Scaling of lumbar vertebrae in anthropoids and implications for evolution of the hominoid axial skeleton

We investigated allometric relationships between vertebral centrum cranial surface areas and body weight and skeletal lumbar length in extant platyrrhine and cercopithecid species. Platyrrhines have smaller lumbar vertebral centra regarding the cranial surface area relative to their body weight than extant catarrhines. However, the stress to the spine of quadrupeds is not only influenced by the body weight but also its length, which contributes to the amount of bending moment. Our results indicated that platyrrhines and cercopithecids have similar lumbar vertebral centrum surface areas when they are scaled on the product of the body weight and skeletal lumbar length. Platyrrhines generally tend to have relatively short lumbar columns for a given body weight. As a result of this tendency, their vertebral centra appear relatively small if only body weight is taken into account. The centrum surface area is rather constant relative to the product of the body weight and skeletal lumbar length within platyrrhines or cercopithecids, despite the fact that skeletal lumbar length is in itself rather variable relative to body weight. This result indicates that the vertebral centrum articular area, the lumbar column length and the body weight are strongly correlated with each other and that such relationships are similar between platyrrhines and cercopithecids. These relationships were observed using both the zygapophyseal and rib definitions of the lumbar vertebrae. However, they were more clearly observed when the zygapophyseal definition was adopted. It appeared that lumbar vertebrae of Proconsul nyanzae (KNM−MW 13142) had distinctively smaller surface areas relative to its body weight and lumbar length than for platyrrhines and cercopithecids, differing from extant hominoids, which have comparatively larger lumbar vertebrae. In the case of Morotopithecus, the lumbar vertebral surface area seems to be as large as in extant platyrrhines and cercopithecids if it had a reduced number of lumbar vertebrae. It is uncertain whether its lumbar vertebral surface area was as large as in extant hominoids. Electronic Publication     

First hominoid from the Miocene of Ethiopia and the evolution of the catarrhine elbow

The first known fossil ape from the early-middle Miocene of Fejej, Ethiopia, is described here. The specimen, FJ-18SB-68, is a partial ulna from a locality dated by ⁴⁰Ar/³⁹Ar and paleomagnetic methods to a minimum age of 16.18 MYA. Compared to a variety of extant and fossil ulnae, FJ-18SB-68 is most similar to Turkanapithecus, Proconsul, and Pliopithecus, and appears to have been an arboreal quadruped with substantial forearm rotational mobility. Among the extant ulnae, canonical variates analysis successfully discriminates platyrrhines from catarrhines and within the latter, cercopithecoids from hominoids. Basal catarrhines (e.g., Aegyptopithecus) are platyrrhine-like in their morphology. Two basic trends appear to evolve from this generalized template: one with less mobile and more habitually pronated forearms, as seen in living and fossil cercopithecoids (including Victoriapithecus and Paracolobus), and another with greater forearm rotational mobility in fossil and modern hominoids. Primitive Miocene apes, including Proconsul, Turkanapithecus, and FJ-18SB-68, share with extant hominoids a more laterally positioned and laterally facing radial notch and an incipient trochlear keel. This morphology, along with a large insertion area for m. brachialis, suggests a departure from the more habitually pronated hand posture of monkeys and may indicate greater climbing abilities in these arboreally quadrupedal apes. Later Miocene apes, such as Oreopithecus and Dryopithecus share additional morphological features with hominoids, indicating considerable suspensory and climbing capabilities. Am J Phys Anthropol 105:257–277, 1998. © 1998 Wiley-Liss, Inc.     

The fossil evidence of anthropoid brain evolution

The endocast of Aegyptopithecus, a 27 million year old ape, reveals that its brain was advanced over that of prosimians and comparable to that of modern anthropoids in relative size and in having expanded visual cortex, reduced olfactory bulbs, and a central sulcus separating primary somatic sensory and motor cortex. The early appearance of those features suggests that they may have been among the adaptations responsible for the evolution of anthropoids from prosimian ancestors. The frontal lobe was relatively smaller in Aegyptopithecus than in modern anthropoids. An endocast of Dolichocebus, one of the oldest known New World monkeys (25–30 million years old), reveals visual cortex expanded as in modern anthropoids. The 19 million year old Napak frontal bone displays a hominoid rather than cercopithecoid sulcal pattern. An 18 million year old endocast of the ape Dryopithecus (Proconsul) was neither monkey-like nor primitive, as originally described, but rather apelike and essentially modern in all observable features. The oldest undoubted Old World monkey endocast, from nine million year old Mesopithecus, reveals that the brain was modern in sulcal pattern and proportions. The sulcal pattern was like that of modern colobines, but that appears to be the more primitive condition, from which features characteristic of modern cercopithecine brains have evolved. The brain of six million year old Libypithecus was similar to that of Mesopithecus. A two million year old endocast of “Dolichopithecus” arvernensis displays a modern cercopithecine sulcal pattern.     

