Economic Botany: Encyclopedia Arctica 6: Plant Sciences (Regional)

Author Stefansson, Vilhjalmur, 1879-1962

Economic Botany

Edible Plants of the Arctic

EA-Plant Sciences (A. E. Porsild)

EDIBLE PLANTS OF THE ARCTIC

CONTENTS

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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

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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,

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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

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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

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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,

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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

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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

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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

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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

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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.

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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

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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.

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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.

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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:

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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

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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

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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:

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Forest tundra 21 acres
Scrubby tundra 23 acres
Non-scrubby tundra 42 acres

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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 ; ^ : ^

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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

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^ 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.

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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)

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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

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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

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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

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LIST OF FIGURES

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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

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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

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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[]^i^chen 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 reside^sta^nce 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

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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

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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^

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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

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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

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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.

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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

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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

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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.

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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, dir^ri^ed 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

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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.

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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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