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PHYLOGENETIC RELATIONSHIPS OF
MORMOOPID BATS (CHIROPTERA:
MORMOOPIDAE) BASED ON
MORPHOLOGICAL DATA
NANCY B. SIMMONS
Associate Curator
Division of Vertebrate Zoology (Mammalogy)
American Museum of Natural History
TENLEY M. CONWAY
Scientific Assistant
Division of Vertebrate Zoology (Mammalogy)
American Museum of Natural History
BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY
Number 258, 97 pp., 12 figures, 4 tables
Issued February 15, 2001
Price: $10.10 a copy
Copyright(cid:1)AmericanMuseumofNaturalHistory2001 ISSN0003-0090
2 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 258
CONTENTS
Abstract ....................................................................... 3
Introduction .................................................................... 3
Historical Background ......................................................... 3
Mormoopid Species: A Synopsis ............................................... 8
Goals of the Present Study .................................................... 12
Materials and Methods ......................................................... 12
Taxonomic Sampling, Outgroups, and Tree Rooting .............................. 12
Sources of Data ............................................................. 13
Definition of Characters and Ordering of Character States ........................ 13
Polarity ..................................................................... 15
Completeness ............................................................... 15
Methods of Phylogenetic Analysis ............................................. 16
Character Descriptions .......................................................... 17
Skull ....................................................................... 17
Dentition ................................................................... 24
Vomeronasal Complex and Brain .............................................. 29
Trachea and Hyoid Apparatus ................................................. 30
Tongue ..................................................................... 35
Face, Ears, and Vibrissae ..................................................... 36
Pelage and Patagia ........................................................... 42
Postcranial Skeleton .......................................................... 45
Postcranial Myology ......................................................... 58
Reproductive Tract ........................................................... 65
Digestive Tract .............................................................. 66
Results ....................................................................... 68
Discussion and Conclusions ..................................................... 73
Intrafamilial Relationships of Mormoopids ...................................... 73
Classification and Taxonomic Diagnoses of Mormoopid Clades ................... 74
Monophyly and Diagnoses of Noctilionoid Families ............................. 81
Interfamilial Relationships .................................................... 83
Directions for Future Research ................................................ 84
Acknowledgments ............................................................. 85
References .................................................................... 85
Appendix 1: Specimens Examined ............................................... 93
Appendix 2: Taxon-Character Matrix ............................................. 96
2001 SIMMONS AND CONWAY: PHYLOGENY OF MORMOOPID BATS 3
ABSTRACT
Mormoopidae is a small family of Neotropical microchiropteran bats that includes two
genera(MormoopsandPteronotus)andtenspecies,twoofwhichareknownonlyfromfossils.
Mormoopidae is typically classified as a member of Noctilionoidea, a group that minimally
includes two other Neotropical families (Phyllostomidae and Noctilionidae) and may also
include Mystacinidae, a taxon endemic to New Zealand and Australia. Phylogenetic relation-
ships of extant mormoopid species and one extinct taxon, Pteronotus pristinus, were investi-
gated in a series of parsimony analyses of 209 morphological characters including featuresof
the skull, dentition, vomeronasal organ complex and brain, trachea and hyoid apparatus,
tongue, face, ears, pelage, patagia, postcranial skeleton, postcranial myology, reproductive
tract,anddigestivetract.Threeextantphyllostomidspecies,twonoctilionids,twomystacinids,
and one emballonurid species were included as outgroupstotestmonophylyofMormoopidae
and to provide a context for determining the sister group of the family. Results of parsimony
analyses under a variety of different assumption sets indicate that Mormoopidae is monophy-
letic, and that Pteronotus and Mormoops are monophyletic sister taxa. Within thegenusPter-
onotus, several clades were repeatedly recovered: (1) P. davyi (cid:1) P. gymnonotus ((cid:2) subgenus
Pteronotus); (2) P. macleayi (cid:1) P. quadridens; (3) P. personatus (cid:1) P. macleayi (cid:1) P. quad-
ridens ((cid:2) subgenus Chilonycteris); (4) P. parnellii (cid:1) P. pristinus (cid:1) P. personatus (cid:1) P.
macleayi (cid:1) P. quadridens; and (5) P. parnellii (cid:1) P. pristinus ((cid:2) subgenus Phyllodia).These
results support monophyly of all subgenera of Pteronotus previously recognized, and addi-
tionally indicate that the subgenera Pteronotus and Chilonycteris are sister taxa. Comprehen-
sive diagnoses for each species and clade of Mormoopidae are provided based on character
optimizations and ancillary morphometric data from the literature.
