Redescription of Mimon koepckeae (Chiroptera: Phyllostomidae)

ZOOLOGIA 31 (4): 377–388, August, 2014http://dx.doi.org/10.1590/S1984-46702014000400009

2014 Sociedade Brasileira de Zoologia | www.sbzoologia.org.br | www.scielo.br/zoolAll content of the journal, except where identified, is licensed under a Creative Commons attribution-type BY-NC.

Mimon koepckeae Gardner & Patton, 1972 is an endemicPeruvian bat whose distribution used to be restricted to thetype locality in Ayacucho Department, Peru, and surround-ings (SIMMONS 2005, WILLIAMS & GENOWAYS 2008, VELAZCO &AGUIRRE 2008). This species is known from three specimens (twofemales and one male), which were captured in the localitiesof Estera Ruana and Huanhuachayo, between 1,600 and 1,900m (GARDNER & PATTON 1972). Natural history information aboutM. koepckeae is scarce, and the description of its habitat offersfew details, for instance a reference to cloud forests of the east-ern 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 theabsence of a dorsal stripe, fewer crenulations and scarce hairson the nose-leaf basis, narrow auditory bullae, and a well de-fined cleft between protocone and hypocone in the first andsecond upper molars. However, KOOPMAN (1976, 1978) consid-ered M. koepckeae as a junior synonym of Mimon crenulatum(Geoffroy St.-Hilaire, 1803), based on a morphological com-parison of skins and skulls of a single specimen of M. koepckeae,collected by Terborgh in 1972 from Estera Ruana (one localityof 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 intraspe-cific variation. Since that revision, KOOPMAN (1993, 1994) keptM. koepckeae as a highland Peruvian sub-species of M. crenulatumwithin 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 anthropo-centric activities, M. koepckeae is listed as Critically Endangeredby 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 yearsafter its description, and present a complete morphological char-acterization of it, including new diagnostic characters support-ing its status of valid species. Based on our results, the distributionof the species is significantly broadened. In addition, the habi-tat of the species, based on analysis of the new collecting local-ity, is characterized. Finally, we also discuss the nomenclaturalstatus of the sub-genera of Mimon, based on comparisons withother species of the genus (sensu SIMMONS 2005).

MATERIAL AND METHODS

Bat inventories were conducted in the locality ofPodocarpus, Santuario Nacional Pampa Hermosa, ChanchamayoDistrict, Chanchamayo Province, in Junin Department of Peru(Fig. 1, 10°59’49.2"S, 75°25’57.1"W, 1890 m). Ten mist nests (12x 2.5 m each one) were set up for seven consecutive nights, witha total sampling of 70 mist nests per hour.

The morphological characterization of Mimon koepckeaepresented here is based on the specimen recorded herein (Ap-pendix 1) and one topotype (AMNH 23222), which were con-

Redescription of Mimon koepckeae (Chiroptera: Phyllostomidae)

Natalí Hurtado1,2,4, Edith Arias1 & Víctor Pacheco1,3

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: [email protected]

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 surround-

ings. 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.

378 N. Hurtado et al.

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trasted with photos of the type specimen (LSUMZ 15676), andthose in the original description (GARDNER & PATTON 1972). Theage 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), andFRACASSO 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 ofMimon (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 placeda graph paper under a petri dish and analyzed stomach con-tents and feces in five randomly selected areas (5 x 5 mm) us-ing a stereomicroscope with 20x magnification.

TAXONOMY

An adult female of Mimon koepckeae was collected fromthe locality of Podocarpus in the Santuario Nacional PampaHermosa, Province of Chanchamayo, Department of Junin, Peru(Fig. 1). The specimen was captured by Edith Arias (EA 216) onOctober 6th, 2011. Its external measurements are: total length78 mm, tail length 23 mm, hindfoot length 11 mm, ear length22 mm, tragus length 9 mm, forearm length 48 mm, and weight14.5 g. This specimen was preserved as skin, with the skull re-moved, and was deposited in the mammal collection of theMuseo de Historia Natural, Universidad Nacional Mayor de SanMarcos, under catalogue number MUSM 41327 (Figs 2-6). Thisspecimen was identified as Mimon koepckeae, based on the origi-nal species description (GARDNER & PATTON 1972).

