Preliminary analysis of<Emphasis Type="Italic"> Nacholapithecus</Emphasis> scapula and clavicle from Nachola,Kenya

The Miocene ape Nacholapithecus is known from rather complete skeletons; some of them preserve the shoulder joint, identified by three scapulae and one clavicle. Comparisons made with other Miocene and living apes (Proconsul, Equatorius, Ugandapithecus) suggest that the mobility of the scapulohumeral joint was important, and scapular features such as the morphology and position of the spine and the morphology of the acromion and axillary border resemble those of climbing arboreal primates except for chimpanzees, gorillas, or orang-utans. From the size of the scapula (male Nasalis size), it is clear that the animal is smaller than an adult chimpanzee, but the clavicle is almost as relatively long as those of chimpanzees. Some features closer to colobine morphology reinforce the hypothesis that Nacholapithecus was probably a good climber and was definitely adapted for an arboreal life.     

New Sivapithecus postcranial specimens from the Siwaliks of Pakistan

Several new postcranial elements of Sivapithecus from the Siwaliks of Pakistan are described. These include a distal femur from the U-level of the Dhok Pathan Formation, a navicular from the Chinji Formation, and seven manual and pedal phalanges from the Nagri Formation. The functional morphology of these elements adds new detail to the reconstruction ofSivapithecus positional behavior. Femoral cross-sectional geometry indicates that the shaft was adapted to support mediolaterally directed loading. Femoral condylar asymmetry and a broad but shallow trochlea are distinctly ape-like, revealing capabilities for both rotation and withstanding eccentric loading in the knee. The navicular is characterized by features relating to a broad mid-tarsus and broad distal articulations for the cuneiforms. It also lacks a navicular tubercle as in Pongo. These features suggest that the foot was capable of a powerful grip on large supports, with an inversion/supination capability that would permit foot placement in a variety of positions. The morphology of the new phalanges, including evidence for a relatively large pollex, similarly suggests powerful grasping, consistent with prior evidence from the hallux and tarsus. The functional features of the new specimens permit refinement of previous interpretations of Sivapithecus positional capabilities. They suggest a locomotor repertoire dominated by pronograde activities and also such antipronograde activities as vertical climbing and clambering, but not by antipronograde suspensory activities as practiced by extant apes.     

New evidence for diet and niche partitioning in Rudapithecus and Anapithecus from Rudabánya,Hungary

Rudabánya is rare among Eurasian Miocene fossil primate localities in preserving both a hominid and pliopithecoid, and as such provides the unique opportunity to reconstruct the nature of sympatry and niche partitioning in these taxa. Rudapithecus and Anapithecus have similar locomotor and positional behavior and overlapping body mass ranges. While prior analyses of molar occlusal anatomy and microwear identify Rudapithecus as a soft-object frugivore, reconstructing the dietary behavior of Anapithecus has been more problematic. This taxon has been interpreted to be more folivorous by some, and more frugivorous by others. Here, we use high-resolution polynomial curve fitting (HR-PCF) to quantify and evaluate the mesiodistal and cervico-incisal curvatures of the incisor crowns of Rudapithecus and Anapithecus to identify diet-specific morphological variation in these taxa. Results are consistent with the interpretation that Anapithecus and Rudapithecus were primarily frugivorous and had diets that included similar resource types. However, Anapithecus may have consumed greater amounts of foliage, similar to extant mixed folivore–frugivores (i.e., Gorilla gorilla gorilla, Symphalangus syndactylus), while Rudapithecus generated elevated compressive loads in the incisor region consistent with a specialized role for the anterior dentition in food processing (i.e., removal of tough protective fruit pericarps). We interpret these findings in light of the paleoecology at Rudabánya and conclude that, if these taxa were indeed sympatric, Anapithecus may have used additional leaf consumption as a seasonal fallback resource to avoid direct competition with Rudapithecus. Conversely, Rudapithecus may have relied on less preferred and harder fruiting resources as a seasonal fallback resource during periods of fruit scarcity.     

