Parasites of Tadarida brasiliensis mexicana (Chiroptera: Molossidae) from Arid Regions of Mexico

Comp. Parasitol. 70(1), 2003, pp. 11–25

Parasites of Tadarida brasiliensis mexicana (Chiroptera: Molossidae) from Arid Regions of Mexico CARMEN GUZMA´N-CORNEJO,1 LUIS GARCI´A-PRIETO,2 GERARDO PE´REZ-PONCE AND JUAN B. MORALES-MALACARA1

DE

LEO´N,2,3

1 Laboratorio de Acarologı ´a, Departamento de Biologı´a Comparada, Facultad de Ciencias, Universidad Nacional Auto´noma de Me´xico, C.P. 04510 Me´xico D.F., Mexico (e-mail: [email protected]) and 2 Laboratorio de Helmintologı ´a, Instituto de Biologı´a, Universidad Nacional Auto´noma de Me´xico, Ap. Postal 70-153, C.P. 04510 Me´xico D.F., Mexico

ABSTRACT: Ninety-eight Mexican free-tailed bats, Tadarida brasiliensis mexicana, collected from 4 locations in Mexico between November 1996 and June 1998, were examined for metazoan parasites. Twenty-one parasitic taxa belonging to 3 phyla (Plathyhelminthes, Nematoda, and Arthropoda) were recovered including 5 helminth taxa (3 digenean, 1 cestode, and 1 nematode) and 16 arthropod taxa (12 mite, 2 tick, and 2 insect). The digenean Ochoterenatrema labda was the most prevalent and abundant helminth in collections from the states of Puebla, Zacatecas, and Durango. Urotrema scabridum and Dicrocoelium rileyi reached the highest values of prevalence and mean abundance of infection in Nuevo Leo´n state. Among arthropod taxa recovered, Chiroptonyssus robustipes was the most prevalent and abundant parasite in all 4 localities, and its nymphal stage was the most abundant form recovered. In total, 10 new host and 43 locality records are presented. KEY WORDS: Tadarida brasiliensis mexicana, Mexican free-tailed bat, Molossidae, helminths, arthropods, Puebla, Zacatecas, Durango, Nuevo Leo´n, Mexico.

MATERIALS AND METHODS

Among the 925 species of bats described around the world, approximately 15% are found in Mexico (Ramı´rez-Pulido et al., 1996; Wilson, 1997). The Mexican free-tailed bat Tadarida brasiliensis mexicana is one of the most widely distributed species, ranging across the whole Mexican territory (Villa and Cockrum, 1962; Hall, 1981). Despite its broad distribution, knowledge about the metazoan parasites of this bat species is fragmentary, comprising only a few isolated records (Caballero, 1940, 1942, 1943; Hoffmann, 1944: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico; Eads et al., 1957; Jameson, 1959; MacDaniel and Baker, 1962; Cain, 1966; Caballero and Caballero-Rodrı´guez, 1969; Martin, 1976; see Pe´rez-Ponce de Leo´n et al. [1996] and references therein). In this study we present results of a survey of endoparasitic and ectoparasitic metazoans recovered from Ta. b. mexicana from several collection localities in arid regions of central and northeastern Mexico and incorporate known faunal and distributional records for these parasites.

Ninety-eight Mexican free-tailed bats from 4 arid localities of Mexico were collected. All bats were examined postmortem for helminths and arthropods. Individual bats were dissected, their organs (liver, gall bladder, spleen, lungs, heart, stomach, and intestine) placed in separate petri dishes with 8.5% saline, and examined with the aid of a stereo microscope. Digeneans and cestodes were fixed with Bouin’s fluid and nematodes with 70% boiling ethanol. Platyhelminthes were stained with Delafield’s haematoxylin, Mayer’s paracarmine, or chlorhidric carmine; nematodes were cleared with Amman’s lactophenol. Voucher specimens were deposited at the Coleccio´n Nacional de Helmintos (CNHE), Instituto de Biologı´a, Universidad Nacional Auto´noma de Me´xico (UNAM), Me´xico D.F., Mexico. Arthropods were collected from the external surface of bats, including the dorsal and ventral body, wings and tail membrane, ears, and nose. All arthropods were fixed in 70% ethanol. Mites and ticks were preserved as semipermanent mounts in Hoyer’s medium with a ring of Glyptal. Fleas were cleared in 10% KOH solution, transferred to hot water, and dehydrated through a gradual alcohol series. Voucher specimens were deposited in the collection of J.B.M.-M., housed at the Laboratorio de Acarologı´a, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico. Abbreviations associated with specimen accession numbers presented in this study refer to the following museums and collections: The Natural History Museum, British Museum, London, United Kingdom (BMNH); Canadian National Collection, Ottawa, Canada (CDA); Coleccio´n Nacional de Helmintos, Insti-

3 Corresponding author (e-mail: ppdleon@servidor. unam.mx).

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COMPARATIVE PARASITOLOGY, 70(1), JANUARY 2003

tuto de Biologı´a, UNAM, Me´xico D.F., Mexico (CNHE); Field Museum of Natural History, Chicago, Illinois, U.S.A. (FMNH); University of Michigan Museum of Zoology, Ann Harbor, Michigan, U.S.A. (HK); Harold W. Manter Laboratory of Parasitology, University of Nebraska, Lincoln, Nebraska, U.S.A. (HWML); collection of Juan Morales-Malacara, Laboratorio de Acarologı´a, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico (JMM); National Museum of Natural History, Washington, D.C., U.S.A. (NMNH or USNM); Rocky Mountain Laboratory, Hamilton, Montana, U.S.A. (RML); United States Department of Agriculture National Veterinary Services Laboratories, Ames, Iowa, U.S.A. (USDA); United States National Parasite Collection, Beltsville, Maryland, U.S.A. (USNPC); United States National Tick Collection, Statesboro, Georgia, U.S.A. (USNTC). Prevalence and mean abundance were calculated and used as defined in Margolis et al. (1982). The Kruskal–Wallis nonparametric test (k $ 3) was used to establish significant differences in mean abundance. Tests were conducted only when parasites were present in all 4 collection localities.

Tadarida brasiliensis mexicana (Saussure, 1860) Nombre de Dios, Durango state, Mexico (238 509170N; 1048149410W): November 1996 and July 1997, n 5 31; Rı´o Salado, Zapotitla´n de las Salinas, Puebla state, Mexico (188209000N; 978289300W): May 1997 and June 1998, n 5 12; Cueva de la Boca, Santiago, Nuevo Leo´n state, Mexico (258259550N; 1008099070W): October 1997 and May 1998, n 5 27; Concepcio´n del Oro, Zacatecas state, Mexico (248379020N; 1018259050W): October 1997 and April 1998, n 5 28. Digenea Dicrocoelium rileyi Macy, 1931 Prevalence and mean abundance: Nombre de Dios, 4 of 31 hosts examined (12.9%, 0.5 6 2.3); Rı´o Salado, 2 of 12 hosts examined (16.6%; 6 6 14.3); Cueva de la Boca, 8 of 27 hosts examined (30%, 4.5 6 10.6); Concepcio´n del Oro, 3 of 28 hosts examined (10.1%, 3 6 3.7); 57/81 (70%) (Macy, 1931); 1/6 (16.6%) (Chandler, 1938); 12/21 (58%) (Jameson, 1959); 6/16 (37.5%) (Cain, 1966); 1/7 (14.3%), and 1/9 (11.1%) (Nickel and Hansen, 1967); 1/28 (3.6%) (Blankespoor and Ulmer, 1970); 1/33 (3%) (Foster and Mertins, 1996). Site of infection: Gall bladder, bile ducts. Type host: Tadarida brasiliensis cynocephala. Reported hosts: Tadarida brasiliensis cynocephala, (Macy, 1931; Martin, 1976; Foster and

