Craniodental morphology and systematics of a new family of hystricognathous rodents (Gaudeamuridae) from the late eocene and early oligocene of Egypt.
ABSTRACT: BACKGROUND: Gaudeamus is an enigmatic hystricognathous rodent that was, until recently, known solely from fragmentary material from early Oligocene sites in Egypt, Oman, and Libya. Gaudeamus' molars are similar to those of the extant cane rat Thryonomys, and multiple authorities have aligned Gaudeamus with Thryonomys to the exclusion of other living and extinct African hystricognaths; recent phylogenetic analyses have, however, also suggested affinities with South American caviomorphs or Old World porcupines (Hystricidae). METHODOLOGY/PRINCIPAL FINDINGS: Here we describe the oldest known remains of Gaudeamus, including largely complete but crushed crania and complete upper and lower dentitions. Unlike younger Gaudeamus species, the primitive species described here have relatively complex occlusal patterns, and retain a number of plesiomorphic features. Unconstrained parsimony analysis nests Gaudeamus and Hystrix within the South American caviomorph radiation, implying what we consider to be an implausible back-dispersal across the Atlantic Ocean to account for Gaudeamus' presence in the late Eocene of Africa. An analysis that was constrained to recover the biogeographically more plausible hypothesis of caviomorph monophyly does not place Gaudeamus as a stem caviomorph, but rather as a sister taxon of hystricids. CONCLUSIONS/SIGNIFICANCE: We place Gaudeamus species in a new family, Gaudeamuridae, and consider it likely that the group originated, diversified, and then went extinct over a geologically brief period of time during the latest Eocene and early Oligocene in Afro-Arabia. Gaudeamurids are the only known crown hystricognaths from Afro-Arabia that are likely to be aligned with non-phiomorph members of that clade, and as such provide additional support for an Afro-Arabian origin of advanced stem and basal crown members of Hystricognathi.
Project description:The early evolutionary and paleobiogeographic history of the diverse rodent clade Hystricognathi, which contains Hystricidae (Old World porcupines), Caviomorpha (the endemic South American rodents), and African Phiomorpha (cane rats, dassie rats, and blesmols) is of great interest to students of mammalian evolution, but remains poorly understood because of a poor early fossil record. Here we describe the oldest well-dated hystricognathous rodents from an earliest late Eocene (approximately 37 Ma) fossil locality in the Fayum Depression of northern Egypt. These taxa exhibit a combination of primitive and derived features, the former shared with Asian "baluchimyine" rodents, and the latter shared with Oligocene phiomorphs and caviomorphs. Phylogenetic analysis incorporating morphological, temporal, geographic, and molecular information places the new taxa as successive sister groups of crown Hystricognathi, and supports an Asian origin for stem Hystricognathi and an Afro-Arabian origin for crown Hystricognathi, stem Hystricidae, and stem Caviomorpha. Molecular dating of early divergences within Hystricognathi, using a Bayesian "relaxed clock" approach and multiple fossil calibrations, suggests that the split between Hystricidae and the phiomorph-caviomorph clade occurred approximately 39 Ma, and that phiomorphs and caviomorphs diverged approximately 36 Ma. These results are remarkably congruent with our phylogenetic results and the fossil record of hystricognathous rodent evolution in Afro-Arabia and South America.
Project description:The Fayum Depression of Egypt has yielded fossils of hystricognathous rodents from multiple Eocene and Oligocene horizons that range in age from ?37 to ?30 Ma and document several phases in the early evolution of crown Hystricognathi and one of its major subclades, Phiomorpha. Here we describe two new genera and species of basal phiomorphs, Birkamys korai and Mubhammys vadumensis, based on rostra and maxillary and mandibular remains from the terminal Eocene (?34 Ma) Fayum Locality 41 (L-41). Birkamys is the smallest known Paleogene hystricognath, has very simple molars, and, like derived Oligocene-to-Recent phiomorphs (but unlike contemporaneous and older taxa) apparently retained dP(4)?4 late into life, with no evidence for P(4)?4 eruption or formation. Mubhammys is very similar in dental morphology to Birkamys, and also shows no evidence for P(4)?4 formation or eruption, but is considerably larger. Though parsimony analysis with all characters equally weighted places Birkamys and Mubhammys as sister taxa of extant Thryonomys to the exclusion of much younger relatives of that genus, all other methods (standard Bayesian inference, Bayesian "tip-dating," and parsimony analysis with scaled transitions between "fixed" and polymorphic states) place these species in more basal positions within Hystricognathi, as sister taxa of Oligocene-to-Recent phiomorphs. We also employ tip-dating as a means for estimating the ages of early hystricognath-bearing localities, many of which are not well-constrained by geological, geochronological, or biostratigraphic evidence. By simultaneously taking into account phylogeny, evolutionary rates, and uniform priors that appropriately encompass the range of possible ages for fossil localities, dating of tips in this Bayesian framework allows paleontologists to move beyond vague and assumption-laden "stage of evolution" arguments in biochronology to provide relatively rigorous age assessments of poorly-constrained faunas. This approach should become increasingly robust as estimates are combined from multiple independent analyses of distantly related clades, and is broadly applicable across the tree of life; as such it is deserving of paleontologists' close attention. Notably, in the example provided here, hystricognathous rodents from Libya and Namibia that are controversially considered to be of middle Eocene age are instead estimated to be of late Eocene and late Oligocene age, respectively. Finally, we reconstruct the evolution of first lower molar size among Paleogene African hystricognaths using a Bayesian approach; the results of this analysis reconstruct a rapid latest Eocene dwarfing event along the lineage leading to Birkamys.
