Zootaxa 3722 (3): 347–360 ISSN 1175-5326 (print edition)

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Copyright © 2013 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3722.3.4
http://zoobank.org/urn:lsid:zoobank.org:pub:FE19D2AF-BBC7-464A-B440-E3C8B0CFC4BB

A new species of Lonchophylla (Chiroptera, Phyllostomidae) from the Atlantic

Forest of southeastern Brazil, with comments on L. bokermanni

DANIELA DIAS1, CARLOS EDUARDO L. ESBÉRARD2 & RICARDO MORATELLI3,4

1
Instituto Oswaldo Cruz, Fundação Oswaldo Cruz, Rio de Janeiro, Brazil. E-mail: dani_dias262@yahoo.com.br
2
Instituto de Biologia, Universidade Federal Rural do Rio de Janeiro, Seropédica, Brazil. E-mail: cesberard@superig.com.br
3
Campus Fiocruz da Mata Atlântica, Fundação Oswaldo Cruz, Rio de Janeiro, Brazil. E-mail: rmoratelli@fiocruz.br
4
Department of Vertebrate Zoology, National Museum of Natural History, Washington D.C., USA

Abstract

We examined Brazilian species of the nectar-feeding bats genus Lonchophylla (Phyllostomidae, Lonchophyllinae) to clar-
ify the identity of Lonchophylla bokermanni and to determine the distribution of this and other species of Lonchophylla
in eastern Brazil. As a result, we have found sufficient differences between Cerrado populations (including the type local-
ity of L. bokermanni) and populations inhabiting the Atlantic Forest of southeastern Brazil,which warrant the treatment
of the Atlantic Forest populations as a separate and new species. We describe this new species here as Lonchophylla per-
acchii, sp. nov. The new species appears to be restricted to the Atlantic Forest, whereas L. bokermanni is found only in
Cerrado habitats.

Key words: Atlantic Rainforest, Cerrado, morphology, morphometrics, South America

Introduction

The Lonchophyllinae (Phyllostomidae) comprises a complex of Neotropical nectar-feeding bats characterized by

elongated skulls with long muzzles and long, extensible tongues that lack a paintbrush tip. Each side of the tongue
has a deep longitudinal groove bordered by short hair-like papillae (Griffiths 1982; Griffiths & Gardner 2008). The
four genera currently recognized are: Lonchophylla Thomas, Lionycteris Thomas, Platalina Thomas, and
Xeronycteris Gregorin and Ditchfield. Molecular analyses have recovered Lonchophylla as paraphyletic (e.g.,
Dávalos & Jansa 2004; Datzmann et al. 2010; Dávalos et al. 2012); but as currently understood, this genus
comprises small to medium-sized species having unreduced molar cusps in contrast with the dentitions of other
lonchophyllines. Other features used to distinguish Lonchophylla from the other three genera include the bases of
dorsal fur paler than tips; absence of conspicuous fur on uropatagium; narrow, anteroposteriorly elongated
premolars, and conspicuously enlarged, procumbent upper inner incisors (Griffiths & Gardner 2008).
Lonchophylla has been the subject of several recent taxonomic studies, with 9 of the 14 valid species described
since 1978, and 6 of these since 2004 (Dávalos 2004; Albuja & Gardner 2005; Woodman & Timm 2006; Woodman
2007; Dávalos & Corthals 2008; Griffiths & Gardner 2008). Four species are known from Brazil (Peracchi et al.
2011; Paglia et al. 2012). Lonchophylla bokermanni Sazima, Vizotto & Taddei, 1978 and L. mordax Thomas, 1903
are endemic; the former recorded in the Cerrado and Atlantic Forest, and the latter in all major habitat zones along
the eastern coast. Lonchophylla dekeyseri Taddei, Vizotto & Sazima, 1983 has been found in low numbers in the
Cerrado of midwestern and northeastern Brazil, and in eastern Bolivia (Emmons et al. 2006; Griffiths & Gardner
2008; Aguiar et al. 2010). Lonchophylla thomasi J. A. Allen, 1904 is widely distributed from northern Bolivia and
adjacent Brazil, northward into Panama (Simmons 2005; Griffiths & Gardner 2008). All recent taxonomic
assessments have focused on the other 10 known species, which are distributed from Peru northward along or
adjacent to the Andes into Central America (Dávalos 2004; Albuja & Gardner 2005; Woodman & Timm 2006;
Woodman 2007; Dávalos & Corthals 2008; Griffiths & Gardner 2008).

