Molecular and morphological identification of Phylloderma stenops Peters, 1865 (Chiroptera, Phyllostomidae) and new records for Colombia Juan M. Martínez-Cerón1, Edilson Patiño-Castillo2, Sara Carvalho-Madrigal3, Juan F. Díaz-Nieto1 1 Grupo Biodiversidad, Evolución y Conservación (BEC), Departamento de Ciencias Biológicas, Escuela de Ciencias, Universidad EAFIT, Carrera 49 No. 7 sur-50, Medellín, Colombia.  2 Grupo Mastozoología, Instituto de Biología, Facultad de Ciencias Exactas y Naturales, Universidad de Antioquia, Calle 67 No. 53-108, AA 1226, Medellín, Colombia.  3 Programa de Biología, Universidad CES, Calle 10 A No. 22-04, Medellín, Colombia. Corresponding author: Juan M. Martínez-Cerón, juanmceron@gmail.com Abstract Based on revisionary work of recently collected material in Colombian museums we confirm the presence of Phyl- loderma stenops Peters, 1865 in 6 new localities for the country, including the first record of the species in the dry lowlands of the northern Caribbean coast, and the increase by more than 800 m of the elevational range of the species in Colombia. DNA-barcoding confirmed our morphological identification, and supported a paraphyletic composition of the cis-Andean populations. Our records exemplify the little knowledge on the ecogeographic distribution of this species and provide further evidence to consider this as a widespread but rare species. Key words DNA barcode, ecogeographic distribution, Pale-faced Bat. Academic editor: Sergio Solari  |  Received 22 January 2018  |  Accepted 14 October 2018  |  Published 18 January 2019 Citation: Martínez-Cerón JM, Patiño-Castillo E, Carvalho-Madrigal S, Díaz-Nieto JF (2019) Molecular and morphological identification of Phylloderma stenops Peters, 1865 (Chiroptera, Phyllostomidae) and new records for Colombia. Check List 15 (1): 37–44. https://doi.org/ 10.15560/15.1.37 Introduction The pale-faced Bat, Phylloderma stenops Peters, 1865, is the only representative of its genus and includes 3 recognized subspecies primarily based on morphometric and distribution data: P. s. boliviensis Barquez & Ojeda, 1979, from southeastern Bolivia, P. s. septentrionalis Goodwin, 1940, distributed in Central America, and the nominal subspecies P. s. stenops Peters, 1865, distributed throughout most of South America from the Guianas to southeastern Brazil (Goodwin 1940, Husson 1962, Hand- ley 1966, Barquez and Ojeda 1979, Simmons and Voss 1998, Emmons and Feer 1999, Simmons 2005, Williams and Genoways 2008). Phylloderma stenops is considered a large and robust bat that externally may be confused with taxa of the genus Phyllostomus (Emmons and Feer 1999, Wetterer et al. 2000, Williams and Genoways 2008), an event that has facilitated its misidentification with species of the latter genus not only in the field but also in biological collections. Check List 15 (1): 37–44 https://doi.org/10.15560/15.1.37 1 15 Copyright Martínez-Cerón et al.  This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. NOTES ON GEOGRAPHIC DISTRIBUTION https://doi.org/10.15560/15.1.37 https://doi.org/10.15560/15.1.37 38 Check List 15 (1) In Colombia, the species is known from a few records primarily from cis-Andean localities which are mostly categorized as terrestrial ecoregions (sensu Dinerstein et al. 2017) of moist forests (Fig. 1: localities 2, 5, 8 and 9) (Marinkelle and Cadena 1972, Lemke et al. 1982, Alberico 1994, Montenegro and Romero-Ruiz 1999), a single locality at the ecotone between montane and dry forests (Fig. 1: locality 11) (Sánchez-Palomino et al. 1993) and a single locality at the llanos (Fig. 1: locality 10). The only two available records with trans-Andean distributions come from northwestern Andean montane forests (Fig. 1: locality 6) (Alberico 1994) and some recent records at the Magdalena Valley dry forest (Fig. 1: local- ity 14) (Ramírez-Fráncel et al. 2015). All the available Figure 1. Collecting localities and ecoregions of Phylloderma stenops for Colombia. Red circles represent new records; blue circles cor- respond to previously published records (see text). Numbers correspond to localities in Table 2. Martínez-Cerón et al.  |  Distribution of Phylloderma stenops in Colombia 39 records are distributed in an elevational range between 0 and 900 m a.s.l. Herein, we review the distribution of the genus in Colombia, providing new localities based on museum specimens as well as recently collected material, and we additionally corroborate our morphology-based identification using a DNA-barcoding approach. Methods In search for additional unpublished or misidentified re- cords and to corroborate the identification of eight previ- ously published records of the species (Sánchez-Palomino et al. 1993, Montenegro and Romero-Ruiz 1999, Ramírez- Fráncel et al. 2015) we visited the Colección Teriológica Universidad de Antioquia, Medellín (CTUA), Colección de Mamíferos “Alberto Cadena García”, Instituto de Cien- cias Naturales, Bogotá (ICN) and the mammal collection of the Museo de Ciencias Naturales de la Salle of the In- stituto Tecnológico Metropolitano, Medellín (CSJ-m). For the morphology-based identification of all specimens, we followed the diagnostic characters described by Goodwin (1940, 1942), Husson (1962), Barquez and Ojeda (1979), and Williams and Genoways (2008). To further corroborate the identification of the spe- cies we DNA-barcoded specimen CSJ-m 995—an adult individual that exhibits all the diagnostic characters of the species (see below)—and performed a phylogenetic analysis. Following a total DNA extraction using the UltraClean Tissue & Cells DNA Isolation Kit (MO BIO Laboratories, Inc.), part of the mitochondrial marker Cytochrome Oxidase Subunit I (COI) was PCR-amplified using the primer cocktail and thermal cycling condi- tions described in Ivanova et al. (2012). Amplicons were sequenced using amplification primers and dye-termina- tor chemistry on an ABI-3730xl automated sequencer. Sequences were edited and assembled in GENEIOUS® 9.1.8 (http://www.geneious.com, Kearse et al. 2012). The obtained sequence was deposited in GenBank with accession number MH298327. We downloaded all the available COI sequences of Phylloderma stenops from GenBank (Table 1) and to corroborate the genus-level identification we also downloaded sequences from at least one species of all the genera within the phyllostomid subfamily Phyllostominae (access numbers JF435806, JF447419, JF447421, JF447429, JF448940, JF449253, JF455242, JF455365, JF455386). All sequences were aligned with MUSCLE (Edgar 2004) and the resulting alignment was analyzed using maximum likelihood as implemented in GARLI 2.0 (Zwickl 2006), specifying 5 independent search replicates, and HKY+G+I as the best fitted nucleotide substitution model—based on the Bayes- ian Information Criterion implement in JMODELTEST Table 1. Sequences of Phylloderma included in this report. * Correspond to unpublished data, ** corresponds to a sequence without voucher. Genbank accession no. Voucher Country COI (bp) Source EF080541 ROM111506 Guyana 608 Clare et al. 2007 EF080542 ROM98903 Guyana 657 Clare et al. 2007 EF080543 ROM106627 Guyana 657 Clare et al. 2007 EF080544 ROM107203 Guyana 657 Clare et al. 2007 EF080545 ROM104694 Guyana 657 Clare et al. 2007 EU096830 ROM117511 Suriname 657 Borisenko et al. 2008 EU096831 ROM117178 Suriname 657 Borisenko et al. 2008 HQ545554 ROM120079 Suriname 657 Lim et al. 2010* HQ545600 ROM120137 Suriname 657 Lim et al. 2010* HQ545647 ROM120189 Suriname 657 Lim et al. 2010* HQ545674 ROM120221 Suriname 657 Lim et al. 2010* HQ919735 ROM120383 Suriname 657 Lim et al. 2010* JF447427 ROM104225 Panama 657 Clare et al. 2011 JF447691 ROM117063 Suriname 657 Clare et al. 2011 JF448984 ROM105186 Ecuador 657 Clare et al. 2011 JF448985 ROM105749 Ecuador 657 Clare et al. 2011 JF448986 ROMF40043 Ecuador 647 Clare et al. 2011  JF455174 ROM116713 Guyana 657 Clare et al. 2011 JF455175 ROM119337 Guyana 657 Clare et al. 2011 JF455176 ROM115679 Guyana 657 Clare et al. 2011 JF455177 ROM112628 Guyana 657 Clare et al. 2011 JF455178 ROM111529 Guyana 657 Clare et al. 2011 JF455179 