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TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)Copyright © 2013 Magnolia Press
Zootaxa 3722 (2): 183–203
www.mapress.com/zootaxa/Article
http://dx.doi.org/10.11646/zootaxa.3722.2.4
http://zoobank.org/urn:lsid:zoobank.org:pub:0620F7A3-C8DD-4AAF-B178-31B462A7DE02
A checklist of sucking lice (Insecta: Phthiraptera: Anoplura) associated with
Mexican wild mammals, including geographical records and a host-parasite list
SOKANI SÁNCHEZ-MONTES1, CARMEN GUZMÁN-CORNEJO1,3, LIVIA LEÓN-PANIAGUA2 & GERARDO
RIVAS1
1Laboratorio de Acarología, Departamento de Biología Comparada; Facultad de Ciencias, Universidad Nacional Autónoma de Méx-
ico. Avenida Universidad 3000, Ciudad Universitaria. C. P. 04510. Distrito Federal, México.
E-mail: [email protected]; [email protected] de Zoología, Departamento de Biología Evolutiva; Facultad de Ciencias, Universidad Nacional Autónoma de México.
Avenida Universidad 3000, Ciudad Universitaria. C. P. 04510. Distrito Federal, México3Corresponding author
Abstract
A checklist of 44 species of sucking lice (Insecta: Phthiraptera: Anoplura) recorded in Mexico, belonging to nine genera
in six families is given, together with a list of the 63 species of Mexican wild mammal hosts with which they are associ-
ated. Summaries of the known geographical records and host relationships for each louse species are presented for each
Mexican state. Data were compiled from published and original records, including three new locality records from the
states of Oaxaca and Guerrero.
Key words: sucking lice, Anoplura, Mexico, checklists, hosts, mammals, geographical records
Introduction
Sucking lice (Insecta: Phthiraptera: Anoplura) are obligate hematophagous ectoparasites of eutherian mammals. They are small, wingless, dorsoventrally flattened hemimetabolous insects, with a conical head that is narrower than the thorax, and with mouthparts that are modified for sucking blood (solenophagous). Additional anopluran adaptations for ectoparasitic living include reduction or absence of eyes (in most species), reduction of antennal segments (3–5), and the development of grasping tibio-tarsal claws. The suborder Anoplura comprises approximately 550 species classified in 16 families and 49 genera (Durden & Lloyd 2009; Light et al. 2010).
Sucking lice are important vectors of microorganisms that cause a variety of diseases, including tularemia, epidemic typhus, louse-borne relapsing fever, trench fever, and murine mycoplasmosis (Durden & Lloyd 2009).
A number of papers published between 1919 and 1986 include important records of Mexican Anoplura (Ferris 1919–1923; Mooser et al. 1931; Rubin 1946; Werneck 1948; Ferris 1951; Werneck 1952; Morlan & Hoff 1957; Johnson 1962; Kim 1965, 1966; Emerson 1971; Kim & Adler 1982; Kim et al. 1986). Whitaker & Morales-Malacara (2005) compiled a catalogue of ectoparasites associated with Mexican mammals, in which they listed 40 species of Anoplura, representing three families (Enderleinellidae, Hoplopleuridae and Polyplacidae) and 6 genera (Enderleinellus, Fahrenholzia, Hoplopleura, Linognathoides, Neohaematopinus and Polyplax) from 50 species of mammals. Recently, Light & Hafner (2007) analyzed the phylogenetics and host associations of members of the genus Fahrenholzia, providing new species records and localities for Mexico, as well as important molecular data from the specimens they studied. Because the anopluran literature is diffuse and often difficult to access, the objective of this study has been to compile all host and distribution records of Anoplura known from Mexico, in order to further elucidate host-parasite relationships and map the geographical distribution of each louse species in Mexico.
Accepted by R. Palma: 16 Sept. 2013; published: 22 Oct. 2013 183
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Material and methods
Our checklist is derived in part from an exhaustive search of the biological literature, utilizing such specialized databases as Biological Abstracts, Biological Sciences, BioOne, and Zoological Record.
This work is divided into three main sections. The first is a checklist of all the louse species arranged taxonomically by family and alphabetically by genus and species within each family. The second section is a louse-host list for each Mexican state, arranged alphabetically. Under each state, louse names appear in boldface, followed by host and locality data, published references (in parentheses), and specimen depositories (in brackets). We use “ND” where data are unavailable. New sucking louse records are presented in a separate paragraph in the following order: louse name, developmental stage (♂ = male, ♀ = female, N = nymph), host, locality, date of collection, and collection number. Additional information (e.g. taxonomic comments) appears in notes directly below a state’s listing. We use the following acronyms for collections where louse specimens have been deposited: CPFC (Facultad de Ciencias, Universidad Nacional Autónoma de México, Mexico City); FMNH (Field Museum of Natural History, Illinois, Chicago, U.S.A.); UCB (University of California, Berkeley, California, U.S.A.); UMC (University of Minnesota Entomology Collection, St. Paul, Minnesota, U.S.A.); USNM (National Museum of
Natural History, Washington, D.C., U.S.A.). Holotype specimens are indicated by a superscript H in boldface following the institutional acronym.
The last section of this work is a host-louse list, arranged alphabetically by mammal orders, families, genera and species. Names of hosts have been updated according to Ramírez-Pulido et al. (2005) and Wilson & Reeder (2005). A list of localities (Locality index) contains geographical coordinates obtained from the “Catalogue of
localities, municipalities and federative entities of Mexico” from the National Institute of Geography and Statistics (INEGI 2012), and electronic gazetteers such as GeoNames. Distribution maps for five anopluran families known from Mexico are included, prepared with the aid of the ArcView 3.2 program (ESRI 2000).
