Flora and Vegetation of Arctic Alaska, Yukon, and Northwestern Canada: Encyclopedia Arctica 6: Plant Sciences (Regional)

Author Stefansson, Vilhjalmur, 1879-1962

Flora and Vegetation of Arctic Alaska, Yukon, and Northwestern Canada

EA-Plant Sciences (A. E. Porsild)

FLORA AND VEGETATION OF ARCTIC ALASKA YUKON, AND NORTHWESTERN CANADA

CONTENTS

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Page
Introduction 1
The Flora 3
Types of Vegetation 6
Rocky Barrens or Fell-fields 8
Tundra 12
Strand Vegetation 17
Vegetation of Fresh Water 18
Bibliography 20

EA-Plant Sciences (A. E. Porsild)

FLORA AND VEGETATION OF ARCTIC ALASKA YUKON, AND NORTHWESTERN CANADA
INTRODUCTION
Floristically the North American Arctic, here taken to include the total land area lying north of the tree line, may be divided into four major provinces, namely ( 1 ) arctic Alaska and Yukon, ( 2 ) the arctic Archipelago, ( 3 ) continental parts of arctic Northwest Territories and Ungava, and ( 4 ) Greenland. The present description however, embraces only arctic Alaska– Yukon and the arctic parts of Mackenzie-Keewatin to which has been added the western and central islands of the Arctic Archipelago. It does not include arctic Ungava, the easternmost islands of Ellesmere, Devon, Cornwallis, Somerset, Baffin, or the islands of Hudson Bay. In this area the writer today recognizes 872 species (and varieties of well-defined geographical range) of vascular plants distributed among 60 families and 194 genera.
Although the two areas under discussion are separated by a rather strongly marked phytogeographical boundary - the Mackenzie Valley - they have many features in common. Approximately one-third of the species which make up the combined floras are widely distributed, circumpolar species. Common to them both, as the most striking feather, is the total absence of trees. There are,

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to be sure, a number of ligneous or woody plants - such as willow, dwarf birch, and various kinds of heath and berry bushes - but even these become low and dwarfed north of the tree line, and are generally restricted to places where protecting snow cover is assured during the winter. In the interior of the mainland some of the these species still are ^ a ^ dominant component of the vegetation, but when we reach the seacoast or cross to the arctic islands their numbers dwindle rapidly. The few woody plants that occur there become low and creeping, with their stems partly buried in the moss, while some of the berry bushes cease to produce fruits, or disappear entirely. the climate of the entire region is arctic but there are, nevertheless, strong regional differences; thus the Bering Sea shores, and also the Canadian Arctic east of Hudson Bay, has a maritime polar climate whereas the Mackenzie and Keewatin districts, and also the Arctic Archipelago, all have distinctly con– tinengal types of climate with relatively warm summers and very cold winters. This is well illustrated by the course of the 50° F. isotherm for the warmest month, July, which in western Alaska first parallels the Bering Sea coast, then swings abruptly eastward across the north slope of Alaska. From the mouth of the Mackenzie, it follows the Arctic coast east to Bathurst Inlet whence, by the cooling influence of Hudson Bay, it is deflected in a southeasterly direction, bisecting the west coast of Hudson Bay between Chesterfield Inlet and Churchill.
Even locally the proximity of the sea strongly affects the temperature and thereby also the vegetation. Thus, at Nome on the south coast of Seward Peninsula, the mean temperature for July is 50° F. whereas Kotzebue, at the head of Kotzebue Sound and only 1 1/2 degrees farther north, but sheltered by Seward Peninsula, has a 2-degree warmer July mean. Of much more importance, however, is that the average numbers of days without killing frost is 52 at

