Upper molar morphology and relationships among higher taxa in bats

Author(s): Maria Paula Aguiar Fracasso , Leandro de Oliveira Salles , and Fernando Arajo Perini
Source: Journal of Mammalogy, 92(2):421-432. 2011.
Published By: American Society of Mammalogists
DOI: http://dx.doi.org/10.1644/09-MAMM-A-415.1
URL: http://www.bioone.org/doi/full/10.1644/09-MAMM-A-415.1

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Journal of Mammalogy, 92(2):421432, 2011

Upper molar morphology and relationships among higher taxa in bats

MARIA PAULA AGUIAR FRACASSO,* LEANDRO DE OLIVEIRA SALLES,

AND

FERNANDO ARAUJO PERINI

Setor de Mastozoologia, Departamento de Vertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro,
Quinta da Boa Vista, s/n, Rio de Janeiro, Rio de Janeiro 20940-040, Brazil (MPAF, LOS, FAP)
Departamento de Genetica, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Avenida Brigadeiro
Trompowski, s/n, Ilha do Fundao, Rio de Janeiro, Rio de Janeiro 21941-617, Brazil (FAP)
* Correspondent: mpa.fracasso@gmail.com
We reviewed topographical homologies in the upper tooth morphology of bats and analyzed the implications to
relationships among higher taxa within Chiroptera. A standardized terminology for the upper molars of bats is
proposed, taking into consideration the nomenclature adopted for tribosphenic mammals. Major patterns of variation
in crown morphology of chiropteran upper molars were reevaluated, and 2 new structures were identified:
mesoconule and mesoconule crista. The main controversies in the literature regarding terminology and structural
identity in the upper molars of chiropterans are discussed. Forty-eight dental morphological characters are presented
for extant bats and the extinct Icaronycteridae, with the exception of Pteropodidae, which has lost the tribosphenic
dental pattern. These were combined with 191 characters of other morphological systems from the literature. The tree
obtained from parsimony analyses mostly agrees with previous proposals based on morphology. However, major
differences were found: the position of Noctilionoidea at the base of the radiation of modern microchiropterans,
which formed a trichotomy with Yinochiroptera (including Emballonuridae) and the remaining Yangochiroptera;
Antrozoinae disassociated from the other Vespertilionidae, forming a poorly supported clade with Mystacinidae and
Molossidae; and the relationship between the sister taxa Myotinae and Miniopterinae within Vespertilionidae.
Key words:

Chiroptera, dentition, homology, nomenclature, phylogeny

E 2011 American Society of Mammalogists

DOI: 10.1644/09-MAMM-A-415.1

Despite past controversies (Baker et al. 1991; Pettigrew

1991; Simmons 1994), the order Chiroptera currently stands as
one of the most strongly supported monophyletic groups
within the eutherian mammals (Gunnell and Simmons 2005;
Simmons et al. 2008; Teeling et al. 2005). Similarly, the extant
families are well supported by both morphological (Gunnell
and Simmons 2005; Simmons 1998) and molecular data
(Teeling et al. 2000, 2002, 2005; Van Den Bussche and
Hoofer 2000, 2001; Van Den Bussche et al. 2002, 2003).
However, suprafamilial and subfamilial ranks remain contentious, especially because of differences among molecular and
morphological data (Miller-Butterworth et al. 2007; Simmons
2005a; Simmons et al. 2008; Teeling et al. 2005).
The study of tooth morphology is particularly relevant to
mammalogy because of its abundance in the paleontological
record, structural diversity, and morphofunctional implications. This is especially true when applied to Chiroptera,
which displays the largest intraordinal radiation of feeding
habits in mammals, including different kinds of herbivory,
insectivory, carnivory, piscivory, and sanguivory (Ferrarezi
and Gimenez 1996). The tribosphenic structure of bat molars,
however, has been poorly explored for patterns of cusp

variation and homology. Studies that evaluate aspects of tooth

morphology in the broader context of Chiroptera did not fully
detail the diversity of cusp forms present in the molar crowns
(Koopman and MacIntyre 1980; Miller 1907; Slaughter 1970).
Moreover, detailed studies usually are restricted to a single
genus, family, or subfamily (Czaplewski et al. 2003a;
Legendre 1984; Menu 1985; Morgan and Czaplewski 2003;
Phillips 1971). Other studies have presented a quantitative or
morphofunctional point of view and ignored or only
superficially mentioned questions related to the homology of
the cusps (Evans 2005; Freeman 1981b, 1998) and its
phylogenetic relevance. The lack of a standardized nomenclature for the chiropteran dentition only adds to the
difficulties regarding its use for taxonomic, phylogenetic,
and paleontological purposes. A need clearly exists for a more
detailed study of cusp homologies in bats, especially in light
of their potential contribution to the elucidation of unresolved
questions in bat phylogeny.

www.mammalogy.org
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JOURNAL OF MAMMALOGY

Simmons and coworkers (Gunnell and Simmons 2005;

Simmons and Geisler 1998; Simmons et al. 2008) conducted
the main morphological analyses of relationships among
higher taxa within Chiroptera. In those papers tooth morphology was not explored in detail. For example, of the 204
characters of the morphology-based matrix, only 8 refer
directly to tooth morphology. Our paper aims to develop
further the understanding of topographical homology in the
upper-tooth morphology of Chiroptera (excluding Pteropodidae) and thereby improve the morphological data set with
potential to infer relationships among higher taxa in bats.

MATERIALS AND METHODS

We analyzed specimens of at least 1 genus of each of the
extant families and subfamilies of Chiroptera, as well as
Icaronycteris, from the extinct family Icaronycteridae (based
on molar illustrations in Hooker [1996]). Exceptions are
Craseonycteridae, a rare monotypic family to which we did
not have access; Pteropodidae, because of extreme modifications that resulted in the loss of the tribosphenic pattern in this
family; and most of the subfamilies of Phyllostomidae (only
Phyllostominae was sampled), because of the extensive dental
modifications within the family, which are regarded as outside
the scope of this work. A total of 50 microbat genera were
examined as part of this study (Appendix I). The classification
of Simmons (2005b) was used as our standard taxonomic
reference, but ranks above family follow Simmons and Geisler
(1998) and Koopman (1985; for Yinochiroptera only).
Descriptions of variation in patterns in the tooth morphology of bats were based on hypotheses of topographic
correspondence relative to shape, position, and association
with other structures and dimensions. The standardization of
the terminology for upper molar cusps was based on a
comparative analysis of bats and other tribosphenic mammals
(Bown and Kraus 1979; Czaplewski et al. 2008; Legendre
1984; MacIntyre 1966; Menu 1985; Wible et al. 2009). The
nonbat tribosphenidans studied included specimens of didelphimorph marsupials (Caluromys, Didelphis, Gracilinanus,
Micoureus, and Thylamys), a eulipotyphlan (Sorex), and the
extinct cimolestans (Pantolambda and Cimolestes), condylarthrans (Choeroclaenus and Litaletes), and Procreodi (Tricentes
and Oxyclaenus; Appendix I). The numbering of teeth refers to
half of the dental arcade, as is customary for mammals. Upper
teeth are designated by I1I2 (incisors), C (canine), P2P3P4
(premolars), and M1M2M3 (molars).
Our ingroup of bats is composed of nearly the same extant
terminal taxa that were used by Gunnell and Simmons (2005).
Additionally, Sorex was chosen as an outgroup representing
Eulipotyphla, a group at the base of Laurasiatheria and closely
related to a clade that includes Chiroptera (Bininda-Emonds et
al. 2007; Onuma et al. 2000; Waddell et al. 1999). Sorex has a
tribosphenic pattern that allows unequivocal recognition of
cusp homologies with bats.
The data matrix combines our 48 characters hypothesizing
primary homology of the upper molar patterns observed within