Mertins, 1996); Nycticeius humeralis, (Chandler, 1938); Ta. b. mexicana (syn. Tadarida mexicana), (Eads et al., 1957; Jameson, 1959; Cain, 1966; Nickel and Hansen, 1967; Caballero and Caballero-Rodrı´guez, 1969; Martin, 1976); Myotis velifer incautus, (Nickel and Hansen, 1967); Eptesicus fuscus, (Blanckespoor and Ulmer, 1970). Other locality records: Cueva del Guano, Nuevo Leo´n, Mexico (Caballero and Caballero-Rodrı´guez, 1969); Alachua County, Florida, U.S.A. (Foster and Mertins, 1996); Aetna, Kansas, U.S.A. (Macy, 1931); Comanche, Kansas, U.S.A. (Nickel and Hansen, 1967); Iowa, U.S.A. (Blankespoor and Ulmer, 1970); New Orleans, Louisiana, U.S.A. (Martin, 1976); Carlsbad Caverns, New Mexico, U.S.A. (Cain, 1966); Freedom, Oklahoma, U.S.A. (Macy, 1931), and Woods, Oklahoma, U.S.A. (Nickel and Hansen, 1967); Bracken Cave, Texas, U.S.A. (Martin, 1976); Central Texas, Texas, U.S.A. (Eads et al., 1957); Frio Cave, Uvalde County, Texas, U.S.A. (Jameson, 1959); Houston, Texas, U.S.A. (Chandler, 1938). Specimens deposited: CNHE 3870–3875. Other known specimens: HWML 37522.

Ochoterenatrema labda Caballero, 1943 (Syn. Prosthodendrium labda (Caballero, 1943) Yamaguti, 1958) Prevalence and mean abundance: Nombre de Dios, 18 of 31 hosts examined (61.3%, 14.2 6 23.8); Rı´o Salado, 4 of 12 hosts examined (33.3%; 70. 1 6 228.3); Cueva de la Boca, 8 of 27 hosts examined (30.8%, 1.4 6 3.3); Concepcio´n del Oro, 2 of 28 hosts examined (7.1%, 3 6 15.3); 6/21 (28.6%) (Jameson, 1959); 6/18 (33.3%) (Cain, 1966); 223/255 (87.8%), 3/42 (7.1%), 5/22 (22.7%), 12/18 (66.7%), 5/52 (9.6%), 15/27 (55.6%), and 8/32 (25%) (Lotz and Font, 1991); 1/33 (3%), 2/7 (29%), and 1/3 (33%) (Foster and Mertins, 1996). Site of infection: Intestine. Type host: Tadarina brasiliensis mexicana. Reported hosts: Natalus stramineus (5Natalus mexicanus), (Caballero, 1943); Ta. b. mexicana, (Caballero, 1943; Jameson, 1959; Cain, 1966; Martin, 1976); Myotis nigricans nigricans, (Caballero and Caballero-Rodrı´guez, 1969); Ta. b.

´ N-CORNEJO ET AL.—PARASITES OF TADARIDA BRASILIENSIS MEXICANA GUZMA

cynocephala, (Martin, 1976; Foster and Mertins, 1996); Myotis austroriparius, Nyctic. humeralis, Pipistrellus subflavus, and Tadarida brasiliensis, (Lotz and Font, 1991). Other locality records: Convento de Acolman, Estado de Me´xico, Mexico, Bosque de Chapultepec, Mexico City, Mexico and Cueva de Xictli, Mexico City, Mexico (Caballero, 1943); Cocoli, Canal Zone, Panama (Caballero and Caballero-Rodrı´guez, 1969); Alachua County, Marion County, and Polk County, Florida, U.S.A. (Foster and Mertins, 1996); East Baton Rouge County, Desoto County, Livingston County, Pointe Coupee County, and Tangipahoa County, Louisiana, U.S.A. (Lotz and Font, 1991); New Orleans, Louisiana, U.S.A. (Martin, 1976); Carlsbad Caverns, New Mexico, U.S.A. (Cain, 1966); Bracken Caves, Texas, U.S.A. (Martin, 1976); Frio Cave, Uvalde County, Texas, U.S.A. (Jameson, 1959). Specimens deposited: CNHE 3864–3869. Other known specimens: CNHE 1198–1200; HWML 37523; USNPC 85415.

Urotrema scabridum Braun, 1900 Prevalence and mean abundance: Nombre de Dios, 1 of 31 hosts examined (3.2%, 0.03 6 0.2); Rı´o Salado, 1 of 12 hosts examined (8.3%; 4.6 6 16.2); Cueva de la Boca, 10 of 27 hosts examined (37%, 4.4 6 15.3); 11/20 (55%) (Ubelaker, 1966); 9/13 (69.2%) (Nickel and Hansen, 1967); 7/28 (25%) (Blankespoor and Ulmer, 1970); 4/15 (26.6%) and 8/29 (27.6%) (Webster, 1971); 14/32 (43.8%), 4/22 (18.2%), 57/255 (22.4%), 31/42 (73.8%), 12/22 (54.5%), 14/18 (77.8%), and 1/52 (1.9%) (Lotz and Font, 1991); 9/33 (27%) and 1/7 (14%) (Foster and Mertins, 1996). Site of infection: Intestine. Type host: Promops centralis (syn. Molossus nasatus). Reported hosts: Promops centralis, Molossus ater (syn. Molossus nigricans), (Braun, 1900); Ta. b. mexicana, (Caballero, 1942; Eads et al., 1957; Martin, 1976); Na. stramineus, (Caballero, 1942); Eptesicus propinquus, (Caballero et al., 1957); Myot. velifer (Eads et al., 1957); Phyllostomus hastatus, (Caballero and Grocott, 1960); Myotis grisescens, (Ubelaker, 1966; Nickel and Hansen, 1967); Molossus molossus

