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Economic Botany: Encyclopedia Arctica 6: Plant Sciences (Regional)
Stefansson, Vilhjalmur, 1879-1962

Economic Botany

Edible Plants of the Arctic

EA-Plant Sciences
(A. E. Porsild)

EDIBLE PLANTS OF THE ARCTIC

CONTENTS
Page
Introduction 1
The Use and Preparation of Arctic Food Plants 5
Some Common Edible Plants of the Arctic 11
Fruits and Berries 11
Potherbs 14
Roots and Root Tubers 21
Beverage Plants 22
Lichens 23
Mushrooms 25
Bibliography 26

EA-Plant Sciences
(A. E. Porsild)

EDIBLE PLANTS OF THE ARCTIC
INTRODUCTION
Plant life, everywhere in the Arctic, is too sparse, dwarfed, and
poorly developed to make any considerable contribution to the food supply
of man. Only a few arctic plants produce edible and nourishing roots or
stems, and only near the southern fringe of the Barren Grounds are there
some that in favorable seasons produce small, edible fruits. All plants,
however, no matter where they grow, and especially those that are green,
have some food value, and many are potential sources of vitamin, besides
containing variable amounts of fat, protein, sugar, or starch.
Primitive man in the Arctic, however, has probably always been carni–
vorous, securing his food by hunting, fishing, or in some instances from
domesticated animals, and only to a very small extent has he ever supple–
mented his food directly from the vegetable kingdom. Possibly he first
came to use plants as food by accident, as a last resort when other sources
of good failed him; perhaps he gradually developed a taste for some of the
plants he had experimented with in this manner; or, conceivably, he may
have observed that some carnivorous animals, as for example, the polar bear,
at certain times of the year “loosens it bowels by eating grass.”
Although he often starved when hunting and fishing failed, his food
habits, from a dietary point of view, must always have been highly satisfactory,

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for not until he began substitute white man’s food for his own did he
begin to suffer from nutritional deficienc i es.
Not so with white men living in the Arctic, for the narratives of most
early arctic expeditions are replete with the tragic accounts of ravages
caused by scurvy; and even in recent times are there numerous instances
when white men wintering in the Arctic have suffered, or even died, from
lack of vitamins.
Different food habits are the cause of this. By preference, all arctic
aborigines, whether by intuition or by experience, have always eaten the
internal organs of animals, that we now know have the highest vitamin contents,
whereas white men have generally declined those parts of the animals, preferring,
instead, the “choice” meaty cuts that make good roasts but contain little
vitamin.
The recent investigations by Rodahl (13) and others of the vitamin
content of arctic plants, have demonstrated, too, that just those arctic
plants that are eaten by preference by nearly all arctic tribes, have the
highest content of ascorbic acid as well as of thiamin, and that the methods
of preparation and of storing of vegetable foods used by these people are
perhaps the best possible for the preservation of the vitamins.
Although, in the aggregate, the amount of vegetable food used by arctic
aborigines has always been small, they have, nevertheless, made use of a large
number of different species. One factor limiting the amount they could use in
that most arctic plants are available only for a very short time each year,
and that primitive man in the Arctic has never learned to grow even those
species that readily respond to makeshift cultivation and fertilizing. He
has, however, even though his methods are often desultory, learned to gather

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and store plant food for winter use, and to improve the palatability of
some species by cooking, and even, if perhaps not at first intentionally,
by a crude form of fermentation.
Among the Eskimos — the most widely distributed race of arctic
aborigines — the dependence on vegetable food varies from group to group
according to tradition and according to what plants are available in the
area occupied by them. Thus, to the most northernly tribes the use of
vegetable food is purely incidental and largely limited to the partly fer–
mented and predigested content of the rumen of caribou and musk oxen,
whereas in the diet of the Eskimos of southwestern Greenland, Labrador,
and western and southwestern Alaska, vegetable food constitutes a regular,
if not very large, item. In northeastern Siberia, Kjellman (7), noted that
vegetable food formed an important part of the food used by the Chukchi.
“Although the flesh of reindeer, seal, walrus and bear, besides blood,
blubber, fish and other animal food forms the bulk of their diet, it cannot
be denied, and must not be overlooked, that not only the nomadic reindeer
Chukchi but also the hunting tibes living along the sea-coast, utilize and
have a definite taste for vegetable food. When available, vegetable food
constitutes a regular part of at least their principal meals, and is eaten
eagerly, and certain kinds even with avidity; furthermore, they consider these
foods important enough each year to gather supplies that will last them through
the long, grim winter.”
In the matter of providence, the Chukchis differ from the Eskimos, to
whom the large-scale gathering and storing of food is not a common or
universal practise. Kjellman related that the inhabitants of Pitlekaj and
the surrounding Chukchi villages, at the beginning of the winter of 1878,

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had accumulated stores of vegetables that were fully comparable to their
stores of meat and blubber. So large, says Kjellman, were some of these stores
that a reindeer Chukchi, whom he visited in March, still had on hand consider–
able quantities of vegetables tha g t had been gathered in the course of the
preceding summer and autumn. The collection of such large quantities of
vegetables would entail an amount of planning and perseverance which is,
indeed, unusual among arctic peoples.
W. Bogoras (3) who, first as a member of Sibiriakov’s party and later
of the Jesup North Pacific Expedition, spend many years among the Chukchi,
has confirmed Kjellman’s observations on the food habits of the Chukchi,
but found that: “On the whole, vegetable food is much more used by women
and children than by men.”
The rather extensive use which the Chukchi make of vegetable food does
not seem to be conditioned by local abundance of edible plants or by the lack
of animal food. In physiographic respect ts, as well as rehards its flora and
fauna, and Chukotsk Peninsula certainly is comparable to northwestern Alaska
where plant food plays a far less important role in the diet of the Eskimo.
Kjellman thinks that the reason is an historical one and that the habit has
been preserved from a time when the Chukchi lived farther south, in a climate
more productive of vegetable food.
It is of interest to note that, although native plants have never been
extensively used by whites living in the Arctic, and then mostly in emergen–
cies, those used have generally been other species, and, in the light of
our present knowledge of vitamins, of lesser value than the ones used by
aboriginal races. Thus there are numerous examples in the narratives of
arctic expeditions of the uses made of lichens — especially “rock tripe”

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or “tripe de roche” of the early Canadian voyageurs — besides mushrooms,
puffballs, and scurvy grass ( Cochlearea ), none of which is ever used by
aboriginal tribes. Likewise, berries such as the alpine cranberry or
cowberry ( Vaccinium Vitis-Idea ), bilberry or whortleberry ( Vaccinium uligino–
sum
), and to a lesser extent baked-apple ( Rubus Chamaemorus ) are perhaps
among the most frequently and most readily used vegetable foods of white men
living in the Arctic whereas these fruits are generally ignored by aboriginal
people who prefer the crowberry ( Empetrum ), which, in tura, is not favored
by whites.
Although a number of arctic plants are greatly favored by manure, and
for this reason often grow more abundantly, and attain larger size, near human
habitations, the cultivation of such plants, or in fact of any plant, is quite
unknown to all aboriginal tribes of the Arctic. This is perhaps not strange
considering that most of these people are nomadic, and that, among primitive
people, the gathering of roots and berries is the work of women and children;
but it is surprising, nevertheless, that the Chukchi, who make such extensive
use of plants, have not learned to take advantage of this fact.
The Use and Preparation of Arctic Food Plants
Generally speaking, no truly arctic plant is poisonous, nor are there
known to be poisonous mushrooms, roots, or berries anywhere beyond the limit
of trees, where, in fact, it is safe to eat any vegetable produce that appears
at all edible. In the northern forest, on the other hand, there are a few
plants that are definitely known to be poisonous. Those chiefly to be on
guard against are the roots of water hemlock of musquash root ( Cicuta spp.),
the fruits of red baneberry ( Actaea rubra ), the death-cup toadstool ( Amanita
phalloides ), and the almost equally poisonous fly amanita ( Amanita muscaria ).

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The latter, however, is in great demand among the Chukchi and other arctic
tribes of eastern Siberia who chew the dried fungus as a narcotic or intoxi–
cabt (2).
Although the amount of vegetable food used by arctic aboriginal races is
not great, it is drawn, nevertheless, from a great many different sources. At
first glance, it might even appear that when vegetable food is used, any plant,
or any part of a plant that happens to be available, and is not too unpalatable,
is used. Such a conclusion, however, would be entirely erroneous. Thus,
certain circumpolar species of plants are used by nearly all arctic tribes,
whereas other, and closely related ones, are not. Furthermore, an examination
of the long list of plants used by Eskimo and Chukchi, shows that the preference
for certain species is not altogether due to local abundance. Kjellman, for
example, found that one of the principal food plants of the Chukchi was a
willow, which was very common near the winter quarters of the Vega and supplied
the bulk of the vegetable food collected. Other, and equally palatable plants,
that to all intent and purpose were just as common and could have been collected
without effort and in equal quantity, were completely ignored.
In this connection it is of interest to note that Rodahl (13) found that
the ascorbic acid content of the leaves and buds of arc g t ic willows exceeds
that of all other arctic plants examined by him.
Some plants on the other hand, that were far less common and, on account
of their scarcity and small size, had relatively small food value, were
collected with an eagerness and perseverance that, in view of the general
indolence of these people, astounded Kjellman. One such plant is Polygonum
viviparum , which, according to the Chukchi, must be collected immediately
after the snow leaves the ground and before the first leaves appear. Only

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the rhizome, which is of the size of an unshelled peanut, is used; but to
find and collect it in early spring certainly is no easy task. Nevertheless,
even full-grown people, according to Kjellman, engaged in the collecting, and
with surprisingly good results.
Kjellman noted that whereas leaves of the willow were the most commonly
used, the flowering stems of the much less common, Rhodiola rosea and especially
the root tubers of the vetch, Hedysarium obscurum , and the rhizomes of Polygonum
viviparum, were considered choice and much favored delicacies. As regards
preparation, he found that only a few were consumed raw; the bulk were eaten
boiled in soup cooked with meat or blood, and often after first having been
made into a form of sauerkraut. Roots, leaves, and stems of plants collected
for winter use were tightly packed into sealskin bags, as a rule, each kind
by itself. In the process of storing, such plants underwent some sort of
fermentation. By their texture, smell, and taste Kjellman was aboe to recog–
nize several kinds of “sauerkraut.” One of them consisted entirely of the
small twigs and leaves of Salix Kolymensis ; a second was composed largely of
the leaves of Petasites frigidus , mixed with a variable quantity of leaves
of Saxifraga punctata , leafy twigs of Salix kolymensis , the flowering axes
of Senecio congestus , and leaves of Oxyris digyna, while, finally, a third
kind consisted entirely of the succulent green leaves and stems of the knotweed,
Polygonum alaskanum ( P. polymorphum ). Other forms of “sauerkraut,” were
prepared from the flowering stems of fernweed ( Pedicularis spp.) and from
the leafy stems of seabeach sandwort ( Arenaria peploides ).
Although several kinds of berries occurred and were known to the Chukchi
by name, Kjellman found that none was used to any great extent and that only
crowberries ( Empetrum ) were eaten occasionally.

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Among Eskimos the amount of plant food used varies from group to group
but to them it nowhere assumed the importance ascribed to it by Kjellman
for the Chukchi. Thus Weyer (19) estimated that in the diet of the Eskimos
of the Bering Sea region vegetable food constituted no more than 5 per cent;
among the Central Canadian Eskimos, Stefansson (17) and Jenness (6) noted
that it was scarcely used at all whereas as in Greenland the use of vegetable
food has always been un-important, except from a dietary point of view.
Probably, it any particular group, the greatest use of plant food is,
at present, made by those who make the least use of imported “white man’s”
food; those who have easy access to trading posts very soon give up the
practice of gathering native plant food and use increasing amounts of imported
plant food in the form of flour, sugar, vegetable fats, preserved fruits, jams,
and so on. Among the more sophisticated Eskimos and Indians it is not uncommon
to find an apologetic attitude, or even a certain amount of condescension,
toward the less “enlightened” and “backward” among their countrymen who still
maintain “native” customs and habits. This attitude was noted some years ago,
at a trading post on the lower Mackenzie River. While waiting for the arrival
of the mail plane the writer had noted, on the riverbank below the post, a
large patch of wild raspberries “loaded” with excellent and fully ripe fruit.
A group of native children were playing hide and seek among the raspberry
canes but did not appear to pick the fruit. When commenting on this to the
mother of one of the children he was told that “ she bought raspberry jam for
her children in the store, where there was lots.”
In the use and preparation of plant food, Eskimo practices differ only
slightly from those of the Chukchi, the chief differences being perhaps, that
more extensive use is made of berries, and that such roots, stems, and leaves of

EA-PS. Porsild: Edible Plants

plants as are used, are not infrequently stored mixed with blubber. Although
oil from the blubber may to some extent act as a preservative, some fermen–
tation undoubtedly takes place as with the “sauerkraut” prepared by the
Chukchi. Twenty-five years ago, the writer found that only a small number
of plants were used by the Eskimos of northwestern Alaska. Amond the more
important were the leaves of Saxifraga punctata , the leaves the flowering
axes of marsh fleabane ( Senecio congestus ) and coltsfoot ( Petasites frigidus ),
all of which were made into a form of sauerkraut mixed with blubber; the root
tubers of Eskimo potato ( Claytonia tuberosa ) and those of the vetch ( Hedysarum
alpinum ) were gathered in considerable quantity and used during the winter
cooked as a vegetable with meat.
Of the several kinds of “berries” used, cloudberry or baked-apple ( Rubus
Chamaemorus ) and crowberry ( Empetrum ) were most favored. Both were eaten
fresh or preserved frozen in sealskin bags, and, besides, were served as
“Eskimo ice-cream” — a dish prepared from a mixture of seal-oil and
masticated reindeer tallow, whipped or beaten to the consistency of whipped
cream, to which the berries were added.
In modern West Greenland only a few native plants are g regularly eaten
by the Greenlanders as seasonal delicacies but from a dietary point of view
they may, nevertheless, be of considerable importance. The more primitive
East Greenlanders, on the other hand, make considerable use of plant food.
In the southern parts of East and West Greenland the kvan Angelica
officinalis — is common along brooks, and in sheltered spots in the fjords
may grow 6 feet tall. The tender, young leaf-stalks and flowering stems
are considered a great delicacy and, when available, are eaten in great
quantities raw. Because the kvan does not grow near the open seacoast,

EA-PS. Porsild: Edible Plants

where most Greenland towns and villages are situated, and because this
vegetable is in such great demand, long journeys are regularly undertaken
to obtain it. The kvan is equally relished by the Denish residents who
generally eat it cooked and creamed.
Of equal importance is the crowberry ( Empetrum ) which is common eve y r y–
where in Greenland where it fruits abundantly at least to latitude 70° N.
In 1857, Rink established that more than 1,000 barrels of crowberries were
consumed annually in Southwest Greenland by 6,100 Eskimos, and that during
the autumn months, when the berries were ripe, they formed a regular part
of the Greenlander’s diet. The berries are either eaten fresh, when picked,
or served with fresh the uncooked seal blubber. They keep well when frozen
and in this state may be kept throughout the winter. In place where Empetrum
is common, the frozen berries may even be collected in winter when a special
scraper or scoop is used.
The flowering stems of roseroot ( Rhodiola rosea ) and the fernweeds,
Pedicularis hirsuta and P. lanata , find a limited use as potherbs. The
alpine cranberry or cowberry ( Vaccinium Vitis-Idaea ) is of local occurrence
in Greenland whereas the bilberry ( Vaccinium uliginosum ) has a distribution
similar to that of the crowberry. While not favored by the Greenlanders,
both are in great demand by Danish residents. Several species of seaweed
are eaten by Greenlanders and have recently been found an important source
of ascorbic acid.
The fermented and half-digested content of caribou rumen, and also that
of the musk ox is considered a delicacy by all Eskimos who hunt these animals.
This vegetable food is eaten raw or added to soup made from meat or blood.
It is frequently preserved frozen for winter use. According to Bogoras, the

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content of the reindeer rumen is eaten by the Chukchis and by other reindeer
nomads of northeastern Siberia. In Greenland, where ptarmigan are hunted
extensively for sale to Danish residents, the content of the crop is usually
eaten at once by the hunter.
In the light of Rodahl’s findings that the arctic willow and ground
birch, summer and winter, are rich sources of ascorbic acid, and that the
latter also is an important source of thiamin (14), the dietary value of
the content of both rumen and ptarmigan crop is probably high. Although
the bulk of the winter food of caribou and reindeer is lichen, the twigs
of willow and ground birch form a not inconsiderable addition, whereas these
plants, summer and winter, are the principal source of food for both musk
oxen and ptarmigan.
SOME COMMON EDIBLE PLANTS OF THE ARCTIC
Fruits and Berries
In late summer several kinds of small fruits may be found in abundance,
especially near the southern fringe of the Arctic. Without exception those
found north of the limit of trees are edible and wholesome. Several kinds
are not damaged, and many even improve in flavor, by frost. Some may be
collected under the snow, or when the snow disappears in spring. In order
of abundance and palatability the more important are as follows:
Black Crowberry or Curlewberry ( Empetrum nigrum ). Depressed and matted,
freely branching, evergreen shrub. Leaves linear, spreading, resembling
those of spruce of juniper. The flowers are inconspicuous and solitary in
the axils. The purplish-black and shiny fruits are very juicy and sweet
but contain a number of large and hard seeds.

