New adapiform primate fossils from the late Eocene of Egypt

Abstract

Caenopithecine adapiform primates are currently represented by two genera from the late Eocene of Egypt (Afradapis and Aframonius) and one from the middle Eocene of Switzerland (Caenopithecus). All are somewhat anthropoid-like in several aspects of their dental and gnathic morphology, and are inferred to have been highly folivorous. Here we describe a new caenopithecine genus and species, Masradapis tahai, from the ~37 million-year-old Locality BQ-2 in Egypt, that is represented by mandibular and maxillary fragments and isolated teeth. Masradapis is approximately the same size as Aframonius but differs in having a more dramatic distal increase in molar size, more complex upper molar shearing crests, and an exceptionally deep mandibular corpus. We also describe additional mandibles and part of the orbit and rostrum of Aframonius which suggest that it was probably diurnal. Phylogenetic analyses place Masradapis either as the sister taxon of Aframonius (parsimony), or as the sister taxon of Afradapis and Caenopithecus (Bayesian methods). Bayesian tip-dating analysis, when combined with Bayesian biogeographic analysis, suggests that a common ancestor of known caenopithecines dispersed to Afro-Arabia from Europe between 49.4 and 47.4 Ma, and that a trans-Tethyan back-dispersal explains Caenopithecus’ later presence in Europe.

For Masradapis: https://www.zoobank.org/urn:lsid:zoobank.org:act:41BC8459-7CCE-487F-BC59-1C34257D5C4E

For Masradapis tahai: https://www.zoobank.org/urn:lsid:zoobank.org:act:C0A620AD-6FCA-4649-A980-FCA237AFE39D     

Astragalar morphology of late Eocene anthropoids from the Fayum Depression (Egypt) and the origin of catarrhine primates

The phylogenetic relationships of the late Eocene anthropoids Catopithecus browni and Proteopithecus sylviae are currently a matter of debate, with opinion divided as to whether these taxa are stem or crown anthropoids. The phylogenetic position of Catopithecus is of particular interest, for, unlike the highly generalized genus Proteopithecus, this taxon shares apomorphic dental and postcranial features with more derived undoubted catarrhines that appear in the same region 1-2 Ma later. If these apomorphies are homologous and Catopithecus is a stem catarrhine, the unique combination of plesiomorphic and apomorphic features preserved in this anthropoid would have important implications for our understanding of the crown anthropoid morphotype and the pattern of morphological character transformations that occurred during the early phases of stem catarrhine evolution.Well-preserved astragali referrable to Proteopithecus, Catopithecus, and the undoubted early Oligocene stem catarrhine Aegyptopithecus have provided additional morphological evidence that allows us to further evaluate competing hypotheses of interrelationships among Eocene-Oligocene Afro-Arabian anthropoids. Qualitative observations and multivariate morphometric analyses reveal that the astragalar morphology of Proteopithecus is very similar to that of early Oligocene parapithecids and living and extinct small-bodied platyrrhines, and strengthens the hypothesis that the morphological pattern shared by these taxa is primitive within crown Anthropoidea. In contrast, Catopithecus departs markedly from the predicted crown anthropoid astragalar morphotype and shares a number of apomorphic features (e.g., deep cotylar fossa, laterally projecting fibular facet, trochlear asymmetry, mediolaterally wide astragalar head) with Aegyptopithecus and Miocene-Recent catarrhines. The evidence from the astragalus complements other independent data from the dentition, humerus and femur of Catopithecus that support this taxon's stem catarrhine status, and we continue to maintain that oligopithecines are stem catarrhines that constitute the sister group of a clade containing propliopithecines and Miocene-Recent catarrhines.     

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.     

New postcranial elements for the earliest Eocene fossil primate Teilhardina belgica

Teilhardina belgica is one of the most primitive fossil primates known to date and the earliest haplorhine with associated postcranials, making it relevant to a reconstruction of the ancestral primate morphotype. Here we describe newly discovered postcranial elements of T. belgica. It is a small primate with an estimated body mass between 30 and 60 g, similar to the size of a mouse lemur. Its hindlimb anatomy suggests frequent and forceful leaping with excellent foot mobility and grasping capabilities. It can now be established that this taxon exhibits critical primate postcranial synapomorphies such as a grasping hallux, a tall knee, and nailed digits. This anatomical pattern and behavioral profile is similar to what has been inferred before for other omomyids and adapiforms. The most unusual feature of T. belgica is its elongated middle phalanges (most likely manual phalanges), suggesting that this early primate had very long fingers similar to those of living tarsiers.     

