Skull and partial skeleton of a new pachycetine genus (Cetacea, Basilosauridae) from the Aridal Formation, Bartonian middle Eocene, of southwestern Morocco.

Pachycetus paulsonii, Pachycetus wardii, and Antaecetus aithai are middle Eocene archaeocete whales found in Europe, North America, and Africa, respectively. The three are placed in the new basilosaurid subfamily Pachycetinae. Antaecetus is a new genus known from Egypt and Morocco, and the only pachycetine known from a substantial postcranial skeleton. The skull of A. aithai described here resembles that of Saghacetus osiris in size, but lacks the narrowly constricted rostrum of Saghacetus. Antaecetus is smaller than Pachycetus and its teeth are more gracile. Upper premolars differ in having two rather than three accessory cusps flanking the principal cusp. Pachycetines differ from dorudontines in having elongated posterior thoracic and lumbar vertebrae like those of Basilosaurus, but differ from basilosaurines and from dorudontines in having conspicuously pachyosteosclerotic vertebrae with dense and thickly laminated cortical bone surrounding a cancellous core. Pachycetinae are also distinctive in having transverse processes on lumbar vertebrae nearly as long anteroposteriorly as the corresponding centrum. We infer from their pachyosteosclerotic vertebrae that pachycetines were probably sirenian-like slow swimmers living in shallow coastal seas and feeding on passing fish and mobile invertebrates.

Pattern and rate in the Plio-Pleistocene evolution of modern human brain size.

Fourteen studies of brain size evolution in Plio-Pleistocene hominins published over the past fifty years show substantial long-term increase in endocranial volume (ECV) for the broad lineage leading to modern humans. The median generation-to-generation step rate for a consensus time series of ECV values, h0 = 0.15 standard deviations per generation, is almost identical to the median step rate observed in modern biological field studies. When specimens are aggregated in a series of 100 k.y. time bins to reflect the precision of their geological ages, temporal scaling identifies four successive phases of stasis and change that are significantly different from random. Phase I from about 3.2 to 2.0 million years before present is an initial phase of relative stasis. Phase II from 2.0 to 1.5 m.y. is a phase of directional brain size increase. Phase III from 1.5 to 0.7 m.y. is a second phase of stasis. Finally, Phase IV from about 0.7 m.y. to 10 k.y. is a second phase of directional increase. The tempo (rate) and the mode (stasis, random, or directional change) of an evolutionary time series are related to each other, and both are related to the time scale appropriate for analysis.

Large-bodied sabre-toothed anchovies reveal unanticipated ecological diversity in early Palaeogene teleosts.

Many modern groups of marine fishes first appear in the fossil record during the early Palaeogene (66-40 Ma), including iconic predatory lineages of spiny-rayed fishes that appear to have originated in response to ecological roles left empty after the Cretaceous/Palaeogene extinction. The hypothesis of extinction-mediated ecological release likewise predicts that other fish groups have adopted novel predatory ecologies. Here, we report remarkable trophic innovation in early Palaeogene clupeiforms (herrings and allies), a group whose modern representatives are generally small-bodied planktivores. Two forms, the early Eocene (Ypresian) †Clupeopsis from Belgium and a new genus from the middle Eocene (Lutetian) of Pakistan, bear conspicuous features indicative of predatory ecology, including large size, long gapes and caniniform dentition. Most remarkable is the presence of a single, massive vomerine fang offset from the midline in both. Numerous features of the neurocranium, suspensorium and branchial skeleton place these taxa on the engraulid (anchovy) stem as the earliest known representatives of the clade. The identification of large-bodied, piscivorous anchovies contributes to an emerging picture of a phylogenetically diverse guild of predatory ray-finned fishes in early Palaeogene marine settings, which include completely extinct lineages alongside members of modern marine groups and taxa that are today restricted to freshwater or deep-sea environments.

Reduction of olfactory and respiratory turbinates in the transition of whales from land to sea: the semiaquatic middle Eocene Aegyptocetus tarfa.

