Coccidian Parasites (Apicomplexa: Eimeriidae) from Insectivores. III. Seven New Species in Shrews (Soricidae: Soricinae) from Canada, Japan, and the United States

Home , Gymnure, Hylomys, Inyo shrew, Vagrant shrew

University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln

Faculty Publications from the Harold W. Manter Laboratory of Parasitology Parasitology, Harold W. Manter Laboratory of

2-1987

Lynn Ann Hertel University of New Mexico

Donald W. Duszynski University of New Mexico, [email protected]

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Hertel, Lynn Ann and Duszynski, Donald W., "Coccidian Parasites (Apicomplexa: Eimeriidae) from Insectivores. III. Seven New Species in Shrews (Soricidae: Soricinae) from Canada, Japan, and the United States" (1987). Faculty Publications from the Harold W. Manter Laboratory of Parasitology. 158. https://digitalcommons.unl.edu/parasitologyfacpubs/158

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COCCIDIANPARASITES (APICOMPLEXA: EIMERIIDAE) FROM INSECTIVORES.III. SEVEN NEW SPECIESIN SHREWS (SORICIDAE:SORICINAE) FROM CANADA, JAPAN, AND THEUNITED STATES

Lynn A. Hertel and Donald W. Duszynski Departmentof Biology,The Universityof New Mexico,Albuquerque, New Mexico87131

ABSTRACT: SinceMay 1979,458 shrews(Blarina sp. and Sorexspp.)representing 20 speciescollected in Canada, Japan,and the United States were examined for coccidia; 110 (24%)had oocysts in their feces, including8 of 21 (38%)B. brevicaudafrom Massachusetts,Ohio, Pennsylvania,and Vermont;2 of 7 (29%)S. caecutiensfrom Hokkaido and Honshu; 14 of 63 (22%) S. cinereus from Colorado, New Mexico, Pennsylvania, Vermont, Manitoba,and Ontario;3 of 7 (43%)S. fontinalis from Pennsylvania;11 of 16 (69%)S. fumeus from Massa- chusetts, Minnesota, Pennsylvania,Vermont, and Ontario; 1 of 4 (25%)S. haydeni from Minnesota; 6 of 8 (75%)S. longirostrisfrom Floridaand Virginia;1 of 2 (50%)S. ornatusfrom California;5 of 12 (42%)S. pacificus from Californiaand Oregon; 13 of 41 (32%)S. palustrisfrom California,Colorado, and New Mexico; 1 of 2 (50%)S. tenellusfrom California;11 of 105 (10%)S. trowbridgiifrom California,Oregon, and Washington;10 of 48 (21%)S. unguiculatusfrom Hokkaido;and 24 of 112 (21%)S. vagransfrom Arizona,California, Colorado, New Mexico, Oregon,and Washington.The followingcoccidians were identifiedfrom infectedshrews: Eimeria brevicaudan. sp. from B. brevicauda;Eimeria fumeus n. sp. from S. fumeus, S. pacificus,S. unguiculatus,and S. vagrans;Eimeria inyoni n. sp. from S. tenellus;Eimeria palustrisn. sp. from S. cinereus,S. fontinalis, S. fumeus, S. haydeni,S. longirostris,S. ornatus,S. pacificus,S. palustris,S. tenellus,S. trowbridgii,and S. vagrans; Eimeria vagrantisn. sp. from S. fumeus, S. trowbridgii,and S. vagrans;Isospora brevicaudan. sp. from B. brevicauda;and Isosporapalustris n. sp. from S. pacificus,S. palustris,S. trowbridgii,S. unguiculatus,and S. vagrans.The world literatureon coccidian parasitesof shrews(16 eimeriansand 3 isosporansexclusive of the 7 new species describedhere) is reviewed.

The mammalian order Insectivora is a zoo- described(Duszynski et al., 1982), but the Japansam- were in 2% logical catchall into which a number of ancient ples stored aqueous(v/v) H2SO4.Methods for and fecal and for con- and related taxa have been storing processing samples seemingly distantly centrating,measuring, and photographingoocysts have placed. However, the family Soricidae (shrews), been describedin detail (Duszynskiet al., 1982; Stout the largest of the insectivore lines, is a rather and Duszynski, 1983). The type locality is that from homogeneous assemblage of small mammals which infectedtype hosts were collected.All measure- ments are in micrometerswith size in paren- which, although virtually unknown to nonscien- ranges theses followingthe means. tists, is important to human agriculture in the control of insect, slug, and snail pests. At the University of New Mexico we have been col- RESULTS lecting shrews and moles since 1979 and gath- The coccidians, the hosts they infected, and ering skeletal, electrophoretic, karyotypic, and collection localities are presented in Table I. parasitic data to help understand the evolution- ary relationship and zoogeographic distribution COCCIDIANS of these poorly studied groups. Here we review Eimeriabrevicauda n. sp. the literature on coccidians from members of the (Figs.1-3, 17) family Soricidae and report 7 new species found Description in Blarina sp. and Sorex spp. collected from Can- Oocystusually spheroid to slightlysubspheroid (Fig. wall < of 2 inner mem- ada, Japan, and the United States. 1), 1.5, composed layers: layer branous,colorless; outer layer sculptured(Fig. 3), col- MATERIALSAND METHODS orless to pale yellow, -2/3 of total thickness; micropyle and oocyst residuum absent; polar body sometimes Fecalsamples were collected from hosts live-trapped present,but usuallyas 2-4 fragmentsof irregularsize in the field using standardpitfall traps. The samples and shape; sporulatedoocysts (n = 50) 20.3 x 19.7 from Canadaand the United Stateswere stored in 2% (18-23 x 17-23) with L:Wratio 1.03 (1.00-1.10); spo- (w/v) potassium dichromate(K2Cr2O7) as previously rocysts(n = 50) ovoid, 11.3 x 7.6 (10-14 x 7-8) with L:Wratio 1.50 (1.27-1.60); Stieda and substiedabodies present,with substiedabody about same width as Received 2 January 1986; revised 9 July 1986; ac- Stieda body or wider (Fig. 2); parastiedabody absent; cepted 9 July 1986. sporocystresiduum occasionally composed of a large 172 HERTELAND DUSZYNSKI-COCCIDIAFROM SHREWS OF THEWORLD 173

TABLE I. Eimeria spp. and Isospora spp. recoveredfrom Blarina sp. and Sorex spp. collected in the United States, Canada, and Japan.