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The crowberry is circumpolar and is found throughout the arctic regions,
in eastern North America south to mountains of the New England states, in the
West south to California. It prefers sandy, rocky, and acid soils and reaches
its best perfection in a rather moist climate. In some parts of the Arctic
the berries are gathered under the snow by the Eskimos who scoop them into a
large sieve made of sealskin through which the snow, leaves, and other
impurities are sifted.
The crowberry or curlewberry, although not as well-flavored as some others,
because of its abundance and hardiness is easily the most important fruit of
the arctic regions, and, with the cloudberry or baked-apple, is the only one
regularly eaten by the natives of the Arctic. The berries are eaten when
picked or stores frozen and eaten with seal blubber or oil. A hundred years
ago (Rink) 1857) reported that at least 1,000 barrels were picked annually
in Greenland and that a mildly alcoholic and most agreeable sparkling white
wine was produced by fermentation of the juice.
Cloudberry, Salmonberry or Baked-Apple (Rubus Chamaemorus ). Herbaceous,
low perennial from a creeping rootstock. Leaves round or kidney-shaped, five
to nine-lo [: b ] ed, stalked. Flowers solitary, terminal 1/2 to 1 inch broad,
and white. The immature fruits are first reddish, then amber, and when
fully ripe became pale yellow and very juicy. Sir John Richardson (11)
aptly described them: “Perhaps the most delicious of the arctic berries,
when in perfection, but cloys if eaten in quantity,” whereas Fernald and
Kinsey (4) praised them with less reserve, saying: “The ripe, fresh berries
of Baked-Apple eaten without sugar or cream are delicious, but with the
addition of these dressings are positively luscious.” The Eskimos, lacking
such refinements, serve them in a mixture of seal oil and chewed caribou

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tallow which has been eaten to the consistency of whipped cream. This culinary
treat in Alaska is known as “Eskimo ice cream.”
Bilberry or Whortleberry (Vaccinium uliginosum ). Low, branching, erect
or decumbent shrub with small oval, deciduous leaves. Flowers small urn–
shaped, pale pink, in the leaf axila. Berries, blue to black with a bloom,
ripen early in August.
The bilberry is common throughout all arctic countries and, in the southern
part of the Arctic usually produces an abundance of sweet, delicious berries.
It grows in acid soil in open places, and inhabits dry as well as moist places.
Although to the European palate of better flavor than the crowberry, the
bilberry is not much esteemed by Eskimos who believe it is liable to cause
dental decay.
Lingon, Mountain-Cranberry, or Cowberry (Vaccinium Vitis-Idaea ). Low,
creeping shrub with dark, leathery, and evergreen leaves. Flowers bell-shaped,
white or pink, in small nodding, terminal clusters. The shiny, dark red
berries ripen in August and September, but remain on the vines throughout
the winter, and the following spring, when the snow disappears, are sweeter
and even better than in the autumn.
The mountain cranberry is widely distributed throughout arctic countries,
occurring north at least to the arctic seacoast, but does not, as a rule,
produce berries far north of the tree line. O It prefers acid soil and
is found in moist as well as in dry rocky places. Its greatest perfection,
however, is reached in open birch or willow thickets where, in some years,
the vines are red with the fruit.
When gathered in the autumn the tart berries, if kept frozen, will keep
until next spring. They are considered better flavored than the southern

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true cranberry, and are excellent for jams and jellies. A very refreshing
drink may be made from the diluted sweetened juice.
Alpine and Red Bearberry ( Arctostaphylos alpina and A. rubra ). Low,
trailing shrubs with shreddy bark, and deciduous, obovate, and finely
serrated leaves. The flowers are small and appear in clusters toward the
end of the branches in early spring before the leaves unfold. In A. alpina
the berries are black and shiny; in A. rubra they are red, juicy but rather
watery and insipid. Although eaten greedily by bears and ptarmigan, the
berries are unattractive to most people, but, according to Fernald and Kinsey,
“in the absence of more attractive berries the fruit is apparently wholesome
and one soon acquires a taste for it.”
Red Bearberry or Kinnikinnick ( Arctostaphylos Uva-Ursi ). Trailing
evergreen shrub with small, bell-shaped pink flowers in nodding, terminal
clusters. The coral-red and somewhat mealy and dry berries are rather
tasteless when raw but quite palatable when cooked. The powdered dry leaves
are occasionally used by natives as a substitute for tobacco, or mixed with it.
Northern Red Current ( Ribes triste ). The northern red current occurs
throughout the wood parts of the Arctic but extends only a short distance
into the barren grounds. The berries that in flavor and appearance are
almost indistinguishable from cultivated red currents, are ripe in August
but last only a short time.
Potherbs
The leaves and flowering stems of a large number of arctic plants may
be used in soups, as potherbs, picked as sauerkraut, or in salads. Descrip–
tions, of some of the most useful and well known, together with brief notes
on their occurrences and uses, are given below.

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Woolly Fernweed or Lousewort ( Pedicularis lanata ). Perennial herb with
a well-developed taproot terminating in one or more dense rosettes of pinnately
lobed leaves that resemble the fronds of certain ferns; the leafy, 5- to 10-inch–
high flowering stem terminates in a dense, white-woolly spi [: ] k e of rose-pink
scented flowers. Toward maturity the stems elongate, and in winter often
protrude through the snow. The root, which is lemon yellow and sweet, like
young carrot, may be eaten raw or cooked; the flowering stem may be eaten
boiled as a potherb. In northern Greenland, Eskimo children pick the flowers
and suck the sweet nectar from the base of the long corolla tube.
The woolly fernweed is one of the earliest spring flowers on the arctic
tundra. It is circumpolar and of arctic range.
Arctic Fernweed ( Pedicularis arctica ). Similar, but with less woolly and
more open spikes of pale pink flowers. The root is pale yellow and more
spindly. Pedicularis arctica is a North American species which ranges from
northwestern Greenland to the north coast of Alaska.
Hairy Fernweed ( Pedicularis hirsuta ). Similar, with still paler flowers in
a shorter spike. Like the preceding species, arctic or high-arctic in range,
but limited to western Asia, Europe, Greenland, and eastern Canada.
Pedicularis sudetica . Similar, but almost glabrous, with dark-colored
leaves and stems. The flowers are dark red, in a dense spike which elongates
as the seeds mature.
According to Kjellman, the Chukchis prepare a sauerkraut from the flower–
ing stems and eat the boiled rootstocks in soup.
Mountain sorrel ( Oxyria digyna ). Low and glabrous, somewhat fleshy
perennial with erect, simple stems from a large, chaffy rootstock. Leaves
are mostly basal, kidney-shaped in outline with from 1- to 2-inch-wide blades

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on long, slender stalks. Flowers small, red or green, in a terminal plume–
like raceme.
The mountain sorrel is a circumpolar, arctic-alpine species, ranging
from the north tip of Greenland south to the limit of trees, and in high
mountains even south into California. It prefers somewhat shaded slopes
and ravines, where snow accumulated during the winter provides moisture
that lasts throughout the growing season. In such places the fresh green
leaves of the mountain sorrel may be found all summer. It responds wonder–
fully to manure and, in the rich soil under bird cliffs and near Eskimo
dwellings, forms luxuriant beds.
The succulent, juicy leaves and young stems are edible. When raw they
are somewhat acid, but most refreshing and thirst-quenching; when cooked
their flavor and appearance resembles spinach. In Greenland a very tasty
dish, not unlike stewed rhubarb, is prepared from the sweetened juice
thickened with a small amount of rice or potato flour.
The Eskimos of Greenland and Alaska eat the fresh leaves of the
mountain sorrel, mixed with seal blubbar.
Broad-leaved Willow Herb ( Epilobium latifolium ). Erect, glabrous,
simple or branching perennial herb from 6 to 18 inches high, with lanceolate,
dark green and somewhat glaucous, sessile and fleshy leaves. The flowers
are purple, very large and showy, and leafy recemes. The long and narrow
seed pods contain four rows of seeds bearing loung, silky, tufts of white
hairs at their summits.
The willow herb is circumpolar in range and is common and even abundant
throughout the Arctic on sandy or gravelly, well-watered soils such as are
found on gravel bars in rivers and on flood plains.

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The flowers — the largest in the Arctic. — may be eaten raw as a salad;
the fleshy leaves are edible when cooked and in taste resemble spinach. In
Greenland the fresh leaves and the flowers are occasionally eaten raw with
seal blubber.
Eskimo rhubarb ( Polygonum alaskanum , P. alpinum var. lapathifolium ).
Freely branching perennial herb from a stout, several-inches-thick fleshy
rootstock, bearing leafy stems from 3 to 6 feet high. The stems are reddish
with thickened, sheath-covered joints from which rise the 2- to 8-inch-long
lanceolate-attenuate leaves. The flowers are small and greenish, in large,
plumose axillary panicles.
Eskimo rhubarb is common in eastern Asia, Alaska, and Yukon, east to
the Mackenzie, and extends north slightly beyond the limit of trees. It
prefers moist, alluvial, or open soil such as is found along river banks
and on land slides in the permafrost area where it may form pure stands
several acres in extent.
The young, finger-thick, bright red and juicy stems appear soon after
the snow melts; in flavor they resemble rhubarb and may be used as stewed
“rhubarb” and as pie-filling. The sweetened juice makes a very refreshing
drink.
Kjellman (under P. polymorphum f. frigida ) reports that the Chukchis
cook the sliced rootstock with meat and prepare sauerkraut from the green stems.
Northern Sweet Coltsfoot ( Petasites frigidus ). Extensively creeping
perennial herb with a slender rootstock; the flowering stems which precede
the leaves and appear soon after the snow leaves the ground, are stout, fleshy,
and cobwebby, from 8 to 18 inches high, with scaly and much reduced leaves,
and terminate in open, racemose corymbs; the flowering heads are about 3/4 inch

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in diam e ter and are composed of creamy white flowers. The basal leaves are
triangular in outline, 2 to 3 inches long, coarsely dentate, green and
glabrous above, white-tomentose beneath, on long, slender petioles. Common
in wet tundra ranging from northern Europe through Asia, western Alaska,
and western Canada almost to Hudson Bay.
According to Kjellman, the northern coltsfoot is a favorite of the
Chukchis, who, from the mature leaves, prepare a special variety of sauerkraut.
Marsh Fleabane ( Senecio congestus ). Biennial with a hollow and wasily
compressed stout simple stem from 1 to 4 feet high, terminating in a dense
corymb of pale, yellow-flowered, woolly heads; leaves ascending, linear to
oblong-lanceolate, undulate, dentate, or more or less pinnatifid.
Circumpolar and common in swampy places on the arctic tundra or by the
edge of lagoons, but attains its best development on open soil such as land
slides in the permafrost area, and on manured soil near human habitations.
The young leaves and flowering stems may be eaten cooked as a potherb,
as a salad, or made into sauerkraut.
Cowslip or March Marigold ( Caltha palustris s. lat.). A marsh plant of
the buttercup family with yellow flowers and rather large, roundish or
kidney-shaped, somewhat fleshy and tender leaves which may be eaten raw as
a salad, or cooked.
There are several races of the circumpolar cowslip; one dwarf and
creeping race inhabits the high-arctic tundra whereas a taller and more robust
plant extends far south into the forested region.
Roseroot ( Rhodiola rosea and R. integrifolia ). Tufted, succulent
perennials, with a large, thick and fleshy rootstock with a fragrance reminis–
cent of roses. The stems are 6 to 12 inches high and bear numerous fleshy

EA-PS. Porsild: Edible Plants

greenish or pink, oblong, toothed leaves. The flowers are pale yellow or
pink, in a terminal cluster.
Two closely related species of roseroot are found in the Arctic. The
first is common in northern Europe, south Greenland and eastern North America;
the second is found in eastern Asia and western North America; both grow in
moist places on cliffs and by brooks, often near the sea; in manured soil below
bird cliffs or near human habitations they attain lush and profuse growth.
The succulent young stems and leaves may be eaten raw as salad, or cooked
as a potherb.
Kvan or Angelica ( Angelica officinalis ). A coarse and glabrous plant,
from 3 to 6 feet high with very large compound leaves on long, hollow, green
stalks; the leafy flowering stalks bear numerous round-topped umbels of small,
greenish-white, sweet-scented flowers.
An Old World species of alpine-boreal rather than arctic range which
from Scandinavia extends west to Greenland, where it is found north to
Disko Island. In North America is found the closely related purple angelica
( Angelica atropurpurea ) and seacoast angelica ( Coelopleurum lucidum ).
The tender young leaf stalks and the peeled young flowering stems are
eaten raw by Lapps and Greenlanders, who consider the kvan their choicest
vegetable delicacy and who undertake long and arduous journeys to obtain it.
To the European palate the raw kvan is rather strong-flavored but when cooked
and creamed as celery is of excellent flavor. Angelica atropurpurea and
Coelopleurum lucidum is still stronger flavored than the kvan and can be
eaten only when cooked.
Sea Purslane or Seabeach Sandwort ( Arenaria peploides ). A coarse,
somewhat fleshy perennial of the chickweed family which is common on sandy

EA-PS. Porsild: Edible Plants

beaches where it often forms dense carpets or large hummocks.
The succulent young stems and leaves may be pickled as sauerkraut or
eaten as a potherb.
Dandelion ( Taraxacum ). A number of species of dandelions are found in
the Arctic where, especially on moist ledges below bird cliffs or near human
habitations, they respond to manure bylush growth. The tender, young leaves,
especially when blanched, make an excellent salad, and throughout the summer
the leaves may be used as a potherb.
Saxifrage (Saxifraga punctata). Low, stemless, mostly glabrous perennial
from a creeping rootstock. The leaves are erect dark green or reddish with a
roundish or kidney-shaped blade on slender stalks; flowering stem 6 to 10
inches high terminating in a short raceme of white or yellowish flowers.
The leaves of this and similar species that are native to eastern Asia
and northwestern America are eaten raw with seal blubber or as sauerkraut by
the Chukchis and western Eskimos.
Willow ( Salix spp .). According to Kjellman, the leaves of the eastern
Siberian willow Salix kolymensis ( S. boganidensis ) which was very common
around Pitlekaj, furnished perhaps the largest amount of vegetable food
consumed by the Chukchis, who, from the young leaves and tender young shoots,
prepared a much-relished sauerkraut. Bogoras adds that the inner bark of
willow roots at one time was an important source of food to the Chikchis.
The leaves of several ractic willows, including those of the closely related
S. pulchra , no doubt would be equally palatable. Weyer (19) states that the
Eskimos of Alaska eat the young leaves of willow. According to Rodahl (13),
the buds and leaves of arctic willow are exceptionally rich in vitamin C.
Scurvy grass ( Cochlearia spp.). Low annual or biennial, diffuse,

EA-PS. Porsild: Edible Plants

branching, and somewhat fleshy, glabrous herb. The lower leaves are bright
green, roundish or kidney-shaped in outline, on short stalks. The flowe [: ] r s
are inconspicuous, white, in few-flowered racemes; the seed pods are globular,
containing a few large seeds.
Scurvy grass is circumpolar and is common along arctic seashores, but
is rarely found inland. On well-manured moist soil under bird cliffs and near
human dwellings itf becomes tall and lush.
The somewhat peppery-flavored leaves when eaten raw as a salad, or when
cooked, are considered a valuable antiscorbutic and as such are mentioned
in the narratives of numerous arctic expeditions; it is not eaten by either
Eskimos or Chukchis.
Roots and Root Tubers
Root tubers, because of their high content of starch and sugar, rank
high in food value but, owing to their small size, rarely can be obtained in
large quantity.
Licorice Root ( Hedysarum alpinum s. lat.). A [: ] nonclimbing perennial of
the pea family with branching, erect leafy stems, 1 to 2 feet high, with
[: ] axillary, long-peduncled racemes of showy but rather small deflexed
pinkish-purple flowers. The seed pods are linear, flat, 1 to 2 inches long,
formed of several roundish net-veined joints. The leaves are short-petioled,
odd-pinnate, with 11 to 21 oblong or oblanceolate leaflets. The half-inch-thick
root tubers are sweet and taste somewhat like young carrots; they mature in
August but may be gathered until the ground freezes. In spring, before the
new growth starts, they are even better than in the autumn, but in summer
become tough and woody. The root tubers during spring and early summer form
the principal food of brown and black bears; several kinds of meadow mice and

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lemmings in autumn harvest and store the tubers for winter use. In order
to obtain a supply of this much-favored vegetable, the Eskimos of Alaska
rob the mice caches, which they locate by means of a dog specially trained
for this purpose. Bogoras reports that the method is practiced also by the
Chukchis.
The species, which includes several geographical races, is circumpolar,
and from the arctic tundra ranges south far beyond the tree line; it is common
in sandy places along river banks and lakes, where it often forms large clumps.
Eskimo Potato ( Claytonia tuberose ). The roundish tubers of this Asiatic
spring beauty, found in eastern Siberia and northern Alaska, is very palatable
and nutritious when boiled. Kjellman states that along the north coast of the
Chukchi Peninsula this is one of the best known and most used vegetable foods
and that even in late spring as much as a barrelful of the tubers might be
found in the storehouses of the more provident Ch i u kchis. In 1926 Porsild
found the Eskimo potato was popular also with the Eskimos of Diomede Island
and northwestern Alaska (9).
Alpine Bistort ( Polygonum viviparum and P. Bistorta spp. plumosum ).
Low perennials with a short and thick tuberlike rootstock and willow-like green
shiny leaves. The small white or pink flowers appear in a terminal rather
showy spike. Below the flowers P. viviparum , bear numerous small bulbs
that take root when detached. The tubers, although slightly astringent,
are rich in starch and have a sweet, nutty flavor.
Beverage Plants
Shrubby cinquefoil ( potentilla fruticosa ). A low, much-branched shrub
with shreddy bark, large yellow flowers, and numerous rather small leaves,
each composed of from 5 to 7 silky pubescent leaflets. The shrubby cinquefoil

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is common throughout the subarctic, in muskegs as well as in rocky places
but is not found far byond the limit of trees. The dried leaves may be used
as a substitute for tea.
Labrador tea ( Ledum decumbens and L. groenlandicum ). Low, branched,
strongly aromatic shrubs with evergreen, leathery, canoe-shaped leaves
covered beneath by a dense, rust-colored felt; the flowers are white, strongly
aromatic and spicy, in umbrella-shaped terminal clusters. One or another
of the several closely related species occur throughout the Arctic in muskegs
or wet tundra. The leaves may be gathered throughout the year and, after
drying, may be used as a substitute for tea.
Spruce tea . An infusion made by steeping in boiling water the young twigs
and leaves of spruce, hemlock, balsam fir, or pine has long been known to be
of value as an antiscorbutic. “Spruce tea,” especially if made from the young
leaves of balsam fir, is a rather agreeable drink when served hot with sugar.
Lichens
About the last source of food we should ordinarily think of are the dry,
juiceless, gray, drab, or brown lichens, often mistakenly called “mosses,”
which carpet sterile ground or expand their flat or crisped surfaces on rocks,
fences, or trees (4). Nevertheless, among the various edible plants occurring
in the North, the greatest potential food value should, perhaps, be assigned
to these uninspiring plants, because they occur so abundantly that 4 to 5 tons
may sometimes be harvested from one acre.
Lichens are low, variously s haped gray, brown, or black plants that, in
many parts of the Arctic (and elsewhere), are important components of the
vegetation. Botanically they are dual organisms consisting of fungi that
are parasitic on, and receive their nourishment from, primitive green or

EA-PS. Porsild: Edible Plants

blue-green algae that, themselves, are completely enveloped by, or diffused
through, the hyphae of the fungus.
Many different kinds of lichens are found in the Arctic and while none
is poisonous, only a few are palatable to man. Most lichens contain an acid
substance that may cause nausea or severe internal irritation unless removed
or neutralized by parboiling in water to which has been added a small amount
of baking soda.
Among the most easily recognized edible lichens are certain rock lichens
of the genera Gyrophora and Umbilicaria — commonly known as “rock tripe” or
“tripe de roche” — and a few species of Cladonia and Cetraria, often mistakenly
referred to as “moss” or “reindeer moss.”
The former, as the name implies, grow on rock or boulders to which their
irregularly shaped, saucerlike, leathery, brown, green, or black fronds are
attached by the center. When dry they are hard and brittle, but in damp
weather become soft and cartilaginous and in this condition are easily
detached from the rocks. The “mossy” kind grown on the ground, often among
other plants, or sometimes form dense and almost pure carpets. The most
important of these are the Iceland moss, ( Cetraria islandica ), said to
contain 80 per cent starch, besides some protein and fat, and “reindeer moss”
( Cladonia rangiferina , Cl. Sylvatica , and Cl. alpestris ). These are low,
bushy, coral-like lichens. The first is dark brown, its fronds straplike,
ciliate on the edge, while the fronds of “reindeer moss” are more coral-like,
composed of round, hollow, gray or greenish-gray branches. These lichens,
too, are brittle when dry and are best collected when moist.
After barboiling with soda, the lichens are oven-dried until brittle
and then powdered, which may be done by rubbing between the palms of the

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of the hands or by pounding, or better still by a grist mill.
The powdered lichen, if put to macerate in water overnight, jellies
when boiled with water or milk. One pound of powdered Iceland moss is said
to produce four quarts of jelly similar to blanc mange and is considered
very nutritious and digestible. In Iceland and in northern Scandinavia,
Iceland moss is used in puddings and in soup; formerly, in times of scarcity,
flour prepared from this and other lichens was added to the bread flour.
The moistened lichen flour will not form a dough unless mixed with a small
quantity of wheat flour. In this manner very tasty biscuits may be prepared.
The starch contained in the lichen may be fermented, and in Scandinavia
it formerly found a limited use in the manufacture of alcohol.
Mushrooms
Many different kinds of edible mushrooms and puffballs occur throughout
the Arctic, especially near the southern fringe of the tundra where, in
midsummer and early autumn, bushels of these fungi may be collected. Thus
far no poisonous species have been detected north of the tree line although
the deadly toadstool ( Amanita phalloides ) has been found in the upper Mackenzie
basin and in the Yukon.
Seaweed . A number of edible species of seaweed or marine algae occur
along rocky shores of the arctic seas and several are used regularly, if
mostly in times of scarcity, by Eskimos. In Greenland, several species,
including ( Rhodymenia palmata and Laminaria spp.) are eaten raw or dipped
in boiling water or with seal oil. Rodahl (15) estimated that 50 per cent
of the vitamin C intake of the East Greenland Eskimos is derived from
marine algae.