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.     

Enigmatic new mammals from the late Eocene of Egypt

A new mammalian genus and species from the earliest late Eocene of Egypt is represented by a lower jaw fragment and two isolated lower molars. A rare combination of features and the fragmentary nature of the materials make their taxonomic assignment to either Marsupialia or Chiroptera uncertain. The holotype of the new genus is the best-preserved specimen in the sample, a dentary fragment with two molars that have uncompressed trigonids, weak buccal cingulids, and a nyctalodont arrangement of the hypocristid. Some traits appear to more clearly support marsupial, as opposed to chiropteran, affinities for this species, including the very likely presence of four molars in the dentary. The enamel of an additional lower molar assigned to this species consists of a single layer of radial enamel without any prism decussation, an enamel type that characterizes marsupials as well as chiropterans. A second taxon is represented by two isolated upper molars. Some traits appear to more clearly support marsupial, as opposed to chiropteran, affinities for these species, including the orientation of pre- and postprotocristae, the moderately slender lingual portion of the trigon, and the absence of any talon expansion. As some of these features can be found in various bat clades, chiropteran affinities for these species cannot be ruled out, but we propose that the new taxa are more likely to represent specialized ‘didelphimorphian’ marsupials.      

A new Late Eocene primate from the Krabi Basin (Thailand) and the diversity of Palaeogene anthropoids in southeast Asia

According to the most recent discoveries from the Middle Eocene of Myanmar and China, anthropoid primates originated in Asia rather than in Africa, as was previously considered. But the Asian Palaeogene anthropoid community remains poorly known and inadequately sampled, being represented only from China, Myanmar, Pakistan and Thailand. Asian Eocene anthropoids can be divided into two distinct groups, the stem group eosimiiforms and the possible crown group amphipithecids, but the phylogenetic relationships between these two groups are not well understood. Therefore, it is critical to understand their evolutionary history and relationships by finding additional fossil taxa. Here, we describe a new small-sized fossil anthropoid primate from the Late Eocene Krabi locality in Thailand, Krabia minuta, which shares several derived characters with the amphipithecids. It displays several unique dental characters, such as extreme bunodonty and reduced trigon surface area, that have never been observed in other Eocene Asian anthropoids. These features indicate that morphological adaptations were more diversified among amphipithecids than was previously expected, and raises the problem of the phylogenetic relations between the crown anthropoids and their stem group eosimiiforms, on one side, and the modern anthropoids, on the other side.     

Taxonomic status of purported primate frontal bones from the Eocene Pondaung Formation of Myanmar

Two isolated cranial fragments from the late middle Eocene Pondaung Formation of central Myanmar have previously been interpreted as frontal bones of the amphipithecid primate Amphipithecus mogaungensis. Aside from a few maxillary fragments, these specimens provide the only potential source of information currently available regarding the cranial anatomy of Amphipithecidae. Were this taxonomic attribution correct, these specimens would indicate that amphipithecids retained numerous primitive skull features, including the absence of a postorbital septum, the retention of a voluminous olfactory chamber, and strong separation between the forebrain and the orbital fossa. However, several anatomical details observable on these specimens are incompatible with their attribution to any primate and strongly suggest that they cannot be ascribed to Mammalia. Particularly problematic in this regard are the extreme thickness of the dermal bone, the odd structure of the alleged "frontal trigon," and the mediolateral orientation and uniquely robust construction of the descending process of the frontal bone (which partially segregates the orbital and temporal fossae). Because these isolated elements can no longer be attributed to Amphipithecus, the anatomical, phylogenetic, and behavioral inferences regarding amphipithecid paleobiology that have been drawn from these specimens can no longer be sustained.     