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Nome but 93 at Kotzebue. The mean annual precipitation at Nome is 18 inches whereas at Kotzebue it is only slightly over 6 inches. Barrow, on the other hand, has an extreme polar climate; the mean for July there is 40° but the number of days without killing frost is only 17, At Coppermine, and at Cambridge Bay on Victoria Island, the mean for July is 10° F. higher and the number of days without killing frost almost seven times as high as at Barrow.
Except for the distinctly moister southern part of the Bering Sea region, precipitation throughout the area is light, averaging perhaps about 10 inches for the year. All records, however, are from stations situated along the coast, and there is reason to believe that the annual precipitation of the interior mainland as well as that of some of the larger islands is consider– ably less. Most of the precipitation falls as snow during the winter but is swept off the ground by frequent gales, so that the plant cover, moreover, is exposed to the drying effect of the wind. The light precipitation is to some measure compensated by the fact that 40 per cent falls as rain during the months of June, July, and August. Of far greater importance, however, is that owing to the presence of permafrost a few inches below the surface, moisture derived from melt water as well as from rain is prevented from penetrating to depths beyond the reach of plant roots. Without permafrost many areas in the Arctic would be a lifeless desert.
THE FLORA
Arctic Alaska and Yukon — largely unglaciated during the Pleistocene — presents the greatest variety of plant habitats; its flora is rich in isolated and endemic species and, undoubtedly, is very old. Through its former land connection across hypothetical Beringia, there has been a free interchange of species so that almost one third of the present flora of 604 species of

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vascular plants is of Asiatic affinity. The much smaller number of Cordilleran species may have reached the area by way of the Mackenzie Mountains. In the Eastern Arctic practically all mountains areas were covered by huge ice caps during the Pleistocene. In consequence, the alpine flora of the Eastern Arctic is youthful, poor in species, and usually entirely lacking in rare or endemic species. In Alaska and Yukon, on the other hand, although some of the high mountains may ^ ^ have had, or even now have, isolated ice caps, such glaciations as existed was largely limited to cirques and valley glaciers; during the Pleistocene many of the large mountains massifs undoubtedly acted as refugia and, indeed, today harbor the richest floras of rare and endemic species (5; 14; 16). Such discontinuous vertical distribution of plants is very noticeable on the Seward Peninsula and, indeed, in most places in arctic Alaska and Yukon wherever mountains or hills exceed one thousand feet in elevation. Below this level the coastal flats are occupied by a wet, marshy tundra that everywhere is inhabited by practically the same assortment of species. The most notable change in the flora, when one travels inland, away from the seashore, is the increasing ration of fruticose species over herbaceous ones.
On the hills, on the other hand, above the one-thousand-foot level, a very different and much more varied flora is encountered; here we find most of the rare and interesting plants for which the region is famous, notably those of Asiatic affinity or at least of bilateral Beringian range. The writer (11) selected 88 species that, in the Bering Sea region, appear to be restricted to levels above one thousand feet. By grouping these species according to their general distribution it is found that, roughly speaking, three-fourths are either Western-Cordilleran, Bering Sea endemics, or essen– tially Asiatic species transgressing into western America, whereas but one-

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fourth are widely ranging, circumpolar, or North American species that are common across the continent from the Atlantic to the Pacific coast. A further breakdown discloses that 17 species are circumpolar or essentially so, 3 are essentially North American species, known from the Atlantic to the Pacific coast, 16 are western American or Cordilleran, 15 endemics of the Bering Sea region, and no less than 37 are Asiatic or Beringian bilateral species transfressing into western America.
The Arctic Archipelago was formerly though to have escaped glaciations. Recent investigations, however, indicate that at least the southern islands have been severely glaciated and that all but the higher parts of the Archi– pelago was submerged in Pleistocene time, (13; 20). The climate is decidedly high-arctic, and the flora, consequently, is comparatively poor in species. The total number of vascular plants species known to occur in the Archipelago has recently been increased to 308, of which 226 occur in the western part and no less than 214 in Banks and Victoria islands alone (13). Almost one-half are widely distributed arctic species. The presence of a compara– tively large number of isolated and endemic species offers some curious phytogeographical problems.
The arctic Canadian mainland is characterized by a distinctly continental and very dry climate which, together with the absence of mountain barriers, accounts for its strikingly uniform and monotonous plant cover. Of the 651 species known to occur north of the tree line, 70 per cent are widely distri– buted circumpolar arctic or North American species. A considerable number of nonarctic forest species reach far north of the present tree line and, together with tree stumps in situ and peat deposits, may indicate past oscillations in the tree line. In terms of postglacial chronology, the flora is