Accepted by M. Weksler: 1 Oct. 2013; published: 23 Oct. 2013 347

 The primary focus of this report is the taxonomic status of populations identified as L. bokermanni.
Distribution records list the species from a few localities in the Cerrado of the state of Minas Gerais and in the
Atlantic Forest of the states of Rio de Janeiro and Espírito Santo. The species was described from Serra do Cipó,
eastern Minas Gerais (Sazima et al. 1978). Additional specimens have been taken near the type locality (Sazima et
al. 1989; L. Geise & C. E. L. Esbérard, unpublished). Specimens identified as L. bokermanni from the Atlantic
Forest of Rio de Janeiro were first reported from Ilha Grande (Taddei et al. 1988), with subsequent records from
other continental islands (Esbérard et al. 2006; Esbérard 2009; Lourenço et al. 2010) and on the mainland from
lowland and montane evergreen forests (Dias et al. 2002; Moratelli & Peracchi 2007; Dias & Peracchi 2008;
Novaes et al. 2010), semideciduous forests (Baptista & Mello 2001), and various second growth forest habitats (C.
E. L. Esbérard, unpublished). A specimen was reported recently from Reserva Biológica de Sooretama, a lowland
Atlantic Forest remnant in Espírito Santo (Pimenta et al. 2010). Specimens identified as L. bokermanni (Baptista &
Oliveira 1998; Griffiths & Gardner 2008) from the state of Bahia are misidentified Xeronycteris vieirai Gregorin &
Ditchfield, 2005 (Gregorin & Ditchfield 2005; Tavares et al. 2008). Other specimens from the Atlantic Forest of
Rio de Janeiro were tentatively assigned to L. mordax (Dias et al. 2002; Esbérard et al. 2006), but reidentified as L.
bokermanni (Dias & Peracchi 2008). We have examined the specimens identified as L. bokermanni from the
Atlantic Forest of Rio de Janeiro and compared them with individuals from the Cerrado of Minas Gerais and find
that they are smaller in size and differ in other characteristics. Herein, we describe a new species from the Atlantic
Forest of southeastern Brazil and provide taxonomic comments on L. bokermanni.

Material and methods

Analyzed specimens are deposited in the collections of the Universidade Estadual Paulista “Júlio de Mesquita
Filho”, São José do Rio Preto, Brazil (DZSJRP); Universidade Federal Rural do Rio de Janeiro (ALP and LMD);
and Museu Nacional, Universidade Federal do Rio de Janeiro (MN), Rio de Janeiro, Brazil (Appendix). Dental
nomenclature of tooth morphology follows Phillips (1971).
We examined 113 specimens of which 2 represent L. dekeyseri (the holotype and a paratype), 32 are L. mordax,
and 79 have been identified as L. bokermanni. Of the latter, 8 are from the Cerrado of Minas Gerais (including 6
from the type series) and 71 from the Atlantic Forest of Rio de Janeiro.
Twelve cranial and one external dimensions (following Sazima et al. 1978) were measured on 77 adults as
determined by epiphyseal fusion of metacarpals and phalanges, and complete closure of basicranial sutures
(Appendix). Measurements (in mm) were taken using a caliper accurate to 0.02 mm as follows: greatest length of
skull (GLS), from the posteriormost point of the occiput to the tip of the upper inner incisors; condylo-incisive
length (CIL), from the line connecting the occipital condyles to the tip of the upper inner incisors; basal length
(BAL), from the anterior margin of the foramen magnum to the tip of the upper inner incisors; palatal length (PAL),
from the posterior rim of the alveolus of the inner incisors to the posterior margin of the bony palate; maxillary
toothrow length (MTL), from the anterior surface of the upper canine, including the cingulum, to the posterior
surface of M3; breadth across canines (BAC), greatest breadth across outer surface crowns of upper canines
including cingulae; breadth across molars (BAM), greatest breadth across outer edges of the crowns of upper
molars; postorbital breadth (POB), least breadth across frontals posterior to the postorbital bulges; braincase
breadth (BCB), greatest breadth of the globular part of the braincase; mastoid breadth (MAB), greatest breadth
across the mastoid region; mandibular length (MAL), from the mandibular symphysis to the condyloid process;
mandibular toothrow length (MAN), from the anterior crown of the lower canine, including cingulae, to the
posterior crown of m3; and forearm length (FA), from the elbow to the distal end of the forearm including carpals,
measured with the wing partially folded. Descriptive statistics (mean, range, and standard deviation) were
calculated for all dimensions.
Principal Component and Discriminant Function Analyses (PCA and DFA, respectively) were used to
summarize patterns of size and shape variation among samples. As multivariate procedures require complete
datasets, missing values (1% of total dataset) were estimated from the existing raw data using the Expectation-
maximization (E-M) algorithm (Little & Rubin 1987; Strauss et al. 2003). Measurements were log-transformed
and the covariance matrices were computed considering all variables. Mahalanobis distances between samples
were portrayed in an UPGMA (unweighted pair-group method using arithmetic averages) dendrogram. The

348 · Zootaxa 3722 (3) © 2013 Magnolia Press DIAS ET AL.

statistical significance of differences among samples was assessed by single and multivariate analyses of variance
(ANOVA and MANOVA, respectively). Mantel’s test was used to assess isolation-by-distance comparing
geographic and morphological distance matrices (Mantel 1967). Statistical procedures were performed in Matlab
(The MathWorks, Inc., Natick, Massachusetts) using functions written by R. Strauss (Strauss 2012).
Samples included in the morphometric analyses are as follows (Figure 1): group 1: Semideciduous seasonal
forest (Cambuci); group 2: Vegetation mosaic within the metropolitan region of Rio de Janeiro (Floresta da Tijuca,
Parque Estadual da Pedra Branca, Reserva do Grajaú); group 3: early successional second growth (Morro de São
João); group 4: Vegetation mosaic in the montane region of Rio de Janeiro (Reserva Biológica do Tinguá); group
5: Costa Verde (Ilha da Gipóia, Ilha de Itacuruça, Ilha Grande, Parati, Reserva de Rio das Pedras, Estrada Rio-
Santos, Vale do Rio Sahy); group 6: Cerrado of southeastern Brazil (Serra do Cipó, Diamantina); group 7:
Caatinga/Atlantic Forest ecotone (Itabaiana); group 8: Caatinga (Parque Natural Grota do Angico); and group 9:
Cerrado of midwestern and northeastern Brazil (Parque Nacional de Brasília, Distrito Federal, and Piracuruca,
Piauí). These samples can be assigned to the following morphoclimatic domains—and whenever appropriate, they
will be referred to as such: Atlantic Forest of southeastern Brazil (groups 1–5 [previously identified as L.
bokermanni]); Cerrado of southeastern Brazil (group 6 [includes holotype and paratypes of L. bokermanni]);
Caatinga of northeastern Brazil (group 7 [L. mordax]); Caatinga/Atlantic Forest ecotone of northeastern Brazil
(group 8 [L. mordax]); and Cerrado of midwestern and northeastern Brazil (group 9 [holotype and paratype of L.
dekeyseri]).

