Lonchophylla inexpectata, Moratelli, Ricardo & Dias, Daniela, 2015

Moratelli, Ricardo & Dias, Daniela, 2015, A new species of nectar-feeding bat, genus Lonchophylla, from the Caatinga of Brazil (Chiroptera, Phyllostomidae), ZooKeys 514, pp. 73-91 : 76-82

Taxon classification Animalia Chiroptera Phyllostomidae

Lonchophylla inexpectata sp. n. Figures 1, 2, 4, 5; Table 1

Lonchophylla mordax : Lima 1926: 76; not Lonchophylla mordax Thomas, 1903.

Lonchophylla mordax : Vieira 1942: 321; not Lonchophylla mordax Thomas, 1903.

Lonchophylla mordax : Taddei, Vizotto and Sazima 1983; not Lonchophylla mordax Thomas, 1903.

Lonchophylla dekeyseri : Woodman and Timm 2006: 450; part, not Lonchophylla dekeyseri Taddei, Vizotto & Sazima, 1983.

Lonchophylla mordax : Woodman and Timm 2006: 475; part, not Lonchophylla mordax Thomas, 1903.

Lonchophylla dekeyseri : Woodman 2007. Part, not Lonchophylla dekeyseri Taddei, Vizotto & Sazima, 1983.

Holotype.

An adult male, USNM 238008, with skin and skull (Figures 1, 2), including mandible, collected by E. Garbe at Barra (12°42'S, 41°33'W), Bahia, Brazil, on January 1908. Skull and mandible are in good condition except for the minimally damaged anteriormost portion of the foramen magnum. The body is prepared as dry skin. Woodman and Timm (2006: 450) described USNM 238008 as a faded skin, but after comparison of its pelage colour with those from other specimens, only mem branes seem to be faded. External and craniodental measurements for the holotype and paratypes are in Table 1.

Paratypes.

The paratype series comprises 46 vouchers. Three paratypes are from the type locality in Barra, Bahia (AMNH 235608, FMNH 21077, 21078), and were collected by R. H. Becker in 1914. One is from Serra do Catimbau, Buíque, Pernambuco (FMNH 137414; 08°37'S, 37°09'W [coordinates for Catimbau National Park]), and was collected by D. Guerra in 1970. Thirty-eight vouchers are from 17 km south of Exu, Pernambuco (CM 99413-99450; 07°41'S, 39°32'W), elevation ca. 480 m, and were collected by M. R. Willig in 1976. Paratypes from Barra (AMNH 235608, FMNH 21077, 21078), and Buíque (FMNH 137414) are in spirits, others are prepared as dry skin.

Other specimen.

One additional specimen (ALP 3686) from the Caatinga of Andaraí, Bahia may represent Lonchophylla inexpectata . The specimen is preserved in spirit, and the dentition is partially worn, preventing its unambiguous identification.

Distribution.

Lonchophylla inexpectata occurs in the Caatinga of North-eastern (NE) Brazil, with confirmed records from Pernambuco (NE), and Bahia (NE) (Figure 3).

Diagnosis.

Lonchophylla inexpectata can be distinguished from all South American species that occur east of the Andes by the following set of traits: presence of a lingual cusp in the P4, absence of a lingual cusp in the P3, absence of a deep longitudinal groove in the posterior face of the upper canine, proximal portion of the dorsal surface of the forearm not furred, and ventral fur pale.

Description and comparisons.

Like other Lonchophylla , the dental formula of Lonchophylla inexpectata is 2/2, 1/1, 2/3, 3/3 = 34. Lonchophylla inexpectata , Lonchophylla dekeyseri and Lonchophylla bokermanni are the three pale-venter Brazilian species of the genus, whereas Lonchophylla mordax and Lonchophylla peracchii have pale-brown ventral pelage. We did not find evidence of Lonchophylla bokermanni and Lonchophylla peracchii in sympatry with Lonchophylla inexpectata - Lonchophylla bokermanni is restricted to a small area in the Serra do Espinhaço, Cerrado of Minas Gerais; and Lonchophylla peracchii occurs in the Atlantic Forest, from Espírito Santo southward to São Paulo. Lonchophylla inexpectata can be distinguished from these two species by the presence of a well-developed lingual cusp in the P4, with lingual root in the median portion of the tooth; absence of a groove along the anterior surface of the upper canines; and proximal portion of the dorsal surface of the forearm not covered with fur.

Based on the samples we have available, Lonchophylla inexpectata resembles Lonchophylla dekeyseri in the pale ventral fur, and Lonchophylla mordax in the dental morphology. These three species overlap partially in external and cranial size, but in general, cranial measurements for Lonchophylla inexpectata average significantly larger than those for Lonchophylla dekeyseri and smaller than those for Lonchophylla mordax (Table 1).