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youthful, as indicated by unstable, preclimax plant communities and by the general paucity of endemic and isolated species. To the west, the Mackenzie Valley forms a strongly marked floristic boundary which has been crossed by comparatively few species of Cordilleran or bilateral Beringian range.
TYPES OF VEGETATION
Land surfaces in the Arctic may be divided into three principal tyles of landscape: ( 1 ) the ice desert, ( 2 ) the rock desert, and ( 3 ) the tundra. The first, which in the form of glaciers, ice caps, or perennial snow fields, occupy 86 per cent of the interior of Greenland, and also some smaller moun– tain regions in Baffin, Devon, Ellesmere, Axel Heiberg, besides several islands of the Sverdrup group, in the area under discussion occurs only as local glaciers in the Brooks Range. The rock desert - the dominating land– scape in heavily glaciated northern Keewatin, northeastern Mackenzie, and the arctic islands - in Alaska and Yukon is found only at high elevations in the mountains. In arctic Cnada the rock desert is everywhere interspersed with lakes and ponds, or with small or large areas of tundra. The tundra, finally, occupies a wide belt between the edge of the forest and the high– arctic rock desert and is endowed with the richest flora. Like the rock desert it may be divided into several distinct plant communities such as marsh, grassland, and dwarf-shrub heath.
The present treatment deals mainly with the vascular plants. The cryp– togamic flora of the arctic landscape is very incompletely known outside Greenland, and its ecological aspects are practically unknown. Among the terrestrial cryptogams, bryophytes and lichens are both very significant

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ecologically in the composition of arctic vegetation.
Thus far comparatively little progress has been made in the study of arctic plant communities in North America. In the heavily glaciated parts of Keewatin, Mackenzie, and the Arctic Archipelago, the comparative youthful– ness of the flora and vegetation is evidenced by unstable plant communities, whereas in unglaciated Alaska and Yukon, and in the coastal plain east of the Mackenzie, climax conditions may have been attained in the lowland “niggerhead” tundra which is so characteristic a feature of the coastal plain landscape.
Four major types of vegetation may be recognized in arctic North America, each capable of subdivision into a number of more or less distinct ecological associations, the composition and interrelation of which again depend on the physiography of the landscape.
Rocky barrens or fell-fields
Vegetation of rock desert Vegetation of unstable screes and frost-activated (congeliturbated) soils Vegetation of gravelly river flats, flood plains, and fans
Tundra
Dwarf-shrub heath Lichen and moss heath Grassland Copses and thickets Marsh and wet tundra Snow-flush herb mats
St ^ r ^ and vegetation
Brackish meadows and marsh subject to floods Vegetation of sand dunes and gravel beaches Vegetation of rocky shores
Vegetation of fresh water
Vegetation of ponds and lakes Vegetation of brooks and rivers

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Rocky Barrens or Fell-fields
Vegetation of Rock Desert . The fell-field, or rocky barren, has been defined ( [: ] ^ 19 ^ ) as having so scanty a vegetation that the ground is only partly covered. To this should be added that the winter snow cover is generally deficient. Fell-field vegetation is found at high elevations and also in high latitudes and is the dominant vegetation type in the rock desert of northern Keewatin and Mackenzie, the greater part of the Arctic Archipelago, and the highest mountains of Alaska and Yukon.
Most striking features of the rock-desert landscape are the flat-rock tables and ridges of pre-Cambrian crystalline rocks of the heavily glaciated Laurentian Shield or peneplain that has been left almost completely denuded of soil, and the extensive, flat, or slightly rolling rock-strewn plateaus of severely weathered Paleozoic or pre-Cambr ian sediments predominant in the western and northern islands of the Archipelago. But although soil is every– where scarce, these wastelands, nevertheless, are not completely lacking in vegetation. Over large areas the rock surfaces of the acid pre-Cambrian rocks are covered by black, brown, and gray species of the crustaceous lichens ( Rhizo–carpo n , Licidea , Lecanora , Buellia , and others), or by the much larger black and green foliose lichens, sometimes called “rock tripe” ( Gyrophora and Umbil– icaria ) that all lend a rather somber and depressing color to the landscape. Some of the Silurian and Paleozoic rocks of the arctic islands, on the other hand, almost completely lack lichen cover.
On loose rocks and gravel grow a number of fruticose species among which may be mentioned the coral-like Stereocaulon , black and gray Alectoria , and yellow, gray or brown species of Cetraria and Cladonia . On cliffs, below

