John O, whitaker,Jr. Department of Life Sciences Indiana StateUniversity Terre Haute, Indiarn 47809 Chris Maset USDI Bureau of Land Managemeot Forestry SciencesLaboratory 3200 Jefferson\Vay Corvallis,Oregon 97331 anq Robert E. Lewis Departmenr of Entomology Iovra State University of Science lo,v/a State University of Science and Technology Ames, Iowa 50011

EctoparasiticMites (ExcludingGhiggers), Fleasn and Lice from Pocket Gophers,Thomomys, in Oregon

Abslract Four of the five sDc{iesof pocket eoohers that occur io Oreqon were examined for ecto@rasites: thesetheseincluded:included: (11( 1) CarriasC-amaspocketiocket gopher,Tbo*ony balbiz,o-r*s,(2) Townseod pocket goprer,7.goprer, 7. tountewli.tountewli,tottn:ewli- (1)(1\ Mazama pocketoocket s(gopher,pc:rlter- T.7. mazama, andaod (4(4) ) northernnortiemrern 1rccket1rcckeroocket gopher.eooher.gophe\ T.Z ,albo;des.,alpo;d.es. Although the Botta pocleir gopher,sooher- T. buttae., occursoca'jfs in southwesterosouthwestertr Oreeon.Oregon, oone was includedi in this study. Major ectoparasites wete Ardrolaelabs geamlts and, Geornldoectts oregonat oi T. b*lbbar*s; E.binon,ystas tbonanv, Hdenosdmat*s on'ycbom,ydit, and Geomtdoecas idahoensis on T. toun- endi; Haenoganarzs reidi, Geinydoecn thomomyi; Androlaelips geonzls, aod, Foxella ignota recrla on T. nazama; Geomytloetu thornon tt, Ecbinon";yswsthomorny, Echinonlssa: longicbelae (?), and Foxella ignata rec k onT. tdlPa;.des.

lntloduclion Five species of pocket gophers occur in Oregon, all in the gerts Tbomamys; tltey include the Camaspocket gopher,7. bulbiaorut, Townsendpocket gopher,7. ,ou&sendi (listed as 7. u.m.btin*s by Hall 1981), Mazama pocket gopher, T, ,zltzdnw, northetn pocket gopher, T. talpoid,el and Botta pocket gopher, T, bortae. The latter is not in- cluded in this study. Pocket gophers are unusual among North American rodents in that they harbor chewing lice, Mallophaga, rathet than sucking lice, Anoplura. Chewing lice are exceedingly bost-specific; those of pocket gophers are in the gents Geomydoecus. Geomydoecxs oregont4Joccttts on T. bdbiaorus, G. id'aboensison T, townsendi, ar.d G. thornomys on both 7. nwzanw ar,d,T. talpoi'des (Price and Emerson 1971, Pdce and Hellenthal 1!80). Mites reported trom T. b*lbfuorus arc And'rol'aelap: fahrenbolzi ar,d A, geoT/t'!r. Those reported ftom T, talpoides are Andtolaelaps fabrenbolzi, A. geomy1 Eulaelaps

Northwest Sciencq Vo1. 59, No. 1, 198) stabularis,IJaetnogatndws atnbulant,H. longitarcls, H. occid,entalis,H, orycbomyd.fu,H. rcicli, Echinory:tut letuuralil E. geomydis,E, itabellinus, E. longichelae,E. reotonwe, E. thotno,myr, E. ahentil ar'd Hypoaspi: miles (Whttaket und Vilror, 1974). Thete apparendy previously are no reported parasitic mites on ?. tnazama ard r, tounsendi, Pocket gophers of the various generaand speciesaie host to 19 speciesof fleas,which are culteotly assignedto rhfee genera. This paper presents information on the later parasitic rnites, chewiog lice, arid fleas, of pocket gophers in Oregon. Although gopbets,Thonzomyl spend more time above ground than do gophers of other genera,pocLet gophers are generally isolatecl from orher animals and from other speciesof goplr"r.. S;ah isolation gives os the op- portunity to examine communities of parasites from four closely related speciesof pocket gophers from a relatively limited area, as well as to provide new data on ectoparasiticmites from two speciesof gophers.