Mimon koepckeae Gardner & Patton, 1972

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 fromreddish brown to golden brown; dorsal stripe absent; noseleafslender, crenulated along proximal margin, and sparsely fringedwith short fine hairs; skull small, with narrow rostrum; audi-tory bullae narrow; first and second upper molars with narrow,well-defined vertical cleft separating protocone and hypocone.

Redescription. Size. Medium-sized bats. Forearm lengthbetween 46.9-50.2 mm; weight 14 g. Pelage. Dorsal fur at rumplevel short (6 mm); bicolored banding pattern is Pale SmokeGray at base and Chestnut at tips. Ventral fur at belly levelslightly shorter (5 mm); tricolored banding pattern with smallPale Smoke Gray at base, large Hair Brown on central portion,and Olive-Buff tips that turn whitish from central abdomentoward flanks. Pale Smoke Gray basal portion on head largerdorsal and ventrally. Hairs of auricular patch exhibit largerwhitish portion and Chestnut tips. Dorsal stripe absent. Head.Comparatively short rostrum, about one third head length orless, mandibular pragmatism not present. Noseleaf limb-horse-

shoe continuous. Limb ovate, length about twice width. Limbmargins pigmented, shallow crenulations at base disappearapically; tuft of long hairs at tip present. Limb-rib compara-tively thin and slimmer than internostrils space. Horseshoecontinuous around nostrils. Anterior noseleaf margin well de-veloped and free. Lips continuous and flat. Four vibrissae placedon each dorsal lip margin; and six vibrissae on each ventralmargin. Chin with two pairs of central pads; one pair placedbelow lip, the other, smaller, behind it; four dermal pad pairsplaced diagonally to central ones; dermal pads pigmented andsymmetrically separated by central shallow cleft, which doesnot extend behind chin. A set of seven vibrissae on each sideof horseshoe; above, a set of swollen continuous papillae. Earsslightly smaller than head length. Pinna delta shaped, fleshyand not translucent, with numerous marked folds, roundedand parallel at tips and edges continuous. Ears bicolored; ex-ternal half pigmented and basal half (including tragus) pale.Pinna poorly developed, inserted the head insertion level. In-ner ear lobe well developed; with long dense hairs, and well-

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.

379Redescription of Mimon koepckeae

ZOOLOGIA 31 (4): 377–388, August, 2014

developed and sparse protuberances on internal surface. Tra-gus long, about one third ear length. Tragus rib thick with scarcelong whitish hairs. Body. Single odor gland placed at center ofthroat. 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 borderof fifth digit. Dactilopatagium major exhibits V shaped slot.Medium and minus dactilopatagium exhibit complete pigmen-tation. 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 blackhairs on dorsal surface. Calcars longer than tibia. Tip of tailfree. Skull. Rostrum short, length average 21.11 mm. In dorsalview, incisor roots form V shaped groove. Anterior edge ofnostril V shaped. Maxillary roots of zygoma swollen. Interor-bital region narrower than width between-canines. Braincaseedge V shaped. In lateral view, a short terrace on the pre-max-illa exhibited at incisors level. Forehead plain without medialdepression. Zygomatic arches flat and thick, with hard andbroad roots. Jugal bone narrower. Sagittal crest wide at frontallevel and tapering towards posterior side. In ventral view, pal-

Figures 2-3. External appearance of Mimon koepckeae (MUSM 41327): (2) dorsal view; (3) ventral view. Scale bar: 3 cm.

Figures 4-6. (4) Dorsal, (5) ventral, and (6) lateral views of the skull of Mimon koepckeae (MUSM 41327). Scale bar: 1 cm.

