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LOS MURCIÉLAGOS DE LAS TIERRAS BAJAS TROPICALES DE ECUADOR OCCIDENTAL
Referencia a publicación del Blgo. Jaime Salas, Jefe de Sección de Procesos de Manejo de Cuencas y MicroCuencas. Los murciélagos de las tierras bajas tropicales de Ecuador Occidental. PUBLICACIONES ESPECIALES. Museo de la Universidad Texas Tech. Número 57. 2010. Esta publicacion es el resultado de las colecciones realizadas durante las expediciones Sowell 2001 y 2004 al occidente del pais y de un trabajo minusioso tanto en el museo como en el laboratorio. Este trabajo es una contribucion util para el conocimiento y conservacion de nuestra fauna y que sirva de motivacion para la investigacion cientifica continue en nuestro lindo Ecuador.
LOS MURCIÉLAGOS DE LAS TIERRAS BAJAS TROPICALES DE ECUADOR OCCIDENTAL
Special PublicationsMuseum of Texas Tech University Number 57 25 May 2010Juan P. Carrera, Sergio Solari, Peter A. Larsen, Diego F. Alvarado, Adam D.Brown, Carlos Carrión B., J. Sebastián Tello, and Robert J. BakerBats of the Tropical Lowlands ofWestern Ecuador
iThis paper is dedicated to the memory of our dear friend and colleague René M.Fonseca, whose leadership and knowledge of the fauna of Ecuador was the foun-dation of the success of the 2001 and 2004 expeditions that are the basis for thispaper.“Para tí Fonquete donde quiera que estés, muchas gracias por todo amigo.”Editorial comment. One extension of this collaborative project included the training of local students whoshould be able to continue with this collaboration and other projects involving Ecuadorian mammals. Ecuador-ian students who have received or are currently pursuing graduate degrees subsequent to the Sowell Expeditionsinclude: Juan Pablo Carrera (completed M.A. degree in Museum Science at Texas Tech University (TTU) in2007; currently pursuing a Ph.D. with Jorge Salazar-Bravo at TTU); Tamara Enríquez (completed M.A. degreein Museum Science at TTU in 2007, Robert J. Baker (RJB), major advisor); René M. Fonseca (received a post-humous M.S. degree from TTU in 2004, directed by RJB); Raquel Marchán-Rivandeneira (M.S. degree in 2008under the supervision of RJB; currently pursuing a Ph.D. at TTU directed by Richard Strauss and RJB); MiguelPinto (M.S. degree at TTU in 2009; currently pursuing a Ph.D. at the Department of Mammalogy and SacklerInstitute for Comparative Genomics at the American Museum of Natural History, City University of New York);Juan Sebastián Tello (completed a Licenciatura at Pontificia Universidad Católica del Ecuador (PUCE) in 2005with Santiago Burneo; currently pursuing a Ph.D. at Louisiana State University directed by Richard Stevens);Diego F. Alvarado (pursuing a Ph.D. at University of Michigan with L. Lacey Knowles); and Carlos Carrión B.(completed Licenciatura in 2006 at PUCE; currently an expedition leader on the Galapagos Islands). These twoSowell expeditions also included students from other SouthAmerican countries who participated in the subsequentscientific efforts to better understand the fauna of Ecuador. These students included Sergio Solari of San Marcos’University, Peru, currently Professor at Universidad de Antioquia, Medellín, Colombia, and Federico Hoffmann,who received an M.S. from University de la Republica Montevideo, Uruguay, with Enrique Lessa, and a Ph.D. atTTU, and is currently at the University of Nebraska. Further, several U.S. undergraduate and graduate studentsparticipated in these expeditions, including Trashanda Johnson (B.S., TTU), Jana Higginbotham (M.S., TTU),Michelle Haynie (Ph.D., TTU), Rex McAliley (Ph.D., TTU), Joel Brant (Ph.D., TTU), Deidre Parish (Ph.D.,TTU), Marcy Revelez (M.S., TTU), Peter Larsen (M.S., TTU; currently pursing Ph.D., TTU), Carl Dick (Ph.D.,TTU), and Adam Brown (M.S., TTU; currently pursuing Ph.D., University of Texas Health Science Center atSan Antonio). The educational foundation of these individuals was undoubtedly strengthened by the culturaland biodiversity experiences encountered during the Sowell Expeditions. At the institutional level, half of thecurated collections, each recognized holotype, and a complete set of tissues are deposited at the Museo de Zoología(QCAZ) of the Pontificia Universidad Católica del Ecuador, where fellow students and professionals can studythem. Other benefits resulting from the Sowell Expeditions include publications with Ecuadorian colleagues,seminars presented to students and faculty at PUCE, environmental education and presentations to local schoolgroups (front cover), media coverage in Ecuadorian newspapers and television networks, and assisting Ecuador-ians in publishing a perspective of the Museo de Zoología Mammal Collection (e.g. Camacho and Burneo 2009).From Texas Tech’s perspective this work has been a positive experience and we thank all of those involved.Robert J. BakerSeries EditorFront cover: René M. Fonseca teaching local people in Manglares Churute Ecological Reserve about thediversity of Ecuadorian bats and their importance to the environment. Photo by Robert J. Baker.
ii2004 Researchers (from left to right): Rear: Carlos Carrión, Peter Larsen, Robert Baker, CarlDick. Front: Juan Pablo Carrera, J. Sebastián Tello, Sergio Solari, Adam Brown, and RenéFonseca. Photo by staff member.2001 Researchers (from left to right): Standing: Trashanda Johnson, Jana Higginbotham,Federico Hoffmann, Michelle Haynie, René Fonseca, Juan Pablo Carrera, Rex McAliley, JoelBrant, Deidre Parish, Sandy Tolan, Marcy Revelez. Seated: Clyde Jones, Robert Baker, CarletonPhillips. Photo by Linda Richardson.
Museum of Texas Tech University and Pontificia Universidad Católica del EcuadorJuan P. Carrera, Sergio Solari, Peter A. Larsen, Diego F. Alvarado, Adam D. Brown,Carlos Carrión B., J. Sebastián Tello, and Robert J. BakerSpecial PublicationsMuseum of Texas Tech UniversityNumber 57Bats of the Tropical Lowlands of Western Ecuador
Layout and Design: Lisa BradleyCover Design: Juan P. Carrera and Robert J. BakerProduction Editor: Lisa BradleyCopyright 2010, Museum of Texas Tech UniversityThis publication is available free of charge in PDF format from the website of the Natural Sciences ResearchLaboratory, Museum of Texas Tech University (nsrl.ttu.edu). The authors and the Museum of Texas TechUniversity hereby grant permission to interested parties to download or print this publication for personalor educational (not for profit) use. Re-publication of any part of this paper in other works is not permittedwithout prior written permission of the Museum of Texas Tech University.This book was set in Times New Roman and printed on acid-free paper that meets the guidelines for per-manence and durability of the Committee on Production Guidelines for Book Longevity of the Council onLibrary Resources.Printed: 25 May 2010Library of Congress Cataloging-in-Publication DataSpecial Publications of the Museum of Texas Tech University, Number 57Series Editor: Robert J. BakerBats of the Tropical Lowlands of Western EcuadorJuan P. Carrera, Sergio Solari, Peter A. Larsen, Diego F. Alvarado, Adam D. Brown, Carlos Carrión B.,J. Sebastián Tello, and Robert J. BakerISSN 0169-0237ISBN 1-929330-21-9ISBN13 978-1-929330-21-8Museum of Texas Tech UniversityLubbock, TX 79409-3191 USA(806)742-2442
1Bats of the Tropical Lowlands of Western EcuadorJuan P. Carrera, Sergio Solari, Peter A. Larsen, Diego F. Alvarado, Adam D.Brown, Carlos Carrión B., J. Sebastián Tello, and Robert J. BakerAbstractIn 2001 and 2004, Sowell Expeditions by Texas Tech University (TTU), in col-laboration with the Pontificia Universidad Católica del Ecuador (PUCE), were carriedout to document the bat fauna west of the EcuadorianAndes. This region of continentalEcuador possesses a unique combination of habitats within the Neotropics due to theconfluence of the Chocó–Darién humid forest and the Peruvian–Chilean desert forma-tion. These features combine to produce high levels of species richness and endemismfor several taxa, including bats. A total of 1,580 specimens were collected from 16localities, providing biological and distributional data for 66 species from six families:Emballonuridae, Phyllostomidae, Noctilionidae, Thyropteridae, Vespertilionidae, andMolossidae. Previously unrecognized species in the genera Lophostoma (Baker et al.2004), Micronycteris (Fonseca et al. 2007), Eumops (Baker et al. 2009), and Thyropterawere identified based on specimens collected during these expeditions. Also, newgeographic records for Anoura aequatoris, A. cultrata, Diaemus youngi, Lophostomabrasiliensis, Macrophyllum macrophyllum, Trinycteris nicefori, and Myotis simus wererecorded for western Ecuador.Key words: Andes Mountains, Chiroptera, Ecuador, natural history, systematicsResumenDurante los años 2001 y 2004 se llevaron a cabo dos expediciones Sowell al Ecua-dor con el objetivo de documentar las especies de murciélagos presentes en la vertienteoccidental de los Andes ecuatorianos en una colaboración científica entre Texas TechUniversity (TTU) y la Pontificia Universidad Católica del Ecuador (PUCE). El occidentedel Ecuador posee una combinación única de hábitats neotropicales debido a la influenciadirecta de bosques húmedos del Chocó-Darién por el norte y formaciones desérticas dePerú y Chile por el sur. Estas características han determinado una particular riquezade especies con altos niveles de endemismo para varios grupos taxonómicos, incluidoslos murciélagos. Un total de 1,580 especímenes fueron colectados en 16 localidades alo largo de la costa proporcionando información biológica y de distribución geográficapara 66 especies de seis familias: Emballonuridae, Phyllostomidae, Noctilionidae,Thyropteridae, Vespertilionidae, y Molossidae. Se reconocen nuevas especies para losgéneros Lophostoma (Baker et al. 2004), Micronycteris (Fonseca et al. 2007), Eumops(Baker et al. 2009), y Thyroptera. Además, se confirman nuevos registros geográficos deAnoura aequatoris, A. cultrata, Diaemus youngi, Lophostoma brasiliensis, Macrophyl-lum macrophyllum, Trinycteris nicefori y Myotis simus para el occidente de Ecuador.Palabras clave: Andes, Chiroptera, Ecuador, historia natural, sistemática
2 Special Publications, Museum of Texas Tech UniversityIntroductionThe tropical forests of the western versant of theEcuadorianAndes are remarkably important for issuespertaining to biodiversity and conservation (Myers etal. 2000). The western versant includes tropical rainforests of the Chocó–Darien humid ecosystem and adeciduous northern extension of the Peruvian–Chileancoastal dry forest (Hershkovitz 1958; Baker 1967;Gentry 1986, 1992; Marquet 1994). These forests arebordered on the west by the Pacific Ocean and on theeast by the Andes Mountains (Cerón et al. 1999). Thecombination of these geographic features has producedmultiple unique ecological formations in western Ecua-dor that support many endemic species (Gentry 1986;Parker and Carr 1992; Lynch and Duellman 1997;Sierra 1999; Gardner 2008 ).Western Ecuador has been categorized as a“hotspot” for conservation activities because risk ofextinction is high for many endemic species (Dodsonand Gentry 1991; Myers et al. 2000; Schipper et al.2008). The tropical region of northwestern Ecuadorhas one of the highest rates of deforestation, at ~4.0%per annum, and thousands of hectares of primary for-est have been destroyed for use by timber and oil palmindustries in the last 50 years (Dodson and Gentry1991; Myers 1993; Sierra-Maldonado 1996; Rudel1999). Only fragments of pristine tropical rain forestnear southern Colombia remain unaltered.Several surveys have focused on the richnessand endemism for groups such as plants, birds, frogs,and mammals in northwestern Ecuador (i.e., Dodsonand Gentry 1991; Parker and Carr 1992; Lynch andDuellman 1997; Jarrín-V. and Fonseca-N. 2001;Ander-son and Jarrín-V. 2002; Albuja and Mena–Valenzuela2004). The reports published by Allen (1916), Bros-set (1965), Albuja (1999), and Albuja and Mena-Valenzuela (2004) provide a baseline of informationon diversity, distribution, taxonomy, and systematicsof bats occurring west of the Ecuadorian Andes. Themost recent revisions recognized the presence of sevenfamilies, 53 genera, and 95 species of bats (Albuja andArcos 2007; Tirira 2007).Recently, several new species of bats have beendescribed from western Ecuador including Lophostomaaequatorialis (Baker et al. 2004), Lonchophylla choc-oana (Dávalos 2004), Anoura fistulata (Muchhala et al.2005), Lonchophylla orcesi (Albuja and Gardner 2005),Platyrrhinus albericoi, P. ismaeli, and P. matapalensis(Velazco 2005), Sturnira koopmanhilli (McCarthy etal. 2006), Lonchophylla cadenai (Woodman and Timm2006), Micronycteris giovanniae (Fonseca et al. 2007),Lonchophylla fornicata (Woodman 2007), Eumopswilsoni (Baker et al. 2009), and Platyrrhinus nitelinea(Velazco and Gardner 2009). Furthermore, Balantiop-teryx infusca (Emballonuridae), Artibeus fraterculus,Choeroniscus periosus, Platyrrhinus chocoensis,Rhinophylla alethina, Vampyriscus nymphaea (Phyl-lostomidae), Amorphochilus schnablii (Furipteridae),Eptesicus innoxius, and Rhogeessa velilla (Vesper-tilionidae) are endemic to the region (Handley 1966;Baker 1967; Koopman 1982; Baud 1982; Albericoand Velasco 1991a; McCarthy et al. 2000; Baird et al.2008). The number of newly described species and theendemic nature of the bat fauna of the western Andesdocument the significance of this region’s biodiversityand the urgency for appropriate conservation measuresin this region.Herein, the results of the Sowell Expeditions con-ducted in Ecuador during 2001 and 2004 by researchers(page ii) from the Museum of Texas Tech Universityand the Museo de Zoología (QCAZ) of the PontificiaUniversidad Católica del Ecuador are reported. In-cluded is information on species diversity, taxonomy,and distribution of the bat fauna of western Ecuador.Materials and MethodsStudy site.—Two fieldtrips (2001 and 2004) wereconducted to sample bat faunas in the provinces ofEsmeraldas, Guayas, and El Oro. The first survey wasconducted from 1 to 8 August of 2001, and the secondfrom 26 June to 12 August of 2004. Sixteen siteswere sampled at localities between 10 and 1100 m in
