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OBJECTIVES One of the most contentious issues in paleoanthropology is the nature of the last common ancestor of humans and our closest living relatives, chimpanzees and bonobos (panins). The numerical composition of the vertebral column has featured prominently, with multiple models predicting distinct patterns of evolution and contexts from which bipedalism evolved. Here, we study total numbers of vertebrae from a large sample of hominoids to quantify variation in and patterns of regional and total numbers of vertebrae in hominoids. MATERIALS AND METHODS We compile and study a large sample (N = 893) of hominoid vertebral formulae (numbers of cervical, thoracic, lumbar, sacral, caudal segments in each specimen) and analyze full vertebral formulae, total numbers of vertebrae, and super-regional numbers of vertebrae: presacral (cervical, thoracic, lumbar) vertebrae and sacrococcygeal vertebrae. We quantify within- and between-taxon variation using heterogeneity and similarity measures derived from population genetics. RESULTS We find that humans are most similar to African apes in total and super-regional numbers of vertebrae. Additionally, our analyses demonstrate that selection for bipedalism reduced variation in numbers of vertebrae relative to other hominoids. DISCUSSION The only proposed ancestral vertebral configuration for the last common ancestor of hominins and panins that is consistent with our results is the modal formula demonstrated by chimpanzees and bonobos (7 cervical-13 thoracic-4 lumbar-6 sacral-3 coccygeal). Hox gene expression boundaries suggest that a rostral shift in Hox10/Hox11-mediated complexes could produce the human modal formula from the proposal ancestral and panin modal formula.

Vertebral formulae, the combination of regional numbers of vertebrae making up the bony spine, vary across vertebrates and within hominoid primates. Reconstructing the ancestral vertebral formulae throughout hominoid evolution has proved a challenge due to limited fossil evidence and disagreement among researchers. Proposed “long-backed” and “short-backed” ancestors have implications for the evolution of bipedalism and human evolutionary history generally. Here, we analyze a large dataset of hominoid vertebral formulae, including previously unstudied species and subspecies. We find more variation within and between species than expected, particularly in hylobatids (gibbons or lesser apes) and in gorilla and chimpanzee subspecies. Our results suggest that combined thoracic and lumbar numbers of vertebrae are somewhat phylogenetically structured, with outgroup taxa (two species of Old World monkeys, or cercopithecoids) retaining the primitive number of 19 thoracolumbar vertebrae, hylobatids generally possessing 18 thoracolumbar vertebrae, and hominids (great apes and humans) having 17 or 16 thoracolumbar vertebrae. When compared to cercopithecoids, and to putative stem hominoids, extant hominoids show evidence for homeotic change at both the lumbosacral (e.g., decrease in lumbar vertebrae; increase in sacral segments) and in the position of the transitional vertebrae. Homeotic changes are probably also responsible for the differences between African apes and modern humans, with differences in the number of thoracic and lumbar within a 17-segment thoracolumbar framework.

Continental gateways occur where mountainous topography interacts with changing climate and sea level to open or close dispersal corridors. The interaction of permeable or impermeable montane barriers with changing or stable climate yields four biogeographic states, each associated with changes in diversification rates and ecological structure of faunas. For example, permeable montane barriers and climatic stability result in low rates of immigration and extinction, elevated endemic speciation, and stable ecological structure. Three examples from the mammalian fossil record test these scenarios. (1) In Miocene faunas of Pakistan, immigration rates peaked and faunal proportions changed during an interval of cooling and open corridors. (2) In Miocene faunas of Spain, elevated extinction and origination rates and changing trophic structure occurred during regional aridification with open corridors. (3) In Quaternary faunas of South Africa, ungulates experienced range reductions and elevated extinction during the transition from glacial to interglacial climates as corridors closed.

