Systematics of Funnel-Eared Bats (Chiroptera: Natalidae)

2011·21.1 MB·English

by  Adrian Tejedor

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D SYSTEMATICS OF FUNNEL-EARED BATS
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On the cover: Portraits of two Cuban funnel-eared L
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bats: Nyctiellus lepidus (top), and Chilonatalus macer (bottom).
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With eight species, the West Indies have the highest diversity 3
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of funnel-eared bats, two of which, the critically endangered

Natalus primus (Cuba) and Natalus jamaicensis (Jamaica), may
soon become extinct in the absence of appropriate conserva-
tion efforts. 2
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1 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY
1
SYSTEMATICS OF FUNNEL-EARED BATS
(CHIROPTERA: NATALIDAE)
ADRIAN TEJEDOR
Asociacio´n para la Conservacio´n de la Cuenca Amazo´nica
Oswaldo Baca 402
Magisterio, Cusco, Peru
BULLETINOFTHEAMERICANMUSEUMOFNATURALHISTORY
Number 353,140pp., 47figures,19plates, 26tables
Issued June3,2011
E
Copyright AmericanMuseumofNaturalHistory2011 ISSN0003-0090
CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Specimens and measurements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Ecomorphological analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Statistical analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Family Natalidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Key for bats of the family Natalidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Species Account . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Genus Nyctiellus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Nyctiellus lepidus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Genus Primonatalus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Primonatalus prattae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Genus Chilonataus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Chilonatalus macer. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Chilonatalus micropus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Chilonatalus tumidifrons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Genus Natalus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Natalus espiritosantensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Natalus jamaicensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Natalus lanatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Natalus major . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Natalus mexicanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Natalus primus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Natalus stramineus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Natalus tumidirostris. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Ecomorphological diversity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Flight morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Molariform dentition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Body size. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Genital morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Appendix 1: Gazetteer and specimens examined . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
2
ABSTRACT
For much of the last third of the 20th century, the diversity of funnel-eared bats (family
Natalidae) was underestimated. Although as many as four genera and 11 species were
recognized by the mid 20th century, by the 1980s the family was traditionally listed as
comprisingasinglegenusoffourorfivespecies.Inthepastdecade,thetaxonomyofNatalidae
hasbeenupdatedbythediscoveryofnewextantandfossiltaxa,therediscoveryoftaxathought
tobeextinct,andtherevalidationofoldsynonymsatthegenusandspecieslevelsonthebasisof
newmorphological and molecular evidence.Here, I provide a complete taxonomicrevision of
Natalidae based on the examination of all taxa described thus far within the family. Thirteen
species are recognized here, including an extinct form. New diagnoses, descriptions, and
illustrations are provided for each species, including summaries of published information on
their natural history, discussions of their conservation status, and an illustrated identification
key.TheNatalidaeappearstobeanadaptivelydiversecladeofbatswithdistinctecomorphsin
termsof foodacquisition, foodprocessing,and matingsystems.
INTRODUCTION Two enigmatic taxa that have been consid-
eredsynonymsofNataluswerealsodescribed
The systematic history of Natalidae has during this time. One was Spectrellum
gone through three distinct stages common macrourum,describedbyGervais(1856)from
to most living groups (Patterson, 2001): (1) Bahia, Brazil, and of which no material or
the stage of alpha taxonomy with numerous illustrations remain. The second was Vesper-
descriptions of species and genera; (2) the tilio splendidus, a taxon apparently referable
stageofrevisionandsynonymization;and(3) to Myotis that was described by Wagner
the evaluation of interrelationships among (1845) from the island of St. Thomas. The
taxonomic units (beta taxonomy), resulting identityofS.macrourumandthereasonswhy
in the revalidation of names previously V. splendidus has continued to be considered
treatedassynonyms.Asmanyasthreeextant a synonym of Natalus are equally puzzling.
genera, one subgenus, 12 species, and six ThefirstrevisionofNatalidaewasmadein
subspecies were described in the family the late 19th century by Dobson in his
Natalidae between 1837 and 1971 (table 1). catalog of Chiroptera in the British Museum
During the last third of the 20th century, (1878).There,DobsonsynonymizedGervais’
however, only one genus and four species of Nyctiellus and Spectrellum, and Wagner’s V.