Resultsof our parsimony analysesalsohaveimplicationsforunderstandinghigher-levelphy-
logeny of noctilionoid bats. Monophyly of each of the traditionally recognized families (i.e.,
Noctilionidae,Mystacinidae,Mormoopidae,andPhyllostomidae)wasstronglysupported.Incon-
gruence with recent analyses of mitochondrial gene-sequence data and DNA hybridization ex-
periments, we found strong support for inclusion of Mystacinidae in Noctilionoidea. Using an
emballonurid species to root the tree, we found the following interfamilialrelationshipsofnoc-
tilionoids: (Noctilionidae (Mystacinidae (Phyllostomidae, Mormoopidae))). Lists of morpholog-
ical synapomorphies of each of these groups are provided based on character optimizations.
INTRODUCTION tropicalforesttosemiaridandaridsubtropical
forest and scrubland (Handley, 1976; Em-
Mormoopidae is a small family of Neo-
mons, 1997; Reid, 1997; Smith, 1972). The
tropical microchiropteran bats that currently
most recent revision of Mormoopidae was
includes two genera (Mormoops and Pteron-
that of Smith (1972), which has formed the
otus) and eight extant species; two additional
basis for most modern treatments of the fam-
species are known only from Quaternaryfos-
ily.
sils (Smith, 1972;Silva-Taboada,1974,1979;
The history of classification and nomen-
Koopman,1993,1994).Commonlyknownas
clature of mormoopids is complex and often
mustached, ghost-faced, or naked-backed
confusing. Smith (1972) provided a compre-
bats, these taxa are characterized by the pres-
hensive systematic literature review for the
ence of flaplike outgrowths below the lower
family, which we will not reproduce here. A
lip and funnel-shaped ears. Mormoopids are
few critical points from the pre-1970 litera-
small to medium in size (e.g., forearm length
ture, together with more recent taxonomic
35–66 mm) and are thought to be exclusively
changes and phylogenetic inferences, are
insectivorous (Koopman, 1984, 1994). Mem-
summarized below.
bers of the family presently range from the
southwesternUnitedStatestosouthernBrazil, HISTORICAL BACKGROUND
and also occur in the Greater Antilles, Lesser
Antilles, Dutch West Indes, and Trinidad and CLASSIFICATION
Tobago (Koopman, 1993, 1994). They live in The taxonomic history of Mormoopidae
awidevarietyofhabitatsrangingfromhumid began with Leach (1821a, 1821b), who de-
4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 258
scribedMormoopsandAello.Pteronotusand sification did not entirely match his phylo-
Chilonycteris were subsequently described genetic tree (fig. 1), which depicted Pteron-
by Gray (1838, 1839), and Lobostoma was otus personatus as more closely related toP.
named by Gundlach (1840). Gray (1843) davyiandP.suapurensisthantoP.macleayii
named a sixth genus, Phyllodia, followed al- and P. fuliginosus. However, Smith’s (1972)
most 60 years later by Gill’s (1901) propo- revision wascompletedinaprecladisticcon-
sition of Dermonotus as a replacement name text, so this discrepancy was not widely rec-
for Pteronotus. Various authors have pro- ognized. Subsequent subgeneric classifica-
posed different synonomies for these taxa tions (e.g., Corbet and Hill, 1980; Herd,
over the last 150 years, and misplaced ho- 1983; Koopman, 1994) retained Smith’s
lotypes, overlooked names, and arguments (1972) subgeneric usage.