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BIBLIOGRAPHY

1. Birket-Smith, Kaj. “Ethnography of the Egedesminde district with
aspects of the general culture of West Greenland.” Medd .
om Grønl . vol.66, 1924.

2. Bogoras, W. “The Chichee” Jesup North Pacific Expedition, Memoir ,
An.Mus.Nat.Hist., vol.vii, part 1, pp.147-149, 1904.

3. Brown, R. N. Rudmose. The Polar Regions , London, 1927.

4. Fernald, M.L. and Kinsey, A.C. Edible wild plants of eastern North
America .” Idlewild Press, New York, 1943.

5. Jenness, Diamond. “The life of the Copper Eskimo.” Rept . Can.Arct.Exp.,
1913-1918, Vol.xii, Ottawa, 1922.

6. ----. People of the Twilight , Macmillan, New York, 1928.

7. Kjellman, F.R. “Om tschuktschennas hushållsväxten,” Ymer , vol.6:e,
Stockholm, 1882.

8. Porsild, A.E. Edible Roots and Berries of Northern Canada . Nat.Mus.
Canada. Ottawa, 1937.

9. ----. “Flora of Little Diomede Island,” Trans . Royal Soc.Can., vol.32,
pp.21-38, 1938.

10. ----. “Emergency food in arctic Canada.” Nat.Mus.Can. Spec.Publ .45-1;
Ottawa, 1945.

11. Richardson, John. Arctic Searching Expedition , London, 1851.

12. Rink, H. Grønland geographisk og statistisk beskrevet . Copenhagen, 1857.

13. Rodahl, Kaare. “Content of vitamin C (1-ascorbic acid) in arctic plants.”
Trans. & Proc . Bot.Soc.Edinb., [: ] . vol.34, no.1.
pp.205-10, 1944.

14. ---. “Vitamin B1 content of arctic plants and animal tissue.” Ibid .
vol.34, no.2, pp.244-51, 1945.

15. ----. “Arctic nutrition,” Can.Geogr.Journ ., vol.30, no.2, pp.52-60, 1950.

16. Stefansson, V. My Life With the Eskimo . Macmillan, New York, 1913.

17. ---. “The Stefansson-Anderson Arctic Expedition of the American Museum
of Natural History, Preliminary ethnological report.”
Anthrop.Papers , Am.Mus.Nat.Hist., vol.14, part 1, 1914.

18. ----. Arctic Manual. Macmillan, New York, 1944.

19. Weyer, E.M. The Eskimos , Yale University Press, New Haven, 1932.

A. E. Porsild

Economic Botany of the Arctic

(EA-Plant Sciences. A. E. Porsild)

ECONOMIC BOTANY OF THE ARCTIC

CONTENTS
Page
Introduction 1
Direct Use of Arctic Plants by Man 2
Plants and Animal Husbandry 5
History of Reindeer Grazing 6
Reindeer Grazing 10
The Winter Range 13
The Summer Range 18
Nutritive Content of Lichens 27
Musk Oxen 29
Gardening 29
Conclusion 30
Bibliography 35

EA-Plant Sciences
[A. E. Porsild]

ECONOMIC BOTANY OF THE ARCTIC
Introduction
That “all flesh is hay” is as true in the Arctic as it is in any other
part of the world. Everywhere does animal life, whether on land or in the
sea, in the final analysis subsist on organic matter that in one way or another
has been produced by plants from inorganic matter. In the polar regions, how–
ever, all biological processes of growth and metabolism are retarded and reduced
in intensity by the low temperatures and, as we approach the Pole, by the de–
creasing length of the growth period. Therefore, there is a very definite
relation between productivity of the land and the distance from the equator.
In terms of abundance of organisms the most productive and fertile parts
of the polar regions undoubtedly are to be found in the sea; but the astounding
if local abundance, near the southern edge of the polar ice pack, of vegetable
and animal plankton that serve as food for larger arctic sea life of economic
importance, is made possible only by inorganic substances which have been carried
north into the polar seas by deep ocean currents from nonarctic regions.
Arctic land surfaces, on the other hand, in terms of productivity, are
everywhere among the lowest in the world. Thus, from the absolute zero of

EA-PS. Porsild: Economic Botany

productivity of land surfaces covered by perennial snow- or ice-fields, to the
relatively fertile flood plains occasionally found near the southern borders
of the arctic zone, the annual yield per unit of surface of arctic land is low.
Arctic vegetation everywhere is affected by the severe climate under which
plants grow. The shortness of the growing season — although to some extent
compensated by long daylight — and the deficiency of soil and precipitation,
have a more profound effect on plant growth than has the relative lowness of the
actual air temperatures as recorded by the meteorologist; for, due to insolation,
the actual microclimate in which the arctic plants live — i.e., the temperature
of the surface soil and the air or water surrounding the growing plant — may on
sunny days, and especially on south-facing slopes, be as much as ten or even
fifteen degrees Centigrade higher than that of the air.
Owing to poor drainage and aeration caused by the presence of permafrost
or bedrock closely below the surface — but particularly due to the low tempera–
tures — arctic and subarctic soils are generally acid and frequently also water–
logged. Therefore, organic decay by bacterial action or by other soil organisms
is extremely slow and the sources of available nitrogen, as well as of other salts
needed by the plant are almost everywhere deficient. Abundant proof of this is
found in the lush and rank growth with which many arctic plants respond in such
places as bird cliffs, near animal burrows, or on refuse neaps near human habita–
tions, where nitrogen and phosphates are plentiful.
Direct Use of Arctic Plants by Man
Everywhere in the Arctic, plant life plays a comparatively minor role in
the economy of man. None of the woody species of plants is of a size sufficiently
large for constructional use, and, before the advent of the white man, the

EA-PS. Porsild: Economic Botany

aboriginal inhabitants of the Arctic obtained such wood as they needed for the
construction of shelters and the manufacture of implements either from drift–
wood or by forays to the fringes of the forest to the south.
Heather and berry bushes, willow, alder, and ground birch, together with
lichen plants, moss, and peat are all used to some extent for cooking purposes.
Of greatest universal importance are, perhaps, the crowberry bush ( Empetrum )
and the white arctic heather ( Cassiope tetragona ), both common or even ubiquitous
in many parts of the Arctic. Nearly all the larger lichens, especially species
of Alectoria , Cladonia , and Cetraria are highly inflammable when dry and may be
used as fuel. Raw peat, particularly heath turf, but also partly decomposed
sphagnum moss which is found in bogs nearly everywhere in the Arctic may be burned;
indeed it provides the bulk of the fuel used by present-day Greenlanders.
The twigs of the larger willow, ground birch, and alder are used by the
peoples of the Arctic as floor coverings for tents or snowhouses or even in more
or less permanent dwellings, and are made into matting used under the sleeping skins.
The bark of several species of willow, and particularly the bark of alder
( Alnus ), is employed for tanning and dyeing purposes. A number of different
kinds of grasses and sedges find various uses. Thus in Greenland lyme grass
( Elymus ) is widely used for insoles in sealskin boots, and both there and elsewhere
for basket-weaving; while bluejoint grass ( Calamagrostis spp.) is used for
insoles, basket-weaving, and for matting under bedding.
Finely carded leaves of the sedge Carex goodenowii (and allied species) serve
the Lapps and various Siberian peoples as lining for winter boots and mittens.
Owing to its absorbent quality, sphagnum moss is widely used as a wick for
seal-oil lamps, as a lining for infants' cradles, and for “diapers.” The long,
silky bristles of several species of arctic cotton g grass ( Eriophorum spp.)
also provide wicks for Eskimo seal-oil lamps.

EA-PS. Porsild: Economic Botany

Several lichens find a limited use for dyeing purposes. Ochrolechia
( Lecanora ) tartarea produces a reddish color, while Parmelia omphalodes .
P. saxatilis , Lobaria pulmonaria , and Cetraria islandica yield yellowish-brown
colors.
Only a small number of arctic plants are regularly used for food by native
and white inhabitants (See article on “Edible Plants”). Of greatest potential
food value, perhaps, are the lichens, although, strangely enough, they are not
known ever to have been used by any of the aboriginal tribes inhabiting either
the New or the Old World. Several species of foliose black, grey, or green
lichens in the genera Umbilicaria and Gyrophora — the tripe de roche of the
Canadian voyageurs — grow on acid rocks throughout the polar regions. Together
with a few of the fruticose species, chiefly Iceland moss ( Cetraria islandica )
and reindeer moss ( Cladonia rangiferina and Cl. alpestris ) they have been used
regularly by arctic travelers, and, on more than one occasion have saved the
lives of field parties. In times of famine, Iceland and reindeer moss were used
in arctic Europe, to ske out the supply of bread flour; in the past, too, they
have been employed to a limited extent in the manufacture of alcohol.
Numerous species of edible fleshy fungi (mushrooms) are found, especially
in the southern parts of the Arctic but, like the lichens, they are not used by
the natives. However, several species of green and red marine algae (dulce or
sea lettuce) are eaten by the Eskimos of Baffin Island and Greenland, as well
as by several tribes inhabiting the seacoasts of Asia.
If arctic vegetation has little direct significance for man, indirectly,
nevertheless, it is of the greatest importance because nearly all sedges, grasses,
and fruticose lichens, besides other herbaceous plants and dwarf shrubs, provide
food for grazing and browsing animals. The seeds, bulbils, winter buds, and roots

EA-PS. Porsild: Economic Botany

of a host of arctic plants are eaten by birds and by small rodents, and these
in turn constitute the food of fur-bearing mammals. Likewise, the comparatively
rich marine plant life indirectly furnishes food for the sea mammals and fish
that are so important in the economy of arctic peoples. The role of plant life
in the balance of Nature, however, is beyond the scope of the present article,
which deals only with the direct use made by man of arctic plants, cultivated or
wild.
Nowhere in the Arctic does the cultivation of land — except on a small scale
and chiefly for experimental purposes — extend north of the forest limit, and
agricultural land use is everywhere restricted to grazing or to a very limited
extent to the harvesting of native wild hay, browse, or lichens.
Plants and Animal Husbandry
In southwestern Greenland the heroic medieval Norse colonization was based
on animal husbandry, and cattle, sheep, goats, and horses were maintained there
for several centuries. But the Norse economy remained marginal and succeeded only
when liberally supplemented by fishing and hunting. Nor was the economy self–
sufficient, and the colonies declined and finally perished when communications
with the motherland ceased.
In modern times sheep farming has been introduced in the parts of Greenland
formerly settled by the Norse. The industry, which is now being developed by
Greenlanders — the modern descendants of Greenland Eskimos — is still in its
infancy. Like the Nose husbandry that preceded it, it is decidedly marginal,
and it remains to be seen if it can survive, without government support, the
vicissitudes of an unfavorable climate and topography combined with a tenuous
and precarious economy.

EA-PS. Porsild: Economic Botany

Thus, such domesticated grazing animals as horses, goats, sheep, and cattle
have been, or are being, maintained in small and insignificant numbers in various
parts of the Arctic. Nevertheless, there is only one animal — the reindeer —
which is being successfully maintained there on any large scale. What is more
important still, it is in its habits a truly arctic animal, and the only one
around which the economy and culture of a large number of arctic and subarctic
peoples have been able to evolve and persist throughout the centuries.
History of Reindeer Grazing
Whereas domesticated reindeer were introduced into North America in compara–
tively recent times, and into Canada only a few years ago, the breeding and
grazing of reindeer is an industry of long standing in the arctic and subarctic
parts of the Old World. Thus, it is known from old Chinese manuscripts (13)
that as early as A.D. 499, reindeer were being employed somewhere in Asia as
draft animals and as beasts of burden, and that reindeer milk, too, was being
used extensively. This is confirmed by the recent discovery of rock paintings
in caves on the upper Yenisei and its tributaries, some of which show reindeer
drawing sledges or mounted by men; they are believed to date back to the be–
ginning of the Christian era (15; 32). In 1945, in the museum of Yakutsk, the
writer saw similar paintings, said to have been copied from the walls of caves
on the upper Lena; they were estimated to be 3,000 years old, and, while reminiscent
of the famous latte paleolithic rock paintings of France, depicted what undoubtedly
were reindeer rather than caribou. According to Mirov (15), Sosnovsky (30)
has reported Neolithic burials on the upper Lena, containing bones and harness
accessories of domesticated reindeer.

EA-PS. Porsild: Economic Botany

In Europe, the earliest historical accounts of domesticated reindeer come
from northern Scandinavia, whence reports of tame reindeer reached the court of
King Alfred, about A.D. 890 (13). Although, as yet, the exact place of origin
of the reindeer industry is not definitely known, it appears, then, that it may
have been somewhere in southern Siberia, east of Lake Baikal and, that, in pre–
historic times, it spread from there westward to arctic Europe, and northeastward
to the Asiatic shores of the Bering Sea, whence, in 1891, the first reindeer were
brought to North America.
Reindeer, like the wild caribou from which it evolved, while preeminently
an arctic animal, is not limited to the arctic tundra, for the earliest known
records come from forests areas deep in Siberia and in modern times there are
reindeer nomads whose herds never leave the forest.
The breeding and grazing of reindeer is thus an old and well established
industry around which a number of distinct nomadic cultures have evolved. It
is still an important industry which is practiced by nearly all aboriginal tribes
of arctic Europe and Asia. According to Mirov (15), the following peoples are
now, or were in the recent past, engaged in the industry:
Uralo-Altaic Group
A. Finno-Ugrian: (1) Lapp, (2) Karelian, (3) Zyrian, (4) Ostyak), (5) Vogul.
B. Tungus: (1) Tungus proper, (2) Lamut, (3) Oroki, and other small tribes.
C. Samoyed.
D. Turko-Tartar: Yakut.
E. Turkicized Samoyed-Yeniseian of southern Siberia: (1) Irkutsk Soyot,
(2) Uriankhai Soyot, (3) Karagas, (4) Kamasin.
Paleo-Asiatic Group : (1) Chukchi, (2) Koryak, (3) Yukaghir, (4) Chuvan, (5) Giliak,
and (6) Yenisei Ostyak.

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Herding practices and the use of reindeer differ materially among the
peoples who are engaged in the reindeer industry. Its highest development has
probably been attained in the Scandinavian peninsula by the Lapps who maintain
their herds under close and constant control, employ dogs for herding, use rein–
deer as draft animals and as beasts of burden, and who make extensive use of
reindeer milk. Opposed to this are the more relaxed herding methods employed
by nomads of the Siberian tundra, including Samoyeds, Zyrians, Ostyaks, Voguls,
and also the Chukchi and Koryaks. These people travel in both winter and summer
in sledges drawn by reindeer, but do not milk the [: ] animals. The Tungus, Karagas,
and Soyot of the Siberian taiga maintain small and closely tended herds, practice
milking, and use reindeer as draft animals and as beasts of burden; only the Tungus
have developed a reindeer that is large and strong enough to support the weight
of a man, and for this reason they alone ride reindeer. The use of dogs for
herding reindeer, apparently, is not practiced east of the Yenisei River.
Only in North America, where the industry was artificially introduced, and
is still largely in its infancy, are Eskimos, and in a few instances white men,
engaged in reindeer herding. The future of this Government-sponsored venture is
still very uncertain and rather closely tied up with the economic and educational
development of the Eskimos for, as their educational and economic standard improves,
reindeer herding to them becomes less and less attractive. In this connection it
is of interest to note that more than fifty years ago, Hahn (7), who probably had
not then heard of the [: ] experimental introduction of reindeer into Alaska, ex–
pressed the belief that the economic importance of this animal had by and large
been overestimated, as demonstrated by the fact that the industry was practiced only
by aboriginal tribes and that, even in regions where the reindeer was of cons i derable

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importance in the local economy, people of European culture and descent either
though the industry unprofitable or lacked themselves the necessary patience
and inclination for its development. The same idea was expressed a year later by
Jackson (1897) who wrote: “The ordinary white man is unwilling to undergo the
drudgery of herding in the rigorous climate, and unwilling to work for the small
compensation that is paid for such services.” Jackson, however, visualized economic
potentialities for the industry, for he goes on to say: “With the increase of
domesticated reindeer in Alaska, it will become possible for white men to own large
herds; but the men that will do the herding and teaming will always be Eskimo and
Lapp.” In Alaska this actually soon came to pass as reported by Palmer (17):
“From the original stock of 1,280 animals imported from Siberia over the period
of ten years up to 1902, the reindeer in Alaska have increased to about 350,000
head, distributed in 110 herds, all but 6 of which are along the coasts of Bering
Sea and the Arctic Ocean… In addition to the numbers in the present herds,
it is estimated that about 125,000 have been killed for food and clothing.”
Reindeer were introduced into Alaska solely for the benefit of the natives,
and for more than twenty years the industry developed as intended, prospering
beyond all expectations. In 1914, commercial exploitation began under white
ownership, with disastrous results to the native interest and, eventually, owing
to marketing difficulties, to commercial interests also. According to Palmer (17)
more than 1,875,000 pounds of reindeer meat was shipped from Alaska during the
period from 1918 to 1925, with approximately 1,000,000 pounds shipped in 1924
and 1925 alone. Nevertheless, the high cost of shipping, in view of the low prices
obtained in the American market, made large-scale operation unprofitable. Following
a period of chaotic decline, the United States Government in 1939 bought all reindeer
owned by whites and, at the same time, prohibited white ownership of reindeer. Since

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that date there has been a slow recovery but, owing to the rapidly changing
economic and social status of Eskimos in Alaska, few natives are now attracted
by the industry. In North America the methods of herd management that were in vogue
among the Chukchi and Koryak owners of the original herds, have been modified and
modernized by the introduction of Lapp methods, but still more by the adaptation
to the special requirements of reindeer of modern American livestock methods as
practiced on Western sheep and cattle reaches. In neither Alaska nor Canada are
reindeer milked, and very limited use is made of them as draft animals. Modern
elective breeding, modern corrals, and modern range utilization is practiced.
Everywhere the trend has been toward permanent settlements and absentee ownership
rather than the maintenance of the nomadic existence inherent in primitive
reindeer culture. The nomadic regime by which the reindeer owner [: ] [] and his
entire household followed the herd during its biannual migration between summer
and winter ranges, obtaining meanwhile practically all food and clothing from
the herd, did not prove compatible with the American way of life.
Reindeer Grazing
The food habits of the reindeer and caribou are essentially the same; for
both animals it is natural to perform seasonal migrations which follow a definite
pattern and route. It is uncertain, however, how far these migrations are dic–
tated by a search for food, and how far they are prompted by an urge to [: ] escape
the insect pests that each summer torment man and beast alike, on the subarctic
tundra and taiga. Seasonal migrations are not undertaken by certain species of
North American woodland caribou, by the barren ground caribou of Greenland, or
by the caribou that inhabit the northernmost islands of the Canadian Arctic

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Archipelago; nor are they practiced by the reindeer of the European and Asiatic
taiga. Regular migrations, on the other hand, are practiced by all reindeer
inhabiting the tundra areas, and also by the barren ground caribou of the North
American mainland.
The common belief that reindeer live exclusively on “moss” in erroneous. On
the contrary, like most other members of the deer family, both reindeer and
caribou are grazing and browsing animals, rather catholic in their choice of food.
Another misconception is the belief that the reindeer “moss” or lichen is abso–
lutely indispensable for reindeer because it possesses certain specific nutritive
qualities. The real reason why lichens are so important to reindeer is that
they occur very abundantly in certain habitats, and are there readily available
even under moderately deep snow. Without this ready source of food, large
herds of reindeer could not be successfully maintained throughout the winter
anywhere in arctic or subarctic regions. Without lichens, in other words, there
could be no reindeer industry.
Actually the nutritive qualities of the lichen plants are below those of
most other forage plants palatable to reindeer; thus, the protein content of
air-dried lichens ( Cladonia spp.) is only about one-third that of native, air–
dried, and summer-out hay; the fat content is about the same and only the starch
or digestible carbohydrate content of the lichen is slightly higher. The great
and all-important difference is that the nutritive value of the lichen plant
remains unimpaired throughout the winter, whereas that of winter-killed grasses,
sedges, and forbs is negligible. For these reasons, lichens are unimportant
on the summer range but, during 8 or 9 months of the year, from the bulk of
the forage available and eaten by reindeer. Therefore, the practical reindeer

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man, or the field botanits engaged in a study of potential reindeer pasture,
must look for winter range having abundant lichen cover which will be available
to reindeer throughout the winter. Likewise, on grazing land already occupied
by reindeer, it is the winter range that must be conserved and carefully protected
against overgrazing or needless destruction by trampling reindeer herds, and
against destruction by fire. The summer range is expendable and will renew it–
self each summer.
Although the food habits of reindeer and caribou are much the same, it
does not follow, however, that domesticated reindeer will thrive in country
that will support caribou. While caribou may roam the length and breadth of
the country in search of their food, the reindeer, in order to stay domesticated,
must remain, at least to some extent, under the constant control and supervision
of its owner, for only thus can it remain useful to him. For this reason,
reindeer can be successfully maintained only in country sufficiently productive
to permit the slow and leisurely grazing of large herds during both summer and
winter. It is of equal importance that the distance between the summer and the
winter ranges should not be too great to permit the leisurely movement of the
herd twice a year, with a resting period allowed in the spring during fawning,
and in the autumn during the rutting season. To prevent the deer from straying
on the range, especially in winter, and to protect them from wolf attacks, the
herders must walk around the herd each day. The available lichen cover, therefore,
must be such as to permit the herd to remain within a fairly limited area for
several days, or even weeks. In order that the range may not become unduly
depleted, the deer man must adopt a rotation grazing system, selecting in
advance a range where conditions are suitable for his deer during each particular
season.

