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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 I1M3 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
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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 Ple...
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Context 1
... 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. ...
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The fossil history of molossids in the North American Tertiary is among the poorest for any family of bats. The oldest definite record is of Wallia scalopidens of middle Eocene (Uintan) age from Saskatchewan, Canada. One of the youngest records is of Eumops cf. E. perotis from the late Pliocene (late early Blancan) of Arizona, USA. New occurrences...
The partially submerged Impossível-Ioiô cave system located in the karst region of Chapada Diamantina in Bahia (Brazil) has recently been the target of extensive paleontological studies. Here we provide the first report of fossil bats from this cave system, in which we recognize six species based on humeral remains: Furipterus horrens, Chrotopterus...
A new genus of vespertilionid bat from Early Miocene of Jebel Zelten, Libya, with comments on Scotophilus and early history of vespertilionid bats (Chiroptera) Nový rod netopýra z basálního miocenu Džebelu Zelten, Libye, s poznámkami k rodu Scotophilus a rané historii vespertilionidních netopýrů (Chiroptera) Abstract. A well preserved mandible of a...
Citations
... 117 Witwatia schlosseri, the largest Eocene fossil bat, was originally described from BQ-2 118 but in Dur At-Talah, it is represented by a larger-sized species. 119 According to Gunnell et al. 120 the molar size of this taxon has decreased over time, suggesting an older age for the Libyan locality. ...
Although the evolutionary history of anthropoid primates (monkeys, apes, and humans) appears relatively well‐documented, there is limited data available regarding their origins and early evolution. We review and discuss here the earliest records of anthropoid primates from Asia, Africa, and South America. New fossils provide strong support for the Asian origin of anthropoid primates. However, the earliest recorded anthropoids from Africa and South America are still subject to debate, and the early evolution and dispersal of platyrhines to South America remain unclear. Because of the rarity and incomplete nature of many stem anthropoid taxa, establishing the phylogenetic relationships among the earliest anthropoids remains challenging. Nonetheless, by examining evidence from anthropoids and other mammalian groups, we demonstrate that several dispersal events occurred between South Asia and Afro‐Arabia during the middle Eocene to the early Oligocene. It is possible that a microplate situated in the middle of the Neotethys Ocean significantly reduced the distance of overseas dispersal.
... A similar biogeographic origin has also been proposed for South American land tortoises (Kehlmaier et al. 2017). An African origin by overwater dispersal during this same time period (Eocene/Oligocene) has also been postulated for several Neotropical bat groups, including Emballonuridae, Molossidae, and Noctilionoidea, the latter including the Phyllostomidae (Teeling et al. 2005;Lim 2007Lim , 2009Gunnell et al. 2014). The occurrence of the Emballonuridae, Molossidae, and three families of noctilionoids, Phyllostomidae, Mormoopidae, and the extinct Speonycteridae in the Oligocene and/or early Miocene of North America, including some faunas that are older than the earliest records of these families in South America, suggests a more thorough analysis of the fossil record and phylogenetic relationships is necessary to determine the ancestry of the South American members of these groups (Czaplewski et al. 2003b;Czaplewski and Morgan 2012;Czaplewski 2012, 2023;Morgan et al. 2019). ...
... There is also a Miocene mystacinid from New Zealand, and late Oligocene and Miocene mystacinids are known from Australia (Hand et al. 1998(Hand et al. , 2005(Hand et al. , 2015b. A seventh family, Myzopodidae, currently endemic to Madagascar, with late Eocene, early Oligocene, and early Pleistocene species from Africa (Gunnell et al. 2014(Gunnell et al. , 2015, also has been regarded as a basal member of the Noctilionoidea (Teeling et al. 2005(Teeling et al. , 2012Gunnell et al. 2014). We follow a phylogenetic analysis by Meredith et al. (2011) that placed the Myzopodidae as a basal branch of the Vespertilionoidea. ...
... There is also a Miocene mystacinid from New Zealand, and late Oligocene and Miocene mystacinids are known from Australia (Hand et al. 1998(Hand et al. , 2005(Hand et al. , 2015b. A seventh family, Myzopodidae, currently endemic to Madagascar, with late Eocene, early Oligocene, and early Pleistocene species from Africa (Gunnell et al. 2014(Gunnell et al. , 2015, also has been regarded as a basal member of the Noctilionoidea (Teeling et al. 2005(Teeling et al. , 2012Gunnell et al. 2014). We follow a phylogenetic analysis by Meredith et al. (2011) that placed the Myzopodidae as a basal branch of the Vespertilionoidea. ...
