Great Basin Naturalist
Volume 48 Number 1 Article 14
1-31-1988
A review of flea collection ecorr ds from Onychomys leucogaster with observations on the role of grasshopper mice in the epizoology of wild rodent plaque
Rex E. Thomas Rocky Mountain Laboratories, Hamilton, Montana
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Recommended Citation Thomas, Rex E. (1988) "A review of flea collection ecorr ds from Onychomys leucogaster with observations on the role of grasshopper mice in the epizoology of wild rodent plaque," Great Basin Naturalist: Vol. 48 : No. 1 , Article 14. Available at: https://scholarsarchive.byu.edu/gbn/vol48/iss1/14
This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. A REVIEW OF FLEA COLLECTION RECORDS FROM ONYCHOMYS LEUCOGASTER WITH OBSERVATIONS ON THE ROLE OF CRASSHOPPER MICE IN THE EPIZOOLOGY OF WILD RODENT PLAGUE
Rex E. Thomas
Abstract —Published records of fleas collected from the northern grasshopper mouse, Ontjchomtj.s leuco^aster, indicate that these mice have a great deal of intimate environmental contact with other rodents or their burrows. Fifty-seven species of fleas have been collected from the omnivorous grasshopper mouse. The range of this mouse overlaps much of the distribution of plague. Yersinia pestis, in the western United States; and nearly one-halt of the flea species collected from O. leuco^astcr arc known to be of importance in the epizoology of plague. This article discusses the importance of the association of fleas with a hospitable secondary host in the maintenance and transmission of wild rodent plague.
Wenzel and Tipton (1966) defined host spe- often kill and consume other small rodents cies that provide suitable conditions for a large (Flake 1973). Hubbard (1947) described them ectoparasite fauna as euxenous or hospitable very colorfully as hosts. For example, certain hosts such as the sa\ age little brutes and miuderous little carnivores whose opossum, Didelphi.s marsiijyialis. collect a dis- marauding habits tell in their fleas, for upon them can be proportionately large number oi ectoparasitic fovuul all the desert species, fleas picked up from their species in comparison to others. Such hosts victims as they pommel and kill them. play potentially important evolutionary and of rodent fleas ecological roles by providing physical, behav- Traub (1985) credits the success adapted to arid habitats as due to the survival ioral, and/or physiological conditions ade- strategies of in the fur of the host quate for a large number of ectoparasite spe- remaining or, unattached, remaining wholly in the cies. Dissemination of ectoparasites between when ecologically restricted populations of pre- burrow. It would certainly be to the advan- ferred hosts by carrier hosts benefits the para- tage of fleas on the victim of a grasshopper to move, at least temporarily, onto the sites by providing opportunities for dispersal mouse and colonization of new host populations or Onycliomys to avoid a harsher second-order gene exchange with established populations. environment. As a species, northern grass- variety Wenzel and Tipton noted that since host spe- hopper mice are certainly exposed to a their semi-carnivorous nature. cies with large flea faunas must have contacts of fleas due to with many other host species or their immedi- This, however, is not adequate to explain the ate habitats, they are likely to acquire patho- phenomenal flea fauna that published records gens from ecologically more restricted hosts collectively indicate O. leucogaster to have or their parasites and thus become carriers or (Table 1). reservoirs themselves. The northern grasshopper mouse, Onij- Host Specificity chomijs leucogaster, is an example of such an animal. Members of the genus Onijchomys In a survey of flea/host associations involved are unique among North American cricetine in the ecology of plague in the western United rodents in that they are true omnivores States, Eskey and Haas (1939) reported over (Landry 1970). The range of O. leucogaster 50 species of fleas from 30,000 small wild ani- extends from Mexico to Canada in the arid mals. They observed that all of the rodent flea plains and deserts of the western United species exhibited some degree of specificity,
States (Fig. 1). While much of their diet prob- which limited them to one host species or a ably consists of arthropods, these mice will group of biologically related rodents. They
National Institutes of Health, National Institute of Alleri;\ and Infectious Diseases, Rock\ Mountain Laboratories. Hamilton, Montana 59840.