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(syn. Molossus major), (Caballero and Caballero-Rodrı´guez, 1969; Groschaft and Del Valle, 1969; Odening, 1969; Zdzitowiecki and Rutkowska, 1980); Ta. b. muscula, (Groschaft and Del Valle, 1969; Odening, 1969); Lasiurus cinereus villosissimus, Myotis chiloensis, Myotis bocagei, (Caballero and Caballero-Rodrı´guez, 1969); Natalus lepidus, (Odening, 1969; Zdzitowiecki and Rutkowska, 1980); Myotis lucifugus, (Blankespoor and Ulmer, 1970; Coggins et al., 1982; Lotz and Font, 1991); Ta. brasiliensis, (Webster, 1971); Pteronotus macleayii (syn. Chilonycteris macleayii), (Odening, 1969; Webster, 1971; Zdzitowiecki and Rutkowska, 1980); Mol. ater (syn. Molossus rufus), (Led and Boero, 1973); Ta. b. cynocephala, (Martin, 1976; Foster and Mertins, 1996); Pteronotus parnellii (syn. Chilonycteris rubiginosa), Macrotus waterhousii, Ep. fuscus, Nyctinomops laticaudatus (syn. Tadarida laticaudata yucatanica), Tadarida sp., (syn. Tadarida minuta), (Zdzitowiecki and Rutkowska, 1980); Myotis keenii, (Coggins et al., 1981); Myot. austroriparius, Nyctic. humeralis, Pi. subflavus, (Lotz and Font, 1991). Other locality records: La Plata, Argentina (Led and Boero, 1973); Brazil (Braun, 1900); Recife, Brazil (Caballero and Caballero-Rodrı´guez, 1969); Volca´n Poas, Costa Rica (Caballero et al., 1957); Habana, Cuba (Odening, 1969; Zdzitowiecki and Rutkowska, 1980); Isla de Pinos, Matanzas, Cuba, and Las Villas, Cuba (Zdzitowiecki and Rutkowska, 1980); Oriente, and Pinar del Rı´o, Cuba (Groschaft and Del Valle, 1969; Zdzitowiecki and Rutkowska, 1980); Santa Clara, Cuba (Odening, 1969); Golden Grove Cave and St. Claire Cave, Jamaica (Webster, 1971); Convento de Acolman, Estado de Me´xico, Mexico, and Cueva Xictli, Mexico City, Mexico (Caballero, 1942); Cueva de Chilibrillo, Panama (Caballero and Grocott, 1960); Montevideo, Uruguay (Caballero and Caballero-Rodrı´guez, 1969); Alachua County and Marion County, Florida, U.S.A. (Foster and Mertins, 1996); Iowa, U.S.A. (Blankespoor and Ulmer, 1970); Crawford, Kansas, U.S.A. (Nickel and Hansen, 1967); Kansas, U.S.A. (Ubelaker, 1966); Albany, Baton Rouge, Hamond, Tangipahoa, and 3 km W of Erwinville, Louisiana, U.S.A. (Lotz and Font, 1991); New Orleans, Louisiana, U.S.A. (Martin, 1976); Bracken Cave, Texas, U.S.A. (Martin, 1976); Dodge County, Wisconsin, U.S.A. (Coggins et al.,

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COMPARATIVE PARASITOLOGY, 70(1), JANUARY 2003

1981, 1982); Eau Claire and Menomonie, Wisconsin, U.S.A. (Lotz and Font, 1991). Specimens deposited: CNHE 3876–3878. Other known specimens: USNPC 83851.

HWML

37527;

Remarks: The species Urotrema shillingeri Price, 1931, Urotrema lasiurensis Alicata, 1932, Urotrema minuta Macy, 1933, Urotrematulum attenuatum Macy, 1933, and Urotrema aelleni Baer, 1957 were declared junior synonyms of Urotrema scabridum by Caballero (1942) and Caballero and Grocott (1960) but were considered valid species by Yamaguti (1971). The elucidation of their taxonomic status either would extend the distribution range of U. scabridum or would establish an interesting pattern of subdistributions divided among a series of presumably closely related taxa. Cestoidea

Vampirolepis sp. Prevalence and mean abundance: Rı´o Salado, 2 of 12 hosts examined (16.6%; 0.4 6 1.2); Cueva de la Boca, 1 of 27 hosts examined (3.7%, 0.04 6 0.2); Concepcio´n del Oro, 2 of 28 hosts examined (7.1%; 0.07 6 0.3). Site of infection: Intestine.

Specimens deposited: CNHE 3857–3863. Remarks: The specimens we collected closely resembles members of the genera Anoplostrongylus Boulenger, 1926 and Tricholeiperia Travassos, 1935, but conform with no described species. Arthropoda Mesostigmata

Chiroptonyssus robustipes (Ewing, 1925) Fonseca, 1948 (Syn. Liponyssus robustipes Ewing, 1925; Liponyssus chilensis Ewing, 1925; Liponyssus nyctinomius Radford, 1938; Liponyssus venezolanus Hoffmann, 1944; Chiroptonyssus texensis Augustson, 1945; Hirstionyssus chilensis Fonseca, 1948; Ichoronyssus robustipes Strandtmann and Hunt, 1951). Prevalence and mean abundance: Nombre de Dios, 31 of 31 hosts examined (100%, 39.8 6 25.5); Rı´o Salado, 12 of 12 hosts examined (100%, 95.5 6 70.8); Cueva de la Boca, 27 of 27 hosts examined (100%, 39.1 6 28.3); Concepcio´n del Oro, 28 of 28 hosts examined (100%, 186.2 6 157.8); 28/28 (100%) (Foster and Mertins, 1996). Site of infestation: Dorsal and ventral body and wing, dorsal tail membrane.

Specimens deposited: CNHE 3879–3882. Remarks: Two species of Vampirolepis Spassky, 1954 have been reported from Ta. b. mexicana: Vampirolepis decipiens (Diesing, 1850), (Rogers, 1965: unpublished thesis, Oklahoma State University, Stillwater, Oklahoma, U.S.A.), and Vampirolepis gertschi Macy, 1947, (Cain, 1966; Martin, 1976). However, our material did not conform to any described species, although is similar to Vampirolepis chiropterophila Pe´rezVigueras, 1941 and Vampirolepis macroti Zdzitowiecki and Rutkowska, 1980. Nematoda

Anoplostrongylinae gen. sp. Prevalence and mean abundance: Nombre de Dios, 5 of 31 hosts examined (16.1%; 0.22 6 0.6); Rı´o Salado, 6 of 12 hosts examined (50%; 1.4 6 1.7); Cueva de la Boca, 5 of 27 hosts examined (18.5%, 1.2 6 3.2); Concepcio´n del Oro, 10 of 28 hosts examined (35.7%; 1 6 2.5). Site of infection: Intestine.

Type host: Tadarida brasiliensis mexicana. Reported hosts: Tadarida brasiliensis mexicana, (Ewing, 1925; Augustson, 1945; Randolph and Eads, 1946; Eads et al., 1957; Davis et al., 1962; Radovsky, 1967; Dooley et al., 1976; Bassols, 1981; Morales-Malacara, 1996, 1998: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico; Ta. brasiliensis, (Ewing, 1925; Hoffmann, 1944: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico; Fonseca, 1948; Jameson, 1959; Bradshaw and Ross, 1961; Yunker and Radovsky, 1966; Mauri, 1967, 1982; Radovsky, 1967; Dusba´bek, 1969a, 1970b; Saunders, 1975; Whitaker and Easterla, 1975; Bassols, 1981; Pence et al., 1981; Wilkins, 1989; Durden et al., 1992; Morales-Malacara, 1996, 1998: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico; Spears et al., 1999); Na. stramineus, (Hoffmann, 1944: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico); Ta. b. cynocephala, (Morlan and Strandtmann,