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ledges providing nesting sites for gulls, hawks, falcons, or ravens, or on isolated boulders that, in the Arctic, are favorite perches for snowy owls and other birds, the dung-loving orange or vermilion lichen ( Calopaca elegans ) often grow in great profusion and lend a touch of color to the otherwise bleak landscape
Flowering plants, however, are not entirely missing in the arctic rock desert, for the rock-strewn barren flats or cliffs that from the air may look entirely devoid of vegetation, on closer inspection from the ground may dis– close small or large patches, or “islands,” of vegetation composed of mountain avens ( Dryas integrifolia ), yellow arnicas, several species of locoweed ( Oxy–tropis arctica , O. pygmaea , O. Maydelliana , and O. arctobia ), or saxifrages. For a short time each summer these flowers transform these cases of the rock desert into incredibly gas miniature rock gardens.
A number of plants that grown on windswept, stony soil develop tussocks or dense, flat, hemispherical cushions, that by their shape provide protection against evaporation and mechanical abrasion by wind, drifting sand, or snow. Examples of such plants are the moss pink ( Silene acaulis ), Diapensia lapponica , Loiseleuria procumbens , a number of saxifrages, Draba Bellii , and many others. In rock crevices grow a number of species that, like the foregoing, are all strongly xerophytic. Among them are several saxifrages ( Saxifraga nivalis and S. reflexa ), tufted fescue ( Festuca brachyphylla ), several sedges including Carex rupestris , and the rock ferns Woodsia ilvensis , W. glabella , and Dryopteris fragans .
On gravelly slopes or in shelter of rocks where some soil has accumulated, the showy, white-flowered avens ( Dryas ) forms large, flat cushions, often many feet in diameter, sometimes to the exclusion of other species. In such places

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we may also find mats of crowberry ( Empetrum ), clumps of the purple-flowered, broad-leaved willow herb ( Epilobium latifolium ) or the vetch Hedysarum Mackenzii , whose showy purple flowers are scented like sweet clover. Other colors are added by the lilac-flowered vetch ( Astragalus alpinus ), by yellow arnicas and locoweeds ( Oxytropis ), by blue campanulas, bluish-white anemones ( Anemone parviflora ), white chickweeds ( Stellaria and Cerastium ), or by the large-flowered and very fragrant Pyrola grandiflora . In the southern part of the arctic rock desert occur a number of dwarf shrubs, including several species of willow, Lapland rho ^ do ^ dendron ( Rhododendron lapponicum ), dwarf azalea ( lois–eleuria procumbens ), Labrador tea ( Ledum decumbens ), and bilberry ( Vaccinium uliginosum ). These are always low and prostrate with their stems bent and gnarled. Furthermore, their shape is often contorted by wind action so that the prostrate crown grows away from the root by which it is anchored. Some of them grow espalierlike against south-facing rocks or cliffs from which they obtain added warmth. The roots of these dwarf shrubs are spread horizontally in the upper layers of the shallow soil, avoiding the permafrost. The majority are evergreen, and in the structure of their leaves possess special features, such as hair covering and concealed stomata to prevent loss of water.
Vegetation of Unstable Screes and Frost-activated (congeliturbated) Soils . Unstable rock screes and stone creeps, together with soil sorting and heaving, usually result from frost action in the soil and are features of high-arctic climates such as are found in high latitudes or at high elevations. The un– stable soil conditions of these habitats make them unsuitable to many plants, especially such as have woody stems or horizontal thizomes, whereas those with superficial runners, strongly developed taproots or adventitious root systems are able to persist. A number of xerophytic species, particularly in the

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families ( Cruciferae and Compositae ( Erysimum pallassii , Lesquerella arctica , Smelowskia calycina , Draba spp., Arabis spp., Crepis nana , Artemisia spp., and Taraxacum spp., to mention only a few) by their growth form seem especially well adapted to slide-rock habitats. In the strongly frost-activated soil of polygon fields in Banks and Victoria islands, avens ( Dryas integrifolia ), cinquefoil ( Potentilla rubricaulis ), and the locoweed ( Oxytropis arctica ) are extremely common and frequently comprise 90 per cent of the plant cover.
Vegetation of Gravelly River Flats, Floodplains, and Fans . Floodplains, river flats, and fans are inhabited by plants whose growth form, like that of the scree plants, permit them to endure unstable soils, but they must also be able to withstand water erosion and inundation. Because the soil movements are less violent and water is abundant, a few woody species, notably willows and several hemicryptophytes with sell-developed rhizomes may grow here as well. [: ] The broad-leaved willow herb ( Epilobium latifolium ), the vetches (Hedysarum alpinum and Astragalus alpinus ), besides several grasses and sedges, are ubiquitous in these habitats.
The predominance of alpine and arctic species on erosion fans and flood– plains has sometimes been taken to indicate that these plants were washed down from adjacent mountains by glacial torrents during spring freshets, and that they were able to gain a foothold only because lowland species are not so well adapted to the edaphic conditions which are peculiar to these habitats. Per– haps the primary reason why alpine and arctic plants frequently dominate flood– plains and erosion fans, is that huge masses of overflow ice (crystocrene) accumulate there in winter and frequently persist until late in the summer, thereby shortening the season to such an extent that only truly alpine or arctic plants can survive.

