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Project Update
- Arthropod Fauna of Soils 2001
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General information and editorial notes News and Notes Field trip to the Onefour grasslands Summary of the Scientific Committee meeting Second spider newsletter published Members of the Scientific Committee
Project Update: Arthropod Fauna of Soils 2001 Requests for Material or Information
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Biodiversity Section, ECORC, Agriculture and Agri-Food Canada, K.W. Neatby Bldg., 960 Carling Ave., Ottawa, Ontario, Canada K1A 0C6
“We know more about the movement of
celestial bodies than about the soil underfoot.”
Soils, wherever they are found, are recognized as habitats for much of the biosphere’s undescribed diversity; they make up an environment where, even in North America, only about 50% of species are described as adults (Behan-Pelletier and Bissett 1992). The Biological Survey has long recognized the importance of soil fauna as essential for maintaining the fertility of Canada’s soils. It regularly comments on relevant developments on soil arthropods, e.g., Behan-Pelletier (1995), although it does not have an active project because of the limited taxonomic resources in Canada. Also, the Survey strongly encourages inclusion of soil arthropods in its Projects, such as the Grassland Project and Insects of the Yukon (Danks and Downes 1997). The Survey first outlined the status and research needs for Canadian soil arthropods in a brief (Marshall et al. 1982), a document as relevant today as then. Subsequently, it helped sponsor a conference on faunal influences on soil structure (Spence 1986), and ensured contributions on soil arthropods in publications associated with the Survey, e.g., Ball and Danks (1993) and Finnamore and Marshall (1994). The following update is not intended to be exhaustive. Rather, it attempts to highlight some recent developments.
Gozmanyina majesta (Marshall and Reeves), an inhabitant of soil and litter in mixed deciduous-coniferous forests of northeastern North America. Since the publication of “Life in the Soil” (Freckman 1994) there has been considerable focus on soil biodiversity in general in North America, and more specifically on documenting the role of soil biodiversity in ecosystem function. Various workshops from the SCOPE Committee on Soils and Sediments have deepened awareness of fundamental similarities in biodiversity between terrestrial soils and the sediments of freshwater and marine environments (Brussard et al. 1997, Wall Freckman et al. 1997, Wall 1999). SCOPE workshops have addressed questions such as the impact of soil biodiversity on plant productivity (SCOPE 2001), have recognized that soils are critical transition zones (Bardgett et al. 2001), and have addressed possible effects of global change on soil biota (Young et al. 1998). Concurrent with SCOPE initiatives on soil biodiversity, the EMAN program of Environment Canada (Finnamore et al. 1998) and various Canadian and international biodiversity assessments, such as the SAGE Grassland Project (Finnamore 1996), and the ALAS Project in Costa Rica ( have helped to formalize sampling methodologies for this biodiversity. In addition, this year FAO has developed a soil biodiversity internet portal (http://www.fao.org/landandwater/agll/soilbiod/biodport.htm) with the overall aim of promoting “a more ecological approach in agricultural systems and the integrated management of land resources with a view to enhancing agricultural productivity and agro-ecosystem sustainability”. In parallel with this surge in interest in soil biodiversity in North America, forest canopies, with