The earliest ossicone and post-cranial record of Giraffa.

The oldest Giraffa material presently known consists of dental specimens. The oldest post-cranial Giraffa material belongs to the Plio-Pleistocene taxon Giraffa sivalensis, where the holotype is a third cervical vertebra. We describe three non-dental specimens from the Early Late Miocene of the Potwar Plateau, including an 8.1 million year old ossicone, 9.4 million year old astragalus, and 8.9 million year old metatarsal and refer them to Giraffa. The described ossicone exhibits remarkable similarities with the ossicones of a juvenile modern giraffe, including the distribution of secondary bone growth, posterior curvature, and concave pitted undersurface where the ossicone would attach to the skull. The astragalus has a notably flat grove of the trochlea, medial twisting between the trochlea and the head, and a square-shaped sustentacular facet, all of which characterize the astragalus of Giraffa camelopardalis. The newly described astragalus is narrow and rectangular, unlike the boxy shaped bone of the modern giraffe. The metatarsal is large in size and has a shallow central trough created by thin medial and lateral ridges, a feature unique to Giraffa and Sivatherium. Our described material introduce the earliest non-dental material of Giraffa, a genus whose extinct representation is otherwise dominated by teeth, and demonstrate that the genus has been morphologically consistent over 9 million years.

A partial hominoid innominate from the Miocene of Pakistan: description and preliminary analyses.

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.

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.

Ecological changes in Miocene mammalian record show impact of prolonged climatic forcing.

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 delta(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.

High resolution transmission electron microscopy of developing enamel in the Australian lungfish, Neoceratodus forsteri (Osteichthyes: Dipnoi).

The permanent tooth plates of the Australian lungfish, Neoceratodus forsteri, are covered by enamel that develops initially in a similar manner to that of other vertebrates. As the enamel layer matures, it acquires several unusual characteristics. It has radially oriented protoprismatic structures with the long axes of the protoprisms perpendicular to the enamel surface. Protoprisms can be defined as aggregations of hydroxyapatite crystals that lack the highly ordered arrangement of the rods of mammalian enamel but are not without a specific structure of their own. The protoprisms are arranged in layers of variable thickness that are deposited sequentially as the tooth plate grows. They may be confined to the separate layers, or may cross the boundary between each layer. Crystals within the protoprisms are long and thin with hydroxyapatite c-axis dimensions of between 30 and 350 nm, and with typical a-b axis dimensions of 5-10 nm. The hydroxyapatite crystals of lungfish enamel have no centre dark lines of octacalcium phosphate, an unusual character among vertebrates. As each crystal develops, arrays of atoms may change direction, and regions exist where dislocations and extra lattice planes are inserted into the long crystal. The resulting hydroxyapatite crystal is not straight, and has a rough surface. The crystals are arranged in tangled structures with their crystallographic c-axes closely aligned with the long axis of the protoprism. Lungfish enamel differs from the enamel of higher vertebrates in that the hydroxyapatite crystals are of different shape, and, in mature enamel, the protoprisms remain as protoprisms and do not develop into the conventional prismatic structures characteristic of mammalian enamel.

Additional material of the enigmatic Early Miocene mammal Kelba and its relationship to the order Ptolemaiida.

Kelba quadeemae, a fossil mammal from the Early Miocene of East Africa, was originally named on the basis of three isolated upper molars. Kelba has previously been interpreted as a creodont, a pantolestid, an insectivoran, and a hemigaline viverrid. The true affinities of this taxon have remained unclear because of the limited material and its unique morphology relative to other Miocene African mammals. New material of Kelba from several East African Miocene localities, most notably a skull from the Early Miocene locality of Songhor in Western Kenya, permits analysis of the affinities of Kelba and documents the lower dentition of this taxon. Morphological comparison of this new material clearly demonstrates that Kelba is a member of the order Ptolemaiida, a poorly understood group whose fossil record was previously restricted to the Oligocene Fayum deposits of northern Egypt. Phylogenetic analysis supports the monophyly of the Ptolemaiida, including Kelba, and recovers two monophyletic clades within the order. We provide new family names for these groups and an emended diagnosis for the order. The discovery of ptolemaiidans from the Miocene of East Africa is significant because it extends the known temporal range of the order by >10 million years and the geographic range by >3,200 km. Although the higher-level affinities of the Ptolemaiida remain obscure, their unique morphology and distribution through a larger area of Africa (and exclusively Africa) lend support to the idea that Ptolemaiida may have an ancient African origin.

Prismatic dentine in the Australian lungfish, Neoceratodus forsteri (Osteichthyes: Dipnoi).

The Australian lungfish, Neoceratodus forsteri, has a dentition consisting of enamel, mantle dentine and bone, enclosing circumdenteonal, core and interdenteonal dentines. Branching processes from cells that produce interdenteonal dentine leave the cell surface at different angles, with collagen fibrils aligned parallel to the long axis of each process. In the interdenteonal dentine, crystals of calcium hydroxyapatite form within fibrils of collagen, and grow within a matrix of non-collagenous protein. Crystals are aligned parallel to the cell process, as are the original collagen fibrils. Because the processes are angled to the cell surface, the crystals within the core or interdenteonal dentine are arranged in bundles set at angles to each other. Apatite crystals in circumdenteonal dentine are finer and denser than those of the interdenteonal dentine, and form outside the fibrils of collagen. In mature circumdenteonal dentine the crystals of circumdenteonal dentine form a dense tangled mass, linked to interdenteonal dentine by isolated crystals. The functional lungfish tooth plate contains prisms of large apatite crystals in the interdenteonal dentine and masses of fine tangled crystals around each denteon. This confers mechanical strength on a structure with little enamel that is subjected to heavy wear.