Natalidae were recognized (table 2). The splendidus(withdoubts)withGray’sNatalus.
recognized diversity of the family is greatest In addition, Dobson (1878: 342) included
at present, with four genera and 13 species, Natalus within Vespertilionidae, noting that
including an extinct form (table 2). thegenuswas‘‘evidentlytherepresentativeof
The first reference to a bat now placed in [the Old World genus] Kerivoula in the New
the family Natalidae was the description of World,’’ but that characters of the upper
Vespertilio lepidus by Gervais (1837) based incisors allied it to the family Emballonur-
on material collected in Cuba during the idae. Two years later, Dobson (1880) de-
Compte de Castelnau scientific expedition to scribed an additional species of Natalus (N.
the New World tropics. The following year, micropus), a small form from Jamaica with
Gray (1838) named a new genus and species, unique dermal outgrowths on the muzzle.
Natalus stramineus, based on a single speci- A sudden increase in the recognized
men(Goodwin,1959;HandleyandGardner, diversity of Natalidae took place during the
1990), but gave no information about the turn of the 19th century and the first few
material used, its geographic origin, or his decades of the20th. Duringthis time 11 new
reasons for choosing the generic name. In natalidtaxa,mostfromtheWestIndies,were
1840, the taxon first described as V. lepidus described, and new taxonomic arrangements
was independently renamed Vespertilio bar- were proposed. The most prolific bat taxon-
batus by Gundlach (1840) and later assigned omistofthisperiodwasGerritS.Millerwho
to the genus Nyctiellus by Gervais (1856). established Natalidae as a distinct bat family
3
4 BULLETIN AMERICAN MUSEUM OFNATURALHISTORY NO.353
TABLE 1
ChronologyoftaxonomicdescriptionsofNatalidaebytaxonomiclevelfromgenustosubspecies,including
authors,knowndistributionattimeofdescription,andcurrenttaxonomicstatus
Author Genus Subgenus Species Subspecies Distribution Currentstatus
Gervais(1837) Vespertilio Cuba distinctspecies
lepidus
Gray(1838) Natalus distinctgenus
Gray(1838) Natalus unknown distinctspecies
stramineus
Gervais(1856) Nyctiellus distinctgenus
Dobson(1880) Natalusmicropus Jamaica distinctspecies
Miller(1898) Chilonatalus distinctgenus
Miller(1898) Natalus Providencia syn.ofC.m.
(Chilonatalus) micropus
brevimanus
Miller(1900) Natalus Curacao distinctspecies
tumidirostris
Miller(1902) Natalusmajor Hispaniola distinctspecies
Miller(1902) Natalus Mexico distinctspecies
mexicanus
Miller(1903) Chilonatalus Bahamas distinctspecies
tumidifrons
Miller(1906) Phodotes syn.ofNatalus
Thomas(1910) Phodotes Venezuela ssp.ofN.
tumidirostris tumidirostris
continentis
Miller(1914) Chilonatalus Cuba ssp.ofC.
macer micropus
Anthony(1919) Natalusprimus Cuba distinctspecies
Shamel(1926) Natalus Dominica syn.ofN.
dominicensis stramineus
Dalquestand Natalus SEMexico syn.ofN.