Significance of primate petrosal from Middle Eocene fissure-fillings at Shanghuang, Jiangsu Province, People's Republic of China

An isolated petrosal bone belonging to a diminutive primate is reported from Middle Eocene fissure-fills near Shanghuang (southern Jiangsu Province, People's Republic of China), the type locality of several newly described primates (Eosimias sinensis, a basal anthropoid; Adapoides troglodytes, a basal adapinan; Tarsius eocaenus, a congener of extant tarsiers; and Macrotarsius macrorhysis, the first Asian representative of an otherwise exclusively North American genus). Because of its fragmentary condition and unique combination of characters, the Shanghuang petrosal cannot be assigned unambiguously to any of the Shanghuang primate taxa known from dental remains. However, the possibility that the petrosal represents either an adapid or a tarsiid can be dismissed because it lacks defining basicranial apomorphines of these groups. By contrast, the element does present arterial features consistent with its being haplorhine. Deciding between the likeliest candidates for its allocation—Omomyidae and Eosimiidae—is difficult, in part because it is not known what (or even whether) basicranial characters can be used to distinguish these clades. If the Shanghuang petrosal is that of an cosimiid, as both direct and indirect evidence appears to indicate, the following implications emerge: (1) as long suspected on other grounds, anthropoids share a closer evolutionary history with Omomyidae (and Tarsiiformes) than they do with Adapidae (and Strepsirhini); (2) the specialised basicranial anatomy of extant anthropoids and their immediate cladistic relatives is derived from a primitive precursor whose otic morphology was like that of omomyids in most known respects; (3) the evolution of the defining dental and basicranial apomorphies of extant Anthropoidea has been distinctly mosaic in pattern.     

Partial skeleton of Theropithecus brumpti (Primates, Cercopithecidae) from the Chemeron Formation of the Tugen Hills, Kenya

Here we describe a complete skull and partial skeleton of a large cercopithecoid monkey (KNM-TH 46700) discovered in the Chemeron Formation of the Tugen Hills at BPRP Site #152 (2.63 Ma). Associated with the skeleton was a mandible of an infant cercopithecoid (KNM-TH 48364), also described here. KNM-TH 46700 represents an aged adult female of Theropithecus brumpti, a successful Pliocene papionin taxon better known from the Omo Shungura Formation in Ethiopia and sites east and west of Lake Turkana, Kenya. While the morphology of male T. brumpti is well-documented, including a partial skeleton with both cranial and postcranial material, the female T. brumpti morphotype is not well-known. This skeleton represents some of the first associated evidence of cranial and postcranial female T. brumpti remains. In addition to the complete skull, postcranial material includes elements of the axial skeleton and lower limb. While aspects of the skeleton conform to those of specimens previously assigned to T. brumpti, other features on the femur and tibia appear to differ from those previously described for this species. It is unclear whether these differences represent general variation within the T. brumpti population, variation between the sexes in T. brumpti, or the incorrect assignment of previous isolated hindlimb specimens. In total, the observable morphological features of the hindlimb suggest that KNM-TH 46700 was a terrestrial quadruped similar to modern savannah baboons (Papio). From the available evidence, it is difficult to assess whether or not KNM-TH 46700 frequently engaged in the specialized squatting and shuffling behavior observed in extant geladas (Theropithecus gelada).     

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.     

A new hominoid hamate and first metacarpal from the Late Miocene Nagri Formation of Pakistan

A hamate and the proximal part of a first metacarpal from the type locality of the Nagri Formation in Pakistan, and attributed to Sivapithecus parvada, are described. In overall proportions, the hamate is rather robust, showing most similarity to that of Gorilla. Unlike extant hominoids it lacks a well-developed hamulus, and its triquetral facet is morphologically dissimilar to that in extant anthropoids. The morphology of the hamate indicates effective weight transmission through the ulnar side of the wrist, limited ulnar deviation and restricted extension in the triquetrohamate joint, and stability of the hamatometacarpal joints. The morphology of the partial first metacarpal is most similar to that of Pan. Previously described postcranial bones of S. parvada indicate that its locomotor behaviour included both quadrupedalism and climbing. This is consistent with the limited evidence of the first metacarpal, whereas the hamate strongly emphasizes the quadrupedal aspect of the locomotor repertoire.     

