and Heather Proctor**
and Heather Proctor**
Birds are host to a diverse fauna of symbiotic arthropods. Mites (Arachnida: Acari) represent one of the most species-rich groups of arthropods associated with birds (Proctor and Owens 2000). Some mite species are relatively benign such as most feather mites (Astigmata: Analgoidea, Pterolichoidea, Freyanoidea), but others are highly detrimental parasites such as the nasal mite Sternostoma tracheacolum Lawrence, 1948 (Mesostigmata: Rhinonyssidae) (Stephan et al. 1950; Proctor and Owens 2000). The Rhinonyssidae are obligate blood-feeding endoparasites in the nasal passages of non-ratite birds worldwide. Rhinonyssids are distributed among eight genera, believed to have descended from ectoparasitic ancestors related to the Macronyssidae (Strandtmann 1948). Rhinonyssid genera vary in their degree of host specificity, with some genera being restricted to a single host family, and others collected in hosts from different orders (Pence 1973c). The genera Ptilonyssus (Berlese and Trouessart), Sternostoma Berlese and Trouessart, Tinaminyssus Strandtmann and Wharton, and Rhinonyssus Trouessart exhibit broad host ranges in North America. Ptilonyssus species have been collected from passeriform, caprimulgiform, falconiform, and apodiform hosts, Sternostoma from passeriform, charadriiform, and piciform birds, Tinaminyssus from ciconiiform and columbiform hosts, and Rhinonyssus from anseriform, charadriiform, and podicipediform birds. Among more host-specific rhinonyssids, Rhinoecius Cooreman species parasitize owls (Strigiformes), with each species generally occurring in a different species of owl; Rallinyssus Strandtmann species parasitize birds of the family Rallidae; and Larinyssus Strandtmann species parasitize gulls and terns (Laridae).
Rhinonyssids are sluggish, slow moving mites that usually occupy the scroll-like and highly vascularized nasal turbinates, but some species invade the lungs, air sacs, tracheal tissues and body cavity (Porter and Strandtmann 1952; Krantz 1978). Rhinonyssid feeding activity may cause trauma to the nasal epithelium (De-Rojas et al. 2002), but generally they are not considered to cause significant pathology. The main exception is S. tracheacolum which invades the lower respiratory tract, lungs, and air sacs of the host (Stephan et al. 1950). Sternostoma tracheacolum has been collected from 37 species, 32 genera and 11 families of captive and wild birds (Bell 1996). Generally the pathology experienced by captive birds is more severe than that of wild birds (Fain and Hyland 1962). The decline of the endangered Gouldian finch, Erythrura gouldiae (Gould), may be the result of S. tracheacolum infesting wild birds from individuals originally held in captivity (Tidemann et al. 1992).
While surveying the parasitic nasal mites associated with birds of Alberta and Manitoba (Knee et al. 2008), we became aware of the need for an image-rich, interactive key to rhinonyssids of Canada. There has never been a key to Canadian rhinonyssids. The best previously published key for nasal mites of North American birds is a paper-based dichotomous key by Pence (1975), but the key lacks a number of rhinonyssid species known to occur in Canada. Because most bird species in Canada are widely distributed and migrate annually (Kaufman 2000), a key to rhinonyssids of Canada should be useful throughout much of northern North America.
Collection of mites from birds took place independently in the laboratories of Heather Proctor at the University of Alberta and Terry Galloway at the University of Manitoba. At the University of Alberta, we examined approximately 450 frozen bird carcasses from Alberta, largely from the contributions of the Alberta Fish & Wildlife Forensic Laboratory, the Royal Alberta Museum, waterfowl hunters, and colleagues at the University of Alberta. Collection data were sparse for many of these specimens, and for some it can only be said that the birds were collected somewhere in Alberta. Bird bodies were maintained at -20ºC until processing. A partially thawed bird was placed in a suitably sized container, ranging from 4-18 L, with a drop of dish detergent, enough 95% ethanol to soak the plumage of the bird, and enough water to submerge it, the nares and mouth were rinsed with 95% ethanol as well. The whole bird was washed in order to collect all mites and lice (Phthiraptera) associated with the host. The sealed container was then shaken vigorously for five minutes. Particularly large birds were washed in a basin and thoroughly massaged while in the solution. Each bird was then removed from the container and rinsed thoroughly over a Fisher Scientific 53 µm mesh sieve; large birds were rinsed over the washing basin. The washing liquid was filtered and the container and lid were rinsed thoroughly over the same 53 µm sieve. The material remaining in the sieve was stored in 30 mL snap cap or scintillation vials in 80% ethanol.
