bioRxiv preprint doi: https://doi.org/10.1101/462051; this version posted November 5, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 1 Camilo A. Calderón-Acevedo 2 [email protected] 3 4 Taxonomy of large Anoura 5 Large Anoura (Chiroptera:Glossophaginae) taxonomy, taxonomic status of Anoura 6 carishina, and implications for the distribution of Anoura latidens in Colombia. 7 Camilo A. Calderón-Acevedo*, Miguel E. Rodríguez-Posada and Nathan Muchhala 8 Department of Biology, University of Missouri – St. Louis, One University Blvd St. Louis, 9 Missouri 63121. USA. (CAC and NM) 10 La Palmita Natural Reserve Foundation, Research Center, Territorial studies for the use 11 and conservation of biodiversity research group, Carrera 4 No 58–59, Bogotá, 12 Colombia. (MRP) 13 14 The Anoura geoffroyi species complex is composed of 3 large species: A. geoffroyi, A. 15 peruana, and A. carishina. Several inconsistencies arise from the description of A. 16 carishina, and given the lack of a comparison with the dentition and external characters 17 of A. latidens, here we compare the taxonomic characters of these species. To understand 18 the position of A. carishina in the morphospace occupied by large Anoura, we conducted 19 a Principal Component Analysis on 12 craniodental and 11 external variables. We 20 complement our results with further analysis of traits thought to be diagnostic for these 21 species, including 1) an elliptical Fourier transformation analysis of the shape of the third 22 upper premolar (P4), 2) a comparison of the area of the second (P3) and third (P4) upper 23 premolars, and 3) a comparison of maxillary toothrow angles. We find that A. carishina 24 is morphologically indistinguishable from A. latidens, and that there is broad overlap in 1 bioRxiv preprint doi: https://doi.org/10.1101/462051; this version posted November 5, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 25 morphology between A. latidens and A. geoffroyi. However several characters found in A. 26 latidens are lacking in A. geoffroyi, including a triangular shape to the P4 caused by a 27 medial-internal cusp enclosed by the base of the tooth, a lack of development of the 28 anterobasal cusp in the P3, a smaller braincase, and a shorter rostrum. We reassess the 29 distribution of Anoura latidens in Colombia, adding new records and correcting 30 previously-published records that were misidentified. Overall, our results suggest that a 31 stable taxonomy for the group should consider A. carishina as a junior synonym of A. 32 latidens, and that, although A. latidens is distinguishable from A. geoffroyi, further 33 genetic and taxonomic work is needed in to clarify species limits within the A. geoffroyi 34 species complex. 35 36 El complejo de especies Anoura geoffroyi se compone de 3 especies , A. geoffroyi, A. 37 peruana, y A. carishina. La descripción de Anoura carishina posee varias 38 inconsistencias, y dado que no se realizó una comparación con A. latidens, realizamos 39 una comparación de los caracteres taxonómicos de ambas especies. Para entender la 40 posición de A. carishina en el morfoespacio ocupado por los Anoura grandes realizamos 41 un Análisis de Componentes Principales usando 12 variables cráneo-dentales y 11 42 variables externas. Complementamos nuestros resultados con 1) un análisis de 43 transformación elíptica de Fourier de la forma del tercer premolar superior (P4) 2) una 44 comparación del área del segundo (P3) y tercer (P4) premolares superiores y 3) una 45 comparación de los ángulos de las hileras dentales maxilares. Encontramos que A. 46 carishina es morfológicamente indistinguible de A. latidens y que existe una amplio 47 superposición en la morfología de A. latidens y A. geoffroyi. Sin embargo, la forma del 2 bioRxiv preprint doi: https://doi.org/10.1101/462051; this version posted November 5, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 48 P4, una cúspide anterobasal no desarrollada en el P3, y caracteres relacionados con una 49 caja craneana menos inflada y un rostro corto son útiles en distinguir A. latidens de A. 50 geoffroyi. Reevaluamos la distribución de Anoura latidens en Colombia, al agregar 51 nuevos registros y corrigiendo registros previamente publicados que se encontraban mal 52 identificados. En general, nuestros resultados sugieren que una taxonomía estable para el 53 grupo debería considerar a A. carishina como un sinónimo junior de A. latidens, que A. 54 latidens es distinguible de A. geoffroyi utilizando medidas cráneo-dentales y resalta la 55 necesidad de estudios genéticos para esclarecer las relaciones filogenéticas entre A. 56 latidens y el complejo de especies A. geoffroyi. 