Postcranial morphology,functional adaptations and palaeobiology of Callistoe vincei,a predaceous metatherian from the Eocene of Salta,north‐western Argentina

Abstract: Callistoe vincei Babot et al., 2002 is a Paleogene borhyaenoid known from exceptionally complete postcranial elements, which provides rare information about the anatomy and evolutionary history of metatherian predators during the South American Cenozoic. The axial skeleton of Callistoe is characterized by the peculiar transverse processes of the cervical vertebrae emphasizing lateral instead of sagittal traction. There is no clavicle and eighteen thoracolumbar vertebrae, of which only five are lumbars. The shoulder and elbow joints suggest movements restricted to parasagittal flexion/extension that are consistent with primarily terrestrial locomotion, as is also emphasized in Borhyaena tuberata and Lycopsis longirostrus. On the manus, the pollex is not reduced and the ungual phalanges indicate very long claws, similar to those observed in some extant digging taxa. This feature is unique to C. vincei among borhyaenoids. The knee joint is characterized by the presence of ossified patellae but shallow femoral trochleae. This joint suggests that the leg was nearly parasagittal, a position also inferred for Borhyaena. The astragalus shape is consistent with parasagittal flexion/extension, as in all Miocene–Pliocene borhyaenoids. The hind foot is characterized by reduced claws in comparison with the manus as well as the slenderness of the first and fifth digits, another peculiarity of C. vincei. The habitat of Callistoe was a temperate humid forest and according to the known fossil record, Callistoe was the largest mammalian predator of its time, sharing the predator ecological niche with crocodiles.     

A highly derived anomalurid rodent (Mammalia) from the earliest Late Eocene of Egypt

Abstract: A new genus and species of diminutive anomalurid rodent, Shazurus minutus, is described on the basis of 15 isolated teeth from the earliest late Eocene (approximately 37 Ma) Birket Qarun Locality 2 in the Fayum Depression of northern Egypt. Shazurus is surprisingly specialized for its age, being most similar in dental morphology to early Miocene Paranomalurus and extant Anomalurus, and is quite different from the roughly contemporaneous anomaluroid genera Nementchamys and Pondaungimys from Algeria and Myanmar, respectively. Parsimony analysis of dental features places Shazurus within crown Anomaluridae as a sister group of two species of Paranomalurus to the exclusion of extant Anomalurus and Idiurus. The marked morphological differences between the two oldest Afro‐Arabian anomaluroids (Shazurus and Nementchamys) suggest that the taxa share a much more ancient common Afro‐Arabian ancestor, possibly derived from Zegdoumyidae. Isolated teeth of Shazurus and other Eocene anomaluroids reveal little about their palaeobiology, but the complete absence of Anomaluroidea from the younger (late Eocene to early Oligocene) Jebel Qatrani Formation is presumably a reflection of environmental change through the late Eocene in northern Africa.     

Oligocene sivaladapid primate from the Bugti Hills (Balochistan, Pakistan) bridges the gap between Eocene and Miocene adapiform communities in Southern Asia

A new species of Guangxilemur (Sivaladapidae, Adapiformes) is described from the early Oligocene Chitarwata Formation (Bugti Member) of the Bugti Hills, Sulaiman geological Province, Balochistan, Pakistan. Guangxilemur singsilai n. sp. provides further diagnostic morphological characters from its newly described upper and lower dentitions, confirming its intermediate phylogenetic position between Eocene and Miocene Asian sivaladapid adapiforms. G. singsilai possesses moderately developed shearing and puncturing molar features and maintains lingual cusps on upper molars as in Eocene hoanghoniines; in contrast, it possesses a typical molariform P(4) as in Miocene sivaladapines. The important paleogeographic changes that have affected South Asia during the Tertiary (related to the collision between the Indian and Eurasian Plates) have played a critical role in reforming circulation and climatic differentiation. The presence in Pakistan of an unique and well-diversified Oligocene primate fauna, clearly demonstrates that South Asia maintained favourable environmental conditions during the middle Caenozoic global climatic deterioration that coincides with drastic changes in faunal structure on the whole Holarctic Province, including the extinction of adapiform primates.     

