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Zootaxa 4623 (3): 485–525 ISSN 1175-5326 (print edition)

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Copyright © 2019 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4623.3.3
http://zoobank.org/urn:lsid:zoobank.org:pub:46B4D4BA-B826-4744-B5E7-865CB9ED8F27

The soft ticks (Parasitiformes: Ixodida: Argasidae) of Mexico: species, hosts,


and geographical distribution
CARMEN GUZMÁN-CORNEJO1, ANGEL HERRERA-MARES1,
RICHARD G. ROBBINS2 & ANDREA REBOLLO-HERNÁNDEZ1
1
Laboratorio de Acarología, Departamento de Biología Comparada, Facultad de Ciencias, Universidad Nacional Autónoma de Méxi-
co, Avenida Universidad 3000, Ciudad Universitaria, 04510 Ciudad de México, México
2
Walter Reed Biosystematics Unit, Department of Entomology, Smithsonian Institution, MSC, MRC 534, 4210 Silver Hill Road, Suit-
land, MD 20746, USA

Abstract

Distribution and host data from published literature and previously unpublished collection records are provided for all
37 species of soft ticks, family Argasidae, that are known to occur in Mexico, including Ornithodoros dugesi, which is
generally regarded as a junior synonym of Ornithodoros talaje, and Ornithodoros marinkellei, a new record for Mexico.
Parasite-host and host-parasite lists are presented, together with keys to adults and larvae, and a gazetteer of collection
localities supplemented by maps showing each species’ distribution in Mexico.

Key words: Argasidae, Mexico, hosts, distribution, keys

Introduction

The family Argasidae comprises approximately 211 species (Luz et al. 2018) that are commonly referred to as
“soft ticks” due to their lack of a scutum as nymphs and adults, although most argasid larvae possess a mid-dorsal
plate. As noted by Guglielmone et al. (2010) and Mans et al. (2019), several competing genus- and subgenus-level
classifications of the Argasidae currently coexist. Recent application of molecular techniques (Burger et al. 2014,
Mans et al. 2019) may eventually yield a unified phylogeny for this family, with predictive value for studies of tick
evolution and host and disease relationships. However, for the purpose of cataloging the argasid species of Mexico,
as well as listing their recorded hosts and describing their current distribution, we have found it convenient to retain
the well-known classification of Hoogstraal (1985), who recognized five argasid genera—Antricola, Argas, Notho-
aspis, Ornithodoros and Otobius—although we do not endorse this or any other classificatory scheme.
Particularly in Mexico, argasid ticks have been known since pre-Hispanic times, under a variety of aboriginal
names and dialects. The first researcher to formally work with these ticks was Alfredo Dugès, who described Argas
turicata (currently classified in the genus Ornithodoros) in 1876, Argas megnini (currently classified in the genus
Otobius) in 1884, and Argas sanchezi in 1887. In the 20th century, several investigators published observations on
the biology and distribution of this tick family in Mexico, e.g., Macías-Valadez (1923), Hoffmann (1930), McIntosh
(1935), and Mazzotti (1940, 1941, 1943a). Others contributed to our understanding of argasid ticks as vectors of
pathogens, e.g., Hoffmann (1930), Pilz & Mooser (1936), Brumpt et al. (1939), Mazzotti (1947, 1949), Silva-Goytia
& Elizondo (1952), and Mazzotti (1953). In her “Monografía de los Ixodoidea de Mexico” (1962), Hoffmann re-
viewed and consolidated our knowledge of the tick fauna of this country, recognizing a total of 14 species of Argasi-
dae (including Ornithodoros dugesi Mazzotti, 1943, which is often regarded as a junior synonym of Ornithodoros
talaje (Guérin-Méneville, 1849)). Hoffmann’s magnum opus set the baseline for future discoveries by national and
foreign investigators: Kohls et al. (1965, 1969), Dusbábek (1970), Jones et al. (1972), Keirans & Clifford (1975),
Endris et al. (1989), Hoffmann & López-Campos (2000), Guzmán-Cornejo et al. (2003), Hoffmann et al. (2004),
Nava et al. (2012) and Herrera et al. (2016), resulting in a total to date of 36 species. In this work we have compiled
all published information on the Argasidae of Mexico, providing new distribution and host records, including the

Accepted by O. Seeman: 9 May 2019; published: 26 Jun. 2019 485


first records for Ornithodoros marinkellei Kohls, Clifford and Jones, 1969, in this country, together with taxonomic
keys to the larvae and adults of all species.

Materials and methods

Bibliographic searches were conducted using a variety of databases, such as CAB Abstracts, ISI Web of Knowledge,
BioOne and Google Scholar, in order to obtain all known references to the soft ticks of Mexico and to confirm all
published tick collections. Records are presented alphabetically by tick genus and species and include: Mexican
state(s) (alphabetized, capitalized and in boldface), locality, host(s) and reference(s).
Within this framework, unavailable data are indicated as ND (Not Determined). Published records referring to
“Distrito Federal” have been updated to “Ciudad de México.” Where we have added new records, we include the
number of specimens, sex or stage, locality, host, date, and collection number (Coleccion Nacional de Ácaros, Insti-
tuto de Biología (CNAC), or Colección Laboratorio de Acarología, Facultad de Ciencias (LAFC), Universidad Na-
cional Autónoma de México (UNAM)). Host names follow Ramírez-Pulido et al. (2014) for mammals and Avibase
(Lepage 2014) for birds. All geographical coordinates for each locality are included in a table; where coordinates are
not available, we reference the coordinates of the nearest municipality, marked with an asterisk (*). Maps showing
the distribution of argasid species were constructed using QGIS Development Team (2018).

Results

We recognize a total of 37 argasid species representing five genera: Antricola (Antricola coprophilus, Antricola
marginatus, Antricola mexicanus), Argas (Argas brevipes, Argas cooleyi, Argas giganteus, Argas miniatus, Argas
persicus, Argas radiatus, Argas sanchezi), Nothoaspis (Nothoaspis reddelli), Ornithodoros (Ornithodoros azteci,
Ornithodoros brodyi, Ornithodoros capensis, Ornithodoros coriaceus, Ornithodoros denmarki, Ornithodoros du-
gesi, Ornithodoros dusbabeki, Ornithodoros dyeri, Ornithodoros hasei, Ornithodoros kelleyi, Ornithodoros knoxjo-
nesi, Ornithodoros kohlsi, Ornithodoros marinkellei, Ornithodoros mormoops, Ornithodoros nicollei, Ornithodoros
parkeri, Ornithodoros puertoricensis, Ornithodoros rossi, Ornithodoros setosus, Ornithodoros stageri, Ornithodo-
ros talaje, Ornithodoros turicata, Ornithodoros viguerasi, Ornithodoros yumatensis) and Otobius (Otobius lagophi-
lus, Otobius megnini).

Key to genera (Adults)

Note: The prominent sexual dimorphism seen in the hard tick family Ixodidae is absent in the Argasidae, adults of
which can only be separated with certainty by examining the genital aperture. In males, the anterior lip of the geni-
tal aperture is more highly chitinized than the posterior lip, appearing to resemble a human thumbnail, whereas in
females such chitinization is not seen, with the result that the two lips of the genital aperture resemble those of the
human mouth. Last-stage nymphs do not possess external genitalia, but a simple pregenital pore may be present. For
additional discussion and illustrations of argasid genitalia, see Nuttall (1921).

1 With a sutural line separating the dorsal and ventral surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argas (Fig. 1A)
- Lacking a sutural line separating the dorsal and ventral surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Integument granular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Otobius (Fig. 1B)
- Integument varied, mammillate, tuberculate or shield-like, but not granular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Dorsum with shield-like false scutum (Fig. 1C); palpal flaps (medial extensions of palpal segment 1 enclosing the hypostome)
present; associated with bats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nothoaspis (N. reddelli)
- Lacking false scutum and palpal flaps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Integument tuberculate; hypostome broad at base and scoop-like (Fig. 4A); associated with bats . . . . . . . . .Antricola (Fig. 1D)
- Integument mammillate; hypostome variously shaped, but denticulate and never scoop-like; associated with reptiles, birds and
mammals, including bats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Ornithodoros (Fig. 1E)

486 · Zootaxa 4623 (3) © 2019 Magnolia Press GUZMÁN-CORNEJO ET AL.


FIGURE 1. A) Argas persicus, female. B) Otobius megnini, female. C) Nothoaspis reddelli, male, dorsal idiosoma with false
scutum. D) Antricola mexicanus, male. E) Ornithodoros brodyi, female.

THE SOFT TICKS OF MEXICO Zootaxa 4623 (3) © 2019 Magnolia Press · 487
Key to species (Adults) (all measurements in millimeters)

Argas (adults of A. giganteus are unknown) (after Kohls et al. 1961, 1970)

1 Dorsal integument with peripheral ridge of small rounded tubercles, with scattered setae arising from indistinct pits or inter-
stices; postpalpal setae absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Dorsal integument with large peripheral rectangular cells, with one or two setiferous pits; postpalpal setae present . . . . . . . . 3
2 Hypostomal dentition apically 2/2, basally 3/3 to 4/4; legs IV not reaching posterior body margin when extended . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. brevipes
- Hypostomal dentition apically 2/2, basally 3/3 to 5/5; legs IV reaching posterior body margin when extended . . . . . A. cooleyi
3 Peripheral cells of dorsum mostly rectangular or subrectangular (maximum number of cells 100), often with pits bearing a
single seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. persicus
- Peripheral cells of dorsum more irregular (more than 150 peripheral cells), with pits bearing one or two setae . . . . . . . . . . . . 4
4 Mammillae conspicuously elevated, not crowded (dorsum verrucose); about 150 peripheral cells . . . . . . . . . . . . . . A. miniatus
- Mammillae not conspicuously elevated, crowded together with ridges (dorsum not verrucose); about 200 peripheral cells . . . .
...................................................................................................5
5 Dorsal integument relatively smooth due to uniform elevation of mammillae and ridges; legs short, not reaching posterior body
margin when extended . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .A. sanchezi (Fig. 2 A)
- Dorsal integument relatively coarse due to slightly greater elevation of mammillae among less elevated, variously shaped
ridges; legs long, reaching posterior body margin when extended . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. radiatus

Otobius (after Cooley & Kohls 1944)

1 Dorsal idiosomal integument with pits separated by two or more times their diameters . . . . . . . . . . . . . . O. megnini (Fig. 1B)
- Dorsal idiosomal integument with pits separated by no more than their diameters . . . . . . . . . . . . . . . . . O. lagophilus (Fig. 2B)

Antricola (based on original descriptions)

1 Dorsum of idiosoma surrounded by elongated digitiform tubercles bearing setae; female length: 7.6–9.0, male length: 7.2–8.3
(Černý, 1966) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. marginatus (Fig. 4B)
- Dorsum of idiosoma not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Tubercles with tufts of ciliated setae located on the posterior margin of the idiosoma and part of lateral walls below coxae IV;
stigmal plates foliaceous and cordiform, protruding from the body; female length: 7.8, male length: 4.5 (Hoffmann 1958) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. mexicanus (Fig. 1D, 4C)
- Lacking tufts of ciliated setae; stigmal plates suboval, not protruding from the body (Hoffmann 1958); female length: 5.6, male
length: 4.0 (McIntosh 1935) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. coprophilus

Ornithodoros (Adults of O. knoxjonesi, O. mormoops, O. rossi and O. setosus are unknown; O. capensis, O. den-
marki and O. dugesi are not included in this key due to a lack of characters to differentiate post-larval stages) (origi-
nal descriptions; Cooley & Kohls 1944; Guzmán-Cornejo et al. 2017; Nava et al. 2017)

1 Eyes present; mammillae reddish gray, almost iridescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. coriaceus (Figs. 2C-D)
- Eyes absent; mammillae not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Cheeks present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Cheeks absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Hood absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Hood present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4 Integument smooth; with large, elliptical spiracular plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .O. marinkellei
- Integument mamillated; spiracular plates not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .O. stageri (males)
5 With ventral sclerotized plates; cheeks atypically small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. viguerasi
- Without ventral sclerotized plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Subapical dorsal protuberance on tarsus IV small or absent; tarsus I with 3-4 dorsal humps . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- Subapical dorsal protuberance on tarsus IV conspicuous; tarsus I proximally with a single prominent dorsal hump . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. nicollei (Fig. 2E)
7 Mammillae large, relatively few in number, not crowded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .O. turicata (Fig. 2F)
- Mammillae small, numerous, and somewhat crowded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. parkeri (Fig. 2G)
8 Body elongate; associated with bats or bat roosts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Body oval; associated with various hosts, including bats or bat roosts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
9 Body with sides subparallel; dorsal surface bounded by two contiguous ridges . . . . . . . . . . . . . . . . . . . . . . . .O. dyeri (Fig. 2H)
- Body pyriform, broadly rounded posteriorly and gradually narrowing anteriorly; dorsal surface not bounded by contiguous
ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. azteci (Fig. 3A)

488 · Zootaxa 4623 (3) © 2019 Magnolia Press GUZMÁN-CORNEJO ET AL.


