Redescription of Mimon koepckeae (Chiroptera: Phyllostomidae)

http://dx.doi.org/10.1590/s1984-46702014000400009 Redescription of Mimon koepckeae (Chiroptera: Phyllostomidae) Natalí Hurtado 1,2,4, Edith Arias 1 & Víctor Pacheco 1,3 1 Departamento de Mastozoología, Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos. Av. Arenales 1256, Lima 14, Lima, Peru. 2 Doctorado en Cs. Mención Ecología y Evolución, Facultad de Ciencias, Universidad Austral de Chile. 3 Instituto de Ciencias Biológicas Antonio Raimondi, Facultad de Ciencias Biológicas, Universidad Nacional Mayor de San Marcos. 4 Corresponding author. E-mail: natalihm@gmail.com ABSTRACT. Mimon koepckeae Gardner & Patton, 1972 is a poorly-known bat species, with only three known specimens, including the holotype. Its distribution is restricted to the type locality in Ayacucho Department, Peru, and surroundings. This species has been synonymized with M. crenulatum by some authors. Based on a new specimen of M. koepckeae collected from Santuario Nacional Pampa Hermosa, Junin Department, Peru, we provide an extensive morphological comparison with M. crenulatum (Geoffroy St.-Hilaire, 1803), Mimom bennettii (Gray, 1838), and Mimon cozumelae Goldman, 1914, concluding that M. koepckeae is a valid species. As a result the distribution range of the species is extended 160 km north of the type locality. In addition, we characterize the habitat of the species, provide current data on feeding behavior, and suggest that M. koepckeae should be categorized as endangered species. KEY WORDS. Conservation; endemic; Junin; Pampa Hermosa; Yungas. Mimon koepckeae Gardner & Patton, 1972 is an endemic Peruvian bat whose distribution used to be restricted to the type locality in Ayacucho Department, Peru, and surroundings (SIMMONS 2005, WILLIAMS & GENOWAYS 2008, VELAZCO & AGUIRRE 2008). This species is known from three specimens (two females and one male), which were captured in the localities of Estera Ruana and Huanhuachayo, between 1,600 and 1,900 m (GARDNER & PATTON 1972). Natural history information about M. koepckeae is scarce, and the description of its habitat offers few details, for instance a reference to cloud forests of the eastern slope of the Peruvian Andes (GARDNER & O NEILL 1971, GARDNER & CARTER 1972, GARDNER & PATTON 1972). Mimon koepckeae has a controversial taxonomic history. GARDNER & PATTON (1972) described the species based on the absence of a dorsal stripe, fewer crenulations and scarce hairs on the nose-leaf basis, narrow auditory bullae, and a well defined cleft between protocone and hypocone in the first and second upper molars. However, KOOPMAN (1976, 1978) considered M. koepckeae as a junior synonym of Mimon crenulatum (Geoffroy St.-Hilaire, 1803), based on a morphological comparison of skins and skulls of a single specimen of M. koepckeae, collected by Terborgh in 1972 from Estera Ruana (one locality of the type series), and a series of 10 specimens of M. crenulatum. Koopman (1978) considered the characters listed by GARDNER & PATTON (1972) for M. crenulatum as corresponding to intraspecific variation. Since that revision, KOOPMAN (1993, 1994) kept M. koepckeae as a highland Peruvian sub-species of M. crenulatum within the sub-genus Anthorhina. Later, SIMMONS & VOSS (1998) and SIMMONS (2005) recognized the validity of M. koepckeae, without supporting their conclusions. Because notorious changes in habitats due to anthropocentric activities, M. koepckeae is listed as Critically Endangered by the Peruvian legislation (MINISTERIO DE AGRICULTURA 2014), contrasting with its classification as Data Deficient by the IUCN (VELAZCO & AGUIRRE 2008, IUCN 2012a). Herein, we report the rediscovery of M. koepckeae 40 years after its description, and present a complete morphological characterization of it, including new diagnostic characters supporting its status of valid species. Based on our results, the distribution of the species is significantly broadened. In addition, the habitat of the species, based on analysis of the new collecting locality, is characterized. Finally, we also discuss the nomenclatural status of the sub-genera of Mimon, based on comparisons with other species of the genus (sensu SIMMONS 2005). MATERIAL AND METHODS Bat inventories were conducted in the locality of Podocarpus, Santuario Nacional Pampa Hermosa, Chanchamayo District, Chanchamayo Province, in Junin Department of Peru (Fig. 1, 10 59 49.2"S, 75 25 57.1"W, 1890 m). Ten mist nests (12 x 2.5 m each one) were set up for seven consecutive nights, with a total sampling of 70 mist nests per hour. The morphological characterization of Mimon koepckeae presented here is based on the specimen recorded herein (Appendix 1) and one topotype (AMNH 23222), which were con- 2014 Sociedade Brasileira de Zoologia www.sbzoologia.org.br www.scielo.br/zool All content of the journal, except where identified, is licensed under a Creative Commons attribution-type BY-NC.

