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Molinari, Jesús, Gutiérrez, Eliécer E., Lim, Burton K. (2023): Systematics and biogeography of Anoura cultrata (Mammalia, Chiroptera, Phyllostomidae): a morphometric, niche modeling, and genetic perspective, with a taxonomic reappraisal of the genus. Zootaxa 5297 (2): 151-188, DOI: 10.11646/zootaxa.5297.2.1, URL: http://dx.doi.org/10.11646/zootaxa.5297.2.1

The study of human relationships with other components of the landscape allows us to understand the dynamics of ecological communities and biodiversity. Although there has been an increase in ethnobiological studies, little is known about the interaction between humans and bats. The objective of this study was to evaluate the perceptions and attitudes about bats by people in the fragile life zone of the tropical dry forest of Colombia. We selected four representative areas and evaluated three different aspects: (1) biological and ecological knowledge, (2) uses and cultural beliefs, and (3) interactions between humans and bats. The data were collected through surveys of 18 questions to 197 people with a gender ratio close to parity. The data obtained are presented as descriptive statistics by using a generalized linear model (GLM). The results showed that people's knowledge and attitude towards bats depend on the covariates of education and gender of the respondents. Women (t= 3.76; P=0.004) and people with a higher education level (t=3.65; P

C. v. villosum Peters, 1860 Synonyms: Chiroderma villosum Peters, 1860:748; type locality “Brasilia.” Chiroderma villosa Jones, 1951: 224; incorrect gender concordance. Chiroderma villosum villosum: Handley, 1960:466; first use of current name combination. Chiroderma trinitatum: Pirlot, 1972: 76, not Chiroderma trinitatum Goodwin, 1958. Chiroderma sp. Nowak, 1994: 160. Chiroderma salvini Nowak, 1994: 160, not Chiroderma salvini Dobson, 1878. Chiroderma [sp.] Czaplewski & Cartelle, 1998: 794. see Garbino & Tavares (2018a) Chiroderma salvini: Medina et al., 2015: 204, not Chiroderma salvini Dobson, 1878. Chiroderma salvini: Rocha et al., 2016: 573; not Chiroderma salvini Dobson, 1878. Chiroderma salvini: Maas et al., 2018: 672; not Chiroderma salvini Dobson, 1878. Distribution and Habitat. The nominal subspecies, C. v. villosum, is known from the tropical region of cis-Andean South America. The taxon has been recorded in Colombia, Venezuela, Trinidad and Tobago, Guyana, Suriname, French Guiana, Brazil, Ecuador, Perú, and Bolivia (Fig. 28). The southern limit of the subspecies is in the Brazilian state of Paraná (not mapped; Reis et al. 1998). The subspecies occurs in a wide variety of environments, including humid and seasonal forests (e.g. Amazonia, Atlantic rainforest), flooded plains (e.g. Pantanal, Llanos), and dry formations (e.g. Caatinga, Cerrado) (Handley 1976; Gregorin et al. 2008; Luz et al. 2011; Fischer et al. 2015). C. v. villosum apparently tolerates some habitat disturbance, as it occurs in small forest fragments and urban areas (Nogueira & Peracchi 2003; Ferreira et al. 2010; Nunes et al. 2017). Chiroderma villosum villosum occurs from sea level to approximately 1,000 m. In Venezuela, 99% of the specimens of C. v. villosum were captured below 500 m (Handley 1976). In the Peruvian Amazonia (Madre de Dios), the species was recorded between 340 and 950 m (Solari et al. 2006)., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on page 54, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Peters, W. (1860) Eine neute Gattung von Flederthieren, Chiroderma villosum, aus Brasilien vor. Monatsberichte der Koniglichen Preussische Akademie des Wissenschaften zu Berlin, 1861,747 - 755.","Jones, T. (1951) Bat records from the islands of Grenada and Tobago, British West Indies. Journal of Mammalogy, 32 (2), 223 - 224. https: // doi. org / 10.1093 / jmammal / 32.2.223 - b","Handley Jr., C. O. (1960) Descriptions of new bats from Panama. Proceedings of the United States National Museum, 112, 459 - 479. https: // doi. org / 10.5479 / si. 00963801.112 - 3442.459","Pirlot, P. (1972) Chiropteres de moyenne Amazonie. Mammalia, 36 (1), 71 - 85. https: // doi. org / 10.1515 / mamm. 1972.36.1.71","Goodwin, G. G. (1958) Three new bats from Trinidad. American Museum Novitates, 1877, 1 - 6.","Nowak, R. M. (1994) Walker's bats of the world. Johns Hopkins University Press, Baltimore, Maryland, 863 pp.","Dobson, G. E. (1878) Catalogue of the Chiroptera in the collection of the British Museum. British Museum of Natural History, London, xxxvi + 567 pp. https: // doi. org / 10.5962 / bhl. title. 55341","Czaplewski, N. J. & Cartelle, C. (1998) Pleistocene bats from cave deposits in Bahia, Brazil. J ournal of Mammalogy, 79 (3), 784 - 803. https: // doi. org / 10.2307 / 1383089","Garbino, G. S. T. & Tavares, V. da C. (2018 a) A Quaternary record of the big-eyed bat Chiroderma villosum (Chiroptera: Phyllostomidae) with a revised lower molar terminology. Mammalia, 82 (4), 393 - 399. https: // doi. org / 10.1515 / mammalia- 2017 - 0037","Medina, C. E., Lopez, E., Pino, K., Pari, A. & Zeballos, H. (2015) Biodiversidad de la zona reservada Sierra del Divisor (Peru): una vision desde los mamiferos pequenos Revista Peruana de Biologia, 22 (2), 199 - 212. https: // doi. org / 10.15381 / rpb. v 22 i 2.11354","Rocha, P. A., Brandao, M. V., Garbino, G. S. T., Cunha, I. N. & Aires, C. C. (2016) First record of Salvin's big-eyed bat Chiroderma salvini Dobson, 1878 for Brazil. Mammalia, 80 (5), 573 - 578. https: // doi. org / 10.1515 / mammalia- 2015 - 0077","Maas, A. C. S., Gomes, L. A. C., Martins, M. A., Dias, D., Pol, A., Chaves, F. G., Schute, M., Araujo, R. M. & Peracchi, A. L. (2018) Bats in a Cerrado landscape of Northern Brazil: Species occurrence, influence of environmental heterogeneity and seasonality, and eight new records for the State of Tocantins. Mammalia, 82 (5), 469 - 480. https: // doi. org / 10.1515 / mammalia- 2017 - 0023","Reis, N. R., Peracchi, A. L., Lima, I. P., Sekiama M. L. & Rocha, V. J. (1998) Updated list of the chiropterians [sic] of the city of Londrina, Parana, Brazil. Chiroptera Neotropical, 4 (2), 96 - 98.","Handley Jr., C. O. (1976) Mammals of the Smithsonian Venezuelan Project. Brigham Young University Science Bulletin, Biological Series, 20 (5), 1 - 89. https: // doi. org / 10.5962 / bhl. part. 5667","Gregorin, R., Carmignotto, A. P. & Percequillo, A. R. (2008) Quiropteros do Parque Nacional da Serra das Confusoes, Piaui, nordeste do Brasil. Chiroptera Neotropical, 14 (1), 366 - 383.","Luz, J. L., Mangolin, R., Esberard, C. E. L. & Bergallo, H. G. (2011) Morcegos (Chiroptera) capturados em lagoas do Parque Nacional da Restinga de Jurubatiba, Rio de Janeiro, Brasil. Biota Neotropica, 11 (4), 161 - 168. https: // doi. org / 10.1590 / S 1676 - 06032011000400016","Fischer, E., Santos, C. F., Carvalho, L. F. A. C., Camargo, G., Cunha, N. L., Silveira, M., Bordignon, M. O. & Silva, M. O. (2015) Bat fauna of Mato Grosso do Sul, southwestern Brazil. Biota Neotropica, 15 (2), 1 - 17. https: // doi. org / 10.1590 / 1676 - 06032015006614","Nogueira, M. R. & Peracchi, A. L. (2003) Fig-seed predation by 2 species of Chiroderma: discovery of a new feeding strategy in bats. Journal of Mammalogy, 84 (1), 225 - 233. https: // doi. org / 10.1644 / 1545 - 1542 (2003) 084 2.0. CO; 2","Ferreira, C. M. M., Fischer, E. & Pulcherio-Leite, A. (2010) Fauna de morcegos em remanescentes urbanos de Cerrado em Campo Grande, Mato Grosso do Sul. Biota Neotropica, 10 (3), 155 - 160. https: // doi. org / 10.1590 / S 1676 - 06032010000300017","Nunes, H., Rocha F. L. & Cordeiro-Estrela, P. (2017) Bats in urban areas of Brazil: roosts, food resources and parasites in disturbed environments. Urban Ecosystems, 20 (4), 953 - 969. https: // doi. org / 10.1007 / s 11252 - 016 - 0632 - 3","Solari, S., Pacheco, V., Luna, L., Velazco, P. M. & Patterson, B. D. (2006) Mammals of the Manu Biosphere Reserve. Fieldiana Zoology, New Series, 110, 13 - 22. https: // doi. org / 10.3158 / 0015 - 0754 (2006) 110 [13: MOTMBR] 2.0. CO; 2"]}

We present a revision of the Neotropical bat genus Chiroderma, commonly known as big-eyed bats. Although species of Chiroderma have a wide distribution from western México to southern Brazil, species limits within Chiroderma are not clearly defined, as attested by identification errors in the literature, and there is no comprehensive revision of the genus that includes morphological and molecular data. Our review is based on phylogenetic analyses of two mitochondrial (COI and CYTB) and two nuclear (RAG2 and DBY) genes, coalescence analyses of mitochondrial genes, and morphological analyses including type specimens of all named taxa. We recognize seven species in three clades: the first clade includes (1) C. scopaeum Handley, 1966, endemic to western México and previously considered a subspecies of C. salvini; and (2) C. salvini Dobson, 1878, a taxon associated with montane forests, distributed from México to Bolivia; the second clade includes (3) C. improvisum Baker and Genoways, 1976, endemic to the Lesser Antilles, and (4) C. villosum Peters, 1860, widely distributed on the continental mainland and polytypic, with subspecies C. v. villosum and C. v. jesupi; and the third clade includes (5) the polytypic C. doriae Thomas, 1891, with C. d. doriae distributed in eastern Brazil and Paraguay, and C. d. vizottoi, occurring in northeastern Brazil; (6) C. trinitatum Goodwin, 1958, distributed from Trinidad to Amazonia; and (7) C. gorgasi Handley, 1960, distributed from Honduras to trans-Andean South America, previously considered a subspecies of C. trinitatum.

Chiroderma improvisum Baker and Genoways, 1976 Synonyms: Chiroderma improvisum Baker and Genoways, 1976: 1; type locality “ Guadeloupe: Basse-Terre; 2 km. S, 2 km. E Baie-Mahault.” Type Material. The type, TTU 19900 (not seen), is a skin, skull and mandible, collected by R. J Baker and H. H. Genoways (field number J. C. Patton 552) in July 29, 1974 on the Basse-Terre island, Guadalupe (Baker & Genoways 1976). It is an adult male captured in a mist net set in a pasture adjacent to gallery forest. The karyotype of the specimen is deposited with the tissue collection of the Texas Tech University under the number TK 8285. The skin, skull, and mandible were examined by means of photographs and are in good condition. Distribution and Habitat. The species is known from four islands in the Lesser Antilles: Guadeloupe, Montserrat, Saint Kitts, and Nevis (Fig. 28). There is also a subfossil specimen collected on the island of Marie-Galante, south of Guadeloupe (Lenoble 2019). Records of C. improvisum are from areas of dry forests and humid forests, from sea level up to approximately 350 m. The few known specimens were captured in mist nets set over streams, in gallery forests, in secondary forests surrounded by pasture and plantations, and in urban and peri-urban areas (Baker et al. 1978; Jones Jr. & Baker 1979; Pierson et al. 1986; Pedersen et al. 2010; Beck et al. 2016). However, a harp trap set across a dry ravine caught the first specimen of C. improvisum from Nevis (Lim et al., 2020). Description and Comparisons. Dorsal pelage varies from grayish to dark brown. Dorsal pelage is long (approximately 13 mm). Individual hairs of the dorsum are tricolored with a grayish base and buff middle band. The interocular stripes are weakly developed and the genal pair was not visible in the two skins we examined, but its presence can be verified in photographs of live animals (Baker & Genoways 1976; Jones Jr. & Baker 1980; Lim et al. 2020). The dorsal stripe is inconspicuous, visible from the mid dorsum to near base of the uropatagium. The ear is moderate to dark brown along most of its length; the base is yellowish. The noseleaf is uniformly dark to medium brown, and the tip may be simple or notched, as evident in the figure in Jones & Baker (1980). The posterior border of the uropatagium has a V-shaped notch. The skull of C. improvisum is the largest among Chiroderma (Tables 7 and 8). The braincase is low and slopes evenly to the frontonasal region in profile. The sagittal and lambdoid crests are conspicuous in both specimens examined and in the type. The nasal notch extends behind the anterior margin of the orbits (Fig. 29). Post-orbital processes are distinct, but not pointed. The orbits are relatively small and the posterior margin approximates the level of the mesial margin of M1 (Fig. 30). A medial accessory foramen between incisive foramina is lacking. There is no posterior palatine process. When skull and mandible are in occlusion, there is a small lateral gap bordered by C, P3, p2, and p4 (Fig. 9). In addition, there is a frontal gap surrounded by I1, I2, c, i1, and i2 (Fig. 14). The I1s are convergent and their tips may or may not be in contact. P3 is separated from P4 by a small gap, or the two teeth contact each other (Fig. 29). The mandibular condyle is above or level with the tooth row. Lower canines are relatively short, their tips below the level of the top of the coronoid process. The basal cingulum of the lower canines is well developed and crowding the lower incisors. The p2 is mesiodistally longer than tall, and is approximately ⅓ of the height of p4 (Fig. 30). The p2 is in contact with lower canines, either touches p4 or the two lower premolars are separated by a small gap. The p4 protoconid is robust, with its distal margin sloping evenly to the tooth’s, not abruptly as in the other species. Compared with C. doriae, which has a similar size, C. improvisum is larger, has grayish or dark brown dorsal pelage (medium brown in doriae), faint facial stripes (bright in doriae), and unicolored noseleaf and ears (in doriae the horseshoe of the noseleaf has pale borders and the ears also have whitish margins). Cranially, C. improvisum can be diagnosed by its wider nasal notch and by the broader posterior border of the palate, not narrow inverted-U shaped as in C. doriae. The P3 of C. improvisum is not expanded buccolingually and is in contact with P4, differing from C. doriae in both aspects. The lower canines of C. improvisum are relatively larger than in C. doriae. The p2 of C. improvisum is mesiodistally longer than tall; whereas, in C. doriae, p2 is taller exceeding ⅔ the height of p4. The p4 of C. improvisum is more robust than in C. doriae, and the protoconid is long mesiodistally. A frontal gap is present in C. improvisum, but lacking in C. doriae. The species most similar to C. improvisum in qualitative characters is C. villosum, from which improvisum can be differentiated by the much larger size, darker pelage (pale brown in villosum), absence of a posterior palatine process, convergent I1s (parallel and separate in villosum), relatively smaller lower canines (taller and more pointed in villosum), large p 2 in contact with p4 (gap between p2 and p 4 in villosum). Geographic Variation and Phylogeography. The genetic distance between the two specimens of C. improvisum in our molecular analysis was 0.22% (Fig. 4). Due to the small sample, we cannot make inferences either on geographic structuring or morphological variation in the taxon. Subspecies. C. improvisum is monotypic. Natural History. There is no information on the diet of C. improvisum, but it probably feeds on fruits, infructescences and their seeds, as the other species of Chiroderma. The type was collected approximately 6 m above the ground and near a forest having a 15-meter-high canopy (Baker et al. 1978), suggesting that the species may be active in the canopy as is the closely-related C. villosum. A mite, Periglischrus iheringi (Spinturnicidae), was recorded on C. improvisum from Saint Kitts (Beck et al. 2016). We lack information on reproduction in C. improvisum. Specimens Examined (N = 2): Montserrat: Saint Anthony Parish, 0.8 km above mouth of Belham river (TTU 31403). Saint Kitts and Nevis: Saint Thomas Parish (Nevis) ¸ Barnes Ghaut (ROM 126002)., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 46-48, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Baker, R. J. & Genoways, H. H. (1976) A new species of Chiroderma from Guadeloupe, West Indies (Chiroptera: Phyllostomatidae). Occasional Papers of the Museum of Texas Tech University, 48, 1 - 9.","Lenoble, A. (2019) The past occurrence of the Guadeloupe big-eyed bat Chiroderma improvisum Baker and Genoways, 1976 on Marie-Galante (French West Indies) with comments on bat remains from pre-Columbian sites in Eastern Caribbean. Acta Chiropterologica, 21 (2), 299 - 308. https: // doi. org / 10.3161 / 15081109 ACC 2019.21.2.005","Baker, R. J., Genoways, H. H. & Patton, J. C. (1978) Bats of Guadeloupe. Occasional Papers of the Museum of Texas Tech University, 50, 1 - 16.","Pierson, E. D., Rainey, W. E., Warner, R. M. & White-Warner, C. C. (1986) First record of Monophyllus from Montserrat, West Indies. Mammalia, 50 (2), 269 - 271. https: // doi. org / 10.1515 / mamm. 1986.50.2.263","Pedersen, S. C., Kwiecinski, G. G., Larsen, P. A., Morton, M. N., Adams, R. A., Genoways, H. H. & Swier, V. J. (2010) Bats of Montserrat: population fluctuation and response to hurricanes and volcanoes, 1978 - 2005. In: Fleming, T. H. & Racey, P. A. (Eds.), Island bats: Evolution, ecology, and conservation. The Chicago University Press, Chicago, Illinois, pp. 302 - 340.","Beck, J. D., Loftis, A. D., Daly, J. L., Reeves, W. K. & Orlova, M. V. (2016) First record of Chiroderma improvisum Baker & Genoways, 1976 (Chiroptera: Phyllostomidae) from Saint Kitts, Lesser Antilles. Check List, 12 (2), 10 - 13. https: // doi. org / 10.15560 / 12.2.1854","Lim, B. K., Loureiro, L. O. & Garbino, G. S. T. (2020) Cryptic diversity and range extension in the big-eyed bat genus Chiroderma (Chiroptera: Phyllostomidae). ZooKeys, 918, 41 - 63. https: // doi. org / 10.3897 / zookeys. 918.48786","Jones Jr., J. K. & Baker, R. J. (1980) Chiroderma improvisum. Mammalian Species, 134, 1 - 2. https: // doi. org / 10.2307 / 3504068"]}

Chiroderma gorgasi Handley, 1960 Synonyms: Chiroderma gorgasi Handley, 1960: 464; type locality “ Tacarcuna Village, 3,200 ft., Río Pucro, Darién, Panama.” Chiroderma trinitatum gorgasi: Barriga-Bonilla, 1965: 246; name combination. Type Material. The type of C. gorgasi, USNM 309903, consists of a stuffed skin, skull and mandible, collected in Tacarcuna Village, Panamá, on March 6, 1959 by C. O. Handley and B. R. Feinstein (field number COHJR 5436). It is an adult male, captured in a mist net over water. The skin is in good condition with the facial and dorsal stripes observable in the specimen. The skull and mandible are in good condition and every tooth is present. The I1 have convergent tips. Distribution and Habitat. Specimens are known from Panamá, western Colombia, and northwestern Ecuador (Fig. 23), and there is a record from eastern Honduras (Turcios-Casco et al. 2020). The unconfirmed record for Costa Rica is based on a bat captured and released by R. LaVal in Tortuguero (Timm & LaVal 1998). Also in Costa Rica, Harvey & González-Villalobos (2007) reported the capture of 18 “ Chiroderma trinitatum ” in Talamanca, but we could not verify if there are voucher specimens to support this claim. The occurrence of the species in Costa Rica is expected, as C. gorgasi has been recorded in western Panamá and eastern Honduras (Handley 1966b; Turcios- Casco et al. 2020). The records of C. gorgasi are from the humid forests of the Chocó of Colombia, the Darién of Panamá, Caribbe- an lowland forests of Honduras, and montane forests of the Rio Magdalena valley in Colombia. The altitude where specimens have been obtained ranges from 30 m in Esmeraldas, Ecuador, to 975 m, in Tacarcuna, Panamá. There are Colombian records of the species occurring at 2,100 in Tolima and between 1,600 and 2,300 m in Risaralda (Galindo-Espinosa et al. 2010; Castaño et al. 2018). Description and Comparisons. Dorsal pelage may vary from pale to dark brown. Individual hairs of the dorsum are tricolored: the base is approximately ¼ of the hair length and dark brown, the middle band is approximately ½ of the length of the hair and varies from buff to pale gray, and the tip is about ¼ of the hair length and varies from pale to dark brown. Both pairs of facial stripes are conspicuous. The dorsal stripe is conspicuous in approximately half of the sample (47%, n=8); whereas, it is barely visible in nine specimens. The ear margins and base are paler than the remainder of the ear conch. The noseleaf has a simple tip, is brown in color, with pale margins on the horseshoe. Dimensions of the skull are similar to those of C. trinitatum, and the two species are the smallest Chiroderma (Tables 7 and 8). Braincase is globose, clearly distinguishable in profile from the frontonasal region. The sagittal crest is poorly developed and was not detected in 9 of the 17 specimens we scored for this character. The nasal notch is short and either does not reach the interorbital region, or extends only the level of the anterior border of the orbit. Similar to C. trinitatum, the post-orbital processes are rhomboid and not pointed as in the other Chiroderma. The posterior palatine process was absent in 13 of 15 specimens and in the other 2, the process was only a small bump. When cranium and mandible are in occlusion, a lateral gap is visible, bordered by C, P3, P4, p2 and p4. The I1s are convergent and their tips are usually in contact. The mandibular condyle is level with or slightly below the toothrow. The lower canines are relatively narrow and high-crowned, with the crown tip level with the top of the coronoid process, when viewed laterally. The p2 is in contact with c, but not with p4, or if not in contact with the lower canine, p2 may be either closer to the lower canine or equidistant from c and p4. The p2 usually is longer mesiodistally than high and the protoconid is shifted anteriorly, not aligned with the base of the tooth when viewed laterally (Fig. 27). Chiroderma gorgasi differs from every other Chiroderma, except C. trinitatum, by its smaller size and nasal notch usually not reaching the interorbital region. Comparisons with C. trinitatum were made in the previous section. Geographic Variation and Phylogeography. Sequences of three individuals of C. gorgasi were analyzed in the phylogeny, precluding making inferences on geographical structuring. Within-species variation was 1.04%, the second highest value in Chiroderma after C. villosum (1.17%). Subspecies. C. gorgasi is monotypic. Remarks. Handley (1960) described Chiroderma gorgasi based on five specimens from Panamá and one C. trinitatum from Trinidad, the type and only known specimen at the time. In the original description, Handley (1960: 465) suggested that, as the sample size increased, the two taxa could prove to be conspecific. Shortly after its description, C. gorgasi was treated as a subspecies of C. trinitatum, based on their morphological similarity (Barriga- Bonilla 1965; Jones & Carter 1976; Hall 1981). Simmons (2005) recognized a monotypic trinitatum with gorgasi as a junior synonym. Recently, Lim et al. (2020) recognized C. gorgasi as a distinct species, because it does not share a most recent common ancestor with C. trinitatum, and has distinguishing morphological characters. Natural History. C. gorgasi is a frugivore, specialized on fruits of Ficus (Bonaccorso 1979). Four species of fruits and infructescences have been recorded in the diet of C. gorgasi: Ficus popenoei, Piper aduncum, Solanum umbellatum, and Vismia sp. (Bonaccorso 1979; Castaño et al. 2018). The vertical distribution suggests that C. gorgasi is a canopy and sub-canopy frugivore, more frequently captured in nets between 3 and 12 m above ground (Bonaccorso 1979). The few reproductive data for the species suggest a pattern of seasonal polyestry. A pregnant female was captured in June in Colombia and lactating individuals were recorded in February and March in Panamá. Literature data for Panamá report pregnancies in February, May and between September and November; whereas lactating females are documented from May and September (Fleming 1973; Bonaccorso 1979). Births apparently occur toward the end of the dry season, between February and May, and in the middle of the rainy season, between July and September, when fruits are most abundant. Specimens Examined (N = 18): Colombia: Antioquia, La Tirana (IAvH-M 917, 934, 974, USNM 499475, 499477, 499479); Chocó, Corregimiento Gilgal (IAvH-M 4932), Finca El Recurso (IAvH-M 3260, 3299, 3323); Valle del Cauca, Río Zabaletas (USNM 483764). Panamá: Darién, Parque Nacional Darién (ROM 104342), Tacarcuna Village Camp (USNM 309902, 309903 [holotype of gorgasi], 309904); San Blas, Armila (USNM 335294, 335296, 335297)., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 44-45, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Handley Jr., C. O. (1960) Descriptions of new bats from Panama. Proceedings of the United States National Museum, 112, 459 - 479. https: // doi. org / 10.5479 / si. 00963801.112 - 3442.459","Barriga-Bonilla, E. (1965) Estudios mastozoologicos colombianos, I: Chiroptera. Caldasia, 9 (43), 241 - 268.","Turcios-Casco, M. A., Medina-Fitoria, A. & Estrada-Andino, N. (2020) Northernmost record of Chiroderma trinitatum (Chiroptera, Phyllostomidae) in Latin America, with distributional comments. Caribbean Journal of Science, 50 (1), 9 - 15. https: // doi. org / 10.18475 / cjos. v 50 i 1. a 2","Timm, R. M. & LaVal, R. K. (1998) A field key to the bats of Costa Rica. Occasional Publication Series, Center for Latin Ameri- can Studies, University of Kansas, 22, 1 - 30.","Harvey, C. A. & Gonzalez-Villalobos, J. A. (2007) Agroforestry systems conserve species-rich but modified assemblages of tropical birds and bats. Biodiversity and Conservation, 16, 2257 - 2292. https: // doi. org / 10.1007 / s 10531 - 007 - 9194 - 2","Handley Jr., C. O. (1966 b) Checklist of the mammals of Panama. In: Wenzel, R. L. & Tipton, V. J. (Eds.), Ectoparasites of Panama. Field Musem of Natural History, Chicago, Illinois, pp. 753 - 795.","Galindo-Espinosa, E. Y., Gutierrez-Diaz, K. A. & Reinoso-Florez, G. (2010) Lista de los quiropteros del departamento del Tolima, Colombia. Biota Colombiana, 11 (1 / 2), 107 - 116.","Castano, H. J., Carranza, A. J. & Perez-torres, J. (2018) Diet and trophic structure in assemblages of montane frugivorous phyllostomid bats. Acta Oecologica, 91, 81 - 90. https: // doi. org / 10.1016 / j. actao. 2018.06.005","Jones Jr., J. K. & Carter, D. C. (1976) Annotated checklist, with keys to subfamilies and genera. In: Baker, R. J. & Jones Jr., J. K. (Eds.), Biology of the bats of the New World family Phyllostomatidae. Part I. Museum of Texas Tech University Special Publications 10. Museum of Texas Tech University Press, Lubbock, Texas, pp. 7 - 38.","Hall, E. R. (1981) The mammals of North America. Vol. 1. John Wiley & Sons, New York, xxx + 546 + 79 pp.","Simmons, N. B. (2005) Order Chiroptera. In: Wilson, D. E. & Reeder, D. M. (Eds.), Mammal species of the world: a taxonomic and geographic reference. Johns Hopkins University Press, Baltimore, Maryland, pp. 312 - 529.","Lim, B. K., Loureiro, L. O. & Garbino, G. S. T. (2020) Cryptic diversity and range extension in the big-eyed bat genus Chiroderma (Chiroptera: Phyllostomidae). ZooKeys, 918, 41 - 63. https: // doi. org / 10.3897 / zookeys. 918.48786","Bonaccorso, F. J. (1979) Foraging and reproductive ecology in a Panamanian bat community. Bulletin of the Florida State Museum Biological Sciences, 24 (4), 359 - 408.","Fleming, T. H. (1973) The reproductive cycles of three species of opossums and other mammals in the Panama Canal Zone. Journal of Mammalogy, 54, (2) 439 - 455. https: // doi. org / 10.2307 / 1379129"]}

