PROC. ENTOMOL.SOC.WASH. 105(4).200.^.pp. 876-882 A REVIEW OF THE NORTH AMERICAN FLEA GENUS SPICATA FOX, I. 1940 (SIPHONAPTERA: CERATOPHYLLIDAE) Robert E. Lewis .^906 Stone BrookeCircle. Ames. lA .'S0()I()-4I74. U.S.A. (e-mail: [email protected]) — Abstract. ThespeciesandsubspeciesofthegenusSplcataarereviewed.Dactylopsylla comis scapoosei Hubbard, D. c. tacomae Hubbard, and D. c. walkeri Hubbard are new synonym.s ofSpiniui coiiiis (Jordan); Dactylopsylla minUloka Prince and Stark is a new synonym ol Spicata luiditeiiaciila (Prince), and Dactylopsylla inoorei nioorei Hubbard and D. m. oregona Hubbard are new .synonyms o\'Spicatapacifica (Hubbard). A key to the remaining species is provided and their diagnt)stic characters are illustrated. Key Words: Spicata. Ceratophyllidae, distribution, host/parasite, pocket gopher There are three genera of fleas that are is an attempt to bring some order to the specific ectoparasites of North American genus with an eye toward a taxonomic re- geomyid rodents commonly known as vision ofthe subfamily towhich itbelongs. pocket gophers. As might be expected they Except for S. comis (Jordan) I have exam- appear to be closely related, have a some- ined the primary types ofall ofthe species what confusing nomenclaturai history, and, and have concluded that there are only six in recent years, have been assigned to the valid species, at most. Following is a ge- subfamily Dactylopsyllinae. Members of neric diagnosis, a key to the species and a the genus Foxclla Wagner 1929 aredistrib- briefresumeofwhat littleisknownofthese uted from western Indiana west to British taxa. Acronyms used are USNM = Nation- ColuTiibiaand south toat leastcentralMex- al MusuemofNatural History. Smithsonian ico. The other two genera, Dactylopsylla Institution, Washington, DC: BMNH = The Jordan 1929 and Spicata I. Fox 1940 are Natural History Museum. Londt)n. tmoucwhesmteorrne pNooorrtlhyAkmneorwincaanfdroarmeCreosltorricatdeod Genus Spicata I. Fox 1940 and Wyoming, west to the coast, and south Spicata I. Fox 1940: 272-276. Type spe- into Mexico. cies: Dactylopsylla (Spicata) rara I. Fox Until now the genus Spicata. which was 1940. Original designation. resurrected by Smit (1983), containedeight This genus contains species with the fol- species, three of which contained named lowing diagnosticcharacters: Head: Witha subspecies, for a total of twelve named ventral seta between the lowest setae in the taxa. Since the name was erected, ten of frontal and ocular rows. Eyes vestigial to these names have been introduced into the absent, when present, unpigmented. Sen- literature with no apparent effort being di- sory pits on antennomeres 4-6, normal for rected toward exploring their validity, hi- the family. Thorax: First pair of lateral deedtwoofthe speciesarestill knownonly plantar setae on tarsomere V not displaced from females, certainly not the diagnostic on to the plantar surface. Abdomen: Spic- sex in this order of insects. The followinii ules on slernites very indistinct or absent. VOl.lIMH 105, NUMBHR4 877 Mcilc iiKxIifieci segments: Tergiim VIII cal lobe ofst IX long and narrow, acuminate without an area spiciilosa on inner surface. \entrally(Fig.9) i<iia SterniteVIIIwithatleastafewlongventral - eMsotvpaobilnte(pFriogc.e4s)s;a-p-e6xXoafssltoVnIgIIaswiwtihdeanatapwiicda-l setae. Posterior portion of distal arm of st extension and bearing long setae laterally as IX with a patch of 3-25 setae which may wellasalongthecaudalmargin(Fig.