Oldest evidence of abundant C4 grasses and habitat heterogeneity in eastern Africa.

The assembly of Africa's iconic C4 grassland ecosystems is central to evolutionary interpretations of many mammal lineages, including hominins. C4 grasses are thought to have become ecologically dominant in Africa only after 10 million years ago (Ma). However, paleobotanical records older than 10 Ma are sparse, limiting assessment of the timing and nature of C4 biomass expansion. This study uses a multiproxy design to document vegetation structure from nine Early Miocene mammal site complexes across eastern Africa. Results demonstrate that between ~21 and 16 Ma, C4 grasses were locally abundant, contributing to heterogeneous habitats ranging from forests to wooded grasslands. These data push back the oldest evidence of C4 grass-dominated habitats in Africa-and globally-by more than 10 million years, calling for revised paleoecological interpretations of mammalian evolution.

Lack of Rule-Adherence During Mountain Gorilla Tourism Encounters in Bwindi Impenetrable National Park, Uganda, Places Gorillas at Risk From Human Disease.

Mountain gorillas (Gorilla beringei beringei) are an endangered primate species, with ~43% of the 1,063 individuals that remain on the planet today residing in Bwindi Impenetrable National Park (BINP) in southwestern Uganda. These primates are at the heart of a growing tourism industry that has incentivized their continued protection, but close proximity between humans and gorillas during such encounters presents well-documented risks for disease transmission. The Uganda Wildlife Authority (UWA) has developed rules to help protect the health of the gorillas, limiting each habituated gorilla group to a single 60 min visit each day by a group of no more than 8 tourists, and emphasizing that humans maintain a >7 m distance from gorillas at all times. A number of studies have documented that not all tour groups respect these rules. This project assesses rule-adherence during gorilla tourism encounters at BINP using both observational and survey-based data collected during the tourism high season between May and August, 2014. Observational data from 53 treks reveal that groups of 1-11 tourists engaged in gorilla viewing encounters between 46 and 98 min in duration. Although 96% of pre-trek briefings conducted by park rangers emphasized the need to maintain >7 m human-gorilla spacing, the 7 m distance rule was violated in over 98% (52 out of 53) of the tours examined in this study. Observational data were collected at 2 min intervals during gorilla-viewing encounters, documenting the nearest distance between any tourist and a gorilla (n = 1,604), of which 1,094 observations (68.2%) took place at a distance less than or equal to 7 m. Importantly, the 7 m rule was violated in visits to all of the gorilla groups habituated during the time of the study. In 224 observations (~14%, per 1,604 total), human-gorilla spacing was 3 m or less. Survey data (n = 243) revealed promising opportunities to improve tourist understanding of and adherence to park rules, with 73.6% of respondents indicating that they would be willing to utilize a precautionary measure of wearing a face-mask during encounters to protect gorilla health.

Fossil lemurs from Egypt and Kenya suggest an African origin for Madagascar's aye-aye.

In 1967 G.G. Simpson described three partial mandibles from early Miocene deposits in Kenya that he interpreted as belonging to a new strepsirrhine primate, Propotto. This interpretation was quickly challenged, with the assertion that Propotto was not a primate, but rather a pteropodid fruit bat. The latter interpretation has not been questioned for almost half a century. Here we re-evaluate the affinities of Propotto, drawing upon diverse lines of evidence to establish that this strange mammal is a strepsirrhine primate as originally suggested by Simpson. Moreover, our phylogenetic analyses support the recognition of Propotto, together with late Eocene Plesiopithecus from Egypt, as African stem chiromyiform lemurs that are exclusively related to the extant aye-aye (Daubentonia) from Madagascar. Our results challenge the long-held view that all lemurs are descended from a single ancient colonization of Madagascar, and present an intriguing alternative scenario in which two lemur lineages dispersed from Africa to Madagascar independently, possibly during the later Cenozoic.

The first hyaenodont from the late Oligocene Nsungwe Formation of Tanzania: Paleoecological insights into the Paleogene-Neogene carnivore transition.

Throughout the Paleogene, most terrestrial carnivore niches in Afro-Arabia were occupied by Hyaenodonta, an extinct lineage of placental mammals. By the end of the Miocene, terrestrial carnivore niches had shifted to members of Carnivora, a clade with Eurasian origins. The transition from a hyaenodont-carnivore fauna to a carnivoran-carnivore fauna coincides with other ecological changes in Afro-Arabia as tectonic conditions in the African Rift System altered climatic conditions and facilitated faunal exchange with Eurasia. Fossil bearing deposits in the Nsungwe Formation in southwestern Tanzania are precisely dated to ~25.2 Ma (late Oligocene), preserving a late Paleogene Afro-Arabian fauna on the brink of environmental transition, including the earliest fossil evidence of the split between Old World monkeys and apes. Here we describe a new hyaenodont from the Nsungwe Formation, Pakakali rukwaensis gen. et sp. nov., a bobcat-sized taxon known from a portion of the maxilla that preserves a deciduous third premolar and alveoli of dP4 and M1. The crown of dP3 bears an elongate parastyle and metastyle and a small, blade-like metacone. Based on alveolar morphology, the two more distal teeth successively increased in size and had relatively large protocones. Using a hyaenodont character-taxon matrix that includes deciduous dental characters, Bayesian phylogenetic methods resolve Pakakali within the clade Hyainailouroidea. A Bayesian biogeographic analysis of phylogenetic results resolve the Pakakali clade as Afro-Arabian in origin, demonstrating that this small carnivorous mammal was part of an endemic Afro-Arabian lineage that persisted into the Miocene. Notably, Pakakali is in the size range of carnivoran forms that arrived and began to diversify in the region by the early Miocene. The description of Pakakali is important for exploring hyaenodont ontogeny and potential influences of Afro-Arabian tectonic events upon mammalian evolution, providing a deep time perspective on the stability of terrestrial carnivore niches through time.

