Hemiptera: Auchenorrhyncha: Cicadellidae)
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311 Chapter 11 Canadian Grasslands and Their Endemic Leafhoppers (Hemiptera: Auchenorrhyncha: Cicadellidae) K.G.A. Hamilton Canadian National Collection of Insects K.W. Neatby Bldg., 960 Carling Ave. Ottawa, Ontario, Canada, K1A 0C6 Abstract. The greatest numbers of leafhopper species endemic to Canada are those of grasslands, 33 of which are known. A far larger number, 263 species, are more broadly endemic to Nearctic grasslands. All 296 demonstrate strong regional differences not widely recognized in ecoregional classifi cations. Most endemics come from the open prairies (174 species). Second most numerous are the various faunas on comparatively tiny intermontane grasslands of southern British Columbia with 126 endemics, including the newly recorded Dikrella nevadensis (Lawson), here reinstated from synonymy. Of these endemics, 96 are found in the Okanagan-Similkameen valley system, 43 belong to the East Kootenays, and another 43 are found near Victoria and Vancouver, with numerous overlaps. One is known only from North Beach on the Haida Gwaii (Queen Charlotte) Islands and this is related to an endemic on Vancouver Island and the adjacent mainland. The fauna of the isolated, more northerly fescue grasslands have 80 endemics (39 in the Peace River district, 25 in the Thompson valley system and 43 in the lower Fraser River valley and its lesser tributaries), and these are similar to the fauna of aspen parkland. An additional 58 endemics are characteristic of the Prairie Peninsula, of which 21 are found on the tallgrass prairie of southernmost Ontario, as contrasted to 23 from alkaline fens, 21 from beaches, 16 from alvars, and 4 from sand plains, many of which are shared with oak savanna. Beringian grasslands have 24 grassland endemics, two of which inhabit arctic areas of the Yukon. Each of these isolated grasslands and their subdivisions have their own endemics and unique histories, many traceable to separate glacial-age refugia. Understanding the rich endemic biota of the many distinctive grassland types in Canada is essential in characterizing their ecology and in setting conservation priorities. Résumé. Les prairies canadiennes renferment le plus grand nombre d’espèces endémiques de cicadelles du pays, dont 33 sont connues. Le nombre d’espèces généralement endémiques aux prairies néarctiques est beaucoup plus considérable, s’établissant à 263. Toutes ces 296 espèces laissent constater d’importantes différences régionales qui ne sont pas largement reconnues dans le système de classifi cation des écorégions. La plupart des espèces endémiques habitent des prairies dégagées (174). Au second rang viennent les diverses entomofaunes vivant dans les prairies intermontagneuses de superfi cie comparativement plus petite du sud de la Colombie-Britannique, où l’on trouve 126 espèces endémiques, y compris une nouvelle mention avec Dikrella nevadensis (Lawson), ici rétabli comme espèce valide. De toutes ces espèces endémiques, 96 se trouvent dans le système de vallées Okanagan-Similkameen, 43 se trouvent dans la portion orientale du bassin de la Kootenay, et 43 autres se trouvent près de Victoria et de Vancouver, plusieurs coexistant dans le même habitat. Une des espèces n’est connue qu’à North Beach, dans l’archipel Haida Gwaii (Îles de la Reine-Charlotte), et s’apparente à une espèce endémique de l’île de Vancouver et de la zone continentale adjacente. La faune de la prairie à fétuque, isolée et plus nordique, compte 80 espèces endémiques (39 dans le district de Peace River, 25 dans la vallée de la rivière Thompson et 43 dans la vallée du bas Fraser et de ses affl uents), et ressemble à la faune de la tremblaie-parc. Cinquante-huit autres espèces endémiques sont caractéristiques de la péninsule de Prairie, parmi lesquelles 21 s’observent dans la prairie à herbes hautes de l’extrême-sud de l’Ontario, alors qu’on en trouve 23 dans les tourbières alcalines, 21 sur les plages, 16 dans les alvars et 4 sur les deltas d’eskers, dont plusieurs se trouvent également dans les chênaies-parcs. Les prairies béringiennes renferment 24 espèces endémiques des prairies, dont deux habitent Hamilton, K. G. A. 2014. Canadian Grasslands and Their Endemic Leafhoppers (Hemiptera: Auchenorrhyncha: Cicadellidae). In Arthropods of Canadian Grasslands (Volume 3): Biodiversity and Systematics Part 1. Edited by H. A. Cárcamo and D. J. Giberson. Biological Survey of Canada. pp. 311-345. © 2014 Biological Survey of Canada. ISBN 978-0-9689321-6-2 doi: http://dx.doi.org/10.3752/9780968932162.ch11 312 K. G. A. Hamilton les régions arctiques du Yukon. Chacune de ces prairies isolées et de leurs subdivisions possèdent leurs propres espèces endémiques et présentent une histoire qui leur est propre et qui remonte souvent à des refuges distincts de l’âge glaciaire. Il est essentiel de bien connaître les riches biotes endémiques des nombreux types distinctifs de prairies du Canada pour pouvoir en caractériser l’écologie et établir les priorités en matière de conservation. Introduction Grasslands in Canada include some of the most endangered