Ecoregions of Canada's prairie grasslands. J.D. Shorthouse
Ecoregions of Canada's prairie grasslands. J.D. Shorthouse
Ecoregions of Canada's prairie grasslands. J.D. Shorthouse
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Chapter 3<br />
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands<br />
Joseph D. <strong>Shorthouse</strong><br />
Department <strong>of</strong> Biology, Laurentian University<br />
Sudbury, Ontario, Canada P3E 2C6<br />
Abstract. The Central Plains <strong>of</strong> Western Canada comprise the Prairies Ecozone and the Boreal Plains Ecozone.<br />
The former has the most <strong>grasslands</strong> and is divided into seven ecoregions: Mixed Grassland, Cypress Upland,<br />
Moist Mixed Grassland, and Fescue Grassland in southern Alberta and Saskatchewan; Aspen Parkland, which<br />
extends in an arc from central Alberta and Saskatchewan southeast to southwestern Manitoba; and Lake Manitoba<br />
Plain and Southwest Manitoba Uplands, but <strong>grasslands</strong> are more abundant in the first six. The Boreal Plains<br />
Ecozone extends across central and northern Alberta and Saskatchewan and arcs southeast to western Manitoba.<br />
This ecozone is divided into 10 ecoregions, but only the Peace Lowland has significant <strong>grasslands</strong>. This chapter<br />
briefly describes the physiography, climate, soils, and prominent flora <strong>of</strong> six <strong>of</strong> the ecoregions in the Prairies<br />
Ecozone and one in the Boreal Plains Ecozone for the benefit <strong>of</strong> future biologists wishing to study the biota <strong>of</strong><br />
<strong>prairie</strong> <strong>grasslands</strong>.<br />
Résumé. Les plaines centrales de l’ouest du Canada englobent l’écozone des <strong>prairie</strong>s et l’écozone des plaines<br />
boréales. La première renferme le plus de <strong>prairie</strong>s; elle se divise en sept écorégions : la <strong>prairie</strong> mixte; les hautes<br />
terres Cypress; la <strong>prairie</strong> mixte humide; la <strong>prairie</strong> à fétuque du sud de l’Alberta et de la Saskatchewan; la<br />
tremblaie-parc qui s’étend en demi-lune du centre de l’Alberta et du sud-est de la Saskatchewan jusqu’au sudouest<br />
du Manitoba; la plaine du lac Manitoba; les hautes terres du sud-ouest du Manitoba. Les <strong>prairie</strong>s sont plus<br />
abondantes dans les six premières de ces écorégions. L’écozone des plaines boréales décrit un arc du centre et<br />
du nord de l’Alberta et de la Saskatchewan vers le sud-est, jusque dans l’ouest du Manitoba; elle se divise en<br />
10 écorégions, mais seules les basses terres de la rivière de la Paix renferment d’importantes zones de <strong>prairie</strong>s.<br />
Ce chapitre décrit brièvement la physiographie, le climat, les sols et les espèces végétales principales de six des<br />
écorégions de l’écozone des <strong>prairie</strong>s et d’une des écorégions de l’écozone des plaines boréales à l’intention des<br />
biologistes qui souhaiteront dans le futur étudier les biotes des <strong>prairie</strong>s canadiennes.<br />
Introduction<br />
The North American Great Plains has the widest latitudinal range <strong>of</strong> any ecological region<br />
on the continent. It extends through parts <strong>of</strong> three Canadian provinces, 18 states in the<br />
United States, and states in Mexico (Gauthier and Wiken 2003). About 16% <strong>of</strong> the Plains<br />
are found in Canada, representing about 5% <strong>of</strong> the country’s landmass. The Central Plains<br />
<strong>of</strong> western Canada comprises two ecozones: the Prairies Ecozone and the Boreal Plains<br />
Ecozone (Fig. 1). Together they cover most <strong>of</strong> Alberta, the lower half <strong>of</strong> Saskatchewan,<br />
and western portions <strong>of</strong> southern Manitoba, and they edge into the Northwest Territories<br />
and the Peace River region <strong>of</strong> British Columbia. These ecozones converge in a unique<br />
ecoregion called the Aspen Parkland, where the southern <strong>prairie</strong> gradually gives way to tree<br />
cover. Some information on the features <strong>of</strong> these ecoregions was taken from websites <strong>of</strong><br />
the governments <strong>of</strong> the three <strong>prairie</strong> provinces and from the federal government document<br />
The State <strong>of</strong> Canada’s Environment (Environment Canada 1996). Information on climate<br />
<strong>Shorthouse</strong>, J. D. 2010. <strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands.<br />
In Arthropods <strong>of</strong> Canadian Grasslands (Volume 1): Ecology and Interactions in Grassland Habitats.<br />
Edited by J. D. <strong>Shorthouse</strong> and K. D. Floate. Biological Survey <strong>of</strong> Canada. pp. 53-81.<br />
© 2010 Biological Survey <strong>of</strong> Canada. ISBN 978-0-9689321-4-8<br />
doi:10.3752/9780968932148.ch3<br />
53
54 J. D. <strong>Shorthouse</strong><br />
Fig. 1. Map <strong>of</strong> the Boreal Plains Ecozone and the Prairies Ecozone. Note the Peace Lowland Ecoregion in<br />
the northwest.<br />
comes from the Ecological Stratification Working Group (1996). Scientific names <strong>of</strong> plants<br />
are from Moss and Packer (1983) and Harms (2003).<br />
The Prairies Ecozone is the most northern extension <strong>of</strong> the Great Plains <strong>of</strong> North<br />
America. The relief is typically subdued, consisting <strong>of</strong> low-lying valleys and plains that<br />
slope eastward. Since European settlement, the Prairies Ecozone has become one <strong>of</strong> the<br />
most extensive agricultural regions in the world. Its total land area is about 47 million ha,<br />
with 70% classified as cropland, 27% as rangeland, and the remaining 3% as deciduous<br />
forest. The natural fertility and good moisture-holding capacity <strong>of</strong> many <strong>of</strong> the soils in<br />
the Prairies Ecozone make the area extremely productive for crops. It is now rare to find<br />
unbroken native <strong>grasslands</strong> that show no evidence <strong>of</strong> European settlement.<br />
The Prairies Ecozone is subdivided into seven ecoregions: Mixed Grassland,<br />
Cypress Upland, Moist Mixed Grassland, and Fescue Grassland in southern Alberta and<br />
Saskatchewan; Aspen Parkland, which is north <strong>of</strong> the Moist Mixed Grassland, and extends<br />
in an arc from southwestern Alberta up toward Edmonton and then southeast to Manitoba<br />
(Fig. 2); and Lake Manitoba Plain and Boreal Transition in southern Manitoba (Fig. 2).
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 55<br />
Tallgrass <strong>prairie</strong> occurs in the southern part <strong>of</strong> Lake Manitoba Plain (Fig. 2). Only Aspen<br />
Parkland is contiguous throughout the three <strong>prairie</strong> provinces (Fig. 2).<br />
The Prairies Ecozone and its ecoregions receive the most attention in this book. The<br />
major towns and cities <strong>of</strong> the <strong>prairie</strong> <strong>grasslands</strong>, which serve as reference points for material<br />
in this and two anticipated volumes, are presented in Fig. 3. The major rivers and dams are<br />
presented in Fig. 4.<br />
The Boreal Plains Ecozone comprises 10 ecoregions with scattered patches <strong>of</strong><br />
<strong>grasslands</strong> in each. The largest <strong>of</strong> these <strong>grasslands</strong> is in the Peace Lowland in northwestern<br />
Alberta and northeastern British Columbia (Fig. 1). Scattered <strong>grasslands</strong> also occur in<br />
the Montane Cordillera and the Boreal Cordillera Ecozones <strong>of</strong> British Columbia and the<br />
Yukon, as well as in southern Ontario (see Chapter 4).<br />
<strong>Ecoregions</strong> <strong>of</strong> the Prairies Ecozone<br />
Mixed Grassland Ecoregion<br />
The Mixed Grassland Ecoregion is the most southern <strong>of</strong> the <strong>prairie</strong> ecoregions. It extends<br />
across southeastern Alberta and southwestern Saskatchewan from a base along the border<br />
with the United States (Fig. 2). This semi-arid ecoregion forms part <strong>of</strong> the shortgrass <strong>prairie</strong><br />
Fig. 2. Map <strong>of</strong> the Prairies Ecozone showing the location <strong>of</strong> ecoregions.
56 J. D. <strong>Shorthouse</strong><br />
Fig. 3. Map <strong>of</strong> the Prairies Ecozone showing the location <strong>of</strong> major communities and highways.<br />
in the Great Plains <strong>of</strong> North America that extends from southern Canada to Mexico (Bragg<br />
and Steuter 1996). Shortgrass <strong>prairie</strong> is characterized by the presence <strong>of</strong> short and midsized<br />
species <strong>of</strong> grass and sedges, as opposed to the much taller grasses found farther to the<br />
east (Coupland 1992).<br />
The Mixed Grassland Ecoregion is a treeless plain (Figs. 5 and 6) dominated by grasses<br />
and forbs. Elevations range from 600 m near Empress, Alberta, to more than 1,300 m<br />
on the lower slopes <strong>of</strong> the Cypress Hills, Sweetgrass Hills, and Milk River Ridge. In<br />
Alberta, the ecoregion covers about 4.8 million ha, or 49% <strong>of</strong> the total grassland area in<br />
the province. It is cultivated for crops or is used for pasture, with extensive irrigation in<br />
the western half. This ecoregion includes several tributaries <strong>of</strong> the South Saskatchewan<br />
River but few large lakes, a notable one being Pakowki Lake south <strong>of</strong> Medicine Hat. In<br />
Saskatchewan, this ecoregion encompasses about 13% <strong>of</strong> the province (Acton et al. 1998),<br />
with about half <strong>of</strong> the region cultivated for crops and the remainder used for pasture or<br />
rangeland. Lakes in the area include Clearwater, Old Wives, Fife, Willow Bunch, Big<br />
Muddy, Crane, and Chaplin. The South Saskatchewan River and Swift Current Creek form<br />
part <strong>of</strong> the Saskatchewan–Nelson drainage basin emptying into Hudson Bay. In contrast,<br />
the Frenchman River and Big Muddy River flow south into the Missouri River drainage<br />
system to empty into the Gulf <strong>of</strong> Mexico. The valleys formed by these waters may be
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 57<br />
entrenched 100 m or more into the plain. The soils are mainly Brown Chernozemic with<br />
significant areas <strong>of</strong> Solonetzic soils.<br />
The mean annual temperature <strong>of</strong> the ecoregion is approximately 3.5 °C. In southern<br />
Alberta, west <strong>of</strong> the Cypress Upland Ecoregion, mean annual temperatures can exceed<br />
5 °C. The mean summer temperature is 16 °C and the mean winter temperature is −10 °C.<br />
The mean annual precipitation ranges from 250 to 350 mm. Moisture deficits in late<br />
summer are caused by low precipitation and high evapotranspiration. Spring is usually the<br />
wettest season, with about two-thirds <strong>of</strong> the annual precipitation falling as rain, most <strong>of</strong> this<br />
occurring during June. During the winter months, the amount <strong>of</strong> snow cover is relatively<br />
low, as is the number <strong>of</strong> days <strong>of</strong> continuous snow cover. More than 30 chinook days per<br />
year occur along the western border <strong>of</strong> this ecoregion.<br />
Grasses and sedges <strong>of</strong>ten form 85–95% <strong>of</strong> the above-ground plant biomass in the<br />
Mixed Grassland Ecoregion, with forbs, sages, and dwarf shrubs making up the remainder<br />
(Rowe and Coupland 1984; Coupland 1992). The most common and widespread grass<br />
communities include mid-sized grasses such as speargrass (Stipa spp.), needle-and-thread<br />
grass (Hesperostipa comata (Trin. and Rupr.) Barkworth), western wheat grass (Agropyron<br />
smithii Rydb.), and June grass (Koeleria macrantha (Ledeb.) Schultes), as well as short<br />
Fig. 4. Map <strong>of</strong> the Prairies Ecozone showing the location <strong>of</strong> major rivers and dams. Note the salt flats from which<br />
there is no drainage.