10 Hood absent; tarsus I without subapical dorsal protuberance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- Hood present; tarsus I with subapical dorsal protuberance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 Cheeks small; leg surfaces nearly smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. stageri (females)
- Cheeks large; leg surfaces micromammillated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. yumatensis (Figs. 4D, 4E)
12 Hood well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- Hood not well developed, small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
13 Large species (length/width: females: 6.0–11.4/3.0–6.1; males: 5.7–7.8/3.0–6.5 . . . . . . . . . . . . . . .O. brodyi (Figs. 1E, 3B, 4F)
- Small species (length/width: females: 3.4–4.1/2.0–2.63; males: 2.7–3.3/1.7–1.9) . . . . . . . . . . . . . . O. puertoricensis (Fig. 3B)
14 Small species (less than 5.0 in length) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. hasei
- Large species (more than 5.0 in length) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Dorsum with small discs; tarsus I with very small subapical dorsal protuberance . . . . . . . . . . . . . . . . . . . . .O. kelleyi (Fig. 3C)
- Dorsum with large discs; tarsus I with moderately developed subapical dorsal protuberance . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 Body pointed anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Body bluntly rounded anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .O. dusbabeki (Fig. 3D)
17 Discs large, occupying much of median area of dorsum . . . . . . . . O. talaje (Fig. 3E) (see discussion for additional comments)
- Discs smaller, occupying less extensive area of dorsum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. kohlsi (Fig. 3F)

(see discussion for additional comments)

Key to genera (Larvae) (all measurements in millimeters)


(after Kohls et al. 1965; Jones & Clifford 1972; Barros-Battesti et al. 2013)

1 Dorsum of idiosoma with 7–10 pairs of setae; venter with 5 pairs of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Otobius
- Dorsum of idiosoma with more than 10 pairs of setae; venter with 6–11 pairs of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Palpal segment IV as long as or longer than other palpal segments; dorsum with 26–30 pairs of setae . . . . . . . . . . . . . . . Argas
- Palpal segment IV shorter than palpal segments II and III; dorsum with 13–21 pairs of setae (except 27–29 pairs in Ornithodo-
ros setosus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Claws absent (except in Antricola marginatus), pulvilli extended (Fig. 5A); three pairs of postcoxal setae . . . . . . . . .Antricola
- Claws present, pulvilli reduced; one pair of postcoxal setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Dorsal plate in shape of isosceles triangle and extending most of length of dorsum; dorsum with 14 pairs of setae; apex of hy-
postome pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nothoaspis (N. reddelli)
- Dorsal plate subrectangular, pyriform or triangular; dorsum with 13–21 pairs of dorsal setae (except 27–29 pairs in Ornithodo-
ros setosus); apex of hypostome rounded or pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros

Key to species (Larvae) (all measurements in millimeters)

Argas (after Kaiser et al. 1964; Kohls et al. 1970; Jones & Clifford 1972; Lindquist et al. 2016)

1 Tarsus I with a trumpet-shaped sensillum extending posteriorly from the capsule of Haller’s organ . . . . . . . . . . . . . . . . . . . . . 2
- Tarsus I without a trumpet-shaped sensillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 With 23–24 pairs of dorsal idiosomal setae; tarsus I without premidventral setae . . . . . . . . . . . . . . . . . . . . . . . . Argas brevipes
- With about 32 pairs of dorsal idiosomal setae; tarsus I with 1 pair of premidventral setae . . . . . . . . . . . . . . . . . . . Argas cooleyi
3 Body when engorged exceptionally large, ca. 4.0 long by 2.6 wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argas giganteus
- Body when engorged not exceptionally large . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Tarsus I with 2 pairs of lateral setae and 3 pairs of ventral setae; posterolateral body setae ca. 0.100 long, almost twice as long
as anterolateral setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argas sanchezi
- Tarsus I with 1 pair of lateral setae and 2 pairs of ventral setae; posterolateral body setae not more than 1.5 times longer than
anterolateral setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Dorsal plate circular; dorsal body setae short, posterolaterals and anterolaterals nearly equal in length, posterolaterals average
ca. 0.039, maximum length ca. 0.047 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argas persicus
- Dorsal plate elongate oval . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Dorsal plate 0.200–0.240 long, 0.168–0.212 wide; posterolateral body setae average ca. 0.050 long; hypostome 0.160-0.180
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argas radiatus
- Dorsal plate 0.188–0.220 long, 0.148–0.180 wide; posterolateral body setae average ca. 0.070 long; hypostome 0.140–0.160
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argas miniatus

Antricola (after Jones & Clifford 1972)

1 Claws present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Antricola marginatus


- Claws absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Antricola coprophilus/Antricola mexicanus

THE SOFT TICKS OF MEXICO Zootaxa 4623 (3) © 2019 Magnolia Press · 489
Otobius (after Jones & Clifford 1972)

1 Eyes present; tarsus I with 4 paracapsular setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Otobius megnini


- Eyes absent; tarsus I with 3 paracapsular setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Otobius lagophilus

Ornithodoros (original descriptions; Kohls et al. 1965; Jones and Clifford 1972; Labruna et al. 2011; Guzmán-
Cornejo et al. 2016; note: larva of O. dugesi is unknown)

1 Dorsum of body with 2 pairs of eyes; palpal article IV about twice as long as any other article . . . . . . Ornithodoros coriaceus
- Dorsum of body without eyes; palpal article IV about equal in length or shorter than any other article. . . . . . . . . . . . . . . . . . . 2
2 Basis capituli ventrally with a pair of cornua-like extensions posteriorly (Fig. 5B), with auriculae-like lateral projections; tarsi
without pulvilli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Basis capituli ventrally without cornua-like extensions posteriorly, without auriculae-like lateral projections; tarsi with pul-
villi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3 Dorsal plate narrow, elongate, 0.320–0.470 long by 0.092–0.133 wide . . . . . . . . . . . . . . . . . . . . . . . .Ornithodoros marinkellei
- Dorsal plate wide, triangular, 0.192–0.220 long by 0.152–0.160 wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Posthypostomal setae 1 long (0.080–0.100); file 1 of hypostome with 22–23 denticles, file 2 with 23–24 . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros mormoops
- Posthypostomal setae 1 short (0.018-0.020); file 1 of hypostome with 18–22 denticles, file 2 with 19–21 . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros viguerasi
5 Dorsum of body with 13 pairs of setae, comprising 11 dorsolateral pairs and 2 central pairs; dorsal plate present or absent, but
if present never pyriform, elongate or pointed; hypostome with denticles only along anterior portion, dentition 2/2 . . . . . . . . 6
- Dorsum of body with 14–30 pairs of setae, comprising 10–25 dorsolateral pairs and 3 or 4 central pairs; dorsal plate present,
elongate or pyriform; hypostome with denticles throughout most of length, dentition 3/3 to 5/5 anteriorly . . . . . . . . . . . . . . . 8
6 Dorsal plate present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros nicollei
- Dorsal plate absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Hypostome short (0.086–0.101); posthypostomal setae 1 moderately long (0.051–0.069); dorsal setae thin . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros parkeri
- Hypostome moderately long (0.130–0.150); posthypostomal setae 1 long (0.074–0.094); dorsal setae thick . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros turicata
8 Capsule of Haller’s organ with reticulations; parasites of bats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Capsule of Haller’s organ without reticulations; hosts various, including bats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
9 Body with 17–30 pairs of dorsal setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Body with 14 (rarely 15 or 16) pairs of dorsal setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10 Body with 17–21 pairs of dorsal setae; hypostome dentition 4/4 at tip, basal denticles crowded and deformed (Fig. 5C) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros azteci
- Body with 27–30 pairs of dorsal setae; hypostome dentition 3/3 at tip, basal denticles not crowded and deformed . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros setosus
11 Venter of body with 7 or 8 pairs of setae plus posteromedian seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- Venter of body with 9 pairs of setae plus posteromedian seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
12 Venter with 7 pairs of setae; hypostome constricted at mid-length (Fig. 5D) . . . . . . . . . . . . . . . . . . . . Ornithodoros knoxjonesi
- Venter with 8 pairs of setae (Fig. 5E); hypostome not constricted at mid-length . . . . . . . . . . . . . . . . . Ornithodoros yumatensis
13 Hypostome length more than 0.290; posterior margin of basis capituli rounded (Fig. 5F); file 1 of hypostome with 22–24 den-
ticles, file 2 with 21–22, file 3 with 12–14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Ornithodoros brodyi
- Hypostome length less than 0.200; posterior margin of basis capituli trapezoidal (Fig. 5G) . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Ornithodoros dyeri and Ornithodoros rossi (Note: comments in discussion section)
14 Hypostome dentition 4/4 or 5/5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- Hypostome dentition 3/3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
15 Dorsum of body with 22–25 pairs of setae; hosts are marine birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros capensis
- Dorsum of body with 19 pairs of setae; hosts are bats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros stageri
16 Posteromedian seta present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Posteromedian seta absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros denmarki
17 Hypostome very long and thin (0.244–0.257); outermost file with 25–27 denticles . . . . . . . . . . . . Ornithodoros puertoricensis
- Without this combination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 Dorsum with 19 pairs of setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros kohlsi
- Dorsum with 16–17 pairs of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 Dorsal plate length 0.222-0.265; file 3 of hypostome with 7–11 denticles; hosts are mammals other than bats. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithodoros talaje
- Without this combination of characters; hosts are bats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 Dorsal plate length 0.331–0.360; file 3 of hypostome with 5–8 denticles . . . . . . . . . . . . . . . . . . . . . . . . . . .Ornithodoros kelleyi
- Dorsal plate length less than 0.300 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 Dorsal plate wide (0.155–0.177); dorsoanterolateral setae ca. 0.125 (separation of these two species based on characters given
by Černý 1967) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Ornithodoros dusbabeki

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- Dorsal plate narrower (0.127–0.150); dorsoanterolateral setae ca. 0.100 (or less); file 3 of hypostome with 8–12 denticles . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Ornithodoros hasei

FIGURE 2. A) Argas sanchezi, male, showing irregular peripheral cells. B) Otobius lagophilus, male, dorsal idiosomal integu-
ment. C) Ornithodoros coriaceus, male, showing iridescent mammillae. D) Ornithodoros coriaceus, male, lateral view of eyes
(arrows). E) Ornithodoros nicollei, male, tarsus I proximally with a single prominent dorsal hump (arrow). F) Ornithodoros
turicata, female, mammillae large and not crowded. G) Ornithodoros parkeri, male, mammillae small and crowded. H) Orni-
thodoros dyeri, female, with dorsal contiguous ridges (arrow).

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FIGURE 3. A) Ornithodoros azteci, female. B) Ornithodoros brodyi, female (upper) compared with Ornithodoros puertoricen-
sis, female (lower). C) Ornithodoros kelleyi, female. D) Ornithodoros dusbabeki, female. E) Ornithodoros talaje, female, discs.
F) Ornithodoros kohlsi, female, discs. G) Ornithodoros talaje, female, ventral view. H) Ornithodoros talaje, female, genital
aperture. I) Ornithodoros talaje, male, ventral view.

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FIGURE 4. A) Antricola marginatus, male, hypostome scoop-like, and B) digitiform tubercles (arrow). C) Antricola mexica-
nus, female, tufts of setae (arrow). D) Ornithodoros yumatensis, female, cheeks large (arrow), and E) micromammillated surface
of legs (arrow). F) Ornithodoros brodyi, female, hood (arrow).

Parasite-Host List

Antricola coprophilus (McIntosh, 1935)


CHIAPAS: Cueva de la Chepa, Tuxtla Gutiérrez, bat guano (Mazzotti 1940; Hoffmann et al. 2004)
(CNAC001931). COLIMA: Cueva de la Fábrica (5 km W of Coquimatlán), bat guano (Mazzotti 1941); Cueva el
Salitre, Los Ortices, Macrotus waterhousii (Herrera et al. 2016). GUERRERO: ND, bat guano (Chavarría 1941).
JALISCO: Cueva “El Chico,” 5 km NW of Tuxcacuesco, bat guano (Hoffmann 1962; Hoffmann et al. 2004).
NUEVO LEÓN: Linares, bat guano (McIntosh 1935). SAN LUIS POTOSÍ: Sótano de la Tinaja, bat guano (Hoff-
mann et al. 2004). TAMAULIPAS: Matamoros, bat guano (Philip 1940); Cueva de los Cuarteles, Aldama, bat gua-
no (Hoffmann 1962; Carrillo et al. 1966; Hoffmann et al. 2004) (CNAC001930); Cueva de la Florida (7.5 km NE
of Antiguo Morelos), bat guano (Hoffmann et al. 2004). VERACRUZ: Cuevas de Laguna Encantada, bat guano
(Hoffmann 1962; Hoffmann et al. 2004) (CNAC001927). YUCATÁN: Cueva de Hoctún, bat guano (Hoffmann
1962; Hoffmann et al. 2004) (CNAC001928).
Note: In this work we have retained the original species determination of specimens deposited at CNAC as
A. coprophilus because their stigmal plates are suboval and do not protrude from the body. However, our female
specimens possess small tufts of setae along the posteroventral edge of the idiosoma, similar to those seen in A.
mexicanus. Further collections near the type localities of both species are needed in order to obtain additional mor-
phological and molecular data that would resolve our specimens’ taxonomic status.

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FIGURE 5. A) Antricola mexicanus, larva, pulvilli extended. B) Ornithodoros marinkellei, larva, cornua-like extensions. C)
Ornithodoros azteci, larva, denticles crowded and deformed. D) Ornithodoros knoxjonesi, larva, hypostome constricted at mid-
length. E) Ornithodoros yumatensis, larva, ventral idiosoma with 8 pairs of setae. F) Ornithodoros brodyi, larva, posterior mar-
gin of basis capituli rounded. G) Ornithodoros dyeri, larva, posterior margin of basis capituli trapezoidal.

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Antricola marginatus (Banks, 1910)
CAMPECHE: Grutas de Xtacumbilxunam (2 km SW of Bolanchenticul), Hopelchén, bat guano (Keirans
& Clifford 1975). VERACRUZ: Cueva cercana a Playa Vicente, Playa Vicente, bat guano (Hoffmann et al.
1972) (CNAC001926). YUCATÁN: Cueva de Calcehtok, bat guano (Labruna et al. 2012) (CNAC006806-6807;
LAFC000189).
New record: CHIAPAS: 3M, 96N, Cueva de la Chepa, Tuxtla Gutiérrez, 20-X-2016 (LAFC000269).

Antricola mexicanus Hoffmann, 1958


CAMPECHE: About 4.8 km S of Champotón, probably Pteronotus davyi fulvus (Kohls et al. 1965); Grutas de
Xtacumbilxunam (2 km SW of Bolanchenticul), Hopelchen, bat guano (Keirans & Clifford 1975). CHIHUAHUA:
Sótano del Sauz, bat guano (Hoffmann et al. 2004). GUERRERO: Grutas de Juxtlahuaca, bat guano (Hoffmann
1958; Hoffmann et al. 2004) (CNAC000257-58). OAXACA: San Pedro Mixtepec, Juquila, Balantiopteryx plicata,
Pteronotus parnelli (Colín-Martínez & García- Estrada 2018). SAN LUIS POTOSÍ: Cueva de Taninul No. 1, bat
guano (Hoffmann et al. 2004). TABASCO: Cueva del Azufre, 3.5 km S of Tapijulapa, bat guano (Keirans et al.
1977). TAMAULIPAS: Cueva de la Florida, bat guano (Hoffmann et al. 2004). YUCATAN: Actún Xpukil, 3 km
S of Calcehtok, bat guano (Keirans et al. 1977).
New records: COLIMA: 1L, Cueva de los Amiales, Jala, Pteronotus parnellii, 22-VI-2015 (LAFC000282).
CHIAPAS: 19M, 7F, 190N, Cueva de la Chepa, Tuxtla Gutiérrez 28-X-2016 (LAFC000270); 5M, 1F, 56N, Cueva
de la Chepa, Tuxtla Gutiérrez 2-IV-2014 (LAFC000271). NAYARIT: 12M, 6F, 10N, Cueva cerro de las Viejas,
Santiago Ixcuintla, 18-VI-2015 (LAFC000272).

Argas brevipes Banks, 1908


BAJA CALIFORNIA: 8.8 km S of San Felipe, woodpecker holes (Kohls et al. 1961); 45.1 km S of Punta
Prieta, nest of Tachycineta thalassina thalassina (Kohls et al. 1961).

Argas cooleyi Kohls and Hoogstraal, 1960


CHIHUAHUA: Cueva del salitre, 13 km W of Villa de Matamoros, ND (Hoffmann et al. 2004).

Argas giganteus Kohls and Clifford, 1968


SONORA: 64.4 km S of Navojoa, Corvus imparatus (Kohls & Clifford 1968); 22.5 km W of Santa Ana, Tox-
ostoma bendirei (Kohls & Clifford 1968); Rancho Carrizal N of Alter, Toxostoma curvirostre (Kohls & Clifford
1968); 30.6 km S of Sonoyta, Kieneria fusca (Kohls & Clifford 1968) (CNAC000256); ND, wild birds (Hoffmann
& López-Campos 2000).