The oldest ape

This paper recounts the history and significance of Morotopithecus bishopi, an early Miocene East African ape. Morotopithecus differs in intriguing ways from its contemporary Proconsul. While craniodental differences are slight, the known elements of its postcranium suggest that Morotopithecus was capable of modern ape–like positional behaviors, including vertical postures, deliberate climbing, and arm hanging. In contrast, Proconsul and other early and middle Miocene hominoids have been reconstructed as above–branch quadrupeds. Paleoanthropologists are currently divided over whether and which of the anatomical features associated with upright posture and suspension in the modern apes are due to inheritance or independent evolution. This debate has important implications for interpreting the phylogenetic positions of both Morotopithecus and Proconsul, as well as for reconstructing the pattern and timing of the emergence of modern ape adaptations.     

A mandible ofBranisella boliviana (Platyrrhini,Primates) from the oligocene of South America

A mandibular specimen from the Bolivian Early Oligocene is provisionally assigned toBranisella boliviana. The crown anatomy of the single preserved tooth, an M₂, indicates platyrrhine affinities and several details of the broken jaw are suggestive of symphyseal fusion. Like the African Oligocene parapithecids,Branisella contrasts with extant anthropoids in the relative shallowness of its mandible.Branisella is the most ancient, and seemingly the most primitive, fossil platyrrhine monkey, lacking any of the derived features of the two major clades of modern ceboids. Taxonomically, it is best regarded as family incertae sedis.     

Stance and gait in the flesh-eating dinosaur Tyrannosaurus

A study of the stance and locomotion of Tyrannosaurus was made for the mounting of the partial skeleton at the British Museum (Natural History). This shows that the posture was much more bird-like than is indicated by previous mounts, and also the tail is shorter. During walking the vertebral column was held nearly horizontal with the tail clear of the ground. The fore-limbs acted as struts to stop the body sliding forward as the animal raised its body from the resting position.     

Knee function through finite element analysis and the role of Miocene hominoids in our understanding of the origin of antipronograde behaviours: the Pierolapithecus catalaunicus patella as a case study

Although extensive research has been carried out in recent years on the origin and evolution of human bipedalism, a full understanding of this question is far from settled. Miocene hominoids are key to a better understanding of the locomotor types observed in living apes and humans. Pierolapithecus catalaunicus, an extinct stem great ape from the middle Miocene (c. 12.0 Ma) of the Vallès-Penedès Basin (north-eastern Iberian Peninsula), is the first undoubted hominoid with an orthograde (erect) body plan. Its locomotor repertoire included above-branch quadrupedalism and other antipronograde behaviours. Elucidating the adaptive features present in the Pierolapithecus skeleton and its associated biomechanics helps us to better understand the origin of hominoid orthogrady. This work represents a new biomechanical perspective on Pierolapithecus locomotion, by studying its patella and comparing it with those drawn from a large sample of extant anthropoids. This is the first time that the biomechanical patellar performance in living non-human anthropoids and a stem hominid has been studied using finite element analysis (FEA). Differences in stress distribution are found depending on body plan and the presence/absence of a distal apex, probably due to dissimilar biomechanical performances. Pierolapithecus’ biomechanical response mainly resembles that of great apes, suggesting a similar knee joint use in mechanical terms. These results underpin previous studies on Pierolapithecus, favouring the idea that a relevant degree of some antipronograde behaviour may have made up part of its locomotor repertoire. Moreover, our results corroborate the presence of modern great ape-like knee biomechanical performances back in the Miocene.     

A closer look at the “Protopithecus” fossil assemblages: new genus and species from Bahia,Brazil

The recently extinct large-bodied New World monkey Protopithecus brasiliensis Lund 1836 was named based on a distal humerus and proximal femur found in the Lagoa Santa cave system in the southeastern Brazilian state of Minas Gerais. These bones are from an animal about twice the size of the largest extant platyrrhines. One hundred and seventy-five years later, a nearly complete skeleton was discovered in the Toca da Boa Vista caves in the neighboring state of Bahia and was allocated to the same taxon as it was the first platyrrhine fossil of comparable size found since the originals. Our detailed study of the equivalent elements, however, reveals important morphological differences that do not correspond to intraspecific variation as we know it in related platyrrhine taxa. The presence of both an expanded brachioradialis flange on the humerus and gluteal tuberosity on the femur of the Bahian skeleton distinguishes it from the Lagoa Santa fossil as well as from all other platyrrhines. Further cranial and postcranial evidence suggests a closer relationship of the former with the alouattine Alouatta, while the limited Lund material fits more comfortably with the ateline clade. Therefore, we propose to limit P. brasiliensis Lund to the distal humerus and proximal femur from Lagoa Santa and erect a new genus and species for the skeleton from Toca da Boa Vista. Cartelles coimbrafilhoi was a large-bodied frugivore with a relatively small brain and diverse locomotor repertoire including both suspension and climbing that expands the range of platyrrhine biodiversity beyond the dimensions of the living neotropical primates.     

