A new species of Lonchophylla (Chiroptera: Phyllostomidae) from the eastern Andes of northwestern South America

2008·1.1 MB·English

by  Liliana M. Dávalos

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3635, 16 pp., 6 figures, 2 tables December 31, 2008
A new species of Lonchophylla
(Chiroptera: Phyllostomidae) from the
eastern Andes of northwestern South America
LILIANA M. DA´VALOS1 AND ANGELIQUE CORTHALS2
ABSTRACT
Since 2004 five new species have been described in the nectar-feeding phyllostomid bat genus
Lonchophylla.AllthenewspeciesareendemictooneNeotropicalecoregion,suggestingthatmore
speciesremaintobediscoveredamongcollectedspecimenscurrentlyreferredtoseveralwidespread
taxa. Herein we describe a new species, Lonchophylla orienticollina, endemic to the middle
elevations of the eastern Andes of Venezuela, Colombia, and Ecuador. The new species
superficially resembles its sympatric congener L. robusta, but its cranial morphology and
combinationofmeasurementsaredistinctive.Throughoutitsrange,L.orienticollinaissympatric
with L. robusta, and it also overlaps with L. handleyi in the Cordillera Oriental of Ecuador. The
evolutionary processes leading to the divergence among Lonchophylla species, as well as the
ecologicalmechanismsthatenablemultiple,subtlydifferentspeciestocoexistwillremainobscure
withoutnew field andphylogenetic studies.
INTRODUCTION and Ditchfield, 2005; Koopman, 1994).
Analyses of morphological (Carstens et al.,
Fourlonchophyllinegenera—Lonchophylla, 2002;GregorinandDitchfield,2005;Wetterer
Platalina, Lionycteris, and Xeronycteris— et al., 2000; Woodman, 2007; Woodman and
form a clade of nectar-feeding phyllostomids Timm, 2006) and molecular data (Da´valos
distributedfromsouthernNicaraguatosouth- and Jansa, 2004) strongly support the mono-
eastern Brazil and, west of the Andes, to phylyofthetribe,butrelationshipsamongthe
northern Chile (Galaz et al., 1999; Gregorin included genera are less certain. Although the
1DepartmentofEcologyandEvolution,650LifeSciencesBuilding,SUNYStonyBrook,StonyBrook,NY11794-5245,
USA([email protected]).
2DepartmentofAnthropology,SBSBuilding,CircleRoad,SUNYStonyBrook,StonyBrook,NY11794-5245,USA
([email protected]).
CopyrightEAmericanMuseumofNaturalHistory2008 ISSN0003-0082
2 AMERICAN MUSEUMNOVITATES NO. 3635
four genera are morphologically distinct sured to the nearest 0.1 mm using manual
(Woodman and Timm, 2006), both morpho- calipers. External measurements are those
logical and molecular data have failed to recorded by the original collectors, with the
recover a monophyletic genus Lonchophylla, exceptions of length of thumb and length of
seeWoodman(2007),GregorinandDitchfield tibia. All measurements reported here are
(2005), and Da´valos and Jansa (2004). from adult individuals with closed epiphyses.
Growing interest in the systematics of the Linear measurements of external and cranio-
Lonchophyllini has uncovered greater species dental dimensions are reported in millimeters
diversity than previously suspected. Since the mm; weights are reported in grams (g).
publication of the latest comprehensive taxo- Measurements are described below:
nomictreatmentofChiropterabySimmonsin
2005, six species in two genera have been Total length Distance from the tip of the
snout to the tip of the last
described and one subspecies has been elevat-
caudalvertebra
ed to species status (Albuja V. and Gardner,
Tail length Measured from the point of
2005;Da´valos,2004;GregorinandDitchfield,
dorsalflexureofthetailwith
2005;Woodman,2007;WoodmanandTimm,
the sacrum to the tip of the
2006).Therecentlydescribedspeciesappearto lastcaudal vertebra
benarrowlyendemic,oftenknownfromfewer Hindfootlength Fromtheanterioredgeofthe
thanfivelocalitiese.g.,AlbujaV.andGardner baseofthecalcartothetipof
(2005), Gregorin and Ditchfield (2006), and the clawof the longest toe
WoodmanandTimm(2006).Therapidrateof Earlength From the notch to the fleshy
species discovery and narrow endemicity of tipof the pinna
Forearmlength Fromtheelbowtipofoleocra-
thetaxainquestionsuggeststhatthediversity
non process to the wrist in-
of this clade is still underestimated (Reeder
cluding the carpals, measured
et al., 2007).
