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j. RaptorRes. 34(3):210-231 ¸ 2000 The Raptor Research Foundation, Inc.

A REVIEW AND CHECKLIST OF THE PARASITIC (ACARINA) OF THE AND STRIGIFORMES

JAMESR. PHILIPS Math/ScienceDivision, Babson College, Babson , MA 02457-0310 U.S.A.

ABSTRACT.--Referencedchecklists are provided of the 86 speciesof parasiticfeather, quill, respiratory, skin, and nest mites (Acarina) that are known from 116 speciesof ,, , and vultures, and the 91 speciesof parasitic mites known from 51 speciesof . KEYWORDS: Falconiformes;SOigiformes; falcons;, hawks; eagles;, vulturex owls; parasites; • Aca•ina.

Un resumeny listado de piojos (Acarina) en Falconiformesy Strigiformes RESUMEN.--Seprovee un listado referenciado de 86 especiesde piojos (Acarina) en plumas, quilla, aparato respiratorio,piel y nidos conocidosa partir de 116 especiesde gavilanes,figuilas, halcones, y buitres y de 91 especiesde piojos conocidosa partir de 51 especiesde bfihos. [Traducci6n de C•sar Mfirquez]

This review summarizesour current knowledge nately, most mites on raptors are not very patho- of the host-parasiterelationships between parasitic genic and feather mites in particular are usually mites and members of the Falconiformes and Stri- more commensal, rarely causingharm unless they giformes (Appendices I and 2). Over the years, become extremely abundant. there have been many taxonomic name changes Feather Mites. There are seven families of fal- within and especiallymites, so my reviewup- coniform feather mites:Analgidae (Ancyralges),Av- dates old host records to current nomenclature, as enzoariidae (Bonnetella),Cheylabididae ( Cheylabis, well as indicating accidental or misidentified re- Hemicheylabis),Gabuciniidae (Aetacarus,Aposolenz- cords that should not be regarded as very signifi- dia, Hieracolichus,Ramogabucinia), Kramerellidae cant. Nonparasitic nest mites and parasitic mites (Pseudogabucinia) , Pterolichidae ( Pseudalloptinus), from prey occur accidentallyon raptors, and birds and Xolalgidae (Analloptes,Dubininia). Ancyralges in captivity or specimen bags may pick up mites occurs only on vultures and Bonnetellaoccurs only from other speciesas well. My review also includes on Ospreys (Pandionhaliaetus). Only a few individ- records of some new ,as yet undescribed, uals of Ancyralgeshave been collected,but over 300 which I have found through necropsiesof raptors. Bonnetellahave been found on an Osprey (Miller There are 21 families of mites that are associated et al. 1997). The cheylabidid,gabuciniid, and pter- with falconiforms and 17 families associated with olichid genera which occur on raptors do not oc- owls.Mites inhabit birds' feathers, quills, skin and cur on other orders of birds, except for Aetacarus subcutaneoustissues, respiratory tracts,and nests, which includes two speciesassociated with the Oti- and feed on blood, tissue fluid, skin and feather didae. In these genera, speciesrange from monox- lipids and debris, keratin, fungi, algae, and other enous to polyxenous. Pseudogabucinia,Analloptes, mites (Philips 1990, 1993). The mite fauna of most and Dubininia are found on several orders of birds falconiform and strigiform speciesis completely but their falconiform speciesare restricted to this unknown, but these raptors can host a diversemite order. Aetacarus,Hieracolichus, Pseudalloptinus, and community with as many as eight mite species Pseudogabucinialive on the wings,especially the pri- known from the Black (Milvus migrans)and maries and upper primary wing coverts. Over 18 speciesknown from the Long-eared (Asio 15 000 Pseudalloptinushave been found on a single otus).Since mites are so small (0.3-1.5 mm long), Bald (Haliaeetusleucocephalus). Feather mites they are often overlooked,but raptors that appear feed on feather fragments,lipids secretions,skin parasite-freeto the can support populationsof debris, and feather fungi, bacteria,and algae. The 15 000 feather mites and 4000 quill mites. Fortu- diet of Aetacarusand Pseudalloptinusincludes fresh-