Country: No. hosts infected/no. Host spp. County, state, or island examined (%) Coccidia spp.

B. brevicauda USA: Franklin Co., MA 1/6 (17) E. brevicauda Ashtabula Co., OH 5/8 (63) E. brevicauda, I. brevicauda Warren Co., PA 1/3 (33) E. brevicauda Lamoille Co., VT 1/4 (25) E. brevicauda S. arcticus Canada: Reynolds Dist., Man. 0/2 S. bendirii USA: Humbolt Co., CA 0/1 S. caecutiens Japan: Hokkaido 0/2 Honshu 2/5 (40) sp.* S. cinereus USA: Boulder Co., CO 1/17 (6) E. palustris Mora Co., NM 1/2 E. palustris Santa Fe Co., NM 0/6 Taos Co., NM 0/6 Westmoreland Co., PA 6/12 (50) E. palustris, sp.* Lamoille Co., VT 0/6 Canada: Piney Dist., Man. 0/1 Reynolds Dist., Man. 5/11 (45) E. palustris, sp.* Hastings Co., Ont. 1/2 E. palustris, sp.* S. fontinalis USA: Cumberland Co., PA 3/7 (43) E. palustris, sp.* S. fumeus USA: Franklin Co., MA 1/3 (33) E. fumeus, E. palustris Clay Co., MN 1/1 E. palustris Warren Co., PA 0/1 Westmoreland Co., PA 6/7 (86) E. palustris, sp.* Lamoille Co., VT 3/3 E. fumeus, E. palustris, E. vagrantis Canada: Hastings Co., Ont. 0/1 S. haydeni USA: Clay Co., MN 1/4 (25) E. palustris S. hosonoi Japan: Honshu 0/1 S. hoyi USA: Cass Co., MN 0/2 Canada: Piney Dist., Man. 0/1 S. longirostris USA: Leon Co., FL 3/3 E. palustris, sp.* Suffolk Co., VA 3/5 (60) E. palustris S. nanus USA: Santa Fe Co., NM 0/1 S. ornatus USA: San Bernardino Co., CA 1/2 E. palustris S. pacificus USA: Humbolt Co., CA 4/11 (36) E. palustris, I. palustris, spp.* Benton Co., OR 1/1 E. fumeus S. palustris USA: Mono Co., CA 1/1 E. palustris Boulder Co., CO 2/9 (22) E. palustris, sp.* Mora Co., NM 3/7 (43) E. palustris, spp.* Sandoval Co., NM 0/6 Santa Fe Co., NM 3/4 (75) E. palustris, I. palustris Taos Co., NM 4/14 (29) E. palustris, sp.* S. preblei USA: Harney Co., OR 0/2 S. tenellus USA: Mono Co., CA 1/2 E. inyoni, E. patustrispalustris 174 THEJOURNAL OF PARASITOLOGY,VOL. 73, NO. 1, FEBRUARY1987

TABLE I. Continued.

Country: No. hosts infected/no. Host spp. County, state, or island examined (%) Coccidia spp.

S. trowbridgii USA: Humbolt Co., CA 2/20 (10) E. palustris, sp.* Benton Co., OR 5/46 (11) E. palustris, E. vagrantis, I. palustris, sp.* Clallam Co., WA 2/13 (15) E. palustris, sp.* Pierce Co., WA 2/26 (8) I. palustris S. unguiculatus Japan: Hokkaido 10/48 (21) E. fumeus, I. palustris, sp.* S. vagrans USA: Apache Co., AZ 2/11 (18) E. vagrantis, E. palustris Cochise Co., AZ 1/17 (6) sp.* Mono Co., CA 3/3 E. palustris Humbolt Co., CA 1/4 (25) E. palustris Boulder Co., CO 0/2 Catron Co., NM 1/12 (8) E. palustris Mora Co., NM 1/1 E. palustris Otero Co., NM 2/16 (13) E. vagrantis, sp.* Sandoval Co., NM 10/25 (40) E. palustris, E. vagrantis, I. palustris, spp.* Santa Fe Co., NM 0/1 Taos Co., NM 0/11 - Torrance Co., NM 2/4 E. fumeus, E. palustris Harey Co., OR 0/1 Clallam Co., WA 0/1 Kittitas Co., WA 0/1 Pierce Co., WA 1/2 E. vagrantis Totals: 20 33 110/458 (24) 7

* Unsporulated oocysts of 1 (sp.) or more (spp.) morphs; unable to identify. homogeneousbody (- 4-5), but moreoften of scattered Prevalence: Found in 8 of 21 (38%)B. brevicauda homogeneousglobules (Fig. 17). Oocysts were 1,568 collected in 4 states (Table I). days old when measured. Site of infection: Unknown,oocysts recoveredfrom feces. Taxonomic summary Etymology: The nomen trivialeis derived from the Diagnosis: Only Eimeria blarinae Todd, French, specificpart of the scientificname of the host. and Levine, 1986, from B. brevicaudain Illinois has been describedfrom all Blarina spp. to date;although similar in size, it differsfrom E. brevicaudaby having Eimeria fumeus n. sp. an oocystresiduum and lackinga substiedabody. There (Figs.4, 5, 16) are 35 Eimeria spp. describedfrom other insectivore et al., 1979;Levine and Ivens, 1979; Description genera(Ahluwalia - Sinhaand Sinha, 1980;Dzerskinskii, 1982; Duszynski, Oocystellipsoid, with wall 1.0, composedof 2 lay- 1985; Duszynskiand Moore, 1986), but oocysts of E. ers: outer layer rough, yellow, -2/3 of total thickness; brevicaudado not closely resembleany of them. usually 1 polar body, but occasionally up to 4 frag- Type host: Blarina brevicauda(Say, 1823), short- ments; micropyle and oocyst residuum absent; spor- tailed shrew, Museum of SouthwesternBiology, Di- ulatedoocysts (n = 100) 27.6 x 22.4 (25-32 x 20-25) vision of Mammalogy,NK 3110 (female),D. W. Moore with L:Wratio 1.24 (1.09-1.49); sporocysts(n = 100) #744, 27 June 1980, MSB 43414. ovoid, 15.1 x 8.8 (13-17 x 8-12) with L:Wratio 1.73 Typelocality: USA. Ohio: AshtabulaCo.; 4 km W, (1.34-2.26); Stieda,sub- and parastiedabodies present 27.4 km S Ashtabula. (Figs. 4, 5), with substiedabody about same width as