Project description:The abrupt appearance of primates and hystricognath rodents in early Oligocene deposits of South America has puzzled mastozoologists for decades. Based on the geoclimatic changes that occurred during the Eocene/Oligocene transition period that may have favoured their dispersal, researchers have proposed the hypothesis that these groups arrived in synchrony. Nevertheless, the hypothesis of synchronous origins of platyrrhine and caviomorph in South America has not been explicitly evaluated. Our aim in this work was to apply a formal test for synchronous divergence times to the Platyrrhini and Caviomorpha splits. We have examined a previous work on platyrrhine and hystricognath origins, applied the test to a case where synchrony is known to occur and conducted simulations to show that it is possible to formally test the age of synchronous nodes. We show that the absolute ages of Platyrrhini/Catarrhini and Caviomorpha/Phiomorpha splits depend on data partitioning and that the test applied consistently detected synchronous events when they were known to have happened. The hypothesis that the arrival of primates and hystricognaths to the New World consisted of a unique event cannot be rejected.
Project description:The Jebel Qatrani Formation of northern Egypt has produced Afro-Arabia's primary record of Paleogene mammalian evolution, including the world's most complete remains of early anthropoid primates. Recent studies of Fayum mammals have assumed that the Jebel Qatrani Formation contains a significant Eocene component ( approximately 150 of 340 m), and that most taxa from that succession are between 35.4 and 33.3 million years old (Ma), i.e., latest Eocene to earliest Oligocene in age. Reanalysis of the chronological evidence shared by later Paleogene strata exposed in Egypt and Oman (Taqah and Thaytiniti areas, Dhofar Province) reveals that this hypothesis is no longer tenable. Revised correlation of the Fayum and Dhofar magnetostratigraphies indicates that (i) only the lowest 48 m of the Jebel Qatrani Formation are likely to be Eocene in age; (ii) the youngest Fayum anthropoids, including well known species such as Aegyptopithecus zeuxis and Apidium phiomense, are probably between 30.2 and 29.5 Ma, approximately 3-4 Ma younger than previously thought; (iii) oligopithecid anthropoids did not go extinct at the Eocene-Oligocene boundary but rather persisted for at least another 2.5 Ma; (iv) propliopithecid anthropoids first appear in the Fayum area at approximately 31.5 Ma, long after the Eocene-Oligocene boundary; and (v) the youngest Fayum mammals may be only approximately 1 Ma older than the 28- to 27-Ma mammals from Chilga, Ethiopia, and not 4-5 Ma older, as previously thought. Whatever gap exists in the Oligocene record of Afro-Arabian mammal evolution is now limited primarily to a poorly sampled 27- to 23-Ma window in the latest Oligocene.
Project description:Throughout the Paleogene, most terrestrial carnivore niches in Afro-Arabia were occupied by Hyaenodonta, an extinct lineage of placental mammals. By the end of the Miocene, terrestrial carnivore niches had shifted to members of Carnivora, a clade with Eurasian origins. The transition from a hyaenodont-carnivore fauna to a carnivoran-carnivore fauna coincides with other ecological changes in Afro-Arabia as tectonic conditions in the African Rift System altered climatic conditions and facilitated faunal exchange with Eurasia. Fossil bearing deposits in the Nsungwe Formation in southwestern Tanzania are precisely dated to ~25.2 Ma (late Oligocene), preserving a late Paleogene Afro-Arabian fauna on the brink of environmental transition, including the earliest fossil evidence of the split between Old World monkeys and apes. Here we describe a new hyaenodont from the Nsungwe Formation, Pakakali rukwaensis gen. et sp. nov., a bobcat-sized taxon known from a portion of the maxilla that preserves a deciduous third premolar and alveoli of dP4 and M1. The crown of dP3 bears an elongate parastyle and metastyle and a small, blade-like metacone. Based on alveolar morphology, the two more distal teeth successively increased in size and had relatively large protocones. Using a hyaenodont character-taxon matrix that includes deciduous dental characters, Bayesian phylogenetic methods resolve Pakakali within the clade Hyainailouroidea. A Bayesian biogeographic analysis of phylogenetic results resolve the Pakakali clade as Afro-Arabian in origin, demonstrating that this small carnivorous mammal was part of an endemic Afro-Arabian lineage that persisted into the Miocene. Notably, Pakakali is in the size range of carnivoran forms that arrived and began to diversify in the region by the early Miocene. The description of Pakakali is important for exploring hyaenodont ontogeny and potential influences of Afro-Arabian tectonic events upon mammalian evolution, providing a deep time perspective on the stability of terrestrial carnivore niches through time.
