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Algae: Encyclopedia Arctica 5: Plant Sciences (General)
Stefansson, Vilhjalmur, 1879-1962


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Corrections in Taylor’s Arctic Bibliography for Algae
#11 Boldt, 1888 - ibid refers to #12
#101 Larsen 1907 - add: pp. 305-364
#106 Lowe 1923- ibid is wrong
#114, 115, 116 - ibid should not cover volume number
#114 volume number is 29
1875 volume number is 32
1885 volume number is 42
#135 Rosenvinge 1898: pages are 129-243, 339-346.
#136 Rosenvinge 1917: volume number is 43
#30, 31 Author’s name is Cedercreutz, C.
H. Croosdale
for Taylor arctic
Ed. Plant Sciences

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EA-Plant Sciences (Wm. Randolph Taylor)

Significance of Arctic Algal Vegetation 1
Representative forms of Arctic Algae 2
Factors controlling algal vegetation 6
Marine Flora of the Arctic and Subarctic 9
Western Soviet Arctic 9
Eastern Soviet Arctic 11
North American Region 12
Baffin Bay and West Greenland 14
East Greenland 15
Iceland 16
Jan Mayen 17
The Faeroes 18
Northern Coast of Norway 19
West Soviet Arctic (Murman Sea) 19
Spitsbergen 20
Summary of Marine Algal Vegetation 22
Freshwater Algal Vegetation 23
Algal Floras of Various Regions 28
Summary of Freshwater Algal Vegetation 32
Bibliography 34

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EA-Plant Sciences
(Wm. Randolph Taylor)

In the Arctic, as in the Antarctic, the great rigors imposed by the
climate force us to consider most attentively the factors controlling life,
and the biological chain by which the more complex forms depend on the
simpler ones. The first source of organic matter is vegetable, and since
all animal life is dependent on it, the nature of the plant life in such a
territory as that which we here consider is very pertinent. It is clear
that the plant life of the exposed land becomes rapidly less as we pass
northward; the forests give place to tundra, beyond which there remains little
continuous vegetation. The sudden extremes of temperature and the destruct–
tive gales which may affect life on land are not so sharply felt under water.
In the sea great current drifts exist which carry warm waters far north, so
the conditions remain fairly suitable for aquatic organisms much farther to
the north than is the case with terrestrial organisms. The truly aquatic
vegetation in fresh water consists only in minor degree of vascular plants,
and in the sea still less.
When we examine the truly aquatic flora of cold northern regions we find
it rapidly diminishes, so that, for instance, Potamogetons reach a northern

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reach a northern limit in Scandinavia, except for a few species in Greenland,
etc., and Myriophyllums and Utricularias also reach only midway on the West
Greenland coast, while in the sea Zostera disappears a little north of the
southernmost portion. We find, however, a very considerable bulk and range
of animal life dependent on the great plant group the Algae, vast in variety
of species and vast in economic importance in all parts of the world, but in
cold regions often the sole ultimate source of organic matter.
Algae are plants of the greatest diversity, often beautiful in form and
color. Because of this wide range of pattern, it is hard to define them
except in technical terms. One may generalize by stating that they all have
the green pigment chlorophyll as their essential photosynthetic tool, just as
land plants do, though a few decadent types may similarly lose it and in many
it is concealed by accessory pigments. Then, the algae have no such system
of food-conducting ducts, or vascular bundles, as is familiar in land plants.
Finally, they lack the particular types of reproductive organs known as
archegonia and antheridia, which many nonvascular land plants possess.
There are several major groups or classes of algae, some ten or twelve,
depending on how the relationships are interpreted. This is far too many to
distinguish in a general account such as this, where we will have occasion to
refer to about six. If we depart from the standard nomenclatorial groupings,
we find that we can designate the types much more simply as freshwater or
marine, as bottom [: ] growng or as free floating (plankton) in the water. The
brackish-water types are few, especially in the Arctic, and the intergrades

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between plankton and bottom-growing types are limited to very small, even
microscopic organisms. In the sea the chief bottom types are often large and
are of the classes of chlorophyceae (green algae), Phaeophyceae (brown algae),
and Rhodophyceae (red algae), although there are some Bacillariophyceae (diatoms)
and Myxophyceae (Cyanophyceae or blue-green algae). The plankton types in
the northern seas are almost exclusively Bacillariophyceae and Dinophyceae
(peridinians), and are, therefore, quite microscopic. These are groups which
must be studied and described separately, so the reader is referred to the
articles “Phytoplankton” and “Algae: Planktonic Groups” in this Encyclopedia,
for they are the greatest food sources in the sea, and of prime importance.
While diatoms and peridinians are also important in freshwater plankton, other
algae of the green and blue-green groups are very important too, and almost
all forms are microscopic, though their abundance makes them a major food
source. The bottom vegetation in freshwater ponds consists chiefly of
organisms belonging to these groups of small species, though they are usually
different from the marine species. The bottom vegetation of the sea includes
numerous small forms as well as intermediate ones, and culminates in the great
rockweeds and kelps several feet in length, which are so conspicuous a feature
of northern shores. We will hardly concern ourselves with the microscopic
marine species, most of which are diatoms.
Blue-Green. Putting aside for the present the freshwater algae, we may
consider the appearance presented by the marine algae, which are more limited
in the Arctic than in warmer regions. First let us deal with the blue-green
algae, the least conspicuous group and the one to be expected highest on the
shore, in the splahs zone above high tides, between high and low tides, or

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rarely below low tide level. The commonest will be forms like Calothrix or
Rivularia , filamentous and, like the rest, microscopic as individuals, the
colonies of which comprise blackish or dark blue-green velvety or gelatinous
crusts of considerable extent on stones or woodwork. There will be a few
others which barely discolor the rocks, and in places Hyella , which lives
in and discolors the substance of mollusk shells. However, the marine blue–
green algal flora of the Arctic Sea is a sparse one.
Green . The green algae also are sparingly represented. They are plants
ordinarily requiring a maximum of light consistent with submergence, and so are
peculiarly susceptible to the long periods of darkness and the added ice and
snow cover. Since they prefer an intertidal habitat, they are subject to
the abrasive action of ice, and consequently green algae are, in general, scarce
in the Arctic, and inconspicuous wherever moving ice occurs. However, in some
districts the large club-shaped cells of Codiolum form a close, slippery,
dark-green coating on stones. Of the Ulvales we have two types, the tubular
Enteromorphas of which several wide-ranging species are present, and the
broad, flat blades of Ulva and Monostroma , the latter being relatively more
prominent than in warmer seas. The filamentous green algae are represented
by the microscopic Ulothrix , which accompanies Calothrix on the rocks, and
the large bushy types Chaetomorpha and Cladophora , reaching a decimeter or
more in height, which chiefly grow in protected shore pools and on littoral
or sublittoral rocks.
Brown . With the remaining two major marine algal groups we cannot concern
ourselves in such detail, because the variety present is too great. The brown
algae or Phaeophyceae are very conspicuous in the Arctic. They would ordinarily

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appear almost exclusively in the littoral and immediate sublittoral zones.
Probably because they cannot withstand ice action well enough, the Fucaceae
or rockweeds alone persist in this zone in any quantity, and chiefly toward
the south. The widespread Ascophyllum is one of these, and the others are
chiefly members of the genus Fucus , which is represented by several species.
These Fucaceae are richly branched, strap-shaped plants with inflations of
the blades serving as floats, swollen fruiting tips and, in Fucus , blades
marked by midribs and tufts of minute whitish hairs from minute pits in
the surface.
In contrast we have bushy filamentous genera such as Ectocarous and
Sphacelaria , and even some microscopic epiphytes (i.e., plants which grow
upon other plants, such as Myrionema . We have also, chiefly in the upper
sublittoral, a number of wide-ranging plant forms, such as crustose Ralfsia ,
narrow strap-shaped and foliaceous blades ( Ilea and Punctaria ), unbranched
tubes ( Scytoeiphon ), soft and branching submucus types ( Aegira , Mesogloia ),
or slenderly branched firmer bushy genera of larger growth ( Dictyosiphon and
Desmarestia ).
The final conspicuous group of brown algae are the kelps. These are quite
varied in aspect, and while occasionally they grow in the upper sublittoral,
they are more often in northern seas forced down into deeper water, where
their relatively long-lived blades and holdfasts may not be destroyed by the
ice. The genera include especially Laminaria , stalked blades which are plane
or ruffled, simple or cleft, and often two to several meters in length. Also
often present are Agarum , having a broader and shorter blade, rarely a meter
long, with midrib, the blade perfora c ted by a multitude of holes up to a

