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Phytogeography and Vegetation of the Bering Sea District: Encyclopedia Arctica 6: Plant Sciences (Regional)
Stefansson, Vilhjalmur, 1879-1962

Phytogeography and Vegetation of the Bering Sea District

EA-Plant Sciences
[Eric Hulten]

PHYTOGEOGRAPHY AND VEGETATION OF THE BERING SEA DISTRICT

TABLE OF CONTENTS
Page
CHARACTER AND DISTRIBUTION OF THE FLORA 1
Circumpolar Plants 2
Arctic Montane Plants 4
American and Eurasiatic Plants 6
Endemic Plants 11
Chukotsk Peninsula 11
Anadyr 11
Aleutian Islands (including Pribilof Islands) 11
Bering Strait District 12
Shores and Islands of the Bering Sea 12
VEGETATION 14
Chukotsk Peninsula 16
Anadyr 16
Penzhina-Gizhiga 18
Kamchatka 18
Commander Islands and the Aleutian Islands 20
Alaska Peninsula 20
Eastern Shore of the Bering Sea 21
Islands in the Bering Sea 22
BIBLIOGRAPHY 24

EA-Plant Sciences
[Eric Hulten]

PHYTOGEOGRAPHY AND VEGETATION OF THE BERING SEA DISTRICT
There can be no doubt that the Bering Sea district occupies a singular
position phytogeographically in the arctic and boreal belts. The distribution
patterns converge toward the Bering Sea. This is certainly not an incidental
phenomenon but the result of the large migrations between Asia and America which
have swept over the region in past geological ages. The Bering Sea area was
to a large extent unglaciated during the Pleistocene glaciations and the climatic
changes there during the Quaternary seem to have been less severe than in most
other places in the arctic and boreal world. Ancient plant populations have
therefore remained more or less undisturbed, or at any rate less depauperated
there than in most places elsewhere. The result is that the vari t ation of species
is larger in the Bering sea area than in most other place in the same latitude.
CHARACTER AND DISTRIBUTION OF THE FLORA
The distributional groups that meet in the Bering Sea area are the following:
The arctic and boreal circumpolar groups.
The circumpolar arctic-montane group.
The boreal Eurasiatic and arctic montane Eurasiatic groups.
The arctic and boreal American groups.
The Eurasiatic-western American group.

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The eastern Asiatic-western American group.
The American-eastern Asiatic group.
The eastern Asiatic and eastern Asiatic Pacific groups.
The western American and western American-Pacific groups.
The northern Pacific group.
The central Asiatic group.
Besides these, there is a considerable endemic group. containing old and
well differentiated endemic species.
Of these the circumpolar groups and the Asiatic-American groups very often
have a more or less large gap in the area of the Bering Strait district, or some–
times the gap is still larger, even comprising the entire Bering Sea area. Plants
belonging to these groups once migrated over Bering Strait when the climate was
more genial than it is now, but they were exterminated there when the climate
deteriorated.
Circumpolar Plants
Many of the circumpolar plants nowadays have a continuous distribution over
Bering Strait. To them belong, among others:
Lycopodium annotinum Saxifraga hirculus
Lycopodium clavatum Parnassia palustris sens.lat.
Lycopodium selago Potentilla palustris
Equisetum arvense Hippuris vulgaris
Equisetum pretense Empetrum nigrum
Eriophorum angustifolium Andromeda polifolia
Luzula multiflora Vaccinium uliginosum

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Allium schoenoprasum Vaccinium vitis-idaea
Stellaria crassifolia Adoxa moschatellina
Caltha palustris Calium boreale
Cardamine pratensis
Circumpolar plants with a gap at Bering Strait are, among others:
Juniperus communis montana Lathyrus palustris
Potamogeton graminous Drosera rotundifolia
Carex limosa Campanula rotundifolia sens.lat.
Scirpus peluatris Naumburgia thyrsiflora
A very large gap in the Bering Sea area contains, among others, the following
circumpolar plants:
Equisetum hiemale Carex lasiocarpa
Dryopteris spinulosa Carex vasicaria
Dryopteris thelyoteris Calla palustris
Pteridium aquilinium Lemna minor
Scheuchzeria palustris Lemna tri s culca
Potamogeton praelongus Ceratophyllum demersum
Potamogeton zosterig l olius Ranunculus paucistamineus
Alisma plantago-quatica Dorsera anglica
Phalaris arundinacea Potentilla norvegica sens.lat.
Poa palustris Rubus idaeus sens.lat.
Phragmites communis Myriophyllum verticillatum
Carex diandra Utricularia minor