over nomenclatural priority have served to Although Smith’s (1972) classification of
further complicate matters (see review in Mormoopidae survived more-or-less intact
Smith, 1972). For most of the 20th century, into the 1990s, several significant nomencla-
classifications of Mormoopidae were based tural changes have occurred since that pub-
on that of Miller (1907), who recognized lication. Silva-Taboada (1976) showed that
three genera (Chilonycteris, Pteronotus, and Lobostoma quadridens Gundlach, 1840 is a
Mormoops), which he placed in the subfam- senior synonym of Chilonycteris fuliginosus
ilyChilonycterinaeinthefamilyPhyllostom- Gray,1843,soPteronotusquadridensisnow
idae. Dalquest and Werner (1954) recom- recognized as the correct name for this spe-
mendedelevationofthegrouptofamilyrank cies. Similarly, Smith (1977) subsequently
as Chilonycteridae, but this suggestion was demonstrated that Chilonycteris gymnonotus
overlooked or ignored by most authors prior Wagner, 1843, is a senior synonym of Der-
to Smith’s (1972) revision. Smith (1972) monotus suapurensis J. A. Allen, 1904, so
agreed that this group deserved familial sta- Pteronotus gymnonotusisnowrecognizedas
tus, but argued that the correct name for this the correct name for this species (see Carter
taxon is Mormoopidae. Most subsequent au- and Dolan [1978] for discussion of author-
thors have followed this recommendation. A ship of gymnonotus). In addition, two fossil
lone dissenter was Hall (1981), who contin- mormoopid species, Mormoops magna and
ued to recognize Chilonycterinae as a sub- Pteronotus pristinus, were described from
family of Phyllostomidaebasedontaxonom- Cuban cave deposits by Silva-Taboada
ic history and his perception that the ‘‘grade (1974). Partial or complete synonomies for
of differentiation’’ of this group was similar all currently recognized species and subspe-
to that of ‘‘some other groups of mammals cies of mormoopids can be found in Smith
currently treated as subfamilies.’’ These ar- (1972, 1977), Silva-Taboada (1976, 1979),
guments did not convince the rest of thesys- Herd (1983), Adams (1989), Rodr´ıguez-Dur-
tematic community, and the group has been a´n and Kunz (1992), Koopman (1993), and
listed under Mormoopidae in most classifi- Rezsutek and Cameron (1993).
cations published since 1972 (e.g., Corbet
and Hill, 1980;Koopman,1984, 1993,1994; PHYLOGENETIC RELATIONSHIPS
McKenna and Bell, 1997; Simmons, 1998;
Simmons and Geisler, 1998). ThefirstexplicitphylogenetictreeofMor-
Smith(1972)recognizedtwogenera,three moopidae was that of Smith (1972; fig. 1).
subgenera, and eight extant species within AlthoughSmithlaterbecamewellknownfor
Mormoopidae. Mormoops was shown to in- his applications of cladistic methodology to
clude two species, M. blainvillii and M. me- bat systematics (e.g., Smith, 1976, 1980;
galophylla (Smith, 1972). Smith (1972) di- Smith and Madkour, 1980; Hood and Smith,
vided Pteronotus into three subgenera: Phyl- 1982, 1983), his tree of mormoopid relation-
lodia (including only Pteronotus parnellii), ships was developed in a precladisticcontext
Chilonycteris (including Pteronotus macle- using methods of numerical taxonomy
ayii, P. fuliginosus, and P. personatus), and (Smith, 1972). This phenetic tree assumed
Pteronotus (including P. davyi and P. sua- monophyly of the family, a hypothesis that
purensis). Interestingly, Smith’s (1972) clas- he supported based on morphological com-
2001 SIMMONS AND CONWAY: PHYLOGENY OF MORMOOPID BATS 5
Fig.1. Smith’s(1972)phylogenetictreeofmormoopidspecies,withthetaxonomychangedtoreflect
modern usage (i.e., P. quadridens (cid:2) P. fuliginosus, P. gymnonotus (cid:2) P. suapurensis). This tree, which
was developed in a pre-cladistic context using methods of numerical taxonomy, was derived from
analysis of 22 skull measurements and 20 qualitative multistate characters of the dentition and external
morphology. The horizontal and vertical spacing of various branches was intended to depict degrees of
similarity and differences among taxa. Redrawn from Smith (1972: fig. 14).
parisons to members of other chiropteran group relationship between Mormoopidae
groups. In large part due to the thoroughness and Noctilionidae was proposed based on
of Smith’s (1972) study, this tree was very five inferred Robertsonian fusion events
influential in shaping subsequent classifica- shared by these taxa(fusionofchromosomes
tions of mormoopids. 21/14, 17/9, 13/8, 22/3, and 18/2; Patton and
Karyotypic studies including mormoopid Baker, 1978). All of these results were con-
specieswerecompletedinthelate1970sand gruent with Smith’s (1972) phylogenetic hy-
early 1980s (e.g., Patton and Baker, 1978; pothesis.
