New Myzopodidae (Chiroptera) from the Late Paleogene of Egypt: Emended Family Diagnosis and Biogeographic Origins of Noctilionoidea

Abstract

Myzopodidae is a family of bats today represented by two extant species of the genus Myzopoda that are restricted to the island of Madagascar. These bats possess uniquely derived adhesive pads on their thumbs and ankles that they use for clinging to smooth roosting surfaces. Only one fossil myzopodid has been reported previously, a humerus from Pleistocene deposits at Olduvai Gorge in Tanzania that was tentatively referred to the genus Myzopoda. Here we describe a new genus and two new species of myzopodids based on dental remains from Paleogene deposits in the Fayum Depression in Egypt, and provide an emended diagnosis for the family Myzopodidae. Phasmatonycteris phiomensis n. sp. is represented by four specimens from the early Oligocene Jebel Qatrani Formation and P. butleri n. sp. is known from a single specimen from the late Eocene Birket Qarun Formation. Together these specimens extend the temporal range of Myzopodidae by 36+ million years, and the geographic range by nearly 4000 kilometers. The new myzopodids, along with previously described bats from the Fayum and Australia, suggest that eastern Gondwana played a critical role in the origin and diversification of several bats clades notably including the superfamily Noctilionoidea, the majority of which live in the Neotropics today.

Full text

New Myzopodidae (Chiroptera) from the Late Paleogene
of Egypt: Emended Family Diagnosis and Biogeographic
Origins of Noctilionoidea
Gregg F. Gunnell
1
*, Nancy B. Simmons
2
, Erik R. Seiffert
3
1Division of Fossil Primates, Duke University Lemur Center, Durham, North Carolina, United States of America, 2Department of Mammalogy, Division of Vertebrate
Zoology, American Museum of Natural History, New York, New York, United States of America, 3Department of Anatomical Sciences, Stony Brook University, Stony Brook,
New York, United States of America
Abstract
Myzopodidae is a family of bats today represented by two extant species of the genus Myzopoda that are restricted to the
island of Madagascar. These bats possess uniquely derived adhesive pads on their thumbs and ankles that they use for
clinging to smooth roosting surfaces. Only one fossil myzopodid has been reported previously, a humerus from Pleistocene
deposits at Olduvai Gorge in Tanzania that was tentatively referred to the genus Myzopoda. Here we describe a new genus
and two new species of myzopodids based on dental remains from Paleogene deposits in the Fayum Depression in Egypt,
and provide an emended diagnosis for the family Myzopodidae. Phasmatonycteris phiomensis n. sp. is represented by four
specimens from the early Oligocene Jebel Qatrani Formation and P. butleri n. sp. is known from a single specimen from the
late Eocene Birket Qarun Formation. Together these specimens extend the temporal range of Myzopodidae by 36+million
years, and the geographic range by nearly 4000 kilometers. The new myzopodids, along with previously described bats
from the Fayum and Australia, suggest that eastern Gondwana played a critical role in the origin and diversification of
several bats clades notably including the superfamily Noctilionoidea, the majority of which live in the Neotropics today.
Citation: Gunnell GF, Simmons NB, Seiffert ER (2014) New Myzopodidae (Chiroptera) from the Late Paleogene of Egypt: Emended Family Diagnosis and
Biogeographic Origins of Noctilionoidea. PLoS ONE 9(2): e86712. doi:10.1371/journal.pone.0086712
Editor: Andrew A. Farke, Raymond M. Alf Museum of Paleontology, United States of America
Received September 25, 2013; Accepted December 9, 2013; Published February 4, 2014
Copyright: ß2014 Gunnell et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: This research was supported by National Science Foundation (http://www.nsf.gov/) Grant DEB 0949859 to NBS and NSF grants BCS-0819186 to ERS and
BCS-0416164 to E.L. Simons and ERS and The Leakey Foundation. The funders had no role in study design, data collection and analysis, decision to publish, or
preparation of the manuscript.
Competing Interests: The authors have declared that no competing interests exist.
* E-mail: gregg.gunnell@duke.edu
Introduction
Myzopodidae is a small family of insectivorous bats that are
today endemic to Madagascar. Two living species are recognized:
Myzopoda aurita, which was described by Milne-Edwards and
Grandidier in 1878 [1], and M. schliemanni, which was named
nearly 120 years later [2]. Both species are characterized by
unique morphological specializations that diagnose the family
including large, non-pedicellate suction pads on the thumb and
ankle, fusion of the tragus to the pinna, and partial obstruction of
the external auditory meatus by a mushroom-shaped process
[2–3]. Long considered a member of the superfamily Vespertilio-
noidea [4–6], Myzopodidae was transferred to the superfamily
Nataloidea by Simmons [3] but subsequent analyses based on
extensive molecular data sets indicate that Myzopodidae is actually
a basal member of the superfamily Noctilionoidea [7–12].
Regardless of whether myzopodids are viewed as basal vesperti-
lionoids or basal noctilionoids, dating analyses unambiguously
place the origin of Myzopodidae in the Eocene [7–10,12].