At the University of Manitoba, nasal cavities were flushed using orthodontic syringes, 15 mL for larger birds and 3 mL for smaller birds. A solution of warm water and mild soap was flushed through each nostril, back out the mouth and into a Petri dish. Occasionally nasal mites were also collected in whole-body washings of birds. Body-washing methods were similar to those described above, except that ethanol was not added to the washing solution, and the washing solution was filtered through a 90 µm sieve. Specimens were preserved in 70% ethanol.
Mites were removed from ethanol and cleared in 85% lactic acid for 1-24 hours depending on the degree of original opacity. Mites were mounted in a polyvinyl alcohol medium (6371A, BioQuip Products, Rancho Dominguez, CA). Slides were cured on a slide warmer at about 40ºC for 3-4 days. Slide-mounted specimens were examined on a Leica DMLB compound microscope with differential interference contrast (DIC) at 400x magnification. Species level identifications were made using keys (Pence 1975) and species descriptions from the primary literature.
The HTML-based dichotomous key to the female Rhinonyssidae species of Canada was made using FrontPage. Illustrations were made on paper using camera lucida, merged in Adobe Photoshop CS version 8.0, and redrawn in Adobe Illustrator CS version 11.0 using an Intuos 2 Graphics Tablet from WACOM Co., Ltd. FrontPage was used to create an HTML ‘species page’ for each species. Each of these species pages includes North American host records, diagnostic characteristics, relevant references, and scaled habitus images. The Sternostoma laniorum Fain, 1956 habitus image was redrawn from Pence (1975) with permission from Pence, due to the poor condition of the single specimen in our possession. Rallinyssus caudistigmus Strandtmann, 1948 and Sternostoma boydi Strandtmann, 1951 were not collected during our survey, but these species are included in the key since specimens were collected in Canada and deposited in the Canadian National Collection of Insects, Arachnids and Nematodes. Most character states are accompanied by a labeled image indicating the structures of interest. Illustrations of the dorsum and venter of a generalized rhinonyssid female are also included. Photos of slide-mounted mites were taken with a Canon Powershot S40 digital camera mounted on a Leica DMLB compound microscope with differential interference contrast (DIC), at 200-400x magnification. Images were captured in the Canon Utilities Remote Capture program, version 22.214.171.124, and edited using Adobe Photoshop CS version 8.0. Voucher specimens are deposited in the University of Alberta E.H. Strickland Museum of Entomology, and in the University of Manitoba J.B. Wallis Museum of Entomology. Host taxonomy and authorities follow Clements (1991) provided by Andrew and McAllan (1998) in Nomina version 1.0, using the ‘Clements 1991-1996’ taxonomy option.
This key provides species level identifications for 47 species and six genera of rhinonyssids collected from birds of Canada (Table 1), illustrated in 116 figures. The key includes all of the known rhinonyssid species from Canada, with 45 species collected from birds of Alberta and Manitoba, five of which were described as new to science (Knee 2008), and two species from birds of the Northwest Territories. The illustrations of the dorsum and venter of a generalized rhinonyssid female, the labeled images indicating the structures of interest, and HTML species pages can be accessed by clicking on the links in the key. Additional keys to species of nasal mites collected from North American birds can be found in Pence (1975). Keys to other blood- and tissue-feeding mites associated with birds of Alberta can be found in Knee and Proctor (2006).
