TEXAS TECH UNIVERSITY Natural Science Research Laboratory Occasional Papers Museum of Texas Tech University Number 305 16 December 2011 Small Mammals of Sangay National Park, Chimborazo Province and Morona Santiago Province, Ecuador Thomas E. Lee, Jr., Carlos Boada-Teran, AmyM. Scott, Santiago F. Burneo, andJ. DeltonHanson Abstract In 2010 a mammal survey was conducted in an ecotone region (including paramo and temperate forest) on the eastern slope of the Andes in Sangay National Park, Ecuador. Sherman traps, Tomahawk traps, pitfall traps, and mist nets were used to collect mammal specimens at two sites (3,400 m elevation and 2,962 m elevation). A total of 169 specimens representing 14 species were collected from the survey area. The species collected include: Caenolestes caniven¬ ter, Cryptotis montivaga, Akodon mollis, Microryzomys altissimus, M. minutus, Thomasomys praetor, T. baeops, T. cinnameus, T. par amor um, T. silvestris, Sturnira bide ns, S. erythromos, Micronycteris megalotis, and Myotis keaysi. One additional species (Mazama rufina) was re¬ corded photographically. In addition, a new national record of a parasitic beetle (Amblyopinus colombiae) was discovered on a specimen of T. praetor. Key words: Chimborazo, distribution range, Ecuador, mammals, Morona Santiago, Sangay National Park Resumen En el ano 2010 se llevo a cabo un inventario de mamiferos en una region de ecotono (que incluye paramo y bosque templado) en el Parque Nacional Sangay en, las estribaciones orientales de los Andes del Ecuador. Se usaron trampas Sherman, Tomahawk, pitfall, y redes de neblina para colectar especimenes de mamiferos en los dos sitios de estudio (a elevaciones entre 2,962 y 3,400 msnm). Un total de 169 especimenes de 14 especies fueron colectados en el area de estudio. Las especies colectadas incluyen: Caenolestes caniventer, Cryptotis montivaga, Akodon mollis, Microryzomys altissimus, M. minutus, Thomasomys praetor, T. baeops, T. cin¬ nameus, T. paramorum, T. silvestris, Sturnira bidens, S. erythromos, Micronycteris megalotis, y Myotis keaysi. Una especie adicional (Mazama rufina) fue registrada fotograficamente. Un nuevo registro nacional de escarabajo parasito (Amblyopinus colombiae) fue descubierto en un especimen de T. praetor. Palabras clave: Chimborazo, Ecuador, mamiferos, Morona Santiago, Parque Nacional Sangay, rango de vida 2 Occasional Papers, Museum of Texas Tech University Introduction The purpose of this study was to document the This survey is important because park officials small mammal assemblage of Parque Nacional San- wanted to know what species of small mammals oc¬ gay in south central Ecuador. Ecuador has had a long curred in Parque Nacional Sangay. The long-term history of mammalian surveys beginning in the early goal of this project and related studies is to acquire 1900s (e.g., Stone 1914; Allen 1916; Anthony 1924a, comparative material to facilitate studies of the bio¬ 1924b; Tate 1931; Barnett 1999; Lee et al. 2006a, geography, ecology, and evolution of mammals in 2006b, 2008, 2010; McDonough et al. 2011). Other Ecuador (Jarrin and Fonseca 2001; Lee et al. 2006a, studies conducted in the same region and elevation as 2006b, 2008, 2010). Parque Nacional Sangay include surveys of Cajas Pla¬ teau and the corridor between Sangay and Llanganates National Parks (Barnett 1999; Haynie et al. 2006). Study Area and Methods This study was conducted from 21 July to 11 Passifloraceae, Fabaceae, Onagraceae, Orchidaceae, August 2010 at two sites (3,400 m and 2,962 m el¬ Polypodiaceae (ferns), and Valerianaceae. In places evation) in the eastern paramo and temperate forests, where the soil is not well drained, Sphagnum moss is respectively, in Parque Nacional Sangay, which is abundant. Stands of reed in the family Cyperaceae located on the border between Chimborazo and Mo- were observed in the lagoons. rona Santiago Provinces on the eastern slope of the Andes. The study area includes mountain lakes, called Site 2 is southeast of Site 1, in Morona San¬ the Atillo Lagoons, and their associated marshes and tiago Province, at 2,962 m elevation (2°12T7.87"S, Polylepis forests. The paramo and temperate forests 78°27'30.92"W) (Fig. 