Hall(1949) mexicanus mexicanus
saturatus
Ruschi(1951) Natalus SEBrazil distinctspecies
espiritosantensis
Goodwin(1959) Natalusmajor Jamaica distinctspecies
jamaicensis
Goodwin(1959) Natalus NEBrazil syn.ofN.
stramineus espiritosantensis
natalensis
Goodwin(1959) Natalus Trinidad ssp.ofN.
tumidirostris tumidirostris
haymani
Linares(1971) Natalus WVenezuela syn.ofN.
stramineus tumidirostris
tronchonii
Morganand Primonatalus{
Czaplewski
(2003)
Morganand Primonatalus Florida distinctspecies
Czaplewski prattae{
(2003)
Tejedor(2005) Nataluslanatus Mexico distinctspecies
Generaandspecieslevelnamecombinationsthatarecurrentlyacceptedareinboldface.{denotesanextincttaxon.
2011 TEJEDOR:FUNNEL-EARED BATS (CHIROPTERA: NATALIDAE) 5
TABLE 2
ClassificationsofextantNatalidae
Miller,1907 Dalquest,1950 Goodwin,1959 Hall,1981
Natalidae Natalidae NorthAmericanNatalidae
GenusNatalus GenusNatalus GenusNatalus GenusNatalus
N.stramineus SubgenusNatalus N.stramineusstramineus N.stramineusstramineus
N.mexicanus N.mexicanus N.stramineusnatalensis N.stramineusmexicanus
N.major mexicanus N.stramineusmexicanus N.stramineussaturatus
GenusPhodotes N.mexicanussaturatus N.majormajor N.stramineusmajor
P.tumidirostris N.major N.majorjamaicensis N.stramineusjamaicensis
GenusChilonatalus N.dominicensis N.majorprimus N.stramineusprimus
C.micropus N.primus N.tumidirostristumidirostris N.micropusmicropus
C.brevimanus N.tumidirostris N.tumidirostriscontinentis N.micropustumidifrons
C.tumidifrons SubgenusChilonatalus N.tumidirostrishaymani N.micropusmacer
GenusNyctiellus N.micropus SubgenusNyctiellus
N.lepidus N.brevimanus N.lepidus
N.macer
N.tumidifrons
SubgenusNyctiellus
N.lepidus
Koopman,1994 Simmons,2005 Thisstudy
Natalidae Natalidae Natalidae
GenusNatalus GenusNatalus GenusNatalus
SubgenusNatalus N.stramineus N.stramineus
N.stramineusstramineus N.major N.major
N.stramineusmexicanus N.jamaicensis N.jamaicensis
N.stramineustronchoni N.primus N.primus
N.stramineusnatalensis N.tumidirostris N.tumidirostris
N.stramineusespiritosantensis GenusChilonatalus N.espiritosantensis
N.stramineusmajor C.micropus N.mexicanus
N.stramineusjamaicensis C.tumidifrons N.lanatus
N.tumidirostristumidirostris GenusNyctiellus GenusChilonatalus
N.tumidirostriscontinentis N.lepidus C.micropus
SubgenusChilonatalus C.tumidifrons
N.micropusmicropus C.macer
N.micropusmacer GenusNyctiellus
N.tumidifrons N.lepidus
SubgenusNyctiellus GenusPrimonatalus
N.lepidus P.prattae
(originally including the genera Natalus, from Hispaniola; Chilonatalus tumidifrons,
Thyroptera, Furipterus, and Amorphochilus; 1903, from San Salvador Island, the Baha-
Miller,1899),anddescribedeighttaxa.These mas; and Chilonatalus macer, 1914, from
includedonesubgenus(Chilonatalus,1898,to Cuba. In additional works, Miller (1903)
include Dobson’s N. micropus), one genus elevated Chilonatalus to genus level, restored
(Phodotes, 1906), and six new species: Nata- thegenericstatusofNyctiellus(Miller,1904),
lus (Chilonatalus) brevimanus, 1898, from the andmovedthegeneraThyroptera,Furipterus,
island of Providencia, Colombia; Natalus and Amorphochilus from Natalidae to other
tumidirostris, 1900 (later transferred to genus bat families (Miller, 1907), ultimately recog-
Phodotes; Miller, 1906), from Curac¸ao, nizing four genera and eight species within
Netherlands Antilles; Natalus mexicanus, Natalidae and giving it its essential modern
1902, from Mexico; Natalus major, 1902, form (table 2). Additional taxa described by
6 BULLETIN AMERICAN MUSEUM OFNATURALHISTORY NO.353
other early 20th-century authors included a tion of only two species of Natalus (N.