^):apical be relatively long and straight or short and lobeofstIXalmostoval,itscephalicandcau- curly. Apical portion of this sternite form- dal marginsparallel (Fig.6) luicliwiuuiilci ing a complete, elongated ellipsoid in lat- 5. CinacuidsailonmaneragrintsheofbsottVtIoImwoifththatelseeasgtmaenstha(lFliogw. eral view. Two acetabularsetae, at least the IS) laia upper one arising remote from the margin - Marginofthissegmentwithoutan incision . . (S ofthe fixed process. Movable process long 6. CaudalmarginofstVIIes.senliallystraightver- and relatively straight, its apex sometimes ticallyformostofitslength(Fig. 14) .... cumis inclined cephalad. Aedeagus without dor- - oCfauvdaarlyimnagrgsihnapoefstVII projecting intoa lobe 7 soapical hooks or lobes. Female: Basal ab- 7. Caudal loberounded <S dt)minal sternite with lateral setae. Base of - Caudal lobe angled although its apex may be bursa copulatrix darkly sclerotized. Bulga rounded 9 of spermatheca globular. Hilla with a large S. Caudal lobe smoothly rounded from \enter papilla. Anal stylet with two ventral and - C(Ia-ulgd.al15)lobe less smoothly rounded, trianWgiu'lialrlnola onedorsal subapical setae. Large fleas,3.5- in the middle(Fig. 13) hiiiiiceps 4.5 mm, in western North America. y. Caudallobebluntlyrounded,itsveiuralmargin The following key will separate the straight,ornearly.so(Fig. lb) ... iiiuliiciuuiiln known species ofthe genus with thecaveat - Caudallobemoresharplyroundedapically, its that only four ofthe six species are known ventral marginslightlyconcave (I-'ig. 17) .p.a.cilua from the diagnostic sex and some are known from so little material that we have SpUutu hottaceps (Hubbard 1943) no knowledge ofthe range of intraspecihc (Fig. 13) variation. Even with accompanying males Dactylopsylla (Fo.xelloides)hottacepsHub- the females remain extremely difficult to bard 1943: 5, I unnumbered Hgure. USA, identify, at least until more material conies California, Del Norte County, school tt) hand. yard at Fort Dick,4I.54N I24.10W,from Thomomys hottae, 3.VII.1943, C. A. Key to Known Species of Spicata Hubbard leg. Holotype 9, USNM No. 1. Male 2 570X4. - Female 5 Nothing appears to have been added to 2. Proximul lobe of distal arm ol si IX with a our knowledge of this species since the densesiibmarginalpatchofeurly setae 3 - Proximallobeofdi.stalarmofstIXwithalew original description. The spermatheca and longsetaethatarenotcurled 4 contours ofthe caudal margin ofst VII are 3. Fixedandmovableprocesseslongandnarrow, reminiscent of S. pacifica. but in the ab- thelatterwithalmostparallelmargins(Fig. 1); sence of additional material, especially apexofstVIIIclavate(Fig. 2); apical lobeof males, the species remains an enigma. Giv- .mstarIgXinlsonsgubapnadralnlaerlro(wF,igi.ts3)cephalicandcaudalkhiiis en the amount ofindividual variation in the - Fixedandmovableprocessesbroadermesally, species ofthe three genera in this subfam- theirmarginsnotparallel (Fig. 10);apexofst ily, it is tempting to assign the two ta\a to VIII not clavate. its margins almost parallel a single species. (Fig. III;apicallobeofstIXshorter,oval, its margins notparallel(Fig. 12) piuiJHd Spiccilii coiiiis (.lordan 1929) 4. Movableprocess~4X aslongaswideatwid- (Figs. 1-3, 14) estpoint(Fig.7);apexofstVIII lackingpro- nouncedapicalextensionandbearinglongse- DcutylopsylUi comis Jordan 1929: 35: 38. taeonlyalongthecaudal margm (Fig. S);api- pi. II, fig. 26. Canada. British Colmnbia. PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON nuditenacula Figs. 1-6. 1-3.Spiciirucomis. nottypes. 1. Maleclasper. 2,ApexofmalestVIII. 3. Ape,\ofmalestIX. -6.Spicauinuditenacula, holotype.4,Maleclasper. 5, Apexofmalest VIII.6.ApexofmalestIX. VOLUME 105.NUMBER4 paciftca Figs. 7-12. 7-9,Spiciira rum. hololype. Mule clusper. 8. Apex eifmale st VIII. 9. Apex ofmalest IX. 10-12.Spicatapucitica. hokitype. 10. Male isper. 11.ApexofmalestVIII. 12, Apex ofmalest L\. Okanagan Landing. 50.12N II9.22W. following data: USA. Oregon. Lane from Tliomomys tolpoides. 2.V.1915. J. County, summit ofMcKenzie Pass, Frog A. Munroleg. Holotype 9, BMNH. Male Camp, --44.15N 121.50W. from Thonio- described by Hubbard (194,^) with the ims mazwmi, 22.Vn.1938. C. A. Hub- PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON Figs. L^-18. SpermathecaandcaudalmarginoffemalesiVII. ]?.Spicatahaniceps. holotype. 14..S".comis. nottypes. 15.S.monlicola(a)holotype,(b)paratype. 16..S'.niuliwiuuulti(alallotype,(blS.minidoka.allotype, (c)S.minidoka.paratype. 17,5.pacifica,(a)allotype,(b)S.innorciiiitiorci.allotype,(c)S.m.oregonu.allotype. 