BakerandBickham,1980;Sitesetal.,1981). Limited immunological studies conducted
All species of Pteronotus and Mormoops around the same time by Honeycutt (1981)
werefoundtohavethesamegrosskaryotype produced divergent results. Analysis of al-
(2N (cid:2) 38, FN (cid:2) 60), and banding compar- bumin immunological distance data for a set
isons indicated that they share a unique fis- of taxa including Mormoops megalophylla,
sion of chromosome 6/7 that occurs in no Pteronotus parnellii, Noctilio leporinus, a
other bat family (Patton and Baker, 1978; mixed sample of several phyllostomid spe-
BakerandBickham,1980;Sitesetal.,1981). cies, and a mixed sample of pteropodids
All extant species of Pteronotus share an (used to root the tree) resulted in a tree in
identical, unique G-band karyotype, thus which Mormoopidae was paraphyletic with
supporting monophyly of the genus (Sites et respect to Phyllostomidae. However, poor
al., 1981). Monophyly of Mormoops was sampling within Mormoopidae and inconsis-
also supported by banding studies, and only tencies in the data led Honeycutt (1981) to
a single difference in banding pattern was exercise caution in interpreting his results,
found between the karyotypes of Mormoops which he regarded as inconclusive.
and Pteronotus (Sites et al., 1981). A sister- In one of the first papers to explicitly ad-
6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 258
dress the problem of combining disparate four data sets (fig. 2). Within Mormoopidae,
data sets in phylogenetic studies, Arnold et the relationships depicted were completely
al. (1982) reviewed availablemorphological, compatible with Smith’s (1972) tree (fig. 1).
immunological, allozyme, and karyotypic However, the Arnold et al. (1982) tree did
data relevant to mormoopid relationships. not include all mormoopid species (Pteron-
Phylogenetic trees for each data set werede- otus gymnonotus was excluded), and rela-
rived by hand byapplyingparsimonycriteria tionships among species of Pteronotus were
to the discrete data sets (morphology, kar- poorly resolved.
yotypes, and allozymes) and distance meth- No new comprehensive phylogeny of
ods to the immunologicaldata(Arnoldetel., Mormoopidae has been published since
1982). The karyotype tree presented by Ar- 1982. However, several studies have ad-
nold et al. (1982: fig. 3) was based on data dressed the relationships of mormoopids to
from PattonandBaker(1978)andBakerand other families. Pierson (1986) and Pierson et
Bickham (1980), which supportedmonophy- al. (1986) proposed a phylogeny of bats
ly of a Mormoopidae (cid:1) Noctilionidae clade, based on a study of transferrin immunologi-
Mormoopidae, and Pteronotus.1 The immu- caldistancedata.TheyfoundthatMormoop-
nological tree was taken from Honeycutt idae, Noctilionidae, and Mystacinidae
(1981), which indicated paraphyly of Mor- formed a clade, with Phyllostomidae as a
moopidae with respect to Phyllostomidae, very close relative to this group. More than
with Noctilionidae as the sister group to a decade later, Kirsch et al. (1998) found
these taxa. The morphology tree, which was verystrongsupportformonophylyofaMys-
based on 16 morphological characters de- tacinidae (cid:1) Noctilionidae (cid:1) Mormoopidae
rived from Smith (1972), supported mono- (cid:1) Phyllostomidae clade in a study of DNA
phyly of Mormoopidae, Mormoops, Pteron- hybridization data. Within this group, mono-
otus, andacladeconsistingofP.personatus, phylyofacladecomprisingMormoopidae(cid:1)
P. quadridens, P. macleayii, and P. davyi Phyllostomidae was also very strongly sup-
(Arnold et al., 1982: fig. 2). Mormoopidae, ported, although only one mormoopid (Pter-
Noctilionidae, and Phyllostomidae together onotus parnellii) was included in the analy-
formed a monophyletic clade Phyllostomo- sis. Relationships of this clade to Noctilion-
idea ((cid:2) Noctilionoidea) in the morphology idae and Mystacinidae remained poorly re-
tree, which was rooted using emballonurids solved (Kirsch et al., 1998).
(Arnold et al., 1982: fig. 2). Finally, Arnold Simmons (1998) and Simmons and Geis-
etal.(1982)presentedanewdatasetderived ler (1998) found support for somewhat dif-
from allozyme studies of 16 populations of ferent relationships in a series of analyses of
mormoopids and 4 populations of noctilion- higher-level bat relationships based on mor-
ids. A cladogram derived from these data phological data. Simmons (1998) and Sim-
supported monophyly of Mormoops, Pter- mons and Geisler (1998) found strong sup-
onotus, and a clade consisting of P. person- port for Noctilionoidea as traditionally rec-
atus, P. quadridens, P. macleayii, and P. ognized (Mormoopidae (cid:1) Phyllostomidae (cid:1)
davyi (Arnold et al., 1982: fig. 1). Noctilionidae). Within this group, weak sup-
Arnold et al. (1982) concluded their paper portwasfound for asister-grouprelationship
by presenting a summary ‘‘composite clad- between Mormoopidae and Noctilionidae in
ogram’’thatwasconstructedusingprinciples both analyses. In each case, Mystacina was
of taxonomic congruence and majority rule tentatively placed as the most basal branch
consensus. The incompatible immunological of the sister clade to Noctilionoidea. How-
data were discounted for a variety of meth- ever, support for this placement was weak,
odologicalreasons,andallnodesinthecom- and the authors noted that several other po-
positecladogramwereultimatelyfoundtobe sitions for Mystacinidae were only slightly
supported by between one and three of the less parsimonious given their data.
