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Number 3426, 14 pp., 3 figures, 3 tables February 27, 2004
A New Chocoan Species of Lonchophylla
(Chiroptera: Phyllostomidae)
LILIANA M. DAVALOS!
ABSTRACT
Lonchophylla is a diverse genus of glossophagines characterized by large, forwardly pro-
jecting inner upper incisors and the absence of zygomatic arches. Seven species are currently
recognized, including the large-bodied (greatest length of skull >24.5 mm) robusta, handleyi,
hesperia, and bokermanni and the small-bodied (greatest length of skull <24.5 mm) thomasi,
dekeyseri, and mordax. Lonchophylla species range throughout the Neotropics and include
endemics in Amazonia, the Cerrado, and the arid regions of coastal Peru and Ecuador. In this
paper I describe a new large-bodied species, Lonchophylla chocoana, from the subtropical
rainforests of the Choc6 in southwestern Colombia and northwestern Ecuador. I also document
the diagnostic external, craniodental, and mitochondrial characters of the new species and
summarize morphological characteristics for the new species and its sympatric congeners.
INTRODUCTION three genera constitute the phyllostomid tribe
Lonchophyllini. Recent analyses of morpho-
The genus Lonchophylla Thomas includes logical and mitochondrial DNA-sequence
seven species of glossophagines character- characters from lonchophyllines have recov-
ized by large, forwardly projecting inner up- ered a paraphyletic Lonchophylla, although
per incisors and the absence of zygomatic low support values indicate that new evi-
arches (Taddei et al., 1983; Thomas, 1903). dence might easily overturn this conclusion
Two closely related monotypic genera distin- (Davalos and Jansa, 2004).
guished primarily by larger size and the mor- As members of the phyllostomid subfam-
phology of premolars are Platalina Thomas ily Glossophaginae, species of Lonchophylla
and Lionycteris Thomas. Together, these have a rostrum roughly equal in length to the
' Division of Vertebrate Zoology (Mammalogy), American Museum of Natural History; and Department of Ecology,
Evolution, and Environmental Biology, Columbia University. e-mail: [email protected]
Copyright © American Museum of Natural History 2004 ISSN 0003-0082
a AMERICAN MUSEUM NOVITATES NO. 3426
braincase and a small but well-developed METHODS
nose leaf (Koopman, 1994). Lonchophylla
STUDY SITE AND COLLECTION METHODS
species range from Nicaragua to Peru, Boliv-
ia and southeastern Brazil, and west of the The known specimens of this new species
Andes through the Choc6é south to the arid were collected 2 km south of Alto Tambo
regions of Ecuador and northwestern Peru. (00°54'N, 78°33'W; 700 m) in Provincia Es-
Large-bodied Lonchophylla species (greatest meraldas, Ecuador, and at La Guaraperia, ca.
length of skull >24.5 mm) include robusta, 3 km NW of Junin (1°21'N, 78°08'W; ca.
handleyi, hesperia, and bokermanni. Loncho- 900 m) in Departamento Narifio, Colombia
Phylla thomasi, dekeyseri, and mordax are (fig. 1; Burton Lim, personal commun.; Cad-
comparatively smaller (greatest length of ena et al., 1998). The average annual rainfall
skull <24.5 mm, Taddei et al., 1983). Lon- at the elevation belt encompassing both Alto
chophylla species are typically found in for- Tambo and La Guaraperia (fig. 1) is estimat-
ests, gardens, and plantations, with L. hes- ed to be around 2800 mm (ESRI, 1997), and
peria restricted to deciduous forest, L. bok- the average annual temperature is estimated
ermanni to the Cerrado, and L. handleyi, L. to be around 25°C (ESRI, 1999). The Alto
robusta, and L. mordax ranging to cloud-for- Tambo locality comprised humid disturbed
est habitat (Cadena et al., 1998; Handley, secondary forest (Burton Lim, personal com-
1976; Koopman, 1994; Sazima et al. 1978). mun.). The vegetation at La Guaraperia had
Lonchophylla feed predominantly on nectar, been selectively logged without ever being
although insects, fruit, and pollen are also fully cleared (Cadena et al., 1998).