Myzopodids have long been considered intriguing for a variety
of reasons including their unusual roosting habits – they use the
suction pads on their wrists and ankles to cling to the smooth
surfaces of broad leaves such as those of Ravenala [13]. In this they
are similar to New World Disk-winged bats of the family
Thyropteridae, but anatomical and evolutionary analyses concur
that the wrist and ankle discs in these two groups evolved
convergently [3,14–15]. The modern geographic distribution of
Myzopodidae is also of interest since close relatives of myzopodids
include lineages endemic to Australia and New Zealand
(Mystacinidae) and the Neotropics (Noctilionidae, Furipteridae,
Thyropteridae, Mormoopidae, and Phyllostomidae) rather than
Africa [7–9].
The modern fauna of Madagascar encompasses six orders of
mammals and includes both endemic and introduced species [16].
The endemic terrestrial mammals (lemuroid primates, tenrecid
afrotherians, euplerid carnivorans, nesomyine muroid rodents) can
trace their ancestry to Africa, where sister groups to these
Malagasy clades occur today [17]. No definitive fossil lemurs
have yet been found on the African mainland but tenrecoids,
euplerid carnivoran sister groups and nesomyines are known from
Africa [18–22].
The extant bat fauna of Madagascar includes 49 species
representing seven families (Pteropodidae, Emballonuridae, Hip-
posideridae, Vespertilionidae, Miniopteridae, and Myzopodidae).
Many species have been described quite recently –17 species since
2005, with more being described every year. Endemism in
Malagasy bats is quite high, with 37 species (55% of the fauna)
known only from Madagascar and nearby islands but not from the
mainland. In the extant Malagasy fauna, Myzopodidae is the only
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endemic family; the other bat families known from Madagascar
have many mainland representatives in Africa and Asia.
The fossil record of bats from Madagascar is sparse and includes
only eleven late Pleistocene and Holocene species (Table 1) [23].
These include two hipposiderid species known only as fossils [24],
seven endemic species (known only from Madagascar, the
Seychelles and Comoro Islands today), and one species also
known from the African mainland (Hipposideros commersoni).
There is no known fossil record of Myzopodidae from
Madagascar. Butler [25] mentioned the presence of a humerus
seemingly from a myzopodid from Olduvai Gorge in Tanzania,
but he did not describe the specimen in detail, only noting that the
family apparently was present on the African mainland in the early
Pleistocene [5,26]. Until now, no other fossil myzopodids have
been reported. Here we document the presence of two new species
of a new genus of myzopodid from the late Eocene and early
Oligocene in North Africa, thus extending the geographic and
temporal range of the family substantially.
Like many bat families, Myzopodidae is broadly accepted as
monophyletic but while simple diagnoses exist [3,6,27], there is no
comprehensive diagnosis or description available that includes the
details of dental morphology necessary for assessing affinities of
incomplete fossils. In addition to describing two new myzopodid
species from the early Paleogene, we here provide a comprehen-
sive diagnosis and description for the family including details of
dental morphology. Because the new myzopodid fossils are known
only from lower dentitions, most of the characters noted are based
on observations from extant Myzopoda. However, lower premolar
and molar characters can also be evaluated in the fossil
myzopodids described here, and the emended diagnosis and
description below reflects morphology of these taxa.
Permissions
Permissions for collecting fossil specimens were obtained from
the Egyptian Mineral Resources Authority and the Egyptian
Geological Museum. All necessary permits were obtained for the
described study, which complied with all relevant regulations.
Methodological Considerations
Nomenclatural Acts: The electronic edition of this article
conforms to the requirements of the amended International Code
of Zoological Nomenclature, and hence the new names contained
herein are available under that Code from the electronic edition of
this article. This published work and the nomenclatural acts it
contains have been registered in ZooBank, the online registration
system for the ICZN. The ZooBank LSIDs (Life Science
Identifiers) can be resolved and the associated information viewed
through any standard web browser by appending the LSID to the
prefix ‘‘http://zoobank.org/’’. The LSID for this publication is:
urn:lsid:zoobank.org:pub: urn:lsid:zoobank.org:pub:DA73AE7E-
88C0-4EC8-B068-6E4FDBEA84B4. The electronic edition of
this work was published in a journal with an ISSN, and has been
archived and is available from the following digital repositories:
PubMed Central, LOCKSS.
Dental Terminology: In order to avoid confusion, we refer to
the posterior two premolars of bats by their traditionally
interpreted homologies rather than those described for placental
mammals by O’Leary et al. [28], thus we recognize these teeth as
P3/p3 and P4/p4 rather than P4/p4 and P5/p5. However, we
recognize the anterior-most premolar of bats as P1/p1, rather
than the more traditional P2/p2 following the analysis of Giannini
and Simmons [29].
Results
Mammalia Linnaeus, 1758.
Chiroptera Blumenbach, 1779.
Noctilionoidea Gray, 1821.
Myzopodidae Thomas, 1904.