Sludy Area The_five speciesof pocket gophers that occur in Oregon do oot, so far as is knowo, overlap (Bailey 1936, Hall 1981, Maser et al. 1()8I) but follow the physiographic provinces of Oregon as defined by Franklin and Dyrness (1!71). For exan.rple,-the southern pocket gopher occurs mainly in the Ump,quaand Rogue Valleys anJ in the extreme southern extensionof the Sitka spruce (picea :itcben:ii) zone of rhe Klamath Mounteins p,ror,ince. The Camaspocket gopher is endemic to the \fillamette Valley within the \ illamette Valley province. The Mazama pocker gopher inhabits meaclow areas of the rWestern Coast Ranges, Cascades,and High Cascadesprovinces, reganclless of major veqetatiofl type_ ]n fact, one sma1l,isolated population just penetratesthe Klamath Mouotains pistol province at River, Curry County. Two speciesoccur east of the High Cascades Province.The most widely spreadis the northerri pocket gopher that occurs jn all legetation types _major within the following foor pr"'uin."r, Basin and Range, High Lava Piains,Blue Mountains,and Owyhee UpUna. in" Townsendpocket gopher occupies two limited areasof the Shrub_steppelwirh bit sagebrush,Artemiia ":*o-)_ and_Desert yL Shrub rnajor vegetationtypes within the Basi-naod Range and Owyhee Upland Provinces. \fhere the northern and Townsend pocket gophers would appear to be symparic, they are separatedby soil type_the Townsend p

Methodsand Materials PocLetgophers were trapped, placed in plastic bags,and quick_frozenin the field with dry ice (Maser et al. 1918). After rhey were transponetl to the laboratory and thawed, staodatd museum data rvere taken They were rhen examineclfot mites and fleas by brushing their fur over a white porcelaio1mn; this method of coliectionselects 1or larger specimens and missesmany of the smaller fo'ns. More detailed examinatronswill show small mites, particularly Geonrylich*.rsp. ( Listrophoridae) , to be abuodant (Fajn et al, 1978). The firr was examioed visually {or lice, rvhich were inclividually collectedqrith the.aid of forceps.Parasites were preservedin 70 percent ethyl alcohol,cleared io Nes_ bitts, and mounted pocket ifl Hoyer's soiution. gopher sample sizes were: g for ?, bulbiooras, Il tot T. townsewli,27 for T. ,rn o*nj id fi6 tir T. tatpoidet.