18.S. ram. nottypes. bard leg, USNM No. 57086. The author Holotype 6. allotype 9, USNM No. erroneously ret'eired to this specimen as 104686. New synonym. the "allotype." DiictylopsyllacomiswalkeriHubbard 1954: Dactylop.sylla comis .scapoosei Hubbard 170, figure unnumbered. USA, Oregon, 1954: 169. figure unnumbered. USA, Curry County, Wedderburn, 42.27N Oregon, Columbia County, Scapoose. 124.26W, from Thomomys monticoUi, 45.46N 122.54W, from Thomomys tcil- 18.IX.1949,C. A. Hubbard leg. Holotype poides. 8.V.1951,C. A. Hubbardleg. Ho- 6. BMNH. New synonym. lotype 6. allotype 9, said to be in the USNM butnot listed inthecomputerized Hubbard(1947)saidthatthisspecieswas typeinventory and not locatedby Adams so infrequently collected that he suspected and Lewis (1995). New synonym. it might be a nest flea or occur as adults Dac1ylop.sylla comis tacomae Hubbard only during the wintermonths. I have been 1954: 170, figure unnumbered. USA, able toexamine eightpairs, and in addition Washington, Pierce County. Tacoma, to the localities cited in the synonymy, 47.16N 122.30W, from Thomomys ma- specimens came from Benton, Deschutes, zamci. 10.XI.1947, C. A. Hubbard le«:. Lane, Linn and Wasco counties in Oregon VOLUME 103.NUMBER4 881 and Pcnd Oreille County in Washington. extendeastwardtoCarson City, thebulkof Hubbard (1947) also cited three females its range extends northward into central from Del Norte County in extreme north- California, almost to the Oregon border. In western California. In passing it should be theabsenceofdocumentedmales,thestatus noted that Fort Dick. California, the type of the species remains obscure, but the locality of S. bottaceps. and Smith River. shape ofthe spermatheca and the contours California, the collection site of Hubbard"s ofSt VII are distinct from other females in females, areonly slightly more than ten km the genus. apart, and both are situated on the coastal plain. Under the circumstance the identity Spicata niiditenacula (Prince 1945) of these specimens is subject to consider- (Figs. 4-6. 16) able question. Dactylopsylla (Foxelloides) niiditenacula Prince 1945: 17. figs. 3-4. USA. Mon- Spicata Dionticola (Prince 1945) tana. GallatinCounty. 14.5 km NW West (Figs. 15) Yellowstone, ~44.40N .07W. from 111 Dactylopsylla inonticoki Prince W1945: 17. Miistelafrenata, 24.V.1938. USPHS per- fig. 7. USA. Nevada. 21 km Carson sonnel leg. Holotype 6. allotype 9. City. ~-39.10N 119.55W. from Thomo- USNM No. 104661. mys nionticola. 19.VIII.1937. USPHS Dactylopsylla minidoka Prince and Stark personnel leg. Holotype 9. USNM No. 1951: 136. figs. 11-15. USA, Utah, Box 104652. Elder County. Raft River Mts. Minidoka National Forest. 12 km SE Yost, 41.59N This species, described from two fe- il3.32W. 1981 m. from Thomoniys tal- males, has evidently not been collected poides. 11.V.1947, K. R. Kelson leg. Ho- since and the male is unknown. Unlike fe- lotype 3. allotype 2. USNM No. males of so many ta.xa in this complex of 104649. New synonym. genera, these females are unequivocally distinct. The caudal margin of st. VII is I have examined the types ofS. niiditen- smoothly roundedandlacksalobeorsinus. acula and S. minidoka. as well as a pairof The spermathecaiselongated, itsbulgaand paratypes ofthe latter, and find them to be hilla are approximately equal in length and conspecific. Unfortunately. I have only the latterbearsaconspicuousapical papilla. been able to examine five males and seven Both the holotype and paratype are in the females from ratherwidely separatedlocal- USNM. ities. In additiontothetype localitiesofthe The description places this species "13 twospeciesIhavematerialfromDeschutes, miles westofCarsonCity.OrmsbyCounty. Harney, Jefferson, and Malheur counties. Nevada." Carson City is evidently no lon- Based on this distribution it seems that the ger associated with a county. There is no species is confined to the Great Basin. Ormsby County in Nevada or any of the contiguous states and thirteen miles westof Spicatapacifica (Hubbard 1943) Carson City places the locality approxi- (Figs. 10-12. 