Phytogeography and Vegetation of the Bering Sea District: Encyclopedia Arctica 6: Plant Sciences (Regional)
Phytogeography and Vegetation of the Bering Sea District
EA-Plant Sciences [Eric Hulten]
PHYTOGEOGRAPHY AND VEGETATION OF THE BERING SEA DISTRICT
TABLE OF CONTENTS
| Page | |
| CHARACTER AND DISTRIBUTION OF THE FLORA | 1 |
| Circumpolar Plants | 2 |
| Arctic Montane Plants | 4 |
| American and Eurasiatic Plants | 6 |
| Endemic Plants | 11 |
| Chukotsk Peninsula | 11 |
| Anadyr | 11 |
| Aleutian Islands (including Pribilof Islands) | 11 |
| Bering Strait District | 12 |
| Shores and Islands of the Bering Sea | 12 |
| VEGETATION | 14 |
| Chukotsk Peninsula | 16 |
| Anadyr | 16 |
| Penzhina-Gizhiga | 18 |
| Kamchatka | 18 |
| Commander Islands and the Aleutian Islands | 20 |
| Alaska Peninsula | 20 |
| Eastern Shore of the Bering Sea | 21 |
| Islands in the Bering Sea | 22 |
| BIBLIOGRAPHY | 24 |
EA-Plant Sciences [Eric Hulten]
PHYTOGEOGRAPHY AND VEGETATION OF THE BERING SEA DISTRICT
There can be no doubt that the Bering Sea district occupies a singular
position phytogeographically in the arctic and boreal belts. The distribution
patterns converge toward the Bering Sea. This is certainly not an incidental
phenomenon but the result of the large migrations between Asia and America which
have swept over the region in past geological ages. The Bering Sea area was
to a large extent unglaciated during the Pleistocene glaciations and the climatic
changes there during the Quaternary seem to have been less severe than in most
other places in the arctic and boreal world. Ancient plant populations have
therefore remained more or less undisturbed, or at any rate less depauperated
there than in most places elsewhere. The result is that the vari
t
ation of species
is larger in the Bering sea area than in most other place in the same latitude.
CHARACTER AND DISTRIBUTION OF THE FLORA
The distributional groups that meet in the Bering Sea area are the following:
The arctic and boreal circumpolar groups.
The circumpolar arctic-montane group.
The boreal Eurasiatic and arctic montane Eurasiatic groups.
The arctic and boreal American groups.
The Eurasiatic-western American group.
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 |
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 |
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 |
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 |
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 |
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:
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 |
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 |
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 |
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 |
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.
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.
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
.
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.
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
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.
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,
EA-PS: Hulten: Phytogeography of the Bering Sea District
this forestless area continues along the western coast of that peninsula to its southernmost tip. Part of this forestless strip consists of former lagoons filled out with Sphagnum bogs, but part of it is covered with a vegetation like that of the southern arctic tundras. The Betula forest reaches an altitude of about 700 meters in the interior, but much less near the coast.
This forest has an undergrowth of tall herbs, such as
Filipendula kamchatica
,
Geranium erianthum
,
Epilobium angustifolium
,
Cirsium kamtchaticum
,
Cimicifuga
simplex
,
lonicera chamissoi
, doubtless remnants of the late Tertiary vegetation.
Along the large rivers there are alluvial meadows characterized by
Thalictrum
kemense; Sanguisorba tenuifolia
and
Majanthemum kamchaticum
and bordering forests
of
Salix sachalinensis
and
Populus suaveolens
; in places also
Salix macrolepis
.
The fringing forests of
Salix sachalinensis
have an undergrowth consisting chiefly
of 2 to 3-meter high
Filipendula Kamchatica
thickets. In the southwest
[:
]
the 3 to
5-meter high
Angelica ursina
covers the alluvial meadows and gives a peculiar, very
luxuriant look to the landscape. In other places groves of
Betula “japonica”
occur, always standing on alluvial soil, while the
Betula ermani
forest begins
higher up on the slopes. At an altitude of about 300 to 400 m. the
Betula
woods
are suddenly replaced by
Alnus
thickets, which are 3 to 4 m. high but gradually
diminish in height father up on the slopes. They alternate, especially at high
altitudes, with open meadowlike spots. On stony ground they are replaced by
Pinus pumila
thickets. At an altitude of about 900 to 1,000 m. the thickets are
very low grown and break up into patches. Higher up, alpine heaths with
Rhododendron
chrysanthum
alternating with
Vaccinium uliginosum
heaths predominate.
In the continental parts of central Kamchatka an isolated
Larix
[:
]
dehurica
forest about 250 km. long and at the broadest place about 100 km. broad occurs,
EA-PS: Hulten: Phytogeography of the Bering Sea District
alternating with groves of Picea jezoensis . Nowhere do these forests reach the coast. A very small isolated grove on an endemic confiner, Abies gracilis , closely related to A. sachalinensis , occurs on the middle part of the eastern coast.
Commander Islands and the Aleutian Islands
The Aleutians are treeless. The subalpine
Alnus
belt is also lacking, except
in the easternmost island, Unimak. The vegetation of the Aleutians is closely re–
lated to that of the alpine parts of Kamchatka Peninsula. In valleys and sheltered
placed meadows with
Geranium erianthum
,
Anemone narcissiflora
,
Artemisia arctica
,
Erigeron peregrines
, and
Calamagrostis canadensis Langsdorffii
occur. Shrubs
are rare, at least in the middle part of the chain, but in the eastern part, at
Unalaska, for instance, about 1 to 2-meter
s
high thickets of
Salix Barclayi
occur
at low altitudes. There, too, meadows with
Thalictrum kemense
, characteristic
of the Kamchatka Peninsula, occur. Above 80 to 100 m. a mosaic of alpine heaths
with abundant
Empetrum
, more or less mixed with
Vaccinium uliginosum
occurs,
alternating with meadowlike spots. On the easternmost island, Unimak,
Alnus
thickets occur from sea level to a few tens of meters above it.
