Patrick M. O’Connor

Palaeontological evidence for an Oligocene divergence between Old World monkeys and apes

Apes and Old World monkeys are prominent components of modern African and Asian ecosystems, yet the earliest phases of their evolutionary history have remained largely undocumented. The absence of crown catarrhine fossils older than ∼20 million years (Myr) has stood in stark contrast to molecular divergence estimates of ∼25–30 Myr for the split between Cercopithecoidea (Old World monkeys) and Hominoidea (apes), implying long ghost lineages for both clades. Here we describe the oldest known fossil ‘ape’, represented by a partial mandible preserving dental features that place it with ‘nyanzapithecine’ stem hominoids. Additionally, we report the oldest stem member of the Old World monkey clade, represented by a lower third molar. Both specimens were recovered from a precisely dated 25.2-Myr-old stratum in the Rukwa Rift, a segment of the western branch of the East African Rift in Tanzania. These finds extend the fossil record of apes and Old World monkeys well into the Oligocene epoch of Africa, suggesting a possible link between diversification of crown catarrhines and changes in the African landscape brought about by previously unrecognized tectonic activity in the East African rift system.

First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism

Previously known only from isolated teeth and lower jaw fragments recovered from the Cretaceous and Palaeogene of the Southern Hemisphere, the Gondwanatheria constitute the most poorly known of all major mammaliaform radiations. Here we report the discovery of the first skull material of a gondwanatherian, a complete and well-preserved cranium from Upper Cretaceous strata in Madagascar that we assign to a new genus and species. Phylogenetic analysis strongly supports its placement within Gondwanatheria, which are recognized as monophyletic and closely related to multituberculates, an evolutionarily successful clade of Mesozoic mammals known almost exclusively from the Northern Hemisphere. The new taxon is the largest known mammaliaform from the Mesozoic of Gondwana. Its craniofacial anatomy reveals that it was herbivorous, large-eyed and agile, with well-developed high-frequency hearing and a keen sense of smell. The cranium exhibits a mosaic of primitive and derived features, the disparity of which is extreme and probably reflective of a long evolutionary history in geographic isolation.

The Earliest Colubroid-Dominated Snake Fauna from Africa: Perspectives from the Late Oligocene Nsungwe Formation of Southwestern Tanzania

The extant snake fauna has its roots in faunal upheaval occurring across the Paleogene - Neogene transition. On northern continents, this turnover is well established by the late early Miocene. However, this transition is poorly documented on southern landmasses, particularly on continental Africa, where no late Paleogene terrestrial snake assemblages are documented south of the equator. Here we describe a newly discovered snake fauna from the Late Oligocene Nsungwe Formation in the Rukwa Rift Basin of Tanzania. The fauna is small but diverse with eight identifiable morphotypes, comprised of three booids and five colubroids. This fauna includes Rukwanyoka holmani gen. et sp. nov., the oldest boid known from mainland Africa. It also provides the oldest fossil evidence for the African colubroid clade Elapidae. Colubroids dominate the fauna, comprising more than 75% of the recovered material. This is likely tied to local aridification and/or seasonality and mirrors the pattern of overturn in later snake faunas inhabiting the emerging grassland environments of Europe and North America. The early emergence of colubroid dominance in the Rukwa Rift Basin relative to northern continents suggests that the pattern of overturn that resulted in extant faunas happened in a more complex fashion on continental Africa than was previously realized, with African colubroids becoming at least locally important in the late Paleogene, either ahead of or as a consequence of the invasion of colubrids. The early occurrence of elapid snakes in the latest Oligocene of Africa suggests the clade rapidly spread from Asia to Africa, or arose in Africa, before invading Europe.

Ecology and caudal skeletal morphology in birds: the convergent evolution of pygostyle shape in underwater foraging taxa.

Birds exhibit a specialized tail that serves as an integral part of the flight apparatus, supplementing the role of the wings in facilitating high performance aerial locomotion. The evolution of this function for the tail contributed to the diversification of birds by allowing them to utilize a wider range of flight behaviors and thus exploit a greater range of ecological niches. The shape of the wings and the tail feathers influence the aerodynamic properties of a bird. Accordingly, taxa that habitually utilize different flight behaviors are characterized by different flight apparatus morphologies. This study explores whether differences in flight behavior are also associated with variation in caudal vertebra and pygostyle morphology. Details of the tail skeleton were characterized in 51 Aequornithes and Charadriiformes species. Free caudal vertebral morphology was measured using linear metrics. Variation in pygostyle morphology was characterized using Elliptical Fourier Analysis, a geometric morphometric method for the analysis of outline shapes. Each taxon was categorized based on flight style (flap, flap-glide, dynamic soar, etc.) and foraging style (aerial, terrestrial, plunge dive, etc.). Phylogenetic MANOVAs and Flexible Discriminant Analyses were used to test whether caudal skeletal morphology can be used to predict flight behavior. Foraging style groups differ significantly in pygostyle shape, and pygostyle shape predicts foraging style with less than 4% misclassification error. Four distinct lineages of underwater foraging birds exhibit an elongate, straight pygostyle, whereas aerial and terrestrial birds are characterized by a short, dorsally deflected pygostyle. Convergent evolution of a common pygostyle phenotype in diving birds suggests that this morphology is related to the mechanical demands of using the tail as a rudder during underwater foraging. Thus, distinct locomotor behaviors influence not only feather attributes but also the underlying caudal skeleton, reinforcing the importance of the entire caudal locomotor module in avian ecological diversification.

