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191 Chapter 6 Springtails (Hexapoda: Collembola) of the Prairie Grasslands of Canada Zoë Lindo Department of Biology, University of Western Ontario, London, Ontario, Canada, N6A 5B7 E-Mail address: [email protected] Telephone: 519 661 2111 ext. 82284 Abstract. Collembola, also known as “springtails”, are common components of many soil systems, including grasslands. The published studies from Canadian grasslands that explore soil microarthropod communities suggest that both the diversity and densities of Collembola are low compared with those in other habitats (e.g., forest systems) and other fauna groups (e.g., Acari). Very few studies, however, document and describe Canadian grassland Collembola at the species level, and all major grassland areas of Canada are undersampled for Collembola. For grasslands outside the Prairies Ecozone, no record of sampling was found for Collembola in southern Ontario tallgrass systems, nor has there been a concerted effort of collection in bunchgrass/sagebrush habitats of British Columbia. Despite the paucity of direct information from Canada, studies from the mid- central United States and elsewhere provide the basis for a potential species-level checklist. In particular, species from the following families are expected to dominate species diversity in Canadian grasslands: Isotomidae, Entomobryidae, Hypogastruridae, and Sminthuridae. The dearth of species-level knowledge is due to the lack of taxonomic expertise for the Collembola in Canada. Résumé. Les collemboles sont présents dans de nombreux systèmes de sols, y compris ceux des prairies. Les études publiées portant sur les prairies canadiennes qui se penchent sur les communautés de microarthropodes terricoles donnent à conclure que la densité et la diversité des collemboles présents dans ces communautés sont faibles comparativement à celles observées dans d’autres habitats (ex : sols forestiers) ou à celles caractérisant d’autres groupes fauniques (ex : acariens). Cependant, rares sont les études qui examinent et décrivent les espèces de collemboles des prairies canadiennes, et aucune des principales zones de prairies du Canada n’a fait l’objet d’échantillonnages exhaustifs des collemboles. À l’extérieur de l’écozone des prairies, il semble que les collemboles n’aient fait l’objet d’aucun travail d’échantillonnage dans les écosystèmes à herbes hautes du sud de l’Ontario, ni d’aucun effort concerté de collecte dans les peuplements de graminées cespiteuses ou d’armoises de la Colombie-Britannique. Malgré la rareté des informations directes provenant du Canada, les études réalisées dans le centre des États-Unis et ailleurs fournissent le point de départ d’une possible liste des espèces. En particulier, on s’attend à ce que les espèces appartenant aux familles suivantes occupent une place dominante dans les prairies canadiennes : isotomidés, entomobryidés, hypogastruridés et sminthuridés. Le manque de connaissances sur les espèces de collemboles découle de l’absence d’experts de ce groupe au Canada. Introduction Collembola, also known as “springtails,” are small (0.5–3 mm), wingless, hexapodous arthropods. The common name springtail is derived from a ventral appendage of many species called the furcula, which is used in predator defence (Hopkin 1997). The diagnostic feature of collembolans, however, is the presence of a tubular collophore used primarily in maintaining moisture balance (Hopkin 1997). Collembola, previously considered within Lindo, Z. 2014. Springtails (Hexapoda: Collembola) of the Prairie Grasslands of Canada. 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. 191-198. © 2014 Biological Survey of Canada. ISBN 978-0-9689321-6-2 doi: http://dx.doi.org/10.3752/9780968932162.ch6 Species checklist available at http://dx.doi.org/10.5886/xd2b2ssq 192 Z. Lindo the class Insecta (Apterygota: Collembola), are now considered as a sister clade to the Insecta within the subphylum Hexapoda (Entognatha: Collembola). Controversy on this placement and the monophyly of Hexapoda (Nardi et al. 2003; Meusemann et al. 2010) continued for many years, when inconsistencies arose depending on whether analyses were based on morphology, mitochondrial DNA sequences, or nuclear genes. However, a subsequent analysis has fi rmly placed Collembola within the Hexapoda and the hexapods as a monophyletic group (Timmermans et al. 2008). As such, Collembola are considered basal hexapods, yet the status of Entognatha as a distinct clade remains uncertain (Meusemann et al. 2010). As the Collembola occur in two distinct body forms, elongate and globular, morphological taxonomy within the group led to the previous distinction of two orders, Arthropleona and Symphypleona (Christiansen and Bellinger 1980), whereas modern taxonomy now recognizes four orders: Poduromorpha, Entomobryomorpha, Symphypleona, and Neelipleona (Deharveng 2004). Collembola are common components of many soil systems (Petersen and Luxton 1982; Behan-Pelletier 2003). Feeding habits, based on both gut enzyme analysis (Berg et al. 2004) and stable isotope ratios (Chahartaghi et al. 2005), suggest that most Collembola species fall within a continuum from phycophages/herbivores to primary and secondary decomposers. Many species display “r-selection” characteristic life history traits (i.e., fast, semelparous reproduction and short lifespan), which allows them to respond quickly to environmental change. The primary abiotic factors affecting collembolan density and diversity are moisture and soil acidity (Hopkin 1997). Collembola play an important, albeit mostly indirect, role in decomposition through their interactions with the microbial community (Visser 1985). Despite recent discussion on placement and classifi cations within the Collembola (Xiong et al. 2008), species-level taxonomy for the group is relatively well-known (Behan- Pelletier and Bisset 1992). There are approximately 7,850 known species worldwide (Janssens 2007), with approximately 700 species described from North America (about 80% of estimated diversity); the known number of species in Canada is 412, whereas the estimated number of Canadian Collembola is roughly 520 species (Behan-Pelletier 2003). Collembola of Canadian Grassland Ecosystems Few Canadian studies document and describe grassland Collembola communities, and only two studies describe the biodiversity of grassland Collembola and document species-level data. Aitchison (1979) found 16 species of winter-active Collembola in a tallgrass Manitoba prairie, where Isotoma manitobae was the most abundant species. This study illustrates how poorly Collembola are known in Canada because the dominant species was previously undescribed. Berg and Pawluk (1984), found 10 Collembola species across seven prairie and agricultural vegetative covers in central Alberta, eight of which were found in native fescue grassland sites. In general, grassland areas have only low-to-moderate diversity (Petersen and Luxton 1982); thus, richness values presented in the aforementioned studies (16 and 10) may be typical of Canadian grassland Collembola diversity. However, species richness reported by Aitchison (1979) is likely an underrepresentation of total Collembola diversity, as sampling was performed in winter when abundances were low, and the collection method used was pitfall traps, which could preclude endogeic (burrowing) species. Collembola densities are better known than richness estimates, both in grasslands and in general. Densities of Collembola typically range from 100 to 670,000 individuals/ Springtails (Hexapoda: Collembola) of the Prairie Grasslands of Canada 193 m2 (Petersen and Luxton 1982) worldwide, with the highest densities associated with moist, organic forest soils. Grasslands typically have low-to-moderate densities, which are infl uenced by factors such as soil moisture and soil organic matter (Petersen and Luxton 1982; Ferguson 2001). For instance, in grassland sites of northeastern Illinois and southwestern Michigan, Brand and Dunn (1998) recorded densities of 104–500 individuals/ m2 in native grasslands, and 27–361 individuals/m2 in restored grasslands. In a study of the relationships among roots, mycorrhizal fungi, and Collembola in a native grassland near Regina, Saskatchewan (50°28´N; 104°22´W), Steinaker and Wilson (2008) used an in situ below-ground “rhizotron” camera to look at collembolan density. They found that densities were greater in forest than in grassland but varied little among horizons within the grassland sites (~250 individuals/m2). This contradicts the results of most vertical distribution studies, which show that densities of Collembola tend to be greatest in the top few centimetres of soil in grassland systems (Berg and Pawluk 1984), in association with greater soil moisture and organic matter at the soil surface. Other factors driving Collembola density in grassland systems include plant species richness (Salamon et al. 2004; Sabais et al. 2011) and disturbance regimes (e.g., grazing; Dombos 2001; Clapperton et al. 2002). Collembola densities also change seasonally, whereby abundance peaks in summer and secondarily in the fall, but remains low in the spring (Berg and Pawluk 1984; Steinaker and Wilson 2008) and winter (Aitchison 1979). Notable Canadian Grassland Collembola Studies • Aitchison (1979) found 16 species of Collembola in aspen parkland and tallgrass prairie near Fort Whyte, Manitoba (49°49´N; 97°13´W). • Berg and Pawluk (1984) found 10