The temperate marine Peruvian Province: How history accounts for its unusual biota.

The Peruvian Province, from 6° S in Peru to 42° S in Chile, is a highly productive coastal marine region whose biology and fossil record have long been studied separately but never integrated. To understand how past events and conditions affected today's species composition and interactions, we examined the role of extinction, colonization, geologic changes to explain previously unrecognized peculiar features of the biota and to compare the Peruvian Province's history to that of other climatically similar temperate coasts. We synthesized all available data on the benthic (or benthically feeding) biota, with emphasis on fossilizable taxa, for the interval from the Miocene (23-5.4 Ma) and Pliocene (5.4-2.5 Ma) to the present. We outline the history of ecological guilds including primary producers, herbivores, predators, and suspension-feeders and document patterns of extinction, colonization, and geographic restriction. We identify twelve unusual attributes of the biota, most of which are the result of repeated episodes of extinction. Several guilds present during the Miocene and Pliocene are not represented in the province today, while groups such as kelps and perhaps intertidal predatory sea stars are relative newcomers. Guilds on soft bottoms and in sheltered habitats were severely affected by extinction, whereas those on hard bottoms were most affected by colonists and held their own in diversity. The Peruvian Province has not served as a biogeographic refuge, in contrast to the coasts of Australasia and Argentina, where lineages no longer present in the Peruvian Province survive. The loss of sheltered habitats since the Pliocene explains many of the present-day peculiarities of the biota. The history of the province's biota explains its unique attributes. High productivity, a rich Southern Hemisphere heritage, and colonization from the north account for the present-day composition and unusual characteristics of the biota.

Combined uranium-series and electron spin resonance dating from the Pliocene fossil sites of Aves and Milo's palaeocaves, Bolt's Farm, Cradle of Humankind, South Africa.

Bolt's Farm is the name given to a series of non-hominin bearing fossil sites that have often been suggested to be some of the oldest Pliocene sites in the Cradle of Humankind, South Africa. This article reports the results of the first combined Uranium-Series and Electron Spin Resonance (US-ESR) dating of bovid teeth at Milo's Cave and Aves Cave at Bolt's Farm. Both tooth enamel fragments and tooth enamel powder ages were presented for comparison. US-ESR, EU and LU models are calculated. Overall, the powder ages are consistent with previous uranium-lead and palaeomagnetic age estimates for the Aves Cave deposit, which suggest an age between ~3.15 and 2.61 Ma and provide the first ages for Milo's Cave dates to between ~3.1 and 2.7 Ma. The final ages were not overly dependent on the models used (US-ESR, LU or EU), which all overlap within error. These ages are all consistent with the biochronological age estimate (<3.4->2.6 Ma) based on the occurrence of Stage I Metridiochoerus andrewsi. Preliminary palaeomagnetic analysis from Milo's Cave indicates a reversal takes place at the site with predominantly intermediate directions, suggesting the deposit may date to the period between ~3.03 and 3.11 Ma within error of the ESR ages. This further suggests that there are no definitive examples of palaeocave deposits at Bolt's Farm older than 3.2 Ma. This research indicates that US-ESR dating has the potential to date fossil sites in the Cradle of Humankind to over 3 Ma. However, bulk sample analysis for US-ESR dating is recommended for sites over 3 Ma.

Pseudopus pannonicus (Squamata), the largest known anguid lizard-Redescription of the type material and new specimens from the Neogene and Quaternary of Hungary and Poland.