Vol. 92, No. 2

Chiroptera and the 191 characters describing other morphological complexes proposed by Gunnell and Simmons (2005).
However, characters that represented redundancies (characters
14, 18, and 20), concerning lower teeth (characters 19, 22, and
23), or did not show any variation for the terminal taxa
considered in our study (characters 13, 24, 118, 152, 159, 175,
and 200) were excluded. In total, 44 characters were regarded as
additive, and 195 characters were considered nonadditive.
Phylogenetic analyses with unweighted characters were
conducted using the parsimony algorithm (traditional search)
implemented by the TNT program (Goloboff et al. 2008)
consisting of a heuristic search with 1,000 replicates of randomaddition sequence followed by tree-bisection-reconnection
branch swapping. The new technology search options of TNT
(sectorial searches, ratchets, tree drifting, and tree fusing) also
were used with default settings to confirm the most-parsimonious trees. A decay (Bremer 1994) and bootstrap analysis (using
heuristic methods of random-addition sequence, 10 repetitions
for each of 3,000 bootstrap replicatesFelsenstein 1985) were
performed to evaluate the relative support for various groupings.
Character-state distributions were examined using TNT.

RESULTS
Standardized terminology for upper molars of bats is proposed
(Table 1) and illustrated (Fig. 1) based on comparison of
tribosphenic mammal dentition (Bown and Kraus 1979; Goin
and Candela 2004; Hershkovitz 1977; MacIntyre 1966; Salles
1996; Szalay 1969; Vandebroek 1967; Van Valen 1966; Wible et
al. 2009) and bat dentition (Czaplewski et al. 2008; Freeman
1998; Legendre 1984; Menu 1985; Slaughter 1970). We present
48 characters based on the upper-tooth morphology of bats
(Appendix II). Most of the characters (30) are original hypotheses
of homology for bats, and characters 7 and 8 refer to structures
that are described here for the 1st time for Chiroptera. Combined
with other morphological characters from Gunnell and Simmons
(2005), the complete matrix has 239 characters (Appendix III).
Analysis of the total matrix resulted in 2 equally mostparsimonious trees with 764 steps, consistency index (CI) of
0.401 and retention index (RI) of 0.532. The strict consensus
was calculated (CI 5 0.398 and RI 5 0.527), producing a tree
with 22 clades (Fig. 2).
The extinct family Icaronycteridae is positioned at the base
of the tree, followed by a trichotomy of 3 clades:
Yinochiroptera (sensu Koopman 1985; includes Emballonuridae), Noctilionoidea, and a large group composed of the
remaining Yangochiroptera families and subfamilies. Chiroptera (excluding Icaronycteridae) is relatively well supported,
with decay values of 7 and bootstrap values of 89%.
Within Yinochiroptera, Emballonuridae is rooted at the
basal node, followed by Rhinopomatidae, Nycteridae, Megadermatidae, Rhinolophidae, and Hipposideridae in a successively branching sequence of families. Although relatively
weak support exists for the basal position of Emballonuridae
and Rhinopomatidae, relationships among more differentiated
yinochiropterans have higher decay and bootstrap values.

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FRACASSO ET AL.UPPER DENTITION AND PHYLOGENY OF BATS

423

TABLE 1.Comparison of commonly used terms for the upper

molars of bats and other tribosphenic mammals with proposed
standardization of molar terminology for Chiroptera.
Tribosphenic mammals

Chiroptera

Standardization

Labial cusps
Paracone
Metacone
Stylocone
Parastyle
Mesostyle
Metastyle
Stylar cusps
Lingual cusps
Protocone
Hypocone
Pericone
Paraconule
Metaconule
Mesoconule

Paracone
Metacone
Absent structure
Parastyle
Mesostyle
Metastyle
Absent structure

Paracone
Metacone
Absent structure
Parastyle
Mesostyle
Metastyle
Absent structure

Protocone
Hypocone
Absent structure
Paraconule
Metaconule
Not mentioned before

Protocone
Hypocone
Absent structure
Paraconule
Metaconule
Mesoconule

Paracrista
Postparacrista

Preparacrista
Postparacrista

Premetacrista

Premetacrista

Metacrista

Postmetacrista

Preprotocrista
Postprotocrista
Not differentiated
Paraloph
Metaloph
Not differentiated
Not mentioned before

Preprotocrista
Postprotocrista
Not differentiated
Postparaconule crista
Premetaconule crista
Not differentiated
Mesoconule crista

Paracingulum
Metacingulum
Precingulum
Postcingulum
Labial cingulum
Lingual cingulum

Paracingulum
Metacingulum
Precingulum
Postcingulum
Labial cingulum
Lingual cingulum

Protoconal basin/trigon
basin
Protofossa
Absent structure
Talon/hypoconal basin
Not named before
External valley
Labial flexus

Trigon basin

Labial crests
Preparacrista
Postparacrista/
centrocrista
Premetacrista/
centrocrista
Postmetacrista
Lingual crests
Preprotocrista
Postprotocrista
Preparaconule crista
Postparaconule crista
Premetaconule crista
Postmetaconule crista
Not named before
Cingula
Paracingulum
Metacingulum
Precingulum
Postcingulum
Ectocingulum
Lingual cingulum
Other structures
Trigon basin
Protofossa
Pretalon
Talon/talon basin
Not named before
Stylar shelf
Ectoflexus

Protofossa
Absent structure
Talon
Paracingulum expansion
Stylar shelf
Ectoflexus

Yangochiroptera (excluding Noctilionoidea) has a dichotomy

at its base between Nataloidea and a group including
Mystacinidae, Molossidae, and Vespertilionidae. Myzopodidae
is positioned at the base of Nataloidea, followed by Natalidae,
which is sister to the clade composed of Thyropteridae and
Furipteridae. The clade Nataloidea is well supported, but decay
and bootstrap values supporting its internal relationships are
relatively low. The remaining Yangochiroptera are composed
of 2 main branches. The 1st is a clade including Antrozoinae,

FIG. 1.Upper molar terminology in bats: a) Myotis albescens

(MN 3380); b) Chaerephon leucogaster (AMNH 115965); and c)
Rhynchonycteris naso (MN 3584). Figures are not to scale.

followed by Mystacinidae, which is sister to Molossidae (with

Molossinae and Tomopeatinae forming a well-supported
monophyletic group). The other main branch consists of
Vespertilionidae (excluding Antrozoinae) standing as a poorly
supported monophyletic group, with Vespertilioninae being the
sister taxon to 2 sister clades, embracing Myotinae and
Miniopterinae and Kerivoulinae and Murininae.