part of their diet (Gardner, 1977; Sazima et Both specimens were captured using mist
al., 1978). nets. At Alto Tambo, nets were set along a
The lowland Lonchophylla species are eas- trail and along a river surrounded by dis-
ily captured by mistnetting (Handley, 1976; turbed secondary growth, and there was rain
Sazima et al., 1978; Simmons and Voss, before the capture of the specimen (B. Lim,
1998), and L. thomasi is widespread through- personal commun.). At La Guaraperia nets
out its range and appears to be the most com- were set across a stream, a trail, and in a
mon glossophagine found in Amazonia (Em- cleared area (Cadena et al., 1998). Mist nets
mons, 1997). Lonchophylla congeners, how- were opened at 1700 hours and stayed open
ever, have not been found in sympatry at overnight until 0500 hours at Alto Tambo,
well-sampled lowland localities (Simmons while at La Guaraperia nets were open at
and Voss, 1998; Voss and Emmons, 1996), dusk and closed around 2200 hours (B. Lim,
although mordax and robusta were recently personal commun.; Cadena et al., 1998).
collected together at 1400 m above sea level Nets varied from 10 to 18 m in length and
in Altaquer, southwestern Choc6é of Colom- were set to reach 3 m above ground at both
bia (Cadena et al., 1998). localities.
In 1999 Sharon A. Jansa and I initiated a
collaboration to sequence cytochrome b from
MEASUREMENTS AND ABBREVIATIONS
lonchophyllines to resolve relationships
All measurements reported here are from
among the bats in this tribe. Among others,
adult individuals with closed epiphyses. The
we obtained samples from 11 individuals
first five measurements listed below were
representing five species of Lonchophylla
taken from skin tags or other records made
(for details and results see Davalos and Jan-
by the collector, but other dimensions were
sa, 2004). One of those species is an unde-
measured using digital calipers. Linear mea-
scribed large Lonchophylla resembling L. ro-
surements of external and craniodental di-
busta and L. handleyi in size, but differing
mensions are reported in millimeters (mm);
from these and other known congeners by a
weights are reported in grams (g). Measure-
unique combination of traits. I describe this
ments follow Simmons et al. (2002), and are
new species below, document its diagnostic
described below:
morphological and molecular attributes, and
summarize available observations in a com- Total length: Distance from the tip of the snout to
parative context. the tip of the last caudal vertebra.
2004 DAVALOS: NEW SPECIES OF LONCHOPHYLLA 3
7 : ; || a y . ‘7 f4 * ‘Le j — Sin" “| ‘o
(| i. ; PN “| ff fr
7 \ ~_ B P f o Hy
SS a ’ J /
= ¥ |\ be # | F, él
XY ‘
~ Yo, ae a i
1 A —— i's 4 !
p a ag x wh VY I
\ | : en Junin* /
pmmsfi/ f my ‘is é/‘ x — : Alt aquear Ny - “he\ \
Cer ~ Bieta 3 Ss Ricaurte \
y i" i ( .— x
i S o—~, 1 \ < a
\ 7 ae 2 { ey \ \ . _ 7 ta SS oF
Ae io oa f % | \ ue “oe, Ay
He ie ~. | ~
4 \ | ie a Colombia
{ KY i“ Alto Tambo#—. ~~
5 / te SY :
( \ ve ow Ecuador Tulcan
5 \’ ‘ ian. =, ;
( My } hy \ \
Af y iSae e A PSee sao . as
f _ é |
Py es \ gon 4 : ‘. Weg \ 7 f
oya } ey s h 8| ()() PT)f e | se. =~ [ A Lo \ | {1 a J
| / | ali ty \ \ i d \ a
{ ~ ioe \ \ , } [
a sn, vo
2 ee se ee eee Fa
Fig. 1. Southern Chocé of Colombia (Departamento Narifio) and Ecuador (Provincia Esmeraldas),
showing the towns closest to the localities at Alto Tambo (Alto Tambo) and La Guaraperia (Junin). Gray
outlines indicate major roads, black outlines correspond to rivers. See text for map coordinates.
Tail length: Measured from the point of dorsal Palatal length: From the anteriormost point be-
flexure of the tail with the sacrum to the tip of hind the incisors to the edge of the bony pala-
the last caudal vertebra. tion.