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.

A detailed assessment of the maxillary morphology of Limnopithecus evansi with implications for the taxonomy of the genus.

Limnopithecus is a small-bodied catarrhine genus that is widespread throughout early Miocene sites in East Africa. Although two species of this genus have been described - Limnopithecus legetet (type species) and Limnopithecus evansi - they are poorly known anatomically and their systematic positions remain unresolved. Here, we provide detailed descriptions and comparisons for two well-preserved maxillary specimens that we attribute to L. evansi. These specimens come from the type locality of the species, Songhor in western Kenya, and add greatly to our knowledge of its dentognathic morphology. Together, they preserve the entire unilateral upper dentition, with overlapping elements demonstrating conspecificity, and provide new information about I(2) morphology and aspects of the palate, nasal aperture, and maxillary sinuses. Detailed morphological comparisons suggest that specimens referred to Limnopithecus from Songhor, Koru, and Rusinga share a unique I(2) morphology not found in any other early Miocene catarrhine. This argues in favor of congeneric status for L. evansi and L. legetet. Moreover, features such as a broad palate, premolar morphology, and the relative proportions of the premolars of L. evansi distinguish it from Lomorupithecus harrisoni, another early Miocene catarrhine from Napak, Uganda. This finding challenges a recently proposed taxonomic interpretation that Lomorupithecus and L. evansi are conspecific. Our results underscore the distinctiveness of L. evansi and Lo. harrisoni, thereby reaffirming the validity of the taxon Lo. harrisoni and indicating that the Songhor and Napak catarrhine communities were relatively distinct, despite their apparent contemporaneity.

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.

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.

Diversity in the later Paleogene proboscidean radiation: a small barytheriid from the Oligocene of Dhofar Governorate, Sultanate of Oman.

Despite significant recent improvements to our understanding of the early evolution of the Order Proboscidea (elephants and their extinct relatives), geographic sampling of the group's Paleogene fossil record remains strongly biased, with the first ~30 million years of proboscidean evolution documented solely in near-coastal deposits of northern Africa. The considerable morphological disparity that is observable among the late Eocene and early Oligocene proboscideans of northern Africa suggests that other, as yet unsampled, parts of Afro-Arabia might have served as important centers for the early diversification of major proboscidean clades. Here we describe the oldest taxonomically diagnostic remains of a fossil proboscidean from the Arabian Peninsula, a partial mandible of Omanitherium dhofarensis (new genus and species), from near the base of the early Oligocene Shizar Member of the Ashawq Formation, in the Dhofar Governorate of the Sultanate of Oman. The molars and premolars of Omanitherium are morphologically intermediate between those of Arcanotherium and Barytherium from northern Africa, but its specialized lower incisors are unlike those of other known Paleogene proboscideans in being greatly enlarged, high-crowned, conical, and tusk-like. Omanitherium is consistently placed close to late Eocene Barytherium in our phylogenetic analyses, and we place the new genus in the Family Barytheriidae. Some features of Omanitherium, such as tusk-like lower second incisors, the possible loss of the lower central incisors, an enlarged anterior mental foramen, and inferred elongate mandibular symphysis and diminutive P(2), suggest a possible phylogenetic link with Deinotheriidae, an extinct family of proboscideans whose origins have long been mysterious.

The evolution of mammal-like crocodyliforms in the Cretaceous Period of Gondwana.

Fossil crocodyliforms discovered in recent years have revealed a level of morphological and ecological diversity not exhibited by extant members of the group. This diversity is particularly notable among taxa of the Cretaceous Period (144-65 million years ago) recovered from former Gondwanan landmasses. Here we report the discovery of a new species of Cretaceous notosuchian crocodyliform from the Rukwa Rift Basin of southwestern Tanzania. This small-bodied form deviates significantly from more typical crocodyliform craniodental morphologies, having a short, broad skull, robust lower jaw, and a dentition with relatively few teeth that nonetheless show marked heterodonty. The presence of morphologically complex, complementary upper and lower molariform teeth suggests a degree of crown-crown contact during jaw adduction that is unmatched among known crocodyliforms, paralleling the level of occlusal complexity seen in mammals and their extinct relatives. The presence of another small-bodied mammal-like crocodyliform in the Cretaceous of Gondwana indicates that notosuchians probably filled niches and inhabited ecomorphospace that were otherwise occupied by mammals on northern continents.