378 N. Hurtado et al. trasted with photos of the type specimen (LSUMZ 15676), and those in the original description (GARDNER & PATTON 1972). The age class was defined following PACHECO & PATTERSON (1992). For morphological characters we followed LEGENDRE (1984), PACHECO & PATTERSON (1992), VELAZCO (2005), WETTERER et al. (2000), PACHECO et al. (2004), GIANNINI & SIMMONS (2007), and FRACASSO et al. (2011). Colors were defined following RIDGWAY (1912). We also followed VELAZCO (2005: 20) for morphometrics. Mimon koepckeae was compared with the other three species of Mimon (sensu SIMMONS 2005): M. crenulatum (É. Geoffroy St.- Hilaire, 1803) (n = 181), M. cozumelae Goldman, 1914 (n = 31), and M. bennettii (Gray, 1838) (n = 5). For diet analysis, we placed a graph paper under a petri dish and analyzed stomach contents and feces in five randomly selected areas (5 x 5 mm) using a stereomicroscope with 20x magnification. TAXONOMY An adult female of Mimon koepckeae was collected from the locality of Podocarpus in the Santuario Nacional Pampa Hermosa, Province of Chanchamayo, Department of Junin, Peru (Fig. 1). The specimen was captured by Edith Arias (EA 216) on October 6 th, 2011. Its external measurements are: total length 78 mm, tail length 23 mm, hindfoot length 11 mm, ear length 22 mm, tragus length 9 mm, forearm length 48 mm, and weight 14.5 g. This specimen was preserved as skin, with the skull removed, and was deposited in the mammal collection of the Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, under catalogue number MUSM 41327 (Figs 2-6). This specimen was identified as Mimon koepckeae, based on the original species description (GARDNER & PATTON 1972). Mimon koepckeae Gardner & Patton, 1972 Figure 1. Records of Mimon koepckeae: (triangle) Huanhuachayo, Ayacucho (type locality) 12 44 00"S, 73 47 00"W and (Circle) Podocarpus, Junin, 10 59 49.2"S, 75 25 57.1"W. Type locality: Huanhuachayo (12 44 00"S, 73 47 00"W), elevation 1,660 m, Ayacucho Department, Peru. Revised diagnosis. Medium-sized bats; pelage color from reddish brown to golden brown; dorsal stripe absent; noseleaf slender, crenulated along proximal margin, and sparsely fringed with short fine hairs; skull small, with narrow rostrum; auditory bullae narrow; first and second upper molars with narrow, well-defined vertical cleft separating protocone and hypocone. Redescription. Size. Medium-sized bats. Forearm length between 46.9-50.2 mm; weight 14 g. Pelage. Dorsal fur at rump level short (6 mm); bicolored banding pattern is Pale Smoke Gray at base and Chestnut at tips. Ventral fur at belly level slightly shorter (5 mm); tricolored banding pattern with small Pale Smoke Gray at base, large Hair Brown on central portion, and Olive-Buff tips that turn whitish from central abdomen toward flanks. Pale Smoke Gray basal portion on head larger dorsal and ventrally. Hairs of auricular patch exhibit larger whitish portion and Chestnut tips. Dorsal stripe absent. Head. Comparatively short rostrum, about one third head length or less, mandibular pragmatism not present. Noseleaf limb-horseshoe continuous. Limb ovate, length about twice width. Limb margins pigmented, shallow crenulations at base disappear apically; tuft of long hairs at tip present. Limb-rib comparatively thin and slimmer than internostrils space. Horseshoe continuous around nostrils. Anterior noseleaf margin well developed and free. Lips continuous and flat. Four vibrissae placed on each dorsal lip margin; and six vibrissae on each ventral margin. Chin with two pairs of central pads; one pair placed below lip, the other, smaller, behind it; four dermal pad pairs placed diagonally to central ones; dermal pads pigmented and symmetrically separated by central shallow cleft, which does not extend behind chin. A set of seven vibrissae on each side of horseshoe; above, a set of swollen continuous papillae. Ears slightly smaller than head length. Pinna delta shaped, fleshy and not translucent, with numerous marked folds, rounded and parallel at tips and edges continuous. Ears bicolored; external half pigmented and basal half (including tragus) pale. Pinna poorly developed, inserted the head insertion level. Inner ear lobe well developed; with long dense hairs, and well-

Redescription of Mimon koepckeae 379 2 3 Figures 2-3. External appearance of Mimon koepckeae (MUSM 41327): (2) dorsal view; (3) ventral view. Scale bar: 3 cm. 4 5 Figures 4-6. (4) Dorsal, (5) ventral, and (6) lateral views of the skull of Mimon koepckeae (MUSM 41327). Scale bar: 1 cm. 6 developed and sparse protuberances on internal surface. Tragus long, about one third ear length. Tragus rib thick with scarce long whitish hairs. Body. Single odor gland placed at center of throat. Forearm slim, fragile, naked; length average 47.7 mm. Fifth phalange larger than third, and fourth larger than fifth. Plagiopatagium inserted at metatarsal level on external border of fifth digit. Dactilopatagium major exhibits V shaped slot. Medium and minus dactilopatagium exhibit complete pigmentation. Dactilopatagium brevis inserted at middle of digit II. Uropatagium wide and long, projected beyond toenail edge. External edge of uropatagium with fringe of scarce and short hairs. Hind feet completely pigmented with dense, long black hairs on dorsal surface. Calcars longer than tibia. Tip of tail free. Skull. Rostrum short, length average 21.11 mm. In dorsal view, incisor roots form V shaped groove. Anterior edge of nostril V shaped. Maxillary roots of zygoma swollen. Interorbital region narrower than width between-canines. Braincase edge V shaped. In lateral view, a short terrace on the pre-maxilla exhibited at incisors level. Forehead plain without medial depression. Zygomatic arches flat and thick, with hard and broad roots. Jugal bone narrower. Sagittal crest wide at frontal level and tapering towards posterior side. In ventral view, pal-