new subspecies from Venezuela (Phodotes stramineus and N. tumidirostris) and raised
tumidirostris continentis Thomas, 1910) and doubts about the distinction of Chilonatalus
twoinsular species:NatalusprimusAnthony, from Natalus even at the subgenus level. In
1919, from Cuba (described from fossil Venezuela, Linares (1971) described N. stra-
remains and considered an extinct form), mineustronchoniifromSierradePerija´ onthe
andNatalusdominicensisShamel,1926,from basis of poorly cleaned skulls in which
Dominica. features of the palate used as diagnostic
TaxonomicinterestinthefamilyNatalidae characters were partially obscured by rem-
resurfaced in the second half of the 20th nant soft tissue. More importantly, Linares
century, often with somewhat contradictory (1971) suggested that all forms of Natalus
approaches: most authors reduced many (subgenus Natalus) represented the single
former names by synonymy, yet often de- species Natalus stramineus (although he
scribed new subspecies based on feeble provisionally maintained N. tumidirostris as
evidence. Dalquest and Hall (1949), for a valid name). This opinion was justified by
example,definedanewsubspeciesofNatalus the observation that all populations of
mexicanus (N. m. saturatus) from eastern Natalus (sensu stricto) showed an overall
Mexico on the basis of questionable size and continuous increase in body size beginning
pelage color differences, but Dalquest (1950) with the smallest forms in Mexico, passing
reduced all natalid genera (except Phodotes through intermediate forms in northern
which he deemed indistinguishable from South America, the Lesser and Greater
Natalus) to subgenera of Natalus. Goodwin Antilles,andendingwiththelargestinCuba.
(1959), in his revision of the subgenus Linares’ (1971) view was widely followed
Natalus, retained N. major as a distinct (e.g., by Varona, 1974; Silva-Taboada, 1979;
speciesbutreducedN.primustoasubspecies Koopman, 1993, 1994). Koopman (1981,
of N. major, N. mexicanus to a subspecies of 1993, 1994), in addition to considering N.
N. stramineus, and considered N. dominicen- stramineus a species widespread throughout
sis indistinguishable from N. stramineus theNeotropics,retainedN. tumidirostris as a
stramineus. Goodwin (1959) also described distinct species. This, therefore, necessitated
three additional subspecies: Natalus major hypothesizing that N. stramineus and N.
jamaicensis (from Jamaica), Natalus strami- tumidirostris occurred as sympatric species
neus natalensis (from Brazil), and Natalus innorthernSouthAmerica,anotionthathas
tumidirostris haymani (from Trinidad). Of survived to the present (Da´valos, 2005;
these,N.tumidirostrishaymaniwastenuously Simmons,2005;seeNoteunderN.stramineus
diagnosedbyitsslightlylargersizecompared in Systematic Account).
to other N. tumidirostris, whereas N. major WorkingintheWestIndies,Varona(1974)
jamaicensis was diagnosed by a suite of furthered the conservative trend of the
unique morphological characters that made period. First, he formalized Cabrera’s
it very distinct from N. major, illustrating an (1957) doubts about the distinctiveness of
inconsistent evaluation of the morphological Chilonatalus and did not list this name even
evidence common in the polytypic species as a subgenus. Also, he grouped all forms of
period. Chilonatalus into the single species Natalus
South American authors joined the con- micropus. Varona’s treatment was followed
servativetrendofthisperiodandsomeadded by Hall (1981), in a radical departure from
newnamestothelistofsubspeciesofNatalus his previous monograph (Hall and Kelson,
stramineus. In Brazil, Ruschi (1951) named 1959). Varona’s (1974) and Hall’s (1981)
several new bat species among which was arrangements thus represented the climax of
Myotis espiritosantensis, which he later rede- the conservative view regarding natalid di-
scribed as Natalus espiritosantensis (Ruschi, versity; Natalidae was seen as a small family
1970). Confirming the generic affinity, Pine of a single genus (Natalus) and four species:
and Ruschi (1976) reduced this taxon to a Natalus (Nyctiellus) lepidus; Natalus (Nata-
subspecies of Natalus stramineus. Cabrera lus) stramineus; Natalus (Natalus) tumidiros-
(1957), in Argentina, suggested the recogni- tris; and Natalus (Natalus) micropus.
