Environment of the Yukon
Transcription
Environment of the Yukon
Environment of the Yukon G.G.E. SCUDDER Department of Zoology, University of British Columbia 6270 University Blvd., Vancouver, British Columbia V6T 1Z4 Abstract. The Yukon Territory, situated in the northwest corner of Canada adjacent to Alaska, and north of 60°, has a rugged and complex physiography and geology, dominated by mountainous terrain. There are five major watersheds, several large rivers, and many smaller, rich river valleys. The area has innumerable small ponds and damp depressions, but few large lakes. Yukon climate is typified by long, cold winters and short, warm summers. The Yukon has the most variable climate in North America, and across the Territory nine distinct climatic regions can be recognized. Most of the Yukon has either permanent, discontinuous or scattered permafrost. Cryosolic soils dominate in the northern permanent permafrost areas, whereas Brunisolic soils occur in the south where there is discontinuous or scattered permafrost. The dominant vegetation types in the Yukon are arctic tundra, alpine tundra, taiga or subarctic forest, boreal forest and subalpine-shrub forest. Arctic tundra and taiga prevail in the northern part of the Territory, whereas the boreal forest and subalpine-shrub forest characterize the southern part. Alpine tundra occurs at higher elevations throughout the Territory. Five distinct ecozones and 23 ecoregions are now recognized in the Yukon. In addition, scattered throughout the Territory are a number of habitats of special interest to entomologists. These include springs, peatlands, saline flats, sand dunes, and distinctive xeric sites with Artemisia frigida-grass communities. During the Pleistocene much of the Yukon remained ice-free and was part of Beringia, the largest refugial area in the north during the last ice age. As such, the Yukon is a central focus of biogeography in the Nearctic region. Résumé. L’environnement au Yukon. Le territoire du Yukon, situé dans la partie nord-ouest du Canada, juste à côté de l’Alaska, au nord du 60e parallèle, a une géologie et une physiographie accidentées et complexes et est dominé par les systèmes de montagnes. On y trouve cinq bassins hydrographiques principaux, plusieurs grandes rivières et de nombreuses petites vallées de cours d’eau plus petits. Le terrain est parsemé d’innombrables petits étangs et dépressions humides mais compte peu de grands lacs. Les hivers sont longs et froids, les étés, courts et chauds. Le Yukon est soumis aux conditions climatiques les plus variables en Amérique du Nord et neuf régions climatiques distinctes peuvent être reconnues. Presque toute la surface du Yukon est recouverte de pergélisol permanent, discontinu ou clairsemé. Les cryosols sont dominants dans les zones de pergélisol permanent du nord, alors que les brunisols dominent dans le sud où le pergélisol est discontinu ou clairsemé. Les principaux types de végétation du Yukon sont ceux de la toundra arctique, de la toundra alpine, de la taïga, ou des forêts subarctique, boréale ou buissonneuse subalpine. La toundra arctique et la taïga dominent dans la partie nord du territoire, alors que la forêt boréale et la forêt buissonneuse subalpine caractérisent la partie australe. La toundra alpine se rencontre en altitude dans tout le territoire. On reconnaît maintenant cinq écozones distinctes et 23 écorégions au Yukon. De plus, un grand nombre d’habitats d’intérêt particulier pour les entomologistes se retrouvent un peu partout dans le territoire: sources, tourbières, plateaux salés, dunes de sable, ainsi que de nombreuses zones xériques à communautés Artemisia frigida-herbacées. Au cours du Pléistocène, le Yukon est resté en grande partie déglacé et formait une partie de la Béringie, la plus grande zone refuge du nord au cours des dernières glaciations. Le Yukon est donc une zone biogéographique d’intérêt primordial dans la région néarctique. Introduction The Yukon Territory occupies an area of 482 681 km2 (Oswald and Senyk 1977) in northwestern Canada (Fig. 1). It is bounded on the north by the Beaufort Sea. In the west along 141°W longitude it abuts Alaska, and along 60°N latitude it meets British Columbia. The eastern boundary with the Northwest Territories is an irregular line following the height of land from about 136°30′W longitude in the north to about 124°W longitude in the south. The Yukon is a rugged territory of mountains, rough plateaus and valleys. Most of the southern three-fifths of the Territory is above 914 metres in elevation (Fig. 2). Only a few major roads traverse the complex terrain (Fig. 3). However, these roads cross the Territory from east to west, and from north to south. In the process they provide access to the major pp. 13 – 57 in H.V. Danks and J.A. Downes (Eds.), Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa. 1034 pp. © 1997 14 G.G.E. Scudder FIG. 1. Geographical reference map for the Yukon. subarctic and boreal ecozones and to more than half of the distinctive ecoregions in the Yukon. There are only about 12 major settlements (Fig. 3), with all but one of them in the southern half of the Territory and connected by the major roads. The one exception is Old Crow in the north, which can be reached by air. Physiography and Geology The physiography of the Yukon has been considered by Bostock (1948, 1961, 1970) and Tarnocai et al. (1993). The geology of the Territory is described in detail by Gabrielse and Yorath (1992) and summarized in map form by Clague (1989a). There are three major physiographic regions in the Yukon (Fig. 4), which have their own distinctive geology (Fig. 6). The Yukon Coastal Plain is part of the Arctic Coastal Plain physiographic region, and is formed of little-deformed sediments and volcanics, mainly of Mesozoic and Cenozoic ages. The Yukon Coastal Plain is a narrow strip of land along the Beaufort Sea, and includes Herschel Island (Fig. 4). The plain, which decreases in elevation to the west, is largely an erosional surface cut into Tertiary sandstone and shale, that is covered with a thin veneer of recent sediment. The unglaciated area west of Herschel Island is an almost flat, lacustrine plain, incised by streams and rivers, with the coastal scarp interrupted by alluvial fans and floodplains at the mouth of the Malcolm River and Firth River (Fig. 13). The glaciated area to the east of Herschel Island, largely covered with late Wisconsinan morainal materials, is spotted with lakes and ponds of thermokarst origin (Rampton 1982). The northeastern part of the Yukon, which includes the Peel Plateau, is part of the Interior Plains physiographic region, and is formed of thick, flat-lying Phanerozoic strata. The Peel Plateau is a striking and rather even land surface lying between the eastern front of the Richardson Mountains and the northern front of the Mackenzie Mountains (Fig. 5). Environment of the Yukon 15 FIG. 2. Yukon showing territory above 914 m elevation. In the extreme southwest corner is a broad, shallow depression, the Bonnet Plume Basin, lying between the Peel River and the first slopes of the mountains to the south. The majority of the Yukon lies in the Cordilleran physiographic region, a large mountain belt of deformed and metamorphosed sedimentary and volcanic rocks (Fig. 6) of mainly Phanerozoic and Proterozoic age. This Cordilleran region in the Yukon is divisible into three major physiographic subdivisions (Bostock 1948) (Fig. 4). To the north, the eastern subdivision, composed almost entirely of folded sedimentary strata, includes the British Mountains, the Richardson Mountains and the northern part of the Mackenzie Mountains (Fig. 5). The British Mountains (Fig. 7), which are the eastern extension of the Brooks Range crossing northern Alaska, reach an elevation of 1680 m near the Alaska/Yukon border, and have steep-sided and sharp-crested peaks and ridges. The Richardson Mountains (Fig. 8) 16 G.G.E. Scudder FIG. 3. Major settlements and major roads in the Yukon. form a belt of closely spaced hills with smooth profiles, a profile also seen in the Canyon Ranges section of the northern Mackenzie Mountains, so named because of the numerous canyons contained therein. In