Habitat use and reproduction of mammals from Tlaxmalac, at Balsas River basin,

Guerrero, Mexico
Author(s): José Alberto Almazán-Catalán, Cornelio Sánchez-Hernández, María de Lourdes Romero-
Almaraz, Leobardo Sánchez-Vázquez, and Sara Beatriz González-Pérez
Source: The Southwestern Naturalist, 60(1):36-44.
Published By: Southwestern Association of Naturalists
DOI: http://dx.doi.org/10.1894/JKF-44.1
URL: http://www.bioone.org/doi/full/10.1894/JKF-44.1

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THE SOUTHWESTERN NATURALIST 60(1): 36–44 MARCH 2015

HABITAT USE AND REPRODUCTION OF MAMMALS FROM

TLAXMALAC, AT BALSAS RIVER BASIN, GUERRERO, MEXICO

JOSÉ ALBERTO ALMAZÁN-CATALÁN, CORNELIO SÁNCHEZ-HERNÁNDEZ,* MARÍA DE LOURDES ROMERO-ALMARAZ,

LEOBARDO SÁNCHEZ-VÁZQUEZ, AND SARA BEATRIZ GONZÁLEZ-PÉREZ

Departamento de Zoologı́a, Instituto de Biologı́a, Universidad Nacional Autónoma de México, A.P. 70-153,
Coyoacán, México, D.F. 04510, México (JAA-C, CS-H, LS-V, SBG-P)
Escuinapa No. 92 bis. Col. Pedregal de Santo Domingo, C.P. 04369, México, D.F., México (MLR-A)
*Correspondent: cornelio@unam.mx

ABSTRACT—Mammals from the state of Guerrero, Mexico, are poorly studied, particularly those from the
Balsas River basin, despite the importance of this area as a biogeographical boundary for flora and fauna. To
contribute to the knowledge of mammals in this state, we compiled information on geographical distribution,
habitat, and reproduction for 41 mammal species from Tlaxmalac in the Balsas River basin in northeastern
Guerrero. These represented 26.9% of mammal species known from Guerrero and 8.6% of mammal species
from Mexico. Eight species were endemic, comprising 19.5% of species surveyed. Mammals from Tlaxmalac
had eight different feeding habits. People from the community use 14 species of mammals as part of their diet,
as pets, for medicinal purposes, and other uses. Overall, Tlaxmalac supports a great diversity of small and
medium-sized mammals.

RESUMEN—Los mamı́feros del estado de Guerrero, México, están pobremente estudiados, en particular los
de la cuenca del rı́o Balsas, no obstante la importancia del área por actuar como lı́mite biogeográfico para la
flora y fauna. Para contribuir al conocimiento de la mastofauna del estado, se compiló información sobre la
distribución geográfica, hábitat y reproducción de 41 especies de mamı́feros de Tlaxmalac, en la cuenca del
rı́o Balsas al noreste de Guerrero. Estas representaron 26.9% de las especies de mamı́feros conocidos para
Guerrero y 8.6% de las especies de mamı́feros de México. Ocho especies fueron endémicas, lo que constituye
19.5% de las especies registradas. Los mamı́feros de Tlaxmalac tuvieron ocho hábitos alimentarios diferentes.
Los habitantes utilizan 14 especies de mamı́feros como parte de su dieta, como mascotas, para fines
medicinales y otros usos. En general, Tlaxmalac mantiene una fauna diversa de mamı́feros de talla pequeña y
mediana.