17) ^ mately in the center of Lake Tahoe. Since Dactylopsylla (Foxelloides) pacifica Hub- the United States Public Health Service bard 1943: 4, figure [unnumbered]. USA, slides usually only indicate the county Oregon, Lincoln County. DeviTs Lake. wherethe specimenswerecollected,theex- -~44.57N 124.00W. from Thomoniys act type localitycan notbeestablishedwith nionticola. 26.V1.1943, C. A. Hubbard any degree ofcertainty. However, the host leg. Holotype 6. allotype 9, USNM No. was the nominate subspecies of T. iiionri- 57085. colci. Although the range ofthis taxon does Dactylopsylla nioorei Hubbard 1949: 47. PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON pi. 6. USA, Washington. Wahkiakum Acknowledgments C1o2u3n.t2y2,W,19frkomm NTEhoCmaothmlyasmett,alp4o6i.d1e3sN. lins1.exCtureantdorm.ySaepcptrieocniaotfioInnvteortJeobhrnatEe.ZRoaowl-- 27.V.1949.A.W.Mooreleg. Holotype cJ. ogy. Carnegie Museum ofNatural History, Allotype 9. USNM No. 104653. New Pittsburgh. PA. Nancy E. Adams, Depart- synonym. ment of Entomology. National Museum of Dactylopsylla moarei oregono Hubbard Natural History. Smithsonian Institution, 1954: 170. figs, unnumbered. USA. Washington, DC, and Michael W. Hastriter, Oregon, Lincoln County, Devil's Lake. Monte L. Bean Life Sciences Museum, ~44.57N 124.00W, from Thomomys Brigham Young University. Provo. UT. for monticola, 8.IX.1949,C. A. Hubbardleg. theloanofsomeofthematerial uponwhich Holotype 6. allotype 9, USNM No. this study was based. 104665. New synonym. Literature Cited I have been able to examine four males Adams. N. E. and R. E. Lewis. 1995. An annotated and five females of this species, including catalog ofprimary types ofSiphonaptera in the the holotype and allotypeofS.pacificaand NationalMuesumofNaturalHistory.Smithsonian holotypes and allotypes ofthe two subspe- Institution.SmithsonianContributionstoZoology 560: 1-86. cies ofS. moorei. While the nominate sub- Fox.I. 1940.SiphonapterafromwesternUnitedStates. species ofthe latterwas describedfrom the Journal ofthe Washington Academy ofSciences most southwestern county of Washington 30: 272-275. astnadte,S.itmsohooruelidobreegnooitueidwtehratebdoetshcrSi.bpeadciffriocma HubbasHcruadbs.geaCn.nudsA—.theT1i9wr4o3s.manOleulwrreswlpeaestctiieveerssn.tgThiehaneFtoaxpleollcolktaeystp.egAoopfnheDew.r the same locality and host species in adja- comis. Pacific UniversityBulletin40: 1-8. cent Clatsop County. Oregon, about six 1947. Fleas ofwestern North America. Iowa years apart. All three taxa conform to the StateCollegePress.Ames.533pp. diagnostic characters ofS. pacifica and the 1949. New fleasandrecordsfromthewestern latter two are thus considered to bejunior selmatyes.ofBSulclieetnicnesof48t:he47S-o5u4t.hern California Acad- synonyms. 1954. MylastnewNorthAmericanfleas.En- tomologicalNews65: 169-175. Spicata rara (\. Fox 1940) Jordan.K. 1929.NotesonNorthAmericanfleas.Nov- (Figs. 7-9. 18) ilatesZoologicae35: 28-39. Prince. F M. 1945. Descriptionsofthreenewspecies Dactylopsylla (Spicata) rara \. Fox 1940: ofDuclyiopsxlla Jordan andone new subspecies 275, figs. 5-6. USA, Colorado. Jackson ofFuxelta Wagner, with recordsofotherspecies County,sineloco, from Thomomysfossor in thegenera(Siphonaptera). Canadian Entomol- ogist77: 15-20. \= talpoides]. 13.VI1.1926. S. C. Mc- Prince.FM.andH.E.Stark. 1951.Fournewfleasof Campbell leg. Holotype S. USNM No. the genus Dactylopsylla Jordan 1929 (Siphonap- 54013. tera). Pan-PacificEntomologist27: 128-139. Smit. F G.A.M. 1983. Keytothegeneraandsubgen- In addition tothe holotypeofthisspecies eraofCeratophyllidae.pp. 1-36.InTraub.R.,M. fIohuarvfeebmaeleensafblreomtoJeacxkasmoinneanfdiveMomnatleeszuamnad ckRooeltylhesclctohiioltndh,eoafgnedfnleeJa.rsaF. aTHnhadeddhofoawsm,tilerydesl.CaetTrihaoentsoRhpiohptyshl,slciwhdiaietl:hd counties, Colorado, and Big Horn County. noteson Iheirevolution,zoogeographyandmed- Wyoming,from ThomomystalpoidesandT. ical importance. Cambridge University Press. bottae. Most were collected in early May U.K.DistributedbyAcademicPress.Inc.London. ssoenstheisthmatayadubletsaar"ewimnotsetr"cosmpemcoiensdiunritnhge Wagnep1hr6y.3liJip..p.1w9e2l9c.heUbaeurfdZiieesneolrndaumnedrikMaunrimseclhteieCreernatloe-- the cooler months. ben. Knowia8: 316-318.