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Tundra
Dwarf-shrub Heath . The tundra differs from the rock desert by having a closed or continuous cover of vegetation and in having a protective cover of snow in winter. The dwarf-shrub heath of the tundra is essentially a continental lowland plant association and reaches its best development in the interior coastal plain of the north slope of Alaska at some distance from the sea, in northern Mackenzie, and in central Keewatin. Its principal components are low willow ( Salix spp.), dwarf birch ( Betula glandulosa , Labrador tea ( Ledum decumbens , and L. groenlandicum ), Lapland rhododendron ( Rhododendron lapponicum ), bearberry ( Arctostaphylos alpina and A. rubra ), besides white heather ( Cassiope tetragona ), bilberry ( Vaccinium uliginosum ), and the alpine cranberry of cowberry ( Vaccinium Vitis-Idaea ). Ubiquitous also are a number of sedges and grasses such as Carex lugens , Arctagrostis latifolia , and Festuca altaica , but notably cotton grass ( Eriophorum vaginatum ) which forms huge tussocks that are known in Alaska as “niggerheads.” These solid, turfy hummocks may stand a foot or a foot and a half above the surrounding ground and are notorious as the chief impediment to travel across the tundra in summer or winter. When viewed from a distance the “niggerhead tundra” has the appearance of a slightly uneven grassy plain, and in mid-July, when the “cotton” matures, may look completely white as if the land were covered with snow.
Among the smaller herbaceous plants growing on the tundra are several fernweeds ( Pedicularis labradorica , P. lanata , P. arctica , and P. sudetica ), baked-apple ( Rubus Chamaemorus ), groundsel ( Senecio atropurpureus and S . lugens ), Saussurea angustifolia , blue lupine ( Lupinus arcticus ), wintergreen ( Pyrola grandiflora ), the knotweed ( Polygonum Bistorta ssp. plumosum ), the

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root of which is edible, yellow saxifrage ( S. Hirculus ), Spiraea Beauverdiana , and many others. Many of these have brightly colored flowers and for a short time in early summer the dwarf-shrib heath is gay with color, but during the rest of the year presents a dreary and desolate aspect.
The ground under and between the stems of the dwarf bushes and between the tussocks is everywhere covered by a dense carpet of mesophytic mosses and lichens into which the foot of the traveler sinks ankle deep at every step, making traveling on foot exceedingly wearisome.
Lichen and Moss Heath . Near the timber line, in drier and better drained parts of the tundra, the dwarf-shrub heath imperceptibly passes into lichen and moss heath. The place of mesophytic species of mosses is here taken by gray and brown fruticose lichens of which the so-called reindeer mosses ( Cladonia rangiferina , Cl. alpestris, Cetraria nivalis ) and Iceland moss ( Cetraria islandica ) dominate to such an extent that the landscape may appear as if snow-covered. In time most herbs and the low dwarf shrubs become sub– merged in the lichen carpet, and only dwarf birch, rhododendron, and willow remain.
Grassland . Extensive grassland or natural meadows are occasionally found in the tundra regions, especially in central Keewatin, on alluvial flats formerly occupied by large glacial lakes. These meadows are still wet in spring but there is reason to believe that they will, in time, be invaded by tundra shrubs and eventually change to dwarf-shrub heath. The principal meadow-forming species are Carex saxatilis and C. membrancea , Arctophila fulva , Arctagrostis latifolia , Deschampsia caespitosa , Juncus balticus , Carex stans and C. canescens , besides a number of herbaceous tundra herbs.
Copses and Thickets . Willow and alder thickets, with their accompanying