58 J. D. <strong>Shorthouse</strong><br />
5<br />
6<br />
Figs. 5 and 6. Typical habitats <strong>of</strong> the Mixed Grassland Ecoregion. Fig. 5. Shortgrass <strong>prairie</strong> near Onefour in the<br />
southeast corner <strong>of</strong> Alberta looking south toward Sweetgrass Hills in neighbouring Montana. Fig. 6. Shortgrass<br />
<strong>prairie</strong> north <strong>of</strong> Medicine Hat. Photographs by the author.
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 59<br />
grasses such as blue grama (Bouteloua gracilis (Willd. ex Kunth) Lag. ex Griffiths), northern<br />
wheat grass (Elymus lanceolatus (Scribn. and Sm.)), and western porcupine grass (Stipa<br />
curtiseta Hitchc.). Blue grama and speargrass dominate at the driest sites, along with low<br />
sedge (Carex duriuscula C. A. Mey) and pasture sage (Artemisia frigida Willd.). Species that<br />
decrease with grazing pressure include wheat grass, green needlegrass (Nassella viridula<br />
(Trin.)), and prickly-pear cactus (Opuntia polyacantha Haw.), whereas blue grama, June<br />
grass, needle-and-thread grass, and pasture sage increase in number (Abouguendia 1996).<br />
In some habitats, shrubs such as hoary sagebrush (Artemisia cana Pursh) and wild roses<br />
(Rosa woodsii Lindl.; R. arkansana Porter) are prominent in scattered patches. Patches<br />
<strong>of</strong> cactus and pasture sage are common in some areas, with shrubs such as wolf willow<br />
(Elaeagnus commutata Bernh. ex Rydb.), Saskatoon berry (Amelanchier alnifolia Nutt.),<br />
snowberry (Symphoricarpos albus (L.)), and chokecherry (Prunus virginiana L.) growing<br />
in low-lying areas and coulees.<br />
Major communities in Alberta include Lethbridge, Taber, Brooks, and Medicine<br />
Hat, and in Saskatchewan, Swift Current, Kindersley, Maple Creek, Leader, Assiniboia,<br />
Shaunavon, and Gravelbourg (Fig. 3).<br />
Cypress Upland Ecoregion<br />
The Cypress Upland Ecoregion is an outlier <strong>of</strong> the montane vegetative zone that occurs on<br />
the lower slopes <strong>of</strong> the Rocky Mountains. The Cypress Hills straddle the border between<br />
Alberta and Saskatchewan (Fig. 2) to cover approximately 20,250 ha. They are the highest<br />
Canadian land elevation east <strong>of</strong> the Rockies and west <strong>of</strong> Labrador and consist <strong>of</strong> three<br />
separate blocks or hills, one in Alberta and two in Saskatchewan. The hills rise 600 m<br />
above the surrounding plains and are home to freshwater streams and creeks, lakes, fescue<br />
<strong>grasslands</strong>, forests, meadows, marshlands, and ranchlands. On July 28, 1850, Captain John<br />
Palliser wrote in his journal: “These hills [the Cypress Hills] are a perfect oasis in the<br />
desert we have travelled” (Beaty, 1975). Hills on the Saskatchewan side are <strong>of</strong> historical<br />
importance (Hildebrandt and Huber 1994) because they were the site <strong>of</strong> the first fort <strong>of</strong> the<br />
North-West Mounted Police (Fig. 7).<br />
The higher elevation <strong>of</strong> the hills provides for cooler temperatures and increased<br />
precipitation compared with the surrounding <strong>prairie</strong>s, thereby facilitating the growth <strong>of</strong><br />
trees. The hills were named “myun-a-tuh-gow” (beautiful highlands) by the Cree and<br />
“ketius netumoo” (hills that shouldn’t be) by the Blackfoot. The name Cypress Hills<br />
originates from an incorrect translation <strong>of</strong> the French “montagne aux cyprès.” The term<br />
“cyprès” was used by French Canadians and Métis to refer to jack pine (Pinus banksiana<br />
Lamb) and lodgepole pine (Pinus contorta Douglas ex Louden); there are no cypress trees<br />
in the Cypress Hills.<br />
The Cypress Upland is a dissected plateau with elevations that rise abruptly above<br />
the surrounding plain (Fig. 8), with steeply sloping escarpments and numerous valleys<br />
and coulees. It formed during the Tertiary period from material that was worn round<br />
on its long river journey from newly formed western mountains (Acton et al. 1998).<br />
Coarse gravels formed erosion-resistant armour that protected the underlying sediments<br />
from removal during the last glacial period. The Cypress Upland consisted <strong>of</strong> nunataks<br />
(unglaciated peaks) during the last Ice Age that are now a series <strong>of</strong> benchlands, <strong>of</strong>ten<br />
referred to as the West, Centre, and East blocks. The hills slope eastward from a maximum<br />
elevation <strong>of</strong> 1,465 m on the west side. This ecoregion is drained by the deeply incised<br />
Frenchman River and the Battle and Swift Current creeks. Several <strong>of</strong> the north-slope<br />
drainage basins are closed, with water flowing into shallow saline lakes from where it
60 J. D. <strong>Shorthouse</strong><br />
evaporates. Most <strong>of</strong> the hills are within two provincial parks that straddle the Alberta–<br />
Saskatchewan border.<br />
The climate <strong>of</strong> the Cypress Hills is cooler and moister than that <strong>of</strong> the surrounding<br />
Mixed Grassland Ecoregion (Acton et al. 1998), with a mean annual temperature <strong>of</strong> 2.7 °C,<br />
a mean July temperature <strong>of</strong> 16 °C, and a mean January temperature <strong>of</strong> −12.1 °C. Mean<br />
annual precipitation ranges from 325 to 450 mm, with 244 mm <strong>of</strong> rainfall occurring from<br />
May to September.<br />
With a rise in elevation, the vegetation changes to a submontane fescue <strong>prairie</strong> at<br />
the top <strong>of</strong> the benches and on the south-facing slopes (Fig. 7) and a mix <strong>of</strong> conifers and<br />
deciduous trees at the upper elevations (Fig. 8). The forest supports a wide array <strong>of</strong> plant<br />
species (see Breitung 1954; Bird 1962; Moss and Packer 1983), including 17 types <strong>of</strong><br />
orchids, and provides habitat for invertebrates and vertebrates that are absent from the<br />
adjacent <strong>grasslands</strong>. Four species <strong>of</strong> trees are native to the Cypress Hills (Newsome and<br />
Dix 1968): lodgepole pine, white spruce (Picea glauca (Moench) Voss), aspen or trembling<br />
poplar (Populus tremuloides Michx.), and balsam poplar (Populus balsamifera L.).<br />
White spruce forests grow on uplands in cooler and moister areas, such as north-facing<br />
slopes (Fig. 8) and near springs. Pure stands occur near wetlands, coulees, and creeks,<br />
but mixed stands with trembling aspen are found on well-drained soils. Aspen woodlands<br />
occur in many habitats in patches below the lodgepole pine woodland on the north side <strong>of</strong><br />
the plateau. Patches <strong>of</strong> fescue <strong>prairie</strong> (Fig. 7) are found on the lower slopes.<br />
The fescue <strong>grasslands</strong> are <strong>of</strong> interest because they do not exist on the dry <strong>grasslands</strong><br />
surrounding the hills and are isolated from those in the foothills farther to the west. Fescue<br />
<strong>grasslands</strong> occur at the highest elevations <strong>of</strong> the Cypress Hills, where the dominant species<br />
is rough fescue (Festuca campestris Rydb.). Other co-occurring species <strong>of</strong> grass include<br />
bluebunch fescue (F. idahoensis Elmer), northern wheat grass, western wheat grass,<br />
awned wheat grass (Agropyron subsecundum (Link) A. S. Hitchc.), June grass, western<br />
porcupine grass, and desert wheat grass (A. desertorum (Fisch. ex Link) Schultes) and<br />
smooth bromegrass (Bromus inermis Leyss.) (Best et al. 1971; Rowe and Coupland 1984).<br />
Common shrubs such as snowberry, cinquefoil (Potentilla spp.), wolf willow, and wild<br />
roses (<strong>Shorthouse</strong> 1991) are found among the fescue <strong>prairie</strong>.<br />
Great Sand Hills and Grasslands National Park<br />
Over 120 areas <strong>of</strong> sand dunes totalling 10,000 km 2 in Alberta, Saskatchewan, and Manitoba<br />
are scattered throughout the Prairies and Boreal Plains ecozones. The Great Sand Hills, the<br />
largest <strong>of</strong> these areas at about 1,900 km 2 , is located in southwestern Saskatchewan between<br />
Leader and Gull Lake and slightly east <strong>of</strong> the Alberta border. The Great Sand Hills area<br />
is characterized by sand dune <strong>grasslands</strong>, low hills, saline lakes, pebble plains, stands <strong>of</strong><br />
cottonwood, aspen bluffs, and individual dunes that exceed 15 m in height (Figs. 9 and 10).<br />
Sand dune fields occur in habitats that are dry and where sand is abundant and subject<br />
to aeolian processes. These conditions create either rolling or active dunes, stabilized dunes,<br />
stabilized blowouts, or dune depressions (Hulett et al. 1966; David 1993). Many sand<br />
dunes in the Great Sand Hills, and elsewhere in Palliser’s Triangle, are classic parabolic<br />
dunes, which consist <strong>of</strong> a slip face that is convex in plain view and has wings that extend<br />
upwind <strong>of</strong> the dune head (Lemmen et al. 1997). In drier years, wind may move the sand to<br />
form parabolic dunes (Wolfe and David 1997).<br />
Although sandy habitats tend to be low in nutrients, the sand’s permeability allows<br />
for the accumulation <strong>of</strong> fine sediment, moisture, and nutrients in lower lying areas. This
7<br />
8<br />
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 61<br />
Figs. 7 and 8. Cypress Hills <strong>of</strong> southwestern Saskatchewan and southeastern Alberta. Fig. 7. Aspen, conifers, and<br />
a field <strong>of</strong> fescue grasses near the Fort Walsh site on the Saskatchewan side. Fig. 8. North-facing slope with white<br />
spruce on the Alberta side. Note the <strong>prairie</strong> below in the distance. Photographs by the author.
