New insights into the ear region anatomy and cranial blood supply of advanced stem Strepsirhini: Evidence from three primate petrosals from the Eocene of Chambi,Tunisia

We report the discovery of three isolated primate petrosal fragments from the fossiliferous locality of Chambi (Tunisia), a primate-bearing locality dating from the late early to the early middle Eocene. These fossils display a suite of anatomical characteristics otherwise found only in strepsirhines, and as such might be attributed either to Djebelemur or/and cf. Algeripithecus, the two diminutive stem strepsirhine primates recorded from this locality. Although damaged, the petrosals provide substantial information regarding the ear anatomy of these advanced stem strepsirhines (or pre-tooth-combed primates), notably the patterns of the pathway of the arterial blood supply. Using μCT-scanning techniques and digital segmentation of the structures, we show that the transpromontorial and stapedial branches of the internal carotid artery (ICA) were present (presence of bony tubes), but seemingly too small to supply enough blood to the cranium alone. This suggests that the ICA was not the main cranial blood supply in stem strepsirhines, but that the pharyngeal or vertebral artery primitively ensured a great part of this role instead, an arterial pattern that is reminiscent of modern cheirogaleid, lepilemurid lemuriforms and lorisiforms. This could explain parallel loss of the ICA functionality among these families. Specific measurements made on the cochlea indicate that the small strepsirhine primate(s) from Chambi was (were) highly sensitive to high frequencies and poorly sensitive to low frequencies. Finally, variance from orthogonality of the plane of the semicircular canals (SCs) calculated on one petrosal (CBI-1-569) suggests that Djebelemur or cf. Algeripithecus likely moved (at least its head) in a way similar to that of modern mouse lemurs.     

Evolutionary and developmental dynamics of the dentition in Muroidea and Dipodoidea (Rodentia, Mammalia)

SUMMARY When it comes to mouse evo‐devo, the fourth premolar–first molar (P4–M1) dental complex becomes a source of longstanding controversies among paleontologists and biologists. Muroidea possess only molar teeth but with additional mesial cusps on their M1. Developmental studies tend to demonstrate that the formation of such mesial cusps could result from the integration of a P4 germ into M1 during odontogenesis. Conversely, most Dipodoidea conserve their fourth upper premolars and those that lost these teeth can also bear additional mesial cusps on their first upper molars. The aim of this study is to assess this developmental model in both Muroidea and Dipodoidea by documenting the morphological evolution of the P4–M1 complex across 50 Ma. Fourteen extinct and extant species, including abnormal and mutant specimens were investigated. We found that, even if their dental evolutionary pathways strongly differ, Dipodoidea and Muroidea retain common developmental characteristics because some of them can present similar dental morphological trends. It also appears that the acquisition of a mesial cusp on M1 is independent from the loss of P4 in both superfamilies. Actually, the progressive decrease of the inhibitory effect of P4, consequent to its regression, could allow the M1 to lengthen and mesial cusps to grow in Muroidea. Apart from these developmental explanations, patternings of the mesial part of first molars are also deeply constrained by morpho‐functional requirements. As there is no obvious evidence of such mechanisms in Dipodoidea given their more variable dental morphologies, further developmental investigations are needed.     

The primate community of Cachoeira (Brazilian Amazonia): a model to decipher ecological partitioning among extinct species

Dental microwear analysis is conducted on a community of platyrrhine primates from South America. This analysis focuses on the primate community of Cachoeira Porteira (Para, Brazil), in which seven sympatric species occur: Alouatta seniculus, Ateles paniscus, Cebus apella, Chiropotes satanas, Pithecia Pithecia, Saguinus midas, and Saimiri sciureus. Shearing quotients are also calculated for each taxon of this primate community. Dental microwear results indicate significant differences between taxa, but are somewhat insufficient when it comes to discriminating between ecologically similar taxa. The primates of Cachoeira Porteira all incorporate a certain amount of fruit in their diet, entailing a definite amount of inter-specific competition as they must share food resources. Alouatta is the most folivorous taxon of this community, which is corroborated by dental microwear analysis. Ateles, although of a similar size to Alouatta, limits inter-specific competition by incorporating more fruit in its diet. Cebus has a very diverse omnivorous diet, which is highlighted in this study, as it compares to both fruit and leaf eating taxa. In some cases, microwear results need to be supplemented by other methods. For example, dental microwear seems insufficient to distinguish between Pithecia and Chiropotes, which eat foods with similar physical properties. However, other methods (i.e. shearing quotients and body mass) provide enough complimentary information to be able to highlight differences between the two taxa. On the other hand, dental microwear can highlight differences between primates which have similar diets, such as Saimiri and Saguinus. In this case, differences could be due to other exogenous factors.     