The Tertiary sediments of northern Pakistan are an exceptional record of terrestrial sedimentation and represent most of Neogene time. Foremost is the Siwalik Group of the Potwar Plateau, for which multiple, superposed fossil levels span ∼18–6 Ma. Well-developed magnetostratigraphic control provides secure dating so that Siwalik fossil horizons may be interpolated into a time scale with resolution to 100,000 years. We describe the geographic setting of the Potwar, give an overview of the temporal distribution of faunas, and discuss changes in paleohabitat and paleoecology with coinciding faunal change, as seen from the Siwalik viewpoint. The long Siwalik biostratigraphy of many successive assemblages with its resolved time scale may be compared directly with other well-dated sequences. Immigrant arrival and timing of faunal change may be traced. The basins of the Iberian Peninsula show somewhat different timing of introduction of hipparionine horses, and faunal turnover in the Siwaliks clearly precedes the Vallesian crisis in Spain. In contrast to the increasingly seasonal precipitation of the late Miocene Potwar, the paleohabitat of coeval North China appears to have been moist and equable, with high diversity faunas. Continued development and comparison of resolved Neogene records allow increasing resolution of the patterns of faunal change on regional to global levels.

We describe a partial innominate, YGSP 41216, from a 12.3 Ma locality in the Siwalik Group of the Potwar Plateau in Pakistan, assigned to the Middle Miocene ape species Sivapithecus indicus. We investigate the implications of its morphology for reconstructing positional behavior of this ape. Postcranial anatomy of extant catarrhines falls into two distinct groups, particularly for torso shape. To an extent this reflects different although variable and overlapping positional repertoires: pronograde quadrupedalism for cercopithecoids and orthogrady for hominoids. The YGSP innominate (hipbone) is from a primate with a narrow torso, resembling most extant monkeys and differing from the broader torsos of extant apes. Other postcranial material of S. indicus and its younger and similar congener Sivapithecus sivalensis also supports reconstruction of a hominoid with a positional repertoire more similar to the pronograde quadrupedal patterns of most monkeys than to the orthograde patterns of apes. However, Sivapithecus postcranial morphology differs in many details from any extant species. We reconstruct a slow-moving, deliberate, arboreal animal, primarily traveling above supports but also frequently engaging in antipronograde behaviors. There are no obvious synapomorphic postcranial features shared exclusively with any extant crown hominid, including Pongo.

Faunal assemblages consist of immigrants, endemics and long-term residents; changes in proportions of these categories through time reflect general aspects of faunal stability and turnover. To study stability and change in the long biostratigraphic succession of Miocene terrestrial mammalian faunas recorded mainly in the Potwar Plateau Siwaliks of Pakistan, we distinguish long-term residents and assess new appearances as endemic vs. immigrant components. Our data represent the biogeography of the northern Indian subcontinent, but because neighboring areas of similar age are not as well known, virtually all species are found only there, and the proportion of the fauna that is immigrant is not apparent. We use preceding faunal assemblages to judge whether individual species might have originated there. We find many long-term residents in the Siwaliks, and of new species, most are immigrants and fewer are likely to have originated endemically. Siwalik rodents and artiodactyls show high rates of immigration f...

Introduction, Allen V. Barker and David J. Pilbeam Essential Elements-Macronutrients Nitrogen, Allen V. Barker and Gretchen M. Bryson Phosphorus, Charles A. Sanchez Potassium, Konrad Mengel Calcium, David J. Pilbeam and Philip S. Morley Magnesium, Donald J. Merhaut Sulfur, Silvia Haneklaus, Elke Bloem, Ewald Schnug, Luit J. de Kok and Ineke Stulen Essential Elements-Micronutrients Boron, Umesh C. Gupta Chlorine, Joseph R. Heckman Copper, David E. Kopsel and Dean A. Kopsell Iron, Volker Roemheld and Miroslav Nikolic Manganese, Julia M. Humphries, James C.R. Stangoulis, and Robin D. Graham Molybdenum, Russell L. Hamlin Nickel, Patrick H. Brown Zinc, J. Benton Storey Beneficial Elements Aluminum, Susan C. Miyasaka, N.V. Hue, and Michael A. Dunn Cobalt, Geeta Talukder and Archana Sharmaz Selenium, Dean A. Kopsell and David E. Kopsell Silicon, George H. Snyder, Vladimir V. Matichenkov and Lawrence E. Datnoff Sodium, John Gorham Vanadium, David J Pilbeam and K Drihem Conclusion, Allen V. Barker and David J Pilbeam