´ N-CORNEJO ET AL.—PARASITES OF TADARIDA BRASILIENSIS MEXICANA GUZMA

1949; Morlan, 1952; Radovsky, 1967); Myot. lucifugus, (Morlan and Strandtmann, 1949; Radovsky, 1967); Antrozous pallidus pallidus, (Herreid, 1961); Myot. nigricans, (Yunker and Radovsky, 1966; Radovsky, 1967); Ta. b. muscular, (Silva, 1965; Dusba´bek, 1969a); Nyctin. laticaudatus, (Dusba´bek, 1969a; Silva, 1979); Nyctinomops macrotis, (Fonseca, 1948; Radovsky, 1967; Whitaker and Easterla, 1975; Bassols, 1981; Milner et al., 1990); Sturnira ludovici, (Saunders, 1975); Mormoops megalophylla, (Whitaker and Easterla, 1975); Myotis californicus californicus, (Dooley et al., 1976); Tadarida sp., (Radovsky, 1967; Bassols, 1981; Morales-Malacara, 1996, 1998: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico); Ep. fuscus, (Radovsky, 1967; Durden et al., 1992); Myot. chiloensis, (Mauri, 1982); Artibeus toltecus (syn. Dermanura tolteca), (Morales-Malacara, 1996, 1998: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico). Locality records: Co´rdoba, La Rioja, Mar del Plata, Patquı´a, and Tucuma´n, Argentina (Mauri, 1967; Bassols, 1981); Buenos Aires City, Buenos Aires, Salta, San Luis and San Juan Provinces, Argentina (Mauri, 1982); Butantan, Sao Paulo, Brazil (Fonseca, 1948; Radovsky, 1967; Bassols, 1981); San Jose´ de la Montan˜a, Heredia, Costa Rica (Radovsky, 1967); Cuba (Silva, 1979); La Municio´n, Gupeyal, Cuba (Dusba´bek, 1970b; Bassols, 1981); Camagu¨ey, Cuba (Silva, 1965); Cairije, Camagu¨ey, Trinidad, Las Villas, and Pinar del Rio, Cuba (Dusba´bek, 1969a); Chile (Ewing, 1925; Dusba´bek, 1969a); Tepoztla´n, Morelos, Mexico (Hoffmann, 1944: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico; Bassols, 1981; MoralesMalacara, 1996, 1998: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico); Island of Dominica, Lesser Antilles (Pence et al., 1981; Wilkins, 1989); Malvern, Jamaica (Fonseca, 1948; Dusba´bek, 1969a; Bassols, 1981); Omiltemi, Guerrero, Mexico (Bassols, 1981; Morales-Malacara, 1996, 1998: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico); Tlilapan, Veracruz, Mexico (Morales-Malacara, 1996, 1998: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico); Cerro Punta, Panama (Yunker and Radovsky, 1966; Radovsky, 1967); Alabama, U.S.A. (Radovsky, 1967); Auburn, Lee

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County, Alabama, U.S.A. (Durden et al., 1992); Cochise County, Arizona, U.S.A. (Bradshaw and Ross, 1961; Radovsky, 1967); Berkeley, Alameda County, Conn Calley, Napa County, near Forestville and Asti, Sonoma County, Sacramento, Sacramento County, Davis, Yolo County, La Grange, Stanishus County, Pioneer, Amador County, Newcastle, Placer County, and Wilbur Springs, Colusa County, California, U.S.A. (Radovsky, 1967); White Springs, Hamilton County, Florida, U.S.A. (Radovsky, 1967); Alachua County, Florida, U.S.A. (Foster and Mertins, 1996); Georgia, U.S.A. (Morlan and Strandtmann, 1949); ‘‘southwest’’ Georgia, U.S.A. (Morlan, 1952); Tomas County, and Brooks County, Georgia, U.S.A. (Radovsky, 1967); Merrihew Cave, Aetna, Kansas, U.S.A. (Radovsky, 1967); Carlsbad Caverns, New Mexico, U.S.A. (Jameson, 1959); Don˜a Ana County, New Mexico, U.S.A. (Dooley et al., 1976); Vickery Cave, Bouse Junction; Selmans Cave, Mooreland, and Woods County, Oklahoma, U.S.A. (Radovsky, 1967); Texas, U.S.A. (Davis et al., 1962; Milner et al., 1990); Fort Sam Houston, Bexar County, Texas, U.S.A. (Augustson, 1945); Lavaca County, Texas, U.S.A. (Randolph and Eads, 1946); Crosbyton and Ney Cave, Medina County, and Ubalde County, Texas, U.S.A. (Jameson, 1959); Camp Mystic, Kerrville County, Texas, U.S.A. (Herreid, 1961); Rock Springs; Justiceberg; Fort Sam Houston; confluence of Rı´o Grande and Pecos rivers; Los Lingos Canyon; Haltville; Kingsville; Clyde Reed; Ney Cave, Medina County, Texas, U.S.A. (Radovsky, 1967); Big Bend National Park, Texas, U.S.A. (Whitaker and Easterla, 1975); El Paso County, Texas, U.S.A. (Dooley et al., 1976); Utah, U.S.A. (Tipton and Saunders, 1971; Whitaker and Wilson, 1974); Me´rida, 4 km E Tabay (La Mucuy), Venezuela (Saunders, 1975). Specimens deposited: JMM361, 390, 479, 573. Other known specimens: RML63562, 63566.

Macronyssus unidens Radovsky, 1967 Prevalence and mean abundance: Nombre de Dios, 1 of 31 hosts examined (3.2%, 0.03 6 0.2). Site of infestation: Ventral body. Type host: Myotis velifer.

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Reported hosts: Leptonycteris nivalis, Myot. lucifugus, Ep. fuscus, Ep. f. fuscus, Ep. f. pallidus, Corynorhinus townsendii (syn. Plecotus townsendii), (Radovsky, 1967; Reisen et al., 1976); Myot. velifer, (Radovsky, 1967; Reisen et al., 1976; Fitch et al., 1981); Co. t. pallescens, Lasionycteris noctivagans, (Turner, 1974); Pi. subflavus, (Reisen et al., 1976); Leptonycteris curasoae (syn. Leptonycteris sanborni), (Webb and Loomis, 1977); Corynorhinus mexicanus (syn. Plecotus mexicanus), (Morales-Malacara and Lo´pez-W, 1990; Tumlison, 1992; Morales-Malacara, 1996, 1998: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico). Locality records: El Tu´nel, Tlaxco, Tlaxcala, Mexico (Morales-Malacara and Lo´pez-W, 1990; Tumlison, 1992; Morales-Malacara, 1996, 1998: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico). Arizona, U.S.A. (Webb and Loomis, 1977); Crystal Cave, Catalina Mountains, Pinal County, Arizona, U.S.A. (Radovsky, 1967); La Salle County, Illinois, U.S.A. (Radovsky, 1967); Leavenworth and Ellsworth, Ellsworth County, Kansas, U.S.A. (Radovsky, 1967); Caves on Roundtop Mountain, Hancock, Maryland, U.S.A. (Radovsky, 1967); Patterson, Wayne County, Missouri, U.S.A. (Radovsky, 1967); Mercury, Nye County, and Pine Nut Mountains, Douglas County, Nevada, U.S.A. (Radovsky, 1967); Reed, Harmon County, Oklahoma, U.S.A. (Radovsky, 1967; Reisen et al., 1976; Fitch et al., 1981); Dark Canyon, Pennington County, South Dakota, U.S.A. (Turner, 1974); Hardeman and Armstrong County, Texas, U.S.A. (Radovsky, 1967); Dyer’s Cave, Wardentown, West Virginia, U.S.A. (Radovsky, 1967). Specimens deposited: JMM553. Metastigmata

Antricola sp. Prevalence and mean abundance: Rı´o Salado, 1 of 12 hosts examined (8.3%, 0.1 6 0.3). Site of infestation: Dorsal wing. Specimens deposited: JMM580. Remarks: Hoffmann and Lo´pez-Campos (2000) record only 3 species of Antricola Cooley and Kohls, 1942 from Mexico: Antricola mexicanus Hoffmann, 1958, Antricola coprophilus (McIntosh, 1935), and Antricola marginatus (Banks,

1910); however, the poor condition of the collected larvae precluded specific determination.