Oligocene Termite Nests with In Situ Fungus Gardens from the Rukwa Rift Basin, Tanzania, Support a Paleogene African Origin for Insect Agriculture.

Based on molecular dating, the origin of insect agriculture is hypothesized to have taken place independently in three clades of fungus-farming insects: the termites, ants or ambrosia beetles during the Paleogene (66–24 Ma). Yet, definitive fossil evidence of fungus-growing behavior has been elusive, with no unequivocal records prior to the late Miocene (7–10 Ma). Here we report fossil evidence of insect agriculture in the form of fossil fungus gardens, preserved within 25 Ma termite nests from southwestern Tanzania. Using these well-dated fossil fungus gardens, we have recalibrated molecular divergence estimates for the origins of termite agriculture to around 31 Ma, lending support to hypotheses suggesting an African Paleogene origin for termite-fungus symbiosis; perhaps coinciding with rift initiation and changes in the African landscape.

Endocranial and Inner Ear Morphology of Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar

ABSTRACT We present the first digital reconstruction of the endocranial cavity and endosseous labyrinth of the Late Cretaceous gondwanatherian mammal Vintana sertichi from the Maevarano Formation of Madagascar. The Malagasy specimen is exceptionally well preserved and represents the only described cranium known for Gondwanatheria, an enigmatic clade from the Late Cretaceous and Paleogene of Gondwana. The endocranial cast of Vintana is relatively small for an animal of its estimated body mass. Its encephalization quotient is 0.28–0.56 for a range of body mass estimates, which is similar to that of basal mammaliaforms. The olfactory bulbs are very large, occupying over 14% of the endocranial volume. The cerebral hemispheres are only slightly expanded, more similar to the condition in Morganucodon than to that in multituberculates and monotremes. Unlike the condition in other Mesozoic mammaliaforms, the endocast is greatly flexed at the circular fissure. The osseous labyrinth displays a mixture of derived and primitive features. The cochlear canal is only slightly curved and short compared with that of extant therians. The ratio between total cochlear canal length and maximum cranial length is smaller than in basal mammaliaforms and approximates that of non-mammaliaform cynodonts. By contrast, the presence of both primary and secondary osseous laminae, the tractus foraminosus, and Rosenthals canal represent derived characteristics of the mammalian inner ear typical of cladotherians. A modern innervation of the cochlea has either evolved independently in Vintana and cladotherians or was already present in the last common ancestor of both clades. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

A New Centrosaurine Ceratopsid, Machairoceratops cronusi gen et sp. nov., from the Upper Sand Member of the Wahweap Formation (Middle Campanian), Southern Utah

The Upper Cretaceous (middle-late Campanian) Wahweap Formation of southern Utah contains the oldest diagnostic evidence of ceratopsids (to date, all centrosaurines) in North America, with a number of specimens recovered from throughout a unit that spans between 81 and 77 Ma. Only a single specimen has been formally named, Diabloceratops eatoni, from the lower middle member of the formation. Machairoceratops cronusi gen. et sp. nov., a new centrosaurine ceratopsid from the upper member of the Wahweap Formation, is here described based on cranial material representing a single individual recovered from a calcareous mudstone. The specimen consists of two curved and elongate orbital horncores, a left jugal, a nearly complete, slightly deformed braincase, the left squamosal, and a mostly complete parietal ornamented by posteriorly projected, anterodorsally curved, elongate spikes on either side of a midline embayment. The fan-shaped, stepped-squamosal is diagnostic of Centrosaurinae, however, this element differs from the rectangular squamosal in Diabloceratops. Machairoceratops also differs in the possession of two anterodorsally (rather than laterally) curved epiparietal ornamentations on either side of a midline embayment that are distinguished by a posteromedially-oriented sulcus along the entire length of the epiparietal. Additionally, the parietosquamosal frill is lacking any other epiossifications along its periphery. Machairoceratops shares a triangular (rather than round) frill and spike-like epiparietal loci (p1) ornamentation with the stratigraphically lower Diabloceratops. Both parsimony and Bayesian phylogenetic analyses place Machairoceratops as an early-branching centrosaurine. However, the parsimony-based analysis provides little resolution for the position of the new taxon, placing it in an unresolved polytomy with Diabloceratops. The resultant Bayesian topology yielded better resolution, aligning Machairoceratops as the definitive sister taxon to a clade formed by Diabloceratops and Albertaceratops. Considered together, both phylogenetic methods unequivocally place Machairoceratops as an early-branching centrosaurine, and given the biostratigraphic position of Machairoceratops, these details increase the known ceratopsid diversity from both the Wahweap Formation and the southern portion of Laramidia. Finally, the unique morphology of the parietal ornamentation highlights the evolutionary disparity of frill ornamentation near the base of Centrosaurinae.