An extensive suite of isotopic data (δ 13 C, δ 15 N, and δ 18 O) from enamel apatite and bone collagen of adult male and female wild chimpanzees establishes baseline values for Pan troglodytes verus in a primary rainforest setting. The Ganta chimpanzee sample derives from a restricted region in northern Liberia. Diet is examined using stable light isotopes at three life stages—infant, young juvenile, and adult—and developmental differences are investigated within and between individual males and females. The isotopic data are very homogeneous with few exceptions. Juvenile females show consistent enrichment in 13 C relative to infants, while juvenile males do not. These data suggest that age at weaning may be more variable for male offspring who survive to adulthood than for female offspring. Alternatively, or additionally, the weaning diet of males and females may differ, with greater consumption of technologically extracted insects and/or nuts by young females. Metabolic differences, including growth and hormone-mediated responses, may also contribute to the observed variation. The Ganta chimpanzee data offer an independent and objective line of evidence to primatologists interested in the dietary strategies of the great apes and to paleoanthropologists seeking comparative models for reconstructing early hominin subsistence patterns. Despite the high diversity of dietary items consumed by chimpanzees, isotopic signatures of chimpanzees from a primary rainforest setting exhibit narrow ranges of variation similar to chimpanzees in more open habitats.

Isotopic analyses of mammalian tooth enamel from a well-defined, laterally extensive 150 k.y. interval (9.15–9.30 Ma) reveal an ecological gradient in vegetation on the late Miocene sub-Himalayan alluvial plain. Two contemporaneous river systems deposited the sediments of this interval, with a mountain-sourced system (herein, Blue-gray) to the southwest interfingering with a foothill-sourced system (Buff) to the northeast. Fossil mammal teeth collected from a 32 km transect across this fluvial gradient are significantly more depleted in 13C from northeastern localities than from southwestern localities. This trend occurs in equids, giraffids, suids, sivapithecine hominoids, and anthracotheres. We propose that the Buff fluvial system provided more equably moist substrate conditions and supported more closed-canopy vegetation than the Blue-gray fluvial system. Herbivores living along the paleovegetation gradient thus acquired different carbon isotopic signatures during the period of tooth enamel formation, resulting from higher δ13C values in the forage supported by the Blue-gray fluvial system compared with forage associated with the Buff system. The data also imply that many Siwalik mammalian herbivores displayed marked fidelity in juvenile home ranges and habitats.

The discovery of the skull known as Toumai four years ago in Chad began a controversy. Faunal studies suggested an age close to 7 million years; a small cranium suggested chimpanzee-like brain size. The team that found Toumai considered it to be a hominid on our side of the chimp-human divide, but others thought it more ape-like. Important finds of teeth and jaw pieces of the Toumai species, Sahelanthropus tchadensis, now help to distance the species from apes, suggesting that it is a hominid closely related to the last common ancestor of chimps and humans. A virtual reconstruction of the Toumai cranium provides more evidence of a close relationship to humans — and this week's cover. You are looking at the face of the earliest known hominid. (Cover by MPFT; M. Brunet, E. Daynes, Ph. Plailly and A. Garaudel contributed). Discoveries in Chad by the Mission Paleoanthropologique Franco-Tchadienne have substantially changed our understanding of early human evolution in Africa1,2,3. In particular, the TM 266 locality in the Toros-Menalla fossiliferous area yielded a nearly complete cranium (TM 266-01-60-1), a mandible, and several isolated teeth assigned to Sahelanthropus tchadensis3 and biochronologically dated to the late Miocene epoch (about 7 million years ago). Despite the relative completeness of the TM 266 cranium, there has been some controversy about its morphology and its status in the hominid clade4,5. Here we describe new dental and mandibular specimens from three Toros-Menalla (Chad) fossiliferous localities (TM 247, TM 266 and TM 292) of the same age6. This new material, including a lower canine consistent with a non-honing C/P3 complex, post-canine teeth with primitive root morphology and intermediate radial enamel thickness, is attributed to S. tchadensis. It expands the hypodigm of the species and provides additional anatomical characters that confirm the morphological differences between S. tchadensis and African apes. S. tchadensis presents several key derived features consistent with its position in the hominid clade close to the last common ancestor of chimpanzees and humans.