2 3

4 5

6


ZOOLOGIA 31 (4): 377–388, August, 2014

ate short and wide, with concave floor. Pre-maxilla triangular,placed below maxillary level; posterior edge exhibits smallrounded projection towards maxillary. Lateral posterior edgesof palate wide, extending towards middle of M2 talonid. Post-palatal process well-marked. Hard palate edge V shaped; ante-rior border extended beyond middle of interpterygoid bone.Medial basisphenoid sharped septum forms basisphenoidal pits.Basioccipital flat and narrow. Basioccipital-basisphenoid sutureperpendicular to basisphenoid septum. Anterior border of fo-ramen magnum shallow. Jugular foramen the same size as ju-gal breath. Occipital condyles short and narrow. Para-occipitalprocesses developed, but shorter than occipital condyles. Au-ditory bullae laterally compressed, wider than diameter of fo-ramen magnum, with few swollen optic rings. Dental

morphology. Dental formula I 2/1, C 1/1, P 2/2, M 3/3. Up-per. Inner incisors rhomboid shaped, half canine length. Tipsand almost all inner borders free. Outer incisors flat and shorterthan inner ones. Canines pyramidal shaped; well-defined me-dial notch on lingual cingulum. Second premolar (P2) reduced,conical, labially placed and one sixth canine length; in ventralview, labial and lingual cingula subequal. Fourth premolar (P4)developed and conical, average size between canines and mo-lars; P4 exhibits well developed main cusp; labial cingulumnarrow, extended towards talon; talon basin wide and deep;secondary cusp absent; on posterior side, metastyle belowparastyle of first molar. Parastyle high of first molar (M1), high,but not reaching paracone level; mesostyle shorter thanparastyle; metacone higher than paracone; preparacrista,

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.

7 8 9 10

11 12 13 14


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postparacrista, premetacrista and postmetacrista with concaveedges; stylar shelves triangular; anterior ectoflexus convex;posterior ectoflexus triangular; protofossa sub equal to ante-rior stylar shelf; protocone developed and pyramidal; hypo-cone less developed; vertical sulcus between protocone andhypocone 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 thirdmolar (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 coni-cal; labial cingulum narrow; mesial cingulum ends in smallconule; parastyle and distal cingulum well developed. Secondpremolar (p2) short, about half size of canine, with single prin-cipal cone, asymmetrically divided. Fourth premolar (p4) largerthan p2, with single principal pyramidal cone, labial cingu-lum restricted to first half. First molar (m1) longer than wider;size average between canines and p4; paraconid, protoconidand metaconid linked by vertical crests with concave edges,

forming deep triangle basin; paraconid below p2 level; proto-conid stands out as the largest cuspid; and paraconid and en-toconid the smallest; hypoconulid placed posteriorly toentoconulid with a small peak; on posterior side, hypoconuliddoes not overlap m2. Second molar (m2) similar to m1, butoblique crest exhibits convex edge; labial cingulum exhibitsdeep cleft between trigonid and talonid; hypoconulid largerthan in m1. Paraconid, metaconid and protoconid of thirdmolar (m3) subequal, connected by vertical crests with con-cave edges shallower than m1 and m2; talonid less developedand shorter than hypoconulid; labial cingulum as m2; labialcingulum developed at trigonid level, but fewer at talonid level.Mandible. In dorsal view, condylar process breadth is largerthan the coronoid process height; coronoid processes slopedlaterally; angular processes shorter than condylar processes. Inlateral view, chin straight with well develop sub-mental pro-cess; posterior edge diagonal with angle less than 30°; toothrowshort; coronoid process short; condylar process thick and twicelarger than angular process, placed over diastema, diastemabetween toothrow and coronoid process; notch between an-gular and condylar processes trapezoidal.

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.