Emended Family Diagnosis and Description
Extant bats referred to this family share many traits that may be
diagnostic for the family as a whole. Characters visible externally
include: large, non-pedicellate suction pads present on the thumb
and ankle; pinna large; tragus fused to the pinna; external auditory
meatus partially obscured by a fleshy, mushroom-shaped process;
digit II of wing reduced to metacarpal only (phalanges absent);
digit III of wing with a fully-ossified nub-like third phalanx; calcar
present; tail long and extends beyond the uropatagium; females
with a transverse genital opening, short clitoris, and lacking pubic
nipples.
Skull with orthoclivous premaxilla; left and right premaxillary
bodies well developed and in contact medially (Figure 1) but
partially separated by a midline notch; palatine process of
premaxilla complete, with lateral and medial flanges enclosing a
pair of incisive foramina; nasal process of premaxilla absent;
nasoincisive suture limited to a point contact between premaxilla
and nasal; maxilloincisive notch absent; jugal small, does not
contact lacrimal; postorbital process absent; hard palate extends
posteriorly into orbital region; medial accessory palatine foramen
absent; malleus with large orbicular apophysis and single tensor
tympani muscle; aqueductus cochleae small or absent; epitym-
panic recess and stapedial fossa both shallow and broad; fenestra
rotundum not enlarged; tympanic annulus inclined, does not form
tubular external auditory meatus; cochlea large and phaneroco-
chlear; stylohyal with large, ax-shaped expansion at cranial tip
where it articulates with the cochlea; basihyal bar shaped and
lacking an entoglossal process; angular process of dentary elongate,
projects at level of occlusal plane of toothrow. Of these cranial
characters, the only feature that is unique is the ax-shaped
expansion of the cranial tip of the stylohyal.
Postcranial skeleton with posteriorly directed ventral accessory
processes present on cervical vertebrae C2-C5; no fusion of
posterior cervical or anterior thoracic vertebrae; ribs not fused to
vertebrae; rib 1 not fused to manubrium; rib 2 contacts sternum at
manubrium-mesosternum joint and articulates via a costal
cartilage; ribs with narrow anterior lamellae and wide posterior
lamellae; manubrium short, length less than 2x width, and with
small anterior face; ventral process of manubrium laterally
compressed to form a keel, projects at obtuse angle from body
of manubrium; mesosternum narrow; xiphisternum without keel
and not laterally flared; acromion process of scapula without
median shelf; tip of acromion process with triangular anteromedial
projection; dorsal articular facet of scapula faces dorsally and
consists of a large, flat surface that is clearly separate from glenoid
fossa; blade of scapula with 3 facets that are subequal in width;
axillary border of scapula with a bladelike lip; anteromedial flange
present on scapula; coracoid process short, stout, curved
ventrolaterally, and lacking a flared tip; suprascapular process
absent; clavicle articulates with scapula between acromion process
and coracoid process; humerus with round head; trochiter extends
beyond head; distal humerus with facets displaced from long axis
of shaft; epitrochlea broad; entepicondylar foramen absent;
olecranon fossa absent; ulna with reduced olecranon process;
ulnar patella present.
Posterior vertebral column with no fusion of lumbar vertebrae;
sacrum extends posterior to level of acetabulum; sacral lamellae
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Table 1. List of extant and extinct bats currently known from Madagascar.
Family Genus Species Type Locality Endemic
Fossil
known Fossil Only Notes & References
Pteropodidae Eidolon duprenum Nossi Be
´X X Fossils from Anjohibe Cave
[24] & d’Andrahomana [50]
Pteropus rufus Madagascar X X Fossils from d’Andrahomana
[50]
Rousettus madagascar-ensis Beforona X X Fossils from Anjohibe Cave
[24] & d’Andrahomana [50]
Hipposideridae Hipposideros commersoni Fort Dauphin X X Fossils from Anjohibe Cave
[24], Tsimanampetsotsa [51] &
d’Andrahomana [50]
Hipposideros besaoka Anjohibe Cave X X [24]
Paratriaenops auritus Die
´go-Suarez X [52]
Paratriaenops furculus Tule
´ar X Fossils from ?Anjohibe Cave
[24], Tsimanampetsotsa [51] &
d’Andrahomana [50]
Triaenops menamena Province de
Mahajanga
X[53]
Triaenops rufus Eastern Madagascar X [54]
Triaenops goodmani Anjohibe Cave X X [24]
Emballonuridae Taphozous mauritianus Non-Malagasy [55]
Coleura kibomalandy Province d’Antsiranana X [56]
Paraemballonura tiavato Province d’Antsiranana X [57]
Paraemballonura atrata Interior Madagascar X X Fossils from
Tsimanampetsotsa [51]
Nycteridae Nycteris madagascar-iensis North of Ankarana X [58]
Myzopodidae Myzopoda aurita Madagascar X [1]
Myzopoda schliemanni Province de Mahajanga X [2]
Molossidae Chaerephon atsinanana Province de
Fianarantsoa
X[59]
Chaerephon jobimena Province d’Antsiranana X [60]
Chaerephon leucogaster Morondava [61]
Chaerephon pumilus Non-Malagasy [62]
Mops leucostigma Antananarivo X X Fossils from d’Andrahomana
[50]
Mops midas Non-Malagasy [63]
Mormopterus jugularis Antananarivo X X Fossils from
Tsimanampetsotsa [51],
Ankilitelo & d’Andrahomana
[50]
Otomops madagascar-iensis Soalala X X Fossils from Ankilitelo [50]
Tadarida fulminans Betsileo [64]
Vespertilionidae Eptesicus matroka Betsileo X [65]
Scotophilus marovaza Province de Mahajanga X [66]
Scotophilus robustus Madagascar X [54]
Scotophilus tandrefana Parc National de
Bemaraha
X[67]
Pipistrellus hesperidus Non-Malagasy [68]
Pipistrellus raceyi Province de
Fianarantsoa
X[69]
Hyposugo anchietae Non-Malagasy [70]
Neoromicia malagasyensis Northeast of Tule
´ar X [71]
Neoromicia melckorum Non-Malagasy [72]
Neoromicia nanus Non-Malagasy [73]
Neoromicia robertsi Anjozorobe X [74]
Neoromicia somalicus Non-Malagasy [75]
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broad; ascending process of ilium does not extend dorsal to
iliosacral joint; dorsal ischial tuberosity absent; pubic spine present
and straight, not curved dorsally; bar-like pubic symphysis present
in males; distal femur shaft straight; fibula thin and threadlike; foot
digits II-IV with only two phalanges on each digit; digital tendon
locking mechanism absent from feet.