Within-species phenotypic variation is the raw material on which natural selection acts to shape evolutionary change, and understanding more about the developmental genetics of intraspecific as well as interspecific phenotypic variation is an important component of the Evo-Devo agenda. The axial skeleton is a useful system to analyze from such a perspective. Its development is increasingly well understood, and between-species differences in functionally important developmental parameters are well documented. I present data on intraspecific variation in the axial postcranial skeleton of some Primates, including hominoids (apes and humans). Hominoid species are particularly valuable, because counts of total numbers of vertebrae, and hence original somite numbers, are available for large samples. Evolutionary changes in the axial skeleton of various primate lineages, including bipedal humans, are reviewed, and hypotheses presented to explain the changes in terms of developmental genetics. Further relevant experiments on model organisms are suggested in order to explore more fully the differences in developmental processes between primate species, and hence to test these hypotheses.

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.

The Siwalik formations of northern Pakistan consist of deposits of ancient rivers that existed throughout the early Miocene through the late Pliocene. The formations are highly fossil- iferous with a diverse array of terrestrial and freshwater vertebrates, which in combination with exceptional lateral exposure and good chronostratigraphic control allows a more detailed and tem- porally resolved study of the sediments and faunas than is typical in terrestrial deposits. Conse- quently the Siwaliks provide an opportunity to document temporal differences in species richness, turnover, and ecological structure in a terrestrial setting, and to investigate how such differences are related to changes in the fluvial system, vegetation, and climate. Here we focus on the interval between 10.7 and 5.7 Ma, a time of significant local tectonic and global climatic change. It is also the interval with the best temporal calibration of Siwalik faunas and most comprehensive data on species occurrences. A methodological focus of this paper is on controlling sampling biases that confound biological and ecological signals. Such biases include uneven sampling through time, differential preservation of larger animals and more durable skeletal elements, errors in age-dating imposed by uncertainties in correlation and paleomagnetic timescale calibrations, and uneven tax- onomic treatment across groups. We attempt to control for them primarily by using a relative-abun- dance model to estimate limits for the first and last appearances from the occurrence data. This model also incorporates uncertainties in age estimates. Because of sampling limitations inherent in the terrestrial fossil record, our 100-Kyr temporal resolution may approach the finest possible level of resolution for studies of vertebrate faunal changes over periods of millions of years. Approximately 40,000 specimens from surface and screenwash collections made at 555 localities form the basis of our study. Sixty percent of the localities have maximum and minimum age esti- mates differing by 100 Kyr or less, 82% by 200 Kyr or less. The fossils represent 115 mammalian species or lineages of ten orders: Insectivora, Scandentia, Primates, Tubulidentata, Proboscidea, Pholidota, Lagomorpha, Perissodactyla, Artiodactyla, and Rodentia. Important taxa omitted from this study include Carnivora, Elephantoidea, and Rhinocerotidae. Because different collecting methods were used for large and small species, they are treated separately in analyses. Small spe- cies include insectivores, tree shrews, rodents, lagomorphs, and small primates. They generally weigh less than 5 kg. The sediments of the study interval were deposited by coexisting fluvial systems, with the larger emergent Nagri system being displaced between 10.1 and 9.0 Ma by an interfan Dhok Pathan sys- tem. In comparison to Nagri floodplains, Dhok Pathan floodplains were less well drained, with smaller rivers having more seasonally variable flow and more frequent avulsions. Paleosol se- quences indicate reorganization of topography and drainage accompanying a transition to a more seasonal climate. A few paleosols may have formed under waterlogged, grassy woodlands, but most formed under drier conditions and more closed vegetation. The oxygen isotopic record also indicates significant change in the patterns of precipitation be- ginning at 9.2 Ma, in what may have been a shift to a drier and more seasonal climate. The carbon isotope record demonstrates that after 8.1 Ma significant amounts of C 4 grasses began to appear and that by 6.8 Ma floodplain habitats included extensive C4 grasslands. Plant communities with predominantly C3 plants were greatly diminished after 7.0 Ma, and those with predominantly C4 plants, which would have been open woodlands or grassy woodlands, appeared as early as 7.4 Ma. Inferred first and last appearances show a constant, low level of faunal turnover throughout the interval 10.7-5.7-Ma, with three short periods of elevated turnover at 10.3, 7.8, and 7.3-7.0 Ma. The three pulses account for nearly 44% of all turnover. Throughout the late Miocene, species richness declined steadily, and diversity and richness indices together with data on body size imply that community ecological structure changed abruptly just after 10 Ma, and then again at 7.8 Ma. Be- tween 10 and 7.8 Ma the large-mammal assemblages were strongly dominated by equids, with more balanced faunas before and after. The pattern of appearance and disappearance is selective with respect to inferred habits of the animals. Species appearing after 9.0 Ma are grazers or typical of more open habitats, whereas many species that disappear can be linked to more closed vege- tation. We presume exceptions to this pattern were animals of the mixed C