62 J. D. <strong>Shorthouse</strong><br />
9<br />
10<br />
Figs. 9 and 10. Great Sand Hills <strong>of</strong> Saskatchewan. Fig. 9. Grasses, shrubs, and creeping juniper growing on flat, sandy<br />
areas between the dunes. Fig. 10. Edge <strong>of</strong> an active dune showing shrubs being buried. Photographs by the author.
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 63<br />
accumulation creates a landscape with sparsely vegetated dunes and sand flats (Fig. 9) that<br />
separate diverse patches <strong>of</strong> shrubby vegetation. Dune grass assemblages occur on active<br />
aeolian ridges and, to a lesser degree, on semi-stable dune ridges. Vegetation cover is sparse<br />
and species composition is variable, depending on moisture and activity <strong>of</strong> dunes.<br />
Common plant species <strong>of</strong> the Sand Hills include sand grass (Calamovilfa longifolia<br />
(Hook.) Scribn.), sand dropseed (Sporobolus cryptandrus (Torr.) A. Gray), Indian rice grass<br />
(Oryzopsis hymenoides (Roem. and Schult.) Barkworth), and prickly-pear cactus. Sand<br />
grass is the only Canadian representative <strong>of</strong> the genus Calamovilfa and is well adapted to<br />
growth on sand dunes, lakeshores, and open sandy forested sites. Scaly rootstocks, the bulk<br />
<strong>of</strong> which mass near the soil surface, bind the sandy soil and keep it from blowing.<br />
Active dunes frequently bury grasses and shrubs (Fig. 10) but can be colonized by<br />
grasses such as northern wheat grass, Indian rice grass, sand grass, and sand dropseed.<br />
Common shrubs on the flats between dunes include creeping juniper (Juniperus horizontalis<br />
Moench) (Fig. 9), wild roses, aspen poplar, and wolf willow. On the border <strong>of</strong> active dunes,<br />
perennials with extensive rootstocks such as lance-leaved psoralea (Psoralea lanceolata<br />
(Pursh) Rydb.) persist, as do annuals that can grow through thick deposits <strong>of</strong> sand, such as<br />
small lupine (Lupinus pusillus Pursh) and skeletonweed (Shinnersoseris rostrata (Gray)<br />
Tomb) (Hulett et al. 1966). More typical <strong>prairie</strong> species such as needle-and-thread grass,<br />
June grass, and creeping juniper grow on stabilized dunes. In the moister and nutrient-rich<br />
low-lying habitats, shrubs and trees are common, for example, Saskatoon, white sagebrush<br />
(Artemisia ludoviciana Nutt.), chokecherry, wild rose, and aspen.<br />
Most <strong>of</strong> the Great Sand Hills landscape has been used for ranching over the last hundred<br />
years, but the majority <strong>of</strong> land remains in provincial ownership. Cattle fill a niche similar<br />
to that <strong>of</strong> bison and, as a result, parts <strong>of</strong> the ecosystems <strong>of</strong> the Sand Hills are likely similar<br />
to what they were in the past. Oil and gas extraction currently pose the greatest threat<br />
to the ecological integrity <strong>of</strong> the Great Sand Hills and, indeed, much <strong>of</strong> Saskatchewan’s<br />
remaining native <strong>grasslands</strong>.<br />
Grasslands National Park, located between Val Marie and Killdeer and abutting the<br />
Saskatchewan–Montana border, is also important for future grassland studies. The Park<br />
comprises two regions (the West Block in the Frenchman River Valley and the East Block in<br />
the Killdeer Badlands) and contains some <strong>of</strong> the best examples <strong>of</strong> the harsh, dry conditions<br />
characteristic <strong>of</strong> the Mixed Grassland Ecoregion. Surface waters are derived mainly from<br />
spring run<strong>of</strong>f such that many creeks (e.g., Breed, Otter, Horse, and Wetheral creeks) and<br />
small lakes in the park are dry later in the summer. The Park has only two permanent water<br />
sources, the Frenchman River in the West Block, which is a prime example <strong>of</strong> a glacial<br />
meltwater channel, and Rock Creek in the East Block.<br />
Moist Mixed Grassland Ecoregion<br />
The Moist Mixed Grassland Ecoregion encircles the Mixed Grassland Ecoregion in an<br />
arc extending from the southwestern interior <strong>of</strong> Alberta northward from the Montana<br />
border and curving into Saskatchewan to the east <strong>of</strong> Red Deer, across to Saskatoon, and<br />
then southeast through Regina back to the border with the United States, west <strong>of</strong> the<br />
Saskatchewan–Manitoba border (Fig. 2). Moisture generally increases and the temperature<br />
decreases toward the north, west, or east compared with the Mixed Grassland Ecoregion,<br />
resulting in a change in associated vegetation. Thus, the Moist Mixed Grassland Ecoregion<br />
supports the more vigorous growth <strong>of</strong> grasses (Fig. 11) and shrubs (Fig. 12) than occurs in<br />
the drier Mixed Grassland Ecoregion. As a result, about 80% <strong>of</strong> the Moist Mixed Grassland<br />
Ecoregion is under cultivation.
64 J. D. <strong>Shorthouse</strong><br />
Physiographically, most <strong>of</strong> the Moist Mixed Grassland Ecoregion is a broad plane<br />
(Fig. 11) interrupted by deep valleys and subdued hilly uplands. Valleys <strong>of</strong> the Qu’Appelle<br />
and South Saskatchewan rivers (Fig. 4) are sometimes sunken 100 m or more into the<br />
plain. Level to gently undulating glacial till plains that seem almost endless typify this<br />
region. A large, level glacial lake plain called the Regina Plain extends from the base <strong>of</strong><br />
the Missouri Coteau south and west <strong>of</strong> Moose Jaw to the Moose Mountain uplands east<br />
<strong>of</strong> Regina and north to the Qu’Appelle Valley. This former glacial lake, known as Lake<br />
Regina, was drained by the Qu’Appelle Valley as the glacier receded from the area. Hills<br />
within the region include Bear Hills north <strong>of</strong> Rosetown and the Allan Hills southeast <strong>of</strong><br />
Saskatoon. Sand hills within the region include those near Douglas Provincial Park north<br />
<strong>of</strong> Moose Jaw. Lakes include Muddy, Trampling, Diefenbaker, Goose, Last Mountain,<br />
Little Manitou, and Buffalo Pound.<br />
The mean annual temperature is about 5 °C, with a mean summer temperature <strong>of</strong><br />
about 15.5 °C, which is 1–2 degrees cooler than the Mixed Grassland Ecoregion. The<br />
mean winter temperature is −11 °C, which is also 1–2 °C cooler than the Mixed Grassland<br />
Ecoregion. The mean annual precipitation ranges from 350 to 450 mm, about 20% greater<br />
than in the Mixed Grassland Ecoregion. The southwestern part <strong>of</strong> the ecoregion in Alberta<br />
lies in the chinook belt along the face <strong>of</strong> the Rocky Mountain foothills. Snow cover is<br />
greater because <strong>of</strong> more winter precipitation in the chinook belt. More intermittent sloughs,<br />
potholes, and streams are present in this ecoregion compared with the drier regions in the<br />
south. Soil types are Dark Brown Chernozemic or Solonetzic associated with sandy to<br />
clayey lacustrine deposits that developed under <strong>prairie</strong> grassland influences.<br />
The vegetation <strong>of</strong> the Moist Mixed Grassland Ecoregion is similar to the drier Mixed<br />
Grassland Ecoregion but is characterized by greater biomass production and a greater<br />
abundance <strong>of</strong> species that favour cooler and moister sites. Species such as western<br />
porcupine grass (Stipa curtiseta Hitchc.), needle-and-thread grass, western wheat grass<br />
(Agropyron smithii (Rydb.)), and northern wheat grass are more common than occurs<br />
farther south. A western wheat grass–northern wheat grass community occurs widely<br />
at mesic sites, whereas fine-textured soils in glacial-lake basins are characterized by<br />
northern wheat grass–June grass communities. On drier, exposed sites, blue grama is more<br />
common. More deciduous shrubs such as buckbrush, chokecherry, wolf willow, wild rose,<br />
and Saskatoon are present here (Fig. 12) than in the Mixed Grassland Ecoregion. The<br />
most abundant species among the forbs and shrubs throughout the region is pasture sage.<br />
Although this ecoregion does not extend into Manitoba, patches <strong>of</strong> <strong>grasslands</strong> similar<br />
to those found in the Moist Mixed Grassland Ecoregion can be found among the aspen<br />
parkland forests <strong>of</strong> the southwestern part <strong>of</strong> the province.<br />
Major communities in this ecoregion include Fort Macleod, Lethbridge, Drumheller,<br />
Biggar, Saskatoon, Moose Jaw, Regina, Estevan, and Weyburn.<br />
Fescue Grassland Ecoregion<br />
This ecoregion lies to the west <strong>of</strong> the Moist Mixed Grassland Ecoregion in southwestern<br />
Alberta (Fig. 2) along the eastern face <strong>of</strong> the Rocky Mountain foothills. It is a small<br />
but ecologically important ecoregion that is dominated by rough fescue grass (Festuca<br />
campestris Rydb.) (Aiken and Darbyshire 1990; Aiken et al. 1997). Most ecologists<br />
consider fescue <strong>grasslands</strong> to be restricted to this part <strong>of</strong> the province (Doormaar and<br />
Willms 1990). However, intermittent patches <strong>of</strong> fescue also occur along the northern edge<br />
<strong>of</strong> the Moist Mixed Grassland Ecoregion and within the Aspen Parkland Ecoregion in<br />
Alberta and Saskatchewan and in southwestern Manitoba. Some ecologists consider this
11<br />
12<br />
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 65<br />
Figs. 11 and 12. Typical habitats <strong>of</strong> the Moist Mixed Grassland Ecoregion. Fig. 11. Mixed grass west <strong>of</strong> Lethbridge<br />
near Stand<strong>of</strong>f on the Blood Indian Reserve. Fig. 12. Mixed grass north <strong>of</strong> Saskatoon showing dense grasses and<br />
deciduous shrubs such as buckbrush, chokecherry, wolf willow, and Saskatoon near some stunted aspen poplars.<br />
Photographs by the author.