Argas miniatus Koch, 1844


SINALOA: ND, hens (Hoffmann & López-Campos 2000).
Note: The presence of this species in Mexico remains doubtful and is based on a single literature record (Hoff-
mann & López-Campos 2000) with limited collection data. According to Muñoz-Leal et al. (2018), A. miniatus is
a Neotropical species, which suggests that it may be present in Mexico, because most of the state of Sinaloa lies
within the Neotropical Zoogeographic Region (Morrone 2014). Nonetheless, the issue of whether A. miniatus oc-
curs in Mexico will not be resolved without additional collections and the examination and deposition of specimens
in scientific collections.

Argas persicus (Oken, 1818)


CIUDAD DE MÉXICO: Tlalpan, chicken coops (Hoffmann 1926); ND, chicken coops, markets (Cha-
varría 1958). CHIAPAS: Tapachula, ND (Chavarría 1941). CHIHUAHUA: Chihuahua, hens (Hoffmann 1962).
COAHUILA: ND, chicken coops (Hoffmann 1926); ND, ND (Bishop 1941); Jalisco, hens (Ortiz Mariotte 1945);
ND, ND (Chavarría 1958). DURANGO: San José de Zaragoza, hens (Ortiz Mariotte 1945). GUANAJUATO: ND,
ND (Nuttall et al. 1908) (referred to as Ar. sanchezi). HIDALGO: ND, ND (Chavarría 1958). NUEVO LEÓN:
ND, ND (Chavarría 1958); near Monterrey, hens (Gibson & Carrillo 1959). SINALOA: Aguacaliente Grande, ND
(Bustamante & Varela 1943); Choix, ND (Bustamante & Varela 1943); El Disparate, ND (Bustamante & Varela
1943); Santa Ana, ND (Bustamante & Varela 1943); San Felipe, ND (Bustamante & Varela 1943); San Joaquín, ND

THE SOFT TICKS OF MEXICO Zootaxa 4623 (3) © 2019 Magnolia Press · 495
(Bustamante & Varela 1943); ND, ND (Ortiz Mariotte et al. 1944); ND, ND (Chavarría 1958); Cacalotán, Rosario,
hens (Hoffmann 1962); Soyita (probably Soyatita), Badiraguato, hens (Hoffmann 1962); Mazatlán, hens (Hoffmann
1962). SONORA: El Carrizal, ND (Bustamente & Varela 1943). TAMAULIPAS: ND, ND (Chavarría 1958); Tam-
pico, hens (Hoffmann 1962).
New record: SONORA: 8F, 5M, 2N, Agua Blanca, Etchojoa, hens, 30-XII-1975 (CNAC001936).
Notes: Chavarría (1941) recorded this species from northern and central Mexico, including the southern por-
tion of Baja California (Baja California Sur). Nuttall et al. (1908) cited Ar. persicus from southern California, but
Hoffmann (1962) recorded this species from Baja California (which can be mistranslated as southern California).
Bishop (1941) provided a distribution map for this species, showing two collection sites in the state of Coahuila,
Mexico. Silva-Goytia & Elizondo (1952) referred to the occurrence of this species in the “Región de la Laguna,”
which includes two states, Durango and Coahuila. Ortiz Mariotte et al. (1944) cite the collection of Ar. persicus
from two Mexican states, Sonora and Sinaloa, without specifying the collection locality. Chavarría (1958) stated
that this species is widely distributed in Mexico, in warm locales, whether wet or dry, and is generally associated
with hens. Hoffmann (1962) cited the occurrence of Ar. persicus in Puebla (Hoffmann 1926); however, this locality
record does not appear in the latter work. Hoffmann (1930) mentioned that Ar. persicus occurs in various parts of
Mexico, and Hoffmann (1961) records Ar. persicus from Baja California, Chiapas, Coahuila, Ciudad de México,
Hidalgo, Estado de México, Puebla, Sinaloa, Sonora, and Tamaulipas. This tick is typically associated with poultry
(specifically hens and turkeys); mammals and humans are exceptional hosts.

Argas radiatus Railliet, 1893


AGUASCALIENTES: ND, chicken coops (Hoffmann & López-Campos 2000). NUEVO LEÓN: ND, chick-
en coops (Hoffmann & López-Campos 2000). PUEBLA: ND, chicken coops (Hoffmann & López-Campos 2000).
TAMAULIPAS: 32.2 km S of Nuevo Laredo, from trees where chickens roost (Kohls et al. 1970; Hoffmann &
López-Campos 2000).

Argas sanchezi Dugès, 1887


AGUASCALIENTES: ND, chicken coops (Hoffmann & López-Campos 2000). BAJA CALIFORNIA:
Santa Agueda (probably Misión Santa Agueda), wild turtle dove (Hunter & Hooker 1907); ND, wild turtle dove
(Hoffmann & López-Campos 2000). BAJA CALIFORNIA SUR: ND, chicken coops (Hoffmann & López-Cam-
pos 2000). CIUDAD DE MÉXICO: ND, chicken coops (Hoffmann & López-Campos 2000). CHIHUAHUA:
Ciudad Juárez, hens (Hoffmann 1962) (CNAC001938); ND, chicken coops (Hoffmann & López-Campos 2000).
COAHUILA: ND, chicken coops (Hoffmann & López-Campos 2000). DURANGO: Lerdo, “from an old sack”
(Kohls et al. 1970); ND, chicken coops (Hoffmann & López-Campos 2000). GUANAJUATO: ND, Zenaida mac-
roura carolinensis (Dugès 1887). HIDALGO: ND, chicken coops (Hoffmann & López-Campos 2000). NUEVO
LEÓN: ND, chicken coops (Hoffmann & López-Campos 2000). PUEBLA: ND, chicken coops (Hoffmann &
López-Campos 2000). QUERÉTARO: ND, chicken coops (Hoffmann & López-Campos 2000). SAN LUIS PO-
TOSÍ: Tamazunchale, hens (Hoffmann, 1962) (CNAC001933); ND, chicken coops (Hoffmann & López-Campos
2000). SINALOA: ND, chicken coops (Hoffmann & López-Campos 2000). SONORA: La Dura, hens (Hoffmann
1962) (CNAC001939-40); Corral, hens (Hoffmann 1962; Carrillo et al. 1966) (CNAC001937, CNAC001941);
La Calera (referred to as Cumuripa), hens (Hoffmann 1962) (CNAC001932, CNAC001935); ND, chicken coops
(Hoffmann & López-Campos 2000). TAMAULIPAS: ND, chicken coops (Hoffmann & López-Campos 2000).
TLAXCALA: ND, chicken coops (Hoffmann & López-Campos 2000). ZACATECAS: ND, chicken coops (Hoff-
mann & López-Campos 2000).
New record: SINALOA: 6F, 4M, 1N, Los Mochis, chicken coops, II-1942 (CNAC001934).
Notes: Macias-Valadez (1923) referred to this species as Argas miniatus and believed that it probably oc-
curred throughout Mexico. Hoffmann (1962) stated that Ar. sanchezi was described by Dugès in 1891; however,
the original description was published in 1887. The records from CNAC—CNAC001932-33, CNAC001935, and
CNAC001937-41—were originally misidentified as Ar. persicus and later re-identified as Ar. sanchezi.

Nothoaspis reddelli Keirans and Clifford, 1975


CAMPECHE: Grutas de Xtacumbilxunam (2 km SW of Bolanchenticul), Holpelchén, bat guano (Keirans &
Clifford 1975; Keirans et al. 1977; Hoffmann et al. 2004; Guzmán-Cornejo et al. 2012). TABASCO: Cueva del

496 · Zootaxa 4623 (3) © 2019 Magnolia Press GUZMÁN-CORNEJO ET AL.


Azufre, 3.5 km S of Tapijulapa, bat guano (Keirans et al. 1977; Hoffmann et al. 2004; Guzmán-Cornejo et al. 2012
(CNAC006769, CNAC006770)). YUCATÁN: Actún Xpukil, 3 km S of Calcehtok, bat guano (Keirans et al. 1977;
Hoffmann et al. 2004); Cueva Toh, Oxcutzcab, 19.5 km SE of Oxcutzcab, ND (Hoffmann et al. 2004).
New record: YUCATÁN: 1M, Opichén, Cueva de Calcehtok, 14-VI-2010 (LAFC000191).

Ornithodoros azteci Matheson, 1935


COLIMA: Cueva el Salitre, Los Ortices, Macrotus waterhousii (Herrera et al. 2016). MORELOS: Cueva
Poza de Moctezuma, Oaxtepec, Desmodus rotundus (Hoffmann 1962); Atlihuayán, Yautepec, Desmodus rotundus
(Hoffmann 1962). QUINTANA ROO: Puerto Juárez, Artibeus jamaicensis yucatanicus (Kohls et al. 1965); ND,
Artibeus jamaicensis yucatanicus (Hoffmann & López-Campos 2000).YUCATÁN: Pisté, Artibeus jamaicensis
yucatanicus (Kohls et al. 1965); ND, Artibeus jamaicensis yucatanicus (Hoffmann & López-Campos 2000).

Ornithodoros brodyi Matheson, 1935


CHIAPAS: Cueva del Naranjo, Rancho el Arco, Cintalapa, Artibeus lituratus (Guzmán-Cornejo et al. 2017);
same data except Desmodus rotundus (Guzmán-Cornejo et al. 2017); Cueva del Guano, San Fernando, Desmodus
rotundus (Guzmán-Cornejo et al. 2017). GUERRERO: Grutas de Cacahuamilpa, on bat guano (Hoffmann 1962;
Hoffmann et al. 2004) (CNAC001947); JALISCO: 25.7 km NE of Tamazula, Pteronotus parnelli (Kohls et al.
1965; Hoffmann & López-Campos 2000). QUINTANA ROO: vicinity of Pueblo Xcan, Artibeus jamaicensis,
Chrotopterus auritus auritus, Myotis nigricans extremus (Kohls et al. 1965; Hoffmann & López-Campos 2000);
Cueva Zereque (actún Zuub), Lázaro Cárdenas, crevices in walls, Carollia sowelli (Guzmán-Cornejo et al. 2017).
TABASCO: Cueva Agua Blanca (Grutas Ixta-Ha), Macuspana, crevices in walls (Guzmán-Cornejo et al. 2017).
YUCATÁN: Cueva Sabacá, Tekax, on the floor (Hoffmann 1962; Hoffmann et al. 2004) (CNAC001965); ND, ND
(Hoffmann & López-Campos 2000); Cueva near Cenote Bal-Mil, Homún, crevices in walls (Guzmán-Cornejo et al.
2017); Cueva Kahua, Kahua, crevices in walls (Guzmán-Cornejo et al. 2017); Cueva Tzabnah, Tecoh, crevices in
walls (Guzmán-Cornejo et al. 2017).
Notes: The specimens labeled as CNAC001947 and CNAC001965 were originally determined as O. talaje,
and relabeled subsequently as O. yumatensis; however, during the research for this review, these specimens were
determined to actually represent O. brodyi.

Ornithodoros capensis Neumann, 1901


BAJA CALIFORNIA: Isla Rasa, ND, under rocks (Hoffmann 1962) (CNAC001949-50). COLIMA: Isla
San Benedicto (Islas Revillagigedo), under stone near nest of Sula dactylatra californica (Kohls 1957; Hoffmann
and López-Campos 2000). QUINTANA ROO: Isla Contoy, Pelecanus occidentalis (Hoffmann & López-Campos
2000). SONORA: Isla del Medio, Guaymas, under rocks (Hoffmann 1962) (CNAC001948). YUCATÁN: Arrecife
Alacranes, on the floor (Hoffmann 1962) (CNAC001964).

Ornithodoros coriaceus Koch, 1844


ND: ND, ND (Koch 1844). COAHUILA: ND, ND (Silva-Goytia & Elizondo 1952). DURANGO: ND, ND
(Silva-Goytia & Elizondo 1952). OAXACA: near San Gerónimo, Istmo de Tehuantepec, ND (Nuttall et al. 1908);
Tehuantepec, ND (Brumpt et al. 1939); Juchitán, houses (Chavarría 1941); Salina Cruz, houses (Chavarría 1941);
Tehuantepec, houses (Chavarría 1941); Tehuantepec, ND (Davis 1942); Juchitán, ND (Mazzotti 1942a); Salina
Cruz, ND (Mazzotti 1942a); Tlacolula, ND (Mazzotti 1947); Espinal, houses (Hoffmann 1962); Huilotepec, houses
(Hoffmann 1962); Tololapa, ND (Hoffmann 1962); Ixtepec, ND (Carrillo et al. 1966); SINALOA: ND, ND (Ortiz
Mariotte et al. 1944). SONORA: ND, ND (Ortiz Mariotte et al. 1944).
New records: OAXACA: 5F, 7N, Ixtepec, ND, XII-1941 (CNAC001953); 2F, 18M, 60N, Ixtepec, ND, 10-IV-
1940 (CNAC001954).
Notes: Hoffmann (1930) stated that this species occurs from California to Chiapas, where it is a parasite of
small rodents and large wild and domestic mammals, such as deer and cattle; it occasionally attacks humans. Later,
Hoffmann (1961) also referred to this species as occurring from California to Oaxaca and Chiapas, again as a pest of
cattle and occasionally humans. Hoffmann and López-Campos (2000) cited this species from Baja California, Baja
California Sur, Chiapas, Coahuila, Durango, Guerrero, Jalisco, Michoacán, Oaxaca, Sinaloa, and Sonora, listing its
hosts as rodents and their burrows, deer, and cattle, while mentioning that it may be found in nests of swallows, and
that it is occasionally a human parasite.

THE SOFT TICKS OF MEXICO Zootaxa 4623 (3) © 2019 Magnolia Press · 497
Ornithodoros denmarki Kohls, Sonenshine and Clifford, 1965
BAJA CALIFORNIA: Isla Raza, under rocks near nesting gulls and terns (Kohls et al. 1965). Isla Calavera,
under rocks (Kohls et al. 1965).
Notes: The records of Kohls et al. (1965) are probably a recapitulation of those cited by Hoffmann and López-
Campos (2000), who used the generic names Larus and Sterna to refer to gulls and terns, respectively.

Ornithodoros dugesi Mazzotti, 1943a


COAHUILA: Near Sabinas, nests of wild rats (Mazzotti 1943a). NUEVO LEÓN: China, nests of wild rats
(Mazzotti 1949); Los Aldamas, ND (Hoffmann 1962); Los Herreras, ND (Hoffmann 1962); General Bravo, ND
(Hoffmann 1962); Villaldama, ND (Hoffmann 1962). SAN LUIS POTOSÍ: Bocas, nests of Neotoma albigula
(Mazzotti 1949).

Ornithodoros dusbabeki Černý, 1967


VERACRUZ: ND, Molossus rufus (Dusbábek 1970).