The large-bodied hominoid from Moroto, Uganda has until recently been known only from proconsulid like craniodental remains and some vertebrae with modern ape like features. The discovery of two partial femora and the glenoid portion of a scapula demonstrates that the functional anatomy of Morotopithecus differed markedly from other early and middle Miocene hominoids. Previous studies have consistently associated the vertebral remains with a short, stiff back and with orthograde postures. Although the proximal femur more closely resembles the femora of monkeys than of apes and suggests a moderate degree of hip abduction, the distal femur resembles those of extant large bodied apes and suggests a varied loading regime and an arboreal repertoire that may have included substantial vertical climbing. The femoral shaft displays uniformly thick cortical bone, beyond the range of thickness seen in extant primates, and signifies higher axial loading than is typical of most extant primates. The glenoid fossa is broad and uniformly curved as in extant suspensory primates. Overall, Morotopithecus is reconstructed as an arboreal species that probably relied on forelimb-dominated, deliberate and vertical climbing, suspension and quadrupedalism. Morotopithecus thus marks the first appearance of certain aspects of the modern hominoid body plan by at least 20 Ma. If the suspensory and orthograde adaptations linking Morotopithecus to extant apes are synapomorphies, Morotopithecus may be the only well-documented African Miocene hominoid with a close relationship to living apes and humans.

Cooking is a human universal that must have had widespread effects on the nutrition, ecology, and social relationships of the species that invented it. The location and timing of its origins are unknown, but it should have left strong signals in the fossil record. We suggest that such signals are detectable at ca. 1.9 million years ago in the reduced digestive effort (e.g., smaller teeth) and increased supply of food energy (e.g., larger female body mass) of early Homo erectus. The adoption of cooking required delay of the consumption of food while it was accumulated and/or brought to a processing area, and accumulations of food were valuable and stealable. Dominant (e.g., larger) individuals (typically male) were therefore able to scrounge from subordinate (e.g., smaller) individuals (typically female) instead of relying on their own foraging efforts. Because female fitness is limited by access to resources (particularly energetic resources), this dynamic would have favored females able to minimize losse...

Fossils from a large-bodied hominoid from early Miocene sediments of Uganda, along with material recovered in the 1960s, show features of the shoulder and vertebral column that are significantly similar to those of living apes and humans. The large-bodied hominoid from Uganda dates to at least 20.6 million years ago and thus represents the oldest known hominoid sharing these derived characters with living apes and humans.

Les inferences a propos du comportement, de l'ecologie et de la vie des hominides ont souvent ete realisees par analogie entre les especes vivantes et les especes fossiles. L'A estime que cette methode est erronee pour trois raisons : l'analogie n'est pas realisee a partir d'especes vivantes, les intervales de confidence pour des valeurs predites font plus d'inferences que les usages minimes et sans pratique, la reduction des variations biologiques des especes fossiles a un determinisme des especes fossiles est statistiquement invalide, est une erreur biologique

Dividing the fossils usually assigned to the taxonHomo habilis sensu latointo two species (as most researchers now accept) necessitates a re-examination of their evolutionary relationships. A cladistic analysis of 48 of the most commonly-used cranial characters from recent studies of Pliocene hominid phylogeny and which distinguish two taxa withinH. habilis sensu latosuggests that these fossils have different evolutionary affinities. One taxon,H. habilis sensu stricto, is represented by KNM-ER 1813 and the fossils from Olduvai Gorge, and is most likely a sister group ofH. erectus. The other taxon,H. rudolfensis, is represented by KNM-ER 1470, and shares many derived characters with the australopithecines. A close analysis of the developmental basis of these characters suggests that many of the australopithecine similarities ofH. rudolfensisare likely to be homologies rather than homoplasies.