the southwestern part of the Yukon, the western subdivision (Fig. 4), composed of plutonic and volcanic rocks, includes the Icefield Ranges of the St. Elias Mountains and the Kluane Ranges (Fig. 9). The Icefield Ranges comprise the main body of the St. Elias Mountains and include many peaks over 4265 m, with Mt. Logan attaining 6050 m. This area is mainly snow and ice, with some nunatak areas evident in summer. Many of the Environment of the Yukon 17 FIG. 4. Main physiographic areas of the Yukon. glaciers reach down into the Alsek River valley, with the Kaskawulsh Glacier sending branches down to the Slims River valley (Fig. 10). The Kluane Ranges form a rugged frontal ridge to the north of the St. Elias Mountains, and have narrow serrated summits rising to over 2130 m, alpine glaciers, and steep talus slopes. Between these eastern and western physiographic systems is the interior subdivision (Fig. 4), composed of a mixture of volcanic, sedimentary and metamorphic rocks with intrusive outcropping. This interior system, principally comprising the Yukon Plateau and forming the Yukon River drainage, is composed of a central series of mountains, namely the Ogilvie, Wernecke, Selwyn and Logan Mountains, with the Pelly Mountains 18 G.G.E. Scudder FIG. 5. Major mountain ranges, plateaus, plains and basins of the Yukon. somewhat offset to the south (Fig. 5). North and south of these mountains is a series of plateaus, parts of the Yukon Plateau. The Ogilvie Mountains are 1525–1830 m in elevation, have a rugged aspect, and as a whole, consist of long, branching ridges connecting precipitous peaks, flanked by deep valleys. The Wernecke Mountains, which reach over 2130 m, were glaciated and contain part of the Mackenzie-Yukon divide; the majority drain north to the Peel River (Fig. 13). The Selwyn Mountains, with peaks up to 2515 m, run almost parallel with the Mackenzie Mountains and are an area of high precipitation. The Logan Mountains, especially in the south, are compact and rugged, with elevations over 2000 m, and with small Environment of the Yukon 19 FIG. 6. Generalized map of the geology of the Yukon (after Oswald and Senyk 1977). glaciers and ice fields. The Pelly Mountains occupy a large, wedge-shaped area in the southern part of the Yukon Plateau, and have a high rugged backbone some 290 km long and 65 km wide. The highest peaks are over 2000 m in elevation, and there are small alpine glaciers. Large U-shaped valleys run deep into all ranges of the Pelly Mountains, and some, such as the valleys of the Big Salmon and Magundy Rivers, traverse the ranges completely. North of the Ogilvie Mountains is the extensive Porcupine Plain and Porcupine Plateau. The Porcupine Plateau is an area of rolling, widely spaced hills, generally above 300 m, that lies between the Old Crow and Porcupine Rivers, and contains the Old Crow Range. The Porcupine Plain is a long, shallow depression between the Porcupine Plateau to the south and the British and Richardson Mountains to the north. The northern part, the Bell Basin, interrupted by a prolongation of the Keele Ranges, forms the Old Crow Plain, a large, flat 20 G.G.E. Scudder FIG. 7. British Mountains, Firth River valley (photograph by S.G. Cannings). FIG. 8. Richardson Mountains, “Erebia Creek” area (photograph by S.G. Cannings). Environment of the Yukon FIG. 9. Kluane Range from Outpost Plateau (photograph by S.G. Cannings). FIG. 10. Kluane Range and Slims River valley from Sheep Mountain (photograph by R.A. Cannings). 21 22 G.G.E. Scudder area thickly spotted with lakes and ponds. The southern part of the Porcupine Plain, termed the Eagle Plain, is an extensive flat, unglaciated area, some 190 km long and 95 km wide, now traversed by the Dempster Highway. The Tintina Trench runs through the centre of the Yukon Plateau, arising in the Pelly Mountains in the southeast and then extending straight towards the northwest through Ross River, and then just south of Mayo and just north of Dawson. Throughout its length it has areas with well defined walls, varies in width from 4.8 km – 22.5 km, and the floor ranges from 425–823 m in elevation. North of the Tintina Trench, the Yukon Plateau comprises the Stewart Plateau, the Macmillan Plateau and the Pelly Plateau. The Stewart Plateau forms a series of rather flat tablelands between the Macmillan River and the southern fronts of the Ogilvie and Wernecke Mountains. The steep-sided, steeply cut, broad valleys through the plateau are typical of unglaciated terrain. The Macmillan Plateau, which extends from near Ross River to the Macmillan River, forms a broken, mountainous area. Pleistocene ice filled the valleys, but many of the mountainous areas between them, particularly in the northwest, stood above the glaciers as nunataks (Bostock 1948). The Pelly Plateau is a glaciated, rolling upland, 1220 –1525 m in elevation, with entrenched valleys that dissect the plateau. South of the Tintina Trench and the Pelly Mountains the southern part of the Yukon Plateau comprises the Lewes Plateau, the Nisultin Plateau and the Teslin Plateau. The Lewes Plateau is a broad depression south of the western ranges of the Pelly Mountains. The Nisultin Plateau lies east of Teslin Lake, while the Teslin Plateau lies west of Teslin Lake. The Shakwak Trench runs diagonally northwest to southeast, parallel with the Tintina Trench, and forms the northern boundary of the Kluane Ranges, and the southern boundary of the western part of the Yukon Plateau. The northernmost section of this part of the plateau constitutes the largely unglaciated Klondike Plateau. The latter is a gently undulating area with a maze of deep, narrow valleys separated, by long, smooth-topped ridges and mountains, the most conspicuous forming the Dawson Range. The more southern section, constituting the Kluane Plateau, is divided into several distinct physiographic parts. The southeasternmost portion, east of the Takhini Valley, is some 1675 m in elevation, with peaks 1980 –2134 m high. Northwest of the Takhini Valley, the plateau consists of two topographic basins, the Aishihik Basin and the Wellesley Basin. The Aishihik Basin, at 915 –1220 m, contains Aishihik Lake and has extensive areas of smooth rolling plateau and some isolated mountainous areas up to 1830 m. The Wellesley Basin, at 610 m, contains Wellesley Lake, which is surrounded by innumerable kettle-holes occupied by ponds and small lakes. Between the Aishihik and Wellesley Basins, the Yukon Plateau forms a wide upland composed of two broad mountainous ridges trending northwest, the Ruby and Nisling Ranges, separated by a high, shallow trough parallel with the Shakwak Trench (Bostock 1948). The Ruby and Nisling Ranges, reaching 2134 m in the former, and 1829 m in the latter, are composed of granitic intrusions. Glacial History During the late Wisconsinan, most of Canada was ice covered. The continental Laurentide ice extended from its eastern centre to cover most of north-central and northeastern North America (Mickelson et al. 1983). To the northwest it extended no farther than a line running roughly along the eastern slopes of the Richardson Mountains and then northwest- Environment of the Yukon 23 erly parallel to the Yukon coast to a point near the coast southwest of Herschel Island (Ritchie 1984; Hughes et al. 1989). The Cordilleran ice sheet, which was centred in British Columbia, in the late Wisconsinan extended south into northern Washington, Idaho and Montana (Waitt and Thorson 1983), and covered virtually all of British Columbia (Clague 1989b), much of southern Alaska (Hamilton and Thorson 1983), and the southern Yukon (Hughes et al. 1989). Mountain glaciers also formed at higher elevations in northern Alaska, especially along the Brooks Range (Heusser 1983; Porter et al. 1983). At various times during the Pleistocene, the Laurentide and Cordilleran ice sheets were contiguous, or nearly so, along the western margin of the Great Plains, but during many of the glacial advances, growth of the two ice sheets evidently was out of phase (Ryder 1983; Clague 1989c). An ice-free corridor existed between the ice sheets at various times, but this was a variable and intermittent entity that existed only