Chiroderma doriae Thomas, 1891 Synonyms: See under subspecies. Type Material. The type of Chiroderma doriae, by original designation, is specimen BMNH 44.9.2.6 (skin) and BMNH 49.8.16.29 (skull and mandible). The holotype is an adult of undetermined sex, but possibly female, as there is no vestige of scrotal sac on the skin. The skin was stitched longitudinally along the dorsum and probably for this reason the descriptions described the dorsal stripe as absent (Dobson 1878, misdentified as C. villosum; Vieira 1942). The left ear, as well as the right tarsus and metatarsus, have been lost, but the right calcar and the tibia are still attached to the skin. Interocular stripes are evident, but the color of the skin appears to be slightly faded. The genal pair of facial stripes is less conspicuous, and the stripe on the right side is more evident than on the left. Part of the medial uropatagium is torn. The skull is damaged, and the region posterior to the mesopterygoid fossa, i.e. the basioccipital, occipital, and the posterior part of the braincase are missing. The upper dental arcade is complete. The right zygomatic arch is broken, and the root of the right upper canine is exposed. Part of the palate is also broken. The inner upper incisors have convergent tips, almost touching each other. The mandible and lower dentition are in good condition, except for the left angular process, which is missing. Information on the label indicates “Minas Geraes” as the locality and that the type was obtained from Parzudaki, referring to Charles Parzudaki and his adopted son Emile, two active dealers in natural history specimens in Paris. As Charles and Emile probably never collected in South America (Gouraud et al. 2016), it is not possible to determine the collector or to estimate a more precise locality for the specimen. Distribution and Habitat. See under Subspecies. Description and Comparisons. Dorsal pelage is pale brown in most specimens of C. d. doriae, but some have dark brown pelage. In C. d. vizottoi the dorsal pelage is pale buff (Fig.18). Dorsal hairs are tricolored; the base is dark brown, the middle is buff, and the tip is pale brown (C. d. doriae) or pale buff (C. d. vizottoi). Two pairs of interocular and genal stripes are present, wide, and formed by entirely white hairs. A median dorsal stripe is present. The stripe is conspicuous in 25 of the 29 (86%) C. d. doriae examined, and usually extends from the interscapular region to the posterior extremity of the rump. One C. d. doriae (ZUFMS 395) has the dorsal stripe beginning on the nape. The ear and tragus are yellowish at the bases, as well as is the margin of the ear. The remainder of the ear is brownish. The spear of the noseleaf is simple-tipped and the lateral borders of the horseshoe are paler than the medial portion and spear. Chiroderma doriae has the second largest cranium among Chiroderma species, smaller only than C. improvisum (Tables 7 and 8). There is some overlap among the measurements of small C. d. doriae and large C. salvini. Measurements of C. d. vizottoi broadly overlap with those of C. villosum, C. salvini, and large C. scopaeum (Tables 7 and 8). The braincase is low in C. d. doriae, relative to the length of the skull, and less globose than in the other species (except for C. improvisum; Figs. 19, 20). In C. d. vizottoi, the braincase is relatively higher and more globose than in the nominal subspecies. A sagittal crest was present in every specimen of C. d. doriae we examined (n=71), being conspicuous in 41 (57%), moderate in 25 (35%), and low or vestigial in 5 (7%). In C. d. vizottoi, the sagittal crest was present in all specimens (n=11), conspicuous in ten (90.9%) and moderate or low in one (9.1%). The nasal notch is long, extending well behind the anterior rim of the orbits. The post-orbital processes are moderately developed, more so than in gorgasi and trinitatum, but less than in salvini, scopaeum and villosum (Fig. 19). The posterior palatine process is absent in 51 (78%) of the 65 specimens of C. d. doriae we examined. When present, the palatine process is small and inconspicuous, except for three specimens (4.6%), in which this structure was conspicuous. In C. d. vizottoi, the palatine process was absent in five of nine specimens in our sample. The posterior border of the palate is U-shaped, differing from other species of Chiroderma, which have an even posterior border. Paraoccipital processes are present in C. d. doriae. When skull and mandible are in occlusion, a lateral gap is formed, as in C. salvini, C. scopaeum, C. gorgasi, and C. trinitatum (Fig. 9). The I1s have converging tips that contact each other in most specimens. One (ZUFMS 395) had parallel I1s, with no contact. The P3, in occlusal view, is wider (buccolingually) than long (mesiodistally). P3 touches C, but does not touch P4 (in the similar-sized C. improvisum, P3 and P4 are in contact; see Fig. 19). The P4 has a welldeveloped disto-lingual cingulum (relatively less-developed in the other species). The mandible is robust, with the condylar processes clearly above the toothrow plane, or at approximately the same level. The coronoid process clearly is higher than the tip of the lower canine, when the mandible is viewed laterally (in C. salvini and C. villosum, the coronoid process is at the same level as the canines). The p2 crown is ½ to ⅔ the height of p4, and is ap- proximately as high as long (longer than tall in C. salvini, C. scopaeum, C. improvisum, and C. villosum) (Fig. 20). The p2 does not touch p4. The mandible with a small p2 identified by Oprea & Wilson (2008) as C. doriae (their Figure 2) is a C. villosum (USNM 309905). The only other species of Chiroderma sympatric with C. doriae is C. villosum, from which doriae can be differentiated by its larger size, the both pairs of facial stripes wide and conspicuous (narrow and inconspicuous in villosum), bicolor noseleaf, and ears with pale margins. C. doriae also lacks posterior palatine processes, has converging I1s (usually parallel in villosum), p2 taller than longer (p2 longer than tall in villosum), relatively short c (tall in villosum), and lacks a frontal gap when cranium and mandible are in occlusion (Fig. 14). Geographic Variation and Phylogeography. The phylogenetic analyses of 16 sequences of C. doriae did not show geographic structuring, with haplotypes from geographically distant regions, e.g. from Paraguay north to Rio Grande do Norte, Brazil grouped together (Fig. 21). Although intraspecific genetic variation (0.49%) is considered low (Table 3), the phenotype of C. doriae varies geographically. Specimens from the Caatinga of Piauí and Ceará, and from the Amazonia—Cerrado ecotone in Maranhão, are significantly smaller and have much paler pelage than specimens from the Mata Atlântica and Cerrado, which are larger and have the pelage color varying from pale brown to dark brown (Figs. 18, 20). Given that population samples of C. doriae from the Caatinga (Ceará and Piauí) and Maranhão are phenotypically distinct from those from Mata Atlântica and Cerrado, but do not compose a distinct haplogroup, we treat this population as a subspecies for which the available name is vizottoi, described by Taddei & Lim (2010). Subspecific recognition followed Patten (2015) for phenotypically distinct and geographically restricted groups that do not form clades. Furthermore, geographic variation in C. doriae appears to be correlated with temperature and rainfall (Fig. 22, Tables 11 and 12). Precipitation in driest month (variable BIO14 of WorldClim), precipitation in driest quarter (BIO17), precipitation in warmest quarter (BIO18), temperature seasonality (BIO4), and latitude were the variables having the strongest correlation with size. This suggests that individuals are larger in regions having a more seasonal climate, or at higher latitudes, and in areas with greater rainfall (Appendix 5). Subspecies. We recognize two subspecies in Chiroderma doriae., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 31-37, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Thomas, O. (1891) Note on Chiroderma villosum with the description of a new species for the genus. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 10, 881 - 883.","Dobson, G. E. (1878) Catalogue of the Chiroptera in the collection of the British Museum. British Museum of Natural History, London, xxxvi + 567 pp. https: // doi. org / 10.5962 / bhl. title. 55341","Vieira, C. O. da C. (1942) Ensaio monografico sobre os quiropteros do Brasil. Arquivos de Zoologia, Museu de Zoologia da Universidade de Sao Paulo, 3, 219 - 471.","Gouraud, C., Chevrier, L. & Mearns, R. (2016) Charles and Emile Parzudaki: Natural history dealers in nineteenth-century Paris. Archives of Natural History, 43 (1), 76 - 89. https: // doi. org / 10.3366 / anh. 2016.0347","Oprea, M., Wilson, D. E. (2008) Chiroderma doriae (Chiroptera: Phyllostomidae). Mammalian Species, 186: 1 - 7. https: // doi. org / 10.1644 / 816.1","Taddei, V. A. & Lim, B. K. (2010) A new species of Chiroderma (Chiroptera, Phyllostomidae) from Northeastern Brazil. Brazilian Journal of Biology, 70 (2), 381 - 386. https: // doi. org / 10.1590 / S 1519 - 69842010000200021","Patten, M. A. (2015) Subspecies and the philosophy of science. The Auk, 132 (2), 481 - 485. https: // doi. org / 10.1642 / AUK- 15 - 1.1"]}

Chiroderma scopaeum Handley, 1966 Synonyms: Chiroderma [sp.]: Anderson, 1960: 7. Chiroderma salvini scopaeum Handley, 1966a:297; type locality “ Pueblo Juárez, Colima, México.” Type Material. The type of C. salvini scopaeum, by original designation, is specimen USNM 338711, an adult female collected by Alfred L. Gardner (field number ALG 1565) in Pueblo Juaréz, Mexican state of Colima, on August 19, 1960. The specimen was previously stored in the University of Arizona collection, under the number 7952. The material consists of a stuffed skin with skull and mandible separated. The skin is in good condition, and both pairs of facial stripes are visible. The median dorsal stripe is also visible and located immediately posterior to the nape and extending to the posterior extremity of the animal. The auditory bullae have separated from the skull and upper inner incisors are missing. The posterior palatine process is absent. Distribution and Habitat. We consider C. scopaeum to be restricted to México, west of the isthmus of Tehuantepec (Fig. 11). The species has been recorded in the states of Chihuahua, Sinaloa, Durango, Nayarit, Jalisco, Colima, México, Morelos, Guerrero, Puebla, Veracruz, and Oaxaca (Anderson 1960; Handley 1966a; Crossin et al. 1973; Alvarez & Alvarez-Castañeda 1996; Valiente-Banuet et al. 1997). Hall (1981) suggests that C. scopaeum would occur from western México to northwestern Costa Rica, and based on this distribution Reid & Langtimm (1993) identified specimen USNM 565812 as C. salvini scopaeum. The morphological characters of the specimen, however, have allowed us to identify it as C. salvini. Records of C. scopaeum are from areas dominated by tropical and subtropical coniferous forests, dry deciduous forests at higher elevations, and shrubby vegetation at lower elevations. Studies suggest that in the arid areas of western México, the species would be restricted to the more humid areas close to the Pacific coast and adjacent montane forests, and along the riparian forests in the canyons that cut through the Sierra Madre Occidental (Anderson 1960, 1972; Crossin et al. 1973; García-Mendoza & López-González 2013). All analyzed specimens were collected within the altitudinal range of the species as reported by Handley (1966a), from sea level to 1,722 m. Description and Comparisons. Dorsal pelage varies from pale brown to dark brown. Most of the 38 specimens examined had pale brown pelage (84.2%, n=32), whereas dark brown pelage was found in 15.8% (n=6). Individually, dorsal hairs are tricolored, with a dark brown base, buff middle band, and light to dark brown tips. The medial dorsal stripe was present in all specimens (n=34), but was weakly developed in 5.8% of the sample (n=2). Usually, the dorsal stripe extends from the interscapular region to the posterior extremity of the body, but in 10 specimens the stripe originated in the region immediately posterior to the nape. Both pairs of facial stripes are bright and wide (interocular stripe> 1.7 mm). The tragus and base of the ears are yellowish, as are the anterior and posterior margins of the ears close to the base. The remainder of the ear is brown. The spear of the noseleaf has a simple tip and is brown, except for the lateral margins of the horseshoe, which are whitish. The dimensions of the skull of C. scopaeum are similar to those of C. villosum, and there is also some overlap between the large C. scopaeum and the small C. doriae vizottoi and C. salvini (Tables 7 and 8). In dorsal view, the brain case is round and less massive than in C. salvini. Approximately ⅓ of the length of the nasal notch extends behind the anterior margin of the orbits. The post-orbital constriction is relatively wide (Table 7); post-orbital processes are small and pointed. A sagittal crest was unambiguously present in 32 of the 38 specimens (84.2%), but not detected in 2 (5.2%), or ambiguous in 4 (10.5%). The posterior palatine process was absent in 32 of the 38 analyzed specimens (83.8%), but small or vestigial in the remaining 6 (16.2%). Out of 35 specimens, 30 (85.7%) had convergent I1s, with the tips touching each other; whereas, 5 had both incisors separated along their entire length. The P3 is approximately oval in occlusal outline and is not in contact with P4. The M2 has well defined main cusps, but lacks a posterolingual cingulum. The lower canine has a relatively low crown, below the level of the coronoid process in lateral view. The anterior cingulum of the lower canine projects rostro-medially and is visible in lateral view (Fig. 17). The p2 is small, approximately ¼ of the height of p4; and is longer than tall and does not touch p4. Compared with C. salvini, C. scopaeum can be distinguished by its smaller size, usually paler dorsal pelage (varying from pale brown to dark brown). C. scopaeum has a relatively broader post-orbital constriction (Fig. 12), and rostro-medially projected cingula of lower canines (Fig. 17). From C. villosum, C. scopaeum can be differentiated by its bicolored noseleaf and spear having a simple tip; paler ear margins; shorter nasal notch (in villosum the notch reaches the post-orbital processes); shorter orbits (in villosum the anterior margin is in line with the middle of P4); I1s with convergent tips (usually parallel in villosum); relatively short lower canine (in villosum the tip of the lower canine is at approximately the same level as the coronoid process); and absence of a frontal gap when cranium and mandible are in occlusion (in villosum there is a frontal gap delimited by C, I1–2, and i1–2). The subspecies C. d. vizottoi differs from C. scopaeum by having pale buff pelage, and larger size (Table 7). The p2 of C. d. vizottoi is larger, about ½ to ⅔ of the height of p4, while in C. scopaeum, p2 is approximately ¼ the height of p4. Geographic Variation and Phylogeography. A clade, here identified as scopaeum, contains six specimens of Chiroderma, of which five were analyzed morphologically (Fig. 16). The two specimens from México (TTU 109703 and TTU 110649) are phenotypically similar to the taxon we defined here as Chiroderma scopaeum, whereas specimens from Panamá (LSUMZ 25470), El Salvador (TTU 62462), and Guatemala (ROM 99703) have the diagnostic characters of Chiroderma salvini. The specimens morphologically diagnosed as salvini that nested in the scopaeum clade may represent a case of incomplete lineage sorting (ILS), a relatively common phenomenon in recently-diverged taxa (Maddison & Knowles 2006). To test the ILS hypothesis between C. salvini and C. scopaeum, we recommend increasing the genetic sample of Chiroderma from western México, and obtaining additional genomic information such as single nucleotide polymorphisms. Also, it is important to note that no specimens, morphologically diagnosed as scopaeum, are nested in the salvini clade, which contains sequences from Central and South American specimens. Subspecies. C. scopaeum is monotypic. Remarks. Anderson (1960) mentioned a record of Chiroderma from Chihuahua, western México, that at the time would considerably increase the known distribution of the genus, suggesting an undescribed species for the region. Based on a larger sample size, Handley (1966a) described the subspecies Chiroderma salvini scopaeum, then considered a smaller variant of C. salvini salvini. In this study, we consider the morphological, genetic, and biogeographic evidence as sufficiently strong to treat scopaeum as a species distinct from salvini, instead of as a geographic variant, or subspecies. Natural History. Information on the diet of C. scopaeum is scarce. In Tahuacán, Puebla, one individual was observed visiting the flowers of the columnar cactus Pachycereus weberi (Pachyceraceae), but the bat was not covered in pollen (Valiente-Banuet et al. 1997). In Sinaloa, C. scopaeum was captured in mist nets set under fruiting fig trees. In Jalisco, mist nets over a stream and under a canopy formed by wild figs and other trees also caught C. scopaeum (Jones et al. 1972; Watkins et al. 1972). Specimens have been captured in altered landscapes, such as cornfields (Almazán-Catalán et al. 2009). Summarizing data from the literature, along with the specimens we examined, C. scopaeum appears to be seasonally polyestrous. Pregnancies occurred in January (Sinaloa), February (Jalisco), and June (Jalisco and Nayarit) (Jones et al. 1972; Watkins et al. 1972). Lactating females have been found in May (Morelos) and June (Nayarit and Jalisco) (Watkins et al. 1972). Females noted as non-reproductive were recorded in July (Chihuahua; Anderson 1972) and August (Colima; Wilson 1979). Specimens Examined (N = 35): México: Colima, La Sidra (TTU 61623), Pueblo Juárez (USNM 338711 [holotype of scopaeum]); Jalisco, 20 km SW Talpa de Allende (AMNH 254647), 9.3 km W Chapala (TTU 38049), 6.4 km NW Autlán de Navarro (TTU 109703), La Cumbre (TTU 40987); Morelos, Oaxtepec (USNM 559607); Nayarit, 12.9 km NE San Miguel del Zapote, 51.5 km W Mesa del Nayar (USNM 559608–559613), 13 km NE San Blas (TTU 110649), 5 km E El Venado (USNM 559614, 559615), 12.9 km E San Blas (TTU 6122), Arroyo La Taberna,, 3.2 km W Mesa del Nayar (USNM 511374–511377), 2.9 km NE (by road), Coapan (USNM 511380–511382), 2.3 km N (by road), El Tacote (USNM 508636), 3.2 km E Jalcotoán (USNM 523258, 523259), Mesa del Nayar (USNM 511378, 511379), Playa Novillero (USNM 553885); Oaxaca, 30 km NW Sala de Veja (AMNH 190006); Veracruz, Ojo de Agua del Rio Atoyac (TTU 9996–9999)., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 29-31, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Anderson, S. (1960) Neotropical bats from Western Mexico. University of Kansas Publications, Museum of Natural History, 14, 1 - 8.","Handley Jr., C. O. (1966 a) Descriptions of new bats (Chiroderma and Artibeus) from Mexico. Anales del Instituto de Biologia, Universidad Nacional Autonoma de Mexico, 36, 296 - 301.","Crossin, R. S., Soule, O. H., Webb, R. G. & Baker, R. H. (1973) Biotic relationships in the Canon Del Rio Mezquital, Durango, Mexico. The Southwestern Naturalist, 18 (2), 187 - 200.","Alvarez, T. & Alvarez-Castaneda, S. T. (1996) Aspectos Biologicos y ecologicos de los murcielagos de Ixtapan del Oro, Mexico. In: Genoways, H. H. & Baker, R. J. (Eds.), Contributions in mammalogy: a memorial volume honoring J. Knox Jones, Jr. The Museum, Texas Tech University, Lubbock, Texas, pp. 169 - 182.","Valiente-Banuet, A., Rojas-Martinez, A., Casas, A., Arizmendi, M. del C. & Davila, P. (1997) Pollination biology of two winterblooming giant columnar cacti in the Tehuacan Valley, central Mexico. Journal of Arid Environments, 37 (2), 331 - 341. https: // doi. org / 10.1006 / jare. 1997.0267","Hall, E. R. (1981) The mammals of North America. Vol. 1. John Wiley & Sons, New York, xxx + 546 + 79 pp.","Reid, F. & Langtimm, C. (1993) Distributional and natural history notes for selected mammals from Costa Rica. The Southwestern Naturalist, 38 (3), 299 - 302. https: // doi. org / 10.2307 / 3671441","Anderson, S. (1972) Mammals of Chihuahua: taxonomy and distribution. Bulletin of the American Museum of Natural History, 148, 153 - 410.","Garcia-Mendoza, D. F. & Lopez-Gonzalez, C. (2013) A checklist of the mammals (Mammalia) from Durango, western Mexico. Check List, 9 (2), 313 - 322. https: // doi. org / 10.15560 / 9.2.313","Maddison, W. P. & Knowles, L. L. (2006) Inferring phylogeny despite incomplete lineage sorting. Systematic Biology, 55 (1), 21 - 30. https: // doi. org / 10.1080 / 10635150500354928","Jones Jr., J. K., Choate, J. R. & Cadena, A. (1972) Mammals from the Mexican state of Sinaloa. II. Chiroptera. Occasional Papers of the Museum of Natural History, the University of Kansas, 6, 1 - 29.","Watkins, L., Jones Jr., J. K. & Genoways, H. H. (1972) Bats of Jalisco, Mexico. Museum of Texas Tech University Special Publications 1. Museum of Texas Tech University Press, Lubbock, Texas, 44 pp. https: // doi. org / 10.5962 / bhl. title. 146912","Almazan-Catalan, J. A., Taboada-Salgado, A., Sanchez-Hernandez, C., Romero-Almaraz, M. L., Jimenez-Salmeron, Y. Q. & Guerrero Ibarra, E. (2009) Registros de murcielagos para el estado de Guerrero, Mexico. Acta Zoologica Mexicana, New Series, 25 (1), 177 - 185.","Wilson, D. E. (1979) Reproductive patterns. In: Baker, J. R., Jones Jr., J. K. & Carter, D. C. (Eds.), Biology of the bats of the New World Family Phyllostomatidae. Part III. Museum of Texas Tech University Special Publications 16. Museum of Texas Tech University Press, Lubbock, Texas, pp. 317 - 378."]}