Hindfoot length: From the anterior edge of the Condylo-incisive length: From the posteriormost
base of the calcar to the tip of the claw of the point on the occipital condyles to the anterior-
longest toe. most point on the upper incisors.
Ear length: From the notch to the fleshy tip of the Postorbital breadth: Least breadth across frontals
pinna. posterior to the postorbital bulges.
Forearm length: From the elbow (tip of oleocra- Braincase breadth: Greatest breadth of the glob-
non process) to the wrist (including the car- ular part of the braincase.
pals), measured with the wing partially folded. Mastoid breadth: Greatest cranial breadth across
Thumb length: From the metacarpal-phalangeal the mastoid region.
joint to the tip of the claw of the thumb. Maxillary toothrow length: From the anteriormost
Tibia length: From the proximal end of the tibia edge of the canine crown to the posteriormost
to the posterior base of the calcar. edge of the crown of M3.
Greatest length of skull: From the posteriormost Breadth across molars: Greatest breadth across
point on the occiput to the anteriormost point the outer edges of the crowns of the upper mo-
on the premaxillae (excluding the incisors). lars.
4 AMERICAN MUSEUM NOVITATES NO. 3426
Height of upper canine: Dorsoventral height from exhaustive mammal inventories in neotropi-
the anterior base of the crown of the upper ca- cal forests west of the Andes have been re-
nine to the tip. stricted to Barro Colorado, Panama, and La
Breadth at the base of upper canine: Greatest an-
Selva, Costa Rica (Voss and Emmons, 1996),
teroposterior breadth at the base of the upper
it is likely that the known distribution is just
canine.
an artifact of inadequate collecting, com-
In attempting to make taxonomic identifi- pounded with superficial similarities to L. ro-
cations I compared the vouchers from Alto busta and L. handleyi. In particular, speci-
Tambo and La Guaraperia with original de- mens of L. handleyi reported from the west-
scriptions, notes on distribution, and com- ern slopes of the Cordillera Occidental of
parative series of specimens of all other Lon- Departamentos of Valle and Narifio, Colom-
chophylla species. The following institution- bia, and specimens of Lonchophylla sp. from
al acronyms correspond to museums in the Pacific lowlands of Ecuador (Alberico
which vouchers and other specimens exam- and Orejuela, 1982; Alberico, 1987; Albuja,
ined are preserved: AMNH, American Mu- 1999) deserve investigation.
seum of Natural History (New York); ICN, ETYMOLOoGy: For the Choc6, an area of en-
Instituto de Ciencias Naturales (Bogota); demism comprising tropical and subtropical
MHN, Museo de Historia Natural (Popayan); humid forests west of the Andes from west-
ROM, Royal Ontario Museum (Toronto); ern Panama through Colombia to northern
TTU, The Museum, Texas Tech University
Ecuador.
(Lubbock); USNM, National Museum of
DIAGNOsIs: A large-sized species of Lon-
Natural History (Washington, DC).
chophylla (forearm 45—48 mm; weight 19—
23 g) with chocolate-brown to chestnut-col-
SYSTEMATICS
ored dorsal fur and brown ventral fur; dorsal
FAMILY PHYLLOSTOMIDAE GRAY, 1825 and ventral hairs bicolored (the former more
distinctly banded than the latter), 7-8 mm
SUBFAMILY GLOSSOPHAGINAE BONAPARTE,
1845 long in shoulder region; pinnae short with
rounded tips; thumb large (7.5—8.3 mm);
GENUS LONCHOPHYLLA THOMAS, 1903
fringe of the uropatagium sparse; calcar
Lonchophylla chocoana, new species shorter than foot; palate long; with postpa-
Figures 2, 3 latine torus; right and left upper I1 meet at
distal third of the crown; large gap present
TYPE MATERIAL: The holotype (ROM
105786), an adult female preserved as skin, between I] and I2, and smaller gap between
skull, skeleton, and tissue, was collected in I2 and C; outer upper incisors (12) pointed
the vicinity of Alto Tambo by Mark Engs- ventrally more than ventromedially; upper
trom, Burton Lim, and Francisco Sornoza canines large; height of P3 slightly less than
(original number F40079) on 3 March 1996. P4; gap present between C and P3; smaller
One paratype is an additional female (CN gap present between P3 and P4; P4 with
13649) collected by Alberto Cadena, Pilar small but well-developed lingual cusp; M1
Rivas, and Robert P. Anderson (original and M2 roughly the same width; lower in-
number ACG 2765) at La Guaraperia on 12 cisors small and trilobed, with crown height
March 1995. approximately equal to crown width.