Current biological inventories that allow us to define
the distribution and natural history of mammals are still
incomplete. This lack of information prevents us from
understanding the evolutionary processes of many re-
gions. A total of 152 species of mammals is reported from
Guerrero, Mexico (Ramı́rez-Pulido et al., 2000; Cervantes
et al., 2004; Almazán-Catalán et al., 2005; Romo-Vázquez
et al., 2005; Tejedor, 2005; Carraway, 2007; Nolasco et al.,
2007; Ruiz-Gutiérrez et al., 2011). The Balsas River basin
is formed by two physiographic provinces, the Trans-
Volcanic Belt and the Sierra Madre del Sur. The genesis
and evolution of these landforms give this basin a wide
variety of physiographic, geological, and climatic charac-
teristics that impose biogeographic limits for much of the
flora and fauna of Mexico. This structural framework
created, in geological time, many ecological islands and
peninsulas in its peaks, slopes, canyons, and isolated
valleys that made the basin a remarkable secondary center
of evolutionary radiation of species (Toledo, 2003).
Despite the importance of the area, there is only one
report on the natural history of bats and rodents from the
coastal region of the Balsas River (Álvarez, 1968). Some
mammals of this region have only been recorded
occasionally by indirect ways (e.g., in pellets of barn owl
Tyto alba; Ramı́rez-Pulido and Sánchez-Hernández, 1972).
As a contribution to the knowledge of mammals from the
upper Balsas River basin, our objectives were to compile a
comprehensive list of mammals from Tlaxmalac, a
community within this area, with information regarding
natural history and habitats, and to compare the richness
found in this area with regions nearby, to understand the
importance of the area.
MATERIALS AND METHODS—Study Area—Tlaxmalac is in the
municipality of Huitzuco de Los Figueroa in the state of
Guerrero. It is located at 18821 0 38 00 N, 99824 0 50 00 W and has an
area of 40 km2. The mean elevation is 911 m with some steep
hills up to 1,600 m (INEGI, 1998). The weather is warm and
semihumid. Average annual temperature is 25.58C and average
annual rainfall is 1,417 mm, with most rain occurring from June
March 2015 Almazán-Catalán et al.—Habitat use and reproduction of mammals from Guerrero, Mexico
to October. The dominant habitat type is tropical deciduous
forest (73%), which covers slopes from 900 to 1,600 m, and is
represented by Jacaratia mexicana (bonete), Amphipterygium
adstringens (cuachalalate), Ipomoea arborescens (cazahuate), and
Bursera copallifera (copal). Oak forest (8%) occurs on hills from
1,200 to above 1,600 m, and is represented by Quercus candicans
(white oak) and Q. glaucoides (lacey oak). The remainder of the
study area (19%) consists of second growth vegetation domi-
nated by plants such as Acacia cymbispina (acacia), A. farnesiana
(huizache, acacia), Haematoxylum brasiletto (Brazilwood), Pith-
ecellobium dulce (guamuchil), and Alvaradoa amorphoides (guachi-
pil; Jiménez Salmerón and Garcı́a, 2002), in addition to field
crops like tamarind (Tamarindus indica), peanut (Arachis
hypogaea), mango (Mangifera indica), and corn (Zea mays).
Mammal Surveys—Fieldwork took place from May 1999
through May 2007 in 22 sessions of 1 to 4 days each, for a total
of 53 sampling days. We surveyed 28 locations representative of all
types of vegetation (Appendix). Each night we set between four
and eight mist nets (6, 9, and 12 m) to capture bats from 1900h to
2300h, checking them approximately every 30 min. For terrestrial
small mammals, we set Sherman traps baited with oats and snap
traps baited with oats mixed with peanut butter, or oats with
bananas. We set small-mammal traps in the afternoon and
checked them early the next morning. Traps were set daily in
transects with intertrap distance of 10 m. Survey effort was 1,500
trap-nights using Sherman traps (3 · 3.5 · 9 in) for small to
medium-size mammals, and 900 trap-nights using snap traps (7 ·
3 1/4 in) for small mammals.
For medium- and large-size mammals, residents provided us
with skins, parts of skulls, or skeletons, whereas others were
found during our fieldwork. We preserved skin and whole
skeleton or skin and skull only. We collected tissue samples
(muscle, liver, and kidney) and preserved them in 96% ethanol.
Voucher specimens were deposited at Colección Nacional de
Mamı́feros, Instituto de Biologı́a, Universidad Nacional Autó-
noma de México; some specimens are still in the cataloguing
process. Some rodents and bats were released at the site of their
capture, if we had already collected voucher specimens. We
recorded sex, reproductive status, and habitat type for animals
that were released.
We followed taxonomy according to Wilson and Reeder
(2005), with three exceptions: Dermanura as separate from
Artibeus (Owen, 1987), Liomys as part of Heteromys (Hafner et al.,
2007), and Spermophilus as part of Otospermophilus (Helgen et al.,
2009). Subspecific designations were used according to Ramı́rez-
Pulido et al. (2005).
We gave four designations according to the traditional uses of
mammals in the area (food, pet, fur, and medicinal). Such
information was obtained from farmers, healers, and house-
wives. Conservation status of species was stated according to
Norma Oficial Mexicana NOM-059-SEMARNAT-2010 (SEMAR-
NAT, 2010), and according to International Union for Conser-
vation of Nature (IUCN, 2014), through the Red List of
Threatened Species.
Nonparametric methods were used to estimate expected
species richness using two incidence-based estimators (Chao 2
and Bootstrap) with EstimateS 8.2 software (Colwell, 2009).
These analyses use sampling numbers of a given area to estimate
the number of species expected to inhabit that region. Chao 2 is
a rigorous and little-biased estimator for small populations, and
Bootstrap makes intensive calculations on the basis of a large
37
number of repetitive calculations for estimating the sampling
distribution of population.
We constructed a matrix of presence and absence of species
to compare the similarity of mammal species from Tlaxmalac
with the nearest regions where mammals have been studied, and
created a dendrogram of hierarchical clustering on the basis of
Jaccard coefficient by unweighted pair-group method using
arithmetic averages. This coefficient standardizes the richness
information of every area minimizing the sampling bias
(sampling effort and sample size) using a binomial datum
(presence 1, absence 0) of the registered species. The
dendrogram graphically represents the results, grouping areas
more similar on the basis of their species composition