DISCUSSION
The homology of upper molar cusps and crests among bats
and other mammals is not entirely clear. Different nomenclatures, sometimes reflecting different assumptions regarding

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JOURNAL OF MAMMALOGY

FIG. 2.A strict consensus of 2 most-parsimonious trees (764

steps each) derived from analysis of our tooth data set together with
the morphological data set of Gunnell and Simmons (2005). Numbers
above the branches are decay values; those below the branches are
bootstrap values.

cusp and crest relationships, are widespread in the literature

(Czaplewski et al. 2008; Freeman 1998; Legendre 1984; Menu
1985; Slaughter 1970). The main controversies regarding
terminology and structural identity in the upper molars of
chiropterans are discussed below.
Hypocone and metaconule.The main posterolingual cusp
in the upper molars is usually identified as a hypocone.
However, this cusp could be the result of an independent
differentiation process (Hunter and Jernvall 1995; Jernvall
1995). The 2 most common interpretations are: the posterolingual cusp is the result of expansion of the postcingulum,
resulting in a hypocone; or the posterolingual cusp emerges as
part of the development and displacement of the metaconule.
In both cases the final tooth morphology might be indistinguishable, and the identity of the hypocone cannot be inferred
from the configuration and position of the posterolingual cusp.
In Chiroptera the hypocone can be absent or vary from tiny
(Chrotopterus and Phyllostomus) to large (Noctilio and Pteronotus). In contrast, the metaconule is usually associated with the
trigon basin and can vary from a minuscule elevation inserted on
the postprotocrista to a large cone similar to the hypocone, or be

Vol. 92, No. 2

absent. The position of the metaconule on the labiallingual axis

also varies from close to the base of the metacone to close to the
tip of the protocone with some intermediate stages. Another
variation in metaconule position occurs on the anteroposterior
axis with insertion on or close to the postprotocrista (Molossus) to
being completely separated (Tadarida and Mormopterus). An
intermediate position is found in Myotis, Thyroptera, Molossops
temminckii, and Chaerephon leucogaster, among others. In M.
temminckii the metaconule is medium sized and displaced from
its original position, although its top is still inserted on the
postprotocrista. In C. leucogaster the metaconule is medium
sized and slightly displaced from its original position in M2
(Fig. 1b) but is large and more displaced in M1, with the top of
the cusp just separate from the postprotocrista. When the
metaconule is well developed (Tadarida and Mormopterus)
and is situated away from the postprotocrista, its general structure
is shaped and positioned similar to a large hypocone. In these
cases we assert that this cusp is a metaconule instead of a
hypocone when it appears in closely related taxa (such as M.
temmicnkii and Cynomops) as a medium cusp closer to the
postprotocrista, and that the hypocone is absent.
Many authors (Czaplewski et al. 2003b; Gregorin 2000;
Legendre 1984) considered the large posterolingual cusp of
some molossid taxa as a hypocone because of its position and
size. Czaplewski et al. (2008) called it a hypocone in
Chiroptera if it was large and separate from the postprotocrista
on the talon. But molossids show all stages of development of
the metaconule: small (Molossus), medium (M. temminckii
and Cynomops), and large (Tadarida and Mormopterus). In
molossid taxa with a more conventional metaconule (small or
medium, in its original position or just a little displaced) the
hypocone is always absent. The small metaconule and the
large hypocone-like metaconule are very different, but the
intermediate stages (M. temminckii) show more continuous
variation between the morphological extremes. Therefore, we
assert that the large posterolingual cusp in some molossids is a
metaconule and not a hypocone.
Many authors have recognized the posterolingual cusp in
Vespertilionidae as a metaconule instead of a hypocone
(Czaplewski 1993; Menu 1985), but it is not as well developed
as in some molossids. The hypocone is absent in all
vespertilionid taxa analyzed in our study.
The hypothesis that the large posterolingual cusp in the
upper molars is in some cases a well-developed metaconule
and not a hypocone, as proposed here for some chiropterans
(molossids, vespertilionids, and thyropterids), has been
proposed before for other unrelated mammal groups (Case et
al. 2005; Goin et al. 2003, 2007; Hunter and Jernvall 1995;
Jernvall 1995) and for some vespertilionid bats (Czaplewski
1993; Menu 1985). Hunter and Jernvall (1995) considered the
posterolingual cusp of bat molars to be a hypocone, but they
used a limited sample of bat taxaonly Phyllostomidae,
Pteropodidae, and Mystacinidae, all of which have a hypocone
associated with the postcingulum.
Mesoconule and mesoconule crista.Some bat species
have a small accessory cusp (or cuspule) on the internal base

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FRACASSO ET AL.UPPER DENTITION AND PHYLOGENY OF BATS

of the metacone, usually situated in the same region as the

internal portion of the premetaconule crista (when the crista is
absent) or associated with the premetaconule crista (Rhynchonycteris). No reference to or citation of this cuspule is found in
the literature on Chiroptera. Because of its position close to
the base of the metacone, this cusp sometimes can be
considered erroneously a vestigial metaconule. However, the
position of the metaconule in bats is associated with the
margin of the trigon, and the small cuspule was observed in
some species in which the metaconule is either vestigial or
well developed.
This small cuspule was observed in a few bats (Rhynchonycteris, Cormura, Chaerephon, Myotis, Furipterus, and
Thyroptera), and was absent in marsupials and other placental
mammals analyzed. The only mention of a similar structure
was MacIntyre (1966:158) on carnivores of the extinct family
Miacidae, who stated that the metaconule marks the point
where the protoconemetastyle crest (5 postprotocrista) gives
off a short branch (5 premetaconule crista) which connects
the metaconule with a small cuspule (mesoconule) near the
lingual base of the metacone. This description is similar in
shape, position, and associated structures to that found in some
bats and has not been described for Chiroptera. Thus we adopt
mesoconule as the preliminary assignment for this tiny
accessory cusp, although we recognize that further research
is needed to confirm its relationships.
The premetaconule crista continues after the mesoconule in
a curved path toward the base of the mesostyle on the floor of
the trigon basin. At this point it can be interrupted by a small
gap before it reaches the labial corner of the trigon basin
nearest to the lingual side of the mesostyle, but in some
specimens the crest completely traverses the trigon basin. We
name this crest the mesoconule crista. It can occur either
associated with or independent of the mesoconule.
Both the mesoconule and the mesoconule crista are small
structures compared to the other molar cusps and potentially
highly susceptible to tooth wear. Additionally, these minuscule
structures show great intraspecific variation in all analyzed taxa,
except Rhynchonycteris, where they seem to be always present
(Fig. 1c). In Myotis the mesoconule crista can be rudimentary or
absent in different specimens of M. albescens, M. levis, M.
ruber, and M. riparius; however, this structure was not found in
M. velifer and M. nigricans. In none of the Myotis species
observed is the mesoconule crista always present (Fig. 1a).
The bats that have a mesoconule are Rhynchonycteris,
Cormura, Chaerephon, Myotis, Furipterus, and Thyroptera.
The mesoconule is a tiny structure and varies intraspecifically
for presence or absence, except in Rhynchonycteris, which has
a small but distinct mesoconule that is always present. The
mesoconule crista always was found in Rhynchonycteris and
can be present or absent (intraspecifically) in Mormopterus,
Chaerephon, Tadarida, and Myotis.
Some extinct miacids have 2 related structures (mesoconule
and mesoconule crista) similar to those found in some bats,
which suggests an evolutionary convergence. Furthermore,
this structure could have evolved convergently at least twice in