New wrist bones of Proconsul africanus and P. nyanzae from Rusinga Island, Kenya

The anatomy of the wrist of two species of the early Miocene hominoid Proconsul is described based on new material collected on Rusinga Island, Kenya. These fossils generally confirm previous findings that the wrist of Proconsul is monkey-like in much of its morphology. However, the structure of the ulnar side of the wrist, particularly the ulnocarpal joint, is significantly different from that of extant monkeys and suggests some functional affinities with extant hominoids. Thus the wrist of Proconsul is neither monkey-like nor ape-like in its total morphology. Instead, it shows a unique combination of features which once again point to the oversimplicity of forcing fossil forms into categories based only on extant taxa.     

Evidence for a grooming claw in a North American adapiform primate: implications for anthropoid origins

Among fossil primates, the Eocene adapiforms have been suggested as the closest relatives of living anthropoids (monkeys, apes, and humans). Central to this argument is the form of the second pedal digit. Extant strepsirrhines and tarsiers possess a grooming claw on this digit, while most anthropoids have a nail. While controversial, the possible presence of a nail in certain European adapiforms has been considered evidence for anthropoid affinities. Skeletons preserved well enough to test this idea have been lacking for North American adapiforms. Here, we document and quantitatively analyze, for the first time, a dentally associated skeleton of Notharctus tenebrosus from the early Eocene of Wyoming that preserves the complete bones of digit II in semi-articulation. Utilizing twelve shape variables, we compare the distal phalanges of Notharctus tenebrosus to those of extant primates that bear nails (n = 21), tegulae (n = 4), and grooming claws (n = 10), and those of non-primates that bear claws (n = 7). Quantitative analyses demonstrate that Notharctus tenebrosus possessed a grooming claw with a surprisingly well-developed apical tuft on its second pedal digit. The presence of a wide apical tuft on the pedal digit II of Notharctus tenebrosus may reflect intermediate morphology between a typical grooming claw and a nail, which is consistent with the recent hypothesis that loss of a grooming claw occurred in a clade containing adapiforms (e.g. Darwinius masillae) and anthropoids. However, a cladistic analysis including newly documented morphologies and thorough representation of characters acknowledged to have states constituting strepsirrhine, haplorhine, and anthropoid synapomorphies groups Notharctus tenebrosus and Darwinius masillae with extant strepsirrhines rather than haplorhines suggesting that the form of pedal digit II reflects substantial homoplasy during the course of early primate evolution.     

The phylogenetic significance of strepsirhinism in Paleogene primates

Two lines of evidence reviewed here argue against the recent proposal that strepsirhinism is an apomorphous feature in primates, shared only by adapiforms and lemuriforms. These are (1) the presence of strepsirhinism in several extant taxa of nonprimate mammals, including Tupaiidae, Tenrecidae, Erinaceus,and Didelphis,and (2) the inferred presence of strepsirhinism not only in adapiforms, but also in all plesiadapiforms and omomyids for which the relevant anatomical regions are known. Therefore, strepsirhinism cannot be invoked as an adaptive innovation underlying the initial strep-sirhine/haplorhine dichotomy. Likewise, the apparent retention of strepsirhinism in omomyids suggests either that the haplorhine oronasal configuration of extant tarsiids and anthropoids was acquired independently or that tarsiids and anthropoids form a clade to the exclusion of omomyids.     

Early Eocene fossil illuminates the ancestral (diurnal) ecomorphology of owls and documents a mosaic evolution of the strigiform body plan

We describe a partial skeleton of a fossil owl (Strigiformes) from the early Eocene London Clay of Walton-on-the-Naze (Essex, UK). The holotype of Ypresiglaux michaeldanielsi, gen. et sp. nov. is one of the most complete specimens of a Palaeogene owl and elucidates the poorly known ecomorphology of stem group Strigiformes. Whereas most of the postcranial bones show the characteristic strigiform morphology, the new species exhibits plesiomorphic features of the skull and cervical vertebrae that differ distinctly from extant owls. A well-developed supraorbital process of the lacrimal bone suggests that the eyes were not as greatly enlarged and forward-facing as in extant owls. A plesiomorphic quadrate morphology indicates differences in the otic region, and a proportionally longer axis suggests that the fossil species was not able to rotate its head to the degree found in crown group Strigiformes. Therefore, the fossil documents a mosaic evolution of the strigiform body plan, with owls developing raptorial adaptations before specializations of the visual and acoustic systems evolved. Because the latter relate to a crespuscular or nocturnal activity pattern, we hypothesize that Ypresiglaux was diurnal. Nocturnality in owls may have evolved in response to the emergence of evolutionary opportunities, which enabled owls to exploit new ecological niches, or owls may have been driven into nocturnal habits by ecological competition.     