Results
Checklist of Lice
We record 44 species of sucking lice from Mexico, belonging to eight genera in five families, as follows:
Echinophthiriidae Enderlein 1904(Fig. 1)
Antarctophthirus Enderlein, 1906Antarctophthirus microchir (Trouessart & Neumann, 1888)
Enderleinellidae Ewing, 1929 (Fig. 1)
Enderleinellus Fahrenholz, 1912Enderleinellus arizonensis Werneck, 1948Enderleinellus deppei Kim, 1966Enderleinellus extremus Ferris, 1919Enderleinellus hondurensis Werneck, 1948Enderleinellus mexicanus Werneck, 1948Enderleinellus nayaritensis Kim, 1966Enderleinellus oculatus Kim, 1966Enderleinellus osborni Kellogg & Ferris, 1915Enderleinellus pratti Kim, 1966Enderleinellus suturalis (Osborn, 1891)
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Hoplopleuridae Ewing, 1929(Fig. 2)
Hoplopleura Enderlein, 1904Hoplopleura acanthopus (Burmeister, 1839)Hoplopleura arizonensis Stojanovich & Pratt, 1961aHoplopleura difficilis Kim, 1965Hoplopleura emphereia Kim, 1965Hoplopleura ferrisi Cook & Beer, 1959Hoplopleura hesperomydis (Osborn, 1891)Hoplopleura hirsuta Ferris, 1916Hoplopleura quadridentata (Neumann, 1909)Hoplopleura reithrodontomydis Ferris, 1951Hoplopleura similis Kim, 1965Hoplopleura trispinosa Kellogg & Ferris, 1915
Linognathidae Webb, 1946(Fig. 1)
Linognathus Enderlein, 1905Linognathus vituli (Linnaeus, 1758)
Polyplacidae Fahrenholz, 1912(Fig. 3)
Fahrenholzia Kellogg & Ferris, 1915Fahrenholzia boleni McDaniel, 1968Fahrenholzia ehrlichi Johnson, 1962Fahrenholzia ferrisi Werneck, 1952Fahrenholzia hertigi Johnson, 1962Fahrenholzia microcephala Ferris, 1922Fahrenholzia pinnata Kellogg & Ferris, 1915Fahrenholzia reducta Ferris, 1922Fahrenholzia texana Stojanovich & Pratt, 1961bFahrenholzia zacatecae Ferris, 1922
Linognathoides Cummings, 1914
Linognathoides laeviusculus (Grube, 1851)Linognathoides marmotae (Ferris, 1923)Linognathoides traubi (Rubin, 1946)
Neohaematopinus Mjöberg, 1910Neohaematopinus citellinus Ferris, 1942Neohaematopinus griseicolus Ferris, 1923Neohaematopinus neotomae Ferris, 1942Neohaematopinus sciuri Jancke, 1932Neohaematopinus sciurinus (Mjöberg, 1910)Neohaematopinus spilosomae Stojanovich & Pratt, 1961a
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FIGURE 1. Geographical distribution of Echinophthiriidae, Enderleinellidae and Linognathidae reported from Mexico.
FIGURE 2. Geographical distribution of Hoplopleuridae reported from Mexico.
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FIGURE 3. Geographical distribution of Polyplacidae reported from Mexico.
Polyplax Enderlein, 1904Polyplax alaskensis Ewing, 1927Polyplax auricularis Kellogg & Ferris, 1915Polyplax spinulosa (Burmeister, 1839)
Louse-host list by Mexican State
The geographic distributions of all anopluran species recorded from Mexico are shown on Figures 1–3.
BAJA CALIFORNIAAntarctophthirus microchir: Zalophus californianus, Lobera Los Cantiles, Isla Ángel de la Guarda, Golfo de
California (Godínez-Reyes et al. 1995).Linognathus vituli: Odocoileus hemionus fuliginatus, Unidad de Manejo y Conservación de la Vida Silvestre
(UMA) “El Tepi,” Sierra de San Pedro Mártir (Contreras et al. 2007).Neohaematopinus griseicolus: Sciurus griseus, ND (Kim et al. 1986).
Note: The record of L. vituli from O. hemionus fuliginatus is considered as an unusual host-louse association, because this louse species is a regular parasite of Bos taurus (see Durden & Musser 1994a).
BAJA CALIFORNIA SURFahrenholzia reducta: Chaetodipus baileyi, Miraflores (Emerson 1971).Neohaematopinus citellinus: Ammospermophilus leucurus extimus, San Antonio (Emerson 1971).
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COAHUILAFahrenholzia boleni: Perognathus merriami, Plan de Guadalupe (Light & Hafner 2007).Fahrenholzia pinnata: Dipodomys sp., Paila (Emerson 1971); Dipodomys merriami merriami, 2 km S Santa
Teresa (Light & Hafner 2007); Dipodomys nelsoni, 5 km S, 16 km W General Cepeda (Light & Hafner 2007); Dipodomys ordii, 3.2 km E Agua Nueva (Light & Hafner 2007).
Fahrenholzia zacatecae: Chaetodipus hispidus, 5 km S, 16 km W General Cepeda (Light & Hafner 2007).Hoplopleura arizonensis: Sigmodon toltecus, Parras (Emerson 1971); S. toltecus, La Paila (Emerson 1971); S.
toltecus, Paila-Parras (Emerson 1971).
COLIMAHoplopleura hirsuta: Xenomys nelsoni, Hacienda Magdalena (Ferris 1921).Neohaematopinus neotomae: Hodomys alleni [USNM 32706], Manzanillo (Ferris 1942).
Notes: The record of H. hirsuta from X. nelsoni is considered an unusual host-louse association, becasue this louse species has been recorded parasitizing rodents of the genus Sigmodon (Ferris 1951; Durden & Musser 1994a). Neohaematopinus neotomae was originally identified as Neohaematopinus sciurinus by Ferris (1923).
CHIAPASEnderleinellus extremus: Sciurus aureogaster nigrescens [USNM 75960], San Cristóbal (Ferris 1919).Enderleinellus hondurensis: Sciurus variegatoides goldmani [USNM 77906], Huehuetán (Werneck 1948).Hoplopleura emphereia: Peromyscus guatemalensis guatemalensis, Unión Juárez (Emerson 1971).Hoplopleura hirsuta: Sigmodon hirsutus, San Clemente (Emerson 1971); S. hirsutus, Tapachula (Emerson 1971).Polyplax auricularis: Peromyscus zarhynchus, Tzontehuitz (Emerson 1971).
Notes: The original record of E. extremus by Ferris (1919) did not include a collection number; however, Kim (1966) referred to the same locality and host, adding the USNM number that is given here. Enderleinellus
hondurensis was originally identified as Enderleinellus kelloggi Ferris, 1916 by Ferris (1919) [USNM 77906].Emerson (1971) originally identified H. emphereia as Hoplopleura ferrisi emphereia.