withthewingpartiallyfolded
While investigating variation within and
Tibialength Fromtheproximalendofthe
among lonchophylline species to describe tibia to the posterior base of
Lonchophylla chocoana (Da´valos, 2004), we the calcar
found distinctive specimens from the foothills Thumblength Fromthemetacarpal-phalan-
of the Cordillera de Me´rida of Venezuela, geal joint to the tip of the
the Serran´ıa del Perija´ of Venezuela, the clawof the thumb
Cordillera Oriental and the Serran´ıa de la Greatestlength Fromtheposteriormostpoint
ofskull ontheocciputtotheanterior-
Macarena of Colombia, and the Cordillera
mostpointonthepremaxillae,
Oriental of Ecuador. These specimens repre-
includingtheincisors
sent an unrecognized Lonchophylla externally
Condylo-incisive Fromtheposteriormostpoint
similar to L. robusta (Miller, 1912), but dif- length on the occipital condyles to
fering from that species, and from other theanteriormostpointonthe
congeners, by a unique combination of traits. upper incisors
Inthisarticle, wedescribethisnewspecies,its Condylo-canine Fromtheposteriormostpoint
diagnostic morphological attributes, and pre- length on the occipital condyles to
sent a summary of available observations on theanteriormostpointonthe
upper canines
its distribution and natural history.
Maxillary toothrow From the anteriormost edge
length of the canine crown to the
MATERIALS AND METHODS posteriormost edge of the
crownof M3
MEASUREMENTS Maxillary breadth Dorsal breadth of the maxil-
larybone at M2
Most measurements follow those of
Breadthacross Greatest breadth across the
Woodman and Timm (2006), who described
molars outer edges of the crowns of
and illustrated skull dimensions relevant to
the upper molars
lonchophyllines, with additional measure- Mastoidbreadth Greatestcranialbreadthacross
ments as described for chocoana (Da´valos, the mastoid region, excluding
2004). Thumbs, tibias, and skulls were mea- themastoidprocesses
2008 DA´VALOS AND CORTHALS: NEWSPECIES OFLONCHOPHYLLA 3
Zygomaticbreadth Greatest breadth of the pos- length, maxillary breadth, breadth across
teriorzygomatic processes molars, and mandibular length. Because there
Postorbital breadth Least breadth across frontals are no estimates of significance associated
posterior to the postorbital
with PC analyses, we used these same mea-
bulges
surements to generate a discriminant function
Braincasebreadth Greatestbreadthoftheglob-
withspeciesidentityasthedependentvariable.
ular partof the braincase
Palatallength From the anteriormost point This allowed us to evaluate both the signifi-
behind the incisors to the cance of the assignment function and the
edgeof the bonypalation accuracy of classification based on these
Mandibular length From the anteriormost edge measurements only. Variables for both anal-
of the mandible to the poste- ysesweremeasuredfrom23individualsofthe
riormostedgeofthemandib- new species from Venezuela, Colombia, and
ular ramus
Ecuador; and 34 L. robusta from Costa Rica,
Mandibular From the anteriormost edge
Panama, Colombia, Venezuela, and Ecuador.