in narrow windows of Quaternary time (Schweger 1989). Central Yukon and central Alaska remained unglaciated throughout the Pleistocene (Hopkins 1967; Hopkins et al. 1982), as they lie in a dry belt northeast of the high St. Elias Mountains and Alaska Range, where most of the moisture in Pacific air masses is precipitated (Prest 1970). This unglaciated area (Fig. 11) was by far the largest refugium in the north during the Pleistocene. Together with the emergent Bering and Chukchi seas, at times it also formed an unbroken corridor for exchange of Palaearctic and Nearctic flora and fauna (Matthews and Telka 1997). The palaeoecology of this ice-age Beringian refugium is somewhat controversial, and is discussed in detail by Schweger (1997). Bostock (1966) inferred four advances of the Cordilleran ice sheet in the southern and central Yukon, with each successive advance being less extensive than its predecessor. Few readily recognizable landforms remain from the first two advances, and so the limits are poorly known. However, marginal features of the penultimate (Reid) glaciation terminating in the Yukon Plateau some 46 600 yr B.P. are moderately well preserved, and those of the final late Wisconsinan (McConnell) glaciation, that terminated 13 000 –14 000 yr B.P. are very well preserved (Hughes et al. 1989). Near the limits of successive glaciations, numerous peaks and plateaus were isolated as nunataks (Hughes et al. 1983). Figure 12 depicts the glacial limits of ice advance in the Yukon as summarized by Oswald and Senyk (1977). Ice covered the whole of the southern and eastern Yukon, with lobes extending westward to the Tintina Trench, Bonnet Plume Basin, and the Arctic coastal plain. Alpine valley glaciers, contemporary with the continental ice masses, originated locally in the Ogilvie, Wernecke, British, and possibly, Richardson Mountains (Hughes 1972; Hughes et al. 1969). The unglaciated portion of the Yukon consists of most of the Klondike, Porcupine and Arctic plateaus, the Porcupine Basin and portions of the Ogilvie, Wernecke and British Mountains, as well as the western slopes of the Richardson Mountains (Oswald and Senyk 1977). It is these unglaciated refugial areas that are central to the discussion of Yukon biogeography, and indeed the present patterns in the fauna and flora of Canada (Scudder 1979), and the origins of the whole North American biota (Downes and Kavanaugh 1988). Drainage Basins The Yukon is drained by rivers that run to both the Pacific and Arctic Oceans. There are five major watersheds, draining to the Beaufort Sea, to the Bering Sea, to the Yukon River, the Alsek River, and the Mackenzie River (2 subsections) (Fig. 13). The largest 24 G.G.E. Scudder FIG. 11. Wisconsinan glaciation in eastern Beringia (redrawn after Heusser 1983 and Hopkins et al. 1982). drainage area is the Yukon River watershed comprising about 66% of the Territory and draining via Alaska to the Bering Sea. Major tributaries include the Stewart River, the Pelly River, the Klondike River, the Donjek River and the White River. Many large lakes lie within the Yukon River watershed, including Teslin Lake, Tagish Lake (Fig. 14), Bennett Lake, Kluane Lake and Lake Laberge. The Porcupine River (Fig. 15) watershed is also part of the Yukon system. Major tributaries include the Eagle River, Bell River, Bluefish River and Old Crow River, the latter draining the Old Crow basin. In the south, the Alsek River watershed occupies about 4% of the Yukon and drains to the Pacific Ocean via Alaska. Major tributaries include the Aishihik River, the Dezadeash River, the Kaskawulsh River and the Dusty River. Aishihik Lake, Dezadeash Lake and Pine Lake lie in the watershed. In the southeast corner of the Territory, the Liard River watershed drains about 12% of the Yukon. Frances Lake, Simpson Lake and Watson Lake lie in this watershed. The Liard River is a major tributary of the Mackenzie River. The Peel River is another tributary of the Mackenzie River and drains most of the Wernecke Mountains, the southwestern portion of the Richardson Mountains and the northwestern part of the Ogilvie Mountains. The Peel watershed drains about 14% of the Yukon, including the Bonnet Plateau Basin. The Firth River, Malwym River, Babbage River and Blow River plus a number of smaller streams, occur on the Arctic coastal plain and drain into the Beaufort Sea. These and other northerly flowing rivers are frequently hampered by the freezing of water at downstream points, often resulting in the rivers overtopping their banks, inundating the surrounding areas. In contrast, the southerly flowing rivers usually freeze in a progressively downstream direction. Environment of the Yukon 25 FIG. 12. Glacial limits in the Yukon (after Oswald and Senyk 1977). The glacial history of these major drainage basins in the Yukon has been discussed by Lindsey et al. (1981). They point out that most basins have had a complex postglacial drainage history. The Peel River, for example, has had a number of drainage reversals during the Pleistocene glaciations. Lindsey et al. (1981) have also documented the physical and biological limnology of many of the lakes in the Yukon. They detail the chemical composition, oxygen and temperature regimes, morphometry, zooplankton and fish fauna of these lakes. Most of the larger lakes are located in the southern part of the Yukon; Kluane Lake is the largest, covering 409.5 km2 with a maximum known depth of 82 m. The lakes in the northern part of the Territory are small, usually less than 5 km2. 26 G.G.E. Scudder FIG. 13. Major rivers, lakes and drainage basins in the Yukon (redrawn after Wahl et al. 1987). Climate Yukon climate is the most variable in North America (Wahl et al. 1987). It is typified by long cold winters and short warm summers, with the annual mean daily temperature below freezing (Fig. 16). It holds the record on the North American continent for extreme low, −62.8°C at Snag. January is usually the coldest month (Fig. 17) and July the warmest (Fig. 18), although the highest maximum temperatures may occur in August. The complexity of the terrain Environment of the Yukon FIG. 14. Tagish Lake, Windy Arm (photograph by R.A. Cannings). FIG. 15. Porcupine River near Old Crow (photograph by R.A. Cannings). 27 28 G.G.E. Scudder FIG. 16. Annual mean daily temperature isotherms in the Yukon (redrawn after Wahl et al. 1987). produces complexity in the climate, with a great deal of local and regional variation. The many closed cells in the southern half of the Territory (Figs. 17, 18) reflect the mountainous terrain, with the flatter terrain in the north being reflected in the more sweeping isotherms. In general, the magnitude of the temperature extremes increases from southeast to northwest. The general circulation pattern in the north is constant, with dominant west to east progression of pressure systems. This results in storm tracks moving from west to east across the whole territory. In winter, a major low pressure centre prevails over the northeastern Pacific, and offshoots of this centre move across the Bering Strait into the Beaufort Sea to Environment of the Yukon 29 FIG. 17. January mean daily temperature isotherms in the Yukon (redrawn after Wahl et al. 1987). affect the Yukon, resulting in frequent storms moving along the Arctic coast. In contrast, by January, a major cell of high pressure prevails over the central Yukon-Mackenzie Mountain sector (Fig. 19). This tends to stabilize the weather, giving clear skies and low temperatures. In the summer, the most persistent feature of the pressure pattern is a large high-pressure cell at mid-latitudes (Fig. 20), giving sunny weather and warm temperatures in the south. The southern half of the Yukon is subject to very changeable weather in the fall and winter as a result of storms that move in from the Gulf of Alaska across the St. Elias-Coast range. In the southwest, the prevailing water-laden clouds moving in from the Pacific leave most of their precipitation on the windward side of the St. Elias Mountains, so that the 30 G.G.E. Scudder FIG. 18. July mean daily temperature isotherms in the Yukon (redrawn after Wahl et al. 1987). leeward slopes and areas in the