425

bats. However, these tiny structures, whose occurrence is

inter- and intraspecifically variable, could be easily overlooked in descriptions of tooth morphology of other
mammalian taxa, as it was for Chiroptera. If it is confirmed
that these structures occur in other chiropteran taxa, they could
be pleisiomorphic for bats or even for a higher taxon. More
thorough analyses of tooth morphology, especially focusing on
Eocene fossil bats and basal placentals, are needed to clarify
this and other questions of structural identities and homology.
Paraconule crista and metaconule crista.Chiropterans
have 1 or 2 small crests beginning in the central region of the
trigon basin and extending directly to its margin, where they
usually connect to the paraconule and metaconule (when these
cusps are present), or to the pre- and postprotocrista (Figs. 1a
and 1b). Some bat studies use the terminology paraloph and
metaloph (Legendre 1984; Menu 1985), or conule wings
(Slaughter 1970), for those crests. However, the crests
associated with the metaconule and paraconule cusps in other
tribosphenic mammals are usually called (pre- and post-)
paraconule crista and (pre- and post-) metaconule crista
(Bown and Kraus 1979; Szalay 1969). The paraconule and
metaconule cusps in bats are very tiny structures that can be
absent in many bat taxa. Also, they wear away fairly quickly
so are difficult to distinguish in some species.
The postparaconule crista and the premetaconule crista of
tribosphenic mammals correspond to the paraloph and
metaloph of bats. The preparaconule crista and the postmetaconule crista of tribosphenic mammals are not evident in
bats, probably having fused with the paracingulum and
metacingulum, or with the preprotocrista and postprotocrista,
respectively, or simply are absent.
To standardize molar terminology we have adopted the
names postparaconule crista and premetaconule crista for bats.
In Chiroptera these crests can be well developed, as in some
Molossidae and Vespertilionidae, but in most taxa they vary
from rudimentary to absent (Myzopoda, Macrophyllum,
Mormoops, and Noctilio). Tooth wear can make these crests
difficult to detect. A few taxa also show a small modification
in the position of these crests, which are directed toward the
lingual end of the trigon basin instead of the trigon margin
(Fig. 1c). Czaplewski et al. (2003a) considered this modification to be an autapomorphic condition in Nyctinomops
(Molossidae), but it occurs convergently in some emballonurids (Rhynchonycteris, Balantiopteryx, and Emballonura).
Postcingulum and talon.The postcingulum follows the
posterior base of the protocone. In Chiroptera this cingulum
varies from rudimentary to large. The large cingulum fills the
space between the molars.
The occlusion surface formed by the expansion of the
postcingulum usually is called the talon (Czaplewski et al.
2008; Legendre 1984), but some authors (Freeman 1998) also
have called it the hypoconal basin. The term talon appears in
both bat and other tribosphenic mammal terminology. It
represents the occlusal surface posterior to the trigon basin.
The term hypoconal basin is not appropriate because the talon
or postcingulum sometimes occurs independently of the

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JOURNAL OF MAMMALOGY

presence of a hypocone. For example, Megadermatidae and

Nycteridae have large talons without hypocones.
The postcingulum is always present in bats, except for
intraspecific variation in Molossus rufus. The talon (expansion
of the postcingulum forming an occlusal surface) can vary
from absent to very large.
Paracingulum expansion.The M1 of some emballonurids
(Rhynchonycteris, Emballonura, and Balantiopteryx) has an
anterolabial expansion of the paracingulum that is not
observed in other mammals studied. This expansion was
described by Plumpton and Jones (1992:1) as a conspicuous
anteroexternal cusp that is distinct from the small parastyle
for Rhynchonycteris. We consider this structure as a small
shelf and not a real cusp (Fig. 1c).
Phylogenetic analyses.Our tree shares the following
phylogenetic patterns with that of Gunnell and Simmons
(2005): Icaronycteridae occupies a basal position; modern
microchiropterans form a monophyletic group; Yinochiroptera
are monophyletic and show the same internal relationships in
both analyses, including Emballonuridae at the base; Molossidae subfamilies are monophyletic with high support;
relationships within Noctilionoidea are the same, but Gunnell
and Simmons (2005) have Mystacinidae as its sister taxon;
Nataloidea is composed of the same families, with Myzopodidae basal to the other taxa, but internal relationships are
different; and Vespertilionidae, to the exclusion of Antrozoinae, is monophyletic and congruent between the 2 analyses.
Some relevant differences between the phylogenetic results
of Gunnell and Simmons (2005) and ours are worth mention.
The inclusion of Noctilionoidea with the other Yangochiroptera is not supported by our data set, which places
Noctilionoidea in a basal radiation of modern microchiropterans forming a trichotomy with Yinochiroptera and the
remaining Yangochiroptera. This arrangement is not supported by other recent phylogenetic analyses (Gunnell and
Simmons 2005; Simmons and Geisler 1998; Simmons et al.
2008) but was proposed earlier by Smith (1976).
In our analysis a sister-taxon relationship between Furipteridae and Thyropteridae is supported by tooth characters 28
(presence of an anterolabial basin of P4), 35 (absence of the
incisor-directed crest on the main cone of C), and 46 (I2 with 2
cusps; convergent with Myotinae). Furipteridae and Thyropteridae also were considered sister groups by Eisenberg (1981) and
Smith (1976). In contrast, Gunnell and Simmons (2005) found a
sister-taxon relationship between Furipteridae and Natalidae,
with Thyropteridae the sister family to this clade. In our
topology Natalidae is basal to Thyropteridae and Furipteridae.
Examination of our data also shows a sister-taxon relationship between Mystacinidae and Molossidae, supported by tooth
characters 11 (reversal of the position of the lingual extremity of
the metacingulum to directly join the postprotocrista), 12
(absence of the lingual cingulum of M1M2), 13 (absence of
precingulum of M1M2), and 16 (the corner between the
postparacrista and premetacrista forms a narrow curve). This
relationship was proposed before by Eisenberg (1981), Novacek
(1991), Simmons and Geisler (1998), and Smith (1976).