C. v. jesupi Allen, 1900 Synonyms: Chiroderma jesupi Allen, 1900: 88; type locality “ Cacagualito, Colombia.” Chiroderma isthmicum Miller, 1912: 25; type locality “ Cabima, Panama.” Chiroderma isthmica Hall and Jackson, 1953: 645; incorrect gender concordance. Chiroderma villosum jesupi: Handley, 1960:466; first use of current name combination. Distribution and Habitat. The subspecies C. v. jesupi occurs in the tropical region of trans-Andean South America and also in Central America and México. The taxon has been recorded in México (Oaxaca, Hidalgo, Veracruz, and states to the south), Guatemala, Belize, El Salvador, Honduras, Nicaragua, Costa Rica, Panamá (including Islas Perlas), western Colombia, western Ecuador, and extreme northwestern Perú (Tumbes; Fig. 28). Collecting sites of C. v. jesupi are in dry and humid tropical forests. The taxon has been recorded from sea level up to approximately 970 m. In northwestern Perú (Tumbes), specimens were captured at 350 m (Novoa et al. 2011). In Colombia (Tolima), the taxon was captured at 900 m (Galindo-Espinosa et al. 2010). In southern México (Chiapas), the maximum capture elevation was 915 m (Davis et al. 1964). Remarks. The original description of Chiroderma villosum was based on specimen ZMB 408 and on a skeleton from the anatomical collection of the Berlin museum that is probably lost (Turni & Kock 2008; Garbino & Nogueira 2017). From the two syntypes, Turni & Kock (2008: 44) selected specimen ZMB 408 as the lectotype, but Thomas (1891: 882) had already designated the same specimen, i.e. the one represented in Peters’ plate, as the lectotype. In the species description, Peters (1860: 748) mentioned only “Brasilia” as the locality and suggested that the lectotype was collected by Friedrich Sellow. In the collection catalogue of the Museum für Naturkunde, the locality of the specimen, handwritten by Peters reads just “ Brasilien ”, and there is no note indicating who collected it (Garbino & Nogueira 2017). Due to the impossibility to further restrict the type locality and the lack of evidence that it was indeed collected by Sellow, we follow most authors in citing “ Brazil ” as the type locality of Chiroderma villosum (Handley 1960; Husson 1962; Carter & Dolan 1978; Simmons 2005; Gardner 2008a; Turni & Kock 2008; Garbino & Nogueira 2017). Cabrera (1958: 85) erroneously designated “ Venezuela ” as the type locality of Chiroderma villosum, followed by Goodwin & Greenhall (1961), and probably by Vieira (1942, 1955), who did not include the species among the Brazilian mammals. Cabrera may have altered the type locality based on the mention of a specimen from St. Esteban, Venezuela by Thomas (1891: 56), which was the first subsequent precise locality reported for the species. Natural History. C. villosum is a secondarily granivorous frugivore, specializing in chewing the seeds of fruits of Ficus to extract nutritive content (Nogueira & Peracchi 2003; Wagner et al. 2015). The species apparently has preference for Ficus, and fruits of this genus may compose 100% of the diet of C. villosum in Panamá (Bonaccorso 1979). Nine species of Ficus have been recorded in the diet of C. villosum in Panamá: Ficus citrifolia, F. dugandii, F. insipida, F. nymphaefolia, F. obtusifolia, F. paraensis, F. pertusa, F. popenoei, and F. trigonata (Bonaccorso 1979; Handley et al. 1991; Wendeln et al. 2000; Wagner et al. 2015). Consumption of infructescences of Cecropia obtusa was recorded in the diet of C. villosum from French Guiana (Lobova et al. 2003; Suárez-Castro & Montenegro 2015). C. villosum have been captured at mineral-rich clay licks (“collpas”) in the Peruvian Amazon (Bravo et al. 2008; Ghanem & Voigt 2014). In the Guianas, Amazonia, and in the Atlantic rainforest, the species was captured more frequently in the canopy, suggesting that C. villosum is a canopy frugivore (Ascorra et al. 1996; Simmons & Voss 1998; Charles- Dominique & Cockle 2001; Kalko & Handley Jr. 2001; Delaval et al. 2005; Gregorin et al. 2017). The only natural day roost recorded for the species are tree hollows in Venezuela (Handley 1976). In addition to natural shelters, C. villosum has been found in buildings in Brazil, Panamá, and Venezuela (Goldman 1920; Handley 1976; data from the DZSJRP collection catalogue). The following ectoparasites have been documented on C. v. jesupi: Aspidoptera busckii (Streblidae), Trichobius joblingi (Streblidae), Paratrichobius sp. A (Wenzel et al. 1966). In Venezuela, two species of streblid bat flies (Trichobius parasiticus and Metelasmus pseudopterus), two spinturnicid mites (Periglischurus acustidens and P. iheringi), and a species of trombiculid mite (Whartonia nudosetosa), are known from C. v. villosum (Herrin & Tipton 1975; Reed & Brennan 1975; Wenzel 1976). Marinkelle & Grose (1981) recorded Megistopoda proxima and Strebla carolliae, two species of streblid bat flies from Colombian C. villosum. The absence or rarity of spinturnicid mites on C. villosum was noted in some studies: in Panamá no mite was recorded on any specimen, in the Peruvian Amazon no mite was collected from 33 specimens, and only two mites were collected from a sample of 724 Venezuelan C. villosum (Furman 1966; Herrin & Tipton 1975; Gettinger 2018). Two flesh fly larvae of the genus Sarcophaga (Sarcophagidae), were found in the abdominal cavity of a freshly-dead female, suggesting that they parasitized the live animal (Goodwin & Greenhall 1961). Blood parasites of the family Trypanosomatidae, subfamily Leishmaninae, are known from C. v. villosum from central Brazil, and Trypanosoma (Schizotrypanum) is documented in this species from Colombia (Marinkelle 1982; Lourenço et al. 2018). One C. v. villosum from southeastern Brazil had traces of hantavirus infection, making C. villosum one of the few species of frugivorous bat to host this virus (Sabino-Santos et al. 2015). Reproductive data from specimens we examined and from the literature (Davis et al. 1964; Jones et al. 1971; Taddei 1976; Bonaccorso 1979; Anderson 1997) suggest seasonal polyestry. In Central America (Nicaragua and Panamá), pregnant females were recorded in January, February, and March, just before the beginning of the rainy season, and lactating C. villosum were found in February, March, and April. In South America, pregnancies were also recorded before the rainy season in July and August (Rondônia, Brazil) and in August and September (Bolivian, Ecuadorian, and Peruvian Amazon). In southeastern Brazil (São Paulo), C. villosum were recorded as pregnant in July and August. In Venezuela, pregnancies were recorded both before the rainy season in January, February, March, and April, and during the rainy season in June, and July; whereas, lactating individuals were documented in February, June, and July. Specimens Examined (N = 509): Belize: Toledo, Bladen Nature Reserve (USNM 583035, 583036). Bolivia: Beni, Río Iténez (AMNH 209529–209533), Vacadiaz (USNM 390606); La Paz, Santa Ana de Madidi (AMNH 261677); Pando, Independencia (AMNH 262526–262529), Río Nareuda (248884, 248885); Santa Cruz, Buenavista (AMNH 61754), Parque Nacional Noel Kempff Mercado (AMNH 264078, 264079). Brazil: without specific locality (ZMB 408 [lectotype of villosum]); Acre, Seringal Lagoinha (DZSJRP 13029–13033), Parque Nacional da Serra do Divisor (ALP 7011, 7018, 7022, 7023, 7050, 7059, 7160, 7308); Alagoas, Mata de Coimbra (UFPB 4349); Amazonas, Comunidade Cachoeirinha (LMUSP [ICA048]), Humaitá (DZSJRP 14793), Igarapé Taracuá (LMUSP [JAP76]), km 27 of BR319 (DZSJRP 14121, 14651), km 5 of BR230 (DZSJRP 13134), opposite to Comunidade São Pedro (LMUSP [ICA169], LMUSP [ICA170]), Vila de Santa Fé (LMUSP [JAP84]); Bahia, Ilhéus (CMU- FLA 1076, 1078, 1119); Espírito Santo, Aracruz Celulose (MZUSP 35032), Fazenda Santa Terezinha (MZUSP 35030, 35031), Reserva Natural Vale (ALP 2806, 2810, 3009, 3249, 3327, 3408, 4560, 4758); Mato Grosso, 264 km N Xavantina (USNM 393712–393714), Aricá (MZUSP 6494), Cláudia (MZUSP [PEV 1225-1226]), Nossa Senhora do Livramento (UFMT 1146, 1147), Parque Nacional do Juruena (CMUFLA 1290, 1299), Sinop (ALP 3419), U.H.E. foz do Apiacás (UFMT 1952, 1953), Usina Teles Pires (UFMT 2137, 2138); Mato Grosso do Sul, Estação Ecológica Dahma (ZUFMS 492), Maciço do Urucum (ZUFMS 208, 209), Pantanal de Aquidauana (ZUFMS 1904), Pantanal de Nhecolândia (ZUFMS 244, 1896); Minas Gerais, Dores do Indaiá (UFMG 3760), Estação Ecológica de Pirapitinga (ALP 9166, 9370), Fazenda Cabriúna (CMUFLA 465), Fazenda Marinheiro (DZSJRP 14480), Parque Nacional do Peruaçu (CMUFLA 907, 1532–1536, 1676–1678, 1680, 1682, 1684, 1686), Parque Estadual do Rio Doce (CMUFLA 1158, 1161, 1834, 1839–1841); Pará, 52 km SSW Altamira (MZUSP 22677), Centro Kaiapó de Estudos Ecológicos (MZUSP 29150–29152), Floresta Nacional de Carajás (UFMG [VCT6305]), Floresta Nacional de Carajás, Serra Norte (UFMG [VCT2303]), Floresta Nacional de Carajás, Serra Sul, Corpo A (UFMG [VCT1959], UFMG [VCT5157]), Floresta Nacional de Carajás, Serra Sul, Corpo C (UFMG [VCT2019], UFMG [VCT5073], UFMG [VCT5079]), Lago Jacaré (MZUSP 13335), Lago Leonardo (MZUSP 13209, 13197), Platô Greig (UFMG 3258), Platô Monte Branco (UFMG 3244), Porto Trombetas (UFMG 3212, 3214, 3221, 3246), Projeto Alemão (UFMG [VCT4394], UFMG [VCT4444]), Rio Xingu, left bank (MPEG 4112), Sta.A, IAN (USNM 361724, 361725), Várzea, Belém (USNM 460128–460130); Paraíba, Fazenda Pacatuba (UFPB 4), João Pessoa (UFPB 10333, 10335, 10336); Pernambuco, Recife (UFMG [M1], UFMG [M2], UFMG [M3]); Piauí, Boqueirão da Esperança (ZUEC 2066), Parque Nacional da Serra das Confusões (MZUSP 33502); Rio de Janeiro, Jardim Botânico (ALP 7419– 7424, 7426–7431, 8278), Parque Estadual Serra da Tiririca (ALP 5578, 5579); Rondônia, Calama (AMNH 37041), Costa Marques (AMNH 209575), Pedra Branca (MZUSP 22827), Pedras Negras (AMNH 209576), Cachoeira de Nazaré, Rio Machado (MZUSP 20200, 20201), U.H.E. Jirau (MZUSP 35408); Roraima, Estação Ecológica da Ilha de Maracá (DZSJRP 11487); São Paulo, São José do Rio Preto (DZSJRP 4586, 4676), Engenheiro Schmidt (DZS- JRP 4804), Fazenda São Paulo (DZSJRP 14302, 14305, 14319), Roberto (DZSJRP 16549), Parque Natural Municipal Grota de Mirassol (DZSJRP 4000, 4032, 4033), Fazenda Paraguassu (ZUEC 968), Sítio Progresso (DZSJRP 3783, 3922, 4337, 4386–4390); Sergipe, Estação Ecológica de Itabaiana (CMUFS 64), Mata do Junco (CMUFS 97), Parque Nacional Serra de Itabaiana (ALP 8971); Tocantins, U.H.E. São Salvador (MZUSP [SSI178]). Colombia: no specific locality (USNM 598086); Amazonas, Puerto Nariño (USNM 483777); Antioquia, La Tirana (IAvH-M 930, USNM 449480–499482), Vereda La Pola, Parque Nacional Natural Los Katíos (IAvH-M 4923); Chocó, Corregimiento Gilgal, P.N.N. Los Katíos (IAvH-M 4924–4926), Reserva Florestal Especial Las Teresitas (IAvH-M 3257), Vereda El Tilupo, P.N.N. Los Katíos (IAvH-M 4927–4929); Magdalena, Cagualito (AMNH 14574 [holotype of jesupi]), Parque Nacional Natural Tayrona (IAvH-M 4198), Vereda El Congo (IAvH-M M-9665); Putumayo, Caño Caucayá (IAvH-M M-624); Sucre, Estación Primates (IAvH-M 9583), Quebrada El Sereno (IAvH-M 9593); Valle del Cauca, Río Zabaletas (USNM 483770–483776); Vaupés, Caño Arara (IAvH-M 1550). Costa Rica : Puntarenas, Corcovado National Park (USNM 565813). El Salvador: La Libertad, Deininger Park (TTU 63906); La Paz, Hacienda Escuintla (TTU 63911); La Unión, El Tamarindo (TTU 63912). Ecuador: Los Ríos, Beata Elvira (USNM 498921, 498922), El Papayo (USNM 498923, 522435–522437), Hacienda Santa Teresita (USNM 522438), Lima Pareja (USNM 498924, 498925, 522434), Río Nuevo (USNM 534315, 534316), Vinces (USNM 534314); Pastaza, Lorocachi (USNM 548240, 548241), Santiago (not located; USNM 548242), Tiguino (USNM 574537, 574539), Yosa (not located, USNM 548237–548239); Pichincha, Río Palenque Science Center (USNM 528541). Guatemala: Jutiapa, Colonia Montufar (AMNH 217417), Santa Rosa, La Avellana (AMNH 235312–235315). Guyana — Barima-Waini, North West, Santa Cruz (ROM 98850); Cuyuni-Mazaruni, 24 km along Potaro road from Bartica (BMNH 1965.645), Namai Creek (ROM 108219); Upper Demerara-Berbice, Dubulay Ranch (USNM 582328); Upper Takutu-Upper Essequibo, Chodikar River (ROM 106644). French Guiana: Sinnamary, Paracou (AMNH 267190, 267191, 268534 – 268536). México: Veracruz, Xalapa (= Jalapa) (BMNH 81.10.27.1). Nicaragua — Zelaya (currently Región Autónoma de la Costa Caribe Sur), 4,5 km NW Rama (TTU 12794). Panamá — Bocas del Toro, Almirante (USNM 315559–315562), Isla Popa (USNM 464304), Sibube (USNM 335298, 335299); Colón, Bohio Peninsula (USNM 503637); Darién, Cana (USNM 179619), El Real (USNM 338045), Jaqué (USNM 362920), Punta Piña (USNM 314718), Tacarcuna Village Camp (USNM 309894, 309896–309900); Los Santos, Cerro Hoya (USNM 323451–323453), Guánico Arriba (USNM 323448–323450); Panamá, Barro Colorado Island (USNM 304904, 304905, 304907–304909, 332053, 457954, 544896), Cabima (USNM 173834 [holotype of isthmicum], USNM 173836), Cerro Azul (USNM 305386), Culebra (USNM 223402), Gamboa (USNM 520557, 520558), Isla San José (USNM 448449), Punta de Cocos (USNM 314719, 314720, 314721), Río Mandinga (USNM 305385); San Blas (currently Guna Yala), Armila (USNM 335300–335316); Veraguas, Isla Cébaco (USNM 360219). Perú: Cusco, Camisea (USNM 582836), Camisea, Armihuari (MUSM 13606, 13631, 13634, 13636, 13637, 13639– 13641), Camisea, Konkariari (MUSM 14742), Camisea, Pagoreni (MUSM 13610, 13615, USNM 582839), Camisea, San Martín (MUSM 13618, 13644, USNM 582840, 582841), Jenaro Herrera (MUSM 6791), Ridge Camp (USNM 588033); Loreto, 13,6 km NW Albarenga (MUSM 26545), Alto Río Pauya (MUSM 17734), Centro de Investigaciones Jenaro Herrera (MUSM 4221, 4222), Estación Biologica Allpahuayo (MUSM 16476), Ninarumi (MUSM 29560, 29561), Paujil (MUSM 29562), Puesto de Vigilancia Castaña (MUSM 21136), Quistococha (USNM 337940), Río Lagartococha (MUSM 21137), Río Pisqui, Campamento Llanura (MUSM 17735), Río Samiria (MUSM 29562), Río Samiria, Flor de Yarina (MUSM 1637), Río Samiria, Tacshacocha (MUSM 1638, 1639), San Lorenzo (BMNH 1924.3.1.75, 1924.3.1.76), Zungarococha (MUSM 29563); Madre de Dios, CICRA (MUSM 26106), Cocha Salvador (MUSM 733), Estación Biológica Cocha Cashu (MUSM 15856), Explorer’s Inn Lodge (MUSM 1640), Hacienda Amazonia (MUSM 9757), Maskoitania (MUSM 19671), Pakitza (MUSM 6781, USNM 566544), Quebrada Aguas Calientes (MUSM 16653, 16660, 16661, 16665–16667), Reserva Cuzco Amazónico (MUSM 6168, 6169), Santuario Nacional Pampas del Heath (MUSM 12827); Pasco, Campamento Río Lobo (MUSM 10232), Cerro Chontiya (MUSM 10225, 10230, 10231), San Juan (USNM 364418), Villa America (MUSM 1641); San Martín, Concesion de Conservación Valle del Biavo (MUSM 43843, 43844, 43485), Juanjuí (MUSM 1642), Saposoa (MUSM 1643), Yurac Yacu (BMNH 27.1.1.63); Tumbes, Carrizalillo (MUSM 22123), Parque Nacional Cerros de Amotape (MUSM 22121, 22122), Quebrada Las Pavas (MUSM 24479); Ucayali, 59 km W Pucallpa (USNM 461256), Concesión de Conservación Río La Novia (MUSM 44186, 44187, 44472). Suriname: Brokopondo, Brownsberg Nature Park (ROM 114212); Sipaliwini, Kushere Landing (ROM 120226). Trinidad and Tobago: Tobago, Charlotteville (USNM 540676); Trinidad, without specific locality (AMNH 256325), Diego Martin (AMNH 183167), Grande Riviere (AMNH 172149), Guaico Tamana (AMNH 172148), Maracas Valley (AMNH 175599), Waterloo (BMNH 1897.6.7.44). Venezuela: Amazonas, 9 km SE Puerto Ayacucho (MZUSP 27168), Cerro Neblina base camp (USNM 560606, 560607, 560767, 560768, 560769, 560770, 560771, 560772), Río Cunucunuma (USNM 405127, 405165, 405167–05171), Río Manapiare (USNM 408624–408660, 408662–408678), Río Mavaca (USNM 405175), San Carlos de Rio Negro (USNM 560604, 560605), Tamatama (USNM 405176, 405177, 408611–408617); Bolívar, El Manaco (USNM 387205–387209, 387212, 387213, 387126, 387210, 387211, 387214, 387215), Hato La Florida (USNM 405164), Hato San José (USNM 405162), Río Supamo (USNM 387204); Sucre, Manacal (USNM 408620); Yaracuy, 19 km NW Urama (USNM 372145, 372146, 372149, 372150)., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 54-56, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Allen, J. A. 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Brigham Young University Science Bulletin, Biological Series, 20 (1, part 1), 1 - 42.","Wenzel, R. L. (1976) The streblid batflies of Venezuela (Diptera: Streblidae). Brigham Young University Science Bulletin, Biological Series, 20, 1 - 177.","Marinkelle, C. J. & Grose, E. S. (1981) A list of ectoparasites of Colombian bats. Revista de Biologia Tropical, 29 (1), 11 - 20.","Furman, D. P. (1966) The spinturnicid mites of Panama (Acarina: Spinturnicidae). In: Wenzel, R. L. & Tipton, V. J. (Eds.), Ectoparasites of Panama. Field Musem of Natural History, Chicago, Illinois, pp. 125 - 166.","Gettinger, D. D. (2018) Checklist of bloodfeeding mites (Acari: Spinturnicidae) from the wings of bats (Mammalia: Chiroptera) in the Manu Biosphere Reserve, Peru. MANTER: Journal of Parasite Biodiversity, 10, 1 - 9. https: // doi. org / 10.13014 / K 2 DJ 5 CVZ","Marinkelle, C. J. (1982) Prevalence of trypanosoma cruzi-like infection of Colombian bats. Annals of Tropical Medicine and Parasitology, 76 (2), 125 - 134. https: // doi. org / 10.1080 / 00034983.1982.11687517","Lourenco, J. L. M., Minuzzi-Sauza, T. T. C., Silva, L. R., Olivei, A. C., Mendonca, V. J., Nitz, N., Aquiar, L. M. S. & Gurgel-Goncalves, R. (2018) High frequency of trypanosomatids in gallery forest bats of a Neotropical savanna. Acta Tropica, 177, 200 - 206. https: // doi. org / 10.1016 / j. actatropica. 2017.10.012","Sabino-Santos Jr., G., Maia, F. G. M., Viera, T. M., Muylaert, R. L., Lima, S. M., Goncalves, C. B., Barroso, P. D., Melo, M. N., Jonsson, C. B., Goodin, D., Salazar-Bravo, J. & Figueiredo, L. T. M. (2015) Evidence of hantavirus infection among bats in Brazil. American Journal of Tropical Medicine and Hygiene, 93 (2), 404 - 406. https: // doi. org / 10.4269 / ajtmh. 15 - 0032","Jones Jr., J. K. Smith, J. D. & Turner, R. W. (1971) Noteworthy records of bats from Nicaragua, with a checklist of the chiropteran fauna of the country. Occasional papers of the Museum of Natural History, the University of Kansas, 2, 1 - 35.","Taddei, V. A. (1976) The reproduction of some Phyllostomidae (Chiroptera) from the northwestern region of the State of Sao Paulo. Boletim de Zoologia, Universidade de Sao Paulo, 1 (1), 313 - 330. https: // doi. org / 10.11606 / issn. 2526 - 3358. bolzoo. 1976.121587","Anderson, S. (1997) Mammals of Bolivia, taxonomy and distribution. Bulletin of the American Museum of Natural History, 231, 1 - 652."]}