FIGURE 1. Map of part of southeastern Brazil showing the geographic distribution of L. peracchii sp. n. in the Atlantic Forest
of Rio de Janeiro (groups 1–5), and L. bokermanni in the Cerrado of Minas Gerais (group 6). Numbers between parentheses
correspond to groups used in the morphometric analyses.

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Results

Qualitative analyses. Specimens from Atlantic Forest (groups 1–5) and Cerrado of southeastern Brazil (group 6)
differ from the remaining samples (groups 7–9) by the proximal portion of the forearm covered with fur;
basisphenoid pits shallow and with broad intervening septum; posteromedial edge of the palate set posteriorly to
posterior border of the optic foramen; pterygoid processes narrow and divergent; upper canines long and distinctly
grooved along the anterior surface; P4 narrow in occlusal view, with inner lobe reduced and lingual root posteriorly
displaced; and coronoid process low, with rounded tip only slightly above the articular condyle.
Considering only specimens tentatively assigned to L. bokermanni (groups 1–6), morphological comparisons
reveal that specimens from Atlantic Forest (groups 1–5) can be distinguished from those from Cerrado (group 6) on
the basis of pelage color, and external and cranial traits. Atlantic Forest specimens have medium-sized ear, with
narrow tip; spatulate tragus with rounded tip; pale-brownish ventral pelage from neck to genital region, with slight
contrast between the medium-brown bases and the pale-brown tips (Figure 2); and ventral and dorsal pelages
contrasting slightly. On the other hand, Cerrado specimens have short ear with more rounded tip; narrow tragus
with pointed tip; pale-grayish ventral pelage from neck to genital region, with strong contrast between the dark-
brown bases and the pale-gray tips (Figure 2; but white tips in MN 79996, 79997); and ventral and dorsal pelage
contrasting strongly. In addition, Atlantic Forest specimens have larger and more elongated noseleaf spear (Atlantic
Forest [N = 21]—length: 6.5–7.5 mm, width: 4.0–5.0 mm; Cerrado [N = 5]—length: 6.0–6.5 mm, width: 3.5–4.5
mm [measurements including the horseshoe]), with indistinct central rib extending up to the tip (contrasting with
the narrower and shorter noseleaf spear, with central rib restricted to the basis in Cerrado specimens); medium-
brown wing membranes (contrasting with almost blackish wing membranes); and shorter interfemoral membrane,
not reaching the ankle (contrasting with elongated interfemural membrane, reaching the ankle).

FIGURE 2. Ventral pelage and tragus of L. peracchii sp. n. (A; ALP 6557), and L. bokermanni (B; MN 79996).

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FIGURE 3. Dorsal, ventral, and lateral views of cranium and lateral view of mandible of L. peracchii sp. n. (A–D; DZSJRP
15162), and L. bokermanni (E–H; DZSJRP 10347). Scale bar: 5.0 mm.

FIGURE 4. Oblique occlusal view of the upper molars of L peracchii (A; DZSJRP 15162), and L. bokermanni (B; DZSJRP
11410). Cusps numbered in the upper second molar are the parastyle (1), mesostyle (2) and metastyle (3). Note the more
developed cusps in L. bokermanni in comparison with L. peracchii sp. n. Maxillary toothrow length ca. 8.0 mm.

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 On the basis of the cranial morphology (Figure 3), Atlantic Forest specimens (groups 1–5) have narrower
rostrum, and not inflated supraorbital region (contrasting with an inflated region in Cerrado specimens [group 6]);
less inflated nasal bones (contrasting with more inflated and conspicuous nasal bones); and narrower, and shorter
braincase (contrasting with a larger, higher, and more globular braincase).
Analyses of tooth morphology reveal differences in the parastyle, mesostyle, and metastyles of the 1st and/or
2nd upper molars (M1 and M2—Figure 4). In Atlantic Forest specimens (groups 1–5) the parastyle of M1 is poorly
developed, and labially oriented in occlusal view; the parastyle of M2 is poorly to moderately developed; the
mesostyles are absent or reduced in both M1 and M2; and the metastyles of both M1 and M2 are poorly developed.
In contrast, in Cerrado specimens (group 6) the parastyle of M1 is well developed, and labially and more anteriorly
oriented in occlusal view; the parastyle of M2, and the mesostyles and metastyles of both M1 and M2 are well
developed (but not visible in the holotype [DZSJRP 10347] because cusps are worn).
Quantitative analyses. The first principal component (PC1) in the morphometric analyses accounted for 61%
of the total variation, and represents a general size axis based on character loadings (Figures 5a and 5b, Table 1).
PC1 scores show that specimens from Cerrado of southeastern Brazil (group 6) are larger than specimens from
Atlantic Forest (groups 1–5), followed by specimens from Caatinga/Atlantic Forest ecotone and Caatinga (groups 7
and 8), with specimens from Cerrado of midwestern and northeastern Brazil being the smaller ones (group 9—
Figure 5a). These results reveal a gradient in size with L. bokermanni (group 6) from Cerrado larger than L.
bokermanni from Atlantic Forest (groups 1–5), followed by L. mordax (group 7, 8), and L. dekeyseri (group 9).
Although ranges of variation overlap in cranial measurements—but not in forearm length—for samples tentatively
assigned to L. bokermanni, specimens from Atlantic Forest are significantly smaller than those from Cerrado,
except in the palatal length, the breadth across canines, and the braincase breadth (Table 2).