210 SEPTEMBER 2000 PARASITIC MITES OF RAPTORS 211 water diatoms which stick to feathers when birds orders.In owls,Paralges colonizes the upper and un- are in water (Dubinin 1956). In great numbers, der trunk feathersbut populationsover 10 havenot feather mites irritate the host with damage result- yet been found on an individual owl. The ing from the 's stressand feather pulling. Vas- Oconnoriidae is known only from the Philippine yukova and Labutin (1990) found that feather BoobookOwl (Ninoxphilippensis) and probablyeats mites occurred on 22% of falconiform birds and the medulla of quills (Gaud et al. 1989). The syrin- 77% of owls in Yakutia. gophilid Bubophilusis known only from the There are three families of owl feather mites: GreatHorned Owl (Philipsand Norton 1978).Two Kramerellidae (Dermonoton,Kramerella, and Petito- thousandBubophilus have been found on one bird, ta), Psoroptoididae (Pandalura), and Xolalgidae inhabiting mainly axillary and nearby wing feathers. (Glaucalges).These genera only occur on owlswith Infestationsof Great quillswith Paralges the exception of one speciesof Glaucalgeswhich and Bubophiluswere photographicallydocumented occurs on Musophagidae. Kramerellaspecies are by Philips (1993). very host specific, occupy primarily wing feathers, Skin Mites. Skin mites of falconiform birds and are often very numerous (thousands) on an which live on the skin surface or burrow into the individual. Philips (1993) photographed Great skin include the families Cheyletiellidae, Epider- Horned Owl (Bubovirginianus) alula feathers with moptidae ( Microlichusand Myialges),Harpyrhynchi- Kramerellainfestations. Kramerellais very common dae (Havpyrhynchus),and Knemidocoptidae(Knem- on owlsand wasfound on 86% of EurasianPygmy- idocoptes).Cheyletiellid mites feed on blood and Owls (Glaucidiumpasserinum) in Thuringia (Cern•, tissuefluid, and most speciesare associatedwith a and Wiesner 1992). Petitota, Pandalura, and Glau- particular family of birds. Microlichusand Myialges calgesspecies are more polyxenousand typicallyoc- are also associatedwith louseflies(Hippoboscidae) cur in smaller numbers on a host (Atyeo and Phil- and are more fly specific. Their bird host range ips 1984). Dermonotonalso is more polyxenousbut tends to correspond to that of their fly host. Fer- population data are lacking. tilized Myialgesfemales parasitize louseflies and lay Quill Mites. There are two families of falconi- their eggson them, but the other stagesof the life form quill mites: Ascouracaridae(Pyonacarus) and cycle are bird parasites.Microlichus is phoretic on Syringophilidae (Peristerophila,undescribed gen- lousefliesand usesthem only for a ride to another era). Ascouracarid mites occur on seven orders of bird host. Microlichuslives in feather bulbs, pro- birds but Pyonacarusis known only from the Black ducing congestionand swelling.These skin mites Kite (Milvus migrans).These mites eat the medulla feed on surface skin debris, keratin, and tissue of quills. Syringophilid mites use their mouthparts fluid. to pierce the quill wall and feed on tissue fluid Skin mites of owls which live on the skin surface from the feather follicle. Feather loss and second- or burrow into the skin include the families Anal- ary bacterial infection can result. Each genus of gidae ( Strelkoviacarus), Epidermoptidae ( Microli- Syringophilidae is primarily or exclusivelyassoci- chus,Myialges, Passeroptes) , Harpyrhynchidae ( Har- ated with a particular order of birds. Peristerophila pyrhynchus), and Knemidocoptidae( Knemidocoptes). is a columbiform mite and P. columbae is known Strelkoviacarus,like Microlichus,is phoretic on - from pigeons ( Columbalivia) and its occurrenceon flies with a broad avian host range. Passeroptesoc- a Red-tailedHawk (Buteojamaicensis)(Casto 1976) curs on Passeriformes and Columbiformes as well is unusual and may be accidental. I have found a as owls,but individual speciesare restrictedto one new genusof syringophilidmite that occurson five order of bird host. North American accipitrid birds. Trunk and scap- Harpyrhynchid and knemidocoptid mites bur- ular feathers are preferred by falconiform syrin- row into the skin, causingitching and mange. Har- gophilid mites. pyrhynchidmite speciesusually have only one avi- There are three families of owl quill mites: Der- an host speciesand occur on the calamus at the moglyphidae (Paralges),Oconnoriidae (Oconnoria), skin surface and in subcutaneous cysts. Schulz and Syringophilidae (Bubophilus).Dermoglyphid (1990) photographicallydocumented feather loss mites can cause extensivemange becauseowls use on the head and neck of a (Aquila their beaksto dig them out. Paralgesoccurs on several chrysaetos)caused by harpyrhynchidmites. This pa- orders of birds, but the undescribedspecies from thology has not been observed in owls. Philips owls (Philips 1993) have not been found on other (1993) photographed a harpyrhynchidembedded 212 PHILIPS VOL. 34, No. 3 in the skin of a (Aegoliusfunereus).Most recordssuggests temporarily successful colonization knemidocoptid mite speciesare polyxenouswithin of other bird hostsin nestingproximity is not un- an order of birds, but those found on owls also common(Pence et al. 1997). This appearsto be the occur on other bird orders. Knemidocoptid mites casewith the record of Hypodectespropus from the live in the stratum corneum of the skin, causing (Speotytocunicularia) and Neottialges hyperplasia, hyperkeratosis, and inflammation. evansifrom the (Tyt0 alba). H. propusis They cause•scaly encrustations on the and associatedwith pigeons,herons, and egretsand N. claws,known as scalyface and scalyleg disease. evansiis a cormorantmite. Gypsodectesis known only This condition in a (Bubo vir- from vultures;Neotytodectes is known only from owls. gtnianus) was documented photographically by Neottialgesis known from four ordersof birds,but the Schulz et al. (1989) and is common in cage birds, specieson falconiformbirds are monoxenous.Tyto- but there is only one record of these mites on fal- dectesoccurs on owls, falcons,and kingfishersand coniform birds (Cooper 1978, 1985). each speciesoccurs on only one host genus.Several Transient skin mites of both bird orders include infestationsin the Barn Owl were photographically the blood-feeding (), documentedby Wurstand Havelka(1997). Macronyssidae (Ornithonyssus),and Laelapidae RespiratoryMites. Respiratorymites of falconi- (Androlaelaps),and tissue-fluid feeding Trombi- form birds include the families Ereynetidae (Boy- culidae (chiggers). Falconiform chiggers include daia, Speleoffnathopsis),Rhinonyssidae (Falconyssus, Blankaartia, Eutrombicula,, Neos- Ptilonyssus,Tinaminyssus), and Turbinoptidae choengastia,Odontacarus, and Ornithogastia,while (Schoutedenocoptes).Ereynetid mites feed on mu- strigiform chiggers include Blankaartia,Euschoen- cous deep in the nasal cavityand nonpasserineer- gastia,Eutrombicula, Hyponeocula, Leptotrombidium, eynetid mites are monoxenousor parasitizevery Miyatrombicula, Neoschoengastia,Odontacarus, Or- few host species.Molted ereynetidmite skinscan nithogastia,and Toritrombicula.Dermanyssid and partially block the nasal cavity.Rhinonyssid mites macronyssidbird parasiteslay eggson the host or feed on blood and occupythe anterior portion of in its nest and chiggers are the parasitic larval the nasal cavity, usually in very small numbers. stage of a predatory soil mite. All four families Each speciesof these rhinonyssidgenera usually usuallyhave relativelylow host specificityand can has only one or several host species.The genus cause dermatitis. Too much blood loss results in Falconyssusoccurs on falconiform and alcedinid energy and weight loss, anemia, and potentially birds, while Ptilonyssusand Tinaminyssusoccur on death. Ornithonyssusoften prefers to feed at the many typesof birds. Turbinoptid mites live in the vent. Dermanyssusfeeds at night. Dermanyssuson a external part of the nares and feed on the corne- Sharp-shinned (Accipiterstriatus) was pho- ous skin there. Most turbinoptid speciesare mo- tographed by Philips (1993). Avian Androlaelaps noxenousor restrictedto one family of birds. species are facultative blood suckerswhich also Respiratorymites of owls include the families prey on other and their , on Cloacaridae( Pneumophagus) , Ereynetidae ( Astrida, birds, and in their nests.Bird chiggersusually re- Aureliania, Neoboydaia),and Rhinonyssidae(Rhin- main attachedfor three to four daysat the thighs, oedus,Sternostoma). The Cloacaridaeis primarily a anus, or under the wings. family of turtle cloaca mites. One genus is a sub- SubcutaneousMites. The Hypoderatidae(Gyps0dec- cutaneous small parasite and one genus tes,Neottialges, and Tytodecteson hawks,kestrels, and is an owl parasite. Pneumophagusis known only vultures;Hypodectes, Neottialges, Neotytodectes, and Tyto- from two dozen individuals from the trachea and decteson owls) are subcutaneous bird mites as bronchi of a Great Horned Owl in Michigan (Fain nymphs.Nonfeeding adults lay eggsin birds' nests. and Smiley1989). Among the ereynetidmites, Au- Nymphscolonize nestlings and adultsand liveon the relianiais known only from Barn Owls (Tyt0 alba), surface of breast and abdominal muscles, in fat tissue Astridais known from owls and Caprimulgiformes, and, occasionally,in respiratory and circulatory and Neoboydaiais known from several orders of tracts. Lacking a mouth, nutrients are absorbed birds. Rhinoedusis restrictedto owls,each species through the skin. Significant pathological effects parasitizing one or several owl species. Philips from these mites remain unproven, however.Most (1993) photographedRhinoecius in the nasalcavity speciesof hypoderatidmites have limited host rang- of a Boreal Owl. The genus Sternostomaparasitizes es,but a significantnumber of unusualone-time host many bird orders but most speciesare restrictedto SEPTEMBER 2000 PARASITIC MITES OF RAPTORS 213 one or several host species.Sternostoma tracheacol- Pages221-226 in R.L. Wenzel and V.J. Tipton. lEDS.I, um, the canary lung mite, parasitizes Ectoparasitesof . Field Mus. Nat. Hist., Chi- cago, IL U.S.A. and parrots, infiltrating the lung sacsand causing BUCHHOLZ,R. 1869. Bermerkungen uber die Arten der mortality, but the other speciesremain in the nares Gattung DermaleichusKoch. Dresden, . and seem to do minimal damage to their hosts. BUNN, D.S., R.B. WARBURTON, AND R.D.S. WILSON. 1982. Fleay (1968) suggestedthat Kytoditesnudus (the air The Barn Owl. Books, Vermillion, SD U.S.A sac mite, Kytoditidae) may occur in Ninox strenua, BUTENKO,O.M. 1971. A summaryof the knowledgeof the nasal mites of birds of the Okskii Reserve Area basedon a report by a veterinarianwho suspected OkskiiGosud. Zapov. Tr. 8:204-223. (In Russian). its presence,but did not find it. This mite feeds on 1976. New speciesof rhinonyssidmites (Gama- serous secretions in the air sacs of and soidea,Rhinonyssidae), parasitic in owls.Parazitolog•ya turkeysand has not been found in owls. 10:303-309. (In Russian). , K.T. YURLOV AND N.M. STOLBOV. 1972ß Distribu- ACKNOWLEDGMENTS tion of avian mites of the Family RhinonyssidaeGa- Funding for this project was provided in part by the masoideaon hosts.Pages 370-372 in A.I. Cherepanov Babson College Board of Research.Most of the new host [ED.], Trans-continental connections of migratory records resulted from my researchwhile Visiting Scientist birds and their role in the distribution of arboviruses. at The Raptor Center of the Universityof Minnesota, and Publ. House "Nauka," Novosibirsk, U.S.S.R. I am extremely grateful to G. Duke, P. Redig, M. Martell, BOTTI•}•, W. ^NDV. CE}•N•Z.1974. Phoresiebei Hippo- D. Rose, W. Crawford, and all the other individuals who bosciden (Diptera) von Saugetieren und Vogeln •n assisted me while I was there. David Ellis contributed der Schweiz. Bull, Soc. Entomol. Suisse 47:319-326. some valuable literature. CANESTRINI, G. AND P. KRAMER. 1899. Demodicidae und Sarcoptidae.Das Tierreich7:1-193. LITERATURE CITED CASTO,S.D. 1976. Host recordsand observationsof qmll AMAR•L,V. 1962. Sternostomaaugei n. sp. (Acarina: Rhi- mites (Acarina: Syringophilidae) from birds. Southwest. Entomol. 1:155-160. nonyssidaeVitz., 1935) and the identification of the species Rhinoeciusbisetosus Strandtmann, 1952 and CE}•N•Z,V. 1967. Camlogode la fauna Cubana--XX--Lista Rhinoeciusgrandis Strandtmann, 1952. Arq. Inst. Biol. de los Acaros parasitosde Aves reportadas de Cuba. 29:69-81. Museo "Felipe Poey" de la Academia de Cienciasde ANONYMOUS. 1963. Mites. Hawk Chalk 3:40. Cuba, Trabajos de Divulgacion No. 45. ASH, J. AND T.E. HUGHES.1952. Further records of the . 1969.The hypopiof hypoderidae() genera MicrolichusTrt. et Neum. and MyialgesSerg. et parasitizingCuban birds. Folia Parasitol. 16:271-274. Trt. Ann. Mag. Nat. Hist. 12:753-759. .1973. Kramerellamrciaki, sp. n., a newfeather mite AT'toO, W.T. AND N.L. BRAASCH.1966. The species ( Pterolichidae) from the Teng- genus Proctophyllodes(Sarcoptiformes: Proctophyllodi- malm's Owl. Acta Fac. Rerum Nat. Univ. Comenianae dae. Bull. Univ. Ne• State Mus. 5:1-354. Zool. 19:9-12. --AND J.R. PHILIPS.1984. The feather mite genus . 1977. The feather mite speciesnew for Czechos- Neopetitota(Pterolichoidea, Kramerellidae). J. Med.En- lovakia. Folia Parasitol. 24:62. tomol. 21:409-411. --AND J. WIESNE}•.1992. Kramerellaglaucidii spec. BANKS, N. 1915. The Acarina or mites. A review of the nov. (, Sarcoptiformes, Kramerellidae)--eine group for the use of economic entomologists.USDA neue Milbenart von Sperlingskauz.Anz. Vez.Thueringer Rep. No. 108. Ornithol. 1:85-87. BECH•mI•, MJ. AND J.K. SCHMUTZ. 1995. Ferruginous CHEK•,R.A. 1972. Birdsof the CheranganiMountain for- Hawk (Buteoregalis). In A. Poole and F. Gill [EDS.], estsand their parasites.E. Aft. Wildl.J. 10:245-249. The birds of .No. 172. The Academy ß 1978. Recordsof birds and their parasitesfrom of Natural Sciences,Philadelphia, and The American the Cherangani Mountains, . East Aft. Wildl. J. Ornithologists' Union, ,DC U.S.A. 16:61-64. BEQUAERT,J. 1953. The Hippoboscidaeor louse-flies(Dip- CooPEa,J.E. 1978. Veterinaryaspects of captivebirds of tera) of mammalsand birds.Part I. Structure,physiology prey. StandfastPress, Saul, Gloucestershire,U.K. and naturalhistory. Entomol. Am. 32:1-209. . 1985. Veterinary aspectsof captivebirds of prey, BREGETOVA,N.G. 1964. Some problems of evolution of 2nd ed. Standfast Press, Saul, Gloucestershire, U.K the rhinonyssidmites. First International Congressof COOREMAN,J. 1946. Rhinoedusoti n. gen., n. sp. (Acarien: Parasitology.Nauka: 1-7. (In Russian). Rhinonyssinae). Bull. Mus. Hist. Nat. Belg.22:1-4. .1965. A new speciesof parasiticmite of the genus DOMROW,R. 1965. New laelapid nasal mites from Austra- RhinoeciusCooreman (:Rhinonyssidae). lian birds. Acarologia7:430-460. Entomol. Rev. 44:116-117. 1966. 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Notes sur quelquesepider- 109-110. moptines des genres Microlichuset Myialges(Acarina: EWING,H.E. 1925. New mites of the family Dermanyssi- Sarcoptiformes).Acarologia 13:532-538ß dae (Acarina). Entomol. News 36:18-22. --AND P. GROOTAERT. 1996. Observations sur des 1936. A short synopsisof the North American Acariens (Acari: Epidermoptidae) parasites speciesof the mite genus Dermanyssus.Proc. Entomol. d'Ornithomyiaavicularia (L.) (Diptera: Hippoboscl- Soc. Wash. 38:47-54. dae) de Belgique. Bull. Ann. Soc.R. BelgeEntomol. 132. FAIN, A. 1956a. Nouvelles observations sur les acariens de 183-186ß la famille Speleognathidaeparasites des fosses nasales -- ANDD.E. JOHNSTON.1966. Nouveaux acariensna- chez les batraciens, les oiseaux et les mammif•res. sicolesd'oiseaux Nord-Americains (Acari: Rhinonys- Ann. Parasitol.Hum. Comp.31:643-662. sidae). Bull. Soc.R. Zool. Anvers 38:25-41ß 1956b. Les acariensde la famille Speleognathi- --^NI) R.L. SMILEYß1989. A new cloacarid mite (Ac- dae Womersleyau Ruanda-Urundi Congo Beige). Rev. ari: Cloacaridae)from the lungs of the Great Horned Zool.Bot. Afr. 53:17-50. Owl, Bubovivginianus, from the U.S.A. Int. J. Acarol --. 1956c. Note complfimentairesur les Rhinonyssi- 15:111-115. dae au Ruanda-Urundi. Rev.Zool. Bot. Afr. 53:392-398. --, J. GAUDAND J. PHILIPSß1987. Notes sur trois es- 1957. Les acariensdes familiesEpidermoptidae p•ces d'Epidermoptidae (Acari, Astigmata) dont et Rhinonyssidaeparasites des fosses nasales d'oiseaux deux nouvelles.Acarologia 28:359-366ß SEPTEMBER2000 PARASITIC MITES OF RAPTORS 215