Molecular genetics has had a major impact on phylogenetics, although many hominoid paleontologists and morphologists ignore or remain unaware of genetic data. However, substantial genetic evidence shows chimpanzees and humans as closest relatives. Living hominoids share many postcranial similarities, many of which are retained from the extant hominoid common ancestor. Miocene hominoid fossils consisted until recently mostly of teeth and jaw fragments which are relatively uninformative phylogenetically. As their postcrania have become better sampled, surprisingly few of these taxa share significant similarities with living apes, suggesting that few if any are related to specific extant lineages. Given the genetically inferred relationships of hominoids and the morphology of the earliest hominids, the common ancestor of humans and chimpanzees was probably chimp-like, a knuckle-walker with small thin-enameled cheek teeth. If correct, this scenario implies that known Miocene hominoids, most of which are postcranially archaic and have large, thickly enameled cheek teeth, throw little if any direct light on hominid origins.

The fluvial Neogene Siwalik formations of northern Pakistan contain a long and richly fossiliferous sequence of terrestrial vertebrate faunas in which patterns of faunal turnover and changes in diversity can be documented and analyzed for intervals having durations of 0.5 m.y. The complete sequence extends from circa 18.5 to 1 Ma, but the part between 18.5 and 5.5 Ma is best sampled, and most intervals within it are well represented. Thirteen orders of Siwalik mammals have been identified, with well-sampled intervals having 50 or more species. Most Siwalik mammals, however, are either rodents or artiodactyls. Bovids are the most common and most speciose of the larger mammals, while murid and “cricetid” rodents dominate the small mammal assemblages. Between 18.5 and 5.5 Ma species diversity varied considerably. Among artiodactyls and rodents the number of species first increased between 15 and 13 Ma and then fell. Data on stratigraphic ranges of rodents and artiodactyls show that faunal change in the Siwaliks was episodic, occurring during short intervals with high turnover followed by longer periods with considerably less change. Maxima of first appearances occurred at approximately 13.5 and 8.5 Ma, while maxima of last occurrences were at 12.5 and 8.0 Ma. Some of the observed faunal events can be correlated to climatic and environmental changes. The Middle Miocene diversification occurred during a period of global cooling, while the latest Miocene decline in diversity and increased turnover accompanied oxygen and carbon isotopic changes that correlate to globally increasing seasonality and aridity. Other correlations are ambiguous. The marked decrease in diversity and the major turnover events between 13 and 8 Ma do not correspond to known local or global events. The Neogene Siwaliks and Paleogene Bighorn-Crazy Mountains sequence in Wyoming and Montana share many similarities. They have equivalent levels of temporal resolution and similar levels of completeness of their fossil records. Siwalik ordinal abundance and diversity patterns differ markedly from those of the Paleogene, but generic, and probably species, diversity was approximately the same, although the Siwalik faunas may have been slightly less diverse. Over time, changes in diversity were of comparable magnitude, with monotonic trends persisting for more than 5 million years. The magnitude of faunal turnover was also similar, ranging from less than half to 3.5 times that expected. In both sequences faunal change appears to have been episodic, with strong pulses between intervals of low turnover. The Siwaliks, in contrast to the Paleogene sequence, may have had more distinct pulses and longer intervals between pulses. Neither sequence has peaks of first occurrences coinciding with peaks of last occurrences.