(Magurran, 1988). We processed the data using the multivariate
statistical package MVSP 3.13r (Kovach, 2009). The areas with
which we compared our richness were Costa Grande (Ramı́rez-
Pulido et al., 1977), Sierra de Taxco (León-Paniagua and Romo-
Vázquez, 1993), Omiltemi in the state of Guerrero (Jiménez-
Almaraz et al., 1993), and Sierra de Huautla in the state of
Morelos (Sánchez-Hernández and Romero-Almaraz, 1995).
SPECIES ACCOUNTS—Didelphis virginiana californica—Lo-
cality 13b. We captured one male with scrotal testes 20 ·
14 mm in October 2000 in a rocky ravine, surrounded by
corn and peanut fields and guamuchil trees; that day we
observed nine more individuals during a ca. 1-h hike close
to open areas in the forest and near bodies of water.
Tlacuatzin canescens—Locality 10, June 2001. We
examined a male with scrotal testes of 11 · 8 mm that
was found dead on the litter of pristine tropical semi-
deciduous forest. This species was frequently seen among
the corn fields and in hanging abandoned Icterus nests.
Dasypus novemcinctus davisi—Locality 7, April 2003.
Residents donated the skin of a male that was captured in
tropical deciduous forest. We observe this species to be
common in corn and peanut fields and occasionally
residents fed them.
Sciurus aureogaster nigrescens—Locality 8a, December
2000. We captured a reproductively inactive adult female
in oak forest. This species was occasionally seen in yards
or in tamarind trees.
Otospermophilus variegatus variegatus—Locality 5, April
2001. We examined a postlactating female (Colección
Nacional de Mamı́feros, Instituto de Biologı́a, Universi-
dad Nacional Autónoma de México 42516). The speci-
men was feeding on guamuchil fruits and was in a flat,
rocky terrain planted with peanuts and corn.
Heteromys irroratus torridus—Localities 2a, 3a, 4, 10a,
10c, 12a, 13, 14b, 15, 16a, 19a, 20, 21, 23, 25, 26, 26a, and
27. We examined 19 females, 22 males, and 9 newborns
from 1999 to 2001 and from 2007. The specimens were
caught in corn- and peanut fields, along roads, dried
streambeds, in grassland, and among roots of trees. Of
the 19 females, 2 were young (July, November), 3 were
subadults (2 in February, 1 in July), and 14 were adults. Of
the adults, eight were reproductively inactive (four in
May, one in June, three in July). A female taken in July
had two embryos (1R, 20 · 14 mm; 1L, 10 · 13 mm), and
38 The Southwestern Naturalist
one taken in October had four embryos (2R, 2L, 21 · 14
mm). Three females caught in September and October
were lactating, and females caught in September had
three and six newborns that were born in the trap. One
female from October had a vaginal plug. Of the 22 males,
3 were young (1 in February, 2 in November), 1 was
subadult (February), and 18 were adults. Of the adults,
eight had scrotal testes, including two in May (7 · 4 mm,
13 · 10 mm), five in July (19 · 12 mm to 20 · 14 mm),
and one in August (20 · 10 mm); five had inguinal testes,
two had abdominal testes, and we did not record data for
three of them.
Baiomys musculus pallidus—Localities 4, 11a, 12a, 13,
14b, 20, 21, and 26a. We examined six females and seven
males from 1999 to 2001 and from 2007. All females were
reproductively inactive, including five adults: one in
January, two in February, one in May, one in August;
and one subadult in November. Seven males were
collected including five adults: one in February, four in
May; and one juvenile and one subadult in November.
One male captured in November and one in May had
scrotal testes (4 · 2 mm and 7 · 3 mm, respectively). The
remaining individuals had abdominal or inguinal testes.
We trapped them along the streambeds, on the limits of
corn- and peanut fields, in cattle ranches, and among
thorny vegetation and secondary vegetation in tropical
deciduous forest.
Hodomys alleni elatturus—Localities 19 and 26a. We
examined four females and one male. One juvenile of
each sex was captured in February 2001. A female
collected in August had two embryos (2L, 9 · 7 mm).
Two females caught in February 2001were reproductively
inactive. The specimens were captured in traps set in rock
slopes near hollows, in tropical deciduous forest, and
thorny vegetation.
Osgoodomys banderanus vicinior—Localities 2a, 3a, 4,
10b, 10c, 11, 11a, 12, 12a, 13, 14, 14a, 15, 16a, 19, 19a,
19b, 19c, 19d, 20, 21, 26, 26a, and 27. We examined 42
females and 55 males from 1999 to 2001 and from 2007.
We trapped all of the specimens among tree roots or
rocks, along dry streambeds, and in corn- and peanut
fields, with surrounding vegetation of tropical deciduous
forest and secondary vegetation.
One of the females was young (February) and four were
subadults (February, one; July, two; August, one). Four
adult females captured in January were pregnant and one
was reproductively inactive; three had two embryos (2R, 7 ·
5 mm; 2L, 15 ·14 mm; 1R, 1L, 2.4 · 1.5 mm) and one had
three embryos (3R, 25 · 17). Of six females caught in
February, two had two embryos (2L, 1 · 1 mm; 2R, 3 · 2
mm), one was lactating, and three were reproductively
inactive. The female caught in March was reproductively
inactive. Of the nine females caught in May, one had two
embryos (2L, 13 · 11 mm), six were lactating, one was
lactating and had three embryos (16 · 12 mm), and one
was inactive. Of three females caught in July, one had two
vol. 60, no. 1
embryos (1R, 1L; 23 · 14 mm), one had three embryos
(3L, 17 · 13 mm), and one was inactive. We captured four
females in August, including one with three embryos (1R,
2L, 6 · 5 mm), one with an early implantation, and two that
were reproductively inactive. Six females from September
were pregnant: two had two embryos (2R, length 14 mm;
1R, 1L, 30 · 13 mm), three had three embryos (3L, 15 · 8
mm; 3R, length 25 mm; 2R, 1L, length 18 mm), and one
had four embryos (4L, 20 · 15 mm). We captured two
females in October including one with three embryos (3L,
9 · 8 mm) and one that was reproductively inactive. In
November, one female was lactating.
Of the males, seven were young (January, two; May,
one; September, three; October, one) and four were
subadults (March, three; August, one). Of the adults, 35
had scrotal testes, including 3 from January (15 · 10 mm
to 20 · 10 mm), 4 from February (16 · 11 mm to 21 · 8
mm), 10 from May (6 · 3 mm to 21 · 10 mm), 1 from
June (17 · 10 mm), 8 from July (17 · 9 mm to 20 · 11
mm), 4 from August (14 · 8 mm to 26 · 10 mm), and 5
from September (7 · 5 mm to 20 · 10 mm).
Peromyscus melanophrys melanophrys—Locality 27. We
captured one pregnant female with four embryos (2R,
2L, 7 · 7 mm) in July 2000. The specimen was caught in a
trap placed among the roots of a guamuchil tree near a
