Parasite 2015, 22, 29
Ó J.M. Caspeta-Mandujano et al., published by EDP Sciences, 2015
DOI: 10.1051/parasite/2015029
urn:lsid:zoobank.org:pub:4C3DCC0F-F1DD-4CB5-9554-D779F1F6419D
Available online at:
www.parasite-journal.org
RESEARCH ARTICLE
OPEN
ACCESS
A new species of Torrestrongylus (Trichostrongylidae,
Anoplostrongylinae) from Macrotus waterhousii (Chiroptera:
Phyllostomidae) in Central Mexico
Juan Manuel Caspeta-Mandujano1,2,*, Jorge Luis Peralta-Rodríguez3, María Guadalupe Galindo-García1,
and Francisco Agustín Jiménez4
1
2
3
4
Laboratorio de Parasitología de Animales Silvestres, Facultad de Ciencias Biológicas, Universidad Autónoma del Estado de Morelos,
Av. Universidad No. 1001, Col. Chamilpa, C.P. 62210 Cuernavaca, Morelos, Mexico
Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de Morelos, Av. Universidad No. 1001, Col. Chamilpa,
C.P. 62210 Cuernavaca, Morelos, Mexico
Facultad de Ciencias Agropecuarias, Universidad Autónoma del Estado de Morelos, Av. Universidad No. 1001, Col. Chamilpa,
C.P. 62210 Cuernavaca, Morelos, Mexico
Department of Zoology, Southern Illinois University, 62901-6501 Carbondale, Illinois, USA
Received 10 August 2015, Accepted 8 October 2015, Published online 29 October 2015
Abstract – A new species of nematode, Torrestrongylus tetradorsalis n. sp., is described herein, based on specimens
recovered from the small intestine of the leaf-nosed bat, Macrotus waterhousii, from the Biosphere Reserve ‘‘Sierra
de Huautla’’ in the state of Morelos, Mexico. The new species is included in Torrestrongylus because it features a
bursa of the type 3 – 2, a divided cephalic vesicle with an anterior half in the shape of an umbrella, and a posterior
widened half. The new species can be distinguished from the only other congener T. torrei Pérez-Vigueras, 1935 by
four key features: first, by the absence of cervical alae in both males and females; second, by the relatively longer
second half of the cephalic cap; third, by the configuration of the dorsal ray, that does not have a medial terminal
ray, and finally, by the structure of the spicules. This is the second species in the genus, previously known from bats
of the families Phyllostomidae and Molossidae in Cuba, and now in Mexico.
Key words: Nematode, Parasite, Torrestrongylus tetradorsalis, Macrotus waterhousii, Chiroptera, Mexico.
Résumé – Une nouvelle espèce de Torrestrongylus (Trichostrongylidae, Anoplostrongylinae) parasite de
Macrotus waterhousii (Chiroptera: Phyllostomidae) du Mexique central. Une nouvelle espèce de nématode,
Torrestrongylus tetradorsalis n. sp. est décrite ici, à partir de spécimens collectés dans l’intestin grêle du
chiroptère Phyllostomidae Macrotus waterhousii, de la Réserve de la Biosphère « Sierra de Huautla » dans l’État
de Morelos au Mexique. La nouvelle espèce est incluses dans Torrestrongylus car elle possède une bourse de type
3-2, une vésicule céphalique divisée avec une moitié antérieure en forme de parapluie et une moitié postérieure
élargie. La nouvelle espèce est distinguée de la seule espèce congénère T. torrei Pérez-Vigueras 1935 par quatre
caractéristiques clés ; premièrement l’absence d’ailes cervicales chez les mâles et les femelles ; deuxièmement, la
seconde moitié de la capsule céphalique relativement plus longue ; troisièmement, la configuration du rayon
dorsal, qui ne possède pas de rayon médian terminal ; finalement, par la structure des spicules. Ceci est la
deuxième espèce du genre, déjà connu des chauves-souris des familles Phyllostomidae et Molossidae à Cuba, et
maintenant au Mexique.