CHIHUAHUAEnderleinellus arizonensis: Sciurus nayaritensis apache [USNM 132347], Colonia García (Werneck 1948).Enderleinellus suturalis: Spermophilus madrensis, Sierra Madre (Ferris 1919); Spermophilus tereticaudus, El
Paso-Chihuahua City highway, about 55 km S Ciudad Juárez (Morlan & Hoff 1957). Fahrenholzia pinnata: Chaetodipus penicillatus, El Paso-Chihuahua City highway, about 55 km S Ciudad Juárez
(Morlan & Hoff 1957); Dipodomys. merriami, El Paso-Chihuahua City highway, about 55 km S Ciudad Juárez (Morlan & Hoff 1957); D. merriami, 9.6 km NW Ricardo Flores Magón (Light & Hafner 2007).
Hoplopleura hesperomydis: Onychomys leucogaster, El Paso-Chihuahua City highway, about 55 km S Ciudad Juárez (Morlan & Hoff 1957).
Hoplopleura hirsuta: Sigmodon ochrognathus [USNM 96268], Parral (Ferris 1921).Neohaematopinus citellinus: Spermophilus tereticaudus, El Paso-Chihuahua City highway, about 55 km S Ciudad
Juárez (Morlan & Hoff 1957).Neohaematopinus sciurinus: Sciurus nayaritensis [USNM 132347], Colonia García (Ferris 1923).
Notes: Specimens of E. arizonensis were originally identified by Ferris (1919) as Enderleinellus longiceps. Hoplopleura hesperomydis from O. leucogaster is considered an unusual host-louse association, because this louse species has been recorded parasitizing mainly rodents of the genus Peromyscus (Durden & Musser 1994a).
DISTRITO FEDERALEnderleinellus oculatus: Sciurus alleni [USNM 116931], Sierra de Guadalupe (Kim 1966).Hoplopleura reithrodontomydis: Reithrodontomys chrysopsis, Ajusco (Ferris 1951).Neohaematopinus sciurinus: S. alleni [USNM 11691], Sierra de Guadalupe (Ferris 1923).
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Polyplax auricularis: Neotomodon alstoni [USNM 50656], ND (Ferris 1923).Polyplax spinulosa: Rattus norvegicus, Cuauhtémoc (Mooser et al. 1931); Rattus rattus, Cuauhtémoc (Mooser et
al. 1931).
Notes: Sierra de Guadalupe encompasses two Mexican states: Distrito Federal and Estado de México. Specimens of E. oculatus were originally identified by Werneck (1948) as Enderleinellus arizonensis. Polyplax
auricularis from N. alstoni is considered an unusual host-louse association, because this louse species has been recorded parasitizing rodents of the genera Onychomys, Peromyscus and Reithrodontomys (Durden & Musser 1994a).
DURANGOFahrenholzia texana: Liomys irroratus, Hacienda Atotonilco (Light & Hafner 2007)Hoplopleura ferrisi: Peromyscus spicilegus, El Salto (Emerson 1971)Polyplax auricularis: Peromyscus sp., Ojo Tongo (Emerson 1971); Peromyscus maniculatus, Ojo Tongo (Emerson
1971); Peromyscus difficilis, Buenos Aires (Emerson 1971).
Note: Specimens of H. ferrisi were originally identified by Emerson (1971) as Hoplopleura ferrisi ferrisi.
ESTADO DE MÉXICOEnderleinellus oculatus: Sciurus alleni [USNM 116931], Sierra de Guadalupe (Kim 1966).Hoplopleura hesperomydis: Peromyscus melanotis, Nevado de Toluca, (Emerson 1971).Fahrenholzia pinnata: Dipodomys phillipsii [USNM 52036], Amecameca (Ferris 1922).Neohaematopinus sciurinus: S. alleni [USNM 11691], Sierra de Guadalupe (Ferris 1923).Polyplax auricularis: P. melanotis, Nevado de Toluca, (Emerson 1971).
Notes: Specimens of E. oculatus were originally identified by Werneck (1948) as Enderleinellus arizonensis.
Sierra de Guadalupe encompasses two Mexican states: Distrito Federal and Estado de México.
GUANAJUATOLinognathoides laeviusculus: Spermophilus mexicanus, ND (Ferris 1923).
Note: Linognathoides laeviusculus was placed in the genus Neohaematopinus by Ferris (1923).
GUERREROHoplopleura emphereia: Peromyscus megalops auritus, Omiltemi (Emerson 1971).Fahrenholzia microcephala: 3♂, 10♀, 2N, from Liomys pictus, Parque Estatal Cerro del Huizteco, Municipio de
Taxco, Guerrero, 30 Jun. & 4 Jul. 2010 [CPFC00001] New locality record.
Note: Hoplopleura emphereia was originally identified as Hoplopleura ferrisi emphereia by Emerson (1971).
HIDALGOFahrenholzia ehrlichi: Liomys irroratus alleni, Ixmiquilpan (Emerson 1971).
JALISCOFahrenholzia ehrlichi: Liomys irroratus jaliscensis, Etzatlán (Emerson 1971); L. irroratus alleni [USNM
120169], Acotlán (Johnson 1962); L. irroratus jaliscensis [USNM 34131], Atamejac (Johnson 1962); L.
irroratus jaliscensis, Encarnación de Díaz (Johnson 1962); L. irroratus, 16 km NNE Ameca (Light & Hafner 2007).
Fahrenholzia microcephala: Liomys pictus, 4.5 km SW Jilotlán (Light & Hafner 2007). Hoplopleura emphereia: Peromyscus sp., Ciudad Guzmán (Emerson 1971).Hoplopleura ferrisi: Peromyscus levipes, Zapotlanejo (Emerson 1971); P. levipes, Nevado de Colima (Emerson
1971).
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Hoplopleura hesperomydis: Peromyscus melanotis, Nevado de Colima (Emerson 1971).Polyplax alaskensis: Microtus mexicanus phaeus, Nevado de Colima (Emerso 1971).Polyplax auricularis: Peromyscus hylocetes, Nevado de Colima (Emerson 1971); P. melanotis, Nevado de Colima
(Emerson 1971).
Notes: Fahrenholzia ehrlichi was originally identified as F. microcephala [USNM 34131] by Ferris (1923). The locality given for F. ehrlichi – Acotlán – does not exist, but this name may refer to Acatlán de Juárez.Hoplopleura emphereia was originally identified by Emerson (1971) as Hoplopleura ferrisi emphereia and Hoplopleura ferrisi was originally identified by Emerson (1971) as Hoplopleura ferrisi ferrisi.
MICHOACÁN
Linognathoides traubi: Spermophilus adocetus [USNM 57684]H, ND (Rubin 1946).