Vol. 92, No. 2

The inclusion of Antrozoinae at the base of the Mystacinidae and Molossidae clade, and disassociated from other
Vespertilionidae, has weak support from only character 41
(absence of I2). Traditional classifications (Hill and Smith
1984; McKenna and Bell 1997; Simmons 2005b) included
Antrozoinae among vespertilionids. Simmons (1998) and
Simmons and Geisler (1998), however, raised this group to
family level and moved it to Molossoidea. Gunnell and
Simmons (2005) placed Antrozoidae as the sister family to
Vespertilionidae, but this relationship had low decay and
bootstrap support. We propose that this taxon should be
treated as a family separate from the other Vespertilionidae
because of its uncertain position among the Yangochiroptera.
The sister-taxon relationship between Myotinae and Miniopterinae contradicts previous proposals to recognize Miniopteridae as a separate family from Vespertilionidae (Gopalakrishna and Chari 1983; Mein and Tupinier 1977; MillerButterworth et al. 2007; Simmons et al. 2008). The clade of
Myotinae and Miniopterinae appears within Vespertilionidae
and is supported by character 1 (presence of a medium to large
metaconule in M1M2), character 2 (intermediate position of
the metaconule in relation to the posterior face of the
protocone in M1M2), character 20 (a reversal to a lingual
position of the metacone in M3 relative to its position in M1
M2), and character 43 (a reversal of I1 to a position anterior to
C). Therefore, we suggest that Miniopterinae be maintained as
a subfamily of Vespertilionidae.
We expect that the molar terminology proposed here can be
useful for future research on comparative tooth morphology
among bats and other tribosphenic mammals. The suprafamilial
and subfamilial relationships of bats are not clearly understood,
and dental attributes might prove to be decisive for more indepth phylogenetic reconstruction. Future developments of this
research will involve amplifying our data set with the inclusion
of Pteropodidae and all Phyllostomidae subfamilies; evaluating
cusp homologies on the lower teeth of Chiroptera; and furthering the study of patterns of tooth variation within tribosphenic
mammals.

RESUMO
Nos revisamos as homologias topograficas relativas a`
morfologia da denticao superior de morcegos e analisamos suas
implicacoes nas relacoes filogeneticas entre grandes grupos de
Chiroptera. E proposta uma padronizacao da terminologia dos
molares superiores de morcegos, levando em consideracao a
nomenclatura padrao adotada para os mamferos tribosfenicos.
Os principais padroes de variacao da morfologia da coroa dos
molares superiores de quiropteros foram reavaliados, e 2 novas
estruturas foram identificadas: mesoconule e crista mesoconular.
Sao tambem discutidas as principais controversias na literatura
com relacao a` terminologia e identidade estrutural dos molares
superiores dos quiropteros. Sao apresentados quarenta e oito
caracteres morfologicos dentarios presentes nas famlias de
morcegos viventes, com excecao de Pteropodidae, que apresenta
uma grande modificacao do padrao dentario tribosfenico original,

April 2011

FRACASSO ET AL.UPPER DENTITION AND PHYLOGENY OF BATS

e na famlia extinta Icaronycteridae. Estes caracteres foram

combinados com 191 caracteres de outros complexos morfologicos obtidos a partir da literatura. A arvore obtida a partir de
analises de parcimonia concorda, em grande parte, com propostas
anteriores baseadas em morfologia. No entanto, diferencas
consideraveis foram encontradas: a posicao do Noctilionoidea
na base da radiacao dos microquiropteros modernos, formando
uma tricotomia com Yinochiroptera (incluindo Emballonuridae)
e os demais Yangochiroptera; Antrozoinae dissociada dos demais
Vespertilionidae, formando um clado pouco suportado com
Mystacinidae e Molossidae; e a relacao entre os grupos-irmaos
Myotinae e Miniopterinae dentro de Vespertilionidae.

ACKNOWLEDGMENTS
We thank C. E. L. Esberard, J. A. Oliveira, L. M. Pessoa, M. R.
Nogueira, and R. Gregorin for previous insights on an earlier version
of this work; C. Tribe for gently reviewing this manuscript; M.
Weksler for kindly taking some photos; and J. da Silva, the artist
responsible for the tooth drawings. We also acknowledge M. de Vivo
(Museu de Zoologia, University of Sao Paulo) and N. B. Simmons
(American Museum of Natural History) for access to specimens in
their care. We extend our acknowledgments to the 2 reviewers for
their valuable comments on the manuscript. Support for this study
was provided by the Brazilian Research Council (CAPES) and
Brazilian National Council for Research Science (CNPq).

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Submitted 18 December 2009. Accepted 6 October 2010.

Associate Editor was Burton K. Lim.

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FRACASSO ET AL.UPPER DENTITION AND PHYLOGENY OF BATS

APPENDIX I
Specimens examined
Classification follows Gardner (2005) for Didelphimorphia,
McKenna and Bell (1997) for extinct placentals (Cimolesta,
Condylarthra, and Procreodi), Waddell et al. (1999) for Eulipotyphla,
and Simmons (2005b) for Chiroptera. Museums in which specimens
are preserved are identified by the following abbreviations: ALP,
Adriano Lucio Peracchi Collection, Universidade Federal Rural do
Rio de Janeiro (Seropedica); AMNH, American Museum of Natural
History (New York); FMNH, Field Museum of Natural History
(Chicago); JAO, field number of Joao Alves de Oliveira, material to
be deposited in the collection of Museu Nacional, Universidade
Federal do Rio de Janeiro (Rio de Janeiro); MN, Museu Nacional,
Universidade Federal do Rio de Janeiro (Rio de Janeiro); MZLSU,
Museum of Zoology, Louisiana State University (Baton Rouge);
MZUSP, Museu de Zoologia, Universidade de Sao Paulo (Sao
Paulo); and MVZ, Museum of Vertebrate Zoology, University of
California (Berkeley); UA, University of Alberta; and USNM, United
States National Museum (Washington, D.C.).
Didelphimorphia.Caluromys lanatus: MN 20963, 66334; Caluromys philander: MN 20785, 20790; Didelphis albiventris: MN
23654, 23666, 23676; Gracilinanus agilis: MN 4787, 4790, 13507;
Gracilinanus microtarsus: MN 8275, 11718; Micoureus demerarae:
MN 24670, 29417; Thylamys macrurus: FMNH 26760; Thylamys
pallidior: FMNH 51003; Thylamys venustus: FMNH 35014, 162507.
Cimolesta.Cimolestes magnus: UA 3793; Pantolambda cavirictum: USNM 21327.
Procreodi.Oxyclaenus sp.: AMNH 113970; Tricentes crassicolidens: AMNH 3101; Tricentes subtrigonus: AMNH 3240.
Condylarthra.Choeroclaenus turgidunculus: AMNH 16489;
Litaletes disjunctus: AMNH 35885.
Eulipotyphla.Sorex vagrans: MN 8149; Sorex palustris: MN 8175.
Chiroptera.Rhinolophidae: Rhinolophus acuminatus: AMNH
107854, 107865; Rhinolophus ferrumequinum: AMNH 160471,
160481; Hipposideridae: Aselliscus stoliczkanus: AMNH 115576,
119465; Aselliscus tricuspidatus: AMNH 110009, 110014; Hipposideros abae: AMNH 49122, 49130; Hipposideros bicolor: AMNH
103323, 216959; Megadermatidae: Macroderma gigas: AMNH
162669, 162672; Megaderma lyra: AMNH 208822, 208823; Megaderma spasma: AMNH 54782, 54815; Rhinopomatidae: Rhinopoma
hardwickii: AMNH 14476, 208126; Rhinopoma microphyllum:
AMNH 170277; Emballonuridae: Balantiopteryx plicata: AMNH
189567, 189572; Centronycteris maxiliani: AMNH 267397; Cormura
brevirostris: AMNH 78804; MZUSP 18860; Diclidurus albus: AMNH
99478, 149167; MN 11189; Diclidurus scutatus: AMNH 99309,
142908; Emballonura monticola: AMNH 216797, 247237; Emballonura beccarii: AMNH 191316, 191317; Mosia nigrescens: AMNH
105065, 105066; Rhynchonycteris naso: AMNH 92642, 92650, 94365,
94380, 209192, 209209, 230051; MN 3584, 3782; Taphozous
mauritianus: AMNH 48807, 48801; Taphozous nudiventris: AMNH
27391; Taphozous perforatus: AMNH 184457, 184458; Nycteridae:
Nycteris arge: AMNH 49133, 49138; Nycteris hispida: AMNH
165819, 165823; Myzopodidae: Myzopoda aurita: AMNH 257130;
Mystacinidae: Mystacina robusta (extinct): AMNH 160269, 214243;
Mystacina tuberculata: MVZ 174825; Phyllostomidae: Chrotopterus
auritus: AMNH 36989, 267852; MZUSP 1000, 28717; Glyphonycteris
daviesi: MZUSP 22532; Glyphonycteris sylvestris: AMNH 183846,
207061; MZUSP 22636; Macrophyllum macrophyllum: AMNH
177664, 177668; MN 37203; MZUSP 22735; Phyllostomus discolor:
AMNH 92189, 92194; MN 37338; Phyllostomus elongatus: MZUSP
22536, 22702; Phyllostomus hastatus: AMNH 134915, 134916; MN