Ethmoturbinates, nasoturbinates, and maxilloturbinates are well developed in the narial tract of land-dwelling artiodactyls ancestral to whales, but these are greatly reduced or lost entirely in modern whales. Aegyptocetus tarfa is a semiaquatic protocetid from the middle Eocene of Egypt. Computed axial tomography scans of the skull show that A. tarfa retained all three sets of turbinates like a land mammal. It is intermediate between terrestrial artiodactyls and aquatic whales in reduction of the turbinates. Ethmoturbinates in A. tarfa have 26% of the surface area expected for an artiodactyl. These have an olfactory function and indicate that early whales retained a sense of smell in the transition from land to sea. Maxilloturbinates in A. tarfa have 6% of the surface area expected for an artiodactyl. These have a respiratory function and their markedly reduced size suggests that rapid inhalation and exhalation was already more important than warming and humidifying air, in contrast to extant land mammals. Finally, the maxilloturbinates of A. tarfa, although greatly reduced, still show some degree of similarity to those of artiodactyls, supporting the phylogenetic affinity of cetaceans and artiodactyls based on morphological and molecular evidence.

Aegicetus gehennae, a new late Eocene protocetid (Cetacea, Archaeoceti) from Wadi Al Hitan, Egypt, and the transition to tail-powered swimming in whales.

Aegicetus gehennae is a new African protocetid whale based on a partial skull with much of an associated postcranial skeleton. The type specimen, Egyptian Geological Museum, Cairo [CGM] 60584, was found near the base of the early-Priabonian-age (earliest late Eocene) Gehannam Formation of the Wadi Al Hitan World Heritage Site in Egypt. The cranium is distinctive in having ventrally-deflected exoccipitals. The vertebral column is complete from cervical C1 through caudal Ca9, with a vertebral formula of 7:15:4:4:9+, representing, respectively, the number of cervical, thoracic, lumbar, sacral, and caudal vertebrae. CGM 60584 has two more rib-bearing thoracic vertebrae than other known protocetids, and two fewer lumbars. Sacral centra are unfused, and there is no defined auricular surface on the ilium. Thus there was no weight-bearing sacroiliac joint. The sternum is distinctive in being exceptionally broad and flat. The body weight of CGM 60584, a putative male, is estimated to have been about 890 kg in life. Long bones of the fore and hind limbs are shorter than expected for a protocetid of this size. Bones of the manus are similar in length and more robust compared to those of the pes. A log vertebral length profile for CGM 60584 parallels that of middle Eocene Maiacetus inuus through the anterior and middle thorax, but more posterior vertebrae are proportionally longer. Vertebral elongation, loss of a sacroiliac articulation, and hind limb reduction indicate that Aegicetus gehennae was more fully aquatic and less specialized as a foot-powered swimmer than earlier protocetids. It is doubtful that A. gehennae had a tail fluke, and the caudal flattening known for basilosaurids is shorter relative to vertebral column length than flattening associated with a fluke in any modern whale. Late protocetids and basilosaurids had relatively long skeletons, longer than those known earlier and later, and the middle-to-late Eocene transition from foot-powered to tail-powered swimming seemingly involved some form of mid-body-and-tail undulation.

Stomach contents of the archaeocete Basilosaurus isis: Apex predator in oceans of the late Eocene.

Apex predators live at the top of an ecological pyramid, preying on animals in the pyramid below and normally immune from predation themselves. Apex predators are often, but not always, the largest animals of their kind. The living killer whale Orcinus orca is an apex predator in modern world oceans. Here we focus on an earlier apex predator, the late Eocene archaeocete Basilosaurus isis from Wadi Al Hitan in Egypt, and show from stomach contents that it fed on smaller whales (juvenile Dorudon atrox) and large fishes (Pycnodus mokattamensis). Our observations, the first direct evidence of diet in Basilosaurus isis, confirm a predator-prey relationship of the two most frequently found fossil whales in Wadi Al-Hitan, B. isis and D. atrox. This extends our understanding of their paleoecology. Late Eocene Basilosaurus isis, late Miocene Livyatan melvillei, and modern Orcinus orca are three marine apex predators known from relatively short intervals of time. Little is known about whales as apex predators through much of the Cenozoic era, and whales as apex predators deserve more attention than they have received.