FIGURES 1-10. Photomicrographs of sporulated oocysts of coccidia recovered from the feces of shrews. x 1,911. NI-Nomarski interference.1-3. Eimeria brevicaudan. sp. 1. Sporulatedoocyst in opticalcross section. 2. Sporocystliberated from brokenoocyst; note substiedabody (arrow).3 (NI). Sculpturednature of outeroocyst wall. 4, 5 (NI). Eimeriafumeus n. sp. showingsmall substiedabody (4, arrow),parastieda body (5, arrow),and largesporocyst residuum (5, *). 6. Eimeria inyoni n. sp.; NI, note highly refractilepolar body (horizontalarrow) and substiedabody (verticalarrow). 7, 8 (NI). Eimeriapalustris n. sp. 9, 10. Eimeria vagrantisn. sp.; note large substiedabody (9, *). HERTELAND DUSZYNSKI-COCCIDIAFROM SHREWS OF THEWORLD 175

* E~~~~~~~~~~~~~~~~~~~~~~~~~~1 / 4k I 176 THEJOURNAL OF PARASITOLOGY, VOL. 73, NO. 1, FEBRUARY1987

Stiedabody; sporocystresiduum a single largeglobule Eimeriapalustris n. sp. (Fig.5), -4.0, or composedof severalglobules. Oocysts (Figs. 7, 8, 18) were 725-1,171 days old when measured. Description Oocystusually spheroid to slightlysubspheroid (Figs. Taxonomic summary 7, 8), with yellow, slightlysculptured wall that gives a Diagnosis: Oocystsof this eimeriando not resemble striated appearancein optical cross section (Fig. 7), those from any species previouslydescribed from in- <1.0, composed of only 1 obvious layer; 1-2 polar sectivores. bodies present;oocyst residuumand micropyleabsent; host: sporulatedoocysts (n = 306); 17.1 x 16.6 (12-22 x Type SorexfumeusMiller, 1895, smokeyshrew, = Museum of SouthwesternBiology, Division of Mam- 12-21) with L:Wratio 1.03 (1.00-1.28);sporocysts (n x x malogy,NK 3032 (sex undetermined),R. M. Sullivan 306) ovoid, 10.8 6.5 (6-14 4-8) with L:W ratio #484, 14 June 1980, MSB 43318; NK 3041 (male), J. 1.66 (1.00-2.58); Stieda body present;sub- and para- A. Cook #361, 15 June 1980, MSB 43319; NK 3042 stieda bodies absent;sporocyst residuum composed of (male), J. A. Cook #362, 15 June 1980, MSB 43320. several dispersedgranules; sporozoites contain 1 or 2 Typelocality: USA. Vermont:Lamoille Co.; 6.4 km refractilebodies. Oocysts were 360-1,433 daysold when E, 3.2 km S Morrisville. measured. Otherhosts and localities: See Table I. Prevalence: Found in 4 of 16 (25%)S. fumeus, 1 of Taxonomic summary 12 (8%)S. pacificus,2 of 48 (4%)S. unguiculatus,and Diagnosis: Sporulatedoocysts ofE. palustrissome- 1 of ll2(<1%) S. vagrans. what resemble those of Eimeria bentongiColley and Site of infection: Unknown,oocysts recoveredfrom Mullin, 1971, from the lesser gymnure(Hylomys suil- feces. lus) from Malaya(SE Asia), Eimeria madagascariensis Etymology: The nomen trivialeis derived from the Uilenberg, 1967, from the largeMadagascar hedgehog specificpart of the scientificname of the host in which (Setifersetosus), Eimeria scapani Henry, 1932, from this species was most prevalent. the westernAmerican mole (Scapanuslatimanus) from California,and Eimeria suncusAhluwalia et al., 1979, from the common house shrew (Suncus murinus)in Eimeriainyoni n. sp. India. Six of the 11 Sorex spp. that harboredE. pa- (Figs.6, 14) lustrisshare some of theirhost rangewith S. latimanus Description and at firstwe thoughtwe might be seeing E. scapani. However ofE. with thin, smooth wall <1.0, Henry'sphotomicrograph scapaniclearly Oocyst subspheroid, shows a smooth wall thatE. lacks composed of only 1 obvious layer; usually 1-2 polar thin, oocyst palustris but sometimes to 7 refractile (see Figs. 7, 8). Another differenceis that E. palustris bodies, up highly frag- is to = ments (Fig. 6); oocyst residuumand micropyleabsent; usuallyspheroid slightlysubspheroid (L:W 1.03), = 21.6 x 19.3 x whereasE. scapaniis subspheroidto ellipsoid (L:W= sporulatedoocysts (n 100) (19-24 More subtle differencesinclude the and 17-23) with L:Wratio 1.12 (1.00-1.31);sporocysts (n = 1.20). shape ovoid 11.9 x 7.4 x with L:W ratio size of their polar bodies and sporocyst residua, but 100) (10-14 6-9) these are structuresthat their 1.60 (1.30-1.86); Stieda and substiedabodies present may change appearance with substieda widerthan Stieda over time and thus may be less reliablein distinguish- (Fig. 6), body slightly of different of E. body; parastiedabody absent; sporocystresiduum of ing oocysts species. Oocysts palustris with 1 obvious refrac- also resemble3 speciesdescribed from distant partsof dispersedgranules; sporozoites the world. the structuraldistinctions between tile body. Oocysts were 359 days old when measured. Although them are often subtle, we think it prudentto call our form a new species because the great geographicdis- Taxonomic summary tance,different host species,and lack of overlapof host Diagnosis: This species most closely resemblesEi- rangesmake it unlikelythese parasitesbelong to a sin- meria suncus Ahluwalia, Singh, Arora, Mandel, and gle species complex. The oocysts we saw differ from Sarkar,1979, from the common house shrew (Suncus those of E. bentongiby having a thinner oocyst wall murinus)from India, but differsby havinglarger oocysts (< 1.0 vs. 1.5) that is spheroidand sculptured(vs. el- (22 x 19 vs. 20 x 15)and by the presenceof a substieda lipsoid and smooth),by the type of sporocystresiduum body, which E. suncus lacks. (a few scatteredgranules vs. compact mass), and by Typehost: Sorextenellus Merriam, 1895, Inyo shrew, having sporozoites with 1-2 refractile bodies (vs. Museum of SouthwesternBiology, Division of Mam- "comma-shaped,without clear globules").They differ malogy, NK 7991 (female), S. B. George #1059, 13 fromoocysts ofE. madagascariensisby havinga rough August 1983, MSB 53229. outer oocyst wall, structureof the sporocystresiduum Typelocality: USA. California:Mono Co.; 22.5 km (few granulesvs. many irregulargranules), and by hav- N, 4 km W Bridgeport. ing sporozoites with refractilebodies that E. mada- Prevalence: Found in 1 of 2 (50%)S. tenellusfrom gascariensisapparently lacks. Finally, they differfrom California. E. suncusby havinga single-layered,rough oocyst wall Site of infection: Unknown,oocysts recoveredfrom (vs. bilayered,smooth), in shape index (1.03 vs. 1.28), feces. by lacking a micropyle cap, and by the presence of Etymology: The nomen trivialeis derived from the polar bodies. common name of the host in which this species was Typehost: SorexpalustrisRichardson, 1828, north- found. ern water shrew, Museum of SouthwesternBiology, HERTELAND DUSZYNSKI-COCCIDIAFROM SHREWS OF THEWORLD 177