Occasionally certain character states were inconsistent intraspecifically, including the presence of unpaired setae, the extent of dorsal or ventral shields, and the form of shield margins; this variation was taken into consideration when building the key. In addition, the posterior pair of “ventral” opisthosomal setae (JV4) are sometimes shifted dorsally or terminally, making them difficult to recognize as being “ventral”, in such cases setae JV4 were excluded from the total count for ventral opisthosomal setae. Intraspecific morphological variation can be quite common, and it must be taken into consideration while creating and using any key. Records of the same species of rhinonyssid in related European and North American birds, or of the same mite species in very different host families, should be interpreted cautiously. Careful morphological and molecular investigations are likely to reveal that many such species are actually clusters of a closely related species, or are more distantly related species that have independently converged on similar morphology. This key is based upon the species collected during a survey of approximately 35% of Canada’s avian fauna, there are many host species left to be examined (e.g., Procellariiformes, marine Charadriiformes) (Knee et al. 2008). It has been estimated that there are 70-80 species of rhinonyssids in Canada; thus, this key is not complete, and it is likely that many more species will be uncovered with further examination of Canada’s avian fauna (Knee et al. 2008). This key will serve as the foundation for any future work on avian nasal mites in Canada.
Table 1. Rhinonyssid species associated with Canadian birds.
|Ptilonyssus acrocephali||Fain, 1964|
|Ptilonyssus angrensis||(Castro, 1948)|
|Ptilonyssus bombycillae||Fain, 1972|
|Ptilonyssus callinectoides||(Brooks and Strandtmann, 1960)|
|Ptilonyssus calvaria||Knee, 2008|
|Ptilonyssus carduelis||Fain, 1962|
|Ptilonyssus cerchneis||Fain, 1957|
|Ptilonyssus coccothraustis||Fain and Bafort, 1963|
|Ptilonyssus echinatus||Berlese & Trouessart, 1889|
|Ptilonyssus euroturdi||Fain & Hyland, 1963|
|Ptilonyssus hirsti||(Castro & Periera, 1947)|
|Ptilonyssus icteridius||(Strandtmann and Furman, 1956)|
|Ptilonyssus japuibensis||Castro, 1948|
|Ptilonyssus morofskyi||Hyland, 1962|
|Ptilonyssus nivalis||Knee, 2008|
|Ptilonyssus nudus||Berlese & Trouessart, 1889|
|Ptilonyssus perisorei||George, 1961|
|Ptilonyssus pinicola||Knee, 2008|
|Ptilonyssus pirangae||(Cerny, 1969)|
|Ptilonyssus plesiotypicus||Knee, 2008|
|Ptilonyssus sairae||Castro, 1948|
|Ptilonyssus troglodytis||Fain, 1964|
|Ptilonyssus tyrannus||(Brooks and Strandtmann, 1960)|
|Ptilonyssus vireonis||(Dusbabek, 1969)|
|Rallinyssus caudistigmus||Strandtmann, 1948|
|Rhinoecius aegolii||Butenko, 1971|
|Rhinoecius alifanovi||Butenko, 1976|
|Rhinoecius brikinboricus||Butenko, 1976|
|Rhinoecius cooremani||Strandtmann, 1952|
|Rhinoecius grandis||Strandtmann, 1952|
|Rhinoecius nycteae||Butenko, 1976|
|Rhinonyssus coniventris||Trouessart, 1894|
|Rhinonyssus rhinolethrum||(Trouessart, 1895)|
|Sternostoma boydi||Strandtmann, 1951|
|Sternostoma cryptorhynchum||Berlese and Trouessart, 1889|
|Sternostoma hylandi||Fain and Johnston, 1966|
|Sternostoma laniorum||Fain 1956|
|Sternostoma longisetosae||Hyland, 1961|
|Sternostoma loxiae||Fain, 1965|
|Sternostoma porteri||Hyland, 1962|
|Sternostoma sialiphilus||Hyland and Ford, 1961|
|Sternostoma technaui||(Vitzthum, 1935)|
|Sternostoma tracheacolum||Lawrence, 1948|
|Sternostoma setifer||Knee, 2008|
|Tinaminyssus columbae||(Crossley, 1950)|
|Tinaminyssus melloi||(Castro, 1948)|
|Tinaminyssus zenaidurae||(Crossley, 1952)|
We particularly thank Bob McClymont and other personnel at the Alberta Fish and Wildlife Forensic Laboratory, as well as Jocelyn Hudon at the Royal Alberta Museum, for providing numerous specimens. We thank Dr. Terry Galloway for providing us with numerous specimens from Manitoba birds. Sergei Mironov forwarded some material from birds from Manitoba that had been sent to the Zoological Institute of the Russian Academy of Sciences. We thank Dr. Valerie Behan-Pelletier for access to her equipment. Stephanie Grundke and Samantha Wojtkiw washed birds during the summer of 2003, and Nikolas Romaniuk and Michael Pedruski did the same during the summer of 2004. We also thank Dave Holder, Lisa Babey, Chérie Dugal, Todd Whiklo and Debra Wytrykush for assistance in the lab in Manitoba, and this work could not have been done without the support of staff at the Manitoba Wildlife Rehabilitation Organization (now Wildlife Haven) and Manitoba Conservation. This research was conducted with scientific salvage permits to take migratory birds from Environment Canada (permit no. CWS03-A009 and CWS99-M023), and permits to salvage found dead wildlife (excluding endangered species) from Alberta Environment (current permit no. 20526) and Manitoba Conservation (permit no. WB02322). This project was funded by NSERC Discovery Grant to H. Proctor, as well as an Alberta Conservation Association Biodiversity Grant to W. Knee.
