And Feather Mites (Acari: Astigmatina: Analgoidea, Pterolichoidea): Ectosymbionts of Grassland Birds in Canada
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139 Chapter 5 Chewing Lice (Insecta: Phthiraptera: Amblycera, Ischnocera) and Feather Mites (Acari: Astigmatina: Analgoidea, Pterolichoidea): Ectosymbionts of Grassland Birds in Canada Terry D. Galloway Department of Entomology, University of Manitoba, Winnipeg, Manitoba, Canada, R3T 2N2 Heather C. Proctor Department of Biological Sciences, University of Alberta, Edmonton, Alberta, Canada, T6G 2E9 Sergei V. Mironov Zoological Institute, Russian Academy of Sciences, Universitetskaya emb. 1, Saint Petersburg, 199034, Russia Abstract. The diversity of bird species nesting in Canadian grasslands is high, and each of them is host to one or more species of lice and mites. A checklist of feather lice (Phthiraptera: Amblycera, Ischnocera) and feather mites (Acari: Analgoidea, Pterolichoidea) from 160 species of birds that nest in terrestrial and aquatic habitats in the grassland biome in Alberta, Saskatchewan, and Manitoba is provided here. The list includes known and expected records. In total, four families, 54 genera, and 183 named species and subspecies of chewing lice are known to occur on these hosts in the Prairie Provinces. At least an additional 63 species are expected to be found eventually, on the basis of records from these hosts elsewhere in North America. Twenty-seven species in six genera have been collected from hosts with no prior louse records; many of these species are undescribed. For feather mites, 20 families, 73 genera, 134 named species, and 38 undescribed species are known to occur, and an additional 13 genera and 39 species are expected to be found eventually on these hosts in Canadian grasslands. Many populations and species of grassland-breeding birds are endangered in North America, and it is clear that should they be lost, many species of lice and mites would disappear with them. Résumé. De très nombreuses espèces d’oiseaux nichent dans les prairies canadiennes et chacune est l’hôte d’une ou de plusieurs espèces de poux et d’acariens. Ce chapitre présente une liste des espèces connues ou prévues de poux d’oiseaux (Phthiraptera : Amblycera, Ischnocera) et d’acariens (Acari : Analgoidea, Pterolichoidea) répertoriées chez 160 espèces d’oiseaux nichant dans les habitats terrestres ou aquatiques du biome des prairies, en Alberta, en Saskatchewan et au Manitoba. La liste recense au total quatre familles, 54 genres et 183 espèces et sous-espèces décrites. Compte tenu des données recueillies sur les hôtes ailleurs en Amérique du Nord, on s’attend par ailleurs à découvrir 63 espèces supplémentaires de ces insectes. Vingt-sept espèces et six genres ont été répertoriés sur des hôtes au sujet desquels il n’existait aucune donnée concernant les poux d’oiseaux. Plusieurs de ces espèces n’avaient jamais été décrites. S’agissant des acariens, on a confi rmé la présence de 20 familles, 73 genres, 134 espèces décrites et 38 espèces non décrites, et on soupçonne la présence sur les hôtes aviaires des prairies canadiennes de 13 genres et de 39 espèces supplémentaires. Plusieurs des populations et des espèces d’oiseaux qui se reproduisent dans les Prairies sont menacées en Amérique du Nord, et il est clair que si ces espèces venaient à disparaître, de nombreuses espèces de poux et d’acariens disparaîtraient également. Galloway, T. D., H. C. Proctor, and S. V. Mironov. 2014. Chewing Lice (Insecta: Phthiraptera: Amblycera, Ischnocera) and Feather Mites (Acari: Astigmatina: Analgoidea, Pterolichoidea): Ectosymbionts of Grassland Birds in 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. 139-188. © 2014 Biological Survey of Canada. ISBN 978-0-9689321-6-2 doi: http://dx.doi.org/10.3752/9780968932162.ch5 140 T. D. Galloway, H. C. Proctor, and S. V. Mironov Introduction Many birds that are grassland specialists nest only in areas of continuous grass. However, grasslands offer a diversity of additional habitats, including bluffs of trees, coulees, wetlands, pothole lakes, streams, and rivers. In combination, this patchwork supports a considerable diversity of birds that nest in suitable habitat wherever it occurs. Cultivation for agricultural production and fragmentation of grassland ecosystems have had a major impact on the welfare of these nesting birds, and indeed, grassland specialists are among the species most seriously affected by human activity in North America (Herkert 1995). As concerns for conservation of grassland birds increase, our understanding of factors affecting their populations has also increased (Koper and Nudds 2011). Considerable attention has been focused on nesting success as affected by habitat fragmentation, especially related to the impact of predation (see