FIGURES11-13. Photomicrographsof sporulatedoocysts of coccidia recovered from the feces of shrews. x 1,911. NI-Nomarski interference.11, 12 (NI). Isosporabrevicauda n. sp.; note substiedabody (12, arrow). 13. Isosporapalustris n. sp.; comparesubstieda body (arrow)with that of I. brevicauda.

Division of Mammalogy,NK 1027 (female),J. M. Ev- havinga coarse,granular sporocyst residuum, and Stie- ans #010, 8 July 1979, MSB 41120; NK 1002 (male), da and substiedabodies that E. dissimilis apparently R. A. Hoyt #098, 6 July 1979, MSB 41122. lacks. It differsfrom E. setosi in oocyst wall thickness Type locality: USA. New Mexico: Santa Fe Co.; 8 (< 1.0 vs. 2), the presenceof a substiedabody, and by km N, 12.9 km E Santa Fe. having sporozoites with obvious refractilebodies. It Otherhosts and localities: See Table I. differs from E. fumeus by lacking a parastiedabody Prevalence: Found in 12 of 63 (19%)S. cinereus,2 and by the size and shape of the substieda body (cf. of 7 (29%)S. fontinalis, 5 of 16 (31%)S. fumeus, 1 of Figs. 4, 5 and 9, 10). 4 (25%)S. haydeni,5 of 8 (63%)S. longirostris,1 of 2 Typehost: Sorex vagransBaird, 1858, vagrantshrew, (50%)S. ornatus, 1 of 12 (8%) S. pacificus, 8 of 41 Museum of SouthwesternBiology, Division of Mam- (20%)S. palustris, 1 of 2 (50%)S. tenellus, 4 of 105 malogy, NK 1589 (female), D. J. Hafner #1671, 17 (4%)S. trowbridgii,and 14 of 112 (12.5%)S. vagrans. August 1979, MSB 40945. Site of infection: Unknown, oocysts recoveredfrom Type locality: USA. New Mexico: Otero Co.; 8.8 feces. km NE Cloudcroft. Etymology: The nomen triviale is derived from the Otherhosts and localities: See Table I. specificpart of the scientificname of the host in which Prevalence: Found in 5 of 112 (4%)S. vagrans, 1 the species was first seen. of 16 (6%)S. fumeus, and 1 of 105 (<1%) S. trowbridgii. Eimeria vagrantis n. sp. Site of infection: Unknown, oocysts recoveredfrom (Figs. 9, 10, 15) feces. Description Etymology: The nomen trivialeis derived from the in Oocystellipsoid, with wall ~ 1.0, composed of 2 lay- specificpart of the scientificname of the host which ers:outer layer golden-yellow, rough, 3/4of total thick- this species was first seen. 5-10 in ness; dispersedgranules present oocyst (oocyst Isospora brevicauda n. sp. residuumor polarfragments?); micropyle absent; spor- (Figs. 11, 12, 19) ulatedoocysts (n = 100)26.6 x 21.8 (24-32 x 20-25) with L:Wratio 1.22 (1.07-1.52); sporocysts(n = 100) Description ovoid, 15.9 x 9.0(13-17 x 7-10) with L:Wratio 1.78 Oocystspheroid (Fig. 11)to slightlysubspheroid (Fig. (1.54-2.22); Stiedaand substiedabodies present(Figs. 12), wall < 1.0, probablycomposed of 2 layersof equal 9, 10), with substiedabody about 2 x widerthan Stieda thickness:outermost layer generallywith a few small body; parastiedabody absent; sporocyst residuum a bumps irregularlyspaced, and with a striatedappear- clusterof 10-15 globules;sporozoites with single pos- ance in optical cross section (Figs. 11, 12); micropyle teriorrefractile body. Oocystswere 932-1,354 days old and oocyst residuum absent; usually 4-6 polar body when measured. fragmentsof irregularshape; sporulatedoocysts (n = 50) 16.5 x 16.2 (15-18 x 15-18) with L:Wratio 1.02 Taxonomic summary (1.00-1.05); sporocysts(n = 50) ovoid, 11.6 x 7.6(10- Diagnosis: In size, this species resemblesEimeria 12 x 7-8) with L:Wratio 1.52 (1.36-1.71); Stiedaand dissimilisYakimoffand Gousseff, 1935, from the Rus- substiedabodies present(Figs. 11, 12), with substieda sian shrew(S. araneus),Eimeria setosi Uilenberg, 1967, body about2 x widerthan Stiedabody; parastieda body from the large Madagascarhedgehog (Setifer setosus), absent;sporocyst residuum of many lipid-likeglobules and E. fumeus describedabove. It differsfrom E. dis- that obscure sporozoites (Figs. 11, 12). Oocysts were similis in oocyst L:W ratio (1.22 vs. 1.47), and by 1,568 days old when measured. 178 THEJOURNAL OF PARASITOLOGY, VOL. 73, NO. 1, FEBRUARY1987

FIGURES14-20. Line drawings of sporulated oocysts of coccidia recovered from the feces of shrews; bar = 10Im. 14. Eimeria inyoni. 15. Eimeria vagrantis. 16. Eimeria fumeus. 17. Eimeria brevicauda. 18. Eimeria palustris. 19. Isospora brevicauda. 20. Isospora palustris. HERTELAND DUSZYNSKI-COCCIDIAFROM SHREWS OF THEWORLD 179