FLEAY,D. 1968. Nightwatchmenof bush and plain. Tap- GIEBEL,C. 1861. Die Milbenarten der Gattung Hypoderas linger Publ., , NY U.S.A. Nitzsch. Zeit. Gesell. Naturw. 18:438-444. FRITSCH,W. 1954. Die Milbengattung HavpyrhynchusMeg- --. 1871. Ueber einige Milben. Z. Gesell.Naturw. 38: nin, 1878. Zool. Anz. 152:177-198. 29-32. FURMAN,D.F. ANDI.B. TARSHIS.1953. Mites of the genera GOFF,M.L. 1979ßThe genus Odontacarus(Acari: Trom- Myialgesand Microlichus(Acarina: Epidermoptidae) biculidae) in New , with descriptionsof four from avian and insecthosts. J. Parasitol.39:70-78. new species.J. Med. Entomol.15:143-154. GAUD,J. 1958. Acariensplumicoles (Analgesoidea) par- GUTII•RREZ,RJ., A.B. FRANKLIN,AND W.S. LAHAYE.1995. asitesdes oiseaux du Maroc II. Analgesidae.Bull. Soc. Spotted Owl ( occidentalis).In A. Poole and F. Gill Sci.Nat. Phys.Maroc 38:27-49. [EDs.], The birds of North America. No. 179. The --. 1966. Ancyralgescometus n.g.n. sp. 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Parasitesdes Ac- HERMAN,C.M. 1945. Hippoboscid flies as parasitesof cipitridae et Sagittariidae (Acariens Gabuciniidae). game in California. Calif Game31:16-25. Rev.Zool. Aft. 97:737-766. HILL, D.S., N. WILSON AND G.B. COP•ET. 1967. Mites as- 1988. Acariens sarcoptiformesplumicoles des sociatedwith British speciesof Ornithomya.J. Med.En- oiseauxFalconiformes d'Afrique. III. Acariens,autres tomol. 4:102-122. que les Gabuciniidae,parasites des .Rev. HOUSTON, C.S., D.G. SMITH, AND C. ROHNER. 1998. Great Zool.Aft. 102:93-102. Horned Owl (Bubovirginianus). InA. Poole and F. G•11 --AND W.T. ATYEO. 1974. Gabuciniidae, famille nou- [EDs.], The birds of North America. No. 372. The velle de Sarcoptiformesplumicoles. Acarologia 16:522- Academyof Natural Sciences,Philadelphia, PA and 561. The American Ornithologists' Union, Washington, -- AND --. 1976. Ascouracarinae, n. sub fam. des DC U.S.A. Syringobiidae,Sarcoptiformes plumicoles. Acarologia HUNTER,J.E., RJ. GUTII•RREZ,A.B. FRANKLIN,AND D. 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Appendix 1. A list of the parasiticmites of the Falconiformes.

RAPTOR MITE REFERENCES

Family Accipitridae Epidermoptidaea skin Herman 1945 Knemidocoptessp. skin Cooper 1978, 1985 ( badius) Coraciacavussp.b feathers McClure and Ratanaworabhan 1971 Hieracolichus nisi feathers Dubinin 1956 Schoutedenocoptesaquilae nasalcavity Fain 1977 B•colored Hawk (Acdpiterbicol- mitesa skin Bequaert 1953, Maa 1969 or) Chestnut-FlankedSparrowhawk Aetacavus andrei feathers Gaud 1983b (Accipiter castanilius) Cooper's Hawk (Accipitercoop- Neonyssussp.b nasalcavity Peters 1936 e•/) Syringophilidaen.g. quills Philips, present work Brown Goshawk (Accipiterfasda-Speleognathopsis accipitris nasal cavity Domrow 1969, Domrow 1991 tus) (Accipiter mites a skin Walter 1989 gentilis) Pseudalloptinusaquilinus •,c feathers Nordberg 1936, Niethammer 1938, Dubinin 1956 Hieracolichus nisi feathers Niethammer 1938, Dubinin 1956 Hieracolichusn. sp. feathers Philips, presentwork Neottialgesvitzthumi subcutaneous Vitzthum 1934, Fain 1967 Ornithonyssus.sylviarum skin/nest Cooper 1978, 1985 Slaty-mantledSparrowhawk (Ac- mites • skin Maa 1966 cipiterluteoschistaceus) Black Goshawk (Accipitermelan- Schoutedenocoptesaquilae nasal cavity Fain 1956c, 1957 oteucus) (Accipiter Aetaecarus andrei feathers Gaud 1983b minullus) Schoutedenocoptesaquilae nasalcavity Fain 1956c, 1957 EurasianSparrowhawk (Accipi- mites a skin Newton 1979, Walter 1989 ter nisus) Cnemidocoptesspp. skin Malley and Whitbread 1996 Dermoglyphuselongatus • quills Dubinin 1956 Dubinia accipitrina feathers Niethammer 1938 Hieracolichus nisi feathers Canestrini and Kramer 1899, Nie- thammer 1938, Radford 1953, Dubinin 1956, Vasilev 1961, 1962, Shumilo et al. 1973, Mi- ronov 1997 Megniniasp. b Niethammer 1938 Microlichus avus • skin Walter 1989 Microlichussp> b skin Ash and Hughes 1952 Myialges?sp. • skin Ash and Hughes 1952 Myialgesuncus • skin Walter 1989 Ornithonyssusbursa skin/nest Anon. 1963 Grey Goshawk (Accipiternovae- mites • skin Maa 1966, 1969 hollandiae) SEPTEMBER 2000 PARASITIC MITES OF RAPTORS 219

Appendix 1. Continued.

