To discover interordinal relationships of living and fossil placental mammals and the time of origin of placentals relative to the Cretaceous-Paleogene (K-Pg) boundary, we scored 4541 phenomic characters de novo for 86 fossil and living species. Combining these data with molecular sequences, we obtained a phylogenetic tree that, when calibrated with fossils, shows that crown clade Placentalia and placental orders originated after the K-Pg boundary. Many nodes discovered using molecular data are upheld, but phenomic signals overturn molecular signals to show Sundatheria (Dermoptera + Scandentia) as the sister taxon of Primates, a close link between Proboscidea (elephants) and Sirenia (sea cows), and the monophyly of echolocating Chiroptera (bats). Our tree suggests that Placentalia first split into Xenarthra and Epitheria; extinct New World species are the oldest members of Afrotheria.
Plesiadapiforms are central to studies of the origin and evolution of primates and other euarchontan mammals (tree shrews and flying lemurs). We report results from a comprehensive cladistic analysis using cranial, postcranial, and dental evidence including data from recently discovered Paleocene plesiadapiform skeletons (Ignacius clarkforkensis sp. nov.; Dryomomys szalayi, gen. et sp. nov.), and the most plesiomorphic extant tree shrew, Ptilocercus lowii. Our results, based on the fossil record, unambiguously place plesiadapiforms with Euprimates and indicate that the divergence of Primates (sensu lato) from other euarchontans likely occurred before or just after the Cretaceous/Tertiary boundary (65 Mya), notably later than logistical model and molecular estimates. Anatomical features associated with specialized pedal grasping (including a nail on the hallux) and a petrosal bulla likely evolved in the common ancestor of Plesiadapoidea and Euprimates (Euprimateformes) by 62 Mya in either Asia or North America. Our results are consistent with those from recent molecular analyses that group Dermoptera with Scandentia. We find no evidence to support the hypothesis that any plesiadapiforms were mitten-gliders or closely related to Dermoptera.Euarchonta ͉ phylogeny ͉ Paromomyidae ͉ Micromomyidae ͉ Paleogene T he origin of Primates represents the first clear step in the divergence of humans from the rest of Mammalia, yet our understanding of this important period in evolutionary history remains limited. The systematic relationships of Paleocene-Eocene plesiadapiforms, which have been considered the ancestors of either Euprimates (primates of ''modern aspect'' or crown-clade primates) (1, 2) or of Dermoptera (3, 4) continue to be debated. Clarifying the position of plesiadapiforms is central to understanding the broader relationships among euarchontan mammals (Primates, Scandentia, Dermoptera), and to testing adaptive hypotheses of primate origins (5, 6) by using direct evidence from the fossil record.Plesiadapiforms are among the most diverse and well sampled Paleogene mammal groups, with Ͼ120 species classified into 11 or 12 families from the Paleocene and Eocene of North America, Europe, Asia, and possibly Africa (7,8). The plesiadapiform dental record is extremely diverse, suggesting correlated diversity in diet and behavior; however, comparatively little is known about the cranial or postcranial morphology of plesiadapiforms [see supporting information (SI) Text, Part 1]. Well preserved crania have been documented for only three families: Plesiadapidae, Microsyopidae, and Paromomyidae (1, 9-11). Postcrania are known from a taxonomically limited sample of North American and European plesiadapids (1), from a sample of North American paromomyids and micromomyids (3, 4, 12) the identification and associations of which are still controversial (13,14), from a recently published North American carpolestid skeleton (15, 16), and from a few other isolated and questionably identified elements (7,17,18). Following the sugges...
In this study, the forelimb of 12 species of tupaiids was analyzed functionally and compared to that of other archontan mammals. Several differences that relate to differential substrate use were found in the forelimb morphology of tupaiids. These differences included shape of the scapula, length and orientation of the coracoid process, size of the lesser tuberosity, shape of the capitulum, length of the olecranon process, and shape of the radial head and central fossa. The forelimb of the arboreal Ptilocercus lowii, the only ptilocercine, is better adapted for arboreal locomotion, while that of tupaiines is better adapted for terrestrial (or scansorial) locomotion. While the forelimb of the arboreal Ptilocercus appears to be habitually flexed and exhibits more mobility in its joints, a necessity for movement on uneven, discontinuous arboreal supports, all tupaiines are characterized by more extended forelimbs and less mobility in their joints. These restricted joints limit movements more to the parasagittal plane, which increases the efficiency of locomotion on a more even and continuous surface like the ground. Even the most arboreal tupaiines remain similar to their terrestrial relatives in their forelimb morphology, which probably reflects the terrestrial ancestry of Tupaiinae (but not Tupaiidae). The forelimb of Urogale everetti is unique among tupaiines in that it exhibits adaptations for scratch-digging. Several features of the tupaiid forelimb reflect the arboreal ancestry of Tupaiidae and it is proposed that the ancestral tupaiid was arboreal like Ptilocercus. Also, compared to the forelimb character states of tupaiines, those of Ptilocercus are more similar to those of other archontans and it is proposed that the attributes of the forelimb of Ptilocercus are primitive for the Tupaiidae. Hence, Ptilocercus should be considered in any phylogenetic analysis that includes Scandentia.