cornfield.
Oryzomys couesi albiventer—Locality 4a, February 2001.
We captured one male with inguinal testes 6 · 4 mm
among the roots of a guamuchil tree near a corn- and
peanut field.
Sigmodon mascotensis—Locality 24, December 2000. We
found one male specimen dead on the road. We
commonly observed this species in grasslands, scrub,
and corn and peanut crops.
Sylvilagus cunicularius cunicularius—Localities 13 and
16b. We examined two females and two males. Both females
had embryos of asymmetric size: one caught in August 2000
had five embryos (4R, 25 · 24 mm; 1L, 20 · 20 mm), and
one caught in June 2001 had three embryos (1R, 27 · 18
mm; 2L, 37 · 30 mm). The males were caught in October
2000 and June 2001 and had scrotal testes 24 · 8 mm and
20 · 10 mm, respectively. The specimens were trapped in
peanut fields, scrub vegetation, and fragmented tropical
deciduous forest. In June 2000, 15 individuals were
observed during the night. People from the area use this
species to supplement their diet.
Balantiopteryx plicata plicata—Localities 4, 6, 12b, and
18. We examined 10 females and 12 males. Of the
females, one caught in June 2001 was a newborn, and one
caught in August 2000 was a juvenile. One female from
August 2000 and three from March 2001 were reproduc-
tively inactive, two from June 2001 and one from August
2000 were lactating, and one from June 2001 had an
embryo 22 · 14 mm. Three males caught in June were
newborns, two from August had inguinal testes (3 · 2
mm), and we did not record reproductive information
March 2015 Almazán-Catalán et al.—Habitat use and reproduction of mammals from Guerrero, Mexico
from the others. The specimens were captured in rock
hollows, inside the bathrooms of a school, and in a small
cave where there was a colony of about 100 individuals.
Surrounding vegetation was tropical deciduous forest.
Desmodus rotundus murinus—Localities 2b and 27. We
examined two adult females and three adult males
captured in 2001, one male in June and the rest in
December. Both females were reproductively inactive.
Males had scrotal testes of 8 · 6 mm (in June) and 7 · 4
mm and 7 · 5 mm (in December). They were caught in a
mist net set over a pond in tropical deciduous forest.
Choeronycteris mexicana—Locality 2b, December 2001.
We examined one adult female caught near a stream with
surrounding tropical deciduous forest.
Glossophaga leachii—Localities 2b and 11a. In 2001, we
captured one reproductively inactive female in December,
and one male with scrotal testes (6 · 5 mm) in May. They
were caught in tropical deciduous forest.
Glossophaga morenoi mexicana—Localities 2b, 11a, and
12b. We examined five females (one in March, two in May,
and two in December) and one male (May) in 2001.
Females were reproductively inactive, and the male had
scrotal testes (6 · 4 mm). We captured them over a body
of water in tropical deciduous forest.
Glossophaga soricina handleyi—Localities 7, 17, and 27.
We examined two females and one male. They were
captured in March, May, and June 2001. The male had
scrotal testes 4 · 3 mm in March; one female from June
was reproductively inactive and we did not record
reproductive data from the other. We captured them
among mango and tamarind trees and over a body of
water in tropical deciduous forest.
Macrotus waterhousii mexicanus—Localities 17 and 27.
We captured one female (December) and four males
(one in May, one in June, and two in December) in 2001.
The female was reproductively inactive, whereas two
males had scrotal testes, one in June (6 · 4 mm) and
one in December (5 · 3 mm). The specimens were
caught among mango and tamarind trees and near ponds
in tropical deciduous forest.
Sturnira lilium parvidens—Localities 2b, 4a, 11a, 12b,
and 27. We examined two females and three males. They
were caught in March (one male with scrotal testes, 5 · 3
mm), May (one young male), June (one lactating
female), and December (one male with inguinal testes;
and one young female) from 2001. The specimens were
caught over a body of water in tropical deciduous forest.
Artibeus hirsutus—Localities 12b and 27. We examined
two females and two males. One young female was
captured in March, and one from December had an
embryo (5 · 4 mm). One male from December had
scrotal testes (8 · 7 mm). The specimens were caught
over a stream in tropical deciduous forest.
Artibeus intermedius intermedius—Localities 2b, 4b, 11a,
17, and 27. We examined three females and nine males.
They were caught in May (n = 2), June (n = 8), and
39
December (n = 2). Of the females, one from June was
receptive, and two were pregnant (June, 6 · 4 mm;
December, 8 · 8 mm). Eight males had scrotal testes
including two from May (6 · 4 mm, 9 · 6 mm), five from
June (5 · 4 mm to 13 · 8 mm), and one from December
(7 · 5 mm). The specimens were caught among mango
and tamarind trees, near a stream, or in tropical
deciduous forest.
Artibeus jamaicensis triomylus—Localities 4b, 11a, 12b,
and 27. We examined four females and four males. They
were caught in March (n = 1), May (n = 1), June (n = 3),
and December (n = 3) 2001. One female from June was
receptive and lactating, and one was lactating, whereas
one in December had an embryo (6 · 5 mm), and one
was reproductively inactive. Males had scrotal testes in
March (7 · 5 mm), May (8 · 5 mm), June (13 · 9 mm),
and December (7 · 4 mm). The specimens were caught
over a body of water in tropical deciduous forest.
Artibeus lituratus palmarum—Localities 12b and 27. We
examined three males with scrotal testes captured in
2001, including two in March (8 · 5 mm, 8 · 6 mm) and
one in December (7 · 5 mm). All were collected over a
water body in tropical deciduous forest.
Dermanura tolteca hespera—Locality 2b. We examined
one lactating female from December 2001. It was caught
near a pond in tropical deciduous forest.
Pteronotus parnellii mexicanus—Localities 2b, 7, 12b, and
27. In 2001, we captured seven females and one adult male,
including three in March, one in June, and four in
December. Of the females, two were pregnant including
one captured in March (1L, 17 · 11 mm) and another in
December (2 · 2 mm); one from December was lactating,
and four were reproductively inactive. The male had scrotal
testes (4 · 3 mm) and was captured in June. We trapped
them near bodies of water in tropical deciduous forest.
Nyctinomops femorosaccus—Locality 12b. We examined
one reproductively inactive adult female (Colección
Nacional de Mamı́feros, Instituto de Biologı́a, Universi-
dad Nacional Autónoma de México, 42514) captured in
March 2001 over a body of water in a rocky canyon in
tropical deciduous forest.