Notes: Linognathoides traubi was originally placed in the genus Neohaematopinus traubi by Rubin (1946). Locality data was not given, but Johnson (1958) cited this specimen (with the same USNM number) from Apatzingán, Michoacán.
MORELOSEnderleinellus mexicanus: Sciurus aureogaster nigrescens [USNM 51156], Huitzilac (Werneck 1948).Hoplopleura difficilis: Peromyscus difficilis, Lagunas de Zempoala (L.A. Durden pers. comm. 2013).Hoplopleura ferrisi: Peromyscus sp., Lagunas de Zempoala (Emerson 1971).Neohaematopinus sciurinus: S. aureogaster nigrescens [USNM 51156], Huitzilac (Ferris 1923).Polyplax alaskensis: Microtus mexicanus mexicanus, Lagunas de Zempoala (Emerson 1971).
Notes: Specimens of E. mexicanus were originally identified by Ferris (1919) as Enderleinellus extremus
Hoploplura ferrisi was originally identified by Emerson (1971) as Hoplopleura ferrisi ferrisi
NAYARIT
Enderleinellus mexicanus: Sciurus colliaei [USNM 96795]H, Chacala (Werneck 1948).
Enderleinellus pratti: S. colliaei [USNM 91245]H, Santiago, Tepic (Kim 1966).Fahrenholzia microcephala: Liomys pictus hispidus, Tepic (Emerson 1971).Hoplopleura ferrisi: Peromyscus spicilegus, “Tepico” (Emerson 1971).Neohaematopinus sciurinus: S. colliaei [USNM 91245], Santiago, Tepic (Ferris 1923).
Notes: Specimens of E. mexicanus and E. pratti were originally identified by Ferris (1919) as Enderleinellus
extremus. The holotype, allotype and five paratypes of E. mexicanus are deposited in the UCB, and two paratypes are in the UMC. The holotype, allotype and five paratypes of E. pratti are deposited in the UCB, and one paratype in the UMC. Hoplopleura ferrisi was originally identified by Emerson (1971) as Hoplopleura
ferrisi ferrisi from “Tepico”, but this locality refers almost certainly Tepic, Nayarit.
NUEVO LEÓNFahrenholzia ehrlichi: Chaetodipus hispidus, San Antonio Allende (Ramos-Casillas 2003); C. penicillatus, San
Antonio Allende (Ramos-Casillas 2003).Hoplopleura acanthopus: Microtus mexicanus subsimus, El Potosí (Emerson 1971).Hoplopleura hirsuta: Sigmodon toltecus, Pesquería (Ramos-Casillas 2003).Linognathoides marmotae: Spermophilus variegatus couchii [USNM 57685], Municipie de Galeand, Ojo de Aqua
(Rubin 1946).Neohaematopinus neotomae: Neotoma leucodon, Doctor Arroyo (Ramos-Casillas 2003); Neotoma micropus,
Lampazos (Ramos-Casillas 2003).Polyplax alaskensis: M. mexicanus subsimus, El Potosí (Emerson 1971).Polyplax auricularis: Peromyscus maniculatus blandus, Cerro Potosí (Emerson 1971).
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Notes: Linognathoides marmotae from Municipie de Galeand, Ojo de Aqua, was originally described as Neohaematopinus mathesoni by Rubin (1946). Later, Johnson (1958) speculated that this locality was probably Municipio Galeana and Agua de Oro. However, Agua de Oro is a locality in the Municipality of Galeana (INEGI 2012).
OAXACAEnderleinellus deppei: Sciurus aureogaster aureogaster [USNM 73297], Juchitán (Kim 1966).Enderleinellus extremus: Sciurus aureogaster nigrescens [USNM 68182], Cerro San Felipe (Ferris 1919); S.
aureogaster aureogaster, Quichicón (Ferris 1919).Fahrenholzia texana: Liomys irroratus, ND (Kim et al. 1986).Hoplopleura emphereia: Megadontomys cryophilus, Cerro Pelón (Emerson,1971).Hoplopleura ferrisi: 5♂, 21♀ 1N, from Peromyscus melanurus, La Yerba Buena, Municipio de Santa Catarina
Juquila, Oaxaca, 30 Apr. and 5 May 2010 [CPFC00002] New locality record.Neohaematopinus sciurinus: S. aureogaster nigrescens [USNM 68182], Cerro San Felipe (Ferris 1923); S.
aureogaster aureogaster [USNM 73297], Quichicón, Oaxaca (Ferris 1923).Polyplax alaskensis: Microtus mexicanus fulviventer, Llano de las Flores (Emerson 1971).Polyplax auricularis: Reithrodontomys mexicanus [USNM 68685], “Tohontepec” (Ferris 1923).Polyplax auricularis: 18♂, 16♀ 17N, from Peromyscus beatae, Km 134.5 on road 175, 21 km N Guelatao,
Oaxaca, 22–25 Nov. 2010 [CPFC00003] New locality record.
Notes: Specimens of E. deppei were originally identified by Ferris (1919) as E. extremus. The original record of E.
extremus Ferris (1919) did not include a collection number; however, Kim (1966) referred to the same locality and host, adding the USNM number that is given here. Hoplopleura emphereia was originally identified by Emerson (1971) as Hoplopleura ferrisi emphereia Polyplax auricularis was recorded from “Tohontepec, Chiapas” by Ferris (1923), who subsequently (Ferris 1951) stated that this locality probably corresponded to “Tehuantepec, Oaxaca.”
PUEBLAEnderleinellus extremus: Sciurus aureogaster, ND (Kim 1966).Fahrenholzia ehrlichi: Liomys irroratus, 6 km N Tilapa (Light & Hafner 2007); L. irroratus, 11 km (by road) SW
Alchichica (Light & Hafner 2007); L. irroratus, 3 km (by road) NE Tilapa (Light & Hafner 2007)Fahrenholzia pinnata: Dipodomys phillipsii, 11 km (by road) SW Alchichica (Light & Hafner 2007); Perognathus
flavus, 3.1 km SW El Veladero (Light & Hafner 2007).
QUERÉTAROHoplopleura trispinosa: Glaucomys volans, Pinal de Amoles (L.A. Durden pers. comm. 2013).
SAN LUIS POTOSÍFahrenholzia ehrlichi: Liomys irroratus texensis, El Salto (Emerson 1971).Hoplopleura hirsuta: Sigmodon toltecus, El Salto (Emerson 1971).