83 84 Great Basin Naturalist Vol. 48, No. 1
Compiled from various sources (McCarty, 1978)
Fig. I. The historical occurrence of liuman and animal plague in the United States (by county) from 1900 to 1986 (A), and the distribution of the northern grasshopper mouse, Onychomys leucogaster, in the United States (B).
also noted that among wild rodents having flea's ecological needs (Holland 1958). Hop- environmental contact with each other, some kins (1957) emphasized the physical condi- degree of flea interchange occurs constantly. tions provided b\' the host in determining
It is not uncommon for survey data to report limitations to host utilization. Many fleas may collection of a flea species from other than its be adapted more to nest conditions favorable primary host. Collection records convention- to eggs and larvae than to other factors ally include the flea species recovered from identified with a particular host (Hopkins various hosts in a restricted area over a rela- 1957). Benton and Miller (1979) noted that tively short period of time. As these data accu- Peromijscopsylla h. hainifer occurs over most mulate, the geographical and seasonal distri- of the range of its normal host, Microttis spp., bution of fleas on host species with which they l)ut appears to be limited to flood plains where are commonly associated are subject to anal\ - the soil type or soil moisture pro\ ides neces- sis over their entire range. Stark (1986) has sary reciuirements. Other flea species are cli- pointed out the difficulty of establishing reli- matically restricted, and climate changes with able terminology to describe host associa- altitude (Barnes et al. 1977, Wenzel and Tip- tions. He suggests use of the terms "primary, ton 1966). "secondary," and "accidental" as used by Hol- Buffer (1965) described several types of land (1964), but notes the freciuent use of burrows constructed by O. leucogaster in a "major" and "principal" as ec^uivalent terms. test arena but (juoted Bailey and Sperr\' (1929)
Collection of a flea from other than its pri- as stating that it is ({uestionable whether they mary host can be interpreted in two ways. dig their own burrows, use abandoned bur-
Either the flea is physically and/or physiologi- rows, or use those of their prey. Bailey (1931) cally adapted to one species of host by specific felt that they occupied an\ burrow found habitat or blood-meal re(juirements (e.g., abandoned or from w Inch the\ could e\ ict the
SpilopsijUns rauictili on Onict(>l(i January 1988 THOMAS; Grasshopper Mouse Fleas 85 Table 1. Species of fleas collected from Oiujcliomys spp., their noniial hosts', location of collection (city, county, state), and collection reference. AnomiopsyUus amphibohis (Neotoma, Pcromysnis and other rodent genera-nest flea) Caliente, Lincoln, Nevada (Barnes etal. 1977) A. hicmalis (Neotoma -nest flea) (Eads and Menzies 1948) , , Texas [several], Texas (Eads 1950) , A. novomexicensis (Neotoma -nest flea) Roswell, Chaves, New Mexico (Graves et al. 1974) Las Cruses, Dona Ana, New Mexico (Barnes et al. 1977) Catallagia decipiens (Small rodents esp. Peromyscus, Microtus, and Clcthrionomys) White Horse Ranch, Malheur, Oregon (Lewis 1976)' Ceylon, Saskatchewan, Canada (Holland 1985) Dactylopsylla bhiei psilos { = psila (Thomomys) Mercury, Nye, Nevada ' (Beck and Allred 1965) D. coinis (Thomomys) (Beck , Utah 1955) , D. digitenua (Geomys) Matador, Motley, Texas (Prince and Stark 1951) D. neomexicana (probablv Geomys) (Smit'l983) D. perce mis (Cratogeomys and Thomomys) (Eads and Menzies 1949) , [several], Texas (Eads 1950) , [several], Texas Diamanus viontantis (Spermophihis) (Stark 1970) Echidnophag,a gcdlinacea (Birds, rodents, large insectivores and carnivores) Albuquerque, Bernalillo, New Mexico (Wifliams and Hoff' 1951) Columbus, [Luna], New