C. d. vizottoi Taddei and Lim, 2010 Synonyms: Chiroderma nov. sp. Cruz, Martínez, and Fernandes, 2007: 615. Chiroderma sp. Gregorin, Carmignotto, and Percequillo, 2008: 372. Chiroderma vizottoi Taddei and Lim, 2010: 384; type locality “Teresina (ca. 05° 02 ′ S and 42° 45 ′ W), in the State of Piauí, Brazil.” Distribution and Habitat. This subspecies is in the Brazilian states of Ceará, Maranhão, and Piauí (Fig. 23). A specimen from the Caatinga of Piauí was captured in an area with low xerophytic trees (3 to 5 meters high), next to a rocky outcrop (Gregorin et al. 2008). Two specimens from Maranhão were captured in mist nets set on the beach, in a mosaic of sandbank and mangrove vegetation. Remarks. Lund (1842a: 134, 1842b: 200), in his studies on the extant and fossil fauna of Lagoa Santa, Minas Gerais, recorded Phyllostoma dorsale among the living species of the region. In both publications, Lund clearly uses Phyllostoma dorsale as a new name, as evidenced by the “m.” (= mihi) following the name. Because the name is present only in a faunal list, and no formal description is given, Phyllostoma dorsale Lund is considered a nomen nudum for nomenclatural purposes (see article 12 of the ICZN 1999). Gray (1866), in his revision of Phyllostomidae, listed two species in Chiroderma, C. villosum and C. pictum. The latter name is a nomen nudum, because it lacks a description. We include Chiroderma pictum in the synonymy of Chiroderma doriae because, in the Dobson’s (1878) catalogue of bats in the British Museum, the only specimen of Chiroderma in the collection at the time of Gray’s (1866) publication was the specimen that later became the type of Chiroderma doriae. The Latin word pictus means “painted”, or “decorated”, suggesting that Gray referred to the conspicuous facial stripes, observable that specimen, a characteristic differentiating this species from Peters’ C. villosum, which was the only other Chiroderma known at that time. Dobson (1878), in his description of Chiroderma salvini, compared the type with a specimen from Minas Gerais (Brazil) identified by him as Chiroderma villosum. However, Thomas (1891), based on an unpublished plate by Wilhelm Peters, available in the Genova Natural History Museum, recognized that the Minas Gerais specimen Dobson identified as Chiroderma villosum did not represent Chiroderma villosum Peters, 1860. Thomas (1891) described that specimen as Chiroderma doriae. Winge (1892), studying the material sent by Lund to the Copenhagen museum, identified as Chiroderma villosum the material referred to Lund as Phyllostoma dorsale. Winge (1892: 9) located, among the Lagoa Santa specimens representing extant taxa, five fluid-preserved specimens, two skeletons, and three skins. Among the fossil material, Winge mentioned only post-cranial elements from Lapa da Escrivaninha and a cave he did not identify. Thomas (1893), after receiving part of the Lagoa Santa material Winge identified as Chiroderma villosum, correctly identified the skull as belonging to C. doriae. This publication is the first to ascertain that Phyllostoma dorsale of Lund was the same taxon that Thomas (1891) described as Chiroderma doriae. The specimen collected by Lund at Lagoa Santa (BMNH 93.1.9.16) and examined by us is probably the same specimen donated by Winge to the British Museum and mentioned by Thomas (1893). Aside from the Lagoa Santa specimens and the holotype, all from Minas Gerais, no additional specimen of C. doriae were mentioned in the literature for 86 years. Probably for this reason, publications that mentioned the species cited only Minas Gerais as a locality of occurrence and repeated the morphological characteristics mentioned in the original description (Vieira 1942; Cabrera 1958). It is important to note, however, that the BMNH houses specimens of C. doriae collected in the beginning of the 20 th century, by Alphonse Robert in Ypanema, currently Floresta Nacional de Ipanema, São Paulo (BMNH 3.7.1.163), and by Wilhelm Ehrhardt in Joinville, Santa Catarina (BMNH 9.11.19.15). Taddei (1979) published the first study including new data on C. doriae since Thomas’s report in 1893. When he redescribed the taxon based on 39 specimens of C. d. doriae collected in northwestern São Paulo, Taddei (1979) confirmed the presence of a conspicuous dorsal stripe previous authors had considered to be absent. Natural History. C. d. doriae is secondarily granivorous, specialized in chewing and grinding seeds of the fruits of Ficus to extract their nutritional content (Taddei 1980; Nogueira & Peracchi 2001, 2003). Four species of Ficus have been documented in the diet of C. d. doriae: Ficus clusiaefolia, F. cyclophylla, F. organensis, and F. tomentella (Sipinski & Reis 1995; Esbérard et al. 1996; Nogueira & Peracchi 2001). Individuals of C. d. doriae have been captured near to or visiting fruiting trees of Cecropia glaziovii (Urticaceae), Ficus enormis, F. gomeleira, and Muntingia calabura (Muntingiaceae), suggesting that this bat also consumes the fruits of these species (Taddei 1980; Esbérard et al. 1996). In addition to figs, fruits and infructescences of Cecropia pachystachya, Chlorophora tinctoria (Moraceae), Piper sp., and Psychotria suterella (Rubiaceae) are also consumed (Taddei 1973; Esbérard et al. 1996; Nogueira & Peracchi 2001; Novaes et al. 2010; Laurindo et al. 2017). Individuals of C. d. doriae have been found covered in the pollen of Mabea fistulifera (Euphorbiaceae) and of an unidentified species, suggesting that the bat also feeds on flowers (Esbérard et al. 1996; Olmos & Boulhosa 2000). Day roosts used by C. d. doriae are not known (see review in Garbino & Tavares 2018b), but there are two records from caves, the use of which may be occasional: one in the state of São Paulo (Arnone 2008) and the other in Minas Gerais (information on specimen UFMG 4953). In the Cerrado of São Carlos, southeastern Brazil, the species was found in pellets of the Stygian Owl (Asio stygius) by Motta-Junior & Taddei (1992). At localities in southeastern Brazil where they occur in sympatry, C. d. doriae is usually captured in much higher numbers than C. v. villosum when mist nets are set at ground level. At the Rio de Janeiro Botanical Garden, eight nights of ground-level mist netting resulted in the capture of 49 C. d. doriae and 5 C. v. villosum (Nogueira & Peracchi 2003). In Mirassol, São Paulo, 24 C. d. doriae and 6 C. v. villosum were captured in ground-level nets (Taddei 1979). In contrast, Gregorin et al. (2017), using canopy mist nets placed between 11 and 19.5 m above ground in the Parque Estadual do Rio Doce, Minas Gerais, captured 1 C. d. doriae and 24 C. v. villosum. The anecdotical report of Taddei (1980) describes the approach by C. d. doriae to fruiting trees as a low flight. Although the literature on the ecology of C. d. doriae is scarce when compared to that for C. gorgasi, C. trinitatum, and C. villosum, it suggests that the species is not as much a canopy frugivore as the other species. Five species of ectoparasites are known from C. d. doriae: Aspidoptera falcata (Streblidae), Megistopoda proxima (Streblidae), Strebla guajiro (Streblidae), and Trichobius joblingi (Streblidae) in Sergipe; and the mite Periglischrus iheringi (Spinturnicidae) recorded on individuals from Mato Grosso do Sul, Rio de Janeiro, and in Paraguay (Presley 2004; Lourenço et al. 2016; Lima Silva et al. 2017; Soares et al. 2017). Lourenço et al. (2018) found a trypanosomatid parasite in the blood of C. d. doriae from Distrito Federal, Brazil. The reproductive pattern is seasonal polyestry (Taddei 1980). Pregnancies were recorded in June through October, with earlier stages reported in June and July, and females with advanced fetuses recorded from August through October (Taddei 1980; Esbérard et al. 1996). Births of single young occur between October and January, suggesting a gestation period of at least three and half months (Taddei 1980). Females simultaneously pregnant and lactating are common, indicating postpartum estrus and the production of two young per year, with a second birth period in February and March (Taddei 1976, 1980; Esbérard et al. 1996). The natural history of C. d. vizottoi is poorly known. In Ceará, seven specimens were mist netted close to an unidentified fruiting Moraceae; one was pregnant and another lactating when captured in January (Silva et al. 2015). In Maranhão, a male and female were caught in a mist net placed along the beach in a small fishing community next to a mangrove swamp near regenerating rain forest and a “babaçual” (Attalea speciosa [Arecaceae]) palm grove. The mist-net was set in front of a tree locally called “agarra” or “amapá” (Apocynaceae?), and one of the bats had its abdomen covered with a sticky material resembling a milky sap, or latex. Almeida et al. (2016) recorded the mite Periglischrus iheringi (Spinturnicidae) on C. d. vizottoi in Ceará. Specimens Examined (N = 136): Brazil: Alagoas, Mata da Cachoeira, São José da Lage (UFPB 4348); Bahia, Reserva Particular do Patrimônio Natural Serra Bonita ([UFMG] RSB 21, [UFMG] RSB 22); Ceará, Reserva Natural Serra das Almas (ALP 10196, 10418, 10421, 10423, 10440, 10451, 10464); Goiás, Itumbiara (MCN-MQ 145); Maranhão, São Luís (ALP 6633–6635, [UFMG] VCT 373, 374); Mato Grosso do Sul, Corumbá ([UFMG] VCT 6069, [UFMG] VCT 6081), Fazenda Barma (MZUSP 28591, 28688), Morro do Paxixi (ZUFMS 2300), Parque Estadual Matas do Segredo (ZUFMS 493), Urucum (ZUFMS 800, 1058–1062, 1069), Urucum, Morro São Domingos (ZUFMS 912); Minas Gerais, without specific locality (BMNH 44.9.2.6 [holotype of doriae]), Área de Proteção Ambiental Coqueiral (CMUFLA 163), Barão de Cocais ([UFMG] VCT 5831), Belo Horizonte (UFMG 3537), Estação Ecológica de Pirapitinga in Morada Nova de Minas (UFMG 3383), Estação Ecológica de Pirapitinga in Três Marias (ALP 9154), Gruta do Salitre in Diamantina (UFMG 4953), João Monlevade (CMUFLA 965), Lagoa Santa (BMNH 93.1.9.16); Mariana (CMUFLA 938, [UFMG] VCT 6058), Parque Estadual do Rio Doce (CMUFLA 1157), Pompéu (MCN-MQ 253), Usina Hidrelétrica Retiro Baixo (CMUFLA 1415), Valos (CMFULA 496); Pará (not mapped; BMNH 7.1.1.723 [probably an error]); Paraíba, Fazenda Pacatuba, 10 km NE Sapé (UFPB 3), Reserva Biológica Guaribas (UFPB 7327, 7341); Pernambuco, Reserva de Saltinho ([MZUSP] AD119); Piauí, Parque Nacional de Sete Cidades (DZSJRP 11460 [paratype of vizottoi]), Teresina (DZSJRP 18054 [holotype of vizottoi]); Rio de Janeiro, Ilha da Marambaia (ALP 6121), Jardim Botânico do Rio de Janeiro (7508–7511, 7513–7515, 7517, 7519, 7520, 8059, 8062, 8064, 8077, 8079–8081), Morro Azul (ALP 9142), Parque Estadual da Pedra Branca (ALP 5784), Parque Natural Municipal da Prainha (ALP 6650); Rio Grande do Norte, Mata da Estrela (MZUSP 35027); Santa Catarina, Joinville (BMNH 9.11.19.15); São Paulo, Barra do Ribeirão Onça Parda (MZUSP 10632), Barão Geraldo (ZUEC 783), Cachoeira dos Índios (DZSJRP 3140, 3141), Cananéia (MZUSP 26354), Estação Experimental de Pindorama (DZSJRP 16506), Fazenda João XXIII ([LMUSP] GTG 01), Fazenda Paraguassu (ZUEC 1002), Fazenda Santa Carlota (MZUSP 35028), Fazenda Silvio Fazoli in Irapuã (DZSJRP 2924, 2937, 2945, 2946, 3611, 3728), Grota de Mirassol (DZSJRP 2640, 3997, 3998, 3999, 4029), Iguape (MZUSP 21802), Ilha do Cardoso (MZUSP 28037, ZUFMS 395, 397), Instituto de Biologia Marinha in São Sebastião (DZSJRP 10050), Itapetininga (USNM 542616), Parque Estadual da Ilha Anchieta (MZUSP 29456, 31582), Parque Estadual de Ilhabela (MZUSP 35029), Parque Estadual Turístico do Alto Ribeira (MZUSP 34012), São José do Rio Preto (DZSJRP 2469), São Roque (MZUSP 15112), Sítio Progresso (DZSJRP 3810, 3825, 3921, 4271–4273, 4381–4385, 4445), Ypanema (BMNH 3.7.1.163), Sergipe, Estação Ecológica de Itabaiana (ALP 6545), Floresta Nacional do Ibura (CMUFS SSB1 - FNI, SSB2 - FNI, SSB376 - FNI, SSB377 - FNI, SSB20 - FNI, SSB33 - FNI), Mata do Junco (CMUFS 51, 71, 73, 94), Parque Nacional Serra de Itabaiana (CMUFS 65). Paraguay: La Cordillera, Estancia Sombrero (TTU 75275, 99569); San Pedro, Yaguarete Forests, 0.5 km W headquarters (TTU 95747). Locality unknown: BMNH 7.1.1.698., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 37-39, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Taddei, V. A. & Lim, B. K. (2010) A new species of Chiroderma (Chiroptera, Phyllostomidae) from Northeastern Brazil. Brazilian Journal of Biology, 70 (2), 381 - 386. https: // doi. org / 10.1590 / S 1519 - 69842010000200021","Cruz, L. D., Martinez, C. & Fernandes, F. R. (2007) Comunidades de morcegos em habitats de uma Mata Amazonica remanescente na Ilha de Sao Luis, Maranhao. Acta Amazonica, 37 (4), 613 - 619. https: // doi. org / 10.1590 / S 0044 - 59672007000400017","Gregorin, R., Carmignotto, A. P. & Percequillo, A. R. (2008) Quiropteros do Parque Nacional da Serra das Confusoes, Piaui, nordeste do Brasil. Chiroptera Neotropical, 14 (1), 366 - 383.","Lund, P. W. (1842 a) Fortsatte Bemaerkninger over Brasiliens uddode Dyrskabning. Det Kongelige Danske videnskabernes selskabs skrifter, 9, 123 - 136.","Lund, P. W. (1842 b) Blik paa Brasiliens Dyreverden for sidste Jordomvaeltning. Det Kongelige Danske videnskabernes selskabs skrifter, 9, 140 - 208, pls 28 - 38.","ICZN (1999) International Code of Zoological Nomenclature. 4 th Edition. International Trust for Zoological Nomenclature, London, xxix + 306 pp.","Gray, J. E. (1866) Revision of the genera of Phyllostomidae, or leaf-nosed bats. Proceedings of the Zoological Society of London, 1866, 111 - 118.","Dobson, G. E. (1878) Catalogue of the Chiroptera in the collection of the British Museum. British Museum of Natural History, London, xxxvi + 567 pp. https: // doi. org / 10.5962 / bhl. title. 55341","Thomas, O. (1891) Note on Chiroderma villosum with the description of a new species for the genus. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 10, 881 - 883.","Peters, W. (1860) Eine neute Gattung von Flederthieren, Chiroderma villosum, aus Brasilien vor. Monatsberichte der Koniglichen Preussische Akademie des Wissenschaften zu Berlin, 1861,747 - 755.","Winge, H. (1892) Jordfundne og nulevende Flagermus (Chiroptera) fra Lagoa Santa, Minas Geraes, Brasilien. E Museo Lundii, Kjobenhavn, 2, 1 - 65.","Thomas, O. (1893). Further notes on the genus Chiroderma. Annals and Magazine of Natural History, Series 6, 11, 186 - 187. https: // doi. org / 10.1080 / 00222939308677490","Vieira, C. O. da C. (1942) Ensaio monografico sobre os quiropteros do Brasil. Arquivos de Zoologia, Museu de Zoologia da Universidade de Sao Paulo, 3, 219 - 471.","Cabrera, A. (1958) Catalogo de los mamiferos de America del Sur. Revista del Museo Argentino de Ciencias Naturales \" Bernardino Rivadavia \", Zoologia, 4, i-xvi + i-iv + 308.","Taddei, V. A. (1979) Phyllostomidae (Chiroptera) do Norte-ocidental do estado de Sao Paulo. III-Stenodermatinae. Ciencia e Cultura, 31 (8), 900 - 914.","Taddei, V. A. (1980) Aspectos da biologia de Chiroderma doriae, Thomas, 1891 (Chiroptera, Phyllostomidae). Sessoes da Academia Brasileira de Ciencias, 52, 643 - 644.","Nogueira, M. R. & Peracchi, A. L. (2001) The feeding specialization in Chiroderma doriae (Phyllostomidae, Stenodermatinae) with comments on its conservation implications. Chiroptera Neotropical, 8 (1 / 2), 143 - 156.","Nogueira, M. R. & Peracchi, A. L. (2003) Fig-seed predation by 2 species of Chiroderma: discovery of a new feeding strategy in bats. Journal of Mammalogy, 84 (1), 225 - 233. https: // doi. org / 10.1644 / 1545 - 1542 (2003) 084 2.0. CO; 2","Sipinski, E. A. B. & Reis, N. R. (1995) Dados ecologicos dos quiropteros da Reserva Volta Velha, Itapoa, Santa Catarina, Brasil. Revista Brasileira de Zoologia, 12 (3), 519 - 528.","Esberard, C. E. L., Chagas, A. S., Baptista, M., Luz, E. M. & Pereira, C. S. (1996) Observacoes sobre Chiroderma doriae Thomas, 1891 no municipio do Rio de Janeiro, RJ (Mammalia, Chiroptera). Revista Brasileira de Biologia, 56 (4), 651 - 654.","Taddei, V. A. (1973) Phyllostomidae da Regiao Norte-Ocidental do Estado de Sao Paulo. PhD dissertation. Universidade Estadual Paulista, Sao Jose do Rio Preto, 233 pp.","Novaes, R. L. M., Menezes Jr., L. F., Duarte, A. C. & Facanha, A. C. S. (2010) Consumo de Psychotria suturella Muell. Arg. (Rubiaceae) por morcegos no sudeste do Brasil. Chiroptera Neotropical, 16 (1), 535 - 538.","Laurindo, R. S., Gregorin, R. & Tavares, D. C. (2017) Effects of biotic and abiotic factors on the temporal dynamic of bat-fruit interactions. Acta Oecologica, 83, 38 - 47. https: // doi. org / 10.1016 / j. actao. 2017.06.009","Olmos, F. & Boulhosa, R. L. P. (2000) A meeting of opportunists: birds and other visitors to Mabea fistulifera (Euphorbiaceae) inflorescences. Ararajuba, 8 (2), 93 - 98.","Garbino, G. S. T. & Tavares, V. da C. (2018 b) Roosting ecology of Stenodermatinae bats (Phyllostomidae): Evolution of foliage roosting and correlated phenotypes. Mammal Review, 48 (2), 75 - 89. https: // doi. org / 10.1111 / mam. 12114","Arnone, I. S. (2008) Estudo da comunidade de morcegos na area carstica do Alto Ribeira - SP: uma comparacao com 1980. MSc thesis, Universidade de Sao Paulo, Sao Paulo, 116 pp.","Motta-Junior, J. C. & Taddei, V. A. (1992) Bats as prey of stygian owls in southeastern Brazil. Journal of Raptor Research, 26 (4), 259 - 260.","Gregorin, R., Lobao, K. W., Oliveira, L. F., Machado, F. S., Gil, B. B. & Tavares, V. da C. (2017) Vertical stratification in bat assemblages of the Atlantic Forest of south-eastern Brazil. Journal of Tropical Ecology, 33 (5), 299 - 308. https: // doi. org / 10.1017 / S 026646741700027 X","Presley, S. J. (2004) Ectoparasitic assemblages of Paraguayan bats: ecological and evolutionary perspectives. PhD dissertation, Texas Tech University, Lubbock, Texas, 326 pp.","Lourenco, E. C., Patricio, P. M. P. & Famadas, K. (2016) Community components of spinturnicid mites (Acari: Mesostigmata) parasitizing bats (Chiroptera) in the Tingua Biological Reserve of Atlantic Forest of Brazil. International Journal of Acarology, 42 (2), 63 - 69. https: // doi. org / 10.1080 / 01647954.2015.1117525","Lima Silva, C., Valim, M. P. & Graciolli, G. (2017) Acaros ectoparasitos de morcegos no estado de Mato Grosso do Sul, Brasil. Iheringia, Serie Zoologia, 107, 1 - 6. https: // doi. org / 10.1590 / 1678 - 4766 e 2017111","Soares, F. A. M., Rocha, P. A., Mikalauskas, J. S., Graciolli, G. & Ferrari, S. F. (2017) Ectoparasitic bat flies (Diptera, Streblidae) of bats (Chiroptera, Mammalia) from Mata do Junco Wildlife Refuge, Sergipe, northeastern Brazil. Oecologia Australis, 21 (4), 385 - 395. https: // doi. org / 10.4257 / oeco. 2017.2104.03","Lourenco, J. L. M., Minuzzi-Sauza, T. T. C., Silva, L. R., Olivei, A. C., Mendonca, V. J., Nitz, N., Aquiar, L. M. S. & Gurgel-Goncalves, R. (2018) High frequency of trypanosomatids in gallery forest bats of a Neotropical savanna. Acta Tropica, 177, 200 - 206. https: // doi. org / 10.1016 / j. actatropica. 2017.10.012","Taddei, V. A. (1976) The reproduction of some Phyllostomidae (Chiroptera) from the northwestern region of the State of Sao Paulo. Boletim de Zoologia, Universidade de Sao Paulo, 1 (1), 313 - 330. https: // doi. org / 10.11606 / issn. 2526 - 3358. bolzoo. 1976.121587","Silva, S. S. P., Dias, D., Martins, M. A., Guedes, P. G., Almeida, J. C., Cruz, A. P., Serra-Freire, N. M., Damascena, J. S. & Peracchi, A. L. (2015) Bats (Mammalia: Chiroptera) from the Caatinga scrublands of the Crateus region, northeastern Brazil, with new records for the state of Ceara. Mastozoologia Neotropical, 22 (2), 335 - 348.","Almeida, J. C., Martins, M. A., Guedes, P. G., Peracchi A. L. & Serra-Freire, N. M. (2016) New records of mites (Acari: Spinturnicidae) associated with bats (Mammalia, Chiroptera) in two Brazilian biomes: Pantanal and Caatinga. Revista Brasileira de Parasitologia Veterinaria, Jaboticabal, 25 (1), 18 - 23. https: // doi. org / 10.1590 / S 1984 - 29612016005"]}

Genus Chiroderma Peters, 1860 Synonyms: Chiroderma Peters, 1860: 747. Type-species Chiroderma villosum Peters, 1860, by monotypy. Mimetops Gray, 1866: 117. Listed in the synonymy of Chiroderma; being a nomen nudum. Chirodesma Thenius, 1989: 113 (not verified). Incorrect spelling of Chiroderma W. Peters, 1860. Distribution. Chiroderma is in western, eastern, and southern México, Central America (Guatemala, Belize, Honduras, El Salvador, Nicaragua, Costa Rica, and Panamá), Lesser Antilles (Guadeloupe, Marie-Galante, Monserrat, Saint Kitts and Nevis), Trinidad and Tobago, and South America (Venezuela, Guyana, Suriname, French Guiana, Colombia, Ecuador, Perú, Bolivia, Paraguay, and Brazil south to the state of Santa Catarina) (Fig. 7). Diagnosis. Chiroderma is a genus of small to large-sized fruit and seed-eating bats (total length of head and body 50–93 mm, length of forearm 34–58 mm, body mass 11–42 g; Tables 7 and 8). Dorsal fur dense, with long guard hairs, standing out above the underfur covering the body, and especially conspicuous on the cephalic region. Dorsal fur varies from buff to dark brown or dark gray; individual hairs with three well-defined bands, with the middle band always wider and paler than basal and terminal bands. Ventral pelage varies from pale gray to plumbous gray. The median dorsal stripe conspicuous or faint; may be absent in some C. villosum. Dorsal stripe begins at interscapular region and extends to the base of uropatagium. Four facial stripes present in most individuals, and may be bright and wide or faint and narrow. Tip of the spear of noseleaf may be notched. Horseshoe of noseleaf with free margins along its entire extension. Ears relatively small and round. Forearm densely furred along the proximal ⅔ of its length. Wing membranes, i.e., propatagium, chiropatagium, plagiopatagium, and uropatagium, are dark and opaque, except for the pale, translucent membrane between digits II and III of the dactylopatagium. Uropatagium relatively well-developed, extending posteriorly to the level of the knees; densely furred dorsally along ⅔ of its length. Plagiopataium inserts at the metatarsus. Tail absent. Calcar shorter than foot. Skull with a conspicuous notch at the region of the nasal bones, which are extremely reduced (Fig. 8). Orbital region relatively large; distinct postorbital processes. Frontonasal region relatively straight, in lateral view (neither concave nor convex; Fig. 8). Hard palate long, extending posteriorly close to glenoid fossa. Basioccipital pits shallow or absent. Dental formula: I 2/2, C 1/1, P 2/2, M 2/2. First upper incisors (I1) conic in cross-section, elongated with simple tips (not bilobed), and more than twice the crown size of the second upper incisors (I2). First upper premolar (P3) and canine (C) in contact; diastema between P3 and second upper premolar (P4). First and second upper molars (M1 and M2) with approximately the same occlusal area, or M2 with slightly larger area than M1; M2 triangular in occlusal view, with protocone distally placed and level with centrocrista. Hypocone absent in both M1 and M2; small hypoconal basin present in M1. With cranium and mandible in occlusion, there is a lateral gap bordered by the upper canine (C), first upper premolar (P3), and the two lower premolars (p2 and p4; Fig. 9). Coronoid process of mandible tall, its height approximately level with the tip of lower canine (c). Angular process conspicuous, projecting ventro-posteriorly in relation to the horizontal ramus of the mandible. Mandibular condyle relatively high, level with or slightly above tooth row. First lower premolar (p2) close to canine, the two teeth usually in contact; p2 shorter in height and length than second lower premolar (p4), ranging from approximately ¼ to ⅔ the height of p4. Diastema between p2 and p4. Second lower molar (m2) is the largest mandibular tooth and approximately twice the mesiodistal length of the first lower molar (m1). Well-developed metaconid, entoconid, protoconid, and hypoconid in m2. There is a fifth cusp between the hypoconid and entoconid, that we identify as the hypoconulid, following Garbino & Tavares (2018a). Discrete morphological comparisons among the species of Chiroderma recognized in this study are summarized in Table 9, and the descriptive statistics of the species is summarized in Tables 7 and 8. The species of Chiroderma for which the karyotype is known, i.e., C. doriae, C. improvisum, C. salvini, C. trinitatum, and C. villosum, have a chromosomal complement of 2n = 26 and FN = 48, a subtelocentric X-chromosome, and a submetacentric or subtelocentric Y-chromosome (Baker 1967, 1973; Baker & Hsu 1970; Baker & Genoways 1976; Varella-Garcia & Taddei 1985)., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 16-20, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Peters, W. (1860) Eine neute Gattung von Flederthieren, Chiroderma villosum, aus Brasilien vor. Monatsberichte der Koniglichen Preussische Akademie des Wissenschaften zu Berlin, 1861,747 - 755.","Gray, J. E. (1866) Revision of the genera of Phyllostomidae, or leaf-nosed bats. Proceedings of the Zoological Society of London, 1866, 111 - 118.","Thenius, E. (1989) Zahne und Gebiss der Saugetiere. In: Walter de Gruyter (Ed.), Handbuch der Zoologie, VIII / 56, pp. i-xi + 1 - 513.","Garbino, G. S. T. & Tavares, V. da C. (2018 a) A Quaternary record of the big-eyed bat Chiroderma villosum (Chiroptera: Phyllostomidae) with a revised lower molar terminology. Mammalia, 82 (4), 393 - 399. https: // doi. org / 10.1515 / mammalia- 2017 - 0037","Baker, R. J. (1967) Karyotypes of bats of the family Phyllostomidae and their taxonomic implications. The Southwestern Naturalist, 12 (4), 407 - 428. https: // doi. org / 10.2307 / 3669608","Baker, R. J. & Hsu, T. C. (1970) Further studies on the sex-chromosome systems of the American leaf-nosed bats (Chiroptera, Phyllostomatidae). Cytogenetics, 9 (2), 131 - 138. https: // doi. org / 10.1159 / 000130083","Baker, R. J. & Genoways, H. H. (1976) A new species of Chiroderma from Guadeloupe, West Indies (Chiroptera: Phyllostomatidae). Occasional Papers of the Museum of Texas Tech University, 48, 1 - 9.","Varella-Garcia, M. & Taddei, V. A. (1985) Analise cariotipica de Chiroderma doriae (Chiroptera, Phyllostomidae). Ciencia e Cultura, 37 (7), 790."]}