Herein, we revise the material of the extinct taxon Pseudopus pannonicus from Central Europe, the largest known anguid lizard and iconic member of herpetofaunas from the Upper Cenozoic of Europe. The geographical position of Polgárdi 2, the type locality of P. pannonicus, as well as several other closely located important localities make Central Europe a valuable area of high interest for studies regarding European Cenozoic palaeoherpetological assemblages. We clarified the nature of the type material of Pseudopus pannonicus, showing that it consisted not only of the five specimens originally figured. Instead, the syntype series also includes a considerable number of specimens from Polgárdi 2 that were only described or figured 12 years after the original description of P. pannonicus. Detailed osteological descriptions are provided for specimens from the type series and Polish specimens, with the aid of high-resolution imaging (SEM and µCT scanning), and intraspecific variability is discussed. The articular surface with the lappet of the parietal overlapping the frontal is discussed as a character potentially relevant for the diagnosis of P. pannonicus. We updated the identification of several specimens of P. pannonicus and discussed the biogeographic implications of such revisions. In Poland, P. pannonicus is an abundant component of Neogene and early Quaternary herpetofaunas, known with certainty from the Middle Miocene of Przeworno (the oldest known occurrence of the species globally), the Early-Late Pliocene of Weze I, the Late Pliocene of Rebielice Królewskie I, the Late Pliocene of Weze II, and the Early Pleistocene of Kadzielnia (one of the youngest occurrences documented globally). An indeterminate anguine with Anguis affinities is newly reported from Rebielice Królewskie II. The taxonomic status of other large anguids from the Neogene of Europe is discussed and we conclude that most are junior synonyms of P. pannonicus. We also show that another purported synonym, that is, Ophisaurus intermedius from the Early Pleistocene of Romania, is instead a nomen nudum. We conducted phylogenetic analyses (18 taxa, 65 characters) to understand the relationship of P. pannonicus relative to other anguid representatives and anguid-related group (i.e., glyptosaurids). A single most parsimonious tree (length: 134 steps) was recovered. The clade Pseudopus is stable, comprising the two distinct sister subclades [Pseudopus laurillardi + Pseudopus ahnikoviensis] and [Pseudopus pannonicus + Pseudopus apodus]. These phylogenetic results are in accordance with previously published works.

A new fossil species of Meliolinites Selkirk associated with Rhodoleia leaves from the Upper Pliocene of southwestern China.

Fossil epifoliar fungi are valuable indicators of paleoenvironment and paleoecology. The Meliolaceae, members of which typically inhabit the surface of living plants as biotrophs or pathogens, is one of the largest groups of epifoliar fungi. In this study, we report a novel fossil species of Meliolinites Selkirk (fossil Meliolaceae), Meliolinites tengchongensis, on the lower epidermis of compressed fossil Rhodoleia (Hamamelidaceae) leaves from the Upper Pliocene Mangbang Formation of Tengchong, Yunnan, southwestern China. Meliolinites tengchongensis is characterized by web-like, superficial, brown to dark brown, septate, and branching mycelia bearing 2-celled appressoria and unicellular phialides. The fungal colonies also include ellipsoidal, 5-celled, 4-septate ascospores and dark brown perithecia with suborbicular outline and verrucose surface. The well-preserved vegetative and reproductive organs help us to explore the potential disease process of the new fossil species. Besides, the presence of fungal remains indicates that the fungal taxon might have maintained its host preference since at least the Late Pliocene. Furthermore, the occurrence of both fossil fungi and their host plants in Tengchong indicate a subtropical-tropical, warm, and humid climate during the Late Pliocene, whereas the distribution pattern of the fungi on the host leaves suggests that Rhodoleia may have been a part of the middle-upper canopies in the Tengchong Late Pliocene multilayered forest.

Leaf ecological stoichiometry and anatomical structural adaptation mechanisms of Quercus sect. Heterobalanus in southeastern Qinghai-Tibet Plateau.