RAPTOR MITE HABITAT REFERENCES

Ovampo Sparrowhawk(Accipiter Aposolenidia anomogonima feathers Gaud and Atyeo 1974 ovampensis) Schoutedenocoptesaquilae nasalcavity Fain 1956c, 1957 Sharp-shinnedHawk (Accipiter mitesa skin Bequaert 1953, Fain 1965, Maa striatus) 1969 Dermanyssusamericanus skin/nest Philips 1993 Ornithonyssusiheringi skin/nest Dusb/tbekand Cern• 1971 Syringophiliaen.g. quills Philips, present work African Goshawk (Accipitertachi- Aetacarus andrei feathers Gaud 1983b ro) Myialgesasturi skin Fain 1965 Myialgesfalconis skin Fain 1965 (Accipiter tri- feather mites feathers Maa and Kuo 1965 virgatus) mitesa skin Maa 1966 (Accipitervirgatus) feather mites feathers Maa and Kuo 1965 mites a skin Maa 1966 Hieracolichus nisi feathers Dubinin 1956 Ornithonyssusbursa skin/nest McClure and Ratanaworabhan 1971 Golden Eagle (Aquila chrysae- Ha50yrhynchussp. skin Schulz 1990 tos) Pseudalloptinusaquilinus feathers Trouessart 1884, L6nnfors 1930, Radford 1953, 1958, Dubinin 1956 •upicaprae b mammals Valentincicand Ku•ej 1989 Greater Spotted Eagle (Aquila Pseudalloptinusaquilinus feathers Dubinin 1956 clanga) Imperial Eagle (Aquila heliaca) Pseudalloptinusaquilinus feathers Dubinin 1956 (Aquila Pseudalloptinusaquilinus feathers Radford 1953, 1958, Dubinin pomarina) 1956 African Tawny-Eagle(Aquila ra- Pseudalloptinusaquilinus feathers Dubinin 1956 pax) Pyonacarussp. quills Atyeo pers. comm. Schoutedenocoptesaquilae nasal cavity Fain 1956c, 1957, Gaud and Till 1961 Verreaux's Eagle (Aquila ver- Hieracolichusdobyi feathers Gaud 1983b reauxii) Wahlberg'sEagle (Aquilawahl- Hieracolichusdobyi feathers Gaud 1983b berg0 Grey-linedHawk (Asturinaniti- Hemicheylabispraecox feathers Trouessart1885, Gaud and Atyeo da) 1984 African Baza (Aviceda cuculo- Aetacarusavicedae feathers Gaud 1983b ides) Hieracolichusostodus feathers Gaud 1983b PacificBaza (Avicedasubcristata) Tinaminyssus epileus nasalcavity Wilson 1964, 1965 GrasshopperBuzzard ( Schoutedenocoptesaquilae nasal cavity Fain 1956b, 1957 rufipennis) Buteosp. Blankaartia allei skin/nest Wharton and Fuller 1952 Zone-tailedHawk (Buteoalbono- Eutrombicula alfreddugesi • skin/nest Philips 1978 tatus) Augur (Buteoaugur) Falconyssusbuteonis nasal cavity Fain 1956b CommonBuzzard (Buteo buteo) Pseudalloptinus aquilinus •,c feathers Nordberg1936, Niethammer 1938 Harpyrhynchustracheatus skin Fritsch 1954 Hieracolichusnisi feathers Canestrini and Kramer 1899, Rad- ford 1953, 1958 Myialgespart • skin Bfittiker and Cern• 1974 Prostigmatalarva a Bfittiker and Cern• 1974 Pseudogabuciniaintermedia feathers Gaud 1988 220 PHILIPS VOL. 34, No. 3

Appendix 1. Continued.

RAPTOR MITE HABITAT REFERENCES

GalapagosHawk (Buteogalapa- mites• skin Maa 1969 goensis) Myialgescaulotoon • skin Harmon et al. 1990, Madden and Harmon 1998 Red-tailedHawk (Buteojamai- mites• skin Bequaert 1953, Maa 1969 censis) Aetacarusn. sp. feathers Philips, present work Eutrombiculaalfreddugesi skin/nest Loomis 1956 Harpyrhynchussp. skin Philips,present work Myialgesfalconis • skin Philips and Fain 1991 Peristerophilacolumbae quills Casto1976 Schoutedenocoptesaquilae nasal cavity Fain 1956b, 1957 Syringophilidaen.g. quills Philips, present work Red-shoulderedHawk (Buteoli- Haemogamasusreidi b mammals Redington1970 neatus) Hieracolichusn. sp. feathers Kurey 1976 Pseudalloptinussp. feathers Kurey 1976 RoadsideHawk (Buteomagniros- mites a skin Maa 1969 tris) Myialgesbombycilla• skin Philipsand Fain 1991 Ptilonyssussouzai nasal cavity Pereira and Castro 1949 Broad-wingedHawk (Buteopla- Hieracolichusn. sp. feathers Philips, present work typterus) Syringophilidaen.g. quills Philips, present work FerruginousHawk (Buteoregal- mites Bechard and Schmutz 1995 is) Swainson'sHawk (Buteoswain- Hieracolichussp. feathers Kurey 1976 soni) Great Black-Hawk ( mites• skin Maa 1969 urubitinga) Eutrombiculabatatas skin/nest Brennan and Yunker 1966 Short-toedSnake Eagle (Circae- Hieracolichusnisi feathers Canestriniand Kramer 1899, Rad- tusgallicus) ford 1953, 1958, Dubinin 1956 WesternMarsh- (Circus Pseudalloptesbisubulatus b feathers Dubinin 1956 aeruginosus) Pseudogabuciniaintermedia feathers Dubinin 1956 CxnereousHarrier (Circusciner- Ingrassiinaesp. a feathers Philips and Fain 1991 eus) (Circuscy- Pseudogabuciniaintermedia feathers Dubinin 1956, Cern• 1967, Kurey aneus) 1976 Pallid Harrier (Circus macrou- mites• skin Maa 1966 rus) Aetacarusleptotrichus feathers Gaud 1983b MyialgesmacdonaldP skin Philips and Fain 1991 Montagu's Harrier (Circuspy- Hieracolichus nisi feathers Canestrini and Kramer 1899, Nie- gargus) thammer 1938, Radford 1953 Pseudogabuciniaintermedia feathers Dubinin 1956 Swallow-tailed Kite (Elanoides Aetacarussp. feathers Meyer 1995 forficatus) Macrochelessp.b litter/nest Meyer 1995 Ornithonyssusbursa skin/nest Meyer 1995 Black-wingedKite (Elanusca- Cheylabislatus feathers Gaud and Atyeo 1984 eruleus) Falconyssuselani nasal cavity Fain 1966a Neottialgeselani subcutaneous Fain 1969 Australian Black-shouldered Cheylabislatus feathers Gaud 1983a Kite (Elanus notatus) Odontacarusaustraliensis skin/nest Domrow 1966, 1991 Ornithonyssusbursa skin/nest Domrow 1966 Palm-nut Vulture (Gypohierax mites a skin Maa 1966 angolensis) Aetacarushyalothrix feathers Gaud and Mouchet 1959b, Gaud and Till 1961, Gaud 1983b Hieracolichus orthochaetus feathers Gaud and Mouchet 1959b, Gaud 1983b Myialgesn. sp.• skin Philips and Fain 1991 Pseudalloptinusafricanus feathers Gaud 1988 Pseudalloptinusodontopus feathers Gaud and Till 1961, Gaud 1988 SEPTEMBER2000 PARASITICMITES OF RAPTORS 221

Appendix 1. Continued.