Chiroderma salvini Dobson, 1878 Synonyms: Chiroderma salvini Dobson, 1878: 532; type locality “ Costa Rica.” Chiroderma salvini salvini: Handley, 1966: 297; name combination. Chiroderma salvini scopaeum Reid and Langtimm, 1993: 300; not Chiroderma salvini scopaeum Handley, 1966. Type Material. The type of C. salvini (BMNH 68.8.16.2), fixed by original designation, is a fluid-preserved adult male with skull removed and tongue still attached to the body. The anterior half of the pelage is more faded than the posterior half. Nevertheless, it is possible to note whitish hairs above the lips and below the eyes. A thin median dorsal stripe extends from the middle dorsum to the posterior extremity of the lower back (contra Dobson 1878: 532, who considered the stripe to be absent). The cranium and mandible are in good condition with all teeth present. Some parts of the basicranium, as well as a large part of the palate have attached soft tissue. The I1s have convergent tips that do not touch each other. The angular processes of the mandible are broken. The species was named after Osbert Salvin, a British zoologist who edited and organized, with Frederick Godman, the 40-volume “Biologia Centrali-Americana”. On the type specimen’s skull label, the locality is given as “ Costa Rica ”, and to the right it is handwritten “O. Salvin [c]”, suggesting that the collector would indeed be Salvin. However, this naturalist collected specimens exhaustively in Guatemala and bordering countries, such as Belize (Godman 1915; Papavero 1973). Enrique Arcé, a Guatemalan field worker trained by Salvin, collected most of the bird specimens from Costa Rica described by the British zoologist (Salvin 1864; Warren & Harrison 1971; Beolens et al. 2014). Therefore, we suggest that the type of C. salvini probably was collected by E. Arcé during his work in Costa Rica. In the volume on mammals of the “Biologia Centrali-Americana”, authored by E. Alston (except for a few pages in the Supplement, authored by O. Thomas) and published between 1879 and 1882 (Lyal 2011), there is a color plate of C. salvini depicting the species as lacking the medial dorsal stripe (Fig. 10), an error that probably originated from Dobson (1878) description. Dobson (1880), however, recorded the presence of the stripe in an additional specimen of C. salvini from Popayán, Colombia. Distribution and Habitat. Chiroderma salvini is in eastern and southern México from Veracruz southeastward through Central America (Guatemala, El Salvador, Honduras, Costa Rica, and Panamá), into South America (northern and western Venezuela, western and northern Colombia, western Ecuador, eastern Perú in the Andean foothills, and western Bolivia; Fig. 11). The absence of records from Nicaragua may be a sampling artifact, possibly related to the fact that this country has lower mean elevations than the neighboring countries, and C. salvini is associated with montane forests. Records of C. salvini are from humid tropical forests, mainly sub-montane and montane. In Guatemala, El Salvador, and Honduras, the species also occurs in seasonally dry tropical forests. The distribution of C. salvini is associated with moderate to high elevations, with records in or close to the Sierra Madre Oriental in México, the cordilleras of Central America (e.g. Sierra Madre de Chiapas in Guatemala, Cordillera de Talamanca in Panamá and Costa Rica), and on both slopes of the Andean cordillera in South America (Fig. 11). Among the 39 specimen localities with precise coordinates, the mean elevation was 1,010 m above sea level (ranging from 73 m to 2,045 m), with 32 localities (82%) above 600 m and 20 (51%) above 1,000 m. In Panamá, C. salvini was more frequently captured between 600 and 1,500 m (Handley 1966b). In Parque Nacional Braulio Carrillo, Costa Rica, the species was recorded at 680 m (Timm et al. 1989). In Venezuela C. salvini was captured between 611 and 2,240 m, with 93% of the captures above 1,000 m (Handley 1976). In Parque Nacional de Manú, Peruvian Amazon, the species was documented between 450 and 1,920 m (Solari et al. 2006). In the Department of Tolima, Colombia, records of C. salvini are between 1,380 and 2,150 m (Bejarano-Bonilla et al. 2007; Galindo-Espinosa et al. 2010), and in the Department of Valle del Cauca, C. salvini was captured at elevations from 1,200 to 1,700 m (Mora-Beltrán & López-Arévalo 2018). Description and Comparisons. The dorsal pelage of C. salvini varies from pale brown to dark brown. Dorsal hairs are tricolored, with a narrow (approximately ¼ of hair length) dark brown base, a wide (approximately ½ of hair length) buff medial band, and a narrow terminal band approximating ¼ of hair length. Basal and terminal bands are usually the same color. Genal and interocular pairs of facial stripes are always present; conspicuous, wide, and brilliant-white. Interocular stripes are large, their widths varying between 1 and 4 mm, and entirely white. The median dorsal stripe is visible in most specimens, not detected in 2 of 174 specimens (1.1% of the sample): one from Venezuela (USNM 415233) had a faint suggestion of the stripe on the middle dorsum, and another from Honduras (TTU 12806) had no trace of a stripe. The spear of the noseleaf has a simple tip. The lateral margins of the horseshoe and the spear are whitish. The base and margins of the ear are yellowish. The dimensions of the cranium of C. salvini are similar to small C. doriae, large C. scopaeum, and large C. villosum (Tables 7 and 8). The braincase is globose, conspicuously standing out from the adjacent frontal and nasal regions. In dorsal view, the nasal notch extends posteriorly to the anterior margin of the orbits (Fig. 12). In lateral view, the anterior margin of the orbits is even with the distal margin of P4 and mesial margin of M1 (Fig. 13). A sagittal crest was present in 87.7% (186 of 212) of the specimens we examined. The sagittal crest was weakly developed in 22 (10.4%) specimens and not detected in four (1.8%). The posterior palatine process was absent in 77% of the sample (159 of 206 specimens), and was poorly developed when present. With cranium and mandible in occlusion, there is no frontal gap (as in C. improvisum and C. villosum; Fig. 14) but there is a small lateral gap, as in C. doriae, C. scopaeum, C. gorgasi and C. trinitatum (Fig. 9). The I1s converge medially in 97% of the specimens (200 of 206) and the tips may or may not be in contact. Six specimens (3%) have parallel I1 crowns that lack any contact. The P3 is oval in occlusal outline, differing from the antero-posteriorly compressed outline of P 3 in C. doriae. The P3 contacts C. but not P4. along = extends posterior to interorbital region; short = does not reach interorbital region or reaches only its anterior margin. bcaudal portion of hard palate. cgap delimited by I1, c, and i1,2, when skull and mandible are occluded. The lower canine is pointed and relatively tall, and the tip is approximately the same height as the coronoid process (as in C. villosum, but differing from C. doriae and C. scopaeum, in which the canines are clearly below the level of the coronoid; Fig. 13). The crown of p2 is low, approximately ¼ of the crown height of p4, longer mesiodistally than tall, and does not contact p4 (similar to the morphology of C. scopaeum and C. villosum). Compared with the allopatric C. doriae, C. salvini can be distinguished by its smaller size, globose braincase (less rounded in doriae), taller lower canines (lower canines in doriae are relatively shorter and below the level of the coronoid process), and smaller p2 (in doriae the p2 is approximately ⅔ of the height of p4). Where sympatric, C. salvini can be confused with C. scopaeum and C. villosum. Externally, C. salvini can be separated from C. villosum by, on average, a longer forearm (Tables 7 and 8); basal and apical bands of dorsal fur the same color (in villosum the base is usually darker than the tip); presence of wide and conspicuous facial stripes (narrow, faint, or absent in villosum), and a simple tip on the noseleaf spear with pale lateral margins (notched tip and noseleaf uniformly brown in villosum). Cranially, C. salvini differs from C. villosum by its relatively longer rostrum and shorter nasal notch (in villosum the notch is longer, ending near the level of the post-orbital constriction); smaller orbits (in villosum the anterior margin of the orbit is even with the P4); post-orbital processes less pointed than in villosum; posterior palatine process small or absent (in villosum process usually present and conspicuous), and absence of a frontal gap when cranium and lower jaw are in occlusion (Fig. 14). Compared with C. scopaeum, C. salvini is larger (Fig. 15, Tables 8 and 10) and externally it differs in pelage color, usually being darker than scopaeum. The skull of C. salvini is more robust, and the lambdoid-suture region of C. scopaeum is rounder, as can be seen in dorsal view (Fig. 12). The nasal notch of salvini is longer, usually reach- ing the interorbital region (Fig. 12). Lower canines are relatively taller and more pointed than in scopaeum, and the medial cingulum of the lower canines is not as well developed as in scopaeum (Fig. 13). Geographic Variation and Phylogeography. Phylogenetic analyses of 18 sequences did not recover any geographical structuring from Costa Rica to Bolivia (Fig. 16). Phenotypically, C. salvini is a relatively homogeneous species across its distribution. Subspecies. C. salvini is monotypic. Remarks. We found four published reports in which C. villosum from localities in Perú and Brazil were misidentified as C. salvini. The record of C. salvini for the Serra do Divisor, in the Peruvian Amazon (Medina et al. 2015), is here reidentified as C. villosum based on the reported forearm length (45.6 mm) and on a photograph of the specimen clearly showing a notched tip on the noseleaf and ears lacking pale margins (C. Medina in litt.). We also reanalyzed the specimens from Porto Velho, Rondônia, (MZUSP 35408) and Aricá, Mato Grosso, (MZUSP 6494) reported by Rocha et al. (2016), and confirmed that they have the diagnostic characters of C. villosum. Also, the record for the Cerrado of Tocantins (Maas et al. 2018) is recognized here as a C. villosum based on the measurements presented in the article and a photo of the skull (L.A.C. Gomes in litt.). Natural History. Four genera and five species of plants are documented in the diet of C. salvini in Colombia: Cecropia telealba (Urticaceae), Ficus insipida, F. cuatrecasana, Poulsenia armata (Moraceae), and Piper phytolaccifolium (Piperaceae) (Castaño et al. 2018). In Bolivia, one C. salvini was captured in a mist net set under a Ficus guaranitica (Aguirre 1994 apud Anderson 1997). In Veracruz, México, individuals were covered in pollen of Pachira aquatica (Malvaceae) (Hernández-Montero & Sosa 2016). In the Peruvian Amazon, Bravo et al. (2008, 2010) recorded C. salvini visting “collpas”, which are mineral licks containing clay-rich water that is ingested by the bats. Diurnal roosts used by C. salvini are unknown, but one animal was recorded flying through a lighted tunnel in a gold mine in Panamá and, in Venezuela specimens, have been captured inside houses (Goldman 1920; Handley 1976). The following ectoparasites have been recorded in C. salvini in Panamá: Amblyomma sp. n. (Ixodidae), Chirnyssoides caparti (Sarcoptidae), Periglischrus iheringi (Spinturnicidae), and Paratrichobius salvini (Streblidae) (Fairchild et al. 1966; Furman 1966; Wenzel et al. 1966; Lourenço et al. 2013). In Venezuela, Periglischrus iheringi and Trichobius persimilis (Streblidae) were collected in C. salvini (Herrin & Tipton 1975; Wenzel 1976), and in México the mites Parichoronyssus lopezi (Macronyssidae), Periglischrus iheringi, and Eudusbabekia vampyrops (Myiobiidae) were recorded on the species (Colín-Martínez et al. 2017). The reproductive pattern of C. salvini in Central America is best described as seasonal polyestry, with birth peaks occurring between March and April, and in August. Based on label information, pregnant females were recorded in Panamá in January (n=1), February (n=41), March (n=1), and June (n=1), while lactating individuals were recorded in March (n=5). In Guatemala, a pregnant C. salvini was recorded in January (Carter et al. 1966). In Honduras, pregnant or lactating C. salvini have been found in July and August (LaVal 1969). In South American populations, the scarcity of data does not permit generalizations. Based on the information we obtained from specimen labels, pregnant females have been recorded in Venezuela in July (n=2), August (n=1), and November (n=1), and in the Colombian Pacific there is a record for June (n=1). Based on literature records, pregnancies in Colombia are known for January, March, April, May, June, October, and December, and there are records of females that were both pregnant and lactating in March and April (Wilson 1979). In cis-Andean South America, two pregnant females were recorded from Perú, in August and September, and a lactating C. salvini was noted in October. Specimens Examined (N = 216): Bolivia: La Paz, Serrania Bellavista (AMNH 246625); Pando, Santa Rosa (AMNH 262537, 262538); Santa Cruz, 4.5 km N and 1.5 km E of Cerro Amboro (AMNH 261666); Santa Cruz, Estancia San Rafael de Amboro (AMNH 261667–261670). Colombia: Quindío, Vereda El Dorado (IAvH-M 7034), Vereda San Juan d’Carolina (IAvH-M 7036, 7039), Valle del Cauca, Pance (USNM 483743–483746), Río Zabaletas (USNM 483747–483762). Costa Rica: without precise locality (BMNH 68.8.16.2 [holotype of salvini]); Cartago, Angostura (USNM 12913/22849), Guanacaste, Rincón de La Vieja (USNM 565812); Heredia, Parque Nacional Braulio Carrillo (USNM 562856); Puntarenas, Cañas Gordas (AMNH 142484). El Salvador: Santa Ana, Los Planes (TTU 62461, 62462). Guatemala: El Progreso, Rio Uyús (ROM 99703). Honduras: Francisco Morazán, 16 km by road N Tegucigalpa (TTU 12800 –12808), La Flor (AMNH 126210, 126211, 126244–126251, 126253, 126255–126264, 126446, 126448–126455), San Marcos (AMNH 123331), Olancho, 50.4 km by road NNE Juticalpa (TTU 12809). México: Veracruz, Las Minas (USNM 329445). Panamá: Bocas del Toro, Río Changena Camp (USNM 319415–319425, 319499, 319500), Rancho Mojica, Río Changena (USNM 319286), Chiriquí, Cuesta de Piedra (USNM 331684–331686), Darién, Cana (USNM 179718), Cerro Malí (USNM 338042, 338043), Cerro Pirre (LSUMZ 25468–25474), Cerro Tacarcuna (USNM 338044), Jaqué (USNM 362919), Tacarcuna Village Camp (USNM 209969, 305387, 309443–309445, 309906, 309908–309910, 309912–309942, 309946–309968, 309972– 309977), Panamá, Cerro Azul (USNM 305388, 323445–323447). Perú: Cajamarca, San Ignacio (MUSM 12637), Cusco, Consuelo (MUSM 19663–19665, 19667), Comunidad Nativa Tangoshiari (MUSM 13377), Ridge Camp (USNM 588032), Madre de Dios, Hacienda Amazonia (MUSM 9742, 9751), Quebrada Aguas Calientes (MUSM 16650), Pasco, Palmira (MUSM 10878–10880), Puno, Yanacocha (MUSM 34980), Tumbes, Quebrada Naranjos (MUSM 19177). Venezuela: Carabobo, La Copa (USNM 440740–440744), La Vega del Río Santo Domingo (USNM 440746), Distrito Federal, Los Venados (USNM 370526, 370527), Hotel Humboldt (USNM 370528, 370530–370532), Miranda, Guatopo Natural Park (USNM 387191), Monagas, Hacienda San Fernando (USNM 415233–415235), San Agustín (USNM 415236, 415237)., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 20-28, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Dobson, G. E. (1878) Catalogue of the Chiroptera in the collection of the British Museum. British Museum of Natural History, London, xxxvi + 567 pp. https: // doi. org / 10.5962 / bhl. title. 55341","Reid, F. & Langtimm, C. (1993) Distributional and natural history notes for selected mammals from Costa Rica. The Southwestern Naturalist, 38 (3), 299 - 302. https: // doi. org / 10.2307 / 3671441","Godman, F. D. (1915). Biologia Centrali-Americana. Introductory Volume. Taylor and Francis, London, vii + 149 pp. https: // doi. org / 10.5962 / bhl. title. 730","Papavero, N. (1973) Essays on the history of Neotropical dipterology, with special reference to collectors (1750 - 1905). Vol. 2. Museu de Zoologia, Universidade de Sao Paulo, Sao Paulo, 230 pp.","Salvin, O. (1864) Descriptions of seventeen new species of birds from Costa Rica. Proceedings of the Zoological Society of London, 1864, 579 - 586.","Warren, R. L. M. & Harrison C. J. O. (1971) Type-specimens of birds in the British Museum (Natural History). Vol. 2. Passerines. Trustess of the British Museum, London, 628 pp. https: // doi. org / 10.5962 / bhl. title. 8584","Beolens, B., Watkins, M. & Grayson, M. (2014) The eponym dictionary of birds. Bloomsbury, London, 1414 pp.","Lyal, C. H. C. (2011) The dating of the Biologia Centrali-Americana. Zoological Bibliography, 1 (2), 67 - 102.","Dobson, G. E. (1880) On some new or rare species of Chiroptera in the colection of the Gottingen Museum. Proceedings of the Zoological Society of London, 48 (3), 461 - 465. https: // doi. org / 10.1111 / j. 1469 - 7998.1880. tb 06584. x","Handley Jr., C. O. (1966 b) Checklist of the mammals of Panama. In: Wenzel, R. L. & Tipton, V. J. (Eds.), Ectoparasites of Panama. Field Musem of Natural History, Chicago, Illinois, pp. 753 - 795.","Timm, R. M., Wilson, D. E., Clauson, B. L., LaVal, R. K. & Vaughan, C. S. (1989) Mammals of the La Selva-Braulio Carrillo Complex, Costa Rica. North American Fauna, 75, i-iii + 1 - 162.","Handley Jr., C. O. (1976) Mammals of the Smithsonian Venezuelan Project. Brigham Young University Science Bulletin, Biological Series, 20 (5), 1 - 89. https: // doi. org / 10.5962 / bhl. part. 5667","Solari, S., Pacheco, V., Luna, L., Velazco, P. M. & Patterson, B. D. (2006) Mammals of the Manu Biosphere Reserve. Fieldiana Zoology, New Series, 110, 13 - 22. https: // doi. org / 10.3158 / 0015 - 0754 (2006) 110 [13: MOTMBR] 2.0. CO; 2","Bejarano-Bonilla, D. A., Yate-Rivas, A. & Bernal-Bautista, M. H. (2007) Diversidad y distribucion de la fauna quiroptera en un transecto altitudinal en el Departamento del Tolima, Colombia. Caldasia, 29 (2), 297 - 308.","Galindo-Espinosa, E. Y., Gutierrez-Diaz, K. A. & Reinoso-Florez, G. (2010) Lista de los quiropteros del departamento del Tolima, Colombia. Biota Colombiana, 11 (1 / 2), 107 - 116.","Mora-Beltran, C. & Lopez-Arevalo, H. F. (2018) Interactions between bats and floral resources in a premontane forest, Valle del Cauca, Colombia. Therya, 9 (2), 129 - 136.","Medina, C. E., Lopez, E., Pino, K., Pari, A. & Zeballos, H. (2015) Biodiversidad de la zona reservada Sierra del Divisor (Peru): una vision desde los mamiferos pequenos Revista Peruana de Biologia, 22 (2), 199 - 212. https: // doi. org / 10.15381 / rpb. v 22 i 2.11354","Rocha, P. A., Brandao, M. V., Garbino, G. S. T., Cunha, I. N. & Aires, C. C. (2016) First record of Salvin's big-eyed bat Chiroderma salvini Dobson, 1878 for Brazil. Mammalia, 80 (5), 573 - 578. https: // doi. org / 10.1515 / mammalia- 2015 - 0077","Maas, A. C. S., Gomes, L. A. C., Martins, M. A., Dias, D., Pol, A., Chaves, F. G., Schute, M., Araujo, R. M. & Peracchi, A. L. (2018) Bats in a Cerrado landscape of Northern Brazil: Species occurrence, influence of environmental heterogeneity and seasonality, and eight new records for the State of Tocantins. Mammalia, 82 (5), 469 - 480. https: // doi. org / 10.1515 / mammalia- 2017 - 0023","Castano, H. J., Carranza, A. J. & Perez-torres, J. (2018) Diet and trophic structure in assemblages of montane frugivorous phyllostomid bats. Acta Oecologica, 91, 81 - 90. https: // doi. org / 10.1016 / j. actao. 2018.06.005","Anderson, S. (1997) Mammals of Bolivia, taxonomy and distribution. Bulletin of the American Museum of Natural History, 231, 1 - 652.","Hernandez-Montero, J. R. & Sosa, V. J. (2016) Reproductive biology of Pachira aquatica Aubl. (Malvaceae: Bombacoideae): A tropical tree pollinated by bats, sphingid moths and honey bees. Plant Species Biology, 31 (2), 125 - 134. https: // doi. org / 10.1111 / 1442 - 1984.12096","Bravo, A., Harms, K. E., Stevens, R. D. & Emmons, L. H. (2008) Collpas: Activity hotspots for frugivorous bats (Phyllostomidae) in the Peruvian Amazon. Biotropica, 40 (2), 203 - 210. https: // doi. org / 10.1111 / j. 1744 - 7429.2007.00362. x","Bravo, A., Harms, K. E. & Emmons, L. H. (2010) Puddles created by geophagous mammals are potential mineral sources for frugivorous bats (Stenodermatinae) in the Peruvian Amazon. Journal of Tropical Ecology, 26 (2), 173 - 184. https: // doi. org / 10.1017 / S 0266467409990472","Goldman, E. A. (1920) Mammals of Panama. Smithsonian Miscellaneous Collections, 69, 1 - 309. https: // doi. org / 10.5962 / bhl. title. 15684","Furman, D. P. (1966) The spinturnicid mites of Panama (Acarina: Spinturnicidae). In: Wenzel, R. L. & Tipton, V. J. (Eds.), Ectoparasites of Panama. Field Musem of Natural History, Chicago, Illinois, pp. 125 - 166.","Wenzel, R. L., Tipton, V. J. & Kiewlicz, A. (1966) The streblid batflies of Panama (Diptera: Streblidae). In: Wenzel, R. L. & Tipton, V. J. (Eds.), Ectoparasites of Panama. Field Musem of Natural History, Chicago, Illinois, pp. 405 - 676.","Lourenco, E. C., Pinheiro, M. da C., Faccini, J. L. H. & Famadas, K. M. (2013) New record, host and localities of bat mite of genus Chirnyssoides (Acari, Sarcoptiformes, Sarcoptidae). Revista Brasileira de Parasitologia Veterinaria, Jaboticabal, 22 (2), 260 - 264. https: // doi. org / 10.1590 / S 1984 - 29612013000200045","Herrin, C. S. & Tipton, V. J. (1975) Spinturnicid mites of Venezuela (Acarina: Spinturnicidae). Brigham Young University Science Bulletin, Biological Series, 20 (2), 1 - 72.","Wenzel, R. L. (1976) The streblid batflies of Venezuela (Diptera: Streblidae). Brigham Young University Science Bulletin, Biological Series, 20, 1 - 177.","Colin-Martinez, H., Morales-Malacara, J. B. & Garcia-Estrada, C. (2017) Epizoic fauna survey on phyllostomid bats (Chiroptera: Phyllostomidae) in a shaded coffee plantation of southeastern Chiapas, Mexico. Journal of Medical Entomology, 55 (1), 172 - 182. https: // doi. org / 10.1093 / jme / tjx 186.","Carter, D. C., Pine, R. H. & Davis, W. B. (1966) Notes on Middle American bats. The Southwestern Naturalist, 11, 488 - 499. https: // doi. org / 10.2307 / 3668862","LaVal, R. K. (1969) Records of bats from Honduras and El Salvador. Journal of Mammalogy, 50 (4), 819 - 822. https: // doi. org / 10.2307 / 1378267","Wilson, D. E. (1979) Reproductive patterns. In: Baker, J. R., Jones Jr., J. K. & Carter, D. C. (Eds.), Biology of the bats of the New World Family Phyllostomatidae. Part III. Museum of Texas Tech University Special Publications 16. Museum of Texas Tech University Press, Lubbock, Texas, pp. 317 - 378."]}

Chiroderma villosum Peters, 1860 Synonyms: See under subspecies. Type Material. The lectotype of Chiroderma villosum (ZMB 408), designated by Thomas (1891), is a stuffed skin. The skull is inside the skin and the skin around the lips is everted, exposing all of the teeth except M2 and m2. The free portion of the noseleaf spear is broken. The wing patagia are crumbly and several fragments have fallen off. The tips of the ears are also crumbly. The skin appears to preserve the original color, the interocular stripes are barely visible, but the genal pair is not visible. Dorsal pelage is pale brown and three bands can be distinguished: a dark brown base, and buff middle. The dorsal stripe appears to be lacking. The I1s are parallel to each other, not in contact, and have slightly diverging tips. The p2 is small and clearly separated from p4. The forearm measures 47.9 mm. According to Thomas (1891), the individual depicted in Peters’ (1906) plate, published posthumously, is probably the type. In the illustration, the skull is separate from the skin, suggesting that the skull could have been removed and later re-inserted in the skin (Fig. 31). The other possibility is that the skull Peters illustrated belonged to the partial skeleton referred to, but not examined by Carter & Dolan (1978:59). Distribution. See under subspecies. Description and Comparisons. Dorsal pelage may be pale brown, dark brown, reddish brown, or grayish brown. Most of the 328 specimens we examined have a pale brown dorsum (61%, n=201). The second most common color of the dorsal pelage is dark brown (38%, n=125). Dorsal hairs are tri-banded, the base is always dark brown, and the middle band is pale buff. Facial stripes were not detected in 191 (58%) of the 328 analyzed specimens. In 36.2% of the sample (n=119), the stripes were detected, but not conspicuous. The interocular pair was the only pair of facial stripes detected in 82.3% of the specimens, while both pairs were present in 17.7%. Only 13 specimens (3.9% of the sample) had conspicuous facial stripes, with the interocular pair more conspicuous than the genal. The dorsal stripe was not detected in 152 of 333 specimens (45.6%) scored for this character. The stripe was faint in 154 specimens (46.2%), and, in some the only evidence of a stripe was a short white mid-dorsal line. Only 22 specimens (6.6%) had a conspicuous dorsal stripe. Five specimens (1.5%) had a median dorsal stripe, but they were not scored as to conspicuousness. The tragus and base of the ears are yellow, but the remainder is uniformly brownish. The tip of the noseleaf is notched in 49 (74.2%) of the 66 specimens scored for this character. The noseleaf is nearly uniformly pale brown, with the central rib of the spear having a more pinkish tone and the lateral margin of the horseshoe slightly paler. The skull of C. villosum is similar in size to C. d. vizottoi and C. scopaeum, and measurements have some overlap with small C. d. doriae, C. salvini, and large C. trinitatum (Tables 7 and 8). The braincase is relatively deep and globose. A sagittal crest is present in 356 (85.5%) of the 416 C. villosum crania scored for this character, and varied from high and conspicuous to low and inconspicuous. The orbits are relatively large. The nasal notch is long, extending back into the interorbital region close to post-orbital processes. A posterior palatine process was present in 280 (83%) of the 336 skulls scored for this character, and varied from a long and conspicuous to a small nub. When cranium and mandible are in occlusion, a wide lateral gap, bordered by C, P3 and P4, and p2 and p4, is evident (Fig. 9). The occluded teeth also form a W-shaped frontal gap rimmed by lower canines, lower incisors, and upper inner incisors (Fig. 14). The I1s were parallel to each other (76.2%, n=314), medially convergent (22.3%, n=92) or divergent (1.4%, n=6) in the 512 crania scored for this character (Fig. 32). M2 usually has a cingulum around the protocone, which projects lingually. Lower canines are narrow and tall, with the tips level with the top of the coronoid process when the mandible is viewed laterally. The crown of p2 is approximately ¼ the height of p4. The p2 is in contact with the canine, but not with p4. Compared to the other species of Chiroderma, the protoconid of p4 is narrow. The subspecies C. v. jesupi and C. v. villosum differ in size, with the former averaging smaller in forearm length and in every cranial dimension we analyzed (Table 9). As previously mentioned, our decision to recognize two subspecies was mainly due to the presence of haplotypes exclusive to the trans-Andean portion of the distribution of C. villosum. The two smallest Chiroderma (C. gorgasi and C. trinitatum) differ from C. villosum by size, presence of conspicuous facial and dorsal stripes, bicolored noseleaf, shorter nasal notch, convergent I1s, and taller p2. The insular C. improvisum is easily distinguished from C. villosum by its larger size, relatively lower braincase, longer rostrum, more robust dentition, and p 2 in contact with p4. Among similar-sized species, C. villosum can be differentiated by the weaker facial and dorsal stripes (conspicuous in C. doriae, C. salvini, and C. scopaeum), paler dorsal pelage (usually darker in C. d. doriae and C. salvini), notched tip of noseleaf (simple tip in C. doriae, C. salvini, and C. scopaeum), deeper braincase (shallow in C. d. doriae), longer nasal notch (shorter in C. doriae, C. salvini, and C. scopaeum), larger orbits (smaller in C. doriae, C. salvini, and C. scopaeum), I1s parallel or divergent (convergent in C. doriae, C. salvini, and C. scopaeum), smaller p2 (large in C. doriae), narrower and taller canines (wider and shorter in C. doriae and C. scopaeum), and presence of a frontal gap when cranium and mandible are in occlusion (gap absent in C. doriae, C. salvini, and C. scopaeum). Geographic Variation and Phylogeography. Phylogenetic analyses of sequences of COI representing 133 C. villosum recovered a clade composed of sequences exclusively from México, Central America, and trans-Andean South America (Figs. 33, 35). The samples from cis-Andean South America, however, form a polytomy, which does not suggest geographic structuring because specimens from distant regions, such as Bolivia and Trinidad, group together. There is geographic variation in size among the subpopulations of C. villosum. Specimens from the Atlantic rainforest of eastern Brazil (n=48) are larger than the other geographic groups (Table 13, Fig. 34). Specimens from Trinidad and Tobago also had larger dimensions; however, the sample size was smaller (n=7). Populations from the Amazon north of the Amazonas River and from México and Central America and trans-Andean South America had the smaller dimensions. We suggest there is clinal variation in size, with larger specimens in the extreme east and southeast part of the distribution and smaller individuals in the northwestern part of the range. There is overlap between measurements from adjacent geographic groups, but the populations from the western portion of the distribution, which would correspond to subspecies C. v. jesupi, are significantly smaller than the others (Fig. 34). Subspecies. We recognize two subspecies in Chiroderma villosum., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 48-51, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Peters, W. (1860) Eine neute Gattung von Flederthieren, Chiroderma villosum, aus Brasilien vor. Monatsberichte der Koniglichen Preussische Akademie des Wissenschaften zu Berlin, 1861,747 - 755.","Thomas, O. (1891) Note on Chiroderma villosum with the description of a new species for the genus. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 10, 881 - 883.","Carter, D. C. & Dolan, P. G. (1978) Catalogue of type specimens of neotropical bats in selected european museums. Museum of Texas Tech University Special Publications 15. Museum of Texas Tech University Press, Lubbock, Texas, pp. 1 - 136."]}