BACKGROUND: With the dramatic uplift of the Qinghai-Tibet Plateau (QTP) and the increase in altitude in the Pliocene, the environment became dry and cold, thermophilous plants that originally inhabited ancient subtropical forest essentially disappeared. However, Quercus sect. Heterobalanus (QSH) have gradually become dominant or constructive species distributed on harsh sites in the Hengduan Mountains range in southeastern QTP, Southwest China. Ecological stoichiometry reveals the survival strategies plants adopt to adapt to changing environment by quantifying the proportions and relationships of elements in plants. Simultaneously, as the most sensitive organs of plants to their environment, the structure of leaves reflects of the long-term adaptability of plants to their surrounding environments. Therefore, ecological adaptation mechanisms related to ecological stoichiometry and leaf anatomical structure of QSH were explored. In this study, stoichiometric characteristics were determined by measuring leaf carbon (C), nitrogen (N), and phosphorus (P) contents, and morphological adaptations were determined by examining leaf anatomical traits with microscopy. RESULTS: Different QSH life forms and species had different nutrient allocation strategies. Leaves of QSH plants had higher C and P and lower N contents and higher N and lower P utilization efficiencies. According to an N: P ratio threshold, the growth of QSH species was limited by N, except that of Q. aquifolioides and Q. longispica, which was limited by both N and P. Although stoichiometric homeostasis of C, N, and P and C: N, C: P, and N: P ratios differed slightly across life forms and species, the overall degree of homeostasis was strong, with strictly homeostatic, homeostatic, and weakly homeostatic regulation. In addition, QSH leaves had compound epidermis, thick cuticle, developed palisade tissue and spongy tissue. However, leaves were relatively thin overall, possibly due to leaf leathering and lignification, which is strategy to resist stress from UV radiation, drought, and frost. Furthermore, contents of C, N, and P and stoichiometric ratios were significantly correlated with leaf anatomical traits. CONCLUSIONS: QSH adapt to the plateau environment by adjusting the content and utilization efficiencies of C, N, and P elements. Strong stoichiometric homeostasis of QSH was likely a strategy to mitigate nutrient limitation. The unique leaf structure of the compound epidermis, thick cuticle, well-developed palisade tissue and spongy tissue is another adaptive mechanism for QSH to survive in the plateau environment. The anatomical adaptations and nutrient utilization strategies of QSH may have coevolved during long-term succession over millions of years.

Out of the desert: Paleoclimatic changes drove the diversification of arid-adapted Ocymyrmex ants in southern Africa.

A highly endemic ant fauna is found in the arid regions of southern Africa, including species in the genus Ocymyrmex. This genus of ants has higher species richness in the western arid regions of southern Africa compared to tropical and subtropical parts of the continent. The processes that have produced these patterns of diversity and distribution of arid adapted ants in southern Africa have never been investigated. The diversification of many other taxa in the region has been associated with past climate fluctuations that occurred during the Miocene epoch. In this study, the nature and timing of historical processes that may have led to the diversification within Ocymyrmex were assessed. We hypothesized that past climate oscillations, characterized by long periods of aridification, have driven the current distribution of Ocymyrmex species that resulted in the highest species richness of the genus in the Deserts & xeric shrublands biome in southern Africa. Ninety-four Ocymyrmex worker specimens from Botswana, Kenya, Namibia, South Africa, Tanzania and Zimbabwe, representing 21 currently described species and six morphospecies, were included in a phylogenomic analysis. Phylogenies for the genus, based on next generation sequencing data from ultraconserved elements, were inferred using Maximum Likelihood, and a dating analysis was performed using secondary age estimates as calibration points. A distribution database of Ocymyrmex records was used to assign species ranges, which were then coded according to major biomes in southern Africa and used as input for biogeographical analysis. We explored the phylogenomic relationships of Ocymyrmex and analysed these within a biogeographical and paleoclimatic framework to disentangle the potential processes responsible for diversification in this group. Dating analyses estimated that the crown age of Ocymyrmex dates to the Oligocene, around 32 Ma. Diversification within this group occurred between the mid-Miocene (∼12.5 Ma) and Pleistocene (∼2 Ma). Our biogeographic analyses suggest that Ocymyrmex species originated in the south-western region of southern Africa, which is now part of the Deserts & xeric shrublands biome and diversified into eastern subtropical areas during the Pliocene. Paleoclimatic changes resulting in increased aridity during the Miocene likely drove the diversification of the genus Ocymyrmex. It is most likely that the diversification of grasslands, because of historical climate change, facilitated the diversification of these ants to the eastern parts of southern Africa when open grasslands replaced forests during the early Miocene.