Sensory Anatomy and Sensory Ecology of Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar

ABSTRACT—Living mammals are distinguished from other extant tetrapods by adaptations for improved senses of hearing, touch, and smell. These adaptations, and concomitant reductions in visual anatomy, evolved during the Mesozoic in the mammalian and therian stem lineages. Here, we present a comparative study of the sensory anatomy of the Late Cretaceous gondwanatherian mammaliaform Vintana sertichi in order to draw inferences regarding its sensory abilities and sensory ecology. Our analyses demonstrate that Vintana has relatively large orbits that may have accommodated large eyes. Vintana also possessed cochlear primary and secondary osseous laminae and a cochlear canal that was relatively longer than in non-mammaliaform cynodonts but shorter than in extant therians. These features suggest that Vintana had some capacity for high-frequency hearing (i.e., >20 kHz), but that its cochlea may have encoded a more limited range of frequencies than the cochleas of most extant therians. The semicircular canals of Vintana have large radii of curvature and are nearly orthogonal, suggesting high sensitivity to angular head accelerations. These vestibular features may have evolved in order to stabilize large eyes during rapid and/or agile locomotion. Combined with evidence for large olfactory bulbs and a large trigeminal endocast, these data reveal that Vintana possessed a unique suite of sensory adaptations that distinguish it from other Mesozoic mammaliaforms. If these inferences are correct, then Vintana was probably a large-eyed and agile species, with a keen sense of smell and better high-frequency hearing than most other Mesozoic mammaliaforms. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Application of U-Pb detrital zircon geochronology to drill cuttings for age control in hydrocarbon exploration wells: a case study from the Rukwa Rift Basin, Tanzania

Precise dating and correlation of drilled wells through continental successions is challenging for hydrocarbon exploration, especially where preservation and recovery of age-diagnostic fossils is poor. As a complement or alternative to biostratigraphic dating we demonstrate the effectiveness of U–Pb geochronology via laser ablation–inductively coupled plasma–mass spectrometry on detrital zircon from well cuttings. In basins with syndepositional volcanic input, the youngest zircons in a stratigraphic interval can refine and serve as a proxy for the age of deposition. We demonstrate the reliability of this technique when applied to hydrocarbon exploration wells by analyzing drill cuttings through a continental interval of the Galula-1 well in the Rukwa Rift Basin, East African rift system, Tanzania, which previously yielded conflicting biostratigraphy results. The lower third of the well section reveals a late Miocene to Pliocene up-hole younging trend in the youngest detrital zircon populations, which matches new radioisotopic ages on volcanic tuffs from a correlative outcrop section. This is followed by an interval with recycled young zircons, followed by a zircon-free interval, interpreted to correspond to changes in magma composition of the nearby Rungwe volcanic province. This study provides the first radioisotopic age constraints for the Lake Beds in the Rukwa rift and demonstrates that sedimentation in the basin began by 8.7 Ma, critical for burial and thermal history modeling and establishing the probability of a working hydrocarbon system. Correspondence in age and zircon preservation between well and outcrop samples from the same intervals provides strong support for applying U–Pb detrital zircon geochronology to well cuttings, as a rapid, inexpensive approach for hydrocarbon exploration.

Abiotic and Biotic Factors as Predictors of Species Richness on Madagascar

Madagascar contains a diversity of endemic species. Yet levels of species richness vary among the isolated habitats scattered across the island. A number of ecological factors have been advanced to account for patterns of species richness. In particular, abiotic factors such as habitat area, latitude, altitude, temperature, and rainfall have been suggested to account for ultimate differences in the number of species a habitat may support. Biotic variables such as vegetation type have been suggested as more proximate factors in determining the diversity of habitats available for animals to occupy. Several studies have included Malagasy locales in evaluating large-scale relationships between ecological variables and species richness (e.g., [Reed and Fleagle, 1995], [Fleagle and Reed, 1996]; [Eeley and Lawes, 1999]; [Emmons, 1999]; [Ganzhorn et al., 1999]). This study combines data on geography, topography, climate, and vegetation with species lists from 27 national parks, reserves, and other protected areas to specifically address biogeographic patterns of species richness on Madagascar. Ecological variables are considered individually in order to determine which biotic and abiotic factors may best predict primate, mammal, bird, reptile, and amphibian richness.