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The Winter Range
Although reindeer and caribou are wonderfully adapted to withstand extreme
low temperatures and high winds, unfavorable climatic conditions nevertheless
affect their general health and well-being no less than the condition and availa–
bility of the range. When in good condition, reindeer can stand prolonged periods
of extreme cold with impunity, but only if well-fed and when some shelter is pro–
vided against wind.
In some parts of theArctic, where winter ice does not normally cover the
sea, or where the shore ice does not extend far [: ] out, rain may occur during the
winter and cause serious crusting of the snow. When this happens, reindeer herds
may suffer severe losses through starvation, when the animals are unable to dig
through the hard, icy crust. Such conditions, however, ra r ely extend far inland.
The average annual precipitation is low everywhere in the Arctic, but especi–
ally in the continental parts, where it is generally less than 7 inches. The
winter snowfall, therefore, it light, and under ordinary conditions the depth of
snow on the coastal tundra does not seriously affect winter [: ] grazing, especially
because the frequent winter gales, to which the area is exposed, sweep the snow
off the tundra or pack it into drifts that fill the depressions. At some distance
from the exposed seacoast, the edge of the forest, open muskeg-filled depressions,
lake basins, and mountain valleys, afford protection from the prevailing wind
and freedom from drifting. In such places the snow, ther f e fore, remains soft.
On comparatively flat or level ground it seldom exceeds 12 to 18 inches in depth,
and only in depressions and ravines does it become too deep for reindeer to paw
through to the vegetation.
The presence of shelter and abundant snow cover affect also the vegetation.

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At no great distance from the coast, and in the greater part of the [: ] treeless
“Barren Grounds” or tundra a very different and more luxuriant type of vegetation
replaces the sedge-grass tundra of the coast. Traveling on foot soon becomes ex–
ceedingly tiresome, because the ground everywhere is covered by low scrub or
heath that grows in a thick and soft carpet of mosses and lichens into which the
traveler [: ] sinks ankle-deep at every step. Low willow, alder, and dwarf birch,
together with other dwarf shrubs of the tundra dominate the herbaceous vegetation;
in sheltered places, along streams and on the steep cut-banks of lakes, they
may even form dense thickets.
Although the tips the young shoots of these shrubs — willow, ground birch,
and alder — are much sought after by the reindeer in spring, autumn, and early
winter, it is the relative abundance, texture, distribution, and composition of
the lichen cover that determines the value of the winter range. For this purpose
the ideal forage cover is a mixture of dwarf shrubs and lichen, usually design–
nated the lichen-browse type. On the coastal winter range of Alaska, Palmer
(17) found the following average composition, expressed in percentages of an
average total ground cover of 71%: Lichen 52%, browse 17%, sedge 20%, herbaceous
nonpalatable species 6%, and nonpalatable moss 5%. On winter range in the deep
interior of Alaska he found a total average ground cover of 80% with the follow–
ing composition: Lichen 52%, browse 21%, sedge 10%, nonpalatable species 5%,
moss 12%.
Because lichens furnish the bulk of the forage on the winter range, and in
general are so indispensable to reindeer that without them there could be no
reindeer industry, it may not be amiss to examine them a little more closely. These
curious plants are really dual organisms, consisting of fungi that are parasitic

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on, and receive their nourishment from certain forms of primitive green or blue–
green algae that, themselves, are completely enveloped by, or diffused through,
the hyphae of the fungus. The alga can, and does, exist independently, but the
fungus cannot: together they live in an intimate partnership known as symbiosis.
The green chlorophyll of the alga assimilates carbon from the carbon dioxide of
the air by photosynthesis; the fungus absorbs water from in the substratum or
from the atmosphere, and provides shelter and protection for the alga.
The relationship between these two entirely different plants is so intimate
and close that for practical reasons, as well as for the purpose of classification,
the symbiotic colonies of fungi and algae called lichens are generally treated as
taxonomical units, and, like normal plants, are divided by the taxonomist into
families, genera, and species.
Reproduction in the lichen plant can be sexual by means of spores produced
by the fungus, or asexual by means of soredia which are singl algal cells or
groups of them, enveloped in hyphal tissue, and capable of growing at once into
a thallus when detached. The soredia generally originate in the gonidal layer
of the lichen thallus and usually appear as fine powder that, when conditions
are favorable, germinate immediately to form new plants. Finally, fragments of
the lichen plant after becoming separated from the mother plant, and dispersed
by wind or water, may continue to grow.
Although the lichen spore and the soredia germinate readily, the lichen
plant itself grows very slowly and never becomes very large in size; because the
growth is terminal and apical, the lichen colony, so to speak, is ageless and may
continue to grow as long as conditions are favorable. The lichen plant is very
long-lived; some species growing on rocks, where they form a hard, leathery crust

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( Lecanora , Pertusaria , Buellia , etc.), may be several hundred years old. Lichens
growing on soil are rather delicate as a rule and, lacking proper roots, are
not firmly anchored to the soil. Therefore, the mat they form is easily removed
or destroyed by picking or by trampling, especially when dry, when it is very
brittle. When moist or wet it is spongy or cartilaginous and not easily injured.
Botanists distinguish a large number of different species of lichens, several
hundred of which occur on the arctic and subarctic tundra. Their palatability
to reindeer varies, but with the exception of a few that are avoided, probably
because of their bitter flavor ( Cetraria nivalis ), all are eaten rather indis–
criminately. On a few species, however, because of their abundance and high
palatability, are of primary importance. Except for minor variations these are
all widely distributed and of circumpolar range.
Lichens grow in a great many different habitats and are adapted to widely
varying climatic conditions; and their geographical as well as vertical range
is very considerable. All boreal and arctic species are capable of enduring
prolonged and intense desiccation as well as extreme ranges in temperature.
Most commonly lichens grow on soil, rocks, or on the bark of trees, but they
may also grow on decaying wood, on the top of mesophytic mosses, on the thallus
of other species, or even on sun-bleached bones. According to the substratum on
which they grow, lichens are said to be berricoline, saxicoline, carticoline,
or muscicoline. Those most useful as forage plants for reindeer nearly all
belong in the first category, although species of Alectoria growing from the
branches of trees are of some importance. Nearly all lichens are light-loving
and develop poorly or not at all in shade. For their growth they require [: ] ,
besides light, moisture and heat. Their very small mineral content may be derived

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in part from the substratum or more likely from air-borne dust, but their prin–
cipal food comes from assimilation and photosynthesis of the carbon dioxide of
the air.
As previously mentioned, the terricoline species of lichens require a
protective snow cover in winter; for this reason lichen range is best developed
on sheltered slopes and in not too moist depressions. On the open tundra where
the competition from grasslike and other herbaceous plant is keen, the lichens
most often grow on the sides of hummocks. In open parklike forest, near the edge
of the forest, almost pure stands of lichen mats frequently form the ground cover.
Rarely does one species alone form extensive colonies; most often several species
grow together, mixed with true mosses such as Sphagnum , and Polytrichum . Often a
handful of lichens, picked at random, may contain half a dozen or more closely
entwined species.
On low and fairly moist tundra and is the lower foothills, the predominant
species are often Cladonia sylvatica and Cl. rangiferina. frequently mixed with
small amounts of Cl. alpestris , Alectoria ochroleuca , Dactylina arctica , Cetraria
islandica , and others. On higher ground with favorable exposure and shelter,
Cladonia alpestris is often the dominant species, whereas on more exposed and
wind-swept ridges Catraria nivalis , Alectoria orchroleuca , and A. nigricans pre–
dominate. Other species of frequent occurrence are: Cladonia gracilis , Cl .
crispata , Cl. amaurocraea , Cl. uncialis , Cl. Delessertii , Cl. decorticata , and
Cl. squamosa , Cetraria cucullata and C. Tilesii , Stereocaulon tomentosum , S .
paschale and Thamnolia vermicularis , Nephroma arcticum , Dactylina arctica and
Sphaerophorus globosus . All these, and many more, are eaten by the reindeer but
only a few of them are really important. A brief description of three of the most
important species, commonly referred to by the inclusive term “reindeer moss,” may

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be appropriate: Cladonia alpestris , when fully developed is usually 3 to 4
inches long but under most favorable conditions may reach 8 or even 10 inches
in length. The growth is terminal and apical and the lower part of the plant
dies off and slowly decays. The color is light gray with a yellowish-green tinge
which is most noticeable when wet. The plant is dense and coral-like and richly
branched, with no clear differentiation between stems and branches; the latter
are round in cross section and hollow, often joined or fused together, forming
small, hemispherical, cupola-like clusters so that, seen from above, the lichen
mat may resemble masses of eggs spread on the ground. Cladonia rangiferina and
Cl. sylvatica attain a similar size and are of a similar structure, but differ
from Cl. alpestris by their less branched growth. Also the tips of the branches
are more forked than spreading, often clawlike and the branches do not rejoin above
the first fork. While structurally alike, the last two are easily distinguished
by their color. Thus, Cladonia rangiferina is of a darker gray, often somewhat
brownish, with a purplish tinge when wet, whereas Cl. sylvatica is pale gray or
yellowish, with a greenish tinge when wet.
The Summer Range
Late in March or in early April, when the backbone of the winter has been
broken and the days are getting long, reindeer herds being to move toward the
summer range. The does are now heavy with fawn, and before the first young are
born in late April, the deer men must select a suitable fawning place, generally in
the foothills adjacent to the summer range. The fawning place must be well shel–
tered, for even in April blizzards may occur; and not infrequently does the night
temperature drop to thirty degrees below zero. Also, there must be abundant forage
of lichen and browse, so that, during the two months spent at the fawning grounds,
the does and young fawns need not travel far to feed. On the fawning grounds the

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males and steers are separated from the does and are generally held in a
separate herd close by.
From the fawning place the reindeer herds slowly move onto the summer pas–
ture. By mid-June most of the snow will have disappeared from the low tundra
which, however, is still very wet; but most of the larger lakes are still ice
covered and the herds can cross at will, keeping to the higher and dry ground
between lakes.
From the end of June to mid-August is the height of the fly season on the
tundra, when, on calm and hot days, mosquitoes and gnats make life unbearable
for the reindeer as well as for the men who tend them. On such days it is im–
portant that places can be found on sand points or headlands near the seashore,
or on high bluffs or hills where cool breezes or fog from the sea provide some
relief against these pests. The first to arrive are the mosquitoes, followed
in July and [: ] August by the blackflies or gnats. About this time the larvae
of the reindeer warble fly ( Oedamagena tarandi ) and the nostril fly ( Cephenomyia
nasalis ) that were already dropped in June on the fawning grounds, have emerged
from their pupal stage and now overtake and torment the reindeer.
For the summer pasture, country of low relief is desirable. The ground
should not be too dry or stony because, otherwise, the reindeer are liable to
foot injury, nor should it be so wet as to cause foot diseases such as foot-rot
and dermatitis. However, if open herding is practiced, little injury to the
reindeer [: ] need result from these causes.
On the summer range the food of the reindeer consists largely of the grasses,
sedges, and other herbaceous perennials of the arctic tundra to which, depending
on the type of range, may be added the young leaves and twigs of willow and [: ]
dwarf shrubs and a variable amount of lichen. The lichen is not necessary for the

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reindeer on the summer range, but is readily eaten when moist. In addition,
all kinds of fleshy fungi are devoured greedily by the reindeer, as are the
eggs and young of birds nesting on the tundra, as well as the young of lemming
and other microtine species of the tundra. This seemingly abnormal carni–
vorous habit of the reindeer, together with their apparent fondness for gnawing
bones and the tips of shed antlers is, perhaps, due to a carving for salt
developed during the winter away from the seacoast. On the summer pasture
reindeer are often seen to be feeding on halophytic seashore plants and
kelp, and even to be drinking sea water.
Although the reindeer on the summer pasture do not show any pronounced or
clearly marked preference for definite species, they to avoid coarse vegetation
and appear to seek out the nibble only the most tender and young shoots. In
their grazing habits they resemble horses more than sheep or cattle. Reindeer
do not bite off the vegetation down to the roots but in a day’s grazing wander
over a considerable area of pasture, nibbling a few leaves of tender grass here
and the young shoots of a willow there. For this reason, the vegetation on
rather wet and soft pasture suffers more from their trampling hoofs than from
their actual grazing. But whereas the continued trampling [: ] of a reindeer
herd within an enclosed pasture quickly destroys the forage and soon also the
complete turf, the over-all effect produced by the mechanical “tilling” of grazing
reindeer on the summer pasture is beneficial rather than otherwise. It is true
that lichens suffer and in time disappear; also that a number of species, especi–
ally woody plants, may become stunted under continued browsing. Grasses and
edges, on the other hand, as well as a number of other perennial herbaceous plants,
benefit and increase by the continued “tilling” of grazing reindeer. Most of the
tundra plants reproduce vegetatively as well as from seed. Consequently, by good

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management, which involves rotation cropping, the summer reindeer pasture will
recover from grazing and even improve in quality.
On the coast summer range of Alaska, Palmer (17) has estimated that the
palatability of the forage cover average about 62%. True mosses undoubtedly
constitute a larger percentage of the unpalatable species than the 4% he has
assigned to them. Among other nonpalatable species are such coarse and tough
plants as the lyme grass ( Elymus ) and most species of rushes ( Juncus ). As far
as is known, no arctic or subarctic plants are poisonous to reindeer. In Alaska
even water hemlock ( Cicuta ), elsewhere poisonous to livestock, is said to be
palatable.
Although reindeer on the summer range eat a great variey variety of range
plants, only a comparatively small number, because of their palatability and
greater abundance, are of primary importance. On the coast range of Alaska,
Yukon, and northwestern Mackenzie some of the more important forage species are:
cotton grass ( Eriophorum vaginatum ), willow ( Salix spp.), ground birch ( Betula
glandulosa ), reindeer “mose” ( Cladonia app.), horstail ( Equisetum arvense ), sedge
( Carex app.); nearly all grasses, including Arctagrostis , Poa , Dupontia , Festuca ,
Alopecurus , Puccinellia , and others; fernweed ( Pedicularia spp.), lupine ( Lupinus
arcticus ), sour dock ( Rumax arcticus ), bilberry ( Vaccinium uliginosum ), sage
( Artemisia spp.), and fireweed ( Epilobium latifolium and E. angustifolium ).
During the migration from the summer range back to the winter range the
reindeer again gradually return to their winter diet of browse and lichen. During
this period they are often seen nibbling the mature seed pods and the ripened
fruiting inflorescences of grasses and sedges. It is during the late summer
and fall that the reindeer put on their back fat.

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Carrying Capacity of Reindeer Pasture
Range studies and controlled grazing experiments which have been carried
out in Alaska by Palmer (17; 18; 19; 20) and in northwestern Canada by Porsild
(21; 22; 23; 24; 25) have shown that the arctic tundra and taiga of North
America is capable of a sustained yield for reindeer that, on a per acre basis,
is comparable or superior to that of short-grass sheep or cattle-range of the
western United State and Canada. Thus Palmer found that, on average tundra–
coast range in Alaska, the minimum year-long grazing area required for each
adult reindeer is 33 acres. On this basis he (1929) gives the following seasonal
land use:
Spring and early summer (2-1/2 months) 4 acres
Summer (2 months) 2 acres
Fall and early winter (4 months) 10 acres
Winter (3-1/2 months) 17 acres
This, however, is a minimum, and for most parts of Alaska, he recommends from 40
to 60 acres per head for year-long grazing. Because the lichen plant is so
easily destroyed by mechanical injury, and because it grows at such slow rate,
the winter range with its all-important constituent on “reindeer moss” is the
first to suffer from overgrazing or from poor range management. When reindeer
were first introduced into Alaska, the immediate coastal areas contained a con–
siderable cover of lichens (10). In these areas lichens have now largely dis–
appeared, owing probably, to the close confinement of the herds to the coast.
The new cover consists almost entirely of herbaceous and shur shrubby vegetation
— sedges, low species of browse, and grasses. On some other ranges similar
changes due to grazing and fire may be expected — changes that on the whole
will probably prove beneficial to summer pasturage but detrimental to the
maintenance of winter forage (18).