Fossils of an insectivorous bat from the early Miocene of Panama are described as a new genus and species, Americanycteris cyrtodon (Chiroptera: Phyllostomidae: Phyllostominae). Americanycteris is a large phyllostomine bat, similar in size to the living species Chrotopterus auritus. Americanycteris cyrtodon can be distinguished from other closely related species by a posteriorly curved p4 and a thick labial cingulum on m1. Americanycteris cyrtodon occurs in two early Miocene vertebrate faunas from Panama. The holotype mandible with p4–m1 and an isolated p3 of A. cyrtodon were recovered from the early Hemingfordian (19–18 Ma) Centenario Fauna, and a mandible with p2 was found in the older late Arikareean (21 Ma) Lirio Norte Local Fauna. A similar large phyllostomine bat is known from the early Miocene Gran Barranca Fauna in Argentina. The presence of early Miocene phyllostomids in both North America and South America confirms the overwater dispersal of bats between the Americas before the late Miocene onset of the Great American Biotic Interchange. Pre-late Miocene chiropteran dispersals between the Americas were previously documented for the Emballonuridae and Molossidae. Although the five endemic New World families in the Noctilionoidea, including Phyllostomidae, were previously thought to be South American in origin, the oldest fossil records of noctilionoids (Mormoopidae and extinct Speonycteridae) are from the early Oligocene of Florida and one of the earliest records of the Phyllostomidae is from the early Miocene of Panama. The currently available fossil records from Panama and Florida suggest a possible North American origin for the Noctilionoidea.
... They recognised two previously described species and also founded four new genera, Witwatia, Qarunycteris, Saharaderma, and Khonsunycteris, and six new species of bats, Witwatia schlosseri, Witwatia eremicus, Dhofarella sigei, Qarunycteris moerisae, Saharaderma pseudovampyrus, and Khonsunycteris aegypticus (Gunnell et al., 2008). Gunnell et al. (2014) erected a new genus of myzopodid bat, Phasmatonycteris, to which he referred two new species Phasmatonycteris phiomensis, from the Early Oligocene site Quarry I of the Gebel Qatrani Formation, and Phasmatonycteris butleri, from the Late Eocene site BQ-2 of the Birket Qarun Formation. Simmons et al. (2016) described the new genus and species Aegyptonycteris knighteae, based on a partial right maxilla from the Late Eocene site BQ-2. ...
The Fayum has yielded one of the oldest and richest records of fossil mammals from Africa. Today, the Fayum Depression represents an oasis in the Western Desert of Egypt, south of Cairo, and contains several localities that are world renowned for their diverse Eocene to Oligocene vertebrate assemblages. The fossil sites of the Fayum area have provided numerous extraordinarily well-preserved vertebrate remains including complete skulls and partial skeletons of turtles, crocodiles, birds, and mammals. Thus, the Fayum Depression has shed light onto the evolution and biogeography of vertebrates during the Paleogene leading to the establishment of two new orders, several new families and subfamilies, and numerous new species of mammals. In recent years, these fossils have provided clues about ecological aspects of some groups using new methods such as stable isotope analysis and µCT scanning. Despite the fact that the Fayum Depression represents a historical excavation site that has been excavated and studied by numerous famous palaeontologists since the nineteenth century, the area continues to provide new insights into the evolution of mammals during the Eocene to Oligocene, greatly improving our understanding of early Cenozoic vertebrate evolution. This chapter presents a historical review of the excavations in the Fayum Depression and a taxonomic overview of its fossil fauna. Special focus was placed on the taxonomy and, where applicable, ecology of terrestrial mammals from the historical Paleogene Fayum localities.KeywordsEgyptFayum DepressionCenozoicPaleogeneMammaliaVertebrate assemblagesFossil fauna
... ; Palaeochiropterygidae: Anatolianycteris in sularis(Jones et al. 2019), Palaeochiropteryx tupaiodon, P. spiegeli, Cecilionycteris prisca, and Stehlinia spp.; Philisidae: Philisis sphingis); family incertae sedis: Necromantis adichaster; Myzopodidae: Phasmatonycteris butleri and P. phiomensis(Gunnell et al. 2014: fig. 3); Mystacinidae: Vulcanops jenny worthyae; Mormoopidae: Koopmanycteris palaeomormoops; Speonycteridae: Speonycteris auran tiadens; Phyllostomidae: Notonycteris magdalenensis and N. sucharadeus; Natalidae: Primonatalus prattae; Molossi dae: Wallia scalopidens (this species was considered a molossid by ...