An Updated Review of Fossil Pollen Evidence for the Study of the Origin, Evolution and Diversification of Caribbean Mangroves.

Recently, the evolutionary history of the Caribbean mangroves has been reconsidered using partial palynological databases organized by the time intervals of interest, namely Late Cretaceous to Eocene for the origin, the Eocene-Oligocene transition for major turnover and Neogene to Quaternary for diversification. These discussions have been published in a set of sequential papers, but the raw information remains unknown. This paper reviews all the information available and provides the first comprehensive and updated compilation of the abovementioned partial databases. This compilation is called CARMA-F (CARibbean MAngroves-Fossil) and includes nearly 90 localities from the present and past Caribbean coasts, ranging from the Late Cretaceous to the Pliocene. Details on the Quaternary localities (CARMA-Q) will be published later. CARMA-F lists and illustrates the fossil pollen from past mangrove taxa and their extant representatives, and includes a map of the studied localities and a conventional spreadsheet with the raw data. The compilation is the most complete available for the study of the origin, evolution and diversification of Caribbean mangroves, and is open to modifications for adapting it to the particular interests of each researcher.

The Deep Past of the White Shark, Carcharodon carcharias, in the Mediterranean Sea: A Synthesis of Its Palaeobiology and Palaeoecology.

The white shark, Carcharodon carcharias, is the main top predator of the present-day Mediterranean Sea. The deep past of C. carcharias in the Mediterranean is witnessed by a rather conspicuous, mostly Pliocene fossil record. Here, we provide a synthesis of the palaeobiology and palaeoecology of the Mediterranean white sharks. Phenetically modern white shark teeth first appeared around the Miocene-Pliocene transition in the Pacific, and soon after in the Mediterranean. Molecular phylogenetic analyses support an origin of the Mediterranean white shark population from the dispersal of Australian/Pacific palaeopopulations, which may have occurred through the Central American Seaway. Tooth dimensions suggest that the Mediterranean white sharks could have grown up to about 7 m total length during the Pliocene. A richer-than-today marine mammal fauna was likely pivotal in supporting the Mediterranean white sharks through the Pliocene and most of the Quaternary. White sharks have seemingly become more common as other macropredators declined and disappeared, notwithstanding the concurrent demise of many potential prey items in the context of the latest Pliocene and Quaternary climatic and environmental perturbations of the Mediterranean region. The overall generalist trophic habits of C. carcharias were likely crucial for securing ecological success in the highly variable Mediterranean scenario by allowing the transition to a mostly piscivorous diet as the regional marine mammal fauna shrank.

New Materials on the Morphology of the Teeth of the Three-Toed Horse (Hipparion houfenense) from the Pliocene of Western Transbaikalia (Russia).

This article is devoted to the morphological features of the teeth of the three-toed horse (Hipparion houfenense) from the Early Pliocene of Western Transbaikalia (Russia). It contains a number of diagnostic features that are unique to this taxon and distinguish it from other Hipparion species, which allows us to speak about their true diversity at the final stage of their existence in the northeast of Inner Asia.

Their fates intertwined: diversification patterns of the Asian gliding vertebrates may have been forged by dipterocarp trees.