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In some cases plants of the same general type but belonging to different
species occur on both sides of the Bering Sea, as for instance in the case of
Populus where the Eurasiatic P. tremula is replaced by P. tremiloides in America.
The arct
Arctic Montane Plants
The arctic-montane plants usually have a continuous distribution in the
Bering Strait area but with gaps in the Aleutain Islands, where there are very
few arctic-montane plants with a continuous distribution. [: ] A only the most fre–
quent arctic-montane species with a good spreading capacity occur all along the
Aleutian chain, such as:
Lycopodium alpinum Polygonum viviparum
Phleum alpinum Sibbaldia procumbens
Luzula perviflora Louiseleuria procumbens
A relatively small gap in the Aleutians contains the following arctic-montane
plants:
Polystichum lenchitis Juncus castaneus
Cryotogramma achrostichoides Salix roticulata
Hierochloe alpina Vernnica wormskjoldii
A larger gap there has the arctic-montane species enumerate below:
Dryopteris fragran Ranunculus hyperboreus
Cvstopteris Montana Ranunculus pygmaeus
Woodsia alpina Thalictrum alpinum
Woodsia glabella Draba alpine

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Eriophorum opacum Saxifraga foliolosa
Eriophorum scheuchzeri Saxifraga nivalis
Arctagrostis latifolia Saxifraga oppositifolia
Cobresia myosuroides Saxifraga cernus
Carex misandra Sedum roseum
Carex rotundata Potentilla nivea
Juncus biglumis Drayas
Lugula spicata Astragalus alpinus
Salix glauca Epilobium anagallidifolium
Koenigia islandica Epilobium dayuricum
Rumex acetosa alpina Arctostanphylos alpina
Minuartia biflora Phyllodoce coerulea
Minuartia verna Diapensia lapponica
Melandryum apetalum Pleurogyne rotate
Melandryum furcatum Gentiana tenella
Sagina linnaei Pinguicula villosa
Silene acaulis Artemisia borealis
Some arctic-montane plants pass over Bering Strait but do not even approach
the Aleutians, such as
Poa alpina Minuartia stricta
Carex capitata Rhododendron lapponicum
Carex microglochin Cassiope tetragona
Carex rupestris Pedicularis lapponica
Cobresia simpliciuscula

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These conditions might indicate that the arctic-montane group is very old
and that is comparatively recent time its members have migrated to the Aleutians
both from the east and from the west.
Another explanation might be that the extremely maritime climate of the
Aleutians does not suit them, but this does not seem very probable as they in–
habit the arctic shores with somewhat similar climatic conditions, although
with less precipitation and cold winters.
American and Eurasiatic Plants
Bering Strait itself is a shallow, rather narrow sound, only about 50 meters
deep [: ] and 56 miles (90 km.)broad, with the two Dio [: ] ede Islands in the middle.
It does not constitute a phytogeographical boundary line, the flora on its eastern
and western shores being very similar. However, many American plants reach west–
ward to central Alaska or a little farther, and many Asiatic plants as far as
to the Anadyr Valley.
The American plants reaching the Seward Peninsula but absent in Asia are
enumerated below. Most of them do not reach the shores of Bering Strait.
Picea mariana Platanthera obtusata
Calamagrostis inexopanss Pipulus tacamahacca
Festuca saximontana Salix fullertonensis
Zygadenus elegans Salix richardsonii
Cypripedium passerinum Nuphar polysepalum
Anemone drummondii Cicuta mackenzieana
Aphargmus eschscholtzianus Gentiana arctophila
Therefon richardsonii Polemonium pulcherrimum

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Saxifraga spicata Mertensia peniculata
Saxifraga tricuspidata Mertensia eastwoodae
Sorbus scopulina Pentstemon procerus
Potentilla virgulata Castilleja hyperborea
Lupinus arcticus Utricularia macrorhiza
Astragalus eucoemus Galium brandegei
Oxytropis gracilia Viburnum peuciflorum
Shepherdia candensis Petasites hyperboreus
Bupleurum americanum Senecio lugens
Of these, Calamagrostis inexpansa , Festuca saximontana , Salix fullertonensis ,
Oxytropis gracilis , Mertensia peniculata , M. eastwoodae , Castilleja hyperborea ,
Galium brandegei , and Petasites hyperboreus belong to critical groups and might
well be found also in easternmost Asia. Zygadenus elegans , Platanthere obtusata ,
Sorbus scopulina , Astragalus eucosmus , Bupleurum americanum , and Utricularia
macrorhiza
have closely related Asiatic counterparts, and Aphragums eschscholtzianus
is an inconspicuous plant easily overlooked. The most remarkable species in the
list are Picea mariana , Populus tacamahacca , Lupinus arcticus , Shepherdia cana–
densis
, and Viburnum pauciflorum . These play a more or less prominent part in
the vegetation and are important purely American components in the flora of the
American side of the Bering Sea distirct.
Asiatic plants reaching the Chuktosk Peninsula but not occurring in America
are the following:

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Pinus [: ] pumila Polygonum peylowskianum
Kobresia shoenoides Polygonum undulatum
Salix pyrolifolia Apemone debilis
Dicentra peregrina Gentiana auriculata
Corydalis ambigua Eritrichium villosum
Ermania parryoides Crepis chrysantha
Oxytropis revuluta
Of these only Pinus [: ] pumilla and Polygonus pavlowskianum play any impor–
tant part in the vegetation.
Many American plants cross the Bering Strait and occupy small areas [: ] in
easternmost Asia, for example:
Salix Richardsonii Oxytropis Maydelliana
Parnassia Kotzebue i [: ] Gentiana propinqua
Dryas integrifolia Chrysanthemum integrifolium
Potentilla Vahliana
Many other American species penetrate farther westward into Asia.
On the other hand, many Asiatic species have crossed over to America, [: ]
such as:
Claytonia eschscholtzii Potentilla elegans
Oxygraphis glacialis Saxifraga nudicaulis
Delphinium brachycentrum Phlox sibirica
Cordydalis pauciflora Primula nivalis
Potentilla biflora

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A number of Asiatic species occupy isolated areas in interior Alaska and
Yukon, e.g.:
Carex bonazensis Ranunculus gelidus
Juncus leucochlamys Trifolium lupinaster
Luzula rufescens Cnidium ajanense
Cerastium maximum Artemisia laciniata
Stellaria Laxmannii Artemisia macrobotrys
Arenaria capillaries
Many other Asiatic plants occupy larger areas in western America.
From the above review the insignificance of Bering Strait as a phytogeo–
graphical boundary line should be clear.
The second connection between Asia and America at present time is the
Aleutian Island, investigated botanically by the present writer in 1932. The
most outstanding component in the flora is the Pacific group, containing plants
distributed along the northern shores of the Pacific Ocean and sometimes protrud–
ing northward also along the shores and island of the Bering Sea. To this
group belong:
Poa hispidula Rubus stellatus
Poa turneri Geum rotundifolium
Deschampsis beringensis Rhododendron kamtschaticum
Carex macrochaeta Phyllodoce aleutica
Carex lyngbyaei cryptocarna Cassiope lycopodioides
Carex pauciflora Trientalis europaea arctica
Orchis aristata Primula cuneifolia

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Salix crassijulis Pedicularis chamissonis
Fritillaria kamtchatcensis Veronica stelleri
Cerastium fischerianum Plantago macrocarpa
Aconitum maximum Campanula dasyanthe
Draba hyperborean Arnica unalaschcensis
Chrysosplenium beringianum Taraxacum trigonolobum
Saxifraga bracteata Hieracium triste
Potentilla nana
Other groups occur mostly at the eastern or at the western end of the
Aleutian chain or at both ends. The middle part of the chain is thus much
poorer in species than the two ends. At the western end of the chain a few
Asiatic species occur which are common in Kamchatka but are not otherwise
found in America. These are:
Veratrum album oxysepalum Sorbus sambucifolia
Allium victorialis platyphylla Primula cuneifolia Dubyi
Cypripedium yatabeanum Mertensia asiatica
Platanthera tipuloides Senecio palmatus
Ranunculus acris frigidus Cirsium kamtschaticum
Cardamine regelliana Picris hieracioides kamtchatica
Aruncus sylyester Calcalia suriculata

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Endemic Plants
More than 200 endemic plants are known from the Bering Sea area in a wide
sense, including Alaska, Yukon, Anadyr, and Kamchatka. Many of them are very
well-marked species, while others, such as those belonging to the apomictic genera
Antennaria and Taraxacum , are very closely related to one another and to more
widespread types. An enumeration of the endemics in some of the districts is
give below.
Chukotsk Peninsula Chukotsk Peninsula
Carex arakemensis Draba chamissonis
Trollius chartosepalus Artemisia laciniatiformis
Anadyr Anadyr
Cardamine victoris Sorbus anadyrensis
Calamagrostis czykczorum Potentilla anadyrensis
Poa soczawai Ranunculus anadyriensis
Salix anadyrensis Claytonia vassilievii
Agrostis anadyrensis Oxytropis dorogastajskyi
Aleutian Islands (including Pribilof Islands) Aleutian Islands (including Pribilof Islands)
Polystichum aleuticum Draba aleutica
Calamagrostis purpurascens arctica Saxifraga aleutica
[: ] Poa turneri Saxifraga punctata insularis
Elymus aleuticus Artemisia aleutica
Stellaria ruscifolia aleutica Artemisia unalaskens is aleutica
Cerastium aleuticum Taraxacum chromocarpum
Ranunculus occidentalis nelsonii Taraxacum everdami
Papayer alaskanum Taraxacum onchophorum

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Bering Strait District Bering Strait District
Colpodium wrightii Smelowkia calycina intergrifolia
Carex jacobi peteri Primula paucifolia
Arenaria dicranoides Artemisia senjavinensis
Melandrium soczavianum Erigeron muirii
Ranunculus glacialis chamissonis
Papaver [: ] walpolei
Shores and Islands of the Bering Sea Shores and Islands of the Bering Sea
Puccinellia laeviuscula Artemisia globularia
Alopecurus alpinus stejnegeri Saussurea viscida
Carex pribylovensis Taraxacum angulatum
Carex karaginensis Taraxacum callorbinorum
Epilobium behringianum Taraxacum chamissonis
Pedicularis penellii Taraxacum demissum
Antennaria monocephala exilis Taraxacum fabbeanum
Artemisia arctica beringensis Taraxacum pribilofense
Along the southern coast of Alaska and especially in the unglaciated central
parts of Alaska and Yukon numerous endemics are found, which are not enumerated here.
To summarize the above: In the Bering Sea area a large number of species
survived the Pleistocene glaciations and later spread eastward as well as west–
ward from there. The areas of these species are usually arranged symmetrically
with Bering Strait as the center. American as well as Asiatic plants in remote
periods crossed Bering Strait or the northern part of the Bering Sea, which was
doubtless dry at certain periods before the last glaciations, and now have isolated
areas on the opposite continent is more southern latitudes.