a4 \flhitaker, Maser,and Lewis Resulls Informatioo on ectoparasiresof rhe four speciesof pocket gophers is given in Table 1. The hosts,with their most numefous parasiteslisted in order of decreasingabun- dance,were (1) T. bzlbi,uotat-Androlaelaps geotnysand Geomydoecusorcgonus, (2) T, townynd,i Echirzonyssatthonornlj, (I) 7. Mzdma FoxeLlaignota recula, Geo- nytloecut thottomls, Haetnoganatu reidi, ar'd Andtolaelapsgeolnjr, ^nd (.4) T, tal- poiiet-Foxella ignota rcula, Geomldoecat tbomomys, Echinony.rut tbomomyt, and E. Longichelae. The louse, Geomyrloec*s orcgonr!, w^s found on 7. bwlbfuotus, ar'd. G. idahoensit was found oo.T. toun:ewli. Geonldoecut thomanys occurredoo both T. rnazamaand T. tdlpoidet. Andtolaelap.rgeou1s is generally a common mite on pocket gophers. It was the most abundant ectoparasiteot T. bulbirorus, fourth io abundanceon T. mazand, bnt nor even found on T, ,oui;e1l.d; or T. tdlpo;det. More than one type of mite keyed o:ut to Echinonyssus longichelae; differences occurred in the sput of coxa I1 of dre female. In typical E, Longichelae,this spur was found to be about 89,m long by 79, m wide. The individuals rve listed as E. longichelae variant, however,had a much latger spur on coxa II; it was about 24p m long and 17p m wide in mites [tom T. ttazama and about 18,um long and 16p,m wide in those from T. talpo;d,er.Dr. C. S. Herin examinedspecimens homT.lDdzd.md and referred to them either as a variaot of EcbioonytsutlongicheLae or as a new species.There is also a prob- len with. Ecbinonlrrr; tho nrrls in that mites fror;, T. trtlpoide.thave setae on the dorsalplate that are ali approximatelythe samelength ( about 30 mm), but individuals ttom T. nazata have nluch shorter setae, more like those in Echinan'ttrtl,J geol7z)tdit of E, fem.aralit,A comparativestudy of Echinonyssurfrom thesegophers is needed. \7e found three raxa of fleas on gaphers,Foxelld ignota recula ar,d Spicatacomis cam)r on T. ln4zana and both of these plrs Foxella ignota utahentis ot T, talpoides. No fleaswere found on 7. bulbirorus or T. tounsewli ( Table 1). The flea, Spicata comi: cotnis, was groupedj until recently, with 14 allied taxa in the lienus DactyLop:yLla.Members of the gennsSpicata, also sFcific to pocket gophers, ate rerely collected in latge numbers.They are rare in collectioos,which leads us to suspectrhar they a{e nest fleas that peak in population during winter when the host caonot be easily collected.The genus Spicatals badly in need of taxooomic revisioo, but lack of sufficient numbers of specimenshas delayedsuch a study. The specimeos we report here (Table 1) were taken from March through September.

Discussion The major parasitesof pocket gophers in Oregon include: fleas Foxella (ignota)1 chewing lice, Geomltloecus,(oregonus, itlahoenit, thrnnomyt); and mites, Androlaelaps (geomlr'), Haenogamarts (reidi, onlchonytlis, thotnomyt), ar'd.Echinonyrut (t'em' utaLit, longicbelae,,hontorryr'). Although there is much similarity betweeo the major ectoparasitesof gopher species!rhere are also someinteresting differences. No specimens of AndrolaeLapt$,ere fouod on T, towntendi ot T. t,tlpoidet. Fleas and Haemogama:at reidi were found on 7, 712,t2d1)Mand T. taLpoidel,but not on the other two gophers. Flaetnogatna:u:onycba ljdi; was conrmononly on 7. townsendi, The fiea complex, Foxella ignota, cootains 11 currently recognizedsubspecies that