Ornithodoros dyeri Cooley and Kohls, 1940


BAJA CALIFORNIA: Isla Pescadero, Myotis vivesi (Kohls et al. 1965); Isla Partida (Cardonosa), Myotis
vivesi (Kohls et al. 1965); Isla Pond, Myotis vivesi (Kohls et al. 1965). CHIAPAS: Cueva La Chepa, Tuxtla Gutiér-
rez, bat guano (Mazzotti 1941). COAHUILA: Cueva Detector perdido, Monclova, cave walls (Guzmán-Cornejo
et al. 2016) (CNAC009228). COLIMA: Cueva La Fábrica, 5km W of Coquimatlán, bat guano (Mazzotti 1941;
Hoffmann et al. 2004). NUEVO LEÓN: Cueva del Diablo, 7 km W of Sabinas Hidalgo, bat guano (Kohls et al.
1965; Hoffmann et al. 2004). OAXACA: Tequistlán, Balantiopteryx plicata (Kohls et al. 1965); Puerto Ángel,
Balantiopteryx plicata (Kohls et al. 1965); San Pedro Mixtepec, Juquila, Balantiopteryx plicata, Pteronotus parnelli
(Colín-Martínez & García- Estrada 2018). QUINTANA ROO: 57.7 km S, 54.5 km W of Chetumal, Trachops cir-
rhosus coffini (Wolfgang & Polaco 1985); 45.9 km S, 43.1 km W of Chetumal, Trachops cirrhosus coffini (Wolf-
gang & Polaco 1985); 39.4 km S, 38.8 km W of Chetumal, Trachops cirrhosus coffini (Wolfgang & Polaco 1985).
SAN LUIS POTOSÍ: Cueva de Taninul, ND (Hoffmann 1962; Hoffmann et al. 2004). VERACRUZ: Cueva Rey
del Oro, Emiliano Zapata, cave walls (Guzmán-Cornejo et al. 2016) (LAFC000127-131, 133-134). YUCATÁN:
Cueva Chocantes, Tekax, cave walls (Guzmán-Cornejo et al. 2016) (LAFC00126); Cueva El Naranjal, Peropteryx
macrotis (Guzmán-Cornejo et al. 2016) (LAFC000209-210); Km 56, Santa Elena-Loltún road (cavity I), Mimon co-
zumelae (Guzmán-Cornejo et al. 2016) (LAFC000205, 206); Santa Elena-Loltún road (cavity II), Peropteryx mac-
rotis (Guzmán-Cornejo et al. 2016) (LAFC000207, 208); Santa Elena-Loltún road (cavity II), cave walls (Guzmán-
Cornejo et al. 2016) (LAFC000211).
New records: COLIMA: 4F, 1M, 2N, Cueva la Finca, Coquimatlán, ND, 20-I-1943 (CNAC001958); 3L, Cue-
va de los Amiales, Jala, Macrotus waterhousii, 22-VI-2015 (LAFC000279); same data except 5L (LAFC000280),
2L (LAFC000281).
Notes: Guzmán-Cornejo et al. (2016) used the acronym LAF instead of LAFC. Collection LAFC000211 was
made from cave walls, not from Peropteryx macrotis. Hoffmann & López-Campos (2000) cited this species from
Chiapas, Colima, Nuevo León and San Luis Potosí, on bat guano and cave walls inhabited by bats.

Ornithodoros hasei (Schulze, 1935)


YUCATÁN: Pisté, Molossus rufus nigricans (Kohls et al. 1965; Hoffmann & López-Campos 2000). SINALOA:
Santa Lucía, Myotis velifer velifer (Kohls et al. 1965; Hoffmann & López-Campos 2000).

Ornithodoros kelleyi Cooley and Kohls, 1941a


DURANGO: Nombre de Dios, Tadarida brasiliensis mexicanus (Guzmán-Cornejo et al. 2003).

Ornithodoros knoxjonesi Jones and Clifford, 1972


MORELOS: ND, Balantiopteryx plicata (Hoffmann & López-Campos 2000).

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Ornithodoros kohlsi Guglielmone and Keirans, 2002
JALISCO: 9.6 km E of Limón, Molossus sinaloae (Jones et al. 1972).

Ornithodoros marinkellei Kohls, Clifford and Jones, 1969


New records: NAYARIT: 1L, Cueva Cerro de las Viejas, Santiago Ixcuintla, Pteronotus personatus, 18-VI-
2015 (LAFC000284); same data except 3L (LAFC000283).

Ornithodoros mormoops Kohls, Clifford and Jones, 1969


COLIMA: Cueva el Salitre, Los Ortices, Macrotus waterhousii (Herrera et al. 2016).

Ornithodoros nicollei Mooser, 1932


ND: ND, rustic houses (Chavarría 1941). COAHUILA: ND, coyote, houses, dogs (Silva-Goytia & Eli-
zondo 1952). COLIMA: Colima, Hodomys alleni (Brumpt et al. 1939); ND, ND (Davis 1942); Coquimatlán,
ND (Mazzotti 1942a); ND, dogs, man (Hoffmann 1961). GUERRERO: Xalitla, huts of natives feeding of man
(Mooser 1932); Iguala, huts of natives feeding of man (Mooser 1932); Mezcala, huts of natives feeding of man
(Mooser 1932); Río Balsas region, huts of natives (Pilz & Mooser 1936), Iguala, ND (Chavarría 1941; Mazzotti
1942); Tuxpan, ND (Chavarría 1941); ND, ND (Davis 1942); ND, ND (Mazzotti 1947); ND, dogs, man (Hoff-
mann 1961); “El Potrero,” Tlalchapa, houses (Hoffmann,1962) (CNAC001962); Tlalchichilpa, houses (Hoffmann
1962) (CNAC001961); Olinalá, houses (Hoffmann 1962); Ometepec, houses (Hoffmann 1962); Río Balsas, houses
(Hoffmann 1962). JALISCO: Autlán, ND (Mazzotti 1942a); ND, ND (Mazzotti 1947); ND, dogs, man (Hoffmann
1961); MICHOACÁN: El Jorullo, on soil (Hoffmann 1962) (CNAC005011). MORELOS: ND, ND (Mazzotti
1947); ND, dogs, man (Hoffmann 1961); Amacuzac, on soil (Hoffmann 1962); Cuevas de Huajintlán, 0.5 km NW
of Huajintlán, walls and on the floor (Hoffmann 1962; Hoffmann et al. 2004) (CNAC001963). OAXACA: Putla,
houses (Hoffmann 1962); Oaxaca, Jamiltepec, houses (Hoffmann 1962); Cacahuatepec, houses (Hoffmann 1962);
Calihuala, Silac, houses (Hoffmann 1962) (CNAC001959); Silac, ND (Carrillo et al. 1966). PUEBLA: Atencingo,
ND (Pilz & Mooser 1936); ND, ND (Davis 1942); ND, Neotoma sp., dogs, man (Davis 1943); ND, dogs, man
(Hoffmann 1961); Tecomatlán, houses (Hoffmann 1962) (CNAC005006); Matamoros, squirrel (Hoffmann 1962)
(CNAC005004). SONORA: Isla Tiburon, lodge of Neotoma sp. (Kohls et al. 1965; Hoffmann & López-Campos
2000).
Notes: Chavarría (1941) stated that this species is common from southern Morelos to southern Mexico. Or-
nithodoros nicollei has been collected from peasant houses, caves, humans, dogs, squirrels, Hodomys alleni, and
Neotoma sp. in the states of Morelos, Coahuila, Colima, Durango, Guerrero, Jalisco, Michoacán, Morelos, Oaxaca,
and Puebla (Hoffmann & López-Campos 2000).
New records: ND: 2M, 5N, ND, ND, ND (CNAC001960). PUEBLA: 21F, 49M, 79N, Tecomatlán, houses,
XII-42 (CNAC005005); same data except 1F, 4M, 1N, II-43, (CNAC005007).

Ornithodoros parkeri Cooley, 1936


BAJA CALIFORNIA SUR: near Santiago, nest of Neotoma sp. (Kohls et al. 1965; Hoffmann & López-Cam-
pos 2000).

Ornithodoros puertoricensis Fox, 1947


COLIMA: Isla Clarion, Islas de Revillagigedo, Athene cunicularia (Endris et al. 1989).

Ornithodoros rossi Kohls, Sonenshine and Clifford, 1965


BAJA CALIFORNIA SUR: San Antonio, Macrotus californicus (Kohls et al. 1965; Hoffmann & López-Cam-
pos 2000); Las Cuevas, Macrotus californicus (Kohls et al. 1965; Hoffmann & López-Campos 2000); Miraflores,
Macrotus californicus (Kohls et al. 1965; Hoffmann & López-Campos 2000). COLIMA: Cueva el Salitre, Los Or-
tices, Macrotus waterhousii (Herrera et al. 2016). MORELOS: ND, Sturnira sp., Leptonycteris sp., Glossophaga
sp. (Hoffmann & López-Campos 2000). PUEBLA: Cueva el Santuario, Tzinacatepec, Macrotus waterhousii (Her-
rera et al. 2016). SONORA: Kino, Macrotus californicus (Kohls et al. 1965; Hoffmann & López-Campos 2000);
ND, Parastrellus hesperus (= Pipistrellus hesperus) (Poché & Keirans 1975; Hoffmann & López-Campos 2000);
Cueva del Tigre, Carbó, Macrotus californicus (Herrera et al. 2016).

THE SOFT TICKS OF MEXICO Zootaxa 4623 (3) © 2019 Magnolia Press · 499
Ornithodoros setosus Kohls, Clifford and Jones, 1969
CHIAPAS: 2.1 km SSE of Zapaluta, Pteronotus parnellii (Kohls et al. 1969; Hoffmann & López-Campos
2000). SINALOA: 2.4 km N of Badiraguato, Pteronotus personatus psilotis (Kohls et al. 1969; Hoffmann &
López-Campos 2000).

Ornithodoros stageri Cooley and Kohls, 1941b


COAHUILA: Mina de San Felipe, Sabinas, walls of tunnels inhabited by bats (Mazzotti 1950b; Hoffmann &
López-Campos 2000).

Ornithodoros talaje (Guérin-Méneville, 1849)


ND: ND, ND (Dugès 1884; Mégnin 1885); ND, man and domestic animals (Chavarría 1941); ND, ND (Davis
1942). BAJA CALIFORNIA: ND, rodents (Hoffmann & López-Campos 2000). BAJA CALIFORNIA SUR: ND,
rodents (Hoffmann & López-Campos 2000). CAMPECHE: ND, rodents (Hoffmann 1930); ND, man and domes-
tic animals (Hoffmann 1961). CIUDAD DE MÉXICO: San Juan de Aragón, horses, pigs, cattle (Macías-Valadez
1923). CHIAPAS: ND, ND, rodents (Hoffmann 1930); Huixtla, ND (Brumpt et al. 1939); Ixtapa, ND (Mazzotti
1942a); Simojovel, ND (Mazzotti 1942a); Tuxtla Gutiérrez, ND (Mazzotti 1942a); Acala, ND (Mazzotti 1942a);
Arriaga, houses (Mazzotti 1942b); Angel Albino Corzo (Xaltenango), ND (Mazzotti 1953); ND, man and domes-
tic animals (Hoffmann 1961); Cintalapa, in houses (Hoffmann 1962); Chicomuselo, in houses (Hoffmann 1962);
Acacoyagua, in houses (Hoffmann 1962); Simojovel, in houses (Hoffmann 1962); Yajalón, in houses (Hoffmann
1962); Tecpatán, in houses (Hoffmann 1962); Huixtla, on soil (Hoffmann 1962) (CNAC001971); Planta de Luz,
Huixtla, in houses (Hoffmann 1962); El Vergel, in houses (Hoffmann 1962); Mapastepec, on soil (Hoffmann 1962);
(CNAC001970); Tonalá, in houses (Hoffmann 1962); Mapastepec (Carrillo et al. 1966). GUERRERO: Mezcala, in
huts of natives (Mooser 1932); Xalitla, in huts of natives (Mooser 1932); Iguala, in huts of natives (Mooser 1932);
Iguala, ND (Brumpt et al. 1939); ND, man and domestic animals (Hoffmann 1961); Olinalá, in houses (Hoffmann
1962); Cuadrilla Cuaulotitlán, on soil (Hoffmann 1962) (CNAC001969); Arcelia, in houses (Hoffmann 1962); ND,
rodents (Hoffmann & López-Campos 2000). JALISCO: Zapotitlán, ND (Macías-Valadez 1923); Río Balsas region,
in huts of natives (Pilz & Mooser 1936). MICHOACÁN: Pueblo de Turicato, horses, pigs, cattle (Macías-Valadez
1923); Morelia, ND (Bustamante et al. 1947); Ario de Rosales, ND (Bustamante et al. 1947); Pátzcuaro, ND (Busta-
mante et al. 1947); Zamora, ND (Bustamante et al. 1947); Coalcomán, ND (Bustamante et al. 1947); Uruapan, ND
(Bustamante et al. 1947); ND, rodents (Hoffmann & López-Campos 2000). MORELOS: ND, man and domestic
animals (Hoffmann 1961); Oaxtepec, on soil (Hoffmann 1962); ND, rodents (Hoffmann & López-Campos 2000).
OAXACA: El Istmo (the author is probably referring to Istmo de Tehuantepec), rodents (Hoffmann 1930); Istmo de
Tehuantepec, in huts of natives (Pilz & Mooser 1936); Matías Romero, ND (Mazzotti 1942a); Pijijiapan, in houses
(Hoffmann 1962); Mapastepec, in houses (Hoffmann 1962); Escuintla, in houses (Hoffmann 1962); Villa Flores, in
houses (Hoffmann 1962); Putla, in houses (Hoffmann 1962). PUEBLA: Tecomatlán, in houses (Hoffmann 1962);
ND, rodents (Hoffmann & López-Campos 2000). QUINTANA ROO: ND, ND (Hoffmann 1962); ND, rodents
(Hoffmann & López-Campos 2000). SINALOA: Mazatlán, ND, horses, pigs, cattle (Macías-Valadez 1923); ND,
man and domestic animals (Hoffmann 1961); Chinobampo, on soil (Hoffmann 1962); ND, rodents (Hoffmann
& López-Campos 2000). SONORA: ND, rodents (Hoffmann & López-Campos 2000); ND, Spermophilus tereti-
caudus (Venzal et al. 2008); ND, Dipodomys deserti (Venzal et al. 2008). TABASCO: ND, rodents (Hoffmann
1930); ND, man and domestic animals (Hoffmann 1961); ND, man and domestic animals (Hoffmann 1961). TAM-
AULIPAS: ND, rodents (Hoffmann 1930; Hoffmann & López-Campos 2000); ND, man and domestic animals
(Hoffmann 1961). VERACRUZ: ND, rodents (Hoffmann 1930; Hoffmann & López-Campos 2000); ND, man and
domestic animals (Hoffmann 1961); Catemaco, under rocks (Hoffmann 1962) (CNAC001968); Los Tuxtlas, on
the soil (Hoffmann 1962). YUCATÁN: ND, rodents (Hoffmann 1930); Xtoloc cenote cave, on the floor (Wharton
1938; Hoffmann et al. 2004); ND, man and domestic animals (Hoffmann 1961); Cueva de Balaam Canché, Chichén
Itzá, on the floor (Hoffmann 1962; Hoffmann et al. 2004); Isla Pérez, under wood (Hoffmann 1962); Yuncu cenote,
Yuncu, Abala, ND (Hoffmann et al. 2004).
Notes: Hoffmann (1930) and Hoffmann & López-Campos (2000) state that this species is associated with ro-
dents and other mammals that live in caves and underground nests. Brumpt et al. (1939) claimed to have collected
this species from an armadillo’s nest in “San Gerónimo” but did not provide the Mexican state. Mazzotti (1947)

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described the geographical distribution of O. talaje in the tropical zones of southern Mexico and the Yucatan Penin-
sula. Hoffmann (1962) cited a record of this species from San Juan Zaichicoban, Oaxaca, based on Mooser (1932);
however, we were unable to find this record in the latter work. Chavarría (1958) mentioned that this species is pres-
ent in Mexico and is associated with mammals and birds, especially pigs and hens.