The repeated evolution of gliding in diverse Asian vertebrate lineages is hypothesized to have been triggered by the dominance of tall dipterocarp trees in the tropical forests of Southeast Asia. These dipterocarp forests have acted as both centres of diversification and climatic refugia for gliding vertebrates, and support most of their extant diversity. We predict similarities in the diversification patterns of dipterocarp trees and gliding vertebrates, and specifically test whether episodic diversification events such as rate shifts and/or mass extinctions were temporally congruent in these groups. We analysed diversification patterns in reconstructed timetrees of Asian dipterocarps, the most speciose gliding vertebrates from different classes (Draco lizards, gliding frogs and Pteromyini squirrels) and compared them with similar-sized clades of non-gliding relatives (Diploderma lizards, Philautus frogs and Callosciurinae squirrels) from Southeast Asia. We found significant declines in net-diversification rates of dipterocarps and the gliding vertebrates during the Pliocene-Pleistocene, but not in the non-gliding groups. We conclude that the homogeneity and temporal coincidence of these rate declines point to a viable ecological correlation between dipterocarps and the gliding vertebrates. Further, we suggest that while the diversification decay in dipterocarps was precipitated by post-Miocene aridification of Asia, the crises in the gliding vertebrates were induced by both events concomitantly.

Sedimentological and palynostratigraphical modeling of sediments penetrated by KW field wells, onshore western Niger Delta Basin, Nigeria.

Sedimentological and palynostratigraphical studies were performed on 189 ditch-cutting samples composited at 50 ft intervals from four well sections located in KW field, onshore western Niger Delta Basin, Nigeria, using sieving and visual microscopic observations, hydrochloric and hydrofluoric acids digestion methods to determine their lithological character and palynologic content of the sedimentary successions. The basis of these data is to interpret the lithofacies and biostratigraphy, establish the age of the strata, establish the palynological zones and infer the environments of deposition. Five lithofacies delineated based on the sedimentological study were very coarse sand, coarse to fine sand, shaly sand, muddy sand, and sandy to silty shale. The age of the sedimentary successions ranges from Early Miocene to Pliocene based on the bio-assemblages of these key species. The Early Miocene age is characterized by the occurrence of Magnastriatites howardi, Psilatricolporites triangularis, Praedapollis africanus, Pachydermites diederixi, Spirosyncoilpites spiralis, Praedapollis flexibility, Racemonocolpites hians, Verrutricolporites rotundiporus, Verrutricolporites microporus, and Psilatricolporites divisus. The Middle Miocene age is characterized by the FAD of Crassoretitriletes vanraadshooveni, the occurrence of Belskipollis elegans, in association with Striatricolporites catatumbus, Verrutricolporites rotundiporus, Verrutricolporites microporus, Crassoretitriletes vanraadshooveni, Racemonocolpites hians, Retibrevitricolpites obodoensis, and Retibrevitricolpites protrudens. The Late Miocene age is characterized by the occurrence of Multiareolites formosus, Cyperaceaepollis sp., Stereiosporites sp., Peregrinipollis nigericus, Gemmamonoporites sp. (Cleistopholis patens), Laevigatosporites sp., Verrutricolporites sp., Nymphaeapollis clarus, and Verrutricolporites usmensis. The Pliocene age is characterized by the basal occurrence of Retistephenocolpites gracilis in association with Nymphaeapollis clarus, Cyperaceaepollis sp., Laevigatosporites sp., Verrutricolporites sp., and Crassoretitriletes vanraadshooveni. The boundary between the Early Miocene and Middle Miocene from the studied oil well sections is marked by the first appearance datum (FAD) and numerical maximum occurrence of Striatricolporites catatumbus and Belskipollis elegans and FAD of Crassoretitrilettes vanraadshooveni. The boundary between the Middle and late Miocene is defined by the LAD of Verrutricolporites rotundiporus, the quantitative top and numerical maximum occurrence of Racemonocolpites hians, and the numerical maximum occurrence of Zonocostites ramonae. The boundary between the late Miocene and the Early Pliocene is marked by the first appearance datum (FAD) and the numerical maximum occurrence of Retistephenocolpites gracilis, the numerical maximum occurrence of Echiperiporites icacinoides, and the highest abundance of Monoporites annulatus. Four palynozones PAZ-1, PAZ-2, PAZ-3, and PAZ-4, and ten sub-zones were erected. Finally, Fluvial-coastal to shallow marine environments of deposition were inferred for the sedimentary successions using sedimentological study, nature of organic matter, and association of key environmentally-sensitive biomarker species.