380 N. Hurtado et al. 7 8 9 10 11 12 13 14 Figures 7-14. Dorsal and ventral views of the skull of four species of Mimon: (7-8) M. koepckeae (MUSM 41327); (9-10) M. crenulatum (MUSM 24723); (11-12) M. cozumelae (FMNH 58151); (13-14) M. bennettii (NMNH 391027). Scale bar: 1 cm. ate short and wide, with concave floor. Pre-maxilla triangular, placed below maxillary level; posterior edge exhibits small rounded projection towards maxillary. Lateral posterior edges of palate wide, extending towards middle of M2 talonid. Postpalatal process well-marked. Hard palate edge V shaped; anterior border extended beyond middle of interpterygoid bone. Medial basisphenoid sharped septum forms basisphenoidal pits. Basioccipital flat and narrow. Basioccipital-basisphenoid suture perpendicular to basisphenoid septum. Anterior border of foramen magnum shallow. Jugular foramen the same size as jugal breath. Occipital condyles short and narrow. Para-occipital processes developed, but shorter than occipital condyles. Auditory bullae laterally compressed, wider than diameter of foramen magnum, with few swollen optic rings. Dental morphology. Dental formula I 2/1, C 1/1, P 2/2, M 3/3. Upper. Inner incisors rhomboid shaped, half canine length. Tips and almost all inner borders free. Outer incisors flat and shorter than inner ones. Canines pyramidal shaped; well-defined medial notch on lingual cingulum. Second premolar (P2) reduced, conical, labially placed and one sixth canine length; in ventral view, labial and lingual cingula subequal. Fourth premolar (P4) developed and conical, average size between canines and molars; P4 exhibits well developed main cusp; labial cingulum narrow, extended towards talon; talon basin wide and deep; secondary cusp absent; on posterior side, metastyle below parastyle of first molar. Parastyle high of first molar (M1), high, but not reaching paracone level; mesostyle shorter than parastyle; metacone higher than paracone; preparacrista,

Redescription of Mimon koepckeae 381 15 16 17 18 Figures 15-18. Lateral view of skull and mandible of four species of Mimon: (15) M. koepckeae (MUSM 41327); (16) M. crenulatum (MUSM 24723); (17) M. cozumelae (FMNH 58151); (18) M. bennettii (NMNH 391027). Scale bar: 1 cm. postparacrista, premetacrista and postmetacrista with concave edges; stylar shelves triangular; anterior ectoflexus convex; posterior ectoflexus triangular; protofossa sub equal to anterior stylar shelf; protocone developed and pyramidal; hypocone less developed; vertical sulcus between protocone and hypocone is present; labial cingulum well developed; metacingulum poorly developed and not in contact with M2; metastyle not in contact with parastyle of M2. Second molar (M2) similar to M1, but paracone and metaestyle of the third molar (M3) below level of parastyle; ectoflexus less developed; stylar shelf flat; protocone less developed than in M1 and M2. Lower. Single pair of incisors (i1) present, flat and bilobed, with diagonal lateral surface wear. Canines developed and conical; labial cingulum narrow; mesial cingulum ends in small conule; parastyle and distal cingulum well developed. Second premolar (p2) short, about half size of canine, with single principal cone, asymmetrically divided. Fourth premolar (p4) larger than p2, with single principal pyramidal cone, labial cingulum restricted to first half. First molar (m1) longer than wider; size average between canines and p4; paraconid, protoconid and metaconid linked by vertical crests with concave edges, forming deep triangle basin; paraconid below p2 level; protoconid stands out as the largest cuspid; and paraconid and entoconid the smallest; hypoconulid placed posteriorly to entoconulid with a small peak; on posterior side, hypoconulid does not overlap m2. Second molar (m2) similar to m1, but oblique crest exhibits convex edge; labial cingulum exhibits deep cleft between trigonid and talonid; hypoconulid larger than in m1. Paraconid, metaconid and protoconid of third molar (m3) subequal, connected by vertical crests with concave edges shallower than m1 and m2; talonid less developed and shorter than hypoconulid; labial cingulum as m2; labial cingulum developed at trigonid level, but fewer at talonid level. Mandible. In dorsal view, condylar process breadth is larger than the coronoid process height; coronoid processes sloped laterally; angular processes shorter than condylar processes. In lateral view, chin straight with well develop sub-mental process; posterior edge diagonal with angle less than 30 ; toothrow short; coronoid process short; condylar process thick and twice larger than angular process, placed over diastema, diastema between toothrow and coronoid process; notch between angular and condylar processes trapezoidal.