We surveyed molar surface morphology of bats of 281 extant and extinct species in 5 archaic and 19 extant families
using scanning microscopy. We note the occurrence of structural features on talonid crests, the cristid obliqua, postcris�tid, and entocristid, and their absence in upper molars, even of the same species having them on lowers. We term the
structures “carnassiform notches” (CN) for their resemblance to similar features on the carnassial teeth of carnivorans.
A CN consists of a small cleft in the edge of a talonid shearing crest accompanied by an adjacent “accessory trough”
on the basinward side of the notch. The CN occur in bats with tribosphenic molar morphology and insectivorous or
insectivorous–omnivorous dietary habits. Of 19 extant families examined, eight include members that possess lower
molars with a CN in at least the cristid obliqua: Megadermatidae, Nycteridae, Mystacinidae, Furipteridae, Thyropteridae,
Phyllostomidae, Natalidae, and Vespertilionidae (Murininae and Kerivoulinae only). An extinct genus of Hipposideridae,
Vaylatsia, shows CN although extant hipposiderids do not. In extinct families for which lower molar fossils are available,
notches were not recognized on the talonids, indicating the condition is not plesiomorphic for bats and probably evolved
convergently in different lineages. Where present, the CN or troughs are morphologically consistent within a family,
and might serve in some cases as characters supporting phylogenetic analyses and clade diagnoses. CN and accessory
troughs probably increase the functional efficiency at sectioning chitin by increasing the effective length of a crest while
maintaining the same cusp-to-cusp distance and precise occlusal relationships, and by improving the food-capture area
of the shearing blade during occlusion. The accessory troughs provide an immediately adjacent fragment-clearance area.
The increased sophistication of this food-processing system might be particularly important in species that must quickly
acquire, chew, and swallow their food and resume echolocating in flight. The common ancestor of bats probably did not
have CN in its molars, and the presence of CN does not signal carnivory in bats.
... BQ-2 has since produced the most diverse African Paleogene mammalian fauna known so far. Mammalian fossils from BQ-2 include Chiroptera (Gunnell et al., 2008(Gunnell et al., , 2014Simmons et al., 2016), Hyaenodonta (Borths and Stevens, 2017), Hyracoidea (Barrow et al., 2010), Macroscelidea (Seiffert et al., 2003), Proboscidea (Liu et al., 2008), Primates (Seiffert et al., 2003(Seiffert et al., , 2005(Seiffert et al., , 2017, and Rodentia (Sallam et al., 2009(Sallam et al., , 2010a(Sallam et al., , 2010b. Several non-mammalian vertebrates have also been unearthed at BQ-2, including teleost fishes (Murray et al., 2010), crocodilians (Stefanic et al., 2020), lizards (Holmes et al., 2010a), and snakes (McCartney and Seiffert, 2016). ...
The Eocene was a key interval for the diversification of squamates worldwide. However, little is known about their early evolution in Africa due to the scarcity of Paleogene fossil squamates from that continent. One of the most important Eocene vertebrate assemblages in Africa is known from the Fayum Depression of Egypt, but only a few lizards and snakes have been reported from these deposits. Here we report the first Paleogene record of the clade Amphisbaenia in Egypt, from the lowest upper Eocene Birket Qarun Locality 2 (BQ-2). It corresponds to a mid-trunk vertebra characterized by a depressed centrum with a flattened base, hemispherical synapophyses, and absence of a neural spine. Additional material from BQ-2 includes seven vertebrae attributed to colubroidean snakes. One trunk vertebra resembles those of early Eocene Procerophis from India while the remaining colubroidean vertebrae are referred to Renenutet enmerwer. Caudal vertebrae of Renenutet are described here for the first time. Similarities in the caudal vertebral morphology between BQ-2 Renenutet and Thaumastophis missiaeni from the lower Eocene of India reinforce the hypothesis of a close affinity between them. The presence of cf. Procerophis and Renenutet in Egypt adds to the evidence for terrestrial faunal exchange between Asia and northern Africa during the early/middle Eocene along the southern margin of the Tethyan Sea.
... Various studies place Myzopodidae as part of basal radiation of either Vespertilionoidea (Van Den Busche & Hoofer, 2004;Eick et al., 2005), Noctilionoidea (Teeling et al., 2005;Agnarsson et al., 2011), or even Yangochiroptera in general (Hoofer & Van Den Busche, 2003), with invariably low support. Given the undoubted antiquity of Myzopodidae (Gunnell et al., 2014), the solution, for some reason never explicitly proposed before, might be to recognize them as a superfamily of their own. ...