Chiroderma trinitatum Goodwin, 1958 Synonyms: Chiroderma trinitatus Goodwin, 1958:1; type locality “ Cumaca, Trinidad, British West Indies. ” Chiroderma trinitatum: Handley, 1960:466; correct gender concordance. Chiroderma trinitatum trinitatum: Barriga-Bonilla, 1965: 247; name combination. Chiroderma trinitratum Linares and Moreno-Mosquera, 2010: 275; incorrect subsequent spelling of Chiroderma trinitatum Goodwin, 1958. Type Material. The holotype, designated in the original publication, is specimen AMNH 175325, a female pregnant when collected in a well-lit cave, by L. Venus and B. Smith on March 22, 1956, in Cumaca, Trinidad and Tobago. The skin is preserved in fluid and nearly all hair has fallen out. There is a transversal cut on the abdomen. The skull has been removed and is in good condition, with all the teeth and cranial bones preserved. The fetus, removed from the type, has a distinct median dorsal stripe and a crown-rump length of approximately 20 mm. Distribution and Habitat. Specimens are known form Guyana; Suriname; French Guiana; Trinidad; northern, central and western Brazil; northern and eastern Bolivia; eastern Perú; eastern Ecuador; southern and eastern Colombia; and southern Venezuela (Fig. 23). Records are from humid tropical forests, in the Amazon basin, Orinoco basin, and the Guianas. Some records are from ecotonal areas between humid forests and drier, more seasonal formations, such as in Serra do Roncador (central Brazil), and in San Ramón (northeastern Bolivia). C. trinitatum have been collected from approximately 20 m above sea level (e.g. Belém, Brazil) to elevations near 1,050 m, in the Andean foothills (e.g. Santa Bibiana, Perú), corroborating the altitudinal amplitude given in the literature (e.g. Handley 1976; Ascorra et al. 1996; Solari et al. 2006, 2019). Description and Comparisons. Dorsal pelage may be pale brown, dark brown, or grayish brown (Fig. 24). A completely white C. trinitatum was recorded in the Peruvian Amazon by Tello et al. (2014). Dorsal hairs are tricolored, the base about ¼ of the hair length and dark brown, middle portion about ½ of the hair length and pale buff or pale gray, and tips about ¼ of the hair length and pale brown, dark brown, or grayish brown. Facial stripes are always present with the interocular pair wider (> 2 mm) than the genal stripes. A median dorsal stripe was present in 111 of the 113 specimens examined (98%). When present, the stripe was barely visible in 13 of 95 specimens (13%). The stripe usually begins in the interscapular region and reaches the posterior rump of the body. In some specimens, the dorsal stripe began more anteriorly in the region immediately behind the nape. The ears have yellowish margins and base, with the remainder brownish. The spear of the noseleaf has a simple tip, is brown in color, with paler lateral borders of the horseshoe. The skull is similar to that of C. gorgasi and it is smaller than every other species of Chiroderma (Tables 7 and 8). The braincase is globose, standing above the frontonasal region in lateral view. A sagittal crest was absent in 37 of 126 specimens (29.4%), weakly developed in 86 (68.2%), and well developed in 3 (2.3%). The nasal notch is relatively short, not reaching the interorbital region (Fig. 25). C. trinitatum has relatively small orbits; the anterior border approximating the mesial margin of M1 (Fig. 26). Post orbital processes are present, but rhomboid instead of pointed as in the other species. A posterior palatine process was absent in 110 (90.9%) of the 121 specimens examined. A small process was present in 10 specimens, and one (AMNH 264076) had an anomalous notch in place of a projection on the margin of the palate. Paraoccipital processes are absent. When cranium and mandible are in occlusion, there is a lateral gap bordered by the C, P3, P4, p2 and p4 (Fig. 9). The I1s have converging tips in most of the specimens (112 out of 122) we examined. The I1s may be in contact at the base, along approximately ⅔ of their length, or only at their tips, the most frequent state. In some C. trinitatum, the I1s are separated throughout their length. The P3 is wider (buccolingually) than long (mesiodistally) and does not touch P4. The crown of the lower canine is relatively short, clearly below the level of the tip of the coronoid, in lateral view (Fig. 26). The p2 is large, approximately ¾ of the height of p4, and higher than long. The p2 may be close to or in contact with the canine, or it may lie approximately equidistant between c and p4 (Fig. 26). Chiroderma trinitatum is easily differentiated from C. doriae and C. improvisum by its much smaller size (Tables 7 and 8). The intermediate-sized species, e.g. C. salvini, C. scopaeum, and C. villosum, are distinguishable from C. trinitatum by having their longer nasal notch that reaches the interorbital region, pointed post-orbital processes and the short p2, which is approximately ¼ the crown height of p4. The species most similar to C. trinitatum is C. gorgasi. C. trinitatum has relatively shorter lower canines, the tips of which are clearly below the level of the top of the coronoid process in lateral view. The lower canines have higher crowns in C. gorgasi, being on the same level or higher than the tip of the coronoid process (Fig. 26). The base of lower canines of C. trinitatum is also longer, but shorter in C. gorgasi. The p2 of C. trinitatum is usually higher than long mesiodistally; whereas, in C. gorgasi the p2 is longer than high (Fig. 27). As recently described by Lim et al. (2020), C. trinitatum tends to have a wider braincase and typically has a third cuspid on p4 that is absent in C. gorgasi. Geographic Variation and Phylogeography. Phylogenetic analyses of 68 sequences of C. trinitatum resulted in two highly-supported clades (Fig. 21). One of the clades is based on sequences from 26 specimens from the Guianas (French Guiana, Guyana, and Suriname), and the island of Trinidad. The other clade consists of 42 specimens, from the Guianas (Guyana and Suriname) and the Amazon basin (Bolivia, Brazil, Ecuador, and Perú) (Fig. 21). However, we could not find any morphological differences between the two clades. Subspecies. C. trinitatum is monotypic. Natural History. C. trinitatum is frugivorous and is known to consume fruits and infructescences of at least five species: Cecropia obtusifolia, Piper elongatum, Solanum riparium (Solanaceae), Ficus sp., and Vismia sp. (Hypericaceae) (Reis & Peracchi 1987; Loayza et al. 2006; Linares & Moreno-Mosquera 2010). C. trinitatum has also been recorded drinking the mineral-rich water at clay licks (“collpas”) in the Peruvian Amazon (Bravo et al. 2008, 2010; Ghanem et al. 2013; Ghanem & Voigt 2014). In most studies comparing the relative abundance of species in the canopy and understory, C. trinitatum was more commonly netted in the higher forest strata, at approximately 20 m, suggesting that the species is a canopy frugivore (Ascorra et al. 1996; Simmons & Voss 1998; Charles-Dominique & Cockle 2001; Kalko & Handley Jr. 2001; Delaval et al. 2005; Rex et al. 2011). Day roosts of C. trinitatum are unknown, and the only information available comes from the type specimen, which was collected on the island of Trinidad in a cave described by Goodwin & Greenhall (1961) as well-lit and co-inhabited by Micronycteris megalotis. In Venezuela, two species of ectoparasites were documented on C. trinitatum, the mite Periglischurus iheringi (Spinturnicidae) and a species of Streblidae of the genus Paratrichobius (salvini complex) (Herrin & Tipton 1975; Wenzel 1976). In Colombia, the tick Ornithodoros azteci (Argasidae) was recorded from C. trinitatum by Marinkelle & Grose (1981). The endoparasite Hasstilesia tricolor (Nematoda) was recorded by Nogueira et al. (2014) in C. trinitatum from Acre, Brazil. Reproductive data suggest seasonal polyestry. Pregnant females are recorded from December through March in the Amazon of Colombia, Perú, and Venezuela, and on the island of Trinidad. Pregnancies are recorded from June through September in Guyana and the Amazon of Brazil, Perú, and Venezuela. Lactating females were documented in February, April, May, and July. Apparently, gestation peaks during the rainy season (December to March), with one birth peak from the middle to the end of the rainy season, and a second during the dry season (June to September) with births occurring close to the beginning of the rainy season. Specimens Examined (N = 146): Bolivia: Beni, Estancia Yutiole (AMNH 210810), Remansos (AMNH 209520, 209521); La Paz, Río Coraico Valley (AMNH 246646), Santa Ana de Madidi (AMNH 261632, 261641); Santa Cruz, 10 km N San Ramón (AMNH 261674), Parque Nacional Noel Kempff Mercado, 23 km S Campamento Los Fierros (AMNH 264077), Parque Nacional Noel Kempff Mercado, 27.5 km S Campamento Los Fierros (AMNH 264076), Parque Nacional Noel Kempff Mercado, 3 km S Campamento Los Fierros (AMNH 264075), Parque Nacional Noel Kempff Mercado, El Refugio (USNM 584492). Brazil: Acre, Parque Nacional da Serra do Divisor (ALP 7020, 7088, 7099, 7124, 7134, 7143, 7144, 7195, 7295, 7311); Amazonas, Comunidade Cachoeirinha (LMSUP [ICA033]), opposite to Comunidade São Pedro (LMUSP [ICA173]); Mato Grosso, 264 km N Xavantina (USNM 393704–393711), Parque Nacional do Juruena, Serra dos Apiacás (CMUFLA 1284), São José do Rio Claro (MZUSP [PEV 896–897]); Pará, Fazenda Bocaina (UFMG [VCT1437]), Fazenda Fartura (MZUSP 36012, 36013), Floresta Nacional de Carajás (UFMG [VCT6342]), rio Xingu, Linha de Transmissão Jurupari (MZUSP 35033), Sta. A, IAN (USNM 361723, 460127), Várzea, Belém (USNM 460124, 460125, 460126); Rondônia, Monte Negro (MZUSP 35026, ZUFMS 1342). Colombia: Amazonas, Puerto Nariño (USNM 483766–483769); Vichada, Territorio Faunistico Tuparro (IAvH-M 2083). Guyana: Cuyuni-Mazaruni, Maz 3 (BMNH 1980.751, 1980.752). Namai Creek (ROM 108144); Potaro-Siparuni, Iwokrama Reserve (ROM 109026); Upper Demerara-Berbice, 3.2 km W Kurupukari (BMNH 1997.39), Tropenbos (ROM 103486). French Guiana: Sinnamary, Paracou (AMNH 266255, 266256, 267189, 268532, 269118). Perú: Amazonas, Cordillera del Condor (USNM 581934); Cusco, Camisea, Armihuari (MUSM 13619, 13622, 13623, USNM 582837), Camisea, Pagoreni (MUSM 13624–13626, USNM 582838), Consuelo (MUSM 19670); Huánuco, Puerto Inca (MUSM 36692); Junín, Santa Bibiana (MUSM 40606); Loreto, Jenaro Herrera (MUSM 4219, 5594), km 22,7 da rodovia Iquitos-Nauta (MUSM 29559), Peña Negra (MUSM 29557), Quebrada Grande (MUSM 21134, 21135), Río Lagartococha (MUSM 21138); Madre de Dios, Albergue Maskoitania (MUSM 19669), Pakitza (MUSM 678), Quebrada Aguas Calientes (MUSM 16651), Refugio Juliaca (MUSM 11662); Pasco, Campamento Río Lobo (MUSM 10229), Cerro Chontiya (MUSM 10226, 10227), Cerro Jonatán (MUSM 10228), San Pablo (AMNH 230656), Yanahuanca (MUSM 10881); Puno, Curva Alegre (MUSM 26658), San Fermín (MUSM 26659); Ucayali, Concesión de Conservación Río La Novia (MUSM 44185); Suriname: Brokopondo, Brownsberg Nature Park (ROM 114213); Sipaliwini, Kushere Landing (ROM 120168). Trinidad and Tobago: Trinidad, Arima (AMNH 205373), Cumaca (AMNH 175325 [holotype of trinitatum]), Fillete (AMNH 205375), Las Cuevas (BMNH 1971.121). Venezuela: Amazonas, Boca Mavaca (USNM 405159), Capibara (USNM 415245), Cerro Neblina base camp (USNM 560764–560766), Guayabal (USNM 415256, 415257), Río Cunucunuma (MZUSP 27166, 27167, USNM 405132–405139, 405141–405148, 405156, 405157), Río Manapiare (USNM 415250–415254); Apure, La Blanquita (USNM 440348); Barinas, Altamira (USNM 418121); Bolívar, 85 km SSE El Dorado (USNM 387192), El Manaco (USNM 387193–387198, 387202, 387203); Yaracuy, Minas de Aroa (USNM 440747)., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 40-44, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Goodwin, G. G. (1958) Three new bats from Trinidad. American Museum Novitates, 1877, 1 - 6.","Handley Jr., C. O. (1960) Descriptions of new bats from Panama. Proceedings of the United States National Museum, 112, 459 - 479. https: // doi. org / 10.5479 / si. 00963801.112 - 3442.459","Barriga-Bonilla, E. (1965) Estudios mastozoologicos colombianos, I: Chiroptera. Caldasia, 9 (43), 241 - 268.","Linares, E. L. & Moreno-Mosquera, E. A. (2010) Morfologia de los frutiolos de Cecropia (Cecropiaceae) del Pacifico colombiano y su valor taxonomico en el estudio de dietas de murcielagos. Caldasia, 32 (2), 275 - 287.","Handley Jr., C. O. (1976) Mammals of the Smithsonian Venezuelan Project. Brigham Young University Science Bulletin, Biological Series, 20 (5), 1 - 89. https: // doi. org / 10.5962 / bhl. part. 5667","Ascorra, C. F., Solari, S. & Wilson, D. E. (1996) Diversidad y Ecologia de los Quiropteros en Pakitza. In: Wilson, D. E. & Sandoval, A. (Eds.), Manu. The biodiversity of Southeastern Peru. Editorial Horizonte, Lima, pp. 593 - 612.","Solari, S., Pacheco, V., Luna, L., Velazco, P. M. & Patterson, B. D. (2006) Mammals of the Manu Biosphere Reserve. Fieldiana Zoology, New Series, 110, 13 - 22. https: // doi. org / 10.3158 / 0015 - 0754 (2006) 110 [13: MOTMBR] 2.0. CO; 2","Solari, S., Medellin, R., Rodriguez-Herrera, B., Tavares, V. da C., Garbino, G. S. T., Camacho, M. A., Tirira, D., Lim, B. K., Arroyo-Cabrales, J., Rodriguez-Duran, A., Dumont, E., Burneo, S., Aguirre, L. F., Tschapka, M. & Espinosa, D. (2019) Family Phyllostomidae (New World leaf-nosed bats). In: Wilson, D. E. & Mittermeier, R. A. (Eds.), Handbook of the Mammals of the World, Bats. Vol. 9. Lynx Edicions, Barcelona, pp. 444 - 583.","Tello, C., Streicker, D. G., Gomez, J. & Velazco, P. M. (2014) New records of pigmentation disorders in molossid and phyllostomid (Chiroptera) bats from Peru. Mammalia, 78 (2), 191 - 197. https: // doi. org / 10.1515 / mammalia- 2013 - 0019","Lim, B. K., Loureiro, L. O. & Garbino, G. S. T. (2020) Cryptic diversity and range extension in the big-eyed bat genus Chiroderma (Chiroptera: Phyllostomidae). ZooKeys, 918, 41 - 63. https: // doi. org / 10.3897 / zookeys. 918.48786","Reis, N. R. & Peracchi, A. L. (1987) Quiropteros da regiao de Manaus, Amazonas, Brasil (Mammalia, Chiroptera). Boletim do Museu Paraense Emilio Goeldi, Serie Zoologia, 3 (2), 1 - 21.","Loayza, A. P., Rios R. S. & Larrea-Alcazar, D. M. (2006). Disponibilidad de recurso y dieta de murcielagos frugivoros en la Estacion Biologica Tunquini, Bolivia. Ecologia en Bolivia, 41 (1), 7 - 23.","Bravo, A., Harms, K. E., Stevens, R. D. & Emmons, L. H. (2008) Collpas: Activity hotspots for frugivorous bats (Phyllostomidae) in the Peruvian Amazon. Biotropica, 40 (2), 203 - 210. https: // doi. org / 10.1111 / j. 1744 - 7429.2007.00362. x","Bravo, A., Harms, K. E. & Emmons, L. H. (2010) Puddles created by geophagous mammals are potential mineral sources for frugivorous bats (Stenodermatinae) in the Peruvian Amazon. Journal of Tropical Ecology, 26 (2), 173 - 184. https: // doi. org / 10.1017 / S 0266467409990472","Ghanem, S. J., Ruppert, H., Kunz, T. H. & Voigt, C. C. (2013) Frugivorous bats drink nutrient-and clay-enriched water in the Amazon rain forest: Support for a dual function of mineral-lick visits. J ournal of Tropical Ecology, 29 (1), 1 - 10. https: // doi. org / 10.1017 / S 0266467412000740","Ghanem, S. J. & Voigt, C. C. (2014) Defaunation of tropical forests reduces habitat quality for seed-dispersing bats in Western Amazonia: An unexpected connection via mineral licks. Animal Conservation, 17 (1), 44 - 51. https: // doi. org / 10.1111 / acv. 12055","Simmons, N. B. & Voss, R. S. (1998) The Mammals of Paracou, French Guiana: a neotropical lowland rainforest fauna Part 1. Bats. Bulletin of the American Museum of Natural Histor y, 237, 1 - 219.","Charles-Dominique, P. & Cockle, A. (2001) Frugivory and seed dispersal by bats. In: Bongers, F., Charles-Dominique, P., Forget, P. - M. & Thery, M. (Eds.), Nouragues: dynamics and plant-animal interactions in a Neotropical rainforest. Springer, Dordrecht, pp. 207 - 215.","Delaval, M., Henry, M. & Charles-Dominique, P. (2005) Interspecific competition and niche partitioning: example of a Neo- tropical rainforest bat community. Revue d'Ecologie, Terre Vie, 60, 149 - 166.","Rex, K., Michener, R., Kunz, T. H. & Voigt, C. C. (2011) Vertical stratification of Neotropical leaf-nosed bats (Chiroptera: Phyllostomidae) revealed by stable carbon isotopes. Journal of Tropical Ecology, 27 (3), 211 - 222. https: // doi. org / 10.1017 / S 0266467411000022","Goodwin, G. G. & Greenhall, A. M. (1961) A review of the bats of Trinidad and Tobago. Bulletin of the American Museum of Natural History, 122, 187 - 301.","Herrin, C. S. & Tipton, V. J. (1975) Spinturnicid mites of Venezuela (Acarina: Spinturnicidae). Brigham Young University Science Bulletin, Biological Series, 20 (2), 1 - 72.","Wenzel, R. L. (1976) The streblid batflies of Venezuela (Diptera: Streblidae). Brigham Young University Science Bulletin, Biological Series, 20, 1 - 177.","Marinkelle, C. J. & Grose, E. S. (1981) A list of ectoparasites of Colombian bats. Revista de Biologia Tropical, 29 (1), 11 - 20."]}

Key to the species and subspecies of genus Chiroderma 1 Size small, forearm equal to or less than 42 mm, greatest length of skull equal to or less than 23 mm................... 2 - Size intermediate to large, forearm equal to or more than 43 mm, greatest length of skull equal to or more than 23 mm......................................................................................................... 3 2 First lower premolar longer than high mesiodistally; third cuspid of first lower premolar absent; crown of lower canine at the same level or higher than the tip of the coronoid process of the mandibular bone.................... Chiroderma gorgasi - First lower premolar higher than long mesiodistally; third cuspid of first lower premolar present; crown of lower canine clearly below the level of the top of the coronoid process of the mandibular bone........................ Chiroderma trinitatum 3 Size large, length of forearm equal to or more than 56 mm, greatest length of skull equal to or more than 28 mm; first lower premolar in contact with the lower canine and second premolar............................... Chiroderma improvisum - Size intermediate to large, forearm length equal to or less than 54 mm (43–54 mm), greatest length of skull usually equal to or less than 28 mm (23–28 mm); diastema between first lower premolar and second lower premolar...................... 4 4 Size large, forearm length 47–54 mm, greatest length of skull 26–29 mm; dorsal pelage brown or dark brown; basal and terminal bands of the dorsal hairs of a same color................................................................ 5 - Size intermediate, forearm length 43–50 mm, greatest length of skull 23–26 mm; dorsal pelage brown, light brown or buff; base of dorsal hairs darker than terminal band................................................................... 6 5 First lower premolar large, with approximately ⅔ the height of second lower premolar; crown of lower canine clearly below the level of the top of the coronoid process of the mandibular bone; paraoccipital processes present....................................................................................................... Chiroderma doriae doriae - First lower premolar small, with approximately ¼ the height of second lower premolar; crown of lower canine approximately the same height as the top of the coronoid process of the mandibular bone; paraoccipital processes absent..................................................................................................... Chiroderma salvini 6 Tip of noseleaf unnotched; facial stripes bright and conspicuous; posterior process of the palate absent; first upper incisors with convergent tips....................................................................................... 7 - Tip of noseleaf notched; facial stripes dark and inconspicuous; posterior process of the palate present; first upper incisors with parallel or divergent tips................................................................................ 8 7 Length of forearm usually more than 48 mm (45–50 mm); greatest length of skull equal to or more than 25 mm (25–27 mm); high-crowned first lower premolar (⅔ the height of second lower premolar; low-crowned lower canine, with the tips clearly below the level of the top of the coronoid process of the mandibular bone.................... Chiroderma doriae vizottoi - Length of forearm equal to or less than 47 mm (43–47 mm); greatest length of skull equal to or less than 25 mm (23–25 mm); low-crowned first lower premolar (¼ the height of second lower premolar); high-crowned lower canine, with the tips approximately the same height as the top of the coronoid process of the mandibular bone................ Chiroderma scopaeum 8 Length of forearm 46 mm on average (41–52 mm); distributed east of the Andes........... Chiroderma villosum villosum - Length of forearm 44 mm on average (41–47 mm); distributed west of the Andes............. Chiroderma villosum jesupi, Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on page 58, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497

DNA barcoding is helping in discovering high levels of cryptic species and an underestimation of biodiversity in many groups of organisms. Although mammals are arguably the most studied and one of the least speciose taxonomic classes, the rate of species discovery is increasing and biased for small mammals on islands. An earlier study found bats in the Caribbean as a taxonomic and geographic deficiency in the International Barcode of Life initiative to establish a genetic reference database to enable specimen identification to species. Recent surveys in Dominican Republic, Jamaica, and Martinique have documented and barcoded half of the 58 bat species known from the Caribbean. I analyze all available barcode data of Caribbean bats to investigate biogeography and cryptic species in the Neotropical region. Analysis of the mitochondrial DNA gene cytochrome c oxidase subunit 1 results in a phylogenetic tree with all but one species as well-supported and reciprocally monophyletic. With a broader sampling across the Neotropics, there are also divergent lineages that exhibit biogeographic structuring: (i) a phylogenetic split between northern and southern Dominican Republic in three species, (ii) two taxa with cryptic species associated with higher degree of island endemism, (iii) populations of two widely distributed species with deep divergence between the Caribbean and North and Central America, and (iv) one species in the Caribbean with affinities to taxa in South America.

Incluye contenido parcial de los autores Biological invasion is one of the main threats to native biodiversity. For a speciesto become invasive, it must be voluntarily or involuntarily introduced by humans into a nonna-tive habitat. Mammals were among first taxa to be introduced worldwide for game, meat, and labor, yet the number of species introduced in the Neotropics remains unknown. In this dataset, we make available occurrence and abundance data on mammal species that (1) transposeda geographical barrier and (2) were voluntarily or involuntarily introduced by humans into theNeotropics. Our data set is composed of 73,738 historical and current georeferenced recordson alien mammal species of which around 96% correspond to occurrence data on 77 speciesbelonging to eight orders and 26 families. Data cover 26 continental countries in the Neotrop-ics, ranging from Mexico and its frontier regions (southern Florida and coastal-central Floridain the southeast United States) to Argentina, Paraguay, Chile, and Uruguay, and the 13 coun-tries of Caribbean islands. Our data set also includes neotropical species (e.g.,Callithrixsp.,Myocastor coypus,Nasua nasua)considered alien in particular areas of Neotropics. The mostnumerous species in terms of records are fromBossp. (n=37,782),Sus scrofa(n=6,730), andCanis familiaris(n=10,084); 17 species were represented by only one record (e.g., Syncerus caf-fer, Cervus timorensis, Cervus unicolor, Canis latrans). Primates have the highest number of spe-cies in the data set (n=20 species), partly because of uncertainties regarding taxonomicidentification of the generaCallithrix,which includes the speciesCallithrix aurita, Callithrixflaviceps, Callithrix geoffroyi, Callithrix jacchus, Callithrix kuhlii, Callithrix penicillata, andtheir hybrids. This unique data set will be a valuable source of information on invasion riskassessments, biodiversity redistribution and conservation-related research. There are no copy-right restrictions. Please cite this data paper when using the data in publications. We alsorequest that researchers and teachers inform us on how they are using the data

Research on mammals in the Guianas of northern South America has had a checkered history. In this review, I summarize the notable contributions to mammalogical study in Guyana, Suriname and French Guiana. These studies began in the mid-18th century with the binomial nomenclature system of scientific classification created by the Swedish naturalist Carl Linnaeus, who described 23 species new to science based on holotype specimens from the Guianas. Notwithstanding popular accounts by amateur naturalists visiting this region, over the next 7 decades there was only sporadic taxonomic work done on Guianan mammals primarily by researchers at European museums. The first comprehensive biological exploration took place in the 1840s during a geographic survey of the boundaries of British Guiana. However, it was not until almost half a century later that scientific publications began to regularly document the increasing species diversity in the region, including the prodigious work of Oldfield Thomas at the British Museum of Natural History in London. Another lull in the study of mammals occurred in the mid-1910s to the early 1960s after which foreign researchers began to rediscover the Guianas and their pristine habitats. This biological renaissance is still ongoing and I give a prospectus on the direction of future research in one of the last frontiers of tropical rainforest. An initiative that would be greatly beneficial is the establishment of a university network in the Guianas with graduate-based research to develop a cadre of professional experts on biodiversity and evolution as seen in other countries of South America.

As a periodic assessment of the mammal collection resource, the Systematic Collections Committee (SCC) of the American Society of Mammalogists undertakes decadal surveys of the collections held in the Western Hemisphere. The SCC surveyed 429 collections and compiled a directory of 395 active collections containing 5,275,155 catalogued specimens. Over the past decade, 43 collections have been lost or transferred and 38 new or unsurveyed collections were added. Growth in number of total specimens, expansion of genomic resource collections, and substantial gains in digitization and web accessibility were documented, as well as slight shifts in proportional representation of taxonomic groups owing to increasingly balanced geographic representation of collections relative to previous surveys. While we find the overall health of Western Hemisphere collections to be adequate in some areas, gaps in spatial and temporal coverage and clear threats to long-term growth and vitality of these resources have also been identified. Major expansion of the collective mammal collection resource along with a recommitment to appropriate levels of funding will be required to meet the challenges ahead for mammalogists and other users, and to ensure samples are broad and varied enough that unanticipated future needs can be powerfully addressed.Aproximadamente cada 10 años, el Comité de Colecciones Sistemáticas (CCS) de la Sociedad Americana de Mastozoologia, evalúa el estado de las colecciones mastozoológicas del hemisferio occidental. En el último censo, el CCS encuestó un total de 429 colecciones y compiló un directorio de 395 colecciones activas que contenían 5,275,155 especímenes catalogados. En comparación con el censo previo, durante la última década 43 colecciones se han cerrado o han sido absorbidas, pero se agregaron al censo 38 nuevas colecciones. Se documentó un incremento en el número total de especímenes, la expansión de la disponibilidad de colecciones de recursos genómicos, además de avances substanciales en digitalización y accesibilidad a la web. También, se detectaron cambios en las proporciones de grupos taxonómicos debido a la representación geográfica cada vez más equilibrada de las colecciones en comparación con encuestas anteriores. Si bien consideramos que las colecciones del hemisferio occidental estan en buen estado en algunas áreas, también identificamos brechas claras en la cobertura espacial y temporal, así como amenazas al crecimiento y vitalidad de estos recursos a largo plazo. Un crecimiento substancial, acompañado de compromisos de adecuado financiamiento, serán necesarios para asegurar que las colecciones incluyan muestras lo suficientemente amplias y variadas como para permitir a mastozoologos y otros científicos abordar las necesidades, muchas de ellas imprevistas, que traerá el futuro.