Arboreal and semiterrestrial guenons show similar osteological features of the limbs across a wide range of species, environments, and geography, while the more terrestrially committed guenons exhibit greater morphological divergence. An ecomorphological comparison of two sympatric guenons living in Kibale Forest, Uganda, reveals an array of anatomical adaptations for terrestriality in the limbs of Cercopithecus lhoesti similar to those found in Erythrocebus patas. In contrast, Cercopithecus aethiops, although also frequent users of the terrestrial environment, generally exhibit fewer morphological adaptations characteristic of a terrestrial lifestyle. It appears that significant morphological modification for terrestriality has occurred twice within the diverse radiation of living guenons with C. aethiops perhaps representing a third group in the making.
In this study, the hindlimb of 12 species of tupaiids was analyzed functionally and compared to that of primates, dermopterans, and chiropterans. Many aspects of the tupaiid hindlimb vary in relation to differential substrate use. These differences include width of the ilium, shape of the acetabulum, size of the anterior inferior iliac spine, size of the greater and third trochanters, depth of the femoral condyles, shape of the patellar groove, and size of the tibial tuberosity. The hindlimb of the arboreal Ptilocercus lowii, the only ptilocercine, is better adapted for arboreal locomotion, whereas that of tupaiines is better adapted for rapid terrestrial (or scansorial) locomotion. The hindlimb of Ptilocercus seems to be habitually flexed and has more joint mobility, a condition necessary for movement on uneven, discontinuous arboreal supports. The tarsus of Ptilocercus facilitates inversion of the foot and its grasping hallux is capable of a great range of abduction. Tupaiines, on the other hand, are characterized by more extended hindlimbs and less mobility in their joints. These restricted joints limit movements more to the parasagittal plane, which increases the efficiency of locomotion on a more even and continuous surface like the ground. The hindlimb of tupaiines is adapted for powerful flexion and extension. Even the most arboreal tupaiines remain similar to terrestrial tupaiines in their hindlimb morphology, which probably reflects the terrestrial ancestry of Tupaiinae (but not Tupaiidae). Many attributes of the tupaiid hindlimb, especially those of the foot, reflect the arboreal ancestry of Tupaiidae and it is proposed that the ancestral tupaiid was arboreal like Ptilocercus. Also, compared to the hindlimb character states of tupaiines, those of Ptilocercus are more similar to those of other archontans, and it is proposed that the hindlimb features of Ptilocercus are primitive for the Tupaiidae. Hence, Ptilocercus should be considered in any phylogenetic analysis that includes Scandentia.
A new species of African monkey, Lophocebus kipunji, was described in 2005 based on observations from two sites in Tanzania. We have since obtained a specimen killed by a farmer on Mount Rungwe, the type locality. Detailed molecular phylogenetic analyses of this specimen demonstrate that the genus Lophocebus is diphyletic. We provide a description of a new genus of African monkey and of the only preserved specimen of this primate. We also present information on the animal's ecology and conservation.
In this study, the axial skeleton of 14 species of tupaiids (tree shrews) was analysed functionally and compared to that of other archontan mammals. Several differences that relate to differential substrate use were found in the ribs and vertebrae. These differences included cranio-caudal width of the ribs; number of thoracic, lumbar, and caudal vertebrae; cranio-caudal width of the atlas; orientation of the spinous process of the axis; length and cranio-caudal width of the spinous processes of the thoracic vertebrae; length of the spinous processes of the lumbar vertebrae; length and orientation of the transverse processes of the lumbar vertebrae; and the number of sacral vertebrae that articulate with the ilia. The ribs and vertebrae of the arboreal Ptilocercus lowii, the only ptilocercine, exhibit adaptations for a stable thorax that probably facilitate bridging locomotion. The vertebral columns of tupaiines, on the other hand, are more mobile and allow more¯exion and extension of the spine; this increased¯exion and extension increases stride length, which in turn increases speed in bounding or galloping mammals such as terrestrial tupaiines. It is proposed here that the attributes of the thorax of Ptilocercus are primitive for the Tupaiidae, that the ancestral tupaiid was arboreal, that the tupaiine condition is derived, and that the ancestral tupaiine was terrestrial. It is also proposed that: Ptilocercus may be primitive for the Archonta in its axial skeletal features; a stable thorax was ®rst evolved in an arboreal ancestral archontan; the adaptations for stability of the thorax were retained in the Volitantia (dermopterans and chiropterans) for certain locomotor types, including gliding or¯ying; a mobile thorax evolved in conjunction with the shift to graspleaping in the ancestral euprimate. These scenarios may be further tested by quantitative analyses of vertebral osteology, as well as myological analyses of the epaxial musculature.
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