TABLE 1. Vector correlation coefficients (loadings) between original variables and principal components (PC1 and
PC2) and between original variables and discriminant functions (DF1 and DF2) for samples of Lonchophylla from
Atlantic Forest and Cerrado of southeastern Brazil, Caatinga and Caatinga/Atlantic Forest ecotone of northeastern Brazil,
and Cerrado of midwestern and northeastern Brazil. Numbers in bold indicate vector correlations with magnitudes >
±0.29.
Loadings of PCA and DFA
Characters PC1 PC2 DF1 DF2
FA 0.57 0.50 0.02 0.50
GLS 0.96 - 0.01 - 0.24 0.60
CIL 0.95 - 0.07 0.78 - 0.56
BAL 0.95 - 0.11 - 0.09 - 0.02
MTL 0.29 0.64 - 0.09 0.08
BAM 0.14 0.74 - 0.16 0.06
BAC 0.66 0.07 0.11 0.01
POB 0.88 - 0.31 0.12 0.06
BCB 0.64 0.08 0.21 - 0.19
MAB 0.38 0.52 - 0.42 0.10
MAL 0.91 0.12 0.05 0.05
MAN 0.39 0.57 -0.18 - 0.12

The first two discriminant functions (DF1 and DF2) summarized 82% of the total variation. Along both axes
three clusters were revealed (Figures 5c, and 6): cluster 1—samples from Atlantic Forest (groups 1–5); cluster 2—
samples from Cerrado of southeastern Brazil (group 6; including type specimens of L. bokermanni); and cluster
3—samples from Caatinga/Atlantic Forest ecotone, Caatinga (groups 7 and 8; both assigned to L. mordax), and
from Cerrado of midwestern and northeastern Brazil (group 9; type specimens of L. dekeyseri). These clusters were
significantly distinct from each other (MANOVA: Wilks’lambda = 0.001, F = 7.29, p < 0.0001). Samples from
Atlantic Forest can be distinguished from Cerrado of southeastern Brazil by negative values along DF2, and from
Caatinga, Caatinga/Atlantic Forest ecotone, and Cerrado of midwestern and northeastern Brazil by positive values

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along DF1 (Figure 5c). The contrast between the forearm length (FA), and the condylo-incisive length (CIL) was
associated with the distinction of samples assigned to L. bokermanni (Figures 5c and 5d); and the ratio between
these measurements (CIL/FA) can be used to distinguish Atlantic Forest (0.64–0.70) and Cerrado (0.59–0.61)
samples. The ratio between the greatest skull length and the forearm length—two measurements revealed
important in the distinction of samples—also revealed to be useful (GLS/FA: 0.65–0.73 [Atlantic Forest]; 0.62–
0.64 [Cerrado]). Considering only samples tentatively assigned to L. bokermanni (groups 1–6), those from Altantic
Forest (groups 1–5) were significantly distinct from Cerrado (group 6—MANOVA: Wilks’ lambda = 0.0192, F =
2.76, p < 0.0001); a result not explained by their geographic distance (Mantel’s test: r = 0.83; p = 0.11). Results of
DFA support the distinction of Atlantic Forest and Cerrado samples assigned to L. bokermanni, as well as their
distinction from both L. mordax and L. dekeyseri. On the other hand, samples assigned to L. mordax and L.
dekeyseri revealed to be morphometrically similar.

FIGURE 5. Plots of multivariate individual scores in the first two principal components (A), and in the first two discriminant
functions (C). Corresponding vector correlations (> ±0.29) of craniometric characters with the first two eigenvectors of the
principal components (B), and of the discriminant functions (D). Groups 1 to 5 (= Lonchophylla peracchii sp. n.) are from the
Atlantic Forest of southeastern Brazil; group 6 (L. bokermanni) is from the Cerrado of southeastern Brazil; groups 7 and 8 (L.
mordax) are from the Caatinga, and Caatinga/Atlantic Forest ecotone of northeastern Brazil; and group 9 (L. dekeyseri) is from
the Cerrado of midwestern and northeastern Brazil.