*Corresponding author: agustinjz@zoology.siu.edu
Juan Manuel Caspeta-Mandujano – urn:lsid:zoobank.org:author:92E35B25-5C33-4C50-A0A4-3D46153F5CDA
Jorge Luis Peralta-Rodríguez – urn:lsid:zoobank.org:author:C33D4786-24A5-4056-B6E6-58A8E4B5FABE
María Guadalupe Galindo-García – urn:lsid:zoobank.org:author:D4CD8DF1-93EA-410E-B864-87AE265FD8A5
Francisco Agustín Jiménez – urn:lsid:zoobank.org:author:C640E07E-20E8-419B-B50A-613013FD36EF
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
2
J.M. Caspeta-Mandujano et al.: Parasite 2015, 22, 29
Introduction
Species in Anoplostrongylinae Chandler, 1938 (Nematoda:
Molineidae) are known to infect xenarthrans and bats [7]. To
our knowledge, 28 species distributed in 15 genera are known
to occur in microchiropterans. The Phyllostomidae contains 57
genera [23], from which the genus Macrotus includes two species, the Californian leaf-nosed bat, M. californicus Baird, and
Waterhouse’s leaf-nosed bat, M. waterhousii Gray. The distribution of the latter is disjunct, including a continental and a
Caribbean range. The continental range includes areas of western and central Mexico and north-central Guatemala, whereas
the Caribbean range includes islands of the Bahamas, Beata,
Cayman, Cuba, Hispaniola, and Jamaica [17]. The species
has not been found in the Yucatan peninsula or in the coastal
plains of the Gulf of Mexico. Waterhouse’s leaf-nosed bats
are cave dwellers and form groups from a few to as many as
500 individuals; they forage twice during the night, 2 h after
sunset, and 2 h before sunrise. As a consequence, some consider them ‘‘gleaning’’ insectivores that capture prey such as
crickets, grasshoppers, beetles, and sphinx moths from the
ground or foliage rather than in flight [1].
During an investigation of helminth parasites of bats in
central Mexico, carried out by the research team of the Laboratorio de Parasitología de Animales Silvestres of the Universidad Autónoma del Estado de Morelos (UAEM), several
nematodes of the genus Torrestrongylus were recovered from
the small intestine of Waterhouse’s leaf-nosed bats in the State
of Morelos, Mexico. A detailed morphological analysis of this
material revealed the presence of a new species, which is
described below.
Materials and methods
From August 2012 to July 2013, 36 Waterhouse’s leafnosed bats were collected by placing a mist net across the
opening of the abandoned ‘‘Mina América’’, located in the
Biosphere Reserve ‘‘Sierra de Huautla’’, municipality of
Tlaquiltenango, Morelos, Mexico (18°270 47" N, 99°000 54" W,
1063 m). Bats were handled and killed following recommendations set forth by the American Society of Mammalogists
[16], and collected under Licencia de Colector Científico
FAUT – 0211 (National Collecting Permit 0211) from the Secretaria del Medio Ambiente y Recursos Naturales (SEMARNAT). Parasitological examinations were performed in
freshly killed bats, following the recommendations and protocols described in Gardner and Jiménez-Ruiz [8].
A thorough helminthological examination resulted in 219
nematodes of an unidentified species of Torrestrongylus that
were recovered from the small intestine of 26 Waterhouse’s
leaf-nosed bats. The worms collected were washed in saline
solution, counted, and fixed in either hot 4% formaldehyde
or 70% ethanol. Nematodes were cleared with glycerin and
temporarily mounted for morphological examination. Two
specimens were progressively dehydrated in a graded ethanol
series, dried by critical point drying using CO2, affixed to a
SEM stub, and sputter-coated with gold palladium. Specimens
were imaged on a scanning electron microscope (Hitachi
S2460 N, Tokyo, Japan). All measurements provided are in
millimeters except where otherwise indicated. Upon identification, all specimens were washed and stored in vials in 70% ethanol and submitted to the Colección Nacional de Helmintos
(CNHE) of the Universidad Nacional Autónoma de México
(Mexico City, Mexico), the Colección Parasitológica de la Universidad Autónoma del Estado de Morelos (COPAUAEM,
Cuernavaca, Mexico), the Harold W. Manter Laboratory of
Parasitology (HWML, University of Nebraska, Lincoln,
Nebraska), and the United States National Parasite Collection
(USNPC, Smithsonian Institution).
Genomic DNA was extracted, isolated, and purified from
vouchered nematodes following standard protocols [10, 15].