Taxonomic summary M. Evans #010, 8 July 1979, MSB 41120; NK 1028 Thirteenisosporans have been described (sex undetermined),J. M. Evans #000, 8 July 1979, Diagnosis: in from all insectivores (Levine and Ivens, 1979; Dus- specimen not deposited MSB. Fe 8 1985; Duszynski and Moore, 1986) although Type locality: USA. New Mexico: Santa Co.; zynski, km E none, as have been found in Blarina spp. Oocysts km N, 12.9 Santa Fe. yet, hosts and localities: See Table I. and sporocystsof this speciesare similar in size to those Other of araneae Golemansky, 1978, but differ by Prevalence: Found in 1 of 12 (8%)S. pacificus,3 of Isospora 1 the striatednature of the oocyst wall and the presence 41 (7%)S. palustris,3 of 105 (3%)S. trowbridgii, of of Stieda and substieda bodies (Figs. 12, 13) that I. 48 (2%)S. unguiculatus,and 1 of 112 (<1%) S. va- araneae lacks. Other isosporanspreviously described grans. from insectivoresdo not closely resembleI. brevicau- Site of infection: Unknown,oocysts recoveredfrom da. feces. The nomen triviale is derived from the Type host: Blarina brevicauda(Say, 1823), short- Etymology: tailed shrew, Museum of SouthwesternBiology, Di- specificpart of the scientificname of the host in which vision of Mammalogy,NK 3122 (female), K. McBee the oocysts were the most abundant. #121, 29 June 1980, MSB 43415. Typelocality: USA. Ohio: AshtabulaCo.; 4 km W, HOSTS 2.7 km S Ashtabula. Of those hosts with coccidia at the time they were in 2 of 21 B. brevicauda. Prevalence: Found (10%) collected,the followingparasites or groupsof parasites Site of infection: Unknown, oocysts recoveredfrom were seen. feces. Blarina brevicauda(Say, 1823). Six of 8 (75%)in- triviale is derived from the Etymology: The nomen fected short-tailedshrews were singly infectedwith E. specificpart of the scientificname of the host in which brevicauda;the other 2 wereinfected with both E. brev- we first saw it. icauda and I. brevicauda. n. sp. Sorex caecutiens Laxmann, 1788. Both infected Isospora palustris shrewscontained a of that failed (Figs.13, 20) single morph oocysts to sporulate. Description Sorex cinereusKerr, 1792. Eleven of 14 (79%)in- Oocyst subspheroid(Fig. 13), wall <1.0, probably fected masked shrewswere singly infectedwith E. pa- composed of 2 layers of equal thickness: outermost lustris(9) or a nonsporulatedform (2); 3 shrews were layer generally with a few small bumps irregularly doubly infectedwith E. palustrisand a nonsporulated spaced,and with a striatedappearance in optical cross species. section (Fig. 20); micropyleand oocyst residuumab- Sorexfontinalis Hollister, 1911. Two of 3 (67%)in- sent; usually 1 polar body, but sometimes (30%)as 2- fected Marylandshrews were singly infected with E. 4 fragments of irregularsize and shape; sporulated palustrisand 1 shrewwas infectedwith a nonsporulated oocysts (n = 100) 17.6 x 16.7 (16-19 x 16-18) with species. L:Wratio 1.06 (1.00-1.19); sporocysts(n = 100)ovoid, Sorex fumeus Miller, 1895. Eight of 11 (73%) in- 12.8 x 8.3 (12-14 x 8-10) with L:Wratio 1.56 (1.25- fected smokey shrewswere singly infected with E. fu- 1.81); Stieda and substieda bodies present (Fig. 13), meus (1), E. palustris(2), or a nonsporulatedspecies with spheroid substieda body same width as Stieda (5); 2 shrewswere doubly infectedwith E. fumeus and body; parastiedabody absent; sporocyst residuum of E. palustris;and 1 shrew was triply infected with E. many small granules(Fig. 20). Oocystswere 360-1,433 fumeus, E. palustris,and E. vagrantis. days old when measured. Sorex haydeniBaird, 1858. The only infectedshrew was singly infected with E. palustris. Taxonomic summary Sorex longirostrisBachman, 1837. Five of 6 (83%) Diagnosis: Oocysts and sporocysts of this species infectedsoutheastern shrews were singly infectedwith are very similar to those of I. brevicauda(above) and, E. palustris,and 1 was infected with a nonsporulated therefore,they are also similar to those of I. araneae. species. They differfrom I. araneae in the same ways as did I. Sorex ornatusMerriam, 1895. The only infectedor- brevicauda.They differ from I. brevicaudain 2 ways: nate shrew was singly infected with E. palustris. (1) oocyst length, oocyst width, sporocystlength, spo- SorexpacificusCoues, 1877. Fourof 5 (80%)infected rocyst width, and oocyst L:W ratio of I. brevicauda Pacificshrews were singly infected with E. palustris(1), from Blarinasp. were highly significantlysmaller (P < I. palustris(1), E. fumeus (1), or a nonsporulatedspecies 0.0001, Student'st-distribution) than those of I. pa- (1); 1 samplecontained 2 typesofnonsporulated oocysts. lustris from Sorex spp., even though the respective Sorex palustrisRichardson, 1828. Nine of 13 (69%) means and rangesappear similar, superficially; and (2) infected northern water shrews were singly infected the width of the substiedabody is about the same as with E. palustris(5) or a nonsporulatedspecies (4); 3 that of the Stieda body in I. palustris, whereas it is shrews were doubly infected with E. palustrisand I. about 2 x wider than the Stieda body in I. brevicauda palustris;and 1 sample contained 2 types of nonspor- (cf. Figs. 12, 13); the shape of the substieda body is ulated oocysts. also differentbetween the 2 species. Sorextenellus Merriam, 1895. The only infectedInyo Typehost: SorexpalustrisRichardson, 1828, north- shrew was doubly infected with E. inyoni and E. pa- ern water shrew, Museum of SouthwesternBiology, lustris. Division of Mammalogy,NK 1002 (male),R. A. Hoyt Sorex trowbridgiiBaird, 1858. All of 11 infected #098, 6 July 1979, MSB 41122; NK 1027 (female),J. Trowbridgeshrews were singly infectedwith E. palus- 180 THEJOURNAL OF PARASITOLOGY,VOL. 73, NO. 1, FEBRUARY1987

TABLEII. Eimeria spp. and Isospora spp. described to date from oocysts recovered from fecal samples of the members of the family Soriciidae.