15

17

16

18


ZOOLOGIA 31 (4): 377–388, August, 2014

Distribution. Mimon koepckeae is known to occur only inthe type locality (Huanhuachayo) and vicinities of EsteraRuana; in Ayacucho Department, and low cloud forest of JuninDepartment, elevation between 1660 and 1890 m.

Remarks. Mimon koepckeae was compared with specimensrepresenting the three recognized species in the genus sensuSIMMONS (2005): M. crenulatum, M. cozumelae, and M. bennettii.

Compared with Mimon koepckeae, M. crenulatum is slightlylarger, average body length is 84.09 mm, dorsal fur is unicoloredDeep Mouse Gray and shorter, the dorsal stripe is alwayspresent, ventral fur is tricolored, auricular patch is bright andconspicuous, noseleaf crenulations and hairs are evenly dis-tributed, ears are shorter, calcar is slightly shorter, skull lengthis slightly larger, nasal depression is U shaped, incisors gumcrests are less swollen, medial depression is present, maxillaryroot of zygoma is less swollen, interorbital region is wider,posterior braincase projection is more developed, palate is largeand narrow, an anterior accessory medial foramen is presentin some populations, lateral palatal edges are narrower, poste-rior border of the hard palate is U-shaped and does not reachthe middle of mesopterygoid fossae, auditory bullae are wider,basioccipital is wider at cochlea level, cleft between protoconeand hypocone is absent in M1 and M2, accessory cuspids oflower canines are less developed, coronoid process is over di-astema level, posterior edge of mandible is heel shaped.

Mimon bennettii is larger, body length is 92 mm, dorsalfur is tricolored and longer, ventral fur is unicolored, noseleafis short and wide, noseleaf crenulations and hairs are absent, afree skin binding of the horseshow is absent, chin exhibits denseand 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, maxil-lary root of zygoma is flat, interorbital region is comparativelynarrower, palate is large and narrow, premaxillary and maxil-lary are at same level on the palate, lateral palatal edges arenarrower, posterior border of the hard palate is U-shaped anddoes not reach the middle of mesopterygoid fossae, auditorybullae are shorter and narrower, basioccipital is wider at co-chlea level, cleft between protocone and hypocone is absentin M1 and M2, accessory cuspids of lower canine are flatted,coronoid process is over diastema level, posterior edge of man-dible is flat, secondary cuspid of P4 is present.

Mimon cozumelae is larger, average body length is 94.5mm, dorsal fur is tricolored and longer, ventral fur is unicolored,noseleaf is short and wide, noseleaf crenulations and hairs areabsent, a free skin binding of the horseshow is absent, chinexhibits dense and long hairs; ears are longer, translucent withmarked folds, ear tips are divergent; forearm is furred, calcar isshorter, plagiopatagium is inserted at tibia level, skull lengthis 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 thepalate, a accessory medial foramen is exhibited behind the in-cisive foramina, lateral palatal edges are narrower, posteriorborder of the hard palate does not reach the middle ofmesopterygoid fossae, auditory bullae are shorter and narrower,basioccipital is wider at cochlea level, a cleft between proto-cone and hypocone is absent in M1 and M2, accessory cuspidsof lower canine are absent, coronoid process is over diastemalevel, posterior edge of mandible is flat, secondary cuspid ofP4 is present.

Table I provides external and cranial measurements foreach species. In addition, Table II provides a summary of ex-ternal and skull-dental characters comparisons among species.

Natural History. Podocarpus is located 160.1 km north-ward from the type locality, Huanhuachayo. The habitat inPodocarpus is dominated by tree species of Podocarpaceae andClusiaceae, which are approximately of 15 m high (Fig. 19).The understory is sparse and covered with Rubiaceae andAraceae shrubs (Fig. 19, LA TORRE-CUADROS et al. 2007). Accord-ing to TOSI (1960) and HOLDRIDGE (1967), this ecosystem is cat-egorized as Very Humid low Montane forest, also known asYungas (TOVAR-NARVÁEZ et al. 2010); it is a steep landscape form-ing an altitudinal belt from 1,600 to 2,800 m elevation.