57 58 Key words: Chiroptera, Colombia, distribution, elliptical Fourier transformation, 59 morphometry, nectarivorous bat, shape analysis 60 *Correspondent: [email protected] 61 62 63 64 65 66 67 68 69 70 3 bioRxiv preprint doi: https://doi.org/10.1101/462051; this version posted November 5, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 71 Anoura is one of the most species-rich genera in the subfamily Glossophaginae. It is 72 currently comprised of 10 species, although not all are widely accepted species (Handley 73 1984, Mantilla-Meluk & Baker 2006, Griffiths & Gardner 2007 [2008], Jarrín-V & Kunz 74 2008, Mantilla-Meluk & Baker 2010, Pacheco et al. 2018). The genus is subdivided into 75 two groups based on dental morphology and size (Allen 1898, Griffiths & Gardner 2007 76 [2008]), with five small species (A. caudifer, A. aequatoris, A. cadenai, A. fistulata and 77 A. luismanueli) and five large species (A. carishina, A. cultrata, A. geoffroyi, A. peruana 78 and A. latidens). Mantilla-Meluk and Baker (2010) designated three of these large species 79 (along with their subspecies) as the A. geoffroyi species complex, including A. carishina, 80 A. geoffroyi geoffroyi, A. geoffroyi lasiopyga and A. peruana, and also elevated A. 81 peruana to a separate species rather than a subspecies of A. geoffroyi. 82 Anoura carishina Mantilla-Meluk and Baker 2010 is only known to date from the 83 5 specimens of the type series deposited at the Mammal Collection Alberto Cadena 84 García at Instituto de Ciencias Naturales (Universidad Nacional, Bogotá, Colombia). Its 85 distribution is limited to 3 localities in the western slopes of the southern Colombian 86 Andes and the Sierra Nevada de Santa Marta, a mountain system isolated from the Andes 87 in the north of Colombia. The type ICN-14530 and paratype ICN-14531 are from 88 Taminango, Nariño department (1.67º, -77.32º). The two other localities are San Pedro de 89 La Sierra, Sierra Nevada de Santa Marta, department of Magdalena (10.90º, -74.04º) for 90 paratypes ICN-5224, 5225 and Cali, Pance, department of Valle del Cauca (3.32º, - 91 76.63º) for paratype ICN-5938. Anoura carishina was described as a large Anoura with 92 the following diagnostic characters: greatest length of skull less than 24.5 mm, small 93 canines, P4 teeth with a wide triangular base, and complete zygomatic arches (although 4 bioRxiv preprint doi: https://doi.org/10.1101/462051; this version posted November 5, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 94 they are broken in several of the type series collections; (Mantilla-Meluk & Baker 2010)). 95 However, in the description it was only explicitly compared to the subspecies of Anoura 96 geoffroyi (A. g. geoffroyi, A. g. lasiopyga) and A. peruana - it was not compared to A. 97 latidens, a species to which it bears resemblance in dental morphology, size, and 98 coloration. 99 Anoura latidens Handley 1984 is described as a large species of Anoura, 100 distinguishable from A. geoffroyi by a relatively short rostrum, an inflated braincase, 101 nearly parallel maxillary toothrows, and smaller and more robust premolars which have a 102 quadrangular appearance when viewed from above. More specifically, Handley (1984) 103 states that the third upper premolar (P4) has a medial-internal cusp enclosed in the 104 triangular base of the tooth (rather than an abruptly protruding cusp as in A. geoffroyi) 105 and that the second upper premolar (P3) possesses a reduced anterobasal cusp. The 106 holotype is from Pico Ávila, Caracas, Venezuela, and the species has been reported for at 107 least 14 localities in Venezuela (Handley 1976, 1984, Linares 1986, 1998), where it 108 occupies a variety of ecosystems with an altitudinal range from 50 to 2600 meters above 109 sea level. Outside of Venezuela A. latidens has only been registered in a handful of 110 localities in Colombia, Guyana, and Peru (Handley 1984, Linares 1998, Solari et al. 111 1999, Lim & Engstrom 2001), suggesting a wide yet discontinuous distribution. 112 In Colombia, Anoura latidens is distributed in the Andean region (eastern, central, 113 and western mountain ranges) and the inter-Andean valleys (Alberico et al.