toothrowlength of the canine crown to the
posteriormost edge of the
crownof m3 DISTRIBUTIONAL MODELING
Heightof the From the lowermost edge of
coronoidprocess the mandible to the upper- To estimate the potential range of the new
most edge of the coronoid species based on climatic variables, we gener-
process
ated an environmental niche model based on
thelocalitydataassociatedwiththespecimens
To identify the specimens from Venezuela,
examined. Nineteen unique localities for the
Colombia, and Ecuador, we compared these
new species were used to infer the environ-
withoriginaldescriptions,notesondistribution,
mental niche. Environmental niche models
and series of specimens from throughout the
(ENMs) are grounded on the relationship
range of Lonchophylla in southeastern Central
between a species’ ecological requirements
America and northwestern South America
and the environmental characteristics of its
(appendix). We examined specimens from the
distribution (Graham et al., 2004). ENMs
following collections: AMNH, American
integrate known records with environmental
Museum of Natural History (New York);
data(e.g.,seasonalityintemperature),andthe
IAvH, Instituto de Investigacio´n de Recursos
resulting statistical models can then be pro-
Biolo´gicos Alexander von Humboldt (Villa de
jected across spatial layers of environmental
Leyva); ICN, Instituto de Ciencias Naturales
variables to infer the suitability of the envi-
(Bogota´); MHN, Museo de Historia Natural
ronment beyond the known distribution, e.g.,
(Popaya´n); NHM, Natural History Museum
(London); ROM, Royal Ontario Museum Jarvis et al. (2003).
(Toronto); USNM, United States National Climatic data including temperature, pre-
Museum(Washington,DC). cipitation, and their extremes and variability
were obtained from published high-resolution
,1 km climate layers compiled from ground
QUANTITATIVE ANALYSESOF MORPHOLOGY
stations from 1950–2000 (Hijmans et al.,
The new species described here is most 2005). The maximum entropy algorithm im-
similar in size (table 1) and morphology to plemented in maxent v. 3.2.1 (Phillips and
Lonchophylla robusta Miller (1912). To com- Dud´ık, 2008) was applied to obtain the
pare the new species with L. robusta in a function relating occurrence points to local
quantitative framework, we extracted princi- climate. The function maximizes the entropy
pal components (PC) from the correlation of the predicted habitat suitability and is
matrix of six measurements most clearly constrained sothat theexpectedvalue ofeach
linked to qualitative differences between the climate variable (or its transform) in the
taxa, and plotted the resulting factor scores. distribution equals its empirical mean
The variables selected for principal compo- (Phillips et al., 2006). This ENM allowed us
nents analyses using SPSS 16.0 for Mac were: to quantify climate suitability the new species
thumb length, greatest length of skull, palatal across northwestern South America.
4 AMERICAN MUSEUMNOVITATES NO. 3635
5
5 6 1 2 2 6 4 1 6
usta 33females (58–96)26(7–12)24(10–29)26(14–19)25(41–46.2)17(12.5–20)15(12.6–16.6)2(5.0–7.4)24(25.5–27.4)2 (24.1–26.0)3 (23.0–24.65) (8.9–9.9)32 (6.05–6.85)3(6.15–6.8)32 (10.2–11.1)2 (10.2–11.3)2(4.8–5.5)32(9.6–10.5)32 (13.4–15.1)3 (17.2–18.8)2(9–10.3)26 (4.2–5.3)26
b 920175484 6 2 6 47 9 036 4 06 3
o 199089104 2 0 4 44 5 710 2 05 8
r 6.8.3.6.3.4.5.6.6. 5. 4. 9. 6.6. 0. 0.5.0. 4. 8.9. 4.
a 7 11411 2 2 2 1 1 1 1 1
America Lonchophyll 6males 60–86)227–11)2010–28)229–19)2143–46)1211–17.9)1612.8–16.7)194.9–6.8)1925.8–27.9)21 24.7–27.6)22 23.6–25.5)21 9.3–10.1)21 6.1–7.0)236.15–7)22 10.3–11.2)22 10.4–11.5)214.7–5.4)239.9–10.6)23 13.4–15.7)22 17.2–19.4)228.8–10.6)22 4.9–6.1)22
h 2 ((((((((( ( ( ( (( ( ((( ( (( (
ut 803019651029087892 77 48 67 5959 85 010330 29 3270 39
So 73.9.13.15.44.14.15.5.26. 25. 24. 9. 6.6. 10. 11.5.10. 14. 18.9. 5.