2011 TEJEDOR:FUNNEL-EARED BATS (CHIROPTERA: NATALIDAE) 7
Althoughsomeconservativeopinionshave differentfromN.stramineus.Finally,Tejedor
survived into recent years, the past three (2006)revalidatedN.espiritosantensisandN.
decades of natalid taxonomy has been mexicanus as species distinct from N. strami-
dominatedby a shifttoward therevalidation neus, but synonymized N. saturatus with N.
of species previously considered synonyms. mexicanus. This later revision, defined N.
In 1982, Ottenwalder and Genoways argued tumidirostris as the only species occurring in
for the separation of Natalus (5 Chilonata- northern South America, a point of view
lus) tumidifrons from Natalus (5 Chilonata- sustained by Gardner (2008), and restricted
lus)micropus,onthebasisofthesmallerskull N. stramineus to the Lesser Antilles. Neither
size of the latter. A few years later, Morgan of these changes was included in the latest
(1989) offered ample evidence for the reval- worldwide list of Chiroptera (Simmons,
idation of Nyctiellus as a distinct genus and 2005).
of Natalus major as a distinct species. In Below I provide a systematic revision
2003, Morgan and Czaplewski described encompassing all described forms of Natali-
Primonatalus prattae, a fossil natalid from dae including thefossilPrimonatalus prattae.
theMioceneofnorthernFlorida,andargued An undescribed Oligocene distal radius as-
strongly for the revalidation of Chilonatalus cribed to Natalidae (Morgan and Cza-
asadistinctgenus.Inaddition,Tejedoretal. plewski,2003)andtheEocenefossilsAegina,
(2004)announcedtherediscoveryofaCuban Chadronycteris, Chamtwaria, Honrovits, and
taxon, Natalus primus, previously thought Stehlinia, excluded from Natalidae by Mor-
extinct and long synonymized with Natalus gan and Czaplewski (2003) are not included
stramineus. On the basis of morphological in this account.
evidence, Tejedor et al. (2005b) revised the
taxonomy of Greater Antillean Natalus,
METHODS
revalidating N. primus and elevating Natalus
jamaicensis to the status of fully distinct SPECIMENSANDMEASUREMENTS:Forthis
species. The separation of N. jamaicensis study I examined 3234 specimens of Natalus
from N. major was corroborated by Da´valos (including all 20 holotypes in the family).
(2005) on the basis of gene sequences. In These specimens are housed in 26 museums,
addition, Da´valos (2005) demonstrated a all of which are listed (including abbrevia-
deepgeneticdivergencebetweenChilonatalus tions) in appendix 1.
and Natalus, supporting the generic distinc- Morphological terminology follows Gian-
tion of the former. Such sweep of old-name nini et al. (2006) for the cranium, Smith
revalidations, however, was countered by (1972) and Morgan and Czaplewski (2003)
Timm and Genoways (2003), and Genoways for the postcranium, Freeman (1998) for the
et al. (2005) who maintained that all greater dentition, and Ridgway (1912) for pelage
Antillean Natalus are not distinct from N. coloration. A total of 1137 specimens were
stramineus at the species level, and retained measured for this study, of which eight were
Chilonatalus as a synonym of Natalus. juveniles or young adults. Additional mea-
Simmons (2005) nonetheless summarized surements were taken from Pedersen et al.