Dental formula I2/3, C1/1, P3/3, M3/3 = 38 (Figs. 1–3); upper
incisors proodont and retroclivous; I1 with crown clearly distinct
from shaft, large main cusp that tapers to a point, and lacking a
distal accessory cusp or lingual cingulum; I1 crown height less than
that of I2; I1 separated from lower incisors by a large gap when
jaw is in occlusion; I2 with well-developed crown including a
lingual cingulum; small diastema present between I2 and C; upper
C tall, height of C .5x the height of I1; upper C with complete,
raised labial cingulum, no anteromedial accessory cusp, and
anteromedial groove present; upper C with narrow lingual
cingulum lacking a cusp or indentation; upper C with narrow
distal cingulum, posteromedial ridge present, posterolingual and
posterolateral surfaces flattened, posterolateral accessory cusp
either absent or just a trace of cusp present; P1 single-rooted, with
well-developed crown that is greater in diameter than the root,
with bluntly pointed central cusp encircled by a complete
cingulum on all sides; P1 either in line with P3 and P4 or slightly
lingually displaced; P1 crown length longer than that of P3; P3
single-rooted, lies in line with P4 and C; P3 with well-developed
cingulum surrounding central cusp and no accessory cusps;
distinct diastema present between P3 and P4; P3 much smaller
than P4 in all dimensions; P4 with large lingual cingulum that
forms a lobe but which lacks a cusp and does not extend as far
medially as protocone of M1; P4 anterior and labial cingula well
developed and lacking accessory cusps; P4 with single postpar-
acrista that decreases sharply in height and does not extend to the
distal edge of the tooth; no diastema present between P4 and M1;
primary cusp of P4 not in line with axis through M1 and M2
paracones, which are more lingually placed; length of P4 less than
length of M1.
Upper molars tribosphenic with W-shaped ectoloph, acute
angle between postparacrista and premetacrista, and lacking an
ectocingulum; no diastema present between adjacent molars; M1
and M2 with protocone and paracone subequal in height,
metacone taller than either of the other major cusps, mesostyle
present at contact between postparacrista and premetacrista,
mesostylar crest absent, paraconule and metaconule absent,
parastyle present and not separated from preparacrista, single
shallow ectoflexus present between parastyle and metastyle,
preparacrista and postparacrista subequal in length, postmeta-
crista longer than premetacrista, postprotocrista oriented distola-
bially toward metacone but terminates before contacting the base
of that cusp, hypoconal shelf and hypocone absent, endoloph
absent; M3 present, reduced to 50–75% the size of M2 when seen
in occlusal view, parastyle and mesostyle present, metacone
developed as a distinct cusp, postparacrista shorter than pre-
metacrista, postmetacrista absent, protocone present, hypoconal
shelf and hypocone absent.
Three somewhat procumbent lower incisors present, each tooth
bilobed or trilobed; no diastema present between right and left i1;
Table 1. Cont.