n
r 7
omnorthweste Lonchophyllachocoana 7females 79(70–84.5)707(7–11)775(10–15)700(14–17)733(42–48)650(19–23)335(16.5–18.2)621(6.1–8.3)700(25.7–26.3)2 74(24.35–27.4) 00(18.3–23.7)2 67(8.89–11.7)7 95(6.5–7.2)704(6.7–7.3)7 80(10.6–11)2 3121(5.1–5.4)728(10.1–10.5)7 69(13.36–15.8)765(17.3–18)220(9.2–9.2)2 90(4.9–4.9)2
fr 78.9.12.16.44.20.17.7.26. 25. 21. 9. 6.7. 10. 11.5.10. 14. 17.9. 4.
a 4
TABLE1sofsimilarlysizedLonchophyll Lonchophyllahandleyi 5males4females 7(66–80)38410(5–7)2617(10–14)31410(17)31713(45–46.8)347.710(12–21)21615(16.4–17.3)216.816(5.3–6.9)35.810(28.9–28.9)127.97(27.4–28.7)35(26.3–27.9)527.08(26.6–27.8)4125.87(25.5–26.2)34(9.9–10.7)510.07(9.88–10.2) 9(6.3–7.3)56.75(6.6–6.8)40(6.4–6.8)56.62(6.4–6.8)4 111.00(10.6–11.5)3110.97(10.9–11)37(4.96–5.3)55.15(5–5.29)42(9.8–10.3)510.30(10.2–10.4)40(15.5–17.1)16.21(15.8–16.6)4119.40(19.1–20)3110.33(10.3–10.4)315.03(4.9–5.2)3
men 72.66.011.617.045.916.516.86.128.9 26.9 26.7 10.2 6.76.6 11.3 11.15.010.1 16.0520.310.8 5.5
surementsofspeci ´entıcollina 16females 2.24(55–80)99.25(7–11.3)86.47(11.5–29)96.52(15–17.3)93.57(43–44)33.85(12.9–16)64.67(14.3–16.2)96.21(5.2–6.8)95.72(24.8–26.9)154.36(23.3–25.4)153.33(22.1–24.4)159.14(8.6–9.4)15 6.34(5.9–6.5)156.28(5.7–6.6)15 0.40(9.8–11.0)15 0.65(10.3–11.0)145.12(4.9–5.4)150.02(9.5–10.5)15 3.60(12.7–14.4)157.24(16.2–18.5)149.29(9.0–9.8)15 4.75(4.2–5.3)15
ea ori 7 11411 2 2 2 1 1 1 1 1
m a 8 7 7 8 7
Summaryof Lonchophyll 19males 3(67.5–81)188(6.5–10)174(10–35)180(11–18)187(40–47)108(11–15.9)178(14.5–16.6)12(5.8–7.3)184(24.7–26.7) 8(23.4–25.4) 2(21.8–24.3) 9(8.5–9.7)19 8(5.9–6.8)199(5.9–6.7)18 8(10.0–11.0)1 9(10.1–11.4)19(4.7–5.4)188(9.6–10.8)18 4(12.2–14.9)1 8(16.3–18.0)16(8.8–10.0)16 8(4.5–5.5)17
574555256738170 33 5 701 5 22 0
6.8.4.5.3.3.5.6.5.14.13.19. 6.6. 0. 0.5.0. 3. 7.9. 5.
