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

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 regions. A total of 152 species of mammals is reported from Guerrero, Mexico (Ramirez-Pulido et al., 2000; Cervantes et al., 2004; Almazan-Catakm et al., 2005; Romo-Vazquez et al., 2005; Tejedor, 2005; Carraway, 2007; Nolasco et al., 2007; Ruiz-Gutierrez et al., 2011). The Balsas River basin is formed by two physiographic provinces, the TransVolcanic 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 characteristics 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 (Alvarez, 1968). Some mammals of this region have only been recorded occasionally by indirect ways (e.g., in pellets of barn owl Tyto alba; Ramirez-Pulido and Sanchez-Hernandez, 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 18[degrees]21'38"N, 99[degrees]24'50"W and has an area of 40 [km.sup.2]. 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 25.5[degrees]C and average annual rainfall is 1,417 mm, with most rain occurring from June 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 dominated by plants such as Acacia cymbispina (acacia), A. farnesiana (huizache, acacia), Haematoxylum brasiletto (Brazilwood), Pithecellobium dulce (guamuchil), and Alvaradoa amorphoides (guachipil; Jimenez Salmeron and Garcia, 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 x 3.5 x 9 in) for small to medium-size mammals, and 900 trap-nights using snap traps (7 x 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 Coleccion Nacional de Mamiferos, Instituto de Biologia, Universidad Nacional Autonoma de Mexico; 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 RamirezPulido 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 housewives. Conservation status of species was stated according to Norma Oficial Mexicana NOM-059-SEMARNAT-2010 (SEMARNAT, 2010), and according to International Union for Conservation 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 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 (RamirezPulido et al., 1977), Sierra de Taxco (Leon-Paniagua and RomoVazquez, 1993), Omiltemi in the state of Guerrero (JimenezAlmaraz et al., 1993), and Sierra de Huautla in the state of Morelos (Sanchez-Hernandez and Romero-Almaraz, 1995).

SPECIES ACCOUNTS--Didelphis virginiana californica--Locality 13b. We captured one male with scrotal testes 20 x 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 x 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 (Coleccion Nacional de Mamiferos, Instituto de Biologia, Universidad Nacional Autonoma de Mexico 42516). The specimen 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 x 14 mm; 1L, 10 x 13 mm), and one taken in October had four embryos (2R, 2L, 21 x 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 x 4 mm, 13 x 10 mm), five in July (19 x 12 mm to 20 x 14 mm), and one in August (20 x 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 x 2 mm and 7 x 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 x 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 x 5 mm; 2L, 15 x 14 mm; 1R, 1L, 2.4 x 1.5 mm) and one had three embryos (3R, 25 x 17). Of six females caught in February, two had two embryos (2L, 1 x 1 mm; 2R, 3 x 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 x 11 mm), six were lactating, one was lactating and had three embryos (16 x 12 mm), and one was inactive. Of three females caught in July, one had two embryos (1R, 1L; 23 x 14 mm), one had three embryos (3L, 17 x 13 mm), and one was inactive. We captured four females in August, including one with three embryos (1R, 2L, 6 x 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 x 13 mm), three had three embryos (3L, 15 x 8 mm; 3R, length 25 mm; 2R, 1L, length 18 mm), and one had four embryos (4L, 20 x 15 mm). We captured two females in October including one with three embryos (3L, 9 x 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 x 10 mm to 20 x 10 mm), 4 from February (16 x 11 mm to 21 x 8 mm), 10 from May (6 x 3 mm to 21 x 10 mm), 1 from June (17 x 10 mm), 8 from July (17 x 9 mm to 20 x 11 mm), 4 from August (14 x 8 mm to 26 x 10 mm), and 5 from September (7 x 5 mm to 20 x 10 mm).