Carrera et al.—Bats of Western Ecuador 3elevation and within several vegetation types, includ-ing primary and secondary forest, disturbed areas, andagricultural plantations (Fig. 1).Gazetteer.—Collecting localities for both fieldseasons are described below. Province, specificlocality, latitude and longitude, elevation, dates ofsampling, and a brief description of the major habitatsare provided.1. ESMERALDAS: Mataje (01°2121.4" N,78°4327.3" W; 87 m). 11 and 13 August 2004.Mataje is located in the northern region of Esmer-aldas Province near the Colombian border. The area ishumid and surrounded by primary and secondary forest.Primary forest is in greater abundance than secondaryforest and consists of palms and other large trees.2. ESMERALDAS: Estación Experimental LaChiquita (01°1355.2" N, 78°4557.7" W; 53 m). 2August 2001 and 5 August 2004.This research station is located in northern Es-meraldas Province and is managed by the Luis VargasTorres University. The area is dominated by primaryrain forest. Most of the forest of Estación ExperimentalLa Chiquita is intact, but human impact was evident inthe form of deforestation and oil palm plantations inthe surrounding areas.3. ESMERALDAS: Banana plantation near SanLorenzo (1°1531" N, 78°4651" W; 53 m). 3 August2001 and 9 August 2004.This site is located approximately 2.3 km S,5.6 km E of San Lorenzo. The surrounding habitat iscomposed of secondary forest and agricultural fieldsassociated with homesteads.4. ESMERALDAS: Finca San José (1°036" N,78°3720" W; 144 m). 6 August 2001.Finca San José is 29 km S, 24 km E of SanLorenzo near the Lita – San Lorenzo highway. Habitatis composed of agricultural crops and shrubs and a fewlarger trees surrounded by human dwellings.5. ESMERALDAS: La Guarapera (1°1613" N,78°4811" W; 44 m). 4 August 2001.La Guarapera is 1.5 km S, 3.4 km E of SanLorenzo. This area is primarily agricultural and in-cludes banana and oil palm plantations and humandwellings.6. ESMERALDAS: San Francisco de Bogotá(01°0536.8" N, 78°4221.5" W; 63 m). 27 July 2004;3–13 August 2004.San Francisco de Bogotá is a small village 21 kmS, 14 km E of San Lorenzo. The area includes primaryforest, disturbed forest, and various types of crop andfruit plantations. Active logging of adjacent forestswas occurring during our period of study.7. GUAYAS: Bosque Protector Cerro Blanco(02°1047.6" S, 80°0117.7" W; 22 m). 4–9 July2004.Bosque Protector Cerro Blanco is approximately8 km W of the city of Guayaquil. The FundaciónProbosque manages this private reserve. This area isa remnant of tropical dry forest consisting of primaryand secondary forest.8. GUAYAS: Reserva Ecológica ManglaresChurute (02°2716.9" S, 79°3738.6" W; 50 m). 30June–3 July 2004.The Ministerio delAmbiente of Ecuador managesthis national reserve. The area is dominated by trees,shrubs, and mangroves, as well as banana, papaya, andmango plantations.9. GUAYAS: Isla Puná (02°4534.3" S,79°5501.5" W; 10 m). 11–13 July 2004.Isla Puná is in the Gulf of Guayaquil. The habitatconsists primarily of agricultural crops, shrubs, andsecondary dry forest.10. ELORO:FuerteMilitarArenillas(03°3848.9"S, 80°0940.8" W; 64 m). 15–20 July 2004.
4 Special Publications, Museum of Texas Tech UniversityFigure 1. Map of Ecuador illustrating 16 sampling localities for bats collected in 2001 and 2004during the Sowell Expeditions. Numbers correspond to the localities in the gazetteer. The lightgrey lines running north-south are isolines that represent the boundaries of the Andes Mountains.AndesMountains
Carrera et al.—Bats of Western Ecuador 5Fuerte Militar Arenillas is near the Peruvianborder. The area is dry and surrounded by primary dryforest, secondary forest, and plantations of crops.11. EL ORO: Bosque Petrificado de Puyango(03°5303.4" S, 80°0441.1" W; 280 m). 21–24 July2004.This forest reserve is in southern Oro Provincenear the border with Loja Province. The Ministerio delAmbiente of Ecuador manages the reserve. The area iscomprised of primary and secondary dry forest.12. EL ORO: Jardín Botánico Moromoro(03°3945.2"S, 79°4441.2" W; 908 m). 30 July 2004.The Jardín Botánico Moromoro Reserve is locat-ed between Machala and Zaruma in the Piñas district.This cloud forest is near the Buenaventura EcologicalReserve (managed by the Fundación Jocotoco) and ismaintained to protect existing primary forest. The habi-tat within the Jardín Botánico Moromoro Reserve in-cludes primary forest, palms, epiphytes, and shrubs.13. EL ORO: Cerro Chiche (03°4559.7" S,79°3850.9" W; 650 m). 25 and 27 July 2004.Cerro Chiche is located in the southern area ofthe province near the border between Portovelo andPiñas districts. The area is dry and consists of disturbedforest, shrubs, and agricultural crops.14. ELORO: Zaruma(03°4123.1" S, 79°3543.7"W; 1,106 m). 26, 29, and 31 July 2004.Zaruma is the highest region sampled during ourstudy. The area includes numerous gold and silvermines. Vegetation is disturbed forest and agriculturalcrops, including banana plantations.15. EL ORO: Presa Tahuín (03°3728" S,80°0007" W; 125 m). 18 July 2004.Presa Tahuín is a dam and associated body of wa-ter located in the Arenillas River basin in the southernregion of El Oro. The lake generates electricity andprovides water for irrigation in the southern region ofthe province and northern Peru. The area is surroundedby disturbed vegetation.16. EL ORO: Portovelo–Quinta Palomares(03°4411.4" S, 79°3541.1" W; 633 m). 27 July2004.This site is located south of the El Tablón area andincludes gold mines and streams. Vegetation consistsof disturbed forest, fruit plantations, and shrubs.Field methods.—Bats were captured at night us-ing mist nets set in flyways and by hand from diurnalroosts. On average, 10–12 nets were set each night at1700–1730h and closed at midnight. Different habitattypes were sampled, including both primary forest andmodified habitats (secondary growth, disturbed forest,and plantations). Bats also were captured from roostslocated in hollow trees, rock crevices, mines, culverts,and abandoned houses. Captured bats were placed inclean, individual cloth bags to prevent parasite transfer.Ectoparasites were removed from the fur and wingsof each individual immediately after euthanization.Voucher specimens were numbered and preserved asskin, skull, and skeleton or fluid preparations (Handley1988). Standard field measurements were recorded foreach specimen. Tissues for molecular studies werepreserved in lysis buffer (Longmire et al. 1997) or 95%ethanol, and blood samples were collected using No-buto blood filter strips (Advantec Manufacturing Inc.,California). As time permitted, representative speci-mens were karyotyped for each species (Baker et al.2003a). Animals were captured and handled followingthe standards of theAmerican Society of Mammalogists(Gannon et al. 2007) and Texas Tech Animal Care andUse Committee (Permit # 02217-02).Species identification.—In the field, each speci-men was tentatively identified to species based onexternal morphology. All voucher specimens were thenbrought to the Natural Science Research Laboratory(NSRL) of the Museum of Texas Tech University tobe positively identified based on external characters,cranial morphology, and selected cranial and dentalmeasurements. Further, genetic analyses were per-formed to confirm the identity of some specimens(Hoffmann and Baker 2001, 2003; Baker et al. 2004;
6 Special Publications, Museum of Texas Tech UniversityHoofer and Baker 2006; Larsen et al. 2007; Fonseca etal. 2007; Porter et al. 2007; Baker et al. 2009; Solari etal. 2009). After specimen identification, all holotypespecimens, one half of the paratype voucher specimens,and tissue samples from all specimens were depositedat the Museo de Zoologia of the Pontificia Universi-dad Católica del Ecuador. The remaining paratypevoucher specimens, as well as tissue samples from allspecimens, were deposited at the NSRL.When a specific taxon was found to be distinct atthe genetic or morphological level, we verified whetherits morphological diagnosis and/or geographic distri-bution agreed with a previously described taxon (i.e.,subspecies or junior synonyms proposed for westernSouthAmerican or MiddleAmerican populations of thespecies). When no agreement existed, we hypothesizedthat this may represent an undescribed species. Theseassessments and their taxonomic outcome are discussedin the “Remarks” section of each species account.ResultsA total of 1,580 specimens were obtained fromthe western side of the Andes Mountains in Ecuador(Table 1). These records increase the number of speciesknown from the western side of the Ecuadorian Andesfrom 95 (see Albuja and Arcos 2007; Tirira 2007) to106. New geographic records were recognized hereinfor Anoura aequatoris, A. cultrata, and Myotis simus insouthwestern Ecuador, whereas Diaemus youngi (Pintoet al. 2007), Lophostoma brasiliensis, Macrophyllummacrophyllum, and Trinycteris nicefori were from thenorthwest. Other taxa including Dermanura rosen-bergi, D. rava, and Rhogeessa velilla were elevatedfrom subspecies to species status based primarily ongenetic analyses of DNA sequences of specimenscollected during this study (Baird et al. 2008; Solariet al. 2009). Additional noteworthy results includedthe discoveries of new species of Lophostoma (Bakeret al. 2004), Micronycteris (Fonseca et al. 2007), andEumops (Baker et al. 2009), as well as a Thryopterathat remains undescribed.The taxonomic arrangement of families presentedherein follows Teeling et al. (2005). Within the fam-ily Phyllostomidae, taxa are arranged by subfamilyand tribe following Baker et al. (2003b). Within theVespertilionidae, subfamilies follow Hoofer and VanDen Bussche (2003). Within families, subfamilies, andtribes, genera and species are listed alphabetically. Foreach species, we include species name with authorityand date followed by the specific collecting localities(and number of specimens), as well as relevant remarksregarding geographic or taxonomic information andecology.Family EmballonuridaePeropteryx kappleri Peters 1867Specimens collected (4♀, 1♂).—Bosque Petrifi-cado de Puyango (5).Remarks.—Peropteryx kappleri was described byPeters in 1867 based on material from Suriname. Werefer our specimens, and all others from Ecuador, tothe nominate form P. k. kappleri (Hood and Gardner2008 ). This species is uncommonly collected inEcuador and limited to the western side of the Andes,where it is known from El Oro Province (Sanborn 1937;Albuja 1999). Albuja and Mena-Valenzuela (2004) alsoreported specimens from Carchi Province (01°0220" N,78°1630" W; 900 m). Our specimens were collectedfrom a small crevice in a hill (280 m) near the bridgethat connects El Oro and Loja Provinces, inside thePuyango Reserve. Desmodus rotundus also was pres-ent in this roost.Saccopteryx bilineata (Temmink 1838)(Plate 1A)Specimens collected (13♀, 6♂).—San Franciscode Bogotá (1), Bosque Protector Cerro Blanco (1),Reserva Ecológica Manglares Churute (11), and PresaTahuín (6).Remarks.—Saccopteryx bilineata previously wasreported from western Ecuador in the provinces of Es-meraldas and Guayas (Allen 1916; Sanborn 1937; Bros-set 1965; Albuja and Mena-Valenzuela 2004; Salas-Z.