their suspended soil habitats, have gained recognition as one of the last biotic frontiers (Winchester 1997; International Canopy Network (ICAN: http://192.211.16.13/individuals/nadkarnn/main/info.htm)). Canopies include a range of soil and litter microhabitats almost homologous to those on the forest floor. These microhabitats, collectively called ‘canopy organic matter’ host diverse assemblages of soil arthropods that can be distinct from and can rival in richness those on the forest floor (Behan-Pelletier and Walter 2000; Winchester 1997). But have these and other initiatives increased taxonomic information AND awareness about soil biodiversity, especially for hyperdiverse taxa such as the arthropods? I think the answer is a qualified Yes. Granted, the “important deficiencies related to soil arthropods” outlined in Marshall et al. (1982) and reiterated by Behan-Pelletier (1993a) have not been addressed, but soil arthropod biodiversity is no longer ignored in meaningful ecosystem analyses. Central to this change is that the taxonomic inadequacies are more widely appreciated, as a result of Kosztarab and Schaefer’s (1990) analysis of status and needs for North American insect and arachnid systematics, subsequent analyses of specific groups, e.g., Bennett (1999), and the Soil Biology Guide (Dindal 1990) which includes keys to most of the major microbial and invertebrate inhabitants of North American soils. We know that Acari, Diptera and Coleoptera comprise over 75% of the arthropod species richness in most North American soils, but that these groups have only about 15%, 68% and 87% of adults identifiable to species, respectively (Behan-Pelletier and Bissett 1992). Furthermore, the immature stages of only 5-10% of species are described, even though these stages are typically the most metabolically active and long-lived in the soil. The few taxonomic experts available are trying to address these inadequacies by developing standard keys, and by using the internet to make data more available. An excellent example is the Checklist of the Collembola (Janssens 2001), the Collembola Key2000 Project (Janssens 2001) and the list of Collembola literature). But the taxonomic impediment remains, and ultimately, taxonomic deficiencies can only be solved by taxonomic expertise. In the meantime these deficiencies are being addressed in a number of ways outlined below:
One or all of these approaches have been used in biodiversity and faunistic studies on soil arthropods in Canada, and have contributed qualitatively and quantitatively to our knowledge of this fauna. As an example, the diversity of studies on Canadian soil arthropods published since 1993 is illustrated for 2 groups, the Acari and Collembola, in Table 1.
Table 1. Recent knowledge on systematics, distribution and ecology of Acari and Collembola in Canada (based on additions to the literature since Behan-Pelletier 1993a).
References Addison, J.A. 1993. Persistence and nontarget effects of Bacillus thuringiensis in soil: a review. Canadian Journal of Forest Research 23: 2329-2342. Addison, J.A. 1996a. Safety of tebufenozide, a new molt-inducing insecticide, for forest soil invertebrates. Ecotoxicol. Environ. Safety 33: 55-61. Addison, J.A. 1996b. Harvesting and site preparation impacts on soil microarthropods. pp.186-189 In: C.R. Smith and G.W Crook (compilers), Advancing Boreal Mixedwood Management in Ontario- Proceedings. 17-18 October 1995, Sault Ste. Marie, Ontario. NRCan, CFS- Sault Ste. Marie, Ontario. Addison, J.A. and S.B. Holmes. 1995. Effect of two commerical formulations of Bacillus thuringiensis subsp. kurstaki (Dipel 8L and Dipel 8AF) on the collembolan species Folsomia candida in a soil microcosm study. Bull. Environ. Contam. Toxicol. 