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TABLE 2. Selected measurements (mm) of pooled samples from the Atlantic Forest (Lonchophylla peracchii) and
Cerrado (L. bokermanni) of southeastern Brazil. F values (ANOVA) with one asterisk (*) indicate statistical significance
with α = 5% (p ≤ 0.05), and with two asterisks (**) indicate statistical significance with α = 0.1% (p ≤ 0.001).
Atlantic Forest of SE Brazil Cerrado of SE Brazil
Characters Mean (Range) SD N Mean (Range) SD N F
FA 35. 8 (34.5–36.9) 0.67 36 40.2 (39.4–41.1) 0.68 7 253.00**
GLS 24.6 (23.8–25.4) 0.51 36 25.3 (24.9–25. 6) 0.23 7 12.94**
CIL 23.6 (22.7–24. 6) 0.48 36 24.2 (23.8–24.4) 0.22 7 7.92*
BAL 21.6 (20.2–22.7) 0.52 36 22.2 (21.9–22.4) 0.20 7 7.64*
PAL 13.9 (13.0–14.6) 0.41 36 14.2 (13.8–14.5) 0.24 7 3.61
MTL 8.0 (7.5–8.4) 0.22 36 8.3 (7.8–8.6) 0.28 7 8.03*
BAM 5.1 (4.8–5.8) 0.17 36 5.4 (5.1–5.7) 0.20 7 9.75*
BAC 3.9 (3.5–4.1) 0.13 36 3.9 (3.8–4.1) 0.10 7 1.92
POB 4.7 (4.4–5.0) 0.16 36 5.0 (4.8–5.2) 0.12 7 12.41**
BCB 9.1 (8.4–9.6) 0.29 36 9.3 (9.0–9.6) 0.25 7 1.62
MAB 9.3 (8.5–9.7) 0.24 36 9.6 (9.4–9.7) 0.13 7 7.45*
MAL 16.9 (16.0–17.8) 0.43 36 17. 5 (17.3–17.6) 0.10 7 12.50**
MAN 8.4 (8.0–8.8) 0.20 36 8.7 (8.2–9.0) 0.26 7 14.13**

Summary statistics: SD = Standard deviation; N = sample size (adults only). See text for a description of measurement
methods.

FIGURE 6. UPGMA dendrogram based on Mahalanobis centroid distances of samples. Groups 1–5 (= L. peracchii sp. n.):
Atlantic Forest of southeastern Brazil; group 6 (L. bokermanni): Cerrado of southeastern Brazil; group 7 and 8 (L. mordax):
Caatinga and Caatinga/Atlantic Forest ecotone of northeastern Brazil; and group 9 (L. dekeyseri): Cerrado of midwestern and
northeastern Brazil.

Due to the similarity with the holotype we assign the sample from Cerrado of southeastern Brazil to L.
bokermanni, and those from Atlantic Forest to a distinct and unnamed species we describe as:

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Lonchophylla peracchii, sp. nov.
Peracchi’s Nectar Bat
Figures 2, 3, 4, and 7; tables 2, and 3

Lonchophylla bokermanni: Taddei, Souza & Manuzzi, 1988: Part; not Lonchophylla bokermanni Sazima, Vizotto and Taddei, 1978.

Holotype. An adult female, DZSJRP 15162, preserved in alcohol, with skull removed and complete, including
mandible (Figures 3a–d), collected by S. A. de Souza and J. L. Manuzzi (original field number DZUFRJ 62) on 09
December 1984.
Type locality. Near Vila do Abraão (ca. 23°07’ S, 44°10’ W), Ilha Grande, Angra dos Reis, Rio de Janeiro
State, Brazil. The type locality is a continental island located (ca.) 2 km from the continent (Esbérard et al. 2006;
see Figure 1).
Paratypes. The paratypes include two adult females, DZSJRP 15159 and DZSJRP 15163 (original numbers
Feema 93 and Feema 94, respectively), collected in 2 August 1980, in the same locality of the holotype, by S. A. de
Souza e J. L. Manuzzi. Dimensions for the type series are reported in table 3.

TABLE 3. Measurements (mm) of the type series (Ilha Grande, Angra dos Reis, Rio de Janeiro, Brazil) of Lonchophylla
peracchii.
Holotype Paratype Paratype
DZSRP 15162 DZSJRP 15159 DZSJRP 15163
Age/sex Adult female Adult female Adult female
FA 35.8 36.0 36.5
GLS 25.5 24.5 25.0
CIL 24.5 23.5 24.0
BAL 22.4 21.5 21.8
PAL 14.7 13.8 13.8
MTL 8.1 8.0 8.0
BAM 5.0 5.0 5.2
BAC 4.0 3.9 3.9
POB 4.8 4.6 4.7
BCB 9.1 9.0 9.3
MAB 9.3 9.1 9.4
MAL 17.6 16.6 17.0
MAN 8.5 8.4 8.3
See text for a description of measurement methods.

Distribution. Lonchophylla peracchii is currently known from different habitat formations along the Atlantic
Forest of Rio de Janeiro. This species occurs in islands near the continent, lowland and mountainous evergreen
forests, semideciduous stational forests, and pioneer formations. Altitudinal records range from the sea level in the
Costa Verde region, to 900 m in the Serra dos Órgãos National Park. Lonchophylla peracchii apparently occurs in
syntopy with the other nectar-feeding species Anoura caudifer (É. Geoffroy, 1818), A. geoffroyi Gray, 1838, and
Glossophaga soricina (Pallas, 1766). We expect the occurrence of the species in other Atlantic Forest localities in
southeastern Brazil. One specimen recently reported from Espírito Santo (Pimenta et al. 2010) fits with
measurements herein reported to L. peracchii.
Etymology. Lonchophylla peracchii is named after Dr. Adriano Lúcio Peracchi—who first questioned the
distinct taxonomic status of the Atlantic Forest populations of L. bokermanni. We are pleased to dedicate this new
species to him due to his outstanding contributions to the Brazilian chiropterology and acarology, and as the former
supervisor of the three authors (see Peracchi 2010).
Diagnosis. The following set of characters distinguish L. peracchii from all congeners: medium-sized ear with
narrow tip; tragus with rounded tip (not pointed); ventral fur brownish, with slight contrast between hair bases and