382 N. Hurtado et al. Distribution. Mimon koepckeae is known to occur only in the type locality (Huanhuachayo) and vicinities of Estera Ruana; in Ayacucho Department, and low cloud forest of Junin Department, elevation between 1660 and 1890 m. Remarks. Mimon koepckeae was compared with specimens representing the three recognized species in the genus sensu SIMMONS (2005): M. crenulatum, M. cozumelae, and M. bennettii. Compared with Mimon koepckeae, M. crenulatum is slightly larger, average body length is 84.09 mm, dorsal fur is unicolored Deep Mouse Gray and shorter, the dorsal stripe is always present, ventral fur is tricolored, auricular patch is bright and conspicuous, noseleaf crenulations and hairs are evenly distributed, ears are shorter, calcar is slightly shorter, skull length is slightly larger, nasal depression is U shaped, incisors gum crests are less swollen, medial depression is present, maxillary root of zygoma is less swollen, interorbital region is wider, posterior braincase projection is more developed, palate is large and narrow, an anterior accessory medial foramen is present in some populations, lateral palatal edges are narrower, posterior border of the hard palate is U-shaped and does not reach the middle of mesopterygoid fossae, auditory bullae are wider, basioccipital is wider at cochlea level, cleft between protocone and hypocone is absent in M1 and M2, accessory cuspids of lower canines are less developed, coronoid process is over diastema level, posterior edge of mandible is heel shaped. Mimon bennettii is larger, body length is 92 mm, dorsal fur is tricolored and longer, ventral fur is unicolored, noseleaf is short and wide, noseleaf crenulations and hairs are absent, a free skin binding of the horseshow is absent, chin exhibits dense and long hairs, ears are longer, translucent with marked folds, ear tips are divergent; forearm is furred, calcar is shorter, plagiopatagium is inserted at tibia level. Skull length is larger, anterior nasal edges are flat, incisors gum crests are flat, maxillary root of zygoma is flat, interorbital region is comparatively narrower, palate is large and narrow, premaxillary and maxillary are at same level on the palate, lateral palatal edges are narrower, posterior border of the hard palate is U-shaped and does not reach the middle of mesopterygoid fossae, auditory bullae are shorter and narrower, basioccipital is wider at cochlea level, cleft between protocone and hypocone is absent in M1 and M2, accessory cuspids of lower canine are flatted, coronoid process is over diastema level, posterior edge of mandible is flat, secondary cuspid of P4 is present. Mimon cozumelae is larger, average body length is 94.5 mm, dorsal fur is tricolored and longer, ventral fur is unicolored, noseleaf is short and wide, noseleaf crenulations and hairs are absent, a free skin binding of the horseshow is absent, chin exhibits dense and long hairs; ears are longer, translucent with marked folds, ear tips are divergent; forearm is furred, calcar is shorter, plagiopatagium is inserted at tibia level, skull length is larger, anterior nasal edges are flat and slightly projected, incisors gum crests are flat, maxillary root of zygoma is flat, interorbital region is comparatively narrow, palate is large and narrow, premaxillary and maxillary are at same level on the palate, a accessory medial foramen is exhibited behind the incisive foramina, lateral palatal edges are narrower, posterior border of the hard palate does not reach the middle of mesopterygoid fossae, auditory bullae are shorter and narrower, basioccipital is wider at cochlea level, a cleft between protocone and hypocone is absent in M1 and M2, accessory cuspids of lower canine are absent, coronoid process is over diastema level, posterior edge of mandible is flat, secondary cuspid of P4 is present. Table I provides external and cranial measurements for each species. In addition, Table II provides a summary of external and skull-dental characters comparisons among species. Natural History. Podocarpus is located 160.1 km northward from the type locality, Huanhuachayo. The habitat in Podocarpus is dominated by tree species of Podocarpaceae and Clusiaceae, which are approximately of 15 m high (Fig. 19). The understory is sparse and covered with Rubiaceae and Araceae shrubs (Fig. 19, LA TORRE-CUADROS et al. 2007). According to TOSI (1960) and HOLDRIDGE (1967), this ecosystem is categorized as Very Humid low Montane forest, also known as Yungas (TOVAR-NARVÁEZ et al. 2010); it is a steep landscape forming an altitudinal belt from 1,600 to 2,800 m elevation. Figure 19. Cloud forest habitat of Podocarpus where Mimon koepckeae was collected in this work. Photo by Edith Arias. Mimon koepckeae was collected in a natural canopy opening during the early rainy season, with Anoura aequatoris (Lönnberg, 1921), Carollia brevicauda (Schinz, 1821), Micronycteris megalotis (Gray, 1842), Myotis keaysi J.A. Allen, 1914, Myotis riparius Handley, 1960, Sturnira magna de la Torre, 1966, and Vampyressa melissa Thomas, 1926. Details for this bat assemblage will be reported elsewhere. The reproductive status of M. koepckeae corresponds to an adult female with perforated genitalia, which indicates recent