These aliquots were used as a template to amplify the mitochondrial gene cytochrome c oxidase subunit 1 (cox1); the
primers and thermal profile used to complete the reactions,
as well as the postamplification processing of these fragments,
are identical to those described elsewhere [10]. Published
sequences of available Anoplostrongylinae and other
strongyles were downloaded from GenBank [10], and these
include Macuahuitloides inexpectans Jiménez, PeraltaRodríguez, Caspeta-Mandujano, and Ramírez-Díaz, 2014 GenBank KF425296, and KF425297, HWML68495 and
HWML68496, and the species described herein with the
accession numbers GenBank KF425294, HWML68493 and
GenBank KF425295, HWML68494. The resulting amplicons
were aligned with ClustalW (http://www.genome.jp/tools/
clustalw/). The model of evolution HKY + G was selected using
the corrected Akaike Information Criterion as implemented in
jModelTest [12]. Parsimony and maximum likelihood were used
as optimality criteria to reconstruct the phylogeny [18], and a
Bayesian inference to estimate the posterior probabilities of
the nodes [13]. In PAUP*, the phylogeny was reconstructed by
means of a heuristic search with tree bisection reconnection
(TBR) branch swapping, 100 random additions of sequences,
and 10 trees held at each replicate. One thousand bootstrap replicates were performed using a heuristic search in PAUP*. For the
Bayesian inference, five chains were set to run for 10 million generations with resampling every 1000 iterations and a burn-in of
25% of the resulting trees. The remaining trees were used to
reconstruct the consensus.
Torrestrongylus tetradorsalis n. sp.
urn:lsid:zoobank.org:act:E25AC858-E5A6-441C-A5CA4B398D656914
Type host: Waterhouse’s leaf-nosed bat, Macrotus waterhousii Gray, 1843.
Type locality: Mexico: Morelos State: municipality of Tlaquiltenango: Mina América, Biosphere Reserve ‘‘Sierra de
Huautla’’ (18°270 47" N, 99°000 54" W).
Site of infection: Small intestine.
Prevalence and range of intensity: 72% (26 of 36 bats
examined); 1–28 nematodes.
Date of collection: From August 2012 to July 2013. Holotype, allotype, and paratypes were collected in April 2013.
Etymology: The species name, tetradorsalis, refers to the
four terminal papillae present in the dorsal ray.
J.M. Caspeta-Mandujano et al.: Parasite 2015, 22, 29
3
Figures 1–12. Torrestrongylus tetradorsalis n. sp. 1: Anterior end of male, ventral view, showing the esophagus and the relative positions of
the nerve ring and deirids; note separation of cuticular projection in anterior vesicle. 2: Anterior end of female, lateral view showing the
relative position of the nerve ring, deirid, and excretory pore; note continuous cuticular expansion in anterior vesicle. 3: Detail of the cephalic
vesicle showing the anterior half with the lateral projections forming an umbrella and the posterior half with ‘‘handles of a pitcher’’
appearance. 4: Ovejector of female showing flap in anterior lip of vulva, vagina vera, infundibulum, sphincters, and uterine branches
containing mature eggs. 5: Face view of male, showing arrangement of papillae and cuticular expansions of the anterior cephalic vesicle.
6: Posterior end of male, showing the relative position of spicules and gubernaculum. 7: Spicule. 8: Gubernaculum. 9: Egg. 10: Caudal bursa
showing bursal arrangement and bifurcation of dorsal ray. 11: Lateral view of tail of female. 12: Ventral view of tail of female.
Deposition of specimens: Holotype, allotype, and paratypes in the Instituto de Biología, UNAM, Mexico City
(CNHE9899 – 9901); USNPC106976 – 79; COPAUAEM N500; HWML68495 – 98.
Description (Figs. 1–26)
General: Small nematodes with synlophe armed with
32–33 ridges at midbody, ridges continuous, of equal size,
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J.M. Caspeta-Mandujano et al.: Parasite 2015, 22, 29
Figures 13–20. Scanning electron micrographs of Torrestrongylus tetradorsalis n. sp. 13: Subventral view of the anterior end, showing
lateral projections of cephalic vesicle; posterior part of cephalic vesicle appears ‘‘deflated’’. 14: Face view showing stoma, papillae, and
amphids. 15: Detail of the vulva featuring conspicuous flap, lateral view. 16: Ventral view of vulva showing the basis of the vulvar flap.