A hominoid distal tibia from the Miocene of Pakistan

A distal tibia, YGSP 1656, from the early Late Miocene portion of the Chinji Formation in Pakistan is described. The fossil is 11.4 million years old and is one of only six postcranial elements now assigned to Sivapithecus indicus. Aspects of the articular surface are cercopithecoid-like, suggesting some pronograde locomotor activities. However, YGSP 1656 possesses an anteroposteriorly compressed metaphysis and a mediolaterally thick medial malleolus, ape-like features functionally related to orthograde body postures and vertical climbing. YGSP 1656 lacks specializations found in the ankle of terrestrial cercopithecoids and thus Sivapithecus may have been primarily arboreal. Nevertheless, the morphology of this tibia is unique, consistent with other interpretations of Sivapithecus postcranial functional morphology that suggest the locomotion of this ape lacks a modern analog. Based on the limited postcranial remains from S. indicus, we hypothesize that this taxon exhibited substantial body size dimorphism.     

Knuckle-walking in Sivapithecus? The combined effects of homology and homoplasy with possible implications for pongine dispersals

Sivapithecus is a Miocene great ape from South Asia that is orangutan-like cranially but is distinctive postcranially. Work by others shows that the humerus resembles large terrestrial cercopithecoids proximally and suspensory hominoids distally, but most functional interpretations nevertheless situate Sivapithecus in an arboreal setting. We present a new quantitative analysis of the Sivapithecus capitate and hamate. Though the functional morphology of both bones suggests some degree of arboreality, the overall morphology is most similar to knuckle-walking African apes. Other features of the Sivapithecus humerus and hind limb are also functionally consistent with knuckle-walking, and we suggest that this locomotor behavior is a valid alternative functional interpretation of the postcranial morphology. We speculate that knuckle-walking in Sivapithecus would have evolved independently from African apes, perhaps for similar ecological reasons. The discovery of a possible pongine knuckle-walker challenges the hypotheses that (1) knuckle-walking evolved only once in hominoids and (2) knuckle-walking is too highly specialized to be the positional behavior from which human bipedalism evolved. The possibility of knuckle-walking in Sivapithecus may help to explain not only the curious combination of characters that typify the postcranium but also the unique postcranial morphology of extant Pongo. Furthermore, it may clarify the distribution of fossil pongines across many ecological zones in Eurasia in the Miocene and Pleistocene, as well as, independently, the spread of African apes across a diversity of environments in equatorial Africa.     

Morphometric analysis of hominoid lower molars from Yuanmou of Yunnan Province, China

Shape analyses of cross-sectional mandibular molar morphology, using Euclidean Distance Matrix Analysis, were performed on 79 late Miocene hominoid lower molars from Yuanmou of Yunnan Province, China. These molars were compared to samples of chimpanzee, gorilla, orangutan,Lufengpithecus lufengensis, Sivapithecus, Australopithecus afarensis, and human mandibular molars. Our results indicate that the cross-sectional shape of Yuanmou hominoid lower molars is more similar to the great apes that to humans. There are few differences between the Yuanmou,L. lufengensis, andSivapithecus molars in cross-sectional morphology, demonstrating strong affinities between these three late Miocene hominoids. All three of the fossil samples show strong similarities to orangutans. From this, we conclude that these late Miocene hominoids are more closely related to orangutants than to either the African great apes or humans.     

Sivapithecus simonsi,a new species of miocene hominoid,with comments on the phylogenetic status of the ramapithecinae

The Ramapithecinae are an extinct, mainly Miocene group of hominoids, whose relationship to modern taxa is disputed. Some regard them as hominids, while others view them as ancestral toPongo,or even as the group ancestral to both hominids and extant apes. In this paper a systematic revision of Ramapithecinae is undertaken. Sivapithecus sivalensis andRamapithecus punjabicus are considered the same species, with the former name having priority. A new Indian species,Sivapithecus simonsi,is recognized. Ramapithecine anatomy is reviewed and compared with that of gracileAustralopithecus, early and middle MioceneProconsul andDryopithecus, and living pongidsPan, Gorilla, andPongo.Ramapithecines are shown to be much more primitive or “ape-like” than some have argued. Anatomical data are evaluated cladistically with several results. Parallel evolution in the jaws, teeth, and facial structure of hominoids appears to be the rule rather than the exception. Bearing this in mind, nevertheless, from the available evidence of anatomy, ramapithecines are cladistically hominids.     