Redescription of Mimon koepckeae 383 Table I. External and cranial measurements ( in millimeters) of Mimon bennettii, M. cozumelae, M. crenulatum, and M. koepckeae. Standard deviation is in parenthesis, following by sample number. Characters M. bennettii M. cozumelae M. crenulatum M. koepckeae Total length 92.00(± 0.00)2 94.50(± 2.60)9 84.09(± 3.80)95 79.00(± 1.41)2 Tail length 20.50(± 0.71)2 20.61(± 2.76)9 23.26(± 2.54)96 19.50(± 4.95)2 Hindfoot length 15.00(± 0.00)2 16.83(± 0.90)9 11.57(± 1.30)96 10.00(± 1.41)2 Tragus length * * 10.22(± 1.00)96 9.50(± 0.71)2 Ear length 36.50(± 0.71)2 35.00(± 1.87)9 24.03(± 1.57)96 22.50(± 0.71)2 Forearm length 59.50(± 0.71)2 57.43(± 1.11)9 48.02(± 3.39)96 47.70(± 0.42)2 Greatest length of skull 25.49(± 0.22)3 25.33(± 0.41)27 21.50(± 0.68)92 21.11(± 0.74)3 Condyle-incisors length 22.61(± 0.35)3 22.67(± 0.41)26 19.09(± 0.61)91 18.79(± 0.21)3 Skull height 11.07(± 0.34)3 11.16(± 0.42)27 9.39(± 0.44)91 9.28(± 0.22)2 Postorbital breadth 4.75(± 0.03)3 4.74(± 0.16)27 4.26(± 0.16)91 4.14(± 0.06)3 Zygomatic breadth 14.02(± 0.15)3 13.82(± 0.33)25 12.13(± 0.46)90 11.67(± 0.24)3 Braincase breadth 9.99(± 0.13)3 10.03(± 0.26)27 8.71(± 0.38)91 8.41(± 0.06)2 Palatal width at canines 5.94(± 0.30)3 5.84(± 0.17)27 5.29(± 0.22)92 5.01(± 0.01)3 Mastoid breadth 11.12(± 0.15)3 10.75(± 0.37)27 9.70(± 0.35)92 9.92(± 0.86)3 Palatal length 11.42(± 1.49)3 11.89(± 0.34)27 9.70(± 0.40)92 8.44(± 0.33)3 Breath across maxilla 4.32(± 0.14)3 4.08(± 0.13)27 3.61(± 0.19)92 3.76(± 0.23)2 Width at M1-M1 9.17(± 0.20)3 8.78(± 0.28)27 7.59(± 0.28)92 7.43(± 0.18)2 Width at M2-M2 9.47(± 0.24)3 9.20(± 0.27)27 8.26(± 0.30)92 8.04(± 0.19)3 Auditory bullas breadth 2.04(± 0.27)3 2.00(± 0.15)26 2.75(± 0.14)91 2.36(± 0.19)2 Maxillary toothrow length 9.10(± 0.88)3 9.44(± 0.25)27 7.88(± 0.23)92 7.71(± 0.36)3 Molariform toothrow length 8.17(± 0.12)3 7.99(± 0.25)27 6.58(± 0.24)92 6.64(± 0.74)3 Condylar breadth 6.20(± 0.12)3 6.52(± 0.23)26 5.39(± 0.32)77 5.58(± 0.15)2 Coronoid height 6.26(± 0.29)3 6.34(± 0.19)27 4.39(± 0.25)91 4.46(± 0.17)2 Dentary length 16.57(± 0.52)2 16.48(± 0.32)27 13.89(± 0.50)91 13.47(± 0.38)3 Mandibular toothrow length 10.79(± 0.44)3 10.72(± 0.34)27 8.94(± 0.27)91 8.69(± 0.06)2 Mandibular height 2.14(± 0.10)3 2.18(± 0.18)27 1.87(± 0.16)91 1.73(± 0.04)2 *Non-available on labels. sexual activity. Stomach content was scarce and unidentifiable, but fecal analysis revealed remains (i.e., elytra, wings, legs, and antennas) of Elateridae (30%) and Scarabaeidae (70%) beetles. DISCUSSION We recognize Mimon koepckeae as a valid species, based on distinctive morphological attributes presented in this work, and thus refuting KOOPMAN s (1976, 1978) arguments. The diagnostic characters of this species are independent, and are not contained within the range of morphological variability of M. crenulatum (i.e. nasal shape, dorsal stripe absent). This species is distributed in low Montane Forests from Oriental slope of Central Andes in Peru, but is not sympatric with M. crenulatum (GARDNER & PATTON 1972, this work), which ranges at lower elevations (300 to 900 m). This new record of Mimon koepckeae is a rediscovery, because this report is more than 40 years after the type series description (see criteria of SCHEFFERS et al. 2011). It is also a distribution range extension of 160 km northward from the type locality and surroundings. The scarcity of records of M. koepckeae reflects the lack of intense fieldwork in the area or not enough sampling, which are largely due to difficulties accessing that zone. Mimon koepckeae and M. crenulatum were placed in the subgenus Anthorhina (GARDNER & PATTON 1972, SIMMONS & VOSS 1998). However, the taxonomic status of Anthorhina has been discussed, since HANDLEY (1960) synonymized it with Mimon. Although HUSSON (1962) considered Anthorhina as a genus, based on differences in the relative size of the upper premolars, height of bullae, and shape and nose-leaf pubescence, CABRERA (1958) and GOODWIN & GREENHALL (1961) considered Anthorhina only as a subgenus of Mimon. The characters of M. koepckeae and M. crenulatum, described here differ from the morphology of M. bennettii and M. cozumleae, and also from the generic description of Mimon provided by GRAY (1847), supporting recent phy-

384 N. Hurtado et al. Table II. External and cranial characters of Mimon bennettii, M. cozumelae, M. crenulatum, and M. koepckeae. Characters M. bennettii M. cozumelae M. crenulatum M. koepckeae Dorsal fur Tricolored, dorsal stripe always absent Tricolored, dorsal stripe always absent Bicolored, dorsal stripe always present Length of dorsal fur 10 mm 9 mm ~ 6 mm ~ 7 mm Unicolored, dorsal stripe always absent Ventral fur Unicolored Unicolored Tricolored Bicolored Auricular patch Whitish and inconspicuous Whitish and inconspicuous Yellowish and conspicuous Whitish and inconspicuous Length of noseleaf Less than twice width Less than twice width Length more than twice width Length twice width Noseleaf edges Crenulations and hairs absent Crenulations and hairs absent Lot of crenulations and long hairs Few crenulations and scarce short hairs at the base Anterior edge of horseshoe Anterior projection absent Anterior projection absent Free skin binding Free skin binding Chin appearance Ornamented Ornamented Nude Nude Ears length and appearance Divergent tips, large, translucent, with well-defined folds Divergent tips, large, translucent, with well-defined folds Parallel tips, smaller, fleshy, with inconspicuous folds Forearm appearance Dense long hairs on first third Dense long hairs on first third Without hairs Without hairs Parallel tips, small, fleshy, with inconspicuous folds Calcar length Short, lesser than one half tibia length Short, lesser than one half tibia length Large, close to tibia length Large, larger than tibia length Plagiopatagium insertion At tibia level, close to ankle At tibia level, close to ankle At tarsus level, on V metatarsal At tarsus level, on V metatarsal Skull length > 25 mm > 25 mm < 23 mm < 23 mm Shape of nasal depression Flat Flat and projected