ROM109322 Guyana 657 Clare et al. 2011 JF455180 ROM108937 Guyana 657 Clare et al. 2011 JF455181 ROM108410 Guyana 657 Clare et al. 2011 JF455182 ROM108529 Guyana 657 Clare et al. 2011 JF455183 ROMF39660 Guyana 657 Clare et al. 2011 JF455184 ROM106673 Guyana 657 Clare et al. 2011 JF455185 ROM104828 Guyana 656 Clare et al. 2011 JF455186 ROM98106 Guyana 657 Clare et al. 2011 KU295479 Isolate C1164 French Guiana 579 Thoisy et al. 2016** http://www.geneious.com 40 Check List 15 (1) (Darriba et al. 2012). Nodal support was evaluated based on 1,000 bootstrap replicates also using GARLI 2.0. Results New records. CSJ-m 995: Antioquia Department: mu- nicipality of Urrao: Parque Nacional Natural Las Or- quídeas: El Macho (06.5414° N, 076.2356° W, 1730 m a.s.l.; Fig. 1: locality 4), collected by J.F. Díaz-Nieto, S. Carvalho-Madrigal, E. Patiño-Castillo, and J.M. Martínez-Cerón, 9 October 2014. CTUA 753: Antio- quia Department: municipality of Maceo: Hacienda Santa Barbara (06.5441° N, 074.6404° W, 500 m a.s.l.; Fig. 1: locality 3), collected by J. Muñoz, 14 April 2004. CTUA 1692: Colombia: Sucre Department: municipality of San Onofre: Reserva Natural Sanguaré (09.7056° N, 075.6815° W, 11 m a.s.l.; Fig. 1: locality 13), collected by D. Gómez, 4 February 2010. ICN 13747: Putumayo Department: municipality of Puerto Leguizamo: Parque Nacional Natural La Paya (00.2667° N, 075.0200° W, 230 m a.s.l.; Fig. 1: locality 12), collected by R. Polanco, 15 May 1994. ICN 14990: Guainía Department: munic- ipality of Inirida: La Ceiba, Bosque alto (03.6388° N, 067.8826° W, 100 m a.s.l.; Fig. 1: locality 7), collected by A. Cadena, C. Ariza, J. Alvarez, 19 March 1998. ICN 21030: Amazonas Department: municipality of Leticia (04.1934° S, 069.9379° W, 84 m a.s.l.; Fig. 1: locality 1), collected by H. Lopez, 29 April 2002. Comments and Identification. As part of a mammal inventory at the Parque Nacional Natural Las Orquídeas on 9 October 2014, an adult female of Phylloderma stenops (Fig. 2) was captured with mist net in secondary forest (northwestern Andean Montane Forest ecoregion) at El Macho, municipality of Urrao, Antioquia Depart- ment, Colombia (Fig. 1: locality 4). The specimen CSJ-m 995 is an adult female with an open pubic symphysis, preserved as skin and skeleton, with liver tissue preserved in ethanol (96%), and deposited in the mammal collection of the Museo de Ciencias Naturales de la Salle of Instituto Tecnológico Metropolitano (CSJ-m). It has the following measurements: total length 122 mm; tail length 18 mm; hind foot length 22 mm; ear length 26 mm; forearm length 73 mm; and maximum length of the skull 33 mm. We confirm the presence of P. stenops in 14 locali- ties (19 specimens) for Colombia, 8 of which were previously published and 6 are new (Fig. 1, Table 2). Specimen CSJ-m 995 collected in northwestern Andean montane forests (Fig. 1: locality 4) represents an increase of 830 m in the elevational distribution of the species for the country, and specimen CTUA 1692 corresponds to the first record of the species in the xeric scrubs of northern Caribbean coast (Fig. 1: locality 13), extend- ing the known distribution of the species in Colombia northwared by 534 km. All examined material conforms to the description of the species and the most diagnostic- relevant characters are illustrated in Figures 2 to 5. This is a large species with reported forearm length between 65–81.7 mm and the maximum length of the skull between 29–34.5 mm (Husson 1962, Barquez and Ojeda 1979, Williams and Genoways 2008). It has a face with prominent chin and lip facial projections (excrescences not warts like those found in Trachops), noseleaf wide at the base with a pointed tip, short dorsal hairs with red- dish-brown color as reported by Salas et al. (2014) (Fig. 2), white wingtips, calcar nearly equal to or shorter than Table 2. List of records of Phylloderma stenops in Colombia. Locality numbers and ecoregions are mapped in Figure 1; asterisk (*) indicates material that was not examined by us (see Methods). Local- ity no. Museum catalog