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])o V/hitaker, Maser, and Lewis occur on pocket gophers throughour the wesrern two-thirds of the United States and southern Caoada.The specieshas not been reported eastof Indiana where the oominate subspeciesoccurs, and the western complex of taxa is grorly understoodand in need of raxonomicrevision. FoxelLa ignota rcctlr hasa broad disrribution in the Pacific North- west, from Idaho west to the coast and from northern Nevada and California to sourhern British Columbia; it occurs oo the vario rs speciesof pocket gophers,as well as on other speciesof mammalsrhat frequent the gopher br-rrrows. The occurrenceof Foxella ig1tot,1lataheoriJ in Oregon was unexpected.The taxon had been previously reported from Utah and Idaho (Hubbard 1947). Ve collectedit in Harney,Malheur, and Union Counties,all in easternOregon The subspecres,utabenth, seemsto be restrictedco more arid regions and was only collectedwith the subspecies, recala,,in M^lheur Covr:.ty. Major parasitesof the plains pocket gopher, Geonyt bursarils, in Indiana include a clrewing louse,Geont3ta,oeca: iLlinoienis; a Ilea,Foxella ignota; ar.d mites,Au.lrolaelapt geomyt, Echitzonystushngichelae, E. geotnytlis,a,nd Geomylichws Jlorioanas (Tuszynski and \Thitaker 1971). Likewise, in Cloloradowhere there are four speciesof gophers repiesenting rhree genela, major parasites were chewing lice, Geotnyd,oecus(srx species), a llea, Foxeila ignota, ar'd the mites, Andtolaelaps,Geomyt, Haernoganatut "atnbulant" (probably H. reidil, and Echinonyssusgeon1d.h. The larter two miceswere abundantooly on one speciesof gopher (Miller and Ward 1!60). Although rhere are great similarities among the parasiresof pocket gophers from the thiee areas,there ate also some differences.The similariries included a complex of chewing lice, Ceotrrytloecutspp.; the flea, FoxelLaignota; and the mites, Anatolaelaps geotnyt, Haettogamalu.r sp., and Enchinonytw.r sp. The major di{ferenceswere: (1) an apparent lack of listrophodd mites io Colorado and Oregon, but an abundance in Indiana, and (2) fewer Haetnogatnasrtsp. on gophers in the east (Iodiana) and a reducecliocidence ol Echinonylrat sp. in the commuoity of ectoparasiteson gophersin Colotado. The apparent lack of listrophorid mites in Coloradoand Oregon may be an artifact of collecting,however, and not a real difference.As noted eadier, these mires are rioy and cling to individual hairs, so are regularly overlooked during routioe sealchesfor ectoparasites.I7e suspect that lisrrophorids may be abundant on gophers of both Coioracloancl Oregon.ln support of this, we have recently examined a seriesof pocket gophers,Thomonyr, from California and found a listrophorid, Geomylichn sp., to be abundant there. Only one speciesol Echinonlssut was found io Colorado,and it was abundant oo only one speciesof gopher (Miller and X(/ard 1960). In Indiana, tuo specieswere involved (Tuszynski and ri/hitaker I91I), E. LongicheLaewith large, rouoded coxal spurs,and E, geonyd,iswith smaller,poinred coxal spurs. Io Oregon, rwo specieswere found on 7, nazanta anclT. td.po;..ler,E. longichelaear,d E. thontomlt, the latter again with smaller pointed coral spr-rrs.Only E. thonronzytv'as forind ot T. touniendi, ar'd only E. fetnural was found on T. bulbitorus. It does seem peculiar the Echi.noxyssul was rret rrroreprominent on gophersfrom Colorado. Many speciesof chewing lice inhabit birds. Chewing lice that inhabit North Amer- ican mammals are primarily parasitesof carnivoresand uogulates.Two narive rodent families (gophers, Geomyidae, and porcrrpines, Erethizontidae) , aod Guinea pigs, Cattia sp., alad nvtrras, Myocartor colp J, alse serve as hosts. The chewing lice of pocket

EctoparasiticMites, Fleas,and Lice 11 gophets, Ceomltdoecus, are bighly host specific. In Oregon, for example, Geomytloecus thomomys occurs on both T. mazama and T. tdtpaide;, which suggeststhar they are closelyrelated on sibling species.The geographicaldistribution of these two speciesof pocket gophers is parapatric (Hall 1981), as one *,oulcl expect of a single species. Differences in their bacula, however, are rnuch greater than woulcl b" op".,"d in members of rhe same species (Johnson and neoson 1960), as are differencesin their habitats (Bailey 1936) . In addition, Geomyd,oecatotegoar: is specific to T. bt bitorat, a narrov'ly endemic species,and GeomlcJoeca.rid.ahoensis is specific to T. tounsencli (Price and Hellentha 1980). Tholtotztyt bottae (:ymbr.i)ut) (Hall 1981) has Ceomycloecas thastensis as a host-specific louse (price an

Acknowtedgemehts Donald K. Grayson (Deparrment of Anthropology, University of \fashington, Seattle), Gerald K. Krantz (Department of Entomology, Oregon State Universiry, Corvallis), and Robert M. Storm (Depatment of Zoology, Orqgon State Universiry, Corvallis), critically read phyllis and improved this paper. Taylor_Hill (USDI tsureau of land Managemenr,Foresuy SciencesLaboratory,3200 Jefferson N(/ay, Corvallis, Oregon) typed the various drafts of this paper. $fe sincerelyappreciate the help.