RAPTOR MITE HABITAT REFERENCES

White-backedVulture (Gypsaf- Ancyralgescometus feathers Gaud 1966, 1988 ricanus) Hemicheylabissikyonemus feathers Gaud 1988 Hieracolichusafricanus feathers Guad and Mouchet 1959b, Gaud and Till 1961, Gaud 1983b Cape Griffon (Gypscoprotheres) Gypsodectes verrucosus • subcutaneous Fain 1984 A ndrolaelapspatersoni skin/nest Zumpt and Till 1956, Till 1963 Ramogabuciniadoleosikya feathers Gaud and Atyeo 1974, Gaud 1983b Eurasian Griffon (Gypsfulvus) Gypsodectesvulturis subcutaneous Dubinin 1953, 1956, Fain 1967 Haliaeetussp. Pseudalloptinusaquilinus feathers Radford 1953, 1958 White-tailed Eagle (Haliaeetus Aetacarusphylloproctus feathers Dubinin 1956 albicilla) Meg'niniapicimajoris b Niethammer 1938 Pseudalloptinusaquilinus feathers Dubinin 1956, Vasilev 1961 Pterolichus obtususb feathers Niethammer 1938 ( Haliaeetusleucoce- Analgesidaeb Spencer 1941 phalus) Pseudalloptinusaquilinus feathers Dubinin 1956, Vasilev 1961 Syringophilidaen.g. quills Philips, present work White-bellied Fish-Eagle(Hal- Aetacatusphylloproctus feathers Canestrini and Kramer 1899, iaeetusleucogaster) Gaud and Petitot 1948b, Rad- ford 1953, 1958, Gaud and Atyeo 1974 Pseudalloptinusodontopus feathers Gaud and Till 1961, Gaud 1988 Pallas'sSea-Eagle (Haliaeetus Aetacarusphylloproctus feathers Dubinin 1956 leucoryphus) Pseudalloptinusaquilinus feathers Dubinin 1956, Vasilev 1961 Pseudalloptinusodontopus feathers Gaud and Till 1961, Gaud 1988 Steller's Sea-Eagle(Haliaeetus Aetacatusphylloproctus feathers Dubinin 1956 pelagicus) Pseudalloptinusaquilinus feathers Dubinin 1956, Vasilev 1961 Solomon Fish-Eagle(Haliaeetus Ornithogastiariversi skin/nest Wharton and Hardcastle 1946, sanfordi) Goff 1979 Pseudalloptinusodontopus feathers Gaud and Till 1961, Gaud 1988 African Fish-Eagle(Haliaeetus Aetacatuspuylaerti feathers Gaud 1983b vocifer) Hemifreyanamarginata b feathers Gaud and Till 1961 Hieracolichusdobyi feathers Gaud and Mouchet 1959b Pseudalloptinusaquilinus feathers Gaud 1988 MadagascarFish-Eagle (Haliaee- Aetacatussp. feathers Atyeo pers. comm. tus vociferoides) Brahminy Kite ( Haliastur indus) Aetacarus haliasturi feathers M6gnin and Trouessart 1884d, Radford 1953, 1958, Dubinin 1956 mites a skin Maa 1969 Pseudalloptinusmilvulinus feathers Radford 1953, 1958, Dubinin 1956 Harpy Eagle ( Harpia harpyja) Hieracolichus hitundo feathers Radford 1953, 1958 SolitaryEagle ( Temnalgessp. a,b feathers Philips and Fain 1991 solitarius) Ayres'sHawk-Eagle (Hieraaetus A etacatuseu rychaetus feathers Gaud and Mouchet 1959b, Gaud ayresii) and Till 1961, Gaud 1983b Hieracolichus orthochaetus feathers Gaud and Till 1961 Bonelli's Eagle ( Hieraaetusfas- Pseudalloptinusaquilinus feathers Gaud 1983b ciatus) ( Hieraaetuspenna- hypopusmites skin Hamerton 1941 tus) Pseudalloptinusaquilinus feathers Dubinin 1956 Grey-headedFish-Eagle (Ichthy- Aetacatusphylloproctus feathers Radford 1958 ophagaichthyaetus) MississippiKtie ( Ictinia mississip-Aetacarus sp. feathers Kurey 1976 piensis) 222 PHILIPS VOL. 34, No. 3

Appendix 1. Continued.

RAPTOR MITE HABITAT REFERENCES

Lizard Buzzard ( Kaupifalcomon- Aetacarus andrei feathers Gaud 1983b ogrammicus) Myialgesanchora • skin Fain 1965 Schoutedenocoptesaquilae nasal cavity Fain 1959a ( Leucopternisalbi- Temnalgessp. a,b feathers Philipsand Fain 1991 coilis) Long-crestedEagle (Lophaetus Falconyssusbuteonis nasal cavity Fain 1956b occipitalis) Pseudalloptinusmilvulinus feathers Gaud and Till 1961, Gaud 1988 Pseudogabuciniaintermedia feathers Gaud 1988 Hawk ( Macheiramphusalci- mites a skin Maa 1966 nits) Myialgesmacdonald• skin Philips and Fain 1991 Gabar Goshawk ( Melieraxgabar) Schoutedenocoptesaquilae nasal cavity Fain 1956b, 1957 Dark Chanting-Goshawk(Meli- Aposolenidiaanomogonima feathers Gaud and Atyeo 1974 erax metabates) ( Milvus migrans) Hemicheylabissp. feathers Atyeo pers. comm. Aetacarushyalothrix feathers Gaud and Mouchet 1959b, Gaud and Till 1961 Aetacarus milvi feathers Gaud 1983b, D'Souza et al. 1986 Hieracolichus nisi feathers Dubinin 1956 Pseudalloptinusmilvulinus feathers Dubinin 1956 Pyonacaruspolysarcus feathers Gaud and Atyeo 1976, Gaud 1988 Schoutedenocoptesaquilae feathers Fain 1956b, 1957 Tinaminyssuscolumbae feathers Fain 1957 ( Milvus milvus) Pseudalloptinusmilvulinus feathers Trouessart 1884, Radford 1953, 1958, Dubinin 1956 (Necrosyrtes Hemicheylabissikyonemus feathers Gaud 1988 monachus) Hieracolichus monachi feathers Gaud and Mouchet 1959b, Gaud and Till 1961, Gaud 1983b Pseudalloptinusglossocercus feathers Gaud 1988 Ramogabuciniafurciseta feathers Gaud 1983b Osprey (Pandion haliaetus) mites skin Bequaert 1953 Analloptesbuettikeri feathers Mironov 1997 Analloptessp. feathers Gaud and Atyeo 1979, Gaud 1983a, Miller et al. 1997 Bonnetellafusca feathers Buchholz 1869, Canestrini and Kramer 1899, L6nnfors 1930, Niethammer 1938, Spencer 1941, Mrciak and Brander 1967, McClure and Ratanawor- abhan 1971, Kurey 1976, Gaud 1983a, Mironov 1991, 1997, Miller et al. 1997 Myialgescaulotoon a skin Philips and Fain 1991 Harris' Hawk ( Parabuteouni- Neoschoengastiaamericana skin/nest Loomis and Crossley1963 cinctus) Pseudalloptinussp. feathers Atyeo pers. comm. European Honey-Buzzard (Per- feather mites feathers Maa and Kuo 1965 nis apivorus) Hieracolichus nisi feathers Canestrini and Kramer 1899, Nie- thammer 1938, Radford 1953 Hieracolichus ramosus feathers Gaud and Mouchet 1959b, Gaud and Till 1961, Gaud 1983b African Harrier-Hawk (Polyboro-Hieracolichus similis feathers Gaud and Mouchet 1959b, Gaud •destypus) and Till 1961, Gaud 1983b Martial Eagle (Polemaetus bellico- Hieracolichus dobyi feathers Gaud 1983b SitS) Pseudalloptinusaquilinus feathers White-backed Vulture (Pseudo- Dermanyssusgallinae skin/nest gypsafricanus) SEPTEMBER2000 PARASITICMITES OF RAPTORS 223

Appendix 1. Continued.

RAPTOR MITE HABITAT REFERENCES

SnailKite (Rostrhamussociabilis) Ornithonyssus lrursa skin/nest Philipset al. 1976,Sykes and For- rester 1983, Sykeset al. 1995 CrestedSerpent-Eagle ( SpilornisCoraciacavus cucuh •' feathers Radford 1958 &eela) ChangeableHawk-Eagle (Spi- Coraciacarussp. b feathers McClure and Ratanaworabhan zaetus cirrhatus) 1971 Crowned Hawk-Eagle(Stephan- Aetacarushyalothrix feathers Gaud and Mouchet 1959b, Gaud oaetuscoronatus) and Till 1961 Hieracolichusdobyi feathers Gaud and Mouchet 1959b, Gaud and Till 1961, Gaud 1983b Pseudalloptinusodontopus feathers Gaud and Till 1961, Gaud 1988 Bateleur (Terath0piusecaudatus) Hieracolichus dobyi feathers Gaud 1983b White-headed Vulture (Trig0n0- Hieracolichusmonachi feathers Gaud 1983b cepsoccipitalis) Long-tailed Hawk ( Urotriorchis Myialgescaulotoon a skin Philips and Fain 1991 macrourus) Pseudogabuciniasp. a feathers Philips and Fain 1991 Family Cathartidae Vulture (Cathartesaura) mites a skin Bequaert 1953, Maa 1969 A ncyralgessp. feathers Kurey 1976 Hieracolichussp. feathers Peters 1936, Kurey 1976 Ornithonyssusbursa skin/nest Peters 1936 Ptilonyssusohioensis nasal cavity Fain and Johnston 1966 Lesser Yellow-headed Vulture Hieracolichussp. feathers Atyeo pers. comm. ( Cathartes burrovianus) Black Vulture (C0ragypsatratus) mites a skin Bequaert 1953 Eutrombiculaalfreddugesi skin/nest Wharton and Fuller 1952 Hieracolichussp. feathers Kurey 1976 Histiogaste••,b trees Philips and Fain 1991 Ptilonyssusdonatoi nasal cavity Pereira and Castro 1949 Sancassaniasp. a,b litter/nest Philips and Fain 1991 Andean Condor (Vultur gry- Hieracolichusspp. feathers Atyeo pers. comm. phus) Family Falcosp. mites a skin Bequaert 1953, Fain 1965 Ptilonyssuscerchneis nasal cavity Bregetova1964 Saker/Peregrine hybrids Dermanyssusgallinae skin/nest Malley and Whitbread 1996 Grey Kestrel (Falcoardosiaceus) Pseudalloptes falconis feathers Gaud 1983a Brown (Falcoberigora) Boydaiafalconis nasal cavity Domrow 1991 Leptotrombidiumnissani skin/nest Domrow and Lester 1985 Ptilonyssuscerchneis nasal cavity Domrow 1965, 1967, 1969 Odontacarus nadchatrami skin/nest Goff 1979 Lanner Falcon (Falco biarmicus) Pseudalloptesfalconis feathers Gaud 1983a Pseudogalruciniamicrodisca b feathers Gaud 1983a Australian Kestrel (Falco cenchro-Leptotrombidium nissani skin/nest Domrow 1974, Domrow and Les- ides) ter 1985 Odontacarus australiensis skin/nest Domrow 1966, 1991 Ornithonyssuslrursa skin/nest Domrow 1977 Ornithonyssussylviarum skin/nest Domrow 1987 Ptilonyssuscerchneis nasal cavity Domrow 1965, 1967, 1969 Red-necked Falcon (Falco chic- Pseudalloptesfalconis feathers Gaud 1983a quera) 224 PHILIPS VOL. 34, No. 3