Repetitive mammalian dwarfing during ancient greenhouse warming events.

Abrupt perturbations of the global carbon cycle during the early Eocene are associated with rapid global warming events, which are analogous in many ways to present greenhouse warming. Mammal dwarfing has been observed, along with other changes in community structure, during the largest of these ancient global warming events, known as the Paleocene-Eocene Thermal Maximum [PETM; ~56 million years ago (Ma)]. We show that mammalian dwarfing accompanied the subsequent, smaller-magnitude warming event known as Eocene Thermal Maximum 2 [ETM2 (~53 Ma)]. Statistically significant decrease in body size during ETM2 is observed in two of four taxonomic groups analyzed in this study and is most clearly observed in early equids (horses). During ETM2, the best-sampled lineage of equids decreased in size by ~14%, as opposed to ~30% during the PETM. Thus, dwarfing appears to be a common evolutionary response of some mammals during past global warming events, and the extent of dwarfing seems related to the magnitude of the event.

Temperature and atmospheric CO2 concentration estimates through the PETM using triple oxygen isotope analysis of mammalian bioapatite.

The Paleocene-Eocene Thermal Maximum (PETM) is a remarkable climatic and environmental event that occurred 56 Ma ago and has importance for understanding possible future climate change. The Paleocene-Eocene transition is marked by a rapid temperature rise contemporaneous with a large negative carbon isotope excursion (CIE). Both the temperature and the isotopic excursion are well-documented by terrestrial and marine proxies. The CIE was the result of a massive release of carbon into the atmosphere. However, the carbon source and quantities of CO2 and CH4 greenhouse gases that contributed to global warming are poorly constrained and highly debated. Here we combine an established oxygen isotope paleothermometer with a newly developed triple oxygen isotope paleo-CO2 barometer. We attempt to quantify the source of greenhouse gases released during the Paleocene-Eocene transition by analyzing bioapatite of terrestrial mammals. Our results are consistent with previous estimates of PETM temperature change and suggest that not only CO2 but also massive release of seabed methane was the driver for CIE and PETM.

Transition of Eocene whales from land to sea: evidence from bone microstructure.

Cetacea are secondarily aquatic amniotes that underwent their land-to-sea transition during the Eocene. Primitive forms, called archaeocetes, include five families with distinct degrees of adaptation to an aquatic life, swimming mode and abilities that remain difficult to estimate. The lifestyle of early cetaceans is investigated by analysis of microanatomical features in postcranial elements of archaeocetes. We document the internal structure of long bones, ribs and vertebrae in fifteen specimens belonging to the three more derived archaeocete families--Remingtonocetidae, Protocetidae, and Basilosauridae--using microtomography and virtual thin-sectioning. This enables us to discuss the osseous specializations observed in these taxa and to comment on their possible swimming behavior. All these taxa display bone mass increase (BMI) in their ribs, which lack an open medullary cavity, and in their femora, whereas their vertebrae are essentially spongious. Humeri and femora show opposite trends in microanatomical specialization in the progressive independence of cetaceans from a terrestrial environment. Humeri change from very compact to spongious, which is in accordance with the progressive loss of propulsive role for the forelimbs, which were used instead for steering and stabilizing. Conversely, hind-limbs in basilosaurids became strongly reduced with no involvement in locomotion but display strong osteosclerosis in the femora. Our study confirms that Remingtonocetidae and Protocetidae were almost exclusively aquatic in locomotion for the taxa sampled, which probably were shallow water suspended swimmers. Basilosaurids display osseous specializations similar to those of modern cetaceans and are considered more active open-sea swimmers. This study highlights the strong need for homologous sections in comparative microanatomical studies, and the importance of combining information from several bones of the same taxon for improved functional interpretation.

Species in the primate fossil record.

Species in the fossil record are population pools of genetic and phenetic variation at a place and time, morphologically recognizable and distinguishable from others by empirical standards. Change through time can be substantial, requiring subdivision of lineages that becomes more arbitrary as they become more complete. Evolution is about form, space, and time; it is about variation and change. Interpretation of species in the fossil record touches all of these.

Endocranial morphology of Palaeocene Plesiadapis tricuspidens and evolution of the early primate brain.