Ornithodoros turicata (Dugès, 1876)


ND: ND, man, pigs, sheep and cattle (Chavarría 1941). AGUASCALIENTES: ND, ND (Hoffmann 1930);
Aguascalientes, on adobe walls of a house (Pilz & Mooser 1936); Aguascalientes, ND (Brumpt et al. 1939; Varela &
Aparicio 1951); ND, ND (Davis 1942); Aguascalientes, ND (Mazzotti 1943b); Calvillo, ND (Mazzotti 1947); ND,
mammals, birds and reptiles (Chavarría 1958); ND, pigs, small mammals, sheep, man, cattle (Hoffmann 1961); Jesús
María, soil of a room infested with moles and rats (Hoffmann 1962); ND, pigs, rats, tuzas (Geomyidae), rabbits,
man (Hoffmann & López-Campos 2000). CIUDAD DE MÉXICO: San Juan de Aragón (near Guadalupe Hidalgo
Municipality = Gustavo A. Madero), ND (Altamirano 1906). COAHUILA: Saltillo, ND (Mazzotti 1942a); Saltillo,
ND (Varela & Aparicio 1951); Región Lagunera, ND (Silva-Goytia & Elizondo 1952); ND, pigs, small mammals,
sheep, man, cattle (Hoffmann 1961); Ramos Arizpe, in houses (Hoffmann 1962); Carneros, in houses (Hoffmann
1962); San Pedro de las Colonias, in houses (Hoffmann 1962); Agua Nueva, in houses (Hoffmann 1962); pigs,
rats, tuzas (Geomyidae), rabbits, man (Hoffmann & López-Campos 2000). DURANGO: Región Lagunera, ND
(Silva-Goytia & Elizondo 1952). ESTADO DE MÉXICO: ND, pigs, small mammals, sheep, man, cattle (Hoff-
mann 1961); pigs, rats, tuzas (Geomyidae), rabbits, man (Hoffmann & López-Campos 2000). GUANAJUATO:
ND, pigs (Dugès 1876) referred to as Argas turicata; ND, ND (Hoffmann 1930); Irapuato, ND (Brumpt et al.
1939); ND, pig sties (Cooley & Kohls 1944); ND, ND (Davis 1942); San Nicolás de los Agustinos, ND (Mazzotti
1942a); pigs, small mammals, sheep, man, cattle (Hoffmann 1961); Valle de Santiago, in houses (Hoffmann 1962);
San Francisco del Rincón, in houses (Hoffmann 1962); pigs, rats, tuzas (Geomyidae), rabbits, man (Hoffmann &
López-Campos 2000). GUERRERO: Arcelia, ND (Varela & Aparicio 1951); ND, pigs, small mammals, sheep,
man, cattle (Hoffmann 1961); pigs, rats, tuzas (Geomyidae), rabbits, man (Hoffmann & López-Campos 2000). HI-
DALGO: ND, mammals, birds and reptiles (Chavarría 1958). JALISCO: Zapotitlán, ND (Macías-Valadez 1923);
Guadalajara, ND (Mazzotti 1942a); Los Altos, pigs (Cooley & Kohls 1944); Encarnación de Díaz, ND (Mazzotti
1947); Jalostotitlán, ND (Mazzotti 1947); ND, mammals, birds and reptiles (Chavarría 1958); pigs, small mammals,
sheep, man, cattle (Hoffmann 1961); ND, pigs, rats, tuzas (Geomyidae), rabbits, man (Hoffmann & López-Campos
2000). MICHOACÁN: Turicato, ND (Macías-Valadez 1923); ND, ND (Hoffmann 1930); Carácuaro, ND (Maz-
zotti 1947); Chucándiro, ND (Varela & Aparicio 1951); ND, pigs, small mammals, sheep, man, cattle (Hoffmann
1961); ND, pigs, rats, tuzas (Geomyidae), rabbits, man (Hoffmann & López-Campos 2000). MORELOS: ND,
pigs, small mammals, sheep, man, cattle (Hoffmann 1961); Yautepec, in houses (Hoffmann 1962); ND, pigs, rats,
tuzas (Geomyidae), rabbits, man (Hoffmann & López-Campos 2000). NUEVO LEÓN: ND, mammals, birds and
reptiles (Chavarría 1958); pigs, small mammals, sheep, man, cattle (Hoffmann 1961). PUEBLA: Tecamachalco,
ND (Hoffmann 1930 (CNAC001975); Brumpt et al. 1939; Carrillo et al. 1966); Huaquechula, pigs (Cooley &
Kohls 1944); Atexcal, pigs (Cooley & Kohls 1944); ND, mammals, birds and reptiles (Chavarría 1958); ND, pigs,
small mammals, sheep, man, cattle (Hoffmann 1961); ND, pigs, rats, tuzas (Geomyidae), rabbits, man (Hoffmann &
López-Campos 2000). QUERETARO: San Clemente, ND (Brumpt et al. 1939); Querétaro, ND (Mazzotti 1942a);
ND, pigs, small mammals, sheep, man, cattle (Hoffmann 1961); San Juan del Río, in houses (Hoffmann 1962); ND,
pigs, rats, tuzas (Geomyidae), rabbits, man (Hoffmann & López-Campos 2000). SAN LUIS POTOSÍ: ND, ND
(Hoffmann 1930); San Luis Potosí, ND (Brumpt et al. 1939); ND, ND (Davis 1942); ND, pigs, small mammals,
sheep, man, cattle (Hoffmann 1961); Santa María del Río, in houses (Hoffmann 1962); ND, pigs, rats, tuzas (Geo-
myidae), rabbits, man (Hoffmann & López-Campos 2000). SINALOA: Mazatlán, ND (Macías-Valadez 1923); El
Parnaso, ND (Mazzotti 1947); La Palma, ND (Mazzotti 1947); Higuera de los Natoches, ND (Mazzotti 1947); ND,
mammals, birds and reptiles (Chavarría 1958); ND, pigs, small mammals, sheep, man, cattle (Hoffmann 1961);
Ocolomé, in houses (Hoffmann 1962); Ampelio, Las Cañas, in houses (Hoffmann 1962); ND, pigs, rats, tuzas (Geo-
myidae), rabbits, man (Hoffmann & López-Campos 2000). ZACATECAS: Espíritu Santo, ND (Mazzotti 1950a);
ND, pigs, rats, tuzas (Geomyidae), rabbits, man (Hoffmann & López-Campos 2000).
Notes: Macías-Valadez (1923) stated that this species is commonly associated with pigs. Hoffmann (1930) later
noted that O. turicata can be associated with pigs, rodents, rats, tuzas (Geomyidae) and man. Hoffmann (1930) also
stated that this species occurs in dry regions of central Mexico and along the west coast of the country. Chavarría

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(1941) confirmed the abundance of this species in central Mexico, but Mazzotti (1947) described a much broader
geographical distribution, from Puebla to Mexico’s northern border. Hoffmann (1962) cited Chavarría (1935, 1936)
and Mooser (1930) as stating that O. turicata occurs in Guanajuato, Puebla, and Jalisco; however, the three latter
papers do not appear among Hoffmann’s references, but instead are in Cooley & Kohls (1944), whose names, locali-
ties and collection years are identical to those listed in Hoffmann (1962).

Ornithodoros viguerasi Cooley and Kohls, 1941c


GUERRERO: Cueva de Juxtlahuaca, Colotlipa, Quelchultenango, Mormoops megalophylla (Nava et al. 2012).
YUCATÁN: Opichén, Cueva de Calcehtok, ND (Nava et al. 2012).

Ornithodoros yumatensis Cooley and Kohls, 1941b


CHIAPAS: Cueva de Zapaluta, 7 km SE of La Tinaja, bat guano, Desmodus rotundus murinus (Kohls et al.
1965; Hoffmann & López-Campos 2000; Hoffmann et al. 2004); Cueva del naranjo, Rancho el Arco, Cintalapa,
crevices in walls (Guzmán-Cornejo et al. 2017); Cueva del Guano, San Fernando, Desmodus rotundus (Guzmán-
Cornejo et al. 2017). COLIMA: Cueva el Salitre, Los Ortices, Macrotus waterhousii (Herrera et al. 2016). DU-
RANGO: Guanaceví, Corynorhinus mexicanus (Villegas-Guzman et al. 2005). SINALOA: Copala, bat guano,
Desmodus rotundus murinus (Kohls et al. 1965; Hoffmann & López-Campos 2000); Panuco, Artibeus lituratus
palmarum (Kohls et al. 1965; Hoffmann & López-Campos 2000); Santa Lucía, A. lituratus palmarum, Dermanura
azteca (Kohls et al. 1965; Hoffmann & López-Campos 2000). TAMAULIPAS: Cueva de los Vampiros, 10 km NE
of Adolfo López Mateos, ND (Hoffmann et al. 2004). YUCATÁN: Chichén Itzá, bat guano, Desmodus rotundus
murinus (Kohls et al. 1965; Hoffmann & López-Campos 2000); Cueva Chocantes, crevices in walls (Sánchez-
Montes et al. 2016); Cueva near Cenote Bal-Mil, crevices in walls (Sánchez-Montes et al. 2016); Cueva de la Santa
María, Homún, crevices in walls (Guzmán-Cornejo et al. 2017); Cueva Kaúa, Kaúa, crevices in walls (Guzmán-
Cornejo et al. 2017); Cueva Tzabnah, Tecoh, crevices in walls (Guzmán-Cornejo et al. 2017); Cueva Chocantes,
Tekax, crevices in walls (Guzmán-Cornejo et al 2017); Cueva el Naranjal, Tekax, Artibeus jamaicensis (Guzmán-
Cornejo et al. 2017).
New records: SAN LUIS POTOSÍ: 4F, 3N, Cueva los Sabinos, Ciudad Valles, San Luis Potosí, ND 3-IX-2015
(LAFC000273); 1L, Cueva del Tigre, Ciudad Valles, San Luis Potosí, Diphylla ecaudata, 2-IX-2015 (LAFC000274);
same data except 21L, Desmodus rotundus (LAFC000275); 2L, Desmodus rotundus (LAFC000276); 1L, Desmodus
rotundus (LAFC000277); 15L, Desmodus rotundus (LAFC000278).

Otobius lagophilus Cooley and Kohls, 1940


COAHUILA: Matamoros, donkeys (Silva-Goytia & Elizondo 1952); ND, rabbits (Hoffmann & López-Cam-
pos 2000). CHIHUAHUA: Delicias, hares (Kohls et al. 1965; Hoffmann & López-Campos 2000).

Otobius megnini (Dugès, 1884)


ND: ND, ND (Chavarría 1941); ND, ears of animals (Bishopp & Trembley 1945). ND, ears of domestic
animals and man (Hoffmann 1961). AGUASCALIENTES: Jesús María, ears of mules (Hoffmann 1962). BAJA
CALIFORNIA: La Acequia, ears of cattle (Hoffmann 1962) (CNAC001982); Los Pozos, ears of cattle (Hoff-
mann 1962) (CNAC001981). CIUDAD DE MÉXICO: ND, cows, horses (Macías-Valadez 1923); Colonia Santo
Tomás, ears of cattle (Hoffmann 1962); Iztacalco, ears of cattle (Hoffmann 1962); Azcapotzalco, ears of a cat
(Hoffmann 1962); Xochimilco, ears of cattle (Hoffmann 1962); Los Morales, ears of dogs (Hoffmann 1962); Cas-
tillo de Chapultepec, ears of dogs (Hoffmann 1962); Tacuba, ears of dogs (Hoffmann 1962); ND, on stables (Car-
rillo et al. 1966). CHIHUAHUA: ND, cows, horses (Macías-Valadez 1923); Chihuahua, ears of cattle (Hoffmann
1962); Rosales, Camargo, ears of cattle (Hoffmann 1962); Bocoyna, ears of cattle (Hoffmann 1962). COAHUILA:
cows, horses (Macías-Valadez 1923); La Flor de Jimulco (Región Lagunera), sheep (Ortiz Mariotte, 1945); Torreón
(Región Lagunera), cows (Ortiz Mariotte 1945); Jalisco (Región Lagunera), dog (Ortiz Mariotte 1945); Región
Lagunera, ND (Silva-Goytia & Elizondo 1952); Mariano Escobedo, ears of cattle (Hoffmann 1962). COLIMA:
Manzanillo, ears of cattle (Hoffmann 1962). DURANGO: Región Lagunera, ND, ND (Silva-Goytia & Elizondo
1952); San Pedro del Gallo, ears of cattle (Hoffmann 1962); Gómez Palacio, ears of a dog (González-Alvarez et al.
2018). ESTADO DE MÉXICO: Texcoco, walls of a cowshed (Gibson & Carrillo 1959). Valle de México, ears of
cattle (Hoffmann 1962) (CNAC001984); Santa Clara, ears of cattle (Hoffmann 1962); Pueblo Nuevo, ears of cattle

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(Hoffmann 1962); Tulpetlac, ears of cattle (Hoffmann 1962); Santa María, ears of cattle (Hoffmann 1962); Temas-
calapa, ears of cattle (Hoffmann 1962); Acambay, ears of cattle (Hoffmann 1962); Texcoco, ears of cattle (Hoff-
mann 1962); San Mateo Atenco, ears of cattle (Hoffmann 1962); Chimalhuacán, ears of cattle (Hoffmann 1962)
(CNAC001978); Carrillo et al. 1966). GUANAJUATO: ND, horses, donkeys, oxen, man (Dugès 1884; Nuttall et
al. 1908) (described in both references as Argas megnini); Apaseo, ears of cattle (Hoffmann 1962); Celaya, ears of
cattle (Hoffmann 1962). GUERRERO: Huitzuco, ears of cattle (Hoffmann 1962); Amealco, ears of cattle (Hoff-
mann 1962). HIDALGO: Zimapán, ears of cattle (Hoffmann 1962); Singuilucan, ears of cattle (Hoffmann 1962);
Tepeji del Río, ears of cattle (Hoffmann 1962); Chapantongo, ears of cattle (Hoffmann 1962) (CNAC001983);
Carrillo et al. 1966). JALISCO: Amatitlán, ears of cattle (Hoffmann 1962); San Juan de los Lagos, ears of cattle
(Hoffmann 1962); Ahualulco, ears of cattle (Hoffmann 1962); Nazareno, ears of donkeys (Hoffmann 1962). MI-
CHOACÁN: Morelia, cows (Bustamante et al. 1947); Ario de Rosales, cows (Bustamante et al. 1947); Pátzcuaro,
cows (Bustamante et al. 1947); Zamora, cows (Bustamante et al. 1947); Coalcomán, cows (Bustamante et al. 1947);
Uruapan, cows (Bustamante et al. 1947); Pajacuarán, ears of cattle (Hoffmann 1962); Jacona, ears of cattle (Hoff-
mann 1962). MORELOS: Ecatepec, ears of cattle (Hoffmann 1962). NUEVO LEÓN: Ojo de Agua, on the ground
(Cooley & Kohls 1944); ND, horses (Zarate-Ramos et al. 2014). OAXACA: Tacachi, Huajuapan, deer (Hoffmann
1962) (CNAC001980). PUEBLA: ND, ND (Hoffmann 1930); Puebla, ears of cattle (Gibson & Carrillo 1959); Am-
alucan, ears of goats (Hoffmann 1962); Tlahuapan, ears of sheep and cattle (Hoffmann 1962). QUINTANA ROO:
Nuevo Tabasco, Bacalar, ears of Tapirus bairdii (Pérez & González 2018). SINALOA: Las Cañas, ears of cattle
(Hoffmann 1962); El Fuerte, ears of dogs (Hoffmann 1962). SONORA: ND, ND, cattle (Cooley & Kohls 1944);
Tonichi, ears of dogs (Hoffmann 1962); Bacoachi, ears of dogs (Hoffmann 1962); Los Molinos, Altar, ears of dogs
(Hoffmann 1962); Río Sonora, Hermosillo, ears of cattle (Hoffmann 1962) (CNAC001989). TLAXCALA: Contla,
ears of cattle (Hoffmann 1962).
Notes: Hoffmann (1930) believed this species to occur in Puebla, in arid and semi-arid areas of central Mexico,
and along the Pacific Coast, parasitizing humans, cattle, horses, donkeys, sheep, cats, dogs, deer and other mam-
mals. Chavarría (1941) and Hoffmann & López-Campos (2000) mentioned that Ot. megnini is common and distrib-
uted throughout Mexico, where it is associated with humans and occurs in the ears of domestic animals.