Carrera et al.—Bats of Western Ecuador 7NorthernLocalitiesCentralLocalitiesSouthernLocalitiesLOCALITY12345678910111213141516TOTALEMBALLONURIDAE Peropteryxkappleri55Saccopteryxbilineata1111619PHYLLOSTOMIDAE MICRONYCTERINAE Micronycterisgiovanniae11Micronycterishirsuta2428Micronycterismegalotis4127621124Micronycterisminuta123DESMODONTINAE Desmodusrotundus322514477163Diaemusyoungi11LONCHORHININAE Lonchorhinaaurita11PHYLLOSTOMINAE Macrophyllummacrophyllum22Trachopscirrhossus2114Lophostomaaequatorialis133963732Lophostomabrasiliense33Mimoncrenulatum512816Phyllodermastenops2114Phyllostomusdiscolor554319316257Phyllostomuselongatus551626Phyllostomushastatus38718743Tonatiasaurophila371920Table1.SpeciescollectedandlocalitiessampledduringtheEcuadorianSowellExpeditions.(1)Mataje,(2)EstaciónExperimentalLaChiquita,(3)bananaplantationnearSanLorenzo,(4)FincaSanJosé,(5)LaGuarapera,(6)SanFranciscodeBogotá,(7)BosqueProtectorCerroBlanco,(8)ReservaEcológicaManglaresChurute,(9)IslaPuná,(10)FuerteMilitarArenillas,(11)BosquePetrificadodePuyango,(12)JardínBotánicoMoromoro,(13)CerroChiche,(14)Zaruma,(15)PresaTahuín,and(16)Portovelo–QuintaPalomares.
8 Special Publications, Museum of Texas Tech UniversityNorthernLocalitiesCentralLocalitiesSouthernLocalitiesLOCALITY12345678910111213141516TOTALChrotopterusauritus11Vampyrumspectrum66GLOSSOPHAGINAE Glossophagasoricina833122221115397Anouraaequatoris11Anouracultrata11Lichonycterisobscura11LONCHOPHYLLINAE Lonchophyllaconcava2333516Lonchophyllarobusta3159Lonchophyllathomasi14611CAROLLIINAE Carolliabrevicauda226221815318178Carolliacastanea14123240Carolliaperspicillata222172245GLYPHONYCTERINAE Trinycterisnicefori11RHINOPHYLLINAE Rhinophyllaalethina244212STERNODERMATINAE Sturniralilium1117182192321277Sturniraluisi1111214135130Sturniraoporaphilum22Artibeusfraterculus523332924143118177Artibeusjamaicensis121471638622132253149Artibeuslituratus3322310532639Chirodermatrinitatum157215Chirodermavillosum9211Table1.(cont.)
Carrera et al.—Bats of Western Ecuador 9Table1.(cont.)NorthernLocalitiesCentralLocalitiesSouthernLocalitiesLOCALITY12345678910111213141516TOTALDermanurarava13911714146Dermanurarosenbergi115125345Mesophyllamacconnelli11Platyrrhinuschocoensis1124Platyrrhinusdorsalis11321228Platyrrhinushelleri21216Platyrrhinusmatapalensis2361315Platyrrhinusnigellus44Urodermabilobatum51513226Vampyressathyone3216Vampyriscusnymphaea333918NOCTILIONIDAE Noctilioleporinus2125THYROPTERIDAE Thyropterasp.2125VESPERTILIONIDAE MYOTINAE Myotisalbescens144162119Myotisnigricans24410822192365Myotisriparius5222314Myotissimus22318VESPERTILIONINAE Eptesicuschiriquinus2125Eptesicusinnoxius681321131Lasiurusblossevillii41117Lasiurusega123Rhogeessavelilla6219
10 Special Publications, Museum of Texas Tech UniversityNorthernLocalitiesCentralLocalitiesSouthernLocalitiesLOCALITY12345678910111213141516TOTALMOLOSSIDAE Eumopswilsoni718Molossusbondae1910Molossusmolossus4304341TOTAL411276423462841092711151521152255876631580Table1.(cont.)2008). Six specimens collected from a diurnal roost inthe tunnels of the Presa Tahuín constitute new recordsfor the El Oro Province in southwestern Ecuador. Allthe Ecuadorian populations correspond to the form S.b. bilineata (Hood and Gardner 2008 ).Family PhyllostomidaeSubfamily MicronycterinaeMicronycteris giovanniae Baker and Fonseca 2007Specimens collected (1♂).—Finca San José (1).Remarks.—Micronycteris giovanniae was de-scribed based on this single specimen from northwest-ern Ecuador. Fonseca et al. (2007) and Porter et al.(2007) provide ecological, morphological, and geneticinformation for this species as well as taxonomic datafor other species of Micronycteris in northwesternEcuador.Micronycteris hirsuta (Peters 1969)Specimens collected (4♀, 4♂).—Mataje (2),Estación Experimental La Chiquita (4), and Finca SanJosé (2).Remarks.—According to Albuja and Mena-Valenzuela (2004), M. hirsuta is rare along the westernside of the Andes Mountains. This species has beenrecorded from El Paraíso in Pichincha Province, AltoTambo and Hacienda la Granada in Esmeraldas Prov-ince, and Espuela Perdida in Manabí Province (Albuja1999; Albuja and Mena-Valenzuela 2004). Molecularanalyses based on sequence variation of mitochon-drial and nuclear genes confirmed the presence of thisspecies west of the Andes and indicated low geneticdivergence among geographic regions, including Southand CentralAmerican populations (Fonseca et al. 2007;Porter et al. 2007). Our specimens were netted in pri-mary forests of the Esmeraldas Province.Micronycteris megalotis (Gray 1842)Specimens collected (12♀, 12♂).—ChiquitaResearch Station (4), La Guarapera (1), San Franciscode Bogotá (2), Bosque Protector Cerro Blanco (7), IslaPuná (6), Bosque Petrificado de Puyango (2), JardínBotánico Moromoro (1), and Cerro Chiche (1).
Carrera et al.—Bats of Western Ecuador 11Remarks.—This species was recorded in Guaya-quil, Guayas Province, by Brosset (1965); in BosqueProtector Cerro Blanco, by Salas (2007); and in Est-ación Experimental La Chiquita, Esmeraldas Province,by Albuja (1999). Porter et al. (2007) identified a ge-netic distance of approximately 5% in the cytochrome-bgene between populations from east and west of theAndes, but this distinction has not been associatedwith any taxonomic changes to date. Micronycterismegalotis was common along the coast in primaryforest, secondary forest, disturbed areas, and planta-tions. Some individuals were collected from a culvertand others were observed roosting in a Ceiba tree andin a log on the forest floor at Bosque Protector CerroBlanco.Micronycteris minuta (Gervais 1856)(Plate 1B)Specimens collected (2♀, 1♂).—San Francisco deBogotá (1) and Bosque Protector Cerro Blanco (2).Remarks.—Micronycteris minuta previously wasrecorded exclusively in the humid forests of north-western Ecuador (Albuja 1999; Tirira 1999, 2004;Albuja and Mena-Valenzuela 2004) based on specimenscollected in Alto Tambo in Esmeraldas Province (Al-buja 1989). The records from Bosque Protector CerroBlanco increase the distribution range for this speciesto the southern part of the coast, which is concordantwith the report by Pacheco et al. (2007) from QuebradaLos Naranjos (03º5015" S, 80º1144.99" W; 550 m) inTumbes Department, Peru. Molecular analyses docu-ment a genetic distance of over 5% among geographicpopulations in SouthAmerica, which may indicate spe-cies level variation (Porter et al. 2007). Micronycterisminuta hypoleuca J.A.Allen 1900, with type locality inMagdalena, Colombia, has not been examined geneti-cally; these data will be needed to understand speciesboundaries if M. minuta as currently recognized provesto be a species complex.Subfamily DesmodontinaeDesmodus rotundus (E. Geoffroy 1810)Specimens collected (29♀, 34♂).—San Franciscode Bogotá (3), Reserva Ecológica Manglares Churute(22), Isla Puná (5), Fuerte Militar Arenillas (14),Bosque Petrificado de Puyango (4), Cerro Chiche (7),Zaruma (7), and Portovelo–Quinta Palomares (1).Remarks.—Desmodus rotundus was observedroosting in a small rock crevice near the Puyango Re-serve and a single specimen was caught inside a goldmine in Zaruma. Also, a colony comprised primarilyof males was collected from a small building within theReserva Ecológica Manglares Churute. Our specimenscorrespond to the subspecies D. r. murinus J.A. Wagner1840 (Kwon and Gardner 2008 ).Diaemus youngi (Jentink 1893)Specimens collected (1♂).—Banana plantationnear San Lorenzo (1).Remarks.—This specimen is the first record ofD. youngi for Ecuador (Pinto et al. 2007). Previously,Cabrera and Yepes (1940), Albuja (1999), and Tirira(2007) hypothesized that this species likely inhabitseastern Ecuador but no voucher specimen documentedsuch distribution. Pacheco et al. (2007) reported thepresence of this species from Angostura (03º4523"S,80º2315" W; 74 m) in Tumbes Department, Peru,from a single specimen, which was the first record ofthis species from western Peru. Our specimen wasnetted in a disturbed habitat surrounded by bananaplantations associated with several occupied housesand domestic animals, including chickens roosting inadjacent trees.Subfamily LonchorhininaeLonchorhina aurita Tomes 1863(Plate 1C)Specimens collected (1♀).—San Francisco deBogotá (1).Remarks.—Lonchorhina aurita has been recordedon both sides of the Andes. Anthony (1923) describedL. aurita occidentalis from Guayas Province, but Wil-liams and Genoways (2008 ) did not recognizeit as a valid subspecies. One specimen was collectedunder a bridge over a small stream in secondary forestnear a fruit plantation.
12 Special Publications, Museum of Texas Tech UniversityA. B.C.E.G.D.F.H.Plate 1. Bats collected during the Sowell Expedition 2004. A. Saccopteryx bilineata; B.Micronycteris minuta; C. Lonchorhina aurita; D. Lophostoma aequatorialis; E. Phyllodermastenops; F. Vampyrum spectrum; G. Glossophaga soricina; H. Sturnia lilium. Photos A, B, D,G, and H by Robert J. Baker. Photo C by Peter A. Larsen. Photos E and F by Carl W. Dick.