Appendix 1. Continued.

RAPTOR MITE HABITAT REFERENCES

Saker Falcon (Falco cherrug) Dermanyssussp. skin/nest Wheeldon pers. comm. Merlin (Falco columbarius) Dubininiaaccipitrina feathers Vasilev1958, Cernp 1967, Gaud 1980, 1983a Gabuciniidae feathers Kurey 1976 Hieracolichussp. feathers Gaud 1983a Microlichus avus • skin Hill et al. 1967 African (Fatcocuvieri) Boydaiafalconis nasal cavity Fain 1956a Eleonora's Falcon (Falco eleono- Acarina sp. Wink et al. 1979 rae) Kramerellamajo• b feathers M•gnin and Trouessart1884a Pseudogabuciniaintermedia feathers Dubinin 1953, 1956, Radford 1958, Gaud 1983a (Falco naumanni) Dubininia accipitrina feathers Gaud and Petitot 1948a, Gaud 1958, 1983a (Fatcoperegri- Aetacarus?sp. feathers Kurey 1976 Glaucalgesattenuatus • Niethammer 1938 Hieracolichus nisi feathers Dubinin 1956, Cernp 1967, Gaud 1983a Pandalurastrigisotz •' Niethammer 1938 Pseudalloptinussp. feathers Kurey 1976, Gaud 1983a Pseudogabuciniaintermedi• feathers Nordberg 1936, Niethammer 1938, Dubinin 1953, Radford 1958, Gaud 1983a Oriental Hobby (Falcoseverus) Ornithogastiariversi skin/nest Wharton and Hardcastle 1946, Coif 1980 American Kestrel (Falcosparver- Dubininia sp.c feathers Philips 1990 zus) Tytodectescerchneis • subcutaneous Philipsand Dindal 1979, Philips 1990 Blankaartia velascoi skin/nest Wharton and Fuller 1952 Boydaiafalconis nasal cavity Pence and Casto 1976 Ptilonyssuscerchneis nasal cavity Strandtmann 1962 EurasianHobby (Falcosubbuteo) Microlichus falco skin Fain and Gaud 1972, Fain et al. 1987, Fain and Grootaert 1996 Neottialgesheteropus subcutaneous Giebel 1861, 1871, Fain 1967 Pseudalloptinusminor feathers M6gnin and Trouessart1884b, Canestrini and Kramer 1899, Dubinin 1956, Gaud 1983a Pseudogabuciniaintermedia skin Niethammer 1938, Radford 1953, 1958, Vasilev 1961, Gaud 1983a (Falcotinnun- Boydaiafalconis nasal cavity Fain 1963a culus) Cheyletiellasp.a Niethammer 1938 Dubininia accipitrina feathers Trouessart 1885, Canestrini and Kramer 1899, Niethammer 1938, Radford 1953, 1958, Gaud 1958, 1983a MeKniniasp. b Neithammer 1938 Myialgessp. nr. par• skin Philips and Fain 1991 Protolichus lunula b feathers Vasilev 1961 Pseudalloptesfalconis nasal cavity Gaud 1983a Ptilonyssuscerchneis nasalcavity Fain 1957 Tytodectescerchneis subcutaneous Fain 1966b Tytodectes falconis subcutaneous Fain 1969 Red-looted Falcon (Falcovesper- Hieracolichus nisi feathers Shumilo et al. 1973, Gaud 1983a tznus) SEPTEMBER2000 PARASITICMITES OF RAPTORS 225

Appendix 1. Continued.

RAPTOR MITE HABITAT REFERENCES Barred Kestrel(Falco zoniven- Aetacarussp. feathers Gaud 1983a tris) BarredForest-Falcon (Micrastur Ingrassiinae sp. a feathers Philipsand Fain 1991 ruficollis) CollaredFalconet (Microhierax Coraciacarussp. b feathers McClure and Ratanaworabhan caerulescens) 1971 Philippine Falconet (Microhier- mitesa skin Maa 1969 ax erythrogenys) Yellow-headedCaracara (Milva- mitesa skin Bequaert1953 go chimachima) Ptilonyssussouzai nasalcavity Pereiraand Castro1949 Crested Caracara (Polyborus Eutrombiculabatatas skin/nest Brennan and Yunker 1966 plancus) Hieracolichussp. feathers Kurey1976 Family Sagittariidae Secretary-bird(Sagittarius ser- Aetacaruslaurencei feathers Gaud 1983b pentarius) These miteswere found on louseflies(Hippoboscidae) on the bird. Theseare incorrectlyidentified or accidentalrecords. Nest record. Cheyletiellaisnow restricted to mammalparasites; there are four other genera of birdparasites in the Family Cheyletiellidae.

Appendix 2. A list of the parasiticmites of the Strigiformes.

OWL MITE HABITAT REFERENCES

Strigiformes Dermanyssusgallinae skin/nest Pfister 1991 Rhinonyssidae nasalcavity Butenko et al. 1972 Family Strigidae Glaucalgesattenuatus feathers Radford 1953, 1958 Kramerella lunulata feathers Radford 1953, 1958 Kramerellalyra feathers Radford 1953, 1958 Kramerellamajor feathers Radford 1953, 1958 Northern Saw-whetOwl (Aego-Dermanyssus americanus skin/nest Philips 1990 lius acadicus) Petitotasp. feathers Philips, present work Boreal Owl (Aegoliusfunereus)Glaucalges attenuatus feathers Niethammer 1938 Harpyrhynchusn. sp. skin Philips 1993 Kramerella lunulate • feathers Niethammer 1938, Dubinin 1953, Radford 1958 Kramerellamajo• ,b feathers M6gnin and Trouessart 1884a, Nordberg 1936, Dubinin 1953, Radford 1958 Kramerella mrciaki feathers Cerm) 1973, Mironov 1997 Kramerellan. sp. feathers Philips, present work Mesalgoidespicimajoris • feathers L6nnfors 1937 Paralgessp. a quills Philips 1993 Passeroptesn. sp. skin Philips 1990 Petitota aluconis feathers L6nnfors 1937 Petitotahaengii feathers Mironov 1997 Rhinoeciusaegolii nasalcavity Butenko1971, Philips 1993 226 PHILI?S VOL. 34, No. 3

Appendix 2. Continued.

OWL MITE HABITAT REFERENCES

Marsh Owl (Asiocapensis) Dermonotonparallelus feathers Gaud and Mouchet 1959b, Gaud and Till 1961, Gaud 1980 Kramerella oti feathers Gaud 1980 Pandalurastrigisoti feathers Gaud 1980 Rhinoeciusafricanus nasal cavity Zumpt and Patterson 1951, Zumpt and Till 1961 Short-earedOwl (Asioflam- mites c skin Maa 1966 Dermonotonsp. feathers Dubinin 1956 Glaucalgesattenuatus feathers Buchholz 1869 Kramerellaflammei feathers Ltnnfors 1937 Kramerellalyra feathers M6gnin and Trouessart1884a, Rad- ford 1958 Kramerellamajor feathers M6gnin and Trouessart1884a, Rad- ford 1958 Kramerella oti feathers Vasyukovaet al. 1996 Kramerellasp. feathers Kurey 1976 Leptotrombidiumakamushi skin/nest Wharton and Fuller 1952 Microlichus trudicola • skin Fain et al. 1987 Pandalurastrigisoti feathers Rothschildand Clay 1952 Petitota aluconis feathers Gaud 1980 Proctophyllodespolyxenus • feathers Atyeo and Braasch1966 Rhinoeciusalifanovi nasal cavity Butenko 1976 Long-eared Owl (Asiootus) chiggers(Trombiculidae) skin/nest Maa and Kuo 1965 mites c skin Walter 1989 Dermanyssusamericanus skin/nest Moss 1978 Dermanyssushiru ndinis skin/nest Kutzer et al. 1982 Dermonotonparallelus feathers Gaud 1980 Eulaelapsstabularis • mammals Kutzer et al. 1982 Glaucalgesattenuatus feathers Buchholz 1869, Canestrini and Kra- mer 1899, Radford 1953, 1958 Glycyphagusdomesticus •,c nest Bfittiker and Cern• 1974 Harpyrhynchusasio skin Fain 1972 Kramerellalyra feathers Mtgnin and Trouessart1884a, Rad- ford 1958 Kramerella oti feathers Ltnnfors 1937, Radford 1958, Vasi- lev 1959, Cern• 1977, Gaud 1980 Kramerellasp. feathers Kurey 1976 Microlichus avus • skin Ash and Hughes 1952 Microlichus charadricola • skin Bfittiker and Cern• 1974 Myialgesmacdonaldi c skin Hill et al. 1967 Myialgesnudus skin Fain 1965, Fain and Grootaert 1996 Neotromln'culalipovskyi skin/nest Loomis 1956 Pandalurastrigisoti feathers Buchholz 1869, Canestrini and Kra- mer 1899, Radford 1953, 1958, Kurey 1976, Cern• 1977 Myialgesuncus • skin Ash and Hughes 1952, Fain 1965 Rhinoedusbrikinboricus nasal cavity Butenko 1976 Rhinoeciusoti nasal cavity Cooreman 1946 Strelkoviacarus critesi• skin Hill et al. 1967 Sternostomastrigitis nasal cavity Butenko 1976 SW•'TEMBER2000 PARASITIC MITES Or R•TO• 227