Ornithodoros kelleyi Cooley and Kohls, 1941 Prevalence and mean abundance: Nombre de Dios, 3 of 31 hosts examined (9.7%, 1 6 0.5). Site of infestation: Ventral body. Type host: Pipistrellus sp. Reported hosts: Myot. l. lucifugus, (Anastos and Clifford, 1956); Ep. fuscus, (Kohls and Ryckman, 1962; Kohls et al., 1965; Whitaker and Wilson, 1971; Wilson and Baker, 1972; Anonymous, 1988; Bonilla et al., 2001); A. pallidus, (Bradshaw and Ross, 1961; Knight and Marr, 1983; Bonilla et al., 2001); A. p. pacificus, Ep. f. pallidus, Myotis subulatus subulatus, (Kohls et al., 1965); Pipistrellus hesperus, (Kohls et al., 1965; Bonilla et al., 2001); Myot. lucifugus, (Whitaker and Wilson, 1971); Nyctic. humeralis, (Jackson and Goddard, 1995); Myot. velifer, Myotis thysanodes, Co. townsendii, (Bonilla et al., 2001). Locality records: Lethbridge, Alberta, and Dundurn, Saskatoon and Waldek, Canada (Wilkinson et al., 1980); Saskatchewan, Canada (Sonenshine and Anastos, 1960; Bonilla et al., 2001); Sonora, Mexico (Bonilla et al., 2001); Saltpeter Cave, Jackson County, Alabama, U.S.A. (Kohls et al., 1965); Continental, Pima County, Arizona, U.S.A. (Bradshaw and Ross, 1961); Oak Grove, Graham County, and Mohave County, Arizona, U.S.A. (Kohls et al., 1965); War Eagle, Benton County, Arkansas, U.S.A. (Kohls et al., 1965); San Jose and near Livermore, California, U.S.A. (Kohls et al., 1965); Colorado, U.S.A. (Sonenshine and Anastos, 1960); Morganton, Fannin County and Clayton, Rabun County, Georgia, U.S.A. (Wilson and Baker, 1972); ‘‘southern New England,’’ U.S.A. (Main, 1979); Illinois, U.S.A. (Sonenshine and Anastos, 1960; Anonymous, 1988); Indiana, U.S.A. (Sonenshine and Anastos, 1960); Donnehue’s Cave, Lawrence County, Indiana, U.S.A. (Whitaker and Wilson, 1971); Iowa, U.S.A. (Sonenshine and Anastos, 1960); Nelson County, Kentucky, U.S.A. (Cilek and Knapp, 1992); Pike County, Louisiana, U.S.A. (Palmer and Gunier, 1975); Louisiana, U.S.A. (Kohls et al., 1965); Chaptico, Maryland, U.S.A. (Anastos and Clifford, 1956); Tishomingo County, Mississippi, U.S.A. (Jack-

´ N-CORNEJO ET AL.—PARASITES OF TADARIDA BRASILIENSIS MEXICANA GUZMA

son and Goddard, 1995); Ann Arbor, Washtenaw County, Michigan, U.S.A. (OConnor and Klompen, 1988); Minnesota, U.S.A. (Sonenshine and Anastos, 1960); Swanson’s Cave, Waynesville, Missouri, U.S.A (Kohls et al., 1965); Fort Benton and near Mayers, Treasure County, Montana, U.S.A. (Kohls et al., 1965); Mercury, Nevada, U.S.A (Kohls et al., 1965); New Mexico, U.S.A. (Bonilla et al., 2001); New York, U.S.A. (Sonenshine and Anastos, 1960); Ohio, U.S.A. (Sonenshine and Anastos, 1960); Pennsylvania, U.S.A. (Sonenshine and Anastos, 1960); near Wall, South Dakota, U.S.A (Kohls et al., 1965); Douglas County, Washington, U.S.A. (Knight and Marr, 1983); Wisconsin, U.S.A. (Sonenshine and Anastos, 1961); West Virginia, U.S.A. (Sonenshine and Anastos, 1962); Utah, U.S.A. (Sonenshine and Anastos, 1960); Childress, Texas, U.S.A (Kohls et al., 1965). Specimens deposited: JMM427. Other known specimens: CDA5389; HK880202-1; RML19857, 21677, 23010, 24329, 32585, 32595, 35137, 37736, 37820-21, 3783035, 37888, 119910; USNTC123071. Remarks: This species has been recorded more frequently in association with other bat species (Kohls et al., 1965) including Ep. fuscus, which was found in sympatry with Ta. b. mexicana in Durango. Prostigmata

Ewingana (Doreyana) inaequalis (Ewing, 1938) Radford, 1948 (Syn. Radfordia inaequalis (Ewing, 1938) Dusba´bek, 1968) Prevalence and mean abundance: Nombre de Dios, 8 of 31 hosts examined (26%, 0.5 6 1.2); Rı´o Salado, 7 of 12 hosts examined (58.3%, 1.6 6 2.3); Cueva de la Boca, 9 of 27 hosts examined (33.3%, 1.1 6 2.2); Concepcio´n del Oro, 16 of 28 hosts examined (57.1%, 2 6 2.3); 1/28 (3.5%) (Foster and Mertins, 1996). Site of infestation: Ventral body, dorsal tail membrane, ear, nose. Type host: Tadarida brasiliensis cynocephala (syn. Tadarida cynocephala). Reported hosts: Tadarida brasiliensis cynocephala, (Ewing, 1938; Foster and Mertins, 1996); Ta. b. muscula, (Dusba´bek, 1968a); A. pallidus

17

(Lavoipierre and Beck, 1970); Ta. b. mexicana, (Beck, 1960 in Paran, 1992). Locality records: Camagu¨ey, Cuba (Dusba´bek, 1968a); California, U.S.A (Beck, 1960 in Paran, 1992; Lavoipierre and Beck, 1970); Alachua County, Florida, U.S.A (Foster and Mertins, 1996); Leon County, Florida, U.S.A (Ewing, 1938). Specimens deposited: JMM358, 549, 571, 577. Other known specimens: USNM1282; USDA9416222.

Ewingana (Mormomyobia) longa (Ewing, 1938) n. comb. (Syn. Myobia longa Ewing, 1938; Acanthophthirius longa (Ewing, 1938); Ewingana longa (Ewing, 1938) Dusba´bek, 1968; Ewingana (Doreyena) longa (Ewing, 1938) Dusba´bek, 1968; Ewingana (Ewingana) longa of Foster and Mertins, 1996). Prevalence and mean abundance: Nombre de Dios, 8 of 31 hosts examined (42%, 1.03 6 1.6); Rı´o Salado, 7 of 12 hosts examined (50%, 1.75 6 2.5); Cueva de la Boca, 17 of 27 hosts examined (63%, 2 6 2.1); Concepcio´n del Oro, 19 of 28 hosts examined (68%, 2 6 3.8); 1/28 (3.5%) (Foster and Mertins, 1996). Site of infestation: Dorsal and ventral body. Type host: Tadarida brasiliensis mexicana. Reported hosts: Tadarida brasiliensis mexicana, (Ewing, 1938; Jameson, 1959); Ta. brasiliensis, (Du´sbabek, 1969b; Durden et al., 1992); Ta. b. cynocephala, (Foster and Mertins, 1996). Locality records: Camagu¨ey, Cuba (Du´sbabek, 1969b); Lee County, Alabama, U.S.A. (Durden et al., 1992); Berkeley, California, U.S.A. (Ewing, 1938); Frio Cave, Uvalde County, Texas, U.S.A. (Jameson, 1959); Alachua County, Florida, U.S.A. (Foster and Mertins, 1996). Specimens deposited: JMM364, 387, 548, 566. Other known specimens: USNM1283, USDA9416226. Remarks: This species was originally described as Myob. longa Ewing, 1938 and later transferred by Dusba´bek (1968b) to Ewingana Radford, 1948 within the subgenus Doreyana Dusba´bek, 1968 on the basis of tarsal claw mor-