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In Alaska Palmer (18) estimated that an area of 300,000 square miles of
tundra and sparsely timbered land was suitable and available for reindeer
grazing, and that, based on year-long grazing requirements of from 40 to 60
acres per animal, this area was capable of supporting indefinitely 4,000,000
reindeer. In the coast section between Bristol Bay and Point Barrow, where
the maximum development of the industry occurred, there was, according to
Palmer, room for 1,000,000 reindeer. In 1926, the official estimate of the
total number of reindeer maintained there was 350,000. There is reason to
believe, however, that this figure, based in part on estimated increases, was
too high, and that the largest number reached before the general decline of
the industry set in could not have been over 250,000. Although the decline
of the reindeer in Alaska was largely due to causes other than depletion of
the grazing range, it seems probable, in view of later developments, that the
earlier estimates were too optimistic and that the total carrying capacity
of all available reindeer grazing land in Alaska would not be in excess of
1,000,000 head.
The arctic tundra and taiga of Canada has been the home of vast numbers of
caribou since it was first explored and probably ever since the retreat of
the great ice sheet that once covered the land. These caribou generally spend
the winter in or near the edge of the forest and in summer migrate north into
the open tundra. Until about 30 years ago large numbers of caribou from the
continent each spring crossed the ice-bound sea to spend the summer on the
larger islands of the Arctic Archipelago, probably to avoid the clouds of
mosquitoes and gnats that tormented them on the mainland. Untol generations
of Indians and Eskimos have depended on them for a large part of their food and
clothing, and it appears that until the introduction of modern firearms a balance

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had been established between the natural increase of the animals on the one
hand and the combined human d and wolf predation on the other. With the
coming of the white man this equilibrium was soon disturbed, and while there
still are large herds of caribou left in the Canadian tundra and taiga, con–
siderable alarm has been felt over the reported decreases. Judging from
personal observation and the reports of earlier travelers, Seton (28) estimated
the northern caribou population at 20,000,000, while more conservative observers
have placed the probable number at between 1,000,000 and 2,500,000. In the
light of recent information obtained from the use of aircraft and aerial photo–
graphy, personal interrogation, and other sources, it would seem that while
perhaps the earlier estimates greatly exaggerated the numbers that could possibly
have subsisted in the Canadian North, there is, nevertheless, abundant evidence
that the herds there are rapidly decreasing in numbers and that the present
caribou population of continental Northwest Territories may not be over 670,000
(3).
In 1919 , m a Royal Commission was appointed to report upon the possibilities of
reindeer and musk-ox industries in arctic and subarctic parts of Canada as a
practical means to preserve Canada’s northern wildlife resources and, at the
same time, to provide the develop new sources of food, clothing, and livelihood
for Canadian Eskimos and Indians. The Commission recommended (1) the estab–
lishment of experimental herds of reindeer in selected locations in the Canadian
North; the exact location of the experiments to be determined by a general botanical
reconnaissance which was to have special reference to reindeer pasture, carrying
capacity, and other general conditions of importance to a future reindeer industry.
In 1927-28 the writer was placed in change of a detailed survey of the grazing
possibilities of the treeless country along the Arctic coast, from the Alaska-Yukon

EA-PS. Porsild: Economic Botany

boundary east to the Coppermine River, and south to the north shore of Great
Bear Lake (21). Following the investigation, arrangements were at once made by
the Dominion Government for the purchase of a herd of reindeer from Alaska, to be
delivered to a selected area lying immediately east of the delta of the Mackenzie
River. In March, 1935, delivery was made of a herd of 2,370 head of reindeer to
the Government corral near Kittigazuit. During 1931-35 the grazing studies
were continued in the country east of the Mackenzie Delta, where, in 1935, a tract
of land of approximately 6,600 square miles was set aside as the Mackenzie Delta
Reindeer Grazing Reserve. In 1930 a similar grazing survey was undertaken in
central Keewatin.
These investigations demonstrated that in the Mackenzie District two areas,
formerly densely populated by caribou, were eminently suitable for the maintenance
of domesticated reindeer. The carrying capacity of an area adjacent to the
Mackenzie Delta was estimated to be at least 250,000 head of reindeer whereas
a much larger, and physiographically rather different area north and east of
Great Bear Lake was estimated to be able to support 300,000 head (21). When the
original survey was made this whole region was practically unmapped. Accurate
maps, based on aerial surveys have since disclosed that in this region more
than half of the land surface is occupied by lakes and ponds. In view of the
much higher ratio of water to land, the writer’s original estimates of the
carrying capacity should be reduced proportionately.
In 1946, the writer again examined the summer and winter grazing range in
the Mackenzie Delta Reserve. An area of approximately 800 square miles of upland
tundra adjacent to the main reindeer station had been grazed each winter, during
the preceding 12 years, by a herd of approximately 5,000 reindeer. Examination
of this range, together with ungrazed control areas, indicated that during the

EA-PS. Porsild: Economic Botany

12-year period no more than one-third of the potential winter pasture within
this area had been utilized and that the remainder had been left almost completely
untouched. A small area of approximately 57 square miles, in close proximity
to the station, showed slight evidence of overgrazing, whereas the rest was
quite unimpaired and capable of providing winter grazing indefinitely for a
herd of 5,000 reindeer (25).
From these observations it would appear that under conditions such as those
described above practically no damage results from winter grazing, provided
that the range is not used until the ground is frozen and covered by snow. The
reason for this is that, when grazing over the frozen lichen carpet, the reindeer
merely nibble the tips of the plants. The tips of the branches are the only
parts of the lichen plant that are capable of growth, and, when lightly grazed,
are able to regenerate in a few years; furthermore, “holes” plucked into the
lichen carpet by grazing, quickly fill in by lateral growth and expansion of
the lichen mass. On the other hand, if the entire upper layer of the lichen
carpet is destroyed by grazing or trampling, regeneration ceases or, at best,
is very slow. Also, when this happens, other plants such as dwarf shrubs,
lichens, and grasses, or true mosses, compete for the space formerly occupied
by the lichens.
Available information indicates that the carrying capacity of reindeer
range in arctic Europe and Asia is similar or at least comparable to that of
North America. Thus, in the Anadyr province of eastern Siberia, which physio–
graphically is comparable to northwestern Alaska, Soviet investigators have
found that one adult reindeer for 8 months of year-long winter grazing s required
the following pasture:
Forest tundra 21 acres
Scrubby tundra 23 acres
Non-scrubby tundra 42 acres

EA-PS. Porsild: Economic Botany

whereas on the summer range 20 acres are needed for each adult deer for a 4–
month period when the daily requirements of green herbage was found to be
about 40 lbs. (6; 34.)
Nutritive Content of Lichens
In the mountain districts of Norway, and perhaps elsewhere, lichens have
long been harvested regularly and fed with native hay to cattle, sheep, goats,
pigs, and horses; fish meal is generally added to make up for their low protein
and fat content. Isaachsen (9) reported that the water content of lichen
( Cladonia spp.) varies under natural conditions from 56% to 80% but when air–
dried sinks to 15%-18%. In the latter condition the lichen contains about
2.5% protein — roughly about one-third of that in wild hay — and an equal
amount of fat, whereas raw lichen of 65% water content averages 0.65% digestible
protein 0.6% fat, 12% carbohydrate, and 8.5% cellulose. Isaachsen further
stated that 6.5 kilo of raw lichen equals in food value 1 kilo barley or 2.5
kilo [: ] wild hay, whereas in air-dried condition (15%-18% water content) it
approximately equals wild hay.
Palmer (19) quotes somewhat higher nutritive values derived from samples
of lichens collected in the interior of Alaska, but, as neither he nor Isachsen
[: ] states the method of analyses, their figures are not directly comparable.
Palmer’s are given in Tables [: ] 1 and 2 below ; :
TABLE 1. - Analyses of samples of lichens collected in the interior of Alaska
TALL-GROWTH TYPE, OR MOIST-SITE LICHENS
Sample Moisture Fat Fiber Protein Ash Nitrogen–
free extract
Italics Cetraria cucullata 12.22 8.70 9.42 1.75 1.27 66.46
C. islandica 11.85 2.08 8.53 3.13 1.89 72.52
Cladonia alpestris 12.35 1.92 43.98 2.13 2.33 37.29
C. amaurocraea celotea 12.61 1.55 35.68 1.73 1.48 46.95
C. a. oxyceras 11.88 1.78 33.56 1.50 1.39 49.89
C. sylvatica (light form) 12.66 1.45 31.98 1.75 1.81 50.35
C. sylvatica (dark form) 13.02 0.57 44.64 1.50 2.05 38.22
C. s. sylvestris 12.93 1.08 48.92 1.67 1.59 33.81
C. rangiferina 12.83 0.69 47.19 1.75 1.78 35.76
C. uncialis 12.89 1.23 37.26 1.50 1.78 45.34
C. gracilis 12.46 0.85 45.72 2.50 1.79 36.68
Average 12.52 1.99 35.17 1.90 1.74 46.68
SHORT-GROWTH TYPE, OR DRY-SITE LICHENS
Italics Cladonia gracilis dilatata and C.
bellidiflora hookeri
12.15 0.89 33.27 3.50 2.92 47.27
C. gracilescens 13.27 0.56 40.08 3.06 2.64 40.39
C. degenerans 12.90 0.76 58.29 3.56 2.21 22.28
C. decorticata 13.04 1.14 40.15 4.25 6.27 35.15
C. crispata 12.56 1.34 43.70 2.25 1.85 38.30
Cetraria hiascens 14.13 5.23 11.18 2.94 1.90 64.62
C. nivalis 13.72 4.27 8.26 1.87 2.69 69.19
Dactylina arctica 13.12 5.94 8.52 2.81 2.54 67.07
Stereocaulon tomentosum 12.66 1.94 27.32 5.44 2.09 50.55
Peltigera spp 13.41 1.12 21.93 17.12 7.91 38.51
Average 13.10 2.32 29.27 4.68 3.30 47.33
ANALYSES OF OTHER FORAGE
For comparison with the lichens, analyses of samples of other fall and winter forages are given
in table 2. Specimens were collected during the first part of December.
TABLE 2. - Analyses of samples of certain fall and winter forage other than lichens
Sample Moisture Fat Fiber Protein Ash Nitrogen–
free extract
Percent Percent Percent Percent Percent Percent
Italics Eriophorum callitrix [ vaginatum ] e) 6.86 1.60 34.84 3.69 2.81 50.20
Salix fluviatilis [ pulchra ] 8.41 2.24 15.74 9.63 3.45 60.53
Chamaedaphne calyculata eaf) 6.10 9.80 23.33 6.25 1.93 52.59
Average 7.12 4.55 24.64 6.52 2.73 54.44

EA-PS. Porsild: Economic Botany

Musk Oxen
The musk ox is adapted to life under extreme arctic conditions to an even
higher degree than the caribou and reindeer. At one period this animal ranged
over the greater portion of the northern half of the North American continent,
but in modern times its ranges has been greatly reduced. From time to time
musk ox calves have been successfully reared in large zoological gardens, and
since 1930 the United States Fish and Wildlife Service in Alaska has conducted
an experiment with East Greenland musk oxen with a view to re-establishing the
species there. Thus far the experiment has shown that musk oxen can indeed be
bred in captivity, but that they increase very slowly. Consequently any attempt
to domesticate the animals, or to raise them commercially, would seem to have
little hope of success. They have responded well, on the other hand, to complete
protection, both on the Canadian mainland and in the islands of the Arctic
Archipelago.
Gardening
Small-scale gardening may be possible in the more favored parts of the Arctic;
in southern West Greenland, at any rate, lettuce, spinach, and rhubarb, besides
potatoes and a number of other root crops may be grown for local consumption;
and even far beyond the Arctic Circle vegetables are grown successfully under
glass. Lack of soil, however, in most parts of the Arctic makes the development
of gardens difficult and costly.

EA-PS. Porsild: Economic Botany

Conclusion
In the Northern Hemisphere that vast tract of land lying north of the
boreal forest or taiga, variously known as Arctic Tundra, Barren Grounds,
Northern Plains, or Arctic Prairie, everywhere is sparsely populated, and from
the point of view of either fur production of grazing, whether by game animals
or by domesticated animals such as reindeer, is among the least productive land
areas in the world.
In the U.S.S.R., Voshchinin (33) estimated that 1,150,000 square miles of
arctic tundra and 2,350,000 square miles of taiga was unsuited to agriculture.
In North America all but a small portion of the million and a half square miles
that comprise the Northwest Territories and Yukon likewise is wasteland from
the point of view of the agriculturist, as are also the northern and north–
western parts of Alaska.
In the Old World the wild caribou that once roamed the tundra and the northern
taiga have long ago disappeared, and in some areas have been replaced by domesti–
cated reindeer. No reliable figures are available giving the area that today
is utilized by reindeer. Balzak, Vasyutin, and Feigin (2), however, state that
“by the end of the Second Five Year Plan there were 1,800,000 reindeer in the
U.S.S.R.,” and on their map (Fig. 65) they show that the greatest concentration
is in the tundra area between the White Sea and th River Ob, and in eastern
Siberia east of the Kolyma River; the map also shows that the southern limit
of reindeer breeding extends far south of the treeless tundra. Unfortunately, no
information is given by them as to the number of people in the U.S.S.R. who
depend on reindeer for a living. Perhaps the most reliable information is
supplied by Mecking (14) who estimated that “In the Arctic fringe, the total

EA-PS. Porsild: Economic Botany

population is probably hardly more than 30,000. This population is made up of
Lapps, Samoyeds, Ostyaks, Dolgans, Tunguses, Yakuts, Yukagirs, Aleuts, Koryaks
and Chukchis.” Most of these, with the exception of the Aleuts, at least
formerly were reindeer nomads.
In Alaska the northern tundra and taiga once was inhabited by large herds
of wild caribou that, in a large measure, contributed to the native economy.
But even before the end of the 19th century these herds had all but disappeared,
and reindeer were introduced into Alaska in order to provide food and clothing
for the Eskimos. On mountain ranges of the interior, the caribou held out
longer than on the coast, but even there the last remnants of the once numerous
caribou population are fast disappearing (12).
Colby (5) estimated that from 13,000 to 15,000 natives of Alaska, including
dependents, rely on reindeer as an essential source of food and clothing. This
figure, even in 1937, may have been an overestimation. At present, at any rate,
the dependence on reindeer is very much smaller for, according to Lantis (12)
there are today only 27,920 reindeer in Alaska.
According to the latest census (1947) the native population of the Northwest
Territories comprises 5,651 Eskimos and 4,334 Indians. Only the few hundred
Eskimos the Indians of the Mackenzie Delta area area directly or indirectly
dependent on the reindeer industry, wh ci ic h, in Canada with a total of only 7,500
reindeer, is still largely in the experimental stage.
In some sections of the Canadian North wild caribou are still plentiful,
although much reduced in numbers. Banfield (3) estimated that, on the Canadian
mainland, in an area of 600,000 square miles of tundra and taiga, the
caribou is one of the basic factors in the economy of approximately 20,000
Canadians.

EA-PS. Porsild: Economic Botany

In Canada the musk oxen, not long ago threatened by extinction, now enjoy
complete and year-long protection. As a result they are again increasing every–
where in Canada, and slowly reoccupying former range.
There are, however, large areas in northern Canada that [: ] today, although
well suited to domesticated reindeer, are no longer occupied by caribou. Probably
more than 1,000,000 head of reindeer could be successfully maintained there,
provided that native or white herders could be found who were willing to live
the strenuous and hard life of a reindeer herder. But for efficient manage–
ment, one million reindeer would have to be divided into at least 500 units,
each requiring about 10 herders for its proper care. There are, however, today
only 9,000 Eskimos in all of Canada, and only a few of them live in areas that
are suitable for reindeer herder. But for efficient management, one million
reindeer would have to be divided into at least 500 units, each requiring about
10 herders for its proper care. There are, however, today only 9k 9,000 Eskimos
in all of Canada, and only a few of them live in areas that are suitable for
reindeer. There are, to be sure, much larger numbers of Indians; but Indians
are forest dwellers, and by temperament and tradition are even less []
inclined than are the Eskimos to give up their traditional hunting habits and
freedom in order to settle down to the monotonous drudgery of reindeer [: ] herding.
Also, as with the Eskimos, the idea of owing and caring for property is quite
foreign to them and can only be acquired slowly and by degrees.
Of the 22,000 native inhabitants of Greenland, only a few hundred are today
directly engaged in sheep farming; however, most of the products from the sheep
industry are used in the country and in this manner benefit a considerably
largely number of Greenlanders. In the fjord district of the west coast caribou
were once numerous, and until the middle of the last century provided an important

EA-PS. Porsild: Economic Botany

source of food and clothing. Thus, in the years between 1820 and 1830 Rink (26)
estimated the annual take at 37,000. As elsewhere in the Arctic, however, the
introduction of firearms seriously depe depleted the stock. In recent years
regulation of the hunting appears to have checked the decline, so that for a
number of years the remaining herds are said to have been [: ] holding their own.
Some writers have, from time to time, been impressed by the economic poten–
tialities of the vast arctic tundra and taiga. They have, no doubt, reasoned
that the untold millions of acres of arctic grassland that at one time supported
vast numbers of wild caribou as well as immense herds of musk oxen, if restocked
with domesticated reindeer or musk oxen could again be made productive, thereby
furnishing substantial sources of food for the meat-hungry world. At first
glance this line of reasoning would seem logical, for have we not here two
large, meat-producing animals that are admirably suited to life in the Arctic,
require no barns of stables to shelter them through the long arctic winter,
and are able to subsist, summer and winter, on native arctic vegetation that,
without cost or effort to the owners of the reindeer, reproduces itself year-in
and year-out? Unfortunately, experience has shown that the musk ox does not lend
itself readily to domestication, and that the raising of reindeer on a c [: ]
commercial scale is not yet practicable. The reason for this is not that the
reindeer is not suitable as a large-scale meat producer, not that the arctic
tundra vegetation will be feed it, but simply that there are not enough people
in the thinly populated Arctic who are willing and satisfied to live the life of
reindeer nomads; nor can the commercial raising the herding of reindeer, owing
to the high cost of marketing as well as to climatic and social disadvantages,
be made attractive enough economically to induce sufficient numbers of people
from elsewhere to go and live in the Arctic as reindeer herders. Nansen (16)

EA-PS. Porsild: Economic Botany

well realized this for, when speaking of the decline of the Samoyeds, he
said: “This decline of the nomads is all the more to be regretted, as they
alone with their [reindeer] culture are able to turn to account the immense
wastes of the tundra; the white race will never learn to do it.”
The most promising the economically practical approach to the problem of
utilization of the vast arctic and subarctic tundra and taiga appears then to
be the wise and careful administration of the remaining wildlife resources.
If given adequate protection against wanton slaughter and against needless
destruction of their natural range, caribou and musk oxen will both respond,
and in the end may be safely counted upon to provide a dependable and lasting
source of food the clothing for the sparsely populated Arctic.

EA-PS. Porsild: Economic Botany

Bibliography

1. Anonymous: Reindeer and Musk-ox . Report of the Royal Commission upon
the possibilities of reindeer and musk-ox industries in the
arctic and sub-arctic regions. Department of the Interior,
Ottawa, 1922.

2. Balzak, S. S., Vasyutin, V. F., and Feigin, Ya. G. Economic Geography of
the U.S.S.R., American Edition edited by Chauncy D. Harris, Mac–
millan Co., N.Y., 1949.

3. Banfield, A. W. F. “The Barren-Ground Caribou.” Can. Wildlife Service
Dept. Resources & Development, Ottawa, 1950. (Mimeogr.)

4. Berg, L. S. Natural Regions of the U.S.S.R. Macmillan Co., N.Y., 1950.

5. Colby, M. A Guide to Alaska . Am. GuideSeries. Macmillan Co., N.Y., 1939.

6. Druri, L. V. “Reindeer-pasture in the chukchee Anadyr district.” (In
Russian with English summary). Trans . Arctic Inst. vol. 62, pp.
105-124. Leningrad, 1936.

7. Hahn, E. Die Hausthiere u. ihre Beziehungen z. Wirtschaft des Menschen .
Leipzig, 1896.

8. Igoshina, K. N. “Contents of the rumen of reindeer in the snow period of
the year.” (In Russian). Publ . Arctic Inst. of the U.S.S.R., no.6,
pp. 63-72, Leningrad, 1936.

9. Isaachsen, H. “Lav som dyrefor.” Landbruksdepartementets Smaaskr . No.7,
pp. 1-10, Kristiania, 1017.

10. Jackson, Sheldon. “Report on introduction of domestic reindeer into Alaska.”
U.S. Senate Misc.Publ . No. 22, Washington, D.C., 1893.

11. ----. Report of the Commissioner of Education for the year 1903, Washington,
D.C., 1903.