Discussion

Of the approximately 211 species of argasids currently known, 36 (including O. dugesi) have previously been re-
corded from Mexico. Here, we add two new records for O. marinkellei associated with P. personatus, bringing to
37 the number of argasid species known from Mexico, a figure that represents 17.53% of the world argasid fauna
(Luz et al. 2018). Prior to our discovery of O. marinkellei, and our concurrent collections of A. mexicanus, Nayarit
was the only one of Mexico’s 32 states (including Ciudad de México) from which argasid tick records had been
missing.
The most widely distributed Mexican argasid species is Ot. megnini (21 states), followed by Ar. sanchezi and
O. talaje (18 states). By contrast, Ar. brevipes, Ar. cooleyi, Ar. giganteus, Ar. miniatus, O. denmarki, O. dusbabeki,
O. kelleyi, O. knoxjonesi, O. kohlsi, O. marinkellei, O. mormoops, O. parkeri, O. puertoricensis and O. stageri are
known only from single Mexican states.
Argasid ticks manifest a relatively high degree of host specificity. Thus, members of the genus Argas are
chiefly associated with birds, while species in the genera Antricola, Nothoaspis and Ornithodoros are mainly or
always found on bats and in bat roosts. However, the large genus Ornithodoros also includes species parasitic on
birds, for example O. capensis and O. denmarki, which have been collected mostly on islands. A dichotomy in host
preference is seen in the genus Otobius, where Ot. lagophilus chiefly parasitizes lagomorphs, while Ot. megnini is
generally associated with ungulates. Only six Mexican argasids are currently known to attack humans: Ar. persicus,
O. coriaceus, O. nicollei, O. talaje, O. turicata, and Ot. megnini (Hoffmann 1930; Mooser 1932; Chavarría 1941;
Hoffmann 1961, 1962; Hoffmann & López Campos 2000).
Of the 37 argasid species that have been recorded from Mexico, 11 have Nearctic affinities (Ar. brevipes, Ar.
cooleyi, Ar. giganteus, Ar. radiatus, Ar. sanchezi, O. denmarki, O. dugesi, O. kelleyi, O. parkeri, O. stageri, and Ot.
lagophilus); 10 are Neotropical (A. marginatus, N. reddelli, O. azteci, O. dusbabeki, O. knoxjonesi, O. kohlsi, O.
mormoops, O. marinkellei, O. puertoricensis, and O. viguerasi), and 16 species are found in both zoogeographic
regions (Figs. 6–10).

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FIGURE 6. Distribution of Antricola species in Mexico.

FIGURE 7. Distribution of Argas species in Mexico.

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FIGURE 8. Distribution of Nothoaspis reddelli in Mexico.

FIGURE 9. Distribution of Ornithodoros species in Mexico.

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FIGURE 10. Distribution of Otobius species in Mexico.

Ten of Mexico’s argasid species were described from specimens collected in this country, but only two of
these are represented by specimens deposited in CNAC: Ar. giganteus (paratype) and A. mexicanus (holotype and
paratype). The remaining types are in foreign collections: Ar. giganteus (United States National Tick Collection,
Statesboro, Georgia, USA (USNTC)), Ar. sanchezi (École Nationale Vétérinaire, Toulouse, France (ENV)), A. cop-
rophilus (United States National Parasite Collection, Beltsville, Maryland, USA (USNPC)), United States National
Museum, Washington, DC, USA (USNM), and USNTC, N. reddelli (USNTC), O. dugesi (Instituto de Salubridad y
Enfermedades Tropicales, now Instituto Nacional de Diagnóstico y Referencia Epidemiológica (INDRE) (holotype
lost), and USNTC), O. coriaceus (Zoologischen Museum, München, Germany (ZMM)), and O. nicollei (USNM).
The types of O. turicata and Ot. megnini were not specified in the original descriptions. We contacted the Museum
of Natural History Alfredo Dugès, where that author’s invertebrate collection is housed, but were told that the types
of these two species could not be found.
The determination of argasid specimens to species is largely based on morphological characters of the larvae
because this stage feeds at length and is therefore more likely to be collected from host animals, but also because lar-
vae possess numerous taxonomically useful structures, such as a rich chaetotaxy. However, in the present study even
larval determinations were sometimes dependent on subtle differences. For example, O. mormoops and O. viguerasi
were separated by the length of posthypostomal setae 1 and the number of hypostomal denticles; O. parkeri and
O. turicata were separated by the length of posthypostomal setae 1 and the length of the hypostome; and larvae of
O. dyeri and O. rossi proved impossible to separate. Kohls et al. (1965) differentiated the last-named pair of spe-
cies based on the length of the subtriangular median extension of the basis capituli; however, this is a subjective
character that cannot be accurately measured. Similarly, Kohls et al. (1965) separated O. talaje from O. kelleyi on
the basis of the relative abruptness of the junction of the subtriangular median extension of the basis capituli with
the hypostome, which is said to be abrupt in O. kelleyi but absent or only suggested in O. talaje; again, this feature
is difficult to discern, and for that reason we separate these species based on their hosts and the dimensions of their
dorsal plates.

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Collections of fresh specimens suitable for scanning electron microscopy (SEM) and the adoption of molecular
genetic methods hold promise for facilitating argasid determinations. Such tools will almost certainly prove valu-
able when attempting to associate larval, nymphal and adult stages, as in the case of O. viguerasi (Nava et al. 2012)
and O. dyeri (Guzmán-Cornejo et al. 2016). In many species, the nymphal and adult stages are unknown, although
in recent years additional developmental stages have been described and molecular techniques have increasingly
been brought to bear on problems in argasid taxonomy (Labruna et al. 2011; Dantas-Torres et al. 2012; Guzmán-
Cornejo et al. 2016).
Three Mexican species of Argas—Ar. miniatus, Ar. radiatus and Ar. sanchezi—were treated as synonyms of Ar.
sanchezi by Neumann (1899); however, after studying the type material of these species, Kohls et al. (1970) con-
cluded that they can be differentiated using larval characters and some characters of the nymphs and adults, such as
the form of the dorsal peripheral cells, the number of setiferous pits, the structure of the mammillae, the number of
setae on the basis capituli, and the length of legs IV. Muñoz-Leal et al. (2018) emphasized that although adults can
be differentiated morphologically through the fine structure of the dorsal integument, molecular characterization is
desirable to confirm species determinations. Particularly when working with fresh specimens, we strongly support
the use of both morphological and molecular methods, which should yield novel insights concerning the identity and
distribution of Mexican Argas species.
Only three species of Antricola have been reported from Mexico: A. coprophilus, A. marginatus, and A. mexica-
nus. The adults of A. marginatus and A. mexicanus are easily separated because the entire body margin of the former
species possesses prominent, digitiform, setiferous tubercles, whereas these tubercles are confined to the posterior
idiosomal margin in A. mexicanus. Hoffmann (1958) states that adults of A. mexicanus and A. coprophilus may be
separated by the presence in the former species of foliaceous, cordiform stigmal plates that protrude from the body,
as well as tufts of setae along the posterior margin of the idiosoma, whereas in A. coprophilus the stigmal plates are
suboval, not protruding from the body, and setal tufts are absent. However, the type of A. coprophilus is lost, which
limits our ability to propose diagnostic characters for this species. Larvae of A. marginatus possess claws, which are
lacking in A. mexicanus. No morphological characters have been found that distinguish larval A. mexicanus from
A. coprophilus.
Ornithodoros talaje (Figs. 3E, 3G-3I) is thought to be the second most widely distributed argasid in Mexico.
The original description of this species is very brief and includes few diagnostic characters (Guérin-Méneville
1849), with the result that adults of O. talaje are easily confused with O. dugesi and O. kohlsi. Mazzotti (1943a)
argued that the larger body and mammillae of O. talaje constitute the chief difference between this species and O.
dugesi. However, Fairchild et al. (1966) note that Kohls examined the larval and postlarval stages of O. dugesi and
O. talaje specimens from the United States, native homes in southern Mexico, and the type locality in Guatemala
but was unable to detect significant morphological differences between the two species, leading him to suggest that
O. dugesi may be a synonym of O. talaje. Subsequently, Buttler and Gibbs (1984) and Venzal et al. (2008) argued
that O. dugesi may represent a valid species, but fresh specimens would be needed from northeastern Mexico and
the southern United States in order to properly evaluate the validity of these taxa. Further complicating this issue is
the fact that the type of O. dugesi is lost.
The characters used to describe O. kohlsi are similar to those that appear in the redescription of O. talaje by
Cooley and Kohls (1944). The chief difference cited for separating these species is the size and arrangement of the
dorsal discs (Kohls & Clifford 1964), which, in the absence of additional specimens, may be a subjective judgment.
Clearly, additional studies are needed to clarify the diagnostic characters of O. talaje, assess the validity of O. dugesi
and O. kohlsi, and define the geographic distribution of these species. In light of this, the distribution of O. talaje as
currently understood may be overestimated, but most of our records are from older literature, and in the absence of
voucher specimens, corroboration of such records is impossible.
The family Argasidae includes a large number of species that are of veterinary or medical importance. For ex-
ample, Ot. megnini, known as the “spinose ear tick,” commonly infests the ear canal of domestic and wild mammals,
as well as humans, causing irritation, allergic or toxic reactions, paralysis, eardrum perforation, muscle spasms,
myotonia and severe otitis (www.sciencedirect.com/topics/immunology-and microbiology/otitis) (Estrada-Peña &
Jongejan 1999; Naudé et al. 2001; Zarate-Ramos et al. 2014; Diyes & Rajakaruna, 2017). This species has also been
considered a potential vector of Q fever (Jellison et al. 1948). However, argasids are particularly important as vec-
tors of pathogens causing human disease. Especially in Mexico, bacteria in the genera Borrelia and Rickettsia have
been detected in various argasids, e.g., Borrelia turicatae from O. turicata (Brumpt et al. 1939), Rickettsia rickettsii

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from O. nicollei and Ot. lagophilus (Silva-Goytia & Elizondo 1952), and Rickettsia lusitaniae from O. yumatensis
(Sánchez-Montes et al. 2016).

Host-Parasite List

“Crevices in walls”
Ornithodoros brodyi
Ornithodoros yumatensis
“Cave walls”
Ornithodoros dyeri
“Domestic animals”
Ornithodoros talaje
“Ears of domestic animals”
Otobius megnini
“Houses”
Ornithodoros coriaceus
Ornithodoros talaje
Ornithodoros turicata
“Huts of natives”
Ornithodoros nicollei
Ornithodoros talaje
“Nests of wild rats”
Ornithodoros dugesi
“Nests of Neotoma albigula”
Ornithodoros dugesi
“On soil”
Ornithodoros nicollei
Ornithodoros talaje
“On the floor”
Ornithodoros talaje
“Under rocks”
Ornithodoros denmarki
Ornithodoros talaje
“Under rocks near nesting gulls and terns”
Ornithodoros denmarki
“Under wood”
Ornithodoros talaje

AVES
“Birds”
Argas giganteus
Ornithodoros turicata
“Chicken coops”
Argas persicus
Argas radiatus
Argas sanchezi
“Wild turtle dove”
Argas sanchezi
“Woodpecker holes”
Argas brevipes
Columbiformes

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Columbidae
Zenaida macroura carolinensis (L.)
Argas sanchezi
Galliformes
Phasianidae
Hens (= Gallus gallus domesticus L.)
Argas miniatus
Argas persicus
Argas sanchezi
Passeriformes
Corvidae
Corvus imparatus Peters
Argas giganteus
Hirundinidae
Tachycineta thalassina thalassina Mearns
Argas brevipes
Mimidae
Toxostoma bendirei (Coues)
Argas giganteus
Toxostoma curvirostre (Swainson)
Argas giganteus
Passerellidae
Kieneria fusca (Swainson)
Argas giganteus
Pelecaniformes
Pelecanidae
Pelecanus occidentalis L.
O. capensis
Sulidae
Near nest of Sula dactylatra californica Rothschild
O. capensis
Strigiformes
Strigidae
Athene cunicularia (Molina)
Ornithodoros puertoricensis

MAMMALIA
“Bat guano”
Antricola coprophilus
Antricola marginatus
Antricola mexicanus
Nothoaspis reddelli
Ornithodoros brodyi
Ornithodoros dyeri
Ornithodoros yumatensis
“Mammals”
Ornithodoros turicata
“Small mammals”
Ornithodoros turicata
Artiodactyla
“Cattle”
Ornithodoros talaje