U-shaped V-shaped Shape of incisors gums Flat Flat Swollen Swollen Forehead appearance Without medial depression Without medial depression With medial depression Without medial depression Shape of maxillary roots of zygoma Flat Flat Swollen More swollen Shape of posterior braincase edge With a little projection With a little projection With a larger projection With a little projection Length of palate Long and narrow Long and narrow Long and narrow Short and wide Pre-maxilla position Same maxilla level Same maxilla level Below maxilla level Below maxilla level Accessory medial foramen position Absent Behind incisive foramina Present Absent Breadth of lateral palatal edges Slightly broad Slightly broad Moderately broad Broad Shape and spread of the hard palate U-shaped, do not reach the middle of the mesopterygoid fossa V-shaped, do not reach the middle of the mesopterygoid fossa U-shaped, do not reach the middle of the mesopterygoid fossa V-shaped, pass the middle of the mesopterygoid fossa Size of auditory bullae Short and narrow Short and narrow High and swollen High and narrow Breadth of basioccipital at cochlea level Broad Broad Broad Narrow Cleft between protocone and hypocone of M1 and M2 Absent Absent Absent Present Accessory cuspids of lower canines Poorly developed Poorly developed Moderately developed Well developed Coronoid process height Larger than breadth between condylar processes Larger than breadth between condylar processes Shorter than breadth between condylar processes Shorter than breadth between condylar processes Coronoid position Below diastema level Below diastema level Below diastema level Above diastema level Shape of mandible profile Straight Straight Heel shaped Diagonal Secondary cuspid on P4 Present Present Absent Absent

Redescription of Mimon koepckeae 385 logenetic analyses based on morphological, molecular, and combined data, which have found Mimon to be polyphyletic (AGNARSSON et al. 2011, DÁVALOS et al. 2012); however, these analyses have only included M. bennettii and M. crenulatum as representatives of the genus. In order to clarify the relationships within the genus, a phylogenetic analysis including all species of Mimon needs to be conducted, because Anthorhina is a synonym of Tonatia (Gardner & Ferrel, 1990); then, if the clade formed by M. koepckeae and M. crenulatum is monophyletic, it would require a new generic name, as suggested by SIMMONS (2005) and WILLIAMS & GENOWAYS (2008). Analyses of fecal samples allocate Mimon koepckeae into the insectivore guild, which agrees with the molar dilambdont pattern that predicts insectivore-feeding behavior in bats (FREE- MAN 1988). Mimon koepckeae feeds on Elateridae and Scarabaeoidae beetles, insects that inhabit upper leaves (BROWNE & SCHOLTZ 1995, COSTA & ROSA 2011). Similarly, M. crenulatum also mainly eats beetles (MELLO & POL 2006), and occasionally some moths (GIANNINI & KALKO 2005), foraging at the understory level (SIMMONS & VOSS 1998). Based on diet items, we assume that M. koepckeae uses the same foraging stratum of M. crenulatum, but our conjecture is based on the analysis of only one specimen. On the other hand, M. bennettii forages both at understory and canopy levels (SIMMONS & VOSS 1998) and its diet includes a wider spectrum of items such as beetles (Scarabaeidae, Elateridae, Passalidae, Lamperidae, Chrysonelidae), lepidopterans (Saturniridae), cicadas (Hemiptera), and some scorpions and spiders (CARVALHO et al. 2007). Finally, M cozumelae feeds on a variety of insects, lizards and fruits (ORTEGA & ARITA 1997). It appears that Mimon also differs in diet and feeding-foraging strategies, because members of an unnamed subgenus (formerly called Anthorhina) eat understory insects, while members of the sub-genus Mimon eat a variety of items from both understory and canopy. Unfortunately, the region where the habitat of M. koepckeae is located is under great threat from the substantial loss of habitat due to demographic expansion, and intensive agriculture. Several roads bisect the Montane forest habitats at high altitudes, and along them there are several settlements (TOSI 1960). This could promote changes in landscape use by deforestation and subsequent lost of the habitat of M. koepckeae. Likewise, the Podocarpus locality is a touristic and recreational zone within the Santuario Nacional Pampa Hermosa area (SERNANP 2012). We suggest that the area of the Podocarpus should be preserved, in order to protect one of the few areas where M. koepckeae has been recorded. According to conservation criteria of the International Union for Conservation for Nature IUCN (2012a), Mimon koepckeae is listed as a Data Deficient species (VELAZCO & AGUIRRE 2008). Considering the endemic condition of M. koepckeae in Peru (PACHECO et al. 2009), the potential loss of its habitat (TOSI 1960, SERNANP 2012), rarity of records (GARDNER & PATTON 1972, PACHECO et al. 2007, and this study), and its categorization as Critically Endangered by the Peruvian legislation (MINISTERIO DE AGRICULTURA 2014) and the B2 criterion (subsections a, bi, bv) of the IUCN (2012b), we suggest revision of the conservation status of M. koepckeae and categorize it as Endangered species. ACKNOWLEDGEMENTS We especially thank to Adela Aguilar, José Alvarez, Anthony Almeyda, Jaime Pacheco, and Alexis Larico for their support in the field expeditions; Anamelba Zambrano Y., Jefe del Santuario Nacional Pampa Hermosa, for granting permit for this research and field facilities; Bruce D. 