review by Stephens et al. 2003). However, the role played by parasites and other symbionts on breeding success and fi tness in avian populations has largely been ignored. From the time it hatches until it dies, each bird is a fl ying menagerie of associated internal and external symbionts. It experiences a dynamic relationship with mites, lice, fl ies, fl eas, bugs, protists, viruses, nematodes, trematodes, tapeworms, and acanthocephalans (Rothschild and Clay 1953), changing in species and numerical composition depending on the age and fi tness of the host, nature of association with conspecifi cs, season, food, habitat, and attack by arthropod vectors. And of course, this impressive diversity of associates is largely unseen by most people, even many of those who work closely with the birds. Conversely, declining host populations may also play a role in the decline and even the extinction of ectosymbionts, but this aspect of the host–symbiont relationship has received relatively little attention (Rózsa 1992; Koh et al. 2004; Whiteman and Parker 2005) compared with that given to declines and extinctions in bird populations. Considering that each bird is host to such tremendous diversity of symbionts (Rothschild and Clay 1953), it is worthwhile to explore the status of this diversity in grassland birds in Canada. Galloway and Danks (1990) emphasized the defi ciency in our knowledge of ectosymbionts of birds in Canada, especially for feather mites and chewing lice. These two groups are taxonomically and ecologically diverse, with the potential to elucidate important evolutionary and phylogenetic relationships in their hosts (Marshall 1981; Johnson and Clayton 2003; Proctor 2003). Some species (i.e., ectoparasites) may affect the health and fi tness of their hosts, an important consideration for host species at risk. Conversely, a reduction in bird host populations may have a major impact on the survival of host-dependent species of ectosymbionts. There has not been a review of species of these arthropod groups in Canada since Wheeler and Threlfall (1989). Given recent published and unpublished studies on the biodiversity of these important ectosymbionts, it is our objective to summarize the current status of our knowledge about chewing lice and feather mites known or likely to be associated with birds nesting in grasslands in the Canadian Prairie Provinces. Scope of the Study The fi rst challenge in this review was to decide what constitutes a “grassland” bird. We chose to include species that breed within areas of continuous grassland in Manitoba, Saskatchewan, and Alberta, including shortgrass, mixedgrass, and tallgrass prairie, as described by Shorthouse and Larson (2010: Fig. 1). We also include forest birds that breed Chewing Lice (Insecta: Phthiraptera: Amblycera, Ischnocera) and Feather Mites 141 (Acari: Astigmatina: Analgoidea, Pterolichoidea): Ectosymbionts of Grassland Birds in Canada in shrubbery, bluffs, and coulees in grasslands, as well as water birds that breed in and along wetlands and waterways within grassland habitats. Species that migrate through grasslands to nest in the boreal, subarctic, or arctic regions are not included in this survey. Species introduced into North America (e.g., ring-necked pheasant, rock pigeon, house sparrow) are not included, though some do nest in our defi ned habitats in grasslands. Our species list was drawn from various sources, primarily Godfrey (1986), Carey et al. (2003), and Penner (2007). In total, we include lice and mites associated with 160 species of birds that nest in grasslands in Canada. Nomenclature for birds follows the American Ornithologists’ Union (1998 and supplements). Sources of Data Both published and new records for ectosymbionts are included in this review. Nomenclature for chewing lice follows Price et al. (2003). Undescribed louse species are included in our list; where genera are in need of extensive taxonomic revision (e.g., Menacanthus (from Galliformes), Anaticola, Brueelia, Craspedorrhynchus, Philopterus), species names for taxa known to occur on certain bird hosts are assigned on the basis of host association, with the understanding that nomenclatural changes are expected. Nomenclature for feather mites (Acari: Astigmatina: Analgoidea, Pterolichoidea) follows Gaud and Atyeo (1996) with these exceptions: recognition of Pteronyssidae as a family rather than as a subfamily of the Avenzoariidae (Mironov 2001), addition of genera described since 1996 (e.g., Mironov et al. 2007; Valim and Hernandes 2010), and placement of those families previously in the Freyanoidea within the Pterolichoidea (OConnor 2009). Feather mite records include described species, undescribed and clearly new species (indicated by “n. sp.” in tables), and genus-level records for species represented