1). This site was sampled for of eastern Ecuador are characterized by high rodent 16 days from 25 July to 9 August 2010. This area diversity and low bat diversity as compared to other consisted of secondary forests and a riparian habitat similarly sampled areas (Lee et al. 2006a, 2006b, 2008, on the Rio Upano. Most of the traps and nets were set 2010). The study sites therefore were chosen because near the edge of the secondary growth managed forest of the lack of documented mammals in Parque Nacional and a riparian forest. At this site the plant families Al- Sangay and to investigate differences in mammalian stroemeriaceae, Bromeliaceae, Clusiaceae, and Poaceae diversity due to changes in elevation within a small also were noted along the stream banks. For both sites, area (Patterson et al. 1996). identification of plant families was conducted using Patzelt (2004), and photos of plants from the sites Two locations, 4.69 km apart, were surveyed dur¬ were examined by Ken Sytsma (Department of Botany, ing this study, one near the Atillo Lagoons and another University of Wisconsin) for confirmation. within the drainage of the Rio Upano (Fig. 1). Site 1, at 3,400 m elevation, includes a series of trap locations Five Tomahawk and 170 Sherman traps were set near the Atillo Lagoons, a basin of lakes surrounded in the grasslands, on the forest floor, in trees, in streams, by mountains that are between 3,900 and 4,200 m and on stream banks for 3,675 trap-nights. Bats were high (2°H'33.4"S, 78°3r29.38"W to 2°10'55.99"S, caught with four, 9-18 m long mist nets set in or near 78°29'57.36"W). We collected at these locations from riparian habitat at the 2,962 site (192 net-hours). Pitfall 21 July to 11 August 2010 (21 days). The habitat is traps were set in forests and along stream banks and primarily alpine grassland dominated by Stipa ichu logs. All voucher specimens (skins, skulls, skeletons, (Poaceae) with patches of Polylepis forests. Other and frozen liver tissue) were deposited in the Abilene plant families that we identified in this habitat were: Christian University Natural History Collection Asteraceae, Bromeliaceae (Puya), Calceolariaceae, (ACUNHC) and at the Museo de Zoologia (QCAZ), Lee et al.—Small Mammals of Sangay National Park, Ecuador 3 79° W 78’W 77°W 76*W COLOMBIA PERU Cotopaxi -rs ungurahu Pastaza 2 s- -2°S Morona Santiago 79°W 73&W 77flW 76°W Legend -- Rivers (cid:9633) Province Boundary , • ] Protected Areas Elevation High: 6260 Low -39 Figure 1. The location of the study sites in Parque Nacional Sangay are designated by the numbers 1 (3,400 m elevation) and 2 (2,962 m elevation) (also see inset map). The other study sites used for comparison are: location 3, the Tandayapa Valley (0°00T3"N, 78o40'70"W); location 4, the Casanga River Valley sites (0°33'00"S, 77°55,00"Wand 0o3T70MS, 77°52,99"W); location 5, the eastern slope of Volcan Sumaco (0°34T9MS, 77°35'64"W); and location 6, the Santa Rosa sites (0o19'51”N, 78o55'55MW and 0°17'33"N, 78°57,38,,W). 4 Occasional Papers, Museum of Texas Tech University Section de Mastozoologia at Pontificia Universidad which follows Weksler et al. (2006), and for bats, which Catolica del Ecuador. Specimens were treated in ac¬ follows Gardner (2007). Some taxonomic identifica¬ cordance with the guidelines of the American Society tions were further corroborated by sequencing the of Mammalogists for the use of wild mammals in cytochrome-# gene using previously published methods research (Sikes 2011). (Hanson and Bradley 2008). The specimens that were sequenced include: Akodon mollis (ACUNHC 1578, Voucher specimens from this study were com¬ 1581,1583,1585,1587,1588,1591,1595,1596,1602, pared with specimens in the collections of Abilene 1603); Thomasomys baeops (ACUNHC 1598); T. cin¬ Christian University Natural History Collection, nameus (ACUNHC 1582); T. paramorum (ACUNHC American Museum of Natural History, and Museo 1600); T. praetor (ACUNHC 1560); and T. silvestris de Zoologia, Seccion de Mastozoologia, Pontificia (ACUNHC 1592). The sequences were then compared Universidad Catolica del Ecuador to confirm species to sequences deposited on GenBank using a neighbor¬ identity. Some