Mexico (Traub and Hoff' 1951) (Miles etal. 1952) , , Texas Roswell, Chaves, New Mexico (Graves etal. 1974) Epitedia stanfordi (primarily Peromysciis Polacca, Navajo, Arizona (Augustson 1955) (Morlan 1955) , Santa Fe, New Mexico , Utah (Stark 1958) , Dugway Proving Ground, Utah (University of Utah 1969) E. tvenmanni (primarilv Peromyscus ' Brigsdale, Weld, Colorado (CDC 1985) Foxella ignota (Geomys and Thomomys) (Eads and Menzies 1949) , Dawson, Texas and Howell 1959) , [Tooele], Utah (Parker (Hansen , Harney, Oregon 1964) Natl. Reactor Testing Sta., Idaho (Allred 1968) Navajo, (CDC 1981) , Arizona (CDC 1983) , Coconino, Arizona F. t. albertensis (Geomys and Thomomys) , Albany, Wyoming (Prince 1945) al. , Bowman, North Dakota (Larson et 1985) F. i. ignota (Geomys and Thomomys) , Grant, Nebraska (Prince 1945) F. i. omissa (Geomys and Thomomys) (Smit 1983) F. i. rectda (Geomys and Thomomys) Fields, [Harney], Oregon (Hubbard 1941) Narrows, , Oregon (Hubbard 1941) Ontario, [Malheur], Oregon (Hubbard 1941) Cody, Park, Wyoming (Wiseman 1955) F. (. utahensis (Geomys and Thomomys) , Big Horn, Wyoming (Prince 1945) Great Salt Lake Desert, Utah (Woodbury 1964) HoplopsyUus affinis =- (EupholopsyUtts glacialis) affinis (Lepus and Sylvilagiis) [several], Texas (Eads and Menzies 1949) , , [several], Texas (Eads 1950) H. anomalus (Ground squirrels and hares) Denio, [Humboldt], Nevada (Hubbard 1947) 86 Great Basin Natur.\list Vol..48, No. 1 Table 1 continued. Alamo, Lincoln, Nevada (Hubbard 1949) Searchlight, Clark, Nevada (Hubbard 1949) Utah (Allred 1952) , (Beck , , Utah 1955) , San Juan, Utah (Stark 19.58) [Tooele], Utah (Parker and Howell 19.59) Great Salt Lake Desert, Utah (Woodbury 1964) (CDC 19S3) , Coconino, Arizona Malaraeus sinomus (Peroimj.scus) Palm Springs, Riverside, California (Augustson 1943) , Riverside, California (Hubbard 1947) Denio, [Humboldt], Nevada* (Hubbard 1947) , Santa Fe, New Mexico (Morlan 1955) Mercury, Nye, Nevada (Beck and Allred 1965) M. telchinus (Peroinyscus: also Microtiis, Clcthnonomijs, and Rcithrudontomtjs) [Tooele], Utah (Parkerand Howell 1959) , , Harney, Oregon (Hansen 1964) Great Salt Lake Desert, Utah (Woodburv 1964) Dugway Proving Ground, Utah (University of Utah 1968) Natl. Reactor Testing Sta., Idaho (Allred 1968) , Colfax, New Mexico (CDC 1982) Mega rth roglossus hisetis (Peroinyscus and Neotoma-nest flea, Eads and Campos 1977) Roswell, Chaves, New Mexico (Rail et al. 1969) Roswell, Chaves, New Mexico (Graves et al. 1974) , , New Mexico or Texas (Tipton et al. 1979) M. divisus (Tamiasciurus) , Colfax, New Mexico (CDC 1982) Meringis altipecten (Dipodomys) Columbus, [Luna], New Mexico (TraubandHoin951) , Socorro, New Mexico (Eads et al. 1987) M. (irnchis (Dipodomys) Hidalgo, New Mexico "(Hul)bard 1947) CJoliunbus, [Luna], New Mexico (Traub and Hoff 1951) , Dona Ana, New Mexico (Eads et al. 1987) , Hidalgo, New Mexico (Eads et al. 1987) M. hilsingi (Dipodomys and Onychomys) [several], (Eads Menzies , Texas and 1949) , [several], Texas (Eads 1950) Roswell, Chaves, New Mexico (Railetal. 1969) Roswell, Chaves, New Mexico (Graves etal. 1974) (Lewis , , Texas 1974a) , New Mexico (Lewis 1974a) , Portales, Roosevelt, New Mexico (Pfattenbcrger and de Bruin 198(i) Lea, New Mexico (Eads et al. 1987) , Dona Ana, New Mexico (Eads etal, 1987) , Sandoval, New Mexico (Eads etal. 1987) M. californiciis (Diixxlouiys) , Kern, C^alilornia (Eads et al. 1987) , Nye, Nevada' (Eads etal. 1987) M. (liiHxhimijs (Dipodomys and man\' other small rodent genera^ Palm Springs, Ri\erside, California (.