Expansion of the brain is a key feature of primate evolution. The fossil record, although incomplete, allows a partial reconstruction of changes in primate brain size and morphology through time. Palaeogene plesiadapoids, closest relatives of Euprimates (or crown-group primates), are crucial for understanding early evolution of the primate brain. However, brain morphology of this group remains poorly documented, and major questions remain regarding the initial phase of euprimate brain evolution. Micro-CT investigation of the endocranial morphology of Plesiadapis tricuspidens from the Late Palaeocene of Europe--the most complete plesiadapoid cranium known--shows that plesiadapoids retained a very small and simple brain. Plesiadapis has midbrain exposure, and minimal encephalization and neocorticalization, making it comparable with that of stem rodents and lagomorphs. However, Plesiadapis shares a domed neocortex and downwardly shifted olfactory-bulb axis with Euprimates. If accepted phylogenetic relationships are correct, then this implies that the euprimate brain underwent drastic reorganization during the Palaeocene, and some changes in brain structure preceded brain size increase and neocortex expansion during evolution of the primate brain.

Evolution of locomotion in Anthropoidea: the semicircular canal evidence.

Our understanding of locomotor evolution in anthropoid primates has been limited to those taxa for which good postcranial fossil material and appropriate modern analogues are available. We report the results of an analysis of semicircular canal size variation in 16 fossil anthropoid species dating from the Late Eocene to the Late Miocene, and use these data to reconstruct evolutionary changes in locomotor adaptations in anthropoid primates over the last 35 Ma. Phylogenetically informed regression analyses of semicircular canal size reveal three important aspects of anthropoid locomotor evolution: (i) the earliest anthropoid primates engaged in relatively slow locomotor behaviours, suggesting that this was the basal anthropoid pattern; (ii) platyrrhines from the Miocene of South America were relatively agile compared with earlier anthropoids; and (iii) while the last common ancestor of cercopithecoids and hominoids likely was relatively slow like earlier stem catarrhines, the results suggest that the basal crown catarrhine may have been a relatively agile animal. The latter scenario would indicate that hominoids of the later Miocene secondarily derived their relatively slow locomotor repertoires.

Cranial asymmetry in Eocene archaeocete whales and the evolution of directional hearing in water.

Eocene archaeocete whales gave rise to all modern toothed and baleen whales (Odontoceti and Mysticeti) during or near the Eocene-Oligocene transition. Odontocetes have asymmetrical skulls, with asymmetry linked to high-frequency sound production and echolocation. Mysticetes are generally assumed to have symmetrical skulls and lack high-frequency hearing. Here we show that protocetid and basilosaurid archaeocete skulls are distinctly and directionally asymmetrical. Archaeocete asymmetry involves curvature and axial torsion of the cranium, but no telescoping. Cranial asymmetry evolved in Eocene archaeocetes as part of a complex of traits linked to directional hearing (such as pan-bone thinning of the lower jaws, mandibular fat pads, and isolation of the ear region), probably enabling them to hear the higher sonic frequencies of sound-producing fish on which they preyed. Ultrasonic echolocation evolved in Oligocene odontocetes, enabling them to find silent prey. Asymmetry and much of the sonic-frequency range of directional hearing were lost in Oligocene mysticetes during the shift to low-frequency hearing and bulk-straining predation.

Continental warming preceding the Palaeocene-Eocene thermal maximum.

Marine and continental records show an abrupt negative shift in carbon isotope values at ∼55.8 Myr ago. This carbon isotope excursion (CIE) is consistent with the release of a massive amount of isotopically light carbon into the atmosphere and was associated with a dramatic rise in global temperatures termed the Palaeocene-Eocene thermal maximum (PETM). Greenhouse gases released during the CIE, probably including methane, have often been considered the main cause of PETM warming. However, some evidence from the marine record suggests that warming directly preceded the CIE, raising the possibility that the CIE and PETM may have been linked to earlier warming with different origins. Yet pre-CIE warming is still uncertain. Disentangling the sequence of events before and during the CIE and PETM is important for understanding the causes of, and Earth system responses to, abrupt climate change. Here we show that continental warming of about 5 °C preceded the CIE in the Bighorn Basin, Wyoming. Our evidence, based on oxygen isotopes in mammal teeth (which reflect temperature-sensitive fractionation processes) and other proxies, reveals a marked temperature increase directly below the CIE, and again in the CIE. Pre-CIE warming is also supported by a negative amplification of δ(13)C values in soil carbonates below the CIE. Our results suggest that at least two sources of warming-the earlier of which is unlikely to have been methane-contributed to the PETM.