specimens of questionable identifica¬ joining tree (Kimura 2-parameter; Kimura 1980). tion were examined by other researchers, named in Acknowledgments, to further corroborate species A Shannon Index value (Shannon 1948) was cal¬ identifications. Nomenclature used in this study fol¬ culated for each site and compared to diversity values lows Wilson and Reeder (2005) for marsupials, shrews, from similar studies (Jarrin and Fonseca 2001; Lee et deer, and rodents, except for the Oryzomyine rodents, al. 2006a, 2006b, 2008, 2010). In total, 169 specimens of 14 species of small smaller in size (head and body 99 to 113 mm). These mammals were collected. An additional species, specimens have an antorbital vacuity that is comma- Mazama rufina, was photographed at the 3,400 m site. shaped and bounded by nasal, maxillary, and frontal Ten species were collected at the 2,962 m site and ten bones. Furthermore, the palatal bridge is curved, which species were collected or observed at the 3,400 m site is diagnostic for Caenolestes caniventer (Timm and (Table 1). No bat species were captured or observed Patterson 2007). The dorsal color is a grizzled gray¬ at the 3,400 m site. Five rodent species, Akodon mol¬ ish brown. There is no conspicuous pectoral spot as lis, Microryzomys minutus, Thomasomys baeops, T. reported by Timm and Patterson (2007). The ventral cinnameus, and T. silvestris, were documented from fur is gray to cream with a dark gray base. The con¬ both locations. Four of the species collected represent spicuous gray ears are lightly furred and have a lighter new records for Parque Nacional Sangay (Casto and colored margin. These specimens represent the first Jacome 1999). record for Morona Santiago (Tirira 2007). Previously reported locations are in western Azuay, Canar, Chim¬ borazo, Pichincha, El Oro, Loja, and Zamora Chinchipe ORDER PAUCITUBERCULATA Provinces (Anthony 1921; Bublitz 1987; Albuja and Family Caenolestidae Patterson 1996; Brown 2004; Tirira 2007). For basic Caenolestes caniventer Anthony 1921 measurements see Table 1. Gray-bellied Caenolestid Eight specimens, all males (ACUNHC 1550, ORDER SORICOMORPHA 1551,1552,1555, QCAZ 11871,11875,11876,11877), Family Soricidae were collected in Sherman traps set in a mixture of sec¬ Cryptotis montivaga (Anthony 1921) ondary growth and primary growth riparian temperate Wandering Small-eared Shrew forest at the 2,962 m site. Two specimens were caught in a trap next to a hole under a tree. Our largest adult One male (QCAZ 12037) was captured at 3,400 specimen is larger (head and body length 129 mm) than m in Chimborazo Province. The species was identified those reported by Timm and Patterson (2007). The by the presence of silver/gray pelage. This specimen rest of our specimens appear to be subadults and are was collected in habitat described by Barnett (1999) as Lee et al.—Small Mammals of Sangay National Park, Ecuador 5 Table 1. Presented are range status, collecting site, and measurements (mm) of mammal specimens collected in or near Parque Nacional Sangay, Ecuador. Range designations are as follows: R = major range records (species not previously recorded in Parque Nacional Sangay or within 70 km of the study sites); E = an elevation distribution record for the eastern Ecuadorean Andes; Known = species previously documented from Parque Nacional Sangay. Site 1 is the 3,400 m site; site 2 is the 2,962 m site. Total Tail Hind Forearm Species Range Site length length foot Ear length Caenolestes caniventer R 2 206-261 114-132 22-28 12-18 N/A Cryptotis montivaga Known 1 122 33 17 0 N/A Micronycteris megalotis E 2 75 19 11 16 33.2 Sturnira bidens Known 2 66-70 0 12-16 13-16 41.8-44 Sturnira erythromos Known 2 68 0 15 15 42 Myotis keaysi E 2 91-101 42-49 9-10 13-15 41.6-42 Akodon mollis Known 1 and 2 163-208 57-98 15-27 14-20 N/A Microryzomys altissimus Known 1 191-196 108-117 24-25 15-16 N/A Microryzomys minutus Known 1 and 2 179-205 108-130 17-25 15-16 N/A Thomasomys praetor R 1 376-432 211-249 38-42 24-25 N/A Thomasomys baeops Known 1 and 2 224-242 117-137 18-26 15-19 N/A Thomasomys cinnameus R 1 and 2 163-205 84-125 19-25 13-17 N/A Thomasomys paramo- Known 1 207-247 113-145 