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centimeter in diameter, and Alaria , again a long entire blade, but with a
midrib and numerous fertile leaflets at the base.
Red . The red algae or Rhodophyceae are distinctly less conspicuous on
northern than on southern shores, and the plants are pretty well restricted
to the sublittoral. There are plants in many of the form categories described
for the brown algae. Of broad bladelike types there are not many; a few are
present and vary from the delicate brown or violet-red blades of Porphyra to
[: ] the larger, coarse, red, forking blades of Rhodymenia , a meter
or more long. Of the tubular types there are still fewer, but branched
Halosaccion and Dumontia are examples. Bushy filamentous species are more
numerous, the moderate-sized, exceedingly delicate, rosy tufts of Callitham–
nion contrasting with the large, coarse ones of Ceramium rubrum or the minute
ones of Rhodochorton , a few millimeters high. Finally, we have the crustose
species, not well represented in the brown and green algae. Hildenbrandia
forms a thin orange-red coat on stones in the littoral, while the opaque white
calcified crusts of various lithothamnioid types, smooth or raised into papillae
or bushy branches, form masses one or two decimeters in diameter which, while
they do not produce “nullipore” reefs as in the tropics, yet in less extreme
northern waters sometimes grow in conspicuous quantities at considerable depths.
Recognizing, then, that we are dealing with a very major food source
even when we have put the diatoms and peridines into other hands for detailed
discussion, we may turn first to the marine and then to the freshwater environ–
ments for discussions of their controlling conditions and plant populations.

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The only comprehensive account of these marine plants and their circumboreal
distribution yet attempted is The Algae of the Arctic Sea (79). Kjellman wrote
it with firsthand knowledge, since he accompanied Nordenskiöld to Spitsbergen,
to Novaya Zemlya, and to northern Siberia on the Vega expedition, during 1872-80.
In the following discussion, the term arctic algae will refer to types characteris–
tically found in the Arctic, but that does not mean that they are exlusively
arctic. The major marine flora of the Arctic is very similar around the whole
circuit. The species found in northern Asia and northern America are much the
same, so the questions to be answered concern, not continental floras, but the
controlling conditions which limit the plants in their abundance.
Light . The first and most striking of the controlling factors is light.
North of the Arctic Circle the summer season has continuous light but because
of its obliquity, the light is not continuously effective under water for
photosynthesis in the submerged marine algae. However, these plants are far
more adapted to use light of low intensity than land plants, and make a seasonal
growth quite comparable with that of similar species in temperate regions. In
fact, where they can grow without interference they often grow in the greatest
luxuriance. It is not surprising that they can manage with a long light period;
the adjustment to the long dark winter is a much more impressive adaptation.
Remembering that oblique illumination from low light near the horizon is not
very effective under water, we see that the marine algae when submerged live in
the dark for much of the year, or in a dim light of an intensity very much
below that which such shoal-water species prefer. Furthermore, penetration of
the light to the algae is by no means unimpeded. Far beyond the [: ] arctic

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night the presence of ice cuts down the light to minute proportions; for the
thick shore ice, fortified by overlying snow, is an effective block.
Temperature . Unfortunately, it is hard to be certain what temperature
prevails at the time and place where growth occurs. For plankton it is rela–
tively easy. Surface temperatures are known for the Subarctic and much of the
Arctic; they range from an average of about 7°C. at the surface in summer to
3°C. in winter near Nordkapp, but farther north the average temperatures drop,
especially the winter temperatures, which generally run well below 0°C.
Temperatures at various depths, too, are known for many localities, and we
know that the cold generally increases with the depth (though the range is
relatively slight), generally to about −1.5°, rarely to −3.0°C. or a little
more. Kjellman states that the temperature at the depth where the richest
algal vegetation occurs does not in general exceed 0°C. at any time of the year.
However, algae, especially nonplankton algae, are much better adapted
to growing at low temperatures than other plants, and in any case in summer
the actual growing temperatures for the shallow-water species (but not the
deep kelp beds) may be considerably above the surface reading for the open sea.
The fact that the winter temperatures, when these species are encased in ice,
may be considerably lower, seems less important, for many algae may be frozen
and survive a remarkable degree of cold, [: ] Even when the chief vegetative
parts are destroyed, regeneration may be possible from holdfasts, or spores,
and sporelings may persist. In the discussion of the algal flora of the
northern coasts, it will be found that temperature is not simply correlated
with latitudes, but that ocean currents alter conditions over large areas.

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Western Soviet Arctic
Novaya Zemlya . Having discussed the conditions under which the marine
algae grow, it is now necessary to select a starting point for a circuit of
the Arctic and an evaluation of the vegetation and controlling conditions in
each sector. The area between Baffin Bay, Spitsbergen, and Norway being by
far the richest and best known, and most continuous with a more temperate
flora to the south, it is decided to leave this to the last, and to start
the discussion with the conditions on the coast of the U.S.S.R. farther to the
east. Here we find considerable differences at once, for the flora of the
Barents Sea area resembles that of northern Norway and Spitsbergen, while that
to the east of Novaya Zemlya is somewhat different. On the whole, the northern
Soviet coast is relatively unproductive. The open character of the coast line
gives little protection, and the rock character is nearly everywhere unfavorable.
The numerous great rivers which discharge into the Arctic Sea along this line
tend to reduce the salinity of the surface water. This is especially marked
in the eastern part of the Kara Sea and the western Siberian Sea where the
salinity down to the level where the algae would grow is essentially less
than the proportion characteristic for the Arctic, or suitable for a general
flora of marine algae. These rivers also affect the temperature of the water,
as during a good part of the year the river is warmer than the sea, with the
result that the surface inshore water is not only less salty but decidedly
warmer than that offshore. The tides are not great, being extremely slight
in the eastern part, but somewhat greater about Novaya Zemlya, where they

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produce violent currents in the Mathochkin Shar Strait, which separates the
islands, with accompanying severe ice action.
Novaya Zemlya . Our knowledge of this district is very incomplete, and
we know details only regarding Novaya Zemlya and parts of the Kara Sea. [: ]
Kjellman (86; 79) and Sinova (147) have made the most significant contributions.
A sparse literature in Russian, to which some references have appeared in
Sinova’s work, is unfortunately unsummarized and not readily inaccessible.
The algal flora is poor in variety and very deficient over much of the coast,
being richest in the western and Novaya Zemlya areas. Sinova reports 123 species
of marine algae for Novaya Zemlya and nearby shores — far more than are re–
ported by Kjellman. The northeast coast is relatively inaccessible, and
glacier-bordered. However, the coast immediately northeast and northwest
of Matochkin Shar has proved unexpectedly rich. The littoral was [: ] nearly
bare, with some Urospora high on the rocks and Enteromorpha a little lower
down, but at the lowest littoral level some dwarfed Fuci, Chordaria and
Pylaiella appear. In the sublittoral appear the kelps: two Laminarias, three
Alarias, Phyllaria lorea (which was luxuriant at 77° N. latitude), a Polysiphonia,
two Delesserias, and a Phyllophora alone were common, but in some places
Lithothamnia were also frequent. Of this list, only Phyllaria lorea (if dis–
tinct from P. dermatodea ) is probably strictly an arctic plant, and there are
few strictly arctic among the less notable species. There are no prominent
species peculiar to these islands, or indeed to the whole Siberian mainland
The Kara Sea . In the western part of the Kara Sea, Euthora , elsewhere a
minor floristic element, becomes abundant. The chief peculiarity of the Siberian