7 11411 2 2 2 1 1 1 1 1
Measurement TotallengthTaillengthHindfootlengthEarlengthForearmlengthWeightTibialengthThumblengthGreatestlengthofskullCondylo–incisivelengthCondylo–caninelengthMaxillarytoothrowlengthMaxillarybreadthBreadthacrossmolarsMastoidbreadth ZygomaticbreadthPostorbitalbreadthBraincasebreadth Palatallength MandibularlengthMandibulartoothrowlengthHeightofcoronoidprocess
2008 DA´VALOS AND CORTHALS: NEWSPECIES OFLONCHOPHYLLA 5
RESULTS del Guayuriba, Acac´ıas, 480 m elevation,
Departamento del Meta, Colombia. One male
(ICN 13839) collected by Alberto Cadena
SYSTEMATICS
(original number ACG2782) on 26 September
FAMILY PHYLLOSTOMIDAE GRAY, 1995 at the forest surrounding the Guayuriba
1825 river, vereda Brisas del Guayuriba, Acac´ıas,
480 m elevation, Departamento del Meta,
SUBFAMILY GLOSSOPHAGINAE
Colombia. Two males (ICN 14399 and 14400)
BONAPARTE, 1845
collected by students of group 5 of the
GENUS LONCHOPHYLLA THOMAS, Introduccio´n Sistema´tica Animal course (orig-
1903 inalnumbers11and15)on5June1996atfinca
La Estrella,veredaElVergel, Cubarral,750 m
Lonchophylla orienticollina, new species
elevation, Departamento del Meta, Colombia.
One male (IAvH 6679) collected by Yaneth
Eastern Cordilleran Nectar Bat
Mun˜ozSaba(originalnumberYMS702)on23
Figures 1, 2 September 1999 at P.N.N. Tama´, Finca
‘‘San Isidro’’ de Pablo Contreras (07u079220N
Lonchophylla robusta: Handley 1976: 21 part, not 72u149420W), 1000 m elevation, Rio Negro,
LonchophyllarobustaMiller,1912. Mpio.Toledo,NortedeSantander,Colombia.
Lonchophylla robusta: Alberico, Cadena, Herna´ndez-
One maleand one female (USNM419409 and
Camacho, and Mun˜oz-Saba 2000: 153 part, not
LonchophyllarobustaMiller,1912. 419410)collectedbyNormanE.Peterson,Fred
Lonchophylla robusta: Da´valos 2004: 14 part, not P. Brown, John O. Matson, and C. E. Yunker
LonchophyllarobustaMiller,1912. (originalnumbers22129and22600)on17April
Lonchophylla robusta: Woodman and Timm 2006: 476
1968atKasmera 9u599N 72u439W,21KmSW
part,notLonchophyllarobustaMiller,1912.
ofMachiques,Zulia,Venezuela.Onemaleand
Lonchophylla robusta: Woodman 2006: 356 part, not
LonchophyllarobustaMiller,1912. one female (USNM 419425 and 419426)
collected by Arden L. Tuttle, Benjamin
HOLOTYPE: Skin and skull of a female
Inquilla, and Ernest L. Stromeyer (original
specimen ICN 10280 collected by M.P. Rivas
numbers 34095, and 34274) in January 1968
(original number RST409) on 9 July 1988 at
at Altamira (8u509N 70u309W), Barinas,
the intersection of can˜o Guamalito and can˜o
Venezuela. Three females (BM-NH 78-1354,
la Cur´ıa, northern part of the Serran´ıa de la
78-1356, and 78-1359) collected by Liam
Macarena, San Juan de Arama, 500 m eleva-
Hutson and R.E. Stebbins on 30 July 1976 at
tion, Departamento del Meta, Colombia.
Yaupi (2u939S 77u549W), Morona Santiago,
REFERREDSPECIMENS: (16)Onemale(ICN
Ecuador.
10278) and one female (ICN 10279) collected
by Mar´ıa del Pilar Rivas (original numbers DISTRIBUTION: Currently known from the
southeastern versant of the Cordillera de
RST041andRST076)on23and26April1988
Me´rida, the eastern versant of the Serran´ıa del
atcan˜olaCur´ıa,northernpartoftheSerran´ıa
Perija´ in Venezuela, the eastern and western
de la Macarena, San Juan de Arama, 500 m
elevation,DepartamentodelMeta,Colombia. versantsofCordilleraOrientalandthenorthern
One female (ICN 9702) collected by the portion of the Serran´ıa de la Macarena of
students of Alberto Cadena (original number Colombia, and the eastern versant of the
ACG1747) on 5 December 1985 at Colegio CordilleraOrientalofEcuador(fig. 3).