Peromyscus melanophrys melanophrys--Locality 27. We captured one pregnant female with four embryos (2R, 2L, 7 x 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 x 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 x 24 mm; 1L, 20 x 20 mm), and one caught in June 2001 had three embryos (1R, 27 x 18 mm; 2L, 37 x 30 mm). The males were caught in October 2000 and June 2001 and had scrotal testes 24 x 8 mm and 20 x 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 reproductively inactive, two from June 2001 and one from August 2000 were lactating, and one from June 2001 had an embryo 22 x 14 mm. Three males caught in June were newborns, two from August had inguinal testes (3 x 2 mm), and we did not record reproductive information 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 x 6 mm (in June) and 7 x 4 mm and 7 x 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 x 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 x 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 x 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 x 4 mm) and one in December (5 x 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 x 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 x 4 mm). One male from December had scrotal testes (8 x 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 December (n = 2). Of the females, one from June was receptive, and two were pregnant (June, 6 x 4 mm; December, 8 x 8 mm). Eight males had scrotal testes including two from May (6 x 4 mm, 9 x 6 mm), five from June (5 x 4 mm to 13 x 8 mm), and one from December (7 x 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 x 5 mm), and one was reproductively inactive. Males had scrotal testes in March (7 x 5 mm), May (8 x 5 mm), June (13 x 9 mm), and December (7 x 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 x 5 mm, 8 x 6 mm) and one in December (7 x 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 x 11 mm) and another in December (2 x 2 mm); one from December was lactating, and four were reproductively inactive. The male had scrotal testes (4 x 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 (Coleccion Nacional de Mamiferos, Instituto de Biologia, Universidad Nacional Autonoma de Mexico, 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 (Coleccion Nacional de Mamiferos, Instituto de Biologia, Universidad Nacional Autonoma de Mexico, 42515) with inguinal testes of 2 x 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 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 x 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 x 17 mm, 18 x 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 November 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 x 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 vegetation is better preserved. They represent a significant 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 (Ramirez-Pulido et al., 2000; Cervantes et al., 2004; Almazan-Catakrn et al., 2005; Romo-Vazquez et al., 2005; Tejedor, 2005; Carraway, 2007; Nolasco et al., 2007; Ruiz-Gutierrez 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 canescens), four rodents (Hodomys alleni, Osgoodomys banderanus, Peromyscus melanophrys, Sigmodon mascotensis), two bats (Glossophaga morenoi, Artibeus hirsutus), and one lagomorph (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 (Ramirez-Pulido et al., 2005).

Relative Abundance--We captured 170 (60.5%) specimens 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 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 (Desmodus 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).

Choeronycteris mexicana and Leopardus wiedii are the only species designated as being at some level of risk of extinction according to the Norma Oficial Mexicana NOM-059-ECOL-2010 (SEMARNAT, 2010). Choeronycteris mexicana might be considered rare in the Balsas River basin because capture rates are variable, depending on several factors such as migration, season, and food availability (Villa, 1967; Fleming and Eby, 2003). On the basis of reported uses by local people, hunting may be a factor threatening L. wiedii.

The estimated richness with Chao2 and Bootstrap was 49 species; this indicates that we registered 88% of the expected mammals in Tlaxmalac (Fig. 1), including the records of Puma concolor and P. yagouaroundi cited by the villagers but not examined or captured by us. According 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 (Sanchez-Hernandez 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%; Sanchez-Hernandez 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 forest. Tlaxmalac exhibited lower similarity with Sierra de Taxco (41.54%), which has temperate forest surrounded by tropical vegetation, and Costa Grande de Guerrero (34.92%). Tlaxmalac was most different from Omiltelmi; these regions shared only 21.79% of species, the latter only with temperate forest. Comparing with studies from the same general area, we found a higher number (n = 41) of mammal species than reported by Alvarez (1968; n = 23), probably due to differences in environmental complexities, with Tlaxmalac having more kinds of microhabitats.

Little is known about the basic ecology of many species of mammals in the Balsas River basin; thus, additional studies are needed in local areas such as Tlaxmalac. Information provided for the traditional use for medium and large mammals in Tlaxmalac represents an additional contribution to their conservation. To the extent that we know the species richness of a given area and compare it with other regions, we can establish priority conservation actions of such regions with special ecological, physiographic, and evolutionary conditions.