A new nothrotheriid xenarthran from the early Pliocene of Pomata‐Ayte (Bolivia): new insights into the caniniform–molariform transition in sloths

Tardigrade xenarthrans are today represented only by the two tree sloth genera Bradypus and Choloepus, which inhabit the Neotropical rainforests and are characterized by their slowness and suspensory locomotion. Sloths have been recognized in South America since the early Oligocene. This monophyletic group is represented by five clades traditionally recognized as families: Bradypodidae, Megalonychidae, Mylodontidae (?), Megatheriidae (?) and Nothrotheriidae (?). A new nothrotheriid ground sloth represented by a dentary and several postcranial elements, Aymaratherium jeani gen. nov. , sp. nov. , from the early Pliocene locality of Pomata‐Ayte (Bolivia) is reported. This small‐ to medium‐sized species is characterized especially by its dentition and several postcranial features. It exhibits several convergences with the ‘aquatic’ nothrotheriid sloth Thalassocnus and the giant megatheriid ground sloth Megatherium (M.) americanum, and is interpreted as a selective feeder, with good pronation and supination movements. The tricuspid caniniform teeth of Aymaratherium may represent a transitional stage between the caniniform anterior teeth of basal megatherioids and basal nothrotheriids (1/1C‐4/3M as in Hapalops or Mionothropus) and the molariform anterior teeth of megatheriids (5/4M, e.g. Megatherium). To highlight the phylogenetic position of this new taxon among nothrotheriid sloths, we performed a cladistic assessment of the available dental and postcranial evidence. Our results, derived from a TNT treatment of a data matrix largely based on a published phylogenetic data set, indicate that Aymaratherium is either sister taxon to Mionothropus or sister to the clade Nothrotheriini within Nothrotheriinae. They further support the monophyly of both the Nothrotheriinae and the Nothrotheriini, as suggested previously by several authors.     

New strepsirrhine primate from the late Eocene of Peninsular Thailand (Krabi Basin)

In this paper, we describe the newly discovered lower jaw of a primate from the late Eocene Krabi coal mine (Bang Mark pit) of Peninsular Thailand. We performed microtomographic examinations at the European Synchrotron Radiation Facility (ESRF, Grenoble, France) to analyze different morphological aspects of the jaw and teeth. Although partially preserved, this fossil mandible reveals a set of distinctive dental traits (e.g., double-rooted P(2) and molarized P(4)) that allow us to describe a new stepsirrhine adapiform: Muangthanhinius siami, new genus and species. This taxon is somewhat atypical among Paleogene adapiforms, and more specialized than the sivaladapid adapiforms (hoanghoniines) that existed in Asia in the same epoch. In fact, Muangthanhinius shows a degree of dental specialization approximating that of some modern strepsirrhine lemuriforms, although it lacks the highly specialized anterior dentition characterizing this living primate group (canine + incisors forming a very procumbent toothcomb). In contrast, Muangthanhinius exhibits a large canine deeply anchored within the dentary that probably protruded high above the toothrow. Finally, despite the development of a molarized P(4) as in Miocene sivaladapid sivaladapines, Muangthanhinius differs in molar morphology from this group, and the position of this new taxon within the Adapiformes remains indeterminate. Clarification of its phylogenetic position will require more morphological evidence than is currently available.     