Departamental Agropecuario, vereda San ETYMOLOGY: FromtheLatinoriens(‘‘east-
Jose´, Acac´ıas, 625 m elevation, Departamento ern’’) and collis (‘‘hill’’), to summarize the
del Meta, Colombia. One female (ICN 10114) known distribution of the species in north-
collected by Alberto Cadena (original number western South America.
ACG2405)on5May1988atbocatomaCaney DIAGNOSIS: A medium-sized species of
Alto, Restrepo, Departamento del Meta, Lonchophylla (forearm 40–47 mm; weight
Colombia. One female (ICN 13839) collected 11–16 g) with dorsal fur ranging from intense
by A. Cadena (original number ACG2784) on ochraceous orange to buckthorn brown to
24 September 1995 at the forest, vereda Brisas snuff brown and light tawny olive ventral fur
6 AMERICAN MUSEUMNOVITATES NO. 3635
Fig. 1. Dorsal (A) and ventral (B) views of the skull, dorsal (C) view of the mandible, and lateral (D)
viewof the skull andmandible of theholotype of Lonchophyllaorienticollina (ICN10280).
2008 DA´VALOS AND CORTHALS: NEWSPECIES OFLONCHOPHYLLA 7
U
Fig.2. DorsalviewofLonchophyllaorienticollinaUSNM419409 (A),anddorsalviewofLonchophylla
U
robustaUSNM 419415 (B). Arrowsindicate fixed differences helpful in distinguishing these species.
(Ridgway, 1912); ventral hairs sometimes between C and P3; narrower to no gaps
bicolored,thebandingisalmostimperceptible between P3 and P4; height of P3 less than
and disappears towards the abdomen; pinnae P4;lingualcusponP4welldefined;M1longer
short with rounded tips; no furry fringe along (anteroposterior axis) than M2; M3 is the
uropatagium; calcar shorter than foot; and smallest of the molars. Lower incisors are
longfeetandthumbsrelativetobodysize.The small, with crowns taller than they are wide,
shape of the skull is distinctive: the braincase bilobed or trilobed.
is tall; the rostrum is inflated and short, Cleaned skulls are necessary to diagnose
appearing thick in profile; the palate is wide, this species from its sympatric congeners
with postpalatine torus. The height of the robusta and handleyi. Five characters are
braincasemakestheslopetotherostrumhave particularly useful in this regard: in dorsal
a relatively high angle, visible in profile. The view the rostrum appears short and wide,
spatial arrangement of teeth in the palate is inflated at the center; in profile the rostrum
diagnostic: P4 is angled outward about 15u appears thick, particularly in the postorbital
with respect to P3, making the palate appear region; the braincase appears tall and forms a
wide and almost round. There are narrow marked angle with respect to the rostrum; in
gaps between I1 and I2, outer upper incisors ventral view P4 is at about a 15u angle
I2pointventromedially;widegapsarepresent outward from P3; and this considerable
8 AMERICAN MUSEUMNOVITATES NO. 3635
Fig. 3. Map of southern Central America and northwestern South America illustrating localities for
specimens of sympatric Lonchophylla of similar size. L. handleyi is included because it has been found in
sympatry withorienticollina at Yaupi, MoronaSantiago,Ecuador.