18

COMPARATIVE PARASITOLOGY, 70(1), JANUARY 2003

phology. However, Uchikawa (1978) discussed the morphology of the members of the subgenus Mormomyobia Fain, 1973, noting that tarsal claw formula is a highly variable character that accurately distinguishes subgenera. Uchikawa (1978) established the presence of a voluminous tubular process as one of the diagnostic characteristics of Mormomyobia. Ewing (1938) superficially noted this character in his original description of Myob. longa. The voluminous tubular process is readily observed in our material, and thus we hereby assign Myob. longa to the subgenus Mormomyobia as Ewingana (Mormomyobia) longa (Ewing, 1938) n. comb.

Leptotrombidium mexicana (Ewing, 1937) (Syn. Trombicula mexicana Ewing, 1937; Trombicula trombicula mexicana Thorn and Willmann, 1947; Trombicula leptotrombidium mexicana Wharton and Fuller, 1952; Chiroptella oudemansidium mexicana of VercammenGrandjean and Langston, 1971). Prevalence and mean abundance: Nombre de Dios, 4 of 31 hosts examined (13%, 0.5 6 1.5); Cueva de la Boca, 4 of 27 hosts examined (15%, 1 6 3.5); Concepcio´n del Oro, 5 of 28 hosts examined (8%, 0.2 6 0.6).

Whartonia (Asolentria) sp. Prevalence and mean abundance: Nombre de Dios, 1 of 31 hosts examined (3.2%, 0.4 6 2.1); Rı´o Salado, 5 of 12 hosts examined (41.7%, 4 6 6.7). Site of infestation: Dorsal and ventral wing. Specimens deposited: JMM362, 389. Remarks: Our specimens exhibit a unique suite of features suggesting an as yet undescribed species. The material resembles Whartonia (Asolentria) carpenteri (Brennan, 1962) but differs in the ramification of sensilas (Brennan, 1962). Astigmata

Dentocarpus macrotrichus Dusba´bek and Cruz, 1966 Prevalence and mean abundance: Nombre de Dios, 1 of 31 hosts examined (3.2%, 0.03 6 0.2); Rı´o Salado, 3 of 12 hosts examined (25%, 8.9 6 29.9); Cueva de la Boca, 4 of 27 hosts examined (14.8%, 0.3 6 0.9); Concepcio´n del Oro, 1 of 28 hosts examined (3.7%, 0.5 6 2.8); 1/28 (3.5%) (Foster and Mertins, 1996). Site of infestation: Dorsal and ventral body, dorsal wing.

Site of infestation: Dorsal and ventral wing.

Type host: Tadarida brasiliensis muscula.

Type host: Bat (specific identity undetermined).

Reported hosts: Tadarida brasiliensis muscula, (Dusba´bek and Cruz, 1966); Nyctin. macrotis, (Cruz and Abreu, 1984); Ta. b. cynocephala, (Foster and Mertins, 1996).

Reported hosts: Myotis velifer, (Bradshaw and Ross, 1961). Locality records: San Luis Potosi, Mexico (Ewing, 1937); Pima County, Arizona, U.S.A. (Bradshaw and Ross, 1961). Specimens deposited: JMM368, 478, 575.

Locality records: Camagu¨ey, Cuba (Dusba´bek and Cruz, 1966); Oriente and Santiago de Cuba, Cuba (Cruz and Abreu, 1984); Alachua County, Florida, U.S.A. (Foster and Mertins, 1996).

Other known specimens: RML35205, USNM1259.

Specimens deposited: JMM359, 385, 554, 563.

Whartonia sp.

Other known specimens: RMNH, USDA9416226.

Prevalence and mean abundance: Concepcio´n del Oro, 1 of 28 hosts examined (3.6%, 0.03 6 0.19). Site of infestation: Ventral wing. Specimens deposited: JMM486. Remarks: This material exhibited traits characteristic of each of the subgenera Whartonia Asolentria and Whartonia whartonia; thus, additional specimens are required to make clear subgeneric and specific determinations.

Notoedres (Notoedres) sp. Prevalence and mean abundance: Cueva de la Boca, 2 of 27 hosts examined (7.4%, 0.1 6 0.3). Site of infestation: Ventral wing. Specimens deposited: JMM568-569. Remarks: The subgenus Notoedres (Notoedres) (Railliet, 1893) is represented by 32 species of which our material most closely resembles No-

´ N-CORNEJO ET AL.—PARASITES OF TADARIDA BRASILIENSIS MEXICANA GUZMA

toedres (Notoedres) yunkeri Fain, 1962. However, our specimens (2 females) exhibit traits that suggest an as yet undescribed species, but confident determination and description is precluded until additional specimens, including males, can be acquired.

Notoedres (Bakeracarus) lasionycteris (Boyd and Bernstein, 1950) Fain, 1965 (Syn. Sarcoptes lasionycteris Boyd and Bernstein, 1950; Teinocoptes lasionycteris Yunker, 1958; Bakeracarus lasionycteris Fain, 1959). Prevalence and mean abundance: Nombre de Dios, 8 of 31 hosts examined (25.8%, 0.74 6 1.9); Rı´o Salado, 7 of 12 hosts examined (58.3%, 2.91 6 3.1); Cueva de la Boca, 14 of 27 hosts examined (51.8%, 2.5 6 3.5); Concepcio´n del Oro, 6 of 28 hosts examined (21.4%, 0.25 6 0.5). Site of infestation: Dorsal and ventral body, dorsal and ventral wing, and dorsal tail membrane. Type host: Lasionycteris noctivagans. Reported hosts: Lasionycteris noctivagans, (Boyd and Bernstein, 1950; Whitaker et al., 1975); Corynorhinus rafinesquii (syn. Plecotus rafinesquii), Myot. l. lucifugus, (Yunker, 1958); Ep. fuscus, Mol. molossus (syn. Mol. m. tropidorhynchus), Nyctin. laticaudatus, Pt. macleayii, Tadarida sp., Ta. b. muscula, (Dusba´bek, 1970a); Eptesicus brasiliensis (syn. Eptesicus melanopterus), (Fain and Lukoschus, 1971); Mol. ater, Mol. molossus, (Fain and Lukoschus, 1975). Locality records: Camagu¨ey, Isla de Pinos, La Habana, Las Villas and Oriente, Cuba (Dusba´bek, 1970a); Lelydorp, Surinam (Fain and Lukoschus, 1971); Meerzog, Moeroekreek, Wageningen, Weg naar Zee, Surinam (Fain and Lukoschus, 1975); Hibernia, New Jersey, U.S.A. (Yunker, 1958); North Carolina, U.S.A. (Whitaker et al., 1975); Jefferson County, Pennsylvania, U.S.A. (Boyd and Berstein, 1950); Shepherdstown and Pendleton, West Virginia, U.S.A. (Yunker, 1958). Specimens deposited: JMM359, 385, 554, 563. Other known specimens: RMNH, USNM1884. Remarks: Several subspecies have been assigned to No. b. lasionycteris since its original description by Boyd and Bernstein (1950). According

19

to Klompen (1992), it is possible additional material will reveal a series of species now hidden within this subspecies complex.

