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271

Chapter 9 of the of British Columbia and the Yukon Territory

James Miskelly Research Associate, Royal BC Museum 675 Belleville St., Victoria, B.C., V8W 9W2

Abstract. Of all the habitats available in British Columbia and the Yukon Territory, grasslands support by far the greatest diversity, 87 , of Orthoptera. Although most of these species have broad distributions in western North America, 23 are not found in other provinces or territories and one is endemic to Yukon. The rarest and most restricted species are those that have their Canadian ranges limited to the arid shrub-steppe in southern British Columbia. Although most Orthoptera in British Columbia and Yukon grasslands are phytophagous, few cause economic damage to cultivated crops. The Orthoptera of British Columbia and Yukon are relatively well studied thanks to the work of a series of researchers over the last century. However, basic inventory and ecological study is needed throughout the region. Since 2005, Orthoptera in British Columbia and Yukon have received renewed attention in fi eld collections and basic research.

Résumé. De tous les habitats disponibles en Colombie-Britannique et au Yukon, ceux des prairies présentent de loin la plus grande diversité d’orthoptères, soit 87 espèces. La plupart de ces espèces sont largement répandues dans l’ouest de l’Amérique du Nord, mais 23 d’entre elles sont inconnues dans les autres provinces et territoires, et une est endémique du Yukon. Les espèces les plus rares et les plus restreintes sont celles dont l’aire de répartition canadienne se limite à la steppe arbustive aride du sud de la Colombie-Britannique. Bien que la plupart des orthoptères des prairies de la Colombie-Britannique et du Yukon soient phytophages, rares sont ceux qui causent des dommages économiques aux cultures. Ces orthoptères sont relativement bien connus grâce au travail accompli par plusieurs chercheurs au cours du dernier siècle. Cependant, il reste nécessaire de procéder à des inventaires de base et à des études écologiques dans l’ensemble de cette région. Depuis 2005, les orthoptères de la Colombie-Britannique et du Yukon retiennent à nouveau l’attention des chercheurs qui effectuent des collectes sur le terrain et des travaux de recherche fondamentale.

Introduction

The Orthoptera of British Columbia and Yukon comprise 108 species, two of which are represented by two subspecies (Vickery 1997; Miskelly 2012). These occur in virtually every habitat available in the region, including arctic tundra, mountain tops, wetlands, and forests of all descriptions. However, by far the greatest number of species can be found in grasslands. Fully 80% (87 species) of the regional orthopteran fauna occurs in grasslands. This includes generalist species that are found in a variety of open habitats, species more common in alpine areas or wetlands, and species that can be considered obligates. The fi rst major contributions to the study of Orthoptera in British Columbia and Yukon came from E.R. Buckell. Buckell worked for the Dominion Entomological Branch, Canada Department of Agriculture, in Vernon and Kamloops from 1920 to 1949. During this time,

Miskelly, J. 2014. Orthoptera of the Grasslands of British Columbia and the Yukon Territory. In of Canadian Grasslands (Volume 3): Biodiversity and Systematics Part 1. Edited by H. A. Cárcamo and D. J. Giberson. Biological Survey of Canada. pp. 271-281. © 2014 Biological Survey of Canada. ISBN 978-0-9689321-6-2 doi: http://dx.doi.org/10.3752/9780968932162.ch9 272 J. Miskelly