DNA sequence data are as yet unavailable
for most mormoopids, but two studies have
1Arnold et al. (1982) completed their study before
addressedrelationshipsofmormoopidsinthe
publicationofSitesetal.(1981),sotheywerenotaware
ofthekaryotypicevidenceformonophylyofMormoops. context of projects designed to resolve the
2001 SIMMONS AND CONWAY: PHYLOGENY OF MORMOOPID BATS 7
Fig. 2. Arnold et al.’s (1982) ‘‘composite cladogram’’ of relationships among noctilionoid species.
This phylogeny was derived from taxonomic-congruence comparisons of trees derived from four data
sets: albumin immunology, chromosomes, allozymes, and morphology. Data sets supporting each clade
are indicated on the tree. Redrawn from Arnold et al. (1982: fig. 5).
phylogenetic position of Mystacina relative support for mormoopid monophyly was es-
to other bat families. Kennedy et al. (1999) sentially nonexistent. Kennedy et al. (1999)
used mitochondrial cytochrome-b gene se- additionally found strong support for place-
quences to investigate relationships among ment of Mystacina withinNoctilionoidea,al-
34 species including Mormoops megalophyl- though the exact position of this taxon rela-
la, Pteronotus parnellii, and P. davyi. The tive to Phyllostomidae, Noctilio, and mor-
most surprising result of this study was the moopids varied under different analysis pa-
finding that Mormoopidae is not monophy- rameters.
letic; Noctilio was placed as the sister group Results of another study of mitochondrial
ofPteronotusinallanalyses,withMormoops genesequencesproducedsomewhatdifferent
falling outside that clade (Kennedy et al., results. Van Den Bussche and Hoofer (2000)
1999).Supportforthisarrangementwasvery analyzed DNA sequence data from three ad-
high in bootstrap and decay analyses of a jacent mitochondrial genes (12S rRNA, t-
weighted data set (codon positions weighted RNAVal, and 16S rRNA) sampled in 11 noc-
5:16:1, transversions/transitions 2:1), while tilionoids and 17 outgroup species. Results
8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 258
of unweighted parsimony analysessupported (1974), Wilkins (1983), Steadman et al.
monophyly of NoctilionoideaincludingMys- (1984), Eshelman and Morgan (1985), Mor-
tacina, and monophyly of each of families gan and Woods (1986), and Morgan (1989,
within this group. Mormoopidae and Phyl- 1991). Tertiary mormoopid fossils are as yet
lostomidae were found to be sister taxa, unknown (Czaplewski, 1997; McKenna and
echoing the results of Kirsch et al. (1998). Bell, 1997).
Mystacinidae was placed as the sister group Mormoops blainvillii Leach, 1821 (Antil-
to the Mormoopidae (cid:1) Phyllostomidae lean ghost-faced bat) is a medium-smallspe-
clade. Mormoopidae, represented by Mor- cies (forearm length (cid:2) 43–50 mm; condy-
moops megalophylla and Pteronotus parnel- lobasal length (cid:2) 12–14 mm; 6–11 g) pres-
lii,wasfoundtobemonophyleticin71–88% ently restricted to the Greater Antillean is-
of bootstrap replicates (higher values were lands of Cuba, Jamaica, Hispaniola, Puerto
obtained using successive approximations Rico, and adjacent small islands (Silva-Ta-
weighting). boada, 1979; Koopman, 1993,1994;Lancas-
ter and Kalko, 1996; Emmons, 1997). No
MORMOOPID SPECIES: A SYNOPSIS subspecies are currently recognized (Smith,
1972; Hall, 1981; Koopman, 1994). Fossils
The principal reference on morphology of M. blainvillii have been reported from
and distribution of mormoopid species is caves in the Bahamas, Antigua and Barbuda
Smith (1972), but this resource is over 25 in the northern Lesser Antilles, and La Gon-
years old and it lacks any substantive con- ave off the west coast of Hispaniola (Koop-
sideration of ecology or the fossil record. To man, 1951, 1955, 1989; Steadman et al.,
help fill these gaps, weprovidebelowabrief 1984; Morgan and Woods, 1986; Morgan,
synopsis of each mormoopid species includ- 1989), indicating that the Late Quaternary
ing information on presentlyrecognizedsub- rangeofthisspecieswasmoreextensivethan
species, geographic range, fossilrecord,hab- it is at present.