Yukon have little precipitation (Fig. 21). This results in the area around Carcross and Kluane having almost desert-like conditions. In the large central area of the Yukon, the precipitation is quite variable, depending on proximity, elevation and position of mountains. July, August or September are normally the wettest months. Wahl et al. (1987) recognize nine distinct climatic regions in the Yukon, as shown in Fig. 22 and outlined below. Arctic Slope. This is a region of low annual precipitation, the total of less than 200 mm occurring as rain or drizzle during the summer. There is very little precipitation in the winter, so snow depths are normally below 25 cm. Winters are characteristically long, lasting from Environment of the Yukon 31 FIG. 19. Mean sea-level pressure isobars for January in the Yukon (redrawn after Wahl et al. 1987). October to early June, but minima reach only from −50° to −55°C, with February the coldest month. Summers are cool and changeable, and dependent on whether the wind is blowing seaward or onshore. Warm spells up to 30°C are infrequent. Winds are strong. Northern Mountains. Annual precipitation is between 300 and 400 mm, most occurring in summer. Winter temperatures tend to be mild owing to inversion effects. Summers tend to have cold changeable weather because of proximity to the frozen Beaufort Sea. The region is subject to strong winds, especially in the region of Wright Pass on the Dempster Highway and in the headwaters of the Blow River. 32 G.G.E. Scudder FIG. 20. Mean sea-level pressure isobars for July in the Yukon (redrawn after Wahl et al. 1987). Porcupine-Peel Basin. In this region annual precipitation is low, and in the 200 –300 mm range. Most precipitation is from summer convective activity, although the region often is affected by well-developed rainstorms passing through. This region has characteristic prolonged cold spells in winter, particularly on the valley floors. Summers are short but variable, often with quite warm spells. Ogilvie-Mackenzie Mountains. Annual precipitation is moderate to heavy in this region, ranging from over 700 mm in the southeast to 400–500 mm in the Ogilvie Mountains. Winter temperatures are moderate, and summers are cool. Central Yukon Basin. This is a region with moderate annual precipitation, ranging from 300 – 400 mm. Most precipitation occurs in summer as showers. Temperatures are quite Environment of the Yukon 33 FIG. 21. Annual mean total precipitation isohyets for the Yukon (redrawn after Wahl et al. 1987). variable, and summers can be quite warm. There can be prolonged cold spells in winter, when well-organized storm centres tend to skirt the region. Liard Basin. In the Liard Basin, there is a moderate annual precipitation of 400 – 600 mm, with a substantial amount in the form of snow. This region has more days with recorded precipitation than anywhere else in the Yukon. There can be protracted cold spells in winter, but the summers are warm. 34 G.G.E. Scudder FIG. 22. Climatic regions of the Yukon (redrawn after Wahl et al. 1987). Pelly-Cassiar Mountains. This region is characterized by relatively high annual precipitation, ranging from 500 –700 mm. The heaviest precipitation occurs in fall and early winter, and coincides with the time that active storm tracks traverse the regions. As a result, winter temperatures are less severe than elsewhere in the Yukon, and there are frequent midwinter mild spells. Summer temperatures are relatively cool. Upper Yukon-Stikine Basin. This region receives less than 300 mm annual precipitation, and many localities, such as Carcross and Kluane Lake, as little as 200 mm. The temperature Environment of the Yukon 35 regime is typically continental, with a great deal of variability on both a daily and seasonal basis. The proximity to the Pacific Ocean results in more frequent midwinter mild spells than occur elsewhere in the Yukon. St. Elias-Coast Mountains. The climate in this region is transitional between the wet maritime climate of the windward coastal area and the dry continental climate northeast of the massive mountains. Annual total precipitation is up to 4000 mm on the coast, but decreases to under 300 mm in the Shakwak Trench. Precipitation is mostly in the form of snow above 3000 m, and major snow accumulation occurs between 1500 and 3000 m. This results in the extensive network of glaciers in the Icefield Ranges, within the St. Elias Mountains. Valley glaciers extend downslope from accumulation zones to the 900 –1200 m level. At low elevations, liquid precipitation occurs from mid-May to September. Temperatures in the region vary widely as a result of the complex topography and storm paths from the Gulf of Alaska. The mean annual temperature within the St. Elias Mountains above 2500 m is −10°C to −15°C. The region is one of the windiest in the Yukon. Permafrost Permafrost is ground (soil or rock) that remains at or below 0°C over at least two consecutive winters and an intervening summer. In general, the overall distribution of permafrost is related to air temperature (Hughes et al. 1983). The extent of permafrost in the Yukon has been described by Brown (1970, 1978), Judge (1973) and French and Heginbottom (1983), and is depicted in Fig. 23. In the extreme north, there is a zone of continuous permafrost. Here permafrost occurs everywhere beneath the exposed land surface, being absent only below the major rivers and lakes. In this zone, the thickness of the frozen material is about 100 m at the southern boundary, and increases towards the north. The active layer, the upper part that freezes in winter and thaws in summer, generally extends to permafrost. Most of the Yukon has either discontinuous or scattered permafrost, where the permafrost is thinner and widely distributed, but not continuous beneath the land surface. The boundary between the continuous and discontinuous permafrost zone corresponds to a mean annual air temperature of −8.3°C (Brown 1978). To the south there is an area of scattered permafrost, where some areas have permafrost but others are free from permafrost. In these latter two permafrost areas, the active layer varies in thickness, and may or may not extend to the permafrost table. In the extreme southwest, in the St. Elias Mountains area, there is often continuous permafrost at high elevations, but it is discontinuous at lower elevations. The lower elevational limit of Cordilleran permafrost coincides approximately with the −1°C average air isotherm (Hughes et al. 1983). Soils The soils in the Yukon have been described by Hughes et al. (1983) and mapped by White et al. (1992). Soil development in the northern half of the Territory has been described in detail by Tarnocai et al. (1993), and soil classification is described by the Agriculture Canada Expert Committee on Soil Survey (1987). Cryosolic soils are the dominant soil type in the northern permafrost zone in the Yukon (Fig. 24), and in the Ogilvie and Richardson Mountains, as well as in the Dawson Range of the Yukon Plateau and other scattered higher elevations. Essentially such soils coincide with 36 G.G.E. Scudder FIG. 23. Permafrost areas of the Yukon (redrawn after Wahl et al. 1987). very cold temperatures throughout the year. They are characteristic of frozen organic soils or peatland terrain, where the thick organic layers insulate the frozen soil from the warm air of the summer. In the north, powerful freeze-thaw cycles lead to cryoturbation of the soils, the mixing and distortion of the soil horizons. Such Turbic Cryosols are associated with patterned ground features, such as earth hummocks, polygons and circles, that are typical of the Arctic slope. Zoltai and Tarnocai (1974) and Zoltai et al. (1978) indicate that cryoturbation has been active in arctic areas since deglaciation, but probably was initially absent from the taiga Environment of the Yukon 37 FIG. 24. General distribution of soil types in the Yukon (after Agriculture Canada Expert Committee on Soil Survey 1987). or subarctic forest areas to the south. However, when climate conditions became cooler about 4500 yr ago, cryoturbation became more widespread, but over the last 200 years there have been alternating active and dormant periods of cryoturbation (Zoltai 1975). The coarse-textured deposits along the Arctic coast are associated with Static Cryosols, cryosols that are mineral-rich, but not cryoturbated. Static Cryosols also dominate the Ogilvie Mountains. Brunisolic soils, with their typical weakly developed brown coloured B horizon, dominate the southern half of the Territory (Fig. 24). Such Brunisolic soils occur under coniferous forest where there is moderate soil development in association with cold temperatures, coarse parent material, and high rates of evaporation. 