DISTRIBUTION: Currently known only from Of the characters listed above, five are par-
northwestern Ecuador and southwestern Co- ticularly useful for field identification of Lon-
lombia (fig. 1), but perhaps widespread chophylla chocoana: size (forearm 44—48
throughout the lower to middle elevations of mm; weight 19—23 g); thumb length (7.5-8.3
the southern biogeographic Choc6. Because mm); dorsal fur color; ventral fur bicolored;
->
Fig. 2. Dorsal (A), ventral (B), and lateral (C) views of the skull of the holotype of Lonchophylla
chocoana (ROM 105768). Scale bar = 5 mm.
2004 DAVALOS: NEW SPECIES OF LONCHOPHYLLA
6 AMERICAN MUSEUM NOVITATES NO. 3426
TABLE 1 probably larger than their male conspecifics.
Measurements of the Specimens of Although this might produce some overlap
Lonchophylla chocoana in measurements between female robusta
and male chocoana, other nonmetric char-
Holotype Paratype acters remain diagnostic.
ROM 105786 ICN 13649 From all other large Lonchophylla, cho-
Sex Female Female coana can be unambiguously distinguished
Weight 23.0 19.0 on the basis of fur coloration, forearm length,
Total length 78.0 84.5 greatest length of skull, and palatal length.
Tail length 7.0 10.5 Additionally, chocoana can be distinguished
Hind foot length 15.0 13.2 from robusta on the basis of size of the
Ear length 17.0 14.0 thumb, greatest length of skull, palatal
Forearm length 48.0 45.0
length, orientation and size of the outer upper
Thumb length dd 8.3
incisors, and the relative size of gaps be-
Tibia length 16.5 18.2
tween I1, I2 and C, which are roughly equal
Greatest length of skull 27.7 28.3
in robusta. L. chocoana can be distinguished
Palatal length 7.0 6.9
from handleyi on the basis of the furry fringe
Condyloincisive length 20.9 27.4
Postorbital breadth 5.3 5.3 along the uropatagium, size of the thumb, the
Braincase breadth 10.2 10.5 relative size of the upper canines, presence
Mastoid breadth 6.9 7.0 of a ridge along the posterior edge of the
Maxillary toothrow length 9.8 11.7 palate (postpalatine torus), and the well-de-
Breadth across molars 7.3 7.3 veloped lingual cusp on P4. Lonchophylla
Height of upper canine 2.6 2.7 chocoana is larger than all remaining con-
Breadth at base of upper canine 1.7 1.7
geners in most anatomical dimensions. L.
hesperia and bokermanni may reach similar
length of skull, but they have greater skull
and trilobed lower incisors. Within the genus length-to-width ratios and are absent from
Lonchophylla this combination of traits is the known range of chocoana (see measure-
unique to L. chocoana. Appendix 1 summa- ments in Taddei et al., 1983).
rizes the molecular data that prompted the As with robusta and handleyi, the dorsal
investigation of morphological differentia- pelage of chocoana is composed of bicolored
tion in L. chocoana. The average uncorrected hairs with cream-white bases and brown tips.
pairwise sequence divergence of the mito- The length of the dorsal fur along the upper
chondrial cytochrome b gene of L. chocoana back in chocoana is approximately 7—8 mm,
with respect to L. robusta from Colombia is slightly longer than in handleyi and robusta
11.6%, to L. robusta from Panama is 12.3%, (4.0-7.5 mm). The ventral pelage of cho-
and to L. handleyi is 11.6%. These genetic coana contrasts with both handleyi and ro-
distances are indicative of species-level dif- busta in being bicolored from neck to genital
ferentiation, according to one recently pro- region, whereas in handleyi the ventral fur is
posed criterion (Bradley and Baker, 2001). exclusively beige-brown unicolored, while
MEASUREMENTS: Dimensions of both spec- some robusta individuals (particularly fe-
imens of Lonchophylla chocoana are provid- males) show bicolored hairs around the neck
ed in table 1, and metrical comparisons with but never in the abdominal region.
representative series of other large congeners The cranial morphology of chocoana is
within its range are summarized in table 2. similar to that of other members of the genus.