Andrew, P., and McAllan, I. 1998. Nomina: Global Bird Dictionary, Relational Taxonomies. CSIRO Publishing, Collingwood, Australia.
Bell, P.J. 1996. Survey of the nasal mite fauna (Rhinonyssidae and Kytoditidae) of the Gouldian finch, Erythrura gouldiae, and some co-occurring birds in the Northern Territory. Wildlife Research, 23: 675-685.
Berlese, A., and E.L. Trouessart. 1889a. Diagnoses d’acariens nouveaux ou peu connus. Bulletin de la Societe de Sciences Naturelles de l’Ouest de la France (Nantes), 2: 126-130.
Berlese, A., and E.L. Trouessart. 1889b. Diagnoses d’acariens nouveaux ou peu connus. Bulletin de la Bibliotheque Scientifique de l’Ouest, 2: 121-143.
Brooks, D.L., and Strandtmann, R.W. 1960. The nasal mites (Acarina) of some West Texas flycatchers (Tyrannidae). Journal of Parasitology, 46: 418-432.
Butenko, O.M. 1971. A summary of the knowledge of the nasal mites of birds of the Okskii Reserve Area. Trudy Okskogo Gosudarstvennogo Zapovednika, 8: 204-223.
Butenko, O.M. 1976. New species of rhinonyssid mites (Gamasoidea, Rhinonyssidae), parasitic in owls. Parazitologiya (St. Petersburg), 10: 303-309.
Butenko, O.M. 1984. Non-passerine nasal mites of Russia. Zool. Inst. Acad. Sciences (St. Petersburg). 187pp.
Castro, M.P. 1948. Reestruturacao generica da familia Rhinonyssidae Vitzthum 1935 (Acari: Mesostigmata: Gamasides) e descraicao de algumas especies novas. Arquivos do Instituto Biologico, 18: 253-284.
Castro, M.P., and C. Pereira. 1947. Acaros nasicolas (Parasitiformes: Rhinonyssidae) do pardal – “Passer domesticus L.”. Arquivos do Instituto Biologico, 18: 125-133.
Cerny, V. 1969. Six new nasal mites (Mesostigmata, Ptilonyssidae) from Cuban birds. Folia Parasitologica, 16: 227-235.
Clements, J.F. 1991. Birds of the World: A Checklist, 4th ed. Ibis Publishing, Vista, California.
Crossley, D.A. 1950. A new species of nasal mite Neonyssus (Neonyssus) columbae, from the pigeon. Proceedings of the Entomological Society of Washington, 52: 309-313.
Crossley, D.A. 1952. Two new nasal mites from Columbiform birds. Journal of Parasitology, 38: 385-390.
De-Rojas, M., Mora, M.D., Ubeda, J.M., Cutillas, C., Navajas, M., and Guevara, D.C. 2002. Phylogenetic relationships in rhinonyssid mites (Acari: Rhinonyssidae) based on ribosomal DNA sequences: insights for the discrimination of closely related species. Parasitology Research, 88: 675-681.
Dusbabek, F. 1969. New species of nasal mites (Acarina: Rhinonyssidae) from Cuban birds. Folia Parasitologica, 16: 213-226.
Estebanes-Gonzalez, M.L. 1997. Acarofauna en nidos de aves silvestres en Mexico. Acta Zoologica Mexicana, 71: 1-15.