38 G.G.E. Scudder Most of the southern half of the Yukon has Eutric Brunisols, which are not strongly acid and have a well developed Ah horizon. However, in the southern Selwyn Mountains and the Logan Mountains, and other scattered areas such as the Coast Mountains and the Cassiar Mountains, Dystric Brunisols are found, brunisols which are strongly acid but lack a well-developed Ah horizon. Brunisolic gray luvisolic soils, with clay accumulation in the B horizon, are confined to the southeast corner of the Territory in the Liard River basin. These luvisolic soils develop in cool to cold soil climates, and typically occur on fine-textured glacio-lacustrine deposits. Chernozemic soils, with well-developed, base-rich, mineral-organic surface horizons (Ah), are grassland soils, and are not common in the Yukon. They occur on the warm, steep south-facing slopes in the south and central Yukon, particularly along the Yukon River system. Regosolic soils, with very weak horizon development, are also scarce, and are largely confined to the valley bottoms of the Peel and Yukon River system. There are extensive areas of the Yukon that lack soil development. Such rock outcrop areas occur mainly in the Richardson, Ogilvie, Selwyn, Logan and St. Elias Mountains. Vegetation The vegetation of the Yukon, like the rest of northern Canada, strongly reflects latitudinal and elevational temperature gradients (Hughes et al. 1983). The dominant vegetation types in the Yukon are arctic tundra, alpine tundra, taiga or subarctic forest, boreal forest and subalpine-shrub forest (Fig. 25). Arctic Tundra. Arctic tundra, which occurs as the dominant vegetation type in the low arctic Yukon Coastal Plain, is treeless, with a continuous vegetative cover (Fig. 26). Typical arctic tundra plants include Eriophorum vaginatum L. (sheathed cottongrass) and Carex aquatilis Wahlenb. (aquatic sedge) (Cyperaceae), Salix arctica Pall. (arctic willow) (Salicaceae), Cassiope tetragona (L.) D. Don (white arctic bell-heather) (Ericaceae), Rumex arcticus Trautv. (arctic dock) (Polygonaceae), and Dryas integrifolia M. Vahl. (arctic avens) (Rosaceae) (Wiken et al. 1981). In upland areas, the vegetation consists primarily of tussocks of cottongrass (Eriophorum spp.), interspersed with trailing shrubs and heaths. In low-lying depressions, sedges and mosses are dominant. Thus, in upland areas, such as the area between Fish Creek and Clarence Lagoon, the elevated and well drained central portion of Herschel Island (Fig. 27), and the higher morainal areas east of Herschel Island, the vegetation consists of expanses of tussock tundra dominated by Eriophorum vaginatum, Ledum decumbens (Ait) Lodd. (narrow-leaved Labrador tea) and Vaccinium vitis-idaea L. ssp. minus (Ludd.) Hult. (mountain cranberry) (Ericaceae), Betula nana L. ssp. exilis (Sukatsch.) Hult. (dwarf birch) (Betulaceae) and mosses (Wiken et al. 1981). Lowland areas, which are dominated by low-centre frost polygons, have cores covered by sedges and Sphagnum spp., along with Ledum decumbens, Andromeda polifolia L. (bog rosemary) (Ericaceae), Vaccinium vitis-idaea, Salix spp., and Betula nana forming elevated rims. High-centre polygons have tussocks of Eriophorum vaginatum in the elevated central portions, and mostly Carex aquatilis in the ice-wedge troughs (Wiken et al. 1981). Alpine Tundra. Alpine tundra (Fig. 28) occurs throughout the British-Richardson Mountains, the Ogilvie, Wernecke and Selwyn Mountains, the St. Elias Mountains, and scattered at higher elevations in the interior Cordilleran region of the Yukon, especially in the Pelly Mountains and Dawson Ranges (Fig. 4). Typical alpine tundra plants include Dryas Environment of the Yukon 39 FIG. 25. Distribution of main vegetation types in the Yukon (redrawn after Wahl et al. 1987). octopetala L. (mountain avens), Arctostaphylos alpina (L.) Spreng. (alpine bearberry) (Ericaceae), Saxifraga caespitosa L. (tufted alpine saxifrage) and S. tricuspidata Rottb. (prickly saxifrage) (Saxifragaceae) and crustose lichens (Wiken et al. 1981). Characteristics and composition of the vegetation of the arctic tundra vary with exposure, slope, soil type, moisture gradient and snow cover. Thus in the British Mountains, lower mountain slopes and ridge tops have patches and stripes of vegetation consisting of Dryas octopetala, along with Salix reticulata L. (net-veined willow), Saxifraga spp. (Saxifragaceae), Antennaria spp. (Asteraceae), Silene acaulis L. (moss campion) 40 G.G.E. Scudder FIG. 26. Arctic tundra, Philip Creek area, Babbage River (photograph by R.A. Cannings). FIG. 27. Well drained tundra on Herschel Island (photograph by S.G. Cannings). Environment of the Yukon 41 FIG. 28. Alpine tundra, Kluane Range, Outpost Mountain (photograph by S.G. Cannings). (Caryophyllaceae), Potentilla uniflora Lebed. (one-flowered cinquefoil) (Rosaceae), Papaver spp. (Papaveraceae) and Senecio cymbalaria Pursh (Asteraceae). On the other hand, toe slopes have a community with Cassiope tetragona, Dryas integrifolia, Betula glandulosa Michx. (ground birch), Salix reticulata, other Salix spp., Vaccinium uliginosum L. (bog bilberry) (Ericaceae), Empetrum nigrum L. (black crowberry) (Empetraceae), Dryas octopetala, and Hylocomium moss (Wiken et al. 1981). In the south Richardson Mountains, Ritchie (1982, 1984) notes that whereas uplands have a very characteristic, sparse, low tundra dominated by patches of Salix phlebophylla Anderss. (skeleton willow), Arctostaphylos alpina, Dryas octopetala, Oxytropis nigrescens (Pall.) Fisch. (Fabaceae), Betula glandulosa, Artemisia norvegica Fries. ssp. saxatilis (Bess. ex Hook.) Hall and Clem. (Asteraceae) and Hierochloe alpina (SW.) Roem. and Schult. (alpine holy grass) (Poaceae), calcareous rocks bear a very different tundra community. This consists of a low vegetation, dominated by Dryas integrifolia and Carex scirpoidea Michx. Hoefs et al. (1976) similarly note that in the alpine tundra on Sheep Mountain in the southwestern part of the Yukon, which occurs above 1500 m, the vegetation on calcareous soils is characterized by plant communities where Dryas integrifolia is predominant. Stanek et al. (1981) point out that in the Richardson Mountains sedge tussock tundra occurs on a variety of terrain types, but where the terrain is hummocky, a closed ground cover of Sphagnum, Polytrichum and Hylocomium mosses persists under widely spaced low shrubs and dwarf willow. On steep northern exposure slopes a lichen-shrub tundra exists, characterized by Cassiope tetragona. On moderate slopes where soil is extremely stony and organic layers are shallow, a lichen low-shrub tundra occurs, dominated by lichens and Betula glandulosa. 