66 J. D. <strong>Shorthouse</strong><br />
to be evidence <strong>of</strong> a second Fescue Grassland Ecoregion (Coupland and Brayshaw 1953).<br />
However, this chapter adopts the view that the Fescue Grassland Ecoregion is restricted to<br />
southwestern Alberta.<br />
The Fescue Grassland Ecoregion covers just over 1.3 million ha (approximately<br />
2%) <strong>of</strong> Alberta’s total land area. Most <strong>of</strong> the ecoregion consists <strong>of</strong> rolling, hummocky,<br />
dissected till left by glaciation. This ecoregion occurs largely on morainal, glaciolacustrine,<br />
and outwash deposits along the lower flanks <strong>of</strong> the foothills at the base <strong>of</strong> the Rocky<br />
Mountains and in the Porcupine Hills, the Milk River Ridge, and adjacent plains areas.<br />
Thus, the ecoregion exists primarily as a narrow band between the Moist Mixed Grassland<br />
Ecoregion to the east and the Aspen Parkland Ecoregion to the northwest in the foothills.<br />
The largest <strong>of</strong> the remaining patches <strong>of</strong> the Fescue Grassland Ecoregion are in Waterton<br />
Lakes National Park (Fig. 13) and the McIntyre Ranch (Fig. 14) on the Milk River Ridge<br />
south <strong>of</strong> Magrath. Disjunct areas <strong>of</strong> rough fescue also grow on the lower slopes <strong>of</strong> the<br />
Sweetgrass Hills, near the Alberta–Montana border, and on the plateaux <strong>of</strong> the Cypress<br />
Hills (see previous section). Elevations in this subregion are much higher than in the other<br />
two grassland ecoregions.<br />
Climate in southwestern Alberta is characterized by warm summers, and winters are<br />
milder than those <strong>of</strong> the Mixed Grassland and Moist Mixed Grassland ecoregions because<br />
<strong>of</strong> the moderating affects <strong>of</strong> chinook winds. The mean annual temperature is approximately<br />
3.5 °C, with a mean summer temperature <strong>of</strong> 14 °C and a mean winter temperature <strong>of</strong> −8 °C.<br />
Mean annual precipitation ranges from 400 to 500 mm. Greater snowfall occurs in late<br />
winter and early spring. The majority <strong>of</strong> precipitation falls during the growing season.<br />
The soils <strong>of</strong> this subregion are predominantly Dark Brown and Black Chernozems.<br />
Solonetzic soils are absent. The rich soils throughout the region led early farmers to quickly<br />
convert most <strong>of</strong> the area into croplands and pastures.<br />
Alberta has the largest area <strong>of</strong> rough fescue grassland in the world (Doormaar and<br />
Willms 1990). Rough fescue <strong>grasslands</strong> are among the most productive <strong>grasslands</strong> in<br />
North America (Doormaar and Willms 1990; Willms et al. 1996), providing valuable<br />
forage for native and non-native ungulates (Stout et al. 1981; Willms 1987). These<br />
<strong>grasslands</strong> have high spring protein and digestible carbohydrate content and are the main<br />
reason bison once overwintered here. Unfortunately, the fescue <strong>grasslands</strong> are some <strong>of</strong><br />
the most threatened plant communities in the Canadian <strong>prairie</strong>s. Concern about their<br />
loss as a result <strong>of</strong> development, woodland encroachment, the introduction <strong>of</strong> exotic<br />
species, and overgrazing has increased because only 5% <strong>of</strong> the <strong>grasslands</strong> remain in presettlement<br />
condition.<br />
Rough fescue was declared Alberta’s provincial grass in 2003. Although scientists<br />
are in disagreement (see Pavlick and Looman 1984; Harms 1985), rough fescue has been<br />
described as a complex <strong>of</strong> three species: mountain rough fescue (Festuca scabrella),<br />
plains rough fescue (F. hallii (Vasey) Piper), and northern rough fescue (F. altaica Trin. ex<br />
Ledeb). If this interpretation is valid, the province is the only jurisdiction in North America<br />
with three co-occurring species <strong>of</strong> rough fescue, which converge in southwestern Alberta,<br />
reflecting a pattern <strong>of</strong> recolonization following glaciation.<br />
Mountain rough fescue is also named foothills rough fescue and buffalo bunchgrass.<br />
This last name refers to the fondness <strong>of</strong> bison for this grass. It is a perennial bunchgrass<br />
(growing in dense tufts) with stiff, narrow leaves (
13<br />
14<br />
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 67<br />
Figs. 13 and 14. Typical habitats <strong>of</strong> the Fescue Grassland Ecoregion. Fig. 13. Hummocky, glacial moraine in<br />
the foothills at the base <strong>of</strong> the Rocky Mountains in Waterton Lakes National Park. Fig. 14. Habitat with dense<br />
forbs and fescue grasses on the McIntyre Ranch south <strong>of</strong> Magrath on the south side <strong>of</strong> the Milk River Ridge.<br />
Photographs by the author.
68 J. D. <strong>Shorthouse</strong><br />
Manitoba and North Dakota. This fescue is a dominant species in the <strong>grasslands</strong> associated<br />
with the Aspen Parkland Ecoregion north <strong>of</strong> the Moist Mixed Grassland <strong>of</strong> Alberta and<br />
Saskatchewan (Pavlick and Looman 1984; Harms 1985; Aiken and Darbyshire 1990).<br />
Northern rough fescue occurs from the Rocky Mountains <strong>of</strong> west-central Alberta and eastcentral<br />
British Columbia north through the Yukon to Alaska and into central Asia. It grows<br />
in open woods and moist meadows and dominates ridgetop <strong>grasslands</strong> in Alberta’s Rocky<br />
Mountain subalpine regions. Mountain rough fescue and plains rough fescue are believed<br />
to have evolved under winter grazing by bison. Bluebunch fescue (F. idahoensis Elmer),<br />
with a range centred in western Montana, also occurs in Alberta’s foothills north <strong>of</strong> Calgary<br />
(Looman 1983).<br />
The northern fescue <strong>grasslands</strong> associated with the Aspen Parkland Ecoregion occur in<br />
an arc passing through Drumheller and Coronation and into Saskatchewan. Compared with<br />
the mixedgrass <strong>prairie</strong>, plant assemblages here have higher species richness for both grasses<br />
and forbs (Rowe and Coupland 1984) than does the Moist Mixed Grassland Ecoregion to<br />
the south. Fescue <strong>grasslands</strong> in the southern <strong>prairie</strong>s have a greater variety and cover <strong>of</strong><br />
forbs than occurs in the Mixed and Moist Mixed Grassland ecoregions.<br />
Major communities in the Fescue Grassland Ecoregion include High River, Pincher<br />
Creek, and Cardston (Fig. 3).<br />
Aspen Parkland Ecoregion<br />
The Aspen Parkland Ecoregion is a north–south transition zone between the dry <strong>grasslands</strong><br />
to the south and the boreal forest to the north. It stretches in a broad arc from southwestern<br />
Alberta northward and then eastward through central and northwestern Alberta, central<br />
Saskatchewan, and central and southern Manitoba, including nearly all <strong>of</strong> southwestern<br />
Manitoba (Fig. 2). In southwestern Alberta, this ecoregion begins north <strong>of</strong> the Fescue<br />
Grassland Ecoregion in the foothills on the east side <strong>of</strong> the Rocky Mountains. The Aspen<br />
Parkland Ecoregion is no wider than 500 km, being widest in Alberta and southeastern<br />
Saskatchewan and southwestern Manitoba. It is also an east–west transition zone between<br />
the mountains and <strong>grasslands</strong> in southwestern Alberta.<br />
In its native state, the Aspen Parkland Ecoregion is characterized by a mosaic <strong>of</strong><br />
trembling aspen (Farrar 1995) found in groves in its southern extent (Fig. 15). Aspen form<br />
thicker stands in more central and northern parts <strong>of</strong> the ecoregion (Fig. 16), interspersed<br />
with northern and plains rough fescues (Pavlick and Looman 1984) or moist mixedgrass<br />
<strong>prairie</strong> (Coupland 1961). Grasslands dominated by plains and northern fescue (see<br />
previous section) are found in open areas between groves in the southern and central parts<br />
<strong>of</strong> the ecoregion and then in intermittent patches in more northern parts. A succession <strong>of</strong><br />
gradually taller vegetation from grassland to forest is common in more northern regions<br />
(Fig. 17). Groves <strong>of</strong> bur oak (Quercus macrocarpa Michx.) are common in southeastern<br />
Saskatchewan and southwestern Manitoba in this ecoregion (Farrar 1995). With only a few<br />
exceptions, the Aspen Parkland Ecoregion is a uniquely Canadian ecoregion.<br />
The Aspen Parkland Ecoregion consists <strong>of</strong> two natural subregions, or sections, referred<br />
to as Central Parkland and Foothills Parkland. A third subregion <strong>of</strong> Aspen Parkland is in the<br />
Peace River Parkland district <strong>of</strong> northwestern Alberta and adjoining northeastern British<br />
Columbia. The Peace River Parkland is located in the Peace River region <strong>of</strong> the province<br />
and is completely surrounded by boreal forest, cut <strong>of</strong>f from the Central Parkland. The Peace<br />
River Parkland is generally cooler than the Central Parkland but still supports <strong>grasslands</strong><br />
and now extensive farmland. All three regions are ecologically similar, being distinguished<br />
mainly by their geographical location.