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

Mimon koepckeae was collected in a natural canopy open-ing during the early rainy season, with Anoura aequatoris(Lönnberg, 1921), Carollia brevicauda (Schinz, 1821), Micronyc-teris megalotis (Gray, 1842), Myotis keaysi J.A. Allen, 1914, Myotisriparius Handley, 1960, Sturnira magna de la Torre, 1966, andVampyressa melissa Thomas, 1926. Details for this bat assem-blage will be reported elsewhere.

The reproductive status of M. koepckeae corresponds to anadult female with perforated genitalia, which indicates recent


ZOOLOGIA 31 (4): 377–388, August, 2014

sexual activity. Stomach content was scarce and unidentifiable,but fecal analysis revealed remains (i.e., elytra, wings, legs, andantennas) of Elateridae (30%) and Scarabaeidae (70%) beetles.

DISCUSSION

We recognize Mimon koepckeae as a valid species, basedon distinctive morphological attributes presented in this work,and thus refuting KOOPMAN’s (1976, 1978) arguments. The di-agnostic characters of this species are independent, and arenot contained within the range of morphological variabilityof M. crenulatum (i.e. nasal shape, dorsal stripe absent). Thisspecies is distributed in low Montane Forests from Orientalslope of Central Andes in Peru, but is not sympatric with M.crenulatum (GARDNER & PATTON 1972, this work), which rangesat lower elevations (300 to 900 m).

This new record of Mimon koepckeae is a rediscovery, be-cause this report is more than 40 years after the type series

description (see criteria of SCHEFFERS et al. 2011). It is also a dis-tribution range extension of 160 km northward from the typelocality and surroundings. The scarcity of records of M.koepckeae reflects the lack of intense fieldwork in the area ornot enough sampling, which are largely due to difficulties ac-cessing that zone.

Mimon koepckeae and M. crenulatum were placed in thesubgenus Anthorhina (GARDNER & PATTON 1972, SIMMONS & VOSS

1998). However, the taxonomic status of Anthorhina has beendiscussed, since HANDLEY (1960) synonymized it with Mimon.Although HUSSON (1962) considered Anthorhina as a genus, basedon differences in the relative size of the upper premolars, heightof bullae, and shape and nose-leaf pubescence, CABRERA (1958)and GOODWIN & GREENHALL (1961) considered Anthorhina onlyas 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 descrip-tion of Mimon provided by GRAY (1847), supporting recent phy-

Table I. External and cranial measurements ( in millimeters) of Mimon bennettii, M. cozumelae, M. crenulatum, and M. koepckeae. Standarddeviation 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.


ZOOLOGIA 31 (4): 377–388, August, 2014

Tab

le II

. Ext

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l and

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of M

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ZOOLOGIA 31 (4): 377–388, August, 2014

logenetic analyses based on morphological, molecular, andcombined data, which have found Mimon to be polyphyletic(AGNARSSON et al. 2011, DÁVALOS et al. 2012); however, these analy-ses have only included M. bennettii and M. crenulatum as repre-sentatives of the genus. In order to clarify the relationshipswithin the genus, a phylogenetic analysis including all speciesof Mimon needs to be conducted, because Anthorhina is a syn-onym of Tonatia (Gardner & Ferrel, 1990); then, if the cladeformed by M. koepckeae and M. crenulatum is monophyletic, itwould 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 dilambdontpattern that predicts insectivore-feeding behavior in bats (FREE-MAN 1988). Mimon koepckeae feeds on Elateridae and Scara-baeoidae beetles, insects that inhabit upper leaves (BROWNE &SCHOLTZ 1995, COSTA & ROSA 2011). Similarly, M. crenulatum alsomainly eats beetles (MELLO & POL 2006), and occasionally somemoths (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, butour conjecture is based on the analysis of only one specimen.On the other hand, M. bennettii forages both at understory andcanopy levels (SIMMONS & VOSS 1998) and its diet includes awider spectrum of items such as beetles (Scarabaeidae,Elateridae, Passalidae, Lamperidae, Chrysonelidae), lepidopter-ans (Saturniridae), cicadas (Hemiptera), and some scorpionsand spiders (CARVALHO et al. 2007). Finally, M cozumelae feedson a variety of insects, lizards and fruits (ORTEGA & ARITA 1997).It appears that Mimon also differs in diet and feeding-foragingstrategies, because members of an unnamed subgenus (formerlycalled Anthorhina) eat understory insects, while members ofthe sub-genus Mimon eat a variety of items from both under-story and canopy.