he collected Orthoptera extensively throughout British Columbia and published many new species records, distribution notes, and other observations (e.g., Buckell 1925, 1929, 1930, 1945). Buckell’s work provided the fi rst basis for checklists of the Orthoptera of British Columbia and established a foundation for future researchers. A number of species collected by Buckell in the 1930s were not collected again in British Columbia until recent years. A few species are yet to be found again. Following the work of Buckell, the next major contributions to the study of the Orthoptera of British Columbia and Yukon were provided by V.R. Vickery. Vickery was an extension entomologist for the province of Nova Scotia from 1949 to 1961. From 1961 to 1986, he was a professor at McGill University and curator of the Lyman Entomological Museum, which contains the largest collection of Orthoptera in Canada. During his time in Montreal, Vickery collected widely and published on the Orthoptera across Canada (e.g., Vickery 1961, 1967; Vickery and Kevan 1967; Vickery and Nagy 1973). Although he conducted little fi eld work in British Columbia or Yukon, Vickery made important contributions there, exemplifi ed by the descriptions of two new species of Orthoptera that occur in Yukon grasslands, Brunneria yukonensis and Xanthippus brooksi (Vickery 1967, 1969). Most important, Vickery (together with D.K.McE. Kevan) authored an Agriculture Canada guide to the orthopteroid insects of Canada (Vickery and Kevan 1985). This monograph greatly facilitated the study of Orthoptera in Canada, for the fi rst time providing a single source for the identifi cation of all taxa and establishing a fi rst checklist of the Orthoptera of Canada and of each of the provinces and territories. The next research of relevance to the Orthoptera of British Columbian grasslands began when G.G.E. Scudder initiated a research project surveying insects in the south Okanagan in the 1990s. More than 6,000 Orthoptera specimens were collected in pitfall traps over the course of 10 years (Royal BC Museum data), representing the most intensive Orthoptera sampling ever conducted in British Columbia. This study provided the fi rst precise location information for many rare species that were previously known only from vague museum labels. Since 2005, Orthoptera have increasingly been targeted in the fi eld collections of the Royal BC Museum and have been collected in eight of British Columbia’s nine terrestrial ecoprovinces (as listed in Demarchi 2011) and 14 of 16 biogeoclimatic zones (as listed in British Columbia Forest Service 2012). These collections and museum research by staff and volunteers at the Royal BC Museum and the BC Ministry of Environment have added several new species to the known fauna of British Columbia, corrected misidentifi cations of earlier specimens, accrued a great deal of new distributional data, and produced an updated checklist for British Columbia (Miskelly 2012). At the same time, workers in Yukon, led by NatureServe Yukon, have collected throughout the territory and produced a great deal of new distributional and ecological data. The Orthoptera of the grasslands of British Columbia and Yukon are presently being inventoried and researched more actively than at any time since the work of Buckell. The conservation status of Orthoptera has recently been ranked nationally and within British Columbia and Yukon (NatureServe 2012). Knowledge of the group is more complete than ever before, but signifi cant gaps remain (see Research Priorities below).

Orthoptera of the Grasslands of British Columbia and Yukon The 87 Orthoptera species known from the grasslands of British Columbia and Yukon (Table 1) are spread among seven families and 14 subfamilies, though only two of these families and four subfamilies occur in Yukon. Most of the species diversity is found in the , with 57 species in British Columbia and 11 species in Yukon. Orthoptera of the Grasslands of British Columbia and the Yukon Territory 273

Feeding patterns in Orthoptera range from primarily phytophagous to including variable amounts of dead or live material in the diet. The majority of grassland Orthoptera species in British Columbia and Yukon are primarily phytophagous. A few species of generalist herbivores have adapted to agricultural habitats, where they are considered pests. The most notorious of these are bivittatus, M. sanguinipes, and Camnula pellucida (Buckell 1945; Vickery and Kevan 1985). A common belief is that all are pests, and most species occurring in Canada are described as occasional pests even in the absence of supporting evidence (Vickery and Kevan 1985). However, in this region at least, the diversity of Orthoptera in cultivated areas is very low and most species are rarely encountered outside of natural habitats. Grasshoppers generally do not cause economic damage to crops or range in British Columbia except when drought conditions have already stressed plants or reduced range productivity (British Columbia Ministry of Agriculture 2009). Many of the phytophagous Orthoptera feed preferentially on a select group of plants, such as graminoids, forbs, or shrubs (Vickery and Kevan 1985). Some species have narrow host-plant specifi city. For example, Melanoplus cinereus feeds almost entirely on the shrub Artemisia tridentata Nuttall 1841, and viridis feeds mainly on a small suite of species in the family Asteraceae (Sheldon and Rogers 1976). The most innocuous herbivores are the pygmy grasshoppers of the family Tetrigidae, which appear to feed mostly on algae that they scrape off soil particles (Vickery and Kevan 1985). Many species of Orthoptera are omnivorous to a certain degree. The camel crickets of the family are scavengers of both plant and animal foods (JM, pers. obs.). Most members of the families Prophalangopsidae, Stenopelmatidae, and capture live prey in addition to feeding on plant material (Vickery and Kevan 1985; JM, pers. obs.). None of the Orthoptera of the grasslands of British Columbia and Yukon are primarily predaceous. Recent collection data have also provided information on distribution patterns of British Columbia and Yukon Orthoptera. Forty-eight of the 87 species recorded in British Columbia and Yukon grasslands are found primarily or exclusively in these grasslands (Table 1) within this region. These species do not form a cohesive ecological group, but rather occur across different types of grasslands and in different parts of the region. The largest group within these primarily grassland species comprises those species associated with the most arid grasslands in British Columbia (Table 2). These shrub-dominated grasslands occur only in the hottest river valleys of the southern third of the province (Nicholson et al. 1991). Because of the uniqueness of this habitat, many of the species in these arid lands are highly restricted in both range and habitat. This group contains some of our rarest species, such as enigma and spurcata. Both species have been collected only a few times in British Columbia and only in the southern Okanagan Valley. Species that are not restricted to grasslands are often associated with more widespread habitats that overlap with grasslands (Table 3). Wetlands of various descriptions support a number of species of Orthoptera, with the species assemblage changing somewhat when these wetlands occur in grassland settings. For example, all three species of Tetrix (Tetrigidae) in British Columbia are associated with sparsely vegetated mud or sand around wetlands. None of these are restricted to grassland settings. Riparian thickets support several species that are likewise not restricted to grasslands, such as fultoni, O. rileyi, and Scudderia furcata. Wet depressions dominated by grasses or sedges provide habitat for several species that are largely restricted to grasslands in British Columbia, such 274 J. Miskelly