Host Coccidian spp. Locality Reference

Blarina brevicauda E. blarinae Illinois, USA Todd et al., 1986 B. brevicauda E. brevicauda Massachusetts, Ohio, Pennsylvania, Vermont, USA Present study B. brevicauda I. brevicauda Ohio, USA Present study Crocidura leucodon E. leucodontis Azerbaidzhan, USSR Musaev and Veisov, 1961 E. chagasi* Bulgaria Golemansky, 1978 C. russulat E. crocidurae Switzerland Galli-Valerio, 1933 C. schweitzeri E. firestonei Liberia Bray, 1958 E. inilleri Liberia Bray, 1958 Neoinms anomalus E. chagasi* Bulgaria Golemansky, 1978 I. neomyi Bulgaria Golemansky, 1978 N. fodiens E. chagasi* Czechoslovakia Cema, 1961 Sorex araneus E. chagasi* Gomel district, USSR Yakimoffand Gousseff, 1935 Czechoslovakia Cernmand Daniel, 1956 Bulgaria Golemansky and Yankova, 1973 E. dissimilis Gomel district, USSR Yakimoff and Gousseff, 1935 E. soricinae Czechoslovakia Galli-Valerio, 1927 I. araneae Bulgaria Golemansky, 1978 I. soricis Bulgaria Golemansky and Yankova, 1973 S. asper E. domnashnevac Kazakhstan, USSR Dzerskinskii, 1982 E. tedoseevi Kazakhstan, USSR Dzerskinskii, 1982 S. californicus E. soricis California, USA Henry, 1932 S. cinereus E. palustris Colorado, New Mexico, Pennsylvania, Vermont, USA; Present study Manitoba, Ontario, CAN S. fontinalis E. palustris Pennsylvania, USA Present study S. furnets E. fumeus Massachusetts, Vermont, USA Present study E. palustris Massachusetts, Minnesota, Pennsylvania, Vermont, Present study USA E. vagrantis Vermont, USA Present study S. havdeni E. palustris Minnesota, USA Present study S. longirostris E. palustris Florida, Virginia, USA Present study S. ornatus E. palustris California, USA Present study S. pacificus E. furneus Oregon, USA Present study E. palustris California, USA Present study I. palustris California, USA Present study S. palustris E. palustris California, Colorado, New Mexico, USA Present study I. palustris New Mexico, USA Present study S. tenellus E. inyoni California, USA Present study E. palustris California, USA Present study S. trowbridgii E. palustris California, Oregon, Washington, USA Present study E. vagrantis Oregon, USA Present study I. palustris Oregon, Washington, USA Present study S. n,giczulatis E. fumeus Hokkaido, Japan Present study I. palustris Hokkaido, Japan Present study S. vagrans E. palustris Arizona, California, New Mexico, USA Present study E. fumneus New Mexico, USA Present study E. vagrantis Arizona, New Mexico, Washington, USA Present study I. palustris New Mexico, USA Present study Suncus murinus E. darjeelingensis West Bengal, India Sinha and Sinha, 1980 E. suncus West Bengal, India Ahluwalia et al., 1979

* Synonyms = E. komarecki. E. neomyi, E. ropotomae; see Discussion, p. 181. t See Remarks, Levine and Ivens (1979), pp. 276-277.

tris (4), I. palustris (3), E. vagrantis (1), or a nonsporulated species (3). DISCUSSION Sorex unguiculatus Dobson, 1890. Nine of 10 (90%) Shrews have a distribution infected shrews were infected with E. cosmopolitan singly fumeus (1) the and Southeast or a nonsporulated species (8); 1 shrew was doubly throughout Holarctic, Africa, infected with I. palustris and E. fumeus. Asia, being absent only from the polar regions, Sorex vagrans Baird, 1858. Twenty-two of 24 (92%) Australia, and South America below the equator infected vagrant shrews were singly infected with E. (Eisenberg, 1981; Yates, 1984). The family has I. E. E. palustris (13), palustris (1), vagrantis (3), fumeus 2 extant the Soricinae ca. 103 (1), or a nonsporulated species (4); 2 shrews were dou- groups, (11 genera, bly infected, 1 with E. palustris and E. vagrantis, the species), mainly Holarctic shrews, and the Croc- other with E. vagrantis and a nonsporulated species. idurinae (9 genera, ca. 189 species), those with HERTELAND DUSZYNSKI-COCCIDIA FROM SHREWS OF THE WORLD 181

