Arthropods of Canadian Grasslands (Volume 1): Ecology and Interactions in Grassland Habitats

Edited by Joseph D. Shorthouse Laurentian University

and

Kevin D. Floate Agriculture and Agri-Food Canada

Biological Survey of Canada Monograph Series No. 3 (2010)

ii J. D. Shorthouse and K. D. Floate

Published by the BIOLOGICAL SURVEY OF CANADA

©2010

All illustrations in this book are copyright by individual authors, and their unauthorized use is prohibited. For further information contact the Biological Survey of Canada.

The Biological Survey of Canada develops and coordinates national initiatives in systematic and faunistic entomology on behalf of the Entomological Society of Canada and the Canadian Museum of Nature.

The monograph series of the Biological Survey of Canada comprises invited, fully reviewed publications of record that are especially relevant to the fauna of Canada.

ISBN 978-0-9689321-4-8

doi:10.3752/9780968932148

Arthropods of Canadian Grasslands (Volume 1): Ecology and Interactions in Grassland Habitats iii

Preface

The Biological Survey of Canada is a national organization based at the Canadian Museum of Nature in Ottawa, Ontario that was established in 1977 to coordinate research on Canada’s arthropod fauna (http://www.biology.ualberta.ca/bsc/bschome.htm). Several years ago, it began a project to coordinate research on the diversity and ecology of the arthropods of Canadian grasslands. This book is the first of a planned three volume set and is an overview of the attributes of Canada’s grasslands and grassland ecosystems, along with an introduction to the kinds of complex interactions exhibited by grassland arthropods.

Canada’s extensive grasslands have only attained their present distribution in the past 10,000 years, since the last glaciation. They extend across most of the prairie regions of southern portions of Alberta, Saskatchewan and Manitoba, with smaller expanses in the Yukon, the interior of British Columbia, and in southwestern Ontario. These regions are dominated by species of grasses that are adapted to thrive in conditions of drought and frequent fires. Many of their associated arthropods also are uniquely adapted for this environment. Some species occupy specific microhabitats that shelter them from prevailing conditions, whereas others are generalists that have the capacity to cope with the vagaries of the environment. This book deals with a sampling of the arthropods that cope in grassland environments.

The physical and biotic attributes of all grasslands in Canada have been shaped by natural events since the last glacial period, such as fire, drought and herbivory by large and small vertebrates; however, it was the arrival of European settlers that precipitated widespread losses in habitat quality and natural biodiversity as habitats were converted to a landscape designed to produce food for humans. History has recorded that within half a century, most of our grasslands were converted to agroecosystems for growing crops and pastures for feeding cattle. These modified grasslands are dominated by plant monocultures, support a relatively sparse arthropod fauna, and require constant human intervention with large farm implements to retain their artificial productivity. Because of their importance for food production, most of our detailed knowledge of grassland arthropods is limited to pest species, some of which are not native to North America. Conversely, we know little about the biodiversity and ecology of native arthropod species and we are rapidly losing the opportunity to learn more. Only a fraction of Canada’s original grasslands remain unmodified, and yet these scattered remnants support hundreds of arthropod species for which many remain unknown to science.

Publication of Arthropods of Canadian Grasslands is intended to increase awareness of Canada’s vanishing grasslands and their associated arthropods. Hopefully it will provide a baseline reference for future studies of arthropods in these environments. We have deliberately targeted a broad audience that includes post-secondary students, teachers, farmers, ranchers and naturalists. To increase the appeal of the series, we have encouraged authors to liberally illustrate their chapters with photographs and images, and to present content in an ‘easily digestible’ manner. There remains, however, considerable variation among chapters, reflecting their contributors’ expertise and interests.

This first volume in our project begins with a series of chapters that provide a context for subsequent chapters, including those in Volume II. Chapter 1 (Shorthouse and Larson) reviews the formation of grasslands in Canada over geological time, and provides general information on the common groups of arthropods associated with grasslands. Chapter 2 (Shorthouse) reviews the effects of glaciation on prairie grasslands, the consequences of soil type and fire, the adaptations of native plants to a grassland environment, and the types

iv J. D. Shorthouse and K. D. Floate

of aquatic habits found on the prairies. Chapter 3 (Shorthouse) reviews the characteristics and extent of the different ecoregions that support the different types of grasslands on the Central Plains of western Canada. Chapter 4 (Shorthouse) provides similar information for the grasslands of British Columbia, the Yukon and southern Ontario. Chapter 5 (McGinn) completes this background context by reviewing climate and weather patterns on the Canadian prairies.