Carrera et al.—Bats of Western Ecuador 13A. B.C.E.G.D.F.H.Plate 2. Bats collected during the Sowell Expedition 2004. A. Artibeus lituratus; B. Chirodermavillosum; C. Uroderma bilobatum; D. Noctilio leporinus; E. Thyroptera sp.; F. Eptesicusinnoxius; G. Lasiurus blossevillii; H. Eumops wilsoni. PhotosA, B, C, D, F, G, and H by RobertJ. Baker. Photo E by Carl W. Dick.
14 Special Publications, Museum of Texas Tech UniversitySubfamily PhyllostominaeTribe MacrophylliniMacrophyllum macrophyllum (Schinz 1821)Specimens collected (2♂).—San Francisco deBogotá (2).Remarks.—The specimens from EsmeraldasProvince constitute the first record for this species inwestern Ecuador (Albuja 1999). Macrophyllum macro-phyllum is known from Bonda, Santa Marta, Colombia(Allen 1900), on the western side of the CordilleraOriental and its distribution extends to MiddleAmerica.This species was netted over a stream in a disturbedforest near agricultural areas.Trachops cirrhosus (Spix 1823)Specimens collected ( 2♀, 2♂).—Estación Ex-perimental La Chiquita (2), banana plantation near SanLorenzo (1), and San Francisco de Bogotá (1).Remarks.—This species was netted in primaryforest, disturbed forest, and a banana plantation, butonly one specimen (from San Francisco de Bogotá)was netted near water, where toads and frogs werecalling around the time of capture. The biology andsystematics of T. cirrhosus were reviewed by Crameret al. (2001); these authors assigned all Ecuadorianpopulations to the subspecies T. c. cirrhosus.Tribe PhyllostominiLophostoma aequatorialis Baker, Fonseca, Parish,Phillips, and Hoffmann 2004(Plate 1D)Specimens collected (18♀, 14♂).—Mataje (1),Estación Experimental La Chiquita (3), San Franciscode Bogotá (3), Reserva Ecológica Manglares Churute(9), Fuerte MilitarArenillas (6), Bosque Petrificado dePuyango (3), and Cerro Chiche (7).Remarks.—Records from Guayas and El OroProvinces increase the distributional range of L. aequa-torialis along the southern coast to considerably drierregions than from the north, where it was described.Baker et al. (2004) provided information regarding thetaxonomic status and systematics of this taxon. Thisspecies was common in primary and secondary foresthabitats within both northern humid and southern dryforests. There is a second species, Lophostoma silvi-colum occidentalis, described from the western versantof theAndes in Peru and Ecuador that is probably sisterto L. aequatorialis. Cytochrome-b gene sequencedata of several individuals collected during our studydoes not provide evidence of significant geographic orgenetic structure typical of two species in this sample.Further study of the nature of this variation within L.aequatorialis/occidentalis is merited. Cytochrome-bsequence data does indicate that whatever species-levelname(s) are ultimately applied, there is a Lophostomaspecies present in western Ecuador that is not distrib-uted east of the Andes.Lophostoma brasiliense Peters 1866Specimens collected (1♀, 2♂).—San Franciscode Bogotá (3)Remarks.—Lophostoma brasiliense has beenrecorded from eastern Ecuador (Albuja 1999; Tirira2004). The specimens collected in Esmeraldas Prov-ince extend the distribution of this species into westernEcuador. This species is considered monotypic byWilliams and Genoways (2008 ).Mimon crenulatum (E. Geoffroy 1803)Specimens collected (9♀, 7♂).—Mataje (5),Estación Experimental La Chiquita (1), La Guarapera(2), and San Francisco de Bogotá (8).Remarks.—Mimon crenulatum was relativelycommon and was found in a diverse range of ecologicalconditions along the western side of theAndes in Ecua-dor, as indicated byAlbuja (1999). Our specimens hada yellowish-cream medial stripe along the back as wellas similarly colored spots behind the ears. Pacheco etal. (2007) collected the first specimen from Peru at Que-brada Los Naranjos (03º5015" S, 80º1144.99" W; 550m) and Quebrada Angostura (03º4523" S, 80º2315"W; 74 m) in Tumbes Department. The subspecies rec-ognized for western Ecuador is M. c. keenani Handley1960 (Williams and Genoways 2008 ).
Carrera et al.—Bats of Western Ecuador 15Phylloderma stenops Peters 1865(Plate 1E)Specimens collected (4♀).—Estación Experimen-tal La Chiquita (2), banana plantation near San Lorenzo(1), and Bosque Petrificado de Puyango (1).Remarks.—This species was known from thewestern side of the Andes based on three previousrecords from Esmeraldas and Pichincha Provinces(Castro and Nolivos 1998; Trujillo and Albuja 2005).Our record from Bosque Petrificado de Puyango con-siderably extends the range of this species to the southin western Ecuador. There was considerable colorvariation within our sample of P. stenops; specimenswere gray-brown in San Lorenzo and Estación Experi-mental La Chiquita, but the specimen from Puyangowas lighter in color. The subspecies recognized forEcuadorian populations is P. s. stenops W. Peters 1865(Williams and Genoways 2008 ).Phyllostomus discolor Wagner 1843Specimens collected (22♀, 35♂).—Banana plan-tation near San Lorenzo (5), La Guarapera (5), SanFrancisco de Bogotá (4), Reserva Ecológica ManglaresChurute (3), Isla Puná (19), Fuerte MilitarArenillas (3),Cerro Chiche (16), and Zaruma (2).Remarks.—Phyllostomus discolor is abundant inthe northern humid forests on both sides of the Andes(Albuja 1999; Albuja and Mena-Valenzuela 2004;Tirira 2007). Phyllostomus discolor was common inour sample and was collected in primary forest, dis-turbed forest, and fruit plantations and was observedroosting inside an abandoned house on Isla Puná. Ourspecimens from El Oro Province extend the southerngeographic range of this species on the western side ofthe Andes in Ecuador.Phyllostomus elongatus (E. Geoffroy 1810)Specimens collected (15♀, 11♂).—EstaciónExperimental La Chiquita (5), La Guarapera (5), andSan Francisco de Bogotá (16).Remarks.—Specimens from San Francisco deBogotá were collected from a roost inside a culvert, 2km outside the village limits. Albuja (1999) lists 22specimens from Esmeraldas. P. elongatus is commonon both sides of the Andes.Phyllostomus hastatus (Pallas 1767)Specimens collected (16♀, 26♂, 1 undet.).—Est-ación Experimental La Chiquita (3), banana plantationnear San Lorenzo (8), La Guarapera (7), San Franciscode Bogotá (18), and Reserva Ecológica ManglaresChurute (7).Remarks.—Santos et al. (2003) included all Ecua-dorian populations in the subspecies P. h. panamensisJ. A. Allen (1904) with the type locality in Chiriqui,Panama. However, Williams and Genoways (2008) restricted the South American distributionof this subspecies to the western side of the AndesMountains and north and west of Lake Maracaibo inVenezuela.Tonatia saurophila Koopman and Williams 1951Specimens collected (10♀, 10♂).—Mataje (3),Estación Experimental La Chiquita (7), La Guarapera(1), and San Francisco de Bogotá (9).Remarks.—Molecular analyses using cyto-chrome-b gene sequence data showed that T. saurophilapopulations from western Ecuador were geneticallydistinct, diverging from populations in Bolivia andGuyana (Baker et al. 2004). The most appropriate namefor these western populations would be T. s. bakeri fromDarién, Panama (Williams et al. 1995). Our specimenswere collected in primary and secondary forests.Tribe VampyriniChrotopterus auritus (Peters 1856)Specimens collected (1♂).—Reserva EcológicaManglares Churute (1).Remarks.—With a previous record from Machalil-la National Park in the Manabí Province (Albuja andMuñoz 2000), our specimen constitutes the secondrecord from western Ecuador and extends the southernlimit of the range of C. auritus to Guayas Province.Pacheco et al. (2007) reported the first record of this
16 Special Publications, Museum of Texas Tech Universityspecies from western Peru from Quebrada Faical(03º4919" S, 80º1530" W; 350 m) in Tumbes Depart-ment, based on a specimen collected in primary forest.Our specimen of this uncommon bat also was collectedin primary forest.Vampyrum spectrum (Linnaeus 1758)(Plate 1F)Specimens collected (5♀, 1♂).—Isla Puná (6).Remarks.—This species was collected in SanRamón–Isla Puná, Guayas Province, by G. H. H. Tatein 1923, based on records in the American Museum ofNatural History database. Five of our six specimensof V. spectrum were collected near a large Ceiba treethat contained bird feathers and bones, suggesting thatthe tree was a feeding roost. Our specimens were col-lected during a single night and we observed at leastone additional individual flying after nets were closed.Although this species has a wide distribution along theNeotropics (Simmons 2005), Pacheco et al. (2007) re-ported it for the first time in northwestern Peru, basedon a single specimen collected in QuebradaAngostura(03º4523" S, 80º2315" W; 74 m) in Tumbes Depart-ment. This species was netted from a disturbed areanear a livestock ranch.Subfamily GlossophaginaeTribe GlossophaginiGlossophaga soricina (Pallas 1766)(Plate 1G)Specimens collected (55♀, 40♂, 2 undet.).—Bosque Protector Cerro Blanco (8), Reserva EcológicaManglares Churute (33), Isla Puná (12), Fuerte MilitarArenillas (22), Bosque Petrificado de Puyango (2),Jardín Botánico Moromoro (1), Cerro Chiche (1),Zaruma (15), and Portovelo–Quinta Palomares (3). .Remarks.—Glossophaga soricina was the mostcommon nectar-feeding bat in our collection. Indi-viduals also were collected at a gold mine in Zaruma.Although considerable pelage color variation was pres-ent across our series of specimens, cytochrome-b genesequences indicated that all specimens represented asingle species (Hoffmann and Baker 2001). It shouldbe noted that populations of G. soricina in westernEcuador were genetically distinct from conspecificsdistributed east of the Andes (Hoffmann and Baker2001). Therefore, the subspecific name G. s. valensMiller 1913 (type locality “Balsas,” Amazonas, Peru)would be available for western populations (Griffithsand Gardner 2008 ).Tribe ChoeronycteriniAnoura aequatoris (Lönnberg 1921)Specimens collected (1♂).—Jardín BotánicoMoromoro (1).Remarks.—We followed Mantilla-Meluk andBaker (2006) in recognition of Anoura aequatoris (Lön-nberg 1921), type locality in western Ecuador (Illambo,Gualea, Pichincha Province), as a valid species andspecifically distinct from A. caudifer (type locality Riode Janeiro, Brazil). This specimen constitutes the firstrecord in the southern region of the western side of theEcuadorian Andes (see Albuja 1999). It was found inthe same habitat and locality as A. cultrata (see below).Mantilla-Meluk and Baker (2009) reported A. aequato-ris for southern Colombia including localities adjacentto Ecuador in the Colombian department of Nariño.Some of the specimens identified as Anoura aequatorisby Mantilla-Meluk and Baker (2006) were previouslyreported by Cadena et al. (1998) as A. caudifer.Anoura cultrata Handley 1960Specimen collected (1♂).—Jardín BotánicoMoromoro (1).Remarks.—This specimen constitutes the firstrecord of A. cultrata for southwestern Ecuador (seeAlbuja 1999; Tirira 2004, 2007). It was captured insympatry with A. aequatoris. Cadena et al. (1998)reported this species from Nariño Department in Co-lombia, andAlbuja (1989) reported a specimen from ElPailón in Carchi Province in northwestern Ecuador.Lichonycteris obscura Thomas 1895Specimen collected (1♀).—Banana plantationnear San Lorenzo (1).Remarks.—Lichonycteris obscura has been re-ported from western Ecuador based on three specimenscollected from the Centro Científico (Scientific Center)