A new species of Anchitherium (Equidae: Anchitheriinae) from the Middle Miocene of Abocador de Can Mata (Vallès-Penedès Basin, NE Iberian Peninsula)

Anchitherium nievei sp. nov. is erected on the basis of dentognathic remains from the late Aragonian (MN7+8, ca. 12.3–12.1 to 11.7–11.6 Ma, Middle Miocene) of three localities from the Abocador de Can Mata local stratigraphic series (els Hostalets de Pierola, Vallès-Penedès Basin, Catalonia, Spain). This taxon differs from endemic species of Anchitherium from the inner Iberian basins, most closely resembling A. aurelianense, A. hippoides and A. steinheimense by its marked brachyodonty and small dental size, although differing by dental proportions and several details of occlusal morphology. From a paleoenvironmental viewpoint, the presence of this taxon fits well with previous inferences that indicated a relatively closed and humid forested habitat, more similar to that of similarly aged French localities.     

Rudabánya: A late miocene subtropical swamp deposit with evidence of the origin of the African apes and humans

Rudabánya, a rich late Miocene fossil site in northern central Hungary, has yielded an abundant record of fossil primates, including the primitive catarrhine Anapithecus and the early great ape Dryopithecus. While the affinities of Anapithecus are not clear, Dryopithecus is clearly a great ape sharing numerous characteristics of its dental, cranial and postcranial anatomy with living great apes. Like all Miocene hominids (great apes and humans), Dryopithecus is more primitive in a number of ways than any living hominid, which is probably related to the passage of time since the divergence of the various lineages of living hominids, allowing for similar refinements in morphology and adaptation to take place independently. On the other hand, Dryopithecus (and Ouranopithecus) share derived characters with hominines (African apes and humans), and Sivapithecus (and Ankarapithecus) share derived characters with orangutans, thus dating the split between pongines and hominines to a time before the evolution of these fossil great apes. Pongines and hominines follow similar fates in the late Miocene, the pongines moving south into Southeast Asia from southern or eastern Asia and the hominines moving south into East Africa from the Mediterranean region, between 6 to 9 Ma.     

Rudabànya: Taphonomic analysis of a fossil hominid site from Hungary

The late Miocene deposits at Rudabànya, Hungary, were laid down in a shallow valley sloping westwards from a range of hills and opening out into the Pannonian Lake. Rise and fall of lake level gave rise to varying conditions, from dry land with soil formation to swamp and lake. The stratigraphic and palaeontological succession has been investigated at one of the sites, Rudabànya 2, where two cycles of deposition and erosion are represented, with soil formation, swamp conditions with lignite formation, and periods of extended high lake level succeeding each other. Both mammal and plant fossils are present at several levels. Taphonomic modifications in the Rudabànya 2 vertebrate faunas include losses of skeletal elements through carnivore selection, fluvial sorting at some levels, and post-depositional destruction by leaching and/or acid soils. The lowest level, the lower lignite, has few fossils. The fossil mammals from the level above, the grey marl, are the least modified but they are mixed with abraded, probably allochthonous, bone fragments and more complete specimens resulting from near-lake deaths. Modifications of bones by carnivores are indicated, but the specimens were too broken post-depositionally for the impact of the carnivores to be assessed. Carnivore action is also indicated for the fauna of the black clay which formed on the surface of the grey marl. The fauna consists of relatively abundant small mammals and the primates Anapithecus hernyaki and Dryopithecus hungaricus, with the latter much less common. The predator accumulating the smaller species was probably a viverrid. The red marl fauna is a transported assemblage from higher up the valley with the fossils extremely fragmentary and abraded and few identifiable specimens, almost all of which are teeth. The black mud fauna is also probably a transported assemblage, lower energy than the red marl environment, and the bones are much modified subsequently by acid corrosion similar to that seen today in bone preserved in peat bogs. Dryopithecus is a major constituent of the fauna, with Anapithecus less common. D. hungaricus is thus associated more strongly with swamp forest and shallow riverine conditions with low energy movement of water, and A. hernyaki is associated with lake shore (probably forest) conditions, accumulating in lake sediments and lake-flat sediments. The palaeoecology of the area as a whole, based on the associated flora and fauna, is a combination of swamp forest, lake shore forest and open mud flats.     