LiteralureCited rq -:G Pa:rey.v. .1. J!1-mqlmqr;,nd zo.resor o,.son.\. ^r,. rdula), : r_,r6. "*'in,*-!, "J..Yl ',l.|; i"."h ;i Mc\;co'pd.,',1 i:.' sl'it*t: llji:'s;,J: i',,f rxrn.,a..J.().whirJLer, Tr..T. C.schn1., anJ t.s Luk,.chu. Io 8. N. lc. on rhegenu.Grnzz;_ Listiophoridae)and de;nprio;s-;;',1){";"* j. 1iiiliff"; \?,tl,.lsrigmata. ,;;.i*.1",i. Imoklin, J. irna C. f. Dyrness.197j-. vegetation rVashinston. !., pNVr ^Natural- of Oregonand USDA For. Serv.ceo. Tcch.Rep. 8. pac.N"iir,-.ri r",.-aii-i".J. nlrp. S,;., p..r""a, ""tL ur1, TheMammals o{ North America.(vol.r). vilc.y & 132] Jotn Sons,Ncw l.ork. pp. Hubbard, C. A. 1947. Fleas r^".^*7Xtr)4,:6!,i;,?.-"i,?1.1;of Western North. Arnerica. The Iowa State Collegc "rpress,;i" Ames.ri;;",'-v "'"-'";':,f;5;-tidb,,li;,fllf'""1i11,fit',1,*.if *t,*:fj,:ii1.,': t$i't#.,!:Jfl;iT;i.+:,','jT:::'ii"if,H:Lt:.:',:"f;"J: Exp. Srn., Poftland, Ore.

3ll rVhitaker, Maser,and Lewis -, R. R. Kindschy, and V. Atlins. 1978. A field ice box. USDI Bureau of I-and Manag. Tech. Note T/N 320. Denver, C,olo. Miller, R- S., and R. A. Ward. 1960. Ectotrrarasitesof pocket gophers ftom C-olotado. Amer. Midl. Nat. 641382-39"1. Patton,J. L., and M. F. Smith. 1981.Molecular evoluti,oni,nTbampnp: phyletic sysrcmatics,para. phyly, and ratesof evolution.J. Mammal. 62:493-500. Price, R. D., and K. C. Emerson.1971. A revision o{ the genus Geonyd.oecas(Mallophaga: Tri- chodectidae)oI the New \trorld pocket gophers (Rodentia ceomyidae). J. Med. Entomol. 8:228-2t7. 'fhe -, and R. A. Hellenthal. 1 0. Geornyd.oec&soregon$ complex (Mallophaga: Tricho- dectidae) of the westernUnited Statespocket gophers (Rodentia Gmmyidae). Proc. Ento- mol. Soc.$?ash. 82:25 38. Tuszynski,R. C., and J. O. \(hi&ker, Jr. 1971. External para:itesof pocket gophers,Geomys bur- tarits, ftom lndiann. Amer. Midl. Nat. 87:545-548. Whitaker, J. O., Jr., and N. Vilson. 1914. Host and distribution lists of mites (Acari), parasitic and phoretic,in the hair of wild marnmaisof North Ametica, north of Mexico- Amer. Midl. Nat. 91:1-67.

Receired 24 Aagwtt 198j Accepted for pubLication 14 Februatlt 1984

Ectoparasitic Mites, Fleas,and Lice 39