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tips; forearm length 37.0 mm or less; mesostyles of M1 and M2 absent or poorly developed; parastyle of M1 poorly
developed, and oriented labially; metastyles of M1 and M2 poorly developed; and ratio of forearm length to
condylo-incisive length 1.57 or less.
Description. Lonchophylla peracchii is a small to medium-sized species (FA: 34.5–36.9 mm; Table 2), with
long and silky pelage; pale-brownish ventral pelage from neck to genital region, with slight contrast between the
medium-brown bases and the pale-brown tips; ventral and dorsal pelages contrasting slightly; medium-brown wing
membranes; pale-brown ears, tragus, noseleaf and uropatagium; elongated and narrow muzzle; medium-sized ears,
with tragus spatulate and rounded at the tip; proximal portion of the dorsal surface of the forearm covered with fur;
large and elongated noseleaf spear, with indistinct central rib extending up to the tip; horseshoe continuous with the
upper lip; and short interfemoral membrane, that not reaches the ankle.
The skull and rostrum are long and narrow, with a postorbital region not inflated and without lateral projections
in dorsal view; supraorbital region and nasals not inflated; posterior border of infraorbital foramen set between the
posterior root of P4 and anterior root of M1 or above the anterior root of M1 in lateral view; mesopterygoid fossa
long, opened, and U-shaped or V-shaped anteriorly; pterygoid processes narrow, divergent and not inflated;
basisphenoid pits shallow, with intervening septum broad; posteromedial edge of the palate positioned posteriorly to
the posterior border of the optic foramen; zygomatic arcs absent; dentary long and slender; articular process long and
slender, but conspicuous; and coronoid process low, with rounded tip slightly above the line of the articular condyle.
Dental formula 2/2, 1/1, 2/3, 3/3, totaling 34 teeth, with inner upper incisors elongated, spatulated, procumbent
and separated at the bases but in contact at the tips; inner and outer upper incisors not in contact; outer upper
incisors small, slender and pointed, and not in contact with canines; upper canines long and distinctly grooved
along the anterior surface, and not in contact with P3 (the first upper premolar); upper premolars triangular,
anteroposteriorly elongated in lateral view, and narrow in occlusal view; P4 with inner lobe reduced, varying from
very low to a small curve, or to a small projection, with a rudimentary cusp, with lingual root displaced posteriorly;
upper molars decreasing in size from M1 to M3; mesostyle absent or reduced in both M1 and M2; parastyle of M1
poorly developed, labially oriented in occlusal view, and not projected over the labial margin of the P4; parastyle of
M2 and metastyles of both M1 and M2 poorly developed; lower incisors trilobed and relatively broad; lower
second premolar (p2) bladelike, with posterior cusp reduced or absent and in contact with canine.

FIGURE 7. Lonchophylla peracchii sp. n. (A) from the Atlantic Forest of Rio de Janeiro, and (B) L. bokermanni from the
Cerrado of Minas Gerais, Brazil.

Comparisons. We directly compare L. peracchii with L. dekeyseri, L. mordax and L. bokermanni due to the
possible sympatry with L. mordax in the Atlantic Forest of southeastern Brazil, possible parapatry with L. dekeyseri
along part of the east coast of Brazil (see Griffiths & Gardner 2008), and cryptic morphology with L. bokermanni.

356 · Zootaxa 3722 (3) © 2013 Magnolia Press DIAS ET AL.