Pierolapithecus and the functional morphology of Miocene ape hand phalanges: paleobiological and evolutionary implications

The partial skeleton of Pierolapithecus, which provides the oldest unequivocal evidence of orthogrady, together with the recently described phalanges from Pa?alar most likely attributable to Griphopithecus, provide a unique opportunity for understanding the changes in hand anatomy during the pronogrady/orthogrady transition in hominoid evolution. In this paper, we describe the Pierolapithecus hand phalanges and compare their morphology and proportions with those of other Miocene apes in order to make paleobiological inferences about locomotor evolution. In particular, we investigate the orthograde/pronograde evolutionary transition in order to test whether the acquisition of vertical climbing and suspension were decoupled during evolution. Our results indicate that the manual phalanges of Miocene apes are much more similar to one another than to living apes. In particular, Miocene apes retain primitive features related to powerful-grasping palmigrady on the basal portion, the shaft, and the trochlea of the proximal phalanges. These features suggest that above-branch quadrupedalism, inherited from stem hominoids, constituted a significant component of the locomotor repertories of different hominoid lineages at least until the late Miocene. Nonetheless, despite their striking morphological similarities, several Miocene apes do significantly differ in phalangeal curvature and/or elongation. Hispanopithecus most clearly departs by displaying markedly-curved and elongated phalanges, similar to those in the most suspensory of the extant apes (hylobatids and orangutans). This feature agrees with several others that indicate orang-like suspensory capabilities. The remaining Miocene apes, on the contrary, display low to moderate phalangeal curvature, and short to moderately-elongated phalanges, which are indicative of the lack of suspensory adaptations. As such, the transition from a pronograde towards an orthograde body plan, as far as this particular anatomical region is concerned, is reflected only in somewhat more elongated phalanges, which may be functionally related to enhanced vertical-climbing capabilities. Our results therefore agree with the view that hominoid locomotor evolution largely took place in a mosaic fashion: just as taillessness antedated the acquisition of an orthograde body plan, the emergence of the latter—being apparently related only to vertical climbing—also preceded the acquisition of suspensory adaptations, as well as the loss of primitively-retained, palmigrady-related features.     

A late divergence hypothesis

The possibility of a Middle-Late Miocene separation of the human lineage from the lineages leading to the extant great apes, based on paleontological and phenetic evidence, is presented. Middle Miocene Sivapithecus, rather then Early Miocene Dryopithecus, is supported as a last common ancestor of Pongo, Pan, Gorilla, and Homo. Estimates for the branching of the lineages are a maximum of 15 m.y.a. for the Pongo lineage and a range from 14-6 m.y.a. for the Pan, Gorilla, and Ausralopithecus/Homo lineages. Weaknesses of the late divergence hypothesis are discussed.     

The phylogenetic placement of the Leucogastrales, including Mycolevis siccigleba (Cribbeaceae), in the Albatrellaceae using morphological and molecular data

The morphology of many hypogeous fungi converges on a homogeneous reduced form, suggesting that disparate lineages are subject to a uniform selection pressure. The primary goal of this study was to evaluate the morphology and infer the phylogeny of the Leucogastrales with Mycolevis siccigleba using a Bayesian methodology. A comprehensive morphological assessment was used for an a priori phylogenetic inference to guide the sequencing effort. All structures except spore ornamentation pointed to the Albatrellaceae as the most likely sister taxon. Polyporoletus sublividus, a close relative of Albatrellus, produces ornamented basidiospores with a similar structure to M. siccigleba basidiospores. The ITS from 30 taxa was used for the molecular phylogenetic analysis. P. sublividus was found sister to Mycolevis. Leucophleps spinispora and L. magnata formed a group sister to the Polyporoletus/Mycolevis group, whereas Leucogaster was polyphyletic with respect to the core of the Leucogastrales and sister to A. caeruleoporus. This relationship was expected as previously undescribed chlamydospores produced by members of Albatrellus had a similar morphology to the basidiospores of L. rubescens.     

