Geology and geochronology of the middle Miocene Kipsaramon site complex, Muruyur Beds, Tugen Hills, Kenya.

The Muruyur Beds are a substantial sedimentary deposit within a middle Miocene sequence of mafic volcanic flows associated with early stages of rifting in the central Kenyan Rift Valley. They are best represented in the Muruyur region, near Bartabwa, north of Kipsaramon, where dates range from 16.0 to 13.4 Ma. At Kipsaramon, located about 10 km south of Muruyur along the crest of the Tugen Hills, the upper Muruyur Beds are absent and the lower part can be divided into three members. Important fossil sites within Member 1 are dated between 15.8 and 15.6 Ma, and within Member 3 between 15.6 and 15.4 Ma. BPRP#89, in Member 1, is a bonebed at least 2500 m(2)in areal extent and up to 30 cm thick, which constitutes one of the richest concentrations of in situ fossil vertebrate bones in eastern Africa. BPRP#91, at approximately the same level at BPRP#89, is the source of a hominoid talus and other mammal and bird fossils. In Member 3, BPRP#122 has produced specimens of at least five individuals of the hominoid Equatorius, including a partial skeleton. The Muyuyur Beds were deposited near the western margin of a lake that was formed during the early stages of faulting and volcanism in the African Rift system. The bonebed in Member 1 appears to represent the influx of fluvially transported vertebrate and plant remains into a shallow portion of the lake. Elements of the fauna as well as stable isotopes that indicate both forest and more open environments occurred in proximity to the lake during the time of deposition of Member 1.     

Major Radiations in the Evolution of Caviid Rodents: Reconciling Fossils,Ghost Lineages,and Relaxed Molecular Clocks

Background

Caviidae is a diverse group of caviomorph rodents that is broadly distributed in South America and is divided into three highly divergent extant lineages: Caviinae (cavies), Dolichotinae (maras), and Hydrochoerinae (capybaras). The fossil record of Caviidae is only abundant and diverse since the late Miocene. Caviids belongs to Cavioidea sensu stricto (Cavioidea s.s.) that also includes a diverse assemblage of extinct taxa recorded from the late Oligocene to the middle Miocene of South America (“eocardiids”).

Results

A phylogenetic analysis combining morphological and molecular data is presented here, evaluating the time of diversification of selected nodes based on the calibration of phylogenetic trees with fossil taxa and the use of relaxed molecular clocks. This analysis reveals three major phases of diversification in the evolutionary history of Cavioidea s.s. The first two phases involve two successive radiations of extinct lineages that occurred during the late Oligocene and the early Miocene. The third phase consists of the diversification of Caviidae. The initial split of caviids is dated as middle Miocene by the fossil record. This date falls within the 95% higher probability distribution estimated by the relaxed Bayesian molecular clock, although the mean age estimate ages are 3.5 to 7 Myr older. The initial split of caviids is followed by an obscure period of poor fossil record (refered here as the Mayoan gap) and then by the appearance of highly differentiated modern lineages of caviids, which evidentially occurred at the late Miocene as indicated by both the fossil record and molecular clock estimates.

Conclusions

The integrated approach used here allowed us identifying the agreements and discrepancies of the fossil record and molecular clock estimates on the timing of the major events in cavioid evolution, revealing evolutionary patterns that would not have been possible to gather using only molecular or paleontological data alone.     

Scolecophidia (Serpentes) of the Late Oligocene and Early Miocene,North America,and a fossil history overview

An abundant fossil record of the snake clade Scolecophidia exists in Europe; however, the minute snake is noticeably absent in reports about the North American Paleogene and Neogene. Presented here are four localities from Florida, USA, that contain scolecophidian remains older than the Pleistocene: Thomas Farm (late Early Miocene, Hemingfordian Land Mammal Age, LMA), Live Oak (Oligocene-Miocene transition, latest Arikareean LMA), White Springs 3B (late Arikareean LMA), and Brooksville 2 (Late Oligocene, middle Arikareean LMA). These remains extend their known existence by about 26 m.y. and are now the oldest reported scolecophidian remains in North America. Molecular evidence on extant scolecophidians concludes that these tiny snakes have a Gondwanan origin. Interestingly, the oldest record of a scolecophidian is from Europe (Belgium) and dates back to the middle Paleocene (MP 1–5). The earliest African record of the snake clade comes from the Paleocene-Eocene boundary in Morocco. The clade is apparently absent from Europe and Middle East deposits dating from the latest Eocene through to the latest Oligocene (MP 19–30) and to the Early Miocene (MN 4). A portion of this time is known as the booid ‘Dark Period’ which represents an apparent response to global aridization and cooling. Scolecophidians appear to re-emerge into the southern Eurasian record in the Early Miocene (MN 4) and become widely dispersed throughout Europe and Middle East. The fossil record of these minute snakes is largely absent in southern Asia and South America. It is possible that the current lack of a decent fossil scolecophidian record outside of Europe and Middle East is due mainly to a bias in the methodology to recover fossils; wet sieving sediments through < 1.0 mm mesh is needed to recover the minuscule vertebrae.     