The remainder of chapters in Volume I have a specific focus on arthropods; either as a general review of a taxonomic group or in association with a specific habitat. Chapter 6 (Scudder) summarizes results of research that identify grassland locations in British Columbia of particular importance for the conservation of potentially rare and endangered species. Chapter 7 (Behan-Pelletier & Kanashiro) examines the diversity of mites (Acari) inhabiting grassland soils and its variation across grassland ecoregions. Chapter 8 (Hamilton & Whitcomb) is a synthesis of the biogeography and ecology of leafhoppers (Homoptera: Cicadellidae) in grassland habitats. Chapter 9 (Paiero, Marshall, Pratt & Buck) reviews the arthropod assemblage of Ojibway Prairie, a remnant tallgrass prairie in south western Ontario. Chapters 10 (Roughley, Pollock & Wade) and 11 (Wade & Roughley) examine the effects on ground beetles (Coleoptera: Carabidae) and spiders (Araneae), respectively, of different fire management strategies on remnant tallgrass prairie near Winnipeg, Manitoba. Chapter 12 (Shorthouse) examines the biology and ecology of cynipid wasps that induce galls on different species of grassland roses. Chapter 13 (Floate) reviews the biology and ecology of aphids and mites that induce galls on cottonwood trees, whose populations often form the only treed areas on the prairies. Chapter 14 (Scudder, Alperyn and Roughley) reviews the occurrence of aquatic Hemiptera in the prairie provinces and factors affecting their distribution. The book ends with Chapter 15 (Shorthouse & Larson) which summarizes the role of grassland arthropods and discusses the plight of grasslands in Canada. It also reviews some of the attempts to preserve tracts of semi-natural and natural grasslands. Joseph D. ShorthouseSudbury, Ontario

Kevin D. FloateLethbridge, Alberta

Arthropods of Canadian Grasslands (Volume 1): Ecology and Interactions in Grassland Habitats v

This book is dedicated to our friend and colleague Robert Edward Roughley, who passed away suddenly on November 9, 2009. Rob was a passionate entomologist who loved learning and talking about all aspects of insect science, especially the inhabitants of tallgrass prairie and the subarctic of northern Manitoba, and his beloved dytiscid beetles. Co-author of three chapters in this book, Rob was able to view the galleys, but sadly did not get to see the finished product.

Rob completed his BSc (Agriculture) in Entomology in the Department of Environmental Biology at the University of Guelph in 1974. He became immersed in beetle taxonomy and systematics and remained at the University of Guelph to revise the genus Hydatiscus (Dytiscidae) for his MSc. He then went to the Department of Entomology at the University of Alberta to revise the genus Dytiscus for his PhD under the mentorship of George Ball. He accepted a faculty position in the Department of Entomology at the University of Manitoba, where he remained until his untimely death.

Rob was an untiring supporter of insect collections and collection management, and as curator of the J. B. Wallis Museum, he was key to its recent expansion and implemented one of the first bar-coded databases for entomological museums in Canada. He was a zealous contributor to the Biological Survey of Canada, the Nature Conservancy of Canada, the Committee on the Status of Endangered Wildlife in Canada, and the Canadian Pollination Initiative (CANPOLIN) based at the University of Guelph.

Robert Edward Roughley, 1950-2009. Photograph courtesy of The University of Manitoba Imaging Services.

Dedication to Robert E. Roughley

vi J. D. Shorthouse and K. D. Floate

Rob was a long-time member of the Scientific Committee of the Biological Survey of Canada (1991-1994, 1999-2009), and a strong supporter of the BSC’s annual BioBlitzes; i.e., surveys of arthropods at grassland sites in Canada. With Kevin Floate, he co-organized the first one in 2001 (Onefour, AB) and participated in BioBlitzes in 2003 (Peace River, AB), 2004 (Aweme, MB), 2005 (Waterton Lakes National Park, AB) and 2007 (Riding Mountain National Park, MB). Rob took the lead in designing the flag of the BSC and proudly displayed it wherever entomologists gathered to investigate arthropod biodiversity. He co-authored the popular BSC brief “Terrestrial Arthropod Biodiversity: Planning a Study and Recommended Sampling Techniques”, and regularly presented at provincial and national meetings. His last presentation was on beetles of the Churchill area, part of a BSC symposium at the Joint Annual Meeting of the Entomological Societies of Canada and Manitoba, held in Winnipeg in October of 2009.

I had the pleasure of visiting Rob in September of 2000 for three days of exploring the remaining tallgrass prairie of southern Manitoba. I was struck by his humour and comradeship as we drove along highways and side roads sharing collecting stories and anecdotes. I was also captivated by the enthusiasm with which he shared his knowledge of the sites we visited including the Criddle/Vane homestead southeast of Brandon, the St. Charles Rifle Range west of Winnipeg, and the Manitoba Tallgrass Prairie Preserve in the southeastern part of the province. Figures 19, 20 and 21 in Chapter 3 were taken during these excursions.

Joe ShorthouseSudbury, Ontario

Rob Roughley (second from right) displaying the flag of the Biological Survey of Canada at the 2005 Waterton Lakes National Park BioBlitz. Also with Rob are from the left, BSC members Joe Shorthouse, Felix Sperling, and Dave Langor. Photograph by Andrea Renelli.