Evolution of the second orangutan: phylogeny and biogeography of hominid origins

Aim To resolve the phylogeny of humans and their fossil relatives (collectively, hominids), orangutans (Pongo) and various Miocene great apes and to present a biogeographical model for their differentiation in space and time. Location Africa, northern Mediterranean, Asia. Methods Maximum parsimony analysis was used to assess phylogenetic relationships among living large‐bodied hominoids (= humans, chimpanzees, bonobos, gorillas, orangutans), and various related African, Asian and European ape fossils. Biogeographical characteristics were analysed for vicariant replacement, main massings and nodes. A geomorphological correlation was identified for a clade we refer to as the ‘dental hominoids’, and this correlation was used to reconstruct their historical geography. Results Our analyses support the following hypotheses: (1) the living large‐bodied hominoids represent a monophyletic group comprising two sister clades: humans + orangutans, and chimpanzees (including bonobos) + gorillas (collectively, the African apes); and (2) the human–orangutan clade (dental hominoids) includes fossil hominids (Homo, australopiths, Orrorin) and the Miocene‐age apes Hispanopithecus, Ouranopithecus, Ankarapithecus, Sivapithecus, Lufengpithecus, Khoratpithecus and Gigantopithecus (also Plio‐Pleistocene of eastern Asia). We also demonstrate that the distributions of living and fossil genera are largely vicariant, with nodes of geographical overlap or proximity between Gigantopithecus and Sivapithecus in Central Asia, and between Pongo, Gigantopithecus, Lufengpithecus and Khoratpithecus in East Asia. The main massing is represented by five genera and eight species in East Asia. The dental hominoid track is spatially correlated with the East African Rift System (EARS) and the Tethys Orogenic Collage (TOC). Main conclusions Humans and orangutans share a common ancestor that excludes the extant African apes. Molecular analyses are compromised by phenetic procedures such as alignment and are probably based on primitive retentions. We infer that the human–orangutan common ancestor had established a widespread distribution by at least 13 Ma. Vicariant differentiation resulted in the ancestors of hominids in East Africa and various primarily Miocene apes distributed between Spain and Southeast Asia (and possibly also parts of East Africa). The geographical disjunction between early hominids and Asian Pongo is attributed to local extinctions between Europe and Central Asia. The EARS and TOC correlations suggest that these geomorphological features mediated establishment of the ancestral range.     

Trabecular bone structure in the primate wrist

Trabecular (or cancellous) bone has been shown to respond to mechanical loading throughout ontogeny and thus can provide unique insight into skeletal function and locomotion in comparative studies of living and fossil mammalian morphology. Trabecular bone of the hand may be particularly functionally informative because the hand has more direct contact with the substrate compared with the remainder of the forelimb during locomotion in quadrupedal mammals. This study investigates the trabecular structure within the wrist across a sample of haplorhine primates that vary in locomotor behaviour (and thus hand use) and body size. High‐resolution microtomographic scans were collected of the lunate, scaphoid, and capitate in 41 individuals and eight genera (Homo, Gorilla, Pan, Papio, Pongo, Symphalangus, Hylobates, and Ateles). We predicted that particular trabecular parameters would 1) vary across suspensory, quadrupedal, and bipedal primates based on differences in hand use and load, and 2) scale with carpal size following similar allometric patterns found previously in other skeletal elements across a larger sample of mammals and primates. Analyses of variance (trabecular parameters analysed separately) and principal component analyses (trabecular parameters analysed together) revealed no clear functional signal in the trabecular structure of any of the three wrist bones. Instead, there was a large degree of variation within suspensory and quadrupedal locomotor groups, as well as high intrageneric variation within some taxa, particularly Pongo and Gorilla. However, as predicted, Homo sapiens, which rarely use their hands for locomotion and weight support, were unique in showing lower relative bone volume (BV/TV) compared with all other taxa. Furthermore, parameters used to quantify trabecular structure within the wrist scale with size generally following similar allometric patterns found in trabeculae of other mammalian skeletal elements. We discuss the challenges associated with quantifying and interpreting trabecular bone within the wrist. J. Morphol. 275:572–585, 2014. © 2013 Wiley Periodicals, Inc.     