Also, we compare L. peracchii with the remaining South American species—all of them either from the Amazon
basin or from the northwestern South America—based on original descriptions, and subsequent reassessments
(e.g., Dávalos 2004; Albuja & Gardner 2005; Woodman & Timm 2006; Woodman 2007; Dávalos & Corthals 2008;
Griffiths & Gardner 2008).
L. peracchii and L. bokermanni can be distinguished by external, cranial, and dental traits reported above in the
distinction of samples previously assigned to L. bokermanni from Atlantic Forest (subsequently assigned to L.
peracchii), and Cerrado (kept as L. bokermanni). Useful traits to distinguish these species are the forearm length
(FA: 34.5–36.9 mm in peracchii; 39.4–41.1 mm in bokermanni); and the ratios of greatest skull length to forearm
length (GLS/FA: 0.65–0.73 in peracchii, 0.62–0.64 in bokermanni), and condylo-incisive length to forearm length
(CIL/FA 0.64–0.70 in peracchii, 0.59–0.61 in bokermanni).
From L. mordax and L. dekeyseri, L. peracchii can be distinguished by the proximal portion of the dorsal surface
of the forearm covered with fur. These species overlap in the forearm length (FA: 34.5–36.9 mm in peracchii, 32.5–
36.7 mm in mordax, 34.7–37.7 mm in dekeyseri), but in the length of skull L. peracchii is larger than L. dekeyseri
and L. mordax, overlapping with L. mordax, but not with L. dekeyseri (GLS: 23.8–25.4 mm in peracchii, 21.9–22.4
mm in dekeyseri, 22.4–24.2 mm in mordax). It can also be distinguished from L. dekeyseri by the ratio of greatest
skull length to forearm length (GLS/FA: 0.65–0.73 in peracchii, 0.60–0.64 in dekeyseri), and by the longer and
narrower rostrum, which is shorter and inflated in L. dekeyseri. From L. mordax, L. peracchii can also be
distinguished by the last premolar (P4) narrower, with inner lobe varying from very low to a small curve, or to a
small projection, with a rudimentary cusp (in contrast with P4 robust, with inner lobe well developed, and lingual
root in the median portion of the tooth in mordax); basisphenoid pits shallow, with intervening septum relatively
broad (basisphenoid pits deep, separated by a narrow septum in mordax); posteromedial edge of the palate
positioned posteriorly to the posterior border of the optic foramen (posteromedial edge of the palate positioned
anteriorly to the posterior border of the optic foramen in mordax); coronoid process low, with rounded tip slightly
above the line of the articular condyle (coronoid processes high, more triangular, above the line of the articular
condyle in mordax); upper canines distinctly grooved along the anterior surface (upper canines with convex anterior
surface not grooved in mordax); and parastyle of M1 oriented labially, and not projected over the labial margin of the
P4 (parastyle of M1 oriented forwardly, projected over the posterior labial margin of the P4 in mordax).
Regarding the remaining species—all of them either from the Amazon basin or northwestern South America—
L. peracchii can be distinguished from L. thomasi, L. cadenai Woodman & Timm, 2006, and L. pattoni Woodman
& Timm, 2006 by the larger external and cranial size (FA ≤ 34.1 mm, and GLS ≤ 22.5 mm in thomasi, cadenai, and
pattoni); from L. concava Goldman, 1914 by the larger mandible (MAL ≤ 15.5 mm in concava; see Woodman &
Timm [2006]); from L. chocoana Dávalos, 2004, and L. orienticollina Dávalos & Corthals, 2008 by the smaller
external size (FA ≥ 40 mm in chocoana and orienticollina); from L. handleyi Hill, 1980, L. hesperia G. M. Allen,
1908, and L. orcesi Albuja & Gardner, 2005 by the smaller skull (GLS: 26.9–29.2 mm in handleyi, 26–28 mm in
hesperia, and > 29 mm in orcesi); also it can be distinguished from L. handleyi, and from L. robusta Miller, 1912
by the smaller and narrower skull and rostrum (BAC > 6 mm, MAB > 10 mm, and MTL > 9 mm in handleyi and
robusta); and from L. fornicata Woodman, 2007 by the more anteriorly positioning of the posterior border of the
anteorbital foramen, and the shallower posterior portion of the palate.
The following set of external characters is useful to field identifications of L. peracchii: forearm length 37.0
mm or less; tragus with rounded tip; ventral pelage slightly bicolor, with dark-brown bases and pale-brown tips,
brownish in the general appearance, and contrasting slightly with the dorsal pelage (see Figures 2 and 7).
Remarks: This research is the first result of a series of studies re-evaluating taxonomic limits among species
of the genus Lonchophylla from the eastern portion of South America. We conclude that populations previously
assigned to L. bokermanni from Cerrado and Atlantic Forest constitute two distinct lineages, with L. bokermanni
restricted to Cerrado, and L. peracchii restricted to southeastern Atlantic Forest.
Due to the similarity in size of the specimen reported by Pimenta et al. (2010) from Reserva Biológica de
Sooretama with L. peracchii, we expect the occurrence of this species in the Atlantic Forest of Espírito Santo; but
based on the restricted distribution ranges and apparent endemism revealed for most congeners (see Dávalos 2004;
Albuja & Gardner 2005, Woodman 2007; Dávalos & Corthals 2008), and the high bat sampling effort in
southeastern Brazil (see Bergallo et al. 2003), we suppose L. peracchii is restricted to Atlantic Forest. Additionally,
after the reidentification of specimens previously assigned to L. mordax from Rio de Janeiro (Dias et al. 2002;
Esbérard et al. 2006) as L. peracchii, we suppose L. mordax does not occur in the State. We recommend the
reidentification of specimens from Atlantic Forest localities assigned to L. mordax (see Pedro & Passos 1995; Faria

NEW LONCHOPHYLLA FROM THE ATLANTIC FOREST Zootaxa 3722 (3) © 2013 Magnolia Press · 357
et al. 2006). Although our morphological comparisons have revealed qualitative traits useful in the distinction
between L. mordax and L. dekeyseri, we examined only two specimens of the latter. Comprehensive samples of
both taxa must be examined to assess the consistency of these traits, and their application in delimiting species.
Due to the high concentration of endemic species and exceptional habitat loss, Atlantic Forest and Cerrado
biomes have been considered biodiversity hotspots for conservation priorities (Myers et al. 2000; Mittermeier et al.
2004; Ribeiro et al. 2009). With the assignment of Atlantic Forest and Cerrado populations of Lonchophylla to distinct
and apparently endemic species—L. peracchii and L. bokermanni, respectively—efforts to increase our background
on the biology and distribution of these species are required to support conservation actions. Among them, we
recommend critical review of museum specimens, and additional capture efforts focused on nectar-feeding bats.
Information on natural history of L. peracchii is available under the name L. bokermanni in Taddei et al.
(1988), Esbérard et al. (1997; 2006; 2010), Baptista & Mello (2001), Dias et al. (2002), Esbérard (2003; 2007;
2009), Brito et al. (2004), Moratelli & Peracchi (2007), Dias & Peracchi (2008), Dias et al. (2008), Lourenço et al.
(2010), and Novaes et al. (2010).