Alaska Peninsula
The vegetation of Alaska Peninsula is very little known. Grigg’s report
on the vegetation of the Katmai district seems to be the only phytogegraphical
survey of a limited area there.
The vegetation is much the same as in the easternmost Aleutians. In the low–
lands, meadows with a high growth of
Calamagrostis canadensis Lengsdorffii
alternate
with
Empetrum
heaths with
Vaccinium uliginosum
,
Betula exilis
,
Ledum decumbens
,
and
Vaccinium vitis-idaea.
EA-PS: Hulten: Phytogeography of the Bering Sea District
Sphagnum
bogs with
Rubus chamaemorus
,
Saxifraga hirculus
and
Eriophorum
occur,
also thickets of willows. In the river valleys, at least in the eastern half of the
peninsula, there are fairly extensive forests of stunted
Populus tacamahacca
. Thus,
at Katmai,
Populus
forests about 50 square miles in area existed before the erup–
tion of the volcano, so that here as well as in Eastern Asia the poplar occurs
for outside the forest region.
The lower slopes of
[:
]
hills and mountains are covered with thickets of
Alnus
crispa sinuata
with undergrowth of
Calamagrostis
and ferns. These thickets are
very similar to those occurring in Kamchatka, although much less luxuriant and
less confluent than there.
At the base of the peninsula,
Betula kenaica
also occurs in scattered groups.
There the tundra district borders on the coastal
Picea sitchensis
forest on the one
hand, and on the interior
Picea glauca-birch
forest on the other. Thus three very
different vegetational types meet at this spot.
Eastern Shore of the Bering Sea
No special study has been made of the vegetation of this district. The shore
is a low plain, covered at least in part with innumerable small lakes. Many of
them have developed into shallow bogs. Further inland the vegetation presumably
resembles that of the Aleutians and Alaska Peninsula. The coastal
Alnus crispa
subsp.
sinuata
is replaced by
Alnus crispa
. To what extent it forms thickets there
is unknown. Probably it plays only a minor part in the vegetation. There, too,
Populus tacamahacca
(
balsam poplar
) forms fringing forests along the rivers in
the eastern part, closest to the interior forest region. East of the tundra belt
bordering the Bering Sea the
Picea glauca-Betula resinifera
woods begin.
Populus
[:
]
tacamahacca
and
P. tremuloides
occur along the streams. In the hills, thickets
EA-PS: Hulten: Phytogeography of the Bering Sea District
of Alnus crispa and, in the muskegs, black spruce, Picea mariana , with under– growth of Salix bebbiana , Alnus crispa , and Ledum groenlandicum are of frequent occurrence. In central Alaska larch occurs, and in the southwestern part Betula kenaica .
These forests do not belong to the Bering Strait district but to the
boreal woods of North America. In the Norton Sound district they come close
to the shores of the Bering Sea.
Islands in the Bering Sea
Of the islands in the Bering Sea, Nunivak Island has a vegetation agreeing
with that of the nearly shore opposite. The Pribilof Islands have a flora simi–
lar to that of the Aleutians, although several arctic plants reach the Pribilofs
but not the Aleutians, such as:
| Phippsia algida | Eutrema edwardsii |
| Arctagrostis latifolia | Saxifraga hieracifolia |
| Panunculus nivalis | Eritrichium chamissonis |
| Panunculus pallasii | Hierochloe pauciflora |
| Panunculus pygmaeus | Gentiana glauca |
| Papaver macounii | Gentiana tenella |
| Coryadalis pauciflora |
On the other hand, many southern plants common on the Aleutians have their
northernmost outposts in the Bering Sea area on the Pribilof Islands, such as:
EA-PS: Hulten: Phytogeography of the Bering Sea District
| Phleum alpinum | Fritillaria camschatica |
| Deschampsia beringensis | Cerastium aleuticum |
| Vahlodea atropurpurea paramushirensis | Stellaria ruscifolia aleutica |
| Carex anthoxantea | Ranunculus eschscholtzii |
| Carex hindsii | Lupinus nootkatensis |
| Carex macrochaeta | Geranium erianthum |
| Streptopus amplexifolius | Veronica stelleri |
St. Matthew Island and St. Lawrence Island have a more arctic flora. Arctic
plants reaching their southern limit in the Bering Sea area at St. Matthew Island
are:
| Eriophorum callitrix | Ranunculus |
| Carex misandra | Draba alpina |
| Claytonia acutifolia | Ligusticum mutellinoides |
| Claytonia tuburosa | Lagotis glauca stelleri |
| Merckia physodes |
In St. Lawrence Island a further number of arctic species lacking farther
south in the Bering Sea area are found, such as:
| Carex rariflora | Diapensia lapponica obovata |
| Salix arctica | Primula borealis |
| Salix chamissonis | Androsace ochotensis |
| Ranunculus glacialis chamissonis | Artemisia glomerata |
| Pyrola grandiflora | Senecio atropurpurea |
| Cassiope tetragona | Saussurea viscida |
EA-PS: Hulten: Phytogeography of Bering Sea District
High-grown shrubs are lacking. St. Lawrence Island seems, therefore, to
have a purely arctic vegetation connecting up with that of Chukotsk Peninsula
and seward Peninsula.
Pinus pumila
is entirely lacking. The island is con–
siderably poorer in species than either Chukotsk Peninsula or Seward Peninsula.
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[:
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----. Das Anadyrgebiet. Zeitschr. Ges. F. Erdkunde 1930, pp.241-263, 1 map.
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Penzhina
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Eric Hulten
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