Redescription of Mimon koepckeae ( Chiroptera : Phyllostomidae )

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.

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 (1976KOOPMAN ( , 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 (1993KOOPMAN ( , 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.

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 .This specimen was identified as Mimon koepckeae, based on the original species description (GARDNER & PATTON 1972).Gardner & Patton, 1972 Type locality: Huanhuachayo (12°44'00"S, 73°47'00"W), elevation 1,660 m, Ayacucho Department, Peru.

Mimon koepckeae
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 46.9-50.2mm; 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-horse-shoe 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-  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.
The reproductive status of M. koepckeae corresponds to an adult female with perforated genitalia, which indicates recent 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 (1976KOOPMAN's ( , 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-   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, andthis 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.

Figure 19 .
Figure 19.Cloud forest habitat of Podocarpus where Mimon koepckeae was collected in this work.Photo by Edith Arias.

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.

Table II .
(Gardner & Ferrel, 1990)racters of Mimon bennettii, M. cozumelae, M. crenulatum, and M. koepckeae.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 &