Cynocephalid dermopterans from the Palaeogene of South Asia (Thailand, Myanmar and Pakistan): systematic, evolutionary and palaeobiogeographic implications

Cynocephalid dermopterans (flying lemurs) are represented by only two living genera (Cynocephalus and Galeopterus), which inhabit tropical rainforests of South‐East Asia. Despite their very poor diversity and their limited distribution, dermopterans play a critical role in higher‐level eutherian phylogeny inasmuch as they represent together with Scandentia (tree‐shrew) the sister group of the Primates clade (Plesiadapiformes + Euprimates). However, unlike primates, for which the fossil record extends back to the early Palaeogene on all Holarctic continents and in Africa, the evolutionary history of the order Dermoptera sensu stricto (Cynocephalidae) has so far remained undocumented, with the exception of a badly preserved fragment of mandible from the late Eocene of Thailand (Dermotherium major). In this paper, we described newly discovered fossil dermopterans (essentially dental remains) from different regions of South Asia (Thailand, Myanmar, and Pakistan) ranging from the late middle Eocene to the late Oligocene. We performed microtomographic examinations at the European Synchrotron Radiation Facility (ESRF, Grenoble, France) to analyse different morphological aspects of the fossilized jaws. The abundant material from the late Oligocene of Thailand (Nong Ya Plong coal mine) allows us to emend the diagnosis of the genus Dermotherium and to describe a new species: Dermotherium chimaera sp. n. This species exhibits an interesting mosaic of plesiomorphic cynocephalid characters shared with Cynocephalus and Galeopterus, and as such, it probably documents a form close to the ancestral morphotype from which the two extant forms are derived (supported by cladistic assessment of the dental evidence). The discovery of Palaeogene cynocephalids is particularly significant since it attests to the great antiquity of the order Dermoptera in Asia, and besides, it provides the first spatio‐temporal glimpse into the evolutionary history of that enigmatic mammal group. In that respect, these fossils testify to a long history of endemism in South Asia for dermopterans, and demonstrate that their modern geographic restriction in south‐eastern Asia is clearly a relictual distribution. Cynocephalids had a more widespread distribution during the Palaeogene, which extended from the Indian subcontinent (the rafting Greater India) to South‐East Asia. Their subsequent extinction on the Indian subcontinent was probably mediated by the major palaeogeographic and geomorphologic events related to the India‐Eurasia collision (retreat of the Paratethys Sea, formation of orogenic highlands) that have strongly affected the climate of South Asia at the end of the Oligocene.     

A new primate from the Eocene Pondaung Formation of Myanmar and the monophyly of Burmese amphipithecids

The family Amphipithecidae is one of the two fossil primate taxa from Asia that appear to be early members of the anthropoid clade. Ganlea megacanina, gen. et sp. nov., is a new amphipithecid from the late middle Eocene Pondaung Formation of central Myanmar. The holotype of Ganlea is distinctive in having a relatively enormous lower canine showing heavy apical wear, indicating an important functional role of the lower canine in food preparation and ingestion. A phylogenetic analysis of amphipithecid relationships suggests that Ganlea is the sister taxon of Myanmarpithecus, a relatively small-bodied taxon that has often, but not always, been included in Amphipithecidae. Pondaungia is the sister taxon of the Ganlea + Myanmarpithecus clade. All three Pondaung amphipithecid genera are monophyletic with respect to Siamopithecus, which is the most basal amphipithecid currently known. The inclusion of Myanmarpithecus in Amphipithecidae diminishes the likelihood that amphipithecids are specially related to adapiform primates. Extremely heavy apical wear has been documented on the lower canines of all three genera of Burmese amphipithecids. This distinctive wear pattern suggests that Burmese amphipithecids were an endemic radiation of hard object feeders that may have been ecological analogues of living New World pitheciin monkeys.     