Fain, A. 1956. Les acariens de la famille Rhinonyssidae Vitzthum 1935, parasites des fosses nasals des oiseaux au Ruanda-Urundi (note preliminaire). Revue de Zoologie et de Botanique Africaines, 53: 131-157.
Fain, A. 1957. Les acariens des familles Epidermoptidae et Rhinonyssidae parasites des fosses nasales d’oiseaux au Ruanda-Urundi et au Congo Belge. Annales du Musee Royal du Congo Belge Tervuren, 60: 1-174.
Fain, A. 1962. Les acariens parasites nasicoles des oiseaux de Belgique. I. Deux especes nouvelles de Rhinonyssidae (Mesostigmata) avec une liste des especes connues de Belgique. Bulletin and Annales de la Societe Royale d'Entomologie de Belgique, 98: 252-270.
Fain, A. 1963. Les acariens nasicoles des oiseaux de Belgique. II. Description de deux espèces nouvelles. Bulletin and Annales de la Societe Royale d'Entomologie de Belgique, 99: 168-181.
Fain, A. 1964. Les acariens parasites nasicoles des oiseaux de Belgique. IV. Notes sur quelques Rhinonyssidae avec description de deux especes nouvelles. Bulletin and Annales de la Societe Royale d'Entomologie de Belgique, 100: 55-61.
Fain, A. 1965. Diagnoses d’acariens parasites nouveux. Revue de Zoologie et de Botanique Africaines, 72: 152-160.
Fain, A. 1972. Nouveaux Rhinonyssidae (Acarina : Mesostigmata). Acarologia, 14: 357-364.
Fain, A., and Bafort, J. 1963. Les acariens parasites nasicoles des oiseaux de Belgique. III. Nouvelles observations sur les rhinonyssides avec description de cinq especes nouvelles. Bulletin and Annales de la Societe Royale d'Entomologie de Belgique, 99: 471-485.
Fain, A., and Hyland, K.E. 1962. The mites parasitic in the lungs of birds. The variability of Sternostoma tracheacolum Lawrence, 1948, in domestic and wild birds. Parasitology, 52: 401-424.
Fain, A., and Hyland, K.E. 1963. Deus nouveaux rhinonyssides communs aux faunes d'Amerique du Nord et de Belgique. Bulletin and Annales de la Societe Royale d'Entomologie de Belgique, 99: 375-386.
Fain, A., and Johnson, D.E. 1966. Nouveaux acariens nasicoles d'oiseaux nord-americains (Acari: Rhinonyssidae). Bulletins de la Societe Royal de Zoologie d'Anvers, 38: 25-41.
Furman, D.P. 1957. Revision of the genus Sternostoma Berlese and Trouessart. Hilgardia, 26: 473-495.
George, J.E. 1961. The nasal mites of the genus Ptilonyssus (Acarina: Rhinonyssidae) occurring on some North American passeriform birds. Kansas Entomological Society, 34: 105-132.
Hood, D.E., and Welch, H.E. 1980. A seasonal study of the parasites of the red-winged blackbird (Agelaius phoeniceus L.) in Manitoba and Arkansas. Canadian Journal of Zoology, 58: 528-537.
Hyland, K.E. 1961. Sternostoma longisetosa, a new species of nasal mite from the eastern kingbird with notes on the occurrence of Tyranninyssus spinosus Brooks and Strandtmann in southern Michigan (Acarina: Rhinonyssidae). Acarologia, 3: 279-284.
Hyland, K.E. 1962. Two new nasal mites, Ptilonyssus morofskyi, n.sp., and Sternostoma porteri n.sp., from North American birds (Acarina; Rhinonyssidae). Bulletin of the Brooklyn Entomological Society, 57: 146-156.
Hyland, K.E., and Ford, H.G. 1961. Sternostoma sialiphilus n. sp. (Acarina: Rhinonyssidae) from the nasal cavities of the eastern bluebird, Sialia sialis (Linnaeus). Journal of Parasitology, 47: 101-104.
Hyland, K.E., and Moorhouse, A. 1970. Nasal mites from Mexican birds. I. Rhinonyssidae (Mesostigmata) from the host family Tyrannidae. Acarologia, 12: 43-58.
Kaufman, K. 2000. Birds of North America. Houghton Mifflin Company, New York, New York.