55: 771-778. Addison, J.A., V.G. Marshall, and J.A. Trofymow. 1998. Soil microarthropod abundance and species richness in successional Douglas-fir forests. Northwest Science 72 (Special Issue No. 2): 96-97. Ball, G. and H.V. Danks. 1993. Systematics and Entomology: diversity, distribution, adaptation, and application. Memoirs of the Entomological Society of Canada 165. 272 pp. Balogh, J. and P. Balogh. 1992. The oribatid mites genera of the world. Hungarian Natural History Museum, Budapest. Vol. I: 263 pp. Vol. II: 375 pp. Bardgett, R.D., J.M. Anderson, V.M. Behan-Pelletier, L. Brussard, D.C. Coleman, C. Ettema, A. Moldenke, and D.H. Wall. 2001. The influence of soil biodiversity on hydrological pathways and the transfer of materials between terrestrial and aquatic ecosystems. Ecosystems in press. Battigelli, J.P., S.M. Berch, and V.G. Marshall 1994. Soil fauna communities in two distinct but adjacent forest types on northern Vancouver Island, British Columbia. Canadian Journal of Forest Research 24: 1557-1566. Behan-Pelletier, V.M. 1993a. Diversity of soil arthropods in Canada: systematic and ecological problems. Memoirs of the Entomological Society of Canada 165. 11-50. Behan-Pelletier, V.M. 1993b. Eremaeidae (Acari:Oribatida) of North America. Memoirs of the Entomological Society of Canada 168. 193 pp. Behan-Pelletier, V.M. 1994. Mycobates (Acari: Oribatida: Mycobatidae) of North America. Canadian Entomologist 126: 1301-1361. Behan-Pelletier, V.M. 1995. Project update: arthropod fauna of soils. Newsletter of the Biological Survey of Canada (Terrestrial Arthropods) 14: 42-43. Behan-Pelletier,V.M. 1997. Oribatid mites (Acari: Oribatida) of the Yukon. pp. 115-149 in H.V. Danks and J.A. Downes (Eds.), Insects of the Yukon. Biological Survey of Canada Monograph Series No. 2. Behan-Pelletier, V.M. 2000. Ceratozetidae (Acari: Oribatida) of arboreal habitats. Canadian Entomologist 132: 153-182. Behan-Pelletier, V.M. and B. Bissett. 1992. Biodiversity of nearctic soil arthropods. Canadian Biodiversity 2:5-14. Behan-Pelletier, V.M. and B. Bissett. 1994. Oribatida of Canadian peatlands. Memoirs of the Entomological Society of Canada 169: 73-88. Behan-Pelletier, V.M., M. Clayton and L. Humble. 2001. Parapirnodus (Acari: Oribatida: Scheloribatidae) of canopy habitats in western Canada. Acarologia in press Behan-Pelletier, V. and D.E. Walter. 2000 Biodiversity of oribatid mites (Acari: Oribatida) in tree-canopies and litter. pp. 187-202 in D.C. Coleman and P. Hendrix (Eds.), Invertebrates as Webmasters. CABI Publication. Behan-Pelletier, V.M. and N.N. Winchester. 1998 Arboreal oribatid mites diversity: colonizing the canopy. Applied Soil Ecology 9:45-51. Bennett, R. 1999. Canadian Spider Diversity and Systematics. Newsletter of the Biological Survey of Canada (Terrestrial Arthropods) 18: 16-27. (http://www.biology.ualberta.ca/esc.hp/bsc/news18_1/spider.htm) Brussard, L., V.M. Behan-Pelletier, D.E. Bignell, V.K. Brown, W. Didden, P. Folgarait, C. Fragoso, D. Wall Freckman, V.V.S.R. Gupta, T. Hattori, D.L. Hawksworth, C. Klopatek, P. Lavelle, D.W. Malloch, J. Rusek, B. Soderstrom, J.M. Tiedje, and R.A. Virginia. 1997. Biodiversity and ecosystem functioning in soil. Ambio 26: 563-570. Cárcamo, H.A., T.A. Abe, C.E. Prescott, F.B. Holl, and C.P. Chanway. 2000. Influence of millipedes on litter decomposition, N mineralization, and microbial communities in a coastal forest in British Columbia, Canada. Canadian Journal of Forest Research 30: 817-826. Chagnon, M., C. Hébert, and D. Paré. 2000a. Community structures of Collembola in sugar maple forests: relations to humus type and seasonal trends. Pedobiologia 44: 148-174. Chagnon, M., D. Paré, and C. Hébert. 