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Many circumpolar plants have the same history here. The northern Pacific
component in the flora is probably very old and occupies the shored and islands
in the Bering Sea, chiefly the Aleutians, stretching far south as well in eastern
Asia as in western America. Many of them nowadays have gaps in their Aleutian
area caused by the deterioration of the climate. On the Asiatic side, eastern
Asiatic species penetrate northward toward Being Strait and eastward along the
Aleutians, just as western American plants penetrate northward along the shore
of the Bering Sea and westward along the Aleutians. In the interior of eastern–
most Siberia continental plants, mostly Siberian, penetrate eastward but most
of them do not reach the coast, just as continental American plants penetrate
westward through interior Yukon and Alaska, likewise for the most part not reach–
ing the Bering Sea. The great age of flora is emphasized by the high percent–
tage of endemics and by the often rather disrupted patterns of distribution.
Examples of species with very disrupted areas belonging to various distri–
butional groups not mentioned above are:
Scirpus rufus , occurring in Eurasia and America with a very isolated locality in
Alaska.
Danthonia intermedia , American species with very isolated localities in Alaska
and Kamchatka.
Streptopus streptopoides , occurring in southern British Columbia and Washington,
on the one hand, and in Sakhalin, Honshu, and Okhotsk, on the
other, with a very isolated locality in Alaska.
Malaxis [: ] paludosa , Eurasiatic species with a very isolated station in
Alaska and a few other places in America.
Smilacina trifolia , occurring in America and in eastern Asia with a very isolated
locality in Alaska.
Lycopus lucidus , occurring in western America and in eastern Asia with a very iso–
lated locality in Alaska.

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VEGETATION
The vegetation of the Bering Sea districts has so far been very little
studied. The mountains along the coast between Anadyr and Kamchatka are as
yet completely unknown botanically, and very few studies of the vegetation have
been made in Alaska.
The soil in permanently frozen in large parts of the northern Bering Sea
area, and, as the summer is rather cold, the soil does not thaw very deeply.
This prevents tree growths. (In places where the soil thaws deeper, for instance,
through fire in the Pinus pumila thickets, the forest grows well, a fact that
demonstrates the influence of the frozen soil.)
The area forms a treeless continuation southward of the arctic tundra belt.
The treeless area stretches from the Chukotsk Peninsula southward to northern
Kamchatka and along the western coast of that peninsula in Asia, and from Seward
Peninsula along the coast to the base of Alaska Peninsula and western Kodiak in
America. The islands in the Bering Sea as well as the Aleutians are treeless
(with the exception of a planted grove of Sitka spruce at Uhalaska). In S
Asia the Larix dehurica forest protrudes into the valleys of the Stanovoi Moun–
tains to slightly west of Markova in Anadyr and reaches the Okhotsk Sea somewhat
west of the Taigonos Peninsula. East of this limit there are alluvial forests
of Salix ( Chosenia) macrolepis , Populus suaveolens , and Betula cajanderi along the
large rivers. Kamchatka is phytogeographically and island, isolated from the
Siberian woods by a broad tundra belt. It is covered with forests of Betula
ermani
, but in its central part there are not very extensive forests of Larix
dahurica
and Picea jezoensis .

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On the American side there is an interior forest, chiefly of Betula
resinifera
, Picea glauca , and Populus tacamabacca , bordering in the most on
the treeless area, and a narrow southern coastal strip of forest, chiefly of
Picea sitchensis and Tauga heterophylla , reaching westward to eastern Kodiak
Island and the base of Alaska Peninsula bordering on the interior forest to the
north and on the treeless area to the west. In central Alaska an isolated area
of Larix laricina occurs which has been regarded as differing from the L. laricina
farther east. In southwestern Alaska, Detula kenaica occurs.
The nature of the large treeless area in the Bering Sea district has been
somewhat widely discussed. It seems to me that it is hardly possible to separate
it in principle from the circumpolar tundra belt. The floristic composition is
different, but this also is true of the Chukotsk Peninsula and northwestern Alaska,
which cannot be excluded from the circumpolar tundra belt. Most of the plants of
the arctic tundras occur also in the treeless Bering Sea area, but in addition
to them there are numerous Pacific, eastern Asiatic, and western American species
superimposed upon the arctic flora, and gradually increasing in number southward.
The heaths, meadows, and wet plant communities of the Arctic also occur to
a large extent in this treeless area and likewise on the mountains of Kamchatka,
the Aleutians, and the Alaskan mainland.
Consequently, the treeless area of the Bering Sea should be regarded as a
southern phase of the arctic tundra zone.