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kelp flora east of Novaya Zemlya is the change in the Alarias, as Alaria
membranacea and A. pylaii are replaced by A. dolichorachis , A. elliptica ,
and A. oblonga . There are a few species lacking on the American side which
are present here, but they are represented in the northern Norwegian or
Spitsbergen floras. Along the general Siberian coast littoral algae appear
to be nearly absent, and kelps even in rather deep water are seldom met with.
Conditions are somewhat better than usual in the eastern parts of the Kara
Sea at Taimyr Island and east of Cape Bolshoi Baranov in the East Siberian Sea.
Eastern Soviet Arctic
With the approach to 170° W. longitude and Bering Strait, one is tempted
to look for a major change in the flora, for if most of the plants so far met
have been of North Atlantic affinities, here i w s the possible point of dis–
persal of a major North Pacific element. No such change occurs. The prominent
elements in the marine flora continue the same. Inspection of Kjellman’s list
shows no conjunction of northern Pacific, Siberian, and northwestern American
records except of very wide-rang i ng types, and the list of Collins (33) offers
only Chondrus affinis , Pterosiphonia bipinnata , and Rhodomela larix as northern
Pacific forms which occur in the American Arctic but not the Siberian. Of
course, the field work in this territory has been altogether inadequate,
but so far as the records go there is little to suggest a spread from the
Pacific into the Arctic or vice versa. A close study of the flora and vegeta–
tion on both sides of Bering Strait from the latitude of St. Lawrence Island
northward is sorely needed, to define sharply the limits of the floras. We
do know that the algal floras of the Okhotsk Sea (138), Kamchatka (151), and

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the Gulf of Alaska (139) are clearly subarctic, having their chief relation–
ships with the floras to the south, and little in common with the arctic flora.
North American Region
The conditions for algal growth on the American coast toward the west are,
like those of northern Eurasia, generally unfavorable. However, there are no
great rivers except the Mackenzie River (at 135° W. long.), so the salinity and
temperatures are relatively little affected from this source. The coast is in
general open and seldom of suitable rocks, so that the substratum is unfavor–
able. The additions made by Collins from the Canadian Arctic Expedition raise
the American records slightly, but this coast and that of Siberia still seem
to possess the poorest floras of the Arctic. As the stations investigated
poorly represent the whole, we note the need for detailed study, especially
by dredging, e.g., in Kotzebue Sound, east and west from Point Barrow, west
of the Mackenzie Delta, and east to both the north and south shores of Amund–
sen Gulf, where as stations investigated by Collins and. What may occur to
the north on the many great islands is practically unknown. It is not until
we pass Melville Peninsula and have the contrast of the subarctic Hudson Bay
to the south and Baffin Bay to the north, that substantial information regard–
ing the marine algae is again available.
Hudson Bay and Hudson Strait . Although from its latitude Hudson Bay cannot
be considered arctic, its connection with the Atlantic is through the arctic
algal zone, and its poverty of opportunity for the growth of algae resembles
much of the Arctic. The literature available is not extensive, Setchell and
Collins (143) in a short paper, Howe (72), and Polunin (124a) report on the

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Canadian Arctic Expedition materials. The first paper dealt with 28 algae in
material from Depot Island in the north and James Bay in the south; Howe had
material from James Bay and as far north as Richmond Gulf on the eastern side,
so that he was able to increase that total to 61 species and varieties — about
a fourth as many as occur in Baffin Bay to the northeast. There is little in–
formation about luxuriance, most of the material having been altogether frag–
mentary, but the writer, from this and other sources, gathers that the character
of the shore and bottom is generally soft and not suitable for good algal
development. One notes a scarcity of kelp, Chorda and a fragment of Alaria
alone being recorded, and only one Fucus . Most of the material was secured
by dredging, and some shallow-water types ( Sphacelaria , Pylaiella , Chordaria ,
etc.) occurred at the surprising depth of 18 meters. A few plants, hitherto
absent in our circuit, presage the appearance of North Atlantic and northern
Norwegian algae, such as Ralfsia deusta and Turnerella pennyi .
Labrador and Newfoundland . Finally, before turning to the North for
a consideration of the relatively rich flora there, we should note the scanty
information avaiable regarding the subarctic flora of Labrador and Newfoundland.
The Labrador Peninsula is represented by one paper (50) dealing with the Ungava
Bay algae of L. M. Turner and the Howgate Polar Expedition algae of L. Kumlien
from nearby. The records are for the most part of types common to the Arctic,
including about 30 species, among which kelps are chiefly represented by Alaria .
Newfoundland many years ago was studied by De la Pylaie, and his classical
observations (37; 36) remain the last catalogues, too old by far to be currently
useful. The flora is subarctic in general, but several typically arctic species,
such as Polysiphonia arctica , occur there; we badly need a modern study,
particularly of the northeastern portions.

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Baffin Bay and West Greenland
The Baffin Bay area represents one of the richer algal districts of the
Arctic. Northward from Hudson Strait on the American side, the shore of Baffin
Island is considerably dissected, and the dissected shore line of Greenland
extends even farther north, which all favors algal growth. The American side
has been very little studied, but there are a few reports for Baffin, Devon,
and Ellesmere Islands recently especially those of Rosenvinge (132), Lund (107),
and Polunin (124a). The south coast of Ellesmere Island frequently has a soft
bottom in the fjords and algal growth is poor, but in Smith Sound the bottom
is rocky and supports good vegetation; however, good littoral vegetation has
not been reported anywhere. Laminarias appear as far north as Flagler Fjord
(79° 04′
The Greenland side is quite well known (131; 135; 130; 132; 107). The
algal flora is an extension of the cold North Atlantic types, with many species
omitted and several more truly arctic ones introduced; the numbers of the
arctic and of the North Atlantic kinds are about equal, the subarctic element
being somewhat greater than either of the others. Considering the small amount
of coast involved, the last-mentioned is richer than might be expected. Hard
rocks help support the algal population, and there is open water on the West
Greenland coast for a considerable annual period quite far to the north, with
fairly high tides. In spite of the ice and tidal action, the dissected Green–
land shores favor the littoral development of Fuci, and these abound unusually
far to the north. Arctic Alarias and Laminarias occur as far north as Uper–
nivik (nearly 73″, and have been formed loose on Bjørling Island far
from the mainland (76°43′ At Foulke Fjord (78° 18′, there

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have been reported a littoral Enteromorpha zone and at the low tide line a
continuous belt of Fucus , with two species of Laminaria in deep water.
The flora of the Baffin Bay area is clearly related with that of the
American coast to the south, and with that of Iceland to the east. Currents
from Iceland sweep around the south coast of Greenland, then curve to the north,
before swinging toward Labrador and the south. In Baffin Bay and Davis Strait
the currents are therefore tending to flow south, and would not be expected
to bring American algae into the area; under conditions such as obtain at
present, forms from Iceland and the east might perhaps reach the boundaries
of the territory; but it is more likely that the similarity of the Baffin
Bay flora to the vegetation of the south is due to spread from the north than
vice versa.
East Greenland
East Greenland would seem much less hospitable to algae than West Green–
land, for the pack ice is driven against the coast by the current from the
Greenland Sea, deflected by the warm current tending northward from the North
Atlantic. There was little information available from the early arctic reports,
but we know considerably more now, and the flora as reported appears only
about one-eighth poorer in the east — for which the lesser proportion of brown
and green algae, forms growing in shallow water, may account. The coast is
now well reported upon in a preliminary way, for Amdrup’s Expedition (76) carried
the stations to 74°32′ N. latitude, while the Scoresby Sound Commission, 1932
expedition, and the Danmark expedition of 1906-08 (136; 133) carried them from
about 68° to 76°30′ N. latitude, about Danmarks Navn, but with notes from farther
north. At Lake Fjord in the south (66°21′, there is very good