Nyctinomops macrotis—Locality 12b. We examined one
adult male (Colección Nacional de Mamı́feros, Instituto
de Biologı́a, Universidad Nacional Autónoma de México,
42515) with inguinal testes of 2 · 1 mm that was captured
in March 2001 over a body of water near a rocky canyon in
tropical deciduous forest.
Myotis velifer velifer—Locality 4b. We examined one
lactating female captured in June 2001 near a pond in
tropical deciduous forest.
Leopardus wiedii glaucula—Locality 9, September 2002.
Local people gave us the carcass of a female that was
captured in a rocky ravine when it was chasing a rabbit
near a pond. There were remains of a lizard (Sceloporus) in
its stomach. In February 1999, we examined the skin of a
female that had been captured in the area. Farmers
40 The Southwestern Naturalist
mentioned that this species is often found in the upper
parts of the hills near the streams.
Canis latrans cagottis—Locality 8, December 2000. A
skin of a male that was caught in tropical deciduous forest
was donated to us. We frequently observed this species in
agricultural land.
Urocyon cinereoargenteus nigrirostris—Localities 13a, 14c,
and 19. We examined a female and a male caught in
February 2001. The female was reproductively inactive
and the male had scrotal testes (20 · 13 mm); they were
captured in a ravine among corn- and peanut fields. In
April 2003, we collected the remains of two specimens,
one in a corn- and watermelon field and the other in
tropical deciduous forest.
Conepatus leuconotus leuconotus—Localities 13a and 22.
We examined two males captured in October 2000 and
February 2001, both with scrotal testes (17 · 17 mm, 18 ·
20 mm). The specimens were caught in a peanut field
and near a pond with thorny vegetation. Local people
capture this species to use its fat to cure skin infections.
Mephitis macroura macroura—Localities 13a and 28.
Local people gave us two reproductively inactive females
that were caught in February 2001, close to a river near
corn- and peanut fields. Individuals of this species are
often observed dead on the road.
Spilogale putorius tropicalis—Locality 16. In June 2001,
local people gave us a skin, skull, and skeleton of a
reproductively inactive female that was captured in
tropical deciduous forest. This species is commonly found
among the stones of walls (tecorrales) and agricultural
land or courtyards.
Bassariscus astutus bolei—Locality 3, December 2000.
The local people gave us the skin of a reproductively
inactive female that was captured in a ravine in tropical
deciduous forest.
Nasua narica molaris—Locality 1, November 2000. The
local people captured and gave us the skin and skull of
two females. One of them was lactating and the other one
was reproductively inactive. The specimens came from a
group of about 14 individuals, most of which were
juveniles. They were captured on rocky ground of tropical
deciduous forest. They are an important source of food
for the people in this community.
Procyon lotor hernandezii—Localities 2 and 3. In Novem-
ber 2000 local people gave us the skin of a young female
that was found with a group of five or six individuals. In
December of the same year they gave us the skin and the
skull of a male with scrotal testes (24 · 18 mm). The
specimens were caught at the edge of a stream in tropical
deciduous forest.
Pecari tajacu humeralis—Locality 19. We examined, but
did not retain, the skins of two specimens captured in
April 2003 in tropical deciduous forest. According to the
farmers, javelinas are rare in the area. However, they were
common at the top of the hill El Jumil, in the
neighboring community of Tuxpan, where natural
vol. 60, no. 1
vegetation is better preserved. They represent a signifi-
cant source of food for the inhabitants.
Odocoileus virginianus mexicanus—Locality 19, April
2003. Local people gave us the skin of a young male that
was caught in tropical deciduous forest. Deer represent a
significant source of food for the people of the region
and are often seen in flat terrain or near water wells; the
inhabitants stated that this species is very abundant in the
area.
RESULTS AND DISCUSSION—We examined a total of 281
specimens of 41 species, 35 genera, 18 families, and 7
orders of mammals that include 27% of the 152 species
reported for Guerrero (Ramı́rez-Pulido et al., 2000;
Cervantes et al., 2004; Almazán-Catalán et al., 2005;
Romo-Vázquez et al., 2005; Tejedor, 2005; Carraway, 2007;
Nolasco et al., 2007; Ruiz-Gutiérrez et al., 2011). The
mammal fauna included 17 species (41.5%) in the order
Chiroptera, nine species (21.9%) each in the orders
Rodentia and Carnivora, two species (4.9%) each in the
orders Didelphimorphia and Artiodactyla, and one species
(2.4%) each in the orders Cingulata and Lagomorpha. We
registered the presence of Puma concolor and P. yagouaroundi
on the basis of comments from villagers.
Eight of the species we documented are endemic to
Mexico including one didelphimorph (Tlacuatzin canes-
cens), four rodents (Hodomys alleni, Osgoodomys banderanus,
Peromyscus melanophrys, Sigmodon mascotensis), two bats
(Glossophaga morenoi, Artibeus hirsutus), and one lago-
morph (Sylvilagus cunicularius). These represent 19.5% of
the 41 mammalian species captured at Tlaxmalac, and
4.7% of the 169 endemic species of mammals reported
for Mexico (Ramı́rez-Pulido et al., 2005).
Relative Abundance—We captured 170 (60.5%) speci-
mens of the order Rodentia, 88 (31.3%) of Chiroptera, 13
(4.6%) of Carnivora, 4 (1.4%) of Lagomorpha, 3 (1.1%)
of Artiodactyla, 2 (0.7%) of Didelphimorphia, 1 (0.4%) of
Cingulata. The most abundant species of order Rodentia
were Osgoodomys banderanus (97 specimens, 34.5%),
Heteromys irroratus (50, 17.8%), and Baiomys musculus
(22, 7.8%). For bats, the most abundant species were
Balantiopteryx plicata (22 specimens, 7.8%), Artibeus
intermedius (12, 4.3%), A. jamaicensis (8, 2.8%), and
Pteronotus parnellii (8, 2.8%). The numbers of specimens
for the orders Didelphimorphia (2, 0.7%) and Cingulata
(1, 0.4%) reflect, in part, that our trapping approach was
less suited for those groups. However, we were able to
record 13 specimens of 9 species of Carnivora, 3
specimens of 2 species of Artiodactyla, and 4 specimens
of 1 species of Lagomorpha.
Of the 41 species documented, 31 were found in tropical
deciduous forest, 19 in agricultural fields, 10 in secondary
vegetation, and 1 in oak forest; some were found in more
than one type of vegetation. Only one species was found in
oak forest (Sciurus aureogaster), probably because of a low
March 2015 Almazán-Catalán et al.—Habitat use and reproduction of mammals from Guerrero, Mexico 41