Dental remains of Equatorius africanus from Kipsaramon, Tugen Hills, Baringo District, Kenya.

Forty-one isolated large hominoid teeth, as well as most of the mandibular and three maxillary teeth associated with a partial skeleton, were recovered from middle Miocene Muruyur sediments near Kipsaramon in the Tugen Hills, Baringo District, Kenya. The isolated teeth were collected as surface finds and the skeleton was excavated in situ at locality BPRP#122 dated between 15.58 Ma and 15.36 Ma. The majority of the teeth recovered at BPRP#122 are referable to a minimum of five individuals of the hominoid Equatorius africanus. Three of the teeth, however, are provisionally assigned to Nyanzapithecus sp. The new hominoids from Kipsaramon add to an increasing inventory of specimens that suggest greater large hominoid taxonomic diversity from the middle Miocene of Kenya than was previously recognized. It is suggested that there are two large-bodied hominoid species present at Mabako, only one of which is assignable to Equatorius.     

Enters the shrew,some considerations on the Miocene palaeobiogeography of Iberian insectivores

The fossil record of the Iberian insectivores forms a subset of those found in central Europe. Comparison of the late early to early late Miocene record of the two areas shows that, particularly during the late Early Miocene, central European taxa have transient occurrences in Spain. Most taxa appear earlier and survive longer in central Europe. A notable exception is the gymnure Galerix, which extirpates earlier in central Europe, except for a transient occurrence in Germany just prior to its extinction. The main period of insectivore migrations is the late middle Miocene, although some of the taxa that enter remain restricted to the coastal areas. Overall, the pattern of distribution in time and space is best explained by the preference of insectivores for humid environments, as were found during the early Miocene and re-appeared at the end of the middle Miocene.     

Miocene cercopithecoidea from the Tugen Hills, Kenya

Miocene to Pleistocene fossiliferous sediments in the Tugen Hills span the time period from at least 15.5 Ma to 0.25 Ma, including time periods unknown or little known elsewhere in Africa. Consequently, the Tugen Hills deposits hold the potential to inform us about crucial phylogenetic events in African faunal evolution and about long-term environmental change. Among the specimens collected from this region are a number of discoveries already important to the understanding of primate evolution. Here, we describe additional cercopithecoid material from the Miocene deposits in the Tugen Hills sequence, including those from securely dated sites in the Muruyur Beds (16-13.4 Ma), the Mpesida Beds (7-6.2 Ma) and the Lukeino Formation (∼6.2-5.7 Ma). We also evaluate previously described material from the Ngorora Formation (13-8.8 Ma). Identified taxa include Victoriapithecidae gen. et sp. indet., cf. Parapapio lothagamensis, and at least two colobines. Specimens attributed to cf. Pp. lothagamensis would extend the species’ geographic range beyond its type locality. In addition, we describe specimens sharing derived characters with modern African colobines (Tribe: Colobina), a finding that is congruent with previous molecular estimates of colobine divergence dates. These colobine specimens represent some of the earliest known members of the modern African colobine radiation and, in contrast to previous hypotheses, suggest that early African colobines were mainly arboreal and that semi-terrestrial Late Miocene and Plio-Pleistocene colobine taxa were secondarily derived in their locomotor adaptations.     

The westernmost tarsier: A new genus and species from the Miocene of Pakistan

As the closest living sister group of anthropoids, tarsiers (Family Tarsiidae) are an important group in primate evolution. However, their fossil record is poor: only four species have been described, two from the Eocene of China and two from the Miocene of Thailand. All are from outside the range of the living species, which occur only on islands off Southeast Asia. Here, we describe a new fossil tarsier from Pakistan, a significant range extension. This record consists of two lower molars, an upper molar, and a lower premolar found in the Miocene Manchar Formation (∼18–16 Ma [millions of years ago]) of Sindh Province, southern Pakistan. The Pakistani tarsier is morphologically distinct from all living and fossil tarsiers, but most similar to the middle Miocene Thai species Tarsius thailandicus. Though living tarsiers have traditionally been classified in a single genus, a recent revision proposed a division into three genera, which is strongly supported by molecular data. The Pakistani species is not referable to any of these genera, and we create for it and T. thailandicus a new tarsiid genus. This discovery broadens our understanding of the geographic range and morphological diversity of Miocene tarsiers and helps to put the living tarsiers into their evolutionary context.     