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flora of herbaceous species, are found chiefly on the continent, in well– drained soils along streams, on south-facing slopes, and on the sloping banks of lakes and rivers where protection against wind and consequent accumulation of snow is assured. The alder ( Alnus crispa ), although it does not extend far north of the tree line, form thickets 18 feet high with trunks up to six inches in diameter, along the lower courses of rivers flowing into Bering Sea. At some inland villages, where timber or driftwood is not available, alder is the principal source of fuel. The principal thicket-forming willows are Salix arbusculoides , S. pulchra , S. Richardsonii , S. glauca , S. planifolia , and S. alaxensis . Only the last may occasionally assume tree like growth, whereas the rest are shrubs that rarely exceed 8 feet in height. In protected places with abundant snow cover, willow thickets may be found even in the arctic islands. Thus, on the south-facing slope at Walker Bay, on the west coast of Victoria Island, in latitude 71° 30′, Salix Richardsonii forms six-foot-high thickets.
Some of the common herbs growing inside the thickets are fireweed. ( Epilobium angustifolium ), horsetail ( Equisetum arvense ), the anemones ( Anemone parviflora and A. Richardsonii ), buttercup ( Ranundulus lapponicus ), coltsfoot ( Petasites frigidus ), groundsel ( Senecio lugens ), wintergreen ( Pyrola grandiflora ), and a number of grasses and sedges. Among the latter, cotton grass ( Eriophorum angustifolium ), blue joint grass ( Calamagrostis canadensis var. Langsdorffi i), and Arctagrostis arundinacea are the most important.
Marsh and WetTUndra . Wet, grassy tundra predominates on low coastal flats of the unglaciated arctic coast of Alaska and probably is a climax type found on thick deposits of raw humus or peat that here often rests on a massive thickness of fossil ice. The often very great thickness of frozen, raw humus suggests that this lowland tundra formation is very old. It was probably

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formed during a period with a climate wetter than the present and may be considered a postclimax relic which is no longer being actively formed. A characteristic feature of thus tundra is the presence of numerous ponds and lakes that have formed in places where the fossil ice below the tundra has melted. Such local erosion is actively going on everywhere in the tundra today; it frequently starts by the widening and cutting, by meltwater, of a frost crack or when, in some other manner, the insulating cover of humus is removed by water or wind erosion. Such tundra ponds usually have steep, slumping banks in which may usually be seen a wall of pure ice (10; 11). This form of tundra occurs only in unglaciated parts of arctic North America but has its counterpart in arctic Eurasia.
Another feature of special interest in the low wet tundra is well-developed patterns of irregular polygonal ridges of peat which have formed along frost cracks. Another peh phenomenon caused by frost action are the conical hills or pingos formed in former lake basins by the local upheaval due to expansion following the progressive downward freezing of a lens or body of water or semifluid mud or silt trapped between the frozen surface soil and a lower permafrost layer. East of the Mackenzie delta some pingos , formed in this manner, are several hundred feet high (10). Around the bases of these hills are concentric peat ridges separated by moatlike ribbons of water.
The flora of the wet, lowland tundra is more varied and far richer in species than is the dwarf-shrub tundra. In the wet tundra back of Atkinson Point on the Arctic Coast, east of the Mackenzie Delta, the writer (8) noted 150 species of flowering plants, more than a third of them grasses and sedges. Dwarf shrubs are almost entirely absent and represented chiefly by creeping willows. Fruticose lichens, too, are scarce whereas bryophytes are well represented.

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Snow-flush Herbmats . An association of particular interest is found in places where large snowdrifts accumulate during winter. Mountain country, especially if the snowfall is heavy, favors the formation of large drifts, but even in the lowland where the snowfall is generally light, large drifts, favorable for the formation of snow-flush vegetation, may be formed by the prevailing wind in river valleys, on sheltered slopes, at the base of cliffs, and along lake shore . s. Such habitats are late, for the snowdrift may last well into the summer or, indeed, may not disappear entirely before winter starts. The snowdrift, on the other hand, throughout the summer provides an unfailing supply of water and, besides, contains a very considerable amount of potential plant food in the form of wind-transported inorganic dust or loess mixed with seeds and plant remains. The accumulation of fertile soil which is often found on screes and slopes is deposited in this manner.
The principal difference between the willow thicket and the snow-flush association is due to the duration of the snowdrift: too deep snowdrifts shorten the season too much for willows. For this reason willow thickets usually are found on the edge of snow-flushes, thus giving the impression that the willows are invading the herbmat area. The assembly of species that occupy snow-flushes are almost entirely herbaceous, and chiefly of the hemicryptophytic type in which the wintering buds are placed below the surface of the soil. The tiny willow ( Salix herbacea ) in the East, and ( S. pseudopolaris ) in the West are among the few woody plants found on snow-flushes. Characteristic, and very necessary for snow-flush plants is their ability to persist vegetatively for several seasons, when thick snowdrifts reduce their growing season to a minimum. Many snow-flush plants may not flower and fruit for a period of years and may even survive several unfavorable seasons completely buried under snow and ice.