Carrera et al.—Bats of Western Ecuador 17Río Palenque (Pichincha Province), Sade (EsmeraldasProvince), and Manta Real (Azuay Province) (Albuja1999;Albuja and Mena-Valenzuela 2004; Griffiths andGardner 2008 ). This species appears to be lo-cally rare throughout its distribution (Hill 1985).Tribe LonchophyllinaeLonchophylla concava Goldman 1914Specimens collected (9♀, 6♂, 1 undet.).—Mataje(2), Banana plantation near San Lorenzo (3), FincaSan José (3), La Guarapera (3), and San Francisco deBogotá (5).Remarks.—Lonchophylla concava was reportedfrom Manabí Province of western Ecuador by Baker(1974). Although it was considered to represent L.mordax by Koopman (1993) andAlbuja (1999),Albujaand Gardner (2005) indicated that L. concava is thecorrect name for this species in western Ecuador, withL. mordax being restricted to eastern South America.The individuals from Mataje were collected from alarge culvert on the military base. This species alsowas collected in disturbed forest and fruit plantationsin Esmeraldas Province.Lonchophylla robusta Miller 1912Specimens collected (7♀, 2♂).—Banana planta-tion near San Lorenzo (3), La Guarapera (1), and SanFrancisco de Bogotá (5).Remarks.—Lonchophylla robusta is similar (ex-ternally) to the newly described L. chocoana Dávalos(2004), but skull and teeth characteristics clearlydistinguish these species. From preliminary observa-tions, our specimens showed slight size differencesfrom typical L. robusta from eastern Ecuador andeastern Peru.Lonchophylla thomasi J. A. Allen 1904Specimens collected (3♀, 8♂).—Banana planta-tion near San Lorenzo (1), Finca San José (4), and SanFrancisco de Bogotá (6).Remarks.—This is the smallest species of Lon-chophylla and it appears to be distributed on both sidesof theAndes. We found no conspicuous morphologicaldifferences between specimens from western Ecuadorand those from Amazonian Ecuador. However, a phy-logenetic analysis of cytochrome-b sequences (Dávalosand Jansa 2004) revealed a geographic division withinL. thomasi, but no specimens from northwestern SouthAmerica were included in that analysis. Recent mor-phological studies (Woodman and Timm 2006; Wood-man 2007) suggested L. thomasi represents a speciescomplex.Subfamily CarolliinaeCarollia brevicauda (Schinz 1821)Specimens collected (35♀, 42♂, 1 undet.).— Es-tación Experimental La Chiquita (2), San Francisco deBogotá (2), Bosque Protector Cerro Blanco (6), ReservaEcológica Manglares Churute (22), Fuerte MilitarAre-nillas (18), Bosque Petrificado de Puyango (15), JardínBotánico Moromoro (3), Cerro Chiche (1), Zaruma (8),and Portovelo–Quinta Palomares (1).Remarks.—Specimens collected in El Oro Prov-ince constitute the first voucher records for southwesternEcuador (Albuja 1999). Genetic data (Hoffmann andBaker 2003) revealed minor intraspecific differences,with 0.2 to 3.2% divergence, and grouped specimensfrom eastern and western Ecuador.Carollia castanea H. Allen 1890Specimens collected (20♀, 20♂).—EstaciónExperimental La Chiquita (14), San José Farm (1),San Francisco de Bogotá (23), and Jardín BotánicoMoromoro (2).Remarks.—This species seems to be common inwestern Ecuador, however the individuals from JardínBotánico Moromoro constitute the first vouchers fromsouthwestern Ecuador (see Albuja 1999; Albuja andMena-Valenzuela 2004). Molecular analyses basedon cytochrome-b sequences indicated high divergenceamong populations from western and eastern Ecuador(Hoffmann and Baker 2003). Subsequent analysesincluding karyotypic and morphological informationindicated that C. castanea is restricted to MiddleAmerica and northwestern South America (includingwestern Ecuador and Colombia), whereas populations
18 Special Publications, Museum of Texas Tech Universityfrom southeastern Ecuador, Peru, Bolivia and Brazilrepresented two species, C. benkeithi and one yet un-described taxon (Solari and Baker 2006).Carollia perspicillata (Linnaeus 1758)Specimens collected (24♀, 21♂).—EstaciónExperimental La Chiquita (22), La Guarapera (2), SanFrancisco de Bogotá (17), Reserva Ecológica Man-glares Churute (2), and Zaruma (2).Remarks.—Adetailed analysis using cytochrome-b gene sequence data showed a close relationshipbetween western Ecuador and MiddleAmerican popu-lations (Hoffmann and Baker 2003) of C. perspicillata.McLellan and Koopman (2008 ) suggested thatthe name C. p. azteca Saussure (1860) may apply toEcuadorian populations.Subfamily GlyphonycterinaeTrinycteris nicefori (Sanborn 1949)Specimen collected (1♀).—San Francisco deBogotá (1).Remarks.—A single specimen of T. nicefori wascollected from the primary forests of Esmeraldas Prov-ince and constitutes the first record of this species fromwestern Ecuador (see Tirira 2007). This genus previ-ously was recognized as a member of the Micronycteriscomplex (Simmons and Voss 1998).Subfamily RhinophyllinaeRhinophylla alethina Handley 1966Specimens collected (6♀, 6♂).—Mataje (2), Es-tación Experimental La Chiquita (4), Finca San José(4), and San Francisco de Bogotá (2).Remarks.—Rhinophylla alethina is endemic tothe Chocó region of Colombia and Ecuador (Handley1966;Albuja 1999) and is morphologically and geneti-cally distinct from other species of the genus (Wright etal. 1999). This species was found only in northwesternEcuador in primary forest, secondary forest, and bananaplantations.Subfamily SternodermatinaeTribe SturniriniSturnira lilium (E. Geoffroy 1810)(Plate 1H)Specimens collected (33♀, 44♂).—Finca SanJosé (1), La Guarapera (1), San Francisco de Bo-gotá (1), Bosque Protector Cerro Blanco (7), ReservaEcológica Manglares Churute (18), Isla Puná (2), FuerteMilitar Arenillas (19), Bosque Petrificado de Puyango(23), Jardín Botánico Moromoro (2), Cerro Chiche (1),and Zaruma (2).Remarks.—This is a common and widely distrib-uted species. Although distinct phylogroups have beenrecognized within S. lilium (e.g., Ditchfield 2000), nosubspecific names (see Jones and Phillips 1976) havebeen proposed for these genetically defined groups.This species was common along the western side ofthe Ecuadorian Andes in primary and disturbed forestand fruit plantations.Sturnira luisi Davis 1980Specimens collected (18♀, 12♂).—EstaciónExperimental La Chiquita (1), banana Plantation nearSan Lorenzo (1), La Guarapera (1), San Franciscode Bogotá (12), Bosque Protector Cerro Blanco (1),Reserva Ecológica Manglares Churute (4), Isla Puná(1), Fuerte Militar Arenillas (3), Cerro Chiche (5), andPortovelo–Quinta Palomares (1).Remarks.—Sturnira luisi was common along thewestern side of the Andes in primary forest, disturbedforest, and fruit plantations. Although Simmons (2005)listed S. luisi as a valid species, based on morphologicalevidence (e.g., Simmons and Voss 1998), some authorsconsider S. luisi as conspecific with S. tildae (e.g.,Brosset and Charles-Dominique 1990).Sturnira oporaphilum (Tschudi 1844)Specimens collected (1♀, 1♂).— Jardín BotánicoMoromoro (2).Remarks.—Although a number of authors recog-nize S. oporaphilum as distinct from S. ludovici (e.g.,
Carrera et al.—Bats of Western Ecuador 19McCarthy et al. 2006), Gardner (2008a ) listsS. o. ludovici H. E. Anthony 1924 as the subspeciesoccurring in Ecuador, Colombia, and Venezuela.Sturnira oporaphilum previously was knownfrom several localities in Esmeraldas and PichinchaProvinces in western Ecuador (Albuja 1999; Albujaand Mena-Valenzuela 2004). Our records from El OroProvince considerably extend the range of this speciesto the south in an undisturbed ecosystem. Morphologi-cal traits distinguishing erythromos, bogotensis, andludovici have limited application and, as a result, thedistribution of theseAndean species from Colombia toPeru is unclear (Pacheco and Patterson 1992). Pres-ently, genetic data from properly identified specimensis not available for further resolution.Tribe StenodermatiniArtibeus fraterculus Anthony 1924Specimens collected (89♀, 88♂).—Bosque Pro-tector Cerro Blanco (5), Reserva Ecológica ManglaresChurute (23), Isla Puná (33), Fuerte Militar Arenillas(29), Bosque Petrificado de Puyango (24), Cerro Chiche(14), Zaruma (31), and Portovelo–Quinta Palomares(18).Remarks.—This species was the most commonlycollected during our study. Nine specimens werehand collected from a termite nest at a farm locatedin Reserva Ecológica Manglares Churute. Artibeusfraterculus has a limited geographic range, restrictedto the dry forests west of the Andes; however, withinthat range it is common and occupies both primary anddisturbed habitats. Specimens collected at the typelocality (Portovelo) were included in recent geneticanalyses by Larsen et al. (2007), the results of whichindicate a close relationship of A. fraterculus to MiddleAmerican species (A. hirsutus and A. inopinatus).Artibeus jamaicensis Leach 1821Specimens collected (84♀, 64♂, 1 undet.).—Mataje (12), Estación Experimental La Chiquita (14),banana plantation near San Lorenzo (7), Finca San José(1), La Guarapera (6), San Francisco de Bogotá (38),Bosque Protector Cerro Blanco (6), Reserva EcológicaManglares Churute (22), Fuerte MilitarArenillas (13),Bosque Petrificado de Puyango (22), Zaruma (5), andPortovelo–Quinta Palomares (3).Remarks.—Following the restricted definitionby Larsen et al. (2007), this species should be rare orabsent east of the Andes in South America. Geneticdata showed that populations of A. jamaicensis fromwestern Ecuador are distinct from A. jamaicensis fromCentral America (Larsen et al. 2007). The appropriatename for these populations would be A. j. aequatorialisAndersen 1906 with type locality Zaruma, Ecuador.Artibeus lituratus (Olfers 1818)(Plate 2A)Specimens collected (18♀, 20♂, 1 undet.).—Mataje (3), Estación Experimental La Chiquita (3), LaGuarapera (2), San Francisco de Bogotá (2), BosqueProtector Cerro Blanco (3), Reserva Ecológica Man-glares Churute (10), Isla Puná (5), Fuerte MilitarArenillas (3), Zaruma (2), and Portovelo–QuintaPalomares (6).Remarks.—Within our sample, this is the onlyspecies of large Artibeus found on both sides of theAndes. In the field, A. lituratus may be confused withA. jamaicensis on the western side of the Andes; how-ever, A. lituratus is larger than A. jamaicensis and thefacial stripes are more distinct in A. lituratus. Somedifferences in skull and dentition also distinguish thesespecies (see Andersen 1908). Additional genetic dataare available in Larsen et al. (2007)Chiroderma trinitatum Goodwin 1958Specimens collected (12♀, 3♂).—Mataje (1),and Estación Experimental La Chiquita (5), bananaplantation near San Lorenzo (7), and San Franciscode Bogotá (2).Remarks.—This species is the smallest in thegenus and it is sometimes confused with species ofVampyriscus (based on external morphology). Geneticdata for C. trinitatum published by Baker et al. (1994)indicated a sister relationship to C. doriae (the largestspecies in the genus) from eastern and southern Braziland Paraguay. Northwestern populations correspondto the subspecies C. t. gorgasi Handley 1960 (Gardner2008b ).