429

3718, 13485, 36236; Trachops cirrhosus: MN 36037, 36720;

Mormoopidae: Mormoops blainvillii: AMNH 176140, 176149;
Mormoops megalophylla: AMNH 27300, 27305; MZUSP 27154;
Pteronotus gymnonotus: AMNH 32083, 32085; MZUSP 16647,
16650; Pteronotus parnellii: AMNH 31569, 31574; MZUSP 22625,
22646, 22716; Pteronotus personatus: MZUSP 16642, 16643;
Noctilionidae: Noctilio albiventris: AMNH 269204, 269206; MN
3370; Noctilio leporinus: AMNH 72315, 72318; MN 3506, 43076;
Furipteridae: Amorphochilus schnablii: AMNH 63723; Furipterus
horrens: AMNH 91903, 93900; MZUSP 20887, 20888, 22726, 22730;
Thyropteridae: Thyroptera discifera: AMNH 46687, 94549; Thyroptera tricolor: AMNH 269501, 269503; MN 46443; Natalidae:
Chilonatalus micropus: AMNH 61191, 216127; Chilonatalus tumidifrons: AMNH 40026; Natalus stramineus: AMNH 148386, 148387;
MZUSP 16640; Natalus tumidirostris: AMNH 176592, 176594;
Nyctiellus lepidus: AMNH 61169, 167130; Molossidae: Chaerephon
jobensis: AMNH 107803, 197157; Chaerephon leucogaster: AMNH
115964, 115965; Cynomops abrasus: AMNH 94625, 239235;
Cynomops greenhalli: AMNH 176286, 207071; Cynomops paranus:
AMNH 94630, 94633; Molossops temminckii: AMNH 181537,
181540; MN 3455; Molossops mattogrossensis: MN 3756; MZUSP
16886; Molossus molossus: AMNH 77679, 91464; MN 3211, 3661,
43856; Molossus rufus: MN 3496, 3603; MZUSP 4437, 21084;
Mormopterus kalinowskii: AMNH 165625, 165627; Mormopterus
planiceps: AMNH 194195, 194196; Tadarida brasiliensis: AMNH
39207, 39241; MN 3517, 3518; MZUSP 1744; Tadarida teniotis:
AMNH 235609, 245384; Tomopeas ravus: MZLSU 25072, 25084,
25148; Vespertilionidae: Antrozous pallidus: AMNH 3754, 121494;
Bauerus dubiaquercus: AMNH 180840, 254685; Eptesicus brasiliensis: MZUSP 27736; Eptesicus diminutus: MZUSP 27767; Eptesicus
furinalis: MZUSP 1746; Harpiocephalus harpia: AMNH 208656;
Histiotus montanus: AMNH 188780, 188781; Histiotus velatus:
AMNH 217565; MN 3373; MZUSP 9047; Kerivoula myrella: AMNH
194462, 194471; Kerivoula picta: AMNH 106704, 106709; Lasionycteris noctivagans: AMNH 99614, 175428; Miniopterus australis:
AMNH 194880, 194882; Miniopterus schreibersii: AMNH 107340,
107345; Murina florium: AMNH 109110, 109111; Murina suilla:
AMNH 103914, 217012; Myotis albescens: AMNH 93927, 93929; MN
3380; Myotis levis: JAO 1771, 1818, 1819; Myotis nigricans: AMNH
61769, 61773; Myotis riparius: ALP 3421; Myotis ruber: JAO 1751;
MN 3414; Myotis velifer: AMNH 203876, 203877; MN 3481;
Phoniscus jagorii: AMNH 106251, 107964; Rhogeessa io: AMNH
175731, 176378; MZUSP 24045; Rhogeessa parvula: AMNH 180522,
180543.

APPENDIX II
Character descriptions
Descriptions of the 48 characters related to the upper-tooth
morphology of bats (24 for molars, 9 for premolars, 7 for canines,
and 8 for incisors). The tooth characters regarded as additive are 1, 2,
4, 5, 20, 22, 24, 25, 30, 31, and 44. Other morphological characters
are described in Gunnell and Simmons (2005).
Character 1.Metaconule of M1M2: (0) absent; (1) minuscule to
small; (2) medium to large; (3) very large. A similar version of this
character was cited by Freeman (1981a: character C68) for
Molossidae, but she considered the large posterolingual cusp of
some molossids to be a hypocone, as opposed to a metaconule.
Character 2.Position of the metaconule in relation to the
posterior face of the protocone in M1M2: (0) close; (1) intermediate;
(2) apart.

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Character 3.Paraconule of M1M2: (0) present; (1) absent.