The Pleistocene archaeology and environments of the Wasiriya Beds, Rusinga Island, Kenya

Western Kenya is well known for abundant early Miocene hominoid fossils. However, the Wasiriya Beds of Rusinga Island, Kenya, preserve a Pleistocene sedimentary archive with radiocarbon age estimates of >33–45 ka that contains Middle Stone Age artifacts and abundant, well-preserved fossil fauna: a co-occurrence rare in eastern Africa, particularly in the region bounding Lake Victoria. Artifacts and fossils are associated with distal volcanic ash deposits that occur at multiple localities in the Wasiriya Beds, correlated on the basis of geochemical composition as determined by electron probe microanalysis. Sediment lithology and the fossil ungulates suggest a local fluvial system and associated riparian wooded habitat within a predominantly arid grassland setting that differs substantially from the modern environment, where local climate is strongly affected by moisture availability from Lake Victoria. In particular, the presence of oryx (Oryx gazella) and Grevy’s zebra (Equus grevyi) suggest a pre-Last Glacial Maximum expansion of arid grasslands, an environmental reconstruction further supported by the presence of several extinct specialized grazers (Pelorovis antiquus, Megalotragus sp., and a small alcelaphine) that are unknown from Holocene deposits in eastern Africa. The combination of artifacts, a rich fossil fauna, and volcaniclastic sediments makes the Wasiriya Beds a key site for examining the Lake Victoria basin, a biogeographically important area for understanding the diversification and dispersal of Homo sapiens from Africa, whose pre-Last Glacial Maximum history remains poorly understood.     

Stratigraphy, age and environments of the late Miocene Mpesida Beds, Tugen Hills, Kenya.

Interpretations of faunal assemblages from the late Miocene Mpesida Beds in the Tugen Hills of the Central Kenyan Rift Valley have figured prominently in discussions of faunal turnover and establishment of the modern East African communities. These faunal changes have important implications for the divergence of the human lineage from the African apes ca. 8-5 Ma. While fossil material recovered from the Mpesida Beds has traditionally been analyzed collectively, accumulating evidence indicates that Mpesida facies span the 7-6 Ma interval and are scattered more than 25 km along the eastern flanks of the Tugen Hills. Stratigraphic distinctions between Mpesida facies and younger sediments in the sequence, such as the Lukeino Formation, are not yet fully resolved, further complicating temporal assessments and stratigraphic context of Mpesida facies. These issues are discussed with specific reference to exposures of Mpesida facies at Rurmoch, where large fossil tree fragments were swept up in an ancient ash flow. Preserved anatomical features of the fossil wood as well as estimated tree heights suggest a wet, lowland rainforest in this portion of the rift valley. Stable isotopic analyses of fossil enamel and paleosol components indicate the presence of more open habitats locally. Overlying air-fall tuffs and epiclastic debris, possibly associated with the ash flow, have yielded an assemblage of vertebrate fossils including two teeth belonging to one of the earliest colombines of typical body size known from Africa, after the rather small Microcolobus. Single-crystal, laser-fusion,(40)Ar/(39)Ar dates from a capping trachyte flow as well as tuffs just below the lava contact indicate an age of greater than 6.37 Ma for the fossil material.     

A new coral‐associated decapod assemblage from the Upper Miocene (Messinian) Upper Coralline Limestone of Malta (Central Mediterranean)

Abstract: A rich coral‐associated decapod assemblage is recorded from the ‘Depiru Beds’ of the upper part of the Upper Coralline Limestone (Messinian, Upper Miocene), from the island of Malta. Nineteen species within 17 genera have been discovered, where 14 genera are new for Malta. Four new species are described, namely Micippa annamariae sp. nov., Pilumnus scaber sp. nov., Panopeus muelleri sp. nov. and Herbstia melitense sp. nov. Herbstia melitense sp. nov. constitutes the first record of the genus from the fossil record in the Mediterranean region. This discovery more than doubles the number of known fossil decapod species from Malta. The fossil bivalve Jouannetia (J.) semicaudata Des Moulins, 1830 and the extant decapod Maja goltziana D’Oliveira, 1888, are also recorded for the first time from Malta. Other Neogene coral‐associated decapod assemblages are investigated and correlated with the new assemblage from Malta. The migration of taxa between the Mediterranean region and the Paratethys, particularly during the Lower Badenian (Langhian), is evidenced by the strong affinity of the Maltese decapod assemblage with that of the Middle Miocene Badenian assemblages from Hungary, Poland and Ukraine. Upper Miocene, Messinian assemblages from Spain, Algeria and Morocco are also similar to that from Malta.     