We thank Y. Q. Jimenez-Salmeron for her help with fieldwork, and G. D. Schnell and R. C. Almazan-NUnez for their suggestions to improve the manuscript. Collecting permits were provided by the Instituto Nacional de Ecologia, Direccion General de Vida Silvestre (02.-6538, 02.-3701, and FAUT.0103) to CS-H.


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Submitted 6 October 2013.

Acceptance recommended by Associate Editor, Jennifer K. Frey, 11 July 2014.

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, 18[degrees]24'32"N, 99[degrees]24'12"W.

2. El Tule, 4.5 km NE de Tlaxmalac, 1,280 m, 18[degrees]24'17"N, 99[degrees]24'00"W.

2a. El Tule, 4 km NE de Tlaxmalac, 1,240 m, 18[degrees]24'06"N, 99[degrees]23'58"W.

2b. El Tule, 4 km NE de Tlaxmalac, 1,240 m, 18[degrees]24'00"N, 99[degrees]24'01"W.

3. El Tecal, 4 km NE de Tlaxmalac, 1,200 m, 18[degrees]23'34"N, 99[degrees]23'05"W.

3a. El Tecal, 3.5 km NE de Tlaxmalac, 1,200 m, 18[degrees]23'27"N, 99[degrees]23'13"W.

4. Barranca de Garcia, 3.8 km NW de Tlaxmalac, 940 m, 18[degrees]23'34"N, 99[degrees]26'12"W.

4a. Barranca de Garcia, 1.5 km SW de Tlaxmalac, 900 m, 18[degrees]22'14"N, 99[degrees]25'36"W.

4b. Barranca de Garcia, 750 m W de Tlaxmalac, 900 m, 18[degrees]21'56"N, 99[degrees]25'24"W.

5. La Cofradia, 2.5 km NW de Tlaxmalac, 960 m, 18[degrees]22'54"N, 99[degrees]25'39"W.

6. Cueva La Cofradia, 2.5 km NW de Tlaxmalac, 1,020 m, 18[degrees]23'06"N, 99[degrees]25'08"W.

7. La Campana, 2.2 km N de Tlaxmalac, 1,060 m, 18[degrees]23'00"N, 99[degrees]23'46"W.

8. Cerro El Papayo, 3.5 km NE de Tlaxmalac, 1,400 m, 18[degrees]23'02"N, 99[degrees]23'00"W.

8a. Cerro El Papayo, 2.5 km NE de Tlaxmalac, 1,120 m, 18[degrees]23'11"N, 99[degrees]23'34"W.

9. Huzicuate, 3 km NE de Tlaxmalac, 1,260 m, 18[degrees]22'56"N, 99[degrees]23'08"W.

10. Cerro El Tenano, 3.5 km N de Tlaxmalac, 1,200 m, 18[degrees]23'24"N, 99[degrees]24'40"W.

10a. Cerro El Tenafio, 3 km N de Tlaxmalac, 1,080 m, 18[degrees]22'53"N, 99[degrees]24'42"W.

10b. Cerro El Tenafio, 2 km N de Tlaxmalac, 920 m, 18[degrees]22'30"N, 99[degrees]24'30"W.

10c. Cerro El Tenano, 1.5 km N de Tlaxmalac, 960 m, 18[degrees]22'34"N, 99[degrees]24'49"W.

11. Barranca Tepeaxtla, 2 km NE de Tlaxmalac, 1,000 m, 18[degrees]22'51"N, 99[degrees]24'23"W.

11a. Barranca Tepeaxtla, 1 km NE de Tlaxmalac, 930 m, 18[degrees]22'38"N, 99[degrees]24'24"W.

12. Poza de Los Santos, 2 km NE de Tlaxmalac, 1,100 m, 18[degrees]22'44"N, 99[degrees]24'07"W.

12a. Poza de Los Santos, 2 km NE de Tlaxmalac, 1,060 m, 18[degrees]22'39"N, 99[degrees]24'00"W.

12b. Poza de Los Santos, 1.5 km NE de Tlaxmalac, 940 m, 18[degrees]22'39"N, 99[degrees]24'13"W.