TABLE 1—Mammals from Tlaxmalac, Guerrero, including common name, feeding strategy, use of vegetation types, use by local
people, and conservation status. Vegetation type: TDF = tropical deciduous forest, AF = agricultural fields, SV = secondary
vegetation, OF = oak forest. Traditional use: Fd = food, P = pet, Fr = fur, Me = medicinal. Conservation status: Norma Oficial
Mexicana (NOM)-059 (SEMARNAT, 2010), A = threatened, P = endangered; International Union for Conservation of Nature
(IUCN, 2014), NT = near threatened.

Species Common name Feeding Vegetation type Traditional use NOM-059 IUCN

Didelphis virginiana Virginia opossum Omnivorous SV, AF Fd — —

Tlacuatzin canescens Grayish mouse opossum Omnivorous TDF — — —
Dasypus novemcinctus Nine-banded armadillo Insectivorous TDF, SV, AF Fd, Fr, P — —
Sciurus aureogaster Red-bellied squirrel Herbivorous OF Fr, P — —
Otospermophilus variegatus Rock squirrel Omnivorous SV, AF P — —
Heteromys irroratus Mexican spiny pocket mouse Granivorous SV, AF — — —
Baiomys musculus Southern pygmy mouse Omnivorous SV, AF —- — —
Hodomys alleni Allen’s wood rat Omnivorous TDF — — —
Osgoodomys banderanus Michoacan deer mouse Omnivorous TDF, SV, AF — — —
Peromyscus melanophrys Plateau deer mouse Omnivorous TDF, AF — — —
Oryzomys couesi Coues’s rice rat Omnivorous SV, AF — — —
Sigmodon mascotensis West Mexican cotton rat Omnivorous AF — — —
Sylvilagus cunicularius Mexican cottontail Herbivorous SV, AF Fd — —
Balantiopteryx plicata Gray sac-winged bat Insectivorous TDF — — —
Desmodus rotundus Common vampire bat Hematofagous TDF, AF — — —
Choeronycteris mexicana Mexican long-tongued bat Nectarivorous TDF — A NT
Glossophaga leachi Gray’s long-tongued bat Nectarivorous TDF, AF — — —
Glossophaga morenoi Western long-tongued bat Nectarivorous TDF — — —
Glossophaga soricina Pallas’s long-tongued bat Nectarivorous TDF, AF — — —
Macrotus waterhousii Waterhouse’s leaf-nosed bat Insectivorous TDF — — —
Sturnira lilium Little yellow-shouldered bat Frugivorous TDF — — —
Artibeus hirsutus Hairy fruit-eating bat Frugivorous TDF — — —
Artibeus intermedius Intermediate fruit-eating bat Frugivorous TDF, AF — — —
Artibeus jamaicensis Jamaican fruit-eating bat Frugivorous TDF — — —
Artibeus lituratus Great fruit-eating bat Frugivorous TDF — — —
Dermanura tolteca Toltec fruit-eating bat Frugivorous TDF — — —
Pteronotus parnellii Parnell’s mustached bat Insectivorous TDF — — —
Nyctinomops femorosaccus Pocketed free-tailed bat Insectivorous TDF — — —
Nyctinomops macrotis Big free-tailed bat Insectivorous TDF — — —
Myotis velifer Cave myotis Insectivorous TDF — — —
Leopardus wiedii Margay Carnivorous TDF Fr P NT
Canis latrans Coyote Omnivorous TDF Fr — —
Urocyon cinereoargenteus Grey fox Omnivorous TDF, SV, AF Fr — —
Conepatus leuconotus American hog-nosed skunk Insectivorous AF Fd, Me — —
Mephitis macroura Hooded skunk Insectivorous AF Me — —
Spilogale putorius Eastern spotted skunk Insectivorous TDF, AF — — —
Bassariscus astutus Ringtail Omnivorous TDF — — —
Nasua narica White-nosed coati Omnivorous TDF Fd, Me, Fr, P — —
Procyon lotor Northern raccoon Omnivorous TDF Fd — —
Pecari tajacu Collared peccary Omnivorous TDF Fd — —
Odocoileus virginianus White-tailed deer Herbivorous TDF, SV, AF Fd, Fr, P — —

effort of capture, since access to oak forest is complicated
because of its rugged topography.
Mammals from Tlaxmalac represented eight different
feeding habits (Table 1), including omnivorous (n = 15,
36.6%), insectivorous (n = 10, 24.4%), frugivorous (n =
6, 14.6%), nectarivorous (n = 4, 9.8%), and herbivorous
(n = 3, 7.4%), which were most abundant, as well as
granivorous (Heteromys irroratus), hematophagous (Desmo-
dus rotundus), and carnivorous (Leopardus weidii), which
had only one species each (2.4%).
People from Tlaxmalac traditionally use 14 species of
wild mammals, some with more than one use. Eight
species are used as a complement of the diet, especially
medium- and large-size species, such as deer, javelinas,
rabbits, and coatis. Seven species are appreciated because
their skins are used to make musical instruments (e.g.,
armadillos and deer) or ornamental objects (e.g., margay,
coyotes, and squirrels). Three species have medicinal
uses: the fat from two species of skunks is used as liniment
and the skin of the coati to prevent hemorrhoids (by
sitting on it); occasionally armadillos, squirrels, coatis,
and deer are kept as pets (Table 1).
42 The Southwestern Naturalist vol. 60, no. 1

FIG. 1—Species accumulation curves comparing values of actual capture for mammals from Tlaxmalac; Bootstrap and Chao 2
estimators.