Unfortunately, the region where the habitat of M.koepckeae is located is under great threat from the substantialloss of habitat due to demographic expansion, and intensiveagriculture. Several roads bisect the Montane forest habitats athigh altitudes, and along them there are several settlements(TOSI 1960). This could promote changes in landscape use bydeforestation and subsequent lost of the habitat of M. koepckeae.Likewise, the Podocarpus locality is a touristic and recreationalzone within the Santuario Nacional Pampa Hermosa area(SERNANP 2012). We suggest that the area of the Podocarpusshould be preserved, in order to protect one of the few areaswhere M. koepckeae has been recorded.

According to conservation criteria of the InternationalUnion for Conservation for Nature – IUCN (2012a), Mimonkoepckeae is listed as a Data Deficient species (VELAZCO & AGUIRRE

2008). Considering the endemic condition of M. koepckeae inPeru (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) ofthe IUCN (2012b), we suggest revision of the conservation sta-tus of M. koepckeae and categorize it as Endangered species.

ACKNOWLEDGEMENTS

We especially thank to Adela Aguilar, José Alvarez, An-thony Almeyda, Jaime Pacheco, and Alexis Larico for their sup-port in the field expeditions; Anamelba Zambrano Y., Jefe delSantuario Nacional Pampa Hermosa, for granting permit forthis research and field facilities; Bruce D. Patterson for his aca-demic support and Mark Hafner for providing photos of thetype specimen. Also to Edgardo Rengifo and André Ampuerofor revisions; and Amelia Corso and Ann-Lloyd Hufstader forEnglish review of an earlier version of this manuscript. We wishto thank the partial support of the FMNH Scholarship Com-mittee to NH. This work was partially supported by grants111001031 and 121001061 of CSI-UNMSM to VP.

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Appendix 1. Specimens examined. Voucher of specimens are deposited in Museo de Historia Natural de la Universidad Nacional Mayorde San Marcos (MUSM), American Museum of Natural History (AMNH), Field Museum of Natural History (FMNH) and National Mu-seum 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, FMNH58152–58153). 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, AMNH92223-92226, 92387-92388). Manaus: (Igarape Cacao Pereira, Negro River, 03°09’00"S, 60°07’00"W, AMNH 79526). São Gabrieldo Cachoeira, (Taua, Uaupes River, 03°36’00"N, 69°12’00"W, AMNH 78651-78658, 78832-78833). BOLIVIA, Beni: Mamore (BauresRiver 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, FMNH58568-58569). 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, Cor-dillera 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, NMNH582790). La Convención (Camisea, Armihuari, 11°51’51"S, 72°46’46"W, MUSM 13738-13739). La Convención (Camisea, San Mar-tin, 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 PicuroYacu (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 (13de 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


ZOOLOGIA 31 (4): 377–388, August, 2014

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 carreteraIquitos-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). ElDorado, 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 carreteraIquitos-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). LaHabana 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 15.2de 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 carreteraIquitos-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, 800m 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 10carretera 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 43de 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, MUSM22172). 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 ZuidRiver, 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, AMNH207063). 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

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Redescription of Mimon koepckeae (Chiroptera: Phyllostomidae)