it may consist of lichens, but of Sphagnum as well.
Open spots in which grow Calamagrostis canadensis langsdorffii , Bromus
sibiricus
, Poa alpigena and other species alternate with the thickets in
their lower parts; higher up the Pinus shrubs alternate with lichen-moss tundra.
At 150 to 250 meters the alpine zone begins with lichen tundra between
the rocks. Cladonia and Alectoria ochroleuca alternate.
On the north coast there are low dunes where Elymus , Hockenya , Cochlearia ,
Arctophila , Catabrosa algida , Artemisia tilesii , and others occur. A vegeta–
tion very similar to that described above stretches southward to Gizhiga and
northern Kamchatka. It is only in the inundation zone of the large river

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

valleys that fringing forests of Salix macrolepis , Populus suaveolens , and
Betula cajanderi occur here and there.
As the section on the flora indicates, the floristic composition in the
Chukotsk Peninsula is very different from that in other parts of the Soviet
Arctic. With the exception of Pinus pumila , the alien components, which are
Pacific, Eastern Asiatic-Western American, or America, play but little part
in the vegetation, which connects up with that of the Eurasiatic Arctic.
Wrangel Island was recently investigated by Gorodkov (unpublished ?).
Before that, only scattered notes on the flora were available. This moun [: ]
tainous island is about 150 km. long. It is regarded by Soviet authors as
belonging to the snow zone, but its rich flora, 150 species, make this doubt–
ful. Probably the low parts of the island should be classed within the lichen–
moss tundra zone, while even low mountains come within the snow zone. Such
species as Salix glauca , S. reptans , S. pulchra , Bromus pumpellianus arcticus ,
Aconitum delphinifolium , Saxifraga bronchialis , Funstonii , Androsace chamaejasme
lehmaniana
, Cassiope tetragona , Vaccinium vitis - idaea , Valeriana capitata , Pedi
cularis verticillata , Aster casaguineus , Saussurea angustifolia , and Senecio
resedifolium
could hardly be expected in the snow zone. The vegetation is un–
known. From Herald Island, east of WrangelIsland, only single plants are
known, while from Big Diomede Island in Bering Strait no botanical data have
been obtained, although the flora of Little Diomede Island a few km. farther
eastward, which belongs to Alaska, was investigated by Porsild.

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

BIBLIOGRAPHY
General
Almquist, E. B. Lichenologiska iakttagelser på Sibiriens nordkust (Lichenological
observations on the North Coast of Siberia). Sv. Vet.
Akad. Förh. Öfvers. 36(1879) Nr 9, pp. 29-59. In German in
Nordenskiöld, A.E. Wissensch. Ergebn. d. Vega Exped. 1 (1882-83)
pp. 50-74. Andersson, G. Zur Pflanzengeographie der Arktis. - Geogr. Zeitschr. 3(1902)
pp. 1-23. Andreev, V.N. Podzoni tundry Severnago kraja. (Underzones in the tundras of
the northern district). - Priroda 1932 No 10 pp. 889-906. Brotherus, V.F. Die Moose des arktischen Küstengebietes von Sibirien nach
der Sammlung der russischen Nordpolar Expedition 1900-1903. -
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(1879) pp. 1-40. ----. Plantae in littore asiatico freti Behringii anno 1876 a
Skalowski lectae. - Acta Hort. Petrop. 8 (1883) pp. 1-22. Vainio, Edv. A. Lichenes in viciniis stationis hibernae expeditionis Vegae
prope pagum Pitlekai in Sibiria septentrionali a D:re E.
Almquist collecti. - Ark. f. Bot., Bd 8:4. Uppsala 1909.
Wrangel Island
Gorodkov, B.N. Manuscript list of the Flora of Wrangel I. Nazarow, M.I. K flore ostrova Wrangelja (To the flora of Wrangel I.) - Hydro–
meteorolog. Committee USSR. III:4 (1934) pp. 1-21.
Eric Hult e é n
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