\ngustson 1943) , Riverside, Cialifornia (Hubbard 1947) St. George, Washington, Utah (Stark 1958) , Washington, Utah" (Tipton and Allred 1951) Mercury, Nye, Nevada (Beck and Allred 1965) Roswell, (Shaves, New Mexico (Rail et al. 1969) Dugway Proving (wound, Utah (University of Utah 1969) Dugway Proving (wound, Utah (University of Utah 197()a) Roswell, Chaves, New Mexico (Graves et al. 1974) , Mohave, Arizona (Eads etal. 1987) , Pima, Arizona (Eads etal, 1987) , Nye, Nevada* (i'.ads.tal. 1987) M. disparalis (Dipodomys ' , Dona Ana, New Mexico (Eads 1978) January 1988 THOMAS: Grasshopper Mouse Fleas 87 Tiihk' 1 continued. Mexico (Fads 1978) , Luna, New Mexico (Eadsetal. 19S7) , Dona Ana, New M. fdcilis (Dipodomiis) (Fads 1978) , Bernalillo, New Mexico Mexico (Eads 1978) , Chaves, New Mexico (Fads 1978) , \alencia. New Mexico (Fadsetal. 1987) , Chaves, New Mexico (Eads et al. 1987) , Bernalillo, New M. huhhardi (Peromijscus, Pcronnathus, and On\ichom\is) Crane, [Harney], Oregon (Hubbard 1947) Denio, [Humboldt], Nevada (Hubbard 1947) Harney, Oregon (Hansen 1964) , Natl. Reactor Testing Sta., Idaho (Allred 1968) (Lewis 1974a) , , Utah White Horse Ranch, Malheur, Oregon (Lewis 1975) Idaho Falls, [Bonneville], Idaho (Eads et al. 1987) (Perognathus and Dipodomifs) M . jamesoni Mexico (Morlan 1955) , Santa Fe, New (Genoways and Jones 1972) , , North Dakota M. nidi {Dipodomiis) Santa Fe, New Mexico "(Morlan 1955) Roswell, Chaves, New Mexico (Rail et al. 1969) Roswell, Chaves, New Mexico (Graves et al. 1974) (Fads et al. 1987) , Coconino, Arizona parked (Dipodomiis: also Oniicliomys and Pero^mithiis) M. ' Crane, [Harney], Oregon (Hubbard 1947) Narrows, , Oregon (Hubbard 1947) Meade, Kansas (Hubbard 1947) [several], Texas (Fads and Menzies 1949) , , , Utah (Tipton 1950) [several], Texas (Eads 1950) , Albuquerque, Bernalillo, New Mexico (Williams and Hofn951) , Santa Fe, New Mexico (Morlan 1955) Dugway, Tooele, Utah (Howell 1957) Kanab, Kane, Utah (Stark 1958) (Hansen , Harney, Oregon 1964) Great Salt Lake Desert, Utah (Woodburv 1964) Mercury, Nye, Nevada (Beck and Allred 1965) Dugway Proving Ground, Utah (Universitv of Utah 1967) Natl. Reactor Testing Sta. , Idaho (Allred 1968) Dugway Proving Ground, Utah (University of Utah 1969) Dugway Proving Ground, Utah (Ecodynamics 1971) , Bighorn, Montana (Jellison and Senger 1973) , Custer, Montana (Jellison and Senger 1973) San Juan, New Mexico (CDC 1981) , Coconino, Arizona (CDC 1982) , Albany, Wyoming (Eads et al. 1987) , Wichita, Kansas (Eads et al. 1987) M. rectus (Dipodomiis) , Santa Fe, New Mexico (Morlan 1953) , Santa Fe, New Mexico (Morlan 1955) Roswell, Chaves, New Mexico (Railetal. 1969) Roswell, Chaves, New Mexico (Graves et al. 1974) M. shannoiii (Oniichomijs, Pero^nathus and Peromijscus) Crane, [Harney], Oregon (Hubtiard 1947) Mabton, [Yakima], Washington (Hubbard 1947) A.L.E. Reserve, Benton, Washington (O'Farrell 1975) , Grant, Washington (Fads et al. 1987) Monopsijllus exilis (Oniichomiis) , Meade, Kansas (Hubbard 1943) , Yavapai, Arizona (Hubbard 1947) Roggen {sic Rugger), Colorado (Hubbard 1947) , Meade, Kansas (Hubbard 1947) Powderville, Powder River, Montana (Hubbard 1947) ) Great Basin Naturalist Vol. 48, No. 1 Table 1 continued. Crane, [Harney], Oregon (Hubbard 1947) [several], Texas (Eads and Menzies 1949) , (Tipton 19.50) , Utah , Texas (Miles etal. 1952) , , (Morlan , Santa Fe, New Mexico 1955) (Kartnian and Prince 19.56) , Nebraska (Holdenreid and Morlan 19.56) , Santa Fe, New Mexico Kanab, Kane, Utah (Stark 19.58) St. George, Washington, Utah (Stark 19.58) [Tooele], Utah (Parker and Howell 19,59) , (Poorbaugh and Gier , Finney, Kansas 1961) Great Salt Lake Desert, Utah (WoodinuA- 1964) Winnett, Petrolenm, Montana (Senger 1966) Natl. Reactor Testing Sta., Idaho (Allred 1968) (Genowavs and 1972) , , North Dakota Jones (CDC 1981) , Navajo, Arizona New Mexico (CDC 1981) , San Juan, (CDC 1982) , Colfax, New Mexico 1982) , Jones, South Dakota (Easton (CDC 198.3) , Weld, Colorado (CDC 1984) , Yavapai, Arizona Alpine, Brewster, Texas (CDC 1984) Brigsdale, Weld, Colorado (CDC 1984) Brigsdale, Weld, Colorado (CDC 1985) Portales, Roosevelt, New Mexico (Pfafifenberger and de Bruin 1986) M. wa^neri (Peromiiscus) (Eads and Menzies 1949) , [several], Texas , Harney, Oregon (Hansen 1964) Natl. Reactor Testing Sta., Idaho (Allred 1968) (CDC 1982) , Colfax, New Mexico , Bowman, North Dakota (Larson et al. 1985) al. 1985) , Mercer, North Dakota (Larson et M. If, ophidius (Peromiiscus) , Lamb, Texas (Eads 19.50) M. IV. systaltiis {PeromijscHs) Ceylon, Saskatchewan, Canada (Holland 1985) M. w. wagneri (Peromijscus) Boardman, [Morrow], Oregon (Hubbard 1941) Two Rivers, , Washington (Hul)bard 1941) Crane, [Harney], Oregon (Hul)i)arcl 1947) Denio, [Huniboldt(, Nevada (Hubl)ard 1947) Mabton, [Yakima], Washington (Hubbard 1947) Steen Mountain Pass, [Harnev], Oregon (Hubbard 1947) (Allred 1952) , , Utah , Santa Fe, New Mexico (Morlan 1955) Utah (Beck 19.55) (Stark 19.58) , , Utah [Tooele [, Utah (Parker and Howell 19.59) Great Salt Lake Desert, I'tah (Woodbiuv 1964) Mercury, Nye, Nevada (Beck and Allred 1965) Dugway Proving Ground, I'tah (UniversitvofUtah 1967) Natl. Reactor Testing Sta., Idaho (Allred 1968) Dugway Proving (Ground, Utah (Uuiversitvoi'Ulah 1969) Opisocrostis liir.siitiis (Ciiuoiniis) (Lads and Menzies 1949) , [several I, Texas (Eads 19,50) , [.several], Texas Navajo, (CDC 1981) , Arizona O. lahis {Spmu(>})}iilu\ I Narrows, Oregon (Hubbard 1947) Natl. Reactor Testing Sta., Idaho (Allred 1968) O. washingtoncnsis (S))cnit(>pliilus} (Smit 198,3) Opisodasys keeni (Peromijscus Dugway Proving (Ground, Utah (UniversitvofUtah !970b) ) January 1988 THOMAS; Grasshopper Mouse Fleas 89 Table 1 continued. Orcho))cas caedeiis (Tamiasciiirus and Glaucomys) Dii^way Proving Ground, Utah (University of Utah 1969) (). Icucopus (Pcromt/scus) (Tipton 1950) , Utah , , Santa Fe, New Mexico (Morlan 1955) Moab, Grand, Utah (Stark 1958) Alpine, Brewster, Texas (CDC 1984) Portales, Roosevelt, New Mexico (Pfaffenberger and de Bruin 1986) O. sexdentatiis (Neotoma ) (Eads and Menzies 1949) , [several], Texas O. s. agilis (Neutonia (Beck _, , Utah 1955) , Santa Fe, New Mexico (Morlan 1955) Kanab, Kane, Utah (Stark 1958) Oropsi/lla iclahocnsis (Spermophihis and other small rodents) (Stark 1970) Peromtjscop.sylla draco (Peromyscus) Roswell, Chaves, New Mexico (Graves et al. 1974) P. hesperomtjs (Peromyscus) , Dawson, Texas (Eads and Menzies 1949) Menzies , Howard, Texas (Eads and 1949) (CDC 1982) , Coconino, Arizona Portales, Roosevelt, New Mexico (Pfaffenberger and de Bruin 1986) P. h. adelpha (Peromyscus) and Traub 1954) , Rosebud, Utah (Johnson , San Juan, Utah (Stark 1958) Howell 1959) , [Tooele], Utah (Parker and Mercury, Nye, Nevada (Beck and Allred 1965) PhalacropsyUa alios (Neotoma) Natl. Reactor Testing Sta., Idaho (Allred 1968) Pidex irritans (Man and large carnivores) (Eads 1950) , , Texas Rhadinopsylla (Acthenophthalmiis) sp. (Eads and Menzies 1949) , Cochran, Texas , Yoakum, Texas (Eads and Menzies 1949) (Eads , Cochran, Texas 1950) (Eads 1950) , Yoakimi, Texas R. fraterna (Spermophihis) Natl. Reactor Testing Sta. , Idaho (Allred 1968) Rock Glen, Saskatchewan, Canada (Holland 1985) R. heiseri (Ammospermopliilus lecurus) (Parker and Howell 1959) , [Tooele], Utah Great Salt Lake Desert, Utah (Woodburv 1964) Mercury, Nye, Nevada (Beck and Allred 1965) R. multidenticulata (Onychomys leuco^aster) , Santa Fe, New Mexico (Morlan and Prince 1954) Dugway Proving Ground, Utah (University of Utah 1970b) Roswell, Chaves, New Mexico (Graves etal. 1974) (Lewis 1974a) , , Arizona , , New Mexico (Lewis 1974a) , , Texas (Lewis 1974a) R. (Micropsylla) sectilis (mainlv Peromyscus) Crane, [Harney], Oregon (Hubbard 1941) R. s. sectilis (mainlv Peromyscus) Mercury, Nye, Nevada (Beck and Allred 1965) Natl. Reactor Testing Sta. , Idaho (Allred 1968) Thrassis