Description of the Pliocene marsupial Ambulator keanei gen. nov. (Marsupialia: Diprotodontidae) from inland Australia and its locomotory adaptations.

Diprotodontids were the largest marsupials to exist and an integral part of Australian terrestrial ecosystems until the last members of the group became extinct approximately 40 000 years ago. Despite the frequency with which diprotodontid remains are encountered, key aspects of their morphology, systematics, ecology and evolutionary history remain poorly understood. Here we describe new skeletal remains of the Pliocene taxon Zygomaturus keanei from northern South Australia. This is only the third partial skeleton of a late Cenozoic diprotodontid described in the last century, and the first displaying soft tissue structures associated with footpad impressions. Whereas it is difficult to distinguish Z. keanei and the type species of the genus, Z. trilobus, on dental grounds, the marked cranial and postcranial differences suggest that Z. keanei warrants genus-level distinction. Accordingly, we place it in the monotypic Ambulator gen. nov. We, also recognize the late Miocene Z. gilli as a nomen dubium. Features of the forelimb, manus and pes reveal that Ambulator keanei was more graviportal with greater adaptation to quadrupedal walking than earlier diprotodontids. These adaptations may have been driven by a need to travel longer distances to obtain resources as open habitats expanded in the late Pliocene of inland Australia.

New Data on Dental Morphology of Hipparion tchicoicum Ivanjev, 1966 from Western Transbaikalia, Russia.

Morphological features of the teeth were studied in the three-toed horse Hipparion tchicoicum from the Pliocene of Western Transbaikalia (Russia). Several diagnostic signs of the Chicoi hipparion were described for the first time to provide criteria for distinguishing the taxon among other fossils of three-toed horses and estimating their real diversity at the final stage of their distribution in Inner Asia.

The earliest most complete skeleton of Theropithecus.

The middle Pliocene site of Woranso-Mille in the Afar Region of Ethiopia has yielded numerous significant early hominin fossils representing multiple, coexisting taxa. Here we report on another significant discovery, the oldest partial skeleton of the papionin, Theropithecus. The specimen was recovered from the Aralee Issie collection area over multiple field seasons from 2004 through 2019. The specimen was unearthed in situ from the fluvial facies of the Mesgid Dora Tuff dated to 3.66-3.57 Ma. The partial skeleton, ARI-VP-1/26, is that of a subadult male lacking a skull. In the absence of unambiguously associated craniodental remains, the male status of the specimen was established from the dimensions of the long bones in comparison to those of other Theropithecus from Woranso-Mille. ARI-VP-1/26 is noteworthy because it preserves partial hand and foot skeletons, including a complete set of metacarpals from the left side. The theropith status of ARI-VP-1/26 was established based on the detailed anatomy of the postcranial skeleton, especially the proximal and distal humerus, proximal radius, and proximal femur. The morphology of the postcranium of ARI-VP-1/26 is consistent with that of recognized Theropithecus from Woranso-Mille and, specifically, with specimens recognized as Theropithecus oswaldi cf. darti from other sites. The ratio of the lengths of the first metacarpal to metacarpals 2-5 in ARI-VP-1/26 is intermediate between that seen in extant Papio and Theropithecus. In Theropithecus gelada and Theropithecus brumpti, the pairing of pollical and indical metacarpals of near equal length contributes to the species' high opposability index and is associated with a 'manual grazing' feeding habit. Cercopithecids constitute 43% of the identified vertebrates at Aralee Issie, and T. oswaldi cf. darti is the most common mammalian species. The monkeys of Aralee Issie lived in an open shrubland habitat, but the specific reasons for their high prevalence at the site are unclear.

Pliocene hominins from East Turkana were associated with mesic environments in a semiarid basin.