SINALOAEnderleinellus nayaritensis: Sciurus nayaritensis apache, Santa Lucía (Kim 1966).Fahrenholzia microcephala: Liomys pictus hispidus, Matatán (Emerson 1971); L. pictus, San Ignacio (Emerson
1971); L. pictus, Santa Lucía (Emerson 1971).Hoplopleura ferrisi: Peromyscus spicilegus, ND (Emerson 1971).Hoplopleura hirsuta: Sigmodon arizonae, Los Mochis (Emerson 1971).Neohaematopinus neotomae: Hodomys alleni alleni, “Conception” (Emerson 1971).
Note: The correct locality record given by Emerson (1971) for N. neotomae is Concepción (La Barrigona).
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SONORAEnderleinellus osborni: Spermophilus tereticaudus neglectus, Kino (Emerson 1971).Fahrenholzia pinnata: Dipodomys deserti sonoriensis, Kino (Emerson 1971); D. merriami merriami, Kino
(Emerson 1971).
TABASCO
Enderleinellus deppei: Sciurus deppei [USNM 100048]H, Teapa (Kim 1966).Neohaematopinus sciurinus: S. deppei [USNM 100048], Teapa (Ferris 1923).
Note: Specimens of E. deppei were originally identified by Ferris (1919) as Enderleinellus extremus.
TAMAULIPASEnderleinellus extremus: Sciurus deppei negligens, Altamira (Ferris 1919).Fahrenholzia ehrlichi: Liomys irroratus, Matamoros (Johnson 1962).Hoplopleura hirsuta: Neotoma micropus, Matamoros (Ramos-Casillas 2003).Hoplopleura quadridentata: Oryzomys rostratus [USNM 92935], Altamira (Ferris 1921).
Note: Hoplopleura hirsuta from N. micropus is considered an unusual host-louse association, because this louse species has been recorded parasitizing rodents of the genus Sigmodon (Ferris, 1951; Durden & Musser 1994a).
VERACRUZEnderleinellus extremus: Sciurus aureogaster [USNM 93016], Papantla (Ferris 1919).
Enderleinellus oculatus: Sciurus oculatus [USNM 54235]H, ND (Kim 1966).
Fahrenholzia ferrisi: Heteromys desmarestianus [USNM 14353]H, Achotal (Werneck 1952); H. desmarestianus
[LSUMZ 36300], 8 km ENE Catemaco (Light & Hafner 2007).Fahrenholzia hertigi: H. desmarestianus, 8 km ENE Catemaco (Light & Hafner 2007).
Fahrenholzia microcephala: Liomys pictus pictus [FMNH 11099]H, San Carlos (Ferris 1922); L. pictus, Biological Station La Mancha (Light & Hafner 2007).
Hoplopleura similis: Oligoryzomys fulvescens [USNM 58259]H, Orizaba (Kim 1965).Neohaematopinus sciurinus: S. oculatus [USNM 54235], ND (Ferris 1923).
Notes: The original record of E. extremus Ferris, 1919 did not include a collection number; however, Kim (1966) referred to the same locality and host, adding the USNM number that is given here, but he gave the locality erroneously as “Papanita”. Specimens of E. oculatus were originally identified by Ferris (1919) as Enderleinellus longiceps. The type specimens of E. oculatus are deposited in the UCB. Fahrenholzia ferrisi
was originally identified as F. microcephala [USNM 14353]H by Ferris (1922). Specimens of H. similis were originally identified by Ferris (1919) as Hoplopleura hesperomydis and Hoplopleura quadridentata.
YUCATÁNEnderleinellus hondurensis: Sciurus yucatanensis, Gruta de “Balankranche” (Kim 1966).Fahrenholzia ferrisi: Heteromys gaumeri, Pisté (Emerson 1971).Hoplopleura arizonensis: Sigmodon hispidus, Pisté (Emerson 1971).
Note: The correct locality record for E. hondurensis is Gruta de Balankanché.
ZACATECAS
Enderleinellus nayaritensis: Sciurus nayaritensis [USNM 90947]H, Sierra Madre (Kim 1966).
Fahrenholzia ehrlichi: Liomys irroratus alleni [USNM 91883]H, Valparaíso (Johnson 1962), L. irroratus, 3.2 km E San Jerónimo (Light & Hafner 2007).
Fahrenholzia pinnata: Dipodomys phillipsii ornatus [USNM 91939], Valparaíso (Ferris 1922); D. ordii, 1.6 k m SE Bañon (Light & Hafner 2007); D. phillipsii, 3.2 km E San Jerónimo (Light & Hafner 2007).
Fahrenholzia zacatecae: Chaetodipus hispidus zacatecae [USNM 91875]H, Valparaíso (Ferris 1922).
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Notes: Specimens of E. nayaritensis were originally identified by Ferris (1919) as Enderleinellus longiceps, and subsequently as Enderleinellus arizonensis by Werneck (1948). The holotype, allotype and seven paratypes of E.
nayaritensis are deposited in the UCB. Fahrenholzia ehrlichi was originally identified as Fahrenholzia
microcephala [USNM 91883]H by Ferris (1922). The holotype and allotype specimens are deposited in the UCB.
MEXICO (no state given)Fahrenholzia microcephala: Liomys irroratus, ND (Ferris 1951).Neohaematopinus sciuri: Sciurus aureogaster, ND (Kim & Adler 1982).Neohaematopinus spilosomae: Spermophilus spilosoma, ND (Durden & Musser 1994a).
Host-louse List
The 44 species of Mexican anopluran lice are associated with 81 taxa (63 species) of mammals in three orders: Artiodactyla, Carnivora and Rodentia.