aridis (Dipodomys) Portales, Roosevelt, New Mexico (Pfaffenberger and de Bruin 1986) Th. aridis campestris (Dipodomys) , Mojave, Arizona (Hubbard 1947) , Hooker, Nebraska (Hubbard 1947) , Cherry, Nebraska (Hubbard 1947) , Hidalgo, New Mexico (Hubbard 1947) [several], (Eads and Menzies , Texas 1949) 90 Great Basin Naturalist Vol. 48, No. 1 Table 1 continued. , Cochran, Texas (Eads 1950) , Terry, Texas (Eads 19.50) , , Texas (Miles et al. 1952) , Santa Fe, New Mexico (Morlan 1955) Camel, Black Mountain, Utah (Stark 19.58) [Tooele], Utah (Parker and Howell 1959) , Roswell, Chaves, New Mexico (Rail et al. 1969) Roswell, Chaves, New Mexico (Graves et al. 1974) Th. a. hoffmani (Dipodomys) * St. Georf^e, Washinjjton, Utah (Tipton and Allred 1951) Mercury, Nye, Nevada (Beck and Allred 1965) Th. ahzonetisis (Neotoma. Spermo))hilu.s. Dip()(h>inijs, Ami)iosi)enn<)))l}ihi.s. and Ontjchomy.s) Palm Springs, Riverside, California (Augustson 1943) Santa Rosa Mountains, California (Augustson 194,3) , Riverside, California (Hubbard 1947) Th. b. bcicchi (Spennophihi.s) Dugway, Tooele, Utah (Howell 19.57) , Morton, North Dakota (Larson etal. 1985) Cheyenne, Laramie, Wyoming (CDC 1985) Th. h. caducus (AnunospennopJiihis and SpciDiophihis ) , Emery, L'tah (Stark 1958) Th. h. consimilis (Ammospennopliihis, Spcnnuphilii.s. and Ontichuniys) Holbrook, Navajo, Arizona (Stark 19.57) Winslow, Navajo, Arizona (Stark 19.57) Th. h. ghidiohis (Ammospennophihi.'i} Searchlight, Clark, Nevada (Hubbard 1949) , Utah (Tipton 19.50) , Dugway Valley, Tooele, L'tah (Stark 19.58) * , Washington, Utah (Stark 19.58) , [Tooele], Utah (Parker and Howell 19.59) , Harney, Oregon (Hansen 1964) Great Salt Lake Desert, Utah (Woodbur\' 1964) Mercury, Nye, Nevada* (Beck and Allred 1965) Dugway Proving Groimd, Utah (University of Utah 1970a) Th. h. johnsoni (Lci^unis) Juarez, Mexico (Trauband Holf 1951) Til. h. pan.su.'i (SfX'rmophilu.s. Amm(>spcrni()i)hihi.s. and iri'(|uentl\ OiiychoDiiis) , Brewster, Texas (Eads and Menzies 1949) , Brewster, Texas (Eads 1950) Albuquerque, Bernalillo, New Mexico (Williams and Hot! 1951) , Santa Fe, New Mexico (Morlan 19.55) Bloomfield, San Juan, New Mexico (Stark 1970) Th. h. setosis (Atnmospennophihts and fr((|utiitl\ Oiu/choiiiti.s) (Stark 1970) Th. fotti.s (Spcnfiophihi.s) Brewster, , Texas (Prince 1944) , [several], Texas (Eads and Menzies 1949) , [several], Texas (Eads 19.50) , , Texas (Miles etal. 19.52) , , Nebraska (Kartmanand Prince 19.56) Roswell, (-haves. New Mexico (Rail etal. 1969) (Juyman, Texas, Oklahoma (Stark 1970) Eunice, Lea, New Mexico (Stark 1970) Loco Hills, Eddy, New Mexico (Stark 1970) Maljamar, Lea, New Mexico (Graves et al. 1974) Roswell, ('haves. New Mexico (Stark 1970) Wellington, Larimer, Colorado (CDC 1984) Brigsdale, Weld, (Colorado (CDC 1984. 1985) Portales, Roosevelt, New Mexico (Pfaffenberger and dc Bruin 1986) Th. francisi rockwoodi (Spc'rm(>i)hihis) Blitzen, [Harney], Oregon (Hubbard 1947) Ontario, Malheur, Oregon Uiubbard 1947) Alamo, Lincoln, Ne\ada (Hubbard 1949) January 1988 THOMAS: Grasshopper Mouse Fleas 91 Table 1 continued. Th. p. pandorae {S))cnnopliilus) Denio, [Humboldt], Nevada (Hubbard 1947) Steen Mountain Pass, [Harney], Oregon (Hubbard 1947) Natl. Reactor Testing Sta., Idaho (Allred 1968) Th. petiolatus (Spcrma))hihi.s) Boardman, [Morrow], Oregon (Hubbard 1941) Denio, [Humboldt], Nevada (Hul)bard 1941) Two Rivers, , Washington (Hubl)ard 1941) 'References to normal hosts are after Lewis (1972, 1974a. 19741), 197.51 except wlu-ic otlu-r-wist i.ted. *Collected from Onijchomtjs torridus. **Probably collected from Onychomys torridus. ^Personal collection data—no reference cited. original ha\ e been added in thi^ I iew, were interred from the original record, or the original Countv records in brackets [ ] were either not in the record