Family Genus Species Type Locality Endemic
Fossil
known Fossil Only Notes & References
Myotis goudoti Madagascar X X Fossils from Anjohibe Cave
[24]
Miniopteridae Miniopterus aelleni Province d’Antsiranana [77]
Miniopterus brachytragos Province de Mahajanga X [78]
Miniopterus egeri Province de Toamasina X [79]
Miniopterus fraterculus Non-Malagasy X [66]
Miniopterus gleni Tule
´ar X X Fossils from Ankilitelo &
d’Andrahomana [50]
Miniopterus griffithsi Province de Toliara X [77]
Miniopterus mahafalie nsis Province de Toliara X [77]
Miniopterus majori Betsileo [79]
Miniopterus manavi Betsileo 79
Miniopterus petersoni Province de Toliara X [76]
Miniopterus sororculus Province de Fianarantsoa X [80]
Erroneous &
unconfirmed
records
Pteropodidae Pteropus niger Mascarene & Reunion
Islands
Madagascar records probably
erroneous
Molossidae Mops niveiventer Congo Madagascar records
erroneous, represent M.
leucostigma
Mormopterus acetabulosus Mauritius No confirmed Malagasy
specimens
Vespertilionidae Scotophilus borbonicus Reunion Island No confirmed Malagasy
specimens
doi:10.1371/journal.pone.0086712.t001
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i1 conspicuously smaller than i2 and i3; i2 and i3 similar in crown
height; alveoli for incisors evenly spaced; small diastema present
between i3 and c; right and left lower canines with bases widely
separated and with laterally divergent tips; c tall and slender with
subterete shaft, anterior cuspule absent, posterior cingulid present
but small, posterobasal cuspule weakly developed, labial cingulid
well developed and lacking cuspules; single-rooted p1 present,
large and cuspidate, with distal cuspule variably present; single-
rooted p3 present and in line with surrounding teeth, not offset
lingually or labially; crown length of p3 much less than that of p1;
p3 premolariform with a single central cusp surrounded by a
complete cingulid that lacks cuspules; crown height of p3 much
less than that of p4; p4 double rooted, with weakly-developed
lingual cingulid that variably present, and a well-developed labial
cingulid that is of roughly uniform width along the length of the
tooth; p4 lacking a distinct paraconid or metaconid; p4 with tall
and sharp protoconid, lingual lobe absent, talonid absent; crown
length of p4 approximately 50–75% of the crown length of m1.
Lower molars tribosphenic, lacking a lingual cingulid, trigonid
fovea open lingually (very broadly on m1), labial cingulid well
developed and complete, labial cingulid continuous with well-
developed precingulid; cristid obliqua on lower molars gently
curved labially where it contacts postvallid ( = posterior wall of
trigonid), lacking a notch, contacts postvallid at a point midway
between metaconid and protoconid; protoconid higher than
hypoconid; paraconid lower in height than metaconid; paraconid
of m1 very robust, anteriorly angled, and positioned labial to line
between metaconid and entoconid; metacristid, if present,
markedly shorter than paracristid; entoconid well developed and
tall, entocristid short, straight, and does not contact trigonid;
paraconid, metaconid, and entoconid of m2–3 in alignment in
occlusal view; hypoflexid shallow; m1 and m2 subequal in length,
with small, low hypoconulid located on lingual edge of tooth and
‘‘twinned’’ with entoconid, either nyctalodont or myotodont,
width of talonid basin greater than that of trigonid; double-rooted
Figure 1. Upper dentitions of extant
Myzopoda
species.
Myzopoda aurita (AMNH 257130), right maxillary dentition with I1-M3
in occlusal view (A) and in close-up occlusal view of right I1-P4 (C).
Myzopoda schliemanni (AMNH 277725), right maxillary dentition with I1-
M3 in occlusal view (B) and in close-up occlusal view of right I1-P4 (D).
Scale bars equal 1 cm.
doi:10.1371/journal.pone.0086712.g001
Figure 2.
Phasmatonycteris phiomensis
, YPM 24198, Holotype,
left dentary with p3-m3. (Note: m3 talonid has been subsequently
damaged, photographs taken before damage occurred). A, Stereo-
photographs of holotype in occlusal view; B, close-up of p3-4 in
occlusal view; C, labial view of holotype; D, lingual view of holotype.
Scale bar for A, C–D = 1 mm, scale bar for B= 0.50 mm.
doi:10.1371/journal.pone.0086712.g002
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m3 with hypoconulid either present or absent, nyctalodont if
hypoconulid is present; m3 crown either subequal in length to m2
or slightly shorter than m2; paraconid robust, approximately the
same size as protoconid; entocristid on m3 oriented parallel to long
axis of tooth; postcristid oriented either perpendicular to long axis
of tooth or directed posterolabially at an oblique angle; m3 talonid
either subequal to or narrower than trigonid.
The morphology of the suction pads, tragus, pinnae, and
stylohyal are unique among bats and serve to distinguish
Myzopodidae from all other families. One dental feature is unique
to myzopodids among bats – the large, anteriorly angled and
slightly labially shifted paraconid on m1. Although the paraconid
is lower than the protoconid, it is of equal size and the anterior
angulation broadly opens the m1 trigonid lingually and produces a
steeply sloping trigonid fovea. Other dental features, while not
unique to myzopodids, are distinctive of the family. The gently
labially curving cristid obliqua is uncommon among bats and the
complete lack of hypocones and hypocone shelves are also
somewhat unusual.
Phasmatonycteris gen. nov. urn:lsid:zoobank.org:act:693EEE77-
DD52-408B-B404-B689A43706AC.