Table 1. Orthoptera of the grasslands of British Columbia and Yukon. Bolded species are found only in grasslands within this region.

Family Subfamily Species Stenopelmatidae Stenopelmatinae Stenopelmatus fuscus Haldeman 1852 Rhaphidophoridae Ceuthophilinae Ceuthophilus agassizii (Scudder 1861) Ceuthophilus alpinus Scudder 1894 Ceuthophilus vicinus Hubbell 1936 Pristoceuthophilus cercalis Caudell 1916 Pristoceuthophilus pacifi cus (Thomas 1872) Prophalangopsidae Cyphoderrinae Cyphoderris buckelli Hebard 1934 Tettigoniidae Anabrus longipes Caudell 1907 Anabrus simplex Haldeman 1852 Apote robusta Caudell 1907 Peranabrus scabricollis (Thomas 1872) Steiroxys undescribed Steiroxys cf. trilineata (Thomas 1870) fasciatus (DeGeer 1773) gladiator Bruner 1891 Phaneropterinae Scudderia furcata Bruner von Wattenwyl 1878 allardi (Alexander and Thomas 1959) (DeGeer 1773) pennsylvanicus Burmeister 1838 (Alexander and Bigelow 1960) Oecanthinae Oecanthus argentinus Saussure 1874 Oecanthus californicus Saussure 1874 Walker 1962 Oecanthus quadripunctatus Beutenmuller 1894 Oecanthus rileyi Baker 1905 Acrididae Asemoplus montanus (Bruner 1885) Bradynotes obesa (Thomas 1872) Buckellacris chilcotinae (Hebard 1922) (Thomas 1872) Melanoplus alpinus Scudder 1897 (Say 1825) (Fieber 1853) Scudder 1897 Melanoplus cinereus Scudder 1878 Scudder 1897 (Scudder 1875) (DeGeer 1773) Scudder 1878 Scudder 1878 Melanoplus kennicotti Scudder 1878 Melanoplus occidentalis (Thomas 1872) Scudder 1878 Orthoptera of the Grasslands of British Columbia and the Yukon Territory 275

Melanoplus rugglesi Gurney 1949 (Fabricius 1798) Melanoplus washingtonius (Bruner 1885) (Scudder 1876) Phaetaliotes nebrascensis (Thomas 1872) Oedipodinae conspersa Scudder 1875 (Thomas 1870) Camnula pellucida (Scudder1862) Chortophaga viridifasciata (DeGeer 1773) carlinianus (Thomas 1870) Rehn and Hebard 1906 Circotettix undulatus (Thomas 1872) Conozoa sulcifrons (Scudder 1876) Cratypedes lateritius (Saussure 1884) Cratypedes neglectus (Thomas1870) (Linnaeus 1758) Dissosteira spurcata Saussure 1884 Metator nevadensis (Bruner 1905) apiculata (Harris 1865) Spharagemon campestris (McNeill 1901) Spharagemon equale (Say 1825) Stethophyma gracile (Scudder 1862) Trachyrhachys kiowa (Thomas1872) Trimerotropis fontana Thomas 1876 Trimerotropis gracilis (Thomas 1872) Trimerotropis pallidipennis (Burmeister 1838) Trimerotropis verruculata suffusa Scudder 1876 Trimerotropis verruculata verruculata (Kirby 1837) Xanthippus brooksi Vickery 1967 Xanthippus corallipes (Haldeman 1852) clavatus (Thomas 1873) deorum (Scudder 1876) coloradus (Thomas 1873) ellioti (Thomas 1870) Brunneria brunnea (Thomas 1871) Brunneria yukonensis Vickery 1969 abdominalis (Thomas 1873) (Harris 1841) pelidna (Burmeister 1838) curtipennis (Harris 1835) brachyptera (Scudder 1862) delicatula (Scudder 1876) Tetrigidae Tetriginae Tetrix brunnerii (Bolivar 1887) Tetrix ornata (Say 1834) Tetrix subulata (Linnaeus 1758) 276 J. Miskelly