Lonchophylla mordax has been reported in the literature as a pale-venter species (e.g., Lima 1926, Vieira 1942, Taddei et al. 1983, Nogueira et al. 2007), and subsequent to the description of Lonchophylla dekeyseri , these taxa have been considered the two pale-venter species from NE Brazil (see Taddei et al. 1983, Nogueira et al. 2007, Dias et al. 2013). However, after examining part of the type series of Lonchophylla mordax (BM 1903.9.5.34 [holotype], USNM 123392 [paratype]), along with one other specimen from the same locality of the type series (MHNG 667.13 [identified as Lonchophylla mordax by Thomas]), and samples from a nearby locality having similar habitat (Itabaiana, Sergipe)-whose external and skull morphology fit with those of the type series of Lonchophylla mordax (ALP 8768-8770, 8812-8819)-we concluded that Lonchophylla mordax has a light-brown ventral pelage, which is consistently darker than the paler ventral pelage of the type material of Lonchophylla dekeyseri and other samples of this species. The ventral pelage of specimens from Barra, Bahia ( Lonchophylla inexpectata ) is similar to that of Lonchophylla dekeyseri . Under "historical remarks" we discuss the reasons for previous assignments of pale-venter samples from the Caatinga of NE Brazil (= Lonchophylla inexpectata ) to Lonchophylla mordax .

Lonchophylla inexpectata averages significantly smaller than Lonchophylla mordax in all cranial dimensions except in POB and BCB (Table 1, Figure 4). This is particularly notable in the length of the mandible (MAL x¯ = 15.6 mm, range [R] = 14.1-16.3 mm [ inexpectata ] versus x¯ = 16.1 mm, R = 15.5-17.0 mm [ mordax ]). Lonchophylla inexpectata can also be distinguished by the ventral pelage, which varies from whitish (e.g., USNM 238008, CM 99415) to pale greyish (near Avelaneous [e.g., CM 99432, 99437]), but near Buffy Brown in Lonchophylla mordax (e.g., BM 1903.9.5.34, USNM 123392). The throat and the posterior region of the belly are consistently paler, tending to whitish, in Lonchophylla inexpectata (Figure 5).

Lonchophylla inexpectata resembles Lonchophylla dekeyseri in the pelage colour, but these species can be distinguished by qualitative and quantitative cranial characteristics. Lonchophylla inexpectata is significantly larger than Lonchophylla dekeyseri in all length measurements of skull and rostrum (GLS, CIL, BAL, MTL, M1M3, MAL, MAN), but Lonchophylla dekeyseri averages slightly larger in those measurements of the width of skull and rostrum (BAC, POB, BCB, MAB), indicating a longer but narrower skull in Lonchophylla inexpectata (Table 1). Lonchophylla inexpectata can be distinguished from Lonchophylla dekeyseri by the narrower first upper premolar (P3) in occlusal view, with lingual lobe absent or obsolete (in contrast with the usually more robust P3, which has a small or moderately developed inner lobe in dekeyseri [Figure 6]); absence of a deep longitudinal groove in the posterior surface of the canine; narrower and uninflated rostrum, with more widely projecting lacrimals (wider and more inflated rostrum, and lacrimal region almost indistinguishable in dekeyseri ); upper molars (M1 and M2) with low crowns in lateral view (molars with higher crowns in dekeyseri ); parastyle of M1 projecting labially over the posterior labial margin of the last upper premolar (P4); mesostyle of M1 shorter; metastyle of M1 well developed (reduced or absent in dekeyseri [Figure 6]); parastyle of M2 well developed but slender (well developed and more rounded in dekeyseri ); mesostyle of M2 shorter; metastyle of M2 distinct, moderate or well developed (reduced or absent in dekeyseri ).

Multivariate analysis.

To test the results obtained from the morphological analyses, we performed a discriminant function analysis including samples we confidently assigned to Lonchophylla dekeyseri (three groups from the Cerrado of Mid-western Brazil), Lonchophylla inexpectata (two groups from the Caatinga of NE Brazil), and Lonchophylla mordax (one group from the Caatinga of NE Brazil, and one group from the Atlantic Forest–Caatinga ecotone in NE Brazil). The first two discriminant functions (DF1, DF2) summarized 47% and 40% of the total variation, respectively (Table 2). All samples grouped as expected, confirming the cohesive pattern retrieved from the morphological analysis. Centroids for samples assigned to Lonchophylla inexpectata were distinct from those of Lonchophylla dekeyseri and Lonchophylla mordax across the first two axes, and only a few scores of Lonchophylla inexpectata are within the dispersal cloud of Lonchophylla mordax (Figure 7). The three species overlap partially across the first axis, but Lonchophylla inexpectata distinguishes from Lonchophylla dekeyseri and Lonchophylla mordax along the second axis. Scores for Lonchophylla inexpectata had very low positive to high negative values along the DF2, whereas those for Lonchophylla dekeyseri and Lonchophylla mordax have low negative to high positive values along this axis.

Etymology.

The name “inexpectata” is Latin for “unexpected”, in allusion to the unexpected (at least for the authors) new taxonomic status of pale-venter populations of Lonchophylla from the Caatinga of North-eastern Brazil.