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    Phytogeography and Vegetation of the Bering Sea District

    Encyclopedia Arctica 6: Plant Sciences (Regional)

    Phytogeography and Vegetation of the Bering Sea District

    Unpaginated      |      Vol_VI-0420                                                                                                                  
    EA-Plant Sciences

    [Eric Hulten]




    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
    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

    001      |      Vol_VI-0421                                                                                                                  
    EA-Plant Sciences

    [Eric Hulten]



            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.



            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



    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

    003      |      Vol_VI-0423                                                                                                                  
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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


    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

    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–

    , 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:

    008      |      Vol_VI-0428                                                                                                                  
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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


            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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            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

    , but in its central part there are not very extensive forests of Larix

    and Picea jezoensis .

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            On the American side there is an interior forest, chiefly of Betula

    , 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.



            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

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    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

    . 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

    , 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

    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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            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.



            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,

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    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

    , 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

    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,

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    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.

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    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

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    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:

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    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


    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

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    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.





    Chamisso, A. and Schlechtendal, D. De Plantis in Expeditione speculatoria

    Romanzoffiana observatis. Linnaea 1-6 (1826-31). Fernow. B. E. The forest of Alaska. Forestly and Irrigation 8 (1902), pp.

    66-70; also in Harriman Alaska Expedition 2 (1901), pp. 235-56. Flora SSSR. 1-14 (1934-49). Harriman Alaska Expedition 1-14. New York 1901-14. Editor: C.H. Merriam. Graves, H.S. The Forests of Alaska. Amer. Forestry 22 (1916) pp. 24-37, 15 illus. Hulten, E. Flora of Alaska and Yukon 1-10. Lunds Univ. Arsskr. 37-47 (1941–

    1950) 1902, pp. 1280, maps (Kungl. Fysiogr. Sällsk. Handl. 52-61). Kellogg, R. S. The Forests of Alaska. U.S. Dept. Agricult. Forest Services Bull.

    81 (1910), 24 pp. 8 pl., 1 fig., 1 map. Ledebour, C. F. v. Flora Rossica 1-4. Stutgartiae 1842-1853. Rothrock, J. T. Sketch of the Flora of Alaska. - Ann. Rep. Smiths. Inst. 1867

    (1868), pp. 433-63. Sargent, C. S. The Forests of Alaska. Gard. & For. 10 (1897), pp. 379-80. Sochawa, V. B. K istorii flory juzhnoi chasti aziatskoi Beringii. Bot.Journ.

    SSSR. 18 (1933), pp. 278-86. Soczawa V. Zur Geschichte der Flora des südlichen Teiles von Asiatisch-Beringien.

    Journ. Bot. U.R.S.S. 18:4 (1933), pp. 278-86.

    025      |      Vol_VI-0445                                                                                                                  
    EA-PS: Hulten: Phytogeography of Bering Sea District

    Turner, L. M. Contrib. Net. Hist. Alaska. - S. Misc. Poc. 155, Congr. 1st

    Sess, 8 (1886) pp.1-226. Vassiljev, V. N. Rastitelnost severnoj chasti vulkanicheskogo koltsa tichogo

    okeana. Vegetation of the northern part of the volcanic ring

    of the Pacific. Izv. Vesecojuz. Geogr. Obshch LXXVI: 5, pp.223-40.


    Chukotsk Peninsula

    Kjellman, F. R. Asiatiska Beringsundskustens fanerogamflora. - Nordenskjöld,

    A.E. Vegaexp. Vetenskapliga arbeten II (1883) pp. 473-572, 5 pl.

    (In German: Die Phanerogamen-Flora an des Asiatischen küste der

    Bering-Strasse. - Nordenskjöld, A.E. Die Wissenschaftliche Ergebnisse

    d. Vega Expedition 1 (1883), pp. 294-379. Kurtz, F. Die Flora der Tachuktschen Halbinsel nach den Sammlungen der Gebrüder

    Krause. - Engler Bot. Jahrb. 19 (1895), pp. 432-493. Trautvetter, E. R. v. Flore terrae Tschuktschorum. Acta. Hort. Petrop. 6:1

    (1879), pp.1-40.



    Ovsiannikov, V. F. Poedzdka v dolinu r. Anadyr letom 1929 goda (A trip to the

    valley of the Anadyr river in summer of 1929). Zapisk Vladiv. Otdj.

    Russk. Geogr. Obshch. V (1930), pp. 41-119. Soczawa V. Von der Waldgrenze in aussersten Nordosten Asiens ( [ ?] in Russian)

    Priroda No.12 (1929). ----. Das Anadyrgebiet. Zeitschr. Ges. F. Erdkunde 1930, pp.241-263, 1 map. Sochawa, V. B. O pjatnistykh tundrakh Anadyrskogo kraja. (On the spotty tundra

    of Anadyr district). Trudy polar. Komm. 2 (1930). ----. Rastenija sobrannye doktorom L. F. Grinevetskim na territorii v. Anadyr–

    skoi okruzi (1890 gg.). (Plants collected by Dr. L. F. Hrynewecki in

    the Anadyr district). Zap. Vladivost. Otdejel Russk. Geogr. Obshch.

    V:2 (1930) pp. 175-77. ----. O nekotorykh rastenijakh Anadyrskogo kraja (On some plants of the Anadyr

    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.

    026      |      Vol_VI-0446                                                                                                                  
    EA-PS: Hulten: Phytogeography of the Bering Sea District



    Gorodkov, N. N. Geobotanicheskii i pochvennyi ocherk penzhiskogo raiona

    Dalnevostochnogo kraja (Review of Phytogeography and soil in the

    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.



    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,

    28 pl., 69 fig., 1 colored map. Komarov, V. L. Flora Peninsulae Kamtschatka 1-3 (1927-30).


    Commander Islands

    Fedschenko, B. Flore des Iles der Commandeur. Acad. Sci. Cracovie (1906), 128 pp. Kjellman, F. R. Om Kommandorskiöarnas Fanerogamflora. Nordenskjöld, A.E.

    Vega Exp. Vetensk. Iaktt. 4 (1885), pp. 281-309. Resume in German

    (Über die Phanerogamenflora der Kommandorski-Inseln. Bot. Centralbl.

    26 (1886), p. 31.


    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.

    027      |      Vol_VI-0447                                                                                                                  
    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.

    Journ. Fac. Agric. Hokkaido Imp. Univ. 36 (1934), 119 p. Turner, L. M. Contributions to the Natural History of Alaska. Senate, Misc.Doc.

    155, 49th Congr. 1st Sess. 8 (1886), List of plants by H. Mann, pp.61-85.


    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.

    028      |      Vol_VI-0448                                                                                                                  
    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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