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littoral development of fuci and kelps, but the fuci grow a few meters below
low tide line instead of in the littoral, and the kelps are still lower.
There is a good vegetation of Lithothamnia at considerable depths, and such
noncalcareous red algae as Turnerella , Phycodrys , Phvllophora , and Polysiphonia
arctica . there were algae in clumps frozen in the ice at Cap Am e é lie (77°32′, and this seems the limit of recorded living specimens on this side,
though there were fragments on the ice at the upper limit of observations,
Hyde Fjord (83°15′ There are about as many species at Danmarks Havn
as are known from Scoresby Sound (70° 21′, and they were the same
species, in general, although with a few notable omissions, such as Scaphosphora,
Chordaria , Dictyosiphon , Punctaria , and Chaetomorpha . The number of North
Atlantic species is far smaller in northeast Greenland, and the whole flora
is more markedly arctic because of this omission.
Iceland, lying close to eastern Greenland as it does, presents sharply
contrasting conditions for algal growth. The north coast is highly dissected,
but receives directly the cold Greenland Sea current bringing the pack ice
south in summer. The south coast is simpler, and receives the remnant of the
warmer current coming north from the mid-Atlantic, which tends to deflect the
ice westward toward Greenland. As this is a long-settled community with rela–
tively accessible coast, its vegetation is perhaps better known than that of
any other northern area, but the algae tend to be less known than the land
plants. Strőmfelt (163) did the pioneering work, Børgesen and J o ó nsson (24; 78)
a thorough study, reporting 200 species, with an elaborate analysis of the
phytogeographic relationships. Where the coast is rocky in Iceland, the rocks

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are chiefly fissured and eroded basalt; sometimes dolerite; sometimes on the
south, however, volcanic tuff. More than half the flora (124 species) is un–
distinctive, appearing in all suitable areas on the island. The small arctic
group of 11 species, including Turnerella , Polysiphonia arctica , and only
Laminaria nigripes to represent the great arctic kelp vegetation of Baffin Bay,
appear almost exclusively on the north and east coasts, where the cold currents
prevail and the tides are [: ] least, being only 0.76 to 2.3 meters in range.
Here the subarctic elements dominate, as they do in northeast Iceland also.
However, the southwest coast is more boreal, though with a considerable sub–
arctic element, the warm boreal being represented by such genera as Chondrus,
Bonnemaisonia , and Leathesia . This element is barely represented in northern
Iceland and is absent in the east. On the south coast, the greater, eastern
portion is sandy and not very productive of algae, but the western border is
better. The southwest coast is rocky and has the greatest [: ] tides, of 1.2
to 4.3 meters. In the southwest, the flora is much like that on the small
rocky area of the south coast. As a whole the flora resembles that of Finmark,
with a tendency for the eastern flora to resemble that of the White Sea in
its subarctic character, while that of the south and southwest resembles the
flora of the Faeroes.
Jan Mayen
The small arctic island of Jan Mayen, with a very inhospitable climate,
shows a comparatively well-developed marine flora, clearly arctic in dominant
characters, which we may piece together from the papers of Kjellman (88),
Rosenvinge (129) and others. None of the collections are adequate to define
the distribution of the vegetation on the coast. Fuci seem to grow [: ] at

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considerable depths (about 5 meters); kelps at 5 to 20 meters; Polysiphonia
and abundant Turnerella still deeper at 50 meters; and the deepest reported
vegetation of Phycodrys and Pantoneura at 110 to 118 meters. This seems to
be the usual shift in growth habit in the Arctic where there is considerable
risk of abrasion by floating ice. Kjellman considers the flora to be inter–
mediate between that of East Greenland and Spitsbergen, but peculiar in several
respects, and with a few species which he considered new and endemic. The variety
of kelps and the presence of Polysiphonia arctica , Turperella , etc., testify to
the arctic characters, as does the absence of the plants which Jonsson ascribes
to the boreal groups.
The Faeroes
The Faeroes resemble southwestern Iceland in flora, but with an even
stronger warm boreal element. There is still a very small arctic element, and
the strong subarctic fraction accounts for more than one-fourth of the species,
but the warm Atlantic current which sweeps around these islands places them in
a marine climate definitely milder than latitude would suggest, and gives them
a considerable number of warm boreal species which are absent from southern
Greenland. The sea temperature is reported to average 5.5°C. in the late
winter about Thorshavn, and about 10.3°C. in the late summer. The flora has
been thoroughly studied by Børgesen (25/ 27). The vegetation is rather luxuriant,
with a large intertidal growth of Fuci , Porphyra , Gigartina , and Rhodymenia ,
while Laminaria and Alaria may even be exposed at spring tides. The algal
growth here is sufficiently rich to promise direct utilization (28), as in
the Orkneys. The Shetland Islands, only a little farther south, lack the arctic
floral element altogether, and the subarctic elements becomes much less important
than the warm boreal element, so reversing the situation found in the Faeroes.

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Northern Coast of Norway
The coast of Norway, lying in the latitudes of south and central Green–
land, offers a marked floristic contrast to the latter. Receiving as it does
a good measure of the warm current from the North Atlantic, which swings around
into Barents Sea, we find that the southern portion has a very distinct boreal
character; but when Nordland Fylke is reached a quarter of the species are sub–
arctic, and in the northernmost Finmark Fylke the change is even more marked,
so that even Kjellman (79b) considers the whale area north of the Arctic Circle
to be in the Norwegian Polar Sea, and the adjoining coast of Finland and
western Russia to front floristically on the Murman Sea.
To the north of the Norwegian Polar Sea is the Spitsbergen Sea, which
floristically is much more extreme in its arctic character. The Norwegian flora
clearly has its affinity with the North Atlantic flora, but in the cold arctic
element may be accounted several species, especially in Monostroma , which are
not known elsewhere. A special aspect is given to the vegetation by the preva–
lence of Phyllaria dermatodea , which replaces in the upper sublittoral the
Laminaria and Alaria vegetation of Greenland, though these genera are not absent.
Several species occur in northern Norway which are not found on the more ice–
bound shores, and some, like Porphyra [: ] amplissima , reach unusual size.
There is abundance and variety of Fucus species and other rockweeds.
West Soviet Arctic (Murman Sea)
The Murman Sea shows the transition from the rather rich flora of the Nor–
wegian Polar Sea, with its prominent littoral flora of rockweeds and other types,
to the sparse flora of the Siberian coast which lacks significant littoral

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vegetation. When we compare the list of species we find that the chief charac–
teristic is a reduction in the variety, for by Kjellman’s listing, the Murman
Sea flora is much less than half as rich as that of northern Norway. There are
a few new types, perhaps a score, but these do not much reduce the loss. In the
western area and the White Sea, the Fuci are still prominent, though Himanthalia
and Halidrys are gone; but in the eastern area, Ascophyllum and Pelvctia have
left the flora, only two species of Fucus remaining instead of the eight or
nine which according to his 500 classification occur in Norway. Though still
in the flora, Rhodymenia has shifted to a sublittoral habitat. Certain other
forms become more prominent. Forming brown crusts on the rocks, we find Ralfsia
deusta widely distributed, while Pantoneura , Halosaccion , and Euthora become
common and luxuriant. The more truly arctic additions to the flora do not appear
in the White Sea and western area, but rather in the east, with the advent
or marked increase in the prominence of Haplospora, Phyllaria lorea, Laminaria
nigripes , L.fissilis , and Sarcophyllis , and the presence of d e structive ice
action which presages the shifting of the brown algal flora into deep water,
as its constitution shifts from a rockweed-dominated to a kelp-dominated list.
The knowledge we have of the flora of Spitsbergen rests mostly on the
important studies of Agardh (4; 2) and Kjellman (82; 84; 79), both most expert
phycologists. These islands lie across the line of 80° N. latitude, being
mostly below it, while Franz Josef Land lies mostly to the north of it, with
marine algal flora unrecorded. The shore line of Spitsbergen is much dissected,
the character of the rocks being uneven as regards suitability for algal growth;
in great part they are of schists and poor, the proximity to the arctic ice

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fields and the position well within the area of drifting pack ice assuring
conditions suitable only for a strongly arctic vegetation. The flora is not
a very poor one, though somewhat smaller than that of [: ] northern Norway;
but it is much more extensive than that developed on the Siberian coast. Certain
general features are apparent. First, it is rare to find many plants in the
littoral, although about 14 species have been so reported, all sporadicall ; y.
There are a few fairly characteristic groupings in deeper water. One, based
on Fucus evanescens and Rhodomela Lycopodioides , with Polsyiphonia arctica and
other species, is quite often met at a few meters depth. Another, found occa–
sionally at 9 to 27 meters depth, is based on Lithoderma , associated with
Phyllophora interrupta , Laminaria solidungula , etc. In fact, the L. solidungula
vegetation is common and luxuriant in Spitsbergen. Another feature is the
occasional presence and apparent continuing growth in detached masses on the
bottom, often in huge bulk, of normally attached species, such as Phyllophora
, Desmarestia aculeata , and Kallymenia rosea .
The relations of the Spitsbergen flora are clearly with that of West Green–
land and arctic Ame b rica, rather than with Siberia, those species occurring in
common with Siberia being generally forms of wide range. Currents bring to
the shores of south Spitsbergen large quantities of debris of southern, often
recognizably Norwegian origin, and algal masses such as Ascophyllum bearing
Polysiphonia lanosa are included, but no boreal element is evident in the flora.
The fact that the temperature is constantly below even that of the northermost
coast of Norway would clearly put such plants at a disadvantage.