MAMMALIA
Artiodactyla
Cervidae
Odocoileus hemionus fuliginatus Cowan
Linognathus vituli
Carnivora
Otariidae
Zalophus californianus (Lesson)
Antarctophthirus microchir
Rodentia
CricetidaeHodomys alleni (Merriam)
Neohaematopinus sciurinus
Hodomys alleni alleni (Merriam)Neohaematopinus neotomae
Megadontomys cryophilus (Musser)Hoplopleura emphereia
Microtus mexicanus fulviventer MerriamPolyplax alaskensis
Microtus mexicanus mexicanus (de Saussure)Polyplax alaskensis
Microtus mexicanus phaeus (Merriam)Polyplax alaskensis
Microtus mexicanus subsimus GoldmanHoplopleura acanthopus
Polyplax alaskensis
Neotoma leucodon MerriamNeohaematopinus neotomae
Neotoma micropus BairdHoplopleura hirsuta
Neohaematopinus neotomae
Neotomodon alstoni MerriamPolyplax auricularis
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Oligoryzomys fulvescens (de Saussure)Hoplopleura similis
Onychomys leucogaster (Wied-Neuwied) Hoplopleura hesperomydis
Oryzomys rostratus MerriamHoplopleura quadridentata
Peromyscus beatae ThomasPolyplax auricularis
Peromyscus difficilis (Allen) Hoploplura difficilis
Polyplax auricularis
Peromyscus guatemalensis guatemalensis MerriamHoplopleura emphereia
Peromyscus hylocetes MerriamPolyplax auricularis
Peromyscus levipes MerriamHoplopleura ferrisi
Peromyscus maniculatus (Wagner)Polyplax auricularis
Peromyscus maniculatus blandus OsgoodPolyplax auricularis
Peromyscus megalops auritus MerriamHoplopleura emphereia
Peromyscus melanotis Allen & ChapmanHoplopleura hesperomydis
Polyplax auricularis
Peromyscus melanurus OsgoodHoplopleura ferrisi
Peromyscus spicilegus AllenHoplopleura ferrisi
Peromyscus zarhynchus MerriamPolyplax auricularis
Peromyscus Gloger sp.Hoplopleura emphereia
Hoplopleura ferrisi
Polyplax auricularis
Reithrodontomys chrysopsis MerriamHoplopleura reithrodontomyis
Reithrodontomys mexicanus (de Saussure)Polyplax auricularis
Sigmodon arizonae MearnsHoplopleura hirsuta
Sigmodon hispidus Say & OrdHoplopleura arizonensis
Sigmodon hirsutus (Burmeister) Hoplopleura hirsuta
Sigmodon toltecus (de Saussure)Hoplopleura arizonensis
Hoplopleura hirsuta
Sigmodon ochrognathus BaileyHoplopleura hirsuta
Xenomys nelsoni MerriamHoplopleura hirsuta
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HeteromyidaeChaetodipus baileyi (Merriam)
Fahrenholzia reducta
Chaetodipus hispidus (Baird)Fahrenholzia ehrlichi
Fahrenholzia zacatecae
Chaetodipus hispidus zacatecae (Osgood)Fahrenholzia zacatecae
Chaetodipus penicillatus (Woodhouse)Fahrenholzia ehrlichi
Fahrenholzia pinnata
Dipodomys deserti sonoriensis GoldmanFahrenholzia pinnata
Dipodomys merriamis merriami MearnsFahrenholzia pinnata
Dipodomys nelsoni MerriamFahrenholzia pinnata
Dipodomys ordii WoodhouseFahrenholzia pinnata
Dipodomys phillipsii GrayFahrenholzia pinnata
Dipodomys phillipsii ornatus MerriamFahrenholzia pinnata
Dipodomys Gray sp.Fahrenholzia pinnata
Heteromys desmarestianus GrayFahrenholzia ferrisi
Fahrenholzia hertigi
Heteromys gaumeri Allen & ChapmanFahrenholzia ferrisi
Liomys irroratus (Gray)Fahrenholzia ehrlichi
Fahrenholzia microcephala
Fahrenholzia texana
Liomys irroratus alleni (Coues)Fahrenholzia ehrlichi
Liomys irroratus jaliscensis (Allen)Fahrenholzia ehrlichi
Liomys irroratus texensis MerriamFahrenholzia ehrlichi
Liomys pictus (Thomas)Fahrenholzia microcephala
Liomys pictus pictus (Thomas)Fahrenholzia microcephala
Liomys pictus hispidus (Allen)Fahrenholzia microcephala
Perognathus flavus BairdFahrenholzia pinnata
Perognathus merriami AllenFahrenholzia boleni
MuridaeRattus norvergicus Berkenhout
Polyplax spinulosa
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Rattus rattus LinnaeusPolyplax spinulosa
SciuridaeAmmospermophilus leucurus extimus Nelson & Goldman
Neohaematopinus citellinus
Glaucomys volans (Linnaeus)Hoplopleura trispinosa
Sciurus alleni NelsonEnderleinellus oculatus
Neohaematopinus sciurinus
Sciurus aureogaster CuvierEnderleinellus extremus
Neohaematopinus sciuri
Sciurus aureogaster aureogaster CuvierEnderleinellus deppei
Enderleinellus extremus
Neohaematopinus sciurinus
Sciurus aureogaster nigrescens BennettEnderleinellus extremus
Enderleinellus mexicanus
Neohaematopinus sciurinus
Sciurus colliaei RichardsonEnderleinellus mexicanus
Enderleinellus pratti
Neohaematopinus sciurinus
Sciurus deppei PetersEnderleinellus deppei
Neohaematopinus sciurinus
Sciurus deppei negligens NelsonEnderleinellus extremus
Sciurus griseus OrdNeohaematopinus griseicolus
Sciurus nayaritensis AllenEnderleinellus nayaritensis
Neohaematopinus sciurinus
Sciurus nayaritensis apache AllenEnderleinellus arizonensis
Enderleinellus nayaritensis
Sciurus oculatus PetersEnderleinellus oculatus
Neohaematopinus sciurinus
Sciurus variegatoides goldmani NelsonEnderleinellus hondurensis
Sciurus yucatanensis AllenEnderleinellus hondurensis
Spermophilus adocetus (Merriam)Linognathoides traubi
Spermophilus madrensis (Merriam)Enderleinellus suturalis
Spermophilus mexicanus (Erxleben)Linognathoides laeviusculus
Spermophilus spilosoma BennettNeohaematopinus spilosomae
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Spermophilus tereticaudus BairdEnderleinellus osborni
Enderleinellus suturalis
Neohaematopinus citellinus
Spermophilus variegatus couchii BairdLinognathoides marmotae
Discussion
Studies of Mexican Anoplura from Ferris (1919) to the present day have yielded a total of 44 species associated with wild mammals, a figure that represents 8% of the world anopluran fauna (Light et al. 2010). In Mexico, sucking lice have been collected from 63 host species, representing 12% of the 525 mammal species known to occur in this country (Ceballos & Oliva 2005).