20 Special Publications, Museum of Texas Tech UniversityChiroderma villosum Peters 1860(Plate 2B)Specimens collected (2♀, 9♂).—Bosque Pro-tector Cerro Blanco (9) and Bosque Petrificado dePuyango (2).Remarks.—Specimens collected in Guayas andEl Oro Provinces represent an extension of the geo-graphic range to southwestern Ecuador (see Albuja1999; Albuja and Mena-Valenzuela 2004). Chiro-derma villosum was netted in primary forest and fruitplantations from central and southern coastal Ecuador.The western populations from Ecuador and Colombiawere assigned to the subspecies C. v. jesupi Allen 1900(Gardner 2008b ).Dermanura rava Miller 1902Specimens collected (22♀, 24♂).—Mataje (1),Estación Experimental La Chiquita (3), banana planta-tion near San Lorenzo (9), La Guarapera (1), San Fran-cisco de Bogotá (17), Reserva Ecológica ManglaresChurute (14), and Jardín Botánico Moromoro (1).Remarks.—Dermanura rava is distinguishedfrom D. phaeotis by Solari et al. (2009) based on ge-netic and morphological data. The northern localitieswhere our specimens were collected are near the typelocality of San Javier, Esmeraldas (Miller 1902). Thisspecies has a broad geographic range, extending fromsouthwestern Ecuador to Panama, and inhabits dry andwet forests, although it is more commonly collected indry forests.Dermanura rosenbergi Thomas 1897Specimens collected (16 ♀, 29 ♂).—Mataje (1),Estación Experimental La Chiquita (15), banana plan-tation near San Lorenzo (1), San Francisco de Bogotá(25), and Jardín Botánico Moromoro (3).Remarks.—Dermanura rosenbergi is distin-guished from D. glauca by Solari et al. (2009) based ongenetic and morphological data. The northern locali-ties where our specimens were collected are near thetype locality of Cachavi, Esmeraldas (Thomas 1897).Dermanura rosenbergi is restricted to the wet forestsof the Chocó, with an apparent isolated population inEl Oro Province.Mesophylla macconnelli Thomas 1901Specimen collected (1♂).—Estación Experimen-tal La Chiquita (1).Remarks.—Asingle specimen was collected fromsecondary forest mixed with agricultural areas of fruitcrops. This species has been associated with Ectophyllaand Vampyressa, but genetic data (chromosomes andDNA sequences) document it is more closely relatedto Vampyressa (sensu Baker et al. 2003b and Hooferand Baker 2006).Platyrrhinus chocoensis Alberico and Velasco 1991Specimens collected (2♀, 1♂, 1 undet.).—Es-tación Experimental La Chiquita (1), San José Farm(1), and San Francisco de Bogotá (2).Remarks.—Platyrrhinus chocoensis is endemicto the Chocó region from Colombia and northwesternEcuador (Velazco 2005). Although externally similarto P. dorsalis, P. chocoensis is distinguished from thatspecies by external and cranial characters (Albericoand Velasco 1991b).Platyrrhinus dorsalis (Thomas 1900)Specimens collected (16♀, 12♂).—EstaciónExperimental La Chiquita (1), banana plantation nearSan Lorenzo (1), Finca San José (3), San Francisco deBogotá (21), and Zaruma (2).Remarks.—The only previous record of P. dor-salis for El Oro Province was from Mina Miranda,almost 1.5 km. N Zaruma (Gardner and Carter 1972).Two individuals caught in culverts near San Franciscode Bogotá showed an interesting pale coloration. Ve-lazco and Patterson (2008) included cytochrome-b,ND2, D-Loop, and RAG 2 sequences from seven ofour specimens from western Ecuador in their reviewof the genus.
Carrera et al.—Bats of Western Ecuador 21Platyrrhinus helleri Peters 1866Specimens collected (2♀, 4♂).—Estación Ex-perimental La Chiquita (2), banana plantation nearSan Lorenzo (1), Finca San José (2), and La Guarapera(1).Remarks.—This species previously had beenrecorded in southwestern Colombia from QuebradaDocordo in Chocó Department, Quebrada Huanquiin Cauca Department, and Rio Sabaletas in Valle delCauca Department (Velazco 2005). Our specimensfrom Esmeraldas Province increase the range of P.helleri to the southern Chocoan region and confirmthe presence of this species in northwestern Ecuador.It appears that this species is restricted, in Ecuador, tothe humid Pacific lowlands, contrary to the situationwith P. matapalensis (see next account).Platyrrhinus matapalensis Velazco 2005Specimens collected (8♀, 7♂).—San Franciscode Bogotá (2), Bosque Protector Cerro Blanco (3),Reserva Ecológica Manglares Churute (6), FuerteMilitar Arenillas (1), and Bosque Petrificado de Puy-ango (3).Remarks.—Velazco (2005) described this spe-cies based on specimens from Matapalo, ZarumillaProvince, Tumbes Department, in northern Peru. Thisspecies is widely distributed along the westernAndeanslopes from northern Peru to Esmeraldas Province inEcuador. Velazco and Patterson (2008) included cyto-chrome-b, ND2, D-Loop, and RAG 2 sequences fromfour of our specimens from southwestern Ecuador intheir molecular analysis of the genus, and hypothesizeda sister relationship between P. matapalensis and Cen-tral American specimens of P. helleri. These westernEcuadorian populations previously were recognized asconspecific with P. helleri (Ferrel and Wilson 1991).Platyrrhinus nigellus (Gardner and Carter 1972)Specimens collected (1♀, 3♂).— Jardín BotánicoMoromoro (4).Remarks.—Platyrrhinus nigellus is intermediatein size between P. dorsalis and P. matapalensis (Velazco2005) and is found primarily in lower montane forests.These bats have darker, almost black, dorsal hair and awhitish mid-dorsal stripe. All of our specimens werecollected from an isolated hill in Moromoro, where oth-er “montane” species were netted. Recently, Velazcoand Gardner (2009) described P. nitelinea from Puentede Moromoro, El Oro Province (03˚44 S, 79˚44 W)based on morphological characters. This new speciesseems to be a larger Platyrrhinus and it is distributedfrom western Colombia to southwestern Ecuador.Uroderma bilobatum Peters 1866(Plate 2C)Specimens collected (12♀, 14♂).—Bananaplantation near San Lorenzo (5), La Guarapera (1),San Francisco de Bogotá (5), Bosque Protector CerroBlanco (13), and Portovelo–Quinta Palomares (2).Remarks.—Although U. bilobatum is common onboth sides of theAndes, no previous records are knownfrom El Oro Province (Albuja 1999;Albuja and Mena-Valenzuela 2004). The records from Portovelo extendthe range of this species to the coastal southwest. Thisspecies previously was reported from Esmeraldas Prov-ince byAllen (1916), Brosset (1965), and Hoffmann etal. (2003). Uroderma bilobatum has been studied ex-tensively using karyotypes (Baker et al. 1972; Baker etal 1975; Baker 1979), morphology (Baker et al. 1972),and gene sequences (Hoffmann et al. 2003), and manyalternative evolutionary scenarios have been debated(Greenbaum 1981; Barton 1982; Lessa 1990; Owenand Baker 2001). The material from the western sideof the Andes in northern South America further docu-ments the complexity of these interpretations. Davis(1968) reported a major morphological break betweenpopulations from Colombia and Ecuador, and suggestedthat CentralAmerican specimens were associated withthose from Colombia, for which the name would be U.b. convexum. On the other hand, he assigned Ecuador-ian specimens from the western side of the Andes toU. b. thomasi, and further indicated that this similaritybetween thomasi-sized specimens from the eastern sideof the Andes and those from the dry deciduous foreston the western side of the Andes documented previ-ous trans-Andean dispersal. Hoffmann et al. (2003)indicated that specimens of U. bilobatum from westernEcuador were genetically indistinguishable (karyotypes
22 Special Publications, Museum of Texas Tech UniversityFamily NoctilionidaeNoctilio leporinus (Linnaeus 1758)(Plate 2D)Specimens collected (3♀, 2♂).—Bosque Protec-tor Cerro Blanco (2), Reserva Ecológica ManglaresChurute (1), and Fuerte Militar Arenillas (2).Remarks.—This species has been recorded inthe Bosque Protector Cerro Blanco (Angel and Salas2003;Alava and Carvajal 2004; Salas-Z. 2008). Basedon their geographic distribution (Davis 1973; Gardner2008d ), the western populations of Colombiaand Ecuador may correspond to the subspecies masti-vus Vahl 1797. Genetic data (Lewis-Oritt et al. 2001)indicate that this fish-eating bat is recently evolvedand little geographic genetic variation distinguishesthe widespread population from the Antilles, CentralAmerica, andAmazonian regions. Pacheco et al. (2007)reported this bat from Quebrada Faical (03º4919" S,80º1530" W; 350 m) in Tumbes Department, confirm-ing the presence of this species on the western side ofthe Peruvian Andes.Family ThyropteridaeThyroptera sp.(Plate 2E)Specimens collected (3♀, 2♂).—Banana planta-tion near San Lorenzo (2), La Guarapera (1), and SanFrancisco de Bogotá (2).Remarks.—These specimens were morphologi-cally similar to T. tricolor (Solari et al. 2004), but someadditional traits appear unique and merit further study.Applicability of the name albigula (G. M. Allen 1923)needs to be evaluated. Although typically associatedwith vegetation near streams, all of our specimens werenetted in secondary forest and crop plantations.Family VespertilionidaeSubfamily MyotinaeMyotis albescens (E. Geoffroy 1806)Specimens collected (11♀, 8♂).—Banana plan-tation near San Lorenzo (1), La Guarapera (4), SanFrancisco de Bogotá (4), Reserva Ecológica Manglaresand cytochrome-b sequences) from Middle Americanpopulations (as far north as Honduras) and assignedthese specimens to U. b. convexum. What is unclearin Davis’s work is whether the material that was thebasis of his conclusions for characteristics of Colom-bian U. bilobatum contained sufficient material fromthe Chocó to define the boundaries of morphologicalchange. Clearly, additional study of the genetic andmorphological characteristics of U. bilobatum fromthe western side of the Andes is merited.Vampyressa thyone Thomas 1909Specimens collected (6♂).—Mataje (3), SanFrancisco de Bogotá (2), and Jardín Botánico Moro-moro (1).Remarks.—Previously reported from GuayasProvince, including the original type locality (Thomas1909a),EsmeraldasProvince(Albuja1989),andPichin-cha Province (Lee et al. 2006). The genus Vampyressawas restricted to include V. melissa, V. pusilla, and V.thyone (Hoofer and Baker 2006). This taxon has beenlisted as a junior synonym of pusilla (Petterson 1968),however Lim et al. (2003) concluded that thyone andpusilla should be accorded specific status.Vampyriscus nymphaea (Thomas 1909)Specimens collected (11♀, 7♂).—Mataje (3),Estación Experimental La Chiquita (3), banana plan-tation near San Lorenzo (3), and San Francisco deBogotá (9).Remarks.—Vampyriscus nymphaea is foundprimarily in the Chocó region of northwestern SouthAmerica that extends to Central America (Albuja andMena-Valenzuela 2004). Most of our specimens camefrom secondary forest near agricultural plots, but somewere collected from patches of primary forest. Molecu-lar data for mitochondrial cytochrome-b and ND3-ND4genes and the nuclear RAG-2 gene confirmed that thisspecies, along with V. bidens, does not form a mono-phyletic lineage with Vampyressa. Thus, to create amonophyletic genus, the name Vampyriscus shouldbe recognized for these species (Baker et al. 2003b;Hoofer and Baker 2006).