15<br />
16<br />
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 69<br />
Figs. 15 and 16. Typical habitats <strong>of</strong> the Aspen Parkland Ecoregion. Fig. 15. Typical aspen grove near Onoway,<br />
Alberta. Fig. 16. Typical stand <strong>of</strong> aspen poplar showing grasses, forbs, and shrubs growing up to the edge <strong>of</strong> an<br />
aspen grove north <strong>of</strong> Saskatoon, Saskatchewan. Photographs by the author.
70 J. D. <strong>Shorthouse</strong><br />
17<br />
18<br />
Figs. 17 and 18. Habitats in the Aspen Parkland Ecoregion. Fig. 17. Fescue grasses, wild roses, and aspen<br />
encroaching on the grassland in Prince Albert National Park in Saskatchewan. Fig. 18. Aspen forests being<br />
cleared and burned east <strong>of</strong> Red Deer, Alberta. Photographs by the author.
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 71<br />
The Central Parkland is the largest section and comprises the main band <strong>of</strong> aspen<br />
parkland extending west to east through central Alberta and Saskatchewan down to<br />
southwestern Manitoba. A continuum <strong>of</strong> grassland occurs from south to north within and<br />
around groves <strong>of</strong> aspen, to aspen parkland, to closed aspen forest in the north. True parkland<br />
vegetation with continuous aspen forest broken by grassland openings is now rare because<br />
<strong>of</strong> large-scale clearing and burning (Fig. 18). In much <strong>of</strong> the rest <strong>of</strong> the Central Parkland<br />
section, native vegetation is scarce because <strong>of</strong> agriculture.<br />
The southern part <strong>of</strong> the parkland is known as grove land, a region dominated by<br />
grasses but with scattered small clumps <strong>of</strong> aspens in depressions that collect moisture. As<br />
the moisture content <strong>of</strong> the soils increases to the north, so, too, does the abundance <strong>of</strong> trees<br />
until, at its northern limit, the Aspen Parkland Ecoregion can be a solid forest. This is a<br />
zone <strong>of</strong> constant competition between grassland and boreal vegetation trying to overtake<br />
one another. Today it is a checkerboard <strong>of</strong> cropland with a few surviving patches <strong>of</strong> original<br />
parkland in the Wainwright area, the Neutral and the Bodo hills, and the Sounding Lake<br />
sand plain.<br />
Climate throughout the Central Parkland is continental, with long, cold winters and<br />
short summers, which tend to be cool to warm. The mean annual temperature for the Central<br />
Parkland region is 2 °C, with a May-to-September average <strong>of</strong> 13 °C. The frost-free period<br />
averages 95 days. Mean annual precipitation is 350–450 mm, with a May-to-September<br />
average <strong>of</strong> 300 mm. Precipitation is generally lower than in the boreal forest but is still<br />
higher than in the <strong>grasslands</strong> to the south.<br />
Black and Dark Brown Chernozems predominate under grassland vegetation, whereas<br />
Dark Gray Chernozems and Luvisolic soils are most common under the moister aspen<br />
woodlands. Dark Brown Chernozems occur under woodland vegetation on sandy parent<br />
materials. A strip <strong>of</strong> Brown Solonetz soils runs through the centre <strong>of</strong> the subregion from<br />
Vegreville through Beaverhill Lake and into the Sullivan and Dowling lakes areas.<br />
Trembling aspen is a clonal species and is well-suited to parkland (Bird 1961).<br />
Trembling aspen can withstand fire by producing suckers from horizontal roots even if<br />
the “mother” tree is destroyed. The characteristic uniform height <strong>of</strong> trees in aspen groves<br />
reflects the common age <strong>of</strong> the individuals that likely developed from suckers after a<br />
disturbance. Aspen produce seeds every year, but reproduction is most <strong>of</strong>ten by vegetative<br />
suckering, or cloning. Suckers can appear 25 m from the parent plant and grow 3 m in a<br />
single year. Hence, aspen can quickly encroach upon grass-dominated areas after a fire.<br />
Aspen woodlands support a dense, entangled, species-rich understory (Fig. 16) <strong>of</strong> midsized<br />
and small shrubs such as red osier dogwood (Cornus stolonifera Michx.), beaked<br />
willow (Salix bebbiana Sarg.), Saskatoon, and chokecherry. Roses and snowberry are also<br />
common shrubs in these <strong>grasslands</strong>.<br />
Most <strong>of</strong> the aspen parkland, like the <strong>prairie</strong>s, has been extensively altered for<br />
agriculture over the last 100 years since settlement first began in the late 19th century.<br />
Of the 7.35 million original hectares <strong>of</strong> aspen parkland in Alberta, 90–95% has been<br />
altered or destroyed, primarily to support agriculture (Bird 1961). The same has occurred<br />
in Saskatchewan. Although the climate is generally cooler in this part <strong>of</strong> Alberta and<br />
Saskatchewan than it is farther south, it is still mild and dry enough to support large-scale<br />
farming. Most <strong>of</strong> the agricultural land is used for growing canola, alfalfa, and wheat. Most<br />
remaining stands <strong>of</strong> original parkland are less than 1,000 ha and are on marginal soils. Oil and<br />
natural gas exploration and drilling have also disturbed the natural habitat in many areas.<br />
The Aspen Parkland Ecoregion in Manitoba occupies the southwestern corner <strong>of</strong><br />
the province in an area nestled between Riding and Duck mountains. It forms part <strong>of</strong> the
72 J. D. <strong>Shorthouse</strong><br />
extensive parkland belt between the closed boreal forest cover to the north and northeast<br />
and treeless <strong>grasslands</strong> to the west and is a mosaic <strong>of</strong> trembling aspen, oak groves, and<br />
rough fescue <strong>grasslands</strong>. The region slopes gently eastward and is drained by the Souris,<br />
Assiniboine, Qu’Appelle, and Pembina rivers (Fig. 4). The average annual precipitation in<br />
the area is about 500 mm. Trembling aspen and shrubs occur on moist sites, with bur oak<br />
and grassland communities occupying drier sites. Dominant grasses here include fescues,<br />
wheat grass, and June grass. Patches <strong>of</strong> the tallgrasses such as big bluestem (Andropogon<br />
gerardii Vitman), once more common south <strong>of</strong> Winnipeg, are also found in some areas.<br />
Of ecological interest is the Shilo area east <strong>of</strong> Brandon, bordered by the Assiniboine<br />
River on the south and the Manitoba Escarpment on the east, with the Spruce Woods<br />
Provincial Forest and Spruce Woods Provincial Park to the north. Here white spruce is<br />
associated with a ground cover <strong>of</strong> mostly mixed <strong>prairie</strong> grasses. Large tracts <strong>of</strong> fine to<br />
medium sands, deposited here by glacial meltwater, have been blown into hummocky<br />
dunes characterized by gentle to strong slopes.<br />
The historically important Criddle/Vane homestead is situated between Shilo and<br />
the Assiniboine River. Established in 1882 on a quarter section <strong>of</strong> virgin grassland that<br />
contained representatives <strong>of</strong> the Moist Mixed Grassland and the Lake Manitoba Plains<br />
ecoregions, it was the former home (Fig. 19) <strong>of</strong> Norman Criddle, one <strong>of</strong> Canada’s pioneers<br />
in entomology. Criddle began his acquaintance with the natural history <strong>of</strong> the <strong>grasslands</strong><br />
and surrounding forests here, resulting in an enormous number <strong>of</strong> insect specimens<br />
labelled with “Aweme, Man., Norman Criddle” in various museums. He established the<br />
first entomological field station (Fig. 20) in western Canada at the site, which now has been<br />
designated as part <strong>of</strong> the Manitoba Provincial Park system. A local historical society has<br />
renovated the field station.<br />
The homestead and its surroundings are important because <strong>of</strong> the broad array <strong>of</strong><br />
information associated with them (Roughley 2000). The Criddles operated a weather<br />
station, which provided one <strong>of</strong> the longest continuous records <strong>of</strong> weather in Canada. They<br />
made observations and collections <strong>of</strong> plants and animals, including insects. The natural<br />
history <strong>of</strong> the area has likely been studied more intensively than any other locality in<br />
Canada. It has been the site <strong>of</strong> important ecological studies, including the interaction<br />
among fire, fire frequency, and competition within the plant communities (Wilson and<br />
Shay 1990; Shay et al. 2001). A list and website has been developed for the insects<br />
collected at Aweme, which by 2004, contained about 2,000 species records and could<br />
well reach 3,000–4,000 records.<br />
Two additional areas within the southwestern Manitoba Aspen Parkland Ecoregion are<br />
collectively known as the Southwest Manitoba Uplands Ecoregion. This ecoregion consists<br />
<strong>of</strong> two distinct uplands: the Turtle Mountain Upland, straddling the border with the United<br />
States in the southwestern portion <strong>of</strong> the province, and the Pembina Hills Upland located in<br />
south-central Manitoba. Prior to settlement, the hills were primarily woodland composed <strong>of</strong><br />
trembling aspen and bur oak, with the latter on the drier and warmer slopes. Very dry sites<br />
on the hills had shrubs with grasses and herbs as ground cover. The Pembina Hills, which<br />
rise about 200 m above the adjacent Manitoba Plain to the east, is a warmer subdivision and<br />
outlier <strong>of</strong> the Boreal Ecoclimatic Region in Manitoba. The Turtle Mountain Ecodistrict is<br />
an outlier <strong>of</strong> the Boreal Plains Ecozone to the north and is a prominent wooded upland. The<br />
vegetation is largely intact and consists <strong>of</strong> deciduous trees and shrubs with an undergrowth<br />
<strong>of</strong> herbs and grasses.<br />
The cities <strong>of</strong> Edmonton and Saskatoon are the largest cities completely in the Aspen<br />
Parkland Ecoregion (Fig. 3). Winnipeg is bordered by tallgrass <strong>prairie</strong> to the west and south
19<br />
20<br />
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 73<br />
Figs. 19 and 20. The historically important homestead <strong>of</strong> the Criddle family southeast <strong>of</strong> Brandon, Manitoba. Fig.<br />
19. Remnant patch <strong>of</strong> tallgrass grassland in a pasture to the south <strong>of</strong> the Criddle farmhouse. Fig. 20. Remains <strong>of</strong><br />
Norman Criddle’s entomological field laboratory as it was in 2000, in bur oak forest to the west <strong>of</strong> the farmhouse.<br />
The laboratory has since been renovated by a local historical society. Photographs by the author.