42 G.G.E. Scudder FIG. 29. Old Crow Flats, view to north from Old Crow Hills (photograph by R.A. Cannings). FIG. 30. Eagle Plain at Arctic Circle. Richardson Mountains in background (photograph by S.G. Cannings). Environment of the Yukon 43 FIG. 31. Boreal forest around Rainbow Lake, Shakwak Trench (photograph by R.A. Cannings). Taiga. This subarctic forest vegetation dominates the non-mountainous areas of the Porcupine Plain and Plateau, and the Interior Plains region. In the former, it covers much of the Old Crow Basin, Old Crow Flats (Fig. 29) and Eagle Plain regions (Fig. 30) and typically is an open woodland with very stunted stands of Picea mariana (Mill.) B.S.P. (black spruce) and Larix laricina (Du Roi) K. Koch (tamarack or larch) (Pinaceae) with lesser quantities of Picea glauca (Moench) Voss (white spruce), and a ground cover of Betula nana, willow (Salix spp.), ericaceous shrubs, cottongrass (Eriophorum spp.), lichens and mosses. In the Eagle Plain, Stanek et al. (1981) note that on gentle western or southwestern slopes an open canopied lichen-spruce taiga occurs, with lichen dominant in the understory and the Picea mariana characterized by a short, ragged growth form. Complex open black spruce-tall shrub taiga is found on steep southern exposures, with Betula glandulosa, Ledum decumbens, and Vaccinium vitis-idaea frequent. Where hummocks are present, a variety of microsites are present in the hummocks and inter-hummock troughs, the terrain as a whole supporting an open spruce taiga with clumps of stunted black spruce, surrounded by low ericaceous shrubs, particularly Ledum groenlandicum Oeder and Vaccinium vitis-idaea. In the Interior Plains region, occupying much of the Peel River Plateau and adjacent Fort MacPherson Plain, the taiga vegetation is similar, but with conspicuous occurrence also of Betula papyrifera Marsh (paper birch), Populus balsamifera L. (balsam poplar) (Salicaceae) and Alnus crispa (Drylander ex Ait.) Pursh (green alder) (Betulaceae). Boreal Forest. Boreal white and black spruce ecosystems occur as the major forest type in the southeastern part of the Yukon (Fig. 31), and occur elsewhere in the southern half of the Territory at lower elevations from valley floors to the bottom of the subalpine forest. The elevational limit is at 100 m in the extreme south around Sheep Mountain in Kluane National Park (Hoefs et al. 1976), and at 800 m along the Dempster Highway in the central Yukon (Kojima and Brooke 1986). 44 G.G.E. Scudder This forest is dominated by Picea glauca in most late seral and climatic climax stands. It is usually accompanied by a number of other trees that have a broad boreal distribution across Canada, and have moved into the Yukon from the east in postglacial time (Delcourt and Delcourt 1987). These include Picea mariana, Populus tremuloides Michx. (trembling aspen), P. balsamifera, Larix laricina and Betula papyrifera (these important plants are illustrated in Stanek and Orloci 1987). However, in the Yukon, two other conifers of western origin are major components of the white and black spruce boreal forests, namely Pinus contorta Dougl. (lodgepole pine) and Abies lasiocarpa (Hook.) Nutt. (alpine fir) (Pinaceae). Species composition and vegetation characteristics vary according to climate, aspect, soil type and drainage, with dense black spruce-moss communities developing in imperfectly drained sites (Meideniger and Pojar 1991). Stanek (1980) has described the vegetation types in the Liard Basin, noting that here Picea glauca is dominant in primary and old secondary stands. Populus tremuloides forms secondary stands on finer soils in warm sites, while Picea mariana and Larix laricina are characteristic in some wetlands, but Salix shrub communities occur in others. Betula papyrifera occurs sporadically. Bonnor and Oswald (1989) in their reconnaissance inventory of Yukon forest resources point out that nearly all of the land below the alpine zone in the southern Yukon has been burnt in the last 300 years, much of it several times; because Picea glauca takes at least 25 years to produce viable seed, lacks serotinous cones, and does not reproduce by layering, this tree is sparse or virtually absent in many areas. Indeed, some tracts of land in the southern Yukon currently support only shrubs owing to the elimination of tree species and their propagules as a result of repeated fires at short intervals (Oswald and Brown 1990). Pinus contorta and Populus tremuloides owe their existence for the most part to the occurrence of fire, because they are initial invaders in recolonization. Pinus contorta possesses serotinous cones which release lots of seeds when opened by fire and can start reproducing in less than 10 years; it thrives best on relatively coarse, textured, moderately well to excessively drained soils that can have a moisture deficit during the summer (Bonnor and Oswald 1989). Populus tremuloides, on the other hand, reproduces after fire largely by suckering, especially on finer textured, moist or somewhat alkaline sites. Oswald and Brown (1986), in their study of the forest around Lake Laberge, describe over 50 distinct plant communities involving Picea glauca, P. mariana, Pinus contorta and Populus tremuloides. Douglas (1974) described the plant communities in the boreal white and black spruce forest ecosystem in the Alsek River region in the southwest Yukon, noting that here is a mosaic of forest, shrub and herb communities. This mosaic results from the variable climate within the region and frequent disturbance by both fire and geomorphological (mainly fluvial) processes. These factors create twenty-one distinct community types, which is extremely diverse for a boreal forest area. Subalpine Forest-shrub. The subalpine zone, with a characteristic spruce-willow-birch ecosystem (Meidinger and Pojar 1991), occurs between the boreal white and black spruce forest and the alpine tundra zone. Krajina (1975) recognized two elevational subzones, a lower forested subzone and an upper shrub subzone. In the former, tree cover is like that in the boreal white and black spruce zone, but the occurrence of Betula glandulosa in forested as well as the non-forested upper subzone indicates the subalpine habitat. Kojima and Brooke (1986) report that the shrub subzone on the Dempster Highway is characterized by the shrub thickets of Betula glandulosa and Salix spp. (mostly S. planifolia Pursh ssp. pulchra (Cham.) Argus). B. glandulosa is common on moderately to well-drained habitats near and above treeline, but is gradually replaced by Salix spp., Environment of the Yukon 45 especially S. planifolia ssp. pulchra and S. glauca L., in more moist habitats near the base of slopes or valley bottoms, where willow may completely dominate the vegetation. In places in the subalpine zone Picea mariana occurs as Krummholtz. Stanek et al. (1981) describe this subalpine shrub vegetation type on the gently sloping terrain of the Eagle Plain as a dwarf shrub tundra, noting that characteristic species in addition to B. glandulosa and S. planifolia ssp. pulchra, and irregularly scattered P. mariana, include Arctostaphylos alpina, A. rubra (Rehd. and Wilson) Fern., Empetrum nigrum L. (Empetraceae), Ledum decumbens, Rubus chamaemorus L. (Rosaceae), Petasites frigidus (L.) Fries. (Asteraceae), Vaccinium uliginosum and V. vitis-idaea, sedges such as Carex spp. and Eriophorum vaginatum, the grasses Arctagrostis latifolia (R. Br.) Griseb. and Calamagrostis canadensis (Michx.) Beauv. (Poaceae) and lichens. In the Alsek River region, the subalpine zone, which occurs up to 1080 –1370 m, is dominated essentially by tall (up to 3 – 4 m) shrubs, mainly Salix spp. with scattered Picea glauca (Douglas 1974). In the Sheep Mountain area of Kluane National Park, the subalpine zone occurs between 1100 and 1500 m (Hoefs et al. 1976), with two distinct plant associations. One occurs on mesic sites and is dominated by Betula glandulosa, Salix glauca, S. reticulata and Arctostaphylos rubra. The other occurs on sites which are more hygric, with longer snow duration, and is dominated by B. glandulosa, S. glauca, S. lanata L., S. reticulata, Arctostaphylos rubra, Cassiope tetragona and Vaccinium uliginosum. Notable Habitats A number of notable habitats in the Yukon are likely to have special and distinctive insect faunas. These habitats include springs, peatlands, saline flats, sand dunes and xeric Artemisia frigida-grass communities. Springs. There are a number of major springs in the Yukon (Souther and Halstead 1969). The hotsprings at Takhini and the sulphur springs at Engineer Creek on the Dempster Highway are just two examples. Unfortunately, there appears to be no detailed and comprehensive study of the characteristics and biota of the Yukon springs. However, one might expect Diptera, Coleoptera, and Trichoptera to occur in such habitats, as these are typical spring inhabitants (Danks and Williams 1991). Peatlands. There are extensive peatlands in the Yukon (Natural Wetlands Working Group 1988; Tarnocai et al. 1995) (Fig. 32). In the Yukon coastal plain, permafrost underlies all the low arctic wetlands at shallow depths, and the wetlands are induced by the development of ice wedges in the ground. These result in either low-centre polygons with soil- and peat-formed dams around shallow pools (Fig. 33), or high-centre polygons that result from peat accumulation (Zoltai 1987). In the subarctic alpine tundra and taiga areas, the predominant form of the peatland is a peat plateau, wherein a perennially frozen peatland is elevated by permafrost above the regional water table (Zoltai 1987). Zoltai (1987) notes that such peat plateaus are dry on the surface, except for small damp depressions, and generally have vegetation that resembles that of the surrounding uplands. However, the surrounding fens are rich in nutrients, and support mainly sedge and moss vegetation communities. In the boreal forest, the peatlands are mostly in the form of fens, with bogs uncommon (Zoltai 1987). Fens are waterlogged peatlands, with water generally rich in mineral nutrients, and with pH 5.5 – 7.5. Bogs in contrast are peat-covered wetlands, generally lacking nutrients and with acidic waters, usually with pH , 4.5 (Zoltai 1987). 