Figure 3. Lower dentitions of fossil and extant myzopodids. A,
Phasmatonycteris butleri, CGM 83761, Holotype, right dentary (reversed)
with m2-3 in occlusal view; B, Myzopoda aurita, Field Museum of
Natural History (FMNH) 194176, left dentary with i1-m3 in occlusal view;
C, Phasmatonycteris phiomensis, YPM 24198, Holotype, left dentary with
p3-m3 in occlusal view; D, Myzopoda schliemanni, FMNH 187604, right
dentary with i1-m3 in occlusal view; E, P. phiomensis, YPM 24195, left
dentary with m3 in occlusal view; F, P. phiomensis, YPM 24196, left
dentary with m2 (broken) in occlusal view; G, P. phiomensis, YPM 24197,
left dentary with m3 in occlusal view. Scale bars = 1 mm.
doi:10.1371/journal.pone.0086712.g003
Figure 4. Eocene- Oligocene stratigraphy in the Fayum
Depression, Egyptian Western Desert. The distribution of fossil
bats from Quarries I, L-41, and BQ-2 is shown in relation to Fayum
magnetostratigraphic polarity zones, epoch and stage boundaries, and
correlated radioisotopic dates.
doi:10.1371/journal.pone.0086712.g004
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Generic Diagnosis
Differs from extant Myzopoda in having a relatively longer p4; a
p3 that is more elongate and narrow, positioned labially and more
closely appressed to p4; m1 and m2 of same length and m3 only
slightly reduced in length; m1 with an especially robust and
anteriorly angled paraconid; and all molars with more robust
labial cingulids.
Etymology
Phasma(to), Greek for apparition or spectre, in reference to the
long ghost lineage connecting Fayum myzopodids with extant
forms, and Nykteris, Greek for bat.
Phasmatonycteris phiomensis sp. nov. urn:lsid:zoobank.org:act:
94812861-7911-438E-BD80-AB849541605B (Figs. 2A–D, 3C, E–
G).
Holotype
Yale Peabody Museum (YPM) 24198, left dentary with p3-m3.
Locality and Horizon
Fayum Quarry I, 242 meter level, Upper Gebel Qatrani
Formation, Early Oligocene, Rupelian (,30 Ma), Fayum
Depression, Western Desert, Egypt (Fig. 4).
Referred Specimens
YPM 24195, left dentary with m3 (Fig. 3E), YPM 24196, left
dentary with m2 (3F), YPM 24197, left dentary with m3 (Fig. 3G),
all from Fayum Quarry I.
Specific Diagnosis
Differs from P. butleri (new species) in having sub-myotodont
lower molars, a relatively larger m3 compared to m2, deeper
hypoflexids on m2-3, more steeply sloping entocristids on m2-3,
and m2 with a more robust, distinct, and anteriorly-oriented
paraconid.
Etymology
Phiom, Greek for the Fayum Region of Egypt’s Western Desert.
Description and Comparison
The lower dentition of P. phiomensis is represented by four
specimens – the holotype dentary preserving p3-m3, and three
referred dentary fragments, two with m3 in place and the other
including a broken m2. Measurements of specimens discussed in
text are included in Table 2.
Lower p3 is dominated by a central cusp surrounded by a
relatively thick, broad cingulid. The base of p3 is rounded
mesiolabially with a straight margin defining the distolingual
portion of the tooth (Fig. 2A–B). The cingulid is broadest
mesiolingually where it forms a small shelf. It is somewhat
narrower mesially, then broadens slightly and wraps around the
labial border to join the distolingual portion of the cingulid. The
posterior portion of p3 is positioned labial to the anterior
mesiolabial margin of p5 and is appressed to the basal cingulid
of p4.
The p3 morphology of P. phiomensis is similar to that of the p3s of
extant Myzopoda but there are some differences. In M. aurita the p3
has a nearly perfectly round base and central cusp with the
cingulid being slightly broader lingually (Fig. 3B). The p3 in M.
aurita is placed directly anterior to p4 and is slightly over-lapped
mesiolingually by p1. The p3 of M. schliemanni (Fig. 3D) has a
rounded central cusp but the base is more ovoid and is wider
labiolingually than it is mesiodistally long. The basal cingulid is
Table 2. Measurements of lower cheek teeth of extant Myzopoda and extinct myzopodid Phasmatonycteris.
Specimen Genus/Species
p4
length
p4
width
p5
length p5 width
m1
length
m1 tri
width
m1 tal
width
m2
length
m2 tri
width
m2 tal
width
m3
length
m3 tri
width
m3 tal
width
FMNH 194176
Myzopoda aurita
0.35 0.42 0.82 0.61 1.48 0.57 0.83 1.39 0.69 0.89 1.36 0.72 0.72
FMNH 187604
Myzopoda schliemanni
0.28 0.36 0.69 0.54 1.42 0.67 0.83 1.33 0.69 0.86 1.22 0.68 0.64
YPM
24198
Phasmatonycteris
phiomensis
0.39 0.41 1.17 0.68 1.64 0.67 0.86 1.58 0.89 0.94 1.38 0.91 0.67
YPM
24195
Phasmatonycteris
phiomensis
1.54 1.01 0.64
YPM
24196
Phasmatonycteris
phiomensis
0.98 0.99
YPM
24197
Phasmatonycteris
phiomensis
1.59 1.02 0.76
CGM
83761
Phasmatonycteris
butleri
1.46 0.93 0.97 1.22 0.74 0.68
doi:10.1371/journal.pone.0086712.t002
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broadest lingually and labiomesially and narrows distolingually. As
in M. aurita the p3 of M. schliemanni is directly anterior to p4 and its
mesiolingual portion is overlapped by p1.