74 J. D. <strong>Shorthouse</strong><br />
and Calgary is bordered by <strong>grasslands</strong> to the east and the Foothills Parkland to the west.<br />
Other communities include Red Deer, Lloydminster, North Battleford, and Humboldt.<br />
The communities <strong>of</strong> Brandon and Souris are in the Manitoba part <strong>of</strong> the ecoregion. Elk<br />
Island National Park in Alberta is near the middle <strong>of</strong> the ecoregion, whereas Prince Albert<br />
National Park in Saskatchewan is in the northern part <strong>of</strong> the ecoregion.<br />
Lake Manitoba Plain Ecoregion<br />
No living man will see again the long-grass <strong>prairie</strong>, where a sea <strong>of</strong> <strong>prairie</strong> flowers<br />
lapped at the stirrups <strong>of</strong> the pioneer.<br />
—Aldo Leopold, 1949<br />
The Lake Manitoba Plain Ecoregion stretches northwestward from the southern border<br />
with North Dakota to Dauphin west <strong>of</strong> Lake Manitoba (Fig. 2). The Manitoba Escarpment<br />
near Morden marks the eastern edge <strong>of</strong> the Aspen Parkland Ecoregion approximately<br />
110 km southwest <strong>of</strong> Winnipeg, which is the start <strong>of</strong> the Lake Manitoba Plain Ecoregion.<br />
The area where tallgrasses were once found occupies the southern half <strong>of</strong> this ecoregion<br />
stretching northward from the border to Winnipeg and westward to Morden. This region is<br />
the northern extension <strong>of</strong> the true North American tallgrass <strong>prairie</strong>, which stretches south<br />
into the United States to Texas (Risser et al. 1981; Madson 1982; Reichman 1987; Kucera<br />
1992; Madson and Oberle 1993). Before settlement, the entire ecoregion in Canada was a<br />
mosaic <strong>of</strong> trembling aspen, oak groves, and tallgrass <strong>grasslands</strong>.<br />
The Lake Manitoba Plain is the lowest and most level <strong>of</strong> three <strong>prairie</strong> steppes. It<br />
is covered by glacial till and by silts and clays, making it lower and smoother than the<br />
Saskatchewan Plain to the west. Slopes range from level to less than 2% and are smooth<br />
and long. Sandy, ancient beaches parallel to the escarpment indicate former shorelines <strong>of</strong><br />
glacial Lake Agassiz (Mayer-Oakes 1967). Heavy clay sediments that settled in the deep<br />
waters <strong>of</strong> Lake Agassiz that once covered the Red River Valley are found to the east. The<br />
Red River meanders northward through the ecoregion and empties into Lake Winnipeg<br />
(Fig. 4), a remnant <strong>of</strong> former glacial Lake Agassiz. Major tributaries <strong>of</strong> the Red River are<br />
the Morris, La Salle, and Assiniboine rivers flowing from the west (Fig. 4) and the Roseau,<br />
Rat, and Seine rivers, which flow from the east. The Red River Valley is characterized<br />
by extensive deep, clayey glaciolacustrine glacial lake sediments. These deposits are<br />
dominantly poorly drained Gleysolic, clay soils characterized by black surface horizons<br />
with a high organic content. The heavy clay tends to remain wet for some time in the<br />
spring. Nearly all <strong>of</strong> these soils are now cultivated and are some <strong>of</strong> the most productive<br />
soils found in western Canada. Soils to the north and northwest are Chernozemic.<br />
The native vegetation in the western and northern parts <strong>of</strong> this ecoregion once<br />
consisted <strong>of</strong> aspen and fescue grasses and many shrubs. In the southern regions,<br />
extensive stands <strong>of</strong> tallgrasses grew, along with groves <strong>of</strong> bur oak (Fig. 21), meadow<br />
<strong>prairie</strong> grass, and meadow grass communities, depending on the drainage conditions.<br />
Fires burning into bur oak-dominated stands maintained the large oaks with a grassy<br />
understory (Reichman 1987).<br />
Tallgrass <strong>prairie</strong> is one <strong>of</strong> the most endangered ecosystems in North America, with less<br />
than 1% remaining (Madson and Oberle 1993; Noss et al. 1995). In Manitoba, the natural<br />
vegetation throughout this ecoregion has largely disappeared as a result <strong>of</strong> cultivation, the<br />
reduction in frequency <strong>of</strong> <strong>prairie</strong> fires, and the development <strong>of</strong> an extensive network <strong>of</strong><br />
drainage ditches. The original 6,000 km 2 <strong>of</strong> tallgrass <strong>prairie</strong> <strong>of</strong> the Red River Basin is today
21<br />
22<br />
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 75<br />
Figs. 21 and 22. Typical remnant patches <strong>of</strong> bur oak in the Lake Manitoba Plain Ecoregion. Fig. 21. Patch <strong>of</strong><br />
tallgrass <strong>prairie</strong> at a site near Tolstoi protected by the Nature Conservancy <strong>of</strong> Canada. Fig. 22. Patch <strong>of</strong> remnant<br />
tallgrass <strong>prairie</strong> alongside a highway in the southeast part <strong>of</strong> the province. Photographs by the author.
76 J. D. <strong>Shorthouse</strong><br />
reduced to about 4 km 2 , or about 0.07% <strong>of</strong> its former area. Recent efforts to identify all<br />
remaining tallgrass <strong>prairie</strong> in Manitoba located only 81 ha <strong>of</strong> prime <strong>prairie</strong>, as well as 220<br />
ha <strong>of</strong> mostly atypical examples, all <strong>of</strong> it under private ownership. Some <strong>of</strong> these lands have<br />
recently received protection under the province’s Critical Wildlife Habitat Program and<br />
the Manitoba Naturalists Society’s Prairie Patron Program. Small pockets remain near the<br />
Canada–US border (Fig. 22) in poorly drained locales that were not broken. Some small<br />
patches with tallgrasses remain as far west as the Criddle/Vane homestead (Fig. 19). Three<br />
blocks totalling about 2,400 ha near Tolstoi and Gardenton in the south (Fig. 21) have been<br />
purchased and are maintained by The Nature Conservancy <strong>of</strong> Canada. The southern block<br />
contains large, deep marshes, whereas the northern block is drier with scattered sedge<br />
meadows. The central block is a mix <strong>of</strong> small <strong>prairie</strong> openings with large tracts <strong>of</strong> aspen<br />
and oak. Manitoba’s tallgrass <strong>prairie</strong> is considered somewhat different from the tallgrass in<br />
the United States because it once thrived on poorly drained lowland exposed after glacial<br />
Lake Agassiz drained.<br />
The Living Prairie Museum, within the city limits <strong>of</strong> Winnipeg (Fig. 23), is 12 ha<br />
<strong>of</strong> tallgrass <strong>prairie</strong> and is home to over 160 species <strong>of</strong> <strong>prairie</strong> plants. It is periodically<br />
burned, as fire is necessary to sustain tallgrass ecosystems, and has become a popular<br />
ecotourism destination. Another urban site, known as the St. Charles Rifle Range, owned<br />
by the Department <strong>of</strong> National Defence, is also within the city limits <strong>of</strong> Winnipeg (see<br />
Chapters 10 and 11). Today, tallgrass <strong>prairie</strong> is difficult to identify in some remaining<br />
sites because heavy grazing has caused the invasion <strong>of</strong> alien species and, in the absence<br />
<strong>of</strong> fire, western snowberry, smooth bromegrass, and aspen poplar commonly overtake<br />
the sites.<br />
The Lake Manitoba Plain Ecoregion is the warmest and most humid in the Canadian<br />
<strong>prairie</strong>s, with an annual temperature <strong>of</strong> around 2.5 °C. The mean summer temperature is<br />
16 °C and the mean winter temperature is −12.5 °C. Mean annual precipitation ranges from<br />
450 to 700 mm.<br />
The dominant species <strong>of</strong> grass in the tallgrass remnants are big bluestem, switch<br />
grass (Panicum virgatum L.), Indian grass (Sorghastrum nutans (L.) Nash), little bluestem<br />
(Schizachyrium scoparium (Michx.) Nash), Canada wild rye, and northern wheat grass<br />
(Scott 1995). Big bluestem is a tall, warm-season, sod-forming, highly nutritious grass that<br />
flourishes under a wide range <strong>of</strong> conditions. Its colour ranges from bluish-green in summer<br />
to reddish-brown in fall (Figs. 19, 21, and 22). It grows 1–2 m tall and has a fibrous root<br />
system down to 2 m. Switch grass is most common on <strong>prairie</strong> lowlands. However, this<br />
warm-season, sod-forming grass is also found in small upland patches. Its ability to remain<br />
in an upright position in snow makes for excellent winter habitat for wildlife. Indian grass<br />
is also a tall, warm-season, sod-forming grass that grows on a variety <strong>of</strong> soils and provides<br />
nutritious forage. It grows to a height <strong>of</strong> 2 m. Little bluestem is a medium-height, warmseason<br />
bunchgrass common on <strong>prairie</strong> uplands. In fall and winter, its reddish-brown colour<br />
makes it one <strong>of</strong> the most beautiful grasses on the <strong>prairie</strong>. Though rarely growing taller than<br />
1 m, it has a dense root system that may reach 2 m deep.<br />
A distinctive feature <strong>of</strong> the Lake Manitoba Plain Ecoregion is the riparian forests <strong>of</strong><br />
deciduous trees (Looman and Best 1987). Driving across the flat lowlands <strong>of</strong> the Lake<br />
Agassiz basin, one comes across what appears to be an abrupt forest margin, which is <strong>of</strong>ten<br />
only 100–200 m <strong>of</strong> trees surrounding the banks <strong>of</strong> the Red and Assiniboine rivers. Dominant<br />
trees here are American white elm (Ulmus americana L.), basswood (Tilia americana L.),<br />
green ash (Fraxinus pennsylvanica Marsh.), Manitoba maple (Acer negundo L.), and many<br />
species <strong>of</strong> Salix.