Comparative functional morphology of the primate peroneal process

The first metatarsal of living Primates is characterized by a well-developed peroneal process, which appears proportionally larger in prosimians than in anthropoids. A large peroneal process has been hypothesized to: 1) reflect powerful hallucal grasping, 2) act as a buttress to reduce strain from loads acting on the entocuneiform-first metatarsal joint during landing and grasping after a leap, and/or 3) correlate with differences in physiological abduction of the hallux. In this study, we address the latter two hypotheses by comparing the morphology of the peroneal process in 143 specimens representing 37 species of extant prosimians, platyrrhine anthropoids, and tupaiids (tree shrews) that engage in different locomotor behaviors. In particular, we compare taxa that vary in leaping frequency and hallucal abduction. Linear and angular measurements on the first metatarsal were obtained to evaluate differences in relative peroneal process thickness and length, first metatarsal abduction angle, and overall first metatarsal shape. Leaping frequency was significantly correlated only with relative peroneal process thickness within extant lorisoids. Relative process length was positively correlated with the angle of hallucal abduction within prosimians; this angle is significantly greater in prosimians than anthropoids. Multivariate analyses of metatarsal shape effectively separate species along phylogenetic lines, but not by locomotor behaviors. The hypothesis that the peroneal process on the first metatarsal reduces the loads on the entocuneiform-first metatarsal joint during landing after a leap is in part supported by data from extant lorisoids (i.e., slow quadrupedal lorises vs. leaping galagos). A peroneal process of greater length within prosimians may serve to increase the lever arm for the peroneus longus muscle in order to prevent hyper-abduction, followed by inversion in locomotor situations where the animal's weight is born on a highly divergent/abducted hallux.     

External morphology of the freshwater prawn Caridina africana collected from Egypt

Abstract

The present paper deals with the external morphology of the freshwater prawn Caridina africana collected from al-Fayum oasis, Egypt. The study includes some morphometric and meristic characters.     

Hindlimb of a giant terrestrial bird from the upper Eocene, Fayum, Egypt

Eremopezus eocaenus Andrews, 1904 is a giant groundbird from upper Eocene deposits of the Fayum, Egypt, which has hitherto been known from non-diagnostic fragmentary material. New fossils collected from quarry L-41 of the Jebel Qatrani Formation include two well-preserved distal tarsometatarsi and an associated whole tarsometatarsus and distal tibiotarsus that allow a more precise evaluation of the phylogenetic position and tarsal function of Eremopezus. Unlike most ratites, the distal tarsometatarsus has a patent distal foramen and a slight hallucal digit. The trochlea for digits II and IV are only slightly reduced in size, are splayed to the right, and the heads lack deep grooving. These features resemble the condition seen in BalaenicepsSagittarius, suggesting active use of the toes in grasping or manipulation, rather than the condition in graviports and cursors, which have reduced medial and lateral trochleae often with distinct grooving of the heads. The limb is relatively long and gracile, another difference from graviports. There is no compelling evidence to link Eremopezus to any known ratite lineage, to the Phorusrhacoidea, or to the extinct predatory birds of the Eocene (e.g. Diatryma, Gastornis). We suggest that Eremopezus represents an endemic African group that independently attained large size and flightlessness.     

Proximal femoral anatomy of a sivaladapid primate from the late middle Eocene Pondaung formation (central Myanmar)

The postcranial anatomy of the Asian sivaladapid adapiforms is still virtually undocumented, whereas dental remains of these primates have been known for several decades. Little is known about their positional behavior as a result. In this article, we describe a partial left femur of a medium-sized primate preserving its entire proximal portion and a significant length of its shaft. This fossil was recently recovered from the fossiliferous locality of Thamingyauk in the late middle Eocene Pondaung Formation (central Myanmar). This femur is considered to pertain to the same individual as two tarsal elements (fragmentary talus and calcaneus) from the same locality (same location), and attributed to a medium-sized sivaladapid adapiform primate (Kyitchaungia takaii). This new postcranial element provides the first documentation of femoral anatomy among Sivaladapidae from Asia. The mechanical implications deriving from the musculoskeletal interpretation of this bone indicate an animal that probably engaged in a kind of active arboreal quadrupedalism with some degree of proficiency in leaping. Even though many musculoskeletal aspects suggest that branch walking and running were important parts of its locomotor repertoire, in other details it appears that relatively complex movements at the hip joint were actually possible and probably associated with climbing or some hindlimb suspensory activities.     