itat preferences, roosting habits, and dietary Like other mormoopids, M. blainvillii
habits.Literaturecitationsareprovidedtofa- roosts in hot, humid caves (i.e., those with
cilitate access to the primary literature. For temperatures 26–40(cid:3)C; Goodwin, 1970; Sil-
identification purposes, readers should con- va-Taboada, 1979; Rodrı´guez-Dura´n and
sult keys to genera and extant species of Lewis, 1987; Rodr´ıguez-Dura´n, 1995, 1998;
mormoopids published in Smith (1972),Hall Lancaster and Kalko, 1996). Colonies may
(1981), Herd (1983), Eisenberg (1989), and number over 40,000 individuals (Rodr´ıguez-
Rodr´ıguez-Dura´n and Kunz (1992). Another Dura´n and Lewis, 1987). Moths are by far
helpful resource is Koopman (1994), which themostcommoniteminthediet,whichalso
includes measurements and some diagnostic includes some beetles, flies, homopterans,
features of each extant species. Useful sum- and hemipterans (Silva-Taboada, 1979; Rod-
maries of morphological and ecological data r´ıguez-Dura´n and Lewis, 1987; Rodr´ıguez-
for various species were provided by Bate- Dura´n et al., 1993).
man and Vaughan (1974), Silva-Taboada Mormoops megalophylla Peters, 1864
(1976, 1979), Herd (1983), Adams (1989), (Peters’ ghost-faced bat; Ghost-faced bat;
Eisenberg (1989), Rodrı´guez-Dura´n and Leaf-chinned bat) is a medium-large species
Kunz (1992), Rezsutek and Cameron(1993), (forearm length (cid:2) 49–61 mm; condylobasal
Emmons (1997), and Reid (1997). Maps length(cid:2)12–14mm;12–20g)thatisbroadly
showing the geographic ranges of extant distributed in the Caribbean and Central and
populations can be found in Hall (1981), South America (Koopman, 1993, 1994; Em-
Koopman (1982), Herd (1983), Adams mons, 1997; Reid, 1997). Four subspecies
(1989), Eisenberg (1989), Rodr´ıguez-Dura´n are currently recognized (Smith, 1972; Rez-
and Kunz (1992), Rezsutek and Cameron sutek and Cameron, 1993; Koopman, 1994):
(1993), Lancaster and Kalko (1996), and M. m. megalophylla (Baja California, south-
Reid (1997). Information on Quaternaryfos- ern Arizona, and southern Texas south to
sils can be found in Koopman (1951, 1955, Honduras); M. m. tumidiceps (northern Co-
1958,1989),Rayetal.(1963),SilvaTaboada lombia,northernVenezuela,MargaritaIsland
2001 SIMMONS AND CONWAY: PHYLOGENY OF MORMOOPID BATS 9
and Trinidad); M. m. intermedia (Aruba,Cu- Pteronotus parnellii Gray, 1843 (Par-
rac¸ao, and Bonaire islands), and M. m. car- nell’s mustached bat; Mustached bat) is a
teri(coastalEcuadorandnorthwesternPeru). medium-sized species(forearmlength(cid:2)48–
Pleistocene fossils of M. megalophylla have 66 mm; condylobasal length (cid:2) 16–22 mm;
been found in Florida, Mexico, Cuba, His- 10–28 g) that is broadly distributed from the
paniola, Jamaica, the Bahamas (Andros), Greater Antilles and tropical Mexico to
Aruba, Curac¸ao, Margarita Island, Trinidad, northeastern Brazil, although it does not oc-
Tobago, and southeastern Brazil (Koopman, curwestoftheAndesinSouthAmerica(Sil-
1958, 1989; Ray et al., 1963; Silva-Taboada, va-Taboada, 1979; Koopman, 1993, 1994;
1974; Wilkins, 1983; Eshelman and Morgan, Emmons, 1997; Reid, 1997). Nine subspe-
1985; Morgan and Woods, 1986; Morgan, cies are currently recognized (Smith, 1972;
1989, 1991; Arroyo-Cabrales and Alvarez, Linares and Ojasti, 1974; Herd, 1983; Koop-
1990; Arroyo-Cabrales, 1992; Czaplewski man, 1994): P. p. parnellii ((cid:2) boothi; Cuba
and Cartelle, 1998). The records from Flor- and Jamaica); P. p. pusillus (Hispaniola); P.