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The opinion expressed by Kjellman that the arctic algal flora is in its
distinctive features endemic, evolved and dispersed within the arctic seas, seems
to have been accepted by most later workers. However, Simmons (146) considers
that, after the evolution of a Tertiary arctic algal flora, it was largely, or
perhaps entirely, driven southward by the progress of glaciation, but that
some representatives afterward moved back. It does not seem that any signifi–
cant interchange with the Pacific has occurred. The fact that several charac–
teristic species also occur well down the American coast suggests more strongly a
dispersal from the Arctic rather than introduction into it. However, it must
be realized that this view can apply only to the species, not the genera or
families to which they belong, which are often very wide-ranging, with the
time of their introduction into the Arctic or dispersal from it accordingly
remote. There are only a very few endemic genera, and some of these are of
doubtful worth.
Of distinctive ecological aspects, perhaps the intertidal rockweed
vegetation so rich in the Norwegian north as well as in deep water on more
northern coasts, the Lithoderma vegetation and the kelp vegetation which is
of very wide range in deep water, are the best examples. The fact that the
vegetation in deep water can develop well even when it is seasonally snow–
and ice-shielded, far beyond the winter-long night, from even the feeble nor–
thern daylight, and at temperatures seldom above zero centigrade, is the most
striking physiological adaptation. Economically there is little promise of a
direct utilization of algae in the Arctic proper, where the biological yield
must be harvested through the cycle of animals in the sea. However, in the

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Subarctic, where the littoral growth is heavy or the sublittoral accessible,
the recent resurgence of utilization of kelp for alginic acid and of red algae
for various gels may well encourage exploitation.
The records of northern peoples using seaweeds for food are few and unsatis–
factory. That people in the tropics use them is well known; so do people in
temperate regions, notably the Asiatics. There is nothing about the coarser
arctic species which would indicate that they could not safely be eaten, with the
minor exception, probably, of the Desmarestias. Where the elaborate preparation
accorded them by the Japanese is impossible they could simply be boiled with other
foods to furnish pleasing bulk and texture, and in some cases a modifying flavor.
Kelps, Porphyra , and Chondrus are groups which so suggest themselves. The nutri–
tive value is probably small, but this is true of many foods; other virtues must
be given due weight.
Kjellman (79) expresses his opinion of the marine algal flora in the Arctic
very aptly at one point in his discussion: “the most prominent features in the
general aspect of the arctic marine Flora are scarcity of individuals, monotony
and luxuriancy.”
The arctic freshwater algal vegetation offers a very different problem from
that of the marine algae. In the first place the expectation of different floras
in different sectors is much less, freshwater algae having, in general, very wide
ranges. Then we have the special problem of the similarity of arctic and alpine
floras. Finally, there is much less literature available, and much of it deals

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only with the areas of interest to Scandinavian scientists. While we have
many more species listed, less is told about even the most characteristic
individual species among them.
The arctic freshwater vegetation may be expected to be the same in all
general features, whether in Greenland, northwestern Europe, Russia, or Alaska.
There will be reports of endemic species, of course, but these do not appear
in any part of the world to play a distinctive role. Therefore, the important
factors are those which produce special environmental conditions and so cause
a characteristic type of flora, which may be expected in various parts of the
The freshwater algal genera found in arctic regions differ very little
from those found in milder zones. The difference lies not in the genera, or
even in the species, but in the way particular types dominate their habitats.
We have the same types of filamentous algae; Nostoc , Spirogvra , Zygnema ,
Cladophora , Rhizoclonium , of unicellular genera such as [: ] Gloeocapsa ,
Cosmarium , of diatoms, and of peridinians. However, in arctic areas among
desmids Cosmaria are disproportionately common, and many of the “arctic”
species of desmid lie in Cosmarium , whereas Euastra and some others are rela–
tively scarce. Indeed, it is only in the specialized groups of desmids that
much prospect exists of designating phytogeographic areas (Boldt 1887).
Algae may notably color the damp rocks and soil, because of the per–
sisting humidity. On the snow they produce the red, green, and brown condi–
tions so often noted in accounts of travel in mountains and arctic lands.
We know little of an exact quantitative nature regarding the productivity of
arctic lakes, but we know that in many subarctic areas the fish production

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of lakes and streams is very generous, and the algae of the subarctic regions
are largely responsible for this. Where the lakes are too shallow for produc–
tion of major fish, algae and submerged mosses and vascular plants (often not
true aquatic genera) will serve as the food source for the lesser fauna.
Ice and Snow Fields . Freshwater algal habitats may be divided into several
categories, for they are responsive to many factors, such as temperature, aera–
tion, pH, available calcium or nitrogen, and light. As these factors have
been little studied in the Arctic, we had best confine ourselves to a brief
mention of a few characteristic aspects. First, let us consider the ice and
snow flora. This has chiefly been studied in alpine and subarctic areas [: ]
[: ] (173; 90), but the observations may be extrapolated to the arctic
situations. Since ice and snow organisms cannot be studied well except in the
living state, and few explorers in the Arctic have taken such care as to do
this, our best knowledge of them comes from the snow fields of mountains parts
of Alaska, Switzerland, and Scandinavia. But comparable conditions in general
prevail in the Arctic, and the at least seasonal continuity of habitat has
tended to a general distribution of the algae concerned, producing a similarity
of ice and snow flora wherever any is known, although there are a number of
different species in the Antarctic.
The classical organism concerned in coloring snow is Chlamydomonas nivalis , which
most often causes red snow, and is well known in the Arctic. On the old snow
fields, resting cells of this plant growing near the surface are sufficiently
abundant to color it, even to a conspicuous pink. Obviously, it multiplies at
about zero centigrade. Other red-snow algae are found, as C. sanguinea and
Smithsoniomonas abbottii . Characteristic organisms produce green and yellow

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snow, as Scotiella . Organisms characteristically forming an ice flora rather
than one on snow are Ancyclonema and Mesotaenium , producing a purplish-brown
color, the former in particular being reported from Greenland. Numerous other
species are minor but normal constituents of the snow and ice flora, which is
by no means an inconsiderable source of organic matter, involving, as it may,
great areas. In the Arctic the long periods of darkness probably limit
growth more than the cold.
Wet Rocks and Ground . Secondly, we may consider the flora of wet rocks
and ground. These are sometimes provided with gelatinous sheaths and so become
resistan se t to drying, but the climate of much of the Subarctic and Arctic in
summer favors algal growth in such situations. The daytime surface temperatures
rise rapidly; the stones lose heat slowly in the evening. There is often
abundant water from springs, bogland, melting snow, etc. on stones, red colora–
tion may be due to Glosocapsa , blue-green to Phormidium , and grass-green to
Rhizoclonium. One of the notable features of subarctic and arctic puddles and
wet ground is the development of Nostoc , especially N. commune colonies, which
have appeared in collections from many countries, including many seen by the
writer from Arctic America.
Brooks and Streams . Thirdly, there is the flora of brooks and streams. In
great part the elements will be like those of alpine districts, and somewhat
different from those of temperate countries. Notable features are the production
of abundant brown gelatinous unpleasant-smelling streamers of Hydrurus , and of
flat green blades of Prasiola fluviatilis . We may even have Batrachospermum
tufts, e.g., in Greenland. [: ] Bordering the brooks and rivulets, Mougeotia ,
Zygnema , Vaucheria , and diatoms may form green or brown mats.