13. Las Juntas, 1 km NE de Tlaxmalac, 1,000 m, 18[degrees]22'43"N, 99[degrees]24'17"W.

13a. Las Juntas, 1 km NE de Tlaxmalac, 900 m, 18[degrees]22'32"N, 99[degrees]24'22"W.

13b. Las Juntas, 500 m W de Tlaxmalac, 900 m, 18[degrees]22'25"N, 99[degrees]24'25"W.

14. Barranca xihuatoxtla, 2.5 km NE de Tlaxmalac, l,060 m, 18[degrees]22'53"N, 99[degrees]24'05"W.

14a. Barranca Xihuatoxtla, 2 km NE de Tlaxmalac, 1,000 m, 18[degrees]22'42"N, 99[degrees]24'05"W.

14b. Barranca Xihuatoxtla, 1.5 km NE de Tlaxmalac, 980 m, 18[degrees]22'36"N, 99[degrees]24'13"W.

14c. Barranca Xihuatoxtla, 1.5 km NE de Tlaxmalac, 960 m, 18[degrees]22'34"N, 99[degrees]24'15"W.

15. Cerro Grande, 1.5 km NE de Tlaxmalac, 1,200 m, 18[degrees]22'16"N, 99[degrees]23'27"W.

16. Cerro El Jumil, 2.5 km W de Tlaxmalac, 1,040 m, 18[degrees]22'10"N, 99[degrees]26'20"W.

16a. Cerro El Jumil, 2.5 km NW de Tlaxmalac, 980 m, 18[degrees]22'19"N, 99[degrees]26'17"W.

16b. Cerro El Jumil, 3.7 km SW de Tlaxmalac, 980 m, 18[degrees]21'09"N, 99[degrees]26'58"W.

17. Calle Rodeo, Tlaxmalac, 900 m, 18[degrees]21'53"N, 99[degrees]25'03"W.

18. Biblioteca, Tlaxmalac, 900 m, 18[degrees]21'31"N, 99[degrees]24'49"W.

19. Cerro Chiltepec, 2 km W de Tlaxmalac, 1,100 m, 18[degrees]21'19"N, 99[degrees]25'20"W.

19a. Cerro Chiltepec, 1.5 km W de Tlaxmalac, 900 m, 18[degrees]21'37"N, 99[degrees]25'13"W.

19b. Cerro Chiltepec, 1 km W de Tlaxmalac, 980 m, 18[degrees]21'44"N, 99[degrees]25'42"W.

19c. Cerro Chiltepec, 500 m W de Tlaxmalac, 920 m, 18[degrees]21'34"N, 99[degrees]25'12"W.

19d. Cerro Chiltepec, 1.5 km SW de Tlaxmalac, 980 m, 18[degrees]21'32"N, 99[degrees]25'13"W.

20. 2.5 km SW de Tlaxmalac, 900 m, 18[degrees]21'09"N, 99[degrees]26'08"W.

21. 3.5 km SW de Tlaxmalac, 900 m, 18[degrees]20'55"N, 99[degrees]26'22"W.

22. La Laguna, 2.5 km SW de Tlaxmalac, 900 m, 18[degrees]20'41"N, 99[degrees]26'06"W.

23. Carretera, 1 km S de Tlaxmalac, 900 m, 18[degrees]21'03"N, 99[degrees]24'54"W.

24. Carretera, 1.5 km S de Tlaxmalac, 900 m, 18[degrees]20'29[degrees]N, 99[degrees]24'42[degrees]W.

25. Carretera, 2 km S de Tlaxmalac, 900 m, 18[degrees]20'05"N, 99[degrees]24'40"W.

26. El (Opalo, 3.8 km SE de Tlaxmalac, 960 m, 18[degrees]20'37"N, 99[degrees]23'01"W.

26a. El (Opalo, 3.6 km SE de Tlaxmalac, 900 m, 18[degrees]20'32"N, 99[degrees]23'12"W.