17: Deirid. 18: Lateral view of caudal bursa, showing projections supported by bursal rays. 19: Ventral view of tail of female, featuring anus
and terminal spines. 20: Subterminal view of tail of female, showing all three cuticular projections and terminal filament.
Table 1. Comparative measurements of key diagnostic traits of Torrestrongylus torrei and T. tetradorsalis. All measurements are in
millimeters.
Measurements
Length of body
Maximum width
Cephalic vesicle
Cervical alae
Nerve ring
Excretory pore
Deirids
Length of esophagus
Vulva to anterior end
Eggs
Tail
Length of terminal spines
Dorsal ray
Torrestrongylus torrei
$
6.2
Length
Width
Length
Width
#
5.6
0.056
0.096
0.096
0.026–0.027
0.160
Length
Width
4.8
0.067
0.027
–
Torrestrongylus tetradorsalis
$
7.6–9.2
0.17–0.23
0.060–0.088
0.130–0.160
0.050–0.090
0.010–0.020
0.260–0.350
0.310–0.400
0.330–0.440
0.570–0.680
5.31–6.45
0.081–0.0110
0.070–0.080
0.060–0.093
#
6.9–8.4
0.160–0.200
0.050–0.080
0.0.110–0.130
0.060–0.080
0.010–0.020
0.210–0.280
0.310–0.360
0.350–0.410
0.430–0.560
–
–
–
0.040
Spicules
supported by strut, barely emerging from cuticle (Figs. 14–16).
Cephalic vesicle divided into two, anterior half in the form of
an umbrella with two lateral cuticular folds, followed
by a posterior half, regularly expanded without striations
Double bifurcation
plus unpaired end
0.147
Trifurcated
Double bifurcation
–
–
0.120–0.150
Bifurcated
(Figs. 1–3, 5, 13, 14). Stoma small, triangular, with esophagorhabdion forming an inward projection or dorsal tooth (Figs. 5
and 14). Stoma flanked by amphids and endowed with four
pairs of cephalic papillae; (Figs. 5, 14); small deirids simple
J.M. Caspeta-Mandujano et al.: Parasite 2015, 22, 29
5
Figures 21–26. Cross section of Torrestrongylus tetradorsalis n. sp. Male at level of 21, nerve ring, 22 midbody and 23, posterior end of
intestine. Female at level of 24, nerve ring, 25 midbody and 26, posterior end of intestine.
and flattened (Fig. 17), located at midlength of esophagus
(Fig. 1). Tail of male ending in subsymmetrical bursa (Figs. 6,
10, 18) and that of female conical, with three conspicuous terminal cuticular processes or tubercles, and a fine terminal
thread (Figs. 11, 12, 19, 20).
Male (based on 10 specimens; ranges, see Table 1): Body
length 6.89–8.38 mm, width at midbody 0.16–0.20 mm. Anterior half of cephalic vesicle 0.05–0.08 mm long and 0.11–
0.13 mm wide; posterior half, 0.06–0.08 mm long and
0.01–0.02 mm wide. Nerve ring, excretory pore, and deirids
0.21–0.28 mm, 0.31–0.36 mm and 0.35–0.41 mm, respectively, from anterior end. Length of esophagus, 0.43–0.56 mm
long and 0.068–0.08 mm at its widest region, close to junction
with intestine. Synlophe starts immediately behind cephalic
vesicle and ends near bursa; 32 ridges at level of nerve ring,
32 at midbody, and 28 near posterior end (Figs. 21–23). Subsymmetric copulatory bursa features a 3 – 2 arrangement and
a prominent dorsal ray (Fig. 10); rays 2 to 4 independent, separated at base, and slightly directed anteriad; rays 5 and 6 connected at base, extend laterally with tips that emerge from
margin; dorsal ray bifurcates at basis and features a prominent
ray 8; rays 9 and 10, subterminal (Fig. 10). Cuneiform spicules,
of equal shape and size, 0.12–0.15 mm long, with lamina
divided into its middle portion ending at two terminal points,
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J.M. Caspeta-Mandujano et al.: Parasite 2015, 22, 29
thick lamina dorsally developed, thin vellum extends into proximal part (Figs. 6, 7). Gubernaculum 0.030–0.037 mm long
(Fig. 8). Ranges for these values are included in Table 1.