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Whereas the general aspect of snow-flush vegetation is similar everywhere in the Arctic, its floristic composition varies greatly according to geo– graphical position. For example, snow-flushes in Greenland and in the Mackenzie Delta have almost no species in common. Examples of widely distri– buted snow-flush plants are the alpine spear grass ( Poa alpina ), sedge ( Carex scirpoidea ), mountain sorrel ( Oxyria digyna ), dwarf buttercup ( Ranunculus pygmaeus ), and alpine club moss ( Lycopodium slpinum ).
Strand Vegetation
Brackish Meadows and Marsh Subject to Floods . Salt marshes subject to floods are found chiefly on the low continental strand flats of western and northern Alaska, and from the delta of the Mackenzie extend east to the mouth of Anderson River. Along this shelving coast large, shallow, and brackish bays or lagoons are cut off from the open sea by long sandspits or by narrow sand or gravel islands formed by sea currents and by ice-push. Rivers empty– ing into the lagoons speed up the silting process, which is further favored by the almost neglibible tide. Back of the lagoons are vast meadow-covered flats that, as one travels away from the coast, imperceptibly change into low tundra. Most common in the meadows are grasses, sedges, and rushes, among them Arctophila fulva , Deschampsia caespitosa , Dupontia psilosantha , D. Fisheri , Puccinallia paupercula var. alaskana , and P. phryganodes , Carex aquatilis , C. glareosa var. amphigena , C. maritima , and C. rariflora , besides Eriophorum Scheuchzeri and E. angustifolium , Juncus balticus and J. castaneus . Among the more common herbs growing in the meadows are the chickweed ( Stellaria humifusa ), scurvy grass ( Cochlearia ), Chrysanthemum arcticum , the sweet-scented white buttercup ( Ranunculus pallassii ), the white cowslip ( Caltha natans ), and marsh fleabane ( Senecio congestus ).

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Vegetation of Sand Dunes and Gravel Beaches . Along the shores, on raised beaches and sandspits, are extensive dunes inhabited by lyme grass ( Elymus arenarius ssp. mollis ), fescue ( Festuca rubra var. arenaria ), spear grass ( Poa arctica and P. eminens ), and Carex Gmelini — the last two do not enter Canada — brome grass ( Bromus Pumpellianus var. arcticua ), seabeach sandwort ( Arenaria peploides ), stonecrop ( Rhodiola integrifolia ), blue lungwort ( Mertensia maritima ), wormwood ( Artemisia Tilesii ), Aster sibericus and several species of dandelion ( Taraxacum spp.). On the edge of the dune, in wet sand near the beach, is often found the delicate blue gentian ( Lomatogenium rotatum ), creep– ing buttercup ( Ranunculus gymgalaria ) and the dwarf sedges, Carex subspathacea and C. ursina .
Vegetation on Rocky Shores . On cliffs and boulder beaches that are not too exposed to the open sea may be found dense swards of Puccinellia phryganodes and similar grasses that occupy crevices and crannies between rocks. Here also grow scurvy grass ( Cochlearia ), chickweed ( Stellaria humifusa ), and a few other seashore plants.
Vegetation of Fresh Water
Vegetation of Ponds and Lakes . Nearly all large and deep lakes in the Arctic are too cold for vascular aquatic plants, but they possess a rich flora of microorganisms such as green and blue-green algae, desmids, diatoms, and flagellates. Different species of these mostly microscopic plants often impart a distinctive colorto the waters of the lakes. This is the best seen from the air, when it is at once noticeable that, apart from the difference in color caused by the depth of the water and the nature of the bottom, the water itself in no two lakes is exactly alike.

