First record of the De Vivo ’ s Disk-winged Bat , Thyroptera devivoi Gregorin , Gonçalves , Lim & Engstrom , 2006 ( Chiroptera , Thyropteridae ) , from Colombia , with comments about the record of Thyroptera lavali Pine , 1993 from the country

We report the first record of Thyroptera devivoi from Colombia in a palm swamp of Mauritia flexuosa in an aeolian seasonally flooded savanna ecosystem. This record is the seventh specimen and the fifth locality known for T. devivoi and extends the known distribution of the species 1250 km northwest from nearest locality in Guyana. We revised the specimen that was previously reported as voucher material of Thyroptera lavali from Colombia, but it matches in all characters with Thyroptera tricolor. We considered the presence of T. lavali in Colombia uncertain.


Introduction
The genus Thyroptera Spix, 1823 is a group of Neotropical insectivorous foliage-roosting bats characterized by the presence of adhesive disks in the base of the thumbs and the soles of their feet.The genus is distributed from southern Mexico across Central America and northern South America to southern Brazil.Currently, 5 species of Thyroptera are recognized (Wilson 2008, Velazco et al. 2014).
Three species of Thyroptera have been recorded in Colombia: T. tricolor Spix, 1823 has a spread distribution in the country; T. discifera (Lichtenstein & Peters, 1854) is known from Andean foothills, Caribbean and Pacific regions; and T. lavali Pine, 1993 has been reported from one locality from the Andean region (Solari et al. 2013, Reyes-Amaya et al. 2016).Additionally, fragmented fossils from middle Miocene from La Venta (Huila department) have been reported as T. lavali (Czaplewski 1996(Czaplewski , 1997)).However, the taxonomic assignment of these fossils is considered uncertain (Velazco et al. 2014).
Thyroptera devivoi is a rare species that inhabits gallery forest in savanna formations.It is known by only 6 specimens from 4 localities (1 from southwestern Guy-ana and 3 from northeastern Brazil).However, the great distance between these localities suggest that the species can be widely distributed (Gregorin et al. 2006, Wilson 2008, Santos et al. 2013, Velazco et al. 2014).
In this report, we present the first record of T. devivoi for Colombia.Additionally, we discuss the previous report of T. lavali from the Andean region of the country.

Methods
The specimen reported here was mist netted in the understory of a palm swamp of Mauritia flexuosa (locally known as "morichales", Fig. 1) at Laguna Encantada in the Hato Viejo farm, vereda Varsovia, Municipality of Paz de Ariporo, Department of Casanare (06°02′15″ N, 070°12′43″ W; 111 m above sea level [a.s.l.]) (Fig. 2).The capture took place on 9 April 2015 under the research permit 500-41-15.07209 of CORPORINO-QUIA.This place is in an aeolian seasonally flooded savanna ecosystem, dominated by grasslands, strips of gallery forest, and palm swamps associated to rivers and wetlands (Sarmiento 1983, Romero-Ruiz et al. 2010).The specimen was captured during the beginning of the rainfall season that extends from April to November in Colombian Llanos (Romero-Ruiz et al. 2010).
The specimen (ICN 21985) is an adult male with scrotal testis preserved in ethanol (96%) with the skull removed and cleaned and deposited in the mammalian collection "Alberto Cadena García" of Instituto de Ciencias Naturales of the Universidad Nacional de Colombia (ICN).
To confirm the taxonomic identification of our specimen and compare with other species of Thyroptera reported for Colombia, we examined the specimens of the genus deposited in ICN and the mammalian collection of the Museo Javeriano de Historia Natural Lorenzo Uribe (MPUJ-MAMM; MPUJ-MAMM 648; 1216).We reviewed all identifications based on characters by Pine (1993), Solari et al. (2004), Gregorin et al. (2006), Wilson (2008) and Velazco et al. (2014).The measurements reported here follow Gregorin et al. (2006) and Velazco et al. (2014).These were measured with a digital caliper to the nearest 0.1 mm.The external measurements were taken from the specimen's tags.