21-27 16-19 N/A rum Thomasomys silvestris R 1 and 2 233-288 121-166 26-34 17-25 N/A Mazama rufina Known 1 N/A N/A N/A N/A N/A quenoa forests dominated by Polylepis. Our specimen (Castro and Roman 2000; Carrera 2003; Tirira 2007, was collected at the base of a cliff with water-logged 2008), although they are found at higher elevations in soils and the ground vegetation was primarily moss other regions (Patterson et al. 1996). The pelage is and grasses. This specimen was found well within its 8-10 mm long and the body length is less than 55 mm range (Tirira 2007). (Simmons and Voss 1998). These characters separate this species from other Micronycteris. For basic mea¬ surements see Table 1. ORDER CHIROPTERA Family Phyllostomidae Micronycteris megalotis (Gray 1842) Sturnira bidens Thomas 1915 Little Big-eared Bat Andean Yellow-shouldered Bat One male (QCAZ 12030) was collected at 2,962 Five specimens, all males (ACUNHC 1565,1566, m. This specimen represents a slight elevation record QCAZ 12032, 12033, 12034), were collected at 2,962 from the previous record of2,950 m in eastern Ecuador m. This species can be distinguished by two lower inci- 6 Occasional Papers, Museum of Texas Tech University sors instead of the typical four for the genus (Giannini given by Hershkovitz (1982). This animal had a black and Barquez 2003). These specimens were found well face with a white mental patch and a white narial patch. within the documented range (Tirira 2007; Gardner The buccal patch or lower cheek was dark red. The 2007). For basic measurements see Table 1. inside of the ears are mostly black with white hair interspersed (Barrio 2010). The posterior side of the neck was black, with most of the body dark red with Sturnira erythromos (Tschudi 1844) black legs. The only other sympatric deer of similar Hairy Yellow-shouldered Bat size is Pudu mephistophiles. Pudu have a completely black face with no white in the mental or narial patches A single male (QCAZ 12036) was collected at and no red on the buccal patch (Hershkovitz 1982). 2,962 m. The specimen had a forearm of 42 mm and Both P. mephistophiles and M. rufina previously have was all gray in color with no shoulder spots. These been documented in Parque Nacional Sangay (Tirira characters indicate that it is a Sturnira erythromos 2007). (Giannini and Barquez 2003). This specimen was found well within the documented range for the species (Tirira 2007; Gardner 2007). For basic measurements ORDER RODENTIA see Table 1. Family Cricetidae Akodon mollis Thomas 1894 Soft-furred Akodont Family Vespertilionidae Myotis keaysi Allen 1914 Fifty specimens (31 males and 19 females, Hairy-legged Myotis ACUNHC 1575, 1576, 1577, 1578, 1579, 1580, 1581, 1583,1584,1585,1586,1587,1588,1591,1595,1596, One male and two females (ACUNHC 1561, 1602, 1603, 1604, 1616, 1618-1620, 1628 QCAZ QCAZ 12038, 12040) were collected from 2,962 m. 11879, 11881, 11888, 11889, 11894, 11895, 11902, These specimens represent a slight elevation record 11904, 11905, 11907, 11908, 11909, 11913, 11914, for the species in Ecuador from the recorded range 11915, 11917, 11918, 11919, 11921, 11922, 11923, of 2,950 m (Tirira 2007). They are found higher in 11924, 11925, 11926, 11927, 11928), were collected other regions (Patterson et al. 1996). The diagnostic from both sites. In the lagoon, paramo, and elfin forest characters of these specimens are that they have a habitats at the 3,400 m site we collected 25 males and sagittal crest, the upper premolar is in the tooth row, 16 females, and in the swampy, cloud forest habitats at and the distance between Cl and Cl is more than the the 2,962 m site we collected 6 males and 3 females. postorbital constriction (Moratelli and Wilson 2011). One individual had a single embryo. For basic mea¬ The two collecting sites are 4.69 km apart. surements see Table 1. Cytochrome-# was sequenced for representatives from both populations and revealed a sequence diver¬ ORDER ARTIODACTYLA gence of 1.88% (Alvarado pers. comm.). This value Family Cervidae is consistent within most intraspecific divergence and Mazama rufina (Pucheran 1851) population level variation as assessed by Baker and Ecuador Red Brocket