and record listed more than one countv of collection and is noted by [several]. indicate that gras.shopper mice frecjuently in- Table 2. Plague vectoring capacities of 25 nominate species and six additional subspecies of fleas collected vestigate or utilize burrows previously con- from Ontjchumij.s leiicogaster. structed by other rodents. This observation is supported by the number of fleas commonly Capable vectors Potential vectors associated with Dipodomys, SpcnnopJiilus, Diamanus montanus AnoiniopsijUus (imphd)ohis^ and geomyids collected from grasshopper Echidnophaiia gaUinacea Cafalla ' likely to enter a burrow of the prey species, Thrassis a. arizonensis MonupsyUii.s wagneri which makes burrow investigation an avenue Pidex irritans^ M. w. W(igneri~ Thra.ssis hacchijohn.wni Opisodasys keeni to interspecific parasitism. ' Th. francisi rockwoodi Orchopeas leucopus^ Some species of fleas considered to have Th. p. pandorae^ \ principal hosts other than O. leiicogaster have O. sexdentatii.s developed what appear to be primary associa- O. .V. aqili.s^ Peromysco)).syUa Jie.spcroniy.s adelpha^ tions with grasshopper mice from secondary Thra.ssi.s h. hacchi associations. was observed Graves et ' This by Th. h. caducus al. (1974) for Meringis nidi, M. bilsingi, and Th. h. gladiohis^ M. dipodomijs in Roswell County, New IVIex- Th. h. pansus" Th.fotus'' ico, and by Morlan and Prince (1954) for Th. petiohitufi^ RhodinopsyUa midtidenticuhita. Several of the flea species reported here irom O. leiico- 'Beck 19.^5.5 ^Pollitzer and Meyer 1961 "Egoscue 1960 'stark 1970 gaster are drawn from a single published ref- Kartman and Prince 19.56 'Thorpe 1962 erence. These may be interpreted as acciden- tal or "trivial collections that do not represent a flea/host association of any natural impor- naturally or experimentally. He defined a ca- tance at present, but they may also represent pable vector as able to transmit the disease rarely documented normal associations. either naturally or experimentally. However, the experimental success or failure of a flea Onychomys, Their Fleas and Plague species tested for vector competency does not necessarily reflect its natural ability to trans- Twenty-six of the 57 nominate species of mit the disease. Jordan (1943) proposed that a fleas collected from O. leiicogaster dre known priori every flea is a "potential" vector of to be potential or capable vectors of plague plague. (Table 2). Beck (1955) defines a potential vec- Individual grasshopper mice vary widely in tor as one that demonstrates the presence of the number of fleas that they carry. The aver- the etiologic agent in its body but has not been age number of fleas per Onychomys (flea in- found to transmit the disease organisms either dex) is often fairly low, and many are found to 92 Great Basin Natur.\list Vol. 48, No. 1 Table 3. Flea index records from Onychomiis leucofiaster collections. Number of" January 1988 THOMAS: Grasshopper Mouse Fleas 93 omnivory and habit of secondary burrow use, 1968. Fleas of the National Reaetor Testing Sta- tion. Great Basin Nat, 28(2): 7.3-87. appear to be likely participants in the mainte- Alicustson, G F 194.3. Preliminary records and discus- of wild rodent plague. As hospitable nance sion of some species of Siphonaptera from the temporary hosts, they are responsible for dis- Pacific southwest. Bull. So. California Acad. Sci. persing a variety of fleas of known importance 42(2): 69-88. in the epizoology of plague. The range of O. 19.5.5. Records of fleas (Siphonaptera) from the Pacific southwest. Bull. So. California Acad. Sci. leucogaster extends into all of the foci of .54(1): 36-39. plague in the western United States described Bailey, V 1931. Mamnuils of New Me.xico. N, Amer. Barnes (compare Fig. 1). In areas by (1982) Fauna No. .53. 