2000b. Relationships between soil chemistry, microbial biomass and the collembolan fauna of southern Quebec sugar maple stands. EcoScience 7: 307-316. Chagnon, M., D. Paré, C. Hébert, and C. Camiré. 2001. Effects of experimental liming on collembolan communities and soil microbial biomass in a southern Quebec sugar maple (Acer saccharum Marsh.) stand. Applied Soil Ecology 17: 81-90. Colloff, M.J. 1993. A taxonomic revision of the oribatid mite genus Camisia (Acari: Oribatida). Journal of Natural History 27: 1325-1408. Colwell R. 1997. Biota: The Biodiversity Database Manager. Sinauer Associates. http://viceroy.eeb.uconn.edu/biota. Cortet, J. and N. Poinsot-Balaguer. 2000. Impact de produits phytopharmaceutiques sur les microarthropodes du sol en culture de maïs irrigué : approche fonctionelle par la méthode des sacs de litière. Canadian Journal of Soil Science 80: 237-249. Danks, H.V. and J.A. Downes (Eds.). 1997. Insects of the Yukon. Biological Survey of Canada Monograph Series No. 2., Ottawa. 1034 pp. Danks, H.V. and N.N. Winchester. 2000. Terrestrial arthropod biodiversity projects - building a factual foundation. Biological Survey Of Canada (Terrestrial Arthropods) Document Series No. 7. 38 pp. Dindal, D.L. (Ed.). 1990. Soil Biology Guide. Wiley & Sons, NY. 1349
pp. Dwyer, E., D.J. Larson and I.D. Thompson. 1998. Oribatid mite communities of old balsam fir (Aibes balsamea (L.)) forests of western Newfoundland, Canada. Pedobiologia 42: 331-347. Fagan, L.L. and N.N. Winchester. 1999. Arboreal arthropods: diversity and rates of colonization in a temperate montane forest. Selbyana 20: 171-178. Ferguson, S.H. 2001. Changes in trophic abundance of soil arthropods along a grass-shrub-forest gradient. Canadian Journal of Zoology 79: 457-464. Fjellberg, A. 1992. Hypogastrura (Mucrella) arborea sp.nov., a tree-climbing species of Collembola (Hypogastruridae) from Vancouver Island, British Columbia. Canadian Entomologist 124: 405-407. Finnamore, A.T. 1996. The SAGE Project: Sampling protocols for microarthropods. EMAN and Partners Publications. http://www.cciw.ca/eman-temp/reports/publications/sage/intro.htm Finnamore, A.T. and S.A. Marshall (Eds.). 1994. Terrestrial arthropods of peatlands with particular reference to Canada. Memoirs of the Entomological Society of Canada 169: 289 pp. Finnamore, A.T., N.N. Winchester, and V.M. Behan-Pelletier. 1998. Protocols for Measuring Biodiversity: Arthropod Monitoring in Terrestrial Ecosystems. Biodiversity Science Board of Canada, Ecological Monitoring and Assessment Network, Burlington, Ontario. Fons, J. and K. Klinka. 1998. Temporal variations of forest floor properties in the coastal western hemlock zone of southern British Columbia. Canadian Journal of Forest Research 28: 582-590. Fox, C.A. 1998. Soil characterization of continuous tomato and wheat-soybean-tomato crop rotation at Leamington site. pp. 115-126, ref 139-140 in R.W. Johnston (Ed.), Crop rotations and cover crop effects on processing tomato yield, soil erosion and soil properties Southwestern Ontario. Research Sub-Program Canada-Ontario Agriculture Green Plan, COESA Report No.: RES/FARM-007/97. 148 pp. Fox, C.A., E.J.A. Fonseca, J.J. Miller, and A.D. Tomlin. 1999. The influence of row position and selected soil attributes on Acarina and Collembola in no-till and conventional continuous corn on a clay loam soil. Applied Soil Ecology 13:1-8. Freckman, D. 1994. Life in the Soil. Soil Biodiversity: its importance to ecosystem processes. Report of a Workshop Held at The Natural History Museum, London, England. August 30-September 1, 1994. Janssens, F. 2001. Checklist of Collembola and Collembola Key2000 Project. Kaneko, N., M.A. McLean, and D. Parkinson. 