New primates from the late Eocene of Thailand: a contribution to primate diversity in the Paleogene of Asia

We report the discovery of two new primates from the late Eocene Krabi Basin, southern Thailand. One isolated upper molar displays morphological features (protocone united with hypocone by the prehypocrista, postprotocrista extended distobuccally) suggesting possible phylogenetic relationships with Amphipithecidae, while an isolated lower molar is tentatively referred to as a new tarsiiform, mainly on the basis of its paraconid and entoconid morphology. Although very scarce, these remains expand the record of Paleogene primates in Southeast Asia, and testify to their successful radiation in that area.     

New remains of Egatochoerus jaegeri (Mammalia,Suoidea) from the late Eocene of Peninsular Thailand

Abstract: We report here additional remains referred to Egatochoerus jaegeri from the late Eocene locality of Ban Mark in the Krabi basin, Thailand. The new material described, comprising upper and lower cheek teeth, deciduous premolars and partial cranial remains, makes E. jaegeri the best‐documented Eocene representative of Old World Suoidea at present. Detailed study and comparison of their cheek teeth structure reveal a homogeneity of the molar crest and groove patterns of Old World and New World Palaeogene suoids.     

Phosphatized rare star-like mouth disc of Punctatus and its functional morphology from the earliest Cambrian of the South Shaanxi China

The Meishucun stage is the prelude in decipher-ing the Cambrian Explosion. In this prominent stage, rapid radioactive evolution and body-plan innovation have taken place and different associations of organism have been shaped. In this paper we report several 3D-preserved rare star-like fossils with finely preserved soft tissues which were recovered from the Kuanchuanpu Member of the Dengying Formation in South Shaanxi, China in 2003. By studying on functional morphology and analogy with mouthpart of Punctatus, there are evidences that this star-like organism approaches the coelenterates in systematic classification and the centre of star-like organism is its mouth. The appearance of coelenterates marks the real beginning of metazoan evolution. Therefore, it has the prominent position in the origin and evolutionary history of organisms. Perhaps the star-like organism represents the early types of coelenterate with original tentacles. These new materials provide new evidence for the origin, evolution and the functional evolution of the metazoan during the early stage of the Cambrian Explosion.     

Functional and adaptive significance of primate pads and claws: Evidence from New World anthropoids

This study tests predicted morphoclines in fingertip morphology among four small-bodied (<1 kg) New World monkeys (Saimiri sciureus, Leontopithecus rosalia, Callithrix jacchus, and Saguinus oedipus) in order to test previous functional and adaptive explanations for the evolution of flattened nails, expanded apical pads, and grasping extremities within the Order Primates. Small-bodied platyrrhines which frequently forage among small-diameter substrates are expected to possess 1) relatively expanded apical pads, 2) well-developed epidermal ridges, 3) distally broad terminal phalanges, and 4) reduced flexor and extensor tubercles compared to those species which use large-diameter arboreal supports more frequently for their locomotor and postural behaviors. Results show that as the frequency of small-branch foraging increases among taxa within this sample, relative distal phalanx breadth also increases but distal phalanx length, height, and flexor tubercle size decrease. Moreover, epidermal ridge development becomes more pronounced as the frequency of small-branch foraging increases. Terminal phalanx breadth and epidermal ridge complexity are both positively correlated with apical pad size. The large, flexible apical pad increases stability of the hand and foot on small-diameter arboreal supports because the pad can contact the substrate in several planes which, in turn, enables the pad to resist disruptive forces from different directions by friction and interlocking (Hildebrand, 1995). The observed morphoclines demonstrate that a gradient in form from claw- to nail-like tegulae exists among these taxa. Thus, the distinction between claw- and nail-bearing platyrrhines is essentially arbitrary. These observations corroborate Cartmill's (1972) functional and adaptive model for the loss of claws in primates: namely, expanded apical pads are required for habitual locomotor and postural behaviors on small-diameter supports whereas claws are more useful for positional behaviors on large-diameter substrates. Finally, results from this study support previous suggestions that the keeled tegulae of callitrichines represent a derived postural adaptation rather than a primitive retention from an ancestral eutherian condition. Am J Phys Anthropol 106:113–127, 1998. © 1998 Wiley-Liss, Inc.