widening makes the palate seem wide and of the genus, larger than mordax, concava,
almost round. Lonchophylla orienticollina is fornicata, thomasi, pattoni, cadenai, and de-
the only species of its size in the genus to keyseri, and smaller than handleyi, chocoana,
display this combination of traits. orcesi and most, but not all, robusta; see
MEASUREMENTS: A summary of all known table 1 (Albuja V. and Gardner, 2005; Da´va-
specimens of Lonchophylla orienticollina is los, 2004; Taddei et al., 1983; Woodman,
provided in table 1. Comparisons with a 2007; Woodman and Timm, 2006). Loncho-
representative series of other similarly sized phylla orienticollina can be unambiguously
congeners are also compiled in table 1. distinguished from the smaller Lonchophylla
DESCRIPTION AND COMPARISONS: Loncho- species on the basis of forearm length
phyllaorienticollinaisamedium-sizedmember (. 40 mm) and greatest length of skull
2008 DA´VALOS AND CORTHALS: NEWSPECIES OFLONCHOPHYLLA 9
(. 24.5 mm). Additionally, orienticollina can of the dorsal fur along the upper back in
be distinguished from mordax and thomasi chocoana is approximately 7–9 mm, slightly
based on dorsal fur coloration. The latter are longer than in robusta, handleyi, and orienti-
darkbrownbisterorsepiaofRidgway(1914), collina 4–8 mm. The ventral pelage of orienti-
whereas orienticollina exhibits the orange- collina, as in robusta, sometimes shows bicol-
brown dorsal fur coloration common in ored hairs around the neck and in the
robusta. abdominal region.
There is overlap in measurements between The cranial morphology of orienticollina is
orienticollina specimens and smaller robusta similartothatofother membersofthegenus.
individuals (table 1). Qualitative characters L. orienticollina has long rostrum relative to
are diagnostic for this comparison. In dorsal non–nectar-feedingphyllostomids,asmallbut
viewtheskulloforienticollinaappearsblunter noticeable anteorbital inflation, and a large,
and shorter, with all features shortened when relatively tall braincase (see Woodman and
compared to robusta of similar size; the Timm, 2006, for diagnosis of the genus).
rostrum clearly inflated above P4 (fig. 2). In Zygomatic arches are almost never preserved
profile, the blunt and shortened appearance after preparation, as in all other lonchophyl-
persists, with the braincase taller and the lines. As all Lonchophylla, orienticollina has a
rostrum shorter and thicker than in robusta; dental formula I2/2, C1/1, P2/3, M3/3 3 2 5
thebraincaseslopesatahigheranglefromthe 34. As found in smaller members of the genus
horizontal plane than in robusta, and in (WoodmanandTimm,2006),L.orienticollina
robusta the postorbital area appears inflated may have supernumerary premolars. The
relative to orienticollina. In ventral view, the inner upper incisors are large compared to
palate of orienticollina is wide, appearing the outer incisors and are separated by a gap
shorter than that of robusta of similar size from each other and from the canine. In this
although palatal length is not diagnostic; P4 orienticollina resembles all other species of
forms an angle about 15u with respect to P3, Lonchophylla.
wideningthepalateandmakingitlookalmost The dentition of orienticollina resembles
round compared to the rectangular appear- that of robusta, even in close detail. Both
ance of the palate of robusta. orienticollina and robusta show a tall gap
Lonchophylla orienticollina is smaller than between the inner upper incisors, meeting at
allremainingcongenersincranialdimensions. the distal quarter of the crown. The upper
Lonchophylla hesperia and L. bokermanni are caninesoforienticollinaaresimilarinabsolute
absent from the range of orienticollina, have size to those of handleyi, robusta, and thomasi
longer skulls, with greater skull length-to- whose canines are large for their body size,
width ratios, and have shorter forearms larger than those of mordax and concava, and
(Taddei et al., 1983). Lonchophylla orienticol- smaller than those of chocoana, which are
lina is smaller than chocoana and orcesi in exceptionally large. The posterior cusp of the
most dimensions (Albuja V. and Gardner, uppercaninesorienticollinaissharp,similarto
2005; Da´valos, 2004), and their distributional that of robusta, thomasi, and mordax. In
ranges do not overlap. Lonchophylla orienti- orienticollina P4 is slightly longer than P3
collinacanbedistinguishedfromitssympatric anteroposterior dimension, as it is in chocoa-
congener handleyi on the basis of size, the na,robusta,handleyi,mordax,andthomasi.P3
furry fringe along the uropatagium, the is shorter than P4 dorsoventral dimension in
definition and size of lingual cusp on P4, orienticollina, as it is in robusta, handleyi, and
presenceofaridgealongtheposterioredgeof less so in chocoana. In mordax, concava and
thepalatepostpalatinetorus,andtheshapeof thomasi P3 is taller than P4.