Knee, W. 2008. Five new species of Rhinonyssidae (Mesostigmata) and one new species of Dermanyssus (Mesostigmata: Dermanyssidae) from birds of Alberta and Manitoba, Canada. Journal of Parasitology, 94: 348-374.
Knee, W., and Proctor, H. 2006. Keys to the families and genera of blood and tissue feeding mites associated with Albertan birds. Canadian Journal of Arthropod Identification No. 2, 28 June 2006, available online at http://www.biology.ualberta.ca/bsc/ejournal/kp02/kp_02.html
Knee, W., Proctor, H., and Galloway, T. 2008. Survey of nasal mites (Rhinonyssidae, Ereynetidae, and Turbinoptidae) associated with birds in Alberta and Manitoba, Canada. Canadian Entomologist, 140: 364-379.
Krantz, G.W. 1978. A manual of acarology, 2nd ed. Oregon State University Book Stores, Corvallis, Oregon.
Lawrence, R.F. 1948. Studies on some parasitic mites from Canada and South Africa. Journal of Parasitology, 34: 364-379.
Mitchell, R.W. 1961. New avian host records for some mesostigmatid mites. The Southwestern Naturalist, 6: 103-105.
Mitchell, R.W., and Rhodes, W.L. 1960. New host records for the mesostigmatid nasal mite Rhinonyssus rhinonlethrum (Acarina: Rhinonyssidae). The Southwestern Naturalist, 5: 107-108.
Owen, B.L. 1958. Records of nasal mites of the mourning dove. The Texas Journal of Science, 10: 447.
Pence, D.B. 1972a. The nasal mite of birds from Louisiana. I. Dermanyssids (Rhinonyssinae) from shore and marsh birds. Journal of Parasitology, 58: 153-168.
Pence, D.B. 1972b. The nasal mites of birds from Louisiana II. The genus Sternostoma (Dermanyssidae: Rhinonyssinae). Journal of Parasitology, 58: 781-889.
Pence, D.B. 1972c. The nasal mites of birds from Louisiana IV. The genus Ptilonyssus (Dermanyssidae: Rhinonyssinae) with a description of two new species. Journal of Parasitology, 58: 1162-1169.
Pence, D.B. 1972d. The nasal mites of birds from Louisiana III. The genus Ptilonyssus (Dermanyssidae: Rhinonyssinae) with description of a new species. Journal of Parasitology, 58: 790-795.
Pence, D.B. 1973a. The nasal mites of birds from Louisiana. VI. New and additional records of Dermanyssids (Rhinonyssinae) with description of a new species. Journal of Parasitology, 59: 359-362.
Pence, D.B. 1973b. The nasal mites of birds from Louisiana. VIII. Additional records and description of a new species (Acarina: Dermanyssidae, Ereynetidae, Epidermoptidae, and Cytoditidae). Journal of Parasitology, 59: 874-880.
Pence, D.B. 1973c. The nasal mites of birds from Louisiana. IX. Synopsis. Journal of Parasitology, 59: 881-892.
Pence, D.B. 1975. Keys, species and host list, and bibliography for nasal mites of North American birds (Acarina: Rhinonyssinae, Turbinoptinae, Speleognathinae, and Cytoditidae). Special Publications of the Museum Texas Tech University, 8: 1-148.
Pence, D.B., and Casto, S. 1975. Two new species and new records of nasal mites of the genus Sternostoma (Acarina: Rhinonyssinae) from birds in Texas. Journal of Parasitology, 61: 360-368.
Pence, D.B., and Casto, S.D. 1976. Studies on the variation and morphology of the Ptilonyssus "sairae" complex (Acarina: Rhinonyssinae) from North American passeriform birds. Journal of Medical Entomology, 13: 71-95.
Pence, D.B., and Young, V.E. 1979. Rallinyssus sorae sp. n. (Acari: Dermanyssidae: Rhinonyssinae) from the nasal passages of the sora, Porzana carolina (Aves: Rallidae). Journal of Parasitology, 65: 791-793.
Porter, J.C. and Strandtmann, R.W. 1952. Nasal mites of the English Sparrow. The Texas Journal of Science, 4: 393-399.