During the middle Pliocene (∼3.8-3.2 Ma), both Australopithecus afarensis and Kenyanthropus platyops are known from the Turkana Basin, but between 3.60 and 3.44 Ma, most hominin fossils are found on the west side of Lake Turkana. Here, we describe a new hominin locality (ET03-166/168, Area 129) from the east side of the lake, in the Lokochot Member of the Koobi Fora Formation (3.60-3.44 Ma). To reconstruct the paleoecology of the locality and its surroundings, we combine information from sedimentology, the relative abundance of associated mammalian fauna, phytoliths, and stable isotopes from plant wax biomarkers, pedogenic carbonates, and fossil tooth enamel. The combined evidence provides a detailed view of the local paleoenvironment occupied by these Pliocene hominins, where a biodiverse community of primates, including hominins, and other mammals inhabited humid, grassy woodlands in a fluvial floodplain setting. Between <3.596 and 3.44 Ma, increases in woody vegetation were, at times, associated with increases in arid-adapted grasses. This suggests that Pliocene vegetation included woody species that were resilient to periods of prolonged aridity, resembling vegetation structure in the Turkana Basin today, where arid-adapted woody plants are a significant component of the ecosystem. Pedogenic carbonates indicate more woody vegetation than other vegetation proxies, possibly due to differences in temporospatial scale and ecological biases in preservation that should be accounted for in future studies. These new hominin fossils and associated multiproxy paleoenvironmental indicators from a single locale through time suggest that early hominin species occupied a wide range of habitats, possibly including wetlands within semiarid landscapes. Local-scale paleoecological evidence from East Turkana supports regional evidence that middle Pliocene eastern Africa may have experienced large-scale, climate-driven periods of aridity. This information extends our understanding of hominin environments beyond the limits of simple wooded, grassy, or mosaic environmental descriptions.

Bone histology of Neogene angulate tortoises (Testudines: Testudinidae) from South Africa: palaeobiological and skeletochronological implications.

Here we examine the tibial microstructure of modern and fossil angulate tortoises to assess the histology and growth from the late Miocene-early Pliocene, Pleistocene through to modern forms. The cross-sections of all the tibiae sampled revealed highly vascularized, uninterrupted, fibrolamellar bone tissue during early ontogeny, which suggests that early growth was fast. However, later in ontogeny, growth was slower, as indicated by the deposition of parallel-fibred bone tissue in the outer cortex, and even ceased periodically, as indicated by lines of arrested growth. Comparative analyses of the growth rates of the tortoises from different time periods showed that the tortoises from the late Miocene-early Pliocene Langebaanweg locality and from Diepkloof Rock Shelter had relatively slower growth rates under less optimal growth conditions. Additionally, these prehistoric specimens show extensive remodelling, and several generations of secondary osteons further suggest functional and/or metabolic stresses on the skeleton. Palaeoenvironmental reconstructions suggest that it was mostly cooler and drier with seasonal fluctuations in late Miocene-early Pliocene, and it is likely that Chersina responded to these conditions by having a lower rate of growth as compared with their modern counterparts, which thrive in the current prevailing more favourable Mediterranean type of climate.

First report on rhinoceros from the late Neogene Qin Basin of Shanxi, China.

Dihoplus is a rhinoceros distributed across East Asia and Europe from the Late Miocene to Pliocene. This study describes a new skull from the Qin Basin in Shanxi Province, China, referred to as Dihoplus ringstroemi, which has long been debated in taxonomic identity. This skull confirms that D. ringstroemi is an independent species and reveals the presence of the upper incisor and variations in the degree of constriction of the two lingual cusps of upper cheek teeth. In addition, the new skull indicates that the Qin Basin has a late Neogene sediment and fauna comparable to that of the Yushe Basin.

Biodiversity of extant snails (Gastropoda, Mollusca) in the Pliocene Mountain Spur Natural Reserve (Northern Apennine, Italy).