Sawfishes and Other Elasmobranch Assemblages from the Mio-Pliocene of the South Caribbean (Urumaco Sequence,Northwestern Venezuela)

The Urumaco stratigraphic sequence, western Venezuela, preserves a variety of paleoenvironments that include terrestrial, riverine, lacustrine and marine facies. A wide range of fossil vertebrates associated with these facies supports the hypothesis of an estuary in that geographic area connected with a hydrographic system that flowed from western Amazonia up to the Proto-Caribbean Sea during the Miocene. Here the elasmobranch assemblages of the middle Miocene to middle Pliocene section of the Urumaco sequence (Socorro, Urumaco and Codore formations) are described. Based on new findings, we document at least 21 taxa of the Lamniformes, Carcharhiniformes, Myliobatiformes and Rajiformes, and describe a new carcharhiniform species (†Carcharhinus caquetius sp. nov.). Moreover, the Urumaco Formation has a high number of well-preserved fossil Pristis rostra, for which we provide a detailed taxonomic revision, and referral in the context of the global Miocene record of Pristis as well as extant species. Using the habitat preference of the living representatives, we hypothesize that the fossil chondrichthyan assemblages from the Urumaco sequence are evidence for marine shallow waters and estuarine habitats.     

Late Miocene continental biota in Northeastern Patagonia (Península Valdés, Chubut, Argentina)

A new vertebrate faunal assemblage was recently discovered from the uppermost part of the late Miocene Puerto Madryn Formation. These deposits crop out along the southwestern coast of the Península Valdés area near Punta Delgada (Chubut Province, Argentina). The exhumed vertebrate fauna includes a range of fish, bird and mammal taxa, of which the latter are most varied and abundant. The new findings represent the first record of continental fossil vertebrates in the Puerto Madryn Formation and this is the first assemblage of late Miocene continental vertebrates recorded to the south of Río Negro Province. It also includes the southernmost record of Hydrochoeridae rodents, Dendrocygninae birds and Loricariidae fishes. The mammals suggest that the fossil-bearing sediments are Huayquerian in age. The climate during the accumulation of late Miocene deposits in this region is inferred to have been warmer and seasonally drier than that of today.     

Fossil fruits of Firmiana and Tilia from the middle Miocene of South Korea and the efficacy of the Bering land bridge for the migration of mesothermal plants

Determining whether the high-latitude Bering land bridge (BLB) was ecologically suitable for the migration of mesothermal plants is significant for Holarctic phytogeographic inferences. Paleobotanical studies provide a critical source of data on the latitudinal positions of different plant lineages at different times, permitting assessment of the efficacy of the BLB for migration. Here we report exceptionally preserved fossils of Firmiana and Tilia endochrysea from the middle Miocene of South Korea. This represents a new reliable record of Firmiana and the first discovery of the T. endochrysea lineage in the fossil record of Asia. The occurrence of these fossils in South Korea indicates that the two lineages had a distribution that extended much farther north during the middle Miocene, but they were still geographically remote from the BLB. In light of the broader fossil record of Asia, our study shows that, in the middle Miocene, some mesothermal plants apparently inhabited the territory adjacent to the BLB and thus they were possibly capable of utilizing the BLB as a migratory corridor. Some other mesothermal plants, such as Firmiana and the T. endochrysea lineages, however, are restricted to more southern regions relative to the BLB based on current fossil evidence. These lineages may have been ecologically unable to traverse the BLB, which raises questions about the efficacy of the BLB as a universal exchange route for mesothermal plants between Asia and North America during the middle Miocene.     