Proctor, H.C., and Owens, I. 2000. Mites and birds: diversity, parasitism and coevolution. Trends in Ecology and Evolution, 15: 358-364.
Spicer, G.S. 1977a. Two new nasal mites of the genus Ptilonyssus (Mesostigmata: Rhinonyssidae) from Texas. Acarologia, 18: 594-601.
Spicer, G.S. 1977b. New host records from avian nasal mites (Acarina: Rhinonyssinae, Speleognathinae). Journal of Medical Entomology, 14: 498.
Spicer, G.S. 1978. A new species and several new host records of avian nasal mites (Acarina: Rhinonyssinae, Turbinoptinae). Journal of Parasitology, 64: 891-894.
Spicer, G.S. 1987. Prevalence and host-parasite list of some nasal mites from birds (Acarina: Rhinonyssidae, Speleognathidae). Journal of Parasitology, 73: 259-264.
Stephan, S.A.R., Kaschula, V.R., and Canham, F.S. 1950. Respiratory acariasis of canaries. Journal of the South African Veterinary Medical Association, 21: 103-107.
Strandtmann, R.W. 1948. The mesostigmatic nasal mites of birds. I. Two new genera from shore and marsh birds. Journal of Parasitology, 34: 505-514.
Strandtmann, R.W. 1951. The mesostigmatic nasal mites of birds. II. New and poorly known species of Rhinonyssidae. Journal of Parasitology, 37: 129-140.
Strandtmann, R.W. 1952. The mesostigmatic nasal mites of birds, III. New species of Rhinoecius from owls. Proceedings of the Entomological Society of Washington, 54: 205-216.
Strandtmann, R.W. 1956. The mesostigmatic nasal mites of birds. IV. The species and hosts of the genus Rhinonyssus. Proceedings of the Entomological Society of Washington, 58: 129-142.
Strandtmann, R.W. 1961. Neonyssus triangulus n.sp. nasal mite (Acarina: Mesostigmata) from the white winged dove (Aves: Columbiformes) and key to the species of the genus Neonyssus. Journal of Parasitology, 47: 323-328.
Strandtmann, R.W. 1962. A ptilonyssid mite from the sparrow hawk, Falco sparverius. Proceedings of the Entomological Society of Washington, 64: 100-102.
Strandtmann, R.W., and Furman, D.P. 1956. A new species of mite, Paraneonyssus icteridius, from the nasal cavities of blackbirds. Pan-Pacific Entomologist, 32: 167-173.
TerBush, L.E. 1963. Incidence of nasal mites in different age classes of herring gulls (Larus argentatus). Journal of Parasitology, 49: 525.
Tidemann, S.C., McOrist, S., Woinarski, J.C.Z., and Freeland, W.J. 1992. Parasitism of wild Gouldian finches (Erythrura gouldiae) by the air-sac mite Sternostoma tracheacolum. Journal of Wildlife Diseases, 28: 80-84.
Trouessart, E.L. 1894. Note sur les acariens parasites des fosses nasals des oiseaux.Comptes Rendus Hebdomadaires des Seances et Memoires de la Societe de Biologie, 2: 723-724.
Trouessart, E.L. 1895. Note sur un acarien parasite des fosses nasals de l’oie domestique (Sternostomum rhinolethrum, n. sp.). Revue des Sciences Naturelles apliquees, Bulletin de la Societe Nationale d’Acclimation, 42: 392-394.
Vitzthum, H.G. 1935. Milben aus der nasenhohle von Vogeln. Journal fur Ornithologie, 83: 563-587.
Wilson, N. 1964. New records and descriptions of Rhinonyssidae, mostly from New Guinea (Acarina: Mesostigmata). Pacific Insects, 6: 357-388.
Wilson, N. 1968. Records of nasal mites (Mesostigmata: Rhinonyssidae) from New Guinea, Philippines and United States. Journal of Medical Entomology, 5: 211-223.
Wilson, N., and Haas, G.E. 1980. Ectoparasites (Mallophaga, Diptera, Acari) from Alaskan birds. Proceedings of the Entomological Society of Washington, 82: 541-552.
Zamudio, M.L. 1988. Desarrollo de Tinaminyssus melloi (Castro, 1948) (Mesostigmata: Rhinonyssidae). Folia Entomologica Mexicana, 74: 205-214.