Palearctic, Oriental, or Ethiopian distribution. the Blarina sp. and Sorex spp. we examined as Only 5 genera in the family (Soricinae: Blarina, to host specificity. On the other hand, there is Neomys, Sorex; Crocidurinae: Crocidura, Sun- considerable geographic overlap between many cus) have been examined for coccidia and the of the Sorex spp. we examined, and coccidians majority of parasites described are from Asia and were shared freely between many host species: Europe (Table II). Prior to our study, only 2 ei- we found oocysts ofE. palustris in 11 host species, merians, E. soricis in S. californicus from Cali- I. palustris in 5 species, E. fumeus in 4 species, fornia and E. blarinae in B. brevicauda from Il- and E. vagrantis in 3 species. In almost every linois, had been reported from North American instance, the hosts that shared coccidians have shrews (Henry, 1932; Todd et al., 1986). geographic distributions that overlap (see Hall, When one assembles the literature on the coc- 1981 for distribution maps). The only exception cidia of any host group, the oocysts of several to our generalization is S. longirostris, found only species often appear to be quite similar and this in the southeastern USA and it was found to be is true for several eimerians described from infected only with E. palustris, the most ubiq- shrews (e.g., E. chagasi, E. komareki, E. neomyi, uitous of the shrew eimerians seen by us. E. ropotamae). The problem that presents itself A similar sharing of eimerians may occur in is that many original descriptions are incomplete European shrews. Yakimoffand Gousseff(1935) and often (but not always) only line drawings are described Eimeria chagasi from Sorex araneus available for comparative purposes. In many in the Gomel district of White Russia. In 1956, ways, those who describe new species of coccidia Cerna and Daniel described similar oocysts that are a century behind other descriptive disciplines they named E. komareki, also from S. araneus, because they are not required to provide type but collected from Czechoslovakia. Both de- specimens of the species they describe. One rea- scriptions are incomplete by today's measure, son for this is that no universal method has been but their brief descriptions and respective line developed to preserve type specimens of oocysts drawings are virtually identical; both pairs of au- although 1 promising technique was published thors were probably seeing the same species, E. amost a decade ago (Marchiondo and Duszynski, chagasi. Cerna (1961) said she saw E. komareki 1978), and the oocysts in this preparation have in Neomysfodiens, but later Golemansky (1978) remained unchanged for over 10 years (pers. obs.). named the form Cerna (1961) saw Eimeria neo- In lieu of type specimens of oocysts for deposit myi. In the same paper Golemansky (1978) also with national museums, we suggest the next best described Eimeria ropotamae from Crocidura thing; namely, that quality photomicrographs of leucodon leucodon in Bulgaria. The descriptions representative oocysts should accompany each provided by Golemanksy (1978) show that only new species description. Additionally, as much a small size difference between the sporulated qualitative and quantitative information as pos- oocysts separate E. neomyi and E. ropotamae; sible should be provided with every description his drawings of the 2 sporulated oocysts are in- of a new coccidium. distinguishable. The difference in oocyst size may We stress these points because as we examine be attributed to the variability in host species hundreds of host species for coccidia worldwide, discussed by Reduker et al. (1985), or to inherent we sometimes see sporulated oocysts that are individual variability in oocyst size over patency identical even though they are found in different (Duszynski, 1971), or both. Based on this argu- (but related, syntopic) host genera (e.g., E. tami- ment we suggest that E. komareki, E. neomyi, asciuri, see Hill and Duszynski, 1986) and in and E. ropotamae are junior synonyms of E. other instances we see oocysts that are nearly chagasi. This concurs with a view held by Levine identical (e.g., I. brevicauda and I. palustris), but and Ivens (1979). which can be separated by quantitative (=statis- Shrews are semifossorial and, as such, they tical) and qualitative differences. As an example inhabit environments (runways, tunnels) that of the latter, the size/shape of the Stieda/sub- could be conducive to the preservation of oocysts stieda body complex may help strengthen the deposited there in host feces (adequate moisture, case to distinguish between species, if these crit- protection from UV radiation, etc.). In addition, ical data are presented. they are reasonably gregarious and are known to The coccidians we saw in this study were not share their tunnels with other shrews and to use shared between host genera, but this may be due tunnels/runways of other small mammals (e.g., as much to lack of geographic overlap between voles), habits that should allow them access to 182 THEJOURNAL OF PARASITOLOGY,VOL. 73, NO. 1, FEBRUARY1987

many different species of oocysts including their collection and processing of shrews or parasites: own. Under such circumstances, we expected that J. A. Cook, J. M. Evans, T. A. Gray, G. L. Gra- shrews, like shrew moles and moles, would have ham, J. Haydock, R. A. Hoyt, L. D. McCurley, a high rate of infection by coccidia. For example, C. A. Stout, and R. M. Sullivan, and Drs. J. H. in recent surveys of moles (unpubl.) and shrew Bandoli, S. B. George, D. J. Hafner, K. McBee, moles (Duszynski, 1985; Duszynski and Moore, D. W. Moore, K. E. Peterson, D. W. Reduker, 1986), 92% of the moles and 75% of the shrew and T. L. Yates. We are most indebted to Dr. moles had at least 1, but usually from 2 to 6 Takashi Setoguchi, his colleagues, and the stu- species of coccidian oocysts in their feces when dents of the Primate Research Institute, Kyoto examined. However, only 24% of the shrews we University, for logistical support while in Japan. examined had coccidian oocysts in their feces and the majority (97%) of these, like most other LITERATURECITED mammalian groups thus far studied (Duszynski S. R. V. G. S. A. K. et Stout and Dus- AHLUWALIA, S., SINGH, ARORA, al., 1982; Duszynski, 1983; MANDAL, AND N. C. SARKAR. 1979. Eimeria zynski and Barkley, 1985; Hill and Best, 1985; suncus sp. nov. (Sporozoa:Eimeriidae) from the Reduker and Duszynski, 1985; Reduker et al., common house shrew, Suncus murinus murinus 1985; Vance and Duszynski, 1985; Wash et al., Linnaeus.Acta Protozoologica18: 451-454. the of Li- Hill and had BRAY,R. S. 1958. On parasiticprotozoa 1985; Duszynski, 1986), only single beria.I. Coccidiaof some small mammals.Journal species infections. The factors responsible for of Protozoology5: 81-83. these different infection patterns among related CERNA,Z. 1961. The development of the coccidia insectivores only can be speculated upon at pres- Eimeriakomareki Cernm-Daniel 1956 from Sorex ent. The and of in- araneus.Acta Societa Zoologica Bohemica 25:181- genetics, physiology, ecology 183. in and in sectivores, general, shrews, particular, , AND M. DANIEL. 1956. K poznini kokcidii are poorly understood. Differences in metabo- volne zijicich drobnych ssavci. Ceskoslovenska lism, length of time spent on the surface, size of Parasitologie3: 19-23. individual home range, defecation behavior, DuszYNSKI,D. W. 1971. Increasein size of Eimeria separataoocysts during patency. Journalof Par- length of time available for coevolution between asitology57: 948-952. host and parasites, etc., probably all contribute 1985. Coccidianparasites (Apicomplexa: Ei- to the distinctly different infection patterns seen meriidae) from insectivores: New species from between shrews, moles, and shrew moles. In the shrewmoles (Talpidae)in the United States.Jour- nal of 32: 577-580. case of shrews, for example, nutritional state, age, Protozoology L. J. BARKLEY.1985. Eimeria from bats and state of sexual all seem to ,AND season, maturity of the world: A new species in Tomopeas ravus affect their physiological state (Gebczynski, 1977), from Peru. Journalof Parasitology71: 204-208. weather (e.g., night rainfall, percent cloud cover, ,G. EASTHAM,AND T. L. YATES. 1982. Eimeria etc.) influences daily activity patterns (Vickery from jumping mice (Zapus spp.): A new species and and featuresof Z. hudson- and Bider, 1977), and at least some shrews are genetic geographic ius luteus.Journal of Parasitology68: 1146-1148. known to maintain a midden separate from the ,AND D. W. MOORE. 1986. Coccidian parasites nest area for deposit of excretory material (Maser (Apicomplexa:Eimeriidae) from insectivores. II. and Hooven, 1974). Sorting out which factors Six new species from Japaneseshrew moles (Tal- Journalof 33: 276-281. the most important role in the patterns pidae). Protozoology may play V. A. 1982. Coccidia from the seen in infected of small DZERSKINSKII, genus naturally populations Eimeria in Sorex asper in Kazakhstan.In Zhi- mammals should provide fertile ground for fu- votnyi mir Kazakhstanai problemyego okhrary. ture research efforts. NaukaKazakhstkoi SSR, Alma-Ata,USSR, p. 64. EISENBERG,J. F. 1981. The mammalian radiations. The Universityof ChicagoPress, Chicago, pp. 116- ACKNOWLEDGMENTS 117. B. 1927. Notes de et This study was supported, in part, by NIH- GALLI-VALERIO, parasitologie de technique parasitologique. Centralblatt fur DHHS grant 5 S06, RR-08139-07 to the Uni- Bakteriologie,I. Abteilung, Originale 103: 177- versity of New Mexico, in part, by NSF grant 182. DEB-8004685 to T. L. Yates and, in part, by a 1933. Notes parasitologiqueset de technique I. grant from the Faculty Research Allocation parasitologique.Centralblatt fur Bakteriologie, Abteilung,Originale 129: 422-433. Committee, of Arts and Sciences, UNM. College GEBCZYNSKI, M. 1977. Body temperature in five We thank the following students and faculty in species of shrews. Acta Theriologica22,35: 521- the Department of Biology for their help in the 530. HERTELAND DUSZYNSKI-COCCIDIA FROM SHREWS OF THE WORLD 183