Anthropoid versus strepsirhine status of the African Eocene primates Algeripithecus and Azibius: craniodental evidence

Recent fossil discoveries have demonstrated that Africa and Asia were epicentres for the origin and/or early diversification of the major living primate lineages, including both anthropoids (monkeys, apes and humans) and crown strepsirhine primates (lemurs, lorises and galagos). Competing hypotheses favouring either an African or Asian origin for anthropoids rank among the most hotly contested issues in paleoprimatology. The Afrocentric model for anthropoid origins rests heavily on the >45 Myr old fossil Algeripithecus minutus from Algeria, which is widely acknowledged to be one of the oldest known anthropoids. However, the phylogenetic position of Algeripithecus with respect to other primates has been tenuous because of the highly fragmentary fossils that have documented this primate until now. Recently recovered and more nearly complete fossils of Algeripithecus and contemporaneous relatives reveal that they are not anthropoids. New data support the idea that Algeripithecus and its sister genus Azibius are the earliest offshoots of an Afro–Arabian strepsirhine clade that embraces extant toothcombed primates and their fossil relatives. Azibius exhibits anatomical evidence for nocturnality. Algeripithecus has a long, thin and forwardly inclined lower canine alveolus, a feature that is entirely compatible with the long and procumbent lower canine included in the toothcomb of crown strepsirhines. These results strengthen an ancient African origin for crown strepsirhines and, in turn, strongly challenge the role of Africa as the ancestral homeland for anthropoids.     

Early Tertiary mammals from North Africa reinforce the molecular Afrotheria clade

The phylogenetic pattern and timing of the radiation of mammals, especially the geographical origins of major crown clades, are areas of controversy among molecular biologists, morphologists and palaeontologists. Molecular phylogeneticists have identified an Afrotheria clade, which includes several taxa as different as tenrecs (Tenrecidae), golden moles (Chrysochloridae), elephant-shrews (Macroscelididae), aardvarks (Tubulidentata) and paenungulates (elephants, sea cows and hyracoids). Molecular data also suggest a Cretaceous African origin for Afrotheria within Placentalia followed by a long period of endemic evolution on the Afro-Arabian continent after the mid-Cretaceous Gondwanan breakup (approx. 105-25 Myr ago). However, there was no morphological support for such a natural grouping so far. Here, we report new dental and postcranial evidence of Eocene stem hyrax and macroscelidid from North Africa that, for the first time, provides a congruent phylogenetic view with the molecular Afrotheria clade. These new fossils imply, however, substantial changes regarding the historical biogeography of afrotheres. Their long period of isolation in Africa, as assumed by molecular inferences, is now to be reconsidered inasmuch as Eocene paenungulates and elephant-shrews are here found to be related to some Early Tertiary Euramerican 'hyopsodontid condylarths' (archaic hoofed mammals). As a result, stem members of afrotherian clades are not strictly African but also include some Early Paleogene Holarctic mammals.     

Spatial and temporal ecological diversity amongst Eocene primates of France: evidence from teeth

Diet is of paramount importance in the life of a primate. It is also highly variable, as potential food sources vary in spatial distribution and availability over time. The fossil record, due to its fragmentary nature, offers few possibilities to assess the dietary range of a given primate across its spatial and temporal distribution. Here we focus on three taxa, Leptadapis magnus (Adapidae, Adapiformes), Necrolemur cf. antiquus (Microchoeridae, Omomyiformes), and Pseudoloris parvulus (Microchoeridae, Omomyiformes). These taxa occur at different localities of the Late Eocene in the south of France ranging from MP16 (Robiac, Lavergne; 39 Ma), MP17a (La Bouffie, Euzet, Fons 4; 38 Ma) to MP17b (Perrière; 37 Ma). Diets of fossil taxa are assessed here by dental microwear analysis using a comparative database of 11 species of living strepsirhines. On the whole, leaves were a preferred food for the large-bodied Leptadapis (4-5 kg). However, the diet of this taxon varied from a mix of leaves and fruit at La Bouffie, a closed tropical rain forest environment, to a strictly leaf-eating one in the more open environment of Perrière. Based on body mass (200-350 g) and dental microwear patterns, Necrolemur had a mainly fruit-based diet, perhaps supplemented by insects. However, the comparison of the different localities reveals the dietary range of this small-bodied omomyiform which seems to vary between insects and a much softer diet. Pseudoloris had a diet strictly based on insects. Contrary to Leptadapis or Necrolemur, its diet seems to have been confined to insects whatever the locality considered.