Background: To date, there is a substantial lack of information about gastropods from the Tuscan-Emilian Apennine in the north of Italy, notwithstanding the availability of detailed and comprehensive literature on this molluscan class. We present a gastropod fauna from the Natural Reserve of the Pliocene Mountain Spur: to our knowledge, this is the first investigation of the extant gastropod fauna in the Tuscan-Emilian Apennine and one of the few in the Apennine's mountain chain as a whole. New information: We describe a gastropod fauna comprised by 25 species, belonging to 18 genera and 10 families: the general figure which is emerging is an assemblage of European and Mediterranean-European species, with a single Asian contribution. Several shells were collected as embedded in sandy-clayey soils and showed fossilisation traces: therefore, we regard these shells as subfossil samples. Namely, subfossil shells are from the species Pomatiaselegans, Granariafrumentum, Retinellaolivetorum, Xerolentaobviaobvia, Oxychiluscf.draparnaudi, Monachacartusiana and Monachacantiana. The present checklist is the first report in the Tuscan-Emilian Apennine and Emilia-Romagna areas for six taxa: Morlinaglabraglabra, Oxychiliusalliarius, Xerosectacespitum, Fruticicolafruticum, Xerogyraspadae and Xerolentaobviaobvia.

Assessment of soil-gas radon concentration over lithologies: a case study from district Karak, Khyber Pakhtunkhwa, Pakistan.

The current study is aimed to determine the variation of soil-gas radon concentrations over different rock formations representing diverse lithologies in the district of Karak, Khyber Pakhtunkhwa, Pakistan. The stratigraphic units were grouped on the basis of lithological contents into four categories, i.e., limestone, evaporites, claystone/mudstone, and sandstone. The highest average soil-gas 222Rn concentration (544 Bq/L) was found in the uranium-bearing Dhok Pathan Formation of the Pliocene age, while the lowest radon levels (0.15 Bq/L) were observed in the salt-bearing strata of Bahadurkhel Salt of Eocene age showing the non-uraniferous nature of the salt. High radon potential associated with the Dhok Pathan Formation is likely to be related to the high degree of uranium mineralization which is contributing to the elevated soil-gas radon levels. The study revealed that the soil-gas radon concentration in all lithologies is varying in the order of RnSandstone > RnLimestone > RnClaystone/Mudstone > RnEvaporites with the highest radon levels in the sandstone unit of uranium-bearing Dhok Pathan Formation. High fluctuations of soil-gas radon levels observed in this study evidently show that lithology and uranium mineralization have strong control over the 222Rn concentrations.

Biochronology of South African hominin-bearing sites: A reassessment using cercopithecid primates.

Despite recent advances in chronometric techniques (e.g., Uranium-Lead [U-Pb], cosmogenic nuclides, electron spin resonance spectroscopy [ESR]), considerable uncertainty remains regarding the age of many Plio-Pleistocene hominin sites, including several in South Africa. Consequently, biochronology remains important in assessments of Plio-Pleistocene geochronology and provides direct age estimates of the fossils themselves. Historically, cercopithecid monkeys have been among the most useful taxa for biochronology of early hominins because they are widely present and abundant in the African Plio-Pleistocene record. The last major studies using cercopithecids were published over 30 y ago. Since then, new hominin sites have been discovered, radiometric age estimates have been refined, and many changes have occurred in cercopithecid taxonomy and systematics. Thus, a biochronological reassessment using cercopithecids is long overdue. Here, we provide just such a revision based on our recent study of every major cercopithecid collection from African Plio-Pleistocene sites. In addition to correlations based on shared faunal elements, we present an analysis based on the dentition of the abundant cercopithecid Theropithecus oswaldi, which increases in size in a manner that is strongly correlated with geological age (r2 ∼0.83), thereby providing a highly accurate age-estimation tool not previously utilized. In combination with paleomagnetic and U-Pb data, our results provide revised age estimates and suggest that there are no hominin sites in South Africa significantly older than ∼2.8 Ma. Where conflicting age estimates exist, we suggest that additional data are needed and recall that faunal estimates have ultimately proved reliable in the past (e.g., the age of the KBS Tuff).