New Oligocene primate from Saudi Arabia and the divergence of apes and Old World monkeys.

It is widely understood that Hominoidea (apes and humans) and Cercopithecoidea (Old World monkeys) have a common ancestry as Catarrhini deeply rooted in Afro-Arabia. The oldest stem Catarrhini in the fossil record are Propliopithecoidea, known from the late Eocene to early Oligocene epochs (roughly 35-30 Myr ago) of Egypt, Oman and possibly Angola. Genome-based estimates for divergence of hominoids and cercopithecoids range into the early Oligocene; however, the mid-to-late Oligocene interval from 30 to 23 Myr ago has yielded little fossil evidence documenting the morphology of the last common ancestor of hominoids and cercopithecoids, the timing of their divergence, or the relationship of early stem and crown catarrhines. Here we describe the partial cranium of a new medium-sized (about 15-20 kg) fossil catarrhine, Saadanius hijazensis, dated to 29-28 Myr ago. Comparative anatomy and cladistic analysis shows that Saadanius is an advanced stem catarrhine close to the base of the hominoid-cercopithecoid clade. Saadanius is important for assessing competing hypotheses about the ancestral morphotype for crown catarrhines, early catarrhine phylogeny and the age of hominoid-cercopithecoid divergence. Saadanius has a tubular ectotympanic but lacks synapomorphies of either group of crown Catarrhini, and we infer that the hominoid-cercopithecoid split happened later, between 29-28 and 24 Myr ago.

Complete primate skeleton from the Middle Eocene of Messel in Germany: morphology and paleobiology.

BACKGROUND: The best European locality for complete Eocene mammal skeletons is Grube Messel, near Darmstadt, Germany. Although the site was surrounded by a para-tropical rain forest in the Eocene, primates are remarkably rare there, and only eight fragmentary specimens were known until now. Messel has now yielded a full primate skeleton. The specimen has an unusual history: it was privately collected and sold in two parts, with only the lesser part previously known. The second part, which has just come to light, shows the skeleton to be the most complete primate known in the fossil record. METHODOLOGY/PRINCIPAL FINDINGS: We describe the morphology and investigate the paleobiology of the skeleton. The specimen is described as Darwinius masillae n.gen. n.sp. belonging to the Cercamoniinae. Because the skeleton is lightly crushed and bones cannot be handled individually, imaging studies are of particular importance. Skull radiography shows a host of teeth developing within the juvenile face. Investigation of growth and proportion suggest that the individual was a weaned and independent-feeding female that died in her first year of life, and might have attained a body weight of 650-900 g had she lived to adulthood. She was an agile, nail-bearing, generalized arboreal quadruped living above the floor of the Messel rain forest. CONCLUSIONS/SIGNIFICANCE: Darwinius masillae represents the most complete fossil primate ever found, including both skeleton, soft body outline and contents of the digestive tract. Study of all these features allows a fairly complete reconstruction of life history, locomotion, and diet. Any future study of Eocene-Oligocene primates should benefit from information preserved in the Darwinius holotype. Of particular importance to phylogenetic studies, the absence of a toilet claw and a toothcomb demonstrates that Darwinius masillae is not simply a fossil lemur, but part of a larger group of primates, Adapoidea, representative of the early haplorhine diversification.

New protocetid whale from the middle eocene of pakistan: birth on land, precocial development, and sexual dimorphism.