The crown of the p4 of P. phiomensis is damaged, with the tip and
the distal wall of the protoconid having been broken away (Fig. 2B).
It is clear, however, that like extant Myzopoda, the p4 of P.
phiomensis lacked both a paraconid and a metaconid. The p4 of P.
phiomensis is relatively longer compared to m1 (71% of m1 length)
than is the p4 in either M. schliemanni (49%) or M. aurita (55%). The
crown is dominated by a relatively large protoconid that is
surrounded by a basal cingulid that is slightly extended into a weak
talonid shelf along the distal margin of the tooth. There is a
distinct preprotocristid present that extends to the margin of the
anterior cingulid as is often found in M. aurita. The cingulid is
broadest anteriorly and narrows both labially and lingually to join
the distal shelf. The anterior-most extension of the cingulid
overlaps the distolingual portion of p3.
The m1 of P. phiomensis shares the distinctive morphology of
extant Myzopoda (Figs. 2A, C–D, 3C). The trigonid is broadly open
with a wide notch separating the paraconid and metaconid. Like
extant myzopodids, the m1 paraconid is large, mesially angled and
connected to the protoconid by a curving, notched paracristid.
The protoconid is the tallest trigonid cusp and is positioned
marginally. The metaconid is nearly as tall as the protoconid and
is located almost directly lingually opposite the protoconid unlike
in M. schliemanni and M. aurita where the metaconid is often slightly
more distal compared to the protoconid.
The hypoconid is low and marginally placed and is connected to
the trigonid by a sharply defined cristid obliqua that joins the
postvallid labial of center like in extant myzopodids. The
entoconid is more elevated and bulbous and is connected to the
postvallid by a relatively short, straight and high entocristid. The
m1 is submyotodont, unlike extant myzopodids that have fully
myotodont molars. Myotodont molars have a postcristid that
connects the hypoconid and entoconid, completely excluding the
hypoconulid from the talonid and leaving this cusp as a small
outlier that is not connected to the postcristid, but lies posterior to
it. In the submyotodont condition, the postcristid extends from the
hypoconid to the entoconid but there is an additional short crest
that connects the postcristid to the hypoconulid as well [30]. While
not common, some bat species exhibit variation in the construc-
tion of the posterior talonid. Some individuals have myotodont,
submyotodont and even nyctalodont molars in the same species,
and occasionally more than one condition in the same specimen
[31–32]. In P. phiomensis the hypoconulid is relatively small,
directly distal to the entoconid and has the postcristid either
extending to it (on the nyctalodont m2) or conjoining it and the
entoconid (on the submyotodont m1). There is a distinct labial
cingulid present that extends around the tooth both mesially and
distally.
The m2 of P. phiomensis (also preserved in YPM 24196, see
Fig. 3F) is similar to m1 but differs by having a wider and more
closed trigonid with a more labially placed paraconid and a
straighter, un-notched paracristid (Fig. 2A, C–D, 3C). Like other
myzopodids, the cristid obliqua turns slightly labially as it joins the
postvallid.
The m3 of P. phiomensis (also present in YPM 24195 and YPM
24197, see Fig. 3E, G) is smaller than m1-2 but is not as relatively
reduced as m3 is in extant myzopodids (Fig. 2A, C–D, 3C). The
trigonid is similar to that of m2 except that it is even more closed
and mesiodistally compressed. Like all myzopodids, the m3
paraconid is robust and as large as the protoconid. The m3
differs from the other lower molars by lacking a hypoconulid
(extant myzopodids either have a very small m3 hypoconulid
[M. schliemanni] or a more distinct one [M. aurita], see Fig. 3B, D),
having a more elongate entocristid, a narrower talonid basin with
the cristid obliqua joining the postvallid more lingually than in m1-
2, and by having the talonid squared-off distally.
Phasmatonycteris butleri sp. nov. urn:lsid:zoobank.org:act:E6B5
FFD8-5250-4C2A-B83F-CC469DD32557 (Fig. 3A).
Holotype
Cairo Geological Museum (CGM) 83761, right dentary with
m2-3, only known specimen.
Locality and Horizon
Fayum Quarry BQ-2, 23 meter level, Birket Qarun Formation,
Late Eocene, Priabonian (,37 Ma), Fayum Depression, Western
Desert, Egypt (Fig. 4).
Specific Diagnosis
Differs from P. phiomensis in having fully myotodont lower
molars, a relatively smaller m3 compared to m2, in having
shallower hypoflexids on m2-3, less steeply sloping m2-3
entocristids, and m2 with a relatively smaller and less anteriorly
oriented paraconid.