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Ornithodoros turicata
Otobius megnini
Bovidae
Cows (= Bos primigenius taurus L.)
Otobius megnini
Sheep (= Ovis orientalis aries L.)
Ornithodoros turicata
Suidae
Pigs (= Sus scrofa domesticus Erxleben)
Ornithodoros talaje
Ornithodoros turicata
Carnivora
Canidae
Dog (= Canis lupus familiaris L.)
Ornithodoros nicollei
Otobius megnini
Felidae
Cat (= Felis silvestris catus Schreber)
Otobius megnini
Chiroptera
“Walls of tunnels inhabited by bats”
Ornithodoros stageri
Emballonuridae
Balantiopteryx plicata Peters
Antricola mexicanus
Ornithodoros dyeri
Ornithodoros knoxjonesi
Peropteryx macrotis (J. A. Wagner)
Ornithodoros dyeri
Molossidae
Molossus rufus É. Geoffroy Saint-Hilaire
Ornithodoros dusbabeki
Molossus rufus nigricans Miller
Ornithodoros hasei
Molossus sinaloae J. A. Allen
Ornithodoros kohlsi
Tadarida brasiliensis mexicanus (de Saussure)
Ornithodoros kelleyi
Mormoopidae
Mormoops megalophylla (Peters)
Ornithodoros viguerasi
Pteronotus davyi fulvus (Thomas)
Antricola mexicanus
Pteronotus parnellii (Gray)
Antricola mexicanus
Ornithodoros brodyi
Ornithodoros dyeri
Ornithodoros setosus
Pteronotus personatus (J. A. Wagner)
Ornithodoros marinkellei
Pteronotus personatus psilotis (Dobson)
Ornithodoros setosus

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Phyllostomidae
Artibeus jamaicensis Leach
Ornithodoros brodyi
Ornithodoros yumatensis
Artibeus jamaicensis yucatanicus J. A. Allen
Ornithodoros azteci
Artibeus lituratus (Olfers)
Ornithodoros brodyi
Artibeus lituratus palmarum J. A. Allen & Chapman
Ornithodoros yumatensis
Carollia sowelli R. J. Baker, Solari & Hoffmann
Ornithodoros brodyi
Chrotopterus auritus auritus (Peters)
Ornithodoros brodyi
Dermanura azteca (Andersen)
Ornithodoros yumatensis
Desmodus rotundus (M. Geoffroy Saint-Hilaire)
Ornithodoros azteci
Ornithodoros brodyi
Ornithodoros yumatensis
Desmodus rotundus murinus J. A. Wagner
Ornithodoros yumatensis
Diphylla ecaudata Spix
Ornithodoros yumatensis
Glossophaga sp.
Ornithodoros rossi
Leptonycteris sp.
Ornithodoros rossi
Macrotus californicus Baird
Ornithodoros rossi
Macrotus waterhousii Gray
Antricola coprophilus,
Ornithodoros azteci
Ornithodoros dyeri
Ornithodoros mormoops
Ornithodoros rossi
Ornithodoros yumatensis
Mimon cozumelae Goldman
Ornithodoros dyeri
Trachops cirrhosus coffini Goldman
Ornithodoros dyeri
Sturnira sp.
Ornithodoros rossi
Vespertilionidae
Corynorhinus mexicanus (G. M. Allen)
Ornithodoros yumatensis
Myotis nigricans extremus (Miller & G. M. Allen)
Ornithodoros brodyi
Myotis velifer (J. A. Allen)
Ornithodoros hasei
Myotis vivesi Menegaux
Ornithodoros dyeri

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Parastrellus hesperus (H. Allen)
Ornithodoros rossi
Lagomorpha
Leporidae
“Hares”
Otobius lagophilus
“Rabbits”
Ornithodoros turicata
Otobius lagophilus
Perissodactyla
Equidae
“Mules”
Otobius megnini
Donkeys (= Equus africanus asinus L.)
Otobius lagophilus
Horses (= Equus ferus caballus L.)
Ornithodoros talaje
Otobius megnini
Tapiridae
Tapirus bairdii (Gill)
Otobius megnini
Primates
Hominidae
Humans (= Homo sapiens L.)
Ornithodoros nicollei
Ornithodoros talaje
Ornithodoros turicata
Otobius megnini
Rodentia
“Moles”
Ornithodoros turicata
“Rats”
Ornithodoros turicata
“Rodents”
Ornithodoros talaje
“Squirrel”
Ornithodoros nicollei
“Tuzas”
Ornithodoros turicata
Cricetidae
Hodomys alleni (Merriam)
Ornithodoros nicollei
Neotoma sp.
Ornithodoros nicollei
Ornithodoros parkeri
Heteromyidae
Dipodomys deserti Stephens
Ornithodoros talaje
Sciuridae
Spermophilus tereticaudus (Baird)
Ornithodoros talaje

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REPTILIA
“Reptiles”
Ornithodoros turicata