Appendix 2. Continued.

OWL MITE HABITAT REFERENCES

Little Owl (Athene noctua) Neotrombiculaautumnalis skin/nest Koptzevet al. 1961 Glaucalgesattenuatus feathers Gaud 1958, 1980 Kramerella lunulata feathers Hailer 1878, Niethammer 1938, Du- binin 1953, Radford 1958, Gaud 1980 Ornithogastiaariadnae skin/nest Hushcha 1982 Pandalurastrigisoti feathers Gaud 1958, 1980 Rhinoeciussubbisetosus nasalcavity Bregetova1965 Bubosp. Dermanyssusgallinae skin/nest Strandtmann and Wharton 1958 Dermonoton bubonis• feathers Gaud 1980 SpottedEagle-Owl ( Buboafri- Astridacaprimulgi nasalcavity Fain 1956a, Zumpt and Till 1961 Dermonotonparallelus feathers Gaud 1980 Glaucalgesattenuatus feathers Gaud 1980 Kramerella maculata feathers Gaud 1980 Pandalurastrigisoti feathers Gaud 1980 Rhinoedusbuboensis nasal cavity Fain 1958, 1959b, 1960, Zumpt and Till 1961, Domrow 1969 EurasianEagle-Owl (Bubo mites c skin Walter 1989 bub0) Dermonotonlongiventer feathers Sohn and Noh 1994 Galucalgesattenuatus feathers L6nnfors 1937, Mumcuogluand Milllet 1974 Kramerella bubonis feathers L6nnfors 1937, Dubinin 1953, Rad- ford 1958, Mrciak and Brander 1967 Kramerellamajor feathers M6gnin and Trouessart1884a, Dubi- nin 1953, Radford 1958 Pandalurastrigisoti feathers L6nnfors 1937, Mumcuogluand Milllet 1974 Petitotabubonis feathers Atyeo and Philips 1984, Sohn and Noh 1994 Verreaux's Eagle-Owl (Bubo Dermonotonapoplax feathers Gaud 1980 lacteus) Dermonotonparallelus feathers Gaud 1980 Glaucalgesattenuatus feathers Gaud 1980 Kramerella lobata feathers Gaud 1980 Kramerella lunulata • feathers Radford 1958 Pandalurastrigisoti feathers Gaud 1980 Akun Eagle-Owl ( Buboleucos- Dermonotonapoplax feathers Gaud 1980 tictus) Glaucalgesattenuatus feathers Gaud 1980 Pandalurastrigisoti feathers Gaud 1980 Fraser'sEagle-Owl ( Bubopoen- Dermonotonapoplax feathers Gaud and Mouchet 1959b, Gaud sis) and Till 1961, Gaud 1980 Glaucalgesattenuatus feathers Gaud and Mouchet 1959b, Gaud and Till 1961, Gaud 1980 Pandalurastrigisoti feathers Gaud 1980 Shelley'sEagle-Owl ( Buboshel- Glaucalgesattenuatus feathers Gaud 1980 /eyi Kramerella lobata feathers Gaud 1980 Pandalurastrigisoti feathers Gaud 1980 228 PHILIPS VOL. 34, No. 3

Appendix 2. Continued.

OWL MITE HABITAT REFERENCES

Great Horned Owl (Bubovirgi- mitesc skin Bequaert1953, Maa 1969 nianus) Blattisociuskeegan? ,c trees Philips and Fain 1991 Bubophilusascalaphus quills Philipsand Norton 1978 Dermonotonsp. d feathers Kurey 1976 Epidermoptidaec skin Herman 1945 Euschoengastianumerosa skin/nest Wrenn and Loomis 1974 Glaucalgesattenuatus feathers Atyeo and Philips 1984 Harpyrhynchussp. skin Philips, present work Knemidocoptesmutans skin Schulz et al. 1989, Schulz 1990, Mal- ley and Whitbread 1996, Houston et al. 1998 Kramerellan. sp. feathers Philipsand Norton 1978,Atyeo and Philips 1984, Philips 1993 Myialgesanchora • skin Furman and Tarshis 1953, Bequaert 1953, Fain 1965 Pandalurastrigisoti feathers Atyeo and Philips 1984 Paralgesn. sp. quills Philips 1993 Petitota bubonis feathers Atyeo and Philips 1984 Pneumophagusbubonis lungs Fain and Smiley 1989 Proctophyllodespolyxenus • feathers Atyeo and Braasch1966 Rhinoeciusgrandis nasal cavity Strandtmann 1952 Glaucidiumsp. Neotytodectesmexicanus b subcutaneousO'Connor 1981 FerruginousPygmy-Owl (Glau- Eutrombiculaalfreddugesi skin/nest Brennan and Jones 1960, Loomis cidium brasilianum) 1969 Asian Barred Owlet (Glauci- Rhinoeciusbisetosus nasal cavity Strandtmann 1960, Wilson 1968 dium cuculoides) Mountain Pygmy-Owl( Glaud- Kramerellasp. feathers Kurey 1976 dium gnoma) EurasianPygmy-Owl ( Glauci- Dermonotoneventratus feathers Canestrini and Kramer 1899, Dubi- dium passerinure) nin 1956, Radford 1953, 1958 Kramerellaglaucidii feathers Mrciak and Brander 1967, Cern9 and Wiesner 1992 Pearl-spottedOwlet (Glauci- Astridacaprimulgi nasalcavity Fain 1956a, Zumpt and Till 1961 diumpertatum) Cuban Pygmy-Owl( GlaucidiumTytodectes glauddii subcutaneousFain 1967, Cern9 1969 siju) BarkingOwl ( Ninoxconnivens) Leptotrombidium nissani skin/nest Domrow 1974, Domrow and Lester 1985 Neoschoengastiaamericana skin/nest Domrow and Lester 1985 Rhinoeciuscooremani nasal cavity Domrow 1969, 1987 Solomon Hawk-Owl (Ninox Odontacarustrisetosus skin/nest Goff 1979 jacquinoti) Morepork ( Ninoxnovaeseelan- Leptotrombidium nissani skin/nest Domrow 1974, Domrow and Lester diae) 1985 Neoschoengastiaamericana skin/nest Domrow and Lester 1985 Rhinoeduscooremani nasal cavity Domrow 1967, 1987 New Britain Hawk-Owl (Ninox mitesc skin Maa 1966 odiosa) Analgidae• skin Philips and Fain 1991 PhilippineHawk-Owl (Ninox Oconnoriainexpectata quills Gaud et al. 1989 philippensis) SnowyOwl (Nycteascandiaca) Knemidocoptessp. skin Cooper 1978, 1985 Rhinoeciusnycteae nasal cavity Butenko 1976 SEPTEMBER 2000 P•S•T•C M•TES OF R•TOP, S 229

Appendix 2. Continued.