Nycteriglyphus bifolium Strandtmann, 1962 Prevalence and mean abundance: Cueva de la Boca, 15 of 27 hosts examined (55.5%, 11.1 6 18.6). Site of infestation: Dorsal and ventral body. Type host: Tadarida brasiliensis. Reported hosts: Tadarida brasiliensis, (Strandtmann, 1962). Locality records: Frio Cave, Uvalde County, Texas, U.S.A. (Strandtmann, 1962). Specimens deposited: JMM358, 388, 480, 495. Other known specimens: NHNM.

Olabidocarpus nyctinomus Fain, 1976 Prevalence and mean abundance: Rı´o Salado, 2 of 12 hosts examined (16.6%, 15 6 51.6); Cueva de la Boca, 8 of 27 hosts examined (29.6%, 5.07 6 17.1). Site of infestation: Dorsal and ventral body. Type host: Nyctinomops laticaudatus. Reported hosts: Nyctinomops laticaudatus, (Fain, 1976, 1982). Locality records: Sapucay, Paraguay (Fain, 1976, 1982). Specimens deposited: JMM358, 388, 480, 495. Other known specimens: BMNH 2.4.7.53-54. Diptera

Trichobius leionotus Wenzel, 1976 Prevalence and mean abundance: Cueva de la Boca, 2 of 27 hosts examined (7.4%, 0.1 6 0.3). Site of infestation: Dorsal body. Type host: Mormoops megalophylla. Reported hosts: Mormoops megalophylla, (Wenzel, 1976; Whitaker et al., 1987; Guerrero, 1994; Guerrero and Morales-Malacara, 1996); Pteronotus davyi, (Wenzel, 1976; Guerrero and Morales-Malacara, 1996). Locality records: La Paz, Ecuador (Wenzel, 1976); San Antonio de Pichincha, Ecuador

20

COMPARATIVE PARASITOLOGY, 70(1), JANUARY 2003

(Guerrero, 1994); Alta Vera Paz, Guatemala (Wenzel, 1976); Cueva Tzinacanoztoc, Puebla, and Cueva Arroyo Bellaco, Veracruz, Mexico (Guerrero and Morales-Malacara, 1996); Mt. Tamana Cave, Trinidad (Wenzel, 1976); Frio Cave, Uvalde County, Texas, U.S.A. (Wenzel, 1976); Big Bend National Park, Texas, U.S.A. (Wenzel, 1976; Whitaker et al., 1987); Bolivar, Falco´n, Sucre, and Yaracuy, Venezuela (Wenzel, 1976); Carabobo, Venezuela (Guerrero, 1994). Specimens deposited: JMM492, 505. Other known specimens: FMNH 64961-83, 65140-65; JMM-Pdav061. Remarks: The streblid fly Tr. leionotus is commonly associated with Mor. megalophylla, which cooccurred with the Mexican free-tailed bat in Cueva de la Boca, Nuevo Leo´n state. Siphonaptera

1944: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico); Arizona, California, and Oklahoma, U.S.A. (Lewis, 1985); Gainesville, Leon, and Tallahassee, Florida, U.S.A. (Fox, 1940); Southest Georgia, U.S.A. (Morlan, 1952); Thomasville, Georgia, U.S.A. (Fox, 1940); Barber County, Kansas (Linley et al., 1994); White Pine County, Nevada, U.S.A. (Egoscue, 1988); Eddy County, New Mexico, U.S.A. (Ewing, 1940); Carlsbad Caverns, New Mexico, U.S.A. (Jameson, 1959); Owl Cave, Woodward County, Oklahoma, U.S.A. (Palmer and Gunier, 1975); South Carolina, U.S.A. (Benton, 1980 in Lewis and Lewis, 1994); Big Bend National Park, Texas, U.S.A. (Whitaker and Easterla, 1975); Crosbyton and Ney Cave, Medina County, and Frio Cave, Uvalde County, Texas, U.S.A. (Jameson, 1959); Pecos, Reeves County, Texas, U.S.A. (Fox, 1914); Utah, U.S.A. (Stark, 1959 in Lewis and Lewis, 1994).

Sternopsylla distincta texana (Fox, 1914)

Specimens deposited: JMM425, 556, 565.

(Syn. Ischnopsyllus texanus Fox, 1914; Sternopsylla texana (Fox, 1914); Aptilopsylla carlsbadensis Ewing, 1940; Sternopsylla carlbadensis Ewing, 1940).

Other known specimens: NMNH18459, 7085.

Prevalence and mean abundance: Nombre de Dios, 5 of 31 hosts examined (16.1%, 0.2 6 0.5); Cueva de la Boca, 5 of 27 hosts examined (18.5%, 0.4 6 0.9); Concepcio´n del Oro, 5 of 28 hosts examined (18%, 0.3 6 0.7). Site of infestation: Dorsal and ventral body. Type host: Tadarida brasiliensis mexicana (syn. Nyctinomus mexicanus). Reported hosts: Tadarida brasiliensis mexicana, (Fox, 1914; Hoffmann, 1944: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico; Eads et al., 1957; Jameson, 1959; Egoscue, 1988); Ta. brasiliensis, (Fox, 1940; Palmer and Gunier, 1975; Whitaker and Easterla, 1975; Lewis, 1985; Ayala-Barajas et al., 1988; Linley et al., 1994); Ta. b. cynocephala, (Fox, 1940; Morlan, 1952); Myot. austroriparius, (Fox, 1940; Lewis, 1985); Pt. parnellii (syn. Chilonycteris rubiginosa mexicana) and Lepton. nivalis, (Ayala-Barajas et al., 1988). Locality records: Bosque de Chapultepec, Mexico City, Convento de Tepoztla´n, Morelos, and Cerro Potosi, Nuevo Leo´n, Mexico (Ayala-Barajas et al., 1988); Michoaca´n, Mexico (Johnson, 1957); El Fuerte, Sinaloa, Mexico (Hoffmann,

DISCUSSION A total of 11,857 parasites were collected from the 98 free-tailed bats sampled. Of the 11,857 parasites recovered 1,918 are helminths. Helminths are represented by 3 major taxa: Digenea (3 species), Cestoidea (1), and Nematoda (1). Three of the 5 helminth taxa (Oc. labda, Di. rileyi, and Anoplostrongylinae gen. sp.) were found in free-tailed bats from all 4 localities sampled. All helminth species recovered from Ta. b. mexicana were adults. Among the helminths, Oc. labda reached the highest prevalence (61.3%), whereas U. scabridum had the lowest (3.2%), both from hosts collected in the locality of Nombre de Dios, Durango state. The highest abundance was observed for Oc. labda in Durango, Puebla, and Zacatecas states; however, the values obtained in the locality of Puebla state are skewed by a single host infected with 795 specimens of Oc. labda. Accordingly, Oc. labda was the only helminth taxon that exhibited significant differences in mean abundance among localities (H 5 20.2101, P , 0.05). A total of 9,938 arthropod specimens representing 16 taxa were collected. Of these, 9,911 specimens are members of the subclass Acari, whereas only 27 are insects. The 32 new locality records presented in this study extend the known geographic distributions of all 16 arthropod taxa.