46 G.G.E. Scudder FIG. 32. Distribution of the major peatland areas of the Yukon (after Tarnocai et al. 1995). Although data on occurrence of insects in fens is very scanty, a number of Odonata and Coleoptera, in particular, are restricted to bogs (Danks and Rosenberg 1987). Because peatlands are extensive in the Yukon, it is expected that peatland-adapted species in at least these two groups of insects will be common in the Territory. Saline Flats. Saline ponds (Fig. 34) and salt-crust basins occur in the Haines Junction area (Stanek 1980). The vegetation in the immediate vicinity of these salt-crust basins contains mostly halophytes. To survive in saline waters, insects must have special osmotic and ionic regulatory physiological adaptations (Shaw and Stobbart 1963). Coleoptera, Diptera and Heteroptera are the predominant insect groups in saline waters (Scudder 1969, 1976; Foster and Treherne 1976). Although inland saline ponds are few in the Yukon, some saline tolerant species can be expected in these habitats. Environment of the Yukon 47 FIG. 33. Low-centre polygon peatlands, Bluefish Basin southwest of Old Crow (photograph by R.A. Cannings). FIG. 34. Saline ponds west of Whitehorse (photograph by S.G. Cannings). 48 G.G.E. Scudder Sand Dunes. The main sand-dune areas in the Yukon occur in the south near Carcross and in the Alsek River region. Douglas (1974) describes two distinct herb communities in the dunes near the junction of the Alsek and Dezadeash Rivers. One occurs on the stabilized sand dunes and is characterized by a continuous but sparse cover of Carex sabulosa Turcz. ssp. leiophylla (Mack.) Pors. The other occurs on aeolian sands in the lee of the semistabilized dune formations and is dominated by Calamagrostis sp. Douglas (1974) found this Calamagrostis community to be the richest (floristically) herb community in the Alsek River region, with a total of 55 plant species. Included in these species are Artemisia norvegica, Thalictrum occidentale Gray (Ranunculaceae), Elymus calderi Barkworth (= Agropyron yukonense auct. plur. non Scribn. and Merr.), Festuca rubra L. (Poaceae), and Hierochloe hirta (Schrank) Borbas ssp. arctica G. Weim. This specialized habitat is poorly collected in the Beringian area, but one lepidopteran sand-dune restricted species is known to occur in the Carcross dunes (Lafontaine and Wood 1988). The sand-dune area may also share species with the xeric Artemisia frigida L.-grass communities. Xeric Artemisia frigida-grass Communities. Distinctive xeric habitats with Artemisia frigida-grass communities occur on warm and south-facing slopes in the south and central Yukon, particularly along the Yukon River, but also in the north on the Firth River and near Old Crow (Scudder 1993) (Fig. 35). These grassland habitats have a distinctive insect fauna (Lafontaine and Wood 1988; Scudder 1993). Douglas (1974) describes such an Artemisia frigida-Poa glauca Vahl community along the Alsek and Dezadeash River, on extremely dry and well-drained sites. The community is recognizable by an open cover of Artemisia frigida and Poa glauca, with the addition of Elymus calderi, Chamaerhodos erecta (L.) Bunge (Rosaceae) and Festuca brachyphylla Schultes and Schultes. There is also a sparse xerophytic cryptogamic stratum, with the mosses Ceratodon purpurens (Hedw.) Brid. and Tortula ruralis (Hedw.) Gaertn., Meyer and Scherb., and the lichens Physconia muscigena (Ach.) Poelt and Cornicularia aculeata (Schreb.) Hoffm. most prominent. Stanek (1980) reports that the pioneer community on regosols of the south-facing arid slopes of Sheep Mountain in Kluane National Park is an Artemisia frigida-Elymus calderiCalamagrostis purpurascens community. Further, Hoefs et al. (1976) note that such Artemisia fridiga-Elymus calderi grassland communities on Sheep Mountain develop mostly on very fine loess aeolian (frequently still active) regosols, in both the boreal white and black spruce as well as the subalpine spruce-willow-birch zones. Hoefs et al. (1976) distinguish five distinct plant associations within this community, which vary in the significance of individual species. The main association in the boreal white and black spruce zone on Sheep Mountain is an Artemisia frigida, Elymus calderi, Carex filifolia Nutt., Erigeron caespitosus Nutt. (Asteraceae), Penstemon gormanii Greene (Scrophulariaceae) association. An Artemisia rupestris L., A. frigida, Elymus calderi association occurs on colluvial material at the bottom of scree slopes in both the boreal forest and the subalpine areas between 1065 and 1370 m, and is the dominant grassland association at these elevations. In the subalpine zone, a Carex filifolia, Artemisia furcata Bieb., A. frigida, Oxytropis viscida Nutt., Calamagrostis purpurascens, Potentilla nivea L., Poa glauca association predominates. Similar Artemisia frigida-grass communities have been described on the arid southfacing slopes of the Yukon River in Alaska by Batten et al. (1979) and Kassler (1979). Kassler (1979) suggests that these Artemisia frigida-grass communities are remnants of an Environment of the Yukon 49 FIG. 35. Known occurrence of xeric Artemisia frigida in the Yukon (redrawn after Cody 1996): major roads are also shown. earlier late Pleistocene arctic steppe ecosystem. Whether this arid steppe-like community dominated Beringia in the late Pleistocene as claimed by Guthrie (1985, 1990), and is now representative of an extinct biome (Matthews 1976), or instead is a remnant of an earlier tundra-steppe mosaic (Ager 1982, 1983), or an upland xeric tundra (Schweger 1982), there is no doubt that this is a unique and special habitat in the Yukon. It contains a number of rare plant species (Batten et al. 1979), many grassland insects with southern affinities (Scudder 1993), and Pleistocene refugial endemic insects (Anderson 1984). G.G.E. Scudder 50 TABLE 1. The terrestrial ecozones and ecoregions of the Yukon (Ecological Stratification Working Group 1996). Ecozone Southern Arctic Taiga Plain Taiga Cordillera Boreal Cordillera Pacific Maritime Ecoregion Yukon Coastal Plain Peel River Plateau Fort MacPherson Plain Muskwa Plateau British-Richardson Mountains Old Crow Basin Old Crow Flats North Ogilvie Mountains Eagle Plains Mackenzie Mountains Selwyn Mountains Klondike Plateau St. Elias Mountains Ruby Ranges Yukon Plateau-Central Yukon Plateau-North Yukon Southern Lakes Pelly Mountains Yukon Stikine Highlands Boreal Mountains and Plateaus Liard Basin Hyland Highland Mount Logan Similar relics of the xerophytic vegetation of Beringia occur in northeastern Asia (Yurtsev 1982). It is certain that the Artemisia frigida-grass communities on warm, arid south-facing slopes are among the most interesting and notable habitats in the Yukon. Ecogeographic Regions, Ecozones and Ecoregions of the Yukon The various physiographic areas of the Yukon, together with their associated climate and vegetation, are major reference points for understanding and interpreting the distribution of the biota. Oswald and Senyk (1977) identified 22 ecoregions in the Yukon, areas of land where the vegetation, soils