First dinosaur tracks from the Arabian Peninsula.

BACKGROUND: The evolutionary history of Mesozoic terrestrial vertebrates from the Arabian Peninsula is virtually unknown. Despite vast exposures of rocky outcrops, only a handful of fossils have yet been described from the region. Here we report a multi-taxon dinosaur track assemblage near Madar village, 47 km north of Sana'a, Republic of Yemen. This represents the first dinosaur tracksite from the Arabian Peninsula, and the only multi-taxon dinosaur ichnosite in the Middle East. METHODOLOGY/FINDINGS: Measurements were taken directly from trackway impressions, following standard ichnological conventions. The presence of bipedal trackmakers is evidenced by a long series of pes imprints preserving smoothly rounded posterior margins, no evidence of a hallux, bluntly rounded digit tips and digital divarication angles characteristic of ornithopod dinosaurs. Nearby, eleven parallel quadrupedal trackways document a sauropod herd that included large and small individuals traveling together. Based on the morphology of manus impressions along with a narrow-gauged stance, the quadrupedal trackways were made by non-titanosauriform neosauropods. Additional isolated tracks and trackways of sauropod and ornithopod dinosaurs are preserved nearby. CONCLUSIONS/SIGNIFICANCE: Taken together, these discoveries present the most evocative window to date into the evolutionary history of dinosaurs of the Arabian Peninsula. Given the limited Mesozoic terrestrial record from the region, this discovery is of both temporal and geographic significance, and massive exposures of similarly-aged outcrops nearby offer great promise for future discoveries.

Body mass distribution and gait mechanics in fat-tailed dwarf lemurs (Cheirogaleus medius) and patas monkeys (Erythrocebus patas).

Most quadrupeds walk with lateral sequence (LS) gaits, where hind limb touchdowns are followed by ipsilateral forelimb touchdowns. Primates, however, typically walk with diagonal sequence (DS) gaits, where hind limb touchdowns are followed by contralateral forelimb touchdowns. Because the use of DS gaits is nearly ubiquitous among primates, understanding gait selection in primates is critical to understanding primate locomotor evolution. The Support Polygon Model [Tomita, M., 1967. A study on the movement pattern of four limbs in walking. J. Anthropol. Soc. Nippon 75, 120-146; Rollinson, J., Martin, R.D., 1981. Comparative aspects of primate locomotion, with special reference to arboreal cercopithecines. Symp. Zool. Soc. Lond. 48, 377-427] argues that primates' use of DS gaits stems from a more caudal position of the whole-body center of mass (COM) relative to other mammals. We tested the predictions of the Support Polygon Model by examining the effects of natural and experimental variations in COM position on gait mechanics in two distantly related primates: fat-tailed dwarf lemurs (Cheirogaleus medius) and patas monkeys (Erythrocebus patas). Dwarf lemur experiments compared individuals with and without a greatly enlarged tail (a feature associated with torpor that can be expected to shift the COM caudally). During patas monkey experiments, we experimentally shifted the COM cranially with the use of a weighted belt (7-12% of body mass) positioned above the scapulae. Examination of limb kinematics revealed changes consistent with systematic deviations in COM position. Nevertheless, footfall patterns changed in a direction contrary to the predictions of the Support Polygon Model in the dwarf lemurs and did not change at all in the patas monkey. These results suggest that body mass distribution is unlikely to be the sole determinant of footfall pattern in primates and other mammals.

Malagasy primate origins: phylogenies, fossils, and biogeographic reconstructions.

The geographic origin of Malagasy primates is a rich source of debate, providing a useful context for understanding effects of differing phylogenetic interpretations upon area of origin reconstructions. This study has evaluated the biogeographic implications of competing primate phylogenies in order to reconstruct the area of origin of Malagasy strepsirhines. The robusticity of biogeographic inference is examined based on sensitivity to tree topology. The results demonstrate extreme vulnerability to both out-group choice and internal tree topology, suggesting caution for area of origin interpretations from phylogenies that exclude fossil taxa. Moreover, even a single taxon can have a powerful effect upon biogeographic interpretations. Perhaps not surprisingly, it is only with greater phylogenetic resolution that a clearer understanding of the biogeographic origins of Malagasy primates will emerge.

Technical note: out-of-plane angular correction based on a trigonometric function for use in two-dimensional kinematic studies.

In two-dimensional (2D) kinematic studies, limb positions in three-dimensional (3D) space observed in lateral view are projected onto a 2D film plane. Elbow and knee-joint angles that are less than 20 degrees out-of-plane of lateral-view cameras generally exhibit very little measurable difference from their 3D counterparts (Plagenhoef 1979 Environment, Behavior, and Morphology; New York: Gustav Fisher, p. 95-118). However, when limb segment angles are more than 20 degrees out-of-plane, as is often the case in locomotor studies of arboreal primates, elbow and knee angles can appear significantly more extended than they actually are. For this reason, a methodology is described that corrects 2D out-of-plane angular estimates using a series of trigonometric transformations.