DESCRIPTION AND COMPARISON: Loncho- L. chocoana has a relatively long rostrum, a
phylla chocoana is the largest member of the small but noticeable anteorbital inflation, and
genus, as large or slightly larger than han- a large braincase. Zygomatic arches are ab-
dleyi, and larger than robusta, hesperia, and sent as in all other lonchophyllines, and the
bokermanni (tables 1—3; Taddei et al., 1983). palate is the longest of any congener. Like
Females are generally larger than males in all Lonchophylla, L. chocoana has a dental
robusta and handleyi (table 2), and the two formula 12/2, C1/1, P2/3, M3/3 X 2 = 34.
chocoana females (table 1) are therefore The inner upper incisors are large compared
2004 DA, VALOS NEW SPECIES OF LONCHOPHYLLA
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8 AMERICAN MUSEUM NOVITATES NO. 3426
TABLE 3
Summary of Principal Diagnostic Characters for Field Identification of Large Species of
Lonchophylla from Northwestern South America
L. robusta L. handleyi L. chocoana
Weight (g) 11.0-20.0 16.0-21.0 19.0-23.0
Forearm length (mm) 41.0-46.2 45.0-46.8 45.0-48.0
Thumb length (mm) 5.2-6.9 5.4-6.3 7.5-8.3
Dorsal fur Light chestnut to orange yellow Light chestnut Dark chestnut to chocolate brown
Ventral fur Uniform beige-brown or Uniform beige-brown Bicolored from neck to genital
slightly bicolored only in region
the neck and thoracic region
Lower outer incisors Not trilobed or slightly trilobed Bilobed Always trilobed or bilobed
to the outer incisors and are separated by a mordax (the latter shows variation between
gap from each other and from the canine. In the two disjunct subspecific populations).
the latter respect chocoana also resembles all In chocoana P4 is longer than P3 (antero-
other species of Lonchophylla. posterior dimension), as it is in robusta, han-
Differences in the dentition are subtle, but dleyi, mordax, and thomasi. The height (dor-
they provide the means to distinguish species soventral dimension) of P3 is either equal to
of Lonchophylla (see Hill, 1980). In cho- or very slightly less than P4. Both robusta
coana the inner upper incisors have a small and handleyi have shorter (dorsoventral di-
triangular gap between them, meeting at the mension) P3 than P4, while mordax and tho-
distal third of the crown. This is unlike ro- masi show upper premolars that are subequal
busta, handleyi, mordax, and thomasi, all of in height. The basal lingual cusp on P4 is
which show a taller gap between the inner present and well developed in chocoana, ro-
upper incisors, meeting at the distal quarter busta, and thomasi, less developed in mor-
of the crown. The upper canines of chocoana dax, and poorly developed or absent in han-
are relatively and absolutely larger than those dleyi. The first molar of chocoana is com-
of robusta and handleyi. The posterior cusp paratively wide, and it is larger (both in an-
of the upper canines of chocoana is blunt, teroposterior and lateral dimensions) than
similar to that of handleyi, while the poste- M2, which in turn is larger than M3. Both
rior cusp is sharp in robusta, thomasi, and robusta and handleyi have similar widths of
Fig. 3. Anterior views of the lower dentition of Lonchophylla chocoana (left, ROM 105786) and
L. robusta (right, ICN 13648, male). Note the shape and height of the gap between inner upper incisors
and the trilobed lower incisors in chocoana. These morphological traits can be used in conjunction with
size and other characters for field identification.