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Lakes . The floras of deep lakes and of shallow ones may differ markedly,
particularly in the more arctic stations, as the shallow ones tend to thaw more
rapidly and completely. The algal flora in these shallow ponds is often in–
conspicuous, and it may require a close inspection to see that the bottom is
completely covered with a living carpet. The margin, usually subject to some
variation in submergence, may support a dense dark-brown felt of Stigonema or
Scytonema , which may extend out over the bottom. However, here the vegetation
is often in the form of a nondescript sludge which, if carefully examined, will
prove to be largely of gelatinous unicellular algae, chiefly blue-greens. When
such ponds are continuously exposed to sunlight they become quite warm, favor–
ing such groups of algae.
The algae are entirely able to withstand freezing and to take advantage of
even a few hours of thawing for rapid growth (68); in fact, a thin ice cover
probably does not inhibit growth, though full encasement and darkening by winter
snow no doubt ends activity for a season. It is not known just what species can
withstand freezing in the vegetative state, and to what degree. One notes in
alpine areas a massive development of Zygnema aplanospores, but commonly complete
absence of zygospores, suggesting that they can well withstand freezing in the
vegetative state.
Though net hauls have occasionally been taken, true plankton studies of
arctic lakes are nearly lacking, and not enough is known upon which to generalize.
Børgesen and Ostenfeld ( 25 29b ) report on the plankton of one deep lake in the
relatively temperate Orkneys; it was dominated by the diatom Asterionella , with
some Dinobryon , and with the diatoms Tabellaria and Fragilaria , the desmids
Xanthidium and Staurastrum (four species), and Sphaerocystis , all frequent on

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occasion. There is nothing peculiar about this. The area is hardly even sub–
arctic, to be sure. Strøm reports (159) various subarctic, alpine, and sub–
alpine lakes with similar floras in which diatoms, Dinobryon , Sphaerocystis ,
and Anabaena were the important plant elements; earlier (160); in discussing
some larger arctic-alpine lakes, he reported from these a very meager plankton
Soil. The algae of the soil have been studied both in Iceland and Greenland.
Petersen extended his studies of surface-growing (but not submerged) algae (121)
to an analysis of the subsurface soil of virgin and cultivated areas (124). The
soil contains in part species more or less peculiar to it, in part species that
may appear in surface waters. The dampness of the soil was found to be a vital
ecological factor in Iceland, a much larger population developing in damp soil.
Exposure to light was also important, soil with a heavy moss or phanerogamic
cover being unfavorable. The degree of acidity of the soil and the persistence
of snow cover seemed relatively unimportant.
Swedish Lappmark . Swedish Lappmark would scarcely deserve consideration were
it not for the valuable study of Borge (14) on the northernmost part at 68°20' N.
latitude in the Torne Träsk area at about 345 meters elevation. Borge’s account
of 442 kinds of freshwater algae is exceptionally informative. In the snow
fields of the area, red snow is reported to be very common, as it is in the
Sarek Mountains (161). Hydrurus is reported in several places; desmids are ex–
ceedingly varied but with southern forms by no means scarce; Pediastrum braunii
is widespread, but although Stigonemas are reported several times, there is no

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indication that they are striking elements in the flora, as they are in high
mountains in Canada (165). The Norwegian reports (158; 157; 159) are chiefly
from alpine and southern areas, but make the Scandinavian Peninsula phycologi–
cally very well known.
Finland . Only in its northernmost part can Finland be considered subarctic.
[: ] Cedercreutz (30; 31) gives the best account from here. he reports an arctic–
alpine element of substantial extent, involving 16 Cosmarium species among
desmids, and Pediastrum braunii , and considers that this high proportion of
Cosmaria (43 per cent on the Fisher Peninsula) is strong evidence of arctic
The Soviet Arctic . The Soviet Arctic has not been well surveyed for fresh–
water algae, and, since even a reconnaissance would require far more detailed
collecting than for marine algae, the situation is very obscure. In the west–
central and southeastern parts of the Murmansk region, the reports given by
Kosinskaia (94; 95) deal especially with desmids; the flora is rather rich and
not particularly arctic, with several Micrasterias and Euastrum species. The
desmids alone have been studies at several islands and stations near the arctic
coast [: ] (92), from the Kara Sea to the Bering Sea. The whole character is
arctic, with no Micrasterias, few Closteria and Euastra, but very many Cosmaria
(50 species).
Jan Mayen, Novaya Zemlya, and Franz Josef Land . The arctic islands of Jan
Mayen, Novaya Zemlya, and Franz Josef Land have all been studied and sere reported
upon by the beginning of the twentieth century (98; 171 a ; 93), in some cases by
several expeditions, but little has appeared recently. Large lakes are not a
feature of these islands, and so the algal flora tends to be one of wet rocks,

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marshy ground, and snow fields, exhibiting rather little variety. The flora is
strongly arctic; for instance on Franz Josef Land, there are, in desmids, no
species of Micrasterias or Closterium, but many Cosmaria, and Prasiola, Gloeocapea,
and especially Nostoc populations are, as usual, common.
Spitsbergen . Spitsbergen has been well studies by Borge (16). The flora
is sharply arctic. Hydrurus and Prasiola fluviatilis are found in streams. On
snow fields, Chlamydomonas nivalis , Raphidonema , and Ancyclonema are reported,
producing characteristic colorations. Though scarce, Pediastrum braunii is re–
ported. Other, subaerial, Prasiolas are reported on rocks. Among desmids, Micra
sterias is absent, and there is a wealth of Cosmaria. The hot springs of [: ]
Spitsbergen, with temperatures from 20.0° to −28.3° C. in a latitude of 79°25′30″
N., are a special feature of the island which has been examined by Strøm (162).
They support quite a large algal flora, including a local from the Chara aspera,
a genus otherwise apparently unknown so far north.
The Faeroes . Of the Faeroes the best account is given by Børgesen (24). On
the whole he finds the relations of the freshwater algae to be strongest with
the west European flora, except in that area of the islands where the arctic–
alpine phanerogams appear. Here there is, in desmids, a strong Cosmarium contin–
gent of the type common in the North. In streams Hyudrurus is common, and, even in
a subalpine stream, Enteromorpha . Though not a feature of alpine flovas, in the
Faeroes, Cladophora forms colonies in the lakes. The Sphagnum bogs and Myriophyllum
communities were rich in algal species. Among notable aerial algae, we have an
important vegetation of Prasiola on rocks.
Iceland . Icelandic freshwater algae have been quite extensively studied. The
general flora, northern in character, needs little additional mention. The most

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distinctive features appear in the studies of the more or less exposed algal
colonies (120; 121). In spite of the prevailing damp, algae on shrubs, masonry,
woodwork were not evident. On turf walls Prasiola was common. Among the hillocks
of the myri, such blue-green algae as Stigonema and Scytonema were common. Meadows
near the sea that were occasionally flooded by salt water were commonly covered
with Vaucheria . Shady mountain clefts were dominated by desmids and diatoms.
About the margins of caves there was much Trentepohlia . Bird cliffs were not
unexpectedly dominated by Prasiola , which favors areas besprinkled by bird dung,
and species of blue-green algae. Hot springs as on Spitsbergen are a feature
of the country; blue-green algae were abundant in them.
Greenland . Greenland in its southern flora partakes much of the character
of Iceland and Scandinavia, and space cannot be spared to discuss it. The more
northern portion has been dealt with by Børgesen (23) and Petersen (122), the
first with samples from about Danmarks Havn [: ] (76°46′, and the
second from 81°15′ to 83°06′ N. latitude, which appears to be the northern
limit of land samples of freshwater algae to date. At 76° there was quite
a varied flora. The desmids as usual were distinctive, with [: ] Cosmarium
dominating (42 species), Staurastrum not far behind, Euastrum (4), and
Closterium (2); Micrasterias were absent. Common and often large Nostoc commune
is reported. At the more northern latitude unfortunately the samples were not
made from very suitable stations, and desmids were few, but the blue-green
algae and diatoms were very well represented. Even on the old ice of the fjord,
the diatoms were freshwater species. Among blue-green algae Nostoc commune
again appears, from the border of the inland ice; Gloeocapsa and Phormidium are
prominent, and, depositing lime on the bottom of a stream, even Schizothrix .