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Chukotsk Peninsula
Easternmost Asia is a mountainous peninsula belonging to the lichen-moss
tundra zone of the Arctic. Polygon tundra is lacking or nearly so. Cladonia
tundra and Sphagnum play a large part in the vegetation. Vast areas are covered
by Eriophorum vaginatum tussock-tundra. Pacific and eastern Asiatic species alien
to other parts of the Arctic occur especially in the meadows. In the mountains
lichen tundra predominates with Pinum pumila thickets up to 150 to 250 meters.
Anadyr
Only in the westernmost part of the Anadyr Valley to the eastern foot of the
Stanovoi Mountain chain do woods of Larix dahurica occur as a narrow strip in the
valleys along the rivers. In post-Pleistocene time Larix forests occurred also
in the valley of Belaya river 150 km. farther east of where they are found now.
A fossil fir, Picea anadyrensis , has been found in Anadyr in layers supposed
to be contemporary with the earliest [: ] Pleistocene glaciations.
East of the present larch woods the lowlands are covered with a vegetation
similar to that of the lichen-moss tundra zone of the Arctic. It is very uniform
and consists mostly of Eriophorum vaginatum tussock-tundra, or of boggy associa–
tions with Sphagnum between the herbs. Lakes bordered with Arctophila fulva and
Senecio congestus are common. The slopes along the shore are usually covered
with Alnus crispa thickets. The seas develop into swamps through the growth of
Carex aquatilis , C. rotundata , C. chordorrhiza , and Potentilla palustris . The
large river valleys are fringed with forests of Salix macrolepis , Populus suaveolens ,
and Betula cajanderi. Salix macrolepis always occurs at the very shore on alluvial
sand, and this tree and Populus , which occurs somewhat higher up on the banks, are

EA-PS: Hulten: Phytogeography of the Bering Sea District

confined to the inundation zone of the river. When the river changes its course
the tree in the drying-out branches [: ] of the river die. In the Populus woods
the undergrowth is very varied. It can consist of mosses, of Alnus fruticosa ,
Betula Middendorfii , Rosa acicularis , Spiraea salicifolia , or Vaccinium
uliginosum
. Between the shrubs there are open spots of abundant grasses,
chiefly Calamagrostis canadensis langsdorffii , with single herbs such as
Mulgedium sibiricum , Aconitum delphinifolium , Bromus richardsonii , Epilobium
angustifolium
, Festuca altaica , Galium boreale , Pyrola asarifolia incarnata ,
and Rubus arcticus. Salix macrolepis attains a height of 15 to 16 meters with
a trunk diameter of 40 to 50 cm. Populus suaveolens attains a height of 10
to 15 m. with a trunk up to 50 cm. in diameter. The birches, which occur
scattered, are 8 to 15 m. high. In the northernmost localities where this
fringing forest occurs, Salix macrolepis predominates. It is, thus, the tree
that penetrates farthest to the northeast in Asia. It is admittedly very
peculiar that this high-grown tree occurs far out in the otherwise treeless
tundra. The reason for this must be historical. It should probably be re–
garded as a relict from the Tertiary woods once occupying the area.
In the hills and mountains, which are especially well developed in the
Stanovoi Mountains, a subalpine zone with thickets of Pinus pumila , Alnus fruticosa ,
and Betula middendorffii occurs up to about 300 meters above the sea. The thickets
are 1/2 to 1 m. high, and Alnus growing on talus slopes alternate with Pinus
pumila
occurring on more stony slopes and open spots.
The undergrowth in the Alnus thickets consists chiefly of grasses and ferns.
In the Pinus pumila thickets it is very variable. Sometimes there is practically
no undergrowth; in other cases mosses, and in still others lichens. predominate.
Above the Alnus and Pinus pumila thickets there is an alpine lichen tundra.

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Penzhina-Gizhiga
The vegetation here has been but little investigated. It is similar to
that of the Anadyr region. In the upper valleys of the Penzhina and Olan rivers,
as well as west of the Taigonos Peninsula, woods of Larix dahurica occur which,
as in Anadyr, are found in the valleys of the Stanovoi Mountains.
East of the Stanovoi Mountains the large river valleys are fringed with
woods of Populus suaveolens , Salix macrolepis , and Betula cajanderi up to approxi–
mately 150 meters. Single specimens of Larix dahurica have been found 150 to 200
km. east of boundary line of the Larix woods, out in the open tundra. In
the lowlands wet meadows with flower mats and masses alternate with shallow bogs,
and drier heaths with lichens in the lowest layer, as well as with Betula exilis
societies with Sphagnum . In hills and mountains Pinus pumila thickets, which are
luxuriant and almost impenetrable in the southern part of the area, alternate
with Alnus thickets, in which single Betula cajanderi trees occur at low altitudes.
The Pinus pumila thickets have an undergrowth of Sphagnum or of lichens, chiefly
Cladonia at low altitudes, but mostly Alectoria above an elevation of 150 to 250
m. In the north. Pinus pumila thickets with Sphagnum girgensohnii undergrowth
are especially common. The thickets cover the mountain sides up to about 300 to
400 m., sometimes even up to 700 m. Above 400 or 500 meters lichen heaths pre–
dominate. The mountains along the west coast are completely unknown botanically.
Kamchatka
Somewhat south of 60°N. latitude, which is approximately the southern limit
of the permanently frozen soil, the Betula ermani forests of Kamchatka interrupt
the forestless area stretching southward from the shore of the Arctic Sea. However,