The Siwalik sequence, particularly the interval from 18 to 7 Ma, provides one of the few terrestrial data sets that allows direct measurement of temporal durations of mammalian species. Its data are drawn from a single biogeographic subprovince and superposed collections likely represent successive samples of single lineages. Observed temporal ranges underestimate total species longevities if (1) species existed in other biogeographic provinces before or after the temporal ranges recorded in the Siwaliks, or (2) the fossil record inadequately samples species durations in the Siwalik subprovince. Some data, notably from Afghanistan, China, and Thailand, bear on the first variable. The second can be controlled by considering data quality, in this case the temporal distribution of good data sets, to assess the scale of accuracy available for defining range endpoints. In general, range endpoints can be estimated to the nearest 0.1 million years.The diverse Rodentia give a mean species longevity of 2.2 million years for the Miocene Siwaliks. This includes single records, but of course ignores unretrieved rare or short-lived taxa. The diverse Artiodactyla yield 3.1 million years. The difference may reflect greater body size and longer generation time; large Perissodactyla and Proboscidea have longer temporal ranges. Carnivorous mammals also show about 3 million year durations. Given these data, the average longevity for Sivapithecus species (1.6 million years) is modest. The deposits of the Clarks Fork Basin, Wyoming, offer a Paleogene data set comparable to that of the Neogene Siwaliks. Paleocene-Eocene mammals of North America yield shorter longevities (most less than one million years).Extinction is the dominant mode of species termination for Siwalik mammals. Most taxa originated by immigration (as at about 13.5 Ma) or abrupt speciation. There are some cases for insitu transformation of lineages, for example in the genera Punjabemys, Antemus, Percrocuta, Dorcatherium, Giraffokeryx, and Selenoportax. The rodent Kanisamys shows a rate of increase in tooth size of 0.5 darwins. This overall rate is moderate by Paleogene standards, but includes an interval of more rapid change between 9.0 and 8.5 Ma.

This short reply summarizes the concerns of the anthropological community about Ashraf and Galor's (Forthcoming) article in the American Economic Review.

The Siwalik Sequence of northern Pakistan contains a 16-Ma record of paleosol carbonate and fossil teeth from which a record of paleovegetation can potentially be reconstructed and compared. The carbon isotopic composition of paleosol carbonate and organic matter from Siwalik strata reflects a major paleoecological change on the floodplains of major rivers beginning7.3 Ma ago. By 6 Ma C 3 -dominated plant communities, probably composed of mostly trees and shrubs, were displaced by nearly continuous C 4 grassland. We find that the carbon isotopic ratios in herbivore tooth enamel reflect this dramatic ecologic shift. Carbonate in enamel older than 7 Ma averages −11‰ in δ 13 C PDB , consistent with a largely C 3 diet. Enamel from the Plio-PIeistocene averages +1.9‰ in δ 13 C, similar to the value displayed by modern C 4 grazers. Analysis of post-burial carbonate cements, and the concordance with isotopic evidence from paleosols argues strongly against major isotopic alteration of the enamel, while coexisting bone may have been altered early in burial. This study confirms that enamel apatite is useful for paleodietary reconstruction much further back in the geologic record than was previously thought.

Until quite recently most evolutionary relationships were inferred from the morphology of hard tissues (bones and teeth) and soft tissues (including muscles, blood vessels, nerves, integument), by using the so-called cladistic approach developed by Hennig (1). In this approach the phenotype is described as a series of characters, which exist in two or more states. These can be of a presence/absence sort or can form a graded series. Character states are categorized as earlier …

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.

The magnetostratigraphy of the Chinji Formation (Siwalik Group, Pakistan) provides age estimates for the earliest occurrence of hominoids attributable toSivapithecus. This report documents a new earliest occurrence for the genus and provides age estimates for other Chinji Formation specimens described previously. These middle Miocene primates have figured prominently in discussions of the course and timing of hominoid evolution.Sivapithecus shares with the living orang-utan (Pongo) a number of derived craniofacial features indicating that they are sister taxa. The newly discovered specimens from the Chinji Formation therefore provide a minimum estimate for the time of the initial divergence between the Asian and African large hominoid clades. Large-bodied hominoids first appear in the Siwaliks at 12·5 m.y.a., with specimens preserving the derived craniofacial features dated at ∼ 12 m.y.a. This age estimate can be used to calibrate branching times of extant hominoids as determined from comparative molecular data.