Currently, the state of Oaxaca has the greatest number (9) of anopluran species recorded, followed by Chihuahua, Jalisco, Nuevo León and Veracruz with 7, but no records exist from four other states (Aguascalientes, Campeche, Quintana Roo, and Tlaxcala).
The genus containing the highest number of species in Mexico is Hoplopleura (11 species) followed by Enderleinellus (10 species), Fahrenholzia (9), Neohaematopinus (6), Linognathoides and Polyplax (3 each), while only one species has been recorded from each of the genera Antarctophthirus and Linognathus. Geographically, the most widespread species is Neohaematopinus sciurinus recorded from 8 states, Followed by Fahrenholzia ehrlichi
and Polyplax auricularis from 7 states.Mexican Anoplura are associated with three orders of mammals – Artiodactyla, Carnivora, and Rodentia – the
latter accounting for the vast majority of host records: 61 species in 22 genera (Hodomys, Megadontomys, Microtus,Neotoma, Neotomodon, Olygoryzomys, Onychomys, Oryzomys, Peromyscus, Reithrodontomys, Sigmodon, Xenomys,Chaetodipus, Dipodomys, Heteromys, Liomys, Perognathus, Rattus, Amnospermophilus, Glaucomys, Sciurus andSpermophilus) and four families (Cricetidae, Heteromyidae, Muridae and Sciuridae). These hosts harbour 95.5% of the total number of anopluran species known from the country. This high percentage agrees with an observation by Durden (2001), who wrote that about two thirds of the world’s Anoplura parasitize rodents.
Among the anopluran species associated with Rodentia, those of the genus Fahrenholzia are specific to heteromyids, those of Hoplopleura mostly with murids, but also with cricetids and sciurids, and those of Enderleinellus and Linognathoides with sciurids. Species of the genera Neohaematopinus and Polyplax are more widespread parasites, the former being associated with cricetids and sciurids, while the latter are most often found on cricetids and murids (Durden & Musser 1994b). In this study, we provide three new locality and host records for three anopluran species (Fahrenholzia microcephala, Hoplopleura ferrisi and Polyplax auricularis), also associated with rodents.
Whitaker & Morales-Malacara (2005) list Glaucomys volans as a host of Hoplopleura trispinosa in Mexico based on Kim et al. (1986), but no Mexican records are mentioned in the latter publication. However, this louse species has been found in association with G. volans from Pinal de Amoles, Quéretaro, Mexico (L.A. Durden pers. comm. 2103). Hoplopleura trispinosa is considered a possible vector of Rickettsia prowazekii, the pathogen for epidemic typhus (McDade 1987). Currently, only four Mexican Anoplura – Hoplopleura acanthopus, Linognathus
vituli, Neohaematopinus sciuri and Polyplax auricularis – are potential vectors of human and animal pathogens. Many pathogenic agents have been detected or isolated from sucking lice, as follows: five bacteria: Bartonella sp. (see Durden et al. 2004; Tsai et al. 2010), Bartonella rattimassiliensis (see Reeves et al. 2006), Bartonella
tribocorum (see Reeves et al. 2006), Brucella brucei (see Parnas et al. 1960), and Haemobartonella muris (see Crystal 1958); two rickettsiae: Rickettsia prowazekii (see Mooser et al. 1931) and Rickettsia typhi (see Azad 1990); three protozoans: Anaplasma spp. (Hornok et al. 2010), Theileria sergenti (see Fujisaki et al. 1993) andTrypanosoma lewisi (see Khachoian & Arakelian 1978); and one fungus: Trichophyton verrucosum (see Kamyszek 1977). However, the transmission of these pathogens by lice remains unproven for most of them.
It is very important that Mexican ectoparasite specimens be deposited in national collections, where they will serve regional investigators. Earlier collectors deposited the type material for 12 (26.6%) Mexican species in United States collections (FMNH, UCB, UMC, USNM). Primary types deposited in the USNM were recently
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cataloged by Durden & Adams (2005), who listed only three types (USNM 57684, 57685, 91883) from Mexico. Although the types of other Mexican Anoplura are cited in the literature, we have not been able to locate them.
Our knowledge of Mexican sucking lice is still far from complete. Indeed, many of the principal hosts of these ectoparasites, i.e. rodents, still have not been sampled for specimens. It is therefore imperative that studies of these insects be encouraged so that a clearer picture of the taxonomy, ecology, biogeography and phylogenetics of Mesoamerican Anoplura may eventually emerge.
Acknowledgments
Partial funding for this research was provided by the Programa de Apoyo a Proyectos de Investigación e
Innovación Tecnológica (PAPIIT No 226010). We thank Griselda Montiel, Rosario Chavarría, Mirna Hernández, Diego Barrales, Luis Darcy, Christina Lynggaard, Ricardo Paredes, Héctor Olguín, Giovani Canchola and Pablo Colunga for their assistance in the field. Richard G. Robbins (Walter Reed Army Medical Center, Washington, D.C., U.S.A.) kindly reviewed and edited an earlier version of our manuscript. We are grateful to Prof. Lance Durden (Georgia Southern University��Statesboro��Georgia��U�S�A�) for his thorough review of our manuscript and for providing useful suggestions that have greatly improved the quality of this paper.