EA-PS: Hulten: Phytogeography of the Bering Sea District

this forestless area continues along the western coast of that peninsula to its
southernmost tip. Part of this forestless strip consists of former lagoons
filled out with Sphagnum bogs, but part of it is covered with a vegetation like
that of the southern arctic tundras. The Betula forest reaches an altitude of
about 700 meters in the interior, but much less near the coast.
This forest has an undergrowth of tall herbs, such as Filipendula kamchatica ,
Geranium erianthum , Epilobium angustifolium , Cirsium kamtchaticum , Cimicifuga
simplex
, lonicera chamissoi , doubtless remnants of the late Tertiary vegetation.
Along the large rivers there are alluvial meadows characterized by Thalictrum
kemense; Sanguisorba tenuifolia
and Majanthemum kamchaticum and bordering forests
of Salix sachalinensis and Populus suaveolens ; in places also Salix macrolepis .
The fringing forests of Salix sachalinensis have an undergrowth consisting chiefly
of 2 to 3-meter high Filipendula Kamchatica thickets. In the southwest [: ] the 3 to
5-meter high Angelica ursina covers the alluvial meadows and gives a peculiar, very
luxuriant look to the landscape. In other places groves of Betula “japonica”
occur, always standing on alluvial soil, while the Betula ermani forest begins
higher up on the slopes. At an altitude of about 300 to 400 m. the Betula woods
are suddenly replaced by Alnus thickets, which are 3 to 4 m. high but gradually
diminish in height father up on the slopes. They alternate, especially at high
altitudes, with open meadowlike spots. On stony ground they are replaced by
Pinus pumila thickets. At an altitude of about 900 to 1,000 m. the thickets are
very low grown and break up into patches. Higher up, alpine heaths with Rhododendron
chrysanthum
alternating with Vaccinium uliginosum heaths predominate.
In the continental parts of central Kamchatka an isolated Larix [: ] dehurica
forest about 250 km. long and at the broadest place about 100 km. broad occurs,

EA-PS: Hulten: Phytogeography of the Bering Sea District

alternating with groves of Picea jezoensis . Nowhere do these forests reach the
coast. A very small isolated grove on an endemic confiner, Abies gracilis , closely
related to A. sachalinensis , occurs on the middle part of the eastern coast.
Commander Islands and the Aleutian Islands
The Aleutians are treeless. The subalpine Alnus belt is also lacking, except
in the easternmost island, Unimak. The vegetation of the Aleutians is closely re–
lated to that of the alpine parts of Kamchatka Peninsula. In valleys and sheltered
placed meadows with Geranium erianthum , Anemone narcissiflora , Artemisia arctica ,
Erigeron peregrines , and Calamagrostis canadensis Langsdorffii occur. Shrubs
are rare, at least in the middle part of the chain, but in the eastern part, at
Unalaska, for instance, about 1 to 2-meter s high thickets of Salix Barclayi occur
at low altitudes. There, too, meadows with Thalictrum kemense , characteristic
of the Kamchatka Peninsula, occur. Above 80 to 100 m. a mosaic of alpine heaths
with abundant Empetrum , more or less mixed with Vaccinium uliginosum occurs,
alternating with meadowlike spots. On the easternmost island, Unimak, Alnus
thickets occur from sea level to a few tens of meters above it.
Alaska Peninsula
The vegetation of Alaska Peninsula is very little known. Grigg’s report
on the vegetation of the Katmai district seems to be the only phytogegraphical
survey of a limited area there.
The vegetation is much the same as in the easternmost Aleutians. In the low–
lands, meadows with a high growth of Calamagrostis canadensis Lengsdorffii alternate
with Empetrum heaths with Vaccinium uliginosum , Betula exilis , Ledum decumbens ,
and Vaccinium vitis-idaea.