Results
Most of the measures of the specimen ICN 21985 (Tables 1, 2) coincide with the ranges of measurements previously reported for T. devivoi (Gregorin et al. 2006, Velazco et al. 2014).Only the length of tail, the condyloincisive length measured to the incisors tips (sensu Gregorin et al. 2006), and zygomatic breadth are slightly bigger in our specimen (11.3 mm, 0.3 mm, 0.1 mm, respectively), but it does not represent a great variation more than individual.Regarding other species of the genus, T. devivoi overlapped in most measurements with T. tricolor, but is slightly greater that T. discifera and T. wynneae Velazco, Gregorin, Voss & Simmons, 2014; and smaller that T. lavali (Gregorin et al. 2006;Velazco et al. 2014; Tables 1, 2).
The specimen ICN 21985 has all diagnostic characters recognized for T. devivoi (Gregorin et al. 2006, Velazco et al. 2014): Externally, it has the ventral hair with 2 bands; the basal band is brown and the distal band is grayish brown, giving a frosted appearance.The adhesive disks of the wings are oblong, and the calcar has no lappets.The other 2 species of Thyroptera confirmed from Colombia, T. discifera and T. tricolor (see below), are easily differentiated externally from T. devivoi because they have unicolored ventral hair.T. discifera has light color dorsum and T. tricolor has whitish ventral hairs.Additionally, both have 2 lappets in the calcar and the adhesive disks are circular (Fig. 3).Cranially, T. devivoi has the rostrum subequal to the braincase, the foramen oval is large (small in T. discifera), and the mesopterygoid fossa is divided in 2 cavities by a midventral blade crest (undivided in T. discifera) (Fig 4).In dental characters, T. devivoi differs because the cusps of the I2 are perpendicularly arranged relative to the long axis of the toothrow, whereas the cusps of I2 are positioned obliquely in T. tricolor and T. discifera (Figs 4,5).In the lower teeth, T. devivoi has the i3 taller and faintly tricuspid than i1 and i2, while in T. tricolor and T. discifera the i3 is less developed.
The other 2 species of Thyroptera, T. lavali and T. wynneae, are not present in Colombia.Both have oblong adhesive disks in the wings as T. devivoi, but they are easily differentiated.T. lavali differs from T. devivoi by Table 1.Range of external and skull measurements (mm) and body mass (g) of Thyroptera devivoi based on all previous records summarized by Gregorin et al. (2006).† To the most anterior point of premaxilla.‡ To the incisors tips.MPUJ-MAMM 1216, has been reported as evidence of the presence of T. lavali in Colombian Andes (Reyes-Amaya et al. 2016).However, different traits of this specimen support its identification as T. tricolor (Solari et al. 2004, Velazco et al. 2014).We considered that the external measurements from the label were taken using a distinct protocol than that used by us.All skull measurements of the specimen MPUJ-MAMM 1216 (Table 3), except for condyloincisive length (CIL), coincide with the previously reported for T. tricolor (Velazco et al. 2014).The greatest value of CIL in the specimen could represent intraspecific variation, considering the specimen has all morphological diagnostic characters of T. tricolor: externally, the specimen has white ventral pelage, circular suction disk of the wing, and 2 lappets in the calcar (Fig. 6).The skull has the rostrum tilted up and the mesopterygoid fossa is divided in 2 cavities by a midventral blade crest.Dental characters included a cusp in the second upper incisive obliquely arranged relative to the long axis of the toothrow (Velazco et al. 2014).
One of the specimens examined for comparisons,