EA-PS. Porsild: Flora and Vegetation

Shallow ponds and lakes that remain water-filled throughout the summer, on the other hand, sustain a surprisingly rich plant life. Because of their shallowness, the water of these ponds warms up more quickly and becomes free of ice long before that of the large and deep lakes. The short season prevents many species of aquatic plants in the Arctic from reproducing by seed, but they maintain themselves successfully, nevertheless, by the production of wintering buds ( Potamogeton spp., Utricularia spp., and others). The most common aquatic plants here are pondweeds ( Potamogeton filiformis ), the arctic buttercup ( Ranunculus hyperboreus ), and mare’s-tail ( Hippuris vulgaris ). Floating and submerged mosses are abundantly represented, as are freshwater algae.
Vegetation of Brooks and Rivers . Small, sluggish lowland streams in the Arctic may sometimes be inhabited by aquatic plants similar to those of fresh– water ponds. Most arctic streams, however, are too cold or too turbulent for vascular species, but a number of marsh plants - chiefly sedges and grasses - inhabit protected and sheltered places on stream margins and flood plains. A number of dark-colored mosses ( Grimmia spp. and Hygrophypnum ) and blue-green and brown algae grow on the rocks of most arctic brooks, even in water that is but a few degrees above freezing. The brown, slimy flagellate ( Hydrurus foetidus ) flourished on the rocks of clear pools in ice-cold meltwater below snow fields.

EA-PS. Porsild: Flora and Vegetation

BIBLIOGRAPHY

1. Anderson, J.P. Flora of Alaska and adjacent parts of Canada , Parts 1 - 8. Iowa State College, Vols. 18, 18, 20, 21 and 23. 1943.

2. Anonymous. Climate and Man . Year Book of Agriculture. U.S. Dept. Agric., Washington, 1941.

3. ----. Meteorology of the Canadian Arctic . Dept. Transp., Met.Div. Ottawa, 1944.

4. Fernald, M. K. “Persistence of plants in unglaciated areas of boreal America,” Mem . Gray Herb., 2. 1925.

5. Hult e ^ é ^ n, E. Outline of the history of arctic and boreal biota during the Quaternary period . Stockholm, 1937.

6. ----. “Flora of Alaska and Yukon,” Parts 1 - 9. Lunds Univ. Årsskr ., vols. 37, 38, 39, 40, [: ] 41, 42, 43, 44, and 45. 1941.

7. Johansen, Frits. “General observations on the vegetation,” Report Can. Arct. Exp. 1913-18, vol.5, Ottawa, 1924.

8. Porsild, A.E. Reindeer Grazing in Northwest Canada . Dept.Interior, Ottawa, 1929.

9. ----. “Flora of Northwest Territories,” In Canada’s Western Northland . Dept. Mines & Res. Ottawa, 1937.

10. ----. “Earth mounds in unglaciated arctic northwestern America,” Geogr. Rev., vol.28, pp.46-58, 1938.

11. ----. “Contributions to the flora of Alaska,” Rhodora , vol.41, pp.141-83, 199-254, and 262-301. 1939.

12. ----. “Materials for a flora of Northwest Territories of Canada,” Sargentia , vol.4, 1943.

13. ----. “A biological exploration of Banks and Victoria islands,” Arctic vol.3, pp.45-54, 1950.

14. ----. “Botany of southeastern Yukon,” Nat.Mus.Can. Bull . 121, 1951.

15. [: ] Porsild, M.P. “The flora of Greenland: Its affinities and probable age and origin,” Torreya vol. 22, pp.53-54, 1922.

16. Raup, Hugh M. “Botanical problems in boreal America,” Bot.Rev ., vol.7, nos. 3 and 4. 1941.

17. Simmons, H.G. “A survey of the phytogeography of the Arctic American Archipelago,” Lunds Univ. Arsskr . N.F. Afdl. 2, Vol.9, 1913.

EA-PS. Porsild: Flora and Vegetation

18. Walker E. P. “Alaska: America’s continental frontier outpost,” Smithsonian Inst. War Backgr.Studies , No.13, Washington, 1943.

19. Warming, Eng. “On Gronlands Vegetation,” Medd. om Gronl ., vol.12, Copenhagen. 1888.

20. Washburn, A. L. “Reconnaissance Geology of portions of Victoria Island and adjacent regions arctic Canada,” Geol.Soc.Am. Mem . 22, 1947.

A. E. Porsild
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