Project description:BACKGROUND: The hystricognath rodents of the New World, the Caviomorpha, are a diverse lineage with a long evolutionary history, and their representation in South American fossil record begins with their occurrence in Eocene deposits from Peru. Debates regarding the origin and diversification of this group represent longstanding issues in mammalian evolution because early hystricognaths, as well as Platyrrhini primates, appeared when South American was an isolated landmass, which raised the possibility of a synchronous arrival of these mammalian groups. Thus, an immediate biogeographic problem is posed by the study of caviomorph origins. This problem has motivated the analysis of hystricognath evolution with molecular dating techniques that relied essentially on nuclear data. However, questions remain about the phylogeny and chronology of the major caviomorph lineages. To enhance the understanding of the evolution of the Hystricognathi in the New World, we sequenced new mitochondrial genomes of caviomorphs and performed a combined analysis with nuclear genes. RESULTS: Our analysis supports the existence of two major caviomorph lineages: the (Chinchilloidea?+?Octodontoidea) and the (Cavioidea?+?Erethizontoidea), which diverged in the late Eocene. The Caviomorpha/phiomorph divergence also occurred at approximately 43 Ma. We inferred that all family-level divergences of New World hystricognaths occurred in the early Miocene. CONCLUSION: The molecular estimates presented in this study, inferred from the combined analysis of mitochondrial genomes and nuclear data, are in complete agreement with the recently proposed paleontological scenario of Caviomorpha evolution. A comparison with recent studies on New World primate diversification indicate that although the hypothesis that both lineages arrived synchronously in the Neotropics cannot be discarded, the times elapsed since the most recent common ancestor of the extant representatives of both groups are different.
Project description:Hyaenodonta is a diverse, extinct group of carnivorous mammals that included weasel- to rhinoceros-sized species. The oldest-known hyaenodont fossils are from the middle Paleocene of North Africa and the antiquity of the group in Afro-Arabia led to the hypothesis that it originated there and dispersed to Asia, Europe, and North America. Here we describe two new hyaenodont species based on the oldest hyaenodont cranial specimens known from Afro-Arabia. The material was collected from the latest Eocene Locality 41 (L-41, ?34 Ma) in the Fayum Depression, Egypt. Akhnatenavus nefertiticyon sp. nov. has specialized, hypercarnivorous molars and an elongate cranial vault. In A. nefertiticyon the tallest, piercing cusp on M1-M2 is the paracone. Brychotherium ephalmos gen. et sp. nov. has more generalized molars that retain the metacone and complex talonids. In B. ephalmos the tallest, piercing cusp on M1-M2 is the metacone. We incorporate this new material into a series of phylogenetic analyses using a character-taxon matrix that includes novel dental, cranial, and postcranial characters, and samples extensively from the global record of the group. The phylogenetic analysis includes the first application of Bayesian methods to hyaenodont relationships. B. ephalmos is consistently placed within Teratodontinae, an Afro-Arabian clade with several generalist and hypercarnivorous forms, and Akhnatenavus is consistently recovered in Hyainailourinae as part of an Afro-Arabian radiation. The phylogenetic results suggest that hypercarnivory evolved independently three times within Hyaenodonta: in Teratodontinae, in Hyainailourinae, and in Hyaenodontinae. Teratodontines are consistently placed in a close relationship with Hyainailouridae (Hyainailourinae + Apterodontinae) to the exclusion of "proviverrines," hyaenodontines, and several North American clades, and we propose that the superfamily Hyainailouroidea be used to describe this relationship. Using the topologies recovered from each phylogenetic method, we reconstructed the biogeographic history of Hyaenodonta using parsimony optimization (PO), likelihood optimization (LO), and Bayesian Binary Markov chain Monte Carlo (MCMC) to examine support for the Afro-Arabian origin of Hyaenodonta. Across all analyses, we found that Hyaenodonta most likely originated in Europe, rather than Afro-Arabia. The clade is estimated by tip-dating analysis to have undergone a rapid radiation in the Late Cretaceous and Paleocene; a radiation currently not documented by fossil evidence. During the Paleocene, lineages are reconstructed as dispersing to Asia, Afro-Arabia, and North America. The place of origin of Hyainailouroidea is likely Afro-Arabia according to the Bayesian topologies but it is ambiguous using parsimony. All topologies support the constituent clades-Hyainailourinae, Apterodontinae, and Teratodontinae-as Afro-Arabian and tip-dating estimates that each clade is established in Afro-Arabia by the middle Eocene.