no. Department Municipality Latitude Longitude Elev. (m) Ecoregions Source 1 ICN 21030 Amazonas Leticia -04.1934 -069.9379 84 Solimões-Japurá moist forests This work 2 TTU 9065* Amazonas Leticia -04.1489 -069.9357 85 Solimões-Japurá moist forests Marinkelle and Cadena 1972 3 CTUA 753 Antioquia Maceo 06.5441 -074.6404 500 Magdalena Valley montane forests This work 4 CSJ-m 995 Antioquia Urrao 06.5414 -076.2356 1730 Northwest Andean montane forests This work 5 ICN 14598 Caquetá Solano 00.0742 -072.4514 130 Caqueta moist forests Montenegro and Romero-Ruiz 1999 6 UV 4516* Chocó San Jose del Palmar 04.9510 -076.2874 900 Northwest Andean montane forests Alberico 1994 7 ICN 14990 Guainía Inirida 03.6388 -067.8826 100 Negro-Branco moist forests This work 8 UV 2747* Guainía Inirida 03.2425 -068.1971 100 Negro-Branco moist forests Alberico 1994 9 IAvH 2217* Meta La Macarena 02.5500 -074.0500 283 Caqueta moist forests Lemke et al. 1982 10 ICN 9612* Meta Puerto López 03.9654 -073.0502 190 Llanos Ramírez-Fráncel et al. 2015 11 ICN 10236*, 10237, 10238*, 10239*, 10240 Meta San Juan de Arama 03.3461 -073.9306 450 Apure-Villavicencio dry forests Sánchez-Palomino et al. 1993 12 ICN 13747 Putumayo Puerto Leguizamo 00.2667 -075.0200 230 Napo moist forests This work 13 CTUA 1692 Sucre San Onofre 09.7056 -075.6815 11 Guajira-Barranquilla xeric scrub This work 14 CZUT-M1330*, 1380 Tolima Ambalema 04.8488 -074.8138 253 Magdalena Valley dry forests Ramírez-Fráncel et al. 2015 Martínez-Cerón et al.  |  Distribution of Phylloderma stenops in Colombia 41 the length of the foot, and the tail reaches only half of the inter-femoral membrane (Fig. 3) (Husson 1962, Barquez and Ojeda 1979, Emmons and Feer 1999,Williams and Genoways 2008, Reid 2009, Díaz et al. 2011, Díaz et al. 2016). Although the species seems to be easily identi- fied, specimen ICN 4465 identified by Ramírez-Fráncel et al. (2015) as P. stenops, was re-identified by us as Phyllostomus elongatus (É. Geoffroy St.-Hilaire, 1810). As previously mentioned, the species is not uncommonly misidentified with members of the genus Phyllostomus. Nonetheless, our examined material of P. stenops can be easily differentiated from the former genus using dental characters: while Phylloderma has 3 lower premolars (the second of which is small and labially oriented) the genus Phyllostomus has only 2 premolars similar in size (Figs 4, 5) (Husson 1962, Emmons and Feer 1999, Wil- liams and Genoways 2008). Our phylogenetic analysis of COI sequences shows 3 salient features (Fig. 6). First, haplotype of specimen CSJ-m 995, identified as P. stenops using morphological characters, is nested with other sequences of the species with strong support—and not with any other Phyllosto- mine genera—implying that the morphological characters are reliable for an accurate identification of this species and delimitation from phyllostomine genera. Second, the Colombian haplotype is sister to the only other avail- able sequence from a trans-Andean locality (Panama), suggesting a trans-Andean clade for the species. Third, cis-Andean haplotypes appear to be paraphyletic and as previously found by other authors (Clare et al. 2011), the material from the Guiana Shield does not form a mono- phyletic group (Fig. 6). Discussion One of our new records, represented by the specimen CSJ-m995, increases the elevational distribution of the species in Colombia by 830 m, as the species had been recorded in the country only up to the 900 m a.s.l. (Alberico 1994); nonetheless, across its distribution (e.g., in Ecuador) this species was known to occur up to 1,750 m a.s.l. (Brito and Arguero 2012), and according to Emmons and Feer (1999) it can reach elevations of 2,600 m a.s.l. Additionally, before our identification of CTUA 1692 from the xeric scrubs in the Caribbean of Colombia (Fig. 1: locality 13), the northernmost known locality of Phylloderma stenops for Colombia was from northwestern Andean montane forests, (Fig. 1: locality 6) (Alberico 1994), and consequently our record increases the latitudinal distribution