Problems of stratigraphic completeness and poor temporal resolution make analysis of faunal change in terrestrial sequences difficult. The fluvial Neogene Siwalik formations of India and Pakistan are an exception. They contain a long vertebrate record and have good chronostratigraphic control, making it possible to assess the influence of biotic interchange on Siwalik fossil communities. In Pakistan, the interval between 18 and 7 Ma has been most intensively studied and changes in diversity and relative abundance of ruminant artiodactyls and muroid rodents are documented with temporal resolution of 200,000 years. Within this interval, diversity varies considerably, including an abrupt rise in species number between 15 and 13 Ma, followed by a decline in ruminant diversity after 12 Ma and a decline in muroid diversity in two steps at 13 and 10 Ma. Significant changes in relative abundance of taxa include an increase in bovids between 16.5 and 15 Ma, a decrease in tragulids after 9 Ma, and a very abrupt increase in murids at 12 Ma. Megacricetodontine rodents also decrease significantly at 12 Ma, and smaller declines are recorded among myocricetodontine and copemyine rodents after 16 Ma. An increase of dendromurine rodents at 15.5 Ma is also observed. There is also a trend of progressive size increase among giraffoids and bovids throughout the sequence.We have also investigated relationships between biotic interchange and diversity, body size, and relative abundance, concluding that (1) the rapid increase in ruminant and muroid diversity was largely due to immigration, whereas in situ speciation had only a secondary role; (2) during intervals of increasing diversity, resident lineages did not have higher than average rates of in situ speciation; (3) during intervals with rising diversity, greater extinction did not accompany increased immigration; (4) during intervals with falling diversity, there may have been greater extinction in recently invading lineages; and (5) change in diversity was independent of changes in relative abundance and body size.

Geohistorical records reveal the long-term impacts of climate change on ecosystem structure. A 5-myr record of mammalian faunas from floodplain ecosystems of South Asia shows substantial change in species richness and ecological structure in relation to vegetation change as documented by stable isotopes of C and O from paleosols. Between 8.5 and 6.0 Ma, C 4 savannah replaced C 3 forest and woodland. Isotopic historical trends for 27 mammalian herbivore species, in combination with ecomorphological data from teeth, show three patterns of response. Most forest frugivores and browsers maintained their dietary habits and disappeared. Other herbivores altered their dietary habits to include increasing amounts of C 4 plants and persisted for >1 myr during the vegetation transition. The few lineages that persisted through the vegetation transition show isotopic enrichment of δ 13 C values over time. These results are evidence for long-term climatic forcing of vegetation structure and mammalian ecological diversity at the subcontinental scale.

Magnetostratigraphy provides unparalleled chronologic precision for long terrestrial sedimentary records. The Siwaliks of the Potwar Plateau, northern Pakistan, present perhaps the best example of a sequence of well dated faunal horizons spanning most of the Neogene in a single terrestrial biogeographic province. Temporal control on fossil localities tied to multiple paleomagnetic sections is ca. 0.1 m.y., less given special conditions of short magnetozone duration or superpositional relationships. New correlations are presented for two long sections in the lower Siwaliks showing the power and the limitations of mag-netostratigraphic correlation within the Siwalik depositional setting. Correlation of Potwar fossil localities to those of an East African section, also dated paleomagnetically, are established with fossil localities from those two biogeographic provinces shown to be coeval on a scale of $$10^{5}$$ yr. The composite biostratigraphy rendered by the thick, well exposed, well dated Potwar Siwalik...

The recent reconstruction of the Sahelanthropus tchadensis cranium (TM 266-01-60-1) provides an opportunity to examine in detail differences in cranial shape between this earliest-known hominid, African apes, and other hominid taxa. Here we compare the reconstruction of TM 266-01-60-1 with crania of African apes, humans, and several Pliocene hominids. The results not only confirm that TM 266-01-60-1 is a hominid but also reveal a unique mosaic of characters. The TM 266-01-60-1 reconstruction shares many primitive features with chimpanzees but overall is most similar to Australopithecus , particularly in the basicranium. However, TM 266-01-60-1 is distinctive in having the combination of a short subnasal region associated with a vertical upper face that projects substantially in front of the neurocranium. Further research is needed to determine the evolutionary relationships between Sahelanthropus and the known Miocene and Pliocene hominids.