Female (based on 10 specimens; ranges, see Table 1):
Body length 7.45–9.23 mm and width 0.17–0.23 mm. Anterior
part of the cephalic vesicle 0.06–0.08 mm long and 0.13–
0.16 mm wide, posterior part 0.05–0.09 mm long and
0.01–0.02 mm wide. Nerve ring, excretory pore, and
deirids 0.26–0.35 mm, 0.31–0.40 mm, and 0.33–0.44 mm,
respectively, from the anterior end. Length of esophagus,
0.57–0.68 mm long and 0.08–0.10 mm wide. Synlophe ridges
start immediately behind cephalic vesicle and at basis of terminus and caudal tubercles; 32 ridges at level of nerve ring, 33 at
midbody, and 21 at level of posterior end of intestine,
close to anus. Amphidelphic. Postequatorial vulva located
5.31–6.45 mm from anterior end; a small extension in the form
of flap present in anterior margin of vulva (Figs. 4, 15, 16).
Vulva connected to short vagina, leads to vestibule; sphincters
short, infundibula elongated, leading to uteri (Fig. 4). Subterminal anus 0.06–0.093 mm from posterior end (Fig. 19). Tail
conical. Posterior end includes three tubercles or cuticular processes of various sizes; the largest is dorsal, two of ventral
medium-sized, and finally, a thin ventral projection or
terminus (Figs. 12, 19, 20). Eggs 0.081–0.11 mm long and
0.07–0.08 mm wide; embryonated (Fig. 9). Ranges for these
values are included in Table 1.
Figure 27. Relative position of Torrestrongylus tetradorsalis and
Macuahuitloides inexpectans as reconstructed in the ML tree used
to estimate the posterior probabilities of the branches. The numbers
to the right of each node indicate bootstrap support for parsimony
and maximum likelihood as optimality criteria and the posterior
probability of the branch. Syngamus trachea belongs to Strongyloidea. The tree and data matrix can be retrieved from
http://opensiuc.lib.siu.edu/zool_data/4/
Phylogeny
According to sequences of available taxa, the genetic difference between the two specimens of T. tetradorsalis is
0.5%; the posterior credibility of the clade T. tetradorsalis + M. inexpectans is 1. This clade has moderate bootstrap
support using maximum likelihood as an optimality criterion,
but no support using parsimony (Fig. 27).
Remarks
The configuration of the cephalic vesicle acts as a reliable
character to help discriminate genera in Anoplostrongylinae.
Torrestrongylus torrei Pérez-Vigueras, 1935 is characterized
by the presence of a cephalic vesicle divided into two parts;
the anterior has the shape of an umbrella and the posterior is
slightly widened, and it has the appearance of ‘‘handles of a
pitcher’’ [11]. Based on the structure of the caudal bursa and
synlophe, as well as the structure of the cephalic vesicle, the
species described herein has been placed in the genus Torrestrongylus. The configuration of the cephalic vesicle is markedly different from the homologous structure in other genera
in the subfamily, which in most cases possess ornate and simple cephalic vesicles.
The specimens described herein show remarkable similarities to T. torrei, especially in the configuration of the cephalic
vesicle, the presence of a vulvar flap, and the 3 – 2 bursal
arrangement. In addition, the specimens used in the present
description were collected from the same species of bat, Macrotus waterhousii. Torrestrongylus tetradorsalis, however, can
be discriminated from T. torrei by the absence of cervical alae
in both sexes, and by the absence of a terminal unpaired end in
the dorsal ray, as seen in T. torrei (Plate 7 in [11]). Other differences between these two species include: the relatively
shorter size of the posterior half of the cephalic cap of T. torrei;