Bats are the second largest mammalian order with an almost worldwide distribution. Bat taxonomy remained almost unchanged for decades, and the diversity of the order was underestimated. The advent of molecular methods brought change to chiropteran taxonomy. The number of families increased from 17–18 to 21, and the relationships between them were revised, as were the composition of suborders and superfamilies. The number of recognized species and genera went up by almost a third. As a discipline, bat taxonomy has changed much methodologically and conceptually. After its long reign, comparative morphology has faded into the background. It has become clear that characters can diverge and converge in related species, masking true phylogenetic relationships. Not writing morphology off entirely, it does necessitate resorting to finer structures or multivariate data analysis. Karyology is of limited use in bat taxonomy, but methods such as FISH add to the understanding of relationships between suprageneric taxa. Mitochondrial DNA sequences are easy to obtain, and their analysis yields well-supported phylogenetic trees, but reticular processes and other factors may mask taxon boundaries. To resolve the uncertainty, nuclear markers are used, and their number and choice depends on taxon characteristics. Overall, building a consistent chiropteran system calls for an integration of all mentioned approaches.
... Simmons et al. (2016) questioned the belonging of these materials to a marsupial. In the same quarry where Ghamidtherium was found, four species of Chiroptera were recognized later: Witwatia schlosseri Gunnell et al., 2008, Witwatia eremicus Gunnell et al., 2008, Qarunycteris moerisae Gunnell et al., 2008 and Phasmatonycteris butleri Gunnell et al., 2014(Gunnell et al., 2008, 2014. The materials described by Sánchez-Villagra et al. (2007) either differ from these species or are not comparable because they were recognized on the basis of lower molars. ...
... To us, the upper molars described in Sánchez-Villagra et al. (2007) have several features that characterize chiropteran species of the Vespertilionidae and Myzopodidae, as Myotis, Pipistrellus or Miostrellus, in the former, or Myzopoda in the second. These features include the shape of the protocone, absence of talon, well-developed mesial and posterior cinguli, M2 having a small hypocone, among other features (Gunnell et al., 2014;Crespo et al., 2018Crespo et al., , 2021. Notwithstanding, these materials also have unique features, as the connection of the preprotocrista with the paracone, which is disconnected to the paracingulum. ...
... Until upper molars of Khonsunycteris are found, we do not discard that the upper molars found in BQ-2 belong to the genus Khonsunycteris. More recently Gunnell et al. (2014) described from the same quarry of Ghamidtherium the myzopodid bat Phasmatonycteris phiomensis on the basis of a lower jaw. We question the taxonomic assignment of this species because it is quite smaller than the material described by Sánchez-Villagra et al. (2007); however, it shouldn't be discarded a referral to the same genus, or even family, until upper molars of Phasmatonycteris are found. ...
The record of extinct African metatherians (Mammalia, Theria) is scanty, restricted in time (Eocene–Miocene), and its taxonomy is still subject of debate. A review of all African metatherians, or alleged metatherians, known up to now, led us to the recognition of only three taxa referable to this group: (1) Kasserinotherium tunisiense (Peradectoidea?), from the early Eocene of Tunisia; (2) Peratherium africanum (Herpetotheriidae), from the early Oligocene of Egypt and Oman, and (3) an indeterminate Herpetotheriidae? from the early Miocene of Uganda. Herpetotheriids probably reached Afro-Arabia from Europe in one or more dispersal waves since the early Oligocene. Kasserinotherium, on the contrary, suggests an earlier (Paleocene) arrival from South America, judging from its alleged affinities with South American and Australian taxa. Such a migration event (probably, through a filter corridor such as the Rio Grande Rise-Walvis Ridge system in the South Atlantic) may also explain the enigmatic presence of polydolopimorphian metatherians in the Cenozoic of central Anatolia (Turkey). A more radical hypothesis is that all European (Eurasian?) Marsupialiformes have an ultimate origin in South America, from where they dispersed via Africa by the Paleocene–earliest Eocene.
... Mammals have been a particular focus of research because of their age, abundance, and diversity. Numerous species of primates (Seiffert et al., 2003(Seiffert et al., , 2005, rodents (Sallam et al., 2009(Sallam et al., , 2010a(Sallam et al., , 2010b, hyraxes (Barrow et al., 2010), bats (Gunnell et al., 2008(Gunnell et al., , 2014Simmons et al., 2016), elephants (Liu et al., 2008), and tenrecoids (Seiffert, 2010), have been documented from BQ-2 with the majority of them being unique to this locality. However, the non-mammalian fauna from BQ-2, including fishes, amphibians, and reptiles (McCartney and Seiffert, 2016;Holmes et al., 2010a), has gained very little attention; only some of the osteichthyan fishes from BQ-2 have been studied and described by Murray et al. (2010). ...