Etymology
Named for Percy Butler in recognition of his work on fossil bats
from Africa and of his 75 year publishing career.
Description and Comparisons
P. butleri shares some distinctive myzopodid characteristics
(Fig. 3A) including fully myotodont lower molars, a m2 cristid
obliqua that turns labially as it joins the postvallid, a high and
relatively short entocristid, and a more elongate m3 entocristid.
However, in some ways this species differs from P. phiomensis and
Myzopoda, especially in having less robust and less mesially angled
lower molar paraconids.
The m2 of P. butleri is similar to that of P. phiomensis in having a
relatively closed trigonid but the paraconid is smaller than in P.
phiomensis and is more closely appressed to the metaconid. It also
differs in having a relatively thick labial cingulid and a less robust
entoconid. The m3 of P. butleri is a reduced version of m2 differing
from the latter tooth mostly in having a narrower talonid.
Discussion
When first described [1], Myzopoda was regarded as a
vespertilionid, a position later supported by Dobson [33]. In
1904, Thomas proposed the family Myzopodidae for Myzopoda
[34], recognizing the distinctiveness of the single known species at
that time, M.aurita. The family was originally viewed as perhaps
rather closely related to Thyropteridae because both families
include species having adhesive pads on thumbs and ankles that
enable them to cling to smooth surfaces such as leaves [13,27,
34–35]. However, the morphological attributes of these adhesive
structures are fundamentally different between the two families,
suggesting that any similarities are a remarkable case of
convergence [2,13–15].
Phylogenetic studies of large nuclear gene data sets indicate that
Myzopodidae is most likely a basal member of the superfamily
Noctilionoidea [7–9], a clade that also includes mystacinids,
thyropterids, furipterids, noctilionids, mormoopids and phyllosto-
mids (Fig. 5). Noctilionoidea is essentially a southern radiation
[7,9], with myzopodids restricted to Africa and Madagascar,
mystacinids limited to New Zealand and Australia [36–37], and
the other families all restricted to South and Central America and
New Fossil Bats from Egypt
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the Caribbean [38]. Teeling et al. [7] indicated that the
biogeographic earliest appearances of myzopodids and mystaci-
nids (either in Laurasia or Gondwana) were equivocal (see their
figure 3) but it is now apparent that both of these families first
appeared in eastern Gondwana with myzopodids being present in
North Africa by the latest Eocene (,37 Ma) and mystacinids being
present in northern Australia [37] by the Oligo-Miocene boundary
(,26 Ma).
Teeling et al. [7] estimated divergence times for various bat
clades using a molecular clock calibrated with several fossils. Based
on their analysis, myzopodids apparently diverged from other
members of Noctilionoidea between 46 and 57 million years ago.
This suggests that the presence of Phasmatonycteris in North Africa
at 37 million years ago is not necessarily a surprising occurrence.
Given that other Malagasy bats (Table 1) and endemic mammals
likely can trace their origin to the African mainland in the early to
late Cenozoic [17,39] a similar expectation should exist for
myzopodids as well. In fact, the presence of a 37 million year old
basal noctilionoid in North Africa suggests that the origins of
Noctilionoidea (Fig. 5) may well be found in eastern Gondwana
with a subsequent dispersal south into Australia (mystacinids) and
then westward on to South America across Antarctica (lineage
leading to the five Neotropical noctilionoid families).
Evidence now available suggests that Antarctica remained
connected to Australia and South America until near the end of
the Eocene (35 Ma) and that ice-free corridors may have existed
well into the Oligocene [40–45]. If true, it is probable that
terrestrial dispersers would have been able to reach South America
from eastern Gondwana before the onset of the development of
the Antarctic Circumpolar Current that mark the final separation
of the three major southern land-masses [41–43]. Based on the
presence of other Fayum and Australian bat taxa, it has been
suggested that the southern continents played a crucial role in the
origin and diversification of Chiroptera [46–49]. The new
myzopodids from Egypt lend even more support to this hypothesis.
Acknowledgments
We thank the Egyptian Mineral Resources Authority and the Egyptian
Geological Museum for their continued support to assure successful
fieldwork in the Fayum Depression. We thank Elwyn L. Simons and
Prithijit Chatrath for organizing and leading the many fossil expeditions to
Egypt upon which this work is based. Walter Joyce provided access to fossil
specimens at the Yale Peabody Museum (New Haven). We thank
Lawrence Heaney, Bruce Patterson and William Stanley at the Field
Museum of Natural History (Chicago) for access to and loan of extant
myzopodid specimens. Eileen Westwig and Aja Marcato at the American
Museum of Natural History (New York) facilitated access to comparative
bat material in the Division of Vertebrate Zoology (Department of
Mammalogy) at the AMNH. Reviews by Thierry Smith and Suzanne J.
Hand were very enlightening and helpful. This is Duke Lemur Center
publication 1258.
Author Contributions
Analyzed the data: GFG NBS ERS. Contributed reagents/materials/
analysis tools: GFG NBS ERS. Wrote the paper: GFG NBS ERS.
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