Choeronycteris mexicana and Leopardus wiedii are the only (34.92%). Tlaxmalac was most different from Omiltelmi;
species designated as being at some level of risk of these regions shared only 21.79% of species, the latter
extinction according to the Norma Oficial Mexicana only with temperate forest. Comparing with studies from
NOM-059-ECOL-2010 (SEMARNAT, 2010). Choeronycteris the same general area, we found a higher number (n =
mexicana might be considered rare in the Balsas River 41) of mammal species than reported by Álvarez (1968; n
basin because capture rates are variable, depending on = 23), probably due to differences in environmental
several factors such as migration, season, and food complexities, with Tlaxmalac having more kinds of
availability (Villa, 1967; Fleming and Eby, 2003). On the microhabitats.
basis of reported uses by local people, hunting may be a Little is known about the basic ecology of many species
factor threatening L. wiedii. of mammals in the Balsas River basin; thus, additional
The estimated richness with Chao2 and Bootstrap was studies are needed in local areas such as Tlaxmalac.
49 species; this indicates that we registered 88% of the Information provided for the traditional use for medium
expected mammals in Tlaxmalac (Fig. 1), including the and large mammals in Tlaxmalac represents an additional
records of Puma concolor and P. yagouaroundi cited by the contribution to their conservation. To the extent that we
villagers but not examined or captured by us. According know the species richness of a given area and compare it
to the predictive model, we did not document six species
that occur in the region. It is highly likely that Mormoops
megalophylla, Pteronotus davyi, Anoura geoffroyi, Leptonycteris
yerbabuenae, Mustela frenata, and Reithrodontomys fulvescens
could inhabit the study area considering that their
distribution includes localities with tropical deciduous
forests in the western part of Mexico, and they have been
recorded in nearby places such as Sierra de Huautla
(Sánchez-Hernández and Romero-Almaraz, 1995).
The dendrogram (Fig. 2) shows the regions grouped
according to similarity of species richness and similarity of
vegetation type. We found that Tlaxmalac was more
similar to Sierra de Huautla, Morelos (53.85%; Sánchez-
Hernández and Romero-Almaraz, 1995), sharing 29
species. These two areas are only separated by 55 km
and the type of vegetation at both is tropical deciduous FIG. 2—Dendrogram of similarity using Jaccard coefficient
forest. Tlaxmalac exhibited lower similarity with Sierra de between mammals from Tlaxmalac and four other study areas:
Taxco (41.54%), which has temperate forest surrounded 1) Costa Grande, 3) Sierra of Taxco, 4) Omiltemi, in Guerrero,
by tropical vegetation, and Costa Grande de Guerrero and 2) Sierra of Huautla, in Morelos.
March 2015 Almazán-Catalán et al.—Habitat use and reproduction of mammals from Guerrero, Mexico
with other regions, we can establish priority conservation
actions of such regions with special ecological, physio-
graphic, and evolutionary conditions.
We thank Y. Q. Jiménez-Salmerón for her help with fieldwork,
and G. D. Schnell and R. C. Almazán-Núñez for their suggestions
to improve the manuscript. Collecting permits were provided by
the Instituto Nacional de Ecologı́a, Dirección General de Vida
Silvestre (02.-6538, 02.-3701, and FAUT.0103) to CS-H.
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44 The Southwestern Naturalist
APPENDIX—Gazeteer of localities where mammals were
surveyed or collected in Tlaxmalac, Guerrero. Localities
identified with lowercase italic letters were geographically
close.
1. La Pedreguera, 5.5 km NE de Tlaxmalac, 1,500 m,
18824 0 32 00 N, 99824 0 12 00 W.
2. El Tule, 4.5 km NE de Tlaxmalac, 1,280 m,
18824 0 17 00 N, 99824 0 00 00 W.
2a. El Tule, 4 km NE de Tlaxmalac, 1,240 m,
18824 0 06 00 N, 99823 0 58 00 W.
2b. El Tule, 4 km NE de Tlaxmalac, 1,240 m,
18824 0 00 00 N, 99824 0 01 00 W.