Character 4.Hypocone of M1M2: (0) absent; (1) minuscule to
small; (2) medium to large. Wetterer et al. (2000: character 61) and
Czaplewski et al. (2003a: character 32) described the development of
the hypocone and talon together in a single character.
Character 5.Talon of M1M2: (0) large; (1) minuscule to small;
(2) absent. Configuration of the talon was discussed together with the
development of the hypocone by Wetterer et al. (2000: character 61)
and Czaplewski et al. (2003a: character 32).
Character 6.Postparaconule and premetaconule cristae of M1
M2: (0) present; (1) absent.
Character 7.Mesoconule of M1M2: (0) present; (1) absent.
Character 8.Mesoconule crista of M1M2: (0) present; (1)
absent.
Character 9.Lingual extremity of the paracingulum of M1M2:
(0) directed toward the preprotocrista; (1) directed toward the
precingulum.
Character 10.Metacingulum of M1M2: (0) present; (1) absent.
Character 11.Lingual extremity of the metacingulum: (0)
directed toward the postprotocrista; (1) directed toward the
postcingulum or talon. Morgan and Czaplewski (2003:751, character
19) discussed the position of the M1 postprotocrista, describing that it
may become confluent with distal cingulum (metacingulum), a
condition similar to our state (0).
Character 12.Lingual cingulum of M1M2: (0) complete,
sometimes interrupted by a small gap; (1) restricted between the
lingual extremity of the protocone base and the hypocone; (2) absent.
We use the term lingual cingulum for the cingulum that is restricted
to the base of the protocone only, but it is usually continuous with the
precingulum (anterolingual) and postcingulum (posterolingual).
Czaplewski et al. (2003a) proposed a similar character (27: M1 and
M2 lingual cingulum), but their states diverge from ours because they
use the term lingual cingulum to encompass all 3 characters of the
lingual cingulum, precingulum and postcingulum.
Character 13.Precingulum of M1M2: (0) present; (1) absent.
Czaplewski et al. (2003a) did not separate the precingulum from the
lingual cingulum, but discussed its presence or absence in their
character 27 (lingual cingulum anterior to the protocone).
Character 14.Lingual projection of the paracone and metacone:
(0) absent; (1) present.
Character 15.Internal faces of the paracone and metacone: (0)
connecting labially, forming a clear dilambdodont pattern; (1)
connecting more lingually, forming a predilambdodont pattern. This
character may be a version of character 45 of Czaplewski et al.
(2003a) on mesostyle position. When the mesostyle falls on a line
drawn between the parastyle and metastyle (state 0 of character 45
of Czaplewski et al. 2003a:280), the internal faces of the metacone
and paracone connect labially, forming a clear dilambdodont pattern;
and when the mesostyle position is indented lingually (state 1 of
character 45 from Czaplewski et al. 2003a:280), the internal faces of
the paracone and metacone connect more lingually, forming a
predilambdodont pattern.
Character 16.Shape between the postparacrista and premetacrista: (0) an acute angle; (1) a narrow curve; (2) a wide curve.
Morgan and Czaplewski (2003: character 17, state 1) described the
mesostyle as a curved mesostylar crest, which may be similar to our
state 1. A wide curve occurs in taxa they did not analyze, such as
Noctilionidae.
Character 17.Labial projection of the metastyle in relation to
mesostyle: (0) absent; (1) present. This character may be similar to
character 35 of Czaplewski et al. (2003a). In their state 1 they

Vol. 92, No. 2

described the 4th commissure (postmetacrista) as being longer than

the other commissures (preparacrista, postparacrista, and premetacrista), which in our character corresponds to a labial projection of
the metastyle in relation to the mesostyle.
Character 18.Number of folds in the ectoflexus: (0) 2; (1) 1.
Character 19.Metacone of M3: (0) present; (1) absent. Freeman
(1981a: character C66), Wetterer et al. (2000: character 64), and
Czaplewski et al. (2003a: character 31) referred indirectly to the
presence or absence of the metacone of M3 when they described the
shape of the M3 ectoloph.
Character 20.Position of the metacone of M3: (0) placed
lingually in relation to the metacone of M1M2; (1) same position as
the metacone of M1M2; (2) placed labially in relation to the
metacone of M1M2.
Character 21.Lingual expansion of inner face of the metacone of
M3: (0) absent; (1) present.
Character 22.Mesostyle of M3: (0) same size as the mesostyle of
M1M2; (1) smaller than the mesostyle of M1M2; (2) absent.
Character 23.Position of the mesostyle of M3: (0) same position
as in M1M2; (1) placed lingually in relation to mesostyle of M1M2.
Character 24.Protocone of M3: (0) same size as the protocone of
M1M2; (1) smaller than the protocone of M1M2; (2) absent.
Character 25.Preparacrista of P4: (0) directed anteriorly; (1)
aligned at the center; (2) directed posteriorly.
Character 26.Lingual basin of P4: (0) large or medium; (1) small.
Czaplewski et al. (2003a: character 30) described the lingual portion of
P4, but the states that they proposed do not coincide with ours.
Character 27.Cusp in the lingual basin of P4: (0) present; (1)
absent. Czaplewski et al. (2003a: character 25) described a lingual
cingular cusp of P4 as absent, weak, or strong.
Character 28.Anterolabial basin of P4: (0) absent; (1) present.
Character 29.Anterolabial accessory cusp of P4: (0) absent; (1)
present. Morgan and Czaplewski (2003: character 14) described a
labial cingular cusp of P4 as absent, weak or strong.
Character 30.Relative size of P3: (0) at least one-third the size of
P4; (1) smaller than one-third the size of P4; (2) absent. Morgan and
Czaplewski (2003: character 49) considered P3 normal, greatly
reduced, or absent. Simmons and Geisler (1998: character 18) and
Gunnell and Simmons (2005: character 20) described the variation in
number of upper premolars as 3, 2, or 1 on each side of the arcade.
Simmons and Conway (2001: character 37) and Wetterer et al. (2000:
character 56) considered P3 present or absent.
Character 31.Relative size of P2: (0) at least one-third the size of
P4; (1) smaller than one-third the size of P4; (2) absent. Morgan and
Czaplewski (2003: character 29) described the size of P2 as normal or
greatly reduced, and Simmons and Conway (2001: character 36)
considered P2 present or absent.
Character 32.Relative position of P2: (0) centered in the
toothrow; (1) displaced lingually or labially in the toothrow.
Character 33.Lingual basin of P2: (0) absent; (1) present.
Character 34.Cross section of the main cone of C: (0) round; (1)
oval; (2) slender.
Character 35.Incisor-directed crest of the main cone of C: (0)
present; (1) absent.
Character 36.Lingual basin of C: (0) present; (1) absent.
Character 37.Lingual fold on the main cone of C: (0) absent; (1)
present.
Character 38.Posterior cusp of C: (0) absent; (1) present.
Character 39.Anterior cusp of C: (0) absent; (1) present.
Character 40.Anterolabial cingulum of C: (0) well marked; (1)
faint; (2) absent.

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FRACASSO ET AL.UPPER DENTITION AND PHYLOGENY OF BATS

Character 41.I2: (0) present; (1) absent. Wetterer et al. (2000:

character 48) also considered I2 present or absent.
Character 42.Diastema between I1 and I1: (0) present; (1)
absent.
Character 43.Position of I1 relative to a line drawn between the
2 upper canines: (0) anterior to C; (1) aligned with C.
Character 44.Relative size of I1: (0) I1 , I2; (1) I1 5 I2; (2) I1
. I2. Wetterer et al. (2000: character 49) and Simmons and Conway
(2001: character 30) described the condition of I2 as being smaller
than I1 or both I1 and I2 being approximately equal in size. The
condition of I1 being smaller than I2 was not present in the taxa
analyzed by those authors.
Character 45.Number of cusps on I1: (0) 1; (1) 2; (2) 3.
Simmons and Conway (2001: character 29) described I1 as tapering
to a blunt point or bilobed.
Character 46.Number of cusps on I2: (0) 1; (1) 2; (2) 3.
Character 47.Crown of I1: (0) in the form of a cone; (1) in the
form of a crest.
Character 48.Labial cingulum of I1: (0) present; (1) absent.