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The American Arctic . The American Arctic at the time of writing (1948 was
too little known for a report on the algal flora to be completely comprehensive.
To the southeast we know (166; 167) that the freshwater algal flora of Newfound–
land has quite a few arctic species, but the variety of Euastra and Micrasterias
found there makes it clear that the arctic floral characteristics do not pre–
dominate. Westward, only Lowe (106) reporting on Canadian Arctic Expedition
(1913-18) algae from the arctic coast of Alaska gives any substantial report.
As was to have been expected, the one species of Micrasterias reported was rare,
Euastrum uncommon, but Closterium was represented by several species and Cosmarium
[: ] dominated as usual (about 40 species), so that the indicator value of the
desmids is again shown. The Nostoc vegetation was prominent, as the writer has
found it represented also in samples from Chesterfield Inlet, Iguliguar Island,
Sarpic Island, and Southampton Island in Hudson Bay; Winter Island to the north,
Cape Wostenholme, Cape Dorset, and Lake Harbour in Hudson Strait; and Minto
Inlet on Victoria Island.
In summary of the freshwater algal distribution, one may say that as far
as land is known to go to the north, and is even briefly exposed by melting
of ice and snow in the summer, there are suitable conditions and a freshwater
algal flora. A very little more exposure, as at Danmarks Havn and Franz Josef
Land, gives a substantial variety. On snow and ice there is also a varied flora,
but very specialized and relatively small. This flora is pretty uniform about
the whole arctic circuit, but we do not know much of its distribution off the
land masses, on the sea ice and the snow which may lie on it. The terrestrial

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and aquatic algae of the land masses also show little distinctive variety
in the different geographical areas apart from that controlled by local climate.
The species concerned are seldom exclusively arctic in distribution, with some
notable exceptions in the desmids. Even here, the most distinctive feature is
the dominance of Cosmarium and considerable suppression of certain other genera.
Among subaerial green alage, Prasiola is important. Among blue-green algae
we not the abundance of Gloeocapsa and especially of Nostoc — not in variety
but of individuals. On the whole one can say that, in terms of reducing variety
and abundance, freshwater algae are among the slowest of plants to respond to
the repressions of the arctic climate.

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1. Agardh, J.C. “Alger insammlade pa Grönland of Dr Sv. Barggren och P. Oberg,
bestämda af Prof. J. G. Agardh. — Redogorelse för en expedition
till Grönland ar 1870 af A. E. Nordenskiöld, Bilaga II,” Svenska
Vetenskapsakad. Öfvers. Förh . vol.27, pp.1080-81, 1871.

2. ----. “Birdrag til kännedomen of Spetsbergens Alger, jemte Tilläg,”
Svenska Vetenskapsakad. Handl, vol.7, no.8, pp.1-49, 1868.

3. ----. “Bidrag till kännedomen of Grőnlands Laminarieer och Fucaceer,”
Svenska Vetenskapsakad. Handl . Vol.10, no.8, pp.1-31, 1872.

4. ----. Om Spetsbergens Alger . Lund, Akademisk Program, 1862.

5. Ashmead, S. “Alage. Enumeration of arctic plants collected by Dr. J. J. Hayes
in his exploration of Smith’s Sound between parallels 78th and 82nd
during the months of July, August and beginning of September 1861,”
Acad. Nat. Sci. Philad. Proc . vol.96, 1864.

6. ----. “Plants from Smith’s Sound. Alage,” Jones, T.R., Manual of the
Natural History, Geology, and Physics of Greenland , London, 1875,

7. Babington, C. “Lichens from Barrow and Davis Straits…alga,” Jones, T.R.
Manual of the Natural History, Geology, and Physics of Greenland . 1875.

8. Bachmann, H. “Beiträge zur Algenflora des Susswassers von Westgrönland,”
Nitt. Naturf, Ges. Luzern , vol.8, pp.1-181, 1921.

9. Bell, H.P. and MacFarlane, C. “Marine algae from Hudson Bay,” Contri. Canada .
Biol. & Fish. Vol.8, no.3, pp.65-68, 1933.

10. Berggren, S. “Alger fran Grönlands inlandsis,” Svenska Vetenskapsakad. Öfvers .
Förh . vol.28, no.2, pp.293-96, 1871.

11. Boldt, R. “Deamideer fran Grönland,” Ibid ., vol.13, no.3, 5, pp.1-48, 1888.

12. ----. “Grundrragen af Desmidieernas Utbredning i Norden,” Svenska Veten–
skapsakad. Bihang. Handl. Vol.13, no.3,6, pp.1-110, 1887.

13. ----. “Nagra sötvattens-alger fran Grönland,” Botaniska Notiser , 1893,
pp.156-8, 1893.

14. Borge, O. “Beiträge zur Algenflora von Schweden, 2. Die Algenflora um den
Torne-Trasksee in Schwedish-Lappland,” Botaniska Notiser , pp.1-110,

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15. ----. “Chlorophyllophyceer fran Norska Finmarken, “Svenska Vetenskap–
sakad. Bihang. Handl . Vol.17, no.3, 4, pp.1-15, 1892.

16. ----. “Die Sűsswasseralgenflora Spitsbergens,” Norske Videnskaps-Akad.
Mat.-Nat. Kl. Skrifter , vol.11, pp.1-39, 1911.

17. ----. “Sűsswasseralgen von Franz Josefs-Land, gesammelt von der Jackson–
Harmsworth’schen Expedition,” Svenska Vetenskapsakad. Öfvers .
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18. ----. “Susswasserchlorophyceen gesammelt von Dr. Osw. Kihlman in nörd–
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19. Børgesen, F. “Algues d’eau douce. (Sauf les Diatomees),” Ostenfeld–
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20. ----. “Conspectus algarum novarum aquae dulcis, quas in insulis Fearoen–
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21. ----. “En för Faerøerne ny Laminaria,” Botanisk Tidsskr . Vol.20, pp.403-5,

22. ----. “Ferskvandsalger fra østgrönland,” Medd. Grønland , vol.18, no.1,
pp.479-81, 1894.

23. ----. “Freshwater algae from the ‘Danmarks-Expedition’ to North-East
Greenland (N. of 76° Lat.),” Danmarks-Ekspedition til Grønlands
Nordøstkyat 1906-1908. vol.3, no.3, pp.71-90, 1910.

24. ----. “Freshwater algae of the Faeroes,” Bot. of the Faeroes , based upon
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25. ----. “The marine algae of the Faeroes,” Ibid ., vol.2,pp.339-532, 1902.

26. ----. “Nogle Ferskvandsalger fra Island,” Botanisk Tidsskr . Vol.22,
pp.131-8, 1899.

27. ----. Om Algevegetationen ved Faerøernes Kyster; en Plantegeografisk
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28. ----, “Om et Par Smaaerhverv før Faerøerne og Island,” Atlanten , vol.2,
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29. ----. “Om Faerøernes Algvegetation. Et Gensvar. 1, 2,” Botaniska Notiser,
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29a. Børgesen, F., and Jonsson, H. “the distribution of the marine algae of
the Arctic Sea and of the northernmost part of the Atlantic,”
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29b. Børgesen, F., and Ostenfeld, C.H. “Phytoplankton of lakes in the Faeroes,”
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29c. Boye, P. “Bidrag til Kundskaben om Algvegetation ved Norges Vestkyst,”
Bergens Mus. Arbog , 1894-1895, vol.16, pp.16-46, 1896.

29d. Brown, R. “Cryptogamic Plants from Baffins Bay,” Jones, T.R. Manual of
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29e. ----. “On the nature of the discoloration of the Arctic Seas,” Ibid .

29f. Cederkrentz, C. “Freshwater Algae from Labrador.” Soc. Fauna flor. Fenn.
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30. ----. “Sűsswasseralgen aus Petsamo,” Soc. Fauna Flor. Fenn. Memor. vol.5,
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31. ----. “----.” Ibid . vol.7, pp.237-48, 1932.

32. Collins, F.S. “Algae of the Neptune Expedition,” Canadian Arctic Expedition,
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33. ----. “Bering Strait and Arctic Ocean Algae,” Ibid ., vol.4, pp.2 [: ] 1B-16B, 1927.

34. Croall, A. “Florula Discoana. Marine Algae,” Jones T.R. Manual of the
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35. ----. “Marine algae,” Brown, A. The Florula Discoana. Contributions to
the phytogeography of Greenland within the parallels of 68° and
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36a 35a . Dall, W.H. “Arctic marine vegetation,” Nature , vol.12, p.166,1875.