23<br />
24<br />
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 77<br />
Figs. 23 and 24. Urban tallgrass and grassland near Peace River, Alberta. Fig. 23. Remnant patch <strong>of</strong> tallgrass<br />
<strong>prairie</strong> within the city <strong>of</strong> Winnipeg. Note that the site to the right <strong>of</strong> the patch was burned the previous year. Fig. 24.<br />
Typical grassy plain in the Peace Lowland Ecoregion west <strong>of</strong> Peace River, Alberta. Photographs by the author.
78 J. D. <strong>Shorthouse</strong><br />
Major communities in the Lake Manitoba Plain Ecoregion include Winnipeg, Portage<br />
la Prairie, Emerson, and Dauphin.<br />
Boreal Plains Ecozone<br />
The Boreal Plains Ecozone lies north <strong>of</strong> the Prairies Ecozone (Fig. 1) and is mostly covered<br />
with either deciduous or coniferous forests. However, intermittent patches <strong>of</strong> <strong>grasslands</strong><br />
are found throughout the southern part <strong>of</strong> the ecozone, such as within the Prince Albert<br />
National Park (Fig. 17), north <strong>of</strong> Prince Albert, Saskatchewan. The most significant <strong>of</strong> these<br />
<strong>grasslands</strong> occurs in northwestern Alberta and adjoining northeastern British Columbia, an<br />
ecoregion known as the Peace Lowland (Fig. 1).<br />
The Boreal Plains Ecozone is part <strong>of</strong> the flat Interior Plains <strong>of</strong> Canada, a northern<br />
extension <strong>of</strong> the Great Plains <strong>of</strong> North America, and belongs to the vast boreal forest biome<br />
that extends across Canada. It occupies a land area <strong>of</strong> about 67.9 million ha extending from<br />
Alberta to Manitoba in a wide band from the Peace River district <strong>of</strong> British Columbia in<br />
the northwest to the southeastern corner <strong>of</strong> Manitoba (Fig. 1). Unlike the neighbouring<br />
Boreal Shield to the east, this ecozone is not bedrock controlled and has few rock outcrops<br />
and considerably fewer lakes. It comprises nine ecoregions, but only the one known as the<br />
Peace Lowland (Fig. 24) will be discussed here.<br />
The subdued relief <strong>of</strong> this ecozone consists <strong>of</strong> low-lying valleys and flat plains. Most<br />
<strong>of</strong> the major rivers have their origin in the Rockies and flow east across the ecozone and<br />
are the products <strong>of</strong> rainfall, snowmelt, and glacial run<strong>of</strong>f at their headwaters. About 7% <strong>of</strong><br />
the ecozone is covered by fresh water, with Lake Winnipeg and Lake Winnipegosis being<br />
the largest lake basins.<br />
The climate <strong>of</strong> the Boreal Plains Ecozone is influenced by its location in the heart <strong>of</strong><br />
North America. The Rocky Mountains to the west block moisture-bearing winds from the<br />
Pacific. The result is short, warm summers and long, cold winters. Mean annual temperatures<br />
vary between −2 and 2 °C; summer mean temperatures range from 13 to 15 °C and winter<br />
mean temperatures range from −17.5 to −11 °C. Northern Alberta receives an average <strong>of</strong><br />
300 mm precipitation annually, whereas southwestern Manitoba receives 625 mm. The<br />
annual precipitation throughout the ecozone is greater than the evaporation rate, resulting in<br />
surplus moisture <strong>of</strong> up to 100 mm near the southern edge <strong>of</strong> the ecozone and up to 300 mm<br />
in the northern and foothills regions.<br />
Soils <strong>of</strong> the forested areas <strong>of</strong> the ecozone are mostly imperfectly drained Gray Luvisols<br />
developed in loamy conditions under a forest canopy. Soils <strong>of</strong> the grassland areas in the<br />
Peace River District and other areas to the east in Saskatchewan and Manitoba are Dark<br />
Gray Chernozemics.<br />
Coniferous and deciduous trees cover about 84% <strong>of</strong> the Boreal Plains Ecozone and<br />
forestry is the primary industry. Common conifers are white spruce (Picea glauca (Moench)<br />
Voss) and black spruce (Picea mariana (Mill.)), balsam fir (Abies balsamea (L.)), jack pine<br />
(Pinus banksiana Lamb.), and tamarack (Larix laricina (Du Roi) K. Koch). The deciduous<br />
trees are aspen poplar (Populus tremuloides Michx.), balsam poplar (Populus balsamifera<br />
L.), and white birch (Betula papyrifera Marsh.). Less than 20% <strong>of</strong> the land area is devoted<br />
to agriculture. Precipitation and surface and groundwater sources are more than adequate<br />
to meet agricultural demand. This ecozone has traditionally been viewed by some people<br />
as the next untapped resource frontier. However, the exploration, production, and delivery<br />
<strong>of</strong> oil and gas products in the area, along with the removal <strong>of</strong> trees for lumbering and<br />
pulp and paper, has resulted in vast roads, railways, and pipeline networks and thousands
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 79<br />
<strong>of</strong> kilometres <strong>of</strong> seismic exploration lines cut through the forests, providing access to<br />
previously remote areas. Even so, the relative remoteness and absence <strong>of</strong> large population<br />
centres has resulted in little comprehensive ecological study in this ecozone.<br />
Peace Lowland Ecoregion<br />
The Peace Lowland Ecoregion is located in northwestern Alberta and northeastern British<br />
Columbia (Fig. 1). It is composed <strong>of</strong> gently undulating or sloping lands <strong>of</strong> mostly till<br />
deposits associated with the Peace River and its tributaries. Isolated patches <strong>of</strong> <strong>grasslands</strong><br />
(Fig. 24) are found throughout the region in close proximity to trembling aspen and balsam<br />
poplar, an understory <strong>of</strong> mixed herbs, and tall shrubs. The aspen and balsam poplars are<br />
in turn surrounded by coniferous forests. Grasslands throughout this ecoregion are more<br />
fragmented than those in the Aspen Parkland Ecoregion. The rivers in this ecoregion are<br />
incised up to 300 m in the foothills <strong>of</strong> northeastern British Columbia, where the elevation<br />
<strong>of</strong> the valleys is about 750 m.<br />
This region has shorter, cooler summers and longer, colder winters than does the Aspen<br />
Parkland Ecoregion to the south. The mean annual temperature is 0.5 °C. The mean summer<br />
temperature is 13 °C and the mean winter temperature is −14 °C. The frost-free period<br />
averages about 95 days each year. The mean annual precipitation ranges from 350 to 600 mm.<br />
Clayey lacustrine deposits are the predominant parent material, along with finetextured<br />
tills and significant areas <strong>of</strong> sandy fluvial deposits. Gray Luvisols are the<br />
predominant soil types, but the abundant vegetative understory and restricted drainage<br />
have resulted in an abundance <strong>of</strong> Dark Gray Luvisols, with some Chernozemic and<br />
Solonetzic soils in the Alberta portion. About 45% <strong>of</strong> the area is farmland, with annual<br />
small grains and grasses dominating.<br />
Grasslands here are dominated by sedges, intermediate oat grass (Danthonia intermedia<br />
Vasey), western porcupine grass (Stipa curtiseta Hitchc.), bearded wheat grass (Elymus<br />
caninus (L.)), inland bluegrass (Poa nemoralis L.), old man’s whiskers (Geum triflorum<br />
Pursh), and low goldenrod (Solidago nana Nutt.). Moss (1952) recorded 154 species <strong>of</strong><br />
vascular plants in the area comprising 29 grasses, 9 sedges, and 96 forbs. Grasslands on<br />
steep, south-facing slopes along river valleys are dominated by western porcupine grass<br />
(Stipa curtiseta Hitch), sedges, and pasture sage (Artemisia frigida Willd.). Other common<br />
species include Columbia needlegrass (Achnatherum nelsonii (Scribn.) Barkworth), June<br />
grass (Koeleria macrantha (Ledeb.) Schult.), and green needlegrass (Nassella viridula<br />
(Trin.) Barkworth). Many <strong>of</strong> the <strong>grasslands</strong> are dotted with willow groves and dense<br />
thickets <strong>of</strong> western snowberry (Symphoricarpos occidentalis Hook.) and wild roses (Rosa<br />
acicularis Lindl. and Rosa woodsii Lindl.).<br />
The isolated <strong>grasslands</strong> <strong>of</strong> the Peace Lowland Ecoregion are remarkable for the presence<br />
and disjunct occurrence <strong>of</strong> species that have a more southerly or westerly distribution,<br />
including prickly-pear cactus Opuntia polyacantha Haw., Richardson’s needlegrass<br />
(Achnatherum richardsonii (Link) Barkworth), short-stemmed thistle (Cirsium drummondii<br />
Torr. and A. Gray), and groundsel (Senecio vulgaris L.). Moss (1952) suggested that the<br />
Peace River <strong>grasslands</strong> are remnants <strong>of</strong> a past period <strong>of</strong> prolonged warmth and dryness,<br />
between 8,000 and 4,000 years ago, and that the region may have been connected with the<br />
<strong>grasslands</strong> <strong>of</strong> south-central Alberta.<br />
The main part <strong>of</strong> the ecoregion is in the Grande Prairie and Peace River areas <strong>of</strong><br />
Alberta, with smaller areas as far north as Fort Vermilion. In British Columbia, <strong>grasslands</strong><br />
are found near Dawson Creek and Fort St. John. Other communities on the Alberta side<br />
include Fairview and Manning.