27. Pozo El Hundido, 4 km S de Tlaxmalac, 900 m, 18[degrees]20'27"N, 99[degrees]23'49"W.

28. Tlatocan, 3 km SW de Tlaxmalac, 900 m, 18[degrees]20'11"N, 99[degrees]25'56"W.


Departamento de Zoologia, Instituto de Biologia, Universidad Nacional Autonoma de Mexico, A.P. 70-153, Coyoacan, Mexico, D.F. 04510, Mexico (JAA-C, CS-H, LS-V, SBG-P)

Escuinapa No. 92 bis. Col. Pedregal de Santo Domingo, C.P. 04369, Mexico, D.F., Mexico (MLR-A)

* Correspondent.

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

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

Species                      Vegetation type   Traditional use

Didelphis virginiana         SV, AF            Fd
Tlacuatzin canescens         TDF                     --
Dasypus novemcinctus         TDF, SV, AF       Fd, Fr, P
Sciurus aureogaster          OF                Fr, P
Otospermophilus variegatus   SV, AF            P
Heteromys irroratus          SV, AF                  --

Baiomys musculus             SV, AF                  --
Hodomys alleni               TDF                     --
Osgoodomys banderanus        TDF, SV, AF             --
Peromyscus melanophrys       TDF, AF                 --
Oryzomys couesi              SV, AF                  --
Sigmodon mascotensis         AF                      --
Sylvilagus cunicularius      SV, AF            Fd
Balantiopteryx plicata       TDF                     --
Desmodus rotundus            TDF, AF                 --
Choeronycteris mexicana      TDF                     --

Glossophaga leachi           TDF, AF                 --
Glossophaga morenoi          TDF                     --

Glossophaga soricina         TDF, AF                 --

Macrotus waterhousii         TDF                     --

Sturnira lilium              TDF                     --

Artibeus hirsutus            TDF                     --
Artibeus intermedius         TDF, AF                 --

Artibeus jamaicensis         TDF                     --

Artibeus lituratus           TDF                     --
Dermanura tolteca            TDF                     --
Pteronotus parnellii         TDF                     --
Nyctinomops femorosaccus     TDF                     --

Nyctinomops macrotis         TDF                     --
Myotis velifer               TDF                     --
Leopardus wiedii             TDF               Fr
Canis latrans                TDF               Fr
Urocyon cinereoargenteus     TDF, SV, AF       Fr
Conepatus leuconotus         AF                Fd, Me

Mephitis macroura            AF                Me
Spilogale putorius           TDF, AF                 --
Bassariscus astutus          TDF                     --
Nasua narica                 TDF               Fd, Me, Fr, P
Procyon lotor                TDF               Fd
Pecari tajacu                TDF               Fd
Odocoileus virginianus       TDF, SV, AF       Fd, Fr, P

Species                      NOM-059   IUCN

Didelphis virginiana           --       --
Tlacuatzin canescens           --       --
Dasypus novemcinctus           --       --
Sciurus aureogaster            --       --
Otospermophilus variegatus     --       --
Heteromys irroratus            --       --

Baiomys musculus               --       --
Hodomys alleni                 --       --
Osgoodomys banderanus          --       --
Peromyscus melanophrys         --       --
Oryzomys couesi                --       --
Sigmodon mascotensis           --       --
Sylvilagus cunicularius        --       --
Balantiopteryx plicata         --       --
Desmodus rotundus              --       --
Choeronycteris mexicana      A         NT

Glossophaga leachi             --       --
Glossophaga morenoi            --       --

Glossophaga soricina           --       --

Macrotus waterhousii           --       --

Sturnira lilium                --       --

Artibeus hirsutus              --       --
Artibeus intermedius           --       --

Artibeus jamaicensis           --       --

Artibeus lituratus             --       --
Dermanura tolteca              --       --
Pteronotus parnellii           --       --
Nyctinomops femorosaccus       --       --

Nyctinomops macrotis           --       --
Myotis velifer                 --       --
Leopardus wiedii             P         NT
Canis latrans                  --       --
Urocyon cinereoargenteus       --       --
Conepatus leuconotus           --       --

Mephitis macroura              --       --
Spilogale putorius             --       --
Bassariscus astutus            --       --
Nasua narica                   --       --
Procyon lotor                  --       --
Pecari tajacu                  --       --
Odocoileus virginianus         --       --
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