3. El Tecal, 4 km NE de Tlaxmalac, 1,200 m,
18823 0 34 00 N, 99823 0 05 00 W.
3a. El Tecal, 3.5 km NE de Tlaxmalac, 1,200 m,
18823 0 27 00 N, 99823 0 13 00 W.
4. Barranca de Garcı́a, 3.8 km NW de Tlaxmalac, 940
m, 18823 0 34 00 N, 99826 0 12 00 W.
4a. Barranca de Garcı́a, 1.5 km SW de Tlaxmalac, 900
m, 18822 0 14 00 N, 99825 0 36 00 W.
4b. Barranca de Garcı́a, 750 m W de Tlaxmalac, 900 m,
18821 0 56 00 N, 99825 0 24 00 W.
5. La Cofradı́a, 2.5 km NW de Tlaxmalac, 960 m,
18822 0 54 00 N, 99825 0 39 00 W.
6. Cueva La Cofradı́a, 2.5 km NW de Tlaxmalac, 1,020
m, 18823 0 06 00 N, 99825 0 08 00 W.
7. La Campana, 2.2 km N de Tlaxmalac, 1,060 m,
18823 0 00 00 N, 99823 0 46 00 W.
8. Cerro El Papayo, 3.5 km NE de Tlaxmalac, 1,400 m,
18823 0 02 00 N, 99823 0 00 00 W.
8a. Cerro El Papayo, 2.5 km NE de Tlaxmalac, 1,120 m,
18823 0 11 00 N, 99823 0 34 00 W.
9. Huzicuate, 3 km NE de Tlaxmalac, 1,260 m,
18822 0 56 0 ’N, 99823 0 08 0 ’W.
10. Cerro El Tenaño, 3.5 km N de Tlaxmalac, 1,200 m,
18823 0 24 00 N, 99824 0 40 00 W.
10a. Cerro El Tenaño, 3 km N de Tlaxmalac, 1,080 m,
18822 0 53 00 N, 99824 0 42 00 W.
10b. Cerro El Tenaño, 2 km N de Tlaxmalac, 920 m,
18822 0 30 00 N, 99824 0 30 00 W.
10c. Cerro El Tenaño, 1.5 km N de Tlaxmalac, 960 m,
18822 0 34 00 N, 99824 0 49 00 W.
11. Barranca Tepeaxtla, 2 km NE de Tlaxmalac, 1,000
m, 18822 0 51 00 N, 99824 0 23 00 W.
11a. Barranca Tepeaxtla, 1 km NE de Tlaxmalac, 930 m,
18822 0 38 00 N, 99824 0 24 00 W.
12. Poza de Los Santos, 2 km NE de Tlaxmalac, 1,100
m, 18822 0 44 00 N, 99824 0 07 00 W.
12a. Poza de Los Santos, 2 km NE de Tlaxmalac, 1,060
m, 18822 0 39 00 N, 99824 0 00 00 W.
12b. Poza de Los Santos, 1.5 km NE de Tlaxmalac, 940
m, 18822 0 39 00 N, 99824 0 13 00 W.
13. Las Juntas, 1 km NE de Tlaxmalac, 1,000 m,
18822 0 43 00 N, 99824 0 17 00 W.
vol. 60, no. 1
13a. Las Juntas, 1 km NE de Tlaxmalac, 900 m,
18822 0 32 00 N, 99824 0 22 00 W.
13b. Las Juntas, 500 m W de Tlaxmalac, 900 m,
18822 0 25 00 N, 99824 0 25 00 W.
14. Barranca Xihuatoxtla, 2.5 km NE de Tlaxmalac,
1,060 m, 18822 0 53 00 N, 99824 0 05 00 W.
14a. Barranca Xihuatoxtla, 2 km NE de Tlaxmalac, 1,000
m, 18822 0 42 00 N, 99824 0 05 00 W.
14b. Barranca Xihuatoxtla, 1.5 km NE de Tlaxmalac, 980
m, 18822 0 36 00 N, 99824 0 13 00 W.
14c. Barranca Xihuatoxtla, 1.5 km NE de Tlaxmalac, 960
m, 18822 0 34 00 N, 99824 0 15 00 W.
15. Cerro Grande, 1.5 km NE de Tlaxmalac, 1,200 m,
18822 0 16 00 N, 99823 0 27 00 W.
16. Cerro El Jumil, 2.5 km W de Tlaxmalac, 1,040 m,
18822 0 10 00 N, 99826 0 20 00 W.
16a. Cerro El Jumil, 2.5 km NW de Tlaxmalac, 980 m,
18822 0 19 00 N, 99826 0 17 00 W.
16b. Cerro El Jumil, 3.7 km SW de Tlaxmalac, 980 m,
18821 0 09 00 N, 99826 0 58 00 W.
17. Calle Rodeo, Tlaxmalac, 900 m, 18821 0 53 00 N,
99825 0 03 00 W.
18. Biblioteca, Tlaxmalac, 900 m, 18821 0 31 00 N,
99824 0 49 00 W.
19. Cerro Chiltepec, 2 km W de Tlaxmalac, 1,100 m,
18821 0 19 00 N, 99825 0 20 00 W.
19a. Cerro Chiltepec, 1.5 km W de Tlaxmalac, 900 m,
18821 0 37 00 N, 99825 0 13 00 W.
19b. Cerro Chiltepec, 1 km W de Tlaxmalac, 980 m,
18821 0 44 00 N, 99825 0 42 00 W.
19c. Cerro Chiltepec, 500 m W de Tlaxmalac, 920 m,
18821 0 34 00 N, 99825 0 12 00 W.
19d. Cerro Chiltepec, 1.5 km SW de Tlaxmalac, 980 m,
18821 0 32 00 N, 99825 0 13 00 W.
20. 2.5 km SW de Tlaxmalac, 900 m, 18821 0 09 00 N,
99826 0 08 00 W.
21. 3.5 km SW de Tlaxmalac, 900 m, 18820 0 55 00 N,
99826 0 22 00 W.
22. La Laguna, 2.5 km SW de Tlaxmalac, 900 m,
18820 0 41 00 N, 99826 0 06 00 W.
23. Carretera, 1 km S de Tlaxmalac, 900 m, 18821 0 03 00 N,
99824 0 54 00 W.
24. Carretera, 1.5 km S de Tlaxmalac, 900 m,
18820 0 29 00 N, 99824 0 42 00 W.
25. Carretera, 2 km S de Tlaxmalac, 900 m, 18820 0 05 00 N,
99824 0 40 00 W.
26. El Ópalo, 3.8 km SE de Tlaxmalac, 960 m,
18820 0 37 00 N, 99823 0 01 00 W.
26a. El Ópalo, 3.6 km SE de Tlaxmalac, 900 m,
18820 0 32 00 N, 99823 0 12 00 W.
27. Pozo El Hundido, 4 km S de Tlaxmalac, 900 m,
18820 0 27 00 N, 99823 0 49 00 W.
28. Tlatocan, 3 km SW de Tlaxmalac, 900 m,
18820 0 11 00 N, 99825 0 56 00 W.