Carrera et al.—Bats of Western Ecuador 23Churute (1), Fuerte Militar Arenillas (6), BosquePetrificado de Puyango (2), and Portovelo–QuintaPalomares (1).Remarks.—Morphological variation has beenrecorded in M. albescens, which occurs on both sidesof the Andes (LaVal 1973). The type locality of M.albescens is in Paraguay and therefore if the materialfrom western Ecuador is deemed conspecific, it wouldbe a remarkable range for a small Vespertilionid.Myotis nigricans (Schinz 1821)Specimens collected (17♀, 47♂, 1 undet.).—Mataje (2), San Francisco de Bogotá (4), Bosque Pro-tector Cerro Blanco (4), Reserva Ecológica ManglaresChurute (10), Isla Puná (8), Fuerte MilitarArenillas (2),Bosque Petrificado de Puyango (2), Cerro Chiche (1),Zaruma (9), and Portovelo–Quinta Palomares (23).Remarks.—Several specific names have beenproposed for populations from northwestern SouthAmerica (reviewed in LaVal 1973). Baker (1974) re-ported M. n. nigricans from Esmeraldas Province, butKoopman (1978) used the combination M. n. punensisfor specimens from northwestern Peru, and suggestedthat populations on either side of the Andes were dis-tinct from one another.Myotis riparius Handley 1960Specimens collected (6♀, 8♂).—Estación Ex-perimental La Chiquita (5), Bosque Protector CerroBlanco (2), Fuerte Militar Arenillas (2), Zaruma (2),and Portovelo–Quinta Palomares (3).Remarks.—Our analyses of mitochondrial cy-tochrome-b sequences contained minimal divergencewithin the geographic localities listed above. Our fieldidentifications included a number of individuals mis-identified as M. nigricans, however the divergence froma specimen of M. nigricans from near the type localityin Brazil indicates it is improbable that the specimenslisted above are referable to M. nigricans. This specieswas reported by Pacheco et al. (2007) based on sixspecimens collected in Angostura, Quebrada CampoVerde (03º5044" S, 80º1111" W; 570 m), Carrizalillo(03º4356.71" S, 80º1110.42"W; 125 m), and QuebradaLos Naranjos (03º5015" S, 80º1144.99" W; 550 m) inTumbes Department, northwestern Peru.Myotis simus O. Thomas 1901Specimens collected (6♀, 2♂).—Bosque Pro-tector Cerro Blanco (2), Fuerte Militar Arenillas (2),Bosque Petrificado de Puyango (3), and Portovelo–Quinta Palomares (1).Remarks.—Myotis simus is endemic to SouthAmerica and appears to be restricted to rain forestsof the Amazon Basin in Colombia, Bolivia, Brazil,Ecuador, and Peru (LaVal 1973; Wilson 2008b ).In Ecuador, this species has been collected in Limon-cocha, Sucumbios Province (00°24 S, 76°38 W) (Al-buja 1999), Río Capahuarí, Napo Province (02°18 S,77°00 W), and Boca del Río Curaray, Pastaza Province(01°22 S, 76°57 W) (LaVal 1973). Our specimensare morphologically and genetically similar to Myotissimus and, if that were the proper species identification,would constitute a new geographic record for westernEcuador. However, our specimens were collected fromdry localities, contrary with the typical habitat typerecorded for the species.Subfamily VespertilioninaeTribe NycticeiniEptesicus chiriquinus Thomas 1920Specimens collected (3♀, 2♂).—Banana planta-tion near San Lorenzo (2), La Guarapera (1), and SanFrancisco de Bogotá (2).Remarks.—Simmons (2005) recognized this spe-cies as valid for Ecuadorian populations. Our speci-mens agree in most details with the revised diagnosisfrom Simmons and Voss (1998), and they are clearlydistinct from E. andinus and E. brasiliensis. Pachecoet al. (2007) reported this species for the first time inwestern Peru based on a specimen collected in Que-brada Los Naranjos (03º5015" S, 80º1144.99" W; 550m) in Tumbes Department.
24 Special Publications, Museum of Texas Tech UniversityEptesicus innoxius (Gervais 1841)(Plate 2F)Specimens collected (17♀, 14♂).—Bosque Pro-tector Cerro Blanco (6), Reserva Ecológica ManglaresChurute (8), Isla Puná (13), Fuerte Militar Arenillas(2), Bosque Petrificado de Puyango (1), and CerroChiche (1).Remarks.—Eptesicus innoxius was reportedfor Ecuador based on 18 specimens from El Oro andGuayas Provinces (Davis 1966). Additionally, Albuja(1999) reported 25 specimens collected from Bar-raganete, Bolívar Province (01º27 S, 79º18 W; 430m). We noticed a large degree of variation in fur color,ranging from light-tobacco to brown, with differentlengths of dorsal hair. Other names used for thesepopulations include E. i. espadae (Cabrera 1901), fromLos Rios Province, and E. i. punicus (Thomas 1909b),from Guayas Province.Tribe LasiuriniLasiurus blossevillii (Lesson and Garnot 1826)(Plate 2G)Specimens collected (4♀, 3♂).—Bosque Protec-tor Cerro Blanco (4), Reserva Ecológica ManglaresChurute (1), Fuerte Militar Arenillas (1), and BosquePetrificado de Puyango (1).Remarks.—This species previously has beenrecorded in primary forest at Bosque Protector CerroBlanco (Salas-Z 2008). This is a remarkably beauti-ful bat with long hair and bright reddish tones/shades,unlike specimens from northern localities in Mexicoand the United States. Previously listed as L. borealis(e.g., Koopman 1993), it is now recognized as a distinctspecies (Baker et al. 1988). Based on distribution, ourrecords from western Ecuador would correspond to L.b. frantzii (see Baker et al. 1988).Lasiurus ega (P. Gervais 1856)Specimens collected (1♀, 2♂).—San Franciscode Bogota (1) and Reserva Ecológica Manglares Chu-rute (2).Remarks.—Within the large distribution of thisspecies (see Kurta and Lehr 1995), these populationswould be referable to L. e. punensis J. A. Allen 1914,with the type locality in Guayas Province. Some varia-tion in dorsal coloration occurs among our specimens,which resembles the pelage of subspecies within therange of L. ega. This species is sometimes listed as adifferent genus, Dasypterus (e.g., Barquez et al. 1999).Gardner and Handley (2008 ) recognize L. egafuscatus as the subspecies in western Colombia andwestern Ecuador.Tribe AntrozoiniRhogeessa velilla Thomas 1903Specimens collected (4♀, 5♂).—Bosque Protec-tor Cerro Blanco (6), Reserva Ecológica ManglaresChurute (2), and Fuerte Militar Arenillas (1).Remarks.—Although this species has been re-corded from western Ecuador as Rhogeessa io (Albujaand Mena-Valenzuela 2004), there are no previousrecords from Guayas and El Oro Provinces. Rhogeessaio has a diploid number of 30 whereas R. velilla fromwestern Ecuador has a diploid number of 42. Ourspecimens considerably extend the geographic range tothe south, and Pacheco et al. (2007) verified the pres-ence of Rhogeessa io in northwestern Peru based onspecimens from Quebrada Los Naranjos (03º5015" S,80º1144.99" W; 550 m), Tumbes Department. Basedon our field observations, this species is an early flyerand was usually netted at dusk. The karyotype fromour specimens is indistinguishable from that describedfor R. genowaysi from Chiapas, Mexico (Baker 1984).Use of the name velilla (Baird et al. 2008) is based onits potential distribution and type locality in Isla Puná,Guayas Province (Goodwin 1958), but a morphologicalcomparison remains to be drawn. Rhogeessa velillais restricted to the western Andean versant of the dryforest of southern Ecuador. This species is part of theR. tumida complex and is sister to other taxa currentlydistributed on the Atlantic coasts of Central Americaand Mexico (Baird et al. 2008). Such data supportsspecific status for R. velilla.
Carrera et al.—Bats of Western Ecuador 25Family MolossidaeEumops wilsoni Baker, McDonough, Swier, Larsen,Carrera, and Ammerman 2009(Plate 2H)Specimens collected (3♀, 5♂).— Bosque Protec-tor Cerro Blanco (7) and Isla Puná (1).Remarks.—These specimens initially were iden-tified as Eumops glaucinus, as reported from westernEcuador (Guayaquil) by Sanborn (1932), Eger (1977),andAlbuja (1999). However, McDonough et al. (2008)revised the E. glaucinus complex and divided it intofour species: E. ferox, distributed in the Caribbean,Mexico, and CentralAmerica; E. floridanus, restrictedto Florida, USA; E. glaucinus, restricted to SouthAmerica east of the Andes; and an undescribed taxonrestricted to the western versant of theAndes in Ecuadorand Peru. Subsequently, Baker et al. (2009) describedthis taxon as E. wilsoni based on genetic uniqueness(karyotypes, cytochrome-b sequence, andAFLPanaly-sis). All specimens collected during our survey werenetted in Guayas Province. The specimen collectionfrom Isla Puná represents a new record for this locality.Morphologically similar specimens have been reportedfrom Guayaquil and genetically similar specimens havebeen reported from northwestern Peru (Sudman et al.1994). The habitat for this species appears to be dryforest with adequate open understory within which thislarge species can forage. It is probable that this speciesis restricted to the dry forests of northwestern Peru andsouthwestern Ecuador.Molossus bondae J. A. Allen 1904Specimens collected (7♀, 3♂).—Bosque Protec-tor Cerro Blanco (1) and Reserva Ecológica ManglaresChurute (9).Remarks.—Eger (2008 ) considered Ec-uadorian populations as M. bondae bondae, althoughLópez-González and Presley (2001) applied the nameM. currentium bondae to populations from westernEcuador. Our specimens were collected inside primaryand secondary forest.Molossus molossus (Pallas 1766)Specimens collected (26♀, 15♂).—BosqueProtector Cerro Blanco (4), Reserva Ecológica Man-glares Churute (30), Isla Puná (4), and Fuerte MilitarArenillas (3).Remarks.—Although this species is common andwidespread across the Neotropics and is associatedwith human dwellings, these specimens constitute thefirst documented records for El Oro Province (Albuja1999). Interesting pelage color variations were foundacross our sample, including strikingly different redand black color phases. This species was found roost-ing in large trees at the Reserva Ecológica ManglaresChurute. Specimens from Isla Puná and Daule weredescribed as daulensis by J.A.Allen (1916); therefore,if applicable, it would be an appropriate name for thesepopulations (Eger 2008 ).DiscussionSome species of bats known from western Ecua-dor were not collected during the Sowell Expeditions.Table 2 summarizes the complete list of bats knownfrom western Ecuador, including both our collectionsand noteworthy records gathered from the literature,including: Balantiopteryx infusca (McCarthy et al.2000), Centronycteris centralis (Simmons and Handley1998), Anoura fistulata (Muchhala et al. 2005), Lon-chophylla chocoana (Dávalos 2004), Lonchophyllaorcesi (Albuja and Gardner 2005), Lonchophyllacadenai (Woodman and Timm 2006), Lonchophyllafornicata (Woodman 2007), Glyphonycteris daviesi(Pine et al. 1996), Platyrrhinus albericoi (Velazco2005), Sturnira koopmanhilli (McCarthy et al. 2006),and Platyrrhinus nitelinea (Velazco and Gardner 2009).Other species on the western versant of Ecuador werecompiled from Brosset (1965), Baker (1974), Albuja(1999), Angel and Salas (2003), Albuja and Mena-Valenzuela (2004), Albuja and Arcos (2007), Tirira(2007), Arroyo-Cabrales (2008a, b ), Davis andGardner (2008 ), Eger (2008 ), Marques-Aguiar (2008a, b ), Gardner (2008c, d, e ),Salas-Z. (2008), and Wilson (2008a, b ). Theserecords indicate that at least 40 bat species are known
26 Special Publications, Museum of Texas Tech UniversityTAXONCollected during SowellExpeditionsData on species recently revised,updated or describedEMBALLONURIDAE Balantiopteryx infusca McCarthy et al. 2000Centronycteris centralis Hood and Gardner 2008 Cormura brevirostris Albuja and Mena-Valenzuela 2004Diclidurus albus Hood and Gardner 2008 Peropteryx kappleri X Hood and Gardner 2008 Rhynchonycteris naso Albuja and Mena-Valenzuela 2004Saccopteryx bilineata X Albuja and Mena-Valenzuela 2004Saccopteryx leptura Hood and Gardner 2008 PHYLLOSTOMIDAE Micronycterinae Micronycteris giovanniae* X Fonseca et al. 2007Micronycteris hirsuta X Porter et al. 2007Micronycteris megalotis X Porter et al. 2007Micronycteris minuta X Porter et al. 2007Desmodontinae Desmodus rotundus X Kwon and Gardner 2008 Diaemus youngi† X Pinto et al. 2007Lonchorhininae Lonchorhina aurita X Williams and Genoways 2008 Phyllostominae Macrophyllum macrophyllum† X Williams and Genoways 2008 Trachops cirrhossus X Williams and Genoways 2008 Lophostoma aequatorialis* X Baker et al. 2004Lophostoma brasiliense† X Williams and Genoways 2008 Lophostoma silvicolum Williams and Genoways 2008 Mimon crenulatum X Williams and Genoways 2008 Phylloderma stenops X Williams and Genoways 2008 Phyllostomus discolor X Williams and Genoways 2008 Phyllostomus elongatus X Williams and Genoways 2008 Phyllostomus hastatus X Santos et al. 2003Tonatia saurophila X Williams and Genoways 2008 Chrotopterus auritus X Williams and Genoways 2008 Vampyrum spectrum X Williams and Genoways 2008 Glossophaginae Glossophaga soricina X Griffiths and Gardner 2008 Table 2. Complete list of species of bats from western Ecuador. An asterisk (*) indicates new spe-cies, and a dagger (†) shows new geographic records collected during the Sowell Expeditions toEcuador.