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& Geociencias Noroeste Argentino, Salta, Argentina Inst Venezolano Invest Cient, Caracas, Venezuela Umweltbundesamt, Vienna, Austria Penclen, Plumelec, France CNRS MNHN UPMC, Ctr Rech Paleobiodivers & Paleoenvironm, Paris, France Staatl Museum Nat Kunde, Stuttgart, Germany Consejo Nacl Invest Cient & Tecn, Museo Argentino Ciencias Nat Bernardino Rivada, Buenos Aires, DF, Argentina Ctr Biol Gest Populat INRA, Montferrier Sur Lez, France Loyola Univ Chicago, Chicago, IL USA Pontificia Univ Catolica Rio do Sul, Porto Alegre, RS, Brazil Durban Museum Nat Sci, Durban, South Africa Univ Estado Amazonas, Manaus, Amazonas, Brazil Brigham Young Univ, Provo, UT 84602 USA Museo Hist Nat La Salle, Caracas, Venezuela Univ Vet Med Hannover, Hannover, Germany Australian Museum, Sydney, NSW, Australia Texas Tech Univ, Lubbock, TX 79409 USA Francis Marion Univ, Florence, SC USA Univ Washington, Seattle, WA 98195 USA Univ Connecticut, Groton, CT USA Fed Univ ABC, Santo Andre, SP, Brazil Univ Fed Uberlandia, Uberlandia, MG, Brazil Fed Univ Para, Belem, Para, Brazil Yale Peabody Museum, New Haven, CT USA Univ Texas Arlington, Arlington, TX 76019 USA Senckenberg Deutsch Entomol Inst, Muncheberg, Germany CSIRO, Natl Res Collect, Canberra, ACT, Australia Univ Venda, Thohoyandou, South Africa Univ Ulsan, Ulsan, South Korea Univ Estadual Sao Paulo, Botucatu, SP, Brazil Senckenberg Forsch Inst & Nat Museum, Frankfurt, Germany Ocean Univ China, Inst Marine Biodivers & Evolut, Qingdao, Shandong, Peoples R China Univ Fed Para, Altamira, Brazil Univ Estadual Paulista, Assis, Brazil UPMC, Ctr Rech Paleobiodiversite & Paleoenvironm, CNRS, MNHN, Paris, France Tech Univ Kaiserslautern, Kaiserslautern, Germany Charles Univ Prague, Dept Zool, Prague, Czech Republic Univ Nacl Expt los Llanos Occident Ezequiel Zamor, Guanare, Venezuela Long Isl Univ, Brooklyn, NY USA Univ Ryukyus, Okinawa, Japan Pontificia Univ Catolica Ecuador, Museo Zool, Escuela Ciencias Biol, Quito, Ecuador Indiana Univ Penn, Indiana, PA USA IRD, Dakar, Senegal State Museum Nat Hist Karlsruhe, Karlsruhe, Germany Univ Toulouse III Paul Sabatier, Toulouse, France Univ Ghent, Ghent, Belgium Univ Antwerp, Antwerp, Belgium Karlsruhe Inst Technol, Karlsruhe, Germany Ctr Cellular & Mol Biol, Hyderabad, India Univ Fed ABC, Santo Andre, Brazil Yale Peabody Museum Nat Hist, New Haven, CT USA Univ Innsbruck, Mondsee, Austria Bethune Cookman Univ, Daytona Beach, FL USA Natl Museum Nat & Sci, Tokyo, Japan Zool Garten Koln, Cologne, Germany Univ Estadual Paulista, Botucatu, SP, Brazil Univ Estadual Paulista, Sao Vicente, Brazil Univ Estadual Paulista, Rio Claro, Brazil Univ Estadual Paulista, Sao Jose Do Rio Preto, Brazil Univ Estadual Paulista, Assis, Brazil

The Guiana Shield of South America contains savannas within one of the largest contiguous expanses of pristine tropical rainforest remaining in the world, but biodiversity in the grasslands is poorly known. In lowland Neotropical areas, bats typically comprise the most species-rich group of mammals. We compare the bat faunal community and phylogeography in the savanna habitats of the Llanos in Venezuela, Rupununi in Guyana, and Sipaliwini in Suriname. Measures of species diversity and relative abundance from standardized field survey methodology enable comparison among these three grassland regions. Genetic variation is summarized by DNA barcoding to examine biogeographic patterns across larger forest&ndash, savanna landscapes. A total of 76 species of bats is documented, of which 18 species are reported from all 3 savannas and 30 species are reported from only 1 of the savannas. Endemism is low with 5 taxa restricted primarily to dry, open habitats. However, 7 other species have divergent phylogeographic lineages associated with savanna populations. Although bat species are usually distributed over wide regions of the Neotropics, the habitat mosaics of the Guiana Shield have different faunal assemblages. Going back into the Miocene, the contractions and expansions of forest&ndash, savanna paleoenvironments over time have contributed to speciation and the current high levels of biodiversity in South America.

We report on the first comprehensive DNA barcoding survey of bats from Jamaica and compare the genetic variation to similar species on South America and Central America. Bats comprise the majority of mammalian diversity in typical lowland forest in the Neotropics, but the Caribbean is one noticeable geographic gap in the International Barcode of Life reference database. Of the 20 known species reported from Jamaica, half were DNA barcoded and were genetically distinct with interspecific variation ranging from 17 to 33%. By contrast, intraspecific variation ranged from 0 to 0.5% indicating that the barcode gap was sufficient in differentiating bat species diversity in Jamaica. The low levels of intraspecific divergence indicate that the populations within each species are relatively homogeneous across the island. There were, however, several cases of high sequence divergence for widely distributed species that occur on both the Caribbean islands and the continental mainland, which warrant further taxonomic study.

There are almost 1,700 species of mammals occurring in the Neotropical region broadly defined as including Mexico, Central America, the Caribbean Islands, and South America. The Barcode of Life Data (BOLD) Systems estab- lished an institutional collaboration with the Royal Ontario Museum (ROM) beginning in 2005 to DNA barcode the mammal tissue collections for the cytochrome oxidase subunit I (COI) gene region. To date, over 14,500 vouchered specimens have been sequenced representing nearly 19% (317 species) of the mammalian diversity in the Neotropics. This includes 11 of the 15 orders found in the Neotropics with the 4 missing orders representing less than 20 species. Al- most half of the bat species (167 of 345) have been barcoded but only 3% (4 of 129) of the primate species have been done. Rodents are the most diverse order in the Neotropics, of which 12% of the species have been barcoded (106 of 898). In terms of geographic coverage, Middle America is the most comprehensively represented with only one country missing (Honduras), but half of the South American countries are not represented and there are no samples from the Caribbean. Although an initial DNA barcode reference library for Neotropical mammals has been established, there are obvious taxo- nomic and geographic areas that need to be surveyed to make it broadly applicable for species identification and discov- ery. Nevertheless, preliminary results indicate an underestimation of approximately one-third of the currently recognized species of mammals in the Neotropics, affirming it as a region of high biodiversity.

The biodiversity of tube-nosed bats (Murininae) from Asia has been underestimated by over 50%. Since 2005, eight taxa have been documented as distinct from the 19 previously known species. We describe three new species of Marina collected in southern China between 2004 and 2007. These species differ morphologically, morphometrically, and genetically from previously described species of Marina. Morphological differences include pelage color, size, skull shape and tooth morphology. Analysis of mitochondrial DNA barcodes of the cytochrome c oxidase subunit I (COI) gene supports species status based on divergent phylogenetic lineages.

Platyrrhinus guianensis new species Guianan Broad-nosed Bat Figures 4���7 P [latyrrhinus]. helleri: Lim, 1993: 162 (part) Platyrrhinus helleri: Smith and Kerry, 1996: 932 (part) Platyrrhinus helleri: Lim and Engstrom, 2000: 121 P [latyrrhinus]. helleri: Lim and Engstrom, 2001 a: 632 (part) Platyrrhinus helleri: Lim and Engstrom, 2001 b: 664 (part) Platyrrhinus helleri: Engstrom and Lim, 2002: 364 (part) Platyrrhinus helleri: Lim and Norman, 2002: 54 P [latyrrhinus]. helleri: Lim et al., 2002: 1239 (part) Platyrrhinus helleri: Lim and Engstrom, 2005: 77 (part) Platyrrhinus helleri: Lim et al., 2005 a: 244 (part) Platyrrhinus helleri: Lim et al., 2005 b: 87 (part) Platyrrhinus helleri: Clare et al., 2007: 187 (part) Platyrrhinus helleri: Borisenko et al., 2008: 475 (part) Platyrrhinus helleri: Lim, 2009: 45 (part) P [latyrrhinus]. recifinus: Velazco, 2009: 259 (part) Platyrrhinus recifinus: Tavares and Velazco, 2010: 119 (part) Platyrrhinus helleri: Clare et al., 2011: 8 (part) Platyrrhinus helleri: Clare, 2011: 4 (part) Platyrrhinus helleri: Lim, 2012: 253 (part) Platyrrhinus helleri: Lim and Tavares, 2012: 115 (part) Holotype. Dried skin, skull and skeleton of an adult pregnant female, Royal Ontario Museum (ROM) number 113465, obtained 20 September 2001 by Burton K. Lim and Zacharias Norman (original field number F 50445). The skin, skull, and skeleton are in good condition. Frozen tissues are deposited at the Royal Ontario Museum (F 50445). Type locality. Pobawau Creek mouth, 100 m; Upper Takutu-Upper Essequibo; Guyana, 3 �� 16 ��� 3.1 ���N, 58 �� 46 ��� 42.7 ���W (Fig. 3). Paratypes. The skin, skull, and skeleton of an adult male (ROM 108487) caught on 8 October 1997 at 38 mi Camp, 35 km SW Kurupukari, 100 m, Iwokrama Forest, Potaro-Siparuni, Guyana, 4 �� 22 ���W, 58 �� 51 ���W; one skin and skull of an adult male (ROM 114070) caught on 16 April 2002 and one skin, skull, and skeleton of an adult pregnant female (ROM 113991) caught on 13 April 2002 at Brownsberg Nature Park headquarters, 500 m, Brokopondo, Suriname, 4 �� 57 ���N, 55 �� 11 ���W; and the skin and skull of an adult male (ROM 114195) caught on 21 April 2002 at Km 2.4 Wittie Kreek trail, 300 m, Brownsberg Nature Park, Brokopondo, Suriname, 4 �� 56 ���N, 55 �� 10 ���W. The holotype and 4 paratypes, along with 31 other specimens from the known distributional range, are listed in Appendix 1 (Fig. 3). Measurements of each specimen of the type series of P. guianensis are provided in Table 4. Distribution. Platyrrhinus guianensis is known from Guyana and Suriname (Fig. 3). Etymology. The species name is derived from the Latin description of its endemic distribution in the Guiana region of South America. Diagnosis. Platyrrhinus guianensis is distinguished from its congeners by a combination of external and craniodental characteristics. The ventral fur is dark gray; ventral fur unicolor; dorsal stripe wide and brilliant white; fringe of hair along margin of uropatagium long, conspicuously dense, and pale yellow. The skull of P. guianensis lacks a fossa on the squamosal root of the zygomatic arch. Dentally, two stylar cuspules are present on the posterior cristid of P 4; and one stylid cuspulid on the anterior cristid of p 4. Description. Platyrrhinus guianensis is a small Platyrrhinus (FA 37���41 mm) distinguished from its sister species P. recifinus by its smaller size and shorter skull (Table 5; Velazco & Gardner 2009, Table 2���4 and 7). However, measurements of P. guianensis overlap with P. angustirostris, P. brachycephalus, P. fusciventris, P. helleri, P. i n c a r u m, and P. matapalensis (Tables 4 ���5). Dorsal fur mostly dark brown, but paler on the upper dorsum; dorsal fur is bicolored with darker tips; facial stripes wide and white; dorsal stripe brilliant white; ventral fur dark gray, individual hairs unicolored; pinnae have well-marked fold lines; tragus and anterior and posterior rims of pinnae bright yellow (Fig. 4); lateral borders of the proximal half of the noseleaf and borders of the horseshoe yellow; inferior border of the horseshoe completely free of upper lip; posterior margin of uropatagium has the shape of an inverted ���U���; hair on upper surface of feet brown, long and dense (Fig. 5); fringe of hair along the trailing margin of uropatagium long, conspicuously dense, and pale yellow; metacarpal III longer than metacarpal V. Rostrum is slender; has a well developed anterior notch in the nasals; postorbital processes moderately developed; paraoccipital processes poorly developed; two infraorbital foramina present; posterior border of hard palate ���V���-shaped (Fig. 6); fossa on the squamosal root of the zygomatic arch absent; and paraoccipital and paracondylar processes poorly developed. Upper inner incisors bilobed and convergent, not in contact, and tips extend below level of cingula of upper canines; upper outer incisors monolobate; two stylar cuspules on posterior cristid of P 4; hypoconal basin fossa of P 4 shallow; M 1 parastyle present; M 1 mesostyle absent; M 1 metacone divided in two cones; M 1 metacone labial cingulum present; stylar cuspule absent on lingual cingulum of M 1 metacone; sulcus on posterior cristid of paracone joined to cingulum of lingual face of metacone on M 1; M 1 metastyle present; M 1 protocone well developed; M 2 parastyle present; labial cingulum present on M 2 paracone; stylar cuspule on lingual face of M 2 paracone absent; M 2 metastyle present; stylar cuspule absent on lingual face of M 2 metacone; lingual cingulum of the M 2 metacone not extending to the paracone; developed M 2 hypoconal basin; M 3 minute; labial and lingual cingulids on p 4; one stylid cuspulid on anterior cristid of p 4; two stylid cuspulids on posterior cristid of p 4; m 1 paraconid poorly developed; labial and lingual cingulids present on m 1; stylid cuspulid present on anterior cristid of m 1 protoconid; m 1 metaconid well developed; m 2 hypoconid absent; stylid cuspulid between the metaconid and protoconid poorly developed on m 2; labial and lingual cingulids present on m 2. Comparisons. Platyrrhinus guianensis can be confused with P. angustirostris, P. brachycephalus, P. fusciventris, P. he l l e r i, P. i nc a r u m, and P. matapalensis because their external and cranial measurements overlap (Table 3���4). But it can be easily distinguished from P. brachycephalus and P. matapalensis by the presence of one accessory cuspulid on the anterolingual cristid of p 4 (Fig. 7) (cuspulid lacking in P. matapalensis and two accessory cuspulids present in P. brachycephalus; Velazco 2005, fig. 27). Therefore, the following comparisons focus on differentiating P. guianensis from P. angustirostris, P. fusciventris, P. helleri, and P. incarum. Externally, ventral fur is dark gray in P. guianensis and P. angustirostris (brownish gray in P. i n c ar u m; pale gray in P. h el l e r i; brown in P. fusciventris); ventral fur unicolored in P. guianensis, P. angustirostris, P. fusciventris, and P. h el l e r i (bicolored in P. i nc a r u m); dorsal stripe wide and brilliant white in P. guianensis and P. he l l e r i (conspicuous but narrow in P. angustirostris, P. fusciventris, and P. incarum); tragus and anterior and posterior rims of pinnae bright yellow in P. guianensis, P. fusciventris, and P. helleri (whitish in P. angustirostris and P. incarum); lateral borders of the proximal half of the noseleaf and borders of the horseshoe yellow in P. guianensis, P. fusciventris, and P. he l l e r i (whitish in P. angustirostris and P. i n c ar um); posterior margin of uropatagium with a shape of an inverted ���U��� in P. guianensis, P. angustirostris, and P. i n c ar u m (���V��� shaped in P. fusciventris and P. helleri); fringe of hair along margin of uropatagium long, conspicuously dense, and pale yellow in P. guianensis (long, dense, and pale brown in P. helleri; long, dense, and whitish in P. fusciventris and P. i nc a r u m; short, dense, and pale brown in P. angustirostris); hair on the upper surface of feet brown, long and dense in P. guianensis, P. angustirostris, and P. i n c a r um (short and intermediate in density in P. fusciventris and P. hell eri); metacarpal III longer than metacarpal V in P. guianensis, P. angustirostris, and P. i n c a r um (metacarpals III and V subequal in P. fusciventris and P. he l l e r i). Cranially, there is a ���V���-shaped posterior border of the hard palate in P. guianensis, P. angustirostris, P. hel leri, and P. i n c ar u m (���V���- or ���U���-shaped in P. fusciventris); fossa on the squamosal root of the zygomatic arch absent in P. guianensis, P. helleri, and P. i ncarum (shallow in P. angustirostris and P. fusciventris). Dentally, there are two stylar cuspules on posterior cristid of P 4 in P. guianensis, P. fusciventris, P. helleri, and P. i nc a r u m (three in P. angustirostris); stylar cuspule on lingual face of M 2 paracone absent in P. guianensis, P. angustirostris, and P. fusciventris (one stylar cuspule in P. he l l e r i and P. incarum); M 3 minute in P. guianensis and P. i n ca r u m (larger in P. h el l e r i, P. angustirostris, and P. fusciventris); one stylid cuspulid on the anterior cristid of p 4 in P. guianensis, P. fusciventris, and P. helleri (one or two in P. i n ca r u m and P. angustirostris); tall m 2 protoconid in P. guianensis, P. angustirostris, P. fusciventris, P. i n c a r um (Fig. 7) (shorter in P. helleri); hypoconid lacking on m 2 in P. guianensis, P. angustirostris, P. fusciventris, and P. helleri (poorly developed in P. i n c a r um); poorly developed stylid cuspulid between the metaconid and protoconid on m 2 in P. guianensis, P. fusciventris, and P. helleri (well developed in P. i ncarum and P. angustirostris). Natural history. Platyrrhinus guianensis has been documented from an elevational range of 60 to 500 m and is found primarily in rainforest (n= 33), but 3 individuals were netted in savanna. Of the 36 specimens examined, 16 are males and 20 females. Testes size (length by width in mm) ranged from 3 by 2 to 5 by 4. From 12 January to 9 February 2006, 8 of 10 females were pregnant with crown-rump (CR) measurements ranging from 4 to 13 mm. A female was pregnant on 13 April 2002 and another on 27 July 2009 with CR of 13 mm and 4 mm, respectively. Three females were pregnant on 20 and 21 September 2001 with CR ranging from 18 to 21 mm. A non-pregnant female was collected on 24 October 1997. A female had an embryo with CR of 26 mm collected on 31 October 2005. Two non-pregnant lactating females were caught on 8 and 11 November 1999 and one non-pregnant female was caught on 19 November 1997., Published as part of Velazco, Pa��l M. & Lim, Burton K., 2014, A new species of broad-nosed bat Platyrrhinus Saussure, 1860 (Chiroptera: Phyllostomidae) from the Guianan Shield, pp. 175-193 in Zootaxa 3796 (1) on pages 181-189, DOI: 10.11646/zootaxa.3796.1.9, http://zenodo.org/record/225163, {"references":["Lim, B. K. (1993) Cladistic reappraisal of stenodermatine bat phylogeny. Cladistics, 9, 147 - 165. http: // dx. doi. org / 10.1006 / clad. 1993.1009","Smith, P. G. & Kerry, S. M. (1996) The Iwokrama Rain Forest Programme for sustainable development: How much of Guyana's bat (Chiroptera) diversity does it encompass? Biodiversity and Conservation, 5, 921 - 942. http: // dx. doi. org / 10.1007 / bf 00054742","Lim, B. K. & Engstrom, M. D. (2000) Preliminary survey of bats from the upper Mazaruni of Guyana. Chiroptera Neotropical, 6 (1 - 2), 119 - 123.","Lim, B. K. & Engstrom, M. D. (2001 a) Species diversity of bats (Mammalia: Chiroptera) in Iwokrama Forest, Guyana, and the Guianan subregion: implications for conservation. Biodiversity and Conservation, 10, 613 - 657. http: // dx. doi. org / 10.1023 / A: 1016660123189","Lim, B. K. & Engstrom, M. D. (2001 b) Bat community structure at Iwokrama Forest, Guyana. Journal of Tropical Ecology, 17, 647 - 665. http: // dx. doi. org / 10.1017 / s 0266467401001481","Lim, B. K. & Norman, Z. (2002) Rapid assessment of small mammals in the eastern Kanuku Mountains, Lower Kwitaro River area, Guyana. In: Montambault, J. R. & Missa, O. (Eds.), A biodiversity assessment of the eastern Kanuku Mountains, Lower Kwitaro River, Guyana. Conservation International, Arlington, Virginia, pp. 51 - 58.","Lim, B. K., Engstrom, M. D., Genoways, H. H., Catzeflis, F. M., Fitzgerald, K. A., Peters, S. L., Djosetro, M., Brandon, S. & Mitro, S. (2005 a) Results of the Alcoa Foundation-Suriname expeditions. XIV. Mammals of Brownsberg Nature Park, Suriname. Annals of Carnegie Museum, 74, 225 - 274. http: // dx. doi. org / 10.2992 / 0097 - 4463 (2005) 74 [225: rotafe] 2.0. co; 2","Lim, B. K., Engstrom, M. D. & Ochoa G., J. (2005 b) Mammals. In: Hollowell, T. & Reynolds, R. P. (Eds.), Checklist of the terrestrial vertebrates of the Guiana Shield. Bulletin of the Biological Society of Washington, 13, 77 - 92.","Clare, E. L., Lim, B. K., Engstrom, M. D., Eger, J. L. & Hebert, P. D. N. (2007) DNA barcoding of Neotropical bats: species identification and discovery within Guyana. Molecular Ecology Notes, 7, 184 - 190. http: // dx. doi. org / 10.1111 / j. 1471 - 8286.2006.01657. x","Borisenko, A. V., Lim, B. K., Ivanova, N. V., Hanner, R. H. & Hebert, P. D. N. (2008) DNA barcoding in surveys of small mammal communities: a field study in Suriname. Molecular Ecology Resources, 8, 471 - 479. http: // dx. doi. org / 10.1111 / j. 1471 - 8286.2007.01998. x","Lim, B. K. (2009) Environmental assessment at the Bakhuis bauxite concession: small-sized mammal diversity and abundance in the lowland humid forests of Suriname. The Open Biology Journal, 2, 42 - 53. http: // dx. doi. org / 10.2174 / 1874196700902010042","Tavares, V. C. & Velazco, P. M. (2010) Platyrrhinus recifinus (Chiroptera: Phyllostomidae). Mammalian Species, 42, 119 - 123. http: // dx. doi. org / 10.1644 / 859.1","Lim, B. K. & Tavares, V. C. (2012) Review of species richness and biogeography of bats (Chiroptera) from the Guiana subregion of South America with comments on conservation. Ecotropica, 18, 195 - 118.","Velazco, P. M. & Gardner, A. L. (2009) A new species of Platyrrhinus (Chiroptera: Phyllostomidae) from western Colombia and Ecuador, with emended diagnoses of P. aquilus, P. dorsalis, and P. umbratus. Proceedings of the Biological Society of Washington, 122, 249 - 281. http: // dx. doi. org / 10.2988 / 08 - 40.1","Velazco, P. M. (2005) Morphological phylogeny of the bat genus Platyrrhinus Saussure, 1860 (Chiroptera: Phyllostomidae) with the description of four new species. Fieldiana Zoology, New Series, 105, 1 - 53. http: // dx. doi. org / 10.5962 / bhl. title. 2689"]}

Velazco, Paúl M., Lim, Burton K. (2014): A new species of broad-nosed bat Platyrrhinus Saussure, 1860 (Chiroptera: Phyllostomidae) from the Guianan Shield. Zootaxa 3796 (1): 175-193, DOI: http://dx.doi.org/10.11646/zootaxa.3796.1.9