Bradley (2006). The populations differ in coloration as well Akodon from the 3,400 m site have an overall The occurrence of Mazama rufina was docu¬ brown dorsum with each hair having black and yel¬ mented with photographs. The animal was found at low banding, and the hair on the dorsum of the feet is the 3,400 m site standing near a house owned by the white. Specimens from the 2,962 m site have a dark national park in a habitat dominated by Polylepis trees. brown (almost black) dorsum and have black hair on The animal’s height was approximately 400 mm at the the dorsum of the feet. Two females had one embryo shoulders, based on the measurements of objects near each. These specimens were found well within the where the deer was standing in the photos. Identifica¬ documented range for the species (Tirira 2007). For tion of the animal was consistent with descriptions basic measurements see Table 1. Lee et al.—Small Mammals of Sangay National Park, Ecuador 7 Microryzomys altissimus (Osgood 1933) Thomasomys cinnameus Anthony 1924 Paramo Colilargo Cinnamon-colored Thomasomys Two males (QCAZ 11929, 11930) were col¬ Seventeen specimens, 12 males and 5 females lected at the 3,400 m site. The incisor tubercle of these (ACUNHC 1564,1582,1601,1610,1611,1621,1627, specimens was short and indistinct, which separates QCAZ 11970, 11971, 11972, 11973, 11975, 11979, this species from M. minutus (Carleton and Musser 11981, 11982, 11983), were collected from both the 1989). These specimens were found well within the 3,400 m and 2,962 m sites. The morphological iden¬ documented range (Carleton and Musser 1989; Tirira tification was confirmed by sequencing cytochrome-# 2007). For basic measurements see Table 1. (Hanson and Bradley 2008). These specimens repre¬ sent the first record for Chimborazo and Morona San¬ Microryzomys minutus (Tomes 1860) tiago Provinces. Previous records are from Pichincha, Tungurahua, and Carchi Provinces (Voss 2003; Tirira Montane Colilargo 2007). These specimens were collected in a variety of habitats, including elfin forest, bogs, and swampy, cloud Six specimens, five males and one female forest. For basic measurements see Table 1. (ACUNHC 1605, 1556, QCAZ 11932, 11933, 11934, 11936), were collected at both the 3,400 m site and the 2,962 m site. These specimens can be distinguished Thomasomysparamorum Thomas 1898 from M. altissimus by a tail length of >145% of head Paramo Thomasomys and body length and by a maxillary toothrow length of <3 mm (Carleton and Musser 1989). These speci¬ Forty-four specimens, 22 males and 22 females mens were found well within the documented range (ACUNHC 1549,1558,1568,1569,1574,1589,1590, (Carleton and Musser 1989; Tirira 2007). For basic 1593,1597,1599,1600,1606,1607,1608,1612,1613, measurements see Table 1. 1615,1622-1626, QCAZ 11995,12002,12004,12006, 12007, 12008, 12009, 12010, 12011, 12013, 12014, 12015, 12016, 12017, 12018, 12019, 12020, 12021, Thomasomys baeops Osgood 1914 12022, 12023, 12024, 12026, 12027, 12028), were Long-tailed Thomasomys collected from the 3,400 m site. All specimens had large inflated auditory bullae, which distinguishes them Nine specimens, six males and three females from other sympatric Thomasomys (Voss 2003). Some (ACUNHC 1570, 1594, 1598, 1609, 1614, QCAZ of these specimens were larger than the measurement 11945,11946,11947,11948), were collected from both range presented by Voss (2003). For example, speci¬ the 3,400 m and 2,962 m sites. The identification was mens in this study had tail lengths of up to 145 mm and confirmed by Robert Voss and characters presented by hind foot lengths of up to 27 mm, whereas Voss (2003) Voss (2003). These characters include: length of hind reported 133 mm for maximum tail length and 25 mm foot < 25 mm; auditory bullae small and uninflated; and for maximum hind foot length. Most of our specimens maxillary molar row >4.2 mm (Voss 2003). However, were collected in Polylepis forests near the lagoons analysis of cytochrome-# indicates that one specimen and paramo, similar to the habitat that Voss (2003) and (ACUNHC 1598) identified morphologically as T. Stone (1914) describe for this species. These specimens baeops would correspond to another unknown taxa were found well within the documented range (Tirira (Fig. 2). Additional investigation is needed to resolve 2007). For basic measurements see Table 1. this dilemma. The specimens were found well within the range of T. baeops in Ecuador (Tirira 2007). One female had three embryos. For basic measurements see Table 1. 