Carrera et al.—Bats of Western Ecuador 27TAXONCollected during SowellExpeditionsData on species recently revised,updated or describedAnoura aequatoris† X Mantilla-Meluk and Baker 2006Anoura cultrata† X Griffiths and Gardner 2008 Anoura fistulata Muchhala et al. 2005Anoura geoffroyi Griffiths and Gardner 2008 Choeroniscus minor Griffiths and Gardner 2008 Choeroniscus periosus Griffiths and Gardner 2008 Lichonycteris obscura X Griffiths and Gardner 2008 Lonchophillinae Lonchophylla cadenai Woodman and Timm 2006Lonchophylla chocoana Dávalos 2004Lonchophylla concava X Griffiths and Gardner 2008 Lonchophylla hesperia Griffiths and Gardner 2008 Lonchophylla fornicata Woodman 2007Lonchophylla orcesi Albuja and Gardner 2005Lonchophylla robusta X Griffiths and Gardner 2008 Lonchophylla thomasi X Griffiths and Gardner 2008 Carolliinae Carollia brevicauda X Hoffman and Baker 2003Carollia castanea X Solari and Baker 2006Carollia perspicillata X Hoffman and Baker 2003Glyphonycterinae Glyphonycteris daviesi Williams and Genoways 2008 Trinycteris nicefori† X Williams and Genoways 2008 Rhinophyllinae Rhinophylla alethina X McLellan and Koopman 2008 Sternodermatinae Sturnira bidens Gardner 2008a Sturnira bogotensis Gardner 2008a Sturnira erythromos Gardner 2008a Sturnira koopmanhilli McCarthy et al. 2006Sturnira lilium X Gardner 2008a Sturnira luisi X Gardner 2008a Sturnira oporaphilum X Gardner 2008a Sturnira sp. A Gardner 2008a Artibeus fraterculus X Marques-Aguiar 2008a Artibeus jamaicensis X Larsen et al. 2007Artibeus lituratus X Marques-Aguiar 2008a Chiroderma salvini Gardner 2008b Table 2. (cont.)
28 Special Publications, Museum of Texas Tech UniversityTable 2. (cont.)TAXONCollected during SowellExpeditionsData on species recently revised,updated or describedChiroderma trinitatum X Gardner 2008b Chiroderma villosum X Gardner 2008b Dermanura rava X Solari et al. 2009Dermanura rosenbergi X Solari et al. 2009Enchistenes hartii Marques-Aguiar 2008b Messophylla macconnelli X Arroyo-Cabrales 2008a Platyrrhinus albericoi Velazco and Patterson 2008Platyrrhinus chocoensis X Velazco and Patterson 2008Platyrrhinus dorsalis X Velazco and Patterson 2008Platyrrhinus helleri X Velazco and Patterson 2008Platyrrhinus ismaeli Velazco and Patterson 2008Platyrrhinus matapalensis X Velazco and Patterson 2008Platyrrhinus nigellus X Velazco and Patterson 2008Platyrrhinus nitelinea Velazco and Gardner 2009Uroderma bilobatum X Hoofer and Baker 2006Vampyressa thyone X Arroyo-Cabrales 2008b Vampyrodes caraccioli Gardner 2008c Vampyryscus nymphaea X Hoofer and Baker 2006NOCTILIONIDAE Noctilio leporinus X Gardner 2008d FURIPTERIDAE Amorphochilus schnablii Gardner 2008e THYROPTERIDAE Thyroptera discifera Wilson 2008a Thyroptera tricolor Wilson 2008a Thyroptera sp.* X VESPERTILIONIDAE Myotinae Myotis albescens X Wilson 2008b Myotis keaysi Wilson 2008b Myotis nigricans X Wilson 2008b Myotis oxyotus Wilson 2008b Myotis riparius X Wilson 2008b Myotis simus† X Wilson 2008b Vespertilioninae Eptesicus andinus Davis and Gardner 2008 Eptesicus chiriquinus X Davis and Gardner 2008 
Carrera et al.—Bats of Western Ecuador 29Table 2. (cont.)TAXONCollected during SowellExpeditionsData on species recently revised,updated or describedEptesicus innoxius X Davis and Gardner 2008 Lasiurus blossevillii X Gardner and Handley 2008 Lasiurus ega X Gardner and Handley 2008 Rhogeessa velilla X Baird et al. 2008MOLOSSIDAE Eumops auripendulus Eger 2008 Eumops perotis Eger 2008 Eumops wilsoni* X Baker et al. 2009Molossops aequatorianus Eger 2008 Molossus bondae X Eger 2008 Molossus molossus X Eger 2008 Nyctinomops macrotis Eger 2008 Promops centralis Eger 2008 Tadarida brasiliensis Eger 2008 to occur in western Ecuador that were not recordedduring this study, bringing the chiropteran diversity ofwestern Ecuador to 106 species.The Sowell Expeditions to western Ecuadorresulted in the addition of 11 species (7 geographicrecords, 3 new species, and 1 unrecognized species) tothe known bat diversity for the area, when combinedwith previous reports published by Albuja (1999),Tirira (1999), Albuja and Mena-Valenzuela (2004),Albuja and Arcos (2007), and Tirira (2007). Althoughour sampling was not repetitious for localities and wasnot geographically even (sampling was concentrated inthe northern and southern end of western Ecuador) ortemporally spread (all sampling was conducted Juneto August), the increase in species diversity and therange extensions reported are noteworthy (Table 2).These results, we believe, come from a comprehensiveapproach that includes the use of molecular systematictools (Baker et al. 2003b), increased taxon sampling(Larsen et al. 2007), and detailed studies of completemuseum voucher specimens (Solari et al. 2009). Weexpect that future field surveys, emphasizing Manabíand Los Ríos Provinces and the mid-elevation slopes ofthe western Ecuadorian Andes, will increase not onlythe known bat diversity but also our understanding ofspecies origins and potential conservation strategies.The Sowell Expeditions resulted in four unrecognizedspecies, three of which have since been described:Lophostoma aequatorialis (Baker et al. 2004), Micro-nycteris giovanniae (Fonseca et al. 2007), and Eumopswilsoni (Baker et al. 2009). The fourth species was aThyroptera that may be a form that has an availablename or may be an undescribed species. Furthermore,the voucher specimens for Macrophyllum macrophyl-lum, Trinycteris nicefori, and Diaemus youngi representnew distributional records for western Ecuador, thusdemonstrating the potential of this region for newscientific discoveries and future work.Western Ecuador constitutes a mosaic of differentenvironments (Hershkovitz 1958). The southwesternportion of the country encloses dry ecosystems that aresimilar to those found along the Pacific Coast of MiddleAmerica, but they are ecologically isolated by thepresence of the humid environments of BiogeographicChocó (Hershkovitz 1958; Rangel 2004). The forma-tion of the Chocoan region is a relatively recent eventassociated with the final period of theAndean uplifting(5 to 3 Mya) (Rangel 2004). Prior to the final upliftingof theAndes, xerophytic environments associated withthe effect of the Alisios winds extended their distribu-tion along the northern coast of South America andproto Central America (Graham 2006). The comple-
30 Special Publications, Museum of Texas Tech Universitytion of the Isthmus of Panama and the deviation of theHumboldt marine current resulted in the predominanthyper-humid environments of Central Chocó. Thus,with the formation of the Chocan region, an ecologicalhiatus exists in western Ecuador (Rangel 2004).Our records indicate differences in bat speciescomposition between northwestern and southwesternEcuador (Table 1) and similarities between northwest-ern Ecuador and western Colombia (see Mantilla-Me-luk et al. 2009). This result is expected considering thedirect influence of the Chocó – Darién humid region inthe Esmeraldas Province in the north and the Peruvian–AcknowledgmentsThe authors sincerely thank the many individualsand institutions that contributed to these studies. Mostsignificantly, James Sowell and Alan Brown providedthe funds that made Sowell Expeditions 2001 and 2004possible. Patricia Galeano, Tatiana Egüez Larrea, andGabriela Montoya at the Ministerio del Ambiente delEcuador authorized the scientific permit (No. 023-IC-FAU-DNBAP/MA) to collect small mammals at eachstudy site. Laura Arcos-Terán, Luis A. Coloma, andSantiago Burneo helped with logistical arrangementsthrough the Department of Biological Sciences at Pon-tificia Universidad Católica del Ecuador. Eric Horst-man, Nelson Zambrano, and Jaime Salas (FundaciónProbosque) provided several facilities at Bosque Pro-tector Cerro Blanco. Special thanks go to Carleton J.Phillips, Federico G. Hoffmann, Jana L. Higginbotham,Deidre A. Parish, Carl W. Dick, Cristina Acuña, Ga-briela Jiménez, Horacio Ponce and Family, and localpeople from: Reserva Ecológica Manglares Churute,Fuerte MilitarArenillas, Isla Puná, Bosque PetrificadoPuyango, San Francisco de Bogotá, San Lorenzo, Mata-je, Portovelo, Jardín Botánico Moromoro (FundaciónJocotoco), and Zaruma, who provided valuable helpand assistance during fieldtrips. Heath Garner, KathyMacDonald, and Annie Spikes at the NSRL helped toprocess and curate the specimens. Faisal Anwarali,Michelle Knapp, Heather Meeks, Steven R. Hoofer, andCalvin Porter assisted in generating DNA sequences.Paul Velazco (FMNH) provided valuable informationand comments about the genus Platyrrhinus. HugoMantilla-Meluk assisted by creating Figure 1. Aspecialrecognition to Luis Albuja, Don E. Wilson, Thomas E.Lee, Miguel Pinto, Jackie B. Chavez, Raquel Marchán,Nicté Ordóñez-Garza, Courtney Thomason, Noe de laSancha, Jaime Salas, and Lisa Bradley who providededitorial assistance and suggestions that improved thispaper. Alava J. J., and R. Carvajal. 2004. Ocurrencia de Noctilio lep-orinus (Chiroptera: Noctilionidae) en la zona urbana yalrededores de Guayaquil, Ecuador. Chiroptera Neotropi-cal 10(1-2):183-187.Alberico, M. S., and E. Velasco. 1991a. Description of a newbroad-nosed bat from Colombia. Bonner ZoologischeBeitrage 42:237-239.Alberico, M. S., and E. Velasco. 1991b. Extended descriptionof Platyrrhinus chocoensis from the Pacific lowlands ofColombia. Trianea 5:343-351.Chilean desert formation in the southern Guayas andEl Oro Provinces. The diversity of vegetation typesand ecosystems in northwestern Ecuador has alloweda large number of species of Emballonuridae, Noctil-ionidae, Phyllostomidae, and Thyropteridae to coex-ist. In the dry environments of southwestern Ecuadorspecies of Molossidae and Vespertilionidae are morehighly represented. The noteworthy results achievedduring the Sowell Expeditions to Ecuador suggest theimportance of continued intensive field surveys of batfaunas along the country, especially in Manabí and LosRíos Provinces as well as middle and higher elevationsof the western versant of the Ecuadorian Andes.Literature CitedAlbuja V., L. 1989. Adiciones a la fauna de quirópteros delnoroccidente del Ecuador. Escuela Politécnica 14:105-111.Albuja V., L. 1999. Murciélagos del Ecuador. 2da edición.Departamento de Ciencias Biológicas. Quito, Ecuador,CICETRONIC. 288 pp.Albuja V., L., and R. Muñoz. 2000. Fauna del Parque NacionalMachalilla. Pp. 32-41 in Compendio de Investigacionesen el Parque Nacional Machalilla. (M. Iturralde and C.Josse, eds.). Centro de Datos para la Conservación yFundación Natura. Quito, Ecuador.