BACKGROUND: Protocetidae are middle Eocene (49-37 Ma) archaeocete predators ancestral to later whales. They are found in marine sedimentary rocks, but retain four legs and were not yet fully aquatic. Protocetids have been interpreted as amphibious, feeding in the sea but returning to land to rest. METHODOLOGY/PRINCIPAL FINDINGS: Two adult skeletons of a new 2.6 meter long protocetid, Maiacetus inuus, are described from the early middle Eocene Habib Rahi Formation of Pakistan. M. inuus differs from contemporary archaic whales in having a fused mandibular symphysis, distinctive astragalus bones in the ankle, and a less hind-limb dominated postcranial skeleton. One adult skeleton is female and bears the skull and partial skeleton of a single large near-term fetus. The fetal skeleton is positioned for head-first delivery, which typifies land mammals but not extant whales, evidence that birth took place on land. The fetal skeleton has permanent first molars well mineralized, which indicates precocial development at birth. Precocial development, with attendant size and mobility, were as critical for survival of a neonate at the land-sea interface in the Eocene as they are today. The second adult skeleton is the most complete known for a protocetid. The vertebral column, preserved in articulation, has 7 cervicals, 13 thoracics, 6 lumbars, 4 sacrals, and 21 caudals. All four limbs are preserved with hands and feet. This adult is 12% larger in linear dimensions than the female skeleton, on average, has canine teeth that are 20% larger, and is interpreted as male. Moderate sexual dimorphism indicates limited male-male competition during breeding, which in turn suggests little aggregation of food or shelter in the environment inhabited by protocetids. CONCLUSIONS/SIGNIFICANCE: Discovery of a near-term fetus positioned for head-first delivery provides important evidence that early protocetid whales gave birth on land. This is consistent with skeletal morphology enabling Maiacetus to support its weight on land and corroborates previous ideas that protocetids were amphibious. Specimens this complete are virtual 'Rosetta stones' providing insight into functional capabilities and life history of extinct animals that cannot be gained any other way.

Rapid Asia-Europe-North America geographic dispersal of earliest Eocene primate Teilhardina during the Paleocene-Eocene Thermal Maximum.

True primates appeared suddenly on all three northern continents during the 100,000-yr-duration Paleocene-Eocene Thermal Maximum at the beginning of the Eocene, approximately 55.5 mya. The simultaneous or nearly simultaneous appearance of euprimates on northern continents has been difficult to understand because the source area, immediate ancestors, and dispersal routes were all unknown. Now, omomyid haplorhine Teilhardina is known on all three continents in association with the carbon isotope excursion marking the Paleocene-Eocene Thermal Maximum. Relative position within the carbon isotope excursion indicates that Asian Teilhardina asiatica is oldest, European Teilhardina belgica is younger, and North American Teilhardina brandti and Teilhardina americana are, successively, youngest. Analysis of morphological characteristics of all four species supports an Asian origin and a westward Asia-to-Europe-to-North America dispersal for Teilhardina. High-resolution isotope stratigraphy indicates that this dispersal happened in an interval of approximately 25,000 yr. Rapid geographic dispersal and morphological character evolution in Teilhardina reported here are consistent with rates observed in other contexts.

Asian gliriform origin for arctostylopid mammals.

Arctostylopids are common in Late Palaeocene and Early Eocene mammal faunas of Asia, but they are rare in North America. From the time of their first discovery, arctostylopids were grouped with the strictly endemic South American Notoungulata based on their strikingly similar dental morphology. This relationship was challenged based on dental morphology of more recently discovered arctostylopids and on supposed tarsal morphology (based on unassociated material) of the Asian arctostylopid Palaeostylops. Therefore, Arctostylopidae were placed in a separate order, Arctostylopida, of enigmatic affinities. Many authors, however, continue to unite Arctostylopidae and Notoungulata based on dental similarities. In the Late Palaeocene Subeng site in Inner Mongolia, we identified the Palaeostylops tarsals based on their size and abundance. This identification is supported by comparison to unpublished Arctostylops tarsals from the North American Clarkforkian, derived from a semi-articulated skeleton also including dental material. Tarsal morphology shows moderate similarity to the gliriform Pseudictops, and strong resemblance to the tarsally conservative gliroid Rhombomylus. Hence, Arctostylopidae may best be grouped with Asian non-gliroid Gliriformes, which we interpret as having dispersed into North America in the Late Palaeocene.