12. Lantis, Margaret. “The reindeer industry in Alaska.” Arctic , vol.3, pp.
27-44, 1950.

13. Laufer, Berthold, “The reindeer and its domestication.” Mem . [: ] Am.Anthrop.
Association, vol.4, pp.91-150, 1917.

14. Mecking, Ludwig. “A regional geography of the Arctic and the Antarctic. The
Geography [: ] of the Polar Regions.” Am.Geogr.Soc. Spec.Publ . No.8, 1928

EA-PS. Porsild: Economic Botany

15. Mirov, N. T. “Notes on the domestication of reindeer,” Am. Anthropologist [: ],
n.s., vol.47, pp.393-408, 1945.

16. Nansen, Fr. Through Siberia the Land of the Future . Wm Heinemann, London,
1914.

17. Palmer, Lawrence J. “Progress of reindeer grazing investigations in
Alaska,” U.S. Dept. of Agric. Bull . 1423, Washington, D.C., 1926.

18. Palmer, Lawrence J. “Improved reindder handling,” U.S. Dept. Agric. Circular
No. [: ] 82, Washington, D.C., 1929.

19. ----. “Raising Reindeer in Alaska,” U.S. Dept. Agric. Misc. Publ . No.207,
Washington, D.C., 1934.

20. ----, and Hadwen, S. “Reindeer in Alaska,” U.S. Dept. of Agric. Bull . No.
1089, Washington, D.C., 1922.

21. Porsild, A. E. Reindeer Grazing in Northwest Canada . Report on an investi–
gation of pastoral possibilities in the area from the Alaska-Yukon
boundary to Coppermine River. Dept. Interior, Ottawa, 1929.

22. ----. “Rener of Eskimoer i Kanada.” Grønl. Selsk. Aaraskr , pp.1-24.
København, 1936.

23. ----. “Reindeer and caribou grazing in Canada.” Trans . N.Am. Wildlife conf.
vol.7, pp.381-91, 1942.

2 4 3 . ----. “The reindeer industry and the Canadian Eskimo.” Geogr.Journ . vol. 88,
pp.1-19, London, 1936.

24. ---. “Reindeer and caribou grazing in Canada.” Trans . N.Am. Wildlife Conf.
vol.7, pp.381-391, 1942.

25. ----. “Report on the reindeer and the Mackenzie Delta Reindeer Grazing Re–
serve” (manuscript), 1947.

26. Rink, H. Grønland geografisk of statistisk beskrevet , 2 vols., Kjøbenhavn, [: ]
1857.

27. Sdobnikov, V. M. “Relation between the reindeer and animal life of the
tundra and forest.” (In Russian.) Trans . Arctic Inst. vol.24,
pp.5-60, 1935.

28. Seton, E. T. The Arctic Prairies , London, 1912.

EA- PS. Porsild: Economic Botany

29. Sochava, V. B. “On the herbage food of the reindeer, as a source of miner–
al nutriment.” In The Soviet Reindeer Industry . (In Russian).
Publ . Arctic Inst. of the U.S.S.R. no.6, 1936.

30. Sosnovsky, G. P. “Ancient traces of animal husbandry in the Baikal region.”
(In Russian) Izv . Gosud. Akad. Istorii Mater. Cult. vol.100, 1933.

31. Stefansson, V. “The resources of the Arctic and the problem of their uti–
lization.” Problems of Polar Research, Am.Geogr. Spec.Publ ., no.7,
pp.209-34, 1928.

32. Tallgren, A.M. “Inner Asiatic and Siberian rock pictures.” Eurasia Septent .
Antiqua , vol.8, pp.174-210, Helsinki, 1933.

33. Voshchinin, V. P. “History, present policies and organization in the U.S.S.R.
Pioneer Settlement.” Am.Geogr.Soc. Spec. Publ . No.14, 1932.

34. Vassiliev, V. N. “The reindeer range in Anadyrland.” (In Russian with English
summary.) Trans . Arctic Inst., vol.62, pp.5-104, Leningrad, 1936.

A. E. Porsild

Utilization of Lichens in the Arctic and Subarctic

(EA-PS. George A. Llano)

UTILIZATION OF LICHENS IN THE ARCTIC & SUBARCTIC

CONTENTS
Page
Character of Range Lands 4
Norphology and Reproduction 8
Growth and Ecesis 11
Composition and Interspersion of Species 13
Components and Biochemistry 16
Food Value of Lichens 22
Industrial Uses 25
Possibilities for Future Research 29
Bibliography 32

EA-PS. Llano: Utilization of Lichens

LIST OF FIGURES
Page
Fig. 1. Usnic acid 20a
Fig. 2. Physcion 20a
Fig. 3. Evernic acid 20a
Fig. 4. Rhizocarpic acid 20a
Fig. 5. Epanorin 20a

EA-Plant Sciences
(George A. Llano)

UTILIZATION OF LICHENS IN THE ARCTIC AND SUBARCTIC *
The growth of human cultures in the temperate, subtropical, and tropical
regions of the world has been proportionate to man’s success in appropriating
and improving on the innumerable species of flowering plants associated with
his environment, aided by generally favorable climatic conditions and fertile
soils. In contrast, the Arctic and Subarctic offer a smaller, but not infre–
quently luxuriant, native flora of vascular plants lacking agricultural signif–
icance, and, in addition, an abundance of nonvascular plants.
To a great extent the vegetation of an area reflects the limitations of
the local environment, and this is well emphasized in the monotonous composi–
tion of the arctic coastal flora. The conditions of low winter temperatures,
a short growing period, low rate of precipitation, relatively high rate of
evaporation, and the acidity of soils brought about by both climate and un–
favorable microbiological development, all present special deterrents to a
normal expansion of agriculture. The presence of permafrost combined with low
relief so impedes the runoff of surface waters that in spite of the low rain–
fall great areas are covered with lakes or marshy ponds, overflowing their
banks at high-water periods. The effect of these factors favors the development
of a rich flora of bryophytes and lichens, which are often the dominant ecolog–
ical forms. Of the two groups, the lichens offer the better possibilities for 1

EA-PS. Llano: Utilization of Lichens

exploitation as forage plants in enormous areas that are absolutely useless
for other purposes.
The northern submarginal lands have never supported more than a sparse,
m n omadic native population, and this has been composed of hunters or herdsmen
indirectly dependent for survival upon the cryptogamic elements of the sea
and land flora. The terrestrial vegetation of the Arctic and Subarctic can
support large numbers of wild herbivores, of which the genus Rangifer has the
widest circumpolar distribution. Seton (23) has stated, in regard to the great
herds of caribou, that their numbers were never proportionate to the available
food supply
The symbiotic relationship between Asiatic man and the gregarious land
mammals, such as the sheep and horse, is bound up with the maintenance of
the grass steppes; reindeer domestication is a contribution of the arctic and
subarctic lichen steppes. It is a culture which still persists across Asia,
extending so u th to latitude 50° N. With the introduction of reindeer into Alaska
for the support of the native Eskimo population, and the emphasis on Siberian
reindeer increment, reindeer husbandry has attained the rank of an industry
extending almost continuously around the polar regions.
The genus Rangifer is characterized by strong migratory habits. The forces
which induce these movements, covering hundreds of miles, are not completely
understood, but they include the tormenting attacks of insects, weather condi–
tions, and a “natural rotation” to new pastures. This apparently aimless wan–
dering resolves itself into two distinct phases; these are the summer and
winter feeding periods, with a stationary period during which calving takes
place. These phases of the migratory cycle are reduced in reindeer “stock” to
a milder expression of movement in domestication. Whether the industry adheres

EA-PS. Llano: Utilization of Lichens

to close-herding practices or to the newer permanent ranching basis, the
animals must still have access to enough grazing land to supply the desired
seasonal fodder, which cannot be cultured or harvested sufficiently to meet
known forage needs,
Although reindeer feed and fatten on grasses, sedges, and variable amounts
of other herbage in summer, supplemented often in greater bulk by browsing on
the tender portions of willows, birch, and other shrubs, lichens may also be
taken, particularly when they are moist; and then in the fall and throughout
the winter, the animals exhibit a marked preference for lichens. Asthis period
composes the greater part of the year, the extent, quality, and quantity of the [: ]
lichen pastures are of paramount importance. Data based on stomach analyses
and field observations show conditions. Thus on Novaya Zemlya,
where heavy snows had made lichen ranges inaccessible, reindeer were able to
subsist on dried summer plants and browse. Under similar conditions on [: ]
Nunivak Island, of Alaska, in the Bering Sea, a semidomesticated reindeer herd
was severly reduced by starvation. In Fennoscandia, when icing or flen condi–
tions prevail, the Lapp herders move their animals into ice-free pastures or
into the forest where they may feed on tree-growing lichens, often eating the
bark of Betula. On the Malozemelskaia tundra, it has been reported that the
amounts of lichens found in the rumen of reindeer range from 3 to 40% of all
food taken. In the case of fawns during the summer and autumn, lichens amount
to 25 to 30% of the total food; adult reindeer average about 26% lichens and
35 to 40% Salix and Betula browse during the autumn.
Reproductive potentialities of domesticated reindeer have proved to be

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high; estimates of the theoretical population that can be carried on circum–
polar grazing lands have ranged up to 100,000,000. However, reindeer
increase, even with progressive government assistance, have failed to achieve
anything near the theoretical maximum; in Alaska, herds have been sever e ly
reduced from the early phenomenal numbers. The industry is best with many
overlapping and interplaying factors which, if disregarded, will reflect the
highly speculative aspects of any individual undertaking. These include the
decimating effects of predators, insects, disease, and weather; the willing–
ness and adaptability of Eskimos to accept a new mode of livelihood, neces–
sitating skills in herding and range management; and the problem of finding
market outlets for meat, hides, and furs. Reindeer management, like every
other form of land cropping, is applied ecology. Therefore, proper evaluation
of the vegetational units for determining carrying capacity of a range is neces–
sary for the successful application of range-management techniques, details of
which are partly available from published floristic, ecological, and phytosociolog–
ical studies of some circumpolar areas. This article concerns only those factors
that influence the lichen ranges.
Character of Range Lands
The limits of the latent northern pastures extend from the shores of the
Arctic Sea inland to the forests, which project as broad, undulating belts
across the Northern Hemisphere lands; the northernmost, spare, coniferous
timberlands have been designated as “taiga” in Siberia, where they extend north
along river valleys to the most northern limits on the Khatanga. Within these
boundaries there are innumerable subdivisions of plant species which reflect
the operative forces of climatic, edaphic, orographic, and biotic factors that

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serve to characterize the plant habitat. These, for the purposes of this
study, may be generalized into several broad types of land cover.
The rolling, treeless, marshy plains along the arctic coast support,
primarily, a rich sedge-grass association; lichens are mere insignificant
associates. The area may be best classified as summer pasture. On the
arctic slope of Alaska, the coastal region merges into the interior uplands,
which, though predominantly sedge-grass, have a better representation of
herbaceous plants and prostrate or dwarf shrubs, particularly along protected
stream depressions. The lichen flora, though more prevalent, is thin and
dispersed. The area is used for winter feeding although it is poor in com–
parison with the best lichen fields — espe c ially the Finmark pastures of
northern Norway. The north slope of the Brooks Range varies from mountain
meadows rich in gras s es and succulent herbs to other which maintain carpet-like
associations of lichens. The mountain slopes, particularly the scress and
talus areas, have well-developed lichen association, though these may be
modified according to such factors as substrate and expo [: ]sure.
The uplands of the Canadian interior are more irregular than northern
Alaska and consequently are more difficult to describe in broad generaliza–
tions. Where timber and brush are absent, sedge-grass associations or natural
grasslands are often present. The uplands are interspersed with open or
short-timbered muskeg tracts set in the vaster northern plains, which often
support heathy shrubs growing out of a thick, well developed carpet of lichens
and mosses. The lichen carpet extends through the scattered timber frontal
to the coniferous forest boundaries and into the open fo [: ]rest edge. The short
timber and low branches of muskegs, scrub, and coniferous forest afford fine

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subst r ates for many fodder-valuable lichens, so th a t lichen feed is available
from both ground and tree strata.
The Asiatic arctic lands have been partially reported on by recent
U.S.S.R. expeditions studying means of expanding the reindeer industry. In
the Yakutia region, grazing areas are given as belonging to 24 associated
vegetational types; the Anadyr reindeer pastures are classified into 11 prin–
cipal types; while the Taimyr Peninsula is stated to consist of 75% tundra
and 25% forest. The orographical variety of the northern Asiatic hinterland
and the northward penetration of forests makes for a greater discontinuity
of vegetational zones. The tundra belt is narrowest in the west, where it
contacts the “wet taiga” of evergreen con i fers, increasing in width eastward,
where it merges into the uplands and highlands of the interior. Open tundra
exists along gentle slopes and forelands of mountains or in poorly drained
lowland areas characterized by sedge and Sphagnum bogs, interspersed with
dwarf birch; this merges into a transitional zone of dwarf birch and spruce,
which in eastern Siberia adjoins the “dry taiga” of deciduous and coniferous
forests. Intervening summits of higher lands assume tundra or mountain-meadow
qualities.
Throughout this area of nother Asia, lichens are a common component of
the flora. The yearly yield of lichen fodder in the Yakutia region is estimated
at 3.7 million tons, with a basic available total of 55 million tons; herbs and
shrubs are estimated at 13 million tons. In the Far Eastern Region, the herbace–
[: ]ous fodder is said to be deficient, with lichens predominating; while the Taimyr
Peninsula is reported to be adequate only in the matter of pastures. In the
Anadyr area, summer and winter pastures are available, but the full utilization

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of these pastures is severely restricted by the presence of a complex swamp
system and by the intervening taiga, which impede normal travel. The inability
completely to tuilize available pasturage is thus due to the disproportionate
ratio between summer and winter pastures, the complications of a topography
disadvantageous to reindeer herding, and the existence of “negative” areas,
including frost-shattered rock uplands, strong polygon shingle formations,
and bare stone ridges, which support an indifferent herbaceous vegetation
and only a thin cover of lichens, these being primarily the smaller rock
species.
Reindeer husbandry in northern scandinavia is strongly competitive for
grazing areas because of the pressure exerted by agricultural and forest
interests and the increased urbanization brought about by mining developments.
The closing of Finnish and Soviet Lapp pastures has aggravated the grazing
problem and resulted in overcongestion of the tracts available in Swedish
Lapland. The Swedish Government has realized the essenti a l nature of this
industry for the independent maintenance of the Lapp culture, and has tried to
comply with the hereditary needs of the Lapp herders. Some relief has been af–
forded by the use of Norwegian fjeld lichen pastures, which are controlled by
restrictive laws invoked jointly by the two governments. These laws regulate
the routes to be used, the time for grazing, the number of reindeer allotted
per area, time of entrance, and compensation for damages. The greatest fric–
tion occurs between the Lapp herders using fields which are also hand-harvested
to supplement domestic animal feed. Nevertheless, some of t h e lichen fields
of Fi [: ] n mark represent the best types of lichen pasturage available, reflecting
the rotational use of the land and a careful analysis of its carrying capacity.

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In those sections of western Eurasia where the coniferous belt approaches
close to the coast, the reindeer culture has developed into two distinct types
of herding practice — that of the mountain Lapps, who hold their herds in
summer on the treeless mountains and in autumn move into forest districts of
eastern Scandinavia, and that of the forest Lapps of Sweden, Finland, and
Soviet Russia, who stay in the lichen forests all the year around. This
[: ]divurgence has not only evolved an alternation of range practices but also
has resulted in the domestication of at least two different races of animals,
of which the forest type is cited as the heavier and larger, more capable of
being used as a mount or for draft purposes. Herding is feasible in the
parklike pine and fir as well as in birch and mixed birch [: ] fir forests, owing
to the well-developed strata of lichens on the ground and on the lower branches
of the trees.
Morphology and Reproduction
Lichens bear so little resemblance to other forms of plant life that it
is difficult to convey to the layman an understanding of their unique struc–
tures. Consequently, they are frequently considered to be a kind of moss.
Moreover, those species which are important to reindeer management have been,
unfortunately, called “reindeer moss,” which is a source of added confusion.
The term, as generally applied, covers numerous species, and it is important that
they be identified as species or at least as genera to give meaning to the art
of reindeer land management.
The lichen structure lacks stems, leaves, roots, flowers, and true see d s;
it constitutes a specialized noncellular system of irregular limitations that
is correctly described as a thallus. Lichenologists have classified thallus

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forms into three types: ( 1 ) the crustaceous lichen thallus which is usually
of small size, frequently lacking an observable thallus, often closely ad–
hering to bark, soil, and rocks, its small size and habit canceling its value
as a grazing plant; ( 2 ) the foliose or leafy lichen thallus, which can grow
to 6 or more inches (15 or more centimeters) in diameter, forming irregular
succulent structures over the substrate; and ( 3 ) the erect or hanging, more
or less cylindrical thallus which may develop on the ground or on trees and
rocks and is aptly described as “bushy” or “fruticose.” The grazing value
of northern pastures in winter is primarily due to the large species of foliose [: ]
and densely growing fruticose lichens.
Microscopically, the internal anatomy of lichens differs markedly from
all other forms of plant life. It consists of a tightly woven web of fungal
threads or hyphae through which may be scattered or oriented innumerable bright–
green, yellow-green, or blue-green algal cells. It is the presence of algae
in lichens that partly gives them their dominant coloring, although when dry
some may appear nondescript. The symbiotic relationship exhibited by this
dual organism is dependent upon the specialized and apparently balanced func–
tions carried out by each partner. The chlorophyll-bearing algae can syn–
thesize sugars and a special form of carbohydrate, lichenin, which can [: ] be
used by the fungus for its own growth; the algae would be easily desiccated
but for the water-absorbing and water-retaining powers of the innumerable
hollow hyphal threads of the fungal partner. As a result of this mutual meta–
bolism, lichens can achi e ve success in environments too barren to support other
forms of plant life, or where an independent existence of either [: ] alga or
fungus would be impracticable.
The methods of reproduction of lichens give a better understanding of

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the relationship of the alga and fungus. Lichens may reproduce by frag–
mentation, that is, a broken part of the thallus may, under suitable condi–
tions, reproduce itself; or they may develop spherical bodies (soredia)
containing fungus threads surrounding a few algal cells which are liberated
as a powdery mass that is easily distributed by the wind. Lastly, the fungal
part may form spores in specialized closed (perthecia) or open (apothecia)
structures after the usual manner of fungi, without the participation of the
algal associates until after the spore germinates. It is because of this
method of sexual reproduction and the fact that the fungus enmeshes the
alga that the former is considered to be the dominant participant. The
ability of these organisms to reproduce themselves as a unit, with specific
morphologic, taxonomic, ecologic, and physiologic properties, makes it con–
venient to treat them as homogeneous units comparable to the true mosses,
hepatics, fungi, or algae.
Lichens are perennial plants of slow growth and insignificant size,
whose importance as forage plants rests partly upon their extensive accumula–
tion under favorable conditions. Their activity is dependent upon the avail–
able air moisture — whether as fog, rain, or snow — even in small quantities.
Under suitable conditions, lichens are active even at low temperatures, par–
tcularly under a covering of snow, so that the vegetative period may be ex–
tended into early spring and late autumn. As heliophytes, they tend to attain
their best development on the southern exposure of mountains, under thin forest
cover, and on the open tundra. On low tundra, their development is often seri–
ously retarded by flooding and by competition from the grass-sedge association.
The chemical composition and thus the edible qualities of lichens may well be
dependent upon the availability of effective illumination.