and permafrost reflect the regional climate. In 1987, I used the Oswald and Senyk (1977) ecoregion scheme, combined with Bostock’s (1948) physiographic divisions of the Yukon, to produce a map of 20 ecogeographic regions that could be used in the description and analysis of the distribution of the different insect groups in the Territory. This map (Fig. 36) is reproduced herein, because many authors in the Insects of the Yukon have referred to it in their individual chapters. More recently, the Ecological Stratification Working Group (1996) elaborating on earlier schemes describing terrestrial ecozones (Wiken 1986) and ecoregions (Ecoregions Working Group 1989), and using the methodology of Ironside (1991), have formalized an ecological framework for the whole of Canada. This procedure identifies 5 ecozones and 23 ecoregions in the Yukon (Table 1; Fig. 37). Ecozones are areas of the earth’s surface representative of large and very generalized ecological units characterized by interactive and adjusting abiotic and biotic factors (Ironside 1991). They have physiographic or macro-landforms, macro-climate, and broad physiognomic vegetation types. The ecozones in the Yukon, listed in Table 1 and depicted in Fig. 37, are briefly described below. There is major road access to the Taiga Cordillera and Boreal Cordillera ecozones, but not to the other three ecozones. Environment of the Yukon 51 FIG. 36. Ecogeographic regions of the Yukon: 1. Arctic Coastal Plain; 2. British Mountains; 3. Arctic Plateau; 4. Porcupine Plain (incl. Old Crow Plain, Old Crow Mts., N. Porcupine Plateau); 5. Richardson Mountains; 6. Eagle Plain (incl. S. Porcupine Plateau); 7. Peel Plateau (incl. Bonnet Plume Basin); 8. Ogilvie Mountains (incl. N. and S. Ogilvie Mountains); 9. Wernecke/Selwyn Mountains; 10. Yukon/Tintina (incl. Lewes Plateau, part of Klondike Plateau, and Tintina Trench); 11. Eastern Plateaus (incl. Stewart, Macmillan, and Pelly Plateaus); 12. Shakwak Trench (incl. Wellesley Basin); 13. Western Ranges (incl. Ruby, Nisling, and Dawson Ranges, part of Klondike Plateau). 14. Pelly Mountains; 15. Logan Mountains; 16. St. Elias/Coast Mountains; 17. Southern Lakes (incl. Aishihik Basin, Takhini Valley, Teslin Plateau, and Nisutlin Plateau); 18. Cassiar Mountains; 19. Liard Plain (incl. Dease Plateau); 20. Hyland/Liard Plateaus. 52 G.G.E. Scudder FIG. 37. The ecozones and ecoregions of the Yukon as recognized by the Ecological Stratification Working Group (1996). The Southern Arctic ecozone (A) in the Yukon coincides with the Arctic Coastal Plain physiographic subdivision of Bostock (1948) and was glaciated during early Wisconsinan and, to a lesser extent, during late Wisconsinan time (Oswald and Senyk 1977; Hughes et al. 1989). It lies within the zone of permanent permafrost and has arctic tundra vegetation dominated by sedge and cottongrass tussocks. Small to moderate-sized lakes are common on this Coastal Plain, and deep, medium-textured morainal material is extensive along the coast. 53 Environment of the Yukon TABLE 2. Comparison of the ecogeographic regions identified in Scudder (1987), with Yukon ecoregions in Ecological Stratification Working Group (ESWG) (1996). ESWG (1996) Scudder (1987) No. Ecogeographic region No. Ecoregion 1. Arctic Coastal Plain 1. Yukon Coastal Plain 2. 3. British Mountains Arctic Plateau 5. British-Richardson Mountains 5. British-Richardson Mountains 4. Porcupine Plain 6. 7. Old Crow Basin Old Crow Flats 6. Eagle Plain 9. Eagle Plain 7. Peel Plateau 2. 3. Peel River Plateau Fort MacPherson Plain 8. Ogilvie Mountains 8. North Ogilvie Mountains 9. Wernecke/Selwyn Mountains 10. 10. Mackenzie Mountains (in part) Mackenzie Mountains (in part) 11. Selwyn Mountains (in part) 10. Yukon/Tintina 15. 12. Yukon Plateau-Central Klondike Plateau (in part) 11. Eastern Plateaus 16. Yukon Plateau-North 12. Shakwak Trench 21. 12. Liard Basin (in part) Klondike Plateau (in part) 14. Ruby Ranges (in part) 13. Western Ranges 12. 14. Klondike Plateau (in part) Ruby Ranges (in part) 14. Pelly Mountains 18. Pelly Mountains (in part) 15. 16. Logan Mountains St. Elias/Coast Mountains 11. 13. Selwyn Mountains (in part) St. Elias Mountains 23. Mount Logan 17. 18. Southern Lakes Cassiar Mountains 17. 18. Yukon Southern Lakes Pelly Mountains (in part) 19. Liard Plain 18. Pelly Mountains (in part) Hyland/Liard Plateaus 21. 22. Liard Basin (in part) Hyland Highland 4. Muskwa Plateau 20. The Taiga Plain ecozone (B) only just enters along the eastern border with the Northwest Territories. For the most part, this ecozone is a rather flat plateau dominated by tundra, subalpine and boreal forest. The Taiga Cordillera ecozone (C) lies within the discontinuous permafrost zone, north of the Tintina Trench. It is underlain by sedimentary rocks to the north and metamorphic rocks to the south. The rugged mountainous areas have predominantly an alpine tundra vegetation, and were largely glaciated by one or more glaciations. The plateau regions have predominately a taiga or subarctic forest vegetation. The Boreal Cordillera ecozone (D) lies generally south of the Tintina Trench, is within the scattered-permafrost zone, and is underlain by a complex mixture of sedimentary, 54 G.G.E. Scudder volcanic, intrusive and metamorphic bedrock. Most of the ecozone lies within the Yukon drainage system, and has predominantly boreal white and black spruce forest, subalpine spruce-willow-birch, and alpine tundra at the higher elevations. The Pacific Maritime ecozone (E) is the extensive wet ecozone of the Pacific coast that just reaches into the southwestern corner of the Yukon. The ecozone is predominantly very high-elevation ice fields, alpine glaciers and summit outcrops. There is no terrestrial vegetation or soil development in this region. Ecoregions are subdivisions of the ecozones, characterized by distinctive large order landforms or assemblages of regional landforms, small order macro- or mesoclimates, and vegetational assemblages. The 23 ecoregions in the Yukon listed in Table 1 and mapped in Fig. 37 are described by the Ecological Stratification Working Group (1996). Table 2 provides comparisons of these ecoregions with the ecogeographic regions identified in Scudder (1987). The Yukon in Summary The general environmental features of the Yukon show it to be a relatively cold, rugged, and mountainous area, much of which is covered by relatively sparse or stunted vegetation, including arctic and alpine tundra, and subalpine and open subarctic forests. Nevertheless, the Yukon has relatively rich river valleys and substantial areas in the south with a surprising diversity of subtypes of forest and other vegetation. Moreover, several specific insect habitats, such as warm, south-facing slopes, have particular interest in the context of current faunal diversity and faunal history. During the Pleistocene much of the Yukon remained ice-free and was part of Beringia, the largest refugial area in the north during the last ice age. As such, the Yukon is a central focus of biogeography in the Nearctic region. Acknowledgements Research for this paper was supported by grants from the National Research Council of Canada. I am indebted to D.A. Demarchi (B.C. MELP), D. Russell (Canadian Wildlife Service, Whitehorse) and S. Smith (Agriculture and Agri-Food Canada, Whitehorse) for providing information and maps. R.A. Cannings and S.G. Cannings kindly provided landscape photographs. I thank L. Lucas for preparing the various illustrations and text. References Ager, T.A. 1982. Vegetational history of western Alaska during the Wisconsinan glacial interval and the Holocene. pp. 75 – 93 in D.M. Hopkins, J.V. Matthews Jr., C.E. Schweger, and S.B. Young (Eds.), Paleoecology of Beringia. Academic Press, New York. 489 pp. ______ 1983. Holocene vegetational history of Alaska. pp. 128 –141 in H.J. Wright (Ed.), Late-Quaternary Environments of the United States. Vol. 2. The Holocene. Univ. Minnesota Press, Minneapolis. 277 pp. Agriculture Canada Expert Committee on Soil Survey. 1987. The Canadian System of Soil Classification. 2nd edition. Res. Brch Agric. Can. Publ. 1646. 164 pp. Anderson, R.S. 1984. 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