ida, the Bahamas, the Greater Antilles, and p. gonavensis (Gonave Island off the east
eastern Brazil (which do not support extant coast of Hispaniola); P. p. portoricensis
populations) indicate that the range of M. (PuertoRico);P.p.mexicanus(Mexicofrom
megalophylla was once more extensive than Sonora and Tamaulipas to Oaxaca and Ve-
it is today. racruz);P.p.mesoamericanus(westerncoast
Mormoops megalophylla has been cap- of Central America from Chiapas to western
tured at sites from sea level to about 3000 m Panama; eastern coast of Central America
in habitats ranging from lowland rainforest from Veracruz and Yucatan to Honduras);P.
to semiarid and arid scrub forest (Webb and p. rubiginosus (Honduras to Panama, Trini-
Baker, 1962; Smith, 1972; Bateman and dad and Tobago, southern Venezuela to Su-
Vaughan, 1974; Albuja, 1982; Graham and rinam,easternPeru,andnortheasternBrazil);
Barkley, 1984; Sanchez-Herrera et al., 1986; P. p. fuscus (northeastern Colombia and
Bonaccorso et al., 1992; Rezsutek and Cam- northern Venezuela except for the Paraguana
eron; 1993; Reid, 1997). This species varies peninsula);andP.p.paraguensis(Paraguana
from uncommon to locally common depend- peninusla of northern Venezuela). P. parnel-
ing on locality (Reid, 1997), typically ap- lii has recently been found on the island of
pearing to be most abundant in hot lowland St. Vincent in the Lesser Antilles, but the
areas in the northern parts of its range (Eas- subspecific status of this population has not
terla, 1973) and moist forested areas in the yet been assessed (Vaughan and Hill, 1996).
tropics (Handley, 1976). Favored roosting The fossil record of P. parnellii within its
sites include caves and abandoned mine currentrangeincludeslatePleistocenefossils
shafts,wherethesebatsoccurincoloniesthat from Puerto Ricoandsubfossilremainsfrom
may include as many as 500,000 individuals Mexico, Cuba, Jamaica, and eastern Brazil
(Barbour and Davis, 1969; Bateman and (Martin, 1972; Silva-Taboada, 1979; Herd,
Vaughan, 1974; Graham and Barkley, 1984; 1983; Koopman, 1989; Morgan, 1989; Cza-
Bonaccorso et al., 1992; Rezsutek and Cam- plewski and Cartelle, 1998). Extinct popu-
eron; 1993). The diet of M. megalophylla lations are known from a number of Carib-
consists of moths, beetles, and flies; most bean islands including Isla de Pinos, New
prey items are relatively large, with body Providence in the Bahamas, La Gonave, An-
lengths of 5–6 mm (Easterla and Whitaker, tigua, Grand Cayman, and Tobago (Koop-
1972; Ceballos and Galindo, 1984). man, 1955, 1989; Silva-Taboada, 1979;
Mormoops magna Silva-Taboada, 1974 Steadman et al., 1984; Eshelman and Mor-
is known only from a pair of fossil humeri gan, 1985; Morgan and Woods, 1986; Mor-
collected from cave in Cuba (Silva-Taboada, gan, 1989). These records indicate that the
1974, 1979). These elements resemble those Late Quaternary range of P. parnellii was
of M. megalophylla but are larger than other somewhat more extensive than it is today.
specimens from the same fossil layer that Pteronotus parnellii has been captured at
have been referred to M. megalophylla (Sil- sites from sea level to about 3000 m in hab-
va-Taboada, 1974, 1979). itatsrangingfromlowlandrainforesttosemi-
10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 258
arid and arid scrub forest (Smith, 1972; a´n and Kunz, 1992; Koopman, 1994): P. q.
Bowles et al., 1979; Silva-Taboada, 1979; quadridens (Cuba) and P. q. fuliginosus ((cid:2)
Herd, 1983; Eisenberg, 1989; Reid, 1997). It inflata; Jamaica, Hispaniola, and Puerto
is a relatively common bat throughout much Rico). The fossil record of P. quadridens in-
of its range and is abundant at some locali- cludes late Pleistocene and Holocene mate-
ties, particularly in moist forests (Smith, rial from several caves in Cuba (Silva-Ta-
1972; Handley, 1976; Silva-Taboada, 1979; boada, 1974, 1979; Woloszyn and Silva-Ta-
Herd, 1983; Reid, 1997). Preferred roosting boada, 1977) and the Bahamas (Andros and
sites include hot, humid caves and mines New Providence; Morgan, 1989). The latter
(Goodwin, 1970; Bowles et al., 1979; Silva- populations are now extinct.