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Redescription of Mimon koepckeae 387 TOSI, J.A. 1960. Zonas de vida natural en el Perú: Memoria explicativa sobre el Mapa Ecológico del Perú. Lima, Instituto Interamericano de Ciencias Agrícolas de la OEA, Zona Andina, Proyecto 39, Programa de Cooperación Técnica, Boletín Técnico 3, 271p. TOVAR-NARVÁEZ, A.; C. TOVAR-INGAR; J. SAITO-DÍAZ; A. SOTO HURTADO; F. REGAL-GASTELUMENDI; Z. CRUZ-BURGA; C. VELIZ-ROSAS; P. VÁSQUEZ RUESTA & G. RIVERA CAMPOS. 2010. Yungas Peruanas Bosques montanos de la vertiente oriental de los Andes del Perú: Una perspectiva ecorregional de conservación. Lima, CDC UNALM, 150p. VELAZCO, P.M. 2005. Morphological phylogeny of the bat genus Platyrrhinus Saussure, 1860 (Chiroptera: Phyllostomidae) with the description of four new species. Fieldiana, Zoology 105: 1-54. VELAZCO, P.M. & L. AGUIRRE. 2008. Mimon koepckeae, IUCN Red List of Threatened Species. Version 2012.2. Available online at: www.iucnredlist.org [Accessed: 03/II/2013]. WETTERER, A.L.; M.V. ROCKMAN & N.B. SIMMONS. 2000. Phylogeny of Phyllostomid bats (Mammalia: Chiroptera): data from diverse morphological systems, sex chromosomes, and restriction sites. Bulletin of the American Museum of Natural History 248: 1-200. WILLIAMS, S.L. & H.H. GENOWAYS. 2008. Genus Mimon, p. 281-286. In A.L. GARDNER (Ed.). Mammals of South America. Chicago, The University of Chicago Press, vol. 1, 669p. Appendix 1. Specimens examined. Voucher of specimens are deposited in Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos (MUSM), American Museum of Natural History (AMNH), Field Museum of Natural History (FMNH) and National Museum of Natural History (NMNH). Mimon bennettii (n = 5). BRAZIL, Minas Gerais: Sete Lagoas (19 27 00"S, 44 14 00"W, NMNH 391027). São Paulo: Bairro do Matodentro, Caverna Tiraprosa (23 26 00"S, 46 39 00"W, AMNH 256294). COLOMBIA: CORDOBA: Tierralta: Socorre, upper Rio Sinu (07 51 00"N, 76 17 00"W, FMNH 69425 69427) Mimon cozumelae (n = 31). Belize, Cayo District: 05 m West Augustine, a Cave along Río Frio Cave (16 58 00"N, 88 59 27"W, FMNH 58152 58153). Barton Creek at Western Highway (17 12 00"N, 88 57 00"W, FMNH 58155 58156). Churchyard, Glenwood Farm (17 17 00"N, 88 34 00"W, FMNH 58154). Listowel School (Baking Pot), along Belize River (17 12 11"N, 89 01 43"W, FMNH 58157). Western Highway at Barton Creek (17 12 00"N, 88 57 00"W, FMNH 108764 108766). Orange Walk District: Gallon Jug (17 33 00"N, 89 01 00"W, AMNH 274575). Lamanai (17 45 50"N, 88 39 00"W, AMNH 277692). Toledo District: Aguacate, along Creek (16 10 12"N, 89 05 24"W, FMNH 108767). Crique Negro (15 58 00"N, 89 06 00"W, NMNH 506467). Pueblo Viejo (16 12 19"N, 89 08 30"W, FMNH 58150 58151). GUATEMALA, Peten: Tikal National Park (17 15 00"N, 89 39 00"W, FMNH 58567). PANAMA: Bocas del Toro: Almirante: Changuinola (09 26 00"N, 82 31 00"W, NMNH 315220). MEXICO: Chiapas: Palenque: Palenque (17 30 33"N, 91 58 56"W, FMNH 150630-150631). Oaxaca: Itsmo: Juchitan, 20 miles north of Matias Romero (17 03 03"N, 95 01 00"W, AMNH 185862 185872). Yucatan: Mérida, Buenavista, Xbac (21 15 00"N, 88 49 30"W, FMNH 5845) Mimon crenulatum (n = 181). BRAZIL, Amazonas: Borba (Santo Antonio da Uayara, Madeira River, 06 42 00"S, 69 52 00"W, AMNH 92223-92226, 92387-92388). Manaus: (Igarape Cacao Pereira, Negro River, 03 09 00"S, 60 07 00"W, AMNH 79526). São Gabriel do Cachoeira, (Taua, Uaupes River, 03 36 00"N, 69 12 00"W, AMNH 78651-78658, 78832-78833). BOLIVIA, Beni: Mamore (Baures River mouth, 12 30 00"S, 64 18 00"W, AMNH 209323). ECUADOR, Manabi: Sucre (Bahia de Caraquez, 00 36 00"S, 80 26 00"W, AMNH 64537 64541). FRENCH GUIANA: Cayenne: Sinnamary (Paracou, 04 56 00"N, 52 20 00"W, AMNH 267109, 267111-267115, 267437, 267880-267881, 267883-267884, 267886-267889). GUATEMALA, Peten: Tikal National Park (17 15 00"N, 89 39 00"W, FMNH 58568-58569). PANAMA: Bocas del Toro: Guabito (Sibube, 09 36 00"N, 82 49 00"W, NMNH 335121). Cusapin, (Peninsula Valiente, 09 08 00"N, 81 55 00"W, NMNH 57846). PERU: Amazonas: Condorcanqui (El Cenepa, Condorcanqui, P.V. 22, Falso Paquisha, Cordillera el Condor, 04 01 01"S, 78 24 00"W, MUSM 351). Cusco: La Convención (Echarate, Camisea, 11 53 00"S, 72 39 00"W, NMNH 582790). La Convención (Camisea, Armihuari, 11 51 51"S, 72 46 46"W, MUSM 13738-13739). La Convención (Camisea, San Martin, 11 47 00"S, 72 42 00"W, MUSM 13740-13741). La Convención (Camisea, Segakiato, 11 50 42"S, 72 35 59"W, MUSM 14771). Huánuco: Puerto Inca (Llullapichis River, tributary of Pachitea River, 09 37 00"S, 74 08 00"W, AMNH 236001). Monte Alegre (09 21 38"S, 75 19 33"W, AMNH 67230-67231). Loreto: Alto Amazonas (Nuevo San Juan, Galvez River, 05 15 00"S, 73 10 00"W, AMNH 272769-272770, 272834). Mariscal Ramon Castilla, (Río Yavari Mirim, San Vicente, 04 21 49"S, 72 29 30"W, FMNH 89038-89040). Maynas (Alto Nanay, Quebrada Agua Blanca, 02 55 26"S, 74 49 01"W, MUSM 24423). Boca del río Curaray (02 22 00"S, 74 05 00"W, AMNH 71690). Fernando Lores, (Estación Biológica Quebrada Blanco 2, 04 21 00"S, 73 09 00"W, MUSM 21198). Indiana (Indiana Chacra de Clayder, 03 29 39"S, 73 02 21"W, MUSM 32080-32081). Mazán, (Sucusari, Quebrada Grande, 03 15 26"S, 72 55 03"W, MUSM 21199). Punchana (Barrio Orosa Estación Madreselva II, río Amazonas, 03 37 40"S, 72 14 24"W, MUSM 32052-32053, 32055). Comunidad de Manacamiri (Río Nanay, Fundo Morropon, 03 42 31"S, 73 18 04"W, MUSM 28593). Iquitos Picuro Yacu (Purma, 03 37 05"S, 73 16 06"W, MUSM 32082). Padrecocha (03 41 29"S, 73 17 07"W, MUSM 28602). San Juan Bautista (13 de Febrero, Fundo Nemith, E km 33 de la carretera Iquitos-Nauta, 04 01 31"S, 73 25 47"W, MUSM 28591). 500 m E km 28.8 de la