GOLEMANSKY,V. 1978. Description de neuf nou- ,L. A. HERTEL,AND D. W. DUSZYNSKI.1985. velles esp6ces de Coccidies (Coccidia: Eimeriidae), Eimeria spp. (Apicomplexa: Eimeriidae) infecting parasites de Micromammiferes en Bulgarie. Acta Peromyscus rodents in the southwestern United Protozoologica 17: 261-270. States and northern Mexico with description of a , ANDP. YANKOVA.1973. Studies on the species new species. Journal of Parasitology 71: 604-613. composition and occurrence of Coccidia (Sporo- SINHA,C. K., AND S. SINHA. 1980. Eimeria darjee- zoa, Coccidia) in some small mammals in Bul- lingensis sp. n. from a house-shrew, Suncus mu- garia. Izvestiya Zoologicheski Institutoi Muzei rinus soccatus (Hodgson). Acta Protozoologica 19: B'lgariyaist Akademii Nauk 37: 5-31. 293-296. HALL,R. E. 1981. The mammals of North America, STOUT,C. A., ANDD. W. DUSZYNSKI.1983. Coccidia Vol. II, 2nd ed. John Wiley and Sons, New York, from kangaroo rats (Dipodomys spp.) in the west- 600 p. ern United States, Baja California, and northern HENRY,D. P. 1932. Observations on the coccidia of Mexico with descriptions of Eimeria merriami sp. small mammals in California, with descriptions n. and Isospora sp. Journal of Parasitology 69: of seven new species. University of California 209-214. Publications in Zoology 37: 279-290. TODD, K. S., JR., R. A. FRENCH,AND N. D. LEVINE. HILL, T. P., AND T. L. BEST. 1985. Coccidia from 1986. Eimeria blarinaen. sp. (Protozoa: Apicom- California kangaroo rats (Dipodomys spp.). Jour- plexa) from the least shrew, Blarina brevicauda. nal of Parasitology 71: 682-683. Transactions of the American Microscopical So- , ANDD. W. DUSZYNSKI.1986. Coccidia (Ap- ciety 105: 182-184. icomplexa: Eimeriidae) from sciurid rodents (Eu- VANCE,T. L., ANDD. W. DUSZYNSKI.1985. Coccid- tamias, Sciurus, Tamiasciurus spp.) from the west- ian parasites (Apicomplexa: Eimeriidae) of Mi- ern United States and northern Mexico with crotus spp. (Rodentia: Arvicolidae) from the United descriptions of two new species. Journal of Pro- States, Mexico, and Japan with descriptions of five tozoology 33: 282-288. new species. Journal of Parasitology 71: 302-311. LEVINE,N. D., AND V. IVENS. 1979. The Coccidia VICKERY,W. L., AND J. R. BIDER. 1977. The effect (Protozoa, Apicomplexa) of insectivores. Revista of weather on Sorex cinereus activity. Canadian Iberica de Parasitologia 39: 261-297. Journal of Zoology 56: 291-297. MARCHIONDO,A. A., AND D. W. DUSZYNSKI. 1978. WASH, C. D., D. W. DUSZYNSKI,AND T. L. YATES. Permanent light microscopy slides of Eimeria 1985. Eimerians from different karyotypes of the nieschulzi oocysts. Journal of Parasitology 64: 163- Japanese wood mouse (Apodemus spp.), with de- 164. scriptions of two new species and a redescription MASER,C., AND E. F. HOOVEN. 1974. Notes on the of Eimeria montgomeryae Lewis and Ball, 1983. behavior and food habits of captive Pacific shrews, Journal of Parasitology 71: 808-814. Sorex pacificus pacificus. Northwest Science 48: YAKIMOFF,W. L., AND W. F. GOUSSEFF. 1935. On 81-95. the coccidia of shrews, grass-snakes, and lizards. MUSAEV,M. A., AND A. M. VEISOV. 1961. Novyi vid Journal of the Royal Microscopical Society 55: koktsidii iz belozubki persidskoi Crocidura leu- 170-173. codon Herm. Doklady Akademii Nauk Azerbaid- YATES,T. L. 1984. Insectivores, elephant shrews, tree zhanskoi SSR 17: 967-969. shrews, and dermopterans. In Orders and families REDUKER,D. W., AND D. W. DUSZYNSKI.1985. Ei- of Recent mammals of the world, S. Anderson and meria ladronensis n. sp. and E. albigulae (Api- J. K. Jones, Jr. (eds.). John Wiley and Sons, pp. complexa: Eimeriidae) from the woodrat, Neot- 117-144. oma albigula (Rodentia: Cricetidae). Journal of Protozoology 32: 548-550.