412 pp, where populations of grasshopper mice have Bailey.V,andC C Sperry 1929. Life history and habits been associated with plague for a period of of the grasshopper mice, genus Onijchomijs. U.S. time, the selection pressures leading to a pop- Dept. Agric. Tech, Bull, No. 145. 19 pp. Barnes, A M. 1982. Surveillance and control of bubonic ulation resistance increase the ability of these plague in the United States. Svmp, Zool, Soc. mice to effectively maintain both the local flea London 50: 237-2.50. populations and the enzootic character of this Barne.s. A M , V J. Tipton, and J A Wilde 1977, The disease in nature. subfamily Anomiopsyllinae (Hystrichopsyllidae: Siphonaptera), 1, A review of the genus Anomtop- sijllus Baker, Great Basin Nat, 37(2): 1,30-206, Taxonomic Treatment Beck. D E 19.55, Distributional studies of parasitic arthropods in Utah, determined as actual and po- The purpose of this review is to report pub- tential vectors of Rocky Mountain spotted fever lished flea collection records from members of and plague, with notes on vector-host relation- the genus Omjchomijs and discuss the impor- ships, Brigham Young Univ, Sci, Bull,, Biol. Ser. 1(1): 1-64. tance of their associations in the epizoology of Beck, D E., and D M Allred. 1965. Siphonaptera plague. It is not my intent to address the (fleas) of the Nevada Test Site. Brigham Young validity of the ciurent taxonomy of North Univ. Sci. Bull., Biol. Ser. 7(2): 1-27. American Siphonaptera. Smit (1983) has re- Benton, A. H.. and D H Miller. 1970. Ecological fac- vised the superspecific taxonomy of fleas of tors in the distribution of the flea Peromyscopsylla h. luimifer. Amer, Midi. Nat. 81: ,301-303. the family Ceratophyllidae. Due to the nature CDC 1981-1985. Annual collection data. Plague of this article, I have adopted Holland's (1985) Branch, Division xif Vector-borne Viral Diseases, use of the conservative taxonomy in order to Centers for Disease Control, Fort Collins, Colo- retain the generic names reported in the origi- rado 80.522, USA. Unpubhshed manuscript. nal literature. For determination of generic Fads. R B 19.50. The fleas of Texas. Control Div., Ento- niol. Bureau, Texas State Health Dept. 85 classification according to Smit, the reader is pp. 1978, Two new species of fleas of the genus referred to Appendix 5 of Traub, Rothschild, Meringis (Siphonaptera: Hystrichopsyllidae), and Haddow (1983). Johnson (1961) reduced Great Basin Nat, 38(4): 447-4,55, the subspecific status of the geographic races Fads. R B , and E G Campo.s. 1977. Notes on the flea of MonopsyUus ivafineri. Again, in order to genus Megartliroglossus (Siphonaptera: Hystri- retain the original taxonomy reported in the chopsyllidae: Anomiopsyllinae), with description ofa new species. Med. Entomol. 14(1): 97-100. collection records, I follow Holland (1985) in J. Eads, R B , E G. Campos, and G O Maupin 1987, A the use of the subspecific epithets. review of the genus Meringis (Siphonaptera: Hys- trichopsyUidae). J. Med. Entomol. 24: 467-476. Acknowledgments Eads, R B , and G C Menzies 1948. An undescribed AnomiopsyUus Baker from the pack rat, Neotoma micropus Baird. J, Kansas Entomol. Soc. 21(4): I am very grateful to Drs. Allan M. Barnes, 1.3,3-1,36. Cluff E. Hopla, Harold Stark, and Robert E. 1949. A preliminary list of the Siphonaptera of Lewis for reviewing this manuscript and mak- .33-39. Texas. Texas J. Sci. 1(4): ing valuable suggestions concerning the text Easton, E. R. 1982. An annotated checklist of the fleas of and format. South Dakota (Siphonaptera). Entomol. News 93(5): 155-158. ECODYNAMICS, Inc 1971. Ecological studies in western Literature Cited Utah, 1970 Annual Report, Ecology and Epizool- ogy Series No, 71-1, Salt Lake City, Utah, 116 pp, Allred, D. 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