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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
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Distribution and host data from published literature and previously unpublished collection records are provided for all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.
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
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).
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)
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
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)
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)
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
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
THE SOFT TICKS OF MEXICO Zootaxa 4623 (3) © 2019 Magnolia Press · 489
Otobius (after Jones & Clifford 1972)
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
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.
Parasite-Host List
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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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(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.
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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.
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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).
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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).
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.
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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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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
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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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
LIST OF LOCALITIES
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Continued.
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
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
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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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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