Platyrrhinus guianensis new species Guianan Broad-nosed Bat Figures 4–7 P [latyrrhinus]. helleri: Lim, 1993: 162 (part) Platyrrhinus helleri: Smith and Kerry, 1996: 932 (part) Platyrrhinus helleri: Lim and Engstrom, 2000: 121 P [latyrrhinus]. helleri: Lim and Engstrom, 2001 a: 632 (part) Platyrrhinus helleri: Lim and Engstrom, 2001 b: 664 (part) Platyrrhinus helleri: Engstrom and Lim, 2002: 364 (part) Platyrrhinus helleri: Lim and Norman, 2002: 54 P [latyrrhinus]. helleri: Lim et al., 2002: 1239 (part) Platyrrhinus helleri: Lim and Engstrom, 2005: 77 (part) Platyrrhinus helleri: Lim et al., 2005 a: 244 (part) Platyrrhinus helleri: Lim et al., 2005 b: 87 (part) Platyrrhinus helleri: Clare et al., 2007: 187 (part) Platyrrhinus helleri: Borisenko et al., 2008: 475 (part) Platyrrhinus helleri: Lim, 2009: 45 (part) P [latyrrhinus]. recifinus: Velazco, 2009: 259 (part) Platyrrhinus recifinus: Tavares and Velazco, 2010: 119 (part) Platyrrhinus helleri: Clare et al., 2011: 8 (part) Platyrrhinus helleri: Clare, 2011: 4 (part) Platyrrhinus helleri: Lim, 2012: 253 (part) Platyrrhinus helleri: Lim and Tavares, 2012: 115 (part) Holotype. Dried skin, skull and skeleton of an adult pregnant female, Royal Ontario Museum (ROM) number 113465, obtained 20 September 2001 by Burton K. Lim and Zacharias Norman (original field number F 50445). The skin, skull, and skeleton are in good condition. Frozen tissues are deposited at the Royal Ontario Museum (F 50445). Type locality. Pobawau Creek mouth, 100 m; Upper Takutu-Upper Essequibo; Guyana, 3 ° 16 ’ 3.1 ”N, 58 ° 46 ’ 42.7 ”W (Fig. 3). Paratypes. The skin, skull, and skeleton of an adult male (ROM 108487) caught on 8 October 1997 at 38 mi Camp, 35 km SW Kurupukari, 100 m, Iwokrama Forest, Potaro-Siparuni, Guyana, 4 ° 22 ’W, 58 ° 51 ’W; one skin and skull of an adult male (ROM 114070) caught on 16 April 2002 and one skin, skull, and skeleton of an adult pregnant female (ROM 113991) caught on 13 April 2002 at Brownsberg Nature Park headquarters, 500 m, Brokopondo, Suriname, 4 ° 57 ’N, 55 ° 11 ’W; and the skin and skull of an adult male (ROM 114195) caught on 21 April 2002 at Km 2.4 Wittie Kreek trail, 300 m, Brownsberg Nature Park, Brokopondo, Suriname, 4 ° 56 ’N, 55 ° 10 ’W. The holotype and 4 paratypes, along with 31 other specimens from the known distributional range, are listed in Appendix 1 (Fig. 3). Measurements of each specimen of the type series of P. guianensis are provided in Table 4. Distribution. Platyrrhinus guianensis is known from Guyana and Suriname (Fig. 3). Etymology. The species name is derived from the Latin description of its endemic distribution in the Guiana region of South America. Diagnosis. Platyrrhinus guianensis is distinguished from its congeners by a combination of external and craniodental characteristics. The ventral fur is dark gray; ventral fur unicolor; dorsal stripe wide and brilliant white; fringe of hair along margin of uropatagium long, conspicuously dense, and pale yellow. The skull of P. guianensis lacks a fossa on the squamosal root of the zygomatic arch. Dentally, two stylar cuspules are present on the posterior cristid of P 4; and one stylid cuspulid on the anterior cristid of p 4. Description. Platyrrhinus guianensis is a small Platyrrhinus (FA 37–41 mm) distinguished from its sister species P. recifinus by its smaller size and shorter skull (Table 5; Velazco & Gardner 2009, Table 2–4 and 7). However, measurements of P. guianensis overlap with P. angustirostris, P. brachycephalus, P. fusciventris, P. helleri, P. i n c a r u m, and P. matapalensis (Tables 4 –5). Dorsal fur mostly dark brown, but paler on the upper dorsum; dorsal fur is bicolored with darker tips; facial stripes wide and white; dorsal stripe brilliant white; ventral fur dark gray, individual hairs unicolored; pinnae have well-marked fold lines; tragus and anterior and posterior rims of pinnae bright yellow (Fig. 4); lateral borders of the proximal half of the noseleaf and borders of the horseshoe yellow; inferior border of the horseshoe completely free of upper lip; posterior margin of uropatagium has the shape of an inverted ‘U’; hair on upper surface of feet brown, long and dense (Fig. 5); fringe of hair along the trailing margin of uropatagium long, conspicuously dense, and pale yellow; metacarpal III longer than metacarpal V. Rostrum is slender; has a well developed anterior notch in the nasals; postorbital processes moderately developed; paraoccipital processes poorly developed; two infraorbital foramina present; posterior border of hard palate ‘V’-shaped (Fig. 6); fossa on the squamosal root of the zygomatic arch absent; and paraoccipital and paracondylar processes poorly developed. Upper inner incisors bilobed and convergent, not in contact, and tips extend below level of cingula of upper canines; upper outer incisors monolobate; two stylar cuspules on posterior cristid of P 4; hypoconal basin fossa of P 4 shallow; M 1 parastyle present; M 1 mesostyle absent; M 1 metacone divided in two cones; M 1 metacone labial cingulum present; stylar cuspule absent on lingual cingulum of M 1 metacone; sulcus on posterior cristid of paracone joined to cingulum of lingual face of metacone on M 1; M 1 metastyle present; M 1 protocone well developed; M 2 parastyle present; labial cingulum present on M 2 paracone; stylar cuspule on lingual face of M 2 paracone absent; M 2 metastyle present; stylar cuspule absent on lingual face of M 2 metacone; lingual cingulum of the M 2 metacone not extending to the paracone; developed M 2 hypoconal basin; M 3 minute; labial and lingual cingulids on p 4; one stylid cuspulid on anterior cristid of p 4; two stylid cuspulids on posterior cristid of p 4; m 1 paraconid poorly developed; labial and lingual cingulids present on m 1; stylid cuspulid present on anterior cristid of m 1 protoconid; m 1 metaconid well developed; m 2 hypoconid absent; stylid cuspulid between the metaconid and protoconid poorly developed on m 2; labial and lingual cingulids present on m 2. Comparisons. Platyrrhinus guianensis can be confused with P. angustirostris, P. brachycephalus, P. fusciventris, P. he l l e r i, P. i nc a r u m, and P. matapalensis because their external and cranial measurements overlap (Table 3–4). But it can be easily distinguished from P. brachycephalus and P. matapalensis by the presence of one accessory cuspulid on the anterolingual cristid of p 4 (Fig. 7) (cuspulid lacking in P. matapalensis and two accessory cuspulids present in P. brachycephalus; Velazco 2005, fig. 27). Therefore, the following comparisons focus on differentiating P. guianensis from P. angustirostris, P. fusciventris, P. helleri, and P. incarum. Externally, ventral fur is dark gray in P. guianensis and P. angustirostris (brownish gray in P. i n c ar u m; pale gray in P. h el l e r i; brown in P. fusciventris); ventral fur unicolored in P. guianensis, P. angustirostris, P. fusciventris, and P. h el l e r i (bicolored in P. i nc a r u m); dorsal stripe wide and brilliant white in P. guianensis and P. he l l e r i (conspicuous but narrow in P. angustirostris, P. fusciventris, and P. incarum); tragus and anterior and posterior rims of pinnae bright yellow in P. guianensis, P. fusciventris, and P. helleri (whitish in P. angustirostris and P. incarum); lateral borders of the proximal half of the noseleaf and borders of the horseshoe yellow in P. guianensis, P. fusciventris, and P. he l l e r i (whitish in P. angustirostris and P. i n c ar um); posterior margin of uropatagium with a shape of an inverted ‘U’ in P. guianensis, P. angustirostris, and P. i n c ar u m (‘V’ shaped in P. fusciventris and P. helleri); fringe of hair along margin of uropatagium long, conspicuously dense, and pale yellow in P. guianensis (long, dense, and pale brown in P. helleri; long, dense, and whitish in P. fusciventris and P. i nc a r u m; short, dense, and pale brown in P. angustirostris); hair on the upper surface of feet brown, long and dense in P. guianensis, P. angustirostris, and P. i n c a r um (short and intermediate in density in P. fusciventris and P. hell eri); metacarpal III longer than metacarpal V in P. guianensis, P. angustirostris, and P. i n c a r um (metacarpals III and V subequal in P. fusciventris and P. he l l e r i). Cranially, there is a ‘V’-shaped posterior border of the hard palate in P. guianensis, P. angustirostris, P. hel leri, and P. i n c ar u m (‘V’- or ‘U’-shaped in P. fusciventris); fossa on the squamosal root of the zygomatic arch absent in P. guianensis, P. helleri, and P. i ncarum (shallow in P. angustirostris and P. fusciventris). Dentally, there are two stylar cuspules on posterior cristid of P 4 in P. guianensis, P. fusciventris, P. helleri, and P. i nc a r u m (three in P. angustirostris); stylar cuspule on lingual face of M 2 paracone absent in P. guianensis, P. angustirostris, and P. fusciventris (one stylar cuspule in P. he l l e r i and P. incarum); M 3 minute in P. guianensis and P. i n ca r u m (larger in P. h el l e r i, P. angustirostris, and P. fusciventris); one stylid cuspulid on the anterior cristid of p 4 in P. guianensis, P. fusciventris, and P. helleri (one or two in P. i n ca r u m and P. angustirostris); tall m 2 protoconid in P. guianensis, P. angustirostris, P. fusciventris, P. i n c a r um (Fig. 7) (shorter in P. helleri); hypoconid lacking on m 2 in P. guianensis, P. angustirostris, P. fusciventris, and P. helleri (poorly developed in P. i n c a r um); poorly developed stylid cuspulid between the metaconid and protoconid on m 2 in P. guianensis, P. fusciventris, and P. helleri (well developed in P. i ncarum and P. angustirostris). Natural history. Platyrrhinus guianensis has been documented from an elevational range of 60 to 500 m and is found primarily in rainforest (n= 33), but 3 individuals were netted in savanna. Of the 36 specimens examined, 16 are males and 20 females. Testes size (length by width in mm) ranged from 3 by 2 to 5 by 4. From 12 January to 9 February 2006, 8 of 10 females were pregnant with crown-rump (CR) measurements ranging from 4 to 13 mm. A female was pregnant on 13 April 2002 and another on 27 July 2009 with CR of 13 mm and 4 mm, respectively. Three females were pregnant on 20 and 21 September 2001 with CR ranging from 18 to 21 mm. A non-pregnant female was collected on 24 October 1997. A female had an embryo with CR of 26 mm collected on 31 October 2005. Two non-pregnant lactating females were caught on 8 and 11 November 1999 and one non-pregnant female was caught on 19 November 1997.

An inventory of mammals in the vicinity of Brownsberg Nature Park, Suriname, incorporated a number of different sampling methods including examining museum voucher specimens, an animal-rescue operation, transect surveys, camera photo-traps, and interviews with local inhabitants. We document a total of 125 mammal species present in the Park. These include ten opossums, five pilosans, four armadillos, 58 bats, eight monkeys, 13 carnivores, one tapir, four artiodactyls, and 22 rodents. Nine of these species are reported for the first time from Suriname: one mouse opossum (Marmosops pinheiroi); one naked-backed moustached bat (Pteronotus gymnonotus); four fruit-eating bats (Artibeus bogotensis, A. gnomus, A. obscurus, and A. planirostris); two evening bats (Eptesicus chiriquinus and Myotis riparius);and one arboreal rice rat (Oecomys auyantepui). There are 191 indigenous non-marine mammals presently known from Suriname.

As part of a larger project on biodiversity and conservation in Guyana, we documented 130 species of mammals from Iwokrama Forest. This included 7 marsupials, 4 xenarthrans, 86 bats, 5 primates, 8 carnivores, 1 perissodactyl, 4 artiodactyls, and 15 rodents. As is typical for most Neotropical sites, the 86 species of bats represent over half of the mammal diversity. Standardized collecting methods implemented in the 1997 faunal survey of Iwokrama Forest in central Guyana enabled us also to investigate species diversity and abundance resulting from the inventory of mammals. Four species of fruit-eating bats (Artibeus lituratus, A. obscurus, A. planirostris, and Carollia perspicillata) were the most abundant and accounted for 43% of the 2,097 total captures in mist nets and harp traps during 79 nights of sampling. For nonvolant mammals, terrestrial spiny rats (Proechimys spp.) represented over half (55%) of the 65 captures in primarily live box-style traps. We estimate that our inventory is approxim...

The genetic algorithm for rule-set prediction (GARP) has beensuccessfully used in modeling species' distributions with environmental data forwell-studied birds in the United States and elsewhere. GARP's efficiency hasbeen demonstrated to be robust even with incomplete occurrence and geographicdata. Thorough biological sampling in conjunction with comprehensive geographicinformation, however, is not the norm for many tropical areas where mostbiodiversity occurs. Mammals from Guyana were used as a test of the robustnessof these approaches in a worst-case scenario of uneven sampling combined withcoarse geographic data. The occurrence of species in poorly surveyed regions,such as the Pakaraima Highlands of west-central Guyana, was consistentlyunder-predicted, whereas presence in well-surveyed areas such as thesouthwestern Rupununi was usually correctly predicted. Comparisons of numbersof species and specimens collected also indicate that lowland forests in thesoutheast and coastal forests in the northwest are under-sampled. For robustdistributional predictions in Guyana, more thorough inventories are needed inthese diverse environments.

With 86 species, Iwokrama Forest in central Guyana has the highest reported bat biodiversity for a protected area in the world. Using standardized capture data for 73 of these 86 species, we document community structure of bats in terms of species diversity, relative abundance, gross biomass, feeding guilds, vertical stratification and a trophic-size niche matrix. Based on faunal surveys in 1997, with similar amounts of effort in the forest canopy and at ground level, the greater fruit-eating bat (Artibeus lituratus) was by far the most ecologically dominant species in terms of frequency of capture and biomass. In total, frugivores comprised 70% of the species diversity and 78% of the biomass. The most common species of bat were fully partitioned in a resource niche matrix of size and trophic guild when vertical stratification was included as a variable. We conclude that resource partitioning and species packing differentially affect relative size in tropical bats, and are better summarized and analysed in three dimensions.

Voss, Robert S., Lim, Burton K., Díaz-Nieto, Juan F., Jansa, Sharon A. (2013): A New Species of Marmosops (Marsupialia: Didelphidae) from the Pakaraima Highlands of Guyana, with Remarks on the Origin of the Endemic Pantepui Mammal Fauna. American Museum Novitates 2013 (3778): 1-28, DOI: 10.1206/3778.2, URL: http://www.bioone.org/doi/abs/10.1206/3778.2

This synthesis provides information related to the Southern Nevada Agency Partnership (SNAP) Science and Research Strategy Goal 1 - to restore, sustain and enhance southern Nevada’s ecosystems - and Goal 2 - to provide for responsible use of southern Nevada’s lands in a manner that preserves heritage resources and promotes an understanding of human interaction with the landscape. The Science and Research Strategy has nine Sub-goals that address the topics of water and water use, fire, invasive species, biological diversity, restoration, cultural resources, historic content, recreation, and science-based management. This synthesis summarizes the state-of-knowledge related to each of these Sub-goals, addresses knowledge gaps, and provides management implications. It builds on previous efforts to develop the necessary scientific understanding for adaptive management of southern Nevada ecosystems.

DNA barcoding using the cytochrome c oxidase subunit 1 gene (COI) is frequently employed as an efficient method of species identification in animal life and may also be used to estimate species richness, particularly in understudied faunas. Despite numerous past demonstrations of the efficiency of this technique, few studies have attempted to employ DNA barcoding methodologies on a large geographic scale, particularly within tropical regions. In this study we survey current and potential species diversity using DNA barcodes with a collection of more than 9000 individuals from 163 species of Neotropical bats (order Chiroptera). This represents one of the largest surveys to employ this strategy on any animal group and is certainly the largest to date for land vertebrates. Our analysis documents the utility of this tool over great geographic distances and across extraordinarily diverse habitats. Among the 163 included species 98.8% possessed distinct sets of COI haplotypes making them easily recognizable at this locus. We detected only a single case of shared haplotypes. Intraspecific diversity in the region was high among currently recognized species (mean of 1.38%, range 0–11.79%) with respect to birds, though comparable to other bat assemblages. In 44 of 163 cases, well-supported, distinct intraspecific lineages were identified which may suggest the presence of cryptic species though mean and maximum intraspecific divergence were not good predictors of their presence. In all cases, intraspecific lineages require additional investigation using complementary molecular techniques and additional characters such as morphology and acoustic data. Our analysis provides strong support for the continued assembly of DNA barcoding libraries and ongoing taxonomic investigation of bats.

We report the discovery of a new species of doglike bat (Peropteryx) from the lowland Amazonian forests of Ecuador and Peru. It has transparent wing membranes that are faintly tinged brown with pale-brown arms and digits; ears that are separated on the forehead; and a skull with small, shallow pterygoid pits that are anterolateral to an undivided basisphenoid pit and that are separated by a mesopterygoid extension. These characters distinguish the new species from morphologically similar species with which it was previously confused (P. leucoptera and P. macrotis). A molecular phylogenetic analysis of unlinked loci from each of the four genetic transmission systems of mammals (mitochondrial, nuclear-autosomal, X, and Y chromosomes) independently corroborated the placement of the new species as the sister taxon to a clade that includes P. kappleri, P. macrotis, and P. trinitatis; the basal lineage for the genus is P. leucoptera. This phylogeny suggests that transparent wings (sometimes described as ‘‘white’’ but actually lacking pigment), the traditional character used to diagnose Peronymus, is not a unique synapomorphy. Furthermore, based on a molecular dating analysis, the depth of divergence of Peropteryx is equivalent to that of another New World emballonurid genus (Balantiopteryx). Therefore, Peronymus does not warrant higher-level recognition as a subgenus or genus.

Peropteryx pallidoptera, new species Pale-winged Doglike Bat Figures 1, 2, 3 HOLOTYPE: An adult male (ROM 104396) collected 66 km S of Pompeya Sur (00 �� 48 9 S, 76 �� 24 9 W), Orellana Province, Ecuador by F.A. Reid, and F. Sornoza (field number F37227) on 8 May 1995. The holotype was prepared as a skin, skull, and partial skeleton. PARATYPES: An adult male (AMNH 74107) prepared as a skin and skull collected at Orosa (03 �� 26 9 S, 72 �� 08 9 W), Amazon River, Loreto, Peru, by Alfonso and Ram��n Olalla on 28 November 1926. Seventeen individuals prepared as fluid-preserved specimens collected at Nuevo San Juan (05 �� 14 9 50 0 S, 73 �� 09 9 50 0 W, 150 m above sea level), R��o G��lvez, Loreto, Peru, by Robert S. Voss and David W. Fleck in 1998 and 1999, including 1 adult male (MUSM 13226), 14 adult females (AMNH 272671, AMNH 272726, AMNH 272854, AMNH 272855, AMNH 272827, AMNH 273042, AMNH 273116, AMNH 273185, MUSM 13228, MUSM 13230, MUSM 15246, MUSM 15249, MUSM 15250, MUSM 15252) and 2 subadult females (MUSM 13227, MUSM 13229). DISTRIBUTION: The new species is currently known from just three localities in primary lowland forests of eastern Ecuador and northern Peru at elevations, 400 m above sea level. It is found sympatrically with P. kappleri and P. leucoptera at Nuevo San Juan. The new species and P. macrotis have overlapping distributions in the western Amazon but their closest points of documented occurrence are approximately 50 km apart (between Orosa and Puerto Indiana on the Amazon River of Peru). ETYMOLOGY: The specific epithet refers to the pale wings and is compounded from the Latin words pallidus (������pale������) and ptera (������wing������). MEASUREMENTS: External measurements are presented in table 1 and cranial measurements are presented in table 2 for Peropteryx pallidoptera and other congeneric species. DIAGNOSIS: Superficially, Peropteryx pallidoptera appears most similar to P. leucoptera because they both have transparent wings. However, the new species has paler brown arms and digits (dark brown in P. leucoptera), the skin from the wing tip to the elbow is translucent and evenly colored with a tinge of brown (gradually darkens in color from wing tip to body in P. leucoptera; fig. 1), the ears are not connected by a band of skin, the rostrum of the skull is not broad, and the lateral pterygoid pits are not as large and deep as they are in P. leucoptera (fig. 2). In contrast, P. pallidoptera is morphologically similar to P. kappleri, P. macrotis, and P. trinitatis with the ears separated, rostrum of the skull not broad, and the lateral pterygoid pits small and shallow (fig. 2). However, the new species does not have dark brown wings and digits, the rostrum is not inflated dorsally, and the posterior border of the mesopterygoid extension is not level with the anterior border of the basisphenoid and lateral pterygoid pits, all of which are characters seen in P. kappleri, P. macrotis, and P. trinitatis. Furthermore, P. pallidoptera has a tiny peglike upper anterior premolar similar to that of P. trinitatis but different from those of P. kappleri and P. macrotis; the latter two species have a slightly larger tooth, bearing a distinct posterior cusp. Peropteryx kappleri is the largest species in the genus with no overlap in forearm length and cranial measurements (except postorbital constriction) with the other species (tables 1 and 2). In general, P. pallidoptera is smaller in cranial size than P. leucoptera, P. macrotis, and P. kappleri, but it is larger than P. trinitatis. This size gradation is most evident in same-sex comparisons of condyloincisive length (table 3). 1 Pregnant (not included in mean or range). 2 Measurements of holotype from Thomas (1924). 3 Holotype. 1 Measurements of holotype from Thomas (1924). 2 Greatest length of skull to front of canine. 3 Length from condyle to front of canine. DESCRIPTION: The new species is a medium-sized Peropteryx (table 2). The dorsal fur is relatively long (approximately 8 mm) and is a uniform medium brown with slightly paler hair bases. The ventral fur is similar but slightly lighter in color. The ears are brown and are not in contact with each other, although they are close together on the forehead. The interfemoral membrane is brown, as is the wing membrane from the body to the elbow, but the distal portion is translucent with a tinge of brown (fig. 1). The arms and digits of the wings are medium brown. There is a poorly developed wing sac situated on the leading edge of the propatagium with an outward lateral opening. The skull of P. pallidoptera has slender postorbital processes and a relatively narrow rostrum, but the rostrum is not dorsally inflated. The relatively shallow, undivided basisphenoid pit has two small anterolateral pterygoid pits and the mesopterygoid extension protrudes posteriorly into the basisphenoid region (fig. 2). The upper anterior premolar is tiny and peglike. The dental formula is i 1/3, c 1/1, p 2/2, m 3/3 with a total of 32 teeth. COMMENTS: The cranial measurements for males of P. pallidoptera are all within the observed range of variation for females (table 2), which suggests that there is little or no sexual dimorphism in skull size. However, although the sample size for male P. pallidoptera is small (N 5 2), males are smaller than females with no overlap in forearm length and weight (table 1). In contrast, females of P. macrotis are significantly larger than males (p, 0.02) for all cranial measurements except breadth of braincase and postorbital constriction. Similarly, females of P. macrotis are significantly larger than males (p, 0.05) for all external measurements except length of tail and length of tragus. Brosset and Charles- Dominique (1990) did not observe sexual dimorphism for P. trinitatis but they only compared 1 male and 1 female. Although our sample size was not much better (at 2 males and 2 females), females were larger for all cranial measurements except postorbital constriction and there was no overlap in range (table 2). The male P. trinitatis reported by Brosset and Charles-Dominique (1990) was large and similar to our females except for maxillary toothrow length. However, with larger sample sizes for external measurements (10 males and 10 females), females of P. trinitatis are significantly larger (p, 0.003) than males except for length of tail and length of tragus. The sample sizes for P. kappleri and P. leucoptera were too small for making meaningful comment on cranial sexual size dimorphism. External measurements suggest that females of P. kappleri are larger than males (p, 0.005 for total length, forearm length, and weight), but there is no indication of sexual dimorphism in any of the external measurements for P. leucoptera. NATURAL HISTORY ECUADOR: Peropteryx pallidoptera was one of 66 species of bats that we captured in Block 16 in 1995 and 1996 (Reid et al., 2000). The holotype was caught in a mist net set at the entrance to a small cave next to a muddy, spring-fed saladero (mineral-rich seep) that periodically flooded the surrounding forest. The cave appeared to have been used primarily as a night roost, because few bats were found to occupy it in the daytime. Another six specimens of Peropteryx pallidoptera (easily recognized as such by their transparent wing tips) were captured and released by Engstrom, who netted them over a larger saladero (at least 1 km in diameter) located near the edge of the Tiputini River about 10 minutes by boat north of the Tiputini Biodiversity Station (00 �� 38 9 S, 76 �� 09 9 W). The small stream that fed this saladero tumbled over the edge of a cliff ca. 10 m high, and a shallow cave was present behind the waterfall. The bats apparently used the cave as a diurnal roost: Engstrom saw small, brown emballonurids in the cave before it got dark, and he caught the six P. pallidoptera in the net nearest the cave exit at dusk. PERU: We recorded a total of 58 species of bats at Nuevo San Juan in 1998 and 1999 (Fleck et al., 2002). Of 503 recorded bat captures at this locality, 372 were in mist nets and 311 were made at diurnal roosts. All 17 specimens of Peropteryx pallidoptera from Nuevo San Juan were taken from roosts. In total, we found 10 roosts occupied by this species in a variety of circumstances as described below (numbers refer to entries in a field catalog of bat roosts). Roost 2: Dark cavities among the roots and buttresses of a fallen tree in well-drained primary forest. Exactly where the bats were roosting is unknown because they had been spooked by kids who were shooting at them with arrows. A total of eight bats were collected as they perched on nearby tree trunks. Three (one adult male, one adult female, and one subadult female) were Peropteryx pallidoptera and five were Saccopteryx bilineata; an unknown number of either or both species might have escaped (22 May 1998). Roost 6: A half-open hollow log (about 50 cm inside diameter) in old secondary growth near a stream. Two bats were found roosting here, only one of which (an adult female Peropteryx pallidoptera) was collected (11 June 1998). Roost 15: An undercut earth bank in the side of a stream gully in well-drained primary forest. Two adult female and one subadult female Peropteryx pallidoptera occupied this roost, where they were found hanging well apart from one another (30 June 1998). Roost 20: A small dark cavity of roots and earth at the base of a large fallen tree in welldrained primary forest. Two bats were found roosting here about 50 cm above the ground, of which only one (an adult female Peropteryx pallidoptera) was collected (8 July 1998). Roost 22: A dark horizontal chamber under a buttress of a fallen tree in well-drained primary forest. Two adult female Peropteryx pallidoptera were found roosting here, about 80 cm above the ground (8 July 1998). Roost 31: A deep armadillo burrow in the side of a stream headwater gully in welldrained primary forest. One adult female Peropteryx pallidoptera was found roosting here in the company of at least two Micronycteris matses and three Carollia brevicauda. No bats were seen to escape this roost by flying out of it, but some might have escaped by flying deeper inside (4 September 1999). Roost 56: The unmodified leaf of a wild banana plant in hilltop primary forest. The single bat (a pregnant female Peropteryx pallidoptera) observed and collected here was hanging from the middle of the leaf about 1 m above the ground (16 September 1999). Roost 73: A hole in the ground in the side of a stream headwater gully on a hillside in well-drained primary forest. Two bats were observed roosting here, of which one (a pregnant female Peropteryx pallidoptera) was collected (22 September 1999). Roost 77: A hole in the side of a stream headwater gully in hilltop primary forest. Four bats were observed using this roost, of which two were collected. One of those collected was an adult female Peropteryx pallidoptera, and the other was an adult male P. leucoptera (23 September 1999). Roost 102: The undercut bank of a small stream in well-drained primary forest. Only a single bat (an adult female Peropteryx pallidoptera) was observed and collected here (11 October 1999). Roost 147: The ������underside of a log������ [as described by the Matses collector; presumably elevated somehow above the ground] in well-drained primary forest in a valley bottom. A single lactating adult female Peropteryx pallidoptera was collected here, but it seems likely that one or more nursing young may have been present but unobserved (27 October 1999). Roost 154: The underside of a fallen tree in primary upland forest near a small stream. A single female Peropteryx pallidoptera was in the roost (2 November 1999). The specimen from Orosa was collected by Alfonso and Ram��n Olalla, members of an Ecuadorean family of professional collectors employed by the American Museum of Natural History, who worked at or near the mouth of the R��o Orosa on the right (south) bank of the Amazon from 30 August to 11 December 1926. According to Wiley (in press), the Olallas��� camp at Orosa must have been in the floodplain of the Amazon, which is so wide at this point (about 15 km) that most of their specimens certainly came from floodplain habitats. Although the forest within several kilometers of the riverbank at the Olallas��� presumed campsite (ca. 3 �� 31 9 31 0 S, 73 �� 11 9 22 0 W; Wiley, in press) is seasonally flooded varzea, the river was probably at or near its lowest annual stage in August��� December, so this habitat would have been easy to traverse on foot., Published as part of Lim, Burton K., Engstrom, Mark D., Reid, Fiona A., Simmons, Nancy B., Voss, Robert S. & Fleck, David W., 2010, A New Species of Peropteryx (Chiroptera: Emballonuridae) from Western Amazonia with Comments on Phylogenetic Relationships within the Genus, pp. 1-20 in American Museum Novitates 3686 on pages 3-14, DOI: 10.1206/691.1, http://zenodo.org/record/4735520, {"references":["Brosset, A., and P. Charles-Dominique. 1990. The bats from French Guiana: a taxonomic, faunistic and ecological approach. Mammalia 54: 509 - 560.","Reid, F. A., M. D. Engstrom, and B. K. Lim. 2000. Noteworthy records of bats from Ecuador. Acta Chiropterologica 2: 37 - 51.","Fleck, D. W., R. S. Voss, and N. B. Simmons. 2002. Underdifferentiated taxa and sublexical categorization: an example from Matses classification of bats. Journal of Ethnobiology 22: 61 - 102."]}

Background Southeast Asia is recognized as a region of very high biodiversity, much of which is currently at risk due to habitat loss and other threats. However, many aspects of this diversity, even for relatively well-known groups such as mammals, are poorly known, limiting ability to develop conservation plans. This study examines the value of DNA barcodes, sequences of the mitochondrial COI gene, to enhance understanding of mammalian diversity in the region and hence to aid conservation planning. Methodology and Principal Findings DNA barcodes were obtained from nearly 1900 specimens representing 165 recognized species of bats. All morphologically or acoustically distinct species, based on classical taxonomy, could be discriminated with DNA barcodes except four closely allied species pairs. Many currently recognized species contained multiple barcode lineages, often with deep divergence suggesting unrecognized species. In addition, most widespread species showed substantial genetic differentiation across their distributions. Our results suggest that mammal species richness within the region may be underestimated by at least 50%, and there are higher levels of endemism and greater intra-specific population structure than previously recognized. Conclusions DNA barcodes can aid conservation and research by assisting field workers in identifying species, by helping taxonomists determine species groups needing more detailed analysis, and by facilitating the recognition of the appropriate units and scales for conservation planning.

A new species of broad-nosed bat Platyrrhinus Saussure, 1860 (Chiroptera: Phyllostomidae: Stenodermatinae) from the Guianan Shield is described based on molecular and morphological data. Previously confused with P. helleri and P. recifinus, the new taxon is currently known from only Guyana and Suriname and is most closely related to P. recifinus from eastern Brazil and not to the two sympatric species (P. fusciventris and P. incarum) also recently recognized as distinct from P. helleri. Morphometrically the new taxon overlaps with the smaller species of the genus (P. angustirostris, P. brachycephalus, P. fusciventris, P. helleri, P. incarum, and P. matapalensis), but forms a different cluster from the larger P. recifinus. Morphologically the new taxon is distinguished from its congeners by a combination of external and craniodental characteristics. Platyrrhinus now includes 21 species making it the most speciose genus in the Neotropical family Phyllostomidae.