Project description:The late Eocene prosimian Wadilemur elegans from the Jebel Qatrani Formation, northern Egypt, was originally interpreted as an anchomomyin adapiform primate based on limited information from the lower molars and distal premolars. Recently recovered fossils attributable to this species, including a proximal femur, the fourth upper premolar and first and second upper molars, and a mandible preserving the lower second premolar and lower canine and incisor alveoli, reveal a number of derived morphological similarities shared with crown strepsirrhines and, in particular, Miocene-to-Recent stem and crown galagids, to the exclusion of known adapiforms. Phylogenetic analysis of 359 morphological features scored across 95 living and extinct crown primate taxa supports a stem galagid placement for Wadilemur and older Saharagalago, and a close relationship between crown strepsirrhines and the Eocene African taxa "Anchomomys" milleri, Djebelemur, and Plesiopithecus (none of which appear to be closely related to European anchomomyins). This scheme of relationships supports the hypothesis that crown Strepsirrhini is of Afro-Arabian origin and that lemuriforms likely colonized Madagascar by crossing the Mozambique Channel. Wadilemur's known dental and postcranial morphology provides additional support for the hypothesis that galagids and lorisids had diverged by the close of the middle Eocene, and, by bolstering the approximately 37 million-year-old calibration point for crown lorisiform origins provided by Saharagalago, indirect support for the hypothesis of an ancient origin of crown Strepsirrhini and crown Primates.
Project description:Reconstructing the origin and early evolutionary history of anthropoid primates (monkeys, apes, and humans) is a current focus of paleoprimatology. Although earlier hypotheses frequently supported an African origin for anthropoids, recent discoveries of older and phylogenetically more basal fossils in China and Myanmar indicate that the group originated in Asia. Given the Oligocene-Recent history of African anthropoids, the colonization of Africa by early anthropoids hailing from Asia was a decisive event in primate evolution. However, the fossil record has so far failed to constrain the nature and timing of this pivotal event. Here we describe a fossil primate from the late middle Eocene Pondaung Formation of Myanmar, Afrasia djijidae gen. et sp. nov., that is remarkably similar to, yet dentally more primitive than, the roughly contemporaneous North African anthropoid Afrotarsius. Phylogenetic analysis suggests that Afrasia and Afrotarsius are sister taxa within a basal anthropoid clade designated as the infraorder Eosimiiformes. Current knowledge of eosimiiform relationships and their distribution through space and time suggests that members of this clade dispersed from Asia to Africa sometime during the middle Eocene, shortly before their first appearance in the African fossil record. Crown anthropoids and their nearest fossil relatives do not appear to be specially related to Afrotarsius, suggesting one or more additional episodes of dispersal from Asia to Africa. Hystricognathous rodents, anthracotheres, and possibly other Asian mammal groups seem to have colonized Africa at roughly the same time or shortly after anthropoids gained their first toehold there.
Project description:The Oligocene Epoch was a time of major radiation of the Odontoceti (echolocating toothed whales, dolphins). Fossils reveal many odontocete lineages and considerable structural diversity, but whether the clades include some crown taxa or only archaic groups is contentious. The New Zealand fossil dolphin "Prosqualodon" marplesi (latest Oligocene, ?23.9 Ma) is here identified as a crown odontocete that represents a new genus, Otekaikea, and adds to the generic diversity of Oligocene odontocetes. Otekaikea marplesi is known only from the holotype, which comprises a partial skeleton from the marine Otekaike Limestone of the Waitaki Valley. Otekaikea marplesi was about 2.5 m long; it had procumbent anterior teeth, and a broad dished face for the nasofacial muscles implicated in production of echolocation sounds. The prominent condyles and unfused cervical vertebrae suggest a flexible neck. A phylogenetic analysis based on morphological features places Otekaikea marplesi in the extinct group Waipatiidae, within the clade Platanistoidea. The phylogeny implies an Oligocene origin for the lineage now represented by the endangered Ganges River dolphin (Platanista gangetica), supporting an Oligocene history for the crown Odontoceti.