´ N-CORNEJO ET AL.—PARASITES OF TADARIDA BRASILIENSIS MEXICANA GUZMA

Chiroptonyssus robustipes and S. d. texana are previously recorded from Ta. b. mexicana (Hoffmann, 1944: unpublished thesis, Facultad de Ciencias, UNAM, Me´xico D.F., Mexico; Hoffmann and Lo´pez-Campos, 2000). The remaining 14 arthropod taxa recovered represent new records for Ta. b. mexicana in Mexico. Fifteen of the 16 taxa are true bat parasites, whereas Nycte. bifolium is considered a commensal species commonly occuring in guano in the caves inhabited by bats (Fain, 1988). The highest arthropod species richness was found in Mexican free-tailed bats from Nuevo Leo´n and Durango states (11 and 10 taxa, respectively) followed by Puebla and Zacatecas states (8 taxa each). Only 5 species of mites were shared by hosts from all 4 localities: Co. robustipes, Ew. d. inaequalis, Ew. m. longa, Dermanura macrotrichus, and No. b. lasionycteris. In contrast, Notoedres Notoedres sp., Nycte. bifolium, Macron. unidens, Or. kelleyi, Antricola sp., and Whartonia sp. were all collected exclusively from single hosts and localities. The streblid fly Tr. leionotus was recovered only from Nuevo Leo´n state, whereas the siphonapteran S. d. texana was found on bats from 3 of the 4 collection localities. Only 5 of the 16 arthropod taxa recovered were exclusively represented by adult forms (Macron. unidens, De. macrotrichus, Notoedres Notoedres sp., S. d. texana, and Tr. leionotus). Five taxa were represented only by larval stages (Or. kelleyi, Antricola sp., Leptot. mexicana, Whartonia Asolentria sp., and Whartonia sp.). The other 6 taxa were represented by mixed metamorphic specimens, i.e., some combination of larvae, nymphs, and adults. The arthropod infestation levels varied among localities, although Ch. robustipes and No. b. lasionycteris were the only species that exhibited significative differences in mean abundance (H 5 24.939, and H 5 14.214, P , 0.05, respectively). Particularly, Ch. robustipes exhibited the highest prevalence and abundance among arthropod parasites in all 4 collection localities. Prevalence was 100% and mean abundance ranged from 39.1 at Cueva de la Boca, Nuevo Leo´n state, to 186.2 at Concepcio´n del Oro, Zacatecas state. Our data is consistent with those of Foster and Mertins (1996), who reported Ch. robustipes in 100% of the Ta. b. cynocephala specimens examined. Protonymphs were the most abundant metamorphic stage reported in

21

their study (Foster and Mertins, 1996). Radovsky (1967) noted that the protonymph is the most abundant metamorphic stage infesting hosts but molts off the host, increasing the potential spread of the infestation among host animals. These observations suggest ready transmission of Ch. robustipes, consistent with its observed high prevalence and abundance. The lowest prevalence and mean abundance levels among arthropod parasites were observed for Macron. unidens from Nombre de Dios, Durango state (3.2% and 0.032, respectively). Foster and Mertins (1996) reported parasitic helminth and arthropod communities from the Brazilian free-tailed bat Ta. b. cynocephala in Florida with species richness and taxonomic compositions very similar to those presented in this study. Foster and Mertins (1996) reported 19 species of parasites (11 helminths and 8 mites), whereas we found 21 species (5 helminths, 12 mites, 2 ticks, and 2 insects). We only recovered less than half the number of helminth taxa recovered by Foster and Mertins (1996), but 3 of the 5 species we recovered are reported from both Ta. b. cynocephala and Ta. b. mexicana, whereas representatives Vampirolepis and Anoplostrongylinae are reported from both subspecies of bats, suggesting that Ta. b. mexicana hosts a robust sample of the larger Ta. b. cynocephala helminth parasite community. Differences among observed helminth communities might be environmentally driven. Our samples were collected in arid regions of Mexico, a habitat much less suited to the completion of helminth life cycles (especially digeneans) than the humid environment of Florida. Interestingly, the mite Ch. robustipes was recovered from all hosts examined in both surveys. In general, the parasites of Ta. b. mexicana we sampled seem to exhibit the ‘‘core’’ fauna (in a biogeographical sense) of this host. For a parasite taxon to be considered part of a biogeographical core, it must not only be widely distributed but also must be associated with and restricted to a monophyletic group of species, even if it is not present in all host species of that group (see Choudhury and Dick [1998]; Pe´rezPonce de Leo´n and Choudhury [2002]). With the exception of Macron. unidens, Or. kelleyi and Tr. leionotus, which represent either instances of host-sharing through ecological host extension or just an accidental infestation, the helminth and arthropod taxa recovered are typically found

22

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in all subspecies of Ta. brasiliensis, with a high degree of host specificity. Additional survey of metazoan parasites of the Mexican free-tailed bat in environments different from the arid habitats of Mexico may reveal a consistent parasitic fauna across the bat’s distributional range. This may be particularly true for ectoparasitic taxa, whose transmission is favored by a direct life cycle. ACKNOWLEDGMENTS We thank the following persons for their help during field and laboratory work: Laura del Castillo, Ada Ruı´z, Sau´l Aguilar, Reyna Lara, Roxana Acosta, Jorge Servı´n, Gerardo Lo´pez, Gerardo Rivas, Eduardo Catala´n, and Fernando Garcı´a; Livia Leo´n for her help in bat taxonomic identification; Agustı´n Jime´nez and Georgina Ortega Leite for providing us some bibliography; Florencia Bertoni for statistical advise. This work was funded by the Programa de Apoyo a Proyectos de Investigacio´n e Innovacio´n Tecnolo´gica (PAPITT-UNAM) grant No. IN215796 and IN214599 to J.B.M.-M. and IN219198 to G.P.-P.L. LITERATURE CITED Anastos, G., and C. M. Clifford. 1956. The occurrence of an Ornithodoros tick in Maryland. Journal of Parasitology 42:146. Anonymous. 1988. Fumigation cancels bat tick infestation. Pest Control 56:72–78. Augustson, G. F. 1945. A new genus, new species of Dermanyssid mite (Acarina) from Texas. Bulletin of the South California Academy of Sciences 44: 46–47. Ayala-Barajas, R., J. C. Morales-Mucin˜o, N. Wilson, J. E. Llorente-Bousquets, and H. E. PonceUlloa. 1988. Cata´logo de pulgas (Insecta: Siphonaptera). Serie de Cata´logos del Museo de Zoologı´a ‘‘Alfonso L. Herrera,’’ I. Universidad Nacional Auto´noma de Me´xico, Me´xico, D.F. 102 pp. Bassols, I. 1981. Cata´logo de los a´caros Mesostigamata de mamı´feros de Me´xico. Anales de la Escuela Nacional de Ciencias Biolo´gicas 24:9–49. Blankespoor, H. D., and M. J. Ulmer. 1970. Helminths from six species of Iowa Bats. Iowa Academy of Sciences 77:200–206. Bonilla, S. D., L. A. Durden, and W. Ll. Gannon. 2001. Tick (Acari) infestations of Bats in New Mexico. Journal of Medical Entomology 38:609– 611. Boyd, E. M., and M. H. Bernstein. 1950. A new species of sarcoptic mite fom a bat. Entomological Society of Washington 52:95–99. Bradshaw, G. V. R., and A. Ross. 1961. Ectoparasites of Arizona bats. Journal of the Arizona Academy of Sciences 1:1–10.

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