1998. Do mites and Collembola affect pine litter fungal biomass and microbial respiration. Applied Soil Ecology 9:209-213. Kevan, P.G., B.C. Forbes, S.M. Kevan and V.M. Behan-Pelletier. 1995. Vehicle tracks on high Arctic tundra: their effects on the soil, vegetation, and soil arthropods. Journal of Applied Ecology. 32: 655-667. Klironomos, J.N. and M. Hart. 2001. Animal nitrogen swap for plant carbon. Nature 410: 652. Klironomos, J.N. and W.B. Kendrick. 1995. Stimulative effects of arthropods on endomycorrhizas of sugar maple in the presence of decaying litter. Functional Ecology 9: 528-536. Klironomos, J.N. and P. Moutoglis. 1999. Colonization of nonmycorrhizal plants by mycorrhizal neighbours as influenced by the collembolan, Folsomia candida. Biology and Fertility of Soils 29: 277-281. Klironomos, J.N. and M. Ursic. 1998. Density-dependent grazing on the extraradical hyphal network of the arbuscular mycorrhizal fungus, Glomus intraradices, by the collembolan, Folsomia candida. Biology and Fertility of Soils 26:250-253. Klironomos, J.N., P. Widden, and I. Deslandes. 1992. Feeding preferences of the collembolan Folsomia candida in relation to microfungal succession on decaying litter. Soil Biology and Biochemistry 24: 685-692. Kosztarab, M. and C.W. Schaefer. 1990. Systematics of the North American insects and arachnids: status and needs. Virginia Agricultural Experimental Station, Information Series 90-1: 247pp. Maraun, M., S. Visser, and S. Scheu. 1998. Oribatid mites enhance the recovery of the microbial community after a strong disturbance. Applied Soil Ecology 9: 175-181. Marshall, SA, R.S. Anderson, R.E. Roughley, V. Behan-Pelletier, and H.V. Danks. 1994. Terrestrial arthropod biodiversity: planning a study and recommended sampling techniques. Bulletin of the Entomological Society of Canada 26(1), Suppl. 33 pp. http://www.biology.ualberta.ca/esc.hp/bsc/briefs/brterrestrial.htm Marshall, V.G., D.K. McE. Kevan, J.V. Matthews Jr., and A.D. Tomlin. 1982. Status and research needs of Canadian arthropods. A brief prepared by the Biological Survey of Canada (Terrestrial Arthropods). Bulletin of the Entomological Society of Canada 14(1), Suppl. 5 pp. McLean, M.A. 1998. Impacts of the epigeic earthworm Dendrobaena octaedra on oribatid mite community diversity and microarthropod abundances in pine forest floor: a mesocosm study. Applied Soil Ecology 7: 125-136. Neave, P. and C.A. Fox. 1998. Response of soil invertebrates to reduced tillage systems established on a clay loam soil. Applied Soil Ecology 9: 423-428. Niedbała, W. 1992. Phthiracaroidea (Acari, Oribatida) systematic studies. Elsevier, Amsterdam. 612 pp. Paquin, P. and D. Coderre. 1997a. Changes in soil macroarthropod communities in relation to forest maturation through three successional stages in the Canadian boreal forest. Oecologia (Berlin) 112: 104-111 Paquin, P. and D. Coderre. 1997b. Deforestation and fire impact on edaphic insect larvae and other macroarthropods. Environmental Entomology 26: 21-30. Puvanendran, D.C., D.J. Larson and I.D. Thompson. 1997. Collembola (Arthropoda) of Balsam fir (Abies balsamea (L.)) forests of western Newfoundland. Canadian Entomologist 129: 505-517. Reeves, R.M. and V. Behan-Pelletier. 1998. The genus Carabodes (Acari: Oribatida: Carabodidae) of North America, with descriptions of new western species. Canadian Journal of Zoology 76: 1898-1921. SCOPE, 2001. http://www.nrel.colostate.edu/soil/SCOPE/scope.html. SCOPE publications relevant to soil biodiversity: http://www.icsu-scope.org/projects/soilsed.htm. Scudder, G.G.E. 1994. An annotated systematic list of the potentially rare and endangered freshwater and terrestrial invertebrates in British Columbia. Entomological Society of British Columbia Occasional Paper 2. 