The geologic history of the Lythraceae

The known fossil record of the Lythraceae has been amplified by recent studies in northern Latin America. A total of 18 genera is recognized in geologic strata ranging in age from lower Eocene to Recent, and among the 22 or 23 modern genera, seven have a documented geologic history. The oldest remains are from an Indo-Malayan Old World warm-temperate to subtropical vegetation preserved in the lower Eocene London Clay flora. The most ancient of extant genera isLagerstroemia and the most recent (among those with an adequate fossil record) isCuphea (middle Miocene to Recent). These represent, respectively, primitive and advanced members of the family, and the paleobotanical record supports current concepts concerning phylogenetic relationships among genera of the Lythraceae. The family apparently had an Old World origin and became differentiated into a distinct modern taxon during Paleocene and early Eocene time.     

The temnospondyl amphibian Cyclotosaurus from the Upper Triassic of Poland

Abstract:  A gap in the Late Triassic fossil record of the capitosaur amphibian Cyclotosaurus is filled by new material from lacustrine deposits at Krasiejów, Poland, corresponding in age to the Lehrberg Beds (late Carnian) of Germany. The skull of the Polish cyclotosaur is intermediate in several respects between that of Cyclotosaurus robustus from the middle Carnian Schilfsandstein of Germany and the younger C. mordax from the early Norian Stubensandstein. It shows a decrease in the width of the skull and in the degree of concavity of the posterior margin of the skull roof. The differences are significant enough to warrant erection of a novel species, the name Cyclotosaurus intermedius sp. nov. being proposed. The pectoral girdle, identified for the first time in Cyclotosaurus , suggests the genus was more fully adapted to an aquatic mode of life than was Paracyclotosaurus .     

Life,death and fossilization on Gran Canaria – implications for Macaronesian biogeography and molecular dating

The Canaries have recently served as a test‐bed island system for evaluating newly developed parametric biogeographical methods that can incorporate information from molecular phylogenetic dating and ages of geological events. To use such information successfully, knowledge of geological history and the fossil record is essential. Studies presenting phylogenetic datings of plant groups on oceanic islands often through necessity, but perhaps inappropriately, use the geological age of the oldest island in an archipelago as a maximum‐age constraint for earliest possible introductions. Recently published papers suggest that there is little chance of informative fossil floras being found on volcanic islands, and that nothing could survive violent periods of volcanic activity. One such example is the Roque Nublo period in Gran Canaria, which is assumed to have caused the extinction of the flora of the island (c. 5.3–3.7 Ma). However, recent investigations of Gran Canaria have identified numerous volcanic and sedimentological settings where plant remains are common. We argue, based on evidence from the Miocene–Pliocene rock and fossil records, that complete sterilization of the island is implausible. Moreover, based on fossil evidence, we conclude that the typical ecosystems of the Canary Islands, such as the laurisilva, the Pinus forest and the thermophilous scrubland, were already present on Gran Canaria during the Miocene–Pliocene. The fossil record we present provides new information, which may be used as age constraints in phylogenetic datings, in addition to or instead of the less reliable ages of island emergences or catastrophic events. We also suggest island environments that are likely to yield further fossil localities. Finally, we briefly review further examples of fossil floras of Macaronesia.     

Divergence time estimation in Cichorieae (Asteraceae) using a fossil-calibrated relaxed molecular clock

Knowing the age of lineages is key to understanding their biogeographic history. We aimed to provide the best estimate of the age of Cichorieae and its subtribes based on available fossil evidence and DNA sequences and to interpret their biogeography in the light of Earth history. With more than 1,550 species, the chicory tribe (Cichorieae, Asteraceae) is distributed predominantly in the northern Hemisphere, with centres of distribution in the Mediterranean region, central Asia, and SW North America. Recently, a new phylogenetic hypothesis of Cichorieae based on ITS sequences has been established, shedding new light on phylogenetic relationships within the tribe, which had not been detected so far. Cichorieae possess echinolophate pollen grains, on the surface of which cavities (lacunae) are separated by ridges. These lacunae and ridges show patterns characteristic of certain groups within Cichorieae. Among the fossil record of echinolophate pollen, the Cichorium intybus-type is the most frequent and also the oldest type (22 to 28.4 million years old). By using an uncorrelated relaxed molecular clock approach, the Cichorieae phylogenetic tree was calibrated with this fossil find. According to the analysis, the tribe originated no later than Oligocene. The species-rich core group originated no later than Late Oligocene or Early Miocene and its subtribes diversified no later than Middle/Late Miocene or Early Pliocene—an eventful period of changing geological setting and climate in the Mediterranean region and Eurasia. The first dispersal from Eurasia to North America, which resulted in the radiation of genera and species in North America (subtribe Microseridinae), also occurred no later than Middle or Late Miocene, suggesting the Bering land bridge as the route of dispersal.