Discussion
The new record of T. devivoi in Colombian Llanos extends the distribution of the species approximately 1250 km northwest from the nearest locality in Guyana, and west to the Orinoco river.This wider distribution of the species has been inferred by the long distance between localities (Gregorin et al. 2006, Velazco et al. 2014).Previous known localities for the species are in areas dominated by savanna formations, and the capture sites were associated with humid forest and palm swamps along rivers and streams (Gregorin et al. 2006).Likewise, other species of Thyroptera have been sampled adjacent to palm swamps of Mauritia in different types of lowland ecosystems as tropical wet forest (Velazco et al. 2014).These formations can be very important habitats for the Table 3. Range of external and skull measurements (mm) and body mass (g) of Thyroptera tricolor based on all previous records summarized by Velazco et al. (2014).conservation of the thyropterid bats.
The distribution of T. devivoi is wide and disjunctive, includes 3 ecosystems of savanna bioma: the Cerrado, the Rupunini savannas, and the aeolian seasonally flooded savannas of the Orinoco river basin, all separated by a gap of wet forest.Yet, the species is scarcely represented in biological collections possibly due to the elusive nature of the thyropterid bats (Velazco et al. 2014).
Two species of Thyropterid bats have been reported for the Orinoco river basin, T. lavali and T. tricolor (Ferrer Pérez et al. 2009).For the Llanos region, only T. lavali was recorded, but we do not know the voucher specimens for these records and the specimen from the supporting reference is not available for revision.In our knowledge, no Colombian specimens exist of this species from the Llanos.On the other hand, Linares and Rivas (2003) reported T. lavali for the Orinoco Delta region well before the description of T. devivoi; as these 2 species may be externally confused, a proper examination of this material is ideal to clarify their occurrence.Hence, we consider the distribution of T. lavali uncertain for the llanos of the Orinoco river basin and possibly the only species of Thyroptera in this region is T. devivoi.
Finally, we discard the presence of T. lavali in Colombian Andes and we agree with Velazco et al. (2014) about the uncertainty identity of the Thyroptera fossils from La Venta.Any of the morphological diagnostic characters for the actual 5 species of Thyroptera are distinguishable in the fossil material of La Venta (Czaplewski 1996(Czaplewski , 1997)); therefore, the actual species assignments of this material is not possible.We recommend the use of T. robusta Czaplewski, 1996 for this material because it is the name under which this fossil taxon was described.Therefore, we considered the presence of T. lavali in Colombia uncertain.

Figure 1 .
Figure 1.Panoramic view of the Palm swamp of Mauritia flexuosa (locally known as Morichales) associated to a strip of gallery forest where the specimen ICN 21985 of Thyroptera devivoi was captured.Photography credit: María C. Calderón-Capote.

Figure 3 .
Figure 3. Comparative figure of Thyroptera discifera, Thyroptera devivoi, and Thyroptera tricolor.These individuals were included in the specimens reviewed for comparisons.A. T. discifera, (ICN 21207).B. T. devivoi, (ICN 21985).C. T. tricolor (ICN temp D3M 400).D. Adhesive disk of wings in T. discifera.E. Adhesive disk of wings in T. devivoi.F. Adhesive disk of wings in T. tricolor.T. devioi has the ventral hair bicolored with frosted appearance (B) while T. discifera is unicolored with the light color of dorsum (A) and T. tricolor is unicolored with the ventral hairs whitish (C).T. devivoi does not have lappets in the calcar (E) while T. discifera and T. tricolor have two lappets in the calcar (A and C, see white arrows).T. devivoi has adhesive disks of wings oblong (E), while these are circular in T. discifera and T. tricolor (D and F).Photography credit: T. discifera (A and D) Miguel E. Rodríguez-Posada, T. devivoi (B and E) María C. Calderón-Capote, and T. tricolor (C and F) Darwin M. Morales-Martínez.

Figure 6 .
Figure 6.Ventral view of the specimen MPUJ-MAMM 1216 shown the external characters for Thyroptera tricolor: Ventral hairs are unicolored with whitish coloration.Adhesive disks of wing are circular (upper box).Calcar has 2 lappets (white arrows in the lower box).

Figure 5 .
Figure 5. Occlusal views of the upper dentition.A. Thyroptera devivoi (ICN 21985; male).B. Thyroptera discifera (ICN 21207; male).The cusps of I2 in T. devivoi are perpendicularly arranged relative to the long axis of the toothrow, whereas, in T. discifera and T. tricolor (no drawing) the cusp of I2 are positioned obliquely to the long axis of the toothrow.Scale bars = 2 mm.Illustration by Valentina Nieto Fernández.