LIST OF LOCALITIES

LOCALITY, MUNICIPALITY Latitude N Longitude W


AGUASCALIENTES
Aguascalientes, Aguascalientes 21°52’47.36’’ 102°17’45.77’’
Calvillo, Calvillo 21°50’48.87’’ 102°43’07.5’’
Jesús María, Jesús María 21°57’40.58’’ 102°20’36.3’’
BAJA CALIFORNIA
8.8 km S of San Felipe*, Mexicali 31°01’22.51’’ 114°50’20.04’’
45 km S of Punta Prieta*, Ensenada 28°55’40.76’’ 114°09’14.32’’
Isla Calaveras, Mexicali 29°01′39.6'' 113°29′55.5''
Isla Partida, Mexicali 28°52’0’’ 113°02’0’’
Isla Pescadero ND ND
Isla Pond, Mexicali 29°03′55.71'' 113°05′ 15.04''
Isla Raza (Isla Rasa), Mexicali 28°49′24.79'' 112°58’48.12''
La Acequia 32°36′25.93'' 115°28’17.38''
Los Pozos, Los Cabos ND ND
Misión Santa Agueda, Ensenada 28°10’10'' 113°04’50''
Near Santiago (Santiago, Ensenada) 31°49’07'' 116°25’30''
BAJA CALIFORNIA SUR
Las Cuevas, Los Cabos 23°32’4.67’’ 109°40’28.51’’
Miraflores, Los Cabos 23°22’0.53’’ 109°46’41.59’’
San Antonio ND ND
CAMPECHE
About 4.8 km S of Champoton* 19°21’22.39’’ 090°43’26.1’’
Grutas de Xtacumbilxunam, 2 km SW of Bolonchenticul* 20°00’15.64’’ 089°44’52.1’’
CHIAPAS
2.1 km SSE of Zapaluta* 16°07’8.61’’ 092°03’1.69’’
Acacoyagua, Acacoyagua 15°20’27.47’’ 092°40’28.62’’
Acala, Acala 16°33’6.02’’ 092°48’27.03’’
Ángel Albino Corzo, Ángel Albino Corzo 15°52’21.52’’ 092°43’30.29’’
Arriaga, Arriaga 16°14’9.83’’ 093°53’58.1’’
Cintalapa, Cintalapa 16°41’55.25’’ 093°43’13.54’’
Chicomuselo, Chicomuselo 15°44’41.77’’ 092°16’59.83’’
Cueva de Zapaluta* 16°07’8.61’’ 092°03’1.69’’
Cueva del Guano, San Fernando 16°48’37.31'' 093°10’30.3''
Cueva de El Naranjo (Rancho “El Arco”), Cintalapa 16°51’6.8'' 093°42’16.3''
Cueva de La Chepa (2 km N, 4 km NE of Tuxtla Gutiérrez), Tuxtla Gutiér- 16°46’41'' 093°8’55''
rez
El Vergel ND ND
Escuintla, Escuintla 15°19’9.37’’ 092°39’30.81’’
Huixtla, Huixtla 15°08’16.9’’ 092°27’57.64’’
Ixtapa, Ixtapa 16°48’12.69’’ 092°54’21.25’’
Mapastepec, Mapastepec 15°26’28.1’’ 092°53’29.78’’
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LOCALITY, MUNICIPALITY Latitude N Longitude W
Pijijiapan, Pijijiapan 15°41’30.02’’ 093°12’33.54’’
Planta de Luz, Huixtla ND ND
Simojovel, Simojovel 17°08’26.97’’ 092°42’54.75’’
Tapachula, Tapachula 14°54’39.86’’ 092°15’51.55’’
Tecpatán, Tecpatán 17°08’11.32’’ 093°18’40.19’’
Tonalá, Tonalá 16°05’28.88’’ 093°45’6.42’’
Tuxtla Gutiérrez, Tuxtla Gutiérrez 16°45’14.3’’ 093°06’54.69’’
Yajalón, Yajalón 17°10’26.08’’ 092°20’2.2’’
CHIHUAHUA
Bocoyna, Bocoyna 27°50’26.67’’ 107°35’21.31’’
Camargo, Camargo 27°41’41.86’’ 105°10’17.83’’
Ciudad Juárez, Juárez 31°44’47.27’’ 106°29’6.067’’
Chihuahua, Chihuahua 28°38’12.46’’ 106°04’34.7’’
Cueva del Salitre, 13 km W of Villa de Matamoros* 26°45’49.96’’ 105°34’59.99’’
Delicias, Delicias 28°11’36’’ 105°28’16’’
Rosales, Rosales 28°11’13’’ 105°33’16’’
Sótano del Sauz (30 km NW of Ejido M. Benavides and 20 km S of Lajitas) ND ND
CIUDAD DE MÉXICO
Azcapotzalco, Azcapotzalco 19°29’2.77’’ 099°11’3.69’’
Castillo de Chapultepec 19°25’13.58’’ 099°11’3.69’’
Col. Santo Tomás ND ND
Los Morales ND ND
San Juan de Aragón (Cerca de Guadalupe Hidalgo), Gustavo A. Madero 19°28’07.1’’ 099°10’54.97’’
Tacuba ND ND
Tlalpan 19°16’44.72’’ 099°12’41.12’’
Xochimilco, Xochimilco 19°15’48.46’’ 099°06’17.69’’
COAHUILA
Agua Nueva, Saltillo 25°12’59’’ 101°05’22’’
Carneros, Saltillo 25°07’19.22’’ 101°06’38.3’’
Coahuila ND ND
Coyote, Región Lagunera ND ND
Cueva del Detector Perdido, Monclova 26°58’27.07’’ 101°03’29.84’’
Jalisco, Torreón 25°13’1.12’’ 103°26’27.94’’
La Flor de Jimulco, Torreón 25°06’12.41’’ 103°19’53.45’’
Mariano Escobedo, Acuña 29°16’12.4’’ 102°10’4.32’’
Matamoros, Matamoros 25°31’41.29’’ 103°13’49.38’’
Mina de San Felipe, N of Sabinas* 27°50’55.1’’ 101°07’11.96’’
Ramos Arizpe, Ramos Arizpe 25°32’35.64’’ 100°57’2.81’’
Sabinas, Sabinas 27°50’55.1’’ 101°07’11.96’’
Saltillo, Saltillo 25°25’17.99’’ 100°59’58.1’’
San Pedro de las Colonias, San Pedro de las Colinas 25°45’28’’ 102°58’59.36’’
COLIMA
Colima, Colima 19°14’34.52’’ 103°43’41.23’’
Coquimatlán, Coquimatlán 19°12’3.53’’ 103°48’37.95’’
Cueva de los Amiales, Jala 19°08’57.84’’ 103°49’58.8’’
Cueva El Salitre, Los Ortices 19°04’0’’ 103°43’0’’
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LOCALITY, MUNICIPALITY Latitude N Longitude W
Cueva de la Fábrica (5 km W of Coquimatlán*) 19°12’3.53’’ 103°48’37.95’’
Isla Clarión (Islas Revillagigego), Manzanillo 18°21’20.29'' 114°43'31.91''
Isla San Benedicto (Islas Revillagigego), Manzanillo 19°18’30.68'' 110°48'39.75''
Manzanillo, Manzanillo 19°03’11.54’’ 104°18’58.97’’
DURANGO
Gómez Palacio 25°35’20.93’’ 103°29’09.1’’
Lerdo, Ciudad Lerdo 25°32’11.59’’ 103°31’29.46’’
Nombre de Dios, Nombre de Dios 23°50’53.42’’ 104°14’41.16’’
San José de Zaragoza, General Simón Bolívar 25°13’37.46’’ 103°28’4.56’’
San Pedro del Gallo, San Pedro del Gallo 25°33’57.11’’ 104°17’34.2’’
ESTADO DE MÉXICO
Acambay, Acambay 19°57’22.42’’ 099°50’38.51’’
Chimalhuacán, Chimalhuacán 19°25’0.87’’ 098°56’43.1’’
Pueblo Nuevo ND ND
San Mateo Atenco, San Mateo Atenco 19°15’51’’ 099°31’47’’
Santa Clara de Juárez, Morelos 19°43’18.32’’ 099°36’31.14’’
Santa María, Huehuetoca 19°50’30.14’’ 099°12’20.93’’
Santa María Tulpetlac, Ecatepec 19°34’27’’ 099°05’56’’
Temascalapa, Temascalapa 19°49’40.03’’ 098°54’3.21’’
Texcoco, Texcoco 19°30’48.38’’ 098°52’53.34’’
Valle de México ND ND
GUANAJUATO
Apaseo, Apaseo el Grande 20°32’43.02’’ 100°41’17.58’’
Celaya, Celaya 20°31’17.49’’ 100°48’48.86’’
Guanajuato, Guanajuato 21°00’57.95’’ 101°15’13.04’’
Irapuato, Irapuato 20°40’22.08’’ 101°20’53.22’’
San Francisco del Rincón, San Francisco del Rincón 21°01’7.25’’ 101°51’30.66’’
San Nicolás de los Agustinos, Salvatierra 20°14’44.05’’ 100°58’1.144’’
Valle de Santiago, Valle de Santiago 20°23’28.76’’ 101°10’53.15’’
GUERRERO
Amealco, Ixcateopan 18°28’40.06’’ 099°42’55.27’’
Arcelia, Arcelia 18°18’56.97’’ 100°17’2.23’’
Cuadrilla Cuaulotitlán, Tlalchapa 18°20’41.65’’ 100°19’40.98’’
Cueva Juxtlahuaca, Quelchultenango* 17°25’40.47’’ 099°08’45.26’’
“El Potrero,” Tlalchapa, Tlalchapa 18°24’39.07’’ 100°26’58.08’’
Grutas de Cacahuamilpa, Pilcaya 18°40’19.87’’ 099°30’38.24’’
Huitzuco, Huitzuco 18°18’21.53’’ 099°20’3.49’’
Iguala, Iguala de la Independencia 18°20’40.38’’ 099°32’18.13’’
Mezcala, Eduardo Neri 17°55’54.81’’ 099°36’16.5’’
Olinalá, Olinalá 17°46’41.1’’ 098°44’28.52’’
Ometepec, Ometepec 16°41’6.32’’ 098°24’15.92’’
Tlachichilpa, Arcelia 18°20’54.54’’ 100°17’0.51’’
Tuxpan, Iguala de la Independencia 18°20’57.68’’ 099°28’44.09’’
Río Balsas, Cocula 18°0’28.15’’ 099°46’6.16’’
Xalitla, Tepeacocuilco de Trujano 17°59’52.1’’ 099°32’25.61’’
HIDALGO
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LOCALITY, MUNICIPALITY Latitude N Longitude W
Chapantongo, Chapantongo 20°17’7.14’’ 099°24’47.63’’
Singuilucan, Singuilucan 19°58’03’’ 098°31’02’’
Tepeji del Río, Tepeji del Río 19°54’18.78’’ 099°20’30.53’’
Zimapán, Zimapán 20°44’17.05’’ 099°22’59.21’’
JALISCO
9.6 km E of Limón 19°45’0’’ 104°01’0’’
27 km NE of Tamazula, Tamazula de Gordiano* 19°40’35’’ 103°15’0’’
Ahualulco de Mercado 20°42’02’’ 103°58’3’’
Amatitlán Sayula 19°52’35.25’’ 103°31’42.67’’
Autlán, Autlán de Navarro 19°46’17’’ 104°21’55’’
Cueva “El Chico,” 5 km N of Tuxcacuesco* 19°41’50’’ 103°59’1’’
Encarnación de Díaz, Encarnación de Díaz 21°31’28’’ 102°13’59’’
Guadalajara, Guadalajara 20°40’35’’ 103°20’32’’
Jalostotitlán, Jalostotitlán 21°10’7’’ 102°27’40’’
Los Altos ND ND
Nazareno ND ND
San Juan de los Lagos 21°14’52’’ 102°19’58’’
Zapotitlán de Vadillo 19°32’51’’ 103°48’33’’
MICHOACÁN
Ario de Rosales, Ario 19°12’28.11’’ 101°42’33.82’’
Carácuaro, Carácuaro 19°01’2.31’’ 101°07’37.17’’
Chucándiro, Chucándiro 19°53’59.87’’ 101°20’0.21’’
Coalcomán, Coalcomán de Vázquez Pallares 18°46’37.56’’ 103°09’34.4’’
El Jorullo ND ND
Morelia, Morelia 19°42’12.18’’ 101°11’31.37’’
Pajacuarán, Pajacuarán 20°07’0.82’’ 102°33’55.85’’
Pátzcuaro, Pátzcuaro 19°30’48.95’’ 101°36’36.42’’
Turicato, Tecámbaro 19°03’13.92’’ 101°25’10.31’’
Uruapan, Uruapan 19°25’15.19’’ 102°03’46.08’’
Zamora, Zamora 19°59’0.89’’ 102°17’9.45’’
MORELOS
Amacuzac, Amacuzac 18°36’0’’ 099°22’10.25’’
Atlihuayán, Yautepec 18°51’46’’ 099°04’18’’
Cueva Poza de Moctezuma, Yautepec* 18°53’9.822’’ 099°03’38.71’’
Cuevas de Huajintlán, 0.5 km NW of Huajintlán* 18°36’35.10’’ 099°25’27’’
Ecatepec, Yecapixtla 18°50’51.17’’ 098°52’45.63’’
Huajintlán, Amacuzac 18°36’35.10’’ 099°25’27’’
Oaxtepec, Yautepec 18°54’27.59’’ 098°58’11.34’’
Yautepec, Yautepec 18°53’9.82’’ 099°03’38.71’’
Yecapixtla, Yecapixtla 18°53’0.31’’ 098°52’5.32’’
NAYARIT
Cueva Cerro de las Viejas, Santiago Ixcuintla 22°51’55.8’’ 105°09’00.72’’
NUEVO LEÓN
China, China 25°42’21.29’’ 099°14’21.54’’
Cueva en Sabinas Hidalgo* 26°29’54.13’’ 100°10’52.13’’
General Bravo 25°47’35.87’’ 099°10’52.44’’
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LOCALITY, MUNICIPALITY Latitude N Longitude W
Linares, Linares 24°51’34.75’’ 099°34’4.33’’
Los Aldamas, Los Aldamas 26°03’50.92’’ 099°11’48.24’’
Los Herreras, Los Herreras 25°54’20.18’’ 099°24’11.87’’
Near Monterrey* 25°39’52.91’’ 100°18’39.21’’
Ojo de Agua ND ND
Villaldama, Villaldama 26°29’53.61’’ 100°25’28.42’’
OAXACA
Cacahuatepec, San Juan Cacahuatepec 16°36’49.44’’ 098°09’21.93’’
Calihuala, Silac ND ND
Ciudad Ixtepec, Ixtepec 16°33’35.61’’ 095°07’11.26’’
El Espinal, El Espinal 16°29’8.55’’ 095°02’34.21’’
Huajuapan, Huajuapán de León 17°48’24.61’’ 097°46’34.45’’
Jamiltepec, Santiago Jamiltepec 16°16’46.05’’ 097°49’10.86’’
Juchitán de Zaragoza 16°25’59.91’’ 095°01’17.88’’
Matías Romero, Matías Romero Avendaño 16°52’40.40’’ 095°02’31.74’’
Puerto Ángel, San Pedro Pochutla 15°39’55.17’’ 096°29’24.27’’
Putla, Putla Villa de Guerrero 17°01’34.34’’ 097°55’41.35’’
Salina Cruz 16°10’33.19’’ 095°11’31.77’’
San Juan Zaichicoban ND ND
San Pedro Huilotepec, San Pedro Huilotepec 16°14’37.17’’ 095°09’10.56’’
San Pedro Mixtepec, Juquila 15°55’0’’ 097°04’0’’
San Pedro Totolapa, San Pedro Totolapa 16°39’59.2’’ 096°18’6.86’’
Santo Domingo Tehuantepec 16°19’55.41’’ 095°13’55.38’’
Silac (Silacayoápam*) 17°30’5.21’’ 098°08’33.31’’
Tacachi, Huajuapan ND ND
Tequisistlán, Magdalena Tequisistlán 16°23’56.10’’ 095°36’10.47’’
Tlacolula de Matamoros, Tlacolula de Matamaros 16°57’20.25’’ 096°28’45.17’’
Villa Flores ND ND
PUEBLA
Amalucan, Puebla 19°03’27.89'' 098°06’38.84’’
Atencingo, Chietla 18°30’42.81’’ 098°36’26.38’’
Atexcal 18°23’54.06’’ 097°44’12.54’’
Cueva El Santurario, Tzinacatepec 18°19’0’’ 097°14’0’’
Huaquechula, Huaquechula 18°46’14.55’’ 098°32’29.51’’
Matamoros (Izúcar de Matamoros) 18°36’8.032’’ 098°27’55.46’’
Puebla, Puebla de Zaragoza 19°02’38.36’’ 098°11’50.98’’
Tecamachalco, Tecamachalco 18°53’4.89’’ 097°43’42.3’’
Tecomatlán, Tecomatlán 18°06’39.46’’ 098°18’51.74’’
QUERÉTARO
San Clemente ND ND
San Juan del Río, San Juan del Río 20°23’11.19’’ 099°59’58.67’’
Querétaro, Querétaro 20°35’36.21’’ 100°23’23.85’’
QUINTANA ROO
Chetumal, Othón P. Blanco (We provide the coordinates of Chetumal for 18°29’38’’ 088°17’52.44’’
the three localities cited by Wolfang & Polaco (1985))
Cueva de Zereque (Actuun Tzuub), Lázaro Cárdenas 20°48’15'' 087°28’34’’
Isla Contoy, Isla Mujeres 21°28’30’’ 086°47’15’’
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LOCALITY, MUNICIPALITY Latitude N Longitude W
Nuevo Tabasco, Bacalar 19°08ʹ 20.66ʺ 089°02ʹ50.76ʺ
Puerto Juárez 21°10’00.88’’ 086°48’40.81’’
Vicinity of Pueblo XCan (Nuevo XCán), Lázaro Cárdenas 20°52’11.75’’ 087°36’6.27’’
SAN LUIS POTOSÍ
Bocas, San Luis Potosí 22°31’13.18’’ 101°01’19.27’’
Cueva de Taninul (13 km SE of Ciudad Valles), Ciudad Valles* 21°59’11.53’’ 099°01’7.4’’
Cueva de los Sabinos, Ciudad Valles 22°05’30.84’’ 098°57’24.84’’
San Luis Potosí, San Luis Potosí 22°09’5.96’’ 100°58’37.05’’
Santa María del Río, Santa María del Río 21°48’1.58’’ 100°44’13.59’’
Sótano de la Tinaja (10.5 km NE of Ciudad Valles*) 21°59’11.53’’ 099°01’7.4’’
Tamazunchale, Tamazunchale 21°15’45.86’’ 098°47’29.46’’
SINALOA
2.4 km N of Badiraquato (Badiraguato*) 25°21’45.55’’ 107°33’3.67’’
Aguacaliente Grande, Choix 26°32’12.08’’ 108°20’55.09’’
Ampelio (Las Cañas), El Fuerte 26°28’58.53’’ 108°35’23.46’’
Cacalotán, Rosario 23°04’4.34’’ 105°50’31.81’’
Chinobampo, El Fuerte 26°23’19.4’’ 108°21’51.59’’
Choix, Choix 26°42’25.24’’ 108°19’20.47’’
Copala, Concordia 23°23’41.12’’ 105°55’58.53’’
El Disparate (Antonio Rosales), Choix 26°33’50.99’’ 108°24’42.02’’
El Fuerte, El Fuerte 26°25’17.76’’ 108°37’11.88’’
El Parmaso, El Fuerte 26°04’19.79’’ 108°54’33.6’’
Higuera de los Natoches, El Fuerte ND ND
La Palma ND ND
Las Cañas, El Fuerte 26°28’58.53’’ 108°35’23.46’’
Los Mochis 25°47’25.68’’ 108°59’09.17’’
Mazatlán, Mazatlán 23°12’1.137’’ 106°25’19.1’’
Ocolome, El Fuerte 26°26’55.22’’ 108°36’27.67’’
Pánuco, Concordia 23°25’24.47’’ 105°54’10.16’’
San Felipe ND ND
San Joaquín ND ND
Santa Ana, Choix 26°35’25.09’’ 108°23’2.72’’
Santa Lucía, Concordia 23°26’6.92’’ 105°51’0.07’’
Soyatita, Badiraguato 25°44’25.54’’ 107°18’33.95’’
SONORA
22.5 km W of Santa Ana* 30°32’39.65’’ 111°07’18.41’’
30.6 km S of Sonoyta* 31°45’9.02’’ 112°40’7.99’’
64.4 km S of Navojoa* 27°04’53.75’’ 109°26’43.97’’
Agua Blanca, Etchojoa 27°06’53.38’’ 109°44’01.93’’
Bacoachi, Bacoachi 30°37’59.79’’ 109°58’9.37’’
Carrizal ND ND
Corral, Cajeme 27°37’47.54’’ 109°57’59.51’’
Cueva de El Tigre, Carbó 29°34’0’’ 110°49’0’’
Isla del Medio, Guaymas ND ND
Isla Tiburón, Hermosillo 29°00’58’’ 112°12’5’’
Kino (Bahía de Kino), Hermosillo 28°49’31.88’’ 111°56’22.86’’
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LOCALITY, MUNICIPALITY Latitude N Longitude W
La Dura ND ND
La Calera (Cumuripa), Cajeme 28°09’7.68’’ 109°54’34.54’’
Los Molinos, Altar 31°08’28.89’’ 111°45’18.83’’
Rancho Carrizal, N of Altar 31°03’11.4’’ 111°42’8.41’’
Río Sonora, Hermosillo 29°05’35.3’’ 111°44’7.11’’
Tónichi, Soyopa 28°35’51.71’’ 109°33’57.07’’
TABASCO
Cueva Agua Blanca (Grutas Ixta-Ha), Macuspana 17°37’11’’ 092°28’12’’
Cueva del Azufre, 3.5 km S of Tapijulapa (= Cueva de las Sardinas, Cueva 17°26’32.53’’ 092°46’31.19’’
de Villa Luz)
TAMAULIPAS
32.2 km S of Nuevo Laredo* 27°29’10.65’’ 099°30’30.57’’
Cueva de La Florida (7.5 km NE of Antiguo Morelos*) 22°32’57.91’’ 099°05’1.02’’
Cueva de los Cuarteles, Aldama* 22°55’9.91’’ 098°04’24.58’’
Cueva de los Vampiros, 10 km NE of Adolfo López Mateos ND ND
Matamoros, Matamoros 25°52’47.69’’ 097°30’17.36’’
Tampico, Tampico 22°12’57.82’’ 097°51’30.19’’
TLAXCALA
Contla 19°20’4.26’’ 098°10’4.46’’
VERACRUZ
Catemaco, Catemaco 18°25’28.61’’ 095°06’44.42’’
Cueva cercana a Playa Vicente* 17°49’46.61’’ 095°48’47.57’’
Cuevas de Laguna Encantada, San Andrés Tuxtla* 18°26’55.21’’ 095°12’45.25’’
Cueva Rey del Oro, Emiliano Zapata* 19°21’55.97’’ 096°39’25.51’’
Los Tuxtlas, San Andrés Tuxtla 18°26’55.21’’ 095°12’45.25’’
YUCATÁN
Arrecife Alacranes 22°22’58.13’’ 089°40’58.29’’
Cave near Cenote Bal-mil, Homún 20°45’10’’ 089°14’65’’
Cenote Yuncu, Abalá* 20°38’51.28’’ 089°40’51.19’’
Chichén Itzá, Tinum 20°40’43.05’’ 088°34’4.25’’
Cueva Calcehtok (Cueva Xpukil), Opichén 20°33’02’’ 089°54’43’’
Cueva Chocantes, Tekax 20°12’10’’ 089°17’58’’
Cueva Kaua, Kaua 20°37´36’’ 089°24’85’’
Cueva Tzabnah, Tecoh 20°43´81’’ 089°28’46’’
Cueva de Balaam Canché (= Grutas de Balankanché), Tinum* 20°46’1.71’’ 088°23’28.45’’
Cueva de Sabacá, Tekax* 20°12’12.47’’ 089°17’18.2’’
Cueva de Hoctum, Hoctún* 20°51’54.76’’ 089°12’7.44’’
Cueva del Cenote Xtoloc (Chichén Itzá), Tinum 20°40’43.05’’ 088°34’4.25’’
Cueva El Naranjal, Tekax 20°11’56’’ 089°18’36’’
Cueva de la Santa María, Homún 20°44’49’’ 089°17’47’’
Cueva Toh, 19.5 km SE of Oxkutzcab* 20°18’6.96’’ 089°25’7.51’’
Cueva Xpukil, 3 km S of Calcetok, Opichén* 20°33’2.34’’ 089°51’22.48’’
Isla Pérez, Progreso 22°22’58.13’’ 089°40’58.29’’
Km 56, Santa Elena-Loltún road (Cavity I) 20°17’34’’ 089°38’42’’
Km 56, Santa Elena-Loltún road (Cavity II) 20°17’34’’ 089°38’42’’
Pisté 20°41’55.28’’ 088°35’21.42’’
ZACATECAS
Espíritu Santo ND ND

THE SOFT TICKS OF MEXICO Zootaxa 4623 (3) © 2019 Magnolia Press · 519
Acknowledgments

We thank Laura Del Castillo-Martínez for her assistance in mounting larval ticks and for helping with field col-
lections. Griselda Montiel-Parra also assisted with our field collections and arranged important loans of specimens
from the Colección Nacional de Ácaros, Instituto de Biología, UNAM. Ana Isabel Bieler Antolín produced our
conventional light photomicrographs, while Georgina Ortega Leite provided bibliographical support. Lorenza Beati
arranged critical loans of specimens from the USNTC and permitted us use of her collection facilities. Financial
support for this study was provided by the Programa de Apoyo a Proyectos de Investigación e Innovación Tec-
nológica of DGAPA-UNAM (PAPIIT-IN214114).

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