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Foliose species grow peripherally; the primary growing portion may
persist or be partially decomposed. Fruticose forms grow apically, with a
continuous decomposition of the basal portion in the case of terrestrial
species; in a sense they possess the power of unlimited growth, although
they rarely exceed 6 to 8 inches (15 to 20 cm.) in height. The cushion
formation of certain fruticose groups, such as Cetraria , Cladonia , and
Thamnolia , on forest soils or open country, is best observed in northern or
alpine regions; such cusion-like formations are often induced by wind action,
the plants being then less sensitive not only to wind but also to variations
of temperature and to intense insolation. The viable part of the fruticose
forms is the upper half or third of the plant, and it is from this portion
that regeneration proceeds.
Growth and Ecesis
Exact kn o wledge of the yearly growth of lichens if basic to a proper eval–
uation of the available winter fodder and consequently to determination of the
carrying capacity of any type of range. Although there are in the literature
scattered casual observations on the growth of lichens, none is of sufficient
exactness to enable computation of the yearly or average growth rates of any
one species in any given latitude. General estimates, based on common knowledge
of past grazing experience in various parts of the Arctic and Subarctic, are,
however, available. It has been shown that fruticose lichens trimmed to a
depth of 1 to 2 centimeters quickly recover; cutting to 5 centimeters is more
detrimental to recovery. Moderate feeding by reindeer — that is, light crop–
ping — permits recovery in a period of from 4 to 5 years. This recovery period
is quicker on young, lightly cropped lichen stands than on the more mature,

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thick cushion stands, which may often be cropped down to the inactive basal
portion.
In judging the range condition of any previously pastured area, it is
well to consider that complete recovery may not be desirable as it would in–
volve forage loss through excessive basal decomposition. In short, moderate
pasturing with a short interval of rotation may be more advantageous than
prolonged abandonment. Successful lichen-range management is thus largely
dependent upon the harder’s ability to evaluate the cropped lichen stand in
terms of the time it will take to recover to a usable state.
Lichens when dry are exceedingly brittle, although they never become
completely desiccated. The use of lichen pastures during dry periods, eith e r
for grazing or for migratory purposes, may result in excessive fragmentation
of the thalli and a serious pasturage loss. This can be further aggravated
if it occurs during the insect season, when the animals are accustomed to make
short, rapid runs to shake off their tormentors. Under optimum conditions of
range use, that is, when the lichen stands are moist or covered with a protective
blanket of snow, there is undoubtedly a normal amount of fragmentation, which is
an important factor in the regeneration of new individuals. Lasting and in–
tense gr [: ] a zing accompanied by excessive trampling in areas of mixed sedge-grass
and lichen forage can result in the total replacement of the lichen association
by grasses and allied higher plants. Trampling of lichens is thus a major item
to observe in estimating available pasturage.
Damage to lichen stands can also be brought about through the destructive
activity of rodents, which during peak years in their cycles may be present in
excessive numbers, comp leting directly or indirectly by using lichens for food or

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by destroying them in their burrowing activities. Such rodents as Lemmus , Dicro
stonyx , Clethrionomys , Arvicola , Microtus , and Lepus have been reported to exert
a detrimental influence on the lichen pasturage of the Bolshezemelsk and Maloze–
melsk tundra of Siberia; a similar phenomenon was noted around Wainwright and
Anaktuvuk Pass of the arctic slope of Alaska. This effect is most noticeable
in spring, especially where surface waters wash plant detritus into windrows
along stream banks or in depressions.
Fire is particularly destructive to lichen fields, frequently resulting
in the total loss of available pasturage for many years. The effect of fire
on reindeer pasturage in Alaska has been most noticeable in the northwest area
around mining camps. Gaps made in l[]ichen pasture vegetation in Norway by fire
fifty or more year old could be readily traced by the comparison of the original
lichen cover with the adventitious and often less desirable species (including
mosses) which may be absent or have limited distribution in the older flora.
Composition and Interspersion of Species
In appraising the comparative worth of winter pastures, the seasonal need
with a possible alternative is the main prerequisite. With this must be consid–
ered the quality and quantity of the growing feed available, the composition of
the flora, the residestance of the pastures to herding, their regrowth rates, and
the distribution of the plant species. The quantity of lichen feed available is
not [: ] determinably by the total numbers of lichen species which may be
recorded as composing the lofra of a given region. The lichen flora of land
areas ranged around the pole may vary from 300 to 700 species, belonging to about
100 genera. Some of these may be locally abundant, others rare; many are rela–
tively unimportant because of their small size or development in inaccessible places.

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The lichens that are important in the composition of reindeer ranges number
about 50 species belonging to some 20 genera, of which 15 species are the
most desirable. The lichen species most fed upon by reindeer belong to the
genera Cladonia , Sterocaulon , Sphaerophorus , Thamnolia , Parmelia , Cornicularia ,
Peltigera , Dufourea , Umbilicaria , Alectoria , and Usnea .
[: Of]Of the terrestrial lichens, the Cladonia are the most important because
of their size, close-growing characteristics, and wide distribution in circum–
polar lands. Of the innumerable species described, the members of the Cladina
group (such as Cladonia rangiferina , C. alpestris , C. sylvatica , and C. impexa )
are characterized by the strong development of the secondary thallus or podetia,
which usually bears the fruiting structure, and by the absence of a primary
thallus; in the Cladina group, however, the development of apothecia, which
usually terminates apical growth, is rare, and apical growth is more or less
continuous while regeneration is dependent upon fragmentation. The species form
cushions or carpetlike mats of closely pressed, richly branched podetia, growing
best in area of definite snow cover and, when damp, are able to withstand con–
siderable trampling; they exhibit a wide tolerance to differences in the substra–
tum, compete well with (and even prevent the normal development of) vascular plant
seedlings, and can grow in a considerable variety of habitats but show slow
recovery after fire. Cladonia amaurocraea , C. gr [: ] a cilis , C. uncialis , C. verti
cillata , and other species are also taken by reindeer, but generally lack the
close-growing habit that is so valuable for forage plants.
The Cetrarias are equally important as they fulfill the quantity require–
ments necessary for forage plants. Although their primary thallus never attains
the podetial size of the Cladina group, they exhibit considerable tolerance to

EA-PS. Llano: Utilization of Lichens

moist situations and thereby give range value to otherwise useless Sphagnum
or other predominantly bryophytic communities. Cetraria hiascens ( C. delisei )
is frequently found in shallow pools, cold bogs, and other wet areas; C. cucullata
and C. nivalis appear in both wet and dryish areas, having their best development
on slopes that are subject to snow cover. All the Cetrarias named and terres–
trial lichens characteristic of open range, with wide circumpolar distributions.
Alectorias compose an important part of the ground- and tree-strata lichen
floras. The terrestrial species, Alectoria ochroleuca , A. nitidula , A. nigricans ,
and A. nidulifera , can be locally dominant [: ] on the tundra or on high mountain meadows,
comp e ting well with higher plants. Their best development is among the short–
growing mosses of driver habitats, such as occur on ledges or rocks or on talus
slopes. Alectoria ochroleuca is a prominent lichen of this last habitat. A. sar
mentosa , A. fremontii , and some of the smaller, darker Alectorias are found hang–
ing from branches of low scrub and are strong lichen elements of the coniferous
forest belt. Usneaceae are present in forested areas; they are in habit much
like the Alectorias, and thus the two may be frequently confused.
Stereocaulon , Sphaerophorus , Thamnolia , Cornicularia , and to a more limited
extent Dufourea , although generally found throughout the circumpolar area never
attain the close-growing habit of the Cladoniaceae. These are all terrestrial
arctic forms with generally short, erect, fruticose thalli.
Of the foliose genera, Parmelia , Peltigera , and Umbilicaria are ubiquitous
northern lichens with relatively large, flat thalli. Umbilicaria grows best on
acidic rock strata — often in such numbers as to give a mountainside a definite
“shadow” coloring effect. Peltigera is common over soil, while Parmelia occurs
over both rocks and soil, adhering closely to the substratum, although it has
been observed that reindeer lick it off.

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The lichen diet of reindeer, as has been proved in the case of other
game-animal foods, may follow a more or less definite sequence. The lichen
species taken may be dependent upon their relative abundance or availability
or they [: ] may follow a descending order of palatability, with the possibility
that seasonal changes and physiological needs may be underlying criteria. If
the staple lichen or other normal fodder is exhausted, reindeer are known to
fill up or “stuff” themselves on plants for which they normally show little
selectivity, including nonnutritive mosses. In Alaska, Nephroma arcticum , a com–
con circumpolar lichen, has been cited as a reindeer lichen; the Lapp herders
of Schandinavia, who have evolved a rich vocabulary of terms for lichens used
by reindeer, and who clearly differentiate between true mosses and lichens,
state that Nephroma is never eaten, or is taken only in the absence of other
foods.
Lichens not only serve to provide ordinary sustenance during the long
and critical winter feeding period, but they are known to be a source of vitamins
and, at a time when water can only be supplied by eating snow, may be the sole
available source from which mineral deficiencies can be made good.
Lichen Components and Biochemistry
The nutritive values of lichen fodder lie in the presence of substances
synthesized in the thallus, which vary in quantity among different species and
even in any one species under dissimilar habitat backgrounds. These include car–
bohydrates, acids, auxiliary vitamins, and minerals; proteins and fats are
present in small amountgs lichens.
The important foodstuffs that compose the greater part of the thallus
are held as reserve carbohydrate; these foodstuffs are cellulose-like

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polysaccharides of the hexose type, sometimes described as polyglucides, of which
lichenin is the most prevalent form. Isolichenin, a starchlike polysaccharide,
occurs along with lichenin and differs from it only in being soluble in cold
water, in giving a blue color with iodine, and in yielding maltose on enzymatic
hydrolysis. Lichenin is soluble in hot water, giving a colloidal solution, and
is easily hydrolyzed to yield D-glucose. Cellulose is less commonly found in
lichens but in the Cladonias it amounts to 6 to 10% of the dry weight of the
plant; this is water-insoluble and less readily hydrolyzed thank lichenin.
Some licen species are characterized by more specialized forms of soluble
carbohydrates, but without necessarily precluding the presence of lichenin; all
posses the common property of yielding glucose when treated with dilute acids.
Thus pustulin replaces lichenin in Umbilicaria which, unlike most lichens, requires
an external source of organic matter. Peltigerin is common to the Peltigeraceae;
while zeorin is found in Anaptychia and Lecanoraceae. Simple reducing sugars are
rare in lichens; erythritol and mannitol have been reported in considerable
amounts in a few tropical species.
Through the action of primarily of aerobic and anaerobic bacteria of the
digestive system, rather than of enzymes, the polyglucides are hydrolyzed to
simple sugars. Lichenin and inulin, though once thought to be peculiar to lichens,
are also present in some of the higher plants. The former has been isolated [: ] from
oats, while the latter occurs in chicory root, dahlia bulbs, dandelion roots, sweet
potatoes, and Jerusalem artichokes. Diachkov and Kursanov (5) have reported that
the total carbohydrate present in Cetraria nivalis , C. islandica , Cladonia al
pestris , C. mitis , C.deformis, and Alectoria ochroleuca composes 80 to 85% of
the dry weight of the plant. In Peltigera and Sterocaulon paschale it is 48 to 72%
A marked contrast was indicated in the formation of lichenin which in Cetraria

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islandica , C. nivalis, and Alectoria ochroleuca made up 45 [: ] to 50% of the water–
soluble carbohydrates; in the rest of the lichens tested it amounted to less
than 5%. The residue of the carbohydrates was in the form of water-insoluble
hemicelluloses, celluloses and pentosans. Cetraria islandica and C. nivalis differed phy–
siologically in human digestibility tests; in the former, 45 to 50% of the
lichenin was found to be digestible, while the latter provoked intestinal
disturbances which necessitated termination of the experiments. White mice
have been reported to utilize about 70% of available lichenin.
The physiological effect of the lichen acids on the feeding habits of
wild herbivores does not appear to be harmful, although it may help to determine
preference in selection of lichen species. At least two species are reported to
be poisonous and have been used in the control of wolves: Evernia ( Letharia )
vulpine , containing vulpinic acid, and Cetraria pinastri, containing pinastrinic
acid, are both bright-yellow species with wide circumboreal distributions. That
the former lichen is not detrimental to herbivores has appeared from field ob–
servations by W. O. Douglas (6) who noted some fifty elk feeding on Evernia vul
pina , Alectoria sarmentosa , and A. fremontii in the mountain region of Grosse
Point, Washington.
R. S. Palmer has noted ( in litt .) that Usnea is one of the best foods for
baiting traps for northern white-tailed deer, that on overbrowsed ranges Usnea
was completely exhausted as high as the animals could obtain it, and that, when
trees are being felled, tame deer will come running to feed on this lichen.
Cowan (3) states that 15% of the annual diet of Odocoileus hemionus columbianus
on Vancouver Island is composed of Usneaceae. In some of these instances, lichen
feeding occurred late in the winter on ovefgrazed or overbrowzed areas or under
unfavorable climatic conditions. However, such lichen feeding may also indicate

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a nutritional deficiency or, as palmer suggested, a physiological need for
some “salt.” Certain minerals have been noted in l [: ] i chens which ordinarily
do not occur in plants. Thus parmelia molliuscula has been reported in the
western United States to contain selenium in sufficient quantities to affect
seriously sheep and cattle. Beryllium has been observed in P. saxatilis and
Xanthoria parietina ; chlorine has been detected in Evernia furfuracea . The
normal development of the colon bacterial flora of ruminants appears to require
the presence of at least trace amounts of certain specific minerals. The need
for investigating this aspect of lichen nutrition becomes more apparent in the
light of the normal selectivity shown by species of Rangifer , the seasonal
selectivity exhibited by members of the Cervidae, and the temporary adaptation
of cattle to lichen fodder. Correlated with these phenomena is the generally
accepted fact that different diets will produce marked and even radical dif–
ferences in the intestinal flora of different species of animals.
The [: ] bright coloring of many lichen species is partly due to the
effects of acidic compounds which occur as crystals or minute granules in greater
or lesser abundance on the outer surface of the hyphal filaments. Some 200
lichen acids have been described, whose chemical structures reveal a unique
pattern unknown in other plants, and which are pressumed to result from the
symbiotic metabolism of alga and fungus — primarily the latter. Dr. Robert L.
Frank of the Department of Chemistry, University of Illinois, has recently com–
municated ( in litt .) the following facts concerning the nature of these acidic
compounds. Chemically, the lichen acids cannot be placed in a single group
because there is great variation in their structure. Some are true carboxylic
acids, others are acidic by virtue of being phenols or enols. Examples are usnic
acid and physcion; the former (Fig. 1) occurs in seventy or more lichens while

EA-PS. Llano: Utilization of Lichens

the latter (Fig. 2) is common to Caloplaca elegans and other species. Evernic
acid is an example of a depside (Fig. 3). Dr. Frank further pointed out that:
“Of the lichen acids, only a few contain nitrogen. The exact environment
of the nitrogen, as far as I know, is known only in the two pigments rhizo [: ]
carpic acid and epanorin. In these the nitrogen is present in amino acid
molecules, an interesting fact from the standpoint … of the nutritional
aspects of lichens. In connection with the preference of the Cervidae for
certain lichens during the winter, [the] assumption that they may crave for
essential metabolite present in the lichens is certainly an interesting and
logical one. Some additional speculation follows. The lichen acids and pig–
ments illustrated are probably not of nutritional importance for the Cervidae,
as none of their structures are at all similar to presently know metabolites
(with the exceptions of the amino acid derivatives, rhizocarpic acid and
epanorin). It would seem more fruitful to look for the necessary metabolites
among the minerals, vitamins, or amino acids. Vitamin D can perhaps be placed
in the same category [as vitamin A] because a good part of an animal’s Vitamin D
is obtained by ultraviolet irradiation of inactive chemical precursors on the
body surfaces … My only other thought is to reiterate the interesting fact
that only two nitrogen-containing lichen pigments, rhizocarpic acid [Fig. 4]
and epanorin [Fig. 5], are pulvinic acid derivatives of the amino acids
[: ]L-pheylalanine and L-leucine, respectively. Both these amino acids are es–
sential dietary constituents of rats and man, and are not synthesized in the
body. The fact that nitrogen-containing pigments having other amino acids tied
up with pulvinic acid, as in rhizocarpic acid and epanorin, have not been reported
suggests that some few of the amino acids such as phenylalanine and leucine may
predominate in lichen flora (and may be present in substantial amounts). Fig 1 Fig 2 Fig 3. fig 4 fig 5

EA-PS. Llano: Utilization of Lichens

Such amino acids could by chance be the needed materials causing the Cer–
vidae to prefer lichen food at certain periods.”
The amount of acid compounds in lichens varies from 2 to 3% to as much
as 25% in some of the economically important dye lichens. Lichen acids
are of taxonomic value in separating certain species or in relating larger
groups by their known reaction to specific chemicals. The distinctive
bitter taste of some Cladonias, due to the presence of fumarprotocetraric
acid, is also of use in the determination of these species.
The vitamin content of several species of fruticose and foliose lichens
eaten by reindeer has been reported upon by L. F. Palmer and his associates
(16). The results indicated that these species yield moderately high di–
gesti bi lity coefficients in reindeer. The selected plants were arbitrarily
divided into short-growth types occurring on dry sites and tall-growth types
occurring on moist sites, and were fed to rats, mixed with alfalfa and oats
or as a pure diet. The rats would not tolerate large amounts of the tall
lichens at concentrations greater than 10% in the all-lichen diets; the
species included Cetraria islandica , C. cucullata , Cladonia alpestris ,
C. sylvestris , C.gracilis , C. amaurocraea , and C. sylvatica . A vitamin-A
response was obtained from this group, although it could not be demonstrated;
vitamin-D response was superior to that shown in short-growth forms. Short–
growth forms were well tolerated at the levels fed; the species included
Alectoria nigricans , A. ochroleuca , Cetraria nivalis , C. hiascens , Cladonia
bellidiflora , C. gracilis , Lobaria linita , Stereocaulon tomentosum , Tham
nolia vermicularis , Nephroma arcticum Parmelia physodes , and Peltigera sp.
Vitamin-B complex was absent in north short- and tall-growth forms. Ergosterol

EA-PS. Llano: Utilization of Lichens

has been reported to have been isolated from Peltigera canina and Cladonia
rangiferina
— the latter, in September, showing only traces of vitamin D.
Food Value of Lichens
It is possible to accustom reindeer to hay and grain by mixing, for a
time, lichens with such feed. Cattle likewise can be [: ] accustomed to lichen
feed by mixing the required amounts with their normal food, or by salting
lichen feed. The bitter taste may be partly removed by boiling or soaking in
water for 24 hours; by the addition of potassium carbonate or weak alkali
solutions, the insoluble lichen acids may be changed to soluble salts. One
kilogram of Cladonia rangiferina is considered equal to a third of a kilogram
of poor fodder or early grass; chemical analysis of this species showed a
composition of 61% lichenin and 1 to 5% proteins, the rest being hemicellu–
loses.
The use of liche n s as food for swine has been highly recommended in Norway,
and it has proved that young pigs thrive better on a combination of “reindeer
lichen” and ordinary feed than on the latter alone. Cattle and pigs are not
grazed but hand-fed with lichen feed, which necessitates harvesting.
It is in the region in Scandinavia where lichen harvesting is conducted
that most of the friction between the agricultural and nomadic elements of the
population arises. The lichens are harvested either with hand rakes or by
hand; in either case the living portion is w h olly removed, necessitating access
to fresh fields for continuous supplies. Reindeer, under proper herding prac–
tices, normally crop about 10% of the available lichen fodder, and even under
more rigorous conditions the range would again be available in 4 to 6 years"
time; harvesting of lichen fields with implements lengthens this period before

EA-PS. Llano: Utilization of Lichens

availability to 25 to 30 years. However, the whole area is never completely
raked up but rather is treated as a series of lanes, the lichens being piled
into small heaps around a branch of birch to facilitate handling. When wet,
the lichens may be compressed in hand presses without any loss, or the lichen
sward when frozen may be cut into blocks for ease in transportation and thus
stored in sheds or under straw. Pure lichen diets are rarely fed to cattle,
but the lichens are used to supplement the more expensively grown feeds.
Ordinarily three to five sledge loads (300 to 600 kilos pre sledge) are col–
lected per cow, one man being able to rake from 300 to 400 kilos in a day.
Lichen fodder is also harvested for feeding reindeer, as in the Petsamo-Kola
region of the U.S.S.R., where reindeer are restricted to enclosures.
The use of lichens for food by man and beasts extends into antiquity,
as indicated by prehistoric remains found near Lake Constance, Switzerland.
Within historic times, lichens have commonly been supplanted by cultivated
grains. Still, in times of poor crops or stravation, their use has been re–
vived. Recent studies of Norse remains in Greenland revealed, in comparison
with contemp o rary Eskimo skulls, an abnormal wear of the teeth due to tritura–
tion — suggesting that during the terminal stage of the settlement the in–
habitants may have been forced to subsist on lichens and/or seaweeds.
Recipes for the preparation of lichen foods are available from early ac–
counts. Cetraria islandica was the most popular of all lichens, and as late
as 1836 there was published an account giving twenty ways of preparing it for
human use — as bread, mixed with potatoes or grain, or as gruel, porridge,
jellies, soups, and in salads. Undoubtedly the presence of lichen acids, which
are known to repel some insects, was an important ingredient of lichens recom–
mended for use in ship’s bread — which was accordingly less subject to weevil
infestation and was nonfriable.