Taboada, 1979; Vizotto et al., 1980; Herd, Pteronotus quadridens is one of the most
1983; Bonaccorso et al., 1992; Rodr´ıguez- abundant bats in Cuba and Puerto Rico,
Dura´n, 1998), but P. parnellii also occurs in where it roosts principally in hot, humid
rainforest regions that have no such refugia caves (Silva-Taboada, 1979; Rodr´ıguez-Dur-
(Simmons and Voss, 1998). In these areas, a´n and Lewis, 1987; Rodr´ıguez-Dura´n and
this species may occupy cavelike hollows in Kunz, 1992; Rodr´ıguez-Dura´n, 1995, 1998).
large trees. Colony size may exceed 140,000 individuals
ThedietofP.parnelliiconsistsprincipally (Rodr´ıguez-Dura´n and Lewis, 1987).
of moths, beetles, orthopterans, and flies, Pteronotus quadridens feeds on flying in-
with relative proportions of each varying sects that are captured primarily in the forest
among populations and seasons (Bateman understory (Rodrı´guez-Dura´n and Kunz,
and Vaughan, 1974; Howell and Burch, 1992). Dietary analyses based on stomach
1974; Silva-Taboada, 1979; Whitaker and contents and fecal samples indicate that a
Findley,1980;Herd,1983).Otherinsectsoc- wide variety of prey are taken by these bats,
casionally taken include odonatans and hy- including beetles, moths, flies, orthopterans,
menopterans (Silva-Taboada, 1979). Seeds homopterans, hymenopterans, dyctiopterans,
were found in the feces of four individuals dermapterans, hemipterans, and heteropter-
(Whitaker and Findley, 1980), but it is not ans (Silva-Taboada, 1979; Rodr´ıguez-Dura´n
known if fruit is intentionally ingested by and Lewis, 1987; Rodr´ıguez-Dura´n et al.,
this species. P. parnellii is unique among 1993). Remains from as many as seven dif-
mormoopids in using long CF (constant fre- ferent families of insects have been found in
quency)echolocationcallsandDopplercom- samples from a single individual P. quadri-
pensation (Novick, 1963, 1965, 1977; Nov- dens, suggesting that these bats may be op-
ick and Vaisnys, 1964; Schnitzler, 1970a, portunistic feeders (Silva-Taboada, 1979;
1970b, 1987; Fenton, 1980, 1982, 1984, Rodr´ıguez-Dura´n and Kunz, 1992). Small
1994, 1995; Schnitzler and Henson, 1980; amounts of pollen have also been found in
Simmons, 1980; Simmons and Stein, 1980; some stomach and fecal samples (Silva-Ta-
Neuweiler, 1984, 1989, 1990; Neuweilerand boada, 1979; Rodr´ıguez-Dura´n and Lewis,
Fenton, 1988; Schnitzler and Kalko, 1998; 1987), but this is thought to be a result of
Kalko and Schnitzler, 1998). All prey is ap- feeding of pollen-covered insects rather than
parentlycapturedonthewingandthereisno intentional feeding directly on flower prod-
evidence that feeding roosts are ever used ucts (Rodr´ıguez-Dura´n and Kunz, 1992).
(Bateman and Vaughan, 1974; Herd, 1983). Pteronotus macleayi Gray 1839 (Ma-
Pteronotus quadridens Gundlach, 1840 cleay’s mustached bat) isanothersmall-sized
(Sooty mustached bat) isthesmallestspecies species (forearm length (cid:2) 41–46 mm; con-
of Pteronotus (forearm length (cid:2) 35–40 mm; dylobasal length (cid:2) 14–16 mm; 4–8 g) that
condylobasal length (cid:2) 12–14 mm; 3–6 g; is presently confined to Cuba and Jamaica
Silva-Taboada, 1979; Koopman, 1994). The (Silva-Taboada, 1979; Koopman, 1993,
geographic range of this species is presently 1994). Two subspecies are currently recog-
restricted to several islands in the Greater nized (Smith, 1972; Koopman, 1994): P. m.
Antilles (Koopman, 1993, 1994). Two sub- macleayi(Cuba)andP.m.griseus(Jamaica).
species are currently recognized (Smith, Fossils of this species are known from the
1972; Silva-Taboada, 1979; Rodr´ıguez-Dur- late Quaternary of the Bahamas (New Prov-
