Phytogeography and Vegetation of the Bering Sea District

Unpaginated      |      Vol_VI-0420                                                                                                                  
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

001      |      Vol_VI-0421                                                                                                                  
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.

002      |      Vol_VI-0422                                                                                                                  
EA-PS: Hulten: Phytogeography of Bering Sea District

        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

003      |      Vol_VI-0423                                                                                                                  
EA-PS: HulteN: Phytogeography of Bering Sea District

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

004      |      Vol_VI-0424                                                                                                                  
EA-PS: Hulten: Phytogeography of Bering Sea District

        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

005      |      Vol_VI-0425                                                                                                                  
EA-PS: Hulten: Phytogeography of Bering Sea District

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

006      |      Vol_VI-0426                                                                                                                  
EA-PS: Hulten: Phytogeography of Bering Sea District

        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

007      |      Vol_VI-0427                                                                                                                  
EA-PS: Hulten: Phytogeography of Bering Sea District

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:

008      |      Vol_VI-0428                                                                                                                  
EA-PS: Hulten: Phytogeography of Bering Sea District

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

009      |      Vol_VI-0429                                                                                                                  
EA-PS: Hulten: Phytogeography of Bering Sea District

        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

010      |      Vol_VI-0430                                                                                                                  
EA-PS: Hulten: Phytogeography of Bering Sea District

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

011      |      Vol_VI-0431                                                                                                                  
EA-PS: Hulten: Phytogeography of the Bering Sea District

       

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

012      |      Vol_VI-0432                                                                                                                  
EA-PS: Hulten: Phytogeography of the Bering Sea District

       

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.

013      |      Vol_VI-0433                                                                                                                  
EA-PS: Hulten: Phytogeography of Bering Sea District

        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.

014      |      Vol_VI-0434                                                                                                                  
EA-PS: Hulten: Phytogeography of the Bering Sea District

       

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 .

015      |      Vol_VI-0435                                                                                                                  
EA-PS: Hulten: Phytogeography of Bering Sea District

        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.

016      |      Vol_VI-0436                                                                                                                  
EA-PS: Hulten: Phytogeography of the Bering Sea District

       

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

017      |      Vol_VI-0437                                                                                                                  
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.

018      |      Vol_VI-0438                                                                                                                  
EA-PS: Hulten: Phytogeography of the Bering Sea District

       

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,

019      |      Vol_VI-0439                                                                                                                  
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,

020      |      Vol_VI-0440                                                                                                                  
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.

021      |      Vol_VI-0441                                                                                                                  
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

022      |      Vol_VI-0442                                                                                                                  
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:

023      |      Vol_VI-0443                                                                                                                  
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

024      |      Vol_VI-0444                                                                                                                  
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.

       

BIBLIOGRAPHY

       

General

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.

       

Anadyr

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

       

Penzhina

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.

       

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,

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