Table 2. Orthopteran species restricted to arid shrub-steppe within the British Columbia portion of their range.

Family Subfamily Species Acrididae Melanoplinae Hesperotettix viridis (Thomas) Melanoplus cinereus (Scudder) Melanoplus rugglesi Gurney Oedaleonotus enigma (Scudder) Phaetaliotes nebrascensis (Thomas) Oedipodinae Dissosteira spurcata Saussure Trimerotropis gracilis (Thomas) Tettigoniidae Tettigoniinae Apote robusta Caudell Stenopelmatidae Stenopelmatinae Stenopelmatus fuscus Haldeman

as and Allonemobius fasciatus, but also support species that occur in a variety of moist, grassy habitats, including two species of meadow katydid, Conocephalus fasciatus and . Mountain meadows support an orthopteran fauna that has a degree of overlap with grasslands. Species that occur from low-elevation grasslands all the way to alpine tundra include Asemoplus montanus, Buckellacris chilcotinae, Melanoplus bruneri, and M. washingtonius. Other species are absent from lower grasslands, but extend from the upper- elevation grasslands into subalpine or alpine regions. These include Melanoplus alpinus and Steiroxys cf. trilineata. A few species seem to be almost restricted to upper-elevation grasslands in British Columbia and are rarely found in either lower grasslands or higher mountain meadows. These include Brunneria brunnea (Fig. 1), , and Peranabrus scabricollis. Some species of Orthoptera that are found in grasslands are associated more usually with wooded habitats. The camel crickets of the genera Ceuthophilus and Pristoceuthophilus are often found in heavily forested settings, but several species have also been found in grasslands, such as Ceuthophilus vicinus, Pristoceuthophilus cercalis, and P. pacifi cus. Open woodlands and forest edges support a diverse assemblage of orthopterans that overlaps with the grassland fauna. A small suite of species is found mainly in dry, open woodlands associated with grasslands. These species are more common in these semi-treed sites than they are in either open grassland or closed forest. Examples include Cyphoderris buckelli, , and C. conspersa.

Biogeographical Aspects The biogeographical affi nities of most orthopterans of British Columbia and Yukon were listed by Scudder and Vickery (1998). The majority of species are described as Cordilleran, Nearctic (excluding Beringia), or Western Nearctic (excluding Beringia). Thus, most of the fauna is made up of species that are relatively widespread, at least in western North America. Most Orthoptera species found in British Columbia and Yukon reach their northern limit in this region. A few range little, or not at all, to the south, whereas many have much larger ranges with the borders of the . Of the Orthoptera found in the grasslands of British Columbia and Yukon, 23 species are not found in other provinces or territories. Of Orthoptera of the Grasslands of British Columbia and the Yukon Territory 277

Table 3. General habitat associations of species found partially in grasslands of British Columbia and Yukon.

Species Wetland/ Woodland/ Montane/ Disturbed/ Arctic Riparian Forest Alpine Generalist Tundra

Ceuthophilus agassizii X Ceuthophilus alpinus X Ceuthophilus vicinus X Pristoceuthophilus cercalis X Pristoceuthophilus pacifi cus X Cyphoderris buckelli X Steiroxys cf. trilineata X Conocephalus fasciatus X Orchelimum gladiator X Scudderia furcata X X X Gryllus veletis X Oecanthus fultoni X Oecanthus rileyi X Asemoplus montanus X Bradynotes obesa X Buckellacris chilcotinae X Melanoplus alpinus X Melanoplus bivittatus X Melanoplus borealis X Melanoplus bruneri X Melanoplus femurrubrum X Melanoplus sanguinipes X Melanoplus washingtonius X Camnula pellucida X Dissosteira carolina X Stethophyma gracile X Trimerotropis pallidipennis X Trimerotropis verruculata X suffusa Trimerotropis verruculata X verruculata Xanthippus brooksi X X Chloealtis abdominalis X Chloealtis conspersa X Pseudochorthippus curtipennis X Tetrix brunnerii X Tetrix ornata X Tetrix subulata X 278 J. Miskelly