Arthropods of Canadian Grasslands (Volume 1): Ecology and Interactions in Grassland Habitats vii

Contents

Preface ...............................................................................................................................iii

Dedication to Robert E. Roughley ................................................................................... v

List of Contributors ......................................................................................................... ix

Chapter 1. Introduction to the grasslands and grassland arthropods of Canada .......................... 1 J. D. Shorthouse and D. J. Larson

Chapter 2. Canada’s grasslands as habitat for arthropods ............................................................ 27 J. D. Shorthouse

Chapter 3. Ecoregions of Canada’s prairie grasslands................................................................... 53 J. D. Shorthouse

Chapter 4. Ecoregions with grasslands in British Columbia, the Yukon, and southern Ontario ..................................................................................................... 83 J. D. Shorthouse

Chapter 5. Weather and climate patterns in Canada’s prairie grasslands ................................. 105 S. M. McGinn

Chapter 6. Grasslands: biodiversity hotspots for some arthropods in British Columbia ......... 121 G. G. E. Scudder

Chapter 7. Acari in grassland soils of Canada .............................................................................. 137 V. M. Behan-Pelletier and D. Kanashiro

Chapter 8. Leafhoppers (Homoptera: Cicadellidae): a major family adapted to grassland habitats ......................................................................................................... 169 K. G. A. Hamilton and R. F. Whitcomb

Chapter 9. Insects of Ojibway Prairie, a southern Ontario tallgrass prairie ............................. 199 S. M. Paiero, S. A. Marshall, P. D. Pratt and M. Buck

Chapter 10. Tallgrass prairie, ground beetles (Coleoptera: Carabidae) and the use of fire as a biodiversity and conservation management tool ..................................... 227 R. E. Roughley, D. A. Pollock and D. J. Wade

viii J. D. Shorthouse and K. D. Floate

Chapter 11. Responses of a tallgrass prairie spider (Araneae) community to various burn seasons and its importance to tallgrass prairie management .......................... 237 D.J. Wade and R.E. Roughley

Chapter 12. Galls induced by cynipid wasps of the genus Diplolepis (Hymenoptera: Cynipidae) on the roses of Canada’s grasslands ............................. 251 J. D. Shorthouse

Chapter 13. Gall-inducing aphids and mites associated with the hybrid complex of cottonwoods, Populus spp. (Salicaceae) on Canada’s grasslands ......................... 281 K. D. Floate

Chapter 14. Aquatic Hemiptera of the prairie grasslands and parkland ..................................... 303 G. G. E. Scudder, M. A. Alperyn and R. E. Roughley

Chapter 15. Arthropods in Canada’s grasslands: synthesis and future directions ...................... 325 J. D. Shorthouse and D. J. Larson

Species index for arthropods ........................................................................................ 345

Species index for plants ................................................................................................ 353

Subject index ................................................................................................................. 356

Arthropods of Canadian Grasslands (Volume 1): Ecology and Interactions in Grassland Habitats ix

List of ContributorsAlperyn, Michael A. Canadian Centre for Mosquito Management in Winnipeg, Manitoba, 471 Craig Street, Winnipeg, Manitoba, R3G 3C2.

Behan-Pelletier, Valerie M. Agriculture and Agri-Food Canada, Eastern Cereal and Oilseed Research Centre, Ottawa, Ontario, K1A 0C6.

Buck, Matthias Department of Environmental Biology, University of Guelph, Guelph, Ontario, N1G 2W1.

Floate, Kevin D. Agriculture and Agri-Food Canada, Lethbridge Research Centre, Lethbridge, Alberta, T1J 4B1.

Hamilton, K. G. Andrew Agriculture and Agri-Food Canada, Eastern Cereal and Oilseed Research Centre, Ottawa, Ontario, K1A 0C6.

Kanashiro, Derrick. Agriculture and Agri-Food Canada, Lethbridge Research Centre, Lethbridge, Alberta, T1J 4B1.

Larson, David J. Box 56, Maple Creek, Saskatchewan, S0N 1N0.

Marshall, Stephen A. Department of Environmental Biology, University of Guelph, Guelph, Ontario, N1G 2W1.

McGinn, Sean M. Agriculture and Agri-Food Canada, Lethbridge Research Centre, Lethbridge, Alberta, T1J 4B1.

Paiero, Steve M. Department of Environmental Biology, University of Guelph, Guelph, Ontario, N1G 2W1.

Pollock, Darren A. Department of Biology, Eastern New Mexico University, Portales, New Mexico, 88130 USA.

Pratt, Paul D. Windsor Parks and Recreation, 2450 McDougall Street, Windsor, Ontario, N8X 3N6.

Roughley, Robert E. (deceased) Department of Entomology, University of Manitoba, Winnipeg, Manitoba, R3T 2NT.

Scudder, Geoffrey G. E. Department of Zoology, University of British Columbia, Vancouver, British Columbia, V6T 1Z4.

Shorthouse, Joseph D. Department of Biology, Laurentian University, Sudbury, Ontario, P3E 2C6.

Wade, David J. Surveillance Program Coordinator, City of Winnipeg Insect Control Branch, 1539 Waverley Street, Winnipeg, Manitoba, R2M 1T4.

Whitcomb, Robert F. (Deceased) Plant Virology Laboratory, Crops Research and Entomology Research Division, USA Department of Agriculture, Agricultural Research Service, Beltsville, Maryland 20705, USA.