EA-PS. Llano: Utilization of Lichens

In all cases, the preparation of lichens for human use entailed boiling
with or without weak alkalies to remove lichen acids, followed by rinsing
in clear water; the resulting mass was than oven-dried and placed in closed
containers until needed. It had good storage qualities of at least one year.
In making bread, this material was mixed with one-fourth grain, producing a
meal from which could be baked a strong bread having a fair taste, although
if the lichen acids had not been thoroughly removed it was said to leave a
sense of heat on the tongue. When boiled until it became thick, it could be
made into a porridge which jelled on cooling; it was eaten with salt and
with or without milk.
The emergency use of Umbilicaria species (rock tripes), for example, by
French-Canadian trappers of colonial America, and by members of the Franklin
expedition in the Arctic, has led to the popular belief that here is a manna
readily available to northern travelers who run short of provisions. This
genus has a distribution restricted to acidic rock substrata throughout its
circumpolar range. Dry thalli are leathery, requiring persistent mastication;
boiling in water increases their palatability. The polyglucide pustulin is
capable of being 51% hydrolyzed with sulfuric acid; but its digestibility
coefficient appears to be lower in human metabolism than that of lichenin
or isolichenin, although one species of Umbilicaria is eaten in Japan as a
delicacy. Cetraria islandica might be the more logical choise as an emergency
food, for it has a wider distribution in the North. At all events, the em–
phasis for arctic survival should be placed more upon birds, mammals, and fish
as a source of food than upon plants generally.
It is striking to note that the culture of primitive arctic and subarctic
peoples who suffered periodically from poor hunting years lacks any suggestion

EA-PS. Llano: Utilization of Lichens

for the of lichens during survival periods. In fact, the only verified
utilizations of lichens by the Alaskan Eskimos, which still prevail today,
are as a fire material and in hunting the northern hoary marmot ( Marmota ).
In the former case, Cetraria richardsonii , a loose, tumbleweed-like and coarse
lichen, is gathered for priming wood fires, and in the latter, the Eskimos
locate the burrows of marmots on mountain slopes by spotting the bright-yellow
lichen Xanthoria . The growth of this nitophilous lichen is materially aided
through the marmot’s habit of evacuating and wetting in a restricted spot
close to the hidden opening of its burrow, which accordingly becomes marked
by the vivid patch of Xanthoria . Tolmache (28) described the Chukchi’s
stone lamps as being fueled with blubber oil and wicked with reindeer moss;
the usual type of wick used among primitive northern peoples is commonly said
to be Sphagnum or the cottony tufts of Eriophorum .
Industrial Uses
The presence of lichenin in lichens has been utilized in the past for
brewing and distilling. Use of lichens instead of hops has been recorded
for one or more monasteries of European Russia and Siberia; the beer was
described as bitter but highly intoxicating. Alcohol production from lichens
is an old art, though now replaced by increased cultivation of potatoes,
importation of sugar, and distillation of wood, and was recommended as a
means of saving grain which otherwise could be diverted into alcohol produc–
tion. It has been stated that 20 pound of lichens would yield 5 liters of
alcohol. An early report published in stockhom in 1868, gives a detailed
account of, as well as plans for, setting up a distillery for the production
of lichen brandy. By 1893, the manufacture of this type of brandy was

EA-PS. Llano: Utilization of Lichens

considered an important local industry; but trough the exhaustion of
neighboring fields, the industry quickly lapsed into obscurity
The first commercial production of lichen-glucose sirup has been re–
ported by Diachkov and Kursanov (5) from two small plants on the Kola Penin–
sula in the U.S.S.R. with a daily yield of 80 kilograms of glucose sirup.
The method consisted of hydrolyzing the lichens with sulfuric acid, boil–
ing the residue with chalk, and finally purifying with charcoal. The bitter
substance of the Cladonias could be eliminated by the usual treatment with
weak alkalies, but it appeared again during the process of hydrolysis. The
nature and elimination of this bitter substance should be the subject t of
further research, for its presence is a serious hindrance to the utilization
of raw stuffs from a common group of lichens. Cetrarias, Alectorias, and
Usnea barbata are reported to yield a sirup with a content of 65 to 70%
glucose, representing 100% of the initial dry weight of the Lichen. The
lichen resources of the Murmansk region are reported to amount to 600,000
tons of dry matter, or about 5 tons of dry lichens per 2.5 acres, which
would suggest sufficient raw materials for the maintenance of local glucose–
producing units. However, in the absence of a practical agricultural plan
for the continual harvesting of lichen thalli, the supply would depreciate
yearly, necessitating longer hauls and conflicting seriously with the needs
for reindeer pasturage.
The use of lichens for tanning never achieved more than a local sig–
nificance in northern Europe. Two boreal lichens having some tanning prop–
erties are Cetraria islandica and Lobaria pulmonaria ; but among primitive
people of the circumpolar regions accustomed to leather and fur clothing,
the use of urine for tanning is a widespread custom, and there is no evidence
that lichens were ever utilized for tanning or dyeing.

EA-PS. Llano: Utilization of Lichens

Synthetic days have largely replaced many formerly common vegetable
days in the textile industry, primarily because of their low production cost
and somewhat better dyeing qualities such as light-fastness. Of the vegetable
dyes, those obtained from lichens were renowned among the peasant days of
old for their high quality and color, but today they are among the least known.
Recent efforst in Scandinavia, Ireland, and Scotland to revive hand-loom
wearing among isolated communities having an otherwise limited or seasonal
source of income have evolved into a lucrative business manly for the luxury
export trade. The use of lichen days, once the mark of genuine Harris tweed,
would probably add to its cost, but it would give further individuality to a
cloth noted for its lasting characteristics. This is partly due to the action
of lichen acids, which effectively and naturally mothproof homespuns. The
utility of lichens for dye production is, possible, one of their most promis–
ing application in arctic and subarctic areas where the introduction of
sheep is feasi a ble. With the increased settlement of northern lands by peoples
accustomed to the use of cloth. Hand looming and home dyeing might provide for
local consumption as well as for outside trade among low-income groups.
Most of the boreal lichens used in dyeing produce brown, gray, or yellow
colors. Parmelia saxatilis is known among Scandinavian farmers as the “dye
lichen,” giving various shades of brown. Cetraria islandica was equally pop–
ular for its brown-dyeing properties and is valued for dying suede, as it
produces the faint pastel tin st ts desired by the trade. Red-yellow coloring
con be extracted from the Usneas or beard lichens that are prevalent in coni–
ferous forests. Ochrolechia tartarea and Lecanova parella , crustaceous lichens
that form thick crusts on rocks, bark, mosses, or on the ground, have long
been highly esteemed for the production of the blue dyes that are more

EA-PS. Llano: Utilization of Lichens

commonly found among the E Roccellas of southern latitudes. Such dyes can also
be extracted to as more limited extent from Umbilicaria pustulata . The colors of
cudbear from Ochrolechica tartarea and archil from the Roccellaceae were com–
mercially indistinguishable, and the former could be varied to a permanent
black by the use of indigo or dye of lungwort. Lichens have long been used
for the commercial preparation of litmus paper. A variety of colors and shades
can be obtained by the use of different species of lichens, varying the treat–
ment with chemicals or other vegetable days. However, it should be noted that
the color of a lichens plant is no indication of the dye which may be obtained
from it.
Early methods used by peasants for dyeing homespuns involved the use of
urine as their only available source of ammonia. The appropriate lichen was
placed in barrels containing urine, and the mass was permitted to fermit for
about ten days, after which the yarn and alum were added prior to boiling. If
the lichen was not immediately utilized, it could be rolled into balls or cakes
with lime or burnt shells, wrapped in dock leaves, dirried over peat fires, and
stored until used. By boiling Parmelia saxatilis in a copper kettle for an
hour or two an extract is obtained which suffices for dyeing. In most cases
the common mordant used is alum.
There lichens, Evernia prunastri , E. furfuracea and Lobaria pulmonaria ,
have long been raw materials for the perfume and cosmetic industries, being
employed in the manufacture of toilet powders, scented sachets, and perfumes.
Their use today (1950) still persists, owing to the demands for the very stable
perfumes of modern extraction. Of these, E. furfuracea , which gros on both
deciduous and coniferous trees, or on rocks, used to be imported in large
bales from Yugoslavia by American processing firms. With the advent of

EA-PS. Llano: Utilization of Lichens

World War II, some supplies were obtained from the eastern United States,
but it was found that Evernia ( Letharia ) vulpina also contained the essential
oil. These lichens are distributed throughout the alpine and coniferous region
of the Subarctic.
The medicinal application of lichens flourished during a period of history
when herbals and country recipes were the main source of information on drugs;
any plants bearing vague resemblances to parts of the human body, or to
diseases, were thought to be specifically intended by a kind Providence for
the cure of ills. The application of so m e lichens still permists in the more
remote areas of the world, but, in the light of modern medical knowledge, their
worth has been justifiably depreciated. However, recent laboratory experiments
have shown some indication of antibiosis among lichens when used against Gram–
positive bacteria, while at least one lichen acid has been reported to inhibit
the growth of tubercular organisms in vitro . Should further experimentation
lead to practical results, and in the absence of synthetic production, the
utilization of lichens having the desired qualities would imply commercial
possibilities for northern lichen fields.
Possibilities for Future Rese ra ar ch
It is in the demand for an increased production of livestock and livestock
products that the practical utilization of the lichen fields of arctic and sub–
arctic lands is fundamental. The introduction of cattle, where practical, is
always dependent upon the difficulty and expense of the production and acces–
sibility of concentrated feeds, and of winter feed generally; lichen sources
[: ] offer feed supplement. Accessory problems include shelter and the need for
available water in winter. The usefulness of reindeer as a source of meat,
hides, and furs has been proved, but the present breeds are inefficient in

EA-PS. Llano: Utilization of Lichens

the production of fluid milk. Nevertheless, reindeer culture has demon–
strated its worth in aiding the Alaskan Eskimos, Scandinavian Lapps, and
Siberian natives to attain a degree of self-sufficiency which cannot be met
by seasonal and haphazard hunting in the face of a steadily dimihishing
source of wild game. It has all the possibilities of a basic industry, and
with local outlet channels should offer more stability and balance than the
present-day Eskimo way of life, which is being increasi ng ly challenged by the
intrusion of modern developments. The feasibility of developing an outside
market for meat will be dependent to a great extent upon the ability of the
industry to settle its own social and organizational problems and to meet the
competition of domestic livestock producers with the full use of modern [: ] p ack–
aging, shipping, advertising, and progressive retail outlets.
The means for improving the character of herds, whether they be raised
for marketing or for draft purposes, is well within the scope of modern animal–
husbandry techniques which could be applied on experimental farms — not only
for the improvement of the breed but to establish confidence and pride among
native owners. The accomplishment of these needs places greater emphases upon
the supplies of wild fodder.
The forage supply for reindeer on the northern submarginal lands is gen–
erally good; the vegetative cover is enormous on lands unsuitable for other
purposes, and theoretically there was some justification for the early optimistic
estimates of their carrying capacity. The distribution of the grazing lands in
relation to the seasonal needs of reindeer serves to reduce these figures to
more reasonable size, but the critical problem lies in the slow regenerative
properties of the lichen forage plants. It is a problem that has been met by

EA-PS. Llano: Utilization of Lichens

restrictive range practices rather than by an expanded program designed to
explore the underlying factors relating to growth and regeneration of the
more desirable species.
Chemical analysis to determine the composition and amount of lichen car–
bohydrates and other components at various times of the year, and upon various
substrata, is essential for the selection of the best qualities of these
forage plants and for a more complete understanding of grazing possibilities.
Very little is known about how to establish, manage, and maintain lichen
pastures agriculturally; efficient application of knowledge would be depend–
ent upon sound research findings. Utilization and management practices,
particularly winter-grazing management, need keen understanding of the limita–
tions of plant species.
The generally low calcium value of some northern native grasses, and of
certain lichens, may have some relationship to the reindeer’s habit of eating
rodents, birds’ eggs and young, and the gnawing of old bones, indicating
deficiencies in either the winter or summer feeds of certain areas which may
result in malnutrition unless otherwise supplemented. The effect of maternal
mineral deficiency upon the prenatal development of offspring has been clearly
proved in domesticated animals in regard to calcium, phosphorus, sodium,
potassium, manganese, copper, iron, and iodine; in the study of trace ele–
ments, the impo r tance of cobalt, moly b lenum, fluorine, etc., suggests fields
for further study in regard to lichen forage. The effect of mineral defici–
ences upon breeding is suggested by a report of the Atka reindeer herd; in
spite of an absence of predators, reasonable slaughter, and an availability
of pasturage, the hard increment has leveled off. The explanation that this
situation has resulted from the dominance of overage bulls would imply improper
culling of herd bucks at slaughtering time; if this is not the case, the
availability of minerals in the fodder may well repay further investigation.

EA-PS. Llano: Utilization of Lichens

BIBLIOGRAPHY

1. Aleksandrova, V.D. “Summer food of the reindeer on Novaya Zemlya,” Lenin–
grad. Arkticheskii Nauchno-Issled. Inst. Trudy , no.22,
pp. 35-50, 1925.

2. - - - -. “Winter forage of reindeer in Novaya Zemlya,” Arkticheskii Nauchno–
Issled. Inst., Sov. Reindeer Ind ., no.9, pp.127-139, 1937.

3. Cowan, I.McT. “The ecological relationships of the food of the Columbia
black-tailed deer, “ Ecological Monogr . vol.15, no.2, pp.109-39,
1945.

4. Dedov, A.A. “Character of the natural feeding ground of the Taimyr circuit,”
Arkticheskii Nauchno-Issled. Inst., Sov. Reindeer Ind ., no.2,
pp.7-48, 1933.

5. Diachkov, N. and Kursanov, A. “The carbohydrate composition of lichens of
the Kola Peninsula in connection with the problem of glucose
production in northern localities,” Akad. Nauk. Comptes Rendus
( Doklady ) vol.46, no.2, pp.60-68, 1945.

6. Douglas, W.O. Of Men and Mountains . N.Y., Harper. 1950.

7. Druri, I.V. “Reindeer pasture in the Chukchee Anadyr district,” Leningrad.
Arkticheskii Nauchno-Issled. Inst. Trudy , no.62, pp.105-24, 1936.

8. Dutilly, A. “A bibliography of Reindeer, Caribou, and Musk-Ox,” Research &
Development Branch, Office Quartermaster General, U. S. Army,
Rep . 129, 1949.

9. Gorodkov, B.N. “Natural grazing lands of the tundra zone of the Far-eastern
Province, “Arkticheskii Nauchno-Issled. Inst., Sov. Reindeer Ind .
no.2, p.161, 1933.

10. - - - -. A study of the growth of lichens. Ibid . no.8, pp.114-115, 1936.

11. Hovgaard, W. “The Norsemen in Greenland,” Geogr. Rev . vol.15, p.605, 1925.

12. Lantis, Margaret “The reindeer industry in Alaska,” Arctic , vol.3, no. 1,
pp.27-44, 1950.

13. Llano, G.A. “Lichens--their biological and economic significance,” Bot. Rev.
vol.10, no. 1, pp.1-65, 1944.

14. - - - -. “Economic uses of lichens,” Econ. Bot . vol.2, no.1, pp.15-45, 1948.

EA-PS. Llano: Utilization of Lichens

15. Mirov, N.T. “Notes on the domestication of reindeer,” Amer. Anthrop . vol.
47, no.3, pp.393-408, 1945.

16. Palmer, L.J. “Food requirements of some Alaskan game mammals,” J. Mammal .
vol. 25, no.1, pp.49-54, 1944.

17. Porsild, A.E. “Reindeer and caribou grazing in Canada,” 7th N. Am. Wild
Life Conf . 6-7, 1941-1942.

18. Rasanen, Veli “ [: ] The lichen flora of Petsamo,” Societas Zool.-Bot.
Fenn. Van. Ann. Bot . vol.18, no.1, pp.1-110, 1942.

19. Salazkin, A.S., and others. “Reindeer pasture and vegetative cover of the
Murmansk District,” Leningrad. Arkticheskii Nauchno-Issled.
Inst. Trudy , no.72, pp.307-12, 1936.

20. Sdobnikov, V.M. “The composition of the reindeer forage in autumn,” Lenin–
grad. Vsesoiuznyi Ark. Inst. Biull . no.24, pp.128-36, 1935.

21. - - - -. “Materials to the problem of winter food for the reindeer,”
Ibid . pp.137-41, 1935.

22. - - - -. “Relations between the reindeer (Rangifer tarandus) and the
animal life of tundra and forest,” Leningrad. Arkticheskii
Nauchno-Issled. Inst. Trudy , no.24, pp.5-66, 1935.

23. Seton, E.T. Lives of Game Animals . Garden City, N. Y., 1929.

24. Smith, P.S. “Exploration in northwestern Alaska,” Geogr. Rev . vol. 2 15, 1925.

25. Soczava, V. “Natural grazing lands of the tundra zone of Yakutia,” Arkti–
cheskii Nauchno-Issled. Inst., Sov. Reindeer Ind ., no.2, pp.47-118,
1933.

26. Sokolov, I.I. “Materials on the character of the exterior life history of
the domestic reindeer of the Bolshesemelskaya tundra, “Leningrad.
Veesoluznyi Ark. Inst. Biull . no. 24, pp.67-127, 1935.

27. Stefansson, V. “Resume of Arctic and problems of utilization. Problems
of polar research,” Amer. Geogr. Soc. Spec. Publ . vol.7,
pp.145-53, 1928.

28. Tolmachev, I.P. Siberian Passage . Rutgers Univ. Press, 1949.

29. Tulina, L. “On the forest vegetation of Anadyr land and its correlation
with the tundra,” Leningrad. Arkticheskii Nauchno-Issled. Inst.
Trudy , no.40, p.204, 1936.

30. Vasiliev, V.N. “The reindeer range in Anadyr region,” Ibid . no.62,
pp.9-104, 1936.

31. Wiklund, K.B. “The Lapps in Sweden,” Geogr. Rev . vol.13, pp.223-42, 1923.

George A. Llano
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