APPENDIX III
Character state matrix including our dental characters and the
characters of Gunnell and Simmons (2005). Polymorphic states are
indicated by square brackets. Missing or unknown states are indicated
by a question mark (?).
Eulipotyphla.
100100110010000000010111?00111100001000001021001000?0000100000000100000???0??00???0??????????????????????????????????00000000000000???1?001000??0000??0?????????????????????????201?100??????00000????00??10?0??100000??0?1????0???10?1?1?1????
Icaronycteridae.
100110110000000110??????????????????????????????????????100????00??1??0?0????1??????????????????????????????????????11000000000101?12100000000?1000010010????????????????????????1000?10?0??0?00001????011001?0?????100?00?????????????????????
Rhinolophidae.
100011110002100000010001?000021101000000100?1?01002000110100111?0113[01]?011011011132[0123]00000101100000020100100000[12]0?101031111021111202110101100030221210100121110010111111110??2101122100111??????311130???00000201????011111011110001000?301020010
Hipposideridae.
100[01]1111[01]012100[01]00[01]1000[01][01]100[01]21[01]010001[01]2101?1?01002000110100111?0113[01]?011011011132[0123]00000101100000020[01]00100000[012]0?10[12]0311111211112021[12]01000000[13]02212101000211?0?????11111?0??2???022100111000100[123]11130011100002011011011111111110001????????20010
Megadermatidae.
1010011111?2[01]110111?000010[01]112[12]10[12]01[01]2211???????012?00110110111?00120111011111102100100010101030002010010000010?100001111001110222110101000030221210100121211010111111111112111121100111???????01130???010102101011011111011110001?00?2110?1001
Rhinopomatidae.
0?02011100?21002000201111001122002010001100?0?010110001100101???01121?00020111103230001010101020000010010101210?1120211100001001220111000000001?020010010010011011100-

431

0101012110011000111???????00130???01010111?????10101011010001?00?3010?[01]001
Nycteridae.
100001110012100000020001?01002200000011201112211013012210100111?0012[01]?1110102111321000010??01010002010010000010?1000011110010102220230110000101?0210100121201010111011101111110111000111?11000?11130???00110110?????11311011100000?????????1000
Emballonuridae.
[01]00[012]0[01][01][01]0012100100[01]20[01][01][01][12]00[01]
[01]2100[12][01][01]0[01][01][012][01]0[01][12]000101000[01][12]
10010010?0012110002110111210000110??01000000010010010110?0?10111100000001[01][12]02[12]1011000102100[01]0100120201101001011101012000111110111?10000011131???01010110?????101110110000000?013?1021100
Myzopodidae.
100020110010000110010001201001100201100000100001110?????20010??211121?000111011???01101011?01131112110011000020?0?00411100000002120221001010302101?110011????????????????????????2000111???????11110???01110110?????1011110??00001?????????????
Mystacinidae.
101[01]10110002100200010011?000020011000001110?0?000100122?20010??101111?001001011???01110110011030001111010100020?101011110000000[01]000121010010311?000010?01????????????????????????2011111??????001111???01011110?????10011001?001010????????????
Phyllostomidae (Phyllostominae).
[01]00[12][01][01]110[01]1[012][01][01][01][01][01]01?0[12]112
[01][01][01]02[01][01][01][01]0[01][01]00[01][01]1[01]21[01]1[01]
01200000200101110111110102000111010111010??110[013]00010
[01]00100101001?0[01]011110000000101021100000031200100101020211010101011101112011022000111111001[03]01110010010101101011110111000?011121111122120100
Mormoopidae.
[01]00[02][01][01]110010000100000111?0[01]102101[02]01100[12]01[01]21011110001[012]02001011101121?011000011???2111000??11000001110010010100?10101111000000002202210100102121011010102021111010101110111201101210011101101110111000011010110?????10111000?011121????????0???
Noctilionidae.
1012[01]1110011110200020110000002200000100001020000010011212001011?01120?011200?111010111010??11030001010010010100?102011110000000121022100001001200100100010211010001011101112001111100111?01111401132???01010110?????10111010?011111101122020100
Furipteridae.
100020[01][01]0000000100010001?10102000211010100011101110?11212010[01]11111121?010111011???01110010011111112110011100000?1102111110000002220231111110211?020021001????????????????????????2000111???????11120???11010110?????1011100??00011?????????0???
Thyropteridae.
2100[12]0[01]10010000100001111200100000111000000021101110110112001011111121?000111?11???0111000??111311121100[01]?000?20?110211111100000222021111011020200000101010110011101001101012000012000111???????11110???01110111?????1011110??000111?1021?1?????
Natalidae.
1000101100[01][02][01]001000[12]00012[01][01]000[01]00101100[01]00020001110?01212001011001121?000011011???0111001001-

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JOURNAL OF MAMMALOGY

1111112110001100020?1000111110000001211201110100211?0201210110210011101000001211001012000111?00010?11120???11010110?????10111001000001?????????0???
Tomopeatinae.
320010110002100200010111?000022000010000101?0?00010?????20100???01121?000211?11???0121001001100000211100000002110?20111110000001010120000011311?000122011????????????????????????2111111???????01120???01111110?????1001101??000???????????????
Molossinae.
[123][012]00[12][01][01][01]000[01][01]00[12]00[01][12]0[01][01]10[01]00[01]2[12]0000[01][01]00[01]1[01][01]?[01]?0[01]01011[01][12]12010111?01121?00021111110101210[01]???1100000211100000002110?2011111000000111122000001131200001221010201110101001001001001042111111110010?011211010111111001111100110110000010000311120100
Antrozoinae.
[01]000211110100101001?0[12]110110022000000000101?0?00011?????20100???01121?000211011???0000101011100000011101100012111120111100000002010121000010311?00011000102???????1000001101001032011111???????01120???01110110?????10111011000001?????????????
Vespertilioninae.
10002[01]110010000110[01]10[12]110[01]000220000[01]0000[01]01?1?0[01]010101212010011?01121100021101111201110011?1101000111101100012100?201111000000021[12]02[12]10100103121-

Vol. 92, No. 2

0001101010210101011001000101001032011111100001?0112001001110110?????10111011[01]0000100002111?0100
Myotinae.
211020[01][01]00100001000010110[01][01]00[01]100[01]00000[01]00011100010101212010011101121?000111011???01110111?1101000111100100012100?201111000000021202111100103121000111001011001000100100100100103200111110?0???01120010011101100111110111011000001000021112????
Miniopterinae.
2100101100100001000001110100020010010001000110000101000020100???01121?00021101110201110111?1101000111100100010100?20111100000002220231110110311?000112101????????????????????????2111111???????01122???01110110?????10111000?00001???????1?0100
Murininae.
[01]000[12]111[01]0100[01][01]00[01]1?02?[12]001002[01]01000
[01]0000010[01]000010?????20100???01121?000211?11???0111010??11010002111001010120?10201111000000031202111100003022020110011????????????????????????2001111???????01120???01110110?????10111011000001?????????????
Kerivoulinae.
100021110010000[01]100101110110[01]0000000000[01]00[01]20000110?????20100??201120?000211?111??0111010??1100000111110100012100?221111000000031202111101103121000110011????????????????????????2001??????????01120???01110110??????0111021000001?????????0100