the palate, which is long and pointed at the Beginning with the description of
incisors in handleyi. Lonchophylla handleyi (Hill, 1980), the degree
Aswithrobusta,handleyi,andchocoana,the ofdevelopmentofthebasallingualcuspofP4
dorsal pelage of orienticollina is composed of has been used as a character to distinguish
bicolored hairs with cream-white bases and among species in this genus. After close
ochraceous orange to brown tips. The length examination of a large series of robusta,
10 AMERICAN MUSEUMNOVITATES NO. 3635
orienticollina, and specimens of chocoana,
handleyi,mordax,concavaandthomasi;differ-
ences in this character deserve further com-
ment.ThebasallingualcusponP4canbeseen
as a small protuberance in mordax, and it has
beendescribedas‘‘welldeveloped’’(Taddeiet
al., 1983: 629). This cusp is absent or
imperceptibleinconcava.Inhandleyithiscusp
has been described as ‘‘small and undevel-
oped, occasionally very small and insignifi-
cant’’ (Hill, 1981: 235), and in chocoana as
‘‘well developed’’ (Da´valos, 2004: 8). All the
handleyi and chocoana specimens examined
here had a basal lingual protuberance on P4,
but the degree of development varied from a
fullyformedcuspprojecting upward(fig. 4B),
to a small shapeless knob (fig. 4D). In both
robusta and orienticollina the basal lingual
cusp is invariably visible and distinct. There
areindividualdifferencesinthesharpnessand
height of the cusp, probably related to dental
wear. A few robusta individuals had rounded
cusps, comparable to those of some chocoana
andhandleyispecimens,thoughneverasblunt
or small (fig. 4). Individual variation might
Fig. 4. Ventral view of P4 of Lonchophylla
thus confound some pairwise comparisons
handleyi: (A) USNM 588021, (B) AMNH 230215;
aimed at identifying large Lonchophylla con- L. chocoana: (C) ROM 105798, (D) ICN 4395; L.
geners based on this character alone. In robusta: (E) ICN 13648; L. orienticollina: (F) ICN
addition to the features of P4, other charac- 10114. Note the differences in size, degree of
ters, e.g., fringe of the uropatagium, size of development, and bluntness across species and
canines, or length and shape of the palate individuals. A and C each have a cusp, but it is
blunt compared to that of E or F. B has a small
should be used for species identification.
cusp, while D has an indistinct protuberance in
As in robusta and handleyi, the M1 and M2
placeof acusp.
of orienticollina have similar widths (lateral
dimension), with M1 longer (anteroposterior
dimension) than M2, and an overall smaller and hooklike in robusta, thomasi, handleyi,
M3. In contrast, in thomasi, mordax, and chocoana, and orienticollina, with some indi-
concava the first two molars are similar in vidual variation in sharpness probably caused
length and height. bywear.Inmordaxthiscusp,ifpresent,isnot
Lowerincisorsmaybetrilobedorbilobedin clearlydistinctorhooklike.Inorienticollinap4
orienticollina, as in robusta. In handleyi, is taller (dorsoventral dimension) and longer
incisors are variably trilobed, bilobed, or (anteroposterior dimension) than p3; p3 was
neither, while both mordax and concava have erroneously reported to be taller than p4 in
trilobed incisors, though this can be difficult robusta (Da´valos 2004: 9). As in robusta, the
to detect in the latter. The height of the lower first molar of orienticollina is the widest
incisorsisgreaterthanthewidthoftheseteeth (lateral dimension), tallest, and longest of the
in orienticollina, and sometimes in robusta. molar series, followed by m2, which is in turn
Height and width of the crown of the lower wider and slightly longer than m3. The molar
incisors are roughly the same in other series varies in width and length more than in
Lonchophylla examined. The lower premolar height. As in robusta, the coronoid process in
dentition of orienticollina resembles that of orienticollina is high and oriented at an angle
robusta. The posterior cusp of p2 is distinct of about 110u with respect to the tooth row.