the rise in recognized diversity of Natalidae (2003), Lo´pez-Gonza´lez et al. (1998), Taddei
listingitascomprisingthreegeneraandeight andUieda(2001),Silva-Taboada(1979),and
species (table 2). Goodwin (1959). Length of forearm and
Mostrecently,thesystematicsofNatalidae length of tibia were measured from study
hascontinuedtochangethroughthedescrip- skins, fluid-preserved specimens, and fresh
tion or revalidation of additional species. In specimens, whereas ear length was measured
2005, Tejedor described Natalus lanatus, a exclusively from fluid-preserved and fresh
species from Mexico that had remained specimens. Given that measurements of
undetected in museum collections for length of tibia taken from fluid-preserved
140years.Da´valos(2005),ontheotherhand, specimenshavebeenfoundtobesignificantly
elevatedthenameN.saturatustospecieslevel larger on average than those taken from dry
to designate the Mexican–Central American skins (Tejedor, 2005), means of length of
populations, which she found genetically tibia taken by those two measures are
8 BULLETIN AMERICAN MUSEUM OFNATURALHISTORY NO.353
reported separately. Lineal measurements Maxillary toothrow length: Greatest crown
were taken with dial and digital calipers and length of the maxillary tooth row, measured
are given in millimeters. Measurements of from the anteriormost edge of the canine to
weight were taken from specimen tags and the posteriormost edge of M3.
are given in grams. Lineal measurements are Mandibular toothrow length: Greatest
defined as follows: crown length of the mandibular tooth row,
Forearm length: Distance from the elbow measured from the anteriormost edge of the
to the distal end of the forearm including canine to the posteriormost edge of m3.
carpals,takenwiththewing atleastpartially Mental length: Least distance from the
folded. mental foramen to the ventral margin of the
Ear length: Maximum distance between dentary between the angular process and the
the ventral point of attachment of the pinna condyle (used only for Natalus primus, a
and the tip of the pinna. species for which numerous fossil specimens
Length of tibia: Distance from the proxi- had incomplete mandibular tooth rows).
mal end of the tibia to the posterior base of Depth of braincase: Distance between the
the calcar (in dry specimens), and distance basioccipito-palatalplaneandthedorsalmost
from the knee to the ankle with both knee edge of the sagittal crest.
and ankle bent to an angle of at least 90u (in Cranial measurements (except mental
fluid-preserved specimens). length) are illustrated in figure 1.
Length of 3rd metacarpal: Distance from In addition, to graphically represent inter-
wrist to distal end of 3rd metacarpal, taken specific variation in cranial shape, several
with the wing folded. indices were calculated: (1) relative breadth
Length of 5th metacarpal: Distance from across canines 5 breadth across canines/
wrist to distal end of 5th metacarpal, taken greatest skull length; (2) relative breadth
with the wing folded. across molars 5 breadth across molars/
Length of penis: Maximum distance from maxillary toothrow length; (3) relative max-
the caudal base to the tip of the penis, illary toothrow length 5 maxillary toothrow
without stretching. length/ greatest skull length; (4) relative
Length of natalid organ: Maximum dis- breadth of postorbital constriction 5 post-
tance from the rostral margin to the caudal orbital breadth/zygomatic breadth.
marginofthenatalidorgan(inNyctiellusand ECOMORPHOLOGICAL ANALYSES: Eco-
Chilonatalus) or from the dorsal point of morphological traits were inferred from a
flexion between rostrum and forehead to the combination of measurement data and field
caudal margin of the natalid organ (in observations. Measurements of forearm
Natalus), taken externally. length, length of tibia, and 3rd and 5th
Greatest skull length: Maximum distance metacarpalswereusedasdescriptorsofflight
between the anteriormost point of the skull, surface shape. In all natalids, the uropata-
including the incisors, and the posteriormost gium is roughly wedge shaped and its free
point of the occiput. marginextendsfromneartheankletothetip
Zygomatic breadth: Greatest distance of the tail; therefore, its extension depends
across the external margins of the zygomatic mostlyon thelength of thelegs and tail.For
processes. thisreason,thelengthofthetibiawasusedas
Braincase breadth: Greatest breadth of the an indicator of the size of the uropatagium.