the general shape and distal tip of spicules (trifurcated in
T. torrei vs. bifurcated in T. tetradorsalis); the larger body size
of specimens of T. tetradorsalis; the location of the nerve ring
relative to the anterior end, and the larger size of eggs in
T. tetradorsalis (Table 1).
Discussion
Other members of Anoplostrongylinae that show similarities to T. tetradorsalis include Anoplostrongylus paradoxus
(Travassos, 1918) Boulenger, 1926 and some species of Tricholeiperia Travassos, 1935 [20]. However, T. tetradorsalis cannot be placed in these genera because of the configuration of
the cephalic vesicle, caudal bursa, and structure of the synlophe. On the one hand, when compared with A. paradoxus,
T. tetradorsalis shows a prominent dorsal ray, more complex
spicules, and a relatively equal length of branches of posterior
and anterior vestibules. On the other hand, species in Tricholeiperia feature an umbrella-shaped cephalic vesicle, and only
Tricholeiperia poeyi Baruš and Valle, 1967 features a divided
cephalic vesicle [2]. The cephalic vesicle of this species, however, appears to be formed by an anterior dome-shaped half
and a widened posterior half. This configuration is markedly
different to the cephalic vesicle present in the other species
of the genus [5, 9, 19, 21], which feature a vesicle barely separated from the rest of the body [9, 19], or conspicuous expansions divided into two lateral halves [21].
The main difference between species of Tricholeiperia and
Torrestrongylus is the configuration of the cephalic cap. In the
present study, one of the specimens (GenBank Accession
Numbers KF425295, HWML68494) features a poorly
developed posterior half of the cephalic vesicle, which suggests
it belongs to Tricholeiperia. However, the genetic
divergence with T. tetradorsalis (GenBank Accession Numbers
KF425294, HWML68493) is 0.5%, which suggests they
J.M. Caspeta-Mandujano et al.: Parasite 2015, 22, 29
belong to the same species. These results show the urgent need
for an extensive phylogenetic analysis including other Anoplostrongylinae featuring the umbrella cephalic vesicle, that
may allow the evaluation and proposition of reliable characters
used in their diagnosis.
Currently, the list of parasitic helminths of M. waterhousii
includes five species, two digenetic trematodes including
Limatulum aberrans Caballero and Bravo, 1950 and
L. oklahomense Macy, 1931, and three species of nematodes,
Aonchotheca viguerasi (Freitas and Lent, 1937), Litomosoides
leonilavazquezae Caballero, 1939 and Torrestrongylus torrei
[3, 4, 11]. The only species of parasite recorded on M. californicus is the nematode Glyptostrongylus collaris [22]; however,
this is considered a nomen nudum since no description is associated with the name. Interestingly, the Waterhouse’s leafnosed bats have a disjunct distribution, with a continental cluster localized in western Mexico and north-central Guatemala,
and a Caribbean cluster that spans several islands in the
Caribbean. Torrestrongylus torrei was described from bats in
La Habana, Cuba, and T. tetradorsalis was collected in bats
from Mexico; therefore, these two localities represent both
clusters. Torrestrongylus torrei is known to infect phyllostomid
and molossid bats [11, 14], which suggests this species may be
a generalist. The results of our present investigations suggest
that in central Mexico, T. tetradorsalis is restricted to the
Waterhouse’s leaf-nosed bat. However, the expansion of the
survey to other areas in the distribution of this species of chiropteran, as well as the inclusion of additional species from
other families, will help in establishing the host spectrum for
T. tetradorsalis. Additional specimens from an unidentified
species of Torrestrongylus have been reported in two species
of phyllostomids in Colombia (Micronycteris schmidtorum
and Sturnira bidens) [6]. An examination of those specimens
should allow an evaluation of the variation of the characters
used in the diagnosis of species in this genus, and in uncovering the diversity of Anoplostrongylinae in Neotropical bats.
Acknowledgements. The authors wish to thank Berenit Mendoza
Garfias, UNAM, and Alejandro de Jesús Medrano Silva, UAEM,
for technical assistance with SEM. We thank Dr. Gabor Racz and
Dr. Scott Gardner, HWML; Luis García Prieto, CNHE, and
Dr. Anna Phillips, Smithsonian Institution, for accepting the donation of specimens and providing collection numbers. Open Access
made available by the COPE program of SIU. This study was supported by a research grant from the Programa de Mejoramiento del
Profesorado (PROMEP-UAEMOR/07/203) to J.M.C.M. and J.A.G.
and by the Programa Integral de Fortalecimiento Institucional
(PIFI-UAEM 2010–2011). J.L.P.R. was supported by a scholarship
from CONACYT (241472).
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Cite this article as: Caspeta-Mandujano JM, Peralta-Rodríguez JL, Galindo-García MG & Jiménez FA: A new species of Torrestrongylus
(Trichostrongylidae, Anoplostrongylinae) from Macrotus waterhousii (Chiroptera: Phyllostomidae) in Central Mexico. Parasite, 2015, 22,
29.
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