these, 22 have their Canadian distribution limited to British Columbia, and one (Brunneria yukonensis) is endemic to Yukon. None of those species that have their Canadian range limited to British Columbia are endemic. Many are typical of the arid lands of the Great Basin, such as Melanoplus rugglesi, Dissosteira spurcata, and Metator nevadensis. A large degree of overlap occurs between the orthopteran faunas of British Columbia and Yukon grasslands and those of the Canadian prairie grasslands. Of the 87 species recorded from British Columbia and Yukon grasslands, at least 50 also occur in the prairies (Vickery and Kevan 1985). The Orthoptera of Canadian prairie grasslands are slightly more diverse, with approximately 100 species recorded (Vickery and Kevan 1985). The similarity in the species richness of the two regions is remarkable, given the comparatively small area that grasslands occupy in British Columbia and Yukon. Several species that occur in British Columbia are more typical of prairie grasslands and range into British Columbia only on the periphery of a range that is more extensive east of the Rocky Mountains. These species may occur in British Columbia in the East Kootenays or the Peace River region (Vickery and Kevan 1985; Royal British Columbia Museum data). Examples include Anabrus simplex, Melanoplus packardii, and Chortophaga viridifasciata. Two other species have a similar distribution pattern, but also extend west over the northern edge of the Rocky Mountains into the southern Yukon and northwestern British Columbia. and Melanoplus kennicotti (Fig. 2) follow this pattern. Both have restricted ranges in British Columbia grasslands, but are characteristic members of the fauna of Yukon grasslands (Royal BC Museum data). Two species found in Yukon grasslands have Beringian distributions and are not known to occur south of the territory (Vickery 1997). Brunneria yukonensis has been recorded only in the grasslands of the southern Yukon. Xanthippus brooksi has been recorded through much of Yukon and western Northwest Territories in a variety of habitats, including grassland, dry tundra, and sparsely vegetated south-facing slopes (Vickery 1997; Royal BC Museum data).

Research Priorities Although the legacy of previous researchers has provided an important base from which to build knowledge on the Orthoptera of British Columbia and Yukon, much remains to be learned about the ecology, habits, and distribution of all species. Most historic specimens are labelled with only vague location information, and these should be refi ned through ongoing fi eldwork. Basic inventory data are needed throughout the region. In particular, additional species are expected to occur in northwestern British Columbia, the Kootenays, and the Peace Region (Miskelly 2012). Even in the most heavily sampled areas (i.e., British Columbia’s Lower Mainland and Okanagan Valley), signifi cant new records are obtained regularly and several historic records have not been replicated. Orthoptera are known for extreme fl uctuations in abundance from year to year, with slight changes in weather patterns causing great changes in the relative abundance of different species (Mulkern 1980). Regular monitoring in well-studied areas and repeated sampling in poorly studied areas are important for the documentation of these fl uctuations and detection of overlooked species. Several species that occur in grasslands within British Columbia and Yukon have been identifi ed as potentially rare (Scudder 1994) or ranked as “at risk” (NatureServe 2012). It is important that more fi eld work be devoted to these species to describe their habitat requirements, assess whether they are truly rare, and determine whether they are in need of conservation measures. The status of many species is likely to change as more complete information is obtained about their distributions and habitat associations. Orthoptera of the Grasslands of British Columbia and the Yukon Territory 279

Fig. 1. Brunneria brunnea (Thomas) is found mainly in upper-elevation grasslands in southern British Columbia. A congener is endemic to grasslands of the southern Yukon. Photo by the author.

Fig. 2. Melanoplus kennicotti Scudder is found in the grasslands of northeastern British Columbia and southwestern Yukon. Photo by the author. 280 J. Miskelly

Although the Orthoptera of Canada are relatively well resolved taxonomically, a few areas of uncertainty exist where revisions are needed, particularly in the family Tettigoniidae. The Steiroxys, in particular, has baffl ed taxonomists for more than a century (Caudell 1907; Rentz and Birchim 1968). At present, three species are believed to occur in British Columbia, but none can be identifi ed to species with certainty and one is likely undescribed (Miskelly 2012). Other taxonomic problems in the family relate to possible synonymies within the genera Apote and Anabrus.

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