globular part of thebraincase, taken perpen- Two indices were calculated to represent
dicular to the skull’s longitudinal axis. shapeofflightsurfaces:(1)shapeofthewing
Postorbital breadth: Least breadth across tip(5 length of 5th metacarpal/lengthof 3rd
the postorbital constriction. metacarpal) and (2) relative size of the
Breadth across molars: Greatest distance uropatagium (5 length of tibia/forearm
between the labial edges of the upper molar length). In addition, two indices were calcu-
series. lated to represent relative size of organs
Breadth across canines: Greatest distance related (or potentially related) to reproduc-
betweenthelabial edgesofthecingulaofthe tion in males: (1) relative length of the penis
upper canines. (5 length of penis/forearm length) and (2)
2011 TEJEDOR:FUNNEL-EARED BATS (CHIROPTERA: NATALIDAE) 9
tion is made between N. primus and the
remaining species of Natalus, which are
hereinaftertreatedasthe‘‘Natalusstramineus
group.’’ Similarly, a distinction is made
between Chilonatalus micropus and a Chilo-
natalus ‘‘tumidifrons group’’ (5 C. tumidi-
frons + C. macer).
In figures, tables, and plates, species are
abbreviated as follows: Cma, Chilonatalus
macer; Cmi, Chilonatalus micropus; Ctu,
Chilonatalus tumidifrons; Nes, Natalus espir-
itosantensis; Nja, Natalus jamaicensis; Nla,
Natalus lanatus; Nma, Natalus major; Nme,
Natalus mexicanus; Npr, Natalus primus;
Ntu, Natalus tumidirostris; and Nst, Natalus
stramineus.
STATISTICAL ANALYSES: I used two-way
ANOVAstodetectoveralldifferencesamong
species and populations (geographic subsets
within a given species) and between sexes, as
well as interaction effects between popula-
tions/species and sex. Where interaction
effects were not found, one-way MANOVAs
(performedoverallvariablessimultaneously)
were used to detect sexual differences within
species. I also used one-way MANOVAs
(performedoverallvariablessimultaneously)
todetectdifferencesbetweentwopopulations
of one species within a single sex when only
one sex was well represented and sexual
differences in that species had been detected
Fig. 1. Cranial dimensions measured in this with previous ANOVAs. In species where
study: A, postorbital breadth; B, zygomatic simultaneous one-way MANOVAs were im-
breadth; C, braincase breadth; D, breadth across practical due to unequal distribution of
canines; E, breadth across molars; F, maxillary missing measurements, one-way ANOVAs
toothrowlength;G,greatestskulllength;H,depth
were performed separately for each variable.
ofskull; I, mandibulartoothrow length.
Post hoc comparisons (Tukey tests for
unequal sample sizes) tested for pairwise
differences among species or populations.
relativelengthofthenatalidorgan(5length ThesignificancelevelwassettoP50.05for
of natalid organ/forearm length). Forearm theTukeytestsandtheone-wayMANOVAs
length was used to define three body size inwhichtestswereperformedsimultaneously
classes: (1) large (mean forearm length 43.5– over the sampling units. The significance
49 mm), including Natalus major, Natalus level was set to P 5 0.01 for all the two-way
jamaicensis, and Natalus primus; (2) medium ANOVAs and the one-way ANOVAs in
(mean forearm length of 37–39 mm), includ- whichmultipleseparatetestswereperformed
ing the remaining species of Natalus; and (3) over the same sampling units. I used canon-
small (mean forearm length of 29–34 mm), ical analyses to detect differences in body
including Nyctiellus and Chilonatalus. Be- dimensions among species and among geo-
cause of the outlying position of Natalus graphical samples within each species. All
primus relative to other Natalus with respect analyseswereperformedusingSTATISTICA
to several ecomorphological traits, a distinc- 5.5 (StatSoft, Inc., 1995).