8 Occasional Papers, Museum of Texas Tech University AY041205 DQ014654 baeops ACUNHC1355 ischyrus AF108675 ACUNHC1592 silvestris DQ914650 australis DQ914645 AF108673 daphne DQ914649 U03540 aureus DQ914653 T. sp. ACUNHC1341 ACUNHC1340 fumeus ACUNHC1339 AF108678 T. sp. AF108676 notatus DQ914652 ladewi DQ914647 DG914648 caudivarius ACUNHC1137 erro AF108674 gracilis DG914646 cinnameus ACUNHC1582 ACUNHC1598 baeops’' ACUNHC1600 QCAZ11704 paramorum QCAZ11705 QCAZ11706 DG914651 oreas AF108677 DG914643 andersoni DG914644 ACUNHC1560 praetori MVZ182704 R. auritus 0.02 Figure 2. Kimura 2-parameter neighbor-joining tree of Thomasomys species based on cytochrome-6 gene sequences. Abilene Christian University Natural History Collections (ACUNHC) and Seccion de Mastozoologia, Museo de Zoologia, PontificaUniversidad Catolica del Ecuador (QCAZ) numbers are listed for specimens collected in Ecuador. Other specimens are listed by their GenBank numbers. Thomasomys species names are listed to the right of the figure. Samples listed from Sangay are in bold, and the Thomasomys baeops from Sangay (ACUNHC 1598) is labeled with an asterisk (*) and shown sister to a clade containing Thomasomys paramorum. The outgroup is a Reithrodon auritus deposited at the Museum of Vertebrate Zoology (MVZ). Lee et al.—Small Mammals of Sangay National Park, Ecuador 9 Thomasomys praetor (Thomas 1900) lected from Thomasomys aureus nicefori in Antioquia, Cajamarca Thomasomys Colombia (Seevers 1955). The species identification must be considered tentative as the males possess the Four specimens, two males and two females most diagnostic characters in Amblyopinus (M.S. En¬ (ACUNHC 1548, 1560, QCAZ 11937, 11938), were gel, pers. comm.). If confirmed, this would be the first collected from the 3,400 m site. These specimens were record of A. colombiae from Ecuador. caught near cliff faces and near the paramo temperate forest ecotone. While Cabrera (1961) indicated that T. Thomasomys silvestris Anthony 1924 praetor is conspecific with T. aureus, Voss (2003) sug¬ Sylvan Thomasomys gested that 4 or possibly 5 distinct species, including T. aureus and T. praetor, may represent this complex. Nineteen specimens, nine males and ten females Although the fur color of our specimens does not match (ACUNHC 1554,1557,1562,1563,1567,1572,1573, that presented in Voss (2003) for T. praetor, sequencing 1592,1617, QCAZ 11952,11955-11957,11960,11961, of the cytochrome-# gene confirmed that our specimens 11963, 11964, 11965, 11966, 11967), were collected are T. praetor and not the morphologically similar T. au¬ from both the 3,400 m and 2,962 m sites. Our speci¬ reus. Interestingly, the neighbor-joining tree resulting mens are similar in color to those reported by Anthony from the cytochrome-# data indicates that T. aureus and (1924a). The dorsal hairs are slate gray but tipped with T. praetor are not sister taxa (Fig. 2). Clearly, further dark brown, giving an overall brown appearance to the investigation is warranted to resolve the taxonomy of dorsum. The ventral fur color is warm light brown at the species within this complex. These specimens con¬ the tip with a bluish gray shaft and base. Most of our firm the presence of T. praetor in Ecuador (Musser and specimens have a patch of light brown behind the ears. Carleton 2005). For basic measurements see Table 1. These specimens represent the most southern record for Ecuador and the first record for Chimborazo and Two female rove beetles (Coleoptera: Staphylini- Morona Santiago Provinces. Previous records were dae) were discovered on the neck of voucher specimen from Pichincha and Napo Provinces (Pacheco 2003; QCAZ 11938. The beetles were tentatively identified Voss 2003). For basic measurements see Table 1. by Michael S. Engel (University of Kansas) as Am- blyopinus colombiae Seevers 1955, a species first