80 J. D. <strong>Shorthouse</strong><br />
Acknowledgements<br />
I thank Léo Larivière <strong>of</strong> the Department <strong>of</strong> Geography at Laurentian University for drawing<br />
the map showing <strong>grasslands</strong> <strong>of</strong> Canada and David Larson for his suggestions on improving<br />
the text. Funds to visit various sites mentioned in this chapter were provided by an NSERC<br />
Discovery Grant and Laurentian University.<br />
References<br />
Abouguendia, Z.M. 1996. Seeded Native Range Plants. Grazing and Pasture Technology Program, Regina,<br />
Saskatchewan.<br />
Acton, D.F., Padbury, G.A., and Stushn<strong>of</strong>f, C.T. 1998. The <strong>Ecoregions</strong> <strong>of</strong> Saskatchewan. Canadian Plains<br />
Research Centre, University <strong>of</strong> Regina, Regina, Saskatchewan.<br />
Aiken, S.G., Dallwitz, M.J., McJannet, C.L., and Consaul, L.L. 1997. Biodiversity among Festuca (Poaceae)<br />
in North America: diagnostic evidence from DELTA and clustering programs, and an INTKEY package<br />
for interactive, illustrated identification and information retrieval. Canadian Journal <strong>of</strong> Botany, 75: 1527–<br />
1555.<br />
Aiken, S.G., and Darbyshire, S.J. 1990. Fescue Grasses <strong>of</strong> Canada. Ministry <strong>of</strong> Supply and Services Canada,<br />
Ottawa, Ontario. Publication 1844E.<br />
Beaty, C.B. 1975. The Landscapes <strong>of</strong> Southern Alberta. The University <strong>of</strong> Lethbridge Printing Services,<br />
Lethbridge, Alberta.<br />
Best, K.F., J. Looman, and J. Baden Campbell, 1971. Prairie Grasses. Information Canada, Ottawa, Ontario.<br />
Bird, C.D. 1962. Bryophytes <strong>of</strong> the Cypress Hills Provincial Parks, Alberta and Saskatchewan. Canadian<br />
Journal <strong>of</strong> Botany, 40: 573–587.<br />
Bird, R.D. 1961. Ecology <strong>of</strong> the Aspen Parkland <strong>of</strong> Western Canada. Queen’s Printer, Ottawa, Ontario.<br />
Publication 1066.<br />
Bragg, T.B., and Steuter, A.A. 1996. Mixed-grass <strong>prairie</strong>s <strong>of</strong> the North American Great Plains. In Prairie<br />
Conservation: Preserving North America’s Most Endangered Ecosystem. Edited by F.B. Samson and F.L.<br />
Knopf. Island Press, Covelo, California. pp. 53–66.<br />
Breitung, A.J. 1954. A botanical survey <strong>of</strong> the Cypress Hills. Canadian Field-Naturalist, 68: 55–92.<br />
Coupland, R.T. 1961. A reconsideration <strong>of</strong> grassland classification in the northern great plains <strong>of</strong> North America.<br />
Journal <strong>of</strong> Ecology, 49: 135–167.<br />
Coupland, R.T. 1992. Mixed <strong>prairie</strong>. In Natural Grasslands: Introduction and Western Hemisphere. Ecosystems<br />
<strong>of</strong> the World 8a. Edited by R.T. Coupland. Elsevier, New York. pp. 151–182.<br />
Coupland, R.T., and Brayshaw, T.C. 1953. The fescue grassland in Saskatchewan. Ecology, 34: 386–405.<br />
David, P.P. 1993. Great Sandhills <strong>of</strong> Saskatchewan: an overview. In Quaternary and Late Tertiary Landscapes<br />
<strong>of</strong> Southwestern Saskatchewan and Adjacent Areas. Edited by D.J. Sauchyn. Canadian Plains Research<br />
Center, Regina, Saskatchewan. pp. 59–81.<br />
Doormaar, J.F., and Willms, W.D. 1990. Sustainable production from rough fescue <strong>prairie</strong>. Journal <strong>of</strong> Soil and<br />
Water Conservation, 45:137–140.<br />
Ecological Stratification Working Group. 1996. A National Ecological Framework for Canada. Agriculture<br />
and Agri-Food Canada, Research Branch, Centre for Land and Biological Resource Research and<br />
Environment Canada, State <strong>of</strong> the Environment Directorate, Ecozone Analysis Branch, Ottawa/Hull,<br />
Ontario. (Map.)<br />
Environment Canada. 1996. The State <strong>of</strong> Canada’s Environment. Public Works and Government Services,<br />
Ottawa, Ontario.<br />
Farrar, J.L. 1995. Trees in Canada. Fitzhenry and Whiteside Limited, Markham, Ontario and Canadian Forest<br />
Service, Ottawa, Ontario.<br />
Gauthier, D.A., and Wiken, E.B. 2003. Monitoring the conservation <strong>of</strong> grassland habitats, <strong>prairie</strong> ecozone,<br />
Canada. Journal <strong>of</strong> Ecological Monitoring and Assessment, 8: 343–364.<br />
Harms, V.L. 1985. A reconsideration <strong>of</strong> the nomenclature and taxonomy <strong>of</strong> the Festuca altaica complex in<br />
North America. Madroña, 32: 1–10.<br />
Harms, V.L. 2003. Checklist <strong>of</strong> the Vascular Plants <strong>of</strong> Saskatchewan and the Provincially and Nationally Rare<br />
Native Plants in Saskatchewan. University <strong>of</strong> Saskatchewan, Saskatoon.<br />
Hildebrandt, W., and Hubner, B. 1994. The Cypress Hills: The Land and Its People. Purich Publishing,<br />
Saskatoon, Saskatchewan.
<strong>Ecoregions</strong> <strong>of</strong> Canada’s Prairie Grasslands 81<br />
Hulett, G.K., Coupland, R.T., and Dix, R.L. 1966. The vegetation <strong>of</strong> dune sand areas within the grassland region<br />
<strong>of</strong> Saskatchewan. Canadian Journal <strong>of</strong> Botany, 44:1307–1331.<br />
Kucera, C.L. 1992. Tall grass <strong>prairie</strong>. In Natural Grasslands: Introduction and Western Hemisphere. Ecosystems<br />
<strong>of</strong> the World 8a. Edited by R.T. Coupland. Elsevier, New York. pp. 227–268.<br />
Lemmen, D.S., Vance, R.E., Wolfe, S.A., and Last, W.M. 1997. Impact <strong>of</strong> future climate change on the southern<br />
Canadian <strong>prairie</strong>s: a paleoenvironmental perspective. Geoscience Canada, 24: 121–133.<br />
Leopold, A. 1949. A Sand County Almanac. Oxford University Press, New York.<br />
Looman, J. 1983. 111 Range and Forage Plants <strong>of</strong> the Canadian Prairies. Agriculture Canada, Ottawa, Ontario.<br />
Publication 1751.<br />
Looman, J., and Best, K.F. 1987. Budd’s Flora <strong>of</strong> the Canadian Prairie Provinces. Research Branch, Agriculture<br />
Canada Publication 1662, Canadian Government Publishing Centre, Hull, Quebec.<br />
Madson, J. 1982. Where the Sky Began: Land <strong>of</strong> the Tallgrass Prairie. Houghton Mifflin, Boston,<br />
Massachusetts.<br />
Madson, J., and Oberle, F. 1993. Tallgrass Prairie. Falcon Press, Helena, Montana.<br />
Mayer-Oakes, W.J. (Editor) 1967. Life, Land and Water. Proceedings <strong>of</strong> the 1966 Conference on Environmental<br />
Studies <strong>of</strong> the Glacial Lake Agassiz Region, University <strong>of</strong> Manitoba Press, Winnipeg.<br />
Moss, E.H. 1952. Grassland <strong>of</strong> the Peace River region, western Canada. Canadian Journal <strong>of</strong> Botany, 30:<br />
98–124.<br />
Moss, E.H., and Packer, J.G. 1983. Flora <strong>of</strong> Alberta, 2nd edition. University <strong>of</strong> Toronto Press, Toronto, Ontario.<br />
Newsome, R.D., and Dix, R.L. 1968. The forests <strong>of</strong> the Cypress Hills, Alberta and Saskatchewan, Canada.<br />
American Midland Naturalist, 80: 118–185.<br />
Noss, R.F., LaRoe, E.T., and Scott, J.M. 1995. Endangered Ecosystems <strong>of</strong> the United States: A Preliminary<br />
Assessment <strong>of</strong> Loss and Degradation. USDI National Biological Service, Washington, D.C. Biological<br />
Report 28.<br />
Pavlick, L.E., and Looman, J. 1984. Taxonomy and nomenclature <strong>of</strong> rough fescues, Festuca altaica, F.<br />
campestris (F. scabrella var. major) and F. hallii, in Canada and the adjacent part <strong>of</strong> the United States.<br />
Canadian Journal <strong>of</strong> Botany, 62: 1739–1748.<br />
Reichman, O.J. 1987. Konza Prairie: A Tallgrass Natural History. University Press <strong>of</strong> Kansas, Lawrence.<br />
Risser, P.G., Birney, P.G., Blocker, H.D., May, S.W., Parton, W.J., and Wiens, J.A. 1981. The True Prairie<br />
Ecosystem. Hutchinson Ross Publishing Company, Stroudsburg, Pennsylvania.<br />
Roughley, R.E. 2000. Aweme, Manitoba: an important historical grassland site. Newsletter, Arthropods <strong>of</strong><br />
Canadian Grasslands, 6: 6–12.<br />
Rowe, J.S., and Coupland, R.T. 1984. Vegetation <strong>of</strong> the Canadian plains. Prairie Forum, 9: 231–248.<br />
Scott, G.A.J. 1995. Canada’s Vegetation: A World Perspective. McGill-Queen’s University Press, Montreal,<br />
Quebec.<br />
Shay, J.M., Kunec, D.L., and Dyck, B. 2001. Short-term effects <strong>of</strong> fire frequency on vegetation composition and<br />
biomass in mixed <strong>prairie</strong> in south-western Manitoba. Plant Ecology, 155: 157–167.<br />
<strong>Shorthouse</strong>, J.D. 1991. An unusual population <strong>of</strong> galls <strong>of</strong> Diplolepis polita (Hymenoptera: Cynipidae) in the<br />
Cypress Hills <strong>of</strong> southeastern Alberta. Canadian-Field Naturalist, 105: 542–549.<br />
Stout, D.G., McLean, A., and Quinton, D.E. 1981. Growth and phenological development <strong>of</strong> rough fescue in<br />
interior <strong>of</strong> British Columbia. Journal <strong>of</strong> Range Management, 34: 455–495.<br />
Willms, W.D. 1987. Response <strong>of</strong> rough fescue (Festuca scarabella) to light, water, temperature, and litter<br />
removal, under controlled conditions. Canadian Journal <strong>of</strong> Botany, 66: 429–434.<br />
Willms, W.D. Adams, B.W., and Doormaar, J.F. 1996. Seasonal changes <strong>of</strong> herbage biomass on the fescue<br />
<strong>prairie</strong>. Journal <strong>of</strong> Range Management, 49: 100–104.<br />
Wilson, S.D., and Shay, J.M. 1990. Competition, fire and nutrients in a mixed-grass <strong>prairie</strong>. Ecology, 71:<br />
1959–1967.<br />
Wolfe, S.A., and David, P.P. 1997. Parabolic dunes <strong>of</strong> the Great Sand Hills, southwestern Saskatchewan.<br />
Canadian Geographer, 41: 207–213.