EA-PS: Hulten: Phytogeography of the Bering Sea District

Sphagnum bogs with Rubus chamaemorus , Saxifraga hirculus and Eriophorum occur,
also thickets of willows. In the river valleys, at least in the eastern half of the
peninsula, there are fairly extensive forests of stunted Populus tacamahacca . Thus,
at Katmai, Populus forests about 50 square miles in area existed before the erup–
tion of the volcano, so that here as well as in Eastern Asia the poplar occurs
for outside the forest region.
The lower slopes of [: ] hills and mountains are covered with thickets of Alnus
crispa sinuata
with undergrowth of Calamagrostis and ferns. These thickets are
very similar to those occurring in Kamchatka, although much less luxuriant and
less confluent than there.
At the base of the peninsula, Betula kenaica also occurs in scattered groups.
There the tundra district borders on the coastal Picea sitchensis forest on the one
hand, and on the interior Picea glauca-birch forest on the other. Thus three very
different vegetational types meet at this spot.
Eastern Shore of the Bering Sea
No special study has been made of the vegetation of this district. The shore
is a low plain, covered at least in part with innumerable small lakes. Many of
them have developed into shallow bogs. Further inland the vegetation presumably
resembles that of the Aleutians and Alaska Peninsula. The coastal Alnus crispa
subsp. sinuata is replaced by Alnus crispa . To what extent it forms thickets there
is unknown. Probably it plays only a minor part in the vegetation. There, too,
Populus tacamahacca ( balsam poplar ) forms fringing forests along the rivers in
the eastern part, closest to the interior forest region. East of the tundra belt
bordering the Bering Sea the Picea glauca-Betula resinifera woods begin. Populus
[: ] tacamahacca and P. tremuloides occur along the streams. In the hills, thickets

EA-PS: Hulten: Phytogeography of the Bering Sea District

of Alnus crispa and, in the muskegs, black spruce, Picea mariana , with under–
growth of Salix bebbiana , Alnus crispa , and Ledum groenlandicum are of frequent
occurrence. In central Alaska larch occurs, and in the southwestern part
Betula kenaica .
These forests do not belong to the Bering Strait district but to the
boreal woods of North America. In the Norton Sound district they come close
to the shores of the Bering Sea.
Islands in the Bering Sea
Of the islands in the Bering Sea, Nunivak Island has a vegetation agreeing
with that of the nearly shore opposite. The Pribilof Islands have a flora simi–
lar to that of the Aleutians, although several arctic plants reach the Pribilofs
but not the Aleutians, such as:
Phippsia algida Eutrema edwardsii
Arctagrostis latifolia Saxifraga hieracifolia
Panunculus nivalis Eritrichium chamissonis
Panunculus pallasii Hierochloe pauciflora
Panunculus pygmaeus Gentiana glauca
Papaver macounii Gentiana tenella
Coryadalis pauciflora
On the other hand, many southern plants common on the Aleutians have their
northernmost outposts in the Bering Sea area on the Pribilof Islands, such as:

EA-PS: Hulten: Phytogeography of the Bering Sea District

Phleum alpinum Fritillaria camschatica
Deschampsia beringensis Cerastium aleuticum
Vahlodea atropurpurea paramushirensis Stellaria ruscifolia aleutica
Carex anthoxantea Ranunculus eschscholtzii
Carex hindsii Lupinus nootkatensis
Carex macrochaeta Geranium erianthum
Streptopus amplexifolius Veronica stelleri
St. Matthew Island and St. Lawrence Island have a more arctic flora. Arctic
plants reaching their southern limit in the Bering Sea area at St. Matthew Island
are:
Eriophorum callitrix Ranunculus
Carex misandra Draba alpina
Claytonia acutifolia Ligusticum mutellinoides
Claytonia tuburosa Lagotis glauca stelleri
Merckia physodes
In St. Lawrence Island a further number of arctic species lacking farther
south in the Bering Sea area are found, such as:
Carex rariflora Diapensia lapponica obovata
Salix arctica Primula borealis
Salix chamissonis Androsace ochotensis
Ranunculus glacialis chamissonis Artemisia glomerata
Pyrola grandiflora Senecio atropurpurea
Cassiope tetragona Saussurea viscida

EA-PS: Hulten: Phytogeography of Bering Sea District

High-grown shrubs are lacking. St. Lawrence Island seems, therefore, to
have a purely arctic vegetation connecting up with that of Chukotsk Peninsula
and seward Peninsula. Pinus pumila is entirely lacking. The island is con–
siderably poorer in species than either Chukotsk Peninsula or Seward Peninsula.
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Journ. Bot. U.R.S.S. 18:4 (1933), pp. 278-86.

EA-PS: Hulten: Phytogeography of Bering Sea District

Turner, L. M. Contrib. Net. Hist. Alaska. - S. Misc. Poc. 155, Congr. 1st
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Anadyr
Ovsiannikov, V. F. Poedzdka v dolinu r. Anadyr letom 1929 goda (A trip to the
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skoi okruzi (1890 gg.). (Plants collected by Dr. L. F. Hrynewecki in
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region). Journ. Soc. Bot. Russ. 15:4 (1930) pp. 305-11. ----. The reindeer range and reindeer pasture in the Anadyrland (In Russian with
English resume). Transact. Arct. Inst. LXII (1936) pp. 1-104, 2 maps. Tjulina, L. On the forest vegetation of Anadyr Land and its correlation with the
tundras. Transact. Arct. Inst. 11, Geobotanic 1936.