36. De la Pylaie, A.J.M.B. Flore de l’Ile de Terre-Neuve et les Iles St. Pierre
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37. ----. “Quelques observations sur les productions d’ile de Terre-Neuve, et
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38. Dickie, G. “Algae and Diatomaceae,” Nares, G.S. Narrative of a Voyage to the
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39. ----. “Algae,” Hooker, J.B. “An account of the plants collected by
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40. ----. “XXIV (VI). Florula Discoana, freshwater algae,” pp.280-83;
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41. ----. “Notes of algae collected on the coast of northwest America, by
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42. ----. “Notes on a collection of algae procured in Cumberland Sound by
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43. ----. “Notes on a collection of plants from the northeast shore of Lan–
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44. ----. “Notes on flowering plants and algae collected during the voyage
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45. ----. “Notes on the algae,” Sutherland, P.C. Journal of a Voyage in
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46. ----. “On the algae found during the Arctic Expedition,” Linnean Soc.
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48. Farlow, W.G. “Alage,” p.214. In ; Wetherell, H.E. Botany. List of Plants
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49. ----. “List of algae collected at points in Cumberland Sound during the
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51. Foerov, B.C. “Sur les algues d’eau douce de Novaia Zemlia,” Wissenach.
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53. ----. “Contribution to the knowledge of the marine algae of Norway. I.
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54. ----. “----. II. Species from different tracts,” Ibid . vol.14,
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55. ----. “The Norwegian forms of Lithothamnion,” Norske Videnskaps-Akad. Skr .
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56. ----. “Om nogle nye arktiske havalger,” Norske Videns Keps-Akad. Forh .
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57. ----. “Remarks on Lithothamnion murmanicum ,” Ibid . 1908, vol.2, pp.1-8.

58. Gardner, G. “Liste annotee des especes de pteridophytes, de phanerogamies
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59. Gobi, C. “Die Algenflora des Weissen Meeres und der demselben zunächts–
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62. Gronlund, C. “Tillaeg til Dr. Kjellmans Afhandling,” Botanisk Tidsskr . Vol.11,
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64. ----. “Note sur les collections cryptogamiques, rapportees par La Manche.
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65. Harvey, W.H. “Alage. Flora of Western Eskimaux-Land,” Seeman, B.L. The
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66. ----. “List of arctic algae, chiefly complied from collections brought
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67. Hauck, F. “Algae” Reichardt, H.W. Flora der Insel Jan Mayen. Beobachtungs
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68. Häyren, E. “Bilder fran Finlands Ishavskust,” Geogr. Sallsk. Finland Terra .
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69. ----. “Notiz über das Überwinttern einiger Algen unter dem Eise,”
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70. ----. “Zwei notizen über das Meereseis und die Algen,” Societas Fauna
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71. Hooker, W.J. “Alagae,” Arnott, G.A.W. The Botany of Captain Beechey’s Voyage;
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72. Howe, M.A. “Hudson Bay algae,” Canadian Arctic Expedition 1913-1918. Report ,
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73. Hubbenet, E.R., and Voblikove, T.A. “Sutochniyi khod fotosinteza u vondoroslei
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74. Jones, T.R. Manual of the Natural History, Geology, and Physics of Greenland
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75. Jonsson, H. “A contribution to the Knowledge of the marine algae of Jan Mayen,”
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76. ----. “The marine algae of East Greenland,” Medd. Grønland vol.30, pp.1-7 [: ] 3 ,

77. ----. “The marine algae of Iceland, I-IV,” Botanisk Tidsskr . vol.24, no.2
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78. ----. “The marine algal vegetation of Iceland,” Botany of Iceland .
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79. Kjellman, F. R. “The algae of the Arctic Sea. A survey of the species,
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80. ----. “Bidrag till kännedomen of Kariska Hafvets Algvegetation,” Svenska
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81. ----. “Bidrag till kännedomen om Islands hafsalgflora,” Botanisk Tidsskr .
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82. ----. “Förberedande anmärkingar om algvegetationen i Mosselbay enligt
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83. ----. “Norra Ishafvets algflora,” Nordenskiöld, A.E. Vega-Expeditions
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84. ---. “Om Spetsbergens marina klorofyllförande Thallophyter 1, 2,”
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85. ----. “Redogörelse för Kariska hafvets växtoch djurvärld,” Nordenskiöld,
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86. ----. “Über die Algenvegetation des Murmanschen Meeres und der Westküste
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87. ----. “Über die Meeresalgenvegetation von Beeren Eiland,” Arkiv. för Bot .
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88. ----. “Zur Kenntnis der marinen Algenflora von Jan Mayen,” Ibid . vol,5,
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89. Kleen, E.A.G. “Om Nordlandens h ö gre hafsalger,” Svenska Vetenskapsakad. Öfvers .
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90. Kol, E. “The snow and ice algae of Alaska,” Smithson. Misc. Coll . Vol.101,
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91. Koldewey, K. “Algae,” The German Arctic Expedition of 1869-70, and narrative
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92. Kosinskaia, E.K. “Desmidievye vodorosli iz Arktiki,” Akad. Nauk. Bot.Inst.
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93. ----. “Kriticheskii spisok presnovodnykh vodoroslei sobrannykh V. P.
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94. ----. “Materiaux pour la connaissance de la flora algologique de la
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95. ----. “Sur la flore des Desmidiees du Lac Montsche,” Ibid . ser.2, no.3,
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96. Krieger, W. “Süsswasseralgen aus Spitzbergen,” Bericht Dtsch. Bot. Ges .
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97. Kuchuck, P. “Meeresalgen von Sermidlet und Kleinen Karajakfjord. Bot. Ergeb
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98. Lagerheim, G. “Beiträge zur Flora der Bären-Insel. 2: Vegetablisches Süss–
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99. ---. “Bidrag til kännedomen om snöfloran i Lulea Lappmark,” Botaniska
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100. ----. “Ein Beitrag zur scheeflora Spitzbergens,” Nouva Notarisia ,
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101. Larsen, E. “Ferskvandsalger fra Vest Gronland,” Medd. Grønland , vol.33, 1907.

102. ----. “The freshwater algae of east Greenland,” Medd. Grønland , vol.30,
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103. Lawson, G. “On the Laminariaceae of the Dominion of Canada and adjacent parts
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104. Lemoine, Mme.P. “Expedition Arctique Canadienne. Melobesiees (Calcareous
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105. ----. “Sur les caracteres generaux des genres de Melobesiees arctiques et
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106. Lowe, C.W. “Freshwater Algae and Freshwater Diatoms,” Ibid . vol.4 (Botany, A),
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107. Lund, S. “The Godthaab Expedition 1928. The Marine Algae,” Medd. Grønland ,
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108. Lyngbye, H. C Tentamen Hydrophytologiae Danicae, continens [: ] omnia hydrophyta
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109. Marr, J.W.S. “Plants collected during the British Arctic Expedition, 1925,”
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110. Merrifield, M.P. “Arctic marine vegetation,” Nature , vol.12, pp.55-58, 1875.

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114. ----. “Desmidiaceae ex insulis Spetsbergen-sibus at Beeren Eiland in ex–
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115. ----. “Desmidieae arctoae,” Ibid , no.6, pp.13-43, 1875.

116. ----. “Desmidieer sammlade af Sv. Berggren under Nordenskiöld’ska Ex–
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120. Petersen, J. “The aerial algae of Iceland,” Bot. of Iceland, vol.2, pp.327-447,

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122. ----. “Freshwater algae from the northern coast of Greenland collected
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126. Richardson, J. “Algae,” Franklin, John. Narrative of a Journey to the Shores
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127. Richter, P. “Süsswasseralgen aus dem Umanakdistrikt. Bot. Ergebn. Der …
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128. Rosenvinge, L.K. “Algues marines,” Ostenfeld-Hansen, C. “Contribution a la
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130. ----. “Deuxieme Memoire sur les algues marines du Grønland,” Medd. Grøn
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175. Zanon, D.V. “Diatomee della Baia del Re (Swalbard),” Mem. Pont. Acad. Sci.
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William Randolph Taylor
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