OWL MITE HABITAT REFERENCES

Otussp. Myialgesbombydllae • skin Philips and Fain 1991 Eastern Screech-Owl (Otus mitesc skin Bequaert 1953, Fain 1965, Maa 1969 asio) Dermanyssusamericanus skin/nest Ewing 1925, 1936, Radford 1950, Evans and Till 1962, Moss 1978 Dermonotonsp. feathers Kurey 1976 Harpyrhynchussp. skin Philips 1993 Miyatrombiculacynos • skin/nest Philips 1978, Philips and Dindal 1990 Neoschoengastiaamericana skin/nest Everett et al. 1972 Syringophilidae quills Johnstonand Kethley 1973 Variable Screech-Owl (Otus Blankaartiasinnamaryi skin/nest Brennan and Yunker 1966 atricapillus) Indian Scops-Owl(Otus bakka- chiggers (Trombiculidae) skin/nest Maa and Kuo 1965 feather mites feathers Maa and Kuo 1965 Dermonotonsp. feathers McClure and Ratanaworabhan 1971 Leptotrombidiumdeliense nest/skin McClure and Ratanaworabhan 1971 Ornithonyssusbursa skin/nest McClure and Ratanaworabhan 1971 Rhinoeciuscavannus nasal cavity Wilson 1968, McClure and Ratana- worabhan 1971 Toritrombiculadensipiliata skin/nest Nadchatram 1967, Vercammen- Grandjeanand Langston1976 Tori vorca skin/nest Vercammen-Grandjean and Lang- ston 1976 White-faced Scops-Owl(Otus Pandalurastrigisoti feathers Gaud 1980 leucotis) ReddishScops-Owl (Otus rufes-Leptotrombidiu m deliense skin/nest McClure and Ratanaworabhan 1971 cens) Toritrombiculadensipiliata skin/nest McClure and Ratanaworabhan 1971 EurasianScops-Owl (Otus chiggers (Trombiculidae) skin/nest Maa and Kuo 1965 scops) feather mites feathers Maa and Kuo 1965 mites c skin Maa 1969 Dermonotonparallelilobus • feathers Radford 1953, 1958 Dermonotonparallelus feathers M6gnin and Trouessart 1884c, Ca- nestrini and Kramer 1899, Dubi- nin 1956, Gaud and Till 1961 Dermonotonsp. feathers McClure and Ratanaworabhan 1971 Kramerella lunulata feathers Dubinin 1953 Kramerellalyra feathers Dubinin 1953 Kramerellamajor feathers Shumilo et al. 1973 Leptotrombidiumdeliense skin/nest McClure and Ratanaworabhan 1971 Neoschoengastialongipes skin/nest McClure and Ratanaworabhan 1971 African Scops-Owl(Otus sene- Astridacaprimulgi nasal cavity Fain 1956a, Zumpt and Till 1961 galensis) Mountain Scops-Owl(Otus Astridacaprimulgi nasal cavity Fain 1963a spilocephalus) Dermonotonsp. feathers McClure and Ratanaworabhan 1971 Leptotrombidiumscutellare skin/nest McClure and Ratanaworabhan 1971 Neoschoengastiasp. skin/nest McClure and Ratanaworabhan 1971 Whiskered Screech-Owl (Otus Dermonotonsp. feathers Kurey 1976 trichopsis) Band-bellied Owl ( Pulsatrix me- Rhinoedusnycteae nasal cavity Amaral 1962 lanota) Vermiculated Fishing-Owl (Sco-Dermonoton parallelilobus feathers Gaud 1980 topeliabouvieri) 230 PHILIPS VOL. 34, No. 3

Appendix 2. Continued.

OWL MITE HABITAT REFERENCES

Pel'sFishing-Owl (Scotopelia Dermonotonparallelilobus feathers Gaud and Mouchet 1959b,Gaud peli) and Till 1961,Gaud 1980 Glaucalgesattenuatus feathers Gaud and Till 1961, Gaud 1980 Pandalurastrigisoti feathers Gaud 1980 Burrowing Owl ( Speotytocuni- Dermonotonsp. feathers Kurey 1976 cularia) Euschoengastoidesgntrneyi skin/nest Loomis 1956 HypodectespropuP subcutaneous Pence and Bergan 1996 Hyponeoculamontanensis skin/nest Loomis 1956 Kramerellamajor feathers M6gnin and Trouessart1884a, Dub•- nin 1953, Radford 1958 Neoschoengastiaamericana skin/nest Loomis 1956, Everett et al. 1972 Proctophyllodespolyxenus • feathers Atyeo and Braasch1966 Rhinoedusbisetosus nasal cavity Strandtmann 1952, Amaral 1962 Sternostomaaugei nasal cavity Amaral 1962 Tytodectesspeotyto subcutaneous Pence and Bergan 1996 ( Strix aluco) Dermanyssushirundinis b skin/nest Zeman and Jurik 1981 Glaucalgesattenuatus feathers Atyeo pers. comm. Kramerella aluconis feathers L6nnfors 1937, Dubinin 1953, Rad- ford 1958, Shumilo et al. 1973 Kramerellamajor feathers Dubinin 1953, Radford 1958 Pandalurastrigisoti feathers Gaud 1958, 1980 Petitota aluconis feathers Buchholz 1869, Radford 1953, 1958 Brown Wood-Owl ( Strix lepto- feather mites feathers Maa and Kuo 1965 grammica) (Strix nebulo- Dermonotonparallelilobus feathers M6gnin and Trouessart1884c, Ca- sa) nestrini and Kramer 1899, Rad- ford 1953, 1958, Dubinin 1956, Gaud and Till 1961, Gaud 1980 Kramerellaaprotuberantia feathers Philips, presentwork Pandalurastrigisoti feathers Philips, presentwork Passeroptesn. sp. skin Philips, presentwork Petitotasp. feathers Philips, presentwork SpottedOwl ( Strixocddentalis) Euschoengastia sp. (probably skin/nest Hunter et al. 1994, Gutierrez et al. numerosa) 1995 Mottled Wood-Owl ( Strix ocel- Myialgesbombydllae • skin Philips and Fain 1991 lata) ( Strix uralensis) Kramerellaaprotuberantia feathers Dubinin 1953, Radford 1958 Petitota aluconis feathers Niethammer 1938 ( Strix varia) Dermonotonsp. feathers Kurey 1976 Dermonotonparallelilobus feathers Banks 1915 Kramerellasp. feathers Kurey 1976 Ornithonyssussp. feathers Peters 1936 Pandalurastrigisoti feathers Atyeo pers. comm. Paralgesn. sp. quills Philips, presentwork Passeroptesn. sp. skin Philips,present work Rhinoeduscooremani nasal cavity Strandtmann 1952, Pence 1973 Mottled Owl ( Strix virgata) Blankaartiasinnamaryi skin/nest Brennan and Yunker 1966 Eutrombiculaalfreddugesi skin/nest Brennan and Jones 1960, Loomis 1969 African Wood-Owl ( Strix wood- Dermonotonspp. feathers Cheke 1972, 1978 fordii) Glaucalgesattenuatus feathers Gaud 1980 Northern Hawk Owl (Surnia Kramerellamajor feathers M6gnin and Trouessart1884a, Dub•- ulula) nin 1953, Radford 1958 SEPTEMBER2000 PARASITICMITES OF RAPTORS 231

Appendix 2. Continued.

OWL MITE HABITAT REFERENCES Kramerellasp. feathers Vasyukovaet al. 1996 Family Tytonidae CongoBay-Owl (Phodilus prigo- Dermonoton parallelus feathers Gaud 1980

Barn Owl (Tyro alba) mitesc skin Maa 1969 Aurelianiaaureliani nasalcavity Fain 1956b,1963b, Zumpt and Till 1961, Domrow 1969, 1991 Dermonoton sclerourus feathers Gaud 1980, D'Souza et al. 1986 Dermonotonsp. feathers McClure and Ratanaworabhan 1971 Glaucalgesattenuatus feathers Rothschildand Clay 1952, Radford 1958, Gaud 1958, 1980, Gaud and Till 1961, Cern• 1967, D'Souza et al. 1986 Glaucalgessp. feathers McClure and Ratanaworabhan1971, Kurey 1976 Harpyrhynchustyto skin Fain 1972 Kramerella lunulata feathers Niethammer 1938, Gaud and Pet•tot 1948a, Cern• 1967, Bunn et al. 1982 Kramerellalyra feathers Radford 1958 Kramerellaquadrata feathers Gaud 1980 Kramerellasp. feathers Kurey 1976 Leptotrombidiumnissani skin/nest Domrow 1974, Domrow and Lester 1985 Neoboydaiasp. nasalcavity Dusb/tbekand Cern• 1970 Neottialgesevans• subcutaneous Pence and Bergan 1996 Ornithonyssusbursa skin/nest Domrow 1977 Ornithonyssussylviarum skin/nest Cooper 1978 Ornithonyssussp. skin/nest Keymer 1972 Pandalurastrigisoti feathers Niethammer 1938, Radford 1958, Gaud 1958, Gaud and Mouchet 1959a, Gaud and Till 1961, Gaud 1980 Rhinoeciustytonis nasalcavity Fain 1956c, 1959a,Zumpt and Till 1961, Domrow 1969 Tytodectesstrigis subcutaneous Gene 1848, Fain 1967, Wurst and Havelka 1997 Tytodectestyto subcutaneous Fain 1966b, 1967, Pence and Ber- gan 1996 African Grass-Owl(Tyt0 capen- Dermonoton sclerourus feathers Gaud 1980 sis) Kramerellaquadrata feathers Gaud 1980 AustralianMasked-Owl (Tyt0 mites c skin Maa 1966 novaehollandiae) Tytodectestyto subcutaneous Domrow 1992 These are incorrectlyidentified or accidentalrecords. Nest record. Thesemites were found on louseflies(Hippoboscidae) on the bird. Dermonotonbubonis from Bubo sp. (Gaud 1980) is actually an invalid nomen nudum for an undescribed species ofDermonoton from Bubovirginianus (Gaud pers. comm.).