2004 DAVALOS: NEW SPECIES OF LONCHOPHYLLA 9
M1 and M2, although M1 is longer (antero- Tambo indicate that it was captured along
posterior dimension) than M2, with overall with other glossophagines (Anoura spp.) in a
smaller M3. The first two molars of thomasi disturbed secondary forest in nets left open
and mordax are similar in length and height. overnight due to continuous rain (Burton
The lower outer incisors of chocoana are Lim, personal commun.). The specimen from
trilobed, as they are in thomasi. Lower inci- La Guaraperia was also captured in second-
sors are rarely trilobed in robusta; m. mordax ary forest, between dusk and 2200 hours,
has trilobed incisors, but m. concava does along with Anoura caudifera, A. cultrata,
not; and incisors are bilobed in handleyi. The and A. geoffroyi (Cadena et al., 1998). The
height of the lower incisors is roughly the survey in Narifio additionally captured Lon-
same as the width of these teeth in all species chophylla robusta and L. mordax within the
of Lonchophylla. The lower premolar denti- same general region at the Altaquer locality.
tion of chocoana resembles that of handleyi. Both specimens were captured in early
In robusta the maximum height (dorsoven- March: at Alto Tambo on 3 March 1996, and
tral dimension) of the series corresponds to at La Guaraperia on 12 March 1995.
p3, and the maximum length (anteroposterior Although these data are scant, some as-
dimension) corresponds to p4. In handleyi, pects are noteworthy. First, L. chocoana cur-
p4 is the highest and longest tooth of the se- rently seems rare in its range; one specimen
ries, and in comparison the p4 of chocoana from Alto Tambo prompted this investiga-
is narrow (lateral dimension). In contrast, the tion, and the search for conspecifics only
longest lower premolar of thomasi and mor- found 1 more among 11 specimens of large-
dax is p2. The height of the premolars in bodied Lonchophylla from west of the Cor-
thomasi is roughly the same throughout the dillera Occidental of Colombia. As noted
premolar series, whereas in mordax p4 is the above, a few large Lonchophylla from west-
highest premolar (dorsoventral dimension). ern Colombia and Ecuador reported in the
The first molar of chocoana is the widest and literature might correspond to chocoana.
longest of the molar series. This is the same Only further sampling can document the
condition as in handleyi, although m1 of complete range and the rarity of chocoana.
handleyi is slender in comparison. In robusta Second, as with other Lonchophylla, cho-
the m1 is slightly wider (lateral dimension) coana can be captured using ground-level
than m2, which in turn is wider than m3. In mist netting. At Paracou, French Guiana, and
robusta m1 is the longest molar (anteropos- in southern Venezuela, most L. thomasi were
terior dimension). The differences among the captured using ground-level mistnets (Han-
molars are less marked in mordax and tho- dley, 1976; Simmons and Voss, 1998), as
masi, particularly the latter. Although m1 is were all L. robusta individuals of the Smith-
the longest molar (anteroposterior dimen- sonian Venezuelan Project (Handley, 1976)
sion) in both species, height and width vary and all L. robusta from Tambito, Colombia
only slightly along the series. (Davalos and Guerrero, 1999). Third, the
The coronoid process in Lonchophylla habitat associated with the capture of L. cho-
chocoana is high and oriented at an angle of coana has been characterized as disturbed,
about 110° with respect to the toothrow, as and the simultaneous capture of other glos-
in robusta. In handleyi the coronoid process sophagines probably indicates the presence
projects at an angle of around 130°. Both tho- of local food resources, regardless of the
masi and mordax show even greater angles, habitat condition.
as well as a longish (anteroposterior dimen- Little is known about the ecological niche
sion) coronoid process in comparison to their of species in this genus, but observed diver-
larger congeners. sity in body size and habitat choice (from dry
scrub to lowland rainforest) indicates that
there is some sort of partitioning in feeding
NATURAL HISTORY
and roosting ecology. These ecological char-
Both known specimens of Lonchophylla acteristics become more intriguing now that
chocoana were captured in ground-level mist three species of Lonchophylla have been
nets. The notes on the specimen from Alto found in similar habitats, as they were in the
10 AMERICAN MUSEUM NOVITATES NO. 3426
western slopes of the Cordillera Occidental American Museum of Natural History and
of Narifio, Colombia (Cadena et al., 1998). Columbia University.
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Uma nova especie de Lonchophylla do Serra do
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