of the species in Colombia by 534 km. Despite our efforts for examining all the avail- able material at the 2 visited collections, we were unable to find several specimens reported by Sánchez-Palomino et al. (1993) and Ramírez-Fráncel et al. (2015). In par- ticular, the single available specimen from the Llanos ecoregion (Fig. 1: locality 10) was not found, and only 2 (out of 5) specimens from the ecotone between montane and dry forests (Fig. 1: locality 11) were found and posi- tively identified as P. stenops (Table 2). Although our phylogenetic analysis using mito- chondrial data had only identification purposes, it does show that despite its wide distribution, this species has a reduced mitochondrial variation—only 2.1% within spe- cies (p-) distance—across haplotypes that cover a wide longitudinal gradient from French Guiana to Panama, and only 0.2% of pairwise (p-) distance between the Colom- bian haplotype and its sister clade of Guyanese sequences (Fig. 6). Although relevant, the issue of the likely para- phyly of Guyanese mitochondrial haplotypes is out of the scope of this paper and should be addressed elsewhere. Many studies have catalogued P. stenops as locally rare but widespread (Handley 1976, Trajano1984, Arita 1993, McDade 1994, Simmons and Voss 1998, Emmons and Feer 1999, Nadkarni and Wheelwright 2000, Reid 2009) and our study seems to provide evidence to rein- forcing this concept. As an example, as part of a mammal inventory at Parque Nacional Natural Las Orquídeas, from more than 300 netted bats only a single individual of P. stenops was captured (Díaz-Nieto unpublished). Although for Colombia the species is only known for 14 localities, these encompass almost the entire latitudinal and longitudinal gradient of the country as well as a great Figures 2, 3. Pale-faced Bat, Phylloderma stenops, adult female (CSJ-m 995). 2. Individual captured at Parque Nacional Natural Las Orquídeas, Colombia. Photograph by CSG. 3. Skin of same specimen; scale bar = 25 mm. 2 3 42 Check List 15 (1) elevational gradient. Such wide geographical distribution includes mostly humid ecoregions (see above) but also at least 2 dry ecoregions, such as dry and xeric shrublands (Fig. 1: localities 11, 13 and 14). New field-based studies with increased sampling effort should be developed to gain knowledge on the ecogeographic limits of the species, in particular in areas with likely suitable habitat where the species has yet to be found (e.g., biogeographic Choco, Andean montane forests of Cordillera Central and Oriental). Additionally, a denser genetic (multiple loci) and geographic sampling should be useful in testing both the ecogeographic and evolutionary limits of the proposed subspecies, in partic- ular, relevant samples such as those from dry ecoregions and those from the northern and southern limits of its distribution would be welcome. Figure 6. A maximum-likelihood topology of COI sequences of Phylloderma stenops. Numbers on branches correspond to boot- strap support values. Each terminal is identified by country of origin and an alphanumeric specimen identifier (see Tables 1, 2). Bolded terminal corresponds to the sequence obtained in this report. Figure 5. Upper and lower incisors of adult female Phylloderma stenops (CSJ-m 995); scale bar = 2 mm. Figure 4. Skull and mandible of adult female Phylloderma stenops (CSJ-m 995); scale bar = 10 mm. Martínez-Cerón et al.  |  Distribution of Phylloderma stenops in Colombia 43 Acknowledgements We thank the curators and staff of institutions that allowed access to voucher material: Sergio Solari (CTUA), Danny Zurc and Andrea Bustamante (CSJ-m), Hugo López-Arévalo and Catalina Cárdenas (ICN). We are particularly grateful to our field crew Alejandro Aguirre, Camilo Sánchez-Giraldo, Flor Martínez, Jorge de Jesús Álvarez, Marcela Gómez, Milena Peñuela and Ovidio Álvarez Cartagena (R.I.P.). Hector Velásquez (Head of Parque Nacional Natural Las Orquídeas) was fundamen- tal for obtaining research permits. 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