The Birket Qarun Formation, sampling upper Eocene deposits of the Fayum Depression, has been the subject of disagreement in terms of the environmental deposition. The formation has traditionally been considered to be a marine deposit but, more recently, one particular site, Birket Qarun Locality 2 (BQ-2), has been reported to have freshwater vertebrates, indicating that at least this locality in the formation contains terrestrial deposits. We here report a new varied assemblage of fossil catfishes from BQ-2, which adds new information with which to interpret the paleoenvironment of the locality. The new BQ-2 catfish assemblage includes the earliest record of the freshwater family Bagridae in Africa, as well as the oldest records of Fajumia and Socnopaea, otherwise known only from the overlying marine deposits of the Qasr el-Sagha Formation. Notable among the finds are remains of Arius fraasi and two additional species of marine ariid catfishes, representing the first such occurrences from the Fayum deposits. These occurrences indicate that BQ-2 likely had marine influences, or perhaps that Fajumia, Socnopaea, and Arius were euryhaline. In addition, the freshwater families Claroteidae and Mochokidae are represented among our collection. The new material documents the presence of a variety of taxa, and in addition it documents the variation in body size of members of the same family. Most evidence from other vertebrate fossils indicates that BQ-2 is a freshwater deposit, but some elements of the new catfish collection suggest the presence of nearshore marine influences, indicating that a much more complex environment was present.
... Thus, we focus on phyllostomid species, which feature significant differences in overall cranial length [15,31] that we could collect from the wild: Carollia perspicillata, a predominantly frugivorous bat [56,78] with a face near the center of cranial shape morphospace (Fig. 2); Artibeus jamaicensis, a predominantly frugivorous bat [41] with a short and wide face; and Glossophaga soricina, a predominantly nectarivorous and pollenivorous bat [12] with an elongated head and narrow face. These dietary specialists were compared to Miniopterus natalensis (family Miniopteridae), a representative insect-feeding outgroup species with a relatively unmodified face from South Africa, the geographic area of origin for Neotropical species in Noctilionoidea [27,44,64]. Embryos from each species were collected during stages CS16, CS17, and CS18 (approximately 50, 54, and 60 days of gestation, respectively) undergoing craniofacial elongation [13]. ...
Background:
Skull diversity in the neotropical leaf-nosed bats (Phyllostomidae) evolved through a heterochronic process called peramorphosis, with underlying causes varying by subfamily. The nectar-eating (subfamily Glossophaginae) and blood-eating (subfamily Desmondontinae) groups originate from insect-eating ancestors and generate their uniquely shaped faces and skulls by extending the ancestral ontogenetic program, appending new developmental stages and demonstrating peramorphosis by hypermorphosis. However, the fruit-eating phyllostomids (subfamilies Carollinae and Stenodermatinae) adjust their craniofacial development by speeding up certain developmental processes, displaying peramorphosis by acceleration. We hypothesized that these two forms of peramorphosis detected by our morphometric studies could be explained by differential growth and investigated cell proliferation during craniofacial morphogenesis.
Results:
We obtained cranial tissues from four wild-caught bat species representing a range of facial diversity and labeled mitotic cells using immunohistochemistry. During craniofacial development, all bats display a conserved spatiotemporal distribution of proliferative cells with distinguishable zones of elevated mitosis. These areas were identified as modules by the spatial distribution analysis. Ancestral state reconstruction of proliferation rates and patterns in the facial module between species provided support, and a degree of explanation, for the developmental mechanisms underlying the two models of peramorphosis. In the long-faced species, Glossophaga soricina, whose facial shape evolved by hypermorphosis, cell proliferation rate is maintained at lower levels and for a longer period of time compared to the outgroup species Miniopterus natalensis. In both species of studied short-faced fruit bats, Carollia perspicillata and Artibeus jamaicensis, which evolved under the acceleration model, cell proliferation rate is increased compared to the outgroup.
Conclusions:
This is the first study which links differential cellular proliferation and developmental modularity with heterochronic developmental changes, leading to the evolution of adaptive cranial diversity in an important group of mammals.
... Third, Volleth (2013) reported chromosomal evolutionary similarities suggestive of a myzopodidemballonuroid association, as did Carter et al. (2008) for placentation data. Fourth, new myzopodid fossils (two species of Phasmatonycteris) have been recovered from up to 37 my old deposits (early Oligocene) of the Fayum Depression, Egypt (Gunnell et al. 2014). Chromosomal and morphological data, and the presence of a myzopodid lineage of Paleogene age in continental Africa, materialize the possibility of an African link between myzopodids and emballonuroids beyond our tree. ...