Orycteropus (Tubulidentata, Mammalia) from Langebaanweg and Baard's Quarry, Early Pliocene of South Africa

Fossil teeth and bones of aardvarks are relatively common at Langebaanweg, an Early Pliocene site in western Cape Province, South Africa. The remains are compatible in size and most details of morphology to extant Orycteropus afer, and are the earliest fossils attributed to this species. Other Late Miocene to Early Pliocene localities in Africa have yielded smaller species of aardvarks, suggesting that the extant lineage evolved in southern Africa. Morphologically the genus Orycteropus has been remarkably conservative since at least the Early Miocene but it witnessed an overall increase in size through the Neogene. The species O. afer has been morphometrically stable since the Early Pliocene. These observations indicate that the evolutionary process in aardvarks is extremely bradytelic. To cite this article: M. Pickford, C. R. Palevol 4 (2005).     

Notes on the floral morphology in Vivianiaceae (Geraniales)

The functional floral morphology of the three genera of Vivianiaceae (= Ledocarpaceae, Geraniales), Rhynchotheca, Viviania and Balbisia, is compared. Likely pollination mechanisms are inferred from morphology and field observations. The flowers of Viviania are nectariferous and apparently zoophilous with nectar as the (primary) pollinator reward. Balbisia has pollen flowers without nectaries, its showy corolla indicates that it is also zoophilous with pollen as sole pollinator reward; bees were observed as flower visitors. One taxon (B. gracilis) may be anemophilous. Rhynchotheca has flowers without petals, with large, pendulous anthers and lacks nectaries. It shows synchronous mass flowering in its natural populations and is evidently anemophilous. A comparison with other Geraniales shows that nectar flowers with small anthers are likely the ancestral condition in Vivianiaceae. This suggests that the pollen flowers with larger anthers of Balbisia and Rhynchotheca may represent an apomorphic condition. The documentation of pollen flowers and anemophily in Vivianiaceae expands the range of known floral and pollination syndromes in Geraniales.     

Evolution of the mandibular third premolar crown in early Australopithecus

The Pliocene hominins Australopithecus anamensis and Australopithecus afarensis likely represent ancestor-descendent taxa—possibly an anagenetic lineage—and capture significant change in the morphology of the canine and mandibular third premolar (P₃) crowns, dental elements that form the canine honing complex in nonhuman catarrhines. This study focuses on the P₃ crown, highlighting plesiomorphic features in A. anamensis. The A. afarensis P₃ crown, in contrast, is variable in its expression of apomorphic features that are characteristic of geologically younger hominins. Temporal variation characterizes each taxon as well. The A. anamensis P₃ from Allia Bay, Kenya expresses apomorphic character states, shared with A. afarensis, which are not seen in the older sample of A. anamensis P₃s from Kanapoi, Kenya, while spatiotemporal differences in shape exist within the A. afarensis hypodigm. The accumulation of derived features in A. afarensis results in an increased level of P₃ molarisation. P₃ molarisation did not evolve concurrent with postcanine megadontia and neither did the appearance of derived aspects of P₃ occlusal form coincide with the loss of canine honing in hominins, which is apparent prior to the origin of the genus Australopithecus. A. afarensis P₃ variation reveals the independence of shape, size, and occlusal form. The evolution of the P₃ crown in early Australopithecus bridges the wide morphological gap that exists between geologically younger hominins on the one hand and extant apes and Ardipithecus on the other.