EA-PS: Hulten: Phytogeography of the Bering Sea District

Penzhina
Gorodkov, N. N. Geobotanicheskii i pochvennyi ocherk penzhiskogo raiona
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Penshina distr. of the Dalnivostakski rayon). Trudy Dalnevost.
filial Akad. Nauk. SSSR. 1 (1935), pp. 7-84. Pavlov. N. V. Materialy k flore Penzhinskogo raiona na Delnem Vastoke.
(Materials for the Flora of the Penzhina distr. In the Far East.),
pp. 601-06. Sochowa, V. B. Po tundra basseina Penzhinskoi gubi. Izv. Gas. Geograf. Ob–
shchestva 64 (1932), pp. 1-24.
Kamchatka
Hulton, E. Flora of Kamchatka and the adjacent Island 1-4. Kungl. Sv. Vet.
Akad. Handl. Ser. 3 vol. 5, 8 (1927-30), 1100 pp., 791km maps,
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Commander Islands
Fedschenko, B. Flore des Iles der Commandeur. Acad. Sci. Cracovie (1906), 128 pp. Kjellman, F. R. Om Kommandorskiöarnas Fanerogamflora. Nordenskjöld, A.E.
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(Über die Phanerogamenflora der Kommandorski-Inseln. Bot. Centralbl.
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Aleutian Islands
Eastwood, A. A. Collection of Plants from the Aleutian Islands. Leafl. West.
Bot. 5 (1947), pp.9-13. Hulten, E. Flora of the Aleutian Islands. Stockholm, 1937, 398 pp., 477 maps,
6 fig., 6 pl. Hutchison I. W. Stepping Stones from Alaska to Asia. London & Glasgow, 1937,
246 pp. Appendix: Hulten, E., List of plants.

EA-PS: Hulten: Phytogeography of the Bering Sea District

Porsild, A. E. Vascular plants collected on Kiska and Great Sitkin Islands…
Can.Field.Nat. 58 (1944), pp. 130-31. Tatewaki, M. Notes on plants of the western Aleutian Islands collected in 1929.
Transact. Sopporo Nat. Hist. Soc. 11 (1930) pp. 152-56; (1931) pp.200-09. ---- [: ] and Kobayashi, J. A contribution to the Flora of the Aleutian Islands.
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Alaska Peninsula
Griggs, R. F. The edge of forest in Alaska and the reasons for its position.
Ecology 15 (1934), pp. 80-96, 6 fig. ----. The vegetation of the Katmai district. - Ecology 17 (1936), p. 380-417, 13 fig.
Eastern Shore of Bering Sea
Collier, A. J. Notes on the vegetation. Brooks, A.H.; Richardson. G.B.; Collier,
A.J.; and Mendenhall, W.C. Reconnaisance in the Cape Nome and Norton
Bay regions Alaska in 1900. H.Doc. 547, 56th Congr., 2nd Sess. (1901),
pp.164-66. Eastwood, A. A. A descriptive list of plants collected by Dr. F. E. Blaisdell at
Nome City, Alaska. Bot. Gaz. 33 (1902), pp.126-194, 199-213, 284-99, 9
fig., 1 map. Flett, J. B. Notes on the Flora about Nome City. Plant word 4 (1901) pp.67-68. Kjellman, F. R. Fanerogamer från vest-eskimåernas land. Nordenskjöld, A.E.
Vega exped. Vetensk. Iakttag. 2 (1883), pp.25-60. Knowlton, F. H. List of Plants collected by C. L. McKay at Nushagak, Alaska in
1881 … Proc. U.S. Nat. Mus. 8 (1885), pp.213-221. Koidzumi, G. Plantae Siphogamae a N. Yokoyama anno 1907 in Alaska arctica,
Tschuktschore et Kamtschatka collectae. Bot.Mag. Tokyo 25 (1911), pp.
203-222. Muir, J. Some botanical notes from the Cruise of the Corwin. Torreya 18 (1918)
pp. 197-210. Porsild, A. E. Contributions to the Flora of Alaska. Rhodora 41 (1939), pp.
141-83, 199-254, 262-301, 4 pl., 1 map.

EA-PS: Hulten: Phytogeography of Bering Sea District

Seeman, B. C. The botany of the voyage of H.M.S. Herald … Flora of western
Eskimaux-land. London, 1852-57, pp.11-56.
Islands in the Bering Sea
Kjellman, F. R. Fanerogamfloran på St. Lawrence -ön. Nordenskjöld, A.E. Vega–
Expedit. Vetensk. Iakttag. 2 (1883) pp.1-23. Macoun, J. M. A list of the plants of the Pribilof Islands, Bering Sea …
Jordan, D.S. The Fur Seals and Fur Seal Islands of the North Pacific
Ocean. 3 (1899) pp. 559-87, 7 pl. Merriam, C. H. Plants of the Pribilof Islands, Bering Sea, with critical notes
by J. N. Rose. Proc. Biol. Soc. Wash. 7 (1892), pp.133-50. Porsild, A. E. Flora of Little Diomede Island is Bering Strait. Trans. Roy.Soc.
Canad., Ser.3, Sect.5, 32 (1938), pp.21-38.
Eric Hulten
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