Functional aspects of the postcranial anatomy of the Permian dicynodont Diictodon and their ecological implications

The postcranial skeleton of the Permian dicynodont Diictodon is described, major skeletal muscles restored and functional aspects of the skeleton analyzed. The humerus was caudolaterally oriented and articulated in a near sagittal plane. This reduced the transverse component of thrust during locomotion. Throughout the stride, the femur, which is relatively longer than the humerus, was held in a parasagittal position. Diictodon probably had a hindlimb gait similar to the high walk of crocodilians. Comparisons between several dicynodont genera, including Diictodon, reveal considerable variation in the limb orientations, articulations, posture and gait among these taxa. Expansion of the preacetabular iliac process and the overall iliac length with respect to its height among these dicynodonts may be correlated with the increase in the number of the sacral vertebrae. Diictodon does not show the cranial and forelimb modifications suitable for digging as seen in Cistecephalus, Kawingasaurus and extant burrowers such as a broad skull, rounded occiput for considerable neck musculature, a robust humerus and a prominent olecranon process. However, a cylindrical body, short limbs with well–developed limb elevators and depressors and a long wide manus with long, blunt claws probably helped in digging. In addition, the hindlimb with well–developed retractors and short, blunt claws participated in soil removal. The caudolaterally oriented humerus and a laterally flexible vertebral column facilitated movement in the narrow confines of the burrows. Forelimb articulation and morphology indicate that its method of digging was probably rotation–thrust, where humeral excursion and not forearm extension played a dominant role. The close associations of articulated paired skeletons of adult individuals suggest that Diictodon was gregarious. A large number of unconnected burrow–casts in a small area of about 500 m², indicates that although the animals lived close together, they did not share a single large communal structure.     

Skull of early Eocene Cantius abditus (primates: Adapiformes) and its phylogenetic implications,with a reevaluation of “Hesperolemur” actius

A substantially complete skull and mandible of the primitive adapiform Cantius is reported from the Early Eocene Willwood Formation of Wyoming. The mandible contains an almost complete lower dentition in which the lower incisors are strongly inclined and have spatulate crowns, I₂ is larger than I₁, and the canine is large and projecting. The cranium shares many features with those of Notharctus and Smilodectes but differs in having nasals that broaden proximally. Presence of a prominent canine and strong sagittal crest may indicate that it represents a male. The basicranium preserves auditory structures almost identical to those in extant noncheirogaleid lemurs, including a large bony tube for the stapedial artery and a small, open sulcus for the distal portion of the promontorial artery. The dentition is sufficiently primitive to be compatible with a relationship to either strepsirrhines or anthropoids, but the anatomy of the auditory region is more consistent with either specific relationship to lemurs or, more likely, a basal position that approximates the euprimate morphotype. Certain features of the basicranium of “Hesperolemur” actius, described by Gunnell ([1995] Am. J. Phys. Anthropol. 98:447–470) as being unlike that of any other adapiform, were either misinterpreted or are apparently no longer present in the holotype. Reassessment of these and other features indicates that in fact “H.” actius differs little from Cantius and should not be separated from the latter at the genus level, although on dental grounds the species appears to be distinct (as C. actius). Am J Phys Anthropol 109:523–539, 1999. © 1999 Wiley-Liss, Inc.     

New material of Qatrania from Egypt with comments on the phylogenetic position of the parapithecidae (primates,Anthropoidea)

New material of the early anthropoid primate Qatrania wingi and a new species of that genus are described. Several features of the dental anatomy show that Qatrania, while quite primitive relative to other anthropoids in many ways, is most likely a parapithecid primate. The new material suggests that several dental features previously thought to ally parapithecids with the catarrhine primates were actually evolved in parallel in catarrhines and some parapithecids. Furthermore, all nonparapithecid anthropoids (including platyrrhines and catarrhines) share a suite of derived dental and postcranial features not found in parapithecids. Therefore, parapithecid origins may predate the platyrrhine/catarrhine split.