92 pp. Scudder, G.G.E. 2000. Arthropod species at risk and livestock grazing in the South Okanagan of British Columbia. Newsletter, Arthropods of Canadian Grasslands 6: 15-20. Setäla, H., V.G. Marshall and J.A. Trofymow. 1995. Influence of micro- and macro-habitat factors on collembolan communities in Douglas-fir stumps during forest succession. Applied Soil Ecology 2: 227-242. Setäla, H., V.G. Marshall, and J.A. Trofymow. 1996. Influence of body size of soil fauna on litter decomposition and 15N uptake by poplar in a pot trial. Soil Biology and Biochemistry 28: 1661-1675. Shorthouse, J.D. and G. Bagatto 1995. Diversity of carabid ground beetles as bioindicators of mine tailings restoration. Report for Inco Ltd., Dec. 1995. Smiley, R. L. 1992. The predatory mite family Cunaxidae (Acari) of the world with a new classification. Indira Publishing House, West Bloomfield, Michigan. 356 pp. Smith, I.M., E.E. Lindquist, and V. Behan-Pelletier. 1996. Mites (Acari). in: I.M. Smith (Ed.), Assessment of Species Diversity in the Mixedwood Plains Ecozone . Ecological Monitoring and Assessment Network. EMAN and Partners Publications. Smith, I.M., E.E. Lindquist, and V. Behan-Pelletier. 1999 Mites (Acari). in: I.M. Smith (Ed.), Assessment of species diversity in the Montane Cordillera Ecozone. Ecological Monitoring and Assessment Network. EMAN and Partners Publications. http://www.eman-rese.ca/eman/reports/publications/99_montane/ Spence, J. (Ed.). 1986. Faunal influences on soil structure. Quaestiones Entomologicae 21: 371.1-700 [1985]. St. John, M. G., V.M. Behan-Pelletier, G. Bagatto, E.E. Lindquist, J.D. Shorthouse, and I. M. Smith. 2001. Soil formation and mite colonization of rehabilitated mine tailings, Sudbury Canada. Poster presented at Soil Ecology Society Conference, Georgia, May 2001. http://res2.agr.ca/ecorc/staff/behan/poster01.pdf Therrien, F., M. Chagnon, and C. Hébert. 1999. Biodiversity of Collembola in sugar maple (Aceraceae) forests. Canadian Entomologist 131: 613-628. Tomlin, A.D., C.M. Tu, and J.J. Miller. 1995. Response of earthworms and soil biota to agricultural practices in corn, soybean and cereal rotation. Acta Zoologica Fennica. 196: 195-199. Wall, D. (Ed.). 1999. BioScience, February 1999, Special Issue, with six articles written by participants of the 1997 AAAS symposium “Global Biodiversity: Is it in the Mud and Dirt” and the First International Workshop of the SSBEF Committee in Wageningen, Netherlands, 1997, addressing the status of biodiversity in soils and sediments and future directions. Wall Freckman, D.H., T.H. Blackburn, L. Brussaard, P. Hutchings, M.A. Palmer, and P.R. Snelgrove. 1997. Linking biodiversity and ecosystem functioning of soils and sediments. Ambio 26: 556-562. Winchester, N.N. 1997. Canopy arthropods of coastal Sitka spruce trees on Vancouver Island, British Columbia, Canada. In: Stork, N.E. Adis, J. and Didham, R.K. (eds.) Canopy Arthropods. Chapman & Hall, London, pp. 151-168. Winchester, N.N., V.M. Behan-Pelletier, and R.A. Ring. 2000. Arboreal specificity, diversity and abundance of canopy-dwelling oribatid mites (Acari: Oribatida). Pedobiologia. 43: 391-400. Young, I.M., E. Blanchart, C. Chenu, M. Dangerfield, C. Fragoso, M. Grimaldo, J. Ingram, and L.J. Monrozie. 1998. The interaction of soil biota and soil structure under global change. Global Change Biology 4: 703-712. Zacharda, M. 1997. New taxa of Rhagidiidae (Acari: Prostigmata) from North America. Part V. Genus Robustocheles Zacharda, with a key to world species of the genus. Journal of Natural History 31: 1075-1103.
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