habitats in the country. They are usually in prime agricultural areas and are also easily converted to other uses. Conserving the remaining areas and their endemic insects is an urgent, ever-growing problem. Unfortunately, grasslands are usually described in ecological generalizations that have little value to conservation priorities. The oldest such classifi cation system differentiates eastern from western prairies according to the height of dominant grass species. This is simple, convenient, and still often cited (e.g., Anderson 2006). Such generalizations are misleading, however, because they are subject to extreme variation. For example, in Manitoba, big bluestem (the most prominent tall grass of the eastern grasslands) grows more than 1 m tall immediately after a burn, or when summer temperatures are higher than usual; otherwise, it may grow prostrate like crab grass (pers. obs.). Biogeographical regions of the North American prairies are more accurately represented by the species of dominant and subdominant plants (Coupland 1952; Munroe 1956; Küchler 1964). This system, in turn, has been replaced in Canada by an Ecozone classifi cation according to substrate type (Smith et al. 1996; Shorthouse 2010; Shorthouse and Larson 2010), which is designed to give a long-term perspective to ecology. The Prairies Ecozone can be subdivided into ecoprovinces, ecoregions, and ecodistricts. This classifi cation system has been adapted and further extended to embrace endemic organisms (Ricketts et al. 1999) as a criterion for delimiting ecoregions with ancient roots. Unfortunately, this ecoregional classifi cation of the Prairies Ecozone does not consider the important contribution made by the numerous endemic leafhoppers to the biodiversity found there (Hamilton 2005a). Leafhoppers (Hemiptera: Cicadellidae) are many and varied, often being easy to collect. There are more than 20,000 species of leafhoppers described worldwide (Oman et al. 1990), of which some 1,200 occur in Canada (Maw et al. 2000). Grassland leafhoppers (Figs. 1–4) are so common that, at any one spot, they usually comprise about 50% of the contents of a sweep net. Their numbers can be so great that 100 sweeps are considered to be an adequate representation of their fauna for a particular site: two historical records from 1920 of Xerophloea in Nebraska reported swarming in such great numbers that their dead bodies gathered under street lights in drifts up to 5 cm thick (Lawson 1931). The fi rst regional list of a grassland leafhopper fauna in Canada, from Alberta, was issued 60 years ago (Strickland 1953). It contained 124 dubiously identifi ed species, of which perhaps as many as 40 valid species were from grasslands and represented only about 9% of the leafhoppers now known from the province (Maw et al. 2000). Some very common species that are prominent among those 40 species are not limited to the Prairies, for example, Draeculacephala feeding on sedges in sloughs as well as in northern fens; the widespread northern leafhoppers Cuerna septentrionalis (Walker), Diplocolenus confi guratus (Uhler), and D. evansi (Ashmead); the generalist Driotura gammaroides (Van Duzee); and the southern migrant or “tramp” species Endria inimica (Say) and Exitianus exitiosus (Uhler). Two common species that are absent from the list came as invasives to Alberta only in recent years: Athysanus argentarius Metcalf and Doratura stylata Canadian Grasslands and Their Endemic Leafhoppers 313 (Hemiptera: Auchenorrhyncha: Cicadellidae) 1 2 3 4 Figs. 1–4. Leafhoppers associated with Canadian grasslands, west to east: 1, Dikrella nevadensis on Purshia tridentata on Cordilleran grasslands; 2, Flexamia graminea on true tallgrass prairie in western Manitoba (this site is dominated by needlegrass and little bluestem on glacial till with big bluestem on the more mesic periphery); 3, Afl exia rubranura on oak savanna of Manitoba; 4, Limotettix (Dryola) elegans on low spikerush in “tallgrass” oak openings at Windsor, Ontario (dominated by big bluestem and cordgrass in mesic areas, little bluestem on sandy uplands). Photograph of Purshia, taken in Washington by Robert Carr, and of insects by J. Elsaesser (1) and Chris Dietrich (2–4). (Boheman). The failure to discover more leafhoppers from an area that was at the time largely resilient native grassland may be attributed in part to the diffi culties of accessing sites. Road maps from the early 1950s show only four paved roads of any extent anywhere on the Canadian Prairies, supplemented by a broken grid work of gravel roads spaced an average of 60 km apart. The majority of early collections were taken along these primitive roads (Fig. 5). Collecting leafhoppers has become much easier since those days (Fig. 6) with the development of extensive paved road networks, especially on the Prairies. Unfortunately, these same roads have also divided native areas, opening up more land to crops and destroying remaining native grasslands. For example, many of the extensive tracts of grassland recorded by Coupland (1973) have vanished or been reduced to small holdings of varying quality, making assessment of the remainder a daunting task.