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Morlan, H.B. & Hoff, C.C. (1957) Notes on some Anoplura from New Mexico and Mexico. Journal of Parasitology, 43, 347–351. http://dx.doi.org/10.2307/3274362
Neumann, L.G. (1909) Notes sur les Pediculidés. Archives de Parasitologie, 13, 497–537.Osborn, H. (1891) The Pediculi and Mallophaga affecting man and lower animals. Bulletin of United States Department of
Agriculture Bureau of Entomology, 7, 1–56.Parnas, J., Zwolski, W. & Burdzy, K. (1960) Infection of lice Polyplax spinulosa and Hoplopleura acanthopus with Brucella
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SÁNCHEZ-MONTES ET AL. 200 · Zootaxa 3722 (2) © 2013 Magnolia Press
TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
Locality index
Locality Latitude N Longitude W
BAJA CALIFORNIA
Lobera Los Cantiles, Isla Ángel de la Guarda, Golfo de California 29°32′00″ 113°29′00″
Unidad de Manejo y Conservación de la Vida Silvestre (UMA) "El Tepi," Sierra de San Pedro Mártir
31°04′36″ 115°16′31″
BAJA CALIFORNIA SUR
Miraflores 23°22'08″ 109°46'47″
San Antonio 26°32′00″ 112°30′00″
COAHUILA
La Paila 25°39'01″ 102°08'44″
Paila 25°38′00″ 102°08′00″
Paila-Parras ND ND
Parras 25°25′00″ 102°11′00″
Plan de Guadalupe 26°12'33" 101°20'40"
2 km S Santa Teresa 26°22'47" 101°21'20"
3.2 km E Agua Nueva 25°11'02" 101°03'29"
5 km S, 16 km W General Cepeda 25°22′35″ 101°28′30″
COLIMA
Hacienda Magdalena ND ND
Manzanillo 19°03′08″ 104°18′57″
CHIAPAS
Huehuetán 15°01′00" 92°23′00"
San Clemente 16°22'14" 93°47'53"
San Cristóbal 16°44′12″ 92°38′18″
Tapachula 14°54′29″ 92°15′38″
Tzontehuitz 16°50′15″ 92°34′00″
Unión Juárez 15º04′00" 92°05′00"
CHIHUAHUA
El Paso-Chihuahua City highway about 55 km S Ciudad Juárez 31°44′22″ 106°29′13″
Colonia García 30°18′34″ 108°04′35″
Parral 26°55′57″ 105°39′47″
Sierra Madre ND ND
9.6 km NW Ricardo Flores Magón 29°59′20″ 107°01′41″
DISTRITO FEDERAL
Ajusco 19°12′00″ 99°16′00″
Cuauhtémoc 19°26′29.81″ 99°09′6.6″
Sierra de Guadalupe ND ND
DURANGO
Buenos Aires 25°44′57″ 105°02′54″
El Salto 23°46′42″ 105°21′37″
Haciendo Atotonilco 24°38′35″ 103°41′56″
Ojo Tongo ND ND
......continued on the next page
Zootaxa 3722 (2) © 2013 Magnolia Press · 201ANOPLURA OF MEXICAN WILD MAMMALS
TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
Locality index (Continued)
Locality Latitude N Longitude W
ESTADO DE MÉXICO
Amecameca 19°07′40″ 98°45′46″
Nevado de Toluca 19°06′30″ 99°45′30″
Sierra de Guadalupe ND ND
GUERRERO
Parque Estatal Cerro del Huizteco, Municipio de Taxco 18°36′08″ 99°36′30″
Omiltemi 17°35′39″ 99°38′46″
HIDALGO
Ixmiquilpan 20°28′55″ 99°13′05″
JALISCO
Acatlán de Juárez 20º14′30″ 103°32′30″
Atamejac 20°05′00″ 103°35′00″
Ciudad Guzmán 19°42′31″ 103°27′51″
Encarnación de Díaz 21°24′00″ 102°25′00″
Etzatlán 20°46′00″ 104°05′00″
Nevado de Colima 19°33′49″ 103°36′29″
Zapotlanejo 20°27′00″ 102°52′00″
4.5 km SW Jilotlán 19°37′00″ 103°02′00″
16 km NNE Ameca 20°55′00″ 104°03′00″
MICHOACÁN
Apatzingán 19°05′19″ 102°21′03″
MORELOS
Huitzilac 19°01′42″ 99°16′02″
Lagunas de Zempoala 19°03′13″ 99°18′54″
NAYARIT
Chacala 21°9′57″ 105°13′28″
Santiago 21°48′41″ 105°12′24″
Tepic 21°31'95″ 104°54′32″
NUEVO LEÓN
Municipio de Galena, Agua del Oro 24°50′00″ 100°04′00″
Cerro Potosí 24°52′18″ 100°13′57″
Doctor Arroyo 24°41′46″ 100°01′52″
El Potosí 24°49'35.23″ 100°20'30.98″
Lampazos 27°01′00″ 100°31′00″
Pesquería 25°47′06″ 100°03′04″
San Antonio Allende 25º17'00″ 100°01'00″
OAXACA
Cerro Pelón 17°01′30″ 96°04′46″
Cerro San Felipe 17°01′32″ 96°04′48″
Juchitán 16°26′00″ 94°56′50″
Km 134.5 road 175, 21 km N Guelatao 17°25'10" 96°29'53"
La Yerba Buena, Municipio de Santa Catarina Juquila 16°13'59" 97°16'59"
......continued on the next page
SÁNCHEZ-MONTES ET AL. 202 · Zootaxa 3722 (2) © 2013 Magnolia Press
TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
Locality index (Continued)
Locality Latitude N Longitude W
Llano de las Flores 17°26'50" 96°30'14"
Quichicón ND ND
Tehuantepec 16°19′28″ 95°14′20″
PUEBLA
3 km (by road) NE Tilapa 18°17′00″ 97°10′00″
3.1 km SW El Veladero 18°57′55″ 97°29′20″
6 km N Tilapa 18°35′38″ 98°33′14″
11 km (by road) SW Alchichica 19°26′09″ 97°23′20″
QUERÉTARO
Pinal de Amoles 21°08′03″ 99°27′31″
SAN LUIS POTOSÍ
El Salto 22°31'14" 101°03'49"
SINALOA
Concepción (La Barrigona) 23°23′39" 106°10′07"
Los Mochis 25°47′37" 108°59′49"
Matatán 23°01′58" 105°43′42"
San Ignacio 23°31′20" 105°44′45"
Santa Lucía 23°26′15" 105°51′10"
SONORA
Kino 30°37′37" 110°58′03"
Nuevo Kino 28°51'24" 112°0'26"
Locality Latitude N Longitude W
TABASCO
Teapa 17°32′00" 92°57′00"
TAMAULIPAS
Altamira 22°23′00" 97°56′00"
Matamoros 25°52′00" 97°30´00"
VERACRUZ
Achotal 17°44′10" 95°07′56"
Biological Station La Mancha 19°36′00" 96°22′00"
Orizaba 18°51′31" 97°05′48"
Papantla 20°26′56″ 97°19′28"
San Carlos 17°40′52" 94°42′24"
8 km ENE Catemaco 18°25′34″ 95°06′45″
YUCATÁN
Gruta de Balankanché 20°40′00″ 88°36′00″
Pisté 20°41′53″ 89°35′19″
ZACATECAS
Sierra Madre ND ND
Valparaíso 22°46′13″ 103°34′04″
1.6 km SE Bañon 23°09'51″ 102°27'56″
3.2 km E San Jerónimo 22°39'06″ 102°29'38″
Zootaxa 3722 (2) © 2013 Magnolia Press · 203ANOPLURA OF MEXICAN WILD MAMMALS