col¬ Discussion The mammalian species diversity and richness ness with increasing elevation (Patterson 1996; Lee et documented for Parque Nacional Sangay can be com¬ al. 2006a, 2006b, 2008, 2010) (Table 2). By taxon, pared with the results of five previously published bat diversity clearly follows this pattern and probably studies in Ecuador (Jarrin and Fonseca 2001; Lee et accounts for the trend in the overall mammalian di¬ al. 2006a, 2006b, 2008, 2010) (Table 2). Shannon versity. This study resulted in the lowest diversity of values for each study, presented in Table 2, are use¬ bats (four species) compared with the other published ful for comparing samples where a common species studies and reflects this trend of low bat species diver¬ could cause species richness values to be misleading sity and richness with increasing elevation (Patterson and for comparing the evenness of diversity (Bacaro et al. 1996) (Table 2). The numbers of individuals of et al. 2011). Identification of these patterns in species bats also were very low compared with other surveys. diversity is important in conservation efforts (Bacaro However, some taxonomic groups, such as the cricetid etal. 2011). rodents, do not show a pattern of decreasing diversity with increasing elevation. For example, in comparing A comparison of overall species diversity be¬ the results from Santa Rosa (lowest elevation 450 m) tween this study and previously published data shows and Parque Nacional Sangay (highest elevation 3,400 a strong trend of decreasing species diversity and rich¬ m), eight species of rodents were collected at each site 10 Occasional Papers, Museum of Texas Tech University Table 2. Comparison of Chiropteran and overall mammalian species composition between this study and previously published studies in Ecuador. Number for Number of all species of Hp for all Location Elevation bat species Hp for bats mammals mammals Imbabura (Lee et al. 450 m 17 1.10 23 1.23 2010) Imbabura (Lee et al. 702 m 10 0.86 15 1.03 2010) Imbabura (Lee et al. 450-702 m 22 1.19 32 1.31 2010) Otonga (Jarrin and 1,300-2,300 m 18 1.02 N/A N/A Fonseca 2001) Tandayapa (Lee et al. 1,500-2,100 m 13 0.88 19 0.99 2006b) Guajalito (Jarrin and 1,800-2,000 m 16 1.03 N/A N/A Fonseca 2001) Cosanga (Lee et al. 1,900-2,100 m 15 0.82 20 0.91 2006c) Volcan Sumaco 2,500 m 8 0.58 12 0.84 (Lee et al. 2008) Sangay National Park 2,962 m 4 0.50 10 0.84 (this study) Sangay National Park 3,400 m 0 N/A 10 0.75 (this study) Sangay National Park 2,962-3,400 m 4 0.50 15 0.88 (this study) (Lee et al. 2010). However, only four species at Santa with our study (Haynie et al. 2006). In contrast, at the Rosa were cricetids (Lee et al. 2010), whereas all eight geographically more distant Papallacta, but at an eleva¬ species at Parque Nacional Sangay were cricetids. tion similar to our study sites, Voss (2003) recorded Therefore, as elevation increases there are fewer fly¬ seven species in common with our study. The differ¬ ing insects and fewer trees with fruit suitable for bats, ence in elevation between our study and Haynie et al. wheras the paramo is dominated by grasses which (2006) is perhaps the reason for the dissimilar species granivore rodents can use to their advantage. Our data composition between the two study areas despite their do show a peak in diversity in a mid-elevation range geographic proximity. for cricetids, which also was found among non-volant mammals in a study by McCain (2005). Cytochrome-# sequence analysis indicates that a specimen identified morphologically as T. baeops (by In 2001, a study was conducted near the northern many researchers) from this study was not most similar border of Parque Nacional Sangay at a lower eleva¬ to other previously identified T. baeops (Fig. 2), sug¬ tion than this study (1,700 m to 2,700 m) (Haynie et gesting that T. baeops may not be monophyletic. Our al 2006). In that study, 15 species were collected, but results suggest the need for a more thoroughly sampled only three taxa (Akodon mollis, Microryzomys minutus, phylogenetic analysis of the genus Thomasomys. and Thomasomysparamorum) were found in common Furthermore, cytochrome-# analysis confirmed the