Acknowledgments

The following researchers, curators and collection staff provided access to specimens under their care: E. Morielle-
Versute (Universidade Estadual Paulista “Júlio de Mesquita Filho”, Brazil), A. L. Peracchi (Universidade Federal
Rural do Rio de Janeiro, Brazil), and L. Geise (Universidade do Estado do Rio de Janeiro, Brazil). We are greatful
to D. E. Wilson (National Museum of Natural History, United States) and A. L. Gardner (United States Geological
Survey Patuxent Wildlife Research Center, United States) for helpfully reviewing a draft of this manuscript. L.
Geise (Universidade do Estado do Rio de Janeiro, Brazil), I. P. de Lima, M. R. Nogueira (Universidade Federal
Rural do Rio de Janeiro, Brazil), and L. Mayumi (Universidade Estadual Paulista “Julio de Mesquita Filho”,
Brazil) provided photographs of museum and live specimens. The National Council for Scientific and
Technological Development (CNPq), Brazil, provided financial support for C. E. L. Esbérard (CNPq 473596/2006-
7), and for R. Moratelli (CNPq 202612/2011-2).

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APPENDIX

Specimens examined. Abbreviations of institutions are as follows: Instituto de Biologia, Universidade Federal Rural do Rio de
Janeiro (ALP, and LMD), Seropédica, Brazil; Universidade Estadual Paulista “Júlio de Mesquita Filho”, São José do Rio Preto,
Brazil (DZSJRP); and Museu Nacional, Univerisdade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (MN). Specimens
marked with an asterisk were used in the morphometric analyses.

Lonchophylla bokermanni (08): Brazil, Minas Gerais: Serra do Cipó (19o16’S, 43o36’W: DZSJRP 10342* [paratype], 10347*
[holotype], 10408* [paratype], 11410* [paratype], 11411* [paratype], 10412* [paratype; referred in the original
description as ZUEC 585]), and Diamantina (18o23’S, 43o61’W: MN 79996*, MN 79997*).
Lonchophylla dekeyseri (02): Brazil, Distrito Federal: Parque Nacional de Brasília (15o41’S, 47o59’W: DZSJRP 10099*
[holotype]); Brazil, Piauí, Sete Cidades, Piracuruca (DZSJRP 11459* [paratype]).
Lonchophylla mordax (32): Brazil, Sergipe: Itabaiana (10o68’S, 37o42W: ALP 8769*, 8770*, 8812*, 8813*, 8814*, 8815*,
8816*, 8817*, 8818*, 8819*), and Parque Nacional Grota do Angico (9o65’S, 37o67’W: ALP 9747*, 9752*, 9755*,
9757*, 9759*, 9761*, 9762*, 9768*, 9769*, 10075*, 10076*, 10077*, 10078*, 10079*, 10080*, 10081*, 10082*,
10084*, 10085*, 10086*, 10087*, 10088*).
Lonchophylla peracchii sp. n. (71): Brazil, Rio de Janeiro: Ilha Grande (23o10’S, 44o12’W: DZSJRP 15159* [paratype], 15160,
15161, 15162 [holotype]*, 15163 [paratype]*, LDM 246, 2090, 3450, 3896, 3897, 4052*, 4233, 4533*), Cambuci
(21o34’S, 41o54’W: LDM 4250*, 4253, 4477*), Reserva do Grajaú (22o55’S, 43o16’W: ALP 1783, 1784, 1785, LDM
237*, 238, 246, 247, 248, 250, 270, 280, 281, 345*, 395, 531, 532*, 533, 1359*, 1495, 1496, 1497, 1499), Parque Estadual
da Pedra Branca (22o52’S, 43o23’W: ALP 5664*, 5820*, 5860*); Reserva Rio das Pedras (22o59’S, 44o06’W: LDM 1781*,
3700*), Morro de São João (22o29’S, 41o58’W: LDM 2219*, 2245, 4113, 4222*, 4226*, 4227*), Floresta da Tijuca
(22o57’S, 43o24W: LDM 1064*, LDM 1460), Jardim Botânico (22o58’S, 43o13’W: LDM 875), Reserva Biológica do
Tinguá (22o39’S, 43o34’W: ALP 6265*, 6560*, 6561*, 6283*, 6284*, 6556*, 6657*, 6658*, 6559*, 6656*), Ilha da
Gipóia (23o02’S, 44o21’W: LDM 4200*, 4423*), Ilha de Itacuruçá (23o56’S, 43o53’W: LDM 5085*), Parati (23o19’S,
44o38’W: LDM 996, 997*), Estrada Rio-Santos (23o55’S, 43o16’W: LDM 5008*, 5010*), and Vale do Rio Sahy (23o55’S,
43o59’W: LDM 5128*), Parque Nacional da Serra dos Órgãos (22o26’S, 42o59’W: ALP 6482).

360 · Zootaxa 3722 (3) © 2013 Magnolia Press DIAS ET AL.