388 N. Hurtado et al. carretera Iquitos-Nauta (03 59 14"S, 73 24 55"W, MUSM 28622). Cahuide km 61 carretera Iquitos-Nauta (04 15 46"S, 73 30 05"W, MUSM 32060). Camino a El Paujil, 1,8 km al W del km 35 de la carretera Iquitos-Nauta (04 01 13"S, 73 26 47"W, MUSM 28592). Caserio Cahuide Km 59 carretera Iquitos-Nauta (04 14 32"S, 73 29 31"W, MUSM 32061 32065). Caserio Cahuide Km 60.4 carretera Iquitos-Nauta, O del camino (04 14 54"S, 73 29 58"W, MUSM 32066). Cerca Allpahuayo, km 28 SO carretera a Nauta (03 59 00"S, 73 24 51"W, MUSM 32067). Cerca de la Aldea, km 18 SO carretera a Nauta (03 54 16"S, 73 22 14"W, MUSM 32068 32069). El Dorado, km 25 de la carretera Iquitos-Nauta, app. 1.5 km al E (03 58 00"S, 73 23 37"W, MUSM 28594). El Triunfo Km 48 carretera Iquitos-Nauta (04 09 02"S, 73 28 03"W, MUSM 32070). Fundo Mery Rojas Km 19.7 carretera Iquitos-Nauta, 10 min. NO (03 54 48"S, 73 22 58"W, MUSM 32072 32073). Habanillo Km 53 carretera Iquitos-Nauta (04 11 30"S, 73 28 45"W, MUSM 32074 32079). La Habana Km 52 carretera Iquitos-Nauta (04 10 52"S, 73 28 52"W, MUSM 32083 32085). Moralillo, 1.5 km E 500 m S del km 15.2 de la carretera Iquitos-Nauta (03 54 23"S, 73 20 37"W, MUSM 28595, 28597). Ninarumi, 7.4 km al W del km 6 de la carretera Iquitos-Nauta (03 50 30"S, 73 22 51"W, MUSM 28598). Ninarumi, 7.4 km al W y 1 km al SE del km 6 de la carretera Iquitos-Nauta (03 50 59"S, 73 22 26"W, MUSM 28599). Ninarumi, 7.4 km al W y 500 m al SE del km 6 de la carretera Iquitos-Nauta (03 50 44"S, 73 22 49"W, MUSM 28601). Paujil, W km 37.45 de la carretera Iquitos-Nauta (04 03 32"S, 73 26 32"W, MUSM 28603 28606). Peña Negra, 600 m al W del km 10 de la carretera Iquitos-Nauta (03 51 19"S, 73 20 42"W, MUSM 28607, 28609). Peña Negra, 800 m al E del km 11 de la carretera Iquitos-Nauta (03 52 24"S, 73 20 08"W, MUSM 28610, 28612 28616). Peña Negra, Km 10 carretera Iquitos-Nauta (03 51 14"S, 73 20 48"W, MUSM 32095 32096). Puerto Almendra Arboretum de CEIFOR (03 50 20"S, 73 22 28"W, MUSM 32097). Quistocoha Km 5 carretera Iquitos-Nauta, 1 km O del camino, Fundo Quistococha (03 49 03"S, 73 19 55"W, MUSM 32101). San Juan (Avenida de la Participación, 03 46 43"S, 73 16 44"W, MUSM 32102). San Lucas (W km 43 de la carretera Iquitos-Nauta, 04 06 15"S, 73 27 48"W, MUSM 28617-28618). Trece de febrero Km 31.5 carretera Iquitos-Nauta, Estación de campo UNAP (03 59 59"S, 73 26 31"W, MUSM 32103). Santo Tomás (6 km al W del km 1 de la carretera Iquitos-Nauta, 03 48 35"S, 73 20 17"W, MUSM 28619). Varillal (400 m W 200 m N del km 14 de la carretera Iquitos-Nauta, 03 52 57"S, 73 21 17"W, MUSM 28621). Torres Causana (Río Lagartococha, Campamento Catalino, 00 31 42"S, 75 15 35"W, MUSM 21200). Requena, (Jenaro Herrera, 04 54 17"S, 73 40 22"W, MUSM 870, 5914). Requena (C.I. Jenaro Herrera, 04 55 01"S, 73 45 00"W, MUSM 869). Ucayali, (Contamana, Sierra de Contamana, Cerros de Canchaguaya, Aguas Calientes, 07 11 20"S, 74 56 54"W, MUSM 17959). Madre de Dios: Manu (Alto Rio Madre de Dios, Hacienda Amazonia, 12 52 38"S, 71 23 11"W, MUSM 9826). Trocha 2 Mirador- Otorongo (12 33 38"S, 70 05 50"W, MUSM 26131). Manu (Pakitza, 11 56 47"S, 71 17 00"W, MUSM 479, 6791, 6792). Pasco: Oxapampa (Palcazu, Cerro Chontiya, 5Km Oeste Shiringamazu, carretera a Iscosazin, 10 15 00"S, 75 10 59"W, MUSM 10250-10251). Cerro Jonatan (5km E Lontananza, 10 21 57"S, 75 11 23"W, MUSM 10252). Hacienda Roca-Lux (Lontananza, Rio Mucñis, afluente del Río Iscosazin, 10 15 00"S, 75 11 00"W, MUSM 10253). Iscozacin (10 11 05"S, 75 08 43"W, MUSM 720). Río Pescado (10 22 48"S, 75 14 24"W, MUSM 24198). San Juan (10 30 00"S, 74 53 00"W, NMNH 364271). San Pablo (10 27 00"S, 74 52 00"W, AMNH 230130-230131, 230133-230136). San Martin: Rioja (Yaracyacu, Aguajal Rio mayo, 05 57 00"S, 77 11 00"W, MUSM 35204-35206). Tumbes: Tumbes (Pampas de Hospital, Qda. Angostura, 03 45 33"S, 80 22 56"W, MUSM 19346-19347). Zarumilla (Matapalo, Quebrada Naranjos, 03 50 31"S, 80 12 08"W, MUSM 19190, 19348). Rio Zarumilla (Carrizalillo, 03 44 38"S, 80 11 25"W, MUSM 22172). Rio Zarumilla (Carrizalillo 2, 03 44 38"S, 80 11 32"W, MUSM 22173). Ucayali: Coronel Portillo (Yarinacocha, 08 18 00"S, 74 36 00"W, FMNH 62120). Padre Abad (Irazola, Padre Abad, B.N. Von Humboldt, 08 46 59"S, 75 07 59"W, MUSM 8473). Purus (Rio Curanja, Balta, 10 08 00"S, 71 13 00"W, MUSM 1145). SURINAME, Sipaliwini: Coeroeni (Kayser Gebergte Airstrip, East of Zuid River, 03 07 00"N, 56 27 00"W, FMNH 93208). TRINIDAD AND TOBAGO, Trinidad: Nariva County (Bush Bush Forest, Nariva Swamp, 10 23 00"N, 61 02 00"W, AMNH 119420-119421). Saint George County (Blanchisseusse, Las Cuevas, 10 47 00"N, 61 23 00"W, AMNH 256308). Port of Spain, Belmont (10 40 00"N, 61 30 00 W, AMNH 175586). Port of Spain (10 39 00"N, 61 31 00"W, AMNH 207063). VENEZUELA, Amazonas: Atabapo (Mount Duida, Middle Camp, 03 25 00"N, 65 40 00"W, AMNH 77524). Falcon: Bolivar (Tocuyo River, 10 16 00"N, 69 56 00"W, AMNH 130687-130699, 130724, 131085-131086). Mimon koepckeae (n = 3). PERU, Ayacucho: La Mar (Huanhuachayo, 12 44 00"S, 73 47 00"W; LSUMZ 15676). Estera Ruana (12 43 00"S, 73 49 00"W; AMNH 233222). Junin: Chanchamayo (Santuario Nacional Pampa Hermosa, Podocarpus